Regrowth and tannin production in woody and succulent karoo shrubs in response to simulated browsing

Allocation of carbon to chemical defences has often been suggested to be a direct response to browsing or grazing by herbivores. This study quantitatively compares total polyphenol and tannin production in response to simulated browsing of three karoo shrubs in order to test this induced defence hypothesis. The three species studied responded to browsing either by rapid regrowth or by increasing polyphenol production in the remaining tissues. The patterns did not follow any phylogenetic relationships but were weakly associated with the palatability of each species. The highly palatable deciduous species Osteospermum sinuatum, which is capable of rapid regrowth, showed no or very low levels of constitutive and browsing-induced total polyphenols, condensed tannins and protein-precipitating tannins. The evergreen sclerophyllous species Pteronia pallens showed a limited regrowth capacity and had intermediate levels of polyphenols, while the evergreen succulent species Ruschia spinosa showed almost no regrowth over the study period. R. spinosa contained the highest constitutive and browsing-induced levels of polyphenols, condensed tannins and protein-precipitating tannins. In two of the species more than one anti-herbivore defence feature co-occur. P. pallens foliage contains both hepatotoxins and polyphenols while R. spinosa has both structural (spines) and chemical defences. Responses of karoo shrubs to simulated browsing are interpreted as the result of passive alterations in plant chemistry rather than as an active defence response to herbivores.     

Vegetation shift from deciduous to evergreen dwarf shrubs in response to selective herbivory offsets carbon losses: evidence from 19 years of warming and simulated herbivory in the subarctic tundra

Selective herbivory of palatable plant species provides a competitive advantage for unpalatable plant species, which often have slow growth rates and produce slowly decomposable litter. We hypothesized that through a shift in the vegetation community from palatable, deciduous dwarf shrubs to unpalatable, evergreen dwarf shrubs, selective herbivory may counteract the increased shrub abundance that is otherwise found in tundra ecosystems, in turn interacting with the responses of ecosystem carbon (C) stocks and CO₂ balance to climatic warming. We tested this hypothesis in a 19‐year field experiment with factorial treatments of warming and simulated herbivory on the dominant deciduous dwarf shrub Vaccinium myrtillus. Warming was associated with a significantly increased vegetation abundance, with the strongest effect on deciduous dwarf shrubs, resulting in greater rates of both gross ecosystem production (GEP) and ecosystem respiration (ER) as well as increased C stocks. Simulated herbivory increased the abundance of evergreen dwarf shrubs, most importantly Empetrum nigrum ssp. hermaphroditum, which led to a recent shift in the dominant vegetation from deciduous to evergreen dwarf shrubs. Simulated herbivory caused no effect on GEP and ER or the total ecosystem C stocks, indicating that the vegetation shift counteracted the herbivore‐induced C loss from the system. A larger proportion of the total ecosystem C stock was found aboveground, rather than belowground, in plots treated with simulated herbivory. We conclude that by providing a competitive advantage to unpalatable plant species with slow growth rates and long life spans, selective herbivory may promote aboveground C stocks in a warming tundra ecosystem and, through this mechanism, counteract C losses that result from plant biomass consumption.     

Mediterranean dwarf shrubs and coexisting trees present different radial-growth synchronies and responses to climate

