The evolution of avian migration

The study of avian migration has reached sophisticated levels in many areas, including ecology, behaviour, and physiology. Traditional discussions of the evolution of migration, however, have been compromised for several reasons. Previous ideas concerning the ancestral home of migrant species, southern or northern, and whether a partially migratory stage always precedes a fully migratory stage, were not expressed as testable hypotheses. Plotting migratory behaviour on phylogenetic trees has become commonplace and allows tests of traditional hypotheses. Some of these studies are reviewed, lending some support for almost all of the previous ideas. Although phylogenetic mapping helps to frame questions about the evolution of migration in a testable framework, there are two serious issues. First, experimental and observational studies reveal that the expression of migratory behaviour can change rapidly within a lineage, which can violate assumptions of character mapping. In addition, a species distribution model is used to show that current conditions for obligate migratory populations of the chipping sparrow were much restricted at the Last Glacial Maximum, and that the species might have been considered a partial migrant at that time. The expression of migratory behaviour in an extant species might be an artefact of the current inter‐glacial period. Only if the rate of gains and losses of migratory behaviour can be incorporated into a phylogenetic mapping exercise will the actual evolutionary pattern of migration be revealed. For example, reconstruction of the ancestral area and the evolutionary history of migratory categories in a clade of New World warblers depended on the assumptions of character state transitions. A second concern is that the trait ‘migratory’ is too broad for evolutionary analysis and that, if possible, the expression of hyperphagia, Zugunruhe, and navigation could be mapped individually. Loss or suppression of any of these components can lead to sedentary populations, revealing how migratory behaviour can appear and disappear rapidly. A report of low levels of Zugunruhe in a sedentary bird, Saxicola torquata, is reconstructed as derived in a clade of otherwise migratory populations, suggesting that the loss of migration was a result of suppression (but not elimination) of Zugunruhe. When researchers mention the independent origin of migration in a clade, they are most likely referring to the gain or loss of the expression of the ancestral migratory programme, not the de novo evolution of migration per se. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 237–250.     

Characterizing population and individual migration patterns among native and restored bighorn sheep (Ovis canadensis)

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Stochastic compartmental modeling and parameter estimation with application to cancer treatment follow-up studies

In this paper we use marginal probabilities to derive expressions for the means, variances and covariances ofm-compartment systems. We also present an efficient algorithm for the estimation of the parameters of the system using time series data when measurements are available fromk of them compartments. An application of the analysis and parameter estimation procedure for a model representing the results of a cancer treatment follow-up study is given. Supported in part by NSF Grant Number DCR74-17282.     

GENETICS OF PRECALLING PERIOD IN THE ORIENTAL ARMYWORM,MYTHIMNA SEPARATA (WALKER) (LEPIDOPTERA: NOCTUIDAE), AND IMPLICATIONS FOR MIGRATION

Migration in insects usually occurs prereproductively so the precalling period, between emergence and the first release of pheromone, determines the number of nights over which females of migrants such as Mythimna separata can express their migratory potential. Precalling periods (PCP's) ranged from 3 to 21 nights in F₁ females from insects collected in the field in eastern China. As technical problems precluded the use of a sib-analysis design, the genetic basis of this variation was investigated by analysis of the correlation between daughters' and mother's PCP's and by selection in isofemale lines followed by reciprocal crosses. The results of these analyses indicated that PCP is strongly influenced by genes located on the X chromosome, inherited from the male parent in Lepidoptera. Genetic regulation of migratory potential, of which PCP is a crucial component, can be expected to evolve in tropical species such as Mythimna separata, whose larval habitats are associated with unpredictable tropical rainfall. This species has no preimaginal diapause but makes regular seasonal incursions every spring and early summer to high temperate latitudes at which it cannot overwinter. Where migrants are carried by winds, as occurs in these migrations, the predominantly poleward winds at this time of year must create a genetic cline across latitude, in which migratory potential of the summer populations founded by the migrants increases with increasing latitude. We suggest that X-linkage of genes influencing PCP may reduce the genetic load associated with this polymorphism by reducing the frequencies of inappropriate PCP genotypes in these summer populations. Thus, in autumn when winds are predominantly northerly, fewer individuals at any latitude will lack the capability to make the return journey southward. Some support for this model is provided by the observed distributions of PCP's of F₁ females from moths collected in the field in central and northern eastern China. If X-linkage of genes influencing PCP contributes to the stability of these migratory cycles, it can be expected in other species with similar migratory strategies.     

