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.     

Spatial distribution of soil nutrients and ephemeral plants underneath and outside the canopy of Porlieria chilensis shrubs (Zygophyllaceae) in arid coastal Chile

Soil nutrients and density and biomass of annual plants underneath and outside the canopy of Porlieria chilensis shrubs were measured at the end of the growing season in a protected arid coastal site in Chile. Levels of soil nitrogen, phosphorus and organic matter were significantly higher underneath than outside the canopies of shrubs. Almost 4 times as many plants occurred outside than underneath shrubs, but no significant differences in total aboveground biomass were found. Several species had higher densities and/or biomass outside rather than underneath shrubs, whereas others showed the oppsite trend. Species richness was lower underneath P. chilensis canopy. The spatial microdistribution of ephemeral species may be explained by differential water and nutrient requirements. Comparison of the patterns observed in our protected site versus surrounding unprotected areas supports the generalization that man, by removing shrubs and trees, has changed a previous heterogeneous spatial distribution of nutrients to a more homogenous one.This is a contribution of the Program of Arid Zone Studies of the Universidad de la Serena     

火烧迹地柽柳灌丛资源岛特征及植被的自然恢复

干旱半干旱地区灌丛资源岛特征及形成机制多有报道,但资源岛土壤对群落稳定性与火烧迹地植被的自然恢复作用尚不明确。以酒泉盐碱地柽柳灌丛地火烧3年后,自然恢复的柽柳(Tamarix ramosissima)及其冠下草本群落为研究对象,构建柽柳枯立株体量指数(SSI,Shrub Size Index)和恢复力综合指数,探讨了资源岛特征与植被恢复、冠下草本群落多样性的关系,量化不同大小柽柳灌丛的恢复力稳定性。结果表明:(1)在不同SSI的柽柳枯立株下土壤有机质和含水率明均显高于灌丛间地,形成了明显的资源岛特征。土壤有机质最大值出现在0—10 cm土层,中灌丛的肥力积聚效果最明显。(2)土壤主要以中性盐为主,在0—40 cm土层,灌丛区域土壤可溶性盐低于冠外,呈明显的盐谷特征,中灌丛的盐谷分布最为明显。(3)随着枯立株SSI的增大,柽柳新生枝条的数量及其高度均有所增大,冠下植物的高度、盖度、密度和地上植物量也明显高于灌丛间地,且灌丛越大恢复效果越明显。(4)灌下植物Simpson指数、Shannon⁃Wiener指数、Margalef指数和Pielou指数随着枯立株SSI的增大均先降后升,多样性指数低,群落结构简单。(5)恢复力综合指数随着枯立株SSI的增大呈先升后降的趋势,当SSI=52.17时灌丛群落的稳定性最强。柽柳较小时主要是冠下草本植物的恢复维持群落稳定性,随着灌丛的增大逐渐以柽柳灌木的恢复来维持群落稳定性。柽柳灌丛形成的高养低盐的土壤环境对植物群落稳定性的维持和植被的恢复有促进作用,对荒漠生态系统火烧迹地植被恢复和生态保护具有重要意义。     

Resource heterogeneity generated by shrubs and topography on coastal sand dunes

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

Influence of a non-native invasive tree on primary succession at Mt. Koma, Hokkaido, Japan

The slopes of Mt. Koma in Japan are undergoing primary succession as a result of a 1929 eruption. Understory vegetation below a non-native invasive tree species, Larix kaempferi, a native tree, Betula ermanii, and in the open were compared to determine if the non-native tree species was influencing species composition. Larix canopies are significantly larger than Betula canopies. Vegetation under Larix canopies had significantly greater richness and diversity than vegetation in the open, vegetation under Betula was intermediate but was significantly greater than the open in diversity. Vegetation cover was highest under Betula and significantly lower in the open. Larix canopy size was positively correlated with size and number of Salix reinii shrubs. Betula canopy size was positively correlated with size but not with number of Salix reinii shrubs. Species assemblages in the three sites are slightly different as shown by DCA. Due to the limited species pool on Mt. Koma the greatest possible extent of differences between the three microsites is not large. At this point Larix certainly appears to be accelerating succession for the non-tree species. If Larix persists on the slopes then succession would be permanently deflected towards a Larix forest. This would be a case of succession being deflected towards dominance by the introduced species. This revised version was published online in June 2006 with corrections to the Cover Date.     

