The evolution of bird migration—a synthesis

We approach the problem of the evolution of bird migration by asking whether migration evolves towards new breeding areas or towards survival areas in the non-breeding season. Thus, we avoid the ambiguity of the usually discussed “southern-home-theory” or “northern-home-theory”. We argue that migration evolved in birds that spread to seasonal habitats through gradual dispersal to enhance survival during the non-breeding season; this in contrast to the alternative idea suggesting that migration evolved towards new breeding areas to increase reproductive success. Our synthesis is based on the threshold model explaining how migratory traits can change rapidly through microevolutionary processes. Our model brings former theories together and explains how bird migration, with the appropriate direction and time program, evolves through selection after genetically non-directed events such as dispersal and colonization. The model does not need the former untested assumptions such as competition as a reason for migration and for the disappearance of sedentary populations or higher reproductive success in temperate breeding areas. Our theory offers answers to questions such as how birds with a southern origin may gradually reach northern latitudes, why migration routes may follow historical expansion routes and why birds leave an area for the non-breeding season and move back instead of breeding on their wintering grounds. The theory proposes gradual change through selection and not sudden changes such as long distance dispersal or mutations and can be applied to migration at all latitudes and in all directions. The scenario provides a reasonable concept to understand most of the existing migratory phenomena on the basis of the ecology and genetics of migratory behaviour.     

Non-migratory breeding by isolated green sea turtles (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) in the Indian Ocean: biological and conservation implications

Green sea turtles (Chelonia mydas) are renowned for their long-distance migrations but have less fame for short-distance migrations or non-migratory behavior. We present satellite telemetric evidence from Cocos (Keeling) Islands, Indian Ocean for the first predominantly non-migratory green sea turtle (C. mydas) population. The mean migration distance from the nesting beach to the foraging grounds was 35.5 km with a maximum mean transit time of 3.4 days. The behavior of this population has major implications for our general understanding of green turtle behavior and their life cycle and for conservation. Firstly, these results indicate a level of juvenile or adult non-breeding homing behavior from the open ocean to foraging grounds adjacent to their natal nesting beach. Secondly, a non-migratory breeding phase reduces the consumption of reproductive energy utilized, potentially resulting in higher fecundity for this population. Thirdly, the close proximity of the nesting and foraging habitats allows for uniformity in management and conservation strategies rarely possible for wide-ranging green turtle populations.     

Specific floater home ranges and prospective behaviour in the European starling,<Emphasis Type="Italic"> Sturnus vulgaris</Emphasis>

In many bird species, floaters are present on the breeding grounds in one or more years before they breed. There is increasing evidence that they have specific home ranges in which they search for information about current and future breeding opportunities. We investigated the role of prospecting in a migratory European starling (Sturnus vulgaris) population. Radio-tracking showed that male starling floaters use specific home range areas during the breeding period. Nest-box observations demonstrated that non-parental nest intrusion is common in the starling and that it is significantly more frequent during the nestling than during the incubation period. In addition, small groups of nest boxes were more likely to be occupied by starlings if they had been put up during the preceding breeding season. The results suggest that floaters try to acquire information about local breeding communities. One specific type of information may be the location of potential breeding sites.     

Delayed colonisation of arable fields by spring breeding ground beetles (Coleoptera: Carabidae) in landscapes with a high availability of hibernation sites

The colonisation of winter barley fields by spring breeding carabids and its temporal modulation by the amount of potential hibernation sites was studied. Species richness of carabids was lower in landscapes with high length of boundaries and a high amount of non-cropped open habitats during early stages of the beetles’ colonisation of arable fields. Species number of beetles with high dispersal potential responded to this landscape features at coarse spatial scales whereas beetles with low dispersal potential responded to intermediate scales. However, the negative impact of potential hibernation sites on colonisation diminished in later sampling phases. The patterns observed may be explained by both overwintering in arable soils in less complex landscapes and delayed colonisation in more complex landscapes. The seasonal patterns of landscape control suggest a need to account for temporal dynamics in interactions between species or functional groups and landscape properties. A high temporal resolution is needed in studies that focus on ecosystem function and services in agricultural landscapes, as direction of effect (positive/negative) of management on animal communities may change across spatial scales and within short time periods.     

