Editing Scorpion (section)

===Phylogeny===

The Scorpiones are a [[clade]] within the pulmonate [[Arachnida]] (those with book lungs). Arachnida is placed within the [[Chelicerata]], a subphylum of [[Arthropoda]] that contains [[sea spider]]s and [[horseshoe crab]]s, alongside terrestrial animals without book lungs such as [[tick]]s and [[harvestmen]].<ref name="Howard Edgecombe 2019"/> The extinct [[Eurypterida]], sometimes called sea scorpions, though they were not all marine, are not scorpions; their grasping pincers were [[chelicerae]], not [[Homology (biology)|homologous]] with the pincers (second appendages) of scorpions.<ref>{{cite web |last=Waggoner |first=B. M. |title=Eurypterida: Morphology |url=https://ucmp.berkeley.edu/arthropoda/chelicerata/eurypteridmm.html |publisher=University of California Museum of Paleontology Berkeley |access-date=20 October 2020 |date=12 October 1999 |archive-date=23 October 2020 |archive-url=https://web.archive.org/web/20201023013411/https://ucmp.berkeley.edu/arthropoda/chelicerata/eurypteridmm.html |url-status=live }}</ref> Scorpiones is [[Sister taxon|sister]] to the [[Tetrapulmonata]], a terrestrial group of pulmonates containing the [[spider]]s and whip scorpions. This 2019 [[cladogram]] summarizes:<ref name="Howard Edgecombe 2019"/>

{{clade
|label1=[[Chelicerata]]
|1={{clade
   |1=[[Pycnogonida]] (sea spiders) [[File:Nymphon signatum 194389384 (white background).jpg|90px]]
   |label2=[[Prosomapoda]]
   |2={{clade
      |1=[[Xiphosura]] (horseshoe crabs) [[File:Limulus polyphemus (aquarium) (white background).jpg|70px]]
      |2={{clade
         |1=†[[Eurypterida]] (sea scorpions) <span style="{{MirrorH}}">[[File:Eurypterus Paleoart (no background).png|70 px]]</span>
         |label2=[[Arachnida]]
         |2={{clade
            |label1=Non-pulmonates
            |1=([[ticks]], [[harvestmen]], etc) [[File:Ixodes scapularis P1170301a (white background).png|70px]]
            |label2=pulmonates
            |2={{clade
               |label1='''Scorpiones'''
               |1=[[File:Buthus_mariefranceae_(10.3897-zookeys.686.12206)_Figure_1.jpg|65px]]
               |label2=[[Tetrapulmonata]]
               |2={{clade
                  |1=[[Araneae]] (spiders) [[File:Aptostichus simus Monterey County.jpg|70px]]
                  |2=Pedipalpi ([[whip scorpions]], etc) [[File:Whip Scorpion body (9672115742) (white background).png|80px]]
                  }}
               }}
            }}
         }}
      }}
   }}
}}

Recent studies place [[pseudoscorpion]]s as the sister group of scorpions in the clade Panscorpiones, which together with Tetrapulmonata makes up the clade Arachnopulmonata.<ref>{{Cite journal|url=https://www.biorxiv.org/content/10.1101/2020.11.18.389098v1|title=Taxonomic sampling and rare genomic changes overcome long-branch attraction in the phylogenetic placement of pseudoscorpions|first1=Andrew Z.|last1=Ontano|first2=Guilherme|last2=Gainett|first3=Shlomi|last3=Aharon|first4=Jesús A.|last4=Ballesteros|first5=Ligia R.|last5=Benavides|first6=Kevin F.|last6=Corbett|first7=Efrat|last7=Gavish-Regev|first8=Mark S.|last8=Harvey|first9=Scott|last9=Monsma|first10=Carlos E.|last10=Santibáñez-López|first11=Emily V. W.|last11=Setton|first12=Jakob T.|last12=Zehms|first13=Jeanne A.|last13=Zeh|first14=David W.|last14=Zeh|first15=Prashant P.|last15=Sharma|journal=Molecular Biology and Evolution|volume=38|issue=6|date=June 2021|pages=2446–2467|doi=10.1093/molbev/msab038|pmid=33565584 |pmc=8136511 }}</ref>

