Significant effects of season and bird age on use of coppice woodland by songbirds

Woodland birds have experienced widespread population declines across Europe, resulting partly from a decrease in management practices such as coppicing. Increasing fuelwood demand may reverse the decline of coppicing, making it timely to attempt a fuller understanding of its effects. Here, the impact of coppicing on year‐round habitat use by adults and juveniles of 16 songbird species was quantified from a quasi‐experimental study over 32 years (1978–2009) in Treswell Wood, Nottinghamshire, UK. Habitat use was inferred using capture rates from more than 10 000 h of mist‐netting (> 25 000 captures) and detailed information on coppicing. Capture rates varied with coppice age in different ways: (1) increases as coppice aged (e.g. Marsh Tit Poecile palustris, juvenile Eurasian Treecreepers Certhia familiaris); (2) declines as coppice aged (e.g. Eurasian Blue Tit Cyanistes caeruleus, Great Tit Parus major); (3) peaks in capture rates at intermediate coppice age (i.e. 5–15 years) (e.g. Garden Warbler Sylvia borin, Willow Warbler Phylloscopus trochilus, adult Treecreepers); and (4) a peak at intermediate ages, followed by a decline, before an increase in use again at the oldest coppice ages (i.e. > 20 years) (e.g. Common Blackbird Turdus merula, Eurasian Blackcap Sylvia atricapilla, Eurasian Bullfinch Pyrrhula pyrrhula). Responses to coppice age were similar in different seasons, although Willow Tits Poecile montana showed little preference during breeding but avoided older coppice at other times. Juveniles and adults often differed in their responses to coppice age. The analyses reveal patterns in habitat use that are relevant to woodland management and conservation policy. They suggest that a mosaic of age structures in woodland is beneficial to a wide range of woodland species, and that management should consider the requirements of all age‐classes of birds at different times of year.     

Migration strategies of sylviid warblers: chance patterns or community dynamics?

The effects of community dynamics in birds on the optimisation of their migratory strategies is a neglected area. For three years, we captured migrating warblers on autumn passage at a coastal site in western Britain. We used canonical correspondence analysis (CCA) to assess spatio‐temporal patterns of occurrence, and principal components analysis (PCA) to assess morphological variation. We calculated Euclidean distance in ordination and morphological space to assess separation between species pairs, and used Monte‐Carlo simulations to assess the probability of pattern occurring by chance.
Ordination revealed five species‐groups separated by habitat type and time of passage. Reed Warbler Acrocephalus scirpaceus and Sedge Warbler A. schoenobaenus (Group 1) occurred in wet habitats and peaked simultaneously. In drier habitats with scrub, a first wave of Blackcap Sylvia atricapilla (Group 2) significantly preceded Grasshopper Warbler Locustella naevia, Willow Warbler Phylloscopus trochilus, Whitethroat Sylvia communis and Lesser Whitethroat Sylvia curruca (Group 3), which in all but one case (Lesser Whitethroat) significantly preceded Garden Warbler Sylvia borin (Group 4); peak numbers of Chiffchaffs Phylloscopus collybita and a second wave of Blackcaps (Group 5) occurred later still. Age effects were found only in Acrocephalus, with adults peaking before juveniles.
For seven out of eight pairings within genera, separation in time of passage increased significantly in species that were morphologically similar. The only exception was Blackcap and Lesser Whitethroat which differed substantially in both passage time and morphology. Monte‐Carlo simulations showed that chance was unlikely to be responsible for ordination patterns, nor for inter‐specific variation in passage time and its relationship with species morphology.
These data provide annually consistent evidence that migrating sylviid warblers are separated ecologically by habitat use, time of passage and morphology: we cannot refute the hypothesis that community dynamics have influenced niche use and autumn migratory strategy. We call for further tests of the ‘migrant interaction’ hypothesis in other geographical locations and taxa, particularly where migrants are allopatric and interact ecologically only on migration.     