锦鸡儿属灌丛化对草原化荒漠区土壤线虫群落组成和代谢足迹的影响

调查了内蒙古草原化荒漠区3种锦鸡儿[垫状锦鸡儿(Caragana tibetica)、荒漠锦鸡儿(C.roborovskyi)和狭叶锦鸡儿(C.stenophylla)]灌丛内、外土壤线虫群落多样性、组成和代谢足迹,以及相关的土壤理化性质和植物群落特征,旨在探讨锦鸡儿属灌丛对土壤线虫群落的影响,并分析这种影响是否具有灌木种间差异,同时从线虫功能团水平上探究灌丛如何通过非生物因素和生物因素影响线虫群落组成。结果表明：锦鸡儿属灌丛对土壤线虫多度、丰富度和多样性无显著影响,但却显著影响土壤线虫群落组成。原因是灌丛对土壤线虫群落具有物种选择性;灌丛可能主要是通过根系分泌物、凋落物质量等因素,而不是通过土壤理化性质和林下植物影响线虫群落组成。土壤线虫组成的变化引起线虫代谢足迹发生明显变化。灌丛内土壤线虫功能代谢足迹大于灌丛外(除狭叶锦鸡儿外),说明灌丛内土壤线虫群落对碳的利用率更高。3种锦鸡儿属灌木中,狭叶锦鸡儿的结构代谢足迹最大,表明狭叶锦鸡儿对捕食杂食类线虫代谢活性的促进作用更强。土壤线虫组成的变化通过线虫代谢足迹导致土壤食物网结构发生显著性变化。狭叶锦鸡儿灌丛土壤干扰程度低,营养富集状况好...     

Species‐specific influences of shrubs on the non‐dormant soil seed bank of native and exotic plant species in central‐northern Monte Desert

Deserts shrubs are well known to facilitate vegetation aggregation, mostly through seed trapping, and stress amelioration during and after plant establishment. Because vegetation aggregation effects are a by‐product of shrub presence, beneficiary species may not only be native, but also exotic. However, despite the high risk that exotic invasive species pose to ecosystem services, little is known of the role of desert shrubs on plant invasions. We assessed the influence of two shrub species on the non‐dormant soil seed bank (i.e. the number of seeds that readily germinate with sufficient water availability) of an invasive annual grass (Schismus barbatus) and of coexisting native species in a central‐northern Monte Desert (Argentina). Soil samples were collected beneath the canopies of two dominant shrub species (Bulnesia retama and Larrea divaricata) and in open spaces (i.e. intercanopies) in May 2001. Overall, the density of germinated seedlings of Schismus and that of the native species were negatively associated across microsite types. Schismus density was similar to that of all native species pooled together (mostly annuals), and was highest in Larrea samples (with no significant differences between Bulnesia and intercanopies). On the contrary, the density of all native species pooled together was highest in Bulnesia samples. Our results suggest that shrubs may contribute to plant invasions in our study system but, most importantly, they further illustrate that this influence can be species specific. Further research is needed to assess the relative importance of in situ seed production (and survival) and seed redistribution on soil seed bank spatial patterns.     

Changes in Spatial Patterns of Caragana stenophylla along a Climatic Drought Gradient on the Inner Mongolian Plateau

Few studies have investigated the influence of water availability on plant population spatial patterns. We studied changes in the spatial patterns of Caragana stenophylla along a climatic drought gradient within the Inner Mongolian Plateau, China. We examined spatial patterns, seed density, “nurse effects” of shrubs on seedlings, transpiration rates and water use efficiency (WUE) of C. stenophylla across semi-arid, arid, and intensively arid zones. Our results showed that patches of C. stenophylla populations shifted from a random to a clumped spatial pattern towards drier environments. Seed density and seedling survival rate of C. stenophylla decreased from the semi-arid zone to the intensively arid zone. Across the three zones, there were more C. stenophylla seeds and seedlings underneath shrub canopies than outside shrub canopies; and in the intensively arid zone, there were almost no seeds or seedlings outside shrub canopies. Transpiration rates of outer-canopy leaves and WUE of both outer-canopy and inner-canopy leaves increased from the semi-arid zone to the intensively arid zone. In the intensively arid zone, transpiration rates and WUE of inner-canopy leaves were significantly lower and higher, respectively, than those of outer-canopy leaves. We conclude that, as drought stress increased, seed density decreased, seed proportions inside shrubs increased, and “nurse effects” of shrubs on seedlings became more important. These factors, combined with water-saving characteristics associated with clumped spatial patterns, are likely driving the changes in C. stenophylla spatial patterns.     

