Experimental study of trophic cascade effect of silver carp (Hypophthalmichthys molitrixon) in a subtropical lake, Lake Donghu: on plankton community and underlying mechanisms of changes of crustacean community

An enclosure experiment was carried out to test trophic cascade effect of filter-feeding fish on the ecosystem: growth of crustacean zooplankton, and possible mechanism of changes of crustacean community structure. Four fish biomass levels were set as follows: 0, 116, 176 and 316 g m^-2, and lake water (containing ca. 190 g m^-2 of filter-feeding fishes) was comparatively monitored. Nutrient levels were high in all treatments during the experiment. Lowest algal biomass were measured in fishless treatment. Algal biomass decreased during days 21–56 as a function of fish biomass in treatments of low (LF), medium (MF) and high (HF) fish biomass. Crustaceans biomass decreased with increasing fish biomass. Small-bodied cladocerans, Moina micrura, Diaphanosoma brachyurum and Scapholeberis kingii survived when fish biomass was high whilst, large-bodied cladocerans Daphnia spp. and the cyclopoids Theromcyclops taihokuensis, T. brevifuratus, Mescyclops notius and Cyclops vicinus were abundant only in NF enclosures. Evasive calanoid Sinodiaptomus sarsi was significantly enhanced in LF, but decreased significantly with further increase of fish biomass. Demographic data indicated that M. micrura was well developed in all treatments. Our study indicates that algal biomass might be controlled by silver carp biomass in eutrophic environment. Changes of crustacean community are probably affected by the age of the first generation of species. Species with short generation time were dominant and species with long generation time survived less with high fish biomass. Evasive calanoids hardly developed in treatments with high fish biomass because of the (bottle neck) effect of nauplii. Species abundance were positively related to fish predation avoidance. Other than direct predation, zooplankton might also be suppressed by filter-feeding fish via competition.     

Long-term changes of planktonic rotifers in a subtropical Chinese lake dominated by filter-feeding fishes

1. The long-term changes (1956–1998) in density and species composition of planktonic rotifers were studied at two sampling stations (I, II) of Lake Donghu, a shallow eutrophic Chinese Lake densely stocked with filter-feeding fishes. Annual average densities of rotifers increased with an increase in fish yield and eutrophication, whilst species number decreased from 82 in 1962–1963 to 62 in 1994–1998.
2. During 1962–98, some species such as Anuraeopsis fissa , Polyarthra spp. (including P. dolichoptera & P. vulgaris ) , Trichocerca pusilla and Synchaeta oblonga increased their percentage in abundance remarkably, whilst the proportion of Keratella cochlearis decreased at two relatively eutrophic stations from 19 to 4.2% at Station I and from 30 to 3.2% at Station II.
3. The high r _max of A. fissa probably made it more successful than other rotifers under high predation pressure by planktivorous fish. The decrease in the K. cochlearis population might be attributed partly to predation by Cyclops vicinus .
4. Small rotifers were less vulnerable to fish predation than large-sized cladocerans. Decreases in cladocerans coincided with increases in rotifers, suggesting that the indirect effect of fish predation on cladocerans might have partly contributed to the population development of rotifers in Lake Donghu during recent decades.
5. We also conducted surveys (1994–1998) of seasonal dynamics of rotifers at four sampling stations (I–IV) which have varied in trophic status after fragmentation of the lake in the 1960s. A total of 75 species were identified at the four stations. Both densities and biomass of rotifers were considerably higher in the two more eutrophic stations than in the two less eutrophic stations. This indicates that the population increase of rotifers at Stations I and II during recent decades might be partly attributed to eutrophication of the lake water.     

