Do non-native invasive fish support elevated lamprey populations?

Summary
1: Managing populations of predators and their prey to achieve conservation or resource management goals is usually technically challenging and frequently socially controversial. This is true even in the simplest ecosystems but can be made much worse when predator–prey relationships are in?uenced by complex interactions, such as biological invasions, population trends or animal movements.
2: Lough Neagh in Northern Ireland is a European stronghold for pollan Coregonus autumnalis, a coregonine ?sh and for river lampreyLampetra ?uviatilis, which feeds parasitically as an adult. Both species are of high conservation importance. Lampreys are known to consume pollan but detailed knowledge of their interactions is scant. While pollan is well known to be a landlocked species in Ireland, the life cycle of normally anadromous river lamprey in Lough Neagh has been unclear. The Lough is also a highly perturbed ecosystem, supporting several invasive, non-native ?sh species that have the potential to in?uence lamprey–pollan interactions.
3: We applied stable isotope techniques to resolve both the movement patterns of lamprey and trophic interactions in this complex community. Recognizing that stable isotope studies are often hampered by high-levels of variability and uncertainty in the systems of interest, we employed novel Bayesian mixing models, which incorporate variability and uncertainty.
4: Stable isotope analyses identi?ed troutSalmo trutta and non-native breamAbramis brama as the main items in lamprey diet. Pollan only represented a major food source for lamprey between May and July.
5: Stable isotope ratios of carbon in tissues from 71 adult lamprey showed no evidence of marine carbon sources, strongly suggesting that Lough Neagh is host to a highly unusual, nonanadromous freshwater population. This ?nding marks out the Lough’s lamprey population as of particular scienti?c interest and enhances the conservation signi?cance of this feature of the Lough.
6: Synthesis and applications.Our Bayesian isotopic mixing models illustrate an unusual pattern of animal movement, enhancing conservation interest in an already threatened population. We have also revealed a complex relationship between lamprey and their food species that is suggestive of hyperpredation, whereby non-native species may sustain high lamprey populations that may in turn be detrimental to native pollan.Long-term conservation of lamprey and pollan in this system is likely to require management intervention, but in light of this exceptional complexity, no simple management options are currently supported. Conservation plans will require better characterization ofpopulation-level interactions and simulation modelling of interventions. More generally, our study demonstrates the importance of considering a full range of possible trophic interactions, particularly in complex ecosystems, and highlights Bayesian isotopic mixing models as powerful tools in resolving trophic relationships.
Key-words: Bayesian, conservation dilemma, Coregonus autumnalis, hyperpredation, Lampetra ?uviatilis, pollan, potamodromous, River lamprey, stable isotope analysis in R, stable isotope

Temporal changes in the distribution of native and introduced freshwater amphipods in Lough Neagh, Northern Ireland

Studies of invasion scenarios over long time periods are important to refine explanations and predictions of invasion success and impact. We used data from surveys in 1958 and 1999 of the macroinvertebrates of Lough Neagh, Northern Ireland, to assess changes in the distribution of native and introduced amphipods in relation to the wider assemblage. In 1958, the invader G. tigrinus dominated the shoreline fauna, with the native G. d. celticus present in very low numbers, whereas in 1999 the reverse was evident. In both surveys, G. tigrinus was the only amphipod present in the mid-Lough. G. tigrinus thus seems to have become established within L. Neagh, perhaps overshot and then senesced, with the native species re-establishing on the shoreline, with the invader mostly restricted to the deep mid-Lough. The non-amphipod macroinvertebrate assemblage was similar between the two surveys, in terms of Bray-Curtis community similarity, assemblage diversity, dominance and the taxa based ASPT water quality index. However, the mean density of macroinvertebrates (all taxa combined) was lower in 1999 compared to 1958, largely accounted for by a decline in oligochaete numbers. Since Gammarus species may be predators of other macroinvertebrates and influence their distribution and abundance, we investigated this trophic link in staged laboratory encounters. Both G. tigrinus and G. d. celticus preyed on isopods, alderflies, mayflies, chironomids and mysids, however, the native G. d. celticus had a significantly greater predatory impact on isopods and chironomids than did the invader G. tigrinus. While we cannot definitively ascribe cause and effect in the present scenario, we discuss how replacement of one amphipod species by another may have impacts on the wider macroinvertebrate assemblage.

