Limnoria lignorum (gribble)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Habitat List
- Biology and Ecology
- Latitude/Altitude Ranges
- Water Tolerances
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Impact Summary
- Economic Impact
- Environmental Impact
- Social Impact
- Risk and Impact Factors
- Uses List
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Limnoria lignorum Rathke 1799
Preferred Common Name
Other Scientific Names
- Cymothoa lignorum Rathke, 1799
- Limnoria terebrans Leach, 1815
- Limnoria uncinata Heller, 1866
Local Common Names
- Denmark: paelekrebs
- Germany: bohrassel
- Netherlands: boorpissebed; paalpissebed
Summary of InvasivenessTop of page
Limnoria lignorum (Rathke, 1799) is a wood-boring isopod (Crustacea) yellowish in colour and up to 5.6 mm in length. The species is distributed within the temperate and boreal Northern Hemisphere (Cookson, 1991), but the history of its introduction and spread have been significantly complicated by centuries of cross oceanic travel in the hulls of wooden ships. L. lignorum damages ship hulls, pilings and other wooden structures in contact with sea water. Damage is most pronounced near the low tide level and typically occurs at depths of 0–30 m sea water. The possibility of rapid population increase due to extended parental care and a high potential of dispersal, through rafting and shipping, ensure its invasive success. L. lignorum attack occurs on the surface of the wood, which makes the wood highly porous and friable causing further deeper erosion. The deterioration is a matter of considerable economic importance along the Pacific and Atlantic coasts of America, Northern Europe and Northwest Pacific. In some parts of its range L. lignorum may be replaced by other species, such as L. quadripunctata. Its replacement may be related to significant warming of coastal waters either due to climate change or to local factors.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Crustacea
- Class: Malacostraca
- Subclass: Eumalacostraca
- Order: Isopoda
- Family: Limnoriidae
- Genus: Limnoria
- Species: Limnoria lignorum
Notes on Taxonomy and NomenclatureTop of page
The family Limnoriidae consists of small isopods which tunnel into wood, seagrasses or macroalgae. It contains three genera: Paralimnoria, Lynseia and Limnoria, (Cookson and Poore, 1994). Cookson (1991) reported 51 species within the family and since the publication of his monograph, several new species have been added (Cookson and Lorenti, 2001; Castello, 2011).
DescriptionTop of page
The body is prolongated oval shape; the length is about 3 times that of the width. Dorsal surface is yellowish in colour. Three main body regions: the head (cephalon), thorax (peraeon) and abdomen (pleon). The cephalon, globe-shaped, deepens in the first segment of peraeon. Eyes small, round, black. Antenna 1 and antenna 2 short, flagellum with 4 articles. Mandible expanded at apex, delicately toothed; with 6 setae. The peraeon consists of 7 segments similar in shape and bears 7 pairs of walking limbs (peraeopods). The pleon bears 5 pairs of flattened swimming or respiratory appendages (pleopods) while the pleotelson carries a pair of uropods. Uropod exopod much shorter than the endopod and with an apical claw; endopod elongate, apex blunt, lacking claw. Pleopod 5 lacks marginal setae. Fifth pleon segment with an anteriorly situated mid-dorsal longitudinal carina, which bifurcates posteriorly and lacks tubercles; posterior dorsal margin without tubercles but with spike-like bristles; pleotelsonic margin with sheathed setae, and stout unsheathed setae. Tegument pitted, especially on pleonite 5 and anterior section of pleotelson; scaly (scales with single large scale centrally).
Size: generally 0.8-4.0 mm, typically 3 mm or less (Cookson, 1991). Kussakin (1963) found a specimen of maximal size from Kamchatka: 5.5 mm in length and 1.4 mm in pleotelsonic width. In the largest specimen from the Kuriles these measurements are 5.6 mm and 1.5 mm respectively, and in that from the Sea of Japan 5.1 mm and 1.2 mm. The smallest oviparous female measures 3.0 mm in length and 0.8 mm in pleotelsonic width. The diameter of a fertilized egg is 0.4 mm.
Holotype not known (type locality: probably Bergen, Norway).
DistributionTop of page
L. lignorum has mainly been reported from temperate and boreal waters in the Northern Hemisphere (Cookson, 1991). Menzies (1957) observed that this species is the only limnorid species found to occupy the fringe of the Arctic, meaning that it has a truly boreal distribution, which is unique for an essentially warm water group of animals. The precise area of origin and distribution remains unclear due to identification difficulties and centuries of transoceanic and interoceanic travel of the species in the hulls of wooden ships (Kussakin, 1963). Some authors argue that L. lignorum has worldwide distribution. However, this is unclear as reports from different locations may be based on specimens of other species (Holthuis, 1949). Menzies (1959) and later authors (Carlton, 1979) confirmed that many species reported to be L. lignorum in the first half of the twentieth century have now been identified as other species.
