Coppice swidden fallows in tropical deciduous forest: Biological,technological, and sociocultural determinants of secondary forest successions

Secondary succession in tropical deciduous forest is often characterized by vegetative reproduction, or coppicing. Coppicing is also observed in forest sites that are disturbed by farming activity. This observation raises questions about the role of established management practices in determining the succession of vegetation on farmsites once they are abandoned to fallow. To what extent is the coppicing succession the result of specific aspects of swidden farming technology and management? And what variations in coppice successions occur in swidden sites following deviations from the standard farming practice? In research on swidden farming among the Susu of Sierra Leone, I examined the successional pathway in an age series of forest fallow sites. I show that the standard pattern of minimal cultivation favors the coppicing of felled trees in the subsequent fallow periods. By contrast, deviation from this pattern results in stump deaths and favors the invasion of fallow sites by grasses and vegetatively reproducing pioneer trees from the savanna. Variations in the environmental outcome of disturbance to plant communities, then, are the result of interactions between processes of tropical tree reproduction and the agricultural practices of local farmers.     

Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau

Background and Aims

Information on soil seed bank processes is crucial for understanding vegetation dynamics. Despite the documented importance of soil seed banks in many ecosystems, their role is not fully understood in some sensitive habitats, such as the alpine meadows of the Tibetan Plateau.

Methods

We studied the seasonal dynamics of the germinable soil seed bank under four disturbance intensities in an alpine meadow on the Tibetan Plateau as well as seed size distribution relative to disturbance intensity. Composition of the seed bank was compared with that of the standing vegetation.

Results

Density of buried seeds increased with disturbance intensity, but species richness and species diversity decreased. Seed density and species richness of the seed bank varied seasonally in all layers (0–2, 2–7, 7–12 cm) and the whole (0–12 cm). The species composition of seed bank was not significantly influenced by season. There was no trend in seed size distribution as disturbance increased. Seasonal seed bank turnover rates increased with increase in disturbance. The result of the NMDS showed that species composition of seed bank and vegetation exhibited a fairly uniform pattern in each season.

Conclusions

Although as a whole the species composition of the vegetation and seed bank showed a relatively low degree of similarity in each season, similarity was highest in the most disturbed habitat. There was no alteration in species composition of seed bank regardless of disturbance intensity, but seed density decreased as disturbance increased. Disturbances in alpine plant communities might increase persistence of regeneration niches. Regeneration from the seed bank together with vegetative reproduction contributed to aboveground vegetation in highly disturbed habitats. Clonal species played an important role in regeneration of vegetation in slightly disturbed areas, where there was little contribution of ruderals from soil seed banks.     

Untangling the causes of vegetation change in a 5,300 year pollen record at Tiniroto Lakes,Gisborne, New Zealand

To address potential causes of disturbances in recent New Zealand vegetation a sediment core from Tiniroto Lakes near Gisborne was used to reconstruct the local history of ecological disturbance and vegetation dynamics. Our approach was to examine the pattern and rate of vegetation change, against known disturbances, in order to identify different causes of disturbance. Despite intermittent disturbances, a steady transformation of vegetation dominates the period from c. 4,900 to c. 2,300 cal. yrs b.p. This is a time of climatic amelioration, with increasing precipitation suggested by the decline of light-adapted taxa, together with the establishment of forest. After c. 2,300 cal. yrs b.p., vegetation change becomes much more irregular, and apparently driven by disturbances, including unusual ones, such as earthquakes. In contrast to earlier disturbances, later vegetation responses are typified by a reduction of forest species and the establishment of semi-open vegetation, which persists for decades. This dichotomy suggests that a change in disturbance regime, especially in terms of fire, characterises the period after c. 2,300 cal. yrs b.p. The rises of fire frequency and of intensity at that time could be a result of severe droughts under climate extremes associated with intensified ENSO frequencies.     

