buthus occitanus , the common yellow scorpion , is a species of scorpion in the buthidae family .

it occurs in the middle east , north africa and europe .

b. occitanus is 60-80 mm in length , has a yellow or yellow-brown color and is venomous producing botit6 toxin , but its toxicity varies markedly across its range .

this scorpion is often found in dry and hot areas with sparse vegetation , where it hides under stones etc. during daytime .

it has also been reported from mediterranean forests in spain at altitudes above 1000 meters ( with snowfall in the winter ) .

the sting in europe is painful but with only mild toxic effects , however , in africa it can be fatal .

buthus occitanus

maggie whitson marked
buthus occitanus
as hidden on the
buthus occitanus
page . reasons to hide : duplicate
maggie whitson set
image of buthus occitanus
as an exemplar on
buthus leach 1815
.
a first exploration of the venom of the buthus occitanus scorpion found in southern france .
kari pihlaviita added the finnish common name
kenttäskorpioni
to
buthus occitanus
.
a first exploration of the venom of the buthus occitanus scorpion found in southern france . - pubmed - ncbi
vargas o , martin mf , rochat h . characterization of six toxins from the venom of the moroccan scorpion buthus occitanus mardochei .
martin mf , rochat h . purification of thirteen toxins active on mice from the venom of the north african scorpion buthus occitanus tunetanus .
gantenbein b , largiadèr cr . the phylogeographic importance of the strait of gibraltar as a gene flow barrier in terrestrial arthropods : a case study with the scorpion buthus occitanus as model organism .
sampieri f , habersetzer - rochat c , martin mf , kopeyan c , rochat h . amino acid sequence of toxin xi of the scorpion buthus occitanus tunetanus . evidence of a mutation having an important effect upon neurotoxic activity .
pictures of the specimens of buthus occitanus studied . a ) animal collected near orange city ( vaucluse county ; photo credit : s . diochot ) and b ) in massif des maures ( var county ; photo credit thierry : hupin ) .
gonzales 1980 : 100 scorpion stings . identification : b . occitanus was identified in 50 % of cases ; criteria not given .
goudet c , chi cw , tytgat j . an overview of toxins and genes from the venom of the asian scorpion buthus martensi karsch .
casal and luque 1985 : 400 scorpion stings . identification : the retrospectively analysed cases were attributed to b . occitanus ; criteria not given .
bouhaouala - zahar b , ducancel f , zenouaki i , ben khalifa r , borchani l , pelhate m , boulain jc , el ayeb m , ménez a , karoui h . a recombinant insect - specific alpha - toxin of buthus occitanus tunetanus scorpion confers protection against homologous mammal toxins .
volkova tm , garsia af , telezhinskaia in , potapenko na , grishin ev . neurotoxins from the venom of the central asian scorpion buthus eupeus .
lourenço wr . compléments à la faune de scorpions ( arachnida ) de l’afrique du nord , avec des considérations sur le genre buthus leach , 1815 .
abroug et al 1999 : 825 patients with scorpion stings . identification : positive history of scorpion sting , with the scorpion being seen or captured with a . australis and b . occitanus being the two most common species in the area .
in addition to long - chain peptides such as α - toxins which constitute about 10 % of the detected masses , we found that 19 % of the masses are situated around 2000 da , 57 % over 3000 da and only 14 % over 4000 da . it is worth mentioning that a peptide with a molecular weight of 3178 da gave a high intensity peak in ms . a toxin with an identical mass , p01 , is thought to be a ligand for the small conductance ca 2 + activated k + channels , and was first purified from the venom of the moroccan scorpion androctonus mauretanicus . a similar peptide was also purified from the venoms of two other north african scorpion species , androctonus australis from algeria and tunisia , and buthus occitanus tunetanus from egypt ( zerrouk et al . 1996 ) . unfortunately , no specific antibodies for this family ( α - ktx8 ) are available to verify the identity of this toxin on our buthus occitanus venoms .
in addition to α - like toxins , two structural homologous to the classic α - toxin aah ii from androctonus australis hector were identified previously in the north african buthus occitanus tunetanus scorpion venom ( martin and rochat 1984 ; sampieri et al . 1987 ) . aah ii is the reference peptide for the structural and immunological group ii and its homologues can be detected using its highly specific anti - serum . aah ii itself has been the target of intensive immunological and pharmacological studies because this toxin constitutes 40 % to 80 % of the total lethality of the androctonus australis venom towards mammals . neutralizing aah ii is crucial to improve serotherapy against androctonus stings in north africa . after elution of our buthus occitanus venoms , a substantial positive reaction was obtained against the anti - aah ii serum with peptidic fraction having a mass of 7304 da . a more precise elisa answer was obtained with the uhplc fraction in which a peptide with a mass of 7465 da was found .
ghalim et al . 2000 : 275 patients with scorpion stings . identification : by the patient or the physician . prevalent in the area : buthus occitanus and androctonus mauretanicus mauretanicus ; venom serum levels were quantified using an elisa . severity scale ( krifi et al . 1998 ) : grade i ( only local symptoms , local pain and a burning sensation ) ( 247 / 275 ) ; grade ii ( local and systemic symptoms ) ( 28 / 275 ) ; grade iii ( local and systemic symptoms with cardiovascular shock , respiratory failure , acute pulmoary oedema , priapism , convulsions ) ( 0 / 275 ) .
abroug et al . 1999 : prospective randomized controlled trial in tunesia to assess the routine administration of 20 ml bivalent ( a . australis , b . occitanus ) f ( ab ' ) 2 scorpion antivenom ( institut pasteur , tunis , tunesia ) , i . v . , versus placebo irrespective of clinical severity . n = 825 ( , patients > 10 years . grading of severity : absence ( grade i ) or presence ( grade ii ) of systemic envenoming . diagnosis of scorpion envenoming based on a positive history of scorpion sting , with the scorpion being seen or captured . a . australis and b . occitanus are the two most common species in the area .
masses found in the sample 2 buthus occinatus venom collected near orange , which are also included at least twice in all the other venoms studied ( a . u . arbitrary unit , m / z , mass - to - charge ratio ; sn , signal - to - noise ratio , i . c . v . intracerebroventricular injection , n . d . not detremined ) .
ghalim et al . 2000 : prospective study of patients with scorpion stings . n = 275 . identification : by the patient or the physician . prevalent in the area : buthus occitanus and androctonus mauretanicus mauretanicus . venom serum levels were quantified using an elisa . severity scale ( krifi et al . 1998 ) : grade i ( only local symptoms , local pain and a burning sensation ) ( 247 / 275 ) ; grade ii ( local and systemic symptoms ) ( 28 / 275 ) ; grade iii ( local and systemic symptoms with cardiovascular shock , respiratory failure , acute pulmoary oedema , priapism , convulsions ) ( 0 / 275 ) . 179 / 275 were treated with equine f ( ab ' ) 2 scorpion antivenom ( androctonus mauretanicus mauretanicus ) ( 27 % 2 - 5 ml ; 73 % 10ml ) . antivenom application 77 . 6 % i . m . , 6 . 2 % s . c . , 16 . 2 % both .
representative chromatograms of collected buthus occitanus venoms . a ) venom sample 1 ( 100 ng ) and b ) venom sample 2 ( 100 ng ) obtained under electric stimulation of the post - abdomen of scorpions collected near orange city ( vaucluse county ) . the chromatogram of the venom sample 3 is very similar to that of sample 2 and it is not shown . c ) venom sample ( 100 ng ) obtained by manual stimulation from a single animal collected in massif des maures ( var county ) . inset : relative proportion of the mass - classes detected . solvents were water plus 0 . 05 % tfa ( a ) and 80 / 20 ( v / v ) acetonitrile / water plus 0 . 04 % tfa ( b ) . the experimental conditions used were : elution gradient 4–50 % of b ; dionex pepswift monolithic column , 200 μm i . d . × 250 mm equipped with a trap column 200 μm i . d . × 5 mm ; flow rate 1 . 5 μl / min ; uv recording at the wavelength of 214 nm .
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the following article is based on an article i wrote for the journal published by the exotiske insekter society in scandinavia ( december issue ( no . 29 ) , 2001 , in danish ) . no reproduction of this article is permitted ( electronic or printed ) without my permit . this article is continuously updated . last updated : 06 . 07 . 18 [ a major update is planned in 2018 ]
a review of the scorpion fauna of europe . jan ove rein medical library & information center norwegian university of science and technology
europe hosts a surprisingly high diversity of scorpions . at the moment , 76 valid species ( one is probably extinct ) [ per 06 . 07 . 18 ] are reported from europe ( the species known from the far eastern part of turkey are excluded ) . scorpions are reported from france , monaco , spain , portugal , italy , san marino , malta , switzerland , austria , cyprus , greece , turkey , russia ( north caucasus ) , ukraine ( crimea only ) , and the balkan countries . in addition , an introduced colony of scorpions in southern england has been known since the 18th . century . scorpion findings have also been reported from germany , holland , norway , denmark and sweden , but no data indicate that these countries host permanent populations ( as england does ) . these scorpions have probably been accidental stowaways .
this family has three representatives in europe , which do not overlap in distribution . they are usually found in hot and dry wasteland with sparse vegetation , where they hide in natural burrows under stones etc . during daytime . as in other buthids , the venom is quite strong and sting is very painful . i have not been able to find any reports of deaths or serious cases attributed to european scorpions in the literature , but see the comments in connection with iurus dufoureius and calchas nordmanni . the european species seem not to pose any great threat to healthy humans .
