Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

The Effects of Established Trees on Woody Regeneration during Secondary Succession in Tropical Dry Forests

Understanding the mechanisms controlling secondary succession in tropical dry forests is important for the conservation and restoration of this highly threatened biome. Canopy‐forming trees in tropical forests strongly influence later stages of succession through their effect on woody plant regeneration. In dry forests, this may be complex given the seasonal interplay of water and light limitations. We reviewed observational and experimental studies to assess (1) the relative importance of positive and negative effects of established trees on regeneration; (2) the mechanisms underlying these effects; and (3) to test the ‘stress gradient hypothesis’ in successional tropical dry forests. The effects of established trees on seed dispersal, seed survival, and seed germination—either through direct changes to moisture and temperature regimes or mediated by seed dispersers and predators—are mainly positive. The balance between positive and negative effects on seedling establishment is more complex and depends on the season and leaf phenology of both trees and seedlings. Seedling survival is generally enhanced by established trees mitigating dry conditions. Established trees have counteracting effects on water and light availability that influence seedling growth. The probability of a positive effect of established trees on seedling survival decreases with increased rainfall, which supports the stress gradient hypothesis. Priorities for future research are experiments to test for facilitation and competition and their underlying mechanisms, long‐term studies evaluating how these effects change with ontogeny, and studies focusing on the species‐specificity of interactions.     

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

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

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Climatic warming strengthens a positive feedback between alpine shrubs and fire

Climate change is expected to increase fire activity and woody plant encroachment in arctic and alpine landscapes. However, the extent to which these increases interact to affect the structure, function and composition of alpine ecosystems is largely unknown. Here we use field surveys and experimental manipulations to examine how warming and fire affect recruitment, seedling growth and seedling survival in four dominant Australian alpine shrubs. We found that fire increased establishment of shrub seedlings by as much as 33‐fold. Experimental warming also doubled growth rates of tall shrub seedlings and could potentially increase their survival. By contrast, warming had no effect on shrub recruitment, postfire tussock regeneration, or how tussock grass affected shrub seedling growth and survival. These findings indicate that warming, coupled with more frequent or severe fires, will likely result in an increase in the cover and abundance of evergreen shrubs. Given that shrubs are one of the most flammable components in alpine and tundra environments, warming is likely to strengthen an existing feedback between woody species abundance and fire in these ecosystems.     

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

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

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。     

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

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

Effects of above- and below-ground competition from shrubs on photosynthesis, transpiration and growth in Quercus robur L. seedlings

For a tree seedling to successfully establish in dense shrubbery, it must maintain function under heterogeneous resource availability. We evaluated leaf-level acclimation in photosynthetic capacity, seedling-level transpiration, and seedling morphology and growth to gain an understanding of the effects of above- and below-ground competition on Quercus robur seedlings. Experimental seedlings were established in a typical southern Swedish shrub community where they received 1 of 4 competition levels (above-ground, below-ground, above- and below-ground, or no competition), and leaf-level responses were examined between two growth flushes. Two years after establishment, first-flush leaves from seedlings receiving above-ground competition showed a maximum rate of photosynthesis (A_max) 40% lower than those of control seedlings. With the development of a second flush above the shrub canopy, A_max of these seedlings increased to levels equivalent to those of seedlings free of light competition. Shrubby competition reduced oak seedling transpiration such that seedlings exposed to above- and below-ground competition showed rates 43% lower than seedlings that were not exposed to competition. The impaired physiological function of oak seedlings growing amid competition ultimately led to a 60-74% reduction in leaf area, 29-36% reduction in basal diameter, and a 38-78% reduction in total biomass accumulation, but root to shoot ratio was not affected. Our findings also indicate that above-ground competition reduced A_max, transpiration and biomass accumulation more so than below-ground competition. Nevertheless, oak seedlings exhibited the ability to develop subsequent growth flushes with leaves that had an A_max acclimated to utilize increased light availability. Our findings highlight the importance of flush-level acclimation under conditions of heterogeneous resource availability, and the capacity of oak seedlings to initiate a positive response to moderate competition in a shrub community.     

Soil as a mediator in plant‐plant interactions in a semi‐arid community

Abstract. Competition and facilitation may occur simultaneously in plant communities, and the prevalence of either process depends on abiotic conditions. Here we attempt a community‐wide approach in the analysis of plant interactions, exploring whether in a semi‐arid environment positive or negative interactions predominate and whether there are differences among co‐occurring shrub species. Most shrubs in our plot exerted significant effects on their understorey communities, ranging from negative to positive. We found a clear case of interference and another case where the effect was neutral, but facilitation predominated and the biomass of annuals under most shrubs in our community was larger than in gaps. Effects on soil water and fertility were revealed as the primary source of facilitation; the build‐up of soil organic matter changed soil physical properties and improved soil water relations. Facilitation by shrubs involved decoupling of soil temperature and moisture. Sheltering from direct radiation had an effect on productivity, but significant differences in understorey biomass did not parallel understorey light environment. A positive balance of the interaction among plants, essentially mediated by changes in soil properties, is the predominant outcome of plant interactions in this semi‐arid community.     

