Multi-species spatially-explicit indicators reveal spatially structured trends in bird communities

Multi-species indicators are often used to assess biodiversity trends. By combining population trends across several species they summarise trends across a community. Composite indicators such as these are useful for examining general temporal patterns and may suggest important drivers of biodiversity change. However, they may also mask substantial spatial variation in population trends, particularly when they are calculated over large spatial regions. We produced spatially-explicit indicators for farmland and woodland bird communities in the UK and further separate these into trends for generalist and specialist species within each group. We found considerable spatial variation in the indicators, which is masked by indicators calculated at the national level. The farmland community indicator showed mostly positive trends in western areas and extensive declines in south-east England. The woodland community indicator showed a north–south divide, with increases in Scotland and northern England and stability in the southern regions. For both communities, indicator trends for specialist species were more negative than those for generalists. We found no significant difference in farmland community indicators between arable land and improved grassland. Woodland specialists had significantly more negative trends in broadleaf compared to coniferous woodlands, suggesting habitat-type is one of the drivers of changes in the woodland community. These spatial patterns in bird population trends may be used to highlight regional conservation priorities and identify where those may differ from the national scale. In combination with information about other environmental changes, they may also be used to develop hypotheses about potential drivers of change. We advocate that this approach is adopted for other taxa and geographical areas.     

Spatio-temporal organization of the bird communities in two Mediterranean montane forests

Seasonal changes of bird communities and use of foraging substrates by the birds were studied in two forests with contrasting vegetation structure in montane and subalpine levels of central Spanish mountains throughout complete annual cycles. Common factors contributing most to foraging segregation among species were found to be the foraging on the ground vs that on vegetation and use of proximal vs distal tree parts. A complete dissimilarity exists between the composition of the respective foliage-gleaning guilds, the converse being true for the trunk-searchers.
Temporal distribution of bird species is to a great extent determined by their use of space, thus existing a gradient of seasonality with maxima for ground-searchers, minima for trunk-gleaners and intermediate positions for foliage-gleaners. A close relationship among temporal variability, mean annual density and niche breadth is found for the subalpine pinewood community, but not for that from the montane oakwood, this being related with their contrasting migratory patterns and the structural stability of vegetation in each wood. The pattern of covariation in community parameters mirrors the summed response of individual bird species to the changing environmental conditions. Increases in community richness, density and diversity are related to temporal invasion by ground and foliage-gleaners of these highly seasonal Mediterranean habitats.     

Tree and bird functional groups as indicators of recovery of regenerating subtropical coastal dune forests

Functional diversity indicators are increasingly used to monitor forest function recovery because they connect biodiversity to ecosystem functions. However, identifying which functions deviate from a reference forest has not received much attention, despite its potential to inform restoration interventions. In this study, we used functional groups to assess the recovery of ecosystem functions in regenerating coastal dune forests. We surveyed birds and trees in forest of different ages and a reference old‐growth forest in KwaZulu‐Natal, South Africa. We classified species into functional groups for each taxa based on functional traits or a priori defined categories (i.e. guilds) and quantified the number of species within functional groups as a proxy of function stability. Bird species density followed an asymptotic trajectory, reaching old‐growth forest values after 25 years. Insectivores and granivores showed saturating trajectories, whereas small frugivores and generalists increased linearly. With the exception of large frugivores, relative abundances of bird functional groups progressed towards old‐growth forest values as the forest aged. Tree species density increased linearly with forest age. In contrast to old‐growth forests, large‐canopy trees and understory shrubs were under‐represented, while mid‐canopy trees dominated regenerating forests. Our result suggests that most bird, but not tree, functions may have been restored. The trend in large frugivore numbers may warrant further investigation, as their low numbers may have hampered the recovery of tree functions. We conclude that functional group trajectories can track functions that deviate from a benchmark, and may therefore direct adaptive actions to recover the stability of regenerating forest.     

Landscape structure and management regime as indicators of calcareous grassland habitat condition and species diversity

This study investigates the importance of spatial landscape characteristics and habitat management on the condition of calcareous grassland in the North Down Natural Area, Kent UK. We used a digitised map of the study area containing shapefiles of all the habitats including 82 patches of calcareous grassland together with management information for each patch and data on the presence and abundance of a range of calcareous grassland indicator plant species. We defined habitat condition by presence of indicator species and used classification trees to generate models with rules for predicting habitat condition from the landscape spatial characteristics and management information. We also applied the same method to investigate the factors affecting presence or diversity of three ecological groups of positive indicator species and dominance of a negative indicator species. All the models except one showed good classification accuracy and high kappa statistic. Favourable habitat condition was predicted by presence of different types of grazing management, presence of woodland around patches of calcareous grassland and shape complexity. These results indicate that calcareous grassland in favourable condition is management-dependent but also located in less intensively managed landscapes. Unfavourable habitat condition was predicted by threat factors such as lack of management and high incidence of arable or improved grassland around patches of calcareous grassland, indicating nutrient enrichment and habitat degradation. Some of these factors also predicted high diversity of the different ecological species groups. The value of this method for predicting habitat condition and species diversity from baseline ecological data for conservation monitoring at the landscape level is emphasised.     

