Shift in predation regime mediates diversification of foraging behaviour in a dragonfly genus

1. Behavioural adaptations to avoid and evade predators are common. Many studies have investigated population divergence in response to changes in predation regime within species, but studies exploring interspecific patterns are scant. Studies on interspecific divergence can infer common outcomes from evolutionary processes and highlight the role of environmental constraints in shaping species traits. 2. Species of the dragonfly genus Leucorrhinia underwent well‐studied shifts from habitats being dominated by predatory fish (fish lakes) to habitat being dominated by predatory invertebrates (dragonfly lakes). This change in top predators resulted in a set of adaptive trait modifications in response to the different hunting styles of both predator types: whereas predatory fish actively search and pursue prey, invertebrate predator follow a sit‐and‐wait strategy, not pursuing prey. 3. Here it is shown that the habitat shift‐related change in selection regime on larval Leucorrhinia caused species in dragonfly lakes to evolve increased larval foraging and activity, and results suggest that they lost the ability to recognise predatory fish. 4. The results of the present study highlight the impact of predators on behavioural trait diversification with habitat‐specific predation regimes selecting for distinct behavioural expression.     

Predators shape distribution and promote diversification of morphological defenses in <Emphasis Type="Italic">Leucorrhinia</Emphasis>, Odonata

Predators strongly influence species assemblages and shape morphological defenses of prey. Interestingly, adaptations that constitute effective defenses against one type of predator may render the prey susceptible to other types of predators. Hence, prey may evolve different strategies to escape predation, which may facilitate adaptive radiation of prey organisms. Larvae of different species in the dragonfly genus Leucorrhinia have various morphological defenses. We studied the distribution of these larvae in relation to the presence of predatory fish. In addition, we examined the variation in morphological defenses within species with respect to the occurrence of fish. We found that well-defended species, those with more and longer spines, were more closely associated with habitats inhabited by predatory fish and that species with weakly developed morphological defenses were more abundant in habitats without fish. The species predominantly connected to lakes with or without fish, respectively, were not restricted to a single clade in the phylogeny of the genus. Our data is suggestive of phenotypic plasticity in morphological defense in three of the studied species since these species showed longer spines in lakes with fish. We suggest that adaptive phenotypic plasticity may have broadened the range of habitats accessible to Leucorrhinia. It may have facilitated colonization of new habitats with different types of predators, and ultimately, speciation through adaptive radiation.     

Food partitioning by lake-dwelling triclads and glossiphoniid leeches: field and laboratory experiments

The triclads Polycelis tenuis and Dugesia polychroa and the glossiphoniid leeches Glossiphonia complanata and Helobdella stagnalis are abundant on the stony shores of productive British lakes. All species are food limited and there is considerable overlap in the diets of these triclads and leeches. This paper investigates interactions between the two groups using field and laboratory experiments to try to identify the mechanism of their co-existence. Triclad and leech numbers were manipulated inside experimental enclosures, mathced by controls, erected on the stony shore of an eutrophic English lake. Increasing the numbers of P. tenuis and D. polychroa prior to the reproductive season in spring resulted in a significant decrease in the numbers and body size of G. complanata and H. stagnalis compared with control populations in the summer months, and vice versa. However, increases and decreases were temporary with a readjustment of numbers and body size to control levels in the autumn after reproduction had ceased. It is suggested that increasing the numbers of either group elevated the severity of both intra- and interspecific competition for food. The condition of prey may, in part, determine the strength of competition, and this was examined in laboratory experiments in which different densities and ratios of P. tenuis and H. stagnalis were offered either live of recently crushed Asellus aquaticus. In monospecific controls, growth rates of P. tenuis were greater when fed on crushed than live Asellus, but there was no significant difference in the growth of H. stagnalis fed either live or crushed prey. In mixed cultures of predators, P. tenuis and H. stagnalis were the superior competitors when fed on crushed and live Asellus, respectively. However, when competitive pressure was low, at low densities of predators, the presence of H. stagnalis in mixed cultures fed on live prey was beneficial to the growth of P. tenuis. These results are explained in terms of the greater ability of triclads to detect damaged prey, leaking body fluids, due to their sophisticated chemosensory system, and the ability of leeches to capture live prey due to the presence of suckers. It is concluded that co-existence of the two groups in British lakes is assisted by the partitioning of food on a live or damaged basis.     

