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Preimaginal feeding habits of Dictyogenus fontium (Plecoptera, Perlodidae) in an alpine brook in NW Italy PDF

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Preview Preimaginal feeding habits of Dictyogenus fontium (Plecoptera, Perlodidae) in an alpine brook in NW Italy

©EntomologicaFennica.15March2007 Preimaginal feeding habits of Dictyogenus fontium (Plecoptera, Perlodidae) in an alpine brook in NW Italy StefanoFenoglio*,TizianoBoandGiorgioMalacarne Fenoglio,S.,Bo,T.&Malacarne,G.2007:PreimaginalfeedinghabitsofDictyo- genusfontium(Plecoptera,Perlodidae)inanalpinebrookinNWItaly.—Ento- mol.Fennica18:27–31. ThefeedinghabitsofDictyogenusfontiumnymphswereinvestigatedintheRio Olen, an alpine brook in NW Italy. In an analysis of the gut contents of 84 nymphs,wedetectedevidenttrophicpreferences:Chironomidae(Diptera)were themajorcomponentofthepreyingested,independentlyoftheiravailabilityin thesubstratum.Plantdetritusandalgaewerealsoanimportantpartofthedietof thiscarnivorousstonefly.Thefindingsarediscussedonthebasisofecological considerations. S.Fenoglio,T.Bo&G.Malacarne,UniversityofPiemonteOrientale,Di.S.A.V., Via Bellini n. 25, 15100 Alessandria, Italy; *corresponding author’s e-mail: [email protected] Received5December2005,accepted16March2006 1.Introduction Plecoptera nymphs are the most abundant predators in many lotic systems. In particular, Benthiccommunitiesinrunningwaterarecom- PlecopteraSystellognathaisthedominantpreda- posedalmostexclusivelyofinsects:theyconsti- tor group in fishless alpine streams, with a key tute an estimated 70–90% of the macro- ecological role as top-down control elements in invertebratespeciesinstreamsandrivers(Min- benthic communities (Zanetell & Peckarsky shall1969).Plecopteraisoneofthemostimpor- 1996,Wipfli&Gregovich2002).Peckarskyand tantordersintheseecosystems,indeedoftenthe Dodson(1980)analysedthecolonisationprocess dominantorder(Zwick2000, Elliott2004). For ofinstreamartificialsubstrataandfoundthatsub- this reason, many studies have investigated dif- stratacontainingpredaceousstoneflieswereless ferent aspects of the ecology and life-history of colonised, confirming that these predators can thisgroup(TiernodeFigueroa&Fochetti2001, significantlyinfluencebenthiccommunitystruc- Stewart&Stark2002). ture. Optimal foraging theory, as formulated by Inrecentyears,therehasbeenincreasingin- Krebs(1978),predictsthatpredatorsoptforthe terestinstoneflyfoodhabits.Sincethepioneer- most advantageous prey according to different ing study of Richardson and Gaufin (1971), re- factors,suchasencounterrate,preydensity,en- search has been conducted on food selection ergy contents, handling time and others. Fuller (Feminella&Stewart1986),ontogeneticshiftsin andStewart(1977,1979)investigatedfoodhab- the diet (Bo & Fenoglio 2005) and functional its of many Plecoptera species, quantifying grouping(Short&Ward1981–feedingmecha- electivityandfoodpreferencesofstoneflylarvae nisms, according to theRCC, River Continuum inrelationtofoodavailability:theydemonstrated Concept,Vannoteetal.1980). theexistenceofseasonalandontogeneticshifts, 28 Fenoglioetal. (cid:127) ENTOMOL.FENNICAVol.18 dietary overlaps,daily periodicity andprey size Table1.Mainabioticparametersinthesamplingsta- electivity. Predaceous stoneflies exhibit a great tion,RioOlenbrook(NWItaly). variety of food habits: some species seemto be Temperature(°C) 4.8 extremely selective, feeding mainly on certain pH 7.7 prey(Fenoglio&Bo2004),whileotherspecies Conductivity(µS/cm) 78.0 aremoreopportunistic,eatingthemostabundant Alkalinity(mgHCO–/l) 26.0 organisms in the benthic community (Allan & Cl–(mg/l) 3 0.5 Flecker1988,Dudgeon2000).Moreover,studies SO––(mg/l) 4.2 PO4–––(mg/l) <0.1 haveshownthatseveralSystellognathahavevar- NO4–(mg/l) 0.6 iedfoodhabits,feedingnotonlyonlivinganimal