Larks

(Alaudidae)

Class Aves

Order Passeriformes

Suborder Passeri (Oscines)

Family Alaudidae

Thumbnail description
Small, highly terrestrial passeriforms with usually cryptic plumage; well known for continuous song and spectacular aerial song-display of males in most species

Size
3.9–9.0 in (10–23 cm); 0.4–2.6 oz (12–73 g)

Number of genera, species
15 genera; 78 species

Habitat
Open landscape, grassland, steppes, arid and semiarid habitats, wastelands, and cultivated fields

Conservation status
Endangered: 2 species; Critical: 2 species; Vulnerable: 4 species; Near Threatened: 3 species

Distribution
Africa, Europe, and Asia; one species through Wallacea to Australia, one species in North America with local population in South America

Evolution and systematics

Larks (Alaudidae) are distinguished from other songbirds by several characters. The back of the tarsus is rounded and covered with individual horny scutes on its rear side; unlike in all other oscines, where the back side of the tarsus is generally covered with one single and sharply edged scute. The lark's syrinx has no pessulus, or small cartilaginous knob at the point of fusion of the bronchiae. The pessulus is present in all other passeriform birds, with the exception of Funarioidea and a few tyrannid species.

Relationships within the Passeri remain uncertain. Larks are predominantly ground-dwelling birds; they evolved, however, from more arboreal ancestors. The recapitulation of arboreal habits can be observed during the development of juvenile larks. When leaving the nest before they can fly, they first hop on the ground like thrushes, but are able to walk within a few days just like their parents. Strong indications for their long adaptation to terrestrial habits are their courtship display and nest building on the ground, egg-rolling behavior, and distraction-display. When alarmed, larks first squat onto the ground, trusting their cryptic plumage to hide them instead of flying away.

The terrestrial lifestyle of the larks is similar to that of pipits and wagtails (Motacillidae), but this is not due to common ancestry. Motacillidae share with Fringillidae, Emberizidae, Parulidae, and Icteridae a sequence of nine additional nucleotides within a specific gene, a sequence missing in larks and in all other birds investigated.

Fifteen genera of larks have been distinguished, and 78 species are listed in Peters Checklist, about a third of which are members of genus Mirafra. However, according to recent reinvestigations, including molecular, morphological, and behavioral studies, eight species should be added which had been thought to be subspecies.

Members of the Alaudidae should not be confused with other birds commonly called larks: meadowlarks (Sturnella) belong to the Icteridae, the magpie-larks (Grallinidae) of Australia and New Guinea are members of the corvid assemblage, and the Australian songlarks (Cinclorhamphus) are thought to be Sylviidae.

Physical characteristics

Larks range in size from the size of a finch to that of a thrush, i.e., 3.9–9.0 in (10–23 cm) and weight 0.4–2.6 oz (12–73 g). In some species males are larger and heavier than females. The plumage is usually inconspicuous, the upper-parts being heavily streaked or unmarked, with a general brownish to grayish color. The plumage is often adapted to the color of local soil so that many larks are remarkably camouflaged. Nonmigrating species with large breeding ranges extending over many habitats with different soil types tend to contain several subspecies that differ in color. The underparts are usually light and without any pattern. Several species have crown feathers which they can raise to a crest; most conspicuous is the crested lark (Galerida cristata) and its congenerics. In most species both sexes look very similar. Even both sexes

of the horned lark (Eremophila alpestris) and Temminck's lark (E. bilopha) possess a contrasting colored pattern of breast and head, and they have tiny elongated feathers above their eyes that form conspicuous horns. However, the breeding plumage of the male black lark (Melanocorypha yeltoniensis) is entirely black. The males of the seven known sparrow-larks (Eremopterix) have black underparts and a species-specific black-and-white pattern on their head. Only the male black-eared sparrow-lark (E. australis) is totally black-headed; females of all these species are colored as larks in general.

Juveniles often have a spotted plumage, which distinguishes them from their parents. This is most conspicuous in the juveniles of horned larks and Temminck's larks, where the feather-horns of the adult plumage are also missing. Adult larks molt completely once per year after the breeding season. With the exception of Alaemon and Eremopterix, the juvenal plumage is replaced by the adult plumage immediately after the bird becomes fully fledged. This juvenal molt is very unusual among nontropical oscines.

In general, the nostrils are concealed with small tufts of feathers. They are clearly visible, however, in the case of Alaemon, Certhilauda, Heteromirafra, Mirafra, and Pinarocorys.

The bills of larks are astonishingly diverse in shape, ranging from short, heavy, and conical to elongated, thin, and pointed. These differences reflect adaptations to a variety of food and feeding techniques. The thick-billed lark (Ramphocoris clotbey) stands out as an extreme; its bill is massive, short, and very deep, reminiscent of the bill of a grosbeak. This lark is, however, unique in having a toothlike projection on its lower mandible which fits into a notch in the upper mandible. The short bills of Calandrella and Eremopterix are less heavy, but similar to those of finches due to their conical shape. All these species feed mainly on seeds, but insects are more common in the diet of other larks. The horned lark has a short, pointed bill like some pipits; the same applies to the wood lark (Lullula arborea) and fawn-colored lark (Mirafra africanoides). Others like the red-winged lark (Mirafra hypermetra) or the crested lark have long and robust bills which are also used for digging. Dupont's lark (Chersophilus duponti), the greater hoopoe lark (Alaemon alaudipes), and the Cape long-billed lark (Certhilauda curvirostris) are characterized by elongated, slightly decurved, and slender bills. Some species are sexually dimorphic in respect to bill length; the male has a longer bill than the female.

