Mechanisms of interaction between a leguminous shrub and its understorey in a semi-arid environment

Retama sphaerocarpa shrubs in semi-and environments often have a dense understorey of annual and perennial herbs forming so-called "islands of fertility" The effect of the canopy on soil fertility and microclimate and the combined effect of canopy and litter on species diversity and productivity were assessed under Retama spliaerocarpa shrubs in a semi-and environment in southeast Spain Soil chemical properties differed significantly among three positions under the canopy, particularly between inner and outer positions The potential mineralization rate of organic matter was significantly higher in soils from an intermediate position under the canopy than in soils from either the centre and the edge Soil chemical fertility and estimated soil seed bank were highest also in soil at an intermediate position and lowest in soil from the edge The understorey flora was favoured by the lower temperature and irradiation and increased soil fertility under the shrubs canopies Species emerging from the soil seed bank separated clearly into two groups which occupied inner and outer positions under the canopy Different levels of addition of Retama litter significantly decreased species richness and the number of emerged seedlings In the field, pots placed near the centre, at an intermediate position and at the edge of the canopy of Retama shrubs differed significantly in species richness and biomass production Overall, micro-climatic conditions combined with increased chemical fertility of the soil and inhibitory effects of litter to produce a large biomass of herbs at intermediate positions between the centre and the edge of the canopy. The high proportion of litter from annual species in that position increases the mineralization rate and hence nutrient dynamics in a process which also could benefit the shrub     

Effects of Prosopis flexuosa on soil properties and the spatial pattern of understorey species in arid Argentina

Abstract. In arid zones dominant woody plants are capable of causing changes in microclimate and soil properties likely to affect species composition, as well as the establishment and spatial distribution of plant species. In North American and European deserts species richness appears to be higher under the canopy of shrubs and trees, in contrast with Chilean deserts where it seems to be lower. Since Prosopis flexuosa (Fabaceae, Mimosoideae) is the most conspicuous tree in the central Monte desert, Argentina, we analysed the effect of this species on the composition and abundance of the shrub and herbaceous layers and on soil properties. We considered two mesohabitats: ‘under P. flexuosa canopy’ and ‘intercanopy areas’. In addition, we analysed the differences between two microhabitats under canopies: ‘northern part of the canopy’ and ‘southern part of the canopy’. Results indicate that species composition and soil properties are affected by both mesohabitats and microhabitats. We found a higher number of shrubs under canopies, whereas that of grasses and perennial forbs increased in intercanopy areas. Concentrations of organic matter, nitrogen, potassium and phosphorus, factors limiting biological productivity in Monte desert soils, were significantly higher under than outside P. flexuosa canopies. Electrical conductivity and concentrations of Na⁺, Ca⁺⁺, Mg⁺⁺ were higher in the northern than in the southern microhabitats. No differences in species richness, evenness or diversity were found between mesohabitats or between microhabitats. We conclude that P. flexuosa modifies the spatial pattern of plant species in the shrub and herbaceous layers and the chemical conditions of the soil, generating spatial heterogeneity on different scales.     

Spatial segregation of plant species caused by Retama sphaerocarpa influence in a Mediterranean pasture: a perspective from the soil seed bank

Retama sphaerocarpa is a leguminous shrub whose important role in the semi-arid regions of south-eastern Spain has already been assessed: shrub canopies reduce light intensity, and thus evapotranspiration; also their extensive radical system take water and nutrients from great volumes of soil, concentrating them in the understorey. Consequently, subcanopy vegetation benefits from these facilitation processes, increasing its productivity. However, these shrublands have been rarely studied at a broader scale, i.e. as a savannah-like system composed of a variable number of shrubs scattered in a herbaceous matrix. As the microenvironmental conditions associated to the understorey are rather different from those of the open spaces among shrubs, species composition of the herbaceous matrix is expected to change accordingly. Thus, R. sphaerocarpa would be an important and still unknown source of spatial heterogeneity to the system. Our main purpose was to evaluate, through the soil seed bank, the heterogeneity in the herbaceous community induced by this shrub species. Soil samples were collected around adult shrubs from three positions relative to the canopy: near the centre of the shrubs, at the edge of the understorey, and completely outside the canopy. Floristic composition was evaluated by germination under greenhouse conditions. The results show that each position has a different floristic composition, characterised by a group of different species. The herbaceous species associated with the external position have functional traits which enable them to resist water stress and herbivore pressure, such as hairs, CAM metabolism, early flowering, horizontal growth or tiny stature. The species associated with the central position lack those traits, and are more competitive in more mesic environments, rich in nitrogen. The lowest number of seedlings and species was found in the internal position, suggesting that in our study the facilitation process may have less importance for community dynamics due to less stressful environmental conditions.     

