Weavers

(Ploceidae)

Class Aves

Order Passeriformes

Suborder Passeri (Oscines)

Family Ploceidae

Thumbnail description
Small to medium-sized passerine birds; bill conical or pointed; plumage plain yellow or black, or these colors in combination with red, brown, or orange, or else sparrowy brown; often there is a seasonal change in plumage, which may include development of greatly elongated tail-feathers; many species highly social, occurring in large flocks

Size
4.3–10 in, up to 28 in with elongated tail (11–25 up to 70 cm); 0.3–2.3 oz (9–65 g)

Number of genera, species
19 genera; 135 species

Habitat
Forest, woodland, swamps, savanna, semi-arid regions

Conservation status
Critically Endangered: 1 species; Endangered: 6 species; Vulnerable: 7 species; Near Threatened: 3 species; Data Deficient: 2 species

Distribution
Sub-Saharan Africa, Arabian Peninsula, South and Southeast Asia, Madagascar, Mauritius, Réunion, Comoros, Seychelles

Evolution and systematics

The fossil record of passerine birds is fragmentary and difficult to interpret. There are no conspicuous skeletal characters in ploceids that distinguish their bones from those of other passerine families, which makes it unlikely that fossils will provide clear evidence of their origins. However, both fossils and molecular data suggest that the passerine birds are an old group, and that many extant families could be as old as 40 million years. Thus the weaver group are likely to have evolved in Africa over a period during which there have been dramatic changes in climate, with the extent of forest cover fluctuating greatly. This would have promoted speciation in both forest and open-country habitats.

Earlier studies based on anatomy and other morphological characters linked the Ploceidae to the Estrildidae, another family of seed-eating birds which is primarily African. This was supported by the DNA-hybridization studies of Charles Sibley, but he placed both these groups as sub-families in a new family (Passeridae), along with the sparrows (Passerinae), wagtails and pipits (Motacillinae), and accentors (Prunellinae). This arrangement remains controversial, and we have followed a more traditional system. The four subfamilies of Ploceidae are most easily defined by their breeding habits: the buffalo weavers, Bubalornithinae (three species), build large nest structures of sticks and have a unique, rigid phalloid organ on the abdomen; the sparrow-weavers, Plocepasserinae (nine species), build nests of straight grass stalks which are not woven but stuck into the nest structure; the "true" weavers Ploceinae (103 species) weave a closed nest, with the entrance either at the side or below; the whydahs, indigobirds, and cuckoo finch Viduinae (20 species) are brood parasites, which lay their eggs in the nests of other birds. The position of the Viduinae is debatable. Behavioral studies by J. Nicolai suggested links to some Ploceinae such as the bishop-birds, whereas skeletal and molecular data imply that the waxbills (Estrildidae) are their closest relatives. The cuckoo finch (Anomalospiza imberbis) has usually been classified as a weaver, but both morphological and molecular data show that it belongs in the Viduinae.

The genus Ploceus (Ploceinae) is one of the largest bird genera at present with more than 60 species. An examination of skull characteristics suggests that there are several distinct groups within this genus, and new studies may lead to it being broken up into several distinct genera. The relationships between the African and Asian Ploceus weavers are not clear, and they have evidently been separated for a long time. The Foudia species on the Indian Ocean islands appear to be derived from the African genera Quelea or Euplectes, whereas the two Madagascar Ploceus species could be African or Asian in origin. These conclusions are based on plumage, nest

structure, carotenoid pigments, and some skeletal characters; no molecular studies had been published by 2001.

Physical characteristics

The weavers have no defining physical characteristics which are shared by all or even most members of the family. The sexes may be virtually indistinguishable, even in the hand, or highly dimorphic. Tails can be short or extravagantly long. The bill is always straight, not curved, but varies from short and heavy to longer and quite slender. At the sub-family level, there is more consistency. Buffalo weavers are either mainly black or mainly white, with heavy seed-eater bills. Sparrow-weavers are all "sparrowy" brown in appearance, with some black or white plumage areas. There is no obvious seasonal plumage change in either of these groups, and little sexual dimorphism, although males are usually larger. Within the parasitic Viduinae, there is marked sexual dimorphism in plumage during the breeding season, after which males molt into a plumage which resembles that of the females. They can usually be disinguished from other small seed-eating birds by black stripes on the crown of the head. Male indigobirds are blackish, with pale or reddish bill and legs, in varying combinations. Male whydahs have mainly black or black-and-white breeding plumage with very long central tail feathers, which may be either narrow or broadened. The male cuckoo finch is canary-yellow in breeding plumage.

Among the Ploceinae, there are conspicuous differences between genera. Males are almost always larger than females, while sexual dimorphism in plumage is especially marked in polygynous species. However, even in dimorphic species, the males do not always have a seasonal plumage change. Eye color often changes with age from brown to red, yellowish, or creamy; in many cases only males have a distinctively colored eye. The bill color of male birds may change seasonally from brown to black, in response to increased levels of male sex hormones. The genus Malimbus is remarkably uniform. All species are predominantly black with some red, or in one case yellow, plumage; males and females differ in plumage, and juvenile birds have a distinctive plumage, different to both adults. There is no seasonal change in plumage. In contrast the open-country bishops and widows (Euplectes) all have sparrowy brown females, while males molt into a breeding plumage which is wholly or partly black, with either red or orange to yellow areas, and in some cases a long, black tail. Young birds resemble females, and males do not usually acquire breeding plumage until at least their second year. The large genus Ploceus includes species that are sexually dimorphic with or without a seasonal change in plumage, and species in which the sexes are identical. Black and/or yellow are the predominant plumage colors in males, with some green, brown, or orange, but never red, feathers.

Distribution

Weavers occur throughout sub-Saharan Africa, where all sub-families are represented. Only two genera of Ploceinae are found outside Africa; the fodies (Foudia) which are endemic to Madagascar and other Indian Ocean islands, and Ploceus with two species on Madagascar and five in Asia. One East African species, Rüppell's weaver (Ploceus galbula), also occurs on the Arabian peninsula. Several species are commonly exported as cage birds, and escapes or deliberate releases have led to their establishment, sometimes temporary, in other regions, including Australia, California, Portugal, Hawaii, St. Helena, and some islands in the West Indies. An Asian species, the streaked weaver (Ploceus manyar), is now established in the Nile delta in Egypt, and is believed to have escaped from Alexandria Zoo.

