Invasive Species Compendium

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Limnoria quadripunctata
(gribble)

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Datasheet

Limnoria quadripunctata (gribble)

Summary

  • Last modified
  • 25 September 2018
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Limnoria quadripunctata
  • Preferred Common Name
  • gribble
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Crustacea
  •         Class: Malacostraca
  • Summary of Invasiveness
  • Limnoria quadripunctata is a wood-boring isopod. Limnoriids generally remain within their burrows in wood, seagrasses or algal hold-fasts, but episodes of probably short-distance migratory behaviour have been o...

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Pictures

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PictureTitleCaptionCopyright
Limnoria quadripunctata, male and female, ventral view. Moustérian, Golfe du Morbihan, Brittany, France. 03/07/2008.
TitleMale and female
CaptionLimnoria quadripunctata, male and female, ventral view. Moustérian, Golfe du Morbihan, Brittany, France. 03/07/2008.
Copyright©Auguste Le Roux-2008: CC BY 3.0
Limnoria quadripunctata, male and female, ventral view. Moustérian, Golfe du Morbihan, Brittany, France. 03/07/2008.
Male and femaleLimnoria quadripunctata, male and female, ventral view. Moustérian, Golfe du Morbihan, Brittany, France. 03/07/2008.©Auguste Le Roux-2008: CC BY 3.0

Identity

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Preferred Scientific Name

  • Limnoria quadripunctata Holthuis, 1949

Preferred Common Name

  • gribble

Local Common Names

  • Germany: Holzbohrassel; Seepocken

Summary of Invasiveness

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Limnoria quadripunctata is a wood-boring isopod. Limnoriids generally remain within their burrows in wood, seagrasses or algal hold-fasts, but episodes of probably short-distance migratory behaviour have been observed (Eltringham and Hockley, 1961; Miranda and Thiel, 2008). Dispersal over longer distances is likely to take place by rafting within the substrate (Thiel and Haye, 2006; Miranda and Thiel, 2008). During the period of large scale trans-ocean transport using wooden ships, limnoriids may have been dispersed in the hulls of these ships. The present distribution of L. quadripunctata in temperate waters of South and North America, Europe, southern Africa and Australasia, but not in tropical waters, is suggestive of invasion from the northern to the southern hemisphere or vice-versa. The status of the species in western North America is not clear, though it is recorded in documentation of non-native species of California (Boyd et al., 2002). In South Africa, it is considered to satisfy the criteria for a cryptogenic species (Robinson et al., 2005). This species infests wooden structures in the intertidal zone (Barnacle, 1987) and as such is at least locally invasive.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Crustacea
  •                 Class: Malacostraca
  •                     Subclass: Eumalacostraca
  •                         Order: Isopoda
  •                             Family: Limnoriidae
  •                                 Genus: Limnoria
  •                                     Species: Limnoria quadripunctata

Notes on Taxonomy and Nomenclature

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The family Limnoriidae consists of small isopods which tunnel into wood, seagrasses or macroalgae. It contains the genera Limnoria, Paralimnoria and Lynseia(Cookson and Poore, 1994). Cookson (1991) reported 51 species within the family and since the publication of his monograph, two new species have been added: an algal borer (Cookson, 1997) and a seagrass borer (Cookson and Lorenti, 2001). Limnoria quadripunctata was described by Holthuis from specimens collected in the Netherlands (Holthuis, 1949). A rather similar species, Limnoria carinata, has subsequently been described from the Mediterranean (Menzies and Becker, 1957). However, it is not clear whether the features used to distinguish the species are consistently distinct and so further anatomical investigations are required which may prove L. carinata to be synonymous with L. quadripunctata (Kühne, 1971; Cookson, 1991), in which case the specific epithet quadripunctata has priority.

Description

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Animals are up to 4 mm in length (Antezana, 1968), though more typically 3 mm or less (Cookson, 1991; Borges et al., 2009). The anatomy of Limnoria quadripunctata (see Pictures) follows the typical isopod pattern, with three main body regions: the cephalon, peraeon and pleon. The peraeon bears seven pairs of walking limbs (peraeopods) and the pleon bears five pairs of flattened swimming/respiratory appendages (pleopods) while the pleotelson carries a pair of uropods.

