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Falcons () are birds of prey in the genus ''Falco'', which includes about 40 species. Falcons are widely distributed on all continents of the world except Antarctica, though closely related raptors did occur there in the Eocene. 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 feather Flight feathers (''Pennae volatus'') are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges (), singular remex (), while those on the tail ...
s, 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 The falcons and caracaras are around 60 species of diurnal birds of prey that make up the family Falconidae (representing all extant species in the order Falconiformes). The family is divided into three subfamilies, Herpetotherinae, which inclu ...
, 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
hawk Hawks are bird of prey, birds of prey of the family Accipitridae. They are widely distributed and are found on all continents except Antarctica. * The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks and others. Th ...
s,
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, just ...
s, and other birds of prey in the Accipitridae, which use their feet. The largest falcon is the gyrfalcon at up to 65 cm in length. The smallest falcon species is the Pygmy falcon which measures just 20 cm. As with hawks and owls, falcons exhibit sexual dimorphism, 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" and some which hover while hunting are called " kestrels". 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 falcon The peregrine falcon (''Falco peregrinus''), also known as the peregrine, and historically as the duck hawk in North America, is a Cosmopolitan distribution, cosmopolitan bird of prey (Bird of prey, raptor) in the family (biology), family Falco ...
s 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 ''Falco'' was introduced in 1758 by the Swedish naturalist Carl Linnaeus in the tenth edition of his ''
Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nomen ...
''. The type species is the
Merlin Merlin ( cy, Myrddin, kw, Marzhin, br, Merzhin) is a mythical figure prominently featured in the legend of King Arthur and best known as a mage, with several other main roles. His usual depiction, based on an amalgamation of historic and le ...
(''Falco columbarius''). The genus name ''Falco'' is Late Latin meaning a "falcon" from ''falx'', ''falcis'', meaning "a sickle", referring to the claws of the bird. In Middle English and Old French, 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 Third or 3rd may refer to: Numbers * 3rd, the ordinal form of the cardinal number 3 * , a fraction of one third * Second#Sexagesimal divisions of calendar time and day, 1⁄60 of a ''second'', or 1⁄3600 of a ''minute'' Places * 3rd Street (d ...
) 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 Falconry is the hunting of wild animals in their natural state and habitat by means of a trained bird of prey. Small animals are hunted; squirrels and rabbits often fall prey to these birds. Two traditional terms are used to describe a person ...
, still in its downy stage, is known as an ''eyas'' (sometimes spelled ''eyass''). The word arose by mistaken division of Old French , from Latin presumed (nestling) from ( nest). The technique of hunting with trained captive birds of prey is known as
falconry Falconry is the hunting of wild animals in their natural state and habitat by means of a trained bird of prey. Small animals are hunted; squirrels and rabbits often fall prey to these birds. Two traditional terms are used to describe a person ...
. Compared to other birds of prey, the fossil record of the falcons is not well distributed in time. The oldest fossils tentatively assigned to this genus are from the Late Miocene, 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, 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 and songbirds.