Habitat

Many weavers are associated with water, since they breed in wetlands, along rivers, dams, and lakes, nesting in reeds or other waterside vegetation. However, in these cases they often move to grassland or savanna during the non-breeding season. Several species may breed in wetlands, but also in trees far from open water, and have adapted well to man-modified habitats such as farmland. Only members of the sparrow-weavers and buffalo weavers are permanent residents of arid and semi-arid areas. Some species are exclusively forest birds, either in lowland or montane evergreen forest, and may spend much of their time in the canopy 100 ft (30 m) above the ground. All members of the genus Malimbus are strictly forest inhabitants.

Behavior

Although many species of weavers move about extensively during the dry season, these are local movements rather than predictable, long-distance migration. The red-billed quelea (Quelea quelea) does carry out predictable movements in many regions, and these seem to be correlated with rainfall patterns. This appears to be the only species that could qualify as a migrant throughout its range.

Although they may have a wide range of different calls, few weavers would be considered "songbirds" in the conventional sense. The songs that male weavers use to advertise their territories are often a harsh, repetitive chatter with no tuneful, musical notes. Some forest species do sing short phrases, sometimes as duets, which are more attractive to our ears. The parasitic indigobirds learn elements of the song of their host species while in the nest, and later incorporate these into the songs which they use in courtship.

Feeding ecology and diet

Categorizing weavers as insectivorous or granivorous is misleading. All species will take insects when they are available, and the young are often fed primarily insects, especially in the first days after hatching. There is frequently a seasonal change in diet, with seeds the main or even the only food source in the dry season, and insects more important in the rainy season. The heavy bill of the grosbeak weaver (Amblyospiza albifrons) enables the birds to open sunflower seeds, but they have also been seen to catch small frogs. Small lizards are on the menu of several other species in the wild. Fruit and berries are eaten readily, and nectar from plants such as Aloe and Erythrina. Here weavers are messy feeders, often eating the whole flower and stripping the plants, leaving with their faces caked with pollen. The Cape weaver (Ploceus capensis) is probably the main pollinating agent for the endemic South African crane flower Strelitzia regina.

Reproductive biology

Social organization in weavers shows clear correlations with habitat and feeding ecology, as J. H. Crook first demonstrated in his innovative comparative studies. Forest weavers are generally insectivorous and remain in pairs throughout the year, whereas seed-eating species of the open savanna associate in flocks, and form colonies for breeding. This, in turn, influences their breeding systems, with monogamy usual in the forest species, while many of the colonial weavers are polygynous, with one male mating in turn with a succession of different females.

Nests and nest construction have attracted most attention in this group of birds. The pioneering work of Nicholas and

Elsie Collias—who observed many species in the field, in captivity, and in museum collections—has provided an excellent framework for the evolution of nest-building in the family. In buffalo weavers and sparrow-weavers the technique is simple, with the nests formed as piles of interlocking material. These birds are associated with nests throughout the year, and thus maintain the structures with periodic building at all seasons; both sexes participate to some degree. In the true weavers (Ploceinae) nest-building is seasonal and these are short-lived structures, which mostly do not survive beyond one breeding season. The commonest pattern is for the male to produce a nest frame by weaving and knotting strips of material collected and prepared for this purpose. Once the female has mated and accepted a particular nest, she then adds the lining. However, the female's contribution varies greatly, depending on the mating system; in Jackson's widow (Euplectes jacksoni) the female is solely responsible for building and lining the nest, which is not on the male's territory.

Courtship in sparrow-weavers and buffalo weavers involves song and visual displays, generally near the nest structures. In

the Ploceinae, among the monogamous forest species, courtship frequently takes place away from the nest, even before construction begins. By contrast the colonial species set up territories and build nests before intensive courtship starts, and the male often displays hanging at the nest entrance. Bursts of display activity may sweep through a colony as groups of females arrive, and the males all appear to be vying for their attention. In the polygynous species, each male will build a series of nests, and try to attract as many females as possible. Nest that are not accepted by females, or are no longer occupied, are often demolished. The male may then rebuild at the same site so that a single male masked weaver (Ploceus velatus) can build more than 20 nests over a three-month season. There is one exceptional species, Jackson's widow, in which males display at dancing grounds which constitute a lek: a male courtship arena which females visit to select a mate, after which they go off to nest and have no further contact with the male. In the parasitic Viduinae, males set up "song-posts" at which they display, and to which females are attracted. The breeding system is thus a form of lek, where the females visit males only to mate, although the males do not occupy a communal display ground.

Parental care is closely correlated with the mating system. In monogamous species, both incubation and feeding of the young may be shared equally between the partners, whereas in polygynous species the female normally does all the parental duties without assistance. In some cases, polygynous males may feed at the nest late in the season, or occasionally feed the young once they have fledged. Compared to some other African bird families such as the starlings, cooperative breeding in which several related or unrelated individuals help the parents to rear the young is rare in weavers. It occurs primarily in some of the sparrow-weavers in arid country, where a nesting tree forms a permanent base for the group, which roosts in the nests throughout the year. The situation in the parasitic Viduinae is especially interesting. Whydahs and indigobirds lay their eggs in the nests of waxbills; the eggs of both host and parasite are plain white, and similar in size. Waxbill young have highly distinctive mouth-markings, which are matched by the young parasites. The young are raised together, so although the waxbill parents have extra mouths to feed, they do not lose their whole brood as is often the case for cuckoo hosts. However, the cuckoo finch parasitizes small grassland warblers, and the host young seldom survive.

Conservation status

BirdLife International has produced a review of globally threatened birds, and an account of the Important Bird Areas of Africa. The major threat to weaver species is habitat loss, since some of them have very restricted ranges. Three island fodies are threatened both by habitat loss and introduced predators on Mauritius, Seychelles, and Rodrigues, respectively. Foudia rubra may be Critically Endangered, whereas F. sechellarum and F. flavicans are currently regarded as Vulnerable. The Asian yellow weaver (Ploceus megarhynchus) is a grassland species with a restricted range in India. Although the Asian golden weaver (Ploceus hypoxanthus) occurs in several countries, it is uncommon and regarded as Near Threatened.

On mainland Africa, the golden-naped weaver (P. aureonucha) and the yellow-footed weaver (P. flavipes) are both known only from the Ituri Forest, and have been seen just a few times in the last 30 years. Their canopy habitat and the political problems in this region make it difficult to obtain accurate information. Four localized species in West Africa, Bannerman's weaver (P. bannermani), Bates's weaver (P. batesi), the Gola malimbe (Malimbus ballmanni) and the Ibadan malimbe (M. ibadanensis), occur in forest that is disappearing rapidly throughout this region. The situation is most critical for the Ibadan malimbe, which has the smallest range. Two little-known species, the Loango weaver (P. subpersonatus) on the coastal strip and the black-chinned weaver (P. nigrimentum) in open savanna, range from Gabon southwards towards Angola.

