Tawny eagle

The tawny eagle (Aquila rapax) is a large bird of prey. Like all eagles, it belongs to the family Accipitridae. Its heavily feathered legs mark it as a member of the subfamily Aquilinae, also known as booted eagles.<ref name= Helbig>Helbig, A. J., Kocum, A., Seibold, I., & Braun, M. J. (2005). A multi-gene phylogeny of aquiline eagles (Aves: Accipitriformes) reveals extensive paraphyly at the genus level. Molecular phylogenetics and evolution, 35(1), 147-164.</ref> Tawny eagles have an extensive but discontinuous breeding range that constitutes much of the African continent as well as the Indian subcontinent, with rare residency in the southern Middle East.<ref name= Ferguson-Lees>Template:Cite book</ref> Throughout its range, it favours open dry habitats such as semideserts, deserts steppes, or savanna plains. Despite its preference for arid areas, the species seldom occurs in areas where trees are entirely absent.<ref name= Hustler>Hustler, K., & Howells, W. W. (1989). Habitat preference, breeding success and the effect of primary productivity on Tawny Eagles Aquila rapax in the tropics. Ibis, 131(1), 33-40.</ref> It is a resident breeder which lays one to three eggs in a stick nest most commonly in the crown of a tree.<ref name= Naoroji>Naoroji, R., & Schmitt, N. J. (2007). Birds of prey of the Indian subcontinent. Om Books International.</ref> The tawny eagle is perhaps the most highly opportunistic of all Aquilinae, and often scavenges on carrion or engages in kleptoparasitism towards other carnivorous animals but is also a bold and active predator, often of relatively large and diverse prey.<ref name= Brown>Brown, Leslie and Amadon, Dean (1986) Eagles, Hawks and Falcons of the World. The Wellfleet Press. Template:ISBN.</ref> It is estimated that tawny eagles can reach the age of 16 years old.<ref name= Wichmann/> Nonetheless, precipitous declines have been detected throughout the tawny eagle's range. Numerous factors, particularly loss of nesting habitat due to logging and global warming, as well as persecution (largely via poisoning) and other anthropogenic mortality (largely through contact with various manmade objects) are driving the once numerous tawny eagle perhaps to the brink of extinction.<ref name="iucn status 12 November 2021" /><ref name= Wichmann>Template:Cite journal</ref><ref>Ogada, D. L. (2014). The power of poison: pesticide poisoning of Africa's wildlife. Annals of the New York Academy of Sciences, 1322(1), 1-20.</ref>

Taxonomy

Dutch naturalist Coenraad Jacob Temminck described the tawny eagle in 1828 from an Indian type specimen.<ref>Temminck, C. J. (1828). Blik op de dierlijke bewoners van de Sunda-eilanden en van de overige Nederlandsche bezittingen in Indië. Bijdr. nat. Wetensch, 3, 64-78.</ref> "Tawny eagle" has been designated the official name by the International Ornithologists' Union (IOC).<ref name="ioclist">Template:Cite web</ref> Aquila is Latin for "eagle" while rapax is also Latin for "to grasp", and relates to the word rapacious, i.e. highly predatory.<ref name= Naoroji/><ref name= Steyn>Steyn, P. (1983). Birds of prey of southern Africa: Their identification and life histories. Croom Helm, Beckenham (UK). 1983.</ref><ref name= Brown2>Brown, L. (1977). Eagles of the World. Universe Books.</ref> The tawny eagle is a member of the booted eagle subfamily (Aquilinae) within the Accipitridae family.<ref name= Ferguson-Lees/> The booted eagle clan are monophyletic and study of karyotypes has indicated that they likely have few to no close external relations within the overall extant accipitrid family.<ref name= Helbig/><ref>Li, Q., Lin, J., Li, S., Wang, Y., Li, W., & Zeng, Y. (2000). Studies on the evolution of mitochondrial DNA in 11 species of Accipitridae. Dong wu xue bao.[Acta zoologica Sinica], 46(2), 209-220.</ref><ref>De Boer, L. E. M., & Sinoo, R. P. (1984). A karyological study of Accipitridae (Aves: Falconiformes), with karyotypic descriptions of 16 species new to cytology. Genetica, 65(1), 89-107.</ref> The booted eagle subfamily all have feathers covering their legs and are distributed in every continent that contains accipitrids.<ref name= Brown/> The genus Aquila has been traditionally defined as largish, dark-hued and long-winged eagles of open country.<ref name= Valli>Väli, Ü. (2002). Mitochondrial pseudo‐control region in old world eagles (genus Aquila). Molecular Ecology, 11(10), 2189–2194.</ref> However, study of genetics have resulted in several reclassifications. These include the moving of smaller, paler and more forest-dwelling eagles in Aquila, the separation of the unique spotted eagles to the genus Clanga as well as the division of some small eagles to the genus Hieraeetus outside of Aquila. Furthermore, genetic research has further revealed a schism in superficially similar eagles between the tawny eagle and its close relatives and other superficially similar Aquila such as the golden eagle (Aquila chrysaetos) species complex. To date, the tawny eagle species group and golden eagle species group are still contained in the same genus despite the apparent lack of close relation.<ref name= Helbig/><ref name= Valli/><ref name= Lerner>Lerner, H. R., & Mindell, D. P. (2005). Phylogeny of eagles, Old World vultures, and other Accipitridae based on nuclear and mitochondrial DNA. Molecular phylogenetics and evolution, 37(2), 327-346.</ref><ref name= Watson>Template:Cite book</ref>

File:Raubadler Aquila rapax.jpg

A tawny eagle sitting on the ground displaying typical features: a mid-sized eagle with a long neck, well-feathered legs, well-proportioned frame and large bill lacking a deep gape.

The tawny eagle was previously treated as conspecific with the migratory steppe eagle (Aquila nipalensis).<ref name= Brown/><ref>Stresemann, E., & Amadon, D. (1979). Order Falconiformes. Check-list of birds of the world, 1, 271-425.</ref> These eagles were considered part of the same species as recently as 1991.<ref>Howard, R., & Moore, A. (1991). A complete checklist of the birds of the world (No. Ed. 2). Academic Press Ltd.</ref> The steppe and tawny eagles were split based on pronounced differences in morphology and anatomy. The steppe eagle is a larger bird, with a much more pronounced gape, and differs by appearance in its blockier frame, bigger wings and evinces different coloring at all stages of development, despite some morphs of the two eagles superficially resembling one another.<ref name= Clark>Template:Cite journal</ref><ref name= Olson>Template:Cite journal</ref><ref>Template:Cite journal</ref><ref>Amadon, D., & Short, L. L. (1976). Treatment of subspecies approaching species status. Systematic Zoology, 25(2), 161-167.</ref><ref>Brooke, R. K., Grobler, J. H, Irwin, M. P. S., & Steyn, P. (1972). A study of the migratory eagles Aquila nipalensis and A. pomarina (Aves:Accipitridae) in Southern Africa, with comparative notes on other large raptors. Occ. Papers of the Nat. Mus. of Rhodesia B5(2): 61–114.</ref> Furthermore, the respective species differ in ecology (dietary biology, nesting habits) and are strongly allopatric in their breeding ground distribution.<ref name= Naoroji/><ref name= Clark/> Two molecular studies, each based on a very small number of genes, indicate that the species are distinct, but disagree over how closely related they are.<ref>Template:Cite web</ref><ref>White, C. M., Olson, P. D., & Kiff, L. F. (1994). Family Falconidae (hawks and eagles). Handbook of the birds of the world, 2: 52-214.</ref> Genetically, the tawny eagle may cluster more closely with the imperial eagle species complex despite the steppe eagle being more sympatric with those northerly Eurasian eagles.<ref name= Helbig/>

Subspecies

File:Tawny Eagle (8409160315).jpg

The distinctly darker subspecies native to India and elsewhere in Asia, A. r. vindhiana.

There are three described races of tawny eagles. The subspecific classification of the species has at times been considered complicated by variations and existence of different morphs; in turn they were once considered tentative.<ref name= Ferguson-Lees/> However, each subspecies is largely allopatric in geography, the primary ambiguities lying in the northern part of east Africa where both African races may intergrade.<ref name= Kemp>Kemp, A.C. and G. M. Kirwan (2020). Tawny Eagle (Aquila rapax), version 1.0. In Birds of the World (J. del Hoyo, A. Elliott, J. Sargatal, D. A. Christie, and E. de Juana, Editors). Cornell Lab of Ornithology, Ithaca, NY, USA.</ref>

• A. r. rapax; distributed in Africa from the southern Democratic Republic of the Congo and central Kenya to all points southwards.<ref name= Ferguson-Lees/> However, this race may range up to as far north as Ethiopia as well (thus likely intergrading considerably with the following subspecies which is widespread in Ethiopia).<ref name= Ash>Ash, C. P., & Atkins, J. D. (2009). Birds of Ethiopia and Eritrea: an atlas of distribution. A&C Black.</ref> Adult often of this nominate subspecies are often relatively more strongly rufous in colour than other races and are sometimes dark streaked below. Meanwhile, the juvenile tends to be light rufous. wing chord lengths in this race have been measured at Template:Convert in males and Template:Convert in females.<ref name= Ferguson-Lees/> The mean wing chord length in two samples of A. r. rapax measured Template:Convert in males and Template:Convert in females.<ref name= Brown/><ref name= Musindo>Musindo, P. T. (2012). Morphological variation in bills and claws in relation to Prey type in Southern African Birds of Prey (Orders Falconiformes and Strigiformes).</ref> The tail length of both sexes in A. r. rapax measures Template:Convert with a tarsus length of Template:Convert. Body mass can range roughly from Template:Convert in overall samples of at least 36 eagles.<ref name= Ferguson-Lees/>
• A. r. belisarius; this race resides in west Africa to Ethiopia and southwest Arabia as well as far south as the northern Democratic Republic of the Congo and northern Kenya.<ref name= Ferguson-Lees/><ref name= Clouet>Clouet, M., Barrau, C., & Goar, J. L. (2000). The diurnal Afro-alpine raptor community of the Ethiopian Balé Highlands. Ostrich, 71(3-4), 380-384.</ref> This race is described to appear "neater" than the nominate subspecies, possibly due to this race often occurring in even more desert-like conditions than the other races and thus having more compact feathering.<ref name= Ferguson-Lees/><ref>Cook, W. E. (2012). Avian desert predators. Springer Science & Business Media.</ref> It is when compared to the nominate, often duller and browner above, showing less of a rufous tinge. Meanwhile, individual pale morph of A. r. belisarius often a shade or two darker than pale nominate but not consistently so. Although said to be slightly larger, measurement data shows this subspecies to of broadly very similar size to the nominate race. In males, the wing chord is Template:Convert and, in females, the wing chord is Template:Convert.<ref name= Ferguson-Lees/> The mean wing chord lengths were reportedly Template:Convert in males and Template:Convert, which indicates a slightly less pronounced sexual dimorphism than in the nominate race.<ref name= Brown/> As for body mass, 1 male was found to weigh Template:Convert while three females weighed from Template:Convert.<ref name= Ferguson-Lees/><ref name= Brown/>
• A. r. vindhiana; excluding the Arabian Peninsula, this race likely comprises all the tawny eagles found in Asia, such as in southeastern Iran and the Indian subcontinent.<ref name= Naoroji/> However, ambiguities exists on where the range of A. r. belisarius ends and of vindhiana begins, especially in Middle Eastern areas.<ref>Al-Sheikhly, O. F., & Al-Azawi, A. J. (2019). The Diurnal Birds of Prey (Raptors) in the Mesopotamian Marshes of Southern Iraq with notes on their Conservation Status. Bulletin of the Iraq Natural History Museum (P-ISSN: 1017-8678, E-ISSN: 2311-9799), 15(4), 381-402.</ref><ref>Crochet, P. A., Raty, L., De Smet, G., Anderson, B., Barthel, P. H., Collinson, J. M., Dubois P.J., Helbig, A.J., Jiguet, F., Jirle, E., Knox, A.G., Le Maréchal, P., Parkin, D.T., Pons, J.-M., Roselaar C.S.,Svensson, L., van Loon, A.J., & Yésou P. (2010). AERC TAC's taxonomic recommendations. July 2010. Disponible sur: http://www. aerc. eu/tac. html.</ref> Sometimes A. r. vindhiana is suggested as full species.<ref>Sibley, C. G., & Monroe, B. L. (1990). Distribution and Taxonomy of Birds of the World. Yale University Press.</ref> This subspecies averages darker than either other race and usually is lacking in warmer rufous tone. In general, it is somewhat more similar in hue to the steppe eagle.<ref name= Ferguson-Lees/> Adult irises are sometimes brown in A. r. vindhiana (again reminiscent of the steppe eagle). The pale morph of this race is greyer and less rufous than African tawny eagles, although generally juveniles and immature are sometimes more rufous. It may be marginally the smallest subspecies, although in general the tawny eagle evinces remarkably little size variation across its wide range. Wing chord measurements are Template:Convert in males and Template:Convert.<ref name= Ferguson-Lees/><ref name= Ali>Ali, S., & Ripley, S. D. (1980). Handbook of the birds of India and Pakistan, together with those of Bangladesh, Nepal, Bhutan, and Sri Lanka. Oxford University Press.</ref><ref name= Baker>Baker, E. C. (1928). Fauna of British India including Ceylon and Burma. Taylor & Francis.</ref> In males, the wing chord reportedly averages about Template:Convert and in females, it averages Template:Convert.<ref name= Brown/> In males the tail length is Template:Convert and in female it is Template:Convert. The tarsus length of male A. r. vindhiana is Template:Convert and in females is Template:Convert.<ref name= Ali/><ref name= Baker/> Unsexed adults in India weighed from Template:Convert.<ref name= Naoroji/>

