Thetree sparrow (Passer montanus) is apasserine bird in thesparrow family with a rich chestnutcrown andnape and a black patch on each pure white cheek. The sexes have similarplumage, and young birds are duller versions of the adult. This sparrow breeds across much oftemperate Eurasia andSoutheast Asia and has been introduced elsewhere, including theUnited States, where it is known as the Eurasian tree sparrow orGerman sparrow to differentiate it from the nativeAmerican tree sparrow. Although several subspecies are recognised, the appearance of this bird varies little across its extensive range.
The untidy nest of the tree sparrow is built in a natural cavity, a hole in a building, or the disused nest of aEuropean magpie orwhite stork. The typicalclutch is five or six eggs, which hatch in under two weeks. This sparrow mainly feeds on seeds, but also consumes invertebrates, particularly during the breeding season. As with other small birds, infection by parasites and diseases, as well as predation bybirds of prey, take their toll, and the typical lifespan is about two years.
The tree sparrow is widespread in the towns and cities of easternAsia, but inEurope, it inhabits lightly wooded open countryside, while thehouse sparrow breeds in more urban areas. The tree sparrow's extensive range and large population mean it is not globally endangered, although there have been significant population declines in Western Europe, partly due to changes in farming practices involving increased use ofherbicides and the loss of winterstubble fields. In eastern Asia and westernAustralia, this species is sometimes considered a pest, though it is also widely celebrated in oriental art.
The tree sparrow is 12.5–14 cm (5–5+1⁄2 in) long,[2] with awingspan of about 21 cm (8.3 in) and a weight of 24 g (0.85 oz),[3] making it roughly 10% smaller than the house sparrow.[4] The adult's crown and nape are richchestnut, and there is a kidney-shaped black ear patch on each pure white cheek; the chin, throat, and the area between the bill and throat are black. The upperparts are light brown, streaked with black, and the brown wings have two distinct narrow white bars. The legs are pale brown, and the bill is lead-blue in summer, becoming almost black in winter.[5]
This sparrow is distinctive within its genus because there are no plumage differences between the sexes. Juveniles also resemble adults, although their colours tend to be duller.[6] Its contrasting face pattern makes this species easily identifiable in all plumages;[4] the smaller size and brown, not grey, crown are additional differences from the male house sparrow.[2] Both adult and juvenile tree sparrows undergo a slow, completemoult in autumn and experience an increase in body mass, despite a reduction in stored fat. This change in mass is due to an increase in blood volume to support active feather growth, as well as a higher overall body water content.[7]
The tree sparrow does not have a true song, but itsvocalisations include an excited series oftschip calls made by unpaired or courting males. Other monosyllabic chirps are used for social interaction, and the flight call is a harshteck.[4] A study comparing the vocalisations of the introduced Missouri population with those of birds fromGermany showed that the US birds had fewer shared syllable types (memes) and more structure within the population than the European sparrows. This may be due to the small size of the original North American population, resulting in a loss ofgenetic diversity.[8]
Description of the house and tree sparrows from theSystema naturae[9]
TheOld World sparrow genusPasser is a group of smallpasserine birds that is believed to have originated in Africa, and which contains 15–25 species depending on the authority.[10] Its members are typically found in open, lightly wooded, habitats, although several species, notably thehouse sparrow (P. domesticus) have adapted to human habitations. Most species in the genus are typically 10–20 cm (3.9–7.9 in) long, predominantly brown or greyish birds with short square tails and stubby conical beaks. They are primarily ground-feeding seed-eaters, although they also consume invertebrates, especially when breeding.[11] The Eurasian species is not closely related to theAmerican tree sparrow (Spizelloides arborea), which is in a different family, theNew World sparrows.[12]
The tree sparrow's binomial name is derived from twoLatin words:passer, "sparrow", andmontanus, "of the mountains" (frommons "mountain").[3] The tree sparrow was first described byCarl Linnaeus in his 1758Systema Naturae asFringilla montana,[13] but, along with the house sparrow, it was soon moved from thefinches (family Fringillidae) into the new genusPasser created by French zoologistMathurin Jacques Brisson in 1760.[14] The tree sparrow's common name is given because it prefers tree holes for nesting. This name, and the scientific namemontanus, do not appropriately describe this species's habitat preferences: theGerman nameFeldsperling and theHungarian namemezei veréb ("field sparrow") comes closer to doing so.[15]
