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Cap i e b eeding o Eu opean eshwa e mussels as a conse a ion ool : A e iew
© 2023 The Au ho s. Aqua ic Conse a ion: Ma ine and F eshwa e Ecosys ems published by John Wiley & Sons L d.
Published e sion
Geis , Jue gen; Thielen, F ankie; La ic oi e, Louise; Hoess, Rebecca; Al muelle ,
Reinha d; Baud imon , Magalie; Blaize, Ch is ine; Campos, Miquel; Ca oll, Paul;
Daill, Daniel; Degelmann, Wol gang; De me , Raine ; Denic, Ma co; Du y,
Pie ick; de Ey o, El i a; G unicke, Felix; Gumpinge , Clemens; Jakobsen, Pe J.;
Kaldma, Ka in; Klaas, Kunna ; Legeay, Alexia; Mage oy, Jon Hamne ; Moo kens,
E elyn A.; Mo e, G égo y; Nakamu a, Keiko; Ondina, Paz; Ös e ling, Ma in;
Pichle ‐Schede , Ch is ian; Spisa , Ondřej; Reis, Joaquim; Schneide , Lea D.;
Schwa ze , A no; Selheim, Heidi; Sole , Joaquín; Taskinen, Jouni; Taylo , John;
S achan, Ben; Wengs öm, Niklas; Zając, Tadeusz
Geis , J., Thielen, F., La ic oi e, L., Hoess, R., Al muelle , R., Baud imon , M., Blaize, C., Campos,
M., Ca oll, P., Daill, D., Degelmann, W., De me , R., Denic, M., Du y, P., de Ey o, E., G unicke,
F., Gumpinge , C., Jakobsen, P. J., Kaldma, K., . . . Zając, T. (2023). Cap i e b eeding o Eu opean
eshwa e mussels as a conse a ion ool : A e iew. Aqua ic Conse a ion : Ma ine and
F eshwa e Ecosys ems, 33(11), 1321-1359. h ps://doi.o g/10.1002/aqc.4018
2023
REVIEW ARTICLE
Cap i e b eeding o Eu opean eshwa e mussels as a
conse a ion ool: A e iew
Jue gen Geis
1
| F ankie Thielen
2
| Louise La ic oi e
3
| Rebecca Hoess
1
|
Reinha d Al muelle
4
| Magalie Baud imon
5
| Ch is ine Blaize
6
|
Miquel Campos
7,8
| Paul Ca oll
9
| Daniel Daill
10
| Wol gang Degelmann
11
|
Raine De me
12
| Ma co Denic
13
| Pie ick Du y
14
| El i a de Ey o
15
|
Felix G unicke
16
| Clemens Gumpinge
10
| Pe J. Jakobsen
17
|
Ka in Kaldma
18,19
| Kunna Klaas
19
| Alexia Legeay
5
|
Jon Hamne Mage oy
20
| E elyn A. Moo kens
21
| G égo y Mo e
22
|
Keiko Nakamu a
23,24
| Paz Ondina
25
| Ma in Ös e ling
26
|
Ch is ian Pichle -Schede
10
| Ondˇ
ej Spisa
27
| Joaquim Reis
28
|
Lea D. Schneide
29
| A no Schwa ze
30
| Heidi Selheim
31
| Joaquín Sole
32
|
Jouni Taskinen
33
| John Taylo
34
| Ben S achan
35
| Niklas Wengs öm
36
|
Tadeusz Zając
37
Co espondence
Jue gen Geis , Aqua ic Sys ems Biology Uni ,
Depa men o Li e Science Sys ems, Technical
Uni e si y o Munich, Mühlenweg 22,
D-85354 F eising, Ge many.
Email: [email p o ec ed]
Funding in o ma ion
COST (Eu opean Coope a ion in Science and
Technology), G an /Awa d Numbe : COST
Ac ion CA18239
Abs ac
1. F eshwa e mussels a e declining h oughou hei ange. Thei impo an
ecological unc ions along wi h insu icien le els o na u al ec ui men ha e
p omp ed cap i e b eeding o popula ion augmen a ion and ques ions abou he
use ulness and applicabili y o such measu es.
2. This a icle e iews he cu en s a e o cap i e b eeding and ea ing p og ammes
o eshwa e mussels in Eu ope. I conside s he a ious species, s a egies, and
echniques o p opaga ion, as well as he di e en le els o e o equi ed
acco ding o ea ing me hod, highligh ing he key ac o s o success.
3. Wi hin he las 30 yea s, 46 b eeding ac i i ies in 16 Eu opean coun ies ha e
been epo ed, mainly o Ma ga i i e a ma ga i i e a and Unio c assus. Some
acili ies p opaga e species ha a e in a e y c i ical si ua ion, such as Pseudunio
au icula ius,Unio mancus, and Unio a oisie i, o mul iple species concu en ly. In
some s eams, he numbe o eleased cap i e-b ed mussels al eady exceeds he
size o he emaining na u al popula ion.
Jue gen Geis , F ankie Thielen and Louise La ic oi e a e he P incipal au ho s.
Fo a ilia ions e e o page 35
Recei ed: 28 Ap il 2023 Re ised: 20 July 2023 Accep ed: 31 Augus 2023
DOI: 10.1002/aqc.4018
This is an open access a icle unde he e ms o he C ea i e Commons A ibu ion License, which pe mi s use, dis ibu ion and ep oduc ion in any medium,
p o ided he o iginal wo k is p ope ly ci ed.
© 2023 The Au ho s. Aqua ic Conse a ion: Ma ine and F eshwa e Ecosys ems published by John Wiley & Sons L d.
Aqua ic Conse : Ma F eshw Ecosys . 2023;1–39. wileyonlinelib a y.com/jou nal/aqc 1
4. Rea ing e o s ange om highly in ensi e labo a o y incuba ion o lowe
in ensi y me hods using in- i e mussel cages o silos. Mos b eeding e o s a e
unded by na ional and EU LIFE(+) g an s, a e well documen ed, and conside he
gene ic in eg i y o he p opaga ed mussels. Limi ed long- e m unding
pe spec i es, he a ailabili y o expe ienced s a , wa e quali y, and eeding/
su i al du ing ea ly li e s ages a e seen as he mos impo an challenges.
5. Success ul cap i e b eeding p og ammes need o be combined wi h es o a ion o
he habi a s in o which he mussels a e eleased. This wo k will bene i om an
e idence-based app oach, knowledge exchange among acili ies, and an o e all
b eeding s a egy comp ising mul iple coun ies and conse a ion uni s.
KEYWORDS
aquacul u e, cap i e b eeding, conse a ion ansloca ion, eshwa e mussel cul u ing,
Ma ga i i e a ma ga i i e a, p opaga ion, ein oduc ion, Unio c assus
1|INTRODUCTION
The global decline o eshwa e biodi e si y has p omp ed many
e o s add essing i s conse a ion (Geis , 2011). This holds
pa icula ly ue o eshwa e mussels, which a e among he mos
h ea ened axa h oughou Eu ope (Lopes-Lima e al., 2017). F om a
conse a ion pe spec i e, endange ed eshwa e mussels, pa icula ly
he eshwa e pea l mussel (Ma ga i i e a ma ga i i e a L.), a e
conside ed a ge species o he conse a ion o aqua ic ecosys ems,
as hey simul aneously ul il he c i e ia o lagship, indica o ,
keys one, and umb ella species (Geis , 2010). The con inuing declines
and lack o na u al ec ui men o eshwa e mussels in Eu ope has
led o an inc easing numbe o ea ing app oaches o popula ion
augmen a ion. I has also led o con o e sies abou he use ulness o
cap i e b eeding in he con ex o mussel conse a ion, pa icula ly i
such ac i i ies a e no p ope ly moni o ed o a e conduc ed wi hou
conside ing habi a es o a ion (P es on, Keys & Robe s, 2007;
Schmid & Vand é, 2010; Gum, Lange & Geis , 2011; Pa e son
e al., 2018).
