Fuxianhuiida

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Fuxianhuiida
Temporal range: Cambrian Stage 3-Cambrian Stage 4 520–514  Ma
20211117 Fuxianhuiida Fuxianhuiids.png
Life restorations of Fuxianhuia (top), Alacaris , (left) and Chengjiangocaris (right)
Guangweicaris restoration.png
Guangweicaris
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Arthropoda
Clade: Mandibulata
Order: Fuxianhuiida
Bousfield, 1995
Genera

Fuxianhuiida is an extinct clade of arthropods from the Cambrian of China. All currently known species are from Cambrian Series 2 aged deposits in Yunnan Province, including the Chengjiang biota. [1] They are known from several specimens with exceptional soft-tissue preservation, including remnants of nerve and brain tissue. [2]

Contents

Description

Diagrammatic reconstruction of Alacaris, with the underside of the head of Chengjiangocaris bottom right. Alacaris restoration.jpg
Diagrammatic reconstruction of Alacaris, with the underside of the head of Chengjiangocaris bottom right.
Fossil of Fuxianhuia Chengjiang Fossil Site (53696047141).jpg
Fossil of Fuxianhuia

Fuxianhuiids reached a size of up to 15 centimetres (5.9 in). The fuxianhuiid exoskeleton was unmineralised, and the number of tergites (plates on the back associated with body segments) ranged from 15 to over 40. The cephalon (head section) was covered by a heart-shaped head shield, and contained stalked eyes connected by the anterior sclerite (a plate-like structure attached to the head), a pair of antennae, and a butterfly shaped hypostome (a plate-like structure), which covered the posterior-facing mouth. Fuxianhuiids possessed specialized appendages near the front of the body posterior to the antennae with serrated edges used for food processing, with chengjiangocardids having gnathobases (basal segments of the limbs close to where they connect with the body with sturdy, thick spines), used for processing food. To the underside of the body were attached numerous pairs of two-branched (biramous) limbs, with the lower branches (the endopods), which served as walking legs, having at least 10 segments (podomeres) and were generally short and similar in appearance to each other along the length of the body, with the frontmost legs of chenjiangocaridids ending in pronounced claws, while the upper branches (the exopods) of fuxianhuiids were flap-shaped. [2] In most fuxianhuiids, the thorax tergites narrowed posteriorly, terminating in either a swimming paddle or paired flukes with a tail spine. In members of Fuxianhuiidae the thorax was divided into two sections, the front wide opisthothorax and the posterior narrow limbless tail-like abdomen. [2]

Labelled diagram of Fuxianhuia 20211113 Fuxianhuiid morphology.png
Labelled diagram of Fuxianhuia

A number of specimens are known with exceptional soft tissue preservation, including preserved guts and neural tissue including brain anatomy, which given their historically assumed basal phylogenetic position has made them a considerable focus of paleoneuranatomy research. [3] [4] Fuxianhuiids had a straight gut tract that ran along the length of the body, with the sides of the gut tract of Fuxianhuia having pairs of lobe-shaped pockets (diverticulae) associated with each body segment. [5] The ventral nerve cord, which in arthropods functions analogously to the vertebrate spinal cord, has reported to have been found preserved in a specimen of Chengjiangocaris, [3] though the degree of certainty of this identification was later questioned. [4] The brains of fuxianhuiids, based on an exceptionally preserved specimen of Fuxianhuia are suggested to have been composed of three parts (tri-partite) and to have been relatively complex. [2] [4]

Paleobiology

Life restoration of Fuxianhuia, engaging in parental care of offspring, as inferred from the fossil record Fu et al. 2018 f14.png
Life restoration of Fuxianhuia, engaging in parental care of offspring, as inferred from the fossil record

