氮素添加对黄土高原典型草原长芒草氮磷重吸收率及C:N:P化学计量特征的影响

 为了解氮素沉降对草地群落的影响, 通过人工氮肥添加模拟试验, 研究了黄土高原天然草地优势植物长芒草(Stipa bungeana)在不同施氮水平下叶片和立枯物碳(C)、氮(N)、磷(P)元素含量的变化特征, 探讨了N素增加对N、P重吸收率和C : N : P化学计量比的影响及其内在联系。结果表明: 氮素添加显著增加了长芒草叶片的C、N和立枯物的N、P含量, 对叶片P和立枯物C含量无显著影响; 氮素添加显著降低了长芒草的N、P重吸收率, 对照处理的N、P重吸收率最高, 分别为60.35%和
71.75%, 并且, 在相同氮素处理条件下P的重吸收率显著大于N重吸收率; 随着氮素添加量的增大, 叶片的C : N降低, N : P和C : P增加, N : P为18.25–29.01。研究表明, 黄土高原天然草地群落主要受P限制, 随氮素沉降增加, P限制进一步加剧; 长芒草较高的N、P重吸收率是保证其在贫瘠的土壤中生存的重要机制。     

Correlations between nutrient concentrations and zooplankton populations in a mesotrophic reservoir

1. A year-round study was conducted in a mesotrophic reservoir to determine the dynamics of zooplankton populations as a function of food availability (edible phytoplankton), nutrient concentration, temperature and hydraulic regime.
2. Rotifer biomass was correlated with soluble reactive phosphorus (SRP) concentration. The abundance of the rotifers Keratella cochlearis and Anuraeopsis fissa were not correlated with food availability (measured by chlorophyll and cell counts) but showed a strong dependence on P availability. Another rotifer, Synchaeta oblonga , and crustacean species were not related to nutrient availability but seemed to be dependent on food concentrations, especially of some phytoplankton taxa.
3. In this field study, rotifers seemed more susceptible than Daphnia or copepods to P-limitation. Among rotifer species, Keratella seemed to be more susceptible than Anuraeopsis to P limitation. Different susceptibilities of zooplankton species to nutrient limitation may be important in explaining the dynamics of these organisms in natural situations. Further analyses are warranted to clarify the interactions between nutrient limitation and energy limitation among zooplankton.     

N:P stoichiometric changes via species turnover in arid versus saline desert environments

1. Aridity and salinity have a key role in driving physiological and ecological processes in desert ecosystems. However, how community‐scale foliar nutrients respond to aridity and salinity, and how these responses might vary with community composition along aridity and salinity gradients is unclear. We hypothesize that the response will be a shift in community stoichiometric values resulting from nutrient variability of shared species and unique species (site‐specific species), but little research has addressed the relative contribution of either component.
2. We analyzed the community‐scale stoichiometric response of a desert community of perennial plants along an aridity and salinity transect by focusing on foliar nitrogen (N) and phosphorous (P) concentrations and N:P ratios. After evaluating the shared and unique species variability, we determined their relative contribution to the community stoichiometric response to aridity and salinity, reflected by changes in nonweighted and weighted community‐average values.
3. Community‐scale stoichiometry decreased significantly under aridity and salinity, with significantly consistent changes in nonweighted and weighted community‐average stoichiometry for most shared and unique species measurements. The relative contribution of unique species shifts to the changes in community stoichiometry was greater (15%–77%) than the relative contribution of shared species shifts (7%–45%), excluding the change in weighted P concentration under aridity. Thus, the shifts of unique species amplified the community stoichiometric response to environmental changes.
4. Synthesis. These results highlighted the need for a more in‐depth consideration of shared and unique species variability to understand and predict the effects of environmental change on the stoichiometry of plant communities. Although variation in community stoichiometry can be expected under extreme aridity and salinity conditions, changes of unique species could be a more important driver of the stoichiometric response of plant communities.

