Highly mobile insectivorous swifts perform multiple intra‐tropical migrations to exploit an asynchronous African phenology

With timely allocated movement phases, mobile organisms can match their space‐use with the seasonality of the environment and thereby optimise their resource utilisation over time. Long‐distance avian migrants are known to move with the seasonal dynamics on an annual basis, but how individuals respond to seasonality within their tropical non‐breeding range has been less studied. Here we analyse the movement pattern of a highly mobile aerial insectivorous bird, the pallid swift Apus pallidus, and its association with the local habitat phenology during the non‐breeding period, using individual‐based light‐level geolocation. We extracted timing and location of 21 birds’ residence periods, as well as characteristics of the intervening movements, such as distance and speed. We used time series of precipitation and vegetation data for each residence area to extract the timing of the local end of the rainy season and the onset of the dry season. The pallid swifts repeatedly upgraded their habitat by undertaking 2–5 intra‐tropical migrations correlated with the withdrawal of the rains and the onset of the local dry season. The birds arrived to the sites on average 12 days after rains ended and departed about two weeks after the onset of dry season suggesting that the birds closely tracked a spatiotemporal window presumably timed with optimal foraging conditions. Our results provide insights in the ways Palaearctic–African migrants respond to the asynchronous phenology within their sub‐Saharan non‐breeding range. We confirmed that pallid swifts actively respond to deteriorating conditions by repeated upgrades in habitat quality, which likely have substantial consequences for an individual's access to an essential, spatiotemporally ephemeral food resource. However, the pallid swifts did not surf an apparent resource wave per se as would be expected in a highly mobile species, indicating that also other factors, such as spatial patchiness of resources, may influence the movement decision.     

NOTES ON BIRDS IN THE GAMBIA

Recent observations in The Gambia have shown that the status of many species has changed in the last forty or fifty years and have provided data on the local breeding seasons for a number of birds. Several species have been found for the first time in this part of West Africa. The various birds concerned are given in a systematic list, together with notes on all the Palaearctic migrants so far recorded from The Gambia. Organized research is recommended on several lines: on the sequence of food sources, especially through the long and severe dry season, when Palaearctic birds are added to resident and when some African species breed. The bird ecology of the mangroves remains to be worked out, and also the relative importance to birds of cleared ground and untouched "bush" at all seasons.     

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.     

THE INUNDATION ZONE OF THE NIGER AS AN ENVIRONMENT FOR PALAEARCTIC MIGRANTS

Of at least 350 species so far recorded within the boundaries of the Niger Inundation Zone, no less than 108 (31%) are wholly or partially of Palaearctic origin. Five main habitats are recognized in the region: wetland, hygrophilous grassland, transition zone, non-flooded areas and aerial. The habitats, the Palaearctic migrants and their possible Ethiopian competitors are described and discussed. The annual Niger flood regime enables Palaearctic waterbirds to find suitable habitats somewhere within the region during all seasons, but most widely during the autumn and winter months of the flood recession. Non-aquatic species inhabiting flood plain grassland are scarce during the Palaearctic autumn, when the growth of vegetation reaches its maximum, becoming commoner and more diverse during the winter months. Wetland warblers of the genera Acrocephalus and Locustella have not been recorded on autumn passage. It is suggested that in some years at least, these and other trans-Saharan migrants from the West Palaearctic overfly the Sahelian latitudes of mid-West Africa. Species inhabiting the transition zone (which increases in area during years of below average rains and floods) are most in evidence at the end of the winter period and during the spring hot dry season, prior to northward trans-Saharan migration. Habitats and species encountered in the non-flooded areas are similar to those recorded in the Sahel zone elsewhere in West Africa. Aerial habitat is utilized by Common Swifts arriving en masse in early August, at the maximum development of the south-westerly monsoon airstream. Several aquatic species and the first few trans-Saharan migrants also occur during the summer rainy season. Some species (e. g. Whiskered Tern, Lesser Kestrel, Turtle Dove, Short-toed Lark and Sand Martin) are most numerous during the late winter and spring hot seasons, when conditions provided by the Inundation Zone may be more suitable for pre-migration feeding than in other parts of the Sahel zone. As yet there is virtually no information available to determine any effects that drought seasons or flood variation may have on the migrant populations.     