Description

File:2011-tawny-eagle-closeup.jpg

Close-up showing gape extending only to below the middle of the eye

The tawny eagle is considered to appear "inelegant, scruffy-looking" but has a fairly characteristic aquiline silhouette.<ref name= Ferguson-Lees/> The species has a fairly long neck and long deep bill with a gape line level with the eye, moderately long wings with fairly pronounced "fingers" and a slightly rounded to almost square-ended and shortish tail, which can be more reminiscent of the tail of a vulture than that of other eagles. The feathering on the legs is extensive and can appear almost baggy-looking.<ref name= Ferguson-Lees/><ref name= Steyn/><ref name= AliS>Template:Cite book</ref> The bill and head are strong and bold, the body well-proportioned and feet are powerful while the countenance is quite fierce-looking.<ref name= Steyn/> While perching, the tawny eagle tends to sit rather upright, often on stumps, posts, low trees or treetops for long periods of the day or may descend to the ground to walk somewhat unsteadily with a more horizontal posture. The wingtips when perched are roughly even with the tip of the tail.<ref name= Ferguson-Lees/> Adults have variably colored eyes, ranging from yellow to pale brown to yellow brown, while those of juveniles are dark brown. Both the cere and feet are yellow at all ages.<ref name= Ferguson-Lees/> The tawny eagle is polymorphic with considerable individual variation in plumage, resulting in occasional disparities in plumages that can engender confusion in some.<ref name= Ferguson-Lees/><ref name= Naoroji/> In adulthood, they can vary in coloration from all dark grey-brown to an occasionally streaky (or more plain) foxy-rufous to buffish-yellow.<ref name= Ferguson-Lees/><ref name= Shirihai>Shirihai, H. (1994). Separation of Tawny Eagle from Steppe Eagle in Israel. British Birds, 87, 396-397.</ref> Most adults are usually a general grey-brown or rufous-tawny color, with occasional pale spotting visible at close quarters on the nape and belly, coverts uniformly toned as the body.<ref name= Naoroji/> The nape is consistently dark and uniform despite the feathers often being tipped paler with other feathers in adults, lacking the contrasting paler feathers often seen in other Aquila.<ref name= Naoroji/> Females, in addition to being slightly larger, may tend to be slightly darker and more streaked than the males.<ref name= Ferguson-Lees/> The most blackish-brown individuals tend to occur in India.<ref name= Ferguson-Lees/><ref name= Naoroji/> Adults often show relatively little varying colors apart from their somewhat blacker wing and tail feathers, though when freshly molted great wing coverts and secondaries may show small pale tips which may form pale lines along closed wing has tawny upper parts and blackish flight feathers and tail.<ref name= Ferguson-Lees/> The head is often similarly tawny in colour as the body but may also sometimes shows darker eyebrows, other thin brown streaks or a darker chin.<ref name= Ferguson-Lees/> Meanwhile, the tail is plain or obscurely dark barred (with around 7 subtle bands). The dark morph adult is essentially all dark, dull brown. Some dark morph tawny eagles with wear may show irregular streaking or molting browns and more blackish feathers.<ref name= Ferguson-Lees/> Intermediate morph are dark to rufous brown above with the mantle and wing coverts variably streaked or molted lighter rufous as is the head with the crown or crown-sides being paler. The intermediate morph's underside is largely rufous (especially farther south in Africa) with breast and flanks very heavily and broadly streaked dark brown, though at times appears all dark brown contrasting with plain trousers and crissum.<ref name= Ferguson-Lees/> Pale morph adult tawny eagles always show a clear contrast between the pale body and wing coverts which bear darker flight feathers and tail. In pale morphs, the underparts are rufous buff to lighty tawny-brown, phasing into somewhat darker lesser and median wing coverts to darker brown to even blackish greater coverts and flight feathers. The head may too be tawny in pale morph tawny eagles but sometimes with thin brown streaks or darker chin. Below pale morph adults are all light rufous to tawny buff or brown, sometimes paler below the belly area. In worn individuals the bodily feathers of pale morph tawny eagles can appear almost whitish.<ref name= Ferguson-Lees/> Dark morph juvenile tawny eagles are generally light rufous to rufous brown with creamier lower back to upper tail coverts. Juveniles show thinly pale-tipped dark brown greater coverts and remiges while the tail is barred grey and brown usually with a narrow creamy tip. Dark morph juveniles may fade to pale buff or creamy often before molting into browner plumage. Subsequent stages are not as well-known but it appears dark morph subadults gradually manifest a darker brown or rufous brown color on the mantle, as well as on the head and upper breast while maintaining a buffish rear body (i.e. lower back and rump patch). Generally other morphs are similar but not as well-known and are perhaps individually inconsistent. Many are rufous or sandy after a molt but have mottling later on, the extent of pale feathers indicative perhaps of their ultimate adult morph.<ref name= Ferguson-Lees/>

File:Tawny eagle (Aquila rapax rapax) streaked morph.jpg

A streaked morph of the nominate subspecies of tawny eagle.

In flight, the tawny eagle appears as a large raptor with a noticeably protruding head on a long neck, with a deep chest, long and broad wings with a somewhat narrower seven-fingered hand.<ref name= Ferguson-Lees/> The trailing edge of the wing is slightly curved outwards, indenting at the junction of primaries and secondaries, whilst the rounded, medium-length tail is usually held spread.<ref name= Steyn/> The deep beats of the kinked wings can make their flight appear rather heavy and slow but they are quicker and more expansive in wing movements and often less forceful-looking than larger Aquila like steppe eagles and can be very agile when chasing other raptors to rob them.<ref name= Ferguson-Lees/><ref name= Naoroji/> Tawny eagles soar with flat wings or very slightly raised and hands only slightly lower, and may fly similarly in a glide but may too arch when in a fast glide.<ref name= Ferguson-Lees/> Adult dark morphs are more or less uniform dark brown above and below, showing indistinctly and slightly paler and greyish primaries on both sides. Above, the main contrast on dark morphs above is paler creamy rump patch while, on the underside, the greyish color is contrasted with blackish tips and a diffused trailing edge along both the wings and tail. Intermediate morph tawny eagles are variably rufous streaked on brown to rufous brown on the back and wing coverts with a similar contrasting pale rump above as dark morphs. Below the intermediate's heavy dark streaks are only subtly different and their coloring can appear almost uniform. The wing quills of intermediate morphs are often greyer with a stronger contrast of the paler inner primaries and blackish wing ends. Pale morph are all pale tawny or buffish on both sides of the wing, which contrasting strongly with demarcated dark brown about the greater coverts, flight feathers and tail and usually the scapulars. The primaries are quite pale on pale morphs with sometimes the hint of a pale carpal comma. Some pale adults have pale bases to all the underprimaries and the quills are sometimes unbarred, but more usually the feathers have dense but narrow dark bars.<ref name= Ferguson-Lees/><ref name= Naoroji/> Dark morph juveniles are light rufous to pale tawny body above which contrasts strongly with dark brown greater coverts, rear scapulars, flight feathers and tail, in turn all highlighting the creamy lower back to tail coverts. Below dark morph juveniles can look similar to pale morph adults apart from trailing whitish edges and often irregular pale diagonals along tips of greater wing coverts, though usually these fade early on. Little is known plumage development but the young eagles moult into brown, becoming patchy with intermediate often showing 1-3 darker bars on wing linings.<ref name= Ferguson-Lees/> The underparts of subadults (i.e. around 2 to 3 years or old) are typically two-toned, with darker brown about the breasts, belly and underwings coverts while the remainder of the underbody is creamy light in colour. This two-toned pattern is evinced in subadult tawny eagles both from India and Africa.<ref name= Naoroji/><ref>Snelling, J. C. (1969). Some information obtained from marking large raptors in the Kruger National Park, Republic of South Africa. Ostrich, 40(S1), 415-427.</ref> Adult plumage is obtained between the 4th and 5th years of life.<ref name= Ferguson-Lees/>

Size

File:Tawny Eagle (Aquila rapax) (33280280216).jpg

A richly rufous-colored tawny eagle.

This is a large bird of prey, though is medium-sized for an eagle and it is one of the smaller species in the genus Aquila. Among currently accepted species in the genus, it is of quite similar size to Bonelli's eagles (Aquila fasciata) (though is notably longer winged), slightly larger than African hawk-eagles (Aquila spilogaster) and much larger than Cassin's hawk-eagles (Aquila africanus). Otherwise, females of the larger species of Aquila are frequently around twice as heavy as an average tawny eagle.<ref name= CRC>CRC Handbook of Avian Body Masses, 2nd Edition by John B. Dunning Jr. (Editor). CRC Press (2008), Template:ISBN.</ref> As is typical in birds of prey, the female tawny eagle is larger than the male, though relatively modestly so and a difference between the sexes is typically up to 15%.<ref name= Ferguson-Lees/> In total length, tawny eagles can measure from Template:Convert.<ref name= Naoroji/><ref>Borrow, N., & Demey, R. (2013). Field guide to the birds of Ghana. Bloomsbury Publishing.</ref> A typical length for a tawny eagle is considered about Template:Convert.<ref>Van Perlo, B. (1999). Birds of southern Africa. Harpercollins Pub Limited.</ref> Wingspans can measure from Template:Convert. Weight can range in fully grown birds from Template:Convert.<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name="ARKive2011">Template:Citation</ref> Average weights were reported in one study as Template:Convert in 5 males and Template:Convert in 5 females.<ref name= Musindo/> In another study, 10 unsexed adult tawny eagles were found to have weighed Template:Convert on average while, for the same data pool, a sample of 15 had an average wingspan of Template:Convert.<ref name= Mendelsohn>Mendelsohn, J. M., Kemp, A. C., Biggs, H. C., Biggs, R., & Brown, C. J. (1989). Wing areas, wing loadings and wing spans of 66 species of African raptors. Ostrich, 60(1), 35-42.</ref> Another small sample of African males, sample size four, averaged Template:Convert while three females averaged Template:Convert.<ref name= Brown/> The mean mass of the species in one estimate was Template:Convert.<ref name= Anderson>Anderson, D. J., & Horwitz, R. J. (1979). Competitive interactions among vultures and their avian competitors. Ibis, 121(4), 505-509.</ref> In all standard measurements combined, the wing chord can vary from Template:Convert, the tail from Template:Convert and the tarsus from Template:Convert.<ref name= Ferguson-Lees/><ref name= Roberts>Hockey, P.A.R., Dean, W.R.J. & Ryan, P.G. (2005). Roberts – Birds of southern Africa, VIIth ed. The Trustees of the John Voelcker Bird Book Fund, Cape Town.</ref> The culmen length of Kenyan tawny eagles was measured at Template:Convert, averaging Template:Convert, while the gape width is Template:Convert on average, ranging from Template:Convert.<ref name= Clark/><ref name= Smeenk>Smeenk, C. (1974). Comparative-ecological studies of some East African birds of prey. Ardea 62 (1-2) : 1-97.</ref> The hallux-claw, the enlarged rear talon often used as a killing instrument on accipitrids, can measure from Template:Convert, averaging Template:Convert in one sample and Template:Convert in another. The talon size is not especially large for a booted eagle and is proportionately similar in size to those of steppe eagles and eastern imperial eagles (Aquila heliaca).<ref name= Brown/><ref name= Clark/><ref name= Smeenk/>

Confusion species

File:Tawny Eagle Aquila rapax.jpg

The slightly darker northern and central African subspecies, A. r. belisarius, in flight displaying the species typical flight profile.

The tawny eagle lives in multiple areas where other broadly similar brownish hued and largish raptors often occur. Thus identification is seldom straightforward.<ref name= Kemp2>Kemp, A., & Kemp, M. (2006). Sasol Birds of Prey; New Edition. Struik.</ref><ref name= Forsman>Forsman, D. (1999). The raptors of Europe and the Middle East: a handbook of field identification. London: T & AD Poyser.</ref>

One source that can especially engender potential confusion in its wintering range is the formerly conspecific steppe eagle. The steppe is larger with a shorter neck, relatively longer and narrower wings, a more massive beak, particularly via the exceptional depth of the gape (although in flight can appear smaller headed due its less protruding neck) and has a longer and rounder tail. Furthermore, steppe eagles tends to have much bolder and widely spaced barring on the wings than tawny eagles and more distinct dark trailing wing edges and paler throats.<ref name="Ferguson-Lees" /><ref name="Clark" /><ref name="Forsman" /><ref name="Shirihai" /><ref>Jankowitz, M. (1976). Tawny or Steppe Eagle. Bokmakierie, 28(3), 64-65.</ref>

Beyond steppe eagles, comparisons to various other groups of sympatric booted eagles may be made. Compared to the spotted eagles, the tawny eagle's tail is longer, the bill more prominent, the wings usually less squared-off in flight, the neck longer and the overall look rangier, despite these species being of often similar size. In contrast to the imperial eagles, the wings of the tawny eagle are broader and have less even trailing edges, the bill is slightly less prominent, and the wings are more likely to be held slightly upwards, while the body size is smaller. When compared to the golden eagle species complex, of which only the golden and the Verreaux's eagle (Aquila verreauxii) are usually relevant (although three dissimilar and sympatric pale-bellied eagles, of a size with tawny eagles or smaller, are found as part of this evolutionary chain), the tawny eagle is considerably smaller, its wings do not taper as much nor are they as likely to be held in a strong dihedral, and proportionately, the bill is notably longer and the tail is rather shorter.<ref name="Helbig" /><ref name="Naoroji" /> Greater spotted eagles (Clanga clanga), like the steppe eagle a Palearctic breeding eagle who often winters in the resident range of tawny eagles, is fairly similar, but that species has a relatively shorter and broader tail, less baggy feathers on the legs and usually a rather darker and more uniform adult plumage.<ref name="Ferguson-Lees" /> The fulvescens form of the greater spotted eagle must be distinguished from the pale forms of the tawny eagle by its underwing pattern, often with completely blackish underwing coverts and usually plain looking dark remiges over the entire primaries with more distinct pale carpal arcs.<ref name="Ferguson-Lees" /><ref name="Davidson">Davidson, I. (1978). Flight identification of southern African raptors. Bokmakierie, 30: 43-48.</ref> The likewise migratory lesser spotted eagle (Clanga pomarina) is smaller than the tawny eagle and more compact with a distinct white U above the tail.<ref name="Ferguson-Lees" /><ref name="Davidson" /> The residential African Wahlberg's eagle (Hieraeetus wahlbergi) can have a similar uniform plumage as in tawny eagles but always has greyer flight feathers and is much smaller than tawny eagles with relatively longer and more rectangular wings and a longer, narrower and straighter-tipped tail.<ref name="Ferguson-Lees" /><ref name="Davidson" /> The eastern imperial eagle in juvenile plumage can appear similar to the pale and intermediate morph tawny eagles, but the imperial eagle is usually visibly larger, with slenderer, longer wings, a longer, broader tail as well as having dark brown streaking on the chest, mantle and wing coverts and bearing more distinct pale trailing edges and wing bars.<ref name="Ferguson-Lees" /><ref name="Shirihai" />

Dark-morph tawny eagles in India may be distinguished from similarly sized black eagles (Ictinaetus malaiensis) by the latter being slenderer and having longer, darker and more paddle-shaped wings with a narrower base and a much longer, narrower and distinctly barred tail.<ref name="Ferguson-Lees" /> More dissimilar eagles such as Circaetinae, i.e. brown snake eagles (Circaetus cinereus), black-breasted snake eagles (Circaetus pectoralis) and juvenile bateleurs (Terathopius ecaudatus), are sometimes mentioned as a potential source of confusion but are usually rather distinct (all larger headed, rather smaller billed, shorter tailed and bare legged with often less uniform coloring) even in their most similar hues.<ref name="Steyn" /><ref name="Davidson" />

