Climate and productivity shape fish and invertebrate community structure in subarctic lakes

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Rapid changes in fish community structure and habitat distribution following the precipitation of lake phosphorus with aluminium

1. Fish community structure and habitat distribution of the abundant species roach, perch and ruffe were studied in Lake Nordborg (Denmark) before (August 2006) and after (August 2007) aluminium treatment to reduce internal phosphorus loading. 2. Rapid changes in fish community structure, abundance and habitat distribution occurred following a decline in in‐lake phosphorus concentrations from 280 to 37 μg P L^?1 and an increase in Secchi depth transparency from 1.1 to 1.9 m (August). The proportion of perch in overnight gill net catches increased, whilst roach decreased, and the average weight of all key species increased. 3. The habitat distribution of perch and roach changed from a high proportion in the upper pelagic and littoral zones in 2006, towards enhanced proportions in the deeper pelagic and profundal zone in 2007. The abundance of large‐bodied zooplankton increased and the abundance of benthic invertebrates decreased in the same period, suggesting that the habitat shift was not induced by food limitation. 4. Ruffe shifted from the littoral and upper profundal zones towards the deep profundal zone, likely reflecting an increased predation risk in the littoral zone and better oxygen conditions in the deep profundal. 5. Our results indicate that enhanced risk of predation in the upper pelagic and the littoral zones and perhaps improved oxygen concentrations in the deeper profundal zone at decreasing turbidity are responsible for the observed habitat shift. The results indicate that fish respond rapidly to changes in nutrient state, both in terms of community structure and habitat use.     

Invertebrate zooplankton predator composition and diversity in tropical lentic waters

Invertebrate zooplankton predators are generally less diverse in average species numbers in tropical than in temperate lakes and reservoirs. Predatory Copepoda which comprise the majority of limnetic predators are particularly low in species numbers in the tropics. Predatory Cladocera are confined to the North Temperate zone. Chaoborus appears to be cosmopolitan. Among Rotifera, only the cosmopolitan predator Asplanchna occurs in tropical waters while the other common limnetic carnivorous genus Ploesoma is restricted to higher latitudes. Hydracarina, and insects besides Chaoborus, are generally restricted to the littoral and appear to be more diverse in the tropics. Lakes Awasa and Zwai, Ethiopia, are almost devoid of predators in the limnetic, which is invaded by a littoral chydorid Alona diaphana. Low diversity of lake types and low production of tropical zooplankton could restrict predator diversity too. Very low diversity of invertebrate predators in the limnetic and higher diversity in the littoral may characterize tropical lakes in contrast to temperate lakes, which have more invertebrate predators in the limnetic and perhaps relatively less in the littoral. Tropical zooplankton in freshwaters, appears to be a very immature community. Hence opportunistic species can readily invade the limnetic and even dominate in isolated situations as has been shown for Alona davidi, Hydracarina and some other unconventional forms.     

Population structure and secondary productivity of Limnodrilus hoffmeisteri (Claparede) and Branchiura sowerbyi Beddard in the profundal zone of Lake Naivasha,Kenya

Lake Naivasha has been well studied since the 1930s but attempts to understand its ecological functioning have had to wait until enough was known about its structure. The energetics of the lake has only been studied to date at primary producer level. Following the identification of the invertebrate components of the littoral and profundal benthos, this study was initiated. The absence of native fish species in Lake Naivasha, combined with a fishery based only on three, introduced species, added an applied dimension to the work. The introduction of additional fish species which will utilize unexploited ecological niches has been suggested. The benthic invertebrates form one such niche. Two oligochaetes dominated the community, Branchiura sowerbyi Beddard and Limnodrilus hoffmeisteri Claparade. The former had a productivity of 7.4 g m^–2 ann^–1 (as dry weight), the latter 0.6. These figures are not particularly high and do not support the introduction of a new fish species on their own.     

Timing of the diet shift from zooplankton to macroinvertebrates and size at maturity determine whether normally piscivorous fish can persist in otherwise fishless lakes

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Depth gradients in food‐web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

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Empirical evaluation of phenotype–environment correlation and trait utility with allopatric and sympatric whitefish, Coregonus lavaretus (L.), populations in subarctic lakes

Adaptive phenotypic divergence of sympatric morphs in a single species may have significant evolutionary consequences. In the present study, phenotypic impacts of predator on zooplankton prey populations were compared in two northern Finnish lakes; one with an allopatric whitefish, Coregonus lavaretus (L.), population and the other with three sympatric whitefish populations. First, we examined whether there were phenotypic associations with specific niches in allopatric and sympatric whitefish. Second, trait utility (i.e. number of gillrakers) of allopatric and sympatric whitefish in utilizing a pelagic resource was explored by comparing predator avoidance of prey, prey size in environment, and prey size in predator diet. The allopatric living large sparsely rakered (LSR) whitefish morph, was a generalist using both pelagic and benthic niches. In contrast, sympatric living whitefish morphs were specialized: LSR whitefish was a littoral benthivore, small sparsely rakered whitefish was a profundal benthivore and densely rakered (DR) whitefish was a pelagic planktivore. In the lake with allopatric whitefish, zooplankton prey did not migrate vertically to avoid predation whereas, in the lake with sympatric whitefish, all important prey taxa migrated significantly. Trait utility was observed as significantly smaller size of prey in environment and predator diet in the lake with DR whitefish than in the lake with only LSR whitefish.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 561–572.     