this new species has so far been collected only in three localities in southern portugal and two locations in southern and easter spain . it is possible that this species is also distributed outside these areas .
described from cyrpus in 2011 . medium sized buthid that can reach 70 mm . color is pale - yellow to yellow with brownish spots on the keels of the trunk . very similar to
, which is the other endemic species on the island . the two species doesn ' t seem to overlap , but very little is known about the new species distribution on the island except for the reported findings in northern cyprus .
is probably distributed all over the island , but this have to be verified .
described from sicily , italy ( and the first buthid from italy ) . the species is probably extinct due to urbanization . more information later .
july 2018 : a new species from the lassithi plateau , crete ( greece ) . more details later .
in addition to these three buthids , michalis & dolkeras ( 1989 ) reported a finding of androctonus bicolor from greece . this is not correct , and the scorpion in question was a m . gibbosus ( fet & braunwalder , 2000 ) . a introduced colony of the south american scorpion centruroides gracilis has been reported from the canary islands ( these islands are not a part of the european fauna element , but the islands are a part of spain , and i therefore choose to mention this ) . armas & báez ( 1988 ) reported about an established colony in la laguna , tenerife island , but i do not know if this scorpion has spread to the other islands in the area .
this family is represented with one genus in europe , belisarius . this genus , with one species , was previsouly included in the small family troglotayosicidae . this family was abolished in 2003 , and belisarius was tranfered to chactidae .
june 2015 : new species from sierra de las nieves region , in southern spain . more details later .
this strange scorpion lacks eyes and pigmentation , which is typical for many cave - living animals . this species is 3 - 4 cm long , and the trunk has a pale yellow color ( almost translucent ) pedipalps are dark brown , and metasoma ( tail ) is also somewhat darker than the trunk . this is the only troglophile (
cave loving
) species in europe . b . xambeui is the rarest scorpion in europe , and is only reported from the southeastern pyrenees in france and in cataluña in spain . it lives in quite high altitudes ( 650 - 1500 m ) . even though it has cave - dwelling adaptations , it is seldom found deep into caves , but rather occupies cave entrances . b . xambeui can also be found under stones etc . , and in connection with abandoned man - made structures like collapsed chimneys , cellars and castle ruins .
euscorpius is the only genus of this family in europe . members of this genus are known from several countries , and are quite common in some places ( in addition to the european distribution , some species have been reported from north africa , georgia and the asian part of turkey , iraq , yemen ( introduced ) and uruguay ( introduced ) ) . because of the synanthropic nature of some of the species in this genus , euscorpius specimens are reported as stowaways from time to time in many countries outside their natural distribution ( e . g . germany , holland , denmark , sweden and norway ) . most species of euscorpius are similar in appearance , and can be difficult to identify without checking trichobothrial structures on the pedipalps . some of the species also display intraspecific variations in colors and size , which also make identification more difficult . an online identification key is available in the scorpion files .
the venom of all species of euscorpius is harmless . these species will rarely sting , and the smaller species have difficulties with penetrating human skin . some sources say that the sting of e . italicus might be very painful ( i . stathi , pers . comm . ) .
november 2016 : a new species from aladaglar mountains , in nigde , kayseri , and adana provinces in southern turkey . more details later .
september 2016 : a new species from taurus mountains in antalya province ( alanya district ) in southern turkey . more details later .
in southern switzerland , northern italy and southern austria revealed the presence of two different forms , separated by the river adige ( etsch ) in the northern italy . the genetic difference between the two forms was large enough to justify an elevation of the western form to a new species ,
by using morphological characteristics , but collection site will tell which species you have .
is a small , black scorpion , which rarely reach more than 30 mm in length . it is usually found in mountain areas with a reasonable high humidity , often under stones , logs etc .
is so far reported from north west italy ( piemonte , valle d ' aosta and lombardy ) and switzerland .
2017 : new species from northeastern sicily and southern calabria , italy . more details later .
january 2017 : new species from amorgos island , greece . more details later .
have now been raised to species status . the new species is restricted to northern and central italy , san marino , vatican city state , western slovenia , northwestern croatia . more details later .
species complex from dilek penninsula ( aydin province ) , westerm turkey . more details later .
this species belongs to the
e . carpathicus species complex
, and was previously recognized as a subspecies of e . carpathicus . e . balearicus is distributed on the balearic islands ( mallorca , menorca , cabrera , ibiza & formentera ) in the mediterranean sea , where it can be found under stones etc . e . balearicus is small to medium sized ( 30 - 37 mm ) . coloration is light brownish ( actually some orange overtones in some specimens ) with little contrasting patterns . metasoma reduced proportionally , and unusual large pedipalps ( compared to other populations of e . carpathicus ) . as a rule of thumb : all euscorpius from the balearic islands are e . balearicus .
this species was recently described from a mountain area ( 1400 - 2400 meters ) in northern albania . some specimens have been collected as high as 2569 meters , something which is very impressive for european scorpions . this is a small species ( appr . 30 mm ) , with a light brownish trunk and brown legs and pedipalps .
species complex
, which is so far poorly investigated . due to this , there is some uncertainty regarding its species status , and more research are needed on
april 2014 : a cave species endemic to the euboea island , greece . more details later .
2017 : new species from calbria and basilicata ( southern part ) , italy . this population was previously synonymized with
september 2013 : the scorpion population in crete ( greece ) is finally elevated to species level from subspecies status . more details later .
2017 : new species from sardinia , italy . this population was previously synonymized with
july 2012 : new species from celano , abruzzo in central italy . more details later .
july 2015 : new species from turkey ( konya and mersin provinces ( eastern part of the central taurus mts . ) ) . more details later .
january 2017 : new species from ios and sikinos islands , greece . more details later .
december 2014 : new species from stara planina mts . , bulgaria , and adjacent areas of bulgaria and serbia . more details later . see also
june 2015 : new species from the western rhodope mts . in southern bulgaria . more details later .
november 2013 : new species from western bosnia & herzegovina , southern croatia and northwestern montenegro . more details later .
this species was described in 2000 , and belongs to the
e . mingrelicus species complex
. e . gamma is reported from austria , italy , slovenia and croatia , and is a dark colored scorpion that reach 32 mm in length . it is usually found in humid mountain areas , where it can be collected under stones , logs and under the bark of old trees . it is very difficult to separate this species from e . mingrelicus and e . germanus .
2017 : new species from puglia , molise , campania and basilicata , italy and pelagosa island , croatia . this population was previously synonymized with
april 2015 : new species from central - western greece . more details later .
april 2014 : new species from antalya province , southern turkey . more details later .
november 2016 : a new species from mount eseler , in denizli province in southwestern turkey . more details later .
april 2016 : new species from mount honaz in southwestern turkey . more details later .
december 2017 : a new species from from balikesir and çanakkale provinces in northwestern turkey . more details later .
october 2014 : species from mt . olympus at the eastern border between thessaly and central macedonia , greece . more details later .
this rare scorpion is only know from mt . athos ( chalkidiki ) in greece . it is a medium to large sized species , which is medium to dark brown in color . the posteriour part of the pody ( including the tail ) and the legs are lighter in color . the metasoma is quite slender with all segments longer than wide . the exceptionally slender and smooth metasoma are key diagnostic characters of this species to be used in separating it from the two related species , e . carpathicus candiota and e . tergestinus , which both share the similar pedipalp patellar external trichobothria series with this species ( eb = 4 , eba = 4 , em = 4 ) . see my online euscorpius identification guide for more information . i have no data on the habitat of this species , but i know that other euscorpius species in greece are found in houses and other human buildings .
november 2013 : new species from southwestern turkey . more details later , but see my blog post
like e . carpathicus , e . mingrelicus is a species with a large intraspecific variation ( 10 subspecies are described ) . this species is probably a
species complex
, and future research might show that this species actually are several closely related species ( e . gamma was separated from this
species complex
) . e . mingrelicus has an eastern distribution , and is reported from bosnia and herzegovina , bulgaria , croatia , georgia , greece , italy , romania , russia , slovenia , turkey and yugoslavia . little information is available on the biology of this species , but it seems that it thrives in high altitude areas with some humidity ( both in mountain forests and in higher areas with sparse vegetation ) . e . mingrelicus is also reported from river valleys in austria . it hides under stones and other suitable objects on the ground , but is also found under the bark of dead trees and logs . it is very difficult to separate this species from e . germanus and e . gamma . e . mingrelicus is also dark ( blackish ) like the other two , but is somewhat larger ( adults reach in average 38 mm in length ) .
april 2014 : new species endemic to the euboea island , greece . more details later .
this species was recently ( 2006 / 2007 ) described from the montecristo island ( tuscan archipelago , italy ) , where it probably is endemic . the island population was orginally described as
( c . l . koch , 1837 ) . a recent study has redescribed this taxa and based on morphological analysis concluded that the monecristo population is a valid species .
species complex ) , with total lenghts up to 43 mm . overall body is light in color with redish pedipalps claws ( fingers ) and a darker colored fifth tail segment .
is a hygrophilic ( humid loving ) species , that is only found in humid areas on the island ( around the only house on the island under stones of small walls , but also under bark of of trees and inside a rotten log .
has a risk of extinction because of its very limited distribution in a small area . it is very vulnerable to climatic changes and loss of humid habitats . the island fauna is protected and it is not possible to visit montecristo without permission .
july 2015 : new species from turkey ( northwest : ankara , bolu , düzce , karabük , and sakarya provinces ) . more details later .
september 2015 : new species from bulgaria ( southwest ) and greece ( northeast , near southwestern bulgarian border ) . more details later .
