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.     

Indirect food web effects of <Emphasis Type="Italic">Bythotrephes</Emphasis> invasion: responses by the rotifer <Emphasis Type="Italic">Conochilus</Emphasis> in Harp Lake,Canada

As a recent invader of North American lakes, Bythotrephes longimanus has induced large changes in crustacean zooplankton communities through direct predation effects. Here we demonstrate that Bythotrephes can also have indirect food web effects, specifically on rotifer fauna. In historical time series data, the densities of the colonial rotifer Conochilus unicornis significantly increased after Bythotrephes invasion in Harp Lake, Ontario. No such changes were observed in a non-invaded reference lake, the nearby Red Chalk Lake. Evidence for two mechanisms explaining the Conochilus increase was examined based on changes to the crustacean zooplankton community over time. Rapid and severe declines in several herbivorous species of cladoceran zooplankton after Bythotrephes detection indicated a decrease in exploitative competition pressure on Conochilus. Secondly, a later and significant decline to virtual extinction of native invertebrate predators (Mesocyclops and Leptodora) could account for the observed Conochilus increase which also began 1–2 years after invasion by Bythotrephes. Ultimately, it appears that both reduced competition followed by a loss of native invertebrate predators were necessary to lead to the large Conochilus densities observed following invader establishment. From this analysis of long-term community data, it appears that Bythotrephes has important indirect, as well as direct, food web effects in newly invaded North American lakes with implications for trophic relationships.     

Patterns of <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis> distribution relative to native macroinvertebrates and zooplankton prey

As exotic species are introduced and spread across a heterogeneous landscape, the abundance and richness of potential competitor and prey species they encounter will vary. Little is known about the interactions between Bythotrephes longimanus and native predatory macroinvertebrates (e.g., Mysis, Chaoborus), which potentially limit the establishment and spread of the invader. An 80-lake survey was conducted in the summer of 2007 to obtain macroinvertebrate abundances across invaded and non-invaded lakes. A subset (15) of these lakes was surveyed more intensively to obtain stratified daytime and night-time distributions of the organisms. Overall co-occurrence of Bythotrephes with native macroinvertebrate predators was widespread across lakes indicating that the presence of native macroinvertebrates alone is unlikely to limit the establishment of Bythotrephes. However, we did find an effect of native macroinvertebrate predators on the vertical distribution of Bythotrephes: as native macroinvertebrate abundances increased, the relative abundance of Bythotrephes in the epilimnion increased. Furthermore, the relative abundance of some zooplankton prey (e.g., Daphnia) was lower in the epilimnion when Bythotrephes abundance was high. Although we cannot rule out consumptive effects, some evidence suggests an avoidance behavioural response in the prey. While the underlying mechanisms of these distributional shifts remain unclear, our results suggest that interactions between Bythotrephes, native macroinvertebrates and zooplankton prey are complex, highlighting the need to further examine these interactions.     

A plague of waterfleas (<Emphasis Type="Italic">Bythotrephes</Emphasis>): impacts on microcrustacean community structure,seasonal biomass,and secondary production in a large inland-lake complex

The spiny cladoceran (Bythotrephes longimanus) is an invasive, predaceous zooplankter that is expanding from Great Lakes coastal waters into inland lakes within a northern latitudinal band. In a large, Boundary Water lake complex (largely within Voyageurs National Park), we use two comparisons, a 2-year spatial and a 12-year temporal, to quantify seasonal impacts on food webs and biomass, plus a preliminary calculation of secondary production decline. Bythotrephes alters the seasonal biomass pattern by severely depressing microcrustaceans during summer and early fall, when the predator is most abundant. Cladoceran and cyclopoid copepods suffer the most serious population declines, although the resistant cladoceran Holopedium is favored in spatial comparisons. Microcrustacean biomass is reduced 40–60 % and secondary production declines by about 67 %. The microcrustacean community shifts towards calanoid copepods. The decline in secondary production is due both to summer biomass loss and to the longer generation times of calanoid copepods (slower turnover). The Bythotrephes “top-down” perturbation appears to hold across small, intermediate, and large-sized lakes (i.e. appears scale-independent), and is pronounced when Bythotrephes densities reach 20–40 individuals L^?1. Induction tests with small cladocerans (Bosmina) suggest that certain native prey populations do not sense the exotic predator and are “blind-sided”. Failure of prey to deploy defenses could explain the disproportionate community impacts in New World versus Old World lakes.     

