Spatial Neophilia and Spatial Neophobia in Resident and Migratory Warblers (Sylvia)

Residents and migrants use their environment very differently – the former remain in a given habitat throughout the year, whereas the latter are repeatedly confronted with unfamiliar environments. The difference in ecology may influence decision-making processes whether, when and to which extent to explore an unfamiliar environment. We have investigated spatial neophobia and spatial neophilia – two important novelty reactions that may underlie decision-making – in two closely related warbler species, the resident Sardinian warbler and the migratory garden warbler. Individuals of both species could access an unfamiliar room from a familiar cage. We assessed the conflict between the motivation to enter the novel room (spatial neophilia) and the motivation to avoid it (spatial neophobia) as the frequency and duration of perching on the dowel in the cage, which led to the unfamiliar room before entering it. Furthermore, we measured the latency to enter the novel room and compared the number of individuals of each species entering the room. The combination of the parameters measured allowed assessing the degree of both neophobia and neophilia. Finally, the time spent on each branch in the novel room was taken as a measure for spatial exploration. The migrants perched less often and spent less time on the dowel leading to the room, and entered the novel room quicker than the residents. Additionally, more migrants than residents entered the room. The migrants' decision to enter the novel room can best be explained with a combination of low spatial neophobia coupled with high spatial neophilia, whereas the residents' decision-making is best explained with high spatial neophobia coupled with high spatial neophilia. The differences in neophobia support the migrant-neophobia hypothesis. When in the room, the migrants spent less time on each branch than the residents, possibly indicating that the former collect less spatial information than the latter.     

Seasonal variation of infestation by ectoparasitic chigger mite larvae (Acarina: Trombiculidae) on resident and migratory birds in coffee agroecosystems of Chiapas, Mexico

Parasitism is not well documented for birds found in tropical habitats. Long-distance migratory birds may face additional risks to an already hazardous journey when infected. This study explores the ecology of an ectoparasite infestation in Chiapas, Mexico. During a mist-netting project in 2 different coffee management systems, chigger mites (Acarina: Trombiculidae), ectoparasitic during the larval stage, were found on both resident and migratory birds. Using a rapid assessment protocol, it was observed that 17 of 26 species of long-distance migrants and 33 of 71 resident species had at least 1 infested individual. Infestation prevalences were unexpectedly high on some long-distance migrants, as high as 0.73 for Swainson's thrush (Catharus ustulatus), a value on par with heavily infested resident species. Prevalence was highest during winter sampling: 0.18 overall, 0.16 of migrants, and 0.23 of residents. Prevalence was 0.14 for resident birds during the summer breeding season. Mean abundance and mean intensity of infestation are reported for 97 species captured and inspected during the course of this study. In this region, chigger mite larvae are relatively common on birds and their abundance varies seasonally. High prevalence for some migratory birds suggests that more research and monitoring of ectoparasites are needed, especially in light of emerging diseases.     

Host immune defence and migration in birds

Migratory birds are exposed to at least two different parasite faunas during their annual cycle, while resident birds only experience a single parasite fauna. Migratory birds should therefore have evolved mechanisms to control or reduce the negative impact of infections from a more diverse parasite fauna. In a comparison of pairs of closely related species of birds that differ with respect to whether they are migratory or residents, the size of two immune defence organs (the bursa of Fabricius and the spleen) was consistently larger in the migratory species. Since the bursa is only found in juvenile, sexually immature birds, we conclude that immune defence adaptations to the impact of a more diverse parasite fauna in migrants already exist before the start of the first migration. Interspecific differences in investment in immune defence between migratory and resident birds have implications for our understanding of complex host–parasite interactions, the acquisition of new hosts by parasites, and the susceptibility of migratory birds to environmental perturbations.     

