Delayed response in a plant-pollinator system to experimental grassland fragmentation

The fragmentation of natural habitat is considered to be a major threat to biodiversity. Decreasing habitat quality and quantity caused by fragmentation may lead to a disruption of plant-pollinator interactions and to a reduction in sexual reproduction in plant species. We conducted a 6-year field experiment to investigate the effects of small-scale fragmentation on plant-pollinator interactions and genetic diversity in the self-compatible Betonica officinalis. We examined the abundance and composition of pollinators, the foraging behaviour of bumblebees and the performance, outcrossing rate and genetic diversity of B. officinalis after 2 and 6years in experimentally fragmented nutrient-poor, calcareous grassland in the northern Swiss Jura mountains. Fragments of different size (2.25 and 20.25m2) were isolated by a 5-m-wide strip of frequently mown vegetation. Control plots of corresponding size were situated in adjacent undisturbed grassland. Experimental grassland fragmentation altered the composition of B. officinalis pollinators and reduced their flower visitation rate. Furthermore, the foraging behaviour of bumblebees was changed in the fragments. After 6years of fragmentation seed weight was higher in fragments than in control plots. However, the densities of B. officinalis rosettes and inflorescences, plant height and inflorescence length were not affected by fragmentation. The outcrossing frequency of B. officinalis growing in fragments was reduced by 15% after 2years and by 33% after 6years of experimental fragmentation. This resulted in a significant reduction of the genetic diversity in seedlings emerging in fragments after 6years. Our study shows that small-scale habitat fragmentation can disturb the interaction between B. officinalis and pollinators resulting in a reduced outcrossing frequency and genetic diversity in plants growing in fragments. However, the response to fragmentation was considerably delayed. This finding strengthens the claim for long-term field experiments with proper replications and controls to assess delayed effects of habitat fragmentatio

Population structure and genetic diversity of relict populations of Alyssum montanum on limestone cliffs in the Northern Swiss Jura mountains

Exposed cliffs in the Northern Swiss Jura mountains harbour a highly diverse flora with numerous rare and relict plant species. The genetic structure of cliff populations is of particular interest because in a variety of plant species the populations are small, isolated and separated from their main distribution area in the Alps and the Mediterranean. We examined possible relationships between population size and size structure, sexual reproduction and genetic diversity of Alyssum montanum, a relict plant species occurring on limestone cliffs in the Northern Swiss Jura mountains. The population size of A. montanum ranged from 8 to 248 plant individuals on the 12 cliffs examined. Most populations contained a high proportion of small-sized (=young) plants indicating a successful establishment of seedlings. Fitness-related traits of A. montanum (percentage of reproductive individuals, number of fruits, fruit weight) varied widely between cliffs and were neither related to the size of the populations nor to the percentage of vegetation cover on the cliffs. RAPD-PCR analysis revealed that A. montanum populations exhibit a remarkably high genetic diversity. However, genetic diversity decreased with decreasing population size. Moreover, the positive relationship found between genetic variability and fitness-related traits indicates that population size is a key factor for the persistence of A. montanum on limestone cliff

Elevated carbon dioxide increases nectar production in Epilobium angustifolium L

Effects of elevated CO2 and nutrient availability on nectar production and onset of flowering in five different seed families (genotypes) of Epilobium angustifolium were investigated in a greenhouse experiment. Elevated CO2 significantly increased nectar production per day (+51%, p > 0.01), total sugar per flower (+41%, p > 0.05), amino acid concentration (+65%, p > 0.05) and total amino acids per flower (+192%, p > 0.001). All other parameters tested, i.e., nectar sugar concentration, proportion of glucose/fructose and proportion of sucrose/(glucose + fructose), were not significantly affected by elevated CO2 and/or fertilization. However, elevated CO2 caused a marginally significant trend for earlier flowering in highly fertilized plants. No significant family x CO2 interaction was found in any of the tested parameters, but the response in nectar production varied considerably among seed families (+10 to +104%) and was significantly positive in two of the five seed families investigated. Our results are not consistent with earlier studies on effects of elevated CO2 on nectar production and flowering phenology in other plant species. It seems, on the other hand, that CO2 effects on nectar production are specific to species and genotype. Hence, no general conclusions about effects of elevated CO2 on these floral traits can be drawn at present, but it must be cautioned that elevated CO2 might not only increase floral rewards as in E. angustifolium, but might also lead to shifts or even disruptions in fine-tuned plant-pollinator interactions