NO3–(mg/l) <0.01 prey but also on plant detritus (Lillehammer NH2+(mg/l) <0.4 4 1988).Forthisreason,StewartandStark(2002) stated that some stonefly species would not fit intoanyofthestereotypedcategoriesdefinedby size of 500 µm) early in the morning, because dominant food or feeding mechanisms, e.g. the Systellognatha are considered mainly nocturnal fivefunctionalfeedinggroupssensuRCC(scrap- feeders (Vaught & Stewart 1974). We also col- ers, shredders, predators, filterers and collector- lectedSurbersamplesinthesameriffletoassess gatherers). Hence, the food habits of unstudied thepresenceandabundanceoftaxainthenatural species cannot beinferred fromtheir phylogen- benthic invertebrate population. Samples were eticposition. preserved in 90% ethanol. In the laboratory, all Dictyogenusfontium(Ris,1896)isatypical organismswerecountedandidentifiedtothege- orophilous species, endemic to the Alps (Con- nuslevel,exceptforLumbriculidaeandearlyin- siglio1980,Ravizza&Ravizza-Dematteis1991, stars of some Diptera, which were identified to Fochetti 1995, Brittain & Saltveit 1996). As its thefamilylevel. specific name suggests, this mountain stonefly The total length of D. fontium nymphs was livesinsmallspringsandbrooksataltitudesrang- measured (0.1 mm accuracy) and the nymphs ingfrom1,000to2,600ma.s.l.,whereitisoften were processed to assess food consumption by theonlyPerlodidae(Ravizza&Ravizza-Demat- means of gut contents analysis. Guts were re- teis1990). moved and the contents of the alimentary canal Theaimofthisstudywastoanalysethedietof were analysed by the transparency method for D. fontium nymphs in a typical alpine environ- slides.Fragmentsofanimalpreywereidentified ment. tothelowesttaxonomiclevelpossible.Identifica- tionofpreywasbasedonsclerotizedbodyparts, particularly head capsules, mouthparts and leg 2.Materialandmethods fragments. Stewart and Stark (2002) stated that thecountofsclerotizedfragments(i.e.headcap- On 29–30 September 2005, D. fontium nymphs sules) can give a reasonably accurate count of werecollectedina100muniformriffleoftheRio preyconsumed,andweightsorvolumescouldbe Olen brook (UTM 414652–5079947; 2,100 m extrapolated, if necessary, from morphometric a.s.l.). The sampling station is a typical alpine weight-volumerelationshipsofprey.Toinvesti- lotic environment, characterized by high slope, gatepossiblefeedingpreferences,wecompared coarseriverbedandfastflowingwater.Riparian thegutcontentswiththenaturalcompositionand vegetation is almost absent, except for some abundanceofmacroinvertebratecommunitiesin Sorbusaucuparia.Thisloticsystemshowsgood theriverbedusingthetrophicelectivityindexof environmentalquality,correspondingtoanenvi- Ivlev(1961): ronmentwithouttracesofhuman-inducedalter- ation(firstclassintheItalianExtendedBioticIn- E=(ri–pi)/(ri+pi) dex, Ghetti 1997). The main abiotic parameters arereportedinTable1.Samplingswerecarried whereri=theproportionofingestedspeciesand out with a kick-net sampler (20 x 20 cm; mesh pi=therelativeabundanceinthebenthiccommu- ENTOMOL.FENNICAVol.18 (cid:127) PreimaginalfeedingofDictyogenusfontium 29 nity.Feedingpreferenceswerealsoquantifiedus- Table2.Relativeabundance(%valueinthecommu- inganotherelectivityindex(McCormick1991): nity)ofmacroinvertebratescollectedinthenatural riverbed. E*=(W–1/N)/W+1/N i i Taxa Relative FFG abundance(%) (*) where Plecoptera W =(r /p)/Sr /p Leuctrasp. 3.68 Sh i i i i i Nemourasp. 4.61 Sh Protonemurasp. 2.63 Sh In this formula, r = the proportion of ingested i Chloroperlasusemicheli 3.16 P species,p =therelativeabundanceinthebenthic i Siphonoperlamontana 1.58 P community, and N = the number of food items. Isoperlasp. 1.05 P Bothindexesrangefrom–1.0to1.0.Avalueof– Dictyogenusfontium 5.05 P 1.0 means total avoidance, 1.0 indicates prefer- Ephemeroptera Ecdyonurussp. 8.95 Sc enceand0indicatesindifference.Thepresenceof Rhithrogenasp. 