Larks usually have short legs, but they are fairly long in Alaemon and Certhilauda. Larks have strong feet. The hind claw is much longer than the toes, especially in species living on soft soil. However, the hind claw is shorter in larks that live on hard ground or are fast runners such as Alaemon.

All lark nestlings have a characteristic brightly colored gape, with one black spot inside the tip of the upper and lower mandible. There are two black spots on the base of the tongue in Ammomanes, some Certhilauda, and Mirafra; and an additional third spot on the tip of the tongue in Alauda, Calandrella, some Certhilauda, Eremophila, Galerida, and Lullula. After hatching, larks are covered with scanty down on their foreheads, napes, backs, shoulders, wings, and thighs.

Distribution

Most lark species occur in Africa, and many have a discontinuous area of distribution and a limited range. Such distribution patterns, in combination with a highly sedentary lifestyle that prevents gene flow between separated populations, are important preconditions for speciation. For example, southwestern Africa is inhabited by six sedentary and closely related Certhilauda species; the short-clawed lark (C. chuana) is probably the sister taxon to a group consisting of five species which were thought to be conspecific until 1999. They are allopatric species; they do not interbreed as their ranges neither overlap, nor are they in contact.

Africa is also inhabited by genera that are primitive among larks in having unconcealed nostrils (Alaemon, Certhilauda, Heteromirafra, Mirafra, and Pinarocorys), so it is probable that

larks are of African origin, and that their first radiation took place on this continent. Madagascar was colonized once by the African ancestor of the Madagascar lark (Mirafra hova). Today, several Mirafra larks occur in Asia. However, it is not possible to say how many species colonized Asia independently, since it is unknown whether the group of Asiatic Mirafra species is monophyletic, i.e., evolved from one common ancestor.

During a second radiation, several lineages derived from an ancestor with concealed nostrils, and perhaps this ancestral species was also of African origin. Some lineages certainly evolved in Africa. One lineage led to Spizocorys, others (probably in Asia) led to Alauda, Eremophila, and Lullula.

The colonization of Australia by the Australasian bushlark (Mirafra javanica) and of North America by the horned lark happened in the Pleistocene at the latest. During Pleistocene glacial periods, sea level was much lower than today. Sumatra, Kalimantan, and Java (Sunda shelf) were therefore part of Asiatic mainland, and New Guinea was connected to north Australia via the Torres land bridge, forming the dispersal route of the Australasian bushlark.

At the same time in the Northern Hemisphere, the Bering land bridge, the continental shelf between Siberia and Alaska, was exposed. The horned lark spread from Asia into North America along this land bridge. Its recent breeding range in North America reaches from northern Alaska to the Gulf of Mexico, and a separated population established itself on the Andean slopes of Bogotá, Colombia.

The skylark (Alauda arvensis) reached Siberia from the west approximately 11,000 years ago, after the Bering Strait was formed due to rising sea-levels. This barrier slowed down its eastward dispersion. However, there are records of possible breeding in the 1970s and two well-documented, but probably unsuccessful, broods in 1995 on the Pribilof Islands, Alaska. The skylark's northward expansion toward Central Europe from southern and eastern Europe is closely connected to increasing deforestation and agriculture during the seventh to thirteenth centuries, which provided appropriate habitats for this species.

The crested lark reached Central Europe from southwestern and eastern Europe later than the skylark and was widespread in the sixteenth century, but receded during the Little Ice Age in the seventeenth and eighteenth centuries. However, it spread into Central Europe again from the middle of the nineteenth century onward, helped by global warming and man-made habitats such as roads and railway stations. Crested larks even reached Scandinavia in 1900, but became extinct in the 1990s, possibly because of climatic change.

Habitat

Larks are characteristic birds of the open landscapes like grasslands, steppes, stony plains, and heaths. Most larks prefer areas with sparse vegetation. Some species, including the wood lark and flapped lark (Mirafra rufocinnamomea), depend on a mixture of vegetation types within the same habitat, such as grasses for nest-building and scattered bushes and small

trees for perching. Human activities provide further suitable habitats for certain species. In North America, skylarks and horned larks regularly breed on cultivated areas such as fields and uncultivated areas such as wastelands.

Larks inhabit extreme regions such as deserts, semideserts, and arctic tundras, and areas varying in altitude from about sea level to high mountains. The horned lark, for example, breeds at 14,750 ft (4,500 m) in the Rocky Mountains, and the skylark and Tibetan lark (Melanocorypha maxima) breed at 14,450 ft (4,400 m) and 15,100 ft (4,600 m) in the Himalayas, respectively. Generally, such extreme habitats are left after the breeding season.

Behavior

Larks are diurnal, ground-dwelling birds, sleeping on the ground in self-made depressions. They scratch their head by indirect method, and frequently take baths in dust or sand like chickens. They may bathe in rain, but not in water. Larks move on the ground by walking and running, reaching high speeds. Their flight is typically strong and undulating, with wings closed periodically.

Larks inhabiting arid climates have evolved certain strategies to cope with these severe conditions. They avoid contact with hot soil by perching on elevated stones and shrubs, and during the hottest part of the day, they become inactive and shelter in the shade provided by vegetation or stones. In the Arabian desert, larks rest in lizard burrows. Parents shade their nestlings by standing over them with wings spread.

Larks are famous for their melodious and continuous songs, which last from minutes to up to almost an hour. Some species even sing at night. Many species enlarge their repertoire

with imitations of other bird songs and calls. Because of this behavior, the Mongolian lark (Melanocorypha mongolica) is called "Hundred Melodies" in China and is a favored cage bird in East Asia. The Latakoo or melodious lark (Mirafra cheniana) is known to imitate 57 different bird species—even ducks, guineafowl, and bee-eaters—and single males can be distinguished by the set of birds they imitate. Some species, such as the crested lark, may even imitate human whistling.

