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.     

Canopy Size Dependent Facilitations from the Native Shrub Rhodomyrtus tomentosa to the Early Establishment of Native Trees Castanopsis fissa and Syzygium hancei in Tropical China

The quantitative role of the canopy size of nurse shrubs on microenvironment and native tree establishment in degraded tropical lands has been seldom studied. In a 21‐month field experiment, we aimed to test the effect of a native shrub with different canopy sizes on the early establishment of native trees as part of the effort of forest restoration in tropical China. We examined the microenvironment, and the seedling establishment and growth of two native trees: Castanopsis fissa and Syzygium hancei in both open space (OS) microsite and microsite under the canopy of the native pioneer shrub Rhodomyrtus tomentosa. Shrub microsite was further divided into large canopy (LC), and medium canopy (MC) microsite, based on the shrub leaf area indices. Results showed that relative to OS, LC had higher soil nutrient concentration and water content, and lower photosynthetic active radiation (PAR), while MC had lower PAR and higher soil exchangeable Mg, K, and Ca. Survival and growth were mostly enhanced, while water stress and photoinhibition reduced for C. fissa seedlings in MC and S. hancei seedlings in LC. It is found that the beneficial effects of the native shrub on seedling establishment and growth result mostly from the improvement in nutrient and water availabilities, the reduction in plant stress caused by harsh summer light, and the specific ecological requirements of different tree species. We suggest that different canopy sizes of native shrub R. tomentosa may be explored to target different native trees and hence promote forest restoration in degraded tropical ecosystems.     

Resource-use efficiencies of three indigenous tree species planted in resource islands created by shrubs: implications for reforestation of subtropical degraded shrublands

Shrub resource islands are characterized by resources accumulated shrubby areas surrounded by relative barren soils. This research aims to determine resource-use efficiency of native trees species planted on shrub resource islands, and to determine how the planted trees may influence the resource islands in degraded shrublands in South China. Shrub (Rhodomyrtus tomentosa) resource islands were left unplanted or were planted with 0.5-year-old indigenous tree species Schima superba, Castanopsis fissa, and Michelia macclurei. The results indicate that, after 2.5?years, the tree seedlings did not modify the physical traits (light, air temperature, and soil water) but tended to increase soil nutrients (soil organic carbon and soil nitrogen) and soil microbial biomass of the resource islands. Relative to S. superba and M. macclurei, C. fissa had greater effects on soil nutrients and microbial biomass but showed lower plant growth, survival, and resource-use efficiencies (for water, light, energy, and nitrogen). These results rejected our initial assumption that shrub resource islands would effectively promote the growth and resource utility of all the indigenous tree species and shorten the reforestation course in subtropical degraded shrubland. C. fissa performed poorly when growing on shrub resource islands, but its role in soil nutrient accumulation might have long-term impacts on the restoration of degraded shrubland. In contrast, S. superba and M. macclurei could make better use of the shrub resource islands therefore accelerating the construction of native plantations.     

Restoration of Dry Afromontane Forest Using Pioneer Shrubs as Nurse-Plants for Olea europaea ssp. cuspidata

