Effects of macrophyte heterogeneity and food availability on structural parameters of the macroinvertebrate community in a Pampean stream

Environmental heterogeneity in natural ecosystems influences several parameters at the population and community levels. In freshwater ecosystems, habitat heterogeneity can be provided by macrophyte species with different structural shapes. Previous studies suggest that aquatic plants with more complex architectures will support higher number, biomass, and taxon richness of macroinvertebrates than plants with simpler shape. We investigated the influence of macrophyte structural heterogeneity (quantified by fractal dimension) and food availability (represented by epiphytic biomass) on several parameters (number of individuals, biomass, body size distribution, taxon richness, and diversity) of the macroinvertebrate community in a Pampean stream. Four submerged macrophyte species (Egeria densa, Elodea ernstae, Ceratophyllum demersum, and Stuckenia striata) and associated macroinvertebrates were sampled in late spring, summer, and autumn. Plants were photographed and fractal dimension was estimated from the images by the box-counting method. Fractal dimension was independent of plant surface area per unit of macrophyte biomass and differed significantly among species. Mean fractal dimension varied between 1.29 and 1.62, and increased following the sequence E. densa → S. striata → E. ernstae → C. demersum. Macrophyte species with higher fractal dimension supported a greater abundance of macroinvertebrates, especially those of small body size (500–1,000 μm); but fractal dimension was unrelated to macroinvertebrate biomass, richness, and diversity. However, overall animal biomass was significantly associated to the epiphytic abundance. Consequently, macrophyte heterogeneity influences macroinvertebrate density and body size distribution, while animal biomass depends on epiphytic food resources provided by plants.     

Aquatic macroinvertebrate richness and diversity associated with native submerged aquatic vegetation plantings increases in longer-managed and wetland-channeled effluent constructed urban wetlands

Macroinvertebrate community structure and assemblages associated with the planted, native submerged aquatic vegetation (SAV) species Heteranthera dubia (Jacq.) MacMillan and Potamogeton nodosus Poiret were examined in a series of constructed urban floodway wetlands, the Dallas Floodway Extension Lower Chain of Wetlands, Dallas, TX, USA. Macroinvertebrate community metrics, including abundance, richness, diversity, and evenness associated with SAV and three different wetlands of varying construction completion dates, water sources (direct or wetland-channeled wastewater effluent), and ecosystem management stage (established/reference or developing) were compared and analyzed. Assemblages at sampling sites were also classified and related to vegetation and wetland physicochemical parameters. Plant species affected only macroinvertebrate abundance, with the less-dissected P. nodosus supporting higher counts than H. dubia. Wetland age and water-effluent type had the most substantial effect on macroinvertebrate communities. The older, longer-managed wetland and wetland-channeled effluent habitat consistently demonstrated higher quality metrics and biodiversity than newly constructed, direct effluent wetland habitat. Increased vegetation cover and wetland age, coupled with moderate water temperature, pH, and DO levels were characteristics of more rich and diverse macroinvertebrate communities, including pollutant-sensitive taxa, such as Ephemeroptera and Trichoptera.

     

Effects of invasive species on plant communities: an example using submersed aquatic plants at the regional scale

Submersed aquatic plants have a key role in maintaining functioning aquatic ecosystems through their effects on the hydrological regime, sedimentation, nutrient cycling and habitat of associated fauna. Modifications of aquatic plant communities, for example through the introduction of invasive species, can alter these functions. In the Sacramento-San Joaquin River Delta, California, a major invasive submersed plant, Brazilian waterweed Egeria densa, has become widespread and greatly affected the functionality of the submersed aquatic plant community. Rapid assessments of the distribution and abundance of this species are therefore crucial to direct management actions early in the season. Given the E. densa bimodal growth pattern (late spring and fall growth peaks), summer assessments of this species may indicate which and where other submersed species may occur and fall assessments may indicate where this and other species may occur in the following spring, primarily because the Delta’s winter water temperatures are usually insufficient to kill submersed aquatic plant species. We assessed community composition and distribution in the fall of 2007 and summer of 2008 using geostatistical analysis; and measured summer biomass, temperature, pH, salinity, and turbidity. In the fall of 2007, submersed aquatic plants covered a much higher proportion of the waterways (60.7%) than in the summer of 2008 (37.4%), with a significant overlap between the seasonal distribution of native and non-native species. Most patches were monospecific, and multispecies patches had significantly higher dominance by E. densa, co-occurring especially with Ceratophyllum demersum. As species richness of non-natives increased there was a significant decrease in richness of natives, and of native biomass. Sustained E. densa summer biomass negatively affected the likelihood of presence of Myriophyllum spicatum, Potamogeton crispus, and Elodea canadensis but not their biomass within patches. Depth, temperature and salinity were associated with biomass; however, the direction of the effect was species specific. Our results suggest that despite native and invasive non-native submersed plant species sharing available niches in the Delta, E. densa affects aquatic plant community structure and composition by facilitating persistence of some species and reducing the likelihood of establishment of other species. Successful management of this species may therefore facilitate shifts in existing non-native or native plant species.     

