Editing Aphid (section)

==Evolution==
[[File:Vosegus.jpg|thumb|Forewing of the early [[Middle Triassic]] (early Anisian) aphid ''Vosegus triassicus''<ref>{{cite journal | last1 = Szwedo | first1 = J. | last2 = Nel | first2 = A. | year = 2011 | title = The oldest aphid insect from the Middle Triassic of the Vosges, France | journal = Acta Palaeontologica Polonica | volume = 56 | issue = 4| pages = 757–766 | doi = 10.4202/app.2010.0034 | doi-access = free }}</ref>]]
[[File:Baltic amber inclusions - Aphid (Hemiptera, Sternorrhyncha, Aphidoidea)7.JPG|thumb|upright|An aphid fossilised in [[Baltic amber]] ([[Eocene]])]]

===Fossil history===
Aphids, and the closely related [[Adelgidae|adelgids]] and [[Phylloxeridae|phylloxerans]], probably evolved from a [[Common descent|common ancestor]] some {{Ma |280}}, in the [[Cisuralian|Early Permian]] period.<ref name=Capinera2008/> They probably fed on plants like [[Cordaitales]] or [[Cycadophyta]]. With their soft bodies, aphids do not fossilize well, and the oldest known [[fossil]] is of the species ''[[Triassoaphis cubitus]]'' from the [[Triassic]].<ref>{{cite journal |author1=Johnson, Christine |author2=Agosti, Donat |author3=Delabie, Jocques H. |author4=Dumpert, Klaus |author5=Williams, D. J. |author6=von Tschirnhaus, Michael |author7=Macshwitz, Ulrich |year=2001 |title=''Acropyga'' and ''Azteca'' ants (Hymenoptera: Formicidae) with scale insects (Sternorrhyncha: Coccoidea): 20 million years of intimate symbiosis |journal=[[American Museum Novitates]] |issue=3335 |pages=1–18 |url=http://research.amnh.org/~cjohnson/Johnson_etal_2001_Novitates.pdf |doi=10.1206/0003-0082(2001)335<0001:AAAAHF>2.0.CO;2 |s2cid=55067700 |access-date=2010-10-18 |archive-date=2012-09-27 |archive-url=https://web.archive.org/web/20120927153952/http://research.amnh.org/~cjohnson/Johnson_etal_2001_Novitates.pdf |url-status=live }}</ref> They do however sometimes get stuck in plant exudates which solidify into [[amber]]. In 1967, when Professor [[Ole Engel Heie|Ole Heie]] wrote his monograph ''Studies on Fossil Aphids'', about sixty species had been described from the Triassic, [[Jurassic]], [[Cretaceous]] and mostly the [[Tertiary]] periods, with [[Baltic amber]] contributing another forty species.<ref>{{cite journal |author=Russell, Louise M. |year=1968 |title=Studies on Fossil Aphids |journal=Bulletin of the Entomological Society of America |volume=14 |issue=2 |pages=139–140 |doi=10.1093/besa/14.2.139a |url=https://academic.oup.com/ae/article-pdf/14/2/139/18806083/besa14-0139a.pdf |access-date=2018-02-04 }}</ref> The total number of species was small, but increased considerably with the appearance of the [[angiosperm]]s {{Ma |160}}, as this allowed aphids to specialise, the speciation of aphids going hand-in-hand with the diversification of flowering plants. The earliest aphids were probably [[polyphagous]], with [[monophagy]] developing later.<ref name="Dixon"/> It has been hypothesized that the ancestors of the [[Adelgidae]] lived on [[conifer]]s while those of the Aphididae fed on the sap of [[Podocarpaceae]] or [[Araucariaceae]] that survived extinctions in the late Cretaceous. Organs like the cornicles did not appear until the Cretaceous period.<ref name="Capinera2008">{{cite book |author=Capinera, John L. |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA193 |year=2008 |publisher=Springer Science & Business Media |isbn=978-1-4020-6242-1 |pages=193–194 |access-date=2018-02-04 |archive-date=2020-08-05 |archive-url=https://web.archive.org/web/20200805231221/https://books.google.com/books?id=i9ITMiiohVQC&pg=PA193 |url-status=live }}</ref><ref>{{Cite journal |last1=Von Dohlen |first1=Carol D. |last2=Moran |first2=Nancy A. |year=2000 |title=Molecular data support a rapid radiation of aphids in the Cretaceous and multiple origins of host alternation|journal=Biological Journal of the Linnean Society |volume=71 |issue=4 |pages=689–717 |doi=10.1111/j.1095-8312.2000.tb01286.x|doi-access=free }}</ref> One study alternatively suggests that ancestral aphids may have lived on angiosperm bark and that feeding on leaves may be a [[derived trait]]. The [[Lachninae]] have long mouth parts that are suitable for living on bark and it has been suggested that the mid-Cretaceous ancestor fed on the bark of angiosperm trees, switching to leaves of conifer hosts in the late Cretaceous.<ref name="Chen Favret Jiang Wang pp. 555–572">{{cite journal | last1=Chen | first1=Rui | last2=Favret | first2=Colin | last3=Jiang | first3=Liyun | last4=Wang | first4=Zhe | last5=Qiao | first5=Gexia | title=An aphid lineage maintains a bark-feeding niche while switching to and diversifying on conifers | journal=Cladistics | volume=32 | issue=5 | date=29 September 2015 | doi=10.1111/cla.12141 | pages=555–572| pmid=34740301 | s2cid=86517289 | doi-access=free }}</ref> The Phylloxeridae may well be the oldest family still extant, but their fossil record is limited to the [[Lower Miocene]] ''[[Palaeophylloxera]]''.<ref>{{cite encyclopedia |author=Gullan, Penny J. |author2=Martin, Jon H. |title=Sternorrhyncha |encyclopedia=Encyclopedia of Insects |edition=2nd |year=2009}}</ref>

