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Kinorhyncha Temporal range: Early Cambrian–Recent PreꞒ Ꞓ O S D C P T J K Pg N
Echinoderes hwiiza
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Clade:	Bilateria
Clade:	Nephrozoa
(unranked):	Protostomia
Superphylum:	Ecdysozoa
Clade:	Scalidophora
Phylum:	Kinorhyncha Reinhard, 1881
Orders
See text

Kinorhyncha /kaɪnoʊˈrɪŋkə, kɪnə-/ (Ancient Greek: κινέω, romanized: kīnéō, lit. 'I move', ῥύγχος rhúnkhos "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are commonly called mud dragons. Modern species are 1 mm (0.039 in) or less, but Cambrian forms could reach 4 cm (1.6 in).

Anatomy

Kinorhynchs are limbless animals, with a body consisting of a head, neck, and a trunk of eleven segments. They are the only members of Ecdysozoa, except from the panarthropoda, with a segmented body. Juveniles have eight or nine segments, depending on genus, with the last two or three being added later during growth. Like other ecdysozoans they do not have external cilia, but instead have a number of spines along the body, plus up to seven circles of spines around the head. These spines are used for locomotion, withdrawing the head and pushing forward, then gripping the substrate with the spines while drawing up the body.

The body wall consists of a thin syncitial layer, which secretes a tough cuticle; this is molted several times while growing to adulthood. The spines are essentially moveable extensions of the body wall, and are hollow and covered by cuticle. The head is completely retractable, and is covered by a set of neck plates called placids when retracted.

Kinorhynchs eat either diatoms or organic material found in the mud, depending on species. The mouth is located in a conical structure at the apex of the head, and opens into a pharynx and then an oesophagus, both of which are lined by cuticle. Two pairs of salivary glands and one or more pairs of "pancreatic glands" connect to the oesophagus and presumably secrete digestive enzymes. Beyond the oesophagus lies a midgut that combines the functions of a stomach and intestine, and lacks a cuticle, enabling it to absorb nutrients. The short hind-gut is lined by cuticle, and empties into an anus at the posterior end of the trunk.

There is no circulatory system, although the body cavity (pseudocoelom) is well developed, and includes amoebocytes. The excretory system consists of two protonephridia emptying through pores in the final segment.

The nervous system consists of a ventral nerve cord, with one ganglion in each segment, and an anterior nerve ring surrounding the pharynx. Smaller ganglia are also located in the lateral and dorsal portions of each segment, but do not form distinct cords. Some species have simple ocelli on the head, and all species have tiny bristles on the body to provide a sense of touch.

Reproduction

There are two sexes that look alike, although some sexual dimorphism in allometry has been reported. A pair of gonads are located in the mid-region of the trunk, and open to pores in the final segment. In most species, the sperm duct includes two or three spiny structures that presumably aid in copulation, although the details are unknown. Individual spermatozoa can reach a quarter of the total body length. The larvae are free-living, but little else is known of their reproductive process. After having laid an egg, the female packs it into a protective envelope of mud and organic material.

Classification

Their closest relatives are thought to be the phyla Loricifera and Priapulida. Together they constitute the Scalidophora.

Taxonomy

The two groups of Kinorhynchs are generally characterized as classes in Sørensen et al. (2015). 270 species have been described and this number is expected to increase substantially. Morphological data has been collected for systematic phylogeny from dozens, and the integration of this with molecular data has led to a new systematic paradigm featuring the order Allomalorhagida (with Homalorhagida being retired). Phylogenomic data has shown Allomalorhagida and Cyclorhagida to be divided in three and two major clades respectively.

The oldest known species is Eokinorhynchus from the Fortunian of China.

Phylum Kinorhyncha

• Eokinorhynchus Zhang et al., 2015
• Class Cyclorhagida (Zelinka, 1896) Chitwood, 1951
  • Order Echinorhagata Sørensen et al., 2015
    • Echinoderidae Zelinka, 1894
  • Order Kentrorhagata Sørensen et al., 2015
    • Antygomonidae Adrianov & Malakhov, 1994
    • Cateriidae? Gerlach, 1956 (following Sørensen et al.)
    • Centroderidae Zelinka, 1896
    • Semnoderidae Remane, 1929
    • Zelinkaderidae Higgins, 1990
  • Order Xenosomata Zelinka, 1907
    • Campyloderidae Remane, 1929
• Class Allomalorhagida Sørensen et al., 2015
  • Pycnophyidae Zelinka, 1986
  • Order Anomoirhaga Herranz et al., 2022
    • Cateriidae? Gerlach, 1956 (following Herranz et al.)
    • Dracoderidae Higgins & Shirayama, 1990
    • Franciscideridae Sørensen et al., 2015
    • Neocentrophyidae Higgins, 1969

