Postdispersal seed predation and seed viability in forest soils: implications for the regeneration of tree species in Ethiopian church forests

Almost all dry Afromontane forests of Northern Ethiopia have been converted to agricultural, grazing or scrub lands except for small fragments left around churches (‘Church forests’). Species regeneration in these forests is limited. We investigated (i) how intense postdispersal seed predation was in church forest, and if this seed predation varied with species and/or habitat, and (ii) for how long tree seeds maintained their viability while buried in forest soil. In the seed predation experiment, we monitored seeds of six tree species in four habitats for a period of 14 weeks (the peak seeding season). In the seed viability experiment, we assessed seed viability of five species in four habitats after being buried 6, 12, or 18 months. Ninety‐two percent of the tree seeds were predated within 3.5 months. Predation was mainly dependent on species whereas habitat had a weaker effect. Seed viability decreased sharply with burial time in soil for all species except for Juniperus. To minimize seed availability limitation for regeneration of such species in the forest, the standing vegetation needs to be persistently managed and conserved for a continuous seed rain supply. Additional seed sowing, and seed and seedling protection (by e.g. animal exclosures) may increase successful regeneration of important species in these forests.     

An ecological paradox: high species diversity and low position of the upper forest line in the Andean Depression

Systematic investigations of the upper forest line (UFL) primarily concentrate on mid and high latitudes of the Northern Hemisphere, whereas studies of Neotropical UFLs are still fragmentary. This article outlines the extraordinary high tree diversity at the UFL within the Andean Depression and unravels the links between the comparatively low position of the local UFL, high tree‐species diversity, and climate. On the basis of Gentry′s rapid inventory methodology for the tropics, vegetation sampling was conducted at 12 UFL sites, and local climate (temperature, wind, precipitation, and soil moisture) was investigated at six sites. Monotypic forests dominated by Polylepis were only found at the higher located margins of the Andean Depression while the lower situated core areas were characterized by a species‐rich forest, which lacked the elsewhere dominant tree‐species Polylepis. In total, a remarkably high tree‐species number of 255 tree species of 40 different plant families was found. Beta‐diversity was also high with more than two complete species turnovers. A non‐linear relationship between the floristic similarity of the investigated study sites and elevation was detected. Temperatures at the investigated study sites clearly exceeded 5.5°C, the postulated threshold value for the upper tree growth limit in the tropics. Instead, quasi‐permanent trade winds, high precipitation amounts, and high soil water contents affect the local position of the UFL in a negative way. Interestingly, most of the above‐mentioned factors are also contributing to the high species richness. The result is a combination of a clearly marked upper forest line depression combined with an extraordinary forest line complexity, which was an almost unknown paradox.     

Structural and Floristic Heterogeneity in a 30-Year-Old Costa Rican Rain Forest Restored on Pasture Through Natural Secondary Succession

Relatively little information exists on neotropical secondary rain forests that have progressed beyond the pioneer stages of succession, or on the potential of natural regeneration to restore forest on large areas. We determined the structural and floristic characteristics (10 cm dbh) of a 30‐year‐old secondary forest developing on a 32.5 ha pasture on hilly terrain, abandoned after use of moderate intensity. Ten 0.24 ha sample plots covered the range of site conditions. The forest was dominated by long‐lived pioneer tree species; overall, the majority of species (70%) was vertebrate dispersed but the majority of individuals (52%) was of wind‐dispersed species. Tree species, including the dominants, were a mixture of those present in old‐growth and adventives colonizing from agricultural land. The forest was very heterogeneous. Vochysia ferruginea‐dominated stands characterized slopes with soils of high exchangeable acidity, while the adventive Cordia alliodora dominated sites with gentler topography and soils of lower acidity. Structural differences between the two forest types were slight, but Cordia forest had significantly greater species diversity and absolute and relative abundances of vertebrate‐dispersed tree species than Vochysia forest, which had significantly greater absolute and relative abundances of wind‐dispersed tree species. These latter differences between forest types, as well as the wide structural variation of the forest as a whole, were probably largely due to spatial and temporal variation in seed rains, some of it linked to the characteristics of the dominant species. Rain forest restoration on large pastures may depend greatly on wind dispersal and adventive tree species, and techniques for silvicultural diagnosis must be developed as a basis for the management of heterogeneous successional stands. Studies of early colonization of pastures should be expanded to focus on the causes of heterogeneity in older forests.     