Overview

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. Kestrels feed chiefly on terrestrial vertebrates and invertebrates of appropriate size, such as rodents,
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s, 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
morph Morph may refer to: Biology * Morph (zoology), a visual or behavioral difference between organisms of distinct populations in a species * Muller's morphs, a classification scheme for genetic mutations * "-morph", a suffix commonly used in tax ...
s), 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 Common may refer to: Places * Common, a townland in County Tyrone, Northern Ireland * Boston Common, a central public park in Boston, Massachusetts * Cambridge Common, common land area in Cambridge, Massachusetts * Clapham Common, originally com ...
and lesser kestrels 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 kestrel The greater kestrel (''Falco rupicoloides'') or white-eyed kestrel, is a bird of prey belonging to the falcon family Falconidae. It is one of the largest kestrels and is found in open country in southern and eastern Africa. Description The pl ...
s 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 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 The hierofalcons are four closely related species of falcon which make up the subgenus ''Hierofalco'': * Lanner falcon, ''Falco biarmicus'' * Laggar falcon, ''Falco jugger'' * Saker falcon, ''Falco cherrug'' * Gyrfalcon, ''Falco rusticolus'' Th ...
s (literally, "hawk-falcons"). They represent taxa with, usually, more phaeomelanins, which impart reddish or brown colors, and generally more strongly patterned plumage reminiscent of
hawk Hawks are bird of prey, birds of prey of the family Accipitridae. They are widely distributed and are found on all continents except Antarctica. * The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks and others. Th ...
s. 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
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s 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 A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
, 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 In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
group–and that
hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
is quite frequent at least in the larger falcon species. Initial studies of mtDNA cytochrome ''b'' sequence data suggested that the hierofalcons are
basal Basal or basilar is a term meaning ''base'', ''bottom'', or ''minimum''. Science * Basal (anatomy), an anatomical term of location for features associated with the base of an organism or structure * Basal (medicine), a minimal level that is nec ...
among living falcons. The discovery of a
NUMT NUMT, pronounced "new might," is an acronym for "''nu''clear ''m''i''t''ochondrial DNA" segment coined by evolutionary geneticist, Jose V. Lopez, which describes a transposition of any type of cytoplasmic mitochondrial DNA into the nuclear genome o ...
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 of the
syrinx In classical Greek mythology, Syrinx (Greek Σύριγξ) was a nymph and a follower of Artemis, known for her chastity. Pursued by the amorous god Pan, she ran to a river's edge and asked for assistance from the river nymphs. In answer, sh ...
, which contributes well to resolving the overall phylogeny of the
Falconidae The falcons and caracaras are around 60 species of diurnal birds of prey that make up the family Falconidae (representing all extant species in the order Falconiformes). The family is divided into three subfamilies, Herpetotherinae, which inclu ...
, 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
biogeographical Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
ly), as well as at least most of the "typical" hobbies, are confirmed to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
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 The Piacenzian is in the international geologic time scale the upper stage (stratigraphy), stage or latest age (geology), age of the Pliocene. It spans the time between 3.6 ± 0.005 year#SI prefix multipliers, Ma and 2.588 ± 0.005 Ma (million yea ...
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 and ...
, ''Falco duboisi'' –
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
(about 1700)


Fossil record

*''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) *''Falco'' sp. (Early Pliocene of Kansas) *''Falco'' sp. (Early Pliocene of Bulgaria – Early Pleistocene of Spain and Czech Republic)A
hierofalcon The hierofalcons are four closely related species of falcon which make up the subgenus ''Hierofalco'': * Lanner falcon, ''Falco biarmicus'' * Laggar falcon, ''Falco jugger'' * Saker falcon, ''Falco cherrug'' * Gyrfalcon, ''Falco rusticolus'' Th ...
(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 In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...
'' *''?Falco bakalovi'' (Late Pliocene of Varshets, Bulgaria) *''
Falco antiquus ''Falco antiquus'' is an extinct species of falcon lived during the Middle Pleistocene in southern Europe. It is thought to be the common ancestor of modern gyrfalcons The gyrfalcon ( or ) (), the largest of the falcon species, is a bird of p ...
'' (Middle Pleistocene of Noailles, France and possibly Horvőlgy, Hungary)Supposedly a saker falcon
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 The Eemian (also called the last interglacial, Sangamonian, Sangamonian Stage, Ipswichian, Mikulin, Kaydaky, penultimate,NOAA - Penultimate Interglacial Period http://www.ncdc.noaa.gov/global-warming/penultimate-interglacial-period Valdivia or Ri ...
origin of that species.
*
Cuban kestrel The Cuban kestrel (''Falco kurochkini'') was a species of small falcon in the family Falconidae that was formerly endemic to the island of Cuba. It was described from fossil remains from late Quaternary deposits from several sites throughout th ...
, ''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 A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
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 Columba or Colmcille; gd, Calum Cille; gv, Colum Keeilley; non, Kolban or at least partly reinterpreted as (7 December 521 – 9 June 597 AD) was an Irish abbot and missionary evangelist credited with spreading Christianity in what is toda ...
'', 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 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 (''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 Falconry is the hunting of wild animals in their natural state and habitat by means of a trained bird of prey. Small animals are hunted; squirrels and rabbits often fall prey to these birds. Two traditional terms are used to describe a person ...
* Ra *
Horus Horus or Heru, Hor, Har in Ancient Egyptian, is one of the most significant ancient Egyptian deities who served many functions, most notably as god of kingship and the sky. He was worshipped from at least the late prehistoric Egypt until the P ...
* Khonsu


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