Pattern and process in the plant community: Fifty years after A.S. Watt

Abstract. This paper is a tribute to A.S. Watt who published his ‘Pattern and process in the plant community’ almost 50 years ago. Watt's interpretation of the plant community “as a working mechanism, which maintains and regenerates itself” is still highly relevant, although the keywords have changed. ‘Process’ in Watt's view involves both upgrade and downgrade aspects, whereas ‘Pattern’ was not specified, neither quantified. Nowadays, process is mainly approached as ‘disturbance’, that is natural disturbance and ‘pattern’ as patch structure. Together they make up the ‘patch dynamics’ of the community. Some implications of patch dynamics for phytosociology are discussed. A ‘Wattian’ concept of the plant community combines the Gleasonian idea of individualistic behaviour of species with the Clementsian (or rather Braun-Blanquetian) notion of community dynamics. Later work by Harper (demography), Grubb (regeneration niche) and earlier work of Sernander (forest gap dynamics) is significant for the understanding of the patch-dynamic nature of the community. Recent interest in plant species mobility can easily be linked to the concept of patch dynamics. Examples of mobility in a limestone grassland are given and a system of mobility types is proposed. Some perspectives for the study of patch dynamics are mentioned. Numerical pattern analysis should have a more prominent place in this type of study; the significance of the study of small permanent plots in a stand is emphasized, and unprejudiced demographic studies, as well as experimental studies of small-scale species replacement are recommended.     

The Dynamic keyhole-key model of coexistence to explain diversity of plants in limestone and other grasslands

Abstract. This paper is a tribute to A.S. Watt who published his ‘Pattern and process in the plant community’ almost 50 years ago. Watt's interpretation of the plant community “as a working mechanism, which maintains and regenerates itself” is still highly relevant, although the keywords have changed. ‘Process’ in Watt's view involves both upgrade and downgrade aspects, whereas ‘Pattern’ was not specified, neither quantified. Nowadays. process is mainly approached as ‘disturbance’, that is natural disturbance and ‘pattern’ as patch structure. Together they make up the ‘patch dynamics’ of the community. Some implications of patch dynamics for phytosociology are discussed. A ‘Wattian’ concept of the plant community combines the Gleasonian idea of individualistic behaviour of species with the Clementsian (or rather Braun-Blanquetian) notion of community dynamics. Later work by Harper (demography), Grubb (regeneration niche) and earlier work of Sernander (forest gap dynamics) is significant for the understanding of the patch-dynamic nature of the community. Recent interest in plant species mobility can easily be linked to the concept of patch dynamics. Examples of mobility in a limestone grassland are given and a system of mobility types is proposed. Some perspectives for the study of patch dynamics are mentioned. Numerical pattern analysis should have a more prominent place in this type of study; the significance of the study of small permanent plots in a stand is emphasized, and unprejudiced demographic studies, as well as experimental studies of small-scale species replacement are recommended.     

Role of disturbed vegetation in mapping the boreal zone in northern Eurasia

Question: Is there a need for disturbance mapping integrated in the CircumBoreal Vegetation Mapping Program? Location: Eurasian boreal forest. Disturbance and mapping: The boreal zone is characterized by a multitude of natural and anthropogenic disturbance agents with importance over a wide range of spatial and temporal scales. Disturbance is a prime driver of succession in most of the boreal zone, producing landscape diversity characterized by a large‐scale vegetation mosaic of early to late succession states. When mapping the circumboreal vegetation, spatial extent, time involved from disturbance to recovered condition and likelihood of interacting disturbance types are crucial for how current vegetation is interpreted and subsequently included as map characteristics. In this paper we present examples from the boreal zone where natural and/or anthropogenic disturbance regimes dominate the state and distribution of vegetation, and possibilities for assessing the nature and extent of the disturbed regions using remotely sensed data. Conclusion: Disturbed vegetation occupies large areas in the boreal zone and related vegetation successions should be adequately represented when mapping the zone. In regions where the ‘potential natural vegetation’ is a hypothetical reconstruction from remnants of ‘natural’ vegetation it would be preferable to use the concept of ‘actual real vegetation’ for which remote sensing at coarse, medium and fine resolution is an efficient tool. The Land Cover Classification System (LCCS) may offer sufficient flexibility to incorporate information about the disturbance of circumboreal vegetation.     

