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The dunnock (''Prunella modularis'') is a small
passerine A passerine () is any bird of the order Passeriformes (; from Latin 'sparrow' and '-shaped'), which includes more than half of all bird species. Sometimes known as perching birds, passerines are distinguished from other orders of birds by th ...
, or perching
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
, found throughout temperate Europe and into Asian Russia. Dunnocks have also been successfully introduced into New Zealand. It is by far the most widespread member of the
accentor The accentors are a genus of birds in the family Prunellidae, which is endemic to the Old World. This small group of closely related passerines are all in the genus ''Prunella''. All but the dunnock and the Japanese accentor are inhabitants of th ...
family; most other accentors are limited to mountain habitats. Other common names of the dunnock include: hedge accentor, hedge sparrow or hedge warbler.


Taxonomy

The dunnock was described by the Swedish naturalist
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
in 1758 in the tenth edition of his '' Systema Naturae''. He coined the binomial name of ''Motacilla modularis''. The specific epithet is from the Latin ''modularis'' "modulating" or "singing". This species is now placed in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
'' Prunella'' that was introduced by the French ornithologist Louis Vieillot in 1816. The name "dunnock" comes from the English ''dun'' (dingy brown, dark-coloured) and the diminutive ''ock'', and "accentor" is from post-classical
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
and means a person who sings with another. The genus name ''Prunella'' is from the German ''Braunelle'', "dunnock", a diminutive of ''braun'', "brown". Eight subspecies are recognised: * ''P. m. hebridium'' Meinertzhagen, R, 1934 – Ireland and the Hebrides (west of Scotland) * ''P. m. occidentalis'' (
Hartert Ernst Johann Otto Hartert (29 October 1859 – 11 November 1933) was a widely published German ornithologist. Life and career Hartert was born in Hamburg, Germany on 29 October 1859. In July 1891, he married the illustrator Claudia Bernadine E ...
, 1910) – Scotland (except the Hebrides), England, Wales and west France * ''P. m. modularis'' (
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, 1758) – north and central Europe * ''P. m. mabbotti'' Harper, 1919 – Iberian Peninsula, south-central France and Italy * ''P. m. meinertzhageni'' Harrison, JM & Pateff, 1937 – Balkans * ''P. m. fuscata'' Mauersberger, 1971 – south Crimean Peninsula (north coast of the Black Sea) * ''P. m. euxina'' Watson, 1961 – northwest and north Turkey * ''P. m. obscura'' ( Hablizl, 1783) – northeast Turkey, Caucasus and north Iran Some taxonomists have recently suggested that dunnock might be better treated as three species, with ''P. m. mabbotti'' and ''P. m. obscura'' being elevated from subspecies status.


Description

A
European robin The European robin (''Erithacus rubecula''), known simply as the robin or robin redbreast in Great Britain & Ireland, is a small insectivorous passerine bird that belongs to the chat subfamily of the Old World flycatcher family. About in len ...
-sized bird, the dunnock typically measures in length. It possesses a streaked back, somewhat resembling a small house sparrow. Like that species, the dunnock has a drab appearance which may have evolved to avoid predation. It is brownish underneath, and has a fine pointed bill. Adults have a grey head, and both sexes are similarly coloured. Unlike any similar sized small brown bird, dunnocks exhibit frequent wing flicking, especially when engaged in territorial disputes or when competing for mating rights. This gives rise to one of their common names, "shufflewing". The main call of the dunnock is a shrill, persistent ''tseep'' along with a high trilling note, which betrays the bird's otherwise inconspicuous presence. The song is rapid, thin and tinkling, a sweet warble which can be confused with that of the
Eurasian wren The Eurasian wren (''Troglodytes troglodytes'') or northern wren is a very small insectivorous bird, and the only member of the wren family Troglodytidae found in Eurasia and Africa (Maghreb). In Anglophone Europe, it is commonly known simply a ...
, but is shorter and weaker.


Distribution and habitat

Dunnocks are native to large areas of Eurasia, inhabiting much of Europe including Lebanon, northern Iran, and the
Caucasus The Caucasus () or Caucasia (), is a region between the Black Sea and the Caspian Sea, mainly comprising Armenia, Azerbaijan, Georgia (country), Georgia, and parts of Southern Russia. The Caucasus Mountains, including the Greater Caucasus range ...
. They are the only commonly found accentor in lowland areas; all the others inhabit upland areas. Dunnocks were successfully introduced into New Zealand during the 19th century, and are now widely distributed around the country and some offshore islands. Favoured habitats include
woodland A woodland () is, in the broad sense, land covered with trees, or in a narrow sense, synonymous with wood (or in the U.S., the ''plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunlight and limited shade (se ...
s, shrubs,
garden A garden is a planned space, usually outdoors, set aside for the cultivation, display, and enjoyment of plants and other forms of nature. The single feature identifying even the wildest wild garden is ''control''. The garden can incorporate bot ...
s, and hedgerows where they typically feed on the ground, often seeking out detritivores as food.


