Taxonomy of the tawny owl

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An individual probably of the western subspecies S. a. sylvatica Tawnyowl03.jpg
An individual probably of the western subspecies S. a. sylvatica

The species was first described by Linnaeus in his Systema naturae in 1758 under its current scientific name. [1] The binomial derives from Greek strix "owl" and Italian allocco, "tawny owl" (from Latin ulucus "screech-owl"). [2] Some early descriptions upon review were found to have somehow conflated the very different barn owl with the scientific name Strix aluco, which in turn engendered some confusion. [3]

An individual from France perched upon a human hand TawnyOwlBeingPointedAt.png
An individual from France perched upon a human hand

The tawny owl is a member of the wood-owl genus Strix , part of the typical owl family Strigidae, which contains all species of owl other than the barn owls. Conservatively, about 18 species are currently represented in this Strix genus, typically being medium to large sized owls, characteristically round-headed and lacking ear tufts, which acclimate to living in forested parts of various climatic zones. [4] [5] Four owls native to the neotropics are sometimes additionally included with the Strix genus though some authors include these in a separate but related genus, Ciccaba . [6] [7] Strix owls have an extensive fossil record and have long been widely distributed. [8] The genetic relationship of true owls is somewhat muddled and different genetic testings has variously indicated that Strix owls are related to disparate appearing genera like Pulsatrix , Bubo and Asio . [9] [10] [7] [11] Tropical species, such as the mottled owl (Strix virgata) and the African wood owl (Strix woodfordii), the latter once considered a close relative to the tawny owl, morphological differ from and have smaller outer ear areas than tawny owls. [12] [13]

The tawny owl is thought to be a close relative of the Ural owl. Authors have hypothesized that the origin of the two species divide followed Pleistocene continental glaciations segregated a southwest or southern group in temperate forest (i.e. the tawny) from an eastern one inhabiting cold, boreal ranges (i.e. the Ural). After retreat of the continental ice masses, the ranges more recently penetrated each other. [12] [14] [15] While the life history details of the tawny and Ural owls are largely corresponding, nonetheless the species have a number of morphological differences and are largely adapted to different climates, times of activity and habitats. [12] [16] Based on Strix fossil species from Middle Pleistocene (given the name Strix intermedia) in variously the Czech Republic, Austria and Hungary show from leg and wing bones indicate an owl of intermediate form and size between Ural and tawny owls. [17] [18] [19] [20] However, fossils of a larger and differently proportioned Strix owl than a tawny owl, identified as Strix brevis, from Germany and Hungary from before the Pleistocene (i.e. Piacenzian) suggest a more complicated evolutionary and distributional history. [12] [18] [19] [20] [21] A hybrid was recorded in captivity between a male Ural and a female tawny owl, which managed to produce two offspring that were intermediate in size and had a more complex song that was also shared some characteristics with both species’ vocalizations. [22]

A number of owls that were considered conspecific with the tawny owl are now widely considered to be separate species via conclusive genetic studies. These consist of the desert or Hume's owl and its sister species, the recently separated and range-restricted Omani owl (Strix butleri). [23] [24] [25] Another species even more recently found to be distinct from tawny owls is the little-known Himalayan owl (Strix nivicolum). [26] [27] In all three separated species there is no evidence that the tawny owl breeds in the same areas as them, making each species allopatric, though the desert and tawny's range nearly abuts in some parts of the Middle East such as northern Israel. [24] [28] Also, in the Western Himalayas both the tawny and Himalayan owls are known to occur but there is likely a gap of several hundred kilometers in distribution with tawny mostly restricted to the Pakistani side while the Himalayan is rarely found west of Himachal Pradesh. [4] [29] [30] Furthermore, the desert and Omani species pair and the Himalayan species are considerably different based on superficial appearance (far more so than true tawny owl subspecies), have distinct voices and appear to have slightly different nesting habits than tawny owls. [31]

