Valuing mangrove ecosystem services: linking nutrient retention function of mangrove forests to enhanced agroecosystem production

Mangroves are highly productive wetland ecosystems strategically located at the interface between land and sea. They play an important role in the biogeochemical cycles of the coastal environment, acting as sources of nutrients to adjacent marine and terrestrial ecosystems through active and passive transport. We examined the nutrient contents in mangrove and nonmangrove soils in and around the Bhitarkanika National Park, India and assessed whether the local agricultural producers were aware of and placed a value on this contribution of mangrove forests in enhancing agroecosystem productivity. Soil samples from both mangrove and nonmangrove areas were analysed and quantity of organic carbon, total nitrogen, available phosphorus and potassium were derived. The replacement cost method was used to derive the value of nutrients in mangrove soils. We estimated that each hectare of mangrove contains additional nutrients worth US$232.49 in comparison to nonmangrove areas. The difference in nutrient content in mangrove versus nonmangrove areas gave the value of US$3.37 million for the nutrients in 145 km² of mangrove forests. The agricultural producers were aware that mangrove forests act as a source of nutrients and were willing to pay a higher price for the land adjoining mangrove forests. Around 92% of the producers ranked nutrient retention as a secondary function of mangrove forests. Despite crop depredation from wild ungulates and conflict with salt water crocodiles the agriculturist finds the benefit to cost ratio of mangrove forests high and more than 76% were in favour of mangrove restoration. This study provides an opportunity to highlight the importance of mangrove ecosystems to the livelihoods of the local people and the urgent need to sustain these through proper policy and market interventions.     

Faunal impact on vegetation structure and ecosystem function in mangrove forests: A review

The last 20 years witnessed a real paradigm shift concerning the impact of biotic factors on ecosystem functions as well as on vegetation structure of mangrove forests. Before this small scientific revolution took place, structural aspects of mangrove forests were viewed to be the result of abiotic processes acting from the bottom-up, while, at ecosystem level, the outwelling hypothesis stated that mangroves primary production was removed via tidal action and carried to adjacent nearshore ecosystems where it fuelled detrital based food-webs. The sesarmid crabs were the first macrofaunal taxon to be considered a main actor in mangrove structuring processes, thanks to a number of studies carried out in the Indo-Pacific forests in the late 1970s and early 1980s. Following these classical papers, a number of studies on Sesarmidae feeding and burrowing ecology were carried out, which leave no doubts about the great importance of these herbivorous crabs in structuring and functioning Old world ecosystems. Although Sesarmidae are still considered very important in shaping mangrove structure and functioning, recent literature emphasizes the significance of other invertebrates. The Ocypodidae have now been shown to have the same role as Sesarmidae in terms of retention of forest products and organic matter processing in New world mangroves. In both New and Old world mangroves, crabs process large amounts of algal primary production, contribute consistently to retention of mangrove production and as ecosystem engineers, change particle size distribution and enhance soil aeration. Our understanding of the strong impact of gastropods, by means of high intake rates of mangrove products and differential consumption of propagules, has changed only recently. The role of insects must also be stressed. It is now clear that older techniques used to assess herbivory rates by insects strongly underestimate their impact, both in case of leaf eating and wood boring species and that herbivorous insects can potentially play a strong role in many aspects of mangrove ecology. Moreover, researchers only recently realized that ant–plant interactions may form an important contribution to our understanding of insect–plant dynamics in these habitats. Ants seem to be able to relieve mangroves from important herbivores such as many insects and sesarmid crabs. It thus seems likely that ants have positive effects on mangrove performance.     