The impact of predation by burbot (<Emphasis Type="Italic">Lota lota</Emphasis> L.) on the macroinvertebrate community in the littoral zone of a large lake

We studied the effects of predation by juvenile burbot (Lota lota) on the macroinvertebrate community in mesocosm experiments in the stony littoral zone of Lake Constance, a large prealpine lake in Central Europe. Although the growth data of the burbot suggest that the benthivorous fish exerts strong predation pressure on the invertebrate community, the predicted level of consumption is only poorly reflected by changes in biomass and abundance of most invertebrate prey taxa. High exchange rates of the prey between the mesocosms and the ambient littoral environment apparently masked the true predation effects of fish. Also, life-history events such as hatching or synchronised emergence of larvae led to temporal effects that obscured the impact of predation. However, for the dominant prey organism, the freshwater amphipod Gammarus roeseli, direct lethal effects appeared when its migration was limited. When exchange with the ambient littoral zone was possible, abundance and biomass of G. roeseli were unexpectedly high in the mesocosms stocked with burbot, indicating behavioural responses to fish presence. During the experiment, the burbot gained ca. 15% in length and about 60% in body mass. According to stomach content analyses at the end of the experiment, the ingested prey consisted mainly of relatively large and abundant invertebrates. Our study indicates that predation by juvenile burbot should be an important factor in structuring the benthic invertebrate community in terms of qualitative and quantitative effects.     

Adaptive Radiation along a Thermal Gradient: Preliminary Results of Habitat Use and Respiration Rate Divergence among Whitefish Morphs

Adaptive radiation is considered an important mechanism for the development of new species, but very little is known about the role of thermal adaptation during this process. Such adaptation should be especially important in poikilothermic animals that are often subjected to pronounced seasonal temperature variation that directly affects metabolic function. We conducted a preliminary study of individual lifetime thermal habitat use and respiration rates of four whitefish (Coregonus lavaretus (L.)) morphs (two pelagic, one littoral and one profundal) using stable carbon and oxygen isotope values of otolith carbonate. These morphs, two of which utilized pelagic habitats, one littoral and one profundal recently diverged via adaptive radiation to exploit different major niches in a deep and thermally stratified subarctic lake. We found evidence that the morphs used different thermal niches. The profundal morph had the most distinct thermal niche and consistently occupied the coldest thermal habitat of the lake, whereas differences were less pronounced among the shallow water pelagic and littoral morphs. Our results indicated ontogenetic shifts in thermal niches: juveniles of all whitefish morphs inhabited warmer ambient temperatures than adults. According to sampling of the otolith nucleus, hatching temperatures were higher for benthic compared to pelagic morphs. Estimated respiration rate was the lowest for benthivorous profundal morph, contrasting with the higher values estimated for the other morphs that inhabited shallower and warmer water. These preliminary results suggest that physiological adaptation to different thermal habitats shown by the sympatric morphs may play a significant role in maintaining or strengthening niche segregation and divergence in life-history traits, potentially contributing to reproductive isolation and incipient speciation.     

The Influence of Invertebrate Predators on Daphnia Spatial Distribution and Survival in Laboratory Experiments: Support for Daphnia Horizontal Migration in Shallow Lakes

The behavioural response of Daphnia cucullata to the presence of the pelagic invertebrate predator Leptodora kindtii, and the predation rate of littoral dragonfly nymphs on this species were investigated under laboratory conditions. Results of this study revealed a strong hiding response of Daphnia cucullata in the presence of the predatory cladoceran, L. kindtii, which was similar to the response of Daphnia in the presence of juvenile perch. This suggests that pelagic invertebrate predators may cause Daphnia to hide in the littoral zone which could result in increased exposure to predation by littoral invertebrates. A strong influence of dragonfly nymphs on D. cucullata, both in the presence and absence of macrophytes, was found. The average predation rate of Odonata larvae was about 5 prey ind^–1 h^–1 and did not differ significantly between treatments. Quantification of dragonfly pressure on Daphnia populations will require cross‐verification with field experiments since in the natural conditions Daphnia seeks a shelter in the vegetation stands against predation by Leptodora, despite the occurrence of odonates. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Discrete foraging niches promote ecological, phenotypic, and genetic divergence in sympatric whitefish (Coregonus lavaretus)

