Invasion History

First Non-native North American Tidal Record: 2004
First Non-native West Coast Tidal Record: 2004
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Polydora hoplura was described from the Gulf of Naples, Italy in 1868, and was regarded as native from southern Britain to the Mediterranean (de Montaudouin,and Sauriau 2000; Dauvin et al. 2003). However, surveys of this worm's genetics and history of its spread suggests that it is native to the Northwest Pacific,,and Japan in particular, and spread by introductions in ship fouling, and with transfers of Pacific Oysters (Magallana = Crassostrea gigas).  Its occurrence in Japan may have been poorly known due to limited studies of shell-boring polychaetes before the 1990s  (Radashevsky et al. 2023). . Polydora hoplura is one of many spionids which burrow into the calcareous shells of mollusks, barnacles and other  organisms, and so can be transported in hull fouling. Reports of its range in Europe are somewhat spotty, possibly due to confusion with a non-boring species, P. ciliata.(Radashevsky et al. 2023). It is known from Belgium (first record 1962, establishment uncertain, Kerckhof et al. 2007); southwest England (Wilson 1928); Atlantic France (Dauvin et al. 2003), the Bay of Biscay (Galicia, Spain; Parapar et al. 2009); the Bay of Naples (Radashevsky and Migotto 2016); the Adriatic Sea (Croatia; Labura and Hrs-Brenko 1990), and the Aegean Sea (GBIF 2017). In Europe, the most common host is Ostrea edulis, the European Oyster (Wilson 1928; Labura and Hrs-Brenko 1990; Radashevsky and Migotto 2016).. However, it is often associated with Pacific Oyster (M. gigas) (Sato-Okoshi et al. 2023).

This polychaete is now widely distributed around the world, especially in culture operations for oysters and abalones, where it can damage shells, thereby affecting the condition and quality of these shellfish. Exotic occurrences are known from: the Canary Islands (Bilbao et al. 2011); South Africa (Simon 2009; Simon 2011); Australia (Blake and Kudenov 1978); New Zealand (Handley and Bergquist 1997); Chile (Moreno et al. 2006); Brazil; Japan (Sato-Okoshi et al. 1998; Sato-Okoshi et al. 2016); and California (Radashevsky and Migotto 2016).

North American Invasion History:

Invasion History on the West Coast:

In May 2004, Polydora hoplura was collected in a marina in San Diego Bay (Sato-Okoshi et al. 2023).  In 2011 it was collected at multiple sites in central and southern California: Monterey Bay, at Breakwater Cove Marina and in Dana Point Harbor, Orange County. The collections were made in 1–2 m depth, within dock fouling. The Monterey fouling community contained oysters, mussels, bryozoans, and sponges, but collection notes from Dana Point mentioned oysters and bryoliths (Radashevsky and Migotto 2016; Radashevsky et al. 2023). This polychaete is established along the coast of centrl and southern California,

Invasion History Elsewhere in the World:

Polydora hoplura was first reported from Table Bay Docks, Cape Town, South Africa, on the Atlantic side of the Cape region in 1947 (Mead et al. 2011). By 1967, this worm reached Plettenberg Bay, on the Indian Ocean side of the Cape (Day 1967, cited by Mead et al. 2011). It is now widely distributed along the southern tip of the continent, from Paternoster to Haga Haga, southwest of Durban, in oyster (Ostrea edulis, Crassostrea gigas) and abalone (Haliotus midas, H. spadaciae) (David et al. 2016; Williams et al. 2016). In 1999, it was discovered in a closed culture operation on Gran Canaria Island, infesting the native abalone Haliotis tuberculata coccinea (Bilbao et al. 2011). Wild populations are patchily distributed in the Canary Islands (Pascual and Nunez 1999, cited by Bilbao et al. 2011) and we consider it cryptogenic there.

Polydora hoplura was collected at many sites in southeastern Australia and New Zealand in the 1970s. It was found in Tiparra Bay, Gulf of St. Vincent, South Australia in 1971 (Atlas of Living Australia 2017), Port Philip Bay in 1975 (Blake and Kudenov 1978), and Simmonds Bay, on the south coast of Tasmania in 1977 (Blake and Kudenov 1978). Later collections were in northern New South Wales in 1992 (Walker 2009, Atlas of Living Australia 2017), and Albany, Western Australia in 2005 (Radashevsky and Migotto 2016). In New Zealand, P. hoplura was first found in Evans Bay, Wellington on the South Island in 1972 (Read 1975). This polychaete has been found in many sites on the North and South Islands (Read 1975; Handley 1995; Handley and Bergquist 1997; Inglis et al. 2006 a,b). It has been associated with the native pen-shell (Atrina pectinata zelandica), a native abalone (Haliotis iris), a native scallop (Pecten novaezelandiae), a native oyster (Ostrea lutaria), and the introduced Pacific Oyster (Crassostrea gigas) (Read 1975; Blake and Kudenov 1978; Handley and Bergquist 1997).