The influence of Prosopis canopies on understorey vegetation: Effects of landscape position

Abstract. The influence of canopy trees and shrubs on under‐storey plants is complex and context‐dependent. Canopy plants can exert positive, negative or neutral effects on production, composition and diversity of understorey plant communities, depending on local environmental conditions and position in the landscape. We studied the influence of Prosopis velutina (mesquite) on soil moisture and nitrogen availability, and understorey vegetation along a topographic gradient in the Sonoran Desert. We found significant increases in both soil moisture and N along the gradient from desert to riparian zone. In addition, P. velutina canopies had positive effects, relative to open areas, on soil moisture in the desert, and soil N in both desert and intermediate terrace. Biomass of understorey vegetation was highest and species richness was lowest in the riparian zone. Canopies had a positive effect on biomass in both desert and terrace, and a negative effect on species richness in the terrace. The effect of the canopy depended on landscape position, with desert canopies more strongly influencing soil moisture and biomass and terrace canopies more strongly influencing soil N and species richness. Individual species distributions suggested interspecific variation in response to water‐ vs. N‐availability; they strongly influence species composition at both patch and landscape position levels.     

Size matters: nest colonization patterns for twig‐nesting ants

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Dominance and species co-occurrence in highly diverse ant communities: a test of the interstitial hypothesis and discovery of a three-tiered competition cascade

The role of competitive exclusion is problematic in highly diverse ant communities where exceptional species richness occurs in the face of exceptionally high levels of behavioural dominance. A possible non-niche–based explanation is that the abundance of behaviourally dominant ants is highly patchy at fine spatial scales, and subordinate species act as insinuators by preferentially occupying these gaps—we refer to this as the interstitial hypothesis. To test this hypothesis, we examined fine-scale patterns of ant abundance and richness according to a three-tiered competition hierarchy (dominants, subdominants and subordinates) in an Australian tropical savanna using pitfall traps spaced at 2 m intervals. Despite the presence of gaps in the fine-scale abundance of individual species, the combined abundance of dominant ants (species of Iridomyrmex, Papyirus and Oecophylla) was relatively uniform. There was therefore little or no opportunity for subordinate species to preferentially occupy gaps in the foraging ranges of dominant species, and we found no relationship between the abundance of dominant ants and nondominant species richness at fine spatial scales. However, we found a negative relationship between subdominant and subordinate ants, a negative relationship between dominant and subdominant ants, and a positive relationship between dominant and subordinate ants. These results suggest that dominant species actually promote species richness by neutralizing the effects of subdominant species on subordinate species. Such indirect interactions have very close parallels with three-tiered trophic cascades in food webs, and we propose a “competition cascade” where the interactions are through a competition rather than trophic hierarchy.     

