Panarthropoda
Panarthropoda Temporal range: Cambrian - Recent PreꞒ Ꞓ O S D C P T J K Pg N | |
---|---|
![]() | |
Panarthropods include onychophorans such as Peripatopsis and arthropods such as polydesmid millipedes | |
Scientific classification ![]() | |
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 |
| ||||||||||||||||||
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
| |||||||||||||||||||||||||||
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 | |
Panarthropoda | |
Panarthropoda |
| ||||||
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
References
- ^ Rota-Stabelli, Omar; Daley, Allison C.; Pisani, Davide (March 2013). "Molecular Timetrees Reveal a Cambrian Colonization of Land and a New Scenario for Ecdysozoan Evolution". Current Biology. 23 (5): 392–398. Bibcode:2013CBio...23..392R. doi:10.1016/j.cub.2013.01.026. PMID 23375891.
- ^ Chen, Zhe; Chen, Xiang; Zhou, Chuanming; Yuan, Xunlai; Xiao, Shuhai (6 June 2018). "Late Ediacaran trackways produced by bilaterian animals with paired appendages". Science Advances. 4 (6): eaao6691. Bibcode:2018SciA....4.6691C. doi:10.1126/sciadv.aao6691. hdl:10919/84444. PMC 5990303. PMID 29881773.
- ^ 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. PMC 2614232. PMID 18192181.
- ^ a b c d e f g h Smith, Martin R.; Ortega-Hernández, Javier (2014). "Hallucigenia's onychophoran-like claws and the case for Tactopoda". Nature. 514 (7522): 363–366. Bibcode:2014Natur.514..363S. doi:10.1038/nature13576. PMID 25132546. S2CID 205239797.
- ^ a b c d e f g 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. S2CID 7751936.
- ^ a b c d e f g Ortega-Hernández, Javier; Janssen, Ralf; Budd, Graham E. (2017-05-01). "Origin and evolution of the panarthropod head – A palaeobiological and developmental perspective". Arthropod Structure & Development. Evolution of Segmentation. 46 (3): 354–379. Bibcode:2017ArtSD..46..354O. doi:10.1016/j.asd.2016.10.011. ISSN 1467-8039. PMID 27989966.
- ^ a b Giribet, Gonzalo; Edgecombe, Gregory D. (2019-06-17). "The Phylogeny and Evolutionary History of Arthropods". Current Biology. 29 (12): R592–R602. Bibcode:2019CBio...29.R592G. doi:10.1016/j.cub.2019.04.057. ISSN 0960-9822. PMID 31211983. S2CID 189926344.
- ^ a b c 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. PMC 6790758. PMID 31575373.
- ^ a b 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. S2CID 225478171.
- ^ a b c d Rota-Stabelli, O.; Kayal, E.; Gleeson, D.; Daub, J.; Boore, J.; Telford, M.; Pisani, D.; Blaxter, M.; Lavrov, D. (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. PMC 2998192. PMID 20624745.
- ^ Dunn, C. W.; Hejnol, A.; Matus, D. Q.; Pang, K.; Browne, W. E.; Smith, S. A.; Seaver, E.; Rouse, G. W.; Obst, M.; Edgecombe, G. D.; Sørensen, M. V.; Haddock, S. H. D.; Schmidt-Rhaesa, A.; Okusu, A.; Kristensen, R. M. B.; Wheeler, W. C.; Martindale, M. Q.; Giribet, G. (10 April 2008). "Broad phylogenomic sampling improves resolution of the animal tree of life". Nature. 452 (7188): 745–749. Bibcode:2008Natur.452..745D. doi:10.1038/nature06614. PMID 18322464. S2CID 4397099.
- ^ 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. S2CID 5260983.
- ^ a b c 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". Current Biology. 23 (5): 392–398. Bibcode:2013CBio...23..392R. doi:10.1016/j.cub.2013.01.026. ISSN 0960-9822. PMID 23375891. S2CID 2510415.
- ^ a b c 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". Current Biology. 29 (2): 312–318.e3. Bibcode:2019CBio...29E.312M. doi:10.1016/j.cub.2018.11.042. ISSN 0960-9822. PMID 30639106. S2CID 58562919.
- ^ a b Ou, Qiang (2012). "Cambrian lobopodians and extant onychophorans provide new insights into early cephalization in Panarthropoda". Nature Communications. 3: 1261. Bibcode:2012NatCo...3.1261O. doi:10.1038/ncomms2272. PMC 3535342. PMID 23232391.
- ^ 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.
- ^ Laumer, Christopher E.; Fernández, Rosa; Lemer, Sarah; Combosch, David; Kocot, Kevin M.; Riesgo, Ana; Andrade, Sónia C. S.; Sterrer, Wolfgang; Sørensen, Martin V.; Giribet, Gonzalo (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. PMC 6650721. PMID 31288696.
- ^ 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. Bibcode:2019BMCEE..19..121S. doi:10.1186/s12862-019-1444-x. ISSN 1471-2148. PMC 6567515. PMID 31195978.
- ^ 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". Nature. 387 (6632): 489–493. Bibcode:1997Natur.387R.489A. doi:10.1038/387489a0. ISSN 1476-4687. PMID 9168109. S2CID 4334033.
