Pinus monophylla establishment in an expanding Pinus‐Juniperus woodland: Environmental conditions,facilitation and interacting factors

Abstract. The tree species comprising Pinus‐Juniperus woodlands are rapidly expanding into shrub‐grasslands throughout their range. Observational studies indicate that establishment is facilitated by nurse plants, but little information exists on the mechanisms involved. I examined both abiotic and biotic factors influencing Pinus monophylla establishment in Artemisia tridentata steppe with expanding populations of P. monophylla and Juniperus osteosperma. I determined soil water contents, temperatures, and nutrient characteristics for the primary establishment microhabitats, i.e. under Pinus, under Juniperus, under Artemisia, tree interspace and sage interspace, and evaluated the emergence and survival response of two seedling cohorts over a 3‐yr period for the different microhabitats. I also examined the effects of seed burial and predation on seedling establishment. Microhabitats under trees and shrubs had higher extractable P and K, higher organic matter, total nitrogen and cation exchange capacity than interspace microhabitats. Soil water contents (0–15 cm) were lower in interspaces than under shrubs or trees due to dry surface (0–5 cm) soils. Soil temperatures (at 1 and 15 cm) were lowest under trees, intermediate under shrubs, and highest in interspaces. Timing and rate of seedling emergence were temperature dependent with the order of emergence paralleling mean growing season temperatures: tree interspace = shrub interspace > under shrub > under Juniperus under Pinus. Seed burial was required for rooting and the highest emergence occurred from depths of 1 and 3 cm indicating that caching by birds and rodents is essential and that animals bury seeds at adequate if not optimal depths for emergence. Seedlings required microenvironmental modification for survival; all seedlings, including those that emerged from seeds and transplants, died within the first year in interspace microhabitats. Survival in under‐tree or under‐shrub microhabitats depended on soil water availability and corresponded closely to soil water contents over the 3‐yr study. Under‐shrub microhabitats had more favourable soil and micro‐environmental characteristics than under‐tree microhabitats and had the highest seedling life spans for the first‐year seedling cohort. Predation of Pinus seedlings by rodents was a significant cause of mortality with caged transplants exhibiting life spans that were 74% longer overall than uncaged transplants. Emergence and survival of P. monophylla within the expanding woodland were dependent upon a complex set of interacting factors including growing season conditions, microhabitat characteristics, and animal species.     

Effects of pioneer shrubs on the recruitment of the fleshy‐fruited tree Olea europaea ssp. cuspidata in Afromontane savanna

Question: Is seedling recruitment of a fleshy‐fruited tree in degraded Afromontane savanna dependent on shelter from pioneer shrubs, and is shelter availability related to shrub traits? Location: Degraded montane savanna in northern Ethiopia (13°36′N, 39°21′E). Method: Nurse plants of Olea europaea ssp. cuspidata seedlings were recorded using T‐square plotless sampling and clustered according to shrub traits, using Ward's method after Principal Components Analysis. Facilitation was further examined through experimental planting and Kaplan‐Meier survival analysis. Results: Both in grazed and protected areas, Olea recruits were found exclusively under shrubs, primarily under Euclea racemosa although Acacia etbaica was more abundant. Olea recruitment is distributed randomly at landscape scale, but depends on shelter at patch scale. Shelter ability is related to shrub shape and species identity. Dense multi‐stemmed shrubs with a wide base and crown on a mulch‐rich mound are key recruitment foci. Euclea shrubs have these favoured traits and probably act as preferential perching sites for avian seed dispersers. Soil and organic matter accumulation under Euclea shrubs may also create favourable conditions for Olea germination and survival. Experimentally planted seedlings had a better chance for survival under Euclea. Conclusions: Olea regeneration is probably subject to both passive (disperser‐mediated) and active facilitation. Small changes of shrub traits can alter the suitability of a patch for Olea recruitment. Protection of shrubs can increase facilitation for seedlings, while pruning may reduce competition for saplings and thus enhance forest succession. Planting of raised Olea seedlings under Euclea shrubs in years with a good rainy season may further assist forest restoration.     

