Changes in patch features may exacerbate or compensate for the effect of habitat loss on forest bird populations

One and a half centuries after Darwin visited Chiloe Island, what he described as "…an island covered by one great forest…" has lost two-thirds of its forested areas. At this biodiversity hotspot, forest surface is becoming increasingly fragmented due to unregulated logging, clearing for pastures and replacement by exotic tree plantations. Decrease in patch size, increased isolation and "edge effects" can influence the persistence of forest species in remnant fragments. We assessed how these variables affect local density for six forest birds, chosen to include the most important seed dispersers (four species) and bird pollinators (two species, one of which acts also as seed disperser), plus the most common insectivore (Aphrastura spinicauda). Based on cue-count point surveys (8 points per fragment), we estimated bird densities for each species in 22 forest fragments of varying size, shape, isolation and internal-habitat structure (e.g. tree size and epiphyte cover). Bird densities varied with fragment connectivity (three species) and shape (three species), but none of the species was significantly affected by patch size. Satellite image analyses revealed that, from 1985 to 2008, forested area decreased by 8.8% and the remaining forest fragments became 16% smaller, 58-73% more isolated and 11-50% more regular. During that period, bird density estimates for the northern part of Chiloé (covering an area of 1214.75 km(2)) decreased for one species (elaenia), increased for another two (chucao and hummingbird) and did not vary for three (rayadito, thrust and blackbird). For the first three species, changes in patch features respectively exacerbated, balanced and overcame the effects of forest loss on bird population size (landscape-level abundance). Hence, changes in patch features can modulate the effect of habitat fragmentation on forest birds, suggesting that spatial planning (guided by spatially-explicit models) can be an effective tool to facilitate their conservation.     

Developmental instability and inbreeding in natural bird populations exposed to different levels of habitat disturbance

As habitat disturbance and inbreeding increasingly stress natural populations, ecologists are in urgent need of simple estimators to measure their impact. It has been argued that developmental instability (DI) could be such a measure. Observed associations between DI and environmental or genetic stress, however, are largely inconsistent. We here test whether an interaction between habitat disturbance and inbreeding could, at least partly, explain these discordant patterns. We therefore studied individual estimates of fluctuating asymmetry (FA) and of inbreeding in three populations of the critically endangered Taita thrush that are differentially exposed to habitat disturbance following severe forest fragmentation. As predicted, the relationship between DI and inbreeding was pronounced under high levels of disturbance, but weak or nonexistent under less disturbed conditions. Examining this relationship with mean d², an allelic distance estimator assumed to reflect ancestral inbreeding, did not reveal any significant trend, hence suggesting that inbreeding effects in the Taita thrush are fairly recent.     

Slow response of plant species richness to habitat loss and fragmentation

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago ( R ² = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km² in the 1930s to 0.21 km² at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.     

Decline of some west-central Florida anuran populations in response to habitat degradation

Recent reports have suggested that a global decline in amphibian populations has taken place during the past few decades. Urban development is thought to affect the richness and abundances of species and, therefore, could be an important cause of decline. We estimated the richness and abundances of anurans in wetlands at a residential development and in similar wetlands at a nearby undeveloped park. The residential development originally was pine flatwoods habitat, as is the undeveloped park curiently. We also compared the anuran species' composition of the park in 1992 with the composition in 1974.Both richness and abundances of anurans in the residential development were different than those in the undeveloped park. Employing the same amount of sampling effort at both sites, we trapped or observed 11 species at the developement and 15 species at the park, and trapped 569 individuals at the development and 1224 individuals at the park. The anuran species richness at the undeveloped park in 1992 was nearly the same as in 1974; a single rare species apparently was not present in 1992. Of the 15 species present in both surveys, 14 showed higher abundances in 1992 than in 1974.We suggest that the current differences between the residential development and the park have resulted from degradation of both the uplands used by many species during the dry season and the temporary wetlands used by many species for reproduction. Four species especially sensitive to such degradation, Bufo quercicus, Scaphiopus h. holbrookii, Hyla femoralis, and H. gratiosa, were the species missing from the residential development.Not all species of anurans typical of pine flatwoods appeared to be affected adversely by development. Three species of ranids, Rana utricularia, R. grylio, and R. catesbeiana, were found in higher abundances at the residential development than at the park. These ranid species breed in a wide variety of aquatic systems, including the permanent bodies of water that are now abundant in the development, and probably use the uplands less than other anurans.If amphibian decline is international in scope, then the decline could be attributable either to global changes caused by humans, or to local, but widespread, environmental degradation, or to a combination of factors. While much recent popular focus has been on potential global causes of decline, we believe that this emphasis may have caused attention to be taken away from local causes that, as our study demonstrated, may be at least as important. We suggest that in many places, local environmental degradation is insidiously chipping away at amphibian diversity, and that more emphasis should be placed on these local causes than is now the case.     

