Amphibian breeding phenology trends under climate change: predicting the past to forecast the future

Global climate warming is predicted to hasten the onset of spring breeding by anuran amphibians in seasonal environments. Previous data had indicated that the breeding phenology of a population of Fowler's Toads (Anaxyrus fowleri) at their northern range limit had been progressively later in spring, contrary to generally observed trends in other species. Although these animals are known to respond to environmental temperature and the lunar cycle to commence breeding, the timing of breeding should also be influenced by the onset of overwintering animals’ prior upward movement through the soil column from beneath the frost line as winter becomes spring. I used recorded weather data to identify four factors of temperature, rainfall and snowfall in late winter and early spring that correlated with the toads’ eventual date of emergence aboveground. Estimated dates of spring emergence of the toads calculated using a predictive model based on these factors, as well as the illumination of the moon, were highly correlated with observed dates of emergence over 24 consecutive years. Using the model to estimate of past dates of spring breeding (i.e. retrodiction) indicated that even three decades of data were insufficient to discern any appreciable phenological trend in these toads. However, by employing weather data dating back to 1876, I detected a significant trend over 140 years towards earlier spring emergence by the toads by less than half a day/decade, while, over the same period of time, average annual air temperature and annual precipitation had both increased. Changes in the springtime breeding phenology for late‐breeding species, such as Fowler's Toads, therefore may conform to expectations of earlier breeding under global warming. Improved understanding of the environmental cues that bring organisms out of winter dormancy will enable better interpretation of long‐term phenological trends.     

Extended season for northern butterflies

Butterflies are like all insects in that they are temperature sensitive and a changing climate with higher temperatures might effect their phenology. Several studies have found support for earlier flight dates among the investigated species. A comparative study with data from a citizen science project, including 66 species of butterflies in Sweden, was undertaken, and the result confirms that most butterfly species now fly earlier during the season. This is especially evident for butterflies overwintering as adults or as pupae. However, the advancement in phenology is correlated with flight date, and some late season species show no advancement or have even postponed their flight dates and are now flying later in the season. The results also showed that latitude had a strong effect on the adult flight date, and most of the investigated species showed significantly later flights towards the north. Only some late flying species showed an opposite trend, flying earlier in the north. A majority of the investigated species in this study showed a general response to temperature and advanced their flight dates with warmer temperatures (on average they advanced their flight dates by 3.8 days/°C), although not all species showed this response. In essence, a climate with earlier springs and longer growing seasons seems not to change the appearance patterns in a one-way direction. We now see butterflies on the wings both earlier and later in the season and some consequences of these patterns are discussed. So far, studies have concentrated mostly on early season butterfly–plant interactions but also late season studies are needed for a better understanding of long-term population consequences.     

Long-term trends in first arrival and first egg laying dates of some migrant and resident bird species in northern Italy

Climate change is affecting the phenology of seasonal events in Europe and the Northern Hemisphere, as shown by several studies of birds’ timing of migration and reproduction. Here, we analyse the long-term (1982–2006) trends of first arrival dates of four long-distance migratory birds [swift (Apus apus), nightingale (Luscinia megarhynchos), barn swallow (Hirundo rustica), and house martin (Delichon urbicum)] and first egg laying dates of two migrant (swift, barn swallow) and two resident species [starling (Sturnus vulgaris), Italian sparrow (Passer italiae)] at a study site in northern Italy. We also addressed the effects of local weather (temperature and precipitation) and a climate index (the North Atlantic Oscillation, NAO) on the interannual variability of phenological events. We found that the swift and the barn swallow significantly advanced both arrival and laying dates, whereas all other species did not show any significant temporal trend in either arrival or laying date. The earlier arrival of swifts was explained by increasing local temperatures in April, whereas this was not the case for arrival dates of swallows and first egg laying dates of both species. In addition, arrival dates of house martins were earlier following high NAO winters, while nightingale arrival was earlier when local spring rainfall was greater. Finally, Italian sparrow onset of reproduction was anticipated by greater spring rainfall, but delayed by high spring NAO anomalies, and swift’s onset of reproduction was anticipated by abundant rainfall prior to reproduction. There were no significant temporal trends in the interval between onset of laying and arrival in either the swift or the barn swallow. Our findings therefore indicate that birds may show idiosyncratic responses to climate variability at different spatial scales, though some species may be adjusting their calendar to rapidly changing climatic conditions.     

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.     

