Biology:Panarthropoda

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Short description: Animal taxon

Panarthropoda
Temporal range: Cambrian - Recent
Molecular clock and ichnofossils indicate a possible Ediacaran origin[1] [2]
Peripatopsis lawrencei 134803788 (cropped).jpg
Panarthropods include onychophorans such as Peripatopsis and arthropods such as polydesmid millipedes
Scientific classification e
Domain: Eukaryota
Kingdom: Animalia
Clade: ParaHoxozoa
Clade: Bilateria
Clade: Nephrozoa
(unranked): Protostomia
Superphylum: Ecdysozoa
(unranked): Panarthropoda
Nielsen, 1995
Phyla

Panarthropoda is a proposed animal clade containing the extant phyla Arthropoda, Tardigrada (water bears) and Onychophora (velvet worms).[3] Panarthropods also include extinct marine legged worms known as lobopodians ("Lobopodia"), a paraphyletic group where the last common ancestor and basal members (stem-group) of each extant panarthropod phylum are thought to have risen.[4][5][6][7][8][9] However the term "Lobopodia" is sometimes expanded to include tardigrades and onychophorans as well.[5]

Common characteristics of the Panarthropoda include a segmented body, paired ladder-like ventral nervous system, and the presence of paired appendages correlated with body segments.[10][5][6][8]

Taxonomy

Bilateria
Deuterostomia

Echinodermata, Chordata and relatives

 
Protostomia
Spiralia

Annelida, Mollusca and relatives

 
Ecdysozoa
Cycloneuralia

Priapulida, Nematoda and relatives

 
Panarthropoda

Onychophora, Tardigrada and Arthropoda

 
 
 
 

Not all studies support the monophyly of Panarthropoda,[11] but most do, including neuroanatomical,[12] phylogenomic[10][13][14] and palaeontological[15][4][5][6] studies. At least a close relationship between onychophorans and arthropods is widely agreed upon, but the position of tardigrades is more controversial.[16] Some phylogenomic studies have found tardigrades to be more closely related to nematodes.[17][18] Traditionally, panarthropods were considered to be closely related to the annelids, grouped together as the Articulata (animals with body segments), but subsequent phylogenomic studies consistently place them closer to cycloneuralians (nematodes, nematomorphs, loriciferans, kinorhynchas & priapulids), grouped together as Ecdysozoa.[19][20][13][14] While annelids are placed among the Spiralia (making them more closely related to mollusks, flatworms and such),[20] having evolved their segmented bodies convergently.[21]

Interrelationship

Panarthropoda
 

(Lobopodian taxa controversial)*

 
 
 

(Lobopodian taxa controversial)*

 
 
 

Antennacanthopodia Antennacanthopodia.jpg*

 
 

Crown-group Onychophora Velvet worm.jpg

 
 
 
 
 

(Lobopodian taxa controversial)*

 
 

Crown-group Tardigrada SEM image of Milnesium tardigradum in active state - journal.pone.0045682.g001-2.png

 
 
 
 

Siberiid lobopodians 20191217 Siberiida Siberion Megadictyon Jianshanopodia.png*

 
 
 

Gilled lobopodians 20210730 Gilled lobopodians Pambdelurion Kerygmachela.png*

 
 
 

Opabinia 20191108 Opabinia regalis.png

 
 

Radiodonta 20191201 Radiodonta Amplectobelua Anomalocaris Aegirocassis Lyrarapax Peytoia Laggania Hurdia.png

 
 

Euarthropoda Arthropoda.jpg

 
 
 
 
 
Summarized interrelationship of Panarthropoda, with focus on phylogeny between lobopodians (asterisk) and extant panarthropods (bold). Relationship between the total-group of extant panarthropods is unresolve (see text for discussion).[5][6]

There are three competing hyphotheses for the interrelationship between the extant panarthropod phyla, each known as Tactopoda (Arthropoda+Tardigrada), Antennopoda (Arthropoda+Onychophora), and the sister relationship between Onychophora and Tardigrada (Lobopodia sensu Smith & Goldstein 2017).[22]

Panarthropoda
 

Onychophora

 
Tactopoda
 

Tardigrada

 
 

Arthropoda

 
 
 
Panarthropoda
 

Tardigrada

 
Antennopoda
 

Onychophora

 
 

Arthropoda

 
 
 
Panarthropoda
 

Arthropoda

 
Lobopodia
sensu Smith & Goldstein 2017
 

Tardigrada

 
 

