Variation in haematozoan parasitism at local and landscape levels in the red-billed quelea Quelea quelea

The red-billed quelea Quelea quelea , one of the most abundant birds in the world, presents two fundamental conundrums that we investigate here with a novel approach using blood parasite assemblages at two spatial scales, landscape and individual. The quelea of southern Africa Q. q. lathamii are split by a hypothesized migratory divide, where birds follow rain fronts in one of two directions (NW or SE). This divide is not detectable in the host population using microsatellite data, and here we show that it is also not apparent from our large-scale phylogeographical analyses of the haematozoan parasite. At a finer scale, the colourful and variable breeding plumage of male red-billed quelea has not previously shown a correlation with predictors of quality, as it does in many other bird species. The male's breeding plumage is partially based on carotenoid colouration, the quality of which has been correlated with haematozoan infection in other bird species. However, we found no correlation between intensity of male carotenoid colouration and haematozoan infection. Our results do not contradict the hypothesis that male breeding plumage in this species serves to identify individuals rather than to indicate quality. Finally, we recovered the greatest number of haematozoan lineages from any phylogenetic survey of a single host species to date. Understanding the reasons for the extreme diversity of parasite lineages in this species may assist in explaining the success of the red-billed quelea in anthropogenic landscapes.     

Lack of genetic and plumage differentiation in the red-billed quelea Quelea quelea across a migratory divide in southern Africa

A migratory divide usually signals the presence of a geographical region over which other traits, such as morphology and genotypes, also undergo rapid change. A migratory divide has been hypothesized in central southern Africa for the abundant migratory weaver, the red-billed quelea Quelea quelea. The positioning of the divide in the region is based on the patterns of rainfall in the region that stimulate the annual migrations of queleas. Evidence indicates that premigratory queleas near the divide show two distinct preferred directions for migration. We used eight polymorphic microsatellite loci and a range of plumage characters to determine whether there was population structure among red-billed queleas in southern Africa, and specifically whether this structure coincided with the location of the migratory divide. There was no evidence of population genetic structure. An amova revealed no significant differences between samples taken either side of the migratory divide. Similarly, there was no geographical variation in plumage patterns across southern Africa. For both microsatellites and plumage characteristics, the variation that does exist occurs within each sampled site, with little differentiation between sites. We were therefore unable to find any evidence that either plumage or microsatellite genotypes varied in a similar way to migratory direction preference in red-billed queleas in southern Africa. This is perhaps because the migratory divide does not act to separate individuals into populations within which genetic and plumage differentiation can be maintained.     

Genetic evidence for male biased dispersal in the red-billed quelea Quelea quelea

In order to detect sex-biased dispersal in the red-billed quelea Quelea quelea in southern Africa, we used the assignment index technique to determine the probability that individuals originated from the population in which they were sampled. This is the first time that this multilocus genetic test has been used in a bird species and is informative despite evidence that the population under study exhibits little genetic structure. There was a pattern of male-biased dispersal, the first example in a passerine, and the first time that evidence of a sex-biased pattern of dispersal has been shown for queleas.     

Morphological changes in the thymus of young and adult red-billed queleas Quelea quelea (aves).

In a wild population of red-billed queleas Quelea quelea L. (Ploceidae: weaver-birds) sampled throughout the year in East Africa, the thymus was found to enlarge in young birds shortly after hatching, remain enlarged during the juvenile stage, and regress towards the end of the postjuvenile moult. In adults, recrudescence occurred in many individuals during the prenuptial and postnuptial moults, and also in most if not all individuals, of both sexes, for a brief period during a breeding session. Thymus enlargement in both young and adults has been found to be accompanied by marked erythropoietic activity within the gland, and it is suggested that this activity is related to an increased demand for erythrocytes which may occur during moult and breeding.     

Histological changes associated with enlargement and regression of the thymus glands of the red-billed quelea Quelea quelea L. (Ploceidae: weaver-birds).

