Steppe eagle
The steppe eagle is a large bird of prey. Like all eagles, it belongs to the family Accipitridae. The steppe eagle's well-feathered legs illustrate it to be a member of the subfamily Aquilinae, also known as the "Booted eagles". This species was once considered to be closely related to the sedentary tawny eagle and the two forms have previously been treated as conspecific. They were split based on pronounced differences in morphology and anatomy; 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.
The Steppe eagle is in many ways a peculiar species of eagle. It is a specialized predator of ground squirrels on the breeding ground, also taking other rather small mammals and other prey, doing so more often when ground squirrels are less consistently found. In rather treeless areas of the steppe habitats, these eagles tend to nest on a slight rise, often on or near an outcrop, but may even be found on flat, wide-open ground, in a rather flat nest. They are the only eagle to nest primarily on the ground. Usually, one to three eggs are laid and, in successful nests, one to two young eagles fledge. The steppe eagle undertakes a massive migration from essentially its entire breeding range, moving en masse past major migration flyways, especially those of the Middle East, Red Sea and the Himalayas. In winter, though less closely studied than during breeding, the steppe eagle is remarkable for its sluggish and almost passive feeding ecology, focusing on insect swarms, landfills, carrion and the semi-altricial young of assorted animals, lacking the bold and predatory demeanor of their cousin species. Although still seen by the thousands at migration sites in larger numbers than other migrating eagles of these areas, the steppe eagle's entire population has declined precipitously. The threats to this species consist of increasing steppe fires and pests around the nests which can cause a large volume of nest failures. Rivaling these factors, declines are being exacerbated by disturbance and persecution by humans, as well as trampling of nests by livestock. Free-flying steppe eagles are also being killed in alarmingly large numbers, especially in the stronghold nation for breeding of Kazakhstan, by electrocutions on dangerous electricial wires and pylons. Due to these and other reasons, the decline of the species is thought to be considerably more than 50%. Therefore, the species is considered to be endangered by the IUCN. The steppe eagle appears on the flag of Kazakhstan and is the national bird of both Kazakhstan and Egypt.
Taxonomy
British naturalist Brian Houghton Hodgson described the steppe eagle in 1833. Aquila is Latin for "eagle" while nipalensis means "from Nepal" based on the location where the type specimen was collected presumably while migrating. Samuel G. Gmelin however, described a species of eagle from Tanais where they were found sitting on ancient mounds or graves of nomads. He called it "Aquila mogilnik," the species name "mogilnik" meaning "burial" in Russian. This was included in the 13th edition of Systema Naturae by his cousin J.F. Gmelin in 1788. The identity of this bird was, however, confused with A. heliaca but research in 2019 suggests that A. mogilnik can reliably be identified and that it would have been a valid senior name for A. nipalensis had it not been declared as a nomen dubium, a doubtful name by Ernst Hartert in 1914. The lack of usage of the name in literature now makes the name nomen oblitum, a valid senior name lost by disuse. The steppe eagle is a member of the booted eagle subfamily within the Accipitridae family. The booted eagle clan are monophyletic and the study of karyotypes has indicated that they likely have few to no close external relations within the overall extant accipitrid family. The booted eagle subfamily all have feathers covering their legs and may be found to some extent on every continent that contains accipitrids. The genus Aquila traditionally comprises large and fairly dark eagles that tend to live in open habitats. However, a significant division was determined to exist between superficially similar eagles such as the golden eagle and its three extant and similar-looking close cousins, as well as three very different smaller and pale-bellied eagles and the species complex which contains the steppe eagle. There is a similar genetic disparity with the golden eagle for the steppe eagle as with the spotted eagles which have been deemed distinct enough to form a separate genus, Clanga. The steppe eagle genetically clusters closely to the tawny eagle as well as, albeit more distantly, with the eastern and Spanish imperial eagles. However, the loci evidenced in the Aquila genera have been found to be relatively homogenous, with general studies of isoenzymes showing their genes as about ten times less distanced than certain owl genera.The steppe eagle has historically been considered conspecific with the tawny eagle, even until as recently as 1991. The latter species resides year-round in the African and Asian areas, often used seasonally as wintering grounds by the steppe eagle. The species were ultimately separated on the grounds of the differences in morphology, disparate coloring, distinct life histories, and behaviours. Testing of genetic materials has reinforced the species distinction of the steppe and tawny eagles. Genetically, the steppe eagle is thought to be basal to related species such as the tawny and imperial eagles. A fossil species, Aquila nipaloides, has been found in Italy, Corsica, Sardinia and France and was hypothesized to most closely related to the steppe eagle based on osteology of the ramus. Despite being even more strongly distinctive from the steppe eagle than the tawny eagle, the eastern imperial eagle has been seen to hybridize with the steppe eagles in the wild, once in Turkey and at least three times in Kazakhstan. Each hybrid with imperial eagles has been known to involve pairs of subadult or juvenile eagles and all known hybrid pairings were between male steppe eagles mated to female imperial eagles. Some of these hybrid pairs also produced seemingly healthy young with roughly intermediate characteristics.
The Steppe eagle has been generally considered to contain two subspecies. One was the nominate subspecies, A. n. nipalensis, which breeds in the eastern portions of the range while the western breeding population, found in most of Kazakhstan and European Russia, was considered as the subspecies, A. n. orientalis. The separation of the two subspecies was largely based on size, with the eastern population being larger and much heavier than the western eagles. The more eastern birds tend to be a shade darker and a have a more extensive nape patch, as well as having a more conspicuously deep gape-line. However, both western and Russian researchers have since made a convincing advocacy that the steppe eagle is actually a monotypical species. It was found that both previously claimed subspecies appear to broadly overlap in the breeding range and become indistinguishable at the Kazakh-Russian meeting point. The primary differences, i.e. in size and mildly in color, can be explained as clinal variations due to the environment. The breeding populations of the eastern and western eagles are insufficiently allopatric and too extensively engage in introgression to be properly regarded as full subspecies. Erroneously, a checklist once included the former subspecies of A. n. orientalis as being part of the subspecies of tawny eagle from Asia, A. r. vindhiana, an error that was later corrected.