Subspecies

The tawny owl subspecies are often poorly differentiated, and may be at a flexible stage of subspecies formation with features related to the ambient temperature, the color tone of the local habitat, and the size of available prey. Consequently, various authors have historically described between 10 and 15 subspecies. [12] The total number of subspecies was once considered as totaling at 11 subspecies but is now reduced due to the separation of the Himalayan owl and its own further two subspecies down to about eight subspecies. [9] [10] [31] The currently recognised subspecies are listed below. [32]

SubspeciesRangeDescribed by (parentheses indicate originally in a different genus)Description
S. a. aluco North & Central Europe from southern Scandinavia to the Mediterranean and Black Sea and European Russia Linnaeus, 1758Markedly polymorphic with all three morphs known. Some brown morph individuals bear indistinct concentric lines on the facial disc and tend to have a disc rim is dark brown. [9] Generally, birds of the nominate subspecies are rather pale below with sparser markings and more creamy base colour showing than other European tawny owls. [33] Study of genetic phylogeography of showed that the population of the nominate race in the Balkans originated as a postglacial occupation of northern territories, although these populations do interbreed with populations of two other clines, in the Alps and Pyrenees. [34] This is a medium-sized subspecies. In wing chord males may measure 259 to 286 mm (10.2 to 11.3 in) and females may measure 268 to 298 mm (10.6 to 11.7 in). [9] [10] The tail measures 148 to 166 mm (5.8 to 6.5 in) in males and 154 to 171 mm (6.1 to 6.7 in) in females. In both sexes, the tarsus may measure 45 to 53 mm (1.8 to 2.1 in) and the bill 28.5 to 34.5 mm (1.12 to 1.36 in). [9] [10] Unlike the species overall, the nominate subspecies neatly corresponds to Bergmann's rule (wherein animal are larger farther from the Equator). In northern Italy, the average wing chord in males and females, respectively, was 267 and 274.5 mm (10.51 and 10.81 in) and body mass averaged 445 and 543 g (0.981 and 1.197 lb) in the two sexes. [33] Much farther north in Finland, nominate race owls were notably larger, averaging 275 and 287 mm (10.8 and 11.3 in) in wing chord and 480 and 699 g (1.058 and 1.541 lb) in body mass. [9] [35] Taken as a whole, the nominate includes both the heavier and lightest recorded birds in the tawny owl species. [10] [33] [35]
S. a. sylvatica West Europe including Great Britain and the Iberian Peninsula Shaw, 1809Generally the appearance of S. a. sylvatica is not dissimilar from the nominate subspecies but on average it is more boldly patterned with considerably less white base colour showing below, particularly with a richer average hue in rufous and intermediate morph individuals. [9] [33] More significantly, the main song of this subspecies differs slightly from that of nominate subspecies based on spectrograms. [36] Linearly, this is a fairly small subspecies, averaging around 10% smaller than S. a. aluco, and may include the smallest known tawny owls going on standard measurements. [9] [37] However, average weights do not significantly differ from those of other subspecies with published weights. [33] [37] Wing chord measurements may range from 248 to 280 mm (9.8 to 11.0 in) in males and from 255 to 296 mm (10.0 to 11.7 in) in females. [9] [10] [38] [39] In Spain, the tail could measure 140 to 191 mm (5.5 to 7.5 in), averaging 167.8 mm (6.61 in), the tarsus could measure 47 to 61 mm (1.9 to 2.4 in), averaging 53.85 mm (2.120 in) and the bill could measure 24 to 31 mm (0.94 to 1.22 in), averaging 28.5 mm (1.12 in). [38] Average wing chord in males from England and France were 260.9 and 268 mm (10.27 and 10.55 in) respectively while those of females were 273.6 and 276 mm (10.77 and 10.87 in). [40] [39] Average weights in England and France were 408.6 and 427.8 g (14.41 and 15.09 oz) for 22 and 66 males and 533 and 567 g (1.175 and 1.250 lb) in 20 and 50 females. [40] [39] Live adult weights can range from 335 to 580 g (0.739 to 1.279 lb) in males and 430 to 780 g (0.95 to 1.72 lb). [39]