Shrubs are often considered competitive barriers for seedlings planted in reforestation programs, although they can facilitate tree recruitment, especially in ecosystems under high abiotic stress. An alternative reforestation technique using pioneer shrubs as nurse‐plants for Olea europaea ssp. cuspidata was tested in exclosures in northern Ethiopia. Seedlings were planted in three different microhabitats, and their survival was monitored. The microhabitats were bare soil patches between shrubs, patches under the dominant shrub Acacia etbaica, and patches under Euclea racemosa, an evergreen shrub, which supports the majority of naturally established Olea recruits. The ability of shrubs to offer protection against browsing goats was tested experimentally. Controlled shading was used to determine whether solar irradiation causes seedling mortality in environments without water stress. Data were analyzed using Kaplan–Meier survival analysis, Kruskal–Wallis analysis of variance (ANOVA), and one‐way ANOVA. Olea survival was significantly higher and shoot damage by goats was lower when planted under shrub cover compared to bare soil patches, particularly under Euclea canopies, although high shade levels reduced seedling performance. Reduction of solar radiation by shrub canopies and thus control of soil–water evaporation and seedling transpiration most likely controlled the observed facilitation. Planting under shrubs may increase seedling survival and assist regeneration of dry Afromontane vegetation. Preserving pioneers also reduces soil erosion and conserves biodiversity. Excluding livestock is essential for Olea woodland restoration and allows persistent but morphologically modified Olea shrubs to develop vigorous regrowth. Facilitative processes are guiding principles for assisted forest restoration, but above‐average rains may be critical to restore higher biomass levels in semiarid areas.     

The role of nurse successional stages on species‐specific facilitation in drylands: Nurse traits and facilitation skills

Plant establishment is a challenge in semiarid environments due to intense and frequent drought periods. The presence of neighboring trees (nurses) can increase the establishment of seedlings (targets) by improving resource availability and microclimate. The nurse effect, however, might vary depending on nurse‐target species combinations but factors that predict this specificity are poorly known. We used a multispecies experiment to investigate the facilitation potential of trees from a range of successional stages, focusing on how nurse functional traits can predict species‐specific interaction outcomes. We conducted a factorial field experiment in a Brazilian semiarid tropical forest during a severe drought period. Sixty pairs of interacting tree species, 20 potential nurses, and three targets were used. Seedlings of all targets were planted both under and far from the nurse canopy, in a randomized block design replicated five times. Target growth and survival were monitored for 275 days from the beginning of the dry season, and interaction outcomes were calculated using the Relative Interaction Intensity (RII) index. Nurse functional traits such as successional stage, height, wood density, and canopy diameter were used as explanatory variables to predict RII values. The average effect of nurse species on target plants was in general positive, that is, seedling survival and growth increased under the nurse canopy. However, for growth pairwise interactions were significantly species specific. Successional stage was the only functional trait explaining RII values, with pioneer tree species being stronger facilitators than later successional trees. However, the explanation power of this variable was low, and positive, negative, or neutral interactions were found among nurse trees of all successional stages. Because seedling mortality during drought in semiarid systems is high, future studies should investigate how nurse traits related to water use could influence nurse facilitation skills.     

Facilitation of seedling establishment by an endemic shrub in tropical coastal sand dunes

Facilitation is predicted to occur on coastal sand dunes as these ecosystems have the harsh physical conditions common during primary succession. In a tropical dune system on the Gulf of Mexico the spatial patterns of plant distribution were analyzed and the hypothesis that facilitation is the responsible mechanism was tested by monitoring changes in the micro-environment and performing a seedling planting experiment under natural conditions. Densities of seedlings and adults of late colonizer grasses were two to six times greater beneath the early colonizer shrub, Chamaecrista chamaecristoides, than in the exposed areas. Temperatures on the sand surface, wind speed and sand accretion were significantly reduced by the shade of Chamaecrista. Only phosphate contents during the dry season were significantly higher in sand in the shade than in exposed sand. In the field experiment, successful establishment of seedlings of two late colonizer grasses (Trachypogon plumosus and Schizachyrium scoparium) was low but was concentrated exclusively beneath the shrubs. The introduced plants were reproductive one year after onset of the experiment and remained vigorous after three years. The observed spatial aggregation among the target species may be a result of improved conditions in the shade of the shrub, which facilitated the survival and establishment of late colonizers. On top of the environmental amelioration, protection from recurrent disturbances such as substrate mobility is an additional beneficial effect of the shrub.     

