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Scalidophora
Taxonomy
Scalidophora was named as a phylum. It is extant.
It was synonymized subjectively with Cephalorhyncha by Zrzavy et al. (1998); it was reranked as the superphylum Scalidophora by Liu et al. (2014); it was reranked as the phylum Scalidophora by Edgecombe et al. (2011), Zhang et al. (2015), Harvey and Butterfield (2017), Mussini et al. (2025).
It was assigned to Ecdysozoa by Edgecombe et al. (2011), Liu et al. (2014), Harvey and Butterfield (2017); and to Cycloneuralia by Zhang et al. (2015), Mussini et al. (2025).
It was synonymized subjectively with Cephalorhyncha by Zrzavy et al. (1998); it was reranked as the superphylum Scalidophora by Liu et al. (2014); it was reranked as the phylum Scalidophora by Edgecombe et al. (2011), Zhang et al. (2015), Harvey and Butterfield (2017), Mussini et al. (2025).
It was assigned to Ecdysozoa by Edgecombe et al. (2011), Liu et al. (2014), Harvey and Butterfield (2017); and to Cycloneuralia by Zhang et al. (2015), Mussini et al. (2025).
Subtaxa
Synonymy list
| Year | Name and author |
|---|---|
| 2011 | Scalidophora Edgecombe et al. p. 153 figs. Fig. 1 |
| 2014 | Scalidophora Liu et al. p. 157 |
| 2015 | Scalidophora Zhang et al. p. 1 |
| 2017 | Scalidophora Harvey and Butterfield p. 1 |
| 2025 | Scalidophora Mussini et al. |
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If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.
Ph. Scalidophora Lemburg 1995
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Unr. †Loricifera
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Ph. Priapulida Théel 1906 [priapulid worm]
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Unr. †Archaeopriapulida Conway Morris 1977
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G. †Lecythioscopa Conway Morris 1977
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Fm. †Miskoidae Walcott 1911
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G. †Louisella
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†Louisella pedunculata Conway Morris 1977
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Invalid names: Miskoia pedunculata Walcott 1911 [replaced]
Fm. †Ottoidae Walcott 1911
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G. †Ottoia Walcott 1911
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†Ottoia minor Walcott 1911
†Ottoia prolifica Walcott 1911
†Ottoia tenuis Walcott 1911
Invalid names: Ancalagonidae Conway Morris 1977 [empty]
G. †Corynetis Luo and Hu 1999
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†Corynetis brevis Luo and Hu 1999
†Corynetis fortis Hu et al. 2012
Invalid names: Anningvermis Huang et al. 2004 [synonym]
G. †Omnidens Hou and Others 2006
Unr. †Palaeoscolecidomorpha Shi et al. 2022
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Fm. †Cricocosmiidae
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G. †Houscolex
G. †Tabelliscolex Han et al. 2003
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†Tabelliscolex chengjiangensis Han et al. 2007
†Tabelliscolex hexagonus Han et al. 2003
G. †Markuelia Valkov 1983
Cl. †Palaeoscolecida Conway-Morris and Robison 1986
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G. †Bohemoscolex
G. †Corallioscolex Müller and Hintz-Schallreuter 1993
G. †Cowiella Hinz 1987
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†Cowiella granulata Hinz 1987
†Cowiella reticulata Hinz 1987
†Cowiella undulata Hinz 1987
†Cowiella verrucata Hinz 1987
Or. †Cricocosmida Han et al. 2007
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Fm. †Maotianshaniidae
Fm. †Palaeoscolecidae Whittard 1953
Invalid names: Tylotitidae Han and Others 2003 [empty]
G. †Gammascolex
G. †Milaculum
G. †Plasmuscolex
G. †Protoscolex
Invalid names: Palaeoscolecida [invalid subgroup]
Fm. Priapulidae Gosse 1855
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G. Acanthopriapulus van der Land 1970
