Artiopoda

Artiopoda Temporal range: Early Cambrian - Late Permian PreꞒ Ꞓ O S D C P T J K Pg N

Misszhouia, a nektaspid

Cheloniellon, a cheloniellid

Scientific classification

KingdomAnimalia

PhylumArthropoda

(unranked)Deuteropoda

Clade †Artiopoda Hou and Bergstrom, 1997

Subgroups

†Siriocaris? †Squamacula †Thulaspis †Urokodia †Falcatamacaris †Molaria †Bailongia †Notchia? †Strabopida? †Protosutura †Acanthomeridion †Zhiwenia †Australimicola †Trilobitomorpha †Retifacies? †Pygmaclypeatus? †Campanamuta? †Kwanyinaspis †Arthroaspis †Tonglaiia †Nektaspida †Trilobita †Agnostida?(Traditionally included within Trilobita) †Conciliterga †Xandarellida (=Petalopleura) †Vicissicaudata †Sidneyia †Emeraldella †Etainia †Kodymirus †Eozetetes †Tardisia †Carimersa †Cheloniellida †Aglaspidida

Artiopoda is a clade of extinct arthropods that includes trilobites and their close relatives. It was erected by Hou and Bergström in 1997 to encompass a wide diversity of arthropods that would traditionally have been assigned to the Trilobitomorpha. Trilobites, in part due to abundance of findings owing to their mineralized exoskeletons, are by far the best recorded, diverse, and long lived members of the clade. Other members, which lack mineralised exoskeletons, are known mostly from Cambrian deposits.

Description

According to Stein and Selden (2012) artiopods are recognised by the possession of filiform (elongate and relatively thin) antennae, limbs with bilobate exopods (upper branches), with the proximal (closest to base of the limb) lobe being elongate and bearing a lamella, while the distal (further from the limb base) lobe is paddle-shaped and setiforous (bearing hair-or bristle like structures). The limb endopod (inner, leg-like branch) has seven podomeres/segments, with first four podomeres bearing inward facing (endite) structures, while podomeres five and six are stenopodous (cylindrical and stout). Common plesiomorphies (ancestral traits) also include the antennules and at least three sets of post-antennular limbs being attached to the head shield, the postantennular limbs having no or little differentiation into distinct morphologies, and broad paratergal folds which contribute to the dorsoventrally (along the up-down axis) flattened look of artiopods. The limbs of artiopods have also been suggested to bear exites, which were described as similar those of the megacheiran Leanchoilia and probably not homologous to those present in crustaceans.

Restoration of the trilobite Triarthrus eatoni
Restoration of Naraoia magna (Nektaspida)
Life restoration of Sidneyia (Vicissicaudata)
Diagram of Sinoburius (Xandarellida)
Diagram of Olenoides serratus (Trilobita)
Life restoration of Aglaspis (Aglaspidida)
Life restoration of Tegopelte (Conciliterga)
Life restoration of Pleuroctenium (Agnostida)

Taxonomy

Internal taxonomy

The Artiopoda have been considered by many studies to consist of two major clades; one reusing Trilobitomorpha to encompass trilobites, nektaspids, concilitergans and xandarellids, and the other called Vicissicaudata encompassing aglaspidids, xenopods and cheloniellids. A small group, Protosutura, comprising Zhiwenia, Acanthomeridion and Australimicola, has been recognised based on the shared presence of simple sutures on the top (dorsal surface) of the carapace where it splits during ecdysis (moulting), and a short tailspine. This group is often placed basal to the Trilobitomorpha-Vicissicaudata split, alongside other taxa like Squamacula. These relationships are not always recovered.

Relationships with other arthropods

The relationship of Artiopoda with the two major clades of modern arthropods, Chelicerata (the group of sea spiders, horseshoe crabs and arachnids such as spiders and scorpions) and Mandibulata (which contains insects, crustaceans, centipedes and millipedes, among others), are unresolved, with some phylogenies recovering Artiopoda as more closely to chelicerates, forming the clade Arachnomorpha, while others recover Artiopoda as more closely related to mandibulates, forming the clade Antennulata. Some studies place them as stem-group euarthropods, with mandibulates and chelicerates more closely related to each other than either is to Artiopoda. Some studies have recovered a close relationship with Marrellomorpha, with the proposed clade including Artiopoda and Marrellomorpha dubbed Lamellipedia, though this relationship is not found in other analyses. The enigmatic artiopodan-like arthropod Kiisortoqia, which bears large "frontal appendages" has been suggested to be closely related to Artiopoda in some analyses.

Phylogeny

Internal phylogeny

After Jiao et al. 2021.

