Editing Grasshopper sparrow

{{Short description|Species of bird}}
{{Speciesbox
| image = Ammodramus savannarum 160849415 (cropped).jpg
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name="iucn">{{cite iucn |author=BirdLife International. |year=2021 |title=''Ammodramus savannarum'' |volume=2021 |page=e.T22721144A138486868 |doi=10.2305/IUCN.UK.2021-3.RLTS.T22721144A138486868.en |access-date=11 November 2021}}</ref>
| genus = Ammodramus
| species = savannarum
| authority = ([[Johann Friedrich Gmelin|Gmelin]], 1789)
| synonyms =
''Coturniculus savannarum''
| range_map = Ammodramus savannarum map.svg
| range_map_caption = {{leftlegend|#FF7F2A|Breeding}}{{leftlegend|#7137C8|Year-round}}{{leftlegend|#5F8DD3|Nonbreeding}}
}}

The '''grasshopper sparrow''' ('''''Ammodramus savannarum''''') is a small [[New World sparrow]]. It belongs to the genus ''[[Ammodramus]],'' which contains three species that inhabit [[grassland]]s and [[prairie]]s. Grasshopper sparrows are sometimes found in crop fields and they will readily colonize reclaimed grassland. In the core of their range, grasshopper sparrows are dependent upon large areas of grassland where they avoid trees and shrubs.<ref>{{Cite journal |last1=Herse |first1=Mark R. |last2=With |first2=Kimberly A. |last3=Boyle |first3=W. Alice |date=September 2018 |editor-last=Stanley |editor-first=Margaret |title=The importance of core habitat for a threatened species in changing landscapes |journal=Journal of Applied Ecology |language=en |volume=55 |issue=5 |pages=2241–2252 |doi=10.1111/1365-2664.13234|s2cid=90975257 |doi-access=free |bibcode=2018JApEc..55.2241H }}</ref> They seek out heterogenous patches of prairie that contain clumps of dead grass or other vegetation where they conceal their nest, and also contain barer ground where they forage for insects (especially grasshoppers), spiders, and seeds.<ref name=":7" /> Grasshopper sparrows are unusual among New World sparrows in that they sing two distinct song types, the prevalence of which varies with the nesting cycle. The primary male song, a high trill preceded by a stereotyped series of short chips, is reminiscent of the sounds of grasshoppers<ref name=":1" /> and is the origin of this species' name. Like some other birds of the central North American grasslands, this species also moves around a lot, not only via annual migrations, but individuals frequently disperse between breeding attempts or breeding seasons. Grasshopper sparrows are in steep decline across their range, even in the core of the breeding distribution in the tallgrass prairies of the central Great Plains.<ref name=SOB2022/> The [[Florida grasshopper sparrow]] (''Ammodramus savannarum floridanus'') is highly endangered.

==Taxonomy==
The grasshopper sparrow was described in 1713 by English parson and naturalist [[John Ray]] based on a specimen collected in Jamaica. It was described again and illustrated in 1725 by the naturalist and collector [[Hans Sloane]]. Both authors used the English name "Savanna bird".<ref>{{ cite book | last=Ray | first=John | author-link=John Ray | year=1713 | title=Synopsis methodica avium & piscium | volume=Avium | language=Latin | location=London | publisher=William Innys | page=188 | url=https://www.biodiversitylibrary.org/page/6355454 }}</ref><ref>{{ cite book | last=Sloane | first=Hans | author-link=Hans Sloane | year=1725 | title=A Voyage to the Islands Madera, Barbados, Nieves, S. Christophers and Jamaica : with the natural history of the herbs and trees, four-footed beasts, fishes, birds, insects, reptiles, &c. of the last of those islands | volume=2 | location=London | publisher=Printed for the author | page=[https://www.biodiversitylibrary.org/page/188477 306], [https://www.biodiversitylibrary.org/page/188770 Plate 259, Fig 5] }}</ref> When the German naturalist [[Johann Friedrich Gmelin]] revised and expanded [[Carl Linnaeus]]'s ''[[Systema Naturae]]'' in 1789 he included the grasshopper sparrow. He placed it with the finches in the [[genus]] ''[[Fringilla]]'', coined the [[binomial nomenclature|binomial name]] ''Fringilla savannarum'' and cited the earlier authors.<ref>{{ cite book | last=Gmelin | first=Johann Friedrich | author-link=Johann Friedrich Gmelin| year=1789 | title=Systema naturae per regna tria naturae : secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis | edition=13th | volume=1, Part 2 | language=Latin | location=Lipsiae [Leipzig] | publisher=Georg. Emanuel. Beer | page=921 | url=https://www.biodiversitylibrary.org/page/2656416 }}</ref> The grasshopper sparrow is now placed with two other American sparrows in the genus ''[[Ammodramus]]'' that was introduced by [[William Swainson]] in 1827.<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | editor3-last=Rasmussen | editor3-first=Pamela | editor3-link=Pamela Rasmussen | date=July 2023 | title=New World Sparrows, Bush Tanagers | work=IOC World Bird List Version 13.2 | url=https://www.worldbirdnames.org/bow/sparrows/ | publisher=International Ornithologists' Union | access-date=14 November 2023}}</ref>

Twelve [[subspecies]] are recognised:<ref name=ioc/>
* ''A. s. perpallidus'' ([[Elliott Coues|Coues]], 1872) – southwest Canada and central, west USA
* ''A. s. ammolegus'' [[Harry C. Oberholser|Oberholser]], 1942 – south Arizona (southwest USA) and northwest Mexico
* ''A. s. pratensis'' ([[Louis Pierre Vieillot|Vieillot]], 1818) – southeast Canada and east USA
* ''A. s. floridanus'' ([[Edgar Alexander Mearns|Mearns]], 1902) – Florida (southeast USA)
* ''A. s. bimaculatus'' [[William Swainson|Swainson]], 1827 – central, south Mexico, Guatemala, Nicaragua and Costa Rica
* ''A. s. beatriceae'' [[Storrs L. Olson|Olson]], 1980 – central Panama
* ''A. s. cracens'' ([[Outram Bangs|Bangs]] & [[Charles Horton Peck|Peck]], 1908) – north, east Guatemala, [[Belize]], east Honduras and northeast Nicaragua
* ''A. s. caucae'' [[Frank Chapman (ornithologist)|Chapman]], 1912 – Colombia
* ''A. s. savannarum'' ([[Johann Friedrich Gmelin|Gmelin, JF]], 1789) – Jamaica
* ''A. s. intricatus'' [[Ernst Hartert|Hartert, EJO]], 1907 – [[Hispaniola]]
* ''A. s. borinquensis'' [[James L. Peters|Peters, JL]], 1917 – [[Puerto Rico]]
* ''A. s. caribaeus'' (Hartert, EJO, 1902) – [[Netherlands Antilles]]