This species varies little in appearance across its large range, and the differences between the seven extantsubspecies recognised by Clement are slight. At least 15 other subspecies have been proposed, but these are considered intermediates of the recognised subspecies.[5][16]
TheCaucasian tree sparrow (P. m. transcaucasicus), described bySergei Aleksandrovich Buturlin in 1906, breeds from the southern regions of the Caucasus east to northernIran. It is duller and greyer than the nominate subspecies.[5]
TheAfghan tree sparrow (P. m. dilutus), described byCharles Wallace Richmond in 1856, is resident in extreme northeastern Iran, northernPakistan and northwesternIndia. It also occurs further north, from Uzbekistan and Tajikistan east toChina. Compared toP. m. montanus, it is paler, with sandy-brown upperparts.[5]
TheTibetan tree sparrow (P. m. tibetanus), the largest subspecies by size, was described byStuart Baker in 1925. It is found in the northernHimalayas, fromNepal east throughTibet to northwestern China. It resemblesP. m. dilutus, but is darker.[5]
P. m. malaccensis, described byAlphonse Dubois in 1885, is found from the southern Himalayas east toHainan andIndonesia. It is a dark-coloured subspecies, likeP. m. saturatus, but is smaller and more heavily streaked on its upperparts.[5]
P. m. hepaticus, described bySidney Dillon Ripley in 1948, breeds from northeasternAssam to northwesternBurma. It is similar in appearance toP. m. saturatus, but redder on its head and upperparts.[5]
Urban nest under a roof tile of a wooden house in JapanTree sparrows inJapan
The tree sparrow's natural breeding range comprises most oftemperate Europe and Asia south of aboutlatitude 68°N (north of this the summers are too cold, with July average temperatures below 12 °C (54 °F)) and through Southeast Asia toJava andBali. It formerly bred in theFaroes,Malta andGozo.[4][5] In South Asia it is found mainly in the temperate zone.[17][18] It is sedentary over most of its extensive range, but northernmost breeding populationsmigrate south for the winter,[19] and small numbers leave southern Europe for North Africa and the Middle East.[4] The eastern subspeciesP. m. dilutus reaches coastal Pakistan in winter and thousands of birds of this race move through eastern China in autumn.[5]
The tree sparrow has beenintroduced outside its native range but has not always become established, possibly due to competition with the house sparrow. It was introduced successfully toSardinia,[20][21] eastern Indonesia, the Philippines andMicronesia, but introductions to New Zealand and Bermuda did not take root.[22] Ship-carried birds colonisedBorneo. This sparrow has occurred as a naturalvagrant toGibraltar, Tunisia, Algeria, Egypt, Israel, the United Arab Emirates,[5] Morocco and Iceland.[1]In North America, a population of about 15,000 birds has become established aroundSt. Louis and neighbouring parts of Illinois and southeastern Iowa.[23] These sparrows are descended from 20 birds (or 12 pairs, depending on the source) imported from Germany and released in late April 1870, probably by members of anacclimatisation society.[22]
Within its limited US range of about 22,000 square kilometres (8,500 sq mi),[24] the tree sparrow has to compete with the house sparrow in urban centres, and is therefore mainly found in parks, farms and rural woods.[8][25] It has been suggested that one factor limiting the spread of tree sparrows in North America is the species' higherneophobia (novelty aversion) compared to house sparrows; one study specifically found that captive tree sparrows showed lower willingness to eat novel foods and habituated more slowly to novel objects than house sparrows.[26] The American population is sometimes referred to as the "German sparrow", to distinguish it from both the native American tree sparrow species and the much more widespread "English" house sparrow.[27]
Despite its scientific name,Passer montanus, this is not typically a mountain species, and only reaches 700 m (2,300 ft) in Switzerland, although it has bred at 1,700 m (5,600 ft) in the northern Caucasus and as high as 4,270 m (14,010 ft) in Nepal.[4][5] In Europe, it is frequently found on coasts with cliffs, in empty buildings, inpollarded willows along slow water courses, or in open countryside with small isolated patches of woodland.[4] The tree sparrow shows a strong preference for nest-sites nearwetland habitats and avoids breeding on intensively managed mixed farmland.[29]
When the tree sparrow and the larger house sparrow are found in the same area, the house sparrow usually breeds in urban areas, while the smaller tree sparrow nests in the countryside.[5] Where trees are scarce, as in Mongolia, both species may use man-made structures for nesting.[30] The tree sparrow is a rural bird in Europe, but is an urban bird in eastern Asia. In southern and central Asia, both species of the genusPasser may be found around towns and villages.[5] In parts of the Mediterranean, such as Italy, both the tree and the Italian or Spanish sparrows may be found in settlements.[31] In Australia, the tree sparrow is largely an urban bird, whereas the house sparrow uses more natural habitats.[28]
