Parrotbill
Encyclopedia
The parrotbills are a group of peculiar bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

s native to East and Southeast Asia
Southeast Asia
Southeast Asia, South-East Asia, South East Asia or Southeastern Asia is a subregion of Asia, consisting of the countries that are geographically south of China, east of India, west of New Guinea and north of Australia. The region lies on the intersection of geological plates, with heavy seismic...

, though feral
Feral
A feral organism is one that has changed from being domesticated to being wild or untamed. In the case of plants it is a movement from cultivated to uncultivated or controlled to volunteer. The introduction of feral animals or plants to their non-native regions, like any introduced species, may...

 populations are known from elsewhere. They are generally small, long-tailed birds which inhabit reedbeds and similar habitat
Habitat
* Habitat , a place where a species lives and grows*Human habitat, a place where humans live, work or play** Space habitat, a space station intended as a permanent settlement...

. They feed mainly on seeds, e.g. of grasses, to which their bill, as the name implies, is well-adapted. Living in tropical to southern temperate
Temperate
In geography, temperate or tepid latitudes of the globe lie between the tropics and the polar circles. The changes in these regions between summer and winter are generally relatively moderate, rather than extreme hot or cold...

 climates, they are usually non-migratory
Bird migration
Bird migration is the regular seasonal journey undertaken by many species of birds. Bird movements include those made in response to changes in food availability, habitat or weather. Sometimes, journeys are not termed "true migration" because they are irregular or in only one direction...

.

The Bearded Reedling or "Bearded Tit", an Eurasia
Eurasia
Eurasia is a continent or supercontinent comprising the traditional continents of Europe and Asia ; covering about 52,990,000 km2 or about 10.6% of the Earth's surface located primarily in the eastern and northern hemispheres...

n species long placed here, is more insectivorous by comparison, especially in summer. It also strikingly differs in morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

, and was time and again placed in a monotypic
Monotypic
In biology, a monotypic taxon is a taxonomic group with only one biological type. The term's usage differs slightly between botany and zoology. The term monotypic has a separate use in conservation biology, monotypic habitat, regarding species habitat conversion eliminating biodiversity and...

 family Panuridae. DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

 data supports this.
As names like "Bearded Tit" imply, their general habitus and acrobatic habits resemble birds like the Long-tailed tit
Long-tailed Tit
The Long-tailed Tit or Long-tailed Bushtit is a common bird found throughout Europe and Asia. There are several extensive accounts of this species, most notably Cramp and Perrins, 1993; Gaston, 1973; and Harrap and Quinn, 1996...

s. Together with these and others they were at some time placed in the titmouse family
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...

 Paridae. Later studies found no justification to presume a close relationship between all these birds, and consequently the parrotbills and Bearded Reedling were removed from the tits and chickadees and placed into a distinct family, Paradoxornithidae. As names like Paradoxornis paradoxus - "puzzling, paradox bird" - suggest, their true relationships were very unclear, although by the latter 20th century they were generally seen as close to Timaliidae ("Old World babblers") and Sylviidae
Sylviidae
Sylviidae is a family of passerine birds that was part of an assemblage known as the Old World warblers. The family was formerly a wastebin taxon with over 400 species of bird in over 70 genera. The family was poorly defined with many characteristics shared with other families...

 ("Old World warblers").

Since 1990 (Sibley & Ahlquist 1990), molecular data has been added to aid the efforts of discovering the parrotbills' true relationships. As Paradoxornis species are generally elusive and in many cases little-known birds, usually specimens of the Bearded Reedling which are far more easy to procure were used for the analyses. Often, the entire group was entirely left out of analyses, being small and seemingly insignificant in the large pattern of bird evolution
Bird evolution
The evolution of birds is thought to have begun in the Jurassic Period, with the earliest birds derived from theropod dinosaurs. Birds are categorized as a biological class, Aves. The earliest known species of class Aves is Archaeopteryx lithographica, from the Late Jurassic period, though...