==Description==
These small sparrows measure {{convert|10|-|14|cm|in|abbr=on}} in length, span about {{convert|17.5|cm|in|abbr=on}} across the wings and weigh from {{convert|13.8|to|28.4|g|oz|abbr=on}}, with an average of {{convert|17|g|oz|abbr=on}}.<ref name=DEC/><ref name=CRC/> Adults have upper parts streaked with brown, grey, black and white; they have a light brown breast, a white belly and a short brown tail. Their face is light brown with a white eye ring and a dark brown crown with a central narrow light stripe. Adults sport bright yellow feathers at the crook of their underwing, and have a yellow-to-amber patch above their lores. Males and females cannot be distinguished by their plumage, and young birds molt into adult plumage within a few months of fledging.<ref name=":8" />

==Distribution and habitat==
[[File:Late-summer nesting habitat for Grasshopper Sparrow in NE Kansas.jpg|thumb|Late-July grasshopper sparrow territory with nest at the Konza Prairie showing preferred patchiness of vegetation including low ground for foraging and denser patches in which to conceal nests.]]
Their breeding habitat is open fields and [[prairie]] grasslands across southern [[Canada]], the United States, [[Mexico]] and [[Central America]], the [[Caribbean]]. There is a small endangered population in the [[Andes]] of [[Colombia]] and (perhaps only formerly) [[Ecuador]]. In the central USA, birds are quick to locate newly available habitat, and can be attracted to sites by playing conspecific song.<ref>{{Cite journal |last1=Andrews |first1=John E. |last2=Brawn |first2=Jeffrey D. |last3=Ward |first3=Michael P. |date=May 2015 |title=When to use social cues: Conspecific attraction at newly created grasslands |journal=The Condor |language=en |volume=117 |issue=2 |pages=297–305 |doi=10.1650/CONDOR-14-172.1 |s2cid=85979571 |issn=0010-5422|doi-access=free }}</ref> The spatial distribution of territories on the landscape can be clumped with several individuals defending territories near to one another interspersed by large areas of unused and apparently suitable habitat. The reasons for this clumping are elusive; aggregation is not related to group defense against predators or brood parasite, cooperative care, extra-pair matings, or kin selection.<ref name=":9">{{Cite journal |last1=Winnicki |first1=S. K. |last2=Munguía |first2=S. M. |last3=Williams |first3=E. J. |last4=Boyle |first4=W. A. |date=February 2020 |title=Social interactions do not drive territory aggregation in a grassland songbird |journal=Ecology |language=en |volume=101 |issue=2 |pages=e02927 |doi=10.1002/ecy.2927 |pmid=31713849 |s2cid=207955738 |issn=0012-9658|doi-access=free |bibcode=2020Ecol..101E2927W }}</ref> Appropriate habitat is strongly influenced by plant structure and precipitation amounts during previous seasons.<ref name="Macías-Duarte 41–49">{{Cite journal |last1=Macías-Duarte |first1=Alberto |last2=Panjabi |first2=Arvind O. |last3=Pool |first3=Duane B. |last4=Ruvalcaba-Ortega |first4=Irene |last5=Levandoski |first5=Greg J. |date=2018-09-01 |title=Fall vegetative cover and summer precipitation predict abundance of wintering grassland birds across the Chihuahuan desert |journal=Journal of Arid Environments |language=en |volume=156 |pages=41–49 |doi=10.1016/j.jaridenv.2018.04.007 |s2cid=90845822 |issn=0140-1963|doi-access=free |bibcode=2018JArEn.156...41M }}</ref> Grasshopper sparrows avoid [[Woody plant|woody vegetation]], and in the wetter parts of their range, prefer fields that are burned every 2–3 years (which reduces encroachment of trees and shrubs) and seek out areas that are moderately grazed by cattle or bison.

==Behavior==

=== Movement ===

==== Migration ====
Subspecies of grasshopper sparrows differ in their [[Migration (ecology)|migratory behavior]]. The birds breeding in most of the eastern US and southern Canada (''A. s. pratensis'') migrate latitudinally up to several thousand kms each year,<ref>{{cite book |last=Pyle |first=Peter |url=https://www.worldcat.org/oclc/38593534 |title=Identification guide to North American birds : a compendium of information on identifying, ageing, and sexing "near-passerines" and passerines in the hand |date=1997 |publisher=Slate Creek Press |others=Steve N. G. Howell, Siobhan Ruck, Institute for Bird Populations, Point Reyes Bird Observatory |isbn=0-9618940-2-4 |location=Bolinas, Calif. |oclc=38593534}}</ref> spending winters along the Gulf of Mexico and the Caribbean.<ref name="Hill 680–692">{{Cite journal |last1=Hill |first1=Jason M. |last2=Renfrew |first2=Rosalind B. |date=January 2019 |title=Migratory patterns and connectivity of two North American grassland bird species |journal=Ecology and Evolution |language=en |volume=9 |issue=1 |pages=680–692 |doi=10.1002/ece3.4795 |pmc=6342103 |pmid=30680148|bibcode=2019EcoEv...9..680H }}</ref> The birds breeding through most of the Western part of their North American range (''A. s. perpallidus'') winter from Texas west to California, and south into the highland desert grasslands of northern and central Mexico. ''A. s. ammolegus'', which breeds in the desert grasslands of SE Arizona, SW New Mexico, and adjoining states in northern Mexico, appears to be a short-distance, partial migrant, with some birds remaining resident year-round, and others likely moving further south in winter. Birds of multiple breeding populations and subspecies winter together in SE Arizona.<ref>{{Cite journal |last=Gordon |first=Caleb E. |date=2000 |title=Movement Patterns of Wintering Grassland Sparrows in Arizona |url=https://www.jstor.org/stable/4089599 |journal=The Auk |volume=117 |issue=3 |pages=748–759 |doi=10.2307/4089599 |jstor=4089599 |issn=0004-8038|url-access=subscription }}</ref> Grasshopper sparrows breeding outside of North America, including those living in the Caribbean, Central America, as well as the Florida grasshopper sparrow are not known to migrate.  

