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Polychaete

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Class of annelid worms
"Polychaeta" redirects here. For the genus of flies, seePolychaeta (fly).
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Polychaetes
Temporal range:Cambrian(or earlier?) – present
Plate titled "A variety of marine worms" from M. J. Schleiden's Das Meer
Plate titled "A variety of marine worms" fromM. J. Schleiden'sDas Meer
Scientific classificationEdit this classification
Kingdom:Animalia
Phylum:Annelida
Class:Polychaeta
Grube, 1850
Groups included
Cladistically included but traditionally excluded taxa

Polychaeta (/ˌpɒlɪˈktə/) is aparaphyletic class of generally marineannelid worms,[1]commonly calledbristle worms orpolychaetes (/ˈpɒlɪˌkts/). Each body segment has a pair of fleshy protrusions calledparapodia which bear manychitinous bristles calledchaetae, hence their name.

More than 10,000 species have beendescribed in this diverse and widespread class; in addition toinhabiting all of the world's oceans, polychaetes occur at allocean depths, fromplanktonic species living near thesurface, to a smallundescribed species observed throughROV at the deepest region in the Earth's oceans,Challenger Deep. In addition, many species live on theabyssal plains,coral reefs,parasitically, and a few withinfresh water.

Commonly encountered representatives include thelugworms,bloodworms, and species ofAlitta such as theclam worm andsandworm or ragworm; these species inhabitshallow water marine environments andcoastlines ofsubtropical andtemperate regions around the world and may be used asfishing bait. More exotic species include thestinging fireworms, thepredatory and large-bodiedbobbit worm, the culturally importantpalolo worm, thebone-eating worms, andgiant tube worms, which areextremophiles that tolerate near-boiling water nearhydrothermal vents.

Description

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Main articles:Annelid § Description, andParapodium

Polychaetes are segmented worms, generally less than 10 cm (4 in) in length, although ranging at the extremes from 1 mm (0.04 in) to 3 m (10 ft), inEunice aphroditois. They can sometimes be brightly coloured, and may beiridescent or evenluminescent. Each segment bears a pair of paddle-like and highly vascularizedparapodia, which are used for movement and, in many species, act as the worm's primaryrespiratory surfaces. Bundles of bristles, thechaetae, project from the parapodia.[2]

However, polychaetebody plans vary widely from this generalized pattern, and can display a range of different body forms. The most generalised polychaetes are those thatcrawl along the bottom, but others have adapted to many differentecological niches, includingburrowing,pelagic swimming,dwelling in self-created tubes or ones bored out of a substrate,commensalism, andparasitism; such varied lifestyles requires adivergence from thebasic body plan of thecommon ancestor.

Pharynx eversion inPhyllodoce lineata
The plumes of afeather duster worm are known asradioles

The head, orprostomium, is relatively well developed, compared with other annelids. It projects forward over the mouth, which is located on the succeeding section; theperistomium. The mouthparts vary in form depending on their diets, since the group includes predators, herbivores, filter feeders, scavengers, and parasites. In general, however, they possess a pair of jaws and apharynx that can be rapidly everted, allowing the worms to grab food and pull it into their mouths. In some species, the pharynx is modified into a lengthyproboscis.[citation needed] Their jaws are formed fromsclerotised collagen.[3] The digestive tract is a simple tube, usually with a stomach partway along.

The head may include two to four pairs of eyes, although some species are eyeless. The eyes are typically fairly simple structures, capable of distinguishing only light and dark, although some species have large eyes withlenses that may be capable of more sophisticated vision,[2] an example being the complex eyes ofAlciopidae, which rival those ofcephalopods andvertebrates.[4][5] The head also includes a pair ofantennae, tentacle-likepalps, and a pair of pits lined withcilia known asnuchal organs, which arechemoreceptors that help the worm to seek out food.[2]

Polychaetecross section

The outer surface of the body wall consists of a simplecolumnar epithelium covered by a thincuticle, constructed fromcross-linkedcollagen fibers and may be 2 to 13 millimetres (0.079 to 0.512 in) thick. Sclerotized collagen makes up their setae.[3]

Underneath the cuticle, in order, are a thin layer ofconnective tissue, a layer ofcircular muscle, a layer of longitudinal muscle, and aperitoneum surrounding thecoelom (body cavity). Additional oblique muscles move the parapodia. In most species the body cavity is divided into separate compartments by sheets of peritoneum between each segment, but in some species it is more continuous.

