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In addition to density-dependent (i.e. food availability) and independent (i.e. abiotic gradients) forces, size-selective harvests of organisms often induce changes to their life history, demography, and reproduction. Shellfish are increasingly overexploited worldwide, and the effect of size-selective harvest on their populations is unclear. The African bloody cockle Senilia senilis is a dominant bivalve in most West African coastal systems, where it is an important source of protein. Here, we studied its population structure (abundance, biomass, and size) and life history traits (age, annual growth rate, and size for age) under contrasting harvesting conditions at the 2 largest intertidal systems in the region: the Bijagós archipelago in Guinea-Bissau (harvested) and Banc d’Arguin in Mauritania (unexploited). To account for local ecological context, we assessed environmental variables that could also affect the abundance and size of S. senilis at these sites. We found that the S. senilis population at Bijagós is significantly denser and older, with lower standing biomass and smaller-sized shells than the population from Banc d’Arguin. In Bijagós, the annual growth is 3-fold lower, and the size-for-age is half of that at Banc d’Arguin. At both sites, S. senilis densities and sizes were correlated with habitat quality indicators: mangrove and mudflat productivity at Bijagós and seagrass and sediment characteristics at Banc d’Arguin. The smaller-sized mature individuals, denser populations, and lower standing biomass at Bijagós compared to Banc d’Arguin are indications of considerable size-selection pressure at Bijagós.

Item does not contain fulltext 01 april 2023

Along Africa’s western coast, many local communities rely on the ocean for their livelihood. Over the last decades, introductions of new fishing techniques along with globalizing trade have strongly changed local fishing practices. The Parc National du Banc d’Arguin (PNBA) in Mauritania had for centuries been subjected to an artisanal, low-impact, fishery. This fishing was exclusively oriented towards migratory bony fish species, mullet (Mugil cephalus) and meagre (Argyrosomus regius). Since the 1980s, these species have been replaced by illegal catches of internationally traded elasmobranchs (sharks and rays) and by non-migratory and relict species (resident) such as tilapias (Sarotherodon melanotheron) and catfishes ( Arius sp.). To date, most monitoring and management efforts have been dedicated to evaluating changes in elasmobranch populations and less focus has been on bony fish species. Data from a fishery monitoring programme are used to analyse the trends in effort, catch and catch per unit of effort of bony fish species by fitting non-parametric generalized additive models to capture changes in the fish community over the last 20 years. Mullet and meagre became overfished early on, and the contribution of resident species (tilapias and catfishes) increased in the catches. Together with a pattern of increased effort on the traditionally targeted species, such a change in the catch could reflect a change in the fish community. These results call for the implementation of sustainable fishing practices within PNBA. We propose the need to implement closures of fisheries during the species’ breeding periods as well as the use of biological reference points such as the size at first capture and maximum sustainable yield targets for resident species.

Determining the drivers of aboveground net primary production (ANPP), a key ecosystem process, is an important goal of ecosystem ecology. However, accurate estimation of ANPP across larger areas remains challenging, especially for savanna ecosystems that are characterized by large spatiotemporal heterogeneity in ANPP. Satellite remote sensing methods are frequently used to estimate productivity at the landscape scale but generally lack the spatial and temporal resolution to capture the determinants of productivity variation. Here, we developed and tested methods for estimating herbaceous productivity as an alternative to labour-intensive repeated biomass clipping and caging of small plots. We compared measures of three spectral greenness indices, normalized difference vegetation index derived from Sentinel-2 (NDVIs) and a handheld radiometer (NDVIg), and green chromatic coordinate derived from digital repeat cameras (GCC) and tested their relationship to biweekly field-measured herbaceous ANPP using movable exclosures. We found that a satellite-based model including average NDVIs and its rate of change (ΔNDVIs) over the biweekly productivity measurement interval predicted herbaceous ANPP reasonably well (Jackknife R2 = 0.26). However, the predictive accuracy doubled (Jackknife R2 = 0.52) when including the sum of day to day increases in camera trap-derived vegetation greenness (tGCC). This result can be considered promising, given the current lack of productivity estimation methods at comparable spatiotemporal resolution. We furthermore found that the fine (daily) temporal resolution of GCC time series captured fast vegetation responses to rainfall events that were missed when using a coarser temporal resolution (>2 days). These findings demonstrate the importance of measuring at a fine temporal resolution for predicting herbaceous ANPP in savanna ecosystems. We conclude that camera traps are promising in offering a reliable and cost-effective method to estimate productivity in savannas and contribute to a better understanding of ecosystem functioning and its drivers.

