the freshwater anchovy ( thryssa scratchleyi ) is a species of fish in the engraulidae family .

it is endemic to australia .

often mistaken for a minnow .

freshwater anchovy

only anchovy in australia restricted to mainly freshwater habitats . form shoals in deep water of lagoons .
freshwater anchovy , thryssa scratchleyi . source : whitehead , nelson & wongratana , 1988 , fao . license : cc by attribution
] . for these analyses , marine and freshwater partitions were considered as separate datasets . as with the clade model analyses , we found clear evidence for positive selection occurring in the freshwater anchovy dataset (
setipinna taty or ikan bilis is the anchovy commonly used in south - east asian cooking to make fish stock or sambals . anchovy is also used to produce budu , by fermentation process . in vietnam , anchovy is the main ingredient to make fish sauce—nước mắm—the unofficial national sauce of vietnam .
the european anchovy , engraulis encrasicolus , is the anchovy of commerce . morocco now leads the world in canned anchovies . the anchovy industry along the coast of cantabria now dwarfs the traditional catalan salters , though the industry was only initiated in cantabria by sicilian salters in the mid - nineteenth century .
the anchovy also is extremely important to marine mammals and birds . for example , there is a strong correlation between anchovy abundance and the breeding success of california brown pelicans and elegant terns ( psmfc 1996 ) .
there is no special fishery for this species , however , it is mainly used as baitfish and to a small extent for anchovy paste ( munroe and nizinski 2002 ) . anchovy are also a target species for semi - industrial pelagic fisheries of argentina with total landings of anchovy in 2003 at 900 tonnes ( elías et al . 2011 ) .
pacific states marine fisheries commission ( psmfc ) . northern anchovy . psmfc , 1996 . retrieved october 3 , 2007 .
harris , j . h . 1984 . zoogeography of the australian freshwater fish fauna . pp . 211–222 .
occurs in brackish estuaries and adjacent marine areas ; penetrates freshwater ( ref . 5217 ) . forms moderate schools . usually migrating from estuaries or the sea , but possibly the limnichthys of lake maracaibo and the amazonensis represent purely freshwater forms . in freshwater , it feeds principally on small fishes , prawns and copepods ; also insect larvae . at sea , it feeds on fishes and various crustaceans . spawns in freshwater . year round occurrence of eggs reported .
allen , g . r . , h . midgley , and m . allen 2002 field guide to the freshwater fishes of australia . field guide to the freshwater fishes of australia . : i - xiv + 1 - 394 .
parker rs , hackney ct , and mf vidrine . 1984 . ecology and reproductive strategy of a south louisiana freshwater mussel , glebula rotundata ( lamarck ) ( unionidae : lampsilini ) . freshwater invertebrate biology 3 : 53 - 58 .
most species of anchovy feed on plankton , including some by filter - feeding , but some larger species each fish ( nelson 1994 ) .
: r . m . mcdowall ( ed . ) freshwater fishes of south - eastern australia , reed , sydney .
the bay anchovy , anchoa mitchilli . illustration by diana rome peebles 1998 . courtesy of florida fish and wildlife conservation commission , division of marine fisheries .
: t . zaret ( ed . ) evolutionary ecology of neotropical freshwater fishes , dr w . junk publishers , the hague .
koehn , j . 1986 . approaches to determining flow and habitat requirements for native freshwater fish in victoria . pp . 95–115 .
overfishing of anchovies has been a problem . since the 1980s , large mechanized anchovy fishing vessels based in france have caught the fish in fine - mesh dragnets .
luo j and ja musick . 1991 . reproductive biology of the bay anchovy in chesapeake bay . transactions of the american fisheries society 120 : 701 - 710 .
anchovies are found in scattered areas throughout the world ' s oceans , as well as some freshwater and brackish water habitats , but are concentrated in temperate waters , and are rare or absent in very cold or very warm seas . of the 139 species in 16 genera recognized by nelson ( 1994 ) , only 17 are freshwater species , occasionally entering brackish water , while the others are marine , rarely entering freshwater .
lake , j . s . 1971 . freshwater fishes and rivers of australia . thomas nelson & sons ltd , melbourne . 61 pp .
allen , g . r . 1989 . freshwater fishes of australia . t . f . h . publications , neptune city . 240 pp .
fossil calibration points ( c1–c7 ) , key divergence times and inferred freshwater colonization events are indicated on the phylogeny along with 95 % reliability intervals .
llewellyn , l . c . & m . c . macdonald . 1980 . family percichthyidae . australian freshwater basses and cods . pp . 142–149 .
llewellyn , l . c . 1971 . breeding studies on the freshwater forage fish of the murray - darling river system . the fisherman , june : 1–12 .
merrick , j . r . & g . e . schmida . 1984 . australian freshwater fishes : biology and management . griffin , adelaide . 409 pp .
dalton pd . 1987 . ecology of bay anchovy , anchoa mitchilli , eggs and larvae in the mid - chesapeake bay . unpublished masters thesis , university of maryland , college park , maryland .
bay anchovies are similar in appearance to two other common anchoa species with which it co - occurs throughout much of florida . the striped anchovy ( anchoa hepsetus ) often grows to be somewhat larger ( to 15 cm ) than the bay anchovy , and both it and the cuban anchovy ( a . cubana ) have a dorsal fin that begins well in front of the anal fin . bay anchovies are the only us anchovies in which the dorsal fin begins at a point right above or only very slightly in front of where the anal fin begins ( robbins et al . 1986 ) .
ancient lakes are home to disproportionate levels of freshwater biodiversity . as standing bodies of water which have existed for at least 100 , 000 years [ 1 ] , these habitats have been remarkably stable when compared to more typically transitory freshwater environments . as a consequence , lakes such as lake baikal ( 25–30 my ) and the african great lakes malawi ( 1–2 my ) and tanganyika ( 9–12 my ) all contain exceptionally high numbers of freshwater taxa , of which up to 99 % are endemics [ 2 ] .
fives jm , warlen sm , and de hoss . 1986 . aging and growth of larval bay anchovy , anchoa mitchilli , from the newport river estuary , north carolina . estuaries 9 : 362 - 367 .
most species of anchovy are small . most are under 20 centimeters in length , but thryssa scratchleyi , which inhabits rivers in papau new guinea and the gulf of carpentaria , reaches 37 centimeters ( nelson 1994 ) .
cadwallader , p . l . & g . n . backhouse . 1983 . a guide to the freshwater fish of victoria . victorian government printing office , melbourne . 249 pp .
olney je . 1983 . eggs and early larvae of the bay anchovy , anchoa mitchilli , and the weakfish , cynoscion regalis , in lower chesapeake bay with notes on associated ichthyoplankton . estuaries 6 : 20 - 35 .
rilling gc and ed houde . 1999 . regional and temporal variability in distribution and abundance of bay anchovy ( anchoa mitchilli ) eggs , larvae , and adult biomass in the chesapeake bay . estuaries 22 : 1096 - 1109 .
harris , j . h . & p . c . gehrke . 1994 . modelling the relationship between streamflow and population recruitment to manage freshwater fisheries . aust . fish . 6 : 28–30 .
koehn , j . d . & w . g . o ' connor . 1990 . biological information for management of native freshwater fish in victoria . government printer , melbourne . 165 pp .
of particular interest in light of the marine incursion scenario put forth here is the grouping of ethmalosa fimbriata , an estuarine species widespread along the coasts of west africa , with the freshwater pellonuline herring found in the region [ 43 ] . this species has been the focus of a recent phylogeographic study [ 45 ] , which suggests that the historical population structure of the species has been strongly influenced by pleistocene sea level fluctuations in the region , when local populations of the species were isolated in freshwater refuges . as this euryhaline species may be the closest living marine relative of the freshwater pellonulines of west africa , future comparisons between the morphology and physiology of ethmalosa and its pellonuline relatives may help to identify key innovations that allowed the ancestor of these groups to successfully colonize freshwater .
habitat includes lower freshwater and estuarine reaches of coastal rivers , bays , sounds , and high salinity nearshore marine waters ; usually it occurs in shallow waters , at depths of less than 20 meters .
positive selection has occurred in the rhodopsin gene of freshwater anchovies , but not their marine relatives . this result is consistent with our hypothesis that the invasion of the much dimmer and red - shifted freshwater rivers of south america is accompanied by visual adaptation in the dim - light sensitive visual pigment rhodopsin . the relatively low d n / d s observed in marine anchovies may be owing to the negligible attenuation of light at depths inhabited by the majority of anchovy lineages [ 15 ] , or because the peak absorbance of rhodopsin is already tuned to the ideal wavelength of light for marine environments . the lack of evidence for positive selection in the three non - vision - related genes indicates that the increased d n / d s found for rhodopsin in freshwater is not owing to differences in population structure or genome - wide shifts in evolutionary rates .
