Impacts of clearing,fragmentation and disturbance on the bird fauna of Eucalypt savanna woodlands in central Queensland,Australia

Abstract This study reports on the responses of bird assemblages to woodland clearance, fragmentation and habitat disturbance in central Queensland Australia, a region exposed to very high rates of vegetation clearance over the last two to three decades. Many previous studies of clearing impacts have considered situations where there is a very sharp management contrast between uncleared lands and cleared areas: in this situation, the contrast is more muted, because both cleared lands and uncleared savanna woodlands are exposed to cattle grazing, invasion by the exotic grass Cenchrus ciliaris and similar fire management. Bird species richness (at the scale of a 1‐ha quadrat) was least in cleared areas (8.1 species), then regrowth areas (14.6 species), then uncleared woodlands (19.9 species). Richness at this scale was unrelated to woodland fragment size, connectivity or habitat condition; but declined significantly with increasing abundance of miners (interspecifically aggressive colonial honeyeaters). At whole of patch scale, richness increased with fragment size and decreased with abundance of miners. This study demonstrates complex responses of individual bird species to a regional management cocktail of disturbance elements. Of 71 individual bird species modelled for woodland fragment sites, the quadrat‐level abundance of 40 species was significantly related to at least one variable representing environmental position (across a rainfall gradient), fragment condition, fragment size and/or connectivity. This study suggests that priorities for conservation management include: cessation of broad‐scale clearing; increased protection for regrowth (particularly where this may bolster connectivity and/or size of woodland fragments); control of miners; maintenance of fallen woody debris in woodlands; increase in fire frequency; and reduction in the incidence of grazing and exotic pasture grass.     

Bird community responses to habitat fragmentation: how consistent are they across landscapes?

Aim The woodland ecosystems of south‐eastern Australia have been extensively disturbed by agriculture and urbanization. Herein, the occurrence of birds in woodland remnants in three distinct landscapes was analysed to examine the effects of different types of landscape matrices on species richness vs. area and species richness vs. isolation relationships and individual species responses to woodland fragmentation. Location The study system comprised three distinct woodland landscapes of the northern Australian Capital Territory and bordering areas of New South Wales. These landscapes (termed agricultural, peri‐urban and urban) are located within 50 km of each other, have remnant fragments of similar age, size, isolation, woodland cover, elevation and climates. The major distinguishing feature of the three landscapes was the properties of the habitats surrounding the numerous woodland remnants. Methods Bird surveys, using an area‐search methodology, were conducted in 1999 and 2000 in 127 remnants in the three landscapes to determine bird species presence/absence. Each remnant was characterized by measures of remnant area, isolation and habitat complexity. To characterize differences between each landscape, we conducted an analysis of the amount of tree cover and human disturbance in each landscape using SPOT imagery and aerial photographs. Linear regressions of woodland‐dependent species richness vs. remnant area and remnant isolation for the three different landscapes were calculated to see if there were any apparent differences. Binomial logistic regressions were used to determine the relationships between the occurrence of each species and the size and isolation of woodland habitat, in each landscape. Results All the landscapes displayed a significant (P < 0.01) species vs. area relationship, but the slope of the urban relationship was significantly greater than those of the other landscapes. In contrast, only the agricultural landscape displayed a significant (P < 0.01) species richness vs. isolation relationship. When individual species were investigated, we found species that were: (1) apparently insensitive to reduction in remnant area and increase in isolation across all landscapes, (2) absent in small remnants in all landscapes, (3) absent in small remnants in all landscapes and also absent in isolated remnants in the agricultural landscape, (4) absent in isolated remnants in the agricultural landscape, and (5) absent in small remnants in the urban landscape. Threshold values (50% probability of occurrence) for area and isolation for individual species were highly variable across the three landscapes. Main conclusions These results indicate that woodland bird communities have a varying response to habitat fragmentation in different landscapes. Whilst we cannot be sure how representative our chosen landscapes are of other similarly composed landscapes, these results suggest that the type of landscape matrix may have a considerable influence on how bird species are affected by woodland fragmentation in the region. For instance, the properties of a matrix may influence both the resources available in the landscape as a whole for different bird species, and the connectivity (dispersal of birds), between woodland remnants. We encourage further research that examines these hypotheses and argue that the management of the matrix should be included in conservation strategies for fragmented landscapes.     

