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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Infraclassis: Archosauromorpha
Cladus: Crurotarsi
Divisio: Archosauria
Cladus: Avemetatarsalia
Cladus: Ornithodira
Subtaxon: Dinosauromorpha
Cladus: Dinosauriformes
Cladus: Dracohors
Cladus: Dinosauria
Ordo: Saurischia
Cladus: Eusaurischia
Subordo: Theropoda
Cladus: Neotheropoda
Cladus: Averostra
Cladus: Tetanurae
Cladus: Avetheropoda
Cladus: Coelurosauria
Cladus: Tyrannoraptora
Cladus: Maniraptoromorpha
Cladus: Maniraptoriformes
Cladus: Maniraptora
Cladus: Pennaraptora
Cladus: Paraves
Cladus: Eumaniraptora
Cladus: Avialae
Infraclassis: Aves
Cladus: Euavialae
Cladus: Avebrevicauda
Cladus: Pygostylia
Cladus: Ornithothoraces
Cladus: Ornithuromorpha
Cladus: Carinatae
Parvclassis: Neornithes
Cohors: Neognathae
Cladus: Neoaves
Ordo: Strigiformes

Familia: Strigidae
Subfamilia: Striginae
Genus: Otus
Species: O. alfrediO. alius – – O. bakkamoena – O. balliO. beccarii – O. brookii – O. brucei – O. capnodes – O. collari – O. cyprius – O. elegans – O. enganensis – O. everetti – O. feae – O. fuliginosus – O. gurneyi – O. hartlaubi – O. icterorhynchus – O. insularis – O. ireneae – O. jolandae – O. lempiji – O. lettia – O. longicornis – O. madagascariensis – – O. manadensis – O. mantananensis – O. mayottensis – O. megalotis – O. mendeni – O. mentawi – O. mindorensis – O. mirus – O. moheliensis – O. nigrorum – O. pamelae – O. pauliani – O. pembaensis – O. podarginus – O. rufescensO. rutilus – O. sagittatus – O. scops – O. semitorques – O. senegalensis – O. siaoensis – O. silvicola – O. socotranus – O. spilocephalus – O. sulaensis – O. sunia – O. tempestatis – O. thilohoffmanni – O. umbraO. magicusO. angelinae

Species extintae (formerly in Mascarenotus): O. grucheti – O. murivorus – O. sauzieri
Species extintae: O. frutuosoi – O. mauli
Name

Otus Pennant, 1769

Typus: Otus bakkamoena Pennant, 1769

Synonyms

Scops Savigny, 1809
Scopus Oken, 1817 (emend.)
Pisorhina Kaup, 1848 Isis col. 769 BHL
Ephialtes Keyserling & Blasius, 1851
Lempijius Bonaparte, 1854
Athenoptera Hume, 1870
Gymnoscops Tristram, 1880
Heteroscops Sharpe, 1889
Mimizuku Hachisuka, 1934
Pyrroglaux Yamashina, 1938
Mascarenotus Mourer-Chauviré, Bour, Moutou & Ribes, 1994

