Lyrodus pedicellatus

Invasion History

First Non-native North American Tidal Record: 1871
First Non-native West Coast Tidal Record: 1871
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Lyrodus pedicellatus was described from Spain but is probably of Indo-Pacific origin. It is now widespread in tropical-to-warm-temperate waters around the globe and its precise native range cannot be determined yet. It was described from the Bay of Biscay, Spain, in 1849 and has been collected and redescribed under many synonymous names (Turner 1966; Turner 1971). One description, as ‘Teredo chlorotica’ was made in Boston Harbor, Massachusetts, from the hull of a whaling ship which had cruised in the Pacific Ocean (Gould 1870). It was first collected on the Pacific coast of the US in the 1870s and appears to be localized in shallow harbors where water temperatures are warm enough for breeding (Carlton 1979). This species and an unidentified Lyrodus sp. were said to have established temporary populations in thermal effluents in England and Barnegat Bay, New Jersey (Coughlan 1977; Hoagland and Turner 1980). It may be that 'Lyrodus pedicellatus' represents a complex of numerous species, but further work needs to be conducted (Borges et al. 2012; Borges et al. 2014b). This shipworm has a short (up to 36 hours) larval duration in the plankton, so long-distance dispersal is largely by ships or driftwood (Lebour 1946).

North American Invasion History:

Invasion History on the West Coast:

Lyrodus pedicellatus was first reported on the West Coast of the US from Los Angeles-Long Beach Harbor, as Teredo navalis in 1871 (Mendell 1871, cited by Carlton 1979). In San Diego Bay, it was collected in 1907 as Xylotrya stuchburyi (Kelsey 1907, cited by Carlton 1979), and reported by Bartsch (1916, cited by Carlton 1979) as Teredo diegensis. In 1920, it was collected in San Bruno Slough, San Mateo County, off South San Francisco Bay (1920, Kofoid 1921, cited by Carlton 1979). Subsequently, it was collected from many sites along the Bay, including Hunters Point, Yerba Buena, Mare Island, and Benicia (Wallour 1960; Carlton 1979). Lyrodus pedicellatus has been found in many of the smaller bays, including Elkhorn Slough (MacGinitie 1935, cited by Carlton 1979), Santa Barbara Harbor, Port Hueneme, Santa Monica Bay (Miller 1951, cited by Carlton 1979), Mugu Lagoon (Burch 1945, cited by Carlton 1979) and Newport Bay (Reish 1972, cited by Carlton 1979).

Invasion History on the East Coast:

Lyrodus pedicellatus was reported (as T. chlorotica) to occur from Florida to Texas (Dall 1889). We regard it as cryptogenic in the Western Atlantic, from North Carolina to Argentina (Turner 1966; Wallour 1960; Farrapeira et al. 2011; Museum of Comparative Zoology 2012), where it may have been introduced in the 16th–19th centuries. However, these tropical shipworms are occasionally transported north of Cape Hatteras by wooden boats and ships. In 1870, A. A. Gould described a specimen as T. chlorotica from a Pacific whaling ship in Boston harbor (Gould 1870). In 1974, a single juvenile Lyrodus sp. (L. pedicellatus or L. floridanus) was found in wood in the effluent of a nuclear power plant in Oyster Creek, Barnegat Bay, New Jersey (Hoagland and Turner 1980).

Invasion History in Hawaii:

The first collection of L. pedicellatus in the Hawaiian Islands is from a dredged, sunken, palm log, from ~ 400 m depth off Oahu in 1902 (as Teredo hawaiensis) (Dall, Bartsch & Rehder 1938, cited by Carlton and Eldredge 2009). Later collections were in Pearl Harbor (in 1935, Coles et al. 1999), Kauai (Dall, Bartsch & Rehder 1938, as T. kauaiensis, cited by Carlton and Eldredge 2009), Midway (Edmondson 1942, cited by Carlton and Eldredge 2009); and Johnston Island, Pacific Ocean (Srinivasan 1968).

Invasion History Elsewhere in the World:

Lyrodus pedicellatus is now widespread in warm-temperate to tropical waters around the world, including the Atlantic Ocean, Mediterranean and Black Seas, Indian Ocean, and West Pacific (Wallour 1960; Turner 1966; Srinivasan 1968; Sen et al. 2010; Museum of Comparative Zoology 2012). We regard it as cryptogenic over most of this range. It is apparently introduced in False Bay, South Africa (in 1931, Mead et al. 2011b), New Zealand (McKoy 1980; Cranfield et al. 1998), and the Galapagos Islands (Cruz 1996; Carlton et al. 2019). In the Tagus estuary, Portugal it replaced and apparently eliminated Teredo navalis from the 1960s–70s to the 1990s–2000s (Borges et al. 2010). In England, specimens are known from ships’ hulls and from thermal discharges (Turner 1966; Coughlan 1977). However, this species is now established and dominant in the English Channel, apparently as a result of increased temperatures (Borges 2007, cited in Borges et al. 2010).

