A comparative analysis of aggression in migratory and non-migratory salmonids

This paper represents a comprehensive test of the hypothesis that aggression in salmonids increases with the duration of stream residence. The intraspecific aggression of eleven juvenile salmonids was compared with their normal duration of stream residence. Salmonids maintained in 1 metre tanks and observed over two days could be separated into four groups based on statistical differences between the frequencies of aggressive behaviour. Non-anadromous Salvelinus fontinalis showed the highest levels of aggressive behaviour, followed by Oncorhynchus masou and O. mykiss (rainbow and steelhead trout). Least aggressive were the early migrants O. nerka (kokanee and sockeye salmon) and O. keta. The remaining species, O. rhodurus, O. kisutch, S. leucomaenis pluvius and Salmo trutta formed a moderately aggressive group. Aggressiveness was significantly and positively correlated with the duration of stream residence.     

Annual routines of non-migratory birds: optimal moult strategies

In a periodically changing environment it is important for animals to properly time the major events of their life in order to maximise their lifetime fitness. For a non-migratory bird the timing of breeding and moult are thought to be the most crucial. We develop a state-dependent optimal annual routine model that incorporates explicit density dependence in the food supply. In the model the birds' decisions depend on the time of year, their energy reserves, breeding status, experience, and the quality of two types of feathers (outer and inner primaries). Our model predicts that, under a seasonal environment, feathers with large effects on flight ability, higher abrasion rate and lower energetic cost of moult should be moulted closer to the winter (i.e. later) than those with the opposite attributes. Therefore, we argue that the sequence of moult may be an adaptive response to the problem of optimal timing of moult of differing feathers within the same feather tract. The model also predicts that environmental seasonality greatly affects optimal annual routines. Under high seasonality birds breed first then immediately moult, whereas under low seasonality an alternation occurs between breeding and moulting some of the feathers in one year and having a complete moult but no breeding in the other year. Increasing food abundance has a similar effect.     

五种迁飞蛾类及非迁飞近似种前翅形状的数值分析

以翅面正投影周缘曲率为指标,对五种蛾类迁飞昆虫及与其近似的非迁飞昆虫的前翅翅形进行数值分析。结果表明,二者之间存在可以相互区别的差异：迁飞种类前翅前缘较平直,特别是前缘1/2处到翅尖的平均曲率绝对值显著小于非迁飞种类;而后缘1/2处到翅尖的平均曲率绝对值偏大,即这段后缘较弯曲,向外凸出,使翅外侧较宽阔。这种翅形特征较适合于利用气流进行远距离迁飞     

Optimal moult strategies in migratory birds

Avian migration, which involves billions of birds flying vast distances, is known to influence all aspects of avian life. Here we investigate how birds fit moult into an annual cycle determined by the need to migrate. Large variation exists in moulting patterns in relation to migration: for instance, moult can occur after breeding in the summer or after arrival in the wintering quarters. Here we use an optimal annual routine model to investigate why this variation exists. The modelled bird's decisions depend on the time of year, its energy reserves, breeding status, experience, flight feather quality and location. Our results suggest that the temporal and spatial variations in food are an important influence on a migratory bird's annual cycle. Summer moult occurs when food has a high peak on the breeding site in the summer, but it is less seasonal elsewhere. Winter moult occurs if there is a short period of high food availability in summer and a strong winter peak at different locations (i.e. the food is very seasonal but in opposite phase on these areas). This finding might explain why only long-distance migrants have a winter moult.     

The scheduling of molt in migratory birds

Summary We model the yearly cycle of small migratory birds to explain the variation in scheduling of complete molt, in particular why some birds molt immediately after breeding on the breeding grounds (summer molt) whereas others migrate to their wintering grounds before molt is initiated (winter molt). We employ the method of dynamic programming, because of its suitability for modelling life history traits. Feather quality and latitude entered the model as state variables and were assumed to affect survival rate and reproductive success. Migration and molt were assumed to be associated with increased mortality risks. By changing the parameters in the model we were able to generate most existing molt patterns, including summer and winter molt, biannual (summer and winter) molt, and molt migration. Our model suggests that the scheduling of molt is basically a result of a trade-off between having a high feather quality during breeding versus during the non-breeding period. A high impact of feather quality on survival rate in combination with low costs of molt resulted in biannual molt. Winter molt became more likely as the survival rateper se increased. A low seasonal amplitude in survival rate is a prerequisite for the occurrence of molt migration. Molt duration, migration costs and reproductive successper se were found to have no impact on the timing of molt. We also investigated the effect of benefits from prior occupancy at breeding and winter grounds.     

