Context-dependent species identity effects within a functional group of filter-feeding bivalves

We asked whether species richness or species identity contributed more to ecosystem function in a trait-based functional group, burrowing, filter-feeding bivalves (freshwater mussels: Unionidae), and whether their importance changed with environmental context and species composition. We conducted a manipulative experiment in a small river examining the effects of mussel assemblages varying from one to eight species on benthic algal standing crop across two sets of environmental conditions: extremely low discharge and high water temperature (summer); and moderate discharge and water temperature (fall). We found strong species identity effects within this guild, with one species (Actinonaias ligamentina) influencing accrual of benthic algae more than other species, but only under summer conditions. We suspect that this effect is due to a combination of the greater biomass of this species and its higher metabolic and excretion rates at warm summer temperatures, resulting in increased nitrogen subsidies to benthic algae. We also found that Actinonaias influenced the condition of other mussel species, likely through higher consumption, interference, or both. This study demonstrates that species within trait-based functional groups do not necessarily have the same effects on ecosystem properties, particularly under different environmental conditions.     

Priority effects and species sorting in a long paleoecological record of repeated community assembly through time

We studied the relative roles of environmental species sorting and priority effects in the assembly of ecological communities on long time scales, by analyzing community turnover of water fleas (Daphnia) in response to strong and recurrent environmental change in a fluctuating tropical lake. During the past 1800 years, Lake Naivasha (Kenya) repeatedly fluctuated between a small saline pond habitat during lowstands and a large freshwater lake habitat during highstands. Starting from a paleoecological reconstruction, we estimated the role of priority effects in Daphnia community assembly across 16 of these habitat turnovers and compared this with the response of the community to reconstructed changes in three environmental variables important for species sorting. Our results indicate that the best predictor of Daphnia community composition during highstands was the community composition just prior to the transition from lowstands to highstands. This reflects a long-lasting priority effect of late lowstand communities on highstand communities, arising when remnant lowstand populations fill newly available ecological space in the rapidly expanding lake habitat. Species sorting and priority effects had a comparable but relatively small influence on community composition during the lowstands. Moreover, these priority effects decayed rapidly with time as Daphnia communities responded to environmental change, in contrast with the highstand communities where priority effects lasted for several decades.     

Running to stand still: temperature effects on species richness, species turnover, and functional community dynamics

The information on temperature-mediated changes in biodiversity in local assemblages is scarce and mainly addresses the change in species richness. However, warming may have more consistent effects on species turnover than on the number of species. Moreover, very few studies extended the analysis of changes in biodiversity and species composition to questions of associated ecosystem functions such as primary production. Here, we synthesize 4 case studies employing microalgal microcosms within the Aquashift priority program to ask (1) do warming-related shifts in species richness correspond to changes in the rate of biomass production, (2) do similar relationships prevail for evenness, and (3) do warming-related shifts in species turnover stabilize or destabilize biomass production? Two of the four cases are previously unpublished, and for a third case, the link between diversity and functional consequences of temperature was not analyzed before. We found accelerated loss of species with warming in all cases. Biomass production was lower with lower species richness in most cases but increased with lower evenness. Most importantly, the relation between functional and compositional stability was different between cases: More rapid extinction resulted in more variable biomass in 2 cases conducted with a limited species pool, indicating that compositional destabilization relates to functional variability. By contrast, the only experiment with a large species pool (30 species) allowed previously rare species to become dominant in the community and showed more stable biomass at high turnover, indicating that compensatory dynamics (turnover) can promote functional stability. These 4 independent experiments highlight the need to consider both compositional and functional consequences of altered temperature regimes.     