Comparison of fish and phantom midge influence on cladocerans diel vertical migration in a dual basin lake

Diurnal vertical migrations (DVM) behaviour of cladocerans was investigated in two mesotrophic Irish lakes connected by a canal, characterised by interesting differences in the presence of zooplanktivorous predators. In Doon Upper, fish (mostly juvenile perch and roach) and a little-studied phantom midge Mochlonyx fuliginosus (Chaoboridae) were found, but Doon Lower was solely inhabited by fish. As the presence of diverse predators may alter spatial avoidance behaviour of zooplankton prey in different ways, the aim of this study was to determine whether and how two predator types, fish and phantom midge larvae, have changed DVM pattern of cladocerans during day and night in Doon lakes. Two sampling series of phytoplankton, zooplankton, fish, and water physical analyses were conducted on 09–10 June and 19–20 September 2007 in both lakes. In the study conducted in June, under a similar distribution of M. fuliginosus and juvenile fish in Doon Upper, a reverse migration of Daphnia galeata was observed as a strategy allowing them to avoid both types of predators. However, in September, when M. fuliginosus lived in a 24 h refugium below the oxycline as a response to increasing predation risk posed by YOY fish penetrated the upper strata of water during day and night, reverse migrations of D. galeata were not clear. In Doon Lower, normal migration was observed as an advantageous behavioural response against visual predators (fish), in both large and small Cladocera species: D. galeata, Diaphanosoma branchyurum and Bosmina sp. Thus, our results indicate dissimilar migration patterns of D. galeata depending on the presence of one (Doon Lower) or two predators with different predation behaviour (Doon Upper).     

Invasive pike establishment in Cook Inlet Basin lakes,Alaska: diet,native fish abundance and lake environment

Northern pike (Esox lucius) were introduced to the northern Susitna Basin of south-central Alaska in the 1950’s, and have since spread throughout the upper Cook Inlet Basin. Extirpations of several native fish populations have been documented in this area. It is hypothesized here that invasive pike remodel the ecology of lakes by removing vulnerable prey types and that these changes are reflected in the diet of invasive pike. Trends in pike diet suggest that pike switch to less desirable but more abundant macroinvertebrate prey as preferred fish prey are eliminated. The impacts of pike introduction were studied in detail for one species of resident fish, the threespine stickleback (Gasterosteus aculeatus). Stickleback abundance decreases as pike invasion progresses. Conductivity is a significant environmental predictor of stickleback abundance, with higher conductivity apparently mitigating population reduction. Higher conductivity water may lessen the physiological costs of developing more robust armor, which reduces vulnerability to predation. Maximum lake depth also appears to predict stickleback abundance, though this trend was only marginally significant. Deeper lakes may provide an open-water refuge from pike predation by allowing stickleback to exist outside of the pike inhabited littoral zone. These findings indicate the importance of diverse habitat types and certain chemical and physical characteristics to the outcome of invasion by fish predators.     

GAPE‐LIMITED PREDATORS AS AGENTS OF SELECTION ON THE DEFENSIVE MORPHOLOGY OF AN INVASIVE INVERTEBRATE

Invasive species have widespread and pronounced effects on ecosystems and adaptive evolution of invaders is often considered responsible for their success. Despite the potential importance of adaptation to invasion, we still have limited knowledge of the agents of natural selection on invasive species. Bythotrephes longimanus, a cladoceran zooplankton, invaded multiple Canadian Shield lakes over the past several decades. Bythotrephes have a conspicuous caudal process (tail spine) that provides a morphological defense against fish predation. We measured viability selection on the longest component of the Bythotrephes spine, the distal spine segment, through a comparison of the lengths of first and second instar Bythotrephes collected from lakes differing in the dominance of gape‐limited predation (GLP) and nongape‐limited predation (NGLP) by fish. We found that natural selection varied by predator gape‐limitation, with strong selection (selection intensity: 0.20–0.79) for increased distal spine length in lakes dominated by GLP, and no significant selection in lakes dominated by NGLP. Further, distal spine length was 17% longer in lakes dominated by GLP, suggesting the possibility of local adaptation. As all study lakes were invaded less than 20 years prior to our collections, our results suggest rapid divergence in defensive morphology in response to selection from fish predators.     