Protected areas as biodiversity benchmarks for human impact: agriculture and the Serengeti avifauna

Protected areas as biodiversity benchmarks allow a separation of the direct effects of human impact on biodiversity loss from those of other environmental changes. We illustrate the use of ecological baselines with a case from the Serengeti ecosystem, Tanzania. We document a substantial but previously unnoted loss of bird diversity in agriculture detected by reference to the immediately adjacent native vegetation in Serengeti. The abundance of species found in agriculture was only 28% of that for the same species in native savannah. Insectivorous species feeding in the grass layer or in trees were the most reduced. Some 50% of both insectivorous and granivorous species were not recorded in agriculture, with ground-feeding and tree species most affected. Grass-layer insect abundance and diversity was much reduced in agriculture, consistent with the loss of insectivorous birds. These results indicate that many species of birds will become confined to protected areas over time. We need to determine whether existing protected areas are sufficiently large to maintain viable populations of insectivorous birds likely to become confined to them. This study highlights the essential nature of baseline areas for assessing causes of change in human-dominated systems and for developing innovative strategies to restore biodiversity.     

SOME ASPECTS OF THE BEHAVIOUR OF THE WATTLED PLOVER AFRIBYX SENEGALLUS (LINNAEUS)

The question of how aridity might influence avian clutch size, through the influences of rainfall seasonality and environmental stochasticity (unpredictability), has received little attention. A marked east-west gradient in aridity across South Africa provides a unique opportunity to test for such influences. Using an extensive collection of nest records for 106 terrestrial bird species from the South African Nest Record Card Scheme, we tested three predictions related to rainfall seasonality and stochasticity. Analyses were conducted at two levels, the first examining each species independently, and the second grouping species into five dietary guilds. The first prediction, that clutch size should generally increase with higher rainfall seasonality (i.e. higher seasonal fluctuation of food availability), was supported, particularly in the most arid environments where food abundance is more closely linked to rainfall. Controlling for rainfall seasonality, the second prediction, that clutch size should generally decrease as a bet-hedging strategy in arid, stochastic environments, was also supported. Although the timing of the rainy season differs among regions in South Africa (winter, early summer, later summer, year-round), birds primarily nest during spring. The relative timing of rainfall and breeding is expected to have different consequences for seasonal variation in clutch size among rainfall regions. The third prediction, of different patterns of seasonal variation in clutch size between rainfall regions, was also supported. In the winter and early-summer rainfall regions, early-nesting birds (breeding with or soon after the rains) generally had a larger clutch size than late-nesting birds. In the late-summer rainfall region, early-nesting birds (breeding well before the rains) had a smaller clutch size than late-nesting birds.     

Garden Warbler Sylvia borin migration in sub-Saharan West Africa: phenology and body mass changes

The Garden Warbler is a classic subject for the study of Palaearctic–African bird migration strategies. Most studies have considered the situation close to the breeding areas, while the African and especially the sub‐Saharan part of the species’ migration have received comparatively little attention. Here we use autumn and spring ringing data from Nigeria and The Gambia to study the movements and energetics of the species in West Africa during the non‐breeding season. The first Garden Warblers arrive south of the desert around the beginning of September, roughly at the same time as the median date for their passage through the Baltic Sea region and c. 3 weeks before their median passage date through southern Italy. In the Nigerian Sahel savannahs, where, owing to the rainy season and its associated increase in food availability, many more Garden Warblers stop over in autumn than in the dry spring, the median date of passage is 1 October. The body mass on arrival south of the desert is normally only a few grams more than the lean body mass (LBM; 15 g) – with a mean of 16.6 g (sd = ±1.8 g) in The Gambia and 17.4 g (sd = ±1.8 g) in the Nigerian Sahel. After resting and refuelling in the Sahel, Sudan and Guinea‐type savannahs the Garden Warblers depart during November–December for wintering areas further south. Before leaving, they again increase their body mass, with an average fuel load of c. 20%, and often more than 50% relative to LBM. Some of the birds passing through Nigeria probably spend midwinter around the Congo Basin. During spring they return northwards to the Guinea savannah zone in April and fuel‐up there for the trans‐Sahara passage. At this time they normally increase their body reserves to around 50% of the LBM, but c. 10% of the birds gain 100%, thus doubling their mass. The passage there peaks around 20 April and continues well into May. That the main take‐off northwards is directly from the Guinea savannahs is indicated by the very low numbers trapped in the Sahel during spring.     

Short‐distance migration of Wrynecks Jynx torquilla from Central European populations

European Wrynecks Jynx torquilla torquilla have generally been considered to be long‐distance Palaearctic–African migrants that spend the non‐breeding season in Sahelian Africa, where they have been reported regularly. Results from tracking individual birds showed that Wrynecks from two Central European populations migrated only relatively short distances to the Iberian Peninsula and northwestern Africa (c. 1500 km and 3000 km, respectively), compared with a minimum distance of about 4500 km to Sahelian Africa. Additionally, differences in wing lengths of populations from Central and Northern Europe support the idea of leap‐frog migration, populations from Northern Europe being long‐distance migrants with a non‐breeding distribution in Sahelian Africa.     