The effects of prey size on diet differentiation of seven passerine species at two spring stopover sites

Prey size was evaluated for seven passerine trans-Saharan migrant species at two spring stopover sites in Sardinia, Italy. The species considered were Pied Flycatcher Ficedula hypoleuca, Spotted Flycatcher Muscicapa striata, Redstart Phoenicurus phoenicurus, Garden Warbler Sylvia borin, Whitethroat Sylvia communis, Willow Warbler Phylloscopus trochilus and Wood Warbler Phylloscopus sibilatrix. The analysis was made for three prominent prey types: beetles (Coleoptera), ants (Hymenoptera: Formicidae) and “flying prey” (wasps and bees [Hymenoptera, excluding ants] and flies, Diptera, pooled), The prey size distribution in the diets of some species was very similar to that caught in our insect traps, showing that our estimates of availability are, at least partially, relevant. On the whole, diets deviated from food available in a species consistent way. The size distributions of “flying prey” differed between the two sites but were almost constant in the birds' diets. In contrast, size distributions of available beetles and ants at the sites were similar but were different in the birds' diets. Different feeding behaviour of the birds, in terms of physiological constraints during migratory stopovers, is discussed. Paired species comparisons show that the diets of most species differ significantly in the distribution of the size classes of at least some prey types. The conclusions drawn from the comparisons of the size distributions of all prey items collected from a bird species in one season are very similar to the conclusions drawn from comparisons based on the presence/absence of a size class per faecal sample. We compared the similarities of diet with and without using information on prey size. With the exception of the Pied Flycatcher and the Redstart at one of the study sites, size information did not add to diet segregation. Certain size classes within prey types tended to be common in the diet of these migrant passerines. However, specialization on certain size classes within broad taxonomic categories was not evident.     

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.     

WEIGHTS OF SOME PALAEARCTIC MIGRANTS IN SOUTHERN UGANDA

Between March 1966 and May 1968 Palaearctic passerines were mist-netted in thick bush and lightly wooded savannah habitats near Kampala, on the northern shore of Lake Victoria. This paper reports weights of the seven principal species involved. Most migrants appeared to be in a lean condition during the winter months, when weights were relatively low and varied little in each species. Birds were not particularly light on arrival. In fact, autumn Garden Warblers Sylvia borin and Willow Warblers Phylloscopus trochilus were sometimes markedly heavy, and for the former species there was some evidence that the individuals concerned were passage migrants. Autumn weights of Swallows Hirundo rustica, Reed Warblers Acrocephalus scirpaceus and Yellow Wagtails Motacilla flava were similar to those recorded in winter. The mean weight of all species rose during late March or early April. Although most Garden Warblers and Willow Warblers trapped at the time of spring migration were within the normal winter weight range, many Acrocephalus warblers and the majority of Sand Martins Riparia riparia and Yellow Wagtails were rather heavy. Spring weights 40% or more above mean winter weight were not uncommon in the Sedge Warbler Acrocephalus schoenobaenus, but were recorded only occasionally in other species. Although most passerine migrants evidently left Kampala with substantial fat reserves, it was concluded that a considerable number of warblers departed at rather low weight. High spring weights were mainly confined to a period of two or three weeks in each of the warbler species. Locally wintering Acrocephalus warblers must have attained full premigratory weights within three weeks, and a number of spring retraps showed substantial gains at minimum mean rates of between 0–1 and 0–35 g per day. Most heavy Garden Warblers were probably on passage. Significant correlations between weight and wing-length were obtained for all species investigated, regressions of weight on wing-length being in the range 011-0-25 g/mm. Spring weights are briefly compared with data from Nigeria, and the northward migration of passerines from Lake Victoria is discussed.     

Seasonal differences in energy requirements of Garden Warblers Sylvia borin migrating across the Sahara desert

The Sahara desert acts as an ecological barrier for billions of passerine birds on their way to and from their African wintering areas. The Garden Warbler Sylvia borin is one of the most common migrants involved. We used body mass of this species from Greece in autumn and spring to simulate the desert crossing and to assess how body mass relates to fuel requirement. The flight range estimates were adjusted to the seasonal extent of the desert, 2200 km in autumn and about 2800 km in spring. In autumn, with an average fuel load of about 100% of body mass without fuel, birds were not able to cross the desert in still air, but northerly winds prevail during September and with the average wind assistance only one in 14 was predicted to fail to make the crossing. Body mass data from spring, after the desert crossing, was used to estimate departure body mass from south of the desert. The average wind assistance in spring is close to zero and departure body mass of the average bird arriving at Antikythira, a small Greek island, under such conditions was estimated to be 34.6 g, which corresponded to a fuel load of 116%. Calculations based on 1% body mass loss per hour of flight showed slightly larger body mass loss than that calculated from flight range estimates. The results suggest that passerine birds about to cross the eastern part of the Sahara desert need to attain a larger fuel load in spring than in autumn.     