Migratory and resident blue tits Cyanistes caeruleus differ in their reaction to a novel object

Individuals differ consistently in their behavioural reactions towards novel objects and new situations. Reaction to novelty is one part of a suit of individually consistent behaviours called coping strategies or personalities and is often summarised as bold or shy behaviour. Coping strategies could be particularly important for migrating birds exposed to novel environments on their journeys. We compared the average approach latencies to a novel object among migrants and residents in partially migratory blue tits Cyanistes caeruleus. In this test, we found migrating blue tits to have shorter approach latencies than had resident ones. Behavioural reactions to novelty can affect the readiness to migrate and short approach latency may have an adaptive value during migration. Individual behaviour towards novelty might be incorporated among the factors associated with migratory or resident behaviour in a partially migratory population.     

Biological exposure models for oil spill impact analysis

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Which regional features of Danish agriculture favour the corn bunting in the contemporary farming landscape?

Corn buntings Miliaria calandra were abundant throughout arable agricultural landscapes in Europe, but have catastrophically declined since the mid 1970s with changes in farming practice and now give serious conservation cause for concern. Corn buntings declined in Denmark during 1976–1993, but (almost unique in Europe) have since increased (by up to 11% per annum) in some areas without specific conservation recovery actions. Based on breeding bird surveys in the mid 1990s, highest corn bunting densities occurred on mixed agriculture in west Denmark (Jylland); the species was rarer or absent in regions of highest arable land cover. Corn bunting density and extent of rotational and permanent grassland were correlated, but not with spring sown barley (all known to constitute important corn bunting winter habitat). The extent of spring barley rapidly declined in Denmark during the 1980s, but since 1990, most counties have since shown 2–3% annual increases in this crop, except in Nordjylland, where high densities of corn buntings have remained stable. Elsewhere in Jylland, corn buntings have increased in counties supporting highest densities during the mid 1990s, contrasting stable or declining trends in south and east Denmark where densities were originally lower. After dramatic decreases everywhere in Denmark, corn buntings retain highest breeding densities associated with mixed agriculture, especially where grassland and spring sown barley remained in greatest extent. Although purely based on land use correlation and bird surveillance, these results show an association between mixed farming and favourable conservation status of a species now red-listed throughout much of Europe. Further investigations of habitat use at small spatial scales and throughout the annual cycle are urgently required to better enlighten specific recommendations for wider applicability of guidelines for corn bunting recovery actions elsewhere.     

Conditions controlling the onset of breeding migration of the Japanese mountain stream frog, <Emphasis Type="Italic">Rana sakuraii</Emphasis>

I studied the conditions controlling the timing of breeding migration of the Japanese mountain stream frog Rana sakuraii, an explosive breeder, over 9 years (1992, 1993 and 1999–2005). I analysed two factors: the daily cumulative temperature (DCT) during hibernation and the triggering temperature on the day of migration onset. Frogs hibernated in shallow running water in streams in December. A total of 53,155 breeding migrating adults were captured by traps. Every year, breeding migration was induced by a rise in daily maximum water temperature to about 5°C. However, its date was limited to 1 February at the earliest, and the onset needed a DCT (from 20 January, using 3°C as the threshold for daily effective temperature) of at least about 15°C. Earlier (e.g. in mid- or late January), even if the maximum temperature rose to 5–8°C, migration did not begin. Moreover, even in early February, if the maximum temperature rose to 5°C, if it had been too cold in January and the DCT was low, migration would not begin until mid- or late February. Thus, the earliest date of readiness for migration varied from 1 February to mid-February, depending on the winter DCT. Logistic regression analyses showed that both factors, the DCT and the daily temperature, were significant. I propose that in temperate-zone amphibian explosive breeders, both the passing of an essential number of days and an essential DCT during hibernation are prerequisites for the onset of breeding migration before the daily temperature rises to the threshold.     

Non-breeding habitat preference affects ecological speciation in migratory waders

Models of ecological speciation predict that certain types of habitat should be more conducive to species diversification than others. In this study, I test this hypothesis in waders of the sub-order Charadrii using the number of morphological sub-species per species as an index of diversity. I classified all members of this clade as spending the non-breeding season either coastally or inland and argue that these represent fundamentally different environments. Coastal mudflats are characterised by high predictability and patchy worldwide distribution, whilst inland wetlands are widespread but unpredictable. The results show that migratory species that winter coastally are sub-divided into more sub-species than those that winter inland. This was not the case for non-migratory species. I argue that coastal environments select for more rigid migratory pathways, whilst inland wetlands favour more flexible movement patterns. Population sub-division could then result from the passive segregation of breeding sites or from the active selection for assortative mating of ecomorphs.     