Physiology

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A simple but well-developedcirculatory system is usually present. The two main blood vessels furnish smaller vessels to supply the parapodia and the gut. Blood flows forward in the dorsal vessel, above the gut, and returns down the body in the ventral vessel, beneath the gut. The blood vessels themselves are contractile, helping to push the blood along, so most species have no need of a heart. In a few cases, however, muscular pumps analogous to a heart are found in various parts of the system. Conversely, some species have little or no circulatory system at all, transporting oxygen in thecoelomic fluid that fills their body cavities.[2] The blood may be colourless, or have any of three different respiratory pigments. The most common of these ishaemoglobin, but some groups havehaemerythrin or the green-colouredchlorocruorin, instead.

The smallest species, and those adapted to burrowing, lackgills, breathing only through their body surfaces (bydiffusion). Most other species haveexternal gills, often associated with the parapodia.

The nervous system consists of a single or double ventral nerve cord running the length of the body, withganglia and a series of small nerves in each segment. The brain is relatively large, compared with that of other annelids, and lies in the upper part of the head. Anendocrine gland is attached to the ventral posterior surface of the brain, and appears to be involved in reproductive activity. In addition to the sensory organs on the head, photosensitive eye spots,statocysts, and numerous additional sensory nerve endings, most likely involved with the sense of touch, also occur on the body.[2]

Polychaetes have a varying number ofprotonephridia ormetanephridia for excreting waste, which in some cases can be relatively complex in structure. The body also contains greenish "chloragogen" tissue, similar to that found inoligochaetes, which appears to function in metabolism, in a similar fashion to that of the vertebrateliver.[2]

Many species exhibitbioluminescence; eight families have luminous species.[6][7]

Ecology

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Polychaetes are predominantly marine, but 168 species (nearing 2% of total species) also live in freshwater,[8] and a few insemiterrestrial environments and evenin caves.[9][10] They are extremely variable in both form and lifestyle, and include a few taxa that swim among theplankton or above theabyssal plain. Most burrow or build tubes in the sediment, and some live ascommensals. A few species, roughly 80 (less than 0.5% of species), are parasitic.[11][12] These include bothectoparasites andendoparasites. Ectoparasitic polychaetes feed on skin, blood, and other secretions, and some are adapted to bore through hard, usually calcerous surfaces, such as the shells of mollusks.[12] These "boring" polychaetes may be parasitic, but may be opportunistic or even obligate symbionts (commensals).[13][12][11]

The mobile forms (Errantia) tend to have well-developed sense organs and jaws, while the stationary forms (Sedentaria) lack them, but may have specialized gills or tentacles used for respiration and deposit or filter feeding, e.g.,fanworms. Polychaete mouthparts are eversible and used to capture prey.[14][self-published source?] A few groups have evolved to live in terrestrial environments, likeNamanereidinae with many terrestrial species, but are restricted to humid areas. Some have even evolved cutaneous invaginations for aerial gas exchange.[9]

Reproduction

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Two examples of epitoky in progress;

Top:Palola viridis (Eunicida)

Bottom: Syllidaesp. (Phyllodocida)

Most polychaetes haveseparate sexes, rather than beinghermaphroditic. The most primitive species have a pair ofgonads in every segment, but most species exhibit some degree of specialisation. The gonads shed immaturegametes directly into the body cavity, where they complete their development. Once mature, the gametes are shed into the surrounding water through ducts or openings that vary between species, or in some cases by the complete rupture of the body wall (and subsequent death of the adult). A few speciescopulate, but most fertilize their eggs externally.

