Family Larks (Alaudidae)

Least Concern

Horned Lark (Eremophila alpestris)

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Taxonomy

French: Alouette hausse-col German: Ohrenlerche Spanish: Alondra cornuda
Taxonomy:

Alauda alpestris

Linnaeus

, 1758,

coast of South Carolina, USA

.

See E. bilopha. Numerous races described over vast range, based mainly on differences in size, ground colour (partly determined by local soil colour) and pattern; recent molecular study suggested taxa in Old World break into five species (excluding E. bilopha), E. elwesi (although unsampled longirostris has priority if it belongs to this clade), E. atlas, E. penicillata, E. brandti and E. flava#R, but sampling incomplete making confident allocation of other taxa to these species impossible; moreover, little consistent fit of morphological characters to proposed species, and vocal analysis reveals no marked differences#R. Race teleschowi sometimes treated as separate species, apparently not interbreeding with other races where ranges overlap. Race kumerloevei perhaps synonymous with penicillata#R. Review of material in NHMUK here suggests a reduction of New World subspecies by synonymising praticola with nominate; enthymia with leucolaema; alpina, merrilli, utahensis and lamprochroma with strigata; sierrae with rubea; actia, ammophila and leucansiptila with occidentalis; lactea and aphrasta with adusta; and oaxacae and diaphora with chrysolaema. Twenty-eight subspecies recognized.

Subspecies and Distribution
  • E. a. arcticola (Oberholser, 1902) – Alaska and NW Canada (Yukon S to British Columbia); winters in W USA.
  • E. a. hoyti (Bishop, 1896) – N Canada (N Baffin I S to N Alberta and W Ontario); winters in N USA.
  • E. a. alpestris (Linnaeus, 1758) – American Horned Lark – E Canada (W Ontario E to Newfoundland and S to Nova Scotia) and E USA (E from Minnesota and S to Kansas and N Carolina); N populations winter in E USA.
  • E. a. strigata (Henshaw, 1884) – W & WC Canada and USA (British Columbia S to N California and E to Idaho, Nevada and Utah); N populations winter in W USA.
  • E. a. leucolaema Coues, 1874 – SW & SC Canada (S Alberta E to Manitoba) S to WC & SC USA (S to New Mexico E to Texas).
  • E. a. rubea (Henshaw, 1884) – NE & C California.
  • E. a. insularis (Dwight, 1890) – Channel Is, off SW California.
  • E. a. occidentalis (McCall, 1851) – SW USA (S California and SW Nevada E to C New Mexico) and NW Mexico (N Baja California and NW Sonora).
  • E. a. adusta (Dwight, 1890) – S Arizona E to S New Mexico S in Mexico to Durango and E to Coahuila.
  • E. a. enertera (Oberholser, 1907) – C Baja California.
  • E. a. giraudi (Henshaw, 1884) – coastal S USA (S Texas) S to NE Mexico.
  • E. a. chrysolaema (Wagler, 1831) – S Mexican Plateau from SE Coahuila and Zacatecas S to Jalisco, Michoacán, Morelos, Veracruz and NE Puebla; also C Oaxaca.
  • E. a. peregrina (P. L. Sclater, 1855) – E Andes of Colombia (Altiplano Cundiboyacense, N of Bogotá).
  • E. a. flava (J. F. Gmelin, 1789) – Shore Lark – N Eurasia E to NE Russia (Anadyrland), S to S Norway, L Baikal and NW Amurland; winters in W & C Europe E to Kazakhstan, Mongolia and C China.
  • E. a. brandti (Dresser, 1874) – Steppe Horned Lark – SE European Russia (lower R Volga) and N Transcaspia E to NE China (Inner Mongolia), S to N Turkmenistan, Tien Shan and Mongolia; N populations migrate S.
  • E. a. atlas (Whitaker, 1898) – Atlas Horned Lark – Morocco (Atlas Mts).
  • E. a. balcanica (Reichenow, 1895) – SE Europe (S Balkans and Greece).
  • E. a. kumerloevei Roselaar, 1995 – W & C Asia Minor.
  • E. a. penicillata (Gould, 1838) – Caucasian Horned Lark – E Turkey and Caucasus E to N & W Iran.
  • E. a. bicornis (C. L. Brehm, 1842) – Lebanon and N Israel–S Syria border (Mt Hermon).
  • E. a. albigula (Bonaparte, 1850) – SW Turkmenistan and NE Iran E to W Tien Shan and S to NW Pakistan.
  • E. a. argalea (Oberholser, 1902) – N Ladakh, extreme W China (W Kunlun Shan) and W Tibetan Plateau.
  • E. a. teleschowi (Przevalski, 1887) – C & E Kunlun Shan from S Xinjiang E to NW Qinghai and S to N Xizang (W China).
  • E. a. przewalskii (Bianchi, 1904) – Qaidam Basin, in N Qinghai.
  • E. a. nigrifrons (Przevalski, 1876) – mountains of NE Qinghai.
  • E. a. longirostris (F. Moore, 1856) – Himalayan Horned Lark – Himalayas from NE Pakistan and Kashmir E to Sikkim.
  • E. a. elwesi (Blanford, 1872) – S & E Tibetan Plateau.
  • E. a. khamensis (Bianchi, 1904) – SC China (SE Xizang, W Sichuan).
  • Descriptive notes

