Genetics of sprouting: effects of long-term persistence in fire-prone ecosystems

Fire functional traits (postfire resprouting and seeding) are considered to be adaptations for persisting in fire-prone environments. Although ecological and evolutionary consequences of sprouting have been extensively discussed, within-species genetic variability and structure are unknown. Here we report levels and distribution patterns of genetic polymorphisms in postfire stands of the predominant sprouter Nothofagus antarctica . Fresh foliage of 50 individuals was collected following a spatially explicit sampling design for isozyme analysis from two replicates of each of four habitat types inhabited by the species in northwestern Patagonia, Argentina: matorral, high elevation, forest, and temporally flooded basins. Average polymorphism per population ranged from 44% to 78% and mean gene diversity per site H _S varied from 0.187 to 0.274. These results show that sprouter populations hold considerable genetic variation. Significant genetic structure over short distances (< 50 m) was found at all locations. Ancient fine-scale genetic structure is preserved by occasional seedling establishment that results in high co-ancestry coefficients. Sprouter populations growing in suboptimal habitats such as matorral, high elevation or basins consist of pairs of heterozygous genets that occur at larger spatial scales as a result of micro-environmental heterogeneity and/or local competition between near neighbour genotypes. In contrast, homozygous pairs of individuals for distinct isozyme loci occurred at larger spatial scales in forest stands. This indicates that biparental inbreeding due to local propagule establishment may take place to some extent in sprouters growing under favourable conditions. Our results show that sprouters follow a long-lasting genet persistence strategy which most probably is selected under unpredictable disturbance regimes, such as fire.     

Rainfall reliability, a neglected factor in explaining convergence and divergence of plant traits in fire-prone mediterranean-climate ecosystems

Aim Rainfall reliability has been neglected as a determinant of plant trait convergence and divergence in mediterranean‐climate ecosystems. This paper reports on patterns of rainfall reliability — quantified as interannual variation in monthly and seasonal rainfall, and as the frequency of individual events in terms of their size, duration and intensity — for four fire‐prone mediterranean‐climate ecosystems. Location The four mediterranean‐climate regions of the world with fire‐prone ecosystems, namely SW Cape (South Africa), SW Australia, California and the Mediterranean Basin (Andalusia, Spain). Methods Using long‐term monthly rainfall data from stations dispersed across the four regions, we computed monthly means and interannual variation for each month of the year — the latter quantified as the coefficient of variation (CV) — and divided these into winter and summer seasons. We also computed the mean number of rainfall events, the mean frequency in various categories of event duration (days), the amount of rainfall per event (mm) and the rainfall intensity per event (mm/day) per year for winter and summer seasons for a subset of the rainfall stations. Results The fraction of rain falling in summer was lowest in California (5%) and similarly low (c. 25%) in the other three regions. The hierarchy of values of coefficient of variation (CV) of monthly rainfall during the winter period was as follows: California > Andalusia >> SW Cape > SW Australia; results for summer were: California > > Andalusia >> SW Australia ～ SW Cape. SW Australian sites experienced the greatest frequency of short, small and low‐intensity rainfall events in both seasons; patterns in the SW Cape were intermediate between Australia and the two northern hemisphere sites which both received fewer, larger and more intense events. Overall, the two southern hemisphere regions (SW Australia and the SW Cape) had significantly more reliable regimes than the two northern hemisphere ones (Mediterranean Basin and California). Main conclusions These differences in rainfall reliability regimes may provide a novel perspective on the distribution of certain plant life‐history traits in mediterranean‐climate ecosystems. Less reliable regimes would select for germination and seedling survival traits that enable persistence of genets in the face of uncertain moisture conditions during the winter and spring establishment phase. Study systems that accommodate for phylogenetic constraints, namely invasive species derived from mediterranean‐climate ecosystems, as well as shared lineages, provide good opportunities to develop and test hypotheses on the implication of different rainfall reliability regimes. One of the novel implications of this study is that the distinctive trait of assemblages in the southern hemisphere regions may be a consequence not so much of their shared nutrient‐poor soils as of their similarly reliable rainfall regimes.     

Decomposition of woody debris in Mediterranean ecosystems: the role of wood chemical and anatomical traits

Plant and Soil - Data about woody debris (WD) decomposition are very scarce for the Mediterranean basin. The specific aim of this work is to explore the relationships between WD traits with the...     