Patterns of phenological changes in migratory birds

The phenology of avian migration appears to be changing in response to climate change. Seemingly contradictory differences in the timing of these annual cycles have been reported in published studies. We show that differences between studies in the choice of songbird species, as well as in the measurements of migration phenology, can explain most of the reported differences. Furthermore, while earlier spring arrival is evident across these studies, trends in timing of departure show large variation between species and according to individual timing of migration (early-arriving vs. late-departing individuals). Much of the variation in departure between species could be explained by each species’ migratory status. We present a detailed analysis of migrants recorded at a Danish migration site, and reveal that although shifts in migration timing can be demonstrated for almost all species, these shifts are either most pronounced in the early arriving/late departing individuals or the changes are similar. Thus most individuals do not seem to change their breeding-area residence time (BART). As BART is likely to reflect ecologically important factors, e.g. number of clutches, we expect that only small effects have been exerted on the breeding ecology of the studied species in the time period investigated. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Genetic control of migratory behaviour in birds

Although it has long been suspected that biannual migration in birds has a direct genetic basis, only in the last decade have details of the inheritance of behavioural traits such as migratory activity and directional preferences been demonstrated. A model has now been developed to estimate how inexperienced first-time migrants manage to reach their unknown winter quarters on the basis of inherited spatio-temporal programs. Furthermore, in obligate partial migrants the decision to migrate or not has been shown to have a strong genetic base. Migratoriness and sedentariness in partial migrants have been shown to have a high potential for rapid evolution. A recent set of results has suggested that novel migratory habits can evolve in less than 25 years. A possible consequence is that environmental changes, including 'greenhouse' effects, might considerably alter avian migration systems by acting on genetic variation for migratory tendencies.     

Protandry and sexual dimorphism in trans-Saharan migratory birds

Earlier arrival to reproductive sites of males relative to females(protandry) is widespread among migratory organisms. Diversemechanisms have been proposed that may select for protandry,including competition for limiting resources (e.g., territories)or mates. In species with large variation in male reproductivesuccess, such as polygamous species and those with intense spermcompetition, early arriving males may accrue a fitness advantagebecause they acquire more mates or have larger chances of paternity.Comparative studies of birds have shown that sexual size dimorphism(SSD) is positively associated with the level of polygyny, whereasintense sperm competition is associated with sexual dichromatism(SD). Positive correlations between protandry and SSD or SDcan therefore be expected to exist across avian species. Becauselarge males are predicted to be better able to cope with adverseecological conditions early in the breeding season, selectionfor protandry, in turn, may have a correlated response on SSDamong migratory species breeding in boreal latitudes. Althoughprevious studies of birds have analyzed the association betweenSSD and protandry, none has analyzed SD in relation to protandry.Here we analyze the association between protandry during springmigration, SSD, and SD in 21 trans-Saharan monogamous migratorybird species. The difference in median migration dates betweenfemales and males, reflecting protandry, was positively associatedwith SD but not with SSD. Because dichromatism is positivelyrelated to sperm competition across species, present resultsare consistent with predictions derived from sexual selectionhypotheses for the evolution of protandry mediated by spermcompetition.     

Seasonal climatic niches diverge in migratory birds

Migratory birds occupy different geographical areas during breeding and non-breeding periods, and thus different factors may determine their range limits depending on each season. One such factor is the spatial climatic component of the niche, which is widely used to model species distributions, yet the temporal component is often neglected and is generally assumed to be constant. We tested the hypothesis that the climatic niche is conserved between breeding and non-breeding areas in 355 bird species migrating through Eurasian–African flyways. For this, we performed niche overlap analyses and compared niche differences between sister or phylogenetically closely related species, as well as linking the differences to migratory distances. For more than 80% of the species, there was no or very little overlap between their breeding and non-breeding climatic niches. For most closely related species, the degree of overlap of their breeding climatic niches was larger than the overlap observed within each species, but not for their wintering climatic niches, suggesting a phylogenetic conservation of breeding climatic niches. Finally, there was a clear negative relationship between migratory distances and climatic niche overlap within each species. Our results confirmed that the climatic niche of most Eurasian–African migratory species differs between both breeding and non-breeding ranges, suggesting distinctive seasonal climatic requirements. Given these results and the geographically uneven effects of climate change, the impact of global change is likely to have different effects in each seasonal range. Hence, both breeding and non-breeding climatic data need to be considered when using species distribution models.     

