Performance as a fish predator of largemouth bass [<Emphasis Type="Italic">Micropterus salmoides</Emphasis> (Lacepède)] invading Japanese freshwaters: a review

This review focuses on how predator performance of the invasive largemouth bass [Micropterus salmoides (Lacepède)] has been, or will be, formed in Japanese freshwaters. Predation impacts of largemouth bass on fish communities appear pervasive in both Japanese as well as North American freshwaters. Factors affecting performance as a piscivorous predator are (1) light intensity and water clarity, (2) oxygen depletion, (3) prey size and gape size, (4) behavioral refuge of prey, (5) weed beds as refuge for prey fish, (6) interaction with bluegill. Size and behavioral refuges requirements are so rigorous that they may have evolved only in some North American prey fish species like bluegill; therefore, most Japanese native fish species are unlikely to be equipped with such refuges. However, refuge habitats like aquatic weed beds could develop in Japanese freshwaters, allowing prey fish species to survive under predation pressure. The density, architecture, and species composition of aquatic plants may affect their suitability as refuges. Studies in Japanese waters have suggested that the presence of rich aquatic vegetation or invasive bluegill in bass-introduced waters have suppressed the predation impact of largemouth bass on fish communities. In addition to these environmental factors, original genotypic and phenotypic traits of the introduced largemouth bass, and hybridization between different lineages of largemouth bass or with Florida bass [Micropterus floridanus (Lesueur)] may be involved in further adaptation of invasive largemouth bass to Japanese freshwaters.     

The influence of littoral zone coarse woody habitat on home range size, spatial distribution, and feeding ecology of largemouth bass (Micropterus?salmoides)

Coarse woody habitat (CWH) may be a critical feature of lakes that influences fish distributions, movement patterns, and feeding habits. We used radio telemetry to examine the role of CWH in determining the movements of largemouth bass (Micropterus salmoides Lacepede) in the context of two whole-lake experiments that provided a gradient of four lake basins varying in natural and manipulated CWH. We also conducted diet studies on largemouth bass in these lakes to test for correlates among consumption rate and prey selectivity with bass behavior. Our results indicated that largemouth bass in basins with lower CWH abundances had larger home ranges, spent more time in deep water, were more selective predators, and showed lower consumption rates. Largemouth bass in basins with higher CWH abundances showed the opposite patterns. Low CWH abundances were correlated with a shift in largemouth bass foraging behavior from sit-and-wait to actively searching. This increased activity, coupled with the potential decline of prey fish species in the absence of CWH, may decrease largemouth bass growth potential regardless of the prey type consumed. Our results suggest that lakeshore residential development and associated removals of CWH from lakes may influence fish behavior, while CWH augmentation may reverse some of those changes. Handling editor: Steven Declerck     

Aquatic vegetation and largemouth bass population responses to water-level variations in Lake Okeechobee,Florida (USA)

A five-year study examined the responses of submerged aquatic vegetation (SAV), emergent vegetation, and largemouth bass (Micropterus salmoides) to variations in water level in a large lake in Florida, USA. SAV was assessed using a combined transect survey/spatial mapping approach, emergent vegetation was quantified with aerial photography and GIS, and bass were surveyed by electro-shocking. During the period leading up to this study (1995–1999), water levels were high in the lake, and the SAV was reduced in spatial extent and biomass, compared to its condition in the early 1990s. Spatial extent of emergent vegetation also was low, and largemouth bass surveys indicated low densities and failure to recruit young fish into the population. This was attributed to the lack of critical vegetative habitat. In spring 2000, the lake was lowered by discharging water from major outlets, and this was followed by a regional drought. Water levels dropped by 1m, and there was widespread development of Chara lawns in shoreline areas, with coincident increases in water clarity. There was some germination of vascular SAV, but Chara was the extreme dominant, such that structural complexity remained low. There was no substantive improvement in bass recruitment. During 2001, water levels declined further, and emergent plants germinated in exposed areas of the lake bottom. SAV was restricted to sites farther offshore, and continued to be dominated by Chara. There again was no bass response. In 2002, conditions changed when water levels increased to a moderate depth, flooding shoreline habitat to 0.5m. Vascular SAV increased in biomass and spatial extent, such that the community developed a high structural complexity. At the same time, emergent aquatic plants developed dense stands along the western shoreline. Largemouth bass had a strong recruitment of young fish for the first time in 5years. Recruitment continued to be successful in 2003, when spatial extent of SAV was somewhat reduced by higher water but total biomass and diversity remained high. These results demonstrate an important effect of inter-annual variation in water depth on the population dynamics of aquatic plants and fish in a subtropical lake.     