Males typically migrate north in the spring a week or two earlier than do females. Data from [[light level geolocator]]s indicate that males are present on the breeding grounds from April to October,<ref name="Hill 680–692"/> consistent with birds completing their annual molt prior to migration.<ref name=":8">Pyle, P., S.L. Jones, and J. M. Ruth. 2008. Molt and aging criteria for four North American grassland passerines. U.S. Department of Interior, Fish and Wildlife Service, Biological Technical Publication, FWS/BTP-R6011-2008, Washington, D.C.</ref> Individuals migrated an average of ~2,500&nbsp;km over ~30 days.<ref name="Hill 680–692" /> Birds migrate mostly over land, make few stop-overs of short duration, and travel at about 82&nbsp;km/day.<ref name="Hill 680–692" /> Data from the [https://motus.org/ Motus network] largely confirm the migratory patterns gleaned from prior work.<ref>{{Cite web |title=Motus Wildlife Tracking System |url=https://motus.org/data/tracksSelect?e=2013-01-01&l=2022-12-31&s=18900 |access-date=2022-10-26 |website=motus.org}}</ref> However, relatively few movement tracks for this species exist so far, potentially due to the sparrows' secretive behavior during most of the year other than the breeding season.<ref>{{Cite journal |last1=Williams |first1=Emily J. |last2=Boyle |first2=W. Alice |date=January 2018 |title=Patterns and correlates of within-season breeding dispersal: A common strategy in a declining grassland songbird |journal=The Auk |language=en |volume=135 |issue=1 |pages=1–14 |doi=10.1642/AUK-17-69.1 |s2cid=90295307 |issn=0004-8038|doi-access=free }}</ref> Few recoveries of banded birds have ever been made, especially at sites other than a birds’ initial banding location.<ref>{{Cite web |title=Bird Banding Laboratory |url=https://www.pwrc.usgs.gov/BBL/Bander_Portal/login/bbl_data_request_summary.php |access-date=2022-10-26 |website=www.pwrc.usgs.gov}}</ref> In Oklahoma, grasshopper sparrows are one of the species most frequently found dead after colliding with windows, despite rarely being detected in migration.<ref>{{Cite web |title=Grasshopper Sparrow |url=https://birdsmack.wordpress.com/category/grasshopper-sparrow/ |access-date=2022-11-17 |website=Bird-Window Collisions at OK State |date=2 August 2018 |language=en}}</ref>

==== Dispersal ====
In Maryland, a study of [[Biological dispersal|natal dispersal]] (the movement from a natal site to the site of first reproduction) revealed that most fledglings remained within their natal habitat during the summer months after fledging, and most recaptured fledglings were encountered within a couple hundred meters of their natal nests.<ref name=":5">{{Cite journal |last1=Anthony |first1=Travis |last2=Gill |first2=Douglas E. |last3=Small |first3=Daniel M. |last4=Parks |first4=Jared |last5=Sears |first5=Henry F. |date=June 2013 |title=Post-Fledging Dispersal of Grasshopper Sparrows (''Ammodramus savannarum'') On A Restored Grassland In Maryland |url=https://bioone.org/journals/the-wilson-journal-of-ornithology/volume-125/issue-2/12-121.1/Post-Fledging-Dispersal-of-Grasshopper-Sparrows-Ammodramus-savannarum-On-A/10.1676/12-121.1.full |journal=The Wilson Journal of Ornithology |volume=125 |issue=2 |pages=307–313 |doi=10.1676/12-121.1 |s2cid=17415980 |issn=1559-4491|url-access=subscription }}</ref> Although distances recaptured from the nest increased with time, the average fledgling was recaptured only 346 meters from its nest and recaptured 33 days after fledging. The longest distance detected was 1.6 kilometers, which was accomplished in less than 20 days.<ref name=":5" /> Data on first-year return rates (or "philopatry", the inverse of natal dispersal) is limited, but in NE Kansas, ~2% of young birds returned to the site to breed in the subsequent year.<ref>Boyle lab, unpublished data</ref>

