Relationship between breeding and wintering ranges in Palaearctic-African migrants

During the northern winter, Palaearctic migrant species are not evenly distributed within sub-Saharan Africa. Species numbers are greatest in a belt of savannah, lying south of the Sahara, and decline southwards. For any one latitude, species numbers are also greater in the east of Africa than in the west. Only about 3% of 187 species winter exclusively south of the equator, but other species migrate from north to south during the course of the northern winter.
For 62 Palaearctic species which winter entirely in Africa, the areas of breeding and wintering ranges are strongly correlated. With some exceptions, species with the largest breeding ranges also have the largest wintering ranges. However, in 69% of species, the breeding range is larger than the wintering range, whereas in 31% of species the wintering range is larger. On average, the wintering ranges of 57 landbird species cover about two-thirds the area of their breeding ranges, and in many species only parts of the wintering range may be occupied at any one time. This implies that the per area carrying capacity of African wintering areas is greater than that of Eurasian breeding areas.
The general correlation between the sizes of breeding and wintering ranges may have its basis in ecology, with generalists able to occupy wider areas than specialists in both breeding and winter quarters. At the same time, the correlation may result partly from an effect of numbers on range size, in that species which have a wide range at one time of the year may then achieve large numbers which occupy a wide range at the other time of year.     

SPRING WEIGHTS OF SOME PALAEARCTIC MIGRANTS AT LAKE CHAD

A visit was made to Malamfatori, on the western shore of Lake Chad, Nigeria, from 22 March to 13 April 1967, with the principal aim of studying Palaearctic migrants in relation to the environment. About 2,400 Palaearctic migrants of 29 species were mist-netted in beds of bulrush Typha australis and thickets of saltbush Salvadora persica. Some 300 of these were collected for fat analysis; the remainder were released after measurement and ringing, and provided 275 retraps during the course of the study. Data were supplemented by further netting by A. J. Hopson in late April and May. Yellow Wagtails Motacilla flava, Sedge Warblers Acrocephalus schoenobaenus and Whitethroats Sylvia communis were abundant and were studied in greater detail than other species. Yellow Wagtails fed almost exclusively on midges, particularly the abundant Tanytarsus spadiceonotatus. Sedge Warblers fed on small insects and spiders, and Whitethroats on Salvadora fruits. Sedge Warblers and some other chiefly insectivorous species turned to a diet including berries shortly before emigrating. Changes in weight during the course of the day were difficult to investigate, but were probably bimodal. Good correlations of weights with wing-lengths were obtained for species for which many data were available. Pre-migratory fattening did not occur synchronously in all populations of a species, but once it started in an individual it proceeded at a constant rate of 0.2 g/day in Sedge Warblers and 0.6 g/day in Whitethroats. Predation pressure probably ensured that individuals emigrated immediately they attained their maximum pre-migratory weight, although some Sedge Warblers and Whitethroats left the area before attaining maximum weight. Differences between the mean weight curves of first-caught and retrapped birds are discussed, and it is concluded that in some species there were both through-migrant and temporarily-resident populations at Malamfatori. The mean weights per day of Whitethroats, Sedge Warblers and also Reed Warblers A. scirpaceus were falling in late March, and slight changes in the weather, particularly temperature, may have been responsible. In mid-April there was a “rush” of lean Whitethroats, which are thought to have originated to the SW in Nigeria. Yellow Wagtails, Sedge Warblers and Whitethroats deposited up to 40% fat/live weight; Sand Martins Riparia riparia had up to 28%, and a small sample of Ruffs Philomachus pugnax up to 17%. In the case of Sedge Warblers, fat reserves were sufficient for crossing the Sahara both to the north and to the northeast. Contrary to the findings in some previous lipid studies, the fat-free dry weight and water content increased during fattening. The fat-free dry weight increase was about 34% in Yellow Wagtails, 18% in Sedge Warblers, and 35% in Whitethroats. The tissues involved in this increase were not investigated, but a study of Yellow Wagtails at Malamfatori in 1968 (in prep.) suggests that the pectoral muscles hypertrophy during fat deposition sufficiently to account for nearly all of the increase in fat-free dry weight and water fraction.     