Song is performed during aerial song-displays while males circle about their territories. Some species rise almost vertically from ground or perch and ascend up to 330 ft (100 m) or more before gliding or dropping with closed wings back to the ground. Continual hovering and singing is characteristic for the skylark. Several lark species frequently utter their songs from the ground and elevated perches such as stones, tops of bushes, or trees.

During its song-flight, the male of the black lark claps its wings above its back, reminiscent of the flight-display of pigeons. Some species within the genus Mirafra, as well as the Dupont's lark, can create rattling sounds with their flight feathers, a sound often compared to the song of cicadas. Wing flapping is generally performed during the lark's ascending phase of song-flight. The frequency of flaps is species-specific and individually variable, and regional dialects can be distinguished. The extent of the wing-flapping behavior is also species specific. Other than its melodious song, the white-tailed lark (Mirafra albicauda) utters only soft instrumental sounds. Nonvocal sounds are more prominent in rufous-naped lark (M. africana), and clapper lark (M. apiata), where clattering-flight is still followed by singing. In the flappet lark, however, sound created by the wings replaces the bird's song almost entirely.

As far as one knows, courtship behavior is displayed on the ground. The male hops and steps around the female in upright posture spreading and cocking its tail-feathers. The undertail-coverts are presented to the female (they are entirely black in the black and the black-crowned sparrow-lark). The wings are drooped and also spread to some degree quivering slightly. The crown feathers are raised even in species without elongated crest feathers. During display, the male utters song fragments. Occasionally, the male presents food items to its mate immediately before copulation (courtshipfeeding).

Several desert-inhabiting larks, including sparrow-larks, are nomadic, and their movements depend on rainfall and food supply. Migratory and nomadic species have to some extent a flocking behavior, and some species form sex-specific flocks during winter months. The granivorous larks, members of the genera Eremopterix and Calandrella, are very gregarious outside their breeding period, forming large flocks.

Feeding ecology and diet

Larks feed on arthropods, as well as seeds, green plant material, buds, and fruits. Food items are taken directly from the ground or pecked from plants. Some larks even prey on venomous insects or arthropods that have chemical defense

strategies, such as ants, darkling beetles, stink bugs, and millipedes. Rarely, flying insects are taken in aerial pursuit. The bill can be used for digging and probing. Depending on their diet, more or less insectivorous and granivorous species can be distinguished, but seasonal changes occur; the crested lark, wood lark, and skylark take fewer seeds during breeding season than in winter. All larks feed arthropods to their young, only Stark's lark (Eremalauda starki) feeds a high proportion of unripe grass seeds, even to newly hatched chicks.

In mainly insectivorous larks, the male is larger and has a longer bill than the female. This is most conspicuous in the greater hoopoe, the long-billed lark, and their relatives, which use their slender and decurved bills for digging in the ground in search of insect larvae. Sexual dimorphism in bill and body size also occurs in the bar-tailed lark (Ammomanes cincturus) and Gray's lark (A. grayi), which feed mainly on seeds. Such differences in size enable both sexes of the same species to exploit different food resources within the same habitat.

Most larks swallow whole seeds, which are crushed in their stomach using grit. Indigestible remains are ejected as small pellets. Larks in the genera Calandrella, Eremopterix, and Melanocorypha de-husk seeds in a finchlike manner, fixing the grain between the tongue and palatine and breaking it up. Crested larks, wood larks, and skylarks remove husks from seeds by beating them against the ground. They use the same technique for removing the legs and wings of large insects. Like the song thrush (Turdus philomelos), greater hoopoe larks crack the shells of snails using stones like an anvil. The same behavior was observed once in the crested lark in Morocco, but never in Central Europe. The greater hoopoe lark also frequently drops snails onto stones until their shells break.

Many larks satisfy their thirst and maintain body weight by drinking dew when water is not available. Various species, including the black, desert, Gray's, and Stark's lark, as well as the black-crowned and black-eared sparrow-lark, drink brackish or even salty water.

Reproductive biology

Larks are monogamous, and pairs stay together at least for one breeding season to raise one or two, rarely three, broods. The breeding period runs from March to July in the Palearctic, in the Tropics and South Africa it depends on the rainy season. Even erratic rainfall can induce egg-laying in some nomadic lark species. Most larks nest on the ground. The only exception is the greater hoopoe lark, which commonly builds its nest in tops of shrubs up to 24 in (60 cm) above the ground. Nests are cup-shaped depressions scratched into the ground, sheltered by grass tufts or stones and mainly lined with plant material. Some Mirafra and Certhilauda species frequently cover their nests with a dome made of grass and supported by vegetation behind the nest. If a nest is built on solid ground and it is impossible to scratch a depression, some larks add a rim or rampart of small pebbles, plant material, or pieces of dung or mud. This behavior has been observed for Ammomanes, Calandrella, Eremophila, Eremopterix, and Ramphocoris.

In most species the female builds the nest alone. However, in the calandra lark (Melanocorypha calandra) and Oriental skylark (Alauda gulgula), as well as several species within the Ammomanes, Calandrella, and Galerida, both sexes are involved in nest building. The male of the chestnut-backed sparrow-lark (Eremopterix leucotis) presents spider webs to its mate, which applies them to its nest. Ritualized nest building behavior as part of the courtship display is reported from two species. The male of the Rudd's lark (Heteromirafra ruddi) walks in front of the female with its bill full of nest material, and the male of the thick-billed lark offers the female small pebbles used for nest building.

