Ignacio Manuel Pérez-Ramos

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Mast seeding and flowering in Mediterranean oak woodlands under increasing drought: results from a long-term dataset and from a rainfall exclusion experiment

Mast seeding, the synchronous, highly variable seed production among years, is very common in tree species, but there is no consensus about its main causes and the main environmental factors affecting it. In this study, we first analyze a long-term data set on reproductive and vegetative growth of Quercus ilex in a mediterranean woodland in order to identify the main environmental drivers of interannual variation in flower and seed production and contrast the impact of climate vs. adaptive factors as main causes of masting. Second, we conducted an experiment of rainfall exclusion to evaluate the effects of an increasing drought (simulating predictions of global change models) on both reproductive processes. The annual seed crop was always affected by environmental factors related to the precipitation pattern, these abiotic factors disrupting the fruiting process at different periods of time. Seed production was strongly dependent upon water availability for the plant at initial (spring) and advanced (summer) stages of the acorn maturation cycle, whereas the final step of seed development was negatively affected by the frequency of torrential-rain events. We also found clear evidence that seed masting in the study species is not only regulated by selective endogenous rhythms, but is mainly a physiological response to the variable environment. Our results from the rainfall exclusion experiment corroborated the conclusions obtained from the 26-year fruiting record and demonstrated that the high interannual variation in seed crop was mainly determined by the success in seed development rather than by the flowering effort. Under a global change scenario, it could be expected that the drier conditions predicted by climate models reinforce the negative effects of summer drought on seed production, leading to negative consequences for tree recruitment and forest dynamics.

Spatial patterns of soil pathogens in declining Mediterranean forests: implications for tree species regeneration

Soil-borne pathogens are a key component of the belowground community because of the significance of their ecological and socio-economic impacts. However, very little is known about the complexity of their distribution patterns in natural systems. Here, we explored the patterns, causes and ecological consequences of spatial variability in pathogen abundance in Mediterranean forests affected by oak decline. We used spatially explicit neighborhood models to predict the abundance of soil-borne pathogen species (Phytophthora cinnamomi, Pythium spiculum and Pythium spp.) as a function of local abiotic conditions (soil texture) and the characteristics of the tree and shrub neighborhoods (species composition, size and health status). The implications of pathogen abundance for tree seedling performance were explored by conducting a sowing experiment in the same locations in which pathogen abundance was quantified. Pathogen abundance in the forest soil was not randomly distributed, but exhibited spatially predictable patterns influenced by both abiotic and, particularly, biotic factors (tree and shrub species). Pathogen abundance reduced seedling emergence and survival, but not in all sites or tree species. Our findings suggest that heterogeneous spatial patterns of pathogen abundance at fine spatial scale can be important for the dynamics and restoration of declining Mediterranean forests.

Relationships between leaf morphological traits, nutrient concentrations and isotopic signatures for Mediterranean woody plant species and communities

Background and aimsSoil factors are driving forces that influence spatial distribution and functional traits of plant species. We test whether two anchor morphological traits—leaf mass per area (LMA) and leaf dry matter content (LDMC)—are significantly related to a broad range of leaf nutrient concentrations in Mediterranean woody plant species. We also explore the main environmental filters (light availability, soil moisture and soil nutrients) that determine the patterns of these functional traits in a forest stand.MethodsFour morphological and 19 chemical leaf traits (macronutrients and trace elements and δ13C and δ15N signatures) were analysed in 17 woody plant species. Community-weighted leaf traits were calculated for 57 plots within the forest. Links between LMA, LDMC and other leaf traits were analysed at the species and the community level using standardised major axis (SMA) regressionsResultsLMA and LDMC were significantly related to many leaf nutrient concentrations, but only when using abundance-weighted values at community level. Among-traits links were much weaker for the cross-species analysis. Nitrogen isotopic signatures were useful to understand different resource-use strategies. Community-weighted LMA and LDMC were negatively related to light availability, contrary to what was expected.ConclusionCommunity leaf traits have parallel shifts along the environmental factors that determine the community assembly, even though they are weakly related across individual taxa. Light availability is the main environmental factor determining this convergence of the community leaf traits.

Acorn removal and dispersal by the dung beetle Thorectes lusitanicus: ecological implications

Abstract 1. Plant–animal interactions, and in particular the processes of seed predation and dispersal, are crucial for tree regeneration and forest dynamics. A novel and striking case of interaction between a dung beetle (Thorectes lusitanicus) and two Quercus species (Q. suber and Q. canariensis) in forests of southern Spain is presented here.

Environmental drivers of mast‐seeding in Mediterranean oak species: does leaf habit matter?

10 paginas.-- 4 tablas.-- 60 referencias.-- Supporting InformationAdditional Supporting Information may be found in the online version of this article

Acorn preference by the dung beetle, Thorectes lusitanicus, under laboratory and field conditions

Processes of seed predation and dispersal are crucial for tree regeneration and forest dynamics. To understand the role of the dung beetle, Thorectes lusitanicus (Col., Scarabaeoidea, Geotrupidae) as secondary seed disperser in Mediterranean oak forests, its food preference was investigated in the field and in the laboratory. This paper had for objectives (1) to explore the feeding preferences for acorns of the two species available in the field: Quercus suber and Quercus canariensis, and (2) to study the olfactory and palatability response of T. lusitanicus to different food resources (oak acorns and dung) under laboratory conditions. The number of beetles in the field was curvilinearly related to the number of acorns, being higher when the number of acorns was intermediate. The maximum values of T. lusitanicus density corresponded to those plots located under Q. suber trees with the highest total weight of acorns. Bioassays with a four-armed olfactometer showed that T. lusitanicus was clearly attracted to volatiles of Q. suber acorns more than to of dung. Palatability bioassays also showed significant preferences for acorns of Q. suber in comparison with the typical food previously described for this beetle species (dung). According to our results, we suggest that a diet based on acorns (due to their high content of polyunsaturated fatty acids) probably satisfies the nutritional requirements of T. lusitanicus during winter, larval development, and metamorphosis.