Effects of Prosopis flexuosa on soil properties and the spatial pattern of understorey species in arid Argentina

Abstract. In arid zones dominant woody plants are capable of causing changes in microclimate and soil properties likely to affect species composition, as well as the establishment and spatial distribution of plant species. In North American and European deserts species richness appears to be higher under the canopy of shrubs and trees, in contrast with Chilean deserts where it seems to be lower. Since Prosopis flexuosa (Fabaceae, Mimosoideae) is the most conspicuous tree in the central Monte desert, Argentina, we analysed the effect of this species on the composition and abundance of the shrub and herbaceous layers and on soil properties. We considered two mesohabitats: ‘under P. flexuosa canopy’ and ‘intercanopy areas’. In addition, we analysed the differences between two microhabitats under canopies: ‘northern part of the canopy’ and ‘southern part of the canopy’. Results indicate that species composition and soil properties are affected by both mesohabitats and microhabitats. We found a higher number of shrubs under canopies, whereas that of grasses and perennial forbs increased in intercanopy areas. Concentrations of organic matter, nitrogen, potassium and phosphorus, factors limiting biological productivity in Monte desert soils, were significantly higher under than outside P. flexuosa canopies. Electrical conductivity and concentrations of Na⁺, Ca⁺⁺, Mg⁺⁺ were higher in the northern than in the southern microhabitats. No differences in species richness, evenness or diversity were found between mesohabitats or between microhabitats. We conclude that P. flexuosa modifies the spatial pattern of plant species in the shrub and herbaceous layers and the chemical conditions of the soil, generating spatial heterogeneity on different scales.     

Interspecific competition for seed production between two naturalized dandelions under different levels of fertilizer supply

Two naturalized dandelions,Taraxacum officinale andTaraxacum laevigatum, were grown in mixed stands at various plant densities and mixing ratios with various levels of fertilizer supply until seeds were produced. The effect of fertilizer supply on the competitive relationship for seed production between the species was analyzed. In pure stands, the simultaneous effects of plant density and fertilizer supply on mean plant weight were formulated for each species. By introducing the density conversion factor, the results of plant weight growth in mixed stands were formulated successfully.T. laevigatum produced more seeds thanT. officinale at the same plant weight. The relationship between mean plant weight and mean number of mature seeds produced per plant was also formulated for each species. Ratio diagrams were predicted with these formulas. The density conversion factors showed thatT. officinale was always dominant overT. laevigatum for plant weight growth within a generation. The ratio diagrams showed thatT. officinale became dominant overT. laevigatum for seed production with repeating generations at higher levels of fertilizer supply, but that the species with a larger ratio of plant density became dominant over the other species at lower levels of fertilizer supply.     

Islands of fertility induce co-occurring negative and positive plant-soil feedbacks promoting coexistence

Positive plant-soil feedback by “ecosystem engineers” is an important driver for the structuring and organization of resource-limited ecosystems. Although ample evidence demonstrates that plant-soil feedbacks can range from positive to strongly negative, their co-occurrence in plant communities have not yet been investigated. We test the hypothesis that the plant-soil feedback generated by the nitrogen-fixer shrub Medicago marina during primary succession in a sand dune community has a positive effect on the coexisting grass Lophochloa pubescens and a negative effect on the shrub species itself. We conducted field measurements and laboratory bioassays to evaluate (1) the effects of islands of fertility on the recruitment and growth of its ecosystem engineer and on the performance of a coexisting species and (2) the mechanisms involved that can explain the opposite effects of islands of fertility on coexisting species. Islands of fertility were present under Medicago crowns evidenced by higher available nitrogen, extractable phosphorus and potassium, organic matter, microbial activity, water holding capacity, soil humidity, and lower salt concentrations. The effects of these islands of fertility were clearly species-specific, with a facilitative impact on Lophochloa and a negative effect on Medicago recruitment. Lophochloa was denser and produced more biomass when rooted inside as compared to outside the crown area of the shrub. Contrarily, the number of seedlings of Medicago was lower inside, despite the higher seed abundance, and higher outside the crown area of adult shrubs as compared to predictions based on random distribution, thus showing a Janzen-Connell distribution. Laboratory experiments demonstrate the occurrence of Medicago negative plant-soil feedback, and that the auto-toxicity of the aboveground senescent plant material is a potentially important underlying mechanism explaining this negative feedback and the resulting Janzen-Connell distribution in the field.     

