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
Synonyms

Canadaspidida Novozhilov in Orlov, 1960

Hymenocarina is an order of extinct 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 some species.

Contents

Taxonomy

Hymenocarines are characterized by the combination of following characters: bivalved, convex carapace covering cephalothoracic region; cephalothorax bearing multisegmented antennules and rounded mandibles, alongside post-maxillular limbs with spiny, subdivided basis and endopods with well-developed terminal claws; absence of appendages between antennules and mandibles; median sclerite and lobate protrusions located between compound eyes; posterior tagma (abdomen) with ring-like segments and terminated by a pair of well-developed caudal rami. [2] [1]

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. [3] [4] [5] [6] [7] [8] [9] [10] They later became widely accepted as mandibulates (jawed arthropods) after the discovery of their mandible-bearing mouthparts in late 2010s. [2] [1] [11] [12] Since then, most phylogenetic analysis suggest hymenocarines represent part of the mandibulate stem-group, [2] [13] [1] [14] [15] [16] [17] [18] with some results suggest a rather crownward position such as stem-pancrustaceans, [1] [19] [20] [18] stem-myriapods, [1] stem-hexapods [16] or somewhere in-between the former taxa. [21]

Several subgroups within the order are recognised, including Waptiidae [1] and Protocarididae. [2] The internal relationships of Hymenocarina are unstable, and it is unclear whether the group is monophyletic or paraphyletic. [22] [23]

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. [17] [24] [25] [26] Chuandianella a bivalved arthropod morphologically similar to Waptia and long thought to be closely related [1] [21] 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. [24]

Diversity

The group was very diverse in shape, with some forms like Waptia somewhat resembling shrimp, [1] and others like Odaraia having a large carapace and trifurcate tail. [27] The appendages showing various degrees of specialization across the group, ranging from the feathery gills of Waptia [1] 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, [28] while largest Balhuticaris reached 24.5 cm (9.6 in) long. [23] Hymenocarines are thought to have been ecologically diverse, with various forms occupying scavenging, predatory, deposit feeding and suspension feeding niches. [23]

Related Research Articles

<i>Canadaspis</i> Extinct genus of arthropods

Canadaspis is an extinct genus of bivalved Cambrian arthropod, known from North America and China. They are thought to have been benthic feeders that moved mainly by walking and possibly used its biramous appendages to stir mud in search of food. They have been placed within the Hymenocarina, which includes other bivalved Cambrian arthropods.

<i>Waptia</i> Cambrian arthropod

Waptia is an extinct genus of arthropod from the Middle Cambrian of North America. It grew to a length of 6.65 cm (3 in), and had a large bivalved carapace and a segmented body terminating into a pair of tail flaps. It was an active swimmer and likely a predator of soft-bodied prey. It is also one of the oldest animals with direct evidence of brood care. Waptia fieldensis is the only species classified under the genus Waptia, and is known from the Burgess Shale Lagerstätte of British Columbia, Canada. Specimens of Waptia are also known from the Spence Shale of Utah, United States.

<i>Parapeytoia</i> Extinct genus of arthropods

Parapeytoia is a genus of Cambrian arthropod. The type and only described species is Parapeytoia yunnanensis, lived over 518 million years ago in the Maotianshan shales of Yunnan, China. Unidentified fossils from the same genus also had been discovered from the nearby Wulongqing Formation.

<span class="mw-page-title-main">Megacheira</span> Extinct class of arthropods

Megacheira is an extinct class of predatory arthropods defined by their possession of spined "great appendages". Their taxonomic position is controversial, with studies either considering them stem-group euarthropods, or stem-group chelicerates. The homology of the great appendages to the cephalic appendages of other arthropods is also controversial. Uncontested members of the group were present in marine environments worldwide from the lower to middle Cambrian.

<i>Plenocaris</i> Extinct genus of arthropods

Plenocaris plena is a Cambrian arthropod with a bivalved carapace, and is known from the Burgess shale and Chengjiang. Originally described as a species of Yohoia by Walcott in 1912, it was placed into its own genus in 1974.

<i>Chuandianella</i> Extinct genus of Cambrian animals

Chuandianella ovata is an extinct bivalved arthropod that lived during Cambrian Stage 3 of the Early Cambrian. It is the only species classified under the genus Chuandianella. Its fossils were recovered from the Chengjiang Biota in Yunnan, China.

<i>Clypecaris</i> Extinct genus of arthropods

Clypecaris is genus of bivalved Cambrian arthropod known from the Chengjiang biota of Yunnan, China. The genus was initially described for the type species C. pteroidea by Hou, 1999. A second species C. serrata was described by Yang et al. in 2016. The species are primarily distinguished by the presence of a serrated edge on the front of the carapace of C. serrata. C. serrata is noted for the modification of an anterior pair of limbs into spined grasping appendages, indicating a predatory lifestyle. It is unknown whether a similar structure was present in C. pteroidea.Clypecaris is considered to likely be a member or a close relative of Hymenocarina, and is closely related to Perspicaris. As well as to Ercaicunia.

