Polydora cornuta

Invasion History

First Non-native North American Tidal Record: 1932
First Non-native West Coast Tidal Record: 1932
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Polydora cornuta was described from Charleston, South Carolina by Bosc in 1802, and later widely found on the East Coast of the United States from the Gulf of St. Lawrence to the Gulf of Mexico (Blake 1971; Blake and Maciolek 1987). It was reported from northern Europe as early as 1859 (Claparede 1861), but its range and invasion status are complicated by confusion with the similar species, P. ciliata, which is usually found boring in shells and calcareous substrates (Wolff 1973; Radashevsky 2005). It is apparently introduced in the Black and Mediterranean Seas (Cinar et al. 2005; Surugiu 2012; Radashevsky and Selifonova 2013), the Southwestern Atlantic (Brazil-Argentina, Orensanz et al. 2002; Radashevsky 2005), the Northeast Pacific (British Columbia to Mexico, Hartman 1941; Light 1977; Carlton 1979; Blake and Ruff 2007), the Northwest Pacific (China-Russia, Sato-Okoshi 1999; Radashvesky 2005), and the Southwest Pacific (Australia-New Zealand, Read 1975; Blake and Kudenov 1978; Walker 2009).

North American Invasion History:

Invasion History on the West Coast:

Polydora cornuta was first collected from Ladysmith, British Columbia (BC) in 1932, and found at the San Juan Islands, Washington (WA), San Francisco Bay, and Mission Bay, California (CA) from 1932 to 1938 (Carlton 1979; Hartman 1941; U.S. National Museum of Natural History 2015). It was subsequently found at many intermediate estuaries, including Grays Harbor, WA (1st record 1953, U.S. National Museum of Natural History 2015); Willapa Bay, WA (1st record 2000, Cohen et al. 2001); the Columbia River estuary, WA – Oregon (OR) (1980, Jones 1983); Coos Bay, OR (Hartman and Reish, 1950; cited by Carlton 1979); Humboldt Bay, CA (Barnhart 1992, cited by Boyd et al. 2002); Elkhorn Slough, CA (1st record 1977, Wasson et al. 2001); and Morro Bay, CA (1st record 1960, Reish and Barnard 1967). Except for the Columbia River, all of these estuaries had received transplants of Eastern Oysters (Crassostrea virginica) (Carlton 1979).

In southern California, although few Eastern Oyster transplants were made, Polydora cornuta has been collected, often in high abundance, both in major harbors for commercial shipping and small bays used by recreational boats. For instance, records are known from Los Angeles-Long Beach Harbor (1st record 1966, Crippen and Reish 1969 cited by Carlton 1979); San Diego Bay (1st record 1994, Levin et al. 1998), Playa del Rey Lagoon, Venice Beach (1st record 1941, Alan Hancock Foundation, cited by Carlton 1979); Alamitos Bay (1st record 1952, Reish and Winter 1954, cited by Carlton 1979); Anaheim Bay (1st record 1972, Reish et al. 1975); and Santa Catalina Island (1st record 1963, Reish et al. 1964, cited by Carlton 1979). Polydora cornuta has also been introduced to the Salton Sea, a land-locked artificial saline lake, and was first collected there in 1961 (Carlton 1979; USNM 39642, U.S. National Museum of Natural History 2015).

It is likely that Polydora cornuta has been transferred from the East to the West Coast many times, with Eastern Oyster transplants, hull fouling, and ballast water, and later moved to additional estuaries by hull fouling, and perhaps by other vectors, such as discarded shellfish, or seaweed used to pack bait.

Invasion History Elsewhere in the World:

In Europe, Polydora cornuta was reported from Scotland in 1859 (Claparede 1861), but was often confused with P. ciliata, although the latter species is predominantly associated with shells and other calcareous substrates (Wolf 1973; Michaelis 1978; Ramberg and Schram 1982; Surugiu 2012). Wolff (1973) notes that it was collected in the Netherlands in 1930, but identified then as P. ciliata. It is known from the Scheldt, Meuse and Rhine estuaries in the Netherlands (Wolff 1973); Langeoog Island and the Jade-Weser Delta, Germany (U.S. National Museum of Natural History 2015; Michaelis 1978); and the inner Oslofjord, Norway (Ramberg and Schram 1982). It is also known from the western Baltic, including the Isefjord, Denmark (Hansen et al. 2010), and the Kiel Bight and Mecklenberg Bay, Germany (Anger 1977; Radashevsky 2005; U.S. National Museum of Natural History 2015). Records on the outer Atlantic coast of Europe are rare and scattered, including the Isle of Skye/Scotland (Claparede 1861), Marennes-Oleron Bay, France (de Montaudouin and Sauriau 2000), and the Óbidos lagoon, Portugal (Carvalho et al. 2011). The frequent confusion between P. ciliata and P. cornuta makes it difficult to determine the invasion status of P. cornuta, at least in northern Europe. Genetic studies would be useful in resolving the origin of these worms.

In the Black Sea, there were apparently no non-boring Polydora collected before 1962, and in the Mediterranean, records of P. cornuta are recent, widely scattered, and located near harbors. These include Valencia, Spain (1st record 1990, Tena et al. 1991); the Venice Lagoon, Italy (1st record 1991, Tagliapietra et al. 1998); Izmir Bay, Turkey (1st record 2003, Cinar et al. 2005); and the Saronikos Gulf, Greece (1st record 2008, Simboura et al. 2008). In the Black Sea proper, P. cornuta was first collected off Odessa, Ukraine in 1962, and initially identified as P. ciliata limicola (Losovskaya and Zolotarev 2003). It is now widespread in the Black Sea, the Sea of Marmara, and the Sea of Azov (Dagli and Ergen 2008; Karhan et al. 2008; Radashevsky and Selifonova 2013).