Body fat influences departure from stopover sites in migratory birds: evidence from whole-island telemetry

Migration remains one of the great mysteries of animal life. Small migratory birds rely on refuelling stopovers after crossing ecological barriers such as deserts or seas. Previous studies have suggested that fuel reserves may determine stopover duration but this hypothesis could not be tested because of methodological limitations. Here, we provide evidence that subcutaneous fat stores determine stopover duration by measuring the permanence of migratory garden warblers (Sylvia borin) on a small Mediterranean island during spring migration with telemetry methods. Garden warblers with large amounts of fat stores departed the island significantly sooner than lean birds. All except one fat bird left the island on the same evening after capture, with a mean total stopover estimate of 8.8 hours. In contrast, the mean estimated total stopover duration of lean birds was 41.3 hours. To our knowledge, this is the first study that measures the true minimum stopover duration of a songbird during migration.     

Spring migration timing of Sylvia warblers in Tatarstan (Russia) 1957–2008

The timing of when migrant birds return to breed is a key component of studies of the impact of climate change upon bird populations. However, such data are not distributed evenly across the World, and in the Northern Hemisphere are underrepresented in Asia and the east of Europe. Therefore, to help rectify this bias, we analysed first arrival dates (FAD) of four species of Sylvia warblers (Blackcap Sylvia atricapilla, Whitethroat S.communis, Lesser Whitethroat S.curruca and Garden Warbler S.borin) collected in the Tatarstan Republic of Russia between 1957 and 2008. Over the whole period the species returned to their breeding sites between three and six days earlier; these trends were significant except for Whitethroat. Advances in arrival were especially apparent in the two earlier species, Blackcap and Lesser Whitethroat, mainly because local temperatures for March had risen substantially. Except for Whitethroat, FADs were significantly related to temperatures in the African wintering ground and/or in Tatarstan. Whilst significant correlations occurred between FADs of some of the species, there was considerable variability in these relationships indicating a species-specific response to rising temperatures. Changes in FADs in this eastern extremity of Europe were smaller than in Central and Western Europe.     

Foraging behavior of three species of songbirds during stopover in southeastern Morocco during spring migration

Investigators studying the stopover ecology of migrating birds typically use the capture–recapture method to examine important parameters such as fuel deposition rates (FDR) and stopover duration. However, such studies can be constrained by the number of recaptures. An alternative method is to calculate a regression of mass over time of day, but this method may not be reliable because patterns of mass change of individual birds through the day may not reflect that of the whole population. Given the potential constraints of these methods, using them in combination with other methods, such as behavioral observations of foraging birds, may improve our understanding of the patterns of fuelling in birds at stopover sites. We observed the foraging behavior of three songbird species, including Western Bonelli's (Phylloscopus bonelli), Subalpine (Sylvia cantillans), and Willow (Phylloscopus trochilus) warblers, from 15 March to 30 April 2011 at a small oasis at the northern border of the Sahara desert in southeast Morocco. Given the location of our study site at the northern edge of the Sahara desert, birds migrating north likely needed to replenish their energy reserves at this stage of their journey. We assessed foraging effort by determining the rate (number per unit time) at which birds pecked at substrates or made aerial forays after flying insects. Peck rates were higher for Western Bonelli's Warblers than for Subalpine and Willow warblers, suggesting either species‐specific adaptations to feeding in arid environments or differences in the motivation to feed. In addition, Western Bonelli's Warblers had FDRs that were negative or close to zero and, therefore, were apparently unable to refuel successfully (i.e., increase their fuel stores) despite greater effort, possibly indicating less efficiency in obtaining food (i.e., more unsuccessful pecks). The lower peck rates of Subalpine and Willow warblers suggest either that they were less efficient at finding prey or were simply foraging at lower rates. For all three species, peck rates were lower at higher wind speeds, suggesting that wind may alter prey availability and detectability, especially of flying insects. Interactions among species‐specific migration strategies, environmental conditions, and habitat quality ultimately define the success of migration. Our results suggest that using observational data in combination with capture data may improve our understanding of these interactions at migration stopover sites.     

