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| name = Tit
| name = Tit
| image = Lophophanes cristatus -Aviemore, Scotland-8 (2).jpg
| image = Lophophanes cristatus -Aviemore, Scotland-8 (2).jpg
| image_caption = [[European crested tit]] in [[Scotland]]
| image_caption = [[Crested tit]] in [[Scotland]]
| taxon = Paridae
| taxon = Paridae
| authority = [[Nicholas Aylward Vigors|Vigors]], 1825
| authority = [[Nicholas Aylward Vigors|Vigors]], 1825
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The '''tits''', [[chickadees]], and [[Titmouse|titmice]] constitute the '''Paridae''', a large family of small [[passerine]] [[bird]]s which occur mainly in the [[Northern Hemisphere]] and [[Africa]]. Most were formerly classified in the genus ''Parus''.
The '''tits''', [[chickadees]], and [[Titmouse|titmice]] constitute the '''Paridae''', a large family of small [[passerine]] [[bird]]s which occur mainly in the [[Northern Hemisphere]] and [[Africa]]. Most were formerly classified in the genus ''Parus''.


Members of this family are commonly referred to as "tits" throughout much of the English speaking world, but North American species are called either "chickadees" ([[onomatopoeic]], derived from their distinctive "chick-a dee dee dee" [[alarm call]])<ref name = "Merriam"/> or "titmice". The name titmouse is recorded from the 14th century, composed of the [[Old English language|Old English]] name for the bird, ''mase'' ([[Proto-Germanic]] ''*maison'', [[Dutch language|Dutch]] ''mezen'', [[German language|German]] ''Meise''), and tit, denoting something small. The former spelling, "titmose", was influenced by ''[[mouse]]'' in the 16th century.<ref>{{cite web|url=http://etymonline.com/index.php?allowed_in_frame=0&search=titmouse&searchmode=term |website=Online Etymology Dictionary |title=titmouse (n.) |access-date=2015-04-28 |author=Douglas Harper (© 2001-2015)}}</ref> Emigrants to [[New Zealand]] presumably identified some of the superficially similar birds of the genus ''[[Petroica]]'' of the family Petroicidae, the Australian robins, as members of the tit family, giving them the title [[tomtit]], although, in fact, they are not related.
Eurasian and African members of this family are referred to as "tits", while North American species are called either "chickadees" ([[onomatopoeic]], derived from their distinctive "chick-a dee dee dee" [[alarm call]])<ref name = "Merriam"/> or "titmice". The name titmouse is recorded from the 14th century, composed of the [[Old English language|Old English]] name for the bird, ''mase'' ([[Proto-Germanic]] ''*maison'', [[Dutch language|Dutch]] ''mees'', [[German language|German]] ''Meise''), and tit, denoting something small. The former spelling, "titmose", was influenced by ''[[mouse]]'' in the 16th century.<ref>{{cite dictionary |url=http://etymonline.com/index.php?allowed_in_frame=0&search=titmouse&searchmode=term |dictionary=Online Etymology Dictionary |title=titmouse (n.) |access-date=2015-04-28 |author=Douglas Harper}}</ref> Emigrants to [[New Zealand]] presumably identified some of the superficially similar birds of the genus ''[[Petroica]]'' of the family Petroicidae, the Australian robins, as members of the tit family, giving them the title [[tomtit]], although, in fact, they are not related.


These birds are mainly small, stocky, woodland [[species]] with short, stout [[Beak|bills]]. Some have crests. They range in length from {{convert|10|to|22|cm|in|abbr=on}}. They are adaptable birds, with a mixed diet including seeds and insects.<ref name=EoB>{{cite book |editor=Forshaw, Joseph|author= Perrins, C.|year=1991|title=Encyclopaedia of Animals: Birds|publisher= Merehurst Press|location=London|pages= 202–203|isbn= 1-85391-186-0}}</ref> Many species live around human habitation and come readily to [[bird feeder]]s for nuts or seed, and learn to take other foods.
These birds are mainly small, stocky, woodland [[species]] with short, stout [[Beak|bills]]. Some have crests. They range in length from {{convert|10|to|22|cm|in|abbr=on}}. They are adaptable birds, with a mixed diet including seeds and insects.<ref name=EoB>{{cite book |editor=Forshaw, Joseph|author= Perrins, C.|year=1991|title=Encyclopaedia of Animals: Birds|publisher= Merehurst Press|location=London|pages= 202–203|isbn= 1-85391-186-0}}</ref> Many species live around human habitation and come readily to [[bird feeder]]s for nuts or seed, and learn to take other foods.


==Description==
==Description==
With the exception of the three monotypic genera ''Sylviparus'', ''Melanochlora'', and ''Pseudopodoces'', the tits are extremely similar in appearance, and have been described as "one of the most conservative avian families in terms of general morphology".<ref>Gosler & Clement (2007) P.669</ref> The typical body length of adult members of the family is between {{convert|10|and|16|cm|in|abbr=on}} in length; when the monotypic genera are added, this range is from {{convert|9|to|21|cm|in|abbr=on}}. In weight, the family ranges from {{convert|5|to|49|g|oz|abbr=on}}; this contracts to {{convert|7|to|29|g|oz|abbr=on}} when the three atypical genera are removed. The majority of the variation within the family is in [[plumage]], and particularly colour.<ref name ="HBW"/>
With the exception of the three monotypic genera ''Sylviparus'', ''Melanochlora'', and ''Pseudopodoces'', the tits are extremely similar in appearance, and have been described as "one of the most conservative avian families in terms of general morphology".<ref>Gosler & Clement (2007) P.669</ref> The typical body length of adult members of the family is between {{convert|10|and|16|cm|in|abbr=on}} in length; when the monotypic genera are added, this range is from {{convert|9|to|21|cm|in|abbr=on}}. In weight, the family ranges from {{convert|5|to|49|g|oz|abbr=on}}; this contracts to {{convert|7|to|29|g|oz|abbr=on}} when the three atypical genera are removed. The majority of the variation within the family is in [[plumage]], and particularly colour.<ref name ="HBW"/>


The [[beak|bills]] of the tits are generally short, varying between stout and fine, depending on diet. The more insectivorous species have finer bills, whereas those that consume more seeds have stouter bills. It is said that tits are evolving longer beaks to reach into bird feeders.<ref>https://news.nationalgeographic.com/2017/10/great-tits-birds-feeders-evolution-beaks/</ref> The most aberrant bill of the family is possessed by [[Hume's ground tit]] of Tibet and the Himalayas, which is long and decurved.<ref name ="HBW"/>
The [[beak|bills]] of the tits are generally short, varying between stout and fine, depending on diet. The more insectivorous species have finer bills, whereas those that consume more seeds have stouter bills. It is said that tits are evolving longer beaks to reach into bird feeders.<ref>{{cite web | url=https://news.nationalgeographic.com/2017/10/great-tits-birds-feeders-evolution-beaks/ | archive-url=https://web.archive.org/web/20171020050410/https://news.nationalgeographic.com/2017/10/great-tits-birds-feeders-evolution-beaks/ | archive-date=October 20, 2017 | title=Great Tits May be Evolving Bigger Beaks. Here's Why | date=19 October 2017 }}</ref> The most aberrant bill of the family is possessed by [[Hume's ground tit]] of Tibet and the Himalayas, which is long and decurved.<ref name ="HBW"/>