Natural populations often vary in their degree of ecological, morphological and genetic divergence. This variation can be arranged along an ecological speciation continuum of increasingly discrete variation, with high inter-individual variation at one end and well defined species in the other. In postglacial fishes, evolutionary divergence has commonly resulted in the co-occurrence of a pelagic and a benthic specialist. We studied three replicate lakes supporting sympatric pelagic and benthic European whitefish (Coregonus lavaretus (L.)) morphs in search for early signs of possible further divergence into more specialized niches. Using stomach content data (recent diet) and stable isotope analyses (time-integrated measure of trophic niche use), we observed a split in the trophic niche within the benthic whitefish morph, with individuals specializing on either littoral or profundal resources. This divergence in resource use was accompanied by small but significant differences in an adaptive morphological trait (gill raker number) and significant genetic differences between fish exploiting littoral and profundal habitats and foraging resources. The same pattern of parallel divergence was found in all three lakes, suggesting similar natural selection pressures driving and/or maintaining the divergence. The two levels of divergence (a clear and robust benthic – pelagic and a more subtle littoral – profundal divergence) observed in this study apparently represent different stages in the process of ecological speciation.     

Distribution and faunal associations of benthic invertebrates at Lake Turkana,Kenya

The benthic environment and fauna of Lake Turkana were studied during 1978–1979 to determine distribution patterns and associations of benthic invertebrates. Lake Turkana is a large, closed-basin, alkaline lake, located in northern Kenya.Detailed environmental information is currently only available for substrate variations throughout Lake Turkana. Water chemistry and other data are currently inadequate to evaluate their effects on the distribution of Lake Turkana benthic invertebrates. Three weak faunal-substrate associations were discovered at Turkana. A littoral, soft bottom association (large standing crop) is dominated by the corixid Micronecta sp. and the ostracod Hemicypris kliei. A littoral, rocky bottom association, also with a large standing crop, is dominated by various gastropods and insects. A profundal, muddy bottom association, with a very small standing crop, is dominated by the ostracods Hemicypris intermedia and Sclerocypris cf. clavularis and several gastropod and chironomid species.     

Dispersal and population expansion in a direct developing marine snail (Littorina saxatilis) following a severe population bottleneck

Most marine benthic invertebrate species have planktonic larvae, and in species in which juveniles and adults have low vagility a larva is obviously an efficient way of active dispersal. A minority of benthic invertebrate species develop without any pelagic phase at all. A largely unsolved question is how and at what rate do these species disperse. We have addressed this question using the marine littoral snail Littorina saxatilis (Olivi) as an example of a species that completely lacks larval dispersal. In the Koster archipelago (north part of the Swedish west coast), L. saxatilis occupies rocky island habitats of different sizes, from large islands to small intertidal skerries (islets). In 1988 an extremely dense bloom of a toxin-producing flagellate killed more than 99% of this snail species in this area. Populations of larger islands were reduced, often to less than 1%, but were restored over 2–4 yr. In contrast, populations of small intertidal skerries were completely wiped out and thus could not increase by local recruitment. Four years later, however, four of 33 skerries (12%) were successfully recolonized with relatively dense populations, and another five had received a few founder individuals. These results indicate recruitment through founder individuals, and are rough estimates of dispersal rate in a snail species that lacks a pelagic developmental stage.     

Context-dependent effects of freshwater mussels on stream benthic communities

1. We asked whether unionid mussels influence the distribution and abundance of co‐occurring benthic algae and invertebrates. In a yearlong field enclosure experiment in a south‐central U.S. river, we examined the effects of living mussels versus sham mussels (shells filled with sand) on periphyton and invertebrates in both the surrounding sediment and on mussel shells. We also examined differences between two common unionid species, Actinonaias ligamentina (Lamarck 1819) and Amblema plicata (Say 1817). 2. Organic matter concentrations and invertebrate densities in the sediment surrounding mussels were significantly higher in treatments with live mussels than treatments with sham mussels or sediment alone. Organic matter was significantly higher in the sediment surrounding Actinonaias than that surrounding Amblema. Actinonaias was more active than Amblema and may have increased benthic organic matter through bioturbation. 3. Living mussels increased the abundance of periphyton on shells and the abundance and richness of invertebrates on shells, whereas effects of sham mussels were similar to sediment alone. Differences in the amount of periphyton growing on the shells of the two mussel species reflected differences in mussel activity and shell morphology. 4. Differences between living and sham mussel treatments indicate that biological activities of mussels provide ecosystem services to the benthic community beyond the physical habitat provided by shells alone. In treatments containing live mussels we found significant correlations between organic matter and chlorophyll a concentrations in the sediment, organic matter concentrations and invertebrate abundance in the sediment and the amount of chlorophyll a on the sediment and invertebrate abundance. There were no significant correlations among these response variables in control treatments. Thus, in addition to providing biogenic structure as habitat, mussels likely facilitate benthic invertebrates by altering the availability of resources (algae and organic matter) through nutrient excretion and biodeposition. 5. Effects of mussels on sediment and shell periphyton concentrations, organic matter concentrations and invertebrate abundance, varied seasonally, and were strongest in late summer during periods of low water volume, low flow, and high water temperature. 6. Our study demonstrates that freshwater mussels can strongly influence the co‐occurring benthic community, but that effects of mussels are context‐dependent and may vary among species.     

