Nurse plants vs. nurse objects: effects of woody plants and rocky cavities on the recruitment of the Pilosocereus leucocephalus columnar cactus

BACKGROUND AND AIMS: Most studies on cactus recruitment have focused on the role of woody plants as seedling facilitators. Although the spatial association of cacti with objects had been described, the mechanisms underlying this association remain unknown. The aims of this study were to identify which mechanisms facilitate the establishment of a columnar cactus under the shade and protection of objects and to compare these mechanisms with those involved in plant-plant facilitation. METHODS: Three split-split-plot field experiments were conducted to compare the effects of two microhabitats (inside rocky cavities and beneath plant canopies) on seed removal, germination, seedling survivorship and dry weight. Flat, open spaces were used as the control. For each microhabitat, the effect of seed or seedling protection and substrate limitation were explored; aboveground microclimate and some soil properties were also characterized. KEY RESULTS: The permanence of superficial seeds was greater inside rocky cavities than beneath woody plant canopies or on flat, open areas. Germination was similar in cavities and beneath plant canopies, but significantly higher than on flat, open areas. Seedling survivorship was greater beneath plant canopies than inside cavities or on flat, open spaces. CONCLUSIONS: The mechanisms of plant facilitation are different from those of object facilitation. There are seed-seedling conflicts involved in the recruitment of P. leucocephalus: nurse plants favour mainly seedling survivorship by providing a suitable microenvironment, while nurse objects mainly favour seed permanence, by protecting them from predators.     

Competition and facilitation structure plant communities under nurse tree canopies in extremely stressful environments

Nurse plant facilitation in stressful environments can produce an environment with relatively low stress under its canopy. These nurse plants may produce the conditions promoting intense competition between coexisting species under the canopy, and canopies may establish stress gradients, where stress increases toward the edge of the canopy. Competition and facilitation on these stress gradients may control species distributions in the communities under canopies. We tested the following predictions: (1) interactions between understory species shift from competition to facilitation in habitats experiencing increasing stress from the center to the edge of canopy of a nurse plant, and (2) species distributions in understory communities are controlled by competitive interactions at the center of canopy, and facilitation at the edge of the canopy. We tested these predictions using a neighbor removal experiment under nurse trees growing in arid environments. Established individuals of each of four of the most common herbaceous species in the understory were used in the experiment. Two species were more frequent in the center of the canopy, and two species were more frequent at the edge of the canopy. Established individuals of each species were subjected to neighbor removal or control treatments in both canopy center and edge habitats. We found a shift from competitive to facilitative interactions from the center to the edge of the canopy. The shift in the effect of neighbors on the target species can help to explain species distributions in these canopies. Canopy‐dominant species only perform well in the presence of neighbors in the edge microhabitat. Competition from canopy‐dominant species can also limit the performance of edge‐dominant species in the canopy microhabitat. The shift from competition to facilitation under nurse plant canopies can structure the understory communities in extremely stressful environments.     

Shrub emergence and seedling growth in microenvironments created by Prosopis glandulosa