2017 : new species from southern puglia ( salento ) , italy . more details later .
december 2014 : new species from southwestern bulgaria and northeastern greece . more details later . see also
october 2014 : species from epirus in the northwestern greece . more details later .
this species , which previously belonged to the
e . carpathicus species complex
, was elevated to species status in october 2003 . this is a rare species , that previously thought to be endemic to ukraina ( crimea , southern coast ) . in 2017 , an investigation revealed a much wider distribution . this species is now belived to be found in cyclades islands ( greece ) and northwestern anatolia ( turkey ) , in addition to ukraine . the pedipalp patellar external trichobothria series eb and series eba have have 4 trichobothria ( this characteristic is shared with e . balearicus , e . koschewnikowi and e . tergestinus . so far , i do not have any information on how these species can be separated , except for using collection site . limited information about e . tauricus ' s habitat are available , but they have been collected under stones and under forest litter in warmer areas .
july 2015 : new species from turkey ( northwest : bursa province ) . more details later .
october 2014 : species from karpathos and kasos islands ( eastern aegean sea ) , greece . more details later .
december 2014 : species from epirus in the northwestern greece . more details later .
two rare representatives of this family are found in europe [ this information is outdated and will be updated soon . also , see below for info on taxa ] . these species are not mentioned in medical literature , but there exists an unpublished report from the greek island kos ( eastern aegean sea ) that a man ( not allergic to scorpion sting ) almost died from a sting from iurus dufoureius ( i . stathi , pers . comm . ) . the symptoms seemed to be systemic . peoples also report about of severe symptoms from calchas nordmanni stings from another greek island ( megisti ) . it is very difficult to conclude anything from these unverified reports about the potential danger of mediterranean iuridae . these scorpions are very rare , and are seldom in contact with peoples . this might explained the lack of cases in the medical literature ( peoples are rarly stung by these scorpions ) . it is also important to note that no other iuridae worldwide are reported to have medical significance . until further research is done , the european iuridae should be treated with respect . peoples who were stung were mostly construction workers who worked in the foundations of old stony buildings ( i . stathi , pers . comm . ) .
these are a small scorpions that reaches 45 mm in length . they are light brown in color . previously , c . nordmanni was the only known species , but in a review article of the genus fet , soleglad & kovarik ( 2009 ) conclude that three distinct , disjunct species exist rather than one widespread species ( c . nordmanni ) as previously thought . the genus is known from from greece ( the islands of megisti and samos ) , parts of turkey and northen iraq . a potential sighting in syria has not been verified . more information is needed on the biology of this genus . as far as i can tell , the species prefer humid habitats in forests , where it is located under stones and other suitable objects . more information will be added later .
in this article i have excluded the species that are reported from the asian ( eastern ) part of turkey . so far , the following species have been reported from eastern part of turkey :
it is possible that other species occur in these areas , as they are not very well investigated .
i ' m very interested in getting european scorpions for my collection ( i ' m interested in purchasing or exchanges ) . it is important for me to know where the scorpions were collected , as some of them might be used for research . please contact me if you are able to collect scorpions for me :
i ' m very grateful to professor victor fet and dr . rolando teruel for reviewing the first edition of this article ( professor fet has also reviewed the seccond edition ) , and to professor victor fet and drs . benjamin gantenbein and valerio vignoli for keeping me updated on the research on euscorpius taxonomy and phylogeny ! dr . iasmi stathi has supplied me with a lot of unpublished information , and also made comments to the manuscript . i ' m very grateful for this ! a special thanks to dave gaban for checking the language and spellings in the article .
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urltoken
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martin - eauclaire mf 1 , bosmans f 2 , céard b 1 , diochot s 3 , bougis pe 4 .
aix marseille université , cnrs , crn2m umr7286 , faculté de médecine , campus nord , cs80011 , bd pierre dramard , f - 13344 marseille cedex 15 , 13015 marseille , france .
department of physiology , johns hopkins university - school of medicine , baltimore , md 21205 , usa ; solomon h . snyder department of neuroscience , johns hopkins university - school of medicine , baltimore , md 21205 , usa .
cnrs , ipmc umr 6097 , 06560 , sophia - antipolis , valbonne , france .
aix marseille université , cnrs , crn2m umr7286 , faculté de médecine , campus nord , cs80011 , bd pierre dramard , f - 13344 marseille cedex 15 , 13015 marseille , france . electronic address : pierre - edouard . bougis @ univ - amu . fr .
pmid : 24418174 pmcid : pmc3952629 doi : 10 . 1016 / j . toxicon . 2014 . 01 . 002
electrophysiological characterization . a ) rna v 1 . 2a current at 0mv before ( black ) and after ( red ) addition of the venom fraction ( 0 . 5 μm ) to the recording chamber . oocyte was held at −90mv and depolarized to test pulse voltage for 50ms . current amplitude is shown on vertical axis , time scale is shown on horizontal axis . b ) normalized steady - state inactivation ( i / i max : open circles ) and conductance ( g ) - voltage ( v ) curves ( filled circles ) before ( black ) and after ( red ) venom fraction addition . g - v curves were deduced from current - voltage relationships obtained by depolarizing rna v 1 . 2a - expressing oocytes in 5mv steps from a holding potential of −90mv over a wide voltage range . the obtained fit values using the boltzmann function for v 1 / 2 are −41 ± 1mv ( control ) and −47 ± 1mv ( venom fraction ) . slope coefficients are 6 . 3 ± 0 . 6 and 5 . 2 ± 0 . 6 , respectively . steady - state inactivation or channel availability curves were obtained by a standard double pulse protocol while holding the oocyte at −100mv . the obtained fit values for v 1 / 2 are −50 ± 1mv ( control ) and −51 ± 1mv ( venom fraction ) . slope coefficients are 6 . 0 ± 0 . 2 and 7 . 8 ± 0 . 6 , respectively . n = 3 for all experiments and data are shown as mean ± sem .
marie - france martin - eauclaire , a frank bosmans , b brigitte céard , a sylvie diochot , c and pierre e . bougis a , *
b department of physiology and solomon h . snyder department of neuroscience , johns hopkins university–school of medicine , baltimore , md 21205 , usa
* corresponding author : pierre e bougis , tel : + 33 ( 0 ) 4 9169 8850 , rf . uma - vinu @ siguob . drauode - erreip , crn2m , umr 7286 , faculté de médecine campus nord , cs80011 , bd pierre dramard , f - 13344 marseille cedex 15 , france
) , two places distant from each other by 200 km ( 125 mi ) . the venoms were individually obtained , by manual stimulations ( one time ) of the post - abdomen of the specimen collected in massif des maures and by electric stimulations ( three times ) for specimens collected near orange city . animals were thereafter released in their natural habitats .
goat - anti - rabbit fab and bovine serum albumin ( bsa ) were from sigma . uv - grade acetonitrile was from carlo - erba and trifluoroacetic acid ( tfa ) was from sigma - aldrich . α - cyano - 4 - hydroxycinnamic acid and sinnapinic acid were from bruker . all other reagents and solvents of analytical grade were obtained from commercial sources , and used without further purification unless otherwise stated .
the lyophilized crude venoms ( 50 to 100 μg for venoms manually obtained and 300 to 2500 μg for venoms obtained by electric stimulations ) were first dissolved in solvent a ( tfa 0 . 1 % in water ) in order to obtain a stock solution of 10 μg / ml . this step was followed by a centrifugation at 13 000 g for 20 min to precipitate the non - soluble high molecular weight proteins , which are not toxic . the supernatant ( 100 ng of venom ) was separated by nano uhplc analyses using a dionex ultimate® 3000 rslc nano system ( thermo ) controlled by chromeleon ® . elution profile was monitored by absorbance readings at 214 nm . hplc grade elution solvents were water plus 0 . 05 % tfa ( a ) and 80 / 20 % acetonitrile / water plus 0 . 04 % tfa ( b ) . pre - column ( 200 μm × 5 mm ) and column ( 100 μm × 250 mm ) were pepswift monolithic nano ps - dvb from dionex . running flow rate was 1 . 5 μl / min . the column pressure was 400 bars and its temperature kept at 60°c .
conventional hplc was conducted as follows : the most abundant venom ( 2 . 5 mg , coming from a scorpion collected near orange city , sample 2 ) was first dissolved in cold milliq - water ( 250 μl ) , and centrifuged at 13 000 g for 20 min at 4°c . part of this material ( 0 . 5 mg ) was loaded onto a reverse - phase hplc column ( 100 rp - 18 , 4×250 mm , lichrospher , 5μm , merck , germany ) using a dionex ultimate 3000 hplc basic systems ( thermo ) . hplc grade elution solvents were water plus 0 . 1 % tfa ( a ) and acetonitrile plus 0 . 1 % tfa ( b ) . the elution ( 0 . 5–1 ml / min ) was monitored by absorbance at both 230 nm and 280 nm , simultaneously .
the nano uhplc effluent was automatically and directly dropped every 20 second ( i . e . 0 . 5 μl by well ) from the tip of the column capillary in a falcon microtest ™ 96 - well elisa plate , each well already containing 50 μl of 50 mm carbonate / bicarbonate buffer , ph 9 . 6 . the plate was shaken overnight at 4°c to achieve the antigen coating process . then , the plate was saturated for 1h with a blocking solution ( 100 μl by well of 5 % dried milk in pbs , 0 . 05 % tween 20 , ph 7 . 4 ) . after an extensive washing ( 3 times 300 μl by well of 0 . 1 % dried milk in pbs , 0 . 05 % tween 20 , ph 7 . 4 ) , the binding of the toxin - specific serum ( 100 μl by well of a 1 / 10 000 dilution ) was allowed for 90 min at room temperature . after washing ( same as above ) , an alkaline phosphatase - coupled goat anti - rabbit fab ( 100 μl by well of a 1 / 40 000 dilution ) was added . after 90 min at room temperature and washing ( same as above ) , the 4 - nitrophenyl phosphate substrate from merck was added ( 300 μl by well of a 3 mg / ml solution in 10 % diethanolamine buffer , 0 . 5 mm mgcl 2 , ph 9 . 8 ) . after the colour had developed ( between 15 and 60 min ) , the absorbance was measured at the wavelength 405 nm using a labsystems iems reader mf .
