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Acta Bot. Neerl. 42(4), December 1993, p. 413-433 Review Mechanisms vegetation succession: of review a of concepts and perspectives J. Andel J.P. van Bakker Department of Plant Biology, and A.P. University of Groningen, Grootjans PO Box 14, 9750 AA Haren, The Netherlands CONTENTS Introduction 413 Concepts 414 414 Vegetation Succession 415 Kinds of succession: 416 phenomenology mechanisms and functions Causes, Clarification of the Vegetation 417 framework 418 mechanisms of succession 419 Seed bank 419 Establishment 419 Facilitation 420 421 Competition Biogenic causes of vegetation succession 422 422 Herbivory Vertebrate herbivores 422 Insect herbivores 423 Parasitism 424 Food webs 424 425 Ecosystem mechanisms of succession Dune Moist 425 valleys 426 grasslands Salt-marshes 426 Mire systems 426 427 References Key-words: causes, mechanisms, succession, vegetation. INTRODUCTION succession Vegetation Nevertheless, vegetation there has been much change suggest knowledge & Slatyer Masing 1987; (Burrows the gradually 1990). Burrows in the many field of and papers terminology. abandoned in favour of the Phenomenological of mechanisms, and 1977; of subject confusion succession has been vegetation (Connell is these are terms labelled with 1990), ‘scenarios’ (Peet Glenn-Lewin & Van der Maarel 413 1992). et The more been terms such or al. reviews. concept of general have 1992), Pickett several term created as ‘classes’ to ‘models’ (Zobel & (1987) distinguished 414 J. between pathways, to attempt autogenic and confusing if In to we only can and has for long succession. The towards mechanistic Grace a & mechanisms Connell & applied are dynamics competition ANDEL, J. P. of BARKER vegetation AND Tilman in guess transient been recognized in approach has 1990), vegetation change (Tilman also in as lit 1988), and made of these use an between is terms from the research a as mechanism of vegetation approach plant on such terms, these notions. Plant to phenomenological important an decennium, competition influenced the positively GROOTJANS distinguishing mechanistic recent way Berendse & Elberse 1988; tition, however, might result which P. organization (vegetation, ecosystem). dynamics (Tilman the last during move, different levels of at A, succession, models. When Slatyer’s allogenic mechanisms, following Tansley (1935), the they addition, soil-driven and causes relate these VAN view The 1990). (reviewed by mechanisms on emphasis on of compe- underestimation of other mechanisms of vegetation an change. Overall, enlarge first to create and conceptual the on of the understanding we review to particular in enables knowledge attempt challenged focusing our purpose, felt we succession, concepts order in the of with concepts of regard perspectives to vegetation they provide succession. vegetation useful hypotheses and predictions. vegetation and succession, of terms, thus use providing a For and this then framework for further repeat earlier reviews, not to such Only concerning work. We will experimental their conclusions for the several evaluation of the mechanisms formulation review some an but rather use present purpose. CONCEPTS Vegetation successional In composition, floristic terms, least at to another) and vegetation change According which in the then been if changes characterize the there is history two association is not communities, Gleason after (1939) aggregate of plant species plant species, of textbooks, rather has created community while a than also species basis ‘species’ one of its within another, to when implies we phytocoenon a labelling together opportunistically an acknowledged vegetation being unit with by its composed does For not ‘a a plant (1926, justified opinion, but unit, coincidental that that merely aspect of a plant association definite successional in an is an status floristic composition’. is considered problems. population this Whittaker not we prevailing vegetation species, and have Gleason recognized by from the a also (cf. of the coin: ‘Are even ecological change to also were side of particular assemblages are come emphasized recognizable number of the one scarcely having forming physiognomy, majority approach have far removed organism, an However, of uniform emphasized conclusion, coincidence?’. In the communities cover aspects, individuality and order, general (from phytocoenosis one the notion of climate conditions continued as the on ‘species replacement’. plant global plant 1926 paper he the to (from what on term (1991), of in change a community present paper, dealing with mechanisms of succession, focus with the Tallis to In his coming we re-ordered 1975). The a we of succession phases Here phytocoenon). of community as worry about the distinction between stages of succession not and plant long In the composition. do 1939). a as of be to plant example, become composed individuals. a species extinct, of The may populations population disappear either from because the MECHANISMS boundaries OF of a the Moreover, using vegetation science, problems be can population term characteristics while the (Grubb of species such the about generalizations al. et life-history vegetation component of dual only not an ecosystem (Odum hierarchy, represented histories and and energy maintain by and evolution) much of individualistic This draws that (Gleasonian) ‘it is al. with succession disturbance of (e.g. vegetation White (e.g. Pickett et al. the has (1992) but also approach’ the both aspects: acknowledges plant populations [in his 1951 in coherence with their sites and growing forming part their site factors and all other forms of life still far from getting to the bottom of into recognize to with of cycling and a life to holistic a in several studies WesthofTs 1992) which of the ecosystem or growing we prefer] together between, of ‘emergent properties’ with definition (1979) largely spontaneously individuals, with reasonable & Pickett in these sites’. In occurring insight which result from interactions between of plant wrote functional a even community 1991). ‘A system he paper al. et considering to (dealing (dealing that successional as (1986) proposed 1987), species shown to attention our 1985; Falinska 1991; Myster 1989; Van Andel plants. 