Variation in the relationship between numbers of breeding pairs and woodland area for passerines in fragmented habitat

Species may differ in the relationship between the numbers of breeding pairs present and woodland area, because the proportion of a wood that forms suitable habitat will vary with woodland size. In this paper, we examine the pattern of variation in abundance with woodland area for eight breeding bird species, and also show how this pattern varied between years. During 1990-1997, we made annual censuses of 53–160 woods, of up to 10 ha in size, and fitted a power function to describe the relationships between numbers of breeding pairs and woodland area. Seven of the eight species, blackbird Turdus merula , dunnock Prunella modularis , wren Troglodytes troglodytes , great tit Parus major , chaffinch Fringilla coelebs , robin Erithacus rubecula and blue tit Parus caeruleus showed a pattern of proportionally higher numbers in smaller woods. Only long-tailed tit Aegithalos caudatus occurred in proportionally higher numbers in larger woods. Blackbird and dunnock showed a trend towards lower numbers in large woods during years with low regional population levels; for these species large woods may provide sub-optimal habitat. Great tit, blue tit, chaffinch and robin showed the opposite trend, towards lower numbers in small woods during years with low regional population levels; for these species small woods may provide sub-optimal habitat. Wren and long-tailed tit, which also showed large annual population fluctuations, showed no change in distribution with regional population level. In great tit and chaffinch, the distribution of pairs in any one year may have been influenced by site fidelity producing a lag in the response associated with regional population levels.     

Influences of population size and woodland area on bird species distributions in small woods

Distributions of individual bird species in 151 small woods (size range 0.02–30 ha) were investigated in 3 consecutive years during which the abundance of certain species varied markedly. Relationships between the probabilities of certain bird species breeding and woodland area were described using incidence functions derived from logistic regression analysis. In general, for species which were largely dependent on woodland and seldom occurred in other habitats (such as hedgerows and gardens), the probability of breeding approached 100% only for woods of 10 ha and more, whereas species with less stringent habitat requirements occurred in the majority of woods, including those of 1 ha and less. The sensitivity of incidence functions to changes in regional abundance and the size distribution of the study woods was examined. For some species, distribution patterns could not be distinguished from those expected if pairs had been distributed in proportion to woodland area (random placement), but the majority did not conform to random placement in at least 1 of the 3 years. This nonconformity was consistent across all 3 years for some species, such as wren (Troglodytes troglodytes), despite substantial fluctuations in population sizes between years, while for others, such as robin (Erithacus rubecula), distribution patterns changed with changes in regional abundance. The results suggested that some species, such as wren and blackbird (Turdus merula), preferred small woods, while other species, such as chiffchaff (Phylloscopus collybita), preferred large woods. For several other species, including robin, great tit (Parus major), long-tailed tit (Aegithalos caudatus) and marsh tit (P. palustris), small woods appeared to be sub-optimal under at least some conditions.     

Synchrony of woodland bird populations: the effect of landscape structure

The influence of environmental stochasticity and dispersal in producing patterns in population synchrony was examined for 53 woods censused annually from 1990 to 1999 for nine resident bird species (wren Troglodytes troglodytes , dunnock Prunella modularis , robin Erithacus rubecula , blackbird Turdus merula , song thrush Turdus philomelos , long-tailed tit Aegithalos caudatus , blue tit Parus caeruleus , great tit Parus major , and chaffinch Fringilla coelebs ) and four migrant bird species (garden warbler Sylvia borin , blackcap Sylvia atricapilla , chiffchaff Phylloscopus collybita and willow warbler Phylloscopus trochilus ). Twelve species showed global synchrony of population counts due to regional population trends and widespread annual population fluctuations. There was a clear link between population fluctuations and winter weather for wren, and three other species showed their largest population declines after the coldest winters. Eight species showed a decline in synchrony with distance between woods, and there was evidence for dispersal causing this pattern in three species. Landscape structure affected patterns of synchrony in several species, with lower synchrony in landscapes with less woodland. For three species, this difference in synchrony across a landscape gradient of decreasing woodland cover accounted for the decline in synchrony over distance. Three species showed greater synchrony between woods with similar amounts of hedgerow in the surrounding landscape, suggesting that the surroundings of a wood influence the population dynamics of some species breeding in the wood. Habitat fragmentation can alter the processes contributing to population synchrony. Loss of woodland reduces the relative abundance of woodland bird species. The remaining patches of habitat are smaller, more isolated and are set in a more hostile landscape, all of which may disrupt dispersal between patches and alter the population dynamics within woods.     

