Relative importance of perch and facilitative effects on nucleation in tropical woodland in Malawi

Individual trees in open vegetation such as woodlands can act as “nuclei” for the colonization of forest tree species, which consequently lead to the formation of forest patches. This phenomenon is known as nucleation. The mechanism of nucleation is generally attributed to two factors: trees provide perches for frugivores that increase seed deposition (perch effect), and tree crowns ameliorate environmental conditions, which improves seedling establishment (facilitative effect). Few studies have attempted to distinguish the relative importance of these two factors. In this study, I separated these two effects in a woodland in northern Malawi. I chose Ficus natalensis as a potential nuclei tree because large individuals of this species are commonly located at the center of forest patches within open woodland at the study site. I monitored several environmental variables, seedling survival, seedling composition, and seed rain at three microsites: under F. natalensis, under Brachystegia floribunda (a dominant woodland species), and in open sites. Both tree species provided similar favorable conditions for the establishment of forest species compared to open sites. Thus, the survival of forest tree seedlings under F. natalensis and B. floribunda was similar, and substantially higher than seedling survival in open sites. However, communities of naturally occurring seedlings differed significantly between F. natalensis and B. floribunda. These results indicate that the facilitative effect alone cannot explain the nucleation pattern. I attribute this result to the perch effect of F. natalensis because the forest seedling species recorded under F. natalensis reportedly have small, brightly colored diaspores, which are indicative of dispersal by birds. Seed deposition of forest species under F. natalensis was significantly higher than that under B. floribunda or in open sites. My findings reinforce the idea that trees will lead to nucleation when they enhance seed deposition and have a positive effect on the post-dispersal stage.     

Nurses experience reciprocal fitness benefits from their distantly related facilitated plants

It is well known that many plants benefit from growing beside a nurse plant of another species, but the possibility that the nurse also benefits has been rarely studied. We hypothesize that positive interactions are maintained not only because of the recruitment benefits for the facilitated plants but also because of fitness benefits for the nurse plant. We tested this hypothesis by comparing seed production, seed predation and seed viability of a dominant nurse plant species (Mimosa luisana) when growing alone and in patches surrounded by its facilitated species. We also tested whether fitness of the nurse species is dependent on the phylogenetic neighborhood formed by their facilitated species using an analysis that accounted for the abundance and pairwise phylogenetic similitude of all species in each patch. Nurses growing associated to their facilitated species produced more seeds (1.86 times) and these seeds were more viable (1.47 times) than those of nurses growing alone. Seed predation did not alter these fitness differences. Seed number and viability increased in phylogenetically diverse neighborhoods. We conclude that distantly related partners are more likely to cause reciprocal increases in fitness, and that such effects contribute to species coexistence.     

Regional and local variation in seedling emergence,mortality and recruitment of a perennial herb in Mediterranean mountain habitats

Spatial patterns of seedling regeneration of woody Mediterranean plants are widely documented, with seedlings and juveniles growing almost exclusively under shrub and tree canopy. Neighbourhood habitat amelioration and consequent facilitative effects on recruitment have been extensively suggested as the major determinant of such spatial pattern for Mediterranean vegetation. Much less is known on recruitment patterns of perennial herbs. As herbs differ from woody plants in many relevant ecological aspects and in their life cycles, we would expect particularities in their recruitment dynamics. We analysed the spatial (regional, local and fine-scale) variation in environment (herb cover, litter depth, air temperature and irradiance) and its relationship with seedling emergence, mortality, and recruitment of the perennial herb Helleborus foetidus (Ranunculaceae) during 2 years. We conducted the study in three distant localities in south-eastern Iberian Peninsula (defining the regional scale). Local scale was defined in terms of microhabitat type based on three categories of vegetation cover (under evergreen shrubs or trees, under deciduous spiny-shrubs, and in open interspaces). Finally, fine-scale variation was defined in terms of 1 m² sampling plots. Our results showed regional, local and fine-scale variations in recruitment, with the major source of variation changing across scales. Seed input was important in determining differences in recruitment both at the largest (regional) and the finest scale. Environment had minor importance in shaping differences in recruitment at these two scales. At the local scale, variation between microhabitats was mainly related to differences in seedling survival through facilitative effects (alleviation of water and irradiance stresses) of shrub and tree cover, although such facilitative effect was not consistent in all localities or years. Our study thus points to a similarity in the spatial patterns of recruitment (and in the environmental factors shaping such patterns) between Mediterranean woody plants and perennial herbs, but much more effort is needed to assess the generality of such similarity.     