1979 and concept of ecosystem function’. The latter aspect has been neglected dealing case. of visible subordinate feasible and of be can the not composed be (Grime approach’ quite concept quality which is in relation plants, ‘process-functional nutrients). They suggested an Brown et the and Muller-Dombois & to plant species O’Neill of determinants of ecosystem related 1987). of all dormant seed bank. a association of individual main succession individual 1971). to (1951) ‘population-community a a an the more of strategies as population thought and oversimplified. composed as are Grime 1982; age of the habitat may have been is being as stability is that Westhoff plant community species-richness Grubb 1977; community boundaries quantitative aspects, to community productivity, as of aspect a the survives suggest may populations. Only Ellenberg (1974) recognized Dominant a with population in addition avoided if overlapping coincide not because the or involved in individuals of do phytocoenosis component populations, Such 415 SUCCESSION with we are communities, organisms (see below). Succession Until now we in rather a According that change, but the if succession into a succession. considering useful to ‘changes does not Tansley of an escape phases’ (Tansley 1935), is important part the sere. which reversible Miles (1979), markedly to changes, have that there is in no whereas as having into a successions Similarly, Finegan (1984) the at mechanisms of of use the all historical involves not which can be abandon the a different patch type’, directional defined succession as of merely phases or separated concept stages. it is vegetation change, vegetation mech- value of the ascertainable laws. Even to of term non-committal species composition (or abundance) defined are to vegetation need fluctuations. the of the concept notion of ‘a series’ alter the appearance of changed this phases subject of change in ‘vegetation change’ include useful when reviewing seem may stated sequence of a confusion sufficiently acknowledge (1935) mechanisms of succession considered state. much differentiate between fluctuation in successional be to ‘a continuous process The so is would term abandon ‘succession’ in favour recognition is series of latter best it succession. the there Though such succession, of concept succession’ and ‘vegetation terms Actually, (1990), seems change’. ‘vegetation of it the using way. Burrows to succession anisms have been arbitrary dynamics vegetation added changes that away in of When quite and general a as such that it can fluctuations are from ‘the directional change an initial with time 416 J. of the progressive, mechanisms hierarchical al' (1993) proposal s from year one succession. For McIntosh (1980), We start Miles from Pickett succession: calculate the to of of the et al.' or particular mechanisms. action pathways are mechanisms, specifying well as Kinds We one another been (cf. the al. et which of presented species of ‘phases’ succession, between pathways, predict or vegetation change; see particular successions (ii) of is cause successional of sets an sere an is and/or causes successional explain by causes, aspects mechanism is (iii) a describe and to of a Prach to (1992). explain on rate or vegetation change. Consequently, particular various combining between the mechanisms and the pathway as 1987). between different types of Burrows some from Muller-Dombois & While pathway. attention information paid we succession. ‘classes’ of vs. prefer clearly separate Glenn-Lewin vegetation ‘to kinds all encompass & Van dynamics, We do kinds der thus scale of time and space: fluctuations, Masing (1987) had proposed community and ecosystem of jargon. Overall, McIntosh (1980) simplicity, which as far that is it of the basis not take this of succession Maarel (1992) providing fine-scale as is a dynamics’, open on not. vegetation causes proposed dynamics, gap vs. The change areas already and mechanisms into consideration and mechanisms of distinguish to on an seven increasing patch dynamics, cyclic and secular succession. Similarly, ‘spatio-temporal classification of several kinds but these authors provided terminology illusion that traditional in areas causes proposal vs. phenomenological an of framework based a succession, secondary succession, primary succession, Zobel & does vegetation’, distinguished five ‘models’ of vegetation change, allogenic influences). to have we terminology, mechanisms, but actually on by vegetation), for another part (e.g. autogenic Ellenberg (1974) literature, phenomenological to the basis of the initial condition (non-vegetated on the studying statement. attempt an in all kinds of covered because in (1990), of amount may illustrate this phenomena partly the by implicitly suggests following of turnover ‘stages’ in change a who (1993), of knowledge our of distinguish to construct relationships Fresco al. et from change detect the concept of might depend of to to for successional patterns; successional course as dilemma describe, to the notion of‘kinds of succession’ surprised fully the of the change conceptual the consider the notions not how al. et of succession (1987) proposal which is GROOTJANS site where climate only not P. of succession: phenomenology adopt distinguish to to a the predict or rate avoiding responsible pathway, Models Huisman to is the temporal pattern interaction that contributes type of s left A. single a is contributing as problem and Glenn-Lewin (1987a) AND between trends and fluctuations, and history vegetation (i) pathway agent, circumstance a thus of succession succession only refer to BAKKER direction. We do some distinguish to review a mechanisms and suffices another, to The 1989). of models set degressive or Bakker type. Here it vegetation et regressive (cf. P. physiognomy that implies but should have climax, to J. ANDEL, vegetation constant’. This effectively pioneer stage of and species composition remains VAN to science, think can be is the us with another concerned, search satisfied by for we regularity replacing set agree with and terms or introducing ‘new-speak’. The distinction of understanding two largely depends past. Secondly, primary vs. secondary succession only of mechanisms of succession. First of all, our on an interpretation mechanisms of of what succession marginally initially might depend much contributes to the difference between the more have on happened the in the initial abiotic MECHANISMS conditions (e.g. driven whereas successions primary debate begin that it is succession on obviously (such kinds of succession and primary be can succession primary or although critical being as (1990), soils relate to on seres). have of in more