On the Atlantic coast the distribution appears restricted between 47°N (Argentia, Newfoundland) and 42°N (Rhode Island) (Menzies, 1957). Jones (1963) suggests that the southern limit may be governed by the requirement of a suitable low temperature (about 8°C). Abood et al. (1995) found the species to be destroying wooden structures in New York Harbor.
The northern limit in the Northeast Atlantic is thought to be determined by the 0ºC isotherm surrounding the Arctic Ocean (Menzies, 1957). The most northern location L. lignorum has been found is near Torsvag (Somme, 1940). In Europe, the species is known from Norway southwards but its limits may extend to a latitude little beyond southern Britain (Somme 1940; Jones, 1963; Santhakumaran, 1984). In the Netherlands, L. lignorum has been found in most areas along the coast except for those with low salinity, such as the Wester Scheldt, or in brackish parts of estuaries of larger rivers and the southern part of the Zuiderzee. L. lignorum is found around the British coast where it may be considered endemic (Jones, 1963) but has not been found in northern Scotland and is absent or rare from the south-western tip of England (Jones, 1963). In the Barents Sea, L. lignorum is found on the Murman coast, having been first reported there in 1908 (Kussakin, 1963). From the White Sea the species has been reported from drowned wood in the Kandalaksha Bay and near Kem.
In the Russian Far Eastern seas L. lignorum is distributed from eastern Kamchatka and Komandorskie Islands to the Russian coast of the Sea of Japan, while it is absent in the colder areas of Okhotsk Sea, except its southern part (Kussakin, 1963). The species is found on the east shore of the Kuril Islands and the Sakhalin (Iljin, 1992). In the Russian Far Eastern seas the first record was made by Grebnitski, who collected specimens of Limnoria in eastern Kamchatka and at the Komandorski Islands in 1880 (Kussakin, 1963).
In Japan, the distribution of the species is poorly known: Shino (1944, 1950) reported L. lignorum from Japan (found near Akkeshi, Onagawa, Chiba, Misaki, Toba, Tanabe Bay, Muhojima, Tomioka, Noto), but Menzies (1957) considered the species as L. tripunctata. Some contemporary authors (Doi et al., 2011) regard L. lignorum from Japan as a cryptogenic species.
Menzies (1957) considers the species to be native to the Pacific coast of America, where it extends from about 58°N (Kodiak Island, Alaska) to 39°N (Point Arena, California). Findings in San Francisco Bay may represent a range extension rather than a true invasion (Ray, 2005). Many findings further south on the Pacific coast, however, which were recorded as L. lignorum before the 1950s (Arnold, 1873) are most likely to be another species (Carlton, 1975, 1979; Cohen and Carlton, 1995), such as L. trupunctata.
There are several reports of L. lignorum from areas outside of its currently accepted distribution. Most of these observations are from early records of the species, many yet to be verified, and include reports from Africa (Stebbing, 1908, 1910; Calman, 1921; Douglas, 1981), the Adriatic Sea (Heller, 1866), Black Sea (Moll, 1915), Falkland Islands (Tattersall, 1914), New Zealand (Calman, 1921), Australia (Whitelegge, 1901; Hale, 1929), China (Wei Chongde, 1991; Huang et al., 1993; Huang, 2001), South Korea (Kuhne, 1976) and Egypt (El-Shanshoury et al., 1994). In Africa, the species is not listed in any of the latest papers indicating that it may not be currently present in the region (Robinson et al., 2005; Griffiths et al., 2009). More recent reports from East Asia and Egypt could suggest a wider area of distribution than is currently acknowledged.