Calls

Tawny eagles are generally silent in most of their range.<ref name= Ferguson-Lees/> However, unlike steppe eagles, which are almost always silent away from their breeding grounds, they are said to occasionally vocalize in any season.<ref name= Shirihai/> They are also more vocal when not breeding than the spotted eagles.<ref name= Ferguson-Lees/> The usual call is a harsh, hollow-sounding, loud bark, variously transcribed as kowk-kowk, kau-kau, kiok-kiok or ki-ark. The call is fairly high-pitched (slightly less deep than the steppe eagle's when the latter is breeding) but is still deeper voiced than spotted eagles.<ref name= Ferguson-Lees/> In Kruger National Park it is said the call is loud and far-travelling.<ref name= Steyn/> Male tawny eagles are the most frequent vocalizers in the species, particularly during sky-dances, but also in other contexts. These include but are not limited to food arguments, disturbances during nesting and males attracting females for food passes.<ref name= Ferguson-Lees/> In nine years of monitoring tawny eagles in Zimbabwe, however, the call was not heard once.<ref name= Steyn2/> Its silence there may be due to the flat landscape.<ref name= Steyn2/> Other call recorded include a harsh grating k eke ke... in aerial courtship displays and a throaty kra in kleptoparastic pursuits. A kra-kra call may emitted at times to warn intruders.<ref name= Ferguson-Lees/><ref name= Naoroji/> The female may also emit an occasional mewing, high shreep-shreep at the nest as well as a rare raucous scream (possible food-begging and alarm calls, respectively).<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name= Davison>Davison, B. (1998). Raptor communities in hill habitats in south-eastern Zimbabwe (Doctoral dissertation, Rhodes University).</ref> The young chick tawny eagle chips initially but once its feathers emerge, it tends to beg with a loud call, i.e. we-yik, wee-yik.<ref name= Steyn/>

Distribution and habitat

Tawny eagles have an extremely extensive natural distribution. The African population can be found in three, fairly discrete populations.<ref name= Ferguson-Lees/> One of these is found in North Africa in south-central Morocco, possibly northern Algeria, southwestern Mauritania, Senegambia, southern Mali, central and southern Niger eastward through southern Chad, northern and central Sudan to most of Ethiopia and Somalia (but for the northeast and central-east).<ref name="iucn status 12 November 2021" /><ref name= Ferguson-Lees/><ref name= Ash/><ref>Isenmann P. & Moali A. (2000). Birds of Algeria. Société d‟Etudes Ornithologiques de France, Paris.</ref><ref>Nikolaus, G. (1987). Distribution atlas of Sudan's birds with notes on habitat and status. Zoologisches Forschungsinstitut und Museum Alexander Koenig.</ref><ref name= Smith>Smith, K. D. (1957). An annotated check list of the birds of Eritrea. Ibis, 99(2), 307-337.</ref><ref>Ash, J. S., & Miskell, J. E. (1998). Birds of Somalia. Pica Press.</ref> The north African population is scarce. In Morocco, they are heavily depleted with a few populations left in some regions such as Tarfaya, Tan-Tan and Souss-Massa.<ref name= Thevenot>Thévenot, M., Vernon, R., & Bergier, P. (2003). The birds of Morocco: an annotated checklist (No. 20). British Ornithologists' Union.</ref> They are likely extirpated from Tunisia, where they were once frequent.<ref>Isenmann, P. (2005). Birds of Tunisia. Société d'études ornithologiques de France, Muséum national d'histoire naturelle.</ref> In West Africa, some tawny eagles occur in Gambia, Togo, Nigeria and (though possibly not breeding) in Ivory Coast and Ghana.<ref>Borrow, N., & Demey, R. (2001). A guide to the birds of western Africa. Princeton, NJ.</ref><ref>Gore, M. E. J. (1990). Birds of the Gambia . Checklist 3. British Ornithologists' Union, Tring.</ref><ref name= Thiollay>Thiollay, J. M. (1985). The birds of Ivory Coast: status and distribution. West African Ornithological Society.</ref><ref>Greig-Smith, P. W. (1976). The composition and habitat preferences of the avifauna of Mole National Park, Ghana. Bull. Nigerian Orn. Soc, 12(42), 45-66.</ref><ref name= Dowsett-Lemaire>Dowsett, R. J., Dowsett-Lemaire, F., & Hester, A. (2008). The avifauna of Ghana: additions and corrections. Bull. Afr. Bird Club, 15, 191-200.</ref><ref name= Cheke>Cheke, R. A., & Walsh, J. F. (1996). The birds of Togo: an annotated check-list (No. 14). British Ornithologists' Union.</ref><ref>Elgood, J. H., Heigham, J. B., Moore, A. M., Nason, A. M., Sharland, R. E., & Skinner, N. J. (1994). The birds of Nigeria: an annotated check-list. British Ornithologists' Union.</ref> In east Africa and central Africa, the tawny eagle is found in central and eastern Democratic Republic of the Congo and throughout the drier portions of Uganda and in the entire nations of Kenya, Tanzania, Zambia (quite often residing in the Luangwa valley and the Chambeshi drainage), Malawi and Mozambique. In east Africa, it is considered perhaps the most widely distributed and regularly sighted brown eagle.<ref>Carswell, M., Pomeroy, D., Reynolds, J., & Tushabe, H. (2005). The Bird Atlas of Uganda. British Ornithologists' Club and British Ornithologists' Union.</ref><ref>Dowsett-Lemaire, F., & Dowsett, R. J. (2006). The birds of Malawi: an atlas and handbook. Tauraco & Aves.</ref><ref>Stevenson, T., & Fanshawe, J. (2002). Field guide to the birds of East Africa. T & AD Poyser.</ref><ref name= Dowsett>Dowsett, R. J., Aspinwall, D. R., & Dowsett-Lemaire, F. (2008). The birds of Zambia: an atlas and handbook. Tauraco Press.</ref> In southern Africa, the tawny eagle is found throughout Zimbabwe (now often rare apart from Matabeleland and Chipinga Uplands), Botswana (still regular in Okavango Delta) and some areas of Namibia, southern and western Angola (Cuando Cubango, Cunene, Huíla Namibe, to Malanje), Eswatini, Lesotho and northern and central parts of South Africa, i.e. mainly north of the Orange River but sometimes down to the Cape Province.<ref name="iucn status 12 November 2021" /><ref name= Steyn/><ref>Dean, W. R. J. (2000). The birds of Angola: an annotated checklist. British Ornithologists' Union.</ref><ref>da Rosa Pinto, A. A. (1983). Ornitologia de Angola (Vol. 1). Instituto de Investigação Científica Tropical.</ref><ref>Parker, V. (2005). The atlas of the birds of central Mozambique. Endangered Wildlife Trust and the Avian Demography Unit.</ref><ref name= Irwin>Irwin, M. P. S. (1981). The birds of Zimbabwe. Quest Pub.</ref><ref name= Penry>Penry, H. (1994). Bird atlas of Botswana. University of Kwazulu Natal Press.</ref><ref>Hancock, P., Muller, M., & Tyler, S. J. (2007). Inventory of Birds of the Okavango Delta Ramsar Site. Babbler, 49, 3-29.</ref><ref name= Barnes>Barnes, K. N. (Ed.). (2000). The Eskom red data book of birds of South Africa, Lesotho and Swaziland. BirdLife South Africa.</ref><ref>Boshoff, A. F., & Vernon, C. J. (1980). The distribution and status of some eagles in the Cape Province. Annals of the Cape Provincial Museums (Natural History), 13(9), 107-132.</ref> The tawny eagle may be extinct as a breeder in Eswatini where it was last confirmed to have bred in 2001.<ref>Monadjem, A., & Rasmussen, M. W. (2008). Nest distribution and conservation status of eagles, selected hawks and owls in Swaziland. Gabar, 19, 1-22.</ref>

File:Steppe Eagle, Pokaran, district Jaisalmer, Rajasthan, India.jpg

A tawny eagle perched in Rajasthan in India.

Out of Africa, the species may possibly be found in the southwestern part of the Arabian Peninsula, i.e. in Yemen and extreme southwestern Saudi Arabia in the Tihamah and 'Asir Regions, but few to none confirmed breeding events have been reported in the last few decades.<ref name="iucn status 12 November 2021" /><ref>Jennings, M. C. (1991). The changing avifauna of Saudi Arabia. Journal of Saudi Arabian Natural History Society, 3, 16-28.</ref><ref>Martins, R. P., & Porter, R. F. (Eds.). (1996). Southern Yemen and Socotra: The Report of the OSME Survey in the Spring 1993. Ornithological Society of the Middle East.</ref> The tawny eagle is considered a rare vagrant in Israel, though some are verified, other reports of them often turn out to be misidentified steppe eagles.<ref>Shirihai, H., Dovrat, E., Christie, D. A., & Harris, A. (1996). The birds of Israel. London: Academic Press.</ref> They are also known as a rare vagrant in Oman.<ref>Eriksen, J., Sargeant, D.E. & Victor, R. 2003. Oman Bird List (6th Edition). Centre for Environmental Studies & Research, Sultan Qaboos University, Muscat, Oman</ref> In Asia, the tawny eagle exists in isolation in southeastern Iran (as in Arabia, verified recent breeding is not known) and somewhat more continuously in eastern Pakistan (often in the Indus valley), much of north and peninsular India, eastward scarcely through southern Nepal and Assam.<ref name= Ferguson-Lees/><ref name= Rasmussen>Rasmussen, P. C., & Anderton, J. C. (2005). Birds of south Asia: the Ripley guide. Washington, DC.</ref><ref>Scott, D. A., & Adhami, A. (2006). An updated checklist of the birds of Iran. Podoces, 1(1/2), 1-16.</ref><ref>Roth, T., Ayé, R., Burri, R., & Schweizer, M. (2005). Bird observations from Iran in February–March 2001, including a new species for the Middle East. Sandgrouse, 27: 63-68.</ref> Though Nepali tawny eagles are rarely recorded, it is thought that the species still resides there in lowland semi-deserts.<ref>Inskipp, C. & Inskipp, T. (1991). A guide to the birds of Nepal. Second edition. Christopher Helm, London.</ref> The Indian range is from Punjab through the Indo-Gangetic Plain and western Bengal, northeastern Bihar, the Deccan Plateau with range continuing down to Andhra Pradesh, Karnataka and (mainly north-central) Tamil Nadu.<ref name= Naoroji/><ref name= Rasmussen/><ref>Donald, C. H. (1919). Some birds of prey of Mesopotamia. Journal of the Bombay Natural History Society, 26: 845-846.</ref> Records of vagrating tawny eagles turning up in Myanmar, northern Vietnam and Thailand are thought to have been likely misidentified steppe eagles or are based on now unidentifiable specimens.<ref name= Duckworth>Duckworth, J. W., Inskipp, T. P., Pasquet, E., Rasmussen, P. C., Rice, N. H., Robson, C. R., & Russell, D. G. D. (2008). A re-evaluation of the status of Tawny Eagle Aquila rapax in South-East Asia.</ref> A small handful of vagrants have been verified to turn up in Sri Lanka (the only known appearance by an Aquila eagle there).<ref>Warakagoda, D. (2015). Photographic Guide to the Birds of Sri Lanka. A & C Black Limited.</ref> Old reports of vagrancy, probably in need of confirmation, are known also from Afghanistan.<ref name= Rasmussen/>

Habitat

File:Bushveld.jpg

Tawny eagles are often at home in the arid and yet rich bushveld.

Tawny eagles occurs in fairly open country at varied elevations but usually live in drier areas. In West Africa, the species breeds often in relatively moist forest-savanna mosaics but can move into dry woodlands and semi-deserts when not breeding.<ref name= Ferguson-Lees/> In Morocco, the species prefers forested areas near mountains with adjacent plains. Elsewhere in Africa, tawny eagles typically inhabit wooded savanna such as dry Acacia savanna and semi-desert to desert areas. However, extreme desert areas, completely lacking in arborescent growth, are avoided nearly as much so as humid tropical rainforests. It also occurs at times in manmade areas such as arable lands, roadsides, dams, farmland, cattle pastures and game areas if feeding opportunities occur in them.<ref name= Ferguson-Lees/><ref name= Ash/><ref name= Dowsett/><ref name= Irwin/><ref name= Penry/><ref>Fry, C. H. (1966). The ecological distribution of birds in northern Guinea savanna, Nigeria. Ostrich, 37 (1): 335-356.</ref> In southern Africa, thornveld is often the preferred habitat with the tawny eagles mostly preferring stands of Acacia. Despite similar climates, within the miombo woodland, the tawny eagle tends to be more scarce.<ref name= Steyn/><ref name= Dowsett/><ref name= Irwin/> In India, similar habitats may largely be used but the tawny eagle may fairly often occur too in the vicinity of villages and cultivations and frequents garbage dumps and slaughterhouses somewhat more so than they do in Africa.<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name= Rasmussen/> In addition to all gradients of arid zones, in India, the tawny eagle frequently is found around thorn forests.<ref name= Naoroji/> Tawny eagles may live from sea level to about Template:Convert but tends to prefer somewhat lower elevations.<ref name= Ferguson-Lees/> Despite a certain level of aridity expected in tawny eagle habitats, they normally will not nest unless a habitat meets certain demands.<ref name= Hustler/> The tawny eagle's presence is predicated on the availability of ephemeral rainfall during the wet season.<ref name= Hustler/><ref name= Hustler2>Hustler, K., & Howells, W. W. (1990). The influence of primary production on a raptor community in Hwange National Park, Zimbabwe. Journal of Tropical Ecology, 6(3), 343-354.</ref><ref name= Herholdt>Template:Cite journal</ref> This reliance on some rainfall is probably key to habitat quality and resulting prey populations to some extent, but also to the availability of nesting sites.<ref name= Steyn/><ref name= Hustler2/> The tawny eagle is by and large an obligate tree nester and so areas that become too arid to support tree growth or where trees are overharvested are unlikely to retain the species.<ref name= Naoroji/><ref name= Steyn/><ref>Malan, G., & Howells, B. (2009). Tree nest surveys. Raptor Survey and Monitoring: A Field Guide for African Birds of Prey, 42.</ref>

Behaviour

File:Tawny Eagle (Aquila rapax) 1.jpg

A tawny eagle in flight.

