Editing Scale insect (section)

==Evolution==

The containing group of the scale insects was formerly treated as the superfamily Coccoidea but taxonomic uncertainties have led workers to prefer the use of the infraorder Coccomorpha as the preferred name for the group.<ref>{{cite journal |last1=Williams |first1=Douglas J. |last2=Hodgson |first2=Chris J. |date=2014 |title=The case for using the infraorder Coccomorpha above the superfamily Coccoidea for the scale insects (Hemiptera: Sternorrhyncha) |url=http://biotaxa.org/Zootaxa/article/view/zootaxa.3869.3.9 |journal=Zootaxa |volume=3869 |issue=3 |pages=348–350 |doi=10.11646/zootaxa.3869.3.9 |pmid=25283922 |doi-access=free }}</ref> Scale insects are members of the [[Sternorrhyncha]]. The [[phylogeny]] of the extant groups, inferred from analysis of small subunit (18S) [[ribosomal RNA]], is shown in the first [[cladogram]].{{Cn|date=May 2023}}

{{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='''Coccomorpha''' (scale insects) [[File:Ceroplastes_ceriferus_from_CSIRO.jpg|90px]]
         |label2=[[Aphidomorpha]]
         |2={{clade
            |1=[[Phylloxeroidea]] (phylloxera bugs) [[File:Daktulosphaira vitifoliae from CSIRO.jpg|60px]]
            |2=[[Aphididae]] (aphids) [[File:Aphid_winged.jpg|90px]]
            }}
         }}
      }}
   }}
}}

[[File:Acropyga glaesaria SMFBE457B2 01.jpg|thumb|Fossil of the [[Pseudococcidae|pseudococcid]] mealybug ''[[Electromyrmococcus]]'' (in the jaws of an ant) in [[Miocene]] [[Dominican amber]]<ref name="Poinar2011">{{cite journal |last1=Poinar |first1=G. |last2=Heiss |first2=E. |year=2011 |title=New Termitaphididae and Aradidae (Hemiptera) in Mexican and Dominican amber |journal=Palaeodiversity |volume=4 |pages=51–62 |url=http://www.palaeodiversity.org/pdf/04/Palaeodiversity_4_PoinarHeiss.pdf}}</ref>]]

The timing of phylogenetic diversification within the Coccomorpha was estimated in a 2016 study based on [[molecular clock]] divergence time estimates, along with fossils being used for calibration. They suggested that the main scale insect lineages diverged before their angiosperm hosts, and suggested that the insects switched from feeding on gymnosperms once the angiosperms became common and widespread in the Cretaceous. They estimated that the Coccomorpha appeared at the start of the [[Triassic]] period, around 245 million years ago, and that the neococcoids appeared during the Early Jurassic, some 185 million years ago.<ref name="VeaGrimaldi2016" /> Scale insects are very well represented in the fossil record, with the oldest known member of the group reported from the [[Late Jurassic]] [[amber]] from Lebanon.<ref name=Jankotejacoccidae>{{Cite journal|last1=Vršanský |first1=P. |last2=Sendi |first2=H. |last3=Kotulová |first3=J. |last4=Szwedo |first4=J. |last5=Havelcová |first5=M. |last6=Palková |first6=H. |last7=Vršanská |first7=L. |last8=Sakala |first8=J. |last9=Puškelová |first9=L. |last10=Golej |first10=M. |last11=Biroň |first11=A. |last12=Peyrot |first12=D. |last13=Quicke |first13=D. |last14=Néraudeau |first14=D. |last15=Uher |first15=P. |last16=Maksoud |first16=S. |last17=Azar |first17=D. |title=Jurassic Park approached: a coccid from Kimmeridgian cheirolepidiacean Aintourine Lebanese amber |year=2024 |journal=National Science Review |at=nwae200 |doi=10.1093/nsr/nwae200 |doi-access=free |pmc=11895504 }}</ref> They are abundantly preserved in amber from the Early Cretaceous, 130 mya, onwards; they were already highly diversified by Cretaceous times. All the families were monophyletic except for the [[Eriococcidae]]. The Coccomorpha are division into two clades the "Archaeococcoids" and "Neococcoids". The archaeococcoid  families have adult males with either compound eyes or a row of unicorneal eyes and have abdominal spiracles in the females. In neoccoids, the females have no abdominal spiracles.<ref>{{cite journal |last1=Williams |first1=D. J. |last2=Gullan |first2=P. J. |last3=Miller |first3=D. R. |last4=Matile-Ferrero |first4=D. |last5=Han |first5=Sarah I. |date=2011 |title=A study of the scale insect genera Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea: Putoidae) and Ceroputo Šulc (Pseudococcidae) with a comparison to Phenacoccus Cockerell (Pseudococcidae) |url=https://biotaxa.org/Zootaxa/article/view/zootaxa.2802.1.1 |journal=Zootaxa |volume=2802 |issue=1 |pages=1 |doi=10.11646/zootaxa.2802.1.1 |hdl=1885/63136 |hdl-access=free }}</ref> In the cladogram below the genus ''Pityococcus'' is moved to the "Neococcoids". A cladogram showing the major families using this methodology is shown below.<ref name="VeaGrimaldi2016">{{cite journal |last1=Vea |first1=Isabelle M. |last2=Grimaldi |first2=David A. |title=Putting scales into evolutionary time: the divergence of major scale insect lineages (Hemiptera) predates the radiation of modern angiosperm hosts |journal=Scientific Reports |volume=6 |issue=1 |pages=23487 |year=2016 |issn=2045-2322 |doi=10.1038/srep23487|pmid=27000526 |pmc=4802209 |bibcode=2016NatSR...623487V }}</ref>