The uropods have a much shorter exopod than endopod, and only the endopod has a claw. These features distinguish the genus Limnoria from Paralimnoria (Cookson, 1991).

The upper surface of the pleotelson has an anteromedial, square array of four punctae, which is characteristic of the species. The punctae are visible when the telson is viewed with a focused light beam directed from the side, but scanning electron microscopy of carefully cleaned specimens is recommended for accurate identification (see Pictures).

 

Distribution

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The species has been reported initially from driftwood on the coast of the Netherlands (Holthuis, 1949) but, in view of the nature of the substrate, the origin of the population studied by Holthuis is not clear. Holthuis pointed out that previous reports of the distribution of wood-boring limnoriids tended to assume that the organisms found were L. lignorum, but that illustrations given of specimens from San Francisco Bay indicate the presence of L. quadripunctata there (Kofoid and Miller, 1927). L. quadripunctata has subsequently been reported from other sites on the coasts of northwestern Europe: Lough Hyne, southwestern Ireland (de Grave and Holmes, 1998); southern UK (Jones, 1963; Oevering et al., 2001), western France (Jones et al., 1972) and Portugal (Borges et al., 2008). It has also been reported from Trieste at the northern end of the Adriatic Sea by authors who report the very similar L. carinata from the west coast of Italy (Menzies and Becker, 1957).

The species has also been recorded on the west coast of the USA, from Humboldt Bay to San Diego (Menzies, 1957; Carlton, 1975; Espinosa-Perez and Hendrickx, 2006), but is apparently absent on the east coast. It occurs in Chile (Antezana, 1968), New Zealand (McQuire, 1964), southern Australia (Cookson, 1991), the southern Indian Ocean Islands of Amsterdam and St Paul (Kensley, 1976) and between Table Bay and Port Elizabeth in South Africa (Stebbing, 1910; Robinson et al., 2005).

Distribution Table

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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/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Africa

South AfricaWidespreadIntroduced Invasive Stebbing, 1910; Robinson et al., 2005Between Table Bay and Port Elizabeth

North America

USAPresentPresent based on regional distribution.
-CaliforniaWidespreadIntroduced Invasive Kofoid and Miller, 1927; Carlton, 1975

South America

ChilePresentIntroduced Not invasive Antezana, 1968Insufficient information to determine status, though more recent studies do not mention this species

Europe

FranceWidespreadNative Not invasive Jones et al., 1972
IrelandLocalised Not invasive Grave and Holmes, 1998Limited information on distribution in Ireland
ItalyLocalised Not invasive Menzies and Becker, 1957Taxonomic status of Mediterranean L. quadripunctata and L. carinata unclear
NetherlandsPresentNative Not invasive Holthuis, 1949
PortugalPresentNative Not invasive Borges et al., 2008
UKWidespreadNative Not invasive Oevering et al., 2001; Borges et al., 2008

Oceania

AustraliaPresentIntroducedCookson, 1991
-New South WalesPresentIntroduced Invasive Barnacle, 1987
-South AustraliaPresentIntroducedCookson, 1991
-TasmaniaPresentIntroducedBarnacle, 1987
-VictoriaPresentIntroducedCookson, 1991
-Western AustraliaPresentIntroducedCookson, 1991
New ZealandPresentIntroducedMcQuire, 1964

History of Introduction and Spread

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The introduction of L. quadripunctata to the coast of California may have resulted from the abandoning of infested wooden sailing ships originating from Atlantic ports during the gold rush that started in 1849 (Carlton, 1975). The introduction to South Africa could have taken place earlier, though the vector of spread is likely to have been the same (Griffiths et al., 2009); wooden sailing ships from Europe would have been provisioning along the coast where L. quadripunctata is found since the early Portuguese, Dutch and British voyages of exploration and trade. Australia and New Zealand are also likely to have been initially colonized by populations established in wooden sailing vessels.