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.     

Restoring Quercus pyrenaica forests using pioneer shrubs as nurse plants

Question: How to improve reforestation success of Quercus pyrenaica. Location: 1800 m a.s.L, southern Spain. Methods: One‐year‐old Quercus pyrenaica seedlings were planted using two treatments: (1) bare soil, using a 30‐cm diameter augur bit (conventional technique) and (2) under the canopy of a pioneer shrub, Salvia lavandulifolia, using a 12‐cm diameter augur bit. Survival and growth were monitored for six years. Our hypothesis is that the use of shrubs as nurse plants is an alternative technique of reforestation with higher success than traditional techniques, in which pre‐existing vegetation is usually considered a source of competition. The rationale for the study was that for environments with a dry season, pre‐existing vegetation buffers summer drought stress, ameliorates the water status of seedlings and thus usually increases seedling recruitment. Results: Quercus survival was 6.3 × higher when planted under individuals of the pioneer shrub as compared to open areas. Quercus seedlings under shrubs also had shoots 1.8 X longer, while the number of shoots per plant did not differ among treatments. The first summer was the period with the highest mortality (49.1% of seedlings). Summer drought was the main cause of mortality. Conclusions: The use of shrubs as nurse plants for Q.pyrenaica reforestation is a viable technique to increase establishment success. The technique could be similarly useful in other environments with a dry period and for other Quercus species. In addition, this technique offers the advantage of following natural succession, thus minimizing the impact in the community.     

Facilitation and interference underlying the association between the woody invaders Pyracantha angustifolia and Ligustrum lucidum

Questions: 1. Is there any post‐dispersal positive effect of the exotic shrub Pyracantha angustifolia on the success of Ligustrum lucidum seedlings, as compared to the effect of the native Condalia montana or the open herbaceous patches between shrubs? 2. Is the possible facilitation by Pyracantha and/or Condalia related to differential emergence, growth, or survival of Ligustrum seedlings under their canopies? Location: Córdoba, central Argentina. Methods: We designed three treatments, in which ten mature individuals of Pyracantha, ten of the dominant native shrub Condalia montana, and ten patches without shrub cover were involved. In each treatment we planted seeds and saplings of Ligustrum collected from nearby natural populations. Seedlings emerging from the planted seeds were harvested after one year to measure growth. Survival of the transplanted saplings was recorded every two month during a year. Half of the planted seeds and transplanted saplings were cage‐protected from rodents. Results: Ligustrum seedling emergence did not differ among treatments while growth was significantly higher in the absence of shrub cover. Sapling survival was significantly higher under the canopy of Pyracantha, intermediate under Condalia, and lowest in the absence of shrub cover. Caging did not affect growth but enhanced seedling emergence and sapling survival. Conclusion: The differential sapling survival in the shrub canopy treatments is consistent with natural sapling distribution. Pyracantha and, less so, Condalia, has a nurse‐plant effect on Ligustrum. This results from contrasting effects of the shrubs on different stages of the life cycle of Ligustrum: no effect on seedling emergence, negative on seedling growth, and positive on sapling survival. This suggests that efforts to control the expansion of Ligustrum over the landscape should tackle Pyracantha as well.     

Canopy vs. soil effects of shrubs facilitating tree seedlings in Mediterranean montane ecosystems

Abstract. Question: Is the facilitative effect of nurse shrubs on early recruitment of trees mediated by a ‘canopy effect’(microclimate amelioration and protection from herbivores), a ‘soil effect’(modification of soil properties), or both? Location: Two successional montane shrublands at the Sierra Nevada Protected Area, SE Spain. Method: Seedlings of Quercus and Pinus species were planted in four experimental treatments: (1) under shrubs; (2) in open interspaces without vegetation; (3) under shrubs where the canopies were removed; (4) in open interspaces but covering seedlings with branches, mimicking a shrub canopy. Results: Both effects benefited seedling performance. However, microclimatic amelioration due to canopy shading had the strongest effect, which was particularly pronounced in the drier site. Below‐ground, shrubs did not modify soil physical characteristics, organic matter, total N and P, or water content, but significantly increased available K, which has been shown to improve seedling water‐use efficiency under drought conditions. Conclusions: We propose that in Mediterranean montane ecosystems, characterised by a severe summer drought, pioneer shrubs represent a major safe site for tree early recruitment during secondary succession, improving seedling survival during summer by the modification of both the above‐and below‐ground environment.     

