Geographic Range
Savannah sparrows ( Passerculus sandwichensis ) are native to the Nearctic, Neotropical, and Oceanic Island biogeographic regions, reflecting their broad and diverse distribution. Their range extends from the northern Canadian provinces and Alaska southward to El Salvador. Longitudinally, they occur across the entire North American continent, from Bristol Bay, Alaska, in the west to St. John’s, Newfoundland, in the east.
During the winter months, all populations breeding in Alaska, Canada, and the northernmost regions of the United States migrate to more temperate areas. Their primary wintering grounds include the Great Plains, non-mountainous areas east of the Appalachian Mountains, and regions south of the Rocky Mountains. Wintering populations also occur in parts of the Neotropical region, including the Caribbean islands such as the Bahamas, Belize, the Cayman Islands, and Cuba.
In Mexico, a resident, non-migratory population occupies highland regions within the Nearctic zone, particularly the Trans-Mexican Volcanic Belt and the Sierra Madre Occidental. This population spans from Guanajuato in the north to Morelos in the south, and from Puebla in the east to Jalisco in the west.
Non-migratory populations are also established in the northeastern United States, from Norfolk, Virginia, to the southern tip of Maine, and along the northwestern coast from central California to the Washington State–Canada border. These resident groups persist year-round within localized coastal environments.
Additionally, vagrant individuals have been observed on Oceanic Islands and in overseas
territories, with records from Bermuda, Japan, the Republic of Korea, the Azores (Portugal),
eastern Siberia (Russian Federation), and the United Kingdom.
- Biogeographic Regions
- nearctic
- neotropical
- oceanic islands
Habitat
Savannah sparrows occupy a diverse range of open habitats, including grasslands, marshes, agricultural fields, coastal meadows, and highland plains. Habitat vary among populations, with migratory individuals breeding in temperate grasslands and tundra, while resident, non-migratory populations persist in specific regions year-round.
During the winter months, migratory populations utilize similar open habitats, including salt marshes, coastal prairies, and inland grasslands. Although Savannah sparrows are mainly terrestrial, they may also be found in wetland environments, including marshes and riparian areas, particularly during migration and the non-breeding season. Habitat selection is strongly influenced by the availability of low vegetation for nesting and cover, as well as open ground for foraging. Some of these areas are at elevations as low as sea level.
In central Mexico, non-migratory population primarily inhabits the high-altitude grasslands
of the Trans-Mexican Volcanic Belt and Sierra Madre Occidental, which are at elevations
ranging from 1,500 to 4,500 meters, with a typical elevation between 2,000 and 3,000
meters. These populations are commonly found in open fields, shrublands, and disturbed
habitat such as pastures and cultivated land.
- Habitat Regions
- temperate
- tropical
- terrestrial
- freshwater
- Terrestrial Biomes
- tundra
- savanna or grassland
- chaparral
- scrub forest
- mountains
- Wetlands
- marsh
- Other Habitat Features
- suburban
- agricultural
- riparian
Physical Description
Savannah sparrows are small, compact birds with short, notched tails and small, conical bills. They measure between 11 and 15.5 cm in length, with wingspans ranging from 19 to 23 cm. They weigh between 15 and 28 grams. This species is sexually monomorphic in plumage, meaning males and females look alike; however, males tend to be slightly larger on average, though specific measurement comparisons between the sexes are limited.
Feather patterns include brown streaks, giving them a distinctive appearance. Across most of their range, they have a pale beige median crown stripe, though this feature is absent in some salt marsh populations. Their eyebrow stripe is typically yellow at the front anteriorly but may appear beige in some individuals. A thin, dark malar stripe is present, and their remiges and rectrices are brown without white markings. They lack distinct wing bars, but a small yellow patch is often visible at the bend of the wing. Their throats, breasts, and bellies range from whitish to pale beige, with brown streaks on the breast that may form a small central spot. The throat and belly may be either streaked or clear. Their legs are pinkish to horn-colored, and they have brown irises.
Juveniles resemble adults but have heavier streaking on the breast, more rufous tones on the wings and tail, and little to no yellow in the eyebrow stripe. Their overall coloration is generally buffier, and they tend to have a less distinct crown stripe. The lower mandible is paler than the upper, and their tails are slightly forked.
Savannah sparrows are often compared to species in the closely related genus Melospiza , such as song sparrows ( Melospiza melodia ) and Lincoln's sparrows ( Melospiza lincolnii ). Members of the genus Melospiza generally exhibit longer tails, more rounded wings, and more heavily streaked plumage overall, especially on the underparts. Additionally, Melospiza species tend to have slightly heavier bills and lack the yellow eyebrow marking characteristic of most Savannah sparrows.
