HBW 9 - Greater Short-toed Lark (Calandrella brachydactyla)

69. Greater Short-toed Lark
Calandrella brachydactyla

French: Alouette calandrelle  German: Kurzzehenlerche  Spanish:Terrera Común

Other common names: (European) Short-toed Lark

Taxonomy. Alauda brachydactila [sic] Leisler, 1814, Montpellier, France. May form a superspecies with C. cinerea, with which often merged, and with C. blanfordi, C. erlangeri and C. acutirostris. Geographical variation slight and clinal, generally becoming paler and greyer to E, more rufous and streaked above to W, but greatly complicated by strong influence of wear on plumage, considerable individual variation and lack of natural boundaries; described ranges of listed taxa somewhat arbitrary. Further, races artemisiana, hermonensis and woltersi all intergrade in S Turkey; birds from Ukraine are intermediate between nominate race and longipennis, and those from Ural-Volga steppes, the Caucasus, E Iran and N Afghanistan between latter race and artemisiana. In addition, populations from E Altai and S Siberia have been separated as race orientalis and those from NW Manchuria as puii, but differences from typical longipennis seem to be minor. Validity of race hungarica sometimes disputed, mainly on grounds that tiny population is apparently boosted by periodic influxes from elsewhere. Eight subspecies currently recognized.

Subspecies and Distribution.
C. b. brachydactyla (Leisler, 1814) - breeds Europe from W, NC & S France and Iberia E (including many Mediterranean islands) to Italy, the Balkans and Romania (possibly to Ukraine and S Russia), apparently also in some parts of Mediterranean coast of NW Africa; winters Africa.
C. b. hungarica Horváth, 1956 - breeds Hungary and N Serbia (Vojvodina), possibly also S Slovakia.
C. b. rubiginosa Fromholz, 1913 - breeds NW Africa from Morocco and N Western Sahara E to NW Libya (W Tripolitania); also Malta.
C. b. hermonensis Tristram, 1864 - S Turkey and Syria S to NE Egypt (Sinai Peninsula).
C. b. woltersi Kumerloeve, 1969 - S Turkey (Amik Gölü, Gaziantep, Birecik) and adjacent extreme NW Syria.
C. b. artemisiana Banjkowski, 1913 - breeds Asia Minor, the Caucasus, Transcaucasia and Iran; winters mainly SW Asia.
C. b. longipennis (Eversmann, 1848) - breeds Ukraine and S Russia E to N Mongolia and NW Manchuria; winters mainly S Asia.
C. b. dukhunensis (Sykes, 1832) - breeds Tibet and C China; winters S Asia.