Explaining bird species composition and richness in eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the factors influencing the distribution of birds in remnants in a fragmented agricultural landscape. Location Forty‐seven eucalypt remnants and six sites in continuous forest in the subhumid Midlands region of Tasmania, Australia. Methods Sites were censused over a two‐year period, and environmental data were collected for remnants. The avifauna of the sites was classified and ordinated. The abundances of bird species, and bird species composition, richness, abundance and diversity were related to environmental variables, using simple correlation and modelling. Results There were two distinct groups of sample sites, which sharply differed in species composition, richness, diversity and bird abundance, separated on the presence/absence of noisy miner (Manorina melanocephala Latham) colonies, remnant size, vegetation structural attributes and variables that reflected disturbance history. The approximate remnant size threshold for the change from one group to another was 20–30 ha. Remnant species richness and diversity were most strongly explained by remnant area and noisy miner abundance, with contributions from structural and isolation attributes in the second case. Segment richness was explained by precipitation, logging history and noisy miner abundance. Bird abundance was positively related to precipitation and negatively related to tree dieback. The 28 individual bird species models were highly individualistic, with vegetation structural variables, noisy miner abundance, climatic variables, variables related to isolation, area, variables related to floristics, disturbance variables, the nature of the matrix and remnant shape all being components in declining order of incidence. Age of the remnant did not relate to any of the dependent variables. Main conclusions Degraded and small remnants may have become more distinct in their avifaunal characteristics than might otherwise be the case, as a result of the establishment of colonies of an aggressive native bird, the noisy miner. The area, isolation and shape of remnants directly relate to the abundance of relatively few species, compared to vegetation attributes, climate and the abundance of the noisy miner. The nature of the matrix is important in the response of some species to fragmentation.     

Synergistic effects of habitat configuration and land-use intensity shape the structure of bird assemblages in human-modified landscapes across three major neotropical biomes

Maximizing biodiversity persistence in heterogeneous human-modified landscapes is hindered by the complex interactions between habitat quality and configuration of native and non-native habitats. Here we examined these complex interactions considering avian diversity across 26 sampling sites, each of which comprised of three sampling points located across a gradient of disturbance: core native habitat fragment, fragment edge, and non-native adjacent matrix. The 78 sampling points were further nested within three neotropical biomes—Amazonia, Cerrado and Pantanal—in central-western Brazil. Matrix type consisted of cattle pastures in the Amazon and teak plantations in the Pantanal and Cerrado. We considered the interactive effects of (1) disturbance-context: fragment core, edge and adjacent matrix, (2) matrix type: tree plantation or cattle pastures, both subject to varying land-use intensity, and (3) native habitat configuration (fragment size, shape and isolation) on bird species richness, abundance and composition. Based on point-count surveys, we recorded 210 bird species. Bird species richness and abundance declined across the disturbance gradient, while genus composition only differed within the adjacent matrix, particularly cattle-pastures. The effect of native habitat area was positive but only detected at fragment edges. Overall bird diversity increased at sites characterized by higher availability of either relict trees within pasture landscapes or old-growth trees within teak plantation landscapes. The core of native fragments played a primary role in ensuring the persistence of bird diversity, regardless of fragment size. In contrast to pastures, tree plantations likely harbour a higher proportion of forest-dependent species while bird diversity can be further enhanced by reduced management intensity in both matrix types. Strategies to maximize avian persistence should not only include retaining native habitats, but also maximizing the size of core native habitats. Likewise, more structurally complex matrix types should be encouraged while maintaining low levels of land-use intensity.

