Invasion History

First Non-native North American Tidal Record: 1921
First Non-native West Coast Tidal Record: 1921
First Non-native East/Gulf Coast Tidal Record: 1951

General Invasion History:

The tubeworm Ficopomatus enigmaticus was first described from Caen, France, on the English Channel Coast (Fauvel 1923, cited by Carlton 1979; Goulletquer et al. 2002). However, the native range of this species is unknown. Fauvel (1923; cited by Carlton 1979) hypothesized an Indian/Indonesian origin, but many authors have ruled this out. It first appeared more or less simultaneously in France, Spain, England, Australia and California (Fauvel 1923; cited by Carlton 1979). The southern coast of Australia is a more likely (though still uncertain) native region (Allen 1953; ten Hove and Weerdenburg 1978; Keough and Ross 1999). Since then it has been recorded from Hawaii, the East and Gulf coasts of the US, the Baltic Sea (Jensen and Knudsen 2005), the Mediterranean (Bianchi and Morri 1996), the Caspian Sea (Grigorovich et al. 2003), Africa (ten Hove and Weerdenburg 1978), South America (Obenat and Pezzani 1991), New Zealand (Read and Gordon 1991), Japan (Asakura 1992) and Taiwan (ten Hove and Weerdenburg 1978).

North American Invasion History:

Invasion History on the West Coast:

Ficopomatus enigmaticus was first reported from San Francisco Bay, California in 1921, when it was found building conspicuous reefs in Lake Merritt lagoon (Carlton 1979). By the 1970s, it was widespread in the Bay, occurring in the central Bay at Alameda and Berkeley, in the south Bay at Palo Alto and Foster City, and in San Pablo Bay at Corte Madera Creek, San Rafael, and San Quentin (Carlton 1979). It has been collected in brackish water in the Petaluma and Napa Rivers (Cohen and Carlton 1995; Cohen et al. 2005). In 1994, F. enigmaticus was collected in the old Salinas River Channel at Moss in the Elkhorn Slough system. It was subsequently found at several locations in Elkhorn Slough during a 1998 survey (Wasson et al. 2001).

This serpulid is established in Southern California, but its distribution is spotty. In 2000, a few specimens of Ficopomatus enigmaticus were collected in Los Angeles Habor (Cohen et al. 2002; Bastida-Zavala et al. 2008). Collections were also reported from San Diego Bay in 2000 (Bastida-Zavala et al. 2008), but these records were erroneous (Bastida-Zavala, pers. comm., 10 November 2015, cited by Pernet et al. 2016). In 2011, 86 specimens were collected on fouling plates in Newport Bay, although none were found in Los Angeles Harbor or San Diego Bay (California Department of Fish and Game 2014). In 2008 and 2015, F. enigmaticus was found in two lagoons (Arroyo Burro Creek, Mission Creek Lagoon) in Santa Barbara County. A population was found along the lower 3 km of the Los Angeles River, in the vicinity of marinas (Pernet et al. 2016). Yee et al. (2019) found unexpectedly high genetic diversity in California F. enigmaticus populations, suggesting multiple introduction.

Invasion History on the East Coast:

Ficopomatus enigmaticus was found on settling plates in Barnegat Bay, New Jersey in 1976 (Hoagland and Turner 1980) and it appears to be established there, although at low densities (Loveland and Shafto 1984). In 1979, it was collected on the Atlantic Coast of Florida, in the Banana River, Indian River Lagoon (Fofonoff, unpublished data). In 1994-1995, F. enigmaticus was found on settling plates in Chesapeake Bay in Norfolk, Virginia; Baltimore Harbor, Maryland; and the Severn River, Maryland (McCann et al. unpublished; Ruiz et al. unpublished). In 2006, a single specimen was collected in the East River, Brunswick, Georgia (USGS Nonindigenous Aquatic Species Program 2007).

Invasion History on the Gulf Coast:

Ficopomatus enigmaticus was found on a boat in Corpus Christi, Texas in 1951 (Hartman 1952), and was subsequently found in the same locality on riprap by Andy Cohen in 1995 (McCann and Carlton, unpublished). It was also reported from Aransas Bay, Texas in 1952 (ten Hove and Weerdenburg 1978); Galveston Bay, Texas in 2003 (Ruiz et al. unpublished data); Pascagoula River, Mississippi in 1997 (USNM 186522, U.S. National Museum of Natural History 2007); and Tampa Bay, Florida in 2002 (Ruiz et al., unpublished data).

Invasion History in Hawaii:

Ficopomatus enigmaticus was first collected in 1937 from Pearl Harbor, Hawaii (Coles et al. 1999b). Subsequently, in 1947, it was found in other Oahu locations, including the Alai Wai Canal and Kewalo Basin (ten Hove and Weerdenburg 1978; Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

Ficopomatus enigmaticus may be native to the southern coast of Australia, where it was first reported from Sydney in 1932 (ten Hove and Weerdenburg 1978). It also occurs in Port Phillip Bay, South Australia, and Western Australia (ten Hove and Weerdenburg 1978; Keough and Ross 1999). However, it is a relatively recent invader to New Zealand, where it was first collected in 1968, and is confined to the North Island (Read and Gordon 1991). Elsewhere in the Pacific, F. enigmaticus was collected from Kojima Bay, Japan in 1966 (Asakura 1992) and Taiwan (ten Hove and Weerdenburg 1978, no location details).

In Europe, as noted above, F. enigmaticus was first reported in 1921, from Caen, France (Fauvel 1923, cited by Carlton 1979; Goulletquer et al. 2002). It spread in a rapid, but spotty fashion throughout European estuaries, appearing most often in brackish lagoons, sometimes near thermal effluents. It was reported from docks in the Thames Estuary, London, England in 1923 (ten Hove 1978; Zibrowius and Thorp 1989); Plymouth, England in 1939 (Zibrowius and Thorp 1989); the Gulf of Cadiz, Spain in 1933 (ten Hove and Weerdenburg 1978); the Po River Delta on the Adriatic Sea in 1934 (Bianchi and Morri 1996); the Black Sea in 1929 (Gomiou et al. 2002); a lagoon in Israel in 1954 (Galil 2007); and the Lac de Tunis, Tunisia in 1969 (ten Hove and Weerdenburg 1978). In the Baltic Sea, it was found in Copenhagen Harbor in 1953, where it is confined to the vicinity of a thermal effluent (Jensen and Knudsen 2005). In 1961, it was first collected in the Caspian Sea, which it reached by shipping transport through canals (Grigorovich et al. 2003).

In 2016, F. enigmaticus was found in a degraded, landlocked, brackish marsh, Praia da Vitória Marsh, on the island of Terceira in the Azores. Since, the marsh is connected to the Atlantic only by water percolating through sand dunes, Costa et al. (2019) consider migratory birds to be likeliest vector.

Ficopomatus enigmaticus has colonized port areas on both sides of the south Atlantic, reaching the Rio de la Plata estuary, Uruguay in 1937 and Port Quequen, Argentina, in 1940 (Rioja 1943; Schwindt et al. 2001; Orensanz et al. 2002). On the other side of the Atlantic, this worm was first collected in Milnerton estuary, in Table Bay, near Cape Town, South Africa in 1951, and ranges around the tip of the continent, to Kosi Bay, on the Indian Ocean (Mead et al. 2011b).


Description

Ficopomatus enigmaticus secretes a calcareous tube, as do other serpulid polychaetes. Serpulids have a feathery crown of modified prostomial palps, called radioles (the prostomium is the first segment, projecting above the mouth). The radioles can be folded and withdrawn into the tube. One of the radioles is modified to form an operculum, which acts as a plug when the animal contracts. The peristomium (segment behind the mouth) is folded back to form a collar, which bears uniramous parapodia, with a distinctive set of collar chaetae, with spines or serrations. The collar is the first of seven thoracic chaeta-bearing segments (setigers). The subsequent segments have biramous parapodia. The dorsal branch of the parapodium is called the notopodium; the ventral is the neuropodium. Chaetae in the two branches and along the body can vary greatly in their morphology, which can be critical in the taxonomy of serpulids (Description from: Barnes 1983; ten Hove and Weerdenburg 1978; Blake and Ruff, in Carlton 2007).