Landscape heterogeneity metrics as indicators of bird diversity: Determining the optimal spatial scales in different landscapes

Species distribution models are often used to study the biodiversity of ecosystems. The modelling process uses a number of parameters to predict others, such as the occurrence of determinate species, population size, habitat suitability or biodiversity. It is well known that the heterogeneity of landscapes can lead to changes in species’ abundance and biodiversity. However, landscape metrics depend on maps and spatial scales when it comes to undertaking a GIS analysis.We explored the goodness of fit of several models using the metrics of landscape heterogeneity and altitude as predictors of bird diversity in different landscapes and spatial scales. Two variables were used to describe biodiversity: bird richness and trophic level diversity, both of which were obtained from a breeding bird survey by means of point counts. The relationships between biodiversity and landscape metrics were compared using multiple linear regressions. All of the analyses were repeated for 14 different spatial scales and for cultivated, forest and grassland environments to determine the optimal spatial scale for each landscape typology.Our results revealed that the relationships between species’ richness and landscape heterogeneity using 1:10,000 land cover maps were strongest when working on a spatial scale up to a radius of 125–250 m around the sampled point (circa 4.9–19.6 ha). Furthermore, the correlation between measures of landscape heterogeneity and bird diversity was greater in grasslands than in cultivated or forested areas. The multi-spatial scale approach is useful for (a) assessing the accuracy of surrogates of bird diversity in different landscapes and (b) optimizing spatial model procedures for biodiversity mapping, mainly over extensive areas.     

Identifying practical indicators of biodiversity for stand-level management of plantation forests

Identification of valid indicators of biodiversity is a critical need for sustainable forest management. We developed compositional, structural and functional indicators of biodiversity for five taxonomic groups—bryophytes, vascular plants, spiders, hoverflies and birds—using data from 44 Sitka spruce (Picea sitchensis) and ash (Fraxinus excelsior) plantation forests in Ireland. The best structural biodiversity indicator was stand stage, defined using a multivariate classification of forest structure variables. However, biodiversity trends over the forest cycle and between tree species differ among the taxonomic groups studied. Canopy cover was the main structural indicator and affected other structural variables such as cover of lower vegetation layers. Other structural indicators included deadwood and distances to forest edge and to broadleaved woodland. Functional indicators included stand age, site environmental characteristics and management practices. Compositional indicators were limited to more easily identifiable plant and bird species. Our results suggest that the biodiversity of any one of the species groups we surveyed cannot act as a surrogate for all of the other species groups. However, certain subgroups, such as forest bryophytes and saproxylic hoverflies, may be able to act as surrogates for each other. The indicators we have identified should be used together to identify stands of potentially high biodiversity or to evaluate the biodiversity effects of silvicultural management practices. They are readily assessed by non-specialists, ecologically meaningful and applicable over a broad area with similar climate conditions and silvicultural systems. The approach we have used to develop biodiversity indicators, including stand structural types, is widely relevant and can enhance sustainable forest management of plantations.     

Tree diversity in 30-year chronosequences of cool-humid forests

Tree data collected in four inventories of twelve permanent sampling plots during 30 years were analyzed to determine to what extent the abundance of the ubiquitous balsam fir (Abies balsamea) drives tree diversity in mixed stands of Eastern Cape Breton Island, Canada. Dominant deciduous species comprised yellow birch (Betula allegheniensis), sugar maple (Acer saccharum), and American beech (Fagus grandifolia). Tree species richness ranged from 2 to 6. The Shannon–Wiener diversity index indicated that high fir proportion decreased tree species diversity (H’) and structural evenness (J’). The basal area ratio for year 30 over year 1 provided an index of resilience (R) for forest stands subject to natural self-thinning and epidemic insect infestations. There was no significant effect of fir admixture on R since basal area losses due to fir decline were offset within the 30-year period largely by hardwood increases. Stand dynamics were however, structurally stratified due to low R-values for low-diameter trees. The indices H’, J’, and R are recommended for assessing the naturalness and apparent resilience function of tree diversity and large trees in mixed forests of low species richness.     