     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

Geography,topography, and history affect realized‐to‐potential tree species richness patterns in Europe

Environmental conditions and biotic interactions are generally thought to influence local species richness. However, immigration and the evolutionary and historical factors that shape regional species pools should also contribute to determining local species richness because local communities arise by assembly from regional species pools. Using the European tree flora as our study system, we implemented a novel approach to assess the relative importance of local and regional mechanisms that control local species richness. We first identified species pools that tolerate particular local environments and quantified the proportion of the pool that is present locally, i.e. the realized/potential (R/P) richness ratio. Because no consensus exists on how to estimate potential richness, we estimated it using three different approaches. Using these three estimates separately and in a combined ensemble estimate, we then analyzed the effects of potential drivers on R/P richness ratios. We predicted that the R/P richness ratio would 1) increase with decreasing distance from glacial refugia (accessibility), 2) and be generally low in geographically fragmented southern Europe because of dispersal limitation; 3) increase with actual evapotranspiration because greater availability of water and energy promotes local population persistence; and 4) increase with topographic heterogeneity because it promotes local species coexistence and facilitates long‐term species survival. There was considerable variation among the three R/P richness ratio estimates, but we found consistent support for a negative effect of regional geographic fragmentation and a positive topographic effect. We also identified fairly broad support for the predicted effect of accessibility. We conclude that local tree assemblages in Europe often fail to realize a large proportion of the potential richness held in the regional species pool, partially reflecting their geographical, historical, and environmental circumstances. The dispersal‐related effects of geographic fragmentation and accessibility exemplify regional controls that combine with local ecological sorting to determine local species richness.     

Long‐term Changes in Bird Communities after Wildfires in the Central Brazilian Amazon

We examined long‐term responses of an Amazonian bird assemblage to wildfire disturbance, investigating how understory birds reacted to forest regeneration 1, 3, and 10 years after a widespread fire event. The bird community was sampled along the Arapiuns and Maró river catchments in central Brazilian Amazonia. Sampling took place in 1998, 2000, and 2008 using mist‐nets in eight plots (four burned, four unburned sites). Species richness did not change significantly in unburned sites. In burned sites, however, we found significantly lower richness in 1998, higher richness in 2000, and similar richness in 2008. Multi‐dimensional scaling ordination showed consistent differences in bird communities both within burned sites sampled in different sampling years, and between burned and unburned sites in all years. Of the 30 most abundant species, 12 had not recovered 10 years after the fires, including habitat specialists such as mixed flocks specialists and ant‐followers. Fire‐disturbance favored three species (two hummingbirds and a manakin) in the short term only. All other species were either favored throughout the study (seven species of omnivores and small insectivores) or did not show a clear response (eight species). In burned sites, we also found significantly lower abundance of species sensitive to disturbances and habitat specialists over the entire study period. Although the bird community seems to be recovering in terms of richness, the overall community composition and abundance of some species in post‐burned and unburned sites remain very different, and have not recovered after 10 years of forest regeneration.     

Factors determining mammal species richness on habitat islands and isolates: habitat diversity, disturbance, species interactions and guild assembly rules

• 1 For over three decades the equilibrium theory of island biogeography has galvanized studies in ecological biogeography. Studies of oceanic islands and of natural habitat islands share some similarities to continental studies, particularly in developed regions where habitat fragmentation results from many land uses. Increasingly, remnant habitat is in the form of isolates created by the clearing and destruction of natural areas. Future evolution of a theory to predict patterns of species abundance may well come from the application of island biogeography to habitat fragments or isolates.
• 2 In this paper we consider four factors other than area and isolation that influence the number and type of mammal species coexisting in one place: habitat diversity, habitat disturbance, species interactions and guild assembly rules. In all examples our data derive from mainland habitat, fragmented to differing degrees, with different levels of isolation.
• 3 Habitat diversity is seen to be a good predictor of species richness. Increased levels of disturbance produce a relatively greater decrease in species richness on smaller than on larger isolates. Species interactions in the recolonization of highly disturbed sites, such as regenerating mined sites, is analogous to island colonization. Species replacement sequences in secondary successions indicate not just how many, but which species are included. Lastly, the complement of species established on islands, or in insular habitats, may be governed by guild assembly rules. These contributions may assist in taking a renewed theory into the new millennium.