Abstract. Facilitation of the establishment of certain plant species by nurse plants is a common phenomenon in arid and semiarid ecosystems. The most commonly reported mechanisms of facilitation include cooler temperatures and increased soil nutrients beneath the nurse plant canopy, which favor establishment of other plant species. During conversion of upland grasslands to thorn woodland in southern Texas, Prosopis glandulosa appears to facilitate establishment of other woody plants, including Celtis pallida, whereas Acacia smallii occurs only in habitats between P. glandulosa canopies. We tested the hypothesis that light intensity and soils under P. glandulosa canopies facilitate seedling emergence and growth of C. pallida but inhibit seedling emergence and growth of A. smallii. In the field, C. pallida and A. smallii seeds were planted under P. glandulosa canopies and in adjacent interspaces. Percent emergence of C. pallida seedlings was greater under the canopy of P. glandulosa, whereas percent emergence of A. smallii seedlings was greater in interspaces. In a greenhouse experiment, seeds of each species were planted in pots filled with soil from under P. glandulosa canopies or from adjacent interspaces. Two treatments, shade and sunlight, were imposed and plants harvested seven weeks later. Seedling mass of both species was greater in canopy soil than in interspace soil in sunlight but mass of the two species did not differ between soil sources in shade. Canopy soils contained more total and available nitrogen than interspace soils. These results suggest that light is more limiting than nutrients under shaded conditions and so neither species can take advantage of the high nutrients beneath P. glandulosa. Shade and greater soil nutrients beneath P. glandulosa do not appear to be the major factors that facilitate C. pallida or inhibit A. smallii. Aggregation of C. pallida beneath P. glandulosa canopies appears to be a complex process that involves both passive facilitation (seed dispersal by birds) and active facilitation (reduction of seed dormancy by under-canopy temperatures) operating only during the seed germination stage with successional mechanisms other than facilitation operating during later stages of shrub establishment and growth.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

The interplay of nurse and target plant traits influences magnitude and direction of facilitative interactions under different combinations of stress and disturbance intensities in Andean dry grassland

Aims Facilitation is a key process in vegetation dynamics, driving the response to natural and anthropogenic pressures. In harsh-grazed systems, palatable plants mainly survive when nested under unpalatable tussocks and shrubs. The magnitude and direction of positive interactions are driven by resource availability, extent of herbivory and type of nurse species. We hypothesized that different combinations of disturbance and environmental stress affect community composition in the dry Puna (southern Peruvian Andes) by modifying nurse types and plant interactions in magnitude and specific associations. We investigated whether different combinations of stress and disturbance influence species richness, type and frequency of occurrence of nurse and beneficiary species and magnitude and patterns of plant interactions; whether nurse species influence these interactions and target species change their interactions under different combinations of stress and disturbance and whether plant functional traits differ in the studied communities and influence the pattern of spatial interactions.Methods We selected three plant communities subject to different precipitation and management regimes: in each we laid a number of transects proportional to its extension. Data collected include species presence/absence, type of spatial interactions with nurse species and functional traits. We calculated species richness and rarefaction patterns, described the patterns of plant–plant spatial interactions and investigated the associations between nurse and other species in the three communities using indicator species analysis (ISA). We performed ISA and correlation analysis to investigate whether plant functional traits influenced facilitative interactions.Important findings We found that different combinations of stress and disturbance shaped a complex set of responses, including changes in the nurse species set. Nurse composition influenced magnitude and direction of plant interactions under different stress intensities. Heavy disturbance increased the relative importance of facilitation, even if the overall number of facilitated species decreased. Under equivalent disturbance regimes, increased abiotic stress led to a greater importance of facilitation. Different combinations of stress and disturbance affected the community assemblage also by changing the behaviour of some non-nurse species. Both heavy disturbance and strong stress led to a decrease of trait states; with certain combinations of stress and disturbance, preferential distribution of these states was observed. We also found that plant traits were of key importance in determining facilitative interactions. Some traits were mainly associated with one type of spatial interaction: plant architecture, life cycle and root type influenced the type of interaction between nurses and beneficiaries under different combinations of stress and disturbance. Our results also demonstrate that in plant interaction research the object of observations (species per se, species percentage, etc.) might influence outputs, and to effectively assess the impact of different stress and disturbance intensities on plant interactions it is necessary to work at the community level to consider the whole species pool.     