References

1. Zhang, Huaqiao; Xiao, Shuhai; Liu, Yunhuan; Yuan, Xunlai; Wan, Bin; Muscente, A. D.; Shao, Tiequan; Gong, Hao; Cao, Guohua; et al. (26 November 2015). "Armored kinorhynch-like scalidophoran animals from the early Cambrian". Nature. 5: 16521. Bibcode:2015NatSR...516521Z. doi:10.1038/srep16521. hdl:10919/74307. PMC 4660871. PMID 26610151.
2. "'Mindblowing' haul of fossils over 500m years old unearthed in China". The Guardian. 21 March 2019.
3. Herranz, M.; Park, T.; Di Domenico, M.; Leander, B. S.; Sørensen, M. V.; Worsaae, K. (2021). "Revisiting kinorhynch segmentation: Variation of segmental patterns in the nervous system of three aberrant species". Frontiers in Zoology. 18 (1): 54. doi:10.1186/s12983-021-00438-5. PMC 8529749. PMID 34674731.
4. Brusca, Richard C; Brusca, Gary J (2003). Invertebrates. Sinauer Associates. ISBN 978-0-87893-097-5. page 347
5. ^ Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 286–288. ISBN 978-0-03-056747-6.
6. Cepeda, Diego; Álamo, David; Sánchez, Nuria; Pardos, Fernando (December 2019). "Allometric growth in meiofaunal invertebrates: do all kinorhynchs show homogeneous trends?". Zoological Journal of the Linnean Society. 187 (4): 1041–1060. doi:10.1093/zoolinnean/zlz083.
7. Neuhaus, B.; Higgins, R. P. (July 2002). "Ultrastructure, Biology, and Phylogenetic Relationships of Kinorhyncha". Integrative and Comparative Biology. 42 (3): 619–32. doi:10.1093/icb/42.3.619. PMID 21708758.
8. Meet the mud dragon—the tiny animal that lives on the beach - Phys.org
9. ^ Sørensen, M. V. et al. Phylogeny of Kinorhyncha based on morphology and two molecular loci. PLoS One 10, 1–33 (2015).
10. Altenburger, Andreas; Rho, Hyun S.; Chang, Cheon Y.; Sørensen, Martin V. (December 2015). "Zelinkaderes yong sp. nov. from Korea – the first recording of Zelinkaderes (Kinorhyncha: Cyclorhagida) in Asia". Zoological Studies. 54 (25): e25. doi:10.1186/s40555-014-0103-6. PMC 6661445. PMID 31966112.
11. "Species list of Kinorhyncha - Hiroshi Yamasaki website". sites.google.com. Retrieved 2018-09-20.
12. Herranz, Maria; et al. (March 2022). "Phylogenomic analyses of mud dragons (Kinorhyncha)". Molecular Phylogenetics and Evolution. 168: 107375. Bibcode:2022MolPE.16807375H. doi:10.1016/j.ympev.2021.107375. hdl:10115/29090. PMID 34952205. S2CID 245419318.
13. Zhang, Huaqiao; Xiao, Shuhai; Liu, Yunhuan; Yuan, Xunlai; Wan, Bin; Muscente, A. D.; Shao, Tiequan; Gong, Hao; Cao, Guohua (December 2015). "Armored kinorhynch-like scalidophoran animals from the early Cambrian". Scientific Reports. 5 (1): 16521. Bibcode:2015NatSR...516521Z. doi:10.1038/srep16521. ISSN 2045-2322. PMC 4660871. PMID 26610151.

External links

• Media related to Kinorhyncha at Wikimedia Commons
• Introduction to the Cephalorhyncha
• Drawings of Kinorhyncha
• "Kinorhyncha". The Encyclopedia of Life.

v t e Kinorhynch families and genera
Kingdom Animalia (unranked) Eumetazoa (unranked) Bilateria (unranked) Protostomia Superphylum Ecdysozoa (unranked) Scalidophora
Stem-group	†Eokinorhynchus
Cyclorhagida	Echinorhagata Echinoderidae Kentrorhagata Antygomonidae Cateriidae? Centroderidae Semnoderidae Zelinkaderidae Tubulideres Wollunquaderes Xenosomata Campyloderidae
Allomalorhagida	Anomoirhaga Cateriidae? Dracoderidae Franciscideridae Neocentrophyidae Pycnophyidae

• Kinorhyncha
• Ecdysozoa phyla