     

Site vegetation characteristics are more important than landscape context in determining bird assemblages in revegetation

The effectiveness of revegetation in providing habitat for fauna is expected to be determined both by within‐site factors and attributes of the landscape in which a revegetation site occurs. Most studies of fauna in revegetation have been conducted in landscapes that have been extensively cleared, modified or fragmented, and in Australia, predominantly in the southern temperate zone. We investigated how within‐site vegetation attributes and landscape context attributes were related to bird species richness and composition in a chronosequence of post‐mining rehabilitation sites within an otherwise intact landscape in tropical northern Australia. Our working hypothesis was that bird species richness in rehabilitating sites would be positively related to site vegetation structure and landscape context including (1) proximity to woodland and (2) the proportion of woodland within a 500‐m buffer of rehabilitation sites. Within each of 67 sites, we sampled vegetation once and surveyed for birds eight times over 16 months. Landscape context variables were calculated using GIS. There were large differences between bird assemblages of woodland and rehabilitation sites and between age classes of rehabilitation. Bird assemblages were strongly related to site vegetation attributes across all rehabilitation sites. Proximity to woodland was only related to bird assemblages in rehabilitation sites older than 4 years old. We conclude that the relative importance of landscape context and site variables at any point in time will be a function of how closely vegetation within the revegetation site matches the habitat resource requirements of individual species.     

Do miners (Manorina spp.) affect bird assemblages in continuous savanna woodlands in north‐eastern Australia?

Overabundant native species can have a significant cascading effect on other components of wildlife, and those that deplete other species, often promoted by anthropogenic change to vegetation cover and habitat, are called reverse keystone species. Birds in the genus Manorina are widely reported as being such species, and in highly disturbed or fragmented environments, and some intact environments, noisy miners Manorina melanocephala can have a strong negative effect on small passerine species via hyper‐aggressive mobbing. The tropical savannas of northern Australia consist of largely unmodified woodlands, and two species of Manorina occur naturally in this region: the noisy miner and the yellow‐throated miner Manorina flavigula. Therefore, what effect do these species have on bird assemblage in predominantly continuous habitats, relative to other typical determinants of avifauna assemblage such as vegetation structure? We used data collected from bird surveys at 511 sites across northern Queensland (179 noisy miner M. melanocephala sites, 332 yellow‐throated miner M. flavigula sites) between 1998 and 2010. We examined the variation in bird composition at each site due to increasing abundance of Manorina spp. using uni‐ and multivariate techniques. We found total bird richness was significantly lower in sites where noisy and yellow‐throated miner abundances were highest, and passerine species seemed most affected. For species, 45 species varied significantly in abundance with increasing miner numbers, and the overall effect of yellow‐throated miners on other birds seemed more pronounced. However, vegetation structure was generally an equal or more important predictor of avifauna richness and abundance. We conclude that despite the superficially intact nature of northern Australian woodlands, pastoral intensification or poor land management might create disturbances that facilitate increases in the abundance of Manorina, causing localized overabundance and a compounding negative effect on other native bird species.     

Inferring relationships between native plant diversity and Lonicera japonica in upland forests in north Mississippi,USA

Question: Do anthropogenic disturbances interact with local environmental factors to increase the abundance and frequency of invasive species, which in turn exerts a negative effect on native biodiversity? Location: Mature Quercus‐Carya and Quercus‐Carya‐Pinus (oak‐hickory‐pine) forests in north Mississippi, USA. Methods: We used partial correlation and factor analysis to investigate relationships between native ground cover plant species richness and composition, percent cover of Lonicera japonica, and local and landscape‐level environmental variables and disturbance patterns in mature upland forests. We directly measured vegetation and environmental variables within 34 sampling subplots and quantified the amount of tree cover surrounding our plots using digital color aerial photography. Results: Simple bivariate correlations revealed that high species richness and a high proportion of herbs were associated with low Lonicera japonica cover, moist and sandy uncompacted soils, low disturbance in the surrounding landscape, and periodic prescribed burning. Partial correlations and factor analysis showed that once we accounted for the environmental factors, L japonica cover was the least important predictor of composition and among the least important predictors of species richness. Hence, much of the negative correlation between native species diversity and this invasive species was explained by soil texture and local and landscape‐level land‐use practices. Conclusions: We conclude that negative correlations between the abundance of invasive species and native plant diversity can occur in landscapes with a gradient of human disturbance, regardless of whether there is any negative effect of invasive species on native species.     

