Succession of phytoplankton in a deep stratifying lake: Mondsee,Austria

Phytoplankton numbers, biovolume, chlorophyll-a and various physico-chemical characteristics were followed at weekly intervals in Mondsee, Austria during the year 1982. Secchi-disk transparency varied from 10 m in winter to 2 m in September. Prior to the onset of stratification phosphate-phosphorus concentration was 4 µg 1^–1 decreasing to undetectable values thereafter. Nitrate-nitrogen dropped from 590 µg 1^–1 to about 100 µg 1^–1 during the same time. The vernal bloom was dominated by Asterionella formosa Hass. which abruptly declined after silicon depletion. Spring growth ceased in early June, when Tabellaria flocculosa (Lyngb.) Kütz var. asterionelloides Grun. dominated. Oscillatoria rubescens D.C. and Microcystis aeruginosa Kütz. dominated summer and early autumn followed by the chrysophyte Dinobryon divergens Imh. and D. sociale Ehr. which formed up to 69% of total biovolume in October. Thereafter diatoms and Cryptophyceae (Rhodomonas lacustris Pascher and Ruttner, Cryptomonas pusilla Bach.) became abundant again.Maximum chlorophyll-a concentration in the epilimnion (16 µg 1^–1) was reached during spring growth of the diatoms. During summer higher chlorophyll-a levels were always associated with the metalimnetic layer of Oscillatoria.Compared with earlier studies, both the total biovolume and the share of Oscillatoria rubescens significantly decreased because of reduced nutrient loading of the lake and wash-out of Oscillatoria (theor. renewal time of the lake: 1.7 years).     

Seasonal subsidy stabilizes food web dynamics: Balance in a heterogeneous landscape

Resource subsidies from external habitats can substantially affect the food web dynamics of local habitats. In this paper, we explore a mathematical model that is tailored for a stream food web, studied by Nakano and colleagues, in which consumers, in situ prey and subsidies all show seasonal fluctuation. The model reveals that the food web dynamics are stabilized if subsidies increase in summer when in situ productivity is low. Consumer dynamics are stabilized because subsidies complement seasonal resource deficiency. In situ prey dynamics are stabilized because subsidies indirectly balance the predation pressure by consumers, with seasonal change in prey carrying capacity. In summer when prey carrying capacity is low, seasonally abundant subsidies indirectly decrease predation pressure, whereas in winter, with high prey carrying capacity, scarce subsidies increase the predation pressure. Our results suggest that temporal productivity differences between spatially linked habitats are important to promote the stability of food web dynamics in a landscape context.     

A comparison of phytoplankton size-fractions in Mondsee,an alpine lake in Austria: distribution,pigment composition and primary production rates

Production rates, abundance, chlorophyll a (Chl a) concentrations and pigment composition were measured for three size classes (<2 μm, 2–11 μm and >11 μm) of phytoplankton from May to December 2000 in deep, mesotrophic, alpine lake Mondsee in Austria. The study focuses on differences among phytoplankton size fractions characterised by their surface area to volume ratio ([mm² l⁻¹: mm³l⁻¹]), pigment distribution patterns and photosynthetic rates. Particular attention was paid to autotrophic picophytoplankton (APP, fraction <2 μm) since this size fraction differed significantly from the two larger size fractions. Among the three fractions, APP showed the highest surface area to volume ratios and a high persistence in the pattern of lipophilic pigments between temporarily and spatially successive samples (about 80% similarity of pigment composition between samples over seasons and depths). The epilimnetic abundance of APP varied seasonally with an annual maximum of 180 × 10³ cells ml⁻¹ in June (at 4–9 m). The minimum (October at 12 m) was more than an order of magnitude lower (4.9 × 10³ ml⁻¹). APP peaked during autumn and contributed between 24% and 42% to the total area-integrated Chl a (10–23 mg m⁻²) and between 16% and 58% to total area-integrated production (5–64 mg m⁻²  h⁻¹) throughout seasons.     