Ecology of plant resprouting: populations to community responses in fire-prone ecosystems

Resprouting has emerged as a key functional trait in plant ecology over the past decade with more than 400 papers published since 2000 (Web of Science). This special issue of Plant Ecology brings together a set of papers that advance our understanding of this functional trait, in the quest for developing a better conceptual framework for predicting community response to disturbance. This special issue highlights current research on all aspects of the effects of fire and other disturbances on plant resprouting behaviour and the importance of the persistence niche in structuring plant communities.     

Plant invasions in undisturbed ecosystems: The triggering attribute approach

The invasion of a target community by a non‐indigenous plant species includes the stages of arrival, establishment and spread, which tend to depend on different characteristics of the invasive species and its context. While the mechanisms behind the invasion of highly disturbed ecosystems are well known, our understanding of the invasion process in undisturbed or weakly disturbed ecosystems is much more limited. Here we propose that, once a non‐indigenous species has arrived to a new ecosystem and become established, the likelihood that it spreads, and thus becomes invasive, may depend on just one or very few characteristics, called‘triggering attributes’(TA). We propose that a TA is a vegetative or regenerative attribute discontinuously distributed in comparison to the resident community. This attribute allows the species to benefit from a resource that is permanently or temporarily unused by the resident community. We present an original study case and examples from the literature to illustrate our approach, and we also propose some ways to test it in different ecosystems.     

Plant traits, litter quality and decomposition in a Mediterranean old-field succession

Human-induced changes in land use lead to major changes in plant community composition which have strong effects on ecosystem processes. Here, we tested the hypothesis that changes in traits of living plants induced by such changes resulted in changes in the quality and decay properties of the litter produced by the different communities. This was done in the context of a secondary succession following land abandonment in the Mediterranean region of Southern France. During the course of succession, species with high specific leaf area (the ratio of leaf area to leaf mass), low leaf dry matter content (the ratio of leaf dry mass to leaf fresh mass) and high leaf nitrogen concentration were progressively replaced by species with opposite characteristics. Accordingly, the initial litter concentrations of carbon (C) and nitrogen (N) decreased, while their C:N ratio and their hemicellulose concentration increased with time after abandonment. Early-successional communities had faster rates of litter decay and N release from litter, but these differences damped out with decomposition time. Nitrogen release from litter was related to initial litter chemical composition, particularly to its N concentration. This also held for litter decay rate, but only during the first 18 months of decomposition. Community functional parameters (i.e. trait values weighed according to the relative abundance of species) were tightly linked to initial litter N concentration, and thereby to litter decay and N loss rates. The strongest correlations were found with leaf dry matter content, which therefore appears as a powerful marker of litter properties. This provides further evidence that characteristics of living leaves persist in litter, and that some ecosystem processes can be inferred from plant functional traits. Responsible Editor: Alfonso Escudero     

Plant traits determine the phylogenetic structure of arbuscular mycorrhizal fungal communities

Functional diversity in ecosystems has traditionally been studied using aboveground plant traits. Despite the known effect of plant traits on the microbial community composition, their effects on the microbial functional diversity are only starting to be assessed. In this study, the phylogenetic structure of arbuscular mycorrhizal (AM) fungal communities associated with plant species differing in life cycle and growth form, that is, plant life forms, was determined to unravel the effect of plant traits on the functional diversity of this fungal group. The results of the 454 pyrosequencing showed that the AM fungal community composition differed across plant life forms and this effect was dependent on the soil collection date. Plants with ruderal characteristics tended to associate with phylogenetically clustered AM fungal communities. By contrast, plants with resource‐conservative traits associated with phylogenetically overdispersed AM fungal communities. Additionally, the soil collected in different seasons yielded AM fungal communities with different phylogenetic dispersion. In summary, we found that the phylogenetic structure, and hence the functional diversity, of AM fungal communities is dependent on plant traits. This finding adds value to the use of plant traits for the evaluation of belowground ecosystem diversity, functions and processes.     

Germination traits explain soil seed persistence across species: the case of Mediterranean annual plants in cereal fields

Background and Aims

Seed persistence in the soil under field conditions is an important issue for the maintenance of local plant populations and the restoration of plant communities, increasingly so in the light of rapidly changing land use and climate change. Whereas processes important for dispersal in space are well known, knowledge of processes governing dispersal in time is still limited. Data for morphological seed traits such as size have given contradictory results for prediction of soil seed persistence or cover only a few species. There have been few experimental studies on the role of germination traits in determining soil seed persistence, while none has studied their predictive value consistently across species. Delayed germination, as well as light requirements for germination, have been suggested to contribute to the formation of persistent seed banks. Moreover, diurnally fluctuating temperatures can influence the timing of germination and are therefore linked to seed bank persistence.