lethality tests were performed in male swiss mice ( n = 48 ) weighting 18 ± 3 g by subcutaneous ( s . c . ) or intracerebroventricular ( i . c . v . ) injections as described ( gordon et al . 1996 ; romi et al . 1993 ) . experiments were carried out in accordance with the european communities council directive 2010 / 63 / eu .
the four collected venoms were individually submitted to a reversed - phase nano uhplc step ( 100 ng each ) in order to separate their components . base on their chromatographic patterns , among the three venoms collected near orange city one ( sample 1 ,
) was rather different from the two others . those were much more similar among themselves ( sample 2 ,
and it shows noticeable differences with the venoms from orange origin . chiefly , the venom complexity is much higher and the relative intensities among peaks equally distributed over the whole chromatogram .
venom collected near orange , which are also included at least twice in all the other venoms studied , as well as elisa cross - reactivity and toxicity in mice .
1 . x potassium channel family , was detected at 18 . 82 min using our elisa test . moreover , a mass of 4149 da is in accordance with that of ktx ( 4150 da ) purified from
venom . similar to this venom , ktx was also found in its deoxidised form ( 4182 da , 20 . 15 min ) (
currents in cerebellum granular cells or cultured striatum neurons . the distribution of binding sites in rat brain and its electrophysiological characteristics suggest that bmtx3 specifically binds to the k
) . a strong positive response was observed from 25 . 48 to 27 . 48 min (
) , which resulted in the mass determination of 3777 da , displaying a very high ms intensity . this mass likely corresponds to a bmtx3 analogue in its z - form ,
we evaluated overall venom toxicity by s . c . injection in mice . to this end , we used the most abundant venom sample available ( 2 . 5 mg , sample 2 collected near orange city ) . similar to the north african
mouse was found , suggesting that the tested venom is indeed lethal for small mammals . it is worth mentioning that the
venom belongs to only one specimen and previous reports demonstrate that large differences in toxicity and composition can arise between individual venoms . due to the low amount of material resulting from uhplc fractionation , it was experimentally impossible to obtain mouse toxicity data of the individual venom components (
0 . 5 μl by well collected from the tip of the column capillary ) . in order to further the biological characterization of individual fractions , conventional hplc was used to fractionate sample 2 ( 500 μg ) . fractions of interest were first analysed using elisa and mass spectrometry in order to verify that masses similar to those detected using nano uhplc were injected into mice . several fractions tested by i . c . v . injection induced moderate convulsions or other toxic signs ( paralysis , twitching , sweating , tachycardia ) in mice lasting 10 min to 1 h (
) . only two fractions , immunoreactive to the anti - bot i serum , were lethal . the most lethal fraction promoting death in less than 45 min and having a mass of 7465 da was also highly positive for the anti - aah ii serum . other fractions giving a strong positive reaction against the anti - bot i serum were able to induce toxicity in mice with envenomation signs corresponding to those of α - like toxins .
oocytes using the two - electrode voltage clamp technique . as is typically seen with α - toxins , a concentration of 0 . 5 μm inhibits fast inactivation of the channel resulting in a persistent current at the end of a 50 ms test pulse (
we used a nano uhplc / maldi - tof - ms automated workflow , toxin specific elisa and electrophysiology .
we would like to thank al goldin ( ucirvine ) for sharing the rna v 1 . 2a clone . we acknowledge claire chaussignand , thierry hupin , marc diochot for collecting and milking scorpions in their natural habitat before being released , and maya belghazi for his valuable experimental support . the centre national de la recherche scientifique ( cnrs ) , marseille - nice génopole and marseille protéomique ( map ) supported this work . parts of the research in this publication were supported by the national institute of neurological disorders and stroke ( ninds ) of the national institutes of health ( nih ) under award number r00ns073797 ( fb ) . the content is solely the responsibility of the authors and does not necessarily represent the official views of the nih .
the authors state that the works presented in this manuscript follow the rules of ethics and respects the duties of authors presented in the elsevier’s ethical guidelines for journal publication .
publisher ' s disclaimer : this is a pdf file of an unedited manuscript that has been accepted for publication . as a service to our customers we are providing this early version of the manuscript . the manuscript will undergo copyediting , typesetting , and review of the resulting proof before it is published in its final citable form . please note that during the production process errors may be discovered which could affect the content , and all legal disclaimers that apply to the journal pertain .
abbas n , rosso jp , céard b , belghazi m , lebrun r , bougis pe , martin - eauclaire mf . characterization of three “birtoxin - like” toxins from the androctonus amoreuxi scorpion venom .
amoreux pj . description méthodique d’une espèce de scorpion roux commune à souvignargues en languedoc et détails historiques à ce sujet .
batista cv , román - gonzález sa , salas - castillo sp , zamudio fz , gómez - lagunas f , possani ld . proteomic analysis of the venom from the scorpion tityus stigmurus : biochemical and physiological comparison with other tityus species .
benguedda ac , laraba - djébari f , ouahdi m , hellal h , griene l , guerenik m , laid y cnles , m . d . expérience de quinze années de lutte contre l’envenimation scorpionique en algérie . envenomation and its treatment in africa . conference proceedings : “immunotherapy in envenomation” dakar , senegal .
bosmans f , martin - eauclaire mf , swartz kj . deconstructing voltage sensor function and pharmacology in sodium channels .
caliskan f , garcía bi , coronas fi , batista cv , zamudio fz , possani ld . characterization of venom components from the scorpion androctonus crassicauda of turkey : peptides and genes .
campos fv , chanda b , beirao ps , bezanilla f . alpha - scorpion toxin impairs a conformational change that leads to fast inactivation of muscle sodium channels .
capes dl , goldschen - ohm mp , arcisio - miranda m , bezanilla f , chanda b . domain iv voltage - sensor movement is both sufficient and rate limiting for fast inactivation in sodium channels .
catterall wa . voltage - gated sodium channels at 60 : structure , function and pathophysiology .
cestèle s , qu y , rogers jc , rochat h , scheuer t , catterall wa . voltage sensor - trapping : enhanced activation of sodium channels by beta - scorpion toxin bound to the s3–s4 loop in domain ii .
cestèle s , yarov - yarovoy v , qu y , sampieri f , scheuer t , catterall wa . structure and function of the voltage sensor of sodium channels probed by a beta - scorpion toxin .
darbon h , zlotkin e , kopeyan c , van rietschoten j , rochat h . covalent structure of the insect toxin of the north african scorpion androctonus australis hector .
gordon d , ilan n , zilberberg n , gilles n , urbach d , cohen l , karbat i , froy o , gaathon a , kallen rg , benveniste m , gurevitz m . an ‘old world’ scorpion beta - toxin that recognizes both insect and mammalian sodium channels .
gordon d , karbat i , ilan n , cohen l , kahn r , gilles n , dong k , stühmer w , tytgat j , gurevitz m . the differential preference of scorpion alpha - toxins for insect or mammalian sodium channels : implications for improved insect control .
gordon d , martin - eauclaire mf , cestèle s , kopeyan c , carlier e , khalifa rb , pelhate m , rochat h . scorpion toxins affecting sodium current inactivation bind to distinct homologous receptor sites on rat brain and insect sodium channels .
granier c , novotny j , fontecilla - camps jc , fourquet p , el ayeb m , bahraoui e . the antigenic structure of a scorpion toxin .
liu zr , ye p , ji yh . exploring the obscure profiles of pharmacological binding sites on voltage - gated sodium channels by bmk neurotoxins .
loret ep , martin - eauclaire mf , mansuelle p , sampieri f , granier c , rochat h . an anti - insect toxin purified from the scorpion androctonus australis hector also acts on the alpha - and beta - sites of the mammalian sodium channel : sequence and circular dichroism study .
martin - eauclaire mf , bougis pe . potassium channels blockers from the venom of androctonus mauretanicus mauretanicus .
martin - eauclaire mf , couraud f . scorpion neurotoxins : effects and mechanisms . in : chang l , dyer rs , editors .
martin - eauclaire mf , granjeaud s , belghazi m , bougis pe . achieving automated scorpion venom mass fingerprinting ( vmf ) in the nanogram range .
oukkache n , rosso jp , alami m , ghalim n , saile r , hassar m , bougis pe , martin - eauclaire mf . new analysis of the toxic compounds from the androctonus mauretanicus mauretanicus scorpion venom .
pedraza escalona m , possani ld . scorpion beta - toxins and voltage - gated sodium channels : interactions and effects .
pimenta amc , stocklin r , favreau p , bougis pe , martin - eauclaire mf . moving pieces in a proteomic puzzle : mass fingerprinting of toxic fractions from the venom of tityus serrulatus ( scorpiones , buthidae )
possani ld , becerril b , delepierre m , tytgat j . scorpion toxins specific for na + - channels .
rogers jc , qu y , tanada tn , scheuer t , catterall wa . molecular determinants of high affinity binding of alpha - scorpion toxin and sea anemone toxin in the s3–s4 extracellular loop in domain iv of the na + channel alpha subunit .
romi r , crest m , gola m , sampieri f , jacquet g , zerrouk h , mansuelle p , sorokine o , van dorsselaer a , rochat h , martin - eauclaire m , van rietschoten j . synthesis and characterization of kaliotoxin . is the 26–32 sequence essential for potassium channel recognition ?
sheets mf , kyle jw , kallen rg , hanck da . the na channel voltage sensor associated with inactivation is localized to the external charged residues of domain iv , s4 .
vacher h , martin - eauclaire mf . antigenic polymorphism of the “short” scorpion toxins able to block k + channels .
valdez - velázquez ll , quintero - hernández v , romero - gutiérrez mt , coronas fi , possani ld . mass fingerprinting of the venom and transcriptome of venom gland of scorpion centruroides tecomanus .
zerrouk h , laraba - djebari f , fremont v , meki a , darbon h , mansuelle p , oughideni r , van rietschoten j , rochat h , martin - eauclaire mf . characterization of po1 , a new peptide ligand of the apamin - sensitive ca2 + activated k + channel .