Bird species turnover and stochastic extinction in woodland fragments

Year-to-year turnover in bird species composition was recorded across, the whole size range (0 02-30 ha) of 146 woods studied The mean number of resident breeding species both lost and gained per wood between consecutive breeding seasons was 2 (range 0-8) No relationship was found between this absolute turnover rate and woodland area, or any other of 24 predictor variables (describing woodland structure, isolation, connectedness and surrounding land use) Extriction and colonisation rates (in terms of numbers of species lost and gained) were also unrelated to woodland area In all sizes of woods, the species most likely to show local extinctions and colonisations were those with small populations within those woods, but the identity of the species concerned changed as woodland area increased In the smallest woods, the majority of turnover involved common species, such as wren and dunnock, which occurred in only small numbers in these small woods As woodland area increased, these species attained sufficient numbers to usually avoid stochastic extinction The majority of turnover was then due to more specialist (and less numerous) woodland species, such as great-spotted woodpecker and marsh tit, which were usually lacking in small woods In Britain, much existing broadleaved woodland falls within the size range studied Thus the numbers of many bird species are liable to be small enough for yearly turnover in woodland bird communities to be appreciable, and for the long-term persistence of individual species in particular woods to depend on dispersal     

Colonization and extinction dynamics of a declining migratory bird are influenced by climate and habitat degradation

Uncovering the mechanisms involved in the decline of long‐distance migrants remains one of the most pressing issues in European conservation. Since the 1980s, the British breeding population of Garden Warbler Sylvia borin has declined by more than 25%. Here we use data from repeated bird surveys of woodland sites in the 1980s and in 2003–2004 to show that, although the overall population declined between the two periods, the probability of occupancy for this species increased at high latitudes and decreased at low latitudes. Range shifts such as this arise from a change in the ratio of colonizations to extinctions at the range margins, and we therefore related colonization and local extinction at the patch level to concurrent changes in temperature and habitat. The probability of patch colonization by this species was significantly lower where the percentage cover of vegetation in the understorey had declined, reducing habitat quality for this species. The probability of local extinction was significantly correlated with increasing mean May temperature, which may reflect a change in phenology, making breeding conditions less suitable. Changed regimes of grazing and woodland management could be used to increase habitat suitability and thereby increase colonization probability at the local scale, which may in turn increase the probability of patch occupancy despite future climatic unsuitability.     

SONGS OF BLUE TITS PARUS CAERULEUS PALMENSIS FROM LA PALMA (CANARY ISLANDS)—A TEST OF HYPOTHESES

ABSTRACT

Parus major and Parus caeruleus palmensis have some song types in common. They consist of whistled element-types in different frequency ranges. Both species show territorial reactions to playback of the song-types belonging to the other species. P.c. palmensis does not share any songtypes with Central European Parus caeruleus. This is confirmed by playback experiments with the different song types, which do not elicit any territorial behaviour. The songs of P.c. palmensis also differ from those of blue tits from any other Canary Island. They may be derived from contact calls of the Lanzarote population. They have been developed by young birds that had not yet learnt their songs when they colonized La Palma from Lanzarote without reaching any of the other Canary Islands. It is suggested that the uniform song repertoire was developed by adapting the already learned contact calls to each other. The similarities to song types of P. major are due to convergence rather than character displacement. All other Canary Island tit populations share the main features of their songs, which are characterized by quick frequency changes. It seems most likely that these islands have been colonized by birds which had already learnt their songs. Differences in song repertoires are mainly caused by drift events which happen to occur because of population bottlenecks during the colonization process.     