Increasing intraspecific facilitation in exposed environments: consistent results from mountain birch populations in two subarctic stress gradients

The stress‐gradient hypothesis predicts that the importance of facilitation relative to competition should increase with increasing stress. The hypothesis has received support from several environments, but multi‐gradient studies on the generality of the hypothesis are exceptionally rare. A within‐species experiment with mountain birch Betula pubescens subsp. czerepanovii was conducted to test the hypothesis in the extreme ends of two subarctic stress gradients (elevation and seashore) in the Kola Peninsula, northwestern Russia. The high stress sites were characterized by strong winds, temperature extremes and potentially drought. The negative effects of abiotic stress on the study seedlings were verified from performance characteristics. Effects of adult hosts as well as seedling–seedlings interactions were studied. Positive host–seedling interactions dominated in each study site, and three out of four performance variables indicated stronger positive net effects in the high stress sites. In the seashore gradient also seedling survival gave similar interpretations. Also a temporal shift towards host–seedling competition was detected in a low stress site after two study years. In seedling–seedling interactions competition dominated, but the effects were weak, likely due to the ‘noise’ caused by genetic and environmental factors. Our results support the stress‐gradient hypothesis and its generality in subarctic environments, as the interpretations were similar for both stress gradients and several fitness‐related variables. The temporal variation in host–seedling interactions and the difference between host–seedling and seedling–seedling effects hint on size‐dependency of plant interactions: facilitation might dominate when the benefactor is substantially larger than the beneficiary, while competition may be stronger when the plants are of similar size and developmental status.     

Shrub facilitation drives tree establishment in a semiarid fog‐dependent ecosystem

Questions

The exceptional occurrence of tall rain forest patches on foggy coastal mountaintops, surrounded by extensive xerophytic shrublands, suggests an important role of plant–plant interactions in the origin and persistence of these patches in semi‐arid Chile. We asked whether facilitation by shrubs can explain the growth and survival of rain forest tree species, and whether shrub effects depend on the identity of the shrub species itself, the drought tolerance of the tree species and the position of shrubs in regard to wind direction.

Location

Open area–shrubland–forest matrix, Fray Jorge Forest National Park, Chile.

Methods

We recorded survival after 12 years of a ~3600 tree saplings plantation (originally ~30‐cm tall individuals) of Aextoxicon punctatum, Myrceugenia correifolia and Drimys winteri placed outside forests, beneath the shrub Baccharis vernalis, and in open (shrub‐free) areas. We assessed the effects of neighbouring shrubs and soil humidity on survival and growth along a gradient related to the direction of fog movement.

Results

B. vernalis had a clear facilitative effect on tree establishment and survival since, after ~12 years, saplings only survived beneath the shrub canopy. Long‐term survival strongly depended on tree species identity, drought tolerance and position along the soil moisture gradient, with higher survival of A. punctatum (>35%) and M. correifolia (>14%) at sites on wind‐ and fog‐exposed shrubland areas. Sites occupied by the shrub Aristeguietia salvia were unsuitable for trees, presumably due to drier conditions than under B. vernalis.

Conclusions

Interactions between shrubs and fog‐dependent tree species in dry areas revealed a strong, long‐lasting facilitation effect on planted tree's survival and growth. Shrubs acted as benefactors, providing sites suitable for tree growth. Sapling mortality in the shrubland interior was caused by lower soil moisture, the consequence of lower fog loads in the air and thus insufficient facilitation. While B. vernalis was a key ecosystem engineer (nurse) and intercepted fog water that dripped to trees planted underneath, drier sites with A. salvia were unsuitable for trees. Consequently, nurse effects related to water input are strongly site and species specific, with facilitation by shrubs providing a plausible explanation for the initiation of forest patches in this semi‐arid landscape.     