Palaearctic migrants in the central Sudan

A brief account is given of the topography and climate of the Sudan and a more detailed survey is made of the vegetation and habitats afforded to wintering Palaearctic migrants in the Sahel and Soudan savannah zones in the central Sudan. Climatic conditions during the migration seasons are mentioned and the utilization of habitats by wintering migrants is broadly surveyed.
The status of 196 Palaearctic migrant species is treated briefly. The basic observational data are tabulated to summarize observed patterns of migration in terms of date, relative abundance and region (for which purpose the central Sudan is divided into four from west to east). Hitherto unpublished field notes of the authors and their collaborators are used, together with dated observations (including specimens collected) of earlier authors, to provide the fullest data on each species.
Moreau's (1967) paradox is discussed as it relates to the Sahel and Soudan savannah zones in the central Sudan, together with the degree of competition with resident African species in those zones. Some disparities between the spring and autumn abundance and d. istribution of Palaearctic migrants are discussed in relation to 40 species ana some tentative views are advanced on the possible reasons for such disparities. Evidence is adduced which points to Lanius minor and Coraczus garrulus (and possibly Circus pygargus , partially at least) having loop migrations, south through the Sudan and north through Arabia, and Falco wespertinus south through the Sudan and north through West Africa. An explanation is offered for the greater volume of water birds observed migrating down-Nile in spring compared with their volume up-Nile in autumn, and for the reversed situation observed in the region west of the Nile for these same species.
Shelduck Tadorna tadorna is included in the Sudan list for the first time.     

Moult of three Palaearctic migrants in their West African winter quarters

Some theories about moult strategies of Palaearctic passerine migrants assume that birds adapt timing of moult to environmental conditions such as rainfall on their African wintering grounds. Species wintering in the northern tropics should limit moult to the period shortly after their arrival at the end of the rainy season. Passerine migrants wintering in West Africa should also moult more rapidly compared to related species or conspecific populations that moult elsewhere. We investigated the moult of melodious warblers Hippolais polyglotta, willow warblers Phylloscopus trochilus and pied flycatchers Ficedula hypoleuca wintering in Comoé National Park, Ivory Coast, between October 1994 and April 1998. In contrast to previous studies we did not restrict our analyses to moult of flight feathers but also included moult of body feathers. The results differed partially from the general assumptions of previous authors. Melodious warblers moulted twice: a complete moult shortly after their arrival, and a moult of body feathers and in some cases some tertials and secondaries in spring. Willow warblers moulting flight feathers were found between December and March with the majority moulting in January and February. Primary moult was not faster compared to populations moulting in central Africa and South Africa. Body feather moult varied strongly among individuals with birds in heavy moult between December and April. Pied flycatchers moulted body feathers and tertials between January and April. Birds with growing feathers were found throughout the whole period including the entire dry season. Moult strategies are thus not readily related to a few environmental factors in general and our results show that factors other than mere resource availability during certain times on the wintering grounds are likely to govern the timing of moult.Communicated by F.Bairlein     

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.     

6.—DATA OF BIRDS' EGGS IN MY COLLECTION

Van Zyl, A.J, Jenkins, A.R. &; Allan, D.G. 1994. Evidence for seasonal movements by Rock Kestrels Falco tinnunculus and Lanner Falcons F. biarmicus in South Africa. Ostrich 65:111-121.

Patterns of seasonal movement by Rock Kestrels Falco tinnunculus and Lanner Falcons F. biarmicus in South Africa were determined using data from the Southern African Bird Atlas Project, road transect counts, ringing recoveries and other sources. Both species were found to be partial, facultative migrants, and Lanner Falcon movements involved mostly immature birds. Rock Kestrels were found to concentrate in the southwest of the country in the breeding season, and to move north and east in the non-breeding season. Lanner Falcons concentrated in the east of the country during the breeding season, and moved into the west in the non-breeding season. The movement by both species into the arid northwest of the country is probably associated with dramatic increases in prey abundance, in turn associated with summer rah events. Lanner Falcon movement into the southern Cape apparently coincides with cereal farming practices, affecting the availability of prey (granivorous birds and rodents). Rock Kestrel movement into the east of the aunt is probably to exploit increases in insect abundance mated with summer rain. The pattern of Rock Kestrel movements suggests that it does not compete for food with the Lesser Kestrel F. naumanni. a Palearctic migrant to the east of the country.     

