SHORT COMMUNICATION
Dese pumpkinseed: die composi ion and b ead h
in a Mo occan i e
Ch is os Gkenas
1,*
, Ma ia Filomena Magalhães
2
, Naza e Campos-Ma in
1
,
Filipe Ribei o
1
and Miguel Cla e o
3
1
MARE, Cen o de Ciências do Ma e do Ambien e, Faculdade de Ciências, Uni e sidade de Lisboa, Campo G ande,
Lisboa, Po ugal
2
cE3c, Cen o de Ecologia, E olução e Al e ações Ambien ais, Faculdade de Ciências, Uni e sidade de Lisboa, Lisboa, Po ugal
3
Depa amen o de Biologia de la Conse acion, Es acion Biologica de Don~ana CSIC, Se illa, Spain
Recei ed: 16 May 2021 / Accep ed: 28 Sep embe 2021
Abs ac –The widely in asi e No h Ame ican pumpkinseed sunfish, Lepomis gibbosus, is cu en ly
es ablished in dese i e s in Mo occo. The success o pumpkinseed in no el ecosys ems has been
associa ed wi h i s gene alis die , bu his ai emains une alua ed in a id egions. Dese i e s a e ha sh
en i onmen s wi h limi ed wa e and p ey a ailabili y which may ad e sely cons ain he die o fish. He e
we s udied he die o pumpkinseed ac oss 4 si es in he D aa Ri e , emb acing a 450 m ele a ional g adien
co e ing om ex emely d y lowlands o ela i ely humid highlands. We desc ibed pumpkinseed die
h ough he analysis o s omach con en s o 82 indi iduals, collec ed in he all o 2013. Pumpkinseed die
was domina ed by Chi onomidae in d y lowlands, while Epheme op e a, He e op e a, T ichop e a and
Odona a we e ele an p ey in mo e humid highlands. Popula ion die b ead h expanded wi h ele a ion, bu
indi idual specializa ion in die was low among all si es. Ou esul s highligh conside able changes in die
composi ion and b ead h wi h a idi y, sugges ing ha eeding plas ici y and use o exclusi e, locally
abundan p ey a he han gene alized eeding may be associa ed wi h he success o pumpkinseed in mos
a id a eas in dese i e s.
Keywo ds: In asi e species / Lepomis gibbosus / a idi y / ele a ion / eeding ecology
Résumé –La pe che-soleil du dése : composi ion e la geu de niche ophique dans une i iè e
Ma ocaine. La la gemen en ahissan e pe che-soleil No d-Amé icaine, Lepomis gibbosus, es é ablie
aujou d’hui dans des i iè es du dése au Ma oc. Le succès de la pe che-soleil dans les nou eaux
écosys èmes a é é associé à son égime alimen ai e géné alis e, mais cela es e non é alué dans les égions
a ides. Les i iè es du dése son des en i onnemen s hos iles a ec peu d’eau e de p oies disponibles ce qui
con ain le égime alimen ai e des poissons. Nous é udions le égime ophique de la pe che-soleil su
qua e si es de la i iè e D aa, le long d’un déni elé de 450 m allan des plaines ex êmemen a ides aux
hau eu s ela i emen humides. Nous a ons déc i le égime alimen ai e de la pe che-soleil pa l’analyse de
l’es omac de 82 indi idus, cap u és à l’au omne 2013. Le égime alimen ai e de la pe che-soleil é ai dominé
pa les Chi onomidae dans les plaines a ides alo s que les Ephémé op è es, Hé é op è es, T ichop è es e
Odona es é aien des p oies impo an es dans les hau eu s plus humides. La la geu de la niche ophique
s’acc oissai a ec l’al i ude mais la spécialisa ion indi iduelle é ai aible en e ous les si es. Nos ésul a s
me en en é idence un changemen considé able dans la composi ion e la la geu de la niche ophique en
onc ion de l’a idi é suggé an que la plas ici é alimen ai e e l’u ilisa ion exclusi e de p oies localemen
abondan es plu ô que l’alimen a ion géné alisée peu ê e associée au succès de la pe che-soleil dans les
zones les plus a ides des i iè es du dése .