In the Southwestern Atlantic, Polydora cornuta was first collected in the Mar Chiquita Lagoon, Mar Del Plata, Argentina in 1967 (Orensanz et al. 2002; Radashevsky 2005; U.S. National Museum of Natural History 2015) and later, in Rio de Janeiro Harbor (in 1986) and Paranagua Bay (in 1998), Brazil (Radashevsky et al. 2005; U.S. National Museum of Natural History 2015). In the Northwest Pacific, it was first reported from the east coast of Taiwan, on the Formosa Straits (in 1990, Radashevsky and Hsieh 2000; Radashevsky 2005). Other early records are from Shihwa Lake, South Korea, on the Yellow Sea (in 1994, Lee and Cha, 1997, cited by Radashevsky 2005) and Vladivostok, Russia (in 1994, Radashevsky 2005). Polydora cornuta was collected at several ports on the southeastern and west coasts of Japan (1984-1997, Sato-Okoshi 2000). One puzzling record came from the Sea of Japan, off Russia, at 3334 m depth (in 2010, Alalykina 2013). This polychaete is also known from two ports in New Zealand: Whangarei and Waitemata Harbors (Read 1975; Read and Gordon 1991), and from Victoria (Blake and Kudenov 1978) and Botany Bay, Australia (in 1973, Walker 2009).

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Description

Adults of the 'Polydora cornuta' species complex have up to 90 segments. The prostomium has a lateral incision, forming two lobes and is prolonged back to chaetiger 3 as a caruncle. Most specimens have 4 eyes in a trapezoidal arrangement, on the prostomium, with a small, triangular, occipital tentacle located posteriorly to them. A ciliated groove runs on either side of the caruncle. The prostomium is flanked by a pair of long, prehensile palps, which stretch back to chaetigers 15-35. The palps each have a ciliated groove along their lengths, bordered by a fine black line, but lack conspicuous pigment.

Chaetiger 1 has well-formed parapodial lobes, but has chaetae (fine, capillary) only on the neuropodia. The notopodia lack chaetae, but have long, finger-like lamellae. Chaetigers 2-4 and 6 have bundles of capillary neurochaetae and notochaetae. However, chaetiger 5 is modified and bears a row of 6-8 stout spines with adhering companion chaetae, and with dorsal musculature running back to chaetigers 6 and 7. Chaetigers from 7 on, have long, thin branchiae and up to 15 stout hooded hooks on the neuropodia, while notopodial chaetae decrease in number posteriorly. The pygidium is flared and resembles a suction cup. Adult worms reach a length of 12-32 mm and are light tan in color. Smaller worms have black melanophores laterally from chaetigers 2-7 to 10-22, and yellow chromatophores ventrally from chaetigers 5-7 to 16-21. Larval development is described (as P. ligni) by Blake (1969). Description based on: Blake 1971, Blake and Kudenov 1978, Light 1978, Blake 1983, Blake and Maciolek 1987, Radashevsky and Hsieh 2000, Cinar et al. 2005, Radashevsky 2005, and Radashevsky and Selifonova 2013; Bertasi 2016.

The species was first described in 1805 from Charleston, North Carolina, but was later known more widely under the name Polydora ligni (Webster 1879, described from New Jersey). The name 'Polydora cornuta' was re-instituted and the animal redescribed by Blake and Maciolek (1987). Populations of P. cornuta around the world show small differences in morphology and substantial differences in size which may indicate both genetic and environmental differences (Radashevsky and Hsieh 2000; Cinar et al. 2005). Rice (1991) and Rice et al. (2008) found that populations of P. cornuta from Florida, Maine, and California each showed genetic differences and low frequency of hybridization indicating their status as cryptic species. However worms from North Carolina and California were interfertile (Rice 1991), and California worms were interfertile with worms from New Zealand (Rice et al. 1998). Further genetic comparisons among populations are needed to trace the likely anthropogenic spread of this species.

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Taxonomy

Ecology

General:

'Polydora cornuta' is an estuarine spionid polychaete, which dwells in mud and mucus tubes in sediment, and in fouling communities on hard surfaces (Blake 1971; Blake and Maciolek 1987). 'Polydora cornuta' is a widely distributed complex of cryptic species, probably varying in environmental tolerances and life history (Rice et al. 2008). The sexes are separate (Rice 1991). Eggs are laid in capsules, linked in strings attached to the interior of the tubes. Capsules in Maine estuaries contained up to 132 eggs, but about half of those were unfertilized and used as food by developing larvae (Blake 1969; Gibson et al. 2012). Larvae are released from the capsules at the 3-chaetiger stage, and feed in the plankton (Blake 1969). Larvae settle at about 12-22 chaetigers, and ~1.1-1.3 mm length (Blake 1969; Orth 1971; Levin 1984; Anger et al. 1986; Radashevsky 2005). Development is strongly effected by temperature, with time to settlement shortening with increasing temperatures (60 days at 10C ; 16-28 days at 12C; 12-41 days at 18C; 7-14 days at 20C, 12 days at 28C) (Orth 1979; Levin 1984, Anger et al. 1986; Takata et al. 2011). Food supply can also effect developmen - two weeks of partial starvation led to an approximate doubling of time to settlement, from ~19 days to 42 days at 11C (Qian and Chia 1993). The wide variation in development times with temperature may reflect laboratory food supply, as well as genetic differences in populations from different regions. Settlement is stimulated by the presence of sediment containing microorganisms (Sebesvari et al. 2006). Growth to first reproduction is rapid, 33 days in culture at 18 C. Adults in culture, survived for an average of 13 months (Anger et al. 1986). In a Brazilian population, some males were mature at 2 mm length and females at 5 mm (Radashevsky 2005), but adult lengths of 10-20 mm are more typical (Levin 1984) and adults can reach 32 mm (Radashevsky 2005).