Demographic response of plant populations to habitat fragmentation and temporal environmental variability

Many plant species currently exist in fragmented populations of different sizes, while they also experience unpredictable climatic fluctuation over time. However, we still understand little about how plant demography responds to such spatial and temporal environmental variability. We studied population dynamics of an understory perennial herb Trillium camschatcense in the Tokachi plain of Hokkaido, Japan, where a significant effect of forest fragmentation on seedling recruitment was previously reported. Four populations across a range of fragment sizes were studied for 6 years, and the data were analyzed using matrix population models. Per capita fecundity (the number of recruits per plant) varied greatly among populations, but the variation in population growth rates (λ) was mainly driven by the variation in stasis and growth rates, suggesting that the general trend of reduced fecundity in fragmented populations may not be readily translated into subsequent dynamics. Temporal variation in λ among years was more than 2 times larger than spatial variation among populations, and this result was likely attributable to the contrasting response of correlation structures among demographic rates. The among-population variation in λ was dampened by negative covariation between matrix elements possibly due to density-dependent regulation as well as an inherent constraint that some elements are not independent, whereas positive covariation between matrix elements resulted in large temporal variation in λ. Our results show that population dynamics responded differently to habitat fragmentation and temporal variability of the environment, emphasizing the need to discriminate these spatial and temporal variations in demographic models. Although no populations were projected to be declining in stochastic simulations, correlation between current habitat size and plant density implies historical λ is positively related to habitat size.     

Atlantic forest bird communities provide different but not fewer functions after habitat loss

Habitat loss often reduces the number of species as well as functional diversity. Dramatic effects to species composition have also been shown, but changes to functional composition have so far been poorly documented, partly owing to a lack of appropriate indices. We here develop three new community indices (i.e. functional integrity, community integrity of ecological groups and community specialization) to investigate how habitat loss affects the diversity and composition of functional traits and species. We used data from more than 5000 individuals of 137 bird species captured in 57 sites in the Brazilian Atlantic Forest, a highly endangered biodiversity hotspot. Results indicate that habitat loss leads to a decrease in functional integrity while measures of functional diversity remain unchanged or are even positively affected. Changes to functional integrity were caused by (i) a decrease in the provisioning of some functions, and an increase in others; (ii) strong within-guild species turnover; and (iii) a replacement of specialists by generalists. Hence, communities from more deforested sites seem to provide different but not fewer functions. We show the importance of investigating changes to both diversity and composition of functional traits and species, as the effects of habitat loss on ecosystem functioning may be more complex than previously thought. Crucially, when only functional diversity is assessed, important changes to ecological functions may remain undetected and negative effects of habitat loss underestimated, thereby imperiling the application of effective conservation actions.     

Home range size of willow tits: a response to winter habitat loss

We examined the behavioral response to habitat loss and fragmentation of willow tits (Parus montanus) in winter in a mosaic forest landscape in northern Finland. We studied habitat preference, flock size and home range size of 16 flocks, half of which had their territories in forests fragmented by forestry and half in continuous forest. We predicted that birds would respond to habitat loss by enlarging their home range and/or diminishing group size. In addition, to compensate for fragmentation effects, willow tits might be expected to include more optimal habitat into their territories. Flocks included on average 3.9 birds and occupied territories of 12.6 ha. Willow tits avoided open areas (clear cuts and young sapling stands) and preferred mature forests and older sapling stands or pine bogs equally. Birds responded to habitat loss by enlarging their home ranges but not by reducing the group size. Large territories included a smaller proportion of mature forests, but the proportion of sapling and pine bog habitat did not change. Birds on territories that included a large proportion of open habitat localized their activity on several distinct habitat patches that were distributed over a wide area. We conclude that willow tits adjust territory use to compensate for the inclusion of unsuitable habitat within home ranges, and older sapling areas and pine bogs serve as surrogates for mature forests. However, birds did not enlarge the proportion of forest habitat in their territories with increasing habitat loss. Thus, our data do not suggest a strong effect of fragmentation, but imply that forestry practices reduce suitable wintering habitat and carrying capacity in the area. Thus winter habitat loss may explain the observed population decline of willow tits in Finland during recent decades.     