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp alpestris   American Horned Lark

    Drawing by Tim Worfolk
    ssp alpestris   American Horned Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    Drawing by Tim Worfolk
    ssp alpestris   American Horned Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp strigata  

    Drawing by Tim Worfolk
    ssp strigata  
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp rubea  

    Drawing by Tim Worfolk
    ssp rubea  
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp peregrina  

    Drawing by Tim Worfolk
    ssp peregrina  
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp flava   Shore Lark

    Drawing by Tim Worfolk
    ssp flava   Shore Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp brandti   Steppe Horned Lark

    Drawing by Tim Worfolk
    ssp brandti   Steppe Horned Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp atlas   Atlas Horned Lark

    Drawing by Tim Worfolk
    ssp atlas   Atlas Horned Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp penicillata   Caucasian Horned Lark

    Drawing by Tim Worfolk
    ssp penicillata   Caucasian Horned Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    Drawing by Tim Worfolk
    ssp penicillata   Caucasian Horned Lark
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp bicornis  

    Drawing by Tim Worfolk
    ssp bicornis  
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    ssp teleschowi  

    Drawing by Tim Worfolk
    ssp teleschowi  
    Descriptive notes:

    14–17 cm; 30–40 g. Medium-sized lark with short bill, fairly long tail, and striking head pattern with elongated lateral crown feathers (“horns”). Male nominate race has yellow forehead and supercilium, yellow from chin and throat extending across lower neck side to meet yellowish ear-coverts (latter darker and browner at rear), all contrasting with blackish central crown with elongated lateral feathers, blackish lores and broad band from eye down to cheek, and black chestband; central crown to upper mantle, also rump and uppertail-coverts, warm ­rufous-brown with pinkish tinge, rest of upper­parts greyer and dark-streaked, wings somewhat darker with pale edgings; tail blackish, central feathers paler and greyish, outer feathers edged white; underparts below chestband whitish, breast side to rear flank washed pinkish-rufous, breast streaked; bill dark grey to black; legs black. Non-breeding male has head pattern partly obscured (pale fringes), “horns” often shorter (often invisible in field). Female resembles male but smaller, slightly duller, with black bands on head and chest a little narrower. Juvenile has head and upperparts heavily mottled blackish and white, throat and lower neck side creamy whitish, chestband mottled; after autumn moult resembles adult. Races differ mostly in plumage colour and pattern, mainly of males, also in size (migratory races larger, darker and relatively longer-winged than resident ones, those in mesic habitats tend to be larger than those in arid habitats, and size also decreases with altitude): in N America, arctic races (e.g. arcticola) are somewhat darker and slightly less streaked above than nominate, nape pinkish, pale areas of face whitish (not yellow), W races (e.g. merrilli, strigata, rubea) have dark reddish nape to upper mantle and breast side, heavily streaked upperparts and bright yellow on head, those in interior W (e.g. utahensis) are paler and greyer with less streaking, palest and unstreaked in S deserts (adusta), greyer and streaked in S Texas (giraudi); Neotropical peregrina has forehead and supercilium whitish, throat yellow, nape greyish, upperparts dark grey with contrasting pinkish lesser and median coverts; in Eurasia, flava is very like American nominate (but with lesser coverts paler, less deep red), atlas is similar but paler, brandti is also similar but has pale areas on head whitish (not yellow); in most of remaining Old World races black of cheekband meets (or more or less meets) black of chestband, balcanica has yellow on face and throat, dark-streaked grey upperparts, penicillata has paler yellowish-white on head, pink hindneck, pink-tinged greyish-buff upperparts with fine streaks on mantle and scapulars, kumerloevi resembles previous but with unstreaked greyish-pink upperparts, bicornis has pale areas on head even whiter, crown to rump uniform plain sandy buff, tail-coverts tinged cinnamon, other E races generally rather pale, greyish to sandy, e.g. albigula has pale parts of head white, grey central crown and hindneck, finely streaked sandy grey upperparts, longirostris is slightly darker than previous, with black of cheekband and chest separated by white lower neck side, teleschowi is pale with black (not white or yellow) forehead.