These combined features, especially the presence of yellow near the lores, compact
body shape, and fine streaking make Savannah sparrows distinguishable from other sparrows
within the same family.
- Other Physical Features
- endothermic
- bilateral symmetry
- Sexual Dimorphism
- sexes alike
- male larger
Development
Savannah sparrows, like all birds, exhibit determinate growth, and sex is determined genetically with Z or W chromosomes.
After mating, females lay 3 to 6 eggs in a well-hidden grass, rootlets, and hair-lined nest on the ground. The eggs are incubated for 10 to 13 days, primarily by the females. During this time, the embryo develops inside the egg, which is nourished by the yolk. The eggs of Savannah sparrows are primarily light in color, ranging from pale white to shades greenish or blueish. Their eggs are typically spotted in a variety of darker colors from red to gray and even brown. Savannah sparrow eggs are noted for their shiny and smooth appearance.
Hatchlings are altricial, meaning they hatch blind, featherless, and completely dependent on parental care. The hatchlings grow rapidly and start developing their feathers. Nestlings remain in the nest for 8 to 13 days before fledging. As nestlings, they open their eyes, their down feathers are replaced by juvenile plumage, and they strengthen their wings and legs. At the fledgling stage, young Savannah sparrows stay close to their parents, who continue feeding them until they learn to forage on their own. This stage lasts a couple of weeks as the young birds refine their flying and survival skills.
Savannah sparrows initially exhibit juvenile plumage. After their first molt, they
start resembling adults. Savannah sparrows reach sexual maturity at about age 1 and
exhibit adult plumage and behaviors, including territoriality and courtship displays.
Reproduction
Savannah sparrows display both monogamous and polygynous mating systems. When breeding in northern regions with shorter seasons, males and females form monogamous pairs. The short months of the tundra breeding season and the necessity of male parental care contribute to the need of monogamy in northern regions. In more southern populations, males will mate with multiple females. The proportion of polygynous males varies annually and regionally, ranging from 15% to 43% across studies. Factors such as territory quality and resource distribution influence the occurrence of polygyny.
Males attract females through song displays. Males perch on shrubs, trees, or fence posts to establish territory boundaries. Males actively defend their territories through different displays and chases to deter rival males, ensuring exclusive access to resources and potential mates within their area.
The mating system influences the social structure and parental roles within Savannah
sparrow populations. In monogamous pairs, both parents share duties such as, building
the nest, egg incubation, and feeding and fostering their offspring. In polygynous
systems, where males have multiple mates, males provide less paternal care to their
offspring. Females in these systems provide practically all the care for their offspring,
which reduces their reproductive success.
- Mating System
- monogamous
- polygynous
The reproductive cycle of Savannah sparrows is closely tied to seasonal changes, with breeding typically occurring between April and August. This period aligns with warmer temperatures and greater food availability, which supports successful nesting. Savannah sparrows are iteroparous, meaning they can breed multiple times throughout their lives.
Within a single breeding season, they produce 1 to 3 broods. Females lay between 3 and 6 eggs per clutch, with typically 4 to 5 eggs. The eggs are incubated primarily by the females for 10 to 13 days (average = 12 days). Once hatched, hatchlings weigh around 1 to 3 g (average = 2 grams), and the altricial chicks are blind, featherless, and entirely dependent on parental care. Both parents participate in feeding and protecting the young.
Fledging occurs 8 to 11 days after hatching, most often at 9 days. Even after leaving the nest, young Savannah sparrows remain dependent on their parents for an additional 14 to 21 days while they learn to forage and evade predators.
Savannah sparrows reach sexual maturity within their first year, typically by the
age of 9 to 12 months. Like all birds, they mate using internal fertilization and
are oviparous.
- Key Reproductive Features
- iteroparous
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
- sexual
- fertilization
- oviparous
Savannah sparrows exhibit parental investment from both parents, with a more prominent role played by females. Particularly in southern populations where polygamy is found and there is little to no parental investment by males.
In northern regions where monogamous pairs are formed, pre-fertilization parental investment is in the form of the females production of healthy eggs and the female's ability to find safe and suitable nesting sites in preparation for breeding. The males parental investment at this stage is mainly defending resource rich territories.
Pre-hatching parental investment is predominantly the females responsibility as she is the main constructor of the nest, and they are the main incubators of egg for 10 to 13 days. Males during this stage help by provisioning the female with food, defending their territory and occasionally incubating the eggs.