Some aspects of the seasonal distribution of flagellates in mountain lakes

In a larger regional survey in Tyrol, phytoplankton species composition and biovolume of mid-altitude and high-mountain lakes was studied. Results from eight lakes showed that flagellates (mainly Chrysophyceae, Dinophyceae, and Cryptophyceae) are important components of the phytoplankton.In the mid-altitude lakes a spring and an autumn maximum of Chrysophyceae as well as a summer maximum of large dinoflagellates are observed, whereas Cryptophyceae and Dinophyceae show irregular distributions. In the high-mountain lakes the seasonal variations of phytoplankton, including flagellates, are limited by the long duration of the winter situation. However similar sequences of phytoplankton assemblages as in the midaltitude lakes can be observed. Flagellates in high-mountain lakes are important to sustain phytoplankton standing crop under the winter snow and ice cover.In order to show similarities and differences of high-mountain and mid-altitude lakes, vertical profiles of phytoplankton from three lakes and seasonal patterns of Gymnodinium uberrimum from two lakes are compared. In addition the patterns of cryptomonads differing in their ecological requirements (Cryptomonas spp. and Rhodomonas minuta) are shown for a meromictic mid-altitude lake.     

Seasonal variations in the condition of two Daphnia species and their hybrid in a eutrophic lake: evidence for food limitation

As seston and chlorophyll concentrations in eutrophic lakesare usually high all year round, it was hypothesized in thepast that food limitation is of major importance for the seasonaldynamics of herbivorous zooplankton in such lakes. Since directmeasurements of food are hampered by the lack of knowledge onthe exact nature of the food in eutrophic conditions, indirectmeasurements are necessary to estimate the degree of food limitationin these circumstances. Hence, we used laboratory- and field-derivedrelationships between the body length and body carbon contentof different species of Daphnia, which were collected from thehighly eutrophic Tjeukemeer, the Netherlands. From a seasonalsurvey of the carbon content of the daphnids. we concluded thatin Tjeukemeer D.galeata, D.galeata x cucullata and D.cucullataare food limited during the largest part of the year. Sincethe condition of the hybrids was relatively high as comparedwith the parental species when the food concentration was high,D.galeata x cucutlaia is expected to be the more successfultaxon during periods of high food availability.     

食物网关键种的判定及其对稳健性的影响

食物网关键种的定量判定方法不仅有助于揭示食物网中的物种关系, 而且有利于对关键物种进行优先保护。本研究以包含49个物种的河流生态系统食物网为对象, 计算网络中表征度性质的指标(点度中心度、中介中心度、紧密中心度), 并根据指标对物种进行聚类, 分析不同类物种影响食物网连接稳健性程度的差异, 探讨不同节点去除方式下节点数量对食物网连接稳健性的影响。结果发现, 聚类将食物网中物种分成三类。I类物种具有相对较高的点度中心度、中介中心度、紧密中心度; II类物种具有相对更低的点度中心度、中介中心度、紧密中心度; III类物种的点度中心度、中介中心度、紧密中心度介于前两者之间。I类物种处于更为重要的地位, 可认为是关键种。本研究为如何判定关键种贡献了新的思路。另外, 升序去除方式下, 连接稳健性呈降低趋势; 降序去除与随机去除方式下, 连接稳健性在持续降低后又有小幅度增加, 这表明去除方式对食物网连接稳健性有影响。三种去除方式下, 食物网连接稳健性发生显著变化对应着几乎相同的去除节点数量, 表明关键种在维持食物网稳健性方面发挥着重要作用。     

Habitat distribution models,spatial autocorrelation,functional traits and dispersal capacity of alpine plant species

