Larks belong to the order Passeriformes, sub-order Passeri and family . Currently there are 17 recognized lark genera and 91 species.
Larks are small to medium-sized birds (11 to 19 cm in length) that reside in open countryside from desert to alpine tundra. They vary in color from light tan to reddish and tend to blend in well with the soil and vegetation in their chosen habitat.
Larks (particularly sky larks (Alauda arvensis)) have provided inspiration for many poets with their complex and beautiful songs. They are primarily Old World inhabitants. Horned larks (Eremophila alpestris) are the only native lark species in North America. (Dean, et al., 1992; Payne, 2003; ; Simms, 1992; Trost, 2001)
Larks primarily live in the Old World. Fifty-seven percent of lark species are found in Africa, 19 percent in Africa and Eurasia, 16 percent in Asia, 6 percent in Eurasia and 1 percent in the New World. Horned larks (Eremophila alpestris) are the only lark species native to North America. Skylarks (Alauda arvensis) were introduced to Vancouver Island in British Columbia, Canada and can still be found there, and occasionally in Washington state. Skylarks were also introduced to Australia and New Zealand. (Campbell and Lack, 1985; Campbell, et al., 1997; Dean, et al., 1992; Simms, 1992)
- Biogeographic Regions
Larks inhabit open countryside in both temperate and tropical regions. Their habitat includes: shrubland, savana, desert, tundra, grassland and farmland. Larks can be found in habitats from coastal areas at sea level to mountainous areas at an elevation of 4000 m. (Campbell and Lack, 1985; Campbell, et al., 1997; Dean, et al., 1992; Mead, 1985; Simms, 1992)
- Other Habitat Features
Larks are small- to medium-sized birds (11 to 19 cm long, 15 to 75 g) with fairly long legs, wings and tail. Most have long straight claws on their hind toes. The length of the claw depends on the bird’s habitat; longer claws are found on birds that live in areas with soft ground and some vegetation, shorter claws and toes are found on species that live in areas with harder ground. Their brown plumage (ranging from light tan to reddish) is often cryptic and matches the soil color. Some species have crests or tufts of feathers on their head. Sexes resemble each other, but males are usually larger and may have brighter, more distinct color and marking than females. Bill shape and length varies between species and can be a good indication of feeding ecology. Razo larks (Alauda razae) show sexual dimorphism in bill length. The male’s bill is 20 percent longer than the female’s, which suggests that males and females exploit different food sources. Larks molt once or twice per year depending on the species. Juveniles have less distinct coloring and patterns than adult birds. (Campbell and Lack, 1985; Dean, et al., 1992; Mead, 1985; Simms, 1992)
- Sexual Dimorphism
- male larger
- sexes colored or patterned differently
- male more colorful
Although there have been some observations of polygyny, larks are largely monogamous. Females do the majority of the nest building, incubation and brooding, and both adults take part in feeding the young. Males perform display flights (high undulating flight accompanied by singing), and will also display with crests, ruffle their plumage, and bow or hop up and down on the ground. Courtship feeding occurs in some species. Males sing from prominent perches; some female larks may also sing during pair formation. Larks are territorial and defend the nest site using song and flight displays.
Cooperative breeding has been observed in one species. The observed group consisted of the breeding pair and a single helper. (Campbell and Lack, 1985; Campbell, et al., 1997; Dean, et al., 1992; Mead, 1985; Simms, 1992)
Larks are seasonal breeders, usually breeding during the season of highest insect and seed abundance. Larks living in areas with long breeding seasons can have two or three clutches, while those in areas with a short window of time for breeding will have only one. Larks are solitary breeders and will defend nesting territories.