Effects of seasonal migration on the latitudinal distribution of west Palaearctic bird species

This paper examines seasonal changes in the latitudinal distribution of birds (excluding seabirds) that breed in the western Palaearctic. Some resident species occupy ranges that span less than 5° of latitude year-round, while some migrant species range over more than 120° of latitude during the course of a year. Among migrant species of land and freshwater habitats, the latitudinal spans of breeding and wintering ranges are correlated. In general, species that breed over a narrow span of latitude also winter over a narrow span of latitude, and vice versa. Among both groups, for any given span of breeding range, species that winter partly in Eurasia and partly in Africa winter over a wider latitudinal span than those that winter entirely within Eurasia or entirely within Africa. Among coastal birds, there is no correlation between the latitudinal spans of breeding and winter areas; most shorebird species breed over a narrow span of (northern) latitude and winter over a wide span distributed linearly along coastlines. Several migration patterns can be distinguished, from complete overlap of breeding and wintering ranges in year-round residents, through partial separation of summer and winter ranges, to complete separation of summer and winter ranges in different geographical regions. In some such species, the gap between breeding and wintering ranges spans up to 55° of latitude (6000 km).     

Migration patterns and habitat use by passerine and near-passerine migrant birds in eastern Africa

Palaearctic migrant passerines and near-passerines which visit eastern Africa can be divided into six groups based on the latitude of their final wintering area. Species wintering further north in Africa tend (a) to breed in more southern parts of the Palaearctic, (b) to prefer drier habitats and (c) to feed more from the ground than those wintering further south. Many species use quite narrow passage routes through eastern Africa and, for some species, passage is centred further east in spring than in autumn. Southward migration can take more than 4 months from the Palaearctic breeding grounds and many birds stopover in the northern tropics from September to November. By contrast, the return migration takes only about 6 weeks. The timing and strategy of migration within Africa can be broadly related to seasonal patterns of rainfall and vegetation.     

PALAEARCTIC MIGRANTS IN NIGERIA

An account is given of the topography, climate and vegetation belts of Nigeria to illuminate the ecology of the Palaearctic migrants to the country. The faunistic relationship of the resident and migrant avifauna is considered and it is shown that approximately one species in six is migrant. It is shown that the more arid vegetation belts support the largest numbers of migrant species, the forest belt supporting only very few species. Some species appear to overfly the most northern areas. Rather few observations of visual migration are available but those known are recorded. A summary of ringing recoveries affecting Nigeria is given and it is shown that there is clear evidence of species returning to the same wintering grounds each year. The status of 135 migrant species is briefly treated and other species are mentioned where negative information is valuable. Calandrella rufescens, Cyanosylvia svecica, and Emberiza hortulana, notmentioned by Bannerman (1930-53) are included and Cettia cetti is recorded from West Africa for the first time. Several species have had their range extended to include Nigeria. A comparison of breeding areas with the Mediterranean/Sahara crossing zones and Nigerian occurrences shows that most terrestrial species in Nigeria move fairly closely to the lines of longitude. Many aquatics, mostly herons and ducks, breed much to the east of the longitude of Nigeria. It is suggested that the unexpected number of Palaearctic migrants which winter in the arid north of Nigeria indicates that food is less scarce than the appearance of the country in the late dry season indicates. There is a great need for an ornithological and ecological research station in Nigeria in an area such as southwest Lake Chad.     

Breeding origin and migration pattern of dunlin (Calidris alpina) revealed by mitochondrial DNA analysis

The large-scale migration of birds has been studied extensively by recoveries of ringed birds. However, there is very little ringing data from the arctic breeding grounds of waders. Here, the migration pattern of the dunlin, Calidris alpina, is studied with population genetic markers, using haplotype frequencies to estimate the breeding origin of migrating and wintering populations. Polymerase chain reaction (PCR) and restriction analysis of DNA from the mitochondrial control region was used to study the breeding origins of morphologically similar winter populations in the western Palaearctic, and to describe the population structure of the dunlin during winter. Also migrating dunlin from various stopover sites in Europe, Africa and Asia, were analysed with respect to their mitochondrial DNA (mtDNA) haplotypes. The genetic markers clearly show that the dunlin has a parallel migration system, with populations breeding in the western Palaearctic wintering mainly in the western part of the wintering range, and dunlin populations breeding further east wintering further east. The results also show that the distance between breeding and wintering area increases eastwards in this region.     

