Root biomass and rooting profile of naturally regenerated beech in mid-elevation Scots pine woodlands

The natural establishment of shade-tolerant forest species such as beech (Fagus sylvatica L.) occurs in naturally regenerated Scots pine (Pinus sylvestris L.) woodlands that develop on former pastures and cultivated lands. To examine possible effects of underground competition in beech establishment, we studied the root biomass and the rooting profile of 53 mixed Scots pine-natural beech woodlands in French mid-elevation volcanic areas. Stands were arranged along a maturation gradient. Roots were sampled using the root-auger technique (0 to 75 cm depth every 15 cm). In addition, 23 young beeches were uprooted to study the entire root system. Total beech fine-root biomass was closely correlated with most beech aerial characteristics, (e.g., height, diameter and girth), and correlated moderately with tree age. However, it correlated poorly with basic competition indices such as stand density and basal area. Conversely, competition indices including vertical dimensions of competing trees were correlated with the underground biomass, probably as a result of redundancy with beech height. The rooting profile (fine roots, < 5 mm) of beech and pine were quite similar, and did not change significantly along the stand maturation gradient. Beech has a heart-shaped root system while pine is more plate-like and dimorphic. Beech fine-root biomass progressively surpassed pine biomass throughout the soil layers, thus confirming that it is dynamic and competitive in mature mixed stands. The coexistence of the root systems of beech and pine in the same soil layers presumably results in strong underground competition.     

Interspecific competition in metapopulations

The assumptions and predictions of metapopulation models for competing species are discussed in relation to empirical studies of colonization and extinction in metapopulations. In three species of Daphnia in rockpools, interspecific competition increased local extinction rates, while no effects on colonization rates were detected. Distributional patterns were consistent with several predictions of the competition model; for example, the number of species on an island increased with the number of pools and the proportion of pools occupied by each species decreased with increasing species number. It is concluded that interspecific competition is important for the distributional dynamics of Daphnia species in rockpools, but the question whether the coexistence of these species depends on metapopulation dynamics is still unresolved. Other studies on the effects of interspecific competition on colonization and extinction rates are discussed.     

The role of climate change in the widespread mortality of holm oak in open woodlands of Southwestern Spain

Forest decline and increasing tree mortality are of global concern and the identification of the causes is necessary to develop preventive measures. Global warming is an emerging factor responsible for the increasing tree mortality in drought-prone ecosystems. In the southwestern Iberian Peninsula, Mediterranean holm oak open woodlands currently undergo large-scale population-level tree die-off. In this region, temperature and aridity have increased during recent decades, but the possible role of climate change in the current oak mortality has not been investigated.To assess the role of climate change in oak die-off in managed open woodlands in southwestern Spain, we analyzed climate change-related signals in century-long tree ring chronologies of dead holm oaks. We examined the high/low-frequency variability in growth and the relationship between growth and climate.Similar to other Mediterranean forests, growth was favored by precipitation from autumn of the year prior to ring formation to spring of the year of ring formation, whereas high temperatures during spring limited growth. Since the 1970s, the intensity of the high-frequency response to water availability increased simultaneously with temperature and aridity. The growth trends matched those of climatic changes. Growth suppressions occurred during droughts in the 1970s, 1980s and 1990s. Widespread stand-level, age-independent mortality occurred since 2005 and affected trees that cannot be considered old for the species standards.The close relationship between growth and climate indicate that climate change strongly controlled the growth patterns. This suggests that harsher climatic conditions, especially increased aridity, affected the tree performance and could have played a significant role in the mortality process. Climate change may have exacerbated or predisposed trees to the impact of other factors (e.g. intense management and pathogens). These observations could suggest a similar future increase in oak mortality which may occur in more northern oak open woodlands if aridity further increases.     

Effects of dominant tree species on insectivorous birds breeding in Mediterranean oak woodlands

Capsule Differences in Cork Oak Quercus suber and Holm Oak Quercus rotundifolia dominance had little influence on bird communities though bark-gleaners showed a foraging preference for Cork Oak.

Aims Examine the use of Cork and Holm Oak trees by insectivorous birds in Mediterranean oak woodlands.

Methods Point-counts were used to compare species abundance among Cork Oak-dominated, Holm Oak-dominated and mixed woodlands. Focal foraging observations were used to evaluate the use of Cork and Holm Oaks in the three habitats and to relate tree characteristics with the foraging time of foliage- and bark-gleaners.

Results Bird densities in the three habitats were not different for most foliage- and bark-gleaners. Tree preference index values and foraging time per tree showed no significant differences between tree species and foraging guilds, however bark-gleaners had positive index values for Cork Oak in the three habitats. The foraging time of foliage- and bark-gleaners on both tree species showed a positive relationship with characteristics associated with arthropod abundance.