Onychophora

 
 
 

Tactopoda had been supported by mitochondrial gene arrangements,[23] palaeontological[24][4] and neuroanatomical evidences, specifically the presence of segmented ganglia shared by arthropods and tardigrades.[25] Antennopodia united by the presence of specialized head appendages and deutocerebrum (additional second section of the brain), but subsequent anatomical studies suggest these features were convergently evolved between onychophoran and arthropod lineages.[15][26] Onychophorans and tardigrades shared some lobopodian traits (e.g. soft cuticle, lobopodous appendages and peripheral nerve roots), but these were generally considered to be plesiomorphies traced back to the last common ancestor of Panarthropoda or Ecdysozoa.[4][5][27][6] While most phylogenomic analyses support the monophyly of Panarthropoda, the results of interrelationship between the three phyla are less correlated - some of them inconsistently placing Tardigrada within Arthropoda,[10] while the others mostly recovering either Antennopoda or Onychophora+Tardigrada.[10][13][14]

  • Anatomy of onychophoran anterior region, showing specialized appendages (ant, md) and deutocerebrum (D).

  • Nervous system of a tardigrade, showing ventral nerve cord with segmented ganglia (ga1-4).

  • Nervous system of a remipede arthropod, showing the presence of both deutocerebrum (dc) and ventral nerve cord (vnc) organized by segmented ganglia.

Within extinct lobopodians, at least Antennacanthopodia are widely accepted as part of the onychophoran stem-group.[28][4][29][30][31][27][32][33][6][34][35] On the other hand, siberiids (Siberion, Megadictyon and Jianshanopodia) and gilled lobopodians (Pambdelurion and Kerygmachela) represent transitional forms between typical lobopodians and basal arthropods (e.g. Opabinia and Radiodonta).[28][4][29][30][31][27][32][5][6][33][34][7][8][35][36][9] The positions of most other lobopodians (e.g. Hallucigenia and luolishaniids as stem onychophorans[4][29][30][31][27][32][35] or stem panarthropods[33][34][36]), including the lobopodian members of tardigrade stem-group (represented by Onychodictyon ferox[4][29][30][31][27][32][35] or Aysheaia[33][34][36]) are more controversial.


Sialomorpha, a genus of microinvertebrate discovered in Dominican amber in 2019, is also considered to be a panarthropod. However, due to the unusual combination of tardigrade and mite-like characteristics, its exact placement is uncertain.[37]