The tube of F. enigmaticus is white, but often covered by a brown layer of algae. It is emicircular to circular in cross-section, and often bears irregularly placed flaring rings, indicating the previous positions of the peristome. Solitary and juvenile tubes may have faint longitudinal keels. The tubes are about 1 mm in diameter. The branchial (gill) radioles arise from paired lobes. There are about 7 (5-9) on the left and 8 (7-10) on the right, forming two semicircles on either side of the mouth. The feathery filaments of the branchiae (pinnulae) are larger towards the ends of the radioles. The operculum is shaped like an elongated funnel, with a concave distal surface and circular rows of brown inwardly curved terminal spines. The gills and operculum account for about 1/6 of the length of the worm. The peduncle of the operculum is D-shaped in cross-section. The collar (first thoracic segment) is high and not lobed, with a smooth edge. The thorax consists of 7 segments. There are two kinds of collar chaetae, thicker coarsely serrated chaetae, and hair-like (limbate). The subsequent thoracic segments bear short, rasplike chaetae, called uncinae. and limbate chaetae. The abdomen has about 60 segments (29-84, n=7, ten Hove and Weerdenburg 1978). The overall length of the worm reaches about 26 mm- the mean of a sample from New Zealand was 14.7 mm (Read and Gordon 1991; Bianchi and Morri 2001). The gills are green, brown, or white, often with the gill filaments barred. This worm may grow in colonies consisting of large masses of brown and white worm tubes. It is typically found in estuaries, particularly in brackish waters. (Description from: ten Hove and Weerdenburg 1978; Hayward and Ryland 1990; Read and Gordon 1991; Blake and Ruff, in Carlton 2007).

Styan et al. (2017) surveyed F. enigmaticus in southern Australia, a possible native region. They expected to see geographical patterns of genetic variation between populations in southeastern and southwestern Australia, a common feature of biogeography in the region. Instead, 2 clades, apparently cryptic species, had a mixed distribution in the two regions. A third clade, morphologically resembling F. uschakovi, was found in two southeastern estuaries (Styan et al. 2017). Genetic comparisons of definitely introduced populations with the cryptogenic Australian populations have not yet been made. Grosse (2021) identifed 3 clades form Majorca. Spain. One was the widespread Clae 1, and the others, Clades 4 and 5, are known only from Majorca but presumably introduced.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Palpata
Order:   Canalipalpata
Suborder:   Sabellida
Family:   Serpulidae
SubFamily:   Serpulinae
Genus:   Ficopomatus
Species:   enigmaticus

Synonyms

Mercierella enigmatica (Fauvel, 1923)

Potentially Misidentified Species

Ficopomatus macrodon
Native to the Indo-Pacific

Ficopomatus miamiensis
Native FL-Atlantic Panama, introduced to Pacific in Mexico and Panama

Ficopomatus uschakovi
Native to the Indo-Pacific, introduced from Jacksonville FL to Corpus Christi TX, Venezuela, Brazil, West Africa, Pacfic Panama

Ecology

General:

The serpulid polychaete Ficopomatus enigmaticus feeds by extending its feathery gills to trap plankton in the water column. The plankton is then transported by cilia to the mouth. In most populations, the two sexes are separate (Tunisia- Vuillemin 1965; Argentina- Obenat and Pezzani 1994), but in at least one population (Thames estuary, England), a small percentage of hermaphroditic individuals were seen (Dixon 1981). Spawning was seasonal (June – October) in the temperate Thames estuary, apparently triggered by temperatures rising above 18C (Dixon 1981). In the Mar Chiquita, Argentina, there were double spawning seasons, November-December and April-May (Obenat and Pezzani 1994). Eggs and sperm are released into the water column, and the larvae are planktotrophic. Based on studies of gametogenesis and settlement, larval development time was estimated at 45-90 days in the Thames estuary (Dixon 1981). However, to our knowledge, the larvae have not been reared in the laboratory.

Ficopomatus enigmaticus tolerates a wide range of temperatures and salinity. At the northern limits of its range in North America and Europe, its establishment seems to be favored by thermal effluents (ten Hove 1974: Turner and Hoagland 1980; Jensen and Knudsen 2005). A temperature of 18C is apparently required for spawning (Dixon 1981). This tubeworm tolerates salinities ranging from 1 to 55 PSU, but seems especially abundant in brackish waters (ten Hove and Weerdenberg 1978), possibly because of a lack of competition. It settles and secretes a calcareous tube on surfaces such as rocks, shells, plant leaves, pilings and pontoon floats. While F. enigmaticus is widespread, reef development does not always occur. Shallow estuaries and lagoons with long retention times appear especially prone to reef formation. The reefs begin as clumps of intertwined tubes, attached to sticks, stone, shells etc., and grow over many generations of settlement. Reefs can reach 0.5 m in thickness and 7 m across (Bianchi and Morri 1996; Schwindt et al. 2001).

Food:

Phytoplankton

Trophic Status:

Suspension Feeder

SusFed

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatOyster ReefNone
General HabitatMarinas & DocksNone
General HabitatUnstructured BottomNone
General HabitatCanalsNone
General HabitatRockyNone
General HabitatVessel HullNone
General HabitatGrass BedNone
Salinity RangeOligohaline0.5-5 PSU
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Salinity RangeHyperhaline40+ PSU
Tidal RangeSubtidalNone
Tidal RangeLow IntertidalNone
Vertical HabitatEpibenthicNone

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)0Based on geographical range
Maximum Temperature (ºC)30Lake Timsah, Israel (Ben-Eliahu and ten Hove 2011); Lagoon of Orbetello, Italy (Bianchi and Morri 2001).
Minimum Salinity (‰)2Vuillemin 1965; Bianchi and Morri, 1996)
Maximum Salinity (‰)55Vuillemin 1965
Minimum Reproductive Temperature18Thames estuary, spawning. Settlement of larvae occurs as temperatures as low as 10 C (Dixon 1981)
Minimum Reproductive Salinity10Vuillemin 1965; Thomas and Thorp 1994
Minimum Duration30Larval period, field estimate (Dixon 1981)
Maximum Duration90Larval period, field estimate (Dixon 1981)
Maximum Length (mm)25Lagoon of Orbetello, Italy (Bianchi and Morri 2001).
Broad Temperature RangeNoneCold temperate-Subtropical
Broad Salinity RangeNoneOligohaline-Euhaline
Minimum Temperature (ºC)1Field data, Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data
Maximum Temperature (ºC)30Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data
Minimum Salinity (‰)1Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data
Maximum Salinity (‰)55Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data

General Impacts

Economic Impacts: In a number of estuaries around the world, the tubeworm Ficopomatus enigmaticus is a major fouling organism, covering man-made surfaces with thick layers of calcareous tubes. It created fouling problems on boats, pilings and pontoons in New Zealand (Read and Gordon 1991), and frequently blocks lock gates and other structures used to control flow in estuaries (Nelson-Smith 1971; Zibrowius 1994).

Ecological Impacts: In many brackish and hypersaline areas around the world, F. enigmaticus has formed large reef-like colonies, greatly affecting estuarine habitats. These calcareous masses can be 1-3 m across, reaching to within a few centimeters of the water's surface, causing reduced circulation, increased sedimentation, increased algal growth, and affecting planktonic and benthic communities through filtering of suspended particles (Davies et al. 1989; Thomas and Thorp 1994; Cohen and Carlton 1995; Bianchi and Morri 1996; Schwindt et al. 2001; Bruschetti et al. 2008).

Herbivory- Ficopomatus enigmaticus reefs can filter significant quantities of phytoplankton and was estimated to clear the volume of the Marina del Gama (near Cape Town, South Africa) in 26 hours (Davies et al. 1989). Phytoplankton chlorophyll decreased by up to 56% near reefs in summer in the Mar Chiquita lagoon, Argentina (Bruschetti et al. 2008).