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Marine Protected Areas (MPAs) are increasingly implemented to facilitate the conservation of marine biodiversity and key habitats. However, these areas are often less effective to conserve mobile marine species like elasmobranchs (i.e., sharks and rays). Industrial fishing near MPA borders possibly impacts vulnerable species utilizing these protected areas. Hence, we aimed to study spatiotemporal patterns of industrial fisheries near MPAs, in relation to the bycatch of elasmobranchs. Specifically, we analyzed the spatiotemporal fishing effort within the West African region, mapped fishing effort in the direct vicinity of the Parc National du Banc d’Arguin (PNBA, Mauritania) and the Bijagós Archipelago (BA, Guinea Bissau), and compared the seasonal overlap between elasmobranch bycatch and fishing effort near these MPAs. We combined Automatic Identification System (AIS) data and local fisheries observer data, and determined fishing effort for each gear type and compared this with bycatch of elasmobranchs. We found that industrial fishing effort was dominated by trawling, drifting longlines, and fixed gear types. Although no industrial fishing was observed within both MPAs, 72 and 78% of the buffer zones surrounding the MPAs were fished for the Banc d’Arguin and Bijagós, respectively. Within the Banc d’Arguin buffer zone, trawling and drifting longlines dominated, with longlines mainly being deployed in fall. In the Bijagós buffer zone, trawling and fixed gears were most prevalent. Fisheries observer data for Mauritania showed that elasmobranch catches increased during the most recent sampling years (2016–2018). Elasmobranch catches within the waters of Guinea Bissau peaked in 2016 and decreased in the following two years. Seasonal patterns in elasmobranch bycatch within the waters of both countries are likely caused by increased catches of migratory species. Catches of rays peaked in May and June for Mauritania, and in October for Guinea Bissau. Shark catches were highest in February and July in Mauritanian waters, and in May and October in the waters of Guinea Bissau. Our study indicates that industrial fisheries near the border of ecologically important MPAs may have potentially major implications for ecosystem functioning by the removal of (migratory) predatory species.

The benthic communities of soft-sediment intertidal ecosystems trophically underpin the migration of birds and fish. Within the East Atlantic Flyway, along the coast of West-Africa, the intertidal mudflats of Banc d’Arguin, Mauritania, host over 2 million migratory waterbirds. Despite the protected status of the Banc d’Arguin, geographical remoteness and seemingly benign human exploitation, we show that large changes have taken place in the intertidal benthic macrofauna across an interval of 28 years. We compared the results of two comparable and spatially comprehensive large-scale benthic surveys in 1986 and 2014. Over this time, the benthos changed from a diverse community to one dominated by a few species of bivalve, with a loss of polychaete worms. The change was associated with a twofold increase in the seagrass cover. Our results suggest that the intertidal habitats of Banc d’Arguin have altered markedly over the last three decades and the estimated benthic secondary production has decreased by a factor of four. These shifts in community structure and production may have contributed to declines in some benthivorous migratory shorebirds.

The fluorochrome calcein has been used in numerous growth studies of molluscs to internally mark calcified structures. Because of interspecific variations in marking success and possible effects on growth performance, methodological assessments of the suitability of calcein as a growth marker, especially in field contexts, remain necessary. Here we report on the effects of different calcein concentrations (100, 200, 400 and 800 mg l(-1) on fluorescent mark deposition, growth rate, density, body condition and size-frequency distribution of an intertidal infaunal bivalve species, Loripes lacteus (Linnaeus, 1758), using an outdoor immersion technique. To avoid stress caused by handling and transportation, in situ enclosures were placed at seagrass-covered patches during low tide, to which calcein solutions were added. After a 1.3- to 2.6-h period of exposure to calcein, the enclosures were removed. Sites were sampled three months later. All calcein concentrations produced live L. lacteus with a clear fluorescent shell mark, but the percentage successfully marked tended to increase with higher calcein concentrations. Furthermore, marking success and growth rate decreased significantly with shell size (i.e., age). Calcein concentration did not measurably affect shell growth rate, body condition and size-frequency distribution, but the numerical densities were lower for sites treated with calcein concentrations >= 400 mg l(-1). Our results suggest that in situ calcein-marking of burrowing lucinid bivalves with low concentrations (100-200 mg l(-1)) is a non-invasive and rapid method to determine growth rate, provided that the lucinids are not too old. (c) 2011 Elsevier B.V. All rights reserved.