'Polydora cornuta' is known from a wide range of climates, and estuarine-marine habitats from the tropics to cold-temperate waters, although some of this range may result from the occurrence of cryptic species. Its habitats include subtidal and intertidal soft substrates, fouling communities in marinas and harbors, ships' hulls, and oyster beds (Graham and Gay 1945; Wolff 1973; Bagaveeva and Zvyagintsev 2000). This worm's habitats are often exposed to temperature and salinity extremes, as well as hypoxia, exposure and toxic chemicals (Wolff 1973; Levin 1984; Cinar et al. 2005). Polydora cornuta occurs in water prone to sea ice formation (Blake 1969), and has been collected from salinities of 29C (Cory 1967) in Chesapeake Bay, and probably occurs at higher temperatures. It has been collected at a salinity range of 2-75 PSU, and is abundant at salinities ranging from 5 to 75 PSU (Wolff 1973; Simmons 1957). Polydora cornuta is also tolerant of hypoxic conditions and is considered an indicator species for pollution (Anger 1977; Rice et al. 1980; Cinar et al. 2005; Levin et al. 2009).

Its larvae are planktotrophic, feeding on phytoplankton (Anger et al. 1986). Adults of P. cornuta are capable of shifting between deposit-feeding detritus and surface-microalgae, and suspension-feeding (phytoplankton and suspended particles). Their ciliated palps can either stretch over the sediment surface to pick up sediment particles, or wave in the current to pick up phytoplankton (Dauer et al. 2005). Feeding and growth rates of P. cornuta increased in simulated tidal currents (Hentschel and Herrick 2005). Although P. cornuta is generally considered a suspension and deposit-feeder, predation on small bivalves (veligers of several species, juveniles of Gemma gemma) has been documented (Breese and Phibbs 1972; Weinberg 1984). The large populations of P. cornuta in eutrophic waters are a potential food source for fishes and invertebrates (Oviatt et al. 1987). Sublethal predation on P. cornuta, with predators biting off parts of worms, with subsequent regeneration, was common in Long Island Sound (Zajac 1995). However, hypoxic conditions can limit the availability of these worms to fishes and other active predators.

Food:

Phytoplankton; benthic microalgae, detritus

Consumers:

Fishes, shrimps, crabs

Competitors:

Other polychaetes, Macoma spp.

Trophic Status:

Deposit Suspension Feeder

DepSusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Unstructured Bottom	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Salt-brackish marsh	None
General Habitat	Grass Bed	None
General Habitat	Vessel Hull	None
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Endobenthic	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	-2	Based on geographical range
Maximum Temperature (ºC)	29	Field, Patuxent River, Chesapeake Bay (Cory 1967). This polychaete probably tolerates higher temperatures.
Minimum Salinity (‰)	2	Field, abundant above 5 PSU (Wolff 1973)
Maximum Salinity (‰)	75	Field, Laguna Madre, Texas (Simmons 1957)
Minimum Dissolved Oxygen (mg/l)	1.4	Field, Dauer et al. 1992, cited by Levin et al. 2009
Minimum Reproductive Temperature	10	Laboratory, Chesapeake Bay (Orth 1971)
Maximum Reproductive Temperature	29	Field, Patuxent River, Chesapeake Bay (Cory 1967). This polychaete probably tolerates ligher temperatures.
Minimum Reproductive Salinity	5	Field, Patuxent River, Chesapeake Bay (Cory 1967). This was the approximate lower limit for settlement
Minimum Duration	9	18 C (Anger et al. 1986) Development is probably much shorter at higher temperatures.
Maximum Duration	60	10 C (Orth 1971)
Minimum Length (mm)	2	Smallest mature male, Brazil. Females were mature at 5.5 mm (Radashevsky 2005).
Maximum Length (mm)	32	Radashevsky 2005
Broad Temperature Range	None	Cold temperate-Tropical
Broad Salinity Range	None	Oligohaline-Hyperhaline

General Impacts

Economic Impacts

On the East of North America, Polydora cornuta is widely regarded as a pest in Easter Oyster (Crassostrea gigas) culture. It can develop dense colonies on the outside of the shells of oysters and mussels, smothering them with mud from their tubes. However, it is not capable of boring into shells, and is frequently confused with the specialized shell-borers P. ciliata and P. websteri (Galtsoff 1964; White and Wilson 1996; Sato-Okoshi 2000; Surugiru 2012). The extent to which P. cornuta affects oyster culture elsewhere in the world is unknown. However, P. cornuta was found consuming clam and oyster larvae in a hatchery in Newport, Oregon (Breese and Phibbs 1972).