Grassland type and seasonal effects have a bigger influence on plant functional and taxonomical diversity than prairie dog disturbances in semiarid grasslands

1. Prairie dogs (Cynomys sp.) are considered keystone species and ecosystem engineers for their grazing and burrowing activities (summarized here as disturbances). As climate changes and its variability increases, the mechanisms underlying organisms'' interactions with their habitat will likely shift. Understanding the mediating role of prairie dog disturbance on vegetation structure, and its interaction with environmental conditions through time, will increase knowledge on the risks and vulnerability of grasslands.
2. Here, we compared how plant taxonomical diversity, functional diversity metrics, and community‐weighted trait means (CWM) respond to prairie dog C. mexicanus disturbance across grassland types and seasons (dry and wet) in a priority conservation semiarid grassland of Northeast Mexico.
3. Our findings suggest that functional metrics and CWM analyses responded to interactions between prairie dog disturbance, grassland type and season, whilst species diversity and cover measures were less sensitive to the role of prairie dog disturbance. We found weak evidence that prairie dog disturbance has a negative effect on vegetation structure, except for minimal effects on C4 and graminoid cover, but which depended mainly on season. Grassland type and season explained most of the effects on plant functional and taxonomic diversity as well as CWM traits. Furthermore, we found that leaf area as well as forb and annual cover increased during the wet season, independent of prairie dog disturbance.
4. Our results provide evidence that grassland type and season have a stronger effect than prairie dog disturbance on the vegetation of this short‐grass, water‐restricted grassland ecosystem. We argue that focusing solely on disturbance and grazing effects is misleading, and attention is needed on the relationships between vegetation and environmental conditions which will be critical to understand semiarid grassland dynamics under future climate change conditions in the region.

     

Temporal patterns in seedling establishment on pocket gopher disturbances

Disturbances often facilitate seedling establishment, and can change the species composition of a community by increasing recruitment of disturbance-adapted species. To understand the effects of pocket gopher disturbances on alpine seedling dynamics, we examined the gopher disturbances effects on seedling emergence and survival on gopher disturbances 0 to 5 years old. In contrast to results from most other ecosystems, these recently created gopher mounds had lower seedling emergence and survival rates than undisturbed areas. A lack of correlation between species abundances on gopher mounds and undisturbed sites in one of the two communities studied suggested that a suite of disturbance-adapted species recruited onto the mounds. To explain low seedling emergence on recent gopher mounds, we quantified gopher mound seed banks and studied recruitment in a site with mounds that ranged from 0 to >20 years old. Seed numbers in first-year gopher mound soils were extremely low relative to undisturbed soils, and this pattern was mirrored in seedling establishment patterns over the long term. Gopher disturbance depressed seedling emergence density for the first 5 years. Subsequently, emergence density increased until at least 20 years following the disturbance. Emergence on disturbances more than 20 years old was higher than on undisturbed sites. Therefore, gopher disturbances probably facilitate seedling establishment in alpine dry and moist meadow; however, this process takes place over decades.     

The disturbance and vegetation dynamics: a review and an alternative framework

Disturbances are common phenomena that occurs in nature and significantly affects the dynamics of vegetation and plant populations. This paper shows that despite considerable progress in the theory of disturbances, for different authors the notion of disturbance has different meanings. The differences follow from an alternative understanding of the term, assumption of the holistic or reductionistic concept of the study or different approaches to equilibrium and non-equilibrium in nature. Different theoretical and methodological assumptions taken when analysing vegetation changes under the effect of disturbances allow relative arbitrariness in the interpretation of results. However, not all biotic interactions and ecological processes taking place in plant communities can be identified as disturbances. The effects of disturbances should be described on the basis of an objective assessment of what is normal for the functioning and stabilisation of a community as well as what disturbs and disrupts this equilibrium. Moreover, assuming the equivalent treatment of ecological systems, the interaction of the disturbing factors should be related to a specific level of life organisation (an individual, population or community), which would permit unambiguous discernment of the disturbance effects and finally generalise its effects on different levels of the hierarchical structure of vegetation.     

Response of two oligohaline marsh communities to lethal and nonlethal disturbance