A typology of indicators of biodiversity change as a tool to make better indicators

Indicators of biodiversity change are of growing political relevance. To be effective, they need to meet a number of scientific requirements, some of which are not easy to fulfil. Here we use a simple typology of biodiversity indicators according to the following two requirements: (1) indicators should reflect changes in biodiversity in general rather than the ups and downs of particular species and species groups, and (2) indicators should be linked to environmental drivers. We apply this typology to existing indicators developed for the Dutch Environmental Data Compendium and describe how the typology may guide the development of indicators of biodiversity change. We conclude that more conceptual and statistical work is needed to make better biodiversity indicators.     

Evaluating the use of Myotis daubentonii as an ecological indicator in Mediterranean riparian habitats

In recent years, interest and concern regarding biodiversity conservation have grown remarkably not only among conservationists but also amongst a wider public beyond scientific institutions. The monitoring of fauna and flora over long periods of time has been satisfactorily proven to be a viable tool for quantifying how environmental changes affect natural communities. Some bat species are regarded as good bioindicators, mainly due to their longevity and high sensitivity to environmental changes. Myotis daubentonii is one of the species most closely associated with riparian habitats in the north-east Iberian Peninsula, and is used as an ecological indicator in specific monitoring programs such as the Waterway Survey (United Kingdom) and the QuiroRius (Spain). Nonetheless, there is still great controversy as to whether M. daubentonii is a good biological indicator or not. While some authors accept it as a bioindicator, others point to the studies carried out in the U.K., Poland, Switzerland and Germany that show a remarkable increase in the numbers of this bat when pollution increases in canalized rivers, which suggest that it is in fact a generalist species.Due to the lack of information regarding habitat-quality requirements in Daubenton’s bats in the Mediterranean region and the species’ potential as a bioindicator in riparian habitats, we aimed to 1) examine how QuiroRius data match other well-established biological indicators (IBMWP for invertebrates and QBR for riparian forests); 2) analyse how environmental variables at both local and landscape scales affect the presence of M. daubentonii; and 3) describe how environmental traits influence the relative abundance of M. daubentonii.A total of 104 streams below 1000 m a.s.l. were simultaneously sampled using bat, macroinvertebrate and vegetation bioindicators. Despite having similar conservation aims, these three bioindicators did not provide consistent images of overall ecosystem quality and thus a multidisciplinary approach is necessary for a full analysis of the health of these riparian ecosystems. M. daubentonii were found more frequently in wide rivers with well-structured native riparian forests; on the other hand, landscape composition at broader scales and altitude had no influence on bat presence/abundance.Thus, we suggest that QuiroRius could be used as a complementary bioindicator for analysing riparian forest quality but cannot be used alone as a tool for evaluating correctly overall riparian ecosystem health. Both relative abundance and/or presence/absence could be used as bioindicator surrogates given that the effect of microhabitat environmental predictors had similar impact on both these measures.     

地中海沿岸沙丘微生境对幼苗出现时空格局的影响

实验样地设在地中海沿岸沙丘,选择了3个不同的微生境代表：(1)稳定沙丘上的开阔地片段,(2)稳定沙丘上的灌丛下区域,(3)不稳定沙丘的路径区域;从2001年11月至2002年4月的整个生长季节,每一个微生境出现的幼苗在4个日期被监测,并在每一个取样日把每一株幼苗鉴定、计数后,用剪刀把地上部分移去;研究调查了3种微生境幼苗出现的时空分布格局,并分析了雨量与幼苗出现数量的关系。结果发现：在地中海沿岸沙丘生态系统,幼苗出现在时间上具有明显的分布特征,大多数幼苗出现在第一观测期,整个生长季幼苗都不断出现,但幼苗出现的数目却逐渐下降;各功能群的幼苗占幼苗总数的比例分别为：1年生阔叶草47．4％,多年生阔叶草2．5％,豆科植物17,0％,菊科植物14．5％,1年生禾草为11．7％,多年生禾草为1．9％,十字花科植物3．7％,伞形科植物1,4％。在空间上,总幼苗密度、物种丰富度和物种多样性等显示出重要的微生境差异,开阔地区域具有最大的幼苗密度、物种丰富度和物种多样性;3个微生境的幼苗出现不是同步的,微生境影响种子萌发的时间分布格局,灌丛下种子萌发具有滞后现象;大多数功能群的幼苗密度分布基本上具有显著的微生境差异,主要物种的幼苗分布也具有显著的微生境差异。雨量和萌发的幼苗数量间未发现显著的关系。     

Hydraulic microhabitats and the distribution of macroinvertebrate assemblages in riffles