The influence of climate on the timing and rate of spring bird migration

Ecological processes are changing in response to climatic warming. Birds, in particular, have been documented to arrive and breed earlier in spring and this has been attributed to elevated spring temperatures. It is not clear, however, how long-distance migratory birds that overwinter thousands of kilometers to the south in the tropics cue into changes in temperature or plant phenology on northern breeding areas. We explored the relationships between the timing and rate of spring migration of long-distance migratory birds, and variables such as temperature, the North Atlantic Oscillation (NAO) and plant phenology, using mist net capture data from three ringing stations in North America over a 40-year period. Mean April/May temperatures in eastern North America varied over a 5°C range, but with no significant trend during this period. Similarly, we found few significant trends toward earlier median capture dates of birds. Median capture dates were not related to the NAO, but were inversely correlated to spring temperatures for almost all species. For every 1°C increase in spring temperature, median capture dates of migratory birds averaged, across species, one day earlier. Lilac (Syringa vulgaris) budburst, however, averaged 3 days earlier for every 1°C increase in spring temperature, suggesting that the impact of temperature on plant phenology is three times greater than on bird phenology. To address whether migratory birds adjust their rate of northward migration to changes in temperature, we compared median capture dates for 15 species between a ringing station on the Gulf Coast of Louisiana in the southern USA with two stations approximately 2,500 km to the north. The interval between median capture dates in Louisiana and at the other two ringing stations was inversely correlated with temperature, with an average interval of 22 days, that decreased by 0.8 days per 1°C increase in temperature. Our results suggest that, although the onset of migration may be determined endogenously, the timing of migration is flexible and can be adjusted in response to variation in weather and/or phenology along migration routes.     

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.     

Temporal trends in phenology of the honey bee Apis mellifera (L.) and the small white Pieris rapae (L.) in the Iberian Peninsula (1952–2004)

Abstract.  1. The first adult appearance of two insect species, the honey bee Apis mellifera (L.) and the small white Pieris rapae (L.), was examined between 1952 and 2004 in Spain.
2. After factoring out the variability resulting from the broad geographical and topographical range of the 798 sampling localities, multiple regression models were used to detect temporal trends in phenology.
3. The best models were repeated, including spring temperature as the explanatory variable to examine the effects of climate on appearance phenology.
4. Both species showed similar temporal trends, delaying their appearance phenology until the mid-1970s and advancing it since that time.
5. The appearance times for both species were negatively related to mean temperature between February and April, with both species appearing earlier in years with warmer springs.
6. The strong dependence of appearance dates on temperature indicates that climatic fluctuations are primarily responsible for the inter-annual variability in spring appearance phenology of both species, and consequently account for the observed long-term trends.
7. This study demonstrates that insect phenology is an accurate and sensitive bioindicator of climate change.     

Local and regional climate variables driving spring phenology of tortricid pests: a 36 year study

1. Insect phenology is driven by local climate variables, most notably temperature. Increased warming has been linked to advancements in critical phenophases such as the spring flight of reproductive adults in the mid‐Atlantic region of the U.S.A. 2. Local climate is governed by the fluctuations of large‐scale climate oscillations. In the northern hemisphere, both the North Atlantic Oscillation (NAO) and the Arctic Oscillation (AO) control the local autumn and winter severity. Low NAO and AO indices are associated with colder autumns and winters, which can delay spring phenology. 3. In this study, 36 years of data from experimental fruit orchards in Biglerville, Pennsylvania, were used to run partial least‐squares regressions in order to determine the climate variables related to the spring phenology of five tortricid pest species. 4. The phenology of the tortricid pests did not advance, even though there was evidence of warming at the research site. 5. Spring temperatures were found to be the most significant climate variables in determining the timing of the spring flights. However, autumn–winter temperatures were also important. 6. For the NAO and the AO, it was found that these oscillations affected the tortricid moths by influencing autumn–winter conditions. The oscillations of the NAO and AO can obscure long‐term changes in phenology. 7. These findings suggest that the inclusion of large‐scale climate oscillations can provide important insights into how climate conditions can influence insect phenology, and presents an opportunity for improving the ability to forecast spring emergence.     

The phenology of spring bird migration to north Norway

Capsule There was no evidence of a long-term trend in arrival dates of spring migrants over the last 20 years in north Norway.

Aims To investigate the effect of climate on the timing of spring arrival of many species at their northern limit of breeding distribution and to seek evidence of any long-term trend.

Methods Observations of spring arrivals of 71 species into Troms, north Norway (69–70°N) between 1970 and 2000 by members of the local branch of the Norwegian Ornithological Society were analysed and related to available weather parameters.

