More and more generalists: two decades of changes in the European avifauna

Biotic homogenization (BH) is a process whereby some species (losers) are systematically replaced by others (winners). While this process has been related to the effects of anthropogenic activities, whether and how BH is occurring across regions and the role of native species as a driver of BH has hardly been investigated. Here, we examine the trend in the community specialization index (CSI) for 234 native species of breeding birds at 10 111 sites in six European countries from 1990 to 2008. Unlike many BH studies, CSI uses abundance information to estimate the balance between generalist and specialist species in local assemblages. We show that bird communities are more and more composed of native generalist species across regions, revealing a strong, ongoing BH process. Our result suggests a rapid and non-random change in community composition at a continental scale is occurring, most likely driven by anthropogenic activities.     

Contrasting effects of habitat reduction,conversion and alteration on neutral and non neutral biological communities

While habitat transformation driven by human activities is the main driver of current biodiversity changes, there is still no framework to explore and forecast the effects of different types of habitat changes on the richness and composition of biological communities. To tackle this issue, we modeled the dynamics of a regional meta‐community, composed either of ecologically equivalent species (neutral model) or of generalist and specialist species (specialization model), and explored the impact of the overall reduction, patch conversion or alteration of an original habitat into one or several other habitats of different total carrying capacity on the community metrics at equilibrium. Our simulations reveal strong interactions between the community model considered (neutral or specialization model) and the type of habitat change. Under neutrality, the impact of habitat changes on richness can be approached by a power law species–individual relationship (SIR), which may at constant density be simplified into the widely used power law species–area relationship (SAR), independent of the type of change. However, in the presence of generalist and specialist species, we found that 1) while habitat reduction in area also leads to approximately power law SIRs and SARs, 2) patch conversion and alteration have more complex effects on regional species richness, and 3) habitat alteration elicits the functional homogenization of communities, involving a decrease of their average level of specialization.     

When species become generalists: on‐going large‐scale changes in bird habitat specialization

Aim Species specialization is often considered as a stable species characteristic over the short term. However, this assumption has hardly been tested, even though it may impair our ability to track the impoverishment of biodiversity induced by the rapid replacement of specialists by generalists. We tested whether species specialization in birds varied over a short period of time, and assessed whether variations in species specialization influence community‐level metrics of biotic homogenization. Location France. Methods We studied the variations in specialization to habitat closure of 94 bird species over the period 2002–08, accounting for species variations in mean density, habitat preference and migratory status. We then quantified the temporal changes in a community specialization index, which measures functional homogenization. Results Specialization decreased over time for 35 species (37%), while 46 (49%) showed non‐significant negative trends and 13 (14%) had null or non‐significant positive trends. The more a species was specialized at the beginning of the study, the more it generalized. We additionally found that changes in the specialization level were density dependent: 34 species (36%) became more generalist in years of higher densities while only one became more specialized. At the community level, accounting for this inter‐annual variability in species specialization accentuated the trend in the functional homogenization of bird communities. Main conclusions Habitat specialization is a labile ecological trait, which may change in the short term following habitat degradation, density dependence and source–sink dynamics. Accounting for short‐term temporal variations in observed habitat specialization of species can increase our understanding of the effects of global changes on species strategies and community dynamics.     

Spatial segregation of specialists and generalists in bird communities

Each species generally has a close relationship with one or more habitats and can therefore be classified as either specialist or generalist. We studied whether specialist and generalist species are spatially distributed independently of each other. Repeating the analysis for 100 of the most frequent terrestrial bird species recorded over the 10 000 sampled sites of the French Breeding Bird survey, we found that specialists were more abundant if the rest of the community was specialized, and that the inverse was also true. This pattern was far subtler than just a simple dichotomy: most species actually presented a maximum abundance at a value of community specialization similar to their own level of specialization. Bird communities appear very well defined along a specialist–generalist gradient. We believe this pattern becomes more apparent with habitat degradation. The consequences on both ecological services and community resilience may well be considerable.     

