BioIn asions Reco ds (2019) Volume 8, Issue 4: 838–847
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 838
CORRECTED PROOF
Resea ch A icle
In asion o he da k alse mussel in sh imp a ms in Venezuela:
species iden i ica ion and gene ic analysis
Césa Lodei os1,2,*, Nie es González-Hen íquez3, Jo ge Cuélla -Anjel4, Dailos He nández-Reyes3,
Ca olina Medina-Alca az3, Ja ie Quin ei o5 and Manuel Rey-Méndez5
1G upo de In es igación en Biología y Cul i o de Moluscos, Escuela de Acuicul u a y Pesque ía, Facul ad de Ciencias Ve e ina ias,
Uni e sidad Técnica de Manabí, Bahía de Ca áquez, Ecuado
2Ins i u o Oceanog á ico de Venezuela, Uni e sidad de O ien e, Cumaná 6101, Venezuela
3Labo a o io BioMol, Depa amen o de Biología, Facul ad de Ciencias del Ma , Uni e sidad de Las Palmas de G an Cana ia, España
4Global Consul ing Inc. Bogo á, D.C., Colombia
5Labo a o io de Sis emá ica Molecula e Ins i u o de Acuicul u a, CIBUS, Campus Vida, Uni e sidade de San iago de Compos ela, 15782-
San iago de Compos ela, Spain
Au ho e-mails: cesa lodei [email protected] (CL), nie [email protected] (NGH), [email p o ec ed] (JCA), [email p o ec ed] (DHR),
[email p o ec ed]s (CMA), [email p o ec ed] (JQ), [email p o ec ed] (MRM)
*Co esponding au ho
Abs ac
An inspec ion was ca ied ou a sh imp a ms loca ed Sou h Wes o Lake Ma acaibo
(Zulia S a e, Venezuela), wi h high incidences o mussel bi al e. Mo phological and
gene ic analysis helped o iden i y he species as he da k alse mussel My ilopsis
leucophaea a Con ad, 1831, and his is he i s eco d o he species om opical
wa e s o no he n Sou h Ame ica. The highes incidences o mussels we e de ec ed
in ponds and channels bu no li e mussels we e obse ed in he coas al in e idal
a ea su ounding he en ances o he a ms, al hough emp y shells we e de ec ed
he e, sugges ing hei o me p esence. The en i onmen al condi ions o he a i icial
sys em o sh imp cul u e, is a niche sui able o he p oli e a ion o he bi al e. The
consequences o he p esence o his bi al e in he p oduc ion o sh imp a e discussed.
Key wo ds: D eissenidae, in asi e alien species, bi al e mollusks, sh imp cul u e
In oduc ion
In aqua ic ecosys ems, bi al e mollusks a e one o he mos in asi e g oups.
They can cause en i onmen al dis u bances due mainly o hei high
il e ing capaci y, which can p oduce quan i a i e and quali a i e changes
in he plank on, causing an imbalance in he ood chain and hus a high
clea ance o wa e . Bi al es also p oduce eces and pseudo eces ha can
al e he biogeochemical cycles and shells can change he physical
p ope ies o sedimen (S aye 1999; Ruesink e al. 2005).
Ponds and channels in sh imp a ms a e a i icial ecosys ems
cha ac e ized by high loads o nu ien s and o ganic ma e in suspension,
including phy oplank on. Op imal condi ions o ac o s such as salini y,
empe a u e, concen a ions o dissol ed oxygen, u bidi y and
phy oplank on, should be main ained o p ope sh imp de elopmen
Ci a ion: Lodei os C, González-Hen íquez
N
, Cuélla
-Anjel J, He nández-Reyes D,
Medina-Alca az C, Quin ei o J, Rey-
Méndez M (2019) In asion o he da k
alse mussel in sh imp a ms in Venezuela:
species iden i ica ion and gene ic analysis.
B
ioIn asions Reco ds 8(4): 838–847,
h ps://doi.o g/10.3391/bi .2019.8.4.12
Recei ed: 21 Augus 2018
Accep ed: 5 Oc obe 2019
Published: 21 No embe 2019
Handling edi o : Ca h yn Abbo
Thema ic edi o : S elios Ka sane akis
Copy igh : © Lodei os e al.