==Distribution and habitat==
==Distribution and habitat==
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==Behaviour==
==Behaviour==
Tits are active, noisy, and social birds. They are [[territory (animal)|territorial]] during the breeding season and often join [[mixed-species feeding flock]]s during the nonbreeding season. The tits are highly adaptable, and after the [[Corvidae|corvids]] (crows and jays) and [[parrot]]s, amongst the most intelligent of all birds.<ref name ="HBW">{{Cite book | first1 = Andrew | last1 = Gosler | first2 = Peter | last2 = Clement | editor-first = Josep | editor-last = del Hoyo | editor2-first = Andrew | editor2-last = Elliott | editor3-first = David | editor3-last = Christie | contribution = Family Paridae (Tits and Chickadees) | title = Handbook of the Birds of the World. Volume 12: Picathartes to Tits and Chickadees | year = 2007 | pages = 662–709| place = Barcelona | publisher = Lynx Edicions | isbn = 978-84-96553-42-2 }}</ref>
Tits are active, noisy, and social birds. They are [[territory (animal)|territorial]] during the breeding season and often join [[mixed-species feeding flock]]s during the nonbreeding season. The tits are highly adaptable, and after the [[Corvidae|corvids]] (crows and jays) and [[parrot]]s, amongst the most intelligent of all birds.<ref name ="HBW">{{Cite book | first1 = Andrew | last1 = Gosler | first2 = Peter | last2 = Clement | editor-first = Josep | editor-last = del Hoyo | editor2-first = Andrew | editor2-last = Elliott | editor3-first = David | editor3-last = Christie | contribution = Family Paridae (Tits and Chickadees) | title = Handbook of the Birds of the World. Volume 12: Picathartes to Tits and Chickadees | year = 2007 | pages = 662–709| place = Barcelona | publisher = Lynx Edicions | isbn = 978-84-96553-42-2 }}</ref> Tits recognize the difference between species that are dangerous or harmless to them, by this they can protect each other or their families. These birds do this by mobbing or escaping, however they also avoid the nest when the predators are present in order to avoid their families to be seen.


===Fission–fusion society===
===Fission–fusion society===
[[Fission–fusion society]] has been documented in a number of avian taxa including this one.<ref>{{cite journal|last1=Kendra|first1=Sewall|title=Social Complexity as a Driver of Communication and Cognition.|journal=Integrative and Comparative Biology|date=2015|volume=55|issue=3|pages=384–395|doi=10.1093/icb/icv064|pmid=26078368|doi-access=free}}</ref><ref>{{cite journal|last1=Alpin|first1=L. M.|last2=Farine|first2=D. R.|last3=Morand-Ferron|first3=J.|last4=Cole|first4=E. F.|last5=Cockburn|first5=A.|last6=Sheldon|first6=B. C.|title=Individual personalities predict social behaviour in wild networks of great tits (Parus major)|journal=Ecology Letters|date=2013|volume=16|issue=11|pages=1365–1372|doi=10.1111/ele.12181|pmid=24047530|hdl=1885/66143|hdl-access=free}}{{dead link|date=February 2018}}</ref> In brief, that means flocks can split into smaller groups or individuals, and subsequently reunite.
[[Fission–fusion society]] has been documented in a number of avian taxa including this one.<ref>{{cite journal|last1=Kendra|first1=Sewall|title=Social Complexity as a Driver of Communication and Cognition.|journal=Integrative and Comparative Biology|date=2015|volume=55|issue=3|pages=384–395|doi=10.1093/icb/icv064|pmid=26078368|doi-access=free}}</ref><ref>{{cite journal|last1=Alpin|first1=L. M.|last2=Farine|first2=D. R.|last3=Morand-Ferron|first3=J.|last4=Cole|first4=E. F.|last5=Cockburn|first5=A.|last6=Sheldon|first6=B. C.|title=Individual personalities predict social behaviour in wild networks of great tits (Parus major)|journal=Ecology Letters|date=2013|volume=16|issue=11|pages=1365–1372|doi=10.1111/ele.12181|pmid=24047530|hdl=1885/66143|s2cid=3400989 |hdl-access=free}}{{dead link|date=February 2018}}</ref><ref>{{Cite web |last=admin |date=2022-08-15 |title=Tufted Titmouse - Best Facts 2022 |url=https://birdsatfirstsight.com/the-tufted-titmouse/ |access-date=2022-09-30 |website=Birds At First Sight |language=en-US}}</ref> In brief, that means flocks can split into smaller groups or individuals, and subsequently reunite.


===Vocalisations===
===Vocalisations===
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[[File:Marsh tit, Leighton Moss, December 2008.jpg|thumb|Hold-hammering is a common way for the family to deal with food items.]]
[[File:Marsh tit, Leighton Moss, December 2008.jpg|thumb|Hold-hammering is a common way for the family to deal with food items.]]
[[File:CyanistesCaeruleusWithForage.jpg|thumb|[[Blue tit]] with prey item]]
[[File:CyanistesCaeruleusWithForage.jpg|thumb|[[Blue tit]] with prey item]]
The tits are generalist [[insectivores]] that consume a wide range of small insects and other invertebrates, particularly small defoliating caterpillars. They also consume [[seed]]s and nuts, particularly in the winter. One characteristic method of foraging in the family is hanging, where they inspect a branch or twig and leaves from all angles while hanging upside down to feed.<ref name ="HBW"/> In areas where numerous species of tit coexist, different species forage in different parts of the tree, their niche determined in no small way by their morphology; larger species forage on the ground, medium-sized species foraging on larger branches, and the smallest species on the ends of branches. Having obtained larger prey items or seeds, tits engage in hold-hammering, where they hold the item between the feet and hammer it with the bill until it opens. In this fashion, they can even open [[hazelnut]]s in around 20 minutes. A number of genera engage in food caching, hoarding supplies of food during the winter.<ref>{{cite journal | doi = 10.2307/1939354 | last1 = Jokinen | first1 = S | last2 = Suhonen | first2 = J | year = 1995 | title = Food Caching By Willow and Crested Tits: A Test of Scatterhoarding Models | jstor = 1939354| journal = Ecology | volume = 76 | issue = 3| pages = 892–898 }}</ref>
The tits are generalist [[insectivores]] that consume a wide range of small insects and other invertebrates, particularly small defoliating caterpillars. They also consume [[seed]]s and nuts, particularly in the winter. One characteristic method of foraging in the family is hanging, where they inspect a branch or twig and leaves from all angles while hanging upside down to feed.<ref name ="HBW"/> In areas where numerous species of tit coexist, different species forage in different parts of the tree, their niche determined in no small way by their morphology; larger species forage on the ground, medium-sized species foraging on larger branches, and the smallest species on the ends of branches. Having obtained larger prey items or seeds, tits engage in hold-hammering, where they hold the item between their feet and hammer it with their bill until it opens. In this fashion, they can even open [[hazelnut]]s in around 20 minutes. A number of genera engage in food caching, hoarding supplies of food during the winter.<ref>{{cite journal | doi = 10.2307/1939354 | last1 = Jokinen | first1 = S | last2 = Suhonen | first2 = J | year = 1995 | title = Food Caching By Willow and Crested Tits: A Test of Scatterhoarding Models | jstor = 1939354| journal = Ecology | volume = 76 | issue = 3| pages = 892–898 }}</ref>