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Australia   Yes Cookson (1991); Griffiths et al. (2009) Wooden boats suspected to be introduced from UK
California 1850s Yes Carlton (1975) Gold rush shipping
South Africa   Yes Wooden boats

Risk of Introduction

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Further spread would probably be reliant on populations in driftwood rather than wooden boats. This species may replace Limnoria lignorum if there is significant warming of coastal waters either due to climate change or to local factors such as release of power station cooling waters.

Habitat

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Wood-boring limnoriids in temperate waters are principally found in wooden structures. Their natural habitat in such waters would probably have been fallen trees and branches brought down to the sea by rivers, but the extensive modification of temperate flood plain ecosystems has markedly reduced the quantities of this sort of substrate entering the sea. Thus the current distribution of L. quadripunctata is constrained by the availability of timber structures.

The world-wide distribution of L. quadripunctata indicates that it is limited to temperate waters. In the UK, for instance, it is restricted to the southern half of the country (Jones, 1963; Oevering et al., 2001).

More locally, salinity also limits colonization; the distribution of L. quadripuncata indicates only limited tolerance of brackish waters. Tunnelling by Limnoria quadripunctata is focused in the intertidal zone (Jones, 1963; Eltringham, 1971), though specimens have been collected from as deep as 30 m (Cookson, 1991).

Habitat List

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CategoryHabitatPresenceStatus
Littoral
Intertidal zone Principal habitat Natural

Biology and Ecology

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Genetics
 
The genetics of L. quadripuncata have not been extensively investigated. The 18S rRNA gene was sequenced as part of a study of isopod phylogeny by Dreyer and Waegele (2002). The potential of microsatellites for examination of populations of limnoriids that was pioneered in Chilean waters (Haye and Marchant, 2007) is now under evaluation in the UK (G Zouganelis, University of Portsmouth, UK, personal communication, 2010).
 
Reproductive Biology
 
Limnoriid life history has been most intensively studied with Limnoria lignorum(Sømme, 1941) and L. tripunctata (Menzies, 1954), but certain life history aspects have been studied with L. quadripunctata. The effects of temperature on development rate have been studied with L. quadripuncata, L. tripunctata and L. lignorum (Eltringham, 1967). Temperature may also affect brood size as L. quadripunctata has a brood size maximum of 17 with a mean of 9.5, which is intermediate between the larger brood sizes of the colder water species L. lignorum and the smaller brood sizes of tropical species (Menzies, 1954). As benthic organisms that inhabit a niche that is slowly destroyed during feeding, limnoriids require a means of dispersal in order to locate new food sources. In Southampton Water on the south coast of the UK, L. quadripuncata juveniles and young males has been shown to migrate (Eltringham and Hockley, 1961).
 
Adult limnoriids are commonly found in pairs with the female at the head of a tunnel and the male behind. Although adults normally stay within their burrows, they have been found (under experimental conditions which would require precise, non-random movement) to be capable of moving to form pairs (G Malyon, University of Portsmouth, UK, personal communication, 2010). Juveniles are retained within parental burrows before establishing their own burrows which often branch off from the parental burrow. The extent of the period of parental care may decrease with increasing population density (Thiel, 2003).
          
Physiology and Phenology
 
The physiology of L. quadripunctata has not been investigated to any great degree. There have been studies of the effects of temperature and salinity on various functions of limnoriids including L. quadripuncata(Eltringham, 1961, 1965 ; Anderson and Reish, 1967). The tolerance of low oxygen levels is a valuable attribute for an organism that lives in a burrow. L. quadripunctata has a lower tolerance limit of oxygen level of between 1 .0 and 0.6 mg l-1, but shows markedly lowered feeding at less than 3 mg l-1 (Eltringham, 1961; Anderson and Reish, 1967).
 
Nutrition
 
Wood consists of cellulose in crystalline arrays bound with hemicelluloses and lignin in a complex known as lignocellulose. Cellulose and hemicelluloses are broken down during digestion to yield glucose that can be exploited as a source of energy. While L. quadripunctata is able to feed on wood alone (Borges et al., 2009), wood-boring limnoriids also consume micro-organisms inhabiting the wood they are feeding on (Daniel et al., 1991). They may also consume microorganisms removed from the exoskeleton during grooming (Boyle and Mitchell, 1981). Ingested microorganisms may provide organic nitrogen compounds that are in very short supply in wood alone.
 