Thymic lobes form over 200 red-billed queleas, Quelea quelea L., were examined histologically. Samples were taken from embryos about to hatch, juveniles and adults. The lobes varied in size from very small to very enlarged (1- greater than 5 mm long). The constituent cell types are described in detail and the occurrence of these cells in different sized lobes is discussed. A cycle of events is proposed which accounts for the observations presented here. It is suggested that the large numbers of erythroid cells found in the cortex of some individuals were developing in situ. The significance of erythropoiesis within the thymus is discussed.     

The migrations of the Red-billed Quelea Quelea quelea and their relation to crop damage

Recent studies on the migrations of the Red-billed Quelea Quelea quelea have shown that while the species is capable of long-distance movements of hundreds of kilometres, the actual migrations probably more often consist of shorter distances with a flexible timing. This flexibility allows adjustments to be made to the rainfall distribution in a particular year. New information comes from studies on the breeding distribution, on the plumage and moult of local populations, and from attempts to apply new techniques such as mass-marking with fluorescent particles. Breeding has been shown to be both more widespread and more frequent than previously recognized, while local populations can prolong breeding in one place if rainfall continues. Local populations have been identified that are apparently distinct, with limited intermixing with other populations, and some data suggest that intermixing may be reduced by flock or group cohesion. The flexibility of the migrations means that predictions of major influxes of quelea into agricultural areas have proved difficult to make, especially in regions where the rainfall patterns are complex and variable, such as East Africa.     

Candidate genes for carotenoid coloration in vertebrates and their expression profiles in the carotenoid-containing plumage and bill of a wild bird

Carotenoid-based coloration has attracted much attention in evolutionary biology owing to its role in honest, condition-dependent signalling. Knowledge of the genetic pathways that regulate carotenoid coloration is crucial for an understanding of any trade-offs involved. We identified genes with potential roles in carotenoid coloration in vertebrates via (i) carotenoid uptake (SR-BI, CD36), (ii) binding and deposition (StAR1, MLN64, StAR4, StAR5, APOD, PLIN, GSTA2), and (iii) breakdown (BCO2, BCMO1). We examined the expression of these candidate loci in carotenoid-coloured tissues and several control tissues of the red-billed quelea (Quelea quelea), a species that exhibits a male breeding plumage colour polymorphism and sexually dimorphic variation in bill colour. All of the candidate genes except StAR1 were expressed in both the plumage and bill of queleas, indicating a potential role in carotenoid coloration in the quelea. However, no differences in the relative expression of any of the genes were found among the quelea carotenoid phenotypes, suggesting that other genes control the polymorphic and sexually dimorphic variation in carotenoid coloration observed in this species. Our identification of a number of potential carotenoid genes in different functional categories provides a critical starting point for future work on carotenoid colour regulation in vertebrate taxa.     

Ornamental plumage does not signal male quality in red-billed queleas.

Sexually selected ornaments often function as condition-dependent signals of quality (or 'indicators'). When ornamentation is costly, only high-quality individuals can afford to produce the most elaborate signals. The plumage ornamentation of male red-billed queleas, Quelea quelea, is an ideal candidate for an indicator because it is continuously variable, conspicuous, sexually dimorphic, is displayed only during breeding and is partially based on carotenoid pigmentation. However, I show here that quelea plumage is not an indicator because first, plumage colour is not correlated with physical condition or age; second, plumage colour is a genetically determined phenotype that is unresponsive to environmental variation; third, different plumage characters have bimodal distributions; fourth, plumage characters vary independently of one another; and finally, plumage colour is not correlated with reproductive success. To my knowledge, this is the first demonstration of non-condition dependence in colourful and sexually dimorphic breeding ornamentation. Instead, plumage variation may function as a sexually selected signal of individual identity among territorial males that nest in huge, densely packed and highly synchronized colonies.     

DETRIMENTAL EFFECTS OF FENTHION (QUELETOX® UL), USED TO CONTROL RED-BILLED QUELEA (QUELEA QUELEA), ON RHEOPHILIC BENTHIC MACROINVERTEBRATES IN THE ORANGE RIVER

Summary

The relative abundance of rheophilic macroinvertebrates in the Orange River was assessed before and 15 hrs after two aerial applications of fenthion (QUELETOX® UL) above roosting sites of the red-billed quelea (Quelea quelea). The first treatment had no detectable effect on benthic macroinvertebrate abundance, whereas the second treatment caused mortality among at least 17 taxa (over half the taxa present). The most sensitive species were nymphs of the mayflies Baetis glaucus and Tricorythus discolor, followed by larvae of the midge Rheotanytarsus fuscus. Fenthion had no detrimental effect on the population densities of the mayfly Afroptilum excisum.