The species richness–productivity relationship in the herb layer of European deciduous forests

Aim In contrast to non‐forest vegetation, the species richness–productivity (SR‐P) relationship in forests still remains insufficiently explored. Several studies have focused on the diversity of the tree layer, but the species richness of temperate deciduous forests is mainly determined by their species‐rich herb layer. The factors controlling herb‐layer productivity may differ from those affecting tree layers or open herbaceous vegetation, and thus the SR‐P relationship and its underlying processes may differ. However, the few relevant studies have reported controversial results. Here we explore the SR‐P relationship in the forest herb layer across different areas from oceanic to continental Europe, and put the effect of habitat productivity on species richness into context with other key factors, namely soil pH and light availability. Location North‐western Germany, Czech Republic, Slovakia and southern Urals (Russia). Methods We measured herb‐layer species richness and biomass, soil pH and tree‐layer cover in 156 vegetation plots of 100 m² in deciduous forests. We analysed the SR‐P relationship and the relative importance of environmental variables using regression models for particular areas and separate forest types. Results We found a consistent monotonic increase in the herb‐layer species richness with productivity across all study areas and all forest types. Soil pH and light availability also affected species richness, but their relative importance differed among areas. Main conclusions We suggest that the monotonically increasing SR‐P relationship in the forest herb layer results from the fact that herb‐layer productivity is limited by canopy shading; competition within the herb layer is therefore not strong enough to exclude many species. This differs fundamentally from open herbaceous vegetation, which is not subject to such productivity limits and consequently exhibits a unimodal SR‐P relationship. We present a conceptual model that might explain the differences in the SR‐P relationship between the forest herb layer and open herbaceous vegetation.     

Dominant forest tree mycorrhizal type mediates understory plant invasions

Forest mycorrhizal type mediates nutrient dynamics, which in turn can influence forest community structure and processes. Using forest inventory data, we explored how dominant forest tree mycorrhizal type affects understory plant invasions with consideration of forest structure and soil properties. We found that arbuscular mycorrhizal (AM) dominant forests, which are characterised by thin forest floors and low soil C : N ratio, were invaded to a greater extent by non‐native invasive species than ectomycorrhizal (ECM) dominant forests. Understory native species cover and richness had no strong associations with AM tree dominance. We also found no difference in the mycorrhizal type composition of understory invaders between AM and ECM dominant forests. Our results indicate that dominant forest tree mycorrhizal type is closely linked with understory invasions. The increased invader abundance in AM dominant forests can further facilitate nutrient cycling, leading to the alteration of ecosystem structure and functions.     

Rehabilitation of Degraded Areas of Central Amazonia Using Direct Sowing of Forest Tree Seeds

Deforestation in the Amazon Basin is still increasing, and the rehabilitation of these lands continues to be a challenge. Autoecological studies of most Amazonian species are rare, and efficient techniques for restoration of forested habitats have yet to be developed. The aim of this study was to test direct sowing as a rehabilitation technique for sites with different degrees of disturbance: bare soil, pasture, and secondary and mature forests in Central Amazonia, Brazil. At each site, we sowed seeds of 11 native tree species. Throughout the following year we evaluated germination and seedling survival. The germination differed according to the study site and species. Seedling survival in degraded sites was higher than in other areas. After 1 year in the bare soil site, 33% of the sown seeds of eight species developed seedlings; in the pasture the establishment was 23%, in secondary forest 15%, and in mature forest 12% of only four species. The only widespread survivor with more than 45% emergence in all perturbed sites was Caryocar villosum. No pioneer seedlings remained after 1 year. There was a positive correlation between seed size and survival. Large‐seeded non‐pioneer species seem to be more suitable for direct sowing than small‐seeded species. We recommend a combination of direct sowing and planting of seedlings as an appropriate means to accelerate the rehabilitation of degraded areas in Central Amazonia.     

