Bird feet and legs

The anatomy of bird legs and feet is diverse, encompassing many accommodations to perform a wide variety of functions.[1]

African jacana. Extremely long toes[1] and claws help distribute the jacana's weight over a wide area to allow it to walk on floating leaves.[2]

Most birds are classified as digitigrade animals, meaning they walk on their toes, rather than the entire foot.[3][4] Some of the lower bones of the foot (the distals and most of the metatarsal) are fused to form the tarsometatarsus – a third segment of the leg, specific to birds.[5][6] The upper bones of the foot (proximals), in turn, are fused with the tibia to form the tibiotarsus, as over time the centralia disappeared.[7][6][4][8] The fibula also reduced.[5]

The legs are attached to a strong assembly consisting of the pelvic girdle extensively fused with the uniform spinal bone (also specific to birds) called the synsacrum, built from some of the fused bones.[8][9]

Bird left leg and pelvic girdle skeleton

Functions

Grey parrot grips the perch with zygodactyl feet.

Palmate feet – Chilean flamingo.

Totipalmate feet – blue-footed booby.

Western grebe presenting a lobate foot.

Lobate feet – a chick of the Eurasian coot.

The great crested grebe. The feet in loons[2] and grebes[2][7] are placed far at the rear of the body - a powerful accommodation to swimming underwater,[7] but a handicap for walking.

The snowshoe-like foot of the willow ptarmigan is an adaptation for walking on snow.[1]

Because avian forelimbs are wings, many forelimb functions are performed by the bill and hindlimbs.[10] It has been proposed that the hindlimbs are important in flight as accelerators when taking-off.[18][19] Some leg and foot functions, including conventional ones and those specific to birds, are:

Locomotion
- Walking, running (ostriches, grouse, wild turkeys) hopping, climbing (woodpeckers, nuthatches, treecreepers)[20]
- Swimming and steering underwater (ducks, grebes, loons)[3]
Perching (as on a branch) or clinging[3]
Carrying (like ospreys holding fish)[3]
Flight-related
- Serving probably as the primary take-off accelerator. In the common vampire bat, by contrast, the required force is generated by the wing.[18][19]
- Absorbing the shock of landing on a perch and on the water, becoming "water skis"[3]
Feeding and related
- Catching and killing prey in raptors (hawks, owls)[3]
- Holding (used like hands in parrots) and pulling apart food (with help from the bill)[3]
- Scratching the ground in search of food[2]
  - Double scratch: hopping forward and then backward using both feet to scratch (often towhees, sparrows, juncos)[2]
  - One-footed scratch (grouse, quails, wild turkeys, domestic chicken)[2]
Reproduction and related
- Cradling and turning eggs during incubation.[3] Birds lacking a brood patch incubate the eggs with their feet – grasping one or even two of them (gannets, boobies) or keeping them on the top surfaces of their feet (penguins under a pouch of belly skin, murres).[1]
- Courtship (sage grouse), including aerial courtship (bald eagles)[3]
- Building nests (with help from the bill)[1]
Preening and cleaning.[10] Sometimes birds use a special claw (for example, barn owls have a so-called "feather comb"). Some herons and nightjars use the claw for cleaning the head.[2]
Heat loss regulation (herons, gulls, giant petrels, storks, New World vultures, ducks, geese)[1][2]

Toe arrangements

Toe arrangement in a bird's right foot

Typical toe arrangements in birds are:

Anisodactyl: three toes in front (2, 3, 4), and one in back (1); in nearly all songbirds and most other perching birds.[4][20]
Zygodactyl: two toes in front (2, 3) and two in back (1, 4) – the outermost front toe (4) is reversed. The zygodactyl arrangement is a case of convergence, because it evolved in birds in different ways nine times.[1][10]
- In many perching birds – most woodpeckers and their allies, ospreys, owls, cuckoos, most parrots, mousebirds, some swifts and cuckoo rollers.[20][4]
  - Woodpeckers, when climbing, can rotate the outer rear digit (4) to the side in an ectropodactyl arrangement. Black-backed woodpeckers, Eurasian three-toed woodpeckers and American three-toed woodpeckers have three toes – the inner rear (1) is missing and the outer rear (4) points always backward and never rotates.[10]
  - Owls, ospreys and turacos can rotate the outer toe (4) back and forth.[10]
Heterodactyl: two toes in front (3, 4) and two in back (2, 1) – the inner front toe (2) is reversed; heterodactyl arrangement only exists in trogons.[20]
Syndactyl: three toes in front (2, 3, 4), one in back (1); the inner and middle (2, 3) are joined for much of their length.[2][1] Common in Coraciiformes, including kingfishers and hornbills.[7]
Pamprodactyl: two inner toes in front (2, 3), the two outer (1, 4) can rotate freely forward and backward. In mousebirds and some swifts. Some swifts move all four digits forward to use them as hooks to hang.[20]