===Taxonomy===
Late 20th-century reclassification within the Hemiptera reduced the old taxon "Homoptera" to two suborders: [[Sternorrhyncha]] (aphids, whiteflies, [[scale insect|scales]], [[Psylloidea|psyllids]], etc.) and [[Auchenorrhyncha]] ([[cicada]]s, [[leafhopper]]s, [[treehopper]]s, [[planthopper]]s, etc.) with the suborder [[Heteroptera]] containing a large group of insects known as the [[true bugs]]. The infraorder Aphidomorpha within the Sternorrhyncha varies with circumscription with several fossil groups being especially difficult to place but includes the Adelgoidea, the Aphidoidea and the Phylloxeroidea.<ref>{{cite book |url=https://archive.org/stream/fundamentalsofpa09rohd#page/267/mode/1up|pages=267–274 |title=Fundamentals of Paleontology. Volume 9. Arthropoda, Tracheata, Chelicerata |editor=Rohdendorf, B. B. |year=1991 |publisher=[[Smithsonian Institution]] and [[National Science Foundation]]}}</ref> Some authors use a single superfamily Aphidoidea within which the [[Phylloxeridae]] and Adelgidae are also included while others have Aphidoidea with a sister superfamily Phylloxeroidea within which the Adelgidae and Phylloxeridae are placed.<ref>{{cite book |pages=27–31 |title=Encyclopedia of Insects |chapter=Aphids |editor1=Resh, Vincent H.|editor2=Cardé, R. T. |edition=2 |publisher=Academic Press |year=2009 |author=Sorensen, J. T.}}</ref> Early 21st-century reclassifications substantially rearranged the families within Aphidoidea: some old families were reduced to subfamily rank (''e.g.'', [[Eriosomatidae]]), and many old subfamilies were elevated to family rank. The most recent authoritative classifications have three superfamilies Adelgoidea, Phylloxeroidea and Aphidoidea. The Aphidoidea includes a single large family [[Aphididae]] that includes all the ~5000<ref name="Zyla2017"/> extant species.<ref name="ASF">{{cite web |url=http://aphid.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1159459 |title=Superfamily Aphidoidea, Latreille, 1802 |publisher=Aphid: species file |access-date=3 February 2018 |archive-date=6 November 2017 |archive-url=https://web.archive.org/web/20171106005419/http://aphid.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1159459 |url-status=live }}</ref>

=== Phylogeny ===

==== External ====

Aphids, adelgids, and [[phylloxera|phylloxerids]] are very closely related within the suborder Sternorrhyncha, the plant-sucking bugs. They are either placed in the insect superfamily Aphidoidea<ref>{{cite book |author1=Blackman, R. L. |author2=Eastrop, V. F. |year=1994 |title=Aphids on the World's Trees. An Identification and Information Guide |publisher=[[CAB International]] |location=[[Wallingford, Oxfordshire|Wallingford]] |isbn=978-0-85198-877-1}}</ref> or into the superfamily [[Phylloxeroidea]] which contains the family Adelgidae and the family Phylloxeridae.<ref name="Dixon" /> Like aphids, phylloxera feed on the roots, leaves, and shoots of grape plants, but unlike aphids, do not produce [[honeydew (secretion)|honeydew]] or [[cornicle]] secretions.<ref>{{cite journal |author1=Granett, Jeffrey |author2=Walker, M. Andrew |author3=Kocsis, Laszlo |author4=Omer, Amir D. |year=2001 |title=Biology and management of grape phylloxera |journal= Annual Review of Entomology |volume=46 |pages=387–412 |doi=10.1146/annurev.ento.46.1.387|pmid=11112174 }}</ref> Phylloxera (''Daktulosphaira vitifoliae'') are insects which caused the [[Great French Wine Blight]] that devastated European [[viticulture]] in the 19th century. Similarly, adelgids or woolly conifer aphids, also feed on plant phloem and are sometimes described as aphids, but are more properly classified as aphid-like insects, because they have no cauda or cornicles.<ref name="McGavin">{{cite book |title=Bugs of the World |author=McGavin, George C. |year=1993 |isbn=978-0-8160-2737-8 |publisher=[[Infobase Publishing]]}}</ref>

