Artiopoda

The Artiopoda is a grouping 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.

Artiopoda Temporal range:
Misszhouia, a nektaspid
Cheloniellon, a cheloniellid
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
(unranked):	†Artiopoda Hou and Bergstrom, 1997
Subgroupings
†Squamacula †Thulaspis †Urokodia Protosutura †Acanthomeridion †Zhiwenia †Australimicola †Campanamuta †Falcatamacaris †Molaria †Retifacies †Pygmaclypeatus †Bailongia †Strabopida? Trilobitomorpha †Retifacies? †Pygmaclypeatus? †Campanamuta? †Kwanyinaspis †Arthroaspis †Tonglaiia †Nektaspida †Trilobita †Agnostida?(Traditionally included within Trilobita) †Conciliterga †Haifengella †Helmetia †Kuamaia †Kwanyinaspis †Rhombicalvaria †Saperion †Skioldia †Tegopelte †Xandarellida (=Petalopleura) †Luohuilinella †Cindarella †Phytophilaspis †Xandarella †Sinoburius †Zhugeia Vicissicaudata †Xenopoda (possibly paraphyletic) †Sidneyia †Emeraldella †Etania †Kodymirus †Eozetetes †Carimersa †Cheloniellida †Aglaspidida

Description

Morphology of Cambrian artiopod Retifacies, showing the morphology of the limbs (E, F) Key for limb morphology: exopod (exo) exite (exi) protopodite (pt) podomere (pd) spine/setae (s) terminal claw (tc)

According to Stein and Selden (2012) artiopods are recognised by the possession of filiform antennulae, limbs with bilobate exopods, with the proximal lobe being elongate and bearing a lamella, while the distal lobe is paddle-shaped and setiforous (bearing hair-or bristle like structures). The limb endopod 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 also include the antennules and at least three sets of post-antennular limbs being incorporated into the head shield, the postantennular limbs having no or little differentiation into distinct morphologies, and broad paratergal folds which contribute to the dorsoventrally 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.

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. There are some taxa, such as Squamacula and the members of "Protosutura", which often are placed near the base of Artiopoda, outside the Vicissicaudata+Trilobitomorpha clade. These relationships are not always recovered.

Relationships with other arthropods

The relationship of Artiopoda with the two major clades of modern arthropods, the Chelicerata and the Mandibulata, 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.

Gallery

• 
  Restoration of the trilobite Triarthrus eatoni
• 
  Fossils of Naraoia compacta (Nektaspida)
• 
  Life restoration of Sidneyia (Vicissicaudata)
• 
  Diagram of Sinoburius (Xandarellida)
• 
  Life restoration of Trimerus (Trilobita)

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