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Quelea

Quelea is a of three of small, gregarious birds in the weaver family , endemic to : the (Quelea quelea), the cardinal quelea (Q. cardinalis), and the red-headed quelea (Q. erythrops). These sparrow-sized birds, typically 11–13 cm in length, feature short tails, stout seed-cracking bills, and streaked plumage that provides in grasslands; males often display vibrant colors during seasons. Primarily granivorous, they form enormous nomadic flocks numbering in the millions, undertaking irregular migrations in response to food availability and rainfall across savannas, semi-arid regions, and cultivated areas. The stands out as the most abundant wild bird species globally, with post-breeding populations estimated at 1.5 billion individuals, enabling massive breeding colonies that can span hundreds of hectares. This species, along with its congeners, is notorious as an agricultural pest, particularly targeting small-grain crops like , millet, and , leading to substantial economic losses and necessitating ongoing control efforts in affected regions. While less numerous, the cardinal quelea favors arid thornveld habitats in southern and eastern , and the red-headed quelea occurs in wetter grasslands of central and eastern , both exhibiting similar and breeding behaviors.

Taxonomy and systematics

Etymology and history

The genus name Quelea derives from the Latin "quelea," denoting a kind of , a term coined by as the specific epithet for the (Quelea quelea) in his 1758 work , where he classified it as Emberiza quelea within the buntings and erroneously designated as the type locality. In 1850, German ornithologist Ludwig Reichenbach established the Quelea and reassigned the species to it within the weaver family , recognizing its affinities with other weavers rather than buntings. Historically, Quelea species have been identified as weaver birds confined to , with early scientific accounts highlighting their widespread distribution across grasslands and savannas. Their recognition as significant agricultural pests emerged prominently in 19th-century colonial reports from administrations in , which documented massive flocks devastating crops like millet and in regions such as southern and eastern . The (Q. quelea), the most studied species in the genus, comprises three recognized differentiated primarily by variations and geographic range: the nominate Q. q. quelea (Linnaeus, 1758), distributed in West and from to and northern ; Q. q. aethiopica (Sundevall, 1850), found in from to and extending south to and ; and Q. q. lathamii (Smith, A., 1836), occurring in Central and from to and .

Phylogeny

The genus Quelea is placed within the family , specifically in the Ploceinae, which encompasses the true weavers characterized by their nest-weaving behaviors. Molecular analyses confirm the of , with Quelea nested in D alongside genera such as Foudia, Ploceus (Asian species), Malimbus, Anaplectes, and Euplectes. This placement is supported by multilocus datasets including three mitochondrial genes (ND2, ND3, ATP6) and four nuclear introns (Myo2, TGFb2, Fib7, MUSK), yielding strong Bayesian posterior probabilities (PP = 1) and maximum-likelihood bootstrap values (>80%) for the family's internal structure. Within Quelea, the three species—red-billed quelea (Q. quelea), cardinal quelea (Q. cardinalis), and red-headed quelea (Q. erythrops)—form a monophyletic with robust support from both mitochondrial and combined datasets. The genus Quelea is strongly supported as to Foudia (the fodies of the islands), with this combined to the Asian Ploceus species; nuclear data weakly suggest potential of Quelea relative to Foudia, but this lacks strong backing. These relationships indicate an African origin for the lineage, followed by dispersal events to and the region, consistent with divergence estimates placing the typical weavers (Clade D) in the (~7.53–10.35 Ma). The classification of Quelea has remained stable, with no major phylogenetic revisions reported through 2025. Historically, the distinct conical morphology of Q. cardinalis (and sometimes Q. erythrops) prompted proposals to separate them into a monotypic or paired Queleopsis, but this taxonomic change has not been widely adopted in modern classifications, which retain all three in Quelea based on molecular evidence.