 (e.g. Barker et al. 2002, 2004). The Bearded Reedling tended to appear close to lark
Lark
Larks are passerine birds of the family Alaudidae. All species occur in the Old World, and in northern and eastern Australia; only one, the Shore Lark, has spread to North America, where it is called the Horned Lark...

s in phylogenies based on e.g. DNA-DNA hybridization (Sibley & Ahlquist 1990), or on mtDNA cytochrome b
Cytochrome b
Cytochrome b/b6 is the main subunit of transmembrane cytochrome bc1 and b6f complexes. In addition, it commonly refers to a region of mtDNA used for population genetics and phylogenetics.- Function :...

 and nDNA c-myc exon
Exon
An exon is a nucleic acid sequence that is represented in the mature form of an RNA molecule either after portions of a precursor RNA have been removed by cis-splicing or when two or more precursor RNA molecules have been ligated by trans-splicing. The mature RNA molecule can be a messenger RNA...

 3, RAG-1 and myoglobin
Myoglobin
Myoglobin is an iron- and oxygen-binding protein found in the muscle tissue of vertebrates in general and in almost all mammals. It is related to hemoglobin, which is the iron- and oxygen-binding protein in blood, specifically in the red blood cells. The only time myoglobin is found in the...

 intron
Intron
An intron is any nucleotide sequence within a gene that is removed by RNA splicing to generate the final mature RNA product of a gene. The term intron refers to both the DNA sequence within a gene, and the corresponding sequence in RNA transcripts. Sequences that are joined together in the final...

 2 sequence data (Ericson & Johansson 2003). Placement in a superfamily
Taxonomic rank
In biological classification, rank is the level in a taxonomic hierarchy. Examples of taxonomic ranks are species, genus, family, and class. Each rank subsumes under it a number of less general categories...

 Sylvioidea
Sylvioidea
Sylvioidea is a superfamily of passerine birds. It is one of at least three major clades within the Passerida along with the Muscicapoidea and Passeroidea. It contains about 1300 species including the Old World warblers, Old World babblers, swallows, larks, bulbuls and perhaps the tits...

 which contained birds such as Sylviidae, Timaliidae and long-tailed tits - but not Paridae - was confirmed.

Cibois (2003a) analyzed mtDNA cytochrome b and 12S/16S rRNA sequences of some Sylvioidea, among them several species of Paradoxornis but not the Bearded Reedling. These formed a robust clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 closer to the Sylvia typical warblers and some presumed "Old World babblers" such as Chrysomma sinense than to other birds. The puzzle was finally resolved by Alström et al. (2006), who studied mtDNA cytochrome b and nDNA myoglobin intron 2 sequences of a wider range of Sylvioidea: The Bearded Reedling was not a parrotbill at all, but forms a distinct lineage on its own, the relationships of which are not entirely resolved at present. The parrotbills' presence in the clade containing Sylvia, on the other hand, necessitates that the Paradoxornithidae are placed in synonymy of the Sylviidae. Cibois (2003b) even suggested that these themselves were to be merged with the remaining Timaliidae and the latter name to be adopted. This has hitherto not been followed and researchers remain equivocal as many taxa in Sylviidae and Timaliidae remain to be tested for their relationships. In any case, it is most likely that the typical warbler-parrotbill group is monophyletic and therefore agrees with the modern requirements for a taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

. Hence, whether to keep or to synonymize it is entirely a matter of philosophy, as the scientific facts would agree with either approach.

