Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Temporal variation in saproxylic beetle assemblages in a Mediterranean ecosystem

One of the main challenges in biological conservation has been to understand species distribution across space and time. Over the last decades, many diversity and conservation surveys have been conducted that have revealed that habitat heterogeneity acts as a major factor that determines saproxylic assemblages. However, temporal dynamics have been poorly studied, especially in Mediterranean forests. We analyzed saproxylic beetle distribution at inter and intra-annual scales in a “dehesa” ecosystem, which is a traditional Iberian agrosilvopastoral ecosystem that is characterized by the presence of old and scattered trees that dominate the landscape. Significant differences in effective numbers of families/species and species richness were found at the inter-annual scale, but this was not the case for composition. Temperature and relative humidity did not explain these changes which were mainly due to the presence of rare species. At the intra-annual scale, significant differences in the effective numbers of families/species, species richness and composition between seasons were found, and diversity partitioning revealed that season contributed significantly to gamma-diversity. Saproxylic beetle assemblages exhibited a marked seasonality in richness but not in abundance, with two peaks of activity, the highest between May and June, and the second between September and October. This pattern is mainly driven by the seasonality of the climate in the Mediterranean region, which influences ecosystem dynamics and imposes a marked seasonality on insect assemblages. An extended sampling period over different seasons allowed an overview of saproxylic dynamics, and revealed which families/species were restricted to particular seasons. Recognizing that seasons act as a driver in modelling saproxylic beetle assemblages might be a valuable tool in monitoring and for conservation strategies in Mediterranean forests.     

Intra-annual patterns of saproxylic beetle assemblages inhabiting Mediterranean oak forests

The temporal factor has been widely neglected in studies related to species distribution. However, knowing the species phenology throughout the year could be a key element in ecological studies, more so considering the current climate change framework. The intra-annual temporal patterns of the saproxylic beetle assemblage inhabiting the oak forests in the western Iberian Peninsula were evaluated in terms of species richness and composition. Moreover, we analysed the temporal patterns of species richness and body size, with respect to the abiotic factors of temperature and relative humidity. The marked seasonality of the Mediterranean weather governed, to a great extent, the temporal patterns of the assemblage, as both species richness and body size showed a positive correlation with temperature and a negative correlation with relative humidity. The high temperatures reached in August limited the activity of certain species, causing a marked decrease in species richness. The results revealed that species body size decreased in the colder months of the year and vice versa, which could fit with the inverse of Bergmann’s rule; nevertheless, studies are needed to address the contribution of species physiology and autoecology to the observed temporal patterns. Regarding species composition, the partition of beta diversity in its two components showed a constant process of species turnover throughout the year. We conclude that, due to global warming, the study of temporal patterns of assemblages is becoming all the more important, since it could significantly affect the spatio-temporal distribution of species and the interactions between them.     

The impact of hedge-forest connectivity and microhabitat conditions on spider and carabid beetle assemblages in agricultural landscapes

Agricultural intensification in terms of decreasing landscape complexity and connectivity has negatively affected biodiversity. Linear landscape elements composed of woody vegetation like hedges may counteract this negative trend by providing habitats and enhancing habitat connectivity for different organisms. Here, we tested the impacts of habitat type (forest edges vs. hedges) and hedges’ isolation (connected vs. isolated hedges) from forests as well as microhabitat conditions (percentage of bare ground and width) on trait-specific occurrence of ground-dwelling arthropods, namely spiders and carabids. Arthropods were grouped by habitat specialisation (forest vs. open-habitat species vs. generalists), hunting strategy (web-building or hunting spiders) and dispersal ability (wing morphology of carabids). Spider and carabid assemblage composition was strongly influenced by habitat type and isolation, but not by microhabitat conditions. Activity density of forest species and brachypterous carabids was higher in forest edges compared to hedges, whereas open-habitat species and macropterous carabids showed reverse patterns, with no effects of isolation. Occurrence of generalist carabids, but not spiders, was higher in hedges compared to forest edges. Habitat type and isolation did not affect spiders with different hunting strategy. Microhabitat conditions were less important for spider and carabid occurrence. Our study concludes that on a landscape scale, type of linear woody habitat is more important for arthropod occurrence than isolation effects and microhabitat conditions, depending on traits. Hedges provide refuges for species specialised to open habitats and species with high dispersal ability, such as macropterous carabids. Forest edges enhance persistence of species specialised to forests and species with low dispersal ability, such as brachypterous carabids.     