In 1997, a boring polychaete was found in Yamada Bay (Iwate Prefecture), Honshu, and Shironohana (Kochi Prefecture), Japan, in the shells of cultured native Pacific Oysters (Crassostrea gigas), and a native top shell (Omphalius rusticus). Sato-Okoshi (1998) described the worm as a new species, P. uncinata. Later, the Japanese specimens were found to be conspecific with P. hoplura from Europe, South Africa, Australia, and New Zealand (Radashevsky and Migotto 2016; Sato-Okoshi et al. 2017).

In 2002, Polydora hoplura, identified then as P. uncinata, was found infesting the shells of cultured Japanese abalones (Haliotis discus hannai), in land-based culture tanks in Coquimbo, Chile (Radashevsky and Olivares 2005, Moreno et al. 2006, Radashevsky and Migotto 2016). We have not found reports of this worm in open waters of Chile. However, P. hoplura has been collected from oysters (Crassostrea rhizophora) from the Atlantic side of South America, near Sao Paulo Brazil in 2015 (Radashevsky and Migotto 2016).

Polydora hoplura has a great potential for transport by humans, in the shells of transported oysters and abalones, on barnacles and bivalves fouling the hulls of ships (Radashevsky and Migotto 2016), or in ballast water. This polychaete is one of a number of invertebrates, displaying poecilogony, meaning that it is capable of switching between lecithotropic brooded development (spending about four days in the plankton at 21ºC), or planktotrophic larvae, spending ~30 days in the plankton (David et al. 2014). Planktotrophic larvae could be transported long distances in ballast water, as long as the water contained sufficient phytoplankton for sustenance.


Polydora hoplura is a shell-boring spionid polychaete, burrowing in molluscan and barnacle shells. Adults of Polydora hoplura have up to 120–160 chaeta-bearing segments (chaetigers). The prostomium has a weak anterior incision, forming two lobes and is prolonged back to chaetiger 3 as a low caruncle. A short occipital tentacle is located on the caruncle. Specimens can have up to four eyes, but can also lack them altogether. When two pairs are present, the anterior pair is further apart than the posterior. A ciliated groove runs on either side of the caruncle. The palps are long and slim and their length is the same as the total length of the first 25th chaetigers of the worm.

Chaetiger 1 has well-formed parapodial lobes, but has short capillary chaetae only in neuropodia. The parapodial lobes are well developed anteriorly with the exception of chaetiger 5. Chaetiger 5 is enlarged, lacks parapodial lobes, and has up to four dorsal superior winged capillary chaetae, and six heavy falcate spines with lateral flanges, alternating with companion bilimbate (tipped with two wings) chaetae, and six ventral winged capillaries. Chaetigers from 7 up to 10 have hooded bidentate (double-toothed) hooks in the neuropodia, not accompanied by capillary setae. The shafts of the hooks are constricted in the middle. The notopodia in 10–15 posterior-most chaetigers each have 1–2 heavy recurved spines and a tuft of slender capillaries. Branchiae begin around chaetiger 7 and continue rearward, but diminish as the posterior spines begin. The pygidium is flared and resembles a suction cup, with a dorsal gap.

Adult worms reach a length of 25–40 mm and are pale yellow in color. Black bands are present on the pair of palps. There is some dusky pigment on the peristomium. The branchiae and the dorsal blood vessel are red. The pygidium is white. Larval morphology is described by Wilson (1928) and Radashevsky and Migotto (2016). The above description is based on Read 1975; Blake and Kudenov 1978; Walker 2009; Radashwvsky and Migotto 2016.


