Climatic responses in spring migration of boreal and arctic birds in relation to wintering area and taxonomy

Large‐scale climate fluctuations, such as the North Atlantic Oscillation (NAO), have a marked effect on the timing of spring migration of birds. It has however been suggested that long‐distance migrants wintering in Africa could respond less to NAO than short‐distance migrants wintering in Europe, making them more vulnerable to climatic changes. We studied whether migratory boreal and arctic bird species returning from different wintering areas show differences in responses to the NAO in the timing of their spring migration. We used data on 75 species from two bird observatories in northern Europe (60°N). By extending the examination to the whole distribution of spring migration and to a taxonomically diverse set of birds, we aimed at finding general patterns of the effects of climate fluctuation on the timing of avian migration. Most species arrived earlier after winters with high NAO index. The degree of NAO‐response diminished with the phase of migration: the early part of a species’ migratory population responded more strongly than the later part. Early phase waterfowl responded strongest to NAO, but in later phases their response faded to non‐significant. This pattern may be related to winter severity and/or ice conditions in the Baltic. In the two other groups, gulls and waders and passerines, all phases of migration responded to NAO and fading with phase was non‐significant. The difference between waterfowl and other groups may be related to differences between the phenological development of their respective macrohabitats. Wintering area affected the strength of NAO response in a complicated way. On average medium distance migrants responded most strongly, followed by short‐distance migrants and partial migrants. Our results concerning the response of long‐distance migrants were difficult to interpret: there is an overall weak yet statistically significant effect, but patterns with phase of migration need further study. Our results highlight the importance of examining the whole distribution of migration and warrant the use of data sets from several sampling sites when studying climatic effects on the timing of avian life‐history events.     

Timing of spring migration in birds: long-term trends, North Atlantic Oscillation and the significance of different migration routes

We studied long-term trends and the yearly variation in mean spring passage time in 36 passerine bird species trapped at Ottenby Bird Observatory in south-eastern Sweden. Between the years 1952–2002, data were available for 22–45 years depending on species. Most long-distance migrant species passed progressively earlier over the study period (range: 2.5 days earlier to 0.7 days later per 10 years, with an average of 0.9 days earlier per 10 years). The annual variation in timing of migration in most species, regardless of migration distance, correlated negatively with the winter index of the North Atlantic Oscillation (NAO), a large-scale climate phenomenon influencing the climate in the North Atlantic region. Birds passed earlier after mild and humid winters, corresponding to the high phase of the NAO. This corroborates the pattern found at a nearby migration site with a comparable dataset (Helgoland, 600 km WSW of Ottenby). However, short/medium-distance migrant species at Ottenby, in contrast to the situation at Helgoland, have shown no general trend of earlier passage in recent years. This was probably a consequence of the shorter study period at Ottenby, which included only the last 22–32 years (41 years at Helgoland), when the NAO showed no significant trend. At the species-specific level, the long-term trends in passage time were similar at the two sites, and there was some congruence to the extent that a given species was affected by NAO. Long-distance migrants wintering south and south-east of the breeding grounds showed some of the strongest changes in long-term trends (passing progressively earlier) at Ottenby, and for some of these species passage time varied negatively with NAO. Obviously, and contrary to previous suggestions, variations in NAO also influence birds migrating through eastern Europe, although the direct or indirect mechanisms through which this is achieved are unknown.     

Influences of the El Niño/Southern Oscillation and the North Atlantic Oscillation on avian productivity in forests of the Pacific Northwest of North America

To model the effects of global climate phenomena on avian population dynamics, we must identify and quantify the spatial and temporal relationships between climate, weather and bird populations. Previous studies show that in Europe, the North Atlantic Oscillation (NAO) influences winter and spring weather that in turn affects resident and migratory landbird species. Similarly, in North America, the El Niño/Southern Oscillation (ENSO) of the Pacific Ocean reportedly drives weather patterns that affect prey availability and population dynamics of landbird species which winter in the Caribbean. Here we show that ENSO‐ and NAO‐induced seasonal weather conditions differentially affect neotropical‐ and temperate‐wintering landbird species that breed in Pacific North‐west forests of North America. For neotropical species wintering in western Mexico, El Niño conditions correlate with cooler, wetter conditions prior to spring migration, and with high reproductive success the following summer. For temperate wintering species, springtime NAO indices correlate strongly with levels of forest defoliation by the larvae of two moth species and also with annual reproductive success, especially among species known to prey upon those larvae. Generalized linear models incorporating NAO indices and ENSO precipitation indices explain 50–90% of the annual variation in productivity reported for 10 landbird species. These results represent an important step towards spatially explicit modelling of avian population dynamics at regional scales.     