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general : cardiotoxicity cardiotoxicity may be direct or indirect , but is a feature of severe envenoming , with cardiac arrhythmias , cardiac failure .
general : other hypovolaemic hypotension possible in severe cases due to fluid loss through vomiting and sweating .
key diagnostic features severe local pain , increased salivation , sweating , irritability , vomiting , cadrdirespiratory effects .
general approach to management all cases should be treated as urgent & potentially lethal . rapid assessment & commencement of treatment including appropriate antivenom ( if indicated & available ) is mandatory . however , most cases will not progress beyond local envenoming with severe pain . be aware young children at extra risk for severe envenoming .
antivenom therapy antivenom is controversial . in some regions it is the standard treatment , but in others is rarely used or avoided . studies have been conflicting on the effectiveness of antivenom for scorpion sting .
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: the principle aim of this site is to provide information useful to improving outcomes for humans suffering from envenoming or poisoning by animals , plants or mushrooms . we make a reasonable attempt to verify accuracy of information listed on this site . however , we cannot access every published paper of potential relevance , either because they are not available to us or are in a language we cannot translate internally . equally , we cannot list knowledge which is not yet reported or known . it should not be assumed that humankind currently knows all there is to know about any species , even for common species . further , we cannot control how users will interpret the information provided on this site . we therefore do not accept legal responsibility for use of the information provided and we require that all users use information from this site at their own risk . the following should also be noted when reading information contained within the databases on this website : italics for scientific nomenclature cannot be displayed , and superscripting and subscripting is absent in some instances .
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experimental and clinical observations suggest that the major systemic effects of envenoming are caused by endogenous catecholamines and acetylcholine , which are released in response to scorpion venom . as these are transmitters in the sympathetic , parasympathetic and somatic nervous systems , the resulting clinical symptoms of envenoming are dealt with in the section
neurological effects
.
however , scorpion venoms are also believed to lead to other indirect effects that are caused by the release of autopharmacologically active substances ( such as kinins , prostaglandins and slow - reacting substances ) . the pathophysiological effects of these substances overlap to a great extent . this makes it difficult to be certain about aetiology . in particular with regard to pulmonary oedema , there has been discussion concerning the effects of mediators on vascular permeability , which might constitute a non - cardiac component of the pulmonary oedema . peripheral blood pressure regulation is also responsive to a variety of different mediators that might be released .


biomphalaria havanensis , common name the ghost rams-horn , is a species of air-breathing freshwater snail , an aquatic pulmonate gastropod mollusk in the family planorbidae , the ram 's horn snails .

the shell of this species , like all planorbids is sinistral in coiling , but is carried upside down and thus appears to be dextral .

biomphalaria havanensis

the biomphalaria havanensis complex ( gastropoda : planorbidae ) in cuba : a morphological and genetic s . . .
. . . the type locality from which paraense and deslandes ( 1958 ) described specimens as b . havanensis is no longer accessible . their description differs from that of yong et al . ( 1997 ) . durand et al . ( 1998 recognized a biomphalaria sp . from several locations in cuba , that was morphologically distinct from b . havanensis of yong et al . ( 1997 ) and very similar to b . havanensis of paraense and deslandes ( 1958 ) . . . .
. . . their description differs from that of yong et al . ( 1997 ) . durand et al . ( 1998 recognized a biomphalaria sp . from several locations in cuba , that was morphologically distinct from b . havanensis of yong et al . ( 1997 ) and very similar to b . havanensis of paraense and deslandes ( 1958 ) . allozymes convincingly distinguished it from b . havanensis of yong et al . ( 1997 ) and from b . helophila . . . .
. . . the type locality from which paraense and deslandes ( 1958 ) described specimens as b . havanensis is no longer accessible . their description differs from that of yong et al . ( 1997 yong et al . ( , 2001 ) . durand et al . ( 1998 ) recognized a biomphalaria sp . from several locations in cuba , that was morphologically distinct from b . havanensis of yong et al . ( 1997 ) and very similar to b . havanensis of paraense and deslandes ( 1958 ) . . . .
. . . their description differs from that of yong et al . ( 1997 yong et al . ( , 2001 ) . durand et al . ( 1998 ) recognized a biomphalaria sp . from several locations in cuba , that was morphologically distinct from b . havanensis of yong et al . ( 1997 ) and very similar to b . havanensis of paraense and deslandes ( 1958 ) . allozymes convincingly distinguished it from b . havanensis of yong et al . ( 1997 ) and from b . helophila . . . .
benson , a . j . , 2018 , biomphalaria havanensis ( pfeiffer , 1839 ) : u . s . geological survey , nonindigenous aquatic species database , gainesville , fl , urltoken revision date : 11 / 1 / 2016 , access date : 7 / 9 / 2018
table 1 . states with nonindigenous occurrences , the earliest and latest observations in each state , and the tally and names of hucs with observations† . names and dates are hyperlinked to their relevant specimen records . the list of references for all nonindigenous occurrences of biomphalaria havanensis are found here .
. . . durand et al . ( 1998 ) recognized a biomphalaria sp . from several locations in cuba , that was morphologically distinct from b . havanensis of yong et al . ( 1997 ) and very similar to b . havanensis of paraense and deslandes ( 1958 ) . allozymes convincingly distinguished it from b . havanensis of yong et al . ( 1997 ) and from b . helophila . sequences from specimens from two of the locations ( hanabanilla lake , guatao ) reported by durand et al . ( 1998 ) were examined and found to be identical , so only the hanabanilla lake specimen was used in the analyses . . . .
fernandez ma , thiengo sc 2006 . susceptibility of biomphalaria amazonica and biomphalaria occidentalis from manso dam , mato grosso , brazil to infection with three strains of schistosoma mansoni . mem inst oswaldo cruz 101 ( suppl . i ) : 235 - 237 . [ links ]
. . . 1 ) . the type locality , and hence the description of b . havanensis , is currently in dispute ( paraense and deslandes 1958 ; yong , pointier , and perera 1997 ) . our aim here is not to determine the correct type locality , but our analyses do show that there are two distinct species in cuba that have been linked to the name b . havanensis . . . .
conchological , anatomical and genetic characteristics were compared among several populations of the biomphalaria havanensis complex from cuba and other localities in the caribbean region . two morphometric measures ( height and diameter ) distinguished two closely similar morphs of the shell . the reproductive system also separated these morphs , particularly upon comparison of sizes of the penis . . . [ show full abstract ]
. . . the observation of two profiles for all analyzed snails demonstrated that among those populations only b . havanensis ( canasi , arroyo arenas , hanabanilla , santa rita , guatao , and hanabanilla vaqueria ) and b . obstructa ( zanja ferrer ) species were found . therefore , it is important to mention that ( a ) zanja ferrer and calle pila have recently been reported as type localities of b . havanensis ( yong et al . 1997 ) , ( b ) yong ( 1998 ) and durand et al . ( 1998 ) reported the presence of biomphalaria sp . in canasi , arroyo arenas , santa rita , guatao , and hanabanilla , and ( c ) b . peregrina has also been reported in hanabanilla ( ) . we showed that pcr - rflp of the biomphalaria rdna its region , using ddei , haeiii , mnli or alui enzymes allowed the differentiation of two cuban biomphalaria species . . . .