A neighborhood analysis of the consequences of Quercus suber decline for regeneration dynamics in Mediterranean forests

In forests, the vulnerable seedling stage is largely influenced by the canopy, which modifies the surrounding environment. Consequently, any alteration in the characteristics of the canopy, such as those promoted by forest dieback, might impact regeneration dynamics. Our work analyzes the interaction between canopy neighbors and seedlings in Mediterranean forests affected by the decline of their dominant species (Quercus suber). Our objective was to understand how the impacts of neighbor trees and shrubs on recruitment could affect future dynamics of these declining forests. Seeds of the three dominant tree species (Quercus suber, Olea europaea and Quercus canariensis) were sown in six sites during two consecutive years. Using a spatially-explicit, neighborhood approach we developed models that explained the observed spatial variation in seedling emergence, survival, growth and photochemical efficiency as a function of the size, identity, health, abundance and distribution of adult trees and shrubs in the neighborhood. We found strong neighborhood effects for all the performance estimators, particularly seedling emergence and survival. Tree neighbors positively affected emergence, independently of species identity or health. Alternatively, seedling survival was much lower in neighborhoods dominated by defoliated and dead Q. suber trees than in neighborhoods dominated by healthy trees. For the two oak species, these negative effects were consistent over the three years of the experimental seedlings. These results indicate that ongoing changes in species’ relative abundance and canopy trees’ health might alter the successional trajectories of Mediterranean oak-forests through neighbor-specific impacts on seedlings. The recruitment failure of dominant late-successional oaks in the gaps opened after Q. suber death would indirectly favor the establishment of other coexisting woody species, such as drought-tolerant shrubs. This could lead current forests to shift into open systems with lower tree cover. Adult canopy decline would therefore represent an additional factor threatening the recruitment of Quercus forests worldwide.

The Comparative Effectiveness of Rodents and Dung Beetles as Local Seed Dispersers in Mediterranean Oak Forests

The process of seed dispersal of many animal-dispersed plants is frequently mediated by a small set of biotic agents. However, the contribution that each of these dispersers makes to the overall recruitment may differ largely, with important ecological and management implications for the population viability and dynamics of the species implied in these interactions. In this paper, we compared the relative contribution of two local guilds of scatter-hoarding animals with contrasting metabolic requirements and foraging behaviours (rodents and dung beetles) to the overall recruitment of two Quercus species co-occurring in the forests of southern Spain. For this purpose, we considered not only the quantity of dispersed seeds but also the quality of the seed dispersal process. The suitability for recruitment of the microhabitats where the seeds were deposited was evaluated in a multi-stage demographic approach. The highest rates of seed handling and predation occurred in those microhabitats located under shrubs, mostly due to the foraging activity of rodents. However, the probability of a seed being successfully cached was higher in microhabitats located beneath a tree canopy as a result of the feeding behaviour of beetles. Rodents and beetles showed remarkable differences in their effectiveness as local acorn dispersers. Quantitatively, rodents were much more important than beetles because they dispersed the vast majority of acorns. However, they were qualitatively less effective because they consumed a high proportion of them (over 95%), and seeds were mostly dispersed under shrubs, a less suitable microhabitat for short-term recruitment of the two oak species. Our findings demonstrate that certain species of dung beetles (such as Thorectes lusitanicus), despite being quantitatively less important than rodents, can act as effective local seed dispersers of Mediterranean oak species. Changes in the abundance of beetle populations could thus have profound implications for oak recruitment and community dynamics.

Measurement of fine root tissue density: a comparison of three methods reveals the potential of root dry matter content

AimsRoot tissue density (RTD, the ratio of root dry mass to root volume) is a fundamental trait in comparative root ecology, being increasingly used as an indicator of plant species’ resource use strategy. However, the lack of standardized method to measure this trait makes comparisons tricky. This study aims to compare three methods commonly used for determining fine RTD and to test whether root dry matter content (RDMC, the ratio between root dry mass and root fresh mass) could be used as a surrogate of fine root tissue density.MethodsRTD of 163 fine root samples was determined using (i) Archimedes’ method, (ii) image analysis (WinRHIZO software), and (iii) using the root dry matter content as a proxy. Root samples belonged to different herbaceous species grown in different conditions.ResultsRTD measured with Archimedes’ method was positively correlated with RTD estimated with image analysis and with RDMC. However we demonstrated that RTD measured with Archimedes’ method was better predicted by RDMC (R2 = 0.90) than by RTD measured with image analysis (R2 = 0.56). The performance and limitations of each method were discussed.ConclusionRDMC is a quick, cheap and relatively easy measurable root attribute; we thus recommended its measurement as a proxy of fine root tissue density.