soils. will we this From to secondary with common succession Therefore, mechanisms on that recognized rich mechanisms secondary focusing may successions secondary mechanisms related primarily by however, type of kinds to of discussing stop different hierarchical levels. at mechanisms and functions Causes, At Tilman and start is (1992), Peet involving environmental change (soil, climate change, confusing primary as succession a view. considers changes, nutrient-impoverished than with this in environmental conditions changes Gleeson & successions whether on illustrate succession would be driven secondary interactions. species small may successional mechanisms by than fertility) examples relatively driving long-term principally etc.), soil following that recognizing 417 SUCCESSION climate, The secondary. factors OF which level According to of community Tilman but idealized ends of enological to a ‘It is spectrum: mechanistic The conceptual occurred In his derived from comparative determine whether correlations related problems ‘because the causal used from for phenom- explaining formulate testable to us any the usual only through manipulative experiments Tilman relationships. short-term mechanisms? absolute entities spectrum optimal allows and of interpretation have the for not are will be found view, theory on search to that is point a observations, based are to ecologists likely that, along optimal point trial-and-error process of science.’ have we and mechanism there will be theory, given ecological pattern. (...) predictions do organization (1989), phenomenology many of which have experiments, experiments can treated (1989) address to long-term questions’. Long-term and Beeftink Only and observations fluctuations 1987; Bakker the basis of on of the relevant mechanisms of 1992). this need as (see ecosystem to be Olff et al. or Roozen & Westhoff succession is & Nisbet attention has been factors, effects’ (Grubb changes in paid to higher plants salinity, nutrient on we & autogenic and biotic and abiotic & Van der Maarel vegetation supply, quantity farther than this and allogenic applied rather than to the successions. are present in all successions’. my succession. suggested It is the 1991; al. not ‘relay to De Leeuw can fact, (...) new I Changes composition ‘In 19261 I think This classical view that has now in terms think, of the a succession’ to species allogenic party the influences of in abiotic factors, groundwater, proposed to e.g. may also distinguish that I should have gone first autogenic not In floristics’ and So-called ‘third 1992) represent succession. and find represent 1992), ‘obligate aspects. sere unravel 1993). 1980). With regard Slatyer allogenically induce succession. Tansley (1935) stated; between successional a does et may refer al. 1993). chronosequences. frequently et 1983; al. ‘life histories’ and ‘vital attributes’ of 1976; Noble 1986; Glenn-Lewin biota other than 1954; i.e. De Leeuw Wilson al. et be tested if we can succession, 1985; concerned, pre-emption’ (Egler 1973; Pickett of et experimentally to Olff & Bakker 1990; zonation spatial ‘competitive hierarchy’ (Horn 1976), (Drury stages Tooren in species attempt we mechanisms of succession that aware sequence of can Van 1982; Turner 1990; Huisman 1990; exact processes, situation, e.g. autogenic ‘initial floristics and and of the knowledge al. et between successions distinguish non to Pickett 1987; 1969, species replacement (e.g. chronosequence far conditio sine qua definitely represent phases context we As a 1989; Andresen In the actual field sites which are Westhoff (e.g. been place and to the factors allogenic factors sufficiently recognized 418 J. until tolerance to al. et Connell & and attempting Peet Pickett recently. proposed by them in that of course additional problem a successional sequence. causes The Pickett (1987), al. et study anisms of view, succession’ mechanisms and and it long and autogenic as individuals the being field in actors which they succession vegetation 1935; Odum the O’Neill 1983; analogy, a and same that reason, called ‘scenarios’ Farrell may An 1991). between the distinguish that the (1987) suggested within vary successional a between distinguishing not environment: ability; a a a or have to al. et fulfil 1986), from interest, in i.e. a allele in the a other these context & Slatkin selection: of the mix of Plant is the mechanisms of discovered (cf. context realized a in the land- re-interpreted that the aver- of other groups and competing of by Tansley of its function- who diseases frequency only the ecosystem context context competitors, particular context ‘Fisher be can be held functioning (1992), in the wider of the However, distinguished. be can of its can a ‘mech- as mechanism words, wider a in vegetation Frank context of clearly be evaluated in natural the In to an delimited. From this view vegetation change, functions. of be according processes a of considered succession’. and 1992), 1987). Therefore, quantity only context of level succession’ and ‘autogenic terms ‘causes can be can study is clearly as the on 1987). Indeed, identify autogenic to ecosystem in the of of influences field the an species an al. depends organization Hobbie e.g. et allogenic useful genotype in the population; (see Allen of the use influences (Pickett quote we group is useful seems fundamental theorem age fitness of the al. et from 1986; if the system but should also ecosystem, and For Fisher’s the be 1987; to mechanism of mechanisms of being reductionistic approach, scape. For the to facilitation, seres continuum of effects of earlier on later level of one allogenic considered ‘an efficient cause’ in a a al. et that the only interact causes termed on implies is useful of is (O’Neill mechanism a succession’ phytocentric point ing of confusion results that addresses another level. This on ‘allogenic action unlikely are the three mechanisms of each of the scenarios Chapin Connell (Crawley 1986). the main interaction specific a as Slatyer (1977) importance & GROOTJANS ‘models’ of productive. (cf. Walker P. and mechanisms. organization effect be to A. about entire hypotheses likely extremes AND that and directions of the interactions could view, our sere BARKER emphasized is that observational studies strength In not the relative mechanisms represent the the species; is P. mainly autogenic alternative way Indeed, different models of succession proposed i.e. the the models of Connell & (1992) suggested in the again once