The fate of the missing spores

It is well-known that many species with small diaspores can disperse far during extended temporal scales (many years). However, studies on short temporal scales usually only cover short distances (in, e.g., bryophytes up to 15 m). By using a novel experimental design, studying the realized dispersal, we extend this range by almost two orders of magnitude. We recorded establishment of the fast-growing moss Discelium nudum on introduced suitable substrates, placed around a translocated, sporulating mother colony. Around 2,000 pots with acidic clay were placed at different distances between 5 m and 600 m, in four directions, on a raised bog, with increased pot numbers with distance. The experiment was set up in April-May and the realized dispersal (number of colonized pots) was recorded in September. Close to the mother colony (up to 10 m), the mean colonization rates (ratio of colonized pots) exceeded 50%. At distances between 10 and 50 m colonization dropped sharply, but beyond 50 m the mean colonization rates stabilized and hardly changed (1-3%). The estimated density of spores causing establishments at the further distances (2-6 spores/m2) was realistic when compared to the estimated spore output from the central colonies. Our study supports calculations from earlier studies, limited to short distances, that a majority of the spores disperse beyond the nearest vicinity of a source. The even colonization pattern at further distances raises interesting questions about under what conditions spores are transported and deposited. However, it is clear that regular establishment is likely at the km-scale for this and many other species with similar spore output and dispersal mechanism

Patch area and current coffee management determine woody plant diversity in patches of semi-forest coffee embedded in an agricultural matrix

AbstractEffective conservation of biodiversity in patches of (semi-) natural vegetation is dependent on an understanding of the influence of management as well as spatial and temporal factors. In southwestern Ethiopia coffee generally grown under a rather dense layer of indigenous trees (so called semi-forest coffee - SFC) often in patches embedded in an open agricultural landscape. The aim of the study was to disentangle what governs the variation in species richness of woody species among such patches. We collected data on species and possible explanatory factors in 40~×~40 m plots centered in 40 SFC patches, measured the patch area for 1987 and 2013, and the amount of surrounding SFC-area for each patch. We recorded the number of coffee stems and the level of disturbance caused by slashing of the understory vegetation. Species richness of large coffee shade trees (>20 cm in diameter) was higher in larger patches with even slightly better fit of the statistical models when the historical area was taken into account. However, most species of large trees also occurred as seedlings showing that there is still a potential to conserve these species in the patches. Coffee management negatively affected the richness and density of woody species, especially in the intermediate size class (1.6–20~cm diameter). Disturbances accompanying coffee management such as slashing of the ground vegetation also negatively affected tree seedling density as well as species richness. There was no effect of connectivity on species richness. Based on the combination of these results we conclude that small patches of semi-forest coffee had fewer species of large trees, not because of a lack of tree seedlings, but probably because of differentiated local extinctions, perhaps during the time when the species were intermediate sized. To maintain the species richness of large trees in semi-forest coffee patches, the sites need to be actively managed

Monthly microclimate models in a managed boreal forest landscape

The majority of microclimate studies have been done in topographically complex landscapes to quantify and predict how near-ground temperatures vary as a function of terrain properties. However, in forests understory temperatures can be strongly influenced also by vegetation. We quantified the relative influence of vegetation features and physiography (topography and moisture-related variables) on understory temperatures in managed boreal forests in central Sweden. We used a multivariate regression approach to relate near-ground temperature of 203 loggers over the snow-free seasons in an area of ∼16,000 km2 to remotely sensed and on-site measured variables of forest structure and physiography. We produced climate grids of monthly minimum and maximum temperatures at 25m resolution by using only remotely sensed and mapped predictors. The quality and predictions of the models containing only remotely sensed predictors (MAP models) were compared with the models containing also on-site measured predictors (OS models). Our data suggest that during the warm season, where landscape microclimate variability is largest, canopy cover and basal area were the most important microclimatic drivers for both minimum and maximum temperatures, while physiographic drivers (mainly elevation) dominated maximum temperatures during autumn and early winter. The MAP models were able to reproduce findings from the OS models but tended to underestimate high and overestimate low temperatures. Including important microclimatic drivers, particularly soil moisture, that are yet lacking in a mapped form should improve the microclimate maps. Because of the dynamic nature of managed forests, continuous updates of mapped forest structure parameters are needed to accurately predict temperatures. Our results suggest that forest management (e.g. stand size, structure and composition) and conservation may play a key role in amplifying or impeding the effects of climate-forcing factors on near-ground temperature and may locally modify the impact of global warming.Peer reviewe

Warm range margin of boreal bryophytes and lichens not directly limited by temperatures

