Hymenocarina

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Hymenocarina
Temporal range: Cambrian
Waptia diagram.jpg
Diagram of Waptia
Canadaspis perfecta USNM 57703.jpg
Fossil of Canadaspis perfecta
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Clade: Mandibulata
Order:† Hymenocarina
Størmer, 1944
Genera

See text

Synonyms

Canadaspidida Novozhilov in Orlov, 1960

Hymenocarina is an order of extinct marine arthropods known from the Cambrian. They possess bivalved carapaces, typically with exposed posteriors. Members of the group are morphologically diverse and had a variety of ecologies, including as filter feeders and as predators. Recent research has generally considered them to be stem or crown group members of Mandibulata, due to the presence of mandibles in well-preserved species.

Contents

Taxonomy

Hymenocarines are characterized by the combination of the following characters: bivalved, convex carapace covering cephalothoracic (combined head and thorax) region; cephalothorax bearing multisegmented antennae (though as an exception antennae are absent in Odaraia [1] ) and rounded mandibles and likely maxillae, post maxillae limbs with spiny, subdivided basis and endopods (lower, leg-like parts) with well-developed terminal claws; absence of appendages between antennae and mandibles; median sclerite and lobate protrusions located between compound eyes; posterior tagma (abdomen) with ring-like segments with the posterior of the body ending with pair of well-developed caudal rami (often in the form of tail flukes). [2] [3]

Based on the interpretation of simple head region that possess only a few segments and appendages, hymenocarine taxa were thought to be part of the upper stem-group euarthropods in early and mid 2010s. [4] [5] [6] [7] [8] [9] [10] [11] They later became widely accepted as mandibulates (jawed arthropods) after the discovery of their mandible-bearing mouthparts in late 2010s. [2] [3] [12] [13] Since then, most phylogenetic analysis suggest hymenocarines represent part of the mandibulate stem-group, [2] [14] [3] [15] [16] [17] [18] [19] with some results suggest a rather crownward position such as stem-pancrustaceans, [3] [20] [21] [19] stem-myriapods, [3] stem-hexapods [17] or somewhere in-between the former taxa. [22]

Several subgroups within the order are recognised, including Waptiidae [3] and Protocarididae. [2] The internal relationships of Hymenocarina are unstable, and it is unclear whether the group is monophyletic or paraphyletic. [23] [24]

Cambrian bivalved arthropods are now recognised to be a polyphyletic group, with other groups of bivalved arthropods such as the Isoxyida, Bradoriida and Phosphatocopina only distantly related to Hymenocarina. [18] [25] [26] [27] Chuandianella a bivalved arthropod morphologically similar to Waptia and long thought to be closely related [3] [22] was reinterpreted as a non-hymenocarine euarthropod based on a restudy published in 2022, which found that it definitely lacked mandibles, characteristic of true hymenocarines. [25]

Diversity

The group was very diverse in shape, with some forms like Waptia somewhat resembling shrimp, [3] and others like Odaraia having a large carapace and trifurcate tail. [28] The appendages showing various degrees of specialization across the group, ranging from the feathery gills of Waptia [3] to the robust claws of Tokummia . [2] They also had a wide range of sizes with some like Fibulacaris reaching a length of up to 2 cm (0.79 in) long, [29] while largest Balhuticaris reached 24.5 cm (9.6 in) long. [24] Hymenocarines are thought to have been ecologically diverse, with various forms occupying scavenging, predatory, deposit feeding and suspension feeding niches. [24]

Cladogram of Hymenocarina, following Izquierdo-López and Caron, (2024): [1]