The tawny eagle, quite unlike the steppe eagle, is largely sedentary and non-migratory. However, in Africa it is at times considered to be fairly nomadic and can engage in some seasonal movements.<ref name= Ferguson-Lees/><ref name= Thiollay2>Thiollay, J. M. (1992). Patterns and ecology of seasonal migrations of Ethiopian raptors in West Africa. In 7th Pan-African Ornithological Congress, Nairobi (Vol. 28).</ref> In west Africa, A. r. belisarius rather regularly travels shorter distance to damp woodlands during October through November, returning north in April, and perhaps at least at times, migrates into Kalahari region of Botswana and may vagrate to southern South Africa.<ref name= Ferguson-Lees/><ref name= Thiollay/><ref name= Dowsett-Lemaire/><ref>Snelling, J. C. (1970). Some information obtained from marking large raptors in the Kruger National Park, Republic of South Africa. Ostrich suppl., 8: 415-427.</ref> Sometimes, tawny eagles seem to ping semi-regularly between Ethiopia and west Africa.<ref name= Thiollay2/> Some long distance wandering has even been reported, such as a vagrant A. r. belisarius in Tunisia and as far as Egypt (where twice recorded in the 1950s) and even Israel (3 winter records in the 1990s) and Oman.<ref name= Ferguson-Lees/><ref>Ławicki, Ł., Jiguet, F., El Din, S. B., van den Berg, A. B., Corso, A., Crochet, P. A., Hoath, R., Schweizer, M. & Waheed, A. (2019). Sixth report of the Egyptian Ornithological Rarities Committee.</ref><ref>Arnold, P. K. (1965). Birds of Israel. Shalit Publishers.</ref><ref>Kinnear, N. B. (1931). On some birds from central South Arabia. Ibis, 73(4), 698-701.</ref> From the Indian range, individuals vagrate not infrequently to nearby Bangladesh, most likely as juveniles post-dispersal wanderings, but reports of the species wandering into southeast Asia such as Thailand are now considered likely apocryphal.<ref name= Duckworth/><ref>Jahan, I., Thompson, P. M., Paul, E., Sultana, N., Roy, K., & Mallick, A. (2017). Birds of Bhawal National Park, Bangladesh. Forktail, (33), 88-91.</ref> Generally, in areas such as southern Africa, tawny eagles usually seldom seem to leave their established breeding territories and juvenile eagles generally wander no more than several dozen kilometres from their original nest.<ref name= Steyn/><ref>Oatley, T.B., Oschadleus, H.D., Navarro, R.A. & Underhill, L.G. (1998). Review of ring recoveries of birds of prey in southern Africa: 1948–1998. Endangered Wildlife Trust, Johannesburg.</ref> A bird banded as a nestling in Esigodini was recovered quite nearby at Fort Rixon more than two years later.<ref name= Irwin/> However, in a rather far dispersal for southern Africa, one eagle banded as a nestling was recovered Template:Convert away from its nest of origin in Zimbabwe four years later.<ref name= Dowsett/> Inconsistent and seemingly unpredictable movements by tawny eagles have been proven via experimental ecological studies to be actually be instances of eagles searching out new areas to compensate for lack of rainfall.<ref name= Wichmann2>Wichmann, M. C., Groeneveld, J., Jeltsch, F., & Grimm, V. (2005). Mitigation of climate change impacts on raptors by behavioural adaptation: ecological buffering mechanisms. Global and Planetary Change, 47(2-4), 273-281.</ref> While non-breeding steppe eagles are often slightly social and flock at opportunistic feeding sources, the tawny eagle is usually considered solitary.<ref name= Ferguson-Lees/><ref name= Naoroji/> However, groups of two to three tawny eagles are sometimes seen, such as in the Indian subcontinent, but occasionally group sizes may even exceed this figure.<ref name= Naoroji/> In the Mirpur Division of Azad Kashmir in Pakistan, small flocks of tawny eagles have reportedly been witnessed gathering in warmer spots between November and February, over three years of study.<ref>Shakil, M. (2004). Aquila Rapax. In: The Eagles of Kashmir. Self-Published.</ref> Small groups or aggregations are known to occur in Africa as well near concentrated foods and even communal roost have been reported in trees, power pylons or on the ground.<ref name= Ferguson-Lees/> Like many large raptors, the tawny eagle probably spends the majority of its day perched but take wing a few times a day.<ref name= Naoroji/><ref name= Steyn/> Unlike most large eagles, in India at least, tawny eagles are often fairly accustomed to humans and may allow fairly close approach by observers.<ref name= Naoroji/><ref>Dharmakumarsinhji, R.S. (1949). Birds of prey as companions. Avicultural Magazine, 55: 81-85.</ref>

Feeding

File:Aquila rapax -Ethiopia -with roadkill-8.jpg

With a black-backed jackal, road traffic victim in Ethiopia.

The tawny eagle is unique as an Aquila eagle in the lack of apparent specialization in its feeding behaviour.<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name= Steyn/> While most other Aquila will opportunistically scavenge, the tawny eagle freely takes to scavenging on carrion, perhaps doing so at all times of the year, though do so somewhat more so when not breeding.<ref name= Naoroji/><ref name= Steyn/> Routine scavenging often drives tawny eagles to refuse dumps in or near villages and slaughterhouses, particularly in India, and to associate quite often with vultures at carrion sites. They are also frequently recorded on roadsides where roadkill provides a steady food source.<ref name= Ferguson-Lees/><ref name= Naoroji/> More routinely than almost any other raptor, perhaps, it is a very skilled pirate, regularly engaging in kleptoparasitism of other birds of prey.<ref name= Naoroji/><ref name= Steyn/> However, descriptions of the tawny eagle as "sluggish", "not very distinguished" and "unimpressive" are not particularly apt as the tawny eagle is a highly rapacious predator that attacks very variable ranges of live prey including particularly large prey.<ref name= Steyn/><ref name= Clark/> This species hunts mainly by a short dive or pounce from a perch or by stoop from up high in a soaring flight.<ref name= Ferguson-Lees/> In the Indian subcontinent, preferred hunting perch trees were Vachellia nilotica, Prosopis cineraria and Capparis decidua.<ref name= Naoroji/> It may also often forage by walking on the ground. Mostly, tawny eagles target live prey that is on the ground, seldom targeting arboreal prey.<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name= Steyn/> However, they will sometimes fly down and take birds on the wing.<ref name= Naoroji/><ref name= Steyn/> Avian prey known to be killed in the air has ranged from speckled pigeons (Columba guinea) to flamingoes.<ref name= Steyn/> Tawny eagles may hunt frequently in pairs during the breeding season, often securing larger prey than in the non-breeding season.<ref name= Naoroji/> Sometimes this tandem hunting by pairs can occur in any season.<ref name= Steyn/> It is likely that tandem hunting by pairs involves one bird engaging in conspicuous flight to distract the quarry while another flies inconspicuously to sneak up on and kill the prey, as has been reported in other tandem hunting raptors.<ref name= Naoroji/><ref name= Steyn/><ref>Ellis, D. H., Bednarz, J. C., Smith, D. G., & Flemming, S. P. (1993). Social Foraging Classes in Raptorial Birds: Highly developed cooperative hunting may be important for many raptors. Bioscience, 43(1), 14-20.</ref> Nocturnal animals such as genets and springhares have been preyed upon by tawny eagles in areas where there was no possibility they were killed by traffic at night. In addition to other observations have been made where tawny eagles drink and bathe at night, some nocturnal subsistence behaviour by this species has been inferred but no irrefutable evidence has been brought forth either.<ref name= Steyn/><ref>Barbour, D. (1975). Do Eagles use moonlight to hunt? Honeyguide, 82: 37.</ref> Semi-regular attendance at grassfires in India, presumably in order to capture displaced creatures, has been reported.<ref name= Rasmussen/> More than 200 species, including both live prey and carrion, are known to be eaten by tawny eagles and they may have one of the most variable diet of all tropical eagles.<ref name= Naoroji/><ref name= Steyn/><ref name= Smeenk/><ref name= Steyn2>Template:Cite journal</ref><ref name= Osborne>Template:Cite journal</ref><ref name= Vernon>Vernon, C. J. (1979). Prey remains from seven Tawny Eagle nests. Honeyguide, 100: 22-24.</ref> Reportedly, most prey the tawny eagle will take alive will weigh not less than Template:Convert and not more than Template:Convert, however live prey has been revealed to be regularly more variable than even that estimate represents.<ref name= Ferguson-Lees/><ref name= Steyn/><ref name= Smeenk/><ref name= Steyn2/> One compilation study showed that, compared to 8 other Aquila and spotted eagles, the tawny eagle's diet was the most evenly spread across all weight classes of prey from under Template:Convert to over Template:Convert, though took prey in the latter prey class slightly less so than the much larger golden and wedge-tailed eagles (Aquila audax). This study further determine that the most often focused on weight class in tawny eagle's diets were Template:Convert and Template:Convert prey class, accounting for a little less than half of the prey by quantity.<ref name= Watson/> According to this authority, the mean prey size falls around approximately Template:Convert, which is around 5 times greater than the mean estimated prey size for the steppe eagle species, around 38% greater than mean estimated prey size of imperial eagles and considerably less only than the golden, wedge-tailed and Verreaux's eagles among the 8 studied Aquila and Clanga species.<ref name= Watson/>

Probable live prey

File:Tawny Eagle (Aquila rapax) juvenile eating a White-throated Monitor Lizard (Varanus albigularis) (16517371167).jpg

A tawny eagle with its prey, a rock monitor.

Determining whether prey has been taken alive at the nests of tawny eagles is generally considered to be difficult, although observations suggest that during breeding tawny eagles usually deliver fresh prey while raising young, indicating that such prey are usually either taken alive or newly pirated from other predators.<ref name= Steyn/><ref name= Steyn2/><ref name= Osborne/> Within a Kenyan study, only 1.9% of prey brought to tawny eagle nests was thought likely to be from carrion.<ref name= Smeenk/> While hunting prey, it often takes a variety of prey usually focusing somewhat on small to medium-sized mammals, usually medium-sized to large birds and occasionally medium to large-sized reptiles.<ref name= Steyn/><ref name= Smeenk/><ref name= Steyn2/> A diet analysis in Zimbabwe indicated that among 160 prey items, 36.9% were mammals, 51.9% were birds, 10% were reptiles and 1.2% were amphibians.<ref name= Steyn2/> The leading prey species here were scrub hare (Lepus saxatilis), at about 15.6% of the total, helmeted guineafowl (Numida meleagris), at about 11.9% of the prey total, both chickens and unidentified snakes at around 6.9% and indeterminate spurfowl and doves at 6.2% and 5%, respectively.<ref name= Steyn2/> A similar dietary study conducted in Lochinvar National Park, Zambia found a higher proportion of birds and amphibians (61.4% & 5.5% respectively), with a surprisingly number of water birds being taken. Of 127 prey items here, the main prey species were determined to be helmeted guineafowl at about 15% of the diet, Swainson's spurfowl (Pternistis swainsonii) at 11%, poultry at about 8.7%, the African openbill (Anastomus lamelligerus) at 7.9% with the top mammals in the diet being scrub hares, at 5.5%, and African marsh rats (Dasymys incomtus) at 3.9%; furthermore indeterminate snakes comprised 5.5% of the diet.<ref name=Osborne/> The variation in diet between the preceding two study sites is due to differences in habitat and prey availability.<ref name= Osborne/> A very detailed study was conducted in Tsavo East National Park, Kenya, of the diet of the local tawny eagles over different years. Of 543 total prey items, 41.2% were mammals, 35.4% were birds, and 23.4% were reptiles and amphibians. Of these the most important prey was Kirk's dik-dik (Madoqua kirkii) at 21.7% of the total, yellow-necked spurfowl (Pternistis leucoscepus) at 6.8% of the total, cape hare (Lepus capensis) and crested francolin (Dendroperdix sephaena) both at 6.3% and red-crested korhaan (Lophotis ruficrista) at 5.9%; additionally, unidentified snakes constituted 21.3% of the foods.<ref name= Smeenk/> The dik-dik is a small antelope and is much larger than a tawny eagle. The tawny eagle certainly took dik-diks weighing up to Template:Convert and possibly even Template:Convert, with this eagle taking about 80 dik-diks in Tsavo East each year.<ref name= Smeenk/> In different areas of South Africa, small dietary studies determined the diet at nests to be highly variable. In the Highveld, about 52% of 60 prey items were mammals, 45% were birds and a small number were fish. In Timbavati and Klaserie, 63% of the diet was birds, 34% of it was mammals and 3% were reptiles. At Highveld, 25% of the diet consisted of yellow mongoose (Cynictis penicillata), 15% Cape ground squirrels (Xerus inauris) and 13% helmeted guineafowl. At Timbavati and Klaserie, various francolins were strongly predominant in foods, at about 44% with another 17% by various mongooses.<ref name= Tarboton>Tarboton, W.R. & Allan, D.G. (1984). The status and conservation of birds of prey in the Transvaal. Transvaal Museum Monograph No. 3. Pretoria.</ref><ref>Boshoff, A. F., Rous, R. C., & Vernon, C. J. (1981). Prey of the tawny-eagle in the colesberg district, Cape Province. Ostrich, 52(3), 187-188.</ref>