G. Priapulopsis Koren and Danielssen 1875
G. Priapulus Lamarck 1816
G. †Xiaoheiqingella Hu 2002
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†Xiaoheiqingella peculiaris Hu 2002
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Invalid names: Yunnanpriapulus halteroformis Huang et al. 2004 [synonym]
Invalid names: Yunnanpriapulus Huang et al. 2004 [synonym]
Or. †Priapulomorpha Slavini-Plawen 1974
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Or. †Selkirkiimorpha Adrianov and Malakhov 1995
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Fm. †Selkirkiidae Conway Morris 1977
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G. †Selkirkia Walcott 1911
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†Selkirkia columbia Conway Morris 1977
†Selkirkia fragilis Walcott 1911
†Selkirkia major Walcott 1908
†Selkirkia sinica Luo and Hu 1999
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Invalid names: Paraselkirkia jinningensis Hou et al. 1999 [synonym]
†Selkirkia spencei Conway Morris and Robison 1986
†Selkirkia transita Wang et al. 2021
†Selkirkia tsering Nanglu and Ortega-Hernández 2024
†Selkirkia willoughbyi Conway Morris and Robison 1986
Invalid names: Paraselkirkia Luo and Hu 1999 [synonym]
G. †Sullulika Peel and Willman 2018
G. †Sicyophorus
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†Sicyophorus rara Luo and Hu 1999
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Invalid names: Protopriapulites haikouensis Hou et al. 1999 [synonym]
Invalid names: Palaeopriapulites Hou et al. 1999 [synonym], Protopriapulites Hou et al. 1999 [synonym]
Invalid names: Halicryptidae Salvini-Plawen 1974 [empty], Maccabeidae Por and Bromley 1974 [empty], Xiaoheiqingidae Chen 2002 [empty]
Invalid names: Kinorhyncha [empty]
Diagnosis
No diagnoses are available
Measurements
No measurements are available
No ecological data are available
Age range
Maximum range based only on fossils: base of the Meishucunian to the top of the Westphalian D or 530.70000 to 307.40000 Ma
Minimum age of oldest fossil (stem group age): 525.5 Ma
Minimum age of oldest fossil (stem group age): 525.5 Ma
Collections (180 total)
| Time interval | Ma | Country or state | Original ID and collection number |
|---|---|---|---|
| Nemakit-Daldynian | China (Shaanxi) | Eolympia pediculata (106454) | |
| Nemakit-Daldynian | Russian Federation (Siberia) | Markuelia sp. (50340) | |
| Early/Lower Cambrian | China (Yunnan) | Paraselkirkia sp., Sicyophorus sp. (185938 185939) Xiaoheiqingella sp. (228573) | |
| Early/Lower Cambrian | China (Sichuan) | Eokinorhynchus rarus (86617) | |
| Waucoban | Canada (Northwest Territories) | Palaeoscolecida indet. (227949) | |
| Caerfai - St Davids | China (Yunnan) | Maotianshania cylindrica (9940 9941) Maotianshania cylindrica, Palaeoscolex sinensis (9937) Palaeoscolex sinensis, Cricocosmia jinningensis (9939) | |
| Meishucunian | China (Shaanxi) | Houscolex lepidotus, Houscolex sp. (6851) | |
| Tommotian | China (Yunnan) | Maotianshania cylindrica (6817 6818 6819) Palaeoscolex sinensis (9784 9785 9786) | |
| Tommotian | Russian Federation (Khabarovsk Krai) | Markuelia secunda (50314) | |
| Tommotian - Caerfai | China (Yunnan) | Maotianshania cylindrica (9783) | |
| Atdabanian | Mongolia (Khovsgol) | Floraconformis egiinensis (235542) | |
| Atdabanian | United Kingdom (England) | Cowiella granulata (79940) Cowiella reticulata, Cowiella verrucata, Cowiella undulata (79933) | |
