Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

Fine-scale structure and cross-taxon congruence of bird and beetle assemblages in an old-growth boreal forest mosaic

Aim Nestedness occurs when species present in depauperate sites are subsets of those found in species‐rich sites. The degree of congruence of site nestedness among different assemblages can inform commonalities of mechanisms structuring the assemblages. Well‐nested assemblages may still contain idiosyncratic species and sites that notably depart from the typical assemblage pattern. Idiosyncrasy can arise from multiple processes, including interspecific interactions and habitat preferences, which entail different consequences for species co‐occurrences. We investigate the influence of fine‐scale habitat variation on nestedness and idiosyncrasy patterns of beetle and bird assemblages. We examine community‐level and pairwise species co‐occurrence patterns, and highlight the potential influence of interspecific interactions for assemblage structure. Location Côte‐Nord region of Québec, Canada. Methods We sampled occurrences of ground‐dwelling beetles, flying beetles and birds at sites within old‐growth boreal forest. We examined the nestedness and idiosyncrasy of sites and sought relationships to habitat attributes. We analysed non‐random species co‐occurrence patterns at pairwise and community levels, using null model analysis and five ‘association’ indices. Results All three assemblages were significantly nested. There was limited congruence only between birds and flying beetles whose nestedness was related to canopy openness. For ground‐dwelling beetles, nestedness was related to high stand heterogeneity and sapling density, whereas site idiosyncrasy was inversely related to structural heterogeneity. For birds, site idiosyncrasy increased with canopy cover, and most idiosyncratic species were closed‐canopy specialists. In all assemblages, species idiosyncrasy was positively correlated with the frequency of negative pairwise associations. Species co‐occurrence patterns were non‐random, and for flying beetles and birds positive species pairwise associations dominated. Community‐level co‐occurrence summaries may not, however, always reflect these patterns. Main conclusions Nestedness patterns of different assemblages may not correlate, even when sampled at common locations, because of different responses to local habitat attributes. We found idiosyncrasy patterns indicating opposing habitat preferences, consistent with antagonistic interactions among species within assemblages. Analysis of such patterns can thus suggest the mechanisms generating assemblage structures, with implications for biodiversity conservation.     

Fat reserves and perceived predation risk in the great tit, Parus major

The fat reserves of small birds are built up daily as insurance against starvation. They are believed to reflect a trade-off between the risks of starvation and predation such that in situations of high predation risk birds are expected either to reduce their fat reserves in response to mass-dependent predation risk or to increase them in response to foraging interruptions. We assessed the effect on fat reserves of experimentally altering the perceived (but not the actual) risk of predation of wild great tits at a winter feeding site. The perceived predation risk was alternated between 'safe' and 'risky'. Increasing the perceived risk of predation involved 'swooping' a model sparrowhawk over the feeder at four unpredictable times each day using a remote mechanism We produce evidence that the experiment was suceessfull in altering the perceived risk of predation. As predicted from the hypothesis of mass-dependent predation risk, great tits (Parus major) carried significantly reduced fat reserves during the 'risky' treatment. Furthermore, dominant individuals were able to reduce their reserves more than subordinates. As birds returned to feeders within seconds after a predator 'attack', the reduction in fat reserves cannot be attributed to an interruption in feeding.     

Patch Size and Tree Species Influence the Number and Duration of Bird Visits in Forest Restoration Plots in Southern Costa Rica

Efforts to restore tropical forest in abandoned pasture are likely to be more successful when bird visitation is promoted because birds disperse seeds and eat herbivorous arthropods that damage leaves. Thus, it is critical to understand bird behavior in relation to different restoration strategies. We measured the likelihood of visitation, number of visits, and duration of visits for all birds and for Cherrie's Tanager ( Ramphocelus costaricensis ), a common seed disperser, in five sizes of forest restoration patches planted with four tree species in southern Costa Rica. We predicted that the largest patches, and the tree species with the greatest canopy cover, would be visited most frequently and have the longest visits because we assumed that these patch types had the greatest food resources and the lowest predation risk. We found that birds were more likely to visit large patches and the tree species with the highest canopy cover ( Inga edulis ). Birds visited Inga trees more often and stayed in Inga and Erythrina poeppigiana trees for longer periods of time than in other tree species. We found similar results for Cherrie's Tanagers. Thus, we identified two factors, tree species and patch size, which may be manipulated in restoration projects to increase bird visitation.     

