Contrasting impacts of a native and an invasive exotic shrub on flood‐plain succession

Abstract. Question: How do Coriaria arborea, an N‐fixing native shrub, and Buddleja davidii, a non‐N‐fixing exotic shrub, affect N:P stoichiometry in plants and soils during early stages of primary succession on a flood‐plain? Location: Kowhai River Valley, northeast South Island, New Zealand. Methods: We measured soil and foliar nutrient concentrations, light levels, plant community composition and the above‐ground biomass of Coriaria and Buddleja in four successional stages: open, young, vigorous and mature. Results: Coriaria occurred at low density but dominated above‐ground biomass by the vigorous stage. Buddleja occurred at 5.3 ± 1.0 stems/m² in the young stage and reached a maximum biomass of 520–535 g.m^‐2 during the young and vigorous stages. Mineral soil N increased with above‐ground Coriaria biomass (r²= 0.45), but did not vary with Buddleja biomass. In contrast, soil P increased with Buddleja biomass (r²= 0.35), but not with Coriaria biomass. In early successional stages, 70–80% of the species present were exotic, but this declined to about 15% by the mature stage. Exotic plant species richness declined with increasing Coriaria biomass, but no other measures of diversity varied with either Coriaria or Buddleja biomass. Conclusion: These results demonstrate that Buddleja dominates early succession and accumulates P whereas Coriaria dominates later succession and accumulates N. A key ecosystem effect of the invasive exotic Buddleja is alteration of soil N:P stoichiometry.     

First report on the occurrence of the uncultivated cluster 2 Frankia microsymbionts in soil outside the native actinorhizal host range area

The occurrence of uncultivated Frankia was evaluated in Tunisian soils by a plant-trapping assay using Coriaria myrtifolia seedlings. Despite the lack of this compatible host plant for more than two centuries, soil-borne Frankia cells were detected in one sampled soil as shown by the development of root nodules on 2-year-old seedlings. Based on glnA sequences, Tunisian trapped Frankia strains belong to the uncultivated cluster 2 strains that associate with other Coriaria species and also with Ceanothus, Datisca and Rosaceae actinorhizal species. This is the first report on the occurrence of Frankia cluster 2 strains in soils from areas lacking compatible host plant groups.     

PCR-amplified 16S rRNA sequence analysis to confirm nodulation of Datisca cannabina L. by the endophyte of Coriaria nepalensis Wall

Different Frankia strains and crushed nodule suspensions were tested for their ability to nodulate Coriaria nepalensis and Datisca cannabina. Datisca cannabina seedlings were nodulated effectively by both crushed nodule suspension from Coriaria nepalensis and Datisca cannabina. The origin of the endophyte in Datisca nodules induced by crushed nodules of Coriaria was confirmed by comparing partial PCR-amplified 16S rRNA sequences with those of the endophytes of both plants. Coriaria seedlings could only be nodulated by crushed nodule suspensions of Coriaria nepalensis. All pure cultures of Frankia used as a single inoculum source or in combinations with a nodule filtrate, failed to induce nodulation on Coriaria. Two atypical Frankia strains Cn3 and Cn7 isolated from Coriaria nodules showed no acetylene reduction activity and did not induce nodulation on the host seedlings.     

Flood‐deposited wood creates regeneration niches for riparian vegetation on a semi‐arid South African river

Question : The formation of large woody debris (LWD) piles during floods has significant impacts on riparian succession through pioneering plants often establishing in association with wood. We assess the importance of LWD for seed regeneration of riparian plants after a century‐scale flood disturbance in a semi‐arid environment. Location : The Sabie River within Kruger National Park in the semi‐arid northeast of South Africa. Methods : Our approach was to quantify the riparian soil seed bank, to record the frequency of establishment of riparian plants in woody debris piles, and to conduct experimental out‐plantings of common riparian trees in plots with and without LWD. Results : We found the abundance and diversity of seedlings were higher in soils taken from wood piles than from open reference areas, and most seedlings were herbaceous species. Surveys indicated that numbers of seedlings recorded within woody debris were significantly greater than in open reference areas or within established vegetation. Seedling establishment in various cover‐types also varied for different riparian tree species. Experimental out‐planting of seedlings of two riparian tree species (Philenoptera violacea and Combretum erythrophyllum) revealed that, after 433 days, planted seedlings survived only in woody debris piles. Conclusion : LWD formed after a large flood creates heterogeneous patches that may influence post‐disturbance regeneration of riparian vegetation by providing a variety of environmental niches for seedlings establishment. We suspect that higher seedling survival in LWD is due to increased moisture (particularly in the dry season) and nutrients, and protection from seasonal flooding and herbivory.     