This is an open access a icle dis ibu ed unde e ms
o he C ea i e Commons A ibu ion License
(A ibu ion 4.0 In e na ional - CC BY 4.0).
OPEN ACCESS.
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 839
(Cuélla -Anjel e al. 2010). Howe e , op imal main enance o balanced
ecosys em condi ions can be al e ed when in asi e bi al es o e come hese
ba ie s and ind a o able niches in he sh imp a ms o coloniza ion and
es ablishmen . This can lead o low yields o sh imp and signi ican
economic losses (Ald idge e al. 2008).
This s udy esul ed om he obse a ion o low pe o mance in sh imp
p oduc ion in cul i a ion ponds loca ed in a eas adjacen o Lake
Ma acaibo, in Zulia S a e, Wes e n Venezuela, associa ed wi h he in asion
o mussels (36% loss o no mal p oduc ion). The co ec iden i ica ion o
he in asi e species and i s incidence in he sh imp a m allowed us o
desc ibe i s po en ial impac on sh imp a ming and o discuss possible
in asion con ol measu es.
Ma e ials and me hods
Loca ion and cha ac e is ics o he sh imp a ms
The inspec ed sh imp a ms a e con iguous and loca ed in he Sou h
Wes e n side o Lake Ma acaibo (Zulia S a e, Venezuela), in he icini y o
Villa del Rosa io de Pe ijá (~ 10°08′N; 71°57′W). The wo a ms a e composed
o se e al cul u e ponds ha ep esen an a ea o app oxima ely 2000 ha,
wi h semi-in ensi e a ming sys em (using densi ies o 8–25 sh imp/m2)
wi h p oduc ions close o 2000–3000 kg/ha pe cycle, wi h 2.5 c op cycles
pe yea . The a ea is cha ac e ized by mang o es in he coas al zone, and
con inen al ege a ion inland, mainly opical d y o semi-d y o es . The
ainy season, whose in ensi y and equency is a iable, o en occu s
be ween Ap il and Decembe . Fa ms ha e pa ial wa e eci cula ion
sys ems and pe o m daily wa e exchange in he ange o 5 o 7%; wa e is
ob ained om he lake, whose salini y anges be ween 3 and 6 PSU, o
eplace losses h ough e apo a ion and seepage.
A mon hly eco d (da a supplied by he sh imp a ms) was aken o he
main en i onmen al ac o s moni o ed du ing 2017 in mo e han 500 ponds,
which we e wi hin he a ms unde s udy. Tempe a u e and dissol ed
oxygen we e moni o ed daily, bo h in he mo ning and in he a e noon,
while u bidi y, pH and ni i es we e moni o ed weekly o biweekly.
Cha ac e iza ion o he in asi e species
A isual inspec ion o ponds and wa e channels was pe o med, p ima ily
de ailing he slopes o he banks and he wa e inle s and ou le s in he
ponds (Figu e 1). The densi y o mussel popula ions in he ponds was
quan i ied by es ima ing hei numbe om 20 quad an s o 25 × 25 cm,
a anged andomly in ou ha es ed ponds (o 3 ha each). The specimens
o he in asi e species we e coun ed and hei an e o-pos e io axis leng h
was measu ed wi h a digi al calipe (0.01 mm o p ecision), o de e mine
he size- equency s uc u e o he es ablished popula ions.
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 840
Figu e 1. Es ablishmen o he in asi e mussel in he ponds and di e en s uc u es in sh imp a ms.
F om he sampled mussels, subsamples (n = 60) we e aken o
mo phological iden i ica ion acco ding o desc ip ions o Ma elli and G ay
(1983), Mackie and Claudi (2010) and he iden i ica ion key o eshwa e
mussels o Williams e al. (2014). Also, isual obse a ions o he gonad
we e made o es ima e ep oduc i e s a us o he o ganisms.
A se o man le issue samples we e p ese ed in 80% e hanol un il DNA
ba coding analysis. To al DNA ex ac ion was pe o med wi h he
“E.Z.N.A. Mollusk DNA Ki ” (Omega Bio-Tek). Pa ial COX1 gene PCR
p oduc s we e ob ained wi h he LCO1490/HCO2198 ba coding p ime s
se (Folme e al. 1994), pu i ied wi h ExoSAP-I ki (A ime ix) and
sequenced using he BigDye ki 3.1 and X e mina o in an ABI 3500
equipmen (The mo Fishe Scien i ic).
