Consistent pattern of habitat and species selection by post-dispersal seed predators in a Mediterranean mosaic landscape

There is still little information on effects of habitat degradation on post-dispersal seed predation at the landscape scale. The aim of this study was to determine the influence of habitat degradation and seed species on the variability of post-dispersal seed-predation rate. Experimental seed removal was investigated in six Mediterranean woody plant species, four trees (Pinus sylvestris, Quercus ilex, Acer opalus ssp. granatense, and Sorbus aria) and two shrubs (Berberis vulgaris and Crataegus monogyna), in an extensively used mosaic landscape on the Sierra Nevada massif (SE Spain). Seed depots were distributed over 2 years in five differently degraded landscape units, each one with three plots: shrubland; native forest; and dense, cleared and fenced reforestation stands. Predation was the highest in native forest, shrubland, and fenced reforestation, and the lowest in dense and cleared reforestation stands, being partially due to a positive correlation between shrub cover and post-dispersal seed predation. However, the main factors driving post-dispersal seed predation were intrinsic to seeds, as species preference explained most of the variance in our model for predation. The plant-species ranking was Quercus > Pinus > Sorbus > Berberis > Acer > Crataegus, the dominant tree species being the most depredated. These findings are novel because they suggest for the first time that species-selection patterns by post-dispersal seed predators tended to remain constant through both study years in all habitats comprising a mosaic landscape, whether native forest, reforestation stands or successional shrubland.     

Seed Dispersal Patterns by Large Frugivorous Mammals in a Degraded Mosaic Landscape

Seed dispersal by Red fox (Vulpes vulpes), Stone marten (Martes foina), and Wild boar (Sus scrofa) was analyzed in an extensively degraded mosaic landscape in Sierra Nevada (SE Spain). The main objective was to determine whether seed dispersal by mammals was related to habitat degradation within a mosaic of adjacent degraded patches mixed with native forest and thereby to determine the potential role of mammals as seed dispersers in degraded landscape units. For three consecutive years, mammal feces were collected in the fruit production period, extracting all seeds of woody species found therein and analyzing their viability. Feces were collected in three different plots for each of five different landscape units: shrubland, native forest, and dense, cleared, and fenced reforestation stands. Seeds from 16 woody species (which represent more than a half of the total fleshy‐fruited woody species available) were recorded, although some agrarian species are also introduced in a low percentage of the scats. Seeds showed a high viability rate for all dispersed species, irrespective of the mammal disperser. No differences in species composition appeared in the overall landscape units or in the seed density between degraded habitats. Due to the small patch size, the high viability of dispersed seeds, and the large home range of the large mammals, these three animal species act as efficient seed dispersers for a diverse assemblage of woody plant species regardless of the habitat type within this degradation framework. This fact has important consequences for the biodiversity recuperation in these degraded habitats, principally in pine plantations.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Direct and indirect interactions among plants explain counterintuitive positive drought effects on an eastern Mediterranean shrub species

Coexistence of woody and herbaceous plants may be governed by a complex set of direct and indirect interactions, whose relative importance have been rarely assessed. We experimentally studied woody species establishment in a mixed plant community by disentangling the potential role of such biotic interactions and the effect of environmental variations on them. Seedling establishment of the common eastern Mediterranean shrub species Sarcopoterium spinosum was investigated under different rainfall and light conditions, combined with the effect of the presence of adult shrubs and annual neighbors. We predicted that seedlings will be directly affected by competition with annuals with increasing water availability, while direct effects of adult shrubs will be positive via amelioration of water stress. Indirect effects were expected beneath shrub canopies due to reduced water stressed and light availability for both annuals and shrub seedlings, which may intensify competition between annuals and shrub seedlings. To test these predictions we performed field and garden experiments in which we combined manipulation of shrub and annual presence with manipulations of water availability and light conditions to simulate the effect of shrub canopy. In contrast to our prediction, shrub seedling establishment was not facilitated but inhibited by adult shrubs because of light limitation. As expected, annuals had direct negative effects on shrub seedlings under wet conditions, which shifted to neutral or positive effects under dry conditions. Thus, interactions among shrubs and annuals, and in particular the release from competition during drought years, leads to a counterintuitive positive effect of drought on shrub seedling establishment. Our findings point to the importance of experimentally studying multidimensional interactions for coexistence of different life forms and to the underestimated role of light for success in water‐limited ecosystems.     