Mo s clés : Espèce en ahissan e / Lepomis gibbosus / a idi é / al i ude / Écologie alimen ai e
*Co esponding au ho : [email p o ec ed]
Knowl. Manag. Aqua . Ecosys . 2021, 422, 34
©C. Gkenas e al., Published by EDP Sciences 2021
h ps://doi.o g/10.1051/kmae/2021033
Knowledge &
Managemen o
A
qua ic
Ecosys ems
www.kmae-jou nal.o g
Jou nal ully suppo ed by O ice
ançais de la biodi e si é
This is an Open Access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License CC-BY-ND (h ps://c ea i ecommons.o g/licenses/by-nd/4.0/), which pe mi s un es ic ed 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. I you emix, ans o m, o build upon he ma e ial, you may no dis ibu e he modified ma e ial.
The pumpkinseed sunfish, Lepomis gibbosus (Linnaeus,
1758), is na i e o he G ea Lakes egion in eas e n No h
Ame ica (Sco and C ossman, 1973) bu has been success ully
in oduced o o e 30 coun ies in Cen al and Sou h Ame ica,
Eu ope, No h A ica, and Ana olia Peninsula (Ya no e al.,
2020). Pumpkinseed has been classified as an equilib ium
s a egis , bu i can be e y plas ic in i s na i e ange and some
non-na i e popula ions exhibi oppo unis ic ai s ha appea
o con ibu e o i s in asi e success (Fox e al., 2007).
Non-na i e popula ions inhabi a wide a ie y o eshwa e
sys ems, and dese i e s in he a id egions o Mo occo
cu en ly cons i u e he limi s o i s dis ibu ion ange. Se e al
s udies on he die o non-na i e popula ions in Eu ope ha e
epo ed mac oin e eb a e and zooplank on consump ion
a ying wi h p ey a ailabili y and habi a cha ac e is ics
(Godinho and Fe ei a, 1998;Van Klee e al., 2008;Gkenas
e al., 2019), bu li le is known abou he die o pumpkinseed
in dese i e s in a id egions.
A id egions a e cha ac e ized by low and unce ain
p ecipi a ion, highly fluc ua ing empe a u es, and wa e
sca ci y (B i o e al., 2014), which may ha e a c i ical
influence on p ey a ailabili y o fish in dese i e s
(Balcombe e al., 2005). Dese i e s a e highly dynamic
sys ems in which d y pe iods, equen ly in ol ing he
cessa ion o supe ficial flow, al e na e wi h iolen flash
floods, p opo iona ely mo e in ense and mo e unp edic able
han in o he i e s (Ha ms e al., 2008). In gene al, aqua ic
mac oin e eb a e abundance and di e si y dec ease wi h
a idi y, due o low p ima y p oduc i i y and habi a complexi y
(Bunn e al., 2003). Fish na i e o dese i e s ha e a guably
de eloped eeding s a egies o cope wi h hese ha sh and
highly fluc ua ing condi ions, bu his may no be he case o
non-na i e species which ha e e ol ed in mo e s able
en i onmen s. The e o e, i is impo an o cla i y whe he
success ul in ade s such as pumpkinseed become mo e o a
gene alis o a he mo e o a specialis in die wi h a idi y.
He e, we analyzed he die o pumpkinseed ac oss ou
si es in he D aa Ri e in No he n Mo occo, which emb ace an
ele a ional g adien co e ing om ela i ely humid highlands
o ex emely d y lowlands. Specifically, we analyzed pa e ns
in he composi ion o he die and in he popula ion and
indi idual die b ead h along his ele a ional g adien .
The D aa Ri e has been desc ibed in de ail by Cla e o
e al. (2015). In b ie , he clima e in he basin is a id wi h
empe a u e anging om 3 °C o34°C, and annual p ecipi a-
ion om 50 o 465 mm (h ps://www.wo ldclim.o g/)
(Fick and Hijmans, 2017). The D aa Ri e flows o
1100 km om he High A las o he A lan ic Ocean bu is
usually d y in he las 600 km. Pumpkinseed we e sampled in
ou si es, in he El Mansou Eddahbi Dam (D1), and 56.4 km
(D2), 88.9km (D3), and 135.6 km (D4) downs eam. To a oid
he pe iod o flash floods, sampling was ca ied ou in
Sep embe and Oc obe 2013. Du ing his pe iod, p ecipi a ion
and empe a u e anged om 19 mm and 19.9–23.8 °CinD1
(1106 m a.s.l.) o 8 mm and 22.8–26.9 °C in D4 (674 m a.s.l).