Associations
 
The tunnelling activity of L. quadripunctata creates an environment favourable for a range of sessile ciliates, each type of which attaches selectively to specific parts of the host animal (Delgery et al., 2006). Most striking of these is the dark green Mirofolliculina limnoriae, which is visible to the naked eye and which interacts with the moult cycle of L. quadripunctata(Delgery et al., 2006).
 
Wood-boring chelurid amphipods are frequently found associated with limnoriid colonies. Chelura terebrans is the species that occurs with L. quadripunctata. These two species have been found in association in South Africa (Robinson et al., 2005), California (Barnard, 1950), Portugal (Borges et al., 2008) and the UK (S Cragg, University of Portsmouth, UK, personal communication, 2010).
 
The tunnels of Limnoria provide conditions suitable for various invertebrates, in particular small crustaceans (Becker, 1971) and annelids (Reish, 1954).
 
Environmental Requirements
 
Water temperature has been recognised as an important factor in determining the distribution of L. quadripunctata. As the organism spends a significant part of the day out of the water at low tide, air temperature may also be a limiting factor, but this has not been investigated. Water temperature affects boring activity and survival (Eltringham, 1965) and life history characteristics, especially egg development (Eltringham, 1967). The distribution of L. quadripunctata indicates that this species requires temperate to warm temperate conditions.

L. quadripunctata is found at full salinity sites and appears to be restricted to marine rather than brackish waters. Lowered salinity is known to affect various aspects of the life history of L. quadripunctata(Eltringham, 1961, 1964).

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Depth (m b.s.l.) Optimum Low intertidal, has been found as deep as 30 m, but normally restricted to the intertidal zone
Dissolved oxygen (mg/l) Optimum Position in the intertidal zone means oxygen levels normally high. Lowest tolerated level about 0.6-1.0 mg/l; feeding markedly reduced below 3 mg/l.
Salinity (part per thousand) Optimum Not tolerant of estuarine conditions
Water temperature (ºC temperature) Optimum Temperate to warm temperate

Notes on Natural Enemies

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There are no known natural enemies that regulate populations of limnoriids (Becker, 1971), but certain polychaete worms are known to predate on them (Reish, 1954).

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

Surface currents are vital in dispersing organisms that colonize fallen wood (Thiel and Haye, 2006; Cragg et al., 2009). Thus the expansion of L. quadripunctata away from established populations will be constrained by local current regimes. Establishment of a population after natural dispersal requires a supply of wood either from fallen trees or human input.

Vector Transmission (Biotic)

Limited dispersal by the animals themselves does occur (Eltringham and Hockley, 1961), but wider dispersal is likely to occur in floating logs, as is suggested to be the case in the Gulf of Aqaba for L. tripunctata (Cragg et al., 2009).

Accidental Introduction

The current distribution of the species is highly suggestive of accidental introduction in the hulls of ocean-going wooden boats. This phase of dispersal has come to an end, but populations remain because of the continued availability of wood in the intertidal zone in the form of piers, wharves and piling.

Intentional Introduction

Intentional introduction of this species is not likely as there is no currently recognized economic benefit to be derived from its introduction. There are no documented cases of intentional introduction.

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Floating vegetation and debrisIntermittent dispersal of adults Yes Yes Thiel and Haye, 2006
Ship hull foulingColonies in hulls of wooden boats is seen as a likely means of dispersal Yes Yes Carlton, 1975

Impact Summary

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CategoryImpact
Cultural/amenity Negative
Economic/livelihood Negative
Environment (generally) Positive

Economic Impact

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Limnoriids are a major threat to wooden structures in the intertidal zone. Problems of attack by limnoriids on wood in Dutch dykes were sufficient to require a royal commission of investigation (Hoek, 1893). These problems were ascribed to the activities of L. lignorum, but at that time, the existence of L. quadripunctata and its occurrence on Dutch coasts had not been recognized. Over £400,000 was required to repair a Grade II-listed (heritage protection listing) wooden pier at Yarmouth on the Isle of Wight, UK, in 2007. This structure had been severely damaged by the activity of L. quadripunctata.

