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.     

Sexual segregation in winter, spring dispersal and habitat use in the pheasant (Phasianus colchicus)

Sexual segregation, spring dispersal and late winter to spring habitat selection were studied in the pheasant Phasianus colchicus L. on two areas for which the proportions of different habitats were similar. On one of the sites (Damerham) hand-reared pheasants were released for shooting. On the other (Fulwell) no releasing or shooting took place.
The sex ratio of birds feeding in fields in autumn favoured males because females fed in groups within woods. Groups of birds were significantly segregated by sex during early to late winter periods, but by late March they were not. Male groups and solitary males in winter were more evenly dispersed than female groups.
At Fulwell, female winter groups consisted of regular group members (mainly adults) and transitory individuals called 'drifters' (mainly immatures).
Females in their first breeding attempt dispersed further from their winter range to their breeding range than did adult females and females dispersed further than males. Males gaining a territory for the first time moved significantly further than old territory owners. Territorial males moved less than non-territorial males between successive spring ranges.
During March June, habitat use by 33 radio-tagged females changed markedly from mixed and deciduous woodland to fields sown with winter and spring corn. The implications for management are outlined.     

Winter climate affects subsequent breeding success of common eiders

The phenology of spring migration depends on the severity of the preceding winter and approaching spring. This severity can be quantified using the North Atlantic Oscillation (NAO) index; positive values indicate mild winters. Although milder winters are correlated with earlier migration in many birds in temperate regions, few studies have addressed how climate‐induced variation in spring arrival relates to breeding success. In northern Europe, the NAO‐index correlates with ice cover and timing of ice break‐up of the Baltic Sea. Ice cover plays an important role for breeding waterfowl, since the timing of ice break‐up constrains both spring arrival and onset of breeding. We studied the effects of the winter‐NAO‐index and timing of ice break‐up on spring migration, laying date, clutch size, female body condition at hatching and fledging success of a short‐distance migrant common eider (Somateria mollissima) population from SW Finland, the Baltic Sea, 1991–2004 (migration data 1979–2004). We also examined the correlation between the NAO‐index and the proportion of juvenile eiders in the Danish hunting bag, which reflects the breeding success on a larger spatial scale. The body condition of breeding females and proportion of juveniles in the hunting bag showed significant positive correlations with the NAO, whereas arrival dates showed positive correlations and clutch size and fledging success showed negative correlations with the timing of ice break‐up. The results suggest that climate, which also affects ice conditions, has an important effect on the fledging success of eiders. Outbreaks of duckling disease epidemics may be the primary mechanism underlying this effect. Eider females are in poorer condition after severe winters and cannot allocate as much resources to breeding, which may impair the immune defense of ducklings. Global climate warming is expected to increase the future breeding success of eiders in our study population.     

Carry‐over effects of winter habitat quality on en route timing and condition of a migratory passerine during spring migration

We examined how conditions prior to migration influenced migration performance of two breeding populations of black‐and‐white warblers Mniotilta varia by linking information on the migrant's winter habitat quality, measured via stable carbon isotopes, with information on their breeding destination, measured via stable hydrogen isotopes. The quality of winter habitat strongly influenced the timing of migration when we accounted for differential timing of migration between breeding populations. Among birds migrating to the same breeding destination, males and females arriving early to the stopover site originated from more mesic habitat than later arriving birds, suggesting that the benefits of occupying high‐quality mesic habitat during the winter positively influence the timing of migration. However, male warblers arriving early to the stopover site were not in better migratory condition than later arriving conspecifics that originated from poor‐quality xeric winter habitat, regardless of breeding destination. The two breeding populations stopover at the study site during different time periods, suggesting that the lower migratory condition of early birds is not a function of the time of season, but potentially a migrant's migration strategy. Strong selection pressures to arrive early on the breeding grounds to secure high‐quality breeding territories may drive males from high‐quality winter habitat to minimize time at the expense of energy. This migration strategy would result in a smaller margin of safety to buffer the effects of adverse weather or scarcity of food, increasing the risk of mortality. The migratory condition of females was the same regardless of the timing of migration or breeding destination, suggesting that females adopt a strategy that conserves energy during migration. This study fills an important gap in our understanding of the linkages between winter habitat quality and factors that influence the performance of migration, the phase of the annual cycle thought to be limiting most migratory bird populations.     