Distribution TableTop of page
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/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Arctic Sea||Present||Invasive||Menzies, 1957; Kussakin, 1963; Iljin, 1992|
|Atlantic, Northeast||Widespread||Invasive||Menzies, 1957; Jones, 1963; Kussakin, 1963; Cookson, 1991; Castello, 2011; Noel, 2011||First recorded in 1790s|
|Atlantic, Northwest||Widespread||Invasive||Menzies, 1957; Cookson, 1991|
|Atlantic, Southwest||Present||NMNH, 2012||Museum record|
|Atlantic, Western Central||Present||NMNH, 2012||Museum record|
|Pacific, Eastern Central||Present, few occurrences||Introduced||Carlton, 1979||In San Francisco bay, identified as L. lignorum, but probably L. tripunctata|
|Pacific, Northeast||Widespread||Invasive||Menzies, 1957; Cookson, 1991||Possibly native|
|Pacific, Northwest||Widespread||Invasive||Cookson, 1991; Iljin, 1992; Ray, 2005||Cryptogenic origin|
|Pacific, Southwest||Present||Introduced||Whitelegge, 1901||Occasional findings in ports|
|China||Present||Wei Chongde, 1991; Huang et al., 1993; Huang, 2001|
|Japan||Present||Invasive||Shino, 1944; Shino, 1950; Doi et al., 2011||Cryptogenic origin|
|-Hokkaido||Present||Invasive||Cookson, 1991||Cryptogenic origin|
|-Honshu||Present||Tsunoda and Nishimoto, 1976|
|Korea, Republic of||Present||Kuhne, 1976||Near Chinhae|
|Egypt||Present||El-Shanshoury et al., 1994||Port Said harbour|
|South Africa||Absent, unreliable record||Douglas, 1981||Near Knysna|
|Canada||Present||Menzies, 1957; Brunel, 1963; Bohn and Walden, 1970||Atlantic and Pacific coast|
|-British Columbia||Widespread||Invasive||Menzies, 1957; Clamp, 1988; Quayle, 1992||Cryptogenic origin|
|-New Brunswick||Widespread||Invasive||Henderson, 1924|
|-Newfoundland and Labrador||Widespread||Invasive||Menzies, 1957|
|-Nova Scotia||Widespread||Invasive||Menzies, 1957|
|USA||Present||Menzies, 1957; Ruiz et al., 2000; NMNH, 2012|
|-Alaska||Widespread||Invasive||Menzies, 1957; Richards and Belmore, 1976|
|-California||Localised||Introduced||mid 19th C||Menzies, 1957; Carlton, 1975|
|-Florida||Present||NMNH, 2012||Preserved specimen, Mayport, Mouth of Saint Johns River|
|-Maine||Absent, unreliable record||Menzies, 1957; NMNH, 2012||Preserved specimen, collected in Aug 1905, Portland Harbor|
|-New Hampshire||Present||Menzies, 1957|
|-New York||Present||Invasive||Abood et al., 1995; NMNH, 2012||Wooden structures in New York Harbor|
|-Rhode Island||Present||Menzies, 1957|
|Argentina||Absent, unreliable record||NMNH, 2012||Preserved specimen, collected Feb 1915, Puerto Madryn|
|Germany||Present||Invasive||Jones et al., 1972|
|Iceland||Present||Stephensen, 1929; Menzies, 1957; Svavarsson, 1982||On the west coast|
|Ireland||Present||Holmes and Jeal, 1987; Grave and Holmes, 1998|
|Netherlands||Present||Holthuis, 1949; NMNH, 2012||Preserved specimen, collected Jan 23 1949, North Sea, South Holland|
|Norway||Present||Rathke, 1799; Sars, 1899; Somme, 1940; Nair, 1959; Santhakumaran, 1984|
|Russian Federation||Present||Invasive||Jesakova, 1961; Kussakin, 1963|
|-Northern Russia||Localised||Introduced||Invasive||Ryabchikov, 1957; Iljin, 1992; Vinogradov et al., 2006||Barents Sea; White Sea|
|-Russian Far East||Widespread||Introduced||Invasive||Kussakin, 1963; Iljin, 1992|
|UK||Widespread||Invasive||Menzies, 1959; Jones, 1963||Rare or absent from the southwestern tip of England and not found around the coast of Northern Scotland|
|Australia||Present, few occurrences||Whitelegge, 1901||Sydney Harbor, occasional findings|
History of Introduction and SpreadTop of page
Although the exact origin of L. lignorum is undetermined, some consider the species native to the Pacific coast of America (Menzies, 1957) and others consider it native to Britain (Jones, 1963). However, most treat the species origin as cryptogenic (Shino, 1950; Ruiz et al., 2000).
L. lignorum may have been introduced to other areas as early as the sixteenth century, when fouling and wood-boring organisms from all continents started to cross oceans on ship hulls (Carlton and Hodder, 1995; Wolff, 2005). In the Netherlands, for example, the invasion became possible in the early seventeenth century with the introduction of direct transportation on ship hulls into the country. However, neither the species nor the damage caused by it were observed until after the nineteenth century (Wolff, 2005). Thus, Limnoria was not mentioned in the references to the Dutch outbreak of the shipworm Teredo navalis around 1730 (Holthuis, 1956), nor was it mentioned by Vrolik et al. (1860). However, Hubrecht et al. (1893) made it clear that the latter authors had overlooked Limnoria. It was probably observed from 1834 and was most likely seen in Dutch waters between 1861 and 1862. The first clear observations were made in 1885-86 in the Westerschelde estuary, the Oosterschelde estuary, along the North Sea coast and in the western Wadden Sea (Hubrecht et al., 1893).
It was discovered in Scandinavia in 1799, in Britain in 1811 and in France in 1868 (Hubrecht et al., 1893). Nineteenth century British authors discussed its possible introduction from America, but concluded that Limnoria is a British species, as did some latter authors (Jones, 1963).