Competition- In the Lagune Orbetello, Italy, F. enigmaticus tubes outnumbered and outgrew those of another introduced serpulid (Hydroides dianthus), which produces thicker and stronger tubes, but with slower growth (Bianchi and Morri 2001).

Habitat Change- In England, F. enigmaticus forms extensive reefs in sheltered lagoons, increasing habitat for attached organisms, and shelter for mobile ones (Thomas and Thorp 1994). In the Po River Delta, Italy, it forms solid habitat for attached organisms. The tubes break down to create gravelly, carbonaceous sediments (Bianchi and Morri 1996). In California, F. enigmaticus was first noticed in Lake Merritt, San Francisco Bay, when large reefs of worm-tubes were discovered. Ecological impacts of these reefs have not been studied and abundances of the worms are now lower than in the early 20th century (Carlton 1979; Cohen and Carlton 1995). Ficopomatus reefs are also spreading in Elkhorn Slough. These reefs support higher densities of organisms than native oyster reefs, predominantly non-native species (Heiman et al. 2008). Reef-building in Mar Chiquita, Argentina, provided new habitat for the crab Cyrtograpsus angulatus (Schwindt et al. 2001). Reefs constructed by F. enigmaticus greatly increased sedimentation and decreased water circulation in the Mar Chiquita Lagoon, Argentina (Schwindt et al. 2001). Reefs had an increased frequency of epibiotic fauna compared to bare mud areas (amphipods, crabs, snails), and provided a more frequently used feeding and resting habitat for migratory shorebirds (Bruschetti et al. 2009).

Trophic Cascade- In the Mar Chiquita Lagoon, Argentina, the reefs supported increased populations of the crab Cyrtograpsus angulatus, resulting in increased predation and reduced abundance of infauna (Schwindt et a. 2001). The reefs also had increased use by resident and migratory shorebirds, but exclusion experiments did not show detectable evidence of increased mortality (Bruschetti et al. 2009).