Structure of the Phytoplankton Community and Its Relationship to Water Quality in Donghu Lake, Wuhan, China

The phytoplankton community structure, in terms of species composition, total standing crop,and abundance of the dominant algal species, at four stations in Donghu Lake, Wuhan, China, was investigated monthly from January 1994 to December 1996. A total of 260 taxa was observed, of which Chlorophyta (106 taxa) contributed the highest portion of the total number of taxa, followed by Bacillariophyta (82 taxa) and Cyanophyta (32 taxa). The total standing crop measured by means of chlorophyll a content, cell density,and cell biovolume, as well as the abundance of the dominant species, declined in the order of Station I to Station IV. Seasonal changes of the standing crop varied greatly among the four stations. Although the cell density at the four stations showed a single peak within a year, the peak density varied from July to November, dependent on the sampling year and the station. For chlorophyll a content and cell biovolume,multiple peaks were observed at Stations I and II, but a single peak was found at Stations III and IV. The phytoplankton community structure indicated that the trophic status was the highest at Station I (most eutrophic), followed by Station II; Stations III and IV were the least trophic areas. The long-term changes in phytoplankton community structure further suggested that changes in phytoplankton community structure were correlated with water quality, and eutrophication of Donghu Lake had been aggravated since the 1950s.     

Structure of the Phytoplankton Community and Its Relationship to Water Quality in Donghu Lake, Wuhan, China

The phytoplankton community structure, in terms of species composition, total standing crop,and abundance of the dominant algal species, at four stations in Donghu Lake, Wuhan, China, was investigated monthly from January 1994 to December 1996. A total of 260 taxa was observed, of which Chlorophyta (106 taxa) contributed the highest portion of the total number of taxa, followed by Bacillariophyta (82 taxa)and Cyanophyta (32 taxa). The total standing crop measured by means of chlorophyll a content, cell density,and cell biovolume, as well as the abundance of the dominant species, declined in the order of Station I to Station Ⅳ. Seasonal changes of the standing crop varied greatly among the four stations. Although the cell density at the four stations showed a single peak within a year, the peak density varied from July to November, dependent on the sampling year and the station. For chlorophyll a content and cell biovolume,multiple peaks were observed at Stations Ⅰ and Ⅱ, but a single peak was found at Stations Ⅲ and Ⅳ. The phytoplankton community structure indicated that the trophic status was the highest at Station Ⅰ (most eutrophic), followed by Station Ⅱ; Stations Ⅲ and Ⅳ were the least trophic areas. The long-term changes in phytoplankton community structure further suggested that changes in phytoplankton community structure were correlated with water quality, and eutrophication of Donghu Lake had been aggravated since the 1950s.     

A study of the zooplankton community of Billings Reservoir — São Paulo

The composition of the zooplankton of the Billings Reservoir and its variation in an eutrophic environment, subject to frequent blooms of algae (chiefly Cyanophyceae) was studied during one year (from October, 1977 to September, 1978) in two stations in the littoral and in the limnetic zone.The zooplankton community in the limnetic zone was dominated by cyclopoid copepods (Thermocyclops crassus and Metacyclops mendocinus) and by rotifers (Brachionus, Polyarthra and others) which represented, respectively 38.5 and 35.5% of the total zooplankton. At the littoral zone, cyclopoids were the most abundant (42.3%).The cladocerans were the least significant group at both stations, and calanoid copepods were found only at the littoral zone.A higher production of small filtrators, such as rotifers, cyclopoid nauplii and Bosmina sp was observed.     