Mammalian herbivores can have pronounced effects on plant diversity but are currently declining in many productive ecosystems through direct extirpation, habitat loss and fragmentation, while being simultaneously introduced as livestock in other, often unproductive, ecosystems that lacked such species during recent evolutionary times. The biodiversity consequences of these changes are still poorly understood. We experimentally separated the effects of primary productivity and herbivores of different body size on plant species richness across a 10-fold productivity gradient using a 7-year field experiment at seven grassland sites in North America and Europe. We show that assemblages including large herbivores increased plant diversity at higher productivity but decreased diversity at low productivity, while small herbivores did not have consistent effects along the productivity gradient. The recognition of these large-scale, cross-site patterns in herbivore effects is important for the development of appropriate biodiversity conservation strategies. [KEYWORDS: Cross-site ; fertility ; grazing ; plant–animal ;species richness]

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Estimating the spatial position of birds in open habitats like intertidal mudflats is important for many studies, for example, detailed density estimates or linking predation pressure to resource availability. To date, several methods have been used to estimate the positions of birds, including density counts in predetermined plots, range finders, photography, and tracking individuals tagged with GPS-equipped transmitters, and each method has advantages and shortcomings. Counts in premarked plots are possible over relatively long distances, but small-scale information is lost due to within-plot averaging. Other methods accurately determine the position of individuals, but can only be used at relatively short distances or involve capturing birds. We describe a simple and low-cost method to estimate the spatial position of individual birds in open habitats using a telescope-mounted instrument that measures the scope's viewing angle. Using this Telescope-Mounted Angulator (TMA), the distance to focal birds can be calculated by simple trigonometry, requiring only the viewing angle and mounting height of the telescope. Laboratory tests revealed that the TMA was most accurate when calibrated for individual observers. Field experiments performed on a 4-m high observation platform showed that the TMA can estimate the position of shorebirds with an accuracy of 18 to 36 m up to a distance of 500 m. By also including the direction, determined with a compass, the spatial position of birds can be reliably estimated. The TMA can be a valuable tool for estimating the spatial position of animals in various flat landscapes, providing detailed measurements in a relatively short period of time.

The species abundance distribution is one of the few “universal” patterns in ecology. Research on this fundamental distribution has been focused to the study of numerical counts, irrespective of the traits of individuals. Here we show that considering a set of Generalized Species Abundance Distributions (GSADs) that encompass several abundance measures, specifically numerical abundance, biomass and resource use can provide novel insights into organizational forces at play in ecological communities. We first leverage a comprehensive dataset to investigate the similarity and difference between GSADs. We then use classical probability theory and allometric scaling laws to explore theoretical linkages among GSADs and between GSADs and other common macroecological distributions. Finally, we show how examining different GSADs in natural systems can help with assessing determinants of community structure. Broadening species abundance distributions to encompass multiple abundance measures opens fresh perspectives in biodiversity research.

Retrieve original file from: http://edepot.wur.nl/121801 High herbivore densities and re-occurring fires are natural phenomenons that determine the structure and functioning of African savannas. Traditional burning practices have been intensified over the past years due to increased herbivore numbers. Insight how animals respond to fire is important to understand ecosystem functioning and for the use of fire as management tool. Until now studies have centred mainly on spatial relationships between herbivores and fires. Very little information exists on effects of temporal fire patterns on herbivore communities. To assess the effect of fire succession on herbivore species greatly differing in body mass we conducted experiments and observational studies on smaller scales and landscape scales. We investigated the response of herbivores on both short-term and long-term succession patterns. Rodent densities were affected by the presence of larger herbivores rather than by seasonal patterns and increased with the absence of large herbivores and increasing vegetation height. Both middle-sized and large herbivore species responded on short-term post-fire patterns with large species being the pioneer species on burnt patches, whereas buffalo was the only species that also responded on long-term post-fire patterns. Our results indicate that larger herbivores in interaction with fire influence rodents by both increasing and prolonging the predation risk they experience in these habitats. Furthermore grazer presence and the amount of rainfall may be two interacting factors that prolong fire patterns in the landscape. Particularly buffalo responded to fire patterns, most likely due to the higher intake levels of digestible plant material and increased forage efficiency. Herbivore response to fire history

