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

Summary 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.