In East Africa, Clarke's weaver (P. golandi) is restricted to the Arabuko-Sokoke Forest in Kenya, while the Tanzanian mountain weaver (P. nicolli) is found in relict forest patches on the Usambara Mountains and a few other sites. Fortunately both areas are now the site of active conservation programs. Agricultural changes in the highland grasslands of Kenya are a potential threat to Jackson's widow, while Fox's weaver (P. spekeoides) is apparently confined to one lake system in central Uganda, but remains unstudied. The Kilombero weaver (P. burnieri) was a surprising discovery in Tanzania, described in 1990 and evidently limited to a small area.

Significance to humans

Several colonial weaver species are closely associated with human settlements, nesting in exotic vegetation, and in forested areas, taking advantage of habitat changes to colonize new clearings. Eggs and nestlings may be utilized for food on occasion, but often the relationship is quite harmonious. The long tail feathers of breeding male long-tailed widows (Euplectes progne) were once used as elements in traditional head-dresses for warrior tribes in South Africa, but otherwise the colored plumages have not been utilized.

For hundreds of years, grain-eating weavers have been a pest for farmers in Africa. M. Adanson, a French botanist for whom the baobab genus Adansonia is named, spent several years in Senegal from 1747, and reported that the inhabitants suffered greatly from the depredations of the weavers. He described several traditional bird-scaring methods which are still in use in Africa today. Since the 1960s the red-billed quelea has been recognized as the major pest of cultivated cereals in Africa. Despite international efforts to reduce its numbers, using aerial spraying and fire-bombs set under roost sites, it remains enormously abundant: in March 2000 the South African department of agriculture reported that an estimated 21 million queleas had been killed in control operations during the past month! It seems that in the past, queleas bred prolifically in good years, and then starved when food supplies declined. Today when wild grass seeds are unavailable, they find crops a very acceptable alternative and consequently agriculture enables them to maintain high population levels. To the interested naturalist, a vast flock of queleas "roller-feeding" (in constant motion, with the birds at the back flying up over those ahead of them to be first at the untouched plants) is one of the great spectacles of Africa, but it is a catastrophe for the small farmer, and there is no simple, effective solution.

Species accounts

List of Species

Red-billed buffalo weaver

Bubalornis niger

subfamily

Bubalornithinae

taxonomy

Bubalornis niger A. Smith, 1836, Kurrichane, South Africa.

other common names

French: Alecto à bec rouge; German: Büffelweber; Spanish: Tejedor Búfalo de Pico Rojo.

physical characteristics

8.7 in (22 cm); unsexed 2.7–2.9 oz (78–82 g). Male dark blackish brown, white patch in wings in flight. White bases to body feathers may show when plumage ruffled. Bill and legs red. Female dark brown, variably flecked with white on underparts. Bill and legs brown. Juveniles paler with more white on underparts. Bill orange-yellow.

distribution

Ethiopia and Somalia through eastern Africa to Angola, Zambia, and northern Mozambique, south to northern South Africa.

habitat

Dry thornveld with large trees.

behavior

In groups or non-breeding flocks up to 50 birds, may associate with other species. Usually present at nest sites throughout the year.

feeding ecology and diet

Mainly insects, also seeds and fruit. Most food collected on the ground.

reproductive biology

Colonial and polygynous, often polygynandrous (each male mates with several females, and female mates with more than one male), since in most broods genetic studies indicate multiple paternity. Nest is a large mass of thorny twigs, containing up to 13 nest chambers, lined with green vegetation. Male builds main structure and starts lining chambers; female adds further lining before laying. Phalloid organ not inserted during copulation, but stimulation from this structure may be essential for successful mating. Lays three to four eggs in spring and summer. Incubation 11 days, fledging 20–23 days. Female alone incubates and does most feeding of chicks; male feeds young occasionally.

conservation status

Not threatened; dependent on large trees, but much habitat is sparsely populated.

significance to humans

None known; may use human-made structures as nest sites, or nest near homesteads.

White-browed sparrow-weaver

Plocepasser mahali

subfamily

Plocepasserinae

taxonomy

Plocepasser mahali A. Smith, 1836, Orange River, South Africa.

other common names

French: Mahali à sourcils blancs; German: Augenbrauenmahali; Spanish: Tejedor Gorrión de Cejas Blancas.

physical characteristics

6.7 in (17 cm); female and male 1.6–1.8 oz (45–52 g). Brown upperparts with broad white eyebrow and white rump, white underparts. Bill brown to black. Sexes alike, juvenile paler than adult with a pale bill.

distribution

Southern Sudan, Ethiopia, and Somalia south through eastern Africa to Zambia, South Angola, and northern South Africa.

habitat

Mopane and acacia savanna in relatively dry country.

behavior

Groups of up to 12 birds resident, defend territory of about 55 yd (50 m) in diameter with complex songs and group displays. Strong dominance hierarchy within group, with a single breeding pair. Roost singly in nests.

feeding ecology and diet

Insects and seeds, in variable proportions. Most food collected on the ground; birds will dig in soil, roll over small stones, clods, and elephant droppings.

reproductive biology

Colonial, cooperative breeder. Nest is an elongated retort made of straight dry grass stems, resting on thin branches, initially with an opening at each end. Both male and female, and other group members, may contribute to building nest. Second entrance closed in breeding nests, which are then lined with feathers. Nest orientation related to prevailing winds. Lays two to three eggs, mainly in spring and summer but also in other months. Incubation 14–16 days, fledging 18–23 days. Juveniles still fed occasionally up to three months after leaving nest. Female alone incubates and feeds young for first three days. Male seldom feeds young; helpers related to breeding pair do much of the feeding. Unrelated helpers assist in territorial defense, but not in raising young.

conservation status

Not threatened; common in many thinly populated areas.

significance to humans

None known, although may feed on wheat or maize in winter, usually only grains left behind in fallow lands.