The tree sparrow reaches breeding maturity within a year of hatching,[32] and typically builds its nest in a cavity in an old tree or rock face. Some nests are not situated in holes per se, but are built among roots of overhanginggorse or similar bushes.[33] Roof cavities in houses may also be used.[33] In the tropics, the crown of a palm tree or the ceiling of a veranda may serve as a nest site.[34] This species will breed in the disused domed nest of aEuropean magpie,[33] or an active or unused stick nest of a large bird such as thewhite stork,[35]white-tailed eagle,osprey,black kite orgrey heron. It will sometimes attempt to take over the nest of other birds that breed in holes or enclosed spaces, such as thebarn swallow,house martin,sand martin orEuropean bee-eater.[36]
Pairs may breed in isolation or loose colonies,[37] and will readily usenest boxes. A Spanish study found that boxes made from a mixture of wood and concrete (woodcrete) had a much higher occupancy rate than wooden boxes (76.5% versus 33.5%). Birds nesting in woodcrete boxes laid their eggs earlier, had a shorter incubation period, and made more breeding attempts per season. There was no difference in clutch size or chick condition between nest box types, but reproductive success was higher in woodcrete, perhaps because the synthetic nests were 1.5 °C warmer than wooden ones.[38]
In spring, the male calls from near the nest site to proclaim ownership and attract a mate. He may also carry nest material into the nest hole.[5] These behaviours are repeated in autumn. The preferred locations for the autumn display are old tree sparrow nests, especially those where nestlings have hatched. Empty nest boxes, and sites used by house sparrows or other hole nesting birds, such astits,pied flycatchers orcommon redstarts, are rarely used for the autumn display.[39]
The untidy nest is made of hay, grass,wool or other materials and is lined withfeathers,[33] which improve its thermal insulation.[40] A complete nest consists of three layers; base, lining and dome.[39] The typical clutch is five or sixeggs (rarely more than four in Malaysia),[34] white to pale grey and heavily marked with spots, small blotches, or speckling;[41] they are 20 mm × 14 mm (0.79 in × 0.55 in) in size and weigh 2.1 g (0.074 oz), of which 7% is shell.[3] The eggs are incubated by both parents for 12–13 days before thealtricial, naked chicks hatch. A further 15–18 days elapse before theyfledge. Two or three broods may be raised each year.[3] Birds breeding in colonies produce more eggs and fledglings from their first broods than solitary pairs do, but the reverse is true for second and third clutches.[42] Females that copulate frequently tend to lay more eggs and have a shorter incubation period. Therefore, within-pair mating may indicate the reproductive ability of a pair.[43] There is a significant level of promiscuity; in a Hungarian study, more than 9% of chicks were sired by extra-pair males, and 20% of the broods contained at least one extra-pair young.[44]
Hybridisation between tree sparrows and house sparrows has been recorded in many parts of the world. Male hybrids tend to resemble tree sparrows, while females tend to resemble house sparrows.[45] A breeding population in theEastern Ghats of India,[46] said to be introduced,[5] may also hybridise with house sparrows.[18] On at least one occasion a mixed pair has resulted in fertile young.[47][48][49] A wild hybridisation with the resident sparrows of Malta, which are intermediate between theSpanish sparrow (P. hispaniolensis) andItalian sparrows (P. italiae), was recorded in Malta in 1975.[5]
Nest box made of wood and concreteA very social bird, no problem sharing abird feeder
The tree sparrow is a bird that predominantly eats seeds and grains. It feeds on the ground in flocks, often alongside house sparrows, finches, or buntings. Its diet includes weed seeds, such aschickweeds andgoosefoot, spilled grain,[5] and it may also visitfeeding stations, especially forpeanuts. During the breeding season, when the young are fed mainly on animal food, it will also feed oninvertebrates, such as insects,woodlice,millipedes,centipedes,spiders andharvestmen.[50]
Adults use a variety of wetlands when foraging for invertebrate prey to feed their young. Aquatic sites play a key role in providing the adequate diversity and availability of suitable invertebrate prey necessary for the successful rearing of chicks throughout the long breeding season of this multi-brooded species. However, large areas of formerly occupied farmland no longer provide these invertebrate resources due to the effects ofintensive farming. The availability of supplementary seed food within 1 kilometre (0.62 mi) of the nest site does not influence nest-site choice, or affect the number of young raised.[29]