Severity is recognized as an important attribute of disturbance in many plant communities. However, the effects of disturbances of different severity on patterns of regeneration in oligohaline marsh vegetation have not been experimentally examined. In these communities, a critical difference in the effects of disturbance severity may be whether the vegetation dies as a result of the disturbance or is merely damaged and hence capable of resprouting. We described the regeneration of vegetation in two Louisiana marsh community types, one dominated by Sagittaria lancifolia L. and the other by Spartina patens (Ait.) Muhl., following three levels of disturbance: no disturbance, a nonlethal disturbance, and a lethal disturbance. In the nonlethal disturbance, aboveground vegetation was clipped to simulate common disturbances such as fire and herbivory that remove aboveground vegetation but leave rhizomes intact. In the lethal disturbance vegetation was killed using herbicide to simulate disturbances causing plant mortality such as wrack deposition, sedimentation, scouring, and flooding following fire or herbivory. Regeneration was assessed over a 2-year period by measuring plant species richness, relative abundance, relative dominance, cover, and final biomass. To elucidate mechanisms for observed responses of vegetation, the species composition of the seed bank, light penetration, water level, salinity, and soil redox potential were evaluated. Despite differences in the structure of undisturbed vegetation in the two community types, they exhibited the same overall pattern of regeneration. Following nonlethal disturbance, the dominant species resprouted and quickly reestablished the structure of the vegetation. In contrast, recolonization following lethal disturbance occurred primarily via seedling recruitment, which resulted in marked shifts in community structure that persisted throughout the study. While the two communities responded similarly overall to disturbance, the response of individual species was not uniform; abundance, dominance, biomass, or cover increased for some species but decreased for others in response to disturbance. Seed bank species occurred in the vegetation following lethal disturbance in the Spartina community and in both disturbed and undisturbed plots in the Sagittaria community, indicating that the seed bank is a source of propagules for regeneration and maintenance of oligohaline marshes. Of the environmental variables measured, light level was most closely related to the effect of disturbance severity on community structure. Our results suggest that lethal and nonlethal disturbances have differential effects on regeneration of vegetation that can create pattern in oligohaline marshes communities. Received: 29 September 1997 / Accepted: 12 May 1998     

A review of similarity between seed bank and standing vegetation across ecosystems

The relationship between above and belowground species composition has been researched in forests, grasslands, and wetlands to understand what mechanisms control community composition. I thoroughly reviewed 108 articles published between 1945 and 2006 that summarized and provided specific values on similarities between above and belowground communities to identify common trends among ecosystems. Using Sørenson's index of similarity, I found that standing vegetation and its associated seed bank was the least similar in forest ecosystems, most similar in grasslands, and of intermediate similarity in wetlands. I also discovered that species richness was not related to seed bank – vegetation similarity in any of the three ecosystems. Disturbances were a common mechanism driving community composition in all ecosystems, where similarity decreased with time since disturbance in forest and wetland ecosystems and increased with time since disturbance in grasslands. Knowing the relationships between seed bank and standing vegetation may help conservationists to manage against exotic species, plan for community responses to disturbances, restore diversity, and better understand the resilience of an ecosystem.     

Disturbance and predation in an assemblage of herbivorous Diptera and algae on rocky shores

Summary Speculation about the effects of disturbance in marine benthic communities is often based on competition theory. Disturbances are thought to provision numerically depleted or competitively inferior species with resources associated with open substrate. However, disturbances that remove entire assemblages of sessile species also alter trophic structure, and thereby, influence the outcome of predator/prey relations. Aspects of community structure may be determined by patterns of disturbance and predation.The influence of disturbance and predation on the distribution and seasonality of blooms of ephemeral algae and associated Diptera was investigated with field experiments at several rocky beaches in central California. Blooms of ephemeral algae developed on high intertidal rock faces that were subject to severe seasonal disturbances caused by shifting sediment. These were subsequently colonized by the herbivorous larvae of several Diptera species for predictable periods each year. Other areas, without blooms, were not so disturbed.Experiments were done to determine if seasonal blooms were caused by seasonal disturbances that remove predators which otherwise might prevent the establishment of the Diptera/algae assemblage. The predators were crabs and limpets which eat both algae and larvae while foraging. Blooms of algae and larvae did not develop when limpets were transplanted to disturbed areas in periods between disturbances. Adjacent control areas did support blooms. Transplanted limpets did not survive periods of burial. When both limpets and crabs were excluded from treatment plots in undisturbed areas, blooms developed where they would not otherwise have occurred; controls remaied unchanged. Crabs and limpets differed in their effects on this assemblage. Crabs recruited quickly to the site of a bloom, but did not crop algal cover as closely, nor decrease larval density as much as the slowly recruiting limpets.The results suggest that disturbances favor blooms of some species by reducing predation. Severe localized disturbances increased the variability of the upper shore community by creating a patchwork of differing predator/prey abundances.     

Environmental fluctuations: How do they affect the topography of the adaptive landscape?

