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Red-billed quelea

The red-billed quelea (Quelea quelea) is a small, sparrow-sized weaver bird in the family , native to , where it is one of the world's most abundant undomesticated avian with an estimated of 1.5–10 billion individuals, often forming flocks numbering in the millions. Measuring about 12 cm in length and weighing 15–26 g, it features a distinctive cone-shaped bill that is bright red in breeding males, accompanied by a black facial mask in over 75% of males (rarely white), while breeding females have yellowish bills and non-breeding birds of both sexes have red bills, with overall grey-brown plumage and streaked underparts in females and non-breeding males. Highly nomadic and migratory, the follows seasonal rainfall patterns to exploit ephemeral grass seed resources, confining its range to semi-arid and arid savannas, grasslands, and thornveld habitats south of the , typically within 30 km of water sources, and increasingly invading agricultural areas. Primarily granivorous, the red-billed quelea feeds on seeds from wild grasses such as species and cultivated cereals including millet, , , and , consuming up to 15 g of seeds per day per adult bird, with nestlings supplemented by ; this diet drives its opportunistic movements and contributes to its reputation as a significant , capable of devastating crops where flocks converge, with estimated losses exceeding annually in affected regions. Ecologically, it breeds colonially in massive aggregations—sometimes exceeding 10 million pairs—constructing compact, cup-shaped nests from grass in shrubs, reeds, or thorny trees like , with socially monogamous pairs laying 1–5 pale blue eggs per clutch (typically 3) up to three times per year in response to favorable wet conditions, achieving at 9–12 months. Diurnally active and intensely social, these birds roost communally at night in reedbeds or trees and exhibit synchronized flocking behaviors that create dramatic aerial displays, aiding predator avoidance and foraging efficiency across their vast distribution from to . Despite its status prompting control efforts such as aerial spraying with avicides like fenthion and manipulation in countries including and , the red-billed quelea's global population remains stable and is classified as Least Concern by the IUCN as of 2018 due to its extremely large range (25,800,000 km²) and adaptability, though ongoing poses localized threats.

Taxonomy

Etymology and names

The binomial name Quelea quelea originates from the species' initial description by Carl Linnaeus in 1758 as Emberiza quelea, where the specific epithet "quelea" derives from Medieval Latin qualea, referring to a quail, possibly evoking the bird's swarming flocks akin to biblical quail plagues (Numbers 11:31). Linnaeus provided no explicit etymology, and the name's type locality was erroneously listed as India, later corrected to Senegal. In 1850, Ludwig Reichenbach established the monotypic genus Quelea for this species, adopting the epithet as the generic name, a practice that highlights its distinctiveness within the weaver family. The common English name "red-billed quelea" directly describes the prominent red bill coloration in breeding adult males, a key identifying feature that contrasts with the paler bills of females and non-breeding males. names vary regionally, reflecting local languages and perceptions of the as a weaver or . In , it is known as Blutschnabelweber (blood-billed weaver); in , as Quéléa à bec rouge (red-billed quelea) or Travailleur à bec rouge (red-billed worker); and in , as kwelea domo-jekundu (red-billed quelea). Other historical English synonyms include "red-billed weaver," "red-billed dioch," and " finch," with "dioch" likely derived from West African indigenous . No major nomenclatural changes have occurred since Reichenbach's classification, though early misplacements in genera like Loxia () were quickly rectified.