Territoriality

Dunnocks are territorial and may engage in conflict with other birds that encroach upon their nests. Males sometimes share a territory and exhibit a strict dominance hierarchy. Nevertheless, this social dominance is not translated into benefits to the alpha male in terms of reproduction, since paternity is usually equally shared between males of the group. Furthermore, members of a group are rarely related, and so competition can result. Female territorial ranges are almost always exclusive. However, sometimes, multiple males will co-operate to defend a single territory containing multiple females. Males exhibit a strong
dominance hierarchy In biology, a dominance hierarchy (formerly and colloquially called a pecking order) is a type of social hierarchy that arises when members of animal social groups interact, creating a ranking system. A dominant higher-ranking individual is so ...
within groups: older birds tend to be the dominant males and first-year birds are usually sub-dominant. Studies have found that close male relatives almost never share a territory. The male's ability to access females generally depends on female range size, which is affected by the distribution of food. When resources are distributed in dense patches, female ranges tend to be small and easy for males to monopolise. Subsequent mating systems, as discussed below, reflect high reproductive success for males and relatively lower success for females. In times of scarcity, female territories expand to accommodate the lack of resources, causing males to have a more difficult time monopolising females. Hence, females gain a reproductive advantage over males in this case.


Breeding


Mating systems

The dunnock possesses variable mating systems. Females are often
polyandrous Polyandry (; ) is a form of polygamy in which a woman takes two or more husbands at the same time. Polyandry is contrasted with polygyny, involving one male and two or more females. If a marriage involves a plural number of "husbands and wive ...
, breeding with two or more males at once, which is quite rare among birds. This multiple mating system leads to the development of
sperm competition Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and ...
amongst the male suitors.
DNA fingerprinting DNA profiling (also called DNA fingerprinting) is the process of determining an individual's DNA characteristics. DNA analysis intended to identify a species, rather than an individual, is called DNA barcoding. DNA profiling is a forensic tec ...
has shown that chicks within a brood often have different fathers, depending on the success of the males at monopolising the female. Males try to ensure their paternity by pecking at the cloaca of the female to stimulate ejection of rival males' sperm. Dunnocks take just one-tenth of a second to copulate and can mate more than 100 times a day. Males provide parental care in proportion to their mating success, so two males and a female can commonly be seen provisioning nestlings at one nest. Other mating systems also exist within dunnock populations, depending on the ratio of male to females and the overlap of territories. When only one female and one male territory overlap,
monogamy Monogamy ( ) is a form of dyadic relationship in which an individual has only one partner during their lifetime. Alternately, only one partner at any one time (serial monogamy) — as compared to the various forms of non-monogamy (e.g., polyg ...
is preferred. Sometimes, two or three adjacent female territories overlap one male territory, and so
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any ...
is favored, with the male monopolising several females. Polygynandry also exists, in which two males jointly defend a territory containing several females. Polyandry, though, is the most common mating system of dunnocks found in nature. Depending on the population, males generally have the best reproductive success in polygynous populations, while females have the advantage during polyandry. Studies have illustrated the fluidity of dunnock mating systems. When given food in abundance, female territory size is reduced drastically. Consequently, males can more easily monopolise the females. Thus, the mating system can be shifted from one that favours female success (polyandry), to one that promotes male success (monogamy, polygynandry, or polygyny).


Nest

The dunnock builds a nest (predominantly from twigs and moss and lined with soft materials such as wool or feathers), low in a bush or conifer, where adults typically lay three to five unspotted blue
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
.


Parental care and provisioning

Broods, depending on the population, can be raised by a lone female, multiple females with the part-time help of a male, multiple females with full-time help by a male, or by multiple females and multiple males. In pairs, the male and the female invest parental care at similar rates. However, in trios, the female and alpha male invest more care in chicks than does the beta male. In territories in which females are able to escape from males, both the alpha and beta males share provisioning equally. This last system represents the best case scenario for females, as it helps to ensure maximal care and the success of the young. A study has found that males tend to not discriminate between their own young and those of another male in polyandrous or polygynandrous systems. However, they do vary their feeding depending on the certainty of paternity. If a male has greater access to a female, and therefore a higher chance of a successful fertilisation, during a specific mating period, it would provide more care towards the young.


References


External links


Xeno-canto: audio recordings of the dunnock

Dunnock videos, photos & sounds
on the Internet Bird Collection
Ageing and sexing (PDF; 2.0 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze


{{Authority control Prunellidae Birds of Europe Birds of New Zealand Birds described in 1758 Taxa named by Carl Linnaeus