Wa e conduc i i y a ied be ween 1318 mS/cm and 6274 mS/
cm a D1 and D4, espec i ely, eflec ing a downs eam
inc ease in salini y. Sampling was conduc ed wi h yke ne s in
D1 and by elec ofishing (50–75 di ec pulses s
1
) in he
emaining si es, and pumpkinseed we e eu hanized wi h an
o e dose o clo e oil and p ese ed in 4% o maldehyde.
We sampled and examined he s omach con en s o
85 pumpkinseed, anging om 40 o 80 mm s anda d leng h.
We ound 3 s omachs wi h less han 3 p eys which we e
conside ed as emp y and disca ded o a oid con ounding
e ec s in he analysis o die s uc u e (Bowen, 1996). The
emaining s omachs we e simila ly ull among si es, as de i ed
om isual es ima ion o pe cen olume ulness anges. P ey
i ems in each s omach we e iden ified o he lowes
ecognizable axon and coun ed.
Fo analysis, p ey we e g ouped in o en ca ego ies, wi h
“o he ”including bo h a e and uniden ified p ey i ems (Tab. 1).
P ey ca ego ies accumula ion cu es based on andom addi ion o
s omachs and 1000 pe mu a ions (Go elli and Colwell, 2001)
eached an asymp o e in all si es sugges ing ha he numbe o
s omachs used we e adequa e o desc ibe local die s (Supplemen-
a yFig.S1).Wedesc ibeddie composi iona eachsi eusing wo
con en ional indices, he equency o occu ence (F%) which is
hep opo iono non-emp ys omachscon ainingapa icula p ey
ca ego y, and he nume ical equency (N%), which is he
Table 1. Va ia ion in he nume ic equency (N%) and in he equency o occu ence (F%; in pa en heses) o p ey ca ego ies in he die o
pumpkinseed Lepomis gibbosus (40–80 mm SL) in he D aa i e in No he n Mo occo. P ey ca ego ies wi h nume ic equency >10% a e
highligh ed in bold. Si es a e o de ed by inc easing a idi y and hus by dec easing ele a ion.
P ey ca ego ies O e all D1 D2 D3 D4
Chi onomidae 53.0 (97.6) 29.2 (100.0) 41.1 (100.0) 70.8 (100.0) 82.4 (90.5)
Epheme op e a 20.3 (69.5) 22.1 (100.0) 43.1 (94.7) 14.3 (81.0) 0.6 (4.8)
Odona a 6.8 (56.1) 11.7 (76.2) 8.1 (57.9) 3.3 (61.9) 1.9 (28.6)
He e op e a 7.0 (45.1) 16.6 (81.0) 3.6 (71.4) 1.9 (23.8)
T ichop e a 5.5 (41.5) 13.7 (90.5) 1.4 (15.8) 0.9 (28.6) 2.5 (28.6)
Cladoce a 1.2 (17.1) 2.2 (28.6) 0.5 (10.5) 2.8 (28.6)
Ce a opogonidae 1.1 (15.9) 3.3 (31.6) 1.5 (33.3)
Copepoda 0.7 (11.0) 1.4 (19.0) 0.2 (5.3) 0.3 (9.5) 0.6 (9.5)
Decapoda 0.2 (6.1) 1.5 (23.8)
O he p ey 4.0 (53.7) 2.9 (42.9) 2.2 (42.1) 5.4 (71.4) 5.9 (57.1)
To al p ey 2257 759 418 756 324
To al fish 82 21 19 21 21
Page 2 o 6
C. Gkenas e al.: Knowl. Manag. Aqua . Ecosys . 2021, 422, 34
p opo ional coun o each p ey ca ego y ela i e o he o al p ey
coun among fish (Hyslop, 1980). We analyzed a ia ion in die
composi ion and b ead h among si es using p ey nume ical
equency o each fish.
To summa ize die s uc u e among si es, we pe o med a
Non-Me ic Mul idimensional Scaling (nMDS), using he
B ay-Cu is dissimila i y coe ficien (Cla ke e al., 2014). To
educe he influence o abundan p ey in he analysis, da a we e
p e iously squa e oo ans o med. We in e p e ed he fi s
wo o dina ion axes, wi h s ess alues <0.2 (Cla ke e al.,
2014). Analysis o Simila i y (ANOSIM) was used o es o
di e ences be ween si es, wi h o e all significance calcula ed
using 9999 pe mu a ions o he da ase and co ec ed o
mul iple es ing using he Bon e oni sequen ial me hod.