Succession and Micro-elevation Effects on Seedling Establishment of Calophyllum brasiliense Camb. (Clusiaceae) in an Amazonian River Meander Forest1

I investigated the effects of successional stage and micro‐elevation on seedling establishment of Calophyllum brasiliense (Clusiaceae), a common canopy tree of seasonally flooded lowland forest along the Manú River meander zone in southeastern Peru. To compare seedling establishment between microhabitat types, I planted C. brasiliense seeds in a fully crossed experimental design of three successional stages (early, mid, and mature) and two micro‐elevations (levees and backwaters). Seedling establishment success in this study was affected by both successional stage and micro‐elevation, but micro‐elevation was most important in mid‐successional habitats. In general, seedlings in early succession experienced better conditions than in mature forest; light levels were higher, herbivory lower, and seedling growth higher. In mid‐successional forest, micro‐elevation determined habitat quality; backwaters had higher light levels, lower herbivory, and higher seedling growth and survival than levees. Mid‐successional backwaters were similar in quality to early successional forest for seedling establishment, while levees in that same successional stage were the poorest microhabitats for establishment. Although mid‐successional backwaters are similar to early succession for seedling establishment, in the long run, seedlings that establish in mid‐succession have a lower chance of reaching reproductive size before their habitat ages to mature forest than members of their cohort that established in early succession. I hypothesize that successful recruitment for C. brasiliense in the Manú River meander system requires dispersal to early successional habitat.     

Canopy vs. soil effects of shrubs facilitating tree seedlings in Mediterranean montane ecosystems

Abstract. Question: Is the facilitative effect of nurse shrubs on early recruitment of trees mediated by a ‘canopy effect’(microclimate amelioration and protection from herbivores), a ‘soil effect’(modification of soil properties), or both? Location: Two successional montane shrublands at the Sierra Nevada Protected Area, SE Spain. Method: Seedlings of Quercus and Pinus species were planted in four experimental treatments: (1) under shrubs; (2) in open interspaces without vegetation; (3) under shrubs where the canopies were removed; (4) in open interspaces but covering seedlings with branches, mimicking a shrub canopy. Results: Both effects benefited seedling performance. However, microclimatic amelioration due to canopy shading had the strongest effect, which was particularly pronounced in the drier site. Below‐ground, shrubs did not modify soil physical characteristics, organic matter, total N and P, or water content, but significantly increased available K, which has been shown to improve seedling water‐use efficiency under drought conditions. Conclusions: We propose that in Mediterranean montane ecosystems, characterised by a severe summer drought, pioneer shrubs represent a major safe site for tree early recruitment during secondary succession, improving seedling survival during summer by the modification of both the above‐and below‐ground environment.     

Facilitation‐ vs. competition‐driven succession: the key role of resource‐ratio

Symbiotic nitrogen (N)‐fixing plants are abundant during primary succession, as typical bedrocks lack available N. In turn, fixed N accumulates in soils through biomass turnover and recycling, favouring more nitrophilous organisms. Yet, it is unclear how this facilitation mechanism interacts with competition for other limiting nutrients such as phosphorus (P) and how this affects succession. Here, we introduce a resource‐explicit, community assembly model of N‐fixing species and analyze successional trajectories along resource availability gradients using contemporary niche theory. We show that facilitation‐driven succession occurs under low N and high enough P availabilities, and is characterised by autogenic ecosystem development and relatively ordered trajectories. We show that late facilitation‐driven succession is sensitive to catastrophic shifts, highlighting the need to invoke other mechanisms to explain ecosystem stability near the climax. Put together with competition‐driven succession, these results lead to an enriched version of Tilman's resource‐ratio theory of succession.     

