# Megacheira > Mediated Wiki article. Canonical URL: https://mediated.wiki/source/Megacheira > Markdown URL: https://mediated.wiki/source/Megacheira.md > Source: https://en.wikipedia.org/wiki/Megacheira > Source revision: 1356413720 > License: Creative Commons Attribution-ShareAlike 4.0 International (https://creativecommons.org/licenses/by-sa/4.0/) {{Short description|Extinct class of arthropods}} {{Use dmy dates|date=May 2024}} {{Automatic taxobox | fossil_range = Middle [Cambrian](/source/Cambrian)-Late [Ordovician](/source/Ordovician) {{Fossilrange|Cambrian Stage 3|Katian|latest=Early Devonian}} Possible --[Silurian](/source/Silurian) and [Devonian](/source/Devonian) records | image = 20191028 Megacheirans Leanchoilia Haikoucaris Yohoia Fortiforceps.png | image_upright = 1.3 | image_caption = ''[Alalcomenaeus](/source/Alalcomenaeus)'' (top left), ''[Fortiforceps](/source/Fortiforceps)'' (top right), ''[Haikoucaris](/source/Haikoucaris)'' (middle), ''[Leanchoilia](/source/Leanchoilia)'' (bottom left) and ''[Yohoia](/source/Yohoia)'' (bottom right). | taxon = Megacheira | authority = [Hou](/source/Hou_Xian-guang) and Bergström, 1997 | subdivision_ranks = Groups | subdivision = See text }} '''Megacheira''' (from [Ancient Greek](/source/Ancient_Greek) μέγας (''mégas''), meaning "great", and χείρ (''kheír''), meaning "hand", also historically '''great appendage arthropods''') is an extinct class of early [Paleozoic](/source/Paleozoic) predatory [arthropod](/source/arthropod)s defined by their possession of spined "great appendages".{{Cite journal| doi = 10.1111/j.1096-3642.2009.00562.x| title = A new arthropod from the Early Cambrian of North Greenland, with a 'great appendage'-like antennula| date = March 2010| orig-date = 26 February 2010 | last1 = Stein | first1 = Martin| journal = [Zoological Journal of the Linnean Society](/source/Zoological_Journal_of_the_Linnean_Society)| volume = 158| issue = 3| pages = 477–500}} Their taxonomic position is controversial, with studies generally either considering them [stem-group](/source/stem-group) euarthropods, or stem-group [chelicerates](/source/Chelicerata).{{Cite journal |last=Aria |first=Cédric |date=26 April 2022 |title=The origin and early evolution of arthropods |url=http://paleorxiv.org/4zmey/download |journal=Biological Reviews |volume=97 |issue=5 |language=en |pages=1786–1809 |doi=10.1111/brv.12864 |pmid=35475316 |s2cid=243269510 |issn=1464-7931 |access-date=8 May 2024 |archive-date=8 May 2024 |archive-url=https://web.archive.org/web/20240508131708/http://paleorxiv.org/4zmey/download |url-status=live }}{{cite journal |last1=Liu |first1=Cong |last2=Fu |first2=Dongjing |last3=Wu |first3=Yu |last4=Zhang |first4=Xingliang |title=Cambrian euarthropod Urokodia aequalis sheds light on the origin of Artiopoda body plan |journal=iScience |date=July 2024 |volume=27 |issue=8 |article-number=110443 |doi=10.1016/j.isci.2024.110443|doi-access=free |pmid=39148713 |pmc=11325232 |bibcode=2024iSci...27k0443L }} The [homology](/source/Homology_(biology)) of the great appendages to the cephalic appendages of other arthropods is also controversial. Uncontested members of the group were present in marine environments worldwide from the lower [Cambrian](/source/Cambrian) to the upper [Ordovician](/source/Ordovician). == Morphology == {{multiple image | align = center | perrow = | image1 = 20190911 Megacheira Great Appendages.png | caption1 = Comparison of megacheiran great appendages | image2 = 20210701 Yohoia tenuis great appendage mobility.gif | caption2 = Movement range of the great appendage of ''[Yohoia](/source/Yohoia)'' | total_width = 525 | direction = horizontal }} [[File:Tanglangia longicaudata reconstruction.png|thumb|3D model of ''[Tanglangia longicaudata](/source/Tanglangia).'' en=endopod, ex=exopod]] Megacheirans are defined by their possession of [uniramous](/source/uniramous) (unbranched) "great appendages", which are their first pair of head appendages. The first one or two proximalmost segments/[podomere](/source/podomere)s closest towards the body are spineless (it has been argued that the supposed first of the two proximal podomeres is actually an arthrodial membrane), while the remaining 3–4 more distal podomeres towards the ends of the limbs each typically bear a single upward pointing spine attached towards the [distal](/source/distal) (towards the tip) end of the segment, with the spineless proximal segment/s typically being connected to the spined distal segments by an elbow-like joint, which curled upwards.{{Cite journal |last1=Haug |first1=Joachim T. |last2=Waloszek |first2=Dieter |last3=Maas |first3=Andreas |last4=Liu |first4=Yu |last5=Haug |first5=Carolin |date=March 2012 |title=Functional morphology, ontogeny and evolution of mantis shrimp-like predators in the Cambrian: MANTIS SHRIMP-LIKE CAMBRIAN PREDATORS |journal=Palaeontology |language=en |volume=55 |issue=2 |pages=369–399 |doi=10.1111/j.1475-4983.2011.01124.x|doi-access=free }} The great appendages have been interpreted as [raptorial](/source/raptorial) limbs involved in predation, with those of some genera such as ''[Yohoia](/source/Yohoia)'' being structurally comparable to the raptorial maxillipeds of [mantis shrimp](/source/mantis_shrimp). The spines on the great appendages of leanchoilid megacheirans such as ''[Leanchoilia](/source/Leanchoilia)'' and ''[Yawunik](/source/Yawunik)'' are elongated and whip-like, suggesting an [antennae](/source/Antenna_(zoology))-like sensory role alongside predatory function.{{cite journal |last1=Aria |first1=Cédric |last2=Caron |first2=Jean-Bernard |last3=Gaines |first3=Robert |date=2015 |title=A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny |journal=Palaeontology |volume=58 |issue=4 |pages=629–660 |doi=10.1111/pala.12161|s2cid=86443516 |doi-access=free |bibcode=2015Palgy..58..629A }} The body is divided into the head and the trunk, with the upper surface of the body being covered in overlapping [tergites](/source/tergites) (upper body plates). On the head behind the great appendages and on the trunk were attached pairs of [biramous](/source/biramous) (divided into two branches) limbs running along the body. The biramous limbs of megacheirans are homonomous (i.e. having little differentiation from each other), with [endopod](/source/endopod)s (the lower, leg-like branches) typically divided into seven segments/podomeres, and paddle-shaped [exopod](/source/exopod)s (the upper limb branches), which are fringed with thin lamellae. The morphology of the terminal [telson](/source/telson) segment is variable.