| Atdabanian | China (Yunnan) | Cricocosmia jiningensis, Maotianshania cylindrica, Palaeoscolex sinensis (86851) Cricocosmia jiningensis, Maotianshania cylindrica, Palaeoscolex sinensis, Tylotites petiolaris, Tabelliscolex hexagonus, Sicyophorus rara, Acosmia maotiania, Corynetis brevis, Paraselkirkia sinica (68083) Cricocosmia jiningensis, Palaeoscolex sinensis, Tabelliscolex hexagonus (86845) Cricocosmia jinningensis, Sicyophorus rara (90136) Mafangscolex sinensis (90138) Mafangscolex sinensis, Cricocosmia jinningensis (90149 90150 90164 90165) Mafangscolex sinensis, Cricocosmia jinningensis, Sicyophorus rara, Corynetis brevis, Paraselkirkia sinica (90133 90135 90137) Mafangscolex sinensis, Cricocosmia jinningensis, Sicyophorus rara, Corynetis brevis, Xiaoheiqingella peculiaris, Paraselkirkia sinica (90151) Mafangscolex sinensis, Paraselkirkia sinica (90152) Maotianshania cylindrica (90494) Maotianshania cylindrica, Mafangscolex sinensis (90161 90163 90166) Maotianshania cylindrica, Palaeoscolex sinensis (9938) Maotianshania cylindrica, Palaeoscolex sinensis, Sicyophorus rara, Palaeopriapulites parvus (86846) Maotianshania sp., Acosmia sp. (86933) Priapulida indet., Cricocosmia sp., Palaeoscolex sp., Maotianshania sp., Acosmia sp., Corynetis sp., Paraselkirkia sp., Sicyophorus sp., Anningvermis sp. (86929) Priapulida indet., Palaeoscolex sp., Maotianshania sp., Corynetis sp., Paraselkirkia sp., Protopriapulites sp. (86930) | |
| Atdabanian | China (Hubei) | Maotianshania cylindrica (68086) | |
| Atdabanian | Mongolia | Markuelia sp. (219046) | |
| Nangaoian | China (Yunnan) | Mafangscolex sinensi, Cricocosmia jinningensis, Corynetis brevis, Archotuba conoidalis, Paraselkirkia jinningensis (194160) Mafangscolex sinensis (132328) Mafangscolex sinensis, Cricocosmia jinningensis (194140) Mafangscolex sinensis, Cricocosmia jinningensis, Archotuba conoidalis (132329 194162) Mafangscolex sinensis, Cricocosmia jinningensis, Archotuba conoidalis, Paraselkirkia jinningensis (194150 194155) Mafangscolex sinensis, Cricocosmia jinningensis, Corynetis brevis, Palaeopriapulites parvus, Xiaoheiqingella peculiaris, Archotuba conoidalis, Paraselkirkia jinningensis (132330) Mafangscolex sinensis, Cricocosmia jinningensis, Palaeopriapulites parvus, Archotuba conoidalis, Paraselkirkia jinningensis (194143 194161 194163) Mafangscolex sinensis, Cricocosmia jinningensis, Xiaoheiqingella peculiaris, Archotuba conoidalis, Paraselkirkia jinningensis (194141) Mafangscolex sinensis, Cricocosmia jinningensis, Yunnanpriapulus halteroformis, Paraselkirkia jinningensis (194153) Mafangscolex sinensis, Tabelliscolex hexagonus, Cricocosmia jinningensis, Archotuba conoidalis (132327) Selkirkia sinica (234719) | |
| Nangaoian | China (Hubai) | Selkirkia sp., Kinorhyncha indet., Archotuba sp., Corynetis sp. (219687) | |
| Stage 3 | China (Yunnan) | Selkirkia transita (235078) | |
| Atdabanian - Caerfai | Greenland | Singuuriqia simoni (9780) | |
| Branchian | United Kingdom (England) | Cowiella reticulata (79935 79937) Hadimopanella apicata (79931) Milaculum sp. indet., Cowiella reticulata, Cowiella verrucata (79928) | |
| Botomian | Australia (South Australia) | Palaeoscolex antiquus, Wronascolex iacoborum (187498) | |
| Botomian | Russian Federation (Siberia) | Hadimopanella knappologica (174262) | |
| Botomian | China (Yunnan) | Guanduscolex minor (90357) Palaeoscolex sp. (90363 90367 90368 90369 90370 90371) Paramaotianshania zijunia, Corynetis fortis (139521) | |
| Botomian | Russian Federation (Yakutia) | Palaeoscolecida indet., Palaeoscolex sp., Corallioscolex sp., Palaeoscolex lubovae, Palaeoscolex spinosus, Sahascolex labyrinthus (90438) | |
| Lenian | Antarctica (South Shetland Islands) | Hadimopanella antarctica (49692) | |
| Dyeran | USA (Pennsylvania) | Ottoia sp., Selkirkia pennsylvanica (192 226194) | |
| Dyeran | Greenland | Hadimopanella apicata (49466 49467 49469 49470 49473 67782 67783 67787 67795 67797 67798 67799 67800) | |
| Dyeran | USA (Nevada) | Ottoia sp. (10560) Selkirkia sp. (9820) | |
| Duyunian | China (Shaanxi) | Palaeoscolecida indet. (90680) | |
| Duyunian | China (Guizhou) | Priapulida new worm? B, Wronascolex sp. (237148) | |
| Stage 4 | USA (Vermont) | Ottoia sp. (237967) | |
| Toyonian | Australia (South Australia) | Kaimenella reticulata (64472 97986 97987 118171 118173) | |