<Emphasis Type="Italic">Bythotrephes</Emphasis> invasion elevates trophic position of zooplankton and fish: implications for contaminant biomagnification

We estimated the effects of Bythotrephes longimanus invasion on the trophic position (TP) of zooplankton communities and lake herring, Coregonus artedi. Temporal changes in lacustrine zooplankton communities following Bythotrephes invasion were contrasted with non-invaded reference lakes, and along with published information on zooplankton and herring diets, formed the basis of estimated changes in TP. The TP of zooplankton communities and lake herring increased significantly following the invasion of Bythotrephes, whereas TP in reference lakes decreased (zooplankton) or did not change significantly (lake herring) over a similar time frame. Elevated TP following Bythotrephes invasion was most prominent in lakes that also supported the glacial relict, Mysis diluvania, suggesting a possible synergistic interaction between these two species on zooplankton community composition. Our analysis indicated that elevated TPs of zooplankton communities and lake herring are not simply due to the presence of Bythotrephes, but rather reflect changes in the zooplankton community induced by Bythotrephes; namely, a major reduction in the proportion of herbivorous cladoceran biomass and a concomitant increase in the proportion of omnivorous and/or predatory copepod biomass in invaded lakes. We demonstrated that increases in TP of the magnitude reported here can lead to substantial increases in fish contaminant concentrations. In light of these results, we discuss potential mechanisms that may be responsible for the disconnect between empirical and theoretical evidence that mid-trophic level species invasions (e.g., Bythotrephes) elevate contaminant burdens of consumer species, and provide testable hypotheses to evaluate these mechanisms.     

Influence of light on the foraging impact of an introduced predatory cladoceran,Bythotrephes longimanus

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The ecological niches of <Emphasis Type="Italic">Bythotrephes</Emphasis> and <Emphasis Type="Italic">Leptodora:</Emphasis> lessons for predicting long-term effects of invasion

We here exploit two large datasets on zooplankton in Norwegian lakes, spanning a wide range of geographical, physical, chemical and biological properties, to assess the ecological niches and habitats of Bythotrephes longimanus and Leptodora kindtii. The species overlapped geographically, yet co-occurred only in a limited number of lakes. Bythotrephes inhabited virtually all types of lakes, except alpine localities and productive lakes dominated by cyprinid communities where the hyaline Leptodora was most abundant. The zooplankton communities also differed in Bythotrephes and Leptodora lakes, probably both reflecting different predatory regimes, but also water quality and other lake-specific properties. We found no evidence for species being excluded by the presence of Bythotrephes, rather the diversity in general was higher in lakes with these predators present compared with those without. We found, however, a very close association between Bythotrephes and Daphnia galeata and to some extent also between Bythotrephes and D. longispina, suggesting that these species also may benefit from Bythotrephes invasion. Both Bythotrephes and Leptodora species occur naturally in this region, and knowledge about the ecological preferences and the zooplankton community composition in Bythotrephes—and Leptodora lakes will provide valuable information about the long-term effects of Bythotrephes invasion and potential interaction with of Leptodora as top invertebrate predator.     

Zooplankton spine induces aversion in small fish predators

Summary The spined cladoceran Bythotrephes cederstroemi is protected from small fish predators due to the difficulty small fish have in ingesting the spine. Juvenile yellow perch (Perca flavescens) 50–60 mm in length were offered Bythotrephes with alternative prey available in two experiments. First, perch were observed as they gained experience with Bythotrephes and developed aversion to the zooplankter. Perch initially attacked and captured Bythotrephes. However, they struggled to ingest the spined zooplankter, rejected and recaptured it many times, and finally ceased to attack it. Second, perch were offered Bythotrephes with varying portions of the spine removed to clarify the spine's role in inducing such behaviors. Perch showed greater preference to attack nospine and half-spine Bythotrephes, and were less likely to reject and more likely to ingest Bythotrephes with the spine removed. For small or young fish that forage on zooplankton in lakes where Bythotrephes is present, aversion is an efficient response to the conspicuous but unpalatable spined cladoceran. However, aversion allows Bythotrephes, also a predator on zooplankton, to more effectively compete with young fish without an increase in predation risk.     