Resilience of Mediterranean shrubland to a severe drought episode: the role of seed bank and seedling emergence

Extreme climate events, such as severe drought episodes, may induce changes in vegetation if they induce species‐specific adult mortality and changes in the seedling recruitment pattern. In 2005 a severe drought occurred in Doñana National Park (south Spain) causing extensive shrubland mortality. Over the following years we monitored the soil seed bank and seedling emergence via a gradient of canopy dieback induced by the drought episode. The canopy dieback corresponded to an increase in emergence of seedlings of woody species in 2007, probably because of the reduced competition induced by canopy loss. The soil seed bank of woody species sampled in 2008 was less abundant on plots with a higher proportion of dead vegetation, probably because of depletion of the seed bank as a result of the increased germination in the previous year and also as a result of a reduction in seed supply in these sites. Accordingly, in 2009 we detected reduced emergence of woody species on plots that had suffered the greatest shrub mortality. We failed to find any significant changes in patterns of the soil seed bank and seedling emergence of short‐lived herbaceous species, indicating greater resilience in these types of species. This study highlights the resilience of Mediterranean shrublands to climate fluctuations at one extreme of the variability characteristic of these ecosystems. An increase in the frequency of severe drought episodes – increasingly probable under the new climate conditions – does have the potential, however, to induce changes in vegetation, especially in woody communities that need more time to replenish their seed banks.     

Community-Wide Spatial and Temporal Discordances of Seed-Seedling Shadows in a Tropical Rainforest

Several factors decrease plant survival throughout their lifecycles. Among them, seed dispersal limitation may play a major role by resulting in highly aggregated (contagious) seed and seedling distributions entailing increased mortality. The arrival of seeds, furthermore, may not match suitable environments for seed survival and, consequently, for seedling establishment. In this study, we investigated spatio-temporal patterns of seed and seedling distribution in contrasting microhabitats (bamboo and non-bamboo stands) from the Brazilian Atlantic Forest. Spatial distribution patterns, spatial concordance between seed rain and seedling recruitment between subsequent years in two fruiting seasons (2004–2005 and 2007–2009), and the relation between seeds and seedlings with environmental factors were examined within a spatially-explicit framework. Density and species richness of both seeds and seedlings were randomly distributed in non-bamboo stands, but showed significant clustering in bamboo stands. Seed and seedling distributions showed across-year inconsistency, suggesting a marked spatial decoupling of the seed and seedling stages. Generalized linear mixed effects models indicated that only seed density and seed species richness differed between stand types while accounting for variation in soil characteristics. Our analyses provide evidence of marked recruitment limitation as a result of the interplay between biotic and abiotic factors. Because bamboo stands promote heterogeneity in the forest, they are important components of the landscape. However, at high densities, bamboos may limit recruitment for the plant community by imposing marked discordances of seed arrival and early seedling recruitment.     