Sociable weaver

Philetairus socius

subfamily

Plocepasserinae

taxonomy

Loxia socia Latham, 1790, Great Namaqualand, South Africa.

other common names

French: Républicain social; German: Siedelweber; Spanish: Tejedor Sociable.

physical characteristics

5.5 in (14 cm); female and male 0.8–1.1 oz (24–32 g). Sandy brown with black chin and throat, dark feathers with pale edges on mantle and flanks. Bill blue-gray. Sexes alike. Juvenile uniform sandy brown, with no darker feathers. Bill pale brown.

distribution

Namibia, southwestern Botswana, northwestern South Africa.

habitat

Open, arid regions with scattered trees and bare ground.

behavior

Gregarious, resident at nest sites, roosting in chambers throughout the year. Predators such as cobras may live within nest structure, also "lodgers" like the pygmy falcon (Polihierax semitorquatus), which is an obligate commensal. Other birds may occasionally roost or breed in vacant nest chambers.

feeding ecology and diet

Seeds and insects, particularly harvester termites. Proportion of insect and seed food varies seasonally, collected primarily on the ground. Feed in flocks within 1 mi (1.5 km) radius of nest site. Seldom drink water.

reproductive biology

Colonial, monogamous with cooperative breeding. Communal nest is a huge mass of dry grass stems, with individual nest chambers entered from below. Up to 13 ft (4 m) deep and 24 ft (7.2 m) long, supported by large branches; in treeless areas may use telephone poles. Pair bond may last for only one breeding attempt, even if both partners resident in same nest mass; helpers chiefly offspring of pair from earlier broods. Lays two to six eggs; season entirely dependent on rainfall, and breeding may start in any month. Incubation 13–14 days, fledging 21–24 days. Both sexes incubate and feed the young; up to nine helpers may feed chicks.

conservation status

Not threatened. Range is thinly populated, includes major conservation areas.

significance to humans

Nest material sometimes used for stock fodder in times of drought.

Blue-billed malimbe

Malimbus nitens

subfamily

Ploceinae

taxonomy

Ploceus nitens J. E. Gray, 1831, Sierra Leone.

other common names

English: Gray's malimbe; French: Malimbe à bec bleu; German: Rotkehlweber; Spanish: Malimbe de Gray.

physical characteristics

5.7–6.7 in (14.5–17.0 cm); female 1.0–1.2 oz (29–36 g), male 1.3–1.7 oz (38–47 g). Black with scarlet throat; female less glossy than male, black tinged with brown, and red less intense. Bill blue-gray, eye red. Juvenile sooty brown with throat and breast dull orange-brown. Eye gray-brown.

distribution

Senegal east to extreme western Uganda, south to Democratic Republic of Congo and northern Angola.

habitat

Lowland forest, oil palms, swamp forest, and mangroves; occasionally dense savanna woodland.

behavior

Usually solitary or in pairs, occasionally groups up to seven birds; regularly joins mixed-species flocks. Very shy except at the nest.

feeding ecology and diet

Mainly insects, also spiders and some fruit. Forages on thin twigs, clusters of dry leaves, and vine tangles, mainly at midlevels of forest.

reproductive biology

Solitary and monogamous, although sometimes several pairs nest close together. Male courtship, singing and posturing, occurs away from the nest. Nest is ball-shaped, woven from palm strips, rootlets, or fibers, with canopy over entrance placed low at one side; always overhanging water. In Ghana the birds appear to select nest sites close to crocodile dens. Lays one to two eggs during late summer to autumn. Incubation 14 days, fledging 16 days. Female alone incubates, and broods small chicks; both male and female feed young.

conservation status

Not threatened, but will not survive without undisturbed forest habitat.

significance to humans

None known.

Spectacled weaver

Ploceus ocularis

subfamily

Ploceinae

taxonomy

Ploceus ocularis A. Smith, 1828, Eastern Cape = Grahamstown, South Africa.

other common names

French: Tisserin à lunettes; German: Brillenweber; Spanish: Tejedor Moteado.

physical characteristics

5.9–6.3 in (15–16 cm); female 0.7–1.0 oz (21–30 g), male 0.8–1.1 oz (22–32 g). Greenish yellow weaver with slender, dark bill and dark "spectacle" line through the eye. Eyes pale cream. Male has dark bib on the throat, lacking in female. Juvenile lacks spectacle line or bib, eye brown, bill pale brown.

distribution

Cameroon east to Sudan, Ethiopia, south to northern Namibia, northern Botswana, and eastern South Africa.

habitat

Open woodland, forest edge, thickets, and gardens.

behavior

Singly or in pairs throughout the year, family groups after breeding. May join mixed-species flocks of insectivorous birds. Territorial, calling regularly, a descending "tee-tee-tee-tee."

feeding ecology and diet

Mainly insectivorous, gleaning leaves and branches and probing bark. Also takes berries, small geckos, nectar, and bread and chicken feed in gardens.

reproductive biology

Nests are finely woven, suspended singly from tip of vegetation, with entrance tunnel 4 in (10 cm) long. Lays one to four eggs, usually two to three, in spring to summer in different regions. Incubation 13–14 days, fledging 15–19 days. Both sexes incubate and feed young. Occasionally parasitized by Diederik cuckoo (Chrysococcyx caprius).

conservation status

Not threatened. Widespread over large area, occurs in manmodified habitats such as suburban gardens.

significance to humans

None known.

Dark-backed weaver

Ploceus bicolor

subfamily

Ploceinae

taxonomy

Ploceus bicolor Vieillot, 1819, 'Senegal' = South Africa.

other common names

English: Forest weaver; French: Tisserin bicolore; German: Waldweber; Spanish: Tejedor Bicolor.

physical characteristics

5.5 in (14 cm); female 1.0–1.3 oz (29–36 g), male 1.1–1.6 oz (32–46 g). Upperparts dark, underparts golden yellow. Bill dark with blue-gray rims. Eye brown or red in different

regions. Sexes alike. Juvenile duller than adult, bill light brown.

distribution

Nigeria east to southern Sudan and eastern Africa, south to eastern South Africa but absent from Botswana, Namibia.

habitat

Forested areas and dense riverine vegetation, subtropical thicket.

behavior

In pairs throughout the year, small family parties after breeding. Joins mixed-species flocks. Song a musical duet, heard at all times of the year.

feeding ecology and diet

Insects, mainly gleaned from branches, bark, and tangled vegetation in mid-story of forest. Also fruit and berries, and nectar.

reproductive biology

Monogamous, solitary nester. Often returns to same site, so several old nests may be in close proximity. Nest is retort-shaped with broad entrance tunnel pointing downwards; suspended at tip of branch or creeper. Woven of thin vines and creepers, appears rough and always looks old and dry. Lays two to four eggs in summer. Incubation 15–17 days, fledging 22 days. Probably both sexes incubate; both feed young.

conservation status

Not threatened, but dependent on preservation of well-wooded habitats.

significance to humans

None known.

