Littorina littorea (common periwinkle)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- 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
- Risk and Impact Factors
- Uses List
- Detection and Inspection
- 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
- Littorina littorea (Linnaeus, 1758)
Preferred Common Name
- common periwinkle
International Common Names
- English: edible periwinkle; winkle; wrinkle
- French: bigorneau; bigorneau commun; bigorneau gris; le vrai bigorneau
Local Common Names
- Germany: Große Strandschnecke
- Ireland: an Giorandán
- Netherlands: gewone alikruik
- Norway: strandsnigel
- Sweden: vanlig strandsnäcka
Summary of InvasivenessTop of page
L. littorea is invasive in North America with a current range from Red Bay, Labrador, to Lewes, Delaware (Blakeslee et al., 2008; Brawley et al., 2009). Recent historical and molecular analyses support Great Britain and Ireland as the region from which L. littorea was introduced into Nova Scotia (NS) (Brawley et al., 2009; also see Blakeslee and Byers, 2008; Blakeslee et al., 2008). L. littorea was common at Pictou (NS) in 1840 (Dawson and Harrington, 1871), and spread southward from Halifax (NS) to Cape May, New Jersey by 1890 (Willis, 1863; Morse, 1880; Verrill, 1880; Ganong 1886, 1887; Bequaert, 1943) and northward to southern Labrador by 1882 (Bequaert, 1943). Rare subfossils of L. littorea exist in the Canadian Maritimes (see History of Invasion/Spread). This snail is the major herbivore of the intertidal zone within its invasive range in the northwestern Atlantic. Although not established, L. littorea is being periodically introduced to bays on the U.S. Pacific coast.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Mollusca
- Class: Gastropoda
- Subclass: Caenogastropoda
- Order: Littorinimorpha
- Unknown: Littorinoidea
- Family: Littorinidae
- Genus: Littorina
- Species: Littorina littorea
Notes on Taxonomy and NomenclatureTop of page
The name Littorina littorea was in common use among scientists by the 1830s-1840s, but the species was originally described by Linnaeus (1758) as Turbo littoreus. No shells or preserved specimens survive as types from Linnaeus’ work, and Reid (1996) established a lectotype from one of Linnaeus’ drawings along with a diagnosis of the species. Reid discusses the taxonomic history involving synonymies and earlier spellings of L. littorea (e.g., Menke (1828) used Litorina litorea). Reid (1996) remarks that a few authors, perhaps even Linneaus, included L. saxatilis within their concept of T. littoreus/L. littorea until the late 1830s, and still fewer authors included L. squalida from the Pacific within L. littorea during part of the 1800s. Lyell (1835) applied the name L. littorea to fossil specimens of L. littorea, and most subsequent authors applied this name to fossil and modern representatives of L. littorea with the same concept of the species as we use today. The common name of L. littorea in English is “the common periwinkle”.
DescriptionTop of page
Reid (1996) gives the mature shell height of L. littorea as 10.6-52.8 mm. Many colour morphs are known, but most shells are dark (brown to black), although when older and eroded, the shell is lighter. The shell is an oblong-turbinate gastropod shell with a large body whorl and pointed spire; sutures and additional whorls are not very prominent except in young snails. Shell width is about two-thirds to three-quartersof the length. The species is unisexual, and, in the breeding season, males have a prominent penis (e.g., Fretter and Graham, 1962; Barroso et al., 2007). Further details and excellent illustrations are given by Fretter and Graham (1962, Chapter 2) and by Reid (1996, see p. 7, 9, 33, 100-105; reproductive details are taxonomically important: males have a penis with 10-42 mamilliform glands, and females have an oviduct that is composed of three loops, two of the albumen gland followed by a capsule gland).