辽宁滨海(环渤海)湿地鸟类多样性调查与研究

本研究于2019－2020年对辽宁滨海（环渤海）湿地开展了春迁徙季、秋迁徙季、繁殖季和越冬季的鸟类多样性调查,共监测到鸟类150种,隶属于16目38科,其中,留鸟25种,夏侯鸟26种,冬侯鸟26种,旅鸟73种;国家一级保护鸟类11种,国家二级保护鸟类20种。调查显示:调查区内鸟类多样性的季节差异明显,春、秋迁徙季,鸟类种类与数量明显高于其他两季,越冬季鸟类的种类数量均最少,丰富度指数、均匀度指数、多样性指数和优势度指数均最低。空间分布显示,盘锦地区鸟类种类和数量均最多,达到78种,473084只;营口地区调查到的鸟类种类最少,为55种;锦州地区鸟类数量最少,只有57723只。从鸟类多样性对比来看,盘锦地区最高,大连地区最低,各地鸟类群落组成的空间差异较为明显,在一定程度上受到该地区鸟类生境种类数量及面积大小的影响。辽宁滨海（环渤海）湿地作为东亚—澳大利西亚候鸟迁徙带上的关键区域,当前正面临气候变化、人类围垦干扰造成的鸟类栖息地减少等不利情况,建议通过扩大保护范围、实施修复和管理对策措施、强化宣传等手段保护鸟类多样性及其栖息生境,巩固渤海综合治理攻坚战的治理成效。     

Bird species migration ratio in East Asia, Australia, and surrounding islands

Bird migration and its relationship with the contemporary environment have attracted long-term discussion. We calculated the avian migration ratio (the proportion of breeding species that migrate) in the areas from 70°E to 180°E and examined its relationship with the annual ranges of ambient temperature, primary productivity (estimated by the Enhanced Vegetation Index), and precipitation, along with island isolation and elevational range. The avian migration ratio increased with increasing latitude in general but varied greatly between the two hemispheres. Additionally, it showed minimal differences between continents and islands. Our analyses revealed that the seasonality of ambient temperature, which represents the energy expenditure of birds, is the dominant factor in determining bird species migration. Seasonality in primary productivity and other environmental factors play an indirect or limited role in bird species migration. The lower avian migration ratio in the Southern Hemisphere can be attributed to its paleogeographical isolation, stable paleoclimate, and warm contemporary environment. Under current trends of global warming, our findings should lead to further studies of the impact of warming on bird migration.     

Effects of landscape complexity on farmland birds in the Baltic States

Data on birds occurring in farmland in the Baltic States of Estonia, Latvia and Lithuania were related to the spatial organisation of farmed habitats in three different agricultural landscape types. Species richness, abundance, and diversity of farmland bird communities, as well as abundance of the most frequently observed species were positively related to the number of residual non-cropped elements within farmland, the local mixture of annual crop and grass fields, and the variety of field types. The positive association of the species richness and abundance of the farmland bird community with richness in residual habitats and crops was most prominent in open landscapes. The results suggest that, by simplifying farmland structure and making it more homogenous, EU agricultural policies will have a detrimental effect on farmland bird populations in Eastern Europe. Ways of better targeting of the agri-environment schemes are suggested.     

Sex- and age-related differences in the timing and body condition of migrating Reed Warblers Acrocephalus scirpaceus and Sedge Warblers Acrocephalus schoenobaenus

The migration strategies of birds may vary strongly between species and also between age and/or sex groups. We studied the autumn migration and body condition of molecularly sexed Reed Warblers Acrocephalus scirpaceus and Sedge Warblers Acrocephalus schoenobaenus (211 and 208 ind., respectively) at a stopover site on Lake Druzno, Northern Poland, in 2008. Immature male Reed Warblers were caught significantly later than females (median dates 9 days later), but in the Sedge Warbler, both sexes of immatures migrated at about the same time. Adult males and females of both species did not differ in their time of migration. Adult and immature males of both species were larger (wing length and body mass) than females. In both species, fat reserves were similar in both sexes of both age classes. Adults of both sexes of Reed and Sedge Warbler were generally caught earlier than immatures. In both species, the body mass and fat reserves of immatures were generally less than in adults. The autumn protogyny of immature Reed Warblers may allow smaller females to limit competition with bigger males during migration and at the wintering grounds. In the Sedge Warbler, which tends to match its migration to peak of occurrence of superabundant food at stopover sites, both sexes gain an advantage from migrating at the same time. Since part of the measured wing length variation in both species was explained by sex differences, temporal trends in wing length recorded at stopover sites should be interpreted with caution.     

