:Falcon

The genus Falco was introduced in 1758 by the Swedish naturalist Carl Linnaeus in the tenth edition of his Systema Naturae.{{cite book | last=Linnaeus | first=Carl | author-link=Carl Linnaeus | year=1758 | title= Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis | volume=1 | edition=10th | page=88 | publisher=Laurentii Salvii | location=Holmiae (Stockholm) | language=Latin | url=https://www.biodiversitylibrary.org/page/726995 }} The type species is the merlin (Falco columbarius).{{ cite book | editor-last1=Dickinson | editor-first1=E.C. | editor1-link=Edward C. Dickinson | editor-last2=Remsen | editor-first2=J.V. Jr. | editor2-link=James Van Remsen, Jr. | year=2013 | title=The Howard & Moore Complete Checklist of the Birds of the World | volume= 1: Non-passerines | edition=4th | location=Eastbourne, UK | publisher=Aves Press | isbn=978-0-9568611-0-8 | page=349 }} The genus name {{lang|la|Falco}} is Late Latin meaning a "falcon" from {{lang|la|falx}}, {{lang|la|falcis}}, meaning "a sickle", referring to the claws of the bird.{{cite book | last=Jobling | first=James A. | year=2010| title=The Helm Dictionary of Scientific Bird Names | publisher=Christopher Helm | location=London | isbn=978-1-4081-2501-4 | page=63 }}{{Cite book|title=Shorter Oxford English dictionary on historical principles|date=2007|publisher=Oxford University Press|editor-last1=Stevenson |editor-first1=Angus |editor-last2=Brown |editor-first2=Lesley |isbn=9780199206872|edition= 6th.|location=Oxford|oclc=170973920}} In Middle English and Old French, the title {{lang|enm|faucon}} refers generically to several captive raptor species.

The traditional term for a male falcon is tercel (British spelling) or tiercel (American spelling), from the Latin {{lang|la|tertius}} (third) because of the belief that only one in three eggs hatched a male bird. Some sources give the etymology as deriving from the fact that a male falcon is about one-third smaller than a female{{OEtymD|tercel}}{{cite web |url= http://dictionary.reference.com/browse/tercel |title= tercel |publisher= Dictionary.reference.com |access-date= 20 March 2010}}"tercel", Oxford Dictionary ({{langx|fro|tiercelet}}). A falcon chick, especially one reared for falconry, still in its downy stage, is known as an eyas{{cite web |url= http://www.thefreedictionary.com/eyas |title=eyas |publisher= Thefreedictionary.com |access-date= 20 March 2010}}{{cite web |url=http://www.tiscali.co.uk/reference/dictionaries/difficultwords/data/d0005484.html |title=Dictionary of Difficult Words – eyas |publisher=Tiscali.co.uk |date=21 September 1964 |access-date=20 March 2010 |url-status=dead |archive-url=https://web.archive.org/web/20090105233550/http://www.tiscali.co.uk/reference/dictionaries/difficultwords/data/d0005484.html |archive-date=5 January 2009}} (sometimes spelled eyass). The word arose by mistaken division of Old French {{lang|fro|un niais}}, from Latin presumed {{lang|la|nidiscus}} (nestling) from {{lang|la|nidus}} (nest). The technique of hunting with trained captive birds of prey is known as falconry.

Compared to other birds of prey, the fossil record of the falcons is not well distributed in time. For years, the oldest fossils tentatively assigned to this genus were from the Late Miocene, less than 10 million years ago.{{Cite journal |last1=Li |first1=Zhiheng |last2=Zhou |first2=Zhonghe |last3=Deng |first3=Tao |last4=Li |first4=Qiang |last5=Clarke |first5=Julia A. |date=June 4, 2014 |title=A falconid from the Late Miocene of northwestern China yields further evidence of transition in Late Neogene steppe communities |url=https://repositories.lib.utexas.edu/server/api/core/bitstreams/26c878fa-ddf5-456b-81ca-afd12dd48b0a/content |journal=The Auk: Ornithological Advances |volume=131 |pages=335–350}} This coincides with a period in which many modern genera of birds became recognizable in the fossil record. As of 2021, the oldest falconid fossil is estimated to be 55 million years old.{{Cite journal |last1=Mayr |first1=Gerald |last2=Kitchener |first2=Andrew C. |date=2021-10-01 |title=New fossils from the London Clay show that the Eocene Masillaraptoridae are stem group representatives of falcons (Aves, Falconiformes) |journal=Journal of Vertebrate Paleontology |language=en |volume=41 |issue=6 |doi=10.1080/02724634.2021.2083515 |issn=0272-4634|doi-access=free |bibcode=2021JVPal..41E3515M }}{{Cite web |last=Kirschbaum |first=Kari |title=Falconidae (falcons) |url=https://animaldiversity.org/accounts/Falconidae/ |access-date=2024-05-01 |website=Animal Diversity Web |language=en}} Given the distribution of fossil and living Falco taxa, falcons are probably of North American, African, or possibly Middle Eastern or European origin. Falcons are not closely related to other birds of prey, and their nearest relatives are parrots and songbirds.{{cite journal |vauthors=Suh A, Paus M, Kiefmann M |title=Mesozoic retroposons reveal parrots as the closest living relatives of passerine birds |journal=Nature Communications |volume=2|pages=443–8 |year=2011 |pmid=21863010 |doi=10.1038/ncomms1448 |pmc=3265382 |issue=8|bibcode=2011NatCo...2..443S |display-authors=etal}}

