Overview

Brief Summary

Polychaeta, composed of about 10,000 species, is the larger (and apparently not monophyletic) of the two generally recognized major groups of segmented worms (phylum Annelida) – the other being the Clitellata (earthworms and leeches).

Polychaete worms are characterized by an elongated, metameric body usually bearing a pair of appendages called parapodia on each metamere (segment), as well as tufts of chaetae (spines served by muscles which typically can be extended and retracted; often the polychaetes are called bristle worms, and their name derives from the Latin for many bristles). Parapodia show vast diversity of form and function, serving purposes such as locomotion, gas exchange, protection, attachment, controlling water flow within a tube, or can be reduced or lost altogether. The polychaete head can be adorned with a multitude of sensory structures such as tentacular palps, antennae, and cirri. Predatory carnivores often have large pharyngeal jaws. At the end of the segmented body is the tail, called the pygidium, which houses the anus (Brusca and Brusca 2003)

Some polychaetes are free-living, with well developed muscles that allow them to move by swimming, crawling, or burrowing, often aided by parapodia adapted as paddles or legs. Burrowers often have a muscular proboscis to aid in digging. In contrast, sedentary polychaetes feed from permanent tubes or burrows, often by suspension feeding, selective deposit feeding, or feeding on detritus. Their parapodia are often adapted for circulating water in the tube. Permanent tube-dwellers have softer and less muscular bodies and frequently lose the septa between segments. This allows for adjustment of hydrostatic pressure within the worm, which is important for functions such as anchoring the end of the body housed in its tube. As well as providing protection, tubes also function as external support for these worms. Tubes can be soft, parchment-like forms constructed from sand and mucus, or hard calcareous tubes, which when many worms are together, form reef structures (Brusca and Brusca 2003)

Reflecting the large diversity of lifestyles and degree of independence of body segments, polychaete circulatory and respiratory systems also show many variations among taxa. Gas exchange in many polychaetes is facilitated by distinct gills, but in others it occurs across the entire body surface (especially in small or sedentary taxa with no parapodia, or in worms with no or partial internal body septa to separate coelomic spaces). Some taxa increase surface respiratory areas with feathery protrusions of the body surface through which the coelom extends. In these taxa, the circulatory system is reduced, and most oxygen and nutrients are distributed in the coelomic fluid. Some taxa have pumping structures to increase blood flow, especially sedentary worms that do not use body movements to circulate the blood. Most (but not all) polychaetes have oxygen-carrying pigments in their circulatory fluid and coelomic fluid, usually a form of haeomoglobin. Polychaetes display a large array of different sensory structures, including touch receptors; photoreceptors which may be developed into one or more pairs of anteriorly positioned eyes or distributed around the body; chemoreceptors, and statocysts.

(Brusca and Brusca 2003; Kozloff 1990)

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Comprehensive Description

Polychaete worms are characterized by an elongated, metameric body usually bearing a pair of appendages called parapodia on each metamere (segment), as well as tufts of chaetae (spines served by muscles which typically can be extended and retracted; often the polychaetes are called bristle worms, and their name derives from the Latin for many bristles). The body segmentation is visible as lateral lines around the worm’s body, reflecting the internal separation of segments with septa (although septa are lost or reduced in some groups, especially tube dwellers, see below). Parapodia show vast diversity of form and function, serving purposes such as locomotion, gas exchange, protection, attachment, controlling water flow within a tube, or can be reduced or lost altogether. The polychaete head can be adorned with a multitude of sensory structures such as tentacular palps, antennae, and cirri. Predatory carnivores often have large pharyngeal jaws. At the end of the segmented body is the tail, called the pygidium, which houses the anus (Brusca and Brusca 2003)

A common organization of the polychaetes is to divide them into sedentary forms and free-living forms. Although this organization does not reflect their genetic relationships, it does illustrate the adaptation of their body form to their habitat and lifestyle. The free-living forms, which include families of carnivorous predators as well as direct deposit feeders – (free-living worms that burrow ingest the sediment sift out food particles in their gut) are commonly composed of a series of identical body segments (a homonomous body plan). They have well developed muscles and move by swimming, crawling, or burrowing with their parapodia adapted as paddles or legs. Burrowers often have a muscular proboscis to aid in digging. In contrast, the body segments of sedentary, tube-dwelling polychaetes show specializations for different functions (heterotomous form). These worms feed from permanent tubes or burrows, often by suspension feeding, selective deposit feeding or feeding on detritus. Their parapodia are often adapted for circulating water in the tube. Permanent tube-dwellers have softer and less muscular bodies, and frequently lose the septa between segments. This allows for adjustment of hydrostatic pressure within the worm, which is important for functions such as anchoring the end of the body housed in its tube. As well as providing protection, tubes also function as external support for these worms. Tubes can be soft, parchment-like forms constructed from sand and mucus, or hard calcareous tubes, which when many worms are together, form reef structures (Brusca and Brusca 2003)