Reductions in connectivity and habitat quality drive local extinctions in a plant diversity hotspot

It is well documented that habitat loss is a major cause of biodiversity decline. However, the roles of the different aspects of habitat loss in local extinctions are less understood. Anthropogenic destruction of an area of habitat causes immediate local extinction but subsequently three additional gradual drivers influence the likelihood of delayed extinction: decreased habitat patch size, lower connectivity and habitat deterioration. We investigated the role of these drivers in local extinctions of 82 declining species in a UK biodiversity hotspot. We combined a unique set of ≈ 7000 vegetation surveys and habitat maps from the 1930s with contemporary species’ occurrences. We extrapolated from these surveys to the whole 2500‐km² study area using habitat suitability surfaces. The strengths of drivers in explaining local extinctions over this 70 yr period were determined by contrasting connectivity, patch size and habitat quality loss for locations at which a species went extinct and those with persisting occurrences. Species’ occurrences declined on average by 60%, with half of local extinctions attributable to immediate habitat loss and half to the gradual processes causing delayed extinctions. On average, locations where a species persisted had a 73% higher contemporary connectivity than those suffering extinctions, but showed no differences in historical connectivity. Furthermore, locations with extinctions experienced a 37% greater decline in suitability associated with changes in habitat type. The strength of the drivers and the proportion of extinctions depended on the species’ habitat specialism, but were affected only minimally by life‐history characteristics. In conclusion, we identified a hierarchy of drivers influencing local extinction: with connectivity loss being the strongest, suitability change being moderately important, but changes in habitat patch size having only weak effects. We suggest conservation efforts could be most effective by strengthening connectivity along with reducing habitat deterioration, which would benefit a wide range of species.     

Effect of Daily Body Mass Variation on the Foraging Behaviour of Tit Species (Parus spp.)

We studied the effect of daily body mass increase on the foraging preferences of two tit species, crested tit, Parus cristatus and blue tit, P. caeruleus, contrasting in morphology and behaviour. We found that both species show a diurnal increase in body mass during winter. Using an experiment with feeders we show that in the crested tit, the proportion of time spent foraging while hanging decreased as body mass increased. In contrast, in the blue tit, foraging behaviour did not change with mass gain. We propose that the species with a morphological design providing a high ecological plasticity in terms of foraging postures can counteract the negative effect of body mass on the ecological options (foraging niche) more than species with a morphological design providing a low ecological plasticity in terms of foraging postures. Our results suggest that blue tits had the advantage of being able to choose to feed on different patches in the habitat throughout the day, which makes resources more predictable for them. In contrast, crested tits might be more restricted in their foraging options as their body mass increases, and this might explain why they hoard food.     

Climate change and breeding parameters of great and blue tits throughout the western Palaearctic

Increasing evidence suggests that climate change has consequences on avian breeding phenology. Here, variations in laying date and clutch size of great tit Parus major and blue tit Parus caeruleus within and between breeding populations through the western Palaearctic are examined in relation to climatic fluctuations, measured by the winter North Atlantic Oscillation (NAO) index. Within and across breeding sites, laying date was related to winter‐NAO index such that great and blue tit females lay earlier after warmer, moister winters (positive values of winter NAO‐index). The present study shows that for most populations there is an advancement of laying date, but the rate of change with respect to NAO significantly differed geographically across the western Palaearctic and did not differ between species. However, clutch size of great and blue tits was not affected by climatic fluctuations, presumably because the whole season is being shifted, but not in relation to food supplies. These combined analyses for the two species controlled for potentially confounding variables such as latitude, longitude, elevation and habitat of each study site.     

Local extinction of British farmland birds and the prediction of further loss

1.  The populations of many UK farmland birds declined between 1970 and 1990, frequently accompanied by contractions in breeding ranges. Ornithological atlas data, land use data and environmental data at the scale of 10-km squares were used to investigate the relationship between local extinctions and habitat suitability for six species, and to predict where future losses are most likely.
2.  For each species we tested the hypothesis that local extinctions were concentrated in environments that were inherently less suitable. We also tested the hypothesis that spatial patterns of loss were not independent between species due to their concurrence in the same habitats.
3.  Multivariate analyses (PCA) showed that areas where each species became extinct between 1970 and 1990 were more similar in land use type, climate and topography to areas where a species was never present than those where it was retained; local extinction was more likely in less suitable environments. Multiple logistic regression showed that for five of the six species the environmental gradient best predicting presence or absence in 1970 was also that best predicting loss between 1970 and 1990. For the six species studied, local extinctions were least likely in lowland arable areas.
4.  For any pair of species, local extinctions were more frequent outside the area of overlap of the two species' ranges than inside. Within the area of overlap, species tended to be lost from the same squares. For each species, likelihood of local extinction declined with increasing number of the other five species present.
5.  We used model parameters to map the probability of future local extinctions of the six species considered, allowing the identification of key areas for conservation management at a spatial scale appropriate to agri-economic incentives.     

