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Falcons () are
birds of prey Birds of prey or predatory birds, also known as raptors, are hypercarnivorous bird species that actively hunt and feed on other vertebrates (mainly mammals, reptiles and other smaller birds). In addition to speed and strength, these predat ...
in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Falco'', which includes about 40 species. Falcons are widely distributed on all continents of the world except
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
, though closely related raptors did occur there in the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
. Adult falcons have thin, tapered wings, which enable them to fly at high speed and change direction rapidly. Fledgling falcons, in their first year of flying, have longer flight feathers, which make their configuration more like that of a general-purpose bird such as a broad wing. This makes flying easier while learning the exceptional skills required to be effective hunters as adults. The falcons are the largest genus in the Falconinae subfamily of Falconidae, which itself also includes another subfamily comprising caracaras and a few other species. All these birds kill with their beaks, using a tomial "tooth" on the side of their beaks—unlike the hawks,
eagle Eagle is the common name for many large birds of prey of the family Accipitridae. Eagles belong to several groups of genera, some of which are closely related. Most of the 68 species of eagle are from Eurasia and Africa. Outside this area, j ...
s, and other birds of prey in the
Accipitridae The Accipitridae is one of the three families within the order Accipitriformes, and is a family of small to large birds with strongly hooked bills and variable morphology based on diet. They feed on a range of prey items from insects to medium-s ...
, which use their feet. The largest falcon is the
gyrfalcon The gyrfalcon ( or ) (), the largest of the falcon species, is a bird of prey. The abbreviation gyr is also used. It breeds on Arctic coasts and tundra, and the islands of northern North America and the Eurosiberian region. It is mainly a resid ...
at up to 65 cm in length. The smallest falcon species is the
Pygmy falcon The pygmy falcon (''Polihierax semitorquatus'') or African pygmy falcon is a bird species native to eastern and southern Africa. It is the smallest bird of prey on the continent. Description Adult pygmy falcons are white below and on the face, ...
which measures just 20 cm. As with hawks and owls, falcons exhibit
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
, with the females typically larger than the males, thus allowing a wider range of prey species. Some small falcons with long, narrow wings are called "
hobbies A hobby is considered to be a regular activity that is done for enjoyment, typically during one's leisure time. Hobbies include collecting themed items and objects, engaging in creative and artistic pursuits, playing sports, or pursuing oth ...
" and some which hover while hunting are called "
kestrel The term kestrel (from french: crécerelle, derivative from , i.e. ratchet) is the common name given to several species of predatory birds from the falcon genus ''Falco''. Kestrels are most easily distinguished by their typical hunting behaviou ...
s". As is the case with many birds of prey, falcons have exceptional powers of vision; the visual acuity of one species has been measured at 2.6 times that of a normal human. Peregrine falcons have been recorded diving at speeds of , making them the fastest-moving creatures on Earth; the fastest recorded dive attained a vertical speed of .


Taxonomy

The
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Falco'' was introduced in 1758 by the Swedish naturalist
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
in the tenth edition of his '' Systema Naturae''. The
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
is the Merlin (''Falco columbarius''). The genus name ''Falco'' is
Late Latin Late Latin ( la, Latinitas serior) is the scholarly name for the form of Literary Latin of late antiquity.Roberts (1996), p. 537. English dictionary definitions of Late Latin date this period from the , and continuing into the 7th century in t ...
meaning a "falcon" from ''falx'', ''falcis'', meaning "a sickle", referring to the claws of the bird. In
Middle English Middle English (abbreviated to ME) is a form of the English language that was spoken after the Norman conquest of 1066, until the late 15th century. The English language underwent distinct variations and developments following the Old English ...
and
Old French Old French (, , ; Modern French: ) was the language spoken in most of the northern half of France from approximately the 8th to the 14th centuries. Rather than a unified language, Old French was a linkage of Romance dialects, mutually intellig ...
, the title ''faucon'' refers generically to several captive raptor species. The traditional term for a male falcon is ''tercel'' (
British spelling Despite the various English dialects spoken from country to country and within different regions of the same country, there are only slight regional variations in English orthography, the two most notable variations being British and American ...
) or ''tiercel'' (
American spelling Despite the various English dialects spoken from country to country and within different regions of the same country, there are only slight regional variations in English orthography, the two most notable variations being British and American ...
), from the Latin ( 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 ( fro, tiercelet). A falcon chick, especially one reared for falconry, still in its downy stage, is known as an ''eyas'' (sometimes spelled ''eyass''). The word arose by mistaken division of
Old French Old French (, , ; Modern French: ) was the language spoken in most of the northern half of France from approximately the 8th to the 14th centuries. Rather than a unified language, Old French was a linkage of Romance dialects, mutually intellig ...
, from Latin presumed (nestling) from (
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materi ...
). The technique of hunting with trained captive birds of prey is known as falconry. Compared to other birds of prey, the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
record of the falcons is not well distributed in time. The oldest fossils tentatively assigned to this genus are from the Late
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
, less than 10 million years ago. This coincides with a period in which many modern genera of birds became recognizable in the fossil record. The falcon lineage may, however, be somewhat older than this, and given the distribution of fossil and living ''Falco''
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, is 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 Parrots, also known as psittacines (), are birds of the roughly 398 species in 92 genera comprising the order Psittaciformes (), found mostly in tropical and subtropical regions. The order is subdivided into three superfamilies: the Psittacoide ...
and
songbirds A songbird is a bird belonging to the suborder Passeri of the perching birds (Passeriformes). Another name that is sometimes seen as the scientific or vernacular name is Oscines, from Latin ''oscen'', "songbird". The Passeriformes contains 5 ...
.