Due to their diversity and dominance in environmentally harsh sites, Mediterranean dwarf shrubs are a valuable tool to understand the consequences of climatic variability on radial growth in woody plants. We evaluate the dendrochronological potential of three Mediterranean dwarf shrubs versus three coexisting tree species inhabiting cold- (Hormathophylla spinosa vs. Pinus sylvestris), mesic- (Ononis fruticosa vs. Abies alba), and xeric sites (Linum suffruticosum vs. Pinus halepensis). Cross-sectional wood sections of the three shrub species and cores in the case of trees were visually cross-dated and ring-widths were measured and converted into residual growth indices. We used linear mixed-effects models to assess how growth indices respond to local factors and climatic variables. The radial growth of the three dwarf shrub species was more asynchronous, i.e., ring-width series differed among conspecific individuals, than that of coexisting tree species. Growth asynchrony was higher for H. spinosa than for O. fruticosa and L. suffruticosum. Similarly, the ring-width series of O. fruticosa and L. suffruticosum was strongly correlated with that of coexisting tree species, while growth series of H. spinosa and P. sylvestris was not related at all. The growth of the three dwarf shrub species was influenced by the regional climatic conditions, but to a lesser degree than coexisting tree species. The highest responsiveness of growth to climate was observed in Mediterranean dwarf shrubs from xeric sites. However, local conditions are also major drivers of growth in Mediterranean dwarf shrubs as indicated by the stronger asynchrony in ring formation of these species as compared with coexisting trees, particularly in cold sites.     

SPRING GROWTH OF SHOOTS AND ROOTS IN SHRUBS OF AN ALASKAN MUSKEG

Early spring shoot and fine-root development of four evergreen and three deciduous shrub species were analyzed in a subarctic muskeg at Fairbanks, Alaska. The overwintered foliage of the evergreen shrubs regreened earlier than new leaves developed on the deciduous species. Likewise, the evergreen shrubs produced new fine roots earlier than the deciduous species. The total nonstructural carbohydrate (TNC) concentration did not decline in the evergreen shrub Ledum palustre during the spring development. This contrasted with the deciduous shrub Betula glandulosa, where a significant TNC reduction in stem tissue coincided with bud break and fine-root growth flush.     

Dwarf shrub and grass vegetation resistant to long‐term experimental warming while microarthropod abundance declines on the Falkland Islands

Dwarf shrubs are a dominant plant type across many regions of the Earth and have hence a large impact on carbon and nutrient cycling rates. Climate change impacts on dwarf shrubs have been extensively studied in the Northern Hemisphere, and there appears to be large variability in response between ecosystem types and regions. In the Southern Hemisphere, less data are available despite dwarf shrub vegetation being a dominant feature of southern South America and mountainous regions of the Southern Hemisphere. Here, we present the response of an Empetrum rubrum dwarf shrub and a Poa grass community to 12 years of experimental climate manipulation achieved using open top chambers on the Falkland Islands, a cold temperate island group in the South Atlantic. The dwarf shrub and grass vegetation did not change significantly in cover, biomass or species richness over the 12 years period in response to climate warming scenarios of up to 1°C reflecting annual warming levels predicted in this region for the coming decades. The soil microarthropod community, however, responded with declines in abundance (37%) under warming conditions in the grass community, but no such changes were observed in the dwarf shrub community. Overall, our data indicate that dwarf shrub communities are resistant to the levels of climate warming predicted over the coming decades in the southern South America region and will, therefore, remain a dominant driver of local ecosystem properties.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Season and light affect constitutive defenses of understory shrub species against folivorous insects

Understory shrubs contribute to overall species diversity, providing habitat and forage for animals, influence soil chemistry and forest microclimate. However, very little is known about the chemical defense of various shrub species against folivorous insects. Using six shrub species, we tested how seasonal changes and light conditions affect their constitutive defense to insect damage. We monitored leaf perforation, concentrations of total phenols, condensed tannins, nitrogen (N), and total nonstructural carbohydrates (TNC). Leaf damage caused by insects was low in Sambucus nigra, Cornus sanguinea, and Frangula alnus, intermediate in Corylus avellana and Prunus serotina, and high in Prunus padus. Leaves of all the species, when growing in high light conditions, had high concentrations of defense metabolites. Except for C. avellana, leaves of the other shrub species growing in full sun were less injured than those in shade. This may be due to higher concentrations of defense metabolites and lower concentrations of nitrogen. Similar patterns of the effects of light on metabolites studied and N were observed for leaves with varying location within the crown of individual shrubs (from the top of the south direction to the bottom of the north), as for leaves from shrubs growing in full sun and shrubs in the shade of canopy trees. A probable cause of the greater damage of more sunlit leaves of C. avellana was the fact that they were herbivorized mostly by Altica brevicollis, a specialist insect that prefers plant tissues with a high TNC level and is not very sensitive to a high level of phenolic compounds.     

