Repercussions of Simulated Climate Change on the Diversity of Woody-Recruit Bank in a Mediterranean-type Ecosystem

Extreme climatic events have the potential to affect plant communities around the world, and especially in the Mediterranean basin, where the frequency of milder and drier summers is expected to be altered under a global-change scenario. We experimentally investigated the effect of three contrasting climatic scenarios on the diversity and abundance of the natural woody-recruit bank among three characteristic habitats in a Mediterranean-type ecosystem: forest, shrubland, and bare soil. The climatic scenarios were dry summers (30% summer rainfall reduction), wet summers (simulating summer storms), and current climatic conditions (control). Seedling emergence and survival after the first summer was recorded during 4 consecutive years. The wet summer boosted abundance and diversity at emergence and summer survival, rendering the highest Shannon H??index. By contrast, the dry summer had no effect on emergence, although survival tended to decline. Nonetheless, the habitat had a key role, bare soil showing almost null recruitment whatever the climatic scenario, and forest keeping the highest diversity in all of them. Our results show that recruit-bank density and diversity depends heavily on extreme climatic events. Community dynamics will depend not only on increased drought but also on the balance between dry and wet years.     

What determines emergence and net recruitment in an early succession plant community? Disentangling biotic and abiotic effects

Question: How do different regeneration scenarios shape species composition at two stages of plant community establishment (emergence and net recruitment) in an early succession? Location: Northern Spain. Methods: In a recently ploughed field, we created eight regeneration scenarios with light, water and nitrogen availability (five replicates each). Seedlings of all species were monitored from emergence to death during one year. Abiotic and biotic variables were measured per quadrat, i.e. soil texture, nutrient contents, seed bank densities and composition, neighbour plant species densitiy and cover. We used partial ordination methods in order to separate the effect of each environmental variable on species composition during emergence and adult net recruitment. Results: Light treatment determined annual plant density at time of emergence and recruitment, while water addition controlled the recruitment of perennials. Resource levels explained the emerged species composition; this effect was not translated into the recruited species composition. N‐addition and N + water addition were strongly associated to species abundances at the time of emergence. Seedling composition in summer was correlated with seed abundance of Cerastium spp. Neighbour species density and cover (mainly Arrhenatherum bulbosum, Agropyron repens and Picris echioides) explained significant fractions of species composition in the emergence and recruitment of the different cohorts. Interactions between species seem to vary in intensity among cohorts and in the key plant species that determined species abundance along succession. Conclusions: Our scenarios exerted contrasting and multilevel effects on the development of our early succession community. Resource availability differently affected plant density and species composition at different life stages. It is relevant to consider different life stages in plant community studies. However, regeneration conditions and other abiotic factors are not enough to explain how community composition varies.     

Sporadic rainy events are more critical than increasing of drought intensity for woody species recruitment in a Mediterranean community

The understanding of the impact of extreme climatic events under a global climate change scenario is crucial for the accurate forecast of future plant community dynamics. We have experimentally assessed the effect of drier and wetter summer conditions on the recruitment probabilities and the growth of seedlings from eight woody species representative of the most important functional groups in the community, pioneer shrubs, mid-successional shrubs and trees, across the main habitats in the study area (open habitat, shrubland, and forest). Our hypothesis proposes that wet summer conditions would represent a good opportunity for tree species regeneration, enhancing both forest maintenance and expansion. A drier summer scenario, on the other hand, would limit forest regeneration, and probably hinder the colonization of nearby habitats. We found a habitat effect on the emergence, survival, and final biomass, whereas different climate scenarios affected seedling survival and biomass. A wet summer boosted growth and survival, whereas greater drought reduced survival only in some cases. These results were modulated by the habitat type. Overall, shrub species presented higher survival and growth and were less affected by more severe drought, whereas some tree species proved to be extremely dependent on wet summer conditions. We conclude that the reduction in frequency of wet summers predicted for the coming decades in Mediterranean areas will have greater consequences for species recruitment than will increased drought. The different response of the species from the various functional groups has the potential to alter the composition and dominance of future plant communities.     