References
Primary references

Indian Zoology p. 3

Additional references

Heidrich, P., König, C. & Wink, M. 1995. Molecular phylogeny of the South American Otus atricapillus complex (Aves Strigidae) inferred from nucleotide sequences of the mitochondrial cytochrome b gene. Zeitschrift für Naturforschung. 50c: 294–302. Full article (PDF)Reference page.
Wink, M. & Heidrich, P. 1999. Molecular evolution and systematics of owls (Strigiformes). In: König, C.; Weick, F. & Becking, J.H. (eds.) Owls: A guide to the owls of the world. pp.39–57. Yale University Press, New Haven. ISBN 0-300-07920-6 Full article (PDF)Reference page.
König, C., Weick, F., & Becking, J.-H. (2008). Owls of the World. Helm ISBN 978-0-7136-6548-2.
Miranda, H.C., Jr., Brooks, D.M. & Kennedy, R.S.. 2011. Phylogeny and taxonomic review of Philippine lowland scops owls (Strigiformes): parallel diversification of highland and lowland clades. Wilson Journal of Ornithology 123: 4441–454. DOI: 10.1676/10-185.1 PDF Reference page.
Rando, J.C., Pieper, H., Alcover, J.A. & Olson, S.L. 2012. A new species of extinct fossil scops owl (Aves: Strigiformes: Strigidae: Otus) from the Archipelago of Madeira (North Atlantic Ocean). Zootaxa 3182: 29–42. Preview Reference page.
Rando, J.C., Alcover, J.A., Olson, S.L. & Pieper, H. 2013. A new species of extinct scops owl (Aves: Strigiformes: Strigidae: Otus) from São Miguel Island (Azores Archipelago, North Atlantic Ocean). Zootaxa 3647(2): 343–357. DOI: 10.11646/zootaxa.3647.2.6 Reference page.
Flint, P., Whaley, D., Kirwan, G.M., Charalambides, M., Schweizer, M. & Wink, M. 2015. Reprising the taxonomy of Cyprus Scops Owl Otus (scops) cyprius, a neglected island endemic. Zootaxa 4040(3): 301–316. DOI: 10.11646/zootaxa.4040.3.3. Preview (PDF) Reference page.
Louchart, A., Bastian, F., Baptista, M., Guarino-Vignon, P., Hume, J.P., Jacot-des-Combes, C., Mourer-Chauviré, C., Hänni, C. & Ollivier, M. 2018. Ancient DNA reveals the origins, colonization histories, and evolutionary pathways of two recently extinct species of giant scops owl from Mauritius and Rodrigues Islands (Mascarene Islands, south-western Indian Ocean). Journal of Biogeography. 45 (12): 2678–2689. DOI: 10.1111/jbi.13450 PaywallReference page.
Salter, J.F., Oliveros, C.H., Hosner, P.A., Manthey, J.D., Robbins, M.B., Moyle, R.G., Brumfield, R.T. & Faircloth, B.C. 2019. Extensive paraphyly in the typical owl family (Strigidae). The Auk. 137 (ukz070). DOI: 10.1093/auk/ukz070 Open access Reference page.

Vernacular names
čeština: Výreček malý
Deutsch: Zwergohreulen
Ελληνικά: Ώτος
English: Scops owls
español: Autillo
suomi: Pöllöset
עברית: שָׂעִיר
日本語: コノハズク属
lietuvių: Apuokėliai
Türkçe: Cüce baykuş
中文: 角鸮属

Scops owls are typical owls in family Strigidae belonging to the genus Otus and are restricted to the Old World. Otus is the largest genus of owls with 58 species. Scops owls are colored in various brownish hues, sometimes with a lighter underside and/or face, which helps to camouflage them against the bark of trees. Some are polymorphic, occurring in a greyish- and a reddish-brown morph. They are small and agile, with both sexes being compact in size and shape. Female scops owls are usually larger than males.

For most of the 20th century, this genus included the American screech owls, which are now again separated in Megascops based on a range of behavioral, biogeographical, morphological and DNA sequence data.

Taxonomy
A well-camouflaged African scops owl (Otus senegalensis)

The genus Otus was introduced in 1769 by the Welsh naturalist Thomas Pennant for the Indian scops owl (O. bakkamoena).[1] The name is derived from the Latin word otus and the Greek word ὦτος ōtos meaning horned or eared owl (cf. οὖς, GEN ὠτός, "ear").[2][3][4][5] The generic name Scops that was proposed by Marie Jules César Savigny in 1809 is a junior synonym[6] and is derived from the Greek σκώψ skōps meaning small kind of owl, Otus scops.[7]

By the mid-19th century, it was becoming clear that Otus encompassed more than one genus. First, in 1848, the screech owls were split off as Megascops. The white-faced owls of Africa, with their huge eyes and striking facial coloration, were separated in Ptilopsis in 1851. In 1854, the highly apomorphic white-throated screech owl of the Andes was placed in the monotypic genus Macabra. Gymnasio was established in the same year for the Puerto Rican owl, and the bare-legged owl (or "Cuban screech owl") was separated in Gymnoglaux the following year; the latter genus was sometimes merged with Gymnasio by subsequent authors. The Palau owl, described only in 1872 and little-known to this day, was eventually separated in Pyrroglaux by Yoshimaro Yamashina in 1938.