Reflecting the large diversity of lifestyles and degree of independence of body segments, polychaete circulatory and respiratory systems also show many variations among taxa. Almost all polychaetes have a closed circulatory system. Many have distinct gills, usually adapted as highly vascularized parts of the parapodia, and circulatory systems are well-developed with a pair longitudinal vessels carrying blood in the anterior (dorsal vessel) and posterior (ventral vessel) directions along the full body of the worm. Gas exchange in others occurs across the entire body surface (especially in small or sedentary taxa with no parapodia, or in worms with no or partial internal body septa to separate coelomic spaces). Some taxa increase surface respiratory areas with feathery protrusions of the body surface through which the coelom extends. In these taxa the circulatory system is reduced, and most oxygen and nutrients are distributed in the coelomic fluid. Some taxa have pumping structures to increase blood flow, especially sedentary worms that do not use body movements to circulate the blood. Most (but not all) polychaetes have oxygen-carrying pigments in their circulatory fluid and coelomic fluid, usually a form of haeomoglobin. Some taxa have more than one pigment. The pigments that are present often have adaptive value for the animal’s lifestyle, for example intertidal dwellers have the ability to hold oxygen during high tides and release it during low tides. Almost all polychaetes have metanephridia allowing for each coelomic space to eliminate waste, osmoregulate, and spawn gametes. The nervous system includes a cerebral ganglion at the head and one or more longitudinal nerves running the length of the body with an associated pair of ganglia in each segment. Polychaetes display a large array of different sensory structures, including touch receptors; photoreceptors which may be developed into one or more pairs of anteriorly positioned eyes or distributed around the body; chemoreceptors, and statocysts.

(Brusca and Brusca 2003; Kozloff 1990)

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Physical Description

Morphology

Sexual Dimorphism

Females generally larger than males and sometimes thicker-bodied; Sexual Dimorphism also in number of nephridial papillae, genital papillae, position of the genital opening and shape of the pygidial appendages; Sexual dimorphism can be extreme with dwarf males showing great reduction in size and simplification of morphology; males may live within the female’s body.
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Type Information

Holotype for
Catalog Number: USNM 32683
Collection: Smithsonian Institution, National Museum of Natural History, Department of Invertebrate Zoology
Preparation: Alcohol (Ethanol)
Collector(s): C. Williamson
Locality: Mitlanatch Island, Mitlenatch Island, British Columbia, Canada, Gulf of Georgia, North Pacific Ocean
Depth (m): 183 to 183
  • Holotype:
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Ecology

Habitat

The polychaete worms (in the traditional inclusion of taxa) are mostly marine. They are commonly found burrowing in sediments on beaches, or live in tubes, which in cases where many worms live together, form calcareous reef structures. Some species are free-swimming, some live as commensals or parasites. Polychaetes are the most abundant macrofauna of the deep sea, and inhabit the world’s oceans at all different depths and water temperatures.

