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:

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 on the Gulf Coast:

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).


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.


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Mollusca
Class:   Bivalvia
Subclass:   Heterodonta
Order:   Myoida
Superfamily:   Pholadoidea
Family:   Teredinidae
Genus:   Lyrodus
Species:   pedicellatus

Synonyms

Teredo arabica (Roch, 1935)
Teredo chlorotica (Gould, 1870)
Teredo pertingens (Iredale, 1932)
Teredo (Lyrodus) hibicola (Koronuma, 1931)
Teredo (Pingoteredo) tristi (Iredale, 1936)
Teredo (Teredo) honoluluensis (Edmondson, 1946)
Teredo (Teredo) india (Nair, 1956)
Teredo (Teredo) madrasensis (Nair, 1956)
Teredo (Teredops) diegensis midwayensis (Edmondson, 1946)
Teredo (Teredops) hawaiensis (Dall, Bartsch, and Rehder, 1938)
Teredo (Teredops) kauiensis (Dall, Bartsch, and Rehder, 1938)
Teredo (Teredops) tateyamensis (Koronuma, 1931)
Teredo calmani (Roch, 1931)
Teredo dagmarae (Roch, 1931)
Teredo dalli (Moll and Roch, 1931)
Teredo franziusi (Roch, 1929)
Teredo helleniusi (Moll, 1936)
Teredo kiiensis (Taki and Habe, 1945)
Teredo lomensis (Roch, 1929)
Teredo malaccana (Roch, 1935)
Teredo nodosa (Roch, 1929)
Teredo pedicellata (de Quatrefages, 1849)
Teredo pedicellata truncata (Jeffreys, 1865)
Teredo pochhammeri (Moll, 1931)
Teredo robsoni (Roch, 1931)
Teredo samoanensis (Bartsch, 1927)
Teredo siamensis (Bartsch, 1927)
Teredo togoensis (Roch, 1929)
Teredo townsendi (Bartsch, 1922)
Teredo yatsui (Moll, 1929)
Teredo hawaiensis (Dall, Bartsch & Rehder, 1938)

Potentially Misidentified Species

Lyrodus bipartitus
Cosmopolitan, tropical, subtropical

Lyrodus floridanus
W Atlantic, subtropical. It was synonymized with L. pedicellatus, but later found to differ in its life history and geneitcs (Borges et al. 2012).

Lyrodus medilobatus
Cosmopolitan, tropical, subtropical

Lyrodus takanoshimensis
Cosmopolitan, tropical, subtropical, temperate, introduced in NE Pacific (British Columbia)

Nototeredo knoxi
Subtropical W Atlantic, oceanic

Psiloteredo megotara
N Atlantic, oceanic

Teredo bartschi
Cosmopolitan, tropical, subtropical, introduced in NE Pacific

Teredo clappi
Cosmopolitan, tropical, subtropical

Teredo fulleri
Cosmopolitan, tropical,

Teredo furcifera
Cosmopolitan, tropical, subtropical

Teredo navalis
Cryptogenic in NE Atlantic, NW Pacific, and Indo-West Pacific, but introduced in NW Atlantic, S Atlantic, NE Pacific, and SW Pacific

Teredora malleolus
Temperate-subtropical Atlantic, oceanic

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

Habitats

General HabitatCoarse Woody DebrisNone
General HabitatMarinas & DocksNone
General HabitatVessel HullNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeLow IntertidalNone
Tidal RangeMid IntertidalNone
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone

Life History


Tolerances and Life History Parameters

Minimum Temperature (ºC)11Experimental 50% survival, 7 days (Ecklebarger and Reish 1972).
Minimum Salinity (‰)21.6Experimental, 50% survival, 7 days (Ecklebarger and Reish 1972).
Minimum Reproductive Temperature14Carlton 1979
Maximum Reproductive Temperature24Experimental, highest tested (Ecklebarger and Reish 1972).
Minimum Reproductive Salinity21.6Experimental, 50% survival, 7 days (Ecklebarger and Reish 1972).
Minimum Duration0.2Larval duration (Turner and Johnson 1971)
Maximum Duration1Larval duration (Turner and Johnson 1971)
Maximum Length (mm)205off Mumbai, India (Raveendram and Wagh 1991)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNonePolyhaline-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.