Rea ing me hods de eloped o No h Ame ican mussel species
and condi ions (Ga enby, Ne es & Pa ke , 1996; Bea y &
Ne es, 2004; Ne es, 2004; Jones, Mai & Ne es, 2005;
Ba nha , 2006; Hua & Ne es, 2007; Pa e son e al., 2018) ha e
o en been ans e ed o Eu opean mussel species, bu wi hou
sys ema ic e alua ion o he success o such ac ions. Recen s udies
e eal ha some cap i e b eeding p ac ices, such as he use o a low
numbe o pa en s, can esul in e osion o he gene ic cons i u ion o
o sp ing compa ed wi h he o iginal popula ions, bo h in eshwa e
mussels (Geis e al., 2021) as well as ishes (S oeckle e al., 2022). Co-
adap a ion o co-e olu ion o mussels o ce ain ish hos s (Geis &
Kuehn, 2008; Taeube e al., 2010; Taeube , Gum & Geis , 2012;
Salonen e al., 2017; Taskinen & Salonen, 2022), he e ec s o hos
ish age (Ma waha e al., 2019), du a ion o he pa asi ic phase
(Ma waha e al., 2017), wa e empe a u e (Taeube , El-Nobi &
Geis , 2014), and ea ing condi ions (Eybe e al., 2013; Eybe
e al., 2015; La ic oi e e al., 2016; La ic oi e e al., 2020) all ha e an
impac on he pe o mance o cap i e-b ed mussels, wi h po en ial
consequences i such in e ac ions a e no aken in o accoun . C oss-
exposu e expe imen s indica e ha s ock o igin and en i onmen al
condi ions a ec bo h he su i al and he g ow h o ju enile
eshwa e mussels a e hei elease (Denic e al., 2015). Based on
an ea lie e iew o Eu opean and No h Ame ican cap i e b eeding
p og ammes, Gum, Lange & Geis (2011) sugges ed ha cap i e
b eeding should only be a escue ool o e ain he e olu iona y
po en ial o p io i y popula ions ha would no pe sis long enough o
bene i om habi a es o a ion p ac ices. This is in line wi h
Ry winski e al. (2021), who iden i ied a need o e idence o e alua e
he e ec i eness o conse a ion-o ien ed cap i e b eeding and
elease p og ammes o impe illed eshwa e mussels. This is
pa icula ly impo an gi en he wo ldwide inc ease in cap i e
b eeding p og ammes o highly h ea ened eshwa e mussels
(S aye e al., 2004; Ba nha , 2006; Thomas, Taylo & Ga cia de
Leaniz, 2010; Gum, Lange & Geis , 2011; Pa e son e al., 2018).
The aim o his a icle is o e iew he cu en s a e o eshwa e
mussel cap i e b eeding p og ammes in Eu ope. The c i ical
challenges associa ed wi h hese ac i i ies we e iden i ied om
pe sonal in e iews wi h key g oups in ol ed in he cap i e b eeding
o eshwa e mussels h oughou Eu ope, including in o ma ion on
he con ex o hese p og ammes, he co e age o species, he
in ensi y and ype o ea ing p ac ices, and hei imelines. This
in o ma ion was hen used o make ecommenda ions o e ec i e
conse a ion o eshwa e mussels h ough cap i e b eeding, and o
iden i y p io i ies o i s u u e use.
2|METHODS
All he ins i u ions and o ganiza ions lis ed in Table 1we e con ac ed
by one o he h ee main au ho s and in e iewed using he same
ques ions, o ga he s anda dized in o ma ion. The in e iews ook
2GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 Cap i e b eeding me hods o eshwa e mussels u ilized by di e en p ojec s in Eu ope. See able legend o explana ion o abb e ia ions.
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
Aus ia Uppe Aus ia/
bla isch e.U.
Consul ancy
Company
M.m. 2011–p esen LI: medium–high
HF: Salmo u a
(ha che y)
MB: kep
pe manen ly a he
b eeding acili y
IC: incuba o boxes
OC: sedimen boxes,
Buddensiek cages,
and mussel silos in
he s eam
In o al: 2,100 ea ed
ind. s ill in caging
sys ems.
30 mussels (11 yea s
old) eleased
(compa ed wi h 19
emaining adul
mussels in his
popula ion)
Su i al o mussels
>1 mm in he i s
win e is abou
50%
Na ional & EU PM and
JM
High mo ali y
among ju eniles
sho ly a e
ha es ing
High mo ali y
among pa en
mussels, wi h low
i ness p esumed
Long- e m unding
unsu e
Gumpinge C., Daill
D. (pe s. comm.,
2022)
Pichle -Schede C.
(pe s. comm.,
2011)
Geis e al. (2021)
Belgium A dennes Region/
Se ice Public de
Wallonie (SPW)
Since 2017
coope a ion wi h
Luxembou g (see
Luxembou g in his
Table)
M.m. 2005–p esen LI: medium
HF: Salmo u a
(ha che y)
MB: glochidia
collec ed in he
ield; mussels s ay
in he i e
IC: none
OC: side channel in
ca chmen o i e s
A lune and Rulles
Be ween 2007 and
2014, 95% o
eshly
me amo phosed
mussels eleased in
wo ou doo
ea ing channels.
No su i al o no
mussels ound
du ing ollow-up
moni o ing in he
yea s 2007, 2014,
and 2018
5% o ju eniles
eleased di ec ly in
Ri e Anlie (2% o
his popula ion
come om
cul u ed ind.)Rulles
Ri e : 99% o his
popula ion come
om cul u ed ind.
Na ional & EU
LIFE-p ojec
PM Limi ed budge and
s a
Cleaning o channels
di icul because o
changing s a
Eme ging bea e
popula ions make
i di icul o ind
good elease si es
Geis e al. (2021);
Mo e G. (pe s.
comm., 2022)
Czech
Republic
Sou h Bohemia/
Na u e
Conse a ion
Agency o he
M.m. 1990–p esen LI: high
HF: Salmo u a (wild
and ha che y)
App ox. 1,000
eleased in o Zla ý
po ok and app ox.
50,000 eleased
Na ional & EU PM Collec ion o na u al
ood (de i us) in
win e di icul
Spisa O., Zelenko á
E., ˇ
S aˇ
íˇ
cko á J. &
Do B. (pe s.
comm., 2022)
(Con inues)
GEIST ET AL.3
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
Czech Republic,
ˇ
Suma a Na ional
Pa k
Adminis a ion;
p ac ical wo k
Gamma us cz, s. .
o., Bohumil Do
Wes Bohemia
BIVALVIA s. .o.:
Ondˇ
ej Spisa
MB: glochidia
collec ed in he
ield
IC: incuba o boxes
OC: sedimen boxes
in side channels
in o Blanice in
1995–; in 2019, 72
ind. 6+; in 2022,
322 ind. 6+
Blanice: eleased
ju eniles make up
10% o he o al
popula ion
Vl a a Ri e :
May/June 2021,
elease o 1,000 8–
10 yea olds;
May/June 2022,
elease o >1,000
6–7 yea olds,
1,000 5-yea -old
ind. a e planned o
be eleased
2023/2027
Malse Ri e : app ox.
300 small ju eniles
ha e al eady been
eleased; app ox.
1,000 ind. a e
planned o be
eleased in
2022/24
Luˇzní Po ok: 810
ju eniles aged 3+
o 5+yea s
eleased in 2016–
2019
Di icul ies in
moni o ing
eleased ju eniles
Vl a a Ri e : ca p-
like ish (cyp inids)
sp ead ups eam
om Lipno dam,
wi h impac s on
ou , which a e
being displaced
om mussel
locali ies
Long- e m unding is
unsu e
Ensu e he quali y
and quan i y o
b eeding s a ion
s a
H uska (1999);
H uska (2000);
ˇ
S anyga e al.
(2013); Simon e al.
(2015); Simon e al.
(2017); Bílý e al.
(2018); ˇ
Ce ná e al.
(2018); Bílý e al.
(2021); Miloˇ
s
Holub (pe s.
Comm., 2022)
Es onia No h o Es onia/
S a e Fo es
Managemen
Cen e
M.m. 2020–p esen LI: medium–high
HF: Salmo u a
(wild)
MB: mussels s ay in
he i e and a e
no dis u bed.
Hos ish a e
In o al 10,000 0+
mussels in
incuba o boxes
and app ox. 440
1+and 150 2+
mussels in he
Na ional & EU-
LIFE p ojec s
No –plan
o do in
nea
u u e
Mussel g ow h in
incuba o boxes is
low (less han 1
mm on a e age) by
he end o he i s
au umn; as a
esul , mo ali y in
Klaas K. & Kaldma K.
(pe s. comm.,
2023)
4GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License

TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
caugh by
elec o ishing and
de ached ju eniles
a e collec ed wi h
ne s. App ox. 100
in es ed ou a e
caugh om
be ween h ee and
se en i e
sec ions and
anspo ed o a
qua an ine cen e.
I possible, hos
ish a e eleased
a e he collec ion
o M.m. ju eniles
IC: incuba o boxes
OC: Buddensiek
cages
na i e i e in
Buddensiek cages
Incuba o boxes:
du ing he i s
summe ju enile
su i al was 70%.
Du ing he i s
yea , mussel
su i al was
app ox. 60%
Buddensiek cages in
i e : in he i s
summe ju enile
su i al was 25%.
In he i s win e
0+mussel su i al
was 17%. In he
second summe
1-yea -old mussel
su i al was 65%.
In he second
win e 1+mussel
su i al was 82%
he cages was high
in he i s win e
Sil and ine
sedimen s in he
i e
Ac i e bea e
popula ion, despi e
cons an hun ing
Finland Uni e si y o
Jy äskylä/
Konne esi
Resea ch S a ion
M.m. 2016–p esen LI: high
HF: Salmo u a
(ha che y), Salmo
sala (ha che y).
Local ish used, i
a ailable in
ha che ies. I local
ish no a ailable,
a leas he
p e e ed hos
(salmon o ou )
used, i he mos
sui able hos ish
species is known. I
i is no known
whe he he M.m.
Ri e Mus ionjoki/
S a å: 200 1-yea -
old ju eniles
in oduced in
g a el boxes in
2019; 93% ali e
a e 3 yea s.
