Interactive effects of productivity and predation on zooplankton diversity

Recent studies suggest the necessity of understanding the interactive effects of predation and productivity on species coexistence and prey diversity. Models predict that coexistence of prey species with different competitive abilities can be achieved if inferior resource competitors are less susceptible to predation and if productivity and/or predation pressure are at intermediate levels. Hence, predator effects on prey diversity are predicted to be highly context dependent: enhancing diversity from low to intermediate levels of productivity or predation and reducing diversity of prey at high levels of productivity or predation. While several studies have examined the interactive effects of herbivory and productivity on primary producer diversity, experimental studies of such effects in predator‐prey systems are rare. We tested these predictions using an aquatic field mesocosm experiment in which initial density of the zooplankton predator Notonecta undulata and productivity were manipulated to test their interactive effects on diversity of seven zooplankton, cladoceran species that were common in surrounding ponds. Two productivity levels were imposed via phosphorus enrichment at levels comparable to low and intermediate levels found within neighboring natural ponds. We used open systems to allow for natural dispersal and behaviorally‐mediated numerical responses by the flight‐capable predator. Effects of predators on zooplankton diversity depended on productivity level. At low and high productivity, prey species richness declined while at high productivity it showed a unimodal relationship with increasing the predator density. Effects of treatments were weaker when using Pielou's evenness index or the inverse Simpson index as measures of prey diversity. Our findings are generally consistent with model predictions in which predators can facilitate prey coexistence and diversity at intermediate levels of productivity and predation intensity. Our work also shows that the functional form of the relationship between prey diversity and predation intensity can be complex and highly dependent on environmental context.     

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.     

Planktivory by larval Odontesthes bonariensis (Atherinidae: Pisces) and its effects on zooplankton community structure

This paper describes the effect of predation by larvae of theatherinid Odontesthes bonariensis upon the zooplankton communityduring a 3-month experiment Three-day-old larvae were stockedin 45 m² concrete tanks at rates of 100 and 200 fish m^–2Gut content analyses showed that the larvae consumed relativelysmall prey all along the experiment Morphological (mouth width)and physiological (gastric inefficiency) constraints seems tohave precluded the capture of the largest prey. The prey community,however, showed all the symptoms generally ascribed to size-selectivepredation upon the largest individuals: decrease in maximumzooplankton size and mean cladoceran size, decrease in cladoceranand copepod biomass, extinction of the largest zooplankton species(Daphnia stmilis), increase in rotifer biomass, etc It is concludedthat size-selective predation is not a necessary condition forthe commonly observed decrease in zooplankton size after theincrease in density of a vertebrate predator. The ecologicalimplications of this result are discussed. Present address: Department of Biology, Lehigh University, WilliamsHall 31, Bethlehem, PA 18015-3189, USA     

Long-term responses of zooplankton to invasion by a planktivorous fish in a subarctic watercourse

1. Introduced or invading predators may have strong impacts on prey populations of the recipient community mediated by direct and indirect interactions. The long-term progression of predation effects, covering the invasion and establishment phase of alien predators, however, has rarely been documented.
2. This paper documents the impact of an invasive, specialized planktivorous fish on its prey in a subarctic watercourse. Potential predation effects on the crustacean plankton, at the community, population and individual levels, were explored in a long-term study following the invasion by vendace ( Coregonus albula ).
3. Over the 12-year period, the density and species richness of zooplankton decreased, smaller species became more abundant and Daphnia longispina , one of the largest cladocerans, was eliminated from the zooplankton community.
4. Within the dominant cladocerans, including Daphnia spp., Bosmina longispina and Bosmina longirostris , the body size of ovigerous females and the size at first reproduction decreased after the arrival of the new predator. The clutch sizes of Daphnia spp. and B. longirostris also increased.
5. Increased predation pressure following the vendace invasion induced many effects on the crustacean zooplankton, and we document comprehensive and strong direct and indirect long-term impacts of an introduced non-native predator on the native prey community.     

Perch or plankton: top‐down control of Daphnia by yellow perch (Perca flavescens) or Bythotrephes cederstroemi in an inland lake?