Organic carbon dynamics in mangrove ecosystems: A review

Our current knowledge on production, composition, transport, pathways and transformations of organic carbon in tropical mangrove environments is reviewed and discussed. Organic carbon entering mangrove foodwebs is either produced autochthonously or imported by tides and/or rivers. Mangrove litter and benthic microalgae are usually the most important autochthonous carbon sources. Depending on local conditions, phytoplankton and seagrass detritus imported with tides may represent a significant supplementary carbon input. Litter handling by the fauna not only affects microbial carbon transformations, but also the amount of organic carbon available for export. Most mangrove detritus that enters the sediment is degraded by microorganisms. Aerobic respiration and anaerobic sulfate reduction are usually considered the most important microbial respiration processes, but recent evidence suggests that iron respiration may be important in mangrove sediments as well. Organic carbon that escapes microbial degradation is stored in sediments and in some mangrove ecosystems, organic-rich sediments may extend to several meters depth. Many mangrove forests also lose a significant fraction of their net primary production to coastal waters. Large differences occur between mangrove forests with respect to litter production and export. Mangrove-derived DOC is also released into the water column and can add to the total organic carbon export. Numerous compounds have been characterized from mangrove tissues, including carbohydrates, amino acids, lignin-derived phenols, tannins, fatty acids, triterpenoids and n-alkanes. Many of these may, together with stable isotopes, exhibit a strong source signature and are potentially useful tracers of mangrove-derived organic matter. Our knowledge on mangrove carbon dynamics has improved considerably in recent years, but there are still significant gaps and shortcomings. These are emphasized and relevant research directions are suggested.     

Diversity,composition, and structure of tropical dry forests in Central America

Tropical dry forests have been reduced to less than 0.1% of their original expanse on the Pacific side of Central America and are considered by some to be the most endangered ecosystem in the lowland tropics. Plots 1000 m² were established in seven tropical dry forests in Costa Rica and Nicaragua in order to compare levels of species richness to other Neotropical dry forest sites and to identify environmental variables associated with species richness and abundance. A total of 204 species and 1484 individuals 2.5 cm were encountered. Santa Rosa National Park was the richest site with the highest family (33), genera (69), and species (75) diversity of all sites. Species richness and forest structure were significantly different between sites. Fabaceae was the dominant tree and shrub family at most sites, but no species was repeatably dominant based on number of stems in all fragments of tropical dry forest. Central American dry forests had similar species richness when compared to other Neotropical forests. There was no correlation between forest cover within reserves, or precipitation and plant species richness. There was a significant correlation between anthropogenic disturbance (intensity and frequency of fire, wood collection, grazing) and total species richness, tree and shrub species richness, and liana abundance. These results suggest controlling levels on anthropogenic disturbance within reserves should be a high priority for resource managers in Central America. Further research in forest fragments which examine individual and a combination of disturbance agents would help clarify the importance of anthropogenic disturbance on species richness and abundance.     

Allometry, biomass, and productivity of mangrove forests: A review

We review 72 published articles to elucidate characteristics of biomass allocation and productivity of mangrove forests and also introduce recent progress on the study of mangrove allometry to solve the site- and species-specific problems. This includes the testing of a common allometric equation, which may be applicable to mangroves worldwide. The biomass of mangrove forests varies with age, dominant species, and locality. In primary mangrove forests, the above-ground biomass tends to be relatively low near the sea and increases inland. On a global scale, mangrove forests in the tropics have much higher above-ground biomass than those in temperate areas. Mangroves often accumulate large amounts of biomass in their roots, and the above-ground biomass to below-ground biomass ratio of mangrove forests is significantly low compared to that of upland forests (ANCOVA, P < 0.01). Several studies have reported on the growth increment of biomass and litter production in mangrove forests. We introduce some recent studies using the so-called “summation method” and investigate the trends in net primary production (NPP). For crown heights below 10 m, the above-ground NPP of mangrove forests is significantly higher (ANOVA, P < 0.01) than in those of tropical upland forests. The above-ground litter production is generally high in mangrove forests. Moreover, in many mangrove forests, the rate of soil respiration is low, possibly because of anaerobic soil conditions. These trends in biomass allocation, NPP, and soil respiration will result in high net ecosystem production, making mangrove forests highly efficient carbon sinks in the tropics.     

Long-term retrospection on mangrove development using transdisciplinary approaches: A review

Large ecosystem processes often take place beyond the observation time of a researcher. Yet, through retrospective research scientists can approach and understand ecosystem changes. This contributes to the fundamental understanding of both human-induced and natural dynamics in ecosystems world-wide. This also holds for fast changing coastal areas with mangrove ecosystems, which are important for biodiversity, for coastal protection, and for the daily livelihood of millions of people in tropical coastal developing countries. In addition, retrospective research generates a basis for predictions that can be used early on to protect an ecosystem. In attempting to protect ecosystems from adverse human-induced change and destruction, and to manage them for sustainability, scientists are only beginning to investigate and understand natural ecosystem dynamics. It is important and advisable to gather, combine and analyse all possible data that allow a researcher to look back in time. This paper reviews the available retrospective methods, and highlights the transdisciplinary way (i.e. combination between basic and applied sciences on one hand, and social and human sciences on the other) in which retrospective research on a scale between months and centuries can be carried out, but it also includes methods on larger scales that may be marginally relevant. The paper particularly emphasizes the lack of transdisciplinary (not interdisciplinary) integration between sciences in retrospective research on mangrove forests in the past.     