A canopy dominant ant affects twig‐nesting ant assembly in coffee agroecosystems

One commonly studied driver of community assembly is the effect of dominant species on subordinate species. Dominant species may impact community assembly during competitive sorting, or recruitment. For ants, important and abundant species in the tropics, several factors may drive community assembly including competition, dispersal, priority effects, and environmental conditions. Although competition is a hallmark of ant ecology, few have examined the influence of patchily distributed dominant ants on other ant species and diversity, especially at the recruitment stage. Here, I consider the impacts of a canopy dominant ant species, Azteca instabilis, and changes in vegetation on twig‐nesting ant colony founding and ant community assembly in a coffee agroecosystem. I added artificial nests to coffee plants in areas with and without A. instabilis four times over a year, and then examined the occupation rate and identity of species colonizing nests. I also examined vegetation characteristics of sites where nests were added. The presence of A. instabilis on coffee plants drastically lowered colonization rates, but nest occupation increased with tree density, and with decreasing proportion of Inga spp. trees in the canopy. The presence of A. instabilis limited the number of nests occupied by six of the ten most common species; most rare species, however, were not affected by A. instabilis presence. Richness of colonizing ants in areas with A. instabilis was lower, but these effects did not significantly affect richness across broader scales. Despite large effects on individual species, species composition did not differ greatly in areas with and without A. instabilis, but some vegetation characteristics (basal area and tree richness) were predictive of ant composition. These results suggest that A. instabilis strongly affects founding events especially for common twig‐nesting species and that both vegetation and influences from this dominant species affect community assembly of twig‐nesting ants at the local scale.     

Canopy effects of the invasive shrub Pyracantha angustifolia on seed bank composition,richness and density in a montane shrubland (Córdoba,Argentina)

Abstract Invasive woody species frequently change the composition of the established vegetation and the properties of the soil under their canopies. Accordingly, invasion may well affect regenerative phases of the community, especially at the seed bank level, likely influencing community restoration. Pyracantha angustifolia (Rosaceae) is an invasive shrub in central Argentina that affects woody recruitment, particularly enhancing the recruitment of other exotic woody species. There is though no information regarding its effect on the soil seed bank within the invaded community. The present study was set up to gain further insight into the canopy effects of P. angustifolia. We aimed to assess whether the invasive shrub affects seed bank composition, richness and seed density as compared with the dominant native shrub Condalia montana (Rhamnaceae), and to relate the observed seed bank patterns with those of the established vegetation. We evaluated the composition of the germinable seed bank and the established vegetation under the canopy of 16 shrubs of P. angustifolia, 16 shrubs of C. montana, and in 16 control plots (10 m²) without shrub cover. The floristic composition of the seed bank differed among canopy treatments. However, seed bank richness did not differ significantly. There was an overall high seed density of exotic species throughout the study site, though exotic forbs showed significantly lower seed densities under the invasive shrub. Pyracantha angustifolia would not promote the incorporation of new species into the seed bank of the invaded community but rather favour the establishment of woody species that do not depend on seed banks. The absence of dominant woody species in the seed bank, the dominance of exotic forbs, and the high similarity between established exotic species and those present in the seed bank may surely affect community restoration following the main disturbances events observed in the region.     

Impacts of the invasive exotic <Emphasis Type="Italic">Prosopis juliflora</Emphasis> (Sw.) D.C. on the native flora and soils of the UAE

The effects of the invasive exotic Prosopis juliflora shrubs on the natural plant communities and soil chemical characters were assessed in two regions of the United Arab Emirates (UAE). Five sites were selected subjectively: three in Sharja with 73 stands and two in Ras Al-Khima with 37 stands. Stands were located randomly within each site to cover density variation in Sharja and size variation in Ras Al-Khima. Density, frequency, richness and evenness of the associated annual and perennial species were studied in nine quadrats distributed under, at the margin and outside the canopy of a P. juliflora shrub located in the center of each stand. The results indicated that the effect of P. juliflora on the associated flora depends significantly on the density and size of the canopy. Larger individuals and greater densities have significantly greater negative impacts on the associated plants. All the studied community attributes were significantly lower under P. juliflora canopies than outside. Annuals were inhibited more than perennials. The number of annuals with significant reductions in density and/or frequency under P. juliflora canopies was significantly greater than the number of perennials. Density of more than 50% of the associated annuals was significantly inhibited under P. juliflora canopies. Density of P. juliflora seedlings was greater underneath the canopy of the same species than away from them, indicating little or no self allelopathic effect (auto-inhibition) under field conditions. P. juliflora ameliorated some soil characters, through significant pH reduction and increase in K, N and P and organic matters. Hence, plant diversity might be enhanced following eradication of P. juliflora.     

