Above‐ground biomass distribution among species during early old‐field succession

Abstract. The interdependence of species richness and plant biomass has widely been accepted as a general biodiversity rule. However, there is no information about how relationships are established during colonization and how total biomass is distributed among plants. The main objective of this study was to determine the role of several factors which we have hypothetized as affecting biomass distribution among species in an early old‐field community. To test this hypothesis, we conducted an experiment in a deeply ploughed Agropyro‐Rumicion crispi community in the Basque Country (Spain). Light, water and soil nitrogen content were factorially manipulated resulting in eight experimental treatments. We also examined several site features, which could potentially influence final biomass distribution: seed bank composition, soil physico‐chemical heterogeneity and cover and density of the dominant plant species in the community. A partition hypothesis testing approach using Redundancy Analysis was conducted to determine the fraction of biomass distribution variability assigned to each treatment and site feature. The most important species, in terms of biomass, were Agropyron repens, Sinapis arvensis, Arrhenatherum bulbosum and Picris echioides. As a general conclusion, biomass distribution among species during early secondary succession primarily depends on light availability and nitrogen supply. Several soil variables, such as magnesium, calcium and clay contents, also explain a relevant fraction of the biomass distribution among plant species. On the contrary, we found no effect of seed bank composition on biomass distribution. Finally, the total species number and cover of dominant species such as Sinapis arvensis, may determine final biomass distribution.     

Rarefaction does not eliminate the species richness‐biomass relationship in calcareous blackland prairies

Abstract. Relationships between species richness and biomass are common, but the causes remain controversial. It has been suggested that a hump‐shaped relationship may be an artifact of few individuals in both low and high biomass samples. We used rarefaction to standardize species richness to a constant number of individuals in 99 quadrat samples from relict calcareous prairies. Animal ecologists have long used rarefaction to standardize species richness to a constant number of individuals. The expected species richness of each quadrat was determined for 10, 25, 50, 100 and 150 individual plant ramets using the EcoSim program which resampled the data in each quadrat 1000 times. Because we had few samples with biomass over 300 g/m², our data showed a positive relationship between species richness and biomass. The relationship was largely unaffected by rarefaction. Even at the most extreme case of rarefying to 10 individual ramets, the relationship was clear. Monte Carlo covariance tests did not change the shape of the richness‐biomass relationship and significant correlations between observed species richness and rarified species richness suggest that the number of individuals had negligible effects on richness patterns. Because the data had no strong declining phase, these results may not apply to the declining phase of a humped relationship. The results are discussed with reference to other methods of standardizing species richness. Most importantly, we suggest that expressing species richness per number of individuals is inappropriate and may lead to incorrect conclusions.     

Effects of surface disturbance and light level on seedling emergence in a Japanese secondary deciduous forest

Abstract. The effects of soil surface disturbance and light level on seedling emergence were examined by creating experimental conditions differing in soil surface treatment and light level in a temperate deciduous broadleaved secondary forest in central Japan. The results of MANOVA showed that soil surface disturbance exerted a favourable influence upon the seedling emergence of many species. Effect of soil disturbance on total density and number of emerged species was greater than that of a high light level. The interaction between soil surface disturbance and light level revealed significant positive effects for seedling emergence, especially for small‐seeded species. The relationship between the percentage of emerged seedlings and seed weight was significantly negative in the soil‐surface disturbed and high light level condition, suggesting that minute‐seeded species largely depend on both factors for seedling emergence. Both soil surface disturbance and high light level had a positive effect on the number of emerged species, suggesting that these factors contributed to species richness at the stage of seedling emergence.     