Results The first migrants arrive in mid-March when air temperatures are still below 0°C and c. 1 m of snow is on the ground, with the main influx in late April and throughout May. By then much of the snow has melted and temperatures are c. 5°C. There was no evidence of a long-term trend in the median dates of arrival across 31 species in Troms between 1980 and 2000. However among 14 species for which detailed data exist in a limited area around Tromsø, the Golden Plover Pluvialis apricaria showed a significant trend towards an earlier arrival and the Willow Warbler Phylloscopus trochilus showed the opposite. Spring migration tended to be advanced during warm springs, and seven of the above subsets of 14 species showed significant negative correlations between their arrival dates and temperature. One showed a positive correlation.

Conclusion The data collected to date are a good baseline for future studies of the effects of climate change on migration phenology.     

Field evidence for earlier leaf-out dates in alpine grassland on the eastern Tibetan Plateau from 1990 to 2006

Worldwide, many plant species are experiencing an earlier onset of spring phenophases due to climate warming. Rapid recent temperature increases on the Tibetan Plateau (TP) have triggered changes in the spring phenology of the local vegetation. However, remote sensing studies of the land surface phenology have reached conflicting interpretations about green-up patterns observed on the TP since the mid-1990s. We investigated this issue using field phenological observations from 1990 to 2006, for 11 dominant plants on the TP at the levels of species, families (Gramineae—grasses and Cyperaceae—sedges) and vegetation communities (alpine meadow and alpine steppe). We found a significant trend of earlier leaf-out dates for one species (Koeleria cristata). The leaf-out dates of both Gramineae and Cyperaceae had advanced (the latter significantly, starting an average of 9 days later per year than the former), but the correlation between them was significant. The leaf-out dates of both vegetation communities also advanced, but the pattern was only significant in the alpine meadow. This study provides the first field evidence of advancement in spring leaf phenology on the TP and suggests that the phenology of the alpine steppe can differ from that of the alpine meadow. These findings will be useful for understanding ecosystem responses to climate change and for grassland management on the TP.     

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.     

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.     

Quo vadis amphibia? Global warming and breeding phenology in frogs,toads and salamanders

As the earth is getting warmer, many animals and plants have shifted their timing of breeding towards earlier dates. However, there is substantial variation between populations in phenological shifts that typically goes unexplained. Identification of the different location and species characteristics that drive such variable responses to global warming is crucial if we are to make predictions for how projected climate change scenarios will play out on local and global scales. Here we conducted a phylogenetically controlled meta‐analysis of breeding phenology across frogs, toads and salamanders to examine the extent of variation in amphibian breeding phenology in response to global climate change. We show that there is strong geographic variation in response to global climate change, with species at higher latitudes exhibiting a more pronounced shift to earlier breeding than those at lower latitudes. Our analyses suggest that this latitude effect is a result of both the increased temperature (but not precipitation) at higher latitudes as well as a greater responsiveness by northern populations of amphibians to this change in temperature. We suggest that these effects should reinforce any direct effect of increasing warming at higher latitudes on breeding phenology. In contrast, we found very little contribution from other location factors or species traits. There was no evidence for a phylogenetic signal on advancing breeding phenology or responsiveness to temperature, suggesting that the amphibians that have been studied to date respond similarly to global warming.     

Temperature and snowfall trigger alpine vegetation green‐up on the world's roof

Rapid temperature increase and its impacts on alpine ecosystems in the Qinghai–Tibetan Plateau, the world's highest and largest plateau, are a matter of global concern. Satellite observations have revealed distinctly different trend changes and contradicting temperature responses of vegetation green‐up dates, leading to broad debate about the Plateau's spring phenology and its climatic attribution. Large uncertainties in remote‐sensing estimates of phenology significantly limit efforts to predict the impacts of climate change on vegetation growth and carbon balance in the Qinghai–Tibetan Plateau, which are further exacerbated by a lack of detailed ground observation calibration. Here, we revealed the spatiotemporal variations and climate drivers of ground‐based herbaceous plant green‐up dates using 72 green‐up datasets for 22 herbaceous plant species at 23 phenological stations, and corresponding daily mean air temperature and daily precipitation data from 19 climate stations across eastern and southern parts of the Qinghai–Tibetan Plateau from 1981 to 2011. Results show that neither the continuously advancing trend from 1982 to 2011, nor a turning point in the mid to late 1990s as reported by remote‐sensing studies can be verified by most of the green‐up time series, and no robust evidence for a warmer winter‐induced later green‐up dates can be detected. Thus, chilling requirements may not be an important driver influencing green‐up responses to spring warming. Moreover, temperature‐only control of green‐up dates appears mainly at stations with relatively scarce preseason snowfall and lower elevation, while coupled temperature and precipitation controls of green‐up dates occur mostly at stations with relatively abundant preseason snowfall and higher elevation. The diversified interactions between snowfall and temperature during late winter to early spring likely determine the spatiotemporal variations of green‐up dates. Therefore, prediction of vegetation growth and carbon balance responses to global climate change on the world's roof should integrate both temperature and snowfall variations.     

