Moving from pattern to process: coexistence mechanisms under intermediate disturbance regimes

Coexistence mechanisms that require environmental variation to operate contribute importantly to the maintenance of biodiversity. One famous hypothesis of diversity maintenance under disturbance is the intermediate disturbance hypothesis (IDH). The IDH proposes patterns of peaked diversity under intermediate disturbance regimes, based on a tension between competitively superior species and species which can rapidly colonize following disturbance. We review the literature, and describe recent research that suggests that more than one underlying mechanism can generate this unimodal diversity pattern in disturbed environments. Several exciting emerging research areas are identified, including interactions between disturbance types, operation of the IDH in multi‐trophic systems, and changes in disturbance regimes. However, empirical work is still focussed on describing the IDH pattern, with little emphasis on identifying its mechanistic basis. We discuss how to extend methods for identifying different coexistence mechanisms, developed in the theoretical literature, to experimental research. In an attempt to operationalize these various ideas we outline a hypothetical IDH research programme. A solid understanding of the life history attributes of the component species and their responses to disturbance will facilitate identification of the coexistence mechanism(s) underlying the IDH pattern, and provide a framework by which empirical and theoretical results can be more fully integrated.     

The effect of colonization and competition processes on the relation between disturbance and diversity in plant communities

Many theoretical and field studies have emphasized the impact of disturbance in the dynamics and diversity of sessile organism communities. This view is best reflected by the Intermediate Disturbance Hypothesis (IDH), which states that a maximum of diversity is found in ecosystems or communities experiencing intermediate disturbance regimes or at an intermediate stage of development since the last major disturbance event. Although theoretical models based on competitive interactions tend to validate this hypothesis, a recent meta-analysis of field experiments revealed that the mono-modal relationship between disturbance and diversity might not be a general pattern. In this article, we investigate the relationship between disturbance and diversity through the study of patch models, combining two types of competitive interactions: with or without competitive hierarchy, with two mechanisms influencing colonization: negative frequency dependence in colonization rates and immigration. These combinations led to various disturbance-diversity patterns. In the model without competitive hierarchy (founder effect model), a decreasing relationship appeared to be the rule as mentioned in previous studies. In the model with competitive hierarchy, the IDH pattern was obtained for low frequency dependence and low immigration. Nevertheless, high negative frequency dependence in colonization rates led to a decreasing relationship between disturbance and diversity. In contrast, high immigration led to an increasing relationship. The coexistence window (the range of disturbance intensity allowing coexistence) was the widest for intermediate immigration rates. For random species assemblages, patterns with multiple peaks were also possible. These results highlight the fact that the mono-modal IDH pattern should not be considered a rule. Competition and colonization mechanisms have a profound impact on the relationship between disturbance and diversity.     

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.     

The intermediate disturbance hypothesis applies to tropical forests, but disturbance contributes little to tree diversity

The intermediate disturbance hypothesis (IDH) predicts local species diversity to be maximal at an intermediate level of disturbance. Developed to explain species maintenance and diversity patterns in species-rich ecosystems such as tropical forests, tests of IDH in tropical forest remain scarce, small-scale and contentious. We use an unprecedented large-scale dataset (2504 one-hectare plots and 331 567 trees) to examine whether IDH explains tree diversity variation within wet, moist and dry tropical forests, and we analyse the underlying mechanism by determining responses within functional species groups. We find that disturbance explains more variation in diversity of dry than wet tropical forests. Pioneer species numbers increase with disturbance, shade-tolerant species decrease and intermediate species are indifferent. While diversity indeed peaks at intermediate disturbance levels little variation is explained outside dry forests, and disturbance is less important for species richness patterns in wet tropical rain forests than previously thought.     

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 Intermediate Disturbance Hypothesis and its applicability to planktonic communities: Comments on the views of Padisak and Wilson

The relevance of Connell's Intermediate Disturbance Hypothesis (IDH) to explanations of diversity and co-existence among plant species generally and in the phytoplankton in particular has been debated recently. Compared to terrestrial vegetation, planktonic communities experience distorted time and space scales, Generation times are in the order of days, not years to decades, Advective fluid transport raises the critical patch size to the order of kilometers, Within these scales, species survival and growth, community assembly and successional development in the phytoplankton conform to all the standards (compositional, strategic, thermodynamic) of conventional community organisation. These processes are known to move toward competitively excluded outcomes, Equally, they are liable to be interrupted by externally imposed disturbances, which reset the succession or alter its potential outcome, These findings are not only illustrative of intermediate disturbance but are instructive in the nature of diversity- disturbance relationships generally, IDH has considerable potential to explaining persistent species co-existence.     