Falcons are roughly divisible into three or four groups. The first contains the kestrels (probably excepting the American kestrel); usually small and stocky falcons of mainly brown upperside colour and sometimes sexually dimorphic; three African species that are generally gray in colour stand apart from the typical members of this group. The fox and greater kestrels can be told apart at first glance by their tail colours, but not by much else; they might be very close relatives and are probably much closer to each other than the lesser and common kestrels. Kestrels feed chiefly on terrestrial vertebrates and invertebrates of appropriate size, such as rodents, reptiles, or insects.

The second group contains slightly larger (on average) species, the hobbies and relatives. These birds are characterized by considerable amounts of dark slate-gray in their plumage; their malar areas are nearly always black. They feed mainly on smaller birds.

Third are the peregrine falcon and its relatives, variably sized powerful birds that also have a black malar area (except some very light color morphs), and often a black cap, as well. They are very fast birds with a maximum speed of 390 kilometres per hour. Otherwise, they are somewhat intermediate between the other groups, being chiefly medium grey with some lighter or brownish colours on their upper sides. They are, on average, more delicately patterned than the hobbies and, if the hierofalcons are excluded (see below), this group typically contains species with horizontal barring on their undersides. As opposed to the other groups, where tail colour varies much in general but little according to evolutionary relatedness,For example, tail colour in the common and lesser kestrels is absolutely identical, yet they do not seem closely related. the tails of the large falcons are quite uniformly dark grey with inconspicuous black banding and small, white tips, though this is probably plesiomorphic. These large Falco species feed on mid-sized birds and terrestrial vertebrates.

Very similar to these, and sometimes included therein, are the four or so species of hierofalcon (literally, "hawk-falcons"). They represent taxa with, usually, more phaeomelanins, which impart reddish or brown colors, and generally more strongly patterned plumage reminiscent of hawks. Their undersides have a lengthwise pattern of blotches, lines, or arrowhead marks.

While these three or four groups, loosely circumscribed, are an informal arrangement, they probably contain several distinct clades in their entirety.

A study of mtDNA cytochrome b sequence data of some kestrels identified a clade containing the common kestrel and related "malar-striped" species, to the exclusion of such taxa as the greater kestrel (which lacks a malar stripe), the lesser kestrel (which is very similar to the common, but also has no malar stripe), and the American kestrel, which has a malar stripe, but its colour pattern – apart from the brownish back – and also the black feathers behind the ear, which never occur in the true kestrels, are more reminiscent of some hobbies. The malar-striped kestrels apparently split from their relatives in the Gelasian, roughly 2.0–2.5 million years ago (Mya), and are seemingly of tropical East African origin. The entire "true kestrel" group—excluding the American species—is probably a distinct and quite young clade, as also suggested by their numerous apomorphies.

File:FalconHeadBeak.png.]]

Other studies have confirmed that the hierofalcon are a monophyletic group–and that hybridization is quite frequent at least in the larger falcon species. Initial studies of mtDNA cytochrome b sequence data suggested that the hierofalcon are basal among living falcons. The discovery of a NUMT proved this earlier theory erroneous. In reality, the hierofalcon are a rather young group, originating at the same time as the start of the main kestrel radiation, about 2 Mya. Very little fossil history exists for this lineage. However, the present diversity of very recent origin suggests that this lineage may have nearly gone extinct in the recent past.