Critical stages in the recruitment process of Rhamnus alaternus L

BACKGROUND AND AIMS: Rhamnus alaternus is a Mediterranean shrub commonly used in reforestation programs. Although several aspects of its reproductive biology have been studied, little is known about the importance of the different recruitment stages in the overall regeneration process of this species, which limits its proper use in Mediterranean forests and shrubland management. The aim of the present work was to quantify the importance of the different recruitment stages in the regeneration process of R. alaternus. METHODS: Two populations of Rhamnus alaternus on the island of Mallorca that differ in climatic conditions, type of habitat and sex ratio were studied. The importance of seed production, seed dispersal and predation, seedling emergence and seedling survival for the regeneration of this species were quantified. KEY RESULTS: In both populations, fruit set and fruit removal by animals were not critical stages, since almost half of the flowers became mature fruits and 90% of those were dispersed. Most seeds were deposited under female conspecifics (86 and 47%, at Lloret and Esporles, respectively), and very few were found in open inter-spaces (1 and 5%). Post-dispersal seed predation (mostly by ants and rodents) was very high in both populations. Seedling emergence took place during autumn and early winter and it ranged from 31 to 68% depending upon year and microhabitat. The majority of emerged seedlings died during the first year, mainly due to desiccation; such mortality was influenced by rainfall and differed among microhabitats (varying from 67 to 100%). The general spatial distribution of seed rain was concordant with the seedling emergence and survival pattern in both populations. CONCLUSIONS: The recruitment of Rhamnus alaternus appeared to be mainly limited by seed and seedling survival, regardless of the type of habitat in which the species is found.     

Seed Dispersers,Seed Predators,and Browsers Act Synergistically as Biotic Filters in a Mosaic Landscape

In this study, we analize the functional influence of animals on the plants they interact with in a mediterranean mountain. We hypothesise that seed dispersers, seed predators, and browsers can act as biotic filters for plant communities. We analyse the combined effects of mutualistic (seed dispersal) and antagonistic (seed predation, herbivory) animal interactions in a mosaic landscape of Mediterranean mountains, basing our results on observational and experimental field. Most of the dispersed seeds came from tree species, whereas the population of saplings was composed predominantly of zoochorous shrub species. Seed predators preferentially consumed seeds from tree species, whereas seeds from the dominant fleshy-fruited shrubs had a higher probability of escaping these predators. The same pattern was repeated among the different landscape units by browsers, since they browsed selectively and far more intensely on tree-species saplings than on the surrounding shrubs. In synthesis, our work identifies the major biotic processes that appear to be favoring a community dominated by shrubs versus trees because seed dispersers, predators, and herbivores together favored shrub dispersal and establishment versus trees.     

The effect of plant litter on ecosystem properties in a Mediterranean semi‐arid shrubland

Abstract. We studied the distribution of litter in a shrubland of the Negev with a semi‐arid Mediterranean climate of less than 200 mm of rainfall per year. Our focus was on the effects of litter on properties of landscape patches relevant to ecosystem processes (water runoff and soil erosion), annual plant community responses (seedling density, biomass production and species richness), and animal activity (soil disturbance by termites). Three 60‐m transects, extending across a pair of opposing north‐ and south‐facing slopes and their drainage channel, showed that litter accumulates not only under shrubs, but to a lesser extent also on the crusted inter‐shrub open areas. We used 35 experimental units (‘cells’, 0.5m × 1 m), each containing a crust and a shrub patch. Because runoff flows from crusted patches and is intercepted by shrub patches, the latter were in the lower third of the cells. Leaf litter was added in single and double amounts providing ca. 0.5 and 1.0 cm litter depth, to either, both, or none of the patches. Litter addition significantly decreased the amount of runoff, regardless of the location and amount of litter applied. Litter on the crust increased species number and seedling density of species with low abundance. Adding a double litter layer increased annual plant biomass production, while a single amount had no effect. Litter addition to the shrub patch affected neither biomass nor species richness. Litter addition to both patches at both quantities caused a large increase in termite activity. Termites caused disturbance by disrupting the crust, which may contribute to the reduction in runoff amounts. In the open, flat crust patches, annual plant communities are limited in their productivity and species richness, as there are few structures stopping the outflow of water, soil and seeds. Litter adds such structures, but affects the plant communities only when added to litter‐free crust. Litter accumulation and its patchy distribution have large impacts on landscape patch properties affecting resource distribution, plant productivity and diversity, and animal activity. Therefore, understanding litter distribution in relation to the patchy structure of the landscape of semi‐arid shrubland should be viewed as an important component of shrubland management.     