Ecological Impacts

Polydora cornuta is widely recognized as a 'pollution indicator' species, known to rapidly increase in eutrophic estuaries, and estuaries receiving toxic pollutants, both in its native range and in areas where it has been introduced. This worm's tolerance of environmental extremes of temperature, salinity, and oxygen, as well as its feeding and habitat flexibility have enabled it to colonize harbors where other species have declined due to natural or anthropogenic stresses (Wolff 1973; Anger 1977; Rice and Simon 1980; Oviatt et al. 1987; Tagliapietra et al. 1998). Polydora cornuta occurs both in soft-bottom communities and fouling communities on marinas, docks, shells, ship hulls, etc. (Graham and Gay 1945; Wolff 1973; Bagaveeva and Zvyagintsev 2000), and can alternate between deposit-feeding from the sediment surface, or suspension-feeding on phytoplankton and particles in the water column (Dauer et al. 2005; Hentshel and Herrick 2005). Consequently, P. cornuta is a strong competitor in eutrophic habitats, and can become overwhelmingly dominant in sewage-polluted habitats (Anger 1977; Oviatt et al. 1987).

Competition- Competitive impacts of invading populations of Polydora cornuta are difficult to evaluate, because this polychaete tends to dominate in strongly disturbed and eutrophic environments, in which many polychaetes do poorly. However, its rapid growth, high fecundity, hypoxic tolerance, and trophic flexibility enable it to outnumber most other polychaetes (Wolff 1973; Anger 1977; Oviatt et al. 1987). Evidence of P. cornuta's competitive impacts is based partly on rapid colonization of polluted areas in the Black Sea, Sea of Marmara, Aegean Sea, and the Seto Inland Sea, Japan (Yamada et al. 2001; Losovskaya and Zolotarev 2003; Dagli and Ergen 2008; Karhan et al. 2008; Cinar et al. 2012), but also on eutrophication experiments, where it can reach extraordinary densities (e.g., Oviatt et al. 1987).

Herbivory- Polydora cornuta, reaches high abundances and becomes the dominant suspension-feeder in eutrophic areas, both as adults, with their palps extended in the water column (Dauer et al. 2005; Hentschel and Herrick 2005; Shimeta et al. 2005), and as planktonic larvae (Watras et al. 1985; Hanson 1999; Yamada et al. 2001). However, we have not found estimates of its grazing rates.

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Regional Impacts

NEP-IV	Puget Sound to Northern California		Economic Impact	Fisheries
IIn a Newport, Oregon shellfish hatchery, in 1970, Polydora cornuta was found to be feeding on larvae of Pacific Oysters (Crassostrea gigas and 'Manila Clams' (Venerupis philippinarum), and causing significant losses (Breese and Phipps 1972).

References

Schmitt, Joseph D.; Peoples, Brandon K.; Castello, Leandro; Orr, Donald J. (2019) Feeding ecology of generalist consumers: a case study of invasive blue catfish Ictalurus furcatus in Chesapeake Bay, Virginia, USA, Environmental Biology of Fishes 192: 443–465

Alalykina, Inna (2013) Preliminary data on the composition and distribution of polychaetes in the deep-water areas of the north-western part of the Sea of Japan, Deep-Sea Research II 86-87: 164-171

Anger, K.; Anger, V.; Hagmeir, E. (1986) Laboratory studies on larval growth of Polydora ligni, Polydora ciliata, and Pygospio elegans (Polychaeta, Spionidae), Helgolander Meeresuntersuchungen 40: 377-395

Anger, Klaus (1977) Benthic invertebrates as indicators of organic pollution in the Western Baltic Sea, Internationale Revue der Gesamten Hydrobiologie 62(2): 245-254

Bagaveeva, E. V.; Zvyagintsev, A. Yu. (2000) Introduction of polychaetes, Hydroides elegans (Haswell), Polydora limicola Annenkova, and Pseudopotamilla occelata Moore to the Northwestern part of the East Sea, Ocean Research 22(1): 25-36

Bancila. Raluca I.; Skolka, Marius; Ivanova, Petya; Surugiu, Victor; Stefanova, Kremena; Todorova. Valentina; Zenetos, Argyro (2022) Alien species of the Romanian and Bulgarian Black Sea coast: state of knowledge, uncertainties, and needs for future research, Aquatic Invasions 17: Published online

Bertasi, F. (2016) The occurrence of the alien species Polydora cornuta Bosc, 1802 (Polychaeta: Spionidae) in North Adriatic lagoons: an overlooked presence, Italian Journal of Zoology Published online: <missing location>

Blake, James A. (1969) Reproduction and larval development of Polydora from northern New England, Ophelia 7: 1-63

Blake, James A. (1971) A revision of the genus Polydora from the East Coast of North America (Polychaeta: Spionidae), Smithsonian Contributions to Zoology 75: 1-32

Blake, James A. (1983) Polychaetes of the family Spionidae from South America, Antarctica, and adjacent seas and islands, Biology of the Antarctic Seas 39(3): 205-288

Blake, James A.; Kudenov, Jerry D. (1978) Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera, Memoirs of the National Museum in Victoria 39: 171-280

Blake, James A.; Maciolek, Nancy J. (1987) A redescription of Polydora cornuta Bosc (Polychaeta: Spionidae) and desgnation of a neotype., Bulletin of the Biological Society of Washington 7: 11-15

Blake, James A.; Ruff, R. Eugene (2007) The Light and Smith Manual: Intertidal invertebrates from Central California to Oregon (4th edition), University of California, Berkeley CA. Pp. 309-410

Bloch. Philip; Novak, Grant (2011) The 2011 International Conference on Ecology & Transportation, International Conference on Ecology & Transportation, <missing place>. Pp. 400-410

Bonar, Scott A.; Bolding, Bruce D.; Divens, Marc AND Meyer, William (2005) Effects of introduced fishes on wild juvenile Coho Salmon in three shallow Pacific Northwest Lakes, Transactions of the American Fisheries Society 134: 641-652
DOI: 10.1577/T04-154.1