Woodland area,species turnover and the conservation of bird assemblages in lowland England

A study over 4 years into the number of breeding bird species and species turnover (extinctions and colonisations) in relation to area was conducted in 35 woodlands, set in an intensively farmed landscape, in north-east Essex, UK. A total of 46 species was recorded. The number of species breeding increased with woodland area; the slope of the species–area relationship did not differ between years. Habitat diversity was the only other measured variable to influence species richness. Absolute species turnover was independent of woodland area but relative turnover declined with increase in woodland area. The numbers of territories of nine species were determined. For four summer visitors the number of woods occupied increased as the overall populations increased but, for the other species, changes in overall population size led to changes in numbers in occupied woods. Chaffinch Fringilla coelebs and Song Thrush Turdus philomelos were more associated with woodland edges, Nightingale Luscinia megarhynchos, Garden Warbler Sylvia borin, Chiffchaff Phylloscopus collybita and Willow Warbler P. trochilus with interiors. Several species showed an inverse relationship between population density and woodland area. Collections of small woods hold similar species richness to single large woods. While the acquisition of large woods for conservation purposes should be a priority, the extension of smaller woods to a size of about 10 ha would be highly beneficial to both the species richness and population stability of regional woodland bird assemblages.     

Density dependent effects between three competitive bird species

Summary Density and breeding success of the great tit Parus major, blue tit Parus caeruleus and collared flycatcher Ficedula albicollis were studied in nest box colony in oak forest over a period of 19 years.Intraspecific density dependent clutch size reduction was found with blue tit and great tit. In interspecific relation the high density of blue tits reduced the clutch size of great tits.In the hatching period neither intraspecific nor interspecific density dependence were showed between the tits when the third competitive species, collared flycatcher was present. The collared flycatcher significantly reduced the hatching success of both tit species and the number of fledglings of great tit.The effects of the great tits and combined density of the great and blue tits on the hatching failure and number of fledglings of collared flycatcher were found when the density of the tits was high. There were not significant relationships to the single density of blue tits.The temporal variability of the competition of the three bird species is discussed.     

Interspecific territoriality in the chaffinch and great tit on islands and the mainland of Scotland: Playback and removal experiments

Territories of the chaffinch (Fringilla coelebs) and great tit (Parus major) overlap on the mainland of Scotland but are mutually exclusive on adjacent islands. Birds on the mainland responded only to playbacks of conspecific song. On the island both species responded to conspecific song and non-conspecific song, but not to the song of the willow warbler. On the island, great tits responded significantly more strongly to songs than did the chaffinches. When chaffinches were removed from part of the island woodland, great tits rapidly expanded into the vacated areas. It is suggested that on the island there is a limit to the co-existence of these two species because they forage in a similar manner and that this has resulted in interspecific territoriality, not seen on the richer mainland environment.     

Rival imprinting: interspecifically cross-fostered tits defend their territories against heterospecific intruders

Failure to recognize conspecifics in social interactions such as mate choice and aggressive encounters will often result in reduced fitness. Studies on mate choice show that the ability to recognize conspecifics as mates is not universally present at birth, but often needs to be learned. In contrast, little is known about the ontogeny of intrasexual species recognition. To test whether learning influences the recognition of sexual rivals, we compared the aggressive response towards intruders of interspecifically cross-fostered individuals and controls reared by conspecific parents. We simulated territorial intrusion by presenting either a caged individual or playback song near the nest of breeding pairs of great tits, Parus major, and blue tits, P. caeruleus. Great tits reared by blue tit parents responded much more to blue tit stimuli than did great tit controls, and furthermore showed stronger responses to blue tit stimuli than to those of their own species. Blue tits reared by great tits responded much more to great tit stimuli than did blue tit controls. In contrast, blue tits cross-fostered to coal tits, P. ater, did not respond more to coal tits than did blue tit controls. There was a species difference in the response to conspecifics: blue tits cross-fostered to great tits responded more to conspecifics than did cross-fostered great tits. The results were similar for males and females. We conclude that learning influences intrasexual species recognition in these tits. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Non‐native salmonids affect amphibian occupancy at multiple spatial scales