Plantation age,understory vegetation,and species‐specific traits of target seedlings alter the competition and facilitation role of Eucalyptus in South China

The Chinese government is considering ways to reconstruct Eucalyptus plantation monocultures in order to improve their ecosystem services. However, the effects of competition or facilitation of Eucalyptus on native species during the reconstruction have not been identified, especially during the formation of novel ecosystems. To better understand these relationships, in this study we tested the effects of competition and facilitation of Eucalyptus on three target species (Schima superba, Michelia macclurei, and Castanopsis fissa), in Eucalyptus plantations with different ages and understory vegetation. The results showed that the survival and growth of S. superba and M. macclurei were not sensitive to plantation age and understory removal treatment. The survival of C. fissa in understory removal was significantly higher and the growth of C. fissa increased gradually with increasing plantation age. The relative neighbor effect (RNE) index for growth indicated that the relationship between Eucalyptus and target species switched from competition to facilitation with an increase in plantation age. A redundancy analysis (RDA) showed that survival was positively correlated with soil water content and soil phosphorus. Growth was positively correlated with soil organic matter and soil nitrogen and negatively correlated with soil bulk density. Our results indicate that plantation age, understory vegetation, and species‐specific traits of target seedlings could change the competition and facilitation roles of Eucalyptus. Higher survival and growth rates indicated that C. fissa was an appropriate species for the reconstruction of mature Eucalyptus plantations and understory removal will benefit the process.     

Effect of shrubs and seasonal variability of rainfall on the establishment of Aspidosperma quebracho‐blanco in two edaphically contrasting environments

Abstract Three experiments were conducted to verify if an increase in environmental stress level would affect the interactions between two species of nurse shrubs and seedlings of Aspidosperma quebracho‐blanco. This is a mesic species with a generalist distribution over an extensive environments gradient. The relationship between Larrea divaricata and seedlings of A. quebracho‐blanco was studied in two contrasting soils, a silty loam soil with higher surface clay content and a sandy loam soil. The effect of seasonal variability of rainfall on the initial establishment of seedlings under the shade of L. divaricata was evaluated in three consecutive years. The effect of nurse plant shade was tested comparing two shrub species with different types of leaf life span (sclerophyllous‐evergreen and leguminous‐deciduous). The natural establishment of A. quebracho‐blanco depended on shaded microsites, but not on the type of shade provided by different nurse shrubs. Emergence and initial establishment depended on interactions of soil type and seasonal rainfall variation with nurse plants. The importance of facilitation increased with clay soil (CS). Sandy soil was ‘less humid’ than CS under shrub shade. However, establishment success depends on opportune even rainfall distribution in interaction with nurse plant presence.     

Seedling Ecology and Restoration of Blackbrush (Coleogyne ramosissima) in the Mojave Desert,United States

Increases in fire frequency are disrupting many ecological communities not historically subjected to fire. In the southwestern United States, the blackbrush (Coleogyne ramosissima) community is among the most threatened, often replaced by invasive annual grasses after fire. This long‐lived shrub is vulnerable because it recruits sporadically, partially due to mast seeding and the absence of a seed bank. The goal of this study was to evaluate if shrub restoration can be enhanced by identifying and ameliorating recruitment limitations. Specifically, we tested the effect of encapsulating seeds in predation‐deterring “seed balls.” We also tested the effects of nurse plants and mammalian exclusion cages on seedling emergence, growth, and survivorship. These experiments were conducted in a full‐factorial design across three sites differing in elevation. Over 2 years, 13% of all planted seeds emerged and the effect of seed balls was overwhelmingly negative because of low emergence. Nurse plants had overall positive effects at Low Elevation, but negative effects at Mid‐ and High Elevation. Emergence and survival were highest in caged plots everywhere, and effect sizes increased with elevation. Interactions between the cage and the nurse plant treatments indicated that nurse plants tended to attract mammalian predators, lowering emergence and seedling survivorship, particularly at higher elevations. Findings conform to the stress‐gradient hypothesis in that interactions among seedlings and mature plants shifted from facilitation to competition as environmental stress decreased with increasing elevation, suggesting that they are transferable to ecologically similar communities elsewhere. Knowledge of site‐specific recruitment limitations can help minimize ineffective restoration efforts.     