    Voice

    Song, from ground or in air, often consists of a few simple, rippling trills followed by short chatter, less fluent than that of many other larks; a second song is more like that of Alauda arvensis but less ebullient and less loud. Usual flight call “eeh” or “ééh-ti”, or liquid “tur-reep”; occasional harsh “tsrr”.

    Habitat

    The only lark to have successfully colonized tundra and alpine habitats; throughout range prefers mainly barren terrain with very short vegetation. In North America, where the only breeding representative of the family, widely distributed across most open habitats, occurring from sea-level to c. 4000 m and from tundra and mountains to steppe, desert and other bare ground, also wide variety of farmland types; shows a preference for recently burned sites; will also feed on sandy beaches and breed in sand dunes; highest densities occur where bare ground predominates, and in grassland and other grazed habitats most numerous in more heavily grazed areas. In Andes, race peregrina found in short-grass pastureland and bare fields, to at least 3100 m, probably higher. In Eurasia, breeds mainly in arctic tundra, dry stony patches in lichen tundra, barren steppes and arctic-alpine zones; breeds up to snow-line, to 5400 m in Himalayas. Also on open coasts and dunes in non-breeding season; in W Europe, migrants of race flava largely confined to open coastal habitats around North Sea in winter.

    Food and feeding

    Wide range of invertebrates in summer and plant material in winter. Main invertebrate prey items are grasshoppers (Acrididae), beetles (Coleoptera), flies (Diptera) and lepidopteran larvae; nestlings fed almost entirely with these invertebrates, also with earthworms (Annelida). Seeds, especially small grass seeds, particularly important in winter, when may comprise entire diet; flocks wintering around North Sea coasts (W Europe) feed largely on energy-rich halophyte seeds found in lower saltmarshes. Forages on ground, singly or in small to large, tightly knit flocks; walks and runs. Obtains most food items directly from soil surface; occasionally digs, pursues flushed prey by running, and takes seeds directly from plants. Occasionally drinks water around dawn.

    Breeding

    In North America, season from mid-Feb in S USA (one of earliest nesters) and from mid-May in Canadian Arctic; in Eurasia, breeds from late May to mid-Jul in Scandinavia but from mid-Jun in arctic Russia; in Colombia, nests with eggs and chicks found in Apr#R; one brood in N of range, 2–3 broods in S. Apparently monogamous; territorial, breeding territory 0·3–5·1 ha depending on habitat. Song-flighting male ascends to great height, up to 250 m, hovers while singing, drops vertically back to earth; generally less aerial than many other larks. Nest, built by female alone, an excavated cavity or natural depression on ground, filled with woven plant material, lined with feathers or other fine material, with stones, bark, clods of earth, animal dung and other material placed around it; nest usually positioned in such a way as to reduce wind flow across it to as little as a tenth of ambient wind speed and to maximize shade. Clutch 1–8 eggs, mostly 2–5, clutch size increasing with increasing latitude; incubation by female alone, period 11–12 days; chicks fed by both parents, leave nest at c. 9–12 days, fly at 16–18 days, reach full adult size at c. 30 days. Success variable; at high altitudes in Asia, productivity in some years reduced to zero by severe cold; drought may also reduce productivity of populations at lower latitudes; nests often parasitized by Brown-headed Cowbird (Molothrus ater) in North America. Recorded longevity at least 8 years.