The eggs are creamy-white or yellowish and are more or less spotted. Egg-laying occurs daily in early morning, and only one egg per day is laid. In favorable habitats the clutch size ranges from two to five, rarely up to seven, eggs. Inhabitants of arid or semiarid areas generally have smaller clutches. Gray-backed sparrow-larks (Eremopterix verticalis) in the Karoo, South Africa, lay significantly larger clutches in years with heavy rainfall. Females of the closely related black-crowned sparrow-lark (Eremopterix nigriceps) lay no more than three eggs, but only two young per brood are reared because each adult feeds only one single chick after they have left the nest. This "split brood-care" behavior, as well as the reduction of clutch size, might be an adaptation to the severe desert conditions, which make egg production and providing food to the young difficult.

Incubation starts with the last egg and, depending on the species, takes 11–16 days. Usually, only the female incubates and broods the young. However, sparrow-larks and a few other species seem to be an exception to this rule.

In all larks, both sexes feed the chicks and eat or remove their feces. After eight to 13 days, before they are able to fly, lark chicks leave the nest, still supplied with food by their parents. If the female starts a second brood, the male cares for the young alone. Steyn observed cooperative breeding in the spike-heeled lark (Chersomanes albofasciata); three adults fed two chicks in one nest in the Karoo.

Conservation status

According to the IUCN, several lark species are in need of special protection. Ash's lark (Mirafra ashi) is known only from six specimens collected in 1981 in a very small range in Somalia. Because its minute population seems to decline, this species is classified as Endangered. Botha's lark (Spizocorys fringillaris) is also Endangered. It is patchily distributed within South Africa, and its population is declining due to habitat destruction by agriculture.

The situation is critical for the Raso lark (Alauda razae) and Rudd's lark. The former species is endemic to the uninhabited Islet of Raso (Cape Verde Islands), and its population is very small (92 birds counted in 1998). Accidentally introduced predators are therefore a serious threat to eggs and nestlings. A rapid population decline of Rudd's lark could be prevented by protecting its breeding range in South Africa.

Four species are classified as Vulnerable because of their restricted and unprotected ranges and small populations. The ferruginous or red lark (Certhilauda burra) breeds in the Cape Province only, where habitat fragmentation continues. Archer's lark (Heteromirafra archeri) is known only from two small areas in Somalia, where it was recorded last in 1955. Only a few specimens and observations of Degodi lark (Mirafra degodiensis) and Sidamo bushlark (Heteromirafra sidamoensis) are known from different localities in Ethiopia. Information about breeding range and population size is even more scanty for Friedmann's lark (Mirafra pulpa) from northeastern Africa, Obbia lark (Spizocorys obbiensis) from Somalia, and Williams's lark (Mirafra williamsi) from Kenya, making it almost impossible to assess the extent of the threat to their extinction.

The population of the Agulhas long-billed lark (Certhilauda brevirostris) is estimated to be around 1,000 mature individuals. Estimates for Sclater's lark (Spizocorys sclateri) are much higher, but the population is severely fragmented throughout Namibia and South Africa. Finally, the population of the Latakoo lark in southeastern Africa could decline drastically during the following decades due to habitat loss. As a result, these three species are considered to be Near Threatened.

In the countries of the European Union (EU), the calandra lark, Dupont's lark, short-toed lark (Calandrella brachydactyla), Thekla lark (Galerida theklae), and wood lark are listed in Annex I of the EU Birds Directive as species which require special conservation efforts. All these species suffer habitat destruction related to agricultural intensification and afforestation, as well as illegal hunting and predation. The European populations of crested larks and skylarks declined during the last years, probably due to climatic change, the loss of suitable habitats, and the increasing use of pesticides. The crested lark became extinct in northern Russia, Norway, Sweden, and Switzerland in the 1990s; declined in Denmark, the Netherlands (3,000–4,000 breeding pairs in 1979 vs. 400 in 1991), France, Germany, and Poland. Skylark numbers in Great Britain declined by 51% between 1968 and 1995, a loss of approximately three million breeding birds. In 1997, only about one million pairs were breeding in Great Britain.

Significance to humans

In Central Europe, larks are regarded as harbingers of spring, and they were honored by the ancient Greeks as mediators between heaven and earth, and by the Celts as being beneficial for their harvest. Their melodious and long song, especially that of the skylark, is celebrated in verse by poets, and Shakespeare called the lark "the herald of the morn" in Romeo and Juliet. Skylarks were successfully naturalized in Australia in the 1850s, in New Zealand and on the Hawaiian Islands in the mid 1860s, as well as on Vancouver Island, British Columbia, in 1903 (although declining seriously in the 1990s). Skylarks were also released at other places in North America, but they were not able to build up stable populations and thus became extinct.

Even today, larks are trapped and shot legally in France and in the Mediterranean region. Spaepen calculated that five times as many skylarks are killed by humans in France than anywhere else in Europe, and the number of skylarks killed annually in southwest France is estimated at five to 10 million birds!

Species accounts

List of Species

Australasian lark

Mirafra javanica

taxonomy

Mirafra javanica Horsfield, 1821, Java.

other common names

English: Singing bushlark; French: Alouette de Java; German: Horsfieldlerche; Spanish: Alondra de Australiana.

physical characteristics

4.7–5.9 in (12–15 cm); 0.7 oz (20 g). Plumage inconspicuous other than rufous wing patch, sexes alike.

distribution

Myanmar, southern and central Thailand, Indochina, Philippine Islands, Kalimantan, Java, Bali, Lesser Sunda Islands, local in New Guinea, northwestern, northern, eastern, and southeastern Australia.

habitat

Open grassland with scattered bushes, salty marshes, and edges of cultivated fields.

behavior

Perches on trees and wires. Song includes imitations of other birds; song-flight takes up to 40 minutes. Forms small groups outside breeding period. Migratory in southern Australia.

feeding ecology and diet

Feeds on seeds and arthropods.

reproductive biology

Monogamous. Builds domed nests in shelter of tussocks; in southern Australia, lays two to four eggs November through January.

conservation status

Not threatened.

significance to humans

None known.