Comparative ecophysiological responses to drought of two shrub and four tree species from karst habitats of southwestern China

Drought stress is one of the most important factors in limiting the survival and growth of plants in the harsh karst habitats of southwestern China, especially at the seedling establishment stage. The ecophysiological response to drought stress of native plants with different growth forms is useful for re-vegetation programs. Two shrub and four tree species were studied, including Pyracantha fortuneana (evergreen shrub), Rosa cymosa (deciduous shrub), Cinnamomum bodinieri (evergreen tree), and other three deciduous trees, Broussonetia papyrifera, Platycarya longipes, and Pteroceltis tatarinowii. The seedlings were randomly assigned to four drought treatments, i.e., well-watered, mild drought stress, moderate drought stress, and severe drought stress. Leaf water relations, gas exchange, chlorophyll fluorescence, and growth of the seedlings were investigated. Under severe drought stress, the two shrubs with low leaf area ratio (LAR) maintained higher water status, higher photosynthetic capacity, and larger percent biomass increase than the most of the trees. The two shrubs also had lower specific leaf area, greater intrinsic water use efficiency, and thermal dissipation than the trees. This suggested that the two shrubs had high tolerance to severe drought and were suitable for re-vegetation in harsh habitats. The evergreen C. bodinieri exhibited higher leaf mass ratio (LMR) and LAR than the deciduous species under mild and moderate stress. However, the low maximum quantum efficiency of PSII photochemistry (F _v/F _m) and net assimilation rate, and the sharp decreases of water potential, LMR, LAR, and biomass under severe stress indicated C. bodinieri’s weak tolerance to severe drought. In response to drought stress, the three deciduous trees revealed sharp reductions of biomass due to the large drought-induced decreases of gas exchange, LAR, and LMR. Under drought conditions, the deciduous trees minimized water loss by stomatal closure and by reducing transpiration leaf area and light harvesting through shedding leaves. This suggested that the three deciduous trees were more sensitive to water availability than the shrubs and used avoidance strategies against drought stress. However, the better growth performance of the deciduous trees than that of the shrubs under favorable conditions suggested that deciduous trees could be suitable for habitats with mild and temporary drought stress.     

Establishing Native Grasses in a Big Sagebrush–Dominated Site: An Intermediate Restoration Step

Many semiarid rangelands in the Great Basin, U.S.A., are shifting dominance to woody species as a consequence of land degradation including intense livestock grazing and fire suppression. Whereas past rehabilitation efforts in Big sagebrush (Artemisia tridentata) steppes removed the shrub and added introduced forage grasses to successfully shift communities from shrublands to grasslands, current consensus is that native species should be included in restoration projects and that retention of some woody plants is desirable. We examined the potential for interseeding grasses into dense shrub communities as a precursor to thinning shrubs and releasing grasses from shrub interference. We compared seedling establishment of the native grass, Bluebunch wheatgrass (Pseudoroegneria spicata), with that of the Eurasia grass, Crested wheatgrass (Agropyron desertorum), in dense Ar. tridentata stands. Shrubs may play an important role as nurse plants for seedling establishment (reduced solar radiation, “island of fertility” effect) but result in highly contrasting light environments and root interference for seedlings. In experimental plots, we examined effects of Ar. tridentata shade levels (0, 40, 70, and 90% reduction of solar radiation) and initial root exclusion (present/absent) on the establishment and growth of P. spicata and Ag. desertorum seedlings. With this design we evaluated the interference effects of Ar. tridentata on the two grasses and identified the most beneficial microsites for grass restoration in Ar. tridentata–dominated communities. We predicted seedling survival and growth to be greater under moderate shade (40% reduction) and limited root competition than under no or strong shade conditions (0 and 90%) and unrestricted root interactions. Fifty to 85% of the P. spicata and Ag. desertorum seedlings survived the dry summer months of 1995 and 1996 and the intervening winter. Neither shading nor root exclusion from Ar. tridentata affected final seedling survival of either species. Seedling biomass of both grass species was negatively affected by initial root interactions with Ar. tridentata. However, the analysis of seedling biomass variability (coefficient of variation) indicated that in all shade and root‐exclusion treatments, some seedlings of both species developed to large individuals to survive in Ar. tridentata–dominated rangelands. Thus, the use of interseeding techniques shows promise for restoring herbaceous species in dense Ar. tridentata stands and should be given further consideration when shrub retention is an important consideration.     