Simila i y Pe cen age (SIMPER) analysis was used o iden i y
he p ey ca ego ies wi h he highes con ibu ion o die
dissimila i y. P ey ca ego ies we e o de ed by hei a e age
con ibu ion o he o al a e age dissimila i y, wi h a cu -o a
50% o cumula i e a e age dissimila i y (Cla ke e al., 2014).
We quan ified die b ead h a he popula ion and indi idual
le els. Popula ion b ead h was de e mined using he Shannon-
Wiene index, which is maximized when popula ions use mo e
p ey, mo e e enly (Colwell and Fu uyma, 1971). Indi idual
b ead h was de e mined using he index o Indi idual
Specializa ion, which equals 1 when all indi iduals consume
he ull se o p ey iden ified a he popula ion-le el and
declines owa ds ze o when indi iduals use smalle subse s o
he popula ion p ey (Bolnick e al., 2002). Es ima es o bo h
indices we e ob ained using boo s apped 95% confidence
in e als (9999 pe mu a ions).
We used linea eg ession analysis o es o a ia ion in
die composi ion and b ead h wi h ele a ion. We used he mean
loca ion (i.e., cen oids) om he nMDS sco es in he fi s s wo
axes and he mean Shannon-Wiene and Indi idual Speciali-
za ion o fish in each si e as esponse a iables in he analysis.
All analyses we e conduc ed using he R so wa e ( .4.0.0,
R Co e Team, 2020), and significance o s a is ical es ing was
assessed a P<0.05.
In o al, we analyzed 82 s omachs and 2257 p ey i ems,
wi h sample sizes pe si e a ying be ween 19 and 21 s omachs
and 324–759 p ey (Tab. 1). Chi onomidae we e ound in 98%
o he s omachs and made up 53% o he o al p ey.
Epheme op e a we e also o en ound (70%) bu in low
abundances (20%), while Odona a, He e op e a and T ichop-
e a occu ed less equen ly in he s omachs (42–56%) and in
small numbe s (<10%). These esul s a e coinciden wi h
Fig. 1. Resul s o Non-Me ic Mul idimensional Scaling (nMDS) o dina ion o p ey nume ic equency in he die o pumpkinseed Lepomis
gibbosus in he D aa i e in No he n Mo occo. a) nMDS biplo , wi h each si e delimi ed by minimum con ex polygons; b and c) box plo s
illus a ing a ia ion in he fi s and second nMDS sco es o each si e in ela ion o ele a ion, espec i ely; boxes ep esen he in e qua ile
ange (IQR; 25 h and 75 h pe cen iles), lines wi hin boxes a e he median, whiske s ep esen he 75 h pe cen ile 1.5 xIQR and he 25 h
pe cen ile þ1.5 xIQR, and do s beyond he whiske s a e ou lie s. S ess alue o o dina ion axes is indica ed in panel a).
Page 3 o 6
C. Gkenas e al.: Knowl. Manag. Aqua . Ecosys . 2021, 422, 34
hose ound in Eu opean i e s (Godinho and Fe ei a, 1998;
Almeida e al., 2009;Gkenas e al., 2019) and ponds
(Van Klee e al., 2008), sugges ing ha mac oin e eb a es
may cons i u e s aple p ey o pumpkinseed ac oss i s non-
na i e ange. Con e sely, only small numbe s o Cladoce a
(1%) and Copepoda (0.7%) we e seldom ound in pumpkin-
seed s omachs (<20%), bu hese ha e been ound o be
ele an p ey o some non-na i e pumpkinseed popula ions in
Lake Banyoles in Spain (Ga cia-Be hou and Mo eno-Amich,
2000).
Die composi ion a ied conside ably among si es (Tab. 1).