===Breeding===
===Breeding===
Tits are cavity-nesting birds, typically using trees, although ''Pseudopodoces''<ref>[http://si-pddr.si.edu/jspui/bitstream/10088/1758/3/VZ_321.Pseudopodoces.pdf "Pseudopodoces humilis, a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones"]</ref> builds a nest on the ground. Most tree-nesting tits excavate their nests,<ref>Mönkkönen, Mikko and Orell, Markku; "Clutch Size and Cavity Excavation in Parids (Paridae): The Limited Breeding Opportunities Hypothesis Tested" in ''[[The American Naturalist]]'', Vol. 149, No. 6 (June 1997), pp. 1164–1174</ref> and [[avian clutch size|clutch size]]s are generally large for altricial birds, ranging from usually two eggs in the [[rufous-vented tit]] of the Himalayas to as many as 10 to 14 in the blue tit of Europe.<ref>[http://www.plosbiology.org/article/info%3Adoi%2F10.1371%2Fjournal.pbio.0060303#pbio-0060303-st001 "List of Species and Data Sources Used for Geographic Distributions and Data on Clutch Sizes and Intrinsic Variables"]</ref><ref name="population">Newton, Ian; ''Population Limitation in Birds''; p. 25. {{ISBN|9780125173667}}</ref> In favourable conditions, this species had laid as many as 19 eggs, which is the largest clutch of any altricial bird.<ref name=EoB/> Most tits are multibrooded, a necessary strategy to cope with either the harsh winters in which they reside in the Holarctic or the extremely erratic conditions of tropical Africa,<ref name="population"/> where typically a single pair cannot find enough food to rear even one nestling and in drought years breeding is likely to be futile.
Tits are cavity-nesting birds, typically using trees, although ''Pseudopodoces''<ref>[http://si-pddr.si.edu/jspui/bitstream/10088/1758/3/VZ_321.Pseudopodoces.pdf "Pseudopodoces humilis, a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones"]{{Dead link|date=September 2023 |bot=InternetArchiveBot |fix-attempted=yes }}</ref> builds a nest on the ground. Most tree-nesting tits excavate their nests,<ref>Mönkkönen, Mikko and Orell, Markku; "Clutch Size and Cavity Excavation in Parids (Paridae): The Limited Breeding Opportunities Hypothesis Tested" in ''[[The American Naturalist]]'', Vol. 149, No. 6 (June 1997), pp. 1164–1174</ref> and [[avian clutch size|clutch size]]s are generally large for altricial birds, ranging from usually two eggs in the [[rufous-vented tit]] of the Himalayas to as many as 10 to 14 in the blue tit of Europe.<ref>[http://www.plosbiology.org/article/info%3Adoi%2F10.1371%2Fjournal.pbio.0060303#pbio-0060303-st001 "List of Species and Data Sources Used for Geographic Distributions and Data on Clutch Sizes and Intrinsic Variables"]</ref><ref name="population">Newton, Ian; ''Population Limitation in Birds''; p. 25. {{ISBN|978-0-12-517366-7}}</ref> In favourable conditions, this species had laid as many as 19 eggs, which is the largest clutch of any altricial bird.<ref name=EoB/> Most tits are multibrooded, a necessary strategy to cope with either the harsh winters in which they reside in the Holarctic or the extremely erratic conditions of tropical Africa,<ref name="population"/> where typically a single pair cannot find enough food to rear even one nestling and in drought years breeding is likely to be futile.


Many African tit species, along with ''Pseudopodoces'', are [[cooperative breeders]],<ref>{{cite journal|doi=10.1111/j.1474-919X.2011.01115.x | volume=153 | issue=2 | title=Geographical variation in patterns of parentage and relatedness in the co-operatively breeding Ground Tit Parus humilis | year=2011 | journal=Ibis | pages=373–383 | last1 = Johannessen | first1 = Lars E.}}</ref> and even pair-breeding parids are often highly social and maintain stable flocks throughout the nonbreeding season.<ref>Stacey, Peter B. and Ligon, J. David; "The Benefits-of-Philopatry Hypothesis for the Evolution of Cooperative Breeding: Variation in Territory Quality and Group Size Effects" in ''[[The American Naturalist]]'', Vol. 137, No. 6 (Jun., 1991), pp. 831–846</ref>
Many African tit species, along with ''Pseudopodoces'', are [[cooperative breeders]],<ref>{{cite journal|doi=10.1111/j.1474-919X.2011.01115.x | volume=153 | issue=2 | title=Geographical variation in patterns of parentage and relatedness in the co-operatively breeding Ground Tit Parus humilis | year=2011 | journal=Ibis | pages=373–383 | last1 = Johannessen | first1 = Lars E.}}</ref> and even pair-breeding parids are often highly social and maintain stable flocks throughout the nonbreeding season.<ref>Stacey, Peter B. and Ligon, J. David; "The Benefits-of-Philopatry Hypothesis for the Evolution of Cooperative Breeding: Variation in Territory Quality and Group Size Effects" in ''[[The American Naturalist]]'', Vol. 137, No. 6 (Jun., 1991), pp. 831–846</ref>
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Tits also have a variety of methods for attracting mates, primarily through their intricate, bouncing mating dance. Only the blue tit is typically polygynous; all other species are generally monogamous.<ref>{{cite journal | pmc = 1688915 | pages=445–450 | volume=265 | issue=1395 | journal=Proc Biol Sci | title=Ultraviolet sexual dimorphism and assortative mating in blue tits | doi=10.1098/rspb.1998.0315| year=1998 | last1=Andersson | first1=S | last2=Rnborg | first2=J | last3=Andersson | first3=M }}</ref> Courtship feeding is typical of pair-breeding tits to deal with the cost of rearing their large broods.
Tits also have a variety of methods for attracting mates, primarily through their intricate, bouncing mating dance. Only the blue tit is typically polygynous; all other species are generally monogamous.<ref>{{cite journal | pmc = 1688915 | pages=445–450 | volume=265 | issue=1395 | journal=Proc Biol Sci | title=Ultraviolet sexual dimorphism and assortative mating in blue tits | doi=10.1098/rspb.1998.0315| year=1998 | last1=Andersson | first1=S | last2=Rnborg | first2=J | last3=Andersson | first3=M }}</ref> Courtship feeding is typical of pair-breeding tits to deal with the cost of rearing their large broods.