The relationship between soil seed bank,above‐ground vegetation and disturbance intensity on old‐field successional permanent plots

Questions: How does disturbance and successional age influence richness, size and composition of the soil seed bank? What is the potential contribution of the soil seed bank to the plant community composition on sites differing in their successional age or disturbance intensity? Location: Experimental Botanical Garden of Göttingen University, central Germany. Methods: Above‐ground vegetation and soil seed bank were studied on formerly arable fields in a 36‐year‐old permanent plot study with five disturbance intensities, ranging from yearly ploughing via mowing to long‐term uninterrupted succession. We compared species compositions, seed densities and functional features of the seed bank and above‐ground vegetation by using several methods in parallel. Results: The seed bank was mainly composed of early successional species typical of strongly disturbed habitats. The difference between seed bank composition and above‐ground vegetation decreased with increasing disturbance intensity. The species of greatest quantitative importance in the seed bank was the non‐native forb Solidago canadensis. Conclusions: The ability of a plant community to regenerate from the soil seed bank dramatically decreases with increasing time since abandonment (successional age) and with decreasing disturbance intensity. The present study underlines that plant species typical of grasslands and woodlands are limited by dispersal capacity, owing to low capacity for accumulation of seeds in the soil and the fact that most species do not build up persistent seed banks. Rare and target species were almost absent from the seed bank and will, after local elimination, depend on reintroduction for continuation of their presence.     

Structure,dynamics and disturbance regime of temperate broad-leaved forests in Japan

Abstract. Structure, diversity and dynamics of five Japanese temperate old-growth forests were compared, differing from each other in, i.a., climate, dominant tree type, topography, disturbance regime. The forests were Aya, Ogawa, Kanumazawa Riparian, Kanumazawa upland and Senju. A permanent plot (1–6 ha) was established in each forest and trees were censused several times at intervals of two years. Mean annual recruitment rates and mortality rates in these forests were both in the range of 0.5 to 4.6 %/yr at the community level. Analyses of the structure and dynamics of populations showed that the underlying process was different among the forests. Some forests experienced compositional shifts in their canopies, others had a constant canopy composition but appeared to lack effective regeneration in recent years. The recruitment rate appeared to be strongly affected by competitive undergrowth vegetation such as dwarf bamboo which has been controlled by natural disturbance or human impact. It is likely that the forests with mostly low recruitment rates had a low species diversity. The current variation in structure, diversity and dynamics of the studied forests might have been determined not only by physical conditions (e.g. climate) but also by chance factors (e.g. disturbance, outbreak of deer population).     

Patch dynamics and local succession in a sandstone area with frequent disturbance

Abstract. A system of sand talus cones in a small forested rocky sandstone area was investigated to determine the importance of disturbance‐related dynamics and mesoclimate to vegetation differentiation. These cones (usually 3–12 m long) are formed by the accumulation of sand at the foot of sandstone rocks and are subject to frequent disturbance by the transport of sand. Vegetation was recorded both at the whole‐cone level and at the within‐cone level; the latter was approached by means of a transect of small squares along each cone. Soil profiles were recorded at the upper, centre and bottom parts of the cones. To express mesoclimatic differentiation among the cones within the rocky area, the potential insolation was calculated from the horizon angular height and the likelihood of thermal inversions was estimated by the height of the cones above the valley bottom. The major environmental factor correlated with variation in cone vegetation is the disturbance dynamics as determined by the structure of the soil profile; active cones (with sand at the surface) are colonized by different species as compared with non‐active cones (with surface covered by humus sediments). There is a clear primary succession gradient from plants able to cope with continuous sand transport to stands of tall bryophytes, small shrubs and herbs. Quite often the same gradient is found within cones, with late successional vegetation confined to the lower, stabilized parts of the cones, while the upper part is still being affected by sand transport. The differentiation along the gradient of disturbance is much stronger than the differentiation due to climatic or other gradients. Indirect data indicate that the long‐term average rate of sand accumulation is ca. 1 mm/yr. If sand transport ceases, Sphagnum peat accumulates on some of the cones; the depth of accumulated Sphagnum remains may reach 50 cm. The development of the peat layer is but weakly correlated with the measured variables suggesting that random processes at the beginning of Sphagnum establishment may be a driving force. By measuring Sphagnum growth and decomposition, and the peat density, we estimated the time needed for their development to be several hundreds of years.     