Identifying practical indicators of biodiversity for stand-level management of plantation forests

Identification of valid indicators of biodiversity is a critical need for sustainable forest management. We developed compositional, structural and functional indicators of biodiversity for five taxonomic groups—bryophytes, vascular plants, spiders, hoverflies and birds—using data from 44 Sitka spruce (Picea sitchensis) and ash (Fraxinus excelsior) plantation forests in Ireland. The best structural biodiversity indicator was stand stage, defined using a multivariate classification of forest structure variables. However, biodiversity trends over the forest cycle and between tree species differ among the taxonomic groups studied. Canopy cover was the main structural indicator and affected other structural variables such as cover of lower vegetation layers. Other structural indicators included deadwood and distances to forest edge and to broadleaved woodland. Functional indicators included stand age, site environmental characteristics and management practices. Compositional indicators were limited to more easily identifiable plant and bird species. Our results suggest that the biodiversity of any one of the species groups we surveyed cannot act as a surrogate for all of the other species groups. However, certain subgroups, such as forest bryophytes and saproxylic hoverflies, may be able to act as surrogates for each other. The indicators we have identified should be used together to identify stands of potentially high biodiversity or to evaluate the biodiversity effects of silvicultural management practices. They are readily assessed by non-specialists, ecologically meaningful and applicable over a broad area with similar climate conditions and silvicultural systems. The approach we have used to develop biodiversity indicators, including stand structural types, is widely relevant and can enhance sustainable forest management of plantations.     

A review of studies on Pichavaram mangrove, southeast India

We studied a tropical mangrove ecosystem, situated at Pichavaram, southeast India. We found 13 species of mangrove trees, with Avicennia marinaand Rhizophora species predominant, besides 73 spp. of other plants, 52 spp. of bacteria, 23 spp. of fungi, 82 spp. of phytoplankton, 22 spp. of seaweeds, 3 spp. of seagrass, 95 spp. of zooplankton, 40 spp. of meiobenthos, 52 spp. of macrobenthos, 177 spp. of fish and 200 spp. of birds. The bacteria performed activities like photosynthesis, methanogenesis, magnetic behaviour, human pathogens and production of antibiotics and enzymes (arysulphatase, L-glutaminase, chitinase, L-asparaginase, cellulase, protease, phosphatase). The microzooplankton included tintinnids, rotifers, nauplius stages of copepods and veliger larvae of molluscs, with a predominance of tintinnids. Tintinopsis spp. alone accounted for 90% of abundance. The macrozooplankton consisted of 95% of copepods and coelenterates. The meiofauna was rich with nematodes (50–70% of the component), followed by foramifera. The macrofauna included polychaetes, bivalves, gastropods, tanaids, isopods, amphipods, cirripedes, crabs, hermit crabs and shrimps. The mangrove harboured a large number of juvenile fishes, especially during summer and post-monsoon. The water was fertile and productive in having several fold-higher levels of nutrients, microbes, plankton and other biological resources, than the adjoining estuarine, backwater and neritic environments. The gross primary production was 8 g cm^-3 d^-1; about 21% of which was contributed by phytoplankton of 5–10 m size. Unfortunately, 90% of the mangrove cover in the study area was degraded. Possible factors that cause degradation of the ecosystem are detailed and remedial measures suggested. Techniques for regeneration of the degraded areas are proposed.     

Investigating the role of impoundment and forest structure on the resistance and resilience of mangrove forests to hurricanes

The present study evaluates how a prolonged artificial flooding regime in impounded mangrove forests influences the regeneration after medium-sized forest destruction created by two hurricanes in 2004 in the Indian River Lagoon, Florida, USA. Disturbance patterns, forest structure, and regeneration were investigated. We found disturbed areas, characterized by uprooted and snapped trees, and intact forests in close proximity. The canopy turnover was greater in forests in higher succession stages with median tree diameter at breast height of 7.6 ± 5.7 cm compared to the intact forest with 3.7 ± 1.2 cm. Larger trees with lower densities were more susceptible to hurricane damage. We observed that regeneration of the open patches was dominated by the flood-tolerant species Rhizophora mangle (89.9%) instead of the faster-growing pioneer species Laguncularia racemosa (7.0%). Some of the disturbed areas created by the hurricanes were not recolonized. The regeneration rate of the disturbed areas expressed by vegetation closure >50% differed between almost zero to 100% in three different impoundments. We concluded that the artificial flooding regime imposed by impoundment is the predominant selective force in the successive process according to the species composition. However, we were not able to detect the driving factors that prevented mangrove establishment in disturbed areas.     

