Editing Anopheles (section)

== Evolution ==
=== Fossil history ===
[[Fossil]]s of the genus ''Anopheles'' are rare; only two had been found by 2015.<ref name="Freitas Russo Voloch Mutaquiha 2015"/> They are ''Anopheles (Nyssorhynchus) dominicanus'' <small>Zavortink & Poinar</small> in Dominican Republic [[amber]] from the Late [[Eocene]] ({{Ma|40.4}} to {{Ma|33.9}}),<ref name="Zavortink Poinar 2000">{{cite journal |last1=Zavortink |first1=Thomas J. |last2=Poinar |first2=George O. |title=Anopheles (Nyssorhynchus) dominicanus sp. n. (Diptera: Culicidae) from Dominican Amber |journal=Annals of the Entomological Society of America |volume=93 |issue=6 |date=2000-11-01 |doi=10.1603/0013-8746(2000)093[1230:ANDSND]2.0.CO;2 |pages=1230–1235|s2cid=86221732 }}</ref> and ''[[Anopheles rottensis]]'' <small>Statz</small> in [[Germany|German]] amber from the Late [[Oligocene]] ({{Ma|28.4}} to {{Ma|23.0}}).<ref name="Freitas Russo Voloch Mutaquiha 2015">{{cite journal |last1=Freitas |first1=Lucas A. |last2=Russo |first2=Claudia A. M. |last3=Voloch |first3=Carolina M. |last4=Mutaquiha |first4=Olívio C. F. |last5=Marques |first5=Lucas P. |last6=Schrago |first6=Carlos G. |title=Diversification of the Genus Anopheles and a Neotropical Clade from the Late Cretaceous |journal=PLOS ONE |volume=10 |issue=8 |date=2015-08-05 |pmid=26244561 |pmc=4526650 |doi=10.1371/journal.pone.0134462 |page=e0134462 |bibcode=2015PLoSO..1034462F |doi-access=free }}</ref>

=== Phylogeny ===

The ancestors of all [[Fly|flies]] including mosquitoes appeared {{Ma|260}}.<ref name="Moreno Marinotti Krzywinski Tadei 2010">{{cite journal |last1=Moreno |first1=Marta |last2=Marinotti |first2=Osvaldo |last3=Krzywinski |first3=Jaroslaw |last4=Tadei |first4=Wanderli P. |last5=James |first5=Anthony A. |last6=Achee |first6=Nicole L. |last7=Conn |first7=Jan E. |title=Complete mtDNA genomes of ''Anopheles darlingi'' and an approach to anopheline divergence time |journal=Malaria Journal |volume=9 |issue=1 |date=2010 |page=127 |pmid=20470395 |pmc=2877063 |doi=10.1186/1475-2875-9-127 |doi-access=free}}</ref> The [[Culicinae|culicine]] and ''Anopheles'' clades of mosquitoes diverged between {{Ma|120}} and {{Ma|150}}.<ref name="Moreno Marinotti Krzywinski Tadei 2010"/><ref name="Calvo 2009">{{cite journal |author1=Calvo, Eric |author2=Pham, Van M. |author3=Marinotti, Osvaldo |author4=Andersen, John F. |author5=Ribeiro, José M. C. |year=2009 |title=The salivary gland transcriptome of the neotropical malaria vector ''Anopheles darlingi'' reveals accelerated evolution of genes relevant to hematophagy |journal=[[BMC Genomics]] |volume=10 |issue=1 |page=57 |doi=10.1186/1471-2164-10-57 |pmc=2644710 |pmid=19178717 |doi-access=free}}</ref> The Old and New World ''Anopheles'' species subsequently diverged between {{Ma|80}} and {{Ma|95}}.<ref name="Moreno Marinotti Krzywinski Tadei 2010"/><ref name="Calvo 2009"/> ''[[Anopheles darlingi]]'' diverged from the African and Asian malaria vectors ~{{Ma|100}}.<ref name="Neafsey Waterhouse Abai 2015">{{cite journal |last1=Neafsey |first1=Daniel E. |last2=Waterhouse |first2=Robert M. |last3=Abai |first3=Mohammad R. |last4=Aganezov |first4=Sergey S. |last5=Alekseyev |first5=Max A. |display-authors=etal |title=Highly evolvable malaria vectors: The genomes of 16 Anopheles mosquitoes |journal=Science |volume=347 |issue=6217 |date=2015-01-02 |page=43 |pmid=25554792 |pmc=4380271 |doi=10.1126/science.1258522|bibcode=2015Sci...347...43N }}</ref>  The cladogram is based on an analysis of mosquito [[genome]]s by Heafsey and colleagues in 2015:<ref name="Neafsey Waterhouse Abai 2015"/>