Old‐field grassland successional dynamics following cessation of chronic disturbance

Question: Does increasing Festuca canopy cover reduce plant species richness and, therefore, alter plant community composition and the relationship of litter to species richness in old‐field grassland? Location: Southeastern Oklahoma, USA. Methods: Canopy cover by species, species richness, and litter mass were collected within an old‐field grassland site on 16, 40 m × 40 m plots. Our study was conducted during the first three years of a long‐term study that investigated the effects of low‐level nitrogen enrichment and small mammal herbivory manipulations. Results: Succession was altered by an increase in abundance of Festuca over the 3‐yr study period. Species richness did not decline with litter accumulation. Instead, Festuca increased most on species‐poor plots, and Festuca abundance remained low on species‐rich plots. Conclusions: Festuca may act as an invasive transformer‐species in warm‐season dominated old‐field grasslands, a phenomenon associated more with invasions of cool‐season grasses at higher latitudes in North America.     

Soil heterogeneity effects on canopy structure and composition during early succession

Early succession from annuals to herbaceous and woody perennials on abandoned fields and grasslands is marked by vegetation heterogeneity; i.e., variation in canopy composition and structure over the field through time. Soil resource heterogeneity could promote vegetation heterogeneity. We created soil resource patchiness on two newly ploughed fields by establishing two types (fertilized, unfertilized) of checkerboard plots with two, four, or eight subplots (trenched, untrenched) to test effects of soil fertility heterogeneity scale on vegetation heterogeneity during early succession. Canopy composition and structure were censused in years 2, 4 and 6. On both fields, soil resource heterogeneity did not affect canopy composition or structure. Differential abundance of dominant species, Ambrosia trifida on one field and Solidago altissima on the other, between fertilized and unfertilized plots led to decreased vegetation heterogeneity among subplots within plots and increased vegetation heterogeneity among plots. Soil enrichment promoted taller and layered, but species-poor, canopy. In general, soil fertility effects on abundance, expansion, and dominance of individuals of dominant species promoted variation in canopy composition and structure, or vegetation heterogeneity, among plots on the two fields.     

Changes in Mediterranean plant succession: old-fields revisited

Abstract. Old-field plots used for a study of succession in Mediterranean France were revisited after 12–14 yr. Our aims were: (1) to verify if predicted patterns of species richness, turnover and composition are confirmed; (2) to compare the development in disturbed plots with that in undisturbed ones; (3) to discuss the impact of management changes. In undisturbed plots species richness and turnover decreased with successional age. Floristic composition changed in a way consistent with the predicted successional development in most plots. Therophytes decreased and phanerophytes increased; anemochorous species decreased and endozoochorous species increased, as expected. In plots disturbed since the first analysis richness decreased with successional age, but generally remained higher than in undisturbed plots. Floristic composition, species turnover and an increase in therophytes indicated changes towards younger successional stages. Thus, disturbance changed succession but not much. This is probably linked with the regeneration abilities typical of mediterranean species, e.g. resprouting. At the landscape scale, richness did not change and species turnover was low. The plots studied were situated in two distinct locations. One had not been disturbed between the two observation periods, while the other is a mosaic of undisturbed and disturbed sites. Observations fitted predictions much more closely at the undisturbed location. We conclude that permanent plot studies are powerful in identifying successional trends and can also provide additional insights into the effects of disturbance some of the mechanisms underlying the dynamics of diversity.     

Tropical forest structure and diversity in relation to altitude and disturbance in a Biosphere Reserve in central India