Grasshopper sparrows are unusual in their particularly high rates of breeding dispersal (i.e., one-way movements between successive breeding sites).<ref name=":6" /> Their high dispersal tendency may be due to the dynamic nature of the grassland environments on which they depend<ref name=":4">{{Cite journal |last1=Jones |first1=Stephanie L. |last2=Dieni |first2=J. Scott |last3=Green |first3=Michael T. |last4=Gouse |first4=Paula J. |title=Annual Return Rates of Breeding Grassland Songbirds |date=March 2007 |url=https://bioone.org/journals/the-wilson-journal-of-ornithology/volume-119/issue-1/05-158.1/ANNUAL-RETURN-RATES-OF-BREEDING-GRASSLAND-SONGBIRDS/10.1676/05-158.1.full |journal=The Wilson Journal of Ornithology |volume=119 |issue=1 |pages=89–94 |doi=10.1676/05-158.1 |s2cid=16966765 |issn=1559-4491}}</ref> Like some other grassland-dependent species, their dispersal movements lead them to be called semi-nomadic, as they opportunistically take advantage of suitable habitats as they appear on the landscape.<ref>{{Cite journal |last1=Dornak |first1=L. Lynnette |last2=Barve |first2=Narayani |last3=Peterson |first3=A. Townsend |date=February 2013 |title=Spatial Scaling of Prevalence and Population Variation in Three Grassland Sparrows |journal=The Condor |volume=115 |issue=1 |pages=186–197 |doi=10.1525/cond.2013.120055 |hdl=1808/13338 |s2cid=53584537 |issn=0010-5422|doi-access=free |hdl-access=free }}</ref> Return rates of adult grasshopper sparrows to their previous breeding sites (site fidelity) differs widely between populations.<ref name=":7">{{Cite taxon|BOW|last=Vickery |first=Peter D. |date=2020 |title=Grasshopper Sparrow (''Ammodramus savannarum'') |version=1.0 |url=https://birdsoftheworld.org/bow/species/graspa/cur/introduction}}</ref> Site fidelity is far higher in eastern parts of the breeding range, with >50% of adult males returning to breed in subsequent seasons at a site in Connecticut<ref>{{cite thesis|author=Crossman TI|year= 1989|title= Habitat use by Grasshopper and Savannah Sparrow at Bradley International Airport and management recommendations|degree= PhD |location=Storrs, CT|publisher= University of Connecticut}}</ref> and over 70% in Maryland.<ref>{{Cite journal |last1=Soha |first1=Jill A. |last2=Lohr |first2=Bernard |last3=Gill |first3=Douglas E. |date=2009-06-01 |title=Song development in the grasshopper sparrow, ''Ammodramus savannarum'' |url=https://www.sciencedirect.com/science/article/pii/S0003347209001213 |journal=Animal Behaviour |language=en |volume=77 |issue=6 |pages=1479–1489 |doi=10.1016/j.anbehav.2009.02.021 |s2cid=18372807 |issn=0003-3472|url-access=subscription }}</ref> In contrast, 0% of adult males in Nebraska returned to breed in subsequent years,<ref name="Kaspari 1988 792–793">{{Cite journal |last1=Kaspari |first1=Michael |last2=O'Leary |first2=Helen |date=1988 |title=Nonparental Attendants in a North-Temperate Migrant |url=https://www.jstor.org/stable/4087399 |journal=The Auk |volume=105 |issue=4 |pages=792–793 |jstor=4087399 |issn=0004-8038}}</ref> 8.9% returned in Montana,<ref name=":4" /> and 20% returned in California.<ref name=Collier1994>{{cite thesis|author= Collier, Christine Lynn|title= Habitat selection and reproductive success of the Grasshopper Sparrow at the Santa Rosa Plateau Ecological Reserve |degree= MS|publisher= San Diego State University|year= 1994}}</ref> In northeastern Kansas, individual birds also commonly disperse within seasons between nesting attempts.<ref name=":6">{{Cite journal |last1=Williams |first1=Emily J. |last2=Boyle |first2=W. Alice |date=January 2018 |title=Patterns and correlates of within-season breeding dispersal: A common strategy in a declining grassland songbird |journal=The Auk |volume=135 |issue=1 |pages=1–14 |doi=10.1642/auk-17-69.1 |s2cid=90295307 |issn=0004-8038|doi-access=free }}</ref> Between 30 and 75% of birds move over 100 m within season, and individuals were detected defending new territories or nesting up to 8.9&nbsp;km from areas they occupied earlier in the season.<ref name=":6" /> Within-season breeding dispersal decisions relate to nest success; more birds moved following nest failure, and birds that dispersed experienced lower brood parasitism and higher nesting success following movements.<ref>{{Cite journal |last1=Williams |first1=Emily J. |last2=Boyle |first2=W. Alice |date=2019-09-01 |title=Causes and consequences of avian within-season dispersal decisions in a dynamic grassland environment |journal=Animal Behaviour |language=en |volume=155 |pages=77–87 |doi=10.1016/j.anbehav.2019.06.009 |s2cid=199544896 |issn=0003-3472|doi-access=free }}</ref> Little is known about the movements of birds during winter, but inter-annual variation in abundance in northern Mexican grasslands is positively related to related to broad-scale variation in vegetation, rainfall the previous summer, and plant productivity.<ref name="Macías-Duarte 41–49" />

=== Breeding ===
Grasshopper sparrows are a [[socially monogamous]] species,<ref name=":7" /> but rates of [[Extra-pair copulation|extra-pair mating]] can be high (e.g., 48.5% of nests contained an extra-pair young in Kansas<ref name=":9" />), and even instances of [[Cooperative breeding|cooperative parenting]]<ref name="Kaspari 1988 792–793"/> have been documented. In the spring, males select and defend [[Territory (animal)|territories]] before being joined by a female. Over the course of the [[breeding season]], birds can raise multiple broods, either with the same mate or a different mate. In the more southern regions of the species range, pairs may produce 2–4 broods while pairs further north are limited by shorter summers leading to 1–2 nesting attempts.<ref name=":7" />

'''Nest building'''
[[File:Nest from 20 cm away.jpg|thumb|Well-concealed grasshopper sparrow nest showing domed structure and side entrance]]
Females construct [[Bird nest|nests]] over the course of a couple of days.<ref>{{cite book |last1=Harrison |first1=H. H. |title=A Field Guide to Birds' Nests of 285 Species Found Breeding in the United States East of the Mississippi River |date=1975 |publisher=Houghton Mifflin Company |location=Boston, MA, USA}}</ref> Grasshopper sparrows build inconspicuous, dome-shaped nests on the ground, typically very well hidden amongst [[grasses]] and [[forb]]s.<ref>{{cite book |last1=Harrison |first1=C. J. O. |title=A Field Guide to the Nests, Eggs and Nestlings of North American Birds |date=1978 |publisher=Collins |location=Toronto, ON, Canada}}</ref> The nests have a small side opening and are usually made of a mix of dead and live grasses. Nests are built on the breeding pair's territory in areas away from shrubs and trees.