沿干旱梯度小叶锦鸡儿灌丛化对草本植物叶片矿质元素浓度及地上生物量累积的影响

草原灌丛化通过改变物种之间的相互作用深刻影响着群落的结构和功能。然而,当前有关灌木如何影响不同功能群草本植物对矿质元素吸收和累积的研究仍明显不足。在内蒙古草原沿干旱梯度选取了4个研究地点,对比分析了小叶锦鸡儿(Caragana microphylla)灌丛斑块内和斑块外群落中禾草和非禾草功能群植物叶中矿质元素浓度以及地上生物量生产的差异,旨在阐明干旱和灌木对群落草本植物叶元素累积的影响,揭示灌草间相互作用的元素利用特征及其随干旱梯度的变化。结果显示：1)随干旱加剧,灌丛斑块内的非禾草功能群植物地上生物量保持恒定,但叶中的K、Ca、Mg、Fe、Mn和Cu元素浓度显著增加(P<0.05)。该结果表明群落中的非禾草功能群植物通过提高叶中的矿质元素水平来抵御和适应干旱胁迫。2)随干旱加剧,灌丛对非禾草功能群植物地上生物量,叶中Ca、Fe、Cu和Zn,禾草功能群植物叶中的P的累积影响从负(RII<0)或中性(RII=0)转变为正效应(RII>0)。该结果与胁迫梯度假说相符,表明灌木对草本植物的促进效应随干旱胁迫加剧而增加。3)灌丛的“沃岛”效应是驱动灌木对草本植物元素累积正效应...     

Distance-dependent pollen limitation of seed set in some insect-pollinated dioecious plants

By applying hand pollination, we studied pollen limitation of seed set of female plants in four dioecious plant species with insect-pollination. The effect of hand pollination increased with distance to the nearest male plant. Distances at which seed set was 50% of its maximal value (after hand pollination) were: 2.3 m for Valeriana dioica, 5.3 m for Salix repens, 8.5 m for Asparagus officinale and 10.6 m for Bryonia dioica. We discuss to what extent the reduced seed set was caused by either fewer visits or by visits of a lower quality. We argue that quantifying distance-dependent seed set in dioecious plants may be a good way of studying effects of habitat fragmentation on the breakdown of mutualistic pollination systems.     

Facilitation by nurse shrubs of resprouting behavior in a post-fire shrubland in northern Patagonia,Argentina

Abstract. The nurse-plant syndrome is a widely recognized example of positive (facilitative) influences of plant species on the establishment and growth of other species. Most studies of the nurse-plant syndrome have been on species that reproduce mainly from seed rather than vegetatively. In this study, we experimentally compared the influences of two species of nurse shrubs, Schinus patagonicus and Berberis buxifolia, on the survival and growth of vegetatively reproducing herbaceous and woody plants in a post-fire shrubland in northern Patagonia, Argentina. The vegetation beneath shrubs was removed by clipping and, in a paired-sample design, one half of the canopy of each shrub was removed. We determined species richness, counted number of resprouts, and measured photon flux density and soil moisture beneath cut and uncut halves of each shrubs. Abundances of resprouts were several times greater beneath the uncut vs. the cut shrubs, as was the mean number of species. Thus, shrubs have a strong facilitating influence as measured by resprout densities and the number of species. Numbers of resprouts and of species were twice as high beneath Schinus as beneath Berberis, implying important differences in the facilitative effects of the two shrubs species. Microsites beneath Schinus were characterized by lower and more heterogeneous light levels but by greater soil moisture. Even though the reproductive mode in this post-fire shrubland is overwhelmingly vegetative, facilitation by nurse shrubs is important and differentially effective for different species of nurse shrubs.     

DENSITY-MEDIATED SEED AND STOLON PRODUCTION IN VIOLA (VIOLACEAE)

Seed and stolon production and spatial distribution were studied in two populations of Viola blanda and two populations of Viola rostrata in West Virginia. Mean individual plant biomass and the proportion of the population producing seeds and/or stolons both decreased with density. Both species possessed a characteristic “threshold” weight required before the onset of seed production; a similar minimum threshold weight was also reached before stolons were produced. It is suggested that competition at higher densities results in fewer individuals reaching the threshold weight for seed or stolon production. Thus, the density response appears as a reduction in the proportion of plants producing seeds or stolons.     

UV-B and PAR in single and mixed canopies grown under different UV-B exclusions in the field