The fertilized eggs typically hatch intotrochophore larvae, which float among theplankton, and eventuallymetamorphose into the adult form by adding segments. A few species have no larval form, with the egg hatching into a form resembling the adult, and in many that do have larvae, the trochophore never feeds, surviving off the yolk that remains from the egg.[2]

However, some polychaetes exhibit remarkable reproductive strategies. Some species reproduce byepitoky. For much of the year, these worms look like any other burrow-dwelling polychaete, but as the breeding season approaches, the worm undergoes a remarkable transformation as new, specialized segments begin to grow from its rear end until the worm can be clearly divided into two halves. The front half, the atoke, is asexual. The new rear half, responsible for breeding, is known as the epitoke. Each of the epitoke segments is packed with eggs and sperm and features a single eyespot on its surface. The beginning of the last lunar quarter is the cue for these animals to breed, and the epitokes break free from the atokes and float to the surface. The eye spots sense when the epitoke reaches the surface and the segments from millions of worms burst, releasing their eggs and sperm into the water.[19]

A similar strategy is employed by the branching deep sea wormSyllis ramosa, which lives inside asponge; the worm develop "stolons" containing eggs or sperm from one of their many rear ends; these stolons detach from the parent worm and rise to the sea surface, where fertilisation takes place.[20]

Evolution

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Stem-group polychaete fossils are known from theSirius PassetLagerstätte, a rich, sedimentary deposit in Greenland tentatively dated to the lateAtdabanian (earlyCambrian). The oldest known polychaete as of 2025 isDannychaeta tucolus, dated to approximately 514 million years ago.[21][22] Many of the more famousBurgess Shale organisms, such asCanadia, may also have polychaete affinities.Wiwaxia, long interpreted as an annelid,[23] is now considered to represent a mollusc.[24][25] An even older fossil,Cloudina, dates to the terminalEdiacaran period; this has been interpreted as an early polychaete, although consensus is absent.[26][27]

Beingsoft-bodied organisms, the fossil record of polychaetes is dominated by their fossilized jaws, known asscolecodonts, and themineralized tubes that some of them secrete.[28] Most importantbiomineralising polychaetes areserpulids,sabellids, andcirratulids. Polychaete cuticle does have somepreservation potential; it tends to survive for at least 30 days after a polychaete's death.[3] Although biomineralisation is usually necessary to preserve soft tissue after this time, the presence of polychaete muscle in the nonmineralised Burgess shale shows this need not always be the case.[3] Their preservation potential is similar to that ofjellyfish.[3]

Taxonomy and systematics

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This section needs to beupdated. The reason given is: Taxonomy list. Please help update this article to reflect recent events or newly available information.(July 2025)
See also:List of annelid families

Taxonomically, polychaetes are thought to beparaphyletic,[29] meaning the group excludes some descendants of its most recent common ancestor. Groups that may be descended from the polychaetes include theclitellates (earthworms andleeches),sipunculans, andechiurans. The Pogonophora andVestimentifera were once considered separate phyla, but are now classified in the polychaete familySiboglinidae.

Much of the classification below matches Rouse & Fauchald, 1998, although that paper does not apply ranks above family.

Older classifications recognize many more (sub)orders than the layout presented here. As comparatively few polychaetetaxa have been subject tocladistic analysis, some groups which are usually considered invalid today may eventually be reinstated.

These divisions were shown to be mostly paraphyletic in recent years.

Below is a phylogenetic tree of annelids from a 2021 review of annelid diversity (clades labeled× are not considered polychaetes);[30]