用鲢鳙直接控制微囊藻水华的围隔试验和湖泊实践

为了控制水体富营养化所形成的水华,国外经典的生物操纵论者提倡放养食鱼性鱼类以控制食浮游生物的鱼类,藉此壮大浮游动物种群,然后借助浮游动物遏制藻类.按照这条思路,以浮游生物为食的鲢和鳙应该是清除的对象.本文作者认为我国湖泊中危害性最大的是微囊藻水华,而浮游动物根本不能摄食这种水华,不如直接利用鲢鳙进行控制,因而1989-1992年间在武汉的东湖进行了三次原位围隔试验,2000年再度重复了一次围隔试验,结果证明,迄今在东湖中已消失18年的微囊藻水华,重新出现在不养鱼的围隔里.在养入一定密度的鲢或/和鳙的围隔中,就没有水华出现.已经出现水华的围隔,放入适当密度的鲢或/和鳙后,在短期内水华又复消失;而放入草鱼的围隔,则水华始终持续,不受影响.说明鲢和鳙能有效地遏制微囊藻水华.鲢、鳙遏制水华的有效放养密度(亦即有效生物量)为46-50g·m^-3.东湖湖泊中鲢、鳙的生物量,直到1985年才达到这个水平,所以1985年湖里反常地没有出现水华.往后的年份,东湖的鱼产量越来越高,也就是湖里鲢、鳙的生物量越来越大,所以迄今没有微囊藻卷土重来的机会.东湖的水质和其他非生物条件仍适于蓝藻水华的发生与发展,如果东湖鲢、鳙的产量下降到阈值以下,亦即降到1985年以前1000t水平,那么微囊藻及其它蓝藻水华的重来将是不可避免的.经典的生物操纵论学说和措施不能解决微囊藻水华猖獗的问题.相反,用滤食浮游生物的鲢和鳙直接遏制微囊藻却是有实验根据并已被渔业生产实践证明行之有效的.     

Crustacean zooplankton assemblages in freshwaters of tropical Australia

Throughout tropical Australia eulimnetic zooplankton is composed of 7 calanoid copepods, 5 cyclopoid copepods and 12 cladocerans. A further 10 cyclopoids and 9 cladocerans occur as littoral strays, making 43 species in all. Dominants include Diaptomus lumholtzi, Mesocyclops notius, Thermocyclops decipiens, Diaphanosoma excisum, Ceriodaphnia cornuta, and Moina micrura. Momentary species composition averages 1.0 calanoids, 1.3 cyclopoids and 2.0 cladocerans. These assemblages are not only less complex than in temperate Australia, but also are not as rich as in most other tropical regions. The main contributors to these differences are calanoids and cladocerans respectively. Nevertheless, as elsewhere, most zooplankters in tropical Australia are eurytropic, widely distributed and small.     

武汉东湖后湖区浮游甲壳动物群落结构的研究

武汉东湖后湖湖区浮游甲壳动物群落结构从60年代至90年代发生了十分显的变化。浮游甲壳动物的物种丰度下降非常明显,枝角类的种类数从60年代的47种下降至90年代的17种,桡足类的种类数由60年代的14种下降为90年代的9种。60年代枝角类的优势种隆线Sou一亚种在90年代被短尾秀体Sou所取代,60年代的两种优势哲水蚤－长江新镖水蚤与特异荡镖水蚤的优势在90年代消失,而60年代很少的一种剑水蚤－台湾温剑水蚤在90年代跃升为优势种。随着滤食性鱼类密度的上升,枝角类的密度从60年代至90年代显下降。哲水蚤1997及1998年的密度明显低于60年代的密度。研究表明,浮游甲壳动物群落物种丰度的下降与后湖湖区沉水植被的退化有关,而优势种的演替主要受滤食性鱼类密度的影响。     

Zooplankton community structure driven by vertebrate and invertebrate predators

Summary A zooplankton community was established in outdoor experimental ponds, into which a vertebrate predator (topmouth gudgeon: Pseudorasbora parva) and/or an invertebrate predator (phantom midge larva: Chaoborus flavicans) were introduced and their predation effects on the zooplankton community structure were evaluated. In the ponds which had Chaoborus but not fish, small- and medium-sized cladocerans and calanoid copepods were eliminated while rotifers became abundant. A large-sized cladoceran Daphnia longispina, whose juveniles had high helmets and long tailspines as anti-predator devices, escaped from Chaoborus predation and increased. In the ponds which had fish but not Chaoborus, the large-sized Daphnia was selectively predated by the fish while small-and medium-sized cladocerans and calanoid copepods predominated. In the ponds containing both Chaoborus and fish, the fish reduced the late instar larvae (III and IV) of Chaoborus but increased the early instar larvae (I and II). Small- and large-sized cladocerans were scarcely found. The former might have been eliminated by predation of the early instar larvae of Chaoborus, while the latter was probably predated by fish. Consequently, the medium-sized cladocerans, which may have succeeded in escaping from both types of predator, appeared abundantly. The results suggest that various combinations of vertebrate and invertebrate predators are able to drive various kinds of zooplankton community structure.     