Annual-Cycle Movements and Phenology of Black Scoters in Eastern North America

Sea ducks exhibit complex movement patterns throughout their annual cycle; most species use distinct molting and staging sites during migration and disjunct breeding and wintering sites. Although research on black scoters (Melanitta americana) has investigated movements and habitat selection during winter, little is known about their annual-cycle movements. We used satellite telemetry to identify individual variation in migratory routes and breeding areas for black scoters wintering along the Atlantic Coast, to assess migratory connectivity among wintering, staging, breeding, and molt sites, and to examine effects of breeding site attendance on movement patterns and phenology. Black scoters occupied wintering areas from Canadian Maritime provinces to the southeastern United States. Males used an average of 2.5 distinct winter areas compared to 1.1 areas for females, and within-winter movements averaged 1,256 km/individual. Individuals used an average of 2.1 staging sites during the 45-day pre-breeding migration period, and almost all were detected in the Gulf of St. Lawrence. Males spent less time at breeding sites and departed them earlier than females. During post-breeding migration, females took approximately 25 fewer days than males to migrate from breeding sites to molt and staging sites, and then wintering areas. Most individuals used molt sites in James and Hudson bays before migrating directly to coastal wintering sites, which took approximately 11 days and covered 1,524 km. Males tended to arrive at wintering areas 10 days earlier than females. Individuals wintering near one another did not breed closer together than expected by chance, suggesting weak spatial structuring of the Atlantic population. Females exhibited greater fidelity (4.5 km) to previously used breeding sites compared to males (60 km). A substantial number of birds bred west of Hudson Bay in the Barrenlands, suggesting this area is used more widely than believed previously. Hudson and James bays provided key habitat for black scoters that winter along the Atlantic Coast, with most individuals residing for >30% of their annual cycle in these bays. Relative to other species of sea duck along the Atlantic Coast, the Atlantic population of black scoter is more dispersed and mobile during winter but is more concentrated during migration. These results could have implications for future survey efforts designed to assess population trends of black scoters. © 2021 The Wildlife Society.     

Examining carry‐over effects of winter habitat on breeding phenology and reproductive success in prairie warblers Setophaga discolor

Winter habitat quality can influence breeding phenology and reproductive success of migratory birds. Using stable isotope ratios of carbon (δ¹³C) from bird claws and red blood cells collected in Massachusetts, USA, we assessed if winter habitat occupancy carried over to affect prairie warbler Setophaga discolor breeding arrival dates, body condition upon arrival, pairing success, first‐egg dates and reproductive success. In two of three years (in 2011 and 2012, but not in 2013), after‐second‐year (ASY) males wintering in drier habitat, as indicated by enriched δ¹³C values, arrived later on the breeding grounds. Based on the North Atlantic Oscillation index, there was likely less rainfall in the Caribbean wintering grounds during the winters of 2011 and 2012 compared to the winter of 2013, suggesting increased winter rainfall in 2013 may have diminished the influence of winter habitat occupancy on arrival date. We did not find any effects of winter habitat on breeding season phenomena for second‐year (SY) males or females, but our sample sizes for these age/sex classes were relatively low. Although winter habitat quality influenced arrival dates of ASY males, there was no evidence that it affected reproductive performance, perhaps because of high rates of nest depredation in our system. Our study adds to a growing body of research that shows the influence of carry‐over effects can differ among species and within populations, and also can be modulated by other environmental conditions. This information enriches our understanding of the role of carry‐over effects in population limitation for migratory birds.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

Satellite tracking of Swedish Ospreys Pandion haliaetus: autumn migration routes and orientation