Aim The introduction of non‐native species into aquatic environments has been linked with local extinctions and altered distributions of native species. We investigated the effect of non‐native salmonids on the occupancy of two native amphibians, the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), across three spatial scales: water bodies, small catchments and large catchments. Location Mountain lakes at ≥ 1500 m elevation were surveyed across the northern Rocky Mountains, USA. Methods We surveyed 2267 water bodies for amphibian occupancy (based on evidence of reproduction) and fish presence between 1986 and 2002 and modelled the probability of amphibian occupancy at each spatial scale in relation to habitat availability and quality and fish presence. Results After accounting for habitat features, we estimated that A. macrodactylum was 2.3 times more likely to breed in fishless water bodies than in water bodies with fish. Ambystoma macrodactylum also was more likely to occupy small catchments where none of the water bodies contained fish than in catchments where at least one water body contained fish. However, the probability of salamander occupancy in small catchments was also influenced by habitat availability (i.e. the number of water bodies within a catchment) and suitability of remaining fishless water bodies. We found no relationship between fish presence and salamander occupancy at the large‐catchment scale, probably because of increased habitat availability. In contrast to A. macrodactylum, we found no relationship between fish presence and R. luteiventris occupancy at any scale. Main conclusions Our results suggest that the negative effects of non‐native salmonids can extend beyond the boundaries of individual water bodies and increase A. macrodactylum extinction risk at landscape scales. We suspect that niche overlap between non‐native fish and A. macrodactylum at higher elevations in the northern Rocky Mountains may lead to extinction in catchments with limited suitable habitat.     

Behavioural responses to changing landscapes: flock structure and anti-predator strategies of tits wintering in fragmented forests

We analysed the effects of forest fragmentation on the flock structure of insectivorous forest passerines (Parus, Aegithalos, Certhia, Regulus, etc.), and on the anti‐predator behaviour and energy management of blue tits in these flocks. We surveyed flocks in Central Spain during two winters. Flocks in fragments comprised fewer individuals and species than flocks in unfragmented forests. The most abundant species in forest flocks (blue tit, Parus caeruleus, and firecrest, Regulus ignicapillus) were also the most abundant in fragments, while the rarest species in the area never occurred in small woodlots. We investigated how fragmentation and related changes in flock structure affect anti‐predator behaviour of blue tits, a widely distributed species in the area. In fragments but not in forests, blue tits increased scanning rates with decreasing flock size. Vigilance was relaxed when great tits, Parus major, were abundant as flock mates, suggesting that the absence of this dominant species in fragments could intensify anti‐predator behaviour of blue tits. Blue tits enhanced anti‐predator behaviour in the second winter parallel to an increase in the abundance of raptors. This behavioural change was stronger in fragments, where blue tits foraged deeper in the canopy and increased scanning and hopping rates. Under increased predation risk, birds are expected to reduce body mass to improve predator avoidance. On average, blue tits weighed similar in fragments and forests the second winter. However, they accumulated fat along the day in fragments only, and adjusted body mass to body size more closely in that habitat type. This suggests that blue tits perceived fragments as unpredictable habitats where fattening would help avoid starvation, but also as dangerous sites where overweight would further increase the risk of predation. In summary, our results support that fragmentation affects individual behaviour of blue tits, and show the potential of behavioural approaches to unravel how different species face the advancing fragmentation of their habitats.     

