{{Short description|Species of bird}} {{EngvarB|date=August 2023}} {{Use dmy dates|date=April 2017}} {{Speciesbox | status = LC | status_system = IUCN3.1 | status_ref = <ref name="iucn status 13 November 2021">{{cite iucn |author=BirdLife International |date=2018 |title=''Prunella modularis'' |volume=2018 |article-number=e.T22718651A132118966 |doi=10.2305/IUCN.UK.2018-2.RLTS.T22718651A132118966.en |access-date=13 November 2021}}</ref> | image = {{CSS image crop |Image = Dunnock (Prunella modularis) 3.jpg |bSize = 360 |cWidth = 220 |cHeight = 200 |oTop = 40 |oLeft = 110 }}
| image2 = Dunnock (Prunella modularis) (W1CDR0001422 BD6).ogg | image2_caption= Song recorded on Dartmoor in Devon, England | taxon = Prunella modularis | authority= (Linnaeus, 1758) | synonyms = * ''Motacilla modularis'' {{small|Linnaeus, 1758}} * ''Accentor modularis'' {{small|(Linnaeus, 1758)}} | range_map = PrunellaModularisIUCN.png | range_map_caption = Global range{{leftlegend|#00FF00|Summer range|outline=gray}}{{leftlegend|#008000|Year-round range|outline=gray}}{{leftlegend|#007FFF|Winter range|outline=gray}}{{leftlegend|#FFFF00|Non-native range|outline=gray}} }} thumb|''Prunella modularis''
The '''dunnock''' ('''''Prunella modularis''''') is a small passerine, or perching bird, found throughout temperate Europe and into Asian Russia. Dunnocks have also been introduced and become common in New Zealand. It is the most widespread member of the accentor family; most other accentors are limited to mountain habitats. Other, largely archaic, English names for the dunnock include '''hedge accentor''', '''hedge sparrow''', '''hedge warbler''', and '''titling'''.<ref>{{cite web | url=https://en.wiktionary.org/wiki/titling#Etymology_2 | title=Titling | date=3 November 2023 }}</ref>
==Taxonomy== The dunnock was described by the Swedish naturalist Carl Linnaeus in 1758 in the tenth edition of his ''Systema Naturae''. He coined the binomial name of ''Motacilla modularis''.<ref>{{cite book | last=Linnaeus | first=Carl | author-link=Carl Linnaeus | year=1758 | title= Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis | volume=1| edition=10th | page=184 | publisher=Holmiae:Laurentii Salvii | language=la | url= https://biodiversitylibrary.org/page/727091 }}</ref> The specific epithet is from the Latin {{Lang|la|modularis}} "modulating" or "singing".<ref name=hbwkey>{{cite web | last=Jobling | first=J.A. | year=2019 | title= Key to Scientific Names in Ornithology | editor1-last=del Hoyo | editor1-first=J. | editor2-last=Elliott | editor2-first=A. | editor3-last=Sargatal | editor3-first=J. | editor4-last=Christie | editor4-first=D.A. | editor5-last=de Juana | editor5-first=E. | work=Handbook of the Birds of the World Alive | publisher=Lynx Edicions | url=https://www.hbw.com/dictionary/definition/modularis | access-date=10 March 2019 }}</ref> This species is now placed in the genus ''Prunella'' that was introduced by the French ornithologist Louis Pierre Vieillot in 1816.<ref>{{cite book | last=Vieillot | first=Louis Pierre | author-link=Louis Pierre Vieillot | year=1816 | title=Analyse d'une Nouvelle Ornithologie Élémentaire | publisher=Deterville/self | location=Paris | page = 43 | language=fr| url=https://gallica.bnf.fr/ark:/12148/bpt6k9745205x/f49.image }}<!--BHL has a scan of an 1883 reprint - same pagination http://biodiversitylibrary.org/page/12830237 --></ref>
The name "dunnock" comes from the English ''dun'' (dingy brown, dark-coloured) and the diminutive form of bird ''ock'' (thus, the original "little brown bird"),<ref name="Lockwood">{{cite book|last = Lockwood| first = W. B.|title = Oxford Book of British Bird Names|publisher = Oxford University Press|year = 1984}}</ref><ref name=OED1>{{ OED |Dunnock}}</ref> while "accentor" is from post-classical Latin and means a person who sings with another.<ref name=OED>{{ OED |Accentor}}</ref> The genus name ''Prunella'' is from the German {{Lang|de|Braunelle}}, "dunnock", a diminutive of {{Lang|de|braun}}, "brown".<ref name =job>{{cite book | last= Jobling | first= James A. | year= 2010| title= The Helm Dictionary of Scientific Bird Names | publisher=Christopher Helm | location = London, United Kingdom | isbn = 978-1-4081-2501-4 | page =318}}</ref>