The results of this study warn of the dangers of fenthion in water supplies. It is suggested that alternative methods of quelea control along the Orange River should be sought.     

Sizes and numbers of nuclei in the cortex of thymus glands of red-billed weavers Quelea quelea

The cortex of the thymus glands of embryos, chicks, juveniles, fledglings and adults from several colonies of Quelea quelea were studied using an image analyser (Quantimet 720) to determine cell populations and nuclear sizes. Just prior to hatching the lobes showed a high level of mitosis and consisted of predominantly small lymphocytes. The larger glands of chicks and juveniles had higher cell populations; pyknotic cells and erythrocytes occurred free in the cortex. The lobes of adults were more variable but in general mitosis occurred in enlarging glands of adults from colonies with eggs; most lobes contained pyknotic cells but not in such high numbers as in lobes from chicks and fledglings. Erythrocytes were common, occurring in large numbers in the cortex in some birds. The factors affecting the interpretation of these data are discussed in detail.     

EMMA-4 analysis of iron in cells of the thymic cortex of a weaver-bird (Quelea quelea).

Combined morphological and analytical studies with the EMMA-4 analytical electron microscope have enabled very early erythroid cells to be identified within the cortex of enlarging thymic lobes of Quelea quelea. These early erythroid cells have pale cytoplasm (sometimes with ferritin-like crystals present), slightly pachychromatic nuclei and have fewer cell organelles (mitochondria) than lymphocytes. Counts for iron were approximately 70% lower than counts from mature erythrocytes found free in the cortex. Iron was also recorded from some epithelial reticular cells and pyknotic nuclei; no iron was recorded from small lymphocytes (thymocytes) in the cortex. The presence of very early erythroid cells is a further indication that erythropoiesis occurs in situ in the avian thymus.     

Traditional capture of Red-billed Quelea Quelea quelea in the Lake Chad Basin and its possible role in reducing damage levels in cereals

Mullié, W.C. 2000. Traditional capture of Red-billed Quelea Quelea quelea in the Lake Chad Basin and its possible role in reducing damage levels in cereals. Ostrich 71 (1 & 2): 15–20.

Three traditional methods of capturing Red-billed Quelea Quelea quelea practised by different ethnic groups, were studied in the Lake Chad Basin in Chad and Cameroon: capture of birds by triangular hand-held nets by Hadjeraï piégeurs (trappers); with standing fishing nets of the “mist net” type by Gambai; and by cast nets (épervier) by Massa and Mousgoum fishermen.

The Hadjeraï net was found to be both selective for queleas and highly efficient. As many as 1.2 million birds were captured from 13 June to 21 August 1994 around N'Djamena, Chad, alone, until activities ceased due to heavy rains. Birds were trapped in tree roosts during moonless periods of the night. They were plucked and fried the following morning, subsequently dried in the sun, and eventually transported to the market in N'Djamena to be sold. The piégeurs operated in teams of about 6 men and each team could process about 20 000 birds per day. Once this number had been captured, they withdrew from further trapping. The catches were almost entirely composed of queleas, with sometimes a small percentage (<1%) of Golden Sparrow Passer luteus. It was estimated that annually at least 5–10 million queleas were trapped, with a market value of about 19–38 million FCFA (USA37 500–75 000).

The impact of trapping on the population of queleas in the Lake Chad Basin, estimated at about 200 million individuals in 1976/77, was insignificant. Since activities of piégeurs are aimed at maximising revenues, and not at minimising damage levels, there is no causal relationship between crop damage and number of birds trapped. However, compared to crop losses in pearl millet due to quelea damage, revenues from selling quelea are up to 40% of capitalized crop losses due to quelea in the same area.     