Analysis of factors affecting bird predation on gypsy moth egg masses by using Holling’s disc-equation

1. Birds’ predation on gypsy moth egg masses deposited on tree trunks was quantitatively evaluated in a birch forest at Bibai, Hokkaido during the four winter seasons from 1974 to 1978.
2. Holling's disc-equation could be successfully applied to describe the relationship between the total egg consumption by birds and the density of egg masses. A rapid method for fitting the disc-equation to observed data was proposed based on the study byShinozaki andKira (1961).
3. Appreciable amounts of predation were observed only during the period when the depth of snow cover on the ground exceeded 1 m, while the egg masses deposited on lower parts of tree trunk below 1 m level suffered from only slight predation. It was therefore presumed that deep snow cover made the predator birds to change their feeding place from ground vegetation to tree trunks.
4. The duration of the period with snow accumulation deeper than 1 m was introduced into the disc-equation as an additional parameter. The values calculated by the resultant formulation agreed with the observed values in respective years satisfactorily.
5. There were circumstantial evidences that the great tit, Parus major, was one of the important predators concerned.

     

Influence of within‐patch habitat quality on high‐Andean Polylepis bird abundance

Habitat restoration strategies for fragmented high Andean forest landscapes must consider the influence of within‐patch habitat quality on bird abundance. I examined vegetation and bird abundance at three locations within a highly fragmented Polylepis forest landscape in the Cordillera Vilcanota, southern Peru. Across the landscape, there was significant variation in the vegetation structure of Polylepis forest patches of different size categories, especially in terms of tree girth, tree height, tree density, and canopy vegetation structure. Principal Component Analysis extracted five factors of habitat quality, which together accounted for 74.2% of the variability within 15 habitat variables. Polylepis bird species differed in their responses to habitat quality but, overall, variation in Polylepis bird abundance was not fully captured by the range of habitat quality variables. Tall, dense vegetation cover was clearly important for 11 conservation‐important species, a high density of large trees was important for 10 species and primary forest ground cover was important for eight species. Habitat quality exhibited no significant influence on the abundance of only one species –Asthenes urubambensis. The abundance of seven species was associated with lower elevation forest, but only one species was associated with higher elevation forest. Management of habitat quality in large and medium remnant forest patches throughout the Cordillera Vilcanota, particularly in the 3800–4200 m elevation range, will be a cornerstone in ensuring the persistence of the majority of conservation‐important bird species populations.     

Compounding effects of habitat fragmentation and predation on bird nests

Habitat fragmentation and invasive species are two of the greatest threats to species diversity worldwide. This is particularly relevant for oceanic islands with vulnerable endemics. Here, we examine how habitat fragmentation influences nest predation by Rattus spp. on cup‐nesting birds in Samoan forests. We determined models for predicting predation rates by Rattus on artificial nests at two scales: (i) the position of the bird's nest within the landscape (e.g. proximity to mixed crop plantations, distance to forest edge); and (ii) the microhabitat in the immediate vicinity of the nest (e.g. nest height, ground cover, slope). Nest cameras showed only one mammal predator, the black rat (Rattus rattus), predating artificial nests. The optimal model predicting nest predation rates by black rats included a landscape variable, proximity to plantations and a local nest site variable, the percentage of low (<15 cm) ground cover surrounding the nest tree. Predation rates were 22 ± 13% higher for nests in forest edges near mixed crop plantations than in edges without plantations. In contrast, predation rates did not vary significantly between edge habitat where the matrix did not contain plantations, and interior forest sites (>1 km from the edge). As ground cover reduced, nest predation rates increased. Waxtags containing either coconut or peanut butter were used as a second method for assessing nest predation. The rates at which these were chewed followed patterns similar to the predation of the artificial nests. Rural development in Samoa will increase the proportion of forest edge near plantations. Our results suggest that this will increase the proportion of forest birds that experience nest predation from black rats. Further research is required to determine if rat control is needed to maintain even interior forest sites populations of predator‐sensitive bird species on South Pacific islands.     