Influence of tree hollow characteristics on saproxylic beetle diversity in a managed forest

Tree hollows are key structures in forest ecosystems constituting long-lasting habitats and nutritional resources for many saproxylic arthropod species. Due to diverse microhabitat structures and conditions in tree hollows, they can support a broad range of species. However, in the past intensive management practices in parts of Europe reduced the abundance of tree hollows, resulting in a decrease and endangerment of species specialised in this tree habitat. We investigated 40 beech trees with hollows in 2014 and a subset of 23 of these trees in 2015 in a managed forest landscape in Germany. Using emergence traps we collected 89 beetle species of which 33% were on the Bavarian Red List. We described the tree characteristics, physical hollow characteristics, and their surrounding environment investigating their influence on α-diversity of non-Red List and Red List species. Furthermore, we investigated spatial (between tree hollows) and temporal (same tree hollow but different years) β-diversity, considering the importance of turnover and nestedness components on β-diversity. α-Diversity decreased with increasing decomposition of wood mould and increased with increasing area of hollow entrance in both years. Additional characteristics differed between years and between non-Red List and Red List species. β-Diversity was related to diameter at breast height, number of surrounding tree hollows, area of hollow entrance and a temperature gradient. We found a higher species turnover than nestedness between tree hollows and between years, indicating highly dynamic beetle communities spatially as well as temporally. To support and maintain the diversity of saproxylic beetles inhabiting tree hollows, the heterogeneity of microhabitats is important and should be supported by maintaining the diversity of differently structured and sized tree hollows.     

Bird assemblages in fragmented agricultural landscapes: the role of small brigalow remnants and adjoining land uses

Agricultural intensification typically leads to changes in bird diversity and community composition, with fewer species and foraging guilds present in more intensively managed parts of the landscape. In this study, we compare bird communities in small (2–32 ha) brigalow (Acacia harpophylla) remnants with those in adjacent uncultivated grassland, previously cultivated grassland and current cropland, to determine the contribution of different land uses to bird diversity in the agricultural landscape. Twenty remnant brigalow patches and adjacent agricultural (‘matrix’) areas in southern inland Queensland, Australia were sampled for bird composition and habitat characteristics. The richness, abundance and diversity of birds were all significantly higher in brigalow remnants than in the adjacent matrix of cropping and grassland. Within the matrix, species richness and diversity were higher in uncultivated grasslands than in current cultivation or previously cultivated grasslands. Forty-four percent of bird species were recorded only in brigalow remnants and 78% of species were recorded in brigalow and at least one other land management category. Despite high levels of landscape fragmentation and modification, small patches of remnant brigalow vegetation provide important habitat for a unique and diverse assemblage of native birds. The less intensively managed components of the agricultural matrix also support diverse bird assemblages and thus, may be important for local and regional biodiversity conservation.     

Unexpectedly diverse forest dung beetle communities in degraded rain forest landscapes in Madagascar

Tropical forests, which harbor high levels of biodiversity, are being lost at an alarming speed. Madagascar, a biodiversity hotspot, has lost more than half of its original forest cover. Most of the remaining forests are small fragments of primary and secondary forest with differing degrees of human impact. These forests, as well as coffee and fruit plantations, may be important in supporting the forest-dependent biodiversity in Madagascar but this has been little studied. In Madagascar, dung beetles, which offer important ecosystem services, are largely restricted to forests. We examined the ability of fragmented and degraded forests to support dung beetle diversity, compared to the large areas of primary forest in eastern Madagascar. We found a general trend of a reduction of species with a loss of forest connectivity. In contrast, a higher level of forest disturbance was associated with higher species diversity. In several sites of low-quality forest as many or more species were found as in less disturbed and primary forests. The average size of dung beetles was smaller in the lower quality localities than in the primary forests. These findings suggest that many forest dung beetles in Madagascar are better adapted to forest disturbance than earlier expected, although they require some level of connectivity to surrounding forest.     

