Drivers and consequences of partial migration in an alpine bird species

1. Partial migration, where a portion of the population migrates between winter and summer (breeding) areas and the rest remain year‐round resident, is a common phenomenon across several taxonomic groups. Several hypotheses have been put forward to explain why some individuals migrate while others stay resident, as well as the fitness consequences of the different strategies. Yet, the drivers and consequences of the decision to migrate or not are poorly understood.
2. We used data from radio‐tagged female (n = 73) willow ptarmigan Lagopus lagopus in an alpine study area in Central Norway to test if (i) the decision to migrate was dependent on individual state variables (age and body weight), (ii) individuals repeated migratory decisions between seasons, and (iii) the choice of migratory strategy was related to reproductive success.
3. Partially supporting our prediction that migratory strategy depends on individual state, we found that juvenile birds with small body sizes were more likely to migrate, whereas large juveniles remained resident. For adult females, we found no relationship between the decision to migrate or stay resident and body weight. We found evidence for high individual repeatability of migratory decision between seasons. Migratory strategy did not explain variation in clutch size or nest fate among individuals, suggesting no direct influence of the chosen strategy on reproductive success.
4. Our results indicate that partial migration in willow ptarmigan is related to juvenile body weight, and that migratory behavior becomes a part of the individual life history as a fixed strategy. Nesting success was not affected by migratory strategy in our study population, but future studies should assess other traits to further test potential fitness consequences.

     

Early springs and breeding performance in two sympatric duck species with different migration strategies

The capacity of migratory species to adapt to climate change may depend on their migratory and reproductive strategies. For example, reproductive output is likely to be influenced by how well migration and nesting are timed to temporal patterns of food abundance, or by temperature variations during the brood rearing phase. Based on two decades (1988–2009) of waterfowl counts from a boreal catchment in southern Finland we assessed how variation in ice break‐up date affected nesting phenology and breeding success in two sympatric duck species, Mallard Anas platyrhynchos and Eurasian Teal Anas crecca. In Fennoscandia these species have similar breeding habitat requirements but differ in migration distance; Teal migrate roughly seven times as far as do Mallard. Annual ice break‐up date was used as a proxy of spring ‘earliness’ to test the potential effect of climate change on hatching timing and breeding performance. Both species were capable of adapting their nesting phenology, and bred earlier in years when spring was early. However, the interval from ice break‐up to hatching tended to be longer in early springs in both species, so that broods hatched relatively later than in late springs. Ice break‐up date did not appear to influence annual number of broods per pair or annual mean brood size in either species. Our study therefore does not suggest that breeding performance in Teal and Mallard is negatively affected by advancement of ice break‐up at the population level. However, both species showed a within‐season decline in brood size with increasing interval between ice break‐up and hatching. Our study therefore highlights a disparity between individuals in their capacity to adjust to ice break‐up date, late breeders having a lower breeding success than early breeders. We speculate that breeding success of both species may therefore decline should a consistent trend towards earlier springs occur.     

Single versus subdivided population strategies in breeding against an adverse genetic correlation

In advanced conifer breeding programmes, the simultaneous genetic improvement of adversely correlated traits constitutes a major challenge. Population subdivision strategies have been proposed to deal with breeding objective uncertainty, to reduce inbreeding depression in production populations and to reduce genetic correlation adversity. We used Monte Carlo simulations based on a finite locus model to study the effect of a two-breeding-population strategy applying selection for each trait in each breeding population on the genetic correlation and on genetic gains in breeding populations (BP) and the production population (PP) within a time frame of ten generations. A single-BP and a two-subline strategy both applying multitrait index selection with equal trait weights were used as references. Two BP strategy simulations indicated that simultaneous genetic gain for the two traits could be achieved in the PP despite adverse pleiotropy. The adversity of the genetic correlations decreased in BPs of the two-BP strategy, in contrast to single-BP and subline strategies, but the adversity reduction came at the cost of a lower rate of aggregated (summed) genetic gain in the PP for the two-BP strategy compared to the single-BP or subline strategies. The subline strategy exhibited increased genetic gain in the PP at equal levels of inbreeding as intended. Two BP strategies could be useful to develop breeds specialised on different traits and to simultaneously reduce adverse genetic correlations. However, if the aggregated genetic gain should be maximised, the single-BP strategy appears a better choice.     