{{clade
|label1='''Coccomorpha'''
|1={{clade
|label1="Archaeococcoids"
|sublabel1=- ''Pityococcus''
|1={{clade
   |1={{clade
      |1=†[[Burmacoccidae]]
      |2=†[[Kozariidae]]
      }}
   |2={{clade
      |1={{clade
         |1=[[Matsucoccidae]] (pine bast scales)
         |2=[[Ortheziidae]] (ensign scales)
         }}
      |2={{clade
         |1=[[Margarodidae]] (ground pearls)
         |2={{clade
            |1=[[Kuwaniidae]]
            |2={{clade
               |1=[[Xylococcidae]]
               |2={{clade
                  |1=[[Coelostomidiidae]]
                  |2=[[Monophlebidae]] (cottony cushion scales)
                  }}
               }}
            }}
         }}
      }}
   }}
|label2="Neococcoids" 
|sublabel2=+ ''Pityococcus''
|2={{clade
   |1=[[Pityococcidae]]
   |2={{clade
      |1=[[Steingeliidae]]
      |2={{clade
         |1={{clade
            |1=[[Phenacoleachiidae]]
            |2=[[Putoidae]] (giant mealybugs)
            }}
         |2={{clade
            |1=[[Pseudococcidae]] (mealybugs)
            |2={{clade
               |1={{clade
                  |1=[[Coccidae]] (soft scales)
                  |2={{clade
                     |1=[[Kermesidae]] (kermes dye scales)
                     |2={{clade
                        |1=[[Asterolecaniidae]] (pit scales)
                        |2=[[Kerriidae]] (lac scales)
                        }}
                     }}
                  }}
               |2={{clade
                  |1={{clade
                     |1=[[Dactylopiidae]] (cochineal insects)
                     |2=''[[Palaearctic]]'' "[[Eriococcidae]]" (felted scales)
                     }}
                  |2={{clade
                     |1=[[Beesoniidae]], [[Stictococcidae]], part of "[[Eriococcidae]]"
                     |2={{clade
                        |1=[[Phoenicococcidae]] (palm scales)
                        |2=[[Diaspididae]] (armoured scales)
                        }}
                     }}
                  }}
               }}
            }}
         }}
      }}
   }}
}}
}}