Macroinvertebrate communities in Phragmites australis (Cav.) Trin. ex Steud. reed beds and open bank habitats in central victorian streams in Australia

Reed invasion is a common phenomenon of open streams with disturbed riparian vegetation in river catchments. Knowledge of the effects of such vegetation change on aquatic communities is fundamental to river management. Macroinvertebrate fauna in Phragmites australis (Cav.) Trin. ex Steud. and open bank habitats were examined in three rivers in central Victoria in order to understand the effect of such littoral habitat on macroinvertebrates. Data were analysed using Partially Nested Factorial ANOVA with season, river and habitats as main effects. Habitat structure had a significant effect (p<0.05) on macroinvertebrate species richness, however this was not seasonally consistent across the three rivers. There was a significant increase (p<0.05) in macroinvertebrate taxa richness in Phragmites habitats during winter and spring seasons. Total abundance of taxa showed no consistent significant differences in the two habitats. Results of Canonical Analysis of Principle Coordinates indicated significant differences (p<0.05) in macroinvertebrate assemblages between Phragmites and bare bank habitats in all seasons. Habitat selection by taxa could be related to the microphysical environment of the habitats. This study suggests that reed beds create important littoral habitat structures which support diverse macroinvertebrate assemblages.     

垫状植物囊种草对群落物种多样性的影响

在甘肃盐池湾国家级自然保护区内海拔4137 m处,选择典型的囊种草垫状植被设置研究样地,研究了垫状植物囊种草对群落物种组成和群落物种多样性的影响,并且定量的研究了囊种草对群落物种丰富度的影响能力和维持潜力。研究结果表明:囊种草为群落中增加了新的植物种类,并且提高了部分生境一般种的多度;囊种草的出现提高了群落物种密度和物种丰富度,进而提高了群落物种多样性;囊种草斑块的增加将会引起景观水平物种丰富度的增加,表明囊种草具有为群落中引入新的植物种类进而提高群落物种丰富度的能力;在景观水平,囊种草所创造的生境多样性则成为一种保障,可以维持景观中物种丰富度从而降低物种损失的风险,表明囊种草具有较高的群落物种丰富度维持潜力。     

Effects of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain

1. We examined the effect of the exotic macrophyte, para grass (Urochloa mutica), on benthic and epiphytic macroinvertebrates of a tropical floodplain in northern Australia. Macroinvertebrates were sampled from four grass communities: (1) para grass, (2) hymenachne (Hymenachne acutigluma), a native perennial; (3) rice (Oryza meridionalis), a native annual, and (4) areas where para grass had been sprayed with herbicide. 2. Macroinvertebrate richness, abundance and community similarity showed very few differences among the grass communities, particularly in the epiphytic habitat. Benthic invertebrates showed some differences among grasses, with lower richness and abundance and different community structure associated with hymenachne. Herbicide control of para grass had no apparent effect on benthic invertebrates but reduced the abundance of epiphytic invertebrates in the short term. 3. The results of this study indicate that para grass has very little impact on macroinvertebrate communities, despite the changes to macrophyte communities. This is probably because para grass has similar physical structure to the native grasses and because none of these grasses contribute directly to aquatic food webs. Control of para grass using herbicide has little impact on aquatic invertebrates. This suggests that predicting the impact of weed invasion in wetlands requires an understanding of both the functional properties of macrophytes and the habitat preferences of the macroinvertebrates.     