Annelida

See also

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References

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Bibliography

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Notes

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  1. ^abStruck, T. H.; Paul, C.; Hill, N.; Hartmann, S.; Hösel, C.; Kube, M.; Lieb, B.; Meyer, A.; Tiedemann, R.; Purschke, G. N.; Bleidorn, C. (2011). "Phylogenomic analyses unravel annelid evolution".Nature.471 (7336):95–98.Bibcode:2011Natur.471...95S.doi:10.1038/nature09864.PMID 21368831.S2CID 4428998.
  2. ^abcdefgBarnes, Robert D. (1982).Invertebrate Zoology. Philadelphia, PA: Holt-Saunders International. pp. 469–525.ISBN 978-0-03-056747-6.
  3. ^abcdeBriggs, Derek E. G.; Kear, Amanda J. (8 February 2016). "Decay and preservation of polychaetes: taphonomic thresholds in soft-bodied organisms".Paleobiology.19 (1):107–135.Bibcode:1993Pbio...19..107B.doi:10.1017/S0094837300012343.JSTOR 2400774.S2CID 84073818.
  4. ^High-resolution vision in pelagic polychaetes
  5. ^"14 Fun Facts About Marine Bristle Worms".
  6. ^Kanie, Shusei; Miura, Daisuke; Jimi, Naoto; Hayashi, Taro; Nakamura, Koji; Sakata, Masahiko; Ogoh, Katsunori; Ohmiya, Yoshihiro; Mitani, Yasuo (2021-09-27)."Violet bioluminescent Polycirrus sp. (Annelida: Terebelliformia) discovered in the shallow coastal waters of the Noto Peninsula in Japan".Scientific Reports.11 (1): 19097.Bibcode:2021NatSR..1119097K.doi:10.1038/s41598-021-98105-6.ISSN 2045-2322.PMC 8476577.PMID 34580316.
  7. ^Zörner, S. A.; Fischer, A. (22 Dec 2006)."The spatial pattern of bioluminescent flashes in the polychaete Eusyllis blomstrandi (Annelida)".Helgoland Marine Research.61 (1):55–66.doi:10.1007/s10152-006-0053-4.ISSN 1438-3888.S2CID 2473677.
  8. ^Glasby, Cristopher; Timm, Tarmo (2008). E. V. Balian; C. Lévêque; H. Segers; K. Martens (eds.). "Global diversity of polychaetes (Polychaeta: Annelida) in freshwater".Hydrobiologia.595 (1: Freshwater Animal Diversity Assessment):107–115.Bibcode:2008HyBio.595..107G.CiteSeerX 10.1.1.655.4467.doi:10.1007/s10750-007-9008-2.S2CID 13143924.
  9. ^abGlasby, Christopher J. (1999).The Namanereidinae (Polychaeta: Nereididae). Part 1, Taxonomy and Phylogeny(PDF). Records of the Australian Museum, Supplement 25.ISBN 0-7313-8856-9. Retrieved21 October 2025.
  10. ^Annelids in Extreme Aquatic Environments: Diversity, Adaptations and Evolution
  11. ^abMartin, Daniel; Nygren, Arne; Cruz-Rivera, Edwin (2017-06-01)."Proceraea exoryxae sp. nov. (Annelida, Syllidae, Autolytinae), the first known polychaete miner tunneling into the tunic of an ascidian".PeerJ.5 e3374.doi:10.7717/peerj.3374.ISSN 2167-8359.PMC 5457667.PMID 28584710.
  12. ^abcMartin, Daniel; Britayev, Temir A. (1998)."SYMBIOTIC POLYCHAETES: REVIEW OF KNOWN SPECIES".Oceanography and Marine Biology: An Annual Review. CRC Press. pp. 225–254.doi:10.1201/b12646-22 (inactive 1 July 2025).hdl:10261/39328.ISBN 9780429210600.{{cite book}}: CS1 maint: DOI inactive as of July 2025 (link)
  13. ^Abe, Hirokazu; Hoshino, Osamu; Yamada, Kazuyuki; Ogino, Tetsuya; Kawaida, Shun; Sato-Okoshi, Waka (2022-06-28)."A novel symbiotic relationship between ascidians and a new tunic-boring polychaete (Annelida: Spionidae: Polydora)".Zootaxa.5159 (1):1–22.doi:10.11646/zootaxa.5159.1.1.ISSN 1175-5334.PMID 36095560.
  14. ^"Bristleworm".MESA.[self-published source]
