Can intensively farmed arable land be favourable for birds during migration? The case of the Eurasian golden plover Pluvialis apricaria

Today's intensive farming practices are known to have affected farmland biodiversity negatively in many different ways. As far as birds are concerned, they are known to have suffered during both summer and winter. Relatively little is known about the effects on birds during migration. We studied the stopover ecology of the Eurasian golden plover Pluvialis apricaria, a species listed in EU Birds Directive, in intensively farmed arable land in southernmost Sweden in the autumns of 2003–2007. We used key ecological variables (length of stay, fat deposition and moult) as fitness proxies to evaluate how the birds manage in this habitat. Eurasian golden plovers were present in large numbers mainly on arable fields from early August to November and radio‐tagged birds were found to stay in the area for up to three months. Adult birds carried out a substantial part of their flight feather moult during their stay. Body mass increased only somewhat during moult, but from the last stages of moult and onwards fuel loads corresponding to 24% above lean body mass (LBM) were accumulated at a rate of 0.5% of LBM per day, before the birds departed. Juveniles arrived later, from mid Sep., and had a similar pattern of fuel deposition. The fact that the birds choose to stay for long periods, moult in the area, and manage to store substantial fuel loads strongly suggests that Eurasian golden plovers do well in this intensively farmed arable land.     

Migration and moult of Dunlin Calidris alpina wintering in the central Mediterranean

Dunlin migration in northeast Italy is described. An attempt to identify the main routes and staging areas used by birds wintering in the central Mediterranean is presented. The results of monthly counts from 1990–1995 revealed that the bulk of the population occupied the wintering area in October and left for the breeding grounds in April and May. The analysis of 342 Italian recoveries of foreign ringed birds showed that 65% were ringed during post-breeding migration through the Baltic Sea, whereas just a few birds had been ringed in western Europe. First-year birds arrived in autumn with a single migratory wave, peaking in October. Two categories of adults were identified during post-breeding migration: birds which directly reached Italian wintering sites and birds which arrived after they had suspended their migration for moulting: the Azov/Black Sea wetlands are suggested as possible moulting areas. Out of 2444 adults and 1627 first-years ringed between 1989 and 1996 at our study area, we obtained a total of 42 recoveries abroad and evidence of direct links between Azov/Black Sea and N Adriatic wetlands, both during autumn and spring migrations. Primary moult was observed only in adults arriving early, the second migratory wave being composed of moulted birds. Locally moulting adults adopted a moult strategy characterized by high raggedness scores, typical of resident moulters. Body mass was not affected by primary moult stage or intensity, winter mass values being reached two weeks after the average date of primary moult completion.     

The moult of Barred Warblers Sylvia nisoria in Kenya—evidence for a split wing-moult pattern initiated during the birds' first winter*

The moult of Barred Warblers Sylvia nisoria was studied during three winter seasons in southeastern Kenya at a southward passage site (Ngulia) and a wintering site (Mtito Andei). Most Barred Warblers migrating through Ngulia in November had yet to commence winter moult. These birds probably moulted subsequently in winter in northern Tanzania. In December, birds were found in heavy moult at Mtito Andei, and some of these birds were known to stay throughout the winter. By contrast, most birds reaching southeastern Kenya from late December onwards had already completed part or all of their winter moult, presumably at stopover sites in northern and eastern Kenya or in Ethiopia. Thus, winter moult in Barred Warblers takes place mainly in late November and December, either just before or soon after the final leg of autumn migration. In general, first-year birds renewed all tertials and tail feathers, about three to five secondaries per wing and commonly also the outer one to four large primaries per wing. Adults renewed all tertials and tail feathers, almost all secondaries and only occasionally an outer primary. The replacement of relatively fresh juvenile secondaries during the birds' first winter implies that the split moult pattern of this species (secondaries, tertials and tail moulted in winter; primaries and tertials in summer) is endogenously controlled.     