Climatic influences on the breeding biology of the agile frog (<Emphasis Type="Italic">Rana dalmatina</Emphasis>)

Severe population declines of amphibians have been shown to be attributed to climate change. Nevertheless, the various mechanisms through which climate can influence population dynamics of amphibians remain to be assessed, notably to disentangle the relative synergetic or antagonistic influences of temperature and precipitations on specific life history stages. We investigated the impact of rainfall and temperature on the egg-clutch abundance in a population of agile frog (Rana dalmatina) during 29 years (1987–2016) on 14 breeding sites located in Brenne Natural Park, France. Specifically, we examined the influence of environmental conditions occurring during five temporal windows of the year cycle corresponding to specific life history stages. Overall, our results suggest that the year-to-year fluctuations of egg-clutch abundances in Brenne Natural Park were partly dependent on local climatic conditions (rainfall and temperature). Climate seemed to influence breeding frogs during the autumn-winter period preceding reproduction. Spring and summer conditions did not influence reproduction. Additionally, we failed to detect effects of climatic conditions on newly metamorphosed individuals. Other factors such as density dependence and inter-specific interactions with introduced predators are likely to play a significant role in reproduction dynamics of the studied frog populations.     

Factors determining presence of passerines breeding within White Stork <Emphasis Type="Italic">Ciconia ciconia</Emphasis> nests

Nests of White Stork Ciconia ciconia are commonly used by various passerines as nesting sites. In this study, we investigated factors determining presence and number of pairs of species breeding within White Stork nests in an extensive farmland in NE Poland. In 133 (57%) out of 233 White Stork nests, we found at least one breeding pair of passerine bird. These were from three species: House Sparrows Passer domesticus (68% of 133 nests with co-breeding), Tree Sparrows Passer montanus (65%), and Starlings Sturnus vulgaris (30%). The probability of breeding passerines within White Stork nests increased with increasing nest thickness, and was significantly higher in currently occupied nests. Sparrows were more likely to breed in White Stork nests located on electricity poles, situated closer to settlements and surrounded mainly by arable fields where meadows were not prevalent. In this paper, we show that White Stork nests are favorable nesting sites for passerines, as they are well insulated and provide an anti-predatory shield.     

1980—2015年京津冀地区雁鸭类潜在栖息地连接度变化及驱动因素

近几十年来,京津冀地区湿地面积萎缩,生态功能下降,严重影响了湿地水鸟的栖息地质量及栖息地间的连接,对水鸟类的迁徙、繁殖产生较大干扰,为了从整体上定量分析近几十年来湿地栖息地质量及连接度的变化,采用MaxEnt模型（maximum entropy model,最大熵模型）,基于京津冀地区雁鸭类的出现点和环境因子反演其潜在栖息地,并根据图论法原理,利用ECA指数（equivalent connected area,等效连接指数）,度量1980-2015年潜在栖息地连接度的变化.结果表明:①1980-2015年京津冀雁鸭类潜在栖息地面积总体呈下降趋势,由1980年的6 898 km2减至2015年的3 764 km2,减少了45.0%,其中,低适宜区面积降幅最为明显,为50.5%;2009-2015年高适宜区有增加趋势.②1980-2015年雁鸭类潜在栖息地连接度呈持续下降趋势,2000年以后的连接度下降速率大于2000年以前;潜在栖息地连接结构的空间分布有向环渤海地区转移的趋势,连接结构变化最为明显的区域为京津冀中部平原地区.研究显示,区域气候趋暖变干、水资源的供需失衡、水利工程设施建设以及人口增加与经济发展带来的土地利用变化是潜在栖息地面积与连接度下降的主要影响因子,而近些年保护区的建立及生态补水措施等生态保护政策对高适宜区的退化有一定缓解作用.      

Inter and intra-population variation in shoaling and boldness in the zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

Population differences in anti-predator behaviour have been demonstrated in several species, although less is known about the genetic basis of these traits. To determine the extent of genetic differences in boldness (defined as exploration of a novel object) and shoaling within and between zebrafish (Danio rerio) populations, and to examine the genetic basis of shoaling behaviour in general, we carried out a study that involved laboratory-raised fish derived from four wild-caught populations. Controlling for differences in rearing environment, significant inter-population differences were found in boldness but not shoaling. A larger shoaling experiment was also performed using one of the populations as the basis of a North Carolina type II breeding design (174 fish in total) to estimate heritability of shoaling tendency. A narrow-sense heritability estimate of 0.40 was obtained, with no apparent dominance effects.     

