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Ji __ _ THE IMPACT OF COMPETITION AND HERBIVORY ALONG A PRODUCTIVITY GRADIENT AN EXPERIMENTAL APPROACH WITH FOUR SALT-MARSH PLANTS herb(+) herb(-) _t_ _1_ 1 LJL. - _1__ J _u_1____j_ 1 _±_ J__t__I_ - L i i'.âii -*- ,±jji.ji, -- - __i_-I- L Ti— ta -1- - _i_ I III I'--t4t-'-- _z_ ,i £.ji £. 1 iiPiji"ii 1.-I-liL - - - ± ------- ---_i , *' ,- -4, / 4_ comp(+) comp(-) comp(-) comp(i-) Carsten F. Dormann c THE IMPACT OF HERBIVORY AND COMPETITION ALONG A PRODUCTIVITYGRADIENT AN EXPERIMENTAL APPROACHWITH FOUR SALT-MARSHSPECIES CarstenDormann Diplomarbejt Groningen, March 1998 Department of VegetationSciences Botanical Institute at the Facultyof Natural Sciences Christian-Albrecht-universjty,Kiel Germany ACKNOWLEDGEMENTS The final part of a German biology study, the "Diplomarbeit", has the function to document the students ability to work scientifically with only little help from his or her supervisor. However, to me, as to others of my colleague students, there is a satisfaction deriving from the research work. For most of us, this is the first time to use the knowledge we gathered during our education and to creatively perform an investigation of our interests. The pressure of own expectations and an always-too-close deadline makes thisfinal work cumbersome for everybody.I am indebted to a number of people for giving me the possibility of nevertheless enjoying this time: This work would not have taken place without the supervision of Prof. K. DierJ3en, Prof.J. P. Bakker and Dr. R. van der Wal. They created the possibility for "science without frontiers", which refers to frontiers between countries as well as between the classically separated fields of plant and animal ecology. René's enthusiasm forfieldexperiments andhisbroad overview enabledthis experiment to take place without major shortcomings. For practical aid I especially like to thank Yzaak de Vries, for raising the seedlings, and Anne Kudlik for helping me with the transplantingofthem;Lukas BouwmanandGuidoHongerboom,Institutvoor Bodemfruchtbarheid, Haren, for their matter-of-cause support with the nematode extraction and determination. Harm van Wijnen provided all different kind of data and he and Adrian van der Veen were always open to my questions and problems. When I came to Schiermonnikoog in April 1997, I was nervous about what would come. After the first two hours in the fieldstation, De Herdershut, I never felt uneasy again for my foreignness. That is due to the amazing politeness and the true warmheartedness of all people I met there during my stay. Daan Bos did the biggest share in comforting me. His friendship sailed me through hard and easy times. I can hardly think of anything he hasn't done for me. My biggest thanx to you! Julia Stahl, Olivier Beucher and Daan Bos were the "field peer group" to me on Schiermonnikoog, as were Lidewij van Katwijk, again Daan, and Quinten van Katwijk in Groningen. Of all the other friendly people with whom I shared biological thoughts of the one or other kind, I want to mention and thank Leo Bruinzeel, Peter Esselink, Dik Heg, Bas Kers, Peter Kunst and Maarten Loonen. Finally, I like to thank my parents and Anne Kudlik for love and faith in whatever I was doing. This study was partly supported by a scholarship of the Dr.-Helmut-Robert-Gedächtnis- stiftung, University of Kiel, which is gratefully acknowledged. - TABLE OF CONTENTS PART 1: THE IMPACT OF COMPETITION AND HERBIVORY ALONG A PRODUCTIVITY GRADIENT AN EXPERIMENTAL APPROACH WITH Foui SALT-MARSH PLANTS ACKNOWLEDGEMENTS 3 TABLE OF CONTENTS 5 INTRODUCTION 7 SITE DESCRIPTION 9 THE SALT MARSH OF SCHIERMONNIKOOG 9 TI-IE VEGETATION 9 THE PLANTS AND THEIR HERBIVORES 11 SETTING THE STAGE: THE FACTORS UNDER MANIPULATION 12 METHODS 15 THE EXPERIMENTAL SET-UP 15 RECORDING PERFORMANCE 16 ADDITIONAL MEASUREMENTS 18 STATISTICAL ANALYSIS 19 METHODOLOGICAL CRITICISM 20 RESULTS 25 COMMUNITY RESPONSE TO HERBIVORE EXCLUSION 25 SURVIVAL ANALYSIS 29 ANALYSIS OF TRANSPLANT BIOMASS 35 FLOWERING 40 THE DIFFERENCES BETWEEN PLANT AGES 43 RELATIVE BIOMASS REDUCTION ALONG THE PRODUCTIVITY GRADIENT 45 DISCUSSION 47 COMPARISON OF MEASURED PARAMETERS 4- ALLOCATION TRADE-OFFS IN THE FOUR INVESTIGATED HALOPHYTES 49 THEIMPORTANCE OF COMPETITION AND HERB! VORY 50 THE INTENSITY OF COMPETITION AND HERBIVORY 54 CONSEQUENCES FOR SALT-MARSH SUCCESSION 57 ABSTRACT 60 ZUSAMMENFASSUNG 60 6 APPENDIX .62 TRANSPLANT-LENGTHANALYSIS .62 REFERENCES 65 PART2: SALT-MARSHNEMATODES Do NOT LIVE U TO THEIR REPUTATION AS REGULATORS OF PLANT PERFORMANCE INTRODUCTION 71 METHODS 73 SITE DESCRIPTION 73 SAMPLINGAND SET-UP 73 DETERMINATIONOF NEMATODES 74 INDICESAND STATISTICS 74 RESULTS 75 NEMATICIDE APPLICATION 75 NUMBEROF NEMATODES IN UNTREATED POTS 75 PLANTBIOMASS 76 DISCUSSION 77 ACKNOWLEDGEMENTS 79 REFERENCES 79 EPILOGUE 83 INTRODUCTION It is in changing that things find response.