Chances and Limitations of Wild Bird Monitoring for the Avian Influenza Virus H5N1 — Detection of Pathogens Highly Mobile in Time and Space

Highly pathogenic influenza virus (HPAIV) H5N1 proved to be remarkably mobile in migratory bird populations where it has led to extensive outbreaks for which the true number of affected birds usually cannot be determined. For the evaluation of avian influenza monitoring and HPAIV early warning systems, we propose a time-series analysis that includes the estimation of confidence intervals for (i) the prevalence in outbreak situations or (ii) in the apparent absence of disease in time intervals for specified regional units. For the German outbreak regions in 2006 and 2007, the upper 95% confidence limit allowed the detection of prevalences below 1% only for certain time intervals. Although more than 25,000 birds were sampled in Germany per year, the upper 95% confidence limit did not fall below 5% in the outbreak regions for most of the time. The proposed analysis can be used to monitor water bodies and high risk areas, also as part of an early-warning system. Chances for an improved targeting of the monitoring system as part of a risk-based approach are discussed with the perspective of reducing sample sizes.     

Acoustic but no genetic divergence in migratory and sedentary populations of blackcaps,Sylvia atricapilla

In songbirds, territorial songs are key regulators of sexual selection and are learned from conspecifics. The cultural transmission of songs leads to divergence in song characteristics within populations, which can ultimately lead to speciation. Many songbirds also migrate, and individual differences in migratory behaviours can influence population genetic structure and local song differentiation. Blackcaps, Sylvia atricapilla, exhibit versatile territorial songs and show diversity in migration behaviours. They therefore comprise a good model for investigating the relationships between migratory patterns, song variation, and genetic diversity. We studied a migratory population (two groups near Paris) and a sedentary population (three groups in Corsica). All of the birds were ringed and blood sampled to investigate genetic relatedness using 17 microsatellite loci. A detailed song analysis showed that this species has a complex repertoire (> 100 syllables), which required the development of a semi‐supervised method to classify different categories of syllables and compare sequences of syllables. Our analysis showed no genetic structuring among populations: individuals belonging to the same group were not genetically closer than those from different groups. However, we found a strong wingsize difference between sedentary and migratory populations. We also showed that geographical variations in songs rely at least on both syllable and sequence content. Unexpectedly, despite a higher turnover of individuals, migratory groups share as many syllables and sequences as sedentary groups, which raises interesting issues on song learning and the maintenance of dialects in migratory birds.     

Live imaging early immune cell ontogeny and function in zebrafish Danio rerio

The success of a robust vertebrate inflammatory response is in part because of the migratory potential of its haematopoietic components. Once these cells converge at an inflammatory site, they interact with each other as well as non‐immune tissues and infectious agents to help manage both the scale and the duration of any ensuing response. Exactly how these blood cells, that constitute the innate and adaptive immune systems, contribute to such immune responses remain largely unknown. Traditionally, assessing these contributions relied upon histological analysis of fixed tissue sections complemented with in vitro dynamic data. Although informative, translating results from these studies into the multicellular whole‐animal setting remain difficult. Recently, non‐invasive live imaging of the immune system in animal models is providing significant insights into how immune cells function within their intact natural environment. Although the majority of these studies have been conducted within mice, another vertebrate, the zebrafish Danio rerio is being recognized as an ideal platform for non‐invasive live imaging applications. The optical transparency, rapid development, genetic tractability and highly conserved innate and adaptive immune systems of this well‐established developmental model have been exploited in a number of recent studies evaluating the immunocompetence of fluorescently tagged blood cells. In addition, similar live imaging studies are helping to dissect the ontogeny of blood‐cell development by tracking various haematopoietic precursor cells to assess their contribution to different blood lineages. This review will examine some recent advances that have helped D. rerio emerge as a live imaging platform as well as its potential to offer valuable insights into the genetics behind diseases associated with immune cell dysfunction.     

Modelling parasite dissemination: host cell subversion and immune evasion by Toxoplasma gondii

Protozoan parasites belong to the most widespread and devastating human pathogens. Their ability to manipulate host responses and establish infection in their hosts continues to puzzle researchers. Recent developments of experimental model systems are contributing to the discovery of new aspects of the biology of parasite dissemination. Here, we review current knowledge on strategies utilized by the apicomplexan parasite Toxoplasma gondii to disseminate and establish infection in its host. Recent findings have revealed intricate mechanisms by which this obligate intracellular protozoan sequesters cellular functions of the immune system to assure propagation. These mechanisms include the hijacking of migratory leucocytes, modulation of migratory properties of infected cells and rapid transfer of parasites between different leucocyte populations by cytotoxicity‐induced parasite egress. Collectively, Toxoplasma strikes a delicate balance, assuring efficient dissemination and establishment of asymptomatic lifelong infection in its host while protecting its intracellular entity and limiting host pathology.     