not are rather than mechanisms. change (1987) J. ANDEL, Slatyer (1977), inhibition, test VAN and food avail- genotypes within competing alleles at a locus’. CLARIFICATION After to having propose will start up with a THE FRAMEWORK reviewed several concepts and the associated reduction of the from the current mechanisms OF phytocentric point views, and jargon along of view but will finish with causes at with a on terminology, we feel the need clarification. In the present vegetation succession considering ecosystem different hierarchical levels are review, in order succession. concerned, we As to we link far as take the following position. (a) Interactions mechanisms between of plants, succession. i.e. autogenic processes, will be termed vegetation MECHANISMS Effects of biota other than (b) the heading ‘biogenic If autogenic (c) VEGETATION established. such we first framework et longer distinguishable and it is al. leave the we mechanisms of succession’. SUCCESSION and competition after only occur the interactions. In broad other emergent have plants mechanisms of colonization and species proposed by Pickett will be treated under that governs succession, facilitation, can treat thereafter mechanisms of only OF as community, Therefore, are no two discussing ‘ecosystem start plants, plant a causes these community a succession’. vegetation MECHANISMS between of properties of interaction between view and phytocentric within higher plants causes mechanisms and allogenic the particularly Interactions 419 OF SUCCESSION become regeneration, outline, and follow the we (1987). Seed bank Plant species can bank or result of as become established immigration These processes 1990). Buried viable attention seed important are context Seed banks of whether Grime their not or related longevity, the to Therefore, al. et (i) transient seeds of adequate seeds that grasslands seed long-term persistent the seed bank. problems et al. Sykora seed bank. 1983; related to McDonald the rate in chalk or This of on nature are useful in of rare is an the less; (ii) in use as or to & far as & they can Blom short-term be 1992). proposed (Bakker distinguished 1989; three types: persistent seeds seeds studies indicate that the (Willems see if we do der Valk (Bakker the process in the 1983; Poschlod Van grasslands of 1991), 1992), 1989) have not dry no of re-establishment of propagules, rather than landscape increasingly poses introduce them (Gartner spatial dispersal urgent research so according (Thompson (iii) long-term persistent immigration species land little information too Voesenek (e.g. 1987; but reserves in the year provides Preliminary that scientific plant communities, ‘transient’ or throughout grasslands means condition. 1992). 5 years; years. Maas Constraints of succession to 1982), and moist availability 1993). of 1 least 5 at depends Fragmentation of lack of seed responses first a received change to These authors 1991). period a species in disturbed ecosystems species on & have They ‘persistent’ as in the soil for 1 year persist & Grime who termed (1916), being bank classification has been 1986; Pfadenhauer & (Van Dijk plant disturbances cf. also Poschlod of characteristic (Vyvey vegetation. of the latter habitat that remain viable in the soil for majority by (‘nudation’) detected were because estimates 1993; the been classified that remain viable in the soil for litter fens of Clements this classification of seed banks frequency a more Thompson have availability Jimenez & Armesto germinable 1979). However, about the site present, either in the seed are neighbouring populations (Hodgson of potential 1989a; species species of conservation and restoration of predictors management (e.g. Peart if viable been mentioned part are only from already ‘ecesis’, banks in the mainly had and ‘migration’, them they a of plant propagules as topic. Establishment Once seed plants have become established availability Hester (Grubb et al. may become 1991). 1977) is a limiting and In many ecosystems, condition to set for the maintain a fruits, microsite regeneration availability from seeds (e.g. rather than Peart 1989c; permanent existence of ‘regeneration species richness, as has been niches’ shown for chalk 420 J. grasslands (Grubb al. et & De be to seems Vries 1992). indicated requirements during the resource of concept ‘safe term P. BARKER & (Harper entire process of A. P. niche 1977), GROOTJANS Rusch 1984; is for 1993), & Russell (Hutchings regeneration site’ AND Verkaar for salt-marshes and 1992) The the by J. ANDEL, Schenkeveld 1982; & Blom flood-plains (Voesenek Bakker VAN much 1989; wider than this niche includes all as from dissemination regeneration, to establishment. Here, do we feel not because concerned, syndromes’ (cf. Rozijn the germination emerged stage is seedling life of active for over germinated quickly for competition productive to thought survive to to (Voesenek in these a in germinating soils, germinated less and at successional thus alternating temperatures, the perceive pioneer species require conditions for 1980; Van Baalen Bouwmeester et a al. 1993). In Fenner species (such colonizing ability Rather 1989b). gradients, et for in example al. 1987). On Gleeson & Tilman suggesting in this a is this for studies on were the different of capable escaping species, delay less on more of the been rich soils, relatively they responded a on having for hay successional Grime & (e.g. 1988; Pons discussed the clearly germination successions bare on not shrubs and as trees), to be the the basis of hypothesis this detected trade-offbetween colonization a In relationship 1992; seeds to of between seed species (Fenner species Werner 1978; successional along (1979) primary angustifolius proposed succession on the poor reproductive allocation, decreasing and ability al. related general, and in Rhinanthus a et as smaller sized than the seeds are phenomenon, of succession’. (1990) similarly in soils, but the within case 1975; Silvertown of seeds among different straightforward seems Jarvis 1991; Voesenek properties Solidago species (Werner 1979) ‘colonization-competition soil, way Therefore, succession: suggesting pioneer species (mainly annual and perennial ruderals) of later successional (Ernst is that the stimulus and/or alternating temperatures Pons (1987) secondary light survival seedling 1982; Van Tooren & succession. vegetation Peart 1992). cut that these slower rates, and a early photosynthetically species, Later autumn. ‘germination