Micro-habitat drivers of saproxylic beetle assemblages in old woodlands of Mediterranean cork oak (Quercus suber)

1. Cork oak landscapes are fascinating ecosystems, historically managed for cork extraction. The persistence in this habitat of many hollow veteran trees provides suitable micro-habitats for saproxylic beetles.
2. We investigated the saproxylic beetle community of two isolated cork oak woodlands of central Italy with different degree of recovery after human transformation: (1) an open woodland and (2) a dense mixed woodland, both dominated by cork oak trees.
3. We found endemic, rare and threatened saproxylic beetles in both the areas, confirming the important conservation value of cork oak landscapes. In the open woodland we observed a higher number of species in all trophic categories, except for mycophagous specialists. Several microhabitat variables reflected the different stage of recovery of the two woodlands.
4. Our findings suggest the crucial role of diversified environments in protected areas: even a small difference in the degree of recovery (i.e., tree closeness) can affect the number of beetle species. Specifically, we found (1) more xerophilous species in the open woodland and (2) more mesophilous species in the dense mixed woodland.

     

Diameter of downed woody debris does matter for saproxylic beetle assemblages in temperate oak and pine forests

Deadwood is widely recognized to be an important issue for biodiversity conservation in forest ecosystems. Establishing guidelines for its management requires a better understanding of relationships between woody debris characteristics and associated species assemblages. Although deadwood diameter has been identified as an important factor predicting occurrence of many saproxylic species, the boundary between small and large diameter has not yet been precisely defined. In commercial forests, it is also of critical importance to know which diameter is large enough to host the beetle species associated with large logs in order to ensure cost-effectiveness of biodiversity conservation measures. We investigated the differences in saproxylic beetle assemblages among four different diameter classes of downed woody oak and maritime pine debris, in France. Beetles were sampled using in situ emergence traps. The diameter of deadwood pieces ranged from 1 to 40 cm. No patterns of nestedness associated with the gradient of diameter size were identified for either tree species. More indicator saproxylic species were observed in large logs and branches than in small logs. A clear distinction appeared in assemblage composition around the 5-cm diameter threshold whereas no similar pattern occurred around the 10 cm value, i.e. the classical threshold used in forestry to distinguish fine woody debris from coarse woody debris. For both tree species, the mean body length of beetles increased with the diameter of deadwood suggesting that the quantity of available resources per piece may constitute a limiting factor for large beetle species. This study confirms that not only large deadwood pieces are relevant for saproxylic biodiversity conservation but also the smallest pieces. Therefore, forest managers would be well advised to maintain a high diversity of deadwoods to maintain saproxylic biodiversity.     

Space use of the Siberian flying squirrel Ptevomys volans in fragmented forest landscapes

We studied the space use of the Siberian flying squirrel in fragmented forest landscapes in southern Finland in 1996- 1999. Twenty-four adult males and 23 adult females were radio trucked in three different study areas. The use of different landscape elements by flying squirrels was compared to their availability using compositional analysis. Effects of fragmentation on flying squirrels were studied by comparing the use of different habitat types, number of used patches, and size of home range to the principal components formed from the landscape variables patch size, patch isolation, area of spruce-dominated forests, and area of other habitats with trees.
Flying squirrels preferred spruce- and deciduous-dominated forests, but other habitats with trees were also used for moving and foraging. These less preferred habitats connected spruce and deciduous forest patches in a matrix of open areas. Males used several separate patches of preferred habitat, but females usually stayed in one patch. The spacing behaviour of males was affected by the structure of the landscape in a way that home ranges were larger and males used more spruce patches in the more highly fragmented landscape. Home-range size of females was not affected by the landscape structure. Basically, the space use of males seemed lo be reflected by the location and number of female home ranges. In the study area where female density was high, the home ranges of males were small. Our results give empirical evidence of the effects of habitat loss and landscape connectivity for the Siberian flying squirrel and the implications of these for the conservation of the species.     