Lowland boreal forests characterization in Algonquin Provincial Park relative to beaver (Castor canadensis) foraging and edaphic factors

Beaver (Caster canadensis) foraging and edaphic conditions can modify the vegetational characteristics of woody plant community in lowland boreal forests. Effective management of these areas requires an understanding of the relative contribution of these factors in shaping the woody plant community structure. Our objective was to quantify the effects of herbivory by beavers and edaphic conditions on woody plant community organization of lowland boreal forests surrounding beaver ponds. Woody vegetation and soils were sampled at 15 ponds occupied by beavers and one other pond abandoned by them in southern Algonquin Park, Ontario. We measured spatial variation in plant diversity, foraging rates and sapling recruitment of trees and shrubs along gradients of beaver foraging intensity and soil moisture, P, K, Mg, and pH. Beavers fed preferentially on a small number of deciduous species and the number of cut stems declined sharply with increasing distance from ponds. Conifers increased in relative dominance to deciduous species in the presence of beavers. Plant species richness and stem and basal area diversity peaked at intermediate distances (about 25 m) from ponds. Sapling recruitment by non-preferred species was positively related to foraging intensity. Total stem abundance and basal area and sapling recruitment by four preferred species (Populus tremuloides, Acer rubrum, Acer saccharum and Corylus cornuta) were negatively related to foraging intensity. However, by including Alnus rugosa and Salix bebbiana (also preferred by beavers) these patterns changed, becoming positively related to foraging intensity. There was also a pronounced gradient in soil moisture, which also decreased with distance from ponds. The other measured edaphic variables did not vary consistently with distance from ponds. Sapling recruitment in mesic versus xeric species varied consistently with hydrid conditions along the moisture gradient, such that variation in moisture also could produce the observed pattern of plant diversity. Diversity patterns changed three years after beaver abandonment of a pond, though sapling recruitment patterns in preferred and non-preferred species around the abandoned pond were similar to the occupied ponds. These observations suggest spatial variation in woody plant richness and diversity could be determined by combined effects of both herbivory (disturbance by beavers) and variable responses of different species to edaphic conditions.     

Oak seedling survival and growth along resource gradients in Mediterranean forests: implications for regeneration in current and future environmental scenarios

Understanding seedling performance across resource gradients is crucial for defining the regeneration niche of plant species under current environmental conditions and for predicting potential changes under a global change scenario. A 2‐year field experiment was conducted to determine how seedling survival and growth of two evergreen and two deciduous Quercus species vary along gradients of light and soil properties in two Mediterranean forests with contrasting soils and climatic conditions. Half the seedlings were subjected to an irrigation treatment during the first year to quantify the effects on performance of an alteration in the summer drought intensity. Linear and non‐linear models were parameterized and compared to identify major resources controlling seedling performance. We found both site‐specific and general patterns of regeneration. Strong site‐specificity was found in the identity of the best predictors of seedling survival: survival decreased linearly with increasing light (i.e. increasing desiccation risk) in the drier site, whereas it decreased logistically with increasing spring soil water content (i.e. increasing waterlogging risk) in the wetter site. We found strong empirical support for multiple resource limitation at the drier site, the response to light being modulated by the availability of soil resources (water and P). Evidence for regeneration niche partitioning among Quercus species was only found at the wetter site. However, at both sites Quercus species shared the same response to summer drought alleviation through water addition: increased first‐year survival but not final survival (i.e. after two years). This suggests that extremely dry summers (i.e. the second summer in the experiment) can cancel out the positive effects of previous wetter summers. Therefore, an increase in the intensity and frequency of summer drought with climate change might cause a double negative impact on Quercus regeneration, due to a general reduction in survival probability and the annulment of the positive effects of (infrequent) ‘wet’ years. Overall, results presented in this study are a major step towards the development of a mechanistic model of Mediterranean forest dynamics that incorporates the idiosyncrasies and generalities of tree regeneration in these systems, and that allow simulation and prediction of the ecological consequences of resource level alterations due to global change.     

