Vegetation dynamics following resource manipulations in herb-rich woodland

Changes in species richness, turnover, composition and above-ground biomass of herb-rich woodland were documented following fertilizer application and water addition over three growing seasons. Addition of fertilizer significantly reduced species richness relative to unmanipulated control and water addition plots after 3 years. This change coincided with significant increases in biomass, which were largely due to increased growth of exotic annual grasses. The reductions in richness observed in the fertilized plots were a consequence of both lower rates of local colonization and enhanced rates of local extinction of the resident species. Species loss was not random; native species were lost after nutrient addition whereas exotic species were not. Nutrient limitation was more important for species coexistence in these communities than was water availability.     

The Relationship between Local Species Richness and Species Pool: A Case Study from the High Mountains of the Greater Caucasus

The causes of linear relationships between local species richness and the size of the actual species pool in closed subalpine meadow communities and open plant communities of the alpine stony substrate (the Greater Caucasus Mountains) were analyzed using a computer simulation model. The results demonstrated that this relationship is insufficient evidence for the variation of local species richness among communities is wholly or partly determined by regional processes (the species-pool hypothesis). A relatively proportional ratio between these variables can also arise where local species richness and the size of the species pool both depend on local processes, or where local species richness is determined by local factors alone while the size of the species pool is determined by both local and regional factors.     

The link between international trade and the global distribution of invasive alien species

Invasive alien species (IAS) exact large biodiversity and economic costs and are a significant component of human-induced, global environmental change. Previous studies looking at the variation in alien species across regions have been limited geographically or taxonomically or have not considered economics. We used a global invasive species database to regress IAS per-country on a suite of socioeconomic, ecological, and biogeographical variables. We varied the countries included in the regression tree analyses, in order to explore whether certain outliers were biasing the results, and in most of the cases, merchandise imports was the most important explanatory variable. The greater the degree of international trade, the higher the number of IAS. We also found a positive relationship between species richness and the number of invasives, in accord with other investigations at large spatial scales. Island status (overall), country area, latitude, continental position (New World versus Old World) or other measures of human disturbance (e.g., GDP per capita, population density) were not found to be important determinants of a country’s degree of biological invasion, contrary to previous studies. Our findings also provide support to the idea that more resources for combating IAS should be directed at the introduction stage and that novel trade instruments need to be explored to account for this environmental externality.

Restoration of a species‐rich meadow on arable land by transferring meadow blocks

Question: Does transplantation of small blocks of turfs contribute to restoration of species‐rich meadows on fallow land? What is the role of vegetative spread and seedling establishment of meadow plants in the neighbourhood of the transplanted turfs? Location: Bílé Karpaty Mountains, SE Czech Republic. Methods: Twenty‐five meadow blocks, 0.4 m × 0.4 m in size, were transplanted to fallow land (unfertile and dry) and species composition was monitored in the source area, in the transferred turfs and in their neighbourhood for 3 yr. Multivariate analyses were used to assess successional trends. Results: The transferred meadow blocks served as a source of diaspores for the seed‐limited fallow land. Out of 80 transplanted species 17 species spread to adjacent plots within 3 yr. The frequency of plants expanding from the transferred turfs, either vegetatively or by seeds, was relatively low and from 2002 to 2004 a total of 22 species declined in the transferred turfs before expanding to the neighbourhood. Successional trends in species composition of the plots adjacent to the transferred blocks were strongly significant compared with the vegetation of the source area despite the short‐term data used for the evaluation. The role of the seed bank in restoration was negligible. Conclusions: In infertile and dry environments, transplanted turfs may significantly speed up restoration, especially if natural sources of target plant seeds are not available in the neighbourhood. However, the restoration process is long‐term and not all transferred plants can be expected to establish in the fallow land.     

