Chaetognatha
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Chaetognatha is 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 or 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. Some deep-sea species are orange. They range in size from 2 mm to 12 cm. The common term for the phylum is Arrow Worms. There are more than 120 modern species assigned to over 20 genera. Despite the limited diversity of species, the number of individuals is staggering.[1]
Chaetognaths are transparent or translucent and are covered by a cuticle. They have fins and a pair of hooked, chitinous, grasping spines on each side of their heads that are used in hunting. The spines are covered with a hood when swimming. They have a distinct head, trunk and tail. All species are hermaphroditic, carrying both eggs and sperm. Some species are known to use the neurotoxin tetrodotoxin to subdue prey.[2]
They have some developmental similarities to nematodes. Although they have a mouth with one or two rows of tiny teeth, compound eyes, and a nervous system, they have no respiratory, circulatory, or excremental systems. Materials are moved about the body cavity by cilia. Waste materials are simply excreted through the skin and anus. Chaetognaths swim in short bursts using a dorso-ventral undulating body motion, where their tail fin assists with propulsion and the body fins for stabilization and steering.[3] At least one species of chaetognath, Caecosagitta macrocephala, has bioluminescent organs on its lateral fins.[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.[5] Molecular phylogenists, however, consider them to be protostomes. Thomas Cavalier-Smith places them in the protostomes in his Six Kingdom classification.[6] The similarities between chaetognaths and nematodes mentioned above may support the protostome thesis - in fact, chaetognaths are sometimes regarded as a basal ecdysozoan.[7] Chaetognatha appears close to the base of the protostome tree in most studies of their molecular phylogeny.[8] 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.[9]
Fossil record
Due to their soft bodies, chaetognaths fossilize poorly. Even so, several fossil chaetognath species have been described.[10] 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[11] and Protosagitta spinosa Hu[12]) and the Middle Cambrian Burgess Shale of British Columbia (Oesia disjuncta Walcott[13]). 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. [14]
ReferencesISBN links support NWE through referral fees
- ↑ Bone, Q. et al. (1991). The biology of chaetognaths. Oxford University Press.
- ↑ Thuesen, E.V. (1991), "The tetrodotoxin venom of chaetognaths", in Bone, Q. et al., The biology of chaetognaths, London: Oxford University Press, at 55-60
- ↑ 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: 119-137.
- ↑ Haddock, S.H.D. & J.F. Case (2004). A bioluminescent chaetognath. Nature 367: 225-226.
- ↑ 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: R575-R576.
- ↑ Marletaz, F. et al. (2006). Chaetognath phylogenomics: a protostome with deuterostome-like development. Current Biology 16: R577-R578.
- ↑ Papillon, Daniel et al. (2004). Identification of chaetognaths as protostomes is supported by the analysis of their mitochondrial genome. Molecular Biology & Evolution 21 (11): 2122-2129.
- ↑ Vannier, J., M. Steiner, E. Renvoise, S.-X. Hu, and J.-P. Casanova. 2007. Early Cambrian origin of modern food webs: evidence from predator arrow worms. Proceedings of the Royal Society B 274:627–633.
- ↑ Chen, J.-Y., and D.-Y. Huang. 2002. A possible Lower Cambrian chaetognath (arrow worm). Science 298:187.
- ↑ 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. Acta Palaeontologica Polonica 50:1-8.
- ↑ Szaniawski, H. 2002. New evidence for the protoconodont origin of chaetognaths. Acta Palaeontologica Polonica 47:405-419.
- Pierrot-Bults, A. C. 2004. Chaetognatha. In B. Grzimek, S. F. Craig, D. A. Thoney, N. Schlager, and M. Hutchins. Grzimek's Animal Life Encyclopedia, 2nd edition, vol. 2, Protosomes. Detroit, MI: Thomson/Gale. ISBN 0787657786.
External links
- Chaetognatha of the World - last retrieved December 13 2006
- List of valid chaetognath species
- Hints for identifying chaetognaths
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