Stability in the patterns of long‐term development and growth of the Canadian spruce–moss forest

Aim The spruce–moss forest is the main forest ecosystem of the North American boreal forest. We used stand structure and fire data to examine the long‐term development and growth of the spruce–moss ecosystem. We evaluate the stability of the forest with time and the conditions needed for the continuing regeneration, growth and re‐establishment of black spruce (Picea mariana) trees. Location The study area occurs in Québec, Canada, and extends from 70°00′ to 72°00′ W and 47°30′ to 56°00′ N. Methods A spatial inventory of spruce–moss forest stands was performed along 34 transects. Nineteen spruce–moss forests were selected. A 500 m² quadrat at each site was used for radiocarbon and tree‐ring dating of time since last fire (TSLF). Size structure and tree regeneration in each stand were described based on diameter distribution of the dominant and co‐dominant tree species [black spruce and balsam fir (Abies balsamea)]. Results The TSLF of the studied forests ranges from 118 to 4870 cal. yr bp . Forests < 325 cal. yr bp are dominated by trees of the first post‐fire cohort and are not yet at equilibrium, whereas older forests show a reverse‐J diameter distribution typical of mature, old‐growth stands. The younger forests display faster height and radial growth‐rate patterns than the older forests, due to factors associated with long‐term forest development. Each of the stands examined established after severe fires that consumed all the soil organic material. Main conclusions Spruce–moss forests are able to self‐regenerate after fires that consume the organic layer, thus allowing seed regeneration at the soil surface. In the absence of fire the forests can remain in an equilibrium state. Once the forests mature, tree productivity eventually levels off and becomes stable. Further proof of the enduring stability of these forests, in between fire periods, lies in the ages of the stands. Stands with a TSLF of 325–4870 cal. yr bp all exhibited the same stand structure, tree growth rates and species characteristics. In the absence of fire, the spruce–moss forests are able to maintain themselves for thousands of years with no apparent degradation or change in forest type.     

Rainfall seasonality and drought performance shape the distribution of tropical tree species in Ghana

Tree species distribution in lowland tropical forests is strongly associated with rainfall amount and distribution. Not only plant water availability, but also irradiance, soil fertility, and pest pressure covary along rainfall gradients. To assess the role of water availability in shaping species distribution, we carried out a reciprocal transplanting experiment in gaps in a dry and a wet forest site in Ghana, using 2,670 seedlings of 23 tree species belonging to three contrasting rainfall distributions groups (dry species, ubiquitous species, and wet species). We evaluated seasonal patterns in climatic conditions, seedling physiology and performance (survival and growth) over a 2‐year period and related seedling performance to species distribution along Ghana's rainfall gradient. The dry forest site had, compared to the wet forest, higher irradiance, and soil nutrient availability and experienced stronger atmospheric drought (2.0 vs. 0.6 kPa vapor pressure deficit) and reduced soil water potential (?5.0 vs. ?0.6 MPa soil water potential) during the dry season. In both forests, dry species showed significantly higher stomatal conductance and lower leaf water potential, than wet species, and in the dry forest, dry species also realized higher drought survival and growth rate than wet species. Dry species are therefore more drought tolerant, and unlike the wet forest species, they achieve a home advantage. Species drought performance in the dry forest relative to the wet forest significantly predicted species position on the rainfall gradient in Ghana, indicating that the ability to grow and survive better in dry forests and during dry seasons may allow species to occur in low rainfall areas. Drought is therefore an important environmental filter that influences forest composition and dynamics. Currently, many tropical forests experience increase in frequency and intensity of droughts, and our results suggest that this may lead to reduction in tree productivity and shifts in species distribution.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Roles of non‐native species in large‐scale regeneration of moist tropical forests on anthropogenic grassland

This paper presents a new synthesis of the role of native and non‐native species in diverse pathways and processes that influence forest regeneration on anthropogenic grassland in the moist tropics. Because of altered species composition, abiotic conditions and landscape habitat mosaics, together with human interventions, these successional pathways differ from those seen in pre‐clearing forests. However, representation of different functional life forms of plant (tree, vine, grass, herb and fern) and animal (frugivorous seed disperser, granivorous seed predator, seedling herbivore and carnivore) shows consistent global variation among areas of pasture, intact forest, and post‐grassland regrowth. Biotic webs of interaction involve complex indirect influences and feedbacks, which can account for wide observed variation in regeneration trajectories over time. Important processes include: limitation of tree establishment by dense grasses; recruitment and growth of pioneer pasture trees (shading grasses and facilitating bird‐assisted seed dispersal); and smothering of trees by vines. In these interactions, species’ functional roles are more important than their biogeographic origins. Case studies in eastern Australia show native rain forest plant species diversity in all life forms increasing over time when pioneer trees are non‐native (e.g., Cinnamomum camphora, Solanum mauritianum), concurrent with decreased grass and fern cover and increased abundance of trees and vine tangles. The global literature shows both native and non‐native species facilitating and inhibiting regeneration. However conservation goals are often targeted at removing non‐native species. Achieving large‐scale tropical forest restoration will require increased recognition of their multiple roles, and compromises about allocating resources to their removal.     