The phylogeny and delimitations of the peregrine and hobby groups are more problematic. Molecular studies have only been conducted on a few species, and the morphologically ambiguous taxa have often been little researched. The morphology of the syrinx, which contributes well to resolving the overall phylogeny of the Falconidae, is not very informative in the present genus. Nonetheless, a core group containing the peregrine and Barbary falcons, which, in turn, group with the hierofalcon and the more distant prairie falcon (which was sometimes placed with the hierofalcon, though it is entirely distinct biogeographically), as well as at least most of the "typical" hobbies, are confirmed to be monophyletic as suspected.

Given that the American Falco species of today belong to the peregrine group, or are apparently more basal species, the initially most successful evolutionary radiation seemingly was a Holarctic one that originated possibly around central Eurasia or in (northern) Africa. One or several lineages were present in North America by the Early Pliocene at latest.

The origin of today's major Falco groups—the "typical" hobbies and kestrels, for example, or the peregrine-hierofalcon complex, or the aplomado falcon lineage—can be quite confidently placed from the Miocene-Pliocene boundary through the Zanclean and Piacenzian and just into the Gelasian, that is from 2.4 to 5.3 Mya, when the malar-striped kestrels diversified. Some groups of falcons, such as the hierofalcon complex and the peregrine-Barbary superspecies, have only evolved in more recent times; the species of the former seem to be 120,000 years old or so.

The sequence follows the taxonomic order of White et al. (1996), except for adjustments in the kestrel sequence.

• Réunion kestrel, Falco duboisi – extinct (about 1700)

• Falco medius (Late Miocene of Cherevichnyi, Ukraine)IZAN 45-4033: left carpometacarpus. Small species; possibly closer to kestrels than to peregrine lineage or hierofalcons, but may be more basal altogether due to its age
• ?Falco sp. (Late Miocene of Idaho)IMNH 27937. A coracoid of a merlin-sized species. It does not seem close to F. columbarius or the Recent North American species (Becker 1987).
• Falco sp. (Early Pliocene of Kansas)
• Falco sp. (Early Pliocene of Bulgaria – Early Pleistocene of Spain and Czech Republic)A hierofalcon (Mlíkovský 2002)? If so, probably not close to the living species, but an earlier divergence that left no descendants; might be more than one species due to large range in time and/or include common ancestor of hierofalcons and peregrine-Barbary complex (Nittinger et al. 2005).
• Falco oregonus (Early/Middle Pliocene of Fossil Lake, Oregon) – possibly not distinct from a living species
• Falco umanskajae (Late Pliocene of Kryzhanovka, Ukraine) – includes "Falco odessanus", a nomen nudum
• ?Falco bakalovi (Late Pliocene of Varshets, Bulgaria)Boev, Z. 1999. Falco bakalovi sp. n. – a Late Pliocene falcon (Falconidae, Aves) from Varshets (W Bulgaria). – Geologica Balcanica, 29 (1–2): 131–35.Boev, Z. 2011. New fossil record of the Late Pliocene kestrel (Falco bakalovi Boev, 1999) from the type locality in Bulgaria. – Geologica Balcanica, 40 (1–3): 13–30.
• Falco antiquus (Middle Pleistocene of Noailles, France and possibly Horvőlgy, Hungary)Supposedly a saker falcon paleosubspecies (Mlíkovský 2002), but this is not too likely due to the probable Eemian origin of that species.
• Cuban kestrel, Falco kurochkini (Late Pleistocene/Holocene of Cuba, West Indies)
• Falco chowi (China)
• Falco bulgaricus (Late Miocene of Hadzhidimovo, Bulgaria)Boev, Z. 2011. Falco bulgaricus sp. n. (Aves, Falconiformes) from the Middle Miocene of Hadzhidimovo (SW Bulgaria). – Acta zoologica bulgarica, 63 (1): 17–35.

Several more paleosubspecies of extant species also been described; see species accounts for these.

"Sushkinia" pliocaena from the Early Pliocene of Pavlodar (Kazakhstan) appears to be a falcon of some sort. It might belong in this genus or a closely related one. In any case, the genus name Sushkinia is invalid for this animal because it had already been allocated to a prehistoric dragonfly relative. In 2015 the bird genus was renamed Psushkinia.Nikita V. Zelenkov; Evgeny N. Kurochkin (2015). "КЛАСС AVES". In E.N. Kurochkin; A.V. Lopatin; N.V. Zelenkov. Ископаемые позвоночные России и сопредельных стран. Ископаемые рептилии и птицы. Часть 3 / Fossil vertebrates of Russia and adjacent countries. Fossil Reptiles and Birds. Part 3. GEOS. pp. 86–290. {{ISBN|978-5-89118-699-6}}.