Questions: Is species diversity affected in protected areas where human activities are permitted or tolerated? On plots of a fixed size, does stem density alone predict number of species? Are differences in density related to disturbance and altitude? Location: Achanakmar‐Amarkantak Biosphere Reserve, central India. Methods: 42 sites, each with three replicate 10‐m radius plots, were examined. All trees (≥ 30 cm GBH) in each plot were measured for girth at breast height. α‐diversity, species richness and evenness were calculated for each site. The sites were ordinated by Nonmetric Multidimensional Scaling (NMS) using relative importance values of component species. Correspondence Analysis was used to broadly delineate communities. Anthropogenic disturbances were recorded in terms of percentage of trees lopped, scale of lopping, number of domestic livestock dung piles and foot trails (both livestock and people) for each plot. Results: The NMS analysis exhibited a near linear arrangement of sites with no evidence of discrete vegetation zones. NMS axes were significantly related to altitude and disturbance scores. With increasing elevation, basal area increased but number of species, α‐diversity and its components declined monotonically. The number of species and indices of species diversity were positively associated with tree lopping and also with total disturbance. Number of species was controlled by stem density only in plots not dominated by Shorea robusta. Conclusions: Recent levels of human disturbance are associated with higher species diversity in this biosphere reserve. There is some evidence that stands at all altitudes follow the same successional pattern to dominance by Shorea, a successional pattern that also results in decreased diversity without disturbance.     

Soil seed bank and floristic diversity in a forest‐grassland mosaic in southern Spain

Abstract. Soil seed bank and floristic diversity were studied in a forest of Quercus suber, a forest of Quercus canariensis and a grassland, forming a vegetation mosaic in Los Alcornocales Natural Park, southern Spain. The soil seed bank was estimated by the germination technique. In each community patch, diversity, woody species cover and herbaceous species frequency was measured. Three biodiversity components – species richness, endemism and taxonomic singularity – were considered in the vegetation and the seed bank. Forest patches had a soil seed bank of ca. 11 200–14 100 seed.m^?2 and their composition had low resemblance to (epigeal) vegetation. The grassland patch had a more dense seed bank (ca. 31 800 seed.m^?2) and a higher index of similarity with vegetation, compared with the forests nearby. The complete forest diversity was 71–78 species on 0.1 ha, including 12–15 species found only in the seed bank; the grassland species richness was higher (113 species on 0.1 ha). We discuss the role of soil seed banks in the vegetation dynamics and in the complete plant biodiversity of the mosaic landscape studied.     

Spatial and temporal variability during primary succession on tropical coastal sand dunes

Abstract. We studied primary succession on mobile tropical coastal sand dunes over an 8‐yr period. Every six months, we monitored changes in species composition and sand movement in permanent quadrats located on the windward slopes, the arms and the crests. Our results indicate that sand movement decreased over time but was significantly higher on the slopes and crests than on the arms. In all cases, there were seasonal fluctuations in sand movement which increased during the period with strong northerly winds and decreased during the rainy season. Sand movement was significantly correlated with species distribution. Plant cover and species richness increased at all three locations. Diversity increased on the slope, decreased on the arms and remained unchanged on the crest. However, the equitability values indicated the dominance of a few species, especially at the end of the study period. Temporal trends and species turnover rates differed among locations. Species turnover occurred first on the arms (1994), then on the crests (1998) and lastly on the slopes (at the end of the study period). In all cases the tall grass, Schizachyrium scoparium var. littoralis, became dominant over the endemic legume, Chamaecrista chamaecristoides var. chamaecristoides. Similar to temperate dunes, primary succession on tropical sand dunes was spatially and temporally heterogeneous.     

Effect of flooding and draw‐down disturbance on germination from a seashore meadow seed bank

Abstract. The objective of this study was to investigate the effects of flooding and draw‐down on the germination from the coastal grassland seed banks and to determine whether the effect of flooding varies between the delta and the seashore. Seed bank samples were collected from three shore transects in SW Finland, two on the shore of the Baltic Sea and one on the delta of River Kokemäenjoki. Samples were germinated in non‐flooded and flooded conditions for over a month, after which both treatments were maintained in non‐flooded conditions. A total of 9267 seedlings of 47 species germinated and mean density of seeds in the soil was ca. 84 000/m². Most of the seedlings were monocots (98%) and perennials (98%). Ca. 30–40% of the species found in the above‐ground vegetation had a seed bank including the majority of the most abundant species. The number of seeds and species richness increased as the organic layer became thicker. The organic layer was thicker in the seashore samples and the seed bank was significantly larger than in the delta. The flooding and draw‐down treatment significantly increased the number of germinating seedlings in the seashore and also increased species richness in two transects, one in the delta and the other in the seashore. Two species, Schoenoplectus tabernaemontani and Typha latifolia, had significantly higher germination in the flooded treatment than in the non‐flooded. Apparently, many species in these coastal grasslands have adapted to flood disturbance and for seeds of some species flooding may work as a positive signal, possibly breaking dormancy.     

