Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide


Ficopomatus enigmaticus



Ficopomatus enigmaticus (tubeworm)


  • Last modified
  • 25 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Ficopomatus enigmaticus
  • Preferred Common Name
  • tubeworm
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Annelida
  •       Class: Polychaeta
  •         Order: Sabellida
  • Summary of Invasiveness
  • F. enigmaticus is an invasive, ecosystem engineering, brackish-water serpulid polychaete that builds calcareous aggregates in estuarine and coastal environments within subtropical/temperate areas throughout the w...

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Most of the reef is made out of calcareous tubes of dead worms. The bottom part of a reef is gradually buried in anoxic black sediment, while the reef grows and the nucleus usually can be found right in the center. Virtually all living worms are around the reef forming a ring and/or on top of it.
CaptionMost of the reef is made out of calcareous tubes of dead worms. The bottom part of a reef is gradually buried in anoxic black sediment, while the reef grows and the nucleus usually can be found right in the center. Virtually all living worms are around the reef forming a ring and/or on top of it.
CopyrightEvangelina Schwindt
Most of the reef is made out of calcareous tubes of dead worms. The bottom part of a reef is gradually buried in anoxic black sediment, while the reef grows and the nucleus usually can be found right in the center. Virtually all living worms are around the reef forming a ring and/or on top of it.
StructureMost of the reef is made out of calcareous tubes of dead worms. The bottom part of a reef is gradually buried in anoxic black sediment, while the reef grows and the nucleus usually can be found right in the center. Virtually all living worms are around the reef forming a ring and/or on top of it.Evangelina Schwindt


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Preferred Scientific Name

  • Ficopomatus enigmaticus (Fauvel, 1923)

Preferred Common Name

  • tubeworm

Other Scientific Names

  • Mercierella enigmatica Fauvel, 1923

International Common Names

  • English: Australian tubeworm; fanworm; reef building tubeworm
  • Spanish: gusano tubícola

Local Common Names

  • Denmark: australsk kalkrørsorm
  • Germany: Tüten-Kalkröhrenwurm
  • Italy: corallina; peoci de fosso; spreo; tubicelli
  • Netherlands: trompetkalkkokerworm
  • Sweden: australisk kalkrörsmask

Summary of Invasiveness

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F. enigmaticus is an invasive, ecosystem engineering, brackish-water serpulid polychaete that builds calcareous aggregates in estuarine and coastal environments within subtropical/temperate areas throughout the world. It was described for the first time as exotic in France in 1923 and soon after that it was recorded in other several European countries, Australia, USA, Uruguay, Australia and Tunisia; the last records of this species were in Ireland and in Croatia in 2006. The native area of origin is unknown. The hypothesis of an Australasian origin is currently the most accepted but most scientists agree that it still needs further evaluation. The most important dispersion worldwide is likely to occur via hull fouling and/or in ballast water in large vessels. For Europe, is listed as one of the 100 worst invasive species (DAISIE, 2009) but so far, there are no unified strategies for its control or management. This species has a fast growth rate, high tolerance to variable environmental conditions, and it is causing important ecological impacts in several regions by modifying the ecological and the physical processes of the ecosystems. In some locations, economic impacts occur due to the prolific growth that can cause blocking of thermal effluents and fouling of aquaculture ponds and leisure crafts.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Annelida
  •             Class: Polychaeta
  •                 Order: Sabellida
  •                     Family: Serpulidae
  •                         Genus: Ficopomatus
  •                             Species: Ficopomatus enigmaticus

Notes on Taxonomy and Nomenclature

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Ficopomatus enigmaticus is a reef-building polychaete that belongs to the recently reviewed Serpulidae family (Kupriyanova et al., 2001; ten Hove and Kupriyanova, 2009). The first name of this species was Mercierella enigmatica until 1978 when ten Hove and Weerdenburg (1978) revised in detail the monotypic brackish-water genera Mercierella, Sphaeropomatus, Mercierellopsis and Neopomatus and unified them under Ficopomatus. Five species are now recognized within this genus: Ficopomatus enigmaticus, Ficopomatus macrodon (type species), Ficopomatus miamiensis, Ficopomatus uschakovi and Ficopomatus talehsapensis, all species except F. enigmaticus live in tropical waters (ten Hove and Kupriyanova, 2009).


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The genus Ficopomatus is characterized by a symmetrical body, operculum and collar chaetae (coarsely serrated and simple) present, opercular peduncle without well developed distal wings and pseudoperculum (rudimentary operculum) absent (ten Hove and Kupriyanova, 2009).

The type species for the genus is F. macrodon (see Notes on Taxonomy and Nomenclature) and the complete description of this species is given by ten Hove and Kupriyanova (2009). Ten Hove and Weerdenburg (1978) made the revision of this genus and they described in detail the morphological characteristics of F. enigmaticus. The body length without the tube is approximately 44 mm; however, this measure is highly variable among sites. Individual lifespan is very variable, Obenat and Pezzani (1994) suggest up to 24 months in the Mar Chiquita coastal lagoon, ten Hove (1979) indicated a 4 to 8 years, and Fox (1963) mentioned 12 years for individuals maintained in an aquarium.
The life cycle of F. enigmaticus follows the general pattern of the Serpulidae. The unfertilized oocytes are cup shaped to irregular (Fischer-Piette, 1937; Vuillemin, 1965). The morphology of the larvae in this species is similar to other Serpulidae species. After the fertilization, the negatively buoyant small eggs (60 µm) sink to the bottom where they undergo cleavage up to the blastula stage. Blastula is ciliated, mobile and develops into a larva with a prototroch consisting of a single ring of cilia. This stage undergoes significant morphological changes until the trochophore stage is reached. The trochophore continues to grow into the metatrochophore and finally to the larva ready to metamorphose to approximately 160 µm in size (Kupriyanova et al., 2001). The development time of larvae is approximately 20-25 days. The survival of the larvae in the water although has not been well determined, it appears to be very variable among habitats, probably due to the differences in environmental conditions. Fischer-Piette (1937) and Vuillemin (1965) estimated in 1-5 weeks, whereas Dixon (1981) indicated that it falls within the range of 1 to 3.5 months. Larvae act as the dispersal phase but the dispersion is mainly driven by currents since their own swimming speed generally does not exceed 5 mm/s (ten Hove, 1979). Then, larvae settle on hard substrates and grow as sedentary suspension feeder worms. The growth rate of the tubes varied among sites where it was measured. In Italy, tubes grew 0.4 mm per day (Bianchi and Morri, 1996) whereas the maximum was reported by Hartman-Schröder (1967) of 1.8 mm per day. For more details on the development see Kupriyanova et al. (2001).