Competitive suppression of Quercus douglasii (Fagaceae) seedling emergence and growth

Reduced recruitment of blue oak (Quercus douglasii) seedlings in California grasslands and woodlands may result from shifts in seasonal soil water availability coincident with replacement of the native perennial herbaceous community by Mediterranean annuals. We used a combination of container and field experiments to examine the interrelationships between soil water potential, herbaceous neighborhood composition, and blue oak seedling shoot emergence and growth. Neighborhoods of exotic annuals depleted soil moisture more rapidly than neighborhoods of a perennial grass or "no-neighbor" controls. Although effects of neighborhood composition on oak seedling root elongation were not statistically significant, seedling shoot emergence was significantly inhibited in the annual neighborhoods where soil water was rapidly depleted. Seedling water status directly reflected soil water potential, which also determined the extent and duration of oak seedling growth during the first year. End-of-season seedling height significantly influenced survival and growth in subsequent years. While growth and survival of blue oak seedlings may be initially constrained by competition with herbaceous species, subsequent competition with adult blue oak trees may further contribute to reduced sapling recruitment.     

Interactions between annuals and woody perennials in a Western Australian nature reserve

Abstract. We studied the interactions between woody perennial species and native and non-native annual species in a number of vegetation types within a nature reserve in the Western Australian wheatbelt. In particular, we examined the responses of annuals to perennial canopy removal, fire, soil disturbance and nutrient additions, and the effects of removal of annuals on perennial seedling regeneration. Experimental shrub removal significantly increased the abundance of annuals in a dense shrubland dominated by Allocasuarina campestris, but had no effect in a more open species-rich sandplain heath. Soil disturbance and nutrient addition in the heath area had no significant influence on annual abundance until three years after treatment. Fire had no clear effect on annual abundance in the heath within the reserve, but promoted a large increase in non-native species within an adjacent roadverge. A pattern of increased soil nutrient levels was accompanied by greatly increased non-native annual abundance beneath individual trees of Santalum spicatum. Exploratory laboratory bioassay experiments indicated that several woody perennials produced leachates that were capable of reducing the germination or growth of the introduced grass Avena fatua, indicating that allelopathy may be an important component of the interaction between the annual and perennial components. Within a woodland community, fire temporarily reduced the abundance of annual species and increased the establishment of perennial seedlings. Field experiments showed that annuals significantly reduced the survival of seedlings of the shrub Allocasuarina campestris. Our results indicate that intact native vegetation canopies effectively prevent invasion by non-native annuals, and that regeneration by native perennials is likely to be inhibited by the presence of an abundant annual cover.     

High N,dry: Experimental nitrogen deposition exacerbates native shrub loss and nonnative plant invasion during extreme drought

Hotter, longer, and more frequent global change‐type drought events may profoundly impact terrestrial ecosystems by triggering widespread vegetation mortality. However, severe drought is only one component of global change, and ecological effects of drought may be compounded by other drivers, such as anthropogenic nitrogen (N) deposition and nonnative plant invasion. Elevated N deposition, for example, may reduce drought tolerance through increased plant productivity, thereby contributing to drought‐induced mortality. High N availability also often favors invasive, nonnative plant species, and the loss of woody vegetation due to drought may create a window of opportunity for these invaders. We investigated the effects of multiple levels of simulated N deposition on a Mediterranean‐type shrubland plant community in southern California from 2011 to 2016, a period coinciding with an extreme, multiyear drought in the region. We hypothesized that N addition would increase native shrub productivity, but that this would increase susceptibility to drought and result in increased shrub loss over time. We also predicted that N addition would favor nonnatives, especially annual grasses, leading to higher biomass and cover of these species. Consistent with these hypotheses, we found that high N availability increased native shrub canopy loss and mortality, likely due to the higher productivity and leaf area and reduced water‐use efficiency we observed in shrubs subject to N addition. As native shrub cover declined, we also observed a concomitant increase in cover and biomass of nonnative annuals, particularly under high levels of experimental N deposition. Together, these results suggest that the impacts of extended drought on shrubland ecosystems may be more severe under elevated N deposition, potentially contributing to the widespread loss of native woody species and vegetation‐type conversion.     

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.     

Survival,growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient

In herbaceous dominated patches and ecosystems, tree establishment is influenced partly by the ability of woody seedlings to survive and grow in direct competition with herbaceous vegetation. We studied the importance of season long wet and dry spells on the competitive interactions between herbaceous vegetation and oak seedlings along a light and nitrogen gradient in an infertile secondary successional grassland in central North America. We conducted a field experiment in which seedlings of bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis) were exposed to two levels of light (full sun and 80% shade), three levels of nitrogen input (0, 5, 15 g m^–1 yr^–1), and three levels of water input (low, medium and high). In addition, seedlings were grown with and without the presence of surrounding herbaceous vegetation under both light and all three water levels. Seedling survival, growth, and rate of photosynthesis were significantly affected by competition with herbaceous vegetation and these effects varied along the multiple resource gradient. Overall, seedling survival of both species was significantly greater in wetter and shaded plots and when surrounding herbaceous vegetation was removed and was lower in nitrogen enriched plots. We found that soil water was significantly affected by varying inputs of water, light, and the presence or absence of herbaceous vegetation, and that seedling survival and rate of photosynthesis were highly correlated with available soil water. Our findings show that the impact of season long wet and dry spells on tree seedling success in grasslands can be affected by light and soil nitrogen availability.     

