Tringa
Encyclopedia
Tringa is a genus
of wader
s, containing the shanks and tattlers
. They are mainly freshwater birds, often with brightly coloured legs as reflected in the English names of six species, as well as the specific names of two of these and the Green Sandpiper
. They are typically associated with northern hemisphere temperate regions for breeding. Some of this group — notably the Green Sandpiper — nest in trees, using the old nests of other birds, usually thrushes
.
The Willet
and the tattlers
have recently been found to belong in Tringa; these genus changes were formally adopted by the American Ornithologists' Union
in 2006.
The present genus in the old, more limited sense was even further subdivided into Tringa proper and Totanus
, either as subgenera or as full genera. The available DNA sequence
data suggests however that neither of these is monophyletic and that the latter simply lumps together a number of more of less closely related apomorphic species. Therefore is seems unwarranted to recognize Totanus even as a subgenus for the time being.
The shanks' and tattlers' closest relatives are sandpipers of the genera Actitis
and Xenus. Together with these, they are related to the phalaropes, as well as the turnstone
s and calidrid
s. The large genus
Tringa and the two very small genera which are most closely related form a phylogeny similar to the situation found in many other shorebird lineages such as calidrid
s, snipe
s and woodcock
s, or gull
s.
The same study has indicated that some morphological
characters such as details of the furcula
and pelvis
have evolved convergently
and are no indicators of close relationship. Similarly, the leg/foot color wildly varies between close relatives, with the Spotted Redshank
, the Greater Yellowlegs
, and the Common Greenshank for example being more closely related among each other than to any other species in the group; the ancestral coloration of the legs and feet was fairly certainly drab buffish as in e.g. the Green Sandpiper
. On the other hand, the molecular phylogeny reveals that the general habitus and size as well as the overall plumage pattern are good indicators of an evolutionary relationship in this group.
The Spotted Greenshank
, a rare and endangered species
, was not available for molecular analyses. It is fairly aberrant and was formerly placed in the monotypic
genus Pseudototanus. It appears closest overall to the semipalmata-flavipes and the stagnatilis-totanus-glareola groups, though it also has some similarities to the Greater Yellowlegs
and Common Greenshank.
shanks are known since the Miocene
, possibly even since the Eo-
/Oligocene
some 33-30 million years ago (mya) which would be far earlier than most extant genera of birds. However, it is uncertain whether Tringa edwardsi indeed belongs into the present-day genus or is a distinct, ancestral form. The time of the Tringa-Actitis
-Xenus-Phalaropus divergence has been tentatively dated at 22 mya, the beginning of the Miocene; even if the dating is largely conjectural, it suggests that T. edwardsi does indeed not belong into the modern genus. Molecular dating - which is not too reliable however - indicates that the diversification into the known lineages occurred between 20 and 5 mya. The fossil record contains species formerly separated in Totanus from the Early Miocene onwards. Although these are usually known from very scant remains, the fact that apparently apomorphic Tringa as well as a putative phalarope are known from about 23-22 mya indicates that the shank-phalarope group had already diverged into the modern genera by the start of the Miocene. The biogeography
of living and fossil species - notably the rarity of the latter in well-researched North America
n sites - seems to suggest that Tringa originated in Eurasia. Time and place neatly coincide with the disappearance of the last vestiges of the Turgai Sea
, and this process may well have been a major factor in the separation of the genera in the shank-phalarope clade
. Still, scolopacids are very similar osteologically, and many of the early fossils of presumed shanks require revaluation.
"Tringa" hoffmanni is now in Ludiortyx
. While its relationships are disputed, it was not a charadriiform.
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
of wader
Wader
Waders, called shorebirds in North America , are members of the order Charadriiformes, excluding the more marine web-footed seabird groups. The latter are the skuas , gulls , terns , skimmers , and auks...
s, containing the shanks and tattlers
Tattler (bird)
The tattlers are the two very similar bird species in the shorebird genus Tringa. They formerly had their own genus, Heteroscelus. The old genus name means "different leg" in Greek, referring to the leg scales that differentiate the tattlers from their close relatives, the shanks.The species are:*...
. They are mainly freshwater birds, often with brightly coloured legs as reflected in the English names of six species, as well as the specific names of two of these and the Green Sandpiper
Green Sandpiper
The Green Sandpiper is a small wader of the Old World. It represents an ancient lineage of the genus Tringa; its only close living relative is the Solitary Sandpiper . They both have brown wings with little light dots and a delicate but contrasting neck and chest pattern...
