Treefall gaps required for establishment,but not survival,of invasive Rubus phoenicolasius in deciduous forest,Maryland, USA

Although plant invasions are often associated with disturbance, localized disturbances can promote invasion either by: (i) creating sites where individuals establish; or (ii) enabling an invader to colonize the entire stand. The former is expected when both establishment and survival to reproductive age require disturbed conditions, whereas the latter should occur in systems when either establishment or survival are limited to disturbed sites. We investigated the role of localized disturbance, specifically treefalls, in the invasion of the Asian Rubus phoenicolasius in a deciduous forest in Maryland, USA. We investigated the density and demography of R. phoenicolasius in treefall gaps of various sizes, but identical age to non‐gap areas, using Classification and Regression Tree (CART) analyses to identify the most important predictors. To explore how the demography of established individuals responds to disturbed versus undisturbed conditions, we carried out a garden experiment with three different levels of shade (5, 12 and 22% full sun). We found vegetative and sexual reproduction, and seedling establishment, to be limited to large gaps in an old stand, but not in a stand in an earlier age of succession. However, in the garden experiment, established plants were able to survive and grow under all shade treatments. These findings indicate that R. phoenicolasius requires disturbances such as treefalls to establish in forests, but established plants will survive canopy closure, leading to stand‐wide invasion. Managers should be able to prevent invasion, however, by inspecting large gaps for new recruits every 3 years.     

Spatial and seasonal heterogeneity in understory light conditions caused by differential leaf flushing of deciduous overstory trees

In a deciduous broad-leaved forest, we investigated the seasonality and heterogeneity of understory light conditions in relation to the leaf phenology of overstory trees. Fisheye photographs were taken from spring to autumn to estimate direct and indirect light conditions above the understory. Spatial variation in daily direct photosynthetic photon flux density (PPFD) was highest in late May, when the early and the intermediate-flushing trees had finished flushing and the late-flushing trees had just started flushing. After whole canopy closure, spatial variation in direct PPFD became low. Thus, asynchronous overstory leaf flushing in spring resulted in spatial heterogeneity in understory light conditions. These results suggest that the leaf phenological patterns of overstory trees are an important factor in the formation of the understory community.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Sapling biomass allocation and growth in the understory of a deciduous hardwood forest

Above- and belowground tissues of co-occurring saplings (0.1-1 m height) of Acer saccharum Marsh. (very shade tolerant), Acer rubrum L. (shade tolerant), Fraxinus americana L. (intermediate shade tolerant), and Prunus serotina Ehrh. (shade intolerant) were harvested from a forest understory to test the hypothesis that the pattern of biomass allocation varied predictably with shade-tolerance rank. The placement and length of branches along the main axis were consistent with the formation of a monolayer of foliage for the tolerant and intermediate species. Other morphological characteristics did not vary predictably with shade-tolerance rank. The maintenance of high specific leaf area (SLA; leaf area/leaf mass) and leaf area ratio (LAR; leaf area/sapling mass) is considered important for growth under extreme shade, yet these traits were not clearly related to the shade-tolerance rank of these species. Fraxinus americana, an intermediate species, had the highest LAR and growth rate in the understory, and with the exception of P. serotina, the very shade-tolerant A. saccharum had the lowest LAR. Prunus serotina maintained a large starch-rich tap root and shoot dieback was common, yielding the largest root/shoot ratio for these species. The observed allocation patterns were not similar to the long-standing expectation for the phenotypic response of juvenile trees to shade, but were consistent with three hypothetical "growth strategies" in the understory: (1) the low SLA and LAR of A. saccharum may provide a measure of defense against herbivores and pathogens and thus promote persistence in the understory, (2) the high SLA for F. americana and high LAR for F. americana and A. rubrum may enable these species to achieve high growth rates in shade, and (3) the large carbohydrate stores of P. serotina may poise this species for opportunistic growth following disturbance. The relative importance of resistance to herbivores and pathogens vs. the maintenance of high growth rates may be important in evaluating the patterns of biomass allocation in the understory.     