Protecting a single endangered species and meeting multiple conservation goals: an approach with Guaiacum sanctum in Yucatan Peninsula,Mexico

Aim New protected areas should consider safeguarding high conservation value sites based on multiple criteria and not just the presence of a single endangered or charismatic species. However, the extent to which complementary criteria coincide is usually unknown. We use the case of Guaiacum sanctum (Zygopyllaceae), an endangered timber tree species, to explore whether the protection of forests where this species is most abundant would meet other complementary conservation goals, such as capturing regional plant biodiversity, protecting other threatened/endemic species or safeguarding ecosystem services. Location Yucatan Peninsula, southern Mexico. Methods We conducted an analysis of the structure, composition and diversity of tree communities (including stems ≥5 cm dbh) at eight G. sanctum forest sites. We identified endemic and threatened tree species and quantified above‐ground tree biomass and carbon storage in these G. sanctum forests. Results Guaiacum sanctum forests contain 35–59 tree species on plots as small as 1000 m². The species composition of tree communities changed rapidly (high β‐diversity) across soil boundaries and rainfall regimes. Twenty‐one endemic and eight threatened tree species were recorded in our inventories. Individuals of G. sanctum represented up to 55% of the above‐ground carbon for trees ≥5 cm dbh. The high basal area of G. sanctum forests plus the high wood density, abundance, large size and longevity (more than 500 years) of G. sanctum and other tree species enhance the potential importance of these forests for carbon storage. Main conclusions A conservation strategy focused on protecting important populations of G. sanctum in the Yucatan Peninsula would have significant co‐benefits for conservation of regional tree species biodiversity and provision of critical ecosystem services. Our study illustrates a multiple criteria approach useful for the selection of areas with high conservation value on the basis of endemic, threatened species, species richness and ecosystem services.     

High value of short rotation coppice plantations for phytodiversity in rural landscapes

The demand for wood from short rotation coppice (SRC) plantations as a renewable energy source is currently increasing and could affect biodiversity in agricultural areas. The objective was to evaluate the contribution of SRC plantations to phytodiversity in agricultural landscapes assessed as species richness, species–area relationships, Shannon indices, detrended correspondence analysis on species composition, Sørensen similarities, habitat preference proportions, and species proportions found in only one land use. Vegetation surveys were conducted on 12 willow (Salix spp.) and three poplar (Populus spp.) coppice sites as well as on surrounding arable lands, grasslands and forests in central Sweden and northern Germany. SRC plantations were richer in plant species (mean: 30 species per 100 m²) than arable land (10), coniferous forests (13) and mixed forests in Germany (12). Comparing SRC plantations with other land uses, we found lowest similarities in species composition with arable lands, coniferous forests and German mixed forests and highest similarities with marginal grassland strips, grasslands and Swedish mixed forests. Similarity depended on the SRC tree cover: at increased tree cover, SRC plantations became less similar to grasslands but more similar to forests. The SRC plantations were composed of a mixture of grassland (33%), ruderal (24%) and woodland (15%) species. Species abundance in SRC plantations was more heterogeneous than in arable lands. We conclude that SRC plantations form novel habitats leading to different plant species composition compared to conventional land uses. Their landscape‐scale value for phytodiversity changes depending on harvest cycles and over time. As a structural landscape element, SRC plantations contribute positively to phytodiversity in rural areas, especially in land use mosaics where these plantations are admixed to other land uses with dissimilar plant species composition such as arable land, coniferous forest and, at the German sites, also mixed forest.     

