Phenological sensitivity to climate change is higher in resident than in migrant bird populations among European cavity breeders

Many organisms adjust their reproductive phenology in response to climate change, but phenological sensitivity to temperature may vary between species. For example, resident and migratory birds have vastly different annual cycles, which can cause differential temperature sensitivity at the breeding grounds, and may affect competitive dynamics. Currently, however, adjustment to climate change in resident and migratory birds have been studied separately or at relatively small geographical scales with varying time series durations and methodologies. Here, we studied differential effects of temperature on resident and migratory birds using the mean egg laying initiation dates from 10 European nest box schemes between 1991 and 2015 that had data on at least one resident tit species and at least one migratory flycatcher species. We found that both tits and flycatchers advanced laying in response to spring warming, but resident tit populations advanced more strongly in relation to temperature increases than migratory flycatchers. These different temperature responses have already led to a divergence in laying dates between tits and flycatchers of on average 0.94 days per decade over the current study period. Interestingly, this divergence was stronger at lower latitudes where the interval between tit and flycatcher phenology is smaller and winter conditions can be considered more favorable for resident birds. This could indicate that phenological adjustment to climate change by flycatchers is increasingly hampered by competition with resident species. Indeed, we found that tit laying date had an additional effect on flycatcher laying date after controlling for temperature, and this effect was strongest in areas with the shortest interval between both species groups. Combined, our results suggest that the differential effect of climate change on species groups with overlapping breeding ecology affects the phenological interval between them, potentially affecting interspecific interactions.     

Restricted dispersal in a long-distance migrant bird with patchy distribution,the great reed warbler

In patchily distributed species dispersal connects local populations into metapopulations. Reliable quantifications of dispersal are therefore crucial to understanding the population dynamics and genetic structure of such metapopulation systems. The great reed warbler (Acrocephalus arundinaceus) inhabits eutrophic lakes and has a patchy breeding distribution. In this study we investigated the dispersal pattern of the great reed warbler based on an extensive capture-recapture effort covering a large census area (22,500 km²). At two adjacent breeding sites (10 km apart) in southern Central Sweden, the "main study area", we ringed the majority of adult and nestling great reed warblers between 1992 and 1999. In 1998 and 1999, we opportunistically searched for territorial males at the majority of the Swedish breeding sites, and were able to examine about 56% of all males in the region. Analyses of recaptured males demonstrated that philopatry predominated. Sixty-nine percent of the recruiting nestlings returned to breed in the main study area (their natal area), and 92% of the resighted adults were found at the same breeding locality in both study years. Breeding dispersal was significantly more restricted than natal dispersal. Additional data on natal and breeding dispersal within the main study area in 1992-1999 suggested that females were as philopatric as males. The overall high level of philopatry, with only occasional longer dispersal distances documented, yielded a root-mean-square dispersal distance of 33 km per generation. High philopatry, short dispersal distances and similar dispersal patterns of male and female great reed warblers contrast the findings among birds in general, but conform to data of species having patchy breeding habitat and isolated populations. Restricted dispersal suggests limited gene flow even among several Swedish populations, which is in line with some previous findings of the population ecology of the great reed warbler: (1) structured mtDNA lineages among European populations; (2) small-scale population differences in song patterns; and (3) low genetic variation and occurrence of inbreeding depression in our main study population.     

Phenological advancement in arctic bird species: relative importance of snow melt and ecological factors