A Phenotypic Adaptive Landscape is defined with fitness as the ordinate, and longevity of the juvenile phase and duration of disturbances to the adult phase as the horizontal axes. The effect of local environmental perturbations on the landscape’s shape is studied using a semistochastic population model. In this model the intrinsic population dynamics takes the form of a differential-delay nonlinear equation and the environmental disturbance appears as a multiplicativetelegraphic noise. We demonstrate that the landscape has no single characteristic scale. Rather, it shows adaptive peaks corresponding to an integer relation between the biological and the environmental periodicities. Since the system is constrained by a finite time and a finite physiological range, the landscape may have different topographies for different local environmental regimes. A very simple fully deterministic model is presented, predicting landscapes that are similar to those obtained by the semistochastic model. Application to life history strategies are discussed.     

Regional productivity mediates the effects of grazing disturbance on plant cover and patch‐size distribution in arid and semi‐arid communities

Patch‐size distribution and plant cover are strongly associated to arid ecosystem functioning and may be a warning signal for the onset of desertification under changes in disturbance regimes. However, the interaction between regional productivity level and human‐induced disturbance regime as drivers for vegetation structure and dynamics remain poorly studied. We studied grazing disturbance effects on plant cover and patchiness in three plant communities located along a regional productivity gradient in Patagonia (Argentina): a semi‐desert (low‐productivity community), a shrub‐grass steppe (intermediate‐productivity community) and a grass steppe (high‐productivity community). We sampled paddocks with different sheep grazing pressure (continuous disturbance gradients) in all three communities. In each paddock, the presence or absence of perennial vegetation was recorded every 10 cm along a 50 m transect. Grazing effects on vegetation structure depended on the community and its association to the regional productivity. Grazing decreased total plant cover while increasing both the frequency of small patches and the inter‐patch distance in all communities. However, the size of these effects was the greatest in the high‐productivity community. Dominant species responses to grazing explained vegetation patch‐ and inter‐patch‐size distribution patterns. As productivity decreases, dominant species showed a higher degree of grazing resistance, probably because traits of species adapted to high aridity allow them to resist herbivore disturbance. In conclusion, our findings suggest that regional productivity mediates grazing disturbance impacts on vegetation mosaic. The changes within the same range of grazing pressure have higher effects on communities found in environments with higher productivity, markedly promoting their desertification. Understanding the complex interactions between environmental aridity and human‐induced disturbances is a key aspect for maintaining patchiness structure and dynamics, which has important implications for drylands management.     

Disturbances and species distribution of the riparian vegetation of a Rift Valley stream

The Njoro River riparian vegetation species composition, distribution, disturbances and uses are presented and discussed. Montane Juniperus procera-Olea europaea spp. africana and submontane Acacia abyssinica forests were identified as the main riparian vegetation groups. Approximately 55% of the riparian vegetation species are used for herbal medicine, treating more than 330 health problems, and only 11% of the plants are edible. Albizzia gummifera in the Syzygium cordatum-Pittosporum abyssinicum-Hibiscus diversifolius forest is cut selectively for herbal medicine preparations. Disturbances on the riparian vegetation zone are broadly classified as those induced by man, livestock and wildlife. Comprehensive effects of disturbance included loss of vegetation vertical strata, increase/decrease of species diversity, introduction of alien plant species, and reduction of plant sizes and vegetation hectarage. The effects of grazing on the vegetation were severe around livestock watering points. Grazing and browsing by wildlife were the main disturbances of the vegetation near the Njoro River estuary at the Lake Nakuru National Park. Periodic flooding, as a natural disturbance, regulates growth and survival of vegetation at the Lake Nakuru drawdown. Quantification of species diversity and the extent of disturbance by humans and livestock is important for future management of the vegetation and, consequently, the river.     

Recurrent habitat disturbance and species diversity in a multiple-competitive species system

To address how species interactions, dispersal and environmental disturbances interplay to affect the spatial distribution and diversity of species, we present a compartment model in which multiple species undergo competitive interaction of Lotka-Volterra type in a patchy environment arranged in a square lattice. Dispersal of species occurs between adjacent patches. Disturbances are periodically imposed on a central part of the environment in a belt-like block or an island-like block of various sizes where each species is killed for a certain time interval and then allowed to recover for the rest of a disturbance cycle. We deal with a case in which the local population dynamics within each patch is analytically determinable and has multiple locally stable equilibrium states in the absence of environmental disturbance. We further assume a trade-off between the reproductive rate of species and its dispersal ability. With these settings, we numerically examine how the spatio-temporal distributions of species are affected by changes in the pattern, size and duration of disturbances. The results demonstrate that: (1) in the undisturbed area, environmental disturbances could generate spatially segregated distributions of species; (2) in the disturbed area, species with higher dispersal abilities quickly invade and preferentially recover their population during the post-disturbance period, being temporarily relieved of competition from other species. These mechanisms collectively lead to increased species diversity in the whole habitat, functioning best when both the size and duration of disturbances are intermediate. In particular, the belt-like disturbance is more effective than the island-like disturbance in sustaining spatial heterogeneity for a wider range of duration of disturbance.     

