Invasion History

First Non-native North American Tidal Record: 1976
First Non-native West Coast Tidal Record: 1976
First Non-native East/Gulf Coast Tidal Record:

General Invasion History:

Clymenella torquata is a tube-dwelling polychaete, native to the northwest Atlantic, where it occurs from the southern Gulf of St. Lawrence and the Bay of Fundy to Texas (Mangum 1962; Hughes and Thomas 1971). This worm was first collected in England in 1947 and later found in Ireland (in 2006, Minchin 2007). It was found to be abundant in mudflats of Boundary Bay (British Columbia-Washington) in 1976 (Swinbanks and Murray 1981), and was collected there again in 1980 (Banse 1981). In 2006, C. torquata was found to be abundant in Samish Bay, Washington, where it can cause problems for oyster-rearing operations (Mach et al. 2012).

North American Invasion History:

Invasion History on the West Coast:

Clymenella torquata was collected in 1976 (Swinbanks and Murray 1981) and 1980 (Banse 1981) in Boundary Bay, British Columbia. It may have been introduced with plantings of Eastern Oysters (Crassostrea virginica) in the 1930s (Mach et al. 2012). In 2008, established populations of this worm were found in Campbell River on Vancouver Island and Roberts Bank on the Fraser River Delta (Mach et al. 2012). A notable range extension was an occurrence in Samish Bay, Washington, 75 km south of Boundary Bay, where this worm was first reported in 2006 (Davis 2007). Clymenella torquata has a benthic larva, so oyster and clam transplants, are the likeliest vector for the spread of this polychaete.

Invasion History Elsewhere in the World:

Clymenella torquata was first reported from Europe in Whitstable, Kent, on the English Channel, in 1947, possibly introduced with a planting of Eastern Oysters in 1936. It is apparently still established there. A single specimen was collected in Northumberland in 1976 (Eno et al. 1997). This worm was collected in Carlingford Lough, Northern Ireland in 2006 (Minchin 2007). Transplants of Eastern Oysters were the likely vector for introduction of C. torquata to the British Isles (Eno et al. 1997).


Description

Clymenella torquata is a head-down tube-dwelling maldanid polychaete. Its body is slender, and resembles a stick of bamboo, with elongated body segments, and poorly defined parapodia. The body consists of a prostomium, a peristomial segment lacking setae, 18 chaetigers, and a funnel-like anal plaque (pygidium). The worm is positioned head-down in its tube, with the pygidium, which has 20-22 short cirri, protruding. The lobes can fold, contract and close off the tube (Lippson and Lippson 1997; Mach et al. 2012).

The head is blunt and concave, with an oval prostomium. The cephalic plaque is wrinkled. The mouth is ventral. The nuchal organs are broad and parallel, running along the surface of the cephalic plate. Chaetiger 1 has 5-10 notochaetae. The neuropodia of the first 3 chaetigers have ridges bearing 9-12 uncini in rows. The 4th chaetiger has a membranous collar, which extends anteriorly to cover part of chaetiger 3. There are three pre-anal segments, lacking chaetae. The anus is central. The anal cirri are all about the same length. The overall length of adult worms ranges from 50 to 195 mm, although more commonly does not exceed 110 mm. The color is variable, but usually slightly iridescent yellow-orange, with red blood-vessels near the joints. Some populations are tinged with dull brown, while others accumulate green pigment, depending on the habitats, sandy or muddy respectively (Leidy 1855; Hartman 1945; Newell 1949; Hartman 1951; Mangum 1962; Banse 1981; Hobson and Banse 1981; Kozloff 1987; Lippson and Lippson 1994; Mach et al. 2012).


Taxonomy

Taxonomic Tree

Kingdom:   Animalia
Phylum:   Annelida
Class:   Polychaeta
Subclass:   Scolecida
Family:   Maldanidae
Genus:   Clymenella
Species:   torquata

Synonyms

Clymene torquatus (Leidy, 1855)
Paraxiothea latens (Webster, 1879)
Axiothea torquata (Lewis, 1897)
Clymenella torquata calida (Hartman, 1951)

Potentially Misidentified Species

Clymenella californica
This species has spines in its neuropodia and 21 chaetigers. It has been reported from low intertidal mud in San Francisco Bay (Blake and Ruff 2007).

Clymenella complanata
This species has spines in its neuropodia and 22-27 chaetigers. It has been reported from crevices in shale rocks in San Mateo County (Blake and Ruff 2007).