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F. enigmaticus is a brackish water species distributed in subtropical-temperate areas worldwide, in both Hemispheres. The first hypothesis of the area of origin for this species was proposed by Fauvel (1933) who noticed that the species was introduced in France, but because it was very abundant in Madras, India, that region according to him must be the native area. Allen (1953) mentioned that by 1932 F. enigmaticus was very abundant and widely spread throughout the southern coast of Australia, and therefore Australia, and not India, must be the area of origin of this species. However, by 1971 Hartmann-Schröder (1971) clarified in part the taxonomic confusion involving this species and found that the records from Madras, some parts of Australia, Nigeria and other tropical areas were of F. uschakovi, not to F. enigmaticus. After the taxonomic revision of ten Hove and Weerdenburg (1978) it was clear that two species of Ficopomatus are present in Australia, F. enigmaticus and F. uschakovi, the former distributed in subtropical/temperate areas and the latter in tropical areas (i.e., the border around Sydney) (ten Hove and Weerdenburg, 1978). It is often assumed that the area with the highest number of representatives of a certain taxon is likely to be its area of origin. Since the Indo-Pacific region concentrates the highest number of Ficopomatus species, thus this region is likely to be the origin of the species (see further discussion on ths hypothesis in Zibrowius 1978). However, unfortunately there is no conclusive evidence to support this hypothesis.

In addition to the Distribution table, there are records in the literature of F. enigmaticus in different areas but unfortunately they do not offer much specific detail. For example, this species was mentioned by Arvanitidis (2000) in the Aegean Sea without further detail, although the author seems to refer to the coast of Greece. Several sources mention the species in the Azov Sea (e.g. Zibrowius, 2002; Surugiu, 2005) but the localities are not specified.
Ben-Eliahu and Dafni (1979) mentioned the local extinction of F. enigmaticus on piers in Israeli Mediterranean estuaries such as in the Alexander River. In 1964 the salinity was monitored in this river and it ranged from 0.5 to 200/00, typically from 2 to 90/00. According to the authors, the Ficopomatuspopulations in the estuaries died, presumably due to pollution of the rivers. However, there was a small population flourishing in an artificial pool, near the Acre (Akko) shore, encrusting on ageing lumber. When the factory stopped operating, the pool dried up along with its biota of brackish species sensitive to industrial pollution. Apparently the species did not disappear since it was mentioned by Galil (2007) as an alien species in the same river (Swamp near Akko, Alexander River). Probably it was not an extinction of Ficopomatus but a reduction in its population density due to variation in the environmental conditions, as also happened in Emsworth, England (Thorp, 1994).
Although F. enigmaticus has not been recorded in Norway (Gollasch et al., 2009), this country may be vulnerable to an invasion in the near future due to climate change. Gjershaug et al. (2009) hypothesized that if sea water temperature increased by 1ºC then F. enigmaticus would reach the southern part of Norway.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Sea Areas

Atlantic, Eastern CentralLocalisedIntroducedHove and Weerdenburg, 1978Morocco
Atlantic, NortheastWidespreadIntroduced Invasive Hove and Weerdenburg, 1978UK, Ireland, Portugal, Spain, France, Belgium, Denmark, Germany, Netherlands
Atlantic, SoutheastPresentIntroduced Not invasive Hove and Weerdenburg, 1978South Africa
Atlantic, SouthwestPresentIntroduced Invasive Hove and Weerdenburg, 1978Argentina, Uruguay
Atlantic, Western CentralLocalisedIntroduced Not invasive Hove and Weerdenburg, 1978USA
Indian Ocean, EasternWidespread Invasive Hove and Weerdenburg, 1978Australia
Mediterranean and Black SeaWidespreadIntroduced Invasive Hove and Weerdenburg, 1978Spain, France, Balearic Islands, Italy, Tunisia, Egypt, Slovenia, Ukraine, Greece, Israel, Turkey, Romania, Bulgaria, Georgia, Corsica, Croatia, Russia
Pacific, Eastern CentralPresentIntroduced Invasive Hove and Weerdenburg, 1978USA including Hawaii
Pacific, NorthwestWidespreadIntroduced Invasive Hove and Weerdenburg, 1978Japan
Pacific, SouthwestPresentIntroduced Invasive Read and Gordon, 1991New Zealand


AzerbaijanAbsent, formerly presentIntroduced Not invasive Khlebovich, 2009The species was present in ports and was not mentioned in the Caspian Sea during last years, but is not considered as disappeared
Georgia (Republic of)PresentIntroducedSurugiu, 2005Poti Harbour, Paleostomi Lake and rivers flowing into it
IsraelPresentIntroducedGalil, 2007Swamp near Akko, Alexander river, Mediterranean coast. First observed in 1954
JapanPresentIntroducedIwasaki et al., 2004Chubu, Chugoku, Kanto, Kinki, Tohoku
-KyushuPresentIntroducedIwasaki et al., 2004Hakata Bay, Fukuoka Prefecture, observed in the 1990s
-Ryukyu ArchipelagoPresentIntroducedIwasaki et al., 2004Ishigaki-jima, Kabira Bay, detected in the 1970s
TurkeyLocalisedIntroduced Not invasive Surugiu, 2005; Çinar et al., 2009; Çinar et al., 2009Golden Horn Estuary, Sea of Marmara. In low densities probably due to pollution. Present since 1952
TurkmenistanPresentIntroducedKhlebovich, 2009The species was present in ports and was not mentioned in the Caspian Sea during last years, but is not considered disappeared


EgyptLocalisedIntroduced Not invasive Samaan and Aleem, 1972Lake Mariut, the low abundance of the species is explained by low salinity
MoroccoLocalisedIntroducedRullier, 1966Near Abrenik
South AfricaWidespreadIntroduced Invasive Hove and Weerdenburg, 1978Several localities around the Cape Town and False Bay
TunisiaAbsent, formerly presentIntroduced Not invasive Rullier, 1966; Diawara et al., 2008; Ayari et al., 2009Tunis Lagoon. This site was subject of many studies about Ficopomatus, however, a recent survey made during 2003 and 2004 by the authors mentioned the disappearance of the species in this lagoon

