Partial migration in birds: tests of three hypotheses in a tropical lekking frugivore

1. Partially migratory species provide opportunities to understand which ecological factors cause some animals to migrate when others remain resident year round. Partial migration in birds has been explained by the dominance, arrival-time, and body-size hypotheses. 2. Testing these hypotheses has proven difficult due to the similarities of the predictions they make in temperate-breeding long-distance migrants. In tropical altitudinal migrants, however, these hypotheses make different predictions regarding the sex, age, and condition of migrants and residents. 3. Among white-ruffed manakins in Costa Rica, young birds were not more likely to migrate (as predicted by the dominance hypothesis), nor were females more likely to migrate (as predicted by the arrival-time hypothesis). All condition-related variables interacted with sex, together explaining much of the variation in migratory behaviour. 4. I re-articulate the body-size hypothesis in the context of tropical altitudinal bird migration, focusing explicitly on how limited foraging opportunities and differences in individual condition affect fasting ability during torrential rains. Despite ample food, the smallest birds or those stressed by parasites or moult may risk starvation at breeding elevations due to a reduction in foraging time. These results highlight how intrinsic and extrinsic factors may interact to produce observed patterns of within- and among-species variation in migratory behaviour.     

Sex differences in settlement behaviour and condition of chiffchaffs Phylloscopus collybita at a wintering site in Portugal. Are females doing better?

In passerine birds, males are generally larger and dominant over females. In line with the dominance theory, in all known differential migrant passerines, females migrate further than males. However, there are alternative explanations to the dominance hypothesis, including the specialisation hypothesis, predicting that females should do better than males in habitats and/or regions to which they are specially adapted, and where they predominate in numbers. Few studies have aimed at comparing the behaviour and condition of males and females wintering in locations largely dominated by female birds. We studied a partly nomadic species, the common chiffchaff Phylloscopus collybita, at a wintering site in Portugal. Most individuals caught at this site were transients. The only consistent and relatively powerful predictor of settlement behaviour was sex. Almost all the birds that settled in the study area were female, while there was an almost even sex ratio in the transient sample. Females had higher muscle and post-juvenile moult scores when first captured. In addition, there was a significant sex-based difference in mass trajectories, with females maintaining body mass and males losing body mass when settled at the study site. Results from this study suggest that the poor performance of males was either due to inferior male competitive ability in a region/habitat where females predominate and/or it resulted from the fact that males that reach these wintering grounds are of inferior individual quality.     

Sexual size dimorphism and timing of spring migration in birds

Sexually selected traits are limited by selection against those traits in other fitness components, such as survival. Thus, sexual selection favouring large size in males should be balanced by higher mortality of larger males. However, evidence from red-winged blackbirds (Agelaius phoeniceus) indicates that large males survive better than small males. A survival advantage to large size could result from males migrating north in early spring, when harsh weather favours large size for energetic reasons. From this hypothesis we predicted that, among species, sex differences in body size should be correlated with sex differences in timing of spring migration. The earlier males migrate relative to females, the larger they should be relative to females. We tested this prediction using a comparative analysis of data collected from 30 species of passerine birds captured on migration. After controlling for social mating system, we found that sexual size dimorphism and difference in arrival dates of males and females were significantly positively correlated. This result is consistent with the hypothesis that selection for survival ability promotes sexual size dimorphism (SSD), rather than opposes SSD as is the conventional view. If both natural selection and sexual selection favour large adult males, then limits to male size must be imposed before males become adults.     

The effects of hatching date on timing of autumn migration in partial migrants – an individual approach

Timing of spring migration and breeding and their interaction with climate change has been widely studied in recent years, but the possible changes in timing of autumn migration have gained less attention. This work focuses on autumn migration and provides the first multi‐species individual‐based study of how hatching date affects the autumn migration date and migration age by using nestling ring data and re‐trappings of the same individuals during the autumn migration at the Hanko Bird Observatory, Finland. We studied three potentially multibrooded passerines (great tit, blue tit and coal tit) and two single‐brooded birds of prey (goshawk, sparrowhawk), all partially migratory short‐distance migrants. Individuals from late broods migrated at a younger age in all tit species and also in hawks the late hatched individuals tended to migrate at a younger age than the early‐hatched individuals. Late‐hatched individuals migrated later than early‐hatched individuals in blue and coal tits, where the latest hatchers represented second brood individuals. Based on our results, the time from hatching to autumn migration is not constant even among individuals of the same population. Our study indicates that climate warming induced advancement of avian breeding may cause changes in the timing of autumn migration through the frequency of second broods.     