Most larks are ground nesters and build open-cup nests in small, excavated hollows in the ground. Some species build domed nests and a few build nests in shrubs to allow for increased air circulation and cooling. Nests built on the ground are situated next to small clumps of vegetation, rocks, or mounds of earth for protection and shade from the sun and prevailing wind. Nests are made of grass, plant fibers, forbs, bark, dead leaves and sedges, and are sometimes lined with plant down or feathers. Eggs are smooth, white or light blue with gray or olive-brown spots and range in size from 19 to 23 by 13 to 17 mm. Clutch size is usually 3 to 5, but can be as low as one and as high as eight. The egg-laying interval is every other day. Females usually do all of the incubation and brooding, although males in some species will help. Incubation lasts 10 to 16 days; chicks hatch synchronously and are brooded for about 4 days depending on the weather. Young larks are altricial and are fed by both adults. Chicks are fed insects (and sometime seeds) and leave the nest after about 10 days. Chicks usually fledge before they can fly and continue to be fed by their parents for 18 to 20 days.
As is common among ground nesting species, most nest failure is due to depredation. Nest success is usually 30 to 60 percent, but can be as low as 10 percent. (Ali and Ripley, 1972; Campbell and Lack, 1985; Campbell, et al., 1997; Dean, et al., 1992; Mead, 1985; Simms, 1992)
- Key Reproductive Features
- seasonal breeding
- gonochoric/gonochoristic/dioecious (sexes separate)
Female larks do most of the incubating and brooding of chicks (males help in some species). Incubation lasts 10 to 16 days and the altricial chicks are brooded for about 4 days after hatching. Chicks are fed insects and occasionally seeds by both parents. Adults remove fecal sacks from the nests. Nestlings usually fledge before they can fly and continue to receive parental care for 18 to 20 days.
If a predator approaches an active nest, the adults will give alarm calls and often feign injury to draw the predator away. Because many larks nest in open desert areas, chicks are often exposed to sun and heat. Adult birds will stand next to the nest to shade it during the hottest parts of the day. (Campbell and Lack, 1985; Campbell, et al., 1997; Dean, et al., 1992; Mead, 1985; Simms, 1992)
Like most small birds, larks probably live on average only two to five years. The longest living known individuals are an 8 year, 5 month old skylark (Alauda arvensis) and a 7 year 11 month old horned lark (Eremophila alpestris). Average annual mortality for skylarks is 33 percent. (Campbell and Lack, 1985; Gill, 1995; ; Trost, 2001)
Most larks are solitary, although some will flock in agricultural fields to feed and others flock after the breeding season prior to migrating. Some larks are sedentary, others are nomadic, moving in response to rainfall and food availability and others (particularly Northern Temperate species) are long-distance migrants. Larks are usually found in low densities, but are found closer together in high quality habitat.
Larks run rather than hop along the ground and rarely perch on shrubs and trees, except occasionally to sing. Many clean by dusting, although some bath in water. Depending on the species, they molt once or twice a year. Larks that live in desert areas will stand in the shade of grass tussocks during the hottest parts of the day.
Larks are territorial and defend their territories through song and flight displays. Some species will also raise their feather crests during a display. (Campbell and Lack, 1985; Campbell, et al., 1997; Dean, et al., 1992; Gill, 1995; Mead, 1985; ; Simms, 1992)
Communication and Perception
Lark flight, feeding, threat and display calls are quite simple, however, their territorial song is very elaborate. In addition to communicating through song, larks will raise the crest of feathers in their head during agonistic and courtship displays.
Larks are omnivorous and forage on the ground. They eat many species of insects in addition to seeds, grasses, leaves, buds, fruits and flowers (especially during the winter when insects are less available). Some species will also eat snails (Gastropoda), which they break open on rocks. Larks' insect prey are diverse and include: Ephemeroptera (mayflies), Diptera (true flies), Hemiptera (true bugs), Coleoptera (beetles), Collembola (springtails), Orthoptera (grasshoppers), Lepidoptera (adult and larval moths) and Isoptera (termites). Skylarks (Alauda arvensis) in England feed on at least 47 species of insect. Desert species acquire water from their food and dew.