Sequences we e edi ed, aligned and dis ances we e es ima ed wi h he
so wa e MEGA 7.0.26 (Tamu a e al. 2013). BLAST (h p://blas .ncbi.nlm.
nhi.go ) and BOLD (www.ba codingli e.o g) we e used o sea ch o
a ailable homologous sequences simila o iden ical o he de ec ed
haplo ypes. The molecula -based axonomy o analyzed samples was
e alua ed by hei posi ion wi hin species-speci ic clades ob ained by
phylogene ic in e ence. Thus, an alignmen o 137 COX1 pa ial sequences
om M. sallei and M. leucophaea a, ob ained in he p esen wo k and
mined om GenBank (Table 1), was used o esol e e olu iona y ela ionships
in ol ing hese species lineages. A phylogene ic ee was in e ed MEGA
7.0.26 (Tamu a e al. 2013) using he neighbo -joining algo i hm and
based in Tamu a-Nei dis ances. To isualize haplo ype ela ionships and
equencies a Median-joining ne wo k (Bandel e al. 1999) was
cons uc ed wi h PopA 1.7 (Leigh and B yan 2015).
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 841
Table 1. Loca ion, GenBank accession numbe , and au ho i y o COX1 My ilopsis leucophaea a
sequences analyzed in he p esen s udy.
A ea Loca ion Accession No. Re e ence
No h Ame ica USA MLU47649 Baldwin e al. 1996
Ma yland Chesapeake Bay, USA KU905843 unpublished
Ma yland Chesapeake Bay, USA KU905894
Ma yland Chesapeake Bay, USA KU905989
Ma yland Chesapeake Bay, USA KU905904
Ma yland Chesapeake Bay, USA KU906056
Eu ope Se illa, Spain MF401396 Clusa e al. 2017
An we p, Belgium EF414477 Alb ech e al. 2007
An we p, Belgium HM100251 Heile e al. 2010
Kiel Canal, Ge many HM100253
Kiel Canal, Ge many HM100254
Uk aine, Sou he n Bug, HM100255
Asia Caspian Sea, I an HM100255 Heile e al. 2010
HM100257
HM100258
HM100259
HM100260
Sou h Ame ica Rio de Janei o, B azil MF139836–MF139866 Fe nandes e al. 2018
Lake Ma acaibo, Venezuela MK843806/VZ05 This S udy
MK843803/VZ06
MK843802/VZ07
MK843802/VZ09
MK843804/VZ23
MK843805/VZ24
Resul s and discussion
Bo h sh imp a ms showed a high incidence o mussels in he d ainage
channels, g ow-ou ponds and ese oi s. They we e a ached o a ious
solid subs a es (shells, conc e e and me al s uc u es, mesh, wood, plas ic,
s ems o he g ass B achia ia sp., e c.) and pa icula ly on conc e e in inle
and ou le s uc u es o ponds, as well as on ae a o equipmen inside he
ponds (Figu e 1). The highes incidence o mussels was obse ed in ponds
and channels. In he in e idal coas al zone, nex o he a m’s en ance, no
li e mussels we e obse ed, al hough he e we e emp y shells, sugges ing
hei o me p esence in his a ea.
Taxonomic iden i ica ion
The in ading species was iden i ied as M. leucophaea a, based on he a io
o he maximum an e o-pos e io o do so- en al leng h o he shell smalle
han 1.3 mm (popula ion mean o 1.27 ± 0.097 mm), which gi es an elonga ed
appea ance and sho in heigh . This ea u e, oge he wi h he absence o
do sal cu a u e o he shell, p esen ing he sep um do sally cu ed and
wi h cle s, as well as he apophysis associa ed o sep um and wide and long
muscle sca in he shell (Figu e 2), de ined mo phologically he s udied
specimens as da k alse mussel, M. leucophaea a (Ma elli and G ay 1983).
