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The Savannah sparrow (''Passerculus sandwichensis'') is a small
New World sparrow New World sparrows are a group of mainly New World passerine birds, forming the family Passerellidae. They are seed-eating birds with conical bills, brown or gray in color, and many species have distinctive head patterns. Although they share t ...
. It was the only member of the genus '' Passerculus'' and is typically the only widely accepted member. Comparison 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 ...
NADH dehydrogenase NADH dehydrogenase is an enzyme that converts nicotinamide adenine dinucleotide (NAD) from its reduced form (NADH) to its oxidized form (NAD+). Members of the NADH dehydrogenase family and analogues are commonly systematically named using the for ...
subunit 2 and 3
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 ...
s indicates that the Ipswich sparrow, formerly usually considered a valid species (as ''Passerculus princeps''), is a well-marked subspecies of the Savannah sparrow, whereas the southwestern large-billed sparrow should be recognized as a distinct species (''Passerculus rostratus''). The species name ''sandwichensis'' is Latin, of place, Sandwich, Unalaska, or Aleutians area, from which came the first subspecies, Aleutian Savannah sparrow, to be described. The common name honors Savannah, Georgia where Alexander Wilson discovered the species in 1811.


Distribution

This
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
breeds in
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S. ...
,
Canada Canada is a country in North America. Its ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, covering over , making it the world's second-largest country by tot ...
, northern, central and
Pacific The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contine ...
coastal United States,
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and Guatemala. The Pacific and Mexican breeders are resident, but other populations are migratory, wintering from the southern United States across
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
and the Caribbean to northern
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
. It is a very rare vagrant to western
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
.


Description

This species has a typically sparrow-like dark-streaked brown back, and whitish underparts with brown or blackish breast and flank streaking. It has whitish crown and supercilium stripes, sometimes with some yellow (more often near the
beak The beak, bill, or rostrum is an external anatomical structure found mostly in birds, but also in turtles, non-avian dinosaurs and a few mammals. A beak is used for eating, preening, manipulating objects, killing prey, fighting, probing for foo ...
). The cheeks are brown and the throat white. The flight feathers are blackish-brown with light brown or white border. The eyes are dark. The feet and legs are horn-colored, as is the lower part of the bill, with the upper part being dark grey. The Savannah sparrow is a very variable species, with numerous subspecies, several of which have been split as separate species at various times. The different forms vary principally in the darkness of the plumage. The variation generally follows
Gloger's rule Gloger's rule is an ecogeographical rule which states that within a species of endotherms, more heavily pigmented forms tend to be found in more humid environments, e.g. near the equator. It was named after the zoologist Constantin Wilhelm Lambe ...
, with Alaskan and interior races the palest, and southwestern coastal forms the darkest. There are some exceptions, though, most conspicuously in some island populations that presumably were strongly affected by founder effects. The general pattern of variation has a fairly clear divide, southwest of which the birds become notably darker; this agrees quite well with the limit between ''P. sandwichensis'' and ''P. (s.) rostratus''. Savannah sparrows show some variation in size across subspecies. The total length can range from , wingspan ranges from and body mass from . In the nominate subspecies, the body weight averages .''CRC Handbook of Avian Body Masses'' by John B. Dunning Jr. (Editor). CRC Press (1992), . The Savannah sparrows proper (see below) are very similar, and migrant birds can not usually be related to a breeding population with certainty. The resident or partially migratory subspecies are well distinguishable by size and, particularly between groups, coloration.


Behavior

These birds forage on the ground or in low bushes; particularly in winter they are also found in grazed low-growth grassland. They mainly eat seeds, but also eat insects in the breeding season. They are typically encountered as pairs or family groups in the breeding season, and assemble in flocks for the winter migration. The flight call is a thin ''seep''. ''
Sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
'', the Savannah sparrow is considered a
threatened species Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of '' critical depen ...
by the IUCN. The song is mixture of ''chirps'' and trills.