Ecological conditions in wintering and passage areas as determinants of timing of spring migration in trans-Saharan migratory birds

1.?Climate change has been associated with shifts in the timing of biological events, including the spring arrival of migratory birds. Early arrival at breeding sites is an important life-history trait, usually associated with higher breeding success and therefore, susceptible to selection and evolution in response to changing climatic conditions. 2.?Here, we examine the effect of changes in the environmental conditions of wintering and passage areas on the mean passage time of 13 trans-Saharan passerines during their spring migration through the western Mediterranean over the 15 years from 1993 to 2007. 3.?We found that most of the species studied have been advancing the timing of their passage in recent years. However, annual variation in the mean date of passage was positively correlated with vegetation growth (measured as the normalized difference vegetation index [NDVI]) both in the Sahel (the region of departure) and in northern Africa (the passage area). Thus, migration dates were delayed in years with high primary productivity in passage and wintering zones. All species seem to respond similarly to NDVI in the Sahel; however, late migrants were less affected by ecological conditions in northern Africa than those migrating earlier, suggesting differences based on species ecology. 4.?Mean timing of passage was not related to the North Atlantic Oscillation (NAO), El Ni?o-Southern Oscillation (ENSO), temperature or NDVI in the species-specific wintering areas (the overwintering region) when analysed in combination with the other covariates. 5.?Our findings show that ecological conditions in the winter quarters (specifically the Sahel) and en route are relevant factors influencing trends in the passage dates of trans-Saharan migratory birds on the southern fringe of Europe. Possible long-term consequences for late arriving spring migrants are discussed.     

Central European Greylag Geese Anser anser show a shortening of migration distance and earlier spring arrival over 60 years

Global climate change can cause pronounced changes in species? migratory behaviour. Numerous recent studies have demonstrated climate‐driven changes in migration distance and spring arrival date in waterbirds, but detailed studies based on long‐term records of individual recapture or re‐sighting events are scarce. Using re‐sighting data from 430 marked individuals spanning a 60‐year period (winters 1956/1957 to 2015/2016), we assessed patterns in migration distance and spring arrival date, wintering‐site fidelity and survival in the increasing central European breeding population of Greylag Geese Anser anser. We demonstrate a long‐term decrease in migration distance, changes in the wintering range caused by winter partial short‐stopping, and the earlier arrival of geese on their breeding grounds. Greylag Geese marked on central Europe moulting grounds have not been recorded wintering in Spain since 1986 or in Tunisia and Algeria since 2004. The migration distance and spring arrival of geese indicated an effect of temperature at the breeding site and values of the NAO index. Greylag Geese migrate shorter distances and arrive earlier in milder winters. We suggest that shifts in the migratory behaviour of Central European Greylag Geese are individual temperature‐dependent decisions to take advantage of wintering grounds becoming more favourable closer to their breeding grounds, allowing birds to acquire breeding territories earlier.     

The influence of year-to-year variations in winter weather on the dynamics of Daphnia and Eudiaptomus in Esthwaite Water, Cumbria

1. Micro-crustacea of the genus Daphnia and Eudiaptomus have evolved different physiological mechanisms for coping with life in a rapidly changing environment. In this paper, we analyse some of the physical and biological factors influencing the winter abundance of the two species in a small lake (Esthwaite Water in Cumbria).
2. The results demonstrate that much of the year-to-year variation in their relative abundance can be related to long-term changes in the weather. The highest numbers of Daphnia were typically found in cold, calm winters when small flagellates were relatively abundant. In contrast, the highest numbers of Eudiaptomus were found in mild, windy winters when the phytoplankton community was dominated by colonial diatoms.
3. Year-to-year variations in the winter abundance of Eudiaptomus had no effect on their subsequent development but the numbers of overwintering Daphnia had a significant effect on the size of their first spring 'cohort'. The most important factor influencing the overwintering performance of the two species was the water temperature.
4. Winter air temperatures over much of Europe are influenced by the atmospheric pressure variation known as the North Atlantic Oscillation (NAO). Winter water temperatures in Esthwaite Water were strongly correlated with this empirical index and there was a significant positive correlation between the NAO and the number of overwintering Eudiaptomus.     