Near‐ground solar radiation along the grassland–forest continuum: Tall‐tree canopy architecture imposes only muted trends and heterogeneity

Solar radiation directly and indirectly drives a variety of ecosystem processes. Our aim was to evaluate how tree canopy architecture affects near‐ground, incoming solar radiation along gradients of increasing tree cover, referred to as the grassland–forest continuum. We evaluated a common type of canopy architecture: tall trees that generally have their lowest level of foliage high above, rather than close to the ground as is often the case for shorter trees. We used hemispherical photographs to estimate near‐ground solar radiation using the metric of Direct Site Factor (DSF) on four sites in north Queensland, Australia that formed a grassland–forest continuum with tree canopy cover ranging from 0% to 71%. Three of the four sites had tall Eucalyptus trees with foliage several metres above the ground. We found that: (i) mean DSF exceeded >70% of the potential maximum for all sites, including the site with highest canopy cover; (ii) DSF variance was not highly sensitive to canopy coverage; and (iii) mean DSF for canopy locations beneath trees was not significantly lower than for adjacent intercanopy locations. Simulations that hypothetically placed Australian sites with tall tree canopies at other latitude–longitude locations demonstrated that differences in DSF were mostly due to canopy architecture, not specific site location effects. Our findings suggest that tall trees that have their lowest foliage many metres above the ground and have lower foliar density only weakly affect patterns of near‐ground solar radiation along the grassland–forest continuum. This markedly contrasts with the strong effect that shorter trees with foliage near the ground have on near‐ground solar radiation patterns along the continuum. This consequence of differential tree canopy architecture will fundamentally affect other ecosystem properties and may explain differential emphases that have been placed on canopy–intercanopy heterogeneity in diverse global ecosystem types that lie within the grassland–forest continuum.     

Prevalence of nest predators in a sub-Arctic ecosystem

High nest loss is an important driver of gallinaceous bird population dynamics. Identifying factors determining the spatial distribution of potential nest predators and thereby indirectly risk of nest losses is therefore essential. The aim of this 1-year study was to estimate relative predation rates on artificial ground nests in willow ptarmigan (Lagopus lagopus) habitats, along replicate altitudinal gradients (transects, n?=?60) spanning from sub-Arctic birch forest to the low-alpine tundra in three locations in northern Norway. In each transect, one artificial nest (track board) was placed in three different habitats: (1) birch forest, (2) edge between birch forest and low-alpine tundra and (3) low-alpine tundra. Total predation rates over all habitats within locations ranged from 47.4% to 77.5% and did not vary systematically in space and time. The average predation rate by avian predators was consistently high (58%), and mammalian predation rate was consistently low (5.6%). The consistently high level of predation inflicted by birds was mainly due to omnipresent corvids, especially the hooded crow (Corvus cornix). Analysis of species-specific predation rates showed that habitat and location effects were insignificant for all species, except for raven (Corvus corax) that showed clearly higher predation in one of the locations. The results indicate that from the perspective of the spatial distribution of potential nest predators in sub-Arctic birch forest, ground nesting birds like willow ptarmigan should not be expected to be selective with respect to nesting habitat in the ecotone between birch forest and the low-alpine tundra.     

Nest predation in forest birds: influence of predator type and predator's habitat quality

We used the introduction of a generalist nest predator, the red squirrel Tamiasciurus hudsonicus, and of a large herbivore, the Sitka black-tailed deer Odocoileus hemionus sitkensis, to the islands of Haida Gwaii (Queen Charlotte Islands, British Columbia, Canada) to study how predator assemblage and habitat quality and structure influenced nest predation in forest birds. We compared losses of natural nests to predators on islands with and without squirrels. We selected nine islands with or without squirrel or deer and used 506 artificial nests put on the ground or in shrubs to further analyse variation of nest predation with predator assemblage and habitat quality for the predators. For both natural and artificial nests predation risk was higher in presence of squirrels. But predation risk varied within island categories. In presence of squirrels it was highest in stands with mature conifers where it fluctuated from year to year, in response to fluctuations in squirrel abundance. Vegetation cover around the nest had little effect on nest predation by squirrels. Where squirrels were absent, nest predation concentrated near predictable food sources for corvids, the main native predators, and increased with decreasing vegetation cover, suggesting that removal of the vegetation by deer increased the risk of predation by native avian nest predators that use visual cues. Predation risk in these forests therefore varies in space and time with predator composition and with quality of the habitat from the predators' perspective. This temporal and spatial variation in predation risk should promote trade-offs in the response of birds to nest predation, rather than fine-tuned adaptations to a given predation pattern.     