North America

USAPresentPresent based on regional distribution.
-CaliforniaLocalisedIntroducedFauvel, 1933; Wasson et al., 2001; Cohen, 2005San Francisco Bay, first record of the species for the West Coast of USA
-FloridaPresentIntroduced Not invasive Fofonoff et al., 2003Banana River in Indian River Lagoon
-GeorgiaPresent, few occurrencesIntroducedUSGS, 2009One specimen was collected in an oyster clump in the Port of Brunswick in East River. Its record needs confirmation
-HawaiiPresentIntroducedHove and Weerdenburg, 1978Ala Wai Canal, near Waikiki
-MarylandPresentIntroduced Invasive Ruiz et al., 2000Baltimore and the Severn River, Chesapeake Bay, observed for the first time in 1994
-New JerseyLocalisedIntroducedHoagland and Turner, 1980Oyster Creek, Barnegat Bay
-TexasPresentIntroduced Invasive Hartman, 1952Corpus Christi Bay
-VirginiaPresentIntroduced Not invasive Fofonoff et al., 2003Scott Creek and Norfolk Harbour

South America

ArgentinaPresentIntroduced Not invasive Rioja, 1943; Bastida, 1971; Orensanz and Estivariz, 1971Quequen Estuary, Buenos Aires Province
UruguayWidespreadIntroducedMonro, 1938a; Brugnoli et al., 2006Arroyo Las Brujas, first record of the species in South America


BelgiumPresentIntroduced Invasive Rullier, 1966Ostende, introduced before 1952
BulgariaPresentIntroducedSurugiu, 2005Varna Lake and Harbour, Balchik Bay, Mandrenckoto Lake
CroatiaWidespreadIntroducedWGITMO, 2007Recorded for the first time in the Bay of Sibenik (Middle Adriatic) in 2006. A possible vector of introduction in ships transporting stone from Middle Dalmatia
DenmarkPresentIntroduced Not invasive Rasmussen, 1958; Rullier, 1966Observed in 1953 in the Copenhagen harbour near power plant. Its presence should be confirmed
FrancePresentIntroducedFauvel, 1923; Rullier, 1966Caen Channel, Normandy. This is the first mention of the species worldwide
-CorsicaWidespreadIntroduced Invasive Zibrowius, 1978Gradugine pond, Cazarello, Biguglia pond
GermanyPresentIntroduced Invasive Wolff, 2005Harbour of Emden near a power plant
GreecePresentIntroducedSimboura and Zenetos, 2002; Antoniadou and Chintiroglou, 2005Mesolongi Lagoon
IrelandLocalisedIntroduced Not invasive Kilty and Guiry, 1973; Minchin, 2007Cork Harbour
ItalyWidespreadIntroduced Invasive Bianchi, 1981aLagoons of Grado-Marano, Venezia, Delta del Po, Comacchio, Lessina-Varano, Burano, Comacchio (Adriatic Sea), Sabaudia, Fondi-Lungo, Fogliano-Monaci-Caprolace-Paola, Patria (Tyrrhenian Sea), Orbetello (Ligurian Sea), Faro-Ganzirri, Palermo (Sicily Island), and Cabras-Mistras-S. Giusta (Mediterranean Sea)
NetherlandsWidespreadIntroduced Invasive Wolff, 2005Observed in the Veerse Meer and the Kanaal door Walcheren in Binnenhaven at Vissingen from 1968 (date of first record) until 2000. Observed in Noordzeekanaal near Velzen in January 1992. Observed in Goesse Meer in 2000
PortugalPresentIntroducedHove and Weerdenburg, 1978Ria Formosa
RomaniaPresentIntroducedSurugiu, 2005Danube-Black Sea Canal, Agigea Harbour, Mangalia Bay, Constanta Harbour
Russian FederationPresentIntroducedSurugiu, 2005; DAISIE, 2009Gelendjik Bay in Black Sea. The author also mentions the species in the Azov Sea
SpainLocalisedIntroduced Invasive Rioja, 1924; Borja et al., 2006; Martínez and Adarraga, 2008Gandia, Valencia. This is the first record for Spain
-Balearic IslandsLocalisedIntroduced Not invasive Fornós et al., 1997Abufera of Menorca
UKWidespreadIntroduced Invasive Zibrowius and Thorp, 1989; Joyce et al., 2005Wales (Abereiddy, Dale, Milford Haven, Swansea and Cardiff). North England (Barrow-in Furness) and South-England (Portishead, Porlock, Falmouth, Plymouth, Weymouth, Cowes, Southampton, Portsmouth, Emsworth, Chichester, Shoreham, Brighton, Dover, Ramsgate, Greenhithe and St. Heleir)
UkrainePresentIntroducedAlexandrov et al., 2007Present in sea and inland saline waters since at least 1961.


AustraliaPresentThe native area of the species might be the Australasian area, thus the correct terminology would be listed as "cryptogenic species". Until this is corroborated here in the table is marked as an introduced species
-New South WalesPresentIntroducedHove and Weerdenburg, 1978Cooke's River, Sydney
-South AustraliaWidespreadIntroduced Invasive Hewitt et al., 2004Port Phillips Bay, reported for the first time in 1975
-VictoriaPresentIntroducedGeddes and Butler, 1984The Coorong Lagoons
-Western AustraliaPresentIntroduced Invasive Monro, 1938bSwan River, this is the first record for Australia but the author mentions that the species was already very abundant
New ZealandWidespreadIntroduced Invasive Read and Gordon, 1991; Probert, 1993Whangarei and Aukland Harbours, Otara Creek. This is the first record of this species in New Zealand but it was probably introduced around 1967/68