The shape of a Lark’s bill is adapted to its diet and feeding technique. For example, hoopoe larks (Alaemon) have long decurved bills that are used for digging for insect larvae, while calandra larks (Melanocorypha) have strong, stout bills that are used for eating seeds. Some can also locate buried insects by ear. (Campbell and Lack, 1985; Dean, et al., 1992; Mead, 1985; Simms, 1992)
Adult larks have numerous avian predators: falcons (Falconiformes), owls (Strigiformes) and shrikes (Laniidae). Adults, chicks and eggs are also taken by mammals. Common mammalian predators include: weasels (Mustelinae), skunks (Mephitinae), squirrels (Sciruidae), raccoons (Procyon lotor) and house cats (Felis domesticus). Additional nest predators include: voles and mice (Rodentia), shrews (Sorex), crows (Corvidae) and western meadowlarks (Sturnella neglecta). Up to 90 percent of lark nests may be lost to predators.
In response to nest predators, incubating females will flush silently when the predator is far from the nest; if the predator is close to the nest she will feign injury to draw it away. Young larks leave the nest early, this is thought to decrease predation and/or decrease the chance that an entire clutch is lost simultaneously.
Larks’ cryptic plumage allows them to blend in with the ground and makes it more difficult for predators to spot them; they will often avoid using patches of ground that do not match their coloration. Foraging in flocks is also thought to be an adaptation to reduce predation. (Beason, 1995; Campbell and Lack, 1985; Dean, et al., 1992; Mead, 1985; Simms, 1992)
- Anti-predator Adaptations
As insectivores, larks affect insect populations throughout their range.
Economic Importance for Humans: Positive
Larks can be important agents in the control of agricultural pests. For example, an adult skylark (Alauda arvensis) was found with 48 weevils (Sitona lineatus) in its stomach. This particular species of weevil is a pest on peas. (Simms, 1992)
- Positive Impacts
- controls pest population
Economic Importance for Humans: Negative
Because they occasionally feed in flocks in agricultural fields, larks are recognized as agricultural pests. In the United States they will damage crops of beets, broccoli, carrots, lettuce, peas, spinach, tomatoes, alfalfa, grain, sugar beets, cantaloupe and watermelon. (Clark and Hygnstrom, 1994)
- Negative Impacts
- crop pest
The IUCN lists two species of larks as critically endangered, two species as endangered and four as vulnerable. North American species are protected under the Migratory Bird Treaty Act. No larks are listed by CITES or ESA.
Declining numbers are the result of habitat loss, degradation and fragmentation, poisoning from chemicals used on crops and introduced species (especially those that are nest predators). Some species may stand to benefit from the clearing of forested areas to create pastures and arable land. ("UNEP-WCMC Species Database: CITES-Listed Species", 2003; Campbell, et al., 1997; IUCN, 2002; Mead, 1985; Simms, 1992; Threatened and Endangered Species System, 2003; U.S. Fish and Wildlife Service, date unknown)
- IUCN Red List [Link]
- Not Evaluated
Alaine Camfield (author), Animal Diversity Web.
Kari Kirschbaum (editor), Animal Diversity Web.
Living in Australia, New Zealand, Tasmania, New Guinea and associated islands.
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.
living in the southern part of the New World. In other words, Central and South America.
living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.
uses sound to communicate
living in landscapes dominated by human agriculture.
young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.
- bilateral symmetry
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
an animal that mainly eats meat
uses smells or other chemicals to communicate
- cooperative breeder
helpers provide assistance in raising young that are not their own
having a worldwide distribution. Found on all continents (except maybe Antarctica) and in all biogeographic provinces; or in all the major oceans (Atlantic, Indian, and Pacific.
having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.
- desert or dunes
in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.
- active during the day, 2. lasting for one day.
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
- female parental care
parental care is carried out by females
union of egg and spermatozoan
an animal that mainly eats leaves.
an animal that mainly eats fruit
an animal that mainly eats seeds
An animal that eats mainly plants or parts of plants.
a distribution that more or less circles the Arctic, so occurring in both the Nearctic and Palearctic biogeographic regions.
Found in northern North America and northern Europe or Asia.
An animal that eats mainly insects or spiders.
- internal fertilization
fertilization takes place within the female's body
referring to animal species that have been transported to and established populations in regions outside of their natural range, usually through human action.