S. a. biddulphi NW India and Pakistan Scully, 1881 [41] This isolated subspecies is fairly distinct for its stark grey morph, with other morphs either rare or non-existent. [9] It has a more stark apparent whitish base colour apparent with a strong grey wash on the head and mantle as well as strong herringbone patterning below. Altogether, it lacks the warmer tones common in more westerly tawny owls and its colouring is not dissimilar from a Ural owl but for the herringbone pattern. [9] [42] [43] Although at times apparently hypothesized as a separate form, [9] most authors continue to retain it as a proper subspecies of tawny owl. [10] [33] Another distinct feature of S. a. biddulphi is its relatively large size and it appears to be the largest bodied race of tawny owl, although published weights are not known. [10] Wing chord in males was found to be 285 to 323 mm (11.2 to 12.7 in) whilst that of females measures 320 to 345 mm (12.6 to 13.6 in). The tail may measure 191 to 210 mm (7.5 to 8.3 in) while a single bird had a tarsal length of 51 mm (2.0 in) and two birds had bill lengths of 33 and 35 mm (1.3 and 1.4 in). [9] [10]
S. a. willkonskii Palestine, Asia Minor to N Iran and the Caucasus up to southeastern Europe (Menzbier, 1896)In likelihood, this subspecies includes the formerly described race of S. a. obscurata. [9] On the whole, this race tends to be somewhat more richly coloured than the nominate subspecies. Particularly unique within this subspecies is a dark morph which is somewhat rufous but can grade into an almost coffee brown hue. [9] [31] Although some authors consider this a small subspecies, [9] measurements suggest it is more so of intermediate size. [10] In fact, average sizes may be exceed those of nominate race tawny owls from further north in Europe and the male song may consequently have a slightly deeper tone as well. [44] Wing chord in males may measure 255 to 296 mm (10.0 to 11.7 in) while females may measure 282 to 305 mm (11.1 to 12.0 in). Furthermore, weight of one male S. a. willkonskii was 510 g (1.12 lb) while one female weighed 582 g (1.283 lb). [10]
S. a. mauritanica NW Africa from Morocco to Tunisia and Mauritania (Witherby, 1905)This race is a rather dark grey-brown with no evidence of morph colour variation. It is slightly dull, cooler and more uniform hue overall than S. a. aluco or S. a. sylvatica. [9] [10] Study of genetic materials, including phylogeography, and the species dispersal behaviour, supports the division of S. a. maurtanica from the European subspecies, and it may even form its own species, with the Strait of Gibraltar as a natural gap between the ranges. [45] Although sometimes suggested as about 5% smaller than S. a. aluco, [12] current data suggest it is of similar size to Italian nominate owls and Spanish S. a. sylvatica found on the other side of the Mediterranean. [10] [33] Standard measurements of both sexes are known to be 272 to 305 mm (10.7 to 12.0 in) in wing chord length, 173 to 189 mm (6.8 to 7.4 in) in tail length, 54 to 61 mm (2.1 to 2.4 in) in tarsal length and 28 to 31 mm (1.1 to 1.2 in) in bill length. Weight of males has been reported at 325 to 470 g (11.5 to 16.6 oz) whilst that of females at 390 to 575 g (0.860 to 1.268 lb). [10]
S. a. sanctinicolaiW Iran, NE Iraq (Zarudny, 1905) [46] This little known subspecies is apparently a rather pale and washed-out form, as excepted for a species that lives in subdesert region. [9] Although the only known measurements obtained have been of wing chord it appears to be one of the smaller forms of the tawny owl. Males may measure from 255 to 273 mm (10.0 to 10.7 in) and females have been known to measure 270 to 285 mm (10.6 to 11.2 in). [10]
S. a. harmsiThe area once known as Turkestan, which today includes portions of six various countries.(Zarudny, 1911) [47] This is a relatively dark hued form, which may be in some way intermediate with the Himalayan owl based on colouring but is still considered part of the tawny owl species. [10] [31] This race is quite large based on wing chord dimensions, and may rival S. a. biddulphi as the largest form of tawny owl. Measurements for males are 303 to 316 mm (11.9 to 12.4 in) while females they are 318 to 332 mm (12.5 to 13.1 in). [9] [10]
S. a. siberiae C Russia from the Urals to about the Irtysh river in Western Siberia Dementiev, 1933This race is paler still than the nominate race with a large amount of dazzling white apparent on the sparsely marked underside, which tends to bare relatively few crossbars. [48] This is a relatively large subspecies, being fairly similar in size to the nominate birds from Scandinavia. [12] This race is up to 12% larger than Central European nominate birds. [12] [49] Wing chord may measure from 280 to 300 mm (11 to 12 in) in males and from 301 to 311 mm (11.9 to 12.2 in) in females. A single owl measured 175 mm (6.9 in) in tail length and 33 mm (1.3 in) in bill length. [10] Unexpectedly, the reported weights for S. a. siberiae are not high relative to most reported in Europe and come in at a similar range as those reported for linearly rather smaller populations such as S. a. sylvatica in France. [10] [39] Reported body mass for S. a. siberiae is 450 to 490 g (0.99 to 1.08 lb) in males and 590 to 680 g (1.30 to 1.50 lb) in females. [10]