Nectar consumption of warblers after long-distance flights during spring migration

The consumption of nectar by European passerines has been reported only occasionally. In this study we investigated the occurrence and significance of nectar consumption of small passerine birds on spring migration after crossing the Mediterranean Sea. On Ventotene Island in the Tyrrhenian Sea, four migrating species of Sylvia warblers [Garden Warbler S. borin , Subalpine Warbler S. cantillans , Whitethroat S. communis , Blackcap S. atricapilld ) regularly foraged on the two most common flowering species at that time of year, Brassica fruticulosa (Cruciferae) and giant fennel Ferula communis (Umbelliferae), while other species visited flowers only occasionally or not at all. Feeding behaviour, pollen traces on the head, and the examination of pollen and sugar remains in droppings indicated that nectar was the main target of the Sylvia warblers, rather than pollen or insects on the flowers. This was confirmed by food choice experiments indicating a clear preference by Garden Warblers and Whitethroats for nectar from artificial flowers over mealworms. Although conclusive experiments are not available, we hypothesize that nectar might be a diet easy to obtain and to absorb for birds after a long-distance flight in which they have incurred a depletion of energy stores and a reduction of the digestive tract.     

Morphological sexing of passerines: not valid over larger geographical scales

Sex determination of birds is important for many ecological studies but is often difficult in species with monomorphic plumage. Morphology often provides a possibility for sex determination, but the characters need to be verified. We tested whether five passerine species can be sexed according to standard morphological measurements applying a forward logistic regression with sex determined by molecular analysis as the dependent variable. Furthermore, we tested whether the results can be used on a larger geographic scale by applying morphological sexing methods gained by similar studies from other regions to our data set. Of the five species of this study only Garden Warblers Sylvia borin could not be sexed morphologically. In the Robin Erithacus rubecula, 87.2% of all individuals were sexed correctly. For Reed Warblers Acrocephalus scirpaceus, Willow Warblers Phylloscopus trochilus and Reed Buntings Emberiza schoeniclus, the respective values were 77.6, 89.4 and 86.4%. When the logistic regression functions from similar studies on Robins and Reed Buntings in Denmark and Scotland were applied to the birds from south-western Germany, they performed less well compared to the original dataset of these studies and compared to the logistic regression function of our own study. The same was the case for Willow Warblers when a wing length criterion used in Great Britain was applied to the birds of our study. These discrepancies may have several explanations: (1) the models are optimised for the dataset from which they were extracted, (2) inter-ringer variation in measurements, (3) the use of different age cohorts, (4) different morphology due to different habitat availability around the study site, or, most likely, (5) different morphology due to different migratory behaviour. We recommend that morphological sex differentiation methods similar to this study (1) be only used population specific, (2) only with one age cohort and (3) to adjust the extracted equations from time to time.     

Flight-range estimates for small trans-Sahara migrants

Arguments in support of the non-stop and the intermittent strategies for crossing the Sahara have been based on data on the fat reserves of birds before the crossing and of birds grounded in the desert. In this paper, flight-range estimates were calculated and the necessary assumptions about air speed, energy input during flight, and energy equivalent of body reserves were evaluated. As examples, Willow Warblers Phylloscopus trochilus and Garden Warblers Sylvia borin were investigated during autumn migration from two study sites north of the Sahara and two study sites in the desert. In still air, the flight-range for both species at all study sites was too short to reach the Sahel zone without refuelling. It is concluded that birds depend on tailwinds for a successful crossing, independent of a non-stop or an intermittent migratory strategy, and that weather conditions in autumn allow them to rely on tailwinds.     