Recognition of scale insect families has fluctuated over time, and the validity of many remains in flux,<ref>{{Cite journal |last1=Gullan |first1=P. J. |last2=Cook |first2=L. G. |year=2007 |title=Phylogeny and higher classification of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea) |journal=Zootaxa |volume=1668 |pages=413–425 |url=http://www.mapress.com/zootaxa/list/2007/Linnaeus.html |doi=10.11646/zootaxa.1668.1.22 |url-access=subscription }}</ref><ref name="Hodgson-2013">{{cite journal |last1=Hodgson |first1=Chris J. |last2=Hardy |first2=Nate B. |year=2013 |title=The phylogeny of the superfamily Coccoidea (Hemiptera: Sternorrhyncha) based on the morphology of extant and extinct macropterous males |journal=Systematic Entomology |volume=38 |issue=4 |pages=794–804 |doi=10.1111/syen.12030 |doi-access=free |bibcode=2013SysEn..38..794H }}</ref> with several recognized families not included in the phylogeny presented above including extinct groups are listed below:<ref>{{cite journal |last=Szwedo |first=J. |year=2018 |title=The unity, diversity and conformity of bugs (Hemiptera) through time |journal=Earth and Environmental Science Transactions of the Royal Society of Edinburgh |volume=107 |issue=2–3 |pages=109–128 |doi=10.1017/s175569101700038x |s2cid=134243346 }}</ref><ref>{{cite web |last1=Ben-Dov |first1=Y. |last2=Miller |first2=D. R. |last3=Gibson |first3=G.A.P. |work=ScaleNet |url=http://www.sel.barc.usda.gov/scalenet/scalenet.htm |title=Home |access-date=2013-04-04 |url-status=dead |archive-url=https://web.archive.org/web/20130404165159/http://www.sel.barc.usda.gov/scalenet/scalenet.htm |archive-date=2013-04-04}}</ref><ref name="johnsonetal1995">{{cite journal |year=2001 |issue=335 |pages=1–18 |title=''Acropyga'' and ''Azteca'' ants (Hymenoptera: Formicidae) with scale insects (Sternorrhyncha: Coccoidea): 20 million years of intimate symbiosis |last1=Johnson |first1=Christine |last2=Agosti |first2=Donat |last3=Delabie |first3=Jacques H. |last4=Dumpert |first4=Klaus |last5=Williams |first5=D. J. |last6=von Tschirnhaus |first6=Michael |last7=Maschwitz |first7=Ulrich |display-authors=3 |journal=American Museum Novitates |doi=10.1206/0003-0082(2001)335<0001:AAAAHF>2.0.CO;2 |s2cid=55067700 |url=https://zenodo.org/record/5370690 }}</ref>