{{clade
|label1=[[Diptera]]
|sublabel1= 260 [[myr|mya]]
|1={{clade
   |label1=[[Culicidae]]
   |sublabel1= ''mosquitoes''
   |1={{clade
      |label1='''''Anopheles'''''
      |sublabel1= 100 [[myr|mya]]
      |1={{clade
         |1={{clade
            |label1=subgenus ''[[Anopheles (Cellia)|Cellia]]''
            |1={{clade
               |1={{clade
               |sublabel1= 30 [[myr|mya]]
                  |label1=subgenus ''Pyretophorus''
                  |1={{clade
                     |1=''[[gambiae]]'' complex (Africa)
                     |2=''A. epirotus'' & spp.
                     }}
                  |2=''A. stephensi'' & spp.
                  }}
               |2=''A. dirus'' & spp.
               }}
            |2=subgenus ''Anopheles''
            }}
         |2=subgenus ''[[Anopheles (Nyssorhynchus)|Nyssorhynchus]]'' (Americas)
         }}
      |2=[[Culicinae]]
      }}
   |2=[[Muscomorpha]]
   |sublabel2=''all other flies''
   }}
}}

=== Taxonomy ===

{{main|Taxonomy of Anopheles}}

The genus name ''Anopheles'' was introduced by the German entomologist [[Johann Wilhelm Meigen]] in 1818. He described two species, ''A. birfurcatus'' and the [[type species]], ''[[Anopheles maculipennis]]''. He stated that the name meant {{lang|de|beschwerlich}}, "burdensome".<ref name="Meigen 1818">{{cite book |last=Meigen |first=Johann Wilhelm |author-link=Johann Wilhelm Meigen |date=1818 |title=Systematische Beschreibung der bekannten Europäischen zweiflügeligen Insekten |language=de |trans-title=Systematic description of the known European two-winged insects |location=Aachen |publisher=Friedrich Wilhelm Forstmann |volume=1 |pages=10–12 |url=https://www.biodiversitylibrary.org/item/45833#page/54/mode/1up}}</ref> The name comes from the [[Ancient Greek]] word {{lang|grc|ἀνωφελής}} {{transliteration|grc|anōphelḗs}} 'useless', derived from {{lang|el|ἀν-}} {{transliteration|grc|an-}}, 'not', 'un-' and {{lang|el|[[:wikt:ὄφελος|ὄφελος]]}} {{transliteration|grc|óphelos}} 'profit'.<ref name="Stevenson2010">{{cite book |author=Stevenson, Angus |title=Oxford Dictionary of English |url=https://books.google.com/books?id=anecAQAAQBAJ&pg=PA64 |date=19 August 2010 |publisher=Oxford University Press |isbn=978-0-19-957112-3 |page=64}}</ref>

The taxonomy of the genus was greatly advanced in 1901 when the English entomologist [[Frederick Vincent Theobald]] described 39 ''Anopheles'' species in his 5-volume monograph on the Culicidae.<ref name=Theobald1901>{{cite book |last=Theobald |first=Frederick Vincent |author-link=Frederick Vincent Theobald |date=1901 |title=A Monograph of the Culicidae, or Mosquitoes |location=London |publisher=[[British Museum (Natural History)]] |volume=1 |isbn=978-1-178-51903-7 |pages=115–214 |url=https://archive.org/details/amonographculic03zoolgoog/page/n135/mode/2up}}</ref> He was provided with mosquito specimens sent in to the [[British Museum (Natural History)]] from around the world, on the 1898 instruction of the [[Secretary of State for the Colonies]], [[Joseph Chamberlain]].<ref name="Harbach Kitching 2016"/>