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.     

Balancing risks? Responses and non-responses of mayfly larvae to fish and stonefly predators

Summary In a series of laboratory experiments we examined the hypothesis that larvae of stream mayflies would respond to the presence of two different types of predators in such a way as to minimize their risk of being consumed by each. Positioning of larvae (whether they frequent the top, sides, or bottom of stones) of Baetis tricaudatus and Ephemerella subvaria was altered by the presence of predaceous stoneflies (Agnetina capitata) with a larger proportion of the population occurring on the upper surfaces, where the probability of encountering the predator was lowest. The presence of a benthivorous fish (Cottus bairdi) had no significant effects on positioning of the mayfly larvae. Lack of fish effects may reflect an inability of the mayflies to detect or respond to sculpins, or alternately may indicate that sculpins do not normally present a important predation risk for these mayflies. Failure of mayfly prey to account for fish predators when responding to the presence of stoneflies appcars to explain facilitation previously observed between stoneflies and sculpins.     

Effects of artificial plant cover on plant colonization of a bare peat surface

Abstract. This paper describes the effect of artificial plant cover on plant colonization of a bare peat surface, resulting from peat harvesting. Plant species colonization was compared on plots supplied with plastic models simulating Vaccinium vitis-idaea plants and plots without this artificial cover. After two growing seasons, species composition and total biomass of the established plant cover were similar in the two plot types. However, the number of established seedlings in the plots with artificial cover was significantly higher than that in the plots without cover. Out of 13 species observed four differed significantly in their performance on the two plot types. Betula spec. had both higher seedling numbers and higher biomass on the test plots; Deschampsia cespitosa had a higher biomass, whereas the seedlings were too numerous to be counted; Salix phylicifolia had higher seedling numbers. On the contrary, the number of seedlings of Epilobium angustifoliwn was lower on plots with artificial cover. It is suggested that colonization by Betula, D. cespitosa and S. phylicifolia was facilitated mainly by the improved microclimatic and soil moisture conditions under the artificial plant cover. On the other hand, germination of E. angustifolium may be negatively influenced by the increased shade on the test plots.     

Intra-generic competition among Nothofagus in New Zealand's primary indigenous forests

Competitive interactions between New Zealand's four Nothofagus or southern beech species were analysed using an extensive dataset describing the composition of natural forests, supplemented by environmental estimates describing both climate and landform. Using multiple regression models of progressively increasing complexity, the analysis first accounted for variation in tree abundance attributable to both environment and regional-scale distributional disjunctions of likely historic origin. Intra-generic competition, expressed as variation in tree abundance dependent on the presence or absence of each congener, was then assessed by adding (1) simple terms to assess the magnitude of gross changes in abundance, and (2) interaction terms to assess variation in abundance along the dominant temperature gradient given different competitive contexts. Results indicate the presence of substantial intra-generic interactions, with simple interaction terms giving marginal increases in explained deviance equal to that explained by initial regressions using environment alone. Addition of interaction terms brought about smaller improvements in model fit, but confirm that variation in abundance along the dominant annual temperature gradient is strongly influenced by the competitive context provided by the remaining congeners. Such results are consistent with current understanding of the niche concept, and underline the difficulty inherent in using current species limits to predict likely changes in species distributions consequent on global warming.     

Spatial relations between shrubs and Prosopis glandulosa canopies

Abstract. Prosopis glandulosa, an arborescent legume, may act as a nurse plant that facilitates the establishment of other woody species. We hypothesized that attenuation of radiant energy and increased soil nutrients beneath P. glandulosa canopies facilitate establishment of subordinate shrubs and shrub cluster development. We determined the spatial distribution pattern of shrubs under P. glandulosa at three locations in southern Texas. Density of Celtis pallida, Zanthoxylum fagara, and total woody plants were comparable among the four cardinal directions at each location, which countered the prediction that shrub density would be greater on the north side of P. glandulosa canopies if attenuation of solar energy was a factor in cluster development. Total woody plant density increased with increasing P. glandulosa basal diameter, canopy radius, and height only at one location. Total woody plant density decreased with increasing total N in the upper 15 cm of soil at two of the three locations. Late in shrub cluster development, extraction of N from the soil and incorporation of N into plant tissue in dense shrub clusters may operate to inhibit further increases in subordinate shrub density.