* Archecoccoidea {{small|Borchsenius, 1958}}
** {{extinct}}[[Apticoccidae]] {{small|Vea & Grimaldi, 2015}}
** {{extinct}}[[Arnoldidae]] {{small|Koteja, 2008}}
** {{extinct}}[[Burmacoccidae]] {{small|Koteja, 2004}}
** [[Callipappidae]] {{small|MacGillivray, 1921}}
** [[Coelostomidiidae]] {{small|Morrison, 1927}}
** {{extinct}}[[Electrococcidae]] {{small|Koteja, 2000}}
** {{extinct}}[[Grimaldiellidae]] {{small|Koteja, 2000}}
** {{extinct}}[[Grohnidae]] {{small|Koteja, 2008}}
** {{extinct}}[[Hammanococcidae]] {{small|Koteja & Azar, 2008}}
** {{extinct}}[[Jankotejacoccidae]] {{small|Szwedo, Azar & Sendi, 2024}}<ref name=Jankotejacoccidae />
** {{extinct}}[[Jersicoccidae]] {{small|Koteja, 2000}}
** {{extinct}}[[Kozariidae]] {{small|Vea & Grimaldi, 2015}}
** {{extinct}}[[Kukaspididae]] {{small|Koteja & Poinar, 2001}}
** [[Kuwaniidae]] {{small|MacGillivray, 1921}}
** {{extinct}}[[Labiococcidae]] {{small|Koteja, 2000b}}
** {{extinct}}[[Lebanococcidae]] {{small|Koteja & Azar, 2008}}
** [[Lithuanicoccidae]] {{small|Koteja, 2008}}
** [[Macrodrilidae]] {{small|Poinar, 2020}}<ref name="Macrodrilus2020">{{cite journal |last1=Poinar |first1=G. O. Jr |year=2020 |title=Macrodrilidae fam. nov. (Hemiptera: Sternorrhyncha: Coccoidea), a new family of scale insects in mid-Cretaceous Burmese amber |journal=Historical Biology: An International Journal of Paleobiology |volume=33 |issue=9 |pages= 1726–1730|doi=10.1080/08912963.2020.1733549 |s2cid=216480240 }}</ref>
** [[Marchalinidae]] {{small|Morrison, 1927}}
** [[Margarodidae]] {{small|Cockerell, 1899}}
** [[Matsucoccidae]] {{small|Morrison, 1927}}
** [[Monophlebidae]] {{small|Morrison, 1927}}
** [[Ortheziidae]] {{small|Amyot & Audinet-Serville, 1843}}
** {{extinct}}[[Pennygullaniidae]] {{small|Koteja & Azar, 2008}}
** [[Phenacoleachiidae]] {{small|Cockerell, 1902}}
** [[Pityococcidae]] {{small|McKenzie, 1942}}
** [[Putoidae]] {{small|Tang, 1992}}
** {{extinct}}[[Serafinidae]] {{small|Koteja, 2008}}
** [[Steingeliidae]] {{small|Morrison, 1927}}
** [[Stigmacoccidae]] {{small|Morrison, 1927}}
** [[Termitococcidae]] {{small|Jakubski, 1965}}
** {{extinct}}[[Weitschatidae]] {{small|Koteja, 2008}}
** [[Xylococcidae]] {{small|Pergande in Hubbard & Pergande, 1898}}
* Neococcoidea {{small|Borchsenius, 1950}}
** [[Aclerdidae]] {{small|Cockerell, 1905}}
** {{extinct}}[[Albicoccidae]] {{small|Koteja, 2004}}
** [[Asterolecaniidae]] {{small|Cockerell, 1896}}
** [[Beesoniidae]] {{small|Ferris, 1950}}
** [[Calycicoccidae]] {{small|Brain, 1918}}
** [[Carayonemidae]] {{small|Richard, 1986}}
** [[Cerococcidae]] {{small|Balachowsky, 1942}}
** [[Cissococcidae]] {{small|Brain, 1918}}
** [[Coccidae]] {{small|Fallen, 1814}}
** [[Conchaspididae]] {{small|Green, 1896}}
** [[Cryptococcidae]] {{small|Kosztarab, 1968}}
** [[Dactylopiidae]] {{small|Signoret, 1875}}
** [[Diaspididae]] {{small|Targioni-Tozzetti, 1868}}
** [[Eriococcoidae]] {{small|Cockerell, 1899}}
** [[Halimococcidae]] {{small|Brown & McKenzie, 1962}}
** {{extinct}}[[Hodgsonicoccidae]] {{small|Vea & Grimaldi, 2015}}
** {{extinct}}[[Inkaidae]] {{small|Koteja, 1989}}
** [[Kermesidae]] {{small|Signoret, 1875}}
** [[Kerriidae]] {{small|Lindinger, 1937}}
** [[Lecanodiaspididae]] {{small|Targioni-Tozzetti, 1869}}
** [[Micrococcidae]] {{small|Silvestri, 1939}}
** [[Phoenicococcidae]] {{small|Stickney, 1934}}
** [[Porphyrophoridae]] {{small|Signoret, 1875}}
** [[Pseudococcidae]] {{small|Cockerell, 1905}}
** [[Rhizoecidae]] {{small|Williams, 1969}}
** [[Stictococcidae]] {{small|Lindinger, 1913}}
** [[Tachardiidae]] {{small|Green, 1896}}