The treatment of the groups especially concerning fossil groups varies greatly due to difficulties in resolving relationships. Most modern treatments include the three superfamilies, the Adelogidea, the Aphidoidea, and the Phylloxeroidea within the infraorder Aphidomorpha along with several fossil groups.<ref name="Favret Eades 2020">{{cite web |url=http://aphid.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1159014 |title=Aphid Species File - Aphidomorpha |last1=Favret |first1=C. |last2=Eades |first2=D.C. |date=2020 |editor-last=Miller |editor-first=G.L. |editor2-last=Qiao |editor2-first=G. |editor3-last=Sano |editor3-first=Masakazu |editor4-last=Stekolshchikov |editor4-first=A.V. |department=Université de Montréal |website=aphid.speciesfile.org |publisher=Aphid Species File |location=Montreal, Canada |access-date=December 13, 2020 |archive-date=February 24, 2021 |archive-url=https://web.archive.org/web/20210224122450/http://aphid.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1159014 |url-status=live }}</ref>

{{clade
|label1=[[Sternorrhyncha]]
|1={{clade
   |1=[[Psylloidea]] (jumping plant lice, etc) [[File:Psyllia_pyricola.png|125px]]
   |2={{clade
      |1=[[Aleyrodoidea]] (whiteflies) [[File:Neomaskellia bergii from CSIRO.jpg|90px]]
      |2={{clade
         |1=[[Coccoidea]] (scale insects) [[File:Ceroplastes_ceriferus_from_CSIRO.jpg|90px]]
         |label2=[[Aphidomorpha]]
         |2={{clade          
            |1={{clade
               |label1=[[Phylloxeroidea]]
               |1={{clade
                  |1=[[Phylloxeridae]] (phylloxerans) [[File:Daktulosphaira vitifoliae from CSIRO.jpg|60px]] 
                  |2=[[Adelgidae]] (woolly conifer aphids) [[File:Chermes pinifoliae.png|110px]]  
                  }}
               |label2='''Aphidoidea'''
               |2='''[[Aphididae]]''' (aphids) [[File:Aphid_icon.png|90px]]
               }}
            }}
         }}
      }}
   }}
}}

====Internal====
The phylogenetic tree, based on Papasotiropoulos 2013 and Kim 2011, with additions from Ortiz-Rivas and Martinez-Torres 2009, shows the internal [[phylogeny]] of the Aphididae.<ref>{{cite journal |author=Papasotiropoulos, Vassilis |author2=Tsiamis, Georgios |author3=Papaioannou, Charikleia |author4=Kilias, George |title=A molecular phylogenetic study of aphids (Hemiptera: Aphididae) based on mitochondrial DNA sequence analysis |journal=Journal of Biological Research-Thessaloniki |year=2013 |volume=20 |pages=1–13 |url=https://www.researchgate.net/publication/249007487 |access-date=2018-01-15 |archive-date=2018-01-16 |archive-url=https://web.archive.org/web/20180116135143/https://www.researchgate.net/publication/249007487 |url-status=live }}</ref><ref name="Kim2011">{{cite journal | last1=Kim | first1=Hyojoong | last2=Lee | first2=Seunghwan | last3=Jang | first3=Yikweon | editor-last=Moreau | editor-first=Corrie S. | title=Macroevolutionary Patterns in the Aphidini Aphids (Hemiptera: Aphididae): Diversification, Host Association, and Biogeographic Origins | journal=PLOS ONE | volume=6 | issue=9 | date=September 2011 | doi=10.1371/journal.pone.0024749 | pmid=21935453 | page=e24749| bibcode=2011PLoSO...624749K | pmc=3174202 | doi-access=free }}</ref><ref name="Ortiz-Rivas Martínez-Torres 2010">{{cite journal | last1=Ortiz-Rivas | first1=Benjamín | last2=Martínez-Torres | first2=David | title=Combination of molecular data support the existence of three main lineages in the phylogeny of aphids (Hemiptera: Aphididae) and the basal position of the subfamily Lachninae | journal=Molecular Phylogenetics and Evolution | volume=55 | issue=1 | year=2010 | doi=10.1016/j.ympev.2009.12.005 | pmid=20004730 | pages=305–317| bibcode=2010MolPE..55..305O }}</ref>