1. Seasonal termination of the vernal clear-water phase in Long Lake, Grand Traverse Co., Michigan coincided with severe size-selective predation on juvenile Daphnia pulicaria from 0.8 to 1.8 mm in length. This could be caused by predation by age-0 yellow perch ( Perca flavescens ) or by the exotic predatory zooplankter Bythotrephes cederstroemi .
2. During the initial decline of Daphnia , Ivlev's electivity coefficient for yellow perch from 15.0 to 20.0 mm in length was 0.50 for copepods and −0.75 for D. pulicaria .
3. Bioenergetics modelling of both yellow perch and Bythotrephes demonstrates that, during the initial Daphnia decline, Bythotrephes consumed 1.5–5 times greater total mass than yellow perch. Furthermore, models in which Bythotrephes consumed juvenile Daphnia were more consistent with the timing of the Daphnia decline than those in which yellow perch consumed juvenile Daphnia .
4. The invasion of Bythotrephes into Long Lake seems to be a significant perturbation, introducing effects that propagate throughout the food chain. Bythotrephes created a possible bottleneck for age-0 yellow perch in late June by suppressing Daphnia .     

Predator presence dramatically reduces copepod abundance through condition‐mediated non‐consumptive effects

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Interpopulation variation in a fish predator drives evolutionary divergence in prey in lakes

Ecological factors are known to cause evolutionary diversification. Recent work has shown that evolution in strongly interacting predator species has reciprocal impacts on ecosystems. These divergent impacts of predators may alter the selective landscape and cause the evolution of prey. Yet, this link between intraspecific variation and evolution is unexplored. We compared the life history of a species of zooplankton (Daphnia ambigua) from lakes in New England in which the dominant planktivorous predator, the alewife (Alosa pseudoharengus), differs in feeding traits and migratory behaviour. Anadromous alewife (seasonal migrants) exhibit larger gapes, gill-raker spacing and target larger prey than landlocked alewife (year-round freshwater resident). In 'anadromous' lakes, Daphnia are abundant in the spring but extirpated by alewife predation in summer. Daphnia are rare year-round in 'landlocked' lakes. We show that Daphnia from lakes with anadromous alewife grew faster, matured earlier but at the same size and produced more offspring than Daphnia from lakes with landlocked or no alewife across multiple temperature and resource treatments. Our results are inconsistent with a response to size-selective predation but are better explained as an adaptation to colder temperatures and shorter periods of development (countergradient variation) mediated by seasonal alewife predation.     

Effects of the non-indigenous cladoceran Cercopagis pengoi on the lower food web of Lake Ontario

1. In North America, the invasive predatory cladoceran Cercopagis pengoi was first detected in Lake Ontario. We explored the impact of Cercopagis on the lower food web of Lake Ontario through assessments of historical and seasonal abundance of the crustacean zooplankton, by conducting feeding experiments on the dominant prey of the invader, and by estimating its food requirements. 2. Between 1999 and 2001, a decrease in the abundance of dominant members of the Lake Ontario zooplankton community (Daphnia retrocurva, Bosmina longirostris and Diacyclops thomasi) coincided with an increase in the abundance of Cercopagis. Daphnia retrocurva populations declined despite high fecundity in all 3 years, indicating that food limitation was not responsible. Chlorophyll a concentration generally increased, concomitant with a decline in the herbivorous cladoceran zooplankton in the lake. 3. Laboratory experiments demonstrated that Cercopagis fed on small‐bodied species including D. retrocurva and B. longirostris. 4. Consumption demand of mid‐summer populations of Cercopagis, estimated from a bioenergetic model of the confamilial Bythotrephes, was sufficient to reduce crustacean abundance, although the degree of expected suppression varied seasonally and interannually. 5. Predatory effects exerted by Cercopagis on the Lake Ontario zooplankton, while initially very pronounced, have decreased steadily as the species became established in the lake.     

Predation by inland silversides on an exotic cladoceran, Daphnia lumholtzi, in Lake Texoma, U.S.A.