The influence of resprouting forest canopy species on richness in Southern Cape forests, South Africa

We investigated the relationship between species richness and numbers and types of individuals and species present in forests with different physiognomies in the southern Cape Province, South Africa. Data were collected from three different ‘plot’ types: 400 m², canopy‐scaled (plot length is directly proportional to canopy height) and per 100 individuals closest to a point. Plots were designed to control for the effect of scale on local richness. Canopy species richness was inversely proportional to the abundance of resprouting species. The strength of the relationship between the abundance of resprouters and canopy species richness increased progressively from the 400 m² plots to the canopy‐scaled plots and finally to the plots of 100 individuals. Resprouter abundance decreased, while canopy species richness increased, with increasing canopy height. Resprouters are able to retain their in situ position in the forests for longer periods of time than do reseeders. This reduces individual and species turnover, thus reducing species richness in resprouter‐dominated forests.     

Extreme drought stress shifts net facilitation to neutral interactions between shrubs and sub‐canopy plants in an arid desert

The stress gradient hypothesis (SGH) predicts that the importance or intensity of competition and facilitation will change inversely along abiotic stress gradients. It was originally postulated that increasing environmental stress can induce a monotonic increase in facilitation. However, more recent models predicted that the relationship between severity and interaction exhibits a hump‐shaped pattern, in which positive interactions prevail under moderate stress but decline at the extreme ends of stress gradients. In the present study, we conducted a field experiment along a temporal rainfall gradient for five consecutive years, in order to investigate interactions in a shrub‐herbaceous plant community at the southern edge of the Badain Jaran Desert, and, more specifically, investigated the effects of Calligonum mongolicum, a dominant shrub species, on both abiotic environmental variables and the performance of sub‐canopy plant species. We found that shrubs can improve sub‐canopy water regimes, soil properties, plant biomass, density, cover, and richness and, more importantly, that the positive effect of shrubs on sub‐canopy soil moisture during the summer diminishes as rainfall decreases, a pattern that partly explains the collapse of the positive interaction between shrubs and their understory plants. These results provide empirical evidence that the positive effect of shrubs on understory plant communities in extreme arid environments may decline and become neutral with increasing drought stress.     

Contribution of cocoa plantations to the conservation of native ants (Insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic Forest fauna of southern Bahia,Brazil

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests. Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.     

Spatial distribution of soil nutrients and ephemeral plants underneath and outside the canopy of Porlieria chilensis shrubs (Zygophyllaceae) in arid coastal Chile

Soil nutrients and density and biomass of annual plants underneath and outside the canopy of Porlieria chilensis shrubs were measured at the end of the growing season in a protected arid coastal site in Chile. Levels of soil nitrogen, phosphorus and organic matter were significantly higher underneath than outside the canopies of shrubs. Almost 4 times as many plants occurred outside than underneath shrubs, but no significant differences in total aboveground biomass were found. Several species had higher densities and/or biomass outside rather than underneath shrubs, whereas others showed the oppsite trend. Species richness was lower underneath P. chilensis canopy. The spatial microdistribution of ephemeral species may be explained by differential water and nutrient requirements. Comparison of the patterns observed in our protected site versus surrounding unprotected areas supports the generalization that man, by removing shrubs and trees, has changed a previous heterogeneous spatial distribution of nutrients to a more homogenous one.This is a contribution of the Program of Arid Zone Studies of the Universidad de la Serena     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.     

Remnant shrubs in Mediterranean semi-arid steppes: effects of shrub size, abiotic factors and species identity on understorey richness and occurrence