THE BREEDING BIOLOGY OF THE BLACK-FACED DIOCH QUELEA QUELEA IN NIGERIA

A study of the breeding biology of Quelea quelea in Nigeria, and particularly at a large breeding colony near Lake Chad, showed that losses of eggs and young were extremely small. 95% of eggs laid hatched successfully, and 87% give rise to fledglings. Nestling deaths were density-dependent and apparently due to starvation.
The incubation period was 10 days or less. By day, eggs were heated to 34°- 37° C. by the sun; at night the females incubated. The nestlings were initially fed mainly on insects, their diet gradually changing to one of seeds—mostly of the grass Echinochloa pyramidalis . The deep body temperatures of young birds were determined. It is suggested that the nestlings left the nest after, on average, 11¹/₂ days to escape intolerable temperature conditions in the nest.
Fat reserves were accumulated by nestlings and fledglings, and were utilized when the young became independent. The adults put on fat during the incubation period and lost it during the time spent feeding nestlings.
It is concluded that the most common clutch-size of Q. quelea , which is everywhere three, corresponds to the largest number of young the parents can normally nourish. This conforms to Lack's theory on the significance of clutch-size, and gives no support to Skutch's opinion that the theory does not apply to tropical birds.     

Disentangling Migratory Routes and Wintering Grounds of Iberian Near-Threatened European Rollers Coracias garrulus

Long-distance migrants are suffering drastic declines in the last decades. Causes beneath this problem are complex due to the wide spatial and temporal scale involved. We aim to reveal migratory routes, stopover areas, wintering grounds, and migratory strategies for the most southwestern populations of the near-threatened European Roller Coracias garrulus in order to identify conservation key areas for the non-breeding stage of this species. To this end, we used tracking data from seven satellite transmitters fitted to birds breeding in different populations throughout the Iberian Peninsula and four geolocators fitted to individuals in a southeastern Iberian population. Precise satellite data were used to describe daily activity patterns and speed in relation to the main regions crossed during the migration. Individuals from the most southwestern Iberian populations made a detour towards the Atlantic African coast whereas those from northeastern populations followed a straight north-to-south route. We identified important stopover areas in the Sahel belt, mainly in the surroundings of the Lake Chad, and wintering grounds on southwestern Africa farther west than previously reported for the species. Concerning the migratory strategy, satellite data revealed: 1) a mainly nocturnal flying activity, 2) that migration speed depended on the type of crossed habitat, with higher average speed while crossing the desert; and 3) that the migration was slower and lasted longer in autumn than in spring. The studied populations showed weak migratory connectivity, suggesting the confluence of birds from a wide range of breeding grounds in a restricted wintering area. Therefore, we suggest to target on defining precisely key areas for this species and identifying specific threats in them in order to develop an appropriate global conservation programme for the European Roller.     

The importance of cytotaxonomy in understanding the biogeography of African rodents: Lake Chad murids as an example

