Process in the evolution of bird migration and pattern in avian ecogeography

Current ideas about the evolution of bird migration equate its origin with the first appearance of fully migratory populations, and attribute its evolution to a selective advantage generated by increased breeding success, gained through temporary emigration from resident populations to breed in under-exploited seasonal areas. I propose an alternative hypothesis in which migration first appears as a temporary directional shift away from the breeding site outside the reproductive period, in response to seasonal variation in the direction and/or severity of environmental gradients. Fully migratory populations then appear through either extinction of sedentary phenotypes, or colonisation of vacant seasonal areas by migrants. Where colonisation occurs, resident ancestral populations can be driven to extinction by competition from migrants which invade their range outside the breeding season, resulting in fully migratory species. An analogous process drives the evolution of migration between high latitudes and the tropics, since extension of breeding range into higher latitudes may drive low latitude populations to extinction, resulting in an overall shift of breeding range. This process can explain reverse latitudinal gradients in avian diversity in the temperate zone, since the breeding ranges of migratory species concentrate in latitudes where they enjoy the highest breeding success. Near absence of forest-dwelling species among Palaearctic-African migrants is attributable to the lack of forest in northern Africa for much of the Tertiary, which has precluded selection both for southward extension of migration by west Palaearctic forest species, and northward breeding colonisation by African forest species.     

The relationship between latitude,migration and the evolution of bird song complexity

For the past several decades it has been proposed that birds show latitudinal variation in song complexity. How universal this variation may be and what factors generate it, however, are still largely unknown. Furthermore, while migration is confounded with latitude, migratory behaviour alone may also be associated with variation in song complexity. In this paper we review the literature to assess current ideas on how latitude and migratory behaviour may drive large‐scale geographical patterns of song complexity. At least seven distinct hypotheses have been proposed in 29 studies of the topic. Four of these hypotheses posit that sexual selection pressures co‐vary with latitude and/or migration, resulting in concordant changes in song. Other hypotheses suggest that mechanisms other than sexual selection, such as large‐scale changes in environmental sound transmission properties, may be at play. Sixteen studies found support for increased song complexity with increased latitude and/or migration, whereas 13 did not. Relatively few studies exist on this topic, and methodological differences between them and variable definitions of ‘complexity’ make it difficult to determine whether results are comparable and concordant. At a minimum, it is possible to conclude there is no strong evidence that song complexity increases with latitude and/or migration in all birds. Future work should focus on examining multiple hypotheses at once to further advance our understanding of how latitude, migration and song complexity may or may not be related.     

A robust tool highlights the influence of bird migration on influenza A virus evolution

One of the fundamental unknowns in the field of influenza biology is a panoramic understanding of the role wild birds play in the global maintenance and spread of influenza A viruses. Wild aquatic birds are considered a reservoir host for all lowly pathogenic avian influenza A viruses (AIV) and thus serve as a potential source of zoonotic AIV, such as Australasian‐origin H5N1 responsible for morbidity and mortality in both poultry and humans, as well as genes that may contribute to the emergence of pandemic viruses. Years of broad, in‐depth wild bird AIV surveillance have helped to decipher key observations and ideas regarding AIV evolution and viral ecology including the trending of viral lineages, patterns of gene flow within and between migratory flyways and the role of geographic boundaries in shaping viral evolution (Bahl et al. 2009 ; Lam et al. 2012 ). While these generally ‘virus‐centric’ studies have ultimately advanced our broader understanding of AIV dynamics, recent studies have been more host‐focused, directed at determining the potential impact of host behaviour on AIV, specifically, the influence of bird migration upon AIV maintenance and transmission. A large number of surveillance studies have taken place in Alaska, United States—a region where several global flyways overlap—with the aim of detecting the introduction of novel, Australasian‐origin highly pathogenic H5N1 AIV into North America. By targeting bird species with known migration habits, long‐distance migrators were determined to be involved in the intercontinental movement of individual AIV gene segments, but not entire viruses, between the Australasian and North American flyways (Koehler et al. 2008 ; Pearce et al. 2010 ). Yet, bird movement is not solely limited to long‐distance migration, and the relationship of resident or nonmigratory and intermediate‐distance migrant populations with AIV ecology has only recently been explored by Hill et al. ( 2012 ) in this issue of Molecular Ecology. Applying a uniquely refined, multidimensional approach, Hill et al. validate the innovative use of stable isotope assays for qualifying migration status of wild mallards within the Pacific flyway. The authors reveal that AIV prevalence and diversity did not differ in wintering mallard ducks with different migration strategies, and while migrant mallards do indeed introduce AIV, these viruses do not circulate as the predominant viruses in resident birds. On the other hand, resident mallards from more temperate regions act as reservoirs, possibly contributing to the unseasonal circulation and extended transmission period of AIV. This study highlights the impact of animal behaviour on shaping viral evolution, and the unique observations made will help inform prospective AIV surveillance efforts in wild birds.     