Seven subspecies are accepted by the IOC list:<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor1-link=Frank Gill (ornithologist) | editor2-last=Donsker | editor2-first=David | year=2024 | title=Waxbills, parrotfinches, munias, whydahs, Olive Warbler, accentors, pipits | work=World Bird List Version 14.1 | url=https://www.worldbirdnames.org/bow/waxbills/ | publisher=International Ornithologists' Union | access-date=1 August 2024 }}</ref>
* ''P. m. hebridium'' Meinertzhagen, R, 1934 – Ireland and the Hebrides (west of Scotland) * ''P. m. occidentalis'' (Hartert, 1910) – Scotland (except the Hebrides), England, Wales and west France * ''P. m. modularis'' (Linnaeus, 1758) – north and central, and southeast Europe (including the formerly accepted ''P. m. meinertzhageni'' of the 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. mabbotti'' Harper, 1919 – Iberian Peninsula, south-central France and Italy * ''P. m. obscura'' (Hablizl, 1783) – northeast Turkey, Caucasus and north Iran
[[File:Prunella modularis - Dağ bülbülü 01-1.jpg|thumb|none|''P. m. obscura'', with its browner head at all ages, is the most distinct subspecies; here at Giresun, northeastern Turkey.]]
Acceptance of these seven subspecies has not been universal; Shirihai & Svensson (2018) accept only three subspecies, ''P. m. occidentalis'' (including ''P. m. hebridium''), ''P. m. modularis'' (including ''P. m. euxina, P. m. mabbotti''), and ''P. m. obscura'' (including ''P. m. fuscata'').<ref name=Shirihai>{{Cite book|title=Handbook of Western Palearctic Birds, Volume 1: Passerines: Larks to Warblers|publisher=Bloomsbury Publishing|date=2018-08-27|isbn=978-1-4729-3758-2|language=en|first1=Hadoram|last1=Shirihai|first2=Lars|last2=Svensson}}</ref> Another study however recently suggested that dunnock might be better treated as three species, with ''P. m. mabbotti'' and ''P. m. obscura'' being elevated from subspecies status to separate species.<ref>{{cite journal |last1=Pavia |first1=Marco |last2=Drovetski |first2=Sergei V. |last3=Boano |first3=Giovanni |last4=Conway |first4=Kevin W. |last5=Pellegrino |first5=Irene |last6=Voelker |first6=Gary |title=Elevation of two subspecies of Dunnock Prunella modularis to species rank |journal=Bulletin of the British Ornithologists' Club |date=15 June 2021 |volume=141 |issue=2 |pages=199–210 |doi=10.25226/bboc.v141i2.2021.a10|doi-access=free }}</ref>
==Description== A robin-sized bird, the dunnock typically measures {{cvt|13|–|14.5|cm}} in length. It has a brown back streaked blackish, somewhat resembling a small house sparrow. Like that species, the dunnock has a drab appearance which may have evolved as camouflage to avoid predation. It is brownish underneath, and has a fine pointed bill. Adults have a dull grey head, and both sexes are similarly coloured; juveniles are browner on the head, looser, 'fluffy' feathering, and more obviously streaked overall.<ref name=Collins>Svensson, L., Mullarney, K., & Zetterström, D. (2009) ''Collins Bird Guide'', ed. 2. {{ISBN|0-00-219728-6}}, pages 274-275</ref><ref name=Shirihai/><ref name="field guide">{{cite book |last1=Heather |first1=Barrie |last2=Rogertson |first2=Hugh |title=The Field Guide to the Birds of New Zealand |year=2005 |edition=Revised |publisher=Viking Press }}</ref> Unlike any similar sized small brown bird in Europe, dunnocks exhibit frequent wing flicking, especially when engaged in territorial disputes or when competing for mating rights.<ref>{{cite web|url=https://www.rspb.org.uk/birds-and-wildlife/bird-and-wildlife-guides/bird-a-z/d/dunnock/|title=Dunnock|publisher= RSPB|access-date=6 May 2017}}</ref> This gave rise to the old nickname of "shufflewing".<ref>{{cite book|title=Webster's Revised Unabridged Dictionary|date=1913|publisher=G. & C. Merriam|url=https://www.websters1913.com/words/Shufflewing|access-date=2 February 2023}}</ref>
The main call of the dunnock is a shrill, persistent ''tseep'' along with a short, weak trilling note, which betrays the bird's otherwise inconspicuous presence. The song is rapid, thin and tinkling, a sweet warble.<ref name=Collins/><ref name=Peterson>{{cite book |last1=Peterson |first1=Roger |author1-link=Roger Tory Peterson |last2=Mountfort |first2=Guy |author2-link=Guy Mountfort |last3=Hollom |first3=P.A.D. |author3-link=Philip Hollom |title=A Field Guide to the Birds of Britain and Europe |year=1954 |publisher=Collins |location=London }}</ref>
==Distribution and habitat== thumb|left|Juvenile, showing the browner, fluffy plumage. Hampshire, UK.