The ecological performance of metallophyte plants thriving in geochemical islands is explained by the Inclusive Niche Hypothesis

Aims The Inclusive Niche Hypothesis has not been validated for plants using ecophysiological performance. The few experiments have measured growth and competition but not the physiological response of plants. A metallophyte plant that hyperaccumulates aluminium (Al), Plantago almogravensis, showed a defined spatial distribution by occurring mostly on vegetation gaps associated with metalliferous areas (geochemical islands). This case was used to determine, in situ, whether the Inclusive Niche Hypothesis was suitable to explain the extent of the species realized niche.Methods The vegetation associated with P. almogravensis geochemical islands in the SW coast of Portugal was mapped. The biotic (neighbouring plants) and abiotic (edaphic) components of the niche were correlated with parameters of the plant's ecological and physiological performances (plant density and cover; leaf C and N concentration and isotopic composition; growth). The results were obtained using image analysis, abundance and morphological measures, isotopic signatures and chemical composition.Important findings The species showed better physiological performance where its ecological performance was lower due to trade-offs with environmental constraints. The species' realized niche was mostly limited by shrub competition and soil Al-toxicity. These limits contribute to explain the rarity status of the species: the species has a poor capacity to compete but, due to an enhanced Al-tolerance and Al-hyperaccumulator trait, has the ability to find refuge in geochemical islands that are too harsh for most other species. This work, based on ecophysiological field studies, provides support for the Inclusive Niche Hypothesis relating to plant species.     

Role of nurse plants in Araucaria Forest expansion over grassland in south Brazil

Abstract Araucaria Forest expansion over grassland takes place under wet climate conditions and low disturbance and it is hypothesized that isolated trees established on grassland facilitate the establishment of forest woody species beneath their canopies. Forest with Araucaria angustifolia is a particular type of Brazilian Atlantic Forest and the main forest type on the highland plateau in south Brazil, often forming mosaics with natural Campos grassland. The objectives of this paper were to evaluate the role of isolated shrubs and trees as colonization sites for seedlings of Araucaria Forest woody species on grassland, to determine which species function as preferential nurse plants in the process and the importance of vertebrate diaspore dispersal on the structure of seedling communities beneath nurse plants. The study was conducted in São Francisco de Paula, Rio Grande do Sul State, where we sampled isolated shrubs and trees in natural grassland near Araucaria Forest edges. Seedlings were counted and identified, and seedling diaspore dispersal syndromes, size and colour were registered. We detected 11 woody species with a potential role in nucleating grassland colonization by forest species. Beneath the canopies of nurse plants more forest species seedlings were found compared with open field grassland and the seedlings had diaspores mostly dispersed by vertebrates. Also, more seedlings were found under the canopy of A. angustifolia than beneath other nurse plant species. We conclude that A. angustifolia trees established on grassland act as nurse plants, by attracting disperser birds that promote colonization of the site by other forest species seedlings, and that under low level of grassland disturbance, conservation of frugivorous vertebrate assemblages may increase forest expansion over natural grassland and also facilitate the regeneration of degraded forest areas.     

Facilitation by nurse shrubs of resprouting behavior in a post-fire shrubland in northern Patagonia,Argentina

Abstract. The nurse-plant syndrome is a widely recognized example of positive (facilitative) influences of plant species on the establishment and growth of other species. Most studies of the nurse-plant syndrome have been on species that reproduce mainly from seed rather than vegetatively. In this study, we experimentally compared the influences of two species of nurse shrubs, Schinus patagonicus and Berberis buxifolia, on the survival and growth of vegetatively reproducing herbaceous and woody plants in a post-fire shrubland in northern Patagonia, Argentina. The vegetation beneath shrubs was removed by clipping and, in a paired-sample design, one half of the canopy of each shrub was removed. We determined species richness, counted number of resprouts, and measured photon flux density and soil moisture beneath cut and uncut halves of each shrubs. Abundances of resprouts were several times greater beneath the uncut vs. the cut shrubs, as was the mean number of species. Thus, shrubs have a strong facilitating influence as measured by resprout densities and the number of species. Numbers of resprouts and of species were twice as high beneath Schinus as beneath Berberis, implying important differences in the facilitative effects of the two shrubs species. Microsites beneath Schinus were characterized by lower and more heterogeneous light levels but by greater soil moisture. Even though the reproductive mode in this post-fire shrubland is overwhelmingly vegetative, facilitation by nurse shrubs is important and differentially effective for different species of nurse shrubs.     