'''Offspring'''
[[File:Newly-hatched Grasshopper Sparrow.jpg|thumb|Nestling grasshopper sparrow within an hour or so of hatching (handled as part of permitted research) at the Konza Prairie, Kansas.]]
Females typically lay between 4–5 eggs per [[Clutch (eggs)|clutch]].<ref>{{cite journal |last1=McNair |first1=D. B. |title=Egg data slips-are they useful for information on egg-laying dates and clutch size |journal=Condor |date=1987 |volume=89 |issue=2 |pages=369–376 |doi=10.2307/1368490|jstor=1368490 }}</ref> Grasshopper sparrow eggs are smooth and oval-shaped. They are a cream-white color, with reddish-brown speckling that is concentrated towards the larger end of the egg.<ref name=":7" /> Females [[Egg incubation|incubate]] the eggs from 10 to 12 days, and nestlings [[fledge]] after 6–9 days.<ref name=":7" /> For the first few days of the nestling period, females primarily feed young, but both parents feed during the latter half of the nestling period. In one study, nonparental helpers were documented attending to broods<ref>{{cite journal |last1=Kaspari |first1=M. |last2=O'Leary |first2=H. |title=Nonparental attendants in a north-temperate migrant |journal=Auk |date=1988 |volume=105 |issue=4 |pages=792–793 |doi=10.1093/auk/105.4.792|doi-broken-date=1 November 2024 |jstor=4087399 }}</ref> and feeding young, but this behavior is apparently not ubiquitous.<ref name=":9" />  Nestlings are fed an [[arthropod]] rich diet, particularly prey ~15–40&nbsp;mm long, and preferentially select [[Acrididae|acridid]] grasshoppers, [[spider]]s, and small [[beetle]]s.<ref>{{Cite journal |last1=Kaspari |first1=Michael |last2=Joern |first2=Anthony |date=1993 |title=Prey Choice by Three Insectivorous Grassland Birds: Reevaluating Opportunism |url=https://www.jstor.org/stable/3544909 |journal=Oikos |volume=68 |issue=3 |pages=414–430 |doi=10.2307/3544909 |jstor=3544909 |bibcode=1993Oikos..68..414K |issn=0030-1299|url-access=subscription }}</ref> Upon leaving the nest, the offspring often remain in the area. Parents provide post-fledging [[Parental care in birds|care]], but the duration and level of care is unknown.
[[File:Grasshopper sparrow nest with four cowbird eggs & one host egg.jpg|thumb|Grasshopper sparrow nest with four cowbird eggs & one host egg]]
'''Brood parasitism'''

Grasshopper sparrows are a common host of the [[brown-headed cowbird]], a [[brood parasite]] which lays their eggs in the nests of other species. Cowbirds can be detrimental to host species because they often remove host eggs and nestlings or destroy nests to initiate another nesting attempt, allowing for future parasitism. However, they do not necessarily reduce the chances of nests successfully fledging young. The rate of brood parasitism by brown-headed cowbirds varies across the grasshopper sparrow's range and depends on habitat characteristics such as the amount of woody vegetation and whether the land is grazed.<ref>{{cite journal |last1=Patten |first1=M. A. |last2=Schochat |first2=E. |last3=Reinking |first3=D. L. |last4=Wolfe |first4=D. H. |last5=Sherrod |first5=S. K. |title=Habitat Edge, Land Management, And Rates Of Brood Parasitism In Tallgrass Prairie |journal=Ecological Applications |date=2006 |volume=16 |issue=2 |pages=687–695 |doi=10.1890/1051-0761(2006)016[0687:HELMAR]2.0.CO;2|pmid=16711055 }}</ref> In surveyed populations, 2–65% of nests were parasitized by cowbirds.<ref name=":7" />

'''Causes of nest failure'''

Only 30–50% of nests successfully raise at least one or more nestlings that fledge the nest.<ref name=":7" /> The majority of grasshopper sparrow nests fail as the result of [[predation]].<ref>{{cite journal |last1=Perkins |first1=D. W. |last2=Vickery |first2=P. D. |last3=Shriver |first3=W. G. |title=Spatial Dynamics of Source-Sink Habitats: Effects on Rare Grassland Birds |journal=The Journal of Wildlife Management |date=2003 |volume=67 |issue=3 |pages=588–599 |doi=10.2307/3802716|jstor=3802716 }}</ref> Snakes, small mammals such as rats, larger mammals such as skunks, armadillos, hogs, and opossums, and even ants have been observed eating eggs and nestlings.<ref>{{cite journal |last1=Hewett Ragheb |first1=E. L. |last2=Miller |first2=K. E. |last3=Hoerl Leone |first3=E. |title=Exclosure fences around nests of imperiled Florida Grasshopper Sparrows reduce rates of predation by mammals |journal=Journal of Field Ornithology |date=2019 |volume=90 |issue=4 |pages=309–324 |doi=10.1111/jofo.12310|s2cid=209591864 }}</ref> Other causes of nest failure include flooding of the nest cup due to rainfall, trampling by livestock or humans, and abandonment by the parents.<ref>{{cite journal |last1=Hovick |first1=T. J. |last2=Miller |first2=J. R. |last3=Dinsmore |first3=S. J. |last4=Engle |first4=D. M. |last5=Debinski |first5=D. M. |last6=Fuhlendorf |first6=S. D. |title=Effects of fire and grazing on grasshopper sparrow nest survival |journal=The Journal of Wildlife Management |date=2012 |volume=76 |issue=1 |pages=19–27 |doi=10.1002/jwmg.243|bibcode=2012JWMan..76...19H }}</ref>

==Vocalizations==
[[File:Grasshopper Sparrow Pawnee National Grasslands CO 2018-06-07 11-48-28 (40892675193).jpg|thumb|right|Grasshopper sparrow singing]]

=== Primary song ===
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The grasshopper sparrow's "primary" or "buzz" song has one to four introductory notes followed by a long high pitched trill,<ref name=":0">{{Cite journal |last1=Bent |first1=Arthur Cleveland |last2=Austin |first2=Oliver L. |date=1968 |title=Life Histories of North American Cardinals, Grosbeaks, Buntings, Towhees, Finches, Sparrows, and Allies |url=https://repository.si.edu/handle/10088/10027 |journal=Bulletin of the United States National Museum |language=en |issue=237 |pages=1–1889 |doi=10.5479/si.03629236.237.1|hdl=10088/10027 |url-access=subscription }}</ref><ref name=":10">{{Cite journal |last1=Lohr |first1=Bernard |last2=Ashby |first2=Sarah |last3=Wakamiya |first3=Sarah M. |date=2013 |title=The function of song types and song components in Grasshopper Sparrows (''Ammodramus savannarum''). |url=https://userpages.umbc.edu/~blohr/Lohr_etal%20Behaviour%202013.pdf |journal=Behaviour |volume=150 |issue=9–10 |pages=1085–1106|doi=10.1163/1568539X-00003094 }}</ref>'tup zeeee' or 'tip tup zeee', and because of its similarity to a grasshopper sound, accounts for the name of the bird.<ref name=":1">{{Cite journal |last=Smith |first=Robert L. |date=1959 |title=The Songs of the Grasshopper Sparrow |journal=The Wilson Bulletin (Wilson Ornithological Society) |volume=71 |issue=2 |pages=141–152}}</ref> There are small variations in the song between individuals, populations, and subspecies.<ref name=":0" /><ref name=":10" /> This song is primarily used for defense, territory, or advertising to the other males and females.<ref name=":2">{{Cite journal |last1=Soha |first1=Jill A. |last2=Lohr |first2=Bernard |last3=Gill |first3=Douglas E. |date=2009 |title=Song development in the grasshopper sparrow, ''Ammodramus savannarum'' |url=https://linkinghub.elsevier.com/retrieve/pii/S0003347209001213 |journal=Animal Behaviour |language=en |volume=77 |issue=6 |pages=1479–1489 |doi=10.1016/j.anbehav.2009.02.021|s2cid=18372807 |url-access=subscription }}</ref> The males will mainly use the primary song from mid-April, when they get to the breeding ground, to mid-August, when they are about to leave the breeding grounds.<ref name=":1" /> It is the principal song heard early in the season, and during each breeding cycle, and the only song that unpaired birds sing.<ref name=":10" /> Birds sing from a grass or forb stalks, poles, fence posts or fence lines, or low shrubs.<ref name=":1" />