The history of introduction in the White Sea is given in Vinogradov et al. (2006). In 1908, L. lignorum was recorded in the Kolsky Bay, the most eastern point of distribution at the time. The first record for the northern area (Pirya Bay) of the White Sea is 1922, probably with the wooden ships during active navigation of the First World War. From this point L. lignorum crossed the Kandalaksha Bay and in 1936 was discovered on the opposite shore in the Kovda Bay, where they found plenty of underwater drown timber. At this time the crustacean was restricted to the Kovda Bay and found only at a depth of 7-9 m (salinity ~23‰), because of brackish (salinity 3-5‰) characteristics of the upper water level. After 1955, the building of a power station caused an increase of salinity up to 21‰ of surface water, which stimulated further dispersal of Limnoria. In 2000, it was found in 7 km from the point of finding in 1939. In 2005, the changing in the vertical distribution was recorded - it was found on all timber from the depth of 1.5 m and deeper.
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Netherlands||USA||19th C||Hitchhiker (pathway cause)||Yes||Wolff (2005)||First observation but may have been present from 17th C|
|Northern Russia||Europe||1908||Hitchhiker (pathway cause)||Yes||Vinogradov et al. (2006)||Introduced to the White Sea|
HabitatTop of page
This species is essentially a wood-boring organism, whose distribution is constrained by the availability of timber structures or drown wood. They mainly bore into the upper thin layer of wood and are often found boring into wood in the intertidal zone. The following variety of substrata where L. lignorum can be found is cited from Cookson (1991): many untreated timbers, wood and piling (Jones, 1963; Kussakin, 1963; Bohn and Walden, 1970); sawn Pinus sylvestris (Jones et al., 1972); unpreserved piles and regions of piles (Richards and Belmore, 1976); lightly creosoted piling (Vind and Hochman, 1961); water-logged stalks of the plant Rumex (Somme, 1940); untreated Alstonia scholaris (Eaton, 1989); light attack on creosote- and ammoniacal copper arsenate – treated pine (Baechler et al., 1970); creosoted timbers (Stevenson, 1862) and greenheart (Stevenson, 1874). Limnoria avoids (Iljin, 1992) iron and rust components in the wood. Softer parts of woods, such as summer growth rings, are destroyed more quickly than others (branches, bark, winter growth rings).
Destructive activity of L. lignorum varies depending on vertical distribution. Menzies (1957) noted that attack was greatest at the bottom rather than at the surface, and this was regardless of the magnitude of the seasonal salinity fluctuations. Similarly, in Norway (Santhakumaran and Sneli, 1984) attack by L. lignorum was more severe near the bottom and the intensity decreased rapidly a few meters above the bottom. Only very sparse settlement was observed at higher levels. A similar pattern of vertical distribution has also been reported by earlier authors (Johnson and Miller 1935; Somme, 1940; Black and Elsey 1948; Nair, 1962). In 1950-1960, there was a significant increase in the damage to wood by L. lignorum in the Barents sea (Kola Bay); the velocity of destruction increasing from the south to the north of the bay. The most active species were in the bottom (10-150 cm) layer of water (Iljin, 1992).
Habitat ListTop of page
|Intertidal zone||Principal habitat||Harmful (pest or invasive)|
|Intertidal zone||Principal habitat||Natural|
Biology and EcologyTop of page
The genetics of L. lignorum have not been widely investigated. Allozyme variation was studied by Henderson (1982) for twelve presumptive gene loci within and between four species in the genus Limnoria. Populations of L. lignorum from the Atlantic and Pacific Oceans, and from opposite sides of the Pacific Ocean, were genetically differentiated. The 18S rRNA gene of a close species L. quadripunctata was sequenced as part of a study of isopod phylogeny by Dreyer and Waegele (2002). The potential of polymorphic microsatellites for examination of the geographical distribution of the genetic diversity of Limnoria was investigated by Haye and Marchant (2007) in Chilean waters. The research revealed a high level of variability at most studied microsatellite loci of Limnoria; a model invertebrate species for the study of marine brooders with high potential of dispersal through rafting. These polymorphic microsatellite loci are to be useful for the study of the geographical distribution of the genetic diversity of this species of Limnoria.
These wood-boring organisms are generally found in heterosexual pairs. Females tend to position themselves at the head of a tunnel with the male behind (Henderson, 1924). Eltringham and Hockley (1961a) suggested that this position of the female might have some significance in copulatory behaviour. It is most likely that the sexes make contact by entering the burrow through the external openings or through the openings on the side walls.
Jones (1963) noted that reproduction of L. lignorum starts at a sea temperature of 8-9ºC and continues until about 14-16ºC. The number of eggs laid can be from 10 to 23 eggs, but numbers vary from region to region (Johnson, 1935; Somme, 1940). The eggs are incubated in a brood pouch formed of four pairs of overlapping oostegites. Once hatched, parents (usually the females) remain in the inner regions of the burrows to protect their brood (Henderson, 1924; Thiel, 2003). Juveniles are thus well sheltered from any adverse outside influences and benefit from this form of extended parental care.