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Palpata
Order:   Canalipalpata
Suborder:   Spionida
Family:   Spionidae
Genus:   Polydora
Species:   hoplura


Polydora hoplura (Claparède, 1868)
Polydora hoplura hoplura (Day, 1967)
Polydora uncinata (Sato-Okoshi, 1998)
Leucodora sanguinea (GiRD, 1881)

Potentially Misidentified Species

Polydora ciliata
Polydora ciliata is a shell-boring polychaete described from the Northeast Atlantic, and native from Northern Europe to the Mediterranean. Records from the East Coast of North America probably refer to P. websteri (Blake 1971). Many records of P. ciliata from other parts of the world probably refer to a related species, and the occurrence of this species in Australia is uncertain (Walker 2009).

Polydora cornuta

Polydora cornuta is a tube-building estuarine polychaete, described from the East Coast of North America, and widely introduced (Blake 1971). It is not a shell-borer, but is very abundant in oyster and mussel beds. It has thin companion setae on chaetiger 5th, flattened, feathery, closely adhering to major spines.

Polydora websteri complex
Polydora websteri is a species complex of shell-boring polychaetes, originally described from Connecticut (Blake 1971), and later reported from both coasts of North America, the West Coast of South America, Australia, and New Zealand (Walker 2009). We consider the West Coast and other outlying populations to be cryptogenic, possibly introduced, and/or possibly cryptic species (Walker 2009). It has no occipital tentacle and no heavy recurved spines on posterior notopodia.



Polydora hoplura has two separate sexes. Eggs are laid in capsules, linked in strings attached to the interior of the burrows. In the Yealm estuary, England there were about 50 eggs per string (Wilson 1928). In South Africa, two modes of reproduction were observed: (1) a short brooded development, with release of large numbers of larvae (mean = 1544.3) at the 3-chaetiger stage followed by a comparatively long planktotrophic development (~13–40 days at 12 to 28 °C); and (2) a long brooded lecithotrophic development, with a small (mean = 19.9) brood of larvae feeding on yolky nurse eggs (adelphophagy), released at 16–18 chaetigers and spending about four to seven days in the plankton (David et al. 2014; David and Simon 2015). The two reproductive forms are conspecific, with shared haplotypes, so this appears to be a case of poecilogony or intraspecific variation in larval developmental mode, which is known in several other spionids (Levin 1984a; David et al. 2014). Larvae of both types settle at about 1 mm length (David et al. 2014).

Polydora hoplura is known from cold-temperate Southwest England and Brittany, to subtropical climates (Sao Paulo, Brazil and Canary Islands) (Bilbao et al. 2011; Radashevsky and Migotto 2016). Its salinity range is unknown, but it appears to be absent from low salinity waters. This worm is associated with a wide range of calcareous habitats, most frequently mollusk shells, but also barnacles, coralline algae, and sponges (Bilbao et al. 2011; David and Simon 2014; Radashevsky and Migotto 2016). It was found on pen shells (Atrina pectinata zelandica) protruding from soft subtidal sediment in New Zealand (Read 1975), but is also found in rocky areas and pilings, on mussels, abalones, other mollusks, barnacles, and coralline algae (Radashevsky and Migotto 2016; David et al. 2016). Polydora hoplura is known to be especially abundant in cultivated oyster beds and in onshore culture tanks for oysters and abalones (Wilson 1928; Handley and Bergquist 1997; Bilbao et al. 2011; Simon 2015). At the same time, its occurrence on barnacles and mussels gives it the potential for transport in ship fouling (Walker 2009). Adults are suspension-feeders, and feed by extending beyond their burrows to capture phytoplankton and detritus on their long palps (Blake 1971).


Phytoplankton, detritus



Other shell-boring spionids

Trophic Status:

Suspension Feeder



General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatUnstructured BottomNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone

Life History

Tolerances and Life History Parameters

Minimum Reproductive Temperature12Lowest tested (David and Simon 2014, South Africa).
Maximum Reproductive Temperature28Highest tested (David and Simon 2014, South Africa).
Minimum Duration7

Polydora hoplura produces either lecithotrophic or planktotrophic larvae. At 28 °C, planktotrophic spend about 13 days in the plankton after release from the egg capsule, while lecithotrophic larvae take about 7 days (David and Simon 2014).

Maximum Duration40

Polydora hoplura produces either lecithotrophic or planktotrophic larvae. At 12 °C, planktotrophic spend about 40 days in the plankton after release from the egg capsule, while lecithotrophic larvae take about 7 days (David and Simon 2014).