Revegetation following soil disturbance in a California meadow: the role of propagule supply

Revegetation following a disturbance event initially should be constrained by the abundance and types of propagules available at the disturbed site. I tested this idea by conducting two experiments in which I created artificial soil disturbances by excavating or burying pre-existing grassland vegetation. In the first experiment, I varied disturbance intensity (depth), to investigate the consequences for revegetation when numbers of surviving propagules (dormant seeds and bulbs) were altered. In the second experiment, I varied the timing of disturbance, to investigate the consequences when disturbed sites experienced differing exposures to seasonal patterns of clonal growth and seed dispersal. I sampled these experiments from 1991 to 1993, and have interpreted their results using measurements of the seed bank, the bulb bank, and the seed rain. In the first (depth) experiment, bulbs declined in abundance with burial depth and were scarcer in deeper excavations. In contrast, numbers of annual graminoids initially showed no trends with respect to disturbance depth. These results reflect the depth distributions of the seed and bulb banks. Since bulbs occur deeply in the soil, progressively deeper disturbances left fewer survivors. Similarly, perennial graminoids could grow through the shallowest burials. In contrast, since the annual-graminoid-dominated seed bank is concentrated near the soil surface, disturbance depth mattered less to these species: any disturbance removing the surface layer was equally destructive. In the second (timing) experiment, more annual graminoids initially occurred in older plots. This result reflects seasonal patterns of seed production: plots exposed to more of the annual-graminoid-dominated seed rain supported higher densities of annual graminoids as a result. In subsequent years, the vegetation of most plots in both experiments was increasingly dominated by annual graminoids, again as a consequence of their great abundance in the seed rain. These results indicate that interactions between soil disturbances and sources of propagules play an important role in controlling early stages of succession in newly created gaps. They also suggest that disturbance may play different roles in communities characterized by species with different reproductive strategies. Understanding sources of colonists will improve our ability to predict the effects of disturbance.     

The Contribution of Tropical Secondary Forest Fragments to the Conservation of Fruit-feeding Butterflies: Effects of Isolation and Age

Concomitant with the rapid loss of tropical mature forests, the relative abundance of secondary forests is increasing steadily and the latter are therefore of growing interest for conservation. We analysed species richness of fruit-feeding nymphalid butterflies in secondary forest fragments of different age and isolation and in mature forest at the eastern margin of the Lore Lindu National Park in Central Sulawesi, Indonesia. From April to August 2001 we collected 2322 individuals of fruit-feeding butterflies, belonging to 33 species. Butterfly species richness increased with succession, but was significantly higher in mature forests than in all types of secondary forest. Isolation of the forest fragments did not have a significant effect on butterfly species richness in the range of distances (up to 1700 m) studied. Rather it appeared to affect only a few species. Species richness of endemic species was higher than of non-endemic species. Although endemic species were most diverse in mature forests, many species captured were restricted to secondary forests. Our results show that mature forest is essential for the conservation of nymphalid butterflies and for the endemic species in this area. However, considering the relatively large number of species found in these rather small habitat islands, secondary forest fragments, especially older successional stages, can be taken into account in conservation efforts and thus contribute to the preservation of tropical biodiversity on a landscape scale.     

Seed supply of native and cultivated grasses in pine forests of the southwestern United States and the potential for vegetation recovery following wildfire

In this study we examined seed supply (seed banks and seed rain) and vegetation of seeded cultivated grasses and naturally occurring native grasses following a wildfire in northern New Mexico, USA. We specifically examined density of native and cultivated grass seeds and plants in areas of high, moderate, and low fire severity. We also examined the similarity between density of native and cultivated grass seeds to density of above ground plants. Density of native grass seed per square meter was higher in areas that burned under low fire severity (85.18 ± 44.83) compared to areas of moderate (18.52 ± 11.26) and high (7.41 ± 4.90) fire severity; however, differences were not statistically significant due, in part, to the high error associated with estimates. Density of cultivated grass seed per square meter was higher than that of native grass seed in areas of high (439.60 ± 117.98) and moderate (437.02 ± 146.50) fire severity, areas that were seeded with cultivated grasses after the wildfire for erosion control. Density of seeded grass plants per square meter was also higher than that of native grass plants in areas of high (18.78 ± 4.59 versus 0.33 ± 0.24) and moderate (8.22 ± 1.76 versus 0.22 ± 0.15) fire severity. There was a higher correspondence between the density of cultivated grass seeds and plants (highest value 0.32 ± 0.11) compared to density of native grass seeds and plants (highest value 0.05 ± 0.04). The high density of seeds, plants, and correspondence indicated that seeds from cultivated grasses are more likely to establish as post-fire vegetation than seeds from native grasses. Seeding with cultivated grasses following a wildfire may slow or inhibit recovery of native grasses in the short term. Longer-term implications for site occupancy deserve further study.     