larson , h . k . and k . c . martin 1989 freshwater fishes of the northern territory . northern territory museum of arts and sciences . handbook series number 1 . darwin . 102 p .
lake , j . s . 1967a . rearing experiments with five species of australian freshwater fishes . i . inducement to spawning . aust . j . mar . freshw . res . 18 : 137–153 .
lake , j . s . 1967b . rearing experiments with five species of australian freshwater fishes . ii . morphogenesis and ontogeny . aust . j . mar . freshw . res . 18 : 155–173 .
in this study , we use the striking marine to freshwater habitat transition as a unique natural experiment to study the effects of different light environments on rhodopsin gene evolution . at the shallow depths occupied by the majority of marine anchovies , the spectral attenuation of light is negligible [ 3 ] . by contrast , at similar depths in south american rivers , the amount of available light is substantially decreased and richer in longer wavelengths . in addition , the degree of spectral attenuation can vary among south american rivers , broadly classified as white water , black water or clear water based on their optical qualities [ 9 ] . given the contrast in spectra of downwelling light between marine and freshwater habitats , as well as the diversity of visual environments in south american rivers , we predicted that the freshwater - invading anchovy lineage would show evidence of positive darwinian selection in the rhodopsin gene . to test this hypothesis , we sequenced rhodopsin from new world anchovies and used codon - based models of molecular evolution to compare the strength of selection acting on freshwater invaders versus their marine relatives .
jordan rc , gospodarek am , schultz et , cowen rk , and k lwiza . 2000 . spatial and temporal growth rate variation of bay anchovy ( anchoa mitchilli ) larvae in the mid hudson river estuary . estuaries 23 : 683 - 689 .
castellanos dl and lp rozas . 2001 . nekton use of submerged aquatic vegetation , marsh , and shallow unvegetated bottom in the atchafalaya river delta , a louisiana tidal freshwater ecosystem . estuaries 24 : 184 - 197 .
south american rivers are also more spectrally diverse than marine ecosystems [ 9 ] . lineages distributed across highly stratified or disparate freshwater light environments may have increased d n / d s values owing to divergent selection pressures acting on the visual systems of species occupying spectrally distinct habitats [ 16 ] . in freshwater anchovies , different adaptations may be required for optimal visual performance in the highly turbid amazon , the clear waters of the rio tapajos or the tannin stained black waters of the rio negro . also , the ecological diversification of freshwater anchovies ( for example , into actively hunting piscivores ) has probably altered visual demands and correspondingly shifted patterns of natural selection in rhodopsin [ 7 ] .
marine ; freshwater ; brackish ; pelagic - neritic ; amphidromous ( ref . 51243 ) ; depth range 0 - 50 m ( ref . 189 ) . tropical ; 24°n - 9°s , 70°e - 118°e ( ref . 189 )
citation : wilson ab , teugels gg , meyer a ( 2008 ) marine incursion : the freshwater herring of lake tanganyika are the product of a marine invasion into west africa . plos one 3 ( 4 ) : e1979 . urltoken
zastrow ce , houde ed , and lg morin . 1991 . spawning , fecundity , hatch - date frequency and young - of - the - year growth of bay anchovy , anchoa mitchilli in mid - chesapeake bay . marine ecology progress series 73 : 161 - 171 .
) , nor for positive selection in either the freshwater or marine partitions for any of the non - vision - related genes ( electronic supplementary material , table s2 ) . to ensure that these results were not owing to artefacts in d
von rintelen t , wilson ab , meyer a , glaubrecht m ( 2004 ) escalation and trophic specialization drive adaptive radiation of freshwater gastropods in ancient lakes on sulawesi , indonesia . proc r soc lond b biol sci 271 : 2541–2549 .
castillo - rivera m , moreno g , and r iniestra . 1994 . spatial , seasonal , and diel variation in abundance of the bay anchovy , anchoa mitchilli ( teleostei : engraulidae ) , in a tropical coastal lagoon of mexico . the southwestern naturalist 39 : 263 - 268 .
albrecht c , trajanovski s , kuhn k , streit b , wilke t ( 2006 ) rapid evolution of an ancient lake species flock : freshwater limpets ( gastropoda : ancylidae ) in the balkan lake ohrid . org divers evol 6 : 294–307 .
select a firm frozen anchovy . soft thawed anchovies can be used most effectively as chum . cut the anchovy into 3 pieces any of which can be used . the head stays on the hook best with the tail being fairly resilient . the trick to making the frozen bait stay on is to only run the hook in one time . do not attempt to turn or reposition the barb . push it in once and leave it for best results . yes the bait will fall off when it thaws but i want to have a fish by then anyway . it should work long enough !
clade model c ( paml ) analyses of rhodopsin and non - vision - related genes . ( lnl , ln likelihood ; ω , d n / d s for site class and percentage of sites in site class ; m / f , marine partition ( background ) / freshwater partition including marine reinvading species ( foreground ) ; asterisk denotes significantly different marine versus freshwater d n / d s values ( * p < 0 . 01 ) ; d . f . , degrees of freedom . )
anchovies generally are very accepting of a wide range of temperatures and salinity . large schools can be found in shallow , brackish areas with muddy bottoms , as in estuaries and bays . most freshwater species are found in south america ( nelson 1994 ) .
the rhodopsin gene was amplified and sequenced from genomic dna extracted from 35 species spanning the taxonomic diversity of freshwater and marine new world anchovies . we used codon - based models of molecular evolution to estimate the ratio of non - synonymous to synonymous substitutions (
roper river drainage , nt , archer river to leichhardt river drainages , gulf of carpentaria , cape york , from mitchell river , qld ( 15°10’s ) ; tropical ; pelagic ; inhabiting freshwater streams and associated lagoons , in clear or turbid water and also in brackish estuaries .
anchovies are an important part of marine food chains , as well as some freshwater food chains . it is a good food source for almost every predatory fish in its environment , including the california halibut , rock fish , yellowtail , sharks , chinook , and coho salmon ( psmfc 1996 ) .
numbers on branches represent bootstrap support for distance , maximum parsimony and maximum likelihood analyses and posterior probabilities from bayesian analysis . traditional clupeoid groups and major african freshwater lineages indicated . clupeid diagrams reprinted with the permission of the food and agriculture organization of the united nations [ 34 ] , [ 75 ] .
phylogeny and molecular evolution in new world anchovies . ( a ) phylogenetic relationships among marine ( blue ) and freshwater ( red ) species of the subfamily engraulinae [ ] . colours on branches show optimization of habitat type ; partitions used for clade model analyses indicated by vertical bars . ( b ) d n / d s estimates for the divergent site class in marine and freshwater partitions from clade model analyses implemented in paml . ( c ) sites with a greater than 0 . 75 posterior probability for inclusion in the positively selected site class as inferred by fubar . fish images after [ ] . ( online version in colour . )
anchovies are most commonly marketed in small tins , either as
flat
fillets , or as
rolled anchovies
where each fillet is rolled around a caper . while both are quite salty , the flat fillets tend to be more so . they are also marketed in jars and tubes as a paste , mostly for use in making sauces , such as anchovy essence .
phylogeny and molecular evolution in new world anchovies . ( a ) phylogenetic relationships among marine ( blue ) and freshwater ( red ) species of the subfamily engraulinae [ 5 ] . colours on branches show optimization of habitat type ; partitions used for clade model analyses indicated by vertical bars . ( b ) d n / d s estimates for the divergent site class in marine and freshwater partitions from clade model analyses implemented in paml . ( c ) sites with a greater than 0 . 75 posterior probability for inclusion in the positively selected site class as inferred by fubar . fish images after [ 13 ] . ( online version in colour . )
anchovy is any member of the more than 100 species of small , largely marine , herring - like fish comprising the family engraulidae . most are characterized by a large mouth , with the upper jaw extending well behind the eye and the jaw articulation well behind the eye , as well as a long , pig - like snout extending beyond the tip of the lower jaw ( nelson 1994 ) .
a strong association between bay anchovies and the atlantic brief squid ( lolliguncula brevis ) in a study by ogburn - matthews and allen ( 1993 ) is likely reflective of a strong predator - prey relationship between these species . gelatinous predators such as sea nettles ( chrysaora quinquecirrha ) end ctenophores ( e . g . , mnemiopsis leidyi ) are known to consume bay anchovy eggs ( breitburg et al . 1997 , rilling and houde 1999 ) .
safina and burger ( 1989 ) indicate bay anchovies are one of two prey fish species most preyed upon by predatory fish and terns near fire island inlet , ny . bluefish ( pomatomus saltatrix ) were identified as a major consumer of bay anchovies by these authors , and appear capable of altering anchovy population numbers through predation . mcginnis and emslie ( 2001 ) report north carolina royal terns ( sterna maxima ) and sandwich terns ( s . sandvicensis ) both prey on bay anchovies .
