Biology:Deuteropoda
Deuteropoda is a clade of arthropods whose members are distinguished from more basal stem-group arthropods like radiodonts by an anatomical reorganization of the head region (which is proposed to include a differentiated, titular "deutocerebral" first appendage pair, a multisegmented head, and a hypostome/labrum complex), and by bearing pairs of segmented biramous (two branched) limbs.[1]
The clade contains all living arthropods (i.e. chelicerates and mandibulates) as well as several fossil groups that share these characteristics (e.g. Fuxianhuiida, Megacheira, Isoxyida, and Artiopoda, the last of which includes the trilobites), while excluding other fossil groups that are more 'basal' or 'primitive' (e.g. radiodonts, opabiniids and lobopodians).[1][2] Members of Deuteropoda that are basal to the last common ancestor of chelicerates and mandibulates (and thus lie outside the crown group of Arthropoda) are often referred to as "upper stem group arthropods".[1][3]
Defining characteristics

Javier Ortega-Hernández in his 2016 study naming the clade, proposed that Deuteropoda is differentiated from more basal total group arthropods by the presence of a differentiated labrum (part of the mouth) and a differentiated first pair of appendages innervated by a part of the brain called the deutocerebrum (making them "deutocerebral").[1] In contrast, lobopodians (part of the "lower stem group") and onychophorans have a pair of pre-ocular (located in front of the eyes) appendages innervated by a part of the brain called the protocerebrum (making them "protocerebral"), which has been proposed by a number of authors to have evolved to be the labrum of modern living arthropods, and to be homologous (derived from the same ancestral structure) to the segmented frontal appendages of radiodonts, which these authors argue was protocerebral.[2] In living arthropods, the deutocerebral appendage corresponds to the first pair of antennae in mandibulates and the chelicerae in chelicerates.[4] All deuteropods are suggested to have a multi-segmented head with at least three segments including the protocerebrum, deutocerebrum and tritocerebrum (the head in non-deuteropodan stem-group arthropods including radiodonts is generally suggested in contrast to have been composed of a single segment including the protocerebrum).[4]
While the clade comprising Deuteropoda (though not necessarily the name itself) is well accepted, the characters Ortega-Hernández proposed as supporting Deuteropoda are somewhat controversial, with number of authors arguing that the transformation of the head from radiodonts to early deuteropods is less radical than proposed,[3] and argue that there is no clear evidence of a switch between a protocerebral head appendage in radiodonts to a deutocerebral one in Deuteropoda,[5] with some authors arguing that brains of radiodonts were two-segmented rather than single segmented and that their frontal appendages were in fact deutocerebral,[6] or that the first appendages of early deuteropodans were in fact protocerebral.[7]
Many early primitive deuteropod first appendages like those in Kylinxia and isoxyids are segmented grasping appendages that are upward (dorsally)-curling, as opposed to downward curling like the frontal appendages of radiodonts, which may represent a shared ancestral trait for most or all of Deuteropoda. It is disputed whether or not the upward-curling grasping appendage of early deuteropods are homologous to the frontal appendage of radiodonts, to which they show strong structural similarities, or that they were convergently evolved separately.[8]
Members of Deuteropoda lack the oral cone mouth present in radiodonts, but instead ancestrally possess a posteriorly facing mouth on the underside of the head, which is at least sometimes covered by a hypostome.[3] Another distinctive characteristic ancestrally present in deuteropodans (with the exception of the radiodont-like Oreinorema[3]) but not more basal stem-arthropods is the presence of two-branched (biramous) segmented (arthropodized) limbs running along the body posterior to the mouth, distinct from the swimming flaps present in radiodonts and "gilled lobopodians" and the lobopods of lobopodians,[1][8] though the appendages of many modern arthropods are secondarily uniramous (unbranched).[9][10] The segmentation of the body limbs may have evolved from the co-option of the genes causing the segmentation of the first head appendage in the Deuteropoda-Radiodonta common ancestor.[4] Most deuteropods (with the exception of Erratus and isoxyids, and secondarily some modern arthropods) also have a segmented and jointed (arthrodized) trunk (main body section),[8]
Phylogeny
The cladogram below is a possible panarthropod phylogeny, taking into account the differentiation between "lower stem group arthropods" (Total group Euarthropoda) and "upper stem group arthropods" (Deuteropoda), based on a phylogenetic analysis published in 2026.[8] Not all parts of this cladogram are universally accepted, as some other studies have found a closer relationship to Chelicerata than to Mandibulata for Artiopoda,[11] and for Megacheira to be outside crown-group Arthropoda.[11]
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References
- ↑ 1.0 1.1 1.2 1.3 1.4 Ortega-Hernández, Javier (2016), "Making sense of 'lower' and 'upper' stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848", Biol. Rev. 91 (1): 255–273, doi:10.1111/brv.12168, PMID 25528950
- ↑ 2.0 2.1 Budd, Graham E. (2021). "The origin and evolution of the euarthropod labrum". Arthropod Structure & Development 62. doi:10.1016/j.asd.2021.101048. ISSN 1467-8039. PMID 33862532.