Climatic warming strengthens a positive feedback between alpine shrubs and fire

Climate change is expected to increase fire activity and woody plant encroachment in arctic and alpine landscapes. However, the extent to which these increases interact to affect the structure, function and composition of alpine ecosystems is largely unknown. Here we use field surveys and experimental manipulations to examine how warming and fire affect recruitment, seedling growth and seedling survival in four dominant Australian alpine shrubs. We found that fire increased establishment of shrub seedlings by as much as 33‐fold. Experimental warming also doubled growth rates of tall shrub seedlings and could potentially increase their survival. By contrast, warming had no effect on shrub recruitment, postfire tussock regeneration, or how tussock grass affected shrub seedling growth and survival. These findings indicate that warming, coupled with more frequent or severe fires, will likely result in an increase in the cover and abundance of evergreen shrubs. Given that shrubs are one of the most flammable components in alpine and tundra environments, warming is likely to strengthen an existing feedback between woody species abundance and fire in these ecosystems.     

Shrub encroachment in Argentinean savannas

Abstract. In the Wet Chaco region of Argentina, increasing shrub encroachment in savannas over the last few decades has led to important changes in the structure and functioning of the landscape. Some sectors of this territory are characterized by the appearance of circular clusters of woody patches, dispersed throughout the grassland matrix. The increasing size of these patches leads to a gradual change from grassland to dense shrubland. We studied these circular woody patches in the eastern region of the Argentine province of Formosa and characterized the variation in terms of floristic composition, diversity and predominant seed dispersal mode in different size patches. We observed an increase in species richness, diversity and compositional heterogeneity among patches with increasing patch size. Seed dispersal by animals, especially birds, is an important factor in the expansion of these woody vegetation patches within the grassland matrix.     

Evaluating Dispersal Limitation in Passive Bottomland Forest Restoration

Dispersal limitation can retard natural establishment of desirable species on restoration sites, especially where landscapes are fragmented, but dispersal limitation is assumed to become less critical with time as early colonists become reproductively mature. Distribution patterns of recruiting trees in a 20‐year‐old passively restored bottomland in northeast Louisiana suggested persistent dispersal limitation in some bottomland hardwood species and influence of dense shrub patches on colonization. To test these hypotheses, we measured seed rain as a function of distance to seed source and association with shrub cover. Seed rain of the wind‐dispersed Fraxinus pennsylvanica was highest near the forest edge, except where mature recruits occurred. Although shrub presence did not influence dispersal of F. pennsylvanica, its negative influence on probability of occurrence in the sapling layer suggests that shrub cover may limit its regeneration. The bird‐dispersed Crataegus viridis and Ilex decidua were found in the seed rain and as reproductive individuals within the field; neither had a positive relationship with shrub presence. Dispersal of heavy‐seeded Quercus spp. and Carya aquatica was limited to within 20 m of the forest edge. These results imply that dispersal limitation is diminishing in wind‐ and bird‐dispersed species with maturation of in‐field recruits and that shrub patches may influence these patterns. Heavy‐seeded species, however, remain restricted to field edges that directly abut a seed source. If canopy closure by wind‐ and bird‐dispersed species precedes dispersal of heavy‐seeded species into the field, establishment of Quercus and Carya spp. may remain low for the foreseeable future.     