Temporal and ontogenetic variation in the diet of squid (<Emphasis Type="Italic">Loligo forbesii</Emphasis> Streenstrup) in Scottish waters

The squid Loligo forbesii is the only cephalopod species currently targeted by fisheries in the northern NE Atlantic. An active predator, it feeds primarily on fish, crustaceans and cephalopods. During 15 years since the only previous large-scale study of the diet of this species in Scottish waters, there have been substantial changes in marine fish abundances. The present study evaluates sources of variation (temporal, ontogenetic) in diet composition and prey size preferences of L. forbesii, including a comparison of contemporary (July 2006–June 2007) and historical (1990–1992) dietary datasets. Results revealed significant size-related and seasonal variation in diet composition and prey size. Teleost fish of the families Ammodytidae and Gobiidae were eaten by squid of all sampled sizes, although occurrence of gobies was generally more frequent in smaller squids, while Gadidae were eaten more frequently by larger squids. Cannibalism was also more frequent in larger squids. Compared to the 1990–1992 dataset, clupeid fish were less important in the diet of squid in 2006–2007, while the importance of gobies increased, and the size of gobies eaten also increased. The trend in gadoids differed according to the index used: their frequency of occurrence was considerably higher in 2006–2007 than in 1990–1992, but their numerical importance was slightly lower. In general, results provided little evidence that changes in the diet of L. forbesii correspond with changes in fish abundance, at least at the scales at which these are measured.     

Modification of the diel vertical migration of Bythotrephes longimanus by the cold-water planktivore, Coregonus artedi

1. The weak diel vertical migration observed in the large cladoceran Bythotrephes longimanus seems contradictory to the predator‐avoidance hypothesis that predicts large zooplankton should have long migration amplitudes. However, cold‐water planktivores, especially Coregonus spp., are a main source of mortality for Bythotrephes and hence a deeper migration would result in a greater overlap with these hypolimnetic planktivores. We hypothesized that Coregonus artedi (cisco) modifies the normal vertical migration pattern of Bythotrephes, such that the latter stays higher in the water column during the day and thus migrates less extremely at night. 2. The vertical distribution of Bythotrephes during the day was determined from single visits to six lakes in Ontario, Canada, all of which contain warm‐water, epilimnetic planktivores but differing in whether they contain cisco. One lake of each fish type was sampled day and night every 2–3 weeks over the ice‐free season to examine daytime depths and migration amplitude. 3. The vertical migration of Bythotrephes differed in the presence and absence of cisco. In the lakes with cisco, there were significantly fewer Bythotrephes in the hypolimnion and they were higher in the water column during the day. Migration amplitude was smaller in the cisco than in the non‐cisco lake. These observations were not attributable to differences in physical factors, and, although not conclusively attributable to cisco, are consistent with an effect of cisco. 4. We suggest that diurnal depth selection by Bythotrephes in lakes containing cisco is a trade‐off between the risk of predation by warm‐ versus cold‐water predators, balanced by the benefits of increased temperature and feeding rates near the surface. Even in lakes without cisco, however, the vertical migration of Bythotrephes was less than expected, suggesting that diurnal depth selection is a balance between the risk from warm‐water planktivores and access to sufficient light to feed effectively.     

Food limitation impacts life history of the predatory cladoceran Bythotrephes longimanus, an invader to North America

North American invasions of the predatory cladoceran Bythotrephes longimanus have resulted in declines in native zooplankton abundance, species richness, and diversity. In the field, population maxima of Bythotrephes are positively correlated to those of their zooplankton prey. To test the hypothesis that increased prey availability enhances Bythotrephes fitness, we reared Bythotrephes in the laboratory on three mixed-species prey densities (equivalent to 15, 30, and 45 prey organisms day^?1; designated “low,” “medium,” and “high” food treatments, respectively) over 22 days at 21°C. Bythotrephes consumed the daily equivalent of 9, 14, and 22 prey organisms at the low, medium, and high food densities. Smaller, slower prey were most often selected. Indeed, with increasing prey density, Bythotrephes’ predation rates increased, resulting in significantly higher population growth rates, net reproductive rates, growth, and first brood clutch and offspring sizes; significantly faster generation times; and shorter maximum life spans. We propose that the positive relationship between Bythotrephes population maxima and prey seen in the field is largely due to increased predation rates by Bythotrephes when prey abundance is high and the fitness benefits that ensue. Our findings may be useful for Bythotrephes risk and impact assessments.     