Stopover strategies in passerine bird migration: a simulation study

Long-distance bird migration consists of a series of stopovers (for refuelling) and flights, with flights taking little time compared to stopovers. Therefore, it has been hypothesized that birds minimize the total time taken for migration through efficient stopover behaviour. Current optimality models for stopover include (1) the fixed expectation rule and (2) the global update rule. These rules maximize the speed of migration by determining the optimal departure fuel load for a given fuel deposition rate. We were interested in simple behavioural rules approaching the stopover behaviour of real birds and how these rules compare to the time minimizing models above with respect to the total time taken for migration. The simple strategies were to stay at a site (1) until a fixed fuel load was reached or (2) for a constant number of days. We simulated migration of small nocturnal passerine birds across an environment of continuously distributed but variable fuel deposition rates, and investigated the influence of different stopover strategies on the duration of migration. Staying for a constant number of days at each stopover site, irrespective of the fuel deposition rate, resulted in only slightly longer than minimum values for migration duration. Additionally, the constant stopover duration, e.g. 10 days, may change by a day or two (per stopover) without having a large effect on total migration duration. There is therefore a possibility that real birds may be close to optimal migration speed without the need for very complex behaviour. When assessing the sensitivity of migration duration to factors other than stopover duration, we found that flight costs, search and settling time, mean fuel deposition rate and the accuracy in the choice of flight direction were the factors with the largest influence. Our results suggest that migrating birds can approximate optimal stopover duration relatively easy with a simple rule, and that other factors, e.g. those above, are more relevant for travel time.     

From phenotyping towards breeding strategies: using in vivo indicator traits and genetic markers to improve meat quality in an endangered pig breed

In endangered and local pig breeds of small population sizes, production has to focus on alternative niche markets with an emphasis on specific product and meat quality traits to achieve economic competiveness. For designing breeding strategies on meat quality, an adequate performance testing scheme focussing on phenotyped selection candidates is required. For the endangered German pig breed ‘Bunte Bentheimer’ (BB), no breeding program has been designed until now, and no performance testing scheme has been implemented. For local breeds, mainly reared in small-scale production systems, a performance test based on in vivo indicator traits might be a promising alternative in order to increase genetic gain for meat quality traits. Hence, the main objective of this study was to design and evaluate breeding strategies for the improvement of meat quality within the BB breed using in vivo indicator traits and genetic markers. The in vivo indicator trait was backfat thickness measured by ultrasound (BF_iv), and genetic markers were allele variants at the ryanodine receptor 1 (RYR1) locus. In total, 1116 records of production and meat quality traits were collected, including 613 in vivo ultrasound measurements and 713 carcass and meat quality records. Additionally, 700 pigs were genotyped at the RYR1 locus. Data were used (1) to estimate genetic (co)variance components for production and meat quality traits, (2) to estimate allele substitution effects at the RYR1 locus using a selective genotyping approach and (3) to evaluate breeding strategies on meat quality by combining results from quantitative-genetic and molecular-genetic approaches. Heritability for the production trait BF_iv was 0.27, and 0.48 for backfat thickness measured on carcass. Estimated heritabilities for meat quality traits ranged from 0.14 for meat brightness to 0.78 for the intramuscular fat content (IMF). Genetic correlations between BF_iv and IMF were higher than estimates based on carcass backfat measurements (0.39 v. 0.25). The presence of the unfavorable n allele was associated with increased electric conductivity, paler meat and higher drip loss. The allele substitution effect on IMF was unfavorable, indicating lower IMF when the n allele is present. A breeding strategy including the phenotype (BF_iv) combined with genetic marker information at the RYR1 locus from the selection candidate, resulted in a 20% increase in accuracy and selection response when compared with a breeding strategy without genetic marker information.     

Informing conservation by identifying range shift patterns across breeding habitats and migration strategies

A species distribution combines the resources and climatic tolerances that allow an individual or population to persist. As these conditions change, one mechanism to maintain favorable resources is for an organism to shift its range. Much of the research examining range shifts has focused on dynamic distribution boundaries wheras the role of species breeding habitat or migration strategies on shift tendencies has received less attention. We expand on previous research by using a large suite of avian species (i.e., 277), analyzing observed abundance-weighted average latitudes, and categorizing species by breeding environment and migration strategy. We used the North American Breeding Bird Survey dataset to address two questions: (1) Has the center of observed abundance for individual species shifted latitudinally? (2) Is there a relationship between migration strategy or breeding habitat and range shifts? Results indicate the majority of species have experienced poleward range shifts over the last 43 years, and birds breeding in all habitat showed trends of poleward shift but only those species breeding in scrub-shrub and grassland environments were different from zero. Additionally, species that are short distance migrants are experiencing significant poleward shifts while Neotropical and permanent residents had shifts that were not different from zero. Our findings do support the general trend expected from climate driven changes (i.e., > 52 % shifting poleward), however, the proportion of species exhibiting equatorial shifts (24 %) or no significant shifts (23 %) illustrates the complex interplay between land cover, climate, species interactions, and other forces that can interact to influence breeding ranges over time. Regardless of the mechanisms driving range shifts, our findings emphasize the need for connecting and expanding habitats for those species experiencing range shifts. This research describes the patterns of breeding birds through central North America and we encourage future research to focus on the mechanisms driving these patterns.     

