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arrowworms: a group of small active transparent marine worms (同)phylum Chaetognatha

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2015/10/03 17:23:54」(JST)

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Chaetognatha, meaning bristle-jaws, and commonly known as arrow worms, are a phylum of predatory marine worms that are a major component of plankton worldwide. About 20% of the known species are benthic, and can attach to algae and rocks. They are found in all marine waters, from surface tropical waters and shallow tide pools to the deep sea and polar regions. Most chaetognaths are transparent and are torpedo shaped, but some deep-sea species are orange. They range in size from 2 to 120 millimetres (0.079 to 4.724 in).

There are more than 120 modern species assigned to over 20 genera.^[2] Despite the limited diversity of species, the number of individuals is large.^[3]

Anatomy

The jaw organ of a Chaetognath of the genus Sagitta

Chaetognaths are transparent or translucent dart-shaped animals covered by a cuticle. The body is divided into a distinct head, trunk, and tail. There are between four and fourteen hooked, grasping spines on each side of their head, flanking a hollow vestibule containing the mouth. The spines are used in hunting, and covered with a flexible hood arising from the neck region when the animal is swimming. All chaetognaths are carnivorous, preying on other planktonic animals.^[4]

The trunk bears one or two pairs of lateral fins incorporating structures superficially similar to the fin rays of fish, with which they are not homologous, however: unlike those of vertebrates, these are composed of a thickened basement membrane extending from the epidermis. An additional caudal fin covers the post-anal tail.^[4] Two chaetognath species, Caecosagitta macrocephala and Eukrohnia fowleri, have bioluminescent organs on their fins.^[5]^[6]

Chaetognaths swim in short bursts using a dorso-ventral undulating body motion, where their tail fin assists with propulsion and the body fins with stabilization and steering.^[7] Some species are known to use the neurotoxin tetrodotoxin to subdue prey.^[8]

The body cavity lacks a peritoneum, and therefore resembles the pseudocoel of animals such as nematodes, but is divided into one compartment on each side of the trunk, and additional compartments inside the head and tail.^[4] Although they have a mouth with one or two rows of tiny teeth, compound eyes, and a nervous system, they have no respiratory or circulatory systems.

The mouth opens into a muscular pharynx, which contains glands to lubricate the passage of food. From here, a straight intestine runs the length of the trunk to an anus just in front of the tail. The intestine is the primary site of digestion and includes a pair of diverticula near the anterior end.^[4]

The nervous system is reasonably simple, consisting of a ganglionated nerve ring surrounding the pharynx. The dorsal ganglion is the largest, but nerves extend from all the ganglia along the length of the body. Chaetognaths have two compound eyes, each consisting of a number of pigment-cup ocelli fused together. In addition, there are a number of sensory bristles arranged in rows along the side of the body, where they probably perform a function similar to that of the lateral line in fish. An additional, curved, band of sensory bristles lies over the head and neck.^[4]

Materials are moved about the body cavity by cilia. Waste materials are simply excreted through the skin and anus.

The arrow worm rhabdomeres are derived from microtubules 20 nm long and 50 nm wide, which in turn form conical bodies that contain granules and thread structures. The cone body is derived from a cilium.^[9]

Reproduction

All species are hermaphroditic, carrying both eggs and sperm.^[3] Each animal possesses a pair of testes within the tail, and a pair of ovaries in the posterior region of the main body cavity. Immature sperm are released from the testes to mature inside the cavity of the tail, and then swim through a short duct to a seminal vesicle where they are packaged into a spermatophore.^[4]

During mating, each individual places a spermatophore onto the neck of its partner after rupture of the seminal vesicle. The sperm rapidly escape from the spermatophore and swim along the midline of the animal until they reach a pair of small pores just in front of the tail. These pores connect to the oviducts, into which the developed eggs have already passed from the ovaries, and it is here that fertilisation takes place.^[4]

The eggs are planktonic, or attached to algae, and hatch into miniature versions of the adult, without a well-defined larval stage.^[4]

Classification

Chaetognaths are traditionally classed as deuterostomes by embryologists. Lynn Margulis and K. V. Schwartz place chaetognaths in the deuterostomes in their Five Kingdom classification.^[10] Molecular phylogenists, however, consider them to be protostomes. Thomas Cavalier-Smith places them in the protostomes in his Six Kingdom classification.^[11] The similarities between chaetognaths and nematodes mentioned above may support the protostome thesis—in fact, chaetognaths are sometimes regarded as a basal ecdysozoan or lophotrochozoan.^[12] Chaetognatha appears close to the base of the protostome tree in most studies of their molecular phylogeny.^[13] This may explain their deuterostome embryonic characters. If chaetognaths branched off from the protostomes before they evolved their distinctive protostome embryonic characters, they may have retained deuterostome characters inherited from early bilaterian ancestors. Thus chaetognaths may be a useful model for the ancestral bilaterian.^[14] Studies of arrow worms' nervous systems suggests they should be placed within the protostomes.^[15]^[16]