     

Glacier melting and stoichiometric implications for lake community structure: zooplankton species distributions across a natural light gradient

Glaciers around the globe are melting rapidly, threatening the receiving environments of the world's fresh water reservoirs with significant changes. The meltwater, carried by rivers, contains large amounts of suspended sediment particles, producing longitudinal gradients in the receiving lakes. These gradients may result in changes in the light : nutrient ratio that affect grazer performance by altering elemental food quality. Thus, glacial melting may induce a shift in the phytoplankton carbon : nutrient ratio and hence influence the dominance of herbivorous zooplankton through stoichiometric mechanisms. To test this hypothesis, we combined field and experimental data, taking advantage of a natural light intensity gradient caused by glacial clay input in a deep oligotrophic Patagonian lake. Across this gradient, we evaluated the abundances of two consumer taxa with different phosphorus requirements, the copepod Boeckella gracilipes and the cladoceran Daphnia commutata, using a six‐station transect along the lake. We found significant differences in light : nutrient ratio and stoichiometric food quality of the seston, together with a switch from dominance of P‐rich Daphnia in low carbon : nutrient stations to dominance of low‐P copepods in high carbon : nutrient stations. The laboratory experiments confirmed that the difference in the carbon : nutrient ratio across the gradient is sufficient to impair Daphnia growth. The overall patterns are consistent with our prediction that shifts in the environmental light : nutrient ratio as a result of glacial melting would contribute to shifts in the dominance of stoichiometrically contrasting taxa in consumer guilds.     

Factors influencing nitrogen and phosphorus excretion rates of fish in a shallow lake

Summary 1. Fish excretion can be an important source of nutrients in aquatic ecosystems. Nitrogen (N) and phosphorus (P) excretion rates are influenced by many factors, including fish diet, fish growth rate and fish size. However, the relative influence of these and other factors on community‐level excretion rates of fish is unknown. 2. We used bioenergetics modeling to estimate excretion rates of eight fish species in a shallow, Minnesota (U.S.A.) lake over four months in both 2004 and 2005. Excretion rates of each species were summed for community‐level N and P excretion rates, as well as the N : P ratio of excretion. We then used a model‐selection approach to assess ability of independent variables to predict excretion rates, and to identify the most parsimonious model for predicting N : P excretion ratios and P and N excretion rates at the community scale. Predictive models were comprised of the independent variables water temperature and average fish density, fish size, fish growth rate, nutrient content of fish and nutrient content of fish diets at the community scale. 3. Fish density and nutrient content of fish diets (either N or P) were the most parsimonious models for predicting both N and P excretion rates, and explained 96% and 92% of the variance in N and P excretion, respectively. Moreover, fish density and nutrient models had 1400‐fold more support for predicting N and 21‐fold more support for predicting P excretion relative to models based on fish density only. Water temperature, fish size, fish growth rates and nutrient content of fish showed little influence on excretion rates, and none of our independent variables showed a strong relationship with N : P ratios of excretion. 4. Past work has focused on the importance of fish density as a driver of fish excretion rates on a volumetric basis. However, our results indicate that volumetric excretion rates at the community scale will also change substantially in response to changes in relative abundance of fish prey or shifts in relative dominance of planktivores, benthivores, or piscivores. Changes in community‐scale excretion rates will have subsequent influences on algal abundance, water clarity, and other ecosystem characteristics.     

中国四种森林类型主要优势植物的C:N:P化学计量学特征

为了评价不同森林类型的生态化学计量特征的差异, 以吉林长白山温带针阔混交林、广东鼎湖山亚热带常绿阔叶林、云南西双版纳热带季雨林和江西千烟洲亚热带人工针叶林为研究对象, 通过对2007年4月-2008年5月4种典型区域森林植物叶片和凋落物的碳(C)氮(N)磷(P)元素质量比与N、P再吸收率的分析, 探讨了4种森林类型N、P养分限制和N、P养分再吸收的内在联系。结果表明: 1)从森林类型上看, 温带针阔混交林叶片的C : N : P为321 : 13 : 1, 亚热带常绿阔叶林叶片的C : N : P为561 : 22 : 1, 热带季雨林叶片的C : N : P为442 : 19 : 1, 亚热带人工针叶林叶片的C : N : P为728 : 18 : 1; 凋落物的C : N : P也是亚热带人工林最高, 达1 950 : 27 : 1, 温带针阔混交林的最低, 为552 : 14 : 1, 热带季雨林的为723 : 24 : 1, 亚热带常绿阔叶林的为1 305 : 35 : 1, 不同森林类型凋落物的C : N : P的计量大小关系与叶片的结果一致; 2)从植物生活型上看, 常绿针叶林叶片的C : N均显著高于常绿阔叶林及落叶阔叶林; 叶片C : P与森林类型的关系并不十分密切; 常绿阔叶林叶片的N : P最高, 常绿针叶林次之, 落叶阔叶林最低; 3)植物叶片的N : P与月平均气温有显著的负相关关系, 但叶片的C : P基本不受月平均气温影响, 叶片的C、N、P计量比与降水的线性关系不显著; 4)高纬度地区的植物更易受N元素限制, 而低纬度地区植物更易受P元素的限制; 但受N或P限制的植物并不一定具有高的N或P再吸收率。研究结果表明, 不同类型森林的叶片与凋落物的化学计量特征具有一致性, 但是环境因子对不同类型植物化学计量比的影响并不相同。     

