Zooplankton,fish and fisheries in tropical freshwaters

About 40% of all fish species occur in freshwater, although only 1% of the globe is occupied by freshwaters. The tropics harbour a high percentage of these fishes. Freshwater zooplankton on the other hand is far less diverse than its marine counterpart and the tropics do not harbour a markedly high percentage of freshwater species either. The antecedents of freshwater zooplankton appear to have come from riverine and terrestrial habitats via temporary habitats (ponds, floodplains). The present zooplankton composition has also been shaped by, among other factors, the highly efficient zooplanktivorous modern teleosts which have restricted the formerly widespread Branchiopoda mainly to fishless freshwaters. Those Branchiopoda frequently co-existing with fishes (Cladocera) have their size composition strongly influenced by fish predation. Circumstantial evidence indicates that pelagic zooplankton (Cladocera, Copepoda, Rotifera) appear to provide a relatively scarce food supply relative to the littoral region for the early stages of fishes. Also, unpredictability of zooplankton may be involved here. These factors have led to the loss of planktonic eggs and the siting of fish breeding in shallow littoral situations, where other animal foods besides zooplankton are also available, especially for later stages of juvenile fish. The Ostariophysi breed in the shallow expanses of flood waters while the Cichlidae, some of which breed like Ostariophysi, also breed in standing waters in the littoral of lakes or floodplains. In all these locations zooplankton and benthic organisms, especially insects, are available. The cichlids are, in addition, provided with parental care. Predation on young fishes is also reduced by these strategies. Young fishes may also be adapted to feed on patches of zooplankton and other food in their breeding grounds. Tropical pelagic clupeids and cyprinids may breed continuously. Some of these clupeids in rivers breed at low water.Zooplankton, supplemented by other animal food is more critical to achieving the potential fish yields in the tropics than in temperate freshwaters because fish yields in the tropics can be very high indeed. The high metabolic rate of young fishes in the tropics adds to the demand for food. Tropical freshwaters have a relatively high primary production but a low zooplankton/phytoplankton ratio. Zooplankton is kept small in size and biomass by continuous predation. Herbivorous fishes can sustain very high fish yields in the tropics but they must have a high fecundity and high survival of young produced seasonally, mainly in rivers or even continuously as in lakes and reservoirs to weather predation. Rich littoral zooplankton and benthos combined with omnivory and a higher efficiency in the use of the available animal food by newly hatched fishes may be critical factors linking fish yields to zooplankton in tropical freshwaters. The ability of herbivorous tilapias to give very high fish yields in shallow tropical lakes may also be due to their efficient use of animal food, algae and microphagy in young stages besides other favourable adaptations like opportunistic feeding on detrities and the ability to feed on and digest cyanobacteria, abundant in the tropics.     

Seasonal variability of secondary production of cladocerans and rotifers,and their trophic role in Lake Tana,Ethiopia, a large,turbid, tropical highland lake

Daily and annual production rates of eight cladoceran and two rotifer species, and their seasonal variation and trophic role in the large, turbid, tropical Lake Tana, Ethiopia, were assessed in 2003–2005. Laboratory cultures were used to infer cladoceran development times, and secondary production was estimated using the growth increment summation and recruitment methods. Production for both taxa was highest in October–November, after the rainy season, and lowest in January–April during the dry season. Cladocerans and rotifers comprised 24% of the metazoan zooplankton biomass of 45.1 mg DW m^?3, but comprised 53% of its production. Daily production for cladocerans and rotifers, respectively, was 1.23 and 0.94 mg DW m^?3 d^?1, and annual production was 447.9 and 353.5 mg DW m^?3 y^?1. Energy transfer efficiency from producers to zooplankton was 1.3% and 4.4% from zooplankton to planktivores. Herbivores consumed 3.4% of primary production and planktivores 36% of zooplankton production. High biomass turnover rates of cladocerans and rotifers sustain planktivores and, after a month's delay, decomposed Microcystis provides their main food source during the pre- and post-rainy months in Lake Tana.     