Patterns of saproxylic beetle succession in loblolly pine

• 1 Patterns of insect succession in dead wood remain unclear, particularly beyond the first several years of decay. In the present study, saproxylic beetles were sampled from loblolly pine (Pinus taeda L.) logs aged between 1 month and 9 years old using both emergence traps attached to logs in the field and rearing bags in the laboratory.
• 2 Species richness peaked within the first year as a result of a diverse assemblage of bark beetles, wood‐borers and predators associated with young logs. After the phloem phase, there were no significant differences in species richness among decay classes.
• 3 Beetle communities differed significantly among decay classes, with 25 and seven species being significantly associated with young and old logs, respectively.

     

Road size and carrion beetle assemblages in a New York forest

In many parts of the world, roads are the most common causes of forest fragmentation. We know roads can affect wildlife, but understand little the extent to which these effects depend on road type and use. We compared the effects of several road types upon a diverse, carrion frequenting beetle assemblage in rural New York State. We found no consistent effects of distance from road on the diversity, abundance or species density of beetles across road types. However, forests near highways and two-lane paved roads were significantly less diverse than were forests near dirt roads. The reduced diversity of beetles near roads was at least in part due to lower species turnover in space near dirt roads than near either type of paved roads. Our data suggest that all roads are not created equal and that comparably sized minimum-use paved roads have a substantially greater affect on fauna than dirt roads. Highways and two-lane paved roads appear to depress biodiversity even among relatively vagile animals like beetles.     

Mammal assemblages in forest fragments and landscapes occupied by black howler monkeys

Species assemblages in disturbed habitats vary as a function of the interaction between species requirements and the spatial configuration of the habitat. There are many reports accounting for the presence of howler monkeys in fragments where other mammals are absent, suggesting that they are more resilient. In the present study we explored this idea and predicted that if howler monkeys were more resilient to habitat loss and fragmentation than other mammals, mammal assemblages in fragments occupied by howler monkeys should include fewer species with decreasing amount of habitat (smaller fragment size and less habitat in the landscape) and increasing number of forest fragments. We explored these relationships by additionally considering the feeding and life habits of mammal species, as well as the isolation and proximity of each fragment to human settlements and roads. We sampled the presence of mammals in five fragments occupied by black howler monkeys (Alouatta pigra) in the Mexican state of Campeche. Through direct sights performed during 240 h in each fragment, we observed 23 species. At the landscape scale, higher fragmentation was associated with a decrease in herbivores, omnivores and total number of species. At the fragment scale semiarboreal, omnivore, and total number of species increased with increasing fragment size. This study supports the idea that howler monkeys are more resilient to forest loss and fragmentation than other native mammals, and our exploratory analyses suggest that the specific mammal assemblages that are found in fragments are related to both landscape and fragment scale spatial attributes, as well as with species-specific characteristics.     

Biodiversity in agricultural landscapes: the ground beetle communities of woody uncultivated habitats

Agricultural landscapes can be defined as mosaics of landscape elements which are affected by farming practices. Woodland habitats, even though they are managed, are amongst the most stable elements of agricultural landscapes and can play a key role in the maintenance of biodiversity. This study of the ground beetle (carabid) communities of woodlands and woody linear features in a Scottish agricultural landscape shows that these habitats contribute significantly to the overall landscape diversity of these beetles. Communities in woods and hedgerows display the same species diversity and are both characterized by the presence of forest species. The main factors constraining carabid communities in both environments are the grazing intensity and, to a lesser extent, the type of soil. Heavily grazed locations are characterized by the occurrence of grassland species while forest species are restricted to ungrazed locations. At the landscape scale, the distribution of the forest species is limited by spatial isolation, indicating that there are insufficient functional links between woodland habitats in the study area. Isolation could be compensated for either by a better control of grazing so that linear features can be used as dispersal corridors for forest carabids or by planting more linear features and woods in the area.     