Restorative mowing on an abandoned semi‐natural meadow: short‐term and predicted long‐term effects

Abstract. When management, in the form of cattle grazing and mowing, ceases the abundance of competitively superior plant species tends to increase in abandoned semi‐natural meadows. Litter accumulation elevates the soil nutrient levels and hinders seedling recruitment. We surveyed changes in plant cover and species composition of a formerly grazed meadow in permanent plots for six years. Some plots were unmown, while others were mown and raked annually in August. The cover of grasses decreased and herb cover remained unchanged regardless of the treatment. Mowing and raking significantly reduced litter accumulation and increased the number of ground layer species. The expected long‐term effects of abandonment and restorative mowing were studied by calculating the transition probabilities for unmown and mown plots and simulating the course of succession as projected by the transition matrices. During a simulation period of 30 yr, abandonment led to (1) a decrease in the cover of small herbs, (2) a slight increase in the cover of tall herbs and (3) a slight decrease in the cover of grasses. In contrast, the cover of small herbs on the mown plots remained unchanged or slightly increased during the course of simulation. These results suggest that mowing late in the season is primarily a management tool for the maintenance of the existing species diversity and composition. However, it may not be an effective restorative tool to induce overall changes in the resident vegetation of abandoned grass‐dominated meadows. Grazing or mowing early in the season may be more effective in this respect. Consequently, mowing early or, alternatively, late in the season may provide management strategies for the maintenance and restoration of species diversity, respectively.     

Spatio-temporal patterns in post-burn succession on Brittany heathlands

Heathlands in the Paimpont area, Brittany, were ravaged by numerous fires in the summer of 1976. The period in which the fires occurred, and their intensity had important consequences for recolonization processes. In the tall Ulex europaeus heathland described in this paper, fire occurred relatively late (mid-July), and was of high intensity. Colonisation was mainly from propagules, often from species foreign to the original heathland. Plant succession together with vertical and horizontal structure was studied in permanent plots over 13 yr. Three stages were distinguished. Vertical structure varied greatly among the three stages. It was very simple in the colonisation phase, became more complicated during the competition phase, but became simpler again in the stabilisation phase. This was particularly evident for the moss layer, which, 13 yr after the fire, was made up of Polytrichum commune alone. Mosses showed interspecific competition with size as the decisive factor: the large mosses gradually eliminated the smaller ones. Horizontal structure was studied using spatial autocorrelation, which provided some indication of the distinctive -ness and scale of spatial pattern. Some species were weakly aggregated such as Ceratodon purpureus, Polytrichum pili-ferum, and P. juniperinum, whileothers, like the P. formosum-P. commune complex, Agrostis curtisii and Ulex minor were strongly aggregated. In the current community, 13 yr after the fire, a mosaic can be distinguished with three components: (1) a moss layer of Polytrichum commune on which a herbaceous layer, dominated by Agrostis curtisii has developed; (2) a birch wood (both Betulapéndula and B. pubescens), which established on the Polytrichum - Agrostis combination; (3) patches of Ulex minor heathland, with a few Frica and Calluna plants. Extreme simplicity and stability are the two prominent features of the present vegetation state.     