Dunnocks are native to large areas of Eurasia, inhabiting much of Europe and southwest Asia including Lebanon, northern Iran, and the Caucasus. It is the only accentor commonly found in temperate lowland areas; the others all inhabit upland areas, or (Siberian accentor) subarctic lowlands.<ref>{{cite web|url=http://www.garden-birds.co.uk/birds/dunnock.htm|access-date=6 May 2017|publisher=British Garden Birds|title=Dunnock}}</ref> Favoured habitats include woodlands, shrubs, gardens, and hedgerows where they typically feed on the ground, often seeking out detritivores as food.<ref name=brittannica>{{cite web|last=Montgomery|first=Sy|title=Dunnock|url=http://www.britannica.com/EBchecked/topic/173831/dunnock|publisher=Encyclopædia Britannica|access-date=23 September 2013}}</ref>
Dunnocks were intentionally introduced into New Zealand during the 19th century, and are now widely distributed around the country and some offshore islands.<ref>{{Cite web|title = Dunnock {{!}} New Zealand Birds Online|url = http://www.nzbirdsonline.org.nz/species/dunnock|website = www.nzbirdsonline.org.nz|access-date = 2015-10-04}}</ref><ref>{{Cite journal |last = Santos|first = Eduardo| year=2012 | title=Discovery of previously unknown historical records on the introduction of dunnocks (''Prunella modularis'') into Otago, New Zealand during the 19th century | journal=Notornis| volume=59 | issue=1 | pages=79–81 | url=https://www.birdsnz.org.nz/wp-content/uploads/2021/12/Santos_2012.pdf }}</ref>
==Territoriality== Dunnocks are territorial and may engage in conflict with other birds that encroach upon their nests.<ref name="brittannica"/> 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.<ref>{{Cite journal|title = Parental care and mating behaviour of polyandrous dunnocks ''Prunella modularis'' related to paternity by DNA fingerprinting| journal=Nature | year=1989 |pages=249–251 | volume=338 | issue=6212 | doi=10.1038/338249a0 | first1=T.| last1= Burke|first2 =N.B. | last2=Davies | first3=M.W.| last3= Bruford | first4=B.J. | last4=Hatchwell | bibcode=1989Natur.338..249B | s2cid=4333938 }}</ref><ref name=":0">{{Cite journal|title = Conflict and co-operation over sex: the consequences of social and genetic polyandry for reproductive success in dunnocks|journal = Journal of Animal Ecology| year=2015 | issn = 1365-2656|pages = 1509–1519|doi = 10.1111/1365-2656.12432|pmid = 26257043|first1 = Eduardo S. A.|last1 = Santos|first2 = Luana L. S.|last2 = Santos|first3 = Malgorzata|last3 = Lagisz|first4 = Shinichi|last4 = Nakagawa|volume=84|issue = 6|doi-access = free| bibcode=2015JAnEc..84.1509S }}</ref> Furthermore, members of a group are rarely related, and so competition can result.<ref name="food patchiness">{{cite journal | last1=Davies | first1=N. B.| last2=Hartley | first2=I.R. |title=Food patchiness, territory overlap and social systems: an experiment with dunnocks ''Prunella modularis'' |journal=Journal of Animal Ecology|year=1996|volume=65|issue=6|pages=837–846|doi=10.2307/5681|jstor=5681| bibcode=1996JAnEc..65..837D}}</ref>
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 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.<ref name="food patchiness"/>
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.<ref name="food patchiness"/><ref name="success">{{ cite journal | last1=Davies | first1=N.B. | last2=Houston | first2=A.I. | year=1986 | title=Reproductive success of dunnocks, ''Prunella modularis'', in a variable mating system | journal=Journal of Animal Ecology | volume=55 | issue=1 | pages=123–138 | doi=10.2307/4697 | jstor=4697 }}</ref>
==Breeding== ===Mating systems=== right|thumb|Male dunnock pecking cloaca of female before mating [[File:Dunnock (Prunella modularis occidentalis).jpg|thumb|On branch with hoar frost]]