The internal [[phylogeny]] of the scorpions has been debated,<ref name="Howard Edgecombe 2019">{{cite journal |last1=Howard |first1=Richard J. |last2=Edgecombe |first2=Gregory D. |last3=Legg |first3=David A. |last4=Pisani |first4=Davide |last5=Lozano-Fernandez |first5=Jesus |title=Exploring the Evolution and Terrestrialization of Scorpions (Arachnida: Scorpiones) with Rocks and Clocks |journal=Organisms Diversity & Evolution |volume=19 |issue=1 |year=2019 |pages=71–86 |issn=1439-6092 |doi=10.1007/s13127-019-00390-7 |doi-access=free|bibcode=2019ODivE..19...71H |hdl=10261/217081 |hdl-access=free }}</ref> but [[genomic]] analysis consistently places the [[Bothriuridae]] as sister to a clade consisting of Scorpionoidea and [[Chactoidea]]. The scorpions diversified between the Devonian and the early [[Carboniferous]]. The main division is into the clades Buthida and Iurida. The Bothriuridae diverged starting before temperate Gondwana broke up into separate land masses, completed by the [[Jurassic]]. The Iuroidea and Chactoidea are both seen not to be single clades, and are shown as "[[paraphyletic]]" (with quotation marks) in this 2018 cladogram.<ref name="Sharma Baker 2018">{{cite journal |last1=Sharma |first1=Prashant P. |last2=Baker |first2=Caitlin M. |last3=Cosgrove |first3=Julia G. |last4=Johnson |first4=Joanne E. |last5=Oberski |first5=Jill T. |last6=Raven |first6=Robert J. |last7=Harvey |first7=Mark S. |last8=Boyer |first8=Sarah L. |last9=Giribet |first9=Gonzalo |title=A Revised Dated Phylogeny of Scorpions: Phylogenomic Support for Ancient Divergence of the Temperate Gondwanan Family Bothriuridae |journal=Molecular Phylogenetics and Evolution |volume=122 |year=2018 |pages=37–45 |issn=1055-7903 |doi=10.1016/j.ympev.2018.01.003 |pmid=29366829 |doi-access=free |bibcode=2018MolPE.122...37S }}</ref>

{{clade
|label1='''Scorpiones'''
|1={{clade
   |label1=Buthida
   |1={{clade
      |1={{clade
         |1=[[Chaeriloidea]] <!--(53 spp.)--> [[File:Chaerilus pseudoconchiformus male (cropped).jpg|55px]]
         |2=[[Pseudochactoidea]] <!--(9 spp.)--> [[File:Vietbocap canhi (white background).jpg|45px]]
         }}
      |2=[[Buthoidea]] <!--(1209 spp.)--> [[File:Buthus mariefranceae (10.3897-zookeys.686.12206) Figure 1.jpg|65px]]
      }}
   |label2=Iurida<!--Parvorder-->
   |2={{clade
      |1="[[Iuroidea]]" (part) <!--14 spp.-->
      |2={{clade
         |1=[[Bothriuroidea]] <!--(158 spp.)--> [[File:Cercophonius squama.jpg|80px]]
         |2={{clade
            |1={{clade
               |1={{clade
                  |1="[[Chactoidea]]" (part)
                  |2="[[Iuroidea]]" (part)
                  }}
               |2="[[Chactoidea]]" (part)
               }}
            |2=[[Scorpionoidea]] <!--(183 spp.)--> <span style="{{MirrorH}}">[[File:Female Emperor Scorpion.jpg|75px]]</span>
            }}
         }}
      }}
   }}
}}