14,000 0+
ju eniles
in oduced in
g a el boxes in
2022. Cu en
popula ion size in
i e is 1,200.
Adul mussels ha e
los hei
Na ional, EU & EU
LIFE p ojec s
PM P oblem wi h coppe
in pipes when new
acili y es ablished
Only wo sui able
de i us sou ces.
Collec ion o
de i us di icul in
win e
P oblems wi h
clogging o cul u e
ins alla ions
De achmen o high
numbe s o
ju eniles wi hin a
sho pe iod o
ime—high
Hy ä inen e al.
(2021)
Taskinen J. (pe s.
comm., 2022).
(Con inues)
GEIST ET AL.5
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
popula ion is
adap ed o salmon
o ou , hen bo h
species a e used a
he beginning
MB: glochidia
collec ed in he
ield and glochidia
collec ed om
adul mussels kep
in he acili y. In
wo cases, mussels
in poo condi ion
we e ehabili a ed
in he acili y o 2
yea s and hey
s a ed eleasing
glochidia. Adul
mussels ed wi h
shell ish die ,
Nanno, and
de i us
IC: incuba o boxes
and ough/ lume.
Pulsed low-
h ough sys em
es ed bu no in
egula use now.
Fed wi h shell ish
die , Nanno, and
de i us
OC: in- i e cages,
sedimen boxes,
and Buddensiek
cages
ep oduc i e
capaci y, bu
eco e ed wi h
2-yea
ehabili a ion in
cap i i y
Ri e Äh ä änjoki/
Esseå: 1,200
2-yea -old
ju eniles
in oduced in
g a el boxes in
2021; 94% su i al
o e 1 yea .
Cu en size o he
na u al popula ion
is 800. Adul
mussels ha e los
hei ep oduc i e
capaci y, bu
eco e ed wi h
2-yea
ehabili a ion in
cap i i y
Ri e Lu o: 2,000
0+ju eniles
in oduced in
g a el boxes in
2021; 60% su i al
o e 1 yea . 4,000
0+-yea -old
ju eniles
in oduced in
g a el boxes in
2022. Cu en size
o he na u al
popula ion is
30,000, bu he
equi ed salmon
hos canno ascend
o he i e
seasonal a ia ion
in labo a o y
Con inua ion and
sus ainabili y o
unding
6GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
Ri e Isojoki:
125,000 0+
ju eniles
in oduced in
g a el boxes and
Buddensiek cages
in 2022. Cu en
size o he na u al
popula ion is 150
F ance B i any &
No mandy/
Fédé a ion de
Pêche du Finis è e
B e agne Vi an e,
CPIE des Collines
No mandes
M.m. 2011–p esen LI: high
HF: Salmo u a
(ha che y)
MB: glochidia
collec ed in he
ield
IC: ough/ lume ed
wi h comme cial
algae
OC: side channel,
sedimen boxes,
mussel silos, in-
i e cages
P oduc ion: since
2012, se e al
million 0+M.m.
ha es ed and
cul u ed in he
ough/ lume (in
six i e s)
Release: se e al
millions, bu 96%
we e 0+, wi h li le
indica ion o
su i al. Olde
mussels eleased a
3–4 cm in size
(kep inside acili y
o 3 yea s and
u he kep in
ou doo sys ems
un il hey each 3–
4 cm)
Release o app ox.
200,000 ind.
anging in age
be ween 1+and
4+yea s
Na u al popula ion
consis s o app ox.
14,000 adul s
Na ional & EU
LIFE-p ojec
PM & JM Cessa ion o g ow h
in indoo sys ems
a e abou 3–4
yea s
Funding di icul ies
a e 10 yea s o
ope a ion
Geis e al. (2021);
Blaize C. (pe s.
comm. 2022)
F ance Dépa emen
Do dogne –
Région Nou elle
Aqui aine/
M.m. 2016–2020 LI: high
HF: Salmo u a
(ha che y)
Du ing he LIFE
p og amme a o al
o app ox. 30,000
ju eniles >8
EU LIFE-p ojec PM No con inua ion o
ea ing ac i i ies a
he end o he LIFE
p ojec
Legeay A. &
Baud imon M.
(pe s. comm.,
2022)
(Con inues)
GEIST ET AL.7
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
Uni e si é de
Bo deaux - Pa c
Na u el Régional
Pé igo d Limousin
MB: glochidia
collec ed in he
ield
IC: ough/ lume ed
wi h comme cial
and eshly
cul u ed algae
OC: in- i e cages
(cylinde cages)
mon hs (3,300
aged 3 yea s and
be ween 5–9mm
+4,300 2 yea s +
14,800 1 yea +
8,000 8 mon hs)
and 115,000 pos -
pa asi ic ind. ha e
been ein oduced
di ec ly in he i e
50% o his
popula ion come
om cul u e
No moni o ing o
eleased mussels
as he LIFE p ojec
ended
Cul u e o esh algae
as ood di icul
F om ime o ime
wa e quali y
issues (pollu ion)
om i e used o
cul u e
P eda ion o ju enile
mussels in sand
ough sys ems
Belamy e al. (2020)
F ance Region Cen e-Val de
Loi e/
Dépa emen
Ind e-e -Loi e/
Uni e si é de
Tou s
P.a. 2014–2018 LI: high
HF: Acipense bae ii
(ha che y)
MB: new mussels
collec ed in he
wild e e y yea
and kep a
b eeding acili y
un il elease o
glochidia
IC: incuba o boxes
and ough/ lume
ed wi h
comme cial algae
die , sel -p epa ed
de i us wi h
lea es. Use o egg
whi e as p o ein
sou ce
OC: none
Ju eniles collec ed/
yea : >10,000
No able o p oduce
animals olde han
4 mon hs
No su i al in
a i icial channel
No elease o olde
ju eniles
F eshly
me amo phosed
ju eniles eleased
(2,000)
No moni o ing o
eleased ju eniles
EU LIFE-p ojec PM & JM Wa e quali y used in
cul u e. Sil ?
Pollu ion?
Su i al low du ing
i s ew mon hs
P ojec du a ion was
sho . No enough
s a o ea ing
ac i i y
Sole J. (pe s. comm.,
2022).
Sole e al. (2018a);
Sole e al.
(2018b); Sole
e al. (2019);
Wan zen e al.
(2019)
Ge many Lowe Saxony/
Uni e si y o
Hanno e , Lowe
Saxony S a e
M.m. 1986–2002;
2009–
p esen
(Ge dau)
LI: Low –medium
HF: Salmo u a
(wild)
Su i al a e 5%–
20% a e 1–2
yea s and <5%
a e 52 mon hs
Na ional and
p i a e
None N/A Buddensiek (1995)
Al mülle & De me
(2006); Al mülle
(2023)
8GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
bo om (s aigh
and ci cula anks
used). Di e en
ank o each yea
class.
Hal we e bu ied in
bes habi a
pa ches and o he
hal eleased in o
open wa e .
Su i al con i med
1 yea la e ; mo e
comp ehensi e
checks o ake
place in 2024
To al was app ox.
15,000 ju eniles
ac oss all coho s
(up o 28 mon hs
old and app ox. 2.5
mm shell leng h)
ju eniles and many
o adul
b oods ock
Dange o ano he
high-sil e en
possible
I eland Coun y Mayo/
Ma ine Ins i u e
M.m. 2020–p esen LI: medium
HF: Salmo sala
(ha che y), Salmo
u a in u u e
MB: adul mussels
kep a acili y o
collec glochidia
IC: no
OC: ju eniles d op
o hos s in o
g a el on he ank
bo om (s aigh
and ci cula anks
used). Di e en
ank o each yea
class
Checked ju eniles in
g a el a bo om o
ank o su i al
and g ow h in
Oc obe and
Decembe 2021.
Some ound and
hey had g own
Wild popula ion
consis s o app ox.
50,000. Plan o pu
some back in o
Newpo Ri e and
pe haps use hese
ju eniles o
pu ing in o o he
i e s, oo. Plan o
be con i med
Na ional None S ill in licensing and
lea ning p ocess
Limi ed expe ience
Ve y s eep lea ning
cu e
De Ey o E. (pe s.
comm., 2022)
I eland Coun y Wa e o d/
Kilmeaden Wa e
T ea men Plan
M.m. 2019–p esen LI: high
HF: Salmo u a (wild
and ha che y in
season 4)
Good encys men on
ish bu ju enile
cap u e le el was
low (none in yea s
Na ional None Cap u e o ju eniles
pos excys men
Ca oll P. (pe s.
comm., 2022)
(Con inues)
GEIST ET AL.15
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License

TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
MB: adul mussels
kep a acili y o
collec glochidia
IC: limi ed success in
cap u ing ju eniles
o da e
OC: none
1 and 2; 50
ju eniles in yea 3)
Poo su i al a es o
hos ish om he
wild
Luxembou g A dennes Ei el
Region/Fonda ion
Hëlle i d'Na u
by na u & ëmwel
M.m. 2008–p esen LI: high
HF: Salmo u a
(ha che y)
MB: glochidia
collec ed in he
ield.