<i>Ercaicunia</i> Extinct genus of arthropods

Ercaicunia is genus of bivalved Cambrian arthropod from the Chengjiang biota of Yunnan, China. It contains a single species, E. multinodosa that was described by Luo et al. in 1999. The total length of the body ranges from 8 to 11 millimetres. The bivalved carapace covered about a third of the total body-length, and has up to six serrations on its forward edge. The head has a pair of large uniramous antennae, as well as a smaller pair of secondary antennae, as well as pair of mandibles and maxillae. The trunk has 16 pairs of biramous appendages. Specimens were CT scanned in 2019, which suggested it to be a stem-group crustacean. Other subsequent studies have recovered it as a member of Hymenocarina, which contains other bivalved Cambrian arthropods.

<i>Haikoucaris</i> Extinct genus of arthropods

Haikoucaris is a genus of megacheiran arthropod that contains the single species Haikoucaris ercaiensis. It was discovered in the Cambrian Chengjiang biota of China.

<span class="mw-page-title-main">Pectocaris</span> Extinct genus of crustaceans

Pectocaris is an extinct genus of bivalved arthropods from the Cambrian Maotianshan Shales, Yunnan Province of China. There are currently three known species within the genus.

<span class="mw-page-title-main">Radiodonta</span> Extinct order of basal arthropods

Radiodonta is an extinct order of stem-group arthropods that was successful worldwide during the Cambrian period. They may be referred to as radiodonts, radiodontans, radiodontids, anomalocarids, or anomalocaridids, although the last two originally refer to the family Anomalocarididae, which previously included all species of this order but is now restricted to only a few species. Radiodonts are distinguished by their distinctive frontal appendages, which are morphologically diverse and used for a variety of functions. Radiodonts included the earliest large predators known, but they also included sediment sifters and filter feeders. Some of the most famous species of radiodonts are the Cambrian taxa Anomalocaris canadensis, Hurdia victoria, Peytoia nathorsti, Titanokorys gainessii, Cambroraster falcatus and Amplectobelua symbrachiata, the Ordovician Aegirocassis benmoulai and the Devonian Schinderhannes bartelsi.

<span class="mw-page-title-main">Marble Canyon (Canadian Rockies)</span>

Marble Canyon is a canyon surrounding Tokumm Creek just above its confluence with the Vermilion River, at the north end of Kootenay National Park in the Canadian Rockies of British Columbia, Canada. South of Marble Canyon on Highway 93 is Numa Falls on the Vermillion River.

<i>Cucumericrus</i> Extinct genus of arthropod

Cucumericrus ("cucumber-leg") is an extinct genus of stem-arthropod. The type and only species is Cucumericrus decoratus, with fossils discovered from the Maotianshan Shales of Yunnan, China.

<i>Kootenichela</i> Extinct genus of arthropods

Kootenichela deppi is an extinct arthropod described from the Middle Cambrian of the Kootenay National Park, Canada. It is originally considered to be a member of "great appendage arthropods", although subsequent studies questioned its affinity. Kootenichela appears to be the sister taxon of Worthenella, from cladistic analysis.

<i>Tokummia</i> Extinct species of arthropod

Tokummia is a genus of fossil hymenocarine arthropod, known only by one species, Tokummiakatalepsis, from the middle Cambrian Burgess Shale as found in a quarry in Marble Canyon in Canada.

<i>Fibulacaris</i> Genus of fossil Arthropod

Fibulacaris is a monotypic genus of fossil arthropod known only by one species, Fibulacaris nereidis, discovered from the Cambrian Burgess Shale of Canada. It was characterized by a bivalved carapace with an inverted rostrum, sandwiching the slender body with stalked eyes and homonomous appendages. It was probably an actively swimming filter feeder and possibly swam upside-down like some branchiopod crustaceans and horseshoe crabs. Phylogenetic analysis suggest it was a relative or member of Hymenocarina, which contains other bivalved arthropods.

<i>Balhuticaris</i> Extinct genus of arthropods

Balhuticaris is a genus of extinct bivalved hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. This extremely multisegmented arthropod is the largest member of the group, and it was even one of the largest animals of the Cambrian, with individuals reaching lengths of 245 mm (9 in). Fossils of this animal suggests that gigantism occurred in more groups of Arthropoda than had been previously thought. It also presents the possibility that bivalved arthropods were very diverse, and filled in a lot of ecological niches.

<i>Nereocaris</i> Genus of bivalved hymenocarine arthropod

Nereocaris is an extinct genus of bivalved hymenocarine arthropod that lived in the Cambrian aged Burgess Shale in what is now British Columbia around 506 million years ago. Two species are known.