1. The influence of hydraulic conditions on the spatial distribution of macroinvertebrate assemblages was investigated in three riffles in a perennial Australian river. 2. Velocity, depth and variability of substrate roughness were measured at each of 56 macroinvertebrate sampling locations. Complex hydraulic variables (roughness Reynolds number, shear velocity, Froude number) were calculated from combinations of two or three of the directly measured variables. The biological significance of directly measured and complex hydraulic variables was determined by a combination of univariate and multivariate statistical procedures. 3. Macroinvertebrate abundance, number of taxa and community composition were significantly different between the identified roughness Reynolds number, Froude number, velocity and shear velocity microhabitats throughout the studied riffles. 4. Regression analysis showed macroinvertebrate abundance and number of taxa were negatively related to roughness Reynolds number, shear velocity, velocity and Froude number. Depth was negatively related to abundance. In general, the majority of the macroinvertebrate community preferred the areas of riffles with the lowest near‐bed turbulence. 5. Roughness Reynolds number explained more of the spatial variation in invertebrate abundance, number of taxa and community composition than the other hydraulic variables, either directly measured or calculated. Of the directly measured variables, velocity had the greatest explanatory power, which was marginally less than roughness Reynolds number and shear velocity. 6. This study demonstrated that small‐scale differences in hydraulic conditions created by combinations of velocity, depth and substrate roughness have an important role in the spatial distribution of macroinvertebrate assemblages in riffle habitats.     

Functional homogenization of bird communities along habitat gradients: accounting for niche multidimensionality

Aim To test whether functional homogenization of bird communities is promoted by anthropogenic landscape transformation, using specialization and habitat preference indices that account for the multidimensionality of niches. Location Catalonia, north‐east Iberian Peninsula. Methods We used data on bird species occurrences and landscape features in 2834 1‐km² squares. Three orthogonal landscape gradients, which were taken as niche dimensions, were defined by means of principal components analysis (PCA). Specialization and habitat preference indices were created for 103 terrestrial bird species on the basis of their frequency of occurrence variation along the landscape gradients. These indices, together with species rarity, were then averaged for bird communities. We then analysed the patterns of variation of communities’ mean specialization, mean rarity and mean habitat preference values along a gradient of agricultural–forest habitat mosaics. Results Wherever we found a significant variation in the degree of specialization along the agricultural–forest gradient, agricultural habitats held more specialized bird communities than did forest ones and bore, on average, rarer species. Thus, results contradicted our initial hypothesis that humanized areas would bear more functionally homogenized bird communities. Higher α‐diversity values tended to be associated with generalist communities and with those having rarer species. Main conclusions Estimations of bird community specialization for different niche dimensions can behave differently along certain landscape gradients, and some of these differences can be explained by the variation of mean habitat preferences. Thus, we argue that a multidimensional approach to assess average niche breadth of communities can be more informative than a unidimensional measure. Our results suggest that widespread land abandonment and current secondary forest expansion throughout the Mediterranean area are promoting functional homogenization of bird communities. It would be desirable to construct larger‐scale indicators of functional homogenization in order to monitor communities’ responses to widespread landscape changes.     

Relations between forest management,stand structure and productivity across different types of Central European forests

In the past 30 years, many stand structural attributes (SSAs) have been suggested and structural indices have been developed to describe the complex structure of forests. Most studies, however, have explored the potential and limits of structural measures to quantify forest structures by applying multiple measures to one stand or few measures to several stands. However, the interdependencies of multiple structural attributes across many stands of different forest management types and developmental stages have not yet been explored. Using 20 structural attributes and 124 completely inventoried 1 ha sized stands we tested to what extent structural characteristics reflect different stand types and management intensities, and how these characteristics change over time. We found that single SSAs do not show the clear gradients that they were intended to reflect, suggesting that stand structure should be described by multiple structural attributes, and that these should represent different structural aspects (including vertical, and horizontal heterogeneity, density, and diversity). A principal component analysis showed that combining several SSAs, allowed us to distinguish between stand types. The structure of mature stands remained rather constant over the observed period of about 6 years, while that of young stands changed more rapidly due to ingrowth and mortality. The older the stands, the less the large trees contributed to stand growth relative to their size. We conclude that multiple stand structural attributes are needed to characterise stand types, management effects and to explain stand productivity.     