THE MIGRATION PATTERNS OF QUELEA QUELEA IN AFRICA

Red-billed Queleas migrate, at the beginning of the wet season, away from their dry-season concentration areas and towards areas where rain started several weeks earlier. Considerable fat deposits are accumulated for this “early-rains migration”. The direction taken by the migrants, the distance they must fly, and the timing of the movement are dependent upon the timing of the rains and the way the rain-front moves. On the return “breeding migration” individuals in breeding condition stop to breed (in large aggregations) wherever they find conditions suitable for the founding of colonies. The location of the colonies can vary greatly from year to year. The migrations performed by several populations, in different parts of Africa, are predicted on the basis of general rainfall patterns, and the predictions tested against the facts available. There is evidence that individual females are able to produce a succession of broods in the same breeding season, at colonies which may be very far apart, and probably with different mates. The adaptive value of this “itinerant breeding” is discussed. Many other bird species, which breed at a particular phase of the wet-season/dry-season cycle, are expected to perform similar “itinerant breeding”.     

海南岛尖峰岭热带山地雨林林下鸟类群落研究

从 2 0 0 0年 5月到 2 0 0 2年 3月 ,用网捕法对海南岛尖峰岭热带山地雨林 (包括原始林和次生林 )林下鸟群落进行研究。共张网 3 992 .7网· h,捕到林下鸟 3 12只 40种 ,隶属于 6个目 11个科。林下鸟群落虽然以小于 2 5g重的雀形目鸟类占优势 ,但许多个体较大的非雀形目鸟类也到林下活动 ,最大个体重达 3 70 g,大于 10 0 g重的个体占整个林下鸟生物量的比例较大。尖峰岭热带山地雨林中的林下鸟以食虫鸟为主 ,食虫鸟占整个林下鸟种数的 70 .0 %、鸟数量的 77.56% ;食果鸟虽然占第 2位 ,但其所占比例较小。林下鸟的种类组成和密度存在季节变化。留鸟是林下鸟最重要的成分 ,留鸟占鸟类种类的比例超过 85% ,留鸟的捕获数量占整个捕获鸟类数量的比例超过 90 % ;迁徙鸟的种类和数量均较少 ,对林下鸟群落的季节变化影响不大。原始林与演替 40 a后的次生林林下鸟平均网捕率相近但林下鸟群落种类组成不同 ,其种的相似性仅为 0 .3 5。     

Waterfowl of a subtropical African floodplain

Many water bodies in arid southern Africa are seasonal and waterfowl habitats become restricted during the dry winters. Nevertheless, some tropical species, such as the white-faced duck, are expanding their ranges into more temperate regions where summer habitats are available, thus increasing pressure on overwintering habitats. The subtropical Pongolo River floodplain is an important overwintering area for a diverse water-fowl community that includes 13 of the 16 species that breed in South Africa and one of the few regular migrants. Historical evidence suggests that numbers of white-faced duck, which represent over 70% of the floodplain community, have increased 10-fold over the last two decades. An abundant, nutritious food (Potamogeton crispus turions) attracts waterfowl to the Pongolo floodplain during winter and an increase in bird fat reserves suggests that they leave for summer breeding grounds in good condition. Unseasonal flood releases from an upstream reservoir may cause low food supplies in late spring before the onset of the summer rainy season when breeding habitats become available. This study emphasizes the need for a better understanding of the role of winter habitats in maintaining waterfowl populations.     

Icacina oliviformis (Icacinaceae): A close look at an underexploited crop. I. Overview and ethnobotany

Icacina oliviformis is an underexploited, suffrutescent, savannah pyrophyte used for its edible seeds, tuberous roots, and fruit mesocarp in many areas of west and north-central Africa. The large seeds provide a relied-upon resource in Senegal, Guinea-Bissau, and the Central African Republic at the end of the dry season when food reserves in the villages are at their lowest annual levels. The tuberous roots, sometimes weighing over 50 kg, provide a rich source of starch in times of famine. This species deserves consideration for use in extensive agroforestry schemes in savannah habitats of west-central Africa.     