Resource availability and growth responses to defoliation in seedlings of three early-successional,tropical, woody species

A growth experiment was conducted using seedlings of three early-successional, tropical, woody species:Dillenia suffruticosa (Dilleniaceae),Macaranga heynei (Euphorbiaceae) andTrema tomentosa (Ulmaceae). These species are characteristic of different positions along a soil fertility gradient in Singapore, withD. suffruticosa being the least andT. tomentosa the most demanding of high nutrient availability. The seedlings were grown in vermiculite at either low or high nutrient availabilities supplied by watering with different concentrations of a commercial plant food. Half the seedlings were subjected to a 50% defoliation at the start of the experiment by cutting off the distal half of each leaf. After 9 weeks the plants were harvested.Macaranga heynei andT. tomentosa seedlings showed no significant difference in parameters of growth such as total dry weight and total leaf area between the defoliated and control seedlings. The seedlings compensated completely for the loss of leaf area.Dillenia suffruticosa did show significant reductions in growth in some parameters due to defoliation, and these were more pronounced under the high nutrient treatment. These findings support the hypothesis that plants characteristic of resource-rich sites can readily recover from herbivory through fast growth, probably associated with a rapid turnover of leaves, whereas species of resource-poor habitats cannot easily replace losses due to herbivory and are adversely effected by defoliation.     

A field test of the stress‐gradient hypothesis along an aridity gradient

Aims: The stress‐gradient hypothesis (SGH) predicts how plant interactions change along environmental stress gradients. We tested the SGH in an aridity gradient, where support for the hypothesis and the specific shape of its response curve is controversial. Location: Almería, Cáceres and Coimbra, three sites in the Iberian Peninsula that encompass the most arid and wet habitats in the distribution range of a nurse shrub species –Retama sphaerocarpa L. (Boiss) – in Europe. Methods: We analysed the effect of Retama on its understorey plant community and measured the biomass and species richness beneath Retama and in gaps. We estimated the frequency (changes in species richness), importance and intensity of the Retama effects, and derived the severity–interaction relationship pattern, analysing how these interaction indices changed along this aridity gradient. Results and conclusions: The intensity and frequency of facilitation by Retama increased monotonically with increasing environmental severity, and the importance tended to have a similar pattern, overall supporting the SGH. Our data did not support predictions from recent revisions of the SGH, which may not apply to whole plant communities like those studied here or when interactions are highly asymmetrical. Facilitation by Retama influenced community composition and species richness to the point that a significant fraction of species found at the most arid end of the gradient were only able to survive beneath the nurse shrub, whereas some of these species were able to thrive in gaps at more mesic sites, highlighting the ecological relevance of facilitation by nurse species in mediterranean environments, especially in the driest sites.     

Feathermoss seedbeds facilitate black spruce seedling recruitment in the forest–tundra ecotone (Labrador,Canada)

As climate warms, conifers are expected to expand their ranges into alpine tundra where ecological factors such as seedbed availability, and post‐dispersal seed and seedling predation may control local recruitment. Seedbed composition may influence microhabitat, nutrients, physical structure, and predation level and, therefore, affect the success of conifer recruitment, thereby providing the template for future expansion. In the boreal forest, seedbed–seedling competition dominates such that seedbed removal increases black spruce recruitment. In the harsher climate of the Mealy Mountains boreal forest–tundra ecotone (Labrador, Canada) the Stress gradient hypothesis (SGH) predicts that facilitation may dominate seedbed–seedling interactions. This study investigated potential mechanisms of seedbed facilitation (temperature, water, nutrients, physical protection) in three conifer seedbeds (Pleurozium schreberi, Cladonia spp., bare soil) and examined whether seed predation and/or seedling herbivory varied among seedbeds over three years. Seed emergence was low overall (< 10% on all treatments), but highest on Pleurozium (6.3%), followed by bare ground (4.6%) and Cladonia (0.3%). Facilitation was observed between Pleurozium and black spruce as seedling height increase (31%) and survival (55%) were highest; herbivory, seed predation and overwinter mortality were lowest compared to both Cladonia and bare ground seedbeds. Unlike in the closed canopy boreal forest, seedlings recruited poorly on bare soil as seedling height increase and survival were 20.5% and 26%. Temperature and water availability were similar across seedbeds, while nutrient availability was higher on Pleurozium. The physical structure of Pleurozium likely protects first to third‐year seedlings from temperature extremes and predators. As climate warms and seed availability increases, Pleurozium may facilitate black spruce recruitment and treeline expansion.     

