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Population Ecology and Functioning of Enchytraeidae in Some Arable

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Population Ecology and Functioning of Enchytraeidae in Some Arable Population ecology and functioning of Enchytraeidae insom e arable farming systems CENTRALE LANDBOUWCATALOGUS 0000 0459 4293 Promotoren: dr. L. Brussaard hoogleraar in de bodembiologie dr. ir. L.J. Pons emeritus hoogleraar ind e regionale bodemkunde 1 tJtióï'lc } N Wim Didden Population ecology and functioning of Enchytraeidae insom e arable farming systems Proefschrift terverkrijgin gva nd egraa dva n doctor ind elandbouw - enmilieuwetenschappen , opgeza gva nd erecto rmagnificus , dr.H.C .va nde rPlas , inhe topenbaa rt everdedige n opwoensda g8 me i199 1 desnamiddag st evie r uuri nd eAul a vand eLandbouwuniversitei tt eWageningen . BIBLIOTHEEK tÄNDBOUWUNIVERSriEflfr WAGENINGEN Cover photograph: Wim van Hof ,oüo^io^ |4)M 1. Het belang van potwormenvoo r nutriëntenstromen in (agro)oecosystemen wordt nietallee n bepaald door biomassae n turnover van de populaties, maar tevens door hunvertical everdelin g over het profiel en temporele dynamiek. Dit proefschrift 2. De indirecte effecten van potwormen op de nutriëntenmobilisatie kunnen aanzienlijk zijn als gevolg van de vermoedelijk lage selectiviteit bij de voedselopname en de hoge selectiviteitwaarme evoedse l wordtgeassimileerd . Abrahamsen, G.,199 0- Biol .Fertil . Soils 9:159-162 Dit proefschrift 3. Potwormen dragen aanmerkelijk bijaa n de dynamiekva n de bodemstructuur en kunnen zodoende een belangrijke rol spelen bijd e lucht- en vochthuishouding van de bodem. Dit proefschrift 4. De fysisch-chemische parameters die ind e meeste oecologische studies gemeten worden, zijnvoo r hetvoorspelle nva n de aan te treffen hoeveelheden potwormen van geringewaarde . Dit proefschrift 5. Laboratoriumgegevens over de levenscyclusva n dieren zijn slechts dan bruikbaar voorvoorspellinge n van develdsituatie ,wannee r ze tevens bepaald zijn onder daar optredende extreme omstandigheden en wanneer veldmetingen op de relevante schaal worden verricht. Dit proefschrift 6. Ruimtelijke heterogeniteit op micro- en macroschaalvorm t de basisvoo r het functioneren van het bodemoecosysteem en tegelijkertijd een van de belangrijkste hinderpalenvoo r bodembiologisch onderzoek. 7. Het gebruik van functionele groepen inee n oecologische analyse, zonder soort- gericht oecologisch onderzoek naar de in deze groepen geplaatste organismen, leidt onvoldoende tot inzicht in hetfunctionere n van oecosystemen. Hunt,H.W . etal. , 19B7 -Biol . Fertil. Soils 3:57-68 Hetfei tda t oecologischonderzoe k aan potwormen slechts relatief iaat is aangepakt, ondanks hetzee r talrijkvoorkome nva n deze dieren, doet vermoeden dat de ontwikkelingva n de wetenschap medeword t bepaald door angstvoo r het onbekende. 9. De huidige praktijkva n voornamelijk kortlopende contractenvoo r wetenschappelijk onderzoek bevordert zowel de produktiviteit als deverschralin g van dat onderzoek. 10. 'No-nonsense beleid'wijs t niet alleen op een povere beheersing van de Nederlandse taal, maar ook op een duidelijke minachting voor afwijkende meningen. 