Species at their warm range margin are potentially threatened by higher temperatures, but may persist in microrefugia. Whether such microsites occur due to more suitable microclimate or due to lower biotic pressure from, for example competitive species, is still not fully resolved. We examined whether boreal bryophytes and lichens show signs of direct climate limitation, that is whether they perform better in cold and/or humid microclimates at their warm range margin. We transplanted a moss, a liverwort and a lichen to 58 boreal forest sites with different microclimates at the species' southern range margin in central Sweden. Species were grown in garden soil patches to control the effects of competitive exclusion and soil quality. We followed the transplanted species over three growing seasons (2016-2018) and modelled growth and vitality for each species as a function of subcanopy temperature, soil moisture, air humidity and forest type. In 2018, we also recorded the cover of other plants having recolonized the garden soil patches and modelled this potential future competition with the same environmental variables plus litter. Species performance increased with warmer temperatures, which was often conditional on high soil moisture, and at sites with more conifers. Soil moisture had a positive effect, especially on the moss in the last year 2018, when the growing season was exceptionally hot and dry. The lichen was mostly affected by gastropod grazing. Recolonization of other plants was also faster at warmer and moister sites. The results indicate that competition, herbivory, shading leaf litter and water scarcity might be more important than the direct effects of temperature for performance at the species' warm range margin. Synthesis. In a transplant experiment with three boreal understorey species, we did not find signs of direct temperature limitation towards the south. Forest microrefugia, that is habitats where these species could persist regional warming, may instead be sites with fewer competitors and enemies, and with sufficient moisture and more conifers in the overstorey.Peer reviewe

Forest edge effects on moss growth are amplified by drought

Forest fragmentation increases the amount of edges in the landscape. Differences in wind, radiation, and vegetation structure create edge-to-interior gradients in forest microclimate, and these gradients are likely to be more pronounced during droughts and heatwaves. Although the effects of climate extremes on edge influences have potentially strong and long-lasting impacts on forest understory biodiversity, they are not well understood and are not often considered in management and landscape planning. Here we used a novel method of retrospectively quantifying growth to assess biologically relevant edge influences likely caused by microclimate using Hylocomium splendens, a moss with annual segments. We examined how spatio-temporal variation in drought across 3 years and 46 sites in central Sweden, affected the depth and magnitude of edge influences. We also investigated whether edge effects during drought were influenced by differences in forest structure. Edge effects were almost twice as strong in the drought year compared to the non-drought years, but we did not find clear evidence that they penetrated deeper into the forest in the drought year. Edge influences were also greater in areas that had fewer days with rain during the drought year. Higher levels of forest canopy cover and tree height buffered the magnitude of edge influence in times of drought. Our results demonstrate that edge effects are amplified by drought, suggesting that fragmentation effects are aggravated when droughts become more frequent and severe. Our results suggest that dense edges and buffer zones with high canopy cover can be important ways to mitigate negative drought impacts in forest edges

Structurally complex farms support high avian functional diversity in tropical montane Ethiopia

Of all feeding guilds, understorey insectivores are thought to be most sensitive to disturbance and forest conversion. We compared the composition of bird feeding guilds in tropical forest fragments with adjacent agro-ecosystems in a montane region of south-west Ethiopia. We used a series of point counts to survey birds in 19 agriculture and 19 forest sites and recorded tree species within each farm across an area of 40 × 35 km. Insectivores (~17 spp. per plot), frugivores (~3 spp. per plot) and omnivores (~5 spp. per plot) maintained species density across habitats, while granivores and nectarivores increased in the agricultural sites by factors of 7 and 3 respectively. Species accumulation curves of each guild were equal or steeper in agriculture, suggesting that agricultural and forest landscapes were equally heterogeneous for all bird guilds. Counter to most published studies, we found no decline in insectivore species richness with forest conversion. However, species composition differed between the two habitats, with certain forest specialists replaced by other species within each feeding guild. We suggest that the lack of difference in insectivorous species numbers between forest and agriculture in this region is due to the benign nature of the agricultural habitat, but also due to a regional species pool which contains many bird species which are adapted to open habitats

Effects of past and present microclimates on northern and southern plant species in a managed forest landscape