The moults of captive Scottish ptarmigan (Lagopus mutus)

The moults of captive Scottish ptarmigan were studied at Banchory, north-east Scotland from December 1968 to February 1971. In the autumn moult (June to September) which included the primaries, cock ptarmigan moulted earlier and more completely than hens. In the winter moult (September to February) hens moulted earlier and both sexes moulted more completely than in spring. In the spring moult (February to June) cocks moulted more rapidly to begin with but by mid-April hens had caught up and thereafter moulted at least as rapidly as cocks. When kept indoors at slightly higher temperatures ptarmigan grew more pigmented feathers during the winter moult. In a colder winter the birds became whiter than in a milder one. First-winter ptarmigan completed the winter moult later than older birds. Birds from the Cairnwell hills had more dark feathers in winter than those from the eastern Cairngorms. There was no correlation between the start or finish of egg-laying and moulting.     

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.     

Juvenile wing shape, wing moult and weight in the family Sylviidae

A study of 56 species of Sylviidae occurring in the western Palaearctic showed a significantly less extensive post-juvenile moult in those species which incur longer migrations to winter quarters than in species that migrate short distances. Species with the more extensive post-juvenile moult also undertook their first full moult in summer compared with the less extensive post-juvenile moult in species with a full moult in the winter. In 11 species, young birds had shorter wings and lower body mass than adults. These differences produced significantly lower wing loadings in young birds compared with adults. Those species undertaking a full moult in the winter had significantly more pointed wings than species with their full moult in the summer. I suggest that this difference is the result of winter-moulting species evolving from the probably basic strategy of a full post-breeding (summer) moult. Some species, such as the Barred Warbler Sylvia nisoria , may be in the process of changing the full-moult season from summer to winter.     

Of squeezers and skippers: factors determining the age at moult of immature African Penguins Spheniscus demersus in Namibia

We used banding and resighting records of 391 African Penguins Spheniscus demersus banded as chicks and later resighted during immature moult to explain the roles of date of fledging and age at moult in determining the season of moult and its timing within the season. Breeding was continuous, but immature moult occurred mainly during spring and summer. Age at immature moult extended over 11 months, from 12 to 23 months after hatching. Birds that fledged during summer and early autumn generally moulted during the next moult season (squeezers), whereas birds that fledged in late autumn, winter and spring skipped the next moult season to moult only the following season (skippers). There was a significant relationship between age at moult and moult date, with young birds moulting later in the season than older birds. The age at moult of immature birds appears to be constrained by minimum age, moult seasonality and plumage wear. Birds that fledged over nearly 2 years moult during one season. Counts of moulting immature African Penguins have not been used to estimate year-class strength and post-fledging survival owing to the wide range of ages at immature moult. Our results provide the means of assigning recruits to specific age groups.     

Seasonal changes in moult,body mass and reproductive condition in siskins Carduelis spinus exposed to daylength regimes simulating different latitudes

Birds use change in daylength during the year to time events during their annual cycles. Individual Eurasian siskins Carduelis spinus can breed and winter in widely separated areas in different years. Birds at different latitudes will experience different changes in photoperiod. So how does latitude affect photoperiodic control? Our aim in this study was to find whether Siskins caught from the wild in Britain and exposed to different photoperiodic regimes, typical of widely separated latitudes, would differ in the subsequent timing and duration of their moults and associated processes. Siskins were caught in late February and early March, and initially kept outside on natural photoperiods. From the spring equinox (21 March), they were divided into three groups kept under photoperiodic regimes that simulated latitudes 40°, 55° and 70°N respectively. All three groups showed highly significant subsequent changes in body mass, fat scores and cloacal protuberance size. Moult of the primary feathers started during June – August (mean 9 July), and lasted 61–99 days (mean 75 days). Birds that started to moult late in the season had shorter moult durations. All individuals showed lower mass and fat levels during moult than before or after moult. Crucially, there were no significant differences in the timing of these events between the three photoperiodic groups. Apparently these birds did not use prevailing absolute photoperiod or the prevailing rate of change in photoperiod to time moult‐related seasonal events, but used instead some other feature of the annual photoperiod cycle or some form of interval timer linked to photoperiod.     