DistributionTop of page
L. littorea occurs abundantly in intertidal and shallow subtidal habitats from the White Sea (Russia) to southern Portugal in the eastern Atlantic; this is its native range (Hawkins et al., 1992; Reid, 1996; Barroso et al., 2007). Rózycki (1991) reported it from southern Spitzbergen. As an invasive species in the northwestern Atlantic, L. littorea occurs from Red Bay, Labrador, to Lewes, Delaware (Reid, 1996; Brawley et al., 2009), where it is abundant on rocky shores, cobbled beaches, salt marshes, and in shallow subtidal areas (Stephenson and Stephenson, 1954; Vadas and Elner, 1992). Occasional recruits on the Virginia (USA) shore are known (Vermeij, 1982). L. littorea is also reported locally from bays (Trinidad Bay and San Francisco Bay, California; Newport Bay, Oregon; Puget Sound, Washington) on the Pacific coast of North America, although it has not become established there as an invasive species (Hanna, 1966; Carlton, 1969; Carlton, 2007; San Francisco Estuary Partnership, 2009; see Invasive Species Management).
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.Last updated: 25 Feb 2021
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Portugal||Present, Widespread||2009||Native||Portugal is the southernmost point of distribution in the northeastern Atlantic|
|Russia||Present||Present based on regional distribution.|
|-Northern Russia||Present, Widespread||2009||Native||Present in White Sea area of northern Russia|
|United Kingdom||Present, Widespread||2009||Native|
|-Channel Islands||Present||1996||Native||Rare on Channel Islands except at Alderney|
|Canada||Present||Present based on regional distribution.|
|-Newfoundland and Labrador||Present||Introduced||Invasive|
|-Prince Edward Island||Present, Widespread||2009||Introduced||Invasive|
|-Quebec||Present, Widespread||2009||Introduced||Invasive||Penetrates St Lawrence River estuary almost to Quebec City|
|United States||Present||Present based on regional distribution.|
|-California||Present, Few occurrences||2009||Introduced||A few individuals found by Carlton in San Francisco Bay. Localised, accidental introductions continue, but an invasive population remains absent|
|-Delaware||Present, Widespread||2009||Introduced||Invasive||Southernmost distribution in NW Atlantic found at Lewes, DL|
|-Maryland||Absent, Formerly present||One living individual found in attached Fucus spiralis|
|-New Hampshire||Present, Widespread||Introduced||Invasive|
|-New Jersey||Present, Widespread||2009||Introduced||1892||Invasive||Reached Cape May in 1892|
|-New York||Present, Widespread||2009||Introduced||Invasive|
|-Oregon||Absent, Formerly present||Occasional introduction but not established|
|-Rhode Island||Present, Widespread||2009||Introduced||Invasive|
|-Virginia||Absent, Formerly present||No longer present|
|-Washington||Absent, Formerly present||Few individuals found in Puget Sound; invasive population not established and no longer present|
History of Introduction and SpreadTop of page
As Wares and Blakeslee (2007) commented, scientists have regarded North American L. littorea as a classic puzzle over time. The puzzle has concerned which of these hypotheses is true: 1) L. littorea in North America is derived completely from invasive introduction(s) of snails from Europe (invasion hypothesis), 2) L. littorea is native to North America, but was rare until the 1800s (native species hypothesis), or 3) L. littorea is both introduced and native to North America. The invasive species hypothesis is supported strongly by recent analyses (Blakeslee and Byers, 2008; Blakeslee et al., 2008; Brawley et al., 2009), but there are still some unresolved issues (see below) that deserve additional research.