''Anopheles'' (with a nearly worldwide distribution) belongs to the subfamily [[Anophelinae]] alongside two other genera: ''[[Bironella]]'' (restricted to [[Australia]]) and ''[[Chagasia]]'' (restricted to the [[Neotropics]]). The taxonomy remains incompletely settled.<ref name="Krzywinski2003">{{cite journal |last1=Krzywinski |first1=Jaroslaw |last2=Besansky |first2=Nora J. |author-link2=Nora J. Besansky |year=2003 |title=Molecular Systematics of Anopheles: From Subgenera to Subpopulations |journal=Annual Review of Entomology |volume=48 |pages=111–139 |doi=10.1146/annurev.ento.48.091801.112647 |pmid=12208816}}</ref><ref name="Foley 1998">{{cite journal |doi=10.1006/mpev.1997.0457 |pmid=9562985 |title=Evolution and Systematics of ''Anopheles'':Insights from a Molecular Phylogeny of Australasian Mosquitoes |journal=Molecular Phylogenetics and Evolution |volume=9 |issue=2 |pages=262–275 |year=1998 |last1=Foley |first1=Desmond H. |last2=Bryan |first2=Joan H. |last3=Yeates |first3=David |last4=Saul |first4=Allan |bibcode=1998MolPE...9..262F }}</ref> Classification into species is based on morphological characteristics – wing spots, head anatomy, larval and pupal anatomy, chromosome structure, and more recently, on DNA sequences.<ref name="Rattanarithikul 2006">{{cite journal |pmid=17262930 |year=2006 |last1=Rattanarithikul |first1=R. |author-link=Rampa Rattanarithikul|title=Illustrated keys to the mosquitoes of Thailand. IV. Anopheles |journal=The Southeast Asian Journal of Tropical Medicine and Public Health |volume=37 |pages=1–128 |last2=Harrison |first2=B. A. |last3=Harbach |first3=R. E. |last4=Panthusiri |first4=P. |last5=Coleman |first5=R. E. |last6=Panthusiri |first6=P. |issue=Suppl 2 }}</ref><ref name="Walton 2007">{{cite journal |doi=10.1016/j.meegid.2006.05.001 |pmid=16782411 |title=Genetic diversity and molecular identification of mosquito species in the Anopheles maculatus group using the ITS2 region of rDNA |journal=Infection, Genetics and Evolution |volume=7 |issue=1 |pages=93–102 |year=2007 |last1=Walton |first1=C. |last2=Somboon |first2=P. |last3=o'Loughlin |first3=S. M. |last4=Zhang |first4=S. |last5=Harbach |first5=R.E. |last6=Linton |first6=Y.-M. |last7=Chen |first7=B. |last8=Nolan |first8=K. |last9=Duong |first9=S. |last10=Fong |first10=M.-Y |last11=Vythilingum |first11=I. |last12=Mohammed |first12=Z. D. |last13=Trung |first13=Ho Dinh |last14=Butlin |first14=R. K. |bibcode=2007InfGE...7...93W |display-authors=5 }}</ref><ref name="Garros Harbach Manguin 2005">{{cite journal |pmid=16119539 |year=2005 |last1=Garros |first1=C. |last2=Harbach |first2=R. E |last3=Manguin |first3=S. |title=Morphological assessment and molecular phylogenetics of the Funestus and Minimus groups of Anopheles (Cellia) |journal=Journal of Medical Entomology |volume=42 |issue=4 |pages=522–536 |doi=10.1093/jmedent/42.4.522 |s2cid=22776177 |doi-access=free }}</ref> In the taxonomy published by Harbach and Kitching in 2016, it was shown that three species of ''Bironella'' (''[[Bironella confusa|B. confusa]]'', ''[[Bironella gracilis|B. gracilis]],'' and ''[[Bironella hollandi|B. hollandi]]'') are phylogenetically more similar to ''[[Anopheles kyondawensis|A. kyondawensis]]'' than other ''Bironella'' species. That phylogeny argues that, based on genetic similarity, ''[[Anopheles implexus|A. implexus]]'' is divergent from the common ancestor of ''Anopheles''.<ref name="Harbach Kitching 2016">{{cite journal |last1=Harbach |first1=R. E. |last2=Kitching |first2=I. |date=January 2016 |title=The phylogeny of Anophelinae revisited: inferences about the origin and classification of ''Anopheles'' (Diptera: Culicidae) |journal=Zoologica Scripta |volume=45 |pages=34–47 |url=https://nhm.openrepository.com/handle/10141/612216 |doi=10.1111/zsc.12137 |hdl=10141/612216 |s2cid=46364692 |hdl-access=free}}</ref>