IC: incuba o boxes,
sand aqua ia, and
ough/ lumes
OC: sedimen boxes,
silos, and loa ing
cages on a pond
Ju eniles >3 mm
p oduced/yea :
3,000–4,000
Released a leng h o
10 mm/yea :
1,000–1,500
Ri e Ou s ain
eleased in cages:
1,300 3–4 yea s
old
Ri e Ou s ain
eleased in cages
ali e in 2021: 570
7–9 yea s old.
100% o his
popula ion om
cul u ed ind.
Belgium s ains
eleased: 8,000 3–
4 yea s old
94% o his
popula ion come
om cul u ed ind.
Ge man s ains
eleased: 800 3–4
yea s old
100% o his
popula ion om
cul u ed ind.
Su i al a acili y
un il elease
app ox. 3%
EU LIFE p ojec
and na ional
unding om
LU, BE & DE
PM & JM Keep su i al high
du ing he i s
yea
Adequa e eeding o
0+,1+, and 2+
animals a acili y
Wa e quali y issues
in he s eams in
he a ea
Eybe e al. (2013);
Eybe e al. (2015)
16 GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
Luxembou g A dennes Ei el
Region/Fonda ion
Hëlle i d'Na u
by na u & ëmwel
U.c. 2012–p esen LI: high
HF: Phoxinus
phoxinus (minnow)
caugh in he wild
MB: new mussels
collec ed in he
wild e e y yea
and kep a
b eeding acili y
un il elease o
glochidia. Mussels
e u ned o i e
IC: incuba o boxes,
sand aqua ia, and
ough/ lumes
OC: sedimen boxes,
silos, and loa ing
cages on a pond
Ju eniles >5 mm
p oduced/yea :
2,500–3,000
Released a leng h o
10 mm/yea :
1,000–1,500
Ri e Ou s ain
2,200 > 2 yea s old
eleased
13% o his
popula ion come
om cul u ed ind.
Ri e Saue s ain
3,400 > 2 yea s old
eleased
22% o his
popula ion come
om cul u ed ind.
Su i al a acili y
un il elease
app ox. 5%
EU LIFE p ojec &
na ional unding
om LU & BE
PM Keep su i al high
du ing he i s
yea
Adequa e eeding o
0+and 1+animals
a acili y
Wa e quali y issues
in he s eams in
he a ea
Eybe e al. (2013);
Eybe e al. (2015)
Luxembou g A dennes Ei el
Region/Fonda ion
Hëlle i d'Na u
by na u & ëmwel
U.p. 2020 single
y
LI: medium
HF: Squalius cephalus
caugh in he wild
MB: mussels
collec ed in he
wild and kep a
b eeding acili y
un il elease o
glochidia. Mussels
e u ned o lake
IC: no used
OC: no used
No success in 2020
Almos comple e loss
o in es ed hos
ish
No able o collec
ju enile mussels
du ing single y in
2020
Na ional None Find adequa e hos
ish s ain o his
species
Thielen F. (pe s.
comm., 2022)
(Con inues)
GEIST ET AL.17
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
No way Aus e oll in Ves land
Coun y/Uni e si y
o Be gen
M.m. 2012–p esen LI: high
HF: Salmo sala and
Salmo u a (2+)
b ed om wild
s ains
MB: adul mussels
collec ed in he
ield and kep a
he acili y o
collec glochidia.
Mussels e u ned
o he wild a e 1
yea in qua an ine
IC: incuba o boxes
and ough/ lume
wi h li e algae and
de i us
OC: sedimen boxes
In 2022, 5,400 2+
mussels eleased
di ec ly in o h ee
i e s. In 2021,
14,000 mussels
eleased di ec ly
in o one i e . In
addi ion, 50,000
ecen ly d opped
mussels we e
eleased di ec ly
in o ano he i e
(excess
p oduc ion).
P e ious eleases
o al 6,400 1- o
7-yea -old
ju eniles in o 22
i e s. Releases
in o boxes o he
i s yea s; now
(since 2021) only
eleases in o i e
g a el
Releasing ind. a 4
mm leng h in
boxes is
accep able, bu
ee elease a
la ge sizes is
be e
Ri e s ha ha e no
ec ui men a e
p io i ies. Many
popula ions ha e
e y ew mussels
le . In mos
popula ions,
p opaga ed
ju eniles accoun
o a leas 10–100
Na ional PM & JM Ini ially ge ing
sys ems in place
o high su i al
was a challenge,
bu sys ems now
e ined and no
longe a p oblem
Releasing ju eniles
in o i e s wi h
poo wa e quali y
limi s su i al in
he wild
E alua ing success
Jakobsen P. &
Mage oy J.H. (pe s.
comm., 2022)
18 GEIST ET AL.
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TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
imes he numbe
o wild adul s
No way S o el a Ri e ,
Ting oll
Municipali y, Mø e
og Romsdal
Coun y/
Na u aglige
Konsulen -
jenes e and Fisk
og miljø-
unde søkelse
M.m. 2017–2020 LI: high
HF: Salmo sala (2+)
b ed om wild
s ains
MB: adul mussels
collec ed in he
ield and kep a
he acili y o
collec glochidia.
Mussels e u ned
o he wild a e 1
yea in qua an ine
IC: incuba o boxes
and ough/ lume
wi h li e algae and
de i us
OC: sedimen boxes
This was done as a
p ecau iona y
measu e, oge he
wi h o he
measu es,
associa ed wi h
oad cons uc ion.
The numbe o
mussels eleased
(28) was low
compa ed wi h he
popula ion
es ima e o a leas
3,000 mussels
Regional None Di icul ies wi h
wa e quali y a
he cul i a ion
acili y led o e y
ew mussels being
p oduced
Mage oy J.H. (pe s.
comm., 2022)
Poland Sou h–Cen al
Poland/The
Ins i u e o Na u e
Conse a ion o
he Polish
Academy o
Sciences in K akow
U.c. 2020–p esen LI: high
HF: Co us gobio and
ials wi h Phoxinus
phoxinus and Gobio
gobio (wild)
MB: new mussels
collec ed in he
wild e e y yea
and kep a
b eeding acili y
un il elease o
glochidia. Mussels
e u ned o i e
IC: incuba o boxes,
sand aqua ia, and
ough/ lumes
OC: none
In he i s yea , 30
ish in es ed wi h
la ae om 30
adul mussels
Ju eniles collec ed
app ox. 100 ind.
App ox. 35 10
mon hs old ind. in
ea ly 2022 le
No elease o
mussels ye
50–60 adul mussels
le in Ri e Nida
EU LIFE+P ojec PM Lack o mussel
cul u e expe ience
a he beginning o
p ojec
B eeding acili y long
dis ance om
p ojec i e
No possible o use
i e wa e o
ea ing sys ems
Cages used o keep
in es ed ish in he
Ri e Nida we e
emo ed and
des oyed
Zajac T. (pe s. comm.,
2022)
Po ugal Dis ic Vila Real/
Bo icas Pa que –
Bo icas
municipali y –
M.m. 2019–p esen LI: high
HF: Salmo u a
(ha che y)
Abou 35,000
ju eniles om he
a ge popula ions
( i e s Beça and
Rabaçal) p oduced
Na ional/p i a e PM Time consuming o
ain echnicians as
no p e ious
expe ience wi h
eshwa e mussels
Reis J. (pe s. comm.,
2022)
(Con inues)
GEIST ET AL.19
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
MARE (Uni e si y
o Lisbon)
MB: glochidia
collec ed in he
ield
IC: incuba o boxes
and ough/ lume
OC: in- i e cages
and encys ed ou
in i e s
so a . To da e, no
ju eniles eleased
o i e . Ve y good
su i al and
g ow h in
aceways. 2020,
i s d op o ,
1,500; 2021,
second d op o ,
5,000. T ying o
g ow o aggable
size
Po ugal No he n Po ugal/
Cas elos aqua ic
ea ing acili y
M.m. 2019–2021 LI: high
HF: Salmo u a
(ha che y)
MB: glochidia
collec ed in he
ield
IC: incuba o boxes
and ough/ lume
OC: elease o
encys ed ou in
i e s
Abou 6,500
ju eniles om he
a ge popula ions
( i e s Pai a,
Nei a, and
Rabaçal) p oduced
in ensi ely.
Es ima ed 60,000
d opped o om
ou in aceways.
Encys ed ou also
eleased in i e s
EU & Na ional PM Time consuming o
ain echnicians as
no p e ious
expe ience wi h
eshwa e mussels
High ine sedimen
con en o wa e
leading o high ish
mo ali y
Reis J. (pe s. comm.,
2022)
Po ugal Dis ic Viseu/
Campelo aqua ic
ea ing acili y
managed by
Que cus
M.m. 2013–2016 LI: high
HF: Salmo u a
(ha che y)
MB: glochidia
collec ed in he
ield
IC: incuba o boxes
and ough/ lume
OC: in- i e cages
and sedimen
boxes
O e 225,000
ju eniles om he
a ge popula ion
(Ri e Pai a)
p oduced in 3
yea s (2013–
2015). High
mo ali y caused
by lack o de i us
in closed sys ems.