Classification and phylogeny

The red-billed quelea ( quelea) belongs to the family , commonly known as the weaverbirds, within the order Passeriformes. It is one of three species in the genus , which is classified in the subfamily Ploceinae. Three are currently recognized, differentiated primarily by geographic distribution and subtle morphological variations: the nominate Q. q. quelea in northern from and to and the ; Q. q. aethiopica in eastern regions from and to and northern ; and Q. q. lathami in southern areas from and the to , , and . Phylogenetic analyses based on multilocus molecular data, including mitochondrial (ND2, ND3, ATP6) and nuclear (Myo2, TGFb2, Fib7, MUSK) markers, have confirmed the of the Quelea with strong support ( = 1, bootstrap > 80). These studies, sampling all three Quelea , place the in a well-supported (Clade D) sister to the fodies (Foudia) and certain Asian Ploceus within Ploceinae, highlighting its close evolutionary ties to other and weavers. The evolutionary adaptations of species, including the red-billed quelea, are closely linked to their granivorous and nomadic , which facilitate exploitation of ephemeral resources in arid and semi-arid savannas. Large-scale has evolved as an efficient strategy in unpredictable environments, allowing synchronized exploitation of patchy grass while minimizing predation risk through collective vigilance. No significant taxonomic revisions or debates regarding the have emerged from genetic analyses since 2020.

Description

Morphology

The Red-billed quelea (Quelea quelea) is a small bird measuring approximately 11–12 cm in total length and weighing 15–26 g, with males typically slightly heavier at around 19 g and females averaging 18 g. The species exhibits a compact build with a short and relatively broad wings suited to its lifestyle, though specific wingspan measurements are not well-documented and estimated at 18–20 cm based on . Plumage is dull and streaked brown overall in both sexes during the non-breeding season, providing in grassy habitats, with finer barring on the underparts and a pale . In the breeding season, males undergo a striking transformation via pre-nuptial moult, developing a variable black mask extending from the face to the throat—ranging from partial white-faced morphs to full dark masks—and often a pinkish wash on the breast, while retaining the brown back and wings. Females and non-breeding males lack this ornamentation, appearing uniformly drab with no significant dimorphism outside breeding. Juveniles resemble adult females but feature buffier, less streaked , a paler gape, and softer feathering that transitions during their first post-juvenile moult. Sexual dimorphism is pronounced in coloration during , with males' black mask and brighter bare parts serving as key identifiers, while females remain monomorphic year-round. Males are also marginally larger, with wings averaging 68 mm in length (67.6 mm in breeding males vs. 67.2 mm in breeding females) and stouter bills, reflecting subtle size differences that may relate to competitive roles. The bill is short, thick, and conical, measuring around 12 mm in length and up to 10 mm in depth at the base, ideally adapted for husking and cracking small grass seeds with its robust structure. Bill color shows seasonal and sex-specific variation: in non-breeding adults of both sexes due to pigments, with males retaining bills in season while females shift to yellow via hormonal influences like ; transitional mottling may occur. Eye-rings and legs are in non-breeding adults of both sexes; in season, males retain while females shift to yellow.

Vocalizations and displays

The red-billed quelea employs a range of vocalizations for social coordination, particularly in large , and for breeding-related communication. The primary contact call is a sharp, repetitive "chirt chirt," which individuals use to maintain group during or flight. In flight, birds produce a high-pitched "tseep" to signal position within the . Softer, twittering calls, often described as "zee-zee," facilitate pair interactions and are more subdued compared to calls. Males possess a simple song repertoire during the breeding season, delivered from perches near nest sites to advertise territories and attract females. These songs consist of short bursts beginning with preliminary chatter, followed by a warbling section of repetitive phrases, typically lasting a few seconds each. The song derives from modified contact calls and is hormonally influenced, increasing in frequency and intensity as breeding progresses. Acoustic studies indicate these vocalizations play a key role in territory defense within dense colonies, where ambient noise from thousands of birds necessitates clear, repetitive signals for effective communication. Visual displays complement vocalizations, especially in and agonistic contexts. During , breeding males raise and flutter their wings while perched at the nest, combining this with bill raising to showcase their and to prospective mates. Wing vibration, a component homologous to precopulatory behaviors in related , often accompanies these displays. In aggressive encounters, such as disputes, males spread their tails and bob their heads, escalating to physical chases if vocal threats like the "chak" call fail to deter intruders. Paired birds greet each other with quivering, half-spread wings upon returning to the nest, reinforcing bonds amid the colony's chaos. These displays are essential in the noisy environment, where visual cues enhance pair recognition and defense.