Species

The Quelea comprises three recognized of small weaver birds endemic to , all belonging to the family . These form a monophyletic within the , characterized by their gregarious nature and adaptations to habitats. The (Quelea quelea) is the most widespread and abundant in the genus, occurring across from to and south to . It is renowned for its enormous population sizes, with peak post-breeding estimates reaching approximately 1.5 billion individuals, making it one of the most numerous wild bird globally. Three are recognized: the nominate Q. q. quelea in , Q. q. aethiopica in , and Q. q. lathamii in , differing primarily in male breeding details such as facial markings and underpart coloration. The cardinal quelea (Quelea cardinalis) has a more restricted distribution in East and Central , including countries such as , , , the , and . This species measures about 10 cm in length and is distinguished by the bright red head of breeding males, which extends onto the throat and variably onto the breast, contrasting with streaky brown upperparts. The red-headed quelea (Quelea erythrops) occupies a broad range across , recorded in over 40 countries including , , , , , , , , , , , , , , , , , , and . It features a rufous-red head in breeding males, often with blackish barring on the throat, and is notable as a specialist on seeds among its grass seed diet, occasionally becoming a in agricultural areas. All three species are classified as Least Concern on the , with no status changes reported as of 2025, reflecting their large populations and extensive ranges despite local agricultural conflicts.

Physical description

Size and morphology

Quelea birds are small weavers in the family , typically measuring 11–13 cm in length and weighing 11–26 g, exhibiting a compact, - or finch-like build across the . Species-specific variations include slightly smaller sizes in the cardinal quelea (Quelea cardinalis) at around 11 cm and 11–15 g, while the (Q. quelea) and red-headed quelea (Q. erythrops) reach up to 13 cm and 26 g. The is characteristically short, stout, and conical, adapted for cracking hard , with a robust structure common to all species for efficient granivory. In the cardinal quelea, the is black, while in the it is red in breeding adults, with non-breeding adults (both sexes) having a that fades post-breeding and breeding females showing a yellowish ; the red-headed quelea shares a similar red form, paler in females. Wings are short and rounded, facilitating agile, maneuverable flight essential for navigation within dense flocks, complemented by a short tail that enhances stability during rapid turns. Legs are strong and sturdy, with adaptations for secure perching and hopping on thorny vegetation or ground substrates. in size is subtle, with males generally larger and heavier than females in species like the red-headed quelea, where males average 18–26 g compared to 15–22 g for females.

Plumage and sexual dimorphism

Queleas display marked in , most evident during the breeding season when males develop vibrant colors while females remain relatively drab. In non-breeding , individuals of both sexes across the typically exhibit streaked brown upperparts and pale underparts, providing a uniform, inconspicuous appearance. Breeding males, however, undergo a pre-nuptial molt to acquire bright red and black elements, such as facial masks, which are absent in females. The (Quelea quelea) exemplifies this pattern. Breeding males feature a red bill, a prominent black , and polymorphic head and neck coloration that varies continuously from black, yellow, pink, purplish, to white, driven partly by pigmentation. Females are duller overall, with streaked brown similar to non-breeding males and a yellowish bill during breeding. Non-breeding adults of both sexes have light underparts, brown-streaked upperparts with yellow-edged , and a reddish bill, though the intensity fades post-breeding. In the cardinal quelea (Quelea cardinalis), breeding males develop a striking red head and breast that does not extend to the streaked , paired with a black bill. Females lack these reds, instead showing a yellowish face accented by superciliary stripes and a yellowish throat, making them harder to distinguish from non-breeding males in the field. Non-breeding males resemble females but retain faint red traces on the head. The red-headed quelea (Quelea erythrops) shows somewhat less pronounced dimorphism. Breeding males have a bright head extending to the with black barring on the , while females exhibit a duller face with yellowish tones, a , and white . Both sexes share similar non-breeding of streaky brown upperparts and pale underparts, with females featuring a short dark moustachial stripe and a paler bill. Following , queleas undergo a post-nuptial molt that transitions males from colorful plumage to the dull non- form, often beginning late in the season and sometimes interrupting to align with reproductive cycles. This molt supports the birds' nomadic by restoring camouflage-effective feathers.