The influence of harvester and Messor capensis nest-mounds on the productivity and distribution of some plant species in the southern Karoo,South Africa

Nest-mounds of the harvester ant Messor capensis occur on and around nutrient-rich patches, along minor drainage lines in nutrient-rich soils, and on the plains, generally in nutrient-poor soils. Nest-site selection is related to the presence of suitable deep soils, the presence of stones and the distance from the nearest neighbouring nest. Two plant species, Galenia fruticosa and Pteronia pallens, were significantly associated with Messor capensis nest-mounds, both in numbers of mounds occupied and in numbers of individuals. A third species, Drosanthemum montaguense, was also more common in numbers of individuals, while a fourth species, Rhinephyllum macradenium was negatively associated with these mounds. The analysis of species guilds by soil type shows that significantly more species of nutrient-rich soils are present on M. capensis nest-mounds. Also, significantly more taller, woody species occurred on nest-mounds than in inter-mound spaces. Two species, Pteronia pallens and Osteospermum sinuatum, growing on ant nest-mounds had significantly longer inter-nodes than the same species growing off mounds. However, two other species, Pteronia cf. empetrifolia and Galenia fruticosa showed no difference in inter-node lengths between plants growing on and off mounds. Six of the nine species of plants sampled on ant nest-mounds had significantly higher seed production than plants of the same species growing in inter-mound spaces. The other three species showed a tendency towards more seeds per plant on ant nest-mounds. The proportions of live and dead plants on mounds differed between species. Only Ruschia spinosa showed a significant difference between the numbers of dead plants in the population on and off mounds, with more dead plants occurring on mounds. Significantly more seeds set on individuals of Pteronia pallens growing on ant nest-mounds than those growing off nest-mounds, but no such difference occurred in P. cf. empetrifolia. There was no significant difference in the proportion of seeds parasitized by the tephritid fly Desmella anceps for individuals of P. pallens and P. cf. empetrifolia growing on and off mounds.     

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.     

Seed dispersal by cattle may cause shrub encroachment of Grewia flava on southern Kalahari rangelands

Abstract. Shrub encroachment, i.e. the increase in woody plant cover, is a major concern for livestock farming in southern Kalahari savannas. We developed a grid‐based computer model simulating the population dynamics of Grewia flava, a common, fleshy‐fruited encroaching shrub. In the absence of large herbivores, seeds of Grewia are largely deposited in the sub‐canopy of Acacia erioloba. Cattle negate this dispersal limitation by browsing on the foliage of Grewia and dispersing seeds into the grassland matrix. In this study we first show that model predictions of Grewia cover dynamics are realistic by comparing model output with shrub cover estimates obtained from a time series of aerial photographs. Subsequently, we apply a realistic range of intensity of cattle‐induced seed dispersal combined with potential precipitation and fire scenarios. Based on the simulation results we suggest that cattle may facilitate shrub encroachment of Grewia. The results show that the severity of shrub encroachment is governed by the intensity of seed dispersal. For a high seed dispersal intensity without fire (equivalent to a high stocking rate) the model predicts 56% shrub cover and 85% cell cover after 100 yr. With fire both recruitment and shrub cover are reduced, which may, under moderate intensities, prevent shrub encroachment. Climate change scenarios with two‐fold higher frequencies of drought and wet years intensified shrub encroachment rates, although long‐term mean of precipitation remained constant. As a management recommendation we suggest that shrub encroachment on rangelands may be counteracted by frequent fires and controlling cattle movements to areas with a high proportion of fruiting Grewia shrubs.     

