Avian responses to forest fragmentation during the breeding and non-breeding seasons

Forest fragmentation represents a threat to several bird species worldwide. Several factors can change across seasons (e.g. bird perception of the landscape, weather conditions, biotic interactions), which can modify the response of bird populations to forest fragmentation. However, most studies have been conducted only during the breeding season. Here we assessed the relationship between forest fragmentation (patch area and patch isolation) with population abundances of resident species during both the breeding and the non-breeding seasons. Bird population abundances (all species in the community, subsets of forest and habitat generalist species and for individual species) were estimated across a gradient of area-isolation in a semi-arid forest in Cordoba, Argentina. Population abundance of the overall avian community and of the subset of forest species declined with patch area reduction independently of the season. By contrast, the subset of habitat generalist species was not affected by patch area reduction or by the increase in patch isolation, either during the breeding or during the non-breeding season. When the analyses were carried out for individual species, we found four forest species and one habitat generalist species whose responses (the relationship between population abundance and patch area or with isolation) were different between breeding and non-breeding seasons. The negative effects of forest fragmentation were found mainly during the breeding season. Our results suggest that reduction of patch area may lead to a reduction of more than 65% of the population abundance of forest bird species, during both the breeding and the non-breeding season. Therefore, there is an urgent need to conserve large forest patches within the region as irreplaceable elements for the conservation of populations of several species.     

Breeding bird species diversity across gradients of land use from forest to agriculture in Europe

Loss, fragmentation and decreasing quality of habitats have been proposed as major threats to biodiversity world‐wide, but relatively little is known about biodiversity responses to multiple pressures, particularly at very large spatial scales. We evaluated the relative contributions of four landscape variables (habitat cover, diversity, fragmentation and productivity) in determining different components of avian diversity across Europe. We sampled breeding birds in multiple 1‐km² landscapes, from high forest cover to intensive agricultural land, in eight countries during 2001?2002. We predicted that the total diversity would peak at intermediate levels of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; forest and open‐habitat specialists would show threshold conditions along gradients of forest cover and fragmentation, and respond positively to increasing habitat diversity and productivity; resident species would be more strongly impacted by forest cover and fragmentation than migratory species; and generalists and urban species would show weak responses. Measures of total diversity did not peak at intermediate levels of forest cover or fragmentation. Rarefaction‐standardized species richness decreased marginally and linearly with increasing forest cover and increased non‐linearly with productivity, whereas all measures increased linearly with increasing fragmentation and landscape diversity. Forest and open‐habitat specialists responded approximately linearly to forest cover and also weakly to habitat diversity, fragmentation and productivity. Generalists and urban species responded weakly to the landscape variables, but some groups responded non‐linearly to productivity and marginally to habitat diversity. Resident species were not consistently more sensitive than migratory species to any of the landscape variables. These findings are relevant to landscapes with relatively long histories of human land‐use, and they highlight that habitat loss, fragmentation and habitat‐type diversity must all be considered in land‐use planning and landscape modeling of avian communities.     

SHOREBIRDS OF COASTAL BUENOS AIRES PROVINCE, ARGENTINA

The Pampas shorebird community is comprised of species from diverse taxonomic and faunal origin. Four major groups can be recognized: North American migrants residing in coastal Buenos Aires Province during their non-breeding season, Patagonian species wintering locally but breeding further south, Pampean species breeding locally, and Brazil–Paraguayan species present during their breeding season. Because of differences among these groups in the timing and migration patterns, community composition shows marked seasonal variation. Both nesting and non-breeding birds use a diversity of habitat types ranging from littoral sandy beaches to upland grasslands. Throughout the year, however, and in all habitat types, non-breeders predominate in species number and abundance. Intra- and inter-specific patterns of aggression suggest both that resources are limiting, and that non-breeding migrants may play an important role in determining breeding community composition.     

Complementary seed dispersal by three avian frugivores in a fragmented Afromontane forest