In the early 1990s, Lake Texoma was invaded by Daphnia lumholtzi , a large, spiny, exoticcladoceran. From April 1994 to April 1995 D. lumholtzi was most abundant in the lake in mid-summer, after native zooplankton declined in early June. Inland silversides Menidia beryllina selectively preyed on D. lumholtzi but appeared to prefer the large native zooplankton when they were present. Daphnia lumholtzi was an important prey item for silversides during summer when large native zooplankton was scarce. The invasion of D. lumholtzi into Lake Texoma may benefit some zooplanktivorous fishes by increasing foraging opportunities during a time of low prey availability.     

Cladoceran Densities,Day-to-Day Variability in Food Selection by Smelt,and the Birth-Rate-Compensation Hypothesis

Late-evening gut inspection of a dominant planktivore (smelt) and evaluation of densities, fecundities, and body-size distributions in dominant zooplankton prey (cladocerans) were made in day-to-day sequences in June–July (24 days in 1999 and 24 days in 2000). This was conducted as a field test of the hypothesis that species-specific population densities in cladocerans result from size-selective predation by a dominant fish assumed to be a general predator, switching from one prey to another as relative abundance changes. Little of the expected coincidence has been revealed between population density declines and increased numbers of a given prey in smelt diet. However, the data were consistent with the notion that fish would switch from one prey to another depending on the prey relative abundance (the number of prey a fish would see in its reaction field volume). Each cladoceran population fluctuated around its species-specific density level, lower or higher, depending on individual susceptibility to smelt predation, from 0.2 ind. l^–1 in large-bodied Daphnia hyalina and Leptodora kindtii, to 30.0 ind. l^–1 in small-bodied Daphnia cucullata andBosmina thersites. In spite of high fish-to-fish and day-to-day variability in both smelt diet and smelt selectivity for different prey, all cladocerans (also copepods and midge larvae) were equally persistent in smelt diet, and smelt selectivity was similar for small- and large-bodied prey categories, but lower for elongated-(Daphnia, Diaphanosoma) than for compact-body (Bosmina, Chydorus) species, when integrated for the entire sampling time. Closer examination of D. cucullata and B. thersites revealed strong smelt selection for later instars and females with greater clutches, showing that size distribution in a cladoceran population might be structured by fish predation in a similar way to that a cladoceran community (species relative abundance) is structured in a lake habitat. The birth-rate-compensation hypothesis is offered to explain why the value of food selectivity index in a planktivorous fish would remain the same for alternate prey categories with similar life-history traits, unless they differ in susceptibility to predation before the time of first reproduction.     

Seasonality and Fecundity of Daphnia Lumholtzi Sars in Lake Phewa,Nepal

Monthly sampling of zooplankton in Lake Phewa, Nepal revealed 18 species of zooplankton excluding protozoans. The seasonality and fecundity of the largest common cladoceran Daphnia lumholtzi Sars was studied. Two distinct peaks of abundance were noted. No active stages were found during May and June. Egg ratio varied from zero to 0.88 in mature females. From the presence of only helmeted individuals of this species it is concluded that the predation pressure by fish was high.     

Predator induced life-history shifts in a freshwater cladoceran

Summary Life-history theory predicts that maturity and resource allocation patterns are highly sensitive to selective predation. Under reduced adult survival, selection will favour genotypes capable of reproducing earlier, at a smaller size and with a higher reproductive effort. When exposed to water that previously held fish, (size selective predators which prefer larger Daphnia), individuals of Daphnia hyalina reproduced earlier, at a smaller size and had a higher reproductive investment. Hence the prey was able to switch its life history pattern in order to become less susceptible to predation by a specific predator. The cue that evokes the prey response is a chemical released by the predator.     