11. Veel bodembiologisch onderzoek blijft aan de oppervlakte. Stellingen behorende bij het proefschrift van Wim Didden: Population ecologyand functioningof EnchytraekJae In some arable fanningsystems. Wageningen, 8 mei 1991 Abstract Didden,W.A.M. ,199 1- Populatio necolog yan dfunctionin g of Enchytraeids insom e arablefarmin gsystem s• Doctora lthesis , Agricultural University,Wageningen , The Netherlands Thepopulatio necolog yo f Enchytraeidae,an dthei rfunctioning ,wer estudie di n somearabl efarmin gsystem san di nlaborator yexperiments . Thesystem sstudie d consistedo fa 'conventional ' (highinpu to fenerg yan dmatter ) andsevera l'integrated ' (reducedinpu to fenerg yan dmatter ) systems. Emphasiswa sgive nt opopulatio n dynamicsan dth erol eo f Enchytraeidae innutrien tcyclin gan dsoi lstructur eevolution . Meanyearl yenchytraei dabundanc ean dbiomas si nth esystem sstudie drange d from1100 0ind/m 2t o4300 0ind/m 2, andfro m0.0 8g C/m 2t o0.4 2g C/m 2,respectively . Itwa scalculate dtha t0.5-3.7 %o fth eyearl yorgani ccarbo ninpu twa srespire d byth e enchytraeid populations. Yearlynitroge nflu xfro mth epopulation s (throughstorag ei n enchytraeidtissue ) rangedfro m0.1 9t o0.6 0g/m 2. Therewer en osignifican t differences betweenth esystem sstudie da sregard s populationdynamica lan dproductio necologica lparameters . However,vertica ldistribu ­ tiono fenchytraei d populationsdiffere d markedlybetwee nth econventiona lfiel dan dth e integratedfields ,probabl yrelate dwit hdifference s insoi ltillag ean dth eresultin gdistri ­ butiono forgani cfertilize r andplan tresidues . Thesedifference s invertica ldistributio n mayhav econsequence sfo rth eris ko f nutrientsleachin gfro mth esystems . Theimpac to fenchytraei d activityo nsoi lstructur eevolutio nwa sstudie d ina fiel d experimentwit hartificia lsoi lcores ,i ncombinatio nwit hfiel dobservations . Itwa sfoun d thatenchytraei dactivit yproduce d measurableeffect s onai rpermeability , poresiz e distribution,an ddistributio n ofaggregat esizes ,probabl ythroug hselectiv eburrowin g andtranspor to f organicbeside s mineralmaterial . Life-history parameterso fEnchytraeus buchholzi, thedominan tspecie sa tth e researchsite ,wer edetermine di na laborator yexperiment ,an dprediction s basedo n thelaborator ytrial swer ecompare dwit hdat ao npopulatio ndevelopmen tunde rmor e naturalconditions . Therewer esignifican tdiscrepancies ,indicatin ga ninfluenc eo fth e physiologicalconditio no fth eexperimenta lanimal so nth elengt ho fth elife-cycl ei nthi s species. Keywords :Enchytraeida e- Soi lfaun a- Agro-ecosystem s -Populatio ndynamic s Productionecolog y- Nutrien tcyclin g- Soi lstructur e- Life-history Contents 1. General introduction 1 1.1. Outlineo fthi sthesi s 5 2. Ecologyo fterrestria lEnchytraeida e 7 2.1. Introduction 7 2.2. Factorsinfluencin g populationsize so fEnchytraeida e 8 2.2.1. Introduction 8 2.2.2. Abioticfactor s 12 2.2.2.1. Moisture 12 2.2.2.2. Temperature 13 2.2.2.3. pH 14 2.2.3. Bioticfactor s 15 2.2.3.1. Foodan dfeedin g 15 2.2.3.2. Competition 17 2.2.3.3. Predatorsan dparasite s 18 2.2.4. Maninduce dfactor s 20 2.2.4.1. Physical habitatchange s 20 2.2.4.2. Chemicals 21 2.2.4.2.1. Pesticides 21 2.2.4.2.2. Fertilizers 24 2.2.4.2.3. Pollution 25 2.3. Therol eo f Enchytraeidae inecosyste mfunctionin g 27 2.3.1. Introduction 27 2.3.2. Soilstructur e 27 2.3.3. Microdistribution of Enchytraeidae 28 2.3.4. Populationdynamic s 29 2.3.5. Production ecology 31 2.4. Concluding remarks 35 3. Populationdynamic san d productionecolog yo f Enchytraeidae inhig han dreduce dinpu tfarmin gsystem s 36 3.1. Introduction 36 3.2. Methodsan dmaterial s 37 3.2.1. Theexperimenta lsit e 37 3.2.2. Sampling procedure 38 