Vegetation dynamics of Mediterranean shrublands in former cultural landscape at Grazalema Mountains,South Spain

Plant community dynamics in Mediterranean basin ecosystems are mainly driven by an alternation of episodes of human intervention and land abandonment. As a result, a mosaic of plant communities has evolved following different stages of degradation and regeneration. Some authors has relate secondary succession to abandoned culture lands and regeneration to natural systems with abandonment of livestock or forestry exploitation. In this paper, the dynamics of shrublands in mid-mountain areas in the South of Spain after disturbance and land abandonment has been studied. The plant cover and 13 environmental variables of 137 selected sites on the Grazalema mountains was analysed to determine the vegetation pattern in relation to environmental factors and the succession types, either regenerative or secondary succession. The results show that today the Grazalema mountains have a heterogeneous vegetation pattern. Besides physical factors such as altitude or soil , human disturbance has modulated current vegetation patterns and dynamics. Two main types of vegetation dynamics can be distinguished in the study area. In areas affected by cutting, regeneration results in rich and dense shrub land, with resprouters as dominant species. In areas affected by recurrent wildfires or agriculture, secondary succession became dominant, resulting in less diverse shrubland, due to the dominance of seeders and decrease in resprouter species richness and cover.     

How disturbance,competition, and dispersal interact to prevent tree range boundaries from keeping pace with climate change

Climate change is expected to cause geographic shifts in tree species' ranges, but such shifts may not keep pace with climate changes because seed dispersal distances are often limited and competition‐induced changes in community composition can be relatively slow. Disturbances may speed changes in community composition, but the interactions among climate change, disturbance and competitive interactions to produce range shifts are poorly understood. We used a physiologically based mechanistic landscape model to study these interactions in the northeastern United States. We designed a series of disturbance scenarios to represent varied disturbance regimes in terms of both disturbance extent and intensity. We simulated forest succession by incorporating climate change under a high‐emissions future, disturbances, seed dispersal, and competition using the landscape model parameterized with forest inventory data. Tree species range boundary shifts in the next century were quantified as the change in the location of the 5th (the trailing edge) and 95th (the leading edge) percentiles of the spatial distribution of simulated species. Simulated tree species range boundary shifts in New England over the next century were far below (usually <20 km) that required to track the velocity of temperature change (usually more than 110 km over 100 years) under a high‐emissions scenario. Simulated species` ranges shifted northward at both the leading edge (northern boundary) and trailing edge (southern boundary). Disturbances may expedite species' recruitment into new sites, but they had little effect on the velocity of simulated range boundary shifts. Range shifts at the trailing edge tended to be associated with photosynthetic capacity, competitive ability for light and seed dispersal ability, whereas shifts at the leading edge were associated only with photosynthetic capacity and competition for light. This study underscores the importance of understanding the role of interspecific competition and disturbance when studying tree range shifts.     

Grassland responses to multiple disturbances on the New England Tablelands in NSW,Australia

Disturbances have played a significant role in shaping vegetation patterns following European settlement and agricultural development in Australia, particularly over the last 100 years. However, little is known about the temporal dynamics of plant communities in relation to disturbances and their interactions. In this study we examined the response and recovery of temperate grassland communities to disturbance, using a multi-factorial experiment involving grazing exclusion (absent and present), fire (absent and present), soil cultivation (none, 5 and 20 cm) and soil amendment (none, fertiliser and fertiliser plus clover seeds) on the New England Tablelands in NSW, Australia. Temporal patterns of plant communities were analysed using detrended correspondence analysis for eight surveys over 24 months. Vegetation patterns at each survey were analysed using principal components analysis. The effects of treatments on malleability (Bray–Curtis dissimilarity) of plant communities were analysed using a linear mixed model, and the relationships between malleability and plant species groups were depicted using a generalised additive model and further analysed using a linear model. Perennial native grasses and a non-native forb (Hypochaeris radicata) initially dominated the vegetation, but after the disturbances H. radicata and other non-native species dominated. Compositional changes from the initial states were greatest in the first spring (7 months after treatment application), and then the vegetation tended to recover towards its original state. Soil cultivation resulted in the greatest deviation in community composition, followed by soil amendment, fire, with grazing exclusion the least. The recovery process and rate of recovery varied with treatment reflecting the dominance of soil cultivation and its interactions with other forms of disturbances. Soil amendment and grazing exclusion tended to reduce the effects of soil cultivation. Malleability was negatively related to perennial native grasses; positively to other non-native species, annual native grasses and perennial native sedges/rushes; and negatively to H. radicata when its cover was below 18%, but positively when above 18%. The degree of malleability reflected the high resilience of the vegetation to disturbance, and was mainly due to the recovery of perennial native grasses and H. radicata. This resilience demonstrated that the small-scale disturbances did not cause vegetation to cross an ecological threshold and that the present vegetation is resilient to common disturbances occurring at small scales. The results also suggest (1) that the present vegetation has developed mechanisms to adapt to these disturbances, (2) the importance of disturbance scale or (3) that the ecological threshold had already been crossed and the present vegetation is in a degraded state compared with its original state before the end of the 19th century.     