Rise of the generalists: evidence for climate driven homogenization in avian communities

Aims Biogeographical evidence suggests a strong link between climate and patterns of species diversity, and climate change is known to cause range shifts. However, there is little understanding of how shifts affect community composition and we lack empirical evidence of recent impacts of climate change on the diversity of vertebrates. Using a long‐term comprehensive dataset on bird abundance, we explore recent patterns of change in different components of species diversity and avian communities, and postulate a process to explain the observed changes in diversity and specialization. Location Britain. Methods We used Breeding Bird Survey data for Britain from 1994 to 2006 to calculate site‐specific diversity and community specialization indices. We modelled these indices using generalized additive models to examine the relationship between local climate and spatial and temporal trends in community metrics and the relationship between changes in diversity and specialization. Results Local temperature was positively associated with alpha diversity, which increased over the study period, supporting empirical and theoretical predictions of the effect of climate warming. Diversity increased in all habitats, but the rate of increase was greatest in upland areas. However, temperature was negatively associated with community specialization indices, which declined over the same period. Our modelling revealed a nonlinear relationship between community specialization and species diversity. Main conclusions Our models of diversity and specialization provide stark empirical evidence for a link between warming climate and community homogenization. Over a 13‐year period of warming temperatures, diversity indices increased while average community specialization decreased. We suggest that the observed diversity increases were most likely driven by range expansion of generalist species and that future warming is likely to increase homogenization of community structure. When assessed in combination, diversity and specialization measures provide a powerful index for monitoring the impacts of climate change.     

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.     

Breakdown of the species-area relationship in exotic but not in native forest patches

We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothesized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns.     

Changes in phytophagous insect host ranges following the invasion of their community: Long‐term data for fruit flies

The invasion of an established community by new species can trigger changes in community structure. Invasions often occur in phytophagous insect communities, the dynamics of which are driven by the structure of the host assemblage and the presence of competitors. In this study, we investigated how a community established through successive invasions changed over time, taking the last invasion as the reference. The community included four generalist and four specialist species of Tephritidae fruit flies. We analyzed a long‐term database recording observed numbers of flies per fruit for each species on 36 host plants, over 18 years, from 1991 to 2009. Community structure before the last invasion by Bactrocera zonata in 2000 was described in relation to host plant phylogeny and resource availability. Changes in the host range of each species after the arrival of B. zonata were then documented by calculating diversity indices. The flies in the community occupied three types of niches defined on the basis of plant phylogeny (generalists, Solanaceae specialist, and Cucurbitaceae specialists). After the arrival of B. zonata, no change in the host range of specialist species was observed. However, the host ranges of two generalist species, Ceratitis quilicii and Ceratitis capitata, tended to shrink, as shown by the decreases in species richness and host plant α‐diversity. Our study shows increased host specialization by generalist phytophagous insects in the field following the arrival of an invasive species sharing part of their resources. These findings could be used to improve predictions of new interactions between invaders and recipient communities.     

Indicator species for avian biodiversity hotspots: Combination of specialists and generalists is necessary in less natural environments

In this work, I tested the premise that the distribution of a group of few common bird species can be used to predict bird species hotspots in Central Italy. The data on bird observations were collected on 530 sampled sites (150 in cultivated, 150 in forest, 150 in grassland and 80 in urban and peri-urban environments). In each environment, sampled sites with values of bird species richness in the upper than third quartile were classified as high species richness spots (HSRS), while sites with lower bird species richness were classified as non-HSRS (binary classification system).Generalized Linear Models (GLM) were applied using HSRS or non-HSRS as binomial response variable and bird species occurrence was used as the predictor variable. All selected models showed “fair” or “good” capacities to predict the avian hotspots, using only few common birds (4–6) species. However, bird species selected as predictors were different on each environment. In more natural environments (grassland, forest), specialist species were selected, while in most disturbed environments (cultivated and urban) both generalist and specialist species were selected. The results are in agreement with other studies which show how homogenization of bird communities is strongly correlated to landscape disturbance. The findings supports the hypothesis that indicators have to incorporate both specialists and generalist’s species simultaneously. Furthermore, the groups of birds selected as surrogates are easy to detect and this makes it possible to involve citizen-science programmes in obtain data. This approach can be a cheap and efficient and can help to significantly speed up the process of assessing ecosystems that might be under threat.     