不同进水有机物浓度下移动床生物膜反应器(SBMBBR)脱氮除磷特性

考察了不同进水有机物浓度下厌氧/好氧序批式移动床生物膜反应器（SBMBBR）污染物去除特性,实验结果表明,SBMBBR能够实现低碳源污水中氮和磷的同步去除,在进水TN和TP浓度分别为116.7 mg.L-1和11.5 mg.L-1、COD浓度为456 mg.L-1的条件下,TN和TP去除率分别达到94.3%和92.2%以上.反应器除磷是基于常规生物除磷和反硝化除磷过程实现的,脱氮主要是基于好氧段发生的同时硝化反硝化（SND）作用而完成.由于生物膜内部存在的DO扩散梯度,在好氧阶段混合液DO浓度不断提高的条件下反应器内具有良好SND反应的发生.进水COD浓度由149 mg.L-1提高至456 mg.L-1的过程中,反应器硝化效果不变,反硝化和除磷效果改善.反应器在好氧阶段pH值基本维持在7.0—7.1之间,为各类菌群的生长创造了条件.碱度变化较pH值更能反映硝化和反硝化反应发生的程度.反应器中微生物相丰富,生物膜以丝状菌为骨架,其上附着大量的球状菌和杆状菌,而悬浮活性污泥中丝状菌较少,形成了由细菌、真菌到原生动物和后生动物的复杂的生态体系,为系统取得稳定的污水处理效果提供了有效的保证.     

Effect of canopy removal on a rocky shore community metabolism and structure

The effects of the permanent removal of the canopy-forming alga Fucus serratus was studied in terms of both functioning and diversity on a mid–low rocky shore ecosystem in the south-west English Channel (48°N 43.686′, 3°W 59.282′). Ecosystem functioning was examined as net or gross primary productivity (NPP or GPP) and respiration (Resp) measured through CO₂ fluxes. Diversity was examined as number and composition of species. Measurements were performed in situ, during emersion times, without altering target assemblages. The experiment was designed with two treatments [control (C) or canopy removed (CR)] and five replicates, and was conducted over an 18-month period (from February 2006 to August 2007) to integrate the seasonal variability. The mean GPP and Resp were severely reduced in CR treatment compared to control throughout the survey. The mean NPP was not affected at first, due to the development of opportunistic green algae, but was drastically reduced after 9 months of experiment. The canopy removal affected neither the number of species nor their distribution among trophic groups, and the algal community was only slightly affected. The abundance and biomass of mobile invertebrates, however, were greatly reduced in the absence of canopy. This indicates an important effect of the dominant alga on the higher trophic levels of the community. At this tidal level, the canopy did not seem to affect the community by dampening the environmental stress but by providing food, habitat or both.     

两级人工湿地用于村镇污水脱氮的长期运行特性研究

对沙田人工湿地稳定运行后的情况进行了长达6 a的监测,探讨了不同流态的两级潜流人工湿地长期运行的脱氮效能、不同形态氮的空间转化规律和几种主要影响因素。研究表明,在整个运行期间人工湿地的脱氮能力呈现低-高-低的变化趋势,这个过程的发生和湿地内部堵塞物的累积关联密切。在华南地区,季节变化对湿地脱氮的影响依然十分强烈：一方面,体现为季节温差对脱氮效果的明显影响,低温月份采用低负荷方式运行,TN去除率也仅有36.0%～47.6%,而高温期采用高负荷的运行方式,大多数月份TN去除率也能达到45.4%以上;另一方面,季节性雨量的不同会引起进水质量浓度的变化,导致湿地系统TN负荷率的改变,进而显著影响脱氮效果。有机氮、NH3-N、NO2--N、NO3--N在湿地内部存在明显的转化迹象,沿流程随着有机物的大量消耗,NO3--N去除能力逐渐下降并在湿地的末端出现了累积现象。通常情况下,由于采用了跌水曝气等措施,DO在湿地内的大部分区域都维持了较高水平,未对脱氮效果产生明显不利影响。     

Sperm removal and ejaculate size correlate with chelae asymmetry in a freshwater crayfish species