Distribution and habitat

Geographic range

The red-billed quelea (Quelea quelea) is native to , with its distribution extending from in the west to and in the east, and southward to , encompassing countries such as , , , , , , , , , and . This broad range covers an extent of occurrence of approximately 25.8 million km², primarily in semi-arid and regions outside dense tropical forests. Three subspecies are recognized based on geographic variation. Q. q. quelea occupies western and central areas from southern and through , , , , , , and northern . Q. q. aethiopica is found in eastern and northeastern regions, including southern , , , northeastern , , , central and eastern , and northwestern and southern . Q. q. lathamii inhabits southern extents from southwestern and southern through , southern , , , western , (excluding arid coastal areas), and central, southern, and eastern . The species supports an estimated post-breeding population of 1.5 billion individuals, rendering it the world's most abundant wild . Recent studies from 2024 and 2025 document range shifts driven by changes, including expansion into agricultural fringes; for example, increased cropland and settlements in Tanzania's Chemba District have altered local distributions, while opportunistic movements into southern Africa's cereal-growing areas, such as the of , have been linked to and developments. Its nomadic movements further influence the effective occupancy within this range.

Habitat requirements

The red-billed quelea primarily inhabits semi-arid savannas, dry grasslands, and thornveld regions south of the , often mixed with scattered woodlands that provide roosting opportunities. These environments typically feature subtropical or tropical dry conditions, including dry savannas, shrublands, and grasslands. The species is also commonly found in human-modified landscapes such as arable lands and pasturelands, demonstrating adaptability to altered ecosystems. Key habitat requirements include access to abundant grass seeds for sustenance, proximity to water sources like rivers or stagnant pools within about 30 km, and suitable nesting sites in reedbeds, thorny shrubs, or trees. Annual rainfall between 300 and 800 mm supports breeding cycles by promoting grass seed availability, though the bird persists in drier areas with reliable water access. While tolerant of farmlands and irrigated agricultural schemes, the red-billed quelea avoids dense forests such as woodlands and rainforests, as well as true deserts and high-altitude zones above 3,000 m. This overlap with agricultural areas frequently results in conflicts with human activities due to foraging. Studies from the 2020s highlight the impacts of driven by changes, including agricultural expansion and settlement growth, which reduce suitable roosting and breeding sites. For example, in Tanzania's Chemba district, forested lands declined by 17.64% and coverage by 5.6% between 2002 and 2022, while increased by 16.89%, contributing to localized population declines and diminished suitability.

Behavior and ecology

Social behavior and movements

The red-billed quelea (Quelea quelea) is highly gregarious, forming enormous that can number in the millions for roosting, , and , with roost sizes occasionally reaching tens of millions of individuals. These massive aggregations facilitate about food sources, as birds observe and follow successful from communal roosts, enhancing group efficiency in locating resources. formation is synchronized, with birds departing roosts in dense streams, often covering daily distances of up to 30 km to foraging sites and water sources. Within these flocks, a hierarchical emerges, particularly evident in colonies where males establish dominance through aggressive interactions and displays, often correlated with the of coloration on their bills as an indicator of and . Allopreening, or mutual grooming between individuals, serves both affiliative and agonistic functions, occurring frequently in non- flocks and helping maintain social bonds while sometimes reinforcing dominance hierarchies. behavior is also observed, with groups collectively harassing potential threats, such as predators near colonies, to deter attacks. The species exhibits nomadic movements driven by rainfall patterns and the availability of grass seeds, with populations shifting across to exploit seasonal resources following the onset of rains. Ringing recoveries from the SAFRING database reveal regional fidelity, with many birds recaptured near ringing sites (e.g., within 4 km at Barberspan, ), yet capable of long-distance displacements exceeding 1,000 km, including movements up to 2,545 km between and the Democratic Republic of Congo. Daily flights typically span tens of kilometers, but nomadic patterns allow for broader seasonal migrations in response to resource pulses. Recent analyses (2023–2025) of climatic variability, including rainfall and temperature fluctuations, indicate shifts in distribution and movement intensity within key wetlands like Hadejia-Nguru in northern , where altered patterns influence flock concentrations and regional fidelity. These responses underscore the species' adaptability to environmental changes, with nomadic strategies buffering against unpredictable seed availability amid climate variability.