Previous studies have documented advancement in clutch initiation dates (CIDs) in response to climate change, most notably for temperate-breeding passerines. Despite accelerated climate change in the Arctic, few studies have examined nest phenology shifts in arctic breeding species. We investigated whether CIDs have advanced for the most abundant breeding shorebird and passerine species at a long-term monitoring site in arctic Alaska. We pooled data from three additional nearby sites to determine the explanatory power of snow melt and ecological variables (predator abundance, green-up) on changes in breeding phenology. As predicted, all species (semipalmated sandpiper, Calidris pusilla, pectoral sandpiper, Calidris melanotos, red-necked phalarope, Phalaropus lobatus, red phalarope, Phalaropus fulicarius, Lapland longspur, Calcarius lapponicus) exhibited advanced CIDs ranging from 0.40 to 0.80 days/year over 9 years. Timing of snow melt was the most important variable in explaining clutch initiation advancement (“climate/snow hypothesis”) for four of the five species, while green-up was a much less important explanatory factor. We found no evidence that high predator abundances led to earlier laying dates (“predator/re-nest hypothesis”). Our results support previous arctic studies in that climate change in the cryosphere will have a strong impact on nesting phenology although factors explaining changes in nest phenology are not necessarily uniform across the entire Arctic. Our results suggest some arctic-breeding shorebird and passerine species are altering their breeding phenology to initiate nesting earlier enabling them to, at least temporarily, avoid the negative consequences of a trophic mismatch.     

Migration speeds among eleven species of long-distance migrating passerines across Europe, the desert and eastern Africa

Based on phenology, passage and median dates gathered from large number of study sites, we measured autumn and spring migration speeds of eleven long distance migratory passerines in four different ecogeographic sectors: Europe, desert, north-eastern and eastern Africa. Results demonstrate that, during the southward autumn migration, late-departing species, such as lesser whitethroat Sylvia curruca , garden warbler S. borin , spotted flycatcher Muscicapa striata , whitethroat S. borin , and willow warbler Phylloscopus trochilus cover their migration route with a slower average migration speed across Europe than do early migrating species. During spring migration, late-departing species (marsh warbler Acrocephalus palustris , garden warbler, spotted flycatcher, red-backed shrike Lanius collurio ) across north-eastern Africa showed a higher speed than early migrating species. Our results show overall shorter migration duration estimates in spring than autumn. Sector-wise seasonal comparisons of duration indicate that migration journey in the African and desert sectors are covered in a relatively shorter time in spring than in autumn. Periods required to cover the distance between northern latitude breeding grounds and desert during both seasons were equivalent.     

Genetic differences among language families in Europe

We investigated whether 59 allele frequencies and 10 cranial variables differed among speakers of the 12 modern language families in Europe. Although this is a classical analysis of variance design, special techniques had to be developed for the analysis because of spatial autocorrelation of both biological and language data. The method examines pooled sums of squares within language families. These are compared with the same quantities obtained by randomly partitioning the available data points in Europe into internally cohesive subsets representing the same sample sizes for each language family as in the originally observed data. Our results suggest that for numerous genetic systems, population samples differ more among language families than they do within families. These findings are considered in relation to two contrasting models: a model of random spatial differentiation of gene frequencies unrelated to language and a model of aboriginal genetic differences among speakers of different language groups. Our observed findings suggest partial validity of both models.     

Body mass of six long-distance migrant passerine species along the autumn migration route

Summary We analysed body mass and moult data of six passerine species along their autumn migration route from northern Europe to North Africa and derived hypothetical models of the organisation of their migration in terms of fuel store accumulation. We analysed data of 46,541 first-year birds from 34 trapping sites, sampled in a network of collaborating European and African ringing stations. After accounting for effects of time of day and size, there were marked differences between the six species examined in the change of body mass along the migration route and in the timing of moult. Garden Warblers (Sylvia borin) and Pied Flycatchers (Ficedula hypoleuca) underwent their postjuvenile moult prior to migration and increased their average body mass along the migration route. Sedge Warblers (Acrocephalus schoenobaenus) also increased body mass towards the south, but started the migration bout without further refuelling well before the Sahara and moulted mainly in the wintering grounds. Reed Warblers (Acrocephalus scirpaceus) and Whitethroats (Sylvia communis) migrated while still moulting and did not increase average body mass towards south. They accumulated the energy needed to fly over the Sahara just before it. Spotted Flycatchers (Muscicapa striata) behaved in the same way, but contrary to Reed Warblers and Whitethroats they did not accumulate much fat stores in North Africa, which might urge them to stop and fuel up regularly in the Sahara. In the course of the season average body mass of all species increased slightly, which enabled them to migrate faster. In general, average body mass of first-year birds in northern and central Europe during the migration period was comparable to that of adults during breeding.

