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Hoopoe

The (Upupa epops), a in the family Upupidae and order , is a medium-sized renowned for its vibrant cinnamon-orange , bold black-and-white barred wings and tail, and a conspicuous fan-like erectile that it raises during displays. Measuring 25–32 cm in length with a of 44–48 cm and weighing 46–89 g, it possesses a slender, downcurved bill ideally suited for foraging from the ground. This distinctive , whose name derives from its onomatopoeic "hoop-hoop" call, inhabits open woodlands, savannas, grasslands, and lightly vegetated areas across and , where it plays a key ecological role as an . Widespread in distribution, the breeds throughout most of continental Eurasia south of 53°N latitude and across much of , excluding high mountains, dense tropical forests, and extreme deserts, with a global population estimated at 5,000,000–10,000,000 individuals classified as Least Concern by the IUCN. Populations in and are largely migratory, wintering in the south of 35°N, while those in , , and tend to be sedentary or nomadic. It favors habitats with soft, sandy soils for and vertical structures like cavities, rock crevices, or even building walls for nesting, though it avoids frozen ground in winter. Behaviorally, the is diurnal and often solitary or in pairs, spending much of its time on the ground probing for prey such as orthopterans, , and lepidopterans, occasionally supplemented by small vertebrates, seeds, or berries. It is serially monogamous, with breeding seasons varying by region; clutches typically number 4–7 eggs in and up to 10 in , incubated for 15–18 days by the , followed by fledging at 26–32 days and after about a month. Notably, Eurasian hoopoe nests produce a strong from antibacterial secretions in the 's , which helps combat parasites and may deter predators, allowing for potentially 2–3 broods per season. Despite its adaptability, populations face declines in some areas due to habitat loss from agricultural intensification and .

Taxonomy and systematics

Classification

The hoopoe is classified within the family Upupidae, a monotypic family containing the single genus , and placed in the order , which encompasses hoopoes, woodhoopoes, and hornbills. In older taxonomic systems, such as the Sibley-Ahlquist classification, hoopoes were assigned to a separate order, Upupiformes, or grouped within alongside rollers and . The (Upupa epops), the most widespread species, was first formally described by in the 10th edition of Systema Naturae in 1758, establishing the binomial name Upupa epops. Historically, hoopoes were classified in the order based on morphological similarities, but molecular phylogenetic analyses in the 2000s, including extensive nuclear , supported their reclassification to due to shared evolutionary traits with hornbills and woodhoopoes. Phylogenetic studies confirm that Upupidae forms a with the families Bucerotidae (hornbills) and Phoeniculidae (woodhoopoes), with hoopoes as the to woodhoopoes. This relationship is reinforced by subsequent phylogenomic research using thousands of loci, highlighting convergent adaptations in bill structure and nesting behaviors. The (Upupa epops) is recognized as comprising eight , differentiated primarily by variations and geographic across and , though some authorities debate the status of certain forms based on .

Species diversity

The genus Upupa includes three extant , each adapted to distinct regions with subtle differences in and vocalizations. The (Upupa epops) is the most widespread, occurring across , , and , with a body length of 25–32 cm and a weight of 46–89 g; it is characterized by its cinnamon-colored and a territorial call consisting of a series of three mellow notes rendered as "hoo-poo-poo." The (Upupa africana) inhabits sub-Saharan south of the , measuring 25–29 cm in length and weighing 38–67 g, distinguished by its slightly smaller size, darker grayish mantle, and a softer, repetitive "oop-oop" call. The (Upupa marginata), endemic to the island of , is the largest at approximately 32 cm long and 57–91 g, featuring a thicker bill, grayer upperparts, and a distinctive long, hollow as its primary vocalization. Fossil evidence indicates additional extinct species within the genus, though the record is sparse and primarily from the Quaternary period rather than earlier epochs. The Saint Helena hoopoe (Upupa antaios), known from subfossil remains on the island of Saint Helena, was notably larger than extant species and likely flightless; it is estimated to have gone extinct in the 16th century following human colonization, due to habitat destruction and introduced predators. No confirmed Miocene fossils of Upupa have been described, with pre-Quaternary records limited to related upupid forms. Hybridization between hoopoe species is rare but documented in zones of , particularly between the (U. epops) and (U. africana) where their ranges overlap in , resulting in intermediate plumage and vocal traits in offspring. All three extant hoopoe species are assessed as Least Concern on the (as of 2024), reflecting their large ranges and adaptable habits.