Identifying Native Vegetation for Reducing Exotic Species during the Restoration of Desert Ecosystems

There is currently much interest in restoration ecology in identifying native vegetation that can decrease the invasibility by exotic species of environments undergoing restoration. However, uncertainty remains about restoration's ability to limit exotic species, particularly in deserts where facilitative interactions between plants are prevalent. Using candidate native species for restoration in the Mojave Desert of the southwestern U.S.A., we experimentally assembled a range of plant communities from early successional forbs to late‐successional shrubs and assessed which vegetation types reduced the establishment of the priority invasive annuals Bromus rubens (red brome) and Schismus spp. (Mediterranean grass) in control and N‐enriched soils. Compared to early successional grass and shrub and late‐successional shrub communities, an early forb community best resisted invasion, reducing exotic species biomass by 88% (N added) and 97% (no N added) relative to controls (no native plants). In native species monocultures, Sphaeralcea ambigua (desert globemallow), an early successional forb, was the least invasible, reducing exotic biomass by 91%. However, the least‐invaded vegetation types did not reduce soil N or P relative to other vegetation types nor was native plant cover linked to invasibility, suggesting that other traits influenced native‐exotic species interactions. This study provides experimental field evidence that native vegetation types exist that may reduce exotic grass establishment in the Mojave Desert, and that these candidates for restoration are not necessarily late‐successional communities. More generally, results indicate the importance of careful native species selection when exotic species invasions must be constrained for restoration to be successful.     

Population dynamics of Echinops gmelinii Turcz. at different successional stages of biological soil crusts in a temperate desert in China

• The effects of biological soil crusts (BSC) on vascular plant growth can be positive, neutral or negative, and little information is available on the impacts of different BSC successional stages on vascular plant population dynamics.
• We analysed seedling emergence, survival, plant growth and reproduction in response to different BSC successional stages (i.e. habitats: bare soil, cyanobacteria, lichen and moss crusts) in natural populations of Echinops gmelinii Turcz. in the Tengger Desert of northwest China. The winter annual E. gmelinii is a dominant pioneer herb after sand stabilisation.
• During the early stages of BSC succession, the studied populations of E. gmelinii were characterised by high density, plant growth and fecundity. As the BSC succession proceeded beyond moss crusts, the fecundity decreased sharply, which limited seedling recruitment. Differences in seedling survival among the successional stages were not evident, indicating that BSC have little effect on survival in arid desert regions. Moreover, E. gmelinii biomass allocation exhibited low plasticity, and only reproductive allocation was sensitive to the various habitats. Our results further suggest that the negative effects of BSC succession on population dynamics are primarily driven by increasing topsoil water‐holding capacity and decreasing rain water infiltration into deeper soil.
• We conclude that BSC succession drives population dynamics of E. gmelinii, primarily via its effect on soil moisture. The primary cause for E. gmelinii population decline during the moss‐dominated stage of BSC succession is decreased fecundity of individual plants, with declining seed mass possibly reducing the success of seedling establishment.

     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

Posthurricane Seedling Structure in a Multi‐aged Tropical Dry Forest: Implications for Community Succession

Hurricane‐caused tree mortality in tropical dry forests occurs predominantly in early successional species. Consequently, hurricanes may accelerate succession in these forests. Forest regeneration, however, must be measured over an extended posthurricane time period to demonstrate this pattern. In this study, we recorded tree seedlings in 19 Florida Keys forests during May–August 1995, 3 years after Hurricane Andrew. For these forests—spanning a chronosequence from 14 to over 100 years since the most recent clearing—we used weighted averaging regression on relative abundances of pre‐hurricane trees to calculate a successional age optimum for each species; and used weighted averaging calibration to calculate inferred successional ages for stands based on pre‐hurricane trees and on posthurricane seedlings. To test the hypothesis that successional stage of seedlings exceeded successional stage of pre‐hurricane trees, we compared inferred stand ages based on posthurricane seedlings with those based on pre‐hurricane trees. Across the study area, inferred stand ages based on posthurricane seedlings were greater than those based on pre‐hurricane trees (P < 0.005); however, more seedlings in the youngest stands were early successional than in older stands. Of 29 species present both as pre‐hurricane trees and posthurricane seedlings, 23 had animal‐dispersed seeds. These results provide evidence that: (1) hurricanes do not ‘reset’ succession, and may accelerate succession; and (2) a strong legacy of stand successional age influences seedling assemblages in these forests.     