Boyd, Milton J.; Mulligan, Tim J; Shaughnessy, Frank J. (2002) <missing title>, California Department of Fish and Game, Sacramento. Pp. 1-118

Breese, Wilbur P.; Phibbs, F. Duane (1972) Ingestion of bivalve molluscan larvvae by the polychete annelid Polydora ligni, Veliger 14(3): 374-375

California Academy of Sciences 2005-2015 Invertebrate Zoology Collection Database. <missing URL>



Carlton, James T. (1979) History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific Coast of North America., Ph.D. dissertation, University of California, Davis. Pp. 1-904

Carlton, James T. (1989) Man's role in changing the face of the ocean: biological invasions and implications for conservation of near-shore environments, Conservation Biology 3(3): 265-273

Carson, Henry S.; Hentschel, Brian T. (2006) Estimating the dispersal potential of polychaete species in the Southern California Bight: implications for designing marine reserves, Marine Ecology Progress Series 316: 105-113

Carvalho, Susana and 5 authors (2011) Factors structuring temporal and spatial dynamics of macrobenthic communities in a eutrophic coastal lagoon (Óbidos lagoon, Portugal), Marine Environmental Research 71: 97-110

Chapman, John W.; Therriault, Thomas; Harris, Leslie; Breitenstein, Ralph (2011) The 2010 PICES Rapid Assessment Survey of shallow water nonindigenous, native and cryptogenic marine species of central Oregon, Oregon State University, Hatfield Marine Center, Newport OR. Pp. 48

Çinar, Melih Ertan; Balkis, Husamttin; Albayrak, Sertan; Dagli, Ertan; Karhan, Unsal Sellahattin (2009) Distribution of polychaete species (Annelida: Polychaeta) on the polluted soft substrate of the Golden Horn estuary, with special emphasis on the alien species, Cahiers de Biologie Marine 50: 1-17

Çinar, Melih Ertan; Dagli, Ertan; Açik, Sermin (2011) Annelids (Polychaeta and Oligochaeta) from the Sea of Marmara, with descriptions of five new species, Journal of Natural History 45(33-34): 2105-2143

Cinar, Melih Ertan; Ergen, Zeki; Dagli, Ertan; Petersen, Mary E. (2005) Alien species of spionid polychaetes (Streblospio gynobranchiata and Polydora cornuta) in Izmir Bay, eastern Mediterranean, Journal of the Marine Biological Association of the United Kingdom 85: 821-827

Cohen, Andrew N. and 22 authors (2001) <missing title>, Washington State Department of Natural Resources, Olympia. Pp. <missing location>

Cohen, Andrew N.; Carlton, James T. (1995) Nonindigenous aquatic species in a United States estuary: a case study of the biological invasions of the San Francisco Bay and Delta, U.S. Fish and Wildlife Service and National Sea Grant College Program (Connecticut Sea Grant), Washington DC, Silver Spring MD.. Pp. <missing location>

Cordell, Jeffrey R. and 5 authors (2009) Factors influencing densities of non-indigenous species in the ballast water of ships arriving at ports in Puget Sound, Washington, United States, Aquatic Conservation: Marine and Freshwater Ecosystems 19(3): 322-343

Cory, Robert L. (1967) Epifauna of the Patuxent River Estuary, Chesapeake Science 8(2): 71-89

Dagli, Ertan; Ergen, Zeki (2008) First record of Polydora cornuta Bosc, 1802 (Polychaeta: Spionidae) from the Sea of Marmara, Turkey basin, Aquatic Invasions 3(2): 231-233

Dauer, Daniel M.; Mahon, Heidi K.; Sarda, Rafael (2003) Functional morphology and feeding behavior of Streblospio benedicti and S. shrubsolii (Polychaeta: Spionidae), Hydrobiologia 496: 207-213

de Montaudouin, Xavier; Sauriau, Pierre-Guy (2000) Contributions to a synopsis of marine species richness in the Pertuis-Charentais Sea with new insights into the soft-bottom macrofauna of the Marennes-Oleron Bay, Cahiers de Biologie Marine 41: 181-222

Dean, Harlan K. (1996) Subtidal benthic polychaetes (Annelida) of the Gulf of Nicoya, Costa Rica, Revista Biologia Tropical 44(Suppl. 3): 69-80

Faasse, Marco (2012) The exotic isopod Synidotea in the Netherlands and Europe, A Japanese or American invasion (Pancrustacea: Isopoda)?, Nederlandse Faunistiche Mededelingen 108: 103-106

Fairey, Russell; Dunn, Roslyn; Sigala, Marco; Oliver, John (2002) Introduced aquatic species in California's coastal waters: Final Report, California Department of Fish and Game, Sacramento. Pp. <missing location>

Galtsoff, Paul S. (1964) The American oyster, Crassostrea virginica Gmelin. Chapter 18: Environmental factors affecting oyster populations, Fisheries Bulletin 64: 397-456

Gartner, Heidi N.; Murray, Cathryn Clarke; Frey, Melissa A.; Nelson, Jocelyn C.; Larson, Kristen J.; Ruiz, Gregory M.; Therriault, Thomas W. (2016) Non-indigenous invertebrate species in the marine fouling communities of British Columbia, Canada, BioInvasions Records <missing volume>: <missing location>

Gibson, Glenys; Hart, Corban; Coulter, Claire; Xu, Haixin (2012) Nurse eggs form through an active process of apoptosis in the spionid Polydora cornuta (Annelida), Integrative and Comparative Biology 52: 151-160