Distribution Calandrella brachydactyla Descriptive notes. 14-15 cm; c. 20-26 g. Small, neat, pale, short-billed lark with long tertials almost cloaking tips of folded wings, narrow and pointed outer primary strongly reduced (7-11 mm short of primary-covert tips), shortish tail; straight hind claw 7-9 mm long. Nominate race has broad whitish supercilium, grey-brown cheeks; warm brown to grey-brown and heavily streaked blackish above, top of head usually densely streaked and often rufous-tinged, with erectile feathers; median coverts with dark centres and buff fringes, contrast with rest of upperwing; tail blackish-brown, centre brown, white outer feathers; throat and underparts whitish, small blackish patch on lower neck side (often obscured in fresh autumn plumage), breast side with faint buff tinge and fine dark streaks (variable, often whitish and unstreaked); undertail blackish (noticeable in flight); bill pale horn-coloured, darker culmen and tip, yellowish on lower mandible; legs pale brownish-flesh to light brown. Sexes similar. Juvenile resembles adult, but upperpart feathers dark brown with buff fringes and whitish tips, outer primary longer (1-6 mm short of primary coverts) and tip more rounded. Races differ mainly in plumage tone above, but greatly affected by influence of wear, and following is only a general guide: hungarica is buffer and greyer than nominate; rubiginosa is more rufous, especially on crown, also more finely streaked; hermonensis is less rufous, more buff, than previous, with crown less contrasting; woltersi resembles last, but paler and greyer; artemisiana is slightly darker than last, more buffish-grey, also more heavily streaked; longipennis is greyish-sandy and finely streaked above, bill relatively small; dukhunensis is distinctive, larger, longer-winged and slightly smaller-billed than other races, dark brown and heavily streaked above, buff-washed below, legs dark. VOICE. Song, in aerial display, less often from ground, clear and musical but rather monotonous, with accelerating simple notes, then several hesitant notes, and finally a long series of persistently repeated phrases each c. 2 seconds long and consisting of 8-10 bubbling notes, pauses of 2-3 seconds between phrases (coinciding with undulations in flight); imitations of other bird species often incorporated. Commonest call a dry "chirrup" or "dreet", often repeated; also "teeoo" or "trilp".
Habitat. Dry areas with sparse and low vegetation cover, on level or undulating terrain, with sandy or stony soils. Breeds mostly in fallow lands in Mediterranean Basin, but also dry pastures, tobacco fields, dirt tracks and olive groves; in Russia also more densely covered pastures, but absent in true steppe; sometimes in semi-arid areas, but avoids true desert. Mainly lowlands; to 1500 m in Tunisia. Occurs in semi-arid areas and farmland in non-breeding areas.
Food and Feeding. Mainly invertebrates during spring, also seeds and green parts of plants in other seasons. Invertebrate food very diverse, variable among areas, chiefly beetles (Coleoptera), ants, bugs (Hemiptera) and snails; seeds mostly of weedy forbs (Polygonum, Amaranthus), also cereal grain. Nestlings fed solely with invertebrates; fledglings take more green material than do adults. Forages on ground, singly or in small or large flocks. Searches slowly, picks items from ground surface.
Breeding. Lays in May-Jul in SW Europe and from mid-Apr in SE Europe; from early Apr in N Africa and Israel; two broods per season. Male song flight typically at heights of c. 30-50 m, undulating and bouncing in a circling but rather meandering path, lasting on average 3-5 minutes. Nest built by female, of grasses, rootlets and similar vegetation, lined with softer material, internal diameter 6 cm, placed in shallow scrape on ground, usually beside shrub or grass tuft, often with small rampart of sticks or stones; orientation in Spain mainly to N. Clutch 2-5 eggs, rarely 6 (mode 3 in N Africa, 4 in Spain, 5 in Hungary), clutch size in Spain larger in middle of season; replacement laid if first clutch lost; incubation by female alone, beginning with last or penultimate egg, period 11-13 days; chicks fed and cared for by both sexes, nestling period 9-12 days, mostly 8-10 days, fledging 12-15 days; dependence period of fledglings 1-3 weeks, shorter for first broods. Nest losses often very high, up to c. 80%; snakes a major predator, and some nests also destroyed by trampling by livestock. First breeding at 1 year.
Movements. Mostly migratory; only partially migratory in S of Palearctic range, and sedentary in Transcaucasia. Those from W & C parts of range winter mainly in sub-Saharan Africa (S Sahel zone) and Red Sea Basin, also in SW Asia; E populations migrate mostly to S & SE parts of Asia; highly gregarious on wintering grounds, occurring in flocks of hundreds or thousands. Autumn departure from mid-Aug through to Sept/Oct, passage on broad front; return begins late Jan, but migration later in E, and arrival on breeding grounds in N of range not until Apr-May; generally common on passage in N Africa, abundant in Egypt, and common to fairly common in Middle East and Arabian Peninsula; in S Israel, up to 14,000 per day recorded in autumn (Sept-Oct) and flock of c. 4000 in Apr. Vagrants regularly W to Scandinavia and British Is, and recorded on Madeira and Canary Is, also N to Iceland and regularly Finland; rare but annual in Japan; in S, recorded to Cameroon, DRCongo and Kenya.