The Freshwater Zooplankton of Sri Lanka,with a Discussion of Tropical Freshwater Zooplankton Composition

Based on the study of over 500 zooplankton samples collected in Sri Lanka (Ceylon) during 1965–1974, the species composition from different habitats is analysed. The zooplankton assemblage is typically tropical with relatively few species of Cladocera and Copepoda. The Rotifera include a large number of species of the genus Brachionus. The limnetic zooplankton resembles the pond zooplankton closely in that all the eurytopic species found in the different types of habitats, including ponds, also occur in the limnetic zooplankton. The large Cladocera belonging to the genus Daphnia are very rare. In general, large zooplankters are absent. The size composition of the zooplankton has a smaller range than in temperate regions. This is due to the absence of large-sized zooplankton species. The reasons for the differences in species variety and size composition between zooplankton of temperate and tropical regions is perhaps due to a number of factors. These include the effects of high and uniform temperatures, food availability and predation by fish and invertebrates.     

Estimating population birth rates of zooplankton when rates of egg deposition and hatching are periodic

Summary I present a general method of computing finite birth and death rates of natural zooplankton populations from changes in the age distribution of eggs and changes in population size. The method is applicable to cases in which eggs hatch periodically owing to variable rates of oviposition. When morphological criteria are used to determine the age distribution of eggs at the beginning and end of a sampling interval, egg mortality can be incorporated in estimates of population birth rate. I raised laboratory populations of Asplanchna priodonta, a common planktonic rotifer, in semicontinuous culture to evaluate my method of computing finite birth rate. The Asplanchna population became synchronized to a daily addition of food but grew by the same amount each day once steady state was achieved. The steady-state rate of growth, which can be computed from the volume-specific dilution rate of the culture, was consistent with the finite birth rate predicted from the population's egg ratio and egg age distribution.     

Ingested prey increase risks of visual predation in transparent Chaoborus larvae

Summary Transparency reduces the chances of detection of large planktonic animals by visual predators. An important constraint on the transparency of planktonic animals may be ingested food which could be seen through the body, thereby increasing the vulnerability of transparent zooplankton to visual predators. To test this hypothesis, we presented fed and un-fed Chaoborus larvae to juvenile coho salmon (Oncorhynchus kisutch). Overall, the presence of prey in the gut of Chaoborus increased their probability of capture by 68%. Predation risks due to the visibility of ingested food increased in proportion to meal size: larvae with nearly full gut were captured about three times faster on the average than larvae which had little food in their gut. Although Chaoborus larvae may be able to reduce this increased predation risk by migrating downward to low light levels, this behavior would reduce feeding opportunities by removing the larvae from surface waters where prey density is generally high. In this way, visual predators may limit the growth and the maximum size that can be achieved by transparent animals.     

Combined effects of environmental factors and predator–prey interactions on zooplankton assemblages in five high alpine lakes

The examination of a yearly cycle of plankton density (rotifer and microcrustacean) and of 419 stomach contents of four species of salmonid fishes, arctic charr, Salvelinus alpinus, brown trout, Salmo trutta, rainbow trout, Onchorynchus mykiss and lake trout, Salvelinus namaycush, living in five high altitude lakes in the French Alps, shows an impact of abiotic variates and the effects of predation on the composition of zooplankton assemblages. The lakes studied may be divided in two groups. The colder, Muzelle and Puy Vachier, are characterised by a low level of food resources, an important impact of predation, and the near absence of planktonic crustaceans. The second group, consisting of Lakes Les Pisses, Petarel and Palluel, is characterised by a low density rotifer assemblage and a more abundant crustacean population controlled by a low level of fish predation.     