Minimum Length (mm)25Radashvesky and Migotto 2015, Brazil
Maximum Length (mm)40Read 1975, New Zealand
Broad Temperature RangeNoneCold temperate-Subtropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

Polydora hoplura is a spionid polychaete which bores into calcareous substrates, usually the shells of mollusks, but also barnacles, coralline algae, sponges, and limestone (Wilson 1928; Read 1975; Simon 2015). In cultivated oyster beds and onshore abalone culture systems, P. hoplura heavily infests shells, producing frequent mud-blisters and riddling the shells with burrows, producing deformations, shell breakage, and lowering the commercial quality of the shellfish (Simon 1995; Handley and Bergquist 1997; Radashevsky and Olivares 2005; Bilbao et al. 2011). The species enters abalone shells through the leading edge and is also capable of burrowing directly through the shell (Lleonard et al. 2003). In open-water culture and natural ecosystems, impacts of P. hoplura are difficult to assess because of the presence of native and cryptogenic shell-boring spionids, including other worms of the P. websteri/ciliata complex (Read 1975; Simon 1995; Sato-Okoshi et al. 2023).

Economic Impacts

Fisheries- Impacts of P. hoplura seem to be concentrated in cultivated oyster beds and closed aquaculture systems, probably because of the high densities of mollusks. Studies found that this polychaete had little effect on condition or survival of the European Oyster (Ostrea edulis) in the Adriatic Sea, Croatia (Labura and Hrs-Brenko 1990), or on Pacific Ooysters (Crassostrea gigas) in New Zealand (Handley and Bergquist 1997). However, more serious infestations of Pacific Oysters by P. hoplura have been reported from South Africa and Tasmania (Lleonard et al. 2003a; David et al. 2015; Simon 2015). Polydora hoplura has had severe impacts on cultured abalones in closed culture, including Japanese abalones (Haliotis discus hannai) in Chile (Radashevsky and Olivares 2005), and in native abalones in the Canary Islands (H. tuberculata coccinea). In South Africa, this polychaete infests cultured native H. midas and H. spadaciae (Simon 2011).

Treatment of commercially reared Crassostrea gigas with fresh water or heated sea water seems to be a practical means to reduce Polydora infestation (Nel et al. 1996) and also air drying was found to be a promising spionid treatment in abalone culture (Lleonard et al. 2003b). The authors recommended about 2–4 h exposure time at greater than approximately 16 °C for treatment of relatively recent infestations, but they mentioned that has to be done prudently to avoid potential suppression of abalone growth. Other chemicals were investigated, but none seems to be the best treatment for both the host and the parasite (Simon et al. 2010).

Regional Impacts

WA-INoneEconomic ImpactFisheries
Polydora hoplura bored into the shells of cultured native abalones, Haliotis tuberculata coccinea, creating holes in the shell, chimneys on the shell-surface, and inducing the formation of blisters, causing breakage of some of shells and death of the abalones. The worm infestations were controlled by the drug Mebendazole, which did not affect the growth of the abalones (Bilbao et al. 2011).
WA-VNoneEconomic ImpactFisheries
Polydora hoplura bored into the shells of cultured Pacific Oysters, Crassostrea gigas, in Algoa Bay. The worms created holes in the shell, inducing the formation of mud blisters inside the shell, and lowering the quality of the oysters. A 12-hour exposure to fresh water or a 40 second exposure to 70ºC heat reduced the infestation after a 2-month recovery period (Nel et al. 1996).
WA-IVNoneEconomic ImpactFisheries
Polydora hoplura is considered to be a serious pest of oysters and to a lesser extent abalones, as far west as Saldanha Bay and Jacobsbaai (Simon et al. 2011; David et al. 2016).
NZ-IVNoneEconomic ImpactFisheries
Polydora hoplura bored into the shells of cultured Pacific Oysters, Crassostrea gigas, in Mahurangi Bay, North Island. The worms created holes in the shells, inducing the formation of mud blisters inside the shell, and lowering the quality of the oysters. A 12-hour exposure to fresh water or a 40 second exposure to 70ºC heat reduced the infestation after a 2-month recovery period (Handley and Bergquist 1997).
SEP-CNoneEconomic ImpactFisheries
Polydora hoplura infested Japanese abalones (Haliotis discus hannai) in a closed culture operation in land-based tanks in Coquimbo, Chile.
AUS-IXNoneEconomic ImpactFisheries
Cultured abalones, Haliotus spp., were infested by Polydora hoplura and native Boccardia knoxi, which caused mud blisters, causing shell damage, reduced weight, and mortality (Lleonart et al. 2003a).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NEP-V Northern California to Mid Channel Islands 2011 Def Estab
P080 Monterey Bay 2011 Def Estab
MED-III None 1868 Def Estab
SA-II None 2015 Def Estab
SEP-C None 2002 Def Estab
WA-I None 1999 Def Estab
WA-IV None 1947 Def Estab
WA-V None 1967 Def Estab
NEA-III None 0 Def Estab
NZ-IV None 1972 Def Estab
NEA-II None 1962 Def Unk
NZ-VI None 2002 Def Estab
NWP-4b None 1997 Native Estab
AUS-V None 2005 Def Estab
NEA-V None 0 Def Estab
NEA-IV None 0 Def Estab
AUS-VIII None 1975 Def Estab
AUS-IX None 1977 Def Estab
AUS-VII None 1971 Def Estab
AUS-XI None 1992 Def Estab
MED-VI None 0 Def Estab
MED-VII None 0 Def Estab
NEP-VI Pt. Conception to Southern Baja California 2011 Def Estab
P027 _CDA_P027 (Aliso-San Onofre) 2011 Def Estab
MED-II None 0 Def Estab
MED-IV None 0 Def Estab
AUS-IV None 2005 Def Estab
NWP-3b None 2007 Native Estab
NWP-3a None 2013 Native Estab
B-II None 1986 Def Estab
MED-IV None 2011 Def Estab
AUS-II None 2013 Def Estab
P050 San Pedro Bay 2017 Def Estab
P020 San Diego Bay 2004 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude


Atlas of Living Australia 2013-2016 Atlas of Living Australia.

Bilbao, Amaia Nunez, Jorge Viera, Beatriz Sosa Maria del Pino Fernandez-Palacios, Hipolito Hernavdez-Cruz, Carmen Maria (2011) Control of shell-boring polychaetes in (Reeve 1846) aquaculture: Species identification and effectiveness of Mebendazole, Journal of Shellfish Research 30: 331-336

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.; 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

Castellanos-Galindo, Gustavo A. ; Robertson, D. Ross ; Torchin, Mark E. (2020) A new wave of marine fish invasions through the Panama and Suez canals, Nature Ecology and Evolution 4: 1444–1446

Dauvin, Jean-Claude; Dewarumrez, Jean-Marie; Gentil, Franck (2003) [Updated list of polychaete annelids reported in the English Channel], Cahiers de Biologie Marine 44: 67-95

David, Andrew A. and Simon, Carol A. (2014) The effect of temperature on larval development of two non-indigenous poecilogonous polychaetes (Annelida:Spionidae) with implications for life history theory, establishment and range expansion, Journa of Experimental Marine Biology and Ecology 461: 20-30

David, Andrew A; Matthee, Conrad A.; Simon, Carol A. (2014) Poecilogony in Polydora hoplura (Polychaeta: Spionidae) from commercially important molluscs in South Africa, Marine Biology 161: 887-898
DOI 10.1007/s00227-013-2388-0

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

GBIF (Global Biodiversity Information Facility) 2017-2023 GBIF (Global Biodiversity Information Facility).

Handley, S. J. (1996) Spionid polychaetes in Pacific oysters, Crassostrea gigas (Thunberg) from Admiralty Bay, Marlborough Sounds, New Zealand, New Zealand Journal of Marine and Freshwater Research <missing volume>: 305-309

Handley, S. J.; Bergquist, P. R. (2009) Spionid polychaete infestations of intertidal Pacific oysters Crassostrea gigas (Thunberg) , Mahurangi Harbour, northern New Zealand, Aquaculture 153: 191-205

Inglis, Graeme and 6 authors (2005a) Dunedin Harbour (Port Otago and Port Chalmers): Baseline survey for non-indigenous marine species, Biosecurity New Zealand Technical Paper 2005/10: 1-49

Inglis, Graeme and 6 authors (2006e) Whangarei Harbour (Whangarei Port and Marsden Point: Baseline survey for non-indigenous species, Biosecurity New Zealand Technical Paper 2005(16): 1-52

Kerckhof, Francis; Haelters, Jan; Gollasch, Stephan G. (2007) Alien species in the marine and brackish ecosystem: the situation in Belgian waters., Aquatic Invasions 2(3): 243-257

Labura, Z.; Hrs-Brenko, M. (1990) Infestation of European Flat Oyster (Ostrea edulis) by polychaete (Polydora hoplura) in the northern Adriatic Sea, Acta Adriatica 31: 173-181