Regional Impacts

NEP-VNorthern California to Mid Channel IslandsEcological ImpactHabitat Change
Ficopomatus enigmaticus was first noticed in Lake Merritt, San Francisco Bay, when large reefs of worm-tubes were discovered. Ecological impacts of these reefs have not been studied and the abundance of worms is now lower than in the early 20th century (Carlton 1979; Cohen and Carlton 1995). Ficopomatus reefs are also spreading in Elkhorn Slough. These reefs support higher densities of organisms than native oyster reefs, predominantly non-native species (Heiman et al. 2008).
NEA-IINoneEcological ImpactHabitat Change
Ficopomatus enigmaticus formed extensive reefs in sheltered lagoons, increasing habitat for attached organisms and shelter for mobile ones (Thomas and Thorp 1994). Ficopomatus enigmaticus fouling provided habitat for 3 introduced species (Amphibalanus improvisus, Rhithropanopeus harrisii, Mytilopsis leucophaeta) (Charles et al. 2018).
NEA-IINoneEcological ImpactHerbivory
The large biomass of F. enigmaticus in a lagoon in England appears to have extensively filtered the water, reducing phytplankton biomass (Thomas and Thorp 1994).
NEA-IINoneEcological ImpactTrophic Cascade
The growth of extensive reefs of F. enigmaticus shifted biomass and nutrients from the water column to the benthos (Thomas and Thorp 1994).
NEA-IINoneEconomic ImpactShipping/Boating
Ficopomatus enigmaticus often fouls boats and blocks canals and tide gates in coastal lagoons (Nelson-Smith 1971; Eno 1996). 'However, at CUMB2 (marina in Cumbria, England) it was super-abundant and a severe fouling nuisance on yacht hulls, pontoons and ropes. This marina has previously requested advice from us as to how to remove it, has advised boat owners on what they can do about it, and has carried out cleaning and procedural' (Wood et al. 2015). Ficopomatus enigmaticus is also perceived as having negative impacts in marinas in Normandy (Charles et al. 2018).
WA-IVNoneEcological ImpactHerbivory
Filtation of phytoplankton by F. enigmaticus was estimated to clear the volume of the Marine del Gama in 26 hours (Davies et al. 1989).
WA-IVNoneEcological ImpactHabitat Change
Ficopomatus enigmaticus was encrusting submerged vegetation (Stukenia pectinata, Sago Pondweed (mentioned by Davies et al. 1989). Increasing water clarity is implied, but not discussed by Davies et al. (1989). A major change is the transformation of the estuary from sand-banks to encrusted concrete walls and reefs, with an increase of infaunal biomass and abundance as well (McQuaid and Griffiths 2014).
WA-IVNoneEconomic ImpactShipping/Boating
Ficopomatus enigmaticus was encrusting docks and concrete walls- a safety hazard to recreational users (Davies et al. 1989).
WA-IVNoneEconomic ImpactAesthetic
Filtration by Ficopomatus improves water clarity (Davies et al. 1989).
MED-VIINoneEcological ImpactHabitat Change
Ficopomatus enigmaticus formed extensive reefs in lagoons of the Po River delta, forming solid habitat for attaching organisms. The tubes break down to create gravelly, carbonaceous sediments (Bianchi and Morri 1996). In the delta of the Neretva River, dense colonies of F. enigmaticus were colonized by Musculista senhousia , Amphibalanus ebureneus, and a variety of naitve fouling organisms (Despalatovic et al. 2013).
NZ-IVNoneEconomic ImpactIndustry
Ficopomatus enigmaticus fouled cooling systems of power plants (Read and Gordon 1991).
NZ-IVNoneEconomic ImpactShipping/Boating
Ficopomatus enigmaticus fouled pleasure boats, pontoons and pilings with encrustations of tubes (Read and Gordon 1991).
MED-IIINoneEcological ImpactCompetition
In the Orbetello lagoon, Italy, Ficopomatus enigmaticus tubes outnumbered and outgrew those of another introduced serpulid (Hydroides dianthus), which produces thicker and stronger tubes, but with slower growth (Bianchi and Morri 2001).
P080Monterey BayEcological ImpactHabitat Change
Ficopomatus reefs are spreading in Elkhorn Slough. These reefs support higher densities of organisms than native oyster reefs and mudflats, predominantly favoring non-native species (Heiman et al. 2008; Heiman and Micheli 2010).
P090San Francisco BayEcological ImpactHabitat Change
Ficopomatus enigmaticus was first noticed in Lake Merritt, San Francisco Bay, when large reefs of worm-tubes were discovered. Ecological impacts of these reefs have not been studied and the abundance of worms is now lower than in the early 20th century (Carlton 1979; Cohen and Carlton 1995).
MED-IXNoneEcological ImpactHabitat Change
'The mass populations of this species induce changes in benthic communities, offering at the same time ecological niches and substrata for various species including other fouling species' (Micu and Micu 2004, cited by Skolka and Preda 2010).
MED-IXNoneEconomic ImpactShipping/Boating
'Fouling organisms, like barnacles or the polychaete Ficopomatus enigmaticus, are harmful because their biomass increases the fuel consumption of ships. Such organisms also increase the corrosion rate of metallic submerged structures.' (Skolka and Preda 2010).
NWP-3bNoneEconomic ImpactFisheries
Ficopomatus enigmaticus interfered with oyster culture in Lake Hamana (Chavanich et al. 2010).
SA-INoneEcological ImpactHabitat Change
Ficopomatus enigmaticus reefs in Mar Chiquita lagoon, Argentina, provided new habitat for the crab Cyrtograpsus angulatus (Schwindt et al. 2001). Reefs also greatly increased sedimentation and decreased water circulation throughout the lagoon (Schwindt et al. 2001).  There was an increased frequency of epibiotic fauna on serpulid reefs compared to bare mud areas (amphipods, crabs, snails), and the reefs provided a more frequently used feeding and resting habitat for migratory shorebirds (Bruschetti et al. 2009).  Reef worms deposit large quantities of organic material as feces and pseudofeces, altering sedimentary habitats (Bruschett et al. 2011). Reef building also increased grazing by herbivores, resulting in the elimination of green algae (Ulva, etc.), while providing habitat for settlement of red algae (Polysiphonia subtilissima) (Bazterrica et al. 2011). This impact may be mutualistic. Ficopomatus colonized by P. subtilissima had increased recruitment, tube length, and body condition (Bazterrica et al. 2014).
SA-INoneEcological ImpactHerbivory
Phytoplankton chlorophyll decreased by up to 56% near reefs in summer, due to suspension-feeding by F. enigmaticus (Bruschetti et al. 2008). Grazing rates, in Mar Chiquita and La Tigra Creek, were greatest for diatoms, resulting in dominance of the phytoplankton by picoplanton (nanochlorophytes) (Pan and Marcoval 2013). In mesocosm experiments, nutrient addicitions promoted phytplankton gorwth, but this was offset by increased biomass and grazing of F. enigmaticus (Bruschetti et al. 2018).
SA-INoneEcological ImpactTrophic Cascade
Increased populations of the crab Cyrtograpsus angulatus  resulted in increased predation and reduced abundance of infauna (Schwindt et al. 2001). Ficopomatus reefs also provided a center of concentration for two native snails (Heleobia conexa and Heleobia australis), resulting in increased prevalence of digenean trematodes (15 spp.) whose life cycle includes parasitism of shorebirds (Etcehgoin et al. 2012).
MED-IIINoneEcological ImpactHabitat Change
In the Lac du Tunis, Tunisia, F. enignmaticus constructed extensive reefs on formerly bare sediments, filling in the basin and reducing circulation. Filtration increased water clarity, stimulating growth of macroalgae. Decaying macroalgae can result in hypoxic conditions (Keene 1980).
MED-IIINoneEcological ImpactTrophic Cascade
Filtration by F. enigmaticus shifts biomass from the water column to the benthos. Filtration increases water clarity, stimulating growth of macroalgae (Keene 1980).
MED-IINoneEcological ImpactHabitat Change
The reefs of Ficopomatus enigmaticus are major features in the Albufera Lagoon, forming calcareous reefs 2-3 m thick. The reefs have the potential to eventually fill up the lagoon (Fornos et al. 1997).
SA-IINoneEcological ImpactHabitat Change
Ficopomatus enigmaticus created reefs in Uruguayan coasta lagoons, which were considered to be an indicator of a perturbed environment (Muniz et al. 2005). Ficopomatus enigmaticus reefs attract dense populations of the introduceed amphipod Melita palmata (Bazterrica et al. 2020).
SA-IINoneEconomic ImpactIndustry
Ficopomatus enigmaticus obstructed the cooling systems of oil refineries in Uruguay (Muniz et al. 2005).
SA-INoneEcological ImpactPredation
In mesocosm experiments, Ficopomatus reefs in the Mar Chiquit, Argentina, did not affect overall zooplantkon biomass, but did affect community composition, especially the abundace of cladocerans (mostly Moina spp. (Bruschetti et al. 2016).
NEP-VNorthern California to Mid Channel IslandsEcological ImpactCompetition
Transplant experiments indicate that competition with Ficopomatus enigmaticus may limit growth and abundance of the introduced bryozoan Conopeum chesapeakensis at the more saline end of Carquinez Strait in the San Francisco estuary (Benicia, ~21 PSU, Newcomer et al. 2018).
P090San Francisco BayEcological ImpactCompetition
Transplant experiments indicate that competition with Ficopomatus enigmaticus may limit growth and abundance of the introduced bryozoan Conopeum chesapeakensis at the more saline end of Carquinez Strait in the San Francisco estuary (Benicia, ~21 PSU, Newcomer et al. 2018).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1951 Def Estab
NEP-V Northern California to Mid Channel Islands 1921 Def Estab
NEP-VI Pt. Conception to Southern Baja California 2000 Def Estab
NA-ET3 Cape Cod to Cape Hatteras 1976 Def Estab
NEA-II None 1921 Def Estab
NEA-III None 1939 Def Estab
MED-VII None 1934 Def Estab
B-III None 1953 Def Estab
NEA-IV None 1922 Def Estab
NEA-V None 1978 Def Estab
MED-II None 1933 Def Estab
MED-IV None 1978 Def Estab
MED-III None 1927 Def Estab
MED-V None 1937 Def Estab
MED-IX None 1929 Def Estab
CASP Caspian Sea 1961 Def Estab
WA-IV None 1951 Def Estab
SP-XXI None 1937 Def Estab
AUS-VIII None 1975 Crypto Estab
AUS-X None 1932 Crypto Estab
AUS-IV None 1935 Crypto Estab
AUS-VII None 1975 Crypto Estab
SA-II None 1917 Def Estab
NZ-IV None 1967 Def Estab
NWP-3b None 1966 Def Estab
NWP-2 None 1978 Def Estab
SA-I None 1940 Def Estab
G070 Tampa Bay 2002 Def Estab
G260 Galveston Bay 2003 Def Estab
G310 Corpus Christi Bay 1951 Def Estab
P050 San Pedro Bay 2000 Def Estab
M130 Chesapeake Bay 1994 Def Estab
S190 Indian River 1979 Def Estab
P080 Monterey Bay 1994 Def Estab
P090 San Francisco Bay 1921 Def Estab
G160 East Mississippi Sound 1997 Def Estab
M070 Barnegat Bay 1976 Def Estab
G300 Aransas Bay 1952 Def Estab
S160 St. Andrew/St. Simons Sounds 2006 Def Unk
CAR-VII Cape Hatteras to Mid-East Florida 2006 Def Estab
P093 _CDA_P093 (San Pablo Bay) 1921 Def Estab
MED-VIII None 1952 Def Estab
MED-VI None 1976 Def Estab
CIO-II None 2004 Crypto Estab
WA-I None 1978 Def Estab
NWP-4a None 2000 Def Estab
S080 Charleston Harbor 2004 Def Estab
MED-X None 1961 Def Estab
RS-3 None 1973 Def Estab
WA-V None 1975 Def Estab
NWP-4b None 0 Def Estab
P040 Newport Bay 2011 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 2008 Def Estab
AUS-XI None 0 Crypto Estab
NEA-VI None 2016 Def Estab
P076 _CDA_P076 (Carmel) 2001 Def Estab
P064 _CDA_P064 (Ventura) 2013 Def Estab
P020 San Diego Bay 2013 Def Unk
NEA-VI None 2016 Def Estab
P060 Santa Monica Bay 2015 Def Estab
MED-V None 2018 Def Estab
B-IV None 2020 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
2720 Hoagland and Turner 1980 1976 1976-01-01 Oyster Creek (Barnegat Bay) Def 39.4092 -74.6632
2721 Hoagland and Turner 1980 1976 1976-01-01 Forked River (Barnegat Bay) Def 39.8397 -74.1606
2722 Ruiz et al. unpublished data 1994 1994-01-01 Scott Creek (Elizabeth River) Def 36.8511 -76.3083
2736 Hartman 1951 1951 1951-01-01 Corpus Christi Bay Def 27.7719 -97.2525
2737 Ruiz et al. unpublished data 2002 2002-09-17 Marina Landing Resort, Galveston Def 29.2756 -94.8481
2738 Ruiz et al. unpublished data 2002 2002-09-16 Payco Marina, Galveston Def 29.2871 -94.8733
2739 Ruiz et al. unpublished data 2002 2002-09-19 Port of Texas City Def 29.3728 -94.8907
2740 Ruiz et al. unpublished data 2002 2002-09-17 West Bay Marina, Def 29.1455 -95.0472
2741 Ruiz et al. unpublished data 2002 2002-07-30 Mariner's Club, Tampa area Def 27.7272 -82.4748
2742 USNMNH catalogue # 186522, ID by Thomas Perkins 1997 1997-11-11 Mississippi Sound Def 30.3958 -88.8853
2745 Carlton 1979 1954 1954-01-01 Berkeley Aquatic Park Lagoon Def 37.8318 -122.3603
2746 Carlton 1979 1971 1971-01-01 Alameda Lagoon Def 37.7652 -122.2416
2747 Carlton 1979 1970 1970-01-01 San Rafael Creek Def 37.9672 -122.4853
2754 Ruiz et al. unpublished 2000 2000-09-09 Port of Richmond Def 37.5121 -122.2108
2755 Ruiz et al. unpublished 2001 2001-09-21 Port of Redwood Def 37.9090 -122.3878
2799 Straughan 1969 1947 1947-01-01 Alai Wai Canal Def 20.4306 -156.3742
2800 Straughan 1969 1943 1943-01-01 Kewalo Basin Def 21.2961 -157.8603
2801 Straughan 1969 1937 1937-01-01 Pearl Harbor Def 21.3550 -157.9722
7230 Ruiz et al. unpublished data 1994 1994-01-01 Baltimore Harbor Def 39.2903 -76.6125
7231 Ruiz et al. unpublished data 1994 1994-01-01 Severn River Def 38.9583 -76.4456
7232 Ruiz et al., unpublished data 2001 2001-09-01 Belle Isle Marina Def 37.0964 -76.2920
7233 Ruiz et al., unpublished data 2001 2001-09-01 Lynhaven Bay Pt. Def 37.1542 -76.3800
7234 Ruiz et al., unpublished data 2004 2004-01-01 Charleston Def 32.7638 -79.8973
7236 Harbor Branch Oceanographic Institiution 1979 1979-01-01 Merritt Island Def 28.2197 -80.6448
26664 Wasson et al, 2001 (Elkhorn Slough Survey) 1998 1998-03-01 Elkhorn Slough Station 9 Def 36.8398 -121.7435
26830 Foss 2009 2005 2005-11-14 California Maritime Academy/Vallejo Def 38.0661 -122.2299
27478 Carlton 1979; Cohen and Carlton, 1995 1969 1969-01-01 Berkeley Marina, San Francisco Bay Def 37.8664 -122.3150
27511 Carlton 1979; Cohen and Carlton, 1995 1974 1974-01-01 Palo Alto, San Francisco Bay Def 37.4584 -122.1052
27557 Wasson et al, 2001 (Elkhorn Slough Survey) 1998 1998-03-01 Elkhorn Slough Station 10 Def 36.8578 -121.7572
27864 Foss 2011 2010 2010-07-01 Loch Lomond Marina Area Def 37.9720 -122.4832
27952 Wasson et al, 2001 (Elkhorn Slough Survey) 1998 1998-03-01 Elkhorn Slough Station 1 Def 36.7908 -121.7906
27966 California Department of Fish and Wildlife 2011 2011 2011-06-21 B-Dock Def 36.8027 -121.7851
28508 Carlton 1979; Cohen and Carlton, 1995 1970 1970-01-01 Corte Madera Creek, San Francisco Bay Def 37.9426 -122.5081
28568 Foss 2009 2005 2005-11-15 Petaluma River Turning Basin Def 38.2344 -122.6354
28573 Carlton 1979; Cohen and Carlton, 1995 1979 1979-01-01 Belvedere Lagoon, San Francisco Bay Def 37.8787 -122.4666
32807 California Department of Fish and Wildlife 2011 2011 2011-04-19 Back Bay Marina Def 33.6194 -117.8933
35687 Pernet et al. 2016 2008 2008-01-01 Mission Creek Lagoon Def 34.4126 -119.6882
35688 Pernet et al. 20162015 None 2015-10-01 Arroyo Burro Creek Def 34.4034 -119.7432
35689 Pernet et al. 2015 2015 2015-10-01 Anaheim-Shoreline Bridges, Los Angeles River Def 33.7792 -118.2047
35690 Pernet et al. 2016 2015 2015-10-01 Long Beach Shoreline Marina Jetty Def 33.7581 -118.1899
767813 Ruiz et al., 2015 2011 2012-09-20 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6979 -122.1912
767845 Ruiz et al., 2015 2011 2011-09-16 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9724 -122.4796
767862 Ruiz et al., 2015 2011 2011-09-27 Petaluma Marina, San Francisco Bay, CA, California, USA Def 38.2304 -122.6136
767869 Ruiz et al., 2015 2011 2011-09-13 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.8046 -122.3985
767898 Ruiz et al., 2015 2011 2011-09-27 Vallejo Marina, San Francisco Bay, CA, California, USA Def 38.1086 -122.2694
767944 Ruiz et al., 2015 2011 2011-09-28 Glen Cove Marina, San Francisco Bay, CA, California, USA Def 38.0663 -122.2130
767959 Ruiz et al., 2015 2011 2011-09-28 Benicia Marina, San Francisco Bay, CA, California, USA Def 38.0453 -122.1561
767966 Ruiz et al., 2015 2011 2011-09-12 Corinthian Yacht Club, San Francisco Bay, CA, California, USA Def 37.8103 -122.3228
768128 Ruiz et al., 2015 2012 2012-09-04 Redwood City Marina, San Francisco Bay, CA, California, USA Def 37.5023 -122.2130
768151 Ruiz et al., 2015 2012 2012-09-06 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9736 -122.4802
768209 Ruiz et al., 2015 2012 2012-08-31 Glen Cove Marina, San Francisco Bay, CA, California, USA Def 38.0663 -122.2130
768231 Ruiz et al., 2015 2012 2012-09-13 San Leandro Marina, San Francisco Bay, CA, California, USA Def 37.6962 -122.1919
768334 Ruiz et al., 2015 2013 2013-08-23 Loch Lomond Marina, San Francisco Bay, CA, California, USA Def 37.9723 -122.4829