{{Cite journal |last1=Aria |first1=Cédric |last2=Zhao |first2=Fangchen |last3=Zeng |first3=Han |last4=Guo |first4=Jin |last5=Zhu |first5=Maoyan |date=December 2020 |title=Fossils from South China redefine the ancestral euarthropod body plan |journal=BMC Evolutionary Biology |language=en |volume=20 |issue=1 |page=4 |doi=10.1186/s12862-019-1560-7 |issn=1471-2148 |pmc=6950928 |pmid=31914921 |doi-access=free |bibcode=2020BMCEE..20....4A }} The biramous limbs of at least some megacheirans have been suggested bear [exite](/source/exite)s.{{Cite journal |last1=Liu |first1=Yu |last2=Edgecombe |first2=Gregory D. |last3=Schmidt |first3=Michel |last4=Bond |first4=Andrew D. |last5=Melzer |first5=Roland R. |last6=Zhai |first6=Dayou |last7=Mai |first7=Huijuan |last8=Zhang |first8=Maoyin |last9=Hou |first9=Xianguang |date=30 July 2021 |title=Exites in Cambrian arthropods and homology of arthropod limb branches |journal=Nature Communications |volume=12 |issue=1 |page=4619 |doi=10.1038/s41467-021-24918-8 |pmid=34330912 |pmc=8324779 |issn=2041-1723|doi-access=free |bibcode=2021NatCo..12.4619L }} == Taxonomy == Several subdivisions within the group are recognised including Jianfengiidae (including ''[Fortiforceps](/source/Fortiforceps)'', ''[Jianfengia](/source/Jianfengia)'', ''[Sklerolibyon](/source/Sklerolibyon)'' and possibly ''[Parapeytoia](/source/Parapeytoia)'') which are known from the Early Cambrian of China, as well as the Cheiromorpha (containing at least ''[Yohoia](/source/Yohoia)'', ''[Haikoucaris](/source/Haikoucaris)'', and Leanchoiliidae), known with certainty from the Early-Mid Cambrian of North America, China and Australia, which is distinguished from Jianfengiidae by having a fewer number of body segments (20+ in Jianfengiidae, as compared to typically only 11 to 13 in Cheiromorpha). The [monophyly](/source/monophyly) of Megacheira is uncertain, with some studies recovering the group as [paraphyletic](/source/paraphyletic). The latest unambiguous megacheiran is the leanchoiliid ''[Lomankus](/source/Lomankus)'' from the Upper Ordovician of North America.{{Cite journal |last=Parry |first=Luke A. |last2=Briggs |first2=Derek E.G. |last3=Ran |first3=Ruixin |last4=O'Flynn |first4=Robert J. |last5=Mai |first5=Huijuan |last6=Clark |first6=Elizabeth G. |last7=Liu |first7=Yu |date=October 2024 |title=A pyritized Ordovician leanchoiliid arthropod |url=https://linkinghub.elsevier.com/retrieve/pii/S0960982224013678 |journal=Current Biology |language=en |doi=10.1016/j.cub.2024.10.013|doi-access=free }} ''[Parapeytoia](/source/Parapeytoia)'' from the Cambrian of China which was formerly misinterpreted as a [radiodont](/source/radiodont) was later suggested to be a member of this group.{{Cite journal |last=Stein |first=Martin |date=1 March 2010 |title=A new arthropod from the Early Cambrian of North Greenland, with a 'great appendage'-like antennula |url=https://academic.oup.com/zoolinnean/article/158/3/477/3798454 |journal=Zoological Journal of the Linnean Society |language=en |volume=158 |issue=3 |pages=477–500 |doi=10.1111/j.1096-3642.2009.00562.x |issn=0024-4082 |url-access=subscription |access-date=29 October 2019 |archive-date=3 November 2019 |archive-url=https://web.archive.org/web/20191103021239/https://academic.oup.com/zoolinnean/article/158/3/477/3798454 |url-status=live }}{{Cite book |last1=Xian-Guang |first1=Hou |url=https://books.google.com/books?id=2YWhDgAAQBAJ&q=Tanglangia&pg=PA184 |title=The Cambrian Fossils of Chengjiang, China: The Flowering of Early Animal Life |last2=Siveter |first2=David J. |last3=Siveter |first3=Derek J. |last4=Aldridge |first4=Richard J. |last5=Pei-Yun |first5=Cong |last6=Gabbott |first6=Sarah E. |last7=Xiao-Ya |first7=Ma |last8=Purnell |first8=Mark A. |last9=Williams |first9=Mark |date=24 April 2017 |publisher=John Wiley & Sons |isbn=978-1-118-89638-9 |language=en |access-date=10 May 2021 |archive-date=8 May 2024 |archive-url=https://web.archive.org/web/20240508131607/https://books.google.com/books?id=2YWhDgAAQBAJ&q=Tanglangia&pg=PA184#v=snippet&q=Tanglangia&f=false |url-status=live }}{{Cite journal |last1=Daley |first1=Allison C. |last2=Budd |first2=Graham E. |last3=Caron |first3=Jean-Bernard |last4=Edgecombe |first4=Gregory D. |last5=Collins |first5=Desmond |date=20 March 2009 |title=The Burgess Shale Anomalocaridid Hurdia and Its Significance for Early Euarthropod Evolution |journal=Science |language=en |volume=323 |issue=5921 |pages=1597–1600 |doi=10.1126/science.1169514 |issn=0036-8075 |pmid=19299617 |bibcode=2009Sci...323.1597D |s2cid=206517995}} Possible megacheirans include ''[Enalikter](/source/Enalikter)'' described from the [Silurian](/source/Silurian) of the [United Kingdom](/source/United_Kingdom), and ''[Bundenbachiellus](/source/Bundenbachiellus)'' from the Early [Devonian](/source/Devonian) of [Germany](/source/Germany);{{Cite journal |last1=Siveter |first1=Derek J. |last2=Briggs |first2=Derek E. G. |last3=Siveter |first3=David J. |last4=Sutton |first4=Mark D. |last5=Legg |first5=David |last6=Joomun |first6=Sarah |date=7 March 2014 |title=A Silurian short-great-appendage arthropod |journal=[Proceedings of the Royal Society B](/source/Proceedings_of_the_Royal_Society_B) |volume=281 |issue=1778 |article-number=20132986 |doi=10.1098/rspb.2013.2986 |pmc=3906945 |pmid=24452026}}{{Cite journal |author1=Derek J. Siveter |author2=Derek E. G. Briggs |author3=David J. Siveter |author4=Mark D. Sutton |author5=David Legg |author6=Sarah Joomun |year=2015 |title=''Enalikter aphson'' is an arthropod: a reply to Struck ''et al.'' (2014) |journal=[Proceedings of the Royal Society B](/source/Proceedings_of_the_Royal_Society_B) |volume=282 |issue=1804 |article-number=20142663 |doi=10.1098/rspb.2014.2663 |pmc=4375861}} due to their possession of great appendage-like cephalic appendages. However, their relationship to megacheirans has been questioned, due to the uncertain [homology](/source/Homology_(biology)) of their appendages.{{Cite journal |last=Aria |first=Cédric |date=October 2022 |title=The origin and early evolution of arthropods |url=http://paleorxiv.org/4zmey/download |journal=Biological Reviews |language=en |volume=97 |issue=5 |pages=1786–1809 |doi=10.1111/brv.12864 |pmid=35475316 |s2cid=243269510 |issn=1464-7931 |access-date=8 May 2024 |archive-date=8 May 2024 |archive-url=https://web.archive.org/web/20240508131708/http://paleorxiv.org/4zmey/download |url-status=live }} ''[Kootenichela](/source/Kootenichela)'' has been suggested to be a [chimera](/source/Chimera_(paleontology)) of various arthropod taxa. Previous inclusion of some "bivalved" genera such as ''[Forfexicaris](/source/Forfexicaris)'', ''[Ovalicephalus](/source/Ovalicephalus)'', and ''[Occacaris](/source/Occacaris)'' to Megacheira was questioned by later investigations.