—HERACLITUS Species composition, species richness and growth forms are observed to vary ina general and predictable way during succession, beginning with species poor colonist communities, leading over diverse grass- and shrub-dominated stages to tall- but slow-growing woodlands (Connell & Slayter 1977, Noble & Slayter 1980, Glenn-Lewin & Van der Maarel 1992). This has caused a wide interest in the factors producing these patterns. Ecologists agreeon plant competition, herbivory and physiological stress to vary with system productivity (Grime 1973, 1979, Tilman 1986, 1988, Coley 1987). In the course of primary succession, the driving factor of species replacement is mostly the accumulation of nutrients, either autochthonous by humus accumulation (e.g. Cowles 1899, Olsson 1958; 01ff et al. 1993, Gerlach et al. 1994), or allochthonous, by weathering of substratum (e.g. Cooper 1923, 1939) or clay sedimentation (e.g. Roozen & Westhoff 1985, 01ff et a!. 1997). Consequently, primary succession is usually accompanied by an increase in productivity and standing biomass (Bazzaz 1979, Pickett et a!.1987). This allows to transfer the literature on biotic interactions along gradient of primary productivity (e.g. Van de Koppel 1997) to primary succession. As productivity is commonly measured as peak standing biomass, I will use the terms synonymously. There are contradicting views on how the intensity of competition changes with increasing productivity: Newman (1973) and Tilman (1985, 1988) predicted competition intensity to remain constant, with a qualitative shift from competition for nutrients in early stages of succession (=lowproductivity) to competition for light in later successional stages (highproductivity). Grime (1973, 1979) argued that in unproductive habitats overall competition is low and increases with standing biomass. Support for the former comes for instance from the studies of Wilson & Tilman (1991, 1993) and Reader et al. (1994), while the latter is supported by observations that competitive ability is related to plant biomass (Gaudet & Keddy 1988, Keddy 1989), leading to intense competition at productive sites. Up to now,'the few available data from natural and experimental productivity gradients are conflicting" (Goldberg & Barton 1992, p.780). As Grace (1995) pointed out, much of this conflict might be clarified when scientists would agree on how to measure competition intensity, either in terms of absolute biomass reduction (as favoured by Grime) or in terms of relative biomass reduction (favoured by Tilman). Tilman's resource-ratio-hypothesis (1982, 1988) was a major step to a mechanistic explanation of succession, which assumes plant species to be superior competitors ata characteristic set of resource conditions, e.g. to low light and high nitrogen (=lowlight- nitrogen-ratio). As the ratio of the resource availabilities changes during succession, species replace each other. The model incorporates herbivory only as a "loss constant" (Grace 8 COMPETITION AND HERBIVORY ALONG A PRODUCTIVITY GRADIENT 1990),however, and thus assumesitto be constant over successionalstages(or productivities). Grime (1979), and especially Oksanen et al. (1981), expect herbivory to increase in importance with biomass available. As soon as herbivore biomass supportsa third trophic level, i.e. carnivore predators (Oksanen et al. 1981), or as food quality for herbivores declines (Van de Koppel et al.1996), their models predicts a decrease in herbivore impact. Experiments assessing these theories have provide evidence for both tendencies: Herbivore impact increased (Rice 1987, Reader 1992) and decreased with plant biomass (McAucliffe 1986, Ellison 1987). Experiments manipulating both competition and herbivory are as abundant from marine environments as they are scarce from terrestrial systems. The former are characterised by a multitude of indirect effects, cascading through all trophic levels (e.g. reviews of Menge & Farrell 1987 and Menge 1995), making the interpretation of results an intellectual challenge (Diamond & Case 1986). Too few field studies are available for terrestrial systems to conclude about the importance of competition versus herbivory (Louda etal.1990, Goldberg & Barton 1992). Those studies have shown, however, that the impact of herbivores might enhance as well as retard succession (Davidson 1993, Crawley 1997), and that herbivores might intensify competitive differences or smoothen them (Dobson & Crawley 1993, Bonser & Reader 1995, Huisman et a!. 1997). Salt marshes are ideal for the examination of structuring forces: The vegetation harbours only a few plant species and, in the case of some back-barrier salt marshes, no direct human impact confoundsoroverrulesnaturalprocesses.Despitetheharsh
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