Barriers and distances as determinants for the evolution of bird migration links: the arctic shorebird system

We analysed migratory connectivity between different winter quarters and breeding sectors in the circumpolar tundra region for arctic shorebirds, in relation to migratory distances and ecological barriers. Total distances and barriers were calculated and measured for all potential migratory orthodrome links between 10 selected winter regions and 12 breeding sectors. The migratory segment between the northernmost stopover site and the breeding area, associated with the entry to and exit from the tundra during spring and autumn migration, respectively, was also identified and measured for each potential link.The analysis indicated that the evolution of migratory links among arctic shorebirds is constrained not by distance as such but by distance across ecological barriers, possibly because of the complex adaptations required for barrier crossing and extensive detour migration (and in a few cases because barrier distances exceed the birds' theoretical flight range capacity). A particularly pronounced barrier effect of the Arctic Ocean, as apparent from a sharp decline in migratory connectivity between the opposite sides of the Arctic Ocean, may reflect a crucial importance of favourable entry and exit conditions for successfully occupying different sectors of the tundra breeding area by shorebirds from winter regions situated at widely different total distances in both the southern and northern hemispheres.     

Detours in bird migration

Bird migration routes often follow detours where passages across ecological barriers are reduced in extent. This occurs in spite of the fact that long barrier crossings are within the birds' potential flight range capacity. Long-distance flights are associated with extra energy costs for transport of the heavy fuel loads required. This paper explores how important the fuel transport costs, estimated on the basis of flight mechanics, may be to explain detours for birds migrating by flapping flight. Maximum detours in relation to expanse of the barrier are predicted for cases where birds travel along the detour by numerous short flights and small fuel reserves, divide the detour into a limited number of flight steps, and where a reduced barrier passage is included in the detour. The principles for determining the optimum route, often involving a shortcut across part of the barrier, are derived. Furthermore, the effects of differences in fuel deposition rates and in transport costs for the profitability of detours are briefly considered. An evaluation of a number of observed and potential detours in relation to the general predictions of maximum detours, indicates that reduction of fuel transport costs may well be a factor of widespread importance for the evolution of detours in bird migration at wide ecological barriers.     

The economics of bird migration

Piersma, T. 2000. The economics of bird migration. Ostrich 71(1): 362.

Through their migrations birds spin a worldwide web of migratory routes. Even at the level of individuals and populations, the migrations can cover a large part of the globe. Many factors are involved in the evolution and maintenance of all these migration systems: the contingencies of earth history, the sensory and metabolic evolutionary potential of flying organisms, the distribution of predators and parasites, and energetic considerations. In this presentation I aim to briefly outline the use of energetics in studies on the maintenance of today's flyways. My overview will focus on the shorebird species seasonally migrating between wintering grounds in coastal African and the breeding grounds in the Canadian, Greenlandic, European and Siberian Arctic.

Every individual animal has to balance its energy budget but depending on size, it can balance the discrepancies between energy expenditure and energy intake through the use or storage of nutrients at various time scales. Energy thus provides a common currency to relate ecological phenomena that are separated in space and time. I will describe methods to estimate energy expenditure, energy intake and the use of energy stores, and how these estimates can be translated as an annual energy budget of a coastal shorebird species. This will then be used to evaluate the repercussions of wintering at tropical latitudes and at temperate latitudes, and how such knowledge might help us to understand the evolution of a migration system. Population-specific energy budgets provide only one of the possible uses of energetics approach. Studies at the individual level, at which it is rarely possible to quantify such budgets completely, nevertheless benefit from energetic considerations when the functional significance of variations in migratory timing and seasonal body mass and plumage changes are the focus of attention.     