Abstract. We evaluate the potential influence of disturbance on the predictability of alpine plant species distribution from equilibrium‐based habitat distribution models. Firstly, abundance data of 71 plant species were correlated with a comprehensive set of environmental variables using ordinal regression models. Subsequently, the residual spatial autocorrelation (at distances of 40 to 320 m) in these models was explored. The additional amount of variance explained by spatial structuring was compared with a set of functional traits assumed to confer advantages in disturbed or undisturbed habitats. We found significant residual spatial autocorrelation in the habitat models of most of the species that were analysed. The amount of this autocorrelation was positively correlated with the dispersal capacity of the species, levelling off with increasing spatial scale. Both trends indicate that dispersal and colonization processes, whose frequency is enhanced by disturbance, influence the distribution of many alpine plant species. Since habitat distribution models commonly ignore such spatial processes they miss an important driver of local‐ to landscape‐scale plant distribution.     

Combining food web and species distribution models for improved community projections

The ability to model biodiversity patterns is of prime importance in this era of severe environmental crisis. Species assemblage along environmental gradients is subject to the interplay of biotic interactions in complement to abiotic filtering and stochastic forces. Accounting for complex biotic interactions for a wide array of species remains so far challenging. Here, we propose using food web models that can infer the potential interaction links between species as a constraint in species distribution models. Using a plant–herbivore (butterfly) interaction dataset, we demonstrate that this combined approach is able to improve species distribution and community forecasts. The trophic interaction network between butterfly larvae and host plant was phylogenetically structured and driven by host plant nitrogen content allowing forecasting the food web model to unknown interactions links. This combined approach is very useful in rendering models of more generalist species that have multiple potential interaction links, where gap in the literature may occur. Our combined approach points toward a promising direction for modeling the spatial variation in entire species interaction networks.     

Seasonal and spatial patterns of population density in the marine macroalga Mazzaella splendens (Gigartinales, Rhodophyta)

Insight into demographic processes that operate at larger spatial scales can be achieved through studying local populations when a particular species of interest is examined over time, by many investigators, in a variety of locations. On the west coast of North America, Mazzaella splendens (Setchell et Gardner) Fredericq is such a species of interest. A synthesis of local demographic studies of M. splendens from the late 1960s to the present reveals a pattern that is potentially common to the larger natural populations. This is the pattern: population density is high in summer and low in winter for both alternate free‐living life history phases of M. splendens. The magnitude of this seasonal change decreases in increasingly wave‐exposed habitats. In wave‐sheltered habitats there is a seasonal alternation from summer haploid to winter diploid dominance. This alternation gradually changes to constant diploid dominance as wave exposure in the habitat increases. Changes in population density are primarily a function of appearances and disappearances of perennating basal crusts (genets), as modules are produced or lost, rather than differential module production by genets of one phase over those of the other. To test the generality of this pattern, we examined seasonal changes in density, in local populations of M. splendens, in both a wave‐sheltered and a wave‐exposed habitat at Second Beach, Barkley Sound. Greater seasonal fluctuation in population density at wave‐sheltered, compared to wave‐exposed habitats is supported as a pattern potentially common to the natural populations of M. splendens. A change from summer haploid dominance in wave‐sheltered areas to summer diploid dominance in wave‐exposed areas is similarly supported. All changes in population density were the result of appearances and disappearances of genets rather than differential module production by haploid versus diploid basal crusts, also consistent with previous observations. A seasonal alternation in phase dominance, however, was absent from the wave‐sheltered site at Second Beach, Barkley Sound for 3 consecutive years. Seasonal alternation in phase dominance of M. splendens appears dependent on local conditions and is not common to all natural populations.     