- ↑ 3.0 3.1 3.2 3.3 Liu, Yu; O’Flynn, Robert J.; Yu, Mengxiao; Mai, Huijuan; Hou, Xianguang; Edgecombe, Gregory D. (May 2026). "A transitional, early Cambrian species bridges radiodonts and upper stem-group euarthropods" (in en). Current Biology. doi:10.1016/j.cub.2026.05.002. https://linkinghub.elsevier.com/retrieve/pii/S0960982226005786.
- ↑ 4.0 4.1 4.2 Lev, Oren; Edgecombe, Gregory D; Chipman, Ariel D (2022-05-24). "Serial Homology and Segment Identity in the Arthropod Head" (in en). Integrative Organismal Biology 4 (1). doi:10.1093/iob/obac015. ISSN 2517-4843. PMID 35620450. PMC 9128542. https://academic.oup.com/iob/article/doi/10.1093/iob/obac015/6571934.
- ↑ Aria, Cédric (29 May 2019). "Reviewing the bases for a nomenclatural uniformization of the highest taxonomic levels in arthropods" (in en). Geological Magazine 156 (08): 1463–1468. doi:10.1017/S0016756819000475. ISSN 0016-7568. https://www.cambridge.org/core/product/identifier/S0016756819000475/type/journal_article.
- ↑ Moysiuk, Joseph; Caron, Jean-Bernard (2022-08-08). "A three-eyed radiodont with fossilized neuroanatomy informs the origin of the arthropod head and segmentation" (in English). Current Biology 32 (15): 3302–3316.e2. doi:10.1016/j.cub.2022.06.027. ISSN 0960-9822. PMID 35809569. https://www.cell.com/current-biology/abstract/S0960-9822(22)00986-1.
- ↑ Aria, Cédric; Caron, Jean-Bernard (2015-06-03). Friedman, Matt. ed. "Cephalic and Limb Anatomy of a New Isoxyid from the Burgess Shale and the Role of “Stem Bivalved Arthropods” in the Disparity of the Frontalmost Appendage" (in en). PLOS ONE 10 (6): e0124979. doi:10.1371/journal.pone.0124979. ISSN 1932-6203. PMID 26038846. PMC 4454494. https://dx.plos.org/10.1371/journal.pone.0124979.
- ↑ 8.0 8.1 8.2 8.3 Liu, Cong; Pates, Stephen; Zhang, Mingjing; Wu, Yu; Ma, Jiaxin; Fu, Dongjing; Zhang, Xingliang (2026-03-21). "3D morphology of the Cambrian bivalved arthropod Sunella informs about head segmentation, arthrodization, and arthropodization" (in en). Communications Biology. doi:10.1038/s42003-026-09909-z. ISSN 2399-3642. https://www.nature.com/articles/s42003-026-09909-z.
- ↑ Dworkin, Ian M.; Tanda, Soichi; Larsen, Ellen (November 2001). "Are entrenched characters developmentally constrained? Creating biramous limbs in an insect" (in en). Evolution & Development 3 (6): 424–431. doi:10.1046/j.1525-142X.2001.01047.x. ISSN 1520-541X. https://onlinelibrary.wiley.com/doi/10.1046/j.1525-142X.2001.01047.x.
- ↑ Lustri, Lorenzo; Gueriau, Pierre; Daley, Allison C. (2024-05-07). "Lower Ordovician synziphosurine reveals early euchelicerate diversity and evolution" (in en). Nature Communications 15 (1). doi:10.1038/s41467-024-48013-w. ISSN 2041-1723. PMID 38714651. PMC 11076625. https://www.nature.com/articles/s41467-024-48013-w.
- ↑ 11.0 11.1 Aria, Cédric (October 2022). "The origin and early evolution of arthropods" (in en). Biological Reviews 97 (5): 1786–1809. doi:10.1111/brv.12864. ISSN 1464-7931. PMID 35475316. https://onlinelibrary.wiley.com/doi/10.1111/brv.12864.
Wikidata ☰ Q112480276 entry