Village weaver

Ploceus cucullatus

subfamily

Ploceinae

taxonomy

Ploceus cucullatus P. L. S. Müller, 1776, Senegal.

other common names

English: Spotted-backed weaver; black-headed weaver, V-marked weaver; French: Tisserin gendarme; German: Textor-weber; Spanish: Tejedor de la Villa.

physical characteristics

5.9–6.7 in (15–17 cm); female 1.1–1.5 oz (31–43 g), male 1.1–1.6 oz (32–45 g). Breeding male has head black; forehead yellow in southern birds, and extent of black on throat and breast varies. All populations have upperparts yellow spotted with black, underparts plain yellow. Bill black, eye red. Female and non-breeding male upperparts dull olive, eyebrow, throat, and breast yellow to buff, belly whitish. Bill brown; older females may have red eye. During breeding season, female more yellow on underparts. Juvenile like female, eye brown.

distribution

Senegal east to Somalia, south to northern Namibia, northern Botswana, eastern South Africa. Introduced to Haiti, Dominican Republic, Puerto Rico, Mauritius, and Réunion.

habitat

Open wooded areas; in forest zone, in clearings and secondary growth, cultivated areas.

behavior

Gregarious, in flocks when foraging and forms large roosts, often with other weavers. May be nomadic in dry season, and possible regular movements in some regions.

feeding ecology and diet

Varied diet includes seeds, insects, flowers, nectar. Forages on the ground or gleans on vegetation and tree trunks.

reproductive biology

Colonial, polygynous. In central Africa often in large mixed colonies with Vieillot's black weaver (Ploceus nigerrimus). Nest is oval with entrance below, may have a short spout. Woven by male, who displays hanging below nest, fluttering wings, and calling. Breeding varies regionally, may continue throughout the year in central Africa. Lays two to five eggs. Incubation 12 days, fledging 17–21 days. Female alone incubates, male may feed nestlings. Often parasitized by Diederik cuckoo.

conservation status

Not threatened, very widespread, common in human-modified habitats and often abundant.

significance to humans

Familiar commensal throughout central Africa; can be significant crop pest for subsistence farmers.

Sakalava weaver

Ploceus sakalava

subfamily

Ploceinae

taxonomy

Ploceus sakalava Hartlaub, 1867, Madagascar.

other common names

French: Tisserin sakalave; German: Sakalavenweber; Spanish: Fodi Sakalava.

physical characteristics

5.1–5.9 in (13–15 cm); adult 0.7–0.9 oz (20–27 g). Breeding male has yellow head, gray underparts and back, brown wings and tail. Bare reddish skin around eye, unique in weavers, blue-gray bill. Female is paler below, with whitish throat and distinct brown stripes on side of head, above and below eye; bill pale gray. Non-breeding male like female, but bare pinkish skin around eye. Juvenile like female, but paler with bill horn-colored.

distribution

Western Madagascar.

habitat

Open, lowland areas including cultivated land, spiny bush, and deciduous dry forests.

behavior

Highly gregarious, typically in flocks of 200 or more.

feeding ecology and diet

Feed primarily on the ground, collecting seeds, but also forage in trees and marshes, and young are fed primarily insects.

reproductive biology

Colonial, some males may be polygynous, but mating system not studied. Nest is retort-shaped, often suspended on a short woven rope, with entrance tunnel up to 16 in (40 cm) long. Both male and female build nest, and colonies usually in trees, often within villages, and even attached to thatched roofs of huts. Small colonies may be placed under nests of crows or large birds of prey. Breeding season varies with rainfall, especially in dry southwest. Lays two to four eggs; incubation and fledging periods unrecorded. Female alone incubates, but both sexes feed young.

conservation status

Not threatened, widespread in open and cultivated areas, and seldom molested.

significance to humans

Appear to take only waste rice, and not regarded as agricultural pests. In many areas weaver colonies in villages, especially those nesting close to a house, are considered a sign of good fortune and consequently protected.

Baya weaver

Ploceus philippinus

subfamily

Ploceinae

taxonomy

Loxia philippina Linnaeus, 1766, Philippines = Sri Lanka.

other common names

French: Tisserin baya; German: Bayaweber; Spanish: Tejedor de Baya.

physical characteristics

5.1–5.9 in (13–15 cm); female 0.7–1.0 oz (20–28 g), male 0.7–0.9 oz (20–26 g). Breeding male, yellow crown, black face mask, mottled brown upperparts, paler, unstreaked underparts; bill black. Female and non-breeding male, mottled rufous-brown upperparts, some streaking on underparts, bill brown. Juvenile like female.

distribution

Pakistan, India, and Sri Lanka east to southwestern China, south throughout Malay Peninsula, Sumatra, Java, Bali.

habitat

Forest edge, open savanna and scrub, cultivated areas. Appears to prefer agricultural land.

behavior

Gregarious in flocks, and forms large communal roosts in reedbeds or sugarcane, together with other weavers, seed-eating birds, starlings, and bulbuls.

feeding ecology and diet

Adults mainly seed-eating, including rice, sorghum, millet, and wheat. Also consumes insects, even frogs recorded, and young fed primarily insects.

reproductive biology

Colonial, polygynous. Nest is retort-shaped with entrance tunnel of varying length, often suspended over water in trees or bushes. Male builds nest, displays there to attract mates. Blobs of mud, sometimes dung, regularly added to inside of nest. Long-standing but unsubstantiated legend that male embeds fireflies in mud, so that they illuminate the inside of the nest. Lays two to five eggs, breeding from spring through to autumn, depending on timing of monsoon. Incubation 14–15 days, fledging 13–16 days. Female alone incubates, male may assist in feeding young. Nests robbed by snakes and crows.

conservation status

Not threatened; locally common to abundant, and regarded as a pest in grain-growing areas.

significance to humans

At times damages crops, so that farmers often systematically destroy nests at egg or nestling stage.

Red-billed quelea

Quelea quelea

subfamily

Ploceinae

taxonomy

Emberiza quelea Linnaeus, 1758, 'India' = Senegal.

other common names

English: Red-billed dioch; French: Travailleur à bec rouge; German: Blutschnabelweber; Spanish: Quelea de Pico Rojo.

physical characteristics

4.7 in (12 cm); female 0.5–0.9 oz (15–25 g), male 0.6–0.9 oz (16–26 g). Breeding male has face mask, either black or white, with pink or yellowish border; upperparts light brown with dark central streaks, underparts whitish. Bill red, legs pink. Non-breeding male and female lack face mask, gray-brown, streaked upperparts and whitish underparts. Bill red; yellow in breeding females. Juvenile like female.

distribution

Throughout unforested sub-Saharan Africa.

habitat

Open grassland and savanna.

behavior

Highly gregarious, flocks sometimes numbering millions. Movements highly synchronized in flocks. Huge roosts may break tree branches. Migratory with clear seasonal patterns in some regions.

feeding ecology and diet

Primarily small seeds about 0.1 in (2 mm) in diameter, also insects. Drinks regularly, even in arid regions; flocks may sweep over water, drinking on the wing.

reproductive biology

Colonial, monogamous. Breeding activities in colony closely synchronized; eggs and chicks may be abandoned when flock moves on. May breed several times in same season, depending on local food supply. Nest built by male, a thin-walled ball with large side entrance. Lays one to five eggs. Incubation 10–12 days, fledging 11–13 days. Both sexes incubate and feed young. Vast colonies with 500 nests per tree attract hundreds of predators, including eagles, vultures, storks, and carnivorous mammals.

conservation status

Not threatened; considered one of the most abundant bird species. Population can tolerate huge losses, and control efforts have had no noticeable effect on numbers.

significance to humans

Queleas are the major animal pest of cereal crops in Africa, and international programs coordinated by the U.N. Food and Agriculture Organization began in the 1960s. In 1989 losses caused by this bird were estimated at $22 million per annum. However, many other factors contribute to crop losses in Africa. Current research focuses on management rather than attempts to eliminate queleas or reduce their overall numbers. In parts of West Africa, traditional hunters net queleas to pluck, dry, and sell in village markets.