Differential predation on the two colour morphs of Nicaraguan Crater lake Midas cichlid fish: implications for the maintenance of its gold‐dark polymorphism

Predation can play an important role in the evolution and maintenance of prey colour polymorphisms. Several factors are known to affect predator choice, including the prey's relative abundance and conspicuousness. In polymorphic prey species, predators often target the most common or most visible morphs. To test if predator choice can explain why in Midas cichlid fish the more visible (gold) morph is also more rare than the inconspicuous dark morph, we conducted predation experiments using two differently coloured wax models in Nicaraguan crater lakes. Contrary to expectations, we observed an overall higher attack rate on the much more abundant, yet less conspicuous dark models, and propose frequency‐dependent predation as a potential explanation for this result. Interestingly, the attack rate differed between different types of predators. While avian predators were biased towards the abundant and less colourful dark morphs, fish predators did not show a strong bias. However, the relative attack rate of fish predators seemed to vary with the clarity of the water, as attack rates on gold models went up as water clarity decreased. The relative differential predation rates on different morphs might impact the relative abundance of both colour morphs and thus explain the maintenance of the colour polymorphism. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 123–131.     

A review of predation as a limiting factor for bird populations in mesopredator‐rich landscapes: a case study of the UK

The impact of increasing vertebrate predator numbers on bird populations is widely debated among the general public, game managers and conservationists across Europe. However, there are few systematic reviews of whether predation limits the population sizes of European bird species. Views on the impacts of predation are particularly polarised in the UK, probably because the UK has a globally exceptional culture of intensive, high‐yield gamebird management where predator removal is the norm. In addition, most apex predators have been exterminated or much depleted in numbers, contributing to a widely held perception that the UK has high numbers of mesopredators. This has resulted in many high‐quality studies of mesopredator impacts over several decades. Here we present results from a systematic review of predator trends and abundance, and assess whether predation limits the population sizes of 90 bird species in the UK. Our results confirm that the generalist predators Red Fox (Vulpes vulpes) and Crows (Corvus corone and C. cornix) occur at high densities in the UK compared with other European countries. In addition, some avian and mammalian predators have increased numerically in the UK during recent decades. Despite these high and increasing densities of predators, we found little evidence that predation limits populations of pigeons, woodpeckers and passerines, whereas evidence suggests that ground‐nesting seabirds, waders and gamebirds can be limited by predation. Using life‐history characteristics of prey species, we found that mainly long‐lived species with high adult survival and late onset of breeding were limited by predation. Single‐brooded species were also more likely to be limited by predation than multi‐brooded species. Predators that depredate prey species during all life stages (i.e. from nest to adult stages) limited prey numbers more than predators that depredated only specific life stages (e.g. solely during the nest phase). The Red Fox and non‐native mammals (e.g. the American Mink Neovison vison) were frequently identified as numerically limiting their prey species. Our review has identified predator–prey interactions that are particularly likely to result in population declines of prey species. In the short term, traditional predator‐management techniques (e.g. lethal control or fencing to reduce predation by a small number of predator species) could be used to protect these vulnerable species. However, as these techniques are costly and time‐consuming, we advocate that future research should identify land‐use practices and landscape configurations that would reduce predator numbers and predation rates.     