Beyond the patch: Disturbance affects species abundances in the surrounding community

The role of disturbance in community ecology has been studied extensively and is thought to free resources and reset successional sequences at the local scale and create heterogeneity at the regional scale. Most studies have investigated effects on either the disturbed patch or on the entire community, but have generally ignored any effect of or on the community surrounding disturbed patches. We used marine fouling communities to examine the effect of a surrounding community on species abundance within a disturbed patch and the effect of a disturbance on species abundance in the surrounding community. We varied both the magnitude and pattern of disturbance on experimental settlement plates. Settlement plates were dominated by a non-native bryozoan, which may have established because of the large amount of initial space available on plates. Percent covers of species within the patch were affected by the surrounding community, confirming previous studies' predictions about edge effects from the surrounding community on dynamics within a patch. Disturbance resulted in lower percent cover in the surrounding community, but there were no differences between magnitudes or spatial patterns of disturbance. Disturbance lowered population growth rates in the surrounding community, possibly by altering the abiotic environment or species interactions. Following disturbance, the recovery of species within a patch may be affected by species in the surrounding community, but the effects of a disturbance can extend beyond the patch and alter abundances in the surrounding community. The dependence of patch dynamics on the surrounding community and the extended effects of disturbance on the surrounding community, suggest an important feedback of disturbance on patch dynamics indirectly via the surrounding community.     

The Intermediate Disturbance Hypothesis of Species Coexistence Is Based on Patch Dynamics

The 'Intermediate Disturbance Hypothesis' (IDH) is one mechanism suggested to explain indefinite species coexistence. Hutchinson's original concept of the IDH was of a mechanism based on patch dynamics, and logical consideration shows that IDH works only if interpreted this way. Dependence on patch dynamics distinguishes IDH from Gradual Climate Change (GCC), though they are distinct also in terms of premature death of individuals, species selectivity, and the suddenness and transience of the perturbation. The application of the concepts of 'disturbance' and of IDH to phytoplankton communities is questioned.     

Unpacking the species conundrum: philosophy,practice and a way forward

The history of ecology and evolutionary biology is rife with attempts to define and delimit species. However, there has been confusion between concepts and criteria, which has led to discussion, debate, and conflict, eventually leading to lack of consistency in delimitation. Here, we provide a broad review of species concepts, a clarification of category versus concept, an account of the general lineage concept (GLC), and finally a way forward for species discovery and delimitation. Historically, species were considered as varieties bound together by reproduction. After over 200 years of uncertainty, Mayr attempted to bring coherence to the definition of species through the biological species concept (BSC). This has, however, received much criticism, and the last half century has spawned at least 20 other concepts. A central philosophical problem is that concepts treat species as ‘individuals’ while the criteria for categorization treats them as ‘classes’. While not getting away from this problem entirely, the GLC attempts to provide a framework where lineage divergence is influenced by a number of different factors (and correlated to different traits) which relate to the different species concepts. We also introduce an ‘inclusive’ probabilistic approach for understanding and delimiting species. Finally, we provide a Wallacean (geography related) approach to the Linnaean problem of identifying and delimiting species, particularly for cases of allopatric divergence, and map this to the GLC. Going one step further, we take a morphometric terrain approach to visualizing and understanding differences between lineages. In summary, we argue that while generalized frameworks may work well for concepts of what species are, plurality and ‘inclusive’ probabilistic approaches may work best for delimitation.     

Effects of prescribed burning and harvesting on ground-dwelling spiders in the Canadian boreal mixedwood forest

The ‘Natural Disturbance Paradigm’ for forest management seeks to meet conservation goals by modeling industrial harvest in fire-driven forest systems on patterns associated with wildfire. Fire suppression and increased forest harvesting may have detrimental effects on biodiversity, and therefore prescribed burning is suggested to retain legacies of wildfire not emulated under natural disturbance based approaches. The merits of this approach are being tested in the EMEND experiment in the Canadian boreal mixedwood forest. We compared responses of ground-dwelling spiders between sites subjected to prescribed post-harvest burning and retention harvest during three seasons during the first 7 years after disturbance. Overall, 38,661 adult spiders representing 190 species were collected. Estimated species richness was highest in undisturbed sites in all 3 years. Burning had the strongest negative effect on species richness 1–2 years after treatment; however, richness was higher in burns than in harvested sites 5–6 years post-disturbance. Species turnover was highest within controls but tended to increase over time between burned and harvested plots. Lower turnover in burned and harvested sites may reflect habitat homogenization by disturbance, suggesting a management and conservation challenge in relation to naturally disturbed and undisturbed areas. Species were grouped into disturbance-specialists, disturbance-tolerant, disturbance-generalists and generalists; 22 species were significant indicators for untreated sites, 18 for the burn and three for the harvest treatments. No major differences were observed in the spider fauna between harvested and burned areas within the first 6–7 years post-disturbance, and little evidence of recovery toward the pre-harvest fauna was evidenced. However, long term experiments may improve understanding of natural disturbance processes and improve management of boreal forests.     

