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.     

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.     

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.     

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.     

Increased abundance of the non-indigenous zooplanktivore, <Emphasis Type="Italic">Bythotrephes longimanus</Emphasis>, is strongly correlated with greater spring prey availability in Canadian Shield lakes

The non-indigenous zooplanktivore, Bythotrephes longimanus, is a large Palaearctic cladoceran that is spreading rapidly in the Great Lakes watershed in North America. As a voracious predator, Bythotrephes can reduce herbivorous cladoceran abundance and diversity; however, the variables that affect its abundance are not well understood. To determine what bottom-up factors are associated with the abundance and seasonal dynamics of established Bythotrephes populations, two Bythotrephes datasets from lakes in south-central Ontario, Canada, were analysed using multiple regression and multivariate analyses: a multi-lake dataset of nine lakes sampled in 2003 and a multi-year dataset of one of these lakes, Harp Lake, sampled from 1994–1998 and 2001–2004. Bottom-up variables tested were Secchi disk depth, epilimnetic temperature, cladoceran (prey) density, total phosphorus, dissolved organic carbon and Chlorophyll a, as well as maximum depth for the multi-lake dataset. In both analyses and datasets, springtime abundance of herbivorous cladocerans was consistently found to be a significant factor associated with Bythotrephes (June–September) abundance; Bythotrephes annual abundance was significantly and positively associated with mean May and June prey abundance, along with mean Secchi disk depth for the multi-lake dataset, and groups of lakes or years with similar Bythotrephes seasonal abundance patterns were predicted by June prey abundance. Additionally, prey availability was the dominant contributor towards changes in weekly Bythotrephes birth rates calculated for two of the study lakes. Our study suggests that prey availability influences Bythotrephes abundance, which provides evidence that Bythotrephes establishment success is affected by the abundance of its prey.     

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.     

Tadpole shrimp structure macroinvertebrate communities in playa lake microcosms

Laboratory microcosms were used to assess whether tadpole shrimp, Triops sp., affect community structure of other native macroinvertebrates in playa lakes of the Southern High Plains of Texas. Removal of tadpole shrimp shortly after hatching reduced abundances of many taxa, and decreased subsequent taxonomic richness and diversity. For many invertebrates, the presence of tadpole shrimp in low numbers had a positive effect on mean abundance. Direct effects of tadpole shrimp include the reduction of prey species abundance, which in turn may alter biotic interactions among other taxa. Indirect effects include physical modification of the environment during foraging through surface sediments. Results suggest that tadpole shrimp may be a key species controlling structure of macroinvertebrate communities in playa lakes.     

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.     

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.     

The relationship between fecundity and adult body weight in Homeotherms

Summary Bythotrephes cederstroemii Schoedler, a predatory freshwater zooplankter (Crustacea: Cladocera), was first found in the Laurentian Great Lakes in December 1984. The first individuals were from Lake Huron, followed in 1985 with records from Lakes Erie and Ontario. By late August, 1986 the species had spread to southern Lake Michigan (43°N). Bythotrephes has not previously been reported from North America, but has been restricted to a northern and central Palearctic distribution. Its dramatic and widespread rise in abundance in Lake Michigan was greatest in offshore regions. Bythotrephes appears to be invading aggressively, but avoiding habitats presently occupied by glacio-marine relict species that became established in deep oligotrophic North American lakes after the Wisconsin glaciation. Because it is a voracious predator its invasion may lead to alterations in the native zooplankton fauna of the Great Lakes. It offers the chance to study how invading plankton species join an existing community. Judging from its persistence and success in deep European lakes, Bythotrephes may now become a permanent member of zooplankton communities in the Nearctic.     

Predator–prey interactions of Procambarus clarkii with aquatic macroinvertebrates in single and multiple prey systems

Understanding the interspecific interactions of Procambarus clarkii with other aquatic macroinvertebrates will help to unveil the mechanisms and processes underlying biological invasiveness. The purpose of this study was to investigate predator–prey interactions of two ontogenic phases of P. clarkii with native and exotic species of aquatic macroinvertebrates at a single and multiple prey level. We performed laboratory experiments to determine the consumption and the behavioral responses of Chironomus riparius, Physa acuta and Corbicula fluminea to P. clarkii. The presence of P. clarkii significantly affected the abundance of C. riparius and P. acuta, but not of C. fluminea whether prey species were provided singly or simultaneously. The consumption of C. riparius by P. clarkii was higher than P. acuta for both crayfish sizes and situations (single/multiple prey systems) and C. fluminea was never consumed. Physa acuta was the only species that exhibited an anti-predator behavior to P. clarkii. Our results show that P. clarkii can have strong consumptive and trait effects on aquatic macroinvertebrate prey at a single and multiple prey level, resulting in differential impacts on different prey species. This study clarifies some aspects of the predator–prey interactions between P. clarkii and native as well as other exotic macroinvertebrate species that have invaded freshwater biocenosis worldwide.     