Vegetation succession at the abandoned Ogushi sulfur mine, central Japan

We surveyed plant community development at the abandoned Ogushi sulfur mine. We found seven communities dominated by the following respective species: Deschampsia flexuosa, Miscanthus sinensis, shrub willow, Gaultheria miqueliana–Betula ermanii, Sasa senanensis–Betula ermanii, willow–Betula ermanii, and Sasa kurilensis–Abies veitchii. We examined the succession of these communities, in which younger communities of low height and ground cover contained seedlings of the successive communities that were taller and had higher ground cover. To understand the development of these different communities, we surveyed damage from mining pollution and effects of immature soils formed by landslides. The average pH (H₂O) was 4.12, and aluminum concentrations were not sufficiently high to damage plant growth, except in areas where sulfur had been mined. The organic carbon and nitrogen content in soil samples were very low because of a delay in soil development caused by a large landslide in 1937. Hence, succession was positively correlated with the soil development stage. The delay in soil development after a large landslide influenced the seven successional steps of the plant communities, but mineral poisons at the abandoned Ogushi sulfur mine had no effect on succession.     

Plant distribution patterns during early succession on an artificial protosoil

Abstract. Plant succession has been studied since 1977 in permanent plots on a calcareous protosoil in the northern outskirts of Göttingen (Central Germany). At the beginning the substrate was extremely poor in organic carbon and nitrogen. On 128 plots of 100 m², the temporal and spatial distribution of the most conspicuous plant species showed remarkable changes and correlations to soil site factors. While presenting data on Melilotus alba, Medicago lupulina, Agropyron repens, Arrhenatherum elatius, Dactylis glomerata, and Picris hieracioides the mechanisms of successional changes on nutrient-poor soils are discussed. It is demonstrated that a high cover of legumes favours the accumulation of soil nitrogen. Grassland species were more abundant in areas with low humus content whereas ruderal plant species preferred those parts of the area which contained more humus in the substrate. Life-form, colonization and population strategies play an important role in the described successional processes, in addition to resource competition. Finally, some methodological conclusions for permanent plot studies are presented.     

The role of successional stage,vegetation type and soil disturbance in the invasion of the alien grass Cortaderia selloana

Question : This paper studies the establishment and performance of Cortaderia selloana (Pampas grass), an alien South American ornamental species that is invading many parts of the world. We asked whether (1) early successional stages were the most susceptible to C. selloana invasion; (2) soil microdisturbances increased invasion at any point of succession, and (3) C. selloana invasion of later successional stages was modulated by vegetation type Location : Delta del Llobregat (Catalonia, NE Spain). Methods : We monitored survival and growth of transplanted C. selloana seedlings in disturbed and non‐disturbed plots throughout a successional gradient with an age range of < 1 to > 10 years in different vegetation types and within the area of influence of coexisting species with similar growth form. Results : Although seedling survival was extremely low in all treatments, our results revealed that early successional stages were not the most easily invaded since we found no significant differences in the percentage survival of C. selloana along the successional gradient. However, survival and seedling bio‐mass were enhanced by soil disturbance at any seral stage. This result suggested that inhibition ruled C. selloana invasion. Invasibility neither depended on the invaded vegetation type nor on the co‐existing species with similar growth form. Finally, C. selloana invasion was not enhanced by decreasing competition with Phragmites australis, a native coexisting species because survival rates after a year were not significantly different. However, Phragmites increased C. selloana leaf length probably due to shading. Conclusions : C. selloana recruitment appears to be positively affected by soil disturbance but it is independent of successional stage or vegetation type.     