Fossil record

Due to their soft bodies, chaetognaths fossilize poorly. Even so, several fossil chaetognath species have been described.^[17] Chaetognaths appear to have originated in the Cambrian Period. Complete body fossils have been formally described from the Lower Cambrian Maotianshan shales of Yunnan, China (Eognathacantha ercainella Chen & Huang^[18] and Protosagitta spinosa Hu^[19]) and the Middle Cambrian Burgess Shale of British Columbia (Oesia disjuncta Walcott^[20]), a view challenged by Conway Morris (2009). A more recent chaetognath, Paucijaculum samamithion Schram, has been described from the Mazon Creek biota from the Pennsylvanian of Illinois. Chaetognaths were thought possibly to be related to some of the animals grouped with the conodonts. The conodonts themselves, however, are thought to be related to the vertebrates. It is now thought that protoconodont elements (e.g., Protohertzina anabarica Missarzhevsky, 1973), are probably grasping spines of chaetognaths rather than teeth of conodonts. Previously chaetognaths in the Early Cambrian were only suspected from these protoconodont elements, but the more recent discoveries of body fossils have confirmed their presence then.^[21]

References

^ "Sagittoidea Claus and Grobben, 1905". Integrated Taxonomic Information System. Retrieved February 8, 2012.
^ "World Register of Marine Species".
^ ^a ^b Bone, Q.; Kapp, H.; Pierrot-Bults, A. C., eds. (1991). The Biology of Chaetognaths. London: Oxford University Press. ISBN 978-0-19-857715-7.
^ ^a ^b ^c ^d ^e ^f ^g ^h Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 1046–1050. ISBN 0-03-056747-5.
^ Haddock, S. H. D. & J. F. Case (2004). "A bioluminescent chaetognath". Nature 367 (6460): 225–226. doi:10.1038/367225a0.
^ Thuesen, E. V., F. E. Goetz & S. H. D. Haddock (2010). "Bioluminescent organs of two deep-sea arrow worms, Eukrohnia fowleri and Caecosagitta macrocephala, with further observations on bioluminescence in chaetognaths". The Biological Bulletin 219 (2): 100–111. PMID 20972255.
^ Jordan, C. E. (1992). "A model of rapid-start swimming at intermediate Reynolds number: undulatory locomotion in the chaetognath Sagitta elegans". Journal of Experimental Biology 163 (1): 119–137.
^ Thuesen, E. V. 1991, "The Tetrodotoxin Venom of Chaetognaths," pp. 55–60
^ "Photoreception". Encyclopædia Britannica from Encyclopædia Britannica 2006 Ultimate Reference Suite DVD . 2009.
^ Systema Naturae 2000 Taxon: Phylum Chaetognatha per Margulis and Schwartz (select Margulis & Schwartz in 'Classification by')—last retrieved November 25, 2006
^ Systema Naturae 2000 Taxon: Phylum Chaetognatha per Cavalier-Smith (select Cavalier-Smith in 'Classification by')—last retrieved November 25, 2006
^ Matus, D. Q.; et al. (2006). "Broad taxon and gene sampling indicate that chaetognaths are protostomes". Current Biology 16 (15): R575–R576. doi:10.1016/j.cub.2006.07.017. PMID 16890509.
^ Marletaz, F.; et al. (2006). "Chaetognath phylogenomics: a protostome with deuterostome-like development". Current Biology 16 (15): R577–R578. doi:10.1016/j.cub.2006.07.016. PMID 16890510.
^ Papillon, Daniel; et al. (2004). "Identification of chaetognaths as protostomes is supported by the analysis of their mitochondrial genome". Molecular Biology and Evolution 21 (11): 2122–2129. doi:10.1093/molbev/msh229. PMID 15306659.
^ Rieger, V.; Perez, Y.; Müller, C. H. G.; Lipke, E.; Sombke, A.; Hansson, B. S.; Harzsch, S. (2010). "Immunohistochemical analysis and 3D reconstruction of the cephalic nervous system in Chaetognatha: Insights into the evolution of an early bilaterian brain?". Invertebrate Biology 129 (1): 77–104. doi:10.1111/j.1744-7410.2010.00189.x.
^ Harzsch, S.; Müller, C. H. (2007). "A new look at the ventral nerve centre of Sagitta: implications for the phylogenetic position of Chaetognatha (arrow worms) and the evolution of the bilaterian nervous system". Frontiers in Zoology 4: 14. doi:10.1186/1742-9994-4-14. PMC 1885248. PMID 17511857.
^ Vannier, J.; Steiner, M.; Renvoise, E.; Hu, S.-X.; Casanova, J.-P. (2007). "Early Cambrian origin of modern food webs: evidence from predator arrow worms". Proceedings of the Royal Society B: Biological Sciences 274 (1610): 627–633. doi:10.1098/rspb.2006.3761. PMC 2197202. PMID 17254986.
^ Chen, J.-Y.; Huang, D.-Y. (2002). "A possible Lower Cambrian chaetognath (arrow worm)". Science 298 (5591): 187. doi:10.1126/science.1075059. PMID 12364798.
^ Hu, S.-X. (2005). "Taphonomy and palaeoecology of the Early Cambrian Chengjiang Biota from Eastern Yunnan, China". Berliner Paläobiologische Abhandlungen 7: 1–197.
^ Szaniawski, H. (2005). "Cambrian chaetognaths recognized in Burgess Shale fossils" (PDF). Acta Palaeontologica Polonica 50 (1): 1–8.
^ Szaniawski, H. (2002). "New evidence for the protoconodont origin of chaetognaths" (PDF). Acta Palaeontologica Polonica 47 (3): 405–419.