History of Introduction and Spread

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F. enigmaticus was first described as Mercierella enigmatica by Fauvel in 1923 in the Canal de Caen (France) and stated as an exotic species for that site where it was collected. Following this publication, F. enigmaticus is mentioned in more than 400 papers and databases in various field of research, and although many of the observations corresponded to other Ficopomatus species distributed in the tropics, this confusion was clarified by ten Hove and Weerdenburg (1978). The routes of dispersion of this species are unknown and further molecular studies would help to understand the areas of origin and the pathways of dispersion. During the first ten years after the first description of F. enigmaticus in France, this species was also suddenly observed North and South America, many countries of Europe, Tunisia and Australia which suggest that the species was not a new invader but undetected in these areas and probably introduced by ship fouling before and/or during the first world war, thus the route of dispersion is still an unsolved puzzle (Zibrowius, 1992). For example it was detected in the London docks (Monro, 1924), Spain (Rioja, 1924), the Black Sea, California (Fauvel, 1933), in coastal lagoons and creeks of Uruguay (Monro, 1938a) and Australia (Monro, 1938b). Rullier in 1966 summarized in detail observations of this species worldwide between 1921 and 1951. Although there are some errors due to the confusion of F. enigmaticus with other tropical Ficopomatus species, most references are correct and complete. After the work of Rullier, this species was detected in several countries including South Africa (Day, 1955), Japan (ten Hove and Weerdenburg, 1978) and New Zealand (Read and Gordon, 1991) and more recently, in 2006, it was found in Croatia (WGITMO, 2007).

Risk of Introduction

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F. enigmaticus is spread primarily by hull fouling and ballast water. Secondarily, since this polychaete is a successful fouling species, it can be potentially dispersed in association with mollusc species used for aquaculture purposes. All the introductions of this species have been reported as accidental.

The risk of spread of F. enigmaticus is highly related to the management of ballast water and hull fouling. It is not only large commercial vessels for interoceanic trips that should be considered as the target vectors, but also the small recreational vessel hulls and the cruise ships which are very important vectors on a regional scale. In fact, a first inoculation and subsequent regional translocations of larvae and/or adult individuals can be done by small rental fishing boats or canoes, since the worms can attach to the bottoms or on the cords, buoys and nets hanging around them, resisting relatively long periods of desiccation while travelling long distances by land from one tourist harbour to another. Thus, if the management is appropriated and effective, the risk of new introductions should be low, but unfortunately different countries are in different levels of development of control and prevention management of invasive species. In the USA there is a National Ballast Information Clearinghouse (2008)that collects, analyzes, and interprets data on the ballast water management practices of commercial ships that operate in the waters of the United States. In Argentina, for example, a recent work by Boltovskoy (2008) revealed that more than 50% of the ships arriving to the most important harbours ignore the Argentinean legislation about protected areas and ballast water management. In addition, the legislation is scarce and the information on ballast water and ship movements is not unified worldwide or even digitalized online. Because of these differences among countries, together with the fact that this species has a worldwide distribution, the risk of new introductions into new areas is high.
F. enigmaticus is listed as one of the 100 worst invasive species for Europe due to the impacts on biodiversity and the socio-economic effects (Streftaris and Zenetos, 2006; European Environmental Agency, 2007; DAISIE, 2009). However, considering the same variables, this species is not on the list of the 100 World's Worst Invasive Alien Species of the Global Invasive Species Database of the Global Invasive Species Programme (ISSG, 2009).
New introductions of F. enigmaticus into new regions are highly probable, especially in calm waters such as estuaries and harbours. This is because the species has a high growth rate, high fecundity and high larval retention in semi-closed and brackish areas. In addition, Bianchi and Morri (1996) proposed also that this species is able to change its life strategy from type R, with high recruitment rate, to a type K with rapid vertical growth during the annual cycle coincidently with changes in temperature and phytoplankton levels in the ecosystem. In addition, Eno et al. (1997) proposed that since this species lives in confined waters with variably salinity and temperature, competition with other serpulids would be low.


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F. enigmaticus is mainly a brackish species that tolerates highly variable salinity, dissolved oxygen and temperature. The typical habitats invaded by this species include estuaries, coastal lagoons, harbours, and inland brackish waters, always in protected wave areas. This species can survive either in polluted or non-polluted habitats. Çinar et al. (2009) found Ficopomatus in low densities in the Golden Horn Estuary, Sea of Marmara and the authors hypothesized that was due to the pollution. Naylor (1959) observed colonies of Ficopomatus in the dock polluted by waste oil in Swansea, England.

F. enigmaticus is commonly found in shallow waters between 0.5 and 2 m where this species has the opportunity to build the reefs. However, it was reported living in deeper waters in Netherlands at 9 m (Sluys et al., 2005) and in Greece at 40 m (Antoniadou and Chintiroglou, 2005), the latter probably being an extreme environmental condition.

Habitat List

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Estuaries Principal habitat Harmful (pest or invasive)
Inland saline areas Secondary/tolerated habitat Harmful (pest or invasive)
Lagoons Principal habitat Harmful (pest or invasive)
Inshore marine Secondary/tolerated habitat Harmful (pest or invasive)