Madagascar fody

Foudia madagascariensis

subfamily

Ploceinae

taxonomy

Loxia madagascariensis Linnaeus, 1766, Madagascar.

other common names

English: Red fody, Madagascar weaver; French: Foudi rouge, Foudi de Madagascar; German: Madagaskarweber; Spanish: Fodi del Madagascar.

physical characteristics

4.7–5.1 in (12–13 cm); both sexes 0.5–0.7 oz (14–19 g). Breeding male bright red, black line through eye, olive-brown wings and tail. Bill black. Female and non-breeding male olive-brown upperparts, gray-brown underparts. Bill horn-brown. Juvenile like female but more buffy in appearance.

distribution

Madagascar; introduced to Amirantes, Comoros, Seychelles, Mauritius, Réunion, St. Helena.

habitat

Open savanna, grassland, forest clearings, and cultivated areas; avoids intact forest.

behavior

Gregarious, foraging in flocks and roosting communally in sugarcane, bamboos, or trees. Solitary and territorial during breeding season.

feeding ecology and diet

Primarily a seed-eater, but also forages for insects in trees, and takes nectar.

reproductive biology

Solitary nests, monogamous. Nest is oval, upright with side entrance near top; woven by male, but female participates from early stages. Lays two to four eggs, breeding season from spring through summer to autumn. Female alone incubates, both sexes feed young. Incubation 11–14 days, fledging 15–16 days.

conservation status

Not threatened; thrives in human-modified habitats and has been introduced successfully to other regions.

significance to humans

An important pest in rice fields in Madagascar. Villagers use traditional cage traps, attempt to scare the birds away from the crops, and destroy nests.

Southern red bishop

Euplectes orix

subfamily

Ploceinae

taxonomy

Loxia orix Linnaeus, 1758, Angola.

other common names

English: Red bishop-bird, Grenadier weaver; French: Euplecte ignicolore; German: Oryxweber; Spanish: Obispo Rojo.

physical characteristics

5.1 in (13 cm); female 0.6–0.9 oz (17–26 g), male 0.7–1.0 oz (21–30 g). Breeding male has red and black plumage, with brown wings and tail. Bill black. Female and non-breeding male sparrowy brown, pale underparts with some streaking. Bill brown. Juvenile like female, buffy edges to feathers before

first molt. Males first molt into breeding plumage in second year.

distribution

Southern Kenya and Uganda south to southern tip of South Africa.

habitat

Tall grassland and cultivation, usually near water.

behavior

Gregarious throughout the year, forming large flocks which feed and roost in association with other seed-eaters. Males return to same breeding localities, often to same territory, in successive years; return rate of females much lower. Both sexes may live more than 10 years in the wild.

feeding ecology and diet

Mainly seed-eating, also takes many insects, particularly when feeding young.

reproductive biology

Colonial and polygynous, often hundreds of males holding territories in a single reed-bed. Male builds a series of nests, oval upright structures with side entrances, typically supported by vertical vegetation. Female lines nest once mated and ready to lay. Old nests not demolished. In courtship male fluffs out plumage, resembling black and red bumble-bee, making short flights towards female. May have up to seven females on territory simultaneously. Breeding season dependent on rainfall; in winter rainfall region of South Africa, starts in winter, ends in early summer. Elsewhere during rainy season, usually summer. Lays one to five eggs (generally three). Incubation 12–13 days, fledging 11–15 days. Female alone incubates and feeds young. Nests often subject to heavy predation, and parasitized by Diederik cuckoo.

conservation status

Not threatened; abundant in many areas, benefits from farming activities and building of dams, which provide additional breeding sites.

significance to humans

Locally an important pest of grain crops; in wheatlands of Western Cape, South Africa, large numbers are killed annually.

Red-collared widow-bird

Euplectes ardens

subfamily

Ploceinae

taxonomy

Fringilla ardens Boddaert, 1783, Cape of Good Hope.

other common names

English: Red-collared widow; French: Veuve noire; German: Schildwida; Spanish: Obispo de Collar Rojo.

physical characteristics

5.1 in (13 cm), with long tail 9.8–11.8 in (25–30 cm); female 0.6–0.7 oz (16–21 g), male 0.7–0.9 oz (20–25 g). Breeding male black with long tail, red collar on upper breast, or red on head and breast; some populations wholly black. Bill black. Female brown with dark streaking above, yellowish eyebrow, underparts buffy and unstreaked; bill brown. Non-breeding adult male like female, but retains black wing feathers. Juvenile with feathers of upperparts broadly edged buff.

distribution

Guinea east to Ethiopia, south to Angola and through Zambia to eastern Zimbabwe and eastern South Africa.

habitat

Open or bushed grassland, cultivated areas; also highland grasslands from 4,900–9,850 ft (1,500–3,000 m).

behavior

Gregarious, forming large roosts even during breeding season, feeding in flocks of 200 birds or more. Often associated with other Euplectes.

feeding ecology and diet

Takes mainly seeds and insects, which may be hawked in the air; rarely berries, nectar.

reproductive biology

Territorial, polygynous, with males well-dispersed. Nest in tall grass, a woven ball with side entrance. Frame started by male, most building done by female. Breeding follows spring or summer rains. Lays two to three eggs; incubation 12–15 days, fledging 14–17 days. Female alone incubates and feeds young.

conservation status

Not threatened; widespread in lowlands, but distinctive montane populations have restricted range.

significance to humans

None known.