Cape long-billed lark

Certhilauda curvirostris

taxonomy

Alauda curvirostris Hermann, 1783, Cape of Good Hope.

other common names

English: Long-billed lark; French: Alouette à long bec; German: Langschnabellerche; Spanish: Alondra Picuda.

physical characteristics

Male 7.5–7.9 in (19–20 cm); female 6.3–6.7 in (16–17 cm); weight unknown. Males clearly larger than females, but sexes do not differ in respect to plumage color or pattern. Plumage brown-grayish, heavily streaked, breast marked with dark spots. Species reminiscent of a thrush with a long, decurved bill. Hind claw is long and straight.

distribution

Small strip from southern Namibia to southwestern South Africa, Cape Province.

habitat

Coastal dunes and arid fields, avoids more rocky areas.

behavior

Solitary or in pairs. Male performs spectacular aerial song-display, characteristic for all Certhilauda species. Male first flies low over the ground, then rises nearly vertically several feet into the air and closes its wings before reaching the high point. After that it nose-dives and descends vertically, not opening the wings before coming close to the ground. Whistles while descending; same song is also uttered from prominent perches.

feeding ecology and diet

Feeds on termites, grasshoppers, beetles, and ants, as well as on seeds and berries. Uses long bill for digging. Since sexes differ in size of bill, they probably exploit different food resources.

reproductive biology

Monogamous. Breeds September through December; nest often domed, clutch size ranges from two to three eggs; both parents feed the young.

conservation status

Not threatened. Common in its limited southwestern African range. Range may have increased due to agriculture.

significance to humans

None known.

Black-crowned sparrow-lark

Eremopterix nigriceps

taxonomy

Pyrrhalauda nigriceps Gould, 1841, São Tiago, Cape Verde Islands.

other common names

English: Black-crowned finch-lark, pallid finch-lark; French: Moinelette à frout-blanc; German: Weißstirnlerche; Spanish: Aloudra gottión de Corouna Negra.

physical characteristics

4.5–4.9 in (11.5–12.5 cm); male 0.5–0.6 oz (14–16 g); female 0.4 oz (12 g); one of the smallest lark species. Sparrow-larks, like finches or sparrows, have a proportionally large head and a strong conical bill. Sexes are dimorphic with respect to plumage color. Female mainly pale brown and streaked, male with white forehead, cheeks, sides of neck, and collar of nape. Crown, stripe through eye to base of bill, and lower border of cheek black. Underparts entirely black, upperparts grayish brown.

distribution

Cape Verde Islands, southern Morocco, northern Mauritania to western Sahara, southeast Egypt. From North Africa, through most of the Arabian Peninsula, including Island of Sokrota, to Pakistan and northwestern India. Expanded its area to northern Nigeria, possibly due to desertification. Vagrant to Israel, Jordan, and Algeria.

habitat

Semidesert and sandy areas with sparse vegetation, also close to cultivated land.

behavior

Territorial during breeding season, gregarious other times. In song-flight, male ascends at a 45° angle with legs dangling up to 66 ft (20 m), then flies in roughly circular paths in an undulating style to descend in stages. Song also performed from elevated perch.

feeding ecology and diet

Feeds more on seeds than arthropods; de-husks seeds; can drink salty or brackish water.

reproductive biology

Monogamous. Female builds cup-shaped nest, male may also be involved in nest-building. Rim of nest frequently surrounded by small lumps of soil or stone. Breeding period irregular; two to three eggs laid, incubated by both sexes for 11–12 days. Both parents feed and brood. Young leave nest after eight days before being able to fly. Each parent cares for one single chick, so that only two young per brood can be reared (split brood-care).

conservation status

Not threatened.

significance to humans

None known.

Greater hoopoe-lark

Alaemon alaudipes

taxonomy

Upupa alaudipes Desfontaines, 1789, between Gafsa and Tozzer, Tunisia.

other common names

English: Hoopoe lark, bifasciated lark; French: Sirli du désert; German: Wüstenläuferlerche; Spanish: Alondra Ibis.

physical characteristics

7.1–7.9 in (18–20 cm); male 1.4–1.8 oz (39–51 g); female 1.1–1.6 oz (30–47 g); one of the largest lark species, females

smaller than males. Bill very long, slender, and slightly decurved. Plumage sand-colored above, underside white, throat and breast with dark spots, similar in both sexes. Wings long and broad with conspicuous black-and-white pattern, comparable to that of African subspecies of hoopoe (Upupa epops). This lark was first described as a hoopoe, and its original scientific name means "hoopoe with legs of a lark."

distribution

Cape Verde Islands, North Africa from Mauritania to Egypt and Sudan, across the Middle East to northwestern India.

habitat

Semideserts and deserts.

behavior

Tame and confiding, allowing humans to approach to within a few feet. Male defends territory against conspecific intruders and other birds by wing-spread behavior. Usually solitary or in pairs. Song ringing and far-carrying. For song-flight, male jumps up from perch and starts to sing, ascending almost vertically up to 33 ft (10 m) and performs somersaults, displaying its contrasting pattern on tail and wings. It then closes its wings, nose-dives back to perch, not opening its wings until landing. Song-flights can be repeated frequently within up to one hour.

feeding ecology and diet

Mainly arthropods and snails. Most food items obtained by digging with bill. Female bill is 30% shorter than bill of male, so sexes probably exploit different food resources. Two methods for cracking snail shells: snails dropped onto stone in flight or smashed against stones directly until shells break.

reproductive biology

Monogamous. Only lark species building nests frequently on top of low shrubs, up to 24 in (60 cm) above ground. Also building cup-shaped nests on ground. Two to four eggs incubated by female for about 14 days. Both parents feed young, which leave nest after 12–13 days, before being able to fly. Young remain for at least one month with parents.

conservation status

Not threatened.

significance to humans

None known.