Root morphology and water transport of Pistacia lentiscus seedlings under contrasting water supply: A test of the pipe stem theory

Restoration of degraded Mediterranean areas often requires the reintroduction of key-stone woody species but the establishment of seedlings of native species is frequently poor. This is partly due to insufficient knowledge of the ecology of these species at the seedling stage. Fast rooting and efficient water supply under water limiting conditions may be crucial to withstand summer drought and ensure establishment. However, knowledge of the relationship between root morphology and the water transport capacity of Mediterranean woody species in response to drought is still scarce. We evaluated the effect of low water availability on biomass allocation, root morphology and transpiration of a common Mediterranean shrub species, Pistacia lentiscus L. Seedlings of this species were grown in pots filled with soil under glasshouse conditions for 6 months, and irrigated either weekly (W+) or monthly (W?). Low water availability strongly reduced all fractions of biomass, and decreased relative biomass allocation belowground. Average diameter of fine roots colonising the soil was higher in W+ plants, but this resulted in only marginal effects on specific root length. Water limitation did not affect the topology of secondary roots colonising the soil. Surprisingly, the ratio of leaf area to coloniser roots surface area was higher in W? seedlings. Sapwood area was strongly correlated with leaf area, secondary roots cross-sectional area, and surface area of fine roots colonising the soil when all seedlings were pooled. In agreement with the pipe stem theory, the ratio of sapwood area to leaf area was not affected by watering regime. Plant water loss when soils were taken to field capacity was significantly correlated with leaf area, sapwood area, secondary roots cross-section area and coloniser roots surface area. Water loss at high water availability was greatly reduced in W? plants, as leaf area decreased and transpiration rates on a leaf area basis were similar in W+ and W? seedlings. P. lentiscus showed limited capacity to acclimate to low water availability by modifying biomass allocation and root morphology. Thus, parallel to what has been observed aboveground, this species can benefit from periods of high water availability by showing relatively high root growth rates, but may respond poorly to water scarcity.     

Spatio‐temporal variation in the effect of herbaceous layer on woody seedling survival in a Chilean mediterranean ecosystem

Questions: What is the effect of herbaceous layer on seedling establishment of three woody pioneer species in open areas of central Chile under a semi‐arid mediterranean climate? How do inter‐annual and habitat conditions (slope aspect) modulate this effect? Under high stress conditions such as the drier year and habitat (north‐facing slope) do herbs reach low abundance and have neutral effects on woody seedlings? Under medium stress conditions for these woody species, such as the wetter year and south‐facing slope, does the herbaceous layer reach greater abundance and have positive effects on woody seedlings due to increasing soil water content? Location: A watershed on the outskirts of Santiago, Chile, subjected to clearing of woody vegetation through firewood extraction and human‐set fires. Methods: In spring 2007, we set up 20 plots (3 m × 2 m). Half of each plot had herbs removed manually and by application of herbicide. In both halves of each plot, one seedling (8 months old) of each of the three native woody species (Colliguaya odorifera, Schinus polygamus and Quillaja saponaria) was planted and survival monitored subsequently. The experiment was repeated in two consecutive growing seasons (2007–2008 and 2008–2009) that differed significantly in total precipitation (152 and 256.5 mm, respectively), and replicated in two sites that differed in aspect and abiotic conditions: a moister south‐ and a drier north‐facing slope. Results: In the first and drier year, the herbaceous layer had low cover and no significant effect on seedling survival of woody species. During the second year, herbs had greater cover and a significant positive effect on spring survival of C. odorifera in the north‐facing slope, which was lost after summer. During this wetter year on the south‐facing slope, herb cover had a positive effect on survival of S. polygamus (mainly during summer). Conclusions: The role of mostly ruderal herbs on woody seedling establishment depended on the species, rainfall of the current year and slope aspect, and may be explained by soil moisture patterns. This suggests that the effect of ruderal herbs on woody seedlings shifts from neutral under high stress conditions produced by drought to positive under moderate stress conditions. Our results contribute to understand interactions between ruderal herbs and woody species under contrasting abiotic conditions. Therefore, control of the herbaceous layer may not be needed in restoration programmes for this region. Moreover, herbs may benefit restoration of woody cover in mesic habitats.