Quantifying patterns and controls of mire vegetation succession in a southern boreal bog in Finland using partial ordinations

Question: How do we distinguish between concurrent allogenic and autogenic forcings behind changing patterns in plant community structures during mire development? Location: Lakkasuo raised bog, southern Finland. Methods: Two radiometrically dated peat profiles were studied using high resolution plant macrofossil analysis. A combination of partial direct and indirect gradient analyses (CCA and DCA) was applied to quantify the role of different drivers of vegetation changes. Results: Autogenic hydroseral succession explained 16% of the compositional variation in the vegetation. Disturbance successions initiated by fire explained 15% of the variation in the hummock, but only 9% in the wetter lawn. The early post‐disturbance successional stages were characterized by Eriophorum vaginatum. After partialling out the effects of peat depth and time since fire, a moisture gradient explained 29% of variation in the hummock core and 26% in the lawn. The analyses also indicated alternation between species with a similar niche. This interaction gradient explained 26% and 31% of the compositional variation in the hummock and lawn, respectively. The similar order of species replacement from both cores supported the existence of general directional succession in mire vegetation, both during the mire development and after fire events. The autogenic succession was slow and gradual while the disturbance successions were episodic and fast. Conclusion: Our results support the paradigm of the complex nature of mire vegetation dynamics where several interlinked agents have simultaneous effects. The approach of combining partial ordinations developed here appeared to be a useful tool to assess the role of different environmental factors in controlling the vegetation succession.     

Effects of natural and artificial disturbance on landscape and forest structure in Tiantong National Forest Park,East China

This paper aims to understand the ecological effects of disturbance on broadleaved evergreen forest in East China. We used a manipulative field experiment approximating the common natural and artificial disturbance types in this area to investigate the community physiognomy, floristic composition, and 5-year recovery dynamics of the post-disturbance forest community. The results indicated that the landscape and forest structure have degraded into shrub communities, structure-damaged evergreen broadleaved communities, and so on. The post-disturbance communities presented different means of plant recruitment and vegetation recovery patterns at an early successional stage. The recovery of disturbed forests primarily depended on external seed sources and re-sprouting from stumps, rather than on soil seed banks, as few buried seeds were found. Re-sprouting thus appears to be key in allowing rapid vegetation recovery in evergreen broadleaved forest. Disturbances seem to be one of the most important factors that can contribute to regional species coexistence across temporal and spatial scales in evergreen broadleaved forests.     

An analysis of succession along an environmental gradient using data from a lawn

Differences in vegetation dynamics over a period of two years along an environmental gradient of shading on a lawn are examined. Communities (groups) recognized by numerical classification are correlated with degree of shading, season and differences between years by means of generalized linear model analysis. More of the variance is explained if environmental position is used instead of degree of shading as spatial distribution of a strongly competitive species (Trifolium repens) is confounded with shading. Transition probabilities are related to environmental position and season. Simulation with Markov matrices for each season and position demonstrate markedly different successions for different positions. These simulations have no predictive value however as accurate estimate of transition probabilities requires knowledge of the state of adjacent quadrats, i.e. individual observations of transition probabilities are not independent.Transition matrices are unlikely to be useful for predictive analysis of succession when spatial pattern is of significance in the community.We thank S. Kendall, S. Witts, A. Howard and C. Helman for their assistance with the data preparation and analysis, and R. Cunningham for advice on statistical analysis.