Biology and Ecology

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The diploid number of chromosomes in this species is 26 and it was studied by Dasgupta and Austin (1960) in Swansea, Mildford, UK. Molecular data on F. enigmaticus can be found in Kupriyanova et al. (2006).
Reproductive Biology
F. enigmaticus has separate sexes but there is evidence of protandric hermaphrodism. True gonads are absent and the germ cells are produced by a germinal epithelium associated with genital blood vessels in the intersegmental septa. This species has external fertilization and spawning occurs through the specialized ducts in abdominal setigers of both males and females (Obenat et al., 2006b).
Temperature is one of the most important factors affecting the reproduction and fecundity in F. enigmaticus and is the most commonly studied variable. In general, development time increases with decreasing temperature (Kupriyanova et al., 2001). The minimum water temperature required for (or associated with) successful reproduction of F. enigmaticus differs among populations. In the Thames estuary (UK) it is about 18ºC (Dixon, 1981), whereas in the Emsworth lagoon (UK) and Tunis lagoon (Tunisia) it is 10ºC (Vuillemin, 1965; Thorp, 1995). In Mar Chiquita lagoon (Argentina) the proportion of sexually undifferentiated and gametogenic, but immature, worms was highest during the winter (June-August), when water temperature is below 16ºC (Obenat et al., 2006b). In this lagoon, the highest proportion of mature worms was registered when temperatures were above 16ºC (September to May); however, individuals at all stages were found all the year round (Obenat et al., 2006b). In Mar Chiquita coastal lagoon sexual maturity is reached at an age of approximately four months, and there are three oocyte generations each year (Obenat et al. 2006b), as observed in some other populations of F. enigmaticus (Vuillemin, 1965; Gambi et al., 2001). Sex ratio was observed to be male-biased throughout the year in Mar Chiquita coastal lagoon (Obenat and Pezzani, 1994). The fecundity of Ficopomatus is reported to vary between 1,000 and 10,000 (Kupriyanova et al., 2001). In Mar Chiquita coastal lagoon the maximum size of sexually undifferentiated worms was 26 mm. Sexually mature males ranged between 5 and 48 mm, and females between 8 mm and 51 mm (Obenat et al., 2006b). In Japan, mature eggs and sperm were first observed in individuals of 6-8 mm (Kupriyanova et al., 2001), whereas in France this species becomes mature at 9-10 mm (Fischer-Piette, 1937).
This species has two periods of spawning and recruitment in most regions where it was studied, one in spring-summer and the other one during the autumn. In Mar Chiquita coastal lagoon, recruitment occurred in November-December and in April-May (Obenat and Pezzani, 1994). Recruitment in southeastern England starts in June and continues through October (Dixon, 1981). Settlement peaks in North Adriatic (Italy) occur in June-July and in September (Bianchi and Morri, 1996) and in Japan occurs in May and October (Kupriyanova et al., 2001).
Bianchi and Morri (1996) also observed that growth and settlement are inversely related; months with heavy settlement showed reduced growth and vice-versa. The growth of the tubes was found to be variable among regions. Bianchi and Morri (1996) in Italy observed that tubes grew 30-35 mm in 90 days, Vuillemin (1965) in Tunisia recorded a tube growth of 54 mm in 108 days and Hartmann-Schröder (1967) in Germany reported 30 mm in 16 days. In some conditions with calm and shallow brackish waters Ficopomatus build reefs with different shapes including fringing reefs and microatolls (Bianch and Morri, 1996; Fornós et al., 1997), and circular shapes up to 7 m in diameter (Schwindt et al., 2004a). In these cases where the reefs are circular is particularly interesting to note that most of the reef structure is dead due to the accumulation of sediment trapped inside the reef, live individuals are found around the edges where larval settlement occurs between the calcareous tubes (Obenat and Pezzani, 1994). The bottom part of a reef is gradually buried in anoxic black sediment while the reef grows and the nucleus usually can be found right in the centre.
Little is known about the growth rate of the reefs. In Mar Chiquita coastal lagoon the growth rate was studied at different spatial and temporal scales. On a large scale, the density of reefs increased by 24% in 24 years (from 1975 and 1999), neighbouring reefs coalesced with each other forming platforms several metres (up to 12 m) long (E Schwindt, CENPAT-CONICET, Argentina, personal communication, 2009) and their density also increased in a 12.5% in the same period (Schwindt et al., 2004a). On a smaller scale, the growth rate of reefs of different sizes was measured during three years. Smaller reefs (average diameter of 0.5 m) increased their size by 24% while the larger reefs (3 m in diameter) only increased by 16% (Schwindt et al., 2004). In addition, the monthly growth rate was measured during a year resulting in a increasing of the reef size at 1.6 cm per month, this growth being higher in summer and lower in winter (Schwindt et al., 2004a)
Physiology and Phenology
F. enigmaticus can survive inside the tube without water for several hours but has no resistant stages. This species is highly successful since can tolerate a wide variability in environmental conditions including temperature, salinity, phytoplankton, water turbidity and pollution. In addition, Ficopomatus has a high growth rate and high fecundity and in enclosed waters the larval retention is high, consequently it would be considered as a potential dominant species occupying the settlement sites.
F. enigmaticus is a filter feeding organism that mainly feeds on phytoplankton and other detritus in suspension (Olivier et al., 1972).
F. enigmaticus is a gregarious polychaete that in many locations builds aggregates commonly called reefs. These reefs generate spaces among the tubes that are available to other epibenthic species. Although this species has no known symbionts, it is interesting to note that the same taxa, and in some cases the same invertebrate species, are usually associated to the Ficopomatus reefs worldwide. For example, the polychaete Spionidae Boccardiella ligerica is also a brackish water species that frequently lives among the tubes of Ficopomatus. Its distribution is very similar to Ficopomatus, as Western Europe, California, Argentina, Uruguay and South Africa (Blake, 1983). The amphipod Melita palmata is very abundant in Ficopomatus reefs of Argentina (Obenat et al., 2006a), England (Thorp, 1994) and Italy (Bianchi and Morri, 1994). Barnacles, crabs and amphipods are also part of the fauna associated to the reefs and in several cases the same species is found with the polychaete reefs in different regions. For example, the barnacle Balanus improvisus [Amphibalanus improvisus] is associated to the reefs in Argentina (Schwindt and Obenat, 2005), Italy (Bianchi and Morri, 1996; Sconfietti et al., 2003,) and the Netherlands (Sluys et al., 2005). The amphipod Monocorophium insidiosum is found in Argentina (Schwindt and Obenat, 2005), England (Thomas and Thorp, 1994), Italy (Bianchi and Morri, 1996), and the USA (Heiman et al., 2008). The polychaete Neanthessuccinea was observed in Argentina (Schwindt and Obenat, 2005) and Italy (Bianchi and Morri, 1996), and the bryozoan Conopeum seurati is very common covering the reefs tubes in England (Thorp, 1995) and Italy (Bianchi and Morri, 1996).
Environmental Requirements
There are several important environmental requirements for the survival and reproduction of Ficopomatus. Firstly, since the tubes have a low wave resistant (Bianchi and Morri, 2001), this species preferentially survives in calm waters such as creeks, channels, estuaries, inland waters and harbours, for example, in Mar Chiquita coastal lagoon, Argentina, where reefs are very successful covering the 86% of the lagoon surface the maximum-minimum current speed is 0.4 – 0.025 m/s-1 (Schwindt et al., 2004b). Occurrences on exposed shores, as at Porlock Weir, are extremely uncommon (Somerset, England) (Harris, 1970).
Secondly, the polychaetes require a hard substratum, or “nucleus”, for the settlement. A nucleus can be small stones, piers, ships, empty mollusc shells (Schwindt and Iribarne, 2000), submerged trees and macrophytes (Fornós et al., 1997) and any human trash such as cans, bottles, tyres, ropes, clothing, etc. (Schwindt and Iribarne, 2000).
Thirdly, temperature, salinity and phytoplankton levels are probably the main environmental factors affecting the reproduction of individuals in a given habitat. Changes in one of these parameters outside the tolerance range for a long period may cause the local extinction of the species. For example, Ficopomatus in the Tunis lagoon (Tunisia) was the subject of several early works (e.g. Heldt, 1944; Vuillemin, 1965; Keene, 1980). This lagoon was one of the most polluted and eutrophic environments of the Mediterranean coast due to the waste discharges that it received (Diawara et al., 2008). During the 1980s the restoration works cleaned, dredged and recovered the eutrophic waters, and as a result, re-established the lagoon communication with the sea and the salinity radically changed from brackish to marine. However, Ficopomatus is no longer present in the lagoon (Diawara et al., 2008; Tlig-Zouari and Maamouri-Mokhtar, 2008). In this case, Ficopomatus seemed to be more resistant to pollution than to a change in salinity. A similar result was obtained when the Peel-Harvey Estuary (Australia) was canalized to increase the tidal exchange with the sea through the Dawesville Channel. This estuary used to be an eutrophic environment and after the restoration works the polychaetes disappeared, without significant changes in temperature or dissolved oxygen; however, salinity increase significantly (Wildsmith et al., in press). Another example of the effect of the salinity on the polychaetes comes from the Mar Chiquita coastal lagoon. Results from a comparative field survey of brackish and seawater areas with an experimental reciprocal transplant of polychaetes between these two areas showed that the permanent contact with seawater affects the biomass and in consequence the growth. Polychaetes exposed to seawater had lower biomass and growth in comparison with polychaetes living in brackish water (Schwindt et al., 2004b). Temperature and phytoplankton are two important variables affecting the reproduction of Ficopomatus. Temperature needs to be higher than 15-18ºC to enhance reproductive success, in addition, Thorp (1994) has shown that even when minimum temperatures are reached the spawning may be delayed in the absence of adequate phytoplankton.
F. enigmaticus seems to be resistant to polluted waters, since it was recorded forming a very abundant population in the highly contaminated Tunis lagoon (Diawara et al., 2008; Tlig-Zouari and Maamouri-Mokhtar, 2008). It successfully lives in waters with high levels of industrial and domestic discharges in the Bilbao estuary (northern Spain) (Bustamante et al., 2007), and it is the main macrobenthic species in a hypereutrophic lagoon in the northwest Adriatic, Italy (Sorokin et al., 2004). However, Ficopomatus shows low densities in the Golden Horn Estuary, Sea of Marmara and this was believed due to be caused by pollution (Çinar et al., 2009). Naylor (1959) in Swansea, England observed colonies of Ficopomatus in the dock polluted by waste oil. It is not surprising that this species can resist polluted waters since one of the common habitats where it is found are harbours, which usually are highly polluted areas.