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Canada||Ireland||early 1800s||Hitchhiker (pathway cause)||Brawley et al. (2009)||See text for discussion of subfossils in North America that predate this period; these subfossils have led to propose that there was also a native population|
|Canada||UK||early 1800s||Hitchhiker (pathway cause)||Brawley et al. (2009)||See text for discussion of subfossils in North America that predate this period; these subfossils have led to propose that there was also a native population|
|Pacific, Northeast||1900s||Hitchhiker (pathway cause)||Carlton (2007); San Francisco Estuary Partnership (2009)||Spot introductions that appear too small to form invasive populations; recent introductions in San Francisco Bay are part of an active eradication effort|
|USA||Canada||late 1800s||Self-propelled (pathway cause)||Reid (1996)||Planktonic dispersal led to rapid and well-chronicled spread from Nova Scotia along the Atlantic shore of New Brunswick to Maine and southward to New Jersey by end of the nineteenth century|
Risk of IntroductionTop of page
L. littorea can survive a wide range of air and water temperatures, is tolerant of brackish water, can withstand brief anoxic periods, and feeds under both damp emersed and immersed conditions in the intertidal zone (see Physiology). Thus, its physiology makes it unusually capable of being transported and established outside its native range, whether in rock ballast on ships that took ~ 21-60 days to reach Pictou in the nineteenth century (Brawley et al., 2009) or, presently, as a hitchhiker as other marine species are transported for aquaculture, food, or fishing bait from the Atlantic to the Pacific.
HabitatTop of page
L. littorea reaches intertidal abundances of hundreds to thousands of animals/m2 on rocky shores, cobbled beaches, mudflats and Spartina marshes (e.g., Lubchenco, 1978; Bertness, 1984; Buschbaum, 2000; Carlson et al., 2006; Tyrell et al., 2008). Generally, it is most common in the lower half of the intertidal zone. L. littorea is found both on rock surfaces and macrophytes (e.g., Fucus vesiculosus, F. serratus, Saccharina latissima, Ascophyllum nodosum,S Brawley, University of Maine, USA, personal communication, 2009) in both the northeastern and northwestern Atlantic. The species is found at considerable depth (e.g., Huntsman, 1918 (36 m, Gulf of St Lawrence); Fretter and Graham, 1980 (60 m, northern Britain)), but predation limits its abundance subtidally (Pettitt, 1975; Perez et al., 2009). L. littorea typically moves into subtidal areas on cold shores (e.g., Maine) in winter, becoming uncommon in the intertidal zone until spring.
Habitat ListTop of page
|Littoral||Coastal areas||Principal habitat||Harmful (pest or invasive)|
|Littoral||Coastal areas||Principal habitat||Natural|
|Littoral||Mud flats||Principal habitat||Harmful (pest or invasive)|
|Littoral||Mud flats||Principal habitat||Natural|
|Littoral||Intertidal zone||Principal habitat||Harmful (pest or invasive)|
|Littoral||Intertidal zone||Principal habitat||Natural|
|Littoral||Salt marshes||Principal habitat||Harmful (pest or invasive)|
|Littoral||Salt marshes||Principal habitat||Natural|
|Brackish||Estuaries||Principal habitat||Harmful (pest or invasive)|
|Marine||Inshore marine||Principal habitat||Harmful (pest or invasive)|
|Marine||Inshore marine||Principal habitat||Natural|
Biology and EcologyTop of page
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)||Optimum||Tolerant of short-term anoxic conditions|
|Salinity (part per thousand)||30||33||Optimum||10-40 tolerated, both values in psu. Euryaline|
|Water temperature (ºC temperature)||8||22||Optimum||0-28 tolerated. Acclimation (phenotypic) and adaptation (genotypic) cause the temperature range to be different across the geographic range, but L. littorea is eurythermal at all sites|
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Calidris canutus||Predator||Aquatic|Adult||not specific|
|Cancer borealis||Predator||Aquatic|Adult||not specific|
|Cancer irroratus||Predator||Aquatic|Adult||not specific|
|Carcinus maenas||Predator||Aquatic|Adult||not specific|