The interesting conclusion from an evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

ary point of view is that the morphologically both internally homogenous and compared to each other highly dissimilar typical warblers and parrotbills form the two extremes in the divergent evolution
Divergent evolution
Divergent evolution is the accumulation of differences between groups which can lead to the formation of new species, usually a result of diffusion of the same species to different and isolated environments which blocks the gene flow among the distinct populations allowing differentiated fixation...

 of the Sylviidae. This is underscored by looking at the closest living relatives of the parrotbills in the rearranged Sylviidae: The genus Chrysomma
Chrysomma
Chrysomma is a songbird genus. Long believed to be Timaliidae , they are actually quite closely related to the typical warblers and even closer to the parrotbills, and therefore a member of the family Sylviidae.Species are:...

are non-specialized species altogether intermediate in habitus, habitat and habits between the typical warblers and the parrotbills. Presumably, the ancestral sylviids looked much like these birds. How dramatic the evolutionary changes wrought upon the parrotbills in their adaptation to feeding on grass caryopses and similar seeds were can be seen by comparing them with the typical fulvetta
Fulvetta
Fulvetta is a songbird genus. Originally proposed in 1877, it was recently reestablished for the typical fulvettas, which were long included with their presumed relatives in the Timaliidae genus Alcippe...

s, which were formerly considered Timaliidae and united with the alcippe
Alcippe (bird)
Alcippe is a genus of birds in the Timaliidae family. It used to be a catchall for the "fulvettas", but has been recently split into 5 independent genera: Alcippe, Schoeniparus, Fulvetta, Pseudominla and Lioparus....

s (Pasquet 2006). These look somewhat like drab fairy-wrens and have none of the parrotbills' adaptations to food and habitat. Yet it appears that the typical fulvettas' and parrotbills' common ancestor evolved into at least two parrotbill lineages independently (Cibois 2003a) & (Yeung et al. 2006). Only the Wrentit
Wrentit
The Wrentit is a small bird that lives in chaparral and bushland. It is the only species in the genus Chamaea....

, the only American
Americas
The Americas, or America , are lands in the Western hemisphere, also known as the New World. In English, the plural form the Americas is often used to refer to the landmasses of North America and South America with their associated islands and regions, while the singular form America is primarily...

 sylviid, resembles the parrotbills much in habitus, though not in color pattern, and of course, as an insectivore
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....

, neither in bill shape.

Species of parrotbills

Paradoxornis is apparently paraphyletic with Conostoma. Deep divergences were found between major clades; basally Conostoma with a clade of large species followed by two clades of smaller species which differ markedly in plumage pattern. This with egg coloration data (Walters 2006) lends considerable support for splitting it up into at least three genera and possibly up to eight. (see http://worldbirdinfo.net/)

Clade of large species

Genus Conostoma
  • Great Parrotbill
    Great Parrotbill
    The Great Parrotbill is a bird species often placed with the Old World babblers or in a distinct family Paradoxornithidae, but it actually seems to belong to the Sylviidae. Its genus Conostoma is monotypic.It is found in Bhutan, China, India, Myanmar, and Nepal.-References:* BirdLife...

    ,
    Conostoma oemodium


Genus Paradoxornis


(Sub)Genus Hemirhynchus or Cholornis
  • Three-toed Parrotbill
    Three-toed Parrotbill
    The Three-toed Parrotbill is a species of bird in the Timaliidae family.It is endemic to China.Its natural habitat is temperate forests.-References:* BirdLife International 2004. . Downloaded on 26 July 2007....

    ,
    Paradoxornis (Hemirhynchus) paradoxus
  • Brown Parrotbill
    Brown Parrotbill
    The Brown Parrotbill is a bird species often placed with the Old World babblers or in a distinct family Paradoxornithidae, but it actually seems to belong to the Sylviidae....

    ,
    Paradoxornis (Hemirhynchus) unicolor


(Sub)Genus Paradoxornis
Eggs white with various amounts of brown sprinkling or speckling. More basal
Basal (phylogenetics)
In phylogenetics, a basal clade is the earliest clade to branch in a larger clade; it appears at the base of a cladogram.A basal group forms an outgroup to the rest of the clade, such as in the following example:...

 lineage, possibly close to Golden-breasted Fulvetta
Golden-breasted Fulvetta
The Golden-breasted "Fulvetta" is a species of songbird found in Bhutan, China, India, Myanmar, Nepal, and Vietnam...