The utility of contemporary and historical estimates of dispersal in determining response to habitat fragmentation in a tropical forest-dependent bird community

It is often assumed that species which exhibit a greater propensity for dispersal are less susceptible to the impacts of habitat fragmentation; however, a growing body of literature suggests that such generalizations should be carefully evaluated as not all species appear to be equally sensitive to fragmentation. In this issue of Molecular Ecology, Callens et al. (2011) take an innovative approach to compare contemporary estimates of dispersal from an extensive mark-recapture and patch occupancy data set with historical estimates derived from multilocus population genetic models for seven sympatric forest-dependent species in the Taita Hills, Africa. As has been observed for forest-dependent species from the Amazon, populations of sedentary species were more strongly differentiated and clustered when compared to those of more dispersive taxa. The most intriguing result recovered though, was that the five species with similar historical estimates of gene flow (dispersal) differed substantially in their contemporary dispersal rates, suggesting that for some species the propensity for dispersal has decreased over time. As a consequence, the authors suggest that post-fragmentation estimates of dispersal on their own may not be the best predictors of how habitat fragmentation could affect forest-dependent animal communities.This work significantly advances our understanding of the dynamics of habitat fragmentation and makes a strong case for the need to integrate data on historical processes with contemporary data.     

Dispersal capacity and diet breadth modify the response of wild bees to habitat loss

Habitat loss poses a major threat to biodiversity, and species-specific extinction risks are inextricably linked to life-history characteristics. This relationship is still poorly documented for many functionally important taxa, and at larger continental scales. With data from five replicated field studies from three countries, we examined how species richness of wild bees varies with habitat patch size. We hypothesized that the form of this relationship is affected by body size, degree of host plant specialization and sociality. Across all species, we found a positive species–area slope (z = 0.19), and species traits modified this relationship. Large-bodied generalists had a lower z value than small generalists. Contrary to predictions, small specialists had similar or slightly lower z value compared with large specialists, and small generalists also tended to be more strongly affected by habitat loss as compared with small specialists. Social bees were negatively affected by habitat loss (z = 0.11) irrespective of body size. We conclude that habitat loss leads to clear shifts in the species composition of wild bee communities.     

Latency in response of barn swallow Hirundo rustica populations to changes in breeding habitat conditions

Inclusion of past, in addition to current ecological conditions can improve the realism of animal‐habitat models, because populations may show delayed responses to changing environments. We studied the relationships between the distribution of barn swallows (Hirundo rustica L.) in 2001 and ecological factors between 1986 and 2001. Livestock farming was the best predictor of swallow distribution. Maximum predictive power of swallow abundance was achieved by livestock farming data 7 years before the census and significantly earlier than 2001, as shown by a bootstrap analysis. Breeding philopatry, social facilitation of breeding habitat choice may delay response of barn swallow populations to rapidly changing ecological conditions in anthropogenic habitats. By bootstrapping the set of farms and applying our predictive distribution models to livestock farming data in the past and those predicted for the near future we forecast no significant change in swallow populations until 2015. This study suggests that population studies should incorporate historical ecological data.     

The robustness of a network of ecological networks to habitat loss

There have been considerable advances in our understanding of the tolerance of species interaction networks to sequential extinctions of plants and animals. However, communities of species exist in a mosaic of habitats, and the vulnerability of habitats to anthropogenic change varies. Here, we model the cascading effects of habitat loss, driven by plant extinctions, on the robustness of multiple animal groups. Our network is constructed from empirical observations of 11 animal groups in 12 habitats on farmland. We simulated sequential habitat removal scenarios: randomly; according to prior information; and with a genetic algorithm to identify best‐ and worst‐case permutations of habitat loss. We identified two semi‐natural habitats (waste ground and hedgerows together comprising < 5% of the total area of the farm) as disproportionately important to the integrity of the overall network. Our approach provides a new tool for network ecologists and for directing the management and restoration of multiple‐habitat sites.     