Autumn migration routes and orientation of Swedish Ospreys Pandion haliaetus were studied by satellite tracking of 18 birds. Of these, 13 could be followed during the entire migration (6 females, 5 males and 2 juveniles). Most birds migrated across western and central Europe to winter in tropical West Africa. However, one juvenile flew to Cameroon and one female used a very easterly route and reached Mozambique. On average, the birds travelled a total distance of about 6700 km, with little variation except for the female wintering in Mozambique, who travelled more than 10 000 km. Of 21 stopovers (of >1 day), only five were made south of 45°N; three of these in Africa. Females departed before males and juveniles and flew to a stopover site they probably were familiar with. After 3–4 weeks there, they continued to their wintering grounds. Also males and juveniles usually made one or more stopovers. Adults seemed to travel to a known wintering site, where they remained stationary, whereas juveniles were more mobile after reaching tropical regions, probably looking for good wintering sites. Males generally left the breeding area in directions similar to the mean migratory direction, whereas a few females departed in diverging initial directions. Apart from these diversions, adult Ospreys followed very straight migratory routes, with overall mean directions of 185–209° and with mean angular deviations of 6–33°. Some juveniles also departed in diverging directions. Moreover, young birds tended to show a larger variability in orientation. Thus, the Ospreys kept a fairly straight direction and did not avoid geographical obstacles such as mountain ranges and desert areas. However, they seemed reluctant to cross large water bodies. There was no correlation between angular deviation and length of the migrational segment, indicating that the principles of orientation by vector summation may not be valid for Osprey migration. Moreover, the geographic direction of migration did not vary in accordance with variations in the magnetic declination, suggesting that the Ospreys did not orient along magnetic loxodromes.     

Body Condition of Wintering Pacific Greater White-Fronted Geese

Extreme changes to key waterfowl habitats in the Klamath Basin (KB) on the Oregon–California border and the Sacramento Valley (SV) in California, USA, have occurred since 1980. The spatial distribution of Pacific greater white-fronted geese (Anser albifrons sponsa; geese) has likewise changed among these areas and population size has grown from 79,000 to >600,000 geese during the same period. To assess the effects of landscape changes and spatial-temporal distribution of geese, we collected Pacific greater white-fronted geese during winters of 2009–2010 and 2010–2011 in the KB and SV and compared their body condition to geese collected during 1979–1980 and 1980–1981. We modeled body and lipid mass to assess body condition for each sex independently and examined the influence of collection day, year, and region. Body condition of geese varied throughout the winter and within years in a nonlinear fashion. We detected an increase in body condition in both sexes during December and January in the SV, which corresponds with improved habitat conditions and increases seen in other species in the region. Body condition upon arrival in fall migration varied by year for females and by year and region for males. Males and females arrived in poorer body condition during 2010–2011 than all other study years and males in the KB during 2010–2011 had extremely low lipid mass, reflecting poor regional habitat conditions induced by drought. Body condition of females varied over spring, by year, and by region and regional effects were evident for males. Body condition was significantly higher for geese in the SV than in the KB during spring. Our results suggest that Pacific greater white-fronted geese have adapted to a changing landscape and have adjusted historical spatial use patterns to take advantage of more favorable conditions in the SV between 1979 and 2010. © 2021 The Wildlife Society.     

Central European Greylag Geese Anser anser show a shortening of migration distance and earlier spring arrival over 60 years

Global climate change can cause pronounced changes in species? migratory behaviour. Numerous recent studies have demonstrated climate‐driven changes in migration distance and spring arrival date in waterbirds, but detailed studies based on long‐term records of individual recapture or re‐sighting events are scarce. Using re‐sighting data from 430 marked individuals spanning a 60‐year period (winters 1956/1957 to 2015/2016), we assessed patterns in migration distance and spring arrival date, wintering‐site fidelity and survival in the increasing central European breeding population of Greylag Geese Anser anser. We demonstrate a long‐term decrease in migration distance, changes in the wintering range caused by winter partial short‐stopping, and the earlier arrival of geese on their breeding grounds. Greylag Geese marked on central Europe moulting grounds have not been recorded wintering in Spain since 1986 or in Tunisia and Algeria since 2004. The migration distance and spring arrival of geese indicated an effect of temperature at the breeding site and values of the NAO index. Greylag Geese migrate shorter distances and arrive earlier in milder winters. We suggest that shifts in the migratory behaviour of Central European Greylag Geese are individual temperature‐dependent decisions to take advantage of wintering grounds becoming more favourable closer to their breeding grounds, allowing birds to acquire breeding territories earlier.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