Spatial patterns in a two-tiered semi-arid shrubland in southeastern Spain

Abstract. Single species and bivariate distribution patterns in a semi-arid shrubland in southeastern Spain, dominated by the tall leguminous shrub Retama sphaerocarpa, were investigated by second-order spatial analysis based on Ripley's K-function. Shrubs were significantly clumped because of a strong association of dwarf shrubs, mostly Artemisia barrelieri, under the canopy of Retama. Retama shrubs were randomly distributed, but when different size-classes were analysed separately, the pattern changed from significantly clumped to random and then to regular with increasing canopy diameter, suggesting increasing intraspecific competition with shrub size. Artemisia was significantly clumped at all scales because of aggregation under the canopy of large Retama shrubs. The association between the species became stronger with increasing canopy diameter of Retama shrubs, suggesting that facilitation prevailed over interspecific competition because of niche separation in different tiers, both above and below ground. Retama shrub size thus determined both the type of pattern for its own size class and tier, and the scale and intensity of the association with its understorey shrubs.     

Field Simulation of Global Change: Transplanting Northern Bog Mesocosms Southward

A large proportion of northern peatlands consists of Sphagnum-dominated ombrotrophic bogs. In these bogs, peat mosses (Sphagnum) and vascular plants occur in an apparent stable equilibrium, thereby sustaining the carbon sink function of the bog ecosystem. How global warming and increased nitrogen (N) deposition will affect the species composition in bog vegetation is still unclear. We performed a transplantation experiment in which mesocosms with intact vegetation were transplanted southward from north Sweden to north-east Germany along a transect of four bog sites, in which both temperature and N deposition increased. In addition, we monitored undisturbed vegetation in control plots at the four sites of the latitudinal gradient. Four growing seasons after transplantation, ericaceous dwarf shrubs had become much more abundant when transplanted to the warmest site which also had highest N deposition. As a result ericoid aboveground biomass in the transplanted mesocosms increased most at the southernmost site, this site also had highest ericoid biomass in the undisturbed vegetation. The two dominant Sphagnum species showed opposing responses when transplanted southward; Sphagnum balticum height increment decreased, whereas S. fuscum height increment increased when transplanted southward. Sphagnum production did not differ significantly among the transplanted mesocosms, but was lowest in the southernmost control plots. The dwarf shrub expansion and increased N concentrations in plant tissues we observed, point in the direction of a positive feedback toward vascular plant-dominance suppressing peat-forming Sphagnum in the long term. However, our data also indicate that precipitation and phosphorus availability influence the competitive balance between Sphagnum, dwarf shrubs and graminoids.     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.     

Fire frequency influences composition and structure of the shrub layer in an Australian subcoastal temperate grassy woodland

Little is known about the relationship between fire regimes and plant diversity in Australia's temperate grassy woodlands. The effect of fire frequency on shrubs in grassy woodland remnants across Western Sydney's Cumberland Plain was examined. Shrub species richness and composition were compared in sites that had experienced a high, moderate or low frequency of fire over the previous 20 years. Nine sites were surveyed, three in each fire frequency category; most sites, including all low‐fire‐frequency sites, had burnt 9–36 months prior to sampling. Fire frequency had a profound effect on the composition and structure of the shrub layer. Per cent frequency and density of the prickly shrub Bursaria spinosa (Pittosporaceae) was considerably higher in low‐fire‐frequency sites than where fires had occurred at least once a decade. In sites where fire had been absent for decades prior to a recent fire, this species dominated the landscape, while elsewhere it occurred as clumps in a grassy matrix. Per cent frequency of other native shrubs, particularly obligate seeders, was greatest at moderate fire frequencies. Exotic shrubs were recorded most often where fire had been rare. While ordination clearly separated out the low‐fire‐frequency sites, complete separation between high‐ and moderate‐fire‐frequency blocks was not achieved. The increase in Bursaria in the absence of fire mirrors the encroachment of woody plants into a range of grassy ecosystems around the world. The sensitivity of obligate seeder species, many of them short‐lived legumes with fire‐cued seeds, to both very frequent and very infrequent fire shows the vulnerability of these species to extreme fire regimes, despite the safeguards conferred by hard‐seededness. Competition from Bursaria, as well as loss of viable seed in the soil, may have contributed to the low frequency of these species after a long inter‐fire interval.     