Biodiversity and community structure of temperate butterfly species within a gradient of human disturbance: An analysis based on the concept of generalist vs. Specialist strategies

We monitored nine butterfly communities with varying degrees of human disturbance by conducting a census twice a month during 1980 by the line transect method in and around Tsukuba City, central Japan. We analyzed the biodiversity and community structures using the generalist/specialist concept. The site (community) order based on decreasing human disturbance was positively correlated with butterfly species diversity (H′), species richness (the total number of species), and the number of specialist species in a community, but not with the number of generalist species. The number of generalist species was rather constant, irrespective of the degree of human disturbance. Thus, both the butterfly species diversity and species richness were more dependent on the specialists than the generalists. Our analyses also showed that the generalist species were distributed widely over the communities, and they maintained high population densities, resulting in high rank status in abundance in a community, with more spatial variation in density per species. Specialist species showed the opposite trends. These results demonstrate that the generalist/specialist concept is a powerful tool applicable to analyse the biodiversity and structure of natural communities.     

Functional biotic homogenization of bird communities in disturbed landscapes

Aim  Worldwide, functional homogenization is now considered to be one of the most prominent forms of biotic impoverishment induced by current global changes. Yet this process has hardly been quantified on a large scale through simple indices, and the connection between landscape disturbance and functional homogenization has hardly been established. Here we test whether changes in land use and landscape fragmentation are associated with functional homogenization of bird communities at a national scale.
Location  France.
Methods  We estimated functional homogenization of a community as the average specialization of the species present in that community. We studied the spatial variation of this community specialization index (CSI) using 1028 replicates from the French Breeding Bird Survey along spatial gradients of landscape fragmentation and recent landscape disturbance, measured independently, and accounting for spatial autocorrelation.
Results  The CSI was very sensitive to both measures of environmental degradation: on average, 23% of the difference in the CSI values between two sample sites was attributed to the difference in fragmentation and the disturbance between sites. This negative correlation between CSI and sources of landscape degradation was consistent over various habitats and biogeographical zones.
Main conclusions  We demonstrate that the functional homogenization of bird communities is strongly positively correlated to landscape disturbance and fragmentation. We suggest that the CSI is particularly effective for measuring functional homogenization on both local and global scales for any sort of organism and with abundance or presence–absence data.     

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.     

Habitat quality is more important than matrix quality for bird communities in protected areas

Protected areas are meant to preserve native local communities within their boundaries, but they are not independent from their surroundings. Impoverished habitat quality in the matrix might influence the species composition within the protected areas through biotic homogenization. The aim of this study was to determine the impacts of matrix quality on species richness and trait composition of bird communities from the Finnish reserve area network and whether the communities are being subject of biotic homogenization due to the lowered quality of the landscape matrix. We used joint species distribution modeling to study how characteristics of the Finnish forest reserves and the quality of their surrounding matrix alter species and trait compositions of forest birds. The proportion of old forest within the reserves was the main factor in explaining the bird community composition, and the bird communities within the reserves did not strongly depend on the quality of the matrix. Yet, in line with the homogenization theory, the beta‐diversity within reserves embedded in low‐quality matrix was lower than that in high‐quality matrix, and the average abundance of regionally abundant species was higher. Influence of habitat quality on bird community composition was largely explained by the species' functional traits. Most importantly, the community specialization index was low, and average body size was high in areas with low proportion of old forest. We conclude that for conserving local bird communities in northern Finnish protected forests, it is currently more important to improve or maintain habitat quality within the reserves than in the surrounding matrix. Nevertheless, we found signals of bird community homogenization, and thus, activities that decrease the quality of the matrix are a threat for bird communities.     

