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Scatterhoarding and Tree Regeneration SCATTERHOARDING AND TREE REGENERATION ECOLOGY OF NUT DISPERSAL IN A NEOTROPICAL RAINFOREST »A/rATALOGUS Promotoren prof.dr . F.J.J.M.Bonger s persoonlijk hoogleraar bijd eleerstoelgroe p bosecologie enbosbehee r prof.dr . H.H.T.Prin s hoogleraar inhe tnatuurbehee r ind etrope n end eecologi eva nvertebrate n Co-promotor dr. S.E.va nWiere n universitairdocen t bijd e leerstoelgroep natuurbeheer ind etrope ne necologi eva nvertebrate n Promotiecommissie: prof.dr . L.E.M.Ve t (Wageningen Universiteit) prof.dr .J.J.M .va nAlphe n (Universiteit Leiden) prof.dr .J.M .va nGroenendae l (Katholieke Universiteit Nijmegen) dr. P.M. Forget (Museum Nationald'Histoir e Naturelle,Paris ) SCATTERHOARDING AND TREE REGENERATION ECOLOGY OF NUT DISPERSAL IN A NEOTROPICAL RAINFOREST PatrickA .Janse n Proefschrift terverkrijgin g vand egraa dva ndocto r opgeza gva nd erecto rmagnificu s vanWageninge n Universiteit, prof.dr . ir. L.Speelma n inhe topenbaa rt everdedige n opvrijda g7 februar i200 3 des namiddagso mvie r uuri nd eAul a Jansen, P.A. Scatterhoarding andtre eregeneration : Ecologyo f nutdispersa l ina Neotropica l rainforest PhDthesis ,Wageninge n University,Wageninge n witha summar y inDutc h ISBN: 90-5808-777-8 Subject headings:scatterhoardin g rodents,natura l regeneration,see ddispersal , seed predation,see dsize ,mas tseeding , naturalselection ,Myoprocta exilis, Carapaprocera, Meliaceae Cover design: Jelled eGruyter , GrafischAtelie rWageninge n Photography: PatrickJanse n Thisstud ywa scarrie dou ta tth e Forest Ecologyan d Forest Managementgrou p (P.O.bo x342 ,670 0A HWageningen ) anda tth eTropica l Nature Conservationan d Vertebrate Ecologygrou p (Bornsesteeg69 ,670 8P DWageningen) , Departmento f Environmental Sciences,Wageninge n University,th e Netherlands Thewor k presented inthi sdissertatio n wassupporte d byth e Netherlands Foundationfo rth eAdvancemen t ofTropica l Research (WOTROgran tW84-407 ) "O •o T3 73 •o 3 H O C» ^1 -vl -vl 3" 3- co o1—•• •t*. 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Zaadetende dierenkunne nzorge nvoo reffectiev e zaadverspreiding,zelf sa lete nzi j hetovergrot edee lva nd ezade nop . Dit proefschrift 2. Terwijlpur ezaadpredatore nzorge nvoo rselecti etege ngrot ezaden ,veroorzake n hamsterendezaadeter sjuis t stabiliserende selectievoo rdergelijk ezaden . Dit proefschrift 3. Hetfenomee nva nmas tbi jnootdragend e plantensoorten ist everklare nvanui t uitsluitendhe tgedra gva nhamsterend ezaadeters . Dit proefschrift 4. Zaadverspreidende dierenvorme nee nbelangrij kelemen tva nbosbeheersysteme n op basisva nnatuurlijk eregeneratie ,e nmoete ndaaro mworde nbeschermd . Dit proefschrift P.D. Moore,2001 . Nature409 :77 5- 777 C.A. Peres,2001 . Conservation Biology15 : 1490-1505 5. Datbloemrijk ewegberme nsteed sbelangrijke r wordenvoo rd e Nederlandse biodiversiteit isee ngevaarlijk e ontwikkeling,nie t ietso mallee nmaa rtrot so pt ezijn . 6. Eenbab yi sgee nlarf ,laa tstaa nd ebest elar fdi ee ro pd ewerel dbestaat . vergelijk M.Dekkers ,2002 .De larf. UitgeverijContact ,Amsterda m 7. Hetgoed ei sd evijan dva nhe tbetere . Voormij nouder se nvoo rKriste l Abstract Jansen,P.A . (2003)Scatterhoardin g andtre e regeneration.Ecolog yo fnu t dispersal ina Neotropica l rainforest. PhDthesis ,Wageninge n University, Wageningen,th e Netherlands. ISBN90-5808-777-8 ;x + 166pp . Seed-eating animalsar erepute d predatorso fseeds ,bu tthe y mayals ofunctio na s seeddispersers .Thi sdissertatio n dealswit hth e interaction of nut-bearingtree s andscatterhoardin g animals,whic hstor e important amountso fseed sa sfoo d reserves inspatiall y scatteredsoi lsurfac ecaches . Itstudie s howlarg ecavi-lik e rodents- inparticula rth e Redacouch y- dispersean d predate uponth eseed so f thecanop ytre e Carapa procera (Meliaceae)a tth e Nouragues BiologicalStation , anundisturbe dtropica l rainforest site inFrenc h Guiana,Sout hAmerica . Videosurveillanc ean dthread-markin gtechnique swer euse dt ofollo wth efat eo f seedsthroughou t thedispersa l process,fro msheddin g untileithe rdeat ho r establishment of aseedling .Thus ,see d productionwa slinke dwit hdispersa l effectiveness andestablishmen t success.Withi nthes esee dfat eexperiments , seedsiz ean dsee dabundanc ewer evarie dt ostud yho wthes e planttrait s affect scatterhoarding andt otes t hypotheses onth eevolutio no f large-seedinessan d mastseeding . Scatterhoarding provedt o bea neffectiv e dispersal mode.Seedling sdi d establishfro mcache dseeds ,eve nthoug hth emajorit y ofseed swer eeventuall y dugu pan dconsumed .Larg eseed swer emor elikel yt ob esuccessfull y dispersed thansmal lseeds ,whic hoppose sth eparadig mtha tth enee dfo r dispersalcause s selectionagains t largeseeds .Larg eseeds ,however ,wer efavoure donl y upt oa certain point beyondwhic hseed sapparentl y became increasingly difficultfo rth e animalst omanipulate .Thi s resulted ina noptimu msee dsiz efo r dispersalb y scatterhoarding animals.A nexplanatio n isgive nfo rth econtrastin g results obtained inpublishe d experiments onsize-dependen t seedpredation . Establishment wasfa r more likely inyear s ofabundan tfruitin gtha ni nlea n years,an dth eselectivit y of rodentsregardin gth esiz eo fscatterhoarde d seedswa s alsogreate r inric hyears .Scatterhoarde r responsest osee dsiz ean dabundanc e aloneca nexplai nwh yman y nut-bearing plantspecie s havemas tseeding ,th e alternationo fyear swit habundan t cropsan dyear swit hfe wo r no seeds. Regeneration ofC. procera innatura lfores t cameexclusivel yfro mseed s cached byscatterhoardin g rodents:see dpredatin g insectsan dmammal s destroyedal lnon-disperse d seeds.Exception swer eseed sshe db yparen ttree s alongo rwithi ntreefal lgaps .Thes e highligh tenvironment s permittedseedlin g establishment evenfro m heavily infestedseeds .Therefore ,regeneratio n needno t bea timmediat e risk inmanage dforest swher escatterhoardin g rodentsar e scarcer, butwher e lightavailabilit ytend st o begreater . Key-words:scatterhoardin g rodents,natura l regeneration,see ddispersal ,see d predation,see dsize ,mas tseeding ,natura lselection ,Myoprocta exilis, Carapa procera, Meliaceae Contents Voorwoord 1. General introduction 1 2. Logging, seed dispersal by vertebrates, and natural regeneration of tropical timber trees 11 with P.A. Zuidema pp.35-5 9 in R.A. Fimbel,J.G .Robinso n andA . Grajal (eds). The cutting edge. Con­ servingwildlife inlogged tropical forests. Columbia University Press, NewYor k(2001 ) 3. Scatterhoarding rodents and tree regeneration 29 with P.M. Forget pp.275-28 8 in F. Bongers, P.Charles-Dominique , P. M.Forge t &M .Ther y(eds) . Nouragues. Dynamics andplant-animal interactions ina Neotropical rainforest. KluwerAcademi c Publishers,Dordrech t (2001) 4. Rodents change perishable seeds into long-term food supplies 45 withF. Bongers andH.H. T. Prins submitted 5. The role of seed size in dispersal by a scatterhoarding rodent 51 withM. Bartholomeus, J.A.Elzinga, F.Bongers, J.den Ouden andS.E. van Wieren Pp.209-22 5 inD.J . Levey,W.R . Silva& M .Galett i (eds).Seed dispersal and frugivory: Ecology, evolution andconservation. CAB International,Wallingfor d (2002) 6. Seed mass and mast seeding enhance dispersal by scatterhoarding rodents withF. Bongers andL. Hemerik 69 submitted 7. Stabilising selection on seed mass by a seed-dispersing rodent 101 with L Hemerik, F.Bongers, F.J.Sterck, S.E.van Wieren andH.H.T. Prins submitted 8. Predator escape, gap colonisation and the recruitment pattern of three rodent-dispersed rainforest tree species 107 withF. Bongers andP.J. van derMeer 9. Synthesis 121 Summary / Samenvatting 135 References 145 Curriculum vitae 165 Voorwoord Onderzoek doen in tropisch regenbos is een droom die vele veldbiologen koesteren. Ook ik deed dat, en met dit promotie-onderzoek is die droom gerealiseerd. Ik mochtwerke n opee ngeweldig e plek- het ongestoorde regenbos van Nouragues in tropisch Frans Guyana. Een plek die ik liefheb vanwege de natuur en tegelijkertijd haat vanwege de maandenlange afzondering van thuis en de eenzame opsluiting in een kleine groep mensen waarmee het toevallig maar moet klikken. Je remerciePierre Charles-Dominique et le Centre Nationalde Recherche Scientifiquepour me permettre detravailler aux Nouragues. Zesjaa r lang heb ik in Frans Guyana enWageninge n kunnen doen waar ik zin in had, onderzocht ik wat ik zelf had uitgedokterd en waarvoor ik zelf de financiering hadgezocht . Helemaalvri j zijn omt e onderzoeken waarj e nieuwsgierig naar bent, als het maar leidt tot een proefschrift: dat is een ongekende luxe. Dank aan de Nederlandse Organisatie voor Wetenschappelijk Onderzoek, die brood zag inmij n ideeene nplannen ,e ngel doverha dvoo rd euitwerkin gervan . Ik benda noo k niet alleen maar blij dat heta f is.Wan t het is nog helemaal niet af.
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