Patch dynamics integrate mechanisms for savanna tree–grass coexistence

Many mechanisms have been suggested to explain the coexistence of woody species and grasses in savannas. However, evidence from field studies and simulation models has been mixed. Patch dynamics is a potentially unifying mechanism explaining tree–grass coexistence and the natural occurrence of shrub encroachment in arid and semi-arid savannas. A patch-dynamic savanna consists of a spatial mosaic of patches. Each patch maintains a cyclical succession between dominance of woody species and grasses, and the succession of neighbouring patches is temporally asynchronous. Evidence from empirical field studies supports the patch dynamics view of savannas. As a basis for future tests of patch dynamics in savannas, several hypotheses are presented and one is exemplarily examined: at the patch scale, realistically parameterized simulation models have generated cyclical succession between woody and grass dominance. In semi-arid savannas, cyclical successions are driven by precipitation conditions that lead to mass recruitment of shrubs in favourable years and to simultaneous collapse of shrub cohorts in drought years. The spatiotemporal pattern of precipitation events determines the scale of the savanna vegetation mosaic in space and time. In a patch-dynamic savanna, shrub encroachment is a natural, transient phase corresponding to the shrub-dominated phase during the successional cycle. Hence, the most promising management strategy for encroached areas is a large-scale rotation system of rangelands. In conclusion, patch dynamics is a possible scale-explicit mechanism for the explanation of tree–grass coexistence in savannas that integrates most of the coexistence mechanisms proposed thus far for savannas.     

Colonization–extinction dynamics of epixylic lichens along a decay gradient in a dynamic landscape

Metapopulation models are often used for understanding and predicting species dynamics in fragmented landscapes. Several models have been proposed depending on e.g. the relative importance of patch dynamics on the metapopulation dynamics. Dead wood is a dynamic substrate patch, and species that are confined to such patches have experienced a high degree of habitat loss in managed forests. Little is, however, known about how the population dynamics of epixylic species are affected by the fast dynamics of their substrate patches. We quantified the effect of local patch conditions and metapopulation processes on colonizations and extinctions of epixylic lichen species in a managed boreal forest landscape. This was done by twice surveying seven lichen metapopulations on 293 stumps in 30 stands of ages covering the duration of the dynamic patches (stumps). We also investigated the relative importance of local stochastic extinctions from stumps that remained available, and deterministic extinctions due to stump surface disappearance. We found importance of a decay gradient, surrounding metapopulation size, and local population sizes, in driving the colonization–extinction dynamics of epixylic lichens. The species were sorted along the stump decay gradient. Increasing surrounding metapopulation size was associated with increased colonization rates, and increasing local population size decreased lichen extinction rates. Finally, both local stochastic extinctions and deterministic extinctions due to patch disappearance occur, confirming that the long‐term persistence of epixylic lichens depends on colonization rates that compensate for stochastic population extinctions as well as deterministic extinctions.     

Neutral communities may lead to decreasing diversity-disturbance relationships: insights from a generic simulation model

Many attempts have been made to confirm or reject the unimodal relationship between disturbance and diversity stated by the intermediate disturbance hypothesis (IDH). However, the reasons why the predictions of the IDH apply or fail in particular systems are not always obvious. Here, we use a spatially explicit, individual-based community model that simulates species coexistence in a landscape subjected to disturbances to compare diversity-disturbance curves of communities with different coexistence mechanisms: neutrality, trade-off mechanism and intraspecific density dependence. We show that the shape of diversity-disturbance curves differs considerably depending on the type of coexistence mechanism assumed: (1) Neutral communities generally show decreasing diversity-disturbance curves with maximum diversity at zero disturbance rates contradicting the IDH, whereas trade-off communities generally show unimodal relationships confirming the IDH and (2) density-dependent mechanisms do increase the diversity of both neutral and trade-off communities. Finally, we discuss how these mechanisms determine diversity in disturbed landscapes.     

Disentangling spatiotemporal dynamics in metacommunities through a species-patch network approach

Colonization and extinction at local and regional scales, and gains and losses of patches are important processes in the spatiotemporal dynamics of metacommunities. However, analytical challenges remain in quantifying such spatiotemporal dynamics when species extinction-colonization and patch gain and loss processes act simultaneously. Recent advances in network analysis show great potential in disentangling the roles of colonization, extinction, and patch dynamics in metacommunities. Here, we developed a species-patch network approach to quantify metacommunity dynamics including (i) temporal changes in network structure, and (ii) temporal beta diversity of species-patch links and its components that reflect species extinction-colonization and patch gain and loss. Application of the methods to simulated datasets demonstrated that the approach was informative about metacommunity assembly processes. Based on three empirical datasets, our species-patch network approach provided additional information about metacommunity dynamics through distinguishing the effects of species colonization and extinction at different scales from patch gains and losses and how specific environmental factors related to species-patch network structure. In conclusion, our species-patch network framework provides effective methods for monitoring and revealing long-term metacommunity dynamics by quantifying gains and losses of both species and patches under local and global environmental change.     