List of genera

References

  1. 1 2 Izquierdo-López, Alejandro; Caron, Jean-Bernard (August 2024). "The Cambrian Odaraia alata and the colonization of nektonic suspension-feeding niches by early mandibulates". Proceedings of the Royal Society B: Biological Sciences. 291 (2027). doi:10.1098/rspb.2024.0622. ISSN   1471-2954. PMC   11463219 . PMID   39043240.
  2. 1 2 3 4 5 Aria, Cédric; Caron, Jean-Bernard (2017-04-26). "Burgess Shale fossils illustrate the origin of the mandibulate body plan". Nature. 545 (7652): 89–92. Bibcode:2017Natur.545...89A. doi:10.1038/nature22080. ISSN   0028-0836. PMID   28445464. S2CID   4454526.
  3. 1 2 3 4 5 6 7 8 9 Vannier, Jean; Aria, Cédric; Taylor, Rod S.; Caron, Jean-Bernard (2018). "Waptia fieldensis Walcott, a mandibulate arthropod from the middle Cambrian Burgess Shale". Royal Society Open Science. 5 (6): 172206. Bibcode:2018RSOS....572206V. doi: 10.1098/rsos.172206 . PMC   6030330 . PMID   30110460.
  4. ↑ Legg, David A.; Sutton, Mark D.; Edgecombe, Gregory D.; Caron, Jean-Bernard (2012-12-07). "Cambrian bivalved arthropod reveals origin of arthrodization". Proceedings of the Royal Society B: Biological Sciences. 279 (1748): 4699–4704. doi:10.1098/rspb.2012.1958. PMC   3497099 . PMID   23055069.
  5. ↑ Legg, David (2013). "Multi-Segmented Arthropods from the Middle Cambrian of British Columbia (Canada)". Journal of Paleontology. 87 (3): 493–501. Bibcode:2013JPal...87..493L. doi:10.1666/12-112.1. ISSN   0022-3360. S2CID   86725173.
  6. ↑ Legg, David A.; Vannier, Jean (2013-09-16). "The affinities of the cosmopolitan arthropod Isoxys and its implications for the origin of arthropods". Lethaia. 46 (4): 540–550. Bibcode:2013Letha..46..540L. doi:10.1111/let.12032. ISSN   0024-1164.
  7. ↑ Legg, David A.; Sutton, Mark D.; Edgecombe, Gregory D. (2013-09-30). "Arthropod fossil data increase congruence of morphological and molecular phylogenies". Nature Communications. 4 (1): 2485. Bibcode:2013NatCo...4.2485L. doi: 10.1038/ncomms3485 . ISSN   2041-1723. PMID   24077329.
  8. ↑ Ortega-Hernández, Javier (2014-12-21). "Making sense of 'lower' and 'upper' stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848". Biological Reviews. 91 (1): 255–273. doi:10.1111/brv.12168. PMID   25528950. S2CID   7751936.
  9. ↑ 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.
  10. ↑ Yang, Jie; Ortega-Hernández, Javier; Legg, David A.; Lan, Tian; Hou, Jin-bo; Zhang, Xi-guang (2018-02-01). "Early Cambrian fuxianhuiids from China reveal origin of the gnathobasic protopodite in euarthropods". Nature Communications. 9 (1): 470. Bibcode:2018NatCo...9..470Y. doi:10.1038/s41467-017-02754-z. ISSN   2041-1723. PMC   5794847 . PMID   29391458.
  11. ↑ Daley, Allison C.; Antcliffe, Jonathan B.; Drage, Harriet B.; Pates, Stephen (2018-05-22). "Early fossil record of Euarthropoda and the Cambrian Explosion". Proceedings of the National Academy of Sciences. 115 (21): 5323–5331. Bibcode:2018PNAS..115.5323D. doi: 10.1073/pnas.1719962115 . ISSN   0027-8424. PMC   6003487 . PMID   29784780.
  12. ↑ 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.
  13. ↑ 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.
  14. ↑ Aria, Cédric; Caron, Jean-Bernard (2017-12-21). "Mandibulate convergence in an armoured Cambrian stem chelicerate". BMC Evolutionary Biology. 17 (1): 261. Bibcode:2017BMCEE..17..261A. doi: 10.1186/s12862-017-1088-7 . ISSN   1471-2148. PMC   5738823 . PMID   29262772.
  15. ↑ Zeng, Han; Zhao, Fangchen; Niu, Kecheng; Zhu, Maoyan; Huang, Diying (2020-12-03). "An early Cambrian euarthropod with radiodont-like raptorial appendages". Nature. 588 (7836): 101–105. Bibcode:2020Natur.588..101Z. doi:10.1038/s41586-020-2883-7. PMID   33149303. S2CID   226248177.
  16. ↑ Aria, Cédric; Zhao, Fangchen; Zhu, Maoyan (2021-09-01). "Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda". Journal of the Geological Society. 178 (5). Bibcode:2021JGSoc.178..246A. doi:10.1144/jgs2020-246. ISSN   0016-7649. S2CID   233952670.
