Spatio‐temporal variation in the effect of herbaceous layer on woody seedling survival in a Chilean mediterranean ecosystem

Questions: What is the effect of herbaceous layer on seedling establishment of three woody pioneer species in open areas of central Chile under a semi‐arid mediterranean climate? How do inter‐annual and habitat conditions (slope aspect) modulate this effect? Under high stress conditions such as the drier year and habitat (north‐facing slope) do herbs reach low abundance and have neutral effects on woody seedlings? Under medium stress conditions for these woody species, such as the wetter year and south‐facing slope, does the herbaceous layer reach greater abundance and have positive effects on woody seedlings due to increasing soil water content? Location: A watershed on the outskirts of Santiago, Chile, subjected to clearing of woody vegetation through firewood extraction and human‐set fires. Methods: In spring 2007, we set up 20 plots (3 m × 2 m). Half of each plot had herbs removed manually and by application of herbicide. In both halves of each plot, one seedling (8 months old) of each of the three native woody species (Colliguaya odorifera, Schinus polygamus and Quillaja saponaria) was planted and survival monitored subsequently. The experiment was repeated in two consecutive growing seasons (2007–2008 and 2008–2009) that differed significantly in total precipitation (152 and 256.5 mm, respectively), and replicated in two sites that differed in aspect and abiotic conditions: a moister south‐ and a drier north‐facing slope. Results: In the first and drier year, the herbaceous layer had low cover and no significant effect on seedling survival of woody species. During the second year, herbs had greater cover and a significant positive effect on spring survival of C. odorifera in the north‐facing slope, which was lost after summer. During this wetter year on the south‐facing slope, herb cover had a positive effect on survival of S. polygamus (mainly during summer). Conclusions: The role of mostly ruderal herbs on woody seedling establishment depended on the species, rainfall of the current year and slope aspect, and may be explained by soil moisture patterns. This suggests that the effect of ruderal herbs on woody seedlings shifts from neutral under high stress conditions produced by drought to positive under moderate stress conditions. Our results contribute to understand interactions between ruderal herbs and woody species under contrasting abiotic conditions. Therefore, control of the herbaceous layer may not be needed in restoration programmes for this region. Moreover, herbs may benefit restoration of woody cover in mesic habitats.     

Plant–plant interactions,environmental gradients and plant diversity: A global synthesis of community-level studies

Previous syntheses on the effects of environmental conditions on the outcome of plant–plant interactions summarize results from pairwise studies. However, the upscaling to the community-level of such studies is problematic because of the existence of multiple species assemblages and species-specific responses to both the environmental conditions and the presence of neighbors. We conducted the first global synthesis of community-level studies from harsh environments, which included data from 71 alpine and 137 dryland communities to: (i) test how important are facilitative interactions as a driver of community structure, (ii) evaluate whether we can predict the frequency of positive plant–plant interactions across differing environmental conditions and habitats, and (iii) assess whether thresholds in the response of plant–plant interactions to environmental gradients exists between “moderate” and “extreme” environments. We also used those community-level studies performed across gradients of at least three points to evaluate how the average environmental conditions, the length of the gradient studied, and the number of points sampled across such gradient affect the form and strength of the facilitation-environmental conditions relationship. Over 25% of the species present were more spatially associated to nurse plants than expected by chance in both alpine and dryland areas, illustrating the high importance of positive plant–plant interactions for the maintenance of plant diversity in these environments. Facilitative interactions were more frequent, and more related to environmental conditions, in alpine than in dryland areas, perhaps because drylands are generally characterized by a larger variety of environmental stress factors and plant functional traits. The frequency of facilitative interactions in alpine communities peaked at 1000 mm of annual rainfall, and globally decreased with elevation. The frequency of positive interactions in dryland communities decreased globally with water scarcity or temperature annual range. Positive facilitation-drought stress relationships are more likely in shorter regional gradients, but these relationships are obscured in regions with a greater species turnover or with complex environmental gradients. By showing the different climatic drivers and behaviors of plant–plant interactions in dryland and alpine areas, our results will improve predictions regarding the effect of facilitation on the assembly of plant communities and their response to changes in environmental conditions.     

Relationships between species’ floral traits and pollinator visitation in a temperate grassland

Knowledge about plant–plant interactions for pollinator service at the plant community level is still scarce, although such interactions may be important to seed production and hence the population dynamics of individual plant species and the species compositions of communities. An important step towards a better understanding of pollination interactions at the community level is to assess if the variation in floral traits among plant species explain the variation in flower visitation frequency among those species. We investigated the relative importance of various floral traits for the visitation frequency of all insects, and bumblebees and flies separately, to plant species by measuring the visitation frequency to all insect-pollinated species in a community during an entire flowering season. Visitation frequency was identified to be strongly positive related to the visual display area and the date of peak flowering of plant species. Categorical variables, such as flower form and symmetry, were important to the visitation frequency of flies only. We constructed floral similarity measures based on the species’ floral traits and found that the floral similarity for all species’ traits combined and the continuous traits separately were positively related to individual visitation frequency. On the other hand, plant species with similar categorical floral traits did not have similar visitation frequencies. In conclusion, our results show that continuous traits, such as flower size and/or density, are more important for the variation in visitation frequency among plant species than thought earlier. Furthermore, differences in visitation frequency among pollinator groups give a poor support to the expectations derived from the classical pollination syndromes.     

From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.