Chi onomidae made up only 30% o he o al p ey in D1 bu
inc eased in D2 (41%) and domina ed he die in D3 (71%) and
D4 (82%). Addi ional impo an p ey in D1 we e He e op e a
(17%), T ichop e a (14%) and Odona a (12%). Epheme op e a
made up 43% o he o al p ey in D2 and we e s ill ele an in
D1 (22%) bu no in D3 (14%) and in D4 (5%). The nMDS
biplo highligh ed conside able die s uc u e despi e he e was
some o e lap in sco es among si es (Fig. 1a). The ANOSIM
e ealed significan a ia ion be ween all si es, wi h he
s onge sepa a ion in die (A D >60%) occu ing be ween
D4 and D1 and D2 (Tab. 2). Dissimila i ies in die we e
consis en ly ela ed o con ibu ions o Chi onomidae, wi h
Epheme op e a being also impo an o he di e en ia ion o
D4, and He e op e a and T ichop e a o he di e en ia ion o
D1. Va ia ion in nMDS1 sco es showed a posi i e associa ion
wi h ele a ion (F
1,2
= 25.07, R
2
= 0.89, p= 0.037; Fig. 1b,)
bu no significan end was ound o nMDS2 (F
1,2
= 0.040,
R
2
=–0.47, p= 0.860; Fig. 1c). Spa ial changes in die ha e
been commonly ound in non-na i e popula ions o pumpkin-
seed eflec ing o aging plas ici y and changes in local p ey
supply (Almeida e al., 2009;Gkenas e al., 2016). This may
also be he case in he D aa Ri e , whe e he ele a ional
g adien in die composi ion p obably eflec s changes in
ichness and abundance o mac oin e eb a es, and he
dec ease o Epheme op e a, He e op e a and T ichop e a
and dominance o ole an Chi onomidae unde mo e d y
and sal y condi ions (Mo eno e al., 2010;Colombe i e al.,
2020). I is also possible ha a ia ion in die can a leas
pa ially be associa ed wi h changes in p ey a ailabili y unde
mo e s able en i onmen al condi ions p e ailing in he dam
(D1), as ound in o he s udies (Almeida e al., 2009).
Popula ion-le el die b ead h inc eased wi h ele a ion
(F
1,3
= 43.89, R
2
= 0.93, p= 0.022; Fig. 2a) bu no significan
Table 2. Resul s o he ANOSIM and SIMPER analyses o p ey
nume ic equency in he die o pumpkinseed Lepomis gibbosus in
he D aa i e in No he n Mo occo. As e isks indica e significan
di e ences in pai wise compa isons based on he sequen ial
Bon e oni me hod (a= 0.05, k= 6). P ey ca ego ies con ibu ing
>20% o a e age dissimila i y (A D) a e highligh ed in bold and
anks o p ey con ibu ions a e shown in pa en heses. P ey codes a e:
CHI, Ch i onomidae; EPH, Epheme op e a; HET, He e op e a; TRI,
T ichop e a.
Si es R
0.440*
A D CHI EPH HET TRI
D1–D2 0.475* 48.11 16.14 (3) 19.86 (1) 18.86 (2)
D1–D3 0.388* 48.35 23.89 (1) 14.03 (3) 16.32 (2)
D1–D4 0.651* 65.86 16.85 (2) 22.93 (1) 15.42 (3)
D2–D3 0.306* 45.63 29.53 (1) 20.55 (2)
D2–D4 0.512* 60.53 21.78 (2) 35.71 (1)
D3–D4 0.332* 54.93 30.27 (1) 21.54 (2)
Fig. 2. Va ia ion in die b ea h o pumpkinseed Lepomis gibbosus in
he D aa i e in No he n Mo occo in ela ion o ele a ion. a)
Popula ion b ead h de e mined om he Shannon-Wiene index; b)
Indi idual b ea h de e mined om he Indi idual Specializa ion
index. Ma ke s and e o ba s a e he mean and he boo s apped 95%
confidence in e als o each index, espec i ely.
Page 4 o 6
C. Gkenas e al.: Knowl. Manag. Aqua . Ecosys . 2021, 422, 34
end was ound o indi idual-le el b ead h (F
1,2
=4.42,
R
2
=0.53,p=0.170;Fig. 2b) which anged be ween 0.65 and
0.74. The alues o popula ion die b ead h a e close o
figu es ound in sou he n Eu ope, namely in he Bullaque
and Es ena i e s in Spain (Almeida e al.,2009) and in he
A dila Ri e in Po ugal (Gkenas e al.,2019). Likewise,
alues o indi idual die b ead h we e wi hin he ange
epo ed o he Guadiana Ri e in Po ugal (Gkenas e al.,
2016). The ends owa ds na owe die s and use o simila
se s o p ey by all indi iduals in d ye lowlands may eflec
condi ions o p ey limi a ion associa ed o d ough as ound
elsewhe e (Magalhães, 1993). Fu he mo e, he use o
exclusi e, likely abundan p ey, may also be associa ed o
changes in fish assemblages ac oss he ele a ional g adien
(see Cla e o e al.,2015), wi h low die b ead h a bo h he
popula ion and indi idual le els a o ing p ey pa i ioning
among species.