新疆开都河流域水生植物多样性及其影响因素

河流是一个连续的、流动的、独特而完整的系统,研究河流生态系统中水生植物的多样性分布格局及其影响因素对河流生态学研究具有重要意义。本文通过野外调查,研究了新疆开都河流域水生植物多样性、主要水生植物群落特征及与环境因子之间的关系,并利用水分-能量动态假说和栖息地异质性假说对该流域水生植物物种多样性的地理格局进行解释。结果表明: 开都河流域共有水生植物71种,隶属于24科39属;聚类分析可将开都河流域水生植物群落划分为10个主要群落类型,其中芦苇群落物种丰富度最高,狭叶香蒲群落和金鱼藻群落物种丰富度最低;流域水生植物群落Shannon指数与pH呈显著负相关,Simpson指数与pH、经度呈显著负相关,与海拔呈显著正相关;流域水生植物群落类型主要受海拔、水深及水温的影响;流域水生植物物种多样性随经纬度无明显变化规律。水分-能量动态假说和栖息地异质性假说共解释开都河流域水生植物多样性格局变量的31.4%,表明这两个假说对于该流域水生植物多样性格局的解释力并不高。     

河西走廊水生植物多样性格局、群落特征及影响因素

水生植物是湿地生态系统重要组成部分,研究水生植物多样性分布格局及其影响因素对地区水生植物资源保护具有重要意义。通过野外调查并结合气候等环境因素,研究了河西走廊主要水生植物群落类型、数量特征、水生植物多样性分布格局及影响因素,并对中域效应假说进行了验证。研究结果表明:(1)河西走廊地区共有水生植物29科42属84种,群落的聚类分析可将河西走廊水生植物群落划分为15个主要群落类型;(2)河西走廊水生植物群落类型主要受到水温、海拔、经纬度等环境因子影响,群落物种多样性指数与盐度以及溶解性固体总量呈显著性相关;(3)河西走廊水生植物多样性空间格局呈现出"∩"型的单峰格局,中域效应模型能较好地解释该地区水生植物多样性水平的纬度格局及海拔垂直分布格局,对该区域水生植物物种丰富度在纬度和海拔梯度上的变异解释率分别为57.56%、63.5%。分析表明,河西走廊水生植物物种丰富度格局由几何(边界)限制和随机过程及其他未知因素共同控制,且几何(边界)限制和随机过程贡献率较大;同时本研究中未考虑的环境异质性、气候、人为干扰等因素也对河西走廊水生植物多样性空间分布产生重要影响。     

Intensive removal of signal crayfish (Pacifastacus leniusculus) from rivers increases numbers and taxon richness of macroinvertebrate species

Invasive species are a major cause of species extinction in freshwater ecosystems, and crayfish species are particularly pervasive. The invasive American signal crayfish Pacifastacus leniusculus has impacts over a range of trophic levels, but particularly on benthic aquatic macroinvertebrates. Our study examined the effect on the macroinvertebrate community of removal trapping of signal crayfish from UK rivers. Crayfish were intensively trapped and removed from two tributaries of the River Thames to test the hypothesis that lowering signal crayfish densities would result in increases in macroinvertebrate numbers and taxon richness. We removed 6181 crayfish over four sessions, resulting in crayfish densities that decreased toward the center of the removal sections. Conversely in control sections (where crayfish were trapped and returned), crayfish density increased toward the center of the section. Macroinvertebrate numbers and taxon richness were inversely correlated with crayfish densities. Multivariate analysis of the abundance of each taxon yielded similar results and indicated that crayfish removals had positive impacts on macroinvertebrate numbers and taxon richness but did not alter the composition of the wider macroinvertebrate community. Synthesis and applications: Our results demonstrate that non‐eradication‐oriented crayfish removal programmes may lead to increases in the total number of macroinvertebrates living in the benthos. This represents the first evidence that removing signal crayfish from riparian systems, at intensities feasible during control attempts or commercial crayfishing, may be beneficial for a range of sympatric aquatic macroinvertebrates.     

河流微生境异质性与大型底栖动物空间分布的关系

大型底栖动物是河流生态系统较好的指示类群,越来越广泛地应用于河流生态状况评价.识别大型底栖动物空间分布的影响因素可以分为宏观尺度因子、中观尺度因子和微观尺度因子.前人的研究较多集中于中观尺度.本研究基于10 m河段微生境因子与大型底栖动物调查,采用地统计学方法,在微观尺度上分析了大型底栖动物空间分布特征及其与微生境的关系.结果表明:不同生境类型中底栖动物指标存在差异性,激流、深潭和急流的Shannon指数平均值分别为2、1.9、1.78;大型底栖动物密度、生物量、丰富度指数及Shannon指数具有空间自相关性,存在一定的扩散效应;微生境因子与大型底栖动物指标及物种的空间关系存在差异性,其中,水深和流速与大型底栖动物关系的拟合度较弱,底质组成与大型底栖动物的拟合关系相对较好,底栖动物密度与粒径<4 mm底质体积百分比呈现正空间自相关,生物量、丰富度指数、Shannon指数与粒径>32 mm底质体积百分比呈现正空间自相关.本研究结果可为底栖动物扩散机制研究、调查样方设置、物理生境修复等提供参考.     