Reproduction strategies in introduced Nymphoides peltata populations revealed by genetic markers

To determine whether introduced Nymphoides peltata populations reproduced vegetatively or sexually, microsatellites were used to study the genetic structure of the species in Sweden. Leaves from 156 plants and seeds from 4 plants were sampled from 7 water-systems and analysed. A total of 10 genotypes were found among the 156 leaves. Seeds accounted for one additional genotype per seed. Lack of genetic variation and entirely vegetative reproduction dominated in the introduced N. peltata populations although sexual reproduction was found in one water-system. However, even where the species reproduced sexually, vegetative reproduction constituted an important part of the total reproduction.     

Fine-scale genetic structure and its consequence in breeding aggregations of a passerine bird

The pattern of fine-scale genetic structure in a population may reflect current biological processes of the species, such as natal dispersal, the breeding system and demography. We investigated the spatial distribution of nests and fine-scale genetic structure during two breeding seasons in a population of a weakly territorial, flock-living passerine bird, the vinous-throated parrotbill, Paradoxornis webbianus . Nest distribution was clustered. There were two peaks of egg laying within each breeding season, and spatial clustering of nests was more extreme during the second peak after controlling for breeding density. The patterns of genetic structure during the breeding season varied with parental sex and season. Genetic structure occurred during the second laying peak: males breeding within 200 m of one another at this time were significantly more closely related than males breeding farther apart. However, no apparent genetic structure was detected in males during the first laying peak or among females in either laying peak. These results reveal male-oriented kin affiliation during part of the breeding season in this species. Furthermore, juvenile recruitment into the winter flocks was positively related to this increased relatedness among males via kin affiliation. This study implies that fine-scale genetic structure during the breeding season could be a factor determining individual fitness and may play an important role in our understanding of the evolution of social systems.     

No downregulation of immune function during breeding in two year‐round breeding bird species in an equatorial East African environment

Some equatorial environments exhibit substantial within‐location variation in environmental conditions throughout the year and yet have year‐round breeding birds. This implies that breeding in such systems are potentially unrelated to the variable environmental conditions. By breeding not being influenced by environmental conditions, we become sure that any differences in immune function between breeding and non‐breeding birds do not result from environmental variation, therefore allowing for exclusion of the confounding effect of variation in environmental conditions. This create a unique opportunity to test if immune function is down‐regulated during reproduction compared to non‐breeding periods. We compared the immune function of sympatric male and female chick‐feeding and non‐breeding red‐capped Calandrella cinerea and rufous‐naped larks Mirafra africana in equatorial East Africa. These closely‐related species occupy different niches and have different breeding strategies in the same grassland habitat. Red‐capped larks prefer areas with short grass or almost bare ground, and breed during low rainfall periods. Rufous‐naped larks prefer areas of tall grass and scattered shrubs and breed during high rainfall. We measured the following immune indices: nitric oxide, haptoglobin, agglutination and lysis, and measured total monthly rain, monthly average minimum (T_min) and maximum (T_max) temperatures. Contrary to our predictions, we found no down‐regulation of immune function during breeding; breeding birds had higher nitric oxide than non‐breeding ones in both species, while the other three immune indices did not differ between breeding phases. Red‐capped larks had higher nitric oxide concentrations than Rufous‐naped larks, which in turn had higher haptoglobin levels than red‐capped larks. T_max was higher during breeding than during non‐breeding for red‐capped larks only, suggesting potential confounding effect of T_max on the comparison of immune function between breeding and non‐breeding birds for this species. Overall, we conclude that in the two year‐round breeding equatorial larks, immune function is not down‐regulated during breeding.     