Good su i al and
g ow h in low-
h ough sys ems
Some mussels
eleased a e 1
yea in cap i i y
(2–4 mm in leng h)
EU LIFE None No sou ce o
de i us collec ion
o eed he
mussels in closed
sys ems
Adminis a i e
cons ain s in
o ganiza ion mean
ollow-up p ojec
no possible
Reis J. (pe s. comm.,
2022)
20 GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License

TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
Po ugal Dis ic Viseu/
Campelo aqua ic
ea ing acili y
managed by
Que cus
U. . 2013–2016 LI: high
HF: Squalius
albu noides and
Squalius a adensis
(ha che y)
MB: collec ion o
glochidia in he
wild
IC: ough/ lume
OC: in- i e cages
Abou 25,000
ju eniles om he
a ge popula ion
(Ri e To gal)
p oduced in 2013
and 5,000 in 2015.
High mo ali y
owing o
inadequa e wa e
quali y (calcium
de iciency) and
need o op imize
die . Ju eniles
could su i e up o
10 weeks and
g ow up o 1 mm
EU LIFE p ojec None No sou ce o
de i us collec ion
o eed he
mussels in closed
sys ems.
Adminis a i e
cons ain s in
o ganiza ion mean
ollow-up p ojec
no possible
Reis J (pe s. comm.,
2022)
Spain Galicia –Lugo/
Uni e si y o
San iago de
Compos ela
M.m. 2012–p esen LI: high
HF: Salmo u a
(ha che y and wild)
and Salmo sala
(ha che y)
MB: adul mussels
kep a acili y o
collec glochidia.
Mussels e u ned
o i e s
IC: incuba o boxes
and suspended
sie es on open-
ci cui aqua ia
supplied by i e
wa e and ed wi h
algae
OC: in- i e cages
(Buddensiek) and
sedimen boxes
Box cul u e: achie ed
a su i al a e
close o 90%
du ing he i s 100
days. Su i al a e
6 mon hs d opped
o app ox. 50%.
Aqua ia: cu en ly,
su i al o 6
mon hs is a ound
90%
Buddensiek cages:
su i al esul s
highly a iable,
su i al a e o
50% in he i s 6
mon hs
P oduc ion:
incuba o boxes,
20,000 (0+yea s);
aqua ia, 30,000
( om 0–2 yea s);
Buddensiek cages,
EU LIFE &
Na ional
PM Pe o m geno yping
o successi e
coho s p io o
elease
Conse a ion s a us
o he species in
he Ulla basin no
clea . Impac o
LIFE p ojec
ac ions no clea
Find sui able elease
si es. Imp o e
b eeding p o ocol
o M.m. ha
imp o es su i al
and g ow h while
being mo e
e icien
Ge long-las ing
unding om
ele an
adminis a ion
Ondina P. & Va ela C.
(pe s. comm.,
2022)
A aujo e al. (2018);
Cas illo e al.
(2020); Cas illo
e al. (2021);
Cas illo e al.
(2022)
(Con inues)
GEIST ET AL.21
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
25,000 (0–8 yea s);
in- i e cages
(sedimen boxes),
300 (7, 8 and 9
yea s old). To al o
75,000 ju eniles
om di e en
coho s, 0–9 yea s
old
In 2017, 25,400
ju eniles a size o
1 mm eleased in
wo s e ches o
he Ulla Ri e
Popula ions in he
wild: Eo Ri e ,
45,500, hope o
elease 200 wi hin
nex 5 yea s;
A nego Ri e ,
20,500, hope o
elease 1,000
wi hin nex 5
yea s; Ulla basin,
11,500, hope o
elease 20,000
wi hin nex 5 yea s
Spain A ag
on/Go e nmen
o A ag
on
P.a. 2001–p esen LI: high
HF: Acipense bae ii
(ha che y) and
Sala ia lu ia ilis
(wild)
MB: new mussels
collec ed in he
wild e e y yea
and kep a
b eeding acili ies
un il elease o
glochidia.
Ju eniles (0+)
collec ed/yea :
2,000,000–
3,000,000 (99%
goes o he Eb o
Ri e and canals;
1% is used o
b eeding in
cap i i y and o
pe o m
eco oxicological
es s)
EU (Feade /
Leade ) &
Na ional
PM one
i e
basin
Mo e
planned
Ques ion on how and
whe e o elease
he cul u ed >50
mm animals needs
o be sol ed
Issues wi h adul
e ili y in he wild
Mo ali y is s ill high
du ing he i s
yea . Space issues
o la ge animals
a he OC acili y
Nakamu a K. (pe s.
comm., 2022)
Nakamu a e al.
(2019)
22 GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
B oods ock
e u ned o i e s
IC: incuba o boxes
and sand aqua ia
OC: ou doo ough/
lume supplied
wi h i e wa e ;
sedimen boxes a
he bo om o he
Eb o Ri e
Ju eniles >1 mm
p oduced/yea :
500–2,000.
Ju eniles 20–50
mm > 1,200 in
2022
Release: ials in
sedimen boxes in
Eb o Ri e wi h
ew ind. (>40 mm)
in summe 2021.
Good esul s
(100% su i al).
Subsequen ly mo e
han 300 ju eniles
>40 mm we e
inco po a ed in
sedimen boxes in
summe 2022.
Once all animals
a e eleased, 20%–
30% in his
popula ion om
cul u ed ind.
Spain Gi ona/Labo a o y
o b eeding naiads
a l'Es any de
Banyoles, managed
by he Conso ci de
l'Es any
U.m. 2010–p esen LI: high
HF: Ba bus
me idionalis (main
use), Lucioba bus
g aellsii,Squalius
laie anus, and
Sala ia lu ia ilis
(wild)
MB: new mussels
collec ed in he
wild e e y yea
and kep a
b eeding acili y
un il elease o
glochidia. Mussels
e u ned o i e s
Te and Flu ià i e
basins (in he i e
and ibu a ies)
Released in es ed
ish: 4,475 ind.
S ocking o eleased
newly excys ed
ju eniles: 87,724
ind.
S ocking o ju eniles
>2 yea s: 3,596
ind.
Banyoles Lake, wa e
in ake s eams and
hei d ainage
channels:
es ima ed
EU LIFE p ojec s &
Na ional
None Adequa e eeding
and au oma ion o
dosage du ing he
i s yea o li e
Technical
imp o emen s in
he eeding and
main enance o
ju eniles du ing
he a ening phase
( om he second
yea o li e)
Implemen a ion o
he in i o cul u e
echnique
Gene ic s udy o
pa en s and
A aujo e al. (2015)
Compos M. (pe s.
comm., 2022)
(Con inues)
GEIST ET AL.23
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
IC: ough/ lume
wi h cages and
sand
OC: pools and
ough/ lume wi h
na u al sand a he
bo om.
Con inuous wa e
eci cula ion and
enewal wi h
na u al wa e om
Lake Banyoles.
Cages placed a
he bo om o he
lake and loa ing
cages
popula ion o
1,000–2,000 ind.
Released in es ed
ish: 3,100 ind.
S ocking o eleased
newly excys ed
ju eniles: 71,651
ind.
S ocking o ju eniles
>2 yea s: 1,267
ind.
An o e all su i al a
acili y o 5% is
es ima ed
cap i e b ed
ju eniles
Spain Gi ona/Labo a o y
o b eeding naiads
a l'Es any de
Banyoles, managed
by he Conso ci de
l'Es any
U. . 2010–p esen LI: high
HF: Ba bus
me idionalis (main
use), Lucioba bus
g aellsii,Squalius
laie anus, and
Sala ia lu ia ilis
(wild)
MB: new mussels
collec ed in he
wild e e y yea
and kep a
b eeding acili y
un il elease o
glochidia. Mussels
e u ned o i e s
IC: ough/ lume
wi h cages and
sand
OC: pools and
ough/ lume wi h
na u al sand a he
bo om, and
con inuous wa e
eci cula ion and
Banyoles Lake, wa e
in ake s eams and
d ainage channels:
Be ween 2010–
2013, a popula ion
o be ween 100–
200 ind. Was
es ima ed.
Released in es ed
ish: 1,910 ind.
S ocking o ju eniles
>2 yea s: 741 ind.
An o e all su i al o
5% is es ima ed o
his acili y
EU LIFE p ojec s &
Na ional
None See Spain U.m. See Spain U.m.
24 GEIST ET AL.
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
TABLE 1 (Con inued)
Coun y Region/ins i u ion Species
Yea s/since
(ending da e)
Le el o in ensi y
and me hod
Rea ing success,
elease, and a io o
wild popula ion age
o size Funding sou ce GM Main challenges Re e ences
En i onmen
Agency
Salmo sala
(ha che y
unsuccess ul)
MB: mussels kep a
ha che y o in es
ish
IC: incuba o boxes
OC: ju eniles mo ed
o low- h ough
ough/ lume
supplied wi h i e
wa e . Release o
a i icially in es ed
ish
glochidia eleased:
825,000. Cu en
li ing ju eniles
om 2017/2018
sp ing/summe
collec ion app ox.