1. Due to the existence of numerous morphologically sibling species among African rodent genera, cytogenetical analyses are essential for correct taxonomic identifications. As an illustration, karyotypes of rodents captured in the Lake Chad area were systematically conducted, to determine them unambiguously. 2. Thirteen species were found, giving a list of at least 18 species in total for the whole Lake Chad region when previously published data were critically re‐examined. Gerbilline rodents dominate in this Sahelian region, with five species of Gerbillus, four of Taterillus, two of Tatera and Desmodilliscus braueri, and six species of murine rodents were also counted. Moreover, two species of Taterillus appeared characterized by unique and hitherto unknown karyotypes, and undoubtedly represent new species. 3. The Lake Chad area, although poorly studied from a biogeographical point of view, occupies a central position in the northern half of Africa which makes its zoogeographical affinities interesting to evaluate. Analysis of the biogeographical characteristics of the rodent species recorded here leads to the conclusion that the murid rodent community of the Lake Chad region is of mixed origin, with a dominant West African component. However, the presence of two species considered as endemics (Mastomys verheyeni and Taterillus lacustris), combined with our discovery of two new chromosomal species of Taterillus, suggest that this region may have acted as a centre of differentiation in relatively recent times, perhaps reflecting the extensive variations of the Lake Chad basin associated with Plio‐Pleistocene cycles of transgression–regression. 4. Without chromosomal data, most of these findings would have remained weakly supported, or even undiscovered, as between five and eight species could have been overlooked.     

SPRING WEIGHTS AND PASSAGE OF SEDGE WARBLERS ACROCEPHALUS SCHOENOBAENUS IN CENTRAL KENYA

Weight and other data were collected on over 1300 Sedge Warblers Acrocephalus schoenobaenus in central Kenya, mainly during spring (April-May). The weight, fattening rate and length of stay of spring passage birds varied from one site to another. In scrub and thicket at Kariobangi, Nairobi, passage and new arrival continued each year into May; few birds stayed to fatten, and weights over 16 g (40% above mean winter weight) were practically never recorded. At Lake Nakuru, thousands of birds were present during late April-early May 1972; many stayed to fatten for 1–3 weeks, and weights over 16 g (maximum 215 g) were not uncommon; new arrivals of lean birds continued into May. At Athi River, in 1971, many of the birds present during mid-late April stayed 1–2 weeks and fattened rapidly; 20% of all late April weights were in the range 16–21 g. Mean fattening rates at Nakuru and Athi River were 0.31 and 0.64 g/d respectively. The migration strategy and potential flight range of Kenyan spring birds is discussed. Active wing moult was rarely recorded. Most locally wintering birds and spring passage birds were judged to have renewed their plumage during the preceding October-January.     

Habitat use by Palaearctic migrant passerine birds in West Africa

Data on passerine migrants in West Africa published since the works of Moreau j 1972) and Curry-Lindhal (1981) are reviewed and new information on wintering ranges are summarized. An important feature of West Africa as a wintering zone is the presence of the two great river basins of the Senegal and the Niger, which are responsible for the Sahel being far richer than would be expected at this latitude. This 'paradoxical' or 'green' Sahel is under the threat of agricultural developments and of increasing desertification. Conversely, Palaearctic migrants can take advantage of the new-savanna/forest mosaics and agricultural developments which have replaced the rain forest.     

Evidence for Megalake Chad, north-central Africa, during the late Quaternary from satellite data

The existence of a very large Lake Chad during the late Quaternary, Megalake Chad, has long been questioned. A Megalake Chad would present strong evidence for climatic fluctuations of great magnitude during the Holocene in tropical Africa. In this paper we used satellite data from Landsat and Modis sensors to collect and analyse new information on landforms in a 2 000 000 km² region of the Lake Chad Basin. We detected 2300 km of remains marking the ancient shoreline of Megalake Chad. The satellite data also indicated many Saharan rivers and relict deltas leading to the long paleoshoreline. Large dunefield flattenings were observed and interpreted as the result of wave-cut erosion by the paleolake. Similarities were noticed between the landforms observed along the paleoshoreline of Megalake Chad and that of the former Aral Sea. This finding has significant consequences for reconstructing paleohydrology and paleoenvironments through the Lake Chad basin, and continental climate change.     