Relationship between leaf characteristics, tree sizes and species distribution along a slope in a warm temperate forest

The relationships between some leaf characteristics, tree size and species distribution were investigated for evergreen tree species along a slope in a warm temperate forest in Japan. Tree species were classified into three groups based on their dominance on the slope: ridge species that were aggregated in an uppersite, valley species that were aggregated in a lowersite, and uniform species that were distributed almost uniformly. The ridge species had a more positive leaf carbon isotope ratio than the valley species, which suggests that the ridge species have larger water use efficiency than the valley species. This may give some advantage to the ridge species over the valley species in the uppersite where water availability would be limited. However, the ridge species had smaller leaf nitrogen content on a mass basis and larger leaf mass per area than the valley species, which suggests that the ridge species had a smaller mass-based leaf photosynthetic capacity than the valley species. This may be disadvantageous to ridge species in the lowersite, because smaller leaf photosynthetic capacities cause lower leaf carbon gain and thus lower growth than the valley species. These differences in leaf characteristics between the ridge and the valley species were affected by microenvironments, and were also partly affected by the difference in species specific responses to microenvironments on the slope.     

雪被斑块对高山森林两种灌木凋落叶质量损失的影响

高山/亚高山森林灌木层植物凋落物的分解对于系统物质循环等过程具有重要意义, 并可能受到冬季不同厚度雪被斑块下冻融格局的影响。该文采用凋落物分解袋法, 研究了高山森林典型灌层植物华西箭竹(Fargesia nitida)和康定柳(Salix paraplesia)凋落物在沿林窗-林下形成的冬季雪被厚度梯度(厚型雪被斑块、较厚型雪被斑块、中型雪被斑块、薄型雪被斑块、无雪被斑块)上在第一年不同关键时期(冻结初期、冻结期、融化期、生长季节初期和生长季节后期)的质量损失特征。在整个冻融季节, 华西箭竹和康定柳凋落叶的平均质量损失分别占全年的(48.78 ± 2.35)%和(46.60 ± 5.02)%。冻融季节雪被覆盖斑块下凋落叶的失重率表现出厚型雪被斑块大于薄型雪被斑块的趋势,而生长季节无雪被斑块的失重率明显较高。尽管如此, 华西箭竹凋落物第一年分解表现出随冬季雪被厚度增加而增加的趋势, 但康定柳凋落物第一年失重率以薄型雪被斑块最高, 而无雪被斑块最低。同时, 相关分析表明冻融季节凋落叶的失重率与平均温度和负积温呈极显著正相关, 生长季节凋落叶的失重率与所调查的温度因子并无显著相关关系, 但全年凋落物失重率与平均温度和正/负积温均显著相关。这些结果清晰地表明, 未来冬季变暖情境下高山森林冬季雪被格局的改变将显著影响灌层植物凋落物分解, 影响趋势随着物种的差异具有明显差异。     

Nitrogen addition increases intraspecific competition in the invasive wetland plant Alternanthera philoxeroides,but not in its native congener Alternanthera sessilis

Nitrogen is often released in pulses with different frequencies, and N supply pulses may affect growth, reproduction, and biomass allocation of plants. However, few studies have examined how N supply pulses affect intraspecific competition of clonal plants and whether such an effect depends on the N supply amount. We grew one (no competition) or 12 ramets (with intraspecific competition) of both an invasive clonal plant Alternanthera philoxeroides and its native congener Alternanthera sessilis in five different N treatments: control (no N addition), low/high amount with low/high frequencies (pulses). Nitrogen addition significantly increased the growth of both species, while intraspecific competition decreased it. Nitrogen addition significantly increased intraspecific competitive intensity of A. philoxeroides as measured by the log response ratio of growth traits, but did not affect that of A. sessilis. Despite the N supply amount, N pulses had little effect on the growth and thus intraspecific competition of the two species. Therefore, increasing N deposition may change population structure and dynamics and the invasion succession of A. philoxeroides, but changes in N pulses may not.     