In the early 20th century, the lumping-together of taxa had come to be preferred. The 3rd edition of the AOU checklist in 1910 placed the screech owls back in Otus. Although this move was never unequivocally accepted, it was the dominant treatment throughout most of the 20th century. In 1988 it was attempted to resolve this by re-establishing all those genera split some 140 years earlier at subgenus rank inside Otus.[8] Still, the diversity and distinctness of the group failed to come together in a good evolutionary and phylogenetic picture, and it was not until the availability of DNA sequence data that this could be resolved. In 1999, a preliminary study of mtDNA cytochrome b across a wide range of owls found that even the treatment as subgenera was probably unsustainable and suggested that most of the genera proposed around 1850 should be accepted.[9] Though there was some debate about the reliability of these findings at first,[10] they have been confirmed by subsequent studies. In 2003, the AOU formally re-accepted the genus Megascops again.[11]
Species

The genus Otus contains 58 species:[12]

Giant scops owl, Otus gurneyi
White-fronted scops owl, Otus sagittatus
Reddish scops owl, Otus rufescens
Serendib scops owl, Otus thilohoffmanni
Sandy scops owl, Otus icterorhynchus
Sokoke scops owl, Otus ireneae
Andaman scops owl, Otus balli
Flores scops owl, Otus alfredi
Mountain scops owl, Otus spilocephalus
Javan scops owl, Otus angelinae
Mindanao scops owl, Otus mirus
Luzon scops owl, Otus longicornis
Mindoro scops owl, Otus mindorensis
São Tomé scops owl, Otus hartlaubi
Torotoroka scops owl, Otus madagascariensis – formerly included in O. rutilus
Rainforest scops owl, Otus rutilus
Mayotte scops owl, Otus mayottensis – formerly included in O. rutilus
Karthala scops owl, Otus pauliani
Anjouan scops owl, Otus capnodes
Moheli scops owl, Otus moheliensis
† Réunion owl, Otus grucheti – extinct, formerly placed in the genus Mascarenotus
† Mauritius owl, Otus sauzieri – extinct, formerly placed in the genus Mascarenotus
† Rodrigues owl, Otus murivorus – extinct, formerly placed in the genus Mascarenotus
Pemba scops owl, Otus pembaensis
Eurasian scops owl, Otus scops
Cyprus scops owl, Otus cyprius – formerly included in O. scops
Pallid scops owl, Otus brucei
Arabian scops owl, Otus pamelae
African scops owl, Otus senegalensis
Annobón scops owl, Otus feae – formerly included in O. senegalensis
Socotra scops owl, Otus socotranus
Oriental scops owl, Otus sunia
Ryūkyū scops owl, Otus elegans
Moluccan scops owl, Otus magicus
Wetar scops owl, Otus tempestatis
Sula scops owl, Otus sulaensis
Biak scops owl, Otus beccarii
Sulawesi scops owl, Otus manadensis
Banggai scops owl, Otus mendeni
Siau scops owl, Otus siaoensis
Sangihe scops owl, Otus collari
Mantanani scops owl, Otus mantananensis
Seychelles scops owl, Otus insularis
Nicobar scops owl, Otus alius
Simeulue scops owl, Otus umbra
Enggano scops owl, Otus enganensis
Mentawai scops owl, Otus mentawi
Rajah scops owl, Otus brookii
Indian scops owl, Otus bakkamoena
Collared scops owl, Otus lettia – formerly included in O. bakkamoena
Japanese scops owl, Otus semitorques – formerly included in O. bakkamoena
Sunda scops owl, Otus lempiji – formerly included in O. bakkamoena
Philippine scops owl, Otus megalotis
Negros scops owl, Otus nigrorum – formerly included in O. megalotis
Everett's scops owl, Otus everetti – formerly included in O. megalotis
Palawan scops owl, Otus fuliginosus
Wallace's scops owl, Otus silvicola
Rinjani scops owl, Otus jolandae'
Palau owl, Otus podarginus – formerly placed in the monotypic genus Pyrroglaux