Impact of internal waves on the spatial distribution of Planktothrix rubescens (cyanobacteria) in an alpine lake

The vertical and horizontal distribution of the cyanobacterium, Planktothrix rubescens, was studied in a deep alpine lake (Lac du Bourget) in a 2-year monitoring program with 11 sampling points, and a 24-h survey at one sampling station. This species is known to proliferate in the metalimnic layer of numerous deep mesotrophic lakes in temperate areas, and also to produce hepatotoxins. When looking at the distribution of P. rubescens at the scale of the entire lake, we found large variations (up to 10 m) in the depth of the biomass peak in the water column. These variations were closely correlated to isotherm displacements. We also found significant variations in the distribution of the cyanobacterial biomass in the northern and southern parts of the lake. We used a physical modeling approach to demonstrate that two internal wave modes can explain these variations. Internal waves are generated by wind events, but can still be detected several days after the end of these events. Finally, our 24-h survey at one sampling point demonstrated that the V1H1 sinusoidal motion could evolve into nonlinear fronts. All these findings show that internal waves have a major impact on the distribution of P. rubescens proliferating in the metalimnic layer of a deep lake, and that this process could influence the growth of this species by a direct impact on light availability.     

Carbon flow in the littoral food web of an oligotrophic lake

Benthic food web dynamics and carbon flow were examined in the littoral zone of Lake Coleridge, a large deep oligotrophic lake, using radioactive and stable isotope techniques in conjunction with analyses of stomach contents of the fauna. We specifically address two hypotheses: (1) that macrophytes only contribute to the carbon flow to higher trophic levels when they have decayed; and (2) that epiphytic algae is the major source of carbon for macroinvertebrates, and thus fish, with only minor contributions from phytoplankton or terrestrial sources. Epiphytic diatoms were a major component of the stomach contents of the gastropod snail Potamopyrgus antipodarum, and of chironomids. Animal remains were also common in the diet of some chironomids, while amorphous organic matter predominated in the stomachs of oligochaetes. A variety of epiphytic algal taxa was found in trichopteran larvae. Feeding rate of P. antipodarum measured with radioactive tracers increased by 10× on decayed macrophytes (Elodea) compared with live material, while feeding rates on characean algae increased by a factor of 3 when decayed material was presented. However, assimilation rates were less than 20% on decayed material compared with 48–52% on live material. Potential carbon sources were easily distinguished based on their ¹³C values, although isotopic ratios showed significant variation among sites. Epiphytic algae showed less variation among sites than macrophytes and were depleted by 4–5 compared with macrophytes. Detrital material, organic matter in the sediments and plankton were significantly depleted in ¹³C relative to macrophytes and slightly depleted relative to epiphytic algae. Most macroinvertebrate taxa showed a similar pattern among sites to macrophytes and epiphytic algae. P. antipodarum and chironomids were slightly enriched compared with epiphytic algae. Ratios for the common bully (Gobiomorphus cotidianus) were generally consistent with a diet dominated by chironomids, while there was some evidence for terrestrial inputs for koaro (Galaxias brevipinnis) and juvenile brown trout. Epiphytic algae appear to underpin much of the production in the littoral zone of this oligotrophic lake, with trichopteran and chironomid larvae mediating carbon flows from algae to fish. Macrophytes do not make a major contribution directly to carbon flow to higher trophic levels even when decayed. The lack of a direct link between macrophytes and higher trophic levels is due to the faunal composition, including a lack of large herbivores.     

Diet, growth and reproduction of the Arctic charr in a high alpine lake

The Arctic charr of Muzelle lake, at 2105 m in the Oisans massif (French Alps) feed most actively, essentially on Chironomidae, during the 6 or 7 weeks following the melting of the ice. The period of maximum growth corresponds to the time when food availability is at its highest, and it does not overlap with the period of maximum temperatures. Males and females mature at age 3 years but only half of the females of age 3 years spawn each year. This ratio of spawning females is constant from 3 to 6 years, the latter being maximum age observed in this lake. Each female spawns only twice at the most during its lifetime.     