Graham, Herbert W.; Gay, Helen (1945) Seasonal attachment and growth of sedentary marine organisms at Oakland, California., Ecology 26(4): 375-386

Hansen, Benni Winding (1999) Cohort growth of planktotrophic polychaete larvae -are they food limited?, Marine Ecology Progress Series 178: 109-119

Hartman, Olga (1941) Some contributions to the biology and life history of Spionidae from California, Allen Hancock Pacific Expeditions 7: 289-322

Hartman, Olga (1969) <missing title>, Allan Hancock Foundation, Los Angeles. Pp. <missing location>

Hentschel, Brian T. (1998) Intraspecific variations in 13C indicate ontogenetic diet changes in deposit-feeding polychaetes, Ecology 79(4): 1357-1370

Hentschel, Brian T.; Herrick, Brenda S. (2005) Growth rates of interface-feeding spionid polychaetes in simulated tidal currents, Journal of Marine Research 63: 983-999

Hernandez-Alcantera, P.; Gonzalez-Ortiz, L.; Solis-Weiss, V. (1994) [Spionids of the Gulf of California and the Gulf of Tehuantepec, Mexico], Revista Biologia Tropical 42(3): 467-477

Hutchings, P. A.; Turvey, S. P. (1978) The Spionidae of South Australia (Annelida: Polychaeta), Transactions of the Royal Society of South Australia 108(1): 1-20

Inglis, Graeme and 6 authors. (2005b) Whangarei Marina- Baseline survey for non-indigenous marine species, Biosecurity New Zealand Technical Paper 2005/15: 1-68

Jones, Kim Koch (1983) <missing title>, Oregon State University, Corvallis OR. Pp. 1-67

Karhan, S. Ü.; Kalkan, E.; N. Simboura; Mutlu, E.; Bekbölet, M. (2008) On the occurrence and established populations of the alien polychaete Polydora cornuta Bosc, 1802 (Polychaeta: Spionidae) in the Sea of Marmara and the Bosporus Strait (Turkey), Mediterranean Marine Science 9(1): 5-19

Kudenov, Jerry D. (1975) Sedentary polychaetes from the Gulf of California, Mexico, Journal of Natural History 9: 205-231

Levin, Lisa A. (1984b) Life history and dispersal patterns in a dense infaunal polychaete assemblage: Community structure and response to disturbance, Ecology 65(4): 1185-1200

Levin, Lisa A.; Talley, Theresa Sinicrope; Hewitt, Judi (1998) Macrobenthos of Spartina foliosa (Pacific cordgrass) salt marshes in southern California: Community structure and comparison to a Pacific mudflat and a Spartina alterniflora (Atlantic smooth cordgrass) marsh., Estuaries 21: 1

Light, William J. (1977) Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: a revised list with nomenclatural changes, new records and comments on related species from the Northeastern Pacific Ocean., Proceedings of the Biological Society of Washington 90(1): 66-88

Light, William J. (1978) <missing title>, Boswood Press, Pacific Grove CA. Pp. <missing location>

Lippson, Alice Jane; Lippson, Robert L. (1997) <missing title>, Johns Hopkins University Press, Baltimore. Pp. <missing location>

Liu, Wenliang; Liang, Xiaoli ; Zhu, Xiaojing (2015) A new record and mitochondrial identification of Synidotea laticauda Benedict, 1897 (Crustacea: Isopoda: Valvifera: Idoteidae) from the Yangtze Estuary, China, Zootaxa 4294: 371-380

Losovskaya, G. V.; Zolotarev, V. N. (2003) The polychaete Polydora limicola in benthic communities of the Black Sea, Russian Journal of Marine Biology 29(4): 248-250

Lu, L.; Levings, C. D.; Piercey, G. E. (2007) Preliminary investigation on aquatic invasive species of marine and estuarine macroinvertebrates on floating structures in five British Columbia harbours, Canadian Manuscript Report of Fisheries and Aquatic Sciences 2814: 2-30

Lunz, G. Robert, Jr. (1941) Polydora, a pest in South Carolina oysters, Journal of the Elisha Mitchell Scientific Society 57(2): 273-283

Lutz-Collins, Vanessa; Quijon, Pedro A. (2014) Animal-sediment relationships in an Atlantic Canada marine protected area: Richness, composition and abundance in relation to sediment food indicators, Marine Biology Research 10(6): 577-588

Michaelis, H. (1978) [On the morphology and ecology of Polydora ciliata and P. ligni (Polychaea, Spionidae)], Helgolander Wissenschaftliche Meeresuntersuchungen 31: 102-116

Miloslavich, Patricia and 16 authors (2010) Marine biodiversity in the Caribbean: Regional estimates and distribution patterns, PLOS ONE 5(8): e11916

Murray, Cathryn Clarke, and 5 authors (2014) Spatial distribution of marine invasive species: environmental, demographic and vector drivers, Diversity and Distributions 20: 824-836

Neira, Carlos; Levin, Lisa A.; Grosholz, Edwin D. (2005) Benthic macrofaunal communities of three sites in San Francisco Bay invaded by hybrid Spartina, with comparison to uninvaded habitats., Marine Ecology Progress Series 292: 111-126

Neves, Carolina Somao; Rocha, Rosana Moreira; Pitombo, Fabio Bettini; Roper, James J. (2007) Use of artificial substrata by introduced and cryptogenic marine species in Paranagua Bay, southern Brazil, Biofouling 23(5): 319-330