Why do some,but not all,tropical birds migrate? A comparative study of diet breadth and fruit preference

Annual migrations of birds profoundly influence terrestrial communities. However, few empirical studies examine why birds migrate, in part due to the difficulty of testing causal hypotheses in long-distance migration systems. Short-distance altitudinal migrations provide relatively tractable systems in which to test explanations for migration. Many past studies explain tropical altitudinal migration as a response to spatial and temporal variation in fruit availability. Yet this hypothesis fails to explain why some coexisting, closely-related frugivorous birds remain resident year-round. We take a mechanistic approach by proposing and evaluating two hypotheses (one based on competitive exclusion and the other based on differences in dietary specialization) to explain why some, but not all, tropical frugivores migrate. We tested predictions of these hypotheses by comparing diets, fruit preferences, and the relationships between diet and preference in closely-related pairs of migrant and resident species. Fecal samples and experimental choice trials revealed that sympatric migrants and residents differed in both their diets and fruit preferences. Migrants consumed a greater diversity of fruits and fewer arthropods than did their resident counterparts. Migrants also tended to have slightly stronger fruit preferences than residents. Most critically, diets of migrants more closely matched their preferences than did the diets of residents. These results suggest that migrants may be competitively superior foragers for fruit compared to residents (rather than vice versa), implying that current competitive interactions are unlikely to explain variation in migratory behavior among coexisting frugivores. We found some support for the dietary specialization hypothesis, propose refinements to the mechanism underlying this hypothesis, and discuss how dietary specialization might ultimately reflect past interspecific competition. We recommend that future studies quantify variation in nutritional content of tropical fruits, and determine whether frugivory is a consequence or a cause of migratory behaviour.     

Niche-tracking migrants and niche-switching residents: evolution of climatic niches in New World warblers (Parulidae)

Differences in life-history traits between tropical and temperate lineages are often attributed to differences in their climatic niche dynamics. For example, the more frequent appearance of migratory behaviour in temperate-breeding species than in species originally breeding in the tropics is believed to have resulted partly from tropical climatic stability and niche conservatism constraining tropical species from shifting their ranges. However, little is known about the patterns and processes underlying climatic niche evolution in migrant and resident animals. We evaluated the evolution of overlap in climatic niches between seasons and its relationship to migratory behaviour in the Parulidae, a family of New World passerine birds. We used ordination methods to measure seasonal niche overlap and niche breadth of 54 resident and 49 migrant species and used phylogenetic comparative methods to assess patterns of climatic niche evolution. We found that despite travelling thousands of kilometres, migrants tracked climatic conditions across the year to a greater extent than tropical residents. Migrant species had wider niches than resident species, although residents as a group occupied a wider climatic space and niches of migrants and residents overlapped extensively. Neither breeding latitude nor migratory distance explained variation among species in climatic niche overlap between seasons. Our findings support the notion that tropical species have narrower niches than temperate-breeders, but does not necessarily constrain their ability to shift or expand their geographical ranges and become migratory. Overall, the tropics may have been historically less likely to experience the suite of components that generate strong selection pressures for the evolution of migratory behaviour.     

Migratory Birds Reinforce Local Circulation of Avian Influenza Viruses

Migratory and resident hosts have been hypothesized to fulfil distinct roles in infectious disease dynamics. However, the contribution of resident and migratory hosts to wildlife infectious disease epidemiology, including that of low pathogenic avian influenza virus (LPAIV) in wild birds, has largely remained unstudied. During an autumn H3 LPAIV epizootic in free-living mallards (Anas platyrhynchos) — a partially migratory species — we identified resident and migratory host populations using stable hydrogen isotope analysis of flight feathers. We investigated the role of migratory and resident hosts separately in the introduction and maintenance of H3 LPAIV during the epizootic. To test this we analysed (i) H3 virus kinship, (ii) temporal patterns in H3 virus prevalence and shedding and (iii) H3-specific antibody prevalence in relation to host migratory strategy. We demonstrate that the H3 LPAIV strain causing the epizootic most likely originated from a single introduction, followed by local clonal expansion. The H3 LPAIV strain was genetically unrelated to H3 LPAIV detected both before and after the epizootic at the study site. During the LPAIV epizootic, migratory mallards were more often infected with H3 LPAIV than residents. Low titres of H3-specific antibodies were detected in only a few residents and migrants. Our results suggest that in this LPAIV epizootic, a single H3 virus was present in resident mallards prior to arrival of migratory mallards followed by a period of virus amplification, importantly associated with the influx of migratory mallards. Thus migrants are suggested to act as local amplifiers rather than the often suggested role as vectors importing novel strains from afar. Our study exemplifies that a multifaceted interdisciplinary approach offers promising opportunities to elucidate the role of migratory and resident hosts in infectious disease dynamics in wildlife.     