Jackson's widow-bird

Euplectes jacksoni

subfamily

Ploceinae

taxonomy

Drepanoplectes jacksoni Sharpe, 1891, Kikuyu, Kenya.

other common names

English: Jackson's widow, Jackson's dancing whydah; French: Euplecte de Jackson; German: Leierschwanzwida; Spanish: Obispo de Jackson.

physical characteristics

5.5 in (14 cm) with long tail 11.4 in (29 cm); female 1.0–1.5 oz (29–42 g), male 1.4–1.7 oz (40–49 g). Breeding male black with

brown wings, curved black tail. Bill steel blue. Female and subadult birds, upperparts dark brown with paler edges to feathers, underparts buffy. Bill brown. Non-breeding adult male like female, but bill dark bluish.

distribution

Central Kenya and northern Tanzania.

habitat

Highland grasslands, above 4,900 ft (1,500 m).

behavior

Gregarious, in flocks when feeding, gathering in communal roosts at night. Breeding areas traditional, and same sites used in successive years.

feeding ecology and diet

Mainly grass seeds, also some insects such as winged termites.

reproductive biology

Polygynous, with a lek mating system. Male displays at dancing ring, a circle of flattened grass about 24 in (60 cm) in diameter, surrounding a central tuft. Facing tuft, male jumps to various heights, up to 1 yd (1 m) into the air while calling. If female lands in ring, courtship and mating may follow. Female builds nest, a ball of woven grass with side entrance, close to the ground in a grass tuft. Nesting area usually about 330 yd (300m) from lek. Lays two to four eggs, usually after main rains. Incubation 12–13 days, fledging 17 days. Female alone incubates, and feeds young on regurgitated grass seeds, not insects.

conservation status

Vulnerable because of limited range, dependence on grasslands which are being altered by agricultural activity.

significance to humans

Flocks may damage grain crops of subsistence farmers.

Thick-billed weaver

Amblyospiza albifrons

subfamily

Ploceinae

taxonomy

Pyrrhula albifrons Vigors, 1831, Algoa Bay, Eastern Cape, South Africa.

other common names

English: Grosbeak weaver; French: Grosbec à front blanc; German: Weißstirnweber; Spanish: Tejedor de Pico Grueso.

physical characteristics

6.7–7.5 in (17–19 cm); female 1.1–1.6 oz (31–45 g), male 1.5–2.1 oz (43–60 g). Breeding male chocolate brown with white forehead, white patch in wing conspicuous in flight. Heavy black bill. White on forehead variable, absent in non-breeding plumage. Female has brown upperparts, underparts white heavily streaked with brown. Heavy yellowish bill. Juvenile like female, more rufous above and buffy below. Bill dull brown.

distribution

Sierra Leone east to southern Sudan, western Ethiopia, south to northern Namibia, northern Botswana, eastern Zimbabwe, and eastern South Africa.

habitat

Reedbeds, cultivated areas, plantation, and forest.

behavior

Gregarious, roosting in reedbeds and breeding there. Flocks move daily up to 19 mi (30 km) to forage when not breeding.

feeding ecology and diet

Fruit, seeds, including large, hard-shelled seeds, and insects. In non-breeding season often forages in forest, feeding on fruit in canopy, also on the ground.

reproductive biology

Colonial, some males polygynous. Nest highly distinctive, woven of very fine reed strips, slung between upright stems, with large side entrance. Built by male; once female accepts nest, male reduces entrance to a narrow circular hole. Female lines nest. Lays two to five eggs in summer. Incubation 14–16 days, fledging 19–22 days. Female alone incubates, feeds young.

conservation status

Not threatened; widespread and range expanding in some areas such as Zimbabwe and South Africa.

significance to humans

None known.

Red-headed weaver

Anaplectes rubriceps

subfamily

Ploceinae

taxonomy

Ploceus (Hyphantornis) rubriceps Sundevall, 1850, Upper Caffraria, near the Tropic = Mohapoani, Rustenburg district, South Africa.

other common names

French: Tisserin écarlate; German: Scharlachweber; Spanish: Tejedor de Cabeza Roja.

physical characteristics

5.9–6.7 in (15–17 cm); female 0.7 oz (20 g); male 0.9 oz (25 g). Breeding male has head, breast, and upper back scarlet, upperparts otherwise gray, underparts white. Wing edged yellow or red in different population; some also have black patch around eye. Bill red, eye red. Female and non-breeding male, scarlet replaced by dull orange on upperparts, below yellow to buff. Bill pink to orange, eye brown. Juvenile like female.

distribution

Senegal east to Somalia, south to Namibia, Botswana, and northeastern South Africa.

habitat

Woodland and acacia savanna, gardens.

behavior

Solitary or in pairs, joins mixed-species flocks. Local movements during dry season. May stay near, and roost in, old nests.

feeding ecology and diet

Mainly insects, spiders, some fruit and seeds. Forages in foliage, on branches and creepers; hawks flying insects.

reproductive biology

Often solitary, monogamous or polygynous; sometimes several males in same tree. Nest built by male, at same site in successive seasons, so that several nests may be close together. Retort-shaped structure, woven from twigs and mid-ribs of leaves with rough appearance; long vertical entrance spout. Suspended from tree, often one in which raptor is nesting. Female lines nest. Lays one to four eggs from late winter through into summer. Incubation 11–13 days, fledging 17 days. Both sexes incubate and feed young, but female contributes more.

conservation status

Not threatened; extensive range, and well-represented in conservation areas.

significance to humans

None known.

Pin-tailed whydah

Vidua macroura

subfamily

Viduinae

taxonomy

Fringilla macroura Pallas, 1764, 'East Indies' = Angola.

other common names

English: King-of-six; French: Veuve dominicaine; German: Dominikanerwitwe; Spanish: Viuda de Cola Aguda.

physical characteristics

4.7–5.1 in (12–13 cm), male with long tail 10.2–13.4 in (26–34 cm); female 0.5–0.6 oz (14–16 g), male 0.5–0.7 oz (14–19 g). Female and non-breeding male, brownish upperparts with broad black stripes on top of head, buff to white underparts. Bill brownish red. Breeding male, black and white with four long, black central tail feathers. Bill bright red. Juvenile plain brown above, buff below.

distribution

Sub-Saharan Africa. Introduced to Hawaii but apparently now extinct.

habitat

Open savanna and grassland, farmland, gardens.

behavior

Male sings from perch, but does not imitate songs of host species. When female arrives, bounces in the air with tail flipping up and down while singing. Aggressive towards other males, but also to other species. Non-breeding birds gregarious, forming small flocks of 20–30 birds, often mixed with other small seed-eaters.

feeding ecology and diet

Mainly seeds, also some insects. Collects most food on the ground. Scratches with backward hops to unearth buried seeds.

reproductive biology

Brood parasite, polygynous. Lays one to two eggs per nest, removing host egg for each egg added. Incubation about 11 days, fledging about 20 days. Most frequent host is common waxbill (Estrilda astrild), also other waxbills and occasionally warblers. Host and parasite young reared together.

conservation status

Not threatened; widespread and common.

significance to humans

None known; annoys those who put out birdseed, as breeding male pin-tailed whydah will attempt to drive all other birds away from feeding site.