Questions: To what extent does species‐specific variation in gut passage time (GPT), habitat use and mobility of three key avian frugivores synergistically affect the distribution of Xymalos monospora seeds within and among isolated forest fragments? Location: Three fragments of a severely fragmented cloud forest, Taita Hills, southeast Kenya. Methods: We experimentally determined GPTs of X. monospora seeds and recorded movements and habitat use by Turdus helleri, Andropadus milanjensis and Tauraco hartlaubi through radiotelemetry, and combined these data to generate species‐specific seed dispersal patterns. Results: Differences in mobility and habitat use among the three frugivores caused significant complementarity in seed dispersal, despite the fact that gut transit times were highly comparable. While the most sedentary and forest‐dependent species mainly led to short‐distance dispersal away from parent trees, two more mobile species dispersed seeds further away from the source trees, both within indigenous forest patches and towards exotic plantations and isolated fruiting trees in the landscape matrix. A. milanjensis inhabiting a very small forest fragment spent significantly more time in the landscape matrix than conspecifics residing in the two larger fragments. Conclusions: By varying distances over which seeds are carried away from parent trees and the habitat types in which they are ultimately deposited, avian frugivores affect the spatial distribution of seeds and early plant recruits in a distinct and complementary manner. Because landscape properties are expected to lead to different constraints on avian mobility for habitat specialists and for generalists, ecosystem processes such as avian seed dispersal are shaped by complex interactions between disperser behaviour and the environment.     

Forest fragmentation relaxes natural nest predation in an Afromontane forest

Nest predation is widely regarded as a major driver underlying the population dynamics of small forest birds. Following forest fragmentation and the subsequent invasion by species from non-forested landscape matrices, shifts in predator communities may increase nest predation near forest edges. However, effects of human-driven habitat change on nest predation have mainly been inferred from studies with artificial nests, despite being regarded as poor surrogates for natural ones. We studied variation in predation rates, and relationships with timing of breeding and characteristics of microhabitats and fragments, on natural white-starred robin Pogonocichla stellata nests during three consecutive breeding seasons (2004–2007) in a Kenyan fragmented cloud forest. More than 70% of all initiated nests were predated during each breeding season. Predation rates nearly quadrupled between the earliest and the latest nests within a single breeding season, increased with distance to the forest edge, and decreased with the edge-to-area ratio of forest fragments. These spatial relationships oppose the traditional perception of edge and fragmentation effects on nest predation, but are in line with results from artificial nest experiments in other East African forests. In case of inverse edge and fragmentation effects on nest predation, such as shown in this study, species that tolerate edges for breeding may be affected positively, rather than negatively, by forest fragmentation, while the opposite can be expected for species restricted to the forest interior. The possibility of inverse edge effects, and its conservation implications, should therefore be taken into account when drafting habitat restoration plans.     

Effects of forest fragmentation on the oak–rodent mutualism

Land‐use changes are expected to affect plant–disperser conditional mutualisms through changes in animal behavior. We analyzed the oak–rodent conditional mutualism in Mediterranean fragmented forests at two climatically different locations. We quantified fragmentation effects on seed dispersal effectiveness and assessed if such effects were due to changes in habitat structure and intraspecific competition for acorns in fragmented areas. Fragmentation decreased cover from predators within mouse territories as well as intraspecific competition for acorns. This resulted in lower dispersal effectiveness in small forest fragments. Globally, habitat structure was the main driver in mouse foraging decisions. In small fragments, low shelter availability precluded mouse movements, leading to short mobilization distances and low caching rates. However, as the proportion of cover from predators increased, mice were able to modulate their foraging decisions depending on intraspecific competition for acorns, resulting in higher dispersal quality. In addition to fragmentation effects, delayed breeding in the southern locality caused lower number of rodents during the dispersal season, which reduced acorn mobilization rates. Our study shows that seed dispersal patterns in managed systems can be analyzed as the result of management effects on key environmental factors in dispersers’ foraging decisions.     

Experimental evidence for extreme dispersal limitation in tropical forest birds

Movements of organisms between habitat remnants can affect metapopulation structure, community assembly dynamics, gene flow and conservation strategy. In the tropical landscapes that support the majority of global biodiversity and where forest fragmentation is accelerating, there is particular urgency to understand how dispersal across habitats mediates the demography, distribution and differentiation of organisms. By employing unique dispersal challenge experiments coupled with exhaustive inventories of birds in a Panamanian lacustrine archipelago, we show that the ability to fly even short distances (< 100 m) between habitat fragments varies dramatically and consistently among species of forest birds, and that this variation correlates strongly with species' extinction histories and current distributions across the archipelago. This extreme variation in flight capability indicates that species' persistence in isolated forest remnants will be differentially mediated by their respective dispersal abilities, and that corridors connecting such fragments will be essential for the maintenance of avian diversity in fragmented tropical landscapes.     