The forests and woodlands of Labrador, Canada: ecology, distribution and future management

Labrador, Canada is the last relatively undeveloped landmass of Boreal and subarctic Canada. Its land area is over 288,000 km², with less than 1% developed, and a human population of below 30,000. Labrador is greater than 60% forest- and woodland-covered and over 30% tundra, soil and rock barrens. We review the ecology and distribution of forests, woodlands, and related vegetation of Labrador within the context of climate, forest site index, landform, soils, and disturbance. Recent ecosystem management through a public planning process with emphasis on past and future comanagement and development with traditional and western scientific principles is currently underway. Plant–animal interactions, traditional uses by aboriginal groups, and early history are also reviewed.     

Patterns of local wood use and cutting of Philippine Mangrove Forests

Small-scale wood harvesting from mangrove forests is a commonplace yet barely studied phenomenon. This paper integrates bio-ecological and ethnographic methods to examine local wood use and cutting of mangrove forests in two areas of the Philippines. Findings reveal considerable site variation in cutting intensity, with heavier cutting typically closer to settlements and in forest stands that are not effectively regulated by government or private interests. Overall, cutting is responsible for almost 90% of stem mortality in both natural and plantation forests. Field measurements confirm ethnographic evidence indicating that harvesting for construction wood, but not fuelwood, is both species- and size-selective. Mangrove management and conservation efforts can be made more effective by better understanding how local people are harvesting wood resources from these forests.     

Liana diversity and distribution in four tropical dry evergreen forests on the Coromandel coast of south India

Liana diversity was inventoried in four tropical dry evergreen forest sites that are characterized by numerous trees, of short stature and small diameter, and a varying degree of anthropogenic disturbance, on the Coromandel coast of south India. A 1-ha plot was established in each of the four sites and was subdivided into 100 quadrats of 10 m× 10 m. All lianas 1 cm diameter at breast height (dbh) rooted within the plot were enumerated. The species richness and density of lianas, with respect to site disturbance and forest stature, varied across the sites. Liana density totaled 3307 individuals (range 497–1163 individuals ha^–1) and species richness totaled 39 species (range 24–29 species ha^–1) representing 34 genera and 24 families. Combretaceae, Asclepiadaceae, Capparaceae and Vitaceae were the well-represented families. The top five species Strychnos minor, Combretum albidum, Derris ovalifolia, Jasminum angustifolium and Reissantia indica contributed 55% of total density. The slopes of the species–area curves were different for each of the four sites and the curve stabilized in only one site. Of the four climbing modes recognized among the total 39 species, 18 were twiners (56% of the total density). Eight species (24% of density) were tendril climbers and 12 species (16% of density) were scramblers. Hugonia mystax was the only hook climber. All the 39 species and 88% of liana density were encountered within a category of 6 cm dbh or less, and a similar pattern prevailed in the individual sites. Of the three diaspore dispersal modes found among the 39 liana species, animal (64%) and wind (23%) dispersal were predominant over the autochorous mode (13%). Liana diversity and distribution in dry forest communities appear to be influenced by forest stature and site disturbance levels. In the light of the extent of liana diversity and sacred grove status of the study sites, the need for forest conservation, involving local people, is emphasized.     

