Ecological benefits of myrmecochory for the endangered chaparral shrub Fremontodendron decumbens (Sterculiaceae)

Fremontodendron decumbens grows in a single county in central California, USA. Prior research showed that its elaiosome-bearing seeds are dispersed by the harvester ant Messor andrei. I tested several hypotheses regarding the positive role of ant-mediated dispersal to F. decumbens: (1) Does ant-mediated seed dispersal facilitate seed escape from rodent predation?; (2) Does ant processing of seeds stimulate germination?; (3) Are ant middens more suitable microsites for seed or seedling survival in unburned chaparral areas?; and (4) Do survival benefits of dispersal occur post-fire in the form of differences in seedling survival probabilities and, if so, why? Results of tests of each hypothesis were: (1) similar percentages of seeds placed on ant middens and under F. decumbens shrub canopies were destroyed by rodents, but seeds from which elaiosomes had been removed were more likely to escape rodent predation; (2) seeds processed by ants did not germinate more readily than seeds removed directly from shrub branches; (3) seedling predation was a major cause of mortality in unburned chaparral on both ant middens and under shrubs, and overall seedling survival did not differ between the two microsites; (4) post-burn seedling survival was significantly greater for seedlings dispersed away from F. decumbens shrub canopies, because dispersed seedlings were both less likely to be killed by predators and more likely to be growing in a gap created by the fire-caused death of an established shrub. I concluded that the major ecological benefit to F. decumbens of ant-mediated seed dispersal was elevated post-fire seedling survival resulting from enhanced escape by dispersed seedlings from both predation and competition.     

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.     

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.     

Legacy microsite effect on the survival of bitterbrush outplantings after prescribed fire: capitalizing on spatial variability to improve restoration

Restoration of shrubs in arid and semi‐arid rangelands is hampered by low success rates. Planting shrub seedlings is a method used to improve success in these rangelands; however, it is expensive and labor intensive. The efficiency of shrub restoration could be improved by identifying microsites where shrub survival is greater. Bitterbrush (Purshia tridentata Pursh DC) is an important shrub to wildlife that has declined because of conifer encroachment, excessive defoliation, wildfires, and low recruitment. We investigated planting bitterbrush seedlings in western juniper (Juniperus occidentalis ssp. occidentalis Hook) encroached shrublands after prescribed fire was used to control trees. Bitterbrush seedlings were planted in under (canopy) and between (interspace) former juniper canopies at five blocks and evaluated for three growing seasons. Bitterbrush survival was greater than 50% in the former canopy, but only 5% in the interspace microsite by the third growing season. Growth of bitterbrush was also greater in the former canopy compared with the interspace, potentially due to markedly less perennial vegetation in this microsite. Exotic annual grasses and annual forbs became prevalent in the former canopy in the second and third growing season, suggesting that soil resource availability was greater in this microsite. These results suggest that restoration success will vary by specific locations within a burned landscape and that this variability can be used to improve restoration efficiency. In this situation, bitterbrush restoration can be improved by planting seedlings in former canopy compared with interspace microsites.     

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.     

Direct and indirect interactions among plants explain counterintuitive positive drought effects on an eastern Mediterranean shrub species

Coexistence of woody and herbaceous plants may be governed by a complex set of direct and indirect interactions, whose relative importance have been rarely assessed. We experimentally studied woody species establishment in a mixed plant community by disentangling the potential role of such biotic interactions and the effect of environmental variations on them. Seedling establishment of the common eastern Mediterranean shrub species Sarcopoterium spinosum was investigated under different rainfall and light conditions, combined with the effect of the presence of adult shrubs and annual neighbors. We predicted that seedlings will be directly affected by competition with annuals with increasing water availability, while direct effects of adult shrubs will be positive via amelioration of water stress. Indirect effects were expected beneath shrub canopies due to reduced water stressed and light availability for both annuals and shrub seedlings, which may intensify competition between annuals and shrub seedlings. To test these predictions we performed field and garden experiments in which we combined manipulation of shrub and annual presence with manipulations of water availability and light conditions to simulate the effect of shrub canopy. In contrast to our prediction, shrub seedling establishment was not facilitated but inhibited by adult shrubs because of light limitation. As expected, annuals had direct negative effects on shrub seedlings under wet conditions, which shifted to neutral or positive effects under dry conditions. Thus, interactions among shrubs and annuals, and in particular the release from competition during drought years, leads to a counterintuitive positive effect of drought on shrub seedling establishment. Our findings point to the importance of experimentally studying multidimensional interactions for coexistence of different life forms and to the underestimated role of light for success in water‐limited ecosystems.     

Shrub emergence and seedling growth in microenvironments created by Prosopis glandulosa

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

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.     