Breeding biology

The red-billed quelea exhibits opportunistic breeding strongly linked to the onset of rainy seasons across its range in , typically initiating nest-building 4–6 weeks after sufficient precipitation (300–800 mm annually) stimulates grass seeding for food availability. This timing allows for 1–3 clutches per year, facilitated by itinerant breeding where pairs or groups relocate hundreds of kilometers to exploit successive rainfall events, as documented in marked individuals moving from southwestern to the Awash Valley. Breeding commences at 9–12 months of age, with pair bonds forming for the season and males using vocal and display behaviors to attract mates near colony sites. Breeding occurs in massive, synchronous colonies spanning 1–100+ hectares, often containing 10,000–100,000 nests or more, with densities reaching 1,000–1,500 nests per tree in thorny acacias, reeds, or shrubs. Males construct oval, roofed nests woven from strips of green grass, featuring a side entrance and compact structure for . Each female lays a clutch of 1–5 whitish-blue eggs, typically averaging 3, with the full reproductive cycle from egg-laying to fledgling independence spanning about 42 days. Incubation lasts 10–12 days, primarily performed by the female, who covers the eggs at night and during midday heat, while the male guards the and supplies food; daytime warming from solar radiation (34–37°C) supplements this in hotter regions. Both parents feed the hatchlings, initially regurgitating protein-rich and soft seeds to support rapid growth, transitioning to harder seeds as nestlings develop; females typically provide more frequent feeds (about 6 per day) than males (4–5 per day) in observed . Chicks at 12–14 days, though early departure (around 11–12 days) occurs in overheated nests, with full independence shortly after; the complete nestling period thus enables quick colony turnover to align with ephemeral resources. Chick mortality is high, often exceeding 80% in some colonies due to predation by , weather extremes, from density-dependent food shortages, and nest abandonment if one parent is lost during . Overall breeding success varies, with up to 95% but only 80–90% fledging rates when both parents survive, dropping sharply if the is removed early. Across Q. q. aethiopica in eastern , Q. q. lathami in southern regions, and nominate Q. q. quelea in the west and patterns are broadly similar, though colony sizes and timing shift with local rainfall variability. Recent studies post-2020 indicate stable or increasing success in some areas despite from agriculture and land-use changes, as queleas adapt by exploiting irrigated crops and shifting to denser, human-modified vegetation for nesting.

Foraging and diet

The red-billed quelea (Quelea quelea) is predominantly granivorous, with its diet consisting primarily of seeds from annual grasses such as sorghum (Sorghum spp.) and millet (Pennisetum spp.), which form the bulk of its intake outside the breeding period. Insects, including beetles, grasshoppers, and termites, supplement this granivorous base, particularly during the breeding season when protein demands increase for egg production and nestling provisioning; in one study from Ethiopia's Awash River Basin, insects comprised about 10.8% of the diet during breeding. This shift supports higher nutritional needs, as females build protein reserves in flight muscles prior to laying. Foraging typically occurs in large, dense flocks on the ground or in low , where employ a "roller feeding" , moving collectively to quickly reduce density in a localized area. The ' stout, conical is adapted for handling small s (1-2 in ), enabling efficient husking by perching on grass stems or heads to extract and process them, though it cannot manage larger grains like whole kernels exceeding 4.8 . Males possess slightly larger bills than females, potentially aiding in and contributing to subtle in foraging efficiency. This flocking behavior enhances social coordination but also amplifies the scale of . An individual red-billed quelea consumes approximately 2-3 grams of unhusked per day, a modest intake that scales dramatically in swarms to enable rapid exploitation of available resources. Seasonal variations in and reflect environmental cues and : during the early , birds target small, fallen grass collected from the ground, shifting to larger as supplies dwindle later in the season. In the rainy season, focuses on ripening grass , with migrations tracking these patches, while become more prominent. Nutritional studies highlight preferences for ripe , which offer optimal energy content (around 18.4 kJ/g for grass ), allowing birds to build fat reserves before lean periods and sustain high reproductive output.