Site tenacity and survival rates of Wrens Troglodytes troglodytes and Treecreepers Certhia familiaris in a Nottinghamshire wood

Mark-recapture data from 20 years of standardized mist-netting were used to investigate site tenacity and adult survival rates of Wrens Troglodytes troglodytes and Treecreepers Certhia familiaris in an English woodland. Within breeding seasons, a high proportion of Wren (but not Treecreeper) recaptures occurred in the same part of the wood as the original capture. Between breeding seasons, a high proportion of recaptures of adult males (but not of adult females) occurred in the same part of the wood as the original capture. Both species experienced exceptionally high mortality during the severe winters of 1978–1979 and 1985–1986, and, in the case of Wren, all or almost all birds which had previously bred in the wood probably died. Outside of these two cold winters, apparent (minimum) adult survival rates of Wrens and Treecreepers showed very different temporal patterns of variation. Wren survival rates were negatively correlated with the number of snow days in winter, whilst Treecreeper survival was negatively correlated with the amount of winter rainfall. After allowing for the influence of winter weather, apparent survival rates of adult Wrens were density dependent, indicating that either mortality or permanent emigration out of the wood was density dependent. There was some evidence from British and Irish ringing recoveries that movements of adult Wrens may be density dependent. The apparent survival rate of adult Wrens was probably an important factor determining the size of the local breeding population.     

Can Experience Alter the Avian Annual Cycle? Results of Migration Experiments with Indigo Buntings

Field experiments were conducted during 2 years testing whether premature release on their familiar, perennial breeding territories would cause indigo buntings (Passerina cyanea) to delete spring migration from their annual cycle. In the first year, male buntings were captured on their breeding territories in summer and held indoors for the winter on the photoregime that they would have encountered in nature as fall migrants and winter residents. The following spring they were released at the time buntings begin spring migration from the subtropics. Half were released on their breeding territories, to which they would normally have returned; half were released far to the south. Some of the males released on their territories remained and passed into breeding condition without having migrated, while some of those released to the south returned north to their territories. The results suggest that endogenous or photoperiodic mechanisms controlling the timing and occurrence of migration in this species can be overridden by experience. In the second year we tested alternative explanations for these results. Our second year's results seem to indicate (1) that birds that deleted migration in year one did not do so because of unfavourable weather and (2) that the birds that returned to their breeding sites in year one were actually migrating and not simply retracing their outward journey.     

Population structure,biometrics and moult of migrant Purple Sandpipers Calidris maritima in southwest Iceland in spring

Capsule Iceland is a stop‐over site for a population of Purple Sandpipers that winter in Britain. Here, they accumulate fuel loads for onward migration along with birds that have wintered in Iceland.

Aims To establish whether Purple Sandpipers from Britain stop‐over in Iceland during spring migration and, if so, to describe their population structure, changes in mass and moult.

Methods Purple Sandpipers were cannon‐netted on the coast of the Reykjanes Peninsula in southwest Iceland during May 2003 and 2005. Birds were aged, sexed (some by DNA) and standard biometric measurements made. Active body moult was scored.