Mismatch between dietary requirements for lipid by a predator and availability of lipid in prey

Growth is an important factor in predicting an organism's overall success as an adult. Understanding how abiotic and biotic factors influence body size is key to predicting how environmental changes will impact organisms and predicting optimal behaviors under varying conditions. Food items can vary widely in nutrient content and this variation can affect growth. We tested how the quantity and macronutrient content of live prey affected the growth of juvenile jumping spiders, Phiddipus audax, using 420 spiderlings raised on one of 21 diets. In addition, we measured the nutrient content of prey from the field and compared average prey nutrient content with the diet at which spiders maximized growth. Our results show that the quantity and nutrient content of prey have significant and interacting effects on the growth of an actively hunting predator. In particular, spiders grew larger in body size (tibia/patella length and posterior lateral eye width) and were heavier on diets with higher lipid content. Yet, data on the nutrient content of prey from the field shows that most potential prey have high protein and low lipid content in their bodies. This apparent mismatch between spider nutritional requirements and prey nutrient availability likely presents a challenge to spiders seeking to maximize growth in the field. Several hypotheses for how these predators overcome this challenge to reach the large body sizes observed in the field are discussed.     

C,N and P fertilization in an Amazonian rainforest supports stoichiometric dissimilarity as a driver of litter diversity effects on decomposition

Plant leaf litter generally decomposes faster as a group of different species than when individual species decompose alone, but underlying mechanisms of these diversity effects remain poorly understood. Because resource C : N : P stoichiometry (i.e. the ratios of these key elements) exhibits strong control on consumers, we supposed that stoichiometric dissimilarity of litter mixtures (i.e. the divergence in C : N : P ratios among species) improves resource complementarity to decomposers leading to faster mixture decomposition. We tested this hypothesis with: (i) a wide range of leaf litter mixtures of neotropical tree species varying in C : N : P dissimilarity, and (ii) a nutrient addition experiment (C, N and P) to create stoichiometric similarity. Litter mixtures decomposed in the field using two different types of litterbags allowing or preventing access to soil fauna. Litter mixture mass loss was higher than expected from species decomposing singly, especially in presence of soil fauna. With fauna, synergistic litter mixture effects increased with increasing stoichiometric dissimilarity of litter mixtures and this positive relationship disappeared with fertilizer addition. Our results indicate that litter stoichiometric dissimilarity drives mixture effects via the nutritional requirements of soil fauna. Incorporating ecological stoichiometry in biodiversity research allows refinement of the underlying mechanisms of how changing biodiversity affects ecosystem functioning.     

黄河三角洲贝壳堤湿地优势灌木碳、氮、磷化学计量特征

为探讨黄河三角洲海岸带湿地不同水盐条件下植物叶片化学计量特征的季节动态及植物生长的限制性营养元素,以滨州贝壳堤岛与湿地国家级自然保护区内的柽柳、杠柳、酸枣3种优势灌木为研究对象,于2017-2018年的生长季（5-10月）每月定期采集叶片样品,测定叶片C、N、P含量。结果表明,生长季内3种灌木叶片C含量呈逐渐上升趋势;叶片N、P含量呈先下降后上升趋势,说明3种灌木采用防御性生活史策略适应盐生和干旱生境。3种灌木叶片C平均含量分别为（399.65±2.66）mg/g、（424.32±1.59）mg/g、（437.47±1.08）mg/g,低于全国（455.1 mg/g）及全球（461.6 mg/g）水平,呈现盐生生境下较低的植物碳储存能力。3种灌木叶片N和P平均含量分别为（30.14±0.26）mg/g和（1.81±0.03）mg/g、（23.18±0.38）mg/g和（2.06±0.04）mg/g、（27.36±0.49）mg/g和（2.01±0.03）mg/g,显著高于全国（N∶20.2 mg/g,P：1.46 mg/g）及全球（N∶19.3-20.1 mg/g,P：1.11-1.42 mg/g）水平。叶片C∶N∶P比呈先上升后下降的趋势,叶片P含量对C∶N∶P比变化具有主导作用。3种灌木叶片C∶N∶P质量比分别为246∶17∶1、224∶12∶1、237∶14∶1,说明柽柳的水盐胁迫适应能力高于杠柳和酸枣。从叶片N、P化学计量特征看,生长季内,柽柳生长一定程度上受土壤P限制,杠柳生长受到土壤N限制,酸枣生长则受土壤N、P共同限制,说明3种灌木的生物地球化学生态位发生了分化,避免了对同种资源的竞争,利于物种共生。     