References

Bastida-Zavala, J. Rolando; McCann, Linda D.; Keppel; Erica; Ruiz, Gregory M. (2017) The fouling serpulids (Polychaeta: Serpulidae) from United States coastal waters: an overview, European Journal of Taxonomy 344: 1-76

Bazterrica, Mar'a Cielo Casariego, Agustina Mendez Alvarez, Graciela Obenat, Sandra Baron, Pedro J. (2021) Reproductive traits of a nonindigenous amphipod associated with alternative habitat structures in presence of an invasive ecosystem-engineering polychaete, None 548: 5051–5066

Shalovenko, N. N. (2020) Tendencies of Invasion of Alien Zoobenthic Species into the Black Sea, Russian Journal of Biological Invasions 11(2): . 164–171

Aladin, Nikolai V.; Plotnikov, Igor S.; Filipov, Andrei A. (2002) Invasive aquatic species of Europe: Distribution, impacts, and management., Kluwer Academic Publishers, Dordrecht. Pp. 351-359

Allen, F. E. (1953) Distribution of marine invertebrates by ships, Australian Journal of Marine and Freshwater Research 4(2): 307-316

Alvarez-Aguilar, A.; Van Rensburg, H.; Simon, C. A. (2022) Impacts of alien polychaete species in marine ecosystems: a systematic review, Journal of the Marine Biological Association of the United Kingdom https://doi.org/10.1017/ S0025315422000315: Published online

Amor, Kounofi-Ben; Rifi, M.; Ghanem, R.; Draief, I.; Zouali, J.; Souissi, J. Ben (2016) Update of alien fauna and new records of Tunisian marine fauna, Mediterranean Marine Science 17(1): 124-143

Asakura, A. (1992) Recent introductions of marine benthos into Tokyo Bay (review): Process of invasion into an urban ecosystem with discussion on the factors inducing their successful introduction, Journal of the Natural History Museum and Institute, Chiba 2(1): 1-14

Associated Press (12/2021) Lummi Nation declares disaster after invasive crab arrives, Seattle Times <missing volume>: <missing location>

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

Barnes, Robert D. (1983) Invertebrate Zoology, Saunders, Philadelphia. Pp. 883

Bastida-Zavala, J. Rolando. (2008) Serpulids (Annelida: Polychaeta) from the Eastern Pacific, including a brief mention of Hawaiian serpulids., Zootaxa 1722: 1-61