random sites ( paml ) analyses of rhodopsin . ( lnl , ln likelihood ; p and q , parameters of beta distribution of site classes in models m8a and m8 ; ω , d n / d s for site class and percentage of sites in site class ; asterisk denotes significantly different marine versus freshwater d n / d s values ( * p < 0 . 01 ) ; d . f . , degrees of freedom . )
field surveys by rilling and houde ( 1999 ) revealed that bay anchovy spawning in chesapeake bay occurred from may through september and peaked during july in the seaward third of the bay . the authors estimate that baywide daily egg production increased from 4 . 25 x 10 12 in june to 8 . 43 x 10 12 in july . olney ( 1983 ) reports that 99 % of fish egg catches and 67 - 88 % of larval catches during this period are bay anchovies . during peak spawning , pelagic egg densities range from 10 - 1 , 000 / m 3 and larval densities reach 1 - 100 / m 3 ( olney 1983 , dalton 1987 ) .
anchovies provide important ecological , commercial , and culinary values . some species of anchovies , especially the european and mediterranean fish engraulis encrasicholus and other engraulis species , are commercially important , being used in various appetizers , sauces , and dishes . as a group , anchovies are an important part of marine food chains , as well as some freshwater food chains , serving as a good food source for predatory fish , as well as marine mammals and birds . some there is some danger of commercial interests leading to their being overfished and thus disrupting the delicate balance in marine ecosystems .
while the frequency of marine incursions into the congo basin is thought to have slowed after the mesozoic due to changes in the geology of the region [ 29 ] , beadle [ 30 ] indicates that the congo basin was dominated by a large inland sea during the pliocene ( 2–5 mya ) . if this is indeed the case , the presence of a large stable water body in central africa at this time may have facilitated the dispersal of freshwater organisms between western and eastern africa . the presence of this palaeolake may have also fostered increased rates of speciation during the pliocene , a pattern recently suggested for the cichlid fishes of palaeolake makgadikgadi in southern africa [ 31 ] .
with marine fisheries’ yield plateauing over the past two decades , and global demand for fish and products continuing to increase , the marine aquaculture sector has grown to meet demand ( frid and paramor 2012 ) . recent estimates suggest that almost 50 % of all fish biomass consumed by humans is produced by aquaculture ( fao 2014 ) . among other environmental impacts , including the dependence on capture fisheries ( primarily of forage fish such as menhaden and anchovy ) to provide fishmeal , aquaculture facilities can negatively impact other fish species . specifically , survival of wild populations can be reduced through the movement of diseases and parasites from farmed to wild individuals , competition with escaped farmed individuals , and hybridisation ( e . g . , salmonids : ford and myers 2008 ) . marine aquaculture was identified as affecting seven species , none of which are currently assessed as threatened .
d 12 - 13 ; a 38 - 42 ; p 13 . body compressed ; abdomen with 27 - 31 scutes ; small spine - like scute present just anterior to dorsal fin origin ; large overshot mouth and very long upper jaw ; maxilla short only reaching to front border of opercle ; teeth small or minute , not canine - like ; lower gill rakers 18 to 23 ; scales large , usually 43 or 44 in lateral series ; vertical striae on scales just interrupted at middle of scale . dorsal fin rays branched ; dorsal fin origin about mid body ; anal fin relatively long , origin at or behind vertical through base of posteriormost dorsal fin ray .
silvery overall , steely - blue dorsally ; no dark blotches on nape ; a dusky area posterior to upper part of gill opening .
breeding biology is poorly known , it is thought that adults move downstream to breed in estuaries .
engraulis scrathleyi ramsay e . p . & ogilby j . d . ( 1886 ) a contribution to the knowledge of the fish fauna of new guinea . proc . linn . soc . n . s . w . ( 2 ) 1 ( 1 ) : 8 - 20 strickland river , new guinea .
carpenter , k . e . and v . h . niem 1999 species identification guide for fisheries purposes . the living marine resources of the western central pacific . batoid fishes , chimeras and bony fishes part 1 ( elopidae to linophrynidae ) . fao , rome .
paxton , j . r . , d . f . hoese , g . r . allen and j . e . hanley 1989 zoological catalogue of australia . volume 7 . pisces . petromyzontidae to carangidae . australian government publishing service , canberra . zoological catalogue of australia . volume 7 . v . 7 : i - xii + 1 - 665 .
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yearsley , g . k . , last , p . r . & morris , g . b . 1997 ,
codes for australian aquatic biota ( caab ) : an upgraded and expanded species coding system for australian fisheries databases
, pp . 15 pp . + appendices
ramsay , e . p . & ogilby , j . d . 1886 ,
a contribution to the knowledge of the fish fauna of new guinea
, proceedings of the linnean society of new south wales , ser . 2 , vol . 1 , no . 1 , pp . 8 - 20
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some species of anchovies , particularly engraulis encrasicholus of europe and the mediterranean , are eaten by humans . they are a small oily fish that typically are filleted , semi - preserved with ten to 12 percent salt , and sometimes benzoic acid , and canned in oil ( herbst 2001 ; bender and bender 2005 ) . canned anchovies can last at least a year at room temperatures , and once opened can be refrigerated for at least two months if sealed airtight , covered with oil ( herbst 2001 ) .
anchovies are both popular and infamous for their strong flavor . in roman times , they were the base for the fermented fish sauce called garum that was a staple of cuisine and an item of long - distance commerce produced in industrial quantities . today , they are a key ingredient in caesar salad and spaghetti alla puttanesca , and are occasionally used as a pizza topping . because of the strong flavor they are also an ingredient in several sauces , including worcestershire sauce and many fish sauces , and in some versions of café de paris butter .
the strong taste that people associate with anchovies is due to the curing process . fresh anchovies , known in italy as alici , have a much softer and gentler flavor . in english - speaking countries , alici are sometimes called
white anchovies ,
and are often served in a weak vinegar marinade . this particular preservation method is associated with the coastal town of collioure in south east france . the white fillets ( a little like marinated herrings ) are sold in heavy salt , or the more popular garlic or tomato oil and vinegar marinade packs .
fishermen also use anchovies as bait for larger fish such as tuna and sea bass .
agbayani , e . family engraulidae : anchovies fishbase , 2004 . retrieved october 3 , 2007 .
bender , d . a . , and a . e . bender . a dictionary of food and nutrition . new york : oxford university press , 2005 . isbn 0198609612
herbst , s . t . the new food lover ' s companion : comprehensive definitions of nearly 6 , 000 food , drink , and culinary terms . barron ' s cooking guide . hauppauge , ny : barron ' s educational series , 2001 . isbn 0764112589
nelson , j . s . fishes of the world , 3rd edition . new york : john wiley & sons , 1994 . isbn 0471547131
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kari pihlaviita added the finnish common name
sapelisardelli
to
lycengraulis grossidens ( spix & agassiz , 1829 )
.
eol content is automatically assembled from many different content providers . as a result , from time to time you may find pages on eol that are confusing .
to request an improvement , please leave a comment on the page . thank you !
justification : listed as least concern in view of the large range extent , large population , apparently stable trend , and lack of major threats .
this fish occurs along the western atlantic and gulf of mexico coasts of north america , from maine to yucatán , mexico ( page and burr 2011 ) .
is distributed in coastal waters in the western atlantic from casco bay , maine south to florida keys and westward around the gulf of mexico south to yucatan ( accessed through fish base , www . fishbase . org , 26 july 2012 ) . this species is a nearshore , coastal , and estuarine species . it seldom occurs in waters deeper than 25 m , but has been collected in 27 - 36 m depths ( houde and zastrow ) .
this species is represented by a large number of occurrences ( subpopulations ) . total adult population size is unknown but presumably exceeds 1 , 000 , 000 . this fish is regarded as abundant in suitable habitat ( page and burr 2011 ) . trend over the past 10 years or three generations is uncertain but probably relatively stable .
feeds on zooplankton , predominately copepods , also on gastropods , isopods , mysid shrimps , and small fishes . in areas where
is abundant it is extremely important in estuarine and coastal food webs because it links secondary plankton production to fisheries output ( munroe and nizinski 2002 ) . in the chesapeake bay this species may live to be slightly more than three years olds and adults may attain a maximum length of 110 mm ( houde and zastrow ) .
anchoa spp . are caught as non - target species and / or bycatch off mexico ( fernández et al . 2011 ) .
currently , this species is of relatively low conservation concern and does not require significant additional protection or major management , monitoring , or research actions .
justification : this species has a wide distribution , no major threats , and occurs in marine protected areas . therefore , this species is listed as least concern .
this species is distributed from southern baja california and the gulf of california to northern peru . it is also found in cocos and the galapagos islands .
this is a pelagic schooling fish . it occurs in coastal waters over sand or gravel in the gulf of nicoya , costa rica , as well as over mud . it is found to 10m . it may spawn throughout the year in the gulf of nicoya , costa rica . it feeds on zooplankton , and it has pelagic eggs and larvae ( whitehead et al . 1988 ; whitehead and rodríguez - sánchez 1995 ) .