On the integrated frameworks of species concepts: Mayden’s hierarchy of species concepts and de Queiroz’s unified concept of species

Richard L. Mayden and Kevin de Queiroz have devised and developed ‘a hierarchy of species concepts’ and ‘a unified species concept’, respectively. Although their integrated frameworks of species concepts are rather different as to how to integrate the diverse modern concepts of species, the end result is that they are likely to agree on species recognition in nature, because they virtually share the same major components (i.e. evolutionary or lineage concept of species; same way of delimiting species), and have the same important consequences. Both the hierarchical and unified frameworks, however, are interpreted to have shortcoming regarding the way of integrating the modern species concepts. I reformulate these ideas into a framework of species concepts as follows: It treats the idea of species as population‐level evolutionary lineages (sensu Wiley 1978 ) as the concept for species category, and it adopts the contingent biological properties of species (e.g. internal reproductive isolation, diagnosability, monophyly) as operational criteria in delimiting species. I also suggest that existing and revised versions of the integrated framework of species concepts all are not new species concepts, but versions of the evolutionary species concept, because they treat the evolutionary (or lineage) species concept as the concept for species category.     

An integrative approach to species delineation incorporating different species concepts: a case study of Limnadopsis (Branchiopoda: Spinicaudata)

The unambiguous delineation and identification of species remain central problems in systematic and taxonomic studies. Species delineation depends on the data utilized and the species concept applied. In recent years, morphology‐based species delineation has been complemented by DNA sequence data, leading to an integrative taxonomy. Such integrative approaches, however, are hampered by the partial incongruence of the various data types with certain species concepts. In this study, we delineated Australian Limnadopsis species employing one mitochondrial (cytochrome c oxidase subunit I, COI) and one nuclear (elongation factor 1α, EF1α) marker and a morphological character apparently part of the specific mate recognition complex, and therefore potentially indicative of reproductive isolation. By integrating the data over various species concepts (e.g. the ‘biological’, ‘Hennigian’, ‘recognition’, ‘phylogenetic’ and ‘evolutionary’ species concepts), the delineation of most species becomes straightforward and unambiguous. Conflicts are particularly interesting as they reveal different aspects of speciation considering the various species concepts. Our study emphasizes the benefits of a truly integrative approach to taxonomy. By combining molecular data with morphological characters indicative of reproductive isolation, it is possible to delineate species integrating not only different data types, but also different underlying species concepts. Overall, 11 Limnadopsis species could be delineated, including all eight currently recognized species, and three so far undescribed species. Most species were congruently delineated under all species concepts. A strict application of the evolutionary species concept, however, would have further split L. parvispinus into two species on the basis of the COI data. In addition, Limnadopsis tatei is consistently split into two sympatrically occurring species under all applied species concepts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 575–599.     

Trophic compensation reinforces resistance: herbivory absorbs the increasing effects of multiple disturbances

Disturbance often results in small changes in community structure, but the probability of transitioning to contrasting states increases when multiple disturbances combine. Nevertheless, we have limited insights into the mechanisms that stabilise communities, particularly how perturbations can be absorbed without restructuring (i.e. resistance). Here, we expand the concept of compensatory dynamics to include countervailing mechanisms that absorb disturbances through trophic interactions. By definition, ‘compensation’ occurs if a specific disturbance stimulates a proportional countervailing response that eliminates its otherwise unchecked effect. We show that the compounding effects of disturbances from local to global scales (i.e. local canopy‐loss, eutrophication, ocean acidification) increasingly promote the expansion of weedy species, but that this response is countered by a proportional increase in grazing. Finally, we explore the relatively unrecognised role of compensatory effects, which are likely to maintain the resistance of communities to disturbance more deeply than current thinking allows.     