Song evolution in the coal tit Parus ater

Since in oscine birds song divergence often leads to premating reproductive isolation, we asked which factors drive the evolution of song in a widespread species, the coal tit Parus ater. Based on an almost taxon‐complete phylogeny we correlated song divergence with morphometric (ecology), genetic (time), and geographic distances (separation). We found eight well‐supported mitochondrial lineages, one of which consists of melanolophus (usually treated as a separate species) and whose relationships remain generally unresolved. Due to intense song variability even sophisticated sonametric analyses failed to separate sub‐specific units. We tested for the role of song in reproductive isolation by playing songs from China and the Himalayas to Central European males. They recognize them as conspecific, but react to their playbacks less aggressively than to local songs. While morphological divergence is correlated with neither time nor separation, song divergence coincides more with geographic than genetic distance. This is mainly driven by differences along longitude and is especially prominent in the southern part of the species’ range, where six lineages persisted during the last glaciation while the north was subsequently re‐occupied by two lineages.     

Relative importance of host plant patch geometry and habitat quality on the patterns of occupancy,extinction and density of the monophagous butterfly <Emphasis Type="Italic">Iolana iolas</Emphasis>

Habitat fragmentation is a major cause of species rarity and decline because it increases local population extinctions and reduces recolonisation rates of remnant patches. Although two major patch characteristics (area and connectivity) have been used to predict distribution patterns in fragmented landscapes, other factors can affect the occurrence of a species as well as the probability of it becoming extinct. In this paper, we study the spatial structure and dynamics of the butterfly Iolana iolas in a 75-patch network of its host plant (Colutea hispanica) to determine the relative importance of patch area, connectivity and habitat quality characteristics on occupancy, extinction and density over the period 2003–2006. Occupancy in 2003, incidence (proportion of years occupied) and probability of extinction were mostly affected by patch area. Smaller patches were less likely to be occupied because they had a higher probability of extinction, partly due to environmental stochasticity. The density of I. iolas was negatively related to patch area in all study years. Only in 2004 was the density of I. iolas positively influenced by fruit production per plant. Our results suggest that for I. iolas, and probably for other specialist butterflies with clearly delimited resource requirements, metapopulation dynamics can be satisfactorily predicted using only geometric variables because most habitat characteristics are subsumed in patch area. However, this hypothesis should be subject to further testing under diverse environmental conditions to evaluate the extent of its generalisation.     

On humans and wildlife in Mediterranean islands

Aim To investigate the effects of human‐induced landscape changes in Mediterranean islands on the ecological and evolutionary responses of bird communities and populations. The combination of mass extinction of large mammals and massive deforestation by humans was hypothesized to produce new selection regimes to which organisms were likely to respond. Habitat selection and niche breadth have been investigated at the scale of species, and phenotypic variation at the scale of local populations. Location The study was carried out along habitat gradients and in habitat mosaics at different spatial scales on the island of Corsica and in areas of similar size and structure in continental France. Methods Two sets of gradients have been used for investigating habitat selection and niche breadth: gradients of altitude, and gradients of vegetation structure. Population studies focused on the blue tit, Cyanistes caeruleus. Large samples of breeding attempts by this species in 10 habitats provided detailed data on phenotypic variation of fitness‐related traits both on Corsica and on the mainland. Results The extent of niche space used by birds differed substantially depending on which habitat gradient was considered. Many species have been found to contract their habitat niche along the elevation gradient on Corsica compared with the mainland, whereas all species in the vegetation gradient broadened their niche on the island. Breeding patterns of the blue tit differed considerably depending on whether they settle in deciduous oaks (Quercus humilis) or in evergreen sclerophyllous oaks (Quercus ilex). Phenotypic variation of breeding traits was much higher on the island, where more populations were correctly timed for the best breeding period than on the mainland, a pattern that is likely to result from lower dispersal of organisms on the island. Main conclusions The differences in observed niche breadth between the two series of habitat gradients is explained both by the species‐specific ecology of the species and the human‐induced environmental history of Corsica. Large‐scale landscape changes provided new opportunities for island colonization by non‐forest species, which are isolated as small, ‘fugitive’ local populations. In both gradients, forest species that are typical components of the Corsican bird fauna definitely expanded their niche and occupied a wider range of habitats on Corsica than on the mainland. At the population scale, landscapes included habitat patches with contrasted selection regimes, which resulted in high phenotypic variation for many fitness‐related traits. Reduced dispersal of birds on the island resulted in a much higher degree of local differentiation on Corsica than on the mainland.