Bazterrica, María Cielo; Bruschetti, Carlos Martín; Alvarez, María Fernanda; Iribarne, Oscar; Botto, Florencia (2014) Effects of macroalgae on the recruitment, growth, and body condition of an invasive reef forming polychaete in a south-western Atlantic coastal lagoon, Journal of Sea Research 88: 121-129

Bazterrica, Maria Cielo; Botto, Florencia; Iribarne, Oscar (2011) Effects of an invasive reef-building polychaete on the biomass and composition of estuarine macroalgal assemblages, Biological Invasions 14: 765-777

Ben-Eliahu, M. Nechama; ten Hove, Harry A. (2011) Serpulidae (Annelida: Polychaeta) from the Suez Canal: From a Lessepsian migration perspective (a monograph), Zootaxa 2848: 1-147

Bianchi, C. N.; Morri, C. (2001) The battle is not to the strong: Serpulid reefs in the lagoon of Orbetello (Tuscany: Italy)., Estuarine, Coastal and Shelf Science 53: 215:220

Bianchi, Carlo N.; Morri, Carla (1996) Ficopomatus 'reefs' in Po river delta (northern Adriatic): their constructional dynamics, biology, and influences on the brackish water biota., Pubblicazioni della Stazione Zoologica di Napoli I: Marine Ecology 17(1-3): 51-66

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

Blakeslee, April M. H.; Miller, A. Whitman; Ruiz, Gregory M.; Kerstin Johannesson · Carl André ·Johannsson, Kerstin;· André, Carl; Panova, Marina (2021) Population structure and phylogeography of two North Atlantic Littorina species with contrasting larval development, Marine Biology 168: <missing location>

Bruschetti, Martin; Luppi, Tomas; Fanjul, Eugenia; Rosenthal, Alan; Iribarne, Oscar (2008) Grazing effect of the invasive reef-forming polychaete Ficopomatus enigmaticus (Fauvel) on phytoplankton biomass in a SW Atlantic coastal lagoon., Journal of Experimental Marine Biology and Ecology 354: 212-219

Bruschetti, Martín; Bazterrica, Cielo; Luppi, Tomás; Iribarne, Oscar (2009) An invasive intertidal reef-forming polychaete affect habitat use and feeding behavior of migratory and local birds in a SW Atlantic coastal lagoon, Journal of Experimental Marine Biology and Ecology 375: 76-83

Bruschetti, Martin; Bazterrica, Cielo; Fanjul, Eugenia; Luppi, Tomas; Iribarne, Oscar (2011) Effect of biodeposition of an invasive polychaete on organic matter content and productivity of the sediment in a coastal lagoon, Journal of Sea Research 66: 20-28

California Department of Fish and Wildlife (2014) Introduced Aquatic Species in California Bays and Harbors, 2011 Survey, California Department of Fish and Wildlife, Sacramento CA. Pp. 1-36

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. (Ed.) (2007) <missing title>, University of California Press, Berkeley. Pp. <missing location>

Carlton, James T.; Eldredge, Lucius (2009) Marine bioinvasions of Hawaii: The introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago., Bishop Museum Bulletin in Cultural and Environmental Studies 4: 1-202

Chainho, Paula and 20 additional authors (2015) Non-indigenous species in Portuguese coastal areas, lagoons, estuaries, and islands, Estuarine, Coastal and Shelf Science <missing volume>: <missing location>

Chavanich, S.; Tan, L. T.; Vallejo, B.; Viyakarn, V. (2010) Report on the current status of marine non-indigenous species in the Western Pacific region, Intergovernmental Oceanographic Commission, Subcommission for the Western Pacific, Bangkok, Thailand. Pp. 1-61

Çinar, M. E.; Noglu, M. Bilece; Özturk, B.; Katagan, T. ; Aysel, V. (2005) Alien species on the coasts of Turkey, Mediterranean Marine Science 6/2: 119-146

Çinar, Melih Ertan (2006) Serpulid species (Polychaeta: Serpulidae) from the Levantine coast of Turkey (eastern Mediterranean), with special emphasis on alien species., Aquatic Invasions 1(4): 223-240

Çinar, Melih Ertan (2013) Alien polychaete species worldwide: current status and their impacts, Journal of the Marine Biological Association of the United Kingdom 93(5): 1257-1278

Cinar, Melih Ertan and 7 authors (2021) Current status (as of end of 2020) of marine alien species in Turkey, PLOS ONE 16: Published online

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; Bilecenoglu, Murat; Öztürk, Bilal; Can, Alp (2006) New records of alien species on the Levantine coast of Turkey, Aquatic Invasions 1(2): 84-90

2005 Exotics Guide. San Francisco Estuary Institute, Oakland, CA, www.exoticsguide.org

Cohen, Andrew N. and 10 authors (2005) <missing title>, San Francisco Estuary Institute, Oakland CA. Pp. <missing location>

Cohen, Andrew N. and 12 authors (2002) Project report for the Southern California exotics expedition 2000: a rapid assessment survey of exotic species in sheltered coastal waters., In: (Eds.) . , Sacramento CA. Pp. 1-23

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>

Coles S. L., DeFelice R. C., Eldredge, L. G. (1999a) Nonindigenous marine species introductions in the harbors of the south and west shores of Oahu, Hawaii., Bishop Museum Technical Report 15: 1-212

Coles, S. L.; DeFelice, R. C.; Eldredge, L. G.; Carlton, J. T. (1999b) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands., Marine Biology 135(1): 147-158

Cranfield, H.J.; Gordon, D.P.; Willan, R.C.; Marshall, B.A; Battershill, C.N.; Francis, M.P.; Nelson, W.A.; Glasby, C.J.; Read, G.B. (1998) <missing title>, The National Institute of Water and Atmospheric Research, New Zealand. Pp. <missing location>

Cukrov, Marijana; Despalatovic, Marija; Žuljevic, Ante; Cukrov, Neven (2010) First record of the introduced fouling tubeworm Ficopomatus enigmaticus (Fauvel, 1923) in the eastern Adriatic Sea, Croatia, Rapportes de la Commision Internationale de la Mer Mediterranee 39: 483

DAISIE (Delivering Alien Invasive Species Inventories to Europe) (2009) Handbook of alien species in Europe, Springer, Dordrecht, Netherlands. Pp. 269-374

Davies, B. R.; Stuart, V.; Villiers, M. de (1989) The filtration activity of a serpuid polychaete population (Ficopomatus enigmaticus) and its effects on water quality in a coastal marina., Estuarine Coastal and Shelf Science 29: 613-620

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

Despalatovic, Marija; Cukrov, Marijana; Cvitkovic, Ivan; Cukrov, Neven; ŽuljeviC, Ante (2013) Occurrence of non-indigenous invasive bivalve Arcuatula senhousia in aggregations of non-indigenous invasive polychaete Ficopomatus enigmaticus in Neretva River Delta on the Eastern Adriatic coast, Acta Adriatica 54(2): 213-220

Dixon, D.R. (1981) Reproductive biology of the serpulid Ficopomatus (Mercierella) enigmaticus in the Thames Estuary, S.E. England, Journal of the Marine Biological Association of the United Kingdom 61: 805-815

Elton, Charles S. (1958) <missing title>, Methuen & Co. Ltd., London. Pp. <missing location>

Eno, N. Clare (1996) Non-native marine species in British waters: effects and controls, Aquatic Conservation: Marine and Freshwater Ecosystems 6: 215-228

Eno, N. Clare; Clark, Robin A.; Sanderson, William G. (1997) <missing title>, Joint Nature Conservation Committee, Peterborough. Pp. <missing location>

1997-2012 Directory of Non-Native Marine Species in British waters. http://jncc.defra.gov.uk/page-1532

Etchegoin, J. A.; Merlo, M. J.; Parietti, M. (2012) The role of the invasive polychaete Ficopomatus enigmaticus (Fauvel, 1923) (Serpulidae) as facilitator of parasite transmission in Mar Chiquita coastal lagoon (Buenos Aires, Argentina), Parasitology 139: 1506-1512