High juvenile mortality during migration in a declining population of a long‐distance migratory raptor

Many populations of long‐distance migrants are declining and there is increasing evidence that declines may be caused by factors operating outside the breeding season. Among the four vulture species breeding in the western Palaearctic, the species showing the steepest population decline, the Egyptian Vulture Neophron percnopterus, is a long‐distance migrant wintering in Africa. However, the flyways and wintering areas of the species are only known for some populations, and without knowledge of where mortality occurs, effective conservation management is not possible. We tracked 19 juvenile Egyptian Vultures from the declining breeding population on the Balkan Peninsula between 2010 and 2014 to estimate survival and identify important migratory routes and wintering areas for this species. Mortality during the first autumn migration was high (monthly survival probability 0.75) but mortality during migration was exclusively associated with suboptimal navigation. All birds from western breeding areas and three birds from central and eastern breeding areas attempted to fly south over the Mediterranean Sea, but only one in 10 birds survived this route, probably due to stronger tailwind. All eight birds using the migratory route via Turkey and the Middle East successfully completed their first autumn migration. Of 14 individual and environmental variables examined to explain why juvenile birds did or did not successfully complete their first migration, the natal origin of the bird was the most influential. We speculate that in a declining population with fewer experienced adults, an increasing proportion of juvenile birds are forced to migrate without conspecific guidance, leading to high mortality as a consequence of following sub‐optimal migratory routes. Juvenile Egyptian Vultures wintered across a vast range of the Sahel and eastern Africa, and had large movement ranges with core use areas at intermediate elevations in savannah, cropland or desert. Two birds were shot in Africa, where several significant threats exist for vultures at continental scales. Given the broad distribution of the birds and threats, effective conservation in Africa will be challenging and will require long‐term investment. We recommend that in the short term, more efficient conservation could target narrow migration corridors in southern Turkey and the Middle East, and known congregation sites in African wintering areas.     

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

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

High variation reduces the value of feather stable isotope ratios in identifying new wintering areas for aquatic warblers Acrocephalus paludicola in West Africa

Stable isotope analysis of feathers can be useful in the study of seasonal interactions and migratory connectivity in birds. For the Palaearctic–African migration system, however, the lack of isotope data from feathers of known origin in Africa renders the geographic assignment of birds captured on European breeding grounds to potential wintering areas problematic. Rectrices of the threatened aquatic warbler Acrocephalus paludicola grown in Africa were sampled across six European countries to assess whether birds in different breeding populations shared similar isotopic signatures and so were likely to have wintered in the same region in Africa. Freshly grown feathers of aquatic warblers collected at the only known wintering site in Senegal showed high variation in carbon, nitrogen, and hydrogen isotope ratios. Due to similarly high variation in isotope ratios of African‐grown feathers within all breeding populations, it was not possible to determine whether different populations wintered in different regions. However, isotope signatures of 20% of birds captured on European breeding grounds fell outside the range of those captured in Senegal, suggesting a wider wintering distribution than is currently known. We therefore assessed whether the origin of these feathers could be estimated by trying to establish isotopic gradients across sub‐Saharan West Africa. Feathers of three ecologically similar surrogate species were sampled from wetlands across a 3000 km east‐west and a 2000 km north–south transect. Within‐site variation in feather isotope ratios was frequently larger than the difference predicted by gradients across West Africa. Thus, predicting the origin of individual feathers using single‐isotope gradients was not reliable. The large within‐site variability of feather isotope ratios of a habitat specialist species like the aquatic warbler indicates that using feather isotope ratios will require large sample sizes from many locations, and may thus not be an efficient tool in identifying wintering areas of Palaearctic–African migrants.     

A NEW SUBSPECIES OF ACROCEPHALUS BAETICATUS FROM LAKE CHAD AND A TAXONOMIC REAPPRAISAL OF ACROCEPHALUS DUMETORUM

A new population of African reed warblers, Acrocephalus baeticatus hopsoni , is described from Lake Chad. Its characters, together with those of unallocated material from Senegal, Eritrea and Tibesti, are intermediate between the Palaearctic A. dumetorum and other races of A. baeticatus . It is concluded that dumetorum is conspecific with A. baeticatus , and by priority becomes A. baeticatus dumetorum . The merits of A. palustris, A. scirpaceus and Hippolais pallida for inclusion in an A. baeticatus superspecies are considered, but allow no firm conclusion.     