External links

Image of Pseudosagitta gazellae with a krill in its gut from the Tasmanian Aquaculture and Fisheries Institute
Chaetognatha of the World – last retrieved December 13, 2006
List of valid chaetognath species
Hints for identifying chaetognaths

English Journal

Contribution to the understanding of the cycle of the protozoan parasite Marteilia refringens.

Arzul I1, Chollet B1, Boyer S2, Bonnet D2, Gaillard J1, Baldi Y3, Robert M1, Joly JP1, Garcia C1, Bouchoucha M4.
Parasitology.Parasitology.2014 Feb;141(2):227-40. doi: 10.1017/S0031182013001418. Epub 2013 Oct 11.
The paramyxean parasite Marteilia refringens infects several bivalve species including European flat oysters Ostrea edulis and Mediterranean mussels Mytilus galloprovincialis. Sequence polymorphism allowed definition of three parasite types 'M', 'O' and 'C' preferably detected in oysters, mussels an
PMID 24128728

Evolutionary history of Chaetognatha inferred from molecular and morphological data: a case study for body plan simplification.

Gasmi S1, Nève G2, Pech N2, Tekaya S3, Gilles A2, Perez Y2.
Frontiers in zoology.Front Zool.2014 Nov 21;11(1):84. doi: 10.1186/s12983-014-0084-7. eCollection 2014.
BACKGROUND: Chaetognatha are a phylum of marine carnivorous animals which includes more than 130 extant species. The internal systematics of this group have been intensively debated since it was discovered in the 18(th) century. While they can be traced back to the earlier Cambrian, they are an extr
PMID 25473413

Neurogenesis in an early protostome relative: progenitor cells in the ventral nerve center of chaetognath hatchlings are arranged in a highly organized geometrical pattern.

Perez Y1, Rieger V, Martin E, Müller CH, Harzsch S.
Journal of experimental zoology. Part B, Molecular and developmental evolution.J Exp Zool B Mol Dev Evol.2013 May;320(3):179-93. doi: 10.1002/jez.b.22493. Epub 2013 Mar 8.
Emerging evidence suggests that Chaetognatha represent an evolutionary lineage that is the sister group to all other Protostomia thus promoting these animals as a pivotal model for our understanding of bilaterian evolutionary history. We have analyzed the proliferation of neuronal progenitor cells i
PMID 23483730

Japanese Journal

Community structure and spatial distribution of chaetognaths in Tosa Bay on the temperate Kuroshio coast of Japan

Plankton and Benthos Research 9(3), 176-187, 2014
NAID 130004688122

Second record of Eukrohnia kitoui (Chaetognatha) with notes on morphological differences from the closely related E. calliops

Plankton & benthos research 8(4), 191-194, 2013-11-01
NAID 10031203405

北太平洋の縁辺海および日本近海における浮遊性毛顎動物相と分布生態に関するレビュー (ヤムシ研究会ワークショップ論文集)

日本プランクトン学会報 59(2), 88-93, 2012-08
NAID 40019436539

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リンク元	「毛顎動物」「毛顎動物門」「arrow worm」

「毛顎動物」

Arrow worms; Temporal range: Lower Cambrian–Recent PreЄ Є O S D C P T J K Pg N
	Spadella cephaloptera
Scientific classification
Kingdom:	Animalia
Phylum:	Chaetognatha; Leuckart, 1854
Class:	Sagittoidea; Claus & Grobben, 1905
Orders and families
	Aphragmophora Bathybelidae; Krohnittidae; Pterokrohniidae; Pterosagittidae; Sagittidae; ; Phragmophora Eukrohniidae; Heterokrohniidae; Krohnittellidae; Spadellidae; ;

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ラ: Chaetognatha、arrow worm
関: 毛顎動物門

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ラ: Chaetognatha
関: 毛顎動物

「arrow worm」

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毛顎動物

関: Chaetognatha

[1] "Sagittoidea Claus and Grobben, 1905". Integrated Taxonomic Information System. Retrieved February 8, 2012.