Physical description

Morphology and size

The hoopoe, specifically the Eurasian hoopoe (Upupa epops), is a medium-sized measuring 25–32 cm in length, with a of 44–48 cm and a weight ranging from 46–89 g. These dimensions position it as a compact yet robust adapted for both terrestrial foraging and aerial mobility. Key anatomical features include a long, thin, downcurved measuring 5–6.3 cm, which facilitates probing into and crevices. The legs are strong and suited for ground-based activities, supporting the bird's habit of striding across open terrain. Its wings are broad and rounded, enabling agile, undulating flight patterns. Skeletal adaptations enhance the hoopoe's efficiency, with a sturdy, pneumatized featuring kinetic hinges and a reinforced system to withstand stresses from bill insertion into substrates. The feet exhibit a semi-zygodactyl arrangement, where the outer toe can rotate backward, aiding in perching on branches and walking on the ground. Sexual dimorphism is minimal, with females generally slightly smaller in body length (averaging 26.23 cm versus 27.88 cm in males) and lighter in mass (54.35 g versus 64.03 g), alongside minor variations in length showing no significant difference between sexes.

Plumage and distinctive features

The hoopoe's is predominantly sandy or on the body, head, and underparts, providing a warm, earthy tone that contrasts sharply with its boldly patterned wings and tail. The wings exhibit striking barring, with broad tips visible both at rest and in flight, while the tail features a similar pattern ending in a white tip bordered by a subterminal . This coloration is consistent across adults, with the sexes showing only subtle differences: females tend to have slightly duller crests, heads, and breasts, often appearing more or overall. A defining feature of the hoopoe is its prominent erectile , composed of 28 elongated feathers that form a fan-like structure up to 10 cm in length when fully raised. These crest feathers are at the base, tipped with black and sometimes showing white patches near the tips, creating a striking visual display. The crest is typically held flat against the head but can be elevated to enhance the bird's . Juvenile hoopoes closely resemble adults in overall pattern but exhibit duller, more plumage with reduced contrast in the barring. Their crests are notably shorter and less developed, contributing to a more subdued appearance; they undergo a post-juvenile molt within 1–2 months, transitioning to adult-like coloration. shows seasonal wear, appearing fresher and more vibrant after molting in late summer or autumn, but fading and becoming worn by the non-breeding season due to exposure. variations primarily involve differences in color intensity and patterning: for instance, populations (such as U. e. africana) and eastern Asian subspecies like U. e. saturata display darker, richer tones.

Distribution and habitat

Geographic range

The hoopoe family (Upupidae) encompasses two recognized with distinct geographic distributions across the . The Common Hoopoe (Upupa epops) has the broadest range, breeding across much of south of approximately 53°N latitude—from the and eastward through (including , , and the region of ), into (extending to , , and ), (from to and the ), and (from and southern southward through , the , central , and , extending to , excluding extreme desert regions and dense equatorial rainforests). This species is partially migratory, with northern populations (in and ) retreating southward in winter to regions south of 35°N, but it occurs year-round in milder regions like the and is sedentary or nomadic in , with limited seasonal movements tied to local wet-dry cycles. Vagrant individuals have been recorded far outside this core range, including at least four sightings in and two or three in . In contrast, the (Upupa marginata) has a highly restricted range, being endemic to the island of , where it occurs abundantly in the north, west, and south, with local abundance on the central plateau (such as between Bemaraha and ) and scarcity in the eastern humid regions. This non-migratory species favors the island's drier zones, with no records beyond . Historically, the Common Hoopoe underwent post-glacial colonization of approximately 10,000 years ago, following a population bottleneck during the , as evidenced by genetic signatures of rapid expansion and low phylogeographic structure indicating continuous across the continent. More recently, in the , the species has shown northward range shifts in , with breeding populations expanding into areas like the , , and , partly attributed to climate warming that has facilitated milder winters and suitable conditions in previously marginal northern habitats (as of 2025).

