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.     

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.     

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.     

Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Question: Insufficient tree regeneration threatens the long‐term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance‐related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.     

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。     

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.     

Shrub influences on seedling performance when restoring the slow‐growing conifer Pilgerodendron uviferum in southern bog forests

Forest restoration is most efficient if it can take advantage of facilitative interactions between established vegetation and planted trees. However, positive and negative interactions have been identified in a number of plant communities. After centuries of anthropogenic fires, forest recovery has been extremely slow in southern bog forests previously dominated by the slow‐growing and vulnerable conifer Pilgerodendron uviferum on Chiloé Island, Chile. Today, the landscape is dominated by secondary shrublands with scattered patches of Sphagnum moss and limited natural tree regeneration. We hypothesized that the retention of secondary shrubs facilitates the early performance of P. uviferum restoration plantings by providing better microsite conditions. To test this hypothesis, we compared the response of seedlings planted on sites prepared at two levels of intervention: after shrubs had been removed or where shrubs were retained. Shrub retention showed a nurse‐plant effect on P. uviferum seedlings 4 years after planting, which resulted in reduced physiological stress (measured as Fv/Fm) for seedlings, as well as reduced browsing. Consequently, the seedlings growing in areas with shrub retention had larger height increment and higher vitality than those in areas where shrubs had been removed. Thus, the more open micro‐site conditions created by shrub removal resulted in generally poorer seedling performance, although seedling mortality—which was low overall (approximately 2–4%)—showed no significant difference between the two levels of intervention. These findings have direct implications for the restoration of slow‐growing conifers that can tolerate extreme wet conditions in highly degraded forests.     

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.     

A Real-Time Quantitative PCR Assay for the Detection of Sphaeropsis sapinea from Inoculated Pinus nigra Shoots

Pinus nigra (Austrian pine) is a tree widely planted in central and southern Europe in the reforestation of areas with poor soils. One of the major constrains of P. nigra is a fungal disease caused by Sphaeropsis sapinea which causes shoot tip dieback and tree mortality. This paper describes a real‐time quantitative PCR assay based on the partial sequence of the small subunit ribosomal RNA of S. sapinea to detect and quantify DNA of the fungal pathogen in S. sapinea inoculated shoots. The study found that real‐time quantitative PCR was a reliable technique to diagnose this disease and suggests that it can also be used to study fungal behaviour in host tissue and particularly to quantify fungal growth in the latent or endophytic phase.     

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.     

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.     

Facilitation and herbivory during restoration of California coastal sage scrub

Using nurse plants to facilitate native plant recruitment in degraded habitats is a common restoration practice across various arid and semiarid environments. Living trees or shrubs are typically considered nurse plants, whereas dead shrubs left in the landscape from prolonged drought are understudied prospective facilitators for native plant recruitment. The interaction between nurse plants and biotic pressures, such as herbivory, on native recruitment is also not well understood in semiarid plant communities. We investigated the effects of facilitation and herbivory on native seedling germination, growth, and survival in the restoration of degraded coastal sage scrub (CSS) habitat. Native shrub seedlings (Artemisia californica and Salvia mellifera) were planted, and native annual species (Amsinckia intermedia, Deinandra fasciculata, Phacelia distans, and Pseudognaphalium californicum) were sown in three Shrub Type treatments (live shrub, dead shrub, and exposed areas), with a nested Cage treatment (no cage and cage) in each Shrub Type treatment. Annual species grew equally well in all Shrub Type treatments; shrub seedlings grew largest in exposed areas. While there was little evidence of facilitation for all species tested, there were strong positive effects of caging on growth and establishment of all species. Caging palatable native species or planting species with anti‐herbivory traits around target plants may be more strategic approaches compared to using nurse plants in restoring degraded CSS after extended drought.     

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.     

Facilitation of Nothofagus antarctica (Fagaceae) seedlings by the prostrate shrub Empetrum rubrum (Empetraceae) on glacial moraines in Patagonia

Abstract Although positive interactions among plants are believed to be common in primary succession, they have rarely been demonstrated on glacial moraines. In Patagonia, the prostrate shrub Empetrum rubrum is dominant in early succession in recently deglaciated valleys, eventually being replaced by the small tree Nothofagus antarctica. This study experimentally evaluates the effects of E. rubrum shrubs on density, biomass, growth and survival of N. antarctica seedlings. The initial density of the seedlings was significantly higher under the canopy of Empetrum than in open areas. A removal experiment was performed to evaluate the influence of Empetrum on survival and growth of seedlings that germinated naturally beneath the canopy of this species. The shrub cover was removed from half of a sample of randomly selected seedlings, and left intact above the remainder. One year later, seedlings with an intact cover of E. rubrum showed significantly higher survival, growth and leaf number than seedlings which had been exposed. The results suggest that the presence of E. rubrum facilitates the establishment of N. antarctica in post‐glacial succession, mainly as a result of more favourable microclimatic conditions under the shrub canopy.     