From logs to landscapes: determining the scale of ecological processes affecting the incidence of a saproxylic beetle

1. Species incidence is influenced by environmental and intrinsic factors operating at multiple scales. The incidence of a dispersal‐limited beetle, Odontotaenius disjunctus (Coleoptera: Passalidae), was surveyed within hierarchically nested organisational levels of its environment (log sections < logs < 10‐m radius subplots < 0.66‐ha plots) in Louisiana, U.S.A. The finest level was the size of a single territory. Passalid beetles are an ecologically prominent group, but little is known of the factors affecting their incidence. 2. Three scale‐sensitive aspects of O. disjunctus incidence were evaluated: (i) the extent (52–3600 ha) within which forest cover was most associated with incidence; (ii) the hierarchical level at which environmental variables best predicted incidence; and (iii) the hierarchical level at which incidence exhibited the greatest spatial autocorrelation as a result of intrinsic factors (e.g. dispersal limitation). 3. Forest cover best predicted incidence at 225 ha, but accounted for only 1.2% of variation in incidence. Incidence was most sensitive to environmental factors measured at the finest scale (i.e. territories). Incidence was positively associated with moderately decayed wood and increased surface area of logs (9.9% and 3.1% of variance, respectively). When environmental factors were accounted for, spatial autocorrelation in incidence was greatest within subplots and logs, consistent with the hypothesis that intrinsic autocorrelation is associated with O. disjunctus average dispersal distance (<5 m). 4. This study indicates the influences of factors acting at multiple scales, but suggests that environmental conditions at the scale of territories may be most important for species incidence.     

Barriers to dispersal of rain forest butterflies in tropical agricultural landscapes

Fragmentation of natural habitats can be detrimental for species if individuals fail to cross habitat boundaries to reach new locations, thereby reducing functional connectivity. Connectivity is crucial for species shifting their ranges under climate change, making it important to understand factors that might prevent movement through human‐modified landscapes. In tropical regions, rain forests are being fragmented by agricultural expansion, potentially isolating populations of highly diverse forest‐dependent species. The likelihood of crossing habitat boundaries is an important determinant of species dispersal through fragmented landscapes, and so we examined movement across rain forest‐oil palm plantation boundaries on Borneo by using relatively mobile nymphalid butterflies as our model study taxon. We marked 1666 individuals from 65 species, and 19 percent (100/527) of recaptured individuals crossed the boundary. Boundary crossing was relatively frequent in some species, and net movement of individuals was from forest into plantation. However, boundary crossing from forest into plantation was detected in less than 50 percent (12/28) of recaptured species and was dominated by small‐sized butterfly species whose larval host plants occurred within plantations. Thus, while oil palm plantations may be relatively permeable to some species, they may act as barriers to the movement of forest‐dependent species (i.e., species that require rain forest habitat to breed), highlighting the importance of maintaining forest connectivity for conserving rain forest species.     

Effects of different tree harvesting intensities on saproxylic beetle diversity in coniferous and broad-leaved mixed forest in the Changbai Mountain,Northeast, China

1. Saproxylic beetles have gained increasing attention due to their role in the decomposition of rotting wood in forests. Studying the response of saproxylic beetles to tree harvesting is important for developing harvesting strategies that consider conservation of saproxylic beetle diversity.
2. We report results from a case study in which we designed four treatment stands to test the effects of forest harvesting intensity on saproxylic beetle diversity, harvest intensities of 0% untreated control (CK), 17.2% light harvest (LT), 34.7% moderate harvest (MT) and 51.9% high harvest (HT). Flight intercept traps were used to collect specimens of saproxylic beetles in each stand at 2, 3 and 4 years post-harvest.
3. The richness and abundance of saproxylic beetles were higher in MT and HT than in CK. Twelve of the 15 indicator species were significantly associated with MT or HT, whereas only three species were significantly associated with CK.
4. We found that moderate and high intensity harvesting affected the composition and increased beetle abundance and richness of saproxylic beetles, and light intensity harvesting had no effect on the beetle community in years 2, 3 and 4 post-harvest. There was no difference in the beetle community composition between moderate and high harvest stands.