Comparison of the ecology and evolution of plants with a generalist bird pollination system between continents and islands worldwide

Thousands of plant species worldwide are dependent on birds for pollination. While the ecology and evolution of interactions between specialist nectarivorous birds and the plants they pollinate is relatively well understood, very little is known on pollination by generalist birds. The flower characters of this pollination syndrome are clearly defined but the geographical distribution patterns, habitat preferences and ecological factors driving the evolution of generalist‐bird‐pollinated plant species have never been analysed. Herein I provide an overview, compare the distribution of character states for plants growing on continents with those occurring on oceanic islands and discuss the environmental factors driving the evolution of both groups. The ecological niches of generalist‐bird‐pollinated plant species differ: on continents these plants mainly occur in habitats with pronounced climatic seasonality whereas on islands generalist‐bird‐pollinated plant species mainly occur in evergreen forests. Further, on continents generalist‐bird‐pollinated plant species are mostly shrubs and other large woody species producing numerous flowers with a self‐incompatible reproductive system, while on islands they are mostly small shrubs producing fewer flowers and are self‐compatible. This difference in character states indicates that diverging ecological factors are likely to have driven the evolution of these groups: on continents, plants that evolved generalist bird pollination escape from pollinator groups that tend to maintain self‐pollination by installing feeding territories in single flowering trees or shrubs, such as social bees or specialist nectarivorous birds. This pattern is more pronounced in the New compared to the Old World. By contrast, on islands, plants evolved generalist bird pollination as an adaptation to birds as a reliable pollinator group, a pattern previously known from plants pollinated by specialist nectarivorous birds in tropical mountain ranges. Additionally, I discuss the evolutionary origins of bird pollination systems in comparison to systems involving specialist nectarivorous birds and reconstruct the bird pollination system of Hawaii, which may represent an intermediate between a specialist and generalist bird pollination system. I also discuss the interesting case of Australia, where it is difficult to distinguish between specialist and generalist bird pollination systems.     

Measuring community responses to large-scale disturbance in conservation biogeography

Aim   Which community metrics should be used to reflect community response to large-scale habitat alterations is unclear. Here, we assess what and how community changes should be measured to accurately track community responses to large-scale disturbance in space and/or time.
Location   France.
Method   We first developed a simulation model to examine temporal changes in the species composition of large-scale metacommunities. Using this model, we assessed how species richness, Shannon index, trends of particular subset of species or community indices of habitat specialization were influenced by different disturbance scenarios, and whether these indices were biased by imperfect detectability. We further used more than 1000 empirical bird communities from the French Breeding Bird Survey recently exposed to disturbances of various intensities as a case study.
Results   Our simulation and empirical results both demonstrate that species richness and diversity measures can show confusing trends and even provide misleading messages of communities' fate. In contrast, reflecting the composition of the community in terms of habitat specialist and generalist species was more robust and powerful to reflect disturbance effects.
Main conclusions   We highlight the weakness of using community metrics that fail to incorporate ecological difference among species when summarizing community-level trends in disturbed landscapes.     

Seed dispersal networks in tropical forest fragments: Area effects,remnant species,and interaction diversity

Seed dispersal interactions involve key ecological processes in tropical forests that help to maintain ecosystem functioning. Yet this functionality may be threatened by increasing habitat loss, defaunation, and fragmentation. However, generalist species, and their interactions, can benefit from the habitat change caused by human disturbance while more specialized interactions mostly disappear. Therefore, changes in the structure of the local, within fragment, networks can be expected. Here we investigated how the structure of seed dispersal networks changes along a gradient of increasing habitat fragmentation. We analyzed 16 bird seed dispersal assemblages from forest fragments of a biodiversity-rich ecosystem. We found significant species–, interaction–, and network–area relationships, yet the later was determined by the number of species remaining in each community. The number of frugivorous bird and plant species, their interactions, and the number of links per species decreases as area is lost in the fragmented landscape. In contrast, network nestedness has a negative relationship with fragment area, suggesting an increasing generalization of the network structure in the gradient of fragmentation. Network specialization was not significantly affected by area, indicating that some network properties may be invariant to disturbance. Still, the local extinction of partner species, paralleled by a loss of interactions and specialist–specialist bird–plant seed dispersal associations, suggests the functional homogenization of the system as area is lost. Our study provides empirical evidence for network–area relationships driven by the presence/absence of remnant species and the interactions they perform.     

Habitat associations and breeding bird community composition within the city of Bristol,UK

Capsule Different urban breeding bird communities are associated with different habitat types, but, although community species diversity varies significantly, total bird density does not.