Landscape‐level tree cover predicts species richness of large‐bodied frugivorous birds in forest fragments

Large‐bodied frugivorous birds play an important role in dispersing large‐sized seeds in Neotropical rain forests, thereby maintaining tree species richness and diversity. Conversion of contiguous forest land to forest fragments is thought to be driving population declines in large‐bodied frugivores, but the mechanistic drivers of this decline remain poorly understood. To assess the importance of fragment‐level versus local landscape attributes in influencing the species richness of large‐bodied (>100 g) frugivorous birds, we surveyed 15 focal species in 22 forest fragments (2.7 to 33.6 ha, avg. = 16.0 ha) in northwest Ecuador in 2014. Fragment habitat variables included density of large trees, canopy openness and height, and fragment size; landscape variables included elevation and the proportion of tree cover within a 1 km radius of each fragment. At both the individual species level, and across the community of 12 species of avian frugivore we detected, there was higher richness and probability of presence in fragments with more tree cover on surrounding land. This tendency was particularly pronounced among some endangered species. These findings corroborate the idea that partially forested land surrounding fragments may effectively increase the suitable habitat for forest‐dwelling frugivorous birds in fragmented landscapes. These results can help guide conservation priorities within fragmented landscapes, with particular reference to retaining trees and reforesting to attain high levels of tree cover in areas between forest patches.     

Social rank and antipredator behaviour of Willow Tits Parus montanus in winter flocks

Field experiments on three free-ranging Willow Tit winter flocks, each consisting of one adult pair and two male and two female juveniles (first-year birds), were performed to examine whether preferences for feeding site and antipredator behaviour are related to social rank. The dominance structure was the same in all flocks; adult male > juvenile male 1 > juvenile male 2 > adult female > juvenile female 1 > juvenile female 2.
The proportion of time spent scanning for predators was positively correlated with distance from cover, and adults scanned relatively more than juveniles at the same controlled distance from cover, especially in the afternoon. Given a choice between feeders placed 1 m, 3 m, 5 m, 10 m and 20 m from the forest edge, the tits preferred feeders close to cover. Low-ranking individuals used feeders farther from cover indicating that higher ranked tits prevented them from using the feeders close to cover by means of social dominance. When only the 10m and 20 m feeders were baited, only low-ranked juveniles visited the feeders, subordinate females slightly more than males. The subordinate juveniles increased their use of exposed feeders at low ambient temperature, suggesting that they are prepared to take greater risks during cold periods.
The sequence of return to a feeder, after a life-like stuffed predator model mounted 1 m from a feeder opening was removed, was positively correlated with dominance status, revealing that subordinates take the greatest predation risks.     

Food resource matching by foraging tits Parus spp. during spring-summer in a Mediterranean mixed forest; evidence for an ideal free distribution

We analysed whether patterns of microhabitat use by Blue Tits Parus caeruleus , Great Tits Parus major and Crested Tits Parus cristatus inhabiting a mixed forest consistently matched the patterns of food availability experienced by foraging birds during spring-summer. The use of five microhabitats by each bird species (the foliage of three tree species, shrubs and ground) and the availability of food in trees during the prebreeding, breeding and post-breeding periods of the birds' annual cycle were measured. All three tit species foraged mainly in the outer part of tree canopies (small branches and leaves or needles). Tit distributions between tree species matched food resource distributions irrespective of overall food resource levels, which varied four-fold between the study periods, and tit species. Tits also exploited secondary microhabitats (shrubs and ground) in periods of low food availability; Blue Tits tended to use shrubs, whereas Great and Crested Tits foraged on the ground. Between-trees distributions fitted that expected from an ideal free distribution, suggesting that food availability and intraspecific exploitative competition were the main factors governing tree use by tits. In contrast, patterns of use of secondary microhabitats (shrubs and ground) seemed to indicate a role for the species-specific morphological configurations of each tit species since Blue Tits are better adapted to hang and tended to forage in shubs, whereas Great and Crested Tits are better adapted to feed on horizontal surfaces and tended to forage on the ground. No evidence of interspecific interactions was observed. Overall, the results pointed to an independent exploitation of Mediterranean mixed forest by each bird species, food availability and food accessibility being the main factors affecting microhabitat use by foraging tits.     