Testing a facilitation model for ecosystem restoration: Does tree planting restore ground layer species in a grassy woodland?

Planning for the restoration of degraded ecosystems has a strong basis in facilitation successional theory, which, as applied in restoration practice, states that planting of structurally dominant tree species will assist the entry of other native species into a restored community. In Australia, tree planting has been widely applied in restoration of grassy woodland ecosystems. Trees have been postulated to reduce the cover and diversity of weed species, thus facilitating recolonization of native woodland species (indirect facilitation). The expected outcomes of this process include reduced species richness and abundance of exotic plant species and increased species richness and abundance/dominance of natives in areas beneath tree canopies, with these trends strengthening with time. To assess whether this was occurring, we carried out a comparative analysis of species assemblages found underneath and outside of planted tree canopies in sites replanted with juvenile canopy tree species 3–5 or 8–10 years previously. We sampled revegetated stands of Cumberland Plain Woodland, an endangered ecological community in Western Sydney, Australia. We found that neither the number nor abundance of native ground layer species beneath canopies increased as a result of trees being planted at sites of both ages. Where seed is limited, we predicted an increase in abundance of existing native species under planted tree canopies. On this point, the results were mixed and showed some natives with an increased abundance while others decreased. Exotic species richness showed the reverse of the expected pattern, being greater under tree canopies. These findings lend no support to the theory of indirect facilitation. We conclude that simple facilitation models may be inadequate to support planning of grassy woodland restoration and that those models incorporating successional time lags and restoration barriers are likely to be more informative about the development of communities initiated by tree planting.     

Effects of plant-plant interactions and herbivory on the plant community structure in an arid environment of Saudi Arabia

There is currently considerable evidence support that plant community structures are driven by plant-plant interactions (e.g., competition and facilitation). In contrast, there is also evidence demonstrating that plant community structure is affected by the impact of consumer pressure (e.g., grazing). In this study, 15 and 10 Acacia gerrardii nurse plants were selected inside and outside Sudyrah natural reserve (protected) area in western Saudi Arabia, respectively. The understory vegetation abundance (e.g. cover and density) was measured among quadrats around the nurse trees in both protected and unprotected areas to examine the impact of grazing and the positive interaction on the understory species. I found that understory vegetation associated with nurse trees (A. gerrardii) has been driven by both the positive impact of nurse plant and the grazing. Although the understory vegetation was positively affected by the impact of facilitation, the composition of such vegetation has been changed due to the impact of herbivory.     

Simultaneous,independent, and additive effects of shrub facilitation and understory competition on the survival of a native forb (Penstemon palmeri)

There is increasing recognition that both competition and facilitation are important drivers of plant community dynamics in arid and semi-arid environments. Decades of research have provided a litany of examples of the potential for shrubs as nurse plants for establishment of desirable species, especially in water-limited environments. However, interactions with the existing understory community may alter the outcome of interactions between shrubs and understory plants. A manipulative experiment was conducted to disentangle interactions between a native forb species (Penstemon palmeri A. Gray), a native shrub (Artemisia tridentata Nutt.), and a diverse understory of exotic and native forbs and grasses in a semi-arid shrubland of Northern Utah, USA. Seedlings of P. palmeri were transplanted in a factorial design: (1) beneath shrub canopies or into their interspaces and (2) with understory interactions retained or removed. Transplant survival was tracked for roughly 1 year. Shrubs appeared to facilitate P. palmeri survival while interactions with the existing understory community were equivalently negative, leading to overall neutral interactions. Further, positive shrub interactions and negative understory interactions appeared to operate independently and simultaneously. While the debate over the importance of facilitation and competition in driving plant community dynamics continues, our observations strongly suggest that both have considerable effects on plant establishment in A. tridentata communities. Furthermore, our results inform the conservation and restoration of P. palmeri populations, and suggest the utility of nurse shrubs and/or understory thinning as strategies for increasing the diversity of desirable species in the arid and semi-arid western United States shrublands.     