Shrub canopies influence soil temperatures but not nutrient dynamics: An experimental test of tundra snow–shrub interactions

Shrubs are the largest plant life form in tundra ecosystems; therefore, any changes in the abundance of shrubs will feedback to influence biodiversity, ecosystem function, and climate. The snow–shrub hypothesis asserts that shrub canopies trap snow and insulate soils in winter, increasing the rates of nutrient cycling to create a positive feedback to shrub expansion. However, previous work has not been able to separate the abiotic from the biotic influences of shrub canopies. We conducted a 3‐year factorial experiment to determine the influences of canopies on soil temperatures and nutrient cycling parameters by removing ~0.5 m high willow (Salix spp.) and birch (Betula glandulosa) shrubs, creating artificial shrub canopies and comparing these manipulations to nearby open tundra and shrub patches. Soil temperatures were 4–5°C warmer in January, and 2°C cooler in July under shrub cover. Natural shrub plots had 14–33 cm more snow in January than adjacent open tundra plots. Snow cover and soil temperatures were similar in the manipulated plots when compared with the respective unmanipulated treatments, indicating that shrub canopy cover was a dominant factor influencing the soil thermal regime. Conversely, we found no strong evidence of increased soil decomposition, CO₂ fluxes, or nitrate or ammonia adsorbtion under artificial shrub canopy treatments when compared with unmanipulated open tundra. Our results suggest that the abiotic influences of shrub canopy cover alone on nutrient dynamics are weaker than previously asserted.     

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.     

毛乌素沙地优势灌丛对两种松树幼苗的保育作用

选择毛乌素沙地优势灌丛柠条(Caragana intermedia)、紫穗槐(Amorpha fruticosa)和沙柳(Salix psammophila)作为保育植物,选择2年生油松(Pinus tabulaeformis)和樟子松(Pinus sylvestris)幼苗作为目标植物,将其分别种植在开阔地、柠条、紫穗槐和沙柳4种微生境下,连续3a监测油松和樟子松幼苗的存活和生长动态,从而确定毛乌素沙地优势灌丛是否对植物幼苗的定居有促进作用。结果表明:柠条、紫穗槐和沙柳冠层下光照强度、大气温度和土壤温度都显著低于开阔地,致使土壤湿度增高,油松幼苗总存活率均显著高于开阔地,而樟子松幼苗总存活率仅紫穗槐冠层下显著高于开阔地(P0.05)。紫穗槐冠层下油松和樟子松幼苗总存活率均最高(P0.05),外来种紫穗槐可选为保育植物。保育灌丛下松树幼苗的主枝生长良好,并未受到抑制,但第1年主枝生长长度最小。光照强度和土壤湿度是决定主枝生长长度的重要因素。保育灌丛高度和冠幅面积与油松和樟子松幼苗存活率呈显著正相关(P0.05)。固氮灌丛(紫穗槐和柠条)对松树幼苗存活有持续的促进作用,但柠条冠幅结构紧密,可能因为对光的竞争,减弱了促进作用。沙柳灌丛下松树幼苗的存活率较低。油松幼苗具有一定程度的耐阴性,樟子松幼苗喜光性强,保育灌丛下油松幼苗存活率显著高于樟子松幼苗(P0.05)。总的来说,保育灌丛对松树幼苗的定居有促进作用,保育植物技术可作为一种有效的恢复措施,应用在毛乌素沙地的植被恢复中来。但是,保育灌丛的选择要考虑是本土种或外来种,以及灌丛的高度、冠幅大小、冠幅结构和化学效应,恢复物种的选择要考虑其自身的生物学特性及其与保育植物的相互作用。     