[2] "World Register of Marine Species".

[Bone-3] Bone, Q.; Kapp, H.; Pierrot-Bults, A. C., eds. (1991). The Biology of Chaetognaths. London: Oxford University Press. ISBN 978-0-19-857715-7.

[IZ-4] ^ ^a ^b ^c ^d ^e ^f ^g ^h Barnes, Robert D. (1982). Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 1046–1050. ISBN 0-03-056747-5.

[5] Haddock, S. H. D. & J. F. Case (2004). "A bioluminescent chaetognath". Nature 367 (6460): 225–226. doi:10.1038/367225a0.

[6] Thuesen, E. V., F. E. Goetz & S. H. D. Haddock (2010). "Bioluminescent organs of two deep-sea arrow worms, Eukrohnia fowleri and Caecosagitta macrocephala, with further observations on bioluminescence in chaetognaths". The Biological Bulletin 219 (2): 100–111. PMID 20972255.

[7] Jordan, C. E. (1992). "A model of rapid-start swimming at intermediate Reynolds number: undulatory locomotion in the chaetognath Sagitta elegans". Journal of Experimental Biology 163 (1): 119–137.

[8] Thuesen, E. V. 1991, "The Tetrodotoxin Venom of Chaetognaths," pp. 55–60

[9] "Photoreception". Encyclopædia Britannica from Encyclopædia Britannica 2006 Ultimate Reference Suite DVD . 2009.

[10] Systema Naturae 2000 Taxon: Phylum Chaetognatha per Margulis and Schwartz (select Margulis & Schwartz in 'Classification by')—last retrieved November 25, 2006

[11] Systema Naturae 2000 Taxon: Phylum Chaetognatha per Cavalier-Smith (select Cavalier-Smith in 'Classification by')—last retrieved November 25, 2006

[12] Matus, D. Q.; et al. (2006). "Broad taxon and gene sampling indicate that chaetognaths are protostomes". Current Biology 16 (15): R575–R576. doi:10.1016/j.cub.2006.07.017. PMID 16890509.

[13] Marletaz, F.; et al. (2006). "Chaetognath phylogenomics: a protostome with deuterostome-like development". Current Biology 16 (15): R577–R578. doi:10.1016/j.cub.2006.07.016. PMID 16890510.

[14] Papillon, Daniel; et al. (2004). "Identification of chaetognaths as protostomes is supported by the analysis of their mitochondrial genome". Molecular Biology and Evolution 21 (11): 2122–2129. doi:10.1093/molbev/msh229. PMID 15306659.

[15] Rieger, V.; Perez, Y.; Müller, C. H. G.; Lipke, E.; Sombke, A.; Hansson, B. S.; Harzsch, S. (2010). "Immunohistochemical analysis and 3D reconstruction of the cephalic nervous system in Chaetognatha: Insights into the evolution of an early bilaterian brain?". Invertebrate Biology 129 (1): 77–104. doi:10.1111/j.1744-7410.2010.00189.x.

[16] Harzsch, S.; Müller, C. H. (2007). "A new look at the ventral nerve centre of Sagitta: implications for the phylogenetic position of Chaetognatha (arrow worms) and the evolution of the bilaterian nervous system". Frontiers in Zoology 4: 14. doi:10.1186/1742-9994-4-14. PMC 1885248. PMID 17511857.

[17] Vannier, J.; Steiner, M.; Renvoise, E.; Hu, S.-X.; Casanova, J.-P. (2007). "Early Cambrian origin of modern food webs: evidence from predator arrow worms". Proceedings of the Royal Society B: Biological Sciences 274 (1610): 627–633. doi:10.1098/rspb.2006.3761. PMC 2197202. PMID 17254986.

[18] Chen, J.-Y.; Huang, D.-Y. (2002). "A possible Lower Cambrian chaetognath (arrow worm)". Science 298 (5591): 187. doi:10.1126/science.1075059. PMID 12364798.

[19] Hu, S.-X. (2005). "Taphonomy and palaeoecology of the Early Cambrian Chengjiang Biota from Eastern Yunnan, China". Berliner Paläobiologische Abhandlungen 7: 1–197.

[20] Szaniawski, H. (2005). "Cambrian chaetognaths recognized in Burgess Shale fossils" (PDF). Acta Palaeontologica Polonica 50 (1): 1–8.

[21] Szaniawski, H. (2002). "New evidence for the protoconodont origin of chaetognaths" (PDF). Acta Palaeontologica Polonica 47 (3): 405–419.

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Chaetognatha