Dispersal,establishment and survival of Ceriops tagal propagules in a north Australian mangrove forest

Studies of the ecology of mangroves show that a wide variety of factors, including salinity, desiccation, disturbance, competition and predation, may affect the distribution and abundance of species. Field studies were done to examine the relative importance of several of these factors in the establishment and early survival of Ceriops tagal, a species common in mid-to high-shore regions of mangrove forests in northern Australia. The fate of marked and tethered propagules was followed to estimate the range of dispersal and the intensity of predation. Propagules were artificially planted under different thicknesses of shade cloth (none, 30%, 80%) and in different habitats (clearing, forest, clearing-forest fringe) to examine the effects of light and soil conditions on survival and growth. Results suggested that dispersal was very limited: only 9% of marked propagules were ever found more than 3 m from the parent tree. Losses to predators were great, with 83% of tethered propagules being damaged or consumed within 3 months. On average, 56% of planted propagules survived for at least 6 weeks and 76% of these initiated growth. Survival in clearings was lower than in other habitats, with 29% fewer surviving six weeks and 48% fewer surviving 15 months. The growth of seedlings was correlated with soil temperature, but the effects of treatments were complex. Overall, results indicated that poor dispersal and establishment were the main factors likely to limit the colonisation and population growth of this species. Received: 12 September 1995/Accepted: 14 June 1996     

Gap formation and regeneration of tropical mangrove forests in Ranong,Thailand

Tropical mangrove forests are characterized by clear zonation along a tidal gradient, and it has been supposed that the zonation is primarily controlled by soil factors. However, effects of disturbance on mangrove forests are still not well understood and may play an important role on the vegetation patterns and forest dynamics in some forest formations. In this study, the pattern of disturbance regime and its effects on regeneration of tropical mangrove forests along a tidal gradient were investigated in Ranong, Thailand. We established one or two 0.5 ha plots in four vegetation zones, i.e. Sonneratia alba–Avicennia alba zone, Rhizophora apiculata zone, Ra – Bruguiera gymnorrhiza zone, Ceriops tagal–Xylocarpus spp. zone. Gap size (percentage gap area to total study area and individual gap size) was the largest in Sa–Aa zone which is located on the most seaward fringe, and it declined from seaward to inland. Canopy trees of S. alba and A. alba had stunted trunks and showed low tree density. On the contrary, canopy dominants in the other three inland zones, e.g. R. apiculata, B. gymnorrhiza, and Xylocarpus spp., had slender trunks and showed high tree density. Accordingly, differences in disturbance regime among the four zones were resulted from the forest structural features of each zone. Disturbance regime matched with regeneration strategies of canopy dominants. Seedlings and saplings of S. alba and A. alba, which need sunny condition for their growth, were abundant in gaps than in understorey. By contrast, R. apiculata, B. gymnorrhiza, and Xylocarpus spp., which can tolerate less light than S. alba and A. alba, had greater seedling and sapling density under closed canopy than gaps. Many large gaps may enhance the abundance of S. alba and A. alba in Sa–Aa zone, and a few small gaps may prevent the light demanding species to establish and grow in the other inland zones. Correspondence of disturbance regime and regeneration strategies (e.g. light requirement) of canopy dominants may contribute to the maintenance of the present species composition in each of the vegetation zones.     

Spatial and temporal variation in the sediment properties of an intertidal mangrove forest: implications for sampling

Studies of the structure or functioning of intertidal soft sediments often involves collection of biogeochemical data over tidal, diel and seasonal time-scales. Little effort has, however, been made to quantify accurately the time-scales at which these properties vary. Many previous studies collected samples from different sites at different times, potentially resulting in the confounding of spatial and temporal variation. This experiment was designed to determine if time of day or time within the exposure period had any significant effect on measurements of 7 different properties of sediments. Samples of sediment were collected using contact cores at the beginning, middle and end of tidal emersion at each of 9 a.m., 12 p.m. and 3 p.m., with two replicate days of each condition, from mangrove forests fringing Glades Bay, Sydney, Australia. These samples were analysed for water-content, pigments, carbohydrates, grain-size and loss on ignition (LOI), to determine the potential effects of time of day and time within the tidal cycle on these properties of the sediments.Whilst both time of day and time within the tidal cycle were found to occasionally have a significant effect upon the measured properties, most of the variation occurred among sites and between replicate days of each set of conditions. The minimal influence of time of day and time within the tidal cycle show that sampling effort should be preferentially placed into replicating days of sampling and sites because these are the scales with the greatest variation. Differences in the patterns found also depended on whether the data were expressed as content or concentration, the consequences of which are briefly described.     