The mo phological-based species iden i ica ion was co obo a ed by
gene ic analysis. We ob ained a se o 6 pa ial COX1 sequence 658 bp
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 842
Figu e 2. Shell o My ilopsis leucophaea a om his s udy, showing he main mo phological
cha ac e is ics o i s iden i ica ion.
leng h and 5 dis inc haplo ypes. Two haplo ypes (VZ05:MK843806,
VZ07/VZ09:MK843802) we e iden ical o haplo ypes assigned o
M. leucophaea a species, ecen ly deposi ed in he GenBank/BOLD
da abase and sampled a Rio de Janei o (B azil) (Fe nandes e al. 2018).
The o he 3 dis inc haplo ypes showed pai wise iden i y alues > 99.7% o
M. leucophaea a sequences, a ypical alue expec ed om in aspeci ic
compa isons. The mean gene ic p-dis ance among hese sequences was
low, only 0.005 (S.E. = 0.001).
A single in asi e haplo ype has been de ec ed in No h Ame ica (Hudson
Ri e ) (Baldwin e al. 1996), Belgium (An we p) (Alb ech e al. 2007),
Ge many (Bal ic and No h Sea), Moldo a, Uk aine, Russia (Black sea), I an,
Russia, Kazakh an (Caspian Sea) (Zhulido e al. 2018) and Spain (Clusa e
al. 2017). I s mean pai wise iden i y espec o Venezuela ( his s udy) and
B azil (Fe nandes e al. 2018) haplo ypes (Table 1) was 97.5% and he ne
p-dis ance be ween he wo haplo ype clus e s was 0.021 (S.E. = 0.001).
The mean in a-speci ic dis ance om all a ailable da a wi hin M.
leucophaea a was 1.2%, a alue close o he mean in a-speci ic alue
(0.96%) es ima ed om 2,801 sequences in 569 species o Paci ic mollusks
(Sun e al. 2016). In con as , wi hin he congene ic My ilopsis sallei, he
mean in a-speci ic alue was highe (5%) wi h he p esence o pa icula ly
di e gen lineages (p-dis ance = 0.1).
The neighbo -joining ee suppo ed he monophyly o M. leucophaea a
(99% boo s apping alue) and exhibi ed wo in a-speci ic clades, one o
hem ep esen ed by he single in asi e haplo ype (95%), and he o he
including samples om B azil and Venezuela (68%) (Figu e 3A). The
haplo ype ne wo k displays a s a -con o ma ion o he B azil/Venezuela
clade, sugges ing a ecen demog aphic expansion, and sha ed common
haplo ypes o he B azilian and Venezuelan samples (Figu e 3B). These
esul s likely sugges a common o igin o he in asi e coloniza ion a
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 843
Figu e 3. A. Neighbo -joining ee o My ilopsis sallei and My ilopsis leucophaea a es ima ed
om a 439 bp alignmen o a ailable COX1 sequence da a mined om Genbank and sequences
gene a ed he e om Venezuela. I was based in Tamu a-Nei (1993) dis ances, wi h a shape
pa ame e o he gamma dis ibu ion = 0.39. Clade suppo was es ima ed by boo s apping
(2000 eplica es) and a e shown (i > 65%) nex o he nodes. As an ou g oup D eissena
bugensis was included. Loca ion speci ic colo s a e de ailed in legend, including ed colo
coding o p esen s udy samples. B. Haplo ype ne wo k using a median joining algo i hm om
he My ilopsis leucophaea a COX1 sequence alignmen . Each geog aphically colo -coded ci cle
ep esen s a single haplo ype, whose a ea is p opo ional o he haplo ype equency. The ha ch
ma ks in he connec ing lines indica e he numbe o mu a ional s eps be ween haplo ypes.
Rio de Janei o and Ma acaibo a he sou h o i s na i e ange and he
disassocia ion om hose in asi e e en s in ol ing he no he n haplo ype.
A main d awback o he e olu iona y analysis o de ec ed clades is he
absence o sequence da a om he na i e geog aphic ange sugges ed o
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 844
Figu e 4. Size s uc u e (an e o-pos e io leng h, mm) o he in asi e mussel in samples om
he wa e supply channel and 4 di e en ponds o he sh imp a m (Ponds A o D).
M. leucophaea a. Ne e heless, he alse mussels ound he e in aqueous
sys ems o Lake Ma acaibo sh imp a ms belong o M. leucophaea a and
he e o e his cons i u es he i s eco d o he species in wa e s o
no he n Sou h Ame ica.