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.     

Shifting invertebrate zooplanktivores: watershed-level replacement of the native <Emphasis Type="Italic">Leptodora</Emphasis> by the non-indigenous <Emphasis Type="Italic">Bythotrephes</Emphasis> in Canadian Shield lakes

The abundance of the native, pelagic macroinvertebrate predator, Leptodora kindtii, is negatively correlated with the abundance of a new invasive competitor, Bythotrephes longimanus, in a small number of Canadian Shield lakes. However, we do not yet know if Bythotrephes is replacing Leptodora on a regional scale. We determined the distribution of both species in 166 lakes in the District of Muskoka, south-central Ontario, Canada—the watershed with the longest history and largest prevalence of Bythotrephes invasions in North America. The frequency of occurrence of Leptodora was substantially reduced (twofold) in the presence of Bythotrephes. We argue that Bythotrephes is responsible for this dramatic reduction in the frequency of occurrence of Leptodora. Lakes in which both species co-occurred could not be distinguished from invaded lakes without Leptodora, suggesting a pattern of species replacement at a watershed level. We believe this is the first account of the widespread replacement of a native, pelagic macroinvertebrate predator by Bythotrephes in North America, and it does not bode well for Leptodora given the rapid, ongoing spread of Bythotrephes.     

Seasonal size variation in the predatory cladoceran Bythotrephes cederstroemii in Lake Michigan

• 1 Dry weight, body length and spine length were measured for the exotic cladoceran Bythotrephes cederstroemii collected from offshore and inshore stations in southeastern Lake Michigan. Average dry weight of each developmental stage exhibited seasonal variation by a factor of more than 5.
• 2 Mean dry weight of Bythotrephes was closely correlated with water temperature. Contrary to the inverse relationship between water temperature and body size frequently observed for other invertebrates, the dry weight of Bythotrephes increased at higher ambient temperatures.
• 3 No significant correlation was observed between abundances of major zooplankton taxa and the dry weight of Bythotrephes. An indirect effect of temperature on prey consumption may cause seasonal variation in dry weight of Bythotrephes in Lake Michigan.
• 4 Distances between adjacent pairs of barbs, added to the caudal spine with each moult, are significantly shorter in Bythotrephes which produce resting eggs. Less material investment in the exoskeleton of sexually reproducing females was observed in favour of growth and reproduction.

     

Invasive Predator, <Emphasis Type="Italic">Bythotrephes</Emphasis>, has Varied Effects on Ecosystem Function in Freshwater Lakes

Bythotrephes longimanus is an invertebrate predator that has invaded the North American Great Lakes and a number of inland lakes, where it preys on crustacean zooplankton. We examined the effect of Bythotrephes on two measures of ecosystem function during a four-month observational study of freshwater lakes on the boreal shield. Bythotrephes-invaded lakes had significantly lower epilimnetic zooplankton abundance and production compared to reference lakes. On average, Bythotrephes consumed 34% of zooplankton production when it was present in lakes. There was some evidence of changes in the timing of zooplankton production, as well as shifts to cooler, less productive habitats, which may lessen the overall effect of the invader on the transfer of energy to higher trophic levels. We experimentally demonstrated a weak trophic cascade where invader predation reduced zooplankton biomass, and subsequently increased phytoplankton growth. However, the response was small in magnitude and not biologically relevant at the whole lake-scale. The most conspicuous effect of Bythotrephes that we measured was a diversion of energy away from native predators at higher trophic levels. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Selective predation on an introduced zooplankter ( Bythotrephes cederstroemi ) by lake herring ( Coregonus artedii ) in Harp Lake,Ontario