<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.     

Response of benthic macroinvertebrates to whole-lake, non-native fish treatments in mid-elevation lakes of the Trinity Alps, California

Introduced fish reduce the abundance and diversity of native aquatic fauna, but the effect can be reduced in complex habitats. We manipulated fish populations in forested mountain lakes to determine whether or not fish affected benthic macroinvertebrate composition across lakes with differing habitat complexity. We compared abundance, biomass, body-length, and community structure of benthic macroinvertebrates from 16 lakes with three treatments (fish stocked, suspended stocking, fish removed) and unstocked fishless “controls”. Over 4 years, we assessed the relative importance of fish and environmental variables influencing the composition of benthic macroinvertebrates. Control lakes had the greatest overall abundance of macroinvertebrates when chironomid midges were excluded. Abundances of insects in the clinger/swimmer functional group and caddisflies were greatest in the control lakes but were primarily influenced by habitat variables including the availability of aquatic vegetation and wood. Total biomass and mean body length of macroinvertebrates were not affected by treatment. Taxon richness of macroinvertebrates was about 40% greater in the control lakes compared to the treatment lakes but did not differ among treatments. Our results suggest that fish reduce susceptible macroinvertebrate richness and abundances, but that changes associated with alterations of fish composition are confounded by other factors in complex lake habitats.     

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.     

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.     

The introduced signal crayfish and native noble crayfish have different effects on sublittoral macroinvertebrate assemblages in boreal lakes

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Mensurative approach to examine potential interactions between age-0 yellow perch (Perca flavescens) and bluegill (Lepomis macrochirus)

Bluegill (Lepomis macrochirus) and yellow perch (Perca flavescens) populations are often sympatric in the Great Plains region of the U.S.A. and portions of Canada; however, very little attention has been given to potential interactions between these species for available resources, especially during the early life stages. Relationships between age-0 bluegill and yellow perch growth and relative abundance were explored across multiple lakes and years within the Nebraska Sandhill region, USA. In addition, four habitat patch types (open water, Phragmites spp., Typha spp., Scirpus spp.) were sampled for age-0 bluegill and yellow perch, and food habits were examined for each species during August, September, and October of 2009 in one of these lakes. Age-0 yellow perch growth was negatively related to age-0 bluegill relative abundance across a spatiotemporal scale. Age-0 bluegill and yellow perch exhibited similar habitat use (moderate–high overlap), but generally consumed different important and dominant prey taxa (bluegill consumed both macroinvertebrates—56?% and zooplankton—44?%, while yellow perch consumed more zooplankton—66?%), which resulted in low overall diet overlap between species. Previous research indicates that age-0 yellow perch diet ontogeny often results in feeding predominately on macroinvertebrates and positively selecting them (and avoiding zooplankton prey) at sizes observed in our study. Therefore, yellow perch growth rates may be compromised by the presence of bluegill because of the need to consume less energetically profitable prey items such as zooplankton.     

Distribution and growth of benthic macroinvertebrates among different patch types of the littoral zones of two arctic lakes

1. To evaluate the effect of habitat patch heterogeneity on abundance and growth of macroinvertebrates in arctic lakes, macroinvertebrate abundance, individual biomass, and potential food resources were studied in three patch types in two arctic lakes on the Alaskan North Slope near the Toolik Lake Field Station. An experiment was conducted to determine which sediment patch type supported higher growth rates for Chironomus sp., a commonly occurring macroinvertebrate. 2. Potential organic matter (OM) resources were significantly higher in both rock and macrophyte patches than in open‐mud patches. Total macroinvertebrate densities in both lakes were highest in rock patches, intermediate in macrophytes and lowest in open‐mud. The open‐mud patches also had lower species richness compared with other patch types. Additionally, individual biomass for one clam species and two chironomid species was significantly greater in rock patches than in open‐mud. 3. In a laboratory experiment, Chironomus showed two to three times greater mass increase in sediments from macrophyte and rock patches than from open‐mud patches. Rock and macrophyte experimental sediments had at least 1.5 × the percentage OM as open‐mud sediments. 4. Chlorophyll a appeared to be the best predictor for invertebrate abundances across all patch types measured, whereas OM content appeared to be the variable most closely associated with Chironomus growth. 5. Our results combined with previous studies show that the relationships between macroinvertebrate community structure, individual growth, and habitat heterogeneity are complex, reflecting the interaction of multiple resources, and biotic interactions, such as the presence or absence of a selective vertebrate predator (lake trout, Salvelinus namaycush).     

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.