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<span class="mw-page-title-main">Dietary biology of the tawny owl</span>

The tawny owl is an opportunistic and generalized predator. Peak hunting activity tends to occur largely between dusk to midnight, with owls often following an erratic hunting pattern, perhaps to sites where previous hunts were successful. When feeding young, hunting may need to be prolonged into daylight in the early morning. Based on hand-reared young owls that re-released into the wild, hunting behaviour is quite innate rather than learned. Normally this owl hunts from a perch. Perching bouts usually last from about 8 to 14 minutes depending largely on habitat. Tawny owl's hunting from a perch or pole can recall a buzzard and the two take similar prey sizes as well. However, high initial speed and maneuvering among trees and bushes with great dexterity may allow it to surprise relatively large prey, more like a goshawk. The tawny owl is capable of lifting and carrying off in flight individual prey weighing up to at least 320 g (11 oz). Their middle talon, the most enlarged claw on owls, measures an average of 19.1 mm (0.75 in). While not as large as those of the Ural owl, the talons are extremely sharp, stout and quite decurved. The claws are considered to be visibly more overdeveloped than those of other European mid-sized owls and the footspan including the claws is fairly larger as well, at an average of about 13.4 cm (5.3 in). The hunting owl often extends its wings to balance and control prey upon impact. Alternatively, this species may hunt from flight. This occurs from 2 to 3 m over the ground, often over open habitats such as bushes, marsh or grassland, forming a quartering or zigzag pattern over the opening. During these flights they cover about 30 to 50 m before changing direction. Hunting from flight was surprisingly prevalent in a Swedish study of two radio-tagged birds, with 34% of study time spent hunting from flight while 40% of the study time was spent on hunting from a perch. In a similar study in England, less than 1% of time was spent hunting from flight. In a more deliberate variation of hunting from flight, the hunting owl may examine crags and nest boxes or also hover around prey roosts. In the latter type of hunts, the tawny owls may strike branches and/or beat their wings together in front of denser foliage, bushes or conifers in order to disturb and flush prey such as small birds and bats, or may dive directly into said foliage. Hovering has also been recorded in differing circumstances, including one incidence of an owl hunting a small bird that was caught on the wing after a hovering flight. Tawny owls have also taken bats on the wing as well and have been seen to hawk large, relatively slow-flying insects such as some beetles and moths in flight. Caterpillars may too be taken from trees. Usually these hunting variations are correlated with poor weather hampering the capture of preferred prey. Tawny owls eat worms with relative frequency, as they often hear them apparently from below the surface and snatch them up from shallow dirt or below leaf litter. Their worm-hunting style recalls worm hunting techniques by most other birds and they were recorded to eat 0.39 worms per minute during an hour of observation in England and were sometimes seen to feed on worms during daylight. Other hunting from the ground has been observed, often of insects such as beetles, but tawny owls have also been reported to "leap" upon from a ground vantage point in order to capture a vole, quite like foxes often do. There are now many accounts of tawny owls feeding on carrion from a wide range of sources, including hares, rats, sheep, and trout.