Distribution and habitat

Geographic range

The genus Quelea is endemic to , with all three distributed across the continent south of the Desert but generally avoiding dense rainforests, elevations exceeding 1,500 m, and extreme arid . Their ranges overlap extensively in and regions, reflecting adaptations to open, seasonally variable environments. The (Quelea quelea) has the broadest distribution, occupying a near-continuous range from in the west to in the south, spanning 37 countries including , , , , , , , , , , Côte d'Ivoire, , , , , , , , , , , , , , , , , , , , , , , , , , and . It was introduced to the island of in 2000, where a self-sustaining population has established. The cardinal quelea (Quelea cardinalis) is more restricted to eastern and central Africa, occurring in 11 countries such as , , , , , , , , , , and . The red-headed quelea (Quelea erythrops) exhibits a wide but patchy distribution across , present in 37 countries from and in the west to in the east, extending south to and including , , , , , , , , Côte d'Ivoire, , , , , , , , , , , , , , , , , , , , São Tomé e Príncipe, , , , , and . Range expansions have been documented for the in , including historical expansion into the province attributed to changes in agricultural practices, as well as more recent steady expansion into the province as of 2024–2025.

Habitat preferences

Quelea birds, comprising the species Quelea quelea (), Q. cardinalis (cardinal quelea), and Q. erythrops (red-headed quelea), predominantly favor open, semi-arid to mesic landscapes across . These s include grasslands, thornbush savannas, and floodplains, where homogenous vegetation supports large flocks and colonies. The , the most widespread species, thrives in dry thornveld and bushed grasslands with scattered trees, often near seasonal water sources that facilitate and . Similarly, the cardinal quelea prefers rank grass and wooded grasslands in drier regions, while the red-headed quelea occupies taller grasslands, particularly in areas of higher rainfall below 1,400 m elevation. Roosting sites are typically selected for protection and density, such as reedbeds along edges, thorny bushes, or thorny trees, allowing massive aggregations—sometimes exceeding a million birds—for safety in numbers. Adaptations to these environments include a reliance on proximity to ephemeral bodies for and nest construction during seasons, as well as for arid conditions outside of reproduction. Quelea avoid dense forests, extreme deserts like Namibia's coastal dunes, and high-altitude zones above 1,500 m, where vegetation structure and climate limit seed availability and nesting opportunities. Human activities have expanded suitable habitats through irrigation in agricultural zones, drawing quelea into farmlands with cereal crops like sorghum and millet, thereby increasing their overlap with human-modified landscapes. This adaptation to irrigated savannas and cultivated floodplains has facilitated population booms in regions previously marginal for the birds.

Behavior and ecology

Diet and foraging

Queleas are primarily granivorous, feeding mainly on seeds of wild grasses such as Echinochloa spp., Sorghum spp., Panicum spp., Digitaria ciliaris*, Dactyloctenium aegyptium, and Setaria spp., which constitute the bulk of their diet in natural habitats. They also consume seeds from cultivated cereals including sorghum, millet, and rice when available, though wild grass seeds predominate outside breeding periods. Nestlings receive a protein-rich supplement of insects, such as beetles (Coleoptera) and grasshoppers (Orthoptera), mixed with soft grass seeds during their early development. Foraging occurs predominantly on the ground or in low vegetation, where queleas glean and strip seeds rapidly in enormous flocks numbering in the millions, enabling them to deplete seed patches efficiently. These flocks exhibit nomadic behavior, tracking seasonal seed availability that surges after post-rain grass growth in semi-arid regions. Their conical bills, adapted for cracking small seeds, facilitate this ground-based feeding strategy. Species within the genus show dietary preferences: the (Quelea quelea) targets small-grain cereals like and millet alongside wild grasses, while the red-headed quelea (Q. erythrops) specializes on panicles in habitats. In contrast, the cardinal quelea (Q. cardinalis) is less oriented toward crops, favoring grass seeds such as and spp., with occasional like caterpillars and from flowers. Daily foraging peaks at dawn and dusk, with birds feeding intensively during cooler periods and resting midday in shaded areas; they drink water 2–3 times per day, often congregating at waterholes shortly after leaving roosts.