Biannual complete moult in the Black-chested Prinia Prinia flavicans

The Black-chested Prinia Prinia flavicans shows two distinctive periods each year during which adult birds undergo a complete moult: there is a fast moult (about 67 days) in spring (September-November) involving all birds simultaneously and a slower moult (about 108 days) in autumn (February-June), when about 95% of adults are moulting during April. A biannual complete moult pattern was also shown to occur in individual birds. The pattern of secondary replacement was variable and unusual for a passerine; the majority replaced S8 to S5/S4 descendantly, or had feathers being renewed ascendantly amongst S4-S7 before the ascendant series starting from the outermost secondary reached the middle secondaries. The descendant series tended to be longer during the autumn moult with S4 most frequently being the last to be replaced in autumn, but S5 last in spring. Breeding was erratic during summer in response to rains and sometimes overlapped extensively with moulting, the onset of which was less variably timed. When breeding occurred during the autumn moult, the new plumage was not the usual winter plumage (without the chest-band), but a new summer plumage.     

THE WINTERING AND MOULT OF RUFFS PHILOMACHUS PUGNAX IN THE KENYAN RIFT VALLEY

Some 5700 Ruffs were ringed in the southern Kenyan rift valley during 1967–79, mainly at Lakes Nakuru and Magadi. These have produced 15 recoveries outside East Africa, 14 in Siberia between 73° and 154°E and one in India. Adult males returned to Kenya mainly during August, and females during late August and early September. Females greatly outnumbered males at all times. Most wintering males departed late in March and early in April, but females not until about a month later. First-year birds appeared from the end of August, but remained in low numbers until late October or November. Most departed during April and May, but a few females oversummered. First-year birds typically accounted for about 25% of the wintering Nakuru females, but about 50% of those at Magadi. At both sites they accounted for a higher proportion of male birds than females. Most of the birds at Nakuru throughout late August to May appeared to be local winterers, and many individuals remained in the area for many months each year. Retrapping indicated that approximately 60% of each season's birds returned the following season. Adult males and most adult females commenced pre-winter wing moult before arrival, but completed most of it in Kenya. Males moulted 3–4 weeks ahead of females, and most had finished before December. Females typically finished during December and early January. Most second year birds timed their pre-winter moult similarly to older adults. Suspension was recorded in over 15% of all moulting birds examined. Adult pre-summer moult involved most or all of the tertials, some or all of the tail feathers, most of the inner wing coverts and the body and head plumage. It occurred mainly during January to March (males) or February to April (females), although tertial renewal commonly began a month earlier. Males showed no sign in Kenya of the supplementary prenuptial moult. First-year birds moulted from juvenile into first winter body plumage during late September to November. They underwent a pre-summer moult similar in extent and timing to that of adults, and again about a month earlier in males than females. Spring feathers acquired were often as brightly coloured as those of adults. About 15% of first-year birds renewed their outer 2–4 pairs of large primaries during January to April. Adult and first-year birds fattened before spring departure, commonly reaching weights 30–60% above winter mean. Weights of adult males peaked early in April, those of adult females early in May, and those of first-winter females later in May. Weights were relatively high also during August and September. This was due to the arrival of wintering birds carrying ‘spare’ reserves, and also apparently to the presence of a late moulting fattening passage contingent. The wing length of newly moulted adults was about 3 mm longer than that of newly arrived first-year birds, but there was no evidence of an increase in the wing kngth of adults with successive moults. Adult wing length decreased by 4–5 mm between the completion of one moult and the middle stages of the next. The migrations and annual timetable of Kenyan wintering Ruffs are discussed, and their moult strategy is compared with that of other Holarctic waders.     