 (Lioparus chrysotis) and/or White-browed Chinese Warbler
White-browed Chinese Warbler
The Chinese Hill Warbler , also known as the White-browed Chinese Warbler or Chinese Bush-dweller, is a species of bird in the monotypic genus Rhopophilus. Its relationships are uncertain and it has been placed in the families Cisticolidae, Timaliidae or Sylviidae...

 (Rhopophilus pekinensis).
  • Spot-breasted Parrotbill
    Spot-breasted Parrotbill
    The Spot-breasted Parrotbill is a species of bird in the Timaliidae family.It is found in Bangladesh, China, India, Laos, Myanmar, Thailand, and Vietnam.-References:...

    , Paradoxornis guttaticollis
  • Black-breasted Parrotbill
    Black-breasted Parrotbill
    The Black-breasted Parrotbill is a 19 cm long, large, thick-billed parrotbill with black patches on the head-sides and throat...

    , Paradoxornis flavirostris


(Sub)Genus Psittiparus
Eggs pale cream or bluish with more intense pattern
  • Grey-headed Parrotbill
    Grey-headed Parrotbill
    The Grey-headed Parrotbill is a bird species often placed with the Old World babblers or in a distinct family Paradoxornithidae, but it actually seems to belong to the Sylviidae....

    ,
    Paradoxornis (Psittiparus) gularis
  • Black-headed Parrotbill, Paradoxornis (Psittiparus) margaritae - formerly included in P. gularis
  • White-breasted Parrotbill, Greater Rufous-headed Parrotbill, Paradoxornis (Psittiparus) ruficeps
    • Rufous-headed Parrotbill, P. (P.) (ruficeps) bakeri


(Sub)Genus
Calamornis
Eggs pale green to white, patterned like in Psittiparus.
  • Reed Parrotbill
    Reed Parrotbill
    The Reed Parrotbill is a species of bird in the Timaliidae family. It is found in China, Mongolia, and Russia. It is threatened by habitat loss....

    , Paradoxornis (Calamornis) heudei
    • Northern Parrotbill, Paradoxornis (Calamornis) (heudei) polivanovi

Clade of small brownish species

Small unmarked eggs, mid-blue or paler. Possibly close to any or all of Fulvetta
Fulvetta
Fulvetta is a songbird genus. Originally proposed in 1877, it was recently reestablished for the typical fulvettas, which were long included with their presumed relatives in the Timaliidae genus Alcippe...

(typical fulvettas), Chrysomma
Chrysomma
Chrysomma is a songbird genus. Long believed to be Timaliidae , they are actually quite closely related to the typical warblers and even closer to the parrotbills, and therefore a member of the family Sylviidae.Species are:...

, or Wrentit
Wrentit
The Wrentit is a small bird that lives in chaparral and bushland. It is the only species in the genus Chamaea....



(Sub)Genus Chleuasicus
  • Black-browed Parrotbill
    Black-browed Parrotbill
    The Pale-billed Parrotbill , also known as the Black-browed Parrotbill or Lesser Rufous-headed Parrotbill, is a species of bird in the Timaliidae family....

    ,
    Paradoxornis (Chleuasicus) atrosuperciliaris


(Sub)Genus Sinoparadoxornis
  • Spectacled Parrotbill
    Spectacled Parrotbill
    The Spectacled Parrotbill is a species of bird in the Timaliidae family. It is endemic to central China. Its natural habitats are temperate forests and subtropical or tropical moist montane forests.-References:...

    , Paradoxornis (Sinoparadoxornis) conspicillatus
  • Brown-winged Parrotbill
    Brown-winged Parrotbill
    The Brown-winged Parrotbill is a bird species often placed with the Old World babblers or in a distinct family Paradoxornithidae, but it actually seems to belong to the Sylviidae....

    , Paradoxornis (Sinoparadoxornis) brunneus
  • Yunnan Parrotbill
    Yunnan Parrotbill
    The Yunnan Parrotbill, Paradoxornis ricketti, is a species of bird in the Old World babbler family. This 10 cm long parrotbill is endemic to China, breeding in northwest Yunnan....