It has been suggested that the phylogeny of the aphid groups might be revealed by examining the phylogeny of their bacterial [[endosymbiont]]s, especially the obligate endosymbiont ''[[Buchnera (bacterium)|Buchnera]]''. The results depend on the assumption that the symbionts are strictly [[Vertical transmission (symbiont)|transmitted vertically]] through the generations. This assumption is well supported by the evidence, and several phylogenetic relationships have been suggested on the basis of endosymbiont studies.<ref>{{cite journal |doi=10.1554/0014-3820(2000)054[0517:CBBEBA]2.0.CO;2 |pmid=10937228 |year=2000 |volume=54 |issue=2 |pages=517–25 |title=Cospeciation Between Bacterial Endosymbionts (Buchnera) and a Recent Radiation of Aphids (Uroleucon) and Pitfalls of Testing for Phylogenetic Congruence |journal=Evolution |last1=Clark |first1=Marta A|last2=Moran |first2=Nancy A. |last3=Baumann |first3=Paul |last4=Wernegreen |first4=Jennifer J.}}</ref><ref>{{cite journal |doi=10.1016/j.ympev.2013.03.016 |pmid=23542003 |title=Reconstructing the phylogeny of aphids (Hemiptera: Aphididae) using DNA of the obligate symbiont Buchnera aphidicola |journal=Molecular Phylogenetics and Evolution |volume=68 |issue=1 |pages=42–54 |year=2013 |last1=Nováková |first1=Eva |last2=Hypša |first2=Václav |last3=Klein |first3=Joanne |last4=Foottit |first4=Robert G |last5=von Dohlen |first5=Carol D. |last6=Moran |first6=Nancy A. |bibcode=2013MolPE..68...42N |url=http://www.canacoll.org/Hemip/Staff/Foottit/PDFs/Novakova_et_al_2013/Novakova_et_al_2013.pdf |access-date=2018-04-27 |archive-date=2017-08-10 |archive-url=https://web.archive.org/web/20170810051059/http://www.canacoll.org/Hemip/Staff/Foottit/PDFs/Novakova_et_al_2013/Novakova_et_al_2013.pdf |url-status=live }}</ref><ref>{{cite journal |doi=10.1038/s41598-017-10761-9 |pmid=28860659 |pmc=5579299 |title=Insect-bacteria parallel evolution in multiple-co-obligate-aphid association: A case in Lachninae (Hemiptera: Aphididae) |journal=Scientific Reports |volume=7 |issue=1 |pages=10204 |year=2017 |last1=Chen |first1=Rui |last2=Wang |first2=Zhe |last3=Chen |first3=Jing |last4=Jiang |first4=Li-Yun |last5=Qiao |first5=Ge-Xia|bibcode=2017NatSR...710204C }}</ref>

{{clade
|label1='''Aphididae'''
|1={{clade
   |1=[[Lachninae]] [[File:Schizolachnus sp 20120528.JPG|100px]]
   |2={{clade
      |1=[[Hormaphidinae]] [[File:Witch Hazel Cone Gall Aphid - Flickr - treegrow (2).jpg|80px]]
      |2={{clade
         |1=[[Calaphidinae]] [[File:Calaphis.flava.-.lindsey.jpg|100px]]
         |2={{clade
            |1=[[Chaitophorinae]] [[File:Periphyllus sp. aphids on sycamore (cropped).jpg|80px]]
            |2={{clade
               |1=[[Eriosomatinae]] (woolly aphids) [[File:Eriosoma ulmi (European Elm Leafcurl Aphid).jpg|80px]]
               |2=[[Anoeciinae]] [[File:Anoecia corni1.jpg|80px]]
               }}
            |3={{clade
               |1=''[[Capitophorus]]'', ''[[Pterocomma]]''
               |label2=[[Aphidinae]]
               |2={{clade
                  |1=[[Macrosiphini]] [[File:Macrosiphum rosae (alate i.e. winged form) on a rose bud.jpg|70px]]
                  |label2=[[Aphidini]]
                  |2={{clade
                     |1=[[Rhopalosiphina]] [[File:Snodgrass Rhopalosiphum prunifoliae (wings closed).jpg|100px]]
                     |2=[[Aphidina]] (''[[Aphis]]'' spp) [[File:Aphids May 2010-3.jpg|60px]]
                     }}
                  }}
               }}
            }} 
         }}
      }}
   }}
}}

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