Breeding biology

Quelea species exhibit opportunistic breeding closely tied to rainfall patterns, initiating nest-building 4–9 weeks after the onset of sufficient (300–800 mm annually) that promotes grass seed availability. In regions like , breeding typically occurs from November to March during the rainy season, while in , up to 3–4 broods per year are possible under favorable conditions. This rapid reproductive cycle allows populations to exploit ephemeral resources, with birds reaching breeding age at 9–12 months. Breeding occurs in massive, highly synchronous colonies that can span 1–100 hectares and contain millions of birds, with nest densities reaching up to 35,000 per hectare in dense thickets or beds. Males construct retort-shaped nests woven from grass strips, typically oval or flask-like structures 100–130 mm wide, suspended from thorny branches for protection; these are built over 2–3 days solely by males to establish territories limited to the nest vicinity. Colonies exhibit strong synchrony, with over 90% of eggs hatching within a 3-day window, facilitating communal defense. Mating displays involve males in facial and breast performing songs—rapid trills, whistles, and —while fluttering wings to attract to incomplete nests. lay clutches of 2–4 (average 3), influenced by their protein reserves, with incubation lasting 9–10 days primarily by the female at night and heating by day. Both parents regurgitate food to nestlings, which after approximately 14 days, though high mortality from predation affects up to 50% of young due to vulnerability in dense colonies. hatching success exceeds 95%, but nestling losses are density-dependent, often from or predators like snakes and birds. The prolific nature of Quelea breeding enables a single large colony to produce several million fledglings per event, with one recorded colony in producing about 4.8 million in 1978, driven by the scale of colonies and multiple annual attempts, underscoring their potential for rapid population explosions.

Migration and social behavior

Queleas exhibit irregular, rainfall-driven nomadism rather than fixed migratory routes, with movements triggered by the onset of wet seasons and the availability of maturing grass seeds. In the (Quelea quelea), birds depart from dry-season concentration areas toward regions where rains began weeks earlier, often covering distances of 300–600 km southward in during the early-rains migration. These nomadic patterns allow flexible exploitation of ephemeral food resources across , with post-breeding dispersal leading to widespread distribution without predictable annual cycles. Flocks of queleas can reach enormous sizes during non-breeding periods, numbering in the millions in the , forming dense, synchronized wheeling formations that enhance predator evasion through collective vigilance and early warning signals. Roosting occurs in large communal groups, occasionally mixed with other species, where birds perch tightly in for safety. These massive aggregations facilitate on resource locations, with birds at the rear of feeding flocks flying overhead to access fresh areas when seeds deplete. Non-breeding social structure in queleas is hierarchical, particularly among males, involving agonistic interactions and individual that maintain within flocks. Vocalizations play a key role in coordination during movements and roosting, while allopreening behaviors strengthen bonds in communal settings. The cardinal quelea (Q. cardinalis) and red-headed quelea (Q. erythrops) are less nomadic than their red-billed counterpart, tending to remain more resident within core ranges in eastern and , though they still form large flocks numbering in the thousands after rains without strict seasonal patterns.

Relationship with humans

Agricultural impact

The (Quelea quelea) is widely regarded as Africa's most significant avian agricultural pest, often dubbed the "feathered " due to its massive flocks that can devastate cereal crops such as millet, , and . These birds target ripening grains, leading to crop losses that can reach up to 100% in small-scale farms during intense invasions, particularly in semi-arid regions where predominates. A single flock can strip fields bare within hours, consuming up to 10 grams of grain per bird daily, which compounds the vulnerability of rural communities reliant on these staples. The red-headed quelea (Quelea erythrops) poses a specialized to rice production, focusing on paddies where it damages panicles and reduces yields by feeding on developing seeds. This species is particularly problematic in irrigated schemes across eastern and , where flocks descend on crops during the vulnerable milky-grain stage, exacerbating losses in flood-prone areas. Across , quelea invasions affect more than 20 countries, translating to economic damages exceeding USD 88.6 million (as of 2025) and severely undermining in hunger-prone regions. Historical records from colonial-era reports, including early accounts, document these impacts on farming, while contemporary outbreaks are intensified by variability, such as erratic rainfall patterns that expand breeding habitats and drive flocks into new agricultural zones.