In vitro dynamics of chromatin organization and migration

The organization of eukaryotic chromatin is not static but changes as a function of cell status during processes such as proliferation, differentiation, and migration. DNA quantification has not been used extensively to investigate chromatin dynamics in combination with cellular migration. In this context, an optimized DNA-specific, nonperturbant method has been developed for studying chromatin organization, using the fluorescent vital bisbenzimidazole probe Hoechst 33342: this property has been described by Hamori et al. (1980). Computer-assisted image analysis was used to follow migratory activity and chromatin organization of L929 fibroblasts during in vitro wound healing. Cell movements were analyzed using an optical flow technique, which consists in the calculation of the velocity field of cells and nuclear movements in the frame. This system allows the correlation of cell migration and position in the cell cycle. It makes it possible to study chromatin dynamics using a quantitative analysis of nuclear differentiation reorganization (nuclear texture) and to correlate this with migration characteristics. The present system would be of interest for studying cell-extracellular matrix interactions using differing substrates, and also the migratory response to chemotactic factors. Such a model is a prerequisite for gaining better understanding of drug action.     

Adult male birds advance spring migratory phenology faster than females and juveniles across North America

Advances in spring migratory phenology comprise some of the most well-documented evidence for the impacts of climate change on birds. Nevertheless, surprisingly little research has investigated whether birds are shifting their migratory phenology equally across sex and age classes—a question critical to understanding the potential for trophic mismatch. We used 60 years of bird banding data across North America—comprising over 4 million captures in total—to investigate both spring and fall migratory phenology for a total of 98 bird species across sex and age classes, with the exact numbers of species for each analysis depending on season-specific data availability. Consistent with protandry, in spring (n = 89 species), adult males were the first to arrive and immature females were the last to arrive. In fall (n = 98), there was little difference between sexes, but adults tended to depart earlier than juveniles. Over 60 years, adult males advanced their phenology the fastest (−0.84 days per decade, 95 CrI = −1.22 to −0.47, n = 36), while adult and immature females advanced at a slower pace, causing the gap in male and female arrival times to widen over time. In the fall, there was no overall trend in phenology by age or sex (n = 57), driven in part by high interspecific variation related to breeding and molt strategies. Our results indicate consistent and predictable age- and sex-based differences in the rates at which species' springtime phenology is shifting. The growing gap between male and female migratory arrival indicates sex-based plasticity in adaptation to climate change that has strong potential to negatively impact current and future population trends.     

Estimating migratory connectivity of birds when re‐encounter probabilities are heterogeneous

Understanding the biology and conducting effective conservation of migratory species requires an understanding of migratory connectivity – the geographic linkages of populations between stages of the annual cycle. Unfortunately, for most species, we are lacking such information. The North American Bird Banding Laboratory (BBL) houses an extensive database of marking, recaptures and recoveries, and such data could provide migratory connectivity information for many species. To date, however, few species have been analyzed for migratory connectivity largely because heterogeneous re‐encounter probabilities make interpretation problematic. We accounted for regional variation in re‐encounter probabilities by borrowing information across species and by using effort covariates on recapture and recovery probabilities in a multistate capture–recapture and recovery model. The effort covariates were derived from recaptures and recoveries of species within the same regions. We estimated the migratory connectivity for three tern species breeding in North America and over‐wintering in the tropics, common (Sterna hirundo), roseate (Sterna dougallii), and Caspian terns (Hydroprogne caspia). For western breeding terns, model‐derived estimates of migratory connectivity differed considerably from those derived directly from the proportions of re‐encounters. Conversely, for eastern breeding terns, estimates were merely refined by the inclusion of re‐encounter probabilities. In general, eastern breeding terns were strongly connected to eastern South America, and western breeding terns were strongly linked to the more western parts of the nonbreeding range under both models. Through simulation, we found this approach is likely useful for many species in the BBL database, although precision improved with higher re‐encounter probabilities and stronger migratory connectivity. We describe an approach to deal with the inherent biases in BBL banding and re‐encounter data to demonstrate that this large dataset is a valuable source of information about the migratory connectivity of the birds of North America.     

Genetic analysis of sympatric migratory ecotypes of Arctic charr Salvelinus alpinus: alternative mating tactics or reproductively isolated strategies?

