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.     

The predator-prey relationship of perch,Perca fluviatilis,larvae and zooplankton in two Scottish lochs

Synopsis Food consumption of perch larvae and the impact of this on zooplankton were examined in two adjacent shallow Scottish lochs. Maximum annual abundance of zooplankton occurred in mid-May at L. Kinord with minimum values in mid-June. Copepods were prominent in spring but were followed by a multi-species community of cladocerans and rotifers in summer. At L. Davan zooplankton biomass remained high through summer with cladocerans dominating andDaphnia longispina the most frequent species. Availability of food items was a principal factor governing feeding behaviour of larvae. Copepodite stages were initially the most common item in the diet in L. Kinord in 1976 and 1977 and rotifers the principal food in June 1977, reflecting the dominance of these items in the zooplankton. Cladocerans were dominant in the plankton community in L. Davan and constituted the greater part of food intake. Overlying this general pattern there was an increase in the size of food items taken by larvae with time and also a definite pattern of food selection for copepods, with initially selection for smaller copepodite stages and later for larger stages and adults. On most occasions larvae selected forCyclops strenuus abyssorum andPolphemus pediculus and selected againstDaphnia longispina. The reduction in the total zooplankton biomass attributed to perch larvae was minimal, with the exception of mid-June at L. Kinord in 1976. However, predation on particular species and copepodite stages was occasionally intense and may have impacted the zooplankton populations.     

A comparison between predation effects on zooplankton communities by Neomysis and Chaoborus

Community level effects of predation by two invertebrate predators, the opossum shrimp (Neomysis intermedia), and the larva of the phantom midge (Chaoborus flavicans) were studied and compared. N. intermedia appeared abundantly in the shallow eutrophic Lake Kasumigaura and had a significant impact on the zooplankton community. The predation pressure by Neomysis was highest on cladocerans, followed by rotifers, and finally copepods. At high densities (maximum nearly 20 000 individuals m^–2), Neomysis excluded almost all cladocerans, rotifers and copepods from the lake.Zooplankton communities were established in experimental ponds, into which C. flavicans was introduced. The predator's density was around 1 individual l^–1, and was probably controled by cannibalism. Although Chaoborus larvae feed on various zooplankton species, their predation impact on zooplankton populations was markedly selective. They eliminated medium- and small-sized cladocerans and calanoid copepods from the ponds, but rotifers increased.Although the feeding selectivities of Neomysis and Chaoborus individuals were similar, the predation effects on zooplankton communities by the two predators were different.     

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.     

Zooplankton dynamics in a gypsum karst lake and interrelation with the abiotic environment

Zooplankton species composition and abundance variation was studied in Lake Amvrakia, which is a deep, temperate, gypsum karst lake situated in the western Greece. The two year survey of zooplankton revealed 33 species (23 rotifers, five cladocerans, four copepods and one mollusc larva). The mean integrated abundance of the total zooplankton ranged between 83.6 and 442.7 ind. L⁻¹, with the higher density to be recorded in the surface 0–20 m layer. Small numbers of specimens of almost all species were found also in the hypoxic or anoxic hypolimnion. Copepods and especially the calanoid Eudiaptomus drieschi dominated the zooplankton community throughout the sampling period, followed by Dreissena polymorpha larvae, rotifers and cladocerans. Seasonal succession among the cladocerans and the most abundant rotifer species was observed. The concentration of chlorophyll-a was the most important factor for the variation of total zooplankton, as well as for the rotifers’ community. Dissolved oxygen affected copepods and cladocerans, water level correlated mainly with the molluscs larvae of D. polymorpha, while temperature influenced the variation of several rotifers, the cladoceran Diaphanosoma orghidani and the mollusc larvae. Negative correlation of conductivity with the cladoceran Daphnia cucullata and the copepods E. drieschi and Macrocyclops albidus was found. The differences in species composition found in Lake Amvrakia in comparison to the nearby lakes are probably ought to the geographical isolation and perhaps to its particular chemistry (e.g., elevated conductivity).     