Predatory impact of the freshwater invader Dikerogammarus villosus (Crustacea : Amphipoda)

To assess the increasing threats to aquatic ecosystems from invasive species, we need to elucidate the mechanisms of impacts of current and predicted future invaders. Dikerogammarus villosus, a Ponto-Caspian amphipod crustacean, is invading throughout Europe and predicted to invade the North American Great Lakes. European field studies show that populations of macroinvertebrates decline after D. villosus invasion. The mechanism of such impacts has not been addressed empirically; however, D. villosus is known to prey upon and replace other amphipods. Therefore, in this study, we used microcosm and mesocosm laboratory experiments, with both single and mixed prey species scenarios, to assess any predatory impact of D. villosus on a range of macro invertebrate taxa, trophic groups, and body sizes. Dikerogammarus villosus predatory behaviour included shredding of prey and infliction of

Predator-prey interactions between brown trout Salmo trutta and native and introduced amphipods; their implications for fish diets

An attempt to improve the food base for brown trout Salmo trutta in Northern Ireland was made in 1958.59 by deliberately introducing English Gammarus pulex into several Irish rivers. In addition. another amphipod Crangonyx pseudogracilis, was later accidently introduced into II ish waters. Our study represents the first attempt to examine the trophic interactions between a native fish predator (S. trutta) and an array of these native (Gammarus duebeni celticus) and introduced (G. pulex and C. pseudogracilis) amphipods. Feeding experiments, involving young brown trout predators and ampiphod prey, revealed that the fish actively selected C. pseudogracilis relative to two alternative Gammarus prey species. Although the trout encountered the Gammarus species more than C. pseudogracilis, they were eaten less than Crangonyx. Difficulties in handling and ingestion of Gammarus by trout may be a. key component of the preference fbr the smaller, more easily handled Crangonyx. The microdistribution of the species was altered by the fish, due to predation being greater in particular microhabitats, Our study showed that the introduction of the herbivorous C. pseudogracilis into Irish freshwaters may represent a useful addition to fish diets. particularly for small and/or juvenile fish. The reprecussions of the deliberate introduction of G. pulex are less clear. It may improve feeding for fish. but only if it can coexist with indigenous macroinvertebrates and thus ultimately improve the range and quantity of possible food items in predator diets. Alternatively, being highly predatory towards other macroinvertebrates including G. d. celticus and C. pseudogracilis. G. pulex may be deleterious to the diversity of the resident benthic community and hence reduce the diversity of prey available to fish predators.

Direct and indirect effects of species displacements; an invading freshwater amphipod can disrupt leaf litter processing and shredder efficiency

Invasive species may threaten the fundamental role played by native macroinvertebrate shredders in determining energy flow and the trophic dynamics of freshwater ecosystems. Functionally, amphipods have long been regarded as mainly shredders, but they are increasingly recognized as major predators of other macroinvertebrate taxa. Furthermore, intraguild predation (IGP) between native and invasive amphipods underlies many species displacements. We used laboratory mesocosms to investigate what might happen to shredders and leaf-litter processing in water bodies invaded by the highly predatory Ponto-Caspian amphipod Dikerogammarus villosus, which is spreading rapidly throughout Europe and may soon invade the North American Great Lakes. The leaf-shredding efficiency of D. villosus was significantly lower than that of 3 Gammarus species (2 native and 1 invasive) that D. villosus has either already displaced or may be currently displacing in The Netherlands. In addition, D. villosus was a major predator of all of these native and invasive amphipod shredders and of a common isopod shredder Asellus aquaticus. Leaf processing in Gammarus and Asellus mesocosms declined rapidly in the presence of D. villosus and ceased altogether within 4 d because by then, all potential shredders had been killed and consumed. Furthermore, the shredding efficiency of surviving amphipods and isopods declined significantly within 2 d of the release of D. villosus, a result indicating that predator-avoidance behavior may override leaf processing. We discuss the implications of these direct and indirect effects of D. villosus invasions and species displacements on community structure and litter processing in aquatic ecosystems. © 2011 The North American Benthological Society.