Multimodality in the individual size distributions of bird communities

Aim Body size often plays a significant role in community assembly through its impacts on the life history and ecological attributes of species. Insight into the importance of size in structuring communities can be gained by examining the distribution of sizes of individuals [i.e. the individual size distribution (ISD) or size spectrum] in a community. ISDs have been studied extensively in aquatic and tree communities, but have received little attention in terrestrial animal communities. Here, we conduct the first macroecological analysis of ISDs in terrestrial animal communities to determine whether they show broad‐scale consistency in shape. Location North America, north of Mexico. Methods Using likelihood‐based methods and Gaussian mixture modelling, coupled with data from the Breeding Bird Survey and Christmas Bird Count, we determine whether the ISDs for thousands of breeding and wintering North American bird communities are: (1) monotonically decreasing, (2) unimodal or (3) multimodal. Results We find that avian ISDs are consistently multimodal, with most characterized by more than five modes in both breeding and wintering communities from local to continental scales. In addition, the positions of these modes along the size axis are remarkably consistent. Main conclusions The striking consistency in the ISD within bird communities, as with tree and aquatic communities, indicates that the ISD is an important and informative characterization of resource utilization within an ecological assemblage. The differences in shape of the ISD among these groups also suggest that differences in body size‐related constraints affect interactions within a group and with the environment. Our results confirm that avian assemblages do exhibit structure along the body size axis, and therefore it will be fruitful to explore this pattern in greater detail.     

Dung beetle communities as biological indicators of riparian forest widths in southern Brazil

Riparian forests provide important habitat for many wildlife species and are sensitive to landscape change. Among terrestrial invertebrates, dung beetles have been used to investigate the effects of environmental disturbances on forest structure and diversity. Since many studies demonstrated a negative response of dung beetle communities to increasing forest fragmentation, and that most dung beetle species had a more pronounced occurrence during warmest seasons, three hypotheses were tested: (1) Scarabaeinae richness, abundance, diversity and evenness are lower in thinner riparian zone widths than in wider widths during the warmest seasons; (2) Scarabaeinae richness and abundance are positively influenced by leaf litter coverage and height and canopy cover; and (3) Scarabaeinae composition varies with the reduction in riparian vegetation and among annual seasons. We selected four fragments with different riparian zone widths in three secondary streams in southern Brazil. In each fragment, four sampling periods were carried out seasonally between spring 2010 and winter 2011. We collected dung beetles using pitfall traps with two types of bait. We collected 1289 specimens distributed among 29 species. In spring and summer, dung beetle richness was higher in fragments with the widest riparian zone than in those with a thinner riparian zone, and it did not vary between fragments in fall and winter seasons. Dung beetle abundance did not differ among fragments with different riparian zone widths, but it was higher in spring and summer than fall and winter. Richness and abundance were positively influenced by leaf litter. While dung beetle diversity was higher in fragments with wider riparian zone widths than in those with thinner widths, the evenness was similar among fragments. Dung beetle composition differed between the fragments with the widest and thinnest riparian zones, and it also varied among the seasons. Our results suggest that decreased riparian zones affect negatively to dung beetle community structure in southern Brazil. Fragments with thinner riparian zones had lower beetle richness in warmest seasons and an altered community composition. In this sense, the dung beetles are potentially good indicators of riparian forest fragmentation since some species were indicators of a particular riparian zone width. From a conservation perspective, our results demonstrate that the new Brazilian Forest Code will greatly jeopardize not only the terrestrial and aquatic biodiversity of these ecosystems, but also countless other ecological functions.     

The potential of military training areas for bird conservation in a central European landscape

European biodiversity has suffered from serious declines during the past few decades, with alterations of land use practices resulting in a loss of fine-scale habitat heterogeneity being a dominant driver. This heterogeneity was maintained by extensive landscape management, which has gradually been replaced by either intensive exploitation or land abandonment. It has been suggested that military training can generate habitat heterogeneity that may support the existence of species of conservation concern, but studies rigorously testing the real importance of military training areas for biodiversity are lacking. Here we address this issue by analyses of two datasets. First, we compared land cover classes between all large military training areas (MTAs) and surrounding control areas (CAs) of the same size in the Czech Republic using multivariate redundancy analysis. We found that the difference in land cover between MTAs and CAs was significant and represented the strongest gradient in land cover classes: from various farmland and artificial habitats typical for CAs to forest and scrubland-grassland mosaic typical for MTAs. Second, we selected one of these areas and compared bird species richness between the MTA and the nearby CA using generalized linear mixed effects models. We found that the number of species of conservation concern was significantly higher in the MTA than in the CA. With respect to habitats, bird species richness was significantly higher in the MTA than in the CA for open habitats, but not for forest habitats. Our results are thus consistent with the view that military training creates areas that are different from the surrounding landscape in terms of land cover, and that this difference translates to a suitability for species of conservation concern. It is remarkable that the positive influence of military training is confined to open habitats, which are subject to the most intensive military activities and also suffer the highest degree of deterioration in European landscapes.