Asymmetry in traits of sexual relevance may impair copulation behaviour and sexual performance of males, ultimately resulting in a fitness cost. Freshwater crayfish males use chelae, a sexually selected trait, to secure and position the female prior to and during mating. Thus, a relatively large chelae asymmetry, resulting from accidental loss and regeneration of one cheliped after autotomy, could have great consequences for male sexual behaviour. We studied copulatory behaviour and sperm expenditure of males paired to a mated female in Austropotamobius italicus, a freshwater crayfish species where both male and female mate multiply and where last-mating males are able to actively remove previously deposited sperm. We aimed at assessing whether male sperm removal and expenditure varied according to sperm allocated by first-mating males, and according to copulation behaviour and phenotypic traits (carapace length, chelae length and relative chelae asymmetry) of second-mating males. Second-mating males did not adjust their ejaculate size in relation to first-mating male ejaculate, nor to the first-mating male’s sperm removed. Moreover, the amount of sperm removed by second-mating males increased with increasing first-mating males ejaculate size, and first-mating male sperm remaining after removal did not correlate with the original first-mating male ejaculate size. Interestingly, the amount of sperm removed by second-mating males decreased with increasing relative chelae asymmetry, while increasing with male body size. However, second-mating (but not first-mating) asymmetric-clawed males produced larger ejaculates than symmetric-clawed ones. Importantly, the proportion of second-mating male sperm remaining after the two matings did not vary with relative chelae asymmetry nor with body size of second-mating males. Thus, small, asymmetric-clawed crayfish males appear to adopt sperm allocation tactics that allow them to fully compensate for their inferior sperm removal ability.     

典型城区与郊区环境两种灌木生理生化特性比较

选取北京城区与郊区典型样点,针对栽植多年的大叶黄杨（Euonymus japonicus）和小叶黄杨（Buxus sinica subsp.sinica var.parvifolia）成龄植株,分别于春、夏、秋3季取样测定叶片的各项生理生化指标,研究了城区与郊区环境下2种灌木的生理生化特性的差异。结果表明：生长前期（春季）城区车公庄2种灌木的叶绿素总量高于郊区植物园,经过一段时期之后低于植物园（夏秋）。Chlb的城郊变化幅度大于Chla。Chla的变化趋势与Chlb相反造成了Chla/b的大幅度变化。大叶黄杨的叶绿素总量和Chla/b比小叶黄杨受城区环境影响更大。城区环境下2种灌木自由水/束缚水降低,脯氨酸质量分数、丙二醛质量摩尔浓度升高,说明城区环境下2种灌木生理代谢水平降低,膜脂过氧化。城区环境下大叶黄杨的自由水/束缚水、脯氨酸质量分数的变化率较小叶黄杨高,2种灌木丙二醛的变化率相近。综合来看,城区环境下大叶黄杨的生理生化反应比小叶黄杨更敏感。     

The role of local and landscape level measures of greenness in modelling boreal plant species richness

Measurements of primary productivity and its heterogeneity based on satellite images can provide useful estimates of species richness and distribution patterns. However, species richness at a given site may depend not only on local habitat quality and productivity but also on the characteristics of the surrounding landscape. In this study we investigated whether the predictions of species richness of plant families in northern boreal landscape in Finland can be improved by incorporating greenness information from the surrounding landscape, as derived from remotely sensed data (mean, maximum, standard deviation and range values of NDVI derived from Landsat ETM), into local greenness models. Using plant species richness data of 28 plant families from 440 grid cells of 25 ha in size, generalized additive models (GAMs) were fitted into three different sets of explanatory variables: (1) local greenness only, (2) landscape greenness only, and (3) combined local and landscape greenness. The derived richness–greenness relationships were mainly unimodal or positively increasing but varied between different plant families, and depended also on whether greenness was measured as mean or maximum greenness. Incorporation of landscape level greenness variables improved significantly both the explanatory power and cross-validation statistics of the models including only local greenness variables. Landscape greenness information derived from remote sensing data integrated with local information has thus the potentiality to improve predictive assessments of species richness over extensive and inaccessible areas, especially in high-latitude landscapes. Overall, the significant relationship between plants and surrounding landscape quality detected here suggests that landscape factors should be considered in preserving species richness of boreal environments, as well as in conservation planning for biodiversity in other environments.     