== Systematics ==
==Systematics==
[[Image:Parus palustris02.jpg|thumb|The [[marsh tit]] was once placed in the genus ''Parus'', but has now been moved to the genus ''Poecile''.]]
[[Image:Parus palustris02.jpg|thumb|The [[marsh tit]] was once placed in the genus ''Parus'', but has now been moved to the genus ''Poecile''.]]
Recently{{when|date=May 2018}}, the large ''Parus'' group has been gradually split into several genera (as indicated below), initially by [[North America]]n ornithological authorities and later elsewhere. Whereas in the mid-1990s, only ''Pseudopodoces'', ''Baeolophus'', ''Melanochlora'', and ''Sylviparus'' were considered well-supported by the available data as distinct from ''Parus''.<ref>Harrap, Simon & Quinn, David (1996): ''Tits, Nuthatches & Treecreepers''. [[Helm Identification Guides|Christopher Helm]], London. {{ISBN|0-7136-3964-4}}</ref> Today, this arrangement is considered [[paraphyletic]] as indicated by [[mtDNA]] [[cytochrome b|cytochrome ''b'']] [[DNA sequence|sequence]] analysis, and ''Parus'' is best restricted to the ''Parus major''—''Parus fasciiventer'' [[clade]], and even the latter species' closest relatives might be considered a distinct genus.<ref name ="Gill">{{cite journal | last1 = Gill | first1 = Frank B. | last2 = Slikas | first2 = Beth | last3 = Sheldon | first3 = Frederick H. | year = 2005 | title = Phylogeny of titmice (Paridae): II. Species relationships based on sequences of the mitochondrial cytochrome-b gene | journal = [[Auk (journal)|Auk]] | volume = 122 | issue = 1| pages = 121–143 | doi = 10.1642/0004-8038(2005)122[0121:POTPIS]2.0.CO;2 }}</ref>
Recently, the large ''Parus'' group has been gradually split into several genera (as indicated below), initially by [[North America]]n ornithological authorities and later elsewhere. Whereas in the mid-1990s, only ''Pseudopodoces'', ''Baeolophus'', ''Melanochlora'', and ''Sylviparus'' were considered well-supported by the available data as distinct from ''Parus''.<ref>Harrap, Simon & Quinn, David (1996): ''Tits, Nuthatches & Treecreepers''. [[Helm Identification Guides|Christopher Helm]], London. {{ISBN|0-7136-3964-4}}</ref> Today, this arrangement is considered [[paraphyletic]] as indicated by [[mtDNA]] [[cytochrome b|cytochrome ''b'']] [[DNA sequence|sequence]] analysis, and ''Parus'' is best restricted to the ''Parus major''—''Parus fasciiventer'' [[clade]], and even the latter species' closest relatives might be considered a distinct genus.<ref name ="Gill">{{cite journal | last1 = Gill | first1 = Frank B. | last2 = Slikas | first2 = Beth | last3 = Sheldon | first3 = Frederick H. | year = 2005 | title = Phylogeny of titmice (Paridae): II. Species relationships based on sequences of the mitochondrial cytochrome-b gene | journal = [[Auk (journal)|Auk]] | volume = 122 | issue = 1| pages = 121–143 | doi = 10.1642/0004-8038(2005)122[0121:POTPIS]2.0.CO;2 | s2cid = 86067032 }}</ref>


In the [[Sibley-Ahlquist taxonomy]], the family Paridae is much enlarged to include related groups such as the [[penduline tit]]s and [[long-tailed tit]]s, but while the former are quite close to the tits and could conceivably be included in that family together with the [[Stenostiridae|stenostirid "warblers"]], the long-tailed tits are not. Indeed, the [[yellow-browed tit]] and the [[sultan tit]] are possibly more distant to the tits than the penduline tits are.<ref name ="Gill"/><ref name ="Jonsson">{{cite journal | last1 = Jønsson | first1 = Knud A. | last2 = Fjeldså | first2 = Jon | year = 2006 | title = Determining biogeographical patterns of dispersal and diversification in oscine passerine birds in Australia, Southeast Asia and Africa | journal = [[Journal of Biogeography|J. Biogeogr.]] | volume = 33 | issue = 7| pages = 1155–1165 | doi = 10.1111/j.1365-2699.2006.01507.x | s2cid = 84542347 }}</ref> If the two current families are lumped into the Paridae, the tits would be a [[subfamily]] '''Parinae'''.
In the [[Sibley-Ahlquist taxonomy]], the family Paridae is much enlarged to include related groups such as the [[penduline tit]]s and [[long-tailed tit]]s, but while the former are quite close to the tits and could conceivably be included in that family together with the [[Stenostiridae|stenostirid "warblers"]], the long-tailed tits are not. Indeed, the [[yellow-browed tit]] and the [[sultan tit]] are possibly more distant to the tits than the penduline tits are.<ref name ="Gill"/><ref name ="Jonsson">{{cite journal | last1 = Jønsson | first1 = Knud A. | last2 = Fjeldså | first2 = Jon | year = 2006 | title = Determining biogeographical patterns of dispersal and diversification in oscine passerine birds in Australia, Southeast Asia and Africa | journal = [[Journal of Biogeography|J. Biogeogr.]] | volume = 33 | issue = 7| pages = 1155–1165 | doi = 10.1111/j.1365-2699.2006.01507.x | s2cid = 84542347 }}</ref> If the two current families are lumped into the Paridae, the tits would be a [[subfamily]] '''Parinae'''.