Effects of microsite conditions on seedling establishment on the foreland of Coleman Glacier,Washington

Abstract. Questions: How do physical microsite conditions of microsites affect germination and seedling survival in different successional stages? Do different species germinate in similar microsites in a given successional stage? Location: Coleman Glacier foreland, Mount Baker, Washington State, USA. Methods: Two methods were used to characterize safe sites. 1. Grids of 300 10 cm × 10 cm plots were located in four different age classes on the foreland. 2.105 pairs of plots, with and without seedlings of Abies amabilis, were located in each age class. For each plot we identified all seedlings and all individuals < 1 m tall. Microsite characteristics such as topography and presence of rocks or woody debris were noted for each plot. Microsite characteristics were compared between plots with and without each species. In addition we examined the effect of distance from seed sources on the presence of Alnus viridis seeds and seedlings in a newly disturbed area. Results: In early successional sites, seedlings of several species were positively associated with depressions and presence of rocks, and negatively associated with ridges. Patterns were generally consistent among species. In later succession, seedlings were not significantly associated with any microsite characteristics. For Alnus viridis, seed density decreased with distance from seed sources but seedling density did not. Conclusions: Because of harsh conditions in early succession, physical microsites are important, and most species have similar microsite requirements. In later succession, physical microsites characteristics are not as important and are more variable. Microsites appear to be more important than seed rain in controlling the distribution of Alnus viridis in early succession.     

Can atmospheric input of nitrogen affect seed bank dynamics in habitats of conservation interest? The case of dune slacks

Questions: Does the increased atmospheric deposition of nitrogen, which can have major effects on the established vegetation of nutrient‐poor habitats, also impact germination from the soil seed bank? Location: Coastal dune slacks at Newborough Warren, Wales, UK. Methods: The effects of nitrogen addition (15 kg.ha^‐1.a^‐1) on seed germination from the soil seed bank were investigated using the seedling emergence method between September 2004 and February 2005. Results: More seedlings emerged from fertilised samples than unfertilised controls. Most species showed enhanced germination after fertilisation with nitrogen, with seedling numbers statistically significantly greater in nitrogen addition samples in a quarter of species abundant enough for analysis. Species that responded positively to fertilisation were species with low Ellenberg indicator values indicative of infertile sites. Conclusions: Most species showed increased germination after fertilisation with nitrogen, including early successional species normally growing in nutrient‐poor conditions. This suggests that the increased atmospheric deposition of nitrogen probably not only impacts on established vegetation, but also has the potential to alter seed bank dynamics.     

Soil conditioning effects of Phragmites australis on native wetland plant seedling survival

Interactions between introduced plants and soils they colonize are central to invasive species success in many systems. Belowground biotic and abiotic changes can influence the success of introduced species as well as their native competitors. All plants alter soil properties after colonization but, in the case of many invasive plant species, it is unclear whether the strength and direction of these soil conditioning effects are due to plant traits, plant origin, or local population characteristics and site conditions in the invaded range. Phragmites australis in North America exists as a mix of populations of different evolutionary origin. Populations of endemic native Phragmites australis americanus are declining, while introduced European populations are important wetland invaders. We assessed soil conditioning effects of native and non‐native P. australis populations on early and late seedling survival of native and introduced wetland plants. We further used a soil biocide treatment to assess the role of soil fungi on seedling survival. Survival of seedlings in soils colonized by P. australis was either unaffected or negatively affected; no species showed improved survival in P. australis‐conditioned soils. Population of P. australis was a significant factor explaining the response of seedlings, but origin (native or non‐native) was not a significant factor. Synthesis: Our results highlight the importance of phylogenetic control when assessing impacts of invasive species to avoid conflating general plant traits with mechanisms of invasive success. Both native (noninvasive) and non‐native (invasive) P. australis populations reduced seedling survival of competing plant species. Because soil legacy effects of native and non‐native P. australis are similar, this study suggests that the close phylogenetic relationship between the two populations, and not the invasive status of introduced P. australis, is more relevant to their soil‐mediated impact on other plant species.     

Phylogeny and assemblage composition of Frankia in Alnus tenuifolia nodules across a primary successional sere in interior Alaska