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Körpermassen von sechs Arten von Langstreckenziehern entlang des Herbstzugweges

Zusammenfassung Wir analysierten Körpermassen- und Mauserdaten von sechs Singvogelarten auf dem Herbstzug entlang ihres Zugweges von Nordeuropa bis Nordafrika. Die Körpermassen von 46 541 diesjährigen Vögeln, die auf 34 Beringungsstationen im Rahmen eines internationalen Projektes gesammelt wurden, sind die Grundlage dieser Untersuchung. Neben der Entwicklung der Körpermassen entlang des Zugweges untersuchten wir den Einfluss der Mauser, des Fangdatums, der Tageszeit und der Körpergröße auf die Körpermasse. Mausernde Vögel waren schwerer als nichtmausernde mit der gleichen Fettmenge. Die Körpermasse nahm im Laufe der Saison und im Verlauf des Tages zu. Große Vögel (mit einer längeren 8. Handschwinge) waren schwerer als kleine. Gartengrasmücken (Sylvia borin) und Trauerschnäpper (Ficedula hypoleuca) wurden entlang des Zugweges kontinuierlich schwerer. Beide Arten machten die Jugendteilmauser hauptsächlich noch im Brutgebiet durch. Schilfrohrsänger (Acrocephalus schoenobaenus) wurden nach Süden ebenfalls etwas schwerer, aber sie begannen mit einem Reiseabschnitt ohne weiteren Fettaufbau schon in Mitteleuropa und mauserten ihr Kleingefieder hauptsächlich erst in Afrika. Die Körpermassen von Teichrohrsängern (Acrocephalus scirpaceus) und Dorngrasmücken (Sylvia communis) blieben nach Süden weitgehend konstant. Beide Arten lagerten die nötigen Energiereserven zum Überfliegen der Sahara erst kurz davor an. Sie trennten Jugendteilmauser und Zug nicht vollständig. Ähnlich verhielten sich Grauschnäpper (Muscicapa striata), doch im Gegensatz zu den zwei vorherigen Arten, wurden sie auch im Mittelmeergebiet nicht wesentlich fetter. Sie scheinen daher regelmässig in der Wüste zu rasten und zu fressen.

     

Geographical differences in seasonal clutch size variation in multi-brooded bird species

The seasonal pattern of clutch size variation in birds varies among species. In single-brooded bird species clutch sizes decline continuously with date from an early season maximum. In resident multi-brooded species, clutch sizes first increase to a mid-season maximum and then decrease again. Limited data for multi-brooded migratory birds suggest that clutches in these species also show a continuous decline throughout the season, but it remains to be resolved whether this applies generally and whether migration adds a constraint to the system. We here report on a study of the Eastern Bluebird Sialia sialis conducted using data collected by volunteers across most of its range. In the southern part of its range, where Eastern Bluebirds are largely year-round residents, clutches initially increase, reach a mid-season maximum and gradually decline later in the season. In the northern part of the range, where the majority of Eastern Bluebirds are migratory, clutch sizes decline continuously throughout the season. To determine whether seasonal changes in the clutch size of multi-brooded species are determined by migratory behaviour of the population we compared our results with published data on the Dunnock Prunella modularis and conclude that in multi-brooded species migratory behaviour is not a sufficient condition for a continuous seasonal decline in clutch size, but it could be a necessary one. We propose two mechanisms for the pattern observed among multi-brooded migrants: (i) a time or energy cost of migration, and (ii) a more abrupt increase in seasonal resources in spring at more northerly latitudes.     