Dramatic changes in the return date of Guillemots Uria aalge to colonies in Shetland, 1962–2005

Capsule Changes in return date coincided with marked changes in population size that probably resulted in fluctuating competition for nest-sites.

Aims To document the changes in return dates over a 44-year period and to identify the factors associated with these changes.

Methods We compared changes in return date at Shetland colonies with those for the Isle of May, southeast Scotland, and with the available information on population size, the abundance of some fish species eaten by Common Guillemots and large-scale changes in the oceanography and climate of the eastern Atlantic as reflected by the winter index of the North Atlantic Oscillation (NAO).

Results Common Guillemots normally return to colonies in Shetland in late winter. However, during the 1960s return dates became gradually earlier with birds present from early October. Autumn return remained the norm for about ten years after which return dates gradually reverted back to late winter. In contrast, Common Guillemots on the Isle of May, 400 km south of Shetland, showed no marked shift, returning in October each year. There was a strong negative correlation between date of return of Shetland birds and population size, whereas on the Isle of May birds came back earlier when there was a large positive winter NAO index. There was no convincing evidence that changes in wintering areas or fish abundance influenced when birds returned to the colonies, although the fish data may not have been collected on the correct spatial scale.

Conclusion Competition for high quality nest-sites is the most likely reason for Common Guillemots returning to the colonies during the autumn and winter.     

Impact of climate change on plant phenology in Mediterranean ecosystems

Plant phenology is strongly controlled by climate and has consequently become one of the most reliable bioindicators of ongoing climate change. We used a dataset of more than 200 000 records for six phenological events of 29 perennial plant species monitored from 1943 to 2003 for a comprehensive assessment of plant phenological responses to climate change in the Mediterranean region. Temperature, precipitation and North Atlantic Oscillation (NAO) were studied together during a complete annual cycle before phenological events to determine their relative importance and potential seasonal carry‐over effects. Warm and dry springs under a positive phase of NAO advance flowering, leaf unfolding and fruiting dates and lengthen the growing season. Spatial variability of dates (range among sites) was also reduced during warm and dry years, especially for spring events. Climate during previous weeks to phenophases occurrence had the greatest impact on plants, although all events were also affected by climate conditions several months before. Immediate along with delayed climate effects suggest dual triggers in plant phenology. Climatic models accounted for more than 80% of variability in flowering and leaf unfolding dates, and in length of the growing season, but for lower proportions in fruiting and leaf falling. Most part of year‐to‐year changes in dates was accounted for temperature, while precipitation and NAO accounted for <10% of dates' variability. In the case of flowering, insect‐pollinated species were better modelled by climate than wind‐pollinated species. Differences in temporal responses of plant phenology to recent climate change are due to differences in the sensitivity to climate among events and species. Spring events are changing more than autumn events as they are more sensitive to climate and are also undergoing the greatest alterations of climate relative to other seasons. In conclusion, climate change has shifted plant phenology in the Mediterranean region.     

Onset of spring starting earlier across the Northern Hemisphere

Recent warming of Northern Hemisphere (NH) land is well documented and typically greater in winter/spring than other seasons. Physical environment responses to warming have been reported, but not details of large‐area temperate growing season impacts, or consequences for ecosystems and agriculture. To date, hemispheric‐scale measurements of biospheric changes have been confined to remote sensing. However, these studies did not provide detailed data needed for many investigations. Here, we show that a suite of modeled and derived measures (produced from daily maximum–minimum temperatures) linking plant development (phenology) with its basic climatic drivers provide a reliable and spatially extensive method for monitoring general impacts of global warming on the start of the growing season. Results are consistent with prior smaller area studies, confirming a nearly universal quicker onset of early spring warmth (spring indices (SI) first leaf date, ?1.2 days decade^?1), late spring warmth (SI first bloom date, ?1.0 days decade^?1; last spring day below 5°C, ?1.4 days decade^?1), and last spring freeze date (?1.5 days decade^?1) across most temperate NH land regions over the 1955–2002 period. However, dynamics differ among major continental areas with North American first leaf and last freeze date changes displaying a complex spatial relationship. Europe presents a spatial pattern of change, with western continental areas showing last freeze dates getting earlier faster, some central areas having last freeze and first leaf dates progressing at about the same pace, while in portions of Northern and Eastern Europe first leaf dates are getting earlier faster than last freeze dates. Across East Asia last freeze dates are getting earlier faster than first leaf dates.     