What shapes the altitudinal range of a high mountain Mediterranean plant? Recruitment probabilities from ovule to seedling stage

Recruitment is a complex process consisting of sequential stages affected by biotic interactions and abiotic factors. Assessment of these sequential stages and corresponding subprocesses may be useful in identifying the most critical stages. Accordingly, to assess the factors that may determine the altitudinal range limits of the high mountain Mediterranean plant Silene ciliata, a set of demographic stages, from flower production to establishment of 2‐yr‐old plants, and their influence on recruitment probability were examined using a step‐by‐step approach. We integrated florivory, pollination and pre‐dispersal seed predation as pre‐dispersal factors, and seedling emergence and survival as post‐dispersal determinants of recruitment. Three populations were monitored at the southernmost margin of the species along its local altitudinal range. Previous studies suggest that seediness is strongly limited by summer drought especially at the lower boundary of the species, a situation that may worsen under current global warming. Our results showed that recruitment was mainly limited by low seed production in the pre‐dispersal stage and low seedling emergence and survival in the post‐dispersal stage, probably due to environmental harshness in summer. By contrast, biotic factors responsible for propagule loss, such as flower and fruit predation, had a minor effect on the probability of plant recruitment. Although the relative importance of transition probabilities was similar among populations along the altitudinal range, comparatively lower flower production significantly reduced the number of recruited plants at the lowest altitude population. This demographic bottleneck, together with increased competition with other species favoured by climate warming, might collapse population growth and limit persistence at the lower altitudinal range of the species, raising its low local altitudinal edge.     

Effects of timing of precipitation and acorn harvest date on emergence of Quercus emoryi

Abstract. Germination of semi-arid Quercus is strongly correlated with the rainy season. However, the timing and amount of summer precipitation in the southwestern United States is extremely variable. Ongoing and impending changes in global and regional climates are likely to increase this variability. Specifically, anthropogenically induced changes in general circulation patterns may alter the seasonal distribution of precipitation in a directional manner. In addition to climatic variability, the inter- and intra-annual variability of Quercus emoryi Torr. (Emory oak) acorn maturation also is high. Therefore, in light of existing climatic variability and potential future climate change, we conducted a greenhouse experiment to investigate the effects of acorn maturation date and the timing of the onset of the ‘monsoon’ on emergence of Q. emoryi seedlings. Acorns were collected at weekly intervals in July of 1995 and 1996, planted in a greenhouse, and subjected to different watering treatments. Watering treatments were (1) acorns watered the same day as planting; (2) acorns watered two weeks after planting; (3) acorns watered four weeks after planting. Emergence in 1995 was significantly greater in the last week of acorn maturation and decreased significantly as time to the onset of the ‘monsoon’ increased. Emergence in 1996 was very low, presumably because of soil moisture contents that were lower than those observed in 1995. The dependence of Q. emoryi on water for emergence has profound implications for Q. emoryi recruitment in the face of current climatic variability and future climate change. Summers with a delayed ‘monsoon’ and decreased soil moisture may severely constrain recruitment of this woody plant.     

Antagonistic effects of large- and small-scale disturbances on exotic tree invasion in a native tussock grassland relict