Learning and signal copying facilitate communication among bird species

Signals relevant to different sets of receivers in different contexts create a conflict for signal design. A classic example is vocal alarm signals, often used both during intraspecific and interspecific interactions. How can signals alert individuals from a variety of other species in some contexts, while also maintaining efficient communication among conspecifics? We studied heterospecific responses to avian alarm signals that drive the formation of anti-predator groups but are also used during intraspecific interactions. In three species-rich communities in the western Himalayas, alarm signals vary drastically across species. We show that, independently of differences in their calls, birds respond strongly to the alarm signals of other species with which they co-occur and much more weakly to those of species with which they do not co-occur. These results suggest that previous exposure and learning maintain heterospecific responses in the face of widespread signal divergence. At an area where only two species regularly interact, one species'' calls incorporate the call of the other. We demonstrate experimentally that signal copying allows strong responses even without previous exposure and suggest that such hybrid calls may be especially favoured when pairwise interactions between species are strong.     

Patterns of endemic extinctions among island bird species

The relationship between island biogeography and the vulnerability of island biota to extinction as a result of human activities was examined. In particular, this study analyzed whether island area, maximum elevation of an island, isolation from the nearest continental landmass, or date of human colonization had statistically significant relationships with the proportion of endemic island bird species that have become endangered or extinct. The study examined islands or island groups with endemic bird species, and which have never been connected to a continental landmass. Both modern and fossil bird species were incorporated into the analysis. Islands that were colonized by humans earliest had the lowest proportion of modern species alone, and modern and fossil species combined, that have gone extinct. However, date of human arrival was not correlated with the proportion of modern species that are endangered. Maximum elevation of an island was negatively correlated with the proportion of modern species that are extinct, and was positively correlated with the proportion of modern species that are endangered. Area was negatively correlated with the proportion of modern species that are endangered. Isolation of islands was not significantly correlated with the proportion of modern species extinct or endangered, but was positively correlated with the proportion of modern and fossil species combined that have gone extinct. These results indicate that the initial spasm of island bird extinctions due to human contact may have, in part, passed. They also indicate that bird species on islands colonized earliest by humans may have had more time to adapt to the presence of man and his commensal species, resulting in reduced extinction rates.     

Climate change and rates of vagrancy of Siberian bird species to Europe

To assess whether increasing numbers of Siberian vagrants observed in Europe in recent autumns can be linked to climate change, we predicted changes in the climatic suitability of the breeding ranges of 46 Siberian bird species known to show vagrancy to Europe and compared these predictions with observed changes in recorded rates of autumn vagrancy across eight European countries during the last three decades. There was a positive correlation between predicted increase in breeding range and vagrancy rates. A positive impact of climate change on range and population size could promote vagrancy, while the increasing use of such alternative migration flyways could provide adaptive advantages in a changing environment.     

Arrival fat and reproductive performance in a long-distance passerine migrant

Long-distance passerine migrants deposit substantial fat stores to fuel their migratory journey. Many of those migratory birds arrive at their northerly breeding grounds with larger fat stores than were necessary to reach their breeding area. Both male and female American Redstarts ( Setophaga ruticilla) arrived to breed in Michigan's Upper Peninsula with fat, and females arrived with more fat than males in 2 out of 3 years. We test the hypothesis that migrants arriving at the breeding grounds with more body fat have higher reproductive success than birds arriving with little or no fat. Females, and to a lesser extent males, that arrive with fat experience gains in reproductive performance as evidenced by increased clutch size, egg volume, and nestling mass. The results have implications for understanding how events occurring during one phase of the annual cycle influence survival and/or reproductive performance in subsequent phases.     