Post-hatch parental investment reaches its peak due to the altricial nature of the hatchlings. Both parents help feed and protect their offspring, females often take the lead in brooding. This behavior helps the young maintain body heat and shields them from environmental hazards. This stage typically lasts 8 to 13 days.
Pre-independence both parents will continue to provision their young while teaching
skills such as foraging and predator recognition. Post-independence juveniles remain
in close proximity of their parental territories for a few weeks before fully dispersing.
- Parental Investment
- altricial
- male parental care
- female parental care
-
pre-fertilization
- provisioning
-
protecting
- male
- female
-
pre-hatching/birth
-
provisioning
- female
-
protecting
- male
- female
-
provisioning
-
pre-weaning/fledging
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
-
pre-independence
-
provisioning
- male
- female
-
protecting
- male
- female
-
provisioning
Lifespan/Longevity
Savannah sparrows in the wild can live up to 6.5 years. The oldest individual in captivity lived to at least 18 years. Their expected life spans in captivity is 12 years and 5 years in the wild.
Factors that limit lifespan in the wild include predation, extended periods of precipitation,
and loss of habitat. Annual adult survival rates are between 28%-42%.
Behavior
Savannah sparrows are semi-social birds and are usually a part of loose flocks in the non-breeding season. During the breeding season, males become territorial and defend their territory through songs and other displays.
Savannah sparrows are migratory birds that move south from September to October. They migrate nocturnally. These sparrows return to breeding grounds in the spring, and arrival time is later at more northern latitudes. For example, they reach breeding grounds in Kentucky in February. In March, they arrive in Missouri and southern Canada. In April, they reach South Dakota and Nova Scotia, Canada. Finally, by early May, they arrive in Alaska. Males typically arrive at breeding grounds one week prior to potential mates.
Savannah sparrows have distinct flight patterns which are short, with flights that are close to the ground. They are also cursorial and walking and running through grasslands rather than flying. They are diurnal which means they are active during the day.
There is no strong evidence of a social hierarchy among Savannah sparrows based on
territory during breeding seasons. Outside their breeding season, these birds forage
in mixed-species flocks and communicate through songs and other displays.
- Key Behaviors
- cursorial
- terricolous
- flies
- diurnal
- motile
- migratory
- territorial
- social
Home Range
Home ranges for Savannah sparrows during the breeding season average 5,000 square
meters (range 2500 to 10000 m^2). The range is dependent on population density and
the overall quality of their habitat. Males are territorial and exhibit fighting and
defensive behaviors. They can chase one another for up to 200 m away from the nesting
area, although the territory size has not been quantified. They fight most early in
the breeding season when pairs are first getting established. Later in the season,
males revert to less aggressive displays (quivering, alarm calls, wing-hits just off
the ground).
Communication and Perception
Savannah sparrows rely on acoustic communication, with males singing to attract mates and mark their territory. Short calls help them stay in contact and signal danger. Numerous studies are currently being conducted on the songs of the Savannah sparrow, examining how variations in tone and pitch influence their meaning. Males have intricate songs that last 2 to 3 seconds. They start with several distinct high-pitched chips and trail into a buzzy sound at 7.2 kHz. It's described as insect-like. The song descends briefly with a few cleared, lower-kHz (4.4 kHz) notes at the end. Although only males can create songs, both sexes can make less complicated alarm calls and nestlings can make simple begging calls.
In addition to vocalizations, they use visual signals like different body postures and wing movements during courtship and territorial defense. These sparrows have tactile behaviors, such as grooming, reinforce bonds between mates and the kin between parents and chicks. They establish territories by visual displays plus tactile wing-tapping and jumping just off the ground to fight.
Their visual and acoustic senses are advanced, allowing them to detect predators, locate food, and interact socially.
Xu et al. (2021) suggest that most songbirds use a magnetic compass in migratory routes.
Food Habits
Savannah sparrows are omnivorous birds that consume both plants and animals. During the breeding season, their diet consists mainly of insects and other small invertebrates, classifying them as insectivores. These birds feed on a variety of arthropods, including grasshoppers (Order Orthoptera), beetles (Order Coleoptera), caterpillars (Order Lepidoptera), flies (Order Diptera), and spiders (Order Araneae).