Alternatively, all tits—save the two monotypic genera discussed in the preceding section and possibly ''Cyanistes'', but including Hume's ground tit—could be lumped in ''Parus''. In any case, four major clades of "typical" tits can be recognized: the dark-capped chickadees and their relatives (''Poecile'' including ''Sittiparus''), the long-crested ''Baeolophus'' and ''Lophophanes'' species, the usually tufted, white-cheeked ''Periparus'' (including ''Pardaliparus'') with more subdued coloration and finally ''Parus sensu stricto'' (including ''Melaniparus'' and ''Machlolophus''). Still, the interrelationship of these, as well as the relationships of many species within the clades, are not well-resolved at all; analysis of [[Morphology (biology)|morphology]] and [[biogeography]] probably gives a more robust picture than the available molecular data.<ref name ="Gill"/>
Alternatively, all tits—save the two monotypic genera discussed in the preceding section and possibly ''Cyanistes'', but including Hume's ground tit—could be lumped in ''Parus''. In any case, four major clades of "typical" tits can be recognized: the dark-capped chickadees and their relatives (''Poecile'' including ''Sittiparus''), the long-crested ''Baeolophus'' and ''Lophophanes'' species, the usually tufted, white-cheeked ''Periparus'' (including ''Pardaliparus'') with more subdued coloration and finally ''Parus sensu stricto'' (including ''Melaniparus'' and ''Machlolophus''). Still, the interrelationship of these, as well as the relationships of many species within the clades, are not well-resolved at all; analysis of [[Morphology (biology)|morphology]] and [[biogeography]] probably gives a more robust picture than the available molecular data.<ref name ="Gill"/>


Tits have settled North America twice, probably at some time during the Early-Mid [[Pliocene]]. The first were the ancestors of ''Baeolophus'', with chickadees arriving somewhat later.<ref name ="Gill"/>
Tits have settled North America twice, probably at some time during the Early-Mid [[Pliocene]]. The first were the ancestors of ''Baeolophus'', with chickadees arriving somewhat later.<ref name ="Gill"/>
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[[File:Tit the bird and thanks for looking.JPG|thumb|right|Tit in a winter tree]]
[[File:Tit the bird and thanks for looking.JPG|thumb|right|Tit in a winter tree]]
[[File:Vogels door Jan van Oort (01).jpg|thumb|250px|Four different tits, although the top-right bird, the [[long-tailed tit]], is not a member of the Paridae]]
[[File:Vogels door Jan van Oort (01).jpg|thumb|250px|Four different tits, although the top-right bird, the [[long-tailed tit]], is not a member of the Paridae]]
'''Family: PARIDAE'''<ref name=ioc>{{cite web| editor1-last=Gill | editor1-first=Frank | editor2-last=Donsker | editor2-first=David | title=Waxwings and their allies, tits & penduline tits | work=World Bird List Version 6.1 | url=http://www.worldbirdnames.org/bow/waxwings/ | publisher=International Ornithologists' Union| access-date= 15 February 2016 }}</ref>
'''Family: PARIDAE'''<ref name="ioc">{{cite web |first=Katerina |last=Twardivoka |date=November 6, 2011 |editor1-last=Gill |editor1-first=Frank |editor2-last=Donsker |editor2-first=David |title=Waxwings and their allies, tits & penduline tits |url=http://www.worldbirdnames.org/bow/waxwings/ |access-date=15 February 2016 |website=World Bird List |version=V. 6.1 |publisher=International Ornithologists' Union}}</ref>
{| class="wikitable"
{| class="wikitable"
|-
|-
! Image !! Genus !! Living species
! Image !! Genus !! Living species
|-
|-
| [[File:Cephalopyrus flammiceps, Bhutan 1.jpg|175px]] || ''Cephalopyrus'' {{small|Bonaparte, 1854}} ||
| [[File:Cephalopyrus flammiceps, Bhutan 1.jpg|175px]] || ''[[Cephalopyrus]]'' {{small|Bonaparte, 1854}} ||
* [[Fire-capped tit]] ''Cephalopyrus flammiceps''
* [[Fire-capped tit]] ''Cephalopyrus flammiceps''
|-
|-
| [[File:Yellow-browed Tit Fambong Lho Wildlife Sanctuary Sikkim 30.03.2014.jpg|175px]]|| ''Sylviparus'' {{small|Burton, 1836}}||
| [[File:Yellow-browed Tit Fambong Lho Wildlife Sanctuary Sikkim 30.03.2014.jpg|175px]]|| ''[[Sylviparus]]'' {{small|Burton, 1836}}||
* [[Yellow-browed tit]], ''Sylviparus modestus''
* [[Yellow-browed tit]], ''Sylviparus modestus''
|-
|-
| [[File:Sultan Tit Mahananda Wildlife Sanctuary Darjeeling, West Bengal, India (cropped).jpg|175px]] || ''Melanochlora'' {{small|Lesson, 1839}} ||
| [[File:Sultan Tit Mahananda Wildlife Sanctuary Darjeeling, West Bengal, India (cropped).jpg|175px]] || ''[[Melanochlora]]'' {{small|Lesson, 1839}} ||
* [[Sultan tit]], ''Melanochlora sultanea''
* [[Sultan tit]], ''Melanochlora sultanea''
|-
|-
| [[File:2017.01.27.-18-Paradiski-La Plagne-Champagny-en-Vanoise--Tannenmeise.jpg|175px]]|| ''[[Periparus]]'' {{small|Sélys Longchamps, 1884}}||
| [[File:2017.01.27.-18-Paradiski-La Plagne-Champagny-en-Vanoise--Tannenmeise.jpg|175px]]|| ''[[Periparus]]'' {{small|Sélys Longchamps, 1884}}||
* [[Rufous-naped tit]], ''Periparus rufonuchalis''
* [[Rufous-naped tit]], ''Periparus rufonuchalis''
* [[Rufous-vented tit]], ''Periparus rubidiventris''
* [[Rufous-vented tit]], ''Periparus rubidiventris''
Line 158: Line 158:
* [[Palawan tit]], ''Pardaliparus amabilis''
* [[Palawan tit]], ''Pardaliparus amabilis''
|-
|-
| [[File:Messange huppée 02.jpg|175px]]|| ''[[Lophophanes]]'' {{small|Kaup, 1829}}||
| [[File:Messange huppée 02.jpg|175px]]|| ''[[Lophophanes]]'' {{small|Kaup, 1829}}||
* [[European crested tit]], ''Lophophanes cristatus''
* [[Crested tit]], ''Lophophanes cristatus''
* [[Grey crested tit]], ''Lophophanes dichrous''
* [[Grey-crested tit]], ''Lophophanes dichrous''
|-
|-
| [[File:Bridled Titmouse (Baeolophus wollweberi) (16875140301).jpg|175px]]|| ''[[Baeolophus]]'' {{small|Cabanis, 1850}}||
| [[File:Bridled Titmouse (Baeolophus wollweberi) (16875140301).jpg|175px]]|| ''[[Baeolophus]]'' {{small|Cabanis, 1850}}||
* [[Bridled titmouse]], ''Baeolophus wollweberi''
* [[Bridled titmouse]], ''Baeolophus wollweberi''
* [[Oak titmouse]], ''Baeolophus inornatus''
* [[Oak titmouse]], ''Baeolophus inornatus''
Line 170: Line 170:
|-
|-
| [[File:Poecile varius on plate.JPG|175px]] || ''[[Sittiparus]]'' {{small|Selys-Longchamps, 1884}} ||
| [[File:Poecile varius on plate.JPG|175px]] || ''[[Sittiparus]]'' {{small|Selys-Longchamps, 1884}} ||
* [[Varied tit]], ''Sittiparus varia''
* [[Varied tit]], ''Sittiparus varius''
* [[Owston's tit]], ''Sittiparus owstoni''
* [[Owston's tit]], ''Sittiparus owstoni''
* [[Iriomote tit]], ''Sittiparus olivaceus''
* [[Iriomote tit]], ''Sittiparus olivaceus''
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* [[White-fronted tit]], ''Sittiparus semilarvatus''
* [[White-fronted tit]], ''Sittiparus semilarvatus''
|-
|-
| [[File:Poecile montanus kleinschmidti.jpg|175px]]|| ''[[Poecile]]'' {{small|Kaup, 1829}}||
| [[File:Poecile montanus kleinschmidti.jpg|175px]]|| ''[[Poecile]]'' {{small|Kaup, 1829}}||
* [[White-browed tit]], ''Poecile superciliosus''
* [[White-browed tit]], ''Poecile superciliosus''
* [[Sombre tit]], ''Poecile lugubris''
* [[Sombre tit]], ''Poecile lugubris''
Line 198: Line 198:
* [[Azure tit]], ''Cyanistes cyanus''
* [[Azure tit]], ''Cyanistes cyanus''
|-
|-
| [[File:Ground Tit north Sikkim India 16.10.2019.jpg|175px]]|| ''Pseudopodoces'' {{small|Zarudny & Loudon, 1902}}||
| [[File:Ground Tit north Sikkim India 16.10.2019.jpg|175px]]|| ''[[Pseudopodoces]]'' {{small|Zarudny & Loudon, 1902}}||
* [[Ground tit]], ''Pseudopodoces humilis'' (previously "Hume's ground jay" in crow family [[Corvidae]]).<ref>James, H. F. et al. (2003). ''Pseudopodoces humilis'', a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones. ''Ibis'' 145: 185–202.[http://www.nmnh.si.edu/vert/birds/pdf/hfj4.pdf pdf file] {{Webarchive|url=https://web.archive.org/web/20060921105947/http://www.nmnh.si.edu/vert/birds/pdf/hfj4.pdf |date=2006-09-21 }}</ref>
* [[Ground tit]], ''Pseudopodoces humilis'' (previously "Hume's ground jay" in crow family [[Corvidae]]).<ref>James, H. F. et al. (2003). ''Pseudopodoces humilis'', a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones. ''Ibis'' 145: 185–202.[http://www.nmnh.si.edu/vert/birds/pdf/hfj4.pdf pdf file] {{Webarchive|url=https://web.archive.org/web/20060921105947/http://www.nmnh.si.edu/vert/birds/pdf/hfj4.pdf |date=2006-09-21 }}</ref>
|-
|-
Line 207: Line 207:
* [[Green-backed tit]], ''Parus monticolus''
* [[Green-backed tit]], ''Parus monticolus''
|-
|-
| [[File:Taiwan tit.jpg|175px]]|| ''[[Machlolophus]]'' {{small|Cabanis, 1850}}||
| [[File:Taiwan tit.jpg|175px]]|| ''[[Machlolophus]]'' {{small|Cabanis, 1850}}||
* [[White-naped tit]], ''Machlolophus nuchalis''
* [[White-naped tit]], ''Machlolophus nuchalis''
* [[Yellow tit]], ''Machlolophus holsti''
* [[Yellow tit]], ''Machlolophus holsti''
* [[Himalayan black-lored tit]], ''Machlolophus xanthogenys''
* [[Himalayan black-lored tit]], ''Machlolophus xanthogenys''
* [[Indian black-lored tit]], ''Machlolophus aplonotus
* [[Indian black-lored tit]], ''Machlolophus aplonotus''
* [[Yellow-cheeked tit]], ''Machlolophus spilonotus''
* [[Yellow-cheeked tit]], ''Machlolophus spilonotus''
|-
|-
| [[File:Parus afer -Namaqua National Park, Northern Cape, South Africa -adult-6.jpg|175px]]|| ''[[Melaniparus]]'' {{small|Bonaparte, 1850}}||
| [[File:Parus afer -Namaqua National Park, Northern Cape, South Africa -adult-6.jpg|175px]]|| ''[[Melaniparus]]'' {{small|Bonaparte, 1850}}||
* [[White-shouldered black tit]], ''Melaniparus guineensis''
* [[White-shouldered black tit]], ''Melaniparus guineensis''
* [[White-winged black tit]], ''Melaniparus leucomelas''
* [[White-winged black tit]], ''Melaniparus leucomelas''