Systematics

Seventeen subspecies (including the large-billed sparrows) are currently recognized, though many are only described from wintering birds and much of the variation seems to be clinal. Four additional subspecies are no longer generally accepted. The complex is usually divided into several groups:


Savannah sparrows proper

All are migratory; wintering ranges overlap widely. * ''P. s. labradorius'', breeds in Newfoundland,
Labrador , nickname = "The Big Land" , etymology = , subdivision_type = Country , subdivision_name = Canada , subdivision_type1 = Province , subdivision_name1 ...
, and N
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...
* ''P. s. oblitus'', breeds in north
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central C ...
and
Manitoba , image_map = Manitoba in Canada 2.svg , map_alt = Map showing Manitoba's location in the centre of Southern Canada , Label_map = yes , coordinates = , capital = Winn ...
* ''P. s. savanna'' (eastern Savannah sparrow), breeds in the northeast US and adjacent Canada (includes ''P. s. mediogriseus'') * ''P. s. sandwichensis'' (Aleutian Savannah sparrow), breeds on the
Aleutian Islands The Aleutian Islands (; ; ale, Unangam Tanangin,”Land of the Aleuts", possibly from Chukchi ''aliat'', "island"), also called the Aleut Islands or Aleutic Islands and known before 1867 as the Catherine Archipelago, are a chain of 14 large v ...
and west
Alaskan Peninsula The Alaska Peninsula (also called Aleut Peninsula or Aleutian Peninsula, ale, Alasxix̂; Sugpiaq: ''Aluuwiq'', ''Al'uwiq'') is a peninsula extending about to the southwest from the mainland of Alaska and ending in the Aleutian Islands. The ...
* ''P. s. anthinus'', breeds in the remainder of Alaska, south and east to central
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
and north of the Great Plains to Manitoba * ''P. s. brooksi'' (dwarf Savannah sparrow), breeds in southernmost British Columbia to northernmost
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
* ''P. s. alaudinus'', breeds in coastal northern and central California * ''P. s. nevadensis'', breeds in the northern Great Plains and the Great Basin * ''P. s. brunnescens'', breeds from central Mexico south to Guatemala (includes ''P. s. rufofuscus'') ''P. s. wetmorei'' is a doubtful subspecies that may breed in the mountains of Guatemala. It is known from only five specimens, collected June 11–17, 1897, in
Huehuetenango Department Huehuetenango is one of the 22 departments of Guatemala. It is situated in the western highlands and shares borders with the Mexican state of Chiapas in the north and west; with El Quiché in the east, with Totonicapán, Quetzaltenango and San ...
.


Ipswich sparrow

Some post-breeding dispersal. Formerly considered a distinct species. * ''P. s. princeps'', breeds almost exclusively on
Sable Island Sable Island (french: île de Sable, literally "island of sand") is a small Canadian island situated southeast of Halifax, Nova Scotia, and about southeast of the closest point of mainland Nova Scotia in the North Atlantic Ocean. The island ...
:The Ipswich sparrow is somewhat larger and paler in color than other eastern Savannah sparrows. The breast streaks are narrower and pale brown. Some birds overwinter on the island; others migrate south along the Atlantic coast, usually departing later and returning sooner than mainland birds. Some birds interbreed with ''P. s. savanna'' in
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
. These birds frequently raise three broods in a year. This bird was first observed in winter on the dunes near the town of Ipswich, Massachusetts.