Changes in stoichiometric constraints on epilithon and benthic macroinvertebrates in response to slight nutrient enrichment of mountain rivers

1. To assess changes in stoichiometric constraints on stream benthos, we measured elemental composition of epilithon and benthic macroinvertebrates in intrinsically P‐limited mountain rivers, upstream and downstream of low‐level anthropogenic nutrient enrichment by effluents of municipal wastewater treatment plants. 2. While there was a broad range in the elemental composition of epilithon (C : P ratios of 200–16 500, C : N ratios of 8–280, N : P ratios of 8–535) and heptageniid mayfly scrapers (C : P ratios of 125–300, C : N ratios of 5.1–7.2, N : P ratios of 20–60), the average C : P ratio of epilithon was 10‐fold lower and the average C : N ratio twofold lower at more nutrient‐rich downstream sites. Nutrient ratios in benthic macroinvertebrates were lower than in epilithon and varied little between relatively nutrient‐poor and nutrient‐rich sites. 3. We modified the existing definition of producer‐consumer elemental imbalance to allow for variation in consumer nutrient content. We defined this ‘non‐homeostatic’ imbalance as the perpendicular distance between the producer and consumer C : P, C : N, or N : P ratios, and the 1 : 1 line. 4. At P‐limited sites, the estimated mayfly N : P recycling ratio was higher than the N : P ratio in epilithon, suggesting nutrient recycling by consumers could accentuate P‐limitation of epilithon. 5. Measuring the degree of producer–consumer nutrient imbalance may be important in predicting the magnitude of effects from nutrient enrichment and can help elucidate the causes and consequences of ecological patterns and processes in rivers.     

Effects of nutrient (N,P, C) enrichment,grazing and depth upon littoral periphyton of a softwater lake

Three field experiments were performed in Lake Lacawac, PA to determine the importance of potentially limiting nutrients relative to other factors (grazing, depth) in structuring shallow water algal periphyton communities. All three experiments measured periphyton growth (as chlorophyll-a, AFDM or biovolumes of the algal taxa) on artificial clay flower pot substrates which released specified nutrients to their outer surfaces.Control of standing crop by nutrient supply rate vs. grazing was examined in Expt. I. Substrates releasing excess N and P, together with one of 4 levels of C (as bicarbonate) were placed either inside or outside exclosures designed to reduce grazer densities. Chlorophyll-a rose from 1.1–25.6 µg.cm^–2, and some dominant taxa (e.g., Oedogonium, Nostoc, Anacystis) were replaced by others (e.g., Scenedesmus, Cryptomonas) as bicarbonate supply increased. Reductions in invertebrate density did not significantly affect chlorophyll-a at any of the nutrient levels.Reasons for the species shift were further evaluated in Expt. II, using a minielectrode to measure the elevation of pH within the periphyton mat through photosynthetic utilization of bicarbonate. The pH adjacent to pots diffusing N, P and large quantities of bicarbonate, and supporting high chlorophyll-a densities of 32 µg cm^–2, averaged 10.0 compared to 6.3 in the water column. Pots diffusing only N and P supported 0.7 µg chlorophyll-a cm^–2 and elevated pH to 8.2. We suspect that bicarbonate addition favored efficient bicarbonate users (e.g., Scenedesmus), while inhibiting other taxa (e.g., Oedogonium) because of the attendant high pH.Expt. III was designed to test effects of depth (0.1 m vs. 0.5 m) and N (NH₄ ⁺ vs. NO₃ ^–) upon the growth response to bicarbonate observed in Expts. I and II. Similar standing crop and species composition were noted on pots at 0.1 m vs. 0.5 m. Enrichment with NH₄ ⁺ vs. NO₃ ^– also appeared to have little effect upon the periphyton community.Shallow water periphyton communities in Lake Lacawac, when supplied with sufficient N and P, appear to show a distinctive response to increasing bicarbonate concentration and pH which is robust to moderate variation in grazer densities, distance from the water surface, and the form of N enrichment.     