Dusky indigobird

Vidua funerea

subfamily

Viduinae

taxonomy

Fringilla funerea de Tarragon, 1847, Natal.

other common names

English: Black widow finch, variable indigobird; French: Combassou variable; German: Purpuratlaswitwe; Spanish: Viuda Variable.

physical characteristics

4.3–4.7 in (11–12 cm), female 0.4–0.5 oz (12–16 g), male 0.5–0.6 oz (14–17 g). Breeding male black, with whitish bill and red legs. Female and non-breeding male buff upperparts with broad black stripes on head, whitish underparts. Juvenile like female but head dark, unstriped.

distribution

Tanzania to eastern South Africa.

habitat

Grassy areas including fringes of cultivation, gardens, and roadsides.

behavior

Male sings from exposed perch for long periods, including song and calls of African firefinch (Lagonosticta rubricata) interspersed with chirping notes. Displays to female in a bobbing flight. In non-breeding season birds associate in flocks, join mixed flocks of other small seed-eaters, and become nomadic.

feeding ecology and diet

Seeds, mainly collected on the ground, including buried seeds dug out by scratching backwards with both feet.

reproductive biology

Brood parasite, polygynous, with males at display sites which females visit for mating. Lays one egg per host nest. Incubation and fledging periods apparently unrecorded. Host is African firefinch, and young of parasite specifically match mouth markings of this species; male indigobirds learn vocalizations of host while in nest, and later incorporate these elements into their own song.

conservation status

Not threatened. Locally common, with an extensive range in Africa.

significance to humans

None known.

Cuckoo finch

Anomalospiza imberbis

subfamily

Viduinae

taxonomy

Crithagra imberbis Cabanis, 1868, East Africa = Zanzibar.

other common names

English: Parasitic weaver; French: Anomalospize parasite; German: Kuckucksfink; Spanish: Tejedor Parásito.

physical characteristics

5.1 in (13 cm); unsexed birds 0.8–0.9 oz (23–26 g). Breeding male, yellow with some streaking, bill black. Non-breeding male, yellowish head, upperparts olive with heavy streaks, bill brown. Female mainly buffy, heavily streaked on upperparts. Short-tailed, with a stubby bill, deep at base. Juvenile resembles female.

distribution

Local in western and central Africa, through eastern Africa to southern Africa.

habitat

Open grassland with scattered trees, wetlands, cultivated lands.

behavior

Little-known and probably nomadic; likely to be overlooked in mixed flocks of seedeaters. When breeding, in pairs or small groups. Male has rasping song in display, defends grassland territory. Non-breeding birds form large roosts in reedbeds, sometimes holding more than 500 birds.

feeding ecology and diet

Seeds, mostly collected while perching on grasses and weeds.

reproductive biology

Brood parasite, mating system not known, but probably polygynous. Lays one to two eggs per nest, removing one or more host eggs. Incubation 14 days, fledging 18 days. Hosts are warblers of the genera Cisticola or Prinia, host young usually trampled in nest, rarely reared with parasite young. Two cuckoo finch young may be reared together.

conservation status

Wide range and not considered threatened.

significance to humans

None known.

Resources

Books

Ali, Salim, and S. Dillon Ripley. Handbook of the Birds of India and Pakistan, Together with Those of Bangladesh, Nepal, Bhutan and Sri Lanka. Vol. 10, Flowerpeckers to Buntings. Delhi: Oxford University Press, 1987.

Bruggers, Richard D., Clive C. H. Elliott. Quelea quelea Africa's Bird Pest. Oxford: Oxford University Press, 1999.

Craig, Adrian J. F. K. "Weaving A Story: The Relationships of the Endemic Ploceidae of Madagascar." In Proceedings of the 22nd International Ornithological Congress, edited by Nigel J. Adams and Robert H. Slotow. Johannesburg: BirdLife South Africa, 1984: 3063–3070.

Fishpool, Lincoln D. C., and Michael I. Evans. Important Bird Areas for Africa and Associated Islands: Priority Sites for Conservation. Newbury and Cambridge, United Kingdom: Pisces Publications and BirdLife International, 2001.

Fry, C. Hilary, Stuart Keith, and Emil K. Urban. The Birds of Africa. Vol. VII. London: Academic Press, in press.

Goodman, Steven M., and John P. Benstead. The Natural History of Madagascar. Chicago: University of Chicago Press, in press.

Sibley, Charles G., and Jon E. Ahlquist. Phylogeny and Classification of Birds. New Haven and London: Yale University Press, 1990.

Stattersfield, Alison J., and David R. Capper. Threatened Birds of the World: The Official Source for Birds on the IUCN Red List. Barcelona and Cambridge: BirdLife International/Lynx Edicions, 2000.

Periodicals

Andersson, Staffan. "Bowers on the Savanna: Display Courts and Mate Choice in a Lekking Widowbird." Behavioral Ecology 2 (1991): 210–218.

Barnard, Phoebe. "Territoriality and the Determinants of Male Mating Success in Southern African Whydahs (Vidua)." Ostrich 60 (1989): 103–117.

Brosset, Andre. "Social Organization and Nest Building in the Forest Weaver Birds of the Genus Malimbus (Ploceinae)." Ibis 120 (1987): 27–37.

Collias, Nicholas E., and Elsie C. Collias. "Evolution of Nest-Building Behavior in the Weaverbirds (Ploceidae)." University of California Publications in Zoology 73 (1964): 1–162.

Crook, John H. "The Evolution of Social Organisation and Visual Communication in the Weaverbirds (Ploceinae)." Behaviour Supplement 10 (1964): 1–178.

Hudgens, Brian R. "Nest Predation Avoidance by the Blue-Billed Malimbe Malimbus nitens (Ploceinae)." Ibis 139 (1997): 692–694.

Nicolai, Jürgen. "Der Brutparasitismus der Viduinae als ethologisches Problem." Zeitschrift für Tierpsychologie 21 (1964): 129–204.

Payne, Robert B. "Brood Parasitism in Birds: Strangers in the Nest." Bioscience 48 (1998): 377–386.

Winterbottom, M., T. Burke, and T. R. Birkhead. "The Phalloid Organ, Orgasm and Sperm Competition in a Polygynandrous Bird: The Red-Billed Buffalo Weaver (Bubalornis niger)." Behavioural Ecology and Sociobiology 50 (2001): 474–482.

Adrian Craig, PhD