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.     

Selective migration from Samoa: a longitudinal study of pre-migration differences in social and psychological characteristics

In 1981 extensive questionnaire and interview data were collected on some 100 young Samoan adults. Five years later in 1986 we determined their whereabouts and divided the data in accordance with migration status. The answers of the 35 who had migrated in the intervening period were contrasted to those 65 who remained in Samoa. The migrants differed in several distinct areas. Migrants reported a higher degree of peer-reliance as a personal adaptive strategy. Migrants also reported larger numbers of individuals in social support networks, a higher quality of support and more community involvement. They also report less expressive display of anger. Those who did not migrate reported a slightly better view of life in Samoa and abroad, as well as better relations with their friends and neighbors. These findings support a hypothesis that migrants are pre-selected to fit into migrant communities and do not appear to be misfits who are unhappy with life in Samoa.     

A comparative study on the evolution of reversed size dimorphism in monogamous waders

Sexual dimorphism in size is common in birds. Males are usually larger than females, although in some taxa reversed size dimorphism (RSD) predominates. Whilst direct dimorphism is attributed to sexual selection in males giving greater reproductive access to females, the evolutionary causes of RSD are still unclear. Four different hypotheses could explain the evolution of RSD in monogamous birds: (1) The ‘energy storing’ hypothesis suggests that larger females could accumulate more reserves at wintering or refuelling areas to enable an earlier start to egg laying. (2) According to the ‘incubation ability’ hypothesis, RSD has evolved because large females can incubate more efficiently than small ones. (3) The ‘parental role division’ hypothesis suggests that RSD in monogamous waders has evolved in species with parental role division and uniparental male care of the chicks. It is based on the assumption that small male size facilitates food acquisition in terrestrial habitats where chick rearing takes place and that larger females can accumulate more reserves for egg laying in coastal sites. (3) The ‘display agility’ hypothesis suggests that small males perform better in acrobatic displays presumably involved in mate choice and so RSD may have evolved due to female preference for agile males. I tested these hypotheses in monogamous waders using several comparative methods. Given the current knowledge of the phylogeny of this group, the evolutionary history of waders seems only compatible with the hypothesis that RSD has evolved as an adaptation for increasing display performance in males. In addition, the analysis of wing shape showed that males of species with acrobatic flight displays had wings with higher aspect ratio (wing span/²wing area) than non-acrobatic species, which probably increases flight manoeuvrability during acrobatic displays. In species with acrobatic displays males also had a higher aspect ratio than females although no sexual difference was found in non-acrobatic species. These results suggest that acrobatic flight displays could have produced changes in the morphology of some species and suggest the existence of selection favouring higher manoeuvrability in species with acrobatic flight displays. This supports the validity of the mechanisms proposed by the ‘display agility’ hypothesis to explain the evolution of RSD in waders.     

Correlates of alternative migratory strategies in western bluebirds

Partial migration occurs when only some animals in a population migrate. While evidence suggests that migratory strategies are partially controlled by genes, individual and environmental conditions which alter the cost‐benefit trade‐off of migration among individuals are also likely to play a role. Three hypotheses have been advanced to explain condition‐dependent partial migration: the arrival time, dominance and body size hypotheses. In this study, we asked whether these hypotheses explained differences in migratory strategy among individuals in a partially migratory population of western bluebirds Sialia mexicana breeding in southern British Columbia, Canada. We used stable hydrogen isotope signatures in claw tissue to determine migratory strategy of individual bluebirds, and examined patterns of migration at both individual and population levels. The proportion of resident bluebirds varied significantly over the three years of the study, and across study sites. Several migrants switched to the resident strategy between years; however, we found no evidence of strategy switching in the opposite direction. Young birds were significantly more likely to be resident than older birds, a pattern which could arise if early arrival is particularly important for birds obtaining a territory for the first time. Furthermore, young females were the most likely of all sex–age classes to be resident, which may reflect a survival advantage of residency for young females. Finally, birds mated assortatively by migratory strategy and isotopic evidence suggests that members of a pair often wintered in the same place. Our results provided no support for the dominance or body size hypotheses, and only limited support for the arrival time hypothesis in bluebirds. However, taken together, we suggest that our findings indicate that social factors may influence migratory strategies in this system.     