Recovery of rain forest soil seed banks under different reforestation pathways in eastern Australia

Summary Seed availability is a major factor limiting the recruitment of rain forest to cleared land, but little is known about the composition of the soil seed bank under different reforestation pathways. We quantified changes in the viable soil seed bank following rain forest clearing and pasture establishment and subsequent reforestation in subtropical eastern Australia. Major reforestation pathways in the region include planting of a diverse suite of native trees for ecological restoration purposes, autogenic regrowth dominated by the non‐native tree Camphor Laurel (Cinnamomum camphora) and management of this regrowth to accelerate the development of a native tree community. These pathways differ considerably in cost: restoration plantings are expensive, autogenic regrowth is free, whilst managing regrowth generally costs much less than restoration plantings. We surveyed five sites within each of three reforestation pathways as well as reference sites in remnant rain forest and pasture. The composition of the seed bank was determined by germinating plants from soil samples collected from each site. Germinants were classified into several functional groups according to life form, origin, dispersal mode and successional stage. The majority of functional groups varied significantly in abundance or richness between rain forest and pasture sites. Most of the functional groups that varied between rain forest and pasture were restored to values similar to rain forest by at least one of the three reforestation pathways examined. The species richness of native woody plants in the soil seed bank was slightly higher in restoration plantings than in autogenic or managed regrowth; nevertheless, the species richness and abundance of native woody plants and vines were higher in the seed bank of autogenic regrowth than pasture, and both attributes were enhanced by the management of regrowth sites. The results of this study show that autogenic regrowth can make an important contribution to rain forest restoration at a landscape scale. The optimal reforestation approach or mix of approaches will depend on the desired rate of recovery and the resources available for restoration.     

Are plant populations seed limited? A critique and meta-analysis of seed addition experiments

We examine the relative importance of processes that underlie plant population abundance and distribution. Two opposing views dominate the field. One posits that the ability to establish at a site is determined by the availability of suitable microsites (establishment limitation), while the second asserts that recruitment is limited by the availability of seeds (seed limitation). An underlying problem is that establishment and seed limitation are typically viewed as mutually exclusive. We conducted a meta-analysis of seed addition experiments to assess the relative strength of establishment and seed limitation to seedling recruitment. We asked (1) To what degree are populations seed and establishment limited? (2) Under what conditions (e.g., habitats and life-history traits) are species more or less limited by each? (3) How can seed addition studies be better designed to enhance our understanding of plant recruitment? We found that, in keeping with previous studies, most species are seed limited. However, the effects of seed addition are typically small, and most added seeds fail to recruit to the seedling stage. As a result, establishment limitation is stronger than seed limitation. Seed limitation was greater for large-seeded species, species in disturbed microsites, and species with relatively short-lived seed banks. Most seed addition experiments cannot assess the relationship between number of seeds added and number of subsequent recruits. This shortcoming can be overcome by increasing the number and range of seed addition treatments.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

The role of the soil seed and seedling bank in the regeneration of diverse plant communities in the subtropical Ailao Mountains,Southwest China

We compared species composition and diversity of the soil seed and seedling banks in three secondary vegetation types (shrubland, Populus bonatii forest, Lithocarpus regrowth forest) and a primary old-growth forest in the subtropical Ailao Mountains of southwestern China to clarify the importance of seed and seedling banks for forest dynamics. The average species richness was the highest in soil samples from the shrubland (26.80 ± 1.98), and the lowest from the primary forest (9.93 ± 0.50). The density of germinable tree seeds increased from the secondary vegetation to the primary forest, and the density of shrub, forb, and graminoid seeds decreased significantly. The most abundant seedlings recorded in soil samples were light-demanding species in the shrubland and Populus bonatii forest. For ground flora, the number of shrub seedlings strongly decreased with the increase in stand age, and shade-tolerant tree seedlings tended to increase. The species similarity between the seed bank and the aboveground vegetation in all sites was low (Sørensen’s index = 0.11–0.33), however, the shrubland had higher similarity compared with the other three plant communities. In the primary forest, light-demanding woody species dominated in soil seed banks, while shade-tolerant species dominated in the overstory and the forest floor. In the primary forest, seedlings of dominant tree species were rare in the understory, and no seeds of the dominant species were found in the soil. Results indicated that the early stages of vegetation recovery should take into account the possibility of recovering soil seed bank processes. However, colonization and establishment of tree seedlings will be difficult once a primary forest is destroyed.     