Rehabilitation of degraded forests in Thailand: policy and practice

Thailand has suffered from severe deforestation during the last century. Forest cover has declined drastically both in terms of area and quality, mostly due to the expansion of human activities. Much of the deforested area has been used for agricultural purposes, but much has also been left in a degraded condition. In late 1980s, the forest declined to a point where the nation decided that the remaining forest should be kept for conservation rather than further exploitation. Consequently, forest policy has shifted its focus from exploitation to sustainable management and protection. Thailand has set a goal of increasing its forest area to 40% of the total land area, while at present, forests occupy around 28.9% of the land. With the intention to retain most of the remaining forest as protected areas and, at the same time, achieve the goal set, several reforestation and rehabilitation initiatives have been implemented, especially on those lands in a degraded condition. This paper focuses on the significant issues affecting both the policy and practice of forest rehabilitation. Given that the large number of people whose livelihood depends on the forests for subsistence and other purposes normally has been excluded from the decision-making process in forest management, most important among these issues are the integration of the socio-economic and environmental needs into rehabilitation initiatives together with the active participation of local communities in the rehabilitation program. Case studies of reforestation and rehabilitation initiatives are also discussed.     

Seed predation by insects in tropical mangrove forests: extent and effects on seed viability and the growth of seedlings

Summary Although insects are known to be important seed predators in most terrestrial forests, their role in marine tidal (mangrove) forests has not been examined. Surveys at 12 sites in tropical Australia showed that between 3.1 and 92.7 percent of the seeds or propagules of 12 mangrove tree species had been attacked by insects. Seeds/propagules of six species (Avicennia marina, Bruguiera gymnorrhiza, B. parviflora, Heritiera littoralis, Xylocarpus australasicus and X. granatum) showed consistently high (>40%) levels of insect damage. Greater than 99% of H. littoralis seeds were attacked by insect predators. The survival and subsequent growth in height and biomass of insect-damaged and non-damaged control seeds/propagules of eight mangrove species were compared in shadehouse experiments. Mangrove species fell into 4 groups with regard to the effect of insect predators on their seeds and seedlings. Xylocarpus australasicus and X. granatum had significantly decreased survival (X 48 and 70%) and growth in height (X 61 and 96%) and biomass (X 66 and 85%). Bruguiera parviflora showed decreased survival (X 59%), but there was no effect of insects on the growth of surviving propagules. In contrast, there was no effect of insect damage on the survival of seedlings of Avicennia marina and Bruguiera exaristata, but decreased growth in height (X 22 and 25%) and biomass (X 22 and 26%). Survival and growth of seedlings of Rhizophora stylosa and Bruguiera gymnorrhiza were not affected. The influence of insect seed predators on the survival and growth of seeds of mangrove species in forests will depend on the relative abundance of seed-eating crabs and intertidal position in mangrove forests.This is Contribution No 499 from the Australian Institute of Marine Science     

Seed bank and vegetation composition of forest stands of varying age in central Belgium: consequences for regeneration of ancient forest vegetation

Vegetation composition differs significantly between ancientand recent forest, due to slow colonization capacity of typical forest speciesand the higher abundance of early successional species in recent forest.However, little is known about differences in persistent seed bank compositionbetween ancient and recent forest and about the interaction between seed bankand vegetation in relation with forest age. We surveyed the seed bank and theunderstorey vegetationcomposition in transects from ancient to recent forest. Seed bank and fieldlayer vegetation characteristics and similarity between seed bank andvegetationwere analysed in relation to recent forest age and distance to the ancientforest. A total of 39 species and 14,911 seedlings germinated, whichcorresponds with a seed density of 12,426 seeds/m².Total seed density is significantly higher in the youngest recent forest parcel(55 years). Also the seed bank composition in the youngest forest parceldifferssignificantly from the other parcels. After a longer period of reforestation,the seed bank approaches that of the ancient forest, suggesting seed bankdepletion, although the seed bank is permanently replenished to some extent byseed bank forming species from local disturbances. Seed bank composition doesnot change significantly with distance to the ancient forest. Similaritybetween seed bank and vegetation composition, nomatter the forest age, is very low, but decreases with increasing forest age.The most frequent species in the vegetation are absent in the seed bank andvice versa. The contribution of forest species is highin the vegetation and they almost not occur in the seed bank, while species offorest edges and clearings, and species of disturbed environments are morefrequent in the seed bank. The seed bank is mainly composed of earlysuccessional species of former forest stages or species which temporary occurinsmall-scale disturbances. The seed bank may enhance the negative effects ofearlysuccessional, mainly competitive species to the forest species richness in therecent forest. In this respect, forest management should minimise forestdisturbances, to prevent germination of competitive species form buriedseeds.