Size‐dependent costs of migration: Migrant bird species are subordinate to residents,but only at small body sizes

Migrant species are commonly thought to be poor competitors in aggressive interactions with resident species. However, no studies have tested whether this relationship is widespread. Here, we compare the behavioural dominance of closely related species of migratory and nonmigratory birds, testing whether migrants are consistently subordinate to resident species in aggressive contests. We compiled published behavioural dominance data involving migrant and resident congeners, gathering additional data on the body mass and migratory distance of each species. Focal species included a diverse array of birds (28 taxonomic families, 12 orders) from around the world. We found that migrant species are usually subordinate to resident species, but that this relationship disappears at larger body sizes. For smaller birds (<500 g), resident species were behaviourally dominant in 83%–88% of comparisons; for larger birds (>500 g), resident species were dominant in only 25%–30% of comparisons. The relative difference in body mass best predicted dominance relationships among species, with larger species dominant in 80%–84% of comparisons. When migrant and resident masses were equal, however, resident species were still more likely to be dominant in smaller birds, suggesting that other factors may also contribute to the subordinate status of migrants. Overall, our results suggest that in smaller species, the evolution of migration is associated with lighter weights and other traits that compromise the competitive abilities of migrants relative to residents. In contrast, larger species appear able to evolve migration without compromising their size or competitive abilities in aggressive contests, suggesting size‐dependent constraints on the evolution of migration.     

Impacts of passage migrant songbirds on behaviour and habitat use of resident Sardinian Warblers Sylvia melanocephala in Gibraltar

Explanations for the coexistence of migratory and sedentary (resident) birds are much debated: some authors have suggested that migrants make use of surplus resources not exploited by residents, whereas others have recorded ecological shifts in residents during periods when migrants are present. These shifts have been ascribed to the presence of migrants but could equally reflect seasonal variation in the distribution of resources required by sedentary species. Here we assess how the abundance of migrants affects the territorial behaviour and habitat use of resident Sardinian Warblers Sylvia melanocephala at a stop‐over site on the Rock of Gibraltar. There was a significant positive relationship between the abundance of migrants and the distances moved, time spent calling and the species and functional diversity of plants used by Sardinian Warblers on a daily basis, suggesting that migrants have a significant impact on residents.     

The Significance of Ecological Factors for Exploration and Neophobia in Parrots

Exploratory behaviour plays an important role in most animals for gathering information about their environment. If it constitutes an adaptation to different environmental conditions exploratory behaviour should differ between species. This has been tested with several hypotheses.
Sixty-one parrot species (Psittacidae) from eight tribes with different diets and habitat preferences were investigated in aviaries. Two tests were carried out. First, a novel object (wooden ring) in the familiar aviary was presented on two test days in the exploration test. Latencies until first contact with the object and the duration of exploration were recorded. Secondly, in the neophobia test, novel objects were placed beside the feeding dish and latencies until first food intake were recorded.
The exploration and neophobia data were related to 12 (13) ecological variables using multiple regression analyses. Phylogenetic relationships were considered.
Species that inhabit complex habitats, such as forest edges, or that feed on buds or species from islands showed the shortest latencies in the exploration test. In contrast, long latencies were related to a diet including a great amount of seeds and/or flowers. The longest duration of exploration occurred in species eating nuts or originating from islands, whereas short durations were related to feeding on seeds. Neophobia was positively related to a diet consisting of insects, and negatively to a diet of leaves. There was no relationship between measures of exploration and neophobia. Exploration and neophobia seem to be tightly related to the ecology of a species.     

Stress hormone dynamics: an adaptation to migration?