Top-down and bottom-up impacts of juvenile fish in a littoral reed stand

1. Research has often focused on pelagic food chains and processes of lakes; less is known about the contribution of benthic energy flows to whole‐lake ecosystem energetics. This stems from the fact that the shoreline and littoral habitats, which provide a key linkage between sediment and water column, have only recently become a significant focus for study. 2. This study aimed to quantify the feeding and phosphorus allocation of a juvenile fish community in a littoral zone of a shallow lake in response to the biomass succession of the invertebrate prey community. Habitats comprising reed and adjacent open water were sampled over two consecutive years during day and night. 3. Although there were substantial year‐to‐year differences in the biomass of invertebrates, the fish community composition, diet consumption rates and phosphorus allocations were very similar in both study years. Biomasses and predation impacts by juvenile fish on prey groups were substantially higher within the reeds than in the adjacent open water habitat. This may be explained by the refuge‐seeking behaviour of the fish. 4. In general, invertebrates were negligibly influenced by fish feeding, with the exception for a strong top‐down control of large cladocerans. In response to the resulting low Daphnia biomass, fish were forced to switch to a higher degree of benthivory. Consequently, juvenile fish in littoral reed stands may shift benthic‐derived energy and phosphorus via the excretion of soluble reactive phosphorus into the open water.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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黄河三角洲芦苇湿地底栖无脊椎动物与环境因子的关系研究——以石油开采区与淡水补给区为例

分别于2018年5月和8月对黄河三角洲芦苇湿地19(淡水补给区11处,石油开采区8处)处采样点的底栖无脊椎动物和水体理化指标进行调查采样,运用统计方法分析两个区域物种组成、优势种、多样性、群落结构以及与环境因子的关系。结果表明:两季共采集到底栖无脊椎动物54种,主要以水生昆虫、腹足纲和软甲纲为主,淡水补给区和石油开采区各类群组成差异明显。独立样本T检验表明淡水补给区和石油开采区水体理化指标间差异显著(P0.05)。双因子方差分析显示,昆虫纲和腹足纲密度在两区域差异显著(P0.05),软甲纲和腹足纲密度在季节上差异明显(P0.05)。底栖无脊椎动物优势种共10种,淡水补给区指示物种8种,而石油开采区未发现有指示物种。聚类和非参数多维排序(nM DS)显示,底栖无脊椎动物群落结构相似性较低; RDA结果表明:淡水补给区底栖无脊椎动物群落结构主要受Cond,TDS,Sal,pH,Eh,HCO_3~-,SO_4~(2-)等环境因子的影响。石油开采区底栖无脊椎动物群落结构影响较大的环境因子为HCO_3~-、NH_4-N。     

Stable isotopes indicate that zebra mussels (Dreissena polymorpha) increase dependence of lake food webs on littoral energy sources

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The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

Acidification of a Small Watershed in Southern Norway and Some Characteristics of Acidic Aquatic Environments

Continuous monitoring of a small watershed in southern Norway shows that pH has decreased significantly by almost half a unit (from 5.9 to 5.4), conductivity increased from 23 to 29 μS/cm within the period from 1965–1975. Changes are also detected in sediment analyses where Daphnia and Chaoborus are only found deeper than 3–4 cm. Forest clear-cutting activities have probably accelerated the acidification processes of the already highly vulnerable freshwater localities. The acidic lakes show a solute content below 0.4 meq/1, and the bicarbonate buffer system may no longer work efficiently. Spring thaw and autumn rain episodes are accompanied by strong alternations in the chemical environment of the lakes; by considerable drop in pH and increasing conductivity and sulphate content. With increasing acidification the planktonic community changes from a fish predation governed system (brown trout and Eurasian perch as dominant species) to an invertebrate predation governed one with dominant predators Heterocope and pelagic corixids. Pelagic zooplankton are primarily composed of littoral species: Bosmina longispina, Diaphanosoma brachyurum, Eudiaptomus gracilis and Heterocope saliens. The littoral plant and invertebrate communities are described and discussed.