. They are typically associated with northern hemisphere temperate regions for breeding. Some of this group — notably the Green Sandpiper — nest in trees, using the old nests of other birds, usually thrushes
Thrush (bird)
The thrushes, family Turdidae, are a group of passerine birds that occur worldwide.-Characteristics:Thrushes are plump, soft-plumaged, small to medium-sized birds, inhabiting wooded areas, and often feed on the ground or eat small fruit. The smallest thrush may be the Forest Rock-thrush, at and...
.
The Willet
Willet
The Willet, Tringa semipalmata , is a large shorebird in the sandpiper family. It is a good-sized and stout scolopacid, the largest of the shanks...
and the tattlers
Tattler (bird)
The tattlers are the two very similar bird species in the shorebird genus Tringa. They formerly had their own genus, Heteroscelus. The old genus name means "different leg" in Greek, referring to the leg scales that differentiate the tattlers from their close relatives, the shanks.The species are:*...
have recently been found to belong in Tringa; these genus changes were formally adopted by the American Ornithologists' Union
American Ornithologists' Union
The American Ornithologists' Union is an ornithological organization in the USA. Unlike the National Audubon Society, its members are primarily professional ornithologists rather than amateur birders...
in 2006.
The present genus in the old, more limited sense was even further subdivided into Tringa proper and Totanus
Totanus
Totanus Bechstein, 1803 is a generic name previously applied to various waders or shorebirds, now subsumed within Tringa. Created by Bechstein, it derives from the species name for the Common Redshank, described by Linnaeus in 1758 as Scolopax totanus, from “totano”, the Italian name for the...
, either as subgenera or as full genera. The available DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
data suggests however that neither of these is monophyletic and that the latter simply lumps together a number of more of less closely related apomorphic species. Therefore is seems unwarranted to recognize Totanus even as a subgenus for the time being.
Living species
These are listed in systematic sequence:- Green SandpiperGreen SandpiperThe Green Sandpiper is a small wader of the Old World. It represents an ancient lineage of the genus Tringa; its only close living relative is the Solitary Sandpiper . They both have brown wings with little light dots and a delicate but contrasting neck and chest pattern...
, Tringa ochropus - Solitary SandpiperSolitary SandpiperThe Solitary Sandpiper is a small wader .-Description:This species measures long, with a wingspan up to and a body mass of . It is a dumpy wader with a dark green back, greyish head and breast and otherwise white underparts. It is obvious in flight, with wings dark above and below, and a dark...
, Tringa solitaria - Grey-tailed TattlerGrey-tailed TattlerThe Grey-tailed Tattler, Tringa brevipes , is a small shorebird.- Description :...
, Tringa brevipes - formerly Heteroscelus brevipes - Wandering TattlerWandering TattlerThe Wandering Tattler, Tringa incana , is a medium-sized wading bird. It is similar in appearance to the closely related Gray-tailed Tattler, T. brevipes...
, Tringa incana - formerly Heteroscelus incanus - Spotted RedshankSpotted RedshankThe Spotted Redshank, Tringa erythropus, is a wader in the large bird family Scolopacidae. It breeds across northern Scandinavia and northern Asia and migrates south to the Mediterranean, the southern British Isles, France, tropical Africa, and tropical Asia for the winter...
, Tringa erythropus - Greater YellowlegsGreater YellowlegsThe Greater Yellowlegs, Tringa melanoleuca, is a large North American shorebird, similar in appearance to the smaller Lesser Yellowlegs. Its closest relative, however, is the Greenshank, which together with the Spotted Redshank form a close-knit group...
, Tringa melanoleuca - Common Greenshank, Tringa nebularia
- WilletWilletThe Willet, Tringa semipalmata , is a large shorebird in the sandpiper family. It is a good-sized and stout scolopacid, the largest of the shanks...
, Tringa semipalmata - formerly Catoptrophorus semipalmatus - Lesser YellowlegsLesser YellowlegsThe Lesser Yellowlegs is a medium-sized shorebird similar in appearance to the larger Greater Yellowlegs. It is not closely related to this bird, however, but instead to the much larger and quite dissimilar Willet; merely the fine, clear and dense pattern of the neck shown in breeding plumage...
, Tringa flavipes - Nordmann's Greenshank, Tringa guttifer
- Marsh SandpiperMarsh SandpiperThe Marsh Sandpiper, Tringa stagnatilis, is a small wader. It is a rather small shank, and breeds in open grassy steppe and taiga wetlands from easternmost Europe to central Asia....
, Tringa stagnatilis - Common RedshankCommon RedshankThe Common Redshank or simply Redshank is an Eurasian wader in the large family Scolopacidae.- Description and systematics :...
, Tringa totanus - Wood SandpiperWood SandpiperThe Wood Sandpiper, Tringa glareola, is a small wader. This Eurasian species is the smallest of the shanks, which are mid-sized long-legged waders of the family Scolopacidae.- Description and systematics :...