The supposed "Falco" pisanus was actually a pigeon of the genus Columba, possibly the same as Columba omnisanctorum, which, in that case, would adopt the older species name of the "falcon". The Eocene fossil "Falco" falconellus (or "F." falconella) from Wyoming is a bird of uncertain affiliations, maybe a falconid, maybe not; it certainly does not belong in this genus. "Falco" readei is now considered a paleosubspecies of the yellow-headed caracara (Milvago chimachima).

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{{cite journal|author=Becker, Jonathan J. |year=1987|title= Revision of "Falco" ramenta Wetmore and the Neogene evolution of the Falconidae|journal=Auk|volume=104|issue=2|pages= 270–76|jstor = 4087033|url=http://sora.unm.edu/sites/default/files/journals/auk/v104n02/p0270-p0276.pdf|doi=10.1093/auk/104.2.270}}

UMMP V27159, V29107, V57508-V57510, V57513/V57514 some limb bones. Slightly smaller than a merlin and more robust than American kestrel, and seems not too distant from F. columbarius. {{cite journal|author=Feduccia, J. Alan |year=1970|title= Some birds of prey from the Upper Pliocene of Kansas|journal=Auk|volume=87|issue=4|pages= 795–97|url=http://sora.unm.edu/sites/default/files/journals/auk/v087n04/p0795-p0797.pdf|doi=10.2307/4083714|last2=Ford|first2=Norman L.|jstor=4083714}}

{{cite journal|author=Griffiths, Carole S. |year=1999|title= Phylogeny of the Falconidae inferred from molecular and morphological data|journal=Auk|volume=116|issue=1|pages= 116–30|url=http://sora.unm.edu/sites/default/files/journals/auk/v116n01/p0116-p0130.pdf|doi=10.2307/4089459|jstor=4089459}}

{{cite journal|author1=Griffiths, Carole S. |author2=Barrowclough, George F. |author3=Groth, Jeff G. |author4=Mertz, Lisa |year=2004|title= Phylogeny of the Falconidae (Aves): a comparison of the efficacy of morphological, mitochondrial, and nuclear data|journal=Molecular Phylogenetics and Evolution|volume=32|issue=1|pages= 101–09|doi=10.1016/j.ympev.2003.11.019|pmid=15186800|bibcode=2004MolPE..32..101G }}

{{cite journal|author1=Groombridge, Jim J. |author2=Jones, Carl G. |author3=Bayes, Michelle K. |author4=van Zyl, Anthony J. |author5=Carrillo, José |author6=Nichols, Richard A. |author7=Bruford, Michael W. |year=2002|title= A molecular phylogeny of African kestrels with reference to divergence across the Indian Ocean|journal=Molecular Phylogenetics and Evolution|volume=25|issue=2|pages= 267–77|doi=10.1016/S1055-7903(02)00254-3|pmid=12414309|bibcode=2002MolPE..25..267G }}

Helbig, A.J.; Seibold, I.; Bednarek, W.; Brüning, H.; Gaucher, P.; Ristow, D.; Scharlau, W.; Schmidl, D. & Wink, Michael (1994): [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/1994/4.%201994.pdf Phylogenetic relationships among falcon species (genus Falco) according to DNA sequence variation of the cytochrome b gene]. In: Meyburg, B.-U. & Chancellor, R.D. (eds.): Raptor conservation today: pp. 593–99

{{cite journal|doi=10.1126/science.1257767|title=Falcon visual acuity|year=1976|last1=Fox|first1=R|last2=Lehmkuhle|first2=S.|last3=Westendorf|first3=D.|journal=Science|volume=192|issue=4236|pages=263–65|pmid=1257767|bibcode=1976Sci...192..263F}}

{{cite journal|author1=Johnson, J.A. |author2=Burnham, K.K. |author3=Burnham, W.A. |author4=Mindell, D.P. |year=2007|title= Genetic structure among continental and island populations of gyrfalcons|journal=Molecular Ecology|volume=16|pages=3145–60|doi=10.1111/j.1365-294X.2007.03373.x|issue=15|pmid=17651193|bibcode=2007MolEc..16.3145J |url=https://deepblue.lib.umich.edu/bitstream/2027.42/71471/1/j.1365-294X.2007.03373.x.pdf|hdl=2027.42/71471 |s2cid=17437176 |hdl-access=free}}

Fox Canyon Local Fauna, 4.3–4.8 million years ago: Martin, R.A.; Honey, J.G. & Pelaez-Campomanes, P. (2000): [http://www.kgs.ku.edu/Publications/OFR/2000/OFR00_61/index.html The Meade Basin Rodent Project; a progress report]. Kansas Geological Survey Open-file Report 2000-61. Paludicola 3(1): 1–32.