It is generally accepted that disturbances increase community invasibility. Yet the role of disturbance in plant invasions may be less predictable than often assumed, due to the influence of environmental stochasticity and interactions between disturbance regimes. We evaluated the single and interactive effects of prescribed burning (large-scale, infrequent event) and animal diggings (small-scale, frequent events) on the invasion success of Gleditsia triacanthos L. in a tussock grassland relict of the Inland Pampa, Argentina. Tree seedling emergence and survival were monitored over 4 years, after adjusting for propagule pressure through copious seed addition to all disturbance treatments. Burning altered community structure by suppressing tussock grasses and promoting exotic forbs, whereas simulated, armadillo-like diggings had little impact on herbaceous composition. Overall, seedling emergence rather than survival represented the main demographic bottleneck for tree invasion. Tree establishment success varied among seedling cohorts emerged in different climatic years. In a dry year, emergence was only slightly affected by disturbances. In contrast, for two consecutive wet years, initial burning and armadillo-like diggings exerted strong, antagonistic effects on tree recruitment. Whereas fire alone increased recruitment, the simulated burrowing regime prevented seedling emergence in both burned and unburned plots. The latter effect might be explained by reduced soil moisture, and increased seed burial and predation in excavated patches. Thus, the impact of a single, large-scale perturbation promoting woody plant invasion was overridden by a regime of small-scale, frequent disturbances. Our results show that grassland invasibility was contingent on inter-annual climatic variation as well as unexpected interactions between natural and anthropogenic disturbance agents.     

Capsule Treatments to Enhance Seedling Emergence of Gaura neomexicana ssp. coloradensis

Management of riparian vegetation is difficult because these communities are frequently impacted by herbivores, invasive weeds, and altered hydrologic regimes. Multiple and intertwined factors affecting rare species recruitment are particularly difficult to identify. Gaura neomexicana ssp. coloradensis Munz (Gaura) is a short‐lived perennial forb endemic to riparian areas in mixed‐grass prairies of Wyoming, Nebraska, and Colorado, U.S.A. It became a federally listed threatened species in October 2000. Because the species is a recruitment‐limited monocarpic perennial, we studied the effects of six capsule‐collection dates, a 2‐month cool‐moist stratification, 24‐hr leaching, and 24‐hr imbibition on Gaura seedling emergence. Seedling emergence did not vary with collection date. Capsules collected from Gaura plants grown at the Bridger Plant Materials Center in Montana exhibited greater emergence than capsules harvested from endemic populations near Cheyenne, Wyoming, suggesting that maternal plant growing conditions impact dormancy. Because cool‐moist stratification enhanced seedling emergence of Gaura and leaching did not, sufficient moisture during cool temperatures may be more critical than leaching of germination inhibitors as might occur with normal stream flows. Spring flooding may enhance Gaura recruitment by increasing the availability of riparian sites that are inundated during periods of cool temperatures. If so, hydrologic and climatic regimes must be considered in restoring the unique conditions needed for germination of this rare riparian endemic.     

Quercus ilex recruitment in a drier world: A multi-stage demographic approach

There is a growing interest in understanding and forecasting the responses of plant communities to projected changes of environmental conditions. Multi-stage demographic approaches, where plant recruitment is explored across multiple and consecutive stages, are essential to obtain a whole overview of the consequences of increasing aridity on tree recruitment and forest dynamics, but they are still rarely used. In this study, we present the results of an experimental rainfall exclusion aimed to evaluate the impact of projected increasing drought on multiple stage-specific probabilities of recruitment in a key tree species typical of late-successional Mediterranean woodlands (Quercus ilex L.). We calibrated linear and nonlinear likelihood models for the different demographic processes and calculated overall probabilities of recruitment along a wide range of microhabitat conditions. Rainfall exclusion altered Q. ilex recruitment throughout ontogeny. Seed maturation, seedling emergence and survival and, to a lesser extent, post-dispersal seed survival were the most sensitive demographic processes to decreased rainfall. Interestingly, both the identity of the most critical stages for recruitment and their specific sensitivity to rainfall manipulation depended largely on the yearly pattern of precipitation. The microhabitat heterogeneity strongly determined the success of recruitment in the study species. The experimental increase in drought displaced the peak of maximum overall recruitment towards the low end of the light gradient, suggesting that the dependence on shrubs for an effective recruitment in Q. ilex could be intensified under future environmental scenarios. In terms of phenotypic plasticity, Q. ilex seedlings responded more strongly to light availability than to experimentally increased drought, which could reduce its ability to persist under on-going environmental conditions due to climate change. Results from this study provide a full picture of the ecological and functional consequences of the projected rainfall reduction on tree recruitment and forest dynamics in two years of contrasting precipitation.