The most common arrangement is the anisodactyl foot, and second among perching birds is the zygodactyl arrangement.[3][7][21]

Claws

All birds have claws at the end of the toes. The claws are typically curved and the radius of curvature tends to be greater as the bird is larger although they tend to be straighter in large ground dwelling birds such as ratites.[22] Some species (including nightjars, herons, frigatebirds, owls and pratincoles) have comb-like serrations on the claw of the middle toe that may aid in scratch preening.[23]

Webbing and lobation

Webbing and lobation in a bird's right foot

Palmations and lobes enable swimming or help walking on loose ground such as mud.[3] The webbed or palmated feet of birds can be categorized into several types:

Palmate: only the anterior digits (2–4) are joined by webbing. Found in ducks, geese and swans, gulls and terns, and other aquatic birds (auks, flamingos, fulmars, jaegers, loons, petrels, shearwaters and skimmers).[20][21] Diving ducks also have a lobed hind toe (1), and gulls, terns and allies have a reduced hind toe.[24]
Totipalmate: all four digits (1–4) are joined by webbing. Found in gannets and boobies, pelicans, cormorants, anhingas and frigatebirds. Some gannets have brightly colored feet used in display.[3][21]
Semipalmate: a small web between the anterior digits (2–4). Found in some plovers (Eurasian dotterels) and sandpipers (semipalmated sandpipers, stilt sandpipers, upland sandpipers, greater yellowlegs and willet), avocet, herons (only two toes), all grouse, and some domesticated breeds of chicken. Plovers and lapwings have a vestigial hind toe (1), and sandpipers and their allies have a reduced and raised hind toe barely touching the ground. The sanderling is the only sandpiper having 3 toes (tridactyl foot).[3]
Lobate: the anterior digits (2–4) are edged with lobes of skin. Lobes expand or contract when a bird swims. In grebes, coots, phalaropes, finfoots and some palmate-footed ducks on the hallux (1). Grebes have more webbing between the toes than coots and phalaropes.[20][4][21]

The palmate foot is most common.

Thermal regulation

Some birds like gulls, herons, ducks or geese can regulate their temperature through their feet.[1][2]

The arteries and veins intertwine in the legs, so heat can be transferred from arteries back to veins before reaching the feet. Such a mechanism is called countercurrent exchange. Gulls can open a shunt between these vessels, turning back the bloodstream above the foot, and constrict the vessels in the foot. This reduces heat loss by more than 90 percent. In gulls, the temperature of the base of the leg is 32 °C (89 °F), while that of the foot may be close to 0 °C (32 °F).[1]

However, for cooling, this heat-exchange network can be bypassed and blood-flow through the foot significantly increased (giant petrels). Some birds, also excrete onto their feet, increasing heat loss via evaporation (storks, New World vultures).[1]