Pest control methods

Traditional methods for managing Quelea populations have centered on lethal interventions targeting large aggregations at breeding colonies and roosts. of the avicide fenthion has been the primary technique since the mid-20th century, enabling rapid destruction of millions of birds in a single operation to curb crop threats. However, fenthion's to non-target species, including aquatic life, mammals, and beneficial birds, prompted its listing in Annex III of the in May 2025 (with amendments entering into force on 22 October 2025), subjecting ultra-low-volume formulations to the Prior procedure for . Complementing spraying, night fire-bombing involves detonating fuel-laden explosives in colonies after , incinerating roosting birds and nesting to disrupt . Contemporary approaches emphasize non-lethal and sustainable alternatives to reduce ecological . In and other East African regions, trained and hawks are deployed to intimidate and disperse Quelea flocks from fields, leveraging natural predation cues without killing birds. Physical barriers like netting over paddies, combined with scaring devices such as makers or reflective tapes, provide localized protection by denying access to ripening grains. Emerging technologies include drone-based systems equipped with lasers or sonic emitters to herd flocks away from crops over larger areas, offering precision and minimal environmental footprint. Harvesting Quelea through traditional traps or nets also serves as a control measure, converting pests into a valuable protein source for local communities facing nutritional deficits. Regional coordination bolsters these efforts through specialized organizations. The Control Organisation for Eastern Africa (DLCO-EA) and the International Red Locust Control Organisation for Central and (IRLCO-CSA) monitor Quelea movements via aerial surveillance and execute joint operations, including avicide applications and habitat assessments across member states. A 2025 FAO webinar highlighted the shift to non-chemical methods, stressing their role in safeguarding raptors from secondary poisoning while promoting . Overall, have yielded limited reductions, often below 20% annually, owing to the ' explosive breeding rates and nomadic patterns that evade complete eradication. Chemical tactics, in particular, have inflicted substantial harm on non-target avifauna, with residues causing widespread mortality through contaminated prey.

Conservation and cultural significance

The three species in the genus Quelea—the red-billed quelea (Quelea quelea), red-headed quelea (Quelea erythrops), and cardinal quelea (Quelea cardinalis)—are all classified as Least Concern on the IUCN Red List, reflecting their large, stable populations across sub-Saharan Africa. These birds are among the most abundant wild avian species globally, with estimates for the red-billed quelea alone exceeding one billion individuals. Populations appear to have increased substantially since the mid-20th century, largely due to the expansion of agriculture, which has created abundant cereal crop habitats and food resources that support larger flock sizes and breeding success. Despite their abundance, queleas face indirect threats from measures, particularly the use of avicides like fenthion, which can non-target species such as endangered raptors that feed on quelea carcasses. In , for instance, such operations have led to documented deaths of , including species like the eagle, highlighting risks to . poses another emerging threat by altering rainfall patterns and vegetation cycles, which disrupt quelea breeding sites and foraging areas in semi-arid regions, potentially leading to shifts in distribution and local population fluctuations. Beyond their status, queleas play positive ecological roles in ecosystems. They contribute to by consuming and excreting seeds of grasses and other , aiding in vegetation regeneration across savannas and farmlands. During the nestling phase, adults feed their young large quantities of , helping control pest populations such as locusts and grasshoppers in localized areas. Large roosts also deposit that enriches soil nutrients, acting as a natural in nutrient-poor regions. Culturally, queleas hold varied significance in African societies. In parts of , such as around , they are traditionally captured and consumed as , providing a protein source during outbreaks. The spectacular aerial displays of massive flocks—sometimes numbering millions—serve as tourist attractions in protected areas like South Africa's and Namibia's , drawing visitors to witness these natural phenomena. In some traditions, large quelea swarms symbolize abundance or impending change, reflecting their dual role as harbingers of plenty in good seasons or destruction during crop raids.

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