See also

  • List of bilaterial animal orders

References

  1. ↑ Rota-Stabelli, Omar; Daley, Allison C.; Pisani, Davide. "Molecular Timetrees Reveal a Cambrian Colonization of Land and a New Scenario for Ecdysozoan Evolution" Current Biology, Volume 23, Issue 5 (31 January, 2013) | https://doi.org/10.1016/j.cub.2013.01.026
  2. ↑ "The Shibantan Lagerstätte: insights into the Proterozoic–Phanerozoic transition". Journal of the Geological Society. January 2021. doi:10.1144/jgs2020-135. 
  3. ↑ Telford, M. J (27 April 2008). "The evolution of the Ecdysozoa". Philosophical Transactions of the Royal Society B: Biological Sciences 363 (1496): 1529–1537. doi:10.1098/rstb.2007.2243. PMID 18192181. 
  4. ↑ 4.0 4.1 4.2 4.3 4.4 4.5 4.6 4.7 Smith, Martin R.; Ortega-Hernández, Javier (2014). "Hallucigenia's onychophoran-like claws and the case for Tactopoda". Nature 514 (7522): 363–366. doi:10.1038/nature13576. PMID 25132546. Bibcode: 2014Natur.514..363S. https://www.researchgate.net/publication/264868021. 
  5. ↑ 5.0 5.1 5.2 5.3 5.4 5.5 5.6 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: Upper and lower stem-Euarthropoda". Biological Reviews 91 (1): 255–273. doi:10.1111/brv.12168. ISSN 1464-7931. PMID 25528950. 
  6. ↑ 6.0 6.1 6.2 6.3 6.4 6.5 6.6 Ortega-Hernández, Javier; Janssen, Ralf; Budd, Graham E. (2017-05-01). "Origin and evolution of the panarthropod head – A palaeobiological and developmental perspective" (in en). Arthropod Structure & Development. Evolution of Segmentation 46 (3): 354–379. doi:10.1016/j.asd.2016.10.011. ISSN 1467-8039. PMID 27989966. 
  7. ↑ 7.0 7.1 Giribet, Gonzalo; Edgecombe, Gregory D. (2019-06-17). "The Phylogeny and Evolutionary History of Arthropods" (in English). Current Biology 29 (12): R592–R602. doi:10.1016/j.cub.2019.04.057. ISSN 0960-9822. PMID 31211983. 
  8. ↑ 8.0 8.1 8.2 Chipman, Ariel D.; Edgecombe, Gregory D. (2019-10-09). "Developing an integrated understanding of the evolution of arthropod segmentation using fossils and evo-devo". Proceedings of the Royal Society B: Biological Sciences 286 (1912): 20191881. doi:10.1098/rspb.2019.1881. ISSN 0962-8452. PMID 31575373. 
  9. ↑ 9.0 9.1 Edgecombe, Gregory D. (2020-11-02). "Arthropod origins: Integrating paleontological and molecular evidence". Annual Review of Ecology, Evolution, and Systematics 51 (1): 1–25. doi:10.1146/annurev-ecolsys-011720-124437. ISSN 1543-592X. https://www.annualreviews.org/doi/10.1146/annurev-ecolsys-011720-124437. 
  10. ↑ 10.0 10.1 10.2 10.3 Rota-Stabelli, O.; Kayal, E.; Gleeson, D.; Daub, J.; Boore, J.; Telford, M.; Pisani, D.; Blaxter, M. et al. (2010). "Ecdysozoan mitogenomics: evidence for a common origin of the legged invertebrates, the Panarthropoda". Genome Biology and Evolution 2: 425–440. doi:10.1093/gbe/evq030. PMID 20624745. 
  11. ↑ Dunn, C. W.; Hejnol, A.; Matus, D. Q.; Pang, K.; Browne, W. E.; Smith, S. A.; Seaver, E.; Rouse, G. W. et al. (10 April 2008). "Broad phylogenomic sampling improves resolution of the animal tree of life". Nature 452 (7188): 745–749. doi:10.1038/nature06614. PMID 18322464. Bibcode: 2008Natur.452..745D. 
  12. ↑ Persson, Dennis K. (November 2012). "Neuroanatomy of Halobiotus crispae (Eutardigrada: Hypsibiidae): Tardigrade brain structure supports the clade panarthropoda". Journal of Morphology 273 (11): 1227–1245. doi:10.1002/jmor.20054. PMID 22806919. 
  13. ↑ 13.0 13.1 13.2 Rota-Stabelli, Omar; Daley, Allison C.; Pisani, Davide (2013-03-04). "Molecular Timetrees Reveal a Cambrian Colonization of Land and a New Scenario for Ecdysozoan Evolution" (in English). Current Biology 23 (5): 392–398. doi:10.1016/j.cub.2013.01.026. ISSN 0960-9822. PMID 23375891. 