Macroinvertebrate assemblages in Andean Patagonian rivers and streams: environmental relationships

Macroinvertebrate communities from 29 streams and rivers of the mountain and the Andean Patagonian Plateau were analyzed. Samples were collected from six river basins, which were part of four different biozones of the Patagonian Ecoregion. Samples from mountain streams were dominated by Plecoptera, Ephemeroptera, Trichoptera and Diptera, while plateau rivers where mainly Diptera, Oligochaeta and Mollusca. Total invertebrate abundance ranged from 7 to 12249 ind.m^–2. Elmidae, Paratrichocladius, Chironomus, Smicridea annulicornis, Parasericostoma ovale and Meridialaris laminata were the most abundant insect taxa, while Nais communis and Hyalella curvispina were the most abundant non-insect taxa. Species-environmental relationships were examined using Canonical Correspondence Analysis. Current speed, conductivity, substrate size and abundance of aquatic plants, were identified as the major variables structuring faunal assemblages. Regression analyses revealed that species richness was negatively correlated with latitude, and positively correlated with water temperature and altitude. Macroivertebrate abundance increased with conductivity, altitude and water temperature. These results suggest that habitat heterogeneity was the strongest predictor of macroinvertebrate assemblages, but species richness could be predicted at a landscape scale using topographical and climatic features.     

Distribution and growth of benthic macroinvertebrates among different patch types of the littoral zones of two arctic lakes

1. To evaluate the effect of habitat patch heterogeneity on abundance and growth of macroinvertebrates in arctic lakes, macroinvertebrate abundance, individual biomass, and potential food resources were studied in three patch types in two arctic lakes on the Alaskan North Slope near the Toolik Lake Field Station. An experiment was conducted to determine which sediment patch type supported higher growth rates for Chironomus sp., a commonly occurring macroinvertebrate. 2. Potential organic matter (OM) resources were significantly higher in both rock and macrophyte patches than in open‐mud patches. Total macroinvertebrate densities in both lakes were highest in rock patches, intermediate in macrophytes and lowest in open‐mud. The open‐mud patches also had lower species richness compared with other patch types. Additionally, individual biomass for one clam species and two chironomid species was significantly greater in rock patches than in open‐mud. 3. In a laboratory experiment, Chironomus showed two to three times greater mass increase in sediments from macrophyte and rock patches than from open‐mud patches. Rock and macrophyte experimental sediments had at least 1.5 × the percentage OM as open‐mud sediments. 4. Chlorophyll a appeared to be the best predictor for invertebrate abundances across all patch types measured, whereas OM content appeared to be the variable most closely associated with Chironomus growth. 5. Our results combined with previous studies show that the relationships between macroinvertebrate community structure, individual growth, and habitat heterogeneity are complex, reflecting the interaction of multiple resources, and biotic interactions, such as the presence or absence of a selective vertebrate predator (lake trout, Salvelinus namaycush).     

Distribution differences and active habitat choices of invertebrates between macrophytes of different morphological complexity

This study explores: (1) whether the abundance of macroinvertebrates differs between macrophytes differing in both morphological complexity and tolerance to nutrient enrichment; (2) whether the distribution of invertebrates between macrophytes is due to active habitat choice; and (3) whether invertebrates prefer structurally complex to simple macrophytes. Macroinvertebrate abundance was compared between two common soft-bottom plants of the Baltic Sea that are tolerant to eutrophication, Myriophyllum spicatum and Potamogeton pectinatus, and one common plant that is sensitive to eutrophication, Chara baltica. Both field sampling and habitat choice experiments were conducted. We recorded higher total macroinvertebrate abundance on the structurally complex M. spicatum than on the more simply structured P. pectinatus and C. baltica, but found no difference in macroinvertebrate abundance between P. pectinatus and C. baltica. In accordance with the field results, our experiment indicated that the crustacean Gammarus oceanicus actively chose M. spicatum over the other macrophytes. Besides, we found that G. oceanicus actively preferred complex to simply structured artificial plants, indicating that the animal distribution was at least partly driven by differences in morphological complexity between plant species. In contrast, the gastropod Theodoxus fluviatilis did not make an active habitat choice between the plants. Our findings suggest that human-induced changes in vegetation composition can affect the faunal community. Increased abundance of structurally complex macrophytes, for example, M. spicatum, can result in increased abundance of macroinvertebrates, particularly mobile arthropods that may actively choose a more structurally complex macrophyte.     