<span class="mw-page-title-main">Breeding biology of the tawny owl</span>

Tawny owls are monogamous and territorial year around. Young birds select territories and look for mates in autumn and tend to be very vocal, especially males. Due to their highly territorial behaviour, young birds frequently struggle to establish a territory unless a nearby adult dies. Males routinely engage in territorial fights. Territories have been known to have been maintained by single tawnys for up to 10 years in Russia and 13 years in Berlin. Of 34 males in Wytham, only one male moved off of territory, due to being disturbed by humans. It appears to be largely up to the male to select territorial boundaries. Despite the aforementioned territorial behaviour, active nests of two separate pairs at as close as 100 m (330 ft), in the Tegel forest, have been reported. This species shows very little extrapair parentage. In Switzerland for example, a study of 137 nestings found that only one, or 0.7%, were from a different father than the mate, females cannot generally raise young without male contribution so the pair structure of these highly residential owls insures little instance of cuckoldry. Cases of bigamy were reported at Wytham in 6 of 34 males, in situations where apparently a neighboring male died and was suffixed subsequently, however, one or the other nesting attempts would completely fail each time. In Pavia, 3 of 22 territories included two mature females.

References

  1. ↑ Linnaeus, C (1758). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata (in Latin). Holmiae. (Laurentii Salvii). p. 93. S. capite laevi, corpore ferrugineo, iridíbus atris, remi-gibus primoribus serratís.
  2. ↑ "Tawny Owl Strix aluco[Linnaeus, 1758]". BirdFacts. British Trust for Ornithology (BTO). Retrieved 31 May 2008.
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  4. 1 2 Holt, D.W., Berkley, R., Deppe, C., Enríquez Rocha, P., Petersen, J.L., Rangel Salazar, J.L., Segars, K.P., Wood, K.L. & Marks, J.S. (2019). Typical Owls. In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (eds.). Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona.
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  6. ↑ Gill, Frank; Donsker, David, eds. (2019). "Owls". World Bird List Version 9.1. International Ornithologists' Union. Retrieved 2 April 2019.
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  8. ↑ Mlíkovský, Jirí (2002): Cenozoic Birds of the World, Part 1: Europe. Ninox Press, Prague.
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  11. ↑ Lee, M. Y., Lee, S. M., Jeon, H. S., Lee, S. H., Park, J. Y., & An, J. (2018). Complete mitochondrial genome of the Northern Long-eared Owl (Asio otus Linnaeus, 1758) determined using next-generation sequencing. Mitochondrial DNA Part B, 3(2), 494–495.
  12. 1 2 3 4 5 6 7 8 Voous, Karel H.; Cameron, Ad (illustrator) (1988). Owls of the Northern Hemisphere. London, Collins. pp. 209–219. ISBN   978-0-00-219493-8.
  13. ↑ van der Weyden, W. J. (1975). Scops and screech owls: vocal evidence for a basic subdivision in the genus Otus (Strigidae). Ardea, 63, 65-77.