this species has no major threats . however , it ( as some other anchovies ) may be affected by overfishing due to bycatch methods . there are no known commercial uses for this species . this species is taken in subsistence fisheries ( whitehead et al . 1988 ; whitehead and rodríguez - sánchez 1995 ) . this species ( as some other anchovies ) may be affected by overfishing due to bycatch methods .
there are no known conservation measures for this species . however , this species ' distribution includes a number of marine protected areas in the central tropical eastern pacific region .
overfishing occurs when more fish are caught than the population can replace through natural reproduction . gathering as many fish as possible may seem like a profitable practice , but overfishing has serious consequences . the results not only affect the balance of life in the oceans , but also the social and economic well - being of the coastal communities who depend on fish for their way of life .
billions of people rely on fish for protein , and fishing is the principal livelihood for millions of people around the world . for centuries , our seas and oceans have been considered a limitless bounty of food . however , increasing fishing efforts over the last 50 years as well as unsustainable fishing practices are pushing many fish stocks to the point of collapse .
more than 30 percent of the world ' s fisheries have been pushed beyond their biological limits and are in need of strict management plans to restore them . several important commercial fish populations ( such as atlantic bluefin tuna ) have declined to the point where their survival as a species is threatened . target fishing of top predators , such as tuna and groupers , is changing marine communities , which lead to an abundance of smaller marine species , such as sardines and anchovies .


cornu aspersum , known by the common name garden snail , is a species of land snail .

as such it is a terrestrial pulmonate gastropod mollusc in the family helicidae , which include the most commonly familiar land snails .

of all terrestrial molluscs , this species may well be the most widely known .

in english texts it was classified under the name helix aspersa for over two centuries , but the prevailing classification now places it in the genus cornu .

cornu aspersum is native to the mediterranean area and western europe , but whether deliberately or accidentally , humans have spread it to temperate and subtropical areas worldwide .

the snail is relished as a food item in some areas , but it is also widely regarded as a pest in gardens and in agriculture , especially in regions where it has been introduced accidentally and where snails are not usually considered to be a menu item .

cornu aspersum

tracing the invasion of the mediterranean land snail cornu aspersum aspersum becoming an agricultural and garden pest in areas recently introduced .
cornu aspersum ( müller , 1774 ) brown garden snail ( jacksonville shell club ) .
cornu aspersum : aggregated on a wall . ( photo : © d . medcalf , wikipedia )
theba pisana ( a ) , cornu aspersum ( b ) , plutonia lamarckii ( c ) .
worms - world register of marine species - cornu aspersum ( o . f . müller , 1774 )
larval development of angiostrongylus chabaudi , the causative agent of feline angiostrongylosis , in the snail cornu aspersum .
note : the surface sculpturing of the shell can be used to distinguish between helix spp . and cornu aspersum . the shell of cornu aspersum is characteristically wrinkled , while the shell surface of helix species lack wrinkles .
mating in cornu aspersum takes place differently to helix po - matia , as the photographer hans josef köhler has observed .
larval development of angiostrongylus chabaudi , the causative agent of feline angiostrongylosis , in the snail cornu aspersum . - pubmed - ncbi
figure 2 . extended adult brown garden snail , cornu aspersum ( müller ) . photograph by bill frank , jacksonville shell club .
the illustration of cornu aspersum in ignaz von born ' s
index rerum naturalium musei caesarei vindobonensis
( 1778 - 1780 ) .
there are also distinct differences between helix pomatia and cornu aspersum concerning the genital apparatus of both species . with helix lucorum , cornu aspersum shares a conspicuous diverticulum , an appendage to the bursa copulatrix channel ( see :
the genital apparatus of helix pomatia
) .
though cornu aspersum is a relative of helix pomatia , it does not belong to the genus helix . however though , it is often misleadingly called helix aspersa . see also :
the cornu problem
.
a cornu aspersum exhibit in the vienna natu - ral history museum . note : the obsolete name is still used here . [ rn ]
also know as cornu aspersum , this species is one of the two main types of snail eaten as a delicacy called escargot in europe .
cornu aspersum . note yellow tubercles on the body of the animal . ( photo : © r . forsyth , smithers , bc , canada )
the latest additions to the distribution pattern of cornu aspersum happened during modern times , the snail introduced practically world wide with fruit and vegetable transports .
during the mating procedure , which , in cornu aspersum other than in helix pomatia , takes place in the horizontal position common among land snails , a love dart may be applied that injects a hormone secretion into the partner ' s body . this is specially being researched in cornu aspersum .
notes helix aspersa müller is also referred to as cantareus aspersus ( müller ) , cryptomphalus aspersus ( müller ) and cornu aspersum ( müller ) .
ansart a , guiller a , madec l ( 2009 ) cabi invasive species compendium : cornu aspersum . in : cabi editor . london . 19p .
photo ( left ) : the land snail cornu aspersum is a common inhabitant of gardens in europe , california and south america . © c . levesque
figure 4 . the brown garden snail , cornu aspersum ( müller ) , showing yellow coloring . photograph by paul m . choate , university of florida .
figure 6 . infestation of brown garden snail , cornu aspersum ( müller ) , on a citrus tree in california . photograph by division of plant industry .
cornu aspersum feeds on a wide variety of vegetables , flowers , fruit , ornamental plants and weeds . dead animal tissue and paper products are also eaten .
figure 3 . shells of the brown garden snail , cornu aspersum ( müller ) , displaying various color shades encountered . photograph by bill frank , jacksonville shell club .
figure 1 . adult brown garden snails , cornu aspersum ( müller ) . photograph taken in southern san diego county , california . photograph by bill frank , jacksonville shell club .
citation : guiller a , martin m - c , hiraux c , madec l ( 2012 ) tracing the invasion of the mediterranean land snail cornu aspersum aspersum becoming an agricultural and garden pest in areas recently introduced . plos one 7 ( 12 ) : e49674 . urltoken
compared to the roman snail ( helix pomatia ) the shell of cornu aspersum is usually a bit smaller . it is between 25 and 40 millimetres wide and between 25 and 35 millimetres high . usually the shell of cornu aspersum , compared to helix pomatia , is a bit more spherical , the last of the 4 ½ to 5 shell whorls is clearly wider than the others .
one thing , finally , remains for sure : cornu aspersum or cantareus aspersus , whichever of both name is used , the brown garden snail ' s correct name definitely is not helix aspersa .
guiller a , madec l ( 2010 ) historical biogeography of the land snail cornu aspersum : a new scenario inferred from haplotype distribution in the western mediterranean basin . bmc evol biol 10 : 18–37
in central europe cornu aspersum has not been able to spread in a continuous area , because the more continental climate bears more severe winters . which is why cornu aspersum , not being able to cope with too low temperatures , is only to be found in the relatively temperate areas of the upper and lower rhine areas , in austria even only introduced in a little spot near vienna simmering .
geographical distribution of cornu aspersum ( orange ) , helix pomatia ( yellow ) and helix lutescens ( green ) in europe . source : kerney et al . ( 1983 ) ( within limits of discussion area ) .
plain pest surveillance unit . ( 2013 , november 1 ) . cornu aspersum * ( european brown garden snail ) - fact sheet . retrieved december 9 , 2014 , from urltoken sheet / eng / 1326334238293 / 1326341285764
the helix aspersa is also known as “european brown garden snail , ” but its scientific is under discussion . some scientists think this snail is “ cornu aspersum , ” “ cantareus aspersus ” or “ cryptomphalus aspersus . ”
size distribution of the de novo assembled contigs from the transcriptome of the foot muscle of aestivating c . aspersum
dekle , g . w . , & fasulo , t . r . 2011 . brown garden snail , cornu aspersum ( müller , 1774 ) ( gastropoda : helicidae ) . university of florida , ifas extension eeny , 240 .
the new study was prompted by the refurbishment of a
small suburban garden
in the spring of 2001 . about 120 new plants were installed , but it took only a few days for cornu aspersum snails to ruin them .
in this study , we have focused on investigating correlations between shell colouration and constitutive as well as inducible po activity in different shell colour morphs of cepaea hortensis , theba pisana and cornu aspersum maximum , applying zymosan a as an immunostimulant .
the common garden snail - helix aspersa / cornu aspersum - crawling up on a window in a cold winter night in israel , 29 . 11 . 2014 שבלול הגינה זוחל במעלה החלון בליל חורף קר בישראל , כ
ט בנובמבר , 2014
chase , r . and darbyson , e . ( 2008 ) . differential survival of allosperm by location within the female storage organ of the snail cornu aspersum ( müller , 1774 ) . canadian journal of zoology 86 , 1244 - 1251 .
nicolai a . the impact of diet treatment on reproduction and thermophysiological processes in the land snails cornu aspersum and helix pomatia . m . sc . thesis , the university of sheffield . 2010 . available : urltoken . accessed 2015 march 5 .