When about 1.0-1.2 mm long the young escape and immediately begin burrowing into the wood where they were born (Johnson, 1935). Therefore, although the reproductive power of this species is relatively small, the hazards of a helpless pelagic existence are eliminated and the total population can increase rapidly. An attack once begun is continued by successive generations (Shino, 1950). In the early stages of attack the species produce burrows running parallel to the surface 1-2 mm below it. The burrow is circular in cross-section and very narrow so that the animal within cannot turn back. If more than one individual enters through the same entrance, they will produce a number of inter-connected burrows, each branch made by a separate individual. The boring activity of young limnorids is usually intensive in its early stages but after about five weeks, activity falls to a lower constant rate (Eltringham, 1961a).
Physiology and Phenology
George (1966) found that L. lignorum does not have a glycogen reserve sufficient to meet prolonged periods of anaerobiosis, so the infestation zone of Limnoria is restricted to the surface of submerged wood.
Some preliminary experiments (Eltringham, 1964) showed the ability of L. lignorum to control over its blood concentrations: blood concentration fell as soon as the animal was introduced to the reduced salinity and levelled off at the hyperosmotic value within a few hours. There was also some evidence of a periodicity in the osmoregulation.
Main migration season and infestation are found to be associated with breeding. Reproductive seasons and consequently infestation patterns vary depending on location. Near its southern limit in England (Naylor, 1972), the species breeds mainly in winter, but it can be from November to May or June. In Japan, Tsunoda and Nishimoto (1976) found that the maximum settlement was from June, when the water temperatures are around 20ºC, and continued for nine months, with a maximum in September. In the Northeastern Atlantic (Friday Harbor, Washington) Johnson (1935) found gravid females throughout the year, with a maximum settlement in April and May and a minimum in January and February, when temperatures were at their lowest for the year. Consequently, the majority of the total yearly settlement occurred over a six-month period from January to July, with maximum in spring. The most active settlement took place during a temperature range of 7.71-9.42°C (Johnson, 1935). Vind and Hochman (1961) concluded that timbers exposed to a marine environment in cold water harbors in the Pacific were attacked by L. lignorum in midsummer only. In Norway, attack by the species was rather erratic on panels immersed for short periods. On long-term panels the species were observed in appreciable numbers, causing damage to the surface of the timber. Thus, migratory activity was noticed during March to May 1977 and also from October 1977 to February 1978, when fresh attack on short-term monthly panels was observed (Santhakumaran and Sneli, 1984).
Population Size and Density
In British waters Limnoria species were present in 7% of the timbers surveyed (Oevering et al., 2001). In the White Sea, Vinogradov et al. (2006) observed up to 10 tunnels of L. lignorum on 1 cm2 of the surface. The tunnels were 5-7 mm deep and did not go further than a rotten layer of wood. Density of the population was about 20 individuals per 10 cm2. Heavily infected wood was found by Johnson (1935) in Friday Harbor, Washington: nearly 400 individuals (juveniles predominating) present in one cubic inch of wood.
L. lignorum is an essentially wood-consuming crustacean. Cellulose is degraded during gut passage to produce glucose but the exact mechanism of this digestion is yet to be studied in detail.
The crustacean starts to feed on wood after the wood has been submerged for some time, and so the wood has already been affected by other wood-degrading organisms. It has been reported that the cellulolytic activities of bacteria and fungi may play an important role in the wood degrading process (Daniel et al., 1991; El-Shanshoury et al., 1994). The microorganisms are ingested by Limnoria and are thought to be an extra source of nitrogen, which is beneficial for the crustacean given that they feed on a carbon-rich substrate containing very little native nitrogen (Cragg et al., 1999). Active feeding on fungal hyphae has also been suggested (Seifert, 1964). Large numbers of bacterial cells have been found associated with the outer exoskeleton surfaces of each individual L. lignorum, the most common being Aeromonas hydrophila, Pseudomonas and Vibrio. Therefore, limnorids may also consume microorganisms removed from the exoskeleton during grooming (Boyle and Mitchell, 1981a,b).
It is suggested that limnorids do not rely on internal symbiotic microbes to help digest lignocellulose (Fahrenbach, 1959; Boyle and Mitchell, 1978a,b; 1981a,b; Cragg et al., 1999). The absence of resident microbes in the digestive tract suggests that they themselves produce all the enzymes necessary for lignocellulose digestion. Ray and Julian (1952) observed that the cells of the intestinal diverticula of Limnoria produce cellulase, so that the species are able to derive their nutritional requirements by assimilation of products resulted from endogenous cellulolytic enzyme digesting activity. The source and functioning of lignocellulose degrading enzymes remains to be demonstrated. King et al. (2010) suggested that limnorid GH7 genes may be important for the efficient digestion of lignocellulose in the absence of gut microbes.