Absence of typical diel vertical migration in Daphnia: varying role of water clarity, food, and dissolved oxygen in Lake Eymir, Turkey

Diel vertical migration (DVM) is a complex and dynamic behaviour against predation because the reaction of migrating organisms to light intensity plays a primary role, but is modified by other factors. In the relatively shallow but thermally stratified Lake Eymir, Daphnia pulex de Geers utilized vertical refugia afforded by the hypolimnion during both day and night. Differences in general vulnerability to fish predation determined the differences in their mean residence depths (MRDs) of different population categories such as most conspicuous and vulnerable individuals of adult with eggs inhabited the deepest depth, whereas juveniles stayed close the thermocline. In late spring, profoundly high amplitude of displacement within the hypolimnion, probably due to the hypolimnion being well-lit and relatively well-oxygenated for the fish and rather unsafe for the large-sized daphnids, was recorded. Therefore, the large-sized daphnids daytime refuge was close to the bottom whereas at night they moved upward to benefit from warmer water temperature along with food availability in the presence of fish predation but still remained below the thermocline. In summer, the insignificant amplitude of the hypolimnetic, which later became epilimnetic, displacements were probably due to the near-anoxic condition found below the thermocline. This might have deterred the fish, thus providing a safer refuge for daphnids in the below thermocline, which afterwards became the above thermocline. Low oxygen availability was regarded as the summer proximate factor. The abundant food and warmer water conditions found in the below/above thermocline also accounted for absence of DVM in summer. Consequently, this study suggests that DVM by Daphnia is an adaptation that is plastic to changing environmental conditions.     

Experimental analysis of the predation impact of the larvae of pond smelt (<Emphasis Type="Italic">Hypomesus transpacificus nipponensis</Emphasis>) on zooplankton populations established in mesocosms

The predation impact of the larvae of pond smelt Hypomesus transpacificus nipponensis on a zooplankton community was studied using mesocosms. The fish significantly depressed the abundances of copepod nauplii and rotifers, especially Hexarthra mira. The vulnerabilities of these prey might be determined by their swimming behavior and population density, suggesting that larval fish selectively prey on zooplankton that have a high encounter rate with the predator. The larvae did not have a negative effect on the densities of cladocerans, but fish predation altered the cladoceran community structure from the dominance of B. longirostris to that of B. fatalis. This result suggests that larval fish predation is an important factor that shifts the species composition of Bosmina in some lakes, the shift occurring in the season when fish larvae are abundant. Our results have shown that predation by the larval fish would control not only the abundance, but also the community structure of the small-sized zooplankton prey.     

Zooplankton of a flooded opencast sulphur mine

The zooplankton community of a flooded opencast sulphur mine was investigated during two years. The complicated physical and chemical relationships in this water-filled pit caused an atypical plankton assemblage. At the time of the investigation 20 species of rotifers, 6 of Cladocera, and 8 species of Copepoda were found. A single Ostracod and Chaoborus flavicans were also noted in the samples. Usually the epilimnion was monopolised by rotifers whereas the hypolimnion had a more diversified assemblage of rotifers, cladocerans and copepods. The result of clustering analysis indicated that each predator was associated with its own group of prey. Discrimination analyses showed weak seasonal differences in the zooplankton and three zones in the vertical profile – epilimnetic, upper hypolimnetic and anaerobic. Principal component analysis reduced the chemical parameters to the main contributors of conductivity (the main anions and cations) plus toxic sulphide and hydrogen sulphide. The animals were classified by the first two components into two groups: cold stenotherms and medium-sized filtrators, such as Bosmina longirostris. A specific analysis of vertical profiles generated five variants of diel vertical migrations. When time–space distributions of the plankters were analysed the highest numbers of different species were usually in different times and depths. The epilimnion was inhabited by small fishes and was dominated by rotifers but was free from C. flavicans which inhabited the cold hypolimnion, together with the remaining invertebrate predators (copepods and Asplanchna).Chaoborus and the other invertebrate predators participated more evenly in the contributions of species to the total density. In the presence of invertebrate predators the structure of the rotifer population shifted from soft-bodied species to spined or fast-moving species. When the predation pressure was too strong, as in the hypolimnion, the soft-bodied species were, in practice, absent. When the pit was in an oligotrophic state the hypolimnion population was composed of up to 60–90% rotifers. When the trophic state shifted to mesotrophy, the proportion of cladocerans increased.     