the example of the brown garden snail ( cornu aspersum ) finely demonstrates , how the scientific name of a snail species has developed over the centuries , what the procedures are in providing scientific names ( or what they are not ) , and that sometimes there are special cases .
gaitán - espitia jd , scheihing r , poulin e , artacho p , nespolo rf ( 2013 ) mitochondrial phylogeography of the land snail cornu aspersum ? : tracing population history and the impact of human - mediated invasion in austral south america . evol ecol res 15 : 1–18 .
the brown garden snail lives in various habitats . besides synanthropic habitats in gardens and parks made by man , it also lives in coastal dunes , grove and bush land , between rocks . in contrary to helix pomatia , cornu aspersum is not that dependant on a limestone underground . but it neither often appears in the fens or in deep forest , nor on open meadows , where there is little visual shelter against predators . instead , it prefers habitats with hiding places in rock and wall crevices or below thick vegetation cover . in the mountains cornu aspersum climbs up to below 1000 metres .
c . aspersum causes extensive damage in orchards ( creating holes in fruit and leaves ) and to vegetable crops , garden flowers and cereals .
citation : gaitán - espitia jd , nespolo rf , opazo jc ( 2013 ) the complete mitochondrial genome of the land snail cornu aspersum ( helicidae : mollusca ) : intra - specific divergence of protein - coding genes and phylogenetic considerations within euthyneura . plos one 8 ( 6 ) : e67299 . urltoken
gaitán - espitia jd , bruning a , mondaca f , nespolo r ( 2013 ) intraspecific variation in the metabolic scaling exponent in ectotherms : testing the effect of latitudinal cline , ontogeny and transgenerational change in the land snail cornu aspersum . comparative biochemistry and physiology part a : molecular & integrative physiology 165 : 169–177
cornu aspersum is an invasive alien snail with shells up to 32 mm in diameter and is originally from europe and is , widespread in new zealand . active at night or during wet weather it is slow moving , using a slime trail . it has a well - developed sense of smell and / taste .
cornu aspersum is a hermaphrodite , meaning it has male and female reproductive organs . it needs a partner to create juveniles . the snails ' hook up ' on the right sides of their heads , this can last for up to 12 hours . they inject each other with spermatozoa and both will become impregnated .
genetic diversity estimates and results of demographic tests for cyt b and 16s genes in main geographical subdivisions of native and invasive c . a . aspersum range .
the brown garden snail ( cornu aspersum ) is one of the most widely spread land snail species in the world . originated from the mediterranean it is today found is as far away areas like northern america ( south carolina , louisiana and california ) , southern america ( south of brazil ) , australia and new zealand , as well as in south africa .
( cornu ) : cryptomphalus charpentier 1837 is a younger synonym of cornu born 1778 . the name has been in frequent use for the last years , though it was based on a teratological specimen , so a suppression in favour of cryptomphalus is hardly possible ( pilsbry 1948 : 1091 , zilch 1962 : 21 anm . 58 , waldén 1976 : 25 ) [ vp , f ] .
being an inhabitant of areas with oceanic temperate climate , the brown garden snail is not able to to endure the severe central european winters as efficiently as helix pomatia does . in contrary , cornu aspersum does not produce a hibernation lid , but instead closes the aperture with a mucus membrane and hides in a sheltered place , where often numerous snail gather for the winter .
cornu aspersum is herein reported for the first time as a suitable intermediate host of c . vulpis . this snail species may play an important role for the infection of animals living in regions of the mediterranean basin . in addition , this study provides more details on the morphological descriptions of l1 and l3 and supports future investigations on the epidemiology of this little known parasite .
although i began my career working with mammals , i have used molluscan models for many years . today , my experimental animal is the brown garden snail cornu aspersum ( formerly known as helix aspersa ) . my research concerns the complicated sex life of this animal . students working in my laboratory pursue either behavioural projects or neurobiological projects , or sometimes a combination of the two .
there is concern about this snail affects indigenous biodiversity through selective feeding on foliage of herbaceous plants and the seedlings of a range of species , cornu aspersum represents a significant threat to natural processes of vegetation regeneration . many of the introduced terrestrial molluscs brought with them to new zealand a range of parasites , including mites , ciliate protozoa , microsporidia , and nematodes . with increasing contact between the alien and indigenous species , there is potential for transmission of these parasites and pathogens to our indigenous fauna . this transmission has already occurred in the case of the parasitic mite riccardoella limacum in urban areas and in egmont national park . cantareus aspersus provides a reservoir of riccardoella that is likely to infest sympatric indigenous species cornu aspersum snails can live for 2 - 3 years .
hallman , g . j . ( 2016 ) . phytosanitary irradiation of the invasive herbivorous terrestrial snail cornu asper - sum ( stylommatophora : helicidae ) . florida entomologist 99 ( s2 ) : 156 - 158 .
juřičková l . & kapounek f . 2009 . helix ( cornu ) aspersa ( o . f . müller , 1774 ) ( gastropoda : helicidae ) in the czech republic . malacologica bohemoslovaca 8 : 53 - 55 .
figure 5 . adult brown garden snails , cornu aspesrum ( müller ) , size comparison on an oversized 160 mm lemon . photograph taken in southern san diego county , california . photograph by bill frank , jacksonville shell club .
from there the species has been distributed synanthropically in garden areas , but also because of its use by man as food in the climatically temperate areas of the mediterranean and adjacent coastal areas of the atlantic . it is assumed today , that cornu aspersum reached the british isles long before the romans did , for example already with the celts . in many other parts of the empire , though , the species was introduced due to the spreading of roman civilisation .
if
helix
aspersa must be placed in one genus with
helix
aperta on the grounds of genital characters , needs further examination . using the genus name cantareus risso 1826 , however , appears to be a problematic solution . it cannot be proved that the name cornu copiae was chosen by von born with the intention and the knowledge of naming a teratological specimen . therefore , iczn ( see above ) is not concerned and as a consequence , cornu is an available genus name
a : phylogenetic relationships of 406 individual sequences of c . a . aspersum . b : focus on the relationships inside the west lineage represented by the w1 and w2 sub - lineages ( see legend of figure 2 for details ) .
different morphs of test snails ; pale and dark morph of c . hortensis ( a ) , pale and dark morphs of t . pisana ( b ) , and pale and dark morph of c . aspersum maximum ( c ) .
the brown garden snail ( european brown snail ) , cornu aspersum , was described by o . f . müller from specimens collected in italy . this plant feeder has been disseminated into many parts of the world intentionally as a food delicacy , and accidentally by the movement of plants and by hobbyists who collect snails . it was introduced to california in the 1850s as a source of escargot . it has adapted well to california and is a very troublesome pest of crops and ornamentals ( capinera 2001 ) .
one - year - old cornu aspersum ( n = 100 ) snails were purchased from a farming centre located in barletta ( puglia , italy ) . the snails were housed in a plastic box in a temperature - controlled room ( 21 ± 1 °c ) and fed every second day with lettuce and water for two months until beginning of the study . at the arrival and one day prior to the infection , ten snails were sacrificed , digested and microscopically examined to assess the absence of any helminth infection .
suitability and developmental time of c . vulpis in the common garden snail cornu aspersum ( = helix aspersa ) was assessed at selected days post - infection ( i . e . 3 , 6 , 10 , 15 , 20 and 180 ) . nematodes were preserved in 70 % ethanol , cleared and examined as temporary mounts in glycerol for morphological descriptions of first - and third - stage larvae . in addition , nematodes collected from the dog and the experimentally infected snails were molecularly analysed by the amplification of the nuclear 18s rrna gene .
cornu aspersum is a quite intriguing species from the point of view of ecology and evolution and its potential use in medical and environmental applications . it is a species of economic importance since it is farmed and used for culinary purposes . however , the genomic tools that would allow a thorough insight into the ecology , evolution , nutritional and medical properties of this highly adaptable organism , are missing . in this work , using next - generation sequencing ( ngs ) techniques we assessed a significant portion of the transcriptome of this non - model organism .
laboratory and field experiments have been conducted in egypt to investigate the molluscicidal activity of weed extracts from ambrosia martine , citrullus colocynthis , cymbopogon proximus and glinus sp . , and of five fertilizers , against c . aspersum . ambrosia martine extract was shown to be the most toxic , followed by c . colocynthis and c . proximus , while glinus sp . was the least effective . snail mortality increased gradually with time . some commonly used fertilizers showed considerable initial and residual molluscicidal effects . ferrous sulfate was the most effective against c . aspersum , followed by ammonium nitrate , ammonium sulfate and superphosphate ( zidan et al . , 1997 ) .