Out of the southern and eastern areas of Africa, less quantitative analysis has been undertaken into the feeding habits of tawny eagles, even around nests. What is known of their prey elsewhere is mainly from wide-ranging surveys, secondary accounts and photographs.<ref name= Naoroji/><ref name= Steyn/> It appears in Ethiopia that the tawny eagle may have a close predatory relationship with the Abyssinian grass rat (Arvicanthis abyssinicus) while one tawny eagle there tore open the nest of the Stresemann's bushcrow (Zavattariornis stresemanni) to access the prey.<ref name= Ash/><ref>Muller, J. P. 1977. Populationsokologie von Arvicanthis abyssinicus in der Grassteppe des Semien Mountain National Park (Athiopien). Z. Saugetierk, 42 (3): 145-172.</ref><ref name="Clouet"/> Unidentified large rats constituted a great majority of prey delivered during the nestling growth stage at some east African nests.<ref name= VanSomeren/> Fewer details are known about the prey of tawny eagles in the Indian subcontinent.<ref name= Naoroji/> One study, without quantitative data known, listed the prey of tawny eagles in Saurashtra as Indian palm squirrels (Funambulus palmarum), eggs of red-wattled lapwings (Vanellus indicus), crows, mongooses, Indian hares (Lepus nigricollis) and even Bengal fox (Vulpes bengalensis).<ref name= Dharmakuarsinhji>Dharmakuarsinhji, K.S. (1955). Birds of Saurashtra. Dil Bahar.</ref> Otherwise, the prey incidentally reported in India is extremely varied, including even the hindlegs of a jungle cat (Felis chaus) (but this may have been scavenged).<ref name= Naoroji/> In general, tawny eagles in south Asia may focus on less varying prey in general, often reportedly favoring desert-dwelling rodents and hares.<ref name= Rasmussen/><ref name= Fitzwater>Fitzwater, W. D., & Prakash, I. (1969). Observations on the burrows, behavior and home range of the Indian desert gerbil, Meriones hurrianae Jerdon. Mammalia, 33(4), 598-606</ref><ref name= Prakash>Prakash, I., & Ghosh, P. K. (Eds.). (2012). Rodents in desert environments (Vol. 28). Springer Science & Business Media.</ref> Tawny eagles may hunt a couple dozen species of rodent from different parts of the range, ranging in size from the Template:Convert Natal multimammate mouse (Mastomys natalensis) to the Template:Convert South African springhare (Pedestes capensis).<ref name= Fitzwater/><ref name= Prakash/><ref>Giban, J. (1978). Control of the multimammate rat, Mastomys natalensis (A. Smith) in the irrigated fields of the Republic of Burundi. In Proceedings of the 8th Vertebrate Pest Conference (1978) (p. 20).</ref> Rock hyrax (Procavia capensis) and yellow-spotted rock hyrax (Heterohyrax brucei) are occasionally preyed upon by tawny eagles.<ref name= Kingdon>Kingdon, J., Happold, D., Butynski, T., Hoffmann, M., Happold, M., & Kalina, J. (2013). Mammals of Africa. A&C Black.</ref><ref>Mbise, F. P., Fredriksen, K. E., Fyumagwa, R. D., Holmern, T., Jackson, C. R., Fossøy, F., & Røskaft, E. (2017). Do hyraxes benefit from human presence in Serengeti? African journal of ecology, 55(4), 672-678.</ref> The tawny eagle may be one of the most accomplished predators of mongoose, many food studies reflecting relatively high numbers of them and they appear to be one of the most feared predators at meerkat (Suricata suricatta) colonies.<ref name= Ash/><ref name= Irwin/><ref>Dean, W. R. J., Huntley, M. A., Huntley, B. J., & Vernon, C. J. (1988). Notes on some birds of Angola. Novitates, Durban Museum, 14(4), 43-92.</ref><ref>Beynon, P., & Rasa, O. A. E. (1989). Do dwarf mongooses have a language-warning vocalizations transmit complex information? South African Journal of Science, 85(7): 447-450.</ref><ref>Manser, M. B., Seyfarth, R. M., & Cheney, D. L. (2002). Suricate alarm calls signal predator class and urgency. Trends in cognitive sciences, 6(2), 55-57.</ref> In the Karoo, the local tawny eagles reportedly live mostly off of mongooses, either meerkats or yellow mongooses.<ref name= Tarboton2>Tarboton, W. R., Pickford, P., & Pickford, B. (1990). African birds of prey. Cornell University Press.</ref> Similarly small or slightly larger carnivores like striped polecats (Ictonyx striatus) and genets are not infrequently prey for tawny eagles as well.<ref name= Steyn/><ref name= Tarboton/> Bat-eared foxes (Otocyon megalotis) may too be taken alive at times.<ref>Malcom, J. R. (1986). Socio‐ecology of Bat‐eared foxes (Otocyon megalotis). Journal of Zoology, 208(3), 457-469.</ref> Various monkeys may be eaten, although not infrequently as carrion, tawny eagles may too in seldom cases attack juveniles of monkeys such as Patas monkeys (Erythrocebus patas), grivets (Chlorocebus aethiops) and vervet monkeys (Chlorocebus pygerythrus) up to the size of juveniles of several species of baboon.<ref name= Smeenk/><ref name= Kingdon/><ref>Mercier, S., Neumann, C., van de Waal, E., Chollet, E., de Bellefon, J. M., & Zuberbühler, K. (2017). Vervet monkeys greet adult males during high-risk situations. Animal Behaviour, 132, 229-245.</ref><ref>Pereira, M. E. (1988). Effects of age and sex on intra-group spacing behaviour in juvenile savannah baboons, Papio cynocephalus cynocephalus. Animal behaviour, 36(1), 184-204.</ref><ref>Pahad, G. (2010). Social behaviour and crop raiding in chacma baboons of the Suikerbosrand Nature Reserve (Doctoral dissertation, University of the Witwatersrand).</ref> However, unlike with larger eagles, the troops of certain baboons do not seem to regard tawny eagles as a threat based on their behavioural responses.<ref>Zinner, D., & Peláez, F. (1999). Verreaux's eagles (Aquila verreauxi) as potential predators of hamadryas baboons (Papio hamadryas hamadryas) in Eritrea. American Journal of Primatology: Official Journal of the American Society of Primatologists, 47(1), 61-66.</ref> While most ungulate prey other than dik-diks is probably largely scavenged as carrion or stolen from other predators, the small calves of ungulates such as Thomson's gazelle (Eudorcas thomsonii) are sometimes apparently killed by tawny eagles.<ref>Roberts, B. A. (2014). The trials of motherhood: Maternal behavior patterns and antipredatortactics in Thomson's gazelle (Gazella thomsonii), a hidingungulate (Doctoral dissertation, Princeton University).</ref> A general picture appears to emerge that tawny eagles quite often takes relatively large mammalian prey, surprisingly often creatures weighing up to Template:Convert such as hares, dik-diks, the young of other antelopes, hyraxes and so on.<ref name= Steyn/><ref name= Clark/><ref name= Steyn2/><ref>Dixon, J. E. W. (1968). Prey of Large Raptors. Ostrich, 39(3), 203-204.</ref>

File:Anagoria Waffenkiebitz Raubadler.jpg

A tawny eagle is mobbed by a blacksmith lapwing; these eagles are a potential menace to many different kinds of birds.

While mammals prey varies from rodents to hares, mongooses and small antelopes, the diversity and size range of bird taken may be even more impressive and more than 120 avian prey species have been reported in the prey spectrum.<ref name= Naoroji/><ref name= Steyn/><ref name= Smeenk/><ref name= Osborne/><ref name= Tarboton/> Included in the prey spectrum are various species of ducks and small or gosling geese,<ref name= Smeenk/><ref name= Tarboton/><ref>Douthwaite, R. J. (1978). Geese and Red‐knobbed Coot on the Kafue Flats in Zambia, 1970–1974. African Journal of Ecology, 16(1), 29-47.</ref> gamebirds, especially francolins and guineafowl,<ref name= Naoroji/><ref name= Roberts/><ref name= Smeenk/><ref name= Steyn2/> many doves and pigeons,<ref name= Steyn/><ref name= Ali/><ref>Hosseini-Moosavi, S. M., Behrouzi-Rad, B., Karimpour, R., & Nasab, S. M. A. (2013). Breeding Biology of Collared Dove (Streptopelia decaocto) in Khuzestan Province, Southwestern Iran. Беркут, 22(1), 51-54.</ref> mostly medium-sized species of bustard and hornbill<ref name= Roberts/><ref name= Smeenk/><ref name= Ali/><ref name= Tarboton/><ref>Combreau, O., & Smith, T. R. (1998). Release techniques and predation in the introduction of houbara bustards in Saudi Arabia. Biological Conservation, 84(2), 147-155.</ref><ref>Runo, M. (2000). Tawny Eagle feeding on Red-billed Hornbill. Honeyguide, 46: 24.</ref> and numerous water birds from small coursers, lapwings, rails and grebes to large flamingoes, storks and herons both small and large.<ref name= Steyn/><ref name= Smeenk/><ref name= Tarboton/> Both young and adult Old World flamingoes of both African species are known to be attacked on occasion, as well as white-breasted cormorants (Phalacrocorax lucidus) and great cormorants (Phalacrocorax carbo), all birds similar in size or somewhat heavier than the tawny eagle itself.<ref>Walters, J. R. (1990). Anti-predatory behavior of lapwings: field evidence of discriminative abilities. The Wilson Bulletin, 49-70.</ref><ref>Robinson, V. (2015). The Ecology of East African Soda Lakes: Implications for Lesser Flamingo (Phoeniconaias minor) Feeding Behavior (Ph.D. Dissertation). Leicester, England: University of Leicester.</ref><ref>Johnson, A., & Cézilly, F. (2009). The greater flamingo. A&C Black.</ref> Further impressive water bird prey includes reportedly spur-winged goose (Plectropterus gambensis), which weighs about twice as much as tawny eagle.<ref name= Steyn2/> Even larger avian prey are taken including a common crane (Grus grus) killed by a pair in Saurashtra (although it was an injured one) and presumably adult female Kori bustard (Ardeotis kori). If average-sized, these prey items likely weighed more than Template:Convert.<ref name= Dharmakuarsinhji/><ref name= CRC/><ref>Lichtenberg, E. M., & Hallager, S. (2008). A description of commonly observed behaviors for the kori bustard (Ardeotis kori). Journal of Ethology, 26(1), 17-34.</ref> More minor avian prey includes nightjars, coucals, sandgrouse, swifts, bee-eaters, kingfishers, rollers, wood hoopoes, turacos, parrots and several passerines.<ref name= Naoroji/><ref name= Steyn/><ref name= Ali/><ref name= Smeenk/><ref name= Tarboton/><ref>Jackson, H. D., & Slotow, R. (2002). A review of Afrotropical nightjar mortality, mainly road kills. Ostrich, 73(3-4), 147-161.</ref><ref>Raju Kasambe, G. (2010). Blue-tailed Bee-eaters are 'winter migrants' in and around Mumbai. Newsletter for Birdwatchers, 50(3), 33.</ref><ref>Brojerová, J. (2013). Referenční vokalizace papouška žako kongo (Psittacus erithacus).</ref><ref>Dean, W. R. J. (2004). Nomadic Desert Birds: With 32 Tables. Springer Science & Business Media.</ref> One small passerine the tawny eagle may routinely hunt is the super-abundant red-billed quelea (Quelea quelea).<ref>van Heerden J. (1977). Quelea predators. Bokmakierie, 29: 45.</ref> Near poultry farms, tawny eagles can take to not infrequently lifting free-range chickens (Gallus gallus domesticus) and other poultry, especially when the eagles must feed their young, thus in turn potentially drawing ire of local farmers.<ref name= Naoroji/><ref name= Steyn/><ref name= AliS/><ref>Vernon, C. J. (1967). Some observations from the journals of KW Greenhow. Ostrich, 38(1), 48-49.</ref>

Various snakes are taken opportunistically by tawny eagles and they can be quite bold about hunting venomous snakes.<ref>Parry-Jones, J. (1997). Eagle. Dorling Kindersley Ltd.</ref> In southern Africa and Kenya the following snakes have been identified in the foods of tawny eagles: Egyptian sand boa (Eryx colubrinus), young African rock python (Python sebae), speckled sand racer (Psammophis punctulatus), rufous beaked snake (Rhamphiophis oxyrhynchus), black-necked spitting cobra (Naja nigricollis), black mamba (Dendroaspis polylepis), boomslang (Dispholidus typus) and puff adder (Bitis arietans).<ref name= Steyn/><ref name= Smeenk/><ref name= Brown3>Brown, L. H. (1952). On the biology of the large birds of prey of the Embu district, Kenya colony. Ibis, 94(4), 577-620.</ref> They also hunt lizards not infrequently given the chance, usually favoring fairly large species but capable of taking those ranging from geckos to rock monitors (Varanus albigularis).<ref name= Steyn/><ref name= Ali/> At one nest in Zimbabwe, monitor lizards made up 29% of 83 prey items, but they were only 8% of 107 prey items of 3 other nests in the same park.<ref name= Steyn2/> One of the most frequently seen prey to be taken by tawny eagles in India have been Indian spiny-tailed lizard (Saara hardwickii).<ref name= Naoroji/><ref>Ramesh, M., & Sankaran, R. (2013). Natural History Observations on the Indian Spiny-tailed Lizard Uromastyx hardwickii in the Thar Desert. In Faunal Heritage of Rajasthan, India (pp. 295-310). Springer, New York, NY.</ref><ref>Baindur, A. (2009). The raptors and the agamid. Indian Birds, 5(1), 11-13.</ref> More minor prey have included turtles, frogs and toads and fish. A tawny eagle in southern Africa was seen to wade into shallow water and successfully pull out a largish catfish.<ref name= Steyn/><ref name= Osborne/><ref name= Tarboton/><ref>Howells, W. W. (1978). Stranded fish as a food source for large birds of prey. Honeyguide, 96: 18.</ref> Tawny eagles can also take communal nesting and swarming insects fairly frequently. These are generally termites, which can attract several of these eagles especially amongst non-breeding eagles and young ones. When visiting termites, the tawny eagles commonly eat alates and may, with an unusual lack of aggression, share the food source with several other birds of prey, including as many as a half dozen conspecifics.<ref>Dial, K. P., & Vaughan, T. A. (1987). Opportunistic predation on alate termites in Kenya. Biotropica, 19(2), 185-187.</ref><ref>Bussia, E., & Wijers, M. (2013). Foraging frenzy: more than 50 raptors at a termite swarm. Biodiversity Observations, 11-18.</ref> At times, tawny eagles can also be attracted to swarms of grasshoppers.<ref>Anene, C., & Vajime, C. G. (1990). Parasites, parasitoids and predators of Oedaleus senegalensis Krauss (Orthoptera: Acrididae) in Nigeria. International Journal of Tropical Insect Science, 11(1), 27-34.</ref> In one case, a tawny eagle was seen consuming the fruit of an Adansonia tree, an unusual instance of frugivory which is very rare in accipitrids other than one unusual species: the palm-nut vulture (Gypohierax angolensis).<ref name= Steyn/> A tawny eagle was once witnessed picking through elephant dung along with a vulture, presumably searching for dung beetles to consume.<ref>van Jaarsveld, J. (1987). Tawny Eagle and Whiteheaded Vulture eat together from elephant dung. Gabar, 2: 26.</ref>

Carrion

File:Tawny Eagle (Aquila rapax) and White-backed Vultures (Gyps africanus) (6001455023).jpg

A tawny eagle perched amongst white-backed vultures, with which they are often obligated to share carrion in Africa.

