Dunnock

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Dunnock
Dunnock (Prunella modularis) 3.jpg
Song recorded on Dartmoor in Devon, England
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Family: Prunellidae
Genus: Prunella
Species:
P. modularis
Binomial name
Prunella modularis
Dunnock ebird data map.png
Global range
  Year-Round Range
  Summer Range
  Winter Range
Synonyms
  • Motacilla modularisLinnaeus, 1758
  • Accentor modularis(Linnaeus, 1758)
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 successfully introduced into New Zealand. It is by far the most widespread member of the accentor family; most other accentors are limited to mountain habitats. Other common names of the dunnock include: hedge accentor, hedge sparrow, hedge warbler, and titling. [2]

Contents

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. [3] The specific epithet is from the Latin modularis "modulating" or "singing". [4] This species is now placed in the genus Prunella that was introduced by the French ornithologist Louis Pierre Vieillot in 1816. [5]

The name "dunnock" comes from the English dun (dingy brown, dark-coloured) and the diminutive ock, [6] and "accentor" is from post-classical Latin and means a person who sings with another. [7] The genus name Prunella is from the German Braunelle, "dunnock", a diminutive of braun, "brown". [8]

Eight subspecies are recognised: [9]

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. [10]

Description

A robin-sized bird, the dunnock typically measures 13.5–14 cm (5.3–5.5 in) 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. [11] Unlike any similar sized small brown bird, dunnocks exhibit frequent wing flicking, especially when engaged in territorial disputes or when competing for mating rights. [12] This gave rise to a common name of "shufflewing". [13]

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, but is shorter and weaker. [14]

Distribution and habitat

Dunnocks are native to large areas of Eurasia, inhabiting much of Europe including Lebanon, northern Iran, and the Caucasus. They are the only commonly found accentor in lowland areas; all the others inhabit upland areas. [15] Dunnocks were successfully introduced into New Zealand during the 19th century, and are now widely distributed around the country and some offshore islands. [16] [17] Favoured habitats include woodlands, shrubs, gardens, and hedgerows where they typically feed on the ground, often seeking out detritivores as food. [18]

Territoriality

Dunnocks are territorial and may engage in conflict with other birds that encroach upon their nests. [18] 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. [19] [20] Furthermore, members of a group are rarely related, and so competition can result. [21]

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. [21]

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. [21] [22]

Breeding

Mating systems

Male dunnock pecking cloaca of female before mating Dunnocks cloaca pecking.jpg
Male dunnock pecking cloaca of female before mating
on branch with hoar frost Dunnock (Prunella modularis occidentalis).jpg
on branch with hoar frost

The dunnock possesses variable mating systems. Females are often polyandrous, breeding with two or more males at once, [23] [24] 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. [20] Males try to ensure their paternity by pecking at the cloaca [25] of the female to stimulate ejection of rival males' sperm. [26] Dunnocks take just one-tenth of a second to copulate and can mate more than 100 times a day. [27] 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. [21] [22]

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). [28]

Nest

Dunnock nest and eggs Dunnock (Hedge Sparrow) Nest 11.04.11.jpg
Dunnock nest and eggs
Egg of Cuculus canorus canorus in a spawn of Prunella modularis - MHNT Cuculus canorus canorus MHNT.ZOO.2010.11.149.30.jpg
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. [18]

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. [28]

Related Research Articles

<span class="mw-page-title-main">Accentor</span> Genus of birds

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 the mountainous regions of Europe and Asia; these two also occur in lowland areas, as does the Siberian accentor in the far north of Siberia. These birds are not strongly migratory, but they will leave the coldest parts of their range in winter and make altitudinal movements.

<span class="mw-page-title-main">Behavioral ecology</span> Study of the evolutionary basis for animal behavior due to ecological pressures

Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for animal behavior due to ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined four questions to address when studying animal behaviors: What are the proximate causes, ontogeny, survival value, and phylogeny of a behavior?

<span class="mw-page-title-main">Alpine accentor</span> Species of bird

The alpine accentor is a small passerine bird in the family Prunellidae, which is native to Eurasia and North Africa.

<i>Ursus</i> (mammal) Genus of bears

Ursus is a genus in the family Ursidae (bears) that includes the widely distributed brown bear, the polar bear, the American black bear, and the Asian black bear. The name is derived from the Latin ursus, meaning bear.

