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Crossbill

From Wikipedia, the free encyclopedia
(Redirected fromLoxia)
Genus of birds
This article is about the genus of birds. For the lens series, seeZeiss Loxia. For the medical condition, seetorticollis.

Crossbill
Red crossbill or common crossbill (Loxia curvirostra)
Scientific classificationEdit this classification
Kingdom:Animalia
Phylum:Chordata
Class:Aves
Order:Passeriformes
Family:Fringillidae
Subfamily:Carduelinae
Genus:Loxia
Linnaeus, 1758
Type species
Loxia curvirostra
Species

Loxia curvirostra
Loxia leucoptera
Loxia megaplaga
Loxia pytyopsittacus
Loxia scotia
Loxia sinesciuris
Loxia patevi

Crossbills arebirds of thegenusLoxia within thefinch family (Fringillidae), with six extant and oneextinct species. These birds are characterized by their mandibles with crossed tips, which gives the group its English name. Adult males tend to be red or orange in color, and females green or yellow, but there is much variation.

Crossbills are specialist feeders onconifer cones, and the unusual bill shape is an adaptation which enables them to extract seeds from cones. These birds are typically found in higher northern hemisphere latitudes, where their food sources grow. They irrupt out of the breeding range when the cone crop fails. Crossbills breed very early in the year, often in winter months, to take advantage of maximum cone supplies.

Systematics and evolution

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The genusLoxia was introduced by the Swedish naturalistCarl Linnaeus in 1758 in the10th edition of hisSystema Naturae.[1] The name is from theAncient Greekloxos, "crosswise".[2] The Swiss naturalistConrad Gessner had used the wordLoxia for a crossbill in 1555 in hisHistoriae Animalium.[3] Thetype species was designated asLoxia curvirostra (red crossbill) byGeorge Robert Gray in 1840.[4][5]

Analysis ofmitochondrialcytochromebsequence data indicates that the crossbills share a common ancestor with theredpoll, which diverged during theTortonian (c. 8mya, LateMiocene).[6] The research suggests that the generaLoxia andCarduelis might be merged into a single genus, for which the nameLoxia would then have priority. But this would imply changing the name of a large number of species, and given that the adaptations of the crossbills represent a unique evolutionary path (seeEvolutionary grade), it seems more appropriate to split up the genusCarduelis as it had already been done during most of the 20th century. The fossil record ofLoxia is restricted to aLate Pliocene (c. 2mya) species,Loxia patevi, found atVarshets, Bulgaria.

The species of crossbills are difficult to separate, and care is needed even with the white-winged and Hispaniolan crossbills, the easiest. The other species are identified by subtle differences in head shape and bill size, and the identification problems formerly led to much taxonomic speculation, with some scientists considering that the parrot and Scottish crossbills and possibly the Hispaniolan and white-winged crossbills are conspecific.

The identification problem is least severe in North America, where only the red, white-winged and (locally) Cassia species occur, and (possibly) the most challenging in theScottish Highlands, where three similar-looking species breed and the two-barred (known as white-winged in North America) is a possiblevagrant.

Work on vocalization in North America suggests that there are eight or nine discrete populations of red crossbill in that continent alone, which do not interbreed and are (like the named species) adapted to specialize in different conifer species. While several ornithologists seem inclined to give these forms species status, no division of the American red crossbills has yet occurred.[7] Preliminary investigations inEurope andAsia suggest an equal, if not greater, complexity, with several different call types identified;[8][9] these call types being as different from each other as from the named species of the parrot and Scottish crossbills - suggesting either that theyare valid species, or else that the parrot and Scottish crossbills maynot be.

Genetic research on theirDNA failed to reveal any difference between any of the crossbills (including the morphologically distinct two-barred), with variation between individuals greater than any difference between the taxa. This led to the suggestion thatlimited interbreeding between the different types prevented significant genetic differentiation, and enabled each type to maintain a degree of morphological plasticity, which may be necessary to enable them to feed on different conifers when their preferred food species has a crop failure. Research in Scotland, however, has shown that the parrot and Scottish crossbills are reproductively isolated from each other and also from the red crossbill, despite irruption of that species into their ranges, and the diagnostic calls and bill dimensions have not been lost. They are, therefore, good species.[10]

Currently accepted species[11] and their preferred food sources are:

ImageScientific nameCommon nameFood sourceDistribution of species
Loxia curvirostraRed crossbillSpruce (Picea) species; some populations (distinct species?) on variouspine (Pinus) species and (in western North America)Douglas firEurasia,North Africa, andNorth America
Loxia leucopteraTwo-barred crossbillLarch (Larix) species, particularlyL. sibirica,L. gmelinii,L. laricina and (in North America) alsohemlock (Tsuga)Eurasia andNorth America
Loxia megaplagaHispaniolan crossbillHispaniolan pine (Pinus occidentalis)Hispaniola (Haiti and theDominican Republic)
Loxia pytyopsittacusParrot crossbillScots pine (Pinus sylvestris)Northern,Eastern, andCentral Europe
Loxia scoticaScottish crossbillScots pine (Pinus sylvestris) andlarch (Larix) species (particularly plantations ofL. decidua)Scotland
Loxia sinesciurisCassia crossbillIsolated population of thelodgepole pine (Pinus contorta latifolia)South Hills andAlbion Mountains,Idaho,United States
Loxia pateviLoxia pateviUnknownFossils dated to theLate Pliocene are known fromVarshets,Bulgaria

Originally, several species such as thechestnut-backed sparrow-lark (Eremopterix leucotis),[12]pine grosbeak (Pinicola enucleator) andnorthern cardinal (Cardinalis cardinalis)[13] were also included inLoxia.