Distance‐dependent effects of soil‐derived biota on seedling survival of the tropical tree legume Ormosia semicastrata

Question: How do seed germination and subsequent seedling survival of O. semicastrata (Hance forma litchiifolia How) vary with respect to distance from parent trees and conspecific density in different types of tropical forest? Are there effects of soil biota on O. semicastrata that systematically depend on distance from parent trees and conspecific density? Do soil pathogens differently affect survival of O. semicastrata in different types of tropical forest? Location: Tropical lowland rain forest and tropical montane rain forest in Jianfengling National Nature Reserve, Hainan Island, China. Methods: Individual adult O. semicastrata trees were selected in lowland rain forest and montane rain forest. Soil was collected at a distance of 0‐5 m or 15‐20 m from the parent tree. Soil samples from each distance were combined into a bulk sample. Half of the soil sample was sterilized by autoclaving. Surface‐sterilized seeds were then added to the soil material in shade‐houses at both forests. Results: Germination of O. semicastrata seeds at low‐ or high‐seed density was barely affected by the sterilization procedure. In both forests, seedlings grown in non‐sterilized soil collected close to parent trees had significantly higher mortality compared to those in sterilized soil. In contrast, seedling survival with soil collected far from parent trees was not affected by the soil sterilization procedure. Conclusions: Host‐specific pathogens concentrated in the soil around parent trees may regulate community structure of tropical trees at the stage of seedling development.     

川西亚高山三种天然次生林土壤微生物多样性变化特征

开展川西亚高山相似土壤母质背景下天然次生林土壤微生物群落结构及其多样性探究,可加深次生林更新过程中土壤微生物群落结构变化的认知。选取川西米亚罗林区20世纪60年代采伐后经自然更新恢复形成的3种天然次生林（槭-桦阔叶林,ABB;桦-槭-冷杉针阔混交林,BAA;岷江冷杉林,AFF）,分析林下表层（0-20 cm）土壤微生物群落结构变化及其影响因素,结果显示：（1）3种林型土壤细菌Chao1和Shannon指数均极显著高于真菌,但仅真菌群落的Shannon指数差异显著,表现为BAA > ABB > AFF;（2）细菌群落优势门主要为变形杆菌门、酸杆菌门、疣微菌门、拟杆菌门、绿弯菌门,相对丰度占比超过82%;真菌群落则为子囊菌门和担子菌门,占比超过85%,AFF担子菌门相对丰度最高而子囊菌门最低。（3） RDA分析显示,土壤pH和乔木物种多样性（Shannon指数）是影响微生物群落结构变化的主导因子;土壤养分元素对细菌群落影响不显著,真菌群落主要受TN、TP含量显著影响。总体上,林型间乔木层物种多样性、土壤酸碱度及其氮磷含量是导致微生物群落结构变化的关键因素。     

Bryophyte Species Diversity in Secondary Forests Dominated by the Introduced Species Spathodea campanulata Beauv. in Puerto Rico

The introduced tree species Spathodea campanulata (Bignoniaceae) forms novel forests in Puerto Rico, these having emerged after the abandonment of fields in the mid‐20th century and resulting in forests with a new species composition. We assessed bryophyte species richness in these novel forests and sought correlations with geological substrate, past land use, forest edge and patch area, forest structure, elevation, microhabitat diversity, tree species richness, and microclimatic conditions. Transects were established (edge and forest interior) in nine moist forest patches dominated by Spathodea in north‐central Puerto Rico. These Spathodea forest patches ranged from 0.6 to 9 ha. ANOVA, Chi‐square, correlation, and cluster analyses were used in data analyses. We found 57 bryophyte species. There was a significant difference in bryophyte richness among patches. Those on karst exhibited highest bryophyte richness due to microhabitat diversity, past land use, and shorter hydroperiods. Alluvial sites scored lowest in bryophyte species richness, and forest structure was important for bryophyte communities on these sites. Significant differences in temperature, relative humidity, and light intensity were observed between edge and forest interior. These appeared important for establishing bryophyte species cover but not richness and composition. Microhabitat diversity, patch area, and forest age were more related to bryophyte species richness than elevation, exposed edge, and tree species richness, regardless of geologic substrate. Collectively, Spathodea patches were similar to mature forests on the Island with respect to bryophyte species richness and composition. Novel Spathodea forests have conservation value due to their habitat suitability for bryophyte communities.     