Fluctuating and directional asymmetry in natural bird populations exposed to different levels of habitat disturbance, as revealed by mixture analysis

While bilateral trait asymmetry is widely recognized to estimate developmental instability, much controversy exists over which types of asymmetry (fluctuating, directional, and/or antisymmetry) to use. Recently it has been hypothesized that the three types are strongly interrelated, and that increased developmental instability may be reflected in a transition from fluctuating to directional asymmetry and/or antisymmetry. Alternatively, habitat disturbance might change the genetic expression of directional asymmetry. We present herein the first empirical evidence for stress-mediated shifts in types of asymmetry in natural populations, by using mixture analysis to model tarsus asymmetry in bird populations exposed to different levels of habitat disturbance. Observed asymmetry patterns almost exclusively consisted of true fluctuating asymmetry in the least disturbed populations, but became progressively mixed with directional asymmetry under increasing disturbance. Failing to unravel these mixtures of different forms of asymmetry may have critical implications for the analysis and interpretation of asymmetry data.     

Genetic and phenotypic differences between thistle populations in response to habitat and weed management practices

Rapid evolutionary change is increasingly being recognized as commonplace, but the evolutionary consequences for species and ecosystems under human‐induced selection regimes have not been explored in detail, although many species occur in such environments. In a common garden experiment and with amplified fragment length polymorphism markers, we examined whether genetic differentiation has taken place between spatially intermixed populations of creeping thistles Cirsium arvense (Asteraceae) collected from a natural habitat (maritime shores), a semi‐natural habitat (road verges) and arable fields under two management regimes: conventional and organic farming. Populations of C. arvense have altered genetically and locally adapted their growth patterns with changed land use. Although plants from different habitats showed similar total biomass production, shoot and root production was higher for maritime populations, suggesting selection for increased competitive ability. Competitive ability then declined in the order semi‐natural, conventional farms and organic farms. Thistles in arable fields may be more selected for tolerance against disturbances from herbicides and mechanical weed control. In addition, early shoot sprouting and genetic analysis showed differentiation between plants originating from conventional farms and farms that were converted to organic 9–30 years ago, suggesting some adaptation to altered crop cultivation practices. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 797–807.     

城市公园生境类型对鸟类群落的影响

2011年12月—2012年11月,在上海世纪公园和滨江森林公园对鸟类群落和植物群落进行调查,通过对12个植被变量进行主成分分析,将两个公园分为8种不同的生境类型。结果表明:2个公园生境构成存在显著差异,滨江森林公园灌木层植物发达的生境(Habitat with developed shrub layer,S型)以及灌木层和地被层植物都发达的生境(Habitat with developed tree layer and shrub layer,T+S型)数量显著多于世纪公园,世纪公园地被层发达的生境(Habitat with developed ground cover layer,G型)以及乔木层和地被层植物都发达的生境(Habitat with developed tree layer and ground cover layer,T+G型)数量显著多于滨江森林公园。世纪公园不同生境中鸟种数差异显著,而滨江森林公园中差异不显著。2个公园有24种共有鸟种,对共有鸟种生境利用率的配对t检验结果表明,滨江森林公园鸟类生境利用率显著高于世纪公园。对2个公园共有生境类型中鸟种数进行分析,发现滨江森林公园鹟科(Muscicapidae)鸟类种数显著大于世纪公园。根据以上结果,上海城市公园不同生境类型对鸟类群落结构存在显著影响。因此,建议在规划和建设大型城市公园时,应构建植被分层结构复杂的生境,多样化种植各类乔木,林下多样化搭配灌木。在保留供游客休憩草坪区域的同时种植各类草本植物,以此提高鸟类生境利用率,增加城市公园的鸟类多样性。     

Variation in the response of an Arctic top predator experiencing habitat loss: feeding and reproductive ecology of two polar bear populations