Nichols, Frederic H.; Thompson, Janet K. (1985a) Time scales of change in the San Francisco Bay benthos., Hydrobiologia 129: 121-138

Orensanz, Jose Maria and 14 other authors (2002) No longer the pristine confines of the world ocean: a survey of exotic marine species in the southwestern Atlantic, Biological Invasions 4(1-2): 115-143

Orth, Robert J. (1971) Observations on the planktonic larvae of Polydora ligni Webster (Polychaeta: Spionidae) in the York River, Virginia, Chesapeake Science 12(3): 121-124

Oviatt, Candace A. et al. (1987) Fate and effects of sewage sludge in the coastal marine environment: a mesocosm experiment, Marine Ecology Progress Series 41: 187-203

Qian, Pei-Yuan; Chia, Fu-Shiang (1993) Larval development as influenced by food limitation in two polychaetes: Capitella sp. and Polydora ligni Webster, Journal of Experimental Marine Biology and Ecology 133: 93-105

Radashevsky, V. I.; Selifonova, Z. P. (2013) Records of Polydora cornuta and Streblospio gynobranchiata (Annelida, Spionidae) from the Black Sea, Mediterranean Marine Science 14(2): 261-269

Radashevsky, Vasily I. (2005) On adult and larval morphology of Polydora cornuta Bosc, 1802 (Annelida: Spionidae), Zootaxa 1064: 1-24

Radashevsky, Vasily I. (2012b) Spionidae (Annelida) from shallow waters around the British Islands: an identification guide for the NMBAQC Scheme with an overview of spionid morphology and biology, Zootaxa 3152: 1-35

Radashevsky, Vasily I.; Hsieh, Hwey-Lian (2000) Polydora (Polychaeta: Spionidae) species from Taiwan, Zoological Studies 39(3): 203-217

Ramberg, Jens Petter; Schram, Thomas (1982) A systematic review of the Oslofjord species of Polydora Bosc and Pseudopolydora Czerniavsky, with some new biological and ecological data (Polychaeta: Spionidae), Sarsia 68: 233-247

Ranasinghe, J. Ananda and 6 authors. (2005) The prevalence of non-indigenous species in southern California embayments and their effects on benthic macroinvertebrate communities, Biological Invasions 7: 679-686

Read, G. B. (1975) Systematics and biology of Polydoroid species (Polychaeta: Spionidae) from Wellington Harbour, Journal of the Royal Society of New Zealand 5: 4

Read, Geoffrey B.; Gordon, Dennis P. (1991) Adventive occurrence of the fouling serpulid Ficopomatus enigmatus (Polychaeta) in New Zealand, New Zealand Journal of Marine and Freshwater Research 25(3): 269-274

Reish, Donald J.; Kauwling, Thomas J.; Schreiber, Traynor C. (1975) Annotated checklist of the marine invertebrates of Anaheim Bay, California Department of Fish and Game Fish Bulletin 165: 41-51

Reish, Donald J.; Barnard, J. Laurens (1967) The benthic Polychaeta and Amphipoda of Morro Bay, California, Proceedings of the United States National Museum 120(3565): 1-26

Rice, Stanley A. (1981) Spermatogenesis and sperm ultrastructure in three species of Polydora and in Streblospio benedicti (Polychaeta: Spionidae), Zoomorphology 97: 1-16

Rice, Stanley A.; Karl, Stephen; Rice, Katherine A. (2008) The Polydora cornuta complex (Annelida: Polychaeta) contains populations that are reproductively isolated and genetically distinct, Invertebrate Biology 127(1): 45-64

Rice, Stanley A.; Simon, Joseph L. (1980) <missing title>, 19 <missing publisher>, <missing place>. Pp. 79-115

Riggs, Sharon R. (2011) <missing title>, Padilla Bay NERR, Padilla Bay WA. Pp. 5

Rodrigues, Ana Maria; Quintino, Victor; Sampaio, Leandro; Freitas, Rosa; Neves, Ramiro (2011) Benthic biodiversity patterns in Ria de Aveiro, Western Portugal: Environmental-biological relationships, Estuarine, Coastal and Shelf Science 95: 338-348

Ruiz, Gregory M.; Geller, Jonathan (2018) Spatial and temporal analysis of marine invasions in California, Part II: Humboldt Bay, Marina del Re, Port Hueneme, and San Francisco Bay, Smithsonian Environmental Research Center & Moss Landing Laboratories, Edgewater MD, Moss Landing CA. Pp. <missing location>

Ruiz, Gregory; Geller, Jonathan (2021) Spatial and temporal analysis of marine invasions: supplemental studies to evaluate detection through quantitative and molecular methodologies, Marine Invasive Species Program, California Department of Fish and Wildlife, Sacramento CA. Pp. 153 ppl.