Questions: Near-ground temperatures can vary substantially over relatively short distances, enabling species with different temperature preferences and geographical distributions to co-exist within a small area. In a forest landscape, the near-ground temperatures may change due to management activities that alter forest density. As a result of such management activities, current species distributions and performances might not only be affected by current microclimates, but also by past conditions due to time-lagged responses. Location: Sweden. Methods: We examined the effects of past and current microclimates on the distributions and performances of two northern, cold-favoured, and two southern, warm-favoured, plant species in 53 managed forest sites. Each pair was represented by one vascular plant and one bryophyte species. We used temperature logger data and predictions from microclimate models based on changes in basal area to relate patterns of occurrence, abundance, and reproduction to current and past microclimate. Results: The two northern species were generally favoured by microclimates that were currently cold, characterised by later snowmelt and low accumulated heat over the growing season. In contrast, the two southern species were generally favoured by currently warm microclimates, characterised by high accumulated heat over the growing season. Species generally had higher abundance in sites with a preferred microclimate both in the past and present, and lower abundance than expected from current conditions, if the past microclimate had changed from warm to cold or vice versa, indicating time-lags in abundance patterns of the species. Conclusions: Our results show a potential importance of past and present microclimate heterogeneity for the co-existence of species with different temperature preferences in the same landscape and highlight the possibility to manage microclimates to mitigate climate change impacts on forest biodiversity

Edge influence on vegetation at natural and anthropogenic edges of boreal forests in Canada and Fennoscandia

Although anthropogenic edges are an important consequence of timber harvesting, edges due to natural disturbances or landscape heterogeneity are also common. Forest edges have been well studied in temperate and tropical forests, but less so in less productive, disturbance-adapted boreal forests. We synthesized data on forest vegetation at edges of boreal forests and compared edge influence among edge types (fire, cut, lake/wetland; old vs. young), forest types (broadleaf vs. coniferous) and geographic regions. Our objectives were to quantify vegetation responses at edges of all types and to compare the strength and extent of edge influence among different types of edges and forests. Research was conducted using the same general sampling design in Alberta, Ontario and Quebec in Canada, and in Sweden and Finland. We conducted a meta-analysis for a variety of response variables including forest structure, deadwood abundance, regeneration, understorey abundance and diversity, and non-vascular plant cover. We also determined the magnitude and distance of edge influence (DEI) using randomization tests. Some edge responses (lower tree basal area, tree canopy and bryophyte cover; more logs; higher regeneration) were significant overall across studies. Edge influence on ground vegetation in boreal forests was generally weak, not very extensive (DEI usually < 20 m) and decreased with time. We found more extensive edge influence at natural edges, at younger edges and in broadleaf forests. The comparison among regions revealed weaker edge influence in Fennoscandian forests. Synthesis. Edges created by forest harvesting do not appear to have as strong, extensive or persistent influence on vegetation in boreal as in tropical or temperate forested ecosystems. We attribute this apparent resistance to shorter canopy heights, inherent heterogeneity in boreal forests and their adaptation to frequent natural disturbance. Nevertheless, notable differences between forest structure responses to natural (fire) and anthropogenic (cut) edges raise concerns about biodiversity implications of extensive creation of anthropogenic edges. By highlighting universal responses to edge influence in boreal forests that are significant irrespective of edge or forest type, and those which vary by edge type, we provide a context for the conservation of boreal forests. Edges created by forest harvesting do not appear to have as strong, extensive or persistent influence on vegetation in boreal as in tropical or temperate forested ecosystems. We attribute this apparent resistance to shorter canopy heights, inherent heterogeneity in boreal forests and their adaptation to frequent natural disturbance. Nevertheless, notable differences between forest structure responses to natural (fire) and anthropogenic (cut) edges raise concerns about biodiversity implications of extensive creation of anthropogenic edges. By highlighting universal responses to edge influence in boreal forests that are significant irrespective of edge or forest type, and those which vary by edge type, we provide a context for the conservation of boreal forests

Multi-scale mosaics in top-down pest control by ants from natural coffee forests to plantations

While top-down control plays an important role in shaping both natural and agricultural food webs, we lack insights into how top-down control effects vary across spatial scales. We used a multi-scale survey of top-down control of coffee pests and diseases by arboreal ants to examine if colony location creates a small-scale mosaic in top-down control around trees and if the strength of that control varies between sites at the landscape scale. We investigated pest and disease levels on coffee shrubs at different distances from shade trees with and without a Crematogaster spp. ant colony in 59 sites along a coffee management intensity gradient in southwestern Ethiopia. Within sites, ants significantly suppressed herbivory and coffee leaf rust at distances less than 10 m from nesting trees. Top-down control varied between sites, with stronger top-down control of free-feeding herbivory near ant colonies at sites with lower management intensity and stronger top-down control of a skeletonizer at sites with higher canopy cover. We conclude that the strength of top-down control by ants is highly heterogeneous across spatial scales, as a consequence of the biology of the predator at the small scale and herbivore density or changes in herbivore-ant interactions at the landscape scale