Status and Conservation. Not globally threatened. Common to locally abundant in many parts of very extensive range. Estimated European population 2,500,000-3,500,000 pairs, vast majority concentrated in Iberia; widespread and numerous in Spain (in early 1990s 2,200,000-2,600,000 pairs, most in Ebro Valley, S Meseta and Andalucia, but absent in N humid belt), with up to 2-5 pairs/ha in best habitats, and common in Portugal (100,000-1,000,000 pairs, mainly in S); much smaller populations in France (1000-5000 pairs), Italy (15,000-30,000), Croatia (1000-1500), Albania (1000-3000), Greece (20,000-50,000), Bulgaria (5000-10,000), Romania (6000-8000), Hungary (8-10), Ukraine (7000-11,000 pairs); on Mediterranean islands, widespread but not abundant in Balearics, scarce Corsica (tens of pairs), common to fairly common Sardinia, Sicily and Crete, very common to common Malta (2000-3000 birds) and Cyprus (perhaps 5000-20,000 pairs); has bred Switzerland (1989) and E Austria (1966), and regularly S Slovakia since 1992. Estimated Russian population 100,000-1,000,000 pairs, with 2-3 pairs/ha on Russian steppes. In C parts of range, 10,000-100,000 pairs in Turkey (widespread and locally common in W, scarce in E), common in Azerbaijan, scarce in Lebanon, fairly common in Israel (a few thousand pairs), local in Jordan (where relatively dense colonization at Azraq in early 1980s, but no recent evidence of breeding); fairly common in China. In N Africa, common in Morocco, widespread in N Algeria, very common and widely distributed in Tunisia, rather scarce in Libya; scarce in NE Egypt, bred in Wadi El Arish area in 1989 and 1990 (breeding formerly suspected only for Wadi El Natrun, in early 19th century). In African non-breeding quarters widespread in Sahel belt, where locally abundant in Mali, widespread in Niger, and common in extreme NE Nigeria, Sudan (abundant on short-grass plains) and NE Ethiopia; common winter visitor in at least Saudi Arabia, Pakistan and parts of India. Declining, and considered "vulnerable", in Europe, where strong decline and range contraction in France (especially in W); over 20% decrease in numbers in 1970-1990 and perhaps over 30% in 1990-2000 in Spain; main threats from agricultural intensification (leading to loss of fallows, increased number of irrigation schemes, increase in surface area covered by crops, etc.) and afforestation of wastelands. Race hungarica threatened with extinction; only 8-10 pairs in Hungary, where numbers in traditional area, the Hortobágy steppe, have declined sharply since early 1980s, although sudden increases in certain years (attributed to influxes from neighbouring populations, hence casting doubts on validity of this race). Reported as causing damage to cultivations of pearl millet (Pannisetum typhoides) on wintering grounds in NW India (Rajasthan).
Bibliography. Adamian & Klem (1999), Ali (1996), Ali & Ripley (1987), Alström et al. (1992), Barlow et al. (1997), Bergmann & Helb (1982), Bindl et al. (1990), Brazil (1991), Brinkman (1987), Brooks, Evans et al. (1987), Bundy (1976), Bundy & Warr (1980), Cave & Macdonald (1955), Cheng Tsohsin (1987), Cheylan et al. (1983), Christen (1983), Clarke (1980), Colston & Sirihai (1986), Cornwallis & Porter (1982), Cramp (1988), Curchod et al. (1990), Dathe (1952), Dementiev et al. (1970), Dennis & Wallace (1975), Díaz (1994b), Dickinson & Dekker (2001a), Dowsett & Dowsett-Lemaire (1993), Dowsett & Forbes-Watson (1993), Dubois et al. (2000), Elgood et al. (1994), Endes (1970), Endes et al. (1967), Étchécopar & Hüe (1964), Flint & Stewart (1992), Fraticelli & Sorace (1988), Glutz von Blotzheim & Bauer (1985), Godlinsky (1996b), Goodman et al. (1989), Goriup (1988a), Grimmett et al. (1998), Guichard (1960), Hagemeijer & Blair (1997), Handrinos & Akriotis (1997), Heath et al. (2000), van Heezik & Seddon (1999), Heim de Balsac (1936), Heim de Balsac & Mayaud (1962), Hollom et al. (1988), Horváth (1956), Hüe & Étchécopar (1970), Inskipp & Inskipp (1991), Isenmann & Moali (2000), Jennings (1981), de Juana & Suárez (2003), de Juana et al. (1988), Karnas (1986), Keith et al. (1992), Kumerloeve (1967, 1969), Lang (1987), Lardelli (1986), Ledant et al. (1981), Mackworth-Praed & Grant (1960, 1970), Magyar et al. (1998), Makatsch (1974), Marchant (1963), Mayaud (1985), Meinertzhagen (1951, 1954a), Meschini & Frugis (1993), Meyer de Schauensee (1984), Mihaly (1980), Moore & Boswell (1956), Muselet (1979, 1981, 1983), Nadler (1974), Niethammer (1963b), Olioso & Cheylan (1999), Olioso et al. (1983), Pesente (1991), Popenko (1979), Ramadan-Jaradi & Ramadan-Jaradi (1999), Richardson (1990), Ripley (1982), Roberts (1992), Roselaar (1995), Rufino (1989), Salvan (1968), Schönwetter (1979), Shirihai (1996), Simms (1992), Smith (1965), Smythies (1986), Snow & Perrins (1998), Suárez et al. (2002), Sultana (1991), Sultana & Gauci (1982), Svensson et al. (1999), Symens & Suhaibani (1993), Tellería (1981), Tellería et al. (1999), Tellini (1987), Thévenot et al. (2003), Thomsen & Jacobsen (1979), Urban & Brown (1971), Vaurie (1951a, 1959), Vinicombe & Cottridge (1996), Vittery et al. (1972), Walker (1981b), Wallace (1983, 1984), Whistler (1941), White (1961a), Zhang Wenguang (1982), Zimmerman et al. (1996), Zink (1987).