Size‐based interactions and trophic transfer efficiency are modified by fish predation and cyanobacteria blooms in Lake Mývatn,Iceland

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Life history strategies of cladocerans: comparisons of tropical and temperate taxa

We review recent works on different life history variables of cladoceran taxa in tropical and temperate freshwater bodies, comparing the strategies that cladocerans have evolved to adapt to contrasting environmental conditions in the two geographical regions. These life-history parameters relate to age and size at maturity, survival, fecundity, life-expectancy at birth, lifespan, gross, and net reproductive rates, generation time, the rate of population increase, peak population density and day of peak abundance. We also discuss the role of photoperiod and temperature on some of these life history parameters. We found a general paucity of experimental work and field data in tropics on cladocerans. There is very limited information on the few Daphnia species found in the tropics. The misconception of low species diversity of cladocerans in the tropics arose due to several reasons including lack of extensive and intensive field collections. Higher water temperatures apparently promote permanent infestation of tropical waters with toxic cyanobacteria, which reduce the zooplankton diversity. In addition to higher temperatures in the tropics, the year-round high predation pressure of planktivorous fish probably causes the tropical species, particularly in pelagic habitats, to reach maturity earlier (< 3 days) than in temperate regions. Species of Daphnia in temperate regions are particularly adapted to living at food concentrations that are much lower and seasonably more variable than those for tropical genera such as Diaphanosoma. This is further corroborated by the more than an order of magnitude higher threshold food concentration (TFC) for tropical Cladocera than for their temperate counterparts. Fecundity patterns differ between tropical and temperate cladoceran taxa: cultured under optimal temperature regimes, tropical taxa have fewer eggs than temperate species of a comparable body size. Predation pressure may act differently depending on the size of the cladoceran neonates and thus on their population size structure. Global warming and climate changes seem to affect the behaviour (migration), distribution, and abundance of cladocerans. Apparently, in direct response to these changes, the possibility of encountering the tropical cladocerans in the northern, temperate hemisphere (bioinvasions) is on the rise.     

Feeding while evading predators by a lotic mayfly: linking short-term foraging behaviours to long-term fitness consequences

Laboratory experiments were performed to quantify the combined effects of food abundance (low, high) and predation threat, imposed using a model fish (safe, risky) on fitness correlates (i.e. growth, time to emergence, adult body mass, fecundity, egg size) of the mayfly Baetis tricaudatus. These effects were determined by rearing larvae under different combinations of food abundance and predator threat. Fitness correlates were significantly affected by food abundance, predation threat or the interaction of these factors. High food abundance and the absence of predation threat significantly increased larval growth rates, adult body mass, fecundity, egg size, and decreased time to emergence. Long-term effects of predator threat and food abundance on fitness correlates of B. tricaudatus were compared to previously conducted short-term patch choice trials to test for concordance between short-term measures of patch choice and their potential longer-term fitness consequences. This comparison indicated that patches that were utilized the most were those that yielded the highest fitness benefits. These results suggest that behavioural costs of balancing mortality risk due to predation against food acquisition by B. tricaudatus can have strong fitness consequences.     

Longitudinal changes in zooplankton distribution below a reservoir outfall with reference to river planktivory

The fate and interactions with river organisms of zooplankton as they drift downriver from a reservoir on a fourth-order mountain stream (Hiji River, Japan) were investigated. Monthly samples were collected at the reservoir and six river sites, simultaneously, from May 2005 to May 2006. Aquatic macroinvertebrates and fish were colleted, and their stomach contents were analyzed in April and May, 2006, respectively. Drift from the reservoir was the primary source for the river plankton community; the abundance of zooplankton, particularly those of cladocerans and large rotifer, rapidly decreased within several kilometers of the dam. Analysis of the contents of fish stomachs showed that drifting zooplankton was the main food for fish, with strong food selectivity for cladocerans and large rotifers. However, fish and insect planktivores showed longitudinally different stomach contents, with progressively fewer zooplankton found in the stomachs at the downriver sites. The results suggest that the outflow of zooplankton from the reservoir is an important food source for the downstream predators, especially fish, but the drift of zooplankton and consequent food availability for the predators at lower sites are strongly limited by concentrated fish predation just below the reservoir dam.     