A neutral terminology to define 'invasive' species

The use of simple terms to articulate ecological concepts can confuse ideological debates and undermine management efforts. This problem is particularly acute in studies of nonindigenous species, which alternatively have been called ‘exotic’, ‘introduced’, ‘invasive’ and ‘naturalised’, among others. Attempts to redefine commonly used terminology have proven difficult because authors are often partial to particular definitions. In an attempt to form a consensus on invasion terminology, we synthesize an invasional framework based on current models that break the invasion process into a series of consecutive, obligatory stages. Unlike previous efforts, we propose a neutral terminology based on this framework. This ‘stage‐based’ terminology can be used to supplement terms with ambiguous meanings (e.g. invasive, introduced, naturalized, weedy, etc.), and thereby improve clarity of future studies. This approach is based on the concept of ‘propagule pressure’ and has the additional benefit of identifying factors affecting the success of species at each stage. Under this framework, invasions can be more objectively understood as biogeographical, rather than taxonomic, phenomena; and author preferences in the use of existing terminology can be addressed. An example of this recommended protocol might be: ‘We examined distribution data to contrast the characteristics of invasive species (stages IVa and V) and noninvasive species (stages III and IVb)’.     

Interactive community responses to disturbance in streams: disturbance history moderates the influence of disturbance types

Both disturbance history and disturbance type act to structure communities through selecting for particular species traits but they may also interact. For example, flooding selects for species with flood‐resistant traits in streams, but those traits could make communities susceptible to other disturbances and so could cause shifts in community composition due to anthropogenic climate change. To better understand the interactive influences of disturbance history and type on community composition, we investigated the response of macroinvertebrate communities to disturbance using in‐stream channels. Using a split‐plot design, individual channels in five ‘stable’ streams and five ‘frequently disturbed’ streams (disturbance history) were subject to different disturbance type treatments (flooding, drying and a control). Disturbance type independently drove effects on species diversity, but all other effects of disturbance type depended on disturbance history. In particular, the interaction of disturbance type and history determined overall community response. Both disturbance types tested produced similar community responses in frequently disturbed streams, including changes in community composition and alterations to the abundance of less mobile taxa, but low‐flow had a significantly greater effect in stable streams. Macroinvertebrate drift was greatest in the rock‐rolling treatments and significantly less in the low‐flow treatment for both disturbance histories. Therefore, disturbance history moderated the effects of disturbance type and determined the mechanism of community response by determining how well species were adapted to disturbance. This outcome suggests that previous disturbances strongly influence how vulnerable communities are to changes in disturbance, and so should be considered when predicting how changes in disturbance regimes will affect future community composition.     

Rethinking patch size and isolation effects: the habitat amount hypothesis

I challenge (1) the assumption that habitat patches are natural units of measurement for species richness, and (2) the assumption of distinct effects of habitat patch size and isolation on species richness. I propose a simpler view of the relationship between habitat distribution and species richness, the ‘habitat amount hypothesis’, and I suggest ways of testing it. The habitat amount hypothesis posits that, for habitat patches in a matrix of non‐habitat, the patch size effect and the patch isolation effect are driven mainly by a single underlying process, the sample area effect. The hypothesis predicts that species richness in equal‐sized sample sites should increase with the total amount of habitat in the ‘local landscape’ of the sample site, where the local landscape is the area within an appropriate distance of the sample site. It also predicts that species richness in a sample site is independent of the area of the particular patch in which the sample site is located (its ‘local patch’), except insofar as the area of that patch contributes to the amount of habitat in the local landscape of the sample site. The habitat amount hypothesis replaces two predictor variables, patch size and isolation, with a single predictor variable, habitat amount, when species richness is analysed for equal‐sized sample sites rather than for unequal‐sized habitat patches. Studies to test the hypothesis should ensure that ‘habitat’ is correctly defined, and the spatial extent of the local landscape is appropriate, for the species group under consideration. If supported, the habitat amount hypothesis would mean that to predict the relationship between habitat distribution and species richness: (1) distinguishing between patch‐scale and landscape‐scale habitat effects is unnecessary; (2) distinguishing between patch size effects and patch isolation effects is unnecessary; (3) considering habitat configuration independent of habitat amount is unnecessary; and (4) delineating discrete habitat patches is unnecessary.     