<i>Jugatacaris</i> Extinct genus of bivalved arthropod

Jugatacaris is an extinct genus of bivalved arthropod known from a single species, Jugatacaris agilis found in the Cambrian Stage 3 aged Chengjiang biota of Yunnan, China. The carapace is around 28 to 37 millimetres in length, with a pronounced ridge at the top of the carapace separating the two valves, which formed a fin-like structure raised above the carapace. The head has a pair of stalked eyes, as well as a dumbbell shaped medial eye between them. The head also bore a pair of mandibles as well as at least one and possibly two pairs of antennules. The trunk had up to 65 segments, each with biramous appendages. The appendages had thin endopods with 30 podomeres, each bearing a spiny endite, with the endopods ending with a terminal claw. The appendages also had overlapping flap-like exopods, which are elongated, being at maximum eight times as long as they are wide, which on their posterior edge are covered with setae. The trunk ended with a forked tail. It was likely an actively swimming filter feeder, using its constantly beating appendages to sift food from the water column, which was then passed forward along the U-shaped food groove between the appendage pairs towards the mouth. While initially placed as a crustaceanomorph, later studies considered to be a member of Hymenocarina, which contains numerous other similar bivalved Cambrian arthropods.

<span class="mw-page-title-main">Isoxyida</span> Extinct order of arthropods

Isoxyids are members of the order Isoxyida and the family Isoxyidae, a group of basal arthropods that existed during the Cambrian period. It contains two genera, Isoxys, with 20 species found worldwide, and Surusicaris known from a single species found in the Burgess Shale of Canada. They are distinguished by their bivalved carapaces and pair of upward curving grasping frontal appendages.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 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. doi: 10.1098/rsos.172206 . PMC   6030330 . PMID   30110460.
  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. doi:10.1038/nature22080. ISSN   0028-0836. PMID   28445464. S2CID   4454526.
  3. ↑ 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.
  4. ↑ Legg, David (2013). "Multi-Segmented Arthropods from the Middle Cambrian of British Columbia (Canada)". Journal of Paleontology. 87 (3): 493–501. doi:10.1666/12-112.1. ISSN   0022-3360. S2CID   86725173.
  5. ↑ 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. doi:10.1111/let.12032. ISSN   0024-1164.
  6. ↑ 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. doi: 10.1038/ncomms3485 . ISSN   2041-1723. PMID   24077329.
  7. ↑ 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.
  8. ↑ 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. doi: 10.1016/j.asd.2016.10.011 . ISSN   1467-8039. PMID   27989966.
  9. ↑ 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.
  10. ↑ 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.
  11. ↑ Giribet, Gonzalo; Edgecombe, Gregory D. (2019-06-17). "The Phylogeny and Evolutionary History of Arthropods". Current Biology. 29 (12): R592–R602. doi: 10.1016/j.cub.2019.04.057 . ISSN   0960-9822. PMID   31211983. S2CID   189926344.
  12. ↑ 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.
  13. ↑ Aria, Cédric; Caron, Jean-Bernard (2017-12-21). "Mandibulate convergence in an armoured Cambrian stem chelicerate". BMC Evolutionary Biology. 17 (1): 261. doi: 10.1186/s12862-017-1088-7 . ISSN   1471-2148. PMC   5738823 . PMID   29262772.
  14. ↑ 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. doi:10.1038/s41586-020-2883-7. PMID   33149303. S2CID   226248177.
  15. ↑ 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). doi:10.1144/jgs2020-246. ISSN   0016-7649. S2CID   233952670.
  16. 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. doi:10.1111/pala.12534. ISSN   0031-0239. S2CID   234812878.
  17. 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. doi:10.1002/spp2.1366. ISSN   2056-2799. S2CID   236284813.
  18. 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.
  19. ↑ Aria, Cédric; Caron, Jean-Bernard (2019). "A middle Cambrian arthropod with chelicerae and proto-book gills". Nature. 573 (7775): 586–589. doi:10.1038/s41586-019-1525-4. ISSN   1476-4687. PMID   31511691. S2CID   202550431.
  20. ↑ 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. doi: 10.1186/s12862-019-1560-7 . ISSN   1471-2148. PMC   6950928 . PMID   31914921.
  21. 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. doi: 10.1126/sciadv.aaz3376 . PMC   7190318 . PMID   32426476. S2CID   218675348.
  22. ↑ 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. doi:10.1098/rsos.220933. ISSN   2054-5703. PMC   9727825 . PMID   36483757.
  23. 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. doi: 10.1016/j.isci.2022.104675 . ISSN   2589-0042. PMC   9283658 . PMID   35845166. S2CID   250057171.
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  25. ↑ 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. doi:10.1098/rsos.191350. PMC   6894550 . PMID   31827867.
  26. ↑ 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.
  27. ↑ 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. JSTOR   2395403.
  28. ↑ 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. doi:10.1098/rsos.191350. PMC   6894550 . PMID   31827867.


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