The social organization of the Scarlet Robin Petroica multicolor and Flame Robin P. phoenicea in southeastern Australia: a comparison between sedentary and migratory flycatchers

Scarlet Robins Petroica multicolor and Flame Robins P. phoenicea occurred sympatrically during the breeding season in southeastern Australia. Scarlet Robins that occur territories in forest habitat remained there throughout the year, while those, mostly first-year birds, that occupied territories in grassland habitat were present only during the non-breeding season. Flame Robins were present for 9 months of the year and were territorial for 6 of those months. The migratory Flame Robin did not show more intense or prolonged territorial or courtship behaviour than the sedentary Scarlet Robin, despite having to obtain territories and mates when it returned each year. It is suggested that the similarities between the two species' social organizations arise partly from high adult survivorship (70–75%) and lengthy breeding seasons (4–5 months). High adult survivorship and scarcity of suitable breeding habitat also caused intense competition for territories and mates, and appears to have led to the evolution of bright, signalling plumages among female and young male Scarlet Robins. Scarcity of females and breeding habitat may have caused delayed maturation of plumage among male Flame Robins.     

Seasonal difference in the effects of fragmentation on seed dispersal by birds in Japanese temperate forests

Many studies have demonstrated that forest fragmentation reduces populations of animal species and causes local extinction, triggering many cascading effects. The effect of fragmentation on animals can be exerted through various processes, but such effects have been understudied. In this study, we posed the possibility of differences in the seasonal effects of fragmentation on frugivorous birds and their dispersal of seeds belonging to five tree species. We hypothesized that these effects may be caused by birds and their habitat selection for suitable breeding forests. We compared the abundance and species richness of frugivorous birds and the number of bird-removed fruits between a well-preserved and a fragmented temperate forest for two consecutive years. The abundance of birds was lower in the fragmented compared to the well-preserved forest during the breeding season, although no clear differences in species richness were observed. In contrast, similar decreases in bird abundance were not observed during the migratory season. After controlling for variation in crop size, the number of bird-removed fruits was lower in the fragmented forest compared to the well-preserved forest during the breeding season, but there was no such tendency during the migratory season. These results indicate that evaluations regarding the effects of fragmentation on seed dispersal that do not consider seasonal factors may lead to erroneous conclusions. This study suggests that the effects of fragmentation can be exerted though various processes, many of which remain poorly studied and warrant further examination.     

Seasonal changes in bird species diversity at the interface between forest and reed-bed

Habitat edges are landscape structures that have a major influence on animal communities. Bird communities' response to habitat edges is influenced by the season and habitat characteristics but it is still poorly understood how communities respond to structural complex (i.e. natural) edges. Inter-seasonal changes in bird species diversity were quantified at a homogeneous, sharp interface between two habitats that host distinct and well-represented bird communities: a mature broad-leaved forest and a compact reed-bed area. Resident species diversity was found higher during winter, both in terms of species richness and evenness. The presence of the reed-bed nesting migrants during the summer season did not notably modify the seasonally reversed diversity pattern; the overall evenness was higher during the winter season while the overall species richness did not differ between the two seasons. Thus, contrary to the expected regional seasonal diversity pattern, the forest – reed-bed interface is, in winter, a local bird diversity hotspot. The possible causes and implications for conservation of this phenomenon are discussed.     

Seasonal use of remnant forest fragments by understorey forest birds in the Uluguru Mountains,Tanzania: a conservation priority

Forest fragmentation can lead to extinctions of some species at local levels and is eroding bird diversity at an increasing rate. While there is information on the distribution of forest bird species in most of the Eastern Arc Mountain forests, some forests, particularly the smaller fragments, have not been adequately surveyed. Using mist netting we surveyed avifauna in some of the poorly known forests (12.5–25 ha) located 320–1 300 m above sea level in the Uluguru Mountains in order to address their conservation importance. Proportions of seasonal altitudinal migrants were significantly higher in these lower-altitude forests during the cold season than the hot season. The results suggest that these forests support bird species of conservation concern, most of which are forest dependent and some of which make seasonal movements between high-altitude montane forests and lowland/ foothill forests. These forests are important cold-season habitat of altitudinal migrants and further fragmentation should be halted as a matter of regional and global priority.     