Landscape host abundance and configuration regulate periodic outbreak behavior in spruce budworm Choristoneura fumiferana

Landscape‐level forest management has long been hypothesized to affect forest insect outbreak dynamics, but empirical evidence remains elusive. We hypothesized that the combination of increased hardwood relative to host tree species, prevalence of younger forests, and fragmentation of those forests due to forest harvesting legacies would reduce outbreak intensity, increase outbreak frequency, and decrease spatial synchrony in spruce budworm Choristoneura fumiferana outbreaks. We investigated these hypotheses using tree ring samples collected across 51 sites pooled into 16 subareas distributed across a large ecoregion spanning the international border between Ontario (Canada), and Minnesota (USA). This ecoregion contains contrasting land management zones with clear differences in forest landscape structure (i.e. forest composition and spatial configuration) while minimizing the confounding influence of climate. Cluster analyses of the 76‐yr time‐series generally grouped by subareas found within the same land management zone. Spatial nonparametric covariance analysis indicated that the highest and lowest degree of spatial synchrony of spruce budworm outbreaks were found within unmanaged wilderness and lands managed at fine spatial scales in Minnesota, respectively. Using multivariate analysis, we also found that forest composition, configuration, and climate together accounted for a total of 40% of the variance in outbreak chronologies, with a high level of shared variance between composition and configuration (13%) and between composition and climate (9%). At the scale of our study, climate on its own did not explain any of the spatial variation in outbreaks. Outbreaks were of higher frequency, lower intensity, and less spatially synchronized in more fragmented, younger forests with a lower proportion of host species, with opposing outbreak characteristics observed in regions characterised by older forests with more concentrated host species. Our study is the first quantitative evaluation of the long‐standing ‘silvicultural hypothesis’ of spruce budworm management specifically conducted at a spatio‐temporal scale for which it was intended.     

You'd better walk alone: Changes in forest composition affect pollination efficiency and pre‐dispersal cone damage in Iberian Juniperus thurifera forests

• Changes in land‐use patterns are a major driver of global environmental change. Cessation of traditional land‐use practices has led to forest expansion and shifts in forest composition. Consequently, former monospecific forests maintained by traditional management are progressing towards mixed forests. However, knowledge is scarce on how the presence of other tree species will affect reproduction of formerly dominant species. We explored this question in the wind‐pollinated tree Juniperus thurifera. We hypothesised that the presence of heterospecific trees would have a negative effect on cone production and on the proportion of cones attacked by specialised predators.
• We assessed the relative importance of forest composition on cone production, seed development and pre‐dispersal cone damage on nine paired pure and mixed J. thurifera forests in three regions across the Iberian Peninsula. The effects of forest composition on crop size, cone and seed characteristics, as well as damage by pre‐dispersal arthropods were tested using mixed models.
• Cone production was lower and seed abortion higher in mixed forests, suggesting higher pollination failure. In contrast, cone damage by arthropods was higher in pure forests, supporting the hypothesis that presence of non‐host plants reduces damage rates. However, the response of each arthropod to forest composition was species‐specific and the relative rates of cone damage varied depending on individual tree crops.
• Larger crop sizes in pure forests compensated for the higher cone damage rates, leading to a higher net production of sound seeds compared to mixed forests. This study indicates that ongoing changes in forest composition after land abandonment may impact tree reproduction.

     

Tree invasion in managed tropical forests facilitates endemic species

Aim To determine whether different abundances of introduced species of Cinchona (Rubiaceae) affect species composition and facilitate species richness in managed tropical forests, to test whether any facilitative effects on understorey species depend on forest type, and to investigate whether facilitative effects can be attributed to the ‘substitutive facilitation model’. Location Makawao Forest Reserve on Maui, Hawai’i, USA. Methods Cinchona species (Cinchona pubescens and Cinchona calisaya) were mapped within various forest types. In three forest types (ageing Eucalyptus and Pinus plantations, and near‐natural Acacia koa forests), we analysed environmental parameters (e.g. canopy cover, litter cover, pH value and soil depth) and the species composition of Cinchona‐invaded and non‐invaded plots; data were compared based on Cinchona cover and forest types. Habitat modelling for several endemic species and tree ferns was carried out to test whether Cinchona cover is an important variable for the probability of occurrence of these endemics. Results Cinchona species have naturalized mainly in Eucalyptus and Pinus plantations and Acacia koa forests and here add an additional shrub layer. In contrast to other studies, we revealed facilitative effects of Cinchona on native species within all forest types. Species richness is about 20% higher in invaded plots than in non‐invaded plots, and these show a nearly 50% higher proportion of endemic species, including tree ferns. The proportion of endemics even increases with increasing Cinchona cover. For several endemics, Cinchona is found to be an important variable for the probability of occurrence, and the removal of Cinchona cover as an explanatory variable lowers the model fit. In addition to Cinchona, variables delineating vegetation structure and light availability have a strong effect on the model fit. Main conclusions In the structurally simplified Hawaiian forests studied, Cinchona facilitated endemic species in accordance with the ‘substitutive facilitation model’. This contrasts with the results of an earlier study in the naturally treeless Galápagos highlands, which revealed a sharp decrease in the abundance of endemics under Cinchona canopy. These results illustrate that, through the same structural change (addition of a vegetation layer), an invasive species may exert divergent effects across different ecosystem types. The facilitation of endemic understorey species by invasive tree species in managed forests leads to a dilemma in conservation but also to new perspectives for ecosystem restoration.     