The shrub Rhodomyrtus tomentosa acts as a nurse plant for seedlings differing in shade tolerance in degraded land of South China

Questions: What are the nurse effects of Rhodomyrtus tomentosa in degraded land of South China? Are canopy or soil factors responsible for the main nurse effect? Do facilitative effects increase with the shade tolerance of the target species? Location: Degraded shrubland in South China. Methods: Seedlings of three native climax woody species (Schima superba, Michelia macclurei, Castanopsis fissa) that differ in shade tolerance were subjected to four treatments by transplantation: (1) under the canopy of R. tomentosa shrubs; (2) in open interspaces without vegetation cover (control); (3) under the canopy of R. tomentosa from which canopies had been removed; and (4) in open interspaces without vegetation but covered by branches and leaves of R. tomentosa. Results: At low soil nutrient levels, increased canopy shade, soil porosity and soil moisture under the canopy of R. tomentosa increased seedling survival of the climax tree species S. superba, C. fissa and M. macclurei, and shoot height of S. superba. The nurse effect (a form of facilitation) of R. tomentosa depended more on canopy shade than on soil amelioration. The magnitude of facilitation or nurse effect was positively correlated with shade tolerance of the target species. Conclusions: Use of nurse plants in restoration differs from traditional reforestation (clearing and/or burning to reduce interspecific competition between target tree species and non‐target species) because it focuses on positive interactions between nurse plants and target plants that increase establishment of target species and reduce time required for restoration. Because nurse effects of R. tomentosa shrubs tended to be larger on target species with greater shade tolerance, shade‐tolerant plants are suggested as target species to accelerate restoration.     

Positive effects of shrubs on plant species diversity do not change along a gradient in grazing pressure in an arid shrubland

Facilitative or positive interactions among species are driven mainly by the environmental amelioration or protection from grazing provided by nurse plants. Some studies have suggested that protection from grazing is inconsequential in water-limited environments because of low herbivore densities and their grazing effects. Others, however, argue that herbivores have a major effect on semi-arid plant communities, and that protection from grazing is a significant factor driving positive plant–plant interactions in such environments. We identified a gradient in grazing pressure in a semi-arid shrubland in south-eastern Australia along which we compared soil condition, incident radiation and plant composition beneath two nurse shrub species with open (shrub-free) interspaces. Our aim was to assess the degree of microclimatic amelioration provided by both shrubs, and changes in the interactions (intensity, importance and frequency) between both nurse shrubs and their understorey species, and their effects on species richness at the community level. Both the relative interaction intensity (RII) and interaction importance (I_imp) indices of plant–plant interactions were generally positive and independent of grazing pressure. Soil beneath both nurse plants had significantly greater indices of nutrient cycling and infiltration, and contained more C and N than soil in the open. Almost twice as many species occurred under the canopies of both shrubs (44 species) than in the open (23 species), and the composition of species differed significantly among microsites. Fifty-four percent of all perennial plant species occurred exclusively under shrubs. Our results suggest that environmental amelioration is a stronger driver of the facilitatory effect of shrubs on their understorey species than protection from grazing. Our conclusions are based on the fact that the substantial effect of plant–plant interactions on plant species richness was largely independent of grazing pressure. Irrespective of the underlying mechanism for this effect, our study illustrates the ecological role of shrubs as refugia for understorey plants in semi-arid environments and cautions against management practices aimed at reducing shrub populations.     