Effects of trampling and vegetation removal on species diversity and micro-environment under different shade conditions

Abstract. The effects of disturbance by trampling and vegetation removal on herbaceous communities and their micro-environments were examined at two sites with different levels of shade. The dominant species of the original vegetation were the perennial herb Artemisia princeps at the sunny site and the dwarf-bamboo Pleioblastus chino at the shady site. With no disturbance and marked dominance of these species, diversity was low. After vegetation removal there was a rapid recovery, leading to a more diverse vegetation with many more species, each with a lower dominance. Continuous trampling induced a short vegetation while the light intensity under the foliage was increased. At the sunny site, species richness was depressed by trampling because the soil water potential decreased markedly during summer and only the stress-tolerant annual Digitaria adscendens grew vigorously. Thus, the lowest species diversity was observed in the heavily trampled vegetation in late summer. At the shady site, soil water availability was not affected by trampling. This allowed the survival of many species and prevented a strong decline in diversity. The results suggest that the pattern of change in diversity in communities subjected to various disturbances, was always determined by the original environments.     

Post‐fire succession and 20th century reduction in fire frequency on xeric southern Appalachian sites

Abstract. We document post‐fire succession on xeric sites in the southern Appalachian Mountains, USA and assess effects of 20th century reduction in fire frequency on vegetation structure and composition. Successional studies over 18 yr on permanent plots that had burned in 1976–1977 indicate that tree mortality and vegetation response varied with fuel load and fire season. In the first three years after fire, hardwood sprouts dominated tree regeneration. On sites where summer and autumn fires reduced litter depth to less than 1 cm, densities of shade‐intolerant Pinus seedlings increased steadily over this period. 4 to 8 yr after fire, large numbers of newly established seedlings and sprouts had grown to 1 – 10 cm DBH. By year 18 growth of these saplings led to canopy closure on most sites. Herbaceous cover and richness peaked in the first decade after fire, then declined. On similar sites that had not burned in more than 50 yr, regeneration of shade‐intolerant Pinus spp. and mean cover and richness of herbs were considerably lower than those observed on recently burned plots. Reconstructions of landscape conditions based on observed post‐fire succession and 20th century changes in fire regime suggest that reductions in fire frequency circa 1940 led to substantial changes in forest structure and decreases in cover and richness of herbaceous species.     

Challenging Theophrastus: A common core list of plant traits for functional ecology

Abstract. Ecologists need a common language of plant traits in order to make comparisons across regions and scales, pool data, and maximize the utility of the data. To develop such a set of traits we began with the primary challenges faced by plants: dispersal, establishment, and persistence in order to identify fundamental traits. Most of these traits are hard to measure, but advances in comparative ecology have suggested a number of easy to measure analogs. Unfortunately, some of the fundamental traits have no simple analog. The common core list includes: seed mass, seed shape, dispersal mode, clonality, specific leaf area, leaf water content, height, above-ground biomass, life history, onset of flowering, stem density, and resprouting ability. Most of the traits can be measured quantitatively, but several traits on the list must still be measured qualitatively due to logistical problems or lack of an easy analog. Key problem areas include: dispersal ability, capacity for vegetative spread, germination, palatability, plasticity, and all the various below-ground traits. Comparative studies need to address these problem areas. The common core list is suggested as a common starting point for studies of functional ecology. The idiosyncrasies of regional floras and specific research agendas will dictate which traits can be ignored and those that need to be added.     

Influence of site conditions on the survival of Fagus sylvatica seedlings in an old‐growth beech forest

Abstract. Following a full mast production in autumn 1995 in the old‐growth beech forest of la Tillaie (France), cupules, live and dead 1‐yr old seedlings were counted in September 1996 in 40 plots, representing a wide range of ecological conditions. The influence of geomorphology and forest stage on mast production and on seedling establishment and survival was studied. Maximum seedling abundance did not occur in areas with the largest cupule abundance, but this result was not statistically significant. The presence of a shallow sandstone table under Fontainebleau sand, where beech is far from its ecological optimum, positively influenced the production of mast but negatively influenced seedling survival. Contrary to expectation, the early biostatic rather than the late biostatic phase seemed to be the best stage for beech regeneration. Gaps also provide good environmental conditions for seedling establishment. Humus and light conditions, water availability and competition between beech individuals may explain the results. Light was the main factor influencing mast production in mature stages, and drought was an important factor causing seedling mortality. This study points out the importance of seedling survival for regeneration patterns in beech forests.