Conclusion Cork and Holm Oak trees are equally preferred by foliage-gleaners but bark-gleaners moderately preferred Cork Oak. Characteristics regarding morphology, phenology and physiological condition of trees can be used to predict habitat quality for insectivorous forest birds in Mediterranean oak woodlands.     

Response of ants and spiders to prescribed fire in oak woodlands of California

Conservation managers of oak woodlands have been reintroducing fire both as an ecological process per se and to assist in restoring native plant communities. To increase our understanding of the impacts of reintroduced fire on ground-dwelling invertebrates we examined the response of ants and spiders to a late season (autumn) prescribed fire conducted in a blue oak (Quercus douglasii) woodland ecosystem in northern California. Twelve 100 m × 100 m plots were established, six plots received a burn treatment and the remaining six plots were unburned controls. Ants and spiders were sampled using pitfall traps left open continuously and collected approximately every 32 days. Sampling was conducted over a year, consisting of four pre-burn and nine post-burn collections. Abundance was analyzed using a repeated measures ANOVA, which showed seed-harvester ants decreased significantly in the two months following the fire. Total spider abundance also showed a significant decrease in two months, although this did not occur immediately after the burn. One spider hunting guild, the ‘diurnal ambush’ group (Thomisidae) remained suppressed for up to nine months. Correspondence analysis measures of ant species abundance with environmental and vegetation variables (percent rock, bare ground, plant species richness and height of herbaceous vegetation) were higher than expected by chance, which assists in explaining some of the responses. Findings from this study revealed that the reintroduction of autumn burns has modest and short-term effect on the invertebrates sampled, suggesting that late season fires are compatible with other conservation goals for oak woodland ecosystems.     

Interspecific competition among macroparasites in a density-dependent host population

 We analyze the dynamics of a community of macroparasite species that share the same host. Our work extends an earlier framework for a host species that would grow exponentially in the absence of parasitism, to one where an uninfected host population is regulated by factors other than parasites. The model consists of one differential equation for each parasite species and a single density-dependent nonlinear equation for the host. We assume that each parasite species has a negative binomial distribution within the host and there is zero covariance between the species (exploitation competition). New threshold conditions on model parameters for the coexistence and competitive exclusion of parasite species are derived via invadibility and stability analysis of corresponding equilibria. The main finding is that the community of parasite species coexisting at the stable equilibrium is obtained by ranking the species according t! o th e minimum host density H ^* above which a parasite species can grow when rare: the lower H ^* , the higher the competitive ability. We also show that ranking according to the basic reproduction number Q ₀ does not in general coincide with ranking according to H ^* . The second result is that the type of interaction between host and parasites is crucial in determining the competitive success of a parasite species, because frequency-dependent transmission of free-living stages enhances the invading ability of a parasite species while density-dependent transmission makes a parasite very sensitive to other competing species. Finally, we show that density dependence in the host population entails a simplification of the portrait of possible outcomes with respect to previous studies, because all the cases resulting in the exponential growth of host and parasite populations are eliminated.. Received: 24 June 1996 / Revised version: 28 April 1998     

Interspecific competition enhances nitrogen fixation in an actinorhizal shrub

In forest understory restoration, the establishment of reintroduced species may be strongly linked to their ability to compete for belowground resources. In this study, we provide isotopic and morphological evidence for competition-induced increases in nitrogen fixation by Morella cerifera (L.) Small (wax myrtle) when planted with Pinus palustris Mill (longleaf pine). Compared to a competition-free treatment, we found no significant differences in tissue N concentrations for M. cerifera. However, ¹⁵N enrichment in leaves, stems and roots, as well as whole-plant values for nitrogen derived from fertilizer were significantly lower when the plants were subject to interspecific competition from P. palustris. Plants in the competition treatment also allocated a significantly greater percentage of belowground biomass to root nodules than those in the competition-free treatment (0.65 vs. 0.41%). This strongly suggests that M. cerifera is capable of upregulating nitrogen fixation in response to interspecific competition. This may help explain why M. cerifera outperformed non-nitrogen-fixing species reintroduced on the same site.     