Size‐specific effects of bighead carp predation across the zooplankton size spectra

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Seasonal variations in stable isotope ratios of two biomanipulation fishes and seston in a large pen culture in hypereutrophic Meiliang Bay, Lake Taihu

This paper reports on seasonal changes in stable carbon and nitrogen isotope ratios of seston and muscle tissue of silver carp and bighead carp during 2004 and 2005, focusing primarily on the carbon sources and trophic relationships among phytoplankton, zooplankton and silver carp and bighead carp in a large fish pen of Meiliang Bay (Lake Taihu, China). δ¹³C showed a minimal value in March 2005 and a maximal value in August 2005 in seston both inside and outside the pen, whereas δ¹⁵N of seston showed the minimum in winter and the maximum during algal blooms. A positive correlation between δ¹³C of silver carp and that of seston suggested that temporal variation of δ¹³C in seston was preserved in fish via the food chain. The differences of δ¹³C among seston, zooplankton and muscle tissue of silver carp and bighead carp ranged only 0.2–1.7%, indicating that plankton production was the primary food source of filter-feeding fishes. According to a mass balance model, we estimated that the contributions of zooplankton to the diets of silver carp and bighead carp were 45.7% and 54.3%, respectively, based on the δ¹⁵N values of zooplankton and planktivorous fishes.     

Effects of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community of a deep,tropical reservoir: an enclosure experiment

1. An in situ enclosure experiment was conducted in a deep reservoir of southern China to examine (i) the effects of a low biomass (4 g wet weight m^?3) of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community and (ii) the response of Daphnia to eutrophication. 2. In the absence of fish, Daphnia galeata dominated the zooplankton community, whereas calanoids were dominant in the fish treatments, followed by D. galeata. Silver carp stocking significantly reduced total zooplankton biomass, and that of D. galeata and Leptodorarichardi, but markedly increased the biomass of smaller cladocerans, copepod nauplii and rotifers. In contrast, nutrient enrichment had no significant effect on the plankton community except for cyclopoids. 3. Chlorophyta, Cryptophyta and Bacillariophyta were dominant phytoplankton groups during the experiment. Chlorophyta with high growth rates (mainly Chlorella vulgaris in the fish enclosures and Ankyra sp. in the fishless enclosures) eventually dominated the phytoplankton community. Total phytoplankton biomass and the biomass of edible phytoplankton [greatest axial linear dimension (GALD) < 30 μm], Chlorophyta, Cryptophyta, Bacillariophyta and Cyanobacteria showed positive responses to fish stocking, while inedible phytoplankton (GALD ≥ 30 μm) was significantly reduced in the fish enclosures. However, there was no significant effect on the plankton community from the interaction of fish and nutrients. 4. Overall, the impact of fish on the plankton community was much greater than that of nutrients. High total phosphorus concentrations in the control treatment and relatively low temperatures may reduce the importance of nutrient enrichment. These results suggest it is not appropriate to use a low biomass of silver carp to control phytoplankton biomass in warmer, eutrophic fresh waters containing large herbivorous cladocerans.     

The effects of predation by juvenile fish on the meiobenthic community structure in a natural pond

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Intraspecific Autochthonous and Allochthonous Resource Use by Zooplankton in a Humic Lake during the Transitions between Winter,Summer and Fall