The purpose of this study was to investigate whether differences in canopy architecture due to the investigated species (planophile versus erectophile, single versus mixed canopies) or to UV-B effects on plant morphology, lead to differences in UV-B and UV-B/PAR doses within canopies.The development of a very small (10 mm diameter) UV-B and PAR sensor on a long 5 mm wide stick allowed us to measure the penetration of UV-B and PAR in single and mixed canopies of the grass Dactylis glomerata and white clover, Trifolium repens. The plants were grown in greenhouses covered with different thicknesses (3 and 5 mm) of UV-transmittant plexi (12 and 18% UV-B exclusion).For clover, a planophile vegetation, radiation penetration was very low for both UV-B and PAR. UV-B penetration was much less than for PAR, resulting in low UV-B/PAR ratio's within the canopy. This is explained by the low UV-B transmittance of the leaves (<0.1 %) in combination with the planophile leaves.In the grass species, both UV-B and PAR penetrated much deeper into the canopy due to the erectophile structure. The difference between UV-B and PAR penetration was generally quite small except in very tall canopies.The mixed species canopies showed results comparable to the clover canopies. Due to the strongly increased grass growth in these plots, light penetration was generally much lower than in the single species cultures. The increased growth of grass in these mixed plots could be linked to the lower UV-B/PAR dose they received.In plots grown under the higher UV-B level there was a relative decrease in UV-B/PAR ratio within the canopy for both species, compared to canopies from the lower UV-B greenhouses. This could not be explained by changes in leaf angle or biomass, but might be linked to the increase in leaf transmittance of PAR.     

Stratospheric Ozone Depletion: High Arctic Tundra Plant Growth on Svalbard is not Affected by Enhanced UV-B after 7 years of UV-B Supplementation in the Field

The response of tundra plants to enhanced UV-B radiation simulating 15 and 30% ozone depletion was studied at two high arctic sites (Isdammen and Adventdalen, 78° N, Svalbard).The set-up of the UV-B supplementation systems is described, consisting of large and small UV lamp arrays, installed in 1996 and 2002. After 7 years of exposure to enhanced UV-B radiation, plant cover, density, morphological (leaf fresh and dry weight, leaf thickness, leaf area, reproductive and ecophysiological parameters leaf UV-B absorbance, leaf phenolic content, leaf water content) were not affected by enhanced UV-B radiation. DNA damage in the leaves was not increased with enhanced UV-B in Salix polaris and Cassiope tetragona. DNA damage in Salix polaris leaves was higher than in leaves of C. tetragona. The length of male gametophyte moss plants of Polytrichum hyperboreum was reduced with elevated UV-B as well as the number of Pedicularis hirsuta plants per plot, but the inflorescence length of Bistorta vivipara was not significantly affected. We discuss the possible causes of tolerance of tundra plants to UV-B (absence of response to enhanced UV-B) in terms of methodology (supplementation versus exclusion), ecophysiological adaptations to UV-B and the biogeographical history of polar plants     

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。     

Size-dependent foraging efficiency,cannibalism and zooplankton community structure

Only recently has the importance of positive interactions among plant species in structuring natural communities been supported by experimental evidence. Most studies have focused on interactions between a pair of species at a single life-history stage. In this study positive interactions between a woody nitrogen-fixing shrub (Myrica pensylvanica) and two herbaceous sand dune species (Solidago sempervirens, Ammophila breviligulata) which frequently grow beneath shrub canopies are examined throughout the life cycles of the herbaceous species. Comparisons of S. sempervirens and A. breviligulata growing beneath and outside M. pensylvanica shrubs show that plants growing in association with shrubs are larger, are more likely to flower, produce greater numbers of flowers and seeds, have higher midday xylem water potentials, have higher tissue nitrogen concentrations, and have higher photosynthetic efficiencies. Measurements of environmental conditions show that areas beneath shrubs are more shaded, have lower soil temperatures, and have higher soil nitrogen levels. The results from experimental manipulations designed to test the effects of Myrica shrubs on understory species suggest that the observed differences in plant performance are strongly influenced by canopy shading and soil nutrient enrichment associated with the shrubs. The results demonstrate that M. pensylvanica facilitates growth, reproduction, and recruitment of S. sempervirens and A. breviligulata growing beneath it. This study, one of the few to examine positive interactions at different life-history stages, supports previous predictions that positive interactions may be particularly important in plant communities characterized by physiologically stressful conditions. Received: 21 July 1999 / Accepted: 18 January 2000     

Apparent competition: an impact of exotic shrub invasion on tree regeneration

Invasion of habitats by exotic shrubs is often associated with a decrease in the abundance of native species, particularly trees. This is typically interpreted as evidence for direct resource competition between the invader and native species. However, this may also reflect indirect impacts of the exotic shrubs through harboring high densities of seed predators––known as apparent competition. Here I present data from separate seed predation experiments conducted with two shrub species exotic to North America; Rosa multiflora, an invader of abandoned agricultural land, and Lonicera maackii, an invader of disturbed or secondary forest habitats. Both experiments showed significantly greater risks of seed predation for tree seeds located under shrub canopies when compared to open microhabitats within the same site. These results indicate the potential importance of indirect impacts of exotic species invasions on native biota in addition to the direct impacts that are typically the focus of research.