a . cantonensis has been detected recently in black rats ( rattus rattus ) in the canary islands [ 14 ] , an archipelago very rich in gastropod species , among which many are endemic [ 15 ] . to our knowledge , this is the only location in europe where the parasite has been detected . nevertheless , there are no data regarding which snail and / or slug species are acting as intermediate hosts in the canary islands . hence , the possible sources of human infection have not been identified until now . the aim of this study was to determine the possible role of three mollusc species , plutonia lamarckii , cornu aspersum and theba pisana , as intermediate hosts of a . cantonensis in tenerife .
here we found that the size of the protein - coding , trna and rrna genes is conserved within eupulmonata ( table 2 ) . the start and stop codon of the protein - coding genes in c . aspersum shows similarities to those of the other species analyzed . the presence of incomplete stop codons in c . aspersum seems to occur frequently in protein - coding genes of most of the mollusc mitochondrial genomes sequenced to date [ 4 ] , [ 12 ] , [ 16 ] , [ 46 ] , and it has been suggested that the transcripts of those genes would be modified to form a complete stop codon via post - transcriptional polyadenylation [ 47 ] .
the brown garden snail is about 1 inch in diameter with a snail that is 25 - 40mm wide with 4 to 5 whorls . and has a distinct brown and grey color pattern on the shell . cornu aspersum is a terrestrial air - breathing snail that is edible . in britain brown garden snail farming is currently a booming cottage industry . likewise , this snail has been used for centuries in traditional medicine to treat sore throats . additionally , this species is an agricultural and garden pest that seems to have a preference for citrus trees and fruits . brown garden snails exhibit a very strong homing ability that causes them to aggregate in large numbers ( pictured above on the right ) , making them difficult to control .
a patented copper complex , formulated as a spray and referred to as copper silicate , shows molluscicidal activity ( davis et al . , 1996 ) . the product was tested in both spray and granular formulations against c . aspersum in a vineyard and a citrus orchard in western australia , where it was compared with copper oxychloride spray . the copper complex spray demonstrated a high level of repellence to c . aspersum for extended periods in the field , and proved superior to the other products tested in reducing snail numbers in vines / trees . the product has potential in the management of pest molluscs , especially as part of ipm programmes , but it seems not to have been further developed .
each mature snail will mate several times in a single season and egg laying starts 3–6 days after first mating but can occur within hours of subsequent matings . cornu aspersum deposit their eggs into pockets in moist soil . these pockets are usually excavated by the snails themselves through movement of the head and anterior foot , though occasionally natural cavities will be used . after oviposition , the nests are covered with soil and abandoned . the number of eggs in each clutch varies from 10 to 170 . the incubation period of the eggs varies according to the relative humidity of the incubation environment . the average is 13 to 19 days . the young hatchling snails remain in the nest for 1–16 days . during this time , cannibalism on sib eggs can occur .
the scientific species name of the brown garden snail , aspersum , means
spotted
, referring to the shell ' s colour pattern . on a light yellowish to brownish base colour there are several darker longitudinal bands , that may be spotted in a lighter colour . laterally there are yellow zig - zagging stripes . the shell surface is characteristically wrinkled .
in order to understand what nematode species were naturally infecting these shells , they were homogenized and using pcr and dna sequencing of the nuclear internal transcribed spacer gene ( its ) of several c . nemoralis shells and 16 c . aspersum shells , the nematodes present in c . nemoralis were identified as p . hermaphrodita , phasmarhabditis californica , eucephalobus sp . and , based on morphology alone , a mermithid sp . ( fig . 3b ) . all c . aspersum shells were found to have c . elegans , a nematode associated with slugs and snails 21 , present in great numbers encased in their shells . therefore , wild caught and farmed snails frequently use their shell to kill a diverse range of nematode species .
in the usa , aphis has produced containment guidelines to assist a researcher , educator , or commercial entity to design , build , maintain and operate a facility for rearing nonindigenous snails in the usa , including c . aspersum ( aphis , 2002 ) , the non - respect of which is subject to civil and / or criminal penalties and loss of permits .
the typical anthropogenic aspersum form is now present in many zones having mediterranean , temperate and even subtropical climates , on the american and african continents , as well as on the mascarene islands , oceania and asia [ 8 ] . more recent events related to human activities are more likely responsible for long distance dispersal . as the ancient romans were very fond of snails , c . a . aspersum may well have been introduced early on by these invaders who would be the first people rearing snails [ 18 ] . besides intentional introductions for farming purposes to produce food , it has been accidentally introduced by the movement of plants and vegetables . quarantine prevention measures for plant material have been adopted in several states in the usa and in canada in the 1970s .
in 1778 , ignaz von born ( 1742 - 1791 ) wrote another book in vienna . he had to publish a description of the imperial natural collections in what is today the vienna natural history museum . in the book ,
index rerum naturalium musei caesarei vindobonensis
he published a snail he named cornu copiae ( the horn of plenty ) . this snail shell clearly was misshaped , what we would today call a scalariform shell .
cornu aspersum is a moderately sized snail with a heliciform shell ranging in height from 20 - 35 mm and width from 25 - 40 mm ( rarely 45 mm ) . the shell is yellow - brown and may possess darker brown spiral stripes interrupted by lighter , irregular markings and streaks , creating a banded appearance . there are irregular dimples on the shell . there are 4 1 / 2 to 5 whorls . the aperture ( mouth ) is large and rounded and has a lip that is white , faintly thickened and slightly recurved . in adults of this species , the umbilicus is normally absent or closed , though it may open to form a narrow slit in rare cases . the body of the animal is grey to pale brown - ochre and the tubercles are yellow . the mantle is somewhat black and speckled with grey - yellow .
the canadian food inspection agency ( cfia , 2014 ) considers c . aspersum to be a plant pest and have quarantines established for preventing the importation of the snail in plant and soil matter . the united states department of agriculture requires a permit to import snails ( or slugs ) into the usa and between states within the usa from aphis , animal plant health and inspection service ( aphis , 2002 ) .
adults and larger sized juveniles are likely to be visible among the host material or attached to the transporting containers . they may also be hidden in protected locations , sealed into their shells , avoiding desiccation . check the underside of containers and their rims , etc . small snails and eggs in soil could be difficult to find . c . aspersum likes to hide in crevices and will over winter in stony ground .
phenoloxidase ( po ) activity levels in different morphs of c . aspersum maximum ; test run 2 with base levels and levels 6 h or 24 h after hemolymph ( hl ) withdrawal considering other individuals as for the base levels , and levels after 6 h or 24 h zymosan a - exposure ( mean + sd ; n = 15 ; p ≤ 0 . 0024 : * after bonferroni - corrections for 21 comparisons ) .
adult individuals of cornu aspersum were collected from three populations in chile across a latitudinal range of approximately 1300 km : la serena ( 29°54′ s , 71°15′ w ) , constitución ( 35°20′ s , 72°25′ w ) and valdivia ( 39° 38′ s , 73° 5′ w ) . we selected these three localities based on their significant genetic [ 27 ] and climatic differentiations [ 25 ] . snails were transported to the laboratory , and maintained at 20°c with a fixed photoperiod of 14l : 10d . additionally , snails were kept in humid litter soil and water was given ad libitum until dna extractions were performed . dna from a single snail of each population was obtained by the isolation of intact mitochondria from approximately 120 mg of fresh foot tissue , using the mitochondrial isolation kit for tissue ( thermo scientific ) . the isolated mitochondrial pellet of each snail was used for the mtdna extraction using the mitochondrial dna isolation kit ( biovision ) .
as terrestrial molluscs have many natural enemies , there has been strong interest in the biological control of c . aspersum using other , predatory snails ( e . g . fisher and orth , 1985 ) . however , as most of these predatory snails are not host - specific , they are not appropriate to use in control programmes in which effects on non - target species are of concern ( cowie , 2001 : barker and watts , 2002 ) .
in gastropods the length of the mitochondrial genomes usually vary from 13 to 17 kb ( e . g . , 13670 bp in biomphalaria glabrata , 17575 bp in diodora aspera ) , however , there are some exceptions such as the ribbed limpet lottia digitalis , with a length of 26835 bp [ 43 ] . here we found that the length of the mitochondrial genome of c . aspersum is similar to the other eupulmonata species examined ( table 2 ) . although the mitochondrial genomes of gastropods are very compact [ 3 ] , they contain intergenic non - coding regions of variable size that are responsible for differences in genome size [ 16 ] . in this regard , c . aspersum , c . nemoralis and c . obtusus have an at - rich intergenic region , that contains the putative origin for mitochondrial dna replication ( por ) , which is almost five times longer than other eupulmonata species . this difference is probably due to insertions of tandem repeats in the por region [ 44 ] , which can be caused by gene rearrangements experienced by these species . accordingly our results show that the por region of c . aspersum , c . nemoralis and c . obtusus are located between cox3 and trna - s1 ( i . e . , first serine ) , whereas in the other pulmonata species is located between cox3 and trna - i ( fig . 3 ) .