Nitrogen cycling in a marine planktonic food chain: Nitrogen fluxes through the principal components and the effects of adding copper

Nitrogen cycling in a natural planktonic food chain was followed before and after perturbation by copper (10 g l^-1). Observations were separated into effects from artificial containment of the food chain and effects from the addition of copper. An initial reduction of seston-N and subsequent recovery, uniformly high settling rates of suspended particulate-N, and general decrease in zooplankton-N were attributed to the physical limitations imposed by the enclosures (e.g. elimination of normal mixing processes). Acute inhibition of nitrate assimilation by the seston, acute elevation of seston-N settling and chronic inhibition of zooplankton feeding, along with accelerated decline in zooplankton stocks, were attributed to copper stress. Since zooplankton stocks and fluxes were most drastically affected by copper, its overall effect on N-cycling through the planktonic food chain would depend largely upon the importance of zooplankton in regulating the production of the remaining seston.     

三江平原小叶章湿地生态系统对氮磷的净化效率

选取面积为1 600 m2的小叶章(Deyeuxia angustifolia)湿地进行小区域模拟试验。试验设割草区、1倍浓度区、静态对照区和2倍浓度区4个处理,研究N、P输入浓度,割草和输出径流对三江平原小叶章湿地生态系统净化N、P污染物的影响。结果表明,在小叶章生长期和成熟期,湿地生态系统对N、P保持较稳定的净化,N、P平均去除率分别为85.62%和85.94%。湿地水体中TN和NH4 -N的累积浓度显著相关。输入浓度、割草和输出径流都影响着湿地生态系统对N、P的净化和N、P在系统内的分配。输入浓度增大时,N、P去除率明显降低;割草会降低N去除率,但对P去除率影响不大;少量输出径流对N、P去除影响不大。三江平原小叶章湿地生态系统对输入的N、P具有显著净化作用,总净化率可达97.97%和99.05%。     

Easier removal of nonylphenol and naphthalene pollutants in wet weather revealed by Markov chains modeling

Environmental Chemistry Letters - The distribution and fate of organic pollutants determines pollution control strategies and definition of environmental standards. So far, few models predict...     

Nitrogen fixation (acetylene reduction) associated with macroalgae in a coral-reef community in the Bahamas

N₂ fixation (C₂H₂ reduction) was associated with several species of macroalgae on a coral reef near Grand Bahama Island. The highest rates were associated with Microdictyon sp. (Chlorophyceae) and Dictyota sp. (Phaeophyceae). Extensive mats of filamentous blue-green algae, not heterotrophic bacteria, were the N₂ fixing agents: in experiments with samples of Microdictyon sp., the activity was lightdependent and not stimulated by organic compounds under either aerobic or anaerobic conditions. Assays in situ, at 20 m depth, and on shipboard, gave similar rates of N₂ fixation; the cyanophytes presumably have pigment adaptations to function in blue light. The maximum rate of N₂ fixation, associated with Microdictyon sp., was 3.8 g N fixed g dry weight^-1 h^-1. Coral-reef communities flourish in nutrientimpoverished waters, and therefore any input of nitrogen is probably important in stabilizing such ecosystems.     

茶树上游猎型蜘蛛功能团对景观低碳管理模式的反应

目前温室效应加剧、生物多样性减少以及农田环境污染加重等问题已成为关注的焦点。由于蜘蛛是环境变化的指示标志和农业生中的重要捕食性天敌。通过选择茶树上游猎型蜘蛛为研究对象,调查分析景观低碳管理模式茶园（JG）与习惯管理模式茶园（XG）对茶树上游猎型蜘蛛组成、结构以及群落数量特征的影响。结果表明：JG茶园茶树上游猎型蜘蛛种类较多,有8个科,18个种;XG茶园有6个科,13个种;优势种和丰盛种中,JG茶园个体数为375头、195头,比XG茶园分别提高1.16、0.33倍。JG与XG两类茶园的蜘蛛种群数量特征在年龄结构、性别比例方面存在显著差异（P＜0.001）;JG茶园雌蛛个体数为157头、雄蛛个体数为320头,是XG茶园的1.12和1.32倍;JG茶园成蛛与幼蛛个体数是XG茶园的1.25和0.87倍,成蛛比幼蛛高出46.32%,而XG茶园成蛛比幼蛛高出21.82%;JG茶园蜘蛛种群数量特征发生动态显著高于XG茶园（P＜0.05）。JG茶园蜘蛛多样性指数和丰富度指数比XG茶园增加16.46%和12.14%。主成分分析表明,第一、二主成分累计贡献率达50.47%, JG茶园位置明显接近光盔蛛科（Liocranidae）、圆颚蛛科（Corinnidae）、跳蛛科（Salticidae）和管巢蛛科（Clubionidae）等蜘蛛群落数量分布较多的方向偏移,而XG茶园则向盗蛛科（Pisauridae）和猫蛛科（Oxyopidae）等蜘蛛群落数量分布较少方向靠近。由此可知,茶园实行低碳管理,实行复合种植,有利于减少CO2气体排放,保护茶树上游猎型蜘蛛种类多样性,提高对害虫的自然控制能力,为茶园固碳增汇,茶叶产值提高打下了基础。     