Levin, Lisa A. (1984a) Multiple patterns of development in Streblospio benedictiWebster (Spionidae) from three coasts of North America., Biological Bulletin 166: 494-508

Lleonart, M.; Handlinger, J.; Powell, M. (2003) Treatment of spionid mud worm (Boccardia knoxi Rainer) infestation of cultured abalone, Aquaculture 217: 1-10

Loosanoff, Victor L.; Engle, James B. (1943) Polydora in oysters suspended in the water, Biological Bulletin 85: 69-78

Mead, A.; Carlton, J. T.; Griffiths, C. L. Rius, M. (2011b) Introduced and cryptogenic marine and estuarine species of South Africa, Journal of Natural History 39-40: 2463-2524

Moreno, Rodrigo A.; Neill, Paula E.; Rozbaczylo, Nicolás (2006) Native and non-indigenous boring polychaetes in Chile: a threat to native and commercial mollusc species., Revista Chilena de Historia Natural 79: 263-278

Nel, Ronel; Coetzee, P. S.; Van Niekerk, G (1998) The evaluation of two treatments to reduce mud worm (Polydora hoplura Claparede) infestation in commercially reared oysters (Crassostrea gigas Thunberg), Aquaculture 146: 31-39

Parapar, Julio Martínez-Ansemil, Enrique Caramelo, Carlos Collado, Rut Schmelz, Rüdig (2009) Polychaetes and oligochaetes associated with intertidal rocky shores in a semi-enclosed industrial and urban embayment, with the description of two new species, Helgoland Marine Research 63: 293-308
DOI 10.1007/s10152-009-0158-7

Radashevsky, Vasily (1999) Redescription of the proposed lectotype for Polydora websteri Hartman in Loosanoff & Engle, 1943 (Polychaeta: Spionidae), Ophelia 2: 107-113

Radashevsky, Vasily I. Migotto, Alvaro E. (2016) First report of the polychaete Polydora hoplura (Annelida: Spionidae) from North and South America and Asian Pacific, Marine Biodiversity 47: 859–868
DOI 10.1007/s12526-016-0515-0

Radashevsky, Vasily I.; . Malyar, Vasily V.; Pankova, Victoria V. ;Choi, Jin-Woo; Yum, Seungshic; Carlton , James T. (2023) Searching for a Home Port in a Polyvectic World: Molecular Analysis and Global Biogeography of the Marine Worm Polydora hoplura (Annelida: Spionidae), Biology 12(780): Published online biology12060780

Radashevsky, Vasily I.; Olivares, Carolina (2005) Polydora uncinata (Polychaeta: Spionidae in Chile: an accidental transportation across the Pacific., Biological Invasions 7: 489-496

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

Rodewald, Nicola; Snyman, Reinette; Simon, Carol A. (2021) Worming its way in Polydora websteri (Annelida: Spionidae) increases the number of non-indigenous shell-boring polydorin pests of cultured molluscs in South Africa, Zootaxa 4949: 255-279

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.

Sato-Okoshi, Waka (1998) Three new species of polydorids (Polychaeta, Spionidae) from Japan, Species Diversity 3: 277-288

Sato-Okoshi, Waka; Abe, Hirokazu (2013) Morphology and molecular analysis of the 18S rRNA gene of oyster shell borers, Polydora species (Polychaeta: Spionidae), from Japan and Australia, Journal of the Marine Biological Association of the United Kingdom 93(5): 1279–1286

Sato-Okoshi, Waka; Abe, Hirokazu; Nishitani, Goh; Simon, Carol A. (2018) And then there was one: Polydora uncinata and Polydora hoplura (Annelida: Spionidae), the problematic polydorid pest species represent a single species, Journal of Marine Biological Association of the United Kingdom 97(8): 1675–1684.

Simon, Carol A. (2011) Polydora and Dipolydora (Polychaeta: Spionidae) associated with Molluscs on the south coast of South Africa, with descriptions of two new species, African Invertebrates 52: 39-50

Simon, Carol A.; Bentley, Matthew G.; Caldwell, Gary S. (2010) 2,4-Decadienal: Exploring a novel approach for the control of polychaete pests on cultured abalone, Aquaculture 310: 52-60

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

Wilson, Douglas P. (1938) The larval development of Polydora ciliata Johnston and Polydora hoplura Claparede, Journal of Marine Biological Association of the United Kingdom 15: 567- 603