=== Alternate song ===
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Male grasshopper sparrows sing "alternate", warble", or "sustained", songs during the breeding season to establish and maintain pair bonds.<ref name=":1" /> The alternate song is more musical than the primary song and is usually delivered from fixed perches or in flight.<ref name=":3">{{Cite journal |last=Walkinshaw |first=L. H. |date=1940 |title=Some notes on the Michigan grasshopper sparrow |journal=The Jack Pine Warbler |volume=18 |pages=50–59}}</ref> The song consists of a 5–15 second series of short notes varying slightly in pitch,<ref>{{Cite book |last=Saunders |first=A. A. |title=A guide to bird songs |publisher=Doubleday and Company |year=1935 |location=Garden City, NY, USA}}</ref> and the entire sequence may be repeated two to four times.<ref name=":2" /> Paired males will start singing alternate song ~5 days after arrival on territory, with frequency of alternate song increasing as birds pair.<ref name=":1" /> The alternate song diminishes towards late-July as the breeding season winds down. Considerable variation in alternate song exists within and between populations and subspecies; further study is needed.<ref name=":10" />

=== Male trill ===
The trill of a male grasshopper sparrow (ti-tu-ti-tu-ti-i-i-i-i) is one of his least common vocalizations and is difficult to detect.<ref name=":0" /> Sometimes called the nesting song, it consists of short, rapid notes and a downward trill.<ref name=":0" /> The song is usually used after two grasshopper sparrows form a pair and is mostly made in response to the female.<ref name=":0" /> The male trills near the nest either from the ground or a perch.<ref name=":0" /> The male trill functions to strengthen pair bonds and to tell the female/young the male is approaching the nest.<ref name=":0" />

=== Female trill ===
The female trill (ti-ti-i-i-i-i-i) is used to announce presence in male's territory.<ref name=":1" /> The primary functions of the female trill is to declare her presence to the male, maintain the pair bond, and signal to both the male and the young that she is approaching the nest.<ref name=":3" /><ref name=":1" /> Given the high rate of extra-pair copulations in this species,<ref name=":9" /> the trill may also function to announce her presence to a territorial male other than the social male. Females usually call from the ground, concealed in grass, and may call independently or in response to primary or sustained song. Females call from pair formation until the end of nesting.<ref name=":1" />

=== Call/chip ===
The call or chip is used by both sexes and is used as an alarm note.<ref name=":0" /> Depending on its intensity, the sound varies. High intensity alarms resemble a slow clicking, and low intensity alarms are a sharp 'tik'.<ref name=":0" /> The chip is used around the nest by both the female and male using single and double 'chip-chip' calls.<ref name=":3" /> This call can be used during feeding when they utter a single, high-pitched chip.<ref name=":0" />

=== Song learning ===
Captive-rearing experiments revealed that grasshopper sparrows are predisposed to learn two distinct songs but must hear species-typical songs to develop a normal sounding song.<ref name=":2" /> Grasshopper sparrows can learn song from pre-recorded tutors but are more inclined to learn songs that are similar to live tutors, suggesting they use information from song models but do not directly imitate conspecific song.<ref name=":2" /> It is likely that in the case of alternate songs, exact notes are imitated but sequences are invented or improvised.<ref name=":2" /> This may be beneficial for pair bonding if females identify individual males by their distinct alternate song.

== Conservation ==

=== Current status ===
The global population of grasshopper sparrows in 2016 was estimated to be around 31,000,000 by the PIF North America Landbird Conservation Plan. It was also estimated that the population had undergone a 68% decrease between the years 1970 and 2014 with an annual decline of about 2.59%. The 2022 State of the Birds reported a long-term, range-wide decline of 2.13% per year, and a more recent decline of 3.48% per year.<ref name=SOB2022>{{Cite web |title=State of the Birds 2022 |url=https://www.stateofthebirds.org/2022 |access-date=2022-11-29 |website=State of the Birds 2022 |language=en-US}}</ref> The leading cause of population decline across its range is linked to habitat loss and management, particularly conversion of grasslands to intensive agriculture and encroachment of shrubs and trees. Despite declining population sizes, grasshopper sparrows are classified as [[Least Concern]] by the [[IUCN]] and are also not included as a "Bird of Conservation Concern" by the U.S. Fish and Wildlife Services (USFWS) on a national level. The [[Florida grasshopper sparrow]] subspecies (''Ammodramus savannarum floridanus'') is classified as endangered by  the USFW.<ref>{{Cite web |title=ECOS: Species Profile |url=https://ecos.fws.gov/ecp/species/32 |access-date=2022-11-16 |website=ecos.fws.gov}}</ref> This subspecies hit an all time population low of 15 breeding pairs in 2017, and reported a population of 120 in 2022.<ref>{{Cite web |title=Florida Grasshopper Sparrow |url=https://www.whiteoakwildlife.org/wildlife/florida-grasshopper-sparrow/ |access-date=2022-11-16 |website=White Oak Conservation |language=en-US}}</ref>