Methods

The role of germination speed measured by T₅₀ (days to germination of 50 % of all germinated seeds), light requirement and reaction to diurnally fluctuating temperatures in determining seed persistence in the soil was evaluated using an experimental comparative data set of 25 annual cereal weed species.

Key Results

It is shown that light requirements and slow germination are important features to maintain seeds ungerminated just after entering the soil, and hence influence survival of seeds in the soil. However, the detection of low diurnally fluctuating temperatures enhances soil seed bank persistence by limiting germination. Our data further suggest that the effect of diurnally fluctuating temperatures, as measured on seeds after dispersal and dry storage, is increasingly important to prevent fatal germination after longer burial periods.

Conclusions

These results underline the functional role of delayed germination and light for survival of seeds in the soil and hence their importance for shaping the first part of the seed decay curve. Our analyses highlight the detection of diurnally fluctuating temperatures as a third mechanism to achieve higher soil seed persistence after burial which interacts strongly with season. We therefore advocate focusing future research on mechanisms that favour soil seed persistence after longer burial times and moving from studies of morphological features to exploration of germination traits such as reaction to diurnally fluctuating temperatures.     

Plant chemical traits define functional and phylogenetic axes of plant biodiversity

To determine which types of plant traits might better explain ecosystem functioning and plant evolutionary histories, we compiled 42 traits for each of 15 perennial species in a biodiversity experiment. We used every possible combination of three traits to cluster species. Across these 11,480 combinations, clusters generated using tissue %Ca, %N and %K best mapped onto phylogeny. Moreover, for the 15 best combinations of three traits, 82% of traits were chemical, 16% morphological and 2% metabolic. The diversity-dependence of ecosystem productivity was better explained by the %Ca, %N and %K clusters: compared to adding a new species at random, adding a species from an absent cluster/clade better-explained gains in productivity. Species number impacted productivity only when all clusters were present. Our results suggest that tissue elemental chemistry might be more phylogenetically conserved and more strongly related to ecosystem functioning than commonly measured morphological and physiological traits, a possibility that merits exploration.     

Plant traits related to leaf decomposition processes in arid ecosystems of northern Patagonia

Aims Plants play an important role in ecosystem processes. Functional meaning of trait variation in wide environmental gradients is well known but is scarcely known across narrow gradients. We analyze the variation of morphological, physical and chemical traits of dominant plant species and the potential rates of dry mass loss and N release/immobilization during senesced leaf decomposition of these species across a narrow aridity gradient, and to identify indicative traits useful to set species functional groups sharing decomposition patterns.     

Social evolution in the genusHalictus: a phylogenetic approach

Summary Halictine bees exhibit an enormous diversity of solitary and social colony structures. To investigate social evolution in the genusHalictus, phylogenies of 15 species of the subgeneraH. (Halictus) andH. (Seladonia) were constructed based on protein electrophoretic data. Solitary, social, and socially polymorphic species were included.Halictus (Seladonia) apparently rendersH. (Halictus) paraphyletic. The common ancestor ofH. (Halictus) andH. (Seladonia) was probably social or socially polymorphic. This implies that some solitary and socially polymorphic species, such asH. confusus andH. tumulorum, represent evolutionary reversals from a completely eusocial condition to the solitary condition that is thought to be primitive for the subfamily as a whole.     

Effective size and the persistence of ecosystems

Summary Ecosystems distributed in space have an effective size, reflecting both their absolute size (extent) and their fine-scale physical structure (viscosity). In this paper, a general mathematical model of a predator-prey interaction is presented via the phase-plane graphs of Rosenzweig and MacArthur (1963) to show one reason why ecosystems of larger effective size should persist longer than smaller ones: oscillations of population densities tend to be displaced farther from extinction thresholds—even in spatially homogeneous systems. Experimental results obtained by Gause and Luckinbill with protozoa and Huffaker with mites are interpreted in this context.     

Response of plant functional types to changes in the fire regime in Mediterranean ecosystems: A simulation approach

Abstract. In the Mediterranean basin, the climate is predicted to be warmer and effectively drier, leading to changes in fuel conditions and fire regime. Land abandonment in the Mediterranean basin is also changing the fire regime through the increase in fuel loads. In the present study, two simulation models of vegetation dynamics were tested in order to predict changes in plant functional types due to changes in fire recurrence in eastern Spain. The two modelling approaches are the FATE-model (based on vital attributes) and the gap model BROLLA (based on the gap-phase theory). The models were arranged to simulate four functional types, based mainly on their regenerative strategies after disturbance: Quercus (resprouter), Pinus (non-resprouter with serotinous cones), Erica (resprouter), and Cistus (non-resprouter with germination stimulated by fire). The simulation results suggested a decrease in Quercus abundance, an increase in Cistus and Erica, and a maximum of Pinus at intermediate recurrence scenarios. Despite their different approaches, both models predicted a similar response to increased fire recurrence, and the results were consistent with field observations.     