Seed‐dispersal interactions in fragmented landscapes – a metanetwork approach

Mutualistic interactions repeatedly preserved across fragmented landscapes can scale‐up to form a spatial metanetwork describing the distribution of interactions across patches. We explored the structure of a bird seed‐dispersal (BSD) metanetwork in 16 Neotropical forest fragments to test whether a distinct subset of BSD‐interactions may mediate landscape functional connectivity. The metanetwork is interaction‐rich, modular and poorly connected, showing high beta‐diversity and turnover of species and interactions. Interactions involving large‐sized species were lost in fragments < 10 000 ha, indicating a strong filtering by habitat fragmentation on the functional diversity of BSD‐interactions. Persistent interactions were performed by small‐seeded, fast growing plant species and by generalist, small‐bodied bird species able to cross the fragmented landscape. This reduced subset of interactions forms the metanetwork components persisting to defaunation and fragmentation, and may generate long‐term deficits of carbon storage while delaying forest regeneration at the landscape level.     

Can agroforest woodlots work as stepping stones for birds in the Atlantic forest region?

In fragmented landscapes, agroforest woodlots can potentially act as stepping stones facilitating movement between forest fragments. We assessed the influence of agroforest woodlots on bird distribution and diversity in the Atlantic forest region (SE Brazil), and also tested which categories of species can use different types of connection elements, and whether this use is influenced by the distance to large forest patches. We studied two fragmented landscapes, with and without stepping stones linking large fragments, and one forested landscape. Using a point count, a bird survey was undertaken in the fragmented landscapes in five different elements: large remnants (>400 ha), agroforest woodlots (0.4–1.1 ha), small patches (0.5–7 ha), riparian corridor, and pasture areas (the main matrix). Generalist and open-area species were commonly observed in the agroforest system or other connection elements, whereas only a few forest species were present in these connections. For the latter species, the distance of woodlots to large patches was essential to determine their richness and abundance. Based on our results and data from literature, we suggest that there is an optimal relationship between the permeability of the matrix and the efficiency of stepping stones, which occurs at intermediate degrees of matrix resistance, and is species-dependent. Because the presence of agroforest system favors a higher richness of generalist species, they appeared to be more advantageous for conservation than the monoculture system; for this reason, they should be considered as a management alternative, particularly when the matrix permeability requirement is met.     

Avian responses to an emergent,wetland weed

African boxthorn (Lycium ferocissimum Solanaceae) is a Weed of National Significance in Australia. It is particularly problematic in Victoria and is thought to not only threaten native wildlife but also provide important habitat, particularly to birds, when there is no native alternative. In a wetland ecosystem such as a saltmarsh, boxthorn has the potential to increase structural complexity because it can stand as an emergent above surrounding vegetation. We compared bird assemblages and behaviour in saltmarsh vegetation with and without boxthorn in a coastal wetland in south‐east Australia. Species assemblage, but not richness, changed with the presence of boxthorn. The presence of singing honeyeaters (Lichenostomus virescens) and white‐fronted chats (Epthianura albifrons), the two most common native bird species (based on numerical and spatial dominance), appeared to drive these differences; singing honeyeaters preferred boxthorn while white‐fronted chats avoided it. The presence of boxthorn increased the seasonal availability of fruit and flowers, which was reflected by a high frequency of foraging for fruit and nectar where boxthorn was present. In saltmarshes without boxthorn, there was a higher frequency of foraging for insects. Some, but not all, species responded to increased structural complexity and fruit/floral resources provided by boxthorn. Consequently, management by reducing boxthorn is likely to alter bird communities and the usage of sites by some native species, thus management success should consider fine‐scale biodiversity objectives, such as managing for particular types or species of birds.     