Brood desertion in Kentish Plover Charadrius alexandrinus: an experimental test of parental quality and remating opportunities

Uniparental male care combined with polyandry is rare in birds, and the best known examples are in shorebirds Charadrii. There are two current hypotheses explaining why males care for the brood, whereas females desert and remate: either males are more capable than females at providing uniparental care (“parental quality hypothesis”) or females gain a greater increase in reproductive success by deserting than do males (“remating opportunity hypothesis”). I experimentally tested both hypotheses in Kentish Plover Charadrius alexandrinus, one of the few avian species in which either parent may desert the brood. By experimentally removing one parent when the chicks hatched, I found that male-tended broods had better survival than female-tended ones, particularly up to 6 days after hatching. It is unlikely that differential brood mortality was caused by chilling of the chicks, since the brooding behaviour of males and females was not different. The results of this study are consistent with the explanation that male-tended broods survived better because males were better able to protect the brood from attacks by conspecifics and predators. The remating opportunity hypothesis was also corroborated because single females acquired new mates faster than did single males. The results of this study suggest that both the better parental capability of males and the greater remating opportunities of females predispose Kentish Plovers for uniparental male care, desertion by the female parent and sequential polyandry.     

Selective migration from Samoa: A longitudinal study of pre‐migration differences in social and psychological characteristics

Abstract

In 1981 extensive questionnaire and interview data were collected on some 100 young Samoan adults. Five years later in 1986 we determined their whereabouts and divided the data in accordance with migration status. The answers of the 35 who had migrated in the intervening period were contrasted to those 65 who remained in Samoa. The migrants differed in several distinct areas. Migrants reported a higher degree of peer‐reliance as a personal adaptive strategy. Migrants also reported larger numbers of individuals in social support networks, a higher quality of support and more community involvement. They also report less expressive display of anger. Those who did not migrate reported a slightly better view of life in Samoa and abroad, as well as better relations with their friends and neighbors. These findings support a hypothesis that migrants are pre‐selected to fit into migrant communities and do not appear to be misfits who are unhappy with life in Samoa.     

Should females of migratory dabbling ducks switch mates between wintering and breeding sites?

Pairing of northern hemisphere dabbling ducks normally occurs in wintering sites. Mate choice occurs when some individuals congregate in courtship parties, in which females mainly choose mates according to male behavioural dominance. The sex ratio of some species is more male-biased at northern than at southern wintering sites, and the age ratio is adult-biased in northern areas. A possible mechanism responsible for this spatial segregation of sex and age classes is behavioural dominance, with males usually dominating females and adults dominating yearlings. Due to this latitudinal segregation of sexes during winter, presumably more males would remain unpaired in northern sites utilized by the more dominant birds than in southern sites utilized by the less-dominant birds, and this is paradoxical. I argue that some females having wintered and paired in southern sites may switch mates at stopover sites during northward migration. This hypothesis provides an explanation as to why, in spite of females being paired when initiating northward migration, some species show an increase in courtship activity during spring at stopover sites, as in common teals (Anas crecca) in southern Spain. In contrast, populations that do not migrate, e.g. mallards (A. platyrhynchos) in southern Spain, do not exhibit an increase in courtship activity during spring.     

Migration of wild and captive‐bred Little Bustards Tetrax tetrax: releasing birds from Spain threatens attempts to conserve declining French populations

Populations of the Little Bustard Tetrax tetrax in the farmlands of Europe have declined greatly over the last century. In Western Europe, France now holds the only remaining migratory population, which currently numbers fewer than 300 displaying males. However, the movements of these birds are virtually unknown, in spite of the important implications of this information for the conservation of this species. We identified migratory movements and overwintering areas of French migratory populations, using wild individuals fitted with satellite or radio‐transmitters. Little Bustards completed their migration journey over a relatively short time period (2–5 days), with nocturnal migration flights of 400–600 km per night. All birds overwintered in Iberia. In addition, we tested the consequences of captive rearing on migratory movements. French wild adults and French captive‐bred juveniles behaved similarly with regard to migration, suggesting that hand‐raising does not alter migratory movements. However, birds originating from eggs collected in Spain and reared in western France did not migrate, suggesting a genetic component to migratory behaviour. These results therefore suggest that a conservation strategy involving the release in France of birds hatched from eggs collected in Spain may imperil the expression of migratory movements of the French population. More generally, to maintain the integrity of native populations, the introduced individuals should mimic their migratory movements and behaviour.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.     