, Tringa glareola
Systematics and evolution
Actitis
Actitis is a small genus of waders, comprising just two very similar bird species:* Common Sandpiper, Actitis hypoleucos, of Eurasia* Spotted Sandpiper, Actitis macularia of North America...
and Xenus. Together with these, they are related to the phalaropes, as well as the turnstone
Turnstone
Turnstones are the bird species in the genus Arenaria in the family Scolopacidae. They are closely related to calidrid sandpipers and might be considered members of the tribe Calidriini....
s and calidrid
Calidrid
The calidrids or typical waders are a group of Arctic-breeding, strongly migratory wading birds. These birds form huge mixed flocks on coasts and estuaries in winter. They are the typical "sandpipers", small to medium-sized, long-winged and relatively short-billed.Their bills have sensitive tips...
s. The large genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
Tringa and the two very small genera which are most closely related form a phylogeny similar to the situation found in many other shorebird lineages such as calidrid
Calidrid
The calidrids or typical waders are a group of Arctic-breeding, strongly migratory wading birds. These birds form huge mixed flocks on coasts and estuaries in winter. They are the typical "sandpipers", small to medium-sized, long-winged and relatively short-billed.Their bills have sensitive tips...
s, snipe
Snipe
A snipe is any of about 25 wading bird species in three genera in the family Scolopacidae. They are characterized by a very long, slender bill and crypsis plumage. The Gallinago snipes have a nearly worldwide distribution, the Lymnocryptes Jack Snipe is restricted to Asia and Europe and the...
s and woodcock
Woodcock
The woodcocks are a group of seven or eight very similar living species of wading birds in the genus Scolopax. Only two woodcocks are widespread, the others being localized island endemics. Most are found in the Northern Hemisphere but a few range into Wallacea...
s, or gull
Gull
Gulls are birds in the family Laridae. They are most closely related to the terns and only distantly related to auks, skimmers, and more distantly to the waders...
s.
The same study has indicated that some morphological
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....
characters such as details of the furcula
Furcula
The ' is a forked bone found in birds, formed by the fusion of the two clavicles. In birds, its function is the strengthening of the thoracic skeleton to withstand the rigors of flight....
and pelvis
Pelvis
In human anatomy, the pelvis is the lower part of the trunk, between the abdomen and the lower limbs .The pelvis includes several structures:...
have evolved convergently
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
and are no indicators of close relationship. Similarly, the leg/foot color wildly varies between close relatives, with the Spotted Redshank
Spotted Redshank
The Spotted Redshank, Tringa erythropus, is a wader in the large bird family Scolopacidae. It breeds across northern Scandinavia and northern Asia and migrates south to the Mediterranean, the southern British Isles, France, tropical Africa, and tropical Asia for the winter...
, the Greater Yellowlegs
Greater Yellowlegs
The Greater Yellowlegs, Tringa melanoleuca, is a large North American shorebird, similar in appearance to the smaller Lesser Yellowlegs. Its closest relative, however, is the Greenshank, which together with the Spotted Redshank form a close-knit group...
, and the Common Greenshank for example being more closely related among each other than to any other species in the group; the ancestral coloration of the legs and feet was fairly certainly drab buffish as in e.g. the Green Sandpiper
Green Sandpiper
The Green Sandpiper is a small wader of the Old World. It represents an ancient lineage of the genus Tringa; its only close living relative is the Solitary Sandpiper . They both have brown wings with little light dots and a delicate but contrasting neck and chest pattern...
. On the other hand, the molecular phylogeny reveals that the general habitus and size as well as the overall plumage pattern are good indicators of an evolutionary relationship in this group.
The Spotted Greenshank
Spotted Greenshank
The Nordmann's Greenshank or Spotted Greenshank Tringa guttifer is a wader in the large family Scolopacidae, the typical waders.It is a medium-sized sandpiper, 29-32 cm long, with a slightly upturned, bicoloured bill and shortish yellow legs...
, a rare and endangered species
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...
, was not available for molecular analyses. It is fairly aberrant and was formerly placed in the monotypic
Monotypic
In biology, a monotypic taxon is a taxonomic group with only one biological type. The term's usage differs slightly between botany and zoology. The term monotypic has a separate use in conservation biology, monotypic habitat, regarding species habitat conversion eliminating biodiversity and...
genus Pseudototanus. It appears closest overall to the semipalmata-flavipes and the stagnatilis-totanus-glareola groups, though it also has some similarities to the Greater Yellowlegs
Greater Yellowlegs
The Greater Yellowlegs, Tringa melanoleuca, is a large North American shorebird, similar in appearance to the smaller Lesser Yellowlegs. Its closest relative, however, is the Greenshank, which together with the Spotted Redshank form a close-knit group...
and Common Greenshank.