Inhibition of growth,and effects on nutrient uptake of arctic graminoids by leaf extracts — allelopathy or resource competition between plants and microbes?

Previous research has shown that plant extracts, e.g. from boreal dwarf shrubs and trees, can cause reduced growth of neighbouring plants: an effect known as allelopathy. To examine whether arctic and subarctic plants could also be affected by leaching of phytochemicals, we added extracts from the commonly occurring arctic dwarf shrubs Cassiope tetragona and Empetrum hermaphroditum, and from mountain birch, Betula pubescens ssp. tortuosa to three graminoid species, Carex bigelowii, Festuca vivipara and Luzula arcuata, grown in previously sterilized or non-sterilized arctic soils. The graminoids in non-sterilized soil grew more slowly than those in sterilized soil. Excised roots of the plants in non-sterilized soil had higher uptake rate of labelled P than those in sterilized soil, demonstrating larger nutrient deficiency. The difference in growth rate was probably caused by higher nutrient availability for plants in soils in which the microbial biomass was killed after soil sterilization. The dwarf shrub extracts contained low amounts of inorganic N and P and medium high amounts of carbohydrates. Betula extracts contained somewhat higher levels of N and much higher levels of P and carbohydrates. Addition of leaf extracts to the strongly nutrient limited graminoids in non-sterilized soil tended to reduce growth, whereas in the less nutrient limited sterilized soil it caused strong growth decline. Furthermore, the N and P uptake by excised roots of plants grown in both types of soil was high if extracts from the dwarf shrubs (with low P and N concentrations) had been added, whereas the P uptake declined but the N uptake increased after addition of the P-rich Betula extract. In contrast to the adverse extract effects on plants, soil microbial respiration and soil fungal biomass (ergosterol) was generally stimulated, most strongly after addition of the Betula extract. Although we cannot exclude the possibility that the reduced plant growth and the concomitant stimulation of microbial activity were caused by phytochemicals, we believe that this was more likely due to labile carbon in the extracts which stimulated microbial biomass and activity. As a result microbial uptake increased, thereby depleting the plant available pool of N and P, or, for the P-rich Betula extract, depleting soil inorganic N alone, to the extent of reducing plant growth. This chain of events is supported by the negative correlation between plant growth and sugar content in the three added extracts, and the positive correlation between microbial activity, fungal biomass production and sugar content, and are known reactions when labile carbon is added to nutrient deficient soils.     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

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.     

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.     

The impact of shrub browsing by mountain hare and reindeer in subarctic Sweden

Background: Climate warming has been causing an increase in tall shrub cover around the Arctic, however, mammalian herbivory has been shown to inhibit shrub expansion. Though the effect of reindeer (Rangifer tarandus) and many other mammals has been widely studied in this context, the role of the mountain hare (Lepus timidus) in subarctic Scandinavia remains unknown.

Aims: To quantify browsing from mountain hare and reindeer on tall shrubs in different vegetation types and to investigate differences in shrub preference between the two.

Methods: In the summers of 2013 and 2014, we counted signs of browsing by hare and reindeer on tall shrub species in 31 study plots at three alpine locations in the Scandes range, Sweden.

Results: Hare browsing was significantly more frequent than that by reindeer in two (dry-mesic heath and dry meadow) out of seven vegetation types studied. Reindeer browsing was significantly higher in the low herb meadow and Långfjället shrub heath. Two shrub species, Betula nana and Salix hastata, were significantly more browsed by hare, while reindeer browsing was significantly higher on S. phylicifolia and S. lapponum.

Conclusions: Our results show that mountain hares can cause extensive damage to tall shrubs in the subarctic and may have a stronger impact on shrub communities than previously recognised.     

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.