Evolution mediates the effects of apex predation on aquatic food webs

Ecological and evolutionary mechanisms are increasingly thought to shape local community dynamics. Here, I evaluate if the local adaptation of a meso-predator to an apex predator alters local food webs. The marbled salamander (Ambystoma opacum) is an apex predator that consumes both the spotted salamander (Ambystoma maculatum) and shared zooplankton prey. Common garden experiments reveal that spotted salamander populations which co-occur with marbled salamanders forage more intensely than those that face other predator species. These foraging differences, in turn, alter the diversity, abundance and composition of zooplankton communities in common garden experiments and natural ponds. Locally adapted spotted salamanders exacerbate prey biomass declines associated with apex predation, but dampen the top-down effects of apex predation on prey diversity. Countergradient selection on foraging explains why locally adapted spotted salamanders exacerbate prey biomass declines. The two salamander species prefer different prey species, which explains why adapted spotted salamanders buffer changes in prey composition owing to apex predation. Results suggest that local adaptation can strongly mediate effects from apex predation on local food webs. Community ecologists might often need to consider the evolutionary history of populations to understand local diversity patterns, food web dynamics, resource gradients and their responses to disturbance.     

Vulnerability of cladoceran species to predation by the copepod Mesocyclops leuckarti: laboratory observations on the behavioural interactions between predator and prey

SUMMARY 1. We analysed the vulnerability of a number of cladoceran species ( Bosmina longirostris , B. fatalis , Diaphanosoma brachyurum , Ceriodaphnia reticulata , Daphnia ambigua and D. pulex ) to predation by Mesocyclops leuckarti in the laboratory.
2. The prey species represented a wide range of body size, morphology, and swimming behaviour. To compare vulnerability, we measured the efficiency of capture and ingestion of each prey species by Mesocyclops . We also measured the rate at which prey were damaged in attacks by Mesocyclops .
3. Mesocyclops preyed effectively on Diaphanosoma and small juvenile Ceriodaphnia but not on Bosmina or Daphnia . Observations suggested that various defence mechanisms, including protruding structures and swimming behaviour and speed, are important in determining prey vulnerability.
4. The body size of Daphnia and Ceriodaphnia seems to be important, because larger animals were better able to escape Mesocyclops attacks. Attacks by Mesocyclops often caused fatal damage, however, even to large Daphnia .     

Effects of <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis> (Crustacea,Cladocera) on the abundance,morphology, and prey community of <Emphasis Type="Italic">Leptodora kindtii</Emphasis> (Crustacea,Cladocera)

We hypothesized that native Leptodora kindtii would be shorter and have smaller feeding baskets in central Ontario lakes with greater abundances of small-bodied zooplankton prey, and that differences in zooplankton size among lakes could be attributed to the invasive cladoceran Bythotrephes longimanus. We evaluated these conjectures by comparing size metrics of Leptodora and the size of their preferred cladoceran prey in lakes invaded or not by Bythotrephes. Leptodora was less abundant in invaded lakes, but were smaller bodied with smaller feeding baskets only in lakes with long invasion histories. Small cladoceran abundance was greater in non-invaded lakes and was directly related to Leptodora abundance although not to Leptodora size. Mean Leptodora body size declined with increasing abundance of Bythotrephes. We evaluated three possible explanations for these patterns in Leptodora—(a) competition with Bythotrephes for zooplankton prey, (b) direct predation by Bythotrephes, and (c) size-selective predation by fish. While we were unable to unequivocally distinguish among these hypotheses, our observations are most consistent with predation by Bythotrephes changing zooplankton community composition and size structure in a manner that is detrimental to Leptodora. Our results indicate that Bythotrephes invasion may trigger more complex and subtle changes in food webs than previously thought.     

Damage released prey alarm substances or predator odours? Risk assessment by an aquatic oligochaete

Although the abilities of prey to detect and respond to chemical substances associated with a predator have been widely reported, the factors promoting the evolution of responses to prey alarm cues vs. predator odours are still vague. In this article, we combined field research with laboratory experiments to explore which chemical substance associated with predator activity (predator odour, conspecific or heterospecific alarm substances) induces defence responses in the aquatic oligochaete Stylaria lacustris, which is vulnerable to common littoral predators. The field results indicated that predators injure the oligochaetes and a great proportion, up to 45% of individuals in the population, were found to be damaged. The results of the laboratory experiments revealed that chemical odours from damselfly larvae feeding on S. lacustris did not induce the defence response in the oligochaetes. On the contrary, oligochaetes detected and responded to alarm substances from damaged conspecifics alone and substances from damaged cladoceran Daphnia magna. We discussed conditions favouring the responses to damage released prey alarm cues instead of predator odours in Stylaria lacustris. Our data suggest that the selection of responses to alarm cues from damaged prey vs. predator odours may be dependent on three factors: (1) non-species-specific predation, (2) divergence of food niche of the different stages of the predator and (3) complex food web with multiple predators. Handling editor: S. Declerk     