Mlíkovský, Jirí (2002): [http://www.nm.cz/download/JML-18-2002-CBE.pdf Cenozoic Birds of the World, Part 1: Europe] {{Webarchive|url=https://web.archive.org/web/20110520101755/http://www.nm.cz/download/JML-18-2002-CBE.pdf |date=20 May 2011 }}. Ninox Press, Prague

{{cite journal|author1=Nittinger, F. |author2=Haring, E. |author3=Pinsker, W. |author4=Wink, Michael |author5=Gamauf, A. |year=2005|title= Out of Africa? Phylogenetic relationships between Falco biarmicus and other hierofalcons (Aves Falconidae)|journal=Journal of Zoological Systematics and Evolutionary Research|volume=43|issue=4|pages= 321–31|doi=10.1111/j.1439-0469.2005.00326.x|url=http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2005/28.2005.pdf}}

NNPM NAN 41-646. Almost complete left tarsometatarsus. Probably a prehistoric hobby, perhaps less specialized for bird hunting: Sobolev, D.V. (2003): [http://www.v-zool.kiev.ua/pdfs/2003/6/11.pdf Новый вид плиоценового сокола (Falconiformes, Falconidae)] [A new species of Pliocene falcon (Falconiformes, Falconidae)] Vestnik zoologii 37 (6): 85–87. [Russian with English abstract]

White, Clayton M.; Olsen, Penny D. & Kiff, Lloyd F. (1994): Family Falconidae. In: del Hoyo, Josep; Elliott, Andrew & Sargatal, Jordi (editors): Handbook of Birds of the World, Volume 2 (New World Vultures to Guineafowl): 216–75, plates 24–28. Lynx Edicions, Barcelona. {{ISBN|84-87334-15-6}}

Wink, Michael; Seibold, I.; Lotfikhah, F. & Bednarek, W. (1998): [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/1998/31.%201998.pdf Molecular systematics of holarctic raptors (Order Falconiformes)]. In: Chancellor, R.D., Meyburg, B.-U. & Ferrero, J.J. (eds.): Holarctic Birds of Prey: 29–48. Adenex & WWGBP

Wink, Michael & Sauer-Gürth, Hedi (2000): [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2000/29.%202000.pdf Advances in the molecular systematics of African raptors]. In: Chancellor, R.D. & Meyburg, B.-U. (eds): Raptors at Risk: 135–47. WWGBP/Hancock House, Berlin/Blaine.

Wink, Michael; Sauer-Gürth, Hedi; Ellis, David & Kenward, Robert (2004): [http://www.uni-heidelberg.de/institute/fak14/ipmb/phazb/pubwink/2004/27.2004.pdf Phylogenetic relationships in the Hierofalco complex (Saker-, Gyr-, Lanner-, Laggar Falcon)]. In: Chancellor, R.D. & Meyburg, B.-U. (eds.): Raptors Worldwide: 499–504. WWGBP, Berlin

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• {{ cite journal | last1=Fuchs | first1=J. | last2=Johnson | first2=J.A. | last3=Mindell | first3=D.P. | year=2015 | title=Rapid diversification of falcons (Aves: Falconidae) due to expansion of open habitats in the Late Miocene | journal=Molecular Phylogenetics and Evolution | volume=82 | pages=166–182 | doi=10.1016/j.ympev.2014.08.010 | pmid=25256056 | bibcode=2015MolPE..82..166F }}

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{{Commons}}

{{wikispecies|Falco|Falcon}}

{{Wiktionary}}

• [http://ibc.lynxeds.com/family/falcons-caracaras-falconidae Falconidae videos] on the Internet Bird Collection, ibc.lynxeds.com
• [https://www.raptorresource.org/ The Raptor Resource Project] – Peregrine, owl, eagle and osprey cams, facts, and other resources, raptorresource.org

• {{Cite NIE|wstitle=Falcon |year=1905 |short=x}}

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Category:Extant Miocene first appearances

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