The effects of salinity on aquatic plant germination and zooplankton hatching from two wetland sediments

1. The effect of increasing salinity on the emergence of zooplankton eggs and the germination of aquatic plant seeds from the sediment of two wetlands was examined. Salinity was found to cause reductions in species richness and abundance of aquatic plants and zooplankton at salinities between 1000 and 5000 mg L^?1. Aquatic plants also had an associated decrease in above ground biomass. 2. Individual taxa showed different responses to salinity, and four response patterns were identified: (i) increased number of organisms emerging at 1000 mg L^?1; (ii) decreased number of organisms emerging above 1000 mg L^?1; (iii) decreased number of organisms emerging between 300 and 1000 mg L^?1; (iv) no difference in number of organisms emerging across the range of salinities. Response patterns (iii) and (iv) were common to both plants and zooplankton, whereas response patterns (i) and (ii) were only identified for zooplankton. 3. Results indicate that there is potential for the increasing salinity in Australian rivers and wetlands to decrease the species richness of aquatic communities resulting in loss of wetland biodiversity.     

Macroinvertebrate community dynamics in a temperate European Atlantic river. Do they conform to general ecological theory?

Spatial and temporal dynamics of macroinvertebrate communities have usually been linked to several environmental and anthropic factors. The aim of this study is to elucidate how important are these factors in structuring macroinvertebrate communities from temperate regions. Regarding the macroinvertebrate number of taxa, the Habitat Template Model, the Dynamic Equilibrium Hypothesis and the Intermediate Disturbance Hypothesis will be tested in order to know how important the diversity of instream elements and the hydrological disturbance frequency are in defining the macroinvertebrate taxonomic richness. Thus, the structure and composition of macroinvertebrate communities were analysed in nine sites of the Pas River basin, a temperate Atlantic basin in northern Spain, during winter, spring, summer and autumn 2005, together with water physicochemical and environmental characteristics. Macroinvertebrate abundance increased downstream and during summer, probably favoured by lower hydraulic stress and water organic enrichment. As predicts the Habitat Template Model, the macroinvertebrate number of taxa was related to habitat heterogeneity. However, no clear relationship amongst macroinvertebrate richness and water quality was found. The macroinvertebrate taxonomic richness did not correspond exactly with the Dynamic Equilibrium Hypothesis and the Intermediate Disturbance Hypothesis because it was relatively high in the absence of hydrological disturbances (summer). Thus, disturbance events may play a secondary role in determining the seasonal dynamic of the number of taxa. However, hydrological disturbances can be considered the most important factors explaining the seasonal pattern of macroinvertebrate abundance. On the other hand, spatial patterns of macroinvertebrate community structure and composition were mainly determined by resource availability, hydraulic conditions, habitat heterogeneity and human alterations, whilst hydrological predictability and resource availability might play a major role in determining seasonal dynamics.     

A comparison between aquatic birds of lakes and coastal rivers in Florida

Aquatic birds were counted on five Gulf coast Florida rivers to determine if these river systems supported densities, biomass and species richness similar to those found on Florida lakes. Forty-two species were identified and for the species that were found on both Florida streams and lakes similar densities and biomass were encountered. As with Florida lakes, stream bird abundance and species richness were higher in winter months than in summer months, a consequence of migratory bird populations. Total bird abundance, biomass per unit of phosphorus, and species richness per unit of area were similar to data collected on Florida lakes. Thus, Florida rivers are capable of supplying sufficient resources to maintain bird densities, biomass and species richness values similar to lakes of equal size and nutrient concentrations and are therefore important habitats for aquatic bird populations. An examination of individual habitat characteristics indicates that water depth was inversely correlated and submersed aquatic vegetation was positively correlated with bird density, biomass and species richness within the river systems. While both habitat characteristics are important they are also inversely related making it difficult to separate the individual significance of each characteristic.     