Aims To investigate the association between breeding bird communities and habitats within Bristol, UK and how these communities vary in terms of species diversity and total bird abundance.

Methods Breeding density data for 70 species in the metropolitan area of Bristol, UK were subjected to de‐trended correspondence analysis to identify the number of different communities present and their indicator species. These data were then used to identify patterns of habitat association with each community and differences in species richness and total bird density.

Results Three communities were identified: a rural community associated with woodland, managed grassland and inland water; a suburban community associated with buildings and residential gardens; and an intermediate community that shared some of these habitat characteristics. Species richness, but not total bird abundance, was lowest in the suburban community.

Conclusion The diversity of species in urban areas appears to be most dependent upon the availability of patches of natural and semi‐natural habitats. Residential gardens support fewer species, but those species that are present may be found at high densities.     

Does habitat loss affect the communities of plants and insects equally in plant–pollinator interactions? Preliminary findings

Habitat loss is a major threat to biodiversity and ecosystem function. As habitats are lost, one factor affecting their community structures is the niche-width demand of species, which ranges from specialist to generalist. This study focused on specialist and generalist species in plant–pollinator interactions and tested the hypothesis that plant and pollinator communities become more generalized as habitat loss increases. The study was made in seven selected sites in southern Ontario, Canada, at the level of landscape that is characterized by distributed forests within intensively managed agricultural fields. We quantified both the degree of habitat loss and the degree of specialization/generalization for each of the plant and insect communities using a sampling method of hexagonal transects. Regression analysis indicated a significant relationship between the increase of habitat loss and the shift to generalization in insect, but not in plant, communities. Our results suggest that, in plant–pollinator interactions, insect communities are more sensitive and/or quicker than plant communities to respond to the effects of habitat loss.     

Context-dependent effects of predator removal from experimental microcosm communities

The loss of a predator from an ecological community can cause large changes in community structure and ecosystem processes, or have very little consequence for the remaining species and ecosystem. Understanding when and why the loss of a predator causes large changes in community structure and ecosystem processes is critical for understanding the functional consequences of biodiversity loss. We used experimental microbial communities to investigate how the removal of a large generalist predator affected the extinction frequency, population abundance and total biomass of its prey. We removed this predator in the presence or absence of an alternative, more specialist, predator in order to determine whether the specialist predator affected the outcome of the initial species removal. Removal of the large generalist predator altered some species' populations but many were unaffected and no secondary extinctions were observed. The specialist predator, though rare, altered the response of the prey community to the removal of the large generalist predator. In the absence of the specialist predator, the effects of the removal were only measurable at the level of individual species. However, when the specialist predator was present, the removal of the large generalist predator affected the total biomass of prey species. The results demonstrate that the effect of species loss from high trophic levels may be very context-dependent, as rare species can have disproportionately large effects in food webs.     

Ecological and Evolutionary Effects of Stickleback on Community Structure

Species’ ecology and evolution can have strong effects on communities. Both may change concurrently when species colonize a new ecosystem. We know little, however, about the combined effects of ecological and evolutionary change on community structure. We simultaneously examined the effects of top-predator ecology and evolution on freshwater community parameters using recently evolved generalist and specialist ecotypes of three-spine stickleback (Gasterosteus aculeatus). We used a mesocosm experiment to directly examine the effects of ecological (fish presence and density) and evolutionary (phenotypic diversity and specialization) factors on community structure at lower trophic levels. We evaluated zooplankton biomass and composition, periphyton and phytoplankton chlorophyll-a concentration, and net primary production among treatments containing different densities and diversities of stickleback. Our results showed that both ecological and evolutionary differences in the top-predator affect different aspects of community structure and composition. Community structure, specifically the abundance of organisms at each trophic level, was affected by stickleback presence and density, whereas composition of zooplankton was influenced by stickleback diversity and specialization. Primary productivity, in terms of chlorophyll-a concentration and net primary production was affected by ecological but not evolutionary factors. Our results stress the importance of concurrently evaluating both changes in density and phenotypic diversity on the structure and composition of communities.