Fine-scale genetic structure of a long-lived reptile reflects recent habitat modification

Anthropogenic habitat fragmentation — ubiquitous in modern ecosystems — has strong impacts on gene flow and genetic population structure. Reptiles may be particularly susceptible to the effects of fragmentation because of their extreme sensitivity to environmental conditions and limited dispersal. We investigate fine-scale spatial genetic structure, individual relatedness, and sex-biased dispersal in a large population of a long-lived reptile (tuatara, Sphenodon punctatus) on a recently fragmented island. We genotyped individuals from remnant forest, regenerating forest, and grassland pasture sites at seven microsatellite loci and found significant genetic structuring (R_ST = 0.012) across small distances (< 500 m). Isolation by distance was not evident, but rather, genetic distance was weakly correlated with habitat similarity. Only individuals in forest fragments were correctly assignable to their site of origin, and individual pairwise relatedness in one fragment was significantly higher than expected. We did not detect sex-biased dispersal, but natural dispersal patterns may be confounded by fragmentation. Assignment tests showed that reforestation appears to have provided refuges for tuatara from disturbed areas. Our results suggest that fine-scale genetic structuring is driven by recent habitat modification and compounded by the sedentary lifestyle of these long-lived reptiles. Extreme longevity, large population size, simple social structure and random dispersal are not strong enough to counteract the genetic structure caused by a sedentary lifestyle. We suspect that fine-scale spatial genetic structuring could occur in any sedentary species with limited dispersal, making them more susceptible to the effects of fragmentation.     

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.     

Landbird community composition varies among seasons in a heterogeneous ponderosa pine forest

ABSTRACT.   There is growing recognition of the need to conserve areas used by birds during migration, including forest and upland habitats. Because extensive thinning and burning treatments are planned for ponderosa pine ( Pinus ponderosa ) forests in the southwestern United States, information on the use of these forests by landbirds during migration is needed for conservation planning. We compared species richness among spring, breeding, and fall seasons at 69 points in a ponderosa pine forest to assess changes in landbird communities and the role of different ponderosa pine cover types in habitat selection among seasons. We detected a total of 64 bird species. Bird community similarity was lowest between the breeding and fall seasons and highest between the spring and breeding seasons. Twenty percent of the species detected were present exclusively in the fall and, of these, over half were Neotropical migrants. Only two species (3%) were detected exclusively during the spring. Although we found little difference in bird species similarity among vegetative cover types during the breeding season, forests that contained a deciduous component exhibited higher bird species similarity with each other than with habitats that did not include a deciduous component in spring and fall. In addition, foliage foragers dominated the community in spring and fall, and all Neotropical migrants detected exclusively in fall were found in ponderosa pine forests with a deciduous component. Our results indicate that ponderosa pine forests may be important to migrating or dispersing landbirds in autumn, especially if there is a deciduous component.     

Forest species in an agricultural landscape in the Netherlands: Effects of habitat fragmentation

Abstract. For 312 forest patches on sandy soils in the Netherlands, effects of fragmentation are studied of forest habitat in the past on the present occurrence of forest plant species. Using regression techniques, the numbers of forest edge, interior, zoochorous and anemochorous species, as well as occurrence of 24 individual species were related to patch area and connectivity measures. Connectivity was defined as the amount of forest habitat around patches within three zones up to 1000 m. Plant categories were distinguished by habitat type and dispersal mechanism. The results showed that number of total species and number of species of all habitat and dispersal categories increased with area. The occurrence of ten individually studied species were also positively related to area. Most of them were interior species. The number of zoochorous species increased with increasing connectivity. Also occurrence of ten individually studied species were affected by connectivity. Interior zoochorous species showed the highest percentage of affected species. The relationship of interior, animal-dispersed plants to connectivity can be explained by the limited distances covered by their dispersal agents (forest birds and ants) in a non-forest habitat. Also, some anemochorous plants appeared to be affected by connectivity, especially those with heavy seeds and potentially short distance dispersal. As not all species within a certain dispersal or habitat category react similar to area or isolation, it is suggested that differences in underlying processes of fragmentation such as local extinction and colonization need more focus.     

Winter bird communities in the northern Negev: species dispersal patterns, habitat use and implications for habitat conservation

Bird habitat conservation may require different management strategies for different seasonal bird assemblages. We studied habitat use by winter birds in forest and scrubland habitat patches in the northern Negev, Israel. Our goal was to assess whether differences in responses to landscape and habitat structure between breeding and non-breeding seasons require changes in future conservation plans that have been suggested for the Negev breeding bird community. We evaluated habitat and area effects on bird abundance and distribution and tested whether species habitat use during winter involves niche shifts. Compared with breeding birds, a larger proportion of winter bird species occupied both scrubland and forest. As in summer, forest bird species responded to habitat structure, whereas scrubland species were associated with both habitat structure and area. Resident birds disperse into habitats in which they were not present during summer. Consequently, for several species, the correlation between bird densities and environmental factors showed a better fit at the landscape rather than at the habitat scale. In addition, rather than niche shift, birds actually extended their niche breadth. Nest site selection may constrain bird distribution into a realized niche, smaller than their fundamental niche. Despite the scale differences in habitat use, the similar species diversity patterns between seasons suggest that both winter and summer birds would benefit from conservation of scrub patches larger than 50 ha, and enrichment of foliage layers within the planted forests.     