Environmental Impact

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Impact on Habitats

In natural environments, marine wood borers play an important role in processing woody debris and releasing energy stored in fallen wood from forests but also in driftwood.

Impact on Biodiversity

The tunnelling of limnoriids creates niches for other small invertebrates (Becker, 1971) and favourable conditions for associates such as the wood-consuming amphipod Cheluraterebrans which often occurs in the outer layers of wood where Limnoria is established (Kühne and Becker, 1964). The reported distributions of L.quadripunctata and Cheluraterebrans in South Africa coincide almost exactly (Robinson et al., 2005). These species also occur together in Los Angeles and San Francisco (Barnard, 1950). Even the animals themselves provide a niche for certain specialist organisms (Delgery et al., 2006).

Social Impact

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Colonization by L. quadripunctata imposes a requirement for maintenance of infrastructure such as wharves, piers and bridges. There is also the potential for impacts on aquaculture and capture fisheries where wooden components are exposed in the water. The costs are felt in direct economic terms but also in terms of loss of important capabilities during maintenance or as a result of unexpected failures of components.

Risk and Impact Factors

Top of page Invasiveness
  • Invasive in its native range
  • Proved invasive outside its native range
  • Has a broad native range
  • Benefits from human association (i.e. it is a human commensal)
  • Fast growing
  • Has high reproductive potential
  • Gregarious
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Difficult to identify/detect in the field
  • Difficult/costly to control

Uses

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Economic Value

These animals are generally viewed as having a negative economic impact; however, they have an unusual digestive system that may yield insights into how to exploit lignocellulosic biomass.

Social Benefit

Effective exploitation of the insights into the digestive process might lead to the development of alternative liquid biofuels.

Environmental Services

Limnoriids play a role in the detrital food chain, converting large refractory detritus into small rather more labile particles and thus hastening the release of stored energy and nutrients. They also create temporary niches for cryptofauna and thus enhance biodiversity.

Uses List

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Fuels

  • Biofuels

General

  • Research model

Genetic importance

  • Gene source
  • Test organisms (for pests and diseases)

Materials

  • Alcohol

Diagnosis

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The nature of the tunnels would enable them to be distinguished from those caused by other borers such as beetles. Limnoriid colonies form an interconnecting superficial complex of tunnels (approximately 1 mm in diameter) on or just below the surface of the infected wood. Tunnels just below the surface tend to have a series of pinhole-sized punctures along their length that improve circulation of water in the tunnel. Juvenile burrows extend at right angles to the parental burrow. L. quadripuncata can be distinguished from the other species likely to occur at the same site (L. lignorum and L. tripuncatata) by the four punctae on the pleotelson, but for a definitive identification, a number of specimens should be sent with full collection details to a specialist.

Detection and Inspection

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Superficial inspection of tunnelled wood followed by laboratory examination of animals would serve to determine whether limnoriid colonization was taking place and if so, by which species.

Similarities to Other Species/Conditions

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Limnoria andamanensis, like L. quadripunctata has an anteromedial array of four punctae on the pleotelson, but differs in the sculpturing on pleonite 5 and other features (Rao and Ganapati, 1969). L. carinata is reported to have similar sculpturing on the upper surface of the pleotelson to L. quadripunctata, though it may have six or more punctae. This species is said to differ from L. quadripunctata in having four rather than five articles on the flagellum of antenna 2 (Menzies and Becker, 1957). Subsequent publications have urged further investigation of the variation in morphological characteristics of L. quadripunctata and especially the possibility of sexual dimorphism, with a view to resolving whether these two species are genuinely distinct (Kühne, 1971; Cookson, 1991).

Prevention and Control

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Prevention
 
Measures need to be taken to prevent infested boats from entering harbours with wooden structures. Where possible, wood treated to prevent borer infestation should be used in initial construction and repairs.

Public awareness
 
Information on selection of raw material and on control measures to be taken against marine borers is available in a format that is usable by port engineers with responsibilities for wooden coastal structures. Cookson et al. (2007) for example reviewed the relative performance of a range of marine piling types. The impact of degradation by limnoriids may extend beyond the concerns of such professionals. The economic impact of damage to a major structure in a small town may require special fund-raising measures that in turn require the raising of public awareness (see Pictures).
 