Use of Shrubs as Nurse Plants: A New Technique for Reforestation in Mediterranean Mountains

Common techniques currently used for afforestation in the Mediterranean basin consider the pre‐existing vegetation (mainly shrubs) as a source of competition for trees, and consequently it is generally eliminated before planting. Nevertheless, it has been demonstrated that woody plants can facilitate the establishment of understory seedlings in environments that, like the Mediterranean area, are characterized by a pronounced dry season. In this study, we experimentally analyze the usefulness of shrubs as nurse plants for afforestation of two native conifers, Pinus sylvestris L. (Scots pine) and Pinus nigra Arnold (black pine). Two‐year‐old seedlings were planted in four microhabitats: (1) open interspaces without vegetation (which is the usual method used in afforestation programs), (2) under individuals of Salvia lavandulifolia, (3) under the north side of spiny shrubs, and (4) under the south side of spiny shrubs. Pine survival was remarkably higher when planted under individuals of the shrub S. lavandulifolia (54.8% for Scots pine, 81.9% for black pine) compared with open areas (21.5% for Scots pine, 56.8% for black pine; chi square, p < 0.05). The survival of both pines was also higher when planted on the north side of spiny shrubs, although the survival on the south side was similar to that found in open areas. In addition, pine growth was not inhibited when planted in association with shrubs. This pattern appears to result from the combination of abiotic conditions imposed by the presence of a nurse shrub, which leads to improvement in seedling water status and therefore reduced summer mortality by drought. The results show that the use of shrubs as nurse plants is a technique that offers both economic and ecological advantages, in terms of savings in labor and plant material and reduced and even negligible impact on the pre‐existing vegetation.     

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.     

Resource heterogeneity generated by shrubs and topography on coastal sand dunes

In early stages of primary succession, colonizing plants can create resource patches that influence the abundance and distribution of other species. To test whether different colonizing shrubs generate contrasting patches on coastal sand dunes, we compared soil characteristics and light availability under the nitrogen-fixing shrub Lupinus arboreus, under the non-nitrogen-fixing shrub Artemisia pycnocephala, and between shrubs on dunes at a site in northern California. Concentrations of inorganic nitrogen and net nitrogen mineralization rates were generally 1–10 times greater in soil under Lupinus than under Artemisia or between shrubs. Soil water content was mostly lower under shrubs. Mean photon flux density near ground level was reduced by at least 80% at 35 cm inside shrub canopies. Topography appeared to have more effect on soil moisture but less direct effect on nitrogen availability than did Lupinus. However, Lupinus probably increases nitrogen levels more on higher, drier dunes. Microhabitats under and between nitrogen-fixing shrubs constitute a mosaic of individually poor but complementary patches in which high levels of light and moderate levels of soil nitrogen are present but tend not to occur together.     

The importance of shrub cover for four cactus species differing in growth form in an Andean semi‐desert

Question: Factors influencing seedling establishment are known to vary between open sites and those protected by plant cover. In many desert regions, protected microhabitats below shrubs are essential for establishment of many cactus species. Very little is known about these factors for Andean cacti and how the importance of vegetation cover varies with cactus species. Are Andean cacti associated more frequently to vegetation cover than to open ground? Are they associated to certain shrub species? Is the distributional pattern in relation to cover similar for different cactus species? In what microhabitat (below or away from shrubs) are cactus seeds more abundant? These questions are addressed for the case of an Andean semi‐desert. Location: Semi‐arid tropical Andes, La Paz department, Bolivia. Methods: We examined 132 isolated shrubs = 50 cm along a line across two microhabitats: areas below and away from shrubs/trees. Shrub crown size was measured. The among‐shrub samples were taken from open spaces contiguous to each of the sampled shrubs. In both microhabitats, all cactus species were recorded. The cardinal direction of the cacti was also registered. Correlation between canopy diameter and number of beneficiaries was evaluated for Prosopis flexuosa. The cactus seed bank in each microhabitat was also studied. Results and Conclusions: The four cactus species found behaved differently in relation to shrub canopies. These distributional differences could be due to differences in growth form. Columnar cacti apparently need the shade of shrubs. Only the columnar species is able to grow near the base of the tallest nurse species. The opuntioid cacti studied seem more facultative: although apparently preferring shrub un‐der‐canopies, they are able to establish in open ground. The globose cactus is the most indifferent to the presence of plant cover. These patterns parallel others found in North America. The capacity of different cacti to appear in open spaces could be related to vegetative propagation, and not necessarily to seedling tolerance of heat.     