Intra- and interspecific predation on the cocoons of Erpobdella octoculata (L.) (Annelida: Hirudinea)

In laboratory experiments, eleven out of twenty-one invertebrate species and all of the three fish species tested fed on freshly laid flaccid cocoons of Erpobdella octoculata, though predation was only severe from a dytiscid beetle, E. octoculata and the fish species. There was a tendency for more cocoons to be eaten by starved than fed predators. Only snail species damaged one-week-old tougher cocoons.Only one out of 471 E. octoculata collected during their breeding season from the stony shores of two eutrophic lakes had a cocoon in its gut.The role of cannibalism of and interspecific predation on cocoons in the control and regulation of lake-dwelling populations of the erpobdellid is discussed, and assessed to be relatively unimportant. A more likely regulatory mechanism may involve high juvenile mortality from as yet unknown causes.     

THE TROPHIC ECOLOGY OF FRESHWATER GAMMARUS SPP. (CRUSTACEA:AMPHIPODA): PROBLEMS AND PERSPECTIVES CONCERNING THE FUNCTIONAL FEEDING GROUP CONCEPT

Gammarus spp. are widespread throughout a diverse range of freshwater habitats and can be the dominant part of many benthic macroinvertebrate assemblages, in terms of both numbers and/or biomass. Although the vast majority of studies have emphasized the herbivorous nature of Gammarus spp. and their ‘shredder’ functional feeding group (FFG) classification, we show that a far wider food base is exploited than has been previously acknowledged. This ‘plasticity’ as herbivore/predator is linked to the success of Gammarus spp. in persisting in and colonizing/invading disturbance-prone ecosystems. Intraguild predation and cannibalism are more common than previously realized. This behaviour appears to be a causal mechanism in many amphipod species replacements. Additionally, Gammarus spp. are major predators of other members of the macroinvertebrate community. Furthermore, while many studies have emphasized fish predation on Gammarus spp., we illustrate how this fish: amphipod, predator: prey interaction may be a two-way process, with Gammarus spp. themselves preying upon juvenile and wounded/trapped fish. We urge that a new realism be adopted towards the trophic ecology of Gammarus spp. and their role as predators and prey and that previously established FFG assumptions of both the food and the feeder be questioned critically.     

Fleeing towards death – leech‐induced behavioural defences increase freshwater snail susceptibility to predatory fish

Prey species are often exposed to multiple predators, which presents several difficulties to prey species. This is especially true when the response to one predator influences the prey’s susceptibility to other predators. Predator‐induced defences have evolved in a wide range of prey species, and experiments involving predators with different hunting strategies allow researchers to evaluate how prey respond to multiple threats. Freshwater snails are known to respond to a variety of predators with both morphological and behavioural defences. Here we studied how freshwater snails Radix balthica responded behaviourally to fish and leech predators, both separately and together. Our aim was to explore whether conflicting predator‐induced responses existed and, if so, what effect they had on snail survival when both predatory fish and leeches were present. We found that although R. balthica increased refuge use when exposed to predatory fish, they decreased refuge use when exposed to predatory leeches. When both predators were present, snails showed a stronger response towards leech than fish and responded by leaving the refuge. This response made the snails more susceptible to fish predation, which increased snail mortality when exposed to both fish and leech compared to fish only. We show that predators that have a relatively low predation rate can substantially increase mortality rates by indirect effects. By forcing snails out of refuges such as rock and macrophyte habitats, leeches can indirectly increase predation from molluscivorous fish and may thus affect snail densities.     

四种鲤科鱼类对捕食胁迫行为响应的种间比较

为了比较早期捕食胁迫经历和当前环境中存在的捕食者对鱼类行为的影响,并考查这些影响是否存在种间差异,研究分别考查了测定环境(有、无捕食者存在)对有、无捕食胁迫经历的鳊(Parabramis pekinensis)、草鱼(Ctenopharyngodon idellus)、鲫(Carassius auratus)和中华倒刺鲃(Spinibarbus sinensis)等4种鲤科鱼类探索性、活跃性和勇敢性的影响。结果发现:早期捕食胁迫经历与当前环境条件对鱼类行为产生截然不同的影响,且存在较大的种间差异。无捕食胁迫经历的鳊、草鱼和中华倒刺鲃均会对陌生的捕食者乌鳢(Channa argus)做出行为响应,提示这3种鱼可能对陌生捕食者具有一定的识别能力,但这种识别与猎物鱼通过捕食胁迫经历获得的识别仍具有一定差距;具有捕食胁迫经历的鳊和中华倒刺鲃在空白环境中未表现出反捕食行为,可能是节约能量的一种策略。总体而言,草鱼对捕食胁迫经历和测定环境处理反应更为敏感,而中华倒刺鲃的反应则相对保守。但当周围环境中存在捕食者时, 4种鲤科鱼类均会通过维持较高运动状态的方式来应对捕食者。维持这种应激状态可能对猎物鱼...     