Limnorid colonies are frequently associated with other boring organisms. Kofoid and Miller (1927) described deceleration of wood degradation in Teredo-Limnoria association. Settlement of Teredo larva is inhibited by the “spongy” surface of Limnoria infested wood. As a result wooden piles were destroyed for longer periods compared to the “Teredo-only” population. Similarly, association with wood-boring chelurid amphipods may slow down wood degradation, because Chelura terebrans, inhabiting limnorids tunnels, prevent effective oxygenculation (Iljin, 1992). Marine fouling also restricts limnorid activity and slows down wood degradation (Weiss, 1948).
The tunnelling activity of L. lignorum often attracts a range of organisms, some of which benefit from the food particles produced by Limnoria to ventilate their burrows. These include the associate ciliates, such as Lagenophrys limnoriae (Clamp, 1988) and Pachyfolliculina gunneri (Brunel, 1963); marine tardigrades (Cantacuzene, 1951), small crustaceans (Brunel, 1963; Boer, 1971; Becker, 1971; Holmes and Jeal, 1987) and annelids (Reish, 1954).
L. lignorum is often observed in association with other species of Limnoria engaged in interspecific competition. Within mixed species populations inhabiting the UK coast, L. lignorum is the least tolerant to environmental changes and has been seen occupying the lower level on vertical piling with the more temperate L. quadripunctata occupying the middle level and the warm-water species L. tripunctata at the upper limit of attack (Eltringham, 1961a,b). In areas free from competition, such as on the north-west and east coast of the UK, L. lignorum may reach the intertidal zone where exposure is 60-65% but in other areas it may be excluded from the intertidal zone due to the competition (Jones, 1963).
Temperature is revealed as one of the main factors affecting L. lignorum activity and distribution. The species can withstand temperatures of up to 20°C, possibly higher by some reports (Becker, 1954). However, Eltringham (1965) found that at 15°C most L. lignorum individuals died earlier than individuals of another species, L. tripunctata. Kudinova-Pasternak (1971) found that in the White Sea a low temperature of 3-4°C caused death and eggs required over 10°C to mature. Somme (1940) reported embryonic development of L. lignorum from Norway to be retarded when temperatures fell below 6°C. At 6°C the entire incubation period was estimated to be one month longer than the usual incubation period of two months at higher temperatures. It was suggested that at freezing, the brood might survive only in a more or less dormant state. Accordingly, Menzies (1957) concluded that the 0ºC isotherm surrounding the Arctic Ocean was the northern limit of the species distribution. The effect of temperature on vertical distribution is known from studies of L. lignorum in the White Sea, where temperature limits distribution to no deeper than 10-20 m. In the Sea of Japan the lower water temperature limits L. lignorum to the surface water level (Iljin, 1992).
Eltringham (1961a) investigated the effect of salinity upon the wood-boring activity of Limnoria. Low salinity was the direct cause of reduced boring activity and boring ceased below 10‰. Over 15-20 days a salinity of 6‰ proved fatal for Limnoria. In freshwater L. lignorum cannot usually survive for more than 36 hours (Menzies, 1957). Eltringham (1961a) found no reduction in survival in concentrated salinities up to a value of 48‰, the highest salinity examined. Although no long-term experiments were conducted, generally, areas with uniformly low salinities (below 10‰) and those having wide ranges of salinity (0-35‰) are considered as unfavorable for the development and establishment of Limnoria (Menzies, 1957). The effects of salinity on wood degradation have been shown in the White and Barents Sea where wood degradation increased with distance from the river mouths. Limnoria can survive without water for about 24 hours and are not affected by a temporary aeration (Iljin, 1992).
Ryabchikov (1957) observed that ice creates favourable conditions for Limnoria in the Sea of Japan. The ice protects the species from the low temperatures in winter and also rubs the upper layer of wood, improving access to newer wood stimulating infestation. The ability of L. lignorum to withstand freezing for 24 hours (Somme, 1940) and regular periods during low tides (Ryabchikov, 1957) is also described.
A water current of more than 1 m/sec prevents Limnoria larva from settlement (Kudinova-Pasternak, 1971); whereas the absence of water current obstructs oxygenation and Limnoria leaves the habitat (Eltringham and Hockley, 1961b; Kudinova-Pasternak, 1971).
Menzies (1957) found that on average dissolved oxygen content below 1.6 ppm might constitute a limiting factor in the survival of Limnoria. The tolerance to low dissolved oxygen concentrations at different temperatures was tested by Anderson and Reish (1967). For L. lignorum 28-day median tolerance limits were 1.0 mg/l of dissolved oxygen at 15-16°C and 19-20°C. The amount of burrowing activity was directly related to the amount of dissolved oxygen, sharply reducing at dissolved oxygen concentrations below 3.0 mg/l.