Isolated shrub patches are a key element for community structure and dynamics in semi-arid ecosystems, and may act as “hot spots” of understorey species diversity. Despite its importance, the relationships between shrub patch characteristics and understorey species richness, and the relative importance of these characteristics against other factors driving understorey species richness, are still poorly understood. We studied perennial species richness under the canopy of late-successional shrubs in semi-arid Stipa tenacissima steppes of SE Spain. The main aim of this study was to evaluate the relationship between shrub size and understorey richness, and to test if this relationship is modified by species identity. We also aimed to know the relative importance of shrub patch characteristics and abiotic factors as predictors of the distribution of understorey species under shrub canopies. Altitude and geographical co-ordinates were able to significantly explain the patterns of shrub occurrence and abundance in the study area. Understorey species richness was significantly related to the size of individual shrubs according to a simple power relationship for all evaluated species. Slope values of linear regressions with log-transformed data, ranging between 0.22 and 0.37, did not differ between species. Site characteristics, patch characteristics and patch species identity significantly explained the occurrence of species on patches. However, the relative importance of site characteristics was higher than that of patch characteristics and species identity in explaining these patterns. Our results agree with the general expectations of the theory of island biogeography and complement previous studies that emphasise the importance of late-successional shrubs in semi-arid Stipa steppes. Environmental management activities within these steppes should promote the conservation of remnant shrubs, as well as its introduction with restoration activities.     

Spatial and temporal variation in islands of fertility in the Sonoran Desert

In many arid and semi-arid ecosystems, canopy trees and shrubs have a strong positive influence on soil moisture and nutrient availability, creating islands of fertility where organic matter and nutrients are high relative to areas outside the canopy. Previous studies of canopy effects on soil processes have rarely considered how landscape context may modulate these effects. We measured the effects of velvet mesquite trees (Prosopis velutina) on soil moisture and the biogeochemistry of nitrogen at different positions along a topographic gradient from upland desert to riparian zone in the Sonoran Desert of central Arizona. We also examined how landscape position and patterns of precipitation interact to determine the influence of P. velutina on soil moisture, N availability assessed using ion exchange resins, net N mineralization and net nitrification, and microbial biomass C and N. P. velutina clearly created islands of fertility with higher soil organic matter, net N mineralization and net nitrification rates, and microbial biomass under mesquite canopies. These effects were consistent across the landscape and showed little temporal variability. Magnitude and direction of effect of mesquite on soil moisture changed with landscape position, from positive in the upland to negative in the terrace, but only when soil moisture was >4%. Resin N showed responses to mesquite that depended on precipitation and topographic position, with highest values during wet seasons and under mesquite on terraces. We suggest changes in proximity of P. velutina to groundwater lead to shifts in biogeochemical processes and species interactions with change in landscape position along a topographic gradient.     

Species Identity and Water Availability Determine Establishment Success Under the Canopy of Retama sphaerocarpa Shrubs in a Dry Environment

Seedling establishment in harsh environments is often enhanced by the proximity of adult shrubs. This information has been used in restoration work by placing seedlings of species being restored under the canopy of some shrubs. However, monitoring this process is often restricted to a single species, and comparisons with practices that protect planted seedlings against harsh conditions are scant. Similarly, few studies have supplied seedlings with water in the summer to observe the effects of water availability on the interaction. We compared sapling survival of three woody species (Olea europaea, Pistacia lentiscus, and Ziziphus lotus) under the leguminous shrub Retama sphaerocarpa and in gaps covered with piled branches that mimicked a shrub canopy. After 3 years, survival of saplings planted under Retama differed depending on species identity and water supply. Survival of Olea saplings placed under Retama shrubs was twice that under piled branches if not watered (35 ± 8 vs. 17 ± 2 %, respectively), whereas survival of saplings under Retama, if watered, was less (48 ± 11 vs. 68 ± 8%, respectively). Retama shrubs had a negative effect on Ziziphus; most saplings died under its canopy, whereas survival in piled branches ranged 10–54%. Pistacia was neither facilitated nor outcompeted by Retama. Facilitation of Olea by Retama shrubs was more apparent under dry conditions where watering increased competition and decreased facilitation. Overall, we conclude that Retama shrubs can help dry land restoration to a greater extent than artificial shade for Olea when not watered. The critical role played by water supply in determining nursing success rates warrants further study.     