The influence of light conditions and interspecific competition on the root foraging traits and seedling growth of two tree species

Abstract

Enriched nutrient patches within natural soil represent an important source of nutrients for tree growth. In the present study, pot experiments in a heterogeneous nutrient environment were conducted to investigate the influence of light conditions and interspecific competition on the root foraging traits and seedling growth of Pinus massoniana and Schima superba. The root foraging scale and the whole-seedling biomass of both species were decreased by shading. The result of this treatment was a lower sensitivity to nutrient heterogeneity in plants that underwent the shading treatment than in plants that were exposed to full-light conditions. The above-ground biomass and whole-seedling biomass of S. superba were not affected by competition with P. massoniana. In contrast, the above-ground biomass and whole-seedling biomass of P. massoniana were negatively affected by competition with S. superba. The more rapid rate of root extension and the more efficient resource uptake of S.superba appear to explain this effect. The species-specific patterns of the influence of environmental factors on foraging ability and seedling growth should be given thorough consideration and should be applied to afforestation and to the management of tree plantations.     

Age-related change in canopy traits shifts conspecific facilitation to interference in a semi-arid shrubland

Shifts between facilitation and interference and their importance in shaping plant population and community dynamics have received wide recognition. Nevertheless, the causes and spatio-temporal scales of these shifts are poorly understood, yet strongly debated. This study tested the hypothesis that age-related changes in canopy structure shift the effect of a nurse shrub on their protégé from facilitation to interference, using as a model system the interaction between the dwarf shrub Sarcopoterium spinosum and conspecific new recruits, in the shrubland of the transition area between the Mediterranean and the semi-arid climatic zones of Israel. Foliation level (i.e. the percentage of canopy surface area covered with leaves), a measure of shrub canopy structure, increased with age. Shading level was significantly and positively related to foliation level. Densities of new recruits in the shrubland showed a unimodal response to canopy structure and cover: the highest densities were associated with canopies presenting low and medium foliation levels (providing 71 and 82% shade, respectively), while high foliation levels (93% shade) and open spaces among canopies were characterized by very low densities. A related field experiment using shading nets revealed that seedling survival rates followed a similar unimodal pattern, with the highest survival (ca 60%) detected in moderate shade (70%), twice as much as in full sun, and the lowest survival (ca 10%) observed in extreme shade (90%). These results support the study hypothesis on age-dependent interactions. Thus, in a semi-arid shrubland ecosystem, the transition of the "nurse shrub" from "young" to "old" stage can shift facilitation to interference. Hence, the age structure of established shrub populations determines a) the availability of suitable sites for seedling recruitment and b) the balance between facilitation versus interference effects on seedling establishment.     

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.