Potential for an Arctic‐breeding migratory bird to adjust spring migration phenology to Arctic amplification

Arctic amplification, the accelerated climate warming in the polar regions, is causing a more rapid advancement of the onset of spring in the Arctic than in temperate regions. Consequently, the arrival of many migratory birds in the Arctic is thought to become increasingly mismatched with the onset of local spring, consequently reducing individual fitness and potentially even population levels. We used a dynamic state variable model to study whether Arctic long‐distance migrants can advance their migratory schedules under climate warming scenarios which include Arctic amplification, and whether such an advancement is constrained by fuel accumulation or the ability to anticipate climatic changes. Our model predicts that barnacle geese Branta leucopsis suffer from considerably reduced reproductive success with increasing Arctic amplification through mistimed arrival, when they cannot anticipate a more rapid progress of Arctic spring from their wintering grounds. When geese are able to anticipate a more rapid progress of Arctic spring, they are predicted to advance their spring arrival under Arctic amplification up to 44 days without any reproductive costs in terms of optimal condition or timing of breeding. Negative effects of mistimed arrival on reproduction are predicted to be somewhat mitigated by increasing summer length under warming in the Arctic, as late arriving geese can still breed successfully. We conclude that adaptation to Arctic amplification may rather be constrained by the (un)predictability of changes in the Arctic spring than by the time available for fuel accumulation. Social migrants like geese tend to have a high behavioural plasticity regarding stopover site choice and migration schedule, giving them the potential to adapt to future climate changes on their flyway.     

Remote estimation of overwintering home ranges in an elusive,migratory nocturnal bird

Due to a long running research bias toward the breeding season, there are major gaps in knowledge on the basic nonbreeding ecology of many species, preventing a full‐annual cycle focus in ecology and conservation. Exacerbating this problem is the fact that many species are extremely difficult to detect outside of breeding. Here, we demonstrate a partial solution to this problem by using archival GPS tags to examine the overwintering ecology of a migratory nocturnal bird, the eastern whip‐poor‐will (Antrostomus vociferous). We deployed tags on 21 individuals and were able to recover 11 (52%) one year later. Tags collected high precision (approx. 10 m) points throughout the nonbreeding period. With continuous time movement models, we used these data to estimate overwintering home ranges. All individuals exhibited at least one bounded home range during this phase of the annual cycle, three of eleven had two wintering locations, and home range area ranged from 0.50 to 10.85 ha. All overwintering home ranges contained closed‐canopy forest land cover (42%–100%), and no other land cover type represented >40% of any home range. We found some evidence, with caveats, that total edge within the landscape surrounding the home range was negatively related to home range area. The prevalence of contiguous closed‐canopy forest cover in overwintering home ranges contrasts with apparent breeding habitat preferences, which includes clear‐cuts and other, more open, habitats. This study is the first to reveal key aspects of overwintering space use in this species by using archival GPS to overcome both logistical and methodological limitations. Expanded use of such technology is critical to gathering basic ecological and distributional data, necessary for achieving a more complete understanding of full‐annual cycles of animal populations.     

Area-sensitivity of three reed bed bird species breeding in Mediterranean marshland fragments

Density and species-specific thresholds for three strictly reed bed (Phragmites australis)—related bird species (Ixobrychus minutus, Acrocephalus scirpaceus, Acrocephalus arundinaceus), known as area-sensitive and breeding in an archipelago of 16 Mediterranean marshland fragments (Central Italy), were studied with mapping method techniques. No individuals of these species occurred in fragments smaller than 1 ha. Marshland fragments larger than 10 ha hosted all the three study species. We observed a significant increase in mean density between 0–1 and 1–10 ha for Acrocephalus scirpaceus and between 1–10 and >10 ha for Acrocephalus arundinaceus. Habitat heterogeneity of the marshland fragments was directly correlated to their size area. Therefore, area-sensitivity for the reed bed species can be interpreted also as a heterogeneity-sensitivity effect at fragment scale. In Mediterranean landscapes, a suitable marshland for the entire selected set of reed bed area-sensitive species should have a size greater than 10 ha with a patchy mosaic of reed beds, open waters, and other edge features.     

Fine‐scale genetic structure and helping decisions in a cooperatively breeding bird

In animal societies, characteristic demographic and dispersal patterns may lead to genetic structuring of populations, generating the potential for kin selection to operate. However, even in genetically structured populations, social interactions may still require kin discrimination for cooperative behaviour to be directed towards relatives. Here, we use molecular genetics and long‐term field data to investigate genetic structure in an adult population of long‐tailed tits Aegithalos caudatus, a cooperative breeder in which helping occurs within extended kin networks, and relate this to patterns of helping with respect to kinship. Spatial autocorrelation analyses reveal fine‐scale genetic structure within our population, such that related adults of either sex are spatially clustered following natal dispersal, with relatedness among nearby males higher than that among nearby females, as predicted by observations of male‐biased philopatry. This kin structure creates opportunities for failed breeders to gain indirect fitness benefits via redirected helping, but crucially, most close neighbours of failed breeders are unrelated and help is directed towards relatives more often than expected by indiscriminate helping. These findings are consistent with the effective kin discrimination mechanism known to exist in long‐tailed tits and support models identifying kin selection as the driver of cooperation.     