|Cliona||Parasite||All Stages||not specific|
|Cryptocotyle lingua||Parasite||Aquatic|Adult||not specific|
|Larus argentatus||Predator||Aquatic|Adult||not specific|
|Platichthys flesus||Predator||Aquatic|Adult||not specific|
|Polydora ciliata||Parasite||Aquatic|Adult||not specific|
Notes on Natural EnemiesTop of page
The heavy shell of L. littorea protects it from many predators but numerous birds, fishes, and crabs are major predators (Pettitt, 1975). Birds that eat L. littorea include herring gulls (Larus argentatus), the knot (Calidris canutus), and oldsquaws (Clangula hymenalis); among the many fishes that eat L. littorea, flounders (e.g., Pleuronectes flesus) are prominent (Pettitt, 1975). Crabs crush shells, and it is common to see L. littorea on the shore that have repaired their shells after escaping crab predation (e.g., Vermeij, 1982). In Europe, the green crab Carcinus maenas is a major predator on L. littorea (e.g., Buschbaum et al., 2007), and this is also true of the crab’s invasive range in North America (Carlson et al., 2006; Ellis et al., 2007; Edgell and Rochette, 2008; Perez et al., 2009). ). L. obtusata is much more affected by C. maenas predation than L. littorea in the Gulf of Maine and Bay of Fundy, which Edgell and Rochette (2008) suggested was due to the prior evolution of predator-resistance by L. littorea in Europe, before its introduction to North America. The Jonah crab Cancer borealis is an important L. littorea predator and is especially important in limiting subtidal abundance of L. littorea (Perez et al., 2009). Lobsters (e.g., Homarus americanus) eat few L. littorea (Jones and Shulman, 2008). The Asian shore crab (Hemigrapsus sanguineus, now invasive on the U.S. shore) consumes few L. littorea directly, but it may eat juveniles as it consumes algae based upon the decreased abundance of L. littorea on one studied shore as H. sanguineus became more abundant (Bourdeau and O’Connor, 2003; Kraemer et al., 2007). Humans should also be considered a natural enemy based upon their use of periwinkles for food (Pettitt, 1975; Reid, 1996).
Means of Movement and DispersalTop of page
Pathway CausesTop of page
|Aquaculture||Accidently introduced with oysters||Yes||Carlton (2007)|
|Fisheries||Accidently introduced with oysters||Yes||Carlton (2007)|
|Food||Hypothesis||Yes||Yes||Department of Marine Resources State of Maine (2009a); Department of Marine Resources State of Maine (2009b); Marine Institute Ireland (2009); Marine Sacs Project UK (2009); Sharp (1998); Sharp et al. (1998)|
|Hitchhiker||With Atlantic seaweeds used as packing for long distance transport of bait worms and shellfish/fish||Yes||Carlton (2007)|
|Hunting, angling, sport or racing||With shipments of bait from Atlantic to Pacific for anglers||Yes||Carlton (2007)|
|Intentional release||Hypothesis||Yes||San Francisco Estuary Partnership (2009); Steneck and Carlton (2001)|
|Interconnected waterways||Yes||Reid (1996)|
|Self-propelled||Planktonic larvae are a highly dispersive stage in life history||Yes||Yes||FRETTER and GRAHAM (1962); Reid (1996)|
|Timber trade||associated with rock ballast||Yes||Brawley et al. (2009)|
Pathway VectorsTop of page
|Aquaculture stock||Adults, in past with oysters||Yes||Carlton (2007)|
|Bait||Adult hitchikers in rockweed (Ascophyllum nodosum) used for packing of bait worms or shellfish||Yes||Carlton (2007); San Francisco Estuary Partnership (2009)|
|Containers and packaging - wood||Hitchikers in rockweed (Ascophyllum nodosum) used for packing of bait worms or shellfish||Yes||Carlton (2007); San Francisco Estuary Partnership (2009)|
|Live seafood||Yes||Yes||Carlton (2007)|
|Ship ballast water and sediment||With rock ballast||Yes||Brawley et al. (2009)|
Impact SummaryTop of page
|Economic/livelihood||Positive and negative|
Economic ImpactTop of page
This snail is a concern to fish aquaculture because, when abundant, it can cause more fish to be infected with trematode parasites (e.g., Kristoffersen, 1991). L. littorea should be considered as a potential threat to the establishment of sea vegetable aquaculture in Europe and North America because it consumes Porphyra spp. and Palmaria spp.