Predators, parasites, and mortality

The red-billed quelea (Quelea quelea) is preyed upon by a diverse array of predators, including raptors such as the (Falco biarmicus) and (Aquila rapax), as well as snakes and various mammals that target nests and roosts. Over 80 species of avian predators have been documented exploiting quelea colonies, particularly during breeding seasons when flocks aggregate in large numbers. Humans also contribute to predation through traditional hunting practices in some regions, though this has limited impact on overall population levels. The formation of massive flocks provides a dilution effect, where the risk to any individual bird decreases as group size increases, thereby reducing per capita predation rates despite the species' abundance. Parasites of the red-billed quelea include both ectoparasites and endoparasites. Ectoparasites such as chewing lice (Phthiraptera) are commonly found on queleas, with species like Machirinius quelea and Columbicola reported in West African populations. Ticks and other external parasites have also been observed feeding on birds during roosting. Endoparasites encompass nematodes, including Ascaris species, which are prevalent in gastrointestinal tracts, alongside protozoans like Giardia. Haematozoan parasites, such as Haemoproteus and Plasmodium, show variable infection rates, often low in queleas compared to other birds. Disease outbreaks include avian pox (avipoxvirus), with seroprevalence reaching 80% in some sampled populations, leading to nodular lesions and potential mortality in affected individuals. Mortality in red-billed queleas arises from multiple natural factors, including weather extremes like high winds that can cause mass deaths by dashing birds against structures, and starvation during prolonged dry seasons when seed availability declines. Annual adult mortality is estimated at around 50%, corresponding to survival rates of approximately 50%, based on ringing recoveries during periods of natural population dynamics. Recent research in 2024 has highlighted increasing parasite loads, particularly gastrointestinal nematodes and haemosporidians, in queleas inhabiting altered habitats due to environmental changes, potentially exacerbating mortality in fragmented ecosystems. Additionally, queleas serve as key prey in African food webs, supporting predator populations across savanna regions.

Human interactions

Agricultural impacts

The red-billed quelea (Quelea quelea) is a major pest of small-grain crops such as , , and millet across , where its granivorous diet enables it to target ripening seeds during breeding seasons. Swarms of these birds, often numbering in the millions, can descend on fields and cause destruction ranging from 20% to complete crop loss in affected areas, particularly in hotspots like the and . Annual economic losses from this damage are estimated at $50–100 million USD, threatening and farmer livelihoods in cereal-dependent economies. Historical outbreaks intensified in the and , with peak colony formations and crop raids reported across southern and eastern , leading to widespread agricultural devastation during periods of favorable rainfall and grass seed availability. In recent years, particularly in the , impacts have escalated due to expanding farmlands and increased production, which have boosted quelea populations by providing more and foraging opportunities; for instance, a 2022 invasion in affected eight regions, ravaging and fields with flocks estimated at 21 million birds. Similar events in in 2023 destroyed up to 40% of harvests in drought-stressed areas. Despite these negative effects, red-billed queleas play a positive role in natural grasslands by consuming large quantities of seeds, thereby providing incidental biological control of invasive grasses and aiding balance outside cultivated areas. This seed predation on wild species like and helps maintain open habitats but becomes problematic when croplands encroach on their native foraging grounds.

