Non‐fire induced seed release ina weakly serotinous pine: climatic factors,maintenance costs or both?

The advantages of canopy seed retention (serotiny) for plants inhabiting fire‐prone ecosystems are well documented. However, very few species are completely serotinous and non‐fire induced opening of serotinous fruits is commonly observed (weak serotiny). Two non‐mutually exclusive causes are envisaged to contribute to this process: mechanical changes in serotinous fruits mediated by climatic conditions (e.g. drought) or the costs of maintenance for the plant of these long‐lasting structures. However, their relative contribution to the spontaneous opening of serotinous fruits remains elusive as well as the consequences for the build‐up of the canopy seed bank and inter‐individual differences in serotiny. In this study we monitored the dynamics of cone production and cone opening in the weakly serotinous Pinus halepensis for five years (2004–2008), including two severe drought episodes (2005, 2006). Drought decreased the production of conelets, increased the abortion of immature cones, reduced the seed quality in the cohorts of cones produced during these years, and increased the opening of serotinous cones. During the first drought episode, a higher proportion of serotinous cones opened in those pines bearing a larger crop of younger cones. This suggests that not only passive changes induced by drought but also competition among cones for resources (e.g. water) might be involved in this process. The opening of serotinous cones in pines bearing more cones made inter‐individual differences in the size of the canopy cone bank to narrow or even to reverse from 2004 to 2008. These results may help to understand the decrease in serotiny when pines grow and accumulate more cones and the large inter‐individual variability in the degree of serotiny observed in P. halepensis forests. In addition, the negative effects of drought episodes for the size of the canopy cone bank and the seeds contained can be an unexplored cause of post‐fire regeneration constraint.     

Influence of clay licks on the diversity and structure of an Amazonian forest

The spatial heterogeneity of resource availability is a major driver of biodiversity patterns. Some environmental conditions and resources are characterized by large‐scale patterns of variation within the landscape. Clumped local discontinuities or discrete elements also increase spatial heterogeneity, promoting local ‘biodiversity hot spots’ by modifying habitat characteristics and promoting plant–animal interactions. Clay licks are faunal attractors owing to their role in the nutritional ecology of the user species; nevertheless, the effect of their presence on the surrounding vegetation has been poorly quantified. Here, we use data from 100 × 10 m transects and evaluate the effects of the presence of clay licks on forest diversity and structure at local and landscape scales. In clay lick areas, there was a higher abundance of certain species, which helps to homogenize species composition between localities counteracting the natural distance‐decay of compositional similarity between transects without clay lick influence (controls). Compared to control sites, clay lick′s forests had higher palm densities, shorter but more variable individuals in the canopy and understory, a thinner canopy layer, and denser herbaceous and ground level covers. These differences were found along the whole length of transects in both sampled areas types. These results reveal that the presence of discrete elements (i.e., clay licks) may help to explain the compositional and structural heterogeneity of Amazonian forests influencing ecological processes such as seed dispersal and trampling. These considerations may be relevant for other biomes where clay licks are present and give weight to their inclusion in conservation initiatives in tropical forests.     

Constraints and trade-offs in Mediterranean plant communities: The case of holm oak-Aleppo pine forests