Feeding behavior

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Red crossbill skull and jaw anatomy fromWilliam Yarrell'sA History of British Birds (1843)

The different species specialize in feeding on different conifer species, with the bill shape optimized for opening that species of conifer. This is achieved by inserting the bill between the conifer cone scales and twisting the lower mandible towards the side to which it crosses, enabling the bird to extract the seed at the bottom of the scale with its tongue.

The mechanism by which the bill-crossing is developed (which usually, but not always, occurs in a 1:1 frequency of left-crossing or right-crossingmorphs), and what determines the direction, has hitherto withstood all attempts to resolve it.

It is very probable that there is a genetic basis underlying the phenomenon (young birds whose bills are still straight will give a cone-opening behavior if their bills are gently pressed, and the crossing develops before the birds are fledged and feeding independently), but at least in the red crossbill (the only species which has been somewhat thoroughly researched regarding this question) there is no straightforward mechanism ofheritability.

While the direction of crossing seems to be the result of at least three genetic factors working together in a case ofepistasis and most probablyautosomal, it is not clear whether the 1:1 frequency of both morphs in most cases is the result of genetics or environmental selection. Populations that feed on cones without removing or twisting them will likely show a 1:1 morph distribution, no matter what the genetic basis may be: thefitness of each morph is inversely proportional to its frequency in the population. Such birds can only access the cone with the lower mandible tip pointingtowards it to successfully extract seeds, and thus a too high number of birds of one morph will result in the food availability for each bird of this morph decreasing.[14]

They can utilize other conifers to their preferred, and often need to do so when their preferred species has a crop failure, but are less efficient in their feeding (not enough to prevent survival, but probably enough to reduce breeding success).

Fossil record

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Loxia patevi was described from theLate Pliocene ofVarshets, Bulgaria.[15]

References

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  1. ^Linnaeus, C. (1758).Systema Naturæ per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis (in Latin). Vol. 1 (10th ed.). Holmiae:Laurentii Salvii. p. 171.{{cite book}}: CS1 maint: publisher location (link)
  2. ^Jobling, James A. (2010).The Helm Dictionary of Scientific Bird Names. London, United Kingdom: Christopher Helm. p. 231.ISBN 978-1-4081-2501-4.
  3. ^Gesner, Conrad (1555).Historiae animalium liber III qui est de auium natura. Adiecti sunt ab initio indices alphabetici decem super nominibus auium in totidem linguis diuersis: & ante illos enumeratio auium eo ordiné quo in hoc volumine continentur (in Latin). Zurich: Froschauer. p. 568.
  4. ^Gray, George Robert (1840).A List of the Genera of Birds : with an Indication of the Typical Species of Each Genus. London: R. and J.E. Taylor. p. 49.
  5. ^Paynter, Raymond A. Jr, ed. (1968).Check-List of Birds of the World. Vol. 14. Cambridge, Massachusetts: Museum of Comparative Zoology. p. 287.
  6. ^Arnaiz-Villena, A.; Guillén, J.; Ruiz-del-Valle, V.; Lowy, E.; Zamora, J.; Varela, P.; Stefani, D.; Allende, L. M. (2001)."Phylogeography of crossbills, bullfinches, grosbeaks, and rosefinches"(PDF).Cellular and Molecular Life Sciences.58 (8):1159–1166.doi:10.1007/PL00000930.PMID 11529508.S2CID 6241573.
  7. ^[1] (2011)
  8. ^Förschler, Marc I.; Kalko, Elisabeth K. V. (2009). "Vocal types in crossbill populations (Loxia spp.) of Southwest Europe".Journal of Ornithology.150:17–27.doi:10.1007/s10336-008-0312-z.S2CID 13074563.
  9. ^Robb, Magnus S (2000). "Introduction to vocalizations of crossbills in north-western Europe".Dutch Birding.22 (2):61–107.
  10. ^Summers, R. W.; Dawson, R. J.; Phillips, R. E. (2007). "Assortative mating and patterns of inheritance indicate that the three crossbill taxa in Scotland are species".Journal of Avian Biology.38 (2):153–162.doi:10.1111/j.0908-8857.2007.03798.x.
  11. ^Gill, Frank; Donsker, David (eds.)."Finches, euphonias".World Bird List Version 5.2. International Ornithologists' Union. Retrieved5 June 2015.
  12. ^"Eremopterix leucotis - Avibase".avibase.bsc-eoc.org. Retrieved2016-11-18.
  13. ^Linnaeus, Carl (1758).Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. T(Laurentii Salvii) (in Latin). Vol. 1. p. 824.
  14. ^Edelaar, Pim; Postma, Erik; Knops, Peter; Phillips, Ron (2005)."No support of a genetic basis of mandible crossing direction in crossbills (Loxia spp.)"(PDF).Auk.122 (4):1123–1129.doi:10.1642/0004-8038(2005)122[1123:NSFAGB]2.0.CO;2.hdl:20.500.11755/488d3732-6dd9-49f2-be2a-efcbfec9e08d.JSTOR 4090517.
  15. ^Boev, Z. 1999. Earliest finds of crossbills (genusLoxia) (Aves: Fringillidae) from Varshets (NW Bulgaria). - Geologica balcanica, 29 (3-4): 51-57.

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