Although the tawny eagle does hunt for food, it also relies extensively on carrion as a food source.<ref name= Steyn/> Although most booted eagles and Aquila will opportunistically feed on carrion, none is known to do so as routinely as the tawny eagle.<ref name= Steyn/><ref name= Anderson/> They have been recorded feeding on a huge array of carcasses as large as African bush elephants (Loxodonta africana) and at least as small as vervet monkeys and perhaps even down to the size of a dove.<ref name= Steyn/><ref name= Kemp3/> Perhaps most frequently in Africa, tawny eagles will feed at carcasses of ungulates such as antelope.<ref name= Steyn/><ref name= Steyn2/> At least 30-40 different species of ungulate have been recorded as carrion food-sources for these eagles.<ref name= Naoroji/><ref name= Steyn/><ref name= Smeenk/><ref name= Steyn2/> At "vulture restaurants" in Ethiopia, feeding stations with dead livestock meant to mitigate the rapid decline in population of most African vultures, the tawny eagle was the second most often recorded scavenger at just under 35% of 1088 of recorded birds to feed at them.<ref>Mulualem, G., Tesfahunage, W., & Ayalew, S. (2015). Abundance and activity pattern of avifauna in ashewa local vulture restaurant dire dawa eastern Ethiopia. International Journal of Avian & Wildlife Biology, 1(1).</ref> The tawny eagle shares its carrion food sources almost invariably with vultures and usually with several other scavengers such as jackals and hyenas. Other birds that frequently also attend carrion are bateleurs, many other eagles (including steppe eagles) and marabou storks (Leptoptilos crumenifer).<ref name= Steyn/><ref name= Mundy>Mundy, P.J. (1982). The comparative biology of southern African vultures. Johannesburg. Vulture Study Group.</ref><ref name= Dean>Dean, W. R. J., & MacDonald, I. A. W. (1981). A review of African birds feeding in association with mammals. Ostrich, 52(3), 135-155.</ref> The producer-scrounger theory predicts that vultures rely on eagles for information on carcasses.<ref name= Ogada>Template:Cite journal</ref> Due to their smaller size, eagles, i.e. tawny eagles and bateleurs, are able to begin foraging earlier in the morning and are thus more likely to locate a carcass first.<ref name= Ogada/> In 91 observed carcass in southern Africa, tawny eagles were verified to be the first to find 5 of them. Furthermore, vultures usually arrived in less than 40 minutes (in 75% of cases) after the tawny eagles found the carcass.<ref name= CJ>Brown, C. J. (1982). The behaviour of a Tawny Eagle at carrion. Madoqua, 14 (1): 95-97.</ref> At large carcasses, there is a hierarchical social structure based on the size of the scavenger.<ref name= Mundy/> At Maasai Mara, the top scavengers were the considerably to slightly heavier mammals, i.e. spotted hyenas (Crocuta croctua), black-backed jackals (Canis mesomelas) and feral dogs (Canis lupus familiaris), then the lappet-faced vulture (Torgos tracheliotos), the Rüppell's griffon (Gyps rueppellii), followed by all other vultures with the tawny eagle and the bateleur in the second most and the most subordinate scavenger positions.<ref name= Kendall>Kendall, C. J. (2013). Alternative strategies in avian scavengers: how subordinate species foil the despotic distribution. Behavioral Ecology and Sociobiology, 67(3), 383-393.</ref> Similar scavenger hierarchies have been reported elsewhere as well.<ref name= Mundy/><ref name= Dean/> Bateleurs were the most likely to first find a carcass of the Maasai Mara scavengers and both the tawny and bateleur were considered as scavengers with "low competitive ability and high search efficiency".<ref name= Kendall/>

However, tawny eagles will at times be able to displace the smaller species of vulture such as hooded vultures (Necrosyrtes monachus) and Egyptian vultures (Neophron percnopterus), both of which are similar in weight to the tawny eagles themselves, with one tawny eagle even reportedly keeping as many as 20 vultures at bay at a carcass. In general, based on the literature, such an event of aggressiveness by this species at a large carcass would surely be unusual.<ref name= Steyn/><ref name= Steyn2/><ref name= Kendall/><ref>Plaskett, G. (1972). Tawny Eagle/vulture behaviour. Witwaterstrand Bird Club News Sheet, 80: 11.</ref> Tawny eagles do tend to be dominant over bateleur at carcasses, however.<ref name= Steyn/> Gyps or griffon vultures are usually the most numerous vultures in attendance at carrion and are considerably larger than tawny eagles but sometimes may briefly tolerate a tawny eagle to feed in their midst depending on the circumstances.<ref name= Steyn/><ref name= Houston>Houston, D.C. (1974). The role of griffon vultures Gyps africanus and Gys rueppellii as scavengers. J. of Zoo, London. 171: 35-46.</ref><ref name= Kemp3>Kemp, A. C., & Kemp, M. I. (1975). Observations on the White-backed Vulture Gyps africanus in the Kruger National Park, with notes on other avian scavengers. Koedoe, 18(1), 51-68.</ref> Usually, the larger the group is of griffon vultures, the less likely the tawny eagle is to get to feed.<ref name= Anderson/><ref name= Kendall/><ref name= Houston/><ref name= Kemp3/> The eagles not infrequently remain on the periphery of the vulture feeding frenzy and wait for pieces of flesh to appear. Often they will be able to pick up small scraps but will wait until the carcass is finished and few vultures remain to feed. The tawny eagle can benefit from leading other scavengers to carrion or feeding subsequently to them since, unlike the largest and most aggressive vultures, such as lappet-faced vultures and cinereous vultures (Aegypius monachus), the tawny eagle cannot tear open large carcasses on their own and tend rely on another source to access any bits of the nutritious viscera.<ref name= Naoroji/><ref name= Steyn/><ref name= Anderson/><ref name= Steyn2/><ref name= Mundy/><ref name= Houston/> The tawny eagles when finding an unopened large carcass have few feeding options although may eat the eyes in such circumstances, as was verified in the circumstance of a tawny eagle finding a horse (Equus ferus caballus) carcass offered by researchers.<ref name= CJ/> Roadkills are another feeding option as they are often torn asunder by impact with automobiles and the eagle may be able to (at least briefly) monopolize the carcass.<ref name= Naoroji/><ref name= Steyn/><ref name= Kemp/> Perhaps not coincidentally, in Maasai Mara, the tawny eagles were found to benefit from a carcass being nearer human habitations and in lower quality habitats relative to the other scavengers.<ref name= Kendall/> In particularly in India, scavenging tawny eagles tend to regularly occur at landfills where vultures seldom come but wintering steppe eagles may often feed alongside them seasonally.<ref name= Naoroji/><ref name= Ali/> Garbage dumps are also visited in different parts of Africa such as Uganda and Ethiopia by hungry tawny eagles.<ref>Pomeroy, D. E. (1975). Birds as scavengers of refuse in Uganda. Ibis, 117(1), 69-81.</ref><ref>Mesfin, K. (2014). Diversity and Abundance of Birds in Dbla Church Forest, Eastern Tigray, North Ethiopia. Journal of Zoology Studies, 1(5), 1-8.</ref> Semi-predaceous and aggressively disposed vultures, like white-headed vultures (Trigonoceps occipitalis) in Africa and red-headed vultures (Sacrogyps calvus) in India as well as the lappet-faced and cinereous vultures, tend to have little tolerance for tawny eagles, with the latter unlikely to approach until these aggressive vultures have had their fill.<ref name= Naoroji/><ref name= Steyn/><ref name= Steyn2/><ref>Samson, A., & Ramakrishnan, B. (2017). Scavenging Mode of Vertebrate Scavengers on Domestic Buffalos Bubalus bubalis (Linnaeus, 1785) Killed by Tiger Panthera tigris and Natural Deaths in Southern India. Journal homepage: www. wesca. net, 12(1).</ref> On the contrary, though, at times white-headed vultures and tawny eagles have been observed peaceably sharing roadkills in some instances.<ref name= Steyn/> Often tawny eagles will come to smaller carcasses of almost any animal, as will other smaller scavengers like bateleurs and hooded vultures as well as crows, perhaps merely to avoid the competition that often occurs at large carcasses.<ref name= Naoroji/><ref name= Anderson/><ref name= Mundy/><ref name= CJ/><ref name= Kendall/> One subadult tawny eagle was observed to be following a pack of African wild dogs (Lycaon pictus), almost certainly in order to scavenge off of their kills.<ref>Lewis, A.D. (1990). Tawny Eagle Aquila rapax following a foraging pack of Hunting Dogs Lycaon pictus. Scopus, 14: 19-20.</ref>

Kleptoparasitism

File:Tawny Eagle (Aquila rapax) and Young Bateleur (Terathopius ecaudatus) (6017682851).jpg

A tawny eagle perched with a young bateleur. The ecology of these species is often broadly similar but the tawny eagle usually dominates the bateleur in food competition.

The tawny eagle steals food from other raptors in addition to catching its own prey and coming to previously dead food sources.<ref name= Wichmann/><ref name= Steyn/> The Afrikaans name for the tawny eagle is a "Roofarend", meaning the "Robber Eagle".<ref name= Wichmann/> This behaviour is not entirely segregated from their scavenging on carrion behaviours but the considerable aggressiveness and boldness of the eagles in this circumstances are very different from their rather retiring disposition in scavenging contexts.<ref name= Steyn/><ref name= Steyn2/><ref>Brockmann, H. J., & Barnard, C. J. (1979). Kleptoparasitism in birds. Animal behaviour, 27, 487-514.</ref> At times the tawny eagle is considered "fearless" in their piratical attacks and is certain to engage in them more frequently than almost any other member of the booted eagle clan or perhaps even birds of prey.<ref name= Brown/><ref name= Steyn/> Other related eagles like the steppe eagle and eastern imperial eagle, as well as most sea eagles, can be locally regular kleptoparasites but tawny eagles rob prey from other birds with some regularity in every part of the range.<ref name= Naoroji/><ref name= Steyn/><ref name= Roberts/> Amongst all birds, only a few types of seabird such as skuas and frigatebirds are likely to derive a majority of their subsistence from kleptoparasitism.<ref>Olsen, K. M. (2013). Skuas and jaegers. A&C Black.</ref><ref>Corre, M. L., & Jouventin, P. (1997). Kleptoparasitism in tropical seabirds: vulnerability and avoidance responses of a host species, the Red-footed Booby. The Condor, 99(1), 162-168.</ref> Generally, tawny eagles will surprise other birds of prey with a dashing stoop and yank away the prey item in a manner of seconds; they will seldom completely land if the prey item is intercepted on the ground so they can take off with the plundered item quickly.<ref name= Steyn/><ref name= Steyn2/><ref name= Brown3/> The size of birds that the tawny eagles have been known to pirate food away from have ranged from species as small as black-winged kites (Elanus caeruleus) and common kestrels (Falco tinnunculus) to those as large as a lammergeier (Gypaetus barbatus).<ref name= Brown/> There seems to few limits to the raptorial birds that the tawny eagle will not pirate from given the opportunity.<ref name= Steyn/><ref name= Roberts/> In one case, a pair of tawny eagles descended on a secretarybird (Sagittarius serpentarius) that had killed a large puff adder and displaced both the secretarybird and an African harrier-hawk (Polyboroides typus) that had tried to enter the fray, after which the eagle pair split the adder between them.<ref name= VanSomeren>Van Someren, V. G. L. (1956). Days with birds: studies of habits of some East African species (Vol. 38). Chicago Natural History Museum.</ref> Other raptors known to be attacked for piracy in well-known and often repeated instances have included dark chanting goshawks (Melierax metabates), bateleurs, lanner falcons (Falco biarmicus) and even the imposing martial eagles (Polemaetus bellicosus) and Verreaux's eagles, the latter eagles having appeared to offer surprisingly little to no contest the tawny eagle's piracy despite their great strength and formidable talons.<ref name= Steyn/><ref name= Steyn2/><ref>Haynes, J.R. (1981). Tawny Eagle pirates food from a Dark Chanting-goshawk. Honeyguide, 50.</ref><ref>Leonardi, G. (2015). The lanner falcon. Italy: Giovanni Leonardi.</ref> Carnivorous birds that are not traditionally considered raptorial birds, such as marabou storks and southern ground hornbills (Bucorvus leadbeateri), are also occasionally kleptoparasitized by tawny eagles.<ref name= Steyn/><ref>Kemp, A. C., & Kemp, M. I. (1980). The biology of the southern ground hornbill Bucorvus leadbeateri (Vigors)(Aves: Bucerotidae). Annals of the Transvaal Museum, 32(4), 65-100.</ref><ref>Kemp, M. I., & Kemp, A. C. (1978). Bucorvus and Sagittarius: Two modes of terrestrial predation. In: Proceedings of the Symposium on African Predatory Birds, Northern Transvaal Ornithological Society, Pretoria.</ref> Interspecific piracy may be most frequent on bateleur despite that species being similarly sized and powered as the tawny eagle. While 5 displacements of tawny eagles by bateleur were reported in a study on their interactions, 26 instances of tawny eagles displacing bateleurs were described, clearly far more.<ref name= Watson2>Watson, R. T., & Watson, C. (1987). Interspecific piracy between Tawny Eagles and Bateleurs: how common is it. Gabar, 2, 9-11.</ref> Several smaller birds of prey were observed to be repeatedly robbed of their catches at a red-billed quelea colony, including queleas crippled but not killed by lanner falcons, although some of the maimed queleas were contested by jackals as well.<ref name= Steyn/><ref name= Steyn2/>

On occasion, a tawny eagle will find itself on the losing end of a kleptoparasitic interaction. Somewhat larger eagles have been seen to displace tawny eagles off of prey. These include African fish eagles (Haliaeetus vociferus), eastern imperial eagles and their cousins, steppe eagles.<ref name= Naoroji/><ref name= Steyn/><ref>Krueger, O. (1997). Population density and intra‐and interspecific competition of the African Fish Eagle Haliæeetus vocifer in Kyambura Game Reserve, southwest Uganda. Ibis, 139(1), 19-24.</ref><ref>Meyburg, B. U., & Kirwan, G. M. (2018). Eastern Imperial Eagle (Aquila heliaca). Handbook of the Birds of the World Alive (J. del Hoyo, A. Elliott, J. Sargatal, DA Christie, and E. de Juana, Editors). Lynx Edicions, Barcelona, Spain.</ref> African fish eagles and Pallas's fish eagles (Haliaeetus leucoryphus) in India both seem to take precedence over tawny eagles at shared feeding sources such as carrion sites and water bird nesting colonies.<ref name= Brown/><ref name= Naoroji/> In the Bale Mountains of Ethiopia, golden eagles appear to engage in displacement of and may dominate the much smaller tawny eagles.<ref>Clouet, M., Barrau, C. & Goar, J.L. (1999). The Golden Eagle (Aquila chrysaetos) in the Bale Mountains, Ethiopia. The Raptor Research Foundation, 33 (2): 102-109.</ref> As aforementioned, a bateleur can succeed in seldom instances in pirating tawny eagles.<ref name= Watson2/> Vultures, especially lappet-faced vultures, may assert themselves at recent tawny eagle kills and certainly can displace the eagles in some circumstances; it is likely but not confirmed that jackals may too opportunistically rob eagles as they have been recorded doing with other eagles. Large kills, which can not infrequently include prey of up to twice the eagle's own weight, are beyond the tawny eagle's ability for wing loading. Such kills are probably frequently lost to other carnivores.<ref name= Mundy/><ref name= Kendall/><ref name= Brown3/> In Ethiopia, Ethiopian wolves (Canis simensis) were seen to rob tawny eagles repeatedly of freshly-caught rodents, succeeding in 5 of 21 attempts to do so.<ref>Sillero-Zubiri, C., & Gottelli, D. (1995). Diet and feeding behavior of Ethiopian wolves (Canis simensis). Journal of Mammalogy, 76(2), 531-541.</ref> Even much smaller birds such as house crows (Corvus splendens) have been seen to successfully rob a tawny eagle of its prey.<ref>Ali, H. (2008). Behaviour and ecology of the House Crow (Corvus splendens) in Islamabad-Rawalpindi and Adjoining areas. University of Agriculture, Faisalabad, Pakistan.</ref>

Interspecific predatory relationship

File:Aquila nipalensis and two Milvus migrans, India.jpg

A tawny eagle in India flies with two black kites.