  14. ↑ 14.0 14.1 14.2 Marlétaz, Ferdinand; Peijnenburg, Katja T.C.A.; Goto, Taichiro; Satoh, Noriyuki; Rokhsar, Daniel S. (2019-01-21). "A New Spiralian Phylogeny Places the Enigmatic Arrow Worms among Gnathiferans" (in English). Current Biology 29 (2): 312–318.e3. doi:10.1016/j.cub.2018.11.042. ISSN 0960-9822. PMID 30639106. 
  15. ↑ 15.0 15.1 Ou, Qiang (2012). "Cambrian lobopodians and extant onychophorans provide new insights into early cephalization in Panarthropoda". Nature Communications 3: 1261. doi:10.1038/ncomms2272. PMID 23232391. Bibcode: 2012NatCo...3.1261O. 
  16. ↑ Giribet, Gonzalo; Edgecombe, Gregory D. (2017). "Current Understanding of Ecdysozoa and its Internal Phylogenetic Relationships". Integrative and Comparative Biology 57 (3): 455–466. doi:10.1093/icb/icx072. ISSN 1557-7023. PMID 28957525. http://nrs.harvard.edu/urn-3:HUL.InstRepos:34253786. 
  17. ↑ Laumer, Christopher E.; Fernández, Rosa; Lemer, Sarah; Combosch, David; Kocot, Kevin M.; Riesgo, Ana; Andrade, Sónia C. S.; Sterrer, Wolfgang et al. (2019-07-10). "Revisiting metazoan phylogeny with genomic sampling of all phyla". Proceedings of the Royal Society B: Biological Sciences 286 (1906): 20190831. doi:10.1098/rspb.2019.0831. PMID 31288696. 
  18. ↑ Smythe, Ashleigh B.; Holovachov, Oleksandr; Kocot, Kevin M. (2019). "Improved phylogenomic sampling of free-living nematodes enhances resolution of higher-level nematode phylogeny". BMC Evolutionary Biology 19 (1): 121. doi:10.1186/s12862-019-1444-x. ISSN 1471-2148. PMID 31195978. 
  19. ↑ Aguinaldo, Anna Marie A.; Turbeville, James M.; Linford, Lawrence S.; Rivera, Maria C.; Garey, James R.; Raff, Rudolf A.; Lake, James A. (1997). "Evidence for a clade of nematodes, arthropods and other moulting animals" (in en). Nature 387 (6632): 489–493. doi:10.1038/387489a0. ISSN 1476-4687. PMID 9168109. Bibcode: 1997Natur.387R.489A. https://www.nature.com/articles/387489a0. 
  20. ↑ 20.0 20.1 Adoutte, André; Balavoine, Guillaume; Lartillot, Nicolas; Lespinet, Olivier; Prud'homme, Benjamin; de Rosa, Renaud (2000-04-25). "The new animal phylogeny: Reliability and implications" (in en). Proceedings of the National Academy of Sciences 97 (9): 4453–4456. doi:10.1073/pnas.97.9.4453. ISSN 0027-8424. PMID 10781043. Bibcode: 2000PNAS...97.4453A. 
  21. ↑ Seaver, Elaine C.; Kaneshige, Lori M. (2006-01-01). "Expression of 'segmentation' genes during larval and juvenile development in the polychaetes Capitella sp. I and H. elegans" (in en). Developmental Biology 289 (1): 179–194. doi:10.1016/j.ydbio.2005.10.025. ISSN 0012-1606. PMID 16330020. 
  22. ↑ Smith, Frank W.; Goldstein, Bob (2017-05-01). "Segmentation in Tardigrada and diversification of segmental patterns in Panarthropoda" (in en). Arthropod Structure & Development. Evolution of Segmentation 46 (3): 328–340. doi:10.1016/j.asd.2016.10.005. ISSN 1467-8039. PMID 27725256. https://www.sciencedirect.com/science/article/pii/S1467803916301487. 
  23. ↑ Hyun Ryu, Shi; Lee, Jimin; Jang, Kuem-Hee; Hwa Choi, Eun; Ju Park, Shin; Chang, Cheon; Kim, Won; Hwang, Ui Wook (2008-01-01). "Partial mitochondrial gene arrangements support a close relationship between Tardigrada and Arthropoda". Molecules and cells 24: 351–7.
  24. ↑ Budd, Graham E. (2001-01-01). "Tardigrades as 'Stem-Group Arthropods': The Evidence from the Cambrian Fauna" (in en). Zoologischer Anzeiger - A Journal of Comparative Zoology 240 (3): 265–279. doi:10.1078/0044-5231-00034. ISSN 0044-5231. https://www.sciencedirect.com/science/article/pii/S004452310470025X. 
  25. ↑ Mayer, Georg; Martin, Christine; Rüdiger, Jan; Kauschke, Susann; Stevenson, Paul A.; Poprawa, Izabela; Hohberg, Karin; Schill, Ralph O. et al. (2013-10-24). "Selective neuronal staining in tardigrades and onychophorans provides insights into the evolution of segmental ganglia in panarthropods". BMC Evolutionary Biology 13 (1): 230. doi:10.1186/1471-2148-13-230. ISSN 1471-2148. PMID 24152256. 