In nitrogen (N) fixing symbioses, host‐symbiont specificity, genetic variation in bacterial symbionts and environmental variation represent fundamental constraints on the ecology, evolution and practical uses of these interactions, but detailed information is lacking for many naturally occurring N‐fixers. This study examined phylogenetic host specificity of Frankia in field‐collected nodules of two Alnus species (A. tenuifolia and A. viridis) in interior Alaska and, for A. tenuifolia, distribution, diversity, spatial autocorrelation and correlation with specific soil factors of Frankia genotypes in nodules collected from replicated habitats representing endpoints of a primary sere. Frankia genotypes most commonly associated with each host belonged to different clades within the Alnus‐infective Frankia clade, and for A. tenuifolia, were divergent from previously described Frankia. A. tenuifolia nodules from early and late succession habitats harboured distinct Frankia assemblages. In early succession, a single genotype inhabited 71% of nodules with no discernable autocorrelation at any scale, while late succession Frankia were more diverse, differed widely among plants within a site and were significantly autocorrelated within and among plants. Early succession Frankia genotype occurrence was strongly correlated with carbon/nitrogen ratio in the mineral soil fraction, while in late succession, the most common genotypes were correlated with different soil variables. Our results suggest that phylogenetic specificity is a significant factor in the A. tenuifolia‐Frankia interaction and that significant habitat‐based differentiation may exist among A. tenuifolia‐infective genotypes. This is consistent with our hypothesis that A. tenuifolia selects specific Frankia genotypes from early succession soils and that this choice is attenuated in late succession.     

Pinus monophylla establishment in an expanding Pinus‐Juniperus woodland: Environmental conditions,facilitation and interacting factors

Abstract. The tree species comprising Pinus‐Juniperus woodlands are rapidly expanding into shrub‐grasslands throughout their range. Observational studies indicate that establishment is facilitated by nurse plants, but little information exists on the mechanisms involved. I examined both abiotic and biotic factors influencing Pinus monophylla establishment in Artemisia tridentata steppe with expanding populations of P. monophylla and Juniperus osteosperma. I determined soil water contents, temperatures, and nutrient characteristics for the primary establishment microhabitats, i.e. under Pinus, under Juniperus, under Artemisia, tree interspace and sage interspace, and evaluated the emergence and survival response of two seedling cohorts over a 3‐yr period for the different microhabitats. I also examined the effects of seed burial and predation on seedling establishment. Microhabitats under trees and shrubs had higher extractable P and K, higher organic matter, total nitrogen and cation exchange capacity than interspace microhabitats. Soil water contents (0–15 cm) were lower in interspaces than under shrubs or trees due to dry surface (0–5 cm) soils. Soil temperatures (at 1 and 15 cm) were lowest under trees, intermediate under shrubs, and highest in interspaces. Timing and rate of seedling emergence were temperature dependent with the order of emergence paralleling mean growing season temperatures: tree interspace = shrub interspace > under shrub > under Juniperus under Pinus. Seed burial was required for rooting and the highest emergence occurred from depths of 1 and 3 cm indicating that caching by birds and rodents is essential and that animals bury seeds at adequate if not optimal depths for emergence. Seedlings required microenvironmental modification for survival; all seedlings, including those that emerged from seeds and transplants, died within the first year in interspace microhabitats. Survival in under‐tree or under‐shrub microhabitats depended on soil water availability and corresponded closely to soil water contents over the 3‐yr study. Under‐shrub microhabitats had more favourable soil and micro‐environmental characteristics than under‐tree microhabitats and had the highest seedling life spans for the first‐year seedling cohort. Predation of Pinus seedlings by rodents was a significant cause of mortality with caged transplants exhibiting life spans that were 74% longer overall than uncaged transplants. Emergence and survival of P. monophylla within the expanding woodland were dependent upon a complex set of interacting factors including growing season conditions, microhabitat characteristics, and animal species.     

Early succession on plots with the upper soil horizon removed

Abstract. Succession was studied on plots with the upper soil horizon removed in an area affected by acidic air pollution in the Kru?né Hory Mts., Czech Republic. 10 permanent 1‐m² plots were marked and vegetation recorded annually using a grid of 100 subplots from 1989 to 1995. Constrained ordination analyses showed that soil texture is the most important environmental factor influencing the course of succession. Its effect on species composition increases with successional age of the plant community. On fine‐grained soils species‐poor communities dominated by grasses (Calamagrostis villosa, Deschampsiaflexuosa) and on coarse‐grained soils species‐rich communities dominated by heather (Calluna vulgaris) developed. Succession proceeded from communities where species composition was determined by diaspore availability towards communities where species composition depended on environmental conditions. Successional communities after 10 yr are more dependent on soil characteristics and consequently environmental determination increases over the course of succession and causes the communities to diverge.