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.     

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.     

Improving restoration success through microsite selection: an example with planting sagebrush seedlings after wildfire

Post‐fire restoration of foundation plant species, particularly non‐sprouting shrubs, is critically needed in arid and semi‐arid rangeland, but is hampered by low success. Expensive and labor‐intensive methods, including planting seedlings, can improve restoration success. Prioritizing where these more intensive methods are applied may improve restoration efficiency. Shrubs in arid and semi‐arid environments can create resource islands under their canopies that may remain after fire. Seedlings planted post‐fire in former canopy and between canopies (interspace) may have different survival and growth. We compared planting Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young) seedlings post‐fire in former sagebrush canopy and interspace microsites at five locations. Four growing seasons after planting, seedling survival was 46 and 7% in canopy and interspace microsites, respectively. Sagebrush cover was 5.8 times greater in canopy compared to interspace microsites. Sagebrush survival and cover were likely greater because of less competition from herbaceous vegetation as well as benefiting from resource island effects in canopy microsites. Initially, post‐fire abundance of exotic annual grasses was less in canopy microsites, but by the third year post‐fire it was substantially greater in canopy microsites, indicating that resource availability to seedlings was greater, at least initially, in canopy microsites. These results suggest microsites with greater likelihood of success should be identified and then utilized to improve restoration success and efficiency. This is important as the need for restoration greatly exceeds resources available for restoration.     

Alleviation of Summer Drought Boosts Establishment Success of <Emphasis Type="Italic">Pinus sylvestris</Emphasis> in a Mediterranean Mountain: An Experimental Approach

We performed an irrigation experiment to study the impact of summer drought on Pinus sylvestris establishment at its southernmost distribution limit. Watering was done during the first growing season simulating mesic summer conditions, and we monitored the consequences for survival and growth during the first growing season and the delayed consequences on the second growing season. In addition, we considered the heterogeneity created by the microhabitats, where seeds are found after dispersal (bare soil, under shrubs, and under adult pines). Summer drought was the main mortality factor in all the microhabitats. Watering increased emergence and doubled seedling survival compared to non-watered control sites. Differences were even higher when the cumulative effect on emergence and survival was considered, with an overall recruitment of 22.4% in watered plots vs. 7.9% in control. Irrigation increased growth in bare soil and under shrubs, but had scant effect on growth under pines, suggesting that radiation was the limiting factor in this microhabitat. The positive effect of irrigation on growth parameters persisted during the second growing season despite water was not added the second year, showing delayed consequences of drought on seedling performance. Summer drought thus limits Pinus sylvestris establishment in these southernmost forests by reducing both recruitment and growth. This might lead to the development of a remnant dynamic in these relict populations under the current regional increase in dryness and rainfall variability associated with global warming.     

Litter decomposition in a sandy Monte desert of western Argentina: Influences of vegetation patches and summer rainfall