豚草入侵对新疆伊犁河谷林下本地草本植物群落结构的影响

为探究外来入侵植物豚草（Ambrosia artemisiifolia）对本地植物群落结构的影响,结合区内林下草本植物调查和室内分析方法,分析了豚草不同入侵压力下（无入侵、轻度入侵、中度入侵、重度入侵）本地草本植物丰富度、盖度等特征,探讨了地形、气候、土壤、光照、人口密度等外部因素与豚草种群特征、本地植物群落结构之间的关系。结果表明：豚草种群盖度与其高度、密度、生物量显著正相关。与对照相比,轻度入侵下本土草本植物Shannon-Wiener多样性指数和Pielou均匀度指数显著增加,丰富度、盖度未出现显著差异。随着豚草入侵程度加剧,本地草本植物物种丰富度、盖度极显著降低,样方Shannon-Wiener多样性指数先升高再降低,本土草本植物Pielou均匀度指数差异不显著。冗余分析（RDA）表明,土壤全氮、与道路距离、土壤全磷、郁闭度对研究区植物群落结构影响最大,土壤全氮含量随豚草盖度和生物量增加而减少,距道路越近或光照越强,豚草盖度和生物量越高。增强伊犁河谷地区交通往来货物豚草检验检疫力度、增加林内植被郁闭度或是抑制区内豚草入侵的有效手段。     

Aquatic macrophytes, macroinvertebrate associations and water levels in a lowland Tasmanian river

Aquatic macrophytes are a common habitat for macroinvertebrates and may occupy depth zones in the littoral region of lowland rivers. Studies have indicated that different species of macrophyte typically support different assemblages, abundances and numbers of species of macroinvertebrates. This has often been attributed to differences in the dissectedness of stems and leaves of the macrophytes, resulting in differences in the surface area and/or the number of microhabitats available to invertebrates. I set out to measure the abundance and taxonomic richness and to describe the macroinvertebrate assemblages associated with three species of aquatic macrophyte in a pool in the Macquarie River, Tasmania and to examine responses of these variables to changes in water levels over summer. The macrophyte species sampled wereMyriophyllum simulans/variifolium, Triglochin procera} and Eleocharis sphacelata, each one differing in the dissectedness of its stems and leaves and its location in the littoral zone. Whereas the greatest abundance of macroinvertebrates was found associated in all months (i.e. at all water levels) with the structurally complex and shallowest macrophyte species, Myriophyllum, the number of taxa associated with this species was in several cases lower than for the structurally simpler and deeper water Triglochin and Eleocharis. While water depth and total plant biomass of samples were often correlated with invertebrate abundance and richness, these relationships were different for each macrophyte species. Of the nine most common invertebrate taxa collected from all samples, the abundances of more than half showed consistent differences among macrophyte species across months, two showed differences among macrophytes, but with an interaction with month and two showed no differences among macrophytes. There were major differences in the invertebrate assemblages associated with each macrophyte species in any one month, however, there was also a large turnover of taxa associated with the species of macrophytes from one month to the next. Changes in water level and concomitant changes in environmental variables are suggested as factors influencing the invertebrate fauna in the littoral zone of the pool of the Macquarie River. It is thus important for river managers to be aware that species of macroinvertebrates are not evenly distributed across species of macrophyte and that water levels and their influence on macrophytes as invertebrate habitat may play an integral part in determining the abundance, richness and assemblage of invertebrates in rivers.     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

Spatial and temporal variability of macroinvertebrates in spawning and non-spawning habitats during a salmon run in Southeast Alaska

Spawning salmon create patches of disturbance through redd digging which can reduce macroinvertebrate abundance and biomass in spawning habitat. We asked whether displaced invertebrates use non-spawning habitats as refugia in streams. Our study explored how the spatial and temporal distribution of macroinvertebrates changed during a pink salmon (Oncorhynchus gorbuscha) spawning run and compared macroinvertebrates in spawning (riffle) and non-spawning (refugia) habitats in an Alaskan stream. Potential refugia included: pools, stream margins and the hyporheic zone, and we also sampled invertebrate drift. We predicted that macroinvertebrates would decline in riffles and increase in drift and refugia habitats during salmon spawning. We observed a reduction in the density, biomass and taxonomic richness of macroinvertebrates in riffles during spawning. There was no change in pool and margin invertebrate communities, except insect biomass declined in pools during the spawning period. Macroinvertebrate density was greater in the hyporheic zone and macroinvertebrate density and richness increased in the drift during spawning. We observed significant invertebrate declines within spawning habitat; however in non-spawning habitat, there were less pronounced changes in invertebrate density and richness. The results observed may be due to spawning-related disturbances, insect phenology, or other variables. We propose that certain in-stream habitats could be important for the persistence of macroinvertebrates during salmon spawning in a Southeast Alaskan stream.