. . . more recently , a revision was undertaken on a particularly confusing group occurring in the caribbean area , which includes two taxa commonly reported in the literature , b . havanensis ( pfeiffer , 1839 ) and b . obstructa ( morelet , 1849 ) . the type locality of b . havanensis is a swampy area in the vicinity of havana , cuba , which was recently redescribed by yong , pointier & perera ( 1997 ) . the morphology ( shell and reproductive system ) of topotypic snails which were collected by these authors , was different from the material previously described by paraense & deslandes ( 1958 ) from another cuban locality erroneously considered by these authors as the type locality for b . havanensis . . . .
paraense wl 1966 . biomphalaria amazonica and b . cousini , two new species of neotropical planorbid molluscs . rev bras biol 26 : 115 - 126 . [ links ]
paraense wl 1988 . biomphalaria kuhniana ( clessin , 1883 ) , planorbid mollusc from south america . mem inst oswaldo cruz 83 : 1 - 12 . [ links ]
paraense wl , corrêa lr 1985 . further experiments on susceptibility of biomphalaria amazonica to schistosoma mansoni . mem inst oswaldo cruz 80 : 259 - 262 . [ links ]
justification : biomphalaria havanensis has been assessed as least concern as this species has a wide distribution , is presumed to be abundant , occurs in a number of protected areas and shows tolerance to a broad range of habitats and to habitat modification . there are no immediate threats and this species is not declining in numbers or is unlikely to be declining at a fast enough rate to qualify for listing in a more threatened category .
paraense wl 1981 . biomphalaria occidentalis sp . n . from south america ( mollusca : basommatophora pulmonata ) . mem inst oswaldo cruz 76 : 199 - 211 . [ links ]
paraense wl , corrêa lr 1978 . differential susceptibility of biomphalaria tenagophila populations to infection with a strain of schistosoma mansoni . j parasitol 64 : 822 - 826 . [ links ]
barbosa fs , coelho mv , carneiro e 1956 . cross - breeding of australorbis glabratus and biomphalaria boissyi . trans r soc trop med hyg 50 : 296 - 297 . [ links ]
corrêa lr , paraense wl 1971 . susceptibility of biomphalaria amazonica to infection with two strains of schistosoma mansoni . rev inst med trop sao paulo 13 : 387 - 390 . [ links ]
. . . nus , oncomelania and biomphalaria ( hope & mcmanus 1994 , caldeira et al . 1998 , jones et al . 1999 , spatz et al . 1999 ) . in cuba , malacological surveys have been considered very important to better know the geographical distribution and also to help the health institution programs , in order to prevent the introduction of schistosomiasis in the country . yong et al . ( 1997 ) discussed about the type locality of b . havanensis suggesting that specimens described by paraense and deslandes ( 1958 ) , as b . havanensis ( collected in the country club of havana ) , may in fact belong to another species . the first authors suggest that the country club of havana species does not correspond to the type locality of b . havan . . .
caldeira rl , teodoro tm , gomes mfb , carvalho os 2010 . preliminary studies investigating the occurrence of biomphalaria cousini in brazil . mem inst oswaldo cruz 105 : 485 - 487 . [ links ]
paraense wl , corrêa lr 1973 . susceptibility of biomphalaria peregrina from brazil and ecuador to two strains of schistosoma mansoni . rev inst med trop sao paulo 15 : 127 - 130 . [ links ]
. . . according to yong ( 1998 ) the topotypic b . obstructa , reviewed by paraense ( 1990 ) , would correspond perfectly to the topotypic b . havanensis and would be a synonym of the latter . in spite of the important questions on b . obstructa ( yong 1998 ) and the type locality of b . havanensis ( yong et al . 1997 ) , in the present work , the cuban snail populations were identified in accordance with the classical morphological systematics adopted by paraense , which has produced the most complete description of the majority of the neotropical biomphalaria species ( paraense 1975a , b , 1990 , 1996 , paraense et al . 1992 ) . as a result of our current analysis , two clear profiles were obtained with ddei enzyme , identifying two species as shown by morphological criteria . . . .
vidigal thda , caldeira rl , simpson ajg , carvalho os 2000 . further studies on the molecular systematics of biomphalaria snails from brazil . mem inst oswaldo cruz 95 : 57 - 66 . [ links ]
paraense wl 1984 . biomphalaria tenagophila guaibensis ssp . n . from southern brazil and uruguay ( pulmonata : planorbidae ) . i . morphology . mem inst oswaldo cruz 79 : 465 - 469 . [ links ]
. . . is the locality for b . havanensis , only one population has been found in nature and it is the least represented species of the genus ( yong et al . 1997 ) . this species has also been reported in other caribbean areas , such as the dominican republic and venezuela ( yong et al . 2001 ) . . . .
lewis fa , richards cs , knight m , cooper la , clark b 1993 . schistosoma mansoni - analysis of an unusual infection phenotype in the intermediate host snail biomphalaria glabrata . exp parasitol 77 : 349 - 361 . [ links ]
. . . the morphology ( shell and reproductive system ) of topotypic snails which were collected by these authors , was different from the material previously described by paraense & deslandes ( 1958 ) from another cuban locality erroneously considered by these authors as the type locality for b . havanensis . moreover , the morphological characteristics of topotypic specimens of b . havanensis described by yong et al . ( 1997 ) appear very similar to those of snails described by paraense ( 1990 ) from isla del carmen , mexico , the type locality for b . obstructa . a multilocus enzyme electrophoresis study ( durand , yong , perera , ducreux & pointier , 1998 ) , showed the presence of two distinct genetic entities within this group of cuban snails . . . .
malek , e . a . 1969 . studies on
tropicorbid
snails ( biomphalaria : planorbdae ) from the caribbean and gulf of mexico areas , including the southern united states . malacologia 7 ( 2 / 3 ) : 183 - 209 .
thus , the identification of biomphalaria snails is of great importance because the presence of the species ( b . amazonica , b . cousini and hybrids between than ) may result in schistosomiasis foci because these species are experimentally susceptible to s . mansoni .
. . . b . havanensis seems very uncommon in cuba because this species was collected only at two sites within the area of the type locality in havana ( yong , pointier & perera , 1997 ) . this snail is more commonly distributed in the southern part of united states and mexico ( malek , 1969 ( malek , , 1985naranjo - garcía , 1983 ) . . . .
teodoro tm , janotti - passos lk , carvalho os , caldeira rl 2010 . occurrence of biomphalaria cousini ( mollusca : gastropoda ) in brazil and its susceptibility to schistosoma mansoni ( platyhelminths : trematoda ) . mol phylogenet evol 57 : 144 - 151 . [ links ]
experiments based on crosses of molluscs of the genus biomphalaria have been performed since the 1950s , using the albinism factor as a genetic marker , which is determined by a pair of recessive alleles ( paraense 1955 , 1956 , paraense & deslandes 1955 ) . crosses between b . glabrata and b . tenagophila were not feasible reporting the reproductive isolation of these species ( paraense & deslandes 1955 ) . nevertheless , crosses between biomphalaria species that produce fertile hybrids have been reported ( barbosa et al . 1956 , 1958 , mello - silva et al . 1998 ) .
carvalho os , jannotti - passos lk , caldeira rl 2008 . importância epidemiológica e biologia molecular aplicada ao estudo dos moluscos do gênero biomphalaria . in os carvalho , pmz coelho , hl lenzi , schistosoma mansoni e esquistossomose : uma visão multidisciplinar , fiocruz , rio de janeiro , p . 311 - 345 . [ links ]
jannotti - passos lk , vidigal thda , dias - neto e , pena sdj , simpson ajg , dutra wo , souza cp , carvalho - parra jf 1997 . pcr amplification of the mitochondrial dna minisatellite region to detect schistosoma mansoni infection in biomphalaria glabrata snails . j parasitol 83 : 395 - 399 . [ links ]
jannotti - passos lk , caldeira rl , carvalho os 2008 . técnicas utilizadas no estudo dos moluscos do gênero biomphalaria e na manutenção do ciclo de schistosoma mansoni . in os carvalho , pmz coelho , hl lenzi , schistosoma mansoni e esquistossomose : uma visão multidisciplinar , fiocruz , rio de janeiro , p . 531 - 544 . [ links ]
genetic examination of molecular phylogeny of global planorboidea at the coi and 18s molecular markers indicates all north american taxa within the subfamily planorbinae form a well - supported clade , as yet unnamed but termed c - clade ( albrecht et al . , 2007 ) . this species is sometimes referred to as biomphalaria obstructa ( morelet , 1849 ) , a likely synonym .
although morphological studies of biomphalaria for species identification have been carried out , differentiation between some species may be complicated . the identification of these molluscs is based on anatomical and morphological characteristics and relies on descriptions by paraense ( 1966 , 1975 , 1981 , 1984 , 1988 ) . currently , the most important tool for specific identification of biomphalaria snails is the researcher ' s capability to dissect and characterise snails based on their morphological differences . the difficulties involved in specific identification based on morphological characters have motivated the use of molecular techniques ( vidigal et al . 2000 ) . therefore , the correct identification of these snails is of great importance because it enables the detection of species in areas where schistosomiasis transmission occurs , as well as in areas without the disease but that might become schistosomiasis foci owing to the presence of natural or experimentally susceptible species .
previous studies have suggested the possibility of natural hybrids between b . amazonica and b . cousini ( teodoro et al . 2010 ) . our results leave no doubt about the possibility of a hybrid between b . amazonica and b . cousini , which we confirmed through a cross between these species using the albinism factor as a genetic marker ( determined by a pair of recessive alleles ) , as standardised to biomphalaria by paraense ( 1955 ) .
experiments based on crosses of molluscs of the genus biomphalaria have been performed since the 1950s ( paraense 1955 , 1956 , paraense & deslandes 1955 ) . crosses between biomphalaria species that produced fertile hybrids have been reported . examples include crosses between b . alexandrina and b . glabrata ( barbosa et al . 1956 ) : b . straminea and b . peregrine , b . tenagophila and b . peregrina ( barbosa et al . 1958 ) and b . glabrata and b . tenagophila ( mello - silva et al . 1998 ) . barbosa ( 1964 ) commented on the natural hybridisation between b . glabrata and b . tenagophila , which generates hybrids with intermediate forms . subsequently , barbosa ( 1973 ) described observations made over three years on the replacement of b . glabrata by b . straminea in northeastern brazil , where four specimens that he called interspecific hybrids were found . only during this process of competitive exclusion were mixed forms found , possibly as a result of crosses between these species . later , new field studies were conducted and these intermediate forms were not found , revealing their inability to persist in nature .