From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.     

Shrub succession and invasibility in a New Zealand montane grassland

Abstract Two successive shrub invasions of a short tussock grassland induced by grazing and burning were examined in montane South Island, New Zealand. The first invasion was by a native shrub, matagouri (Discaria toumatou Raoul). The second invasion was by an exotic shrub, Scotch broom (Cytisus scoparius (L.) Link), which invaded the matagouri shrubland that had developed over the grassland. The invasions were investigated using analysis of spatial patterns of both shrubs and tussocks, and age, growth rates and size structure of the shrubs. Competition between the two shrub species was examined using spatial patterns and comparing allometric relationships. After initial invasion by matagouri of the grasslands, stand density increased by consolidation about its initial colonization points. Current matagouri distribution is often negatively associated with tussocks. Scotch broom occurs most frequently in a dense sward of introduced grasses and occasionally in tussocks in interstices among matagouri shrubs. Despite the palatability of Scotch broom to sheep that graze the site, there was no evidence that the spiny matagouri facilitates invasion by protecting Scotch broom seedlings; rather there was negative association between the shrub species. The two species probably compete for above-ground space. However, diameter and height growth rates of Scotch broom far exceed those of matagouri so Scotch broom is likely to increase in biomass rapidly at the site. The autogenic organization and disturbance history of the resident plant communities have rendered each vulnerable to successive invasions.     

Fine-scale spatial heterogeneity and seed size determine early seedling survival in a desert perennial shrub (<Emphasis Type="Italic">Ericameria nauseosa:</Emphasis> Asteraceae)

In heterogenous landscapes, seeds settle in some microsites more readily than others, independently of whether those microsites are suitable for germination and subsequent growth, and survival of seedlings. Wild seeds and seedlings of Ericameria nauseosa var. oreophila in the sand-dune ecosystem near Mono Lake, California show both concordance and conflict in where seeds are retained and where seedlings flourish. Using a field experiment, we followed performance of seeds and seedlings (i.e., the rate of germination and successful emergence, initial seedling size, seedling growth and survival over first growing season), by planting seeds of known sizes in pre-identified microsites (windward and lee aspects of interspaces between shrubs, under shrub canopies, and under snag canopies, respectively). Heavier seeds were more likely to germinate and emerge successfully and resulted in initially larger seedlings than lighter seeds. These initially larger seedlings subsequently remained larger over the growing season and lived longer than initially smaller seedlings. Independent of seed and initial seedling size, seedlings that germinated under the canopy of adult shrubs grew larger and survived longer than seedlings that germinated in interspaces (open space with little or no vegetation) or under snags (dead adult shrubs). Seedlings grown under the canopy of adult shrubs experienced significantly less solar radiation and wind-deposited sand than seedlings grown in interspace or snag microsites. Sand burial more than 1 cm was lethal for seedlings. The two variables of seed size and microsite type had by far greater impact on eventual growth and survival of seedlings than did aspect, and the effects of the former two variables were independent of each other. This study amplifies the body of work on E. nauseosa at this site demonstrating that the earliest events in the life history of this long-lived perennial shrub appear to persist through time despite the harsh and variable environment.     