Responses of plant and bird communities to prescribed burning in tallgrass prairies

Historic losses and fragmentation of tallgrass prairie habitat to agriculture and urban development have led to declines in diversity and abundance of plants and birds associated with such habitat. Prescribed burning is a management strategy that has potential for restoring and rejuvenating prairies in fragmented landscapes, and through such restoration, might create habitat for birds dependent upon prairies. To provide improved data for management decision-making regarding the use of prescribed fire in tallgrass prairies, we compared responses of plant and bird communities on five burned and five unburned tallgrass prairie fragments at the DeSoto National Wildlife Refuge, Iowa, USA, from 1995 to 1997. Overall species richness and diversity were unaffected by burning, but individual species of plants and birds were affected by year-treatment interactions, including northern bobwhite (Colinus virginianus) and ring-necked pheasant (Phasianus colchicus), which showed time-delayed increases in density on burned sites. Analyses of species/area relationships indicated that, collectively, many small sites did make significant contributions to plant biodiversity at landscape levels, supporting the overall conservation value of prairie fragments. In contrast, most birds species were present on larger sites. Thus, higher biodiversity in bird communities which contain area-sensitive species might require larger sites able to support larger, more stable populations, greater habitat heterogeneity, and greater opportunity for niche separation.     

Song parameters of the fuscous honeyeater Lichenostomus fuscus correlate with habitat characteristics in fragmented landscapes

Both avian abundance and species richness decline in response to habitat loss and fragmentation. Studying variation in bird song structure across modified landscapes can provide insights into the effects of habitat alterations on coherence of social interactions within populations. Here, we tested whether fragmentation or change of habitat quality within box‐ironbark forest of central Victoria impacted cultural connectivity and song characteristics in fuscous honeyeater, a declining common Australian bird. First, we tested whether geographic distance and/or spatially‐explicit landscape connectivity models can explain patterns of song similarity across fragmented landscapes. We found no evidence that distance or habitat fragmentation impacts the nature and transmission of fuscous honeyeater song, and concluded that acoustic connectivity at the scale of our study is high. Second, we tested whether variation in habitat quality explains variation in song characteristics. In accordance with acoustic adaptation to habitat structure, birds sang longer songs in sites with more large trees and produced longer common song elements in sites with greater tree height. However, the acoustic adaptation hypothesis cannot explain the finding that in less‐disturbed landscapes with higher tree‐cover birds sang songs (and song elements) with higher maximum frequency and wider frequency bandwidth. We also found that birds sing longer and more variable songs of wider frequency bandwidth in less disturbed sites with a greater number of large mature trees, which may represent better feeding resources. Our study suggests that changes in song structure with habitat degradation could signal disturbed population processess, such as changes in the acoustic communication among resident birds.     

Edge effects as the principal cause of area effects on birds in fragmented secondary forest

Bird communities in tropical forests are strongly affected by both patch area and habitat edges. The fact that both effects are intrinsically confounded in space raises questions about how these two widely reported ecological patterns interact, and whether they are independent or simply different spatial manifestations of the same phenomenon. Moreover, do small patches of secondary forest, in landscapes where the most sensitive species have gone locally extinct, exhibit similar patterns to those previously observed in fragmented and continuous primary forests? We addressed these questions by testing edge‐related differences in vegetation structure and bird community composition at 31 sites in fragmented and continuous landscapes in the imperilled Atlantic forest of Brazil. Over a two‐year period, birds were captured with mist nets to a standardized effort of 680 net‐hours at each site (～22 000 net‐hours resulting in 3381 captures from 114 species). We found that the bird community in patches of secondary forest was degraded in species composition compared to primary continuous forest, but still exhibited a strong response to edge effects. In fragmented secondary forests, edge and area effects also interacted, such that the magnitude of edge to interior differences on bird community composition declined markedly with patch size. The change in bird species composition between forest interiors and edges was similar to the change in community composition between large and small patches (because species had congruent responses to edge and area), but after controlling for edge effects community composition was no longer affected by patch area. Our results show that although secondary forests hold an impoverished bird community, ecological patterns such as area and edge effects are similar to those reported for primary forests. Our data provide further evidence that edge effects are the main drivers of area effects in fragmented landscapes.     