耐热石油烃降解混合菌的筛选及其群落结构分析

对克拉玛依采集的部分石油污染土壤进行了筛选,得到了5组石油烃高效降解混合菌,其中混合菌KL9-1在45℃的条件下,通过7 d的降解,稀油的降解率达到43.27%,稠油的降解率达到20.09%。混合菌KL9-1经过多次分离纯化后,获得3株具有石油烃降解能力的优势单菌,3株单菌对稀油的降解率都在30%以上。结合分离单菌株的形态、生理生化特征和16S rDNA基因序列的分析结果,初步鉴定KL9-1-1为Pseudomonas putida,KL9-1-2和KL9-1-3为Pseudomonas sp.。     

三江平原小叶章湿地生态系统对氮磷的净化效率

选取面积为1600m^2的小叶章（Deyeuxia angustifolia）湿地进行小区域模拟试验。试验设割草区、1倍浓度区、静态对照区和2倍浓度区4个处理，研究N、P输入浓度，割草和输出径流对三江平原小叶章湿地生态系统净化N、P污染物的影响。结果表明，在小叶章生长期和成熟期，湿地生态系统对N、P保持较稳定的净化，N、P平均去除率分别为85．62％和85．94％。湿地水体中TN和NH4^＋-N的累积浓度显著相关。输入浓度、割草和输出径流都影响着湿地生态系统对N、P的净化和N、P在系统内的分配。输入浓度增大时，N、P去除率明显降低；割草会降低N去除率，但对P去除率影响不大；少量输出径流对N、P去除影响不大。三江平原小叶章湿地生态系统对输入的N、P具有显著净化作用，总净化率可达97．97％和99．05％。     

Global change shifts vegetation and plant-parasite interactions in a boreal mire

The aim of this study was to detect vegetation change and to examine trophic interactions in a Sphagnum-dominated mire in response to raised temperature and nitrogen (N) addition. A long-term global-change experiment was established in 1995, with monthly additions of N (30 kg x ha(-1) x yr(-1)) and sulfur (20 kg x ha(-1) x yr(-1)) during the vegetation period. Mean air temperature was raised by 3.6 degrees C with warming chambers. Vegetation responses were negligible for all treatments for the first four years, and no sulfur effect was seen during the course of the experiment. However, after eight years of continuous treatments, the closed Sphagnum carpet was drastically reduced from 100% in 1995 down to 41%, averaged over all N-treated plots. Over the same period, total vascular plant cover (of the graminoid Eriophorum vaginatum and the two dwarf-shrubs Andromeda polifolia and Vaccinium oxycoccos) increased from 24% to an average of 70% in the N plots. Nitrogen addition caused leaf N concentrations to rise in the two dwarf-shrubs, while for E. vaginatum, leaf N remained unchanged, indicating that the graminoid to a larger extent than the dwarf-shrubs allocated supplemented N to growth. Concurrent with foliar N accumulation of the two dwarf-shrubs, we observed increased disease incidences caused by parasitic fungi, with three species out of 16 showing a significant increase. Warming caused a significant decrease in occurrence of three parasitic fungal species. In general, decreased disease incidences were found in temperature treatments for A. polifolia and in plots without N addition for V. oxycoccos. The study demonstrates that both bryophytes and vascular plants at boreal mires, only receiving background levels of nitrogen of about 2 kg x ha(-1) x yr(-1), exhibit a time lag of more than five years in response to nitrogen and temperature rise, emphasizing the need for long-term experiments. Moreover, it shows that trophic interactions are likely to differ markedly in response to climate change and increased N deposition, and that these interactions might play an important role in controlling the change in mire vegetation composition, with implications for both carbon sequestration and methane emission.     