- ^ a b Adoutte, André; Balavoine, Guillaume; Lartillot, Nicolas; Lespinet, Olivier; Prud'homme, Benjamin; de Rosa, Renaud (2000-04-25). "The new animal phylogeny: Reliability and implications". Proceedings of the National Academy of Sciences. 97 (9): 4453–4456. Bibcode:2000PNAS...97.4453A. doi:10.1073/pnas.97.9.4453. ISSN 0027-8424. PMC 34321. PMID 10781043.
- ^ 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". Developmental Biology. 289 (1): 179–194. doi:10.1016/j.ydbio.2005.10.025. ISSN 0012-1606. PMID 16330020.
- ^ Smith, Frank W.; Goldstein, Bob (2017-05-01). "Segmentation in Tardigrada and diversification of segmental patterns in Panarthropoda". Arthropod Structure & Development. Evolution of Segmentation. 46 (3): 328–340. Bibcode:2017ArtSD..46..328S. doi:10.1016/j.asd.2016.10.005. ISSN 1467-8039. PMID 27725256.
- ^ Ryu, Shi Hyun; Lee, Ji Min; Jang, Kuem-Hee; Choi, Eun Hwa; Park, Shin Ju; Chang, Cheon Young; Kim, Won; Hwang, Ui Wook (2007-12-31). "Partial mitochondrial gene arrangements support a close relationship between Tardigrada and Arthropoda". Molecules and Cells. 24 (3): 351–357. doi:10.1016/S1016-8478(23)07350-8. ISSN 1016-8478. PMID 18182850.
- ^ Budd, Graham E. (2001-01-01). "Tardigrades as 'Stem-Group Arthropods': The Evidence from the Cambrian Fauna". Zoologischer Anzeiger - A Journal of Comparative Zoology. 240 (3): 265–279. Bibcode:2001ZooAn.240..265B. doi:10.1078/0044-5231-00034. ISSN 0044-5231.
- ^ Mayer, Georg; Martin, Christine; Rüdiger, Jan; Kauschke, Susann; Stevenson, Paul A.; Poprawa, Izabela; Hohberg, Karin; Schill, Ralph O.; Pflüger, Hans-Joachim; Schlegel, Martin (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. Bibcode:2013BMCEE..13..230M. doi:10.1186/1471-2148-13-230. ISSN 1471-2148. PMC 4015553. PMID 24152256.
- ^ 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. PMC 4549126. PMID 26303946.
- ^ a b c d e 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". Proceedings of the National Academy of Sciences. 113 (11): 2988–2993. Bibcode:2016PNAS..113.2988Y. doi:10.1073/pnas.1522434113. ISSN 0027-8424. PMC 4801254. PMID 26933218.
- ^ a b 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. S2CID 220463025.
- ^ a b c d Smith, Martin R.; Caron, Jean-Bernard (2015). "Hallucigenia's head and the pharyngeal armature of early ecdysozoans". Nature. 523 (7558): 75–78. Bibcode:2015Natur.523...75S. doi:10.1038/nature14573. ISSN 1476-4687. PMID 26106857. S2CID 205244325.
- ^ a b c d 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. Bibcode:2015PNAS..112.8678Y. doi:10.1073/pnas.1505596112. ISSN 0027-8424. PMC 4507230. PMID 26124122.
- ^ a b c d 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 (1): 19. Bibcode:2016BMCEE..16...19M. doi:10.1186/s12862-016-0582-7. ISSN 1471-2148. PMC 4722706. PMID 26801389.
- ^ a b c d 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. PMC 5046927. PMID 27677816.
- ^ a b c d Caron, Jean-Bernard; Aria, Cédric (2017-01-31). "Cambrian suspension-feeding lobopodians and the early radiation of panarthropods". BMC Evolutionary Biology. 17 (1): 29. Bibcode:2017BMCEE..17...29C. doi:10.1186/s12862-016-0858-y. ISSN 1471-2148. PMC 5282736. PMID 28137244.
- ^ a b c d 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. PMC 6124121. PMID 30224988.
- ^ a b c d Howard, Richard J.; Hou, Xianguang; Edgecombe, Gregory D.; Salge, Tobias; Shi, Xiaomei; Ma, Xiaoya (2020-04-20). "A Tube-Dwelling Early Cambrian Lobopodian". Current Biology. 30 (8): 1529–1536.e2. Bibcode:2020CBio...30E1529H. doi:10.1016/j.cub.2020.01.075. ISSN 0960-9822. PMID 32109391. S2CID 211542458.
- ^ a b c Caron, Jean-Bernard; Aria, Cédric (2020). "The Collins' monster, a spinous suspension-feeding lobopodian from the Cambrian Burgess Shale of British Columbia". Palaeontology. 63 (6): 979–994. Bibcode:2020Palgy..63..979C. doi:10.1111/pala.12499. ISSN 1475-4983. S2CID 225593728.
- ^ 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. S2CID 204157733.
- v
- t
- e
- Domain
- Archaea
- Bacteria
- Eukaryota
- (major groups
- Excavata
- Diaphoretickes
- Hacrobia
- Cryptista
- Rhizaria
- Alveolata
- Stramenopiles
- Plants
- Hacrobia
- Amorphea
- Amoebozoa
- Opisthokonta
- Animals
- Fungi
- Mesomycetozoea)
Animalia |
| ||||||||
---|---|---|---|---|---|---|---|---|---|
|
|
Major groups within phyla |
---|
- Phyla with ≥1000 extant species bolded
- Potentially dubious phyla †