Species abundances are undoubtedly the most widely available macroecological data, but can we use them to distinguish among several models of community structure? Here we present a Bayesian analysis of species-abundance data that yields a full joint probability distribution of each model's parameters plus a relatively parameter-independent criterion, the posterior Bayes factor, to compare these models. We illustrate our approach by comparing three classical distributions: the zero-sum multinomial (ZSM) distribution, based on Hubbell's neutral model, the multivariate Poisson lognormal distribution (MPLN), based on niche arguments, and the discrete broken stick (DBS) distribution, based on MacArthur's broken stick model. We give explicit formulas for the probability of observing a particular species-abundance data set in each model, and argue that conditioning on both sample size and species count is needed to allow comparisons between the two distributions. We apply our approach to two neotropical communities (trees, fish). We find that DBS is largely inferior to ZSM and MPLN for both communities. The tree data do not allow discrimination between ZSM and MPLN, but for the fish data ZSM (neutral model) overwhelmingly outperforms MPLN (niche model), suggesting that dispersal plays a previously underestimated role in structuring tropical freshwater fish communities. We advocate this approach for identifying the relative importance of dispersal and niche-partitioning in determining diversity of different ecological groups of species under different environmental conditions.

Current neutral theory in community ecology views local biodiversity as a result of the interplay between speciation, extinction and immigration. Simulations and a mean-field approximation have been used to study this neutral theory. As simulations have limitations of convergence and the mean-field approximation ignores dependencies between species' abundances when applied to species-abundance data, there is still no final conclusion whether the neutral theory or the traditional lognormal model describes community structure best. We present a novel analytical framework, based on the genealogy of individuals in the local community, to overcome the problems of previous approaches, and show, using Bayesian statistics, that the lognormal model provides a slightly better fit to the species-abundance distribution of a much-discussed tropical tree community. A key feature of our approach is that it shows the tight link between genetic and species diversity, which creates important perspectives to future integration of evolutionary and community ecological theory.

Aim Tropical rain forests are often regarded as pristine and undisturbed by humans. In Central Africa, community-wide disturbances by natural causes are rare and therefore current theory predicts that natural gap phase dynamics structure tree species composition and diversity. However, the dominant tree species in many African forests recruit poorly, despite the presence of gaps. To explain this, we studied the disturbance history of a species-rich and structurally complex rain forest. Location Lowland rain forest in Southern Cameroon. Methods We identified the recruitment conditions of trees in different diameter classes in 16 ha of species-rich and structurally complex 'old growth' rain forest. For the identification of recruitment preference we used independent data on the species composition along a disturbance gradient, ranging from shifting cultivation fields (representing large-scale disturbance), to canopy gaps and old growth forest. Results In nine of sixteen 1-ha forest plots the older trees preferred shifting cultivation fields for recruitment while younger trees preferred gaps and closed forest conditions. This indicates that these nine sites once experienced large-scale disturbances. Three lines of evidence suggest that historical agricultural use is the most likely disturbance factor: (1) size of disturbed and undisturbed patches, (2) distribution of charcoal and (3) historical accounts of human population densities. Main conclusions Present-day tree species composition of a structurally complex and species-rich Central African rain forest still echoes historical disturbances, most probably caused by human land use between three to four centuries ago. Human impact on African rain forest is therefore, contrary to common belief, an issue not of the last decades only. Insights in historical use will help to get a more balanced view of the 'pristine rain forest', acknowledging that the dualism between 'old growth' and 'secondary' forest may be less clear than previously thought.

Secondary forests in Central Africa are increasing in importance for biodiversity conservation as old growth forests outside the few protected areas are disappearing rapidly. We examined vegetation recovery in a lowland rain forest area in Cameroon based on a detailed botanical survey of old growth forest and different-aged logging gaps (5-27 years) and shifting cultivation fields (10-60 years). Our analysis focuses on the long-term recovery of botanical conservation values by analysing trends in vegetation structure, species composition, species diversity and levels of endemism and rarity. In the total survey (4.25 ha), we recorded 834 species of which 23% were endemic to the Lower Guinea forest region. The proportion of endemic species was high in shrubs and low in herbs. Geographic range and (local) rarity were not significantly associated. The proportion of rare species (relative frequency