Artiopoda	Squamacula clypeata Protosutura Zhiwenia coronata Australimicola spriggi Acanthomeridion serratum Acanthomeridion anacanthus Bailongia longicaudata Trilobitomorpha Retifacies abnormalis Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia Conciliterga Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum Xandarellida Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena Nektaspida Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata Vicissicaudata Sidneyia inexpectans Emeraldella brocki Cheloniellida Cheloniellon calmani Triopus draboviensis Duslia insignis Kodymirus vagans Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Squamacula clypeata

	Protosutura Zhiwenia coronata Australimicola spriggi Acanthomeridion serratum Acanthomeridion anacanthus Bailongia longicaudata Trilobitomorpha Retifacies abnormalis Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia Conciliterga Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum Xandarellida Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena Nektaspida Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata Vicissicaudata Sidneyia inexpectans Emeraldella brocki Cheloniellida Cheloniellon calmani Triopus draboviensis Duslia insignis Kodymirus vagans Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
Protosutura	Zhiwenia coronata Australimicola spriggi Acanthomeridion serratum Acanthomeridion anacanthus
	Zhiwenia coronata Australimicola spriggi
	Zhiwenia coronata

	Australimicola spriggi


	Acanthomeridion serratum Acanthomeridion anacanthus
	Acanthomeridion serratum

	Acanthomeridion anacanthus



	Bailongia longicaudata Trilobitomorpha Retifacies abnormalis Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia Conciliterga Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum Xandarellida Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena Nektaspida Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata Vicissicaudata Sidneyia inexpectans Emeraldella brocki Cheloniellida Cheloniellon calmani Triopus draboviensis Duslia insignis Kodymirus vagans Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Bailongia longicaudata

Trilobitomorpha	Retifacies abnormalis Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia Conciliterga Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum Xandarellida Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena Nektaspida Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata
	Retifacies abnormalis

	Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia Conciliterga Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum Xandarellida Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena Nektaspida Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata
	Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia Conciliterga Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum
	Kwanyinaspis maotiashanensis Trilobita Olenoides serratus Eoredlichia intermedia
	Kwanyinaspis maotiashanensis

Trilobita	Olenoides serratus Eoredlichia intermedia
	Olenoides serratus

	Eoredlichia intermedia



Conciliterga	Haifengella corona Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum
	Haifengella corona

	Kuamaia lata Helmetia expansa Tegopelte gigas Skioldia aldna Saperion glumaceum
	Kuamaia lata Helmetia expansa
	Kuamaia lata

	Helmetia expansa


	Tegopelte gigas Skioldia aldna Saperion glumaceum
	Tegopelte gigas

	Skioldia aldna

	Saperion glumaceum





	Xandarellida Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena Nektaspida Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata
Xandarellida	Cindarella eucalla Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena
	Cindarella eucalla

	Luohuilinella deletres Luohuilinella rarus Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena
	Luohuilinella deletres

	Luohuilinella rarus

	Sinoburius lunaris Xandarella spectaculum Phytophilaspis pergamena
	Sinoburius lunaris

	Xandarella spectaculum Phytophilaspis pergamena
	Xandarella spectaculum

	Phytophilaspis pergamena





Nektaspida	Buenaspis forteyi Tariccoia arrusensis Soomaspis splendida Liwia convexa Naraoia spinosa Naraoia compacta Misszhouia longicaudata
	Buenaspis forteyi

	Tariccoia arrusensis Soomaspis splendida Liwia convexa
	Tariccoia arrusensis

	Soomaspis splendida

	Liwia convexa


	Naraoia spinosa Naraoia compacta Misszhouia longicaudata
	Naraoia spinosa

	Naraoia compacta Misszhouia longicaudata
	Naraoia compacta

	Misszhouia longicaudata







Vicissicaudata	Sidneyia inexpectans Emeraldella brocki Cheloniellida Cheloniellon calmani Triopus draboviensis Duslia insignis Kodymirus vagans Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Sidneyia inexpectans

	Emeraldella brocki Cheloniellida Cheloniellon calmani Triopus draboviensis Duslia insignis Kodymirus vagans Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Emeraldella brocki

Cheloniellida	Cheloniellon calmani Triopus draboviensis Duslia insignis
	Cheloniellon calmani

	Triopus draboviensis Duslia insignis
	Triopus draboviensis

	Duslia insignis



	Kodymirus vagans Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Kodymirus vagans

	Eozetetes gemmelli Aglaspidida Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Eozetetes gemmelli

Aglaspidida	Beckwithia typa Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Beckwithia typa

	Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Chlupacaris dubia Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Chlupacaris dubia

	Quasimodaspis brentsae Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis
	Quasimodaspis brentsae

	Tremaglaspis unite Tremaglaspis vanroyi Brachyaglaspis singularis
	Tremaglaspis unite

	Tremaglaspis vanroyi Brachyaglaspis singularis
	Tremaglaspis vanroyi

	Brachyaglaspis singularis




	Cyclopites vulgaris Australaglaspis stonyensis Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Cyclopites vulgaris

	Australaglaspis stonyensis

	Uarthrus instabilis Flobertia kochi Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Uarthrus instabilis

	Flobertia kochi

	Aglaspella granulifera Gogglops ensifer Glypharthrus simplex Aglaspis spinifer Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Aglaspella granulifera

	Gogglops ensifer Glypharthrus simplex Aglaspis spinifer
	Gogglops ensifer

	Glypharthrus simplex Aglaspis spinifer
	Glypharthrus simplex

	Aglaspis spinifer



	Chraspedops modesta Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Chraspedops modesta

	Glypharthrus thomasi Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Glypharthrus thomasi

	Glypharthrus trispinicaudatus Glypharthrus magnoculus
	Glypharthrus trispinicaudatus

	Glypharthrus magnoculus

Implied weights parsimony phylogeny after Berks et al. 2023.