The moult of the Little Stint Calidris minuta in the Kenyan rift valley

Moult data were collected during 1967–80 from some 6900 Little Stints in the southern Kenyan rift valley.
Adults typically moulted from summer to winter body and head plumage during September and early October, soon after arrival. The complete pre-winter wing and tail moult began in most adults between mid-September and early October. Some birds finished by December, but others continued until February and March. Individual duration was usually between 100 and 150 days. Adults which completed this moult early often remoulted outer primaries between January and early April.
Young birds acquired first-winter body plumage during October and early November. Some 90% had a complete pre-winter wing and tail moult. This usually began between December and early February, and finished during March or early April, taking about 70–100 days. In about 10% of young birds, flight feather moult was restricted to the outer primaries and inner secondaries. Birds adopting this strategy typically began moult late, during January or February. Short periods of suspension were common during pre-winter wing moult, particularly in adults. The difference in moult speed between adult arid first-winter birds was attributable in the primary, secondary and tail tracts to differences in numbers of growing feathers.
Practically all birds completed a pre-summer moult involving the entire body and head plumage, most of the tertials, some or all of the tail feathers and many wing coverts. Most birds began this moult between early February and late March, and finished between mid-April and early May. It was typically later and more rapid in first-year birds than adults. In late birds, the onset of pre-summer moult was linked to the final stages of pre-winter moult.
The wing moult of the Little Stint in different wintering areas is discussed. First-winter moult strategy is compared with that in other small Calidris species.     

The winter fattening model: a test at low latitude using the Clamorous Reed Warbler

Small birds at high latitudes accumulate fat during the day so that they can survive long and cold winter nights. The winter fattening model suggests that birds increase their minimum (morning) mass in cold weather, build up their mass during the day, and then rely on the energy reserves so accumulated until the morning. While data from mid and high latitudes support this model, little is known about the strategies of birds inhabiting lower latitudes (< 40°N). We use an 18-year data set to investigate whether the winter fattening model holds in a mid-latitude (32°N) population of the Clamorous Reed Warbler Acrocephalus stentoreus . We show that morning mass and fat score vary as predicted by the winter fattening model. In addition, adults appear to be better adapted to long and cold winter nights than first-year birds. A long-term trend of increased body mass may be attributable to character release following reduced interspecific competition.     

Timing and duration of moult in three populations of Purple Sandpipers Calidris maritima with different moult/migration patterns

Timing and duration of primary moult in three populations of Purple Sandpipers Calidris maritima were described and discussed in relation to the birds’ need to complete moult before the onset of winter, when resources are required for survival. We predicted that moult would be completed earlier by birds wintering at higher latitudes. The south Norwegian breeding population, which moults and winters along the coast of east Britain (54–57°N) had a mean starting date of 21 July for primary moult (16 July for females and 24 July for males), a mean duration of 61 days, and completed on 20 September. Resident Icelandic (64–65°N) birds had a mean starting date of 22 July for primary moult (17 July for females and 25 July for males), a mean duration of 51 days, and completed on 11 September. Birds moulting in north Norway (70°N) arrived in north Norway in suspended primary moult or without having started moult, and completed it there. They had a mean completion date of 2 November for primary moult (31 October for females and 3 November for males). Starting date and duration could not be estimated because some suspended moult for an undetermined period, but it was thought that they started in late August. It is likely that most originated from Russia. The onset of moult appears to be set by the end of breeding and there is little overlap in these two events. The earlier start of moult by females in all three populations may be because they abandon the males when the chicks hatch, leaving the males to attend the chicks. Although the duration of primary moult followed the expected trend, being fastest in north Norway and slowest in Britain, the onset of moult was so late in north Norway that they had an unexpectedly late completion date, despite their rapid moult. The late completion of primary moult in north Norway suggests that wintering in the far north may not pose the energetic constraints on Purple Sandpipers that had previously been supposed.     

The effects of delaying the start of moult on the duration of moult, primary feather growth rates and feather mass in Common Starlings Sturnus vulgaris

In many species of birds there is a close relationship between the end of breeding and the start of moult. Late-breeding birds therefore often start to moult late, but then moult more rapidly. This is an adaptive mechanism mediated by decreasing day lengths that allows late-breeding birds to complete moult in time. This study asked how these birds complete moult of the primary feathers more rapidly, and the consequences of this on the mass of primary feathers. Common Starlings Sturnus vulgaris were induced to moult rapidly in one of two ways. In the first experiment, one group was exposed to artificially decreasing photoperiods from the start of moult, whereas the control group remained on a constant long photoperiod. The second experiment was a more realistic simulation. Two groups were allowed to moult in an outdoor aviary. One group started to moult at the normal time. In the other, the start of moult was delayed by 3 weeks with an implant of testosterone. The duration of moult was significantly reduced in both the group experiencing artificially decreasing photoperiods and the group in which the start of moult was delayed. The faster moult rate was achieved by moulting more feathers concurrently. The rate of increase in length of each of the primary feathers, and their final length, did not differ between groups. The rate at which total new primary feather mass was accumulated was greater in more rapidly moulting birds, but this was insufficient to compensate for the greater numbers of feathers being grown concurrently. Consequently, the rate of increase in mass of individual feathers, and the final feather mass, were less in the rapidly moulting birds. A 3-week delay in the start of moult is not an unrealistic scenario. That this caused a measurable decrease in feather mass suggests that late-breeding birds are indeed likely to suffer a real decrease in the quality of plumage grown during the subsequent moult.     