Description
The steppe eagle is a large, bulky and robust-looking eagle. It is mainly dark brown in color with a longish but very thick neck and a relatively small head that nonetheless features a strong bill and long gape-line. It appears long-winged and has a longish and rather rounded tail and markedly well-feathered legs. Steppe eagles tend to perch somewhat upright and usually do so in the open, often utilizing isolated trees, posts, rocks or other suitable low lookouts such as mounds or straw-piles. The species often is seen on the ground where may stand for long periods of the day and walk with horizontal posture and with wingtips just exceed the tail-tip. Steppe eagles, like tawny eagles, can be relatively tame and approachable, at least compared to many of the other Aquila eagles. The adult is a somewhat variable brown with darker centers to the greater coverts. More pronouncedly in the eastern part of the range, adults have normally prominent pale rufous to dull orange-yellow to yellow-brown patches on the nape and hindcrown. Any other paler areas are obscured on perched adults. The massive gape-line runs to level with the rear of eye and is longer than in any other Aquila eagles including tawny eagles. Combined with their deep-set eyes, it lends steppe eagles an altogether rather fierce facial expression. Steppe eagle juveniles are almost invariably paler than adults, with some ranging overall from umber-brown to tawny-buff but then some are darker and more deeply brown. Juveniles tend to be brown to grey-brown on the upperparts but for generally rufous-buff nape patch. The juveniles bear conspicuously and broadly white-tipped black about the greater coverts, wings and tail and a bold but narrow cream band on the brown medians. The juvenile steppe eagle's white uppertail coverts is generally concealed when perched; the underparts are usually the same as the upperparts but may be somewhat paler tawny-buff hue. Upon their 2nd year, the plumage is still much as the 1st year appearance but show the pale tips to secondaries, median coverts and tail as often well-worn and narrower; by the start of 2nd winter the, tips of retained juvenile flight-feathers and coverts are heavily abraded and very thin. By the end of 2nd winter, often the immatures look very worn and have nearly lost pale tips altogether and from 3rd year onward manifest a variable mix of old and new feathers. Generally, immatures are often rather scruffy in appearance until adult-like plumage attained at year five, after which the feathers generally appear more compact. Adults have brown to hazel eyes, while juveniles have distinctly dark brown eyes; the cere and feet are yellow at all ages.In flight, the steppe eagle appears as a large, impressive and visibly heavy raptor with a well-projecting large head and bill and rather broad neck and long, broad wings. They evidence proportionately long arms, especially in the larger eastern birds. The wings tend to be held almost parallel-edged and square-ended with 7 very elongated emarginations. Often juveniles can tend to appear somewhat narrower winged. The broad body of the species often looks suspended underneath and the tail appears rounded or even wedge-shaped, measuring about 3/4 of the length of wing-base. The wingspan is about 2.6 times greater than the total body length. On the upperwings, steppe eagles show a pale greyish primary patch that is often quite large and obvious, often being pale at the base on the greater primary coverts but on adults much less marked. On the underwing, a very small carpal crescent may be present but can vary from invisible to slightly more marked. The flight feathers are greyish and all have 7–8 well-spaced blackish bars, while the fingers are plain blackish. Adults are basically all fairly uniform dark brown. Adults may evidence in flight some whitish patches on back and tail coverts that are varying from insignificant to fairly prominent. Adult eagles that do show a dark-barred greyish primary patch usually have that confined to a wedge-shape on inner primaries though can sometimes be rather more prominent. Below adults show dark-barred grey flight feathers and tail with the broad blackish trail edges and wing ends being rather distinctive; the wing linings are often slightly paler to darker than remiges and often with an obscure remnant of broken paler central band. Juveniles are quite distinctive in flight if seen in reasonable view. Above, juveniles are pale greyish-brown to yellow-brown about the body and forewing-coverts, have a broad whitish U above the tail. They possess broad white tips to the blackish greater coverts, flights feathers and tail creating obvious whitish bars on the wings and trailing edges as well as a large and prominent whitish patch covering much of the inner primaries. On its underside, the juvenile is mid-brown to brownish-yellow with a paler throat and creamy crissum. Below, the creamy central wing band is even broader than above, while the greater coverts all white with some dark centres on primaries. Despite reports that some juvenile 1st years have subtle or no central wing bands, these are believed to be cases where these feathers exist but are obscured by long median coverts. At the end of the first year, the young steppe eagle tends to have pale tips to wings, tail and upperwing coverts become rather abraded; thereafter the development young evidence much variation due to individual differences. Usually, by the end of 2nd winter, the wing looks even more worn and uneven in pattern, with any newly acquired narrowly white-tipped quills clearly longer than old worn juvenile ones that have lost their pale tips. From the 3rd winter on, the pale parts clearly reduced, flight feathers and tail often appear quite ragged and by the 4th year start to more resemble adults. From the end of the 3rd year to when they obtain adult plumage, the eagles tend to have adult-like broad blackish trailing edges and tail often coupled with dark-barred grey base to black fingers and traces of the pale band along greater underwing-coverts. Maturity is obtained between the 4th and 5th years, not at 6–7 years as previously reported despite some presumed five-year-old eagles still have flecks of pale on the wing coverts and the throat and more subtle nape patches than they will ultimately manifest.