  17. 1 2 Anderson, Evan P.; Schiffbauer, James D.; Jacquet, Sarah M.; Lamsdell, James C.; Kluessendorf, Joanne; Mikulic, Donald G. (2021-04-19). "Stranger than a scorpion: a reassessment of Parioscorpio venator, a problematic arthropod from the Llandoverian Waukesha Lagerstätte". Palaeontology. 64 (3): 429–474. Bibcode:2021Palgy..64..429A. doi:10.1111/pala.12534. ISSN   0031-0239. S2CID   234812878.
  18. 1 2 Izquierdo-López, Alejandro; Caron, Jean-Bernard (2021-06-15). "A Burgess Shale mandibulate arthropod with a pygidium: a case of convergent evolution". Papers in Palaeontology. 7 (4): 1877–1894. Bibcode:2021PPal....7.1877I. doi:10.1002/spp2.1366. ISSN   2056-2799. S2CID   236284813.
  19. 1 2 O'Flynn, Robert; Williams, Mark; Yu, Mengxiao; Harvey, Thomas; Liu, Yu (2022-02-11). "A new euarthropod with large frontal appendages from the early Cambrian Chengjiang biota". Palaeontologia Electronica. 25: a6. doi: 10.26879/1167 . S2CID   246779634.
  20. ↑ Aria, Cédric; Caron, Jean-Bernard (2019). "A middle Cambrian arthropod with chelicerae and proto-book gills". Nature. 573 (7775): 586–589. Bibcode:2019Natur.573..586A. doi:10.1038/s41586-019-1525-4. ISSN   1476-4687. PMID   31511691. S2CID   202550431.
  21. ↑ Aria, Cédric; Zhao, Fangchen; Zeng, Han; Guo, Jin; Zhu, Maoyan (2020-01-08). "Fossils from South China redefine the ancestral euarthropod body plan". BMC Evolutionary Biology. 20 (1): 4. Bibcode:2020BMCEE..20....4A. doi: 10.1186/s12862-019-1560-7 . ISSN   1471-2148. PMC   6950928 . PMID   31914921.
  22. 1 2 Ou, Qiang; Vannier, Jean; Yang, Xianfeng; Chen, Aiiln; Huijuan, Mai; Shu, Degan; Han, Jian; Fu, Dongjing; Wang, Rong; Mayer, Georg (2020-04-29). "Evolutionary trade-off in reproduction of Cambrian arthropods". Science Advances. 6 (18): 3376–3405. Bibcode:2020SciA....6.3376O. doi: 10.1126/sciadv.aaz3376 . PMC   7190318 . PMID   32426476. S2CID   218675348.
  23. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (December 2022). "The problematic Cambrian arthropod Tuzoia and the origin of mandibulates revisited". Royal Society Open Science. 9 (12): 220933. Bibcode:2022RSOS....920933I. doi:10.1098/rsos.220933. ISSN   2054-5703. PMC   9727825 . PMID   36483757.
  24. 1 2 3 Izquierdo-López, Alejandro; Caron, Jean-Bernard (2022-07-15). "Extreme multisegmentation in a giant bivalved arthropod from the Cambrian Burgess Shale". iScience. 25 (7): 104675. Bibcode:2022iSci...25j4675I. doi: 10.1016/j.isci.2022.104675 . ISSN   2589-0042. PMC   9283658 . PMID   35845166. S2CID   250057171.
  25. 1 2 Zhai, Dayou; Williams, Mark; Siveter, David J.; Siveter, Derek J.; Harvey, Thomas H. P.; Sansom, Robert S.; Mai, Huijuan; Zhou, Runqing; Hou, Xianguang (2022-02-22). "Chuandianella ovata: An early Cambrian stem euarthropod with feather-like appendages". Palaeontologia Electronica. 25 (1): 1–22. doi: 10.26879/1172 . ISSN   1094-8074. S2CID   247123967.
  26. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (2019). "A possible case of inverted lifestyle in a new bivalved arthropod from the Burgess Shale". Royal Society Open Science. 6 (11): 191350. Bibcode:2019RSOS....691350I. doi:10.1098/rsos.191350. PMC   6894550 . PMID   31827867.
  27. ↑ Zeng, Han; Zhao, Fang-Chen; Yin, Zong-Jun; Zhu, Mao-Yan (September 2021). "A new early Cambrian bivalved euarthropod from Yunnan, China and general interspecific morphological and size variations in Cambrian hymenocarines". Palaeoworld. 30 (3): 387–397. doi:10.1016/j.palwor.2020.09.002.
  28. ↑ Briggs, D. E. G. (1981). "The Arthropod Odaraia alata Walcott, Middle Cambrian, Burgess Shale, British Columbia". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 291 (1056): 541–582. Bibcode:1981RSPTB.291..541B. doi:10.1098/rstb.1981.0007. JSTOR   2395403.
  29. ↑ Izquierdo-López, Alejandro; Caron, Jean-Bernard (2019). "A possible case of inverted lifestyle in a new bivalved arthropod from the Burgess Shale". Royal Society Open Science. 6 (11): 191350. Bibcode:2019RSOS....691350I. doi:10.1098/rsos.191350. PMC   6894550 . PMID   31827867.
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