In colder months, they shift toward a granivorous diet, relying more on grains and
seeds for sustenance. In these months, they consume seeds from different grasses and
forbs, such as (
Panicum
), ragweed (
Ambrosia
), and foxtail (
Setaria
)
- Primary Diet
-
carnivore
- insectivore
- eats non-insect arthropods
- herbivore
- omnivore
- Animal Foods
- terrestrial non-insect arthropods
- Plant Foods
- seeds, grains, and nuts
Predation
Savannah sparrows possess cryptic adaptations which affect their distinctive coloration. These sparrows hide in grasslands and possess brown and beige streaked plumage to blend into their surrounds. Savannah remains low to the ground and freezes in place when threatened. If detected, they employ swift, erratic flight patterns in short bursts to evade predators.
During early stages of their life, Savannah sparrows are venerable to snakes and foxes
such as eastern racer snakes (
Coluber constrictor
) and red foxes (
Vulpes vulpes
). As adults, Savannah sparrows face threats from predators such as red-tailed hawks
(
Buteo jamaicensis
), domestic cats (
Felis catus
), and short-eared owls (
Asio flammeus
).
- Anti-predator Adaptations
- cryptic
Ecosystem Roles
Savannah sparrows play important roles in their ecosystems, primarily through being prey for larger birds and consuming insects and seeds.
Brown-headed cowbirds ( Molothrus ater ) are brood parasites that rely on Savannah sparrows to raise their young; this parasitism can impact the reproductive success of Savannah sparrows.
Ectoparasites include blowflies ( Protocalliphora hirudo ), which are rare but can affect a small percentage of nestlings. Louse flies in the family Hippoboscidae were found in fledglings at rates of 10%. Adults may rarely host ticks of unreported species.
Protozoal parasites (
Plasmodium
) are endoparasite blood protozoans that cause avian malaria. prevalence varies widely
among Savannah sparrow populations, with infection rates ranging from 0% to 60% across
14 studied groups. Infections are more frequently observed in mainland males and are
associated with increased sexual size dimorphism, suggesting a possible a possible
role of parasite-mediated sexual selection.
- brown-headed cowbirds ( Molothrus ater )
- protozoal parasites ( Plasmodium )
- blowflies ( Protocalliphora hirudo )
- louse flies (family Hippoboscidae)
- ticks
Economic Importance for Humans: Positive
Savannah sparrows serve as a small part of the large bird watching industry that generates
around $108 billion annually across the United States. Local economies benefit economically
through guided bird watching tours and other ecotourism activities.
- Positive Impacts
- ecotourism
Economic Importance for Humans: Negative
There are no known adverse economic effects of Savannah sparrows on humans.
Conservation Status
Savannah sparrows are listed as "Least Concern" on the IUCN Red List. They are protected under the United States Migratory Bird Treaty Act, which makes it illegal to kill, take or have possession of any migratory bird. While the species is not listed as a whole under the Endangered Species Act, there are some subspecies that have protections. Passerculus sandwichensis beldingi is a subspecies that inhibits coastal salt marshes in California is listened as an endangered species. Passerculus sandwichensis princeps inhabits Sable Island, Nova Scotia and is designated as "Special Concern" by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC).
The primary threats to vulnerable subspecies, such as Passerculus sandwichensis beldingi , stem from habitat loss, particularly the destruction of coastal salt marshes for development and agricultural expansion. These habitats are essential for nesting and foraging. Similarly, individuals in the subspecies Passerculus sandwichensis princeps face substantial risks due to storm exposure and shoreline erosion in their breeding areas. Passerculus sandwichensis princeps in Manitoba declined 77% in 25 years, largely due to habitat altercations.
Other threats documented include mowing practices in agricultural fields and at airports. Hay fields can result in up to 99% nest failure, this is due to fledging times occurring around June 21st which is optimal mowing days. Airports offer great grassland habitats which are suitable for Savannah sparrows, but due to mowing regimes their results are often high nest failure.
To safeguard Savannah sparrows and their vulnerable subspecies, several protections
have been in place. At the Federal level Savannah sparrows are protected under the
Migratory Bird Treaty Act, which prohibits the capturing, possession or killing of
Savannah sparrows without proper authorization. For subspecies at greater risk, such
as
Passerculus sandwichensis beldingi
, listing under the Endangered Species Act ensures targeted conservation efforts,
including habitat preservation and restoration initiatives in California’s coastal
salt marshes. Similarly,
Passerculus sandwichensis princeps
, designated as a species of Special Concern by COSEWIC in Canada, benefits from monitoring
programs and conservation planning aimed at minimizing habitat degradation from storm
impacts and shoreline erosion. Additionally, land management practices are being adjusted
in some areas such as delaying mowing during nesting season to reduce nest failure.