Regeneration traits are structuring phylogenetic diversity in cork oak (Quercus suber) woodlands

Question: What factors determine the deviations from the relationship between species richness (which considers species as independent entities) and phylogenetic diversity (PD) (which considers species relatedness)? What are the implications for community composition and phylogenetic structure? Location: Los Alcornocales Natural Park (36°03′–36°45′N and 5°20′–5°45′W), in southern Iberian Peninsula (Spain). Methods: We recorded all woody species and geographical features on 94 (20 m × 20 m) plots of cork oak woodlands. Disturbance information was obtained from the Park records; precipitation was estimated from local maps. PD was computed as the minimum total length of all the phylogenetic branches spanning the set of species on each site. Then, PD was regressed against species richness to test to what extent the unexplained variance in this relationship could be accounted for by environmental variables and disturbances, and against the representation of species with different regeneration strategies. Results: Species richness and PD are strongly related; however, the remaining variability can be explained by: (1) precipitation and disturbance, and (2) the proportion of seeder species. Thus, the PD both of areas with low precipitation and high disturbance, and of areas with a high representation of seeder species, is lower than what would be expected from the species richness. Conclusions: Regeneration traits are important in structuring plant community composition; specifically, they contribute to shaping biodiversity in Mediterranean ecosystems. Species richness tends to overestimate biodiversity in highly disturbed systems.     

Interspecific competition constrains local abundance in highly suitable areas

Sites with high environmental suitability for species’ occurrence, in terms of abiotic conditions, may hold populations with higher local abundances by increasing reproductive and survival rates and decreasing extinction rate. Interspecific competition, however, may affect this relationship. Here we tested the hypothesis that local abundance of the gray slender opossum Marmosops incanus is affected by the local richness of potential competitors and environmental suitability derived from ecological niche models (ENMs). We also discuss the ability of distinct modelling methods to predict species’ abundance. We compiled occurrence records and information about M. incanus’ relative abundance from museums and published articles. Environmental suitability was derived from five algorithms using seven environmental predictors. To assess our hypothesis, we chose the best statistical models among generalised linear models and quantile regressions, and then tested whether the effects of richness of competitors on local abundance are stronger under highly suitable conditions. We found that environmental suitability given by presence-only methods are positively related to the maximum abundance of M. incanus. That is, species’ local abundance is low when suitability is low but can be either low or high when suitability is high. The richness of competitors, in turn, explains the abundance variation within sites with high environmental suitability. We strongly recommend that the relationship between abundance and suitability must be carefully interpreted when using ENMs to predict species’ distribution because biotic interactions can be the main driver of local abundance within highly suitable environments.     

Interspecific competition alters leaf stoichiometry in 20 grassland species

The extensive use of traits in ecological studies over the last few decades to predict community functions has revealed that plant traits are plastic and respond to various environmental factors. These plant traits are assumed to predict how plants compete and capture resources. Variation in stoichiometric ratios both within and across species reflects resource capture dynamics under competition. However, the impact of local plant diversity on species‐specific stoichiometry remains poorly studied. Here, we analyze how spatial and temporal diversity in resource‐acquisition traits affects leaf elemental stoichiometry of plants (i.e. the result of resource capture) and how flexible this stoichiometry is depending on the functional composition of the surrounding community. Therefore, we assessed inter‐ and intraspecific variations of leaf carbon (C), nitrogen (N), and phosphorus (P) (and their ratios) of 20 grassland species in a large trait‐based plant diversity experiment located in Jena (Germany) by measuring leaf elemental concentrations at the species‐level along a gradient in plant trait dissimilarity. Our results show that plants showed large intra‐ and interspecific variation in leaf stoichiometry, which was only partly explained by the functional group identity (grass or herb) of the species. Elemental concentrations (N, P, but not C) decreased with plant species richness, and species tended to become more deviant from their monoculture stoichiometry with increasing trait dissimilarity in the community. These responses differed among species, some consistently increased or decreased in P and N concentrations; for other species, the negative or positive change in P and N concentrations increased with increasing trait difference between the target species and the remaining community. The strength of this relationship was significantly associated to the relative position of the species along trait gradients related to resource acquisition. Trait‐difference and trait‐diversity thus were important predictors of how species’ resource capture changed in competitive neighbourhoods.     

Interspecific competition in small rodents: from populations to individuals

The role of interspecific competition in shaping animal and plant communities has formed one of the major issues in ecology for decades. Small mammals, mainly rodents, have been among the model systems used for research on interspecific competition. Most studies within small mammal systems in the past have examined effects of competition on population attributes such as on population size, habitat use, or population dynamics. Population-level responses are the cumulative effects of individual responses, however, the influence of competition on individual life-history traits has rarely been studied. Research on life-histories may bridge gaps between population biology and effects of competition on individual behaviour. In this paper, we review recent research approaches to interspecific competition in rodents based on census data and species assemblages, that use regression analysis, time series analysis, removal and exclusion experiments, and showcase our own experimental research on the effects of interspecific competition on individual life-history traits in boreal voles.     

Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Question: Insufficient tree regeneration threatens the long‐term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance‐related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.     

Interspecific competition in perennial sedentary organisms: An individual-based model

We investigated a mathematical model of the dynamics of the ecological system consisting of two competing perennial species, each of which leads a sedentary life. It is an individual-based model, in which the growth of each individual is described. The rate of this growth is weakened by competition from neighboring individuals. The strength of the competitors' influence depends on their size and distance to them. The conditions, in which the competitive exclusion of one of the competitors and the coexistence of both competitors take place are provided. The influence of the parameters responsible for the strength of competition, the degree of competitive asymmetry, and consideration of the importance of specific elements of the spatial structure of this ecological system on the results of the competition were analyzed. Both species co-exist when they are equal competitors. Permanent coexistence is possible only when interspecific competition is weaker than intraspecific. When interspecific competition is stronger, the coexistence of equal interspecific competitors is random. Both species have equal probability of extinction. If species are not equal competitors, the stronger one wins. This result can be modified by different strengths of intraspecific competition. The weaker interspecific competitor can permanently coexist with stronger one, when its individuals suffer stronger intraspecific competition.     

沙区植物种的一类种间竞争模型及应用

以沙坡头人工固沙植物油蒿和柠条为研究对象,针对水分是沙坡头植物生r长的限制因子的现实,研究了在水分条件限制下两种植物的种间关系,建立了混播油蒿、柠条种间竞争模型,验证了油蒿、柠条既竞争又稳定共存的关系,其目的为充分有效利用水资源、稳定人工生态系统提供科学依据。     

小麦/大豆间作中作物种间的竞争作用和促进作用

春小麦／春大豆间作是西北一熟制灌区广泛采用的高产种植形式．本文采用田间小区和微区根系分隔试验研究了这种种植形式作物种间的竞争作用和促进作用．结果表明,小麦／大豆间作具有明显的间作优势．土地当量比为１．２３～１．２６．小麦为优势种,竞争力强于大豆,具有明显的间作边行优势．小麦边行优势的１／３贡献来自于地下部．小麦收获后,大豆生长具有恢复作用,认为这种恢复作用是间作优势的机制之一．间作相对于单作两种作物的收获指数均有显著提高．收获指数的种间促进作用是间作优势的另一机制．     

The influence of vegetation on bird distribution in dry forests and oak woodlands of western Mexico

The bird species distribution along a dry forest-oak woodland vegetation gradient was studied in autumn and spring in two consecutive years. Intra-seasonal comparisons showed that bird species had similar distributions in each of the two years. Inter-seasonal changes were mainly due to compositional differences even though resident species generally used similar habitats in both seasons. Ordination analyses, based on the first year bird species abundances, showed a clearly segregated distribution between forest and woodland birds. Within these two vegetation types, the distribution tended to be more individualistic. Nevertheless further habitats could be identified according to groups of birds having similar distributions. These habitats did not correspond to the plant associations which resulted from a previous classification of the vegetation. Observations of the plant use by the birds during the study period showed that, in most cases, the plant variables associated with ordination analyses are unlikely to be very important for the bird species life cycles.     

Testing the effect of restoration‐focused silviculture on oak regeneration and groundlayer plant communities in urban–exurban oak woodlands

Throughout their global range, oak‐dominated ecosystems have undergone state changes in stand structure and composition. Land managers face an especially acute challenge in restoring oak ecosystems and promoting oak regeneration in urban–exurban areas, where high‐intensity silvicultural treatments are often not feasible. To investigate low‐intensity management alternatives which could be widely applied in urban–exurban forests, a large‐scale adaptive management experiment was implemented in Lake County, IL, in 2012. Five canopy manipulation treatments of varying intensity, timing, and spatial aggregation were replicated across three study areas and oak seedlings were under‐planted into treatment units following management. Responses of understory light environment, shrub and groundlayer plant communities, and survival and growth of underplanted oak seedlings were evaluated. Understory light availability, canopy openness, total groundlayer plant cover, and groundlayer species diversity all differed among treatments. However, although understory light availability was significantly increased by canopy manipulation, groundlayer communities and oak seedling survival and growth did not differ among treatments. High overall seedling survival rates suggest current conditions are amenable to oak regeneration, but long‐term monitoring will be needed to assess the potential for seedlings to transition to the sapling and canopy layers. Early results demonstrate that canopy‐focused silvicultural treatments can affect the understory light environment and, to some degree, groundlayer plant communities. However, underplanting of oak seedlings paired with subcanopy thinning may be sufficient to restore an oak seedling layer, and (when necessary or preferred) canopy manipulation could potentially be deferred until later in the restoration timeline to promote oak recruitment.