  26. ↑ Martin, Christine; Mayer, Georg (2015-08-25). "Insights into the segmental identity of post-oral commissures and pharyngeal nerves in Onychophora based on retrograde fills". BMC Neuroscience 16 (1): 53. doi:10.1186/s12868-015-0191-1. ISSN 1471-2202. PMID 26303946. 
  27. ↑ 27.0 27.1 27.2 27.3 27.4 Yang, Jie; Ortega-Hernández, Javier; Butterfield, Nicholas J.; Liu, Yu; Boyan, George S.; Hou, Jin-bo; Lan, Tian; Zhang, Xi-guang (2016-03-15). "Fuxianhuiid ventral nerve cord and early nervous system evolution in Panarthropoda" (in en). Proceedings of the National Academy of Sciences 113 (11): 2988–2993. doi:10.1073/pnas.1522434113. ISSN 0027-8424. PMID 26933218. Bibcode: 2016PNAS..113.2988Y. 
  28. ↑ 28.0 28.1 Ma, Xiaoya; Edgecombe, Gregory D.; Legg, David A.; Hou, Xianguang (2013). "The morphology and phylogenetic position of the Cambrian lobopodian Diania cactiformis". Journal of Systematic Palaeontology 12 (4): 445–457. doi:10.1080/14772019.2013.770418. ISSN 1477-2019. https://www.researchgate.net/publication/247777382. 
  29. ↑ 29.0 29.1 29.2 29.3 Smith, Martin R.; Caron, Jean-Bernard (2015). "Hallucigenia's head and the pharyngeal armature of early ecdysozoans" (in en). Nature 523 (7558): 75–78. doi:10.1038/nature14573. ISSN 1476-4687. PMID 26106857. Bibcode: 2015Natur.523...75S. https://www.nature.com/articles/nature14573. 
  30. ↑ 30.0 30.1 30.2 30.3 Yang, Jie; Ortega-Hernández, Javier; Gerber, Sylvain; Butterfield, Nicholas J.; Hou, Jin-bo; Lan, Tian; Zhang, Xi-guang (2015-07-14). "A superarmored lobopodian from the Cambrian of China and early disparity in the evolution of Onychophora". Proceedings of the National Academy of Sciences of the United States of America 112 (28): 8678–8683. doi:10.1073/pnas.1505596112. ISSN 0027-8424. PMID 26124122. Bibcode: 2015PNAS..112.8678Y. 
  31. ↑ 31.0 31.1 31.2 31.3 Murdock, Duncan J. E.; Gabbott, Sarah E.; Purnell, Mark A. (2016-01-22). "The impact of taphonomic data on phylogenetic resolution: Helenodora inopinata (Carboniferous, Mazon Creek Lagerstätte) and the onychophoran stem lineage". BMC Evolutionary Biology 16: 19. doi:10.1186/s12862-016-0582-7. ISSN 1471-2148. PMID 26801389. 
  32. ↑ 32.0 32.1 32.2 32.3 Zhang, Xi-Guang; Smith, Martin R.; Yang, Jie; Hou, Jin-Bo (2016). "Onychophoran-like musculature in a phosphatized Cambrian lobopodian". Biology Letters 12 (9): 20160492. doi:10.1098/rsbl.2016.0492. ISSN 1744-9561. PMID 27677816. 
  33. ↑ 33.0 33.1 33.2 33.3 Caron, Jean-Bernard; Aria, Cédric (2017-01-31). "Cambrian suspension-feeding lobopodians and the early radiation of panarthropods". BMC Evolutionary Biology 17 (1): 29. doi:10.1186/s12862-016-0858-y. ISSN 1471-2148. PMID 28137244. 
  34. ↑ 34.0 34.1 34.2 34.3 Siveter, Derek J.; Briggs, Derek E. G.; Siveter, David J.; Sutton, Mark D.; Legg, David (2018-08-08). "A three-dimensionally preserved lobopodian from the Herefordshire (Silurian) Lagerstätte, UK". Royal Society Open Science 5 (8): 172101. doi:10.1098/rsos.172101. PMID 30224988. 
  35. ↑ 35.0 35.1 35.2 35.3 Howard, Richard J.; Hou, Xianguang; Edgecombe, Gregory D.; Salge, Tobias; Shi, Xiaomei; Ma, Xiaoya (2020-04-20). "A Tube-Dwelling Early Cambrian Lobopodian" (in English). Current Biology 30 (8): 1529–1536.e2. doi:10.1016/j.cub.2020.01.075. ISSN 0960-9822. PMID 32109391. 
  36. ↑ 36.0 36.1 36.2 Caron, Jean-Bernard; Aria, Cédric (2020). "The Collins' monster, a spinous suspension-feeding lobopodian from the Cambrian Burgess Shale of British Columbia" (in en). Palaeontology 63 (6): 979–994. doi:10.1111/pala.12499. ISSN 1475-4983. https://onlinelibrary.wiley.com/doi/abs/10.1111/pala.12499. 
  37. ↑ Poinar, George; Nelson, Diane R. (2019). "A new microinvertebrate with features of mites and tardigrades in Dominican amber". Invertebrate Biology 138 (4): e12265. doi:10.1111/ivb.12265. ISSN 1744-7410. 

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