1. Bythotrephes cederstroemi (Crustacea: Onychopoda: Cercopagidae) invaded Harp Lake, Ontario in 1993, since when the zooplankton community has shifted from dominance of small-bodied to large-bodied species. During 1995 diets of adult lake herring (Coregonus artedii), Harp Lake’s primary planktivorous fish, were examined to determine the extent to which this conspicuous zooplankter has become integrated into the lake’s foodweb and whether fish condition has been affected in consequence. 2. Bythotrephes and Daphnia galeata mendotae were strongly preferred prey, whilst Holopedium gibberum and calanoid and cyclopoid copepods were negatively selected by lake herring. Predation on Bythotrephes and Holopedium was not size-selective, although D. galeata mendotae and calanoid copepods (Leptodiaptomus minutus and L. sicilis) consumed by herring were significantly larger than co-occurring conspecifics in the lake. 3. Caudal spines of Bythotrephes may form boluses in lake herring stomachs. However, the number of caudal spines in fish digestive tracts did not differ significantly from the number of Bythotrephes’ mandible pairs, indicating that the former were not differentially retained. 4. Lake herring weight-at-length relationships in lakes in Muskoka, Ontario, invaded by Bythotrephes did not differ from those in adjacent non-invaded lakes, indicating that Bythotrephes invasion of lakes apparently has not affected condition of lake herring.     

Planktivory in the changing Lake Huron zooplankton community: Bythotrephes consumption exceeds that of Mysis and fish

1. Oligotrophic lakes are generally dominated by calanoid copepods because of their competitive advantage over cladocerans at low prey densities. Planktivory also can alter zooplankton community structure. We sought to understand the role of planktivory in driving recent changes to the zooplankton community of Lake Huron, a large oligotrophic lake on the border of Canada and the United States. We tested the hypothesis that excessive predation by fish (rainbow smelt Osmerus mordax, bloater Coregonus hoyi) and invertebrates (Mysis relicta, Bythotrephes longimanus) had driven observed declines in cladoceran and cyclopoid copepod biomass between 2002 and 2007. 2. We used a field sampling and bioenergetics modelling approach to generate estimates of daily consumption by planktivores at two 91‐m depth sites in northern Lake Huron, U.S.A., for each month, May–October 2007. Daily consumption was compared to daily zooplankton production. 3. Bythotrephes was the dominant planktivore and estimated to have eaten 78% of all zooplankton consumed. Bythotrephes consumption exceeded total zooplankton production between July and October. Mysis consumed 19% of all the zooplankton consumed and exceeded zooplankton production in October. Consumption by fish was relatively unimportant – eating only 3% of all zooplankton consumed. 4. Because Bythotrephes was so important, we explored other consumption estimation methods that predict lower Bythotrephes consumption. Under this scenario, Mysis was the most important planktivore, and Bythotrephes consumption exceeded zooplankton production only in August. 5. Our results provide no support for the hypothesis that excessive fish consumption directly contributed to the decline of cladocerans and cyclopoid copepods in Lake Huron. Rather, they highlight the importance of invertebrate planktivores in structuring zooplankton communities, especially for those foods webs that have both Bythotrephes and Mysis. Together, these species occupy the epi‐, meta‐ and hypolimnion, leaving limited refuge for zooplankton prey.     

Biotic resistance of impact: a native predator (<Emphasis Type="Italic">Chaoborus</Emphasis>) influences the impact of an invasive predator (<Emphasis Type="Italic">Bythotrephes</Emphasis>) in temperate lakes

Introduced predators have caused some of the largest documented impacts of non-native species. Interactions among predators can have complex effects, leading to both synergistic and antagonistic outcomes. Complex interactions with native predators could play an important role in mediating the impact of non-native predators. We explore the role of the native predator context on the effect of the introduced predatory cladoceran Bythotrephes longimanus. While post-invasion impacts have been well described, studies have largely ignored the role of native predators. We used a field mesocosm experiment to determine whether Bythotrephes’ impact on prey communities is influenced by the presence of the ubiquitous native predatory insect larvae Chaoborus. The two predators exhibited niche complementarity as no change in total zooplankton prey abundance was detected across predator treatments. Rather, copepod abundances increased with decreasing abundances of Chaoborus, while cladocerans decreased with increasing abundances of Bythotrephes. Thus, the replacement of Chaoborus with Bythotrephes led to changes in the overall community structure of the zooplankton prey, but had little effect on prey total abundance. More interestingly, we found evidence of biotic resistance of impact, that is, the impact of Bythotrephes on the cladoceran community was altered when the two predators co-occurred. Specifically, the predation effect of Bythotrephes was more restricted to the shallower regions of the water column in the presence of Chaoborus, leading to a reduced impact on deeper dwelling prey taxa. Overall, our results demonstrate that the native predator context is important when trying to understand the effect of non-native predators and that variation in native predator abundances and assemblages could explain variation in impact across invaded habitats.     

