Ficopomatus enigmaticus

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

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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).

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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.

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Taxonomy

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

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Unstructured Bottom	None
General Habitat	Canals	None
General Habitat	Rocky	None
General Habitat	Vessel Hull	None
General Habitat	Grass Bed	None
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Salinity Range	Hyperhaline	40+ PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	0	Based on geographical range
Maximum Temperature (ºC)	30	Lake Timsah, Israel (Ben-Eliahu and ten Hove 2011); Lagoon of Orbetello, Italy (Bianchi and Morri 2001).
Minimum Salinity (‰)	2	Vuillemin 1965; Bianchi and Morri, 1996)
Maximum Salinity (‰)	55	Vuillemin 1965
Minimum Reproductive Temperature	18	Thames estuary, spawning. Settlement of larvae occurs as temperatures as low as 10 C (Dixon 1981)
Minimum Reproductive Salinity	10	Vuillemin 1965; Thomas and Thorp 1994
Minimum Duration	30	Larval period, field estimate (Dixon 1981)
Maximum Duration	90	Larval period, field estimate (Dixon 1981)
Maximum Length (mm)	25	Lagoon of Orbetello, Italy (Bianchi and Morri 2001).
Broad Temperature Range	None	Cold temperate-Subtropical
Broad Salinity Range	None	Oligohaline-Euhaline
Minimum Temperature (ºC)	1	Field data, Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data
Maximum Temperature (ºC)	30	Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data
Minimum Salinity (‰)	1	Bianchi and Morri 1996; Dixon 1981;Ruiz et al. unpublished data
Maximum Salinity (‰)	55	Bianchi 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).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Habitat 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).
P080	Monterey Bay		Ecological Impact	Habitat 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).
P090	San Francisco Bay		Ecological Impact	Habitat 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).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Competition
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).
P090	San Francisco Bay		Ecological Impact	Competition
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).
CA	California		Ecological Impact	Competition
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)., 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).
CA	California		Ecological Impact	Habitat 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)., 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)., 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).

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