Extrapair paternity and the evolution of bird song

Bird song is usually considered to have evolved in the contextof sexual selection. Because extrapair paternity is a majorcomponent of sexual selection, mating advantages at the sociallevel for males that produce songs of high quality may be transformedinto higher success in extrapair paternity. Therefore, maleswith longer and more complex songs should suffer less from extrapairpaternity intraspecifically, whereas species with high ratesof extrapair paternity, reflecting intense sperm competition,should produce more elaborate songs. Although some intraspecificstudies demonstrated a negative link between features of songsand extrapair paternity in own nest, others failed to detectsuch a relationship. Contrary to expectation, a meta-analysisof all studies revealed no significant intraspecific evidencefor songs being associated with extrapair paternity. In addition,in comparative analyses based on generalized least squares (GLS)models, we found that no measures of song complexity and temporaloutput were significantly related to extrapair paternity interspecifically,even when potentially confounding factors such as social matingsystem, life history, migration, habitat, or sexual dichromatismwere held constant. Only plumage dichromatism was significantlyrelated to extrapair paternity. The absence of both intra- andinterspecific relationships between measures of song variabilityand extrapair paternity suggests that factors other than postmatingsexual selection have been the important evolutionary forcesshaping differences in song.     

Genetic divergence and migration patterns in a North American passerine bird: implications for evolution and conservation

Like many other migratory birds, the black-throated blue warbler (Dendroica caerulescens) shows pronounced differences in migratory behaviour and other traits between populations: birds in the southern part of the breeding range have darker plumage and migrate to the eastern Caribbean during the winter, whereas those in the north have lighter plumage and migrate to the western Caribbean. We examined the phylogeography of this species, using samples collected from northern and southern populations, to determine whether differentiation between these populations dates to the Pleistocene or earlier, or whether differences in plumage and migratory behaviour have arisen more recently. We analysed variation at 369 bp of the mitochondrial control region domain I and also at seven nuclear microsatellites. Analyses revealed considerable genetic variation, but the vast majority of this variation was found within rather than between populations, and there was little differentiation between northern and southern populations. Phylogeographic analyses revealed a very shallow phylogenetic tree, a star-like haplotype network, and a unimodal mismatch distribution, all indicative of a recent range expansion from a single refugium. Coalescent modelling approaches also indicated a recent common ancestor for the entire group of birds analysed, no split between northern and southern populations, and high levels of gene flow. These results show that Pleistocene or earlier events have played little role in creating differences between northern and southern populations, suggesting that migratory and other differences between populations have arisen very recently. The implications of these results for the evolution of migration and defining taxonomic groups for conservation efforts are discussed.     

Phenology and the changing pattern of bird migration in Britain

 Britain has a huge mass of data on the timing of bird migration, although much of this remains in a form that is inaccessible for immediate scientific study. In this paper, I undertake a preliminary examination of data from a number of historical and current sources. Among these are the Marsham family records from Norfolk, dating back to 1736, and post-World War II records from coastal bird observatories. The majority of the examined time series displayed a negative relationship to temperature indicating a tendency for the earlier arrival of the studied birds in warmer springs. In addition to temperature effects, trends through time and some sampling effects (through population size) have become apparent. Identification and curation of data sources and further analysis is still required to produce a clearer picture of climate effects on bird migration timing and on subsequent bird population dynamics. Received: 26 May 1998 / Accepted: 28 October 1998     

Urban noise and the cultural evolution of bird songs

In urban environments, anthropogenic noise can interfere with animal communication. Here we study the influence of urban noise on the cultural evolution of bird songs. We studied three adjacent dialects of white-crowned sparrow songs over a 30-year time span. Urban noise, which is louder at low frequencies, increased during our study period and therefore should have created a selection pressure for songs with higher frequencies. We found that the minimum frequency of songs increased both within and between dialects during the 30-year time span. For example, the dialect with the highest minimum frequency is in the process of replacing another dialect that has lower frequency songs. Songs with the highest minimum frequency were favoured in this environment and should have the most effective transmission properties. We suggest that one mechanism that influences how dialects, and cultural traits in general, are selected and transmitted from one generation to the next is the dialect''s ability to be effectively communicated in the local environment.     

A polar system of intercontinental bird migration

Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64 degrees N in Bering Strait up to latitude 75 degrees N far north of Wrangel Island, with eastward flights making up 79% of all track directions.The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian-American bird migration system in a wide Arctic sector between longitudes 110 degrees E and 130 degrees W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3-5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40-80 degrees of longitude, corresponding to distances and durations of 1400-2600 km and 26-48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.     

A review of bird migration over Israel

The sheer number of migrating birds that pass over Israel give the country a unique situation in the study of the phenomenon, which was, not surprisingly, noticed even in biblical times. This paper is a review of current knowledge of bird migration in Israel. It includes a brief summary of migration in the Mediterranean basin and the Middle East and new information from an extensive migration survey done in the northern Arava Valley and Negev Highlands.