crossing trials - the pcr amplification of the biomphalaria its region with the primers etts2 and etts1 resulted in a product of approximately 1 . 3 kb ( data not shown ) . the rflp profiles obtained by digesting rdna its with dde i are represented in figure , with the profile of the albino snail corresponding to the profile of b . amazonica ( lane 12 ) and the pigmented snail corresponding to b . cousini ( lane 13 ) . the profiles of the 10 pigmented offspring obtained from each albino snail correspond to the profile of the hybrid control ( lane 14 ) . the five crosses generated the same result and only one gel is represented .
studies using biomphalaria snails found three variant molecular profiles for b . amazonica ( vidigal et al . 2000 ) . later , studies using phylogeny , morphological and molecular taxonomy of brazilian , bolivian and colombian snails previously identified as b . amazonica showed that some populations studied were b . cousini and established the occurrence of b . cousini in brazil . three different molecular profiles were observed , enabling the differentiation of b . amazonica from b . cousini . the third profile showed associations with the two b . cousini and b . amazonica profiles , suggesting the possibility of hybrids between these species ( caldeira et al . 2010 , teodoro et al . 2010 ) . experiments on susceptibility have shown that b . amazonica and b . cousini are potential hosts of s . mansoni ( corrêa & paraense 1971 , paraense & corrêa 1985 , fernandez & thiengo 2006 , teodoro et al . 2010 ) .
lynx haven crossing , south fork , guadalupe river , kerr co . , texas . ( 8 . 5 mm . ) . digital images by david kirsh
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the atlantic drainages of georgia , comprising approximately 40 % of the waters of the state , are presently covered . . . ››
the four ecoregions of north carolina drain toward the atlantic through six major river systems . . . ›› more
although perhaps not as environmentally heterogeneous as neighboring states , south carolina does include . . . ›› more
the survey reported here is focused on the eastern 75 % of the state , draining from the blue ridge to the atlantic . . . ›› more
all waters draining into the tennessee river above the alabama line , including sw virginia , w north carolina , and n georgia . . . ›› more
a study area of approximately 150 , 000 km2 , including delaware , maryland , new jersey , eastern pennsylvania and the west virginia panhandle . . . ›› more
in a us atlantic drainage was that of dillon & dutra - clarke ( 1992 ) from a public park in charleston , sc . there is also an older ( 1960 ) record from
mcintosh co , ga
in the georgia museum of natural history that we have been unable to confirm . in 2015 we discovered a third
population on hilton head island ( beaufort co , sc ) . see my essay of 16dec15 from the link below for more .
as texas , louisiana , florida , mexico , puerto rico , and cuba , adding that “it has also been reported from california , but was probably introduced there . ” it seems quite likely to us that all the
populations in our study area result from artificial introductions as well . the ponds and ditches in charleston and hilton head are largely artificial , and the areas heavily disturbed .
population has historically been closely associated with floating vegetation . in years when park managers have allowed the vegetation
have been targeted for extensive research ( malek 1958 , dillon 2000 : 61 - 66 ) . the charleston population seems to display the annual , semelparous life cycle
sampled from cuba may be susceptible ( michelson 1976 ) . i myself have not observed any parasitism in the charleston population .
back in the early 1990s dr . emile malek confirmed that our charleston population was specifically identical to populations widespread in florida , texas , and louisiana . but for years there was no consensus regarding the identity of any
type population in cuba ( paraense & deslandes 1958 , yong et al . 1997 ) . the more recent genetic and morphological studies of yong et al . ( 2001 ) and dejong et al . ( 2001 ) have , however , firmed up the identity of
in cuba and convincingly demonstrated a match of our u . s . populations to it .
. but the classification of the planorbidae proposed by the tag team of baker ( 1945 ) and hubendick ( 1955 ) remains , after 50 years , the basis for our understanding of this large and diverse family of pulmonates worldwide . see my essay of 11apr08 ( available below ) for more .
see my post to the fwgna blog of 11apr08 for a review of the classification of the planorbidae .
or view the ( hubendick 1955 ) classification of north american planorbids in a tabular format [ here ] .
index to f . c . baker ' s
the molluscan family planorbidae .
nautilus , 59 , 127 - 41 .
north american freshwater snails . malacological publications , hamburg , mi . 365 pp .
the ecology of freshwater molluscs . cambridge , cambridge university press . 509 pp .
phylogeny in the planorbidae . trans . zool . soc . london 28 : 453 - 542 .
i ( saudi arabia strain ) . j . parasitol . , 64 : 940 - 941 .
factors conditioning the habitat of bilharziasis intermediate hosts of the family planorbidae . bull . w . h . o . , 18 : 785 - 818 .
: planorbidae ) from the caribbean and gulf of mexico areas , including the southern united states . malacologia , 7 : 183 - 209 .
snail hosts of schistosomiasis and other snail - transmitted diseases in tropical america : a manual . washington , d . c . , pan american health organization . 325 pp .
( pulmonata : planorbidae ) . revista brasileira de biologia 18 : 87 - 91 .
yong , m . , gutierrez , a . , perera g . , durand p . & pointier j - p . ( 2001 )
complex ( gastropoda : planorbidae ) in cuba : a morphological and genetic study . journal of molluscan studies 67 : 103 - 111 .
yong , m , pointier j - p . & perera , g . ( 1997 )
please cite as : dillon , r . t . , jr . , b . t . watson , t . w . stewart & w . k . reeves 2006 . the freshwater gastropods of north america . internet address : urltoken
department of biology , college of charleston , charleston , sc 29424 p : 843 . 953 . 8087 f : 843 . 953 . 5453
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thank you for taking the time to provide feedback on the iucn red list of threatened species website , we are grateful for your input .
johnson , p . d . , bogan , a . e . , brown , k . m . , burkhead , n . m . , cordeiro , j . r . , garner , j . t . , hartfield , p . d . , lepitzki , d . a . w . , mackie , g . l . , pip , e . , tarpley , t . a . , tiemann , j . s . , whelan , n . v . and strong , e . e . 2013 . conservation status of freshwater gastropods of canada and the united states . fisheries 38 ( 6 ) : 247 - 282 .
dyer , e . , soulsby , a . - m . , whitton , f . , mcguinness , s . , de silva , r . , milligan , h . t . , kasthala , g . , thorley , j . , herdson , r . , mcmillan , k . & collins , a .
this species is widely distributed in the west indies and central america ( thompson 1999 ) . in north america , it is distributed in parts of florida , louisiana and texas , arizona , and mexico ( burch 1989 ) . it is also known from cuba , but only from havana ( pointier
this species is abundant , particularly in the southern united states and mexico ( yong et al . 2001 ) .
2009 ) as a result of its ability to serve as an intermediate trematode host .
this species occurs in a number of protected areas and shows tolerance to a broad range of habitats and to a certain degree of habitat modification . there are no immediate threats and this species is not declining in numbers or is unlikely to be declining at a fast enough rate to impact the global population of this species .
this species occurs in a number of protected areas . there are no species - specific conservation measures in place .
to make use of this information , please check the < terms of use > .
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the nonindigenous occurrences section of the nas species profiles has a new structure . the section is now dynamically updated from the nas database to ensure that it contains the most current and accurate information . occurrences are summarized in table 1 , alphabetically by state , with years of earliest and most recent observations , and the tally and names of drainages where the species was observed . the table contains hyperlinks to collections tables of specimens based on the states , years , and drainages selected . references to specimens that were not obtained through sighting reports and personal communications are found through the hyperlink in the table 1 caption or through the individual specimens linked in the collections tables .
snails in the family planorbidae have a discoidal shape and flattened spire , adults have 5 or more whorls ( burch and tottenham 1980 ) .
southern florida , central and southern louisiana , and much of texas ( malek 1969 ) ; arizona , mexico , and central america ( bequaert and miller 1973 as cited in burch and tottenham 1980 ) ; puerto rico and cuba ( burch and tottenham 1980 ) .
bequaert , j . c . and w . b . miller . 1973 . the molluscs of the arid southwest , with an arizona checklist . the university of arizona press , tucson , xvi , 271 pp .
bowler , p . a . , and t . j . frest . 1991 . the non - native snail fauna of the middle snake river , southern idaho . proceedings of the desert fishes council 23 : 28 - 44 .
burch , j . b . , and j . l . tottenham . 1980 . north american freshwater snails : species list , ranges and illustrations . walkerana - transactions of the poets society 1 ( 3 ) : 81 - 215 .
this information is preliminary or provisional and is subject to revision . it is being provided to meet the need for timely best science . the information has not received final approval by the u . s . geological survey ( usgs ) and is provided on the condition that neither the usgs nor the u . s . government shall be held liable for any damages resulting from the authorized or unauthorized use of the information .
the data represented on this site vary in accuracy , scale , completeness , extent of coverage and origin . it is the user ' s responsibility to use these data consistent with their intended purpose and within stated limitations . we highly recommend reviewing metadata files prior to interpreting these data .
citation information : u . s . geological survey . [ 2018 ] . nonindigenous aquatic species database . gainesville , florida . accessed [ 7 / 9 / 2018 ] .
contact us if you are using data from this site for a publication to make sure the data are being used appropriately and for potential co - authorship if warranted . for queries involving fish , please contact pam fuller . for queries involving invertebrates , contact amy benson .
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turgeon , d . d . , a . e . bogan , e . v . coan , w . k . emerson , w . g . lyons , w . pratt , et al .
turgeon , d . d . , j . f . quinn , jr . , a . e . bogan , e . v . coan , f . g . hochberg , w . g . lyons , et al .
common and scientific names of aquatic invertebrates from the united states and canada : mollusks , 2nd ed .
disclaimer : itis taxonomy is based on the latest scientific consensus available , and is provided as a general reference source for interested parties . however , it is not a legal authority for statutory or regulatory purposes . while every effort has been made to provide the most reliable and up - to - date information available , ultimate legal requirements with respect to species are contained in provisions of treaties to which the united states is a party , wildlife statutes , regulations , and any applicable notices that have been published in the federal register . for further information on u . s . legal requirements with respect to protected taxa , please contact the u . s . fish and wildlife service .