A RINGING STUDY OF THE MIGRATORY BROWN SHRIKE IN WEST MALAYSIA

Wintering Brown Shrikes frequent open lowland country, a habitat that is largely man-made in Malaya. Individuals are sedentary during the winter season, each occupying a restricted area and exhibiting territorial behaviour. The earliest observations each year in different parts of Southeast Asia indicate that the southward migratory journey is relatively rapid. In Malaya, migrants arrive from the first week of September to the third week of October. At a lowland netting station during 1964–68 a major part of the total catch was taken in the months of September and October. Only a small proportion of these early shrikes wintered in the netting area. No distant recoveries were reported, and the subsequent movements of birds that were not retrapped are unknown. Shrikes netted in September—October comprised 29% adults, 54% full grown (i.e. immatures plus poorly—characterized adults), and 17% juveniles. The mean wing-length was significantly longer among adults than among both other classes, which did not differ significantly. During the winter, all ages showed a progressive decline in wing-length until the flight feathers were renewed in a premigratory moult falling in February—early April. Moult recorded in four Brown Shrikes taken in October-November is interpreted as the completion of a post-nuptial moult, commenced on the breeding grounds before autumn migration. In April, after the premigratory moult, confirmed adults constituted 70% of the total trapped and apparent immatures 30%. The mean weight of September birds was lower than any other month except November. The low weight in November is partly correlated with the shorter mean wing-length of the sample; it may also reflect the seasonally unfavourable weather of this month. The mean weight in February was high, although all birds were moulting; the weather in this month is typically hot and dry. Highest weights were recorded in April, indicating the premigratory deposition of fat. Weights of birds trapped more than once at different intervals showed a small initial weight loss (2 g), followed by a recovery within four days and no long-term adverse effects. A comparison of September weights in Taiwan and Malaya provides a tentative basis for the calculation of fat reserves utilized on the migratory flight. The proportion of returns after one year was 11%, and after two years 1 % only. Most returning birds were present in the netting area during the latter part of the winter of initial ringing; it is suggested tentatively that imprinting of the wintering grounds may occur during this period. Ecologically in Malaya the Brown Shrike occupies a new habitat only gradually being filled by the resident Rufous-backed Shrike. There is no evidence of interaction between the two species.     

Primary moult, body mass and migration of Grey Plovers Pluvialis squatarola in Britain

Long-distance migrants have evolved complex strategies for the timing of their annual moult, fattening and migration cycles. These strategies are likely to vary at different stages of a bird's life. Ringing data on 6079 Grey Plovers Pluvialis squatarola , caught on the Wash, England, between 1959 and 1996, were analysed to relate migratory strategies to patterns of primary moult and body mass changes. Adults returning from breeding grounds had a shorter and delayed primary moult (duration 90 days, starting date 19 August) in comparison with over-summering birds (duration 109 days, starting date 5 June). Three categories of migrant adults were identified on the basis of primary moult and body mass: (1) birds which did not moult, but increased body mass and migrated further south; (2) birds which moulted 1–3 inner primaries, suspended moult, increased body mass and migrated; and (3) birds which completed or suspended moult and wintered locally. In birds of the second category, timing of primary moult and body mass increase overlapped. Among wintering birds, 38% were in suspended moult. Ninety-six per cent of birds that suspended moult at the beginning of winter were males and almost all completed moult in spring. Grey Plovers which left Britain in autumn had an average body mass of 280 g, enough to reach southern Morocco without refuelling. Both wintering adults and first-year birds showed a prewinter body mass increase, peaking in December. Adults had a synchronized premigratory body mass increase in May, which suggested a negligible presence of African migrants. The average departure mass for spring migration, estimated at 316 g, would allow birds to fly non-stop to the Siberian breeding grounds in western Taymyr.     