20 > 1.5 cm, 2019
sp ing/summe
collec ion app ox.
200 > 5 mm, 2021
sp ing/summe
collec ion app ox.
80 > 1 mm, 2022
win e collec ion
app ox. 200 < 1
mm. High mo ali y
in each yea g oup,
especially in i s
ew mon hs,
common ob ious
cause o mo ali y
Sap olegnia
in ec ion
Plan o elease
mussels when >15
mm
encys men
densi y)
High mo ali y in
each yea g oup
especially in i s
ew mon hs,
common ob ious
cause o mo ali y:
Sap olegnia
in ec ion (mo ali y
a es >60%)
No e: Me hod le el o in ensi y (LI) is p o ided oge he wi h de ails abou hos ish used (HF), mussel b oods ock si ua ion (MB), indoo cul u e (IC), and ou doo cul u e (OC) sys ems used. GM, gene ic
moni o ing (PM, pa en mussels; JM, ju enile mussels).
Abb e ia ions: A.a., Anodon a ana ina; ind., indi iduals; M.d., Ma ga i i e a du o ensis;M.m., Ma ga i i e a ma ga i i e a;P.a., Pseudunio au icula ius;P.l., Po omida li o alis;U.c., Unio c assus;U.m., Unio mancus;U.p.,
Unio pic o um;U. ., Unio a oisie i;U. ., Unio umidi o mis.
GEIST ET AL.31
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License

place p ima ily ia ideo-con e encing, by phone, o , o a lesse
deg ee, by email. The answe s gi en we e o ma ed in a s anda dized
way, o enable compa ison, and a e p esen ed in Table 1.
3|RESULTS AND DISCUSSION
3.1 |Exis ing ea ing and cul u ing p og ammes o
eshwa e mussels in Eu ope
In his e iew, 46 cap i e ea ing and cul u ing ini ia i es da ing back
o 1989 we e iden i ied (Table 1). These ini ia i es co e 16 coun ies
and 10 species o eshwa e mussels. This includes he endange ed
eshwa e pea l mussel M. ma ga i i e a and he I ish Ma ga i i e a
du o ensis, as well as he gian eshwa e pea l mussel Pseudunio
au icula ius, occu ing only in Spain and F ance. Fi e Unio species ( he
hick-shelled i e mussel Unio c assus, he pain e ’s mussel Unio
pic o um, as well as Unio mancus,Unio a oisie i, and Unio umidi o mis)
and Po omida li o alis ha e also been p opaga ed. Among he pond
mussels (Anodon inae), only Anodon a ana ina has been p opaga ed.
The e o s o cap i e b eeding a e e y une enly dis ibu ed
among hese 10 species, wi h he as majo i y o ac i i ies being
exclusi ely ocused on M. ma ga i i e a, and, o a lesse bu inc easing
ex en , on U. c assus. Some ac i i ies ocus on p opaga ing species in
a e y c i ical si ua ion, such as P. au icula ius,U. mancus, and
U. a oisie i, o mul iple species concu en ly. In p opo ion wi h he
e o s pu in o cap i e b eeding, mos is known abou glochidial
elease imes as well as cul u ing and ea ing p ac ices o he
eshwa e pea l mussel, ollowed by he hick-shelled i e mussel
and he gian eshwa e pea l mussel, whe eas a less is known
abou hese aspec s o all o he species.
In mos cases, exis ing b eeding p og ammes co e he co e a eas
o species dis ibu ion, wi h some excep ions. Fo ins ance, i is
ala ming ha cu en ly only he Spanish cap i e b eeding p og amme
is in place o he highly endange ed P. au icula ius, as p e ious
p og ammes co e ing he emaining co e dis ibu ion a ea o his
species in F ance ha e come o an end. Se e al coun ies ope a e
mul iple b eeding s a ions o he same species o add ess
geog aphical co e age and isk mi iga ion, whe eas some p og ammes
se e popula ions in se e al coun ies. Mos b eeding p og ammes
ha e ecei ed unding h ough ei he na ional o EU LIFE(+) p ojec s
o limi ed du a ion ( ypically a maximum o 5–8 yea s; Table 1).
In M. ma ga i i e a, he species upon which he majo i y o cap i e
b eeding e o s ha e been ocused, he e is al eady e idence ha
some o he eleased cap i e-b ed mussels ha e s a ed o ep oduce,
e.g. in he Ri e Lu e (Ge many). In some s eams, he numbe o
eleased cap i e-b ed mussels al eady exceeds he size o he
emaining na u al popula ion.
3.2 |Di e en in ensi ies o ea ing echniques
Mos ea ing echniques y o b idge he c i ical ju enile s ages in he
li e cycle o mussels (i.e. he pa asi ic and ea ly pos -pa asi ic phases)
by p o iding op imal condi ions du ing his ime. B eeding app oaches
a y g ea ly in he in ensi y, size, species, numbe s o ju eniles
p oduced, and he age/size a which hey a e eleased in o he wild
(Figu e 1; Table 1,). They also di e in moni o ing in ensi y wi h
ega ds o he gene ic moni o ing o he o sp ing, which is in place in
abou hal o he p og ammes. A he highes in ensi y le el, pa en
mussels a e collec ed in he wild and ans e ed in o a ha che y whe e
hey emain ei he pe manen ly (a k- ype sys em) o o ex ended
pe iods o ime. Examples a e he Winde me e s a ion in he UK,
whe e eshwa e pea l mussels om i e i e s a e kep , ypically o
be ween 2 and 36 mon hs, be o e being e u ned o he wild, al hough
one b oods ock popula ion has been p esen a he A k o 13 yea s.
Upon glochidial elease, hos ish, also main ained a he acili y, a e
in es ed, and a e d op-o ju enile mussels a e main ained in a
a ie y o eci cula ing and low- h ough sys ems un il hey each a
size o abou 15–20 mm. Sizes can a y depending on species and
loca ion. Such highly in ensi e sys ems a e usually ee om many o
he ad e se condi ions in mussel i e s, bu also bea a g ea e isk o a
o al loss o mul iple popula ions owing o he agg ega ion o mussels
in one place (e.g. when sys ems ail and sui able eme gency sys ems
a e absen ). These high-in ensi y p og ammes end o be looked upon
less a ou ably by conse a ionis s, unless habi a and ca chmen
es o a ion is being ca ied ou concu en ly wi h he cap i e b eeding
ac i i ies. The eeding o ju eniles is conside ed a majo challenge as
main aining mussels in he ha che y o en equi es he labou -
in ensi e collec ion o de i us om na u al sou ces. In addi ion,
comme cially a ailable algal ood is o en used (Ga enby, Ne es &
Pa ke , 1996), which oge he wi h wa e and de i us om he same
habi a as he pa en mussels, ha e yielded success ul esul s o
ju enile su i al and g ow h.
A he o he ex eme, some p og ammes, such as he eshwa e
pea l mussel conse a ion p og amme in he Ri e Lu e , a e la gely
ield based and independen o echnical acili ies. Each yea a small
numbe o ipe mussels a e collec ed om he i e and used o in es
wild elec o ished o ha che y- ea ed hos ish wi h hose glochidia,
be o e eleasing he in es ed ish in o he i e . This sys em has a low
isk o a ec ing he gene ic cons i u ion o he o sp ing, especially i
di e en pa en mussels a e used each yea , bu i bea s he isk o
being la gely unsuccess ul i s ages o he han he pa asi ic phase
o he li e cycle a e he main bo leneck. This is o en he case wi h
he eshwa e pea l mussel, whe e colma ed and oxygen-de icien
s eam beds a e conside ed he main bo leneck o ec ui men in
Eu ope (Geis & Aue swald, 2007; Denic & Geis , 2015; Simon
e al., 2015), as low oxygen le els e iden ly comp omise he su i al
o ju eniles (Hy ä inen e al., 2022).
The me hods used in any pa icula acili y depend upon ac o s
such as he in as uc u e a ailable, he wa e sou ce, and he amoun
o s a ime and unding a ailable. Whe e specialis acili ies a e
a ailable and he e a e s a o moni o b oods ock, ish, and ju eniles,
high and medium in ensi y me hods, as desc ibed in Figu e 1,canbe
used. Pulsed low- h ough sys ems (Pa e son e al., 2018; Hy ä inen
e al., 2021), incuba o /de i us boxes (Eybe e al., 2013; Schede
e al., 2014; Nakamu a e al., 2018; G unicke e al., 2023), lumes,
aqua ia sys ems (La ic oi e e al., 2016; La ic oi e e al., 2020), and he
32 GEIST ET AL.
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Floa ing Upwelle Sys em (FLUPSYS) (Pa e son e al., 2018) all equi e
medium o high le els o a en ion. The mos esou ce-in ensi e
sys ems pe ju enile ea ed a e he incuba o /box and he pulsed
low- h ough sys ems (Kunz e al., 2020; Hy ä inen e al., 2021). In he
incuba o sys em, ju eniles a e kep in s a ic wa e in boxes (wi h
usually 200–500 indi iduals pe box) and a e cleaned and supplied
wi h esh wa e , ood, and de i us one o wo imes pe week. The
pulsed low- h ough sys em p o ides wa e (and ood) changes a
egula in e als (as much as once pe hou ), bu he e a e usually no
mo e han 50–100 ju eniles pe beake . The lowe esou ce-in ensi e
sys ems a e he downwelling aqua ium sys em (wi h 1000 ju eniles
pe sie e, and con aining 12–15 sie es, cleaned e e y 2 weeks), he
sand aqua ia (wi h 150–500 ju eniles pe sys em, and a weekly wa e
exchange), FLUPSYS (wi h se e al hund ed o 1000 ju eniles pe
bucke , and cleaned when necessa y), and lumes (wi h se e al
housand indi iduals pe lume, and cleaned when necessa y).