{{Cite journal |last1=Ortega-Hernández |first1=Javier |last2=Janssen |first2=Ralf |last3=Budd |first3=Graham E. |date=1 May 2017 |title=Origin and evolution of the panarthropod head – A palaeobiological and developmental perspective |journal=Arthropod Structure & Development |volume=46 |issue=3, ''Evolution of Segmentation'' |pages=354–379 |doi=10.1016/j.asd.2016.10.011 |issn=1467-8039 |pmid=27989966|doi-access=free |bibcode=2017ArtSD..46..354O }} The Late Cambrian [Orsten](/source/Orsten) taxon ''[Oelandocaris](/source/Oelandocaris)'' typically considered to be a crustacean relative, has also been suggested in some studies to be a megacheiran.{{Cite journal |last1=Aria |first1=Cédric |last2=Caron |first2=Jean-Bernard |date=May 2017 |title=Burgess Shale fossils illustrate the origin of the mandibulate body plan |url=http://www.nature.com/articles/nature22080 |journal=Nature |language=en |volume=545 |issue=7652 |pages=89–92 |doi=10.1038/nature22080 |pmid=28445464 |bibcode=2017Natur.545...89A |s2cid=4454526 |issn=0028-0836 |url-access=subscription |access-date=12 January 2023 |archive-date=23 September 2022 |archive-url=https://web.archive.org/web/20220923174326/https://www.nature.com/articles/nature22080 |url-status=live }} {{gallery|Artistic reconstructions of Fortiforceps foliosa and Sklerolibyon maomima gen. et sp. nov. in early Cambrian Gondwanian seas.png|Life reconstruction of ''[Fortiforceps](/source/Fortiforceps)'' (left) and ''[Sklerolibyon](/source/Sklerolibyon)'' (right)|Jianfengia reconstruction.png|Life restoration of ''[Jianfengia](/source/Jianfengia)''|2012 Leanchoilia superlata 06.png|Closeup of great appendages of ''[Leanchoilia](/source/Leanchoilia) superlata''|Y._kootenayi.png|Life restoration of ''[Yawunik](/source/Yawunik)''|File:Parry et al. 2024 f03 (reconstruction).png|Life restoration of ''[Lomankus](/source/Lomankus)''|File:Oestokerkus megacholix reconstruction.png|Life restoration of ''[Oestokerkus](/source/Oestokerkus)''|||width=200|height=180|align=center}} === List of genera === * †''[Tanglangia](/source/Tanglangia)'' * †''[Oelandocaris](/source/Oelandocaris)''? * †Jianfengiidae ** ''[Sklerolibyon](/source/Sklerolibyon)'' ** ''[Jianfengia](/source/Jianfengia)'' ** ''[Parapeytoia](/source/Parapeytoia)''? ** ''[Fortiforceps](/source/Fortiforceps)'' * †Kootenichelidae? ** ''[Kootenichela](/source/Kootenichela)''? ** ''[Worthenella](/source/Worthenella)''? ** ''[Pseudoiulia](/source/Pseudoiulia)''? * †Cheiromorpha ** ''[Yohoia](/source/Yohoia)'' ** ''[Haikoucaris](/source/Haikoucaris)'' ** Leanchoiliida *** [Enaliktidae](/source/Enaliktidae)? **** ''[Enalikter](/source/Enalikter)''? **** ''[Bundenbachiellus](/source/Bundenbachiellus)''? *** Leanchoiliidae **** ''[Actaeus](/source/Actaeus_(animal))'' **** ''[Alalcomenaeus](/source/Alalcomenaeus)'' **** ''[Leanchoilia](/source/Leanchoilia)'' **** ''[Oestokerkus](/source/Oestokerkus)'' **** ''[Yawunik](/source/Yawunik)'' **** ''[Kanoshoia](/source/Kanoshoia)'' **** ''[Lomankus](/source/Lomankus)'' === Relationship to other arthropods === {{Main|Arthropod head problem}} Megacheirans are either suggested to be stem-group [chelicerate](/source/chelicerate)s (the group containing [arachnids](/source/Arachnid), [sea spiders](/source/Sea_spider) and [horseshoe crabs](/source/Horseshoe_crab), among other extinct groups) or stem-group arthropods outside the split between Chelicerata and the other major group of living arthropods, [Mandibulata](/source/Mandibulata) (which includes [crustaceans](/source/Crustacean), [insects](/source/Insect), [centipedes](/source/Centipede) and [millipedes](/source/Millipede), among others). The former hypothesis is based on the [chelicerae](/source/chelicerae)-like morphology of the great appendages,{{Cite journal |last1=Chen |first1=Junyuan |last2=Waloszek |first2=Dieter |last3=Maas |first3=Andreas |date=2004 |title=A new 'great-appendage' arthropod from the Lower Cambrian of China and homology of chelicerate chelicerae and raptorial antero-ventral appendages |journal=Lethaia |language=en |volume=37 |issue=1 |pages=3–20 |bibcode=2004Letha..37....3C |doi=10.1080/00241160410004764 |issn=1502-3931}}{{Cite journal |last=Chen |first=Jun-Yuan |date=2009 |title=The sudden appearance of diverse animal body plansduring the Cambrian explosion |journal=The International Journal of Developmental Biology |volume=53 |issue=5–6 |pages=733–751 |doi=10.1387/ijdb.072513cj |issn=1696-3547 |pmid=19557680 |doi-access=free}}{{Cite journal |last1=Haug |first1=Joachim T. |last2=Waloszek |first2=Dieter |last3=Maas |first3=Andreas |last4=Liu |first4=Yu |last5=Haug |first5=Carolin |date=March 2012 |title=Functional morphology, ontogeny and evolution of mantis shrimp-like predators in the Cambrian: MANTIS SHRIMP-LIKE CAMBRIAN PREDATORS |journal=Palaeontology |language=en |volume=55 |issue=2 |pages=369–399 |doi=10.1111/j.1475-4983.2011.01124.x |s2cid=82841481 |doi-access=free}} alongside neuroanatomy{{Cite journal |last1=Tanaka |first1=Gengo |last2=Hou |first2=Xianguang |last3=Ma |first3=Xiaoya |last4=Edgecombe |first4=Gregory D. |last5=Strausfeld |first5=Nicholas J. |date=17 October 2013 |title=Chelicerate neural ground pattern in a Cambrian great appendage arthropod |journal=[Nature](/source/Nature_(journal)) |volume=502 |issue=7471 |pages=364–367 |bibcode=2013Natur.502..364T |doi=10.1038/nature12520 |pmid=24132294 |s2cid=4456458}} and the presence of a reduced [labrum](/source/Labrum_(arthropod_mouthpart)){{Cite journal |last1=Liu |first1=Yu |last2=Ortega-Hernández |first2=Javier |last3=Zhai |first3=Dayou |last4=Hou |first4=Xianguang |date=25 June 2020 |title=A Reduced Labrum in a Cambrian Great-Appendage Euarthropod |journal=Current Biology |language=en |volume=30 |issue=15 |pages=3057–3061.e2 |bibcode=2020CBio...30E3057L |doi=10.1016/j.cub.2020.05.085 |issn=0960-9822 |pmid=32589912 |s2cid=220057956 |doi-access=free}} resembling those of modern chelicerates. The proponents of this hypothesis argue that chelicerae and the great appendages are homologous structures.{{Cite journal |last1=Tanaka |first1=Gengo |last2=Hou |first2=Xianguang |last3=Ma |first3=Xiaoya |last4=Edgecombe |first4=Gregory D. |last5=Strausfeld |first5=Nicholas J. |date=17 October 2013 |title=Chelicerate neural ground pattern in a Cambrian great appendage arthropod |url=http://www.nature.com/articles/nature12520 |url-status=live |journal=Nature |language=en |volume=502 |issue=7471 |pages=364–367 |bibcode=2013Natur.502..364T |doi=10.1038/nature12520 |issn=0028-0836 |pmid=24132294 |s2cid=4456458 |url-access=subscription |archive-url=https://web.archive.org/web/20221016211810/https://www.nature.com/articles/nature12520 |archive-date=16 October 2022 |access-date=12 January 2023}} Other studies suggest that the megacheirans are stem-group arthropods based on the argument that the great appendages are homologous to the frontal appendages of stem-group arthropods like ''[Isoxys](/source/Isoxys)'' and [radiodonts](/source/radiodonts). This identity is disputed, with other authors suggesting that the frontal appendages of radiodonts are homologous to the labrum of modern arthropods. Some studies have suggested based on brain and nervous system anatomy that while some megacheirans are stem-chelicerates, others are stem-mandibulates.