Shrubs as a source of spatial heterogeneity—the case of Retama sphaerocarpa in Mediterranean pastures of central Spain

The processes underlying the positive effect exerted by woody species on other plant species can be grouped into a physical effect due to the canopy itself (mainly amelioration of temperature and light extremes), and an edaphic effect (nutrient enrichment). Both groups of effects are present in the savanna-like systems originated by Retama sphaerocarpa in the Iberian Peninsula. The consequences of these shrubs for the spatial heterogeneity induced in the herbaceous community are not well known. The herbaceous community was sampled by means of radial transects around 20 adult shrubs, from the canopy centre to the open areas among the shrubs, both northwards and southwards. Floristic composition, diversity and pasture mean height change both along the inside–outside gradient, and from north to south. As a nutrient enriched environment, the centre of the understorey showed the lowest species richness and evenness, and the highest standing crop. In contrast, the environmentally stressful open areas showed the highest number of species and evenness, and the lowest standing crop. This general effect is only partially due to nutrient availability, and is not homogeneous around the shrub, as open area conditions penetrate into the understorey driving the herbaceous community towards that of open areas. An individual shrub effect is also suggested by our results. Different conditions are thus provided by R. sphaerocarpa understoreys, favouring different subsets of species from the general pool, and therefore defining shrub canopies as sources of spatial heterogeneity in the whole savanna-like system.     

Benefits of Using Shrubs as Nurse Plants for Reforestation in Mediterranean Mountains: A 4-Year Study

Shrubs are commonly considered competitors of planted seedlings in reforestation programs. However, shrubs can facilitate the establishment of understory seedlings in environments that, like Mediterranean‐type ecosystems, are characterized by harsh environmental conditions. In 1997, an experiment was set up in the Sierra Nevada mountains (southeast Spain) to test the use of shrubs as nurse plants for an alternative reforestation technique. Two‐year‐old seedlings of Pinus sylvestris and Pinus nigra were planted in four microhabitats: (1) open interspaces without vegetation (which is the usual method employed in reforestation programs), (2) under individuals of the shrub Salvia lavandulifolia, (3) under the north side of spiny shrubs, and (4) under the south side of spiny shrubs. Seedlings were also distributed in plots with and without ungulates to test the effect of herbivore damage. We report here the results of survival and growth after four growing seasons, a time span long enough to draw robust conclusions concerning the suitability of this technique. Pine survival was remarkably higher when planted under individuals of S. lavandulifolia as compared with open areas (2.6 times for P. sylvestris and 1.8 for P. nigra). The survival of both pine species was also higher when planted on the north side of spiny shrubs, while mortality on the south side was similar to that found in open areas. The reduction of solar radiation by the canopy of shrubs was likely the main factor determining shrub facilitation. The growth of the pines differed among years. However, growth was not inhibited when planted with shrubs as compared with open areas in any of the years. Herbivore damage was low but was mostly concentrated in the leader shoot, exacerbating the deleterious effect of ungulate herbivores on pine growth. We conclude that the use of shrubs as nurse plants for reforestation is a viable technique to increase establishment success of reforestation in Mediterranean‐type ecosystems and that it might be similarly useful in other water‐stressed environments. In addition, this technique offers the advantage of following natural succession, thus minimizing the impact in the community.     

Invasional 'meltdown' on an oceanic island

Islands can serve as model systems for understanding how biological invasions affect community structure and ecosystem function. Here we show invasion by the alien crazy ant Anoplolepis gracilipes causes a rapid, catastrophic shift in the rain forest ecosystem of a tropical oceanic island, affecting at least three trophic levels. In invaded areas, crazy ants extirpate the red land crab, the dominant endemic consumer on the forest floor. In doing so, crazy ants indirectly release seedling recruitment, enhance species richness of seedlings, and slow litter breakdown. In the forest canopy, new associations between this invasive ant and honeydew‐secreting scale insects accelerate and diversify impacts. Sustained high densities of foraging ants on canopy trees result in high population densities of host‐generalist scale insects and growth of sooty moulds, leading to canopy dieback and even deaths of canopy trees. The indirect fallout from the displacement of a native ‘keystone’ species by an ant invader, itself abetted by introduced/cryptogenic mutualists, produces synergism in impacts to precipitate invasional ‘meltdown’ in this system.