Effects of regular and irregular temporal patterns of disturbance on biomass accrual and species composition of a subtidal hard-bottom assemblage

Assessing patterns of species distribution and abundance is important to understand the driving processes of, and predict future changes in, biodiversity. To this date, ecological studies have been mainly designed to investigate the effects of the mean magnitude of predictor variables, although ecological factors naturally vary in space and time. In a nine month long field experiment, we tested the effects of different temporal patterns (regular, lowly and highly irregular) in biomass removal (=disturbance event) on the diversity, species composition, and biomass accrual of macrobenthic assemblages grown on 15 × 15 cm² PVC-panels. For each pattern of disturbance, disturbance events were timed at three sequences to control for possible confounding effects with recruitment patterns. Disturbance intensity was kept identical among treatments. Assemblages developed in the absence of disturbance for 3 months prior to a 150-day manipulation period, during which the biomass from 20% of the panel area was removed at each of ten disturbance events. Additional undisturbed settlement panels were deployed in the field to assess monthly recruitment rates and species succession over a one year period. Disturbance (i) reduced biomass and total species cover, (ii) changed species composition during the first half of the manipulation period significantly, and (iii) was without effect on species richness and evenness. Irregular disturbance regimes enhanced the abundance of the ascidian Ciona intestinalis, biomass accrual, and total species cover of assemblages relative to the regular disturbance regime, but had either no or only transient effects on diversity and species composition, respectively. Neither the degree of irregularity in disturbance nor the sequence of disturbance events affected any of the response variables significantly. Recruitment of species was strongly seasonal with almost only diatoms recruiting during winter, while recruitment was most intense during summer. Our results suggest that the temporal patterns of predictor variables might be of low explanatory power for the variance of responses in communities with seasonal recruitment patterns that are exposed to a high level of disturbance. Thus the need to include temporal patterns of predictor variables in experimental designs may depend on community dynamics and the characteristics of the process under investigation. S. Wollgast and M. Molis contributed equally to this work.     

Competitive displacement or biotic resistance? Disentangling relationships between community diversity and invasion success of tall herbs and shrubs

Questions: Are negative invasion–diversity relationships due to biotic resistance of the invaded plant community or to post‐invasion displacement of less competitive species? Do invasion–diversity relationships change with habitat type or resident traits? Location/species: Lowlands and uplands of western and southern Germany, Heracleum mantegazzianum; mountain range in central Germany, Lupinus polyphyllus; and coastal dunes of northwest Germany, Rosa rugosa. Methods: We tested the significance and estimated regression slopes of invasion–diversity relationships using generalized linear (mixed effects) models relating invader cover and habitat type to species richness in different plant groups, stratified based on size, life cycle and community association. Results: We found negative, positive and neutral relationships between invader cover and species richness. There were negative linear correlations of invader cover with small plant species throughout, but no negative linear correlation with tall species. Invasion–diversity relationships tended to be more negative in early‐successional habitats, such as dunes or abandoned grasslands, than in late‐successional habitats. Conclusions: Invasion diversity–relationships are complex; they vary among habitat types and among different groups of resident species. Negative invasion–diversity relationships are due to asymmetric competitive displacement of inferior species and not due to biotic resistance. Small species are displaced in early‐successional habitats, while there is little effect on persistence of tall species.     

Variation in exotic and native seed arrival and recruitment of bird dispersed species in subtropical forest restoration and regrowth

Invasive bird-dispersed plants often share the same suite of dispersers as co-occurring native species, resulting in a complex management issue. Integrated management strategies could incorporate manipulation of dispersal or establishment processes. To improve our understanding of these processes, we quantified seed rain, recruit and seed bank density, and species richness for bird-dispersed invasive and native species in three early successional subtropical habitats in eastern Australia: tree regrowth, shrub regrowth and native restoration plantings. We investigated the effects of environmental factors (leaf area index (LAI), distance to edge, herbaceous ground cover and distance to nearest neighbour) on seed rain, seed bank and recruit abundance. Propagule availability was not always a good predictor of recruitment. For instance, although native tree seed rain density was similar, and species richness was higher, in native plantings, compared with tree regrowth, recruit density and species richness were lower. Native plantings also received lower densities of invasive tree seed rain than did tree regrowth habitats, but supported a similar density of invasive tree recruits. Invasive shrub seed rain was recorded in highest densities in shrub regrowth sites, but recruit density was similar between habitats. We discuss the role of microsite characteristics in influencing post-dispersal processes and recruit composition, and suggest ways of manipulating these processes as part of an integrated management strategy for bird-dispersed weeds in natural areas.