<span class="mw-page-title-main">House mouse</span> Species of mammal

The house mouse is a small mammal of the order Rodentia, characteristically having a pointed snout, large rounded ears, and a long and almost hairless tail. It is one of the most abundant species of the genus Mus. Although a wild animal, the house mouse has benefited significantly from associating with human habitation to the point that truly wild populations are significantly less common than the semi-tame populations near human activity.

Polygynandry is a mating system in which both males and females have multiple mating partners during a breeding season. In sexually reproducing diploid animals, different mating strategies are employed by males and females, because the cost of gamete production is lower for males than it is for females. The different mating tactics employed by males and females are thought to be the outcome of stochastic reproductive conflicts both ecologically and socially.

<span class="mw-page-title-main">Sperm competition</span> Reproductive process

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 variety of mates increases a female's chance to produce more viable offspring. However, multiple mates for a female means each individual male has decreased chances of producing offspring. Sperm competition is an evolutionary pressure on males, and has led to the development of adaptations to increase male's chance of reproductive success. Sperm competition results in a sexual conflict between males and females. Males have evolved several defensive tactics including: mate-guarding, mating plugs, and releasing toxic seminal substances to reduce female re-mating tendencies to cope with sperm competition. Offensive tactics of sperm competition involve direct interference by one male on the reproductive success of another male, for instance by physically removing another male's sperm prior to mating with a female. For an example, see Gryllus bimaculatus.

<span class="mw-page-title-main">Parental investment</span> Parental expenditure (e.g. time, energy, resources) that benefits offspring

Parental investment, in evolutionary biology and evolutionary psychology, is any parental expenditure that benefits offspring. Parental investment may be performed by both males and females, females alone or males alone. Care can be provided at any stage of the offspring's life, from pre-natal to post-natal.

<span class="mw-page-title-main">Reproductive synchrony</span>

Reproductive synchrony is a term used in evolutionary biology and behavioral ecology. Reproductive synchrony—sometimes termed "ovulatory synchrony"—may manifest itself as "breeding seasonality". Where females undergo regular menstruation, "menstrual synchrony" is another possible term.

<span class="mw-page-title-main">Parental care</span>

Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal kingdom. There is great variation in different animal groups in terms of how parents care for offspring, and the amount of resources invested by parents. For example, there may be considerable variation in the amount of care invested by each sex, where females may invest more in some species, males invest more in others, or investment may be shared equally. Numerous hypotheses have been proposed to describe this variation and patterns in parental care that exist between the sexes, as well as among species.

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Bateman's principle, in evolutionary biology, is that in most species, variability in reproductive success is greater in males than in females. It was first proposed by Angus John Bateman (1919–1996), an English geneticist. Bateman suggested that, since males are capable of producing millions of sperm cells with little effort, while females invest much higher levels of energy in order to nurture a relatively small number of eggs, the female plays a significantly larger role in their offspring's reproductive success. Bateman's paradigm thus views females as the limiting factor of parental investment, over which males will compete in order to copulate successfully.

<span class="mw-page-title-main">Nicholas Barry Davies</span>

Nicholas Barry Davies is a British field naturalist and zoologist, and Emeritus Professor of Behavioural Ecology at the University of Cambridge, where he is also a Emeritus Fellow of Pembroke College.

Polygyny is a mating system in which one male lives and mates with multiple females but each female only mates with a single male. Systems where several females mate with several males are defined either as promiscuity or polygynandry. Lek mating is frequently regarded as a form of polygyny, because one male mates with many females, but lek-based mating systems differ in that the male has no attachment to the females with whom he mates, and that mating females lack attachment to one another.

<span class="mw-page-title-main">Sexual selection in birds</span>

Sexual selection in birds concerns how birds have evolved a variety of mating behaviors, with the peacock tail being perhaps the most famous example of sexual selection and the Fisherian runaway. Commonly occurring sexual dimorphisms such as size and color differences are energetically costly attributes that signal competitive breeding situations. Many types of avian sexual selection have been identified; intersexual selection, also known as female choice; and intrasexual competition, where individuals of the more abundant sex compete with each other for the privilege to mate. Sexually selected traits often evolve to become more pronounced in competitive breeding situations until the trait begins to limit the individual's fitness. Conflicts between an individual fitness and signaling adaptations ensure that sexually selected ornaments such as plumage coloration and courtship behavior are "honest" traits. Signals must be costly to ensure that only good-quality individuals can present these exaggerated sexual ornaments and behaviors.