In winter, seed resources are most likely to be a key limiting factor.[29] At this time of year, individuals in a flock establish lineardominance hierarchies, but the size of the throat patch does not strongly correlate with position in the hierarchy. This is in contrast to the house sparrow, in which fights to establish dominance are reduced by displaying the throat patch. The size of this patch acts as asignalling "badge" offitness.[51] Although there is evidence that the black throat patch of male, but not female, tree sparrows predicts fighting success in foraging flocks.[52]
The risk of predation affects feeding strategies. One study showed that when the distance between shelter and food sources increased, birds visited feeders in smaller flocks, spent less time there, and became more vigilant when far from shelter. Sparrows can feed as "producers", searching for food directly, or as "scroungers", just joining other flock members who have already found food. Scrounging was 30% more likely at exposed feeding sites, although this was not due to increased anti-predator vigilance. One possible explanation is that individuals with lower fat reserves use riskier places.[53]
Many species ofbird lice are present on the birds and in their nests,[63][64] and mites of the genusKnemidocoptes have been known to infest populations, resulting in lesions on the legs and toes.[65] Parasitisation of nestlings byProtocalliphorablow-fly larvae is a significant factor in nestling mortality.[66] Egg size does not influence nestling mortality, but chicks from large eggs grow faster.[67]
Tree sparrows are susceptible to bacterial and viral infections. It has been shown that bacteria play an important role in eggs failing to hatch and in nestling mortality,[68] and mass deaths due toSalmonella infection have been noted in Japan.[69]Avian malaria parasites have been found in the blood of many populations,[70] and birds in China were found to harbour a strain ofH5N1 that was highly virulent to chickens.[71]
The immune response of tree sparrows is weaker than that of house sparrows, which has been suggested as an explanation for the latter's greater invasive potential.[72] The house sparrow and tree sparrow are the most frequent victims ofroadkill on the roads of Central, Eastern and Southern Europe.[73] The maximum recorded age is 13.1 years,[32] but three years is a typical lifespan.[3]
Tree sparrows exhibit various physiological adaptations to seasonal changes in temperature and photoperiod. When exposed to shorter photoperiods and colder temperatures in winter, tree sparrows increase their body mass andbasal metabolic rate, as well as the activity of thecytochrome c oxidase enzyme andthyroid hormone hormones. Temperature has a greater effect than photoperiod.[74] Increasing body mass allows these birds to store more energy, while their increased basal metabolic rate is due to the higher amount of energy needed to maintain their body temperature in colder conditions. Another study also found that during the winter, tree sparrows also exhibited higher levels of thyroid hormone T3, which helps regulate the body's use of energy.[75] The birds' increased T3 levels was correlated with increased heat production.
The tree sparrow has an estimated range of 98.3 million square kilometres (38.0 million sq mi) and a population of 190–310 million individuals. Although its population is declining, the species is not believed to be approaching the thresholds for the population decline criterion of theIUCN Red List (that is, declining more than 30% in 10 years or three generations). For these reasons, the species'conservation status is evaluated at the global level as being ofLeast Concern.[1]
After the ending of theFour Pests campaign, the species was extirpated from China, which afterwards imported tree sparrows from Russia to recover its population.[77] Although the tree sparrow has been expanding its range inFennoscandia and eastern Europe, populations have been declining in much of western Europe,[4][78] a trend reflected in other farmland birds such as theskylark,corn bunting andnorthern lapwing. From 1980 to 2003, common farmland bird numbers fell by 28%.[76] The collapse in populations seems to have been particularly severe in Great Britain, where there was a 95% decline between 1970 and 1998,[79] and Ireland, which had only 1,000–1,500 pairs in the late 1990s.[4][80] In the British Isles, such declines may be due to natural fluctuations, to which tree sparrows are known to be prone.[31] Breeding performance has improved substantially as population sizes have decreased, suggesting that decreases in productivity were not responsible for the decline and that survival was the critical factor.[81] The large decline in tree sparrow numbers is probably the result of agricultural intensification and specialisation, particularly the increased use of herbicides and a trend towards autumn-sown crops (at the expense of spring-sown crops that produce stubble fields in winter). The change from mixed to specialised farming and the increased use of insecticides has reduced the amount of insect food available for nestlings.[76]