In this study we analysed the effects of large herbivores on small rodent communities in different habitats using large herbivore exclosures. We studied the effects of three year grazing introduction by red deer (Cervus elaphus L.) in previously ungrazed pine and oak woodland and the exclusion of grazing by red deer, roe deer (Capreolus capreolus L.) and mouflon (Ovis ammon musimin L.) in formerly, heavily grazed pine woodland and heathland. At eight exclosure sites within each habitat type, small rodents were captured with live traps using trapping grids. At each trapping grid, seed plots of beechnuts (Fagus sylvatica L.) and acorns (Quercus robur L.) were placed to measure seed predation by rodents. Exclusion of grazing by large herbivores in formerly, heavily grazed habitats had a significant effect on small rodent communities. Inside exclosures higher densities of mainly wood mice (Apodemus svlvaticus L.) and field voles (Microtus agrestis L.) were captured. Introduction of grazing by red deer appeared to have no significant negative effects on small rodent communities. The seed predation intensity of beechnuts and acorns by small rodents was significantly higher in ungrazed situations, particularly in habitats that were excluded from grazing. The differences between grazing introduction and exclusion effects on small rodent communities can be explained by differences in vegetation structure development. The recovery of heavily browsed understory vegetation after large herbivore grazing exclusion proceeded faster than the understory degradation due to grazing introduction. Small rodents depend on structural rich vegetations mainly for shelter. We conclude that large herbivores can have significant effects on vegetation dynamics not only via direct plant consumption but also through indirect effects by reducing the habitat quality of small rodent habitats.

Three grasses (Holcus lanatus, Anthoxanthum odoratum and Festuca ovina) and three herbs (Rumex obtusifolius, Plantago lanceolata and Hieracium pilosella) were grown in a greenhouse at 3 nutrient levels in order to evaluate plant allocation, architecture and biomass turnover in relation to fertility level of their habitats. Four harvests were done at intervals of 4 weeks. Various plant traits related to biomass partitioning, plant architecture, biomass turnover and performance were determined. Differences in nutrient supply induced a strong functional response in the species shoot:root allocation, but architecture and turnover showed little or no response. Architectural parameters like specific leaf area and specific root length, however, in general decreased during plant development. Species from more nutrient-rich successional stages were characterized by a larger specific leaf area and longer specific shoot height (height/shoot biomass), resulting in a higher RGR and total biomass in all nutrient conditions. There was no evidence that species from nutrient-poor environments had a longer specific root length or any other superior growth characteristic. The only advantage displayed by these species was a lower leaf turnover when expressed as the fraction of dead leaves and a shorter specific shoot height (SSH) which might prevent herbivory and mowing losses. The dead leaf fraction, which is a good indicator for biomass and nutrient loss, appeared to be not only determined by the leaf longevity, but was also found to be directly related to the RGR of the species. This new fact might explain the slow relative growth rates in species from a nutrient-poor habitat and should be considered in future discussions about turnover. [KEYWORDS: architecture; allocation; biomass turnover; nutrient availability; specific leaf area; specific root length Relative growth-rate; dry-matter; grassland; leaf; root; allocation; nitrogen; shoot; availabilities; competition]

We studied the degree and scale of patchiness of vegetation and selected soil variables along a gradient of herbivore impact. The gradient consisted of a radial pattern of 'high', 'intermediate' and 'low' herbivore impact around a watering point in a semi-arid environment in Burkina Faso (West Africa). We hypothesised that, at a certain range of herbivore impact, vegetated patches alternating with patches of bare soil would occur as a consequence of plant-soil feedbacks and run-off-run-on patterns. Indeed, our transect data collected along the gradient showed that vegetated patches with a scale of about 5-10 m, alternating with bare soil, occurred at intermediate herbivore impact. When analysing the data from the experimental sites along the gradient, however, we also found a high degree of patchiness of vegetation and soil variables in case of low and high herbivore impact. For low herbivore impact, most variation was spatially explained, up to 100% for vegetation biomass and soil temperature, with a patch scale of about 0.50 m. This was due to the presence of perennial grass tufts of Cymbopogon schoenanthus. Patterns of soil organic matter and NH4-N were highly correlated with these patterns of biomass and soil temperature, up to r = 0.7 (P < 0.05) for the in situ correlation between biomass and NH4-N. For high herbivore impact, we also found that most variation was spatially explained, up to 100% for biomass and soil temperature, and 84% for soil moisture, with three distinct scales of patchiness (about 0.50 m, 1.80 m and 2.80 m). Here, microrelief had a corresponding patchy structure. For intermediate herbivore impact, again most variation was spatially explained, up to 100% for biomass and soil temperature, and 84% for soil moisture, with a patch scale of about 0.95 m. Here, we found evidence that vegetated patches positively affected soil moisture through less run-off and higher infiltration of rainwater that could not infiltrate into the bare soil elsewhere, which was not due to microrelief. Thus, we conclude that our findings are in line with our initial hypothesis that, at intermediate herbivore impact, vegetated patches alternating with patches of bare soil persist in time due to positive plant-soil feedbacks.