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seasonal studies of two populations of tarebia granifera . mary yong , ricardo sanchez , gloria perera , . . .
turgeon , d . d . , j . f . quinn , jr . , a . e . bogan , e . v . coan , f . g . hochberg , w . g . lyons , p . m . mikkelsen , r . j . neves , c . f . e . roper , g . rosenberg , b . roth , a . scheltema , f . g . thompson , m . vecchione , and j . d . williams . 1998 . common and scientific names of aquatic invertebrates from the united states and canada : mollusks . 2nd edition . american fisheries society special publication 26 , bethesda , maryland : 526 pp .
it is widely distributed in the west indies and central america , as well as mexico and the american southeast . this species has a wide distribution , presumed large population , occurrence in a number of protected areas , tolerance of a broad range of habitats , tolerance to habitat modification , lack of substantial immediate threats , and because it is not in decline or is unlikely to be declining fast enough to qualify for listing in a more threatened category .
due to latency between updates made in state , provincial or other natureserve network databases and when they appear on natureserve explorer , for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data . please refer to
this species is widely distributed in the west indies and central america ( thompson , 1999 ) . in north america , it is distributed in parts of florida , louisiana and texas ; arizona , and mexico ( burch , 1989 ) . it is also known from cuba but only from havana ( pointier et al . , 2005 ) . burch ( 1989 ) also cites puerto rico .
common in ditches and ponds in southeast florida from brevard and okeechobee cos . south to monroe co . , florida ( thompson , 1999 ) . an introduced population was discovered in south carolina pond at charles towne landing state park in charlestown ( dillon and stewart , 2003 ) . dillon et al . ( 2006 ) report a specimen lot at the georgia museum of natural history from mcintosh co . , but this occurrence has not been confirmed .
( 200 , 000 - 2 , 500 , 000 square km ( about 80 , 000 - 1 , 000 , 000 square miles ) ) this species is widely distributed in the west indies and central america ( thompson , 1999 ) . in north america , it is distributed in parts of florida , louisiana and texas ; arizona , and mexico ( burch , 1989 ) . it is also known from cuba but only from havana ( pointier et al . , 2005 ) . burch ( 1989 ) also cites puerto rico .
occurrences are based on some evidence of historical or current presence of single or multiple specimens , including live specimens or recently dead shells ( i . e . , soft tissue still attached without signs of external weathering or staining ) , at a given location with potentially recurring existence . weathered shells constitute a historic occurrence . evidence is derived from reliable published observation or collection data ; unpublished , though documented ( i . e . government or agency reports , web sites , etc . ) observation or collection data ; or museum specimen information .
separation barriers are largely based on permanent hydrological discontinuity between water bodies , with distances of 30 meters or greater between maximum high water marks constituting a separation barrier . additional barriers are chemical and / or physical and include any connecting water body ( regardless of size ) with one or more of the following on a permanent basis : no dissolved calcium content , acidity greater than ph 5 , lack of dissolved oxygen , extremely high salinity such as that found in saline lakes and brine waters , or temperature greater than 45 an additional physical barrier , particularly for flowing water , is presence of upland habitat between water connections . high waterfalls and anthropogenic barriers to water flow such as dams are barriers as they limit movement in an upstream direction .
use the generic guidelines for the application of occurrence ranks ( 2008 ) . the key for ranking species occurrences using the generic approach provides a step - wise process for implementing this method .
zoological data developed by natureserve and its network of natural heritage programs ( see local programs ) and other contributors and cooperators ( see sources ) .
albrecht , c . , k . kuhn , and b . streit . 2007 . a molecular phylogeny of planorboidea ( gastropoda , pulmonata ) : insights from enhanced taxon sampling . zoologica scripta , 36 : 27 - 39 .
burch , j . b . 1989 . north american freshwater snails . malacological publications : hamburg , michigan . 365 pp .
dillon , r . t . , jr . and t . w . stewart . 2003 . the freshwater gastropods of south carolina . created 26 august 2003 . last updated september 2007 . available online : urltoken
dillon , r . t . , jr . , w . k . reeves , and t . w . stewart . 2006b [ 2007 ] . the freshwater gastropods of georgia . created 26 august 2003 . last updated september 2007 . available online : urltoken
pointier , j . - p . , m . yong , and a . gutierrez . 2005 . guide to the freshwater molluscs of cuba . conchbooks : hackenheim , germany . 120 pp .
thompson , f . g . 1999 . an identification manual for the freshwater snails of florida . walkerana 10 ( 23 ) : 1 - 96 .
thomson , j . w . 1984 . american arctic lichens . 1 . the macrolichens . columbia university press , n . y . 504 pp .
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ridgely , r . s . , t . f . allnutt , t . brooks , d . k . mcnicol , d . w . mehlman , b . e . young , and j . r . zook . 2003 . digital distribution maps of the birds of the western hemisphere , version 1 . 0 . natureserve , arlington , virginia , usa .
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mem . inst . oswaldo cruz vol . 106 no . 7 rio de janeiro nov . 2011
i laboratório de helmintologia e malacologia médica ii moluscário lobato paraense , instituto de pesquisas rené rachou - fiocruz , av . augusto de lima 1715 , 30190 - 001 belo horizonte , mg , brasil iii biomedical sciences department , tropical disease institute , college of osteopathic medicine , ohio university , athens , oh , usa iv center for infectious disease research , school of biological sciences , pontifical catholic university of ecuador , quito , ecuador
in this context , the aim of this work was to investigate the presence of hybrids between b . cousini from the type locality santo domingo de los colorados , ecuador and b . amazonica from the type locality igarape da cachoeirinha , manaus , state of amazonas ( am ) , brazil and to verify if these species are susceptible to s . mansoni .
snail populations - the molluscs used for crosses experiments were from type localities defined by paraense ( 1966 ) . b . cousini snails were collected from santo domingo de los colorados ( 0024303s 07913934w ) . b . amazonica snails were collected from igarape da cachoeirinha ( 3073274s 60003261w ) .
the molluscs used for susceptibility experiments were from benjamin constant , am ( b . cousini and b . amazonica ) , which exhibited the same profile as the snails from the type locality belo horizonte , minas gerais ( mg ) , brazil ( b . glabrata ) and descended from the cross between b . amazonica and b . cousini from benjamin constant ( hybrids ) .
the specimens were identified by polymerase chain reaction - restriction fragment length polymorphism ( pcr - rflp ) using the internal transcribed spacer ( its ) region of the rdna with the primers etts2 ( 5 ' taacaaggtttccgtaggtgaa3 ' ) and etts1 ( 5 ' tgcttaagttcagcgggt3 ' ) , which were anchored in the conserved extremities of the 18s and 28s ribosomal genes , respectively ( kane & rollinson 1994 ) . dde i enzyme was used for each digestion reaction . the snails were maintained in the lobato paraense mollusc rearing of rené rachou research institute / oswaldo cruz foundation under appropriate environmental conditions and were fed lettuce leaves and calcium carbonate .
cross trials - populations of b . cousini and b . amazonica that were kept in the lobato paraense mollusc rearing and had been previously identified by pcr - rflp were used . using albinism as the genetic marker , five crosses were performed using albino b . amazonica and pigmented b . cousini snails as described below .
one adult albino b . amazonica and one pigmented b . cousini were kept separate until they had laid eggs . after hatching , 10 offspring from each species were identified by pcr - rflp and were kept isolated until they had laid eggs for the first time . five albino b . amazonica and five pigmented b . cousini snails were selected for the crosses . each couple was kept in the same container for one week and then was separated . ten pigmented offspring obtained from each albino snail were individually identified by pcr - rflp to verify the profile of these descendents .
susceptibility experiments - the hybrids were susceptible to the le , sj and al strains of s . mansoni , ( 4 . 4 % , 5 . 6 % and 2 . 2 % , respectively ) . all infected snails shed cercariae , which were determined to be viable by mouse inoculation . b . amazonica and b . cousini were also susceptible to the le , sj and al strains of s . mansoni ( 22 . 2 % , 18 . 9 % and 26 . 6 % to b . amazonica and 11 . 1 % , 8 . 9 % and 3 . 3 % to b . cousini , respectively ) ( table ) .
confirming our speculations , the pigmented descendents obtained from each albino snail had the hybrid profile ( figure ) . thus , this study confirms that these two species produce hybrids . mello - silva et al . ( 1998 ) commented that the results of fertility and fecundity tests of fertile hybrids revealed that the f2 generation was not as efficient as the f1 , involving the formation of normal , vigorous and fertile f1 generation hybrids , but a weak f2 generation with the presence of some sterile individuals . however , further studies are necessary to better understand the biology of these hybrids .
mortality was observed in the snails used as the negative control . this relatively short survival can be attributed to the long period of experimentation and to adjustments to the artificial environment .
to the lobato paraense mollusc rearing of cpqrr / fiocruz , to the support for this research , and to josé geraldo amorim da silva and jairo , cpqrr / fiocruz , for technical support .
barbosa fs 1964 . the renal ridge a disputed feature of anatomy of planorbid snail australorbis tenagophilus . rev inst med trop sao paulo 6 : 64 - 70 . [ links ]
barbosa fs 1973 . possible competitive displacement and evidence of hybridization between two brazilian species of planorbid snails . malacologia 14 : 401 - 408 . [ links ]
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