into important aspect from several been ecological spring. next size and al. et of the had grasslands The in such secondary grassland high percentage, suggesting a evolved for that, arranged 1992). report the results (1994) species be to vulnerable part most 25-years period. Early light by Seeds of many to al. et purposes, a stratification and until OUT plant. agricultural of time periods considered the characteristics of germination used a chance a 1985; (1979) be can Van der Valk is germination physiology as Chabot physiologies are far as & Angevine germination physiologies into detail go & Van Andel have seedlings to with agree species-specific purposes, that the need we competitive ability (for nitrogen case). Facilitation It is well two-way, matous 1991), known plants positive phenomenon water and Robinson that plants of different species effect. This (Joenje 1978; exemplified by Schat interactions between of hydraulic nutrients 1991), (De Hullu 1985) ‘Experiments may be in and or 1984; Laan different lift in semi-arid through interspecific benefits from can et al. a beneficial, areas (Caldwell mycorrhizal N-fixing plants, soils 1989; Schat & Vaccinium after their death (Akkermans dry grasslands exhibit the aeration of anoxic species et al. as Van 1991), or Beckhoven (Maillette networks either one-way by aerenchy- 1988), the the transport of (Grime et al. 1987; hosts for hemi-parasites 1971). We quote from Gigon (1990): in Switzerland show that adult plants can have positive MECHANISMS OF microclimatic and SUCCESSION effects edaphic Other experiments species. 421 and between adult individuals of many not direct but via other of YAM composition to abandoned fields Within the scope of the Connell & if Only plant species, know does wood not plants, concern (Shigo species As 1975). far cite the (1982) N-fixation which succession. on europaeus the by Bay (Crocker et al. (Joenje facilitating of the of plants are et initially et al. (1980) The in al. majority of et al. fungal species 1993), an Miles areas and (1987a) from which shrub increase in the examples dead on Gerlach 1958; and in reclaimed are in Glacier e.g. (Olson of forest, of facilitation 1974), we Ulex rate Australian open Major in dune 1967), of successional examples, an to (Horn of facilitation we example showed (see is related succession’ N-fixing legume fire a poor soils OUT 1991; few a Facilitation term establishment clear the effects of the on Lawrence ‘obligate concerned, mentioned only succession. on in soil indirect facilitation indirect Farrell In polder this under the leaving the but (Miles Tilman 1991), analogous prefer we exhibit their beneficial effects formation (sensu practice, however, distinguished. Therefore, term plants (direct facilitation), soil dynamics’ (see competition. will discuss and changes ‘soil-driven of processes interaction plant-to-plant a plant-induced resulting time most This aspect. and between distinguish areas 1978). Many result secondary of between soil succession. to viz. the succession of Grove Westhoff & Van Oosten 1989; The should we leading the latter to Macrozamia riedlei after Major 1955; & al. successions primary to et Similarly, cycad facilitate may related of Bradshaw study those adequate. (1992) are species positively was inter-relationship facilitates the fungi, Glenn-Lewin & Van der Maarel the succession during secondary (Egler 1954) higher as that fungal community succession, on and directly but changed showed (1991) density. floristics’ facilitation term plant the beneficial effects most cases al. concomitant a YAM coexistence 1916), ‘relay plant one is the indicating 1977) is confined Slatyer In et of the YAM present paper result in ‘reaction’ (Clements 1976). diversity and properties, plant productivity (sensu Johnson communities fungal forest. The and establishment of other germination plant species. organisms’. related with soil C and N, thus interactions which the on literature survey show that positive interactions exist a The the heading ‘ecosystem ‘facilitation’ in Hobbie latter favour as be a via thus 1992), might called distinction between direct and direct and indirect facilitation consider facilitation to not indirect way, an 1987b; 1988). to in as an mechanisms of of ecosystem hardly can be ecosystem process and succession’, terms such the at same feedback as mechanisms. Competition for Competition succession al. 1987; Tilman but mechanism, importance 1986). of Grace the have they 1991) (i) resource outcome knowledge long Horn 1976; None competition as: has for differ in in of and been their or the not claim the plant an 1977; communities views of Grime ‘stress-tolerators’ The 1985) is question of the mechanism of to Grime intensity nature, (cf. and (Grime the main mechanism of important competition whether ‘maximum resource requirement’ (Tilman of succession. of as Slatyer authors view contrasting (ii) recognized Connell & these discussed whether (Tilman 1987), ‘minimum ing 1916; 1990). (1990, questions such nutrients resources (Clements and be Welden & Tilman to the the et only relative Slauson by considering 1979) do compete for capture’ (Grime 1987) plant property understand its role as or in determin- be discussed here is whether competition Pickett 1979; to a we should mechanism 422 of J. succession. vegetation plant properties competition cession, as the quantify of intensity growth, population provides as (1993), Elberse feedback Bobbink in chalk which to Berendse & is requirement’, we competition for nutrients establishment gain in it and the of the on of replacement & nutrient increase in is abilities of species may not (Marrs & Hicks the context of BIOGENIC a & the on (1982, Weissing Slatyer can and (1991) tetralix by Erica species as Agnew (1992) in cycling the ecosystem. become to It is worth for nutrients concept of 1988) more much as result in a proposed by model. general In than is complex in a of ‘positive-feedback which succession, change, 1988). The But be it term switches in the represents inhibition, due to no example, for competition a change cyclic (Watt inhibition at there is dominance. For competitive plant competitive community modifies a way, this allelopathic competition. of prevention particular aspect treated processes successional predomi- mentioning ‘relative nutrient more the meaning may inhibit further to a to 1993). 