Habitat preferences

The hoopoe (Upupa epops) primarily inhabits open and semi-open landscapes that provide suitable foraging and nesting opportunities, including savannas, grasslands, farmlands, and parklands with short vegetation cover. These environments feature sparsely vegetated or bare ground, such as pastures, orchards, and olive groves, which facilitate ground foraging, while the species avoids dense forests and extensive woodlands due to limited access to prey and nesting sites. In agricultural mosaics, hoopoes favor areas with a mix of short grass and bare patches, typically selecting sites with 30-70% bare ground coverage at the foraging scale for optimal insect access. The occurs across a broad altitudinal range from to 3,650 m, though it is more commonly found below 2,000 m; in populations, it reaches up to 2,500 m in dry savannas, whereas populations are generally restricted to lower elevations under 1,500 m. Microhabitat requirements emphasize proximity to insect-rich, penetrable soils like sandy or loamy grounds near scattered trees, cliffs, or walls for nesting cavities, often within 200-300 m of areas. Some populations have adapted to settings, utilizing lawns and parks in areas like the for while nesting in nearby structures. Studies from the indicate that influences hoopoe distribution, with a preference for landscape edges over or interiors, as these transitional zones offer complementary resources for in wooded patches and in adjacent open areas. This edge affinity is evident in mosaic farmlands, where fragmented habitats with hedgerows and bare ground patches support higher efficiency compared to uniform dense vegetation.

Behavior and ecology

Diet and foraging

The hoopoe (Upupa epops) is primarily insectivorous, with insects accounting for the majority of its diet across various studies. Favored prey includes large soil-dwelling invertebrates such as orthopterans (e.g., crickets and grasshoppers), coleopterans (beetles and their larvae), and lepidopterans (moth and butterfly larvae and pupae), along with ants, earwigs, cicadas, and ant lions. In one Mediterranean study, insect larvae comprised 84% of the identified diet, dominated by cicada (Cicada orni) nymphs at 61% of that portion. Occasionally, the diet incorporates small vertebrates like lizards, frogs, snakes, and even bird eggs, as well as plant matter such as seeds and berries. Foraging occurs mainly on the ground in open, sparsely vegetated areas like bare , short grasslands, or field margins, where the bird's long, curved —adapted for soil penetration—allows it to probe for hidden prey. The hoopoe walks steadily, pausing to visually scan for movement before inserting its bill vertically into soft , often employing a "gaping" by opening and closing the bill tip to loosen soil and expose . It is diurnal, with activity peaking at dawn and dusk when prey is more active, though it may forage solitarily or in pairs throughout the day. Diet composition shows some seasonal variation, remaining insect-focused year-round but shifting toward greater reliance on available large invertebrates during breeding periods, such as mole crickets in alpine regions or processionary moth larvae in certain forests. Adults can consume substantial quantities, with provisioning rates during active periods exceeding 14 prey items per hour, supporting high energetic demands.

Social structure and vocalizations

Hoopoes (Upupa epops) are generally solitary outside the season, often observed or moving alone or in pairs, reflecting a that emphasizes individual or monogamous pair bonds lasting typically one season. During non-breeding periods, including , they may form small groups or loose flocks, occasionally exceeding 100 individuals, though these groups lack strong cohesion compared to more . Territorial behavior is prominent in males, who defend nesting and foraging areas through a combination of vocal and physical displays to deter rivals and assert dominance. The simplest threat posture involves the male standing erect with its crest raised but not fully fanned, wings slightly opened, and tail spread to appear larger and more intimidating. Escalations may include bowing motions, aggressive chases, or even physical clashes with bills, sometimes resulting in injury to intruders. These displays serve both aggressive and functions, with males using them to advertise territory ownership and attract potential mates. The hoopoe's vocal repertoire supports its and dynamics, featuring an iconic trisyllabic rendered as "hoop-hoop-hoop" or "oop-oop-oop," which primarily functions to defend intrasexually and attract females intersexually, often audible up to 1.5 away. Softer chatters and contact calls maintain pair or group proximity during non-breeding interactions, while alarm calls—such as raspy croaks, mechanical rattles, or harsh hisses—signal threats during chases or disturbances, indicating varying levels of urgency. These vocalizations, combined with visual displays, enable effective communication in their open habitats.