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

Fornos, J.J.; Forteza, V.; Martinez-Taberner, A. (1997) Modern polychaete reefs in Western Mediterranean lagoons: Ficopomatus enigmaticus (Fauvel) in the Albufera of Menorca, Balearic Islands, Palaeogeography, Palaeoclimatology, Palaeoecology 128: 175-186

Foss, Stephen (2009) <missing title>, California Department of Fish and Game, Sacramento CA. Pp. <missing location>

Foss, Stephen (2011) <missing title>, California Department of Fish and Game, Office of Spill Prevention and Response, Sacramento. Pp. 54

Galil, Bella S. (2007) Seeing Red: Alien species along the Mediterranean coast of Israel., Aquatic Invasions 2(4): 281-312

Ghobashy, Abdel Fattah A.; Ghobashy, Mahi A. F. (2005) Marine fouling studies in Egypt: A- Serpulids, Egyptian Journal of Aquatic Research 31(2): 89-102

Gittenberger, Adriaan; Rensing, Marjolein; Stegenga, Herre; Hoeksema, Bert (2010) Native and non-native species of hard substrata in the Dutch Wadden Sea, Nederlandse Faunistiche Mededelingen 33: 20-76

Gollasch, Stephan; Nehring, Stefan (2006) National checklist for aquatic alien species in Germany., Aquatic Invasions 1(2): 245-269

Gomiou, Marian-Traian; Alexandrov, Boris; Shadrin, Nikolai; Zaitsev, Yuvenaly (2002) The Black Sea- a recipient, donor, and transit area for alien species., In: Leppakoski, E.; Gollasch, S.; Olenin, S.(Eds.) Invasive aquatic species of Europe: Distribution, impacts, and management.. , Dordrecht. Pp. 341-350

Goulletquer, Philippe; Bachelet, Guy; Sauriau, Pierre; Noel, Pierre (2002) Invasive aquatic species of Europe: Distribution, impacts, and management., Kluwer Academic Publishers, Dordrecht. Pp. 276-290

Green, Stephanie J. and 7 authors (2021) Broad-scale acoustic telemetry reveals long-distance movements and large home ranges for invasive lionfish on Atlantic coral reefs, Marine Ecology Progress Series 673: 117-134

Grigorovich, Igor A.; Therriault, Thomas W.; MacIsaac, Hugh J. (2003) History of aquatic invertebrate invasions in the Caspian Sea., Biological Invasions 5: 103-115

Grosse, Maël; Pérez, Roberto; Juan-Amengual, Mateo; Pons; Joan; Capa, María (2021) The elephant in the room: first record of invasive gregarious species of serpulids (calcareous tube annelids) in Majorca (western Mediterranean), Scientia Marina 85(1): 15-28

Hartman, O. (1951) The littoral marine annelids of the Gulf of Mexico, Publications of the Institute of Marine Science 2(1): 7-124

Hartman, Olga (1952) Fouling serpulid worms, new to the Gulf of Mexico., Texas Journal of Science 4(1): 63-64

Hayward, P.J.; Ryland, J. S. (1990) <missing title>, 1 Clarendon Press, Oxford. Pp. <missing location>

Heiman, Kimberly W.; Micheli, Fiorenza (2010) Non-native ecosystem engineer alters estuarine communities, Integrative and Comparative Biology 50(2): 226-236

Heiman, Kimberly W.; Vidargas, Nicholas; Micheli, Fiorenza (2008) Non-native habitat as home for non-native species: Comparison of communities associated with invasive tubeworm and native oyster reefs., Aquatic Biology 2: 47-56

Hewitt, C.L.; Campbell, M.L.; Thresher, R.E.; Martin, R.B. (1999) Marine Biological Invasions of Port Phillip Bay, Victoria, In: (Eds.) . , Hobart, Tasmania. Pp. <missing location>

Hewitt, Chad L. and 14 authors. (2004) Introduced and cryptogenic species in Port Phillip Bay, Victoria, Australia., Marine Biology 144: 183-202

Hille, Sven; Kunz, Friederike; Markfort, Greta; Ritzenhofen, Lukas; Zettler, Michael L. (2021) First record of mass occurrence of the tubeworm Ficopomatus enigmaticus (Fauvel, 1923) (Serpulidae: Polychaeta) in coastal waters of the Baltic Sea, Bioinvasions Records 10: In press

Hoagland, K. E.; Turner, R. D. (1980) Range extensions of teredinids (shipworms) and polychaetes in the vicinity of a temperate-zone nuclear generating station., Marine Biology 58(1): 55-64

Holmes, Samuel; Callaway, Ruth (2021) Fouling communities and non-native species within five ports along the Bristol Channel, South Wales, UK, Estuarine Coastal and Shelf Science 252(1107295): Publiished online

ICES Advisory Committee on the Marine Environment (2012) <missing title>, International Council for the Exploration of the Seas, Copenhagen. Pp. <missing location>

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

Iwasaki, Keiji (2006) Assessment and Control of Biological Invasion Risks., Shoukadoh Book Sellers,and IUCN, Kyoto and Gland, Switzerland. Pp. 104-112

Jensen, Kathe R.; Knudsen, Jorgen (2005) A summary of alien marine benthic invertebrates in Danish waters., Oceanological and Hydrobiological Studies 34 (suppl. 1): 137-161

Keene, William C. Jr. (1980) The importance of a reef-forming polychaete, Mercierella enigmatica in the oxygen and nutrient dynamics of a hypereutrophic subtropical lagoon., Estuarine and Coastal Marine Science 11: 167-178

Keough, M. J. ; Ross, J. (1999) Introduced fouling species in Port Phillip Bay., In: Hewitt, C. L.; Campbell; M.;Thresher, R.; Martin,(Eds.) Marine Biological Invasions of Port Phillip Bay, Victoria. , Hobart, Tasmania. Pp. 193-225

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

Kim, Tae Won; Micheli, Fiorenza (2013) Decreased solar radiation and increased temperature combine to facilitate fouling by marine non-indigenous species, Biofouling 29(5): 501-512

Lavigne, Andrea S. Sunesen, Ines Sar, Eugenia A. (2015) Morphological, taxonomic and nomenclatural analysis of species of Odontella, Trieres and Zygoceros (Triceratiaceae, Bacillariophyta) from Anegada Bay (Province of Buenos Aires, Argentina), Diatom Research 30: 307-331

LeBlanc, Francis; Steeves, Royce; Irlich, Ulrike; Bourque, Daniel; Akaishi, Fabiola; Gagné, Nellie (2021) Scoping the distribution of Smallmouth Bass (Micropterus dolomieu) in the Miramichi River Watershed in 2019 and 2020 using environmental DNA, Canadian Manuscript Report of Fisheries and Aquatic Sciences 3222: 1-22

Leppäkoski Erkki, Mihnea, Pia E. (1996) Enclosed seas under man-induced change: a comparsion between the Baltic and Black Seas., Ambio 25(6): 380-389

Li, Xinzhengl Shenz. Anglv; Xu, zhaou (2008) Report on some shrimos from dontou Island, Zhejiang, China (Decapoda, Natantia, Crustaceana (Leiden) <missing volume>: <missing location>

Lipej, L.; Mavric, B.; Orlando-Bonaca, M.; Malej, A. (2012) State of the art of the marine non-indigenous flora and fauna in Slovenia, Mediterranean Marine Science 13(2): 243-249

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

Loveland, Robert E.; Shafto, Sylvia S. (1984) Fouling Organisms, In: Kennish, Michael J., and Lutz, Richard A.(Eds.) Ecology of Barnegat Bay, New Jersey.. , Berlin. Pp. 226-20

Lutaenko, Konstantin A.; Furota,Toshio; Nakayama, Satoko; Shin, Kyoungsoon; Xu, Jing (2013) <missing title>, Northwest Pacific Action Plan- Data and Information Network Regional Activity Center, Beijing, China. Pp. <missing location>