Sahin, Güley Kurt; Çinar, Melih Ertan (2012) A check-list of polychaete species (Annelida: Polychaeta) from the Black Sea, Journal of the Black Sea/Mediterranean Environment 18(1): 10-48

Sato-Okoshi, Waka (2000) Polydorid species (Polychaeta: Spionidae) in Japan, with descriptions of morphology, ecology and burrow structure. 2. Non-boring species, Journal of the Marine Biological Association of the United Kingdom 80: 443-456

Sato-Okoshi, Waka; Okoshi, Kenji (1997) Survey of the genera Polydora, Boccardiella and Boccardia (Polychaeta, Spionidae) in Barkley Sound (Vancouver Island, Canada), with special reference to boring activity, Bulletin of Marine Science 60(2): 482-493

Sebesvári, Zita (2007) <missing title>, University of Oldenberg, <missing place>. Pp. 1-106

Sebesvari, Zita; Esser, Ferdinand; Harder, Tilmann (2006) Sediment-associated cues for larval settlement of the infaunal spionid polychaetes Polydora cornuta and Streblospio benedicti, Journal of Experimental Marine Biology and Ecology 337: 109-120

Selifonova, Zh. P. (2015) Marine biological invasions in the liman 'Zmeinoe Ozero' (Snake Lake) ecosystem, the north-eastern Black Sea, Journal of the Marine Biological Association of the United Kingdom 95(3): 453-459

Simboura, N.; Sigala, K.; Voutsinas, E.; Kalkan, E. (2008) First occurrence of the invasive alien species Polydora cornuta Bosc, 1802 (Polychaeta: Spionidae) on the coast of Greece (Elefsis Bay; Aegean Sea), Mediterranean Marine Science 9(2): 119-124

Simmons, Ernest G. (1957) An ecological survey of the Upper Laguna Madre of Texas, Publications in Marine Science of the University of Texas 4(2): 156-200

Smith, L. David; Wonham, Marjorie J.; McCann, Linda D.; Ruiz, Gregory M.; Hines, Anson H.; Carlton, James T. (1999) Invasion pressure to a ballast-flooded estuary and an assessment of inoculant survival, Biological Invasions 1: 67-87

Son, Mikhail O. ; Bondarenko, Olena; Kvach, Yuriy (2023) New data on aquatic alien invertebrates in the Ukrainian Danube Delta with special emphasis on the first records of the trumpet ram’s-horn, Menetus dilatatus (Gould, 1841), BioInvasions Records 12(3): 787–795
https://doi.org/10. 3391/bir.2023.12.3.14

Surugiu, Victor (2008) Zoogeographical origin of the polychaete fauna of the Black and Azov Seas, Cahiers de Biologie Marine 49: 351-354

Surugiu, Victor (2009) The Influence of sewage pollution on polychaetes associated with mussel beds of the southern Romanian Black Sea coast, Geo-Eco-Marina 15: 77-87

Surugiu, Victor (2012) Systematics and ecology of species of the Polydora -complex (Polychaeta: Spionidae) of the Black Sea, Zootaxa 3518: 45-65

Tagliapietra, D.; Pavan, M.; Wagner, C. (1998) Macrobenthic community changes related to eutrophication in Palude della Rosa (Venetian Lagoon, Italy), Estuarine, Coastal and Shelf Science 47: 217-226

Takata, Norimi (2011) Physiological characteristics of Polydora cornuta inhabiting the eutrophic port of Fukuyama, Eco-Engineering 23(2): 41-46

Talley, Theresa Sinicrope; Dayton, Paul K.; Ibarra-Obando, Silvia E. (2000) Tidal flat macrofaunal communities and their associated environments in estuaries of southern California and northern Baja California, Mexico, Estuaries 23(1): 97-114

U.S. National Museum of Natural History 2002-2021 Invertebrate Zoology Collections Database. http://collections.nmnh.si.edu/search/iz/



Walker, Lexie Margaret (2009) Polydora and Dipolydora (Polychaeta: Spionidae) of estuaries and bays of subtropical eastern Australia: A review and morphometric investigation of their taxonomy and distribution, Southern Cross University, Lismore, New South Wales, Australia. Pp. 1-179

Wasson, Kerstin; Zabin, C. J.; Bedinger, L.; Diaz, M. C.; Pearse J. S. (2001) Biological invasions of estuaries without international shipping: the importance of intraregional transport, Biological Conservation 102: 143-153

Watras, Carl J.; Garcon, Veronique; Olson, Robert J.; Chisolm. Sallie W; Anderson, Donald M. (1985) The effect of zooplankton grazing on estuarine blooms of the toxic dinoflagellate Gonyaulax tamarensis, Journal of Plankton Research 7(6): 891-908

Weinberg, James R. (1984) Interactions between functional groups in soft substrata: Do species differences matter?, Journal of Experimental Marine Biology and Ecology 80(1): 11-28

White, Marie E.; Wilson, Elizabeth A. (1996) The Eastern Oyster: Crassostrea virginica, Maryland Sea Grant, College Park. Pp. 559-579

Wolff, W. J. (1973) The estuary as a habitat: An analysis of data on the soft-bottom macrofauna of the estuarine area of the rivers Rhine, Meuse, and Scheldt, Zoologische Verhandelingen 126: 4-242

Wong, Eunice; Kupriyanova, Elena K.; Hutchings, Pat; Capa, María; Radashevsky, Vasily; ten Hove, Harry A. (2014) A graphically illustrated glossary of polychaete terminology: invasive species of Sabellidae, Serpulidae and Spionidae, Memoirs of Museum of Victoria 71: 327-342

Wonham, Marjorie J.; Carlton, James T. (2005) Trends in marine biological invasions at local and regional scales: the Northeast Pacific Ocean as a model system, Biological Invasions 7: 369-392

Yamada, Hiroshi; Imabayashi, Hiromichi; Takat, Norimi (2001) Effect of oxygen-deficient water on the settlement and survival of spionid polychaete Polydora cornuta on the organically enriched bottom in Fukuyama Harbor, Japan, Nippon Suisan Gakkaishi 67(5): 814-820

Zajac, R. N. (1995) Sublethal predation on Polydora cornuta (Polychaeta: Spionidae): patterns of tissue loss in a field population, predator functional response and potential demographic impacts, Marine Biology 123(3): 531-541

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