How landscape modulators function: woody plant impact on seed dispersal and abiotic filtering

The Mediterranean landscape is characterized by a heterogeneous structure: a mosaic of woody plants (trees or shrubs) with scattered patches of herbaceous vegetation. Although the herbaceous and woody patches are adjacent to each other, plant species composition in them is substantially different. This could be attributed to either differences in environmental conditions between patch types (i.e., abiotic filters), or to dispersal limitations caused by the woody plants acting as dispersal filters. In this article, we focus on the relative impact of woody plants, applying these two filter types, in determining plant species composition in Mediterranean woodland. We experimentally manipulated shade and litter cover and examined the effect of each of these factors on plant species composition. We used seed-traps to evaluate seed arrival in the patches, and experimentally removed the shrub canopy to study the effect of the shrub as a physical barrier to seed entry. Results showed that plant species number and composition were not significantly affected by shade and litter manipulation. The number of trapped seeds were significantly higher in the open patches than in the woody patches, and removal of woody plants increased the number of trapped seeds in both open and woody patches, as a result of eliminating the physical obstacle to free seed movement. Our findings show that woody plants affect the herbaceous plant community by influencing seed dispersal, and highlight that they affect other organisms not only by modifying resource availability but also through the creation of a new landscape structure.     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

Multiple Recruitment Limitation Causes Arrested Succession in Mediterranean Cork Oak Systems

Lack of tree regeneration and persistency of species-poor shrublands represent a growing problem across Mediterranean evergreen oak forests. What constrains forest regeneration is poorly understood, and restoration attempts have been largely unsuccessful. We assessed the contribution of four different mechanisms of tree recruitment limitation (that is, source, dispersal, germination, and establishment) in a cork oak (Quercus suber) system in southern Portugal. Using a combination of field studies and experiments, we quantified seed production, seed removal and dispersal, seed survival and germination, seedling establishment and survival, as well as cork oak natural regeneration for the three dominant vegetation types in this system (Cistus ladanifer shrubland, oak forest, and oak savanna). We found that all four forms of cork oak recruitment limitation were significantly more severe in shrublands than in oak forests and savannas, so that oak seedling recruitment in shrubland was impeded in multiple ways. Our results explain why transitions from shrublands to oak savannas and forests are extremely difficult, and that the release from arrested succession in this system requires the simultaneous relief of multiple constraints on recruitment limitation in the early life history of oaks. These results have important implications for the restoration and conservation of Mediterranean oak systems.     

Restoration of Araucaria Forest: The Role of Perches, Pioneer Vegetation, and Soil Fertility

The Araucaria forest is Brazil's highly threatened subtropical forest ecosystem that has been disappearing in recent decades. Restoration programs involving this forest type are scarce, and there is a lack of scientific information on how ecological processes such as competition, facilitation, and seed dispersal influence natural forest restoration. This work aims to investigate how use of perches to attract seed dispersers and the influence of pioneer vegetation and soil fertilization could affect the colonization of woody species in a degraded area. An experiment was conducted in an abandoned field where the natural establishment of seeds and seedlings of woody species was monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control); and (3) perches (presence and absence). Seed and seedling abundance, seed and seedling species richness, and seedling mortality were recorded monthly during 12 months. Seed abundance and species richness were significantly greater in places with perches than in control plots. These results were consistent over the year and more pronounced when the surrounding forest produced a higher amount of fruit. Species richness and abundance of seedlings were significantly greater in places with perches than in control plots, and in places with vegetation than without. Soil fertility did not influence seedling establishment. Facilitation and seed dispersal are important factors affecting the colonization of woody species in this subtropical area. Nutrient availability neither regulates the facilitation process nor influences species replacement during the early stages of Araucaria forest succession.