Role of phosphorus and nitrogen for bacteria and phytopankton development in a large shallow lake

Nutrient (P and N) enrichment experiments in small enclosures (20 l) were carried out to determine P and/or N limitation of bacterioplankton in Lake Võrtsjärv. The specific interest of the study was to test if it is possible to detect nutrient `physiological' or growth (rate) limitation of bacterioplankton and competition for nutrients (N and P) with phytoplankton in generally nutrient rich lake. Thymidine and leucine incorporation; leucine aminopeptidase, -D-glucosidase and alkaline phosphatase activity, total count of bacteria, chlorophyll a concentration and primary production as well as the concentrations of different chemical forms of N and P were followed during 4–5 days of the experiment. To address the question of the interactions between nutrients, bacterio- and phytoplankton, experimental and seasonal data sets were included in the analyses. Phosphorus (P) had a positive effect on bacterioplankton in enclosure experiments in June 1997; no effects of nutrients were found in September 1996, while in May 1996, P affected mainly the phytoplankton. On the seasonal scale, the development of bacterioplankton was connected to primary production, total phosphorus and temperature. In enrichment experiments, bacterioplankton was mainly related with primary productivity but the possible importance of bacterial grazers could be presumed. Thus, no evidence was found for nutrient growth limitation and/or competition for N and/or P, rather bacterioplankton depended on organic food supply originating from phytoplankton.     

滇中亚高山森林植物叶-凋落叶-土壤生态化学计量特征

为了深入认识滇中亚高山区域5种典型森林生态系统养分循环规律和系统稳定机质,通过测定植物叶、凋落叶和土壤C、N、P含量,掌握该区域典型森林植物叶-凋落叶-土壤生态化学计量特征。结果显示：（1）5种森林类型C、N、P含量差异显著,其中各林型植物叶和凋落叶C、N、P含量均高于土壤。（2）5种森林类型植物N、P再吸收率均为华山松林 > 云南松林 > 常绿阔叶林 > 高山栎林 > 滇油杉林;5种森林类型的再吸收率均为P（均值为61.20%）高于N（均值为36.48%）,表明了该区域土壤P的相对匮乏。（3）5种森林类型C/N、C/P、N/P均表现为凋落叶 > 植物叶 > 土壤;各林型植物叶N/P范围为10.17-15.31。（4）5种森林类型植物叶与凋落叶C、N、P含量、C/N和C/P均呈极显著或显著正相关;凋落叶与土壤的C、N含量及N/P呈显著正相关;5种森林类型植物叶N与P含量呈显著正相关关系;土壤N与P含量呈显著负相关关系。本文探究养分元素在"植物叶-凋落叶-土壤"之间的化学计量特征,为了解该区域森林生态系统的养分状况和揭示生物地球化学循环过程提供了理论数据。     

Genetically-based trade-offs in response to stoichiometric food quality influence competition in a keystone aquatic herbivore

The genetic basis of organism response to stoichiometric mismatches between environmental availability and somatic demand is still poorly understood. This study reports a consistent genotype × environment interaction related to phosphorus : carbon availability to Daphnia . In multiple pairs of Daphnia pulicaria clones, genetic variation at the phosphoglucose isomerase ( Pgi ) locus indicated that Pgi- heterozygotes out competed Pgi -homozygotes under high P : C conditions, whereas the opposite outcome was observed under low P : C conditions. Estimates of phosphorus use efficiency indicated that homozygotes were significantly more efficient. However, homozygotes were comparatively less homeostatic. We hypothesize that lower specific activity of Pgi from homozygotes, which results in lowered energetic efficiency during the second glycolytic step, may underlie the competitive advantage enjoyed by homozygotes under low P : C (i.e. excess C) conditions. Our results show that analysing stoichiometric mismatches between diet and consumer should advance our quest for a fundamental understanding of the mechanisms driving genotype-environment interactions.     