Investigations on fecundity of Bythotrephes longimanus in Lake Lucerne (Switzerland) and on Niche Segregation of Leptodora kindti and Bythotrephes longimanus in Swiss lakes

In Lake Lucerne, Switzerland, the predaceous cladocerans Leptodora kindti and Bythotrephes longimanus segregate along spatial and temporal dimensions. In spring (April–May/June), Bythotrephes longimanus occurs below 0–20 m, while Leptodora is absent. In summer and early autumn (July–September/October), when Leptodora dominates during daytime in the 0–20 m depth, Bythotrephes longimanus also lives in deeper zones. Food competition and fish predation pressure may be the cause of differences in ecology of Leptodora and Bythotrephes acquired during evolution. Due to its transparency and tolerance of higher temperature, Leptodora could avoid fish predation and, therefore, competes with Bythotrephes longimanus successfully. In addition, the differences between the two species may account for the spatial and temporal niche segregation in oligotrophic Swiss Lakes. But spatial niche segregation is less important in mesotrophic lakes with high prey density than in oligotrophic lakes with low prey density. In small, eutrophic lakes importance of temporal niche segregation also decreases, and Bythotrephes is seldom or not present. The preference of Bythotrephes to live in deeper water to avoid fish predation during summer may be the cause of its difficulties to establish itself in small and eutrophic lakes with high prey densities, where the hypolimnion is missing or anoxic.In the spring, Bythotrephes exhibits r-strategy (smaller body size and a higher fecundity), the female is already fertile after the first molt. In the summer, a K-strategy prevails (larger body length and lower fecundity than in the spring), and female Bythotrephes are fertile only after the second molt. Shortage of prey (biomass of Bosmina and Daphniadecreased after June especially in the surface layers) and the maximum fish predation pressure in summer may change the life strategy of Bythotrephes: while fecundity decreases from generation to generation, body length increases. Enhanced prey densities (e.g. during mesotrophic conditions in L. Lucerne) lead to larger individuals in summer and autumn.     

The invasive predator <Emphasis Type="Italic">Bythotrephes</Emphasis> induces changes in the vertical distribution of native copepods in Lake Michigan

Invasive predators can have large negative effects on native prey populations. The susceptibility of native prey to invasive predators may depend on their ability to respond behaviorally to the presence of these non-native predators. In a field survey conducted in Lake Michigan over several years, we found that high densities of the invasive predatory cladoceran Bythotrephes were correlated with lower vertical distributions of some species and age classes of native copepods; moving from inhabiting primarily the epiliminion at low Bythotrephes density to primarily the hypolimnion at high Bythotrephes density. Five groups showed this pattern; diaptomid copepodites, adult cyclopoids, Diacyclops thomasi, and the adult diaptomids Leptodiaptomus ashlandi and L. minutus. In contrast, Bythotrephes density was not correlated with the vertical distribution of copepod nauplii and adult L. sicilis. Laboratory experiments suggest that the changes in the vertical distribution in the field at high Bythothrephes are due to an inducible, plastic response to predation threat from Bythotrephes signaled by water-borne cues. Species that were lower in the field at high Bythotrephes densities responded behaviorally to water-borne cues from Bythotrephes by moving to lower levels of experimental water columns. These species included D. thomasi and L. minutus, with L. ashlandi displaying a non-significant trend in the same direction. In contrast, L. sicilis, which was not correlated with Bythotrephes density in the field, was unaffected by the water-borne cues. Differences in vertical distribution shifts among these native copepod species and life-history stages are likely due to species-specific differences in spatial overlap with Bythotrephes and their relative ability to migrate large distances or employ alternative avoidance strategies. The varied responses exhibited among the copepod groups likely alter their interactions with each other, their resources and other predators, thus revealing the complex effects Bythotrephes can have on invaded communities.