Ou esul s indica e ha pumpkinseed may display a
na owe die wi h a idi y. I would be impo an o u he
unde s and how eeding pa e ns a e influenced by p ey
a ailabili y and in e specific ophic in e ac ions and how
hese a ec he sp ead o pumpkinseed in a id en i onmen s
which a e expec ed o expand unde u u e wa me and d ye
clima es (see e e ences in B i o e al., 2014). In eg a i e
s udies dealing wi h pumpkinseed popula ion ai s and habi a
cha ac e is ics addi ional o die , should hus be encou aged in
dese i e s.
Supplemen a y Ma e ial
Figu e S1. P ey ca ego ies accumula ion cu es o pumpkin-
seed Lepomis gibbosus in he D aa i e in No he n Mo occo,
based on andom addi ion o s omachs and he a e age o 1000
pe mu a ions.
The Supplemen a y Ma e ial is a ailable a h ps://www.kmae-
jou nal.o g/10.1051/kmae/2021033/olm.
Acknowledgemen s. This wo k was unded by he Na ional
Geog aphic Socie y’s Commi ee o Resea ch and Explo a-
ion h ough g an #9188-12 and by he EBD-CSIC h ough a
mic o-p ojec wi hin he Spanish Se e o Ochoa P og am
(SEV-2012-0262), also benefi ing by he suppo o Land
Ro e Jagua Espan~a. Pe missions we e ob ained om he
Hau Commissa ia aux Eaux e Fo e^ s e ala Lu e Con e la
Dése ifica ion du Royaume du Ma oc. We g ea ly acknow-
ledge he company and field assis ance p o ided by Raulo
A oyo, Ca li Pe ez, I ia So o, Fali Bece a, Ma iangeles
Ma inez Panke, Pepa Bo e o and Manu Pe ez. Addi ional
suppo was p o ided by he Fundação pa a a Cie^ncia e
Tecnologia (FCT) h ough he s a egic p ojec UID/MAR/
04292/2019 g an ed o MARE and he pos -doc g an s o
Ch is os Gkenas (SFRH/ BPD/ 84859/2012) and o Filipe
Ribei o (SFRH/BPD/46761/2008) om he P og ama
Ope acional Po encial Humano/Fundo Social Eu opeu
(POPH/FSE).
Re e ences
Almeida D, Almodo a A, Nicola GG, El i a B. 2009. Feeding ac ics
and body condi ion o wo in oduced popula ions o pumpkinseed
Lepomis gibbosus: aking ad an ages o human dis u bances? Ecol
F eshw Fish 18: 15–23.
Balcombe SR, Bunn SE, McKenzie-Smi h FJ, Da ies PE. 2005.
Va iabili y o fish die s be ween d y and flood pe iods in an a id
zone floodplain i e . J Fish Biol 67: 1552–1567.
Bolnick DI, Yang LH, Fo dyce JA, Da is JM, S anba
ck R. 2002.
Measu ing indi idual-le el esou ce specializa ion. Ecology 83:
2936–2941.
Bowen SH. 1996. Quan i a i e desc ip ion o he die . In Mu phy BR,
Willis DW, eds. Fishe ies echniques. Be hesda, MD: Ame ican
Fishe ies Socie y, pp. 513–532.
B i o JC, Godinho R, Ma inez-F ei iaF,e al. 2014. Un a elling
biodi e si y, e olu ion, and h ea s o conse a ion in he Saha a-
Sahel. Biol Re 89: 215–231.
Bunn SE, Da ies PM, Winning M. 2003. Sou ces o o ganic ca bon
suppo ing he ood web o an a id zone floodplain i e . F eshw
Biol 48: 1–17.
Cla ke KR, Go ley RN, Some field PJ, Wa wick RM. 2014.
Change in ma ine communi ies: An app oach o s a is ical
analysis and in e p e a ion, 3 d edn. Plymou h: P ime -E L d,
256 p.
Cla e o M, Esqui ias J, Qninba A, e al. 2015. Fish in ading dese s:
non-na i e species in a id Mo occan i e s. Aqua Conse Ma
F eshw Ecosys 25: 49–60.
Colombe i P, Calde on M, Tello J, González P, Jo é M. 2020.