Artiopoda	Thulaspis Squamacula Protosutura Zhiwenia Australimicola Acanthomeridion Vicissicaudata Sidneyia Cheloniellida Emeraldella Kodymirus Eozetetes Aglaspidida Trilobitomorpha Retifacies Pygmaclypeatus Xandarellida Nektaspida Conciliterga Kwanyinaspis Phytophilaspis Trilobita
	Thulaspis Squamacula
	Thulaspis

	Squamacula


	Protosutura Zhiwenia Australimicola Acanthomeridion Vicissicaudata Sidneyia Cheloniellida Emeraldella Kodymirus Eozetetes Aglaspidida Trilobitomorpha Retifacies Pygmaclypeatus Xandarellida Nektaspida Conciliterga Kwanyinaspis Phytophilaspis Trilobita
Protosutura	Zhiwenia Australimicola Acanthomeridion
	Zhiwenia

	Australimicola

	Acanthomeridion


	Vicissicaudata Sidneyia Cheloniellida Emeraldella Kodymirus Eozetetes Aglaspidida Trilobitomorpha Retifacies Pygmaclypeatus Xandarellida Nektaspida Conciliterga Kwanyinaspis Phytophilaspis Trilobita
Vicissicaudata	Sidneyia Cheloniellida Emeraldella Kodymirus Eozetetes Aglaspidida
	Sidneyia

	Cheloniellida Emeraldella Kodymirus Eozetetes Aglaspidida
	Cheloniellida

	Emeraldella Kodymirus Eozetetes Aglaspidida
	Emeraldella

	Kodymirus Eozetetes Aglaspidida
	Kodymirus

	Eozetetes Aglaspidida
	Eozetetes

	Aglaspidida






Trilobitomorpha	Retifacies Pygmaclypeatus Xandarellida Nektaspida Conciliterga Kwanyinaspis Phytophilaspis Trilobita
	Retifacies Pygmaclypeatus
	Retifacies

	Pygmaclypeatus


	Xandarellida Nektaspida Conciliterga Kwanyinaspis Phytophilaspis Trilobita
	Xandarellida

	Nektaspida Conciliterga Kwanyinaspis Phytophilaspis Trilobita
	Nektaspida

	Conciliterga Kwanyinaspis Phytophilaspis Trilobita
	Conciliterga

	Kwanyinaspis Phytophilaspis Trilobita
	Kwanyinaspis

	Phytophilaspis Trilobita
	Phytophilaspis

	Trilobita

External phylogeny

Cladogram supporting the Antennulata hypothesis after Liu et al; 2026:

Panarthropoda	† "Lobopodia" (paraphyletic, ancestral to tardigrades, onychophorans and arthropods) Total group Euarthropoda †Megadictyon †Kerygmachela †Pambdelurion †Opabiniidae † Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	† "Lobopodia" (paraphyletic, ancestral to tardigrades, onychophorans and arthropods)

Total group Euarthropoda	†Megadictyon †Kerygmachela †Pambdelurion †Opabiniidae † Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Megadictyon

	†Kerygmachela †Pambdelurion †Opabiniidae † Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Kerygmachela †Pambdelurion †Opabiniidae † Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Kerygmachela

	†Pambdelurion †Opabiniidae † Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Pambdelurion

	†Opabiniidae † Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Opabiniidae

	† Radiodonta (e.g Anomalocaris) Deuteropoda †Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	† Radiodonta (e.g Anomalocaris)

Deuteropoda	†Erratus †Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Erratus

	†Sunella †Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Sunella

	†Isoxyidae + "Great appendage bivalved forms" †Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Isoxyidae + "Great appendage bivalved forms"

	†Fengzhengia †Kylinxia †Megacheira Pan-Chelicerata †Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Fengzhengia †Kylinxia
	†Fengzhengia

	†Kylinxia


	†Megacheira Pan-Chelicerata
	†Megacheira

	Pan-Chelicerata


	†Kiisortoqia †Bushizheia Antennulata †Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Kiisortoqia

	†Bushizheia

Antennulata	†Artiopoda Mandibulata †Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Artiopoda

Mandibulata	†Fuxianhuiida †Hymenocarina Myriapoda Pancrustacea
	†Fuxianhuiida

	†Hymenocarina Myriapoda Pancrustacea
	†Hymenocarina

	Myriapoda Pancrustacea
	Myriapoda

	Pancrustacea