    , Paradoxornis (Sinoparadoxornis) ricketti
  • Ashy-throated Parrotbill
    Ashy-throated Parrotbill
    The Ashy-throated Parrotbill, Paradoxornis alphonsianus, is a songbird. In old sources, it may be called Alphonse's Crow-tit; though superficially resembling a tit it is not a member of the Paridae...

    , Paradoxornis (Sinoparadoxornis) alphonsianus
  • Vinous-throated Parrotbill
    Vinous-throated Parrotbill
    The Vinous-throated Parrotbill is a species of bird in the Timaliidae family. It is found in China, Japan, North Korea, South Korea, Mongolia, Russia, Taiwan, and Vietnam...

    , Paradoxornis (Sinoparadoxornis) webbianus
  • Grey-hooded Parrotbill
    Grey-hooded Parrotbill
    The Grey-hooded Parrotbill is a species of bird in the Timaliidae family.It is endemic to China.Its natural habitats are temperate forests and temperate shrubland.It is threatened by habitat loss.-References:...

    , Paradoxornis (Sinoparadoxornis) zappeyi
  • Rusty-throated Parrotbill
    Rusty-throated Parrotbill
    The Przevalski's Parrotbill or Rusty-throated Parrotbill is a species of bird in the Timaliidae family. It is endemic to a small area of central China. Its natural habitat is temperate forests. It is threatened by habitat loss.-References:* BirdLife International 2004. . Downloaded on 26 July...

    , Paradoxornis (Sinoparadoxornis) przewalskii

Clade of small yellowish species

Small unmarked eggs, mid-blue or paler. Possibly close to any or all of Fulvetta
Fulvetta
Fulvetta is a songbird genus. Originally proposed in 1877, it was recently reestablished for the typical fulvettas, which were long included with their presumed relatives in the Timaliidae genus Alcippe...

(typical fulvettas), Chrysomma
Chrysomma
Chrysomma is a songbird genus. Long believed to be Timaliidae , they are actually quite closely related to the typical warblers and even closer to the parrotbills, and therefore a member of the family Sylviidae.Species are:...

, or Wrentit
Wrentit
The Wrentit is a small bird that lives in chaparral and bushland. It is the only species in the genus Chamaea....


(Sub)Genus Suthora
  • Fulvous Parrotbill
    Fulvous Parrotbill
    The Fulvous Parrotbill is a species of bird in the Timaliidae family.It is found in Bhutan, China, India, Myanmar, and Nepal.Its natural habitat is temperate forests.-References:...

    ,
    Paradoxornis (Suthora) fulvifrons
  • Black-throated Parrotbill
    Black-throated Parrotbill
    The Black-throated Parrotbill is a bird species often placed with the Old World babblers or in a distinct family Paradoxornithidae, but it actually seems to belong to the Sylviidae....

    ,
    Paradoxornis (Suthora) nipalensis
    • Blyth's Parrotbill, Paradoxornis (Suthora) (nipalensis) poliotis
  • Golden Parrotbill
    Golden Parrotbill
    The Golden Parrotbill is a species of bird in the Timaliidae family. It is found in China, Laos, Myanmar, Taiwan, and Vietnam. Its natural habitat is subtropical or tropical moist montane forests....

    ,
    Paradoxornis (Suthora) verreauxi

Paradoxornithinae?

Conceivably, the parrotbills and their closest relatives might be considered a distinct subfamily Paradoxornithinae; they appear to form a fairly well-supported clade though the position in regard to basal Sylviidae is unclear (Cibois 2003a, Jønsson & Fjeldså 2006).
  • Genus Lioparus - formerly in Alcippe (Timaliidae)
    • Golden-breasted Fulvetta
      Golden-breasted Fulvetta
      The Golden-breasted "Fulvetta" is a species of songbird found in Bhutan, China, India, Myanmar, Nepal, and Vietnam...