Reproduction

Breeding biology

The common hoopoe (Upupa epops) exhibits a monogamous , with pair bonds typically lasting for a single season, although pairs may occasionally attempt a second brood with the same or a different partner. In northern populations, is seasonal and occurs primarily in (March to June in ), synchronized with abundance following from wintering grounds in the . In contrast, tropical populations are largely resident and breed year-round or during favorable wet seasons, allowing multiple clutches without long-distance . Courtship begins with males attracting s through vocalizations and visual displays, including raising their flamboyant while emitting soft calls to signal availability and quality. Males also perform pursuit flights, chasing potential mates in undulating patterns characteristic of the species, and engage in feeding where they offer prey but repeatedly withdraw it in a manner to elicit female interest and strengthen pair bonds. Females lay clutches of 4–9 eggs, with an average of about 7 in temperate regions and slightly smaller sizes in ; laying is asynchronous, typically one egg every 1–2 days. , which begins after the first or second egg is laid, lasts 15–18 days and is performed exclusively by the , during which the provisions her with at the nest entrance. Reproductive success varies but typically achieves 50–70% fledging rates per clutch, with overall nest success influenced heavily by prey availability, particularly outbreaks of soil-dwelling insects like () that boost chick survival in resource-rich years. In one study, 73% of hatched chicks fledged, though rates decline with later-season broods or poor insect densities.

Nesting and parental care

Hoopoes are secondary cavity nesters that do not construct nests but instead utilize preexisting natural cavities, such as tree hollows, rock crevices, cliffs, sandy banks, or holes excavated by other species like woodpeckers. They also frequently occupy man-made structures, including walls, , pipes, and nest boxes, often reusing the same sites across seasons with accumulated remains of soft materials from prior occupations. This reuse contributes to the nest environment's microbial and parasitic profile but provides no structural addition by the birds themselves. The clutch typically consists of 4–9 pale blue-grey, unmarked, oval eggs measuring approximately 2.5–3 cm in length. Females apply a cosmetic secretion from their to the eggshells shortly after laying, which alters their appearance to a more saturated greyish-blue during , potentially serving or signaling functions. Only the incubates the eggs for 15–18 days, leaving the nest infrequently; during this period, the provisions her with food at the cavity entrance to sustain her. Hatchlings are altricial, emerging helpless, blind, and covered in downy . Both parents feed the chicks a diet primarily of , including large prey items like beetles and caterpillars that are beaten and broken down before delivery to the nest. The nestling period lasts 26–32 days, after which the young , though they remain dependent on parental provisioning for up to several weeks post-fledging as they develop flight and skills. Hoopoes exhibit defenses against by the , a potential parasite, primarily through aggressive nest protection and against non-kin at the nestling and fledgling stages rather than routine ejection.

Conservation

Population status and threats

The common hoopoe (Upupa epops) is classified as Least Concern on the due to its extensive global range spanning over 104 million km² and a large overall population. Global population estimates range from 5 to 10 million individuals, with the Eurasian subspecies being the most abundant across its wide in , , and . In specifically, the breeding population is estimated at 1.37 to 3.21 million pairs, equivalent to 2.6 to 6.4 million mature individuals, though data quality is considered poor. Despite the stable global status, regional population trends indicate declines, particularly in where long-term decreases (1980–2018) have been observed in several countries, including 30–60% reductions in , 20–30% in , and 10–30% in . These declines are attributed to and loss of grounds, exacerbated by agricultural intensification that reduces open, insect-rich areas essential for the . Key threats include the widespread use of pesticides, which diminish invertebrate prey populations such as mole crickets and beetles that form the hoopoe's primary diet. Climate change further compounds these pressures by altering weather patterns, with cool, rainy conditions during breeding seasons reducing chick provisioning and reproductive success, potentially leading to rapid population drops if adverse years persist. Emerging concerns involve increased road mortality along migratory routes in fragmented landscapes, though quantitative data specific to hoopoes remains limited.

Protection and management

The hoopoe (Upupa epops) is protected under the European Union's Birds Directive, which safeguards all wild species native to the EU, including their habitats, eggs, and nests, through measures such as the designation of Special Protection Areas under the network. In , where occur, the benefits from protections within national parks, such as in . The species is not listed under appendices, reflecting its global Least Concern status, though populations are monitored through frameworks like the African-Eurasian Migratory Landbirds Agreement. Conservation programs in have focused on mitigating agricultural impacts, including the EU-wide ban on insecticides implemented in , which aims to reduce declines in insectivorous birds like the hoopoe by protecting prey availability, though recovery has been slow in contaminated landscapes. Habitat restoration efforts incorporate agri-environment schemes that promote bare ground patches and low-intensity farming practices, creating foraging habitats essential for the hoopoe in intensively managed farmlands. Monitoring initiatives include BirdLife International's European Red List assessments and population surveys, which track breeding trends and distribution across the EU using data from member states to inform conservation priorities. platforms like eBird contribute to tracking by aggregating observer reports, enabling real-time mapping of seasonal movements and range shifts for the hoopoe. Nest box programs have demonstrated success in stabilizing local populations; for instance, intensive supplementation in agricultural landscapes since 1998 increased breeding density and by providing artificial cavities in areas lacking natural sites, leading to . Similar initiatives in , such as those in the Hoya de Guadix region, have supported breeding through provision, contributing to stabilized numbers in modified habitats since the early .