FISHER'S SEX RATIO THEORY MAY EXPLAIN HATCHING PATTERNS IN BIRDS

I present the hypothesis that asynchronous hatching is a means of ensuring an equal degree of parental investment in the progeny of each sex in altricial species of birds that are sexually dimorphic in size. In a comparative analysis of bird species of Africa and the Western Palearctic, I find a positive relationship between hatching asynchrony and sexual size dimorphism, in support of the hypothesis. The relation is significant for species in which males are larger than females, and in species in which females are larger than males. In addition, it holds even if allometric effects of body size are controlled for. No such relationship is found in species with self-feeding young. Alternative hypotheses to explain asynchronous hatching in altricial birds are discussed. The results of the comparative study are also consistent with some of these hypotheses. For instance, asynchronous hatching may be a mechanism used by parents of dimorphic species to deal with unpredictable primary sex ratios; it may be a way of avoiding simultaneous peak food demands by the young; or it may be a way of advancing the time of hatching so that the division of labor between the parents is optimized.     

Evolution of prey holding behaviour and large male body size in Ninox owls (Strigidae)

The hawk owl genus Ninox is unique among raptorial birds in that it includes three species in which males are substantially larger than females. This is a reversal of the normal pattern observed in both diurnal and nocturnal raptorial birds in which females are larger. Interestingly, these three Ninox species also are both the largest of the 22 species in the genus and the only species that exhibit the striking behaviour of ‘prey holding’ in which large (> 600 g) mammalian or avian prey is captured at night and held with body parts intact, and draped below a roost for the entire day without being consumed. Because explanations of the evolution of large male size suggest that it results from competition among males, the adaptive significance of prey holding was studied in a wild population of powerful owl Ninox strenua. Prey holding is largely confined to breeding males and its occurrence varies significantly across the breeding cycle, being most frequent during incubation and brooding. The study did not clearly resolve whether prey holding is a form of food storage or territorial display; however, both functions can select for large male body size and therefore play a significant role in the evolution of nonreversed size dimorphism. Although female‐only incubation and brooding is typical of Ninox owls and other owl species, prey holding appears to occur only in the large Ninox species because of the unique combination of large body size, large prey size, separate sex roles, and obligate cavity nesting. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 284–292.     

Sex‐related variation in migration phenology in relation to sexual dimorphism: a test of competing hypotheses for the evolution of protandry

Timing of arrival/emergence to the breeding grounds is under contrasting natural and sexual selection pressures. Because of differences in sex roles and physiology, the balance between these pressures on either sex may differ, leading to earlier male (protandry) or female (protogyny) arrival. We test several competing hypotheses for the evolution of protandry using migration data for 22 bird species, including for the first time several monochromatic ones where sexual selection is supposedly less intense. Across species, protandry positively covaried with sexual size dimorphism but not with dichromatism. Within species, there was weak evidence that males migrate earlier because, being larger, they are less susceptible to adverse conditions. Our results do not support the ‘rank advantage’ and the ‘differential susceptibility’ hypotheses, nor the ‘mate opportunity’ hypothesis, which predicts covariation of protandry with dichromatism. Conversely, they are compatible with ‘mate choice’ arguments, whereby females use condition‐dependent arrival date to assess mate quality.     

Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast

The blackcap Sylvia atricapilla shows a complex migratory pattern and is a suitable species for the studies of morphological migratory syndrome, including adaptations of wing shape to different migratory performance. Obligate migrants of this species that breed in northern, central, and Eastern Europe differ by migration distance and some cover shorter distance to the wintering grounds in the southern part of Europe/North Africa or the British Isles, although others migrate to sub-Saharan Africa. Based on ˃40 years of ringing data on blackcaps captured during autumn migration in the Southern Baltic region, we studied age- and sex-related correlations in wing pointedness and wing length of obligate blackcap migrants to understand the differences in migratory behavior of this species. Even though the recoveries of blackcaps were scarce, we reported some evidence that individuals which differ in migration distance differed also in wing length. We found that wing pointedness significantly increased with an increasing wing length of migrating birds, and adults had longer and more pointed wings than juvenile birds. This indicates stronger antipredator adaptation in juvenile blackcaps than selection on flight efficiency, which is particularly important during migration. Moreover, we documented more pronounced differences in wing length between adult and juvenile males and females. Such differences in wing length may enhance a faster speed of adult male blackcaps along the spring migration route and may be adaptive when taking into account climatic effects, which favor earlier arrival from migration to the breeding grounds.     