MORPHOLOGICAL EVOLUTION MEDIATED BY BEHAVIOR IN THE DAMSELFLIES OF TWO COMMUNITIES

Behavior can play a mediating role in determining the selective pressures that influence the evolution of morphological structures. To examine this, I quantified patterns of morphological variation among larvae of Enallagma damselfly species (Odonata: Coenagrionidae) that use different behaviors to avoid the major predators found in each of two communities, lakes with and without fish. Specifically, I quantified the sizes and shapes of the abdomens and caudal lamellae (used for swimming) and legs for three species from fishless lakes and six species from lakes with fish. A preliminary cladistic analysis indicates that species within each lake type are not members of a single clade, which supports the conclusions of previous odonate taxonomists. Previous studies have shown that species in fishless lakes are very active, running and swimming frequently and at high rates of speed in the absence of predators, and they avoid their primary predators, large dragonflies, by swimming. These species have the widest abdomens, the largest caudal lamellae relative to overall body size, and the longest legs of the species studied, which should make them powerful swimmers and runners. Furthermore, species in fishless lakes are morphologically very similar to one another and differ greatly from fish-lake species, although each is more closely related to species in fish lakes. In contrast, species from lakes with fish move very slowly and infrequently in the absence of predators and do not attempt to evade attacking predators. However, despite their behavioral similarity, large interspecific variation in morphology exists among the fish-lake species, and the only morphological patterns were differences associated with membership in the two primary clades identified in the cladistic analysis. A modification of Felsenstein's (1985) method of evolutionary contrasts which allows character change to be isolated along single branches is introduced and is used to reconstruct the evolutionary histories of these characters. This analysis suggests that large increases in caudal lamella size, abdominal segment lengths and widths, and leg length accompany speciation events associated with habitat shifts from fish-lakes to fishless lakes. Following habitat shifts selection pressures exerted by dragonfly predation apparently favored swimming as an escape tactic, which mediated selection pressures onto morphologies used in swimming to increase swimming performance; morphological patterns in extant species reflect this adaptation to a new environment. Mechanisms by which behaviorally mediated selection could have accelerated evolutionary dynamics following founder events are discussed.     

Fishing spiders, green sunfish, and a stream-dwelling water strider: male-female conflict and prey responses to single versus multiple predator environments

Many studies have experimentally addressed the effects of a particular predator species on prey behavior. In nature, however, prey frequently face multiple species of predators that often vary in their predatory mode and in their level of predation risk. Relatively few studies have considered prey responses under these complex conditions. In Kentucky, the stream-dwelling water strider (Aquariusremigis) coexists with many potentially dangerous predators, two of which are the green sunfish (Lepomiscyanellus) and the fishing spider (Dolomedesvittatus). Green sunfish occupy stream pools and attack water striders from below. In contrast, fishing spiders hunt along stream shorelines where they perch on overhanging vegetation or rocks and attack water striders near shore. We compared how A. remigis individuals respond to these two very different predators in pools with one or both predators. The presence of sunfish in pools had strong effects on male water strider behavior, including increased use of three types of refuge from sunfish (riffles, climbing out of the water, sitting on the water but at the edges of pools), decreased activity and a decreased number of aggressive males on the water. Spiders also influenced water strider behavior; male water striders avoided spiders by shifting away from the edges of pools. Comparisons of the effects of the two predator species showed that in general, antipredator responses by male water striders were stronger in pools with fish alone than in those with spiders alone. In the presence of both predators, male water strider behavior (microhabitat use and activity) was generally similar to behavior in the presence of fish alone. In contrast, female water striders showed no significant response to the presence of sunfish, and little response to the presence of spiders. This lack of response could be because females spent much of their time in refuges even in the absence of predators (apparently hiding from harassment by males). Both spiders and fish caused decreases in water strider mating activity. The presence of fish reduced both the number of matings per pool (mating frequency), and mean mating durations. Spiders induced a decrease in mean mating duration, but not in mating frequency. The largest reductions in mating activity occurred in pools with both predators present. Pools with either spiders or fish alone suffered 15–20% water strider mortality during our experiment (versus no mortality in predator-free pools). Extant theory suggests that when prey face conflicting microhabitat responses to two predators (as in this study), the predators should have facilitative effects on predation rates (i.e., prey that avoid one predator are often killed by the other and vice versa). Mortality rates in pools with both predators present, however, were not significantly different from that predicted by a null model of multiple predator effects. The lack of predator facilitation can be explained by the compensatory reductions in water strider activity and mating activity in the presence of both predators. Received: 26 August 1996 / Accepted: 12 June 1998     