Temporal and spatial variation in the occurrence of Palearctic warblers around Lake Victoria

Goudswaard, K. & Wanink, J.H. 2000. Temporal and spatial variation in the occurrence of Palearctic warblers around Lake Victoria. Ostrich 71 (1 & 2): 210–212.

Only one Palearctic warbler species, the Sedge Warbler, wintered at the south-eastem shores of Lake Victoria. Willow Warbler and Garden Warbler occurred mainly on passage, with peak values in the first lunar quarter, the period in which lakeflies would normally swarm. Higher warbler densities have been reported from the north-western shores. This agrees with our idea that the prevailing eastern winds will result in a higher availability of lakeflies to warblers at the western side of the lake.     

Fall migration of Empidonax flycatchers in northwestern Colombia

ABSTRACT South of Mexico, little is known about the fall migration patterns of most Neotropical migrants. I studied the migration of Empidonax flycatchers using mist‐net surveys in northwestern Colombia from late September to mid‐October in 4 yr (2003–2005, 2008). Empidonax species were identified using linear measurements and color patterns. About 62% of captured individuals were reliably identified to species, with 86% identified as Willow Flycatchers (Empidonax traillii) and 14% as Alder Flycatchers (E. alnorum). No Acadian Flycatchers (E. virescens) were identified. Most birds captured were adults (84.9%) and, due to overlap in measurements, I was only able to determine the sex of 16.3% of the birds. Most Empidonax flycatchers migrated through northwestern Colombia during September and October, with individuals migrating through my study area over a period of at least 1 mo. Willow Flycatchers tended to migrate earlier than Alder Flycatchers, a pattern consistent with the fall movements of these two species at other locations. No captured flycatchers were molting either remiges or rectrices, and most (89%) had either no or slight traces of subcutaneous fat. No Empidonax flycatchers were recaptured, suggesting that stopover duration at my study site was brief. My results show that many Empidonax flycatchers can be identified as Willow and Alder flycatchers during the nonbreeding period, and such identification will enhance our knowledge of their geographical distribution and improve our understanding of possible patterns of segregation on their wintering grounds.     

Endemic Cyprus Warbler Sylvia melanothorax and colonizing Sardinian Warbler Sylvia melanocephala show different habitat associations

Anthropogenic habitat change and assisted colonization are promoting range expansions of some widespread species with potential consequences for endemic fauna. The recent colonization of Cyprus by breeding Sardinian Warblers Sylvia melanocephala has raised concerns that it might be displacing the closely related and endemic Cyprus Warbler Sylvia melanothorax. Habitat associations of both species were examined using models of abundance within the 95% density kernel of the Sardinian Warbler’s range and also outside this range for Cyprus Warbler. Within the Sardinian Warbler’s range, the two species were associated with subtly different scrub habitats. Outside the Sardinian Warbler’s range, the Cyprus Warbler differed again in its habitat association, but this probably resulted from marked differences in habitat extent and availability in different parts of the island rather than from competitive displacement, as none of the habitat or land‐use elements differentially associated with Cyprus Warblers was positively associated with Sardinian Warbler occurrence. This suggests that the Sardinian Warbler has exploited a different niche, rather than displacing the endemic species, and has perhaps benefitted from changing land‐use patterns, particularly recent fallows and abandoned agriculture, in contrast to the stronger association of Cyprus Warblers with semi‐natural scrub.     

Some aspects of frugivory by bird populations using coastal dune scrub in Lincolnshire

The fruit diets of Sylvia warblers, and some thrushes, were studied during 1989 at a coastal scrubland site, using faecal examination. Fruit feeding commenced as the earliest berries ripened in late July, but there were differences of timing between species. Amongst both warblers and thrushes, the larger species commenced earlier and ate more fruit. Almost all Blackcap and Garden Warbler faeces examined from early August contained some fruit remains, as did most Whitethroat faeces from mid-month. Mean fruit contents of faecal samples exceeded 80% infrequently, indicating that even the most frugivorous species always took other food, normally insects, to supplement their diet. Each species differed in their fruit diet: Blackcaps, and to a lesser extent Garden Warblers, ate Woody Nightshade berries in preference to Blackberries, as they are structurally adapted for feeding on the former plant. Only inexperienced juveniles of other species regularly chose the Nightshade berries. Elder berries, abundant from late August, became the preferred fruit for all Sylvia species. Using data from an earlier investigation, 1 it is shown that by feeding on Elder berries, migrating Blackcaps can obtain 75–90% of their daily energy requirements in 10% of daylight hours.     