The hormone corticosterone (CORT) is an important component of a bird’s response to environmental stress, but it can also have negative effects. Therefore, birds on migration are hypothesized to have repressed stress responses (migration-modulation hypothesis). In contrast to earlier studies on long-distance migrants, we evaluate this hypothesis in a population containing both migratory and resident individuals. We use a population of partially migratory blue tits (Cyanistes caeruleus) in southern Sweden as a model species. Migrants had higher CORT levels at the time of capture than residents, indicating migratory preparations, adaptation to stressors, higher allostatic load or possibly low social status. Migrants and residents had the same stress response, thus contradicting the migration-modulation hypothesis. We suggest that migrants travelling short distances are more benefited than harmed by retaining the ability to respond to stress.     

Is the range size of migratory birds constrained by their migratory program?

Aim Intuitively, species in which the individuals migrate long distances between summer and winter quarters should be more likely to disperse and colonize new breeding areas than resident species. However, it has repeatedly been noted that many bird species with large ranges are residents. This paradox was tested on land birds breeding in the boreal forest of the Palaearctic, the largest uninterrupted stretch of habitat on earth. Methods The longitudinal distribution of two land bird communities on each side of the Eurasian continent, in Scandinavia and eastern Siberia, were used to test whether migratory birds indeed have a lower colonization success than resident birds. Results The migratory species are significantly less likely than resident species to have a range including both regions. The pattern held true even after controlling for latitudinal effects and local abundance, and was also observed at the level of genus and family. Main conclusions The relatively low colonization success of migratory species into new breeding areas may be because these new areas require novel migratory programs (migratory distance, direction and timing) in order for the birds to reach suitable wintering grounds.     

Foraging ecology of a tropical aerial feeding bird guild

The feeding ecology of the constituent members of the aerial feeding bird guild was studied in Malaya. Although numbers changed with the movements of migrants, more birds were seen feeding over forested habitats than open areas. Methods of ecological isolation were seen in morphology, flight behaviour and food selection. Seasonal changes in foraging behaviour associated with breeding and migration were found in Pacific Swallows and Barn Swallows. Migrant species were observed to mix freely with residents and no evidence was found for competitive exclusion between the two groups.     

Temperate migrants and resident bird species in Afro‐tropical savannahs show similar levels of ecological generalism

The specificity of an animal's habitat requirements will determine its ability to deal with anthropogenic climate and habitat change. Migratory birds are thought to be particularly vulnerable to such change, but theory predicts that they should be largely generalists. This prediction was tested with the aim of assessing whether migratory Palaearctic‐breeding birds wintering in the savannah biome of Africa are more or less generalist in their habitat use compared with taxonomically and ecologically similar Afro‐tropical resident species. The degree of specialization of these species groups to certain habitat characteristics was assessed and compared by calculating the relative occurrence of the species along habitat gradients, where wide occurrence indicates generalism and narrow occurrence indicates specialism. Palaearctic migrants as a group could not clearly be distinguished as generalists relative to Afro‐tropical residents with respect to habitat attributes. The only indication of greater flexibility in Palaearctic migrants was a significant tendency to use habitats over a wider latitudinal range. The results suggest that migrants are generalists, but not necessarily more generalist than taxonomically similar resident species that also occur over a wide range of habitat types within the savannah biome. The availability of specific habitat requirements on the wintering grounds in Africa is therefore unlikely to be a primary limiting factor for many Afro‐Palaearctic migratory bird species.     

The Flight Apparatus of Migratory and Sedentary Individuals of a Partially Migratory Songbird Species

Variations in the geometry of the external flight apparatus of birds are beneficial for different behaviors. Long-distance flight is less costly with more pointed wings and shorter tails; however these traits decrease maneuverability at low speeds. Selection has led to interspecific differences in these and other flight apparatuses in relation to migration distance. If these principles are general, how are the external flight apparatus within a partially migratory bird species shaped in which individuals either migrate or stay at their breeding grounds? We resolved this question by comparing the wing pointedness and tail length (relative to wing length) of migrant and resident European blackbirds (Turdus merula) breeding in the same population. We predicted that migrant blackbirds would have more pointed wings and shorter tails than residents. Contrary to our predictions, there were no differences between migrants and residents in either measure. Our results indicate that morphological differences between migrants and residents in this partially migratory population may be constrained.     