In this paper we review those aspects that are relevant to the development of a mechanistic ecological theory to account for the structure and dynamics of Mediterranean forests, focusing our attention on mixed forests of holm oak (Quercus ilex L.), a shade-tolerant, slowgrowing species that resprouts vigorously after disturbance, and Aleppo pine (Pinus halepensis M.), a fast-growing, nonresprouting, shade-intolerant species. The main objectives of this report are: to introduce some of the primary features of these forests, showing their structural complexity and historical peculiarities; to show that much of this complexity can be conceptually reduced to two main factors of variation, soil-moisture gradients and a complex interaction of historical management and disturbance regimes; and to contrast the unique features of Mediterranean systems with other communities that have inspired generalization in ecology. Plants in Mediterranean-climate regions must face several environmental constraints during their life cycle: water limitation, competition for light, and a complex set of disturbance regimes, mainly fire, herbivory, and human exploitation. The response of co-occurring species to a given set of environmental constraints depends on a combination of physiological and morphological traits. In holm oak-Aleppo pine forests, the lower limit of distribution along a soil-moisture gradient appears to be controlled by dry-season water stress on seedling performance, and the upper limit seems to be controlled by shade tolerance relative to competitors. The processes that generate and maintain these patterns are related to the responses of the two species to the water and light environments that result from interacting gradients of disturbance and resource availability. The dynamics of mixed holm oak-Aleppo pine forests may be represented along two major environmental axes: water availability and light intensity; namely, time since last disturbance. At the regional scale, the presence of holm oak and Aleppo pine is expected to be driven mainly by the precipitation regime, with the proportion of Aleppo pine increasing toward the driest border and with holm oak being the dominant species in areas with higher precipitation. Changes of dominance of holm oak and Aleppo pine also respond to water availability at the local scale. In this case, variations between species depend on different factors in a complex way, because reduced soil-moisture levels may result either from low precipitation or from topography and edaphic features. The dynamics of holm oak-Aleppo pine forests are also determined by temporal changes in canopy closure; that is, forest recovery after disturbance. In this case, the proportion of Aleppo pine would increase in recently disturbed stands (i.e., with high light intensity reaching the forest floor), whereas regeneration of holm oak would be dominant under partially closed canopies. Theories of forest dynamics developed in humid regions may apply only poorly to Mediterranean plant communities, where vegetation change is qualitatively or quantitatively different. Thus, succession in temperate forests appears to be driven by differences in light availability and shade tolerance; but in Mediterranean plant communities, water limitation is of greater importance for the distribution of forest species. In Mediterranean landscapes the interaction of life-history strategies with changing environments is difficult to infer from observational and experimental studies. A mechanistic approach, in which competition or plant performance is measured as a function of resource availability, seems more feasible. The idea should be to develop multispecies models calibrated specifically for Mediterranean forests in a combined program of modeling, field research, and experimentation.

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Resumen  En este trabajo se revisan los aspectos más relevantes en el desarrollo de una teoría ecológica mecanicista sobre la estructura y dinámica de los bosques mediterráneos, en concreto sobre los bosques mixtos de encina (Quercus ilex L.), una especie tolerante a la sombra, de crecimiento lento, y que rebrota vigorosamente después de una perturbación, y pino carrasco (Pinus halepensis M.), una especie de crecimiento rápido, no rebrotadora, e intolerante a la sombra. Los principales objetivos de este estudio son: introducir algunas de las principales características de estos bosques, mostrando su complejidad estructural y sus peculiaridades históricas; mostrar que gran parte de esta complejidad puede reducirse conceptualmente a dos principales factores de variación, la disponibilidad de agua en el suelo y una compleja interaction de regímenes de gestión y perturbación; y contrastar las características únicas de los sistemas mediterráneos con las de otros sistemas en los que se han basados gran parte de las generalizaciones que se han hecho en ecologia. En las regiones de clima mediterráneo, las plantas deben enfrentarse a diferentes restricciones: limitación hídrica, competencia por la luz, y un conjunto complejo de regímenes de perturbación, principalmente fuego, herbivoría, y explotación humana. La respuesta de las especies a un conjunto de limitaciones ambientales depende de la combinación de caracteres fisiológicos y morfológicos. En los bosques de pino carrasco y encina, el límite inferior de distribución de las especies viene controlado por el estrés hídrico de la estación, mientras que el límite superior viene controlado por la tolerancia a la sombra. Los procesos que generan y mantienen estos patrones están relacionados con las respuestas de ambas especies a lo largo de su ciclo de vida a los ambientes de luz y agua que se generan como resultado de la interacción de los gradientes de perturbación y disponibilidad de recursos. La dinámica de los bosques mixtos de pino carrasco y encina se puede representar a lo largo de dos principales ejes ambientales: disponibilidad de agua e intensidad de luz; o, lo que es lo mismo, tiempo desde la última perturbación. A una escala regional, la presencia de pino carrasco y encina viene determinada por el régimen de precipitación: el pino carrasco aumenta hacia el extremo más seco, mientras que la encina se hace dominante en áreas con mayor precipitación. A escala local, los cambios en la dominancia de ambas especies también responden a la disponibilidad de agua. En este caso las variaciones entre especies dependen de diferentes factores, ya que los niveles bajos de humedad del suelo pueden ser debidos a la baja precipitación, pero también a la topografía o a las características edáficas. La dinámica de los bosques de pino y encina también depende de los cambios temporales en el cierre de la cubierta, es decir, de la recuperación del bosque después de la perturbación. En este caso, la proporción de pino carrasco se incrementa en los rodales recién perturbados (en los cuales llega mucha luz al suelo del bosque), mientras que la regeneración de encina domina bajo cubiertas parcialmente cerradas. Las teorías de dinámica forestal desarrolladas en las regiones húmedas se aplican con dificultad en las comunidades de plantas mediterráneas, donde los cambios de la vegetación suelen ser cualitativamente y cuantitativamente distintos. Así, en los bosques templados la sucesión viene determinada por diferencias en la disponibilidad de luz y la tolerancia a la sombra de las diferentes especies, mientras que en las comunidades mediterráneas la limitación hídrica es más importante a la hora de explicar la distribución de las especies forestales. En los ecosistemas mediterráneos, la interacción de las estrategias del ciclo vital con los cambios en el ambiente dificilmente pueden deducirse a partir de estudios observacionales y experimentales. Se requiere una aproximación mecanicista, en la cual la competencia o el éxito de las plantas se pueda medir en función de la disponibilidad de recursos. La idea sería desarrollar modelos multiespecíficos calibrados específicamente para los bosques mediterráneos, en los que se combinara la modelización con los estudios de campo y la experimentación.

