Desert pumpkinseed: diet composition and breadth in a Moroccan river

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. 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