Major changes in pelagic rotifers during natural and forced recovery from acidification

Pelagic rotifers were studied in lakes with contrasting acidification histories situated in an acid-stressed region of southern Norway. Life histories and spatial distribution varied considerably between the investigated species, and influenced the recovery processes. Most headwater lakes have experienced strongly acidified environments during the last five decades, whereas lakes close to the Skagerrak coast have been stable within the same period. Rotifer diversity and abundance were reduced in the most acidic sites and increased towards the coast. Most surveyed species are known to possess sediment egg-banks, and after chemical recovery most rotifers dispersed into the plankton from these egg-banks and produced viable populations. Some species of the genera Polyarthra and Collotheca, and the species Kellicotta longispina and Keratella serrulata showed a striking ability to tolerate acidification, and were the dominant taxa in the acidmost environments. K. serrulata characterised, but did not numerically dominate, acid rotifer communities especially in the most coloured sites, and decreased following liming. The predominantly bacteriophageous genus Conochilus exploded in numbers shortly after liming, most probably because bacteria increased strongly during this transition phase. Planktivorous fish influenced indirectly rotifer abundance by consuming invertebrate predators and important rotifer competitors such as filter feeding cladocerans. Invertebrate predators, such as larvae of Chaoborus spp. and Heterocope saliens probably influenced rotifer distributional patterns in a complex top-down manner, both during chronic acidification and liming in environments with low fish predation. Important rotifer predators such as pelagic cyclopoid copepods, Bythotrephes longimanus and Leptodora kindti, were absent from the most acidic fishless lakes. Considerable populations of large-sized Daphnia longispina probably suppressed several rotifer species in sites with low fish predation, as did large populations of Bosmina longispina and Ceriodaphina quadrangula in lakes with intense fish predation.     

Subtle top‐down control of a freshwater meiofaunal assemblage by juvenile fish

1. Top‐down control of prey assemblages by fish predators has been clearly demonstrated in lakes (for zooplankton prey) and rivers (for macroinvertebrate prey). Fish predation can have a significant impact on the body size of prey assemblages; often large‐bodied prey are reduced in abundance, and indirect facilitation of small‐bodied prey occurs potentially initiating a trophic cascade. 2. Benthic communities in aquatic ecosystems also include a numerous and functionally important meiofaunal‐sized component, but in freshwaters the impact of fish predation on meiofaunal assemblages is unknown. We used a laboratory microcosm study to explore the impact of juvenile fish predation on the abundance and size structure of a riverine meiofaunal assemblage. 3. The presence of fish in our microcosms had no significant effect on overall meiofaunal (temporary and permanent) abundance. However, for the Copepoda, we found the first evidence of top‐down control of freshwater meiofaunal assemblages; in microcosms with juvenile fish, the abundance of large‐bodied Copepoda was significantly reduced, whereas small‐bodied Copepoda were significantly more abundant suggesting indirect facilitation. 4. We conclude that predation by juvenile fish can alter the structure of freshwater meiofaunal assemblages, although we do not yet know whether these relatively subtle changes are overwhelmed by large‐scale events such as flow disturbances.     