Plant functional traits and diversity in sand dune ecosystems across different biogeographic regions

Plant species of a functional group respond similarly to environmental pressures and may be expected to act similarly on ecosystem processes and habitat properties. However, feasibility and applicability of functional groups in ecosystems across very different climatic regions have not yet been studied. In our approach we specified the functional groups in sand dune ecosystems of the Mediterranean, Hyrcanian and Irano-Turanian phytogeographic regions. We examined whether functional groups are more influenced by region or rather by habitat characteristics, and identified trait syndromes associated with common habitat types in sand dunes (mobile dunes, stabilized dunes, salt marshes, semi-wet sands, disturbed habitats). A database of 14 traits, 309 species and 314 relevés was examined and trait-species, trait-plot and species-plot matrices were built. Cluster analysis revealed similar plant functional groups in sand dune ecosystems across regions of very different species composition and climate. Specifically, our study showed that plant traits in sand dune ecosystems are grouped reflecting habitat affiliation rather than region and species pool. Environmental factors and constraints such as sand mobility, soil salinity, water availability, nutrient status and disturbance are more important for the occurrence and distribution of plant functional groups than regional belonging. Each habitat is shown to be equipped with specific functional groups and can be described by specific sets of traits. In restoration ecology the completeness of functional groups and traits in a site may serve as a guideline for maintaining or restoring the habitat.     

Directional biases in phylogenetic structure quantification: a Mediterranean case study

Recent years have seen an increasing effort to incorporate phylogenetic hypotheses to the study of community assembly processes. The incorporation of such evolutionary information has been eased by the emergence of specialized software for the automatic estimation of partially resolved supertrees based on published phylogenies. Despite this growing interest in the use of phylogenies in ecological research, very few studies have attempted to quantify the potential biases related to the use of partially resolved phylogenies and to branch length accuracy, and no work has examined how tree shape may affect inference of community phylogenetic metrics. In this study, we tested the influence of phylogenetic resolution and branch length information on the quantification of phylogenetic structure, and also explored the impact of tree shape (stemminess) on the loss of accuracy in phylogenetic structure quantification due to phylogenetic resolution. For this purpose, we used 9 sets of phylogenetic hypotheses of varying resolution and branch lengths to calculate three indices of phylogenetic structure: the mean phylogenetic distance (NRI), the mean nearest taxon distance (NTI) and phylogenetic diversity (stdPD) metrics. The NRI metric was the less sensitive to phylogenetic resolution, stdPD showed an intermediate sensitivity, and NTI was the most sensitive one; NRI was also less sensitive to branch length accuracy than NTI and stdPD, the degree of sensitivity being strongly dependent on the dating method and the sample size. Directional biases were generally towards type II errors. Interestingly, we detected that tree shape influenced the accuracy loss derived from the lack of phylogenetic resolution, particularly for NRI and stdPD. We conclude that well‐resolved molecular phylogenies with accurate branch length information are needed to identify the underlying phylogenetic structure of communities, and also that sensitivity of phylogenetic structure measures to low phylogenetic resolution can strongly vary depending on phylogenetic tree shape.     

Scavenging in Mediterranean ecosystems: effect of the invasive Argentine ant

Above-ground invertebrates may represent a high proportion of animal biomass, but few data are available on their fate after death. In Mediterranean ant communities, they are frequently scavenged by ants. Here, we assessed the consequences of Argentine ant invasion on the removal of arthropod corpses in Doñana National Park (SW Spain). In three natural habitats that differed in their degree of vegetation cover (i.e. protection for ants against high temperatures), we experimentally provided dead Drosophila, and observed their disappearance over a 60-min period at different times of day and year. The habitats used were isolated cork oak trees, pine tree forest and dry scrubland; we compared invaded with uninvaded plots in each. Oak trees were the most invaded habitat, while scrubland was the least and the only one where the Argentine ant coexisted with native ant species. In accordance with this degree of invasion, the Argentine ant removed the highest percentage of dead flies in oak trees and the lowest in scrubland. Its performance as scavenger was higher than uninvaded ant communities, but it was reduced at high temperatures, when native species were highly efficient. The saturated distribution of the Argentine ant colony seems to be the key to its efficiency. We discuss how the occurrence and scavenger efficiency of the Argentine ant could affect the nutrient cycling and the progression of its invasion.