Winter climate affects subsequent breeding success of common eiders

The phenology of spring migration depends on the severity of the preceding winter and approaching spring. This severity can be quantified using the North Atlantic Oscillation (NAO) index; positive values indicate mild winters. Although milder winters are correlated with earlier migration in many birds in temperate regions, few studies have addressed how climate‐induced variation in spring arrival relates to breeding success. In northern Europe, the NAO‐index correlates with ice cover and timing of ice break‐up of the Baltic Sea. Ice cover plays an important role for breeding waterfowl, since the timing of ice break‐up constrains both spring arrival and onset of breeding. We studied the effects of the winter‐NAO‐index and timing of ice break‐up on spring migration, laying date, clutch size, female body condition at hatching and fledging success of a short‐distance migrant common eider (Somateria mollissima) population from SW Finland, the Baltic Sea, 1991–2004 (migration data 1979–2004). We also examined the correlation between the NAO‐index and the proportion of juvenile eiders in the Danish hunting bag, which reflects the breeding success on a larger spatial scale. The body condition of breeding females and proportion of juveniles in the hunting bag showed significant positive correlations with the NAO, whereas arrival dates showed positive correlations and clutch size and fledging success showed negative correlations with the timing of ice break‐up. The results suggest that climate, which also affects ice conditions, has an important effect on the fledging success of eiders. Outbreaks of duckling disease epidemics may be the primary mechanism underlying this effect. Eider females are in poorer condition after severe winters and cannot allocate as much resources to breeding, which may impair the immune defense of ducklings. Global climate warming is expected to increase the future breeding success of eiders in our study population.     

Lagged effects of North Atlantic Oscillation on spittlebug Philaenus spumarius (Homoptera) abundance and survival

The North Atlantic Oscillation (NAO) is a large‐scale pattern of climate variability that has been shown to have important ecological effects on a wide spectrum of taxa. Studies on terrestrial invertebrates are, however, lacking. We studied climate‐connected causes of changes in population sizes in island populations of the spittlebug Philaenus spumarius (L.) (Homoptera). Three populations living in meadows on small Baltic Sea islands were investigated during the years 1970–2005 in Tvärminne archipelago, southern Finland. A separate analysis was done on the effects of NAO and local climate variables on spittlebug survival in 1969–1978, for which survival data existed for two islands. We studied survival at two stages of the life cycle: growth rate from females to next year's instars (probably mostly related to overwintering egg survival), and survival from third instar stage to adult. The latter is connected to mortality caused by desiccation of plants and spittle masses. Higher winter NAO values were consistently associated with smaller population sizes on all three islands. Local climate variables entering the most parsimonious autoregressive models of population abundance were April and May mean temperature, May precipitation, an index of May humidity, and mean temperature of the coldest month of the previous winter. High winter NAO values had a clear negative effect on late instar survival in 1969–1978. Even May–June humidity and mean temperature of the coldest month were associated with late instar survival. The climate variables studied (including NAO) had no effect on the growth rate from females to next year's instars. NAO probably affected the populations primarily in late spring. Cold and snowy winters contribute to later snow melt and greater spring humidity in the meadows. We show that winter NAO has a considerable lagged effect on April and May temperature; even this second lagged effect contributes to differences in humidity. The lagged effect of the winter NAO to spring temperatures covers a large area in northern Europe and has been relatively stationary for 100 years at least in the Baltic area.     