  15. ^"'Zombie worms' found off Sweden".BBC News. 18 October 2005. Retrieved12 February 2010.
  16. ^Accessed Oct. 8, 2009Archived 1996-10-27 at theWayback Machine Geography of the ocean floor near Guam with some notes on exploration of the Challenger Deep.
  17. ^A tiny worm sheds light into genome compaction
  18. ^Martín-Durán, José M.; Vellutini, Bruno C.; Marlétaz, Ferdinand; Cetrangolo, Viviana; Cvetesic, Nevena; Thiel, Daniel; Henriet, Simon; Grau-Bové, Xavier; Carrillo-Baltodano, Allan M.; Gu, Wenjia; Kerbl, Alexandra; Marquez, Yamile; Bekkouche, Nicolas; Chourrout, Daniel; Gómez-Skarmeta, Jose Luis; Irimia, Manuel; Lenhard, Boris; Worsaae, Katrine; Hejnol, Andreas (2020)."Conservative route to genome compaction in a miniature annelid".Nature Ecology & Evolution.5 (2):231–242.Bibcode:2020NatEE...5..231M.doi:10.1038/s41559-020-01327-6.PMC 7854359.PMID 33199869.
  19. ^Piper, Ross (2007).Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals.Greenwood Press.ISBN 9780313339226.
  20. ^Frost, Emily; Waters, Hannah (1 July 2015)."Some polychaetes have sex lives out of a science fiction movie".14 fun facts about marine bristle worms. Smithsonian.com. Retrieved9 August 2017.
  21. ^"Oldest relative of ragworms and earthworms discovered | University of Oxford".www.ox.ac.uk. 2020-06-11. Retrieved2025-08-19.
  22. ^Chen, Hong; Parry, Luke A.; Vinther, Jakob; Zhai, Dayou; Hou, Xianguang; Ma, Xiaoya (July 2020)."A Cambrian crown annelid reconciles phylogenomics and the fossil record".Nature.583 (7815):249–252.Bibcode:2020Natur.583..249C.doi:10.1038/s41586-020-2384-8.ISSN 1476-4687.PMID 32528177.
  23. ^Butterfield, N. J. (1990). "A reassessment of the enigmatic Burgess Shale fossilWiwaxia corrugata (Matthew) and its relationship to the polychaeteCanadia spinosa Walcott".Paleobiology.16 (3):287–303.Bibcode:1990Pbio...16..287B.doi:10.1017/S0094837300010009.JSTOR 2400789.S2CID 88100863.
  24. ^Smith, M. R. (2012)."Mouthparts of the Burgess Shale fossilsOdontogriphusandWiwaxia: Implications for the ancestral molluscan radula".Proceedings of the Royal Society B.279 (1745):4287–4295.doi:10.1098/rspb.2012.1577.PMC 3441091.PMID 22915671.
  25. ^Smith, M. R. (2014)."Ontogeny, morphology and taxonomy of the soft-bodied Cambrian 'mollusc'Wiwaxia".Palaeontology.57 (1):215–229.Bibcode:2014Palgy..57..215S.doi:10.1111/pala.12063.S2CID 84616434.
  26. ^Miller, A. J. (2004).A revised morphology ofCloudina with ecological and phylogenetic implications.CiteSeerX 10.1.1.526.5035.
  27. ^Vinn, Olev; Zatoń, Michał (March 2012)."Inconsistencies in proposed annelid affinities of early biomineralized organismCloudina (Ediacaran): structural and ontogenetic evidences".Carnets de Géologie (Lettres).doi:10.4267/2042/46095.
  28. ^Vinn, O; Mutvei, H (2009)."Calcareous tubeworms of the Phanerozoic".Estonian Journal of Earth Sciences.58 (4): 286.Bibcode:2009EsJES..58..286V.doi:10.3176/earth.2009.4.07.
  29. ^Westheide, W. (1997). "The direction of evolution within the Polychaeta".Journal of Natural History.31 (1):1–15.Bibcode:1997JNatH..31....1W.doi:10.1080/00222939700770011.
  30. ^Capa, Maria; Hutchings, Pat (March 2021)."Annelid Diversity: Historical Overview and Future Perspectives".Diversity.13 (3): 129.Bibcode:2021Diver..13..129C.doi:10.3390/d13030129. Retrieved5 July 2025.

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