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As - Tropical savanna climate with dry summer Tolerated < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25]) Although this species is not from equatorial areas
C - Temperate/Mesothermal climate Preferred Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Preferred Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
55 39 0 0

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Ammonium [ionised] (mg/l) 0.0756 0.522 Optimum This parameter is not commonly obtained in ecological works. Data comes from two works in two different locations, France and Spain (Camus et al., 2000; Lucena-Moya et al., 2009)
Chlorophyll-a (mg/l) Optimum 1-167 tolerated, these values are from 5 works from 4 different worldwide locations
Depth (m b.s.l.) 0.1 2 Optimum 0.05-40 tolerated
Dissolved oxygen (mg/l) 6 8 Optimum 1-14 tolerated, data obtained from reviewing the available information of the literature of 11 countries including 29 ecosystems where this species invaded
Salinity (part per thousand) 10 30 Optimum 0.2-45 tolerated, data obtained from reviewing the available information of the literature of 11 countries including 29 ecosystems where this species invaded
Suspended solids (mg/l) 0.0086 3.68 Optimum 0.0086-3.68 tolerated, data comes from Schwindt et al. (2004b). The lowest value corresponds to area with full interchange of seawater
Turbidity (JTU turbidity) 2 24 Optimum Few papers reported this parameter and most of them used different methods. Data came from two works in two different areas invaded by the polychaete
Water pH (pH) 7 9 Optimum 4-10 tolerated. The tolerated pH values were obtained from laboratory measures (Bianchi, 1981b)
Water temperature (ºC temperature) 10 20 Optimum 0-30 tolerated. Data obtained from reviewing the available information of the literature of 12 countries including 30 ecosystems where this species invaded

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Anguilla anguilla Predator Adult to species Bianchi and Morri, 1996; Fornós et al., 1997; Froese R Pauly D, 2009
Carcinus maenas Predator Adult to species Carlton and Cohen, 2003; Hidalgo et al., 2005; Thorp, 1994
Gobiosoma parri Predator Adult to species Olivier et al., 1972
Gobius niger Predator Adult to species Bianchi and Morri, 1996; Froese R Pauly D, 2009

Notes on Natural Enemies

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Few natural enemies are known for F. enigmaticus and less known are the effects of these enemies on the polychaetes. At least four species, one crab and three fishes, were cited as predators of Ficopomatus (See Table of Natural Enemies). One fish species, the grey mullet Liza saliensis, does not feeds on the polychaetes; however, it grazes on the algae attached to the reefs and breaks the tube edges by “biting” (Bianchi and Morri, 1996). Several species of predators of Ficopomatus were reported feeding on the polychaetes in their native areas (See Table of Natural Enemies) and, although several of these predators species were also introduced in the same areas where Ficopomatus is invasive, nothing was mentioned in the literature about the potential predation in these invasive areas. No other enemies were reported in the literature

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

The capability of the larvae to swim is too poor (lower than 5 mm/s) (ten Hove, 1979) and they rely on local water currents (eventually wind-mediated). Since the survival of the larvae in the water is low and the swimming capability poor, the dispersal of Ficopomatus by natural means is only to short distances.
Accidental Introduction
Almost all introductions of this species were accidental. The most likely vectors are via ship/boat hull fouling and ship ballast water/sediment. However, transport of this polychaete in association with the aquaculture species is also a potential vector. In fact, there is one example of aquaculture as a potential vector of dispersal; one specimen was collected in an oyster clump in the Port of Brunswick in East River in Georgia, USA (USGS, 2009); however, the information is scarce.
Ballast water and hull fouling are the main vectors at the international level, whereas hull fouling seems to be the most important vector at the regional and local levels. For example, the introduction of Ficopomatus to the Caspian Sea occurred as fouling on vessels going from the Sea of Azov and the Black Sea via the Volga-Don Canal (Zevina and Kuznetsova, 1965). No other vectors were reported as a dispersal source.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
HitchhikerOne specimen was collected in an oyster clump in the Port of Brunswick in East River Yes Yes USGS, 2009
Interconnected waterwaysThe opening of the Volga-Don Canal in 1952 allowed the entrance of several exotic species Yes Yes Zevina and Kuznetsova, 1965