Influences of the atmospheric patterns on unstable climate-growth associations of western Mediterranean forests

Quantifying climate-growth associations is needed to evaluate how forest productivity will respond to climate change. Year-to-year fluctuations in forest productivity and radial growth are partly explained by local climatic conditions driven by large-scale atmospheric patterns. This is illustrated by Iberian forests in the western Mediterranean Basin, which are subjected to complex climatic and atmospheric influences such as Atlantic and Mediterranean cyclogenesis. The North Atlantic Oscillation (NAO) is one of the major atmospheric circulation patterns affecting Iberian forests since positive winter NAO phases lead to dry and warm conditions. The Western Mediterranean Oscillation (WeMO) may also explain Iberian forest growth in some areas since this index captures Mediterranean cyclogenesis and WeMO negative phases are linked to warm and wet spring to summer conditions. Here, we analyze the associations between atmospheric patterns, climate and tree growth and we determine if they are changing through time. We use dendrochronology to relate radial growth of four tree species (Pyrenean oak, Sweet chestnut, Maritime pine and Scots pine) growing in western Spain to climate conditions and the NAO and WeMO indices. Winter and early spring temperatures increased since the 1950s in the area whereas the negative association between winter precipitation and the NAO strengthened since then. However, mean temperature rise was particularly evident since the 1970s. Growth was reduced by dry conditions during the growing season (spring and summer), but also by cold and dry conditions during the previous autumn and winter. This explains why the NAO January and the WeMo April indices were negative to growth of three species excluding Pyrenean oak. The early 1970s reflected an inflection point in the instability of climate-growth associations in the study area. We conclude that the winter NAO is a relevant driver of forest growth in the western Iberian Peninsula forests but additional atmospheric patterns (WeMO) also affect, albeit to a minor extent, these forests.     

The relative importance of conditions in wintering and passage areas on spring arrival dates: the case of long-distance Iberian migrants

Remote sensing data have been used in previous studies to assess the effects of winter ecological conditions in Africa on biological parameters recorded in bird populations during the following breeding season in Europe. Based on the results of these studies, we hypothesized that a high productivity of vegetation during the winter and, thus, high resource availability, should advance the arrival of long-distance migrants to the European breeding areas due to enhanced ecological conditions. To test this hypothesis, between 1982 and 2000 we examined the first arrival date to the Iberian Peninsula of five species (White Stork, Cuckoo, Common Swift, Barn Swallow and Nightingale) in relation to several explanatory variables: ecological conditions in their African wintering grounds and passage areas, as reflected by the normalized difference vegetation index (NDVI), temperature and precipitation in their passage areas and the winter North Atlantic Oscillation (NAO). Ecological conditions in the wintering areas were important for White Stork, Cuckoo and Barn Swallow phenology, while both NDVI in passage areas and NAO did not have an effect on any species. Migratory birds arrived earlier after winters with high vegetation productivity in Africa. Temperature in passage areas was important for the later species (i.e. Cuckoo, Common Swift and Nightingale), although in all cases the true relevance of this factor was scarce due to the poor explanatory capacity of the models. These species were recorded in the Iberian Peninsula earlier in the spring of those years with warmer temperatures in passage areas. The nexus between African NDVI and arrival phenology is hypothesized through increases in wintering survival rates and/or the faster acquisition of pre-migratory body condition and progression through sub-Saharan areas. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Long-term responses in arctic ungulate dynamics to changes in climatic and trophic processes

 Following predictions from climatic general circulation models, the effects of perturbations in global climate are expected to be most pronounced in the Northern Hemisphere. Elaborating on a recently developed plant–herbivore–climate model, we explore statistically how different winter climate regimes and density-dependent processes during the past century have affected population dynamics of two arctic ungulate species. Our analyses were performed on the dynamics of six muskox and six caribou populations. In muskoxen, variation in winter climate, mediated through the North Atlantic Oscillation (NAO), explained up to 24% of the variation in interannual abundance, whereas in caribou up to 16% was explained by the NAO. Muskoxen responded negatively following warm and snowy winters, whereas caribou responded negatively to dry winters. Direct and delayed density dependence was recorded in most populations and explained up to 32% and 90% of variations in abundance of muskoxen and caribou, respectively. Received: November 19, 2001 / Accepted: May 28, 2002     

Global climate change and reindeer: effects of winter weather on the autumn weight and growth of calves