Two extinct species are sometimes placed in the genus:

† Madeiran scops owl, Otus mauli (extinct, c. 15th century)
† São Miguel scops owl, Otus frutuosoi (extinct, c. 15th century)

An apparent Otus owl was heard calling at about 1,000 meters ASL south of the summit of Camiguin in the Philippines on May 14, 1994. No scops owls had previously known from this island, and given that new species of Otus are occasionally discovered, it may have been an undescribed taxon.[13][14]

In July 2016, an unknown Otus species was photographed on Príncipe. The image was published on Ornithomedia.[15]
Formerly placed here

As noted above, the fossil record of scops owls gives an incomplete picture of their evolution at present. While older sources cite many species of supposed extinct Otus (or "Scops"), these are now placed in entirely different genera:[16]

"Otus" henrici was a barn owl of the genus Selenornis
"Otus" providentiae was a burrowing owl, probably a paleosubspecies
"Otus" wintershofensis may be close to extant genus Ninox and some material assigned to it belongs into Intutula
"Scops" commersoni is a junior synonym of the recently extinct Mauritius owl, referring to pictures and descriptions which mention ear tufts; the subfossil material of this species had been erroneously assigned to tuftless owls.

Evolution

The evolutionary relationships of the scops and screech owls are not entirely clear. What is certain is that they are very closely related; they may be considered sister lineages which fill essentially the same ecological niche in their allopatric ranges. A screech-owl fossil from the Late Pliocene of Kansas[17] – which is almost identical to eastern and western screech owls – indicate a long-standing presence of these birds in the Americas, while coeval scops owl fossils very similar to the Eurasian scops-owl have been found at S'Onix on the Spanish island Majorca.[16] The scops and screech owl lineage probably evolved at some time during the Miocene (like most other genera of typical owls), and the three (see below) modern lineages separated perhaps roughly 5 million years ago. Note that there is no reliable estimate of divergence time, as Otus and Megascops are osteologically very similar, as is to be expected from a group that has apparently conserved its ecomorphology since before its evolutionary radiation. As almost all scops and screech owls today, their common ancestor was in all probability already a small owl, with ear tufts and at least the upper tarsus ("leg") feathered.

However that may be, the hypothesis that the group evolved from Old World stock[18] is tentatively supported by cytochrome b sequence data.[9][19]
Ecology and behaviour

While late 19th-century ornithologists knew little of the variation of these cryptic birds which often live in far-off places, with every new taxon being described a few differences between the Old and New World "scops" owls became more and more prominent. Namely, the scops owls give a whistling call or a row of high-pitched hoots with less than four individual hoots per second. This call is given in social interaction or when the owl tries to scare away other animals. The screech owls on the other hand are named for their piercing trills of more than four individual notes per second. They also have a kind of song, which is a short sequence of varying calls given by the males when they try to attract females to their nests, or between members of a pair. There are a few other differences such as the screech owls almost never being brown below which is common in scops owls, but the difference in vocalizations is most striking.

Scops owls hunt from perches in semi-open landscapes. They prefer areas which contain old trees with hollows; these are home to their prey which includes insects, reptiles, small mammals such as bats and mice and other small birds. The owls will also eat earthworms, amphibians and aquatic invertebrates.[20] Scops owls have a good sense of hearing which helps them locate their prey in any habitat. They also possess well-developed raptorial claws and a curved bill, both of which are used for tearing their prey into pieces small enough to swallow easily.

Scops owls are primarily solitary birds. Most species lay and incubate their eggs in a cavity nest that was originally made by another animal. During the incubation period, the male will feed the female. These birds are monogamous, with biparental care, and only fledge one young per year. The young of most scops owls are altricial to semialtricial.