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Rapid sympatric ecological differentiation of crater lake cichlid fishes within historic times

Background: After a volcano erupts, a lake may form in the cooled crater and become an isolated aquatic ecosystem. This makes fishes in crater lakes informative for understanding sympatric evolution and ecological diversification in barren environments. From a geological and limnological perspective, such research offers insight about the process of crater lake ecosystem establishment and speciation. In the present study we use genetic and coalescence approaches to infer the colonization history of Midas cichlid fishes (Amphilophus cf. citrinellus) that inhabit a very young crater lake in Nicaragua-the ca. 1800 year-old Lake Apoyeque. This lake holds two sympatric, endemic morphs of Midas cichlid: one with large, hypertrophied lips (~20% of the total population) and another with thin lips. Here we test the associated ecological, morphological and genetic diversification of these two morphs and their potential to represent incipient speciation.
Results: Gene coalescence analyses [11 microsatellite loci and mitochondrial DNA (mtDNA) sequences] suggest that crater lake Apoyeque was colonized in a single event from the large neighbouring great lake Managua only about 100 years ago. This founding in historic times is also reflected in the extremely low nuclear and mitochondrial genetic diversity in Apoyeque. We found that sympatric adult thin- and thick-lipped fishes occupy distinct ecological trophic niches. Diet, body shape, head width, pharyngeal jaw size and shape and stable isotope values all differ significantly between the two lip-morphs. The eco-morphological features pharyngeal jaw shape, body shape, stomach contents and stable isotopes (d15N) all show a bimodal distribution of traits, which is compatible with the expectations of an initial stage of ecological speciation under disruptive selection. Genetic differentiation between the thin- and thick-lipped population is weak at mtDNA sequence (FST = 0.018) and absent at nuclear microsatellite loci (FST < 0.001).
Conclusions: This study provides empirical evidence of eco-morphological differentiation occurring very quickly after the colonization of a new and vacant habitat. Exceptionally low levels of neutral genetic diversity and inference from coalescence indicates that the Midas cichlid population in Apoyeque is much younger (ca. 100 years or generations old) than the crater itself (ca. 1 800 years old). This suggests either that the crater remained empty for many hundreds of years after its formation or that remnant volcanic activity prevented the establishment of a stable fish population during the early life of the crater lake. Based on our findings of eco-morphological variation in the Apoyeque Midas cichlids, and known patterns of adaptation in Midas cichlids in general, we suggest that this population may be in a very early stage of speciation (incipient species), promoted by disruptive selection and ecological diversification.

delta C-13 and delta N-15 reveal significant differences in the coastal foodwebs of the seas surrounding Trinidad and Tobago

This study assessed nearshore, marine ecosystem function around Trinidad and Tobago (TT). The coastline of TT is highly complex, bordered by the Atlantic Ocean, the Caribbean Sea, the Gulf of Paria and the Columbus Channel, and subject to local terrestrial runoff and regional riverine inputs (e.g. the Orinoco and Amazon rivers). Coastal organisms can assimilate energy from allochthonous and autochthonous Sources, We assessed whether stable isotopes delta C-13 and delta N-15 Could be used to provide a rapid assessment of trophic interactions in primary consumers around the islands. Filter-feeding (bivalves and barnacles) and grazing organisms (gastropods and chitons) were collected from 40 marine sites during the wet season. The flesh of organisms was analysed for delta C-13 and delta N-15. Results indicate significant variation in primary consumers (by feeding guild and sampling zone). This variation was linked to different energy Sources being assimilated by consumers. Results suggest that offshore production is fuelling intertidal foodwebs; for example, a depleted delta C-13 signature in grazers from the Gulf of Paria, Columbus Channel and the Caribbean and Atlantic coastline of 9 Tobago indicates that carbon with an offshore origin (e.g. phytoplankton and dissolved organic matter) is more important than benthic or littoral algae (luring the wet season. Results also confirm findings from other studies indicating that much of the coastline is subject to Cultural eutrophication. This Study revealed that ecosystem function is spatially variable around the coastline of TT, This has clear implications for marine resource management, as a single management approach is unlikely to be successful at a national level.