Ecology

General:

Clymenella torquata is a marine, tube-dwelling polychaete, found in mud, sand, and gravel in intertidal and subtidal habitats. The sexes are separate. The period of spawning in Massachusetts and Britain is brief, over a few days in April or May, possibly determined by tidal conditions. Eggs are deposited in the mud near the mouth of the tube, and are fertilized by sperm shed into the water (Mead 1897; Newell 1951). The larvae are demersal, developing through a trochophore stage and reaching a length of 0.6 mm and several segments in about a week. They are capable of feeding on benthic diatoms and constructing tubes in about 10 days. Metamorphosis is gradual, and is more or less complete in 24-25 days (Newell 1951). In the northern part of the range, at least, spawning appears to be annual.

Clymenella torquata tolerates a wide range of temperature and salinity as indicated by its native range and experimental data, from near-freezing to 40.5°C, and salinities of 15-40 PSU (Mangum 1964; Wass 1972; Kenny 1969a; Kenny 1969b). It can withstand relatively low oxygen and/or high hydrogen sulfide concentrations (Fuller 1994). The substrates this worm inhabits include fine gravel, sand, and mud (Mangum 1964). The worm dwells head-down in a vertical tube, up to 200 mm long, composed of sand or mud particles, detritus, and mucus. The cylindrical tube extends 5-30 cm deep, and projects above the surface of the sediment, with the anal cirri projecting when the animal is undisturbed (Lippson and Lippson 1997). Tube building was most efficient with sand grains of 0.25 to 0.84 mm, at 25 C and 35 PSU ( (Kenny 1969). In estuaries with wide tidal ranges, C. torquata is abundant in the lower intertidal zone, but in estuaries with a small tide range, it is limited to subtidal waters (Mangum 1964).

Clymenella torquata can obtain surface food in two wayS: ingesting sediment directly from the mouth deeply in the sediment or using the posterior body to hoe the surface material directly into the tube (Dobb and Whitlatch 1982; Weinberg 1988). The worms often show missing and/or regenerating segments, indicating sublethal predation (Mangum 1964), by fishes (Schneider & Harrington 1981), shorebirds (Schneider & Harrington 1981) or other predators. The regneration ability of this worm has been studied. It seems to have have different tail- and head-determining factors according to the body region involved and it usually tends to maintain its original 22 segments (Sayles 1942; Smith 1963).

Food:

Diatoms, sediment, detritus

Consumers:

Fishes, shorebirds, crabs

Trophic Status:

Deposit Feeder

DepFed

Habitats

General HabitatGrass BedNone
General HabitatUnstructured BottomNone
General HabitatOyster ReefNone
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEndobenthicNone


Tolerances and Life History Parameters

Minimum Temperature (ºC)5 Field, South Carolina (Kenny 1969b) but lower in its northern geographical range. This species is stablished in estuaries with winter ice cover
Maximum Temperature (ºC)39Experimental, 50% survival, heating at 1 C per 5 min, (Kenney 1969b)
Minimum Salinity (‰)15Experimental, 50% survival, 45 hrs (Kenney 1969b)
Maximum Salinity (‰)40Experimental, highest tested (Kenney 1969b)
Minimum Length (mm)50Hartman 1945; Newell 1949; Mangum 1962
Maximum Length (mm)195Hartman 1945; Newell 1949; Mangum 1962
Broad Temperature RangeNoneCold temperate-Warm temperate
Broad Salinity RangeNoneMesohaline-Euhaline

General Impacts

In dense populations, Clymenella torquata can function as an ecosystem engineer, by modifying the quality of sediments, sub-ducting organic and inorganic matter, oxygenating and loosening packed sand, and mixing sediments with detritus and mucus (Sanders et al. 1962; Levin et al. 1997). Moreover its role can be considered positive for the retention and burial of eelgrass seeds, thereby favoring germination (Luckenbach and Orth 1999). This worm has recently invaded Samish Bay, Washington, and in this new environment it has had adverse effects on oyster culture by destabilizing the sediment through bioturbation (Davis 2007; Hancock et al. 2008; Mach et al. 2012). In its native range, it can increase the surface microbial mat leading to microalgal enrichment which stabilizes surface sediments (Dobb and Whitlatch 1982; Fuller 1994; Leven et al. 1997; Lukenbach and Orth 1999; Campbell and Lindsay 2014).  

Economic Impacts

Fisheries - Clymenella torquata has had adverse impacts on Pacific Oyster (Crassostrea gigas) culture in Samish Bay, where oysters were cultivated on a packed sand bottom. The worms developed dense populations, excreting fine sand and detritus out of their tubes, and pumping in water, softening the sediment, and creating areas of soft muck, which can suffocate oysters. Several control methods have been tried to reduce worm densities and increase the firmness of the sediment (Davis 2007; Hancock et al. 2008; Mach et al. 2012).