Disturbance reinforces community assembly processes differentially across spatial scales

Background and AimsThere is a paucity of empirical research and a lack of predictive models concerning the interplay between spatial scale and disturbance as they affect the structure and assembly of plant communities. We proposed and tested a trait dispersion-based conceptual model hypothesizing that disturbance reinforces assembly processes differentially across spatial scales. Disturbance would reinforce functional divergence at the small scale (neighbourhood), would not affect functional dispersion at the intermediate scale (patch) and would reinforce functional convergence at the large scale (site). We also evaluated functional and species richness of native and exotic plants to infer underlying processes. Native and exotic species richness were expected to increase and decrease with disturbance, respectively, at the neighbourhood scale, and to show similar associations with disturbance at the patch (concave) and site (negative) scales.MethodsIn an arid shrubland, we estimated species richness and functional dispersion and richness within 1 m² quadrats (neighbourhood) nested within 100 m² plots (patch) along a small-scale natural disturbance gradient caused by an endemic fossorial rodent. Data for the site scale (2500 m² plots) were taken from a previous study. We also tested the conceptual model through a quantitative literature review and a meta-analysis.Key ResultsAs spatial scale increased, disturbance sequentially promoted functional divergence, random trait dispersion and functional convergence. Functional richness was unaffected by disturbance across spatial scales. Disturbance favoured natives over exotics at the neighbourhood scale, while both decreased under high disturbance at the patch and site scales.ConclusionsThe results supported the hypothesis that disturbance reinforces assembly processes differentially across scales and hampers plant invasion. The quantitative literature review and the meta-analysis supported most of the model predictions.     

Ethnonationalism,ethnic nationalism,and mini‐nationalism: A comparison of Connor,Smith and Snyder

Some terms have been coined to highlight the important relation between nationalism and ethnicity. However, there has been much confusion as to the meaning of such concepts as ‘ethnonationalism’, ‘ethnic nationalism’ and ‘mini‐nationalism’. This article compares the three terms, as used respectively in the works of Walker Connor, Anthony Smith and Louis Snyder. These scholars are selected not only because of their well‐respected position in the study of nationalism but also because their works are representative of each of the three concepts. The comparison finds that since both Connor and Smith emphasized the ethnic dimension of nationalism, their ideas can be presented in one analytical framework. In fact, Connor's ‘ethnonationalism’ and Smith's ‘ethnic nationalism’ overlap with each other. However, Smith's theory of nationalism is comprehensive enough to take care of the fact that the nationalisms of many new states of today's world have no ethnic base at all, whereas Connor's analysis makes no mention of this. In this respect, Smith's model is preferred to that of Connor. Finally, Snyder's ‘mini‐nationalism’ is seen as the least useful concept, since the use of size rather than ethnicity to classify nationalisms does not increase our understanding of the concept.     

Minireviews: Neighborhood Effects, Disturbance Severity, and Community Stability in Forests

A theoretical framework and conceptual model for temporal stability of forest tree-species composition was developed based on a synthesis of existing studies. The model pertains primarily to time periods of several tree lifetimes (several hundred to a few thousand years) at the neighborhood and stand spatial scales (0.01–10 ha), although a few extensions to the landscape scale are also made. The cusp catastrophe was chosen to illustrate compositional dynamics at the stand level for jack pine, northern hardwood, and white pine forests in the Great Lakes Region of the United States and for tropical rainforests in the northern Amazon basin. The models feature a response surface (degree of dominance by late-successional species) that depends on two variables: type of neighborhood effects of the dominant tree species and severity of disturbances. Neighborhood effects are processes that affect the chance of a species replacing itself at the time of disturbance (they can be positive, neutral, or negative) and are of two types: overstory–undestory effects, such as the presence of advanced reproduction; and disturbance-activated effects, such as serotinous seed rain. Disturbance severity is the proportion of trees killed during a disturbance. Interactions between neighborhood effects and disturbance severity can lead to either punctuated stability (dramatic but infrequent change in composition, in those forests dominated by species with positive neighborhood effects) or succession (continuous change, in those forests dominated by species with neutral-negative neighborhood effects). We propose that neighborhood effects are a major organizing factor in forest dynamics that provide a link across spatial scales between individual trees and disturbance/patch dynamics at the stand and landscape scales. Received 23 June 1998; accepted 16 December 1998.     