Fossil record
FossilFossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
shanks are known since the Miocene
Miocene
The Miocene is a geological epoch of the Neogene Period and extends from about . The Miocene was named by Sir Charles Lyell. Its name comes from the Greek words and and means "less recent" because it has 18% fewer modern sea invertebrates than the Pliocene. The Miocene follows the Oligocene...
, possibly even since the Eo-
Eocene
The Eocene Epoch, lasting from about 56 to 34 million years ago , is a major division of the geologic timescale and the second epoch of the Paleogene Period in the Cenozoic Era. The Eocene spans the time from the end of the Palaeocene Epoch to the beginning of the Oligocene Epoch. The start of the...
/Oligocene
Oligocene
The Oligocene is a geologic epoch of the Paleogene Period and extends from about 34 million to 23 million years before the present . As with other older geologic periods, the rock beds that define the period are well identified but the exact dates of the start and end of the period are slightly...
some 33-30 million years ago (mya) which would be far earlier than most extant genera of birds. However, it is uncertain whether Tringa edwardsi indeed belongs into the present-day genus or is a distinct, ancestral form. The time of the Tringa-Actitis
Actitis
Actitis is a small genus of waders, comprising just two very similar bird species:* Common Sandpiper, Actitis hypoleucos, of Eurasia* Spotted Sandpiper, Actitis macularia of North America...
-Xenus-Phalaropus divergence has been tentatively dated at 22 mya, the beginning of the Miocene; even if the dating is largely conjectural, it suggests that T. edwardsi does indeed not belong into the modern genus. Molecular dating - which is not too reliable however - indicates that the diversification into the known lineages occurred between 20 and 5 mya. The fossil record contains species formerly separated in Totanus from the Early Miocene onwards. Although these are usually known from very scant remains, the fact that apparently apomorphic Tringa as well as a putative phalarope are known from about 23-22 mya indicates that the shank-phalarope group had already diverged into the modern genera by the start of the Miocene. The biogeography
Biogeography
Biogeography is the study of the distribution of species , organisms, and ecosystems in space and through geological time. Organisms and biological communities vary in a highly regular fashion along geographic gradients of latitude, elevation, isolation and habitat area...
of living and fossil species - notably the rarity of the latter in well-researched North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
n sites - seems to suggest that Tringa originated in Eurasia. Time and place neatly coincide with the disappearance of the last vestiges of the Turgai Sea
Turgai Sea
The Turgai Sea or Turgai Strait, also known as the West Siberian Sea, was a large shallow body of salt water of the Mesozoic and Cenozoic Eras...
, and this process may well have been a major factor in the separation of the genera in the shank-phalarope clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
. Still, scolopacids are very similar osteologically, and many of the early fossils of presumed shanks require revaluation.
- ?Tringa edwardsi (Quercy Late Eocene/Early Oligocene of Mouillac, France)
- ?Tringa gracilis (Early Miocene of WC Europe) - calidridCalidridThe calidrids or typical waders are a group of Arctic-breeding, strongly migratory wading birds. These birds form huge mixed flocks on coasts and estuaries in winter. They are the typical "sandpipers", small to medium-sized, long-winged and relatively short-billed.Their bills have sensitive tips...
? - ?Tringa lartetianus (Early Miocene of Saint-Gérand-le-Puy, France)
- Tringa spp. (Early Miocene of Ravolzhausen, Germany - Early Pleistocene of Europe)
- ?Tringa grivensis (Middle Miocene of Grive-Saint-Alban, France)
- ?Tringa majori (Middle Miocene of Grive-Saint-Alban, France)
- ?Tringa minor (Middle Miocene of Grive-Saint-Alban, France) - includes "Erolia" ennouchii; calidriid?
- ?Tringa grigorescui (Middle Miocene of Ciobăniţa, Romania)
- ?Tringa scarabellii (Late Miocene of Senigallia, Italy)
- Tringa sp. 1 (Late Miocene/Early Pliocene of Lee Creek Mine, USA)
- Tringa sp. 2 (Late Miocene/Early Pliocene of Lee Creek Mine, USA)
- ?Tringa numenioides (Early Pliocene of Odessa, Ukraine)
- Tringa antiqua (Late Pliocene of Meade County, USA)
- Tringa ameghini (Late Pleistocene of Talara Tar Seeps, Peru)
"Tringa" hoffmanni is now in Ludiortyx
Ludiortyx
Ludiortyx is a bird genus from the Late Eocene. Its remains have been found in the Montmartre Formation at the Montmartre . A single species is accepted, Ludiortyx hoffmanni....
. While its relationships are disputed, it was not a charadriiform.