The lowe in ensi y and mo e ield-based sys ems (boxes 6-10,
Figu e 1) can be used when space in specialis acili ies is limi ed o
absen . The smalles ju eniles ( eshly excys ed om ish) can be
placed in o in- i e cage sys ems. These sys ems, placed in mussel
i e s, s ill need egula cleaning and main enance, bu a e ela i ely
low e o in ela ion o acili y-based sys ems. Ju eniles o >4 mm
can be placed in o mussel silos, again wi h egula cleaning (mon hly),
whe eas sligh ly la ge ju eniles (>10 mm) can be placed in sedimen
boxes (Bílý e al., 2018) o in o side channels (o en e e ed o as
H uˇ
ska channels; H uska, 1999; H uska, 2001; Gum, Lange &
Geis , 2011), as a so - elease me hod be o e being s ocked in o inal
elease si es. Sedimen boxes need some cleaning and main enance,
bu he side-channel me hod equi es almos no main enance.
3.3 |Challenges associa ed wi h mussel cap i e
b eeding p og ammes
The p ima y challenges iden i ied in he majo i y o cap i e b eeding
p og ammes a e ela ed o limi a ions in budge o s a ing, and o
ea ing condi ions such as wa e quali y, eeding, and he su i al o
ju eniles du ing ea ly li e s ages (Table 1). Conce ning budge
limi a ions, he sho du a ion o unding schemes o e only a ew
yea s, especially o long-li ed species such as M. ma ga i i e a,isa
mo e se ious cons ain han he unding le el when s a ing new
p og ammes. As M. ma ga i i e a only become ma u e a an age o
10–15 yea s (Young & Williams, 1984), he ypical unding schemes,
wi h a maximum du a ion o 3–6 yea s in his species, only co e he
ea ing o a ew coho s o sexually imma u e ju eniles. Fu he mo e,
unce ain y in he con inua ion o unding dec eases he e en ion o
expe ienced s a and is ou o s ep wi h he biological pace o some
eshwa e mussel species. F equen ly, se ing up and main aining
FIGURE 1 Cap i e b eeding me hods used o eshwa e mussels h oughou Eu ope. Colou ed do s on he map e e o he di e en
echniques desc ibed in he no e boxes. Boxes 1 and 2 desc ibe high-in ensi y me hods, equi ing main enance a leas once pe week; boxes 3–5
desc ibe medium-in ensi y me hods, equi ing main enance a leas once pe o nigh ; and boxes 6–10 desc ibe low-in ensi y me hods, equi ing
main enance app oxima ely mon hly, o as needed
GEIST ET AL.33
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b eeding p og ammes can also be complica ed by wa e quali y issues,
he p o ision o na u al o pu chased eed o ea ly li e s ages, and
high mo ali y a es, some imes culmina ing in he loss o an en i e
yea coho . Rea ing sys ems dependen on an ex e nal wa e supply
especially ace a g ea e isk o un o eseen wa e quali y issues, such
as cyanobac e ia blooms (No wegian b eeding s a ion Aus e oll),
excess u bidi y, and ine sedimen loading (Aus ian b eeding s a ion),
as well as wa e quan i y issues (as obse ed in d y summe s in he
Ge man b eeding s a ion nea Ho ). A la e s ages o he mussel li e
cycle, he iden i ica ion and limi ed a ailabili y o habi a o su icien
quali y in o which he ju eniles can be eleased a e epo ed as
addi ional challenges (Table 1). In addi ion o he challenges di ec ly
associa ed wi h he cap i e b eeding o mussels, moni o ing hem
a e hei elease is ano he key issue. Such moni o ing is essen ial
o assessing he sui abili y o ecipien wa e bodies o cap i e-b ed
ju eniles, iden i ying he ideal elease si es, and de e mining he ideal
cap i e b eeding p ocedu es ha esul in he g ea es su i al in he
wild.
3.4 |Recommenda ions o he u u e
Despi e p og ess in he cap i e b eeding o eshwa e mussels, hei
sus ainable conse a ion will always depend on he conse a ion o
es o a ion o habi a s and ca chmen s. While conse a ion and
es o a ion p ojec s con inue, cap i e b eeding p og ammes can help
o sa e small popula ions om ex inc ion and boos he numbe o
indi iduals o e en ual elease back in o he wild once he habi a s
a e capable o suppo ing ea ly-s age ju enile mussels. Based on
in o ma ion om cu en eshwa e mussel b eeding s a ions in
Eu ope, he ollowing measu es a e sugges ed o imp o emen .
3.4.1 | P o ision o species- and basin-wide
Eu opean conse a ion s a egies o eshwa e mussel
species wi h long- e m unding commi men s
The p o ision o long- e m unding op ions o secu e acqui ed
speci ic knowledge and con inuous ac ion o long-li ed mussel
species is c ucial and key o success. The auna discussed he e a e
long-li ed, slow-g owing species, and mussel ca chmen s a e so
signi ican ly deg aded ha in mos cases habi a es o a ion may
ake mo e han a decade. Also, om a gene ic poin o iew,
con inued cap i e b eeding o e mul iple gene a ions can help o
p e en he e osion o gene ic di e si y (Geis e al., 2021). Owing
o he dominance o unding o cap i e b eeding p o ided by he
Eu opean Union LIFE/LIFE+p og ammes and na ional unds wi h
limi ed unning imes o 5 yea s o less, easy op ions o p ojec
ex ensions ollowing an objec i e and independen e iew a
in e media e s ages would be mos welcome and use ul. Species-
and basin-wide Eu opean conse a ion s a egies o eshwa e
mussels will ensu e ha p og ammes ha e he bes chance o
success.
3.4.2 | Science- and e idence-based suppo in he
de elopmen and e alua ion o b eeding p og ammes
A signi ican olume o wo k has been ca ied ou by b eeding
p og ammes o imp o e e iciencies and inc ease he numbe o
ju eniles su i ing in cap i i y. Howe e , much o his aluable wo k
does no ge published in he p ima y li e a u e, limi ing i s impac .
Al hough he e is a s ong and collabo a i e Eu opean ne wo k o
mussel b eeding p og ammes, mo e science- and e idence-based
suppo (wi h he subsequen publica ion and dissemina ion o
esul s) is needed o d i e as e paced posi i e ou comes. The
de elopmen o b eeding p og ammes equi es he in eg a ion o
gene ic in o ma ion, as demons a ed o he eshwa e pea l mussel
(Geis e al., 2021), as well as a c i ical e alua ion o he impac s o
cap i e b eeding p ocedu es on he p ogeny, and ul ima ely on he
long- e m success o di e en ea ing me hods in aiding he e-
es ablishmen o unc ional popula ions in he wild.
3.4.3 | Inc eased knowledge exchange and aining
oppo uni ies
Al hough he di e si y o di e en app oaches o mussel b eeding is
gene ally use ul, by inc easing he o e all sys em esilience and
minimizing he high losses o ju enile mussels, e.g. ela ed o eeding,
hese issues could be educed by an inc eased le el o in o ma ion
exchange and aining. The e a e o en e y limi ed o no unds o
exchange and aining oppo uni ies. The e o e, he e is a need o
p o ide ans-Eu opean unding, which allows scien i ic and p ac ical
exchange among exis ing and newly planned b eeding p og ammes. In
he pas , mos o he exchange among b eeding s a ions has
happened h ough local con e ences held, o example, wi hin exis ing
LIFE(+) p ojec s, hus being spo adic in ime and place. Wi hin he
Eu opean COST p ojec CONFREMUS (2023), an in ensi ied
exchange among mussel expe s and mussel b eede s has been
ini ia ed o he i s ime, albei ime-limi ed o when his p ojec
ends in 2024. Con inua ion and expansion (whe e app op ia e) o
such success ul ini ia i es is impo an o ensu ing a well-connec ed
ne wo k o expe s who communica e new indings in a imely
manne and who a e a ailable o aining hose new o mussel
conse a ion.