Rela ionships be ween aqua ic in e eb a e assemblages and
en i onmen al quali y on saline we lands o an a id en i onmen .
J A id En i on 181: 104245.
Colwell RK, Fu uyma DJ. 1971. On he measu emen o niche b ead h
and o e lap. Ecology 52: 567–576.
Fick SE, Hijmans RJ. 2017. Wo ldclim 2: new 1-km spa ial
esolu ion clima e su aces o global land a eas. In J Clima ol
37: 4302–4315.
Fox MG, Vila-Gispe A, Copp GH. 2007. Li e his o y ai s o
in oduced Ibe ian pumpkinseed (Lepomis gibbosus) ela i e o
na i e popula ions: can di e ences explain coloniza ion success?
J Fish Biol 71: 56–69.
Ga cia-Be hou E, Mo eno-Amich R. 2000. Food o in oduced
pumpkinseed sunfish: on ogene ic die shi and seasonal a ia ion.
J Fish Biol 57: 29–40.
Godinho FN, Fe ei a MT. 1998. Spa ial a ia ion in die composi ion
o pumpkinseed sunfish, Lepomis gibbosus, and la gemou h bass,
Mic op e us salmoides, om a Po uguese s eam. Folia Zool 47:
205–213.
Gkenas C, Magalha~es MF, Cuche ousse J, Domingos I, Ribei o F.
2016. Long e m pa e ns in he la e summe die a y niche o he
in asi e pumpkinseed sunfish Lepomis gibbosus.Knowl Manag
Aqua Ecosys 417: 1–5.
Gkenas C, Magalhães MF, Cuche ousse J, O juela RL, Ribei o F.
2019. Die a y niche di e gence be ween wo in asi e fish
in Medi e anean s eams. Knowl Manag Aqua Ecosys 420:
1–10.
Go elli NJ, Colwell RK. 2001. Quan i ying biodi e si y: p ocedu es
and pi alls in he measu emen and compa ison o species ichness.
Ecol Le 4: 379–391.
Ha ms TK, Sponselle RA, G imm NB. 2008. Dese s eams. In
Jø gensen SE, Fa h BD, eds. Ecosys ems. Encyclopedia o
Ecology. Ox o d: Else ie , pp. 871–879.
Hyslop EJ. 1980. S omach con en s analysis a e iew o me hods
and hei applica ion. J Fish Biol 17: 411–429.
Magalhães MF. 1993. Feeding o an Ibe ian s eam cyp inid
assemblage—seasonali y o esou ce use in a highly a iable
en i onmen . Oecologia 96: 253–260.
Page 5 o 6
C. Gkenas e al.: Knowl. Manag. Aqua . Ecosys . 2021, 422, 34

Mo eno JL, Angele DG, De las He as J. 2010. Seasonal dynamics
o mac oin e eb a e communi ies in a semia id saline sp ing
s eam wi h con as ing en i onmen al condi ions. Aqua Ecol 44:
177–193.
R De elopmen Co e Team. 2020. R: A language and en i onmen o
s a is ical compu ing. Vienna, Aus ia: R Founda ion o S a is ical
Compu ing. h ps://www.R-p ojec .o g/.
Sco WB, C ossman EJ. 1973. F eshwa e fishes o Canada. Bulle in
Fishe ies Resea ch Boa d o Canada, O awa, 966 p.
Van Klee HH, Van de Velde G, Leu en RSE. 2008. Pumpkinseed
sunfish (Lepomis gibbosus) in asions acili a ed by in oduc-
ions and na u e managemen s ongly educe mac oin e e-
b a e abundance in isola ed wa e bodies. Biol In asions 10:
1481–1490.
Ya no S, Gobin J, Wilson CW, Vila-Gispe A, Copp GH, Fox MG.
2020. New and Old Wo ld phylogeog aphy o pumpkinseed
(Lepomis gibbosus): he No h Ame ican o igin o in oduced
popula ions in Eu ope. Hyd obiologia 847: 345–364.
Ci e his a icle as: Gkenas C, Magalhães MF, Campos-Ma in N, Ribei o F, Cla e o M. 2021. Dese pumpkinseed: die composi ion and
b ead h in a Mo occan i e . Knowl. Manag. Aqua . Ecosys ., 422, 34.
Page 6 o 6
C. Gkenas e al.: Knowl. Manag. Aqua . Ecosys . 2021, 422, 34