Mathieson, Arthur C.; Dawes, Clinton J. (2017) Seaweeds of the Northwest Atlantic, University of Massachusetts Press, Amherst MA. Pp. <missing location>

McQuaid, K. A.; Griffiths, C. L. (2014) Alien reef-building polychaete drives long-term changes in invertebrate biomass and diversity in a small, urban estuary, Estuarine, Coastal and Shelf Science 138: 101-106

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

Mineur, Frederic and 5 authors (2012) Changing Coasts: Marine aliens and artificial structures, Oceanography and Marine Biology, an Annual Review 50: 189-234

Miranda, Nelson A. F.; Kupriyanova, Elena K.; Rishworth, Gavin M.; Peer, Nasreen; Bornman, Thomas G.; Bird, Matthew S.; Perissinotto, Renzo (2016) An invasive polychaete species found in living marine stromatolites, Aquaitc Invasions 11: Published online

Munari, C.; Bocchi, N.; Mistri, M. (2015) Epifauna associated to the introduced Gracilaria vermiculophylla (Rhodophyta: Floridophyceae; Graciliariales) and comparison with the native Ulva rigida in an Adriatic lagoon, Italian Journal of Zoology 82(3): 436-445

Muniz, Pablo; Clemente, Juan; Brugnoli, Ernesto (2005) Benthic invasive pests in Uruguay: a new problem or an old one recently perceived?, Marine Pollution Bulletin 50: 1014-1018

Nelson-Smith, Anthony (1971) Annelids as fouling organisms., In: Gareth Jones, E. B. and Eltringham, S. K.(Eds.) Marine Borers, Fungi, and Fouling Organisms of Wood.. , Paris. Pp. 171-184

Norris, James N. (2010) Marine Algae of the northern Gulf of California: Chlorophyta and Phaeophyceae, Smithsonian Contributions to Botany 94: 1276

Obenat, S. M.; Pezzani, S. E. (1994) Life cycle and population structure of the polychaete Ficopomatus enigmaticus in Mar Chiquita coastal lagoon, Argentina, Estuaries 17(1B): 263-270

2000-2016 Inventory of Baltic Sea alien species. http://www.ku.lt/nemo/species.htm

Oliva, Matteo; Mennillo, Elvira; Barbaglia, Martina; Monni, Gianfranca Tardelli, Federica; Casu, Valentina; Pretti, Carlo (2015) The serpulid Ficopomatus enigmaticus (Fauvel,1923) as candidate organisms for ecotoxicological assays in brackish and marine waters, Ecotoxicology and Environmental Safety <missing volume>: <missing location>

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

Pan, Jerónimo; Marcoval, M. Alejandra (2013) Top-down effects of an exotic serpulid polychaete on natural plankton assemblage of estuarine and brackish systems in the SW Atlantic, Journal of Coastal Research 30(6): 1226-1235

Pancuci-Papadopoulou., M. A.; Zenetos, A , Corsini-Foka; Politou, Ch. (2005) Update of marine alien species in Hellenic waters., Mediterranean Marine Science 6(2): 1-11

Partaly, Ye. M. (2003) Peculiarities of successions in biocenoses of fouling in the Sea of Azov, Hydrobiological Bulletin 39(1): 103-112

Pernet, Bruno; Barton, Michelle; Fitzhugh, Kirk; Harris, Leslie H.; Lizárraga, David; Ohl, Ryan; Whitcraft, Christine R. (2016) Establishment of the reef-forming tubeworm Ficopomatus enigmaticus (Fauvel, 1923) (Annelida: Serpulidae) in southern California, BioInvasions Records 5: In press

Preda, Cristina; Memedemin, Daniyar; Skolka, Marius; lniceanu, Dan Coga (2012) Early detection of potentially invasive invertebrate species in Mytilus galloprovincialis Lamarck, 1819 dominated communities in harbours, Helgoland Marine Research 66: 545-556

Raymont, J. E. G. (1976) Harvesting Polluted Waters, Plenum, New York. Pp. 185-199

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.; Gerlinger, Thomas V.; Ware, Robert R. (2018) Comparison of the polychaetous annelids populations on suspended test panels in Los Angeles Harbor in 1950-1951 with the populations in 2013-2014, Bulletin of the Southern California Academy of Sciences 17(1): 82-90

Rioja, E. (1943) Estudios Anelidológicos IX: La presencia de la Mercierella enigmatica Fauvel, en las costas argentinas, Anales del Instituto de Biología de la Universidad Nacional Autónoma de México 14: 547-551

Ruiz, G.M.; Fofonoff, P. W.; Carlton, J. T.; Wonham, M. J.; Hines, A. H. (2000) Invasion of coastal marine communities in North America: Apparent patterns, processes, and biases., Annual Review of Ecology and Systematics 31: 481-531

Ruiz, Gregory M., Fofonoff, Paul, Hines, Anson H. (1999) Non-indigenous species as stressors in estuarine and marine communities: assessing invasion impacts and interactions, Limnology and Oceanography 44(3, part 2): 950-972

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

Schwindt, Evangel; Bortolus, Alejandro; Iribarne, Oscar Osvaldo (2001) Invasion of a reef-builder polychaete: direct and indirect impacts on the native benthic community structure, Biological Invasions 3: 137-149

2000 List of exotic introduced species in the Black Sea. http://ibss.iuf.net/people/shadrin/introduc.html

Skolka, Marius; Preda, Cristina (2010) Alien invasive species at the Romanian Black Sea coast: present and perspectives, Travaux du Muséum National d’Histoire Naturelle «Grigore Antipa» 53: 443-467

Straughan, Dale (1969) Intertidal zone-formation in Pomatoleios kraussi(Annelida: Polychaeta), Biological Bulletin 136: 469-482

Straughan, Dale (1969) Serpulidae (Annelida: Polychaeta) from Oahu, Hawaii., Bulletin of the Southern California Academy of Sciences 68(4): 229-240

Tarakanadha, B.; Morrell, Jeffrey J.; Rao, K. Satyanarayana (2004) Environmental impacts of preservative-treated wood., Florida Center for Environmental Solutions, Orlando. Pp. 320-335

Ten Hove, H. A.; Weerdenburg, J. C. A. (1978) A generic revision of the brackish-water serpulid Ficopomatus Southern 1921 (Polychaeta : Serpulinae) including Mercierella Fauvel 1923, Sphaeropomatus Treadwell 1934, Mercierellopsis Rioja 1945 and Neopomatus Pillai 196, Biological Bulletin 154: 96-120

ten Hove, Harry (1974) Notes on Hydroides elegans (Haswell, 1883) and Mercieriella enigmatica Fauvel 1923, alien serpulid polychaetes introduced into the Netherlands., Bulletin Zoologisch Museum, Universiteit van Amsterdam 4(6): 45-51

Thomas, Nigel S.; Thorp, Clifford H. (1994) Cyclical changes in the fauna associated with tube aggregates of Ficopomatus enigmaticus (Fauvel), Memoires du Museum Nationale d'Histoire Naturelle 162: 575-584

2002-2021 Invertebrate Zoology Collections Database. <missing description>

2003-2015 Nonindigenous Aquatic Species Database. Gainesville, FL. http://nas.er.usgs.gov

Vuillemin, Simone (1965) Contribution a l'etude ecologique du lac de Tunis: Biologie de Mercierella enigmatica Fauvel, Theses a la Faculte des sciences de lUniversite de Paris A4622(5469): 1-276

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

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

Zaitsev, Yuvenali; Ozturk, Bayram (2001) <missing title>, Turkish Marine Research Foundation Publication, <missing place>. Pp. 1-265

Zibrowius, Helmut (1994) Introduced Species in European Coastal Waters, European Commission, Brussels. Pp. 44-65

Zibrowius, Helmut, Thorp, Clifford H. (1989) A review of the alien serpulid and spirorbid polychaetes in the British Isles, Cahiers de Biologie Marine 30(3): 271-285