Abstract: We tested the hypothesis that shrub canopies interact with monthly rain pulses to control litter decomposition in a sandy Monte desert, in Argentina. We assessed (i) the potential for litter decomposition of soils beneath the canopies of two dominant shrub species (Larrea divaricata and Bulnesia retama, Zygophyllaceae R. Br.) and from bare‐ground microsites or ‘openings’; (ii) litter decomposition at different spatial patches over the summer rainy season; and (iii) the interaction between vegetation patches and monthly rain pulses on short‐term litter decomposition, or decomposition pulses. In a greenhouse experiment, we found buried litter decomposition to be higher in soils from under the canopies of a dominant shrub species compared with soils from openings and sterilized controls. This, and higher nutrient concentration under shrub soils, suggest undercanopy soils may support a microbial community capable of decomposing litter at higher rates than soils in bare openings. However, ?eld trials showed that shrub patches did not affect leaf litter decomposition over the rainy season, at least for short periods. We found an interaction between shrub patches and incubation time at the end of the ?eld experiment, with higher litter decomposition rates under B. retama canopies. In a monthly ?eld experiment, we found monthly rain pulses signi?cantly explained decomposition pulses, irrespective of patch type. Our ?ndings support the hypothesis that shrub soils have a greater potential for litter decomposition, but this is not directly translated to the ?eld possibly due to interactions with abiotic factors. Rain pulses create conditions for decomposition pulses to occur at shorter time scales, whereas rainfall may interact with a dominant shrub undercanopy to control litter mass loss over longer time scales.     

Factors Limiting the Survival of Native Tree Seedlings Used in Conservation Efforts at the Edges of Forest Fragments in Upland Madagascar

We tested four reforestation techniques in tropical forest fragments that were damaged by fire in upland Madagascar. We conducted a full‐factorial experiment on the survival of transplanted seedlings of five native tree species in grassland plots adjacent to the forest fragments in the Ambohitantely Forest Reserve. The species studied were Dodonaea madagascariensis, Filicium decipiens, Olea lancea, Podocarpus madagascariensis, and Rhus taratana. A total of 480 seedlings were planted; 207 survived the 15 months of the experiment. The factors examined were distance of the reforestation plots from the forest, mixing of forest soil into the plots, application of chemical fertilizers, experimental shading of plots, and the cover of naturally establishing shrubs. Both increasing the distance of plots from the forest edge and adding chemical fertilizers significantly reduced the survival of all seedlings. The surprising negative effects of fertilization may be partly due to increased competition from naturally establishing shrubs that are adapted to exploit high nitrogen levels. Mixing soil from the forest areas into the plots did not change seedling survival. Shading reduced the survival of D. madagascariensis seedlings and did not increase the survival of any species. These findings suggest that the success of reforestation projects can be increased by planting seedlings close to the existing forest fragments. Reforestation of similar tropical forests is likely to be more successful if efforts are focused on expanding the size of existing fragments of tropical forest, rather than on establishing new fragments in grassland openings.     

Multiple effects of shrubs on annual plant communities in arid lands of South Australia

Abstract The presence of shrubs in arid lands creates spatial heterogeneity that affects the distribution and performance of annual plants; several possible mechanisms have been implicated. A preliminary survey in a chenopod shrubland in South Australia showed differences in the distribution of annual plants under canopies of Atriplex vesicaria and Maireana sedifolia (the two dominant shrub species) and open spaces. A series of experiments were conducted to test the potential contribution to these patterns of nutrient enrichment under shrubs, differential seed accumulation, stress reduction by the canopy, competition by shrub roots, and protection against grazing. The germinable soil seed‐bank under A. vesicaria and M. sedifolia was different from that in open spaces, but these differences can only explain a fraction of the differences observed in the growing annual plant community in different microsites. The soil under A. vesicaria had higher total nitrogen content than soil in open spaces, whereas soil under M. sedifolia had lower available phosphorus than open spaces. Although annual plant densities under A. vesicaria were higher than in open spaces, experimental removal of shrubs increased their density, suggesting that shrub canopies inhibit annual plants in this system. Surprisingly, trenching of open areas close to shrubs (severing lateral shrub roots) decreased annual plant density. We suggest that water moves laterally through shrub roots, in a process akin to a hydraulic lift, increasing water availability for the annual plants. Exclusion of vertebrate grazers had a stronger effect on annual plant biomass in open spaces than under M. sedifolia, suggesting that this shrub provides shelter against herbivory. Overall our results show that shrubs can have simultaneously facilitative and inhibitory effects on the annual plant community through different mechanisms, but more importantly that different shrub species have different effects. This is a potential mechanism allowing for species coexistence of annual plants.