Continuous variation in the pattern of marine v. freshwater foraging in brown trout Salmo trutta L. from Loch Lomond, Scotland

Carbon stable-isotope analysis showed that individual brown trout Salmo trutta in Loch Lomond adopted strategies intermediate to that of freshwater residency or anadromy, suggesting either repeated movement between freshwater and marine environments, or estuarine residency. Carbon stable-isotope (delta C-13) values from Loch Lomond brown trout muscle tissue ranged from those indicative of assimilation of purely freshwater-derived carbon to those reflecting significant utilization of marine-derived carbon. A single isotope, two-source mixing model indicated that, on average, marine C made a 33% contribution to the muscle tissue C of Loch Lomond brown trout. Nitrogen stable isotope, delta N-15, but not delta C-13 was correlated with fork length suggesting that larger fish were feeding at a higher trophic level but that marine feeding was not indicated by larger body size. These results are discussed with reference to migration patterns in other species. (c) 2008 The Authors Journal compilation (c) 2008 The Fisheries Society of the British Isles.

Biological influences on inter- and intraspecific isotopic variability among paired chondrostome fishes

This study aimed at examining resource partitioning both at the inter- and intraspecific levels between paired chondrostome fishes: Chondrostoma nasus, the nase, C. toxostoma, the sofie, and their hybrid. The study was performed in the south of France and concerned a main river (the Durance River) and a tributary (the Buech River). In these rivers, C. nasus was an introduced species, originating in central Europe, and C. toxostoma was an endemic congener, in the south of France. Stable isotope analysis was used to analyse trophic and spatial niches. Isotopic differences indicated that individuals from the three taxa (C. nasus, C. toxostoma and their hybrid) have different spatial origins. At the interspecific level, the different chondrostomes originating from the Buech River showed a high level of trophic niche overlap. At the intraspecific level, nase individuals originating from the different spatial origins showed a resource polymorphism; differences in morphology were associated with variation in behaviour and life history traits. Their coexistence was a likely outcome of resource polymorphism. This study provides an example of the importance of considering the link between intra- and interspecific interactions to gain an understanding of the mechanisms driving the coexistence of species-pairs. (C) 2010 Academie des sciences. Published by Elsevier Masson SAS. All rights reserved.

Biodiversity loss and ecosystem functioning: Distinguishing between number and identity of species

Given currently high rates of extinction, it is critical to be able to predict how ecosystems will respond to loss of species and consequent changes in community structure. Much previous research in this area has been based on terrestrial systems, using synthetically assembled communities. There has beer! much less research on inter-trophic effects in different systems, using in situ removal experiments. Problems with the design of early experiments have made it difficult to determine whether reductions in ecosystem functioning in low diversity treatments were due to the number of species present or merely to the reduced likelihood of including particular (

Simulated predator extinctions: Predator identity affects survival and recruitment of oysters

The rate of species loss is increasing at a global scale, and human-induced extinctions are biased toward predator species. We examined the effects of predator extinctions on a foundation species, the eastern oyster (Crassostrea virginica). We performed a factorial experiment manipulating the presence and abundance of three of the most common predatory crabs, the blue crab (Callinectes sapidus), stone crab (Menippe mercenaria), and mud crab (Panopeus herbstii) in estuaries in the eastern United States. We tested the effects of species richness and identity of predators on juvenile oyster survival, oyster recruitment, and organic matter content of sediment. We also manipulated the density of each of the predators and controlled for the loss of biomass of species by maintaining a constant mass of predators in one set of treatments and simultaneously using an additive design. This design allowed us to test the density dependence of our results and test for functional compensation by other species. The identity of predator species, but not richness, affected oyster populations. The loss of blue crabs, alone or in combination with either of the other species, affected the survival rate of juvenile oysters. Blue crabs and stone crabs both affected oyster recruitment and sediment organic matter negatively. Mud crabs at higher than ambient densities, however, could fulfill some of the functions of blue and stone crabs, suggesting a level of ecological redundancy. Importantly, the strong effects of blue crabs in all processes measured no longer occurred when individuals were present at higher-than-ambient densities. Their role as dominant predator is, therefore, dependent on their density within the system and the density of other species within their guild (e.g., mud crabs). Our findings support the hypothesis that the effects of species loss at higher trophic levels are determined by predator identity and are subject to complex intraguild interactions that are largely density dependent. Understanding the role of biodiversity in ecosystem functioning or addressing practical concerns, such as loss of predators owing to overharvesting, remains complicated because accurate predictions require detailed knowledge of the system and should be drawn from sound experimental evidence, not based on observations or generalized models.