<span class="mw-page-title-main">Sexual selection in amphibians</span> Choice of and competition for mates

Sexual selection in amphibians involves sexual selection processes in amphibians, including frogs, salamanders and newts. Prolonged breeders, the majority of frog species, have breeding seasons at regular intervals where male-male competition occurs with males arriving at the waters edge first in large number and producing a wide range of vocalizations, with variations in depth of calls the speed of calls and other complex behaviours to attract mates. The fittest males will have the deepest croaks and the best territories, with females making their mate choices at least partly based on the males depth of croaking. This has led to sexual dimorphism, with females being larger than males in 90% of species, males in 10% and males fighting for groups of females.

Polyandry in fishes is a mating system where females mate with multiple males within one mating season. This type of mating exists in a variety of animal species. Polyandry has been found in both oviparous and viviparous bony fishes and sharks. General examples of polyandry occur in fish species, such as green swordtails and Trinidadian guppies. Specific types of polyandry have also been classified, such as classical polyandry in pipefish cooperative polyandry in cichlids and convenience polyandry in sharks.

<span class="mw-page-title-main">Polyandry in animals</span> Class of mating system in non-human species

In behavioral ecology, polyandry is a class of mating system where one female mates with several males in a breeding season. Polyandry is often compared to the polygyny system based on the cost and benefits incurred by members of each sex. Polygyny is where one male mates with several females in a breeding season . A common example of polyandrous mating can be found in the field cricket of the invertebrate order Orthoptera. Polyandrous behavior is also prominent in many other insect species, including the red flour beetle and the species of spider Stegodyphus lineatus. Polyandry also occurs in some primates such as marmosets, mammal groups, the marsupial genus' Antechinus and bandicoots, around 1% of all bird species, such as jacanas and dunnocks, insects such as honeybees, and fish such as pipefish.

<span class="mw-page-title-main">Parental care in birds</span>

Parental care refers to the level of investment provided by the mother and the father to ensure development and survival of their offspring. In most birds, parents invest profoundly in their offspring as a mutual effort, making a majority of them socially monogamous for the duration of the breeding season. This happens regardless of whether there is a paternal uncertainty.