Grass-Shrub Associations over a Precipitation Gradient and Their Implications for Restoration in the Great Basin,USA

As environmental stress increases positive (facilitative) plant interactions often predominate. Plant-plant associations (or lack thereof) can indicate whether certain plant species favor particular types of microsites (e.g., shrub canopies or plant-free interspaces) and can provide valuable insights into whether “nurse plants” will contribute to seeding or planting success during ecological restoration. It can be difficult, however, to anticipate how relationships between nurse plants and plants used for restoration may change over large-ranging, regional stress gradients. We investigated associations between the shrub, Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis), and three common native grasses (Poa secunda, Elymus elymoides, and Pseudoroegneria spicata), representing short-, medium-, and deep-rooted growth forms, respectively, across an annual rainfall gradient (220–350 mm) in the Great Basin, USA. We hypothesized that positive shrub-grass relationships would become more frequent at lower rainfall levels, as indicated by greater cover of grasses in shrub canopies than vegetation-free interspaces. We sampled aerial cover, density, height, basal width, grazing status, and reproductive status of perennial grasses in canopies and interspaces of 25–33 sagebrush individuals at 32 sites along a rainfall gradient. We found that aerial cover of the shallow rooted grass, P. secunda, was higher in sagebrush canopy than interspace microsites at lower levels of rainfall. Cover and density of the medium-rooted grass, E. elymoides were higher in sagebrush canopies than interspaces at all but the highest rainfall levels. Neither annual rainfall nor sagebrush canopy microsite significantly affected P. spicata cover. E. elymoides and P. spicata plants were taller, narrower, and less likely to be grazed in shrub canopy microsites than interspaces. Our results suggest that exploring sagebrush canopy microsites for restoration of native perennial grasses might improve plant establishment, growth, or survival (or some combination thereof), particularly in drier areas. We suggest that land managers consider the nurse plant approach as a way to increase perennial grass abundance in the Great Basin. Controlled experimentation will provide further insights into the life stage-specific effectiveness and practicality of a nurse plant approach for ecological restoration in this region.     

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.     

Rodent-favored cache sites do not favor seedling establishment of shade-intolerant wild apricot (Prunus armeniaca Linn.) in northern China

Seed dispersal by scatter hoarders is widely assumed to involve directed dispersal, in which microhabitats selected for caching also benefit seedling establishment and growth. However, in many systems, this may not be true if animal-favored cache sites do not match the safe sites for plants, or if cache sites benefit the plant in one life stage and not another. Here, we investigated whether cache sites selected by rodents are favorable for seedling establishment and growth of shade-intolerant wild apricot (Prunus armeniaca Linn.) in northern China. We tracked tagged seeds and compared the germination and growth of seedlings from rodent-cached seeds with that of naturally established seedlings in a secondary forest and shrubland stand. Rodents preferred to cache seeds under shrubs with medium canopy cover (31–60 %) in litter substrate in the secondary forest, and under shrubs with high canopy cover (>60 %) in soil or litter substrate in the shrubland stand, neither of which conveyed an advantage for seedling establishment. Although fewer caches were made along shrub edges, or under low canopy cover (≤30 %) in the secondary forest, or along shrub edges, open areas of grass, or under low canopy cover in the shrubland stand, these cache sites consistently contributed to higher survival rates. The microhabitats of grass, soil, or low canopy cover significantly promoted the emergence, survival, and growth of naturally established seedlings. Our results are best explained by the conflicting demands of rodents for caching seeds in more secure sites and P. armeniaca’s high-light requirements for seedling recruitment. We argue that the relationship between favored cache sites for seed hoarders and safe sites for plants will often not match but may still allow a reasonable rate of establishment and regeneration.     

Post‐fire shrub recruitment in a semi‐arid grassland: the role of microsites

Question: (1) Which factors regulate post‐fire recruitment and spread of the shrub Senecio bracteolatus in Patagonian grasslands? (2) What is the role of the grass Stipa speciosa on S. bracteolatus establishment in the post‐fire succession? Location: Northwest Patagonia, Argentina. Methods: We studied the effect of fire on S. bracteolatus recruitment and density by comparing these variables between burned and unburned grasslands. In burned areas, we compared abiotic characteristics and seedling establishment under the canopy of grasses (S. speciosa) and in gaps (inter‐tussock areas). Post‐fire interactions between S. bracteolatus seedlings and S. speciosa were studied using field and greenhouse experiments. Results: Density of S. bracteolatus was higher in burned than in unburned areas. In burned sites, seedlings were more abundant under tussock grasses, whereas juveniles were more abundant in gaps. Tussocks generated more attenuated micro‐environmental conditions than gaps during stressful summers. Gaps were more abundant in burned sites, while “under tussock” microsites were more frequent in unburned sites. In burned areas, tussocks allowed higher establishment of seedlings (facilitation), but gaps allowed more seedling growth and higher persistence of juveniles. Conclusions: Fire promoted S. bracteolatus recruitment in Patagonian grasslands by increasing the availability of favourable gap microsites. Grass protection for shrub seedlings became negative with time, probably due to competition with grasses. Gaps led to better performance and persistence of shrub plants. Six years after fire, higher shrub recruitment and adult density (observed as a trend) in burned grassland provides an opportunity for potential S. bracteolatus invasion.     