Regeneration niche differentiates functional strategies of desert woody plant species

Plant communities vary dramatically in the number and relative abundance of species that exhibit facilitative interactions, which contributes substantially to variation in community structure and dynamics. Predicting species’ responses to neighbors based on readily measurable functional traits would provide important insight into the factors that structure plant communities. We measured a suite of functional traits on seedlings of 20 species and mature plants of 54 species of shrubs from three arid biogeographic regions. We hypothesized that species with different regeneration niches—those that require nurse plants for establishment (beneficiaries) versus those that do not (colonizers)—are functionally different. Indeed, seedlings of beneficiary species had lower relative growth rates, larger seeds and final biomass, allocated biomass toward roots and height at a cost to leaf mass fraction, and constructed costly, dense leaf and root tissues relative to colonizers. Likewise at maturity, beneficiaries had larger overall size and denser leaves coupled with greater water use efficiency than colonizers. In contrast to current hypotheses that suggest beneficiaries are less “stress-tolerant” than colonizers, beneficiaries exhibited conservative functional strategies suited to persistently dry, low light conditions beneath canopies, whereas colonizers exhibited opportunistic strategies that may be advantageous in fluctuating, open microenvironments. In addition, the signature of the regeneration niche at maturity indicates that facilitation expands the range of functional diversity within plant communities at all ontogenetic stages. This study demonstrates the utility of specific functional traits for predicting species’ regeneration niches in hot deserts, and provides a framework for studying facilitation in other severe environments.     

Nurse plants have the potential to accelerate vegetation recovery in Lapalala Wilderness old fields,South Africa

Nurse plants offer microclimates that are favourable to plant growth of understory native vegetation, thus facilitating ecological restoration in degraded old fields. This study examined the impact of three nurse plants on vegetation diversity and soil physical properties in old fields located at Lapalala Wilderness, South Africa. Vegetation surveys in plots measuring 5 m x 5 m under and outside the canopy of nurse plants in both old field and natural sites were conducted. Top soils under and outside the nurse plants canopy were collected in all plots and quantified for soil moisture, soil penetration resistance and soil water repellency. Results indicate that species diversity was high under plant canopy compared to outside plant canopy for all nurse plants. Soils under nurse plants canopy showed improved soil moisture and soil penetration resistance compared to soils outside plant canopy, but no differences were reported for soil water repellency. The study concludes that the presence of native plants under nurse plants canopy points to a positive vegetation recovery trajectory. For vegetation and soil restoration to be effective in Lapalala Wilderness old fields, nurse plants should be protected and active restoration, e.g. seeding or seedling sowing under nurse plants canopy should be considered.     

Directed dispersal and decomposition drive cyclic succession in arid subtropical thicket

The influence of animal-mediated processes on the purported cyclical succession model of vegetation dynamics in arid subtropical thicket of southern South Africa was examined. We used spatially explicit sampling methods to evaluate the influence of dispersal and decomposition on plant community structure. The harvester ant Messor capensis was the only ant species recorded harvesting Portulacaria afra fruit and the elaiosome bearing seed of the tree Gymnosporia polyacantha from experimental piles. Propagules were dispersed to nest mounds positioned beneath woody shrubs. Vertebrates dispersed fleshy-fruited tree species to regeneration sites beneath P. afra clumps. The minute seeds of succulent shrubs were secondarily dispersed from beneath shrubs to ant nest sites located in the open. The results from two experiments indicate that soil nutrients and decomposition activity are spatially localised. The rate of below-ground decomposition, and the incidence of an invertebrate detritivore assemblage, was high beneath P. afra clumps compared with exposed nutrient-enriched and grassy sites. Animal dispersal vectors are integral to the regeneration of major plant groups in arid subtropical thicket by moving seeds from competitively harsh environments to safe sites beneath nurse plants. Decomposition and nutrient release are localised beneath P. afra clumps due to the unique conditions that support a suite of detritivore taxa recorded nowhere else at the study site.     