3.2.3. Analysiso fth edat a 39 3.2.4. Productionecologica lcalculation s 39 3.3. Results 41 3.3.1. Compositiono fth eenchytraei dfaun a 41 3.3.2. Verticaldistributio nan drelatio nt oplan tpositio n 42 3.3.3. Thecanonica lcorrespondenc eanalyse s 44 3.3.4. Relationsbetwee nbod yweigh tan dlengt h 46 3.3.5. Population dynamics 46 3.3.6. Respirationan dproductio n 49 3.3.7. Nitrogenturnove r 50 3.4. Discussion 52 4. Life-history ofEnchytraeus buchholziVejdovsky ,187 9 (Oligochaeta)an dpopulatio ndevelopmen t inartificia lsoi l 57 4.1. Introduction 57 4.2. Methods 58 4.2.1. Life-history characteristics 58 4.2.2. Population development inartificia lsoi lcore s 59 4.2.3. Themode lfo rpotentia lpopulatio ndevelopmen t 59 4.3. Results 60 4.3.1. Life-history characteristics 60 4.3.2. Growthrate s 62 4.3.3. Comparisono f modelprediction swit hdat afro martificia lcore s 62 4.4. Discussion 66 4.4.1. Comparisono fth epresen tdat awit hothe rlife-histor y studies 66 4.4.2. Evaluationo fth epresen tlife-histor ydat a 67 4.4.3. Synthesis 70 5. Ametho dt oconstruc tartificia lsoi lcore sfro mfiel dsoi lwit h a reproduciblestructur e 72 Agriculture, Ecosystems and Environment, 34 (1991):329-333 5.1. Introduction 72 5.2. Descriptiono fth emetho d 73 5.3. Practicalconsideration s 74 5.4. Results 75 5.5. Discussion 75 6. Involvement ofEnchytraeida e(Oligochaeta )i nsoi lstructur e evolutioni nagricultura lfield s 76 Biology and Fertility ofSoils ,9 (1990) :152-15 8 6.1. Introduction 76 6.2. Methodsan dmaterial s 77 6.2.1. Theexperimenta lfiel d 77 6.2.2. Thefiel dsamplin g 78 6.2.3. Thefiel dexperimen t 78 6.2.4. Treatments infiel dexperimen t 80 6.3. Results 81 6.3.1. Thefiel dsamplin g 81 6.3.2. Thefiel dexperimen t 83 6.4. Discussion 88 7. Synthesis 90 8. Summary 94 9. Samenvatting 98 10. References 102 Woordvoora f Toeni ki n198 6bego nme the tbestudere nva nd eoecologi eva n potwormen, wist iknauwelijk swa tpotworme nware ne nwelk evrage no pdi tterrei n liggen. Nu,vij fjaa r later,be ni ke rmi jdaa ri niede rgeva lbete rva nbewust . Ditproefschrif t vormtd eneer ­ slagva ndi tleerproces . Hoewelslecht sbi jéé nhoofdstu kva ndi tproefschrif t anderenzij ngenoem dal s mede-auteur, houdtda tgeenszin si nda talle so peige nkrach ti sgedaan . Zonderd e steun,aanmoedigin g enkritie kva nvelen ,zo udi tproefschrif t erwaarschijnlij k nietzij n geweest. Opd eeerst e plaatswi li khie rmij nouder s bedanken,di emi jsteed shebbe n aangemoedigd ommij ninteresse st evolgen ,e nspeciaa l mijnvader ,di emi ja lvroe gi n contactbrach tme td enatuu re nmi jleerd ekijke ne nluisteren . Helaasheef thi jnie t meermoge nmeemake nwa tdaarui ti svoortgekomen . Lijbert Brussaard heeft,doo rd evel estimulerend e gesprekkene ndiscussie sove r hetonderzoe k zelf enhe tbeschrijve nervan ,ee nhee lbelangrijk ero lgespeel d bijd e totstandkomingva ndi t proefschrift. LeenPons ,hoewe lnie tvaa kaanwezig ,heef tdoo r zijnonbevange nkij ko pd eproblematie k mijdez esom svanui tee nande rperspectie f latenzien . Tweemense ni nmij ndirect ewerkomgevin g zijnsteed shee lbelangrijk ,e nvaa k onmisbaargewees tgedurend ed eafgelope njaren . JokeMarinisse nheeft , inonz e tallozediscussie s overon sbeide ronderzoek ,ee nzee rbelangrijk epositiev ero l gespeeld,e nRijnder td eFluite r heeftme tvee lgedul de ninze ttalloz e klussengeklaard , vanhe ttelle nva npotwormeiere nto the ti nwee re n windbemonstere no pd eLovink - hoeve.
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