Corticosterone,food intake and refueling in a long-distance migrant

Elevated baseline corticosterone levels function to mobilize energy in predictable life-history stages, such as bird migration. At the same time, baseline corticosterone has a permissive effect on the accumulation of fat stores (fueling) needed for migratory flight. Most migrants alternate flight bouts with stopovers, during which they replenish the fuel used during the preceding flight (refueling). The role of corticosterone in refueling is currently unclear. In a fasting–re-feeding experiment on northern wheatears (Oenanthe oenanthe) in autumn we found that baseline total and free corticosterone levels were negatively related with both food intake and the rate of fuel deposition after fasting. This confirms our earlier findings in wild conspecifics in spring and indicates that corticosterone does not stimulate stopover refueling. Whether the negative relationship between baseline corticosterone level and fuel deposition rate is causal is questionable, because within-individual comparison of corticosterone metabolite levels in droppings did not reveal differences between refueling and control periods. In other words, corticosterone does not appear to be down-regulated during refueling, which would be expected if it directly hampers refueling. We discuss possible correlates of corticosterone level that may explain the negative association between corticosterone and stopover refueling. Additionally, we found that fasting decreases total corticosterone level, which contrasts with previous studies. We propose that the difference is due to the other studies being conducted outside of the migration life-history stage, and provide a possible explanation for the decrease in corticosterone during fasting in migrating birds.     

Climate change and timing of spring migration in the long-distance migrant Ficedula hypoleuca in central Europe: the role of spatially different temperature changes along migration routes

Coinciding with increasing spring temperatures in Europe, many migrants have advanced their arrival or passage times over the last decades. However, some species, namely long-distance migrants, could be constrained in their arrival dates due to their largely inherited migratory behaviour and thus a likely inflexibility in their response to exogenous factors. To examine this hypothesis for pied flycatchers (Ficedula hypoleuca), we tested the effects of the temperature regimes along their migration routes north of the Sahara on their arrival times in central Europe. To do so, we developed a site-independent large-scale approach based on temperature data available on the Internet. Temperature regimes along the migration routes of pied flycatchers within Europe convincingly correlate with their first arrival times. It can be concluded that the progression of spring migration in this species is strongly influenced by temperature en route. Because of the recent inconsistent climatic changes in various parts of Europe, we hypothesize that individuals migrating along different routes will be unequally affected by further climatic changes.     

Flight speeds among bird species: allometric and phylogenetic effects

Flight speed is expected to increase with mass and wing loading among flying animals and aircraft for fundamental aerodynamic reasons. Assuming geometrical and dynamical similarity, cruising flight speed is predicted to vary as (body mass)^1/6 and (wing loading)^1/2 among bird species. To test these scaling rules and the general importance of mass and wing loading for bird flight speeds, we used tracking radar to measure flapping flight speeds of individuals or flocks of migrating birds visually identified to species as well as their altitude and winds at the altitudes where the birds were flying. Equivalent airspeeds (airspeeds corrected to sea level air density, U_e) of 138 species, ranging 0.01–10 kg in mass, were analysed in relation to biometry and phylogeny. Scaling exponents in relation to mass and wing loading were significantly smaller than predicted (about 0.12 and 0.32, respectively, with similar results for analyses based on species and independent phylogenetic contrasts). These low scaling exponents may be the result of evolutionary restrictions on bird flight-speed range, counteracting too slow flight speeds among species with low wing loading and too fast speeds among species with high wing loading. This compression of speed range is partly attained through geometric differences, with aspect ratio showing a positive relationship with body mass and wing loading, but additional factors are required to fully explain the small scaling exponent of U_e in relation to wing loading. Furthermore, mass and wing loading accounted for only a limited proportion of the variation in U_e. Phylogeny was a powerful factor, in combination with wing loading, to account for the variation in U_e. These results demonstrate that functional flight adaptations and constraints associated with different evolutionary lineages have an important influence on cruising flapping flight speed that goes beyond the general aerodynamic scaling effects of mass and wing loading.     

Genetic and plant host differences of Fomes fomentarius in selected parts of Southern Europe

Abstract

Our study included a total of 36 Fomes fomentarius specimens obtained from Portugal, Spain, Italy, Greece and the Czech Republic. After multiple sequence alignment analysis we observed that samples collected in Southern Europe belong to the lineage B with the exception of one specimen from northern Greece.     