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
- male parental care
parental care is carried out by males
makes seasonal movements between breeding and wintering grounds
eats mollusks, members of Phylum Mollusca
Having one mate at a time.
having the capacity to move from one place to another.
This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.
- native range
the area in which the animal is naturally found, the region in which it is endemic.
generally wanders from place to place, usually within a well-defined range.
an animal that mainly eats all kinds of things, including plants and animals
found in the oriental region of the world. In other words, India and southeast Asia.
reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.
having more than one female as a mate at one time
- seasonal breeding
breeding is confined to a particular season
remains in the same area
reproduction that includes combining the genetic contribution of two individuals, a male and a female
associates with others of its species; forms social groups.
uses touch to communicate
that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).
Living on the ground.
defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
- tropical savanna and grassland
A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.
A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.
- temperate grassland
A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.
A terrestrial biome with low, shrubby or mat-like vegetation found at extremely high latitudes or elevations, near the limit of plant growth. Soils usually subject to permafrost. Plant diversity is typically low and the growing season is short.
uses sight to communicate
2003. "UNEP-WCMC Species Database: CITES-Listed Species" (On-line). Accessed November 14, 2003 at http://www.cites.org/eng/resources/species.html.
Ali, S., S. Ripley. 1972. Handbook of the Birds of Indian and Pakistan, Volume 5. London: Oxford University Press.
Beason, R. 1995. Horned Lark (Eremophila alpestris). A Poole, F Gill, eds. The Birds of North America, Vol. 195. The Academy of Natural Sciences, Philadelphia, and The American Ornithologist's Union, Washington, D.C.
Campbell, B., E. Lack. 1985. A Dictionary of Birds. Vermillion: Buteo Books.
Campbell, R., N. Dawe, I. McTaggart-Cowan, J. Cooper, G. Kaiser, M. McNall, G. Smith. 1997. The Birds of British Columbia, Volume 3, Passerines, Flycatchers through Vireos. Vancouver: UBC Press.
Clark, J., S. Hygnstrom. 1994. "Horned Larks" (On-line). Prevention and Control of Wildlife Damage. Accessed October 29, 2003 at http://wildlifedamage.unl.edu/handbook/handbook/allPDF/bird_e63.pdf.
Dean, W., C. Fry, S. Keith, P. Lack. 1992. Family Alaudidae: Larks. Pp. 13-124 in S Keith, E Urban, C Fry, eds. The Birds of Africa, Vol. IV. London: Academic Press.
Gill, F. 1995. Ornithology, Second Edition. New York: W.H. Freeman and Company.
IUCN, 2002. "The IUCN Red List of Threatened Species" (On-line). Accessed October 29, 2003 at http://www.redlist.org/.
Mead, C. 1985. Larks, Wagtails and Pipits. Pp. 336-338 in C Perrins, A Middleton, eds. The Encyclopedia of Birds. New York: Facts on File Publications.
Payne, R. 2003. "Bird Families of the World" (On-line). Accessed October 27, 2003 at http://www.ummz.lsa.umich.edu/birds/Bird_Families_of_the_World.html.
Sibley, C., J. Ahlquist. 1990. Phylogeny and Classification of Birds, A study in Molecular Evolution. New Haven: Yale University Press.
Simms, E. 1992. British Larks, Pipits & Wagtails. London: Harper Collins Publishers.
Threatened and Endangered Species System, 2003. "U.S. Listed Vertebrate Animal Species Report by Taoxonomic Group" (On-line). Accessed November 14, 2003 at http://ecos.fws.gov/tess_public/TESSWebpageVipListed?code=V&listings=0#B.
Trost, C. 2001. Larks. Pp. 416-418 in C Elphick, J Dunning, D Sibley, eds. The Sibley Guide to Bird Life and Behavior. New York: Alfred A. Knopf.
U.S. Fish and Wildlife Service, date unknown. "Birds Protected by the Migratory Bird Treaty Act" (On-line). Accessed November 14, 2003 at http://migratorybirds.fws.gov/intrnltr/mbta/mbtintro.html.