considering all the above , c . aspersum is a quite intriguing species from the point of view of ecology and evolution and its potential use in medical and environmental applications . at the same time , it is a species with an already acknowledged economic importance as a farmed species used for culinary purposes . however , the genomic tools that would allow a thorough insight into the ecology , evolution , nutritional and medical properties of this highly adaptable organism , are missing .
suitability and developmental time of c . vulpis in c . aspersum was assessed by artificially digesting two portions ( i . e . muscular foot and viscera ) of two snails per time - point ( i . e . 3 , 6 , 10 , 15 , 20 days post - infection , dpi ) . in addition , six infected snails were gradually placed at 4 °c in order to favour gastropods hibernation , replaced at laboratory condition , and analysed at 180 dpi .
the go terms relating to ( a ) biological process , ( b ) molecular function and ( c ) cellular component categories assigned to the contigs of the de novo assembly of c . aspersum transcriptome . the arrow and the ( − ) and ( + ) signs on the left of each pie - chart legend , indicate the direction of increase in the number of contigs corresponding to the biological process , molecular function or cell compartment denoted in the respective pie - chart legend
sampling characteristics ( status , origin , geographic coordinates , population code , collectors , sampling date ) of cornu aspersum aspersum populations analyzed in this study and in [ 10 ] , [ 13 ] . population code : ( i ) native : a , algeria ; ba , balearic islands ; c , corsica ; can , canary islands , ct , crete ; cr , croatia ; e , spain ; f , france mainland ; g , greece mainland ; gb , great - britain ; i , italia ; m , morocco ; mal , malta ; p , portugal ; sa , sardinia ; si , sicily ; t , tunisia ; tu , turkey ; ( ii ) : invasive : ca , california ; ch , chile ; ha , haiti ; hw , hawaiian islands ; nz , new zealand ; saf , south africa ; tx , texas ; ut , utah ) . mitochondrial diversity overall samples analyzed . sample size ( n ) , name of individuals sequenced ( population code and individual number ) and their corresponding cyt b and 16s rna haplotypes and genbank accession numbers . two kinds of haplotype codes are provided : ( i ) the sequence name for unique haplotype , ( e . g . a1 . 1 ) , ( ii ) hk ( k up to 71 distinct types ) and sk ( k up to 56 distinct types ) for cyt b and 16s respectively for haplotypes shared by distinct individuals ( * : haplotypes published in [ 13 ] ; # haplotypes published in [ 10 ] ) .
specimens of c . aspersum digested before the infection ( n = 10 ) were negative for helminth infections . out of 115 larvae recovered from infected gastropods ( mean of 9 . 58 larvae per snail ) , 36 ( 31 . 3 % ) were localised in the foot and 79 ( 68 . 7 % ) in the viscera . the 18s rdna sequences obtained from larvae collected from the dog and the snail tissues displayed 100 % identity to the nucleotide sequence of c . vulpis .
handpicking with subsequent destruction of animals is the oldest method of control of pest molluscs . even though it is time intensive , it can be effective as c . aspersum is gregarious and can be found in huge numbers under snail traps . other places around susceptible plants where snails can hide during daily or seasonal dormancy , such as boards , debris and dense cover such as ivy , branches growing near the ground , stones , must be removed ( barker and watts , 2002 ; dreistadt et al . , 2004 ) .
in south african viticultural regions , c . aspersum feeds essentially on the developing foliar buds and young leaves of the vines . in kiwifruit vineyards ( california , new zealand ) , damage occurs on the flowers , not the fully developed fruit , since snails consume only the sepal tissue around the receptacle area . damage to the sepals can be detrimental by increasing the development of the fungus botrytis cinerea during cold storage of fruits , and moreover , the slime trail mucus stimulates germination of b . cinerea conidia ( michailides and elmer , 2000 ) .
the annotation success of the transcriptome of c . aspersum was relatively low . this is probably due to the very limited number of annotated reference genomes existing for mollusc species , especially terrestrial ones . several biological processes being active in the aestivating species were revealed through the association of the transcripts to enzymes relating to the pathways . the genomic tools provided herein will eventually aid in the study of the global genomic diversity of the species and the investigation of aspects of the ecology , evolution , behavior , nutritional and medical properties of this highly adaptable organism .
under laboratory conditions c . nemoralis can encapsulate and kill parasitic nematodes but does this process occur in nature ? shells of c . nemoralis and its congener ( c . hortensis ) were collected from sand dunes in north west england and the north of scotland , respectively ( figure s1 , table s1 ) and examined for presence of nematodes . nematodes were found fixed in shells of both wild c . nemoralis and c . hortensis shells ( fig . 3a–c ) . the numbers of nematodes found in c . nemoralis shells collected from several locations in north west england and c . hortensis shells from northern scotland differed significantly ( c . nemoralis x 2 ( 4 ) = 93 . 06 , p < 0 . 0001 and c . hortensis χ 2 ( 3 ) = 430 . 54 , p < 0 . 0001 ) . in some cases 60 % of the shells collected had nematodes present in numbers ranging from 1 to 101 nematodes per shell . also cornu aspersum shells purchased from an escargot farm in the u . k . had nematodes present within 100 % of shells ( n = 136 ) , having a mean of 31 ± 2 nematodes per shell . thus , several snail species isolated from the wild can encase and kill nematodes .
phenoloxidase ( po ) activity levels in different morphs of c . aspersum maximum ; test run 1 with base - levels and levels 6 h or 24 h after hemolymph ( hl ) withdrawal considering the same individuals as for the base levels , and base levels and levels after hemolymph ( hl ) withdrawal plus 24 h zymosan a - exposure considering the same individuals as for the base levels ( mean + sd ; n = 15 ; 0 . 000435 < p ≤ 0 . 0022 : * ; p ≤ 0 . 000435 : * * after bonferroni - corrections for 23 comparisons ) .
aphis ( 2009 ) encourages people using c . aspersum in classrooms , nature facilities , or keeping them as pets to turn them in voluntarily . some us states have created an ‘invasive species hotline ' ( e . g . oregon , hawaii , michigan ) for early detection of potential pest invaders . residents of the state of hawaii are asked to be on the lookout for this pest and to call a resident agricultural specialist on their respective island if they see this snail . it is currently recorded on only two of the hawaiian islands ( cowie , 1997 ; cowie et al . , 2008 )
the size of the mitochondrial genome of c . aspersum is 14050 bp ( fig . 2 ) , and contains 13 protein - coding genes , 22 transfer rna genes , and two ribosomal rna genes ( fig . 2 ) . from these 37 genes , 13 are coded on the minus strand : trna - q , trna - l2 , atp8 , trna - n , atp6 , trna - r , trna - e , 12s - rna , trna - m , nadh3 , trna - s1 , trna - t , and cox3 ( fig . 2 ) . the mitochondrial genome of c . aspersum also includes nine small size - variable intergene spacer regions ranging from 2 to 47 bp , and a large at - rich ( 81 . 2 % ) intergenic region ( 186 bp ) located between cox3 and trna - s2 ( i . e . , second serine ) . this large intergene spacer contains three tandem repeated sequences ( attatta , attagtg and tataaatat ) of 7 bp length , distributed across the non - coding region . there are small gene overlaps at 14 gene borders , the largest has a length of 10 nucleotides and is located between nadh3 and trna - s1 ( i . e . , first serine ) . the overall base composition of these mitochondrial genomes shows a high at content ( 69 . 9 % ; table 2 ) .
the overall at - content of the mitochondrial genome of c . aspersum is higher in comparison to c . nemoralis and c . obtusus ( similar to that one observed in e . herklotsi and a . caerulea ) but lower in comparison to s . putris ( table 2 ) . nevertheless , the average at - content of the whole stylommatophora clade is higher in comparison to other eupulmonata orders ( table 2 ) . the high at - content is a common feature of most animal mitochondrial genomes [ 3 ] , and it has been suggested that is the result of directional mutation pressure due to the lower energy requirement for opening the dna strands with high at content [ 45 ] .
this edible snail lives in gardens and along roadsides and consumes both living and dead plant material . this species is recorded as a pest of citrus in california . in many parts of the world , c . aspersum attacks vegetables ( carrot , cabbage , lettuce , onion , tomato ) , cereals ( oats , wheat , barley ) , flowers ( sweet - pea , lilies , carnation , aster , pansy ) , ornamental ( california boxwood , hibiscus , rose ) and fruit trees ( peach , plum , apple , apricot ) . the garden snail can lay as many as 80 eggs per clutch . the juveniles of this species will achieve maturity in 1 - 2 years . longevity is approximately 5 years .
each circle represents a haplotype , and circle size is proportional to haplotype frequency . colors indicate native ( grey ) versus invasive ( red ) status of haplotypes . branch lengths are approximately equal to inferred mutational steps . a : phylogenetic relationships of 390 individual sequences of c . a . aspersum and schematic geographic location of the main haplogroups defined ( east versus west lineages are represented by pink and green color respectively , the kabylia putative hybrid zone is in blue ) . c . a . maximum is used as outgroup . b : focus on the relationships inside the west lineage represented by the w1 and w2 sub - lineages . dashed lines represent superflous links deleted using the network’s mp option . haplotype codes according to those in table s1 .