Complex patterns of genetic and phenotypic divergence in an island bird and the consequences for delimiting conservation units

Substantial phenotypic and genetic variation is often found below the species level and this may be useful in quantifying biodiversity and predicting future diversification. However, relatively few studies have tested whether different aspects of intraspecific variation show congruent patterns across populations. Here, we quantify several aspects of divergence between 13 insular populations of an island endemic bird, the Vanuatu white-eye ( Zosterops flavifrons ). The components of divergence studied are mitochondrial DNA (mtDNA), nuclear DNA microsatellites and morphology. These different aspects of divergence present subtly different scenarios. For instance, an mtDNA phylogenetic tree reveals a potential cryptic species on the most southerly island in Vanuatu and considerable divergence between at least two other major phylogroups. Microsatellite loci suggest that population genetic divergence between insular populations, both between and within phylogroups, is substantial, a result that is consistent with a low level of interisland gene flow. Finally, most populations were found to be strongly morphologically divergent, but no single population was morphologically diagnosable from all others. Taken together, our results show that, although many measures of divergence are concordant in this system, the number of divergent units identified varies widely depending on the characters considered and approach used. A continuum of divergence and a degree of discordance between different characters are both to be expected under simple models of evolution, but they present problems in terms of delimiting conservation units.     

Long-term population dynamics of breeding bird species in the German Wadden Sea area

For no other group of organisms in coastal areas are there so exact and long-term data available as there are for seabirds. Since the beginning of the 20th century, documentation of population size, especially for species breeding in colonies from the groups gulls, terns and auks, is almost complete. These species act as bio-indicators, and data on fluctuations in their population size are useful as they reflect changes in the state of the marine ecosystem. The population development of some of these seabird species (Herring Gull, Guillemot, Common, Arctic and Sandwich Tern) from the German North Sea coast, which primarily feed on fish, is given. Common to all these species is an exponential increase in numbers in recent years (1970–1985). Possible causes for this development, e.g. pressure from enemies or competitors, availability of breeding places, anthropogenic stress and mortality factors, as well as the direct and indirect anthropogenic-influenced changes in the trophic system due to the increasing eutrophication of coastal waters, are evaluated. Signs of a collapse in the stocks of seabrids resulting from environmental pollution are discussed. Consequences resulting from the ecosystem changes, such as reduction of nutrient discharge into the North Sea and the expansion of biological monitoring, are described. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Social genetic and social environment effects on parental and helper care in a cooperatively breeding bird

Phenotypes expressed in a social context are not only a function of the individual, but can also be shaped by the phenotypes of social partners. These social effects may play a major role in the evolution of cooperative breeding if social partners differ in the quality of care they provide and if individual carers adjust their effort in relation to that of other carers. When applying social effects models to wild study systems, it is also important to explore sources of individual plasticity that could masquerade as social effects. We studied offspring provisioning rates of parents and helpers in a wild population of long-tailed tits Aegithalos caudatus using a quantitative genetic framework to identify these social effects and partition them into genetic, permanent environment and current environment components. Controlling for other effects, individuals were consistent in their provisioning effort at a given nest, but adjusted their effort based on who was in their social group, indicating the presence of social effects. However, these social effects differed between years and social contexts, indicating a current environment effect, rather than indicating a genetic or permanent environment effect. While this study reveals the importance of examining environmental and genetic sources of social effects, the framework we present is entirely general, enabling a greater understanding of potentially important social effects within any ecological population.     

A new westward migration route in an Asian passerine bird

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濒危物种巴东木莲的等位酶遗传多样性及其保护策略

为了确定中国特有濒危植物巴东木莲(Manglietia patungensis)的就地保护优先单元和制定迁地保护的取样策略,采用超薄平板聚丙烯酰胺等电聚焦电泳技术对巴东木莲的7个野生居群的等位酶遗传多样性及遗传结构进行了分析。通过对8个酶系统19个酶位点的分析结果来看,巴东木莲具有较高的遗传多样性和一定程度的遗传分化。每位点平均等位基因数（A）为1.57,多态位点百分率（P）为48.1%,平均预期杂合度（H_e）为0.192。巴东木莲居群间的遗传分化系数（G_ST）为0.165,说明其16.5％的遗传变异存在于居群间;基于遗传分化系数计算的基因流（N_m）为1.27,说明居群间存在适度的基因流;固定指数(F)为-0.191,显示巴东木莲居群杂合体轻微过量,纯合体略显不足。本研究表明,目前巴东木莲仍具有较高的遗传多样性并且在其适宜生境中可以完成自然更新,因此应采取以就地保护为主的综合保育策略。