Elevated carbon dioxide increases nectar production in Epilobium angustifolium L

Effects of elevated CO2 and nutrient availability on nectar production and onset of flowering in five different seed families (genotypes) of Epilobium angustifolium were investigated in a greenhouse experiment. Elevated CO2 significantly increased nectar production per day (+51%, p < 0.01), total sugar per flower (+41%, p < 0.05), amino acid concentration (+65%, p < 0.05) and total amino acids per flower (+192%, p < 0.001). All other parameters tested, i.e., nectar sugar concentration, proportion of glucose/fructose and proportion of sucrose/(glucose + fructose), were not significantly affected by elevated CO2 and/or fertilization. However, elevated CO2 caused a marginally significant trend for earlier flowering in highly fertilized plants. No significant family × CO2 interaction was found in any of the tested parameters, but the response in nectar production varied considerably among seed families (+10 to +104%) and was significantly positive in two of the five seed families investigated. Our results are not consistent with earlier studies on effects of elevated CO2 on nectar production and flowering phenology in other plant species. It seems, on the other hand, that CO2 effects on nectar production are specific to species and genotype. Hence, no general conclusions about effects of elevated CO2 on these floral traits can be drawn at present, but it must be cautioned that elevated CO2 might not only increase floral rewards as in E. angustifolium, but might also lead to shifts or even disruptions in fine-tuned plant-pollinator interaction

Effects of Long-Term Trampling on the Above-Ground Forest Vegetation and Soil Seed Bank at the Base of Limestone Cliffs

Exposed limestone cliffs in central Europe harbor a highly divers flora with many rare and endangered species. During the past few decades, there has been increasing recreational use of these cliffs, which has caused local environmental disturbances. Successful restoration strategies hinge on identifying critical limitations. We examined the composition of aboveground forest vegetation and density and species composition of seeds in the soil seed bank at the base of four limestone cliffs in mixed deciduous forests that are intensively disturbed by human trampling and at four undisturbed cliffs in the Jura Mountains in northwestern Switzerland. We found that long-term human trampling reduced total aboveground vegetation cover at the base of cliffs and caused a significant shift in the plant-species composition. Compared with undisturbed cliffs, total seed density was lower in disturbed cliffs. Human trampling also altered the species composition of seeds in the soil seed bank. Seeds of unintentionally introduced, stress-tolerant, and ruderal species dominated the soil seed bank at the base of disturbed cliffs. Our findings indicate that a restoration of degraded cliff bases from the existing soil seed bank would result in a substantial change of the original unique plant composition. Active seed transfer, or seed flux from adjacent undisturbed forest areas, is essential for restoration succes

Forest Perception and Knowledge of Hikers and Mountain Bikers in Two Different Areas in Northwestern Switzerland

Recreational activities can have major impacts on vegetation and wildlife in frequently visited forests. We assessed forest perception and knowledge (state, functions, and species diversity) among hikers and mountain bikers in a frequently visited, seminatural suburban recreation forest (Muttenz) and in a more distantly situated, naturally grown excursion forest (Wasserfallen) in northwestern Switzerland. In all, 239 hikers and 126 mountain bikers were interviewed. Mountain bikers in both forests and hikers in the more intensely used recreation forest at Muttenz assessed the state of the forest less optimistically and showed a higher awareness of the negative impact of recreational activities on the flora and fauna than hikers at Wasserfallen. Furthermore, mountain bikers seemed aware of the social conflicts caused by their activity, since they appreciated neutral or positive encounters with other forest visitors. In contrast, 57% of hikers at Muttenz reported on negative experiences with other forest visitors, particularly with mountain bikers. In general, the interviewees' ecological and biological forest knowledge (forest type and function, species diversity) was rather high. A large proportion was aware of the pros and cons of different forest conditions for plants and animals, and could name or recognize at least some plant and/or animal species typical for the visited forest. The forest knowledge was neither influenced by the type of recreational activity carried out nor by any aspect of forest visit behavior (frequency and duration of forest visits, means of transportation and travel distance to forest). However, the interviewees' forest knowledge was influenced by their age and educational leve

Small-scale experimental habitat fragmentation reduces colonization rates in species-rich grasslands