Part-whole relations between food webs and the validity of local food-web descriptions

This work analyzes the relationship between large food webs describing potential feeding relations between species and smaller sub-webs thereof describing relations actually realized in local communities of various sizes. Special attention is given to the relationships between patterns of phylogenetic correlations encountered in large webs and sub-webs. Based on the current theory of food-web topology as implemented in the matching model, it is shown that food webs are scale invariant in the following sense: given a large web described by the model, a smaller, randomly sampled sub-web thereof is described by the model as well. A stochastic analysis of model steady states reveals that such a change in scale goes along with a re-normalization of model parameters. Explicit formulae for the renormalized parameters are derived. Thus, the topology of food webs at all scales follows the same patterns, and these can be revealed by data and models referring to the local scale alone. As a by-product of the theory, a fast algorithm is derived which yields sample food webs from the exact steady state of the matching model for a high-dimensional trophic niche space in finite time. (C) 2008 Elsevier B.V. All rights reserved.

The top-down mechanism for body-mass-abundance scaling

Scaling relationships between mean body masses and abundances of species in multitrophic communities continue to be a subject of intense research and debate. The top-down mechanism explored in this paper explains the frequently observed inverse linear relationship between body mass and abundance (i.e., constant biomass) in terms of a balancing of resource biomasses by behaviorally and evolutionarily adapting foragers, and the evolutionary response of resources to this foraging pressure. The mechanism is tested using an allometric, multitrophic community model with a complex food web structure. It is a statistical model describing the evolutionary and population dynamics of tens to hundreds of species in a uniform way. Particularities of the model are the detailed representation of the evolution and interaction of trophic traits to reproduce topological food web patterns, prey switching behavior modeled after experimental observations, and the evolutionary adaptation of attack rates. Model structure and design are discussed. For model states comparable to natural communities, we find that (1) the body-mass-abundance scaling does not depend on the allometric scaling exponent of physiological rates in the form expected from the energetic equivalence rule or other bottom-up theories; (2) the scaling exponent of abundance as a function of body mass is approximately -1, independent of the allometric exponent for physiological rates assumed; (3) removal of top-down control destroys this pattern, and energetic equivalence is recovered. We conclude that the top-down mechanism is active in the model, and that it is a viable alternative to bottom-up mechanisms for controlling body-mass-abundance relations in natural communities.

Soil resource supply influences faunal size-specific distributions in natural food webs

The large range of body-mass values of soil organisms provides a tool to assess the ecological organization of soil communities. The goal of this paper is to identify graphical and quantitative indicators of soil community composition and ecosystem functioning, and to illustrate their application to real soil food webs. The relationships between log-transformed mass and abundance of soil organisms in 20 Dutch meadows and heathlands were investigated. Using principles of allometry, maximal use can be made of ecological theory to build and explain food webs. The aggregate contribution of small invertebrates such as nematodes to the entire community is high under low soil phosphorus content and causes shifts in the mass-abundance relationships and in the trophic structures. We show for the first time that the average of the trophic link lengths is a reliable predictor for assessing soil fertility responses. Ordered trophic link pairs suggest a self-organizing structure of food webs according to resource availability and can predict environmental shifts in ecologically meaningful ways.

Universal power-law diet partitioning by marine fish and squid with surprising stability–diversity implications

A central question in community ecology is how the number of trophic links relates to community species richness. For simple dynamical food-web models, link density (the ratio of links to species) is bounded from above as the number of species increases; but empirical data suggest that it increases without bounds. We found a new empirical upper bound on link density in large marine communities with emphasis on fish and squid, using novel methods that avoid known sources of bias in traditional approaches. Bounds are expressed in terms of the diet-partitioning function (DPF): the average number of resources contributing more than a fraction f to a consumer's diet, as a function of f. All observed DPF follow a functional form closely related to a power law, with power-law exponents indepen- dent of species richness at the measurement accuracy. Results imply universal upper bounds on link density across the oceans. However, the inherently scale-free nature of power-law diet partitioning suggests that the DPF itself is a better defined characterization of network structure than link density.