As opposed to screech owls, scops owls have only a single type of call. This consists of a series of whistles or high-pitched hoots, given with a frequency of 4 calls per second or less, or of a single, drawn-out whistle. Calls differ widely between species in type and pitch, and in the field are often the first indication of these birds' presence, as well as the most reliable means to distinguish between species. Some, like the recently described Serendib scops owl (Otus thilohoffmanni), were discovered because their vocalizations were unfamiliar to experts in birdcalls.
See also

Mascarene owls

References

Pennant, Thomas (1769). "Otus bakkamoena". Indian Zoology. London. p. 3.
Jobling, J.A. (2010). "Otus". The Helm Dictionary of Scientific Bird Names. London: Christopher Helm. p. 286. ISBN 978-1-4081-2501-4.
otus. Charlton T. Lewis and Charles Short. A Latin Dictionary on Perseus Project.
ὦτος. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project.
οὖς in Liddell and Scott.
Savigny, M.J.C. (1809). "Scops Ephialtes. Le petit duc". Description de l'Égypte, ou recueil des observations et des recherches qui ont été faites en Égypte. Vol. I. Paris: L'Imprimerie Impériale. p. 107.
σκώψ in Liddell and Scott.
Marshall, J. T.; King, B. (1988). "Genus Otus". In Amadon, D.; Bull, J. (eds.). Hawks and owls of the world: A distributional and taxonomic list. Proceedings of the Western Foundation of Vertebrate Zoology. Vol. 3. pp. 296–357.
Heidrich, P.; König, C. & Wink, M. (1995). "Molecular phylogeny of the South American Otus atricapillus complex (Aves Strigidae) inferred from nucleotide sequences of the mitochondrial cytochrome b gene" (PDF). Zeitschrift für Naturforschung C. 50 (3–4): 294–302. doi:10.1515/znc-1995-3-420. PMID 7766262. S2CID 28746107.
South American Classification Committee (SACC) (2003). "Proposal (#58): Elevate subgenus Megascops (New World Otus) to full generic status". Archived from the original on 2008-05-16.
Banks, R. C.; Cicero, C.; Dunn, J. L.; Kratter, A. W.; Rasmussen, P. C.; Remsen, J.V. Jr.; Rising, J. D. & Stotz, D. F. (2003). "Forty-fourth supplement to the American Ornithologists' Union check-list of North American birds" (PDF). Auk. 120 (3): 923–931. doi:10.1642/0004-8038(2003)120[0923:fsttao]2.0.co;2.
Gill, Frank; Donsker, David; Rasmussen, Pamela, eds. (January 2021). "Owls". IOC World Bird List Version 11.1. International Ornithologists' Union. Retrieved 20 May 2021.
Balete, D. S.; Tabaranza, B. R. Jr. & Heaney, L. R. (2006). "An Annotated Checklist of the Birds of Camiguin Island, Philippines". Fieldiana Zoology. New Series. 106: 58. doi:10.3158/0015-0754(2006)106[58:AACOTB]2.0.CO;2.
Heaney, L. R. & Tabaranza, B. R. Jr. (2006). "Mammal and Land Bird Studies on Camiguin Island, Philippines: Background and Conservation Priorities". Fieldiana Zoology. New Series. 106: 1–13. doi:10.3158/0015-0754(2006)106[1:MALBSO]2.0.CO;2.
"An unknown bird of the island of Príncipe has been photographed" (in French). Ornithomedia. 2016. Retrieved 10 August 2016.
Mlíkovský, J. (2002). Cenozoic Birds of the World, Part 1: Europe. Prague: Ninox Press.
Ford, N. L. (1966). "Fossil Owls From the Rexroad Fauna of the Upper Pliocene in Kansas" (PDF). Condor. 68 (5): 472–475. doi:10.2307/1365319. JSTOR 1365319.
Johnson, D. (2003). "Owls in the Fossil Record". The owl pages.
Wink, M. & Heidrich, P. (1999). "Molecular evolution and systematics of owls (Strigiformes)" (PDF). In König, C.; Weick, F. & Becking, J.H. (eds.). A guide to the owls of the world. New Haven: Yale University Press. pp. 39–57. ISBN 0-300-07920-6.
Marchesi, L. & Sergio, F. (2005). "Distribution, density, diet and productivity of the Scops Owl Otus scops in the Italian Alps". Ibis. 147 (1): 176–187. doi:10.1111/j.1474-919x.2004.00388.x.

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