Reindeer/caribou (Rangifer tarandus), which constitute a biological resource of vital importance for the physical and cultural survival of Arctic residents, and inhabit extremely seasonal environments, have received little attention in the global change debate. We investigated how body weight and growth rate of reindeer calves were affected by large-scale climatic variability [measured by the North Atlantic Oscillation (NAO) winter index] and density in one population in central Norway. Body weights of calves in summer and early winter, as well as their growth rate (summer to early winter), were significantly influenced by density and the NAO index when cohorts were in utero. Males were heavier and had higher absolute growth than females, but there was no evidence that preweaning condition of male and female calves were influenced differently by the NAO winter index. Increasing NAO index had a negative effect on calves' body weight and growth rate. Increasing density significantly reduced body weight and growth rate of calves, and accentuated the effect of the NAO winter index. Winters with a higher NAO index are thus severe for reindeer calves in this area and their effects are associated with nutritional stress experienced by the dams during pregnancy or immediately after calving. Moreover, increased density may enhance intra-specific competition and limits food available at the individual level within cohorts. We conclude that if the current pattern of global warming continues, with greater change occurring in northern latitudes and during winter as is predicted, reduced body weight of reindeer calves may be a consequence in areas where winters with a high NAO index are severe. This will likely have an effect on the livelihood of many northern indigenous peoples, both economically and culturally.     

Differential response to climatic variation of free‐floating and submerged macrophytes in ditches

1. Experimental studies have indicated in freshwater ecosystems that a shift in dominance from submerged to free‐floating macrophytes may occur with climate change because of increased water surface temperatures and eutrophication. Field evidence is, however, rare. 2. Here, we analysed long‐term (26 years) dynamics of macrophyte cover in Dutch ditches in relation to Dutch weather variables and the North Atlantic Oscillation (NAO) winter index. The latter appears to be a good proxy for Dutch weather conditions. 3. Cover of both free‐floating macrophytes and evergreen overwintering submerged macrophytes was positively related to mild winters (positive NAO winter index). On the other hand, high cover of submerged macrophytes that die back in winter coincided with cold winters (negative NAO winter index). Our results therefore suggest that the effect of weather on macrophyte species depends strongly on their overwintering strategy. 4. The positive relation of free‐floating macrophytes with the NAO winter index was significantly stronger in ditches in organic soil than in those in inorganic soil. This may be because of increased nutrient loading associated with increased decomposition of organic matter and increased run‐off to these ditches during mild wet winters. 5. Our results suggest that mild winters in a changing climate may cause submerged macrophytes with an evergreen overwintering strategy and free‐floating macrophytes to outcompete submerged macrophytes that die back in winter.     

Winter North Atlantic Oscillation, temperature and ischaemic heart disease mortality in three English counties

As cold weather is an ischaemic heart disease (IHD) risk factor, year-to-year variations of the level of IHD mortality may be partly determined by inter-annual variations in winter climate. This paper investigates whether there is any association between the level of IHD mortality for three English counties and the winter North Atlantic Oscillation (NAO), which exerts a fundamental control on the nature of the winter climate over Western Europe. Correlation and regression analysis was used to explore the nature of the association between IHD mortality and a climate index (CI) that represents the interaction between the NAO and temperature across England for the winters 1974–1975 to 1989–1999. Statistically significant inverse associations between the CI and the level of IHD mortality were found. Generally, high levels of winter IHD mortality are associated with a negative CI, which represents winters with a strong negative phase of the NAO and anomalously low temperatures across England. Moreover, the nature of the CI in the early stages of winter appears to exert a fundamental control on the general level of winter IHD mortality. Because winter climate is able to explain a good proportion of the inter-annual variability of winter mortality, long-lead forecasting of winter IHD mortality appears to be a possibility. The integration of climate-based health forecasts into decision support tools for advanced general winter emergency service and capacity planning could form the basis of an effective adaptive strategy for coping with the health effects of harsh winters.     