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aquaculture stockTransported as adults. See text in 'movement and dispersal' Yes Yes Fofonoff et al., 2003
Ship ballast water and sedimentTransported as adult and larvae Yes Yes Hewitt et al., 2009
Ship hull foulingTransported as adult Yes Yes Hewitt et al., 2009

Impact Summary

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Cultural/amenity Negative
Economic/livelihood Negative
Environment (generally) Positive and negative

Economic Impact

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F. enigmaticus is one of the major fouling agents on artificial surfaces and has become a nuisance in ports and marinas throughout the Mediterranean (Streftaris and Zenetos, 2006). It has impacts not only on ships and piers but also on harbour management and structures where it clogs pipes and blocks tide-gates (WGITMO, 2001). In New Zealand, Ficopomatus reached nuisance levels on artificial structures including pleasure craft, and the cooling water intake pipes for the Otahuhu Power Station, Otara Creek, after which the station changed to a freshwater cooling source (Read and Gordon, 1991). In Montevideo, Uruguay, Ficopomatus obstructed the cooling system of an oil refinery (Muniz et al., 2005). In other countries this species has also been reported affecting power station installations, such as in Denmark (Rasmussen, 1958), the Netherlands (Sluys et al., 2005), England (Markowski, 1960), the USA (Hoagland and Turner, 1980) and Italy (Bianchi and Morri, 1996). Nothing is known about the economic impact in terms of costs of maintaining the systems, ships and tide gates free of this fouling species.

Environmental Impact

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F. enigmaticus has been shown to have both negative and positive environmental impacts. Davies et al. (1989) studied the filter feeding effect of the species on the water transparency in enclosed waters in Marina da Gama (South Africa). They found that the clearance time was 0.025x106 m3 in 1.1 days which increases the light penetration by reducing the suspended particulate loads and the phytoplankton biomass. Similar results were found in Mar Chiquita coastal lagoon (Argentina) in mesocosm experiments where Ficopomatus decreased the concentration of chlorophyll a by 56% and 19% in summer and winter, respectively (Bruschetti et al., 2008). Conversely, in this same lagoon, the reefs were often observed dramatically increasing the abundance of green algae by providing a substrate for attachment. According to Sorokin et al. (2004), in a study performed in a hypereutrophic lagoon in the northwest Adriatic, the high filtering capacity of this species explained the nuisance monospecific blooms typical of eutrophication. Its long-term functioning resulted in hyper-accumulation of organic matter and nutrients and in formation of sulphide mud on the bottom (Sorokin et al., 2004).

In England, Thomas and Thorp (1994) argued if the reef density was too high filter-feeders can also deplete phytoplanktonic resources and suspended particulate organic material too much which might otherwise be utilised by other, native, filter-feeders. In addition, Thomas and Thorp (1994) mention that through production of faeces and pseudofaeces in large quantities polychaetes would also concentrate contaminants from the water column and pass them into the sediment and hence up the food chain.
Reefs of F. enigmaticus also alter the hydrodynamic flow by affecting the bedload transport and deposition of the sediment entering to the Mar Chiquita coastal lagoon (Argentina). Reefs can act as efficient traps for sediments depending on their density. Since reefs become obstacles to water movement, they generate a topographic heterogeneity and ameliorate physical conditions by accumulating and stabilizing sediments. In a coastal lagoon without reefs the water and/or the sediment coming from rainfall, river drainage, floods, and artificial channels would discharge into the sea. However, considering that 86% of the lagoon surface is colonized by reefs, the sediments entering are progressively deposited inside the lagoon rather than discharging them out (Schwindt et al., 2004a).
Impact on Biodiversity
F. enigmaticus has been repeatedly suggested to be an ecosystem engineer, an organism able to create new habitats for other species. The spaces among the reef tubes are occupied by epibenthic species, and the spaces generated below the reefs are also occupied by other benthic species. For example, in Mar Chiquita coastal lagoon (Argentina) the survival of the native crab Cyrtograpsus angulatus during the settlement process is highly dependent on the availability of small refuges in-between the tubes of the polychaete (Mendez Casariego et al., 2004), as the size of the crabs increases the availability of refuges decreases. Once the crabs reach the adult size, they live in the space available between the reefs and the mud at high densities (up to 167 individuals/m2) (Schwindt and Iribarne, 2000). These crabs use the reefs as a refuge from predation and at the same time they negatively affect the macroinfauna of the soft-bottom areas surrounding the reefs, since they can prey upon the free-living polychaetes. In short, Ficopomatus has a double effect on biodiversity in this coastal lagoon; firstly reefs positively affect the density of the native crabs by providing them essential refuge for the survival. Then, by doing this, reefs negatively affect other polychaetes species that are eaten of the crabs (Schwindt et al., 2001).
Ficopomatus also has a positive effect of epibenthic species. For example in Mar Chiquita coastal lagoon, more than 15 macroinvertebrate species live among the reef tubes plus several undetermined green algae (Schwindt and Obenat, 2005). The algae attached to the reefs increase the local primary productivity and shelter for herbivores that can vary from small invertebrates to waterfowl. Seven of the sixteen species reported living among the tubes can survive in the mudflats surrounding the reefs without the shelter they provide (Schwindt and Obenat, 2005). In Emsworth millpond (England), although there is a high temporal variation in number of species, 24 macroinvertebrates species inhabits between the reefs tubes, being the dominant residents being polychaetes, amphipods and isopods (Thomas and Thorp, 1994). In the Po River Delta (Italy) more than 15 macroinvertebrates species plus several algae species are associated to the reefs, the most abundant being the amphipods, polychaetes, barnacles and bryozoans (Bianchi and Morri, 1996).
It is interesting to note that several species associated with the reefs in Mar Chiquita coastal lagoon were also observed in association with Ficopomatus in other regions of the world where this species has invaded (see Associations).
Interaction with other exotic species was observed in California (USA). Exotic invertebrate species are more abundant amongst the Ficopomatus reef tubes than with the native oysters (Heiman et al., 2008). Reefs also have positive effects on bird communities in Mar Chiquita coastal lagoon, since they provide resting sites for local birds during high tides and also feeding sites for migratory birds (Schwindt and Iribarne, 1998; Bruschetti et al., 2009).