Polar bears (Ursus maritimus) have experienced substantial changes in the seasonal availability of sea ice habitat in parts of their range, including the Beaufort, Chukchi, and Bering Seas. In this study, we compared the body size, condition, and recruitment of polar bears captured in the Chukchi and Bering Seas (CS) between two periods (1986–1994 and 2008–2011) when declines in sea ice habitat occurred. In addition, we compared metrics for the CS population 2008–2011 with those of the adjacent southern Beaufort Sea (SB) population where loss in sea ice habitat has been associated with declines in body condition, size, recruitment, and survival. We evaluated how variation in body condition and recruitment were related to feeding ecology. Comparing habitat conditions between populations, there were twice as many reduced ice days over continental shelf waters per year during 2008–2011 in the SB than in the CS. CS polar bears were larger and in better condition, and appeared to have higher reproduction than SB bears. Although SB and CS bears had similar diets, twice as many bears were fasting in spring in the SB than in the CS. Between 1986–1994 and 2008–2011, body size, condition, and recruitment indices in the CS were not reduced despite a 44‐day increase in the number of reduced ice days. Bears in the CS exhibited large body size, good body condition, and high indices of recruitment compared to most other populations measured to date. Higher biological productivity and prey availability in the CS relative to the SB, and a shorter recent history of reduced sea ice habitat, may explain the maintenance of condition and recruitment of CS bears. Geographic differences in the response of polar bears to climate change are relevant to range‐wide forecasts for this and other ice‐dependent species.     

Changes in bird populations on sample lowland English farms in relation to loss of hedgerows and other non-crop habitats

Farmland bird population trends were examined on a sample of lowland English farms to assess the relative importance of habitat loss and habitat degradation. Data were extracted from 11 farms surveyed by territory mapping between 1966 and 1986 as part of the British Trust for Ornithology's Common Birds Census. The population size of 38 bird species was quantified for each farm in each year. The extents of five non-crop habitats were measured at 4-yearly intervals on each farm. The farms were selected because some had undergone extensive removal of non-crop habitats while others had undergone little or none. Although declines were commonest on farms where the severest habitat loss had taken place, we found no evidence that habitat loss was the main factor causing population declines: all 11 farms had significant numbers of declining species, even where habitat loss was minimal. Furthermore, general linear modelling found no significant effects of habitat loss on population trends and principal-components analysis found limited effects of habitat extent on community composition. These results suggest that habitat loss is of secondary importance in causing farmland bird population declines. We suggest that other processes, such as habitat degradation, may have caused a baseline population decline in at least 10 farmland bird species and that declines may have been exacerbated by localised habitat loss. Received: 4 February 1998 / Accepted: 1 April 1998     

Multi-scale habitat models for reintroduced bird populations: a case study of South Island saddlebacks on Motuara Island

Understanding resource selection by animals is important when considering habitat suitability at proposed release sites within threatened species recovery programmes. Multi-scale investigatory approaches are increasingly encouraged, as the patchy distribution of suitable habitats in fragmented landscapes often determines species presence and survival. Habitat models applied to a threatened New Zealand forest passerine, the South Island saddleback (Philesturnus carunculatus carunculatus), reintroduced to Ulva Island (Stewart Island) found that at landscape scale breeding pairs? preferences for sites near the coast were driven by micro-scale vegetation structure. We tested these results by examining models of breeding site selection by a reintroduced saddleback population on Motuara Island (Marlborough Sounds) at two scales: (1) micro-scale, for habitat characteristics that may drive breeding site selection, and (2) landscape scale, for variations in micro-scale habitat characteristics that may influence site colonisation in breeding pairs. Results indicated that birds on Motuara Island responded similarly to those on Ulva Island, i.e. birds primarily settled at the margins of coastal scrub and forest and later cohorts moved into larger stands of coastal forest where they established breeding territories. Plant species composition was also important in providing breeding saddleback pairs with adequate food supply and nesting support. However, Motuara Island birds differed in their partitioning of habitat use: preferred habitats were used for nesting while birds were foraging outside territorial boundaries or in shared sites. These differences may be explained because Motuara has a more homogeneous distribution of microscale habitats throughout the landscape and a highly bird-populated environment. These results show that resource distribution and abundance across the landscape needs to be accounted for in the modelling of density?bird?habitat relationships. In the search for future release sites, food (invertebrates and fruiting tree species) should be abundant close to available nesting sites, or evenly spread and available throughout the landscape.