Long‐term changes in the migration phenology of UK breeding birds detected by large‐scale citizen science recording schemes

The timing of migration is one of the key life‐history parameters of migratory birds. It is expected to be under strong selection, to be sensitive to changing environmental conditions and to have implications for population dynamics. However, most phenological studies do not describe arrival and departure phenologies for a species in a way that is robust to potential biases, or that can be clearly related to breeding populations. This hampers our ability to understand more fully how climate change may affect species’ migratory strategies, their life histories and ultimately their population dynamics. Using generalized additive models (GAMs) and extensive large‐scale data collected in the UK over a 40‐year period, we present standardized measures of migration phenology for common migratory birds, and examine how the phenology of bird migration has changed in the UK since the 1960s. Arrival dates for 11 of 14 common migrants became significantly earlier, with six species advancing their arrival by more than 10 days. These comprised two species, Blackcap Sylvia atricapilla and Chiffchaff Phylloscopus collybita, which winter closest to Britain in southern Europe and the arid northern zone of Africa, Common Redstart Phoenicurus phoenicurus, which winters in the arid zone, and three hirundines (Sand Martin Riparia riparia, House Martin Delichon urbicum and Barn Swallow Hirundo rustica), which winter in different parts of Africa. Concurrently, departure dates became significantly later for four of the 14 species and included species that winter in southern Europe (Blackcap and Chiffchaff) and in humid zones of Africa (Garden Warbler Sylvia borin and Whinchat Saxicola rubetra). Common Swift Apus apus was the exception in departing significantly earlier. The net result of earlier arrival and later departure for most species was that length of stay has become significantly longer for nine of the 14 species. Species that have advanced their timing of arrival showed the most positive trends in abundance, in accordance with previous studies. Related in part to earlier arrival and the relationship above, we also show that species extending their stay in Great Britain have shown the most positive trends. Further applications of our modelling approach will provide opportunities for more robust tests of relationships between phenological change and population dynamics than have been possible previously.     

Garden Warbler Sylvia borin migration in sub-Saharan West Africa: phenology and body mass changes

The Garden Warbler is a classic subject for the study of Palaearctic–African bird migration strategies. Most studies have considered the situation close to the breeding areas, while the African and especially the sub‐Saharan part of the species’ migration have received comparatively little attention. Here we use autumn and spring ringing data from Nigeria and The Gambia to study the movements and energetics of the species in West Africa during the non‐breeding season. The first Garden Warblers arrive south of the desert around the beginning of September, roughly at the same time as the median date for their passage through the Baltic Sea region and c. 3 weeks before their median passage date through southern Italy. In the Nigerian Sahel savannahs, where, owing to the rainy season and its associated increase in food availability, many more Garden Warblers stop over in autumn than in the dry spring, the median date of passage is 1 October. The body mass on arrival south of the desert is normally only a few grams more than the lean body mass (LBM; 15 g) – with a mean of 16.6 g (sd = ±1.8 g) in The Gambia and 17.4 g (sd = ±1.8 g) in the Nigerian Sahel. After resting and refuelling in the Sahel, Sudan and Guinea‐type savannahs the Garden Warblers depart during November–December for wintering areas further south. Before leaving, they again increase their body mass, with an average fuel load of c. 20%, and often more than 50% relative to LBM. Some of the birds passing through Nigeria probably spend midwinter around the Congo Basin. During spring they return northwards to the Guinea savannah zone in April and fuel‐up there for the trans‐Sahara passage. At this time they normally increase their body reserves to around 50% of the LBM, but c. 10% of the birds gain 100%, thus doubling their mass. The passage there peaks around 20 April and continues well into May. That the main take‐off northwards is directly from the Guinea savannahs is indicated by the very low numbers trapped in the Sahel during spring.     