Conditional outcomes in plant–herbivore interactions: neighbours matter

Spatial distribution of palatable and unpalatable plants can influence the foraging behaviour of herbivores, thereby changing plant‐damage probabilities. Moreover, the immediate proximity to certain plants can benefit other plants that grow below them, where toxicity or spines act as a physical barrier or concealment against herbivores. This paper presents the results of a multi‐scale experiment performed to test the effect of shrubs as protectors of tree saplings against herbivores and the mechanism involved in Mediterranean ecosystems. We performed a factorial design in two mountain ranges, similar in physiognomy and vegetation, planting saplings of a palatable tree, the maple (Acer opalus subsp. granatense), and an unpalatable tree, the black pine (Pinus nigra), under three different types of shrubs. We considered four experimental microhabitats: highly palatable shrub (Amelanchier ovalis), palatable but spiny shrub (Crataegus monogyna or Prunus ramburii), unpalatable spiny shrub (Berberis vulgaris subsp. australis) and control (gaps of bare soil without shrubs). Three main factors were found to determine the probability of sapling attack: sapling palatability, experimental microhabitat and plot. Palatable saplings (maples) were browsed much more than unpalatable ones (pines). The degree of protection provided by the shrub proved greater as its palatability decreased with respect to sapling palatability, the unpalatable spiny shrub being the safest microhabitat for palatable saplings and bare soil for unpalatable ones. The differences found in number of attacked saplings between plots may be attributable to differences in herbivore pressure. The community context in which interaction takes place, namely the characteristics of the neighbours and the intensity of herbivore pressure, are determining factors for understanding and predicting the damage undergone by a target plant species. The mechanism that best explains these results is associational avoidance of saplings that grow near to unpalatable shrubs. It is necessary to introduce this neighbour effect in theoretical models and food‐web approaches that analyse the plant–herbivore relationships, since it can strongly determine not only the intensity of the interaction, but also the spatial distribution and diversity of the plant community.     

When do nurse plants stop nursing? Temporal changes in water stress levels in Austrocedrus chilensis growing within and outside shrubs

Question: Does the proximity of shrubs affect seasonal water stress of young Austrocedrus chilensis trees (a native conifer of the Austral Temperate Forest of South America) in xeric sites? Location: A. chilensis xeric forest in northwest Patagonia, Argentina. Methods: We examined the dependence of predawn twig water potential on tree development (seedling to adult) and proximity to nurse shrubs during spring and summer. We analysed spatial associations of seedlings, saplings and adult trees with nurse shrubs, and also evaluated if trees affected shrub canopy vitality. Results: Water stress in Austrocedrus trees was affected by shrub presence. Small trees (i.e.<0.5 m in height) growing in the open were most stressed, particularly in summer. Small trees growing within a shrub canopy had low water stress and little change between spring and summer. The opposite trend, however, was true for the medium‐height category (i.e. 0.5‐1.5 m in height); trees in this size category were more stressed when growing within the shrub canopy than in the open. Larger Austrocedrus trees (i.e.>2 m in height) were not affected by shrub presence. Austrocedrus trees were spatially associated with shrubs in all height classes; however, the percentage of living shrub canopy decreased with tree height. Conclusions: In xeric areas of northwest Patagonia, the strength and direction of interactions between A. chilensis and shrubs, in terms of tree water stress, are dynamic and modulated by tree size and environmental conditions. Overall, positive effects of shrubs on early developmental stages appear to be more important than subsequent negative interactions, since nursing effects could generate a spatial association of shrubs and Austrocedrus trees that persists through later successional stages. These findings shed light on mechanisms behind successional changes, and have important conservation and management implications.     

Avian Response to Bottomland Hardwood Reforestation: The First 10 Years

Bottomland hardwood forests were planted on agricultural fields in Mississippi and Louisiana predominantly using either Quercus species (oaks) or Populus deltoides (eastern cottonwood). We assessed avian colonization of these reforested sites between 2 and 10 years after planting. Rapid vertical growth of cottonwoods (circa 2–3 m/year) resulted in sites with forest structure that supported greater species richness of breeding birds, increased Shannon diversity indices, and supported greater territory densities than on sites planted with slower‐growing oak species. Grassland birds (Spiza americana[Dickcissel] and Sturnella magna[Eastern Meadowlark]) were indicative of species breeding on oak‐dominated reforestation no more than 10 years old. Agelaius phoeniceus (Red‐winged Blackbird) and Colinus virginianus (Northern Bobwhite) characterized cottonwood reforestation no more than 4 years old, whereas 14 species of shrub‐scrub birds (e.g., Passerina cyanea[Indigo Bunting]) and early‐successional forest birds (e.g., Vireo gilvus[Warbling vireo]) typified cottonwood reforestation 5 to 9 years after planting. Rates of daily nest survival did not differ between reforestation strategies. Nest parasitism increased markedly in older cottonwood stands but was overwhelmed by predation as a cause of nest failure. Based on Partners in Flight prioritization scores and territory densities, the value of cottonwood reforestation for avian conservation was significantly greater than that of oak reforestation during their first 10 years. Because of benefits conferred on breeding birds, we recommend reforestation of bottomland hardwoods should include a high proportion of fast‐growing early successional species such as cottonwood.