Erratum to: Effects of fish farming on plankton structure in a Brazilian tropical reservoir

This study investigated the effects of fish farming on phytoplankton and zooplankton structure in a Brazilian tropical reservoir. Samplings were undertaken among 15 net cages and upstream and downstream of the cages over a 120-day period. Soon after the tilapia feeding, we observed the highest density of phytoplankton, with dominance of Cyanobacteria which probably increased due to the increase in predation pressure by cladocerans and copepods on the Cryptophyceae. However, after 30 days from that, a reduction in zooplankton density was registered due to the dominance of Cyanobacteria. The absence of a clear spatial difference in the phytoplankton and zooplankton densities was observed. The effects of the net cages on the nutrients and planktonic communities were small, probably due to the low number of net cages and fish employed, the seasons (autumn/winter) and the hydrodynamics, as lotic traits, the strong influence of the wind, and the large extension of the mixture zone.     

Relative significance of direct and indirect effects of predation by planktivorous fish on zooplankton

One of the most obvious features of tropical lakes and reservoirs is the small body size of their zooplankton taxa. It is believed that this is the result of high and persistent predation by abundant planktivorous fish, which select large-bodied zooplankton prey thus making them more vulnerable to extinction in tropical as compared to temperate habitats. Do these extinctions result directly from fish predation? Could the high predation-induced mortality alone be responsible for an extermination of the population from a habitat? Or could indirect effects of predation be responsible? Some important indirect effects can be seen at the demographic level; these include reduced reproduction in the population resulting from higher vulnerability of ovigerous females to predation by visually oriented planktivores. Other important indirect effects can be observed at the individual level; these include shifts in behavior (from foraging to predator avoidance) and adjustments in physiology (from high to low feeding rate) in those planktonic animals which detect danger from their predators by sensing either the ‘predator odor’ or an ‘alarm substance’ originating from injured conspecific prey. Although a zooplankton species density may mostly result from the brutal force of direct predator impact on the population (mortality), it is more likely that its distribution in time and space could be attributed to a combination of indirect effects of predation on individual behavior and physiology. An example of periodicity in density and depth distribution patterns of Cahora Bassa zooplankton species and their periodic exterminations seems to confirm the role of indirect effects of predation by planktivorous fish.     

Seasonal succession of zooplankton in the north basin of Lake Biwa

To examine the seasonal succession of the entire zooplankton community in Lake Biwa, zooplankton biomass (on an areal basis) and its distribution patterns among crustaceans, rotifers and ciliates were studied in the north basin from April 1997 to June 1998. Seasonal changes in phytoplankton and population dynamics of Daphnia galeata were also examined to assess food condition and predation pressure by fish. From March to November, crustaceans dominated zooplankton biomass, but rotifers and ciliates were dominant from December to February. Among crustaceans, Eodiaptomus japonicus was the most abundant species, followed by D. galeata. Zooplankton biomass increased from January to a peak in early April, just before the spring bloom of phytoplankton, then decreased in mid-April when mortality rate of D. galeata increased. From mid-June, zooplankton increased and maintained a high level until the beginning of November. During this period, both birth and mortality rates of D. galeata were relatively high and a number of rotifer and crustacean species were observed. However, their abundances were very limited except for E. japonicus which likely preys on ciliates and rotifers. In Lake Biwa, food sources other than phytoplankton, such as resuspended organic matter from the sediments, seems to play a crucial role in zooplankton succession from winter to early spring, while zooplankton community seems to be regulated mainly by fish predation from summer to fall.     

Dynamics of Daphnia hyalina × galeata in Castelo-do-Bode Reservoir: The effect of food availability and flatworm predation

Daphnia hyalina × galeata in Castelo-do-Bode Reservoir was regulated mainly by food availability during the summers of 1993 and 1994. Because of high temperatures and low rainfall in 1994, the reservoir was, apparently, more eutrophic relatively to 1993. Higher chlorophyll-a concentrations and higher clutch size in 1994 suggest that Daphnia population benefited from better feeding conditions in 1994. The analysis of the contributions from fecundity and adult proportion to birth rate variation suggest that vertebrate predation and predation by copepods were negligible factors as population organisers. A general decrease in the abundance of large crustaceans during the periods of high density of typhloplanid flatworms, in both years, points to the importance of flatworm predation on zooplankton. A decrease in the number of individuals in all Daphnia size classes was observed during the periods of flatworms population increase.     