      , Lioparus chrysotis

  • Genus Fulvetta
    Fulvetta
    Fulvetta is a songbird genus. Originally proposed in 1877, it was recently reestablished for the typical fulvettas, which were long included with their presumed relatives in the Timaliidae genus Alcippe...

     - typical fulvettas. Formerly in
    Alcippe (Timaliidae)
    • Spectacled Fulvetta
      Spectacled Fulvetta
      The Spectacled Fulvetta is a bird species in the family Sylviidae. Like the other typical fulvettas, it was long included in the Timaliidae genus Alcippe....

      ,
      Fulvetta ruficapilla
    • Chinese Fulvetta
      Chinese Fulvetta
      The Chinese Fulvetta is a bird species in the family Sylviidae. Like the other typical fulvettas, it was long included in the Timaliidae genus Alcippe.It is found in China and India.-References:...

      ,
      Fulvetta striaticollis
    • White-browed Fulvetta
      White-browed Fulvetta
      The White-browed Fulvetta is a bird species in the family Sylviidae. Like the other typical fulvettas, it was long included in the Timaliidae genus Alcippe....

      ,
      Fulvetta vinipectus
    • Grey-hooded Fulvetta
      Grey-hooded Fulvetta
      The Grey-hooded Fulvetta is a bird species in the family Sylviidae. Like the other typical fulvettas, it was long included in the Timaliidae genus Alcippe....

      ,
      Fulvetta cinereiceps
    • Taiwan Fulvetta
      Taiwan Fulvetta
      The Taiwan Fulvetta is a bird species in the family Sylviidae. Like the other typical fulvettas, it was long included in the Timaliidae genus Alcippe. In addition, it was long included in F. cinereiceps as a subspecies....

      ,
      Fulvetta formosana - formerly in F. cinereiceps
    • Streak-throated Fulvetta
      Streak-throated Fulvetta
      The Streak-throated Fulvetta or Manipur Fulvetta is a bird species in the family Sylviidae. It is named for the state of Manipur in northeastern India. Like the other typical fulvettas, it was long included in the Timaliidae genus Alcippe. In addition, it was long included in F...

      ,
      Fulvetta manipurensis - formerly in F. cinereiceps
    • Ludlow's Fulvetta
      Ludlow's Fulvetta
      The Brown-throated Fulvetta or Ludlow's Fulvetta is a species of bird in the Timaliidae family.It is found in Bhutan, China, India, Myanmar, and Nepal.-References:...

      ,
      Fulvetta ludlowi - tentatively placed here

  • Genus Chrysomma
    Chrysomma
    Chrysomma is a songbird genus. Long believed to be Timaliidae , they are actually quite closely related to the typical warblers and even closer to the parrotbills, and therefore a member of the family Sylviidae.Species are:...

     - formerly in Timaliidae
    • Yellow-eyed Babbler
      Yellow-eyed Babbler
      The Yellow-eyed Babbler Chrysomma sinense is a passerine bird species found in open grass and scrub in south Asia. Its common name refers to its traditional placement with the Old World babbler family Timaliidae...

      , Chrysomma sinense
    • Jerdon's Babbler
      Jerdon's Babbler
      Jerdon's Babbler is an endangered passerine bird from South Asia. Formerly placed in the Timaliidae family – hence the common name "babbler" –, the genus Chrysomma and its relatives are actually closer to the typical warblers and parrotbills in the Sylviidae.-Description:Measuring...

      , Chrysomma altirostre
      • Burmese Jerdon's Babbler, Chrysomma altirostre altirostre - extinct (1940s)
    • Rufous-tailed Babbler
      Rufous-tailed Babbler
      The Rufous-tailed Babbler is a bird species in the family Sylviidae. It was formerly included with the Old World babblers , but is actually very closely related to the typical warblers . It is endemic to China.-References:* BirdLife International 2004. . Downloaded on 25 July 2007.* Collar,...

      , Chrysomma poecilotis

  • Genus Chamaea - Wrentit

  • Genus Rhopophilus - White-browed Chinese Warbler

External links

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