Identifying multiple wildlife species-crop interactions using network analysis

Managing crop damage by wildlife is a complex challenge in agrarian-wildlife landscapes. Losses to farmers and their negative attitude towards crop-raiding wildlife, compromise wildlife conservation and farmer-wildlife coexistence. Priority crop types, crop-raiding wildlife and socio-ecological factors are highly integrated. Here we use cross-sectional network analysis as a conservation planning tool to simultaneously identify the crop types used by crop-raiding wildlife, and evaluate the importance of these crop-wildlife interactions relative to the socio-ecological factors that affect these interactions. The most vulnerable crops were maize, millet and fruit trees, while the most problematic crop-raiding wildlife were African elephant, bushpig, hippo, warthog and red-billed quelea. Crop damage declined with countermeasure effectiveness, farm size, and distance from river to farm, but increased with richness of attractive crops. Undamaged crops included cash crops, such as chili, onion, ginger, lemon grass and garlic. Applying network analysis to different crops and damage scenarios enables identification of the most important and influential crops, crop-raiding wildlife and socio-ecological factors needed to develop effective crop protection strategies.     

Ultrastructure of the spermatozoon of Anomotaenia quelea (Mettrick, 1961) (Cestoda,Cyclophyllidea, Dilepididae), an intestinal parasite of Quelea quelea (Aves,Ploceidae) in Senegal

The mature spermatozoon of Anomotaenia quelea exhibits an apical cone of electron-dense material and two helicoidal crest-like bodies. The apical cone near its base is surrounded by a lucent cytoplasm and a spiraled layer of cortical microtubules. The crest-like bodies are of different lengths, spiraled and make an angle of 30–40° to the hypothetical spermatozoon axis. The axoneme is of the 9 + ‘1’ trepaxonematan pattern and is surrounded by a periaxonemal sheath of electron-dense material. The cytoplasm contains in regions III and IV numerous electron-dense granules situated between the periaxonemal sheath and the cortical microtubules. The posterior extremity of the spermatozoon of A. quelea exhibits a nucleus and a disorganized axoneme and cortical microtubules. This type of posterior extremity of the mature spermatozoon has never been described previously in a Dilepididae. Similarly, two crest-like bodies have not been observed before in a dilepidid cestode.     

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

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

Autumn migratory orientation and displacement responses of two willow warbler subspecies (Phylloscopus trochilus trochilus and P. t. acredula) in South Sweden

Topography and historical range expansion has formed a so-called migratory divide between two subspecies of willow warbler (Phylloscopus trochilus) in central Scandinavia. The autumn migratory directions of individuals assigned molecularly to both subspecies and possible hybrids were recorded using orientation cage experiments in southwest and southeast Sweden. We found pronounced differences in willow warblers’ orientation in respect to genotype. The mean directions registered in the control experiments were in accordance with the ringing recoveries and analyses of stable isotopes for Scandinavian willow warblers. With the same individuals we performed displacement experiments between both sites. They resulted in non-significant orientation, which could be explained by the intermediate distance of the displacement or reactions to housing, transportation and location. On a separate set of birds we tested whether stress following transportation could explain the disorientation and found that orientation before and after transport was unchanged. Experimental studies of effects of intermediate displacements across longitudes and studies of orientation of hybrid individuals in the zones of migratory divides are crucial for understanding the mechanisms underlying orientation behaviour. Our work further stresses the importance of knowing the migration genotype of a particular bird under study, in order to correctly evaluate expected migration routes.     

Haematocrit values of breeding Red-billed queleas Quelea quelea (Aves: Ploceidae) in relation to body condition and thymus activity

In a Red-billed quelea colony in north-eastern Nigeria that was later abandoned as a result of the poor nutritional condition of the adults, laying females showed low haematocrit values compared with normal pre-breeding haematocrit levels but nestbuilding males did not. In females the low haematocrits were correlated with extremely low levels of reserve lipid and labile protein but males in equally poor condition did not have such low haematocrits. This difference must result in some way from the additional nutritional burden for females of mobilizing sufficient protein to form eggs. Earlier observations of massive red blood cell production in the thymus of queleas during incubation suggest that low haematocrits, indicating anaemia, may be a normal occurrence in queleas during breeding, and were not peculiar to this abnormal abandoned colony.     