Late‐moulting Black‐necked Grebes Podiceps nigricollis show greater body mass in the face of failing food supply

From August to December, thousands of Black‐necked Grebes Podiceps nigricollis concentrate during the flightless moult period in salt ponds in the Odiel Marshes, southern Spain, where they feed on the brine shrimp Artemia parthenogenetica. We predicted that because Black‐necked Grebes moulted in a food‐rich, predator‐free environment, there would be no net loss of body mass caused by the use of fat stored to meet energy needs during remigial feather replacement (as is the case for some other diving waterbirds). However, because the food resource disappears in winter, we predicted that grebes moulting later in the season would put on more body mass prior to moult because of the increasing risk of an Artemia population crash before the moult period is completed. Body mass determinations of thousands of birds captured during 2000–2010 showed that grebes in active wing‐moult showed greater mass with date of capture. Early‐moulting grebes were significantly lighter at all stages than late‐moulting birds. Grebes captured with new feathers post‐moult were significantly lighter than those in moult. This is the first study to support the hypothesis that individual waterbirds adopt different strategies in body mass accumulation according to timing of moult: early‐season grebes were able to acquire an excess of energy over expenditure and accumulate fat stores while moulting. Delayed moulters acquired greater fat stores in advance of moult to contribute to energy expenditure for feather replacement and retained extra stores later, most likely as a bet hedge against the increasing probability of failing food supply and higher thermoregulatory demands late in the season. An alternative hypothesis, that mass change is affected by a trophically transmitted cestode using brine shrimps as an intermediate host and Black‐necked Grebes as final host, was not supported by the data.     

DISTRIBUTION,MIGRATION AND TIMING OF MOULT IN THE SOUTH AFRICAN CLIFF SWALLOW

Earle, R. A. 1987. Distribution, migration and timing of moult in the South African Cliff Swallow. Ostrich 58:118-121. The South African Cliff Swallow Hirundo spilodera breeds in South Africa mainly between 25 and 31S and 24 and 31E. In some years with exceptionally high rainfall the breeding range is more extensive. Man has probably had a pronounced influence on the present-day distribution of this species. Seven winter recovery/collecting localities are known from the lower Congo basin in Zaire. Possible migrating birds were observed in Zambia and Malawi in the east, and on the Namibian coast in the west. The few sight records suggest a direct migrational route over Botswana. Moult takes place between March and September primarily in the winter quarters, but about 2% of the birds handled during March and April started moult before migrating.     

Body weight, gonad development and moult in the Tawny owl (Strix alum)

Gonad development, moult and seasonal changes in body weight and composition in the Tawny owl Strix aluco were studied by examining the carcasses of 369 owls (mostly road casualties) supplemented by 112 weights of live birds. In breeding females laying was preceded by the accumulation of fat and to a lesser extent protein which meant that they weighed more at this time (February/March) than at any other. Females declined in weight after laying but were still heavy during incubation. In contrast, males and non-breeding females did not increase in weight before the start of the breeding season. Juveniles reached or even exceeded adult weight well before independence due to the deposition of fat. Even after the exclusion of diseased or contaminated individuals, 9·4% of the birds examined were identified as starving; most of these were in the autumn and were probably newly-independent young wandering in search of territories. In both sexes gonad maturation was of brief duration coinciding with the period (mid-March to mid-April) in which eggs are normally laid. Ovarian growth was biphasic. In the three months prior to the breeding season ovarian condition in different birds was positively correlated with body weight and it appeared that the largest ovarian follicles of females in poor condition failed to attain the size from which rapid growth to final ovulation occurs. in males testis size in the breeding season was correlated with pectoral muscle weight (an index to protein condition) but not body weight. The majority of adults commenced wing moult in June. The average duration of primary moult was estimated to be 77 days. Healthy birds replaced the primaries of both wings at the same rate but most diseased birds moulted asymmetrically and/or out of season. First-year birds renewed their body feathers between September and November. In the Tawny owl territory establishment, breeding and moult are temporally separated.     