     

Patterns of seedling recruitment in West-Mediterranean Quercus ilex forest influenced by canopy development

Abstract. Quercus ilex (holm oak) coppice forests belong to the most representative communities in the western part of the Iberian Peninsula. Due to the high sprouting potential of holm oak after coppicing, the dynamics of recruitment of new individuals in existing populations has been largely overlooked. In these forests, the density of seedlings increases in old stands, as a result of the remarkable increase in the recruitment of 1-yr and 2-yr old seedlings, probably due to better environmental conditions for germination. Older seedlings show a tendency towards increasing survival under some degree of canopy closure, since their density appears to be higher just before complete canopy closure takes place. Nevertheless, the reduced growth of seedlings in older stands due to low light levels causes age distributions of holm oak seedlings to change drastically, with saplings older than 15 yr becoming very unusual in these stands. Therefore, a bottleneck of recruitment appears at this sapling stage and prevents vigorous regeneration just before thinning. Present conditions of genet density and management tend to prevent population renewal. Dynamics of the seedling bank reinforce the role of gap formation through small or large scale perturbations in population turnover.     

The Interplay among Acorn Abundance and Rodent Behavior Drives the Spatial Pattern of Seedling Recruitment in Mature Mediterranean Oak Forests

The patterns of seedling recruitment in animal-dispersed plants result from the interactions among environmental and behavioral variables. However, we know little on the contribution and combined effect of both kinds of variables. We designed a field study to assess the interplay between environment (vegetation structure, seed abundance, rodent abundance) and behavior (seed dispersal and predation by rodents, and rooting by wild boars), and their contribution to the spatial patterns of seedling recruitment in a Mediterranean mixed-oak forest. In a spatially explicit design, we monitored intensively all environmental and behavioral variables in fixed points at a small spatial scale from autumn to spring, as well as seedling emergence and survival. Our results revealed that the spatial patterns of seedling emergence were strongly related to acorn availability on the ground, but not by a facilitationeffect of vegetation cover. Rodents changed seed shadows generated by mother trees by dispersing most seeds from shrubby to open areas, but the spatial patterns of acorn dispersal/predation had no direct effect on recruitment. By contrast, rodents had a strong impact on recruitment as pilferers of cached seeds. Rooting by wild boars also reduced recruitment by reducing seed abundance, but also by changing rodent’s behavior towards higher consumption of acorns in situ. Hence, seed abundance and the foraging behavior of scatter-hoarding rodents and wild boars are driving the spatial patterns of seedling recruitment in this mature oak forest, rather than vegetation features. The contribution of vegetation to seedling recruitment (e.g. facilitation by shrubs) may be context dependent, having a little role in closed forests, or being overridden by directed seed dispersal from shrubby to open areas. We warn about the need of using broad approaches that consider the combined action of environment and behavior to improve our knowledge on the dynamics of natural regeneration in forests.     

Complex selection on life-history traits and the maintenance of variation in exaggerated rostrum length in acorn weevils

Trophic interactions can trigger the development of exaggerated specialized characters and promote morphological diversification. For example, acorn weevils (genus Curculio) present strikingly long rostrums, which are used by females to perforate oviposition holes through the seed coat. Species exhibiting longer rostrums are known to exploit larger acorns, and therefore rostrum length is thought to be subject to selection to match the preferred acorn type. However, rostrum length is strongly correlated with body size, and morphological divergence could result from either selection on rostrum length for optimal food exploitation or from other pressures acting on body size. We collected infested acorns at oak forests where the large Curculio elephas and the small-bodied Curculio glandium co-occur. There were no interspecific differences in adult female body size to rostrum length allometric relationships, and rostrum length is equally correlated with body size in either species. MtDNA-based species identification showed that C. glandium larvae were present within acorns of all sizes, whereas C. elephas larvae were restricted to acorns above a minimum size, irrespective of oak species. Hence, exploitation of large acorns can hardly have triggered rostrum enlargement, as the small sized C. glandium adults (with short rostrums) could perforate and oviposit in both small and large acorns. Rather, increased rostrum length is probably a by-product of the larger body sizes of individuals emerging from bigger acorns, which allow increased larval size and enhance larval survival likelihood. Summarizing, when exaggerated feeding traits co-vary with other body features, interspecific morphological variability may result from contrasting selective pressures acting on these correlated characters.     