    Movements

    Migratory or partially migratory in N; mostly resident or altitudinal migrant in S. Across Holarctic Region, races in far N wholly migratory, with much or all of breeding range abandoned in winter (usually by late Oct, earlier in Russia): arcticola and hoyti migrate to W & N USA, nominate to SE USA, and flava to coasts of W Europe (S North Sea, W Baltic Sea), less regularly inland in C & E Europe, and in large numbers across C Asia (S to Sea of Azov, N Caucasus, N Kazakhstan and NE China); returning birds arrive as early as late Feb in S Canada, but not until Apr or early May in Alaska and N Canada, and in Eurasia from late Apr in W (N Norway) and from mid-May in extreme N Russia; males arrive back on breeding grounds before females. Several races are partial or altitudinal migrants in N of respective ranges, e.g. strigata, praticola, enthymia and leucolaema from S Canada move S after breeding, some reaching N Mexico, and those breeding in high mountains of Asia (e.g. albigula, longirostris) descend to lower elevations in winter. In addition, some populations of S USA deserts (ammophila, occidentalis) undertake short-distance movements to NW & N Mexico. Generally in flocks, usually of up to 50 individuals but sometimes to several hundred in hard weather, during migration and outside breeding season. Nominate race recorded as vagrant in UK and Bermuda; flava recorded annually in Japan.

    Status and conservation

    Not globally threatened (Least Concern). Common to locally very common; scarce in some areas. Has one of the largest world ranges of any songbird, and is the only alaudid occurring naturally in the New World (apart from Alauda arvensis, which has recently gained a tenuous foothold in W Alaska). Across much of range, and particularly in North America and arctic Eurasia, one of the commonest bird species in a wide range of open habitats; spread of cultivation led to considerable range expansion in North America at end of 19th century and beginning of 20th century. Global population almost certainly very large, but few reliable data; estimates at end of 1990s include c. 6370–18,560 pairs in Europe (most in Norway), 10,000–100,000 pairs in Turkey and 100,000–1,000,000 pairs in Russia; fairly common in SE Europe, and density in Caucasus up to 4·5–6 birds/km²; locally common in Middle East and N Africa, and common in China. No figures for North America but widespread and common, probably several million pairs; fairly common to common in Mexico. Race peregrina, restricted to Altiplano Cundiboyacense, in Colombian Andes, is rare, and declining rapidly as a result mainly of spread of introduced Kenyan kikuyu grass (Pennisetum clandestinum), unfavourable agricultural practices and unrestricted use of pesticides; in 2002 total of 401 individuals located, and population estimated at fewer than 1000. Some North American races also have small ranges and populations, although trends across the continent appear stable. Declining in some areas, largely because of changes in agriculture; replacement of arable crops by biomass-fuel production, and of conventional tillage by minimum tillage, poses threats in some parts of North America, as does agricultural abandonment; direct poisoning by pesticides has also been recorded, and this species may be particularly susceptible to this risk as in some areas it is a significant pest of crops, and chemical repellents are used to reduce crop damage. Other populations are increasing; for example, deforestation for farmland or airstrips has allowed the species to colonize formerly forested regions across much of North America. In Europe, drastic declines noted since 1950s in N Fennoscandia, especially Finland, believed possibly due to overgrazing of lichen by reindeer (Rangifer tarandus), but increase and range extension recorded in Balkans; expansion in area of coastal marshes in Netherlands as a result of improved flood defences led to increase in numbers of this species wintering there.

    Recommended citation

    Donald, P. & de Juana, E. (2019). Horned Lark (Eremophila alpestris). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (eds.). Handbook of the Birds of the World Alive. Lynx Edicions, Barcelona. (retrieved from https://www.hbw.com/node/57664 on 23 August 2019).