The relationship between cladoceran body size and the growth of underyearling roach (Rutilus rutilus) (L.) in two shallow lowland lakes: a mechanism for density-dependent reductions in growth

Large planktonic Cladocera are typically the most important components of the diet of underyearling roach. Selection for large species and individuals by fish can result in a shift in the species composition of the cladoceran community as well as a reduction in the mean size of the individuals of large species and in the assemblage as a whole. Laboratory experiments demonstrated that underyearlings feeding on Daphnia hyalina smaller than 1 mm in length has a significantly lower intake of prey volume per unit time than when feeding on prey greater than 1.5 mm. A decrease in the nutritional quality of zooplankton prey, brought about by increasing predation pressure is suggested as the mechanism for density-dependent reductions in the growth of underyearling roach in eutrophic water bodies.     

Preference for Cannibalism and Ontogenetic Constraints in Competitive Ability of Piscivorous Top Predators

Occurrence of cannibalism and inferior competitive ability of predators compared to their prey have been suggested to promote coexistence in size-structured intraguild predation (IGP) systems. The intrinsic size-structure of fish provides the necessary prerequisites to test whether the above mechanisms are general features of species interactions in fish communities where IGP is common. We first experimentally tested whether Arctic char (Salvelinus alpinus) were more efficient as a cannibal than as an interspecific predator on the prey fish ninespine stickleback (Pungitius pungitius) and whether ninespine stickleback were a more efficient competitor on the shared zooplankton prey than its predator, Arctic char. Secondly, we performed a literature survey to evaluate if piscivores in general are more efficient as cannibals than as interspecific predators and whether piscivores are inferior competitors on shared resources compared to their prey fish species. Both controlled pool experiments and outdoor pond experiments showed that char imposed a higher mortality on YOY char than on ninespine sticklebacks, suggesting that piscivorous char is a more efficient cannibal than interspecific predator. Estimates of size dependent attack rates on zooplankton further showed a consistently higher attack rate of ninespine sticklebacks compared to similar sized char on zooplankton, suggesting that ninespine stickleback is a more efficient competitor than char on zooplankton resources. The literature survey showed that piscivorous top consumers generally selected conspecifics over interspecific prey, and that prey species are competitively superior compared to juvenile piscivorous species in the zooplankton niche. We suggest that the observed selectivity for cannibal prey over interspecific prey and the competitive advantage of prey species over juvenile piscivores are common features in fish communities and that the observed selectivity for cannibalism over interspecific prey has the potential to mediate coexistence in size structured intraguild predation systems.     

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.     

Different Predation Impacts of Two Cyclopoid Species on a Small-sized Zooplankton Community: An Experimental Analysis with Mesocosms

We used mesocosms to analyze predation impacts on the prey populations and prey community structures by two cyclopoid copepod species, the larger Mesocyclops pehpeiensis and the smaller Thermocyclops taihokuensis, who coexist with small-sized herbivorous zooplankton species in a fish-abundant lake. The overall predation impact on the prey populations was stronger for Mesocyclops than for Thermocyclops. Mesocyclops had a strong and less selective impact on the rotifer community but a selective impact on the crustaceans. In contrast, Thermocyclops had a selective predation impact on rotifers but a weak and less selective impact on the crustacean community. As a result, the former predator reduced the diversity of the crustacean community but not the rotifer community, while the latter had an opposite impact on the diversities of the two communities. It has been suggested that fish induce development of a zooplankton community dominated by the small-sized zooplankton species in fish-abundant lakes. Our results demonstrated that cyclopoid copepods altered species composition and diversity of the small-sized zooplankton community in such lakes. Thus, the results have given an important suggestion on the role of the invertebrate predator cyclopoid copepods, which often coexist with fish, that they determine population dynamics and community structures of small-sized zooplankton in fish-abundant lakes.