Latest revision as of 19:05, 12 December 2024

Tit
Crested tit in Scotland
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Infraorder: Passerida
Family: Paridae
Vigors, 1825
Genera

5–10, see text.

Global range (In green)
Synonyms

See text

The tits, chickadees, and titmice constitute the Paridae, a large family of small passerine birds which occur mainly in the Northern Hemisphere and Africa. Most were formerly classified in the genus Parus.

Eurasian and African members of this family are referred to as "tits", while North American species are called either "chickadees" (onomatopoeic, derived from their distinctive "chick-a dee dee dee" alarm call)[1] or "titmice". The name titmouse is recorded from the 14th century, composed of the Old English name for the bird, mase (Proto-Germanic *maison, Dutch mees, German Meise), and tit, denoting something small. The former spelling, "titmose", was influenced by mouse in the 16th century.[2] Emigrants to New Zealand presumably identified some of the superficially similar birds of the genus Petroica of the family Petroicidae, the Australian robins, as members of the tit family, giving them the title tomtit, although, in fact, they are not related.

These birds are mainly small, stocky, woodland species with short, stout bills. Some have crests. They range in length from 10 to 22 cm (3.9 to 8.7 in). They are adaptable birds, with a mixed diet including seeds and insects.[3] Many species live around human habitation and come readily to bird feeders for nuts or seed, and learn to take other foods.