1977), be considered 1986; Gimingham climax process- experimental of the latter N-supply. competition based between inhibition and and nutrients in addition this words, opposite of facilitation (or negative feedback switches). However, distinguishing which its to other In (1986). Examples that the suitable for itself. In more stands of Pteridium aquilinum light al. competitive ability of Tilman’s which indicate all environment, making to suc- important most the level resources). competition for light is (Huisman Wilson & 1989b). plant communities’, discussing we want vegetation The Brachypodium pinnatum an which case by existing plants, Peart shown showed that Connell (sensu in importance be found in Tilman & Wedin that the model of aware if a is that it relates species responses soil et when hand, as question refer here. increase in 1987), particular a should be ‘Inhibition’ ability (cf. (1984, competition species On the other GROOTJANS avoid the population-community can only result of a can P. habitat, and that this replacement is irreversible due (1988) as of A. treat we parameter (R*: O’Neill grasses we to should be known. limiting by AND levels, relative ecosystem Molinia caerulea al. shows (1993) Aerts an of the of et an have (1990) grasslands that Huisman least the its between the successional on eutrophication positive between, and reduce advocated Molinia caerulea results from Similarly, to can interface an research Berendse & nant insufficient succession. plant competition allocation) approaches, Wedin & Tilman a it BARKER competitive ability vegetation competition equilibrium in functional result of Is P. of Tilman’s (1988) mechanistic approach (dynamics, follow-up words: determine the mechanism of a the mechanisms of advantage concept other J. ANDEL, in this context? In view of the hierarchical phenomenon of which In VAN in competitive 1947, seems 1955) useful only or in stage of succession. CAUSES OF VEGETATION SUCCESSION Herbivory Vertebrate herbivores. vegetation depends productivity (due to herbivores. is assumed to level. In such to According on the to Oksanen limiting nutrients or At intermediate levels of be close a 1987). Probably, to the still levels of the impact soil-plant of herbivores system. At low temperatures), vegetation is herbivore is higher (1990), the primary productivity carrying capacity, situation the at of productivity so that the ‘prisoner the primary productivity not crop on the primary attractive consumption standing of its food low is kept supply’ (Drent pressure at a low & Prins the herbivore densities are MECHANISMS the longer regulated by no vegetation This abiotic has been (Joenje 1985; Olff and 1992) by lemmings (Batzli herbivores 1993). As Bakker the soon as & of (deflections in Verkaar species-rich way) a & and Brown competition a Fresco and Carex savannas grazed Prins of rate 1989; among lesser responsible on longer a herbivores. In a sheep could grazing succession, vulgaris after grasslands, Trirhabda spp. succession was area while snow to (Kerbes the grazing al. et vulgaris (1990) acidic soil. for of plant and increases. supports 1988, a for on feed Diptera A similar but in their by modifications the intensity of distinctive and a review). individual on a of cover al. et increase in relative on dominant conclusion, foliar-feeding by recycling and the case its and light of a the forbs, was a or is applied prairie even a cyclic underground, chewing period to on com- detect the succession vegetation (Brown larvae of mainly bare sandy and of 3 years first resulted thereafter in a relative increase of by above-ground Hemiptera annual forbs. a in by chrysomelid beetles, fluctuation which resulted insects caused in beetle microfloral/microfaunal enhanced on result heather observed was early successional plants during may faeces, quality. irreversibly replacing grasses have been on via food geese densities of complex plants grasses, for ‘optimization 1992). canadensis Insecticides of annual as higher photosyn- the of nutrient These authors showed that roots of studies majority of increased fitness consequences regrowth high this preferential sap-feeding Cicadellidae feed In the phenomenon Elimination of these soil herbivores to avoided intended for in on temporal a grubbing 1990; Hik heathland, 1983), Gange 1991, 1992). and due were result showed that speeding-up and forbs. The latter effect grasses herbivory, to a 1987). al. et relative increase of the richness. large Jeugd external factor, land, induced. In grasslands, most of the primary production of insect herbivores Coleoptera phases, by der dominated by Puccinellia phryganodes attack of the dominant Solidago (McBrien patches examples consider not swards phenology time-scale (Berdowski an Stanton Brown & perennial & Van succession In the latter case, account did on few a evidence. Jefferies geese related the for only ratio may Hik & Calluna rhizosphere munity (see a dune 1990), browsed some depending Lochmaea suturalis may result in subordinate perennial in and review of the literature that the convincing was destruction of the habitat 1990; This waterfowl by al. et Prins vegetation while other kept intact, defoliation. He subspathacea, was However, impact either as 1977), coastal tundras On calcareous former arable defoliation, species. grazing intensity’ by and the that the 1992). leaf higher following rate Calluna so long as grazed 1989; Andresen & 1986; the direction of concluded from (1988) rate and grasslands, of the result of defoliation have Insect and 1980), succession. indicate adverse effects of which salt-marshes e.g. 1985, Georgiadis stable relatively 1957; Zeevalking carrying capacity, enabling grazing (Gibson growth is density of herbivores becomes reduced by of the thus sheep, thetic parasites, al. (1983) showed that moderate sheep grazing intensity resulted in patches of et restoration a ecosystems, al. et grassland being intensively grazed the and least retard further succession. at or cattle (Bakker (see McNaughton below the level of variety a by rabbits (Watt grazed by predators vegetation biotic factors inhibit or shown for grasslands grazed indicates that the general picture factors but rather vegetation, into woodland. develop can stress 423 OF SUCCESSION slowing in In later successional increase of species an of the rate of succession grass-dominated sward, and thus resemble the well-documented effect of vertebrate herbivores, whereas root-feeding insects effected an increase in the rate of succession. 