Thick-billed lark

Ramphocoris clotbey

taxonomy

Melanocorypha clot-bey Bonaparte, 1850, Egyptian desert.

other common names

English: Clotbey lark; French: Alouette de Clotbey; German: Knackerlerche; Spanish: Alondra de Pico Grueso.

physical characteristics

6.7 in (17 cm); male 1.8–1.9 oz (52–55 g); female 1.6 oz (45 g). Strong lark with large head. Short but massive bill most conspicuous. Blunt, toothlike projection on middle of lower mandible fits into notch on upper mandible. Upperparts uniformly pink/gray-brown, chest to vent with large black spots, sides of head blackish, throat and eye-ring white; plumage of females of less contrasting color and not so heavily streaked.

distribution

Patchily distributed in North Africa, north of Sahara from northern Mauritania and Morocco to Algeria, Tunisia, and Libya. Further east, distribution uncertain, but reported as breeding bird from Egypt, Syria, Jordan, and Arabia.

habitat

Stony deserts with sparse vegetation, avoids sand dunes.

behavior

Solitary or in small groups during breeding season, but large flocks observed in winter. Male rises from ground and starts to sing before descending in parachute style.

feeding ecology and diet

Feeds on seeds and green plant material as well as on insects. Uses bill to cut off plant material and to dig for food, and maybe for cracking solid cuticles of large beetles. However, even hard seeds are swallowed whole and not husked with bill, instead grit is taken with food to aid digestion.

reproductive biology

Monogamous. During ground display, male presents to female pebbles that are used in nest building. Nest cup-shaped, frequently surrounded by small lumps of soil or small stones. Two to three eggs incubated by female March through May, both parents feed young.

conservation status

Not threatened.

significance to humans

None known.

Calandra lark

Melanocorypha calandra

taxonomy

Alauda calandra Linnaeus, 1766, Pyrenees.

other common names

French: Alouette calandre; German: Kalanderlerche; Spanish: Calandria Común.

physical characteristics

7.1–7.5 in (18–19 cm); male 2.0–2.6 oz (57–73 g); female 1.8–2.2 oz (50–65 g); larger and stronger than skylark. Bill conical and heavy. Upperparts brown and streaked, black patches on each side of upper breast characteristic. Sexes alike.

distribution

North Africa, southern Europe east to Ural steppes, from Asia Minor to central Asia, missing between Caspian Sea and Lake Aral.

habitat

Open lowlands, steppe, grasslands, cultivated farmland, and meadows.

behavior

Resident in southern Europe, Near East, and North Africa, migratory in Russia. Forms flocks of up to 2,500 individuals autumn and winter; frequently associated with other larks and corn bunting (Miliaria calandra). Male utters continuous song from ground or perch. Song-flight performed in circles, ascending in spirals. Several males often sing close together. Song similar to skylark, contains imitations of other birds.

feeding ecology and diet

Diet changes from insects during summer to seeds in winter. Bill used for digging.

reproductive biology

Monogamous. Two broods April through June; both sexes build cup-shaped nest, clutch size ranges from three to six eggs, incubated by female, but brood patch also observed in several males. Young hatch after 16 days; fed by both parents. Leave nest after 10 days before being able to fly.

conservation status

Not threatened, though population is declining in southern Europe due to agricultural intensification and possibly hunting; listed in Annex I of the European Birds Directive.

significance to humans

Hunted in the Mediterranean region.

Greater short-toed lark

Calandrella brachydactyla

taxonomy

Alauda brachydactila Leisler, 1814, France and Italy = Montpellier, France.

other common names

English: Short-toed lark; French: Alouette calandrelle; German: Kurzzehenlerche; Spanish: Terrera Grande.

physical characteristics

5.1–5.5 in (13–14 cm); male 0.7–1.0 oz (21–28 g); female 0.6–0.9 oz (17–26 g). Small lark with dull, cryptic plumage, no streaks on chest. Bill short and finchlike. Sexes alike.

distribution

North Africa, southern Europe, eastward from Asia Minor through Mongolia to China.

habitat

Steppe with sparse vegetation, cultivated land, seashores, and saline areas. Avoids desert, moist areas, and vicinity of forests.

behavior

Populations of Europe and Middle East migratory, wintering south to Sahel and Red Sea. Birds from central Asia winter in India. Highly gregarious outside breeding season. Song-flight performed by male in wide circles in sequence of deep and shallow undulations. Male ascends stepwise, uttering first phrase of melodious song with imitations of other birds, then stops singing and beating wings, drops down, ascends, and drops down again before next ascent, while singing starts again. Song-flight ends with descent or glide-in stages. Song sometimes performed from ground or perch.

feeding ecology and diet

Diet changes from insects and seeds in summer to nearly exclusively seeds during winter. Can go for months without drinking, even drinks brackish water.

reproductive biology

Monogamous. Breeds April through June; cup-shaped nest often surrounded by pieces of mud, dung, and small stones. Female lays four to five eggs, incubates and broods alone, but both parents feed young.

conservation status

Not threatened, though decreasing in France; listed in Annex I of the European Birds Directive.

significance to humans

None known.