Densi y, size dis ibu ion and onse o mussel ep oduc ion in sh imp ponds
In ponds analyzed, a co e age o he mussel agg ega ions o 10–30% was
es ima ed, and popula ion densi y in he agg ega ions a e aged 13,200 ±
1,905 indi iduals/m2, wi h densi ies ha exceeded 20,000 indi iduals/m2.
The dis ibu ion o sizes in he di e en places whe e mussels we e
collec ed in he sh imp a ms (Figu e 4), showed se e al modes, sugges ing
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 845
Table 2. Mon hly a e age (± SD) o physical and chemical pa ame e s om wa e in cul i a ion ponds du ing 2017. T anspa ency
was es ima ed wi h Secchi disc dep h.
Mon h Tempe a u e a.m. (°C) Tempe a u e p.m. (°C) Oxygen a.m. (mg/L) Oxygen p.m. (mg/L) T anspa ency (cm)
Janua y 27.8 ± 0.8 29.7 ± 1.2 3.5 ± 0.3 9.5 ± 0.5 44.5 ± 37.0
Feb ua y 27.9 ± 0.7 29.9 ± 1.1 3.6 ± 0.4 9.5 ± 0.5 31.2 ± 16.2
Ma ch 27.9 ± 0.8 29.7 ± 1.0 3.5 ± 0.3 9.5 ± 0.6 32.6 ± 15.3
Ap il 29.4 ± 0.9 31.3 ± 1.5 3.4 ± 0.3 9.4 ± 0.6 35.5 ± 16.9
May 30.3 ± 0.8 32.1 ± 1.4 3.4 ± 0.4 9.2 ± 0.8 32.8 ± 15.1
June 30.2 ± 0.6 31.8 ± 1.2 3.4 ± 0.4 9.1 ± 0.9 34.9 ± 18.1
July 30.6 ± 0.6 32.3 ± 1.4 3.5 ± 0.5 9.2 ± 0.8 36.5 ± 16.8
Augus 30.6 ± 0.6 32.3 ± 1.3 3.5 ± 0.4 9.1 ± 0.8 35.4 ± 15.0
Sep embe 30.5 ± 0.6 32.3 ± 1.4 3.4 ± 0.6 9.1 ± 0.7 28.4 ± 10.0
Oc obe 30.1 ± 0.8 31.8 ± 1.5 3.5 ± 0.4 9.1 ± 0.8 23.5 ± 6.30
No embe 30.1 ± 0.8 31.9 ± 1.5 3.5 ± 0.4 9.1 ± 0.8 23.5 ± 6.3
Decembe 28.7 ± 0.8 30.8 ± 1.3 3.5 ± 0.4 9.5 ± 0.9 25.0 ± 7.7
di e en imes o g ow h and in asion. The channel showed a wide
popula ion s uc u e indica ing di e en ec ui men s. Un o una ely, o
his analysis he e we e no eco ds o he ime when he ponds we e illed
up; only pond A, which had been in ope a ion o 36 days. This pond
showed a unimodal dis ibu ion o small size mussels wi h an a e age
leng h o 4.9 ± 1.53 mm. The la al de elopmen o he species akes place
wi hin 6–8 days (Siddall 1980), es ima ing abou 30 addi ional days o
each a leng h o 4 mm, which sugges s he la ae in aded he ponds a he
beginning o pond seeding wi h sh imp. The i s size o o ganisms
o ming ma u e gonads is 8–10 mm (Mackie and Claudi 2010), which
could be eached in abou 50 days, demons a ing ha he mussel can
ep oduce se e al imes wi hin he cul u e pond, because he sh imp
cul u e cycle is be ween 90 and 120 days.
The o ma ion o ep oduc i e issue in small-sized indi iduals was
e i ied; gonad issue was obse ed in mussels o 14–16 mm. Mos
indi iduals had li le o no ep oduc i e issue, indica ing low ep oduc i e
ac i i y a he ime o sampling.