Refuge availability and sequence of predators determine the seasonal succession of crustacean zooplankton in a clay-turbid lake

The contribution of predators and abiotic factors to the regulation of the biomass and seasonal succession of crustacean zooplankton was studied in Lake Rehtijärvi (southern Finland). Field data in combination with bioenergetics modeling indicated that invertebrate planktivory by Chaoborus depressed cladoceran populations during early summer. In particular, bosminids that generally form the spring biomass peak of cladocerans in stratified temperate lakes did not appear in the samples until July. In July, predation pressure by chaoborids was relaxed due to their emergence period and cladoceran population growth appeared to be limited by predation by planktivorous fish. The effect of fish predation was amplified by reduced refuge availability for cladocerans. The concentration of dissolved oxygen below the epilimnion was depleted, forcing cladocerans to move upward to less turbid and thus more dangerous water layers. The effect of size selective predation by fish resulted in reduced mean size of cladocerans during the period when refuge thickness (thickness of the water layer with oxygen concentration <1 mg l^?1 and water turbidity >30 NTU) was lowest. The results confirmed that in clay-turbid lakes, invertebrate predators could be the main regulators of herbivorous zooplankton even when cyprinid fish are abundant.     

Effects of flood on the functioning of the Dobczyce reservoir ecosystem

The effects of two summer floods, in 1997 and 2001 on phytoplankton, zooplankton and fish in the Dobczyce reservoir are presented. Shifts in phytoplankton distribution (from hypolimnion into the whole water column) and species composition (domination of diatoms after the flood) were observed. High water flow eliminated large species of cladocerans and copepods (the most effective filtrators) and favoured development of rotifers. Both, the total zooplankton biomass and chlorophyll a concentration after the flood dropped considerably. In the case of fish, the observed changes in their distribution and decrease in concentration were attributed to their behaviour. During the flood, fish were avoiding open water also during the night, but two weeks following the flood they returned to their usual migratory behaviour. The Dobczyce reservoir ecosystem showed great regeneration abilities to recover after the flood.     

Direct and indirect impacts of predation by fish on the zooplankton community: an experimental analysis using tanks

The impact of Pseudorasbora parva, a common zooplanktivorous fish species in Japan, on a zooplankton community was analyzed in experimental tanks, half of which were stocked with the fish. Different zooplankton species showed different responses to the introduction of the fish. In the presence of the fish, the populations of the large cladoceran Ceriodaphnia and the predatory copepod Mesocyclops were reduced, but the population of the herbivorous copepod Eodiaptomus and the small cladocerans Bosmina fatalis and Bosminopsis deitersi increased relative to the controls. The increase of Mesocyclops seen in the control tanks might have suppressed the populations of the small cladocerans, which are vulnerable to invertebrate predation. The results suggest that the population densities of the large prey items preferred by the fish, Ceriodaphnia and Mesocyclops, were controlled directly by fish predation, but the population densities of the smaller and less preferred zooplankton were controlled indirectly through the food-web cascade.     

Finding the optimal vertical distribution: behavioural responses of Daphnia pulicaria to gradients of environmental factors and the presence of fish

1. The vertical distribution of zooplankton results from active habitat choice aiming to optimise fitness gain in a system of trade‐offs. 2. Using large, controlled indoor mesocosms (Plön Plankton Towers), we monitored the behavioural response of Daphnia pulicaria to vertical gradients of temperature, food, oxygen and light, in the presence and absence of fish predation. 3. In the absence of fish, Daphnia distributed as predicted by an ideal ‘free distribution with costs’. If the food was distributed homogeneously, they stayed in the warm epilimnion, while they balanced their time dwelling in epi‐ and hypolimnion if the food was concentrated in a deep‐water maximum. 4. However, oxygen depletion in the hypolimnion, representing an additional cost, prevented Daphnia from completely exploiting the hypolimnetic food maximum. Consequently, the proportion dwelling in the hypolimnion was larger if oxygen was not limiting. 5. Fish predation had an overwhelming effect, driving Daphnia into the hypolimnion under all experimental conditions. If permitted by oxygen availability, Daphnia used the whole hypolimnion, but oxygen depletion reduced their possible habitat to the upper hypolimnion with oxygen concentrations above c. 0.7 mg L^?1. As fish were less tolerant of low oxygen, the layer below the thermocline formed a predation refuge for Daphnia.     