Management and control

The management and control of red-billed quelea populations are primarily driven by their substantial impacts on small grain crops such as , millet, and across , where swarms can cause losses equivalent to millions of dollars annually. Control strategies emphasize targeting breeding colonies and roosting sites to disrupt , with a shift in recent decades toward integrated approaches that minimize environmental harm. Chemical controls have historically relied on aerial or ground spraying of avicides, particularly fenthion (Queletox®), which was effective against large swarms but highly toxic to non-target organisms, including and mammals, leading to secondary poisoning and disruption. Efforts to phase out fenthion have been ongoing since the early due to its toxicity, with international bodies like the FAO and promoting alternatives, though it remains in use in several countries as of 2025. These alternatives achieve variable mortality rates, often with delayed effects that reduce immediate swarm reduction to 50-70%, though they pose lower risks to . Physical methods include netting at breeding colonies, which can capture thousands of birds daily—such as over 19,000 in trials at Iyoli, —and chick harvesting using poles or hooks, yielding up to 3.78 kg of birds per person per hour. Frightening devices, including explosions and fire-bombs, have been deployed in countries like and , while emerging technologies such as laser scarecrows and AI-powered drones for monitoring and repellent application were tested in the , with drones covering up to 40 hectares in 30 minutes. These methods limit damage to 0-4% in protected fields compared to 2.7-18.8% without intervention, though they are labor-intensive and less scalable for massive swarms. Biological approaches focus on promoting natural predators through initiatives like trained falcons in , , and , which deter without chemicals, and (IPM) programs led by the FAO since the 1990s, incorporating cultural practices such as early planting and resistant cultivars alongside trapping. IPM has demonstrated reductions in crop damage from 85% to 30% using repellents like mesurol in trials. In 2024-2025, environmental concerns over residues—such as fenthion's persistence up to 188 days—have accelerated the of eco-friendly technologies, with studies in reporting 70-90% swarm reductions through combined drone surveillance and nest destruction, saving up to 500,000 tonnes of crops. For example, in from December 2024 to March 2025, aerial operations destroyed over 50 million birds, saving approximately 500,000 tonnes of crops using avicides like Bathion 60% ULV and Quelea Tox. Challenges persist, including high costs and by , necessitating ongoing monitoring and international collaboration.

Cultural and ecological roles

In various African communities, particularly in Tanzania's Chemba and Kondoa districts, the red-billed quelea serves as a vital source, with people harvesting adults and chicks using nets and traps to provide affordable protein during seasonal scarcities. These birds are consumed roasted or fresh, often sold at markets or bus stops for low prices such as TZS 100–300 (approximately USD 0.06–0.18) per portion, and are valued for their nutritional benefits, including high energy content that supports livelihoods and even perceived enhancements to physical vitality. Harvesting yields significant income, with households earning up to USD 18 weekly from live birds or USD 15 daily from roasted ones, fostering positive cultural attitudes toward the despite its pest reputation. The red-billed quelea also holds minor roles in , where it is occasionally kept and bred in captivity by enthusiasts in large aviaries to mimic its flocking behavior, though such practices remain limited due to the bird's nomadic nature. Ecologically, it contributes to grassland regeneration through , as undigested grass seeds pass through its digestive system and are deposited across landscapes, promoting plant diversity in semi-arid habitats. As a primary prey item, it supports a wide array of predators, including over a dozen species such as the (Falco biarmicus), (Aquila rapax), and (Circus macrourus), along with and storks, thereby bolstering food webs in sub-Saharan ecosystems. Its massive, responsive populations further serve as an indicator of , reflecting rainfall patterns and productivity. The species' conservation status is classified as Least Concern by the IUCN, with a stable global population estimated in the billions across its vast 25.8 million km² range, showing no evidence of significant declines. However, recent studies highlight the need for ongoing of climate impacts, such as shifting rainfall and temperature variability altering breeding distributions in regions like northern Nigeria's Hadejia-Nguru Wetlands and Tanzania's Chemba District, potentially affecting long-term habitat suitability.

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