Hypertrophic phytoplankton and the Intermediate Disturbance Hypothesis

The provisions of Connell's Intermediate Disturbance Hypothesis (IDH) were investigated in relation to the behaviour of freshwater phytoplankton in a hypertrophic lagoon, paying special attention to the link between species-diversity and environmental disturbances. Phytoplankton diversity yielded different indices depending on the basic unit of calculations (cells, particles, phytoplankton units, biomass). Although their ranges were approximately equal, they did not covary so could not be considered mutually substitutable. For the purpose of IDH testing, biomass diversity was chosen.Equilibrium states were considered to obtain in those periods with a very high fraction of total phytoplankton biomass, shared by no more than three phytoplankton species. Disturbances were considered as counterparts of equilibrium states. Disturbance factors were mostly abiotic, environmental features of the lake operating on the phytoplankton community at different time scales (co-occurring and with 1- and 2-weeks' lags). These scales may relate to the time required to establish phytoplankton community structure.IDH could be suspected not to hold for the phytoplankton of this hypertrophic lake, which experienced seven near-equilibrium phases and six disturbance periods throughout the study. As a rule, equilibrium states lasted longer than disturbance periods. The expected relationships between both disturbance intensity or frequency were not shown. Furthermore, no relationship was demonstrated between diversity (and hence IDH) and the phytoplankton community change rate. Wind stress probably played a minor role in triggering disturbance events. Disturbances were shown partly to promote small-sized phytoplankton communities.Finally, a plea for studying hypertrophic phytoplankton in greater detail is stressed if its responses to disturbances are to be fully understood.     

Shade tolerance,canopy gaps and mechanisms of coexistence of forest trees

The belief that canopy gaps are important for the maintenance of tree species diversity appears to be widespread, but there have been no formal theoretical models to assess under what conditions gap phase processes allow coexistence. Much of the empirical research on niche differentiation in response to gaps has focused on evidence for an interspecific tradeoff between low light survival and high light growth. The objectives of this study are first to distinguish the possible mechanisms allowing coexistence based on this tradeoff, and second, to explore their limitations. We present a theory of forest dynamics driven by small‐scale disturbances as a special case of the theory of coexistence in variable environments. We demonstrate that temporal and spatial heterogeneity in light conditions that results from canopy gaps can allow stable coexistence as a result of three previously documented general mechanisms: ‘relative non‐linearity’, ‘the successional niche’ and the ‘storage effect’. We find that temporal fluctuations in light availability alone allow the stable coexistence of only two species. Spatial variation in disturbance synchronicity and intensity allows three species to coexist in a narrow parameter space. The rate of extinction is, however, extremely slow and there is transient coexistence of a larger number of species for a long period of time. We conclude that while the low light survival/high light growth tradeoff may be ubiquitous in forest tree species, it is unlikely to function as an important mechanism for the stable coexistence of several tree species.     

Low impact of disturbance on ecological success of invasive tree and shrub species in temperate forests

Disturbance is claimed to be one of the most important triggers of biological invasions. There is a lack of data about disturbance impacts on the youngest life stage of invasive trees and shrubs. Thus, we aimed to assess the role of disturbances in shaping responses of natural regeneration of three model invasive species—Prunus serotina Ehrh., Quercus rubra L., and Robinia pseudoacacia L.—to disturbances in forest plant communities. Our study was conducted over 3 years on 372 study plots (100 m²), across nine types of temperate forests in Wielkopolski National Park (Poland). Disturbance was assessed using ecological indicator values for disturbance severity and frequency. Our study revealed the high importance of disturbance on species composition of understory vegetation. We also found relationships between ecological success (density and biomass of natural regeneration) of invaders and disturbance indices. These models were statistically significant but their effect sizes were low. Due to the low effect sizes, we can state only limited conclusions about impact of disturbance on ecological success of invasive species natural regeneration. The results suggest that for seedlings (up to 50 cm height—threshold between understory and shrub layer) disturbance, a leading factor in biological invasions of numerous taxa, has a small role in this case. Thus, we may assume that their ecological success is connected with stochastic processes in populations of the invader’s seedlings, rather than with stochastic release from competition caused by disturbances.