1.58 Sc algae and detritus (e.g. fragments of terrestrial Epeorusalpicola 2.11 Sc vegetation) was recorded and quantified on a Baetismelanonyx 5.79 Cg scaleof0–3(0=nopresence;3=highestabun- Baetissp. 2.63 Cg danceclass). Trichoptera Rhyacophilasp. 11.32 P Hyporhyacophilasp. 2.11 P Drusussp. 5.26 Sh 3.Results Monocentralepidoptera 2.63 Sh Cryptothrixnebulicola 2.63 Sh Weexaminedthegutcontentsof84Dictyogenus Diptera Chironomidae 11.58 Cg fontium nymphs and we collected 760 macro- Limoniidae 2.11 P invertebratesbelongingto26taxa.Thetaxalist Empididae 1.05 P andrelativemacroinvertebrateabundanceinthe Simuliidae 8.11 F riverbedarereportedinTable2.Themeanlength Lispesp. 1.45 P ofD.fontiumimmaturestageswas12.9mm,with Prionocerasp. 0.53 Sh Oligochaeta a minimumof 8.0 mmand a maximumof 17.5 Lumbriculidae 3.68 Cg mm. Eiseniellatetraedra 0.53 Cg Duringthelaboratoryanalysis,wefoundfour Crustacea Niphargussp. 0.13 Cg Platyhelminthes Crenobiaalpina 2.50 P Arachnida Hydracarina 1.58 P (*)FFG:f unctionalfeedinggroups(Cg=collector-gatherers;F= filterers;P=predators;Sc=scrapers;Sh=shredders.seeMerrittand Cummins1996). completelyemptyguts.Fineplantfragmentsand algaewerepresentin82%oftheexaminedguts. Furthermore,plantdetrituswasvirtuallytheonly element found in five nymphs; interestingly, these five nymphs were of very different size, from10.4to17.5mm(totallength). Resultsofthefeedingpreferenceanalysisare Fig.1.Electivityindexesformacroinvertebratetaxain thedietofDictyogenusfontiumnymphsinRioOlen reportedinFig.1.Consideringallspecimenscol- brook. lected,themostimportantpreygroupwasChiro- 30 Fenoglioetal. (cid:127) ENTOMOL.FENNICAVol.18 nomidae(Diptera): it constituted 68.8% oftotal theRioOlenbrook,thetwoindexesrevealedthat ingesteditemsandwaspresentin65.5%ofguts. theirpresenceinthedietwasrelatedtoactivese- Ephemeroptera,mainlyBaetidae,wasthesecond lection and preference (Ivlev index = 0.79, mostfrequentgroup,occurringin9.5%ofguts. McCormick index = 0.76). This consistent pat- Intwospecimens,wefoundheadsandotherbody ternofelectionofChironomidaeandavoidance partsofLimnephilidae(Trichoptera)andinone ofotherabundantorganismscouldberelatedto gut there was an entire specimen of Hyporhya- ecologicalandbehaviouralfactors;forexample, cophila sp. (Trichoptera). Plecoptera legs and Simuliidae could be difficult to catch because fragmentswerepresentinonlyonegut.Interest- they inhabit microhabitats difficult to explore ingly,someorganismsthatwerequiteabundant (uppersurfaceofstonesinfastflowingenviron- inthenaturalenvironment,suchasRhyacophila ments). On the other hand, the relatively abun- sp.,Hyporhyacophilasp.(Trichoptera)andPro- dant caddisflies of the family Rhyacophilidae simuliumsp.(Diptera),werealmostabsentinthe maybeadifficultpreybecauseoftheirlargesize dietofD.fontium. andpredaceoushabits. In conclusion, this study provides evidence thatthedietofD.fontiuminalpineenvironments 4.Discussion isnotstrictlycarnivorousandischaracterisedby trophicselectionforafewselectedprey. Several studies have examined food habits in stoneflies (Fuller & Stewart 1979, Peckarsky Acknowledgements.WethankPaoloAgostaandMassimo 1980,Allan&Flecker1988).Theproblemofan Pessino for help during field sampling. We are also in- debtedtoPeterZwickandananonymousrefereefortheir accurate assessment of feeding habits and food constructivecomments. electivity has been pointed out in many papers and summarized in the work of Peckarsky (1984): the exclusion of particular prey items References fromthepredator’sdietcouldberelatedtosev- eralcauses,suchaseffectivepreyescapemecha- Allan, J. D. & Flecker, A. S. 1988: Prey preference in nisms, low prey availability or lack of habitat stoneflies:acomparativeanalysisofpreyvulnerabil- overlapbetweenpreyandpredator. ity.—Oecologia76:495–503. 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