  14. ↑ Voous, K. H. (1964). Wood owls of the genera Strix and Ciccaba. Zoologische Mededelingen, 39(46), 471–478.
  15. ↑ Salomonsen, F. (1931). Beretning om en Rejse til Færøerne. Dansk Orn. Foren. Tidsskr, 25, 3–37.
  16. ↑ Scherzinger, W. (1980). Zur Ethologie der Fortpflanzung und Jugendentwicklung des Habichtkauzes (Strix uralensis) mit Vergleichen zum Waldkauz (Strix aluco). Bonn. Zool. Mongr, 15.
  17. ↑ Jánossy D. (1972). Die mittelpleistozäne Vogelfauna der Stránská skála. – In: Musil R. (ed.): Stránská skála I. – Anthropos (Brno) 20: 35–64.
  18. 1 2 Jánossy D. (1976). Die Felsnische Tarkő und die Vertebratenfauna ihrer Ausfüllung. Karsztés Barlangkutatás 8: 3–106.
  19. 1 2 Jánossy, D. 1978. Új finomrétegtani szint Magyarország pleisztocén őslénytani sorozatában [A new fine stratigrafic level in Paleontological series at Hungarian Pleistocene]. Földrajzi Közlemények 26(1–3): 161–174.
  20. 1 2 Mourer-Chauvire, C. (1975). Faunes d'oiseaux du Pléistocene de France: systématique, évolution et adaptation, interprétation paléoclimatique. Geobios, 8(5), 333-IN11.
  21. ↑ Becker, C., & Pieper, H. (1982). Zum Nachweis des Habichtkauzes Strix uralensis in einer neolithischen Seeufersiedlung der Schweiz. Der Ornithologische Beobachter, 79, 159–162.
  22. ↑ Scherzinger, W. (1983). Beobachtungen an Waldkauz-Habichtskauz-Hybriden:(Strix aluco x Strix uralensis).
  23. ↑ Heidrich, P., & Wink, M. (1994). Tawny owl (Strix aluco) and Hume's tawny owl (Strix butleri) are distinct species: evidence from nucleotide sequences of the cytochrome b gene. Zeitschrift für Naturforschung C, 49(3–4), 230–234.
  24. 1 2 Kirwan, G. M., Schweizer, M., & Copete, J. L. (2015). Multiple lines of evidence confirm that Hume’s Owl Strix butleri (AO Hume, 1878) is two species, with description of an unnamed species (Aves: Non-Passeriformes: Strigidae). Zootaxa, 3904(1), 28–50.
  25. ↑ Robb, M. S., Sangster, G., Aliabadian, M., van den Berg, A. B., Constantine, M., Irestedt, M., Khani, A., Musavi, S.B., Nunes, J.M.G., Willson, M.S. & Walsh, A. J. (2016). The rediscovery of Strix butleri (Hume, 1878) in Oman and Iran, with molecular resolution of the identity of Strix omanensis Robb, van den Berg and Constantine, 2013. Avian Research, 7(1), 7.
  26. ↑ Del Hoyo, J., Collar, N. J., Christie, D. A., Elliott, A., & Fishpool, L. D. C. (2014). HBW and BirdLife International Illustrated Checklist of the Birds of the World: non-passerines (Vol. 1). Barcelona: Lynx Edicions.
  27. ↑ Sibley, C. G., & Monroe Jr, B. L. (1993). A world checklist of birds. Ann Arbor, MI: Edwards Brothers Inc.
  28. ↑ Dov, A.B., Atar, A., Baruchi, E, Levi, K. & Sapir, N. (2017). Breeding biology of Desert Owl in Israel. Dutch Birding, 39, 285–307.
  29. ↑ BirdLife International (2016). "Strix aluco". IUCN Red List of Threatened Species . 2016: e.T22725469A86871093. doi: 10.2305/IUCN.UK.2016-3.RLTS.T22725469A86871093.en .