Eradication
 
Eradication is generally not attempted except by replacement of timber. The effects of biocide sprays etc. could not be restricted to the infested timbers. Wood protection by means of impregnation with creosote and/or a mixture of copper chromium and arsenic salts (CCA) has been widely used (Cookson, 2007), but environmental legislation is restricting the use of these broad spectrum biocides and alternatives have yet to achieve widespread acceptance. Furthermore, L. quadripunctata achieves a degree of tolerance to CCA by sequestering excess copper in the hepatopancreas (Tupper et al., 2000). Contact toxins such as synthetic pyrethroids offer a possible alternative (Rutherford et al., 1979). Some naturally durable timbers are either toxic to L. quadripunctata, or markedly reduce feeding rates (Borges et al., 2008). If available in suitable quantities and dimensions, these timbers can be used as an alternative to biocide-impregnated timber. Also, they may give insights enabling the development of tailored, rather than broad spectrum biocides.
 
Control
 
Physical/mechanical control
 
Ultraviolet-resistant PVC sheathing has proved a commercially viable means of protecting timber piling. Such a physical barrier will prevent attack by wood-boring Limnoria species.
 
Movement control
 
The movements of infested boats should be tightly restricted and any disposal of infested wood should be to dry land, rather than by dumping at sea.
 
Biological control
 
No suitable agents of biological control have been identified.
 
Chemical control
 
While spraying with anti-arthropod biocides at low tide would probably give a measure of control over populations of Limnoria, this approach would not permit sufficient control over the biocide to prevent unwanted side-effects.
 
Monitoring and Surveillance
 
Regular inspection by port engineers of wooden structures in the intertidal zone using the features described in the Diagnosis section should permit early detection of infestation.
 
Mitigation
 
Currently, the effects of infestation are mitigated by the removal of infested wood to dry land.
 
Ecosystem Restoration

Ecosystem restoration is not an approach that seems relevant for limnoriid control, as the organisms exploit anthropogenic changes in the marine environment (installation of wooden structures etc.) but do not themselves bring about adverse environmental changes.

Gaps in Knowledge/Research Needs

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The uncertainty as to whether Limnoria carinata is a synonym of L. quadripunctata needs to be resolved by anatomical studies that take account of possible sexual dimorphism of pleotelson ornamentation as reported by Cookson (1991), and of within- and between-population variations. This area of uncertainty would also benefit from the application of an approach that compares DNA sequences, as has been shown with limnoriids of the Chilean coast (Haye and Marchant, 2007). This approach could also be used to give insights into the extent to which limnoriid populations are isolated. This would give indications of the extent of mobility of the species in the absence of the extensive movements of wooden ocean going ships.

Cellulase activity has been demonstrated in limnoriids (Ray and Comita, 1952) and the digestive enzyme systems of limnoriids have been shown to preferentially degrade cellulose and hemicelluloses in lignocellulose (Seifert, 1964). This was later shown to take place without any symbiotic gut-resident micro-organisms (Boyle and Mitchell, 1978). The wood digestion capabilities of Limnoria need to be probed using techniques of modern molecular biology.

References

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Anderson JW; Reish DJ, 1967. The effects of varied dissolved oxygen concentrations and temperature on the wood-boring isopod genus Limnoria. Marine Biology, 1(1):56-9.

Antezana T, 1968. [English title not available]. (Limnoria (Limnoria) quadripunctata Holthuis (Crustacea,Isopoda), nuevo exponente de la fauna del Pacifico Sur-Oriental.) Revista de Biologia Marina, 13:293-301.

Barnacle JE, 1987. Limnoria quadripunctata Holthuis - a review of its status as a marine wood borer. Material und Organismen, 22(1):7-23.

Barnard JL, 1950. The occurrence of Chelura terebrans Philippi in Los Angeles and San Francisco harbors. Bulletin, Southern California Academy of Sciences, 49(3):90-97.

Becker G, 1971. The biology, physiology and ecology of marine wood-boring crustaceans. In: Marine borers, fungi and fouling organisms of wood [ed. by Jones, E. B. G. \Eltringham, S. K.]. Paris, France: OECD.