References

  1. ↑ BirdLife International (2018). "Prunella modularis". IUCN Red List of Threatened Species . 2018: e.T22718651A132118966. doi: 10.2305/IUCN.UK.2018-2.RLTS.T22718651A132118966.en . Retrieved 13 November 2021.
  2. ↑ "Titling". 3 November 2023.
  3. ↑ Linnaeus, Carl (1758). Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (in Latin). Vol. 1 (10th ed.). Holmiae:Laurentii Salvii. p. 184.
  4. ↑ Jobling, J.A. (2019). del Hoyo, J.; Elliott, A.; Sargatal, J.; Christie, D.A.; de Juana, E. (eds.). "Key to Scientific Names in Ornithology". Handbook of the Birds of the World Alive. Lynx Edicions. Retrieved 10 March 2019.
  5. ↑ Vieillot, Louis Pierre (1816). Analyse d'une Nouvelle Ornithologie Élémentaire (in French). Paris: Deterville/self. p. 43.
  6. ↑ "Dunnock" . Oxford English Dictionary (Online ed.). Oxford University Press.(Subscription or participating institution membership required.)
  7. ↑ "Accentor" . Oxford English Dictionary (Online ed.). Oxford University Press.(Subscription or participating institution membership required.)
  8. ↑ Jobling, James A. (2010). The Helm Dictionary of Scientific Bird Names. London, United Kingdom: Christopher Helm. p. 318. ISBN   978-1-4081-2501-4.
  9. ↑ Gill, Frank; Donsker, David, eds. (2019). "Waxbills, parrotfinches, munias, whydahs, Olive Warbler, accentors, pipits". World Bird List Version 9.1. International Ornithologists' Union. Retrieved 11 March 2019.
  10. ↑ Pavia, Marco; Drovetski, Sergei V.; Boano, Giovanni; Conway, Kevin W.; Pellegrino, Irene; Voelker, Gary (15 June 2021). "Elevation of two subspecies of Dunnock Prunella modularis to species rank". Bulletin of the British Ornithologists' Club. 141 (2): 199–210. doi: 10.25226/bboc.v141i2.2021.a10 .
  11. ↑ Heather, Barrie; Rogertson, Hugh (2005). The Field Guide to the Birds of New Zealand (Revised ed.). Viking Press.
  12. ↑ "Dunnock". RSPB. Retrieved 6 May 2017.
  13. ↑ Webster's Revised Unabridged Dictionary. G. & C. Merriam. 1913. Retrieved 2 February 2023.
  14. ↑ Peterson, Roger; Mountfort, Guy; Hollom, P.A.D. (1954). A Field Guide to the Birds of Britain and Europe. London: Collins.
  15. ↑ "Dunnock". British Garden Birds. Retrieved 6 May 2017.
  16. ↑ "Dunnock | New Zealand Birds Online". www.nzbirdsonline.org.nz. Retrieved 4 October 2015.
  17. ↑ Santos, Eduardo (2012). "Discovery of previously unknown historical records on the introduction of dunnocks (Prunella modularis) into Otago, New Zealand during the 19th century" (PDF). Notornis. 59 (1): 79–81.
  18. 1 2 3 Montgomery, Sy. "Dunnock". Encyclopædia Britannica. Retrieved 23 September 2013.
  19. ↑ Burke, T.; Davies, N.B.; Bruford, M.W.; Hatchwell, B.J. (1989). "Parental care and mating behaviour of polyandrous dunnocks Prunella modularis related to paternity by DNA fingerprinting". Nature. 338 (6212): 249–251. Bibcode:1989Natur.338..249B. doi:10.1038/338249a0. S2CID   4333938.
  20. 1 2 Santos, Eduardo S. A.; Santos, Luana L. S.; Lagisz, Malgorzata; Nakagawa, Shinichi (2015). "Conflict and co-operation over sex: the consequences of social and genetic polyandry for reproductive success in dunnocks". Journal of Animal Ecology. 84 (6): 1509–1519. Bibcode:2015JAnEc..84.1509S. doi: 10.1111/1365-2656.12432 . ISSN   1365-2656. PMID   26257043.
  21. 1 2 3 4 Davies, N. B.; Hartley, I.R. (1996). "Food patchiness, territory overlap and social systems: an experiment with dunnocks Prunella modularis". Journal of Animal Ecology. 65 (6): 837–846. Bibcode:1996JAnEc..65..837D. doi:10.2307/5681. JSTOR   5681.
  22. 1 2 Davies, N.B.; Houston, A.I. (1986). "Reproductive success of dunnocks, Prunella modularis, in a variable mating system". Journal of Animal Ecology. 55 (1): 123–138. doi:10.2307/4697. JSTOR   4697.
  23. ↑ Davies, Nicholas (1992). Dunnock behaviour and social evolution. Oxford: Oxford University Press. ISBN   978-0198546740.
  24. ↑ Santos, Eduardo S. A.; Nakagawa, Shinichi (9 July 2013). "Breeding Biology and Variable Mating System of a Population of Introduced Dunnocks (Prunella modularis) in New Zealand". PLOS ONE. 8 (7): e69329. Bibcode:2013PLoSO...869329S. doi: 10.1371/journal.pone.0069329 . PMC   3706400 . PMID   23874945.
  25. ↑ Attenborough, D. 1998. p.215. The Life of Birds BBC ISBN   0563-38792-0
  26. ↑ Davies, N. B. (1983). "Polyandry, cloaca-pecking and sperm competition in dunnocks". Nature. 302 (5906): 334–336. Bibcode:1983Natur.302..334D. doi:10.1038/302334a0. S2CID   4260839.
  27. ↑ Birkhead, Tim (2012). Bird Sense.
  28. 1 2 Davies, N.B.; Lundberg, A. (1984). "Food distribution and a variable mating system in the dunnock, Prunella modularis" (PDF). Journal of Animal Ecology. 53 (3): 895–912. Bibcode:1984JAnEc..53..895D. doi:10.2307/4666. JSTOR   4666.
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