Emergence of long distance bird migrations: a new model integrating global climate changes

During modern birds history, climatic and environmental conditions have evolved on wide scales. In a continuously changing world, landbirds annual migrations emerged and developed. However, models accounting for the origins of these avian migrations were formulated with static ecogeographic perspectives. Here I reviewed Cenozoic paleoclimatic and paleontological data relative to the palearctic–paleotropical long distance (LD) migration system. This led to propose a new model for the origin of LD migrations, the ‘shifting home’ model (SHM). It is based on a dynamic perspective of climate evolution and may apply to the origins of most modern migrations. Non-migrant tropical African bird taxa were present at European latitudes during most of the Cenozoic. Their distribution limits shifted progressively toward modern tropical latitudes during periods of global cooling and increasing seasonality. In parallel, decreasing winter temperatures in the western Palearctic drove shifts of population winter ranges toward the equator. I propose that this induced the emergence of most short distance migrations, and in turn LD migrations. This model reconciliates ecologically tropical ancestry of most LD migrants with predominant winter range shifts, in accordance with requirements for heritable homing. In addition, it is more parsimonious than other non-exclusive models. Greater intrinsic plasticity of winter ranges implied by the SHM is supported by recently observed impacts of the present global warming on migrating birds. This may induce particular threats to some LD migrants. The ancestral, breeding homes of LD migrants were not ‘northern’ or ‘southern’ but shifted across high and middle latitudes while migrations emerged through winter range shifts themselves. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Satellite Remote Sensing of Terrestrial Net Primary Production for the Pan-Arctic Basin and Alaska

We applied a terrestrial net primary production (NPP) model driven by satellite remote sensing observations of vegetation properties and daily surface meteorology from a regional weather forecast model to assess NPP spatial and temporal variability for the pan-Arctic basin and Alaska from 1982 to 2000. Our results show a general decadal trend of increasing NPP for the region of approximately 2.7%, with respective higher (3.4%) and lower (2.2%) rates for North America and Eurasia. NPP is both spatially and temporally dynamic for the region, driven largely by differences in productivity rates among major biomes and temporal changes in photosynthetic canopy structure and spring and summer air temperatures. Mean annual NPP for boreal forests was approximately 3 times greater than for Arctic tundra on a unit area basis and accounted for approximately 55% of total annual carbon sequestration for the region. The timing of growing season onset inferred from regional network measurements of atmospheric CO₂ drawdown in spring was inversely proportional to annual NPP calculations. Our findings indicate that recent regional warming trends in spring and summer and associated advances in the growing season are stimulating net photosynthesis and annual carbon sequestration by vegetation at high latitudes, partially mitigating anthropogenic increases in atmospheric CO₂. These results also imply that regional sequestration and storage of atmospheric CO₂ is being altered, with potentially greater instability and acceleration of the carbon cycle at high latitudes.     

Mate guarding and territorial aggression vary with breeding synchrony in golden whistlers (<Emphasis Type="Italic">Pachycephala pectoralis</Emphasis>)

Male paternity assurance behaviour during the female fertile period has been widely documented amongst birds. In contrast, how sex-specific behavioural strategies vary with local breeding synchrony levels remains largely unknown. This is important because, in many species, intra-population patterns of extra-pair fertilisation rates, and hence cuckoldry risk, are known to vary with the number of simultaneously fertile females. Each sex may therefore differ in how they behave towards male conspecifics during different degrees of breeding synchrony. Here I provide evidence of such sex-specific differences in the golden whistler (Pachycephala pectoralis), a species in which within-pair paternity assurance is negatively associated with breeding synchrony. Via simulated territorial intrusions using decoy males, I show that males, but not females, increase levels of aggression to male intruders during periods of low synchrony, possibly because cuckoldry risk is greatest during this period. In addition, males appear to invest more effort into mate guarding after, but not before, territorial intrusions during this period. These inter-sexual differences may reflect conflicts in interest between the sexes, with females consistently showing interest in males during the fertile period regardless of synchrony levels and males investing more resources into expelling intruders when the risk of paternity loss is greatest. This study thus provides evidence that males may be able to detect variation in breeding synchrony and cuckoldry risk and adjust their paternity assurance behaviour accordingly.