Differential Migration by Sex in the Great Bustard: Possible Consequences of an Extreme Sexual Size Dimorphism

We explored migration patterns in Great bustards ( Otis tarda ), a species that shows strong sexual selection and the most extreme sexual size dimorphism among birds. The aim was to explain differential migration, examining whether Great bustards fulfil the main predictions of bird migration theory hypotheses and sexual segregation theory hypotheses. We radio-tracked the seasonal movements of 65 males and 68 females in central Spain. We found four main sexual differences. First, the proportion of migratory males was higher than that of migratory females (86% vs. 51%). Second, males abandoned the leks immediately after the mating season (late May to early Jun.), whereas females remained there for another 3–7 mo. Third, 54% of the migratory males used two different post-breeding areas, the first located northwards at 82 km from the breeding sites in summer, and the second southwards at 50 km in autumn–winter. Migratory females used only one area in autumn–winter which coincided geographically with that of males. And fourth, males returned to the breeding areas earlier (between Sep. and Mar.) than females (between Jan. and Apr.). These results show that the Great bustard is a differential migrant by sex in central Spain and support the weather sensitivity hypothesis (males were less tolerant to summer heat) and the specialization hypothesis (exclusive maternal care of the brood by females). Sexual differences in migratory behaviour are probably ultimately determined by the strong sexual selection in this species.     

Intrasexual mounting in the beetle Diaprepes abbreviatus (L.).

The weevil Diaprepes abbreviatus shows three kinds of same-sex mountings: males mount other unpaired males, males mount males already engaged in copulation and females mount other females. Four hypotheses were evaluated in order to explain same-sex matings by males: (i) female mimicry by inferior males, (ii) dominance of larger males which affects the behaviour of small males, (iii) sperm transfer in which smaller males gain some reproductive success by 'hitchhiking' their sperm with the sperm of larger males, and (iv) poor sex recognition. Data from mate choice and sperm competition experiments rejected the female mimicry, dominance and sperm transfer hypotheses and supported the poor sex recognition hypothesis. We tested three hypotheses in order to explain female mounting behaviour: (i) females mimic male behaviour in order to reduce sexual harassment by males, (ii) females mount other females in order to appear larger and thereby attract more and larger males for mating, and (iii) female mimicry of males. The results of our mate choice experiments suggested that the female mimicry of males hypothesis best explains the observed female mounting behaviour. This result is also consistent with the poor sex recognition hypothesis which is the most likely explanation for male and female intrasexual mating behaviour in many insect species.     

ABNORMAL MIGRATION AND HYBRIDIZATION OF LARUS ARGENTATUS AND L. FUSCUS AFTER INTERSPECIES FOSTERING EXPERIMENTS

Between 1962 and 1966 eggs of Larus argentatus and L. fuscus were interchanged andalmost 900 young were reared by the wrong species. Many of these cross-fostered young were later recovered or retrapped on and away from the colonies.
The British population of argentatus is sedentary and ringed birds are not recovered outside Britain, whereas fuscus normally migrates. Many cross-fostered argentatus migrated to France, Spain and Portugal, areas where fuscus is common, but they did not migrate as far as the control fuscus . It is possible that these cross-fostered argentatus had followed their foster parents when these migrated, but this is unlikely as the cross-fostered fuscus also migrated although their foster parents would have remained in Britain.
Despite wide ecological and behavioural overlaps, interbreeding between L. argentatus and L.fuscus is exceedingly rare. However, as a result of cross-fostering experiments, 31 and 40 mixed pairs were found on Skokholm in 1968 and 1969 respectively. Although some of the birds involved were unringed it is probable that all the adults in mixed pairs had been cross-fostered. Other cross-fostered birds were found mated with their own species and it appears that the sex of the imprinted birds was important. Female gulls will usually only mate with males of their own species, or in the case of the cross-fostered birds, with males of their foster species. Males will mate with either species.
Evidence is given that suggests that the colour of the mantle and wings is important in species recognition at long range, and the colour of eye-ring and join of the mandibles for recognition at short range. The role of voice is uncertain but general behaviour is probably unimportant.