ClimateTop of page
|C - Temperate/Mesothermal climate||Preferred||Average temp. of coldest month > 0°C and < 18°C, mean warmest month > 10°C|
|D - Continental/Microthermal climate||Preferred||Continental/Microthermal climate (Average temp. of coldest month < 0°C, mean warmest month > 10°C)|
|E - Polar climate||Tolerated||Polar climate (Average temp. of warmest month < 10°C)|
Latitude/Altitude RangesTop of page
|Latitude North (°N)||Latitude South (°S)||Altitude Lower (m)||Altitude Upper (m)|
Water TolerancesTop of page
|Parameter||Minimum Value||Maximum Value||Typical Value||Status||Life Stage||Notes|
|Dissolved oxygen (mg/l)||3||Optimum|
|Dissolved oxygen (mg/l)||1||Harmful|
|Salinity (part per thousand)||6||15||Harmful|
|Salinity (part per thousand)||12||16||Optimum|
|Water temperature (ºC temperature)||0||20||Harmful|
|Water temperature (ºC temperature)||8||16||Optimum|
Natural enemiesTop of page
Notes on Natural EnemiesTop of page
Some species of polychaete, such as Nereis vexillosa and Halosydna johnsoni, are known to predate on limnorids(Reish, 1954). Fungal infection was also observed by Iljin (1992) but this has not been explored in detail.
Means of Movement and DispersalTop of page
The spread of L. lignorum should be seen as a combination of active and passive migration. Limnoria is not a strong swimmer and does not possess the pelagic egg or larval stage allowing wide and rapid dispersal. Instead, regular seasonal migration, associated with the main breeding season may be the principal means of dispersal within an infected area (Johnson, 1935; Eltringham and Hockley, 1961a). These migrations, however, occur over a few meters or less (Vinogradov et al., 2006). Local current regimes may impact significantly on dispersal (Thiel and Haye, 2006; Cragg et al., 2009). Thus, water currents are very important for the distribution of the Limnoria larvae (Ryabchikov, 1957).
The spread of infection to more remote areas occurs through mechanical transportation by drifting of infected wood (Thiel and Haye, 2006; Vinogradov et al., 2006). The ecology of rafting is known mainly for kelp-dwelling limnorids, but may be similar for wood-rafting. Miranda and Thiel (2008) hypothesized that limnorids could successfully reproduce during rafting journeys facilitating long-distance dispersal. Nikula et al. (2010) support rafting as the most plausible recolonization mechanism, and which may explain similarities in the species composition of intertidal marine communities across the sub Antarctic.
L. lignorum can be easily transported in the hulls of wooden boats and ships (Iljin, 1992; Vinogradov et al., 2006). Shino (1950) commented that distribution of the species may be affected more by the navigation routes of wooden ships, rather than environmental restrictions. The abundance of wood in the intertidal zone, such as piling, piers and drift wood may also support establishing populations.
Pathway VectorsTop of page
|Floating vegetation and debris||Drift of infected wood||Yes||Yes||Thiel and Haye, 2006; Vinogradov et al., 2006|
|Ship hull fouling||Yes||Yes||Iljin, 1992; Shino, 1950; Vinogradov et al., 2006|
|Water||Dispersal of larva and adults with local water currents||Yes||Yes||Cragg et al., 1999; Ryabchikov, 1957; Thiel and Haye, 2006|
Impact SummaryTop of page
|Economic/livelihood||Positive and negative|
Economic ImpactTop of page
Since the nineteenth century (Hoek, 1893) the damage caused by this species to wooden structures has been considered as a serious economic issue and since the mid-twentieth century the species has been viewed as a serious pest. In many areas, such as Kamchatka (Avacha Bay), the Barents Sea (Kola Bay), Vladivostok and the Sea of Japan (De-Casri Bay) L. lignorum has been reported as destroying wooden constructions at a rate of 0.4-1.4 cm per year (Ryabchikov, 1957; Kussakin, 1963; Iljin, 1992). In the Sea of Okhotsk, it was estimated that the combined effect of L. lignorum and L. magadanensis were destroying wooden constructions within three years.
The associated cost of replacing wooden structures is usually significant. The Los Angeles Harbor Department Testing Laboratory estimated that borers, mainly Limnoria, caused about one million dollars worth of damage each year (Menzies, 1957). In America, the cost of replacing a seawall and its wooden supports along the Seattle waterfront was around $700 million (Roach, 2004).
Environmental ImpactTop of page
Impact on Habitats
Limnorids process wood debris, releasing energy stored in submerged wood and drift wood. Colonies create a complex of interconnecting tunnels, which usually have a series of pinhole-sized punctures along their length, are about 1 mm in diameter and may be found just below the surface of the infected wood. These tunnels are numerous and give the wood a characteristic lace-like appearance and a sponge-like structure.
The use of anti-fouling compounds to control L. lignorum may have unwanted side effects on the environment.
Impact on Biodiversity
The tunnels of the borer create niches for other organisms including annelids, marine tardigrades and other small crustaceans (Cantacuzene, 1951; Reish, 1954; Brunel, 1963; Boer, 1971; Becker, 1971; Holmes and Jeal, 1987).