Plankton composition, abundance and dynamics in a severely stressed cooling reservoir

The effects of severe thermal stress imposed by the intermittentoperation of a nuclear reactor on plankton abundance and dynamicswere investigated in Pond C, a cooling reservoir on the SavannahRiver Site in South Carolina, USA. Temperatures in Pond C rangedup to 58°C during reactor operation. The thermal effluenteliminated zooplankton from regions where the temperature exceeded45°C, reduced zooplankton abundance by 1–3 ordersof magnitude and typically halved the number of taxa. Reactoroperation also reduced phytoplankton biovolume, often by >70%.During intermittent reactor operation, the rotifer Filinia longisetadominated the zooplankton and two cladocerans of the genus Moinawere abundant. These species were not abundant during extendedreactor shutdowns. The success of Filinia and Moina was dueprimarily to their tolerance of high temperatures. Sparse phytoplanktonprobably limited some zooplankton taxa, although other taxa,such as Filinia may have utilized bacterial resources. Reactoroperation may have intensified predation on crustacean zooplanktonwhen fish were concentrated in refuge areas with zooplankton.Processes by which zooplankton repopulated the reservoir afterreactor shutdown were inferred from zooplankton distributionpatterns, and population growth and birth rates. Repopulationtypically occurred within a few days due to rapid growth ofpopulations from refuge areas within the reservoir and colonistsbrought in through a tributary canal. Mechanisms of zooplanktonrepopulation in Pond C suggest that refuges or colonizationcorridors should be maintained when the re-establishment ofcommunities following cessation of stresses is desired.     

Linking herbivore-induced defences to population dynamics

1. Theoretical studies have shown that inducible defences have the potential to affect population stability and persistence in bi‐ and tritrophic food chains. Experimental studies on such effects of prey defence strategies on the dynamics of predator–prey systems are still rare. We performed replicated population dynamics experiments using the herbivorous rotifer Brachionus calyciflorus and four strains of closely related algae that show different defence responses to this herbivore. 2. We observed herbivore populations to fluctuate at a higher frequency when feeding on small undefended algae. During these fluctuations minimum rotifer densities remained sufficiently high to ensure population persistence in all the replicates. The initial growth of rotifer populations in this treatment coincided with a sharp drop in algal density. Such a suppression of algae by herbivores was not observed in the other treatments, where algae were larger due to induced or permanent defences. In these treatments we observed rotifer population densities to first rise and then decline. The herbivore went extinct in all replicates with large permanently defended algae. The frequency of herbivore extinctions was intermediate when algae had inducible defences. 3. A variety of alternative mechanisms could explain differential herbivore persistence in the different defence treatments. Our analysis showed the density and fraction of highly edible algal particles to better explain herbivore persistence and extinctions than total algal density, the fraction of highly inedible food particles or the accumulation of herbivore waste products or autotoxins. 4. We argue that the rotifers require a minimum fraction and density of edible food particles for maintenance and reproduction. We conjecture that induced defences in algae may thus favour larger zooplankton species such as Daphnia spp. that are less sensitive to shifts in their food size spectrum, relative to smaller zooplankton species, such as rotifers and in this way contributes to the structuring of planktonic communities.     

Ecological determinants of rotifer vertical distribution in a coastal karst lake (Vrana Lake,Cres Island,Croatia)

Vrana Lake is a coastal karst lake on Cres Island within the eastern Adriatic coast. It appears to have formed as a cryptodepression on karstified carbonate rocks, and is filled with fresh water. It is classified as an oligotrophic hydrosystem and serves as a natural water supply reservoir (max. depth 74.5 m; surface area 5.5 km²; water volume 220 × 10⁶ m³). The aim of this study was to assess the influence of seasonally and spatially varying ecological conditions (i.e., thermal stratification, dissolved oxygen, orthophosphate and chlorophyll a concentrations) on the vertical distribution of the rotifer plankton community in this geomorphologically and hydrologically specific coastal lake. The rotifer community was surveyed monthly over a one year period. During the study period, a total of 21 rotifer taxa were identified. Among them Ascomorpha spp. (including A. ecaudis, A. ovalis and A. saltans), Filinia terminalis, Keratella cochlearis, Synchaeta gr. tremula-oblonga and Polyarthra vulgaris prevailed, representing 92% of the total rotifer abundance. Measured seasonal and spatial patterns of the rotifer assemblage along a vertical profile of Vrana Lake were considerably affected by temperature, food availability and biotic (competition, predation) interactions in the zooplankton community.     