Food resources of the wild Barbary macaque (Macaca sylvanus) in high-altitude fir forest, Ghomaran Rif, Morocco

A small geographically isolated population of the Barbary macaque inhabits a high-altitude fir forest habitat ( Abies pinsapo ) in the Ghomaran region of the Rif mountains of northern Morocco. The climate of this region is Mediterranean, but the altitude (1600–2100 m) causes winters to be cold (as low as -8.0 C) with snow occurring from November to May (snowfall as deep as 1.5 m). The primary winter feeding adaptation is the ability to ingest high quantities of fir foliage; in spring, the macaques took a high diversity of leafy food items from all vegetation layers; in summer, the macaques foraged terrestrially for a high diversity of food items including seeds, small fruits, bulbous geophytes, and animal foods (including tadpoles from small streams); in autumn, the macaques returned to arboreal foraging, primarily feeding on oak acorns ( Quercus ilex ), fir seeds and yew fruit ( Taxus baccata ). The macaques were capable of ingesting 100 of 195 (51%) of all identified plant species in the region, although during the four-month winter, the macaques only averaged 12.5 common food items. A comparison of the study area with the prime habitat of the Barbary macaque-the high-altitude cedar forests of the Moroccan Moyen Atlas-indicates that climate and vegetation physiognomy are highly similar in both regions. Correspondingly, there is a high degree of similarity in macaque diet in both regions in terms of feeding behaviour by season, food diversity and specific feeding techniques. In the Ghomara, the winter feeding adaptation of fir foliage eating parallels that of the Barbary macaque in cedar forest (winter foraging for cedar foliage). This enables the Barbary macaque to exploit the Ghomaran fir forest habitat during the cold, snowy winters much the same as it does cedar forest habitat throughout a major portion of its geographical range.     

Does climate change explain the decline of a trans-Saharan Afro-Palaearctic migrant?

There is an urgent need to understand how climate change will impact on demographic parameters of vulnerable species. Migrants are regarded as particularly vulnerable to climate change; phenological mismatch has resulted in the local decline of one passerine, whilst variations in the survival of others have been related to African weather conditions. However, there have been few demographic studies on trans-Saharan non-passerine migrants, despite these showing stronger declines across Europe than passerines. We therefore analyse the effects of climate on the survival and productivity of common sandpipers Actitis hypoleucos, a declining non-passerine long-distant migrant using 28 years’ data from the Peak District, England. Adult survival rates were significantly negatively correlated with winter North Atlantic Oscillation (NAO), being lower when winters were warm and wet in western Europe and cool and dry in northwest Africa. Annual variation in the productivity of the population was positively correlated with June temperature, but not with an index of phenological mismatch. The 59% population decline appears largely to have been driven by reductions in adult survival, with local productivity poorly correlated with subsequent population change, suggesting a low degree of natal philopatry. Winter NAO was not significantly correlated with adult survival rates in a second, Scottish Borders population, studied for 12 years. Variation in climatic conditions alone does not therefore appear to be responsible for common sandpiper declines. Unlike some passerine migrants, there was no evidence for climate-driven reductions in productivity, although the apparent importance of immigration in determining local recruitment complicates the assessment of productivity effects. We suggest that further studies to diagnose common sandpiper declines should focus on changes in the condition of migratory stop-over or wintering locations. Where possible, these analyses should be repeated for other declining migrants.     

Spring arrival response to climate change in birds: a case study from eastern Europe

This paper analyses the dependence of the first spring arrival dates of short/medium- and long-distance migrant bird species on climate warming in eastern Europe. The timing of arrival of the selected species at the observation site correlates with the North Atlantic Oscillation (NAO) index, air temperature, atmospheric pressure, precipitation and wind characteristics. A positive correlation of fluctuations in winter and spring air temperatures with variations in the NAO index has been established in eastern Europe. Positive winter NAO index values are related to earlier spring arrival of birds in the eastern Baltic region and vice versa—arrival is late when the NAO index is negative. The impact of climate warming on the bird’s life cycle depends on local or regional climate characteristics. We tested the hypothesis that differences in climate indices between North Africa and Europe can influence the timing of spring arrival. Our results support the hypothesis that differences in first spring arrival dates between European populations occur after individuals cross the Sahara. We assume that the endogenous programme of migration control in short/medium-distance migrants synchronises with the changing environment on their wintering grounds and along their migration routes, whereas in long-distance migrants it is rather with environmental changes in the second part of their migratory route in Europe. Our results strongly indicate that the mechanism of dynamic balance in the interaction between the endogenous regulatory programme and environmental factors determines the pattern of spring arrival, as well as migration timing.     