==== Habitat management ====
The majority of the former breeding range of grasshopper sparrows in North America has been converted to intensive agriculture<ref>{{Citation |last=Tivy |first=Joy |title=Grassland ecosystems |url=https://www.taylorfrancis.com/chapters/mono/10.4324/9781315845227-11/grassland-ecosystems-joy-tivy |work=Biogeography |year=2018 |pages=212–227 |doi=10.4324/9781315845227-11 |isbn=9781315845227 |s2cid=240017487 |access-date=2022-11-16|url-access=subscription }}</ref> or encroached by shrubs and trees.<ref>{{Cite web |title=The effects of management practices on grassland birds—Grasshopper Sparrow (''Ammodramus savannarum'') {{!}} U.S. Geological Survey |url=https://www.usgs.gov/publications/effects-management-practices-grassland-birds-grasshopper-sparrow-ammodramus-savannarum |access-date=2022-11-16 |website=www.usgs.gov |language=en}}</ref> Likewise, these birds are experiencing a loss of wintering habitats, particularly within the Chihuahuan grasslands, due to similar impacts (conversion to agriculture and woody encroachment).<ref>{{Cite journal |last1=Pool |first1=Duane B. |last2=Panjabi |first2=Arvind O. |last3=Macias-Duarte |first3=Alberto |last4=Solhjem |first4=Deanna M. |date=2014-02-01 |title=Rapid expansion of croplands in Chihuahua, Mexico threatens declining North American grassland bird species |url=https://www.sciencedirect.com/science/article/pii/S0006320713004400 |journal=Biological Conservation |language=en |volume=170 |pages=274–281 |doi=10.1016/j.biocon.2013.12.019 |bibcode=2014BCons.170..274P |issn=0006-3207|url-access=subscription }}</ref> Because grasshopper sparrows require areas of dense dead grass for nesting, sparse vegetation for foraging, and little to no woody plants, they are vulnerable to management practices such as high-intensity grazing accompanied by annual burning, fire suppression leading to woody encroachment, and haying that can destroy nests if it occurs during the breeding season. Ideal grazing intensities and fire frequencies vary across their range depending on climate, but areas with moderate grazing by cattle or bison, prescribed burning every 2–3 years, and removal of woody plants tend to support the highest densities of grasshopper sparrows in the Southern great plains.<ref>{{Cite journal |last=Powell |first=Alexis F. L. A. |date=March 2008 |title=Responses of breeding birds in tallgrass prairie to fire and cattle grazing |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1557-9263.2008.00144.x |journal=Journal of Field Ornithology |language=en |volume=79 |issue=1 |pages=41–52 |doi=10.1111/j.1557-9263.2008.00144.x |issn=0273-8570|url-access=subscription }}</ref>

==Gallery==
<gallery>
File:Grasshopper Sparrow, Gerald R. Ford Int'l airport, 13 July 2014 (14691040876).jpg|Grasshopper sparrow
File:Grasshopper Sparrow Pawnee National Grasslands CO 2018-06-06 17-33-21 (40892558283).jpg|Grasshopper sparrow flying
File:Juvenile Grasshopper Sparrow in hand.jpg|Juvenile grasshopper sparrow
</gallery>

== References ==
<!-- WilsonBull18:47 (compare to current Ohio checklist http://www.ohiobirds.org/publications/OBRClist.pdf) -->
{{Reflist|refs=
<ref name=CRC>{{cite book |title=CRC Handbook of Avian Body Masses |editor-first=John B. |editor-last=Dunning Jr. |publisher=CRC Press |year=1992 |isbn=978-0849342585}}</ref>

<ref name=DEC>{{cite web |url=http://www.dec.ny.gov/animals/59577.html |title=Grasshopper Sparrow Fact Sheet |publisher= [[New York State Department of Environmental Conservation]]}}</ref>
}}

== External links ==
{{Commons category|Ammodramus savannarum}}
{{Wikispecies|Ammodramus savannarum}}
* [http://www.birds.cornell.edu/AllAboutBirds/BirdGuide/Grasshopper_Sparrow.html Grasshopper sparrow Species Account] - Cornell Lab of Ornithology
* [http://www.mbr-pwrc.usgs.gov/id/framlst/i5460id.html Grasshopper sparrow - ''Ammodramus savannarum''] - USGS Patuxent Bird Identification InfoCenter
* {{VIREO|grasshopper+sparrow}}
* {{BirdLife|22721144|Ammodramus savannarum}}
* {{Avibase|name=Ammodramus savannarum}}
* {{InternetBirdCollection|grasshopper-sparrow-ammodramus-savannarum|Grasshopper sparrow}}
* {{IUCN_Map|22721144/138486868|Ammodramus savannarum}}
* {{Xeno-canto species|Ammodramus|savannarum|Grasshopper sparrow}}

<!--
==Further reading==
===Book===

* Vickery, P. D. 1996. ''Grasshopper Sparrow (Ammodramus savannarum)''. In ''The Birds of North America'', No. 239 (A. Poole and F. Gill, eds.). The Academy of Natural Sciences, Philadelphia, PA, and The American Ornithologists’ Union, Washington, D.C.

===Report===

* Cannings RJ. (1995). ''Status of the grasshopper sparrow in British Columbia''. Canadian Research Index. p. n/a.

===Theses===

* Ahlering MA. Ph.D. (2005). ''Settlement cues and resource use by Grasshopper Sparrows and Baird's Sparrows in the Upper Great Plains''. University of Missouri - Columbia, United States, Missouri.
* Ammer FK. Ph.D. (2003). ''Population level dynamics of grasshopper sparrow populations breeding on reclaimed mountaintop mines in West Virginia''. West Virginia University, United States, West Virginia.
* Dillery DG. Ph.D. (1961). ''FOOD HABITS OF SAVANNAH AND GRASSHOPPER SPARROWS IN RELATION TO FOODS AVAILABLE''. The Ohio State University, United States, Ohio.
* Gamble K. M.S. (2005). ''Habitat use in Baird's and grasshopper sparrows''. University of Missouri - Columbia, United States, Missouri.