DISPERSION, POPULATION ECOLOGY AND MIGRATION OF EASTERN GREAT REED WARBLERS ACROCEPHALUS ORIENTALIS WINTERING IN MALAYSIA

A population of 400–600 Acrocephalus orientalis wintering in a Phragmites habitat at 3°N in West Malaysia was studied during four northern hemisphere winters, by means of systematic mist-netting. Data from other study-areas, other habitats and other winters are also used. Intensive mist-netting appears to have made birds move over longer distances than they did in the absence of disturbance, and to have led to the emigration of marked birds from the study-area. Trapping also affected feeding behaviour, resulting in weight-loss; repeated trapping may have increased mortality. Males and females could be separated by means of wing-length in fresh plumage. Females were largely confined to Phragmites; males were more numerous on the edge of reed-beds and in scrub vegetation. Males suffered greater feather-wear than females. As measured by the trapping rate, birds were uniformly distributed throughout the Phragmites habitat, at the same density in different winters. Undisturbed birds used a “home-range” of 1–4 ha, overlapping with 15–50 other individuals. Disturbed birds overlapped with 100–200 others. Individual birds returned to exactly the same “home-range” in successive winters. After correcting for the effects of disturbance and incomplete sampling, the proportion of adults ringed in one winter which returned in the next is estimated as 65% in each of two study-areas. This is a minimum estimate of the annual survival rate for adults. Mean total body-weights were at a minimum in midwinter (November-February). Fat-free weights were also lower in midwinter than in autumn and spring. Body-moult was observed in March and April. Moult of the flight-feathers takes place between July and September, on the breeding grounds or slightly to the south. Females departed on spring migration between 10 and 25 May; males some 11–14 days earlier. Adults arrived in autumn between 8 September and 7 October; males and females often came in in separate “waves”. Females were absent for only about 127 days, about the minimum required for migration, breeding and moult. Dates of migration match those of the more northern breeding populations. Spring departure is later than dates of passage recorded in south China; hence birds of this population appear to make long nights. On average, birds departing in spring carried about 9 g of fat, roughly 40% of total fat-free body-weight. This is about half the energy reserve required for the entire journey. Dates of passage in central China are consistent with a hypothesis that they make the journey (4,500-5,000 km) in two “hops”. A few birds which remained light until very late in the spring showed a significantly lower return rate in the next year. Most birds arriving in autumn appear to have carried 1–2 g of fat, but some were at or below the normal fat-free weight. Many birds appear to have lost weight soon after arrival. Returning ringed adults were amongst the very first birds trapped in September. Individual birds appear to have migrated on very similar dates in different years: many of the dates of trapping differed by 2 days or less in successive years. Trapping rates reached a peak in early October and then declined rapidly, reaching the midwinter level by 21 October. The decline coincided with the differential disappearance of juvenile birds. However, birds collected at this time had adequate fat reserves, and the disappearance appears to have preceded the period of food-shortage. It is suggested that the loss of juvenile birds resulted from behavioural interactions favouring the more dominant individuals, as has been described for several temperate zone residents. The first few weeks in the wintering area may thus be the critical period of mortality during the year. Because birds from different breeding areas are expected to be mixed in the winter-quarters, and vice versa, local mortality factors in winter may affect a number of breeding populations. High adult survival rates have been recorded in several other birds which breed in the temperate zones and winter in the tropics. In general their breeding success appears to be high, so the first-year mortality must be high. The closely related A. arundinaceus, which winters in Africa, differs from A. orientalis in size, wing-shape, timing of spring migration and timing of moult. These differences can be interpreted as adaptations to different environmental (primarily climatic) factors experienced during migration and on the breeding grounds. The segregation of males and females into different habitats probably reduces inter-sexual competition in winter, but this is not necessarily its primary function. Males collected in the evening in Phragmites had smaller fat reserves than females, suggesting that the females are better adapted to this habitat. The large size of the males is probably maintained in part by sexual selection in the breeding season. On the other hand, the size of females and their habitat is probably limited by the specialisation of their nest. These factors would suffice to explain the sexual dimorphism in size and habitat.