Climatic effects on arrival and laying dates in a long-distance migrant, the Collared Flycatcher Ficedula albicollis

Long‐distance migrants may respond to climate change in breeding, wintering or staging area by changing their phenology. The geographical variation in such responses (e.g. coastal vs. continental Europe) and the relative importance of climate at different spatial scales remain unclear. Here we analysed variation in first arrival dates (FADs) and laying dates of the Collared Flycatcher Ficedula albicollis in a central European population, from 1973 to 2002. The North Atlantic Oscillation (NAO) index correlated weakly with local temperature during the laying period. Decreasing spring temperatures until 1980 were associated with a trend towards later laying. The rate of warming (0.2 °C per year) and laying advancement (0.4 days per year) since 1980 are amongst the highest values reported elsewhere. This long‐term trend in laying date was largely explained by the change in climatic factors. The negative effect of local spring temperature on laying was relatively stronger than that of NAO. The number of clutches initiated on a particular day was marginally affected by the temperature 3 days prior to laying and the response of females to daily variation in temperature did not change over years. Correspondence between the average population‐level and the individual‐level responses of laying date to climate variation suggests that the advancement of laying was due to phenotypic plasticity. Despite warmer springs and advanced laying, FADs did not change over years and were not correlated with local spring temperature. Marginal evidence suggests later departure from wintering grounds and faster migration across staging areas in warmer conditions. Advancement of arrival was probably constrained by low local temperatures in early spring just before arrival that have not changed over years. The interval between first arrival and laying has declined since 1980 (0.5 days per year), but the increasing temperature during that period may have kept the food supply approximately unchanged.     

The nature of migration in the red admiral butterfly Vanessa atalanta: evidence from the population ecology in its southern range

1. The migrant Vanessa atalanta (L.) occurs throughout Europe and North Africa. In autumn, populations emigrate from northern and central Europe to the Mediterranean region to overwinter. In the spring, the northern range is recolonised by migrants from the south. The dynamics of the species in the winter range is poorly known. 2. From 1994 to 1999, adults and immatures of V. atalanta were monitored all year round in Mediterranean habitats in north‐east Spain. 3. Data showed that the Catalonia lowlands is an area to which V. atalanta migrates to breed during the winter. Migrants arrive in October and early November and initiate a period of intensive breeding. Larval development occurs throughout the winter until a first annual generation of adults appears in early spring. 4. Most of the butterflies emerging in the spring emigrate and leave the area without breeding. The data suggest strongly that recolonisation of the northern range is by these butterflies not by wintering adults. Altitudinal migration also seems to be a common phenomenon, allowing a further summer generation of adults to occur at high elevations within the Mediterranean region. 5. The complex phenology of V. atalanta in its southern range has evolved as a strategy to track larval resources through space and time. Autumn migration coincides with the greatest availability of the main food plant, Urtica dioica L. Late spring migration occurs by the time food quality is decreasing.     

Spring arrival of birds depends on the North Atlantic Oscillation

The timing of arrival of 81 migratory species in response to the North Atlantic Oscillation (NAO) was studied at two Finnish bird observatories (1970–99). Timing was determined for the first migrants and for the peak of migration, as well as for the early, median and late phases of migration, defined as the dates when the seasonal cumulative sum of birds reached 5%, 50% and 95%, respectively. For most species, the timing of arrival correlated negatively with the NAO in all phases of migration: the correlation was significant for 79% of species studied. Thus, most species arrived in Finland early when the NAO was positive and indicative of mild and rainy winters in northern Europe. Although all phases of migration correlated negatively with the NAO, the correlations were more negative for the early than for the late phases of migration. Since the NAO did not show a significant trend during the study period, the correlations indicate that the timing of birds followed stochastic fluctuations in the NAO. This finding suggests that most Finnish migratory birds are able to adjust the timing of spring arrival in response to climatic change without time delay.