     

Rapid root extension during water pulses enhances establishment of shrub seedlings in the Atacama Desert

Questions: (1) What is the water threshold for shrub seedling establishment in arid scrubland? (2) How do seedling root growth morphological traits affect the water threshold required for seedling establishment? Location: Arid scrubland, Atacama Desert, north‐central Chile. Methods: We conducted a field experiment with nine native shrub species under a gradient of simulated rainfall to test if species with deep root architecture have higher seedling survival rates and establish more successfully during water pulses. Results: Seedling survival rate was very low; only 2% of the 12 150 planted seedlings survived the first summer drought. Seedling survival required at least 206 mm of water, which is twice the average rainfall and roughly equivalent to the precipitation during an ENSO (El Niño Southern Oscillation) event in this region. Seedling survival at the onset of the summer drought was best explained by leaf habit, root length and initial seedling size. However, only Senna cumingii seedlings survived through the first year. S. cumingii seedlings had distinctively longer roots than the other shrub species, enabling them to reach moister soil layers. Conclusions: Water conditions resembling rainy years enhance shrub seedling establishment in the Atacama Desert, but the effects of higher water availability are strongly dependent on the shrub species. Rapid and deep rooting appears to be a very important functional trait for successful first‐year survival in this arid system where water availability is highly episodic.     

Shrub–tree interactions and environmental changes drive treeline dynamics in the Subarctic

Treelines have drawn persistent research interest as they can respond markedly to climate. However, the mechanisms that determine tree seedling recruitment and the response of the forest‐tundra ecotone to environmental changes remain poorly understood. We hypothesise that treeline tree seedling performance depends on the interplay between climatic and soil nutritional changes and facilitative and competitive interactions between trees and shrubs. We conducted a seedling transplantation experiment with Betula pubescens at a subarctic treeline, in northern Sweden, which followed a full factorial design with four treatment factors relating to environmental regimes of stress and resource availability: site (forest vs treeline); temperature (+/? passive warming); shrub presence (+/?Vaccinium myrtillus removal); and nutrient availability (+/? NPK addition). During three growing seasons we assessed the establishment and performance of Betula. The experimental manipulations caused highly significant effects on seedling performance. Although Vaccinium enhanced seedling survival and reduced the effects of excessive solar radiation and insect herbivory, the seedlings growing with the shrub had a poorer performance by the end of the experimental period. Also, seedlings in the forest had a poorer performance than those at the treeline. Betula seedlings showed a very pronounced and positive response to passive warming and to nutrient addition, but such effects were more evident at the treeline site and often interacted with the presence of Vaccinium. This experiment shows that shrub–tree interactions are important drivers of subarctic treeline dynamics and that they vary with time and space. Facilitation, competition, herbivory and environmental changes at the tree seedling stage act as important filters in structuring the forest–tundra ecotone. We demonstrate that changes in this ecotone cannot be simply predicted from changing temperature patterns alone, and that complex interactions need to be considered, not only between shrubs and trees, but also with herbivores and between warming and soil nutrient availability.     

Competition by herbs as a limiting factor in shrub invasion in grassland: a test with different growth forms

Abstract. We tested the hypothesis that seedling establishment, the critical stage in the invasion of grassland by shrubs, is limited by competition with perennial grasses in seasonally wet/dry savannas. We placed seeds of two invasive exotic shrubs – Cryptostegia grandiflora, a woody vine, and Acacia nilotica, an arborescent legume – into pots with a wide range of existing above- and below-ground herbaceous biomass provided by either a tussock or a stoloniferous perennial grass. We also imposed different levels of watering frequency (5, 10 and 21 d), nutrient addition (+ and -) and grass clipping intensity (no clipping, clipped to 5 cm and clipped to 25 cm). There was no effect of any treatment on shrub seedling emergence or survival and all of the seedlings that emerged survived the 90-d growing period. Herbaceous competition also failed to have an effect on biomass accumulation in shrub seedlings. More frequent watering significantly increased above- and below-ground biomass accumulation for both shrub species and nutrient addition significantly increased Cryptostegia biomass accumulation. Based on these results, we question the proposition that reduction in competition by herbs via livestock grazing has been a significant factor in determining the rate or pattern of exotic shrub increase in the seasonally wet/dry tropics. We also question the suitability of the two-layer soil moisture hypothesis as a basis for management practices to control the ingress of woody species into grasslands and open savannas.