亚热带不同树种凋落叶分解对氮添加的响应

为探究不同质量凋落物对氮(N)沉降的响应, 该研究采用尼龙网袋分解法, 在亚热带福建三明格氏栲(Castanopsis kawakamii)自然保护区的米槠(Castanopsis carlesii)天然林, 选取4种本区常见的具有不同初始化学性质的树种凋落叶进行模拟N沉降(N添加)分解实验(施N水平为对照0和50 kg·hm ^-2·a ^-1)。研究结果表明: 在2年的分解期内, 对照处理的各树种凋落叶的分解速率依次为观光木(Michelia odora, 0.557 a ^-1)、米槠(0.440 a ^-1)、台湾相思(Acacia confusa, 0.357 a ^-1)、杉木(Cunninghamia lanceolata, 0.354 a ^-1); N添加处理凋落叶分解速率依次为观光木(0.447 a ^-1)、米槠(0.354 a ^-1)、杉木(0.291 a ^-1)、台湾相思(0.230 a ^-1), 除杉木凋落叶外, N添加显著降低了其他3种凋落叶分解速率。N添加不仅使4种树木凋落叶分解过程中的N释放减慢, 同时还抑制凋落叶化学组成中木质素和纤维素的降解; N添加在凋落叶分解过程中总体上提高β-葡萄糖苷酶(βG)和酸性磷酸酶活性, 对纤维素水解酶的活性影响不一致, 而降低β-N-乙酰氨基葡萄糖苷酶活性和酚氧化酶活性。凋落叶分解速率与凋落叶中的碳获取酶(βG)活性以及其化学组分中的可萃取物含量极显著正相关, 与初始碳浓度、纤维素和木质素含量极显著负相关, 与初始N含量没有显著相关性。凋落物类型和N添加的交互作用虽未影响干质量损失速率, 但对木质素和纤维素的降解具有显著效应。综上所述, 化学组分比初始N含量能更好地预测凋落叶分解速率, 而N添加主要通过抑制分解木质素的氧化酶(如PHO)来降低凋落叶分解速率。     

Paddlefish (Polyodon spathula) in Europe: An aquaculture species and a potential invader

The paddlefish (Polyodon spathula) was first introduced to Europe in 1974, mainly due to its potential for rearing in natural polyculture ponds and large temperate reservoirs. The information on the history of paddlefish aquaculture efforts in Europe is scarce, as well as data on current paddlefish aquaculture status and trends. In addition, there is a lack of data on its presence and potential establishment in the wild, while its invasive potential and associated risks and impacts are largely unknown. In order to evaluate its current status in Europe, we conducted a survey among scientists, aquaculture producers and other stakeholders, and reviewed literature and data on the Internet. Based on the results obtained, we discuss the potential and the challenges in European paddlefish aquaculture development, and analyze paddlefish invasive potential and risks associated with its naturalization. Paddlefish aquaculture is well established only regionally in Europe, but offers relatively high potential for further development in pond farms. Nevertheless, future development will require careful planning, especially regarding market development and improved marketing strategies. While paddlefish likely represents a low‐risk invader, improved control and reporting on trade and intentional and unintentional releases will be required. Given the lack of knowledge on potential impacts following its introduction, due caution seems highly advisable.     

The influence of chlorophyll fluorescence on the light gradients and the phytochrome state in a green model leaf under natural conditions

Abstract. The effect of chlorophyll fluoresence on the spectral light gradients within a model green leaf was examined under different light qualities (day-light, sunset, canopy) and different quantum efficiencies. Light fluxes within the leaf tissue are nearly doubled in the emission domain of fluorescence but the effect on the phytochrome photoequilibrium is very small.     

Effect of native and exotic leaf litter on macroinvertebrate communities and decomposition in a western Montana stream

Invasion by exotic trees into riparian areas has the potential to impact terrestrial and aquatic systems. To test the effect of different streamside tree species on the aquatic food web in a stream in Montana, we compared decomposition rates of leaf litter and invertebrate assemblages associated with the leaf litter of the exotic Acer platanoides and the dominant native Populus trichocarpa trees. Macroinvertebrate family richness, evenness, and diversity increased with days of aquatic processing; however, there was no effect of leaf species. Leaves of the A. platanoides were associated with 70% greater density of macroinvertebrates of the family Nemouridae. This family consists primarily of detritivores and had the greatest overall density and frequency of occurrence relative to other macroinvertebrate families. The density of a family of predatory macroinvertebrates (Rhyacophilidae) was also generally (73%) greater in association with A. platanoides than P. trichocarpa leaves. The density of Ephemerellidae and Rhyacophilidae increased over time. In contrast to studies comparing leaves of exotic vs. native trees, we observed no difference in leaf decomposition rates; however, the amount of leaf inputs are likely to differ between native and invaded forests. The results indicate that replacement of native riparian trees with exotics affected the most common family of macroinvertebrates and possibly a common family of predatory macroinvertebrates (Rhyacophilidae), which may affect the detrital food web.     