| Toyonian | Türkiye (Konya) | Hadimopanella oezgueli (93621) | |
| Delamaran | USA (Utah) | Selkirkia columbia, Palaeoscolex ratcliffei, Selkirkia spencei (77218) Selkirkia willoughby, Selkirkia sp. (77219) | |
| Delamaran | USA (Idaho) | Selkirkia sp. (207) Selkirkia spencei (76096) | |
| Delamaran | Canada (British Columbia) | Ottoia prolifica, Selkirkia columbia (84414 84416 84417 84434 84437) Ottoia prolifica, Selkirkia columbia, Louisella pedunculata (84423 84436) Priapulida indet. A, Selkirkia sp. A, Ottoia prolifica, Selkirkia columbia, Louisella pedunculata (84425) Priapulida indet. B, Fieldia lanceolata, Ottoia prolifica, Selkirkia columbia, Louisella pedunculata (84418) Priapulida indet., Ottoia prolifica, Selkirkia columbia (121959) Selkirkia columbia (84413 84415 84420) Selkirkia sp. A, Ottoia prolifica, Selkirkia columbia (84421) Selkirkia sp. A, Ancalagon minor, Fieldia lanceolata, Ottoia prolifica, Selkirkia columbia, Louisella pedunculata (84424) Selkirkia sp. A, Fieldia lanceolata, Ottoia prolifica, Selkirkia columbia (84427) Selkirkia sp. A, Fieldia lanceolata, Ottoia prolifica, Selkirkia columbia, Louisella pedunculata (84422) Selkirkia sp. A, Ottoia prolifica, Selkirkia columbia (84412 84419 84426 84430 84431 84433) Selkirkia sp. A, Ottoia prolifica, Selkirkia columbia, Louisella pedunculata (84428 84429 84432 84435) | |
| Amgan | Kyrgyzstan | Hadimopanella oezgueli (156378) | |
| Amgan | Russian Federation (Siberia) | Hadimopanella knappologica (110619) | |
| Solvan | Russian Federation (Siberia) | Hadimopanella knappologica (49429) | |
| Solvan | USA (Nevada) | Ottoia sp. (9215) | |
| Taijangian | China (Guizhou) | Ottoia sp. (79770) | |
| Middle Cambrian | Canada (Brithish Columbia) | Selkirkia sp. (188664) | |
| Middle Cambrian | Canada (British Columbia) | Priapulida indet., Selkirkia sp. (171308) | |
| St Davids | Canada (British Columbia) | Ottoia sp. (122017) Ottoia sp., Ottoia prolifica (122094) | |
| Marjumian | Canada (British Columbia) | Ottoia sp. (122072) | |
| Marjumian | USA (Utah) | Ottoia prolifica (77206) Ottoia sp., Selkirkia sp., Palaeoscolecida gen. et sp. nov. A (230681) Scathascolex minor, Ottoia prolifica, Selkirkia willoughbyi (230679) | |
| Guzhangian | Canada (Alberta) | Selkirkia sp., Priapulomorpha indet., Ottoia prolifica, Badgiromorpha delicata (238940) | |
| Guzhangian | China (Yunnan) | Palaeoscolecida indet. (219647) | |
| Furongian | Canada (Alberta) | Eolorica deadwoodensis (197354) | |
| Merioneth | Russian Federation (Siberia) | Protoscolex tchopkoensis (10009) | |
| Tremadocian | Morocco (Zagora) | Selkirkia tsering (235072) | |
| Floian | China (Hunan) | Ottoia sp., Palaeoscolecida indet. (226639) | |
| Darriwilian | United Kingdom (Wales) | Priapulida NMW.2021.3 G.97, Palaeoscolecida NMW.2021.3 G.89, Kinorhyncha NMW.2021.3 G.69, Archaeopriapulida NMW.2021.3 G.19 (230215) | |
| Llandeilo | United Kingdom (Wales) | Palaeoscolecidae gen. et sp. nov. Botting et al. in press (216511) | |
| Turinian | Canada (British Columbia) | Milaculum sp. (192195) | |
| Rocklandian | USA (Wisconsin) | Milaculum sp. (90362) | |
| Rocklandian - Shermanian | USA (Wisconsin) | Milaculum sp. (90360) | |
| Shermanian | USA (Wisconsin) | Milaculum sp. (90366) | |
| Katian | USA (New York) | Protoscolex sp. (340) | |
| Edenian | USA (Ohio) | Protoscolex sp. (383 384) | |
| Maysvillian | Canada (Ontario) | Protoscolex sp. (26302) | |
| Richmondian | Canada (Nunavut) | Milaculum ethinclarki (360) | |
| Rhuddanian - Aeronian | USA (Iowa) | Protoscolex sp. (26860) | |
| Telychian | USA (Wisconsin) | Palaeoscolecida indet. (91415) | |
| Ludlow | USA (Illinois) | Protoscolex ruedemanni (76450) Protoscolex sp. (26861) | |
| Ludfordian | United Kingdom | Radnorscolex latus (236353 236354 236355) | |
| Westphalian D | USA (Illinois) | Priapulites konecniorum (122493 122538 151852 168004) |