Oak trees and soil interactions in Mediterranean forests: a positive feedback model

Questions: What is the spectrum of variability of chemical elements in a Mediterranean forest ecosystem across the different compartments? Do co‐existing tree species with different leaf chemical composition and nutrient cycling distinctly modify soil conditions? Could these species‐specific, tree‐generated soil changes create a potential positive feedback by affecting long‐term species distribution? Location: Mixed oak forests of southern Spain, Los Alcornocales Natural Park. Methods: We sampled and chemically analysed five different ecosystem components: leaves, leaf fall, litter and superficial (0–25 cm) and sub‐superficial (25–50 cm) soil beneath the canopies of evergreen Quercus suber and deciduous Q. canariensis trees. We used multiple co‐inertia analysis (MCoA) to conjointly analyse the patterns of variability and covariation of eight macro‐ and micronutrients determined in each of the sampled ecological materials. We implemented a path analysis to investigate alternative causal models of relationships among the chemical properties of the different ecosystem components. Results: Variability in the concentration of chemical elements was related to the nature of their biogeochemical cycles. However, the rank of element concentration was consistent across ecosystem components. Analysis of co‐inertia (MCoA) revealed that there was a common underlying multivariate pattern of nutrient enrichment in the ecosystem, which supported the hypothesis of a separation in biogeochemical niches between the two co‐existing oak species, with Q. canariensis having richer plant tissues and more fertile soil directly under each tree than Q. suber. The feasibility of a potential tree–soil positive feedback model was the only statistically validated among several alternative (non‐feedback) models tested. Conclusions: In the studied Mediterranean forests, oak species distinctly modify soil fertility conditions through different nutrient return pathways. Further investigation is needed to address whether these tree‐generated soil changes could affect seedling establishment and ultimately influence species distribution.     

Livestock grazing and forest structure regulate the assembly of ecological clusters within plant networks in eastern Australia

Questions

How do changes in grazing intensity by different herbivores and differences in forest structure affect the assembly of ecological clusters within plant ecological networks in dryland plant communities?

Location

Eastern Australia across an area of 0.4 million km².

Methods

We used correlation network analysis and structural equation modelling to examine how changes in grazing intensity, by different herbivores, and differences in forest structure (tree canopy cover, basal area and density) and soil fertility influenced the assembly of ecological clusters of plant communities (i.e. relative abundance of ecological clusters formed by co‐occurring plant species within an ecological network) in three forested communities from eastern Australia.

Results

Livestock grazing and forest structure regulated the relative abundance of ecological clusters within plant networks, but their effects on these plant assemblies were highly dependent on the ecological cluster and forest community type, with no single winner or loser across forest types, conditions or grazing intensities. Thus, the relative abundance of some ecological clusters increased under grazing while others declined, a response that was maintained across different forest structures. The relative importance of grazing, forest structure and soil fertility varied across forest community type. The two eucalypt communities exhibited mixed effects of grazing and forest structure (Eucalyptus largiflorens ) or forest structure only (Eucalyptus camaldulensis ). In the third (Callitris glaucophylla ) community, grazing played a larger role in controlling the plant community assembly. Soil fertility (soil C and P) effects were of a similar magnitude to grazing and forest structure, but the effects differed among clusters.

Conclusions

Livestock grazing and forest structure regulated the relative abundance of ecological clusters within networks of plant communities in forests in eastern Australia. Our study uses a novel approach of ecological clusters to show that differences in grazing and forest structure will always disadvantage some plant ecological clusters. Furthermore, changes in one cluster will ultimately affect other clusters. Any changes in management therefore will have varied effects on different ecological plant clusters.