Description

[edit]

With the exception of the three monotypic genera Sylviparus, Melanochlora, and Pseudopodoces, the tits are extremely similar in appearance, and have been described as "one of the most conservative avian families in terms of general morphology".[4] The typical body length of adult members of the family is between 10 and 16 cm (3.9 and 6.3 in) in length; when the monotypic genera are added, this range is from 9 to 21 cm (3.5 to 8.3 in). In weight, the family ranges from 5 to 49 g (0.18 to 1.73 oz); this contracts to 7 to 29 g (0.25 to 1.02 oz) when the three atypical genera are removed. The majority of the variation within the family is in plumage, and particularly colour.[5]

The bills of the tits are generally short, varying between stout and fine, depending on diet. The more insectivorous species have finer bills, whereas those that consume more seeds have stouter bills. It is said that tits are evolving longer beaks to reach into bird feeders.[6] The most aberrant bill of the family is possessed by Hume's ground tit of Tibet and the Himalayas, which is long and decurved.[5]

Distribution and habitat

[edit]
The tufted titmouse is restricted to North America.

The tits are a widespread family of birds, occurring over most of Europe, Asia, North America, and Africa. The genus Poecile occurs from Europe through Asia into North America, as far south as southern Mexico. American species in this genus are known as chickadees. Some species in this genus have quite large natural distributions; one, the grey-headed chickadee, is distributed from Scandinavia to Alaska and Canada. The majority of the tits in the genus Periparus are found in the southeastern portion of Asia. This includes two species endemic to the Philippines. The coal tit, also in this genus, is a much more widespread species, ranging from the British Isles and North Africa to Japan. The two crested tits of the genus Lophophanes have a disjunct distribution, with one species occurring in Europe and the other in central Asia.[5]

The genus Baeolophus is endemic to North America. The genus Parus includes the great tit that ranges from Western Europe to Indonesia. Cyanistes has a European and Asian distribution (also into northern Africa), and the three remaining genera, Pseudopodoces, Sylviparus, and Melanochlora, are all restricted to Asia.[5]

Behaviour

[edit]

Tits are active, noisy, and social birds. They are territorial during the breeding season and often join mixed-species feeding flocks during the nonbreeding season. The tits are highly adaptable, and after the corvids (crows and jays) and parrots, amongst the most intelligent of all birds.[5] Tits recognize the difference between species that are dangerous or harmless to them, by this they can protect each other or their families. These birds do this by mobbing or escaping, however they also avoid the nest when the predators are present in order to avoid their families to be seen.

Fission–fusion society

[edit]

Fission–fusion society has been documented in a number of avian taxa including this one.[7][8][9] In brief, that means flocks can split into smaller groups or individuals, and subsequently reunite.

Vocalisations

[edit]
A great tit calling in Finland.

The tits make a variety of calls and songs. They are amongst the most vocal of all birds, calling continuously in most situations, so much so that they are only ever silent for specific reasons such as avoiding predators or when intruding on a rival's territory. Quiet contact calls are made while feeding to facilitate cohesion with others in their social group.[5] Other calls are used for signalling alarm—a well-known example being the "chic-a-dee-dee" of North American species in the genus Poecile, the call which gives them their local common name, the chickadee. The call also serves as a rallying call to summon others to mob and harass the predator. The number of "dee" syllables at the end of the call increases with the level of danger the predator poses.[10]

Diet and feeding

[edit]
Hold-hammering is a common way for the family to deal with food items.
Blue tit with prey item

The tits are generalist insectivores that consume a wide range of small insects and other invertebrates, particularly small defoliating caterpillars. They also consume seeds and nuts, particularly in the winter. One characteristic method of foraging in the family is hanging, where they inspect a branch or twig and leaves from all angles while hanging upside down to feed.[5] In areas where numerous species of tit coexist, different species forage in different parts of the tree, their niche determined in no small way by their morphology; larger species forage on the ground, medium-sized species foraging on larger branches, and the smallest species on the ends of branches. Having obtained larger prey items or seeds, tits engage in hold-hammering, where they hold the item between their feet and hammer it with their bill until it opens. In this fashion, they can even open hazelnuts in around 20 minutes. A number of genera engage in food caching, hoarding supplies of food during the winter.[11]

Breeding

[edit]

Tits are cavity-nesting birds, typically using trees, although Pseudopodoces[12] builds a nest on the ground. Most tree-nesting tits excavate their nests,[13] and clutch sizes are generally large for altricial birds, ranging from usually two eggs in the rufous-vented tit of the Himalayas to as many as 10 to 14 in the blue tit of Europe.[14][15] In favourable conditions, this species had laid as many as 19 eggs, which is the largest clutch of any altricial bird.[3] Most tits are multibrooded, a necessary strategy to cope with either the harsh winters in which they reside in the Holarctic or the extremely erratic conditions of tropical Africa,[15] where typically a single pair cannot find enough food to rear even one nestling and in drought years breeding is likely to be futile.

Many African tit species, along with Pseudopodoces, are cooperative breeders,[16] and even pair-breeding parids are often highly social and maintain stable flocks throughout the nonbreeding season.[17]

Tits also have a variety of methods for attracting mates, primarily through their intricate, bouncing mating dance. Only the blue tit is typically polygynous; all other species are generally monogamous.[18] Courtship feeding is typical of pair-breeding tits to deal with the cost of rearing their large broods.

Systematics

[edit]
The marsh tit was once placed in the genus Parus, but has now been moved to the genus Poecile.

Recently, the large Parus group has been gradually split into several genera (as indicated below), initially by North American ornithological authorities and later elsewhere. Whereas in the mid-1990s, only Pseudopodoces, Baeolophus, Melanochlora, and Sylviparus were considered well-supported by the available data as distinct from Parus.[19] Today, this arrangement is considered paraphyletic as indicated by mtDNA cytochrome b sequence analysis, and Parus is best restricted to the Parus majorParus fasciiventer clade, and even the latter species' closest relatives might be considered a distinct genus.[20]

In the Sibley-Ahlquist taxonomy, the family Paridae is much enlarged to include related groups such as the penduline tits and long-tailed tits, but while the former are quite close to the tits and could conceivably be included in that family together with the stenostirid "warblers", the long-tailed tits are not. Indeed, the yellow-browed tit and the sultan tit are possibly more distant to the tits than the penduline tits are.[20][21] If the two current families are lumped into the Paridae, the tits would be a subfamily Parinae.