Novelty Responses in a Bird Assemblage Inhabiting an Urban Area

Neophobia, or the hesitancy to approach a novel food item, object, or place, is an important factor influencing the foraging behavior of animals. Environmental factors (e.g. rapid anthropogenic changes, migration into new habitats) are associated with novelty in feeding ecology and may affect neophobic responses. Mechanisms that underlie the differential neophobic response may involve complex interactions with the environment: post-fledging experience in a greater diversity of habitats or in habitats that are more complex may contribute to reduced neophobia. In a previous study, it was observed that some urbanized species, in particular house sparrows ( Passer domesticus ) and shiny cowbirds ( Molothrus bonariensis ) show high levels of neophobia. This study was carried out in a suburban marsh of Cortaderia selloana , a relatively simple and predictable ecosystem as compared to urban areas. For this reason, in the present study, we explored novelty responses of bird species inhabiting an urban area, representing a complex environment. The results were compared to those obtained previously in the suburban marsh. We found unexpectedly high levels of neophobia in house sparrows, but shiny cowbirds showed a somewhat neophilic response. In the presence of novel objects, house sparrows tended to enter the feeders alone, while shiny cowbirds tended to forage in groups. We found no differences in latencies to forage or in visit duration between habitat types, but the proportion of individuals that visited the feeders when novel objects were present was lower in the urban area for house sparrows and eared doves ( Zenaida auriculata ). The results are discussed in the context of invasion success and feeding innovation in shiny cowbirds.     

Migrant birds disperse haemosporidian parasites and affect their transmission in avian communities

Migration has an important impact on the transmission of pathogens. Migratory birds disperse parasites through their routes and may consequently introduce them to new areas and hosts. Hence, haemosporidian parasites, which are among the most prevalent, diverse and important bird pathogens, are potentially dispersed when infecting migrant hosts. Further, migrant hosts could enhance local parasite prevalence and richness by transporting new parasite strains to new areas. Here, we hypothesize and aim to evaluate if 1) migratory birds spread parasite lineages along their routes, and 2) localities crossed by more migratory birds have greater prevalence and richness of haemosporidians. For the first hypothesis, we tested whether parasite lineages found 1) in both migrants and residents, and 2) only in residents, differ in their frequencies of occurrence among localities. For the second hypothesis, we tested for a relationship among localities between the overall local haemosporidian parasite richness and prevalence, and the proportion of migratory bird individuals present in a locality. We combined a dataset on 13 200 bird samples with additional data from the MalAvi database (total: ~2800 sequenced parasites comprising 675 distinct lineages, from 506 host species and 156 localities) from South America, and used Bayesian multi-level models to test our hypotheses. We demonstrate that parasites shared between resident and migratory species are the most spatially widespread, highlighting the potential of migrants to carry and transmit haemosporidians. Further, the presence of migrants in a locality was negatively related to local parasite richness, but not associated with local prevalence. Here, we confirm that migrants can contribute to parasite dispersal and visiting migrants are present in regions with lower Plasmodium prevalence. Also, we observed their presence might raise Haemoproteus community prevalence. Therefore, we demonstrate migrants enhance pathogens spread and their presence may influence parasite community transmission.     

Brain size, innovative propensity and migratory behaviour in temperate Palaearctic birds

The evolution of migration in birds remains an outstanding, unresolved question in evolutionary ecology. A particularly intriguing question is why individuals in some species have been selected to migrate, whereas in other species they have been selected to be sedentary. In this paper, we suggest that this diverging selection might partially result from differences among species in the behavioural flexibility of their responses to seasonal changes in the environment. This hypothesis is supported in a comparative analysis of Palaearctic passerines. First, resident species tend to rely more on innovative feeding behaviours in winter, when food is harder to find, than in other seasons. Second, species with larger brains, relative to their body size, and a higher propensity for innovative behaviours tend to be resident, while less flexible species tend to be migratory. Residence also appears to be less likely in species that occur in more northerly regions, exploit temporally available food sources, inhabit non-buffered habitats and have smaller bodies. Yet, the role of behavioural flexibility as a response to seasonal environments is largely independent of these other factors. Therefore, species with greater foraging flexibility seem to be able to cope with seasonal environments better, while less flexible species are forced to become migratory.     