Three populations of Arctic charr Salvelinus alpinus from southern Baffin Island were previously identified to display variable migratory phenotypes, with an anadromous component of the population and another remaining resident in fresh water. In this study, 14 microsatellite markers were used to help distinguish between two alternative hypotheses to explain the co‐existence of the two ecotypes: that the two ecotypes originate from a single population and are the result of a conditional mating tactic or that the migratory ecotypes are reproductively isolated populations utilizing alternative migratory strategies. In two of the three replicate systems, F_ST values between the resident and anadromous individuals were non‐significant, while they were significant in a third sampling location. Bayesian clustering analysis implemented in structure, however, failed to identify any within‐location clustering in all three sampling locations. It is concluded from these analyses that the life‐history ecotypes are most likely conditional mating tactics, rather than reproductively isolated populations. Other evidence in favour of the alternative mating tactic hypothesis is briefly reviewed, and implications for management of those populations are discussed.     

A Matrix Model to Study the Colonization by Brown Trout of a Virgin Ecosystem in the Kerguelen Islands

We present a matrix model for the study of the population dynamics of brown trout Salmo trutta L., introduced in the '60s in the virgin aquatic ecosystems of the Kerguelen Islands. This species clearly acclimatized very well: a portion of the population became migratory and spent a part of its life cycle in the sea, which allowed the rapid colonization of two rivers close to the stream of origin in the same bay (Baie Norvégienne).These migratory trout can become a smolt at 2, 3, or 4 years of age. The model takes into account age and smolt age structures and in a first step considers the fish from the Baie Norvégienne as belonging to a single population. The transition matrix looks like a 32 × 32 Leslie matrix in which some survival rates are not on the subdiagonal. They represent survival after the first sea migration and are particularly important for the dynamics of the whole population.The estimate of demographic parameters was obtained from a data base containing information collected in the field since 1970. The model was calibrated on the population size estimate and the stock structure of the migratory trout in 1979. Population size was estimated by tagging-recapture and monitoring of the migratory trout in freshwater when they returned to overwinter or reproduce. Under the hypothesis of a constant survival rate for all ages and categories of fish, it was possible to determine relationships between the annual population growth rate and survival rates at first downstream migration, for which no direct estimate was available.These constraints on the model induced paradoxical results. For instance, an increase in survival rate reduced migratory trout numbers in the first years of colonization. These data suggested that the average survival rate should be around 0.3 and not 0.5 as surmised in previous studies. However, the model systematically underestimated numbers of migratory trout during the first years of development. Thus, to improve the model it will be necessary to introduce survival rates varying with time. Another possible approach would be to consider the population as three subunits corresponding to the three colonized rivers of the Baie Norvégienne.     

Diel migration and vertical distribution of Cladocera in Lake D. Helvecio (Minas Gerais,Brazil)

Six genera of Clad ocera (Diaphanosoma, Daphnia, Ceriodaphnia, Moina, Bosmina, Bosminopsis), each of them usually with only one species were found in Lake D. Helvecio, a natural valley lake located in the eastern part of Brazil. Diurnal migratory movements of the organisms observed in this lake showed a different pattern in different species. Closely related species, which explore the same food source, live in different layers, thus avoiding interspecific competition. The migratory behaviour of the species was studied mainly in relation to temperature and oxygen distribution in the lake. Thus, analyses were made in the summer (January, 1978) when a strong stratification occurs with the establishment of a thermocline and an oxycline. Comparisons were made also with the data obtained in winter (July, 1978), when a complete mixing of water occurs.     

Accounting for heterogeneity when estimating stopover duration,timing and population size of red knots along the Luannan Coast of Bohai Bay,China

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Migratory strategies of waterbirds shape the continental‐scale dispersal of aquatic organisms

Long distance dispersal (LDD) of propagules is an important determinant of population dynamics, community structuring and biodiversity distribution at landscape, and sometimes continental, scale. Although migratory animals are potential LDD vectors, migratory movement data have never been integrated in estimates of propagule dispersal distances and LDD probability. Here we integrated migratory movement data of two waterbird species (mallard and teal) over two continents (Europe and North America) and gut retention time of different propagules to build a simple mechanistic model of passive dispersal of aquatic plants and zooplankton. Distance and frequency of migratory movements differed both between waterbird species and continents, which in turn resulted in changes in the shapes of propagule dispersal curves. Dispersal distances and the frequency of LDD events (generated by migratory movements) were mainly determined by the disperser species and, to a lesser extent, by the continent. The gut retention time of propagules also exerted a significant effect, which was mediated by the propagule characteristics (e.g. seeds were dispersed farther than Artemia cysts). All estimated dispersal curves were skewed towards local‐scale dispersal and, although dispersal distances were lower than previous estimates based only on the vector flight speed, had fat tails produced by LDD events that ranged from 230 to 1209 km. Our results suggest that propagule dispersal curves are determined by the migratory strategy of the disperser species, the region (or flyway) through which the disperser population moves, and the propagule characteristics. Waterbirds in particular may frequently link wetlands separated by hundreds of kilometres, contributing to the maintenance of biodiversity and, given the large geographic scale of the dispersal events, to the readjustment of species distributions in the face of climate change.     