Crested lark

Galerida cristata

taxonomy

Alauda cristata Linnaeus, 1758, "in Europae viis" = Vienna.

other common names

French: Cochevis huppé; German: Haubenlerche; Spanish: Cogujada Común.

physical characteristics

6.7 in (17 cm); male 1.3–1.8 oz (37–52 g); female 1.3–1.7 oz (37–48 g); not as large as skylark, more robust, with stronger bill and longer crest. Uniformly dull-colored plumage, upper-parts and breast heavily streaked, sexes alike. Very similar to Thekla lark, its sibling species.

distribution

From southern Europe as far as southern Scandinavia. North Africa south to Senegambia, eastward through Chad and Sudan to Ethiopia and Somalia. Does not occur in the Sahara. Breeds from Arabian Peninsula toward India, Nepal, Mongolia, China,

and Korea. In southern France and North Africa east of Libya, sympatric with Thekla lark.

habitat

Variety of habitats, mainly open areas with sparse vegetation, also cultivated land and other man-made semideserts such as railways, airfields, and wastelands. Where it co-occurs with the Thekla lark, the crested lark occupies the plains, the Thekla inhabits rocky and bushy slopes.

behavior

Largely sedentary, not gregarious, low flight distance. Song-flight starts from perch, male ascends silently at angle up to 230 ft (70 m), then utters loud and melodious song ascending further up to 330 ft (100 m) and more, flying wide circles over territory. Song, uttered on ground or from perch, lasts four minutes on average, but up to 30 minutes have been observed. Known to perfectly imitate other bird songs and calls. One extraordinary example, the imitation of a shepherd's whistle reproduced so accurately that sheep dogs obey the signals as if the shepherd has given them.

feeding ecology and diet

Diet consists mainly of vegetal food; seeds and green plant material taken from ground or picked directly from plants; food remains in horse-droppings also exploited. Can husk seeds. Animal food taken to larger extent during breeding season, but proportion is negligible during rest of the year. Young fed with insects and worms.

reproductive biology

Monogamous. Breeds April–June in Europe; female builds cup-shaped nest alone, lays three to five, rarely seven, eggs. Incubation by female alone, 11–17 days. Both parents feed young, which leave nest after eight to 11 days before being able to fly.

conservation status

Not threatened.

significance to humans

None known.

Wood lark

Lullula arborea

taxonomy

Alauda arborea Linnaeus, 1758, Europe-Sweden.

other common names

French: Alouette lulu; German: Heidelerche; Spanish: Totovía.

physical characteristics

5.9 in (15 cm); male 0.7–1.2 oz (21–35 g); female 1.1–1.2 oz (30–35 g); smaller and more slender than skylark and crested lark. Plumage buff brown, upperparts and chest streaked, distinguished from other larks by broad, white supercilium which continues to nape. Black-and-white pattern of alula (first digit of wing) feathers very conspicuous. Crown feathers can be raised to a small crest.

distribution

Northern West Africa, Europe from the Mediterranean to southern Scandinavia, Asia Minor east to Iran and Turkmeniya.

habitat

Requires habitat with short grass for feeding, higher vegetation for nesting, exposed trees or bushes as song-perches.

behavior

Not gregarious, even during migration. Northern populations migratory, wintering in southern Europe and North Africa; southern populations mostly resident. Arboreal, walks on branches and perches on treetops, bushes, and wires. For song-flight, males take off from perch in tree top, ascend at an angle, spiral upward and fly circles on nearly same level 165–330 ft (50–100 m) above ground, singing all the time (unpaired males fly higher than paired ones). Descent either in stages while still singing or sometimes silently with wings closed. Song-flight takes two minutes on average, but unpaired males have been watched singing for 70 and 94 minutes. Males also sing from perch or ground, and frequently on moonlit nights. Song composed of pleasing and soft phrases, hesitant at the beginning, then increasingly forceful and louder. High site-fidelity of males has been shown by ringing experiments.

feeding ecology and diet

Feeds on small insects and spiders during breeding season, otherwise mainly granivorous.

reproductive biology

Monogamous. Breeds March through June; cup-shaped nest built by female. Two, sometimes three broods annually; clutch size normally three to five eggs, female incubates for 11–15 days. Young fed by both parents, leave nest after eight days before being able to fly. If female starts incubation of second brood, male cares for young of first clutch alone.

conservation status

Not threatened, though declining populations in Europe caused by habitat loss; listed in Annex I of the European Birds Directive.

significance to humans

None known.

Sky lark

Alauda arvensis

taxonomy

Alauda arvensis Linnaeus, 1758, Europe.

other common names

French: Alouette des champs; German: Feldlerche; Spanish: Alondra Común.

physical characteristics

7.1–7.5 in (18–19 cm); male 0.9–1.9 oz (27–55 g); female 0.6–1.7 oz (17–47 g). Extensively streaked brown plumage, crown feathers can be raised to a short crest; bill stronger than bill of wood lark; sexes alike. Most common lark species within its western Palearctic part of distribution.

distribution

North Africa, Europe, and Asia; introduced in Australia, Canada (Vancouver Island, British Columbia), Hawaii, and New Zealand.

habitat

Dense grasslands, cultivated farmland, airfields and sports grounds.

behavior

Perches only rarely on wires and trees. High-level song-flight in breeding season. Male rises with rapid wing-beats, after prolonged hovering falls parachute style and drops to ground suddenly, singing all the time loudly and forcefully. Populations of the western Palearctic are migratory, hibernate in the Mediterranean region; breeding birds of the British Isles are sedentary. Eastern Asiatic populations migrate to southeastern China; populations of central Asia hibernate in northern India, Afghanistan, and Iran; and eastern Palearctic populations migrate to Turkey, Syria, and Jordan.

feeding ecology and diet

Mainly arthropods during summer, but seeds in winter.

reproductive biology

Monogamous. Breeds April–July; two, rarely three, broods. Cup-shaped nest in depression on ground. Three to five, exceptionally up to seven, eggs. Female incubates for 12–14 days, chicks leave nest when eight to 10 days old. Both parents care for young.

conservation status

Decreasing populations in Europe, but not considered threatened.

significance to humans

Still hunted legally in France, Greece, and Italy.