The Influence of Habitat Variables on Bird Communities in Forest Remnants in Costa Rica

This study examined the effects of forest structure (tree species richness, canopy height, percent canopy cover, understory density, tree density and DBH) and avian species traits (nest type and indicator list status) on the diversity, abundance and dissimilarity of bird communities in forest remnants and reforestation areas adjacent to Costa Rican banana plantations. Bird species richness and abundance were significantly related to tree species richness, canopy height and canopy cover in multiple linear regressions, the latter two forest structure variables being the best statistical predictors. Stratification of analyses by bird species indicator categories improved fits of regressions, because correlations with environmental variables differed in sign for different guilds of birds, a result likely to hold for other avifaunas. Analysis of avifauna dissimilarities among sites demonstrated that the species composition of bird communities was highly correlated with forest structure and tree species composition. Logistic regressions indicated that birds making protected (cavity, burrow, pendant, sphere and covered) nests were 2–6 times more likely to be present in the study avifauna than birds making open (cup, saucer, platform and scrape) nests and indicators of disturbed habitats were 11 times more likely to be present than indicators of primary forest. The forest structure data used were simple and inexpensive to collect, and data on avian traits were drawn from the literature. Thus, these methods could easily be replicated at other locations and would be valuable management aids and biodiversity assessment tools for conservation planning.     

Short‐term changes in the structure of termite assemblages associated with slash‐and‐burn agriculture in Côte d'Ivoire

Termites are major decomposers in tropical regions and play critical roles in many soil‐related processes. Studies conducted in Asia and the Neotropics suggest that habitat modification can strongly affect termite assemblages, but data on termite communities from forests in Africa, especially West Africa, are scarce. Here, we measured the short‐term impact of slash‐and‐burn agriculture on termite assemblages in an agricultural region of central Côte d'Ivoire. We assessed termite diversity and relative abundance in four habitat types: secondary forest, cleared forest, burned forest, and crop fields. The secondary forest had higher species richness compared with the other habitats, but all habitat types had similar assemblage structures. Fungus‐growing termites were the most abundant feeding group in all habitats. Soil feeders were most abundant in secondary forest, intermediately abundant in cleared and burned forests, and almost entirely absent in crop fields. Wood‐feeding species showed clear responses to burning; their abundances decreased after fire. We conclude that slash‐and‐burn agriculture does not appear to severely erode the diversity of termite assemblages. This could be due to the dominance of ecologically versatile fungus growers or to the relatively long time between clearing and burning. However, forest clearing negatively affects soil feeders, with the Apicotermitinae most affected by canopy loss.     

老君山自然保护区四川山鹧鸪非繁殖期对栖息地的利用(英文)

四川山鹧鸪(Arborophila rufipectus)是中国西南山地特产珍稀鸟类,由于亚热带阔叶林破碎化,其种群数量显著下降。2005年11—12月在老君山自然保护区,共记录四川山鹧鸪非繁殖期栖息地20个。以其新鲜粪便为中心设一个10m×10m大样方和4个1m×1m的小样方,测量每个样方中与栖息地利用有关的12个生境因子(海拔、坡向、坡度、乔木层盖度、灌木高、灌木密度、灌木层盖度、竹盖度、竹密度、雪盖度、林缘距离和道路距离);并在距栖息地100m处任意设对照样方1—2个,共获取30个对照样方,测量相同的生境因子。结果表明,四川山鹧鸪非繁殖期以靠近林间小道和森林边缘的次生落叶阔叶林为栖息地,主要利用海拔高度为1000—1600m,坡度为2—15°的南坡;其偏爱的栖息地竹盖度、竹密度和雪盖度明显小于对照样方,而灌木盖度明显大于对照样方。对栖息地变量进一步分析表明,影响四川山鹧鸪栖息地利用的主要因子依次为地面层食物因子、地型因子、隐蔽因子、温度因子等,其中食物条件的贡献率为29.407%。四川山鹧鸪在利用栖息地时可能面对食物资源与天敌风险的权衡。因此,为了更好地保护四川山鹧鸪栖息地,应重视阔叶林的保护和恢复。     