Using value of information to prioritize research needs for migratory bird management under climate change: a case study using federal land acquisition in the United States

In response to global habitat loss, many governmental and non‐governmental organizations have implemented land acquisition programs to protect critical habitats permanently for priority species. The ability of these protected areas to meet future management objectives may be compromised if the effects of climate change are not considered in acquisition decisions. Unfortunately, the effects of climate change on ecological systems are complex and plagued by uncertainty, making it difficult for organizations to prioritize research needs to improve decision‐making. Herein, we demonstrate the use of qualitative value of information analysis to identify and prioritize which sources of uncertainty should be reduced to improve land acquisition decisions to protect migratory birds in the face of climate change. The qualitative value of information analysis process involves four steps: (i) articulating alternative hypotheses; (ii) determining the magnitude of uncertainty regarding each hypothesis; (iii) evaluating the relevance of each hypothesis to acquisition decision‐making; and (iv) assessing the feasibility of reducing the uncertainty surrounding each hypothesis through research and monitoring. We demonstrate this approach using the objectives of 3 U.S. federal land acquisition programs that focus on migratory bird management. We used a comprehensive literature review, expert elicitation, and professional judgement to evaluate 11 hypotheses about the effect of climate change on migratory birds. Based on our results, we provide a list of priorities for future research and monitoring to reduce uncertainty and improve land acquisition decisions for the programs considered in our case study. Reducing uncertainty about how climate change will influence the spatial distribution of priority species and biotic homogenization were identified as the highest priorities for future research due to both the value of this information for improving land acquisition decisions and the feasibility of reducing uncertainty through research and monitoring. Research on how changes in precipitation patterns and winter severity will influence migratory bird abundance is also expected to benefit land acquisition decisions. By contrast, hypotheses about phenology and migration distance were identified as low priorities for research. By providing a rigorous and transparent approach to prioritizing research, we demonstrate that qualitative value of information is a valuable tool for prioritizing research and improving management decisions in other complex, high‐uncertainty cases where traditional quantitative value of information analysis is not possible. Given the inherent complexity of ecological systems under climate change, and the difficulty of identifying management‐relevant research priorities, we expect this approach to have wide applications within the field of natural resource management.     

Migratory herds of wildebeests and zebras indirectly affect calf survival of giraffes

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Participatory wildlife surveys in communal lands: a case study from Simanjiro,Tanzania

It is widely accepted that protected areas alone are not sufficient to conserve wildlife populations particularly for migratory or wide‐ranging species. In this study, we assess the population density of migratory species in the Tarangire–Simanjiro Ecosystem by conducting a ground census using DISTANCE sampling. We focus on the Simanjiro Plains which are used as a dispersal area by wildebeest (Connochaetes taurinus) and zebra (Equus burchellii). We demonstrate that DISTANCE sampling can provide precise estimates of population density and is an affordable method for monitoring wildlife populations over time. We stress the importance of involving local communities in monitoring programmes across landscapes that incorporate communal lands as well as protected areas.     

Sea trout Salmo trutta in the subarctic: home-bound but large variation in migratory behaviour between and within populations

Anadromous brown trout (sea trout), Salmo trutta, is currently in decline throughout its range, largely due to anthropogenic stressors in freshwater and marine habitats. Acoustic telmetry was utilized to study the marine migration of sea trout post-smolts from three populations in a relatively pristine subarctic fjord system. While at sea, the sea trout spent a substantial part of their time close to their natal river, preferred near shore over pelagic habitats and were strongly surface oriented. Despite a fidelity towards local areas, the sea trout utilized various parts of the fjord system, with maximum dispersion >30 km and total migration distance >300 km. Almost half of the sea trout (44%) migrated between river outlets, indicating that a metapopulation approach may be appropriate when managing neighbouring sea trout populations at high latitudes. Furthermore, the different populations displayed different migratory behaviours in terms of distance migrated, dispersion from origin and the likelihood of leaving their home area. This variation in migratory behaviour is likely influenced by spatiotemporal differences in habitat quality between sites, indicating that local habitat variations may promote population-specific behavioural responses even in relatively confined fjord systems.