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Associations

Known predators

Polychaeta (Zooplankton) is prey of:
Actinopterygii
mixed-food consumers
secondary carnivores
Pleuronichthys verticalis
Citharichthys xanthostigma
Engraulidae
Calidris ferruginea
Calidris alba
Ovalipes
Erynnis japonica
Limanda yokohamae
Limanda herzensteini
Chaeturichthys hexanema
Concholepas concholepas
Sicyases sanguineus
Nematoda
Gambusia
Heterandria formosa
Decapoda
Floridichthys carpio
Lophogobius cyprinoides
high carnivores
Copepoda
Callinectes sapidus
Chondrichthyes
Scombridae
decomposers/microfauna
organic stuff
Octopus
Cephalopoda
Stomatopoda
Anomura
Asteroidea
Echinoidea
Gastropoda
Priapula
Polychaeta
Ophiuroidea
Cancer
Brachyura
Ammodytes marinus
Clupea harengus
Alosa pseudoharengus
Scomber
Peprilus triacanthus
Actinonaias ellipsiformis
Tridonta arctica
Pollachius pollachius
Merluccius bilinearis
Urophycis regia
Urophycis tenuis
Urophycis chuss
Gadidae
Melanogrammus aeglefinus
Hemitripterus americanus
Myoxocephalus octodecemspinosus
Leucoraja erinacea
Leucoraja ocellata
Amblyraja radiata
Macrozoarces americanus
Brosme brosme
Anarhichas
Tautogolabrus adspersus
Triglidae
Sebastes marinus
Pleuronectes ferrugineus
Scophthalmus aquosus
Paralichthys dentatus
Glyptocephalus cynoglossus
Hippoglossina oblonga
Pleuronectes americanus
Hippoglossoides platessoides
Hippoglossus hippoglossus
Mustelus canis
Squalus acanthias
Lophius americanus
Cynoscion
Pomatomus saltatrix
Other suspension feeders
Mya arenaria
Crassostrea virginica
Nereis
meiofauna
Alosa chrysochloris
Anchoa mitchilli
Brevoortia tyrannus
Alosa sapidissima
Micropogonius undulatus
Trinectes maculatus
Morone americana
Arius felis
Paralichthyes albigutta
Strongylura marina
Urophycis floridana
Prionotus scitulus
Prionotus tribulus
Gobiosoma robustum
Microgobius gulosus
Lagodon rhomboides
Leiostomus xanthurus
Syngnathus scovelli
Hippocampus zosterae
Laridae
Cyprinodon variegatus
Anatidae
Fundulus confluentus
Fundulus similis
Adinia xenica
suspended particulate carbon
Sabellidae
Serpulidae

Based on studies in:
Antarctic (Estuarine)
USA: California, Southern California (Marine, Sublittoral)
South Africa (Desert or dune)
USA: Florida, Everglades (Estuarine)
Puerto Rico, Puerto Rico-Virgin Islands shelf (Reef)
USA: Maryland, Chesapeake Bay (Estuarine)
unknown: Black Sea (Marine)
Pacific (Marine)
Japan (Coastal, mesopelagic zone)
Chile, central Chile (Littoral, Rocky shore)
USA: Florida (Estuarine)
USA, Northeastern US contintental shelf (Coastal)

This list may not be complete but is based on published studies.
  • A. C. Brown, Food relationships on the intertidal sandy beaches of the Cape Peninsula, S. Afr. J. Sci. 60:35-41, from p. 39 (1964).
  • Baird D, Ulanowicz RE (1989) The seasonal dynamics of the Chesapeake Bay ecosystem. Ecol Monogr 59:329–364
  • Christian RR, Luczkovich JJ (1999) Organizing and understanding a winter’s seagrass foodweb network through effective trophic levels. Ecol Model 117:99–124
  • G. A. Knox, Antarctic marine ecosystems. In: Antarctic Ecology, M. W. Holdgate, Ed. (Academic Press, New York, 1970) 1:69-96, from p. 87.
  • J. C. Castilla, Perspectivas de investigacion en estructura y dinamica de communidades intermareales rocosas de Chile Central. II. Depredadores de alto nivel trofico, Medio Ambiente 5(1-2):190-215, from p. 203 (1981).
  • Link J (2002) Does food web theory work for marine ecosystems? Mar Ecol Prog Ser 230:1–9
  • M. A. Hatanaka, Sendai Bay. In: Productivity of Biocenoses in Coastal Regions of Japan, K. Hogetsu, M. Horanaka, T. Hatanaka, T. Kawamura, Eds. (Japanese Committee for the International Biological Program Synthesis, Tokyo, 1977), 14:173-221, from p. 190.
  • M. E. Vinogradov and E. A. Shushkina, Some development patterns of plankton communities in the upwelling areas of the Pacific Ocean. Mar. Biol. 48:357-366, from p. 359 (1978).
  • Opitz S (1996) Trophic interactions in Caribbean coral reefs. ICLARM Tech Rep 43, Manila, Philippines
  • T. A. Clark, A. O. Flechsig, R. W. Grigg, Ecological studies during Project Sealab II, Science 157(3795):1381-1389, from p. 1384 (1967).
  • T. S. Petipa, E. V. Pavlova, G. N. Mironov, The food web structure, utilization transport of energy by trophic levels in the planktonic communities. In: Marine Food Chains, J. H. Steele, Ed. (Oliver and Boyd, Edinburgh, 1970), 142-167 from p. 155.
  • T. S. Petipa, E. V. Pavlova, G. N. Mironov, The food web structure, utilization transport of energy by trophic levels in the planktonic communities. In: Marine Food Chains, J. H. Steele, Ed. (Oliver and Boyd, Edinburgh, 1970), 142-167, from p. 154.
  • W. E. Odum and E. J. Heald, The detritus-based food web of an estuarine mangrove community, In Estuarine Research, Vol. 1, Chemistry, Biology and the Estuarine System, Academic Press, New York, pp. 265-286, from p. 281 (1975).
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Known prey organisms