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.     

集合群落及其在河流鱼类群聚研究中的应用

集合群落(metacommunity)是指多个潜在相互作用的物种通过它们之间的扩散而连接在一起的一组局域群落,目前已成为斑块生境下生物群落结构、格局和动态的重要理论基础之一。斑块动态、物种排序、群体效应和中性模型等4种理论模型,可用于解释不同情形下集合群落内物种的迁移状况,描述集合群落的动态。可采用群落结构或生态学机制等途径,来阐述所研究的群落是属于哪一种特定的集合群落类型。集合群落可用于研究河流鱼类群聚,解释鱼类的群落结构等问题。另外本文还结合我国水域生态环境及水生生物现状,对今后集合群落的研究作了展望。     

Central Die-back of Monoclonal Stands of Reynoutria japonica in an Early Stage of Primary Succession on Mount Fuji

Reynoutria japonica is a common perennial pioneer species onJapanese volcanoes. In a volcanic desert (1500m above sea level)on Mount Fuji (3776m), central Japan, this species forms circularstands (patches). As a patch develops, shoot density decreasesin its centre (‘central die-back’). The centraldie-back has been considered a key process in the early stagesof primary succession, though its mechanism has been unknown. The pattern of patch development, population dynamics of aerialshoots, and growth patterns of below-ground organs were analysedin order to investigate the mechanism of die-back, and the followingtraits are clarified: (1) central die-back areas occur in mostsmall patches (approx. 1m²) without later successional species;(2) shoot characteristics are dependent both on their positionwithin a patch and on patch size; (3) despite the large differencesin shoot density, neither time course of shoot growth nor theirmortality differs between the centre and periphery of patches;and (4) rhizomes ofR. japonicagrow outwards with regular sympodialbranching. From these results, it is concluded that neither interspecificnor intraspecific competition is likely to be a primary causeof the die-back phenomenon, but that central die-back is broughtabout intrinsically by the growth pattern of the rhizome systems.We also discuss the importance of the central die-back in facilitatingestablishment of later successional species in the early stagesof primary succession. Clonal plant; central die-back; competition; facilitation; Japanese knotweed; Mount Fuji; primary succession;Reynoutria japonica ; rhizome growth; volcanic desert     

Local sex-ratio dynamics: a model for the dioecious liverwort Marchantia inflexa

In many dioecious bryophyte species, population sex ratios range from all female to all male. The focal species of the present study, the liverwort Marchantia inflexa, forms patches on rock and bark surfaces, and these differ widely in sex ratio at a rainforest field site in Trinidad. This analysis – to our knowledge the first modeling study of sex-ratio dynamics in a dioecious clonal organism – addresses abundances of male and female M. inflexa through time within an individual patch. We represent the life history of this species using seven different stages (non-reproductive, asexually reproductive, sexually reproductive males, non-reproductive, asexually reproductive, unfertilized and fertilized sexual females) and express their dynamics using ordinary differential equations. Some of the stages become more abundant as thalli extend over the substrate and may overgrow each other to capture space. Our simple representation of dynamics within the patch failed to stabilize the sex ratio: females gradually eliminated males at low to moderate disturbance frequency and males eliminated females at high disturbance frequency. This pattern did not hinge on whether sexual propagules could germinate within the patch, but asexual reproduction (via gemmae dispersed within the patch) played an important role. This suggests that the maintenance of sex in these populations may hinge on metapopulation structure and dynamics. Though sexual reproduction appears to be unimportant within patches, spores provide the primary means of recolonizing patches eliminated by large-scale disturbances. We found that shortly after the patch was fully occupied, the production of these wind-dispersed spores was maximized, but spore production declined thereafter as the sex ratio became increasingly biased toward one sex or the other. Much additional modeling and empirical work is needed to link within-patch dynamics across patches and account for dynamics at the metapopulation level.     

Forests affected by frequent and intense typhoons challenge the intermediate disturbance hypothesis

Tropical cyclones (hurricanes and typhoons) are extreme disturbances that have a significant impact on ecosystem structure and processes. The intermediate disturbance hypothesis (IDH) generalizes disturbance–diversity relationships but its validity is hotly debated. The IDH has been challenged both theoretically and with results from experimental studies; however, few studies have empirically tested the proposed mechanisms of IDH using results from the actual ecosystems. In this commentary, based on empirical observations from studies on the interactions between tropical cyclones and forest dynamics, we outlined seven possible outcomes of gap dynamics and tree diversity as a result of different frequency and intensity combinations. We argue that the lack of distinction and differentiation between disturbance intensity and severity, and the overlooked role of tree mortality, seedling recruitment, and tree adaptations limit the applicability of IDH in predicting the disturbance–diversity relationship. In the era of climate change characterized with more frequent climate extremes and natural disturbances, we should move beyond the generalizations and directly address the processes leading to the observed disturbance–diversity relationships to make reliable predictions. Abstract in Chinese is available with online material.     