424 VAN J. J. ANDEL, BARKER P. AND A. P. GROOTJANS Parasitism In the Dutch cleared away influence the pathogens they die from or The or Alexander mentioned (1987) parasitizing plant the example, and & the soils. result Rosa N-fixing in on arenaria in coastal and soils, appeared also thus contrast research to (semi)natural culture, on dune amount on have might of rust to et for thus This 1981). nigra, Ligustrum on reduced Festuca rubra ssp. root system, Otten Empetrum nigrum responsible al. reduce the mycorrhizal of capable are Sambucus to, e.g. or acidified of vitality arenaria (Van 1993). interactions almost unfortunately are For willows stands pure This They areas. related favouring plant-parasite ecosystems of the Longidorus genus the of succession dunes, selectively succession. the succession, 1988; plant- well. Zadoks as seedlings and other nutrients (Oremus & Putten & Troelstra 1990; Van der Putten In the in coastal calcareous nematodes species, fungi, vegetation if became dominantand killed the willows fungi nodules, few formed nearly the or interactions in & Van Damme a the ice had receded. During proved (Den develops differently community many be whether plants not pathogenic which once spruce. acceleration rubiginosa the were to of whether pests be, only by that killed species Nematodes of 1939). of phosphate Endoparasitic Ammophila der the to Nooij may accelerate Hippophaë rhamnoides uptake may and vulgare birch and to Wadsworth including reducing showed river Yukon in Alaska, stands of damaging damage pioneer but various canker and heart rot declined, and populations which De dominated community, a alexensis, accelerated succession system, is been never test supposed plant-pathogen level (e.g. Melampsora bigelowii rust S. banks of the (Baxter in species willow pulchra gravel both studies few a concerning two-species plant community a that ‘the crucial plant communities has of studies the relations may affect pathogen Salix If 1990). but this has pests, but whether the community on is Labyrinthula macrocystis catastrophy, of submarine Zostera beds evergreen Harper (1990) majority natural communities focus of pathogen composition disease excluded’. are 1932. The We agree with 1987). Hartog 15 000 ha some for this almost irreversible responsible suffer estuaries, around in the of context devoid of such a agri- type of investigations. Food webs McNaughton (1992) has that disturbance proposed at level in one a trophic have effects far removed from the initiallocus of disturbance. Because of the of food webs, only a few case studies are to be referred in the to of context web can complexity vegetation succession. Prins Africa & ungulates section, the such (1991) central increased an effect with species on events reconstructed of bush encroachment in savanna rinderpest pandemic having treated the be reported aware on coast. of the Depending symmetric or on in limpet the an asymmetric, on Acacia tortilis plants intertidal in the former community activity competition but it is East among effects, via animals. foraging limpets, epidemics even-aged parasites succession decreased of of indirect occurrence of study Barnacles abundance. could be a direct effects of anthrax or impala, which enabled the establishment Oregon algal not and on thereby between two clear whether this has algal abundance. Robertson herbivores Jeugd (1993) should also we barnacle as after So, Farrell der coinciding as stands. Van (1991) studied mangrove forest ecosystems consumed overall less than 5% of the canopy in tropical production, Australia. but the Insect rates of MECHANISMS herbivory OF SUCCESSION individual on species ranged tree Heritiera marina, (Avicennia 425 differences in chemical composition. common are mangrove species tree also contribute messa) playing role in the key a References to ECOSYSTEM For of reasons case autogenic latter, confined ourselves we clearly be level of the succession to of entity The and case in such an examples succession and ecosystem Dune seres in meant are case of abiotic causes two thus that the recognize to the of context complex processes. have been we succession. distinguishing With regard it is allogenic factors, and interaction (feedback to succession make a can longer no at the mechanisms) is involved choice for the the to mechanisms autogenic types of process indistinguishable. vegetation illustrate the to with succession, primary Westhoff & Van Oosten and causes Cenlaurio 1991) interface increasing emphasis This is in ecosystem ‘ecosystem’ between as vegetation the latter. on nigricantis Saginetum governing the rates of more pathways matter (indirect facilitation). blowing The saline various allogenic origin, becomes but they by (Stal 1985; (De Wit the early 1989) Visscher 1992). establishment of the main plants, the autogenically 1988; Roelofs et al. an rate Dieren induced oxic environment around the et al. from 1984) retard the succession mollusc shells, and cause, either determinants of how 1991). High pH values, to secondary a the bare and (Pluis 1993), on beach plains 1990). After hydrological regime long the pioneer stage low input of nutrients, and plant roots (Ernst higher productive precipitation are organic fixation of C and Pluis & De Winder of decalcification and the of seres mats important part by 1934; i.e. These factors autogenic build-up primary allogenic (Van baltici- hydrological regime, blowing. formation of microbial allogenic (and partly autogenic) persist (Grootjans carbonates This is the in blown-out dunes or Junco beginning of these successional which green algae and cyanobacteria play 1991; allogenic communities, the local on and sand may be modified by the 1988, the Parnassio-Juncetum atri- conditions, depend mainly At the very stabilized pioneer al. et between association calciphilous from develop acidification, desalination, mainly among exemplify the complex interaction under or, of sand valleys (Westhotf 1947; Grootjans may may Further successional capilli. dune mechanisms. autogenic Schoenetum can Sesarma leaves, valleys Successional are of mangrove study. few next where (e.g. because community components causes, that it is advisable extent in place vegetation continuous soil renders the wetlands, of allogenic the influence the can crabs the need stress takes existing literature, causes in the between and SUCCESSION biogenic to because vegetation, vegetation generally the distinguish to the between allogenic distinguished. However, worthwhile OF in touch with remaining to sesarmid crabs addition, These decomposition serve to vegetation MECHANISMS In zone. due propagules of eight of the propagules processes and ecosystem community mechanisms and insects. 