Results Bill and wing lengths showed that the Purple Sandpipers we caught were similar to one of the populations that winter in Britain rather than Icelandic breeding birds. There were more males than females throughout the migration period (63% males for first‐year‐birds and 67% for adult birds). Accounting for a bias due to a higher percentage of males in a less usual habitat (muddy/sandy bays), the values for rocky sites were 52% males for first‐year birds and 62% for adults. The percentage of first‐year birds was 19% in 2003 and 32% in 2005, though the latter figure was biased by catches in muddy/sandy bays where there was a higher percentage of young birds. The percentage of first‐year birds was 25% on just the rocky shores in 2005. Many birds were in latter stages of body moult, and males were slightly in advance of females. Increasing mass showed that they were preparing for onward migration. The average increase of 0.58 g per day was similar to the rate measured in Orkney at an earlier point on the migration route. However, a high turnover of birds could be the reason for these low values. By late May, and close to the assumed departure date, the Purple Sandpipers of the different age/sex classes had fuel indices of 24–29% (33–42% of the lean mass). This was lower than that for the high Arctic sandpipers (Knots and Sanderlings) leaving southwest Iceland for Greenland and Canada.

Conclusions Our study confirmed that Purple Sandpipers do stop‐over in Iceland, and the possible lower rate of fuel accumulation and smaller amount stored, compared with Knots and Sanderlings, suggests a different migration pattern.     

Field ecology of a brood-caring dung beetleKheper nigroaeneus-habitat predictability and life history strategy

Summary The life history strategy of the dung beetleKheper nigroaeneus was studied in Mkuzi Game Reserve, South Africa.K. nigroaeneus, a large ball-rolling dung beetle, brood-cares a single offspring per nesting occasion. Adults emerged from the ground following the first spring rain (≥17 mm) in October or November, occasionally September, and trap catches were at a maximum 1–2 weeks later. Females outnumbered males by 3:1 at the time of maximum catches, but thereafter declined steadily, as the numbers of females involved in brood-care increased. Brood-care lasted 12 weeks, and parental females then (a) entered a second brood-care period, or (b) remained underground in diapause, or (c) emerged to feed. Reproductive activity stopped after February, ensuring that the entire population reached the adult stage by winter. Females active above ground after February, in reproductive diapause, had low fat levels and resorbed oocytes. Females buried underground, in reproductive diapause prior to and during winter, had high fat levels, resorbed oocytes, a reduced metabolic rate and empty guts. The Mkuzi habitat is predictable forK. nigroaeneus in spring, since only one good fall of rain is sufficient to drive population events for the next three months. In the latter half of the season the life history ofK. nigroaeneus is more flexible, and hence less constrained by climate.     

Hatching date influences winter habitat occupancy: Examining seasonal interactions across the full annual cycle in a migratory songbird

Birds experience a sequence of critical events during their life cycle, and past events can subsequently determine future performance via carry‐over effects. Events during the non‐breeding season may influence breeding season phenology or productivity. Less is understood about how events during the breeding season affect individuals subsequently in their life cycle. Using stable carbon isotopes, we examined carry‐over effects throughout the annual cycle of prairie warblers (Setophaga discolor), a declining Nearctic–Neotropical migratory passerine bird. In drier winters, juvenile males that hatched earlier at our study site in Massachusetts, USA, occupied wetter, better‐quality winter habitat in the Caribbean, as indicated by depleted carbon isotope signatures. For juveniles that were sampled again as adults, repeatability in isotope signatures indicated similar winter habitat occupancy across years. Thus, hatching date of juvenile males appears to influence lifetime winter habitat occupancy. For adult males, reproductive success did not carry over to influence winter habitat occupancy. We did not find temporally consecutive “domino” effects across the annual cycle (breeding to wintering to breeding) or interseasonal, intergenerational effects. Our finding that a male''s hatching date can have a lasting effect on winter habitat occupancy represents an important contribution to our understanding of seasonal interactions in migratory birds.     

THE MIGRATION PATTERNS OF THE PURPLE SANDPIPER CALIDRIS MARITIMA

Summers, R. W. 1994. The migration patterns of the Purple Sandpiper Calidris maritima. Ostrich 65: 167–173.