Environmental ImpactTop of page
Impact on Habitats
Impacts on Biodiversity
Grazing by L. littorea quantitatively reduces recruitment of many benthic intertidal organisms; larger sessile organisms (e.g., rockweeds) size-escape grazing and then benefit from being cleaned by surficial grazing of L. littorea on their surfaces (Lubchenco, 1983; Vadas and Elner, 1992). Locally, the quantity of green algae (in particular) is markedly reduced by the presence of L. littorea, but the overall biodiversity (species richness) of a shore is rarely affected because there are microhabitat refuges where such species escape grazing.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Has a broad native range
- Abundant in its native range
- Highly adaptable to different environments
- Is a habitat generalist
- Capable of securing and ingesting a wide range of food
- Highly mobile locally
- Has high reproductive potential
- Has high genetic variability
- Altered trophic level
- Ecosystem change/ habitat alteration
- Modification of natural benthic communities
- Modification of successional patterns
- Negatively impacts animal health
- Soil accretion
- Threat to/ loss of native species
- Competition - monopolizing resources
- Pest and disease transmission
- Interaction with other invasive species
- Highly likely to be transported internationally accidentally
- Difficult/costly to control
UsesTop of page
Uses ListTop of page
Animal feed, fodder, forage
- Attractant in fish/shrimp feed
Human food and beverage
- Emergency (famine) food
- Meat/fat/offal/blood/bone (whole, cut, fresh, frozen, canned, cured, processed or smoked)
Detection and InspectionTop of page
L. littorea is easy to spot on shipped live seafoods or potential aquaculture targets, and to prevent spread to the North American Pacific coast, this should be carried out by traders in seafoods, shellfish, and bait worms. However, there are no regulations in the State of Maine that enforce such inspections presently (Department of Marine Resources, State of Maine, personal communication to S Brawley, University of Maine, USA, December 2009), and the best strategy to change behaviour (e.g., type of packing materials) is probably for regulation to occur at the level of receiving states and countries.
Similarities to Other Species/ConditionsTop of page
In its invasive range in the Atlantic, L. littorea is most similar to L. saxatilis, but is easily distinguished when mature by its larger size and the less pronounced ridges (roughness) of the L. littorea shell compared to L. saxatilis. Also, L. saxatilis is rarely found below the high intertidal zone, and L. littorea becomes more common below the high zone (but is also found in abundance in many high intertidal zones and pools). In the Pacific, L. squalida, which is considered to be L. littorea’s sister species (Reid, 1996), has a similar shell shape, but L. littorea’s shell is a little narrower and less incised. These two species are also distinguished by differences in reproductive anatomy, and the number of zygotes/capsule that are spawned (14-15 zygotes/capsule in L. squalida; 1-3 [less commonly to 9] zygotes/capsule in L. littorea; see Reid, 1996, Table 3, p. 91). When in doubt (e.g., with immature specimens), molecular techniques will separate L. littorina from other species (Reid et al., 1996).
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Once a large breeding population is established, eradication is not feasible, and the snail can be expected to extend its range substantially. Thus, control of this potentially invasive snail in the Pacific is best based upon care in avoiding accidental transport with other live organisms from the intertidal zone. The San Franscisco Bay Estuary Partnership (2009) is sponsoring local eradication efforts for L. littorea, which appear to be helpful to preventing invasion.
Gaps in Knowledge/Research NeedsTop of page
1. More information is needed on the paleooceanography of the North Atlantic, especially in the last interglacial period.
ReferencesTop of page
Barroso CM; Gonçalves C; Moreira MH, 2007. Growth, reproductive cycle and penis shedding of Littorina littorea in the Ria de Aveiro (north-west Portugal). Journal of the Marine Biological Association of the United Kingdom, 87:547-550.