{{Cite journal |last1=Strausfeld |first1=N. J. |last2=Andrew |first2=D. R. |last3=Hou |first3=X. |last4=Hirth |first4=F. |year=2025 |title=Brain anatomy of the Cambrian fossil ''Jianfengia multisegmentalis'' informs euarthropod phylogeny |journal=Nature Communications |volume=16 |doi=10.1038/s41467-025-62849-w |pmc=12394709 |pmid=40877252 |doi-access=free |article-number=7938}} Cladogram of Arthropoda after Aria et al. 2020, showing megacheirans (as indicated by the orange bar) as a paraphyletic stem group to modern arthropods: {{clade |label1=Arthropoda (''s. l.'') |1={{clade |1=†''[Opabinia](/source/Opabinia)'' 100px |2={{clade |1=†''[Anomalocaris](/source/Anomalocaris)'' 100px |2={{clade |1=†[Hurdiidae](/source/Hurdiidae) 100px |label2=[Deuteropoda](/source/Deuteropoda) |2={{clade |1=†[Isoxyidae](/source/Isoxyidae) 100px |2={{clade |bar1=orange |label1=†Jianfengiidae |1={{clade |1=†''[Jianfengia](/source/Jianfengia)'' 100px |2={{clade |1=†''[Sklerolibyon](/source/Sklerolibyon)'' 100px |2={{clade |1=†''[Fortiforceps](/source/Fortiforceps)'' 100px |2=†''[Parapeytoia](/source/Parapeytoia)'' 100px }}}}}} |2={{clade |bar1=orange |1=†''[Yohoia](/source/Yohoia)'' 100px |2={{clade |bar1=orange |1={{clade |1=†''[Haikoucaris](/source/Haikoucaris)'' 100px |2=†[Leanchoiliidae](/source/Leanchoiliidae) 100px }} |2={{clade |label1=[Chelicerata](/source/Chelicerata) |1={{clade |1=[Pycnogonida](/source/Pycnogonida) (sea spiders) 100px |2={{clade |1=†[Habeliida](/source/Habeliida) 100px |2=[Euchelicerata](/source/Euchelicerata) 100px 100px }}}} |label2=[Antennulata](/source/Antennulata) |2={{clade |1={{clade |1=†''[Kiisortoqia](/source/Kiisortoqia)'' 100px |2=†[Artiopoda](/source/Artiopoda) (including [Trilobita](/source/Trilobita)) 100px }} |2={{clade |1=†''[Marrella](/source/Marrella)'' 100px |2={{clade |1=†''[Aquilonifer](/source/Aquilonifer)'' 100px |2=[Mandibulata](/source/Mandibulata) 100px 100px }}}}}}}}}}}}}}}}}}}}}}|style=font-size:85%}} Placement of monophyletic Megacheira at the base of Chelicerata, after Parry et al. 2024:supplemental material {{clade |label1=Arthropoda (''s. l.'') |1={{clade |1=†''[Opabinia](/source/Opabinia)'' 100px |2={{clade |1=†[Radiodonta](/source/Radiodonta) (e.g. ''[Anomalocaris](/source/Anomalocaris)'') 100px |label2=[Deuteropoda](/source/Deuteropoda) |2={{clade |1={{extinct}}''[Fengzhengia](/source/Fengzhengia)'' 100px |2={{extinct}}''[Kylinxia](/source/Kylinxia)'' 100px |3={{clade |label1=Total group Mandibulata |1={{clade |1={{clade |1=†''[Kiisortoqia](/source/Kiisortoqia)'' 100px |2={{extinct}}''[Bushizheia](/source/Bushizheia)'' 100px }} |2={{clade |1=†[Isoxyidae](/source/Isoxyidae) 100px |2={{clade |1=†[Artiopoda](/source/Artiopoda) (including [Trilobita](/source/Trilobita)) 100px |2={{clade |1=†[Fuxianhuiida](/source/Fuxianhuiida) 100px |2={{clade |1=†[Hymenocarina](/source/Hymenocarina) 100px |2=[Mandibulata](/source/Mandibulata) (crown group) 100px 100px }}}}}}}}}} |label2=Pan-Chelicerata |2={{clade |1={{clade |1=†''[Habelia](/source/Habelia)'' 100px |2={{clade |1=†''[Mollisonia](/source/Mollisonia)'' 100px |2={{clade |1=[Xiphosura](/source/Xiphosura) (horseshoe crabs) 100px |2={{clade |1=†''[Dibasterium](/source/Dibasterium)'' 100px |2=†''[Offacolus](/source/Offacolus)'' 100px }}}}}}}} |label2=[Megacheira](/source/Megacheira) |2={{clade |1={{clade |1=†''[Haikoucaris](/source/Haikoucaris)'' 100px |label2=Leanchoiliidae |2={{clade |1=†''[Oestokerkus](/source/Oestokerkus)'' 100px |2={{clade |1=†''[Yawunik](/source/Yawunik)'' 100px |2=†''[Alalcomenaeus](/source/Alalcomenaeus)'' 100px }} |3={{clade |1=†''[Lomankus](/source/Lomankus)'' 100px |2=†''[Leanchoilia](/source/Leanchoilia)'' 100px }}}}}} |2={{clade |1={{clade |1=†''[Tanglangia](/source/Tanglangia)'' 100px |2=†''[Yohoia](/source/Yohoia)'' 100px }} |label2=Jianfengiidae |2={{clade |1= †''[Fortiforceps](/source/Fortiforceps)'' 100px |2={{clade |1=†''[Sklerolibyon](/source/Sklerolibyon)'' 100px |2=†''[Jianfengia](/source/Jianfengia)'' 100px }}}}}}}}}}}}}}}}}}|style=font-size:85%}}Placement of megacheirans (represented by orange bar) as a paraphyletic assemblage at the base of total-group Chelicerata, after Lerosey-Aubril and Ortega-Hernández (2026):{{Cite journal |last=Lerosey-Aubril|first=Rudy|last2=Ortega-Hernández|first2=Javier|date=2026-04-01|title=A chelicera-bearing arthropod reveals the Cambrian origin of chelicerates|url=https://www.nature.com/articles/s41586-026-10284-2|journal=Nature|language=en|pages=1–7|doi=10.1038/s41586-026-10284-2|issn=1476-4687|doi-access=free}}{{clade|{{clade |label1=Pan-Chelicerata |1={{clade | bar1=orange |1={{clade |1=†''[Jianfengia](/source/Jianfengia)'' 100px |2=†''[Sklerolibyon](/source/Sklerolibyon)'' 100px }} |2={{clade |1=†''[Fortiforceps](/source/Fortiforceps)'' 100px | bar1=orange |2=†''[Yohoia](/source/Yohoia)'' 100px | bar2=orange |3={{clade | bar1=orange |1=†''[Haikoucaris](/source/Haikoucaris)'' 100px |2=†[Leanchoiliidae](/source/Leanchoiliidae) 100px | bar2=orange }} |4={{clade |1=†[Mollisoniida](/source/Mollisoniida) 100px |2={{clade |1=†''[Wisangocaris](/source/Wisangocaris)'' 100px |2=†''[Habelia](/source/Habelia)'' 100px |3={{clade |1=†''[Sanctacaris](/source/Sanctacaris)'' 100px |2=†''[Utahcaris](/source/Utahcaris)'' }} |label4=[chelicerae](/source/chelicerae) present |4={{clade |1=†''[Megachelicerax](/source/Megachelicerax)'' 100px |2={{clade |1=†''[Offacolus](/source/Offacolus)'' 100px |2={{clade |1={{clade |1=†''[Setapedites](/source/Setapedites)'' 100px |2=†''[Dibasterium](/source/Dibasterium)'' 100px }} |2={{clade |1=†''[Weinbergina](/source/Weinbergina)'' 100px |2={{clade |1=†''[Venustulus](/source/Venustulus)'' 100px |2={{clade |1=[Pycnogonida](/source/Pycnogonida) (sea spiders) 100px |2={{clade |1=[Xiphosura](/source/Xiphosura) (horseshoe crabs) 100px |2={{clade |1=[Arachnida](/source/Arachnida) 100px |2=†[Eurypterida](/source/Eurypterida) (sea scorpions) 70px }}}}}}}}}}}}}}}}}}}}}}}}}}|style=font-size:85%}} ==References== {{Reflist|30em}} {{Megacheira}} {{Taxonbar|from=Q1946246}} {{Authority control}} Category:Megacheira Category:Arthropod classes Category:Prehistoric protostome classes --- Adapted from the Wikipedia article [Megacheira](https://en.wikipedia.org/wiki/Megacheira) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Megacheira?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.