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).     

Effect of vertebrate predation on the spatio-temporal distribution of cladocerans in a temperature eutrophic lake

We analysed the spatio-temporal distribution of zooplankton along a profile of 10 stations from the shore to the pelagic zone from April to September 1988, the period when the larvae and juveniles Rutilus rutilus, the most abundant species in the Lake, are in the littoral zone. The digestive tracts of the young roach were analysed. They fed essentially on rotifers and on cladocerans. For comparison, zooplankton was also analysed at one littoral area without fish fry. There was an increase of cladoceran density from the vegetated nearshore zone to the offshore zone. Considering the density of Bosmina longirostris, Daphnia longispina, Chydorus sphaericus and Ceriodaphnia quadrangula, we observed a different distribution pattern in the course of the year. In the nearshore zone, the relative abundance of small species, Bosmina and Chydorus, was much higher than that of the larger Daphnia. From April to September, predation pressure mainly affected the smallest species: in contrast to the inshore station without fish fry, the density of Bosmina decreased in May in the littoral with fish. Chydorus was concentrated in the littoral between February and April, then grew into the pelagic zone, where predation pressure obviously was low during the warm season. The number of Daphnia, which was eaten by the fish fry at any time, remained low in the nearshore zone, which suggests that the presence of fish may cause Daphnia to avoid this zone. Ceriodaphnia which was not affected by this predation, was scarce in the nearshore zone during mid-summer. The low density of the cladocerans in the nearshore zone is likely associated with vertebrate predation by roach fry and juveniles, the result of such a process being either a depletion in density of the prey, or an avoidance behaviour.     

A Review of Predation Impact by 0+ Fish on Zooplankton in Fresh and Brackish Waters of the Temperate Northern Hemisphere

To assess potential differences in predation impact on zooplankton communities by small (larva) and large 0+ juvenile fish, 18 studies were reviewed from fresh water and the brackish Baltic Sea of the northern hemisphere temperate region. These case studies were performed either in the field or in mesocosm experiments. Larva stocks were found to exert only minor impact on small zooplankton species such as rotifers, copepodids and small cladocerans. In contrast, stocks of 0+ juveniles were found to have the potential to depress populations of large cladocerans and copepods, especially during late summer and autumn. However, studies where both 0+ juvenile fish consumption and zooplankton dynamics and production were exactly quantified are still very rare, and therefore final evaluation of this interaction cannot be made. In addition, papers were summarized that describe differences in morphological and physiological performance between larva and 0+ juvenile fish. The greater impact of 0+ juvenile fish on large zooplankton may be explained by their larger mouth gape and by their better developed abilities to detect and consume their prey items. However, this partly is lessened by the lower energy requirements of juvenile fish compared with identical biomasses of fish larvae, although larva bioenergetics remains only fragmentarily understood. Consequently, selective predation by fish larvae on particular small zooplankton prey may be more important than has been detected so far.     

Creating conditions for changes in prey community structure by Chaoborus spp. in a lake in Sweden

In Lake G»rdsjön (Southwest Sweden), liming as an experimental improvement of living conditions for pelagic algae, resulted in a significant increase of algal biomass and a reduction of mean cell size. The algal development was beneficial for small sized filter feeding zooplankton, particularly rotifers, which showed a significant increase. The increase in abundance of small sized zooplankton created better food conditions for the smaller instars, and thus a much better overall survival of Chaoborus larvae. The resulting, 6–7 times larger population of Chaoborus larvae significantly changed the structure of the crustacean zooplankton community. Bosmina coregoni, the fastest swimmer of the crustacean species suffered most and was strongly reduced by the increased predation from Chaoborus. The share of cladocerans decreased, while copepods increased in importance.     

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.     