The dunnock possesses variable mating systems. Females are often polyandrous, breeding with two or more males at once,<ref>{{Cite book|title = Dunnock behaviour and social evolution|last = Davies|first = Nicholas|publisher = Oxford University Press|year = 1992|isbn = 978-0-19-854674-0|location = Oxford}}</ref><ref>{{Cite journal|title = Breeding Biology and Variable Mating System of a Population of Introduced Dunnocks (''Prunella modularis'') in New Zealand|journal = PLOS ONE|date = 2013-07-09|pmc = 3706400|pmid = 23874945|article-number = e69329|volume = 8|issue = 7|doi = 10.1371/journal.pone.0069329|first1 = Eduardo S. A.|last1 = Santos|first2 = Shinichi|last2 = Nakagawa|bibcode = 2013PLoSO...869329S|doi-access = free}}</ref> which is quite rare among birds. This multiple mating system leads to the development of sperm competition amongst the male suitors. DNA fingerprinting has shown that chicks within a brood often have different fathers, depending on the success of the males at monopolising the female.<ref name=":0" /> Males try to ensure their paternity by pecking at the cloaca<ref>Attenborough, D. (1998). ''The Life of Birds''. BBC. p. 215. {{ISBN|0563-38792-0}}.</ref> of the female to stimulate ejection of rival males' sperm.<ref>{{cite journal |last1= Davies|first1= N. B.| year=1983 |title=Polyandry, cloaca-pecking and sperm competition in dunnocks |journal=Nature |volume= 302|issue= 5906|pages=334–336 |doi= 10.1038/302334a0|bibcode= 1983Natur.302..334D|s2cid= 4260839}}</ref> Dunnocks take just one-tenth of a second to copulate and can mate more than 100 times a day.<ref>{{cite book |last1=Birkhead |first1=Tim |title=Bird Sense |year=2012}}</ref> 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 is preferred. Sometimes, two or three adjacent female territories overlap one male territory, and so polygyny is favoured, 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.<ref name="food patchiness"/><ref name="success"/>
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).<ref name="food distribution">{{ cite journal | last1=Davies | first1=N.B. | last2=Lundberg | first2=A. | year=1984 | title=Food distribution and a variable mating system in the dunnock, ''Prunella modularis'' | journal=Journal of Animal Ecology |volume=53 | issue=3 | pages=895–912 | doi=10.2307/4666 | jstor=4666 | bibcode=1984JAnEc..53..895D | url=http://www.webpages.uidaho.edu/wlf314/Homeworks/Davies_Lundberg_1984.pdf }}</ref>
===Nest=== thumb|right|Dunnock nest and eggs [[File:Cuculus canorus canorus MHNT.ZOO.2010.11.149.30.jpg|thumb|Egg of ''Cuculus canorus canorus'' in a spawn of ''Prunella modularis ''- MHNT]]
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.<ref name="brittannica"/>
===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.<ref name="food distribution"/>
==References== {{Reflist}}
==External links== {{Commons}} * [https://www.xeno-canto.org/species/Prunella-modularis Xeno-canto: audio recordings of the dunnock] * [https://search.macaulaylibrary.org/catalog?taxonCode=dunnoc1 Dunnock videos, photos & sounds] on the Internet Bird Collection * [http://blascozumeta.com/wp-content/uploads/aragon-birds/passeriformes/324.dunnock-pmodularis.pdf Ageing and sexing (PDF; 2.0 MB) by Javier Blasco-Zumeta & Gerd-Michael Heinze] {{Webarchive|url=https://web.archive.org/web/20211212002915/http://blascozumeta.com/wp-content/uploads/aragon-birds/passeriformes/324.dunnock-pmodularis.pdf |date=12 December 2021 }} * [http://www.ornithos.de/Ornithos/Feather_Collection/Prunella_modularis/Prunella_modularis.htm Feathers of dunnock (Prunella modularis)] {{Webarchive|url=https://web.archive.org/web/20140808061516/http://www.ornithos.de/Ornithos/Feather_Collection/Prunella_modularis/Prunella_modularis.htm |date=8 August 2014 }}
{{Taxonbar|from=Q26698}} {{Authority control}}
Category:Animal taxa named by Carl Linnaeus Category:Birds described in 1758 Category:Birds of Europe Category:Birds of New Zealand Category:Prunellidae