Large-billed sparrows

The large-billed sparrows proper are two dark, large and strong-billed subspecies: * ''Passerculus rostratus/sandwichensis rostratus'', which breed on the Gulf Coast of northeast
Baja California Baja California (; 'Lower California'), officially the Free and Sovereign State of Baja California ( es, Estado Libre y Soberano de Baja California), is a state in Mexico. It is the northernmost and westernmost of the 32 federal entities of Mex ...
and northwest Sonora (some post-breeding dispersal). * ''P. r./s. atratus'', resident on the coast of central Sonora to central Sinaloa (resident) The Belding's (Savannah/large-billed) sparrows are all-year residents of
salt marsh A salt marsh or saltmarsh, also known as a coastal salt marsh or a tidal marsh, is a coastal ecosystem in the upper coastal intertidal zone between land and open saltwater or brackish water that is regularly flooded by the tides. It is domin ...
es of the
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
n Pacific coast. They are dark,
rufous Rufous () is a color that may be described as reddish-brown or brownish-red, as of rust or oxidised iron. The first recorded use of ''rufous'' as a color name in English was in 1782. However, the color is also recorded earlier in 1527 as a dia ...
, and have rather long but not very hefty bills. * ''P. r./s. beldingi'', resident on the Pacific coast from
Morro Bay Morro Bay (''Morro'', Spanish for "Hill") is a seaside city in San Luis Obispo County, California. Located on the Central Coast of California, the city population was 10,757 as of the 2020 census, up from 10,234 at the 2010 census. The town ...
, California, to El Rosario, Baja California (includes ''P. r./s. bryanti'') * ''P. r./s. anulus'', resident around
Sebastián Vizcaíno Bay Sebastián Vizcaíno Bay (Spanish: ''Bahía de Sebastián Vizcaíno'') is a prominent bay along the west coast of the Baja California Peninsula in northwestern Mexico. It is named after explorer Sebastián Vizcaíno. Geography The bay lies on ...
, Baja California * ''P. r./s. guttatus'', resident around San Ignacio Lagoon * ''P. r./s. magdalenae'', resident around
Magdalena Bay Magdalena Bay ( es, Bahía Magdalena) is a long bay in Comondú Municipality along the western coast of the Mexican state of Baja California Sur. It is protected from the Pacific Ocean by the unpopulated sandy barrier islands of Isla Magdalena ...
The San Benito (Savannah/large-billed) sparrow is a resident bird of the
Islas San Benito The Islas San Benito lie in the Pacific Ocean off the west coast of the Mexican state of Baja California, 25 km west of Cedros Island. They are part of the Cedros Island ''delegación'', a subdivision of Ensenada (municipality), Baja Califo ...
off Baja California; a stray bird was observed on
Cedros Island Cedros Island (''Isla de Cedros'', "island of cedars" in Spanish) is an island in the Pacific Ocean belonging to the state of Baja California, Mexico. The dry and rocky island had a population of 1,350 in 2005 and has an area of which include ...
on April 21, 1906. * ''P. r./s. sanctorum'' :This is a large-bodied and large-billed subspecies, similar to ''rostratus''. They utilize different habitat and their breeding season does not seem to coincideFor late April 1906, Thayer & Bangs (1907) report nestlings, young birds molting into adult plumage, but no eggs anymore. Rising ''in'' Zink ''et al.'' (2005) found no breeding activity in late April 1999. Consequently, breeding seems to take place in the winter months, with the last young fledging in April/May. with that of Belding's sparrows. However, their bill size is due to
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
and their habitat choice simply to the lack of alternatives on their barren island home; altogether, it appears to be a fairly recent offshoot from the Belding's sparrows group. It appears as distinct
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 ...
arily from these as does the Ipswich sparrow from the Savannah sparrow proper group, only that there seems to have been more gene flow and/or a larger
founder population In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, using ...
in the case of the latter.


Gallery

File:Savannah Sparrow.jpg, Probably ''P. s. sandwichensis'', Alaska File:Passerculus sandwichensis.JPG, Probably ''P. s. anthinus'', Alaska File:Savannah Sparrow chicks in nest.jpg, ''P. s. labradorius'' nest File:Savannah Sparrow, Newfoundland.jpg, ''P. s. labradorius'', Newfoundland File:Passerculus sandwichensis.jpg, Probably a Belding's sparrow, wintering in California


Notes


References


Further reading

*


External links


Government of Canada Species at Risk Public Registry – Savannah sparrowSavannah sparrow species account
– Cornell Lab of Ornithology

– USGS Patuxent Bird Identification InfoCenter * * * {{Taxonbar, from=Q287893 savannah sparrow American sparrows Birds of North America savannah sparrow savannah sparrow