Ecological Impacts

Clymenella torquata can develop dense populations in its native and invaded ranges, and can have significant effects on the qualities of the sediment it inhabits. The worms feed head-down in their tubes in the sediment, ingesting and excreting fine sediment and detritus, which is replaced by water, creating a loose, spongy texture in the sediment. The effects were seen in Barnstable Harbor, Massachusetts, where C. torquata is native, but also recently in Samish Bay, Washington (Sanders et al. 1962; Mach et al. 2012). In addition to adverse effects on oyster culture, the altered sediment is considered a threat to eelgrass, important as food and habitat for fishes and waterfowl (Davis 2007; Hancock et al. 2008).

Regional Impacts

NEP-IIIAlaskan panhandle to N. of Puget SoundEconomic ImpactFisheries
In Samish Bay, WA high densities of Clymenella torquata have altered the texture of mudflats, releasing fine excreted feces, turning mid-sand tidelands into soft muck. The suspended sediments are detrimental to Pacific Oysters (Crassostrea gigas) being cultivated in the Bay (Davis 2007; Hancock et al. 2008; Mach et al. 2012). Hancock et al. (2008) tested five control methods, adding shells to sediment, rototilling, hydraulic pumping, covering, and compression in reducing worm biomass, and increasing sediment firmness. These treatments did not reduce worm abundance, but some of them did increase sediment firmness (Hancock et al. 2008).
NEP-IIIAlaskan panhandle to N. of Puget SoundEcological ImpactHabitat Change
In Samish Bay, WA high densities of Clymenella torquata have altered the texture of mudflats, creating a spongy texture, due to the tubes, and fine, excreted feces, turning mid-sand tidelands into soft muck. These suspended sediments are detrimental to oysters and Eelgrass (Zostera marina) (Davis 2007; Hancock et al. 2008; Mach et al. 2012).
P293_CDA_P293 (Strait of Georgia)Economic ImpactFisheries
In Samish Bay, WA high densities of Clymenella torquata have altered the texture of mudflats, releasing fine excreted feces, turning mid-sand tidelands into soft muck. The suspended sediments are detrimental to Pacific Oysters (Crassostrea gigas) being cultivated in the Bay (Davis 2007; Hancock et al. 2008; Mach et al. 2012). Hancock et al. (2008) tested five control methods: adding shells to sediment, rototilling, hydraulic pumping, covering, and compression for reducing worm and tube biomass, and increasing sediment firmness. These treatments did not reduce worm abundance, but some of them did increase sediment firmness (Hancock et al. 2008).
P293_CDA_P293 (Strait of Georgia)Ecological ImpactHabitat Change
In Samish Bay, WA high densities of Clymenella torquata have altered the texture of mudflats, creating a spongy texture, due to the tubes, and fine, excreted feces, turning mid-sand tidelands into soft muck. These suspended sediments are detrimental to oysters and to Eelgrass (Zostera marina) (Davis 2007; Hancock et al. 2008; Mach et al. 2012).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
NA-ET2 Bay of Fundy to Cape Cod 0 Native Estab
NA-ET3 Cape Cod to Cape Hatteras 0 Native Estab
CAR-VII Cape Hatteras to Mid-East Florida 0 Native Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 0 Native Estab
NEP-III Alaskan panhandle to N. of Puget Sound 1976 Def Estab
P293 _CDA_P293 (Strait of Georgia) 2006 Def Estab
NEA-II None 1936 Def Estab
P297 _CDA_P297 (Strait of Georgia) 1976 Def Estab
G074 _CDA_G074 (Crystal-Pithlachascotee) 0 Native Estab
M080 New Jersey Inland Bays 0 Native Estab
G080 Suwannee River 0 Native Estab
N010 Passamaquoddy Bay 0 Native Estab
N020 Englishman/Machias Bay 0 Native Estab
N135 _CDA_N135 (Piscataqua-Salmon Falls) 0 Native Estab
M040 Long Island Sound 0 Native Estab
N165 _CDA_N165 (Charles) 0 Native Estab
N195 _CDA_N195 (Cape Cod) 0 Native Estab
M026 _CDA_M026 (Pawcatuck-Wood) 0 Native Estab
S030 Bogue Sound 0 Native Estab
M010 Buzzards Bay 0 Native Estab
N170 Massachusetts Bay 0 Native Estab
M130 Chesapeake Bay 0 Native Estab
S020 Pamlico Sound 0 Native Estab
S140 St. Catherines/Sapelo Sounds 0 Native Estab
G330 Lower Laguna Madre 0 Native Estab
G300 Aransas Bay 0 Native Estab
G200 Barataria Bay 0 Native Estab
NA-S3 None 0 Native Estab
NA-ET1 Gulf of St. Lawrence to Bay of Fundy 0 Native Estab
CAR-IV None 0 Native Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude

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