Biogenic methane contributes to the food web of a large,shallow lake

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Seasonal variation in the composition and rate of change of the phytoplankton community in a deep subalpine lake (Lake Garda, Northern Italy). An application of nonmetric multidimensional scaling and cluster analysis

Seasonal variations and spatial homogeneity of the phytoplankton community were followed, from spring 1991 to spring 1992, in four pelagic stations of a large deep subalpine lake (Lake Garda, Northern Italy). Both cluster analysis and nonmetric multidimensional scaling (NMDS) applied to Bray & Curtis' dissimilarity matrices computed on density and biovolume data were employed; the Bray & Curtis index, calculated between pairs of chronologically contiguous samples, was also used as a measure of the community change rate over the temporal succession. In the tree diagrams obtained both from density and biovolume data, six groups of different size have been identified, with ordered sequences of samples within them. Superimposition of the results of cluster analysis on the NMDS configurations has allowed interpretations of the trajectories of the samples as a chronological and cyclical succession of compositionally homogeneous groups. A clear relationship between the community change rate and stability of the water column has been assessed. The specific composition of the six groups has been discussed in relation to environmental variables and in terms of adaptive strategies. During winter (winter-spring group) the turbulence of the water and the availability of nutrients favour the development of colonial Bacillariophyceae (Fragilaria crotonensis Kitton, Tabellaria fenestrata (Lyngb.) Kütz., Aulacoseira spp.), Cyanophyceae (Planktothrix agardhii (Gom.) Anagn. et Kom. and Planktolyngbya limnetica (Lemur.) Kom.-Legn. et Cronb.) and Cryptophyceae. In late spring group, with the stabilisation of the water column and silicon depletion, the diatoms give way to small, opportunistic species (Ankyra judayi (G. M. Smith) Fott, Cyclotella spp., Chroomonas acuta Utermhöl) and larger species (Ceratium hirundinella (O. F. Müll.) Dujardin). The three summer groups are characterised by a development of Chlorophyceae (chiefly Chlorococcales), Cyanophyceae (mainly Chroococcales), Cyclotella spp. and Dinophyceae. With the autumn destratification the summer community undergoes a rearrangement principally in favour of Cyanophyceae and Cryptophyceae.     

群落水平食物网能流季节演替特征

稳定碳、氮同位素比值分析技术是研究生态系统中物质循环与能量流动的有效技术.δ13C可以用来判断食物网中不同生物的能量来源;δ15N主要用于确定生物在食物网中所处的营养位置.通常用δ13C—δ15N图来表征某一特定时间或空间的食物网结构,但是这种方法在比较不同时间和空间食物网结构中功效较差.同时这种定性描述食物网结构也无法满足食物网复杂变化下的假说验证.应用环形统计方法,以太湖梅梁湾鱼类群落为例,定量评价了群落水平食物网能流季节演替特征.结果表明太湖梅梁湾鱼类营养生态位移动的季节特征明显.进一步物种水平分析结果表明,各种鱼类角度和幅度随季节均有变化.Rayleigh检验结果表明,群落中不同种类在秋冬、冬春和夏秋均有显著的一致的方向变化;而春夏期间不显著.Watson-William检验结果表明,群落水平的鱼类营养生态位移动在秋冬和冬春季节转换中有显著差异.引起鱼类群落水平营养生态位在食物网空间中季节性移动的主要因素为可利用资源稳定同位素的季节变化和不同鱼类种群自身的食性季节性转变.同时,由于梅梁湾食物网鱼类群落杂食性水平高,季节性浮游初级生产力成为食物网能量流动的重要驱动作用.因此,在富营养化生态系统中,食物网群落水平营养生态位季节波动也暗示了系统稳定性的下降.定量评价食物网变化有助于认识和理解食物网结构与功能在生态学和生态系统管理等方面的重要.     