Macroecology of habitat choice in long-distance migratory birds

Patterns of habitat choice in ecological communities are not only influenced by present-day selective forces but also by historical processes, such as the biogeographical history of the lineages they are composed of. Nevertheless, it has been very difficult to test historical factors. The possible tropical origin of long-distance migratory birds provides an opportunity for such a test. If habitat choice of long-distance migrants is inherited from their tropical ancestors then Nearctic long-distance migrants might have acquired their habitat choice from Neotropical forest species and European long-distance migrants from African savannah species. Here we use a macroecological approach to show that this hypothesis can be confirmed. Long-distance migrants in the Nearctic are found in forested habitat types, while those in Europe are found in open ones. In comparison, the habitat choice of residents and short-distance migrants (in genera without long-distance migration) does not differ between the Nearctic and Europe. These results demonstrate that habitat choice in temperate bird communities can be explained by the tropical history of long-distance migrants. Thus, habitat choice seems to be shaped not only by local mechanisms, but also by processes acting on much larger spatial and temporal scales.     

Orientation of migratory birds under ultraviolet light

In view of the finding that cryptochrome 1a, the putative receptor molecule for the avian magnetic compass, is restricted to the ultraviolet single cones in European Robins, we studied the orientation behaviour of robins and Australian Silvereyes under monochromatic ultraviolet (UV) light. At low intensity UV light of 0.3 mW/m², birds showed normal migratory orientation by their inclination compass, with the directional information originating in radical pair processes in the eye. At 2.8 mW/m², robins showed an axial preference in the east–west axis, whereas silvereyes preferred an easterly direction. At 5.7 mW/m², robins changed direction to a north–south axis. When UV light was combined with yellow light, robins showed easterly ‘fixed direction’ responses, which changed to disorientation when their upper beak was locally anaesthetised with xylocaine, indicating that they were controlled by the magnetite-based receptors in the beak. Orientation under UV light thus appears to be similar to that observed under blue, turquoise and green light, albeit the UV responses occur at lower light levels, probably because of the greater light sensitivity of the UV cones. The orientation under UV light and green light suggests that at least at the level of the retina, magnetoreception and vision are largely independent of each other.     

Highly pathogenic avian influenza virus H5N1 infection in a long-distance migrant shorebird under migratory and non-migratory states

Corticosterone regulates physiological changes preparing wild birds for migration. It also modulates the immune system and may lead to increased susceptibility to infection, with implications for the spread of pathogens, including highly pathogenic avian influenza virus (HPAIV) H5N1. The red knot (Calidris canutus islandica) displays migratory changes in captivity and was used as a model to assess the effect of high plasma concentration of corticosterone on HPAIV H5N1 infection. We inoculated knots during pre-migration (N = 6), fueling (N = 5), migration (N = 9) and post-migration periods (N = 6). Knots from all groups shed similar viral titers for up to 5 days post-inoculation (dpi), peaking at 1 to 3 dpi. Lesions of acute encephalitis, associated with virus replication in neurons, were seen in 1 to 2 knots per group, leading to neurological disease and death at 5 to 11 dpi. Therefore, the risk of HPAIV H5N1 infection in wild birds and of potential transmission between wild birds and poultry may be similar at different times of the year, irrespective of wild birds'' migratory status. However, in knots inoculated during the migration period, viral shedding levels positively correlated with pre-inoculation plasma concentration of corticosterone. Of these, knots that did not become productively infected had lower plasma concentration of corticosterone. Conversely, elevated plasma concentration of corticosterone did not result in an increased probability to develop clinical disease. These results suggest that birds with elevated plasma concentration of corticosterone at the time of migration (ready to migrate) may be more susceptible to acquisition of infection and shed higher viral titers—before the onset of clinical disease—than birds with low concentration of corticosterone (not ready for take-off). Yet, they may not be more prone to the development of clinical disease. Therefore, assuming no effect of sub-clinical infection on the likelihood of migratory take-off, this may favor the spread of HPAIV H5N1 by migratory birds over long distances.     