Polychaeta (Zooplankton) preys on:
benthic filter feeding invertebrates
detritus
phytoplankton
saprophagous plankton
meroplankton
Appendicularia
Doliolidae
Calanoida
Euphausiidae
Cyclopoidea
Cnidaria
Tomopteridae
Cumacea
Turbellaria
protozoa
suspended or deposited organic matter
benthic invertebrates
Mysidacea
Ostracoda
Euphausiacea
Hyperiidea
Cyclopoida
algae
fungi
bacteria
Isopoda
Amphipoda
Pycnogonidae
Tanaidae
Gastropoda
Priapula
Polychaeta
Ophiuroidea
Bivalvia
Ectoprocta
Cirripedia
Ascidia
Porifera
Anthozoa
Bacteria attached to suspended POM
Bacteria attached to sediment POM
Bacillariophyceae
microzooplankton
zooplankton
Ctenophora
Chrysaora quinquecirrha
Other suspension feeders
Mya arenaria
Nereis
Macoma
meiofauna
Crustacea
Callinectes sapidus
bacterioplankton
Microprotozoa

Based on studies in:
Antarctic (Estuarine)
unknown: Black Sea (Marine)
USA: California, Southern California (Marine, Sublittoral)
Pacific (Marine)
South Africa (Desert or dune)
Chile, central Chile (Littoral, Rocky shore)
USA: Florida, Everglades (Estuarine)
USA, Northeastern US contintental shelf (Coastal)
Puerto Rico, Puerto Rico-Virgin Islands shelf (Reef)
USA: Maryland, Chesapeake Bay (Estuarine)
USA: Florida (Estuarine)
Japan (Coastal, mesopelagic zone)

This list may not be complete but is based on published studies.
  • A. C. Brown, Food relationships on the intertidal sandy beaches of the Cape Peninsula, S. Afr. J. Sci. 60:35-41, from p. 39 (1964).
  • Baird D, Ulanowicz RE (1989) The seasonal dynamics of the Chesapeake Bay ecosystem. Ecol Monogr 59:329–364
  • Christian RR, Luczkovich JJ (1999) Organizing and understanding a winter’s seagrass foodweb network through effective trophic levels. Ecol Model 117:99–124
  • E. A. Shushkina and M. E. Vinogradov, Trophic relationships in communities and the functioning of marine ecosystems: II. Some results of investigations on the pelagic ecosystem in tropical regions of the ocean. In: Marine Production Mechanisms, M. J. Dun
  • G. A. Knox, Antarctic marine ecosystems. In: Antarctic Ecology, M. W. Holdgate, Ed. (Academic Press, New York, 1970) 1:69-96, from p. 87.
  • J. C. Castilla, Perspectivas de investigacion en estructura y dinamica de communidades intermareales rocosas de Chile Central. II. Depredadores de alto nivel trofico, Medio Ambiente 5(1-2):190-215, from p. 203 (1981).
  • Link J (2002) Does food web theory work for marine ecosystems? Mar Ecol Prog Ser 230:1–9
  • M. A. Hatanaka, Sendai Bay. In: Productivity of Biocenoses in Coastal Regions of Japan, K. Hogetsu, M. Horanaka, T. Hatanaka, T. Kawamura, Eds. (Japanese Committee for the International Biological Program Synthesis, Tokyo, 1977), 14:173-221, from p. 190.
  • M. E. Vinogradov and E. A. Shushkina, Some development patterns of plankton communities in the upwelling areas of the Pacific Ocean. Mar. Biol. 48:357-366, from p. 359 (1978).
  • Opitz S (1996) Trophic interactions in Caribbean coral reefs. ICLARM Tech Rep 43, Manila, Philippines
  • T. A. Clark, A. O. Flechsig, R. W. Grigg, Ecological studies during Project Sealab II, Science 157(3795):1381-1389, from p. 1384 (1967).
  • T. S. Petipa, E. V. Pavlova, G. N. Mironov, The food web structure, utilization transport of energy by trophic levels in the planktonic communities. In: Marine Food Chains, J. H. Steele, Ed. (Oliver and Boyd, Edinburgh, 1970), 142-167 from p. 155.
  • T. S. Petipa, E. V. Pavlova, G. N. Mironov, The food web structure, utilization transport of energy by trophic levels in the planktonic communities. In: Marine Food Chains, J. H. Steele, Ed. (Oliver and Boyd, Edinburgh, 1970), 142-167, from p. 154.
  • W. E. Odum and E. J. Heald, The detritus-based food web of an estuarine mangrove community, In Estuarine Research, Vol. 1, Chemistry, Biology and the Estuarine System, Academic Press, New York, pp. 265-286, from p. 281 (1975).
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Life History and Behavior