diversity measures from microsatellite loci comparing 10 native and 10 invasive populations of c . a . aspersum . he , expected heterozygosity per locus and per population ; na , number of alleles per locus and per population , ae : allelic richness per locus and per population ( based on minimum sample size of 6 diploid individual ) . population code : ( i ) native : ba , balearic islands ; e , spain ; f , france mainland , si , sicily ; ( ii ) : invasive : ca , california ; ch , chile ; hw , hawaiian islands ; nz , new zealand ; saf , south africa . n : number of individuals analyzed per population ; n * , number of individuals used for analyses after discarding individuals with too much missing data .
in this study , we report the first transcriptome originating from the foot muscle of an aestivating c . aspersum individual . the annotation success was relatively low ( 30 . 6 % ) . this is due to the very limited number of annotated reference genomes existing for mollusc species , especially terrestrial ones . the 9445 contigs were classified into biological and molecular processes as well as into cellular component categories . several biochemical pathways being active in the aestivating species were revealed through the association of the transcripts to enzymes relating to the pathways . after scanning the transcriptome for ssrs , 563 ssr markers were recovered . the genomic tools provided herein will eventually aid in the study of the global genomic diversity of the species and the investigation of aspects of the ecology , evolution , behavior , nutritional and medical properties of this highly adaptable organism .
shells of c . aspersum were homogenized using sterile plastic eppendorf pestles and dna was then extracted using a qiagen dna extraction kit . individual nematodes in c . nemoralis shells were extracted using sterile scalpel blades . dna was extracted using genejet genomic dna purification kit ( thermo fisher scientific ) and the its gene was amplified using the primer pair n93 5′ - ttgaaccgggtaaaagtcg - 3′ and n94 5′ - ttagtttcttttcctccgct - 3′ . pcr conditions were : 2 mins at 95 °c , followed by 35 cycles of 15 s at 95 °c , 15 s at 50 °c and 2 mins at 72 °c and finally 7 mins at 72 °c . pcr products were then purified using genejet pcr purification kit ( thermo fisher scientific ) . the pcr product was sequenced in forward and reverse directions using the same primers previously described and the sequence was compared with genbank sequences using blastn searches .
nucleotide translated sequences for the protein - coding genes from the three mtdna sequences of c . aspersum , and other euthyneuran species ( table 1 ) were aligned using the l - ins - i strategy from mafft [ 37 ] . nucleotide alignments were generated using the amino acid alignment as a template using the web - based program translatorx [ 38 ] . our taxonomic sampling included representative species of pulmonata , opistobranchia , caenogastropoda and vetigastropoda clades ( table 1 ) . the last two groups were used as outgroups . the nucleotide and amino acid composition were estimated by geneious v 4 . 8 . 5 [ 32 ] . individual alignments were concatenated prior to phylogenetic analysis . additionally , intra - specific divergences for each protein - coding gene were calculated based on these alignments . a p - distance method was performed using 1000 bootstrap replications for variance estimation using the program mega v . 5 . 1 [ 31 ] .
the gene order on the c . aspersum mitochondrial genome is the same as previously described for cepaea nemoralis ( fig . 3 ) . however , few differences were found when it was compared to c . obtusus , in which two trnas ( trna - a and trna - p ) are switched ( fig . 3 ) . similarly , when it was compared to e . herklotsi , s . putris and a . caerulea ( i . e . , stylommatophora ) , some additional switches were detected in nine trnas ( trna - d , trna - c , trna - f , trna - y , trna - w , trna - g , trna - h , trna - t , trna - s2 ) and one protein - coding gene ( nadh 4 ) ( fig . 3 ) . almost all of these differences are conserved in other pulmonata species with the exception of trna - p that is inverted ( fig . 3 ) .
intra - specific divergence analysis for each of the 13 protein - coding genes in c . aspersum revealed the existence of genes with high ( nadh1 , nadh3 and nadh4 ) intermediate ( cox2 , cox3 , cytb , nadh2 , nadh4l , nadh5 , nadh6 , atp6 and atp8 ) and low ( cox1 ) degrees of genetic differentiation ( fig . 4 ) . in agreement with our results it has been shown that the cox1 gene is one of the most conserved protein - coding genes in the mitochondrial genome of all metazoans [ 48 ] , [ 49 ] , while the most variable are the subunits of the nadh oxidoreductase and atp synthase complexes [ 50 ] – [ 52 ] . the relatively low divergence in the mitochondrial genes that belong to the cytochrome c oxidase , the complex that catalyze the transfer of electrons from cytochrome c to oxygen , is most probably explained because this complex is the rate limiting step of the electron transport chain [ 53 ] .
next , to understand whether nematodes were permanently fixed in shells when killed , i examined collections of c . nemoralis housed in liverpool and manchester museums , a large part of which contains arthur cain’s u . k . wide survey of c . nemoralis from 1950 22 . from 1 , 406 c . nemoralis shells , specimens collected in 1960 , 1918 , 1909 , 1866 and even 1864 had nematodes encased in their shells ( figure s2 , table s2 ) . also sub - fossil snail shells of c . aspersum , c . nemoralis and c . hortensis , estimated to be > 500 years old 23 , were also examined and nematodes were found to be present , albeit in low amounts ( n = 76 shells , 1 c . nemoralis shell had 2 nematodes present , figure s3 ) . therefore , when nematodes are encapsulated they are fixed for hundreds of years as a permanent record of parasitism , which permits an evolutionary analysis of shells across the stylommatophora .
given that the mitochondrial genome of pulmonate snails and slugs contain very little intergenic non - coding sequences and / or gene overlaps [ 11 ] , [ 12 ] , gene rearrangements are very rare considering that they would most likely disrupt some of the genes involved , rendering them non - functional . as a result of this , gene order in these species is well conserved [ 5 ] . it has been described that trna genes are more prone to switch their position than larger protein - coding and rrna genes [ 5 ] , [ 11 ] , [ 16 ] . this pattern is observed in our study where c . aspersum has identical gene order and content to c . nemoralis , but both have small differences in comparison to the other stylommatophorans . these differences are explained by the inversions of some trna genes , and the nadh4 gene ( fig . 3 ) . most of these inversions are maintained within eupulmonata , but comparisons with basommatophora revealed additional rearrangements that involve two additional trnas , and protein - coding genes ( cox2 and nadh4l ) [ 4 ] , [ 16 ] .
fragments of the whole mitochondrial dna sequence were analyzed in macclade 4 . 08 [ 30 ] and mega v . 5 . 1 [ 31 ] . to control for sequencing errors , each partial sequence was evaluated at least twice . ambiguous base pairs were resolved manually according to roche’s flowcharts and the corresponding quality score values for each base in the reads . annotations and editing procedures of the mitochondrial genomes of c . aspersum were done in geneious v 4 . 8 . 5 [ 32 ] . mitochondrial genes were identified by sequence comparison using dogma [ 33 ] and blast searches at ncbi ( with the blastn and blastx algorithms [ 34 ] ) against other eupulmonata sequences ( table 1 ) . the limits of both protein coding and ribosomal rna genes were adjusted manually based on location of adjacent genes , and the presence of start and stop codons . transfer rna genes were located using arwen v . 1 . 2 [ 35 ] , dogma [ 33 ] and trnascan - se v . 1 . 21 [ 36 ] , by means of the generalized invertebrate mitochondrial trna settings , after which they were manually adjusted based on specific anticodons in regions between identified genes .
born , i . 1778 . index rerum naturalium musei cæsarei vindobonensis . pars i . ma . testacea . verzeichniß der natürlichen seltenheiten des k . k . naturalien cabinets zu wien . erster theil . schalthiere . - pp . [ 1 - 40 ] , 1 - 458 , [ 1 - 82 ] . vindobonæ . ( kraus ) .
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brieva , a . ; philips , n . ; tejedor , r . ; guerrero , a . ; pivel , j . p . ; alonso - lebrero , j . l . ; gonzalez , s .
a screen for natural products bearing pharmacological properties has yielded a secretion of the mollusk cryptomphalus aspersa ( sca ) , which possesses skin - regenerative properties . in this report , we outline some of the cellular and molecular effects underlying this observation . first , we found that sca contained antioxidant sod and gst activities . in addition , sca stimulated fibroblast proliferation and rearrangement of the actin cytoskeleton . additional mechanisms involved in the regenerative effect of sca included the stimulation of extracellular matrix assembly and the regulation of metalloproteinase activities . together , these effects provide an array of molecular mechanisms underlying sca - induced cellular regeneration and postulate its use in regeneration of wounded tissue . copyright © 2007 s . karger ag .