Social Impact

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Reefs built by F. enigmaticus have been found to affect the recreational activities in Mar Chiquita coastal lagoon (Schwindt and Iribarne, 1998). Tourism is one the main activities that supports the economy of Mar Chiquita village and the main portion of the lagoon is used for sport fishing in small rental boats. However, due to the lagoon’s shallow water regime and the high density of reefs (estimated in 1999 at 89 reefs per hectare (SE + 7.41)) with average reef sizes of 4 m in diameter) navigation can be made impossible (Schwindt et al., 2004a). As a consequence, recreational fishing has slowly decreased over the years. In some cases local people tried to eradicate the Ficopomatus reefs by breaking them into small parts, unaware that each small part would regenerate a new reef.

Keene (1980) also mentioned that reefs of Ficopomatus in Tunis lagoon (Tunisia) affected the small boat navigation making it virtually impossible.

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Long lived
  • Fast growing
  • Has high reproductive potential
  • Gregarious
Impact outcomes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Infrastructure damage
  • Modification of hydrology
  • Modification of natural benthic communities
  • Negatively impacts cultural/traditional practices
  • Negatively impacts aquaculture/fisheries
  • Negatively impacts tourism
Impact mechanisms
  • Filtration
  • Fouling
  • Herbivory/grazing/browsing
  • Interaction with other invasive species
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult/costly to control


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F. enigmaticus has no known economic value or social benefit. However, it was involved in pilot projects developed in Argentina in order to study its potential as compost material to feed hens, and other farm animals.

Uses List

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  • Research model

Similarities to Other Species/Conditions

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Within the Serpulidae, the most similar organism to F. enigmaticus is the tube worm of the genus Hydroides. This is the largest serpulid genus, and it is composed by 89 species, including one with two subspecies. Several species of Hydroides share the same subtropical and temperate distribution of Ficopomatus, and both genera have several similar morphological characteristics such as the symmetrical body, the presence of collar chetae (coarsely serrated and limbate in Ficopomatus and bayonet-type and limbate in Hydroides) and the opercular peduncle without well developed distal wings. However, a pseudoperculum (rudimentary operculum) is present in Ficopomatus and not in Hydroides (ten Hove and Kupriyanova, 2009).

Prevention and Control

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Public awareness

Among all the marine invasive species, F. enigmaticus is not one of the target species used in public awareness materials. However, this species is mentioned in over 15 invasive species web pages, most of these sites have a specific fact sheet for Ficopomatus and they offer information on contacts in case people want to report a new record.


There are no reported cases of planned eradication of F. enigmaticus. This polychaete apparently disappeared from several locations only accidentally after restoration management of the ecosystem. See Environmental Requirements with the examples in Tunis Lagoon (Tunisia) and Peel-Harvey Estuary (Australia). In both locations the ecosystems used to be highly polluted and hyper-eutrophic. After restoration the species disappeared probably due to the increase in seawater input. Thus, Ficopomatus apparently disappeared from other locations where it was previously observed but it was not as a result of specific eradication management (as in Azerbaijan, Turkmenistan, some parts of Denmark, see the Distribution Table for details).


Physical/mechanical control

Since F. enigmaticus is a successful fouling organism, it is able to grow on harbour structures, tide gates, ships, and power plants cooling systems. The only method of mechanical control reported is removal by scraping to clean all structures surfaces (Eno et al., 1997). At the Otahuhu Power Station on the Tamaki Estuary, Auckland (New Zealand) they changed from seawater to freshwater in the cooling system to control the invasion of Ficopomatus in the power plant (Read and Gordon, 1991). This last method is cited in most revisions and databases, but it is unknown if is still used or if it was successful.

Chemical control

Anti-fouling paints reduce the fouling on ships and boat hulls. Although there are experimental works investigating chemical treatment for the control of Ficopomatus settlement and larval survival, up to now, they are not regularly used. Brown et al. (2000) investigated the effect of the anti-marine-borer treatment of wood using a pressure impregnated solution of copper, chromium and arsenic compounds (CCA) on different fouling species in several coastal sites as in Ría Formosa (Portugal) where Ficopomatus was found. They observed that the polychaete was resistant to the anti-marine borer timber preservative.

Tamburri et al. (2002) studied the effect of the oxygen level in ballast water on larvae survival of different invasive species. They found that 21% of the larvae of F. enigmaticus from Elkhorn Slough (USA) survived after 2 days of exposure to oxygen levels below 0.8 mg/L.

Ecosystem Restoration

There are several examples of ecosystem restoration where F. enigmaticus used to live, however these works were not performed with the purpose of eradicate this species or because of the invasion of this species. Those restoration works were performed due to environmental problems caused by the pollution and hyper-eutrophication of the waters (see Environmental Requirements and Eradication).

Gaps in Knowledge/Research Needs

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F. enigmaticus is an invasive species in most brackish waters worldwide that is still spreading and causing environmental and economic impacts. Further knowledge on native area and the routes of introduction would be an interesting tool to compare the ecology and biology of this species in both invaded and native habitat.


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Links to Websites

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Baltic Sea Alien Species Database
Global Invasive Species Database GISD aims to increase awareness about invasive alien species and to facilitate effective prevention and management. It is managed by the Invasive Species Specialist Group (ISSG) of the Species Surviva
International Council for the Exploration of the Sea (ICES)
Red de Informacion sobre especies invasoras (I3N)


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Acre: International Polychaeotology Association (IPA), Web page maintained by Dr Geoffrey Read (NIWA, New Zealand),


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31/08/09 Original text by:

Evangelina Schwindt, Centro Nacional Patagonico (CENPAT-CONICET), Grupo de Ecología en Ambientes Costeros, Blvd. Brown 2915, Puerto Madryn (U9120ACD), Argentina

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