Two facets of the putative power of herbivores to manipulate their food supply according to the ideas of McNaughton are reviewed with special reference to a young salt marsh in the Dutch Wadden Sea. A general feature of vegetations described as 'grazing lawns' seems to be that return visits by the herbivore enhance the quality of food on offer. In the salt marsh we studied, intensive and repeated grazing by staging geese in spring protracted the period of high nutritive status of the food plants, effectively extending the exploitation window for the herbivores by 7 weeks. Detailed quantification of the visit interval in relation to regrowth of previously grazed ramets indicate that the geese time their grazing cycle in order to maximize the intake of nutritious regrowth. However, the observed visit interval tends to fall on the early side of the model outcome for intake maximization, suggesting that the competitive interaction between flocks on the marsh needs further attention. In the longer term, vertebrate herbivory on the marsh extends the period of persistence of the younger successional stages of the vegetation supporting the heaviest level of exploitation. In our island study, the year-round presence of the brown hare, Lepus europaeus, retarded establishment of the dwarf shrub Atriplex on the young marsh, and thus facilitated access by the geese to the palatable species otherwise screened by the bushy Atriplex. Exclosures placed on the marsh at sites of different age up to 40 years, combined with experimental removal or addition of Atriplex plants, allow the conclusion that hares facilitate grazing by geese and extend the goose exploitation period by 25 years. Why the herbivores fail to arrest the onward march of plant succession on the marsh is unresolved but may be caused by ongoing changes in the nutrient status of the soil. The grazing lawns are thus not permanent in our system, but the general features of the plant-herbivore interaction agree with the framework hypothesized by McNaughton.

A study was conducted to analyse the effect of habitat productivity on woody species colonisation. Three soil types were distinguished: a relatively poor sandy soil type (1), a somewhat richer sandy type (2) and a relatively rich sandy loamy type (3). Chronosequences were established on these three soil types of 38 (type 1), 20 (type 2) and 54 years (type 3) after abandonment. In total 117 vegetation relevées were used to analyse life form change and species responses during old field succession via regression models. On the rich soil type the colonisation rate of woody species was slower than on the poor soil type. This can be explained by higher abundances of perennial species during the first 20 years after abandonment on the rich soil type in contrast to the poor soil type. Perennial species may delay the woody species colonisation. First they close the bare ground which inhibits germination and next they compete with woody seedlings for light, water and nutrients. The effect of habitat productivity on woody species colonisation can only be determined appropriately by taking life history traits into account. Early successional ‘pioneer’ woody species dispersed by wind have less difficulties colonising old fields than late successional ‘forest’ species; they colonise old fields prior to the development of a dense perennial sward. Forest species depend on animals to be dispersed which are attracted by vegetation structure. In ± 30 years on the poor soil type and in ± 45 years on the rich soil type woody species become dominant relative to other life forms. Forest species like Quercus robur L. invaded relatively early (

Vegetation and soil development during succession in coastal dune slacks on Terschelling island, the Netherlands, was investigated, by comparing neighbouring ecosystems on similar substrates that had been developing for 1, 5, 35 and 76 years since the vegetation and organic soil layer had been removed. In this successional sequence, soil organic matter accumulated rapidly due to the production of litter and dead roots. N mineralization was extremely low, increasing from 0.2 g m-2 yr-1 after 5 years to 0.8 g m-2 yr-1 after 76 years. It was accompanied by a decline in the pH (KCl) in the upper 10 cm of the soil from 6.8 to 4.4. Most of the above-ground biomass accumulated in the shrub species Oxycoccus macrocarpos and Salix repens. The 5- year-old plots harboured many plant species (18 species per 0.25 m2), but plant species diversity was much lower in the older plots. It is concluded that most changes in species composition and the decline in diversity occurred because early successional plant species were gradually outshaded by the thick litter layer and the accumulated shrub biomass.