Horned lark

Eremophila alpestris

taxonomy

Alauda alpestris Linnaeus, 1758, North America = coast of North Carolina.

other common names

English: Shore lark; French: Alouette hausse-col; German: Ohrenlerche; Spanish: Alauda Cornuda.

physical characteristics

5.9–6.7 in (15–17 cm); male 1.1–1.7 oz (30–48 g); female 0.9–1.5 oz (26–42 g). Named after elongated black feathers at sides of crown reminiscent of horns; head and breast contrastingly colored, somewhat brighter in males than in females.

distribution

Holarctic; widespread in North America, northern and southeastern Europe, northern and central Asia. Separated populations in Morocco, Lebanon, northern Israel, and South America (Colombia).

habitat

Steppes, semideserts, arctic tundra; prefers open, bare, and stony areas.

behavior

Perches on stones, fence posts, or shrubs. Song uttered while perching and during song-flight up to 300 ft (91 m) above ground. Populations of arctic tundra are migratory, hibernate between 55° and 35° N. Present year round in most areas in North America, from southern Canada southward; some may be permanent residents. Migratory in far north, one of the earliest spring migrants. Horned larks breeding in Alaska and Canada winter south as far as the Gulf of Mexico; populations from the tundras of Northern Europe migrate annually to wintering areas around the North Sea.

feeding ecology and diet

Feeds mainly on insects during breeding season, but takes more seeds during winter months.

reproductive biology

Monogamous. Nest cup-shaped, frequently surrounded with pebbles. Breeds March through July; one to two broods; three to five eggs, incubation takes 13 days; chicks leave nest after eight to 10 days. Both parents care for young.

conservation status

Not threatened.

significance to humans

None known.

Resources

Books

Burton, John F. Birds and Climate Change. London: Christopher Helm, 1995.

Cramp, Stanley, ed. Handbook of the Birds of Europe, The Middle East and North Africa: The Birds of the Western Palearctic. Vol.5. Oxford and New York: Oxford University Press, 1988.

Glutz von Blotzheim, Urs N. (Hrsg.). Handbuch der Vögel Mitteleuropas. Bd. 10 Passeriformes (1. Teil). Wiesbaden: Aula-Verlag, 1985.

Keith, S., E. K. Urban, and C. H. Fry, eds. The Birds of Africa. Vol. 4. London: Academic Press, 1992.

Pätzold, Rudolf. Die Lerchen der Welt. Magdeburg: Westarp-Wissenschaften, 1994.

Sibley, Charles G., and Burt L. Monroe. Distribution and Taxonomy of Birds of the World. New Haven: Yale University Press, 1990.

Periodicals

Alström, Per. "Taxonomy of the Mirafra assamica Complex." Forktail 13 (1998): 97–107.

Baicich, Paul J., Steven C. Heinl, and Mike Toochin. "First Documented Breeding of the Eurasian Skylark in Alaska." Western Birds 27 (1996): 86–88.

Browne, Stephen, Juliet Vickery, and Dan Chamberlain. "Densities and Population Estimates of Breeding Skylarks Alauda arvensis in Britain in 1997." Bird Study 47 (2000): 52–65.

Ericson, Per G., Ulf S. Johansson, and Thomas J. Parsons. "Major Divisions in Oscines Revealed by Insertions in the Nuclear Gene c-myc: A Novel Gene in Avian Phylogenetics." The Auk 117 (2000): 1069–1078.

Lloyd, Penn. "Rainfall as a Breeding Stimulus and Clutch Size Determinant in South African Arid-Zone Birds." Ibis 141(1999): 637–643.

Ratcliffe, Norman, Luis R. Monteiro, and Cornelis J. Hazevoet. "Status of Raso Lark Alauda razae with Notes on Threats and Foraging Behaviour." Bird Conservation International 9 (1999): 43–46.

Riley, Steve. "Crested Lark Using 'Anvil'." British Birds 82 (1989): 30–31.

Ryan, Peter G., Ian Hood, Paulette Bloomer, Joris Komen, and Timothy M. Crowe. "Barlow's Lark: A New Species in the Karoo Lark Certhilauda albescens Complex of Southwest Africa." Ibis 140 (1998): 605–619.

Ryan, Peter G., and Paulette Bloomer. "The Long-Billed Lark Complex: A Species Mosaic in Southwestern Africa." Auk 116 (1999): 194–208.

Spaepen, J. F. "A Study of the Migration of the Skylark Alauda arvensis, Based on European Ringing Data." Gerfault 85(1995): 63–89.

Williams, Joseph B. "Lizard Burrows Provide Thermal Refugia for Larks in the Arabian Desert." Condor 101 (1999): 714–717.

Wilson, J. D., J. Evans, S. J. Browne, and J. R. King. "Territory Distribution and Breeding Success of Skylarks Alauda arvensis on Organic and Intensive Farmland in Southern England." Journal of Applied Ecology 34 (1997): 1–20.

Other

The European Commission. "Discover the Species of the Birds Directive." 1 January 2002. 13 March 2002 <http://europa.eu.int/comm/environment/nature/directive/birdshome_en.htm>

The IUCN Species Survival Commission. "2000 Red List of Threatened Species." 1 January 2002. 13 March 2002 <http://www.redlist.org/>

Albrecht Manegold

Walter Sudhaus, PhD