Relationship with humans

Cultural symbolism and emblems

The hoopoe serves as a prominent national emblem in , where it was designated the national bird in May 2008 following a public vote organized by the Society for the Protection of Nature in to mark the country's 60th anniversary; over 155,000 participants selected it from a list of ten species for its striking appearance and long-standing presence in the region. In ancient Egyptian iconography, the hoopoe featured prominently in art and reliefs, where its elaborate crest was sometimes likened to the rays of , symbolizing solar attributes and ; it also represented the legitimate heir and successor in depictions from around 2575–2400 BCE. Persian cultural traditions elevate the hoopoe as a symbol of virtue, spiritual perfection, and guidance, often portrayed as a wise messenger leading other birds toward enlightenment in classical literature. In modern emblems, the African hoopoe appears in the design of the University of Johannesburg's logo, incorporating two stylized representations of the bird to reflect regional biodiversity, and inspires the university's sports mascot, Hoepie, which embodies the institution's vibrant identity. The hoopoe also functions as a tourism icon in birdwatching hotspots, such as Andalusia in Spain—where its vivid plumage and crest make it a highlight of local wildlife tours—and Kruger National Park in South Africa, drawing visitors to observe its distinctive foraging behaviors in savanna habitats. In ancient Jewish texts, the is listed among the unclean birds prohibited for consumption in , reflecting ritual purity laws that deemed it impure due to its habits and appearance. This classification underscores the bird's association with impurity in biblical dietary restrictions, where it is grouped with other species considered ritually unfit. Greek mythology links the hoopoe to the tragic tale of Tereus, Procne, and Philomela, where Tereus is transformed into a hoopoe as punishment for his crimes, symbolizing betrayal and vengeance. In this myth, the gods intervene to turn the principals into birds, with the hoopoe's distinctive crest evoking Tereus's crown of guilt, a motif echoed in classical literature like Ovid's Metamorphoses. In African folklore, particularly among the Ndebele people of , the hoopoe is revered as a harbinger of rain, its calls believed to summon the rainy season and ensure bountiful harvests. This role positions it as a protector spirit in agrarian societies, where harming the bird is taboo due to its perceived influence over weather and fertility. In Islamic tradition, the hoopoe (hudhud) appears in the Quran's Surah an-Naml (27:20-28) as a messenger to Prophet , reporting on of Sheba's sun worship and delivering a divine letter that leads to her conversion. This narrative highlights the bird's wisdom and role as a divine intermediary, emphasizing themes of and obedience. Hoopoes feature in Middle Eastern folk medicine, particularly in Coptic Egyptian remedies where warm blood from the bird, mixed with , treats eye conditions like excessive tearing or inflammation. Among communities in northern , the bird's eyes are used in amulets to ward off the , a practice rooted in pre-Islamic traditions and documented in medieval texts like those of for treating ailments including . These uses, often involving the bird's body parts in potions or rituals, lack scientific validation and reflect cultural beliefs in its protective and curative powers. In , the hoopoe serves as the enlightened guide in Farid ud-Din Attar's 12th-century Sufi epic , leading a flock on a spiritual quest to find the , symbolizing the soul's journey toward divine union. This allegorical role has influenced mystical traditions, portraying the bird as a wise shepherd of . In modern , hoopoes appear in animations like the short film Gift of the Hoopoe (2009), where the bird embodies imagination and personal growth in a fantasy for children. They also feature in nature documentaries, and in video games like Feather Family on , where the hoopoe is a playable character inspired by its real-world traits. In fantasy settings, such as Magic: The Gathering's River Hoopoe card from the Hour of Devastation set, it represents adaptive survival in mystical worlds.

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