Life Cycle

Polychaete eggs undergo spiral cleavage. Many develop into a free-swimming trochophore larva. This larva grows at the “growth zone” on the posterior end, by forming sequential serial segments in a process called teloblastic growth. After a period of larval elongation, the larvae settles from the plankton and becomes a juvenile worm. Other polychaetes undergo direct development (without a larval stage), or short, non-feeding trochophore stage (Brusca and Brusca 2003)

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Reproduction


Most polychaetes can regenerate to some degree – from regenerating lost appendages to posterior body segments. Some polychaetes reproduce asexually by breaking into two or more groups of segments, and some reproduce very efficiently this way by “multiple fragmentation”, in which each segment becomes a new individual (for example Dodecaceria (Cirratulidae)).

Polychaetes are almost all dioecious, although they do not have distinct gonads. Instead, patches of the peritoneum lining the coelem (in one or many segments, depending on the taxon) divide to produce prospective gametes, which then break off and fully mature into eggs or sperm in the coelom. Eggs and sperm are released from the worm’s coelom through nephridial or coelomic ducts to the outside, or through breaks in the worm’s body wall during spawning. Most polychaetes have external fertilization, although some species brood their young. In order to maximize fertilization some primarily-benthic polychaetes (characteristic in the families Nereidae, Eunicidae, and Sillidae) spawn while swarming at the top of the water column. To do this, these worms transform into a swimming, sexual form quite different from the benthic form (called epitoky). One way this is done, as seen in members of the families Nereidae and Eunicidae, is by completely transforming the whole body into a sexual individual. Some notable modifications include enlarging swimming parapodia at the anterior end and often developing large eyes. Epitoke formation is stimulated by hormones in the brain, which are found only in older worms. Another method is to bud off the posterior portion of the body to form the sexual epitoke. This happens in the Syllidae. A third method of forming a sexual form is for a hind portion with the gametes to break off. This is not a true epitoke because the swimming section is not a complete worm. Palola viridis (Eunicidae) is an example of a species that makes this type of swarmer. Lunar cycles trigger the swarming event, and this species swarms in such huge numbers that natives of the samoan islands, where they are found, collect and feast on them (Kozloff 1990; Brusca and Brusca 2003)

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Evolution and Systematics

Evolution

Classification

Several minor orders presented in Fauchald (1977)are now regarded as superfluous. All orders so tagged do not have child taxa that are currently in use. Basically the classification of Rouse & Pleijel (2001: Fig.1.1) is presented here, with old name Errantia substituting for their Aciculata and old name Sedentaria including their Canalipalpata and Scolecida, but if the indications of placements in Zrzavý et al (2009) are implemented, then Scolecida is polyphyletic and its families should disperse into Terebellida and Sabellida. Further, Struck (2011) has introduced Clade Pleistoannelida defined as the last common ancestor of a revived Errantia and Sedentaria, with the latter containing Clitellata (leeches and earthworms). This view is partly integrated into the WoRMS arrangement of Annelida taxa, although his new Pleistoannelida may be superfluous if Polychaeta is retained. Palpata as a subclass appears likely to be superfluous and is not currently used here. Some interstitial annelid families (including former 'archiannelids') are under 'Polychaeta incertae sedis' awaiting assignment, together with some other taxa of obscure or Polychaeta-basal affinities.
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Systematics and Taxonomy

The polychaete worms are problematic in that molecular phylogenetic analyses carried out the last 20 years indicate that they are not a natural (monophyletic) group; rather, this clade includes the phylums Sipunculata and Echiura, as well as a small enigmatic group of worms called the beard worms (pogonophorans+vestimentiferans; about 100 species). In addition, many analyses also now suggest that Clitellata, which has long been considered the sister group to the polychaetes, is actually a derived group of worms within the polychaetes (McHugh 1997, 2005, Struck et al 2007).

Fossil polychaetes have been found dating back to the early Cambrian. They are known mostly from fossilized jaws and mineralized tubes (Brusca and Brusca 2003).

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
Specimen Records:18693
Specimens with Sequences:12902
Specimens with Barcodes:11575
Species:1949
Species With Barcodes:1501
Public Records:10739
Public Species:791
Public BINs:1996
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Genomic DNA is available from 1 specimen with morphological vouchers housed at Museums and Art Galleries of the Northern Territory
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