Migration patterns of Hoopoe Upupa epops and Wryneck Jynx torquilla: an analysis of European ring recoveries

For many bird species, recovery of ringed individuals remains the best source of information about their migrations. In this study, we analyzed the recoveries of ringed European Hoopoe (Upupa epops) and the Eurasian Wryneck (Jynx torquilla) from 1914 to 2005 from all European ringing schemes. The aim was to define general migration directions and to make inferences about the winter quarters, knowing that hardly any recoveries are available from sub-Saharan Africa. For the autumn migration, there is evidence of a migratory divide for the Hoopoe in Central Europe, at approximately 10–12°E. Autumn migration directions of Wrynecks gradually change from SW to SE depending on the longitude (west to east) of the ringing place. In both species, only a few recoveries were available indicating spring migration directions, but they showed similar migration axes as for autumn migration, and hence no evidence for loop-migration. Due to a paucity of recoveries on the African continent, we can make only limited inferences about wintering grounds: extrapolating migration directions are only indicative of the longitude of the wintering area. The directions of autumn migration indicate a typical pattern observed in European long-distance migrants: west-European Hoopoes and Wrynecks are likely to winter in western Africa, while central- and east-European birds probably winter more in the east. Due to the migratory divide, for the Hoopoe, this phenomenon is more pronounced.     

How does a first year passerine migrant find its way? Simulating migration mechanisms and behavioural adaptations

In the Palaearctic-African migration system, birds face several trade-offs on their first autumn migration. The shortest route, minimising travelling time, would lead them directly south across the Mediterranean Sea and the Sahara Desert, involving long stretches of no refuelling possibilities. This route is risky because of flight range constraints. Besides taking longer, a detour along the Iberian peninsula may require a more complex orientation mechanism. We simulated migrants with stopover and orientation behaviour and investigated the effect of flight costs and behavioural rules (e.g. crossing or flying along coast lines, a shift in migratory direction) on the resulting flight path and especially on the evolution of endogenous directions. The simulation of autumn migration from southern Scandinavia to south of the Sahara showed that it would be possible to reach the winter quarters by vector summation with a constant endogenous direction, but then either orientation must be very accurate or flight costs must be small. For small passerines both requirements are so far not corroborated by empirical studies. Alternatively, flying along coast lines or shifting direction in northern Africa from south-westerly to southerly, resulted in similar survival rates as with a constant south-westerly endogenous direction, but with a larger range of feasible values. Although weather factors were not included, our results suggest that the Mediterranean Sea and the Sahara desert had a dominating influence on the evolution of endogenous directions. This influence is probably acting through flight range constraints.     

Age-dependent migration strategy in honey buzzards Pernis apivorus tracked by satellite

Six adult and three juvenile honey buzzards Pernis apivorus were radio-tracked by satellite during autumn migration from southwestern Sweden. All adults crossed the Mediterranean Sea at the Strait of Gibraltar and continued across the Sahara desert to winter in West Africa, from Sierra Leone to Cameroon. Analysing three main steps of the migration, (1) from the breeding site to the southern Mediterranean region, (2) across the Sahara and (3) from the southern Sahara to the wintering sites, the adults changed direction significantly between these steps, and migrated along a distinct large-scale detour. In contrast, the juveniles travelled in more southerly directions, crossed the Mediterranean Sea at various places, but still ended up in the same wintering areas as the adults. Average speeds maintained on travelling days were similar for the two age groups, about 170 km/day in Europe, 270 km/day across Sahara and 125 km/day in Africa south of Sahara. However, as the adults used fewer stopover days en route, they maintained higher mean overall speeds and completed migration in a shorter time (42 days) than the juveniles (64 days). Although the juveniles set out on more direct courses towards the wintering grounds, they did not cover significantly shorter distances than the adults, as they tended to show a larger directional scatter between shorter flight segments. The results corroborate previous suggestions that adult and juvenile honey buzzards follow different routes during autumn migration, and that the birds change migration strategy during their lifetime. While juveniles may use individual vector orientation, social influences and learning may be of great importance for the detour migration of adults. The remarkable and distinct age-dependent shift in migratory route and orientation of the honey buzzard provides a challenging evolutionary problem.