Lake size and fish diversity determine resource use and trophic position of a top predator in high‐latitude lakes

Prey preference of top predators and energy flow across habitat boundaries are of fundamental importance for structure and function of aquatic and terrestrial ecosystems, as they may have strong effects on production, species diversity, and food‐web stability. In lakes, littoral and pelagic food‐web compartments are typically coupled and controlled by generalist fish top predators. However, the extent and determinants of such coupling remains a topical area of ecological research and is largely unknown in oligotrophic high‐latitude lakes. We analyzed food‐web structure and resource use by a generalist top predator, the Arctic charr Salvelinus alpinus (L.), in 17 oligotrophic subarctic lakes covering a marked gradient in size (0.5–1084 km²) and fish species richness (2–13 species). We expected top predators to shift from littoral to pelagic energy sources with increasing lake size, as the availability of pelagic prey resources and the competition for littoral prey are both likely to be higher in large lakes with multispecies fish communities. We also expected top predators to occupy a higher trophic position in lakes with greater fish species richness due to potential substitution of intermediate consumers (prey fish) and increased piscivory by top predators. Based on stable carbon and nitrogen isotope analyses, the mean reliance of Arctic charr on littoral energy sources showed a significant negative relationship with lake surface area, whereas the mean trophic position of Arctic charr, reflecting the lake food‐chain length, increased with fish species richness. These results were supported by stomach contents data demonstrating a shift of Arctic charr from an invertebrate‐dominated diet to piscivory on pelagic fish. Our study highlights that, because they determine the main energy source (littoral vs. pelagic) and the trophic position of generalist top predators, ecosystem size and fish diversity are particularly important factors influencing function and structure of food webs in high‐latitude lakes.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.     

Predator identity and consumer behavior: differential effects of fish and crayfish on the habitat use of a freshwater snail

Predators can alter the outcome of ecological interactions among other members of the food web through their effects on prey behavior. While it is well known that animals often alter their behavior with the imposition of predation risk, we know less about how other features of predators may affect prey behavior. For example, relatively few studies have addressed the effects of predator identity on prey behavior, but such knowledge is crucial to understanding food web interactions. This study contrasts the behavioral responses of the freshwater snail Physellagyrina to fish and crayfish predators. Snails were placed in experimental mesocosms containing caged fish and crayfish, so the only communication between experimental snails and their predators was via non-visual cues. The caged fish and crayfish were fed an equal number of snails, thereby simulating equal prey mortality rates. In the presence of fish, the experimental snails moved under cover, which confers safety from fish predators. However, in the presence of crayfish, snails avoided benthic cover and moved to the water surface. Thus, two species of predators, exerting the same level of mortality on prey, induced very different behavioral responses. We predict that these contrasting behavioral responses to predation risk have important consequences for the interactions between snails and their periphyton resources. Received: 1 June 1998 / Accepted: 12 October 1998     

Adaptive evolution to novel predators facilitates the evolution of damselfly species range shifts

Most species have evolved adaptations to reduce the chances of predation. In many cases, adaptations to coexist with one predator generate tradeoffs in the ability to live with other predators. Consequently, the ability to live with one predator may limit the geographic distributions of species, such that adaptive evolution to coexist with novel predators may facilitate range shifts. In a case study with Enallagma damselflies, we used a comparative phylogenetic approach to test the hypothesis that adaptive evolution to live with a novel predator facilitates range size shifts. Our results suggest that the evolution of Enallagma shifting from living in ancestral lakes with fish as top predators, to living in lakes with dragonflies as predators, may have facilitated an increase in their range sizes. This increased range size likely arose because lakes with dragonflies were widespread, but unavailable as a habitat throughout much of the evolutionary history of Enallagma because they were historically maladapted to coexist with dragonfly predators. Additionally, the traits that have evolved as defenses against dragonflies also likely enhanced damselfly dispersal abilities. While many factors underlie the evolutionary history of species ranges, these results suggest a role for the evolution of predator‐prey interactions.