The tawny eagle occurrence in Africa and the Indian subcontinent places it in arguably two of the most competitive environments for birds of prey in the world.<ref name= Naoroji/><ref name= Steyn/> In turn, the tawny eagle seems to adapt via a lack of specialization on any particular prey type, hunting style or food source, and via including carrion in the diet quite often.<ref name= Steyn/><ref name= Steyn2/> Many other raptors and eagles overlap in habitat use with tawny eagles.<ref name= Wichmann/><ref name= Steyn/> However, wintering and residential Aquila and spotted eagles that bear some relation usually use slightly differing habitats in contrast to the tawny eagle.<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name= Smeenk/> In Tsavo East National Park, the ecology of this eagle was studied at length in contrast to bateleurs and much larger martial eagles, which can appear to have broadly similar habitat and prey preferences, as well as the slightly smaller but larger-clawed African hawk eagles, which tends to habituate to slightly more wooded dry areas.<ref name= Smeenk/> In general within this study, all four eagle species derived a majority of their prey biomass from Kirk's dik-dik but that the martial eagles tended to take slightly larger dik-diks than the bateleur and tawny eagles, took slightly more in the park per pair based on annual estimates and were more unlikely to scavenge the prey while the African hawk-eagle tends to take younger dik-diks.<ref name= Smeenk/> The diet of the tawny eagle and bateleur in Tsavo East overlapped by around 64%, whereas the diet of the tawny and martial eagles only overlapped by 29%. The tawny eagle was the only eagle here to heavily supplement their diet with alternate prey like snakes, although bateleurs also took a wide range of prey.<ref name= Smeenk/> The Tsavo East study further indicated that the predatory pressure on dik-diks is mitigated temporally by the slightly staggered nesting seasons of each eagle, with bateleurs tending to nest rather earlier, the hawk-eagle slightly later, so the peak reliance on the prey did not generally overlap.<ref name= Smeenk/> Furthermore, habitat differs, with the African hawk-eagle foraging in more wooded areas while the bateleur can forage in more open, treeless areas than tawny eagles because the bateleur is an aerial hunter while the tawny eagle typically requires perches to hunt from.<ref name= Smeenk/> Further study has indicated that, in Africa, the bateleur broadly mirrors the tawny eagle in most respects of ecology.<ref>Watson, R. T. (1986). The ecology, biology and population dynamics of the Bateleur Eagle (Terathopius ecaudatus) (Doctoral dissertation, RT Watson).</ref> One stark difference from virtually any other known eagle is the tawny eagle's nesting habits. That is that this eagle nests almost invariably on the top of the canopy of a tree, rather than a main trunk or large sturdy branch of trees (or on cliffs or, in steppe eagles, the ground).<ref name= Steyn/><ref name= Bildstein>Bildstein, K. L., & Parry-Jones, J. (2011). The eagle watchers: Observing and conserving raptors around the world. Cornell University Press.</ref> The nesting location of tawny eagles runs more parallel to those of vultures.<ref name= Mundy/> Study in Kruger National Park has shown that the tawny eagle and white-backed vulture (Gyps africanus) will freely nest in the treetop nest built by the other species.<ref name= Kemp3/> Furthermore, other species, including large owls and snake eagles, will use old nests built by tawny eagles.<ref name= Steyn/> Although the habitats used by martial and tawny eagles have been reported as broadly similar, detailed study in the Karoo found that the tawny species preferred areas with higher and more predictable summer rainfall and with higher primary productivity than the martial.<ref>Machange, R. W., Jenkins, A. R., & Navarro, R. A. (2005). Eagles as indicators of ecosystem health: Is the distribution of Martial Eagle nests in the Karoo, South Africa, influenced by variations in land-use and rangeland quality? Journal of arid environments, 63(1), 223-243.</ref>

Opportunistically, the tawny eagle may prey upon smaller birds of prey but this is fairly infrequent and the capture of raptorial birds has thus far been seldom reported. A hungry or food-gathering male tawny eagle may infrequently plunder the nests of other raptorial birds.<ref name= Smeenk/><ref name= VanSomeren/> Incautious, injured or distracted birds of prey may too be vulnerable to being killed as well.<ref name= Steyn2/><ref name= Engel>Engel, J. I. (2011). Possible predation of a Pygmy Falcon by a Tawny Eagle in Namibia. Biodiversity Observations, 34-35.</ref> Diurnal birds of prey known to be preyed upon by tawny eagle in Africa have included black-winged kites, hooded vultures, pale chanting goshawks (Melierax canorus) and African pygmy falcons (Polihierax semitorquatus).<ref name= Smeenk/><ref name= Roberts/><ref name= Tarboton/><ref name= Engel/> In India, the tawny eagle has been known to prey upon western marsh harriers (Circus aeruginosus), shikras (Accipiter badius) and white-eyed buzzards (Butastur teesa).<ref name= Ali/><ref name= Dharmakuarsinhji/> Owls are apparently fairly vulnerable to tawny eagle predation. Species that they have been known to prey on are barn owls (Tyto alba), spotted eagle-owls (Bubo africanus), little owls (Athene noctua), pearl-spotted owlets (Glaucidium perlatum) and marsh owls (Asio capensis).<ref name= Smeenk/><ref name= Engel/><ref>Van Nieuwenhuyse, D., Genot, J. & Johnson, D. (2008). The Little Owl: Conservation, Ecology and Behavior of Athene noctua. Cambridge, New York: Cambridge University Press.</ref> The fresh remains of a secretarybird were found in one tawny eagle nest in Africa but, if the eagles killed the bird rather than scavenged it, this would need confirmation.<ref name= Steyn/> The tawny eagle, despite being an eagle of intermediate size, does not seem to be subject to natural predators in adulthood as far as is known and can be said to fulfill the role of an apex predator.<ref name= Steyn/><ref name= Roberts/><ref name= Osborne/> Nestling tawny eaglets and young tawny eagles are commonly vulnerable to assorted natural predators but these are little known. A partial list of probable nest predators are likely corvids, snakes and carnivores capable of climbing.<ref name= Ferguson-Lees/><ref name= Steyn2/><ref name= Osborne/> One confirmed predator of nestling tawny eagles is the honey badger (Mellivora capensis).<ref>Begg, C. M., Begg, K. S., Du Toit, J. T., & Mills, M. G. L. (2003). Sexual and seasonal variation in the diet and foraging behaviour of a sexually dimorphic carnivore, the honey badger (Mellivora capensis). Journal of Zoology, 260(3), 301-316.</ref>

Breeding biology

Pairing and Territories

The tawny eagle often seems to pair for life. Like most birds of prey, they are quite territorial towards conspecifics.<ref name= Ferguson-Lees/><ref name= Steyn/> The commonest display is single or mutual high circling or soaring often in wide spiral. Males will sometimes dive and stoop repeatedly around the female, though she does not usually respond by turning over.<ref name= Ferguson-Lees/> Pairs may engage in the display each year to strengthen pair bonds.<ref name= Naoroji/> Occasionally two tawny eagles will interlocks talons to descend rapidly, cartwheeling down 30 m or more within a few seconds, sometimes disengaging just before the ground.<ref name= Naoroji/> In other related eagles of the Aquilinae subfamily, cartwheeling interactions are usually considered to be aggressive fights between a territorial eagle and an intruder of the same sex. Prior studies thought this to be the case for the tawny eagle, with an estimated 82% of cartwheeling instances thought to be aggressive, 11% for courtship and 7% for apparent play. However, through closer observations evidence has been made of frequent cartwheeling between males and females as a regular part of the courtship display.<ref>Murn, C., Betchley, P., & Robert, C. (2009). Talon-locking and cartwheeling as a prelude to copulation in Tawny Eagles Aquila rapax. Gabar, 20, 12-14.</ref><ref>Borello, W. D., & Borello, R. M. (2004). Two incidents of talon-grappling and cartwheeling in the Tawny Eagle Aquila rapax. Ostrich-Journal of African Ornithology, 75(4), 320-321.</ref><ref>Simmons, R. E., & Mendelsohn, J. M. (1993). A critical review of cartwheeling flights of raptors. Ostrich, 64(1), 13-24.</ref> Undulating sky dances are sometimes performed too by males with a series of descents and upward swoops on partially close wings, accompanied by calling. However, instances of this seem to be rare.<ref name= Ferguson-Lees/> In one instance, two males appeared to engage in a display for a single female.<ref name= Dharmakuarsinhji/> Per one author's opinion the aerial displays of the tawny eagle are "not particularly spectacular compared to other eagles".<ref name= Steyn/> The breeding season tends to fall in March to August in northeastern Africa, October to June in west Africa and in almost all months of the year but in central, east and southern Africa, but mainly from May to November in Kenya and April to January in central and southern Africa.<ref name= Ferguson-Lees/><ref name= Vincent>Vincent, A. W. (1945). On the breeding habits of some African birds. Ibis, 87(3), 345-365.</ref> In India, the breeding season is usually November to May, but occasionally can vary from any time from October to August.<ref name= Ferguson-Lees/><ref name= Naoroji/> Mating generally occurs in and around the nest vicinity.<ref name= Naoroji/> The density very variable on the African continent overall of breeding pair which were estimated to occupy about Template:Convert each.<ref name= Ferguson-Lees/> Zimbabwe nest spacing was found to be Template:Convert in one study. On the border of Kruger National Park, 7 pairs found in a Template:Convert area but in regular spaced pylon nests in western Transvaal, nests were Template:Convert apart.<ref name= Steyn/> In Hwange National Park, over 11 years of study, 92 pairs on were found to be nesting over basalt in a Template:Convert area while 84 pairs on Kalahari sands in a Template:Convert area. Mean nest distances on basalt were around Template:Convert while on sands it was around Template:Convert.<ref name= Hustler/> In Zambia, the nesting density was considered high for the species at a pair per Template:Convert.<ref name= Osborne/>

Nest

The nests of the tawny eagle are large platforms, composed of sticks but sometimes incorporating animal bones.<ref name= Ferguson-Lees/> Nesting sites tend to be open to the sky, in flat, open or hilly country, and offer a commanding, good view of the surrounding country. The sites are not infrequently close to watering holes and, more so in India, close to villages.<ref name= Ferguson-Lees/><ref name= Naoroji/><ref name= Steyn/> Nests are usually Template:Convert above the ground, though seldom can be up to Template:Convert high.<ref name= Ferguson-Lees/> Nests are located at the top crown of the tree and only very rarely are placed beneath the canopy or on a lateral branch.<ref name= Ferguson-Lees/><ref name= Steyn/> In Kenya, tawny eagles showed no nesting preference according to tree height or spatial distribution of trees;<ref name= Kendall/> however, they preferred Euphorbia, Boscia and Euclea tree species.<ref name= Kendall/> In India, commonly used trees used in the northern areas are Ficus religiosa, Dalbergia sissoo and mango trees while in the arid Kutch and western Rajasthan areas they often nest in rather stunted Vachellia nilotica and Prosopis chilensis (i.e. usually the nests here are Template:Convert high but are sometimes down to Template:Convert ).<ref name= Naoroji/><ref>Anderson A. (1871). Notes on the raptorial birds of India. Journal of Zoology, 44: 675-690.</ref> Trees are usually selected that have prickly branches, presumably for protection.<ref name= Steyn/> Despite their prominent position in the trees, the nests can be surprisingly hard to perceive peering from the ground level.<ref name= Steyn/><ref name= Vincent/> In Kgalagadi Transfrontier Park, South Africa, tawny eagles build nests that are positioned in the canopy of large Vachellia erioloba trees.<ref name= Herholdt/> These Kgalagadi pairs tend to be the largest and tallest trees, averaging at Template:Convert.<ref name= Herholdt/> Tawny eagles in India reportedly often nest in a tree over successive years, but the species is threatened by the lopping and cutting of all remaining suitable trees for fuel and fodder.<ref name= Naoroji/> Tawny eagles in Kgalagadi Transfrontier Park, on the other hand, build new nests yearly and only 2% of nests are reused for breeding purposes the following year.<ref name= Herholdt/> Potential risk of collapse and growth of branches around the nest are thought to be factors limiting nest reuses in Africa. Usually new nests are not more than Template:Convert away from the prior nest.<ref name= Steyn/> Both sexes participate in nest building and the repair of a nest takes up to 4 to 7 weeks, though most of the construction can be completed within about a week.<ref name= Steyn/> For an eagle, their nests are relatively wide, flat and shallow.<ref name= Steyn/> Nests may measure just under Template:Convert in diameter and Template:Convert deep but can easily reach over Template:Convert and Template:Convert with repeated uses.<ref name= Ferguson-Lees/><ref name= Steyn2/> Nests are usually lined with grass, leaves, seedpods and fur as well as odd objects such as newspapers, paper packets and polythene bags.<ref name= Ferguson-Lees/><ref name= Steyn/> In Kruger National Park, tawny eagles have been recorded using nests of other species of raptor such as white-backed vulture and white-headed vulture.<ref name= Steyn2/> At times, tawny eagles have been known to nest on top of the large communal nests of the white-headed buffalo weaver (Dinemellia dinemelli).<ref name= Steyn/> In the Central Karoo region of South Africa, tawny eagles build their nests in large electric transmission towers.<ref name= Jenkins/> Populations of large eagles like the martial eagle and Verreaux's eagle have been recorded breeding on these power pylons since the 1970s.<ref name= Jenkins/> Between 2002 and 2003, 39% of electrical faults recorded on transmission lines were due to large eagle nests.<ref name= Jenkins/> As a result, problem nests were dismantled and rebuilt below the electrical conductors.<ref name= Jenkins/>

Eggs

File:Aquila rapax MHNT.ZOO.2010.11.91.1.jpg

Aquila rapax – MHNT

File:Aquila rapax belisarius MHNT.ZOO.2010.11.91.2.jpg

Aquila rapax belisarius – MHNT

Eggs are laid at intervals of several days, mainly timed to the dry season but at times also in the wet season.<ref name= Ferguson-Lees/> Evidence from the Kalahari Desert shows that egg-laying is timed to exploit a number of food resources with warmer weather in sync with young in the nest, such as various small mammals and the springbok (Antidorcas marsupialis) lambing season.<ref>Maclean, G. L. (1969). The breeding seasons of birds in the south-western Kalahari. Ostrich, 40(1): 179-192.</ref> In India, intervals were more prolonged when the habitat was less optimal.<ref name= Naoroji/> Of 26 tawny eagle nests monitored between 1988 and 1996 in the Kgalagadi Transfrontier Park, 84.6% of the laying dates occurred between May and June.<ref name= Herholdt/> These laying dates are similar to populations in Zambia, Zimbabwe<ref name= Osborne/><ref name= Steyn/> and the Maasai Mara region in Kenya as well as elsewhere in southern and southeastern Africa.<ref name= Smith/><ref>Tarboton, W. R., Kemp, M. I., & Kemp, A. C. (1987). Birds of the Transvaal. Transvaal Museum.</ref><ref name= Kendall2>Template:Cite journal</ref> Cases of eggs being laid in southern Africa in July and August may be cases of replacement clutches.<ref name= Steyn/> Incidental data on laying phenology from north and West Africa shows the tawny eagles of the area lay eggs usually in the earlier part of the year, i.e. January to April; in Ghana egg-laying may range from October to February, though largely is in December–January; November–February egg-laying occurred in Ethiopia and apparently around April in Morocco.<ref name= Ash/><ref name= Cheke/><ref name= Thevenot/><ref>de Balsac, H. H., & Mayaud, N. (1962). Les oiseaux du nord-ouest de l'Afrique: distribution géographique, écologie, migrations, reproduction (Vol. 10). P. Lechevalier.</ref><ref>Bannerman, D. A. (1951) The birds of tropical West Africa. Crown Agents, London.</ref><ref>Bannerman, D. A. (1953). The Birds of West and Equatorial Africa. Edinburgh & London.</ref> Showing the variation in India, in Punjab, Uttar Pradesh and Bihar, tawny eagles are mainly laying eggs in January while in Kutch and Jaisalmer, the young are already leaving the nests.<ref name= Naoroji/> Clutch sizes range from 1 to 3 eggs per nest, but average 1.7 eggs per clutch.<ref name= Steyn/><ref name= Osborne/> In drier years in Hwange National Park, clutch sizes appear to become reduced.<ref name= Hustler/> The eggs are white but variously and usually faintly marked with brown, varying from unmarked sometimes to quite well-marked with spots and blotches of reddish brown.<ref name= Brown/> In 67 eggs of the nominate subspecies, the eggs were Template:Convert in height by Template:Convert in diameter, with an average of Template:Convert in the sample while another 30 from the same race averaged Template:Convert. In A. r. vindhiana, 80 eggs measured from Template:Convert by Template:Convert, with an average of Template:Convert.<ref name= Brown/><ref name= Steyn/>