35-0% to probably area, function’ of mangrove habitats. studies may of leaf 20-80% of the intertidal the process of (Excoecaria agallocha) expanded mangrove the across to development interactions among on ‘nursery ground these few and patterning largely the damaged by were species from 03% Moreover, important post-dispersal predators distribution patterns of N of littoralis) an & Van der Ham communities. Both of calcite may act as 426 J. in the buffering agents formation Moist P. (F. thus zone, J. ANDEL, P. BAKKER preventing organic AND matter A. P. GROOTJANS deposition and soil Sival, personal communication). grasslands When intensively used abandonedand in changes vegetation to restore be to first the former semi-natural decreased. The system largely governed by nutrients limiting was hay plant-soil peaty soils appeared in the Drentse A brook agricultural grasslands for cut of the productivity to root VAN limited N by succession secondary and such and K, by later P the species of the entire addition. in Holcus lanatus and as were these on the response of the successional 1987; Olff 1992). The productivity (Pegtel indicators of eutrophic conditions, valley system species-rich hayfields, As result, a Agrostis stolonifera, were replaced by Plantago lanceolata, Anthoxanthum odoratum and Rhinanthus angustifolius, which in had turn to make room for a acutiflorus Juncus dominated community 1989). Here, the competitive abilities of the successional species nism) depend on the nutrient autogenic influences the status of the soil. regard the to and thus demand indistinguishable, are With an (Bakker mecha- (an autogenic latter, allogenic and ecosystem approach of soil system. plant Salt-marshes Olff reconstructed 200 years of (1992) Here, plant species replacement of the parts of progress of gradient, probably course which nitrogen accumulation of succession the relative supposed was be could to short in succession than had the to in the of higher observed rate for the replaced were of study derived from studying the lower on clay. (an the the (an sea autogenic and as responses In result influences of understanding a mechanism) allogenic develop- geomorphological species The allogenic succession. species, taller autogenic case evidence for more be faster vegetation by competition this to of sedimentation of sediments from the clay light in retrospect, been was appeared responsible Though provided ecosystem of coastal bar island salt-marshes. development succession plant species importance increase. distinguished of the ment due rather than the desalination cause) the during and vegetation competitive abilities. Mire systems Zobel the reviewed (1988) ‘autogenic term processes bog succession in boreal mires. He succession’ or ‘self-development rather than vegetation succession in itself. He comparison with mineral land deterministic (though not communities, bog tiate between external and internal factors. determine autogenic processes, for peat formation processes, e.g. (Wheeler 1992) Hornets (1984) development logical and over a wider example area, it is when the water table or when a showed may conditions allogenically occur of a in bog bog it is likely rising very Sphagnum it difficult sometimes may for we comparable dealing are faster than the However, difficult growth. to without and by to that, in a more differen- possible to sequences in with are allogenic changes encountered. autogenic knowledge distinguish Small of peat could accumulate drained and unusual sequences allogenic seem use ecosystem is characterized where it is that regards the conclusion came to searching development simultaneously. induced also was bog has been that Though when mires’ succession unidirectional) development, showed that his clearly of between in the types of the of hydro- autogenically hydrological MECHANISMS regime al. OF SUCCESSION 427 may result in acidification of the top Wirdum layer (Van Van 1991; Diggelen et 1991). succession Bog and drain may structures perform of growing contribute to respect processes flow, water of random a regime, the The the terms arise communicate which functions with of of succession, discussions species moss autogenic on undirectional in met Masing (1987) put it: is it e.g. caused changes the ‘In unidirectional a and hydrologists feedback mechanisms with by carried be the bogs autogenic certain a should literature, of case or allogenic or type of research, because it should be simultaneously As Zobel & these the continuous explain to autogenic terms structures hydrological hydrological Phytocentric frequently are mire. easily can that so formation, structural characteristics of individual on nature complex which bog hydrological by identifying spatial scales. discussions about the or a the in ecologist species. such entangle of parts based moss During expressed An bog. various on be can such research to carpets even out the other development, ecosystem meaningless. flood or is actually become allogenic process disturbance considered as artificial’. In the above of ecosystem succession ecosystem appeared it examples, succession. could only another review (see that of ecosystem the study vegetation ‘In choosing that understand the to 1979). mechanisms of But these does enable an analysis of simulate a interest, is a one may focus on the this However, unit and limits one’s single size one of (1983): functions. We sometimes forcing that require emphasize understanding functions. forcing details of to would this as are meant to much as context vegetation succession, We quote from Odum of system a on system least at influences outside concentrates examples approach. interplay of pathways model and must one al. processes is reductionistic and procedure on receive additional attention, et all its by making succession in the study vegetation to does the mechanistic as the boundaries for unit of interest ability e.g. Gorham succession useful In the present paper than the larger say of one interest.’ REFERENCES Akkermans, nodulation conditions. Alexander, pathogen munities. Pests, (1990): interactions In: Burdon, and Scientific T.F.H. of nation of Leiden. J.J. natural & Plant and natural plant , (eds): 31-45. Oxford. Hierarchical B. & salt-marsh M.W. & data a space. in U. Brongers, (1990): M., Long-term communities by cattle 137-148. 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