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。     

Ecology of Aglaia mackiana (Meliaceae) Seedlings in a New Guinea Rain Forest1

The large seeds of Aglaia mackiana (Meliaceae) germinate and produce vigorous seedlings under closed canopies or in large gaps. To assess seedling ecology after germination, we measured growth, herbivore damage, and survivorship of seedlings over one year. The sample included shaded seedlings from dispersed seeds, undispersed seeds under parent trees, and seedlings transplanted to gaps. We quantified the light environment using hemispherical canopy photographs taken above seedlings at the beginning and end of the one–year study. Seedlings transplanted to gaps grew faster and had more leaves, larger total leaf surface area, longer secondary roots, and greater root mass than shaded seedlings. Seedlings in gaps did not differ from shaded seedlings in survivorship or amount of herbivore– and pathogen–caused leaf damage. The canopy photographs taken one year apart suggest there is a rough equilibrium in closed canopies with slight changes occurring around an average light level. Sites with < 0.06 ISF (a unitless, relative measure of canopy openness or reflected sunlight) tended to remain the same with minor fluctuations toward brighter or darker. Sites with canopy openness > 0.06 ISF tended to close; few gaps grew larger. Seedlings under parenr trees and seedlings away from parent trees had similar amounts of leaf damage and virtually identical survivorship after 18 months, but seedlings under parent trees had slower growth rates and smaller total leaf surface areas. Dispersal did not strongly benefit seeds via escaping high levels of mortality or comperition around the parent.     

Density‐dependent Survival in Seedlings Differs among Woody Life‐forms in Tropical Wet Forests of a Caribbean Island

Negative density dependence contributes to seedling dynamics in forested ecosystems, but the relative importance of this factor for different woody plant life‐forms is not well‐understood. We used 1 yr of seedling survivorship data for woody seedlings in 17 different plots of lower to mid‐montane rain forests on the island of Dominica to examine how seedling height, abiotic factors, and biotic factors such as negative density dependence are related to seedling survival of five different life‐forms (canopy, midstory, and understory trees; shrubs; and lianas). Across 64 species, taller seedlings in seedling plots with higher canopy openness, greater seedling density, lower relative abundance of conspecific seedlings, and lower relative abundance of conspecific adults generally had a greater probability of surviving. Height was the strongest predictor of seedling survival for all life‐forms except lianas. Greater seedling density was positively related to survival for canopy and midstory trees but negatively related to survival for the other life‐forms. For trees, the relative abundance of conspecific seedling and adult neighbors had weak and strong negative effects on survival respectively. Neither shrub nor liana seedling survival was affected by the relative abundance of conspecific neighbors. Thus, negative density dependence is confirmed as an important structuring mechanism for tree seedling communities but does not seem to be important for lianas and shrubs in Dominican rain forests. These results represent the first direct assessment of controls on seedling survival of all woody life‐forms – an important step in understanding the dynamics and structure of the entire woody plant community.     

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.     

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.     

Canopy manipulations of exotic Bitter Willow (Salix elaeagnos) forest for indigenous seedling recruitment: A pilot study

The invasive exotic tree species Bitter Willow (Salix elaeagnos; Salicaceae) has colonised areas of rank exotic grassland and has been found to contain indigenous seed, dispersed by frugivorous birds into the monospecific stands. This small pilot study examined whether indigenous seedlings that have germinated in the understorey of exotic Bitter Willow stands could be stimulated to establish through the creation of small‐scale canopy gaps. In Bitter Willow forest, four single Bitter Willow trees were poisoned to create canopy gaps. Light transmission and seedling regeneration of tree and shrub species were assessed beneath both the four manipulated and three comparable intact Bitter Willow canopies. Over 3 years, seedling height and density increased more beneath opened compared to intact Bitter Willow canopies. These results suggest that Bitter Willow can fill the roles of both a facilitative nurse and a perch tree. Larger‐scale canopy manipulation experiments of both Bitter Willow and other Salix species are needed to determine the full potential of canopy manipulations for forest restoration.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

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.