内蒙古典型草原生态系统中N素添加对羊草和黄囊苔草N:P化学计量学特征的影响(英文)

过度放牧是中国典型草原生态系统日趋退化的主要原因。由于养分输入-输出的不均衡,将导致土壤养分库的耗竭,而确定退化草原受何种养分的制约是对其进行恢复的重要途径之一。应用NP化学计量学的原理和方法,有望代替传统的野外养分添加实验,来研究不同草地受养分限制的状况。本文采用这两种方法在物种水平上研究限制性养分。此外,陆地植物器官中的NP比相对恒定是植物在地球上生存的重要适应机制,养分添加为验证这一假说提供了一种有效手段。为此,我们采用野外N素添加的方法,研究了内蒙古典型草原两种演替系列样地中(围封22年的样地A和围封2年的样地B)羊草(Leymuschinensis(Trin.)Tzvel.)和黄囊苔草(CarexkorshinskyiKom)生物量和NP化学计量学特征的变异。N素添加梯度分别为0、5、15、30、50、80gNH4NO3.m-2.a-1。研究结果表明,在施肥第一年,两个物种的地上生物量和P含量均不受N素添加的影响;相关分析结果表明,在施肥第二年两种植物的NP比不受氮素添加的影响;施肥可以显著提高羊草和黄囊苔草地上器官的含N量,P含量只是在第二年有显著增大的趋势;2001年,两块样地中羊草和黄囊苔草的氮磷含量在不同施肥处理下均呈极显著地正相关。这表明,样地A中黄囊苔草缺乏P,样地B中羊草缺乏N,施肥两年后,两个物种器官中的N     

内蒙古典型草原生态系统中N素添加对羊草和黄囊苔草N:P化学计量学特征的影响

过度放牧是中国典型草原生态系统日趋退化的主要原因.由于养分输入-输出的不均衡,将导致土壤养分库的耗竭,而确定退化草原受何种养分的制约是对其进行恢复的重要途径之一.应用N:P化学计量学的原理和方法,有望代替传统的野外养分添加实验,来研究不同草地受养分限制的状况.本文采用这两种方法在物种水平上研究限制性养分.此外,陆地植物器官中的N:P比相对恒定是植物在地球上生存的重要适应机制,养分添加为验证这一假说提供了一种有效手段.为此,我们采用野外N素添加的方法,研究了内蒙古典型草原两种演替系列样地中(围封22年的样地A和围封2年的样地B)羊草(Leymus chinensis(Ttin.)Tzvel.)和黄囊苔草(Carex korshinskyi Kom)生物量和N:P化学计量学特征的变异.N素添加梯度分别为0、5、15、30、50、80 gNH4NO3·m-2·a-1.研究结果表明,在施肥第一年,两个物种的地上生物量和P含量均不受N素添加的影响;相关分析结果表明,在施肥第二年两种植物的N:P比不受氮素添加的影响;施肥可以显著提高羊草和黄囊苔草地上器官的含N量,P含量只是在第二年有显著增大的趋势;2001年,两块样地中羊草和黄囊苔草的氮磷含量在不同施肥处理下均呈极显著地正相关.这表明,样地A中黄囊苔草缺乏P,样地B中羊草缺乏N,施肥两年后,两个物种器官中的N、P含量具有显著的协同关系,从物种水平上验证了我们提出的假说.同时,两年的实验结果还表明,生态系统中不同物种对添加N素的响应不同,笼统地界定一个生态系统受某种元素的制约是不恰当的.     

Generation time of Acanthocyclops robustus in relation to food availability and temperature in a shallow eutrophic lake

The generation time of the predatory cyclopoid copepod Acanthocyclops robustus was estimated on 11 occasions during the years 1980 to 1982 in Alderfen Broad. In a multiple regression model, generation time was found to be uncorrelated with temperature, positively correlated (p < 0.05) with the densities of Bosmina longirostris and rotifers, and negatively correlated (p < 0.001) with the density of nauplii of the calanoid copepod Eudiaptomus gracilis. It is suggested that generation time was determined largely by the availability of calanoid nauplii as prey, even though these constituted only 2% of zooplankton standing biomass.     

Responses of benthic algae to nutrient enrichment in a shallow lake: Linking community production,biomass, and composition

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