Significance of a low oxygen layer for a Daphnia population in Lake Yunoko,Japan

Population dynamics and vertical migration of Daphnia longispina in Lake Yunoko were studied. The Daphnia population was small in spring and early summer, probably because of high predation pressure by fish. The population grew in midsummer, when thermal stratification developed and the dissolved oxygen became very low in the deeper layer of the hypolimnion. In this season, adults of D. longispina concentrated in the daytime near the lake bottom, where fish were absent because of the anoxic conditions, but ascended at night to the upper layer of the hypolimnion, where food was most abundant. The low oxygen layer near the bottom kept out the predators and protected Daphnia from predation, and consequently contributed to the built-up of its population. However, the low oxygen layer was unfavorable for reproduction of Daphnia, as reflected in the low egg ratio and high percentage of males in the population. The population decreased in the fall, when thermal stratification disappeared and predation pressure seemed to increase.     

Zooplankton dynamics in the upstream part of Stratos reservoir (Greece)

The investigation of the zooplankton community in the upstream part of Stratos reservoir during a 24 months survey (September 2004–August 2006) revealed 26 invertebrate species (14 rotifers, 6 cladocerans, 5 copepods and one mollusk larva). The mean abundance of the total zooplankton was higher in the first sampling period (2004–2005) and ranged between 8.81 and 47.74 ind. L⁻¹, than the second period (2005–2006) when fluctuated between 1.91 and 43.09 ind. L⁻¹. The seasonal variation was strongly influenced by the presence of rotifers, which accounting on average for 68.4% in total. Among them Keratella cochlearis and the order Bdelloidea were numerically the most important, while Macrocyclops albidus prevailed among the copepods and Bosmina longirostris among the cladocerans. Dreissena polymorpha was the only mollusk found in the zooplankton community. Rotifers, copepods and cladocerans showed a seasonal succession with the former preceding in the abundance having their first maximum in spring, while copepods and cladocerans followed, having peaks of abundance in early summer and in autumn, respectively. No seasonal succession among the cladoceran species was observed. The intense water flow in the upstream part of the reservoir, as well as temperature, conductivity, DO, pH, phosphates and silicates, were significant parameters controlling abiotic and biotic elements of the ecosystem and consequently influencing the seasonal variation and the dynamics of the zooplankton community.     

Zooplankton in Rainbow Bay, a Carolina bay pond: population dynamics in a temporary habitat

SUMMARY.

• 1 Rainbow Bay is a temporary freshwater pond in South Carolina, U.S.A., that typically fills in winter and dries in spring or summer. We studied population dynamics of zooplankton in 1984 and 1985 when it held water for 282 and 57 days, respectively. The zooplankton were diverse, including one species of calanoid copepod, nine or more cyclopoid copepods, nineteen or more cladocerans, and twelve or more rotifers. The community was initially dominated by the copepods Diaptomus stagnalis and Acanthocyclops vernalis or Diacyclops haueri, later by cyclopoids and cladocerans including Daphnia laevis and Simocephalus spp.
• 2 Laboratory experiments with sediments from the dry pond support the inference from field data that time of emergence from resting stages was a proximate cause of the initial succession of species. The experiments also showed that eggs of the calanoid copepod, Diaptomus stagnalis, were inhibited from hatching in early November, but became ready to hatch by late November. A similar inhibition was not observed for cladocerans or cyclopoid copepods.
• 3 Food limitation appeared to influence population dynamics following emergence. For the cladocerans Daphnia laevis and Simocephalus spp., great reductions in size-specific fecundity occurred 1 month (1985) or 3 months (1984) after the pond filled and were associated with disappearance of the rotifer Conochilus hippocrepis. The cladocerans produced ephippia when their fecundities declined. Reproduction of Diaptomus occurred before cladoceran fecundities decreased in 1984, but during their decline in 1985. Broods of Diaptomus were smaller in 1985.
• 4 The impact of invertebrate predators was probably greater than that of vertebrate predators, but neither was sufficient to prevent over-exploitation of food resources. Prey consumption rates by salamander larvae were estimated from diets and population densities of the larvae in 1984 (Taylor et al., 1988). Their impact on the zooplankton was low in February, increased in March and April as the larvae grew, and then