L. lignorum may compete with other species for space, including other Limnoria species (Eltringham, 1961a,b).
The use of anti-fouling compounds to control the crustacean may negatively impact upon other species and the native biodiversity.
Many protozoa and other organisms associated with Limnoria are likely to have been transported around the world and thus introduced to new environments.
Social ImpactTop of page
The damage caused by the crustacean may result in unexpected failures of wooden constructions, which may also lead to health and safety risks. Damage to constructions of important historical or heritage value has also been a major negative effect of the organism.
Risk and Impact FactorsTop of page Invasiveness
- Invasive in its native range
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Fast growing
- Has high reproductive potential
- Negatively impacts aquaculture/fisheries
- Negatively impacts tourism
- Reduced amenity values
- Highly likely to be transported internationally accidentally
- Difficult to identify/detect in the field
UsesTop of page
In the White Sea, Limnoria activity disposed of submerged waste and drift wood from timber mills. Therefore, their activity was viewed positively as it cleared the bottom of the Kovda Bay (Vinogradov et al., 2006).
The mechanism of digestion of lignocellulose and hemicelluloses by Limnoria is attracting attention both in terms of basic research into metabolism by microorganisms, and also as a means of converting plant biomass into bio-fuels, which may be exploited as a source of energy (King et al., 2010). Recent research has identified the structure and function of a cellulase enzyme with some unusual properties (Kern et al., 2013). It is hoped that the robust nature of this enzyme, which allows it to be compatible with sea water, will lower the costs of making biofuels and make it a more cost effective alternative to other energy sources. Costs can be reduced further by reproducing the enzyme on an industrial scale using an industrial microbe that can produce it in large quantities; in a similar way to how enzymes for biological washing powders are produced (BBSRC, 2013).
Uses ListTop of page
- Miscellaneous fuels
- Research model
Similarities to Other Species/ConditionsTop of page
Although similar, Limnoria can be distinguished from the genus Paralimnoria by its uropods, which have a much shorter exopod than endopod, and only the endopod has a claw.
More than 50 species of Limnoria have been described, some of them morphologically very similar. This can cause identification diffculties in the field but inspection of tunnelled wood and microscopic examination can determine if limnorid infestation is taking place and by which species. Holthuis (1949) gives details of the differences between L. lignorum and the similar species Limnoria quadripunctata. The differences between the European species of Limnoria may be found in Castello (2011).
Prevention and ControlTop of page
Chemical control is often used to protect against Limnonia attack on submerged timbers. Creosote or a mixture of creosote, copper chromium and arsenic salts are the most popular chemicals to use (Eaton, 1989; Cookson, 2007). Nylinder-Norman et al. (1974) reported that treatment with Zn-Cr-As extended the service life of Pinus sylvestris panels from 2.5 to 26 years. In Sweden, chemical treatments with copper components were most effective against L. lignorum (Henningsson and Norman, 1979). Some methods of chemical wood protection are also discussed in Eaton and Cragg (1996) and Eaton and Hale (1993). All these methods may be used as a temporary measure but most have unwanted side-effects.
A number of other methods have been suggested for the prevention and control of this species. Henderson (1998) and Cragg et al. (1999) suggested biological control where factors derived from conspecifics and wood-inhabiting microorganisms could limit Limnoria settlement. Bjoerdal and Nilsson (2008) showed that reburial of shipwrecks in marine sediment could be used as a simple and efficient method for long-term preservation of these historical events. Iljin (1992) pointed out that the right choice of the depth, location and engineering of a construction could significantly decrease wood-borers' damage. Economically feasible and effective means of protecting timber piling is to create a physical barrier, such as ultraviolet-resistant PVC sheathing. Norman (1976) found that bark provided good protection against attack; wood weight loss caused by Limnoria of unbarked logs was 17-52% verses only 6-18% for barked logs.
Regular inspection of wooden structures in the intertidal zone with an aim of early detection of infestation could reduce the effects of L. lignorum. Furthermore, disposing of infested wood on dry land rather than dumping at sea and controlling the movements of infested boats could help prevent the spread of the borer.
Gaps in Knowledge/Research NeedsTop of page
Much remains unknown in terms of the phenomena, patterns, and processes of invasions of the genus in general. Further study of intraregional human-mediated dispersal vectors and experimental ecology of invasions are needed in order to establish more effective management plans (Cohen and Carlton, 1995). In addition, research focusing on the nutrition, migratory and substrate seeking behaviour of L. lignorum may provide key information beneficial for their control (Cragg et al., 1999; Cragg, 2003).
The source or sources of wood-degrading enzymes that permit digestion of wood particles by Limnoria requires further study, most likely using techniques of modern molecular biology. Furthermore, the role of wood degrading bacteria and fungi associated with tunnelling activity of Limnoria remains to be fully explained.
ReferencesTop of page
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ContributorsTop of page
10/10/12 Original text by:
Ekaterina Shalaeva, South Croydon, Greater London, UK
Distribution MapsTop of page
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