Regional Impacts

NEA-VNoneEcological ImpactCompetition
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-INoneEconomic ImpactShipping/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-IINoneEcological ImpactHerbivory
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-IINoneEconomic ImpactFisheries
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-INoneEconomic ImpactFisheries
Wooden rafts used in mussel and pearl-oyster culture in Vizhinjam Bay are attacked by L. pedicellatus and other shipworms.
NWP-3bNoneEconomic ImpactIndustry
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-3aNoneEconomic ImpactIndustry
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-4aNoneEconomic ImpactIndustry
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-VNoneEconomic ImpactShipping/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).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
RS-3 None 0 Crypto Estab
RS-2 None 0 Crypto Estab
RS-1 None 0 Crypto Estab
IP-1 None 0 Crypto Estab
CIO-I None 0 Crypto Estab
CIO-II None 0 Crypto Estab
EAS-VI None 0 Crypto Estab
CIO-V None 0 Crypto Estab
AUS-XI None 0 Crypto Estab
AUS-X None 1931 Crypto Estab
AUS-VII None 1931 Crypto Estab
AUS-VI None 0 Crypto Estab
NZ-IV None 1962 Def Estab
SP-XXI None 1902 Def Estab
SP-VIII None 1924 Crypto Estab
EAS-I None 0 Crypto Estab
MED-III None 0 Crypto Estab
WA-II None 1929 Crypto Estab
NWP-3a None 0 Crypto Estab
NWP-2 None 0 Crypto Estab
MED-VII None 0 Crypto Estab
NEA-V None 1849 Crypto Estab
MED-VI None 0 Crypto Estab
MED-VIII None 0 Crypto Estab
MED-II None 0 Crypto Estab
MED-I None 0 Crypto Estab
MED-IV None 0 Crypto Estab
MED-V None 0 Crypto Estab
SA-I None 1963 Crypto Estab
CAR-V None 0 Crypto Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1889 Crypto Estab
NEA-II None 1865 Def Estab
NEP-V Northern California to Mid Channel Islands 1920 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1871 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 1889 Crypto Estab
SEP-H None 1960 Def Estab
SP-XII None 0 Crypto Estab
NA-ET2 Bay of Fundy to Cape Cod 1870 Def Failed
NWP-3b None 0 Crypto Estab
NA-ET4 Bermuda 0 Crypto Estab
CAR-II None 0 Crypto Estab
CAR-IV None 0 Crypto Estab
CAR-III None 0 Crypto Estab
GA None 0 Crypto Estab
WA-I None 0 Crypto Estab
EAS-III None 0 Crypto Estab
WA-IV None 1931 Def Estab
AUS-XII None 1936 Crypto Estab
AUS-IX None 0 Crypto Estab
AUS-IV None 0 Crypto Estab
AUS-VIII None 0 Crypto Estab
P020 San Diego Bay 1876 Def Estab
G070 Tampa Bay 0 Crypto Estab
G310 Corpus Christi Bay 0 Crypto Estab
G260 Galveston Bay 0 Crypto Estab
S190 Indian River 0 Crypto Estab
P050 San Pedro Bay 1871 Def Estab
P040 Newport Bay 1972 Def Estab
P060 Santa Monica Bay 1951 Def Estab
P061 _CDA_P061 (Los Angeles) 1945 Def Estab
P062 _CDA_P062 (Calleguas) 1951 Def Estab
P065 _CDA_P065 (Santa Barbara Channel) 1951 Def Estab
P080 Monterey Bay 1935 Def Estab
P090 San Francisco Bay 1920 Def Estab
S206 _CDA_S206 (Vero Beach) 0 Crypto Estab
G080 Suwannee River 0 Crypto Estab
S200 Biscayne Bay 0 Crypto Estab
G330 Lower Laguna Madre 0 Crypto Estab
S050 Cape Fear River 0 Crypto Estab
S045 _CDA_S045 (New) 0 Crypto Estab
S080 Charleston Harbor 0 Crypto Estab
N170 Massachusetts Bay 1870 Def Failed
NA-ET3 Cape Cod to Cape Hatteras 1974 Def Failed
M070 Barnegat Bay 1974 Def Failed
SP-IX None 1968 Crypto Estab
SP-XIV None 1968 Crypto Estab
NWP-4a None 0 Crypto Estab
SA-IV None 0 Crypto Estab
SA-III None 0 Crypto Estab
SA-II None 0 Crypto Estab
P058 _CDA_P058 (San Pedro Channel Islands) 1964 Def Estab
MED-VIII None 0 Crypto Estab
MED-IX None 0 Crypto Estab
EA-III None 0 Crypto Estab
NEA-IV None 2009 Crypto Estab
NEA-III None 1946 Def Unk
NEA-VI None 1940 Crypto Estab
PAN_CAR Panama Caribbean Coast 0 Crypto Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1965 Def Failed
WA-V None 0 Def Estab
CIO-III None 0 Crypto Estab
AUS-II None 0 Crypto Estab
AUS-I None 0 None None
SP-I None 0 None None
SEP-Z None 1989 Def Estab
SEP-I None 1987 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
27052 Eckelbarger and Reish 1972) 1968 1968-01-01 Alamitos Bay Def 33.7502 -118.1185
28147 Carlton 1979 1920 1920-01-01 San Bruno Slough- San Bruno Def 37.6527 -122.3719
29679 Carlton 1979 1945 1945-01-01 Mugu Lagoon Def 34.1016 -119.0999
30098 Carlton 1979 1871 1871-01-01 Los Angeles/Long Beach Harbor Complex Def 33.7632 -118.2526
30182 Cohen and Carlton, 1995 1972 1972-01-01 Newport Bay Def 33.6092 -117.9067
30414 MacGinitie 1935, cited by Carlton 1979 1935 1935-01-01 Elkhorn Slough General Location Def 36.8086 -121.7856
31880 Cohen and Carlton, 1995 1876 1876-01-01 San Diego Bay Def 32.6717 -117.1439
32270 Miller 1951, cited by Carlton 1979 1951 1951-01-01 Santa Barbara Harbor Def 34.4057 -119.6913
32581 Miller 1951, cited by Carlton 1979 1951 1951-01-01 Port Hueneme Def 34.1496 -119.2082
32721 Miller 1951, cited by Carlton 1979 1951 1951-01-01 Santa Monica, General Location Def 34.0077 -118.4985

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