Controls on the release of dissolved organic carbon and nitrogen from a deciduous forest floor investigated by manipulations of aboveground litter inputs and water flux

Although dissolved organic matter (DOM) released from the forest floor plays a crucial role in transporting carbon and major nutrients through the soil profile, its formation and responses to changing litter inputs are only partially understood. To gain insights into the controlling mechanisms of DOM release from the forest floor, we investigated responses of the concentrations and fluxes of dissolved organic carbon (DOC) and nitrogen (DON) in forest floor leachates to manipulations of throughfall (TF) flow and aboveground litter inputs (litter removal, litter addition, and glucose addition) at a hardwood stand in Bavaria, Germany. Over the two-year study period, litter manipulations resulted in significant changes in C and N stocks of the uppermost organic horizon (Oi). DOC and DON losses via forest floor leaching represented 8 and 11% of annual litterfall C and N inputs at the control, respectively. The exclusion of aboveground litter inputs caused a slight decrease in DOC release from the Oi horizon but no change in the overall leaching losses of DOC and DON in forest floor leachates. In contrast, the addition of litter or glucose increased the release of DOC and DON either from the Oi or from the lower horizons (Oe + Oa). Net releases of DOC from the Oe + Oa horizons over the entire manipulation period were not related to changes in microbial activity (measured as rates of basal and substrate-induced respiration) but to the original forest floor depths prior to manipulation, pointing to the flux control by the size of source pools rather than a straightforward relationship between microbial activity and DOM production. In response to doubled TF fluxes, net increases in DOM fluxes occurred in the lower forest floor, indicating the presence of substantial pools of potentially soluble organic matter in the Oe + Oa horizons. In contrast to the general assumption of DOM as a leaching product from recent litter, our results suggest that DOM in forest floor leachates is derived from both newly added litter and older organic horizons through complex interactions between microbial production and consumption and hydrologic transport.     

Variability in leaf optical properties among 26 species from a broad range of habitats

Leaves from 26 species with growth forms from annual herbs to trees were collected from open, intermediate, and shaded understory habitats in Mississippi and Kansas, USA. Leaf optical properties including reflectance, transmittance, and absorptance in visible and near infrared (NIR) wavelengths were measured along with leaf thickness and specific leaf mass (SLM). These leaf properties and internal light scattering have been reported to vary with light availability in studies that have focused on a limited number of species. Our objective was to determine whether these patterns in leaf optics and light availability were consistent when a greater number of species were evaluated. Leaf thickness and SLM varied by tenfold among species sampled, but within-habitat variance was high. Although there was a strong trend toward thicker leaves in open habitats, only SLM was significantly greater in open vs. understory habitats. In contrast, leaf optical properties were strikingly similar among habitats. Reflectance and reflectance/transmittance in the NIR were used to estimate internal light scattering and there were strong relationships (r1 > 0.65) between these optical properties and leaf thickness. We concluded that leaf thickness, which did not vary consistently among habitats, was the best predictor of NIR reflectance and internal light scattering. However, because carbon allocation to leaves was lower in understory species (low SLM) yet gross optical properties were similar among all habitats, the energy investment by shade leaves required to achieve optical equivalence with sun leaves was lower. Differences in leaf longevity and growth form within a habitat may help explain the lack of consistent patterns in leaf optics as the number of species sampled increases.     

Elevated CO2 reduces leaf damage by insect herbivores in a forest community

By altering foliage quality, exposure to elevated levels of atmospheric CO(2) potentially affects the amount of herbivore damage experienced by plants. Here, we quantified foliar carbon (C) and nitrogen (N) content, C : N ratio, phenolic levels, specific leaf area (SLA) and the amount of leaf tissue damaged by chewing insects for 12 hardwood tree species grown in plots exposed to elevated CO(2) (ambient plus 200 microl l(-1)) using free-air CO(2) enrichment (FACE) over 3 yr. The effects of elevated CO(2) varied considerably by year and across species. Elevated CO(2) decreased herbivore damage across 12 species in 1 yr but had no detectable effect in others. Decreased damage may have been related to lower average foliar N concentration and SLA and increased C : N ratio and phenolic content for some species under elevated compared with ambient CO(2). It remains unclear how these changes in leaf properties affect herbivory. Damage to the leaves of hardwood trees by herbivorous insects may be reduced in the future as the concentration of CO(2) continues to increase, perhaps altering the trophic structure of forest ecosystems.     