Fat reserves of migrating passerines at arrival on the breeding grounds in Swedish Lapland

Mist-net capture data taken during 6 years (1988–1990 and 1992–1994) of field work were used to describe the arrival sequences and fat loads of nine species of migratory passerines which breed in a near-Arctic environment in Swedish Lapland. Long-distance migrants arrived with significantly larger mean fat reserves than did medium- and short-distance migrants. Long-distance migrants carried fat loads at arrival which corresponded to potential remaining flight distances between 242 and 500 km. When females and males arrived on the breeding grounds simultaneously, females carried significantly larger energy reserves than did males in seven out of nine species. In contrast, when the sexes showed a significant difference in median arrival date (two out of nine species), there was no difference in mean fat load carried into the breeding area. A relationship was found between the migratory habits and foraging ecology of each species and the amount of fat reserves at arrival, suggesting that species-specific migratory distances and feeding habits determine the amount of fat that is needed during the transition period between migration and onset of breeding. The short growing season in the study area restricts the time available for breeding and moult, and large energy reserves at arrival may speed up the breeding schedule to counteract possible time constraints. Overloading at the last stopover site during spring migration may be an adaptation allowing birds to cope with a restricted time frame for breeding and moult at high latitudes.     

Does the migration programme constrain dispersal and range sizes of migratory birds?

Aim  It is generally believed that the migration programme constrains the dispersal and hence range sizes of migratory bird species. This conclusion is based on analyses of breeding ranges of migratory versus non-migratory (resident) terrestrial bird species, and rests on the assumption that there are no ecological or evolutionary constraints on extending the non-breeding range. To investigate this assumption, the abilities of migrant and resident terrestrial species to colonize new wintering areas were compared.
Location  Three major wintering regions of long-distance migrants: South America, sub-Saharan Africa and the Indian Subcontinent.
Methods  It was determined whether the relative numbers of residents and short- and long-distance migrants were the same in those species that have dispersed to a novel wintering region as in the source species pools.
Results  At the species level, long-distance migratory species are more likely to have non-breeding ranges that include more than one of the above regions than resident species. This indicates that the dispersal of migratory species is less constrained than that of resident species. The pattern holds irrespective of the inclusion or exclusion of species associated with coastal, freshwater and wetland habitats, and also holds for ecological groups such as aerial feeders. The pattern is most pronounced between the regions separated by the strongest dispersal barriers (South America and sub-Saharan Africa).
Main conclusions  It is unlikely that the migration programme per se constrains dispersal, but rather that difficulties in establishing new non-breeding areas prevent range expansions in migrant species.     

Perceived wintering latitude determines timing of song output in a migratory bird

Migratory bird populations frequently consist of individuals that overwinter variable distances from the breeding site. Seasonal changes in photoperiod, which varies with latitude, underlie seasonal changes in singing frequency in birds. Therefore, migratory populations that consist of individuals that overwinter at different latitudes with large overwintering ranges could experience within‐population variation in seasonal production of song. To test the influence of overwintering latitude on intrapopulation variance in song production in the spring, we subjected two groups of Eastern Song Sparrows (Melospiza melodia melodia) from the same partially migratory breeding population to different photoperiodic schedules associated with a 1,300‐km difference in overwintering location. One group remained on the natural photoperiodic schedule of the breeding site (resident group) while the other group experienced a nonbreeding photoperiod that mimicked a southern migration in the fall followed by a northern migration back to the breeding site in the spring (migratory group). We compared song output between the two groups in three different stages (nonbreeding, prebreeding, and breeding). Little singing occurred during nonbreeding stage sample dates (20 November, 6 December) for the resident group, and no singing occurred for the migrant group. During the prebreeding stage (27 January, 7 February), significantly more singing occurred in the resident group than in the migrant group. During the breeding stage (21 March, 4 April), after a simulated migration for the migrants, song output was similar in both groups. These results suggest that within‐population variation in wintering latitude may contribute to variation in seasonal changes in singing behavior, which may covary with readiness to breed. Studies utilizing confirmed migrants and residents, rather than merely simulated migrants and residents, are also needed to better understand these processes.