References

Gill, Frank B. (2001). Ornithology (2md ed.). New York: W.H. Freeman and Company. ISBN 978-0-7167-2415-5.
Kochan, Jack B. (1994). Feet & Legs. Birds. Mechanicsburg: Stackpole Books. ISBN 978-0-8117-2515-6.CS1 maint: ref=harv (link)
Kochan 1994; Proctor 1993; Elphick 2001
Kowalska-Dyrcz, Alina (1990). "Entry: noga [leg]". In Busse, Przemysław (ed.). Ptaki [Birds]. Mały słownik zoologiczny [Small zoological dictionary] (in Polish). I (I ed.). Warsaw: Wiedza Powszechna. pp. 383–385. ISBN 978-83-214-0563-6.
Proctor 1993; Kowalska-Dyrcz 1990; Dobrowolski 1981
Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 205–208. ISBN 978-0-03-910284-5.
Proctor, Noble S.; Lynch, Patrick J. (1993). "Chapters: 6. Topography of the foot, 11. The pelvic girdle, and 12. The bones of the leg and foot Family". Manual of Ornithology. Avian Structure & Function. New Haven and London: Yale University Press. pp. 70–75, 140–141, 142–144. ISBN 978-0-300-07619-6.
Dobrowolski, Kazimierz A.; Klimaszewski, Sędzimir M.; Szelęgiewicz, Henryk (1981). "Chapters: Gromada: Ptaki - Aves: Układ kostny; Pas miednicowy i kończyna tylna [Class: Birds: The skeletal system; The pelvic girdle and the hindlimb]". Zoologia [Zoology] (in Polish) (4th ed.). Warsaw: Wydawnictwo Szkolne i Pedagogiczne. pp. 462–464, 469. ISBN 978-83-02-00608-1.
Kowalska-Dyrcz, Alina (1990). "Entry: synsakrum [synsacrum]". In Busse, Przemysław (ed.). Ptaki [Birds]. Mały słownik zoologiczny [Small zoological dictionary] (in Polish). II (I ed.). Warsaw: Wiedza Powszechna. p. 245. ISBN 978-83-214-0563-6.
Elphick, John B.; Dunning, JR., Jack B.; Sibley, David Allen (2001). National Audubon Society: The Sibley Guide to Bird Life & Behavior. New York: Alfred A. Knopf. ISBN 978-0-679-45123-5.
Munn, Philip W. (1 January 1894). "On the Birds of the Calcutta District". Ibis. 36 (1): 39–77. doi:10.1111/j.1474-919x.1894.tb01250.x. ISSN 1474-919X.
Chasen, F. N. (1923). "On The Heel-Pad in certain Malaysian Birds". Journal of the Malayan Branch of the Royal Asiatic Society. 1 (87): 237–246. JSTOR 41559544.
Wedel, Mathew J. (2003). "Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs" (PDF). Paleobiology. 29 (2): 243–255. doi:10.1666/0094-8373(2003)029<0243:vpasat>2.0.co;2.
Schorger, A. W. (September 1947). "The deep diving of the loon and old-squaw and its mechanism" (PDF). The Wilson Bulletin. 59 (3): 151–159.
Fastovsky, David E.; Weishampel, David B. (2005). The Evolution and Extinction of the Dinosaurs (2nd ed.). Cambridge, UK: Cambridge University Press. ISBN 978-0-521-81172-9.
Gier, H. T. (1952). "The air sacs of the loon" (PDF). The Auk. 69 (1): 40–49. doi:10.2307/4081291. JSTOR 4081291.
Bezuidenhout, A.J.; Groenewald, H.B.; Soley, J.T. (1999). "An anatomical study of the respiratory air sacs in ostriches" (PDF). Onderstepoort Journal of Veterinary Research. 66 (4): 317–325. PMID 10689704.
Earls, Kathleen D. (Feb 2000). "Kinematics and mechanics of ground take-off in the starling Sturnis vulgaris and the quail Coturnix coturnix" (PDF). The Journal of Experimental Biology. 203 (Pt 4): 725–739. PMID 10648214.
Whitfield, John (10 March 2000). "Off to a flying jump-start : Nature News". Nature. Nature Publishing Group. doi:10.1038/news000316-1. Retrieved 17 January 2014.
Gill 2001; Kochan 1994; Proctor 1993; Elphick 2001
Kalbe, Lothar (1983). "Besondere Formen für spezielle Aufgaben der Wassertiere [Special adaptations of aquatic animals to specific lifestyles]". Tierwelt am Wasser [Wildlife by the Water] (in German) (1st ed.). Leipzig-Jena-Berlin: Urania-Verlag. pp. 72–77.
Pike, A. V. L.; Maitland, D. P. (2004). "Scaling of bird claws". Journal of Zoology. 262: 73–81. doi:10.1017/S0952836903004382.
Stettenheim, Peter R. (August 2000). "The Integumentary Morphology of Modern Birds—An Overview". American Zoologist. 40 (4): 461–477. CiteSeerX 10.1.1.559.1172. doi:10.1668/0003-1569(2000)040[0461:timomb]2.0.co;2. ISSN 0003-1569.
Kochan 1994; Elphick 2001

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[Gill01-1] Gill, Frank B. (2001). Ornithology (2md ed.). New York: W.H. Freeman and Company. ISBN 978-0-7167-2415-5.

[Kochan94-2] Kochan, Jack B. (1994). Feet & Legs. Birds. Mechanicsburg: Stackpole Books. ISBN 978-0-8117-2515-6.CS1 maint: ref=harv (link)

[kochan-proctor-sibleybeh-3] Kochan 1994; Proctor 1993; Elphick 2001

[Dict90_1-4] Kowalska-Dyrcz, Alina (1990). "Entry: noga [leg]". In Busse, Przemysław (ed.). Ptaki [Birds]. Mały słownik zoologiczny [Small zoological dictionary] (in Polish). I (I ed.). Warsaw: Wiedza Powszechna. pp. 383–385. ISBN 978-83-214-0563-6.

[proctor-dict-dobrowolski-5] Proctor 1993; Kowalska-Dyrcz 1990; Dobrowolski 1981

[Romer77-6] Romer, Alfred Sherwood; Parsons, Thomas S. (1977). The Vertebrate Body. Philadelphia, PA: Holt-Saunders International. pp. 205–208. ISBN 978-0-03-910284-5.