Borges LMS; Cragg SM; Bergot J; Williams JR; Shayler B; Sawyer GS, 2008. Laboratory screening of tropical hardwoods for natural resistance to the marine borer Limnoria quadripunctata: the role of leachable and non-leachable factors. Holzforschung, 62(1):99-111. http://www.atypon-link.com/WDG/doi/abs/10.1515/HF.2008.015

Borges LMS; Cragg SM; Busch S, 2009. A laboratory assay for measuring feeding and mortality of the marine wood borer Limnoria under forced feeding conditions: a basis for a standard test method. International Biodeterioration & Biodegradation, 63(3):289-296. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6VG6-4VH8Y1R-1&_user=6686535&_coverDate=04%2F30%2F2009&_rdoc=8&_fmt=high&_orig=browse&_srch=doc-info(%23toc%236030%232009%23999369996%23931093%23FLA%23display%23Volume)&_cdi=6030&_sort=d&_docanchor=&_ct=19&_acct=C000066028&_version=1&_urlVersion=0&_userid=6686535&md5=43c28304035fa0544286a863012f2e28

Borges LMS; Nunes L; Valente AA; Palma P, 2008. Wood boring speceis present in the Tagus Estuary and the severity of their attack on woodne piles exposed in the area: a case study, International Research Group on Wood Protection.

Boyd MJ; Mulligan TJ; Shaughnessy FJ, 2002. Non-indigenous marine species of Humboldt Bay, California. California, USA: California Department of Fish and Game, 118 pp.

Boyle PJ; Mitchell R, 1978a. Absence of microorganisms in crustacean digestive tracts. Science, 200(4346):1157-1159.

Boyle PJ; Mitchell R, 1981b. External microflora of a marine wood-boring isopod. Applied and Environmental Microbiology, 42(4):720-729.

Carlton JT, 1975. Introduced intertidal invertebrates. In: Light's manual: intertidal invertebrates of the central California coast [ed. by Smith, R. I. \Carlton, J. T.]. California, USA: University of California Press, 17-25.

Cookson LJ, 1991. Australasian species of Limnoriidae (Crustacea: Isopoda). Memoirs of the Museum of Victoria, 52(2):137-262.

Cookson LJ, 1997. Additions to the taxonomy of the Limnoriidae (Crustacea: Isopoda). Memoirs of the Museum of Victoria, 56:129-143.

Cookson LJ, 2007. Comparative performance of commercially available timber marine piles in Australia. Forest Products Journal, 57(7/8):27-31.

Cookson LJ; Lorenti M, 2001. A new species of limnoriid seagrass borer (Isopoda) from the Mediterranean. Crustaceana, 74(4):339-346.

Cookson LJ; Poore GCB, 1994. New species of Lynseia and transfer of the genus to Limnoriidae (Crustacea: Isopoda). Memoirs of the Museum of Victoria, 54:197-189.

Cookson LJ; Scown DK; McCarthy KJ; Chew N, 2007. The effectiveness of silica treatments against wood-boring invertebrates. Holzforschung, 61(3):326-332. http://www.extenza-eps.com/WDG/loi/hfsg

Cragg SM; Jumel MC; Al-Horani FA; Hendy IW, 2009. The life history characteristics of the wood-boring bivalve Teredo bartschi are suited to the elevated salinity, oligotrophic circulation in the Gulf of Aqaba, Red Sea. Journal of Experimental Marine Biology and Ecology, 375(1-2):99-105.

Daniel G; Nilsson T; Cragg S, 1991. Limnoria lignorum ingest bacterial and fungal degraded wood. Holz als Roh- und Werkstoff, 49(12):488-490.

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Links to Websites

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Organizations

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Sweden: International Research Group on Wood Protection, IRG Secretariat, Box 5609, SE-114 86, Stockholm, http://www.irg-wp.com/

Contributors

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13/01/10 Original text by:

Simon Cragg, Institute of Marine Sciences, School of Biological Sciences, University of Portsmouth, Ferry Road, Portsmouth PO4 9LY, UK

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