The tree sparrow is considered a pest in some areas. In Australia, for example, it damages many cereal and fruit crops, and its droppings spoil cereal crops, animal feed and stored grain.Quarantine rules prohibit the transport of this species into Western Australia.[28]
In the 19th century, mass campaigns to catch and kill sparrows were launched in several European countries, including France, Prussia, Hungary, Baden.[82]
In April 1958, theParamount leader of China,Mao Zedong, attempted to reduce the damage to crops caused by tree sparrows, which was estimated to be 4.5 kg (9.9 lb) of grain per bird each year, by mobilising millions of people and many scarecrows to drive the birds to death by exhaustion. Although this was successful in reducing the sparrow population, the "Four Pests campaign" overlooked the fact that the birds also consumed large numbers of locusts and other insect pests. Crop yields subsequently collapsed, exacerbating a famine that led to the deaths of 30 million people between 1959 and 1961.[23][83] The tree sparrow's consumption of insects has led to its use in agriculture to control fruit tree pests and thecommon asparagus beetle (Crioceris aspergi).[84]
The tree sparrow has long been a popular subject in Chinese and Japanese art. It is often depicted either perched on a plant spray or flying in a flock.[83] Representations by Oriental artists, includingHiroshige, have featured on postage stamps issued byAntigua and Barbuda, the Central African Republic, China, and the Gambia. More straightforward illustrations were used on stamps issued by Belarus, Belgium, Cambodia, Estonia, and Taiwan.[85] The fluttering of the bird gave rise to a traditional Japanese dance, theSuzume Odori, developed inSendai, which was depicted by artists such asHokusai.[86]
In the Philippines, where it is one of several species referred to asmaya, and is sometimes specifically referred to as the "mayang simbahan" ("church maya" or "church sparrow"), the tree sparrow is the most common bird in the cities. Many urban Filipinos confuse it with the former national bird of the Philippines, theblack-headed munia – also called amaya, but specifically differentiated infolk taxa as the "mayang pula" ("red maya").[87]
^Although Linnaeus gives the location as simplyin Europa, the type specimen was fromBagnacavallo, Italy (Clancy, Philip Alexander (1948). "Remarks onPasser montanus (Linnaeus) in the Western Palaearctic Region with special reference toPasser catellatus Kleinschmidt, 1935: England".Bulletin of the British Ornithologists' Club.68: 135.) Linnaeus's text for the tree sparrow translates "F[inch]. With dusky wings and tail, black and grey body paired white wing bars."
^del Hoyo, Josep; Elliott, Andrew; Sargatal, Jordi; Christie, David A.; de Juana, Eduardo, eds. (2013)."Eurasian Tree Sparrow (Passer montanus)".Handbook of the Birds of the World Alive. Barcelona: Lynx Edicions. Retrieved25 October 2018.
^Barlow, Jon C.; Leckie, Sheridan N. Pool, A. (ed.)."Eurasian Tree Sparrow (Passer montanus)".The Birds of North America Online. Ithaca: Cornell Laboratory of Ornithology. Retrieved30 January 2009.
^abcMassam, Marion."Sparrows"(PDF).Farmnote No. 117/99. Agriculture Western Australia. Archived fromthe original(PDF) on 12 August 2008. Retrieved1 February 2009.
^Barta, Zoltán; Liker, András; Mónus, Ferenc (February 2004). "The effects of predation risk on the use of social foraging tactics".Animal Behaviour.67 (2):301–308.doi:10.1016/j.anbehav.2003.06.012.S2CID53180647.
^Bertolino, Sandro; Ghiberti, Elena; Perrone, Aurelio (2001). "Feeding ecology of the long-eared owl (Asio otus) in northern Italy: is it a dietary specialist?".Canadian Journal of Zoology.79 (12):2192–2198.Bibcode:2001CaJZ...79.2192B.doi:10.1139/z01-182.
^Cordero, P. J.; Salaet, M. "Breeding season, population and reproduction rate of the tree sparrow (Passer montanus, L.) in Barcelona, NE Spain." inPinowski & Summers-Smith 1990, pp. 169–177
^Mainka, S. A.; Melville D. S.; Galsworthy A.; Black, S. R. (1994). "Knemidocoptes sp. on wild passerines at the Mai Po nature reserve, Hong Kong".Journal of Wildlife Diseases.30 (2):254–256.doi:10.7589/0090-3558-30.2.254.PMID8028111.S2CID40000445.
^Puchala, Peter (2004). "Detrimental effects of larval blow flies (Protocalliphora azurea) on nestlings and breeding success of Tree Sparrows (Passer montanus)".Canadian Journal of Zoology.82 (8):1285–1290.Bibcode:2004CaJZ...82.1285P.doi:10.1139/z04-111.
^Zheng, Wei-Hong; Li, Ming; Liu, Jin-Song; Shao, Shu-Li; Xu, Xing-Jun (2014). "Seasonal variation of metabolic thermogenesis in Eurasian tree sparrows (Passer montanus) over a latitudinal gradient".Physiological and Biochemical Zoology.87 (5):704–718.doi:10.1086/676832.ISSN1537-5293.PMID25244382.
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