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Description

Lyrodus pedicellatus is commonly known as the Blacktip Shipworm. It belongs to the family Teredinidae (shipworms), which are highly modified mollusks, hardly recognizable as bivalves, adapted for boring into wood. The shell is reduced to two small, ridged valves, covering the head, and is used for grinding and tearing wood fibers. The body is naked and elongated, and ends with two siphons, protected by elaborate calcareous structures called pallets (Turner 1966).

In the genus Lyrodus, the shell resembles that of the Naval Shipworm (Teredo navalis), but is smaller and more finely sculptured. The pallets have a calcareous base, which is conical distally and a brown to black cap made of periostracum. The outline resembles that of the stem and cup of a wineglass. The cap has straight sides, while the outer margin varies from a shallow U-shape to being deeply excavated, so that the conical, calcareous base protrudes (Turner 1966; Turner 1971; Coan et al. 2000). In one study, off Mumbai, India, the largest specimens reached 205 mm (Raveendram and Wagh 1991).

Genetic analysis of Lyrodus pedicellatus in the northeast Atlantic and Mediterranean Sea indicates that these two populations may represent two cryptic species (Borges et al. 2012). Further genetic analysis is needed to evaluate the possibility of cryptic species and cryptic invasions throughout the reported range of L. pedicellatus.

Potentially misidentified species - The diversity of shipworms in tropical waters is very great. Many are now widely distributed in the Atlantic, Pacific, and Indian Oceans, largely as a result of shipping. The species listed have been reported in Florida, the Caribbean, the West Coast of North America, or Hawaiian waters.

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Taxonomy

Ecology

General:

Shipworms dig long burrows in submerged wood in marine environments. They burrow by rocking and abrading the wood fibers. The mantle covers most of the length of the body, and secretes a calcareous lining along the interior of the burrow. They normally have their anterior end, with head and shells inside the burrow, and their siphons protruding. The pallets plug the burrow when the siphons are retracted (Barnes 1983).

Shipworms are protandrous hermaphrodites, beginning life as male and transforming to female, but they have no capacity for self-fertilization. Males release sperm into the water column, which fertilizes eggs for the female. The fertilized eggs are then brooded in the gills. Larvae are retained in the gills to the veliger stage. Lyrodus pedicellatus, releases its larvae in an advanced stage, as pediveligers, which spend only 2 to 24 hours in the plankton (Turner and Johnson 1971). The larvae settle in the pediveliger stage, and then rapidly metamorphose and begin boring into wood within 2–3 days. They quickly develop a calcified shell, pallets, and burrow lining (Turner and Johnson 1971). Shipworms may obtain some (or most, Paalvast and van der Velde 2013) of their nutrition from plankton, but some comes from wood, which consists largely of cellulose. Symbiotic bacteria fix nitrogen, essential for protein synthesis (Turner and Johnson 1971; Barnes 1983).

Lyrodus pedicellatus is known from fixed wood structures and panels, and from driftwood in tropical and subtropical climates. In New Zealand, it was absent in mangrove habitats (Turner 1966; McKoy 1980), but it is reported to be abundant, together with five other shipworm species, in mangrove forests of southeastern India (Nair 1984). In experiments, L. pedicelllatus from California and New Guinea did not tolerate salinities below 20–25 PSU (Eckelbarger and Reish 1972; Rayner 1979). However, this shipworm is present in the Black Sea (~18–20 PSU) (Sen et al. 2010). This shipworm survived winter temperatures as low as 4.7 °C in England (Borges 2007, cited by Borges et al. 2011).