Fuxianhuiids are interpreted as benthic (seafloor dwelling) animals. [2] Although initially interpreted as passive animals that ingested mud to feed on seafloor detritus, [5] [2] they are now considered to have probably been scavengers and possibly predators of slow moving prey such as worms. [2] The presence of gnathobases in members of Chengjiangocardidae suggests that they were capable of durophagy (crushing hard prey). [1] A specimen of Fuxianhuia was found with the remains of the trilobite Pagetia in its gut tract, indicating that it took relatively hard prey at least on occasion. [6] [1] The diverticulae in the gut tract of Fuxianhuia may indicate that it only intermittently fed. [2] As fuxianhuiids grew, new segments were added to the posterior of the body. The findings of somewhat advanced juvenile individuals of Fuxianhuia associated with adults suggests that in this taxon the juveniles were subject to extended post-hatching parental care. [7]

Taxonomy

Internal taxonomy of Fuxianhuiida per Liu et al., 2020: [8]

Fuxianhuiida

It has been suggested that Shankouia zhenghei is synonymous with Liangwangshaniabiloba, with sexual dimorphism accounting for variation between specimens. [9]

Relationship with other arthropods

Although historically suggested to be members of the arthropod stem group, [1] studies from the early 2020s onwards have supported their placement as closely related to living mandibulates, which include crustaceans, insects, and Myriapoda (the group which contains centipedes and millipedes, among others), due to the suggestion that their head feeding appendages are homologous to the mandibles of mandibulates. [10] [11] [12] [13] [14] [15] Cladogram of the position of Fuxianhuiida in relation to living and extinct arthropods after O’Flynn et al, 2023: [11]

Total group Arthropoda

"Gilled lobopodians" ( Pambdelurion , Kerygmachela ) 20191022 Kerygmachela kierkegaardi without lobopods.png

Opabinia 20191108 Opabinia regalis.png

Radiodonta (e.g Anomalocaris ) 20191203 Anomalocaris canadensis.png

Deuteropoda

Megacheira 20191020 Yohoia tenuis.png

Habeliida Habelia reconstruction from Aria & Caron 2017 modified.png

Mollisonia 20191003 Mollisonia plenovenatrix side.png

Chelicerata (horseshoe crabs, sea spiders, arachnids, etc) 20200813 Lunataspis aurora.png

"Great appendage bivalved forms" ( Occacaris , Forfexicaris )

Isoxyida Artistic reconstruction of Isoxys curvirostratus.jpg

Artiopoda (including Trilobita) Estonian Museum of Natural History - trilobite - Hydrocephalus.png

Mandibulata

Fuxianhuiida 20211117 Alacaris mirabilis.png

Myriapoda (millipedes, centipedes, etc) Scolopendra multidens Guang Xi Sheng Chan .jpg

Hymenocarina 20211025 Waptia fieldensis.png

Pancrustacea (crustaceans, insects, etc)