===Articles===

* Arguedas-Negrini N. (2001). ''Distribution, habitat and behavior of grasshopper sparrows, Ammodramus savannarum (Passeriformes : Emberizidae) in northeastern Nicaragua''. Revista De Biologia Tropical. vol '''49''', no 2. pp.&nbsp;703–707.
* Balent KL & Norment CJ. (2003). ''Demographic characteristics of a Grasshopper Sparrow population in a highly fragmented landscape of western New York State''. Journal of Field Ornithology. vol '''74''', no 4. pp.&nbsp;341–348.
* Behrend FW. (1973). ''Occurrences of the Grasshopper Sparrow Ammodramus-Savannarum at High Altitude on a Southern Appalachian Grass Bald''. Journal of the Tennessee Academy of Science. vol '''48''', no 2.
* Bulgin NL, Gibbs HL, Vickery P & Baker AJ. (2003). ''Ancestral polymorphisms in genetic markers obscure detection of evolutionarily distinct populations in the endangered Florida grasshopper sparrow (Ammodramus savannarum floridanus)''. Molecular Ecology. vol '''12''', no 4. pp.&nbsp;831–844.
* Cortelyou RG. (1970). ''Grasshopper Sparrow Actions''. Nebraska Bird Review. vol '''38''', no 4.
* Delany MF & Cox JA. (1986). ''Florida USA Grasshopper Sparrow Breeding Distribution and Abundance in 1984''. Florida Field Naturalist. vol '''14''', no 4. pp.&nbsp;100–104.
* Delany MF, Giesel JT & Brazeau DA. (2000). ''Genetic variability among populations of the Florida grasshopper sparrow''. Journal of Wildlife Management. vol '''64''', no 3. pp.&nbsp;631–636.
* Delany MF & Linda SB. (1994). ''Characteristics of occupied and abandoned Florida grasshopper sparrow territories''. Florida Field Naturalist. vol '''22''', no 4. pp.&nbsp;106–109.
* Delany MF & Linda SB. (1998). ''Characteristics of Florida Grasshopper Sparrow nests''. Wilson Bulletin. vol '''110''', no 1. pp.&nbsp;136–139.
* Delany MF, Stevenson HM & McCracken R. (1985). ''Distribution Abundance and Habitat of the Florida Grasshopper Sparrow Ammodramus-Savannarum-Floridanus''. Journal of Wildlife Management. vol '''49''', no 3. pp.&nbsp;626–631.
* Herkert JR. (1998). ''The influence of the CRP on grasshopper sparrow population trends in the mid-continental United States''. Wildlife Society Bulletin. vol '''26''', no 2. pp.&nbsp;227–231.
* Joern A. (1988). ''Foraging Behavior and Switching by the Grasshopper Sparrow Ammodramus-Savannarum Searching for Multiple Prey in a Heterogeneous Environment''. American Midland Naturalist. vol '''119''', no 2. pp.&nbsp;225–234.
* Joern A. (2002). ''Context-dependent foraging and enemy-free space: Grasshopper sparrows (Ammodramus savannarum) searching for grasshoppers (Acrididae)''. Ecoscience. vol '''9''', no 2. pp.&nbsp;231–240.
* Kaspari M. (1991). ''PREY PREPARATION AS A WAY THAT GRASSHOPPER SPARROWS (AMMODRAMUS-SAVANNARUM) INCREASE THE NUTRIENT CONCENTRATION OF THEIR PREY''. Behavioral Ecology. vol '''2''', no 3. pp.&nbsp;234–241.
* Lohr, B., * Ashby, S., & Wakamiya, S.M. (2013). The function of song types and song components in Grasshopper Sparrows (Ammodramus savannarum). Behaviour 150: 1085-1106.
* McNair DB. (1986). ''Clutch Information for the Florida Grasshopper Sparrow from Oological Collections''. Florida Field Naturalist. vol '''14''', no 2. pp.&nbsp;48–49.
* Miller P. (2005). ''Long distance dispersal of a Florida grasshopper sparrow''. Florida Field Naturalist. vol '''33''', no 4. pp.&nbsp;123–124.
* Olson SL. (1980). ''The Subspecies of Grasshopper Sparrow Ammodramus-Savannarum in Panama Aves Emberizinae''. Proceedings of the Biological Society of Washington. vol '''93''', no 3. pp.&nbsp;757–759.
* Perkins DW & Vickery PD. (2001). ''Annual survival of an endangered passerine, the Florida Grasshopper Sparrow''. Wilson Bulletin. vol '''113''', no 2. pp.&nbsp;211–216.
* Perkins DW & Vickery PD. (2005). ''Effects of altered hydrology on the breeding ecology of the Florida Grasshopper Sparrow and Bachman's Sparrow''. Florida Field Naturalist. vol '''33''', no 2. pp.&nbsp;29–40.
* Perkins DW, Vickery PD, Dean TF & Scheuerell MD. (1998). ''Florida grasshopper sparrow reproductive success based on nesting records''. Florida Field Naturalist. vol '''26''', no 1. pp.&nbsp;7–17.
* Pranty B. (2000). ''Three sources of Florida Grasshopper Sparrow mortality''. Florida Field Naturalist. vol '''28''', no 1. pp.&nbsp;27–29.
* Shriver WG & Vickery PD. (1999). ''Aerial assessment of potential Florida Grasshopper Sparrow habitat: Conservation in a fragmented landscape''. Florida Field Naturalist. vol '''27''', no 1. pp.&nbsp;1–9.
* Whitmore RC. (1979). ''Short-Term Change in Vegetation Structure and Its Effect on Grasshopper Sparrows Ammodramus-Savannarum in West-Virginia USA''. Auk. vol '''96''', no 3. pp.&nbsp;621–625.
* Whitmore RC. (1981). ''Structural Characteristics of Grasshopper Sparrow Ammodramus-Savannarum Habitat''. Journal of Wildlife Management. vol '''45''', no 3. pp.&nbsp;811–814.
* Wiens JA. (1971). ''Egg Dumping by the Grasshopper Sparrow in a Savannah Sparrow Nest''. Auk. vol '''88''', no 1. pp.&nbsp;185–186.
-->

{{Taxonbar|from=Q1588210}}

[[Category:Ammodramus|grasshopper sparrow]]
[[Category:Birds of North America]]
[[Category:Birds of the United States]]
[[Category:Birds of the Greater Antilles]]
[[Category:Birds of Jamaica]]
[[Category:Birds of the Dominican Republic]]
[[Category:Birds of Haiti]]
[[Category:Birds of Puerto Rico]]
[[Category:Birds of Mexico]]
[[Category:Birds of Belize]]
[[Category:Birds of Costa Rica]]
[[Category:Birds of Panama]]
[[Category:Birds described in 1789|grasshopper sparrow]]
[[Category:Taxa named by Johann Friedrich Gmelin|grasshopper sparrow]]