Plankton dynamics in a tropical floodplain lake: fish, nutrients, and the relative importance of bottom-up and top-down control

1. Two enclosure experiments were carried out in Laguna Bufeos, a neotropical várzea lake located in the floodplain of River Ichilo (Bolivia). The experiments aimed (i) to assess the relative importance of bottom‐up and top‐down control on the plankton community, (ii) to assess the relative impact of direct and indirect effects of planktivorous fish on the zooplankton, and (iii) to attempt to identify the mechanisms responsible for these effects. 2. During the first experiment, bottom‐up control seemed to dominate the planktonic food web. Compared with fishless enclosures, oxygen concentrations, chlorophyll a levels and the population densities of all cladoceran zooplankton taxa increased in enclosures with fish. Birth rates of Moina minuta, the dominant taxon, were substantially higher in the presence than in the absence of fish, whereas death rates did not differ between treatments. These results are the first to suggest that the positive effects of fish on crustacean zooplankton via effects on nutrient cycling and the enhancement of primary production can compensate for losses because of fish‐related mortality. 3. During the second experiment, the direction of control appeared to vary between trophic levels: the phytoplankton appeared to be bottom‐up controlled whereas the zooplankton was mainly top‐down controlled. Chlorophyll a concentrations were enhanced by both fish and nutrient additions. The majority of the zooplankton taxa were reduced by the presence of fish. Birth rates of most cladoceran taxa did not differ between treatments, whereas death rates were higher in the enclosures with fish than in the fishless enclosures. Bosminopsis deitersi reached higher densities in the presence of fish, probably because of a release from predation by Chaoborus. 4. We convincingly showed strong deviations from trophic cascade‐based expectations, supporting the idea that trophic cascades may be weak in tropical lakes.     

Summer zooplankton dynamics in the limnetic and littoral zones of a humic acid lake

The limnetic and littoral zooplankton were studied during summer 1989 in Triangle Lake, a humic acid (pH 4.9) bog lake in Ohio, USA. The limnetic zooplankton showed low species richness and biomass, and dominance by the rotifers Kellicottia bostoniensis and Polyarthra vulgaris. In the littoral, species richness and biomass were markedly higher, and the crustaceans Alona guttata, Ceriodaphnia quadrangula, Chydorus sphaericus, Simocephalus serrulatus and Tropocyclops prasinus, and the rotifer Ascomorpha ecaudis were the dominants. The extreme rotifer dominance and lack of cladocerans in the limnetic zone were likely the result of Chaoborus predation. A pronounced mid-summer decline in cladoceran biomass in the littoral was likely due to predation by T. prasinus and Utricularia (bladderwort).     

Mechanisms regulating zooplankton populations in a high-mountain lake

SUMMARY 1. We studied the seasonal succession of phyto- and zooplankton and the potential impact of predation by salmonids on zooplankton population dynamics in a high-mountain Swiss lake.
2. A comparison of patterns in the abundance, body length, fecundity and age structure in the Daphnia galeata population strongly suggests that trout predation had little impact on the population and was not the cause for a decline in summer.
3. The dominance in the lake of adult trout that feed mainly on benthic prey may buffer the effect of predation on the larger zooplankton. Further, the relatively high amount of phytoplankton after spring thaw could be important for sustaining the Daphnia population under moderate fish predation.
4. Partial correlation analyses proved circumstantial evidence for both exploitative and interference competition between some zooplankton taxa. D. galeata depressed performance of other plankton species through exploitative competition.
5. Our study shows that the impact of fish on zooplankton in high-mountain lakes depends strongly on food web structure and trophic state of the lake. Where fish predation is weak, invertebrate predation combined with competition for food may be responsible for the dominance of large-bodied zooplankton species.