Development of young and parental behaviour

Eggs are incubated by the female for 40–44 days, with extreme records of 30 to 45 days, before hatching.<ref name= Ferguson-Lees/> Incubation tends to begin with the first egg and may be done exclusively by the female in India but, in African data, the male sometimes briefly relieves her.<ref name= Ferguson-Lees/><ref name= Naoroji/> When the nests are climbed up to, the females are tight sitters, often flying at the very last minute.<ref name= Naoroji/> Green lining may still be added at the incubation stage.<ref name= Steyn/> Upon hatching, the young apparently have to be constantly brooded or shaded from strong sun in the very open nests.<ref name= Ferguson-Lees/><ref name= Brown/> The chicks are initially covered with white down, with a black bill, yellow cere and feet and brown eyes; a thicker white coat is acquired at 2 weeks and 1 week later the 1st feathers appear on scapulars and wing coverts.<ref name= Steyn/> The young eaglets can stand weakly at about 3 weeks, walk around the nest at 4 weeks and start to wing-flap about a week later.<ref name= Steyn/><ref name= Steyn2/> Wing and tail quills sprout rapidly, with feathers appearing down the side of the breast at 4 weeks. By 5 weeks, feathers cover much of the body except for the head and underparts. By week 7, the chick has only a small amount of down feathers remaining and weighs around Template:Convert.<ref name= Steyn/><ref name= Steyn2/> The rapid development of dorsal feathers is comparable to other raptors that use open nest sites such as snake eagles and secretarybirds.<ref name= Steyn/> Only one chick usually survives after hatching.<ref name= Steyn/><ref name= Roberts/> This is often due to siblicide, where the older chick fatally wounds the younger chick in the first few days of life. The older sibling weighs around Template:Convert while the younger one weighs about Template:Convert frequently at the point of its demise or disappearance.<ref name= Steyn/> In southern Africa, there are at least four cases of two fledglings occurring in one nest, however.<ref name= Steyn/><ref>McGowan, G. (1992). Tawny Eagle pair raise two chicks. Babbler, 23: 50.</ref> At around 5 weeks, the eaglet adopts anti-predator behaviours, some laying prone while a novel animal approaches while others adopt a truculent threat posture with feathers raised, gape opened, wings poised to slap and talons barred for slashing.<ref name= Steyn/> For the first 10 days, the adult female observes the chick very closely, relying on food provisioned by the male. After two weeks, the chick is left alone for 2.5 hours each day, whilst the adults forage. At this point the male may start making direct food deliveries to the eaglet. This is considered a relatively early point to stop attending to the nest for an eagle this size and the nest often soon becomes unsuitably foul with remains.<ref name= Steyn/><ref name= Steyn2/> One 39-day-old eaglet was able to tear up its own food already but was still primarily fed by the female.<ref name= Steyn/> The first flight attempts are around 7–10 weeks but the chick is fully grown and capable of fledging the nest fully after 10–12 weeks.<ref name= Ferguson-Lees/><ref name= Steyn2/> However, the female may remain to shelter during rainstorms around to as late as the fledging stage.<ref name= Ferguson-Lees/> The full stage of dependence is ongoing for about 6 weeks after fledging. The young tawny eagle may stay with the parents even until next breeding season.<ref name= Ferguson-Lees/> In India at least, after the nesting period, the pairs disperse and leave the nesting area, seldom being seen near the nest until pairing off again initiates in October.<ref name= Naoroji/> A juvenile tawny eagle that was shot at 2 years old was Template:Convert away from its original nest while 2 juveniles at 5 months and 7 months old were Template:Convert away, respectively.<ref name= Steyn/>

Nesting success and failures

Nest losses of eggs and young appears to be quite high.<ref name= Ferguson-Lees/> Young eaglets often die, at times by their siblings, and if poorly guarded nests are often predated by a probably wide range of predators.<ref name= Ferguson-Lees/> Nesting success is driven by quality of habitats and food access.<ref name= Naoroji/> Breeding efforts in Zimbabwe produced 19 young in 26 pair years with a replacement rate of 0.73 young per pair per year.<ref name= Steyn2/> In India, tawny eagles pairs seem to adapt to suboptimal overly sandy habitats by more dispersing nests, and can show similar productivity of chicks per nest as a result.<ref name= Naoroji/> In Hwange National Park, 72.4% of pairs present were thought to breed on average in the course of a year, with an average of 0.61 fledglings produced per effort.<ref name= Hustler3>Hustler, K., & Howells, W. W. (1986). A population study of tawny eagles in the Hwange National park, Zimbabwe. Ostrich, 57(2), 101-106.</ref> This is and other studies support that rainfall is key to productive success in tawny eagles of this area, with far more two egg clutches rather than one egg ones (which usually failed) and less confined breeding periods in years that had greater rainfall.<ref name= Steyn2/><ref name= Hustler3/> Breeding success, recorded as young per pair per year (ypy), was lower still in Namibia and Tsavo East National Park than in Zimbabwe (0.4, 0.5 and 0.78 ypy respectively).<ref name= Steyn2/><ref name= Osborne/> Higher nesting success was found in Zambia, where pair produced a mean of 1 fledgling per nest.<ref name= Osborne/> Although an extensive study of lifespan are not known to have taken place for the tawny eagle, it is known that these eagles can live up to at least 16 years of age in the wild.<ref name= Wichmann/><ref>Flower, S. S. (1923). List of Birds of Prey 1898–1923: With Notes on Their Longevity. Government Press.</ref>

File:Aquila rapax.jpg

An unusually light juvenile tawny eagle from India.

Status

Conservation

The tawny eagle still occupies a large range.<ref name="iucn status 12 November 2021" /> In Africa, it has been estimated that the range of the species covers about 15 million square kilometers, in addition to a range of about 3.1 million square kilometers in Asia.<ref name= Ferguson-Lees/> As recently as the 1990s, the global population was thought to possibly range into six figures with a population in Asia at that time thought to be in the hundreds of thousands alone.<ref name= Ferguson-Lees/> However, the species is currently listed as Vulnerable on the IUCN list of Threatened species.<ref name="iucn status 12 November 2021" /> The current population is far less than half of what it was once thought to be, with only about 100,000 to just under 500,000 individuals thought to persist worldwide.<ref name="iucn status 12 November 2021" /> There was a clear decrease in tawny eagle sightings between SABAP and SABAP2 in Southern Africa, occurring in only 323 of 1440 quarter degree grid cells.<ref name= Underhill>Template:Cite journal</ref> During close study of the tawny and martial eagle in central Namibia, a precipitous decline was detected in both, with a tawny eagle population that was once regionally numbered about 19 pairs down to 2 known pairs.<ref name= CJ2>Brown, C. J. (1991). Declining Martial Polemaetus bellicosus and Tawny Aquila rapax Eagle populations and causes of mortality on farmlands in central Namibia. Biological conservation, 56(1), 49-62.</ref> The once seemingly innumerous population of this species within Kgalagadi Transfrontier Park was known by the 1990s to be down to merely 40 known pairs.<ref>Herholdt, J. J. (1998). Survival threats and conservation management of raptors in the Kgalagadi Transfrontier Park. Transactions of the Royal Society of South Africa, 53(2), 201-218.</ref> Roadside counts conducted in Mali, Niger and Burkina Faso show that although the majority of raptor species are in drastic population decline, only the tawny eagle and snake eagles are surviving outside of protected areas.<ref name= Thiollay3>Template:Cite journal</ref> In India, the tawny eagle was once considered "our commonest eagle"<ref name= AliS/> but strong declines have been detected with surveys indicating strongholds like Rajasthan have shown reductions of observed pairs by up to half.<ref name= Naoroji/> According to the producer-scrounger foraging theory, vultures are to some extent reliant on tawny eagles to help locate carcasses.<ref name= Ogada/> Thus, the conservation of eagles outside protected areas is of vital importance to ensure the survival of vultures.<ref name= Ogada/>

Threats

File:Serengeti Raubadler.jpg

A tawny eagle in the Serengeti.

Tawny eagles face a number of threats that affect their breeding behaviour, foraging success and ultimately the survival of individual birds. The most recent and devastating threat to survival occurred on 20 June 2019. The carcasses of 468 white-backed vultures, 17 white-headed vultures, 28 hooded vultures, 14 lappet-faced vultures and 10 cape vultures were found alongside 2 tawny eagles. A total of 537 vultures and 2 eagles were found poisoned in northern Botswana. It is suspected that they died after eating the carcasses of 3 elephants that were laced with poison by poachers. Carcasses are poisoned to ensure that scavengers are unable to aid rangers in the effort to locate poached wildlife. By circling above dead animals, large raptors act as an early detection system for anti-poaching rangers.<ref name="NDTV AFP 06-2019">Template:Cite news</ref><ref name="CNN 06-2019">Template:Cite news</ref><ref name="Smithsonian 06-2019">Template:Cite magazine</ref><ref name="Afrik21 06-2019">Template:Cite news</ref> Poisoning events are far from restricted to Botswana and are thought to be a direct factor in the reduction of tawny eagles as well even in the protected areas of Kruger National Park.<ref>Loftie-Eaton, M. (2014). Geographic range dynamics of South Africa's bird species. South Africa: Department of Biological Sciences, University of Cape Town.</ref> In central Namibia, all 5 of the juvenile tawny eagles that were radio-tagged were poisoned by strychnine baits, completely decimating all recruitment of the species in the area.<ref name= CJ2/> Mysteriously, the populations of bateleurs and tawny eagles in the Maasai Mara appear to be bumping up as opposed to the declines reported elsewhere, seemingly in sync with the worsening declines of vultures on the Maasai.<ref>Virani, M. Z., Kendall, C., Njoroge, P., & Thomsett, S. (2011). Major declines in the abundance of vultures and other scavenging raptors in and around the Masai Mara ecosystem, Kenya. Biological Conservation, 144(2), 746-752.</ref>

Further threats to tawny eagles include habitat loss and land-use changes such as intensified cattle grazing, firewood collection and sale and the charcoal industry.<ref name= Wichmann/> Such culling of the spare trees of arid India seem to be the primary driver of less understand decline of tawny eagles in India.<ref name= Naoroji/> A seemingly higher instance of bacterial infections also seems to affecting the tawny eagles of India.<ref>Dutta, T. K., Sudhan, N. A., & Azmi, S. (2008). PCR detection of Pasteurella multocida in eagle PCR based detection, characterization and antibiogram of Pasteurella multocida from wild eagle (Aquila rapax) in Jammu and Kashmir. The Indian Journal of Animal Sciences, 78(12).</ref> Raptor populations are reliant on seasonal rainfall events which influence the survival of prey populations.<ref name= Wichmann/> Climate change is alternating rainfall patterns in the arid regions of Southern Africa and impacting on prey populations. There is a clear correlation between rainfall events and breeding success of tawny eagles. It was found that the projected decline of tawny eagles from climate change, which is already underway, begun via impact population persistence, first effecting population dynamics, the composition of biological communities and finally biodiversity.<ref name= Wichmann/><ref name= Wichmann3>Wichmann, M. C., Jeltsch, F., Dean, W. R. J., Moloney, K. A., & Wissel, C. (2003). Implication of climate change for the persistence of raptors in arid savanna. Oikos, 102(1), 186-202.</ref> Electrocutions and collision risks associated with overhead power lines remain a constant threat to large eagles and vultures. Furthermore, the powerline nesting raptors were found to be a significant source of line faulting in the area, causing substantial financial issues.<ref name= Jenkins>Template:Cite journal</ref><ref>Van Rooyen, C. S. (2000). Raptor mortality on power lines in South Africa. In Raptors at risk: proceedings of the 5th world conference on birds of prey and owls. Hancock House Publishers and the World Working Group on Birds of Prey and Owls, Blaine, Washington (pp. 739-750).</ref> Occasionally, tawny eagles are also killed by flying into various manmade objects such as reservoirs or are killed by automobiles and are at risk at wind turbines in India.<ref name= CJ2/><ref>Anderson, M. D., Maritz, A. W., & Oosthuysen, E. (1999). Raptors drowning in farm reservoirs in South Africa. Ostrich, 70(2), 139-144.</ref><ref>Pande, S., Padhye, A., Deshpande, P., Ponkshe, A., Pandit, P., Pawashe, A., Pednekar , S., Pandit, R. & Deshpande, P. (2013). CEPF Western Ghats Special Series: Avian collision threat assessment at Bhambarwadi Wind Farm Plateau in northern Western Ghats, India. Journal of Threatened Taxa, 5(1), 3504–3515.</ref> The overarching threat to any raptor population is human population increase which causes competition for habitat and food resources.<ref name= Thiollay/> Key to conservation of the tawny eagle population is mitigating the effects of global warming.<ref name= Wichmann/><ref>Crick, H. Q., & Sparks, T. H. (1999). Climate change related to egg-laying trends. Nature, 399(6735), 423-423.</ref><ref>Aumann, T. (2001). The structure of raptor assemblages in riparian environments in the south-west of the Northern Territory, Australia. Emu, 101(4), 293-304.</ref><ref>Post, E., Peterson, R. O., Stenseth, N. C. & McLaren, B. E. (1999). Ecosystem consequences of wolf behavioural responses to climate. Nature, 401: 905–907.</ref><ref>Moss, R., Oswald, J., & Baines, D. (2001). Climate change and breeding success: decline of the capercaillie in Scotland. Journal of Animal Ecology, 47-61.</ref> Also, clearly, the banning of poison baits and the mitigation of dangerous powerlines in eagle-utilized areas is key for the survival of the tawny eagles.<ref name= CJ2/><ref name="CNN 06-2019"/><ref name= Jenkins/>

External links

• Tawny eagle – Species text in The Atlas of Southern African Birds
• Tawny eagle at Animal Diversity Web