Consequences of Fragmentation of Tropical Moist Forest for Birds and Their Role in Predation of Herbivorous Insects

The consequences of habitat alteration on the role of understory insectivorous birds as predators of herbivorous insects in tropical forests are poorly understood. To examine whether fragmentation may affect the top–down controls of herbivory, we compared the number of species, individuals, and the community structure of insectivorous birds between fragments and continuous tropical moist forest in Mexico. We also registered insect herbivore abundances and conducted a larvae predation experiment to evaluate the potential role of insectivorous birds as predators of herbivorous insects. We recorded 63 bird species from 22 families, 43 percent of which were insectivorous birds. Species richness, abundance, and diversity of the avian community were higher in continuous forest compared with forest fragments. For insectivorous birds in particular, there was low similarity in avian insectivore communities between forest types, and forest fragments had more heavily dominated communities of avian insectivores. During the dry season, forest fragments presented significantly higher predation rates on artificial caterpillars, and lower abundance of herbivorous Lepidoptera larvae, compared with continuous forest. Furthermore, there was a significant negative correlation between artificial caterpillar predation rate and larval Lepidoptera abundance, with higher rates of predation in sample sites of low Lepidoptera abundance. Hence, the potentially greater light in the dry season combined with a more dominated avian insectivore community in forest fragments may facilitate increased predation by avian insectivores, resulting in a decline in abundance of larval Lepidoptera, with implications for the process of insect‐driven herbivory in forest fragments.     

西双版纳白背桐次生林土壤种子库、种子雨研究

 本文讨论了西双版纳常见的次生林白背桐(Mallotus paniculatus )林的土壤种子库和种子雨的组成及动态。结果表明该林地的种子雨年输入量为745粒/m2,土壤种子库储量在雨季末期为3345±438粒/m2,旱季末期为4555±554粒/m2,种子雨的输入高峰出现在旱季,导致旱季末期土壤种子库储量大于雨季末期。森林破碎化导致的隔离效应阻碍了种子的传播途径,使土壤种子库储量下降,种子雨和土壤种子库中鸟类传播的种子比例增大。传播成熟林树种的大型动物到达不了该样地,植被的演替受到了阻碍。     

THE FORMATION, STRUCTURE AND FUNCTION OF MIXED-SPECIES INSECTIVOROUS BIRD FLOCKS IN WEST AFRICAN SAVANNA WOODLAND

Mixed-species flocks of birds were observed during the wet season (July to September 1975) in savanna woodland in Ghana. Thirty-four flocks contained birds of 56 species in 20 families, including insectivorous, granivorous, and nectarivorous species, using a wide range of foraging methods. Only two species occurred in more than half the flocks. There was no correlation between the number of flocks joined by a species and its abundance in the community. Among insectivores, but not granivores, the species which joined most flocks were those which habitually occurred in the largest single-species groups. All stages of breeding activity were represented by the various members. Some species joined flocks only while these were passing through their territories. Of the two species which were present most frequently, there were no differences between mixed and single-species flocks for Eremomela pusilla, but Parus leucomelas foraged and called on more occasions in mixed flocks than single-species flocks, though the rates of foraging and calling were related only to the number of P. leucomelas present. Groups of P. leucomelas appeared to initiate some flocks by attraction due to their conspicuous wing-bars, active movement, and loud calls. Black-and-white species joined them first, followed by birds of other plumage patterns. The advantages of mixed flocking are thought to be connected with finding patches of the food of bark- and foliage-searching insectivores, which were the only species regularly seen foraging in the flocks. Because of dry season burning which leaves small unburnt patches of savanna, these insect species may share a common, patchy distribution. Birds may also gain protection from predators, and some species probably gain no advantages. The species composition and behaviour of flocks previously recorded elsewhere in African savannas are similar to the Ghana flocks.     

Process in the evolution of bird migration and pattern in avian ecogeography

Current ideas about the evolution of bird migration equate its origin with the first appearance of fully migratory populations, and attribute its evolution to a selective advantage generated by increased breeding success, gained through temporary emigration from resident populations to breed in under-exploited seasonal areas. I propose an alternative hypothesis in which migration first appears as a temporary directional shift away from the breeding site outside the reproductive period, in response to seasonal variation in the direction and/or severity of environmental gradients. Fully migratory populations then appear through either extinction of sedentary phenotypes, or colonisation of vacant seasonal areas by migrants. Where colonisation occurs, resident ancestral populations can be driven to extinction by competition from migrants which invade their range outside the breeding season, resulting in fully migratory species. An analogous process drives the evolution of migration between high latitudes and the tropics, since extension of breeding range into higher latitudes may drive low latitude populations to extinction, resulting in an overall shift of breeding range. This process can explain reverse latitudinal gradients in avian diversity in the temperate zone, since the breeding ranges of migratory species concentrate in latitudes where they enjoy the highest breeding success. Near absence of forest-dwelling species among Palaearctic-African migrants is attributable to the lack of forest in northern Africa for much of the Tertiary, which has precluded selection both for southward extension of migration by west Palaearctic forest species, and northward breeding colonisation by African forest species.