References

  1. 1 2 3 4 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.
  2. 1 2 3 4 5 6 7 8 Ortega-Hernández, Javier; Yang, Jie; Zhang, Xi-guang (2018-07-01). "Fuxianhuiids". Current Biology. 28 (13): R724 –R725. Bibcode:2018CBio...28.R724O. doi: 10.1016/j.cub.2018.04.042 . PMID   29990450.
  3. 1 2 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.
  4. 1 2 3 Aria, Cédric; Vannier, Jean; Park, Tae-Yoon S.; Gaines, Robert R. (March 2023). "Interpreting fossilized nervous tissues". BioEssays. 45 (3) 2200167. doi:10.1002/bies.202200167. ISSN   0265-9247. PMID   36693795.
  5. 1 2 Ortega-Hernández, Javier; Fu, Dongjing; Zhang, Xingliang; Shu, Degan (2018-02-19). "Gut glands illuminate trunk segmentation in Cambrian fuxianhuiids". Current Biology. 28 (4): R146 –R147. Bibcode:2018CBio...28.R146O. doi:10.1016/j.cub.2018.01.040. ISSN   0960-9822. PMID   29462577.
  6. Zhu, M.–Y.; Vannier, J.; Iten, H. V.; Zhao, Y.–L. (2004-08-07). "Direct evidence for predation on trilobites in the Cambrian". Proceedings of the Royal Society of London. Series B: Biological Sciences. 271 (suppl_5): S277-80. doi:10.1098/rsbl.2004.0194. ISSN   0962-8452. PMC   1810081 . PMID   15503993.
  7. Fu, Dongjing; Ortega-Hernández, Javier; Daley, Allison C; Zhang, Xingliang; Shu, Degan (December 2018). "Anamorphic development and extended parental care in a 520 million-year-old stem-group euarthropod from China". BMC Evolutionary Biology. 18 (1) 147. Bibcode:2018BMCEE..18..147F. doi: 10.1186/s12862-018-1262-6 . ISSN   1471-2148. PMC   6162911 . PMID   30268090.
  8. Liu, Yu; Ortega-Hernández, Javier; Chen, Hong; Mai, Huijuan; Zhai, Dayou; Hou, Xianguang (1 June 2020). "Computed tomography sheds new light on the affinities of the enigmatic euarthropod Jianshania furcatus from the early Cambrian Chengjiang biota". BMC Evolutionary Biology. 20 (1): 62. Bibcode:2020BMCEE..20...62L. doi: 10.1186/s12862-020-01625-4 . ISSN   1471-2148. PMC   7268425 . PMID   32487135.
  9. Chen, Ailin; Chen, Hong; Legg, David A.; Liu, Yu; Hou, Xian-guang (September 2018). "A redescription of Liangwangshania biloba Chen, 2005, from the Chengjiang biota (Cambrian, China), with a discussion of possible sexual dimorphism in fuxianhuiid arthropods". Arthropod Structure & Development. 47 (5): 552–561. Bibcode:2018ArtSD..47..552C. doi:10.1016/j.asd.2018.08.001. PMID   30125735. S2CID   52053402.
  10. Aria, C.; Zhao, F.; Zhu, M. (2021). "Fuxianhuiids are mandibulates and share affinities with total-group Myriapoda". Journal of the Geological Society. 178 (5) jgs2020-246. Bibcode:2021JGSoc.178..246A. doi:10.1144/jgs2020-246. S2CID   233952670.
  11. 1 2 O’Flynn, Robert J.; Liu, Yu; Hou, Xianguang; Mai, Huijuan; Yu, Mengxiao; Zhuang, Songling; Williams, Mark; Guo, Jin; Edgecombe, Gregory D. (August 2023). "The early Cambrian Kylinxia zhangi and evolution of the arthropod head". Current Biology. 33 (18): 4006–4013.e2. Bibcode:2023CBio...33E4006O. doi:10.1016/j.cub.2023.08.022. hdl: 10141/623137 . ISSN   0960-9822. PMID   37643622.
  12. 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) 20240622. doi:10.1098/rspb.2024.0622. ISSN   1471-2954. PMC   11463219 . PMID   39043240.
  13. Pulsipher, Mikaela A.; Anderson, Evan P.; Wright, Lauren S.; Kluessendorf, Joanne; Mikulic, Donald G.; Schiffbauer, James D. (2022-12-31). "Description of Acheronauta gen. nov., a possible mandibulate from the Silurian Waukesha Lagerstätte, Wisconsin, USA". Journal of Systematic Palaeontology. 20 (1): 1–24. doi:10.1080/14772019.2022.2109216. ISSN   1477-2019.
  14. Strausfeld, Nicholas J.; Andrew, David R.; Hou, Xianguang; Hirth, Frank (2025-08-28). "Brain anatomy of the Cambrian fossil Jianfengia multisegmentalis informs euarthropod phylogeny". Nature Communications. 16 (1) 7938. Bibcode:2025NatCo..16.7938S. doi:10.1038/s41467-025-62849-w. ISSN   2041-1723. PMC   12394709 . PMID   40877252.
  15. Liu, Yao; Zeng, Han; Zhao, Fangchen; Zhu, Yuyan; Li, Yimeng; Yin, Zongjun; Zhu, Maoyan (2025-05-31). "A tiny Cambrian stem-mandibulate reveals independent evolution of limb tagmatization and specialization in early euarthropods". Scientific Reports. 15 (1) 19115. Bibcode:2025NatSR..1519115L. doi:10.1038/s41598-025-03544-0. ISSN   2045-2322. PMC   12126567 . PMID   40450053.