The component community structure of larval trematodes in the pulmonate snail Helisoma anceps

Factors that affected the component community structure of larval trematodes in the pulmonate snail Helisoma anceps in Charlie's Pond, North Carolina, were studied over a 15-mo period using a multiple mark-recapture protocol. Patent infections of 8 species were observed in 1,485 of 4,899 snails examined. Reproductive activity, population size, and survival rate of the snail population were estimated to evaluate the extent of resource availability for the parasites. Antagonistic interactions between trematode species that occurred at the infracommunity level had a neglible effect on the composition and structure of the component community. The patterns observed at this level were related to temporal heterogeneity in the abundance of infective stages (mostly miracidia), differential responses of trematode species to the diverse and constantly changing distribution of snail size and abundance, differential mortality of snails infected with certain trematode species, constant recruitment of 1 trematode species over time, and the existence of predictable disturbances such as the complete mortality of the host population and recruitment of a replacement population during a 6-8 wk period. The last factor operated as a reset mechanism for this snail-trematode system once each year. A model of patch dynamics, with snails as patch resources, best explains the organization and dynamics of this system.     

Patch quality and context, but not patch number, drive multi-scale colonization dynamics in experimental aquatic landscapes

Colonization and extinction are primary drivers of local population dynamics, community structure, and spatial patterns of biological diversity. Existing paradigms of island biogeography, metapopulation biology, and metacommunity ecology, as well as habitat management and conservation biology based on those paradigms, emphasize patch size, number, and isolation as primary characteristics influencing colonization and extinction. Habitat selection theory suggests that patch quality could rival size, number, and isolation in determining rates of colonization and resulting community structure. We used naturally colonized experimental landscapes to address four issues: (a) how do colonizing aquatic beetles respond to variation in patch number, (b) how do they respond to variation in patch quality, (c) does patch context affect colonization dynamics, and (d) at what spatial scales do beetles respond to habitat variation? Increasing patch number had no effect on per patch colonization rates, while patch quality and context were critical in determining colonization rates and resulting patterns of abundance and species richness at multiple spatial scales. We graphically illustrate how variation in immigration rates driven by perceived predation risk (habitat quality) can further modify dynamics of the equilibrium theory of island biogeography beyond predator-driven effects on extinction rates. Our data support the importance of patch quality and context as primary determinants of colonization rate, occupancy, abundance, and resulting patterns of species richness, and reinforce the idea that management of metapopulations for species preservation, and metacommunities for local and regional diversity, should incorporate habitat quality into the predictive equation.     

Temporally explicit habitat ecology and the coexistence of species

Habitats may have dynamics that exist independently of the population densities of species occupying the habitat. For example, ephemeral habitat patches may disappear regardless of whether a particular species is present or not. Such habitat dynamics are frequently modelled by ignoring age-related variation in patch turnover rates. This can be thought of as a temporally implicit approach. An alternative, temporally explicit approach involves using age-structured models in order to describe variations in habitat dynamics. Simple models of coexistence between competing species show that temporally implicit models may be misleading where there is age-related variation in patch dynamics. Changing the shape of the patch survivorship function but not the average patch survivorship can result in mutual extinction, monocultures or coexistence of an inferior and a superior competitor. An explicit treatment of habitat demography may therefore offer improved predictive models and alternative landscape management strategies.     

Patch dynamics and metapopulation theory: the case of successional species

We present a mathematical framework that combines extinction-colonization dynamics with the dynamics of patch succession. We draw an analogy between the epidemiological categorization of individuals (infected, susceptible, latent and resistant) and the patch structure of a spatially heterogeneous landscape (occupied-suitable, empty-suitable, occupied-unsuitable and empty-unsuitable). This approach allows one to consider life-history attributes that influence persistence in patchy environments (e.g., longevity, colonization ability) in concert with extrinsic processes (e.g., disturbances, succession) that lead to spatial heterogeneity in patch suitability. It also allows the incorporation of seed banks and other dormant life forms, thus broadening patch occupancy dynamics to include sink habitats. We use the model to investigate how equilibrium patch occupancy is influenced by four critical parameters: colonization rate, extinction rate, disturbance frequency and the rate of habitat succession. This analysis leads to general predictions about how the temporal scaling of patch succession and extinction-colonization dynamics influences long-term persistence. We apply the model to herbaceous, early-successional species that inhabit open patches created by periodic disturbances. We predict the minimum disturbance frequency required for viable management of such species in the Florida scrub ecosystem.