Adding ecology into phylogeography: ecological niche models and phylogeography in tandem reveals the demographic history of the subalpine warbler complex

Capsule: This study documents evidence of interglacial refugia during the Last Interglacial for birds in the Mediterranean region, and emphasizes the importance of the Last Interglacial on the geographic distribution and genetic structure of Mediterranean species.

Aims: We focused on the historical biogeography of the subalpine warbler complex: Subalpine Warbler Sylvia cantillans and Moltoni’s Warbler Sylvia subalpina; we tested if this Mediterranean bird complex shared a similar demographic fate as the present-day widespread species in the temperate zones of Europe, through the late Quaternary glacial-interglacial cycles.

Methods: An ecological niche model was developed to predict the geographic distribution of the subalpine warblers under the past (the Last Interglacial and the Last Glacial Maximum) and the present bioclimatic conditions. Additionally, Bayesian Skyline Plot analysis was used to assess effective population size changes over the history of the subalpine warbler complex.

Results: During the Last Glacial Maximum, the subalpine warblers almost reached their current distribution in the Mediterranean region; yet, unlike the widespread temperate bird species, they survived the Last Interglacial in allopatric refugia in the Mediterranean region.

Conclusion: A unique biogeographic pattern was revealed, indicating the importance of the Last Interglacial on current distributional patterns and demographic histories of common bird species in the Mediterranean region. This study suggests that Mediterranean biogeography is far more complex than previously assumed, and so deserves further study and more attention.     

Numerical and functional responses of breeding passerine species to mass occurrence of geometrid caterpillars in a subalpine birch forest: a 30-year study

In northern Fennoscandia, the geometrid moths Epirrita autumnata and Operophtera brumata have cyclicities in density with mass occurrence at 10‐year intervals. The larvae of Epirrita and Operophtera attain a size of 2–3 cm and 1.5–2 cm, respectively, and are nutritious food items for passerine birds. To examine whether these larvae have any numerical and/or functional influence on a passerine bird community (mountain birch forest in Budal, central Norway) during a 30‐year period (1972–2001), I estimated their abundance (number of larvae per 100 sweeps) in the birch canopy, and the densities of breeding birds in the passerine community. In addition, from 1972 to 1998, I monitored the nesting success of five of the bird species. The foraging pattern of the most abundant bird species and their gizzard contents (adults and nestlings) were examined in 1972–78 (covering population peaks of both the geometrids). Population peaks of Epirrita occurred in 1975–76, 1985–86 and 1996, and of Operophtera in 1976–77, 1986–87 and 1997–98. The passerine community consisted of eight species that were territorial in all 30 years, one species in 26 years, three species in 14–21 years and three species in 1–4 years. Only the Brambling Fringilla montifringilla population responded numerically to the fluctuations of Epirrita and Operophtera. Brambling was also the only species whose mean clutch size varied between years, and this correlated positively with the density of Epirrita. The mean annual nesting success of Willow Warbler Phylloscopus trochilus, Bluethroat Luscinia svecica and Common Redpoll Carduelis flammea tended to be higher in years with mass outbreaks of Epirrita, but was significantly so only for Reed Bunting Emberiza schoeniclus. The abundance of Operophtera larvae showed no influence on the nesting success of any bird species. The passerines foraged more frequently in the birch canopy in the Epirrita outbreak years (1975–76) than in the years before or after. Gizzard analyses of five adult passerine species and their nestlings showed that Epirrita was the main food item in 1974–76. Even though Operophtera occurred in large numbers in birch trees in 1976 and 1977, only a few larvae were found in the gizzards of the passerines. None of the passerines showed an increase in their population density in the year following the larval outbreaks, but the densities of Willow Warbler and Bluethroat increased in the succeeding year, indicating a higher return rate for these species. The study shows the existence of a dietary response and also indicates a reproductive response to the changes in the abundance of Epirrita in mountain birch forest. The lack of numerical response in the passerines (except the Brambling) to the fluctuation in Epirrita contrasts with the pattern described for passerine communities in northern temperate deciduous forests in North America, where Lepidoptera caterpillars periodically have mass outbreaks.