Overview

Falcons are roughly divisible into three or four groups. The first contains the
kestrel The term kestrel (from french: crécerelle, derivative from , i.e. ratchet) is the common name given to several species of predatory birds from the falcon genus ''Falco''. Kestrels are most easily distinguished by their typical hunting behaviou ...
s (probably excepting the
American kestrel The American kestrel (''Falco sparverius''), also called the sparrow hawk, is the smallest and most common falcon in North America. It has a roughly two-to-one range in size over subspecies and sex, varying in size from about the weight of ...
); 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. Kestrels feed chiefly on
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () (chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with c ...
s and
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s of appropriate size, such as
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are n ...
s, reptiles, or
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...
s. 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
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary relatedness,For example, tail colour in the common and
lesser kestrel The lesser kestrel (''Falco naumanni'') is a small falcon. This species breeds from the Mediterranean across Afghanistan and Central Asia, to China and Mongolia. It is a summer migrant, wintering in Africa and Pakistan and sometimes even to Indi ...
s is absolutely identical, yet they do not seem closely related.
However, the
fox Foxes are small to medium-sized, omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull, upright, triangular ears, a pointed, slightly upturned snout, and a long bushy tail (or ''brush''). Twelve sp ...
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. The tails of the large falcons are quite uniformly dark grey with inconspicuous black banding and small, white tips, though this is probably
plesiomorph In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, a ...
ic. 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 hierofalcons (literally, "hawk-falcons"). They represent taxa with, usually, more
phaeomelanin Melanin (; from el, μέλας, melas, black, dark) is a broad term for a group of natural pigments found in most organisms. Eumelanin is produced through a multistage chemical process known as melanogenesis, where the oxidation of the amino ...
s, 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 Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
cytochrome ''b''
sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is calle ...
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 The Gelasian is an age in the international geologic timescale or a stage in chronostratigraphy, being the earliest or lowest subdivision of the Quaternary Period/System and Pleistocene Epoch/Series. It spans the time between 2.58 Ma (million ye ...
, 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
apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
ies. Other studies have confirmed that the hierofalcons 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 hierofalcons are basal among living falcons. The discovery of a NUMT proved this earlier theory erroneous. In reality, the hierofalcons 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 Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines * Morphology (archaeology), study of the shapes or forms of artifacts * Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies ...
of the syrinx, which contributes well to resolving the overall
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
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 hierofalcons and the more distant prairie falcon (which was sometimes placed with the hierofalcons, 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 An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. Radiations may affect one clade or many, and be rapid ...
seemingly was a
Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...
one that originated possibly around central Eurasia or in (northern) Africa. One or several lineages were present in North America by the
Early Pliocene Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...
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 The aplomado falcon (''Falco femoralis'') is a medium-sized falcon of the Americas. The species' largest contiguous range is in South America, but not in the deep interior Amazon Basin. It was long known as ''Falco fusco-coerulescens'' or ''Fal ...
lineage—can be quite confidently placed from the
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
-
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Zanclean The Zanclean is the lowest stage or earliest age on the geologic time scale of the Pliocene. It spans the time between 5.332 ± 0.005 Ma (million years ago) and 3.6 ± 0.005 Ma. It is preceded by the Messinian Age of the Miocene Epoch, and ...
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.


Species

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


Extinct species

*
Réunion kestrel The Réunion kestrel (''Falco duboisi'') is an extinct bird of prey which belongs to the falcon family. It inhabited the Mascarene island of Réunion and was part of the Western Indian Ocean radiation of kestrels. Known from subfossil bones ...
, ''Falco duboisi'' – extinct (about 1700)


Fossil record

*''Falco medius'' (Late Miocene of Cherevichnyi, Ukraine) IZAN 45-4033: left
carpometacarpus The carpometacarpus is a bone found in the hands of birds. It results from the fusion of the carpal and metacarpal bone, and is essentially a single fused bone between the wrist and the knuckles. It is a smallish bone in most birds, generally flat ...
. 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) *''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) *'' Falco antiquus'' (Middle Pleistocene of Noailles, France and possibly Horvőlgy, Hungary)Supposedly a
saker falcon The saker falcon (''Falco cherrug'') is a large species of falcon. This species breeds from central Europe eastwards across the Palearctic to Manchuria. It is mainly migratory except in the southernmost parts of its range, wintering in Ethiopia ...
paleosubspecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
(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) 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. . 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 The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
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 A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
of the
yellow-headed caracara The yellow-headed caracara (''Daptrius chimachima'') is a bird of prey in the family Falconidae. It is found in tropical and subtropical South America and the southern portion of Central America. Unlike the falcons in the same family, the caraca ...
(''Milvago chimachima'').


See also

*
Abu Dhabi Falcon Hospital The Abu Dhabi Falcon Hospital (ADFH) is the first public medical institution exclusively for falcons in the United Arab Emirates. Established by Environment Agency – Abu Dhabi and opened on 3 October 1999, Abu Dhabi Falcon Hospital has become ...
* Falconry * Ra * Horus *
Khonsu Khonsu ( egy, ḫnsw; also transliterated Chonsu, Khensu, Khons, Chons or Khonshu; cop, Ϣⲟⲛⲥ, Shons) is the ancient Egyptian god of the Moon. His name means "traveller", and this may relate to the perceived nightly travel of the Moon ...


Notes


References


Further reading

*


External links


Falconidae videos
on the Internet Bird Collection, ibc.lynxeds.com
The Raptor Resource Project
– Peregrine, owl, eagle and osprey cams, facts, and other resources, raptorresource.org * {{Authority control Birds of prey Extant Miocene first appearances Falconry Taxa named by Carl Linnaeus