Interspecific and intraspecific differences in habitat use and their conservation implications for Palaearctic harriers on Sahelian wintering grounds

The floodplains of the West‐African Sahel region have experienced extensive habitat transformation during the past four decades, coinciding with an impoverishment of raptor populations. We investigated foraging patterns of Palaearctic migratory Eurasian Marsh Harriers Circus aeruginosus, Pallid Harriers C. macrourus and Montagu’s Harriers C. pygargus on a floodplain system in northern Cameroon to assess species, sex‐ and age‐related habitat preferences. Sex and age have rarely been incorporated into general studies of raptor habitat associations, despite clear evidence of intrasexual and age‐related differences in foraging strategies and diet composition, potentially carrying strong conservation implications. We found evidence of sexual differences in foraging preference related to land use, particularly in the most sexually dimorphic Pallid Harrier, and evidence that juveniles used different habitats to adults. This constitutes the first quantitative documentation of such differentiation by Palaearctic raptors on African wintering grounds, indicating that general patterns of habitat use in wintering raptors may obscure sex‐ and age‐specific preferences. Contrary to expectations, we found limited evidence for interspecific foraging segregation. Food partitioning by prey mass was related to harrier body mass and facilitated by a diverse availability of prey on human‐transformed floodplains. Anticipated further large‐scale conversion of floodplain habitat into predominantly desiccated grasslands raises concerns about the survival of wintering harriers.     

The nature and origin of the crustacean zooplankton of Sahelian Africa,with a note on the Limnomedusa

The zooplankton of the major Sahel river basins Nile, Shari (Chad), Niger, and Senegal, is different from that found in the Sahara and in Equatorial Africa. Similarities and differences between the individual basins are numerous as well. Many species are shared by the Nile and Lake Chad, by Lake Chad and the Niger (plus Senegal), or occur in all four basins, or are restricted (endemic) to only one basin.These patterns are identical to patterns found in fish, molluscs, and macrophytes and show that crustacean zooplankton obeys the same laws of dispersal as these groups, in spite of its apparent preadaptation to passive dispersal.The patterns can be explained by the climatic fluctuations of the Upper Pleistocene and,Holocene. Following a general dry period over Africa between 20 000 and 13 000 BP, high river and lake levels prevailed between 12 500 and 8 400 BP. This was the period of maximum faunal interchange between all basins, and even with the Zaire basin. After a regression (8 000–7 000 BP), wetter conditions returned around 6 000 BP, but the Sahel itself remained dry, although its rivers and lakes, fed by waters of southern origin, showed higher levels than today. They flooded large areas of the southern Sahara, permitting aquatic animals and plants to reach the Adrar of Mauretania, the Tibesti, and the Ennedi mountains.Since 3 000 BP, present day conditions developed. This last period is characterized by species extinctions, as exemplified by the droughts in Lake Chad in historical times, and in spite of the tremendous diversity still extant here today. Between 6 000 BP and the present, however, very little speciation took place, and faunal exchange between basins was very limited.     

4. GAME RESERVES IN SOUTHERN AFRICA

Steyn, P. &; Grobler, J. H. 1981. Breeding biology of the Booted Eagle in South Africa. Ostrich 52:108-118.

The Booted Eagle Hieraaetus pennatus is a breeding visitor to the Cape Province of South Africa, wintering mostly in Namibia on present knowledge. Palaearctic birds probably also reach the Cape but arrive later. Two nests in different localities in the Cape were studied. The birds breed soon after arrival. Both sexes build the nest on a cliff ledge. Incubation, which lasts 40 days, is done mostly by the female. The female spends most of her time on the nest during the first four weeks of the nestling period, but considerably less time thereafter. The male provides nearly all the prey until near the end of the nestling period, and helps to feed the young. Details of nestling growth and behaviour and of parental care are given. The nestling period was 50 and 54 days in two cases. Post-nestling dependence is about two months. Prey preferences in the two study areas were very similar: 54% birds, 33% lizards and 13% rodents. Breeding biology in South Africa is basically the same as that of Palaearctic populations, with the main differences being the habitat and cliff nest site.