Additional Links
Contributors
Alec Correa (author), Radford University, Natalie May (editor), Radford University, Alexander McVicker (editor), Radford University, Karen Powers (editor), Radford University, Tanya Dewey (editor), University of Michigan-Ann Arbor.
- Nearctic
-
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- Neotropical
-
living in the southern part of the New World. In other words, Central and South America.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- oceanic islands
-
islands that are not part of continental shelf areas, they are not, and have never been, connected to a continental land mass, most typically these are volcanic islands.
- native range
-
the area in which the animal is naturally found, the region in which it is endemic.
- temperate
-
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
- tropical
-
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- terrestrial
-
Living on the ground.
- freshwater
-
mainly lives in water that is not salty.
- tundra
-
A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.
- tropical savanna and grassland
-
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
- savanna
-
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
-
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
- chaparral
-
Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.
- scrub forest
-
scrub forests develop in areas that experience dry seasons.
- mountains
-
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- marsh
-
marshes are wetland areas often dominated by grasses and reeds.
- suburban
-
living in residential areas on the outskirts of large cities or towns.
- agricultural
-
living in landscapes dominated by human agriculture.
- riparian
-
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
- monogamous
-
Having one mate at a time.
- polygynous
-
having more than one female as a mate at one time
- iteroparous
-
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- seasonal breeding
-
breeding is confined to a particular season
- sexual
-
reproduction that includes combining the genetic contribution of two individuals, a male and a female
- fertilization
-
union of egg and spermatozoan
- oviparous
-
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
- altricial
-
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- male parental care
-
parental care is carried out by males
- female parental care
-
parental care is carried out by females
- diurnal
-
- active during the day, 2. lasting for one day.
- motile
-
having the capacity to move from one place to another.
- migratory
-
makes seasonal movements between breeding and wintering grounds
- territorial
-
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
- social
-
associates with others of its species; forms social groups.
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- visual
-
uses sight to communicate
- tactile
-
uses touch to communicate
- acoustic
-
uses sound to communicate
- chemical
-
uses smells or other chemicals to communicate
- magnetic
-
(as perception channel keyword). This animal has a special ability to detect the Earth's magnetic fields.
- cryptic
-
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- ecotourism
-
humans benefit economically by promoting tourism that focuses on the appreciation of natural areas or animals. Ecotourism implies that there are existing programs that profit from the appreciation of natural areas or animals.
- carnivore
-
an animal that mainly eats meat
- insectivore
-
An animal that eats mainly insects or spiders.
- herbivore
-
An animal that eats mainly plants or parts of plants.
- granivore
-
an animal that mainly eats seeds
- omnivore
-
an animal that mainly eats all kinds of things, including plants and animals
- endothermic
-
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- bilateral symmetry
-
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
References
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COSEWIC, 2023. "Savannah Sparrow princeps subspecies (Passerculus sandwichensis princeps): COSEWIC rapid review of classification 2023" (On-line). Government of Canada. Accessed April 01, 2025 at https://www.canada.ca/en/environment-climate-change/services/species-risk-public-registry/cosewic-assessments-status-reports/savannah-sparrow-2023.html. .
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Harrison, C. 1978. A Field Guide to Nests, Eggs, and Nestlings of North American Birds . Cleveland, Ohio: Collins Press.
Headstrom, R. 1970. A Complete Field Guide to Nests in the United States (including those of Birds, Mammals, Insects, Fishes, Reptiles, and Amphibians) . New York, New York: Van Rees Press.
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Rising, J. 1988. Geographic variation in sex ratios and body size in wintering flocks of Savannah sparrows (Passerculus sandwichensis). The Wilson Bulletin , 100/2: 183-203.
Rising, J. 2000. Geographic variation in size and shape of Savannah sparrows (Passerculus sandwichensis). Studies In Avian Biology , 23/12: 1-65.
Sharma, S., S. Dobney, R. Norris, S. Doucet, A. Newman, J. Burant, I. Moran, S. Mueller, H. Spina, D. Mennill. 2024. Effects of age, breeding strategy, population density, and number of neighbors on territory size and shape in Passerculus sandwichensis (Savannah sparrow). Ornithology , 141/4: ukae025. Accessed February 03, 2025 at https://doi.org/10.1093/ornithology/ukae025 .
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Wheelwright, N., J. Rising. 2020. "Savannah sparrow  (Passerculus sandwichensis), version 1.0. In Birds of the World (A. F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA." (On-line). Accessed February 03, 2025 at doi: 10.2173/bow.savspa.01 .
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