The hunting of migratory birds in Europe: a ringing recovery analysis

The hunting of 20 species of migratory birds in Europe and countries bordering the Mediterranean Sea was investigated using ringing recovery data. The intensity of the hunting of birds in each country was measured by the calculation of an index which controls for ringing effort and reporting rates. The hunting of birds was shown to be consistently high in western Mediterranean countries, particularly in southern France, northern and southwestern Iberia, northern Italy and in northwest Africa. Geographical patterns in the relative magnitude of hunting indices for different breeding populations were species-specific. In Europe most birds are taken during autumn and winter but in North Africa hunting is almost equally prevalent in spring. All species investigated, other than those that remain legitimate quarry species, showed a general reduction in index values after 1980. Analysis of long-term trends in index values since 1950 indicated a statistically significant overall decrease in the hunting of the majority of species. These changes are thought to be at least partly attributable to a real decline in the taking of birds but they may also reflect changed attitudes to reporting the hunting of species which are now protected. This analysis provides the first quantitative Europe-wide assessment of geographical and temporal trends in the hunting of migratory birds.     

A highly invasive malaria parasite has expanded its range to non-migratory birds in North America

Parasite range expansions are a direct consequence of globalization and are an increasing threat to biodiversity. Here, we report a recent range expansion of the SGS1 strain of a highly invasive parasite, Plasmodium relictum, to two non-migratory passerines in North America. Plasmodium relictum is considered one of the world''s most invasive parasites and causes the disease avian malaria: this is the first reported case of SGS1 in wild non-migratory birds on the continent. Using a long-term database where researchers report avian malaria parasite infections, we summarized our current understanding of the geographical range of SGS1 and its known hosts. We also identified the most likely geographical region of this introduction event using the MSP1 allele. We hypothesize that this introduction resulted from movements of captive birds and subsequent spillover to native bird populations, via the presence of competent vectors and ecological fitting. Further work should be conducted to determine the extent to which SGS1 has spread following its introduction in North America.     

Climate patterns and the stochastic dynamics of migratory birds

We analyse time series data of 17 bird species trapped at Ottenby Bird Observatory, Sweden, during spring migration 1972–1999. The species have similar demography but respond differently to variation in the North Atlantic Oscillation (NAO) – a strong determinant of winter climate in the northern Hemisphere. Species wintering in northern Europe, compared to species having winter quarters in the Mediterranean area, tend to respond positively to variation in NAO. The variation within each group is high due to wide-ranging winter-distribution in many species, probably smoothing out the effect of spatial variation in NAO. Whereas mild winters (high NAO) is benign for many – but not all – birds wintering in northern Europe, the effect of drier-than-normal conditions in the Mediterranean area during high NAO index winters are uncertain. The work presented here goes beyond simple correlative studies and help identifying which species that are most affected by variation in winter climate. This is a first important step that calls for a more mechanistic approach when analysing possible changes to climate change.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.     

Climate change and the optimal arrival of migratory birds

Recent climate change has sparked an interest in the timing of biological events, which is a general problem in life-history evolution. Reproduction in many organisms breeding in seasonal environments, e.g. migratory birds, is dependent on the exploitation of a short but rich food supply. If the seasonal timing of the food peak advances owing to climate change, then one would expect the bird to track those changes, hence, initiate migration and breeding earlier. However, when there is competition for territories and a risk of pre-breeding mortality, the optimal response to a shifting food distribution is no longer obvious. We develop a theoretical model to study how the optimal arrival time depends on the mean and variance of the food distribution, the degree of competition for territories and the risk of mortality. In general, the optimal shift in arrival date should never be as extreme as the shift in food peak date. Our results also show that we should expect the high variation of trends in arrival date observed among migratory birds, even if migration and information about climate change were unconstrained.