Blakeslee AMH; Byers JE; Lesser MP, 2008. Solving cryptogenic histories using host and parasite molecular genetics: the resolution of Littorina littorea's North American origin. Molecular Ecology, 17(16):3684-3696. http://www3.interscience.wiley.com/cgi-bin/fulltext/120779938/HTMLSTART
Brawley SH; Coyer JA; Blakeslee AMH; Hoarau G; Johnson LE; Byers JE; Stam WT; Olsen JL, 2009. Historical invasions of the intertidal zone of Atlantic North America associated with distinctive patterns of trade and emigration. Proceedings of the National Academy of Sciences of the United States of America, 106(20):8239-8244. http://www.pnas.org/
Byers JE; Blakeslee AMH; Linder E; Cooper AB; Maguire TJ, 2008. Controls of spatial variation in the prevalence of trematode parasites infecting a marine snail. Ecology, 89(2):439-451. http://www.esajournals.org/perlserv/?request=get-abstract&doi=10.1890%2F06-1036.1
Chapman JW; Carlton JT; Bellinger MR; Blakeslee AMH, 2007. Premature refutation of a human-mediated marine species introduction: the case history of the marine snail Littorina littorea in the northwestern Atlantic. Biological Invasions, 9(6):737-750. http://www.springerlink.com/content/7jk8k72622456598/?p=90d9db51ce5547e18ef85ee971c0decf&pi=10
Chase ME; Thomas MLH, 1995. The effect of the rate and onset of temperature increase on spawning of the periwinkle, Littorina littorea (L.). Journal of Experimental Marine Biology and Ecology, 186:277-297.
Cunningham CW, 2008. How to use genetic data to distinguish between natural and human-mediated introduction of Littorina littorea to North America. Biological Invasions, 10(1):1-6. http://www.springerlink.com/link.asp?id=103794
Davenport J; Davenport JL, 2007. Interaction of thermal tolerance and oxygen availability in the eurythermal gastropods Littorina littorea and Nucella lapillus. Marine Ecology Progress Series, 332:167-170.
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Jones PL; Shulman MJ, 2008. Subtidal-intertidal trophic links: American lobsters [Homarus americanus (Milne-Edwards)] forage in the intertidal zone on nocturnal high tides. Journal of Experimental Marine Biology and Ecology, 361:98-103.
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Perez KO; Carlson RL; Shulman MJ; Ellis JC, 2009. Why are intertidal snails rare in the subtidal? Predation, growth and the vertical distribution of Littorina littorea (L.) in the Gulf of Maine. Journal of Experimental Marine Biology and Ecology, 369:79-86.
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Tyrrell MC; Dionne M; Edgerly JA, 2008. Physical factors mediate effects of grazing by a non-indigenous snail species on saltmarsh cordgrass (Spartina alterniflora) in New England marshes. ICES Journal of Marine Science [5th International Conference on Marine Bioinvasions: Integrating Knowledge for Managing Impacts, Cambridge, Massachusetts, USA, 21-24 May 2007.], 65(5):746-752. http://icesjms.oxfordjournals.org/cgi/content/abstract/65/5/746
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Wood CL; Byers JE; Cottingham KL; Altman I; Donahue MJ; Blakeslee AMH, 2007. Parasites alter community structure. Proceedings of the National Academy of Sciences of the United States of America, 104(22):9335-9339. http://www.pnas.org/
Barroso C M, Gonçalves C, Moreira M H, 2007. Growth, reproductive cycle and penis shedding of Littorina littorea in the Ria de Aveiro (north-west Portugal). Journal of the Marine Biological Association of the United Kingdom. 547-550.
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CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Sertkaya G, Martini M, Musetti R, Osler R, 2007. Detection and molecular characterization of phytoplasmas infecting sesame and solanaceous crops in Turkey. Bulletin of Insectology. 60 (2), 141-142. http://www.bulletinofinsectology.org/
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15/12/09 Original text by:
Susan Brawley, University of Maine, USA
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