Food:

Phytoplankton, detritus

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Marinas & Docks	None
General Habitat	Vessel Hull	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Mid Intertidal	None
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)	11	Experimental 50% survival, 7 days (Ecklebarger and Reish 1972).
Minimum Salinity (‰)	21.6	Experimental, 50% survival, 7 days (Ecklebarger and Reish 1972).
Minimum Reproductive Temperature	14	Carlton 1979
Maximum Reproductive Temperature	24	Experimental, highest tested (Ecklebarger and Reish 1972).
Minimum Reproductive Salinity	21.6	Experimental, 50% survival, 7 days (Ecklebarger and Reish 1972).
Minimum Duration	0.2	Larval duration (Turner and Johnson 1971)
Maximum Duration	1	Larval duration (Turner and Johnson 1971)
Maximum Length (mm)	205	off Mumbai, India (Raveendram and Wagh 1991)
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Polyhaline-Euhaline

General Impacts

Specific impacts of Lyrodus pedicellatus are difficult to determine, because this shipworm often co-occurs with Teredo navalis (Naval Shipworm) and a few other species (Psiloteredo megotara, North Atlantic; Bankia gouldi, Northwest Atlantic; Bankia setacea, Northeast Pacific) in temperate waters, and with a more diverse community of shipworms in subtropical and tropical waters. However, L. pedicellatus was the dominant, or one of the most abundant, shipworm species in many tropical, subtropical, and warm-temperate harbors and bays (Wallour 1960; Turner 1966; Ibrahim 1981; Nair 1984; Raveendran and Wagh 1991).

Lyrodus pedicellatus is a common shipworm in the warmer waters of the world. Its economic impacts include the devouring of historic and modern wooden boats, ships, wharves, pilings, and other structures (Atwood 1922; Turner 1971; Turner and Johnson 1971; Wallour 1960). They can affect structures used in fisheries and aquaculture (Nair 1984), and affect the timber industry in regions where logs are stored in seawater before processing (Tsunoda 1979).

Ecological effects of the L. pedicellatus invasion include competition with other shipworm species (Borges et al. 2011); wood breakdown and the incorporation of wood into marine food webs (Barnes 1983); and providing habitat by riddling wood with tunnels and galleries used by many small mobile animals (polychaetes, amphipods, isopods, etc.) (Carlton 1979). Measures to control the impacts of shipworms on vessels and marine infrastructure include the use of metal, plastic, or toxic substances (creosote, copper salts, etc.) to avoid shipworm attacks (Atwood 1922; Sen et al. 2010), with adverse effects on other marine organisms and possible toxicity to animal and human consumers.

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Regional Impacts

NEA-V	None		Ecological Impact	Competition
Lyrodus pedicellatus replaced Teredo navalis as the dominant shipworm attacking wood in the Tagus estuary, Portugal, in surveys (~1994-2010), compared to earlier surveys conducted in the 1960s-1970s. Increases in average temperature and salinity in the estuary may favor L. pedicellatus (Borges et al. 2011).
CIO-I	None		Economic Impact	Shipping/Boating
'Reported to be particularly destructive in coastal waters of Bombay' (Mumbai), rare in panels at 2m, most abundant in attacking wood at 22-62 m (Raveendran and Wagh 1991).
CIO-II	None		Ecological Impact	Herbivory
Lyrodus pedicellatus was reported to be major wood-borer, together with five other shipworm species, in the mangrove forests of Pichavaram on the southeast coast of India (Nair 1984).
CIO-II	None		Economic Impact	Fisheries
Lyrodus pedicellatus, Teredo furcifera, and other species attack wooden stakes and rafts used in oyster, pearl-oyster, fish and seaweed culture in the Gulf of Mannar (Nair 1984).
CIO-I	None		Economic Impact	Fisheries
Wooden rafts used in mussel and pearl-oyster culture in Vizhinjam Bay are attacked by L. pedicellatus and other shipworms.
NWP-3b	None		Economic Impact	Industry
In Japan, logs from the timber industry were stored in harbors. Lyrodus pedicellatus was the second-most abundant shipworm attacking these stored logs, often inflicting significant damage (Tsunoda 1979).
NWP-3a	None		Economic Impact	Industry
In Japan, logs from the timber industry were stored in harbors. Lyrodus pedicellatus was the second-most abundant shipworm attacking these stored logs, often inflicting significant damage (Tsunoda 1979).
NWP-4a	None		Economic Impact	Industry
In Japan, logs from the timber industry were stored in harbors. Lyrodus pedicellatus was the second-most abundant shipworm attacking these stored logs, often inflicting significant damage (Tsunoda 1979).
MED-V	None		Economic Impact	Shipping/Boating
Lyrodus pedicellatus was, together with T. naviais, one of the two major wood-borers in Turkish waters. Lyrodus pedicellatus produced more damage in tropical hardwoods than T. navalis, which tended to produce more damage in softwoods (Sen et al. 2010).

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