3.4.4 | Coo dina ed ac ion ela ed o species and
geog aphical ep esen a ion as well as gene ic aspec s
Despi e he success s o ies o many cap i e b eeding e o s a he
Eu opean scale, he e is s ill no coo dina ed app oach among hem in
e ms o species, geog aphical, and gene ic ep esen a ion. The
de elopmen o a coo dina ed Eu opean s a egy o cap i e mussel
b eeding, ideally in eg a ed in o habi a and ca chmen es o a ion
plans –all o which should be e idence-based and suppo ed by
scien i ic esea ch –would be needed o inc ease he e ec i eness o
34 GEIST ET AL.
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such p og ammes. This should ideally be in eg a ed in o he
de elopmen o minimal s anda ds o cap i e mussel b eeding,
conce ning animal wel a e aspec s o hos ish use, disease
p e en ion, main aining gene ic in eg i y o cap i e-b ed mussels, and
minimizing ex inc ion isk, as well as equi emen s o acking
and assessing he success o mussel eleases. I may also be necessa y
o p io i ize conse a ion ac ions (Geis , 2015) o ensu e he bes
emaining occupied si es wi hin each ca chmen a e secu ed, a he
han sp eading conse a ion ac ion oo hinly o a emp o conse e
ma ginal popula ions and isk ailu e owing o a lack o esou ces.
3.4.5 | Guidelines o assessmen o success o he
b eeding ac ion and isk mi iga ion
Toge he wi h he p oduc ion o coun y-wide/Eu opean
conse a ion s a egies, he p oduc ion o guidelines o assess
b eeding p og amme success and how o mi iga e isks would be
aluable o new p og ammes. Guidelines would also help o
s anda dize he way in which cu en b eeding p og ammes
communica e hei ou pu s, d i ing an inc eased awa eness o ac o s
a ec ing success and how isks can be minimized. In u u e,
in o ma ion on which species, numbe s, and sizes/ages o mussels a e
being s ocked a which loca ions needs o be collec ed mo e
sys ema ically and beyond he le el o indi idual s a ions. The
success ul de elopmen and implemen a ion o a s anda d app oach
o moni o ing eshwa e pea l mussel (M. ma ga i i e a) popula ions
in Eu opean i e s (Boon e al., 2019) and he con inuing de elopmen
o a new Eu opean Commi ee o S anda diza ion (CEN) s anda d on
mussel moni o ing o a wide ange o species illus a e ha such
ac ion is possible, e en wi h limi ed le els o unding. This pa allels
s a egic conse a ion app oaches in No h Ame ica, whe e a na ional
s a egy o he conse a ion o na i e eshwa e molluscs has been
de eloped (FMCS, 2016).
CONFLICT OF INTEREST
The au ho s ha e no con lic s o in e es associa ed wi h his wo k.
AUTHOR CONTRIBUTIONS
JG, LL, and FT join ly led he s udy, conduc ed and analysed he
in e iews, and a e he p incipal au ho s. Concep ualiza ion: JG, LL,
and FT. Me hodology: JG, LL, and FT. Da a cu a ion: all au ho s.
Visualiza ion: LL, FT, and RH. In es iga ion: JG, LL, and FT. W i ing—
o iginal d a : JG. W i ing— e iew and edi ing: all au ho s. All au ho s
ha e ead and ag eed o he published e sion o he a icle.
AFFILIATIONS
1
Aqua ic Sys ems Biology Uni , TUM School o Li e Sciences,
Technical Uni e si y o Munich, F eising, Ge many
2
Fonda ion Hëlle i d'Na u /na u & ëmwel , Ma nach, Luxembou g
3
F eshwa e Biological Associa ion, YMCA No h Campus, Newby
B idge, UK
4
Lachendo , Ge many
5
Uni . Bo deaux, CNRS, Bo deaux INP, Pessac, F ance
6
B e agne Vi an e –Socié é pou l’
E ude e la P o ec ion de la
Na u e en B e agne (SEPNB), B es , F ance
7
F eshwa e Mussel B eeding Labo a o y o Lake Banyoles, Banyoles,
Spain
8
Wildli e Conse a ion Medicine Resea ch G oup (WildCoM),
Depa amen de Medicina i Ci u gia Animals, Uni e si a Au ònoma
de Ba celona, Ca alonia, Spain
9
F iend o he Ri e Clodiagh G oup, Kilmeaden Wa e Plan ,
Kilmeaden, I eland
10
Consul an s in Aqua ic Ecology and Enginee ing –bla isch e.U.,
Wels, Aus ia
11
Bund Na u schu z Ho , Ho , Ge many
12
Hanno e , Ge many
13
Landscha sp lege e band Passau e.V., Passau, Ge many
14
Fédé a ion de Pêche du Finis è e, Piscicul u e du Fa o , B aspa s,
F ance
15
Ma ine Ins i u e, Fu nace, Newpo , Co. Mayo, I eland
16
Ins i u e o Hyd obiology, Technical Uni e si y o D esden,
D esden, Ge many
17
Depa men o Bioscience, Uni e si y o Be gen, Be gen, No way
18
Ins i u e o Ag icul u al and En i onmen al Sciences, Es onian
Uni e si y o Li e Sciences, Ta u, Es onia
19
S a e Fo es Managemen Cen e, Haljala Municipali y, Lääne-Vi u
Coun y, Es onia
20
No wegian Ins i u e o Na u e Resea ch, Oslo, No way
21
T ini y Cen e o he En i onmen , School o Na u al Sciences,
Depa men o Zoology, T ini y College Dublin, The Uni e si y o
Dublin, Dublin, I eland
22
Public Se ice o Wallonia, Ag icul u e, Na u al Resou ces and he
En i onmen , DEMNA, Di ec o a e o Na u e and Wa e , Gembloux,
Belgium
23
En i onmen al Se ice Depa men , Sociedad A agonesa de Ges i
on
Ag oambien al (SARGA), Za agoza, Spain
24
Ca anilles Ins i u e o Biodi e si y and E olu iona y Biology,
Uni e si y o Valencia, Valencia, Spain
25
Depa men o Zoology, Gene ics and Physical An h opology,
Uni e si y o San iago de Compos ela, Campus Te a, Lugo, Spain
26
Biology, Ins i u ion o En i onmen al and Li e Sciences, Ka ls ad
Uni e si y, Ka ls ad, Sweden
27
BIVALVIA s. .o., Zábˇ
eh, Czech Republic
28
Ma ine and En i onmen al Sciences Cen e (MARE)/Aqua ic
Resea ch Ne wo k (ARNET), Faculdade de Ciências da Uni e sidade
de Lisboa, Lisbon, Po ugal
29
The Ru al Economy and Ag icul u al Socie y, Eldsbe ga, Sweden
30
Lüsslingen, Swi ze land
31
Biologische S a ion S äd e Region Aachen e.V., S olbe g, Ge many
32
Uni e si é de Tou s, Cen e Na ional de la Reche che Scien i ique
(CNRS) l’UMR 7324 CI és, TER i oi es, En i onnemen e Socié és
(CITERES), Tou s, F ance
33
Depa men o Biological and En i onmen al Science, Uni e si y o
Jy äskylä, Jy äskylä, Finland
34
Cyn ig Fish Cul u e Uni , Llan yanch, UK
GEIST ET AL.35
10990755, 0, Downloaded om h ps://onlinelib a y.wiley.com/doi/10.1002/aqc.4018 by Uni e si y O Jy äskylä Lib a y, Wiley Online Lib a y on [02/10/2023]. See he Te ms and Condi ions (h ps://onlinelib a y.wiley.com/ e ms-and-condi ions) on Wiley Online Lib a y o ules o use; OA a icles a e go e ned by he applicable C ea i e Commons License
35
Kielde Salmon Cen e, En i onmen Agency, No humbe land, UK
36
Swedish Angle s Associa ion, Gö ebo g, Sweden
37
Ins i u e o Na u e Conse a ion, Polish Academy o Sciences,
K ak
ow, Poland
ACKNOWLEDGEMENTS
This publica ion is based upon wo k om COST Ac ion CA18239,
suppo ed by COST (Eu opean Coope a ion in Science and
Technology). We also hank F. Jecke, M. Koes e , M. Pagel, and
E. Zelenko á o suppo ing his s udy. Open Access unding enabled
and o ganized by P ojek DEAL.
DATA AVAILABILITY STATEMENT
The da a ha suppo he indings o his s udy a e a ailable om he
co esponding au ho , upon easonable eques .
ORCID
Jue gen Geis h ps://o cid.o g/0000-0001-7698-3443
F ankie Thielen h ps://o cid.o g/0000-0001-8137-3877
Louise La ic oi e h ps://o cid.o g/0000-0002-9751-8505
Magalie Baud imon h ps://o cid.o g/0000-0001-9901-8435
Jon Hamne Mage oy h ps://o cid.o g/0000-0002-5027-4398
Keiko Nakamu a h ps://o cid.o g/0000-0001-5037-3516
Paz Ondina h ps://o cid.o g/0000-0003-4392-9250
Ma in Ös e ling h ps://o cid.o g/0000-0001-6758-5857
Joaquim Reis h ps://o cid.o g/0000-0002-6404-0875
Joaquín Sole h ps://o cid.o g/0000-0002-1606-6675
Jouni Taskinen h ps://o cid.o g/0000-0003-0098-9560
Ben S achan h ps://o cid.o g/0009-0001-6154-3696
Tadeusz Zająch ps://o cid.o g/0000-0003-2048-9205
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