  30. ↑ Marcot, B. G. (1995). Owls of old forests of the world. Gen. Tech. Rep. PNW-GTR-343. Portland, OR: US Department of Agriculture, Forest Service, Pacific Northwest Research Station. 64 p, 343.
  31. 1 2 3 4 Owls of the World: A Photographic Guide by Mikkola, H. Firefly Books (2012), ISBN   9781770851368
  32. ↑ "Tawny Owl Strix aluco". Owl Information. World Owl Trust. Archived from the original on 13 June 2008. Retrieved 7 November 2017.
  33. 1 2 3 4 5 6 7 Galeotti, P. (2001). Strix aluco Tawny Owl. Birds of Western Palearctic. Update, 3(1), 43–77.
  34. ↑ Brito, P. H. (2007). Contrasting patterns of mitochondrial and microsatellite genetic structure among Western European populations of tawny owls (Strix aluco). Molecular Ecology, 16(16), 3423–3437.
  35. 1 2 Solonen, T. (2011). Effects of variable food supply on the body condition of breeding Tawny Owls Strix aluco in southern Finland. ISRN Zoology, 2011.
  36. ↑ Galeotti, P. R., Appleby, B. M., & Redpath, S. M. (1996). Macro and microgeographical variations in the ‘hoot’of Italian and English tawny owls (Strix aluco). Italian Journal of Zoology, 63(1), 57–64.
  37. 1 2 Mikkola, H. (1983). Owls of Europe. T. & AD Poyser.
  38. 1 2 Martinez, J.A., Zuberogoitia, I. & Alonso, R. (2002). Guía para la determinación de la edad y el sexo en las estrigiformes ibéricas- CÁRABO COMÚN TAWNY OWL Strix aluco. Monticola: 95–102.
  39. 1 2 3 4 5 Baudvin, H., & Dessolin, J. L. (1992). Analyse de la morphométrie de la chouette hulotte Strix aluco en Bourgogne. Alauda, 60(2), 93–104.
  40. 1 2 Hardy, A. R., Hirons, G. J. M., Stanley, P. I., & Huson, L. W. (1981). Sexual dimorphism in size of tawny owls (Strix aluco): a method for sexing in field studies. Ardea, 69, 181–184.
  41. ↑ Scully, J. (1881). A Contribution to the Ornithology of Gilgit. Ibis, 23(3), 415–453.
  42. ↑ Grewal, B., & Pfister, O. (1998). A photographic guide to birds of the Himalayas. New Holland.
  43. ↑ Blanford, W. T. (1894). Notes on the Indian Owls. Ibis, 36(4), 524–531.
  44. ↑ Shekhovtsov, S. M., & Sharikov, A. V. (2015). Individual and geographical variation in the territorial calls of Tawny Owls Strix aluco in Eastern Europe. Ardeola, 62(2), 299–311.
  45. ↑ Doña, J., Ruiz-Ruano, F. J., & Jovani, R. (2016). DNA barcoding of Iberian Peninsula and North Africa Tawny Owls Strix aluco suggests the Strait of Gibraltar as an important barrier for phylogeography. Mitochondrial DNA Part A, 27(6), 4475–4478.
  46. ↑ Zarudny, N. 1905. [Itinerary of the voyage to western Iran in the year 1903–1904]. Ezhegodnik’ Zoologicheskago Muzeya Imperatorskoi Akademii Nauk 9.
  47. ↑ Zarudny, N.A. (1911) Verzeichnis der Vögel Persiens. Journal für Ornithologie, 59: 185–241.
  48. ↑ Dementiev, G. P., Gladkov, N. A., Ptushenko, E. S., Spangenberg, E. P., & Sudilovskaya, A. M. (1966). Birds of the Soviet Union, vol. 1. Israel Program for Scientific Translations, Jerusalem.
  49. ↑ Bergmann, Carl (1847). "Über die Verhältnisse der Wärmeökonomie der Thiere zu ihrer Grösse". Göttinger Studien (in German). 3 (1): 595–708.
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