Early-life behaviour predicts first-year survival in a long-distance avian migrant

Early-life conditions have critical, long-lasting effects on the fate of individuals, yet early-life activity has rarely been linked to subsequent survival of animals in the wild. Using high-resolution GPS and body-acceleration data of 93 juvenile white storks (Ciconia ciconia), we examined the links between behaviour during both pre-fledging and post-fledging (fledging-to-migration) periods and subsequent first-year survival. Juvenile daily activity (based on overall dynamic body acceleration) showed repeatable between-individual variation, the juveniles'' pre- and post-fledging activity levels were correlated and both were positively associated with subsequent survival. Daily activity increased gradually throughout the post-fledging period, and the relationship between post-fledging activity and survival was stronger in individuals who increased their daily activity level faster (an interaction effect). We suggest that high activity profiles signified individuals with increased pre-migratory experience, higher individual quality and perhaps more proactive personality, which could underlie their superior survival rates. The duration of individuals’ fledging-to-migration periods had a hump-shaped relationship with survival: higher survival was associated with intermediate rather than short or long durations. Short durations reflect lower pre-migratory experience, whereas very long ones were associated with slower increases in daily activity level which possibly reflects slow behavioural development. In accordance with previous studies, heavier nestlings and those that hatched and migrated earlier had increased survival. Using extensive tracking data, our study exposed new links between early-life attributes and survival, suggesting that early activity profiles in migrating birds can explain variation in first-year survival.     

Migratory connectivity and population-specific migration routes in a long-distance migratory bird

Knowledge about migratory connectivity, the degree to which individuals from the same breeding site migrate to the same wintering site, is essential to understand processes affecting populations of migrants throughout the annual cycle. Here, we study the migration system of a long-distance migratory bird, the Montagu''s harrier Circus pygargus, by tracking individuals from different breeding populations throughout northern Europe. We identified three main migration routes towards wintering areas in sub-Saharan Africa. Wintering areas and migration routes of different breeding populations overlapped, a pattern best described by ‘weak (diffuse) connectivity’. Migratory performance, i.e. timing, duration, distance and speed of migration, was surprisingly similar for the three routes despite differences in habitat characteristics. This study provides, to our knowledge, a first comprehensive overview of the migration system of a Palaearctic-African long-distance migrant. We emphasize the importance of spatial scale (e.g. distances between breeding populations) in defining patterns of connectivity and suggest that knowledge about fundamental aspects determining distribution patterns, such as the among-individual variation in mean migration directions, is required to ultimately understand migratory connectivity. Furthermore, we stress that for conservation purposes it is pivotal to consider wintering areas as well as migration routes and in particular stopover sites.     

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.     

萼花臂尾轮虫种复合体内三个姐妹种的形态差异

应用单因子方差分析和判别分析方法,分别比较了采自芜湖市3个自然水体的萼花臂尾轮虫种复合体内3个姐妹种(分别以克隆HE1和HE5、克隆HE3和SE1、克隆LE9为代表)的成体和幼体的形态差异。结果表明,克隆HE3成体的各形态参数均最小,而克隆HE1和LE9成体的各形态参数间无显著的差异;克隆HE1和HE3幼体的前中棘刺间距离和后侧棘刺长度、克隆LE9和HE1幼体的前侧棘刺间距离以及前中棘刺和侧棘刺间距离均无显著性差异,轮虫幼体的其它形态参数均以克隆LE9最长,克隆HE1次之,克隆HE3最短。对克隆HE1、HE3和LE9幼体有明显差异的形态参数所作的分布图显示,它们的分布区有很高程度的重叠。以判别分析建立的判别函数对3姐妹种成体的综合判别率中形态分析Ⅰ为47.8%,形态分析Ⅱ为55.6%;对3姐妹种幼体的综合判别率中形态分析Ⅰ和Ⅱ均为71.1%;两种形态分析对克隆HE1判别准确率均较低,分别为0-76.7%和0-74.2%;依据形态特征无法准确甄别萼花臂尾轮虫姐妹种。