[Proctor93-7] Proctor, Noble S.; Lynch, Patrick J. (1993). "Chapters: 6. Topography of the foot, 11. The pelvic girdle, and 12. The bones of the leg and foot Family". Manual of Ornithology. Avian Structure & Function. New Haven and London: Yale University Press. pp. 70–75, 140–141, 142–144. ISBN 978-0-300-07619-6.

[Dobrowolski81-8] Dobrowolski, Kazimierz A.; Klimaszewski, Sędzimir M.; Szelęgiewicz, Henryk (1981). "Chapters: Gromada: Ptaki - Aves: Układ kostny; Pas miednicowy i kończyna tylna [Class: Birds: The skeletal system; The pelvic girdle and the hindlimb]". Zoologia [Zoology] (in Polish) (4th ed.). Warsaw: Wydawnictwo Szkolne i Pedagogiczne. pp. 462–464, 469. ISBN 978-83-02-00608-1.

[Dict90_2-9] Kowalska-Dyrcz, Alina (1990). "Entry: synsakrum [synsacrum]". In Busse, Przemysław (ed.). Ptaki [Birds]. Mały słownik zoologiczny [Small zoological dictionary] (in Polish). II (I ed.). Warsaw: Wiedza Powszechna. p. 245. ISBN 978-83-214-0563-6.

[SibleyBeh01-10] Elphick, John B.; Dunning, JR., Jack B.; Sibley, David Allen (2001). National Audubon Society: The Sibley Guide to Bird Life & Behavior. New York: Alfred A. Knopf. ISBN 978-0-679-45123-5.

[11] Munn, Philip W. (1 January 1894). "On the Birds of the Calcutta District". Ibis. 36 (1): 39–77. doi:10.1111/j.1474-919x.1894.tb01250.x. ISSN 1474-919X.

[12] Chasen, F. N. (1923). "On The Heel-Pad in certain Malaysian Birds". Journal of the Malayan Branch of the Royal Asiatic Society. 1 (87): 237–246. JSTOR 41559544.

[Wedel03-13] Wedel, Mathew J. (2003). "Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs" (PDF). Paleobiology. 29 (2): 243–255. doi:10.1666/0094-8373(2003)029<0243:vpasat>2.0.co;2.

[Schorger47-14] Schorger, A. W. (September 1947). "The deep diving of the loon and old-squaw and its mechanism" (PDF). The Wilson Bulletin. 59 (3): 151–159.

[Fastovsky05-15] Fastovsky, David E.; Weishampel, David B. (2005). The Evolution and Extinction of the Dinosaurs (2nd ed.). Cambridge, UK: Cambridge University Press. ISBN 978-0-521-81172-9.

[Gier52-16] Gier, H. T. (1952). "The air sacs of the loon" (PDF). The Auk. 69 (1): 40–49. doi:10.2307/4081291. JSTOR 4081291.

[Bezuidenhout99-17] Bezuidenhout, A.J.; Groenewald, H.B.; Soley, J.T. (1999). "An anatomical study of the respiratory air sacs in ostriches" (PDF). Onderstepoort Journal of Veterinary Research. 66 (4): 317–325. PMID 10689704.

[Earls00-18] Earls, Kathleen D. (Feb 2000). "Kinematics and mechanics of ground take-off in the starling Sturnis vulgaris and the quail Coturnix coturnix" (PDF). The Journal of Experimental Biology. 203 (Pt 4): 725–739. PMID 10648214.

[Whitfield00-19] Whitfield, John (10 March 2000). "Off to a flying jump-start : Nature News". Nature. Nature Publishing Group. doi:10.1038/news000316-1. Retrieved 17 January 2014.

[gill-kochan-proctor-sibleybeh-20] Gill 2001; Kochan 1994; Proctor 1993; Elphick 2001

[Kalbe83-21] Kalbe, Lothar (1983). "Besondere Formen für spezielle Aufgaben der Wassertiere [Special adaptations of aquatic animals to specific lifestyles]". Tierwelt am Wasser [Wildlife by the Water] (in German) (1st ed.). Leipzig-Jena-Berlin: Urania-Verlag. pp. 72–77.

[22] Pike, A. V. L.; Maitland, D. P. (2004). "Scaling of bird claws". Journal of Zoology. 262: 73–81. doi:10.1017/S0952836903004382.

[Stettenheim-23] Stettenheim, Peter R. (August 2000). "The Integumentary Morphology of Modern Birds—An Overview". American Zoologist. 40 (4): 461–477. CiteSeerX 10.1.1.559.1172. doi:10.1668/0003-1569(2000)040[0461:timomb]2.0.co;2. ISSN 0003-1569.

[kochan-sibleybeh-24] Kochan 1994; Elphick 2001