The Purple Sandpiper breeds largely in the Arctic, and winters (boreal season) on the rocky shores of the north Atlantic, further north than any other sandpiper. As the populations from Canada, Greenland, Iceland, Svalbard, Norway and Russia differ in wing and bill lengths it is possible to match measurements taken from breeding birds with samples of birds caught in winter. Ringing recoveries, especially from colour marked birds, have also helped to determine migration routes and wintering areas. Four populations move to the nearest ice-free coast. Two populations move south of the nearest ice-free coast, being replaced by larger birds from a more northerly population (“chain migration”). Only the north Canadian population is believed to migrate a long distance, “leap-frogging” other winter populations. These patterns are discussed in relation to theories for the migration patterns of waders.     

The apparent rôle of temperature in breeding initiation and winter population structure in Hyale nilssoni Rathke (Amphipoda): Field observations 1972–1983

Data are presented on the population structure of Hyale nilssoni Rathke (an amphipod inhabitant of high littoral seaweeds) over the period of breeding initiation (late February) of 11 yr. By sampling, as near as possible, the same date annually, photoperiodic effects are controlled. The percentage of ovigerous females on this date was significantly correlated with post-winter solstice mean sea temperature. February sex ratios (M : F) did not correlate with either preceding August or September mean sea temperature such as would have been expected was sex determination temperature sensitive. Sex ratio, however, was significantly positively related to mean winter sea temperatures, with warmer winters having male-dominated Hyale populations in late February. Behavioural hypotheses involving amphipod migration and/or size specific elimination of larger males (c.f. females) during harsher winters are offered in explanation. Population size-frequency distributions over 11 yr were remarkably consistent. Male size was more heterogeneous between years than female size. Mean male size (late February) proved to be significantly negatively correlated with mean winter sea temperatures. Mean female size showed no significant relationship. Given that large males are necessary for precopulatory carrying behaviour it is important that, as shown, they are available to initiate spring breeding, even after harsh winters.     

Breeding and movements of goshawks in boreal forests in Sweden

Goshawks were studied in their boreal forest breeding habitat in central Sweden from 1977 to 1982. Nests were regularly spaced, with a nearest neighbour distance of 6.3 km. corresponding to a density of about 3 pairs 100 km⁻². Of initiated breedings. 68% were successful. There was no difference in breeding performance between years. Average brood size was 2.7. Sex ratio among first captures of adults was even, but males were recaptured more often than females. Nearly all juveniles left the area in winter. Among radio-tagged adults 20% of the males, and 50% of the females moved out of the area. When moving, both adults and juveniles moved into agricultural areas, the juveniles generally covering longer distances. From autumn to spring, body condition of females improved, whereas that of the males deteriorated. It is argued that the males' stronger tendency to stay in winter is due to the advantage of possessing a territory in early spring. Females, on the other hand, can move to areas with as good food conditions as possible.     

Family-based winter territoriality in western bluebirds, Sialia mexicana: the structure and dynamics of winter groups

Winter residency is characteristic of the majority of cooperatively breeding birds, but the composition and dynamics of winter groups have been examined in relatively few. In 1996-1998, we examined winter territoriality in the western bluebird, a year-round resident that shows a limited degree of helping behaviour in central coastal California, U.S.A. In spring, most western bluebirds breed as socially monogamous pairs, but a small proportion of pairs (3-16%) have additional breeding-age males helping at the nest, usually assisting parents or brothers. We found that year-round residents commonly wintered in family groups that defended territories similar to those used in spring. Winter groups had an even sex ratio and formed early in the autumn, when hatch-year birds dispersed. More females than males left their natal groups to be replaced by an influx of immigrant hatch-year birds. Winter groups typically consisted of breeders and one or two sons from the prior breeding season along with one or more immigrant females. A second period of dispersal occurred in spring when winter groups broke up and most birds other than the breeding pair left the winter territory. When they bred, yearling males and females often bred with unrelated individuals from their winter groups. Sons were more likely to remain on the study area as yearlings when they wintered with both parents than when they wintered with just one parent. We suggest that young males stay the winter due to benefits of remaining in family groups on mistletoe-based winter territories. Subsequent localized dispersal of sons then leads to opportunistic kin-based interactions later in life. Copyright 2001 The Association for the Study of Animal Behaviour.