Changes in spring arrival of Nearctic-Neotropical migrants attributed to multiscalar climate

Climate-related changes associated with the California marine ecosystem have been documented; however, there are no studies assessing changes in terrestrial vertebrate phenology on the Pacific coast of western North America. We analyze the spring phenology of 21 Nearctic-Neotropical migratory songbird species in central and northern CA. Using observational and banding data at multiple sites, we evaluate evidence for a change in arrival timing being linked to either nonclimatic or multiscalar climatic explanations. Using correlation analysis, of the 13 species with a significant ( P <0.10) change in arrival, the arrival timing of 10 species (77%) is associated with both temperature and a large-scale climate oscillation index (El Niño Southern Oscillation, ENSO; North Atlantic Oscillation, NAO; and/or Pacific Decadal Oscillation, PDO) at least at one location. Eight of the 13 species (62%) are advancing their migratory timing. All species for which spring arrival is associated with climate at multiple locations are exhibiting changes ( n =5) and all species lacking evidence for association between migration phenology and climate ( n =3) exhibit no change. Migrants tend to arrive earlier in association with warmer temperatures, positive NAO indices, and stronger ENSO indices. Twelve species negatively correlate ( P ≤0.05) with local or regional temperature at least at one location; five species negatively correlate with ENSO. Eleven species' arrival is correlated ( P ≤0.05) with NAO; 10 are negatively associated. After an exhaustive literature search, this is apparently the first documentation of an association between NAO and migratory phenology in western North America.     

Long-term changes in breeding phenology at two seabird colonies in the western North Sea

There is compelling evidence that the breeding phenology of many species has changed substantially in recent decades. However, taxonomic and spatial variation in the direction and rate of change is still not well understood. We explored these issues by analysing a dataset containing information on first egg dates of 10 species of seabird at two major breeding colonies (86 km apart) in the western North Sea over a period of 35 years. Within a species, timing of breeding was positively correlated between the two colonies, suggesting that factors affecting the phenology of these species operated at a regional rather than a colony scale. Comparison of time trends among the species revealed contrasting patterns, with some showing no systematic change, others becoming earlier and others later. The clearest species groupings appeared to be among the terns with arrival and/or first egg dates becoming earlier in Arctic Terns Sterna paradisaea , Common Terns Sterna hirundo and Sandwich Terns Sterna sandvicensis , and among the auks (Common Guillemot Uria aalge , Razorbill Alca torda and Atlantic Puffin Fratercula arctica ) and Black-legged Kittiwake Rissa tridactyla where the trend was in the opposite direction towards later breeding. This general trend towards later breeding in the latter group of species contrasts with correlational evidence from many other organisms indicating that breeding phenology is advancing in response to climate change.     

Challenging a 15‐year‐old claim: The North Atlantic Oscillation index as a predictor of spring migration phenology of birds

Many migrant bird species that breed in the Northern Hemisphere show advancement in spring arrival dates. The North Atlantic Oscillation (NAO) index is one of the climatic variables that have been most often investigated and shown to be correlated with these changes in spring arrival. Although the NAO is often claimed to be a good predictor or even to have a marked effect on interannual changes in spring migration phenology of Northern Hemisphere breeding birds, the results on relations between spring migration phenology and NAO show a large variety, ranging from no, over weak, to a strong association. Several factors, such as geographic location, migration phase, and the NAO index time window, have been suggested to partly explain these observed differences in association. A combination of a literature meta‐analysis, and a meta‐analysis and sliding time window analysis of a dataset of 23 short‐ and long‐distance migrants from the constant‐effort trapping garden at Helgoland, Germany, however, paints a completely different picture. We found a statistically significant overall effect size of the NAO on spring migration phenology (coefficient = ?0.14, SE = 0.054), but this on average only explains 0%–6% of the variance in spring migration phenology across all species. As such, the value and biological meaning of the NAO as a general predictor or explanatory variable for climate change effects on migration phenology of birds, seems highly questionable. We found little to no definite support for previously suggested factors, such as geographic location, migration phenology phase, or the NAO time window, to explain the heterogeneity in correlation differences. We, however, did find compelling evidence that the lack of accounting for trends in both time series has led to strongly inflated (spurious) correlations in many studies (coefficient = ?0.13, SE = 0.019).