Spatial patterns of woody plant and bird diversity: functional relationships or environmental effects?

Aim To understand cross‐taxon spatial congruence patterns of bird and woody plant species richness. In particular, to test the relative roles of functional relationships between birds and woody plants, and the direct and indirect environmental effects on broad‐scale species richness of both groups. Location Kenya. Methods Based on comprehensive range maps of all birds and woody plants (native species > 2.5 m in height) in Kenya, we mapped species richness of both groups. We distinguished species richness of four different avian frugivore guilds (obligate, partial, opportunistic and non‐frugivores) and fleshy‐fruited and non‐fleshy‐fruited woody plants. We used structural equation modelling and spatial regressions to test for effects of functional relationships (resource–consumer interactions and vegetation structural complexity) and environment (climate and habitat heterogeneity) on the richness patterns. Results Path analyses suggested that bird and woody plant species richness are linked via functional relationships, probably driven by vegetation structural complexity rather than trophic interactions. Bird species richness was determined in our models by both environmental variables and the functional relationships with woody plants. Direct environmental effects on woody plant richness differed from those on bird richness, and different avian consumer guilds showed distinct responses to climatic factors when woody plant species richness was included in path models. Main conclusions Our results imply that bird and woody plant diversity are linked at this scale via vegetation structural complexity, and that environmental factors differ in their direct effects on plants and avian trophic guilds. We conclude that climatic factors influence broad‐scale tropical bird species richness in large part indirectly, via effects on plants, rather than only directly as often assumed. This could have important implications for future predictions of animal species richness in response to climate change.     

Frugivoría por Aves en un Paisaje Fragmentado: Consecuencias en la Dispersión de Semillas1

Few data exist on seed dispersal by frugivorous birds in fragmented landscapes, originating from tropical dry forests, in contrast to more abundant data from tropical rain forests. In this study, we assessed the effect of frugivorous birds in a fragmented landscape of Veracruz, Mexico, now occupied by remnant fragments of tropical semi‐deciduous forest and dry deciduous forest, grassland, and shrubby patches on sand dunes. We determined four characteristics related to seed dispersal by birds: the interacting species of plants and birds, the characteristics of these species, spatio‐temporal variation in the dispersal system, and the outcome of the process. During one year, we recorded 54 frugivorous bird species and 33 ornithochorous plant species, which engaged in 176 different bird‐plant species interactions. Similarity (Sorensen index) of frugivorous bird communities using different vegetation types was high (>70%), suggesting that many bird species used all of the vegetation types. In contrast, the similarity of ornithochorous plant communities among vegetation types commonly was low (<37%), suggesting that most plant species were restricted to particular sites in this landscape. At the landscape level, as well as for tropical deciduous forest, we detected a significant positive relationship (Spearman's correlation of rank coefficient >0.65, P <0.05) among richness per month of frugivorous birds and plant species bearing fleshy fruits. Seeds of many plant species previously detected in studies of seed rain at the site were eaten by birds during this study. Most seeds of zoochorous species, which are deposited in the dry and decidous tropical forests patches, are produced within these vegetation types (i.e., they are autochthonous species), whereas bird‐dispersed seeds arriving in grassland and shrubby patches are produced outside (i.e., allochthonous) and are mostly woody species. Birds are important seed dispersers among vegetation types in this landscape but they have different effects in each one. The four characteristics studied, as well as the landscape approach of this research, allowed us to detect spatial and temporal patterns that otherwise would have remained undetected.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.