Habitat fragmentation is one of the most important threats to biodiversity. Decreasing patch size may lead to a reduction in the size of populations and to an increased extinction risk of remnant populations. Furthermore, colonization rates may be reduced in isolated patches. To investigate the effects of isolation and patch size on extinction and colonization rates of plant species, calcareous grasslands at three sites in the Swiss Jura Mountains were experimentally fragmented into patches of 0.25, 2.25, and 20.25m2 by frequent mowing of the surrounding area from 1993 to 1999. Species richness in the fragment plots and adjacent control plots of the same sizes was recorded during these 7years. In agreement with the theory of island biogeography, colonization rate was reduced by 30% in fragments versus non-isolated controls, and extinction increased in small versus large plots. Habitat specialists, in contrast to generalists, were less likely to invade fragments. In the last 4years of the experiment, extinction rates tended to be higher in fragment than in control plots at two of the three sites. Despite reduced colonization rates and a tendency of increased extinction rates in fragments, fragmented plots had only marginally fewer species than control plots after 7years. Hence, rates were a more sensitive measure for community change than changes in species richness per se. From a conservation point of view, the detected reduced colonization rates are particularly problematic in small fragments, which are more likely to suffer from high extinction rates in the long ru

Disturbance of forest by trampling: Effects on mycorrhizal roots of seedlings and mature trees of Fagus sylvatica

The effects of disturbance by recreational activities (trampling) on changes in soil organic matter (SOM) and on mycorrhizal roots of seedlings and mature trees were studied in four stands of a beech (Fagus sylvatica L.) forest near Basel, Switzerland. At each site, comparable disturbed and undisturbed plots were selected. Disturbance reduced ground cover vegetation and leaf litter. Beech seedlings had lower biomass after disturbance. Ergosterol concentration in seedling roots, an indicator of mycorrhizal fungi, was lower in two of the four disturbed plots compared to undisturbed plots; these two disturbed sites had especially low litter levels. Based on ergosterol measurements, mycorrhizas of mature trees did not appear to be negatively affected by trampling. Total fine roots and SOM were higher in the disturbed than in the undisturbed plots at three sites. At the fourth site, fine roots and SOM in the disturbed areas were lower than in the undisturbed areas most probably due to nutrient input following picnic activities. Principal component analysis revealed a close correlation between SOM and fine roots of mature trees as well as litter and seedling biomass. Trampling due to recreational activities caused considerable damage to the vegetation layer and in particular to the beech seedlings and their mycorrhizal fine roots, whereas, roots of mature trees were apparently resilient to tramplin

Effects of Fireplace Use on Forest Vegetation and Amount of Woody Debris in Suburban Forests in Northwestern Switzerland

Urban forests are popular recreation areas in Europe. Several of these temperate broad-leaved forests also have a high conservation value due to sustainable management over many centuries. Recreational activities, particularly the use of fireplaces, can cause extensive damage to soil, ground vegetation, shrubs, and trees. Firewood collection depletes woody debris, leading to a loss of habitat for specialized organisms. We examined the effects of fireplace use on forest vegetation and the amount of woody debris by comparing disturbed and control plots in suburban forests in northwestern Switzerland. At frequently used fireplaces, we found reduced species densities in the ground vegetation and shrub layer and changes in plant species composition due to human trampling within an area of 150-200m2. Picnicking and grilling also reduced the height and changed the age structure of shrubs and young trees. The amount of woody debris was lower in disturbed plots than in control plots. Pieces of wood with a diameter of 0.6-7.6cm were preferentially collected by fireplace users. The reduction in woody debris volume extended up to a distance of 16m from the fire ring, covering an area of 800m2 at each picnic site. In order to preserve the ecological integrity of urban forests and to maintain their attractiveness as important recreation areas, we suggest depositing logging residues to be used as firewood and to restrict visitor movements near picnic site

Owners' perceptions do not match actual ground-dwelling invertebrate diversity in their gardens

Urban gardens are important for human well-being, biodiversity and other ecosystem functions. Biodiversity-promoting initiatives would benefit from their owners being aware of the state of biodiversity in their gardens. We examined whether garden owners' perceptions match actual biodiversity in their gardens and whether perceptions are influenced by the owners' ecological knowledge. We used a structured interview to assess the motivations and biodiversity knowledge of owners of 33 domestic gardens in the city of Basel (Switzerland) and related them to a survey of native plants and several groups of ground-dwelling invertebrates in their gardens. Owners showed different priorities, with promotion of habitat for biodiversity, receiving, on average, higher scores than cultivation, recreation and garden designing. Owners prioritizing biodiversity promotion had gardens with high habitat richness. The garden owners' perceptions of both native plant and overall invertebrate diversity were not correlated with actual diversity data for native plants and ground-dwelling invertebrates. The perceptions of the abundance of invertebrate groups by garden owners with good biodiversity knowledge were not more accurate than those from owners with less knowledge. Despite their willingness, many owners do not know all the opportunities to promote biodiversity. Initiatives to further biodiversity-friendly gardening should thus transfer knowledge