Habitat fragmentation shapes natal dispersal and sociality in an Afrotropical cooperative breeder

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.     

Do forest plants conform to the theory of island biogeography: the case study of bog islands

Biodiversity conservation is confronted with increasing risk of extinction in isolated small-area remnants and the limitation of species to colonize recently formed habitats. We hypothesized that the equilibrium pattern of forest herb layer in long-term fragmented landscape should comply with the theory of island biogeography. Forests on mineral soil islands located in large mires of western Estonia were considered as dispersal target habitats, and forests on mainland and peninsulas in mires as sources. Species richness was the lowest in mainland forests and the effect was confounded by habitat structure, suggesting a negative effect of silvicultural management in easily accessible forests. We observed the ‘small island effect’, i.e. greater overall species richness in small-area habitats, which was determined by the habitat preference of shade tolerant generalists. The average species richness of common mainland forest specialists varied little, but capitalizing on the traditional approach and analyzing only island data, weak effects of distance and habitat quality were detected. At single species level, unexpectedly, many habitat specialists were observed to have successfully dispersed to islands, indicating insufficient knowledge of the long-distance dispersal mechanisms of forest-dwelling plants. In fragmented forest landscapes the theory of island biogeography can be applied to habitat specialist plant species, but only regarding the effect of isolation and in conditions of persistent forest structural quality. In the light of global changes, optimized conservation planning should primarily target on (i) the conservation of ancient habitat fragments independent of their current area, and (ii) the promotion of diversity of potential dispersal vectors in the landscape.     

The roles of ecology,behaviour and effective population size in the evolution of a community

Organismal traits such as ecological specialization and migratory behaviour may affect colonization potential, population persistence and degree of isolation, factors that determine the composition and genetic structure of communities. However, studies focusing on community assembly rarely consider these factors jointly. We sequenced 16 nuclear genes and one mitochondrial gene from Caucasian and European populations of 30 forest‐dwelling avian species that represent diverse ecological (specialist–generalist) and behavioural (migratory‐resident) backgrounds. We tested the effects of organismal traits on population divergence and community assembly in the Caucasus forest, a continental mountain island setting. We found that (i) there is no concordance in divergence times between the Caucasus forest bird populations and their European counterparts, (ii) habitat specialists tend to be more divergent than generalists and (iii) residents tend to be more divergent than migrants. Thus, specialists and residents contribute to the high level of endemism of Caucasus forest avifauna more than do generalists and migrants. Patterns of genetic differentiation are better explained by differences in effective population sizes, an often overlooked factor in comparative studies of phylogeography and speciation, than by divergence times or levels of gene flow. Our results suggest that the Caucasus forest avifauna was assembled through time via dispersal and/or multiple vicariant events, rather than originating simultaneously via a single isolation event. Our study is one of the first multilocus, multispecies analyses revealing how ecological and migratory traits impact the evolutionary history of community formation on a continental island.     

Population structure and mitochondrial DNA variation in sedentary Neotropical birds isolated by forest fragmentation

The current worldwide concern about tropical deforestation raises questions about the sustainability of avian populations in isolated forest fragments. One of the most important issues concerns the sizes of forest fragments necessary to maintain populations and the genetic variation within them. We address this by: (1) using mtDNA sequence variation to infer aspects of the population structure of four species of understory birds from four sites in southern Costa Rican rainforest; and (2) determining whether forest fragmentation that has occurred in the last 50 years has had an effect on the amount of within-population variation for the species in question. High levels of between-population differentiation (D _xy ) were found over a relatively small geographic scale (<130 km) for white-breasted wood-wren (Henicorhina leucosticta), bicolored antbird (Gymnopithys leucaspis), and gray-headed tanager (Eucometis penicillata), suggesting that these species are highly sedentary and exhibit strong female philopatry. No mtDNA variation was found in Plain Antvireo (Dysithamnus mentalis). In all three of the polymorphic species there was a significant decrease in mtDNA nucleotide diversity in populations isolated by forest fragmentation as compared to populations in contiguous primary forest. Even in relatively large (250–1000 ha) forest reserves, sedentary avian species have lost roughly half (range 43–85) of the nucleotide diversity in mtDNA over a relatively short period of time. Our results indicate that sedentary avian species in forest fragments isolated by clearing have undergone severe reductions in effective population size due to population bottlenecks perpetuated by prolonged isolation and potential edge effects.