Effects of Flow Restoration and Exotic Species Removal on Recovery of Native Fish: Lessons from a Dam Decommissioning

Flow diversion and invasive species are two major threats to freshwater ecosystems, threats that restoration efforts attempt to redress. Yet, few restoration projects monitor whether removal of these threats improve target characteristics of the ecosystem. Fewer still have an appropriate experimental design from which causal inferences can be drawn as to the relative merits of removing exotic fish, restoring flow, or both. We used a dam decommissioning in Fossil Creek, Arizona, to compare responses of native fish to exotic fish removal and flow restoration, using a before‐after‐control‐impact design with three impact treatments: flow restoration alone where exotics had not been present, flow restoration and exotic fish removal, and flow restoration where exotics remain and a control reach that was unaffected by restoration actions. We show that removal of exotic fish dramatically increased native fish abundance. Flow restoration also increased native fish abundance, but the effect was smaller than that from removing exotics. Flow restoration had no effect where exotic fish remained, although it may have had other benefits to the ecosystem. The cost to restore flow ($12 million) was considerably higher than that to eradicate exotics ($1.1 million). The long‐term influence of flow restoration could increase, as travertine dams grow and re‐shape the creek increasing habitat for native fish. But in the 2‐year period considered here, the return on investment for extirpating exotics far exceeded that from flow restoration. Projects aimed to restore native fish by restoring flow should also consider the additional investment required to eradicate exotic fish.     

不同光环境下亚热带常绿阔叶树种和落叶阔叶树种幼苗的叶形态和光合生理特征

由于地球环境的演变,亚热带常绿阔叶林中常有落叶阔叶成分的存在,但从生态角度,其存在的机理尚不清楚.通过比较研究常绿阔叶优势树种大头茶与落叶阔叶树种枫香幼苗的叶形态和光合生理特征对不同光环境(旷地、林窗、林下)的响应,尝试解释落叶成分遗留的基础.结果表明:①枫香具有较小的比叶重(LMA)和较高的光合氮利用效率(PNUE),在高光环境中,并未受到光抑制,具有较高的光合可塑性,尤其在林窗,表现出较大头茶高3倍的光合能力(Pnmax),保证了其在较短的生长季节积累更多的光合产物,增强与常绿树种的竞争力;②常绿阔叶树种大头茶具有较大的LMA,在旷地受到严重的光抑制,将更多的氮用于化学防御中.在林窗和林下具有相对高的Pnmax,生长幅度较广,但在林下将更多的氮投入到比生长更为重要的生存消耗中.而枫香在林下响应表现为Rubisco活性降低,光合受阻;③两树种在林窗均表现出较高的光合适应性,具有最大的光合能力(Pnmax)及合理的氮在光合机构中的分配系数,说明林窗是它们更新的最佳环境.但枫香的光合作用更依赖于光照,而大头茶则更依赖于CO2浓度;④总之,落叶阔叶树种以其较高的形态可塑性和对高光的光合生理可塑性能够在常绿阔叶林种生存、生长,并成为常绿阔叶林中固有成分的主要原因之一.     

Climate controls the distribution of a widespread invasive species: implications for future range expansion

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Variable host phenology does not pose a barrier to invasive weevils in a northern hardwood forest

• 1 A suite of invasive weevils has established in hardwood forests of the North American Great Lakes Region. We quantified patterns of host availability and the capacity of adults to succeed in a system with high host variability both within and between seasons in Michigan, U.S.A.
• 2 We quantified phenological development of foliage on three host species [sugar maple, Acer saccharum Marshall; ironwood, Ostrya virginiana (Mill.) K. Koch; and raspberry, Rubus spp.]. We estimated adult abundance using emergence traps and sweep net sampling over 3 years, and compared field host associations with laboratory choice assays.
• 3 Host plant phenology varied among species, between years, and in their interactions. The four most common weevils, Phyllobius oblongus (L.), Polydrusus sericeus (Schaller), Barypeithes pellucidus (Boheman) and Sciaphilus asperatus (Bonsdorff), emerged in early to mid‐June, in approximately that order. After emergence, each species showed evidence of host preference based on their abundances on foliage. Overall, P. oblongus and B. pellucidus were most prevalent on sugar maple, P. sericeus was most prevalent on ironwood, and S. asperatus was relatively evenly distributed. Laboratory choice tests with P. oblongus and P. sericeus confirmed these preferences.
• 4 These four invasive species comprised over 99% of all 12 845 weevils obtained, suggesting displacement of native species. The optimal sampling methods varied among weevil species.
• 5 These invasive weevils contend with the highly variable conditions of their environment, and also potential phenological asynchrony, via relatively late emergence, even at the cost of lower host quality. Annual variation is greater for numbers of adults than larvae, suggesting that mortality of late instars or pupae is particularly important.