Asymmetrical food web responses in trophic-level richness, biomass, and function following lake acidification

We tested for disproportional changes in annual and seasonal species richness and biomass among five trophic levels (phytoplankton, herbivorous, omnivorous, and carnivorous zooplankton, and fish) as well as altered trophic structure and ecosystem function following the 5-year experimental acidification of Little Rock Lake (Wisconsin, USA) from pH 6.1 to 4.7. Abiotic and biotic controls of trophic level response during acidification were also identified. Asymmetric reductions of species richness among trophic levels, separated by life stage and feeding type, were evident and changes in trophic structure were most pronounced by the end of the acidification period. Relative declines in richness of fish and zooplankton were greater than phytoplankton, which were generally unaffected, leading to a reduction of upper trophic level diversity. Each of the lower four trophic levels responded to a distinct combination of abiotic and biotic variables during acidification. pH was identified as a direct driver of change for only carnivorous zooplankton, while all other trophic levels were affected more by indirect interactions caused by acidification. Fluctuations in ecosystem function (zooplankton biomass and primary production) were also evident, with losses at all trophic levels only detected during the last year of acidification. The acidified basin displayed a tendency for greater variation in biomass for upper trophic levels relative to reference conditions implying greater unpredictability in ecosystem function. Together, these results suggest that trophic asymmetry may be an important and recurring feature of ecosystem response to anthropogenic stress.     

Seasonal and interspecific variation in the biochemical composition of some British fern species and their effects on Spodoptera littoralis larvae

Fern-feeding insects in Britain are mainly found on mature fronds in late summer. Six fern species (Dtyopteris filix-mas, D. dilatata, D. borreri, Phyllitis scolopendrium, Polyslkhum setiferum and Polypodium vulgare) were analysed for fibre, lignin, cellulose, tannins, cyanogenesis and thiaminase activity in an attempt to determine the biochemical basis for this seasonal pattern of attack. A bioassay was also carried out, using frond material incorporated in the diet of Spodoptera littoralis, to determine the effects of seasonal changes in fern toxins on a non-adapted, insect herbivore. The young fronds of all six species had a high protein content and low fibre-lignocellulose; protein levels decreased and cell wall materials increased as the fronds matured. Tannin concentrations and thiaminase activity showed a less consistent seasonal pattern. Tannins were present in the highest concentration in the young fronds of D. borreri, Polystichum and Polypodium and decreased with season; D. dilatata, Polystichum and Polypodium showed a similar pattern of thiaminase activity. The other species showed no strong seasonality in the levels of either tannins or thiaminase. No cyanogenic activity was detected in any of the fern species. Frond material from all six species decreased survivorship and growth rates of Spodoptera larvae. Larval performance and growth on diets containing fern material from young immature fronds was generally poorer than on diets containing material from mature fronds, collected later in the season. Neither tannin concentrations nor thiaminase activity levels closely corresponded to the observed interspecific and seasonal patterns of larval development and mortality, and other toxins must be involved. It is concluded that ferns are highly toxic to non-adapted herbivorous insects but the effects on adapted species are unknown. The restricted seasonal occurrence of a small number of insect species exhibiting specialized feeding habits suggests, however, that biochemical properties of the ferns determine this pattern of attack and limit the number of species which are able to exploit ferns as food resources.     

Effects of temperature and food supply on the growth of whitefish Coregonus lavaretus larvae in an oligotrophic peri‐alpine lake

The relative roles of temperature and food availability on the seasonal and daily growth of whitefish Coregonus lavaretus larvae were investigated in the oligotrophic peri‐alpine Lake Annecy, France. During the spring from 2004 to 2007, surface water temperature and density of potential zooplankton prey were monitored, and 2688 larvae were caught and measured. In addition, the daily growth of 130 larvae was estimated retrospectively by investigating the microstructure of their otoliths. Temperature played the predominant role in controlling both seasonal and daily growth of early larvae. In contrast, the abundance of Mesocyclops leuckarti and larval density was only slightly correlated to larval growth, suggesting no food limitation nor strong interindividual competition over the study period. Overall, these findings run counter to concerns about potential food limitation, but sound a warning about the potential impact of climate change on fish ecology and fisheries management.