Alternatively, all tits—save the two monotypic genera discussed in the preceding section and possibly Cyanistes, but including Hume's ground tit—could be lumped in Parus. In any case, four major clades of "typical" tits can be recognized: the dark-capped chickadees and their relatives (Poecile including Sittiparus), the long-crested Baeolophus and Lophophanes species, the usually tufted, white-cheeked Periparus (including Pardaliparus) with more subdued coloration and finally Parus sensu stricto (including Melaniparus and Machlolophus). Still, the interrelationship of these, as well as the relationships of many species within the clades, are not well-resolved at all; analysis of morphology and biogeography probably gives a more robust picture than the available molecular data.[20]

Tits have settled North America twice, probably at some time during the Early-Mid Pliocene. The first were the ancestors of Baeolophus, with chickadees arriving somewhat later.[20]

Species in taxonomic order

[edit]
 
Phylogeny of the Paridae based on Johansson et al. 2013[22]
Tit in a winter tree
Four different tits, although the top-right bird, the long-tailed tit, is not a member of the Paridae

Family: PARIDAE[23]

Image Genus Living species
Cephalopyrus Bonaparte, 1854
Sylviparus Burton, 1836
Melanochlora Lesson, 1839
Periparus Sélys Longchamps, 1884
Pardaliparus Sélys Longchamps, 1884
Lophophanes Kaup, 1829
Baeolophus Cabanis, 1850
Sittiparus Selys-Longchamps, 1884
Poecile Kaup, 1829
Cyanistes Kaup, 1829
Pseudopodoces Zarudny & Loudon, 1902
Parus Linnaeus, 1758
Machlolophus Cabanis, 1850
Melaniparus Bonaparte, 1850

References

[edit]
  1. ^ The Merriam-Webster New Book of Word Histories. Springfield, MA, USA: Merriam-Webster. 1991. p. 362. ISBN 0-87779-603-3.
  2. ^ Douglas Harper. "titmouse (n.)". Online Etymology Dictionary. Retrieved 2015-04-28.
  3. ^ a b Perrins, C. (1991). Forshaw, Joseph (ed.). Encyclopaedia of Animals: Birds. London: Merehurst Press. pp. 202–203. ISBN 1-85391-186-0.
  4. ^ Gosler & Clement (2007) P.669
  5. ^ a b c d e f g Gosler, Andrew; Clement, Peter (2007). "Family Paridae (Tits and Chickadees)". In del Hoyo, Josep; Elliott, Andrew; Christie, David (eds.). Handbook of the Birds of the World. Volume 12: Picathartes to Tits and Chickadees. Barcelona: Lynx Edicions. pp. 662–709. ISBN 978-84-96553-42-2.
  6. ^ "Great Tits May be Evolving Bigger Beaks. Here's Why". 19 October 2017. Archived from the original on October 20, 2017.
  7. ^ Kendra, Sewall (2015). "Social Complexity as a Driver of Communication and Cognition". Integrative and Comparative Biology. 55 (3): 384–395. doi:10.1093/icb/icv064. PMID 26078368.
  8. ^ Alpin, L. M.; Farine, D. R.; Morand-Ferron, J.; Cole, E. F.; Cockburn, A.; Sheldon, B. C. (2013). "Individual personalities predict social behaviour in wild networks of great tits (Parus major)". Ecology Letters. 16 (11): 1365–1372. doi:10.1111/ele.12181. hdl:1885/66143. PMID 24047530. S2CID 3400989.[dead link]
  9. ^ admin (2022-08-15). "Tufted Titmouse - Best Facts 2022". Birds At First Sight. Retrieved 2022-09-30.
  10. ^ Templeton, Christopher; Greene, Erick; Davis, Kate (2005). "Allometry of Alarm Calls: Black-Capped Chickadees Encode Information About Predator Size". Science. 308 (5730): 1934–1937. Bibcode:2005Sci...308.1934T. doi:10.1126/science.1108841. PMID 15976305. S2CID 42276496.
  11. ^ Jokinen, S; Suhonen, J (1995). "Food Caching By Willow and Crested Tits: A Test of Scatterhoarding Models". Ecology. 76 (3): 892–898. doi:10.2307/1939354. JSTOR 1939354.
  12. ^ "Pseudopodoces humilis, a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones"[permanent dead link]
  13. ^ Mönkkönen, Mikko and Orell, Markku; "Clutch Size and Cavity Excavation in Parids (Paridae): The Limited Breeding Opportunities Hypothesis Tested" in The American Naturalist, Vol. 149, No. 6 (June 1997), pp. 1164–1174
  14. ^ "List of Species and Data Sources Used for Geographic Distributions and Data on Clutch Sizes and Intrinsic Variables"
  15. ^ a b Newton, Ian; Population Limitation in Birds; p. 25. ISBN 978-0-12-517366-7
  16. ^ Johannessen, Lars E. (2011). "Geographical variation in patterns of parentage and relatedness in the co-operatively breeding Ground Tit Parus humilis". Ibis. 153 (2): 373–383. doi:10.1111/j.1474-919X.2011.01115.x.
  17. ^ Stacey, Peter B. and Ligon, J. David; "The Benefits-of-Philopatry Hypothesis for the Evolution of Cooperative Breeding: Variation in Territory Quality and Group Size Effects" in The American Naturalist, Vol. 137, No. 6 (Jun., 1991), pp. 831–846
  18. ^ Andersson, S; Rnborg, J; Andersson, M (1998). "Ultraviolet sexual dimorphism and assortative mating in blue tits". Proc Biol Sci. 265 (1395): 445–450. doi:10.1098/rspb.1998.0315. PMC 1688915.
  19. ^ Harrap, Simon & Quinn, David (1996): Tits, Nuthatches & Treecreepers. Christopher Helm, London. ISBN 0-7136-3964-4
  20. ^ a b c d Gill, Frank B.; Slikas, Beth; Sheldon, Frederick H. (2005). "Phylogeny of titmice (Paridae): II. Species relationships based on sequences of the mitochondrial cytochrome-b gene". Auk. 122 (1): 121–143. doi:10.1642/0004-8038(2005)122[0121:POTPIS]2.0.CO;2. S2CID 86067032.
  21. ^ Jønsson, Knud A.; Fjeldså, Jon (2006). "Determining biogeographical patterns of dispersal and diversification in oscine passerine birds in Australia, Southeast Asia and Africa". J. Biogeogr. 33 (7): 1155–1165. doi:10.1111/j.1365-2699.2006.01507.x. S2CID 84542347.
  22. ^ Johansson, Ulf S; Ekman, Jan; Bowie, Rauri C.K; Halvarsson, Peter; Ohlson, Jan I; Price, Trevor D; Ericson, Per G.P (2013). "A complete multilocus species phylogeny of the tits and chickadees (Aves: Paridae)". Molecular Phylogenetics and Evolution. 69 (3): 852–860. doi:10.1016/j.ympev.2013.06.019. PMID 23831453.
  23. ^ Twardivoka, Katerina (November 6, 2011). Gill, Frank; Donsker, David (eds.). "Waxwings and their allies, tits & penduline tits". World Bird List. V. 6.1. International Ornithologists' Union. Retrieved 15 February 2016.
  24. ^ James, H. F. et al. (2003). Pseudopodoces humilis, a misclassified terrestrial tit (Paridae) of the Tibetan Plateau: evolutionary consequences of shifting adaptive zones. Ibis 145: 185–202.pdf file Archived 2006-09-21 at the Wayback Machine
[edit]