Seasonal patterns in assimilation of externally produced, allochthonous, organic matter into aquatic food webs are poorly understood, especially in brown-water lakes. We studied the allochthony (share biomass of terrestrial origin) in cladoceran, calanoid and cyclopoid micro-crustacean zooplankton from late winter to fall during two years in a small humic lake (Sweden). The use of allochthonous resources was important for sustaining a small population of calanoids in the water column during late winter. However, in summer the calanoids shifted to 100% herbivory, increasing their biomass several-fold by making efficient use of the pelagic primary production. In contrast, the cyclopoids and cladocerans remained at high levels of allochthony throughout the seasons, both groups showing the mean allochthony of 0.56 (range in mean 0.17-0.79 and 0.34-0.75, for the respective group, depending on model parameters). Our study shows that terrestrial organic matter can be an important resource for cyclopoids and cladocerans on an annual basis, forming a significant link between terrestrial organic matter and the higher trophic levels of the food web, but it can also be important for sustaining otherwise herbivorous calanoids during periods of low primary production in late winter.     

Feeding at different plankton densities alters invasive bighead carp (<Emphasis Type="Italic">Hypophthalmichthys nobilis</Emphasis>) growth and zooplankton species composition

Invasive Asian carps Hypophthalmichthys spp. are an ecological threat to non-native aquatic ecosystems throughout the world, and are poised to enter the Laurentian Great Lakes. Little is known about how these filter-feeding planktivores grow and impact zooplankton communities in mesotrophic to oligotrophic systems like the Great Lakes. Our purpose was to determine how different plankton densities affect bighead carp H. nobilis biomass and how bighead carp affect zooplankton species composition. We conducted a 37-day indoor mesocosm experiment (volume = 678 l) with high and low plankton treatments (zooplankton dry mass ≈ 1,900 and 700 μg l⁻¹; chlorophyll a = 25 and 14 μg l⁻¹, respectively) in the presence and absence of juvenile bighead carp (mean = 5.0 g, 8.5 cm). Carp lost weight in the low plankton treatment and gained weight in the high plankton treatment, suggesting that food availability may be a limiting factor to bighead carp growth in regions of low plankton densities. In the presence of carp, zooplankton shifted from Daphnia to copepod dominance, while in the absence of carp, Daphnia remained dominant. Chydorids and ostracods increased in the presence of carp, but only in the low plankton treatment, suggesting that the impact of bighead carp on zooplankton species composition may vary with zooplankton density. Chlorophyll was higher in the absence of carp than in the presence. Chlorophyll and zooplankton densities in many Great Lakes ecosystems are substantially lower than our low treatment conditions, and thus our results suggest that Asian carp establishment in these regions may be unlikely. Handling editor: S. Declerck     

滤食杂食性鱼类放养对浮游动物群落结构的影响

滤食杂食性鱼类是我国南方水体中常见鱼类,本文调查了放养滤食杂食性鱼类后浮游动物的丰度和种类组成的变化,以探讨其对浮游动物群落的影响.结果显示,滤食杂食性鱼类放养之后,枝角类模糊秀体溞(Diaphanosoma dubium)、微型裸腹溞(Moina micrura)和桡足类蒙古温剑水蚤(Thermocyclops mongolicus)、南方近镖水蚤(Tropodiaptomus australis)等丰度迅速下降,而轮虫丰度在鱼类放养一周后迅速增加,且保持较高的密度;轮虫优势种也发生了变化,调查前两周主要以臂尾轮虫(Brachionus)为主,两周后以暗小异尾轮虫(Trichocereidae pusilla)、广布多肢轮虫(Polyarthra trigla)和裂痕龟纹轮虫(Anuraeopsisfissa)等个体更小的种类为主.本研究说明,滤食杂食性鱼类能够显著影响浮游动物群落结构,个体较大的种类受到的影响较大,最后导致浮游动物群落的小型化.     