Functional identity and functional structure change through succession in a rocky intertidal marine herbivore assemblage

Despite the great interest in characterizing the functional structure and resilience of functional groups in natural communities, few studies have examined in which way the roles and relationships of coexisting species change during community succession, a fundamental and natural process that follows the release of new resources in terrestrial and aquatic ecosystems. Variation in algal traits that characterize different phases and stages of community succession on rocky shores are likely to influence the magnitude, direction of effects, and the level of redundancy and complementarity in the diverse assemblage of herbivores. Two separate field experiments were conducted to quantify per capita and population effects and the functional relationship (i.e., redundancy or complementarity) of four herbivore species found in central Chile during early and late algal succession. The first experiment examined grazer effects on the colonization and establishment of early-succession algal species. The second experiment examined effects on the late-successional, dominant corticated alga Mazzaella laminarioides. Complementary laboratory experiments with all species and under natural environmental conditions allowed us to further characterize the collective effects of these species. We found that, during early community succession, all herbivore species had similar effects on the ephemeral algae, ulvoids, but only during the phase of colonization. Once these algae were established, only a subset of the species was able to control their abundance. During late succession, only the keyhole limpet Fissurella crassa could control corticated Mazzaella. The functional relationships among these species changed dramatically from redundant effects on ephemeral algae during early colonization, to a more complementary role on established early-successional algae, to a dominant (i.e., keystone) effect on late succession. This study highlights that functional relationship within consumer assemblages can vary at different phases and times of community succession. Differentiation in herbivore roles emphasizes the need to evaluate consumer's impacts through different times of community succession, and through experimental manipulations to make even broad predictions about the resilience or vulnerability of diverse intertidal assemblages to human disturbances.     

Population and species diversity fluctuations in a rocky intertidal community relative to severe aerial exposure and sediment burial

From a series of 10 quarterly assessment between October 1975 and May 1978 (inclusive), fluctuations in abundance were determined for macroinvertebrates and macrophytes of a rocky intertidal community on Santa Cruz Island, California, USA. During afternoon low tides in late fall and winter of the first 2 yr of study, organisms of the upper and middle intertidal zones were subjected to prolonged aerial exposure. Many species there tolerated this exposure, but die-backs occurred for a barnacle (Chthamalus fissus/C. dalli) and several algae (Endocladia muricata, Pelvetia fastigiata f. gracilis, Corallina officinalis var. chilensis, Corallina vancouveriensis, Cylindrocarpus rugosus and Codium fragile). These die-backs were succeeded by blooms of Ulva californica and Porphyra perforata. In the upper and middle intertidal zones, the major cover organisms that could tolerate prolonged aerial exposure were disproportionately prevalent and appeared to be maintained by the periodic repression of species intolerant to such exposure. In February 1978, heavy rainfall caused sediment inundation of the middle and lower intertidal zones. This event was followed by declines in abundance of a barnacle (Tetraclita rubescens) and several algae (Pelvetia fastigiata f. gracilis and corallina spp.). Shannon-Wiener H species diversity fluctuated from a high in October to a low the following May during both 1975–1976 and 1976–1977 in conjunction with the period of increased daytime aerial exposure in late fall and winter. A further decline in diversity following sediment inundation in February 1978 contributed to a long-term trend of declining H species diversity (3.06 in October 1975 to 1.87 in May 1978). We hypothesize that predictable late fall to winter aerial exposure stresses, in combination with a random physical disturbance (sediment burial), exceeded an optimal intermediate level of disturbance predicted for maximal species diversity.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.