En i onmen al ac o s in sh imp a ms
Salini y is no eco ded on a egula basis in he sampled sh imp a ms, as
a e he o he ac o s lis ed in Table 2. Howe e , alues on a e age o 4.8
PSU we e obse ed, wi hin an in e al o 3 o 6.2 PSU du ing 2017. The
echnicians o he sh imp a m sugges an associa ion o he p esence o
da k alse mussels wi h consecu i e d ough s, which could ha e allowed a
highe salini y in he sh imp a ms, acili a ing he p oli e a ion o he
in asi e species, which can suppo om eshwa e (< 0.5 PSU) o ma ine
condi ions (Siddall 1980; The iaul e al. 2004). Howe e , hei popula ions
no mally occu in oligohaline o mesohalines condi ions, in gene al anges
o 0.5–10 PSU (Kennedy 2011), which coincides wi h he no mal salini y
eco ded in he e alua ed sh imp a ms and in Lake Ma acaibo (annual
ange o 2–8 PSU; José-B acho e al. 2016). Measu emen s o empe a u e,
oxygen, and u bidi y ound in ponds (Table 2) we e consis en wi h hose
My ilopsis leucophaea a in asion in sh imp a ms in Venezuela
Lodei os e al. (2019), BioIn asions Reco ds 8(4): 838–847, h ps://doi.o g/10.3391/bi .2019.8.4.12 846
known o be app op ia e sys ems o he p oli e a ion o M. leucophaea a
(Kennedy 2011).
The geog aphic dis ibu ion o M. leucophaea a (bo h na i e and in aded)
anges om loca ions o cold empe a u e in New England, USA (Hall
1964), o opical a eas such as es ua ine sys ems in he S a e o Pe nambuco
in B azil (de Souza e al. 2005; Fa apei a e al. 2007). Acco ding o hese
da a and s udies on empe a u e ole ance, he species can be conside ed
an eu y he m (Kennedy 2011).
Conclusion and ecommenda ions
The p esence o M. leucophaea a epo ed in his s udy ep esen s he i s
eco d o he species in opical wa e s o no he n Sou h Ame ica. The
species is an in ade ha can p oduce mul iple e ec s on pond sys ems,
and may con ibu e o dec eases in he p oduc ion o a med sh imp.
S udies on he biological cycle, popula ion dynamics, ep oduc ion, and
g ow h, in associa ion wi h en i onmen al bio ic and abio ic ac o s; as
well as physiological e ec s o molluscicides o he use o biocon ols and
possible mussel-sh imp in e ac ions a e ecommended. In e ms o
managemen and ope a ion o he sh imp a ms, p e en ion and apid
con ol is desi ed, hence he ea ly moni o ing o la ae in he wa e and
isualiza ion o he in ade s on subs a es is ecommended, pa icula ly a
he beginning o each g owing cycle. Species iden i ica ion wi h he help o
molecula ools could aise he abili y o de ec , e en a a la al s age, an
incipien ea ly in asion ha may imp o e he chances o quick esponses
o e adica e o con ain he sp ead o he in asi e species.
Acknowledgemen s
The collabo a ion o he e alua ed sh imp a ms is app ecia ed, as well as ha o J. Fe nández,
J. Ma ínez, M.G and M.A. He e a o hei echnical suppo in he collec ion and p ocessing
o samples. We a e g a e ul o J. Alió o English e iewing and p o iding help ul commen s.
Also we hank CYTED P og am (AquaCibus ne wo k 318RT0549 “S eng hening aquacul u e
in Ibe oame ica: quali y, compe i i eness and sus ainabili y”) o p omo ing in e ac ions among
au ho s. We app ecia e he sugges ions and co ec ions made by he e iewe s and edi o s,
which subs an ially imp o ed he a icle.
Con lic o In e es
The au ho s decla e ha hey ha e no con lic o in e es .
Re e ences
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Molecula phylogene ics, biogeog aphy and sys ema ics o D eissena in he Balkans.
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Ald idge DC, Salaza M, Se na A, Cock J (2008) Densi y dependen e ec s o a new in asi e
alse mussel, My ilopsis au wineana (T yon 1866), on sh imp, Li openaeus annamei
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j.aquacul u e.2008.05.022
Baldwin BS, Black M, Sanju O, Gus a son R, Lu z RA, V ijenhoek RC (1996) A diagnos ic
molecula ma ke o zeb a mussels (D eissena polymo pha) and po en ially co-occu ing
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