Spatial and seasonal variation of the zooplankton in the coastal zone and main khors of Lake Nasser (Egypt)

A prominent feature of Lake Nasser is the presence of khors (dendritic side extensions). We studied the zooplankton of the larger khors and coastal zone of the main lake in 1996 and 1997, and found an assemblage of rotifers, cladocerans and copepods that was partly tropical, partly temperate, at relatively high biomasses. Spatial differences were weak, but the upstream khors (Toushka and Korosko) were consistently richer than the downstream khors (El-Ramla and Kalabsha), with a rather sudden transition around km 150 at El Madiq. Summer standing crops were higher than those in winter by a factor 2–3. The zooplankton of the littoral of the main channel showed the same spatial pattern as that in the khors, being more abundant in spring (average 82300 ind m^–3) than in autumn (average 72700 ind m^–3). Zooplankton dry weight increased from 4 g m^–2 at khor El-Ramla to 7 g m^–2 at khor Toushka. These rather high values had low variation. The number of species, diversity and evenness all showed a high degree of similarity among the khors and in the littoral of the main lake. The lake fish fauna is poor, lacking a pelagic planktivore. The predominance of medium-sized Copepoda (one calanoid, two cyclopoids) in the zooplankton suggests that fish predation is moderate. This is confirmed by the persistence of two Daphnia species at low abundance, although rather strong variations in time suggest that Cladocera suffer from summer predation by invertebrates (copepods) as well as vertebrates (mostly larval fish). Because the zooplankton is underutilised by higher trophic levels, we suggest to assess the benefits of introducing a pelagic zooplanktivorous fish.     

Does stocking of filter-feeding fish for production have a cascading effect on zooplankton and ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient concentrations

Stocking of filter-feeding fish is a common tool used in Chinese reservoirs to increase fish production because of low natural recruitment. Whether such stocking has important negative effects on zooplankton with cascading effects on phytoplankton is debated. We compared the zooplankton communities in fourteen reservoirs with different nutrient concentrations and fish densities. Both chlorophyll a (Chla) and fish catch were positively related with total phosphorus (TP), whereas zooplankton biomass did not show a similar relationship with TP. Zooplankton seemed to be influenced by fish as high fish catches coincided with a low proportion of calanoids of the total copepod biomass, a high proportion of rotifers of the total zooplankton biomass, a low zooplankton:phytoplankton biomass ratio, and the absence of Daphnia irrespective of TP concentration. Both zooplankton biomass and most of the zooplankton:phytoplankton biomass ratios were among the lowest reported in the literature for the nutrient range studied. Furthermore, the Chla:TP ratio was higher than what is typically observed in temperate lakes. We conclude that top-down control of zooplankton is of key importance in reservoirs in South China where frequent stocking of filter-feeding fish seems to contribute to poor water quality in the form of higher algal biomass and reduced clarity.     

鲤鱼种和鲢鳙对池塘浮游生物的影响

本文根据１９９０年－９月对９个鱼池的研究,报道鲤鱼种和鲢鳙对浮游生物的影响,结果表明：鲤鱼种使浮游植物和浮游动物生物量增加,浮游植物大型化,浮游动物小型化,并使浮游植物多样性增加,浮游动物多样性下降;鲢鳙密度增加,浮生物小型明显,微型藻类和超微在在浮植物中所占比重显著增加,小型浮游动物（原生动物和轮虫）在浮游动物中比重也明显增;鲢鳙密度对浮游生物多样性和浮游生物量的影响具有阶段性,当鲢鳙密度低时,     

The impact of planktivorous flsh on the structure of a plankton community

SUMMARY. 1. The abundance of pianktivorous juvenile yellow perch, Perca flavescens , was manipulated in three 750 m³ enclosures in a eutrophic lake.
2. There was a significant negative relationship between fish and zoopiankton biomasses. At high fish densities the zooplankton community was dominated by small filter-feeding cladocera. primarily bosmi- nids. At low fish densities the zooplankton community was dominated by large filter-feeding cladocera, primarily daphnids.
3. There was no significant relationship between zooplankton and phytoplankton biomasses when considered over the whole experiment but there was a trend towards lower phytoplankton biomass in the enclosure dominated by daphnids during mid-summer.
4. We conclude that although planktivorous fish have a strong negative impact on zooplankton community biomass and size structure, the relationship at the next lower trophic level, zooplankton and phytoplankton, is much weaker. Therefore, the biomanipulation of planktivorous fish populations as a management technique to control phytoplankton abundance is largely ineffective.