广西黑颈长尾雉对夜宿地的选择

2003年12月、2004年5月和12月、2005年4月,通过野外调查及设置样方的方法对广西金钟山鸟类保护区黑颈长尾雉的夜宿地选择进行了研究。共记录到12个夜宿地,以夜宿树为中心做一个10m×10m和5个1m×1m的样方,记录夜宿树特征(种类、胸径、栖枝高度、栖枝直径、栖枝上方盖度),测量每个样方的11个生境因子(海拔、林型、坡向、坡度、乔木层盖度、灌木层盖度、草本层盖度、距水源距离、距林缘距离、距道路距离和乔木密度);并随机设10个对照点,测量相同的生境因子。结果表明,黑颈长尾雉夜宿地主要位于阔叶林中,一般,夜间在乔木上休息,一树栖息一只,也有2—3只同宿一树。黑颈长尾雉选择有一定坡度、林下灌木和草本较少以及在森林内部远离林缘并且乔木密度较大的地方作为夜宿地,其夜栖树主要为生境内的中等大小的阔叶树,栖枝直径约为2·5—4cm,栖处高度为2—5m,黑颈长尾雉夜宿地的选择与距水源距离的远近无关。影响黑颈长尾雉夜宿地选择的因素依次为:位置及林下植被、栖枝、天气。影响黑颈长尾雉夜宿地选择的最终原因可能是安全、舒适和栖息地转换的方便程度。     

Predation risk alters interactions among species: competition and facilitation between ants and nesting birds in a boreal forest

Although interactions between species are often assumed to be fixed, theory and empirical evidence suggest that they may be quite variable, changing in the presence of other species or environmental conditions. The interaction between ants and nesting birds exhibits such variability, ants sometimes being predators of bird nests and other times protectors of them. Hypothesizing that predation risk might be a critical factor in altering the interaction of ants with birds, I investigated the interaction of wood ants Formica aquilonia with nesting birds under different levels of predation risk. In a controlled field experiment, I allowed tits ( Parus major, P. caeruleus, P. ater ) and pied flycatchers ( Ficedula hypoleuca ) to select nest boxes in trees with ants (ant trees) or trees without ants. I found that birds usually nested in trees without ants, apparently to avoid the danger of injury from encounters with ants. Nesting in ant trees occurred mainly in the habitat where risk of predation was highest (along the forest edge), and with the bird taxa that lost nests most frequently in trees without ants (tits). Tits nesting on the forest edge achieved significantly greater nesting success, and fledged significantly more young, in ant trees compared with trees without ants. As the season progressed, ant traffic increased in trees without nesting birds, but decreased in trees with nesting birds, indicating that the outcome of interference competition between ants and nesting birds was reversed under increased predation risk. These results support the idea that predation risk can shift species interactions from predominately competitive processes to predominately facilitative processes.     

Habitat-mediated predation risk and decision making of small birds at forest edges

We studied the effect of four components of predation risk, namely predation pressure, spatial variation in woody cover, visibility, and flock size, on the behaviour of four species of the Parus guild (crested tit, P. cristatus , willow tit, P. montanus , coal tit, P. ater , and goldcrest, Regulus regulus ) at the edge of patches of mature boreal forest. Birds used the exposed side of the edge (matrix) mainly during periods with low levels of predation pressure by pygmy owls ( Glaucidium passerinum ). Some species avoided edges under low light conditions. Birds in large groups were more prone to cross the edge, whereas group cohesion increased in risky situations, especially in the most vulnerable species (coal tit and goldcrest). The effects of these three components of predation risk were not general in that only some species responded to them, and in that intra-specific responses were not always consistent. In contrast, all behaviours examined in all four bird species (occurrence, matrix crossing, edge crossing, and group cohesion) appeared to be strongly affected by habitat-mediated predation risk. Mature boreal forests appeared to be qualitatively superior to any type of matrix in terms of protection against predators. Birds generally avoided open matrix, and seemed to move towards and across more developed stages in forest regeneration according to the ratio between food intake and predation risk attributable to a given matrix type. Open matrix (farmland and clearcuts) and very young plantations strongly restricted the rate of bird movement between old forest patches. Our results contrast with the widespread thought that birds have a great potential to use fragmented landscapes in a fine-grained manner. These limited movements across the landscape during winter might have important repercussions on the occupation of some forest patches by tits, their subsequent reproduction, and in turn their population dynamics.     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.