Endozoochory and Fire as Germination Triggers in Neotropical Dry Forests: an Experimental Test

Endozoochory and fire are crucial ecological factors determining germination success and recruitment in many plant species. Fire is a well‐known germination trigger while endozoochory may allow seed dispersal along with an increase in germination. Their interaction has rarely been addressed, however, even though both factors are pervasive in human‐transformed ecosystems like most Neotropical Dry Forests (NDF). For three common Mesoamerican tree species (Acacia pennatula, Enterolobium cyclocarpum, and Guazuma ulmifolia), we used feeding trials to assess the preference of cattle, which are their main seed dispersal agent. We also experimentally tested the interaction between gut passage and fire as triggers of germination. The fruits of the three species were eaten by cattle, but the small seeds of G. ulmifolia were ingested 10‐fold more than those of the other species. While gut passage did not have any effect on germination, heat‐shocks above 90 °C increased the number of germinating seeds by 15 percent. These results suggest that cattle may be a key dispersal vector in NDF, but that fire may be an important germination trigger. Physical dormancy in these species may have been selected for by extinct megaherbivores because it was a key trait ensuring seed survival after gut passage. However, in light of the recent expansion of cattle‐ranching and fire occurrence in NDF, it has become a useful exaptation facilitating the colonization of disturbed areas.     

Of niche differentiation,dispersal ability and historical legacies: what drives woody community assembly in recent Mediterranean forests?

Community assembly rules have been extensively studied, but its association with regional environmental variation and land use history remains largely unexplored. Land use history might be especially important in Mediterranean forests, considering their historical deforestation and recent afforestation. Using forest inventories and historical (1956) and recent (2000) land cover maps, we explored the following hypotheses: 1) woody species assembly is driven by environmental factors, but also by historical landscape attributes; 2) recent forests exhibit lower woody species richness than pre‐existing due to the existence of colonization credits; 3) these credits are modulated by species’ life‐forms and dispersal mechanisms. We examined the association of forest historical type (pre‐existing versus recent) with total species richness and that of diverse life‐forms and dispersal groups, also considering the effects of current environment and past landscape factors. When accounting for these effects, no significant differences in woody species richness were found between forest historical types except for vertebrate‐dispersed species. Species richness of this group was affected by the interaction of forest historical type with distance to coast and rainfall: vertebrate‐dispersed species richness increased with rainfall and distance to the coast in recent forests, while it was higher in dryer sites in pre‐existing forests. In addition, forest historical types showed differences in woody species composition associated to diverse environmental and past landscape factors. In view of these results we can conclude that: 1) community assembly in terms of species richness is fast enough to exhaust most colonization credit in recent Mediterranean forests except for vertebrate‐dispersed species; 2) for these species, colonization credit is affected by the interplay of forest history and a set of proxies of niche and landscape constraints of species dispersal and establishment; 3) woody species assemblage is mostly shaped by the species’ ecological niches in these forests.     

Regeneration by sprouting of holm-oak (Quercus ilex) stands exploited by selection thinning

Holm-oak forests of the Montseny Massif have traditionally been exploited by selection thinning, which consists of a partial removal of standing trees and biomass. Cutting percentages within each size class indicate that this management technique represents a compromise between forest improvement and harvesting.Sprouting is the mean regeneration process of these holm-oak stands. The mean number of living sprouts per stool is considerably higher than in other coppiced hardwood species, and decreases along the cutting-cycle according to a negative exponential function. Mean number of surviving sprouts per stool is significantly correlated with regenerative age and stool size. Death of sprouts throughout the cutting cycle is compensated by growth of surviving sprouts. Thus, the weight of living sprouts per stool increases linearly with time, and also with the basal area coppiced.Sprouts developing within the first fifteen years of regeneration belong to a single cohort that appears during the first year after cutting. After that period of time, new sprouts appear from dormant buds, so that after 30 years of regeneration a wide range of cohorts of different ages are present.