Relationship between soil nutrient availability and plant species richness in a tropical semi‐arid environment

Question: What is the relationship between soil fertility and plant species richness in the ‘fertile islands’ occurring beneath two species of legume (Cercidium praecox and Prosopis laevigata)? Location: Tehuacán‐Cuicatlán region, central Mexico. Methods: Plant richness was measured in three micro‐environments (below canopies of C. praecox, below canopies of P. laevigata and in areas without canopies). The concentration of soil nutrients (C, N and P), C and N in the microbiota, and processes of ecosystem functioning (net C mineralization rate and N mineralization) were measured. The relationship between soil variables and plant richness were assessed with ANCOVAs. Results: Soil nutrients and species richness increases markedly under fertility islands. There were higher concentrations of C and N in the soil, faster rates of C mineralization, and higher species richness under P. laevigata canopies. The relationship between soil fertility and species richness was always positive except for total N, ammonium and net C mineralization rate under C. praecox, and for available P under P. laevigata. Conclusions: The sign of the relationship between soil fertility and species richness varies according to the nutrient and the micro‐environment. Positive relationships could result from between species complementarity and facilitation. Negative relationships could be explained by a specific limitation threshold for some soil resources (P and N for plants and C for the soil microbiota) which eliminate the possibilities of between species complementarity and facilitation above that threshold. As in all observational studies, these relationships should be considered only correlational.     

The shift from plant–plant facilitation to competition under severe water deficit is spatially explicit

The stress‐gradient hypothesis predicts a higher frequency of facilitative interactions as resource limitation increases. Under severe resource limitation, it has been suggested that facilitation may revert to competition, and identifying the presence as well as determining the magnitude of this shift is important for predicting the effect of climate change on biodiversity and plant community dynamics. In this study, we perform a meta‐analysis to compare temporal differences of species diversity and productivity under a nurse plant (Retama sphaerocarpa) with varying annual rainfall quantity to test the effect of water limitation on facilitation. Furthermore, we assess spatial differences in the herbaceous community under nurse plants in situ during a year with below‐average rainfall. We found evidence that severe rainfall deficit reduced species diversity and plant productivity under nurse plants relative to open areas. Our results indicate that the switch from facilitation to competition in response to rainfall quantity is nonlinear. The magnitude of this switch depended on the aspect around the nurse plant. Hotter south aspects under nurse plants resulted in negative effects on beneficiary species, while the north aspect still showed facilitation. Combined, these results emphasize the importance of spatial heterogeneity under nurse plants for mediating species loss under reduced precipitation, as predicted by future climate change scenarios. However, the decreased water availability expected under climate change will likely reduce overall facilitation and limit the role of nurse plants as refugia, amplifying biodiversity loss.     

Ecological longevity of Polaskia chende (Cactaceae) seeds in the soil seed bank,seedling emergence and survival

• Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5‐year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico.
• P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12‐h photoperiod under white light, far‐red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years.
• The species forms long‐term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling.
• Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence.

     

Love of Nurse Plants is Not Enough for Restoring Oak Forests in a Seasonally Dry Tropical Environment

The highest concentration of oak species in the world occurs in Mexico, but human activities have strongly degraded these oak forests. Mexican oaks have high economic, social, and cultural value, and restoring these forests is of paramount importance for the people of Mexico. Here, we propose a method for restoring oak forests using native shrubs that colonize degraded areas as nurse plants for oak seedlings. To test the viability of this proposal, seedling transplant experiments were performed in a degraded area near a protected oak forest relict. Two pioneer shrubs were identified as potential nurse species: Mimosa luisana and Senecio sp. The target oak species was Quercus castanea . Oak seedlings were located beneath the canopies of both shrubs and in the surrounding area without shrub cover. Water is a limiting resource for oak establishment in seasonally dry environments; therefore, we included irrigation systems in our experimental design to determine whether the combination of nurse plants plus watering led to higher rate of survival than the presence of nurse species alone. Seedling survival without watering was less than 20% both beneath nurse species and in the surrounding habitat. When water was supplied, survival rate beneath nurse species increased up to 58% while survival in the surrounding habitat did not differ from that observed in treatments without watering. Our results indicate that survival rate of oak seedlings is increased by the presence of nurse plants only when water is supplied. This suggests that restoration of oak forests in these degraded areas requires both nurse plants and watering.