Variations in piscivory of invasive largemouth bass <Emphasis Type="Italic">Micropterus salmoides</Emphasis> associated with pond environments

Introductions of non-native predatory fishes can be a major driver of aquatic biodiversity loss. The largemouth bass Micropterus salmoides (L.) has been introduced throughout much of the world, thereafter negatively affecting native faunal communities owing to its predatory impact. To investigate the environmental factors affecting the predatory performance of invasive bass, we examined the stomach contents and habitat characteristics of bass in 15 irrigation farm ponds in northeastern Japan. The food habits of the bass populations differed among the studied ponds: the predominant prey items were fishes among bass in seven of the ponds, whereas aquatic invertebrates (mainly insects and zooplankton) were the predominant taxa in the diets of bass in the eight remaining ponds, with the onset of piscivory related to body size. The results of multivariate analysis indicated that the extent to which the bass consumed fish was positively associated with fish prey abundance and negatively associated with percentage of aquatic vegetation coverage. We suggest that the extent of aquatic vegetation coverage strongly influenced the predation efficiency of bass in the ponds. These findings might be employed to assess a pond ecosystem’s vulnerability to invasive largemouth bass and to reduce the predator’s impact on native fish species by improvements to the habitat.     

Effects of crayfish size,orientation, and movement on the reactive distance of largemouth bass foraging in clear and turbid water

Laboratory experiments were performed in clear and turbid water to determine the effects of prey size, orientation, and movement on the reactive distance of largemouth bass (Micropterus salmoides) when feeding on crayfish (Procambarus acutus). In clear water, the reactive distance increased linearly with an increase in prey size, and prey movement resulted in a significant increase in the reactive distance. Prey orientation (head-on versus perpendicular) did not change the reactive distances. In moderately turbid water, the reactive distance did not increase with increased prey size, and prey movement did not result in any changes in the reactive distance. The absence of any effects of prey orientation in clear water or prey movement in turbid water is inconsistent with results from studies using different species (primarily planktivorous fish). I propose that largemouth bass change their foraging tactics as prey visibility changes. When prey are highly visible (low turbidity), predators attack (react) only after prey recognition, which is based on multiple cues such as prey size (length, width) and movement. When prey are less visible (high turbidity), predators attack immediately upon initial prey sighting, which does not depend on prey size or movement.     

Phenotypic variation and vulnerability to predation in juvenile bluegill sunfish (<Emphasis Type="Italic">Lepomis macrochirus</Emphasis>)

Bluegill sunfish (Lepomis macrochirus) are known to diversify into two forms specialized for foraging on either limnetic or littoral prey. Because juvenile bluegills seek vegetative cover in the presence of largemouth bass (Micropterus salmoides) predators, natural selection should favor the littoral body design at size ranges most vulnerable to predation. Yet within bluegill populations, both limnetic and littoral forms occur where vegetation and predators are present. While adaptive for foraging in different environments, does habitat-linked phenotypic variation also influence predator evasiveness for juvenile bluegills? We evaluate this question by quantifying susceptibility to predation for two groups of morphologically distinct bluegills; a limnetic form characteristic of bluegills inhabiting open water areas (limnetic bluegill) and a littoral form characteristic of bluegills inhabiting dense vegetation (littoral bluegill). In a series of predation trials, we found that bluegill behaviors differed in open water habitat but not in simulated vegetation. In open water habitat, limnetic bluegills formed more dense shoaling aggregations, maintained a larger distance from the predator, and required longer amounts of time to capture than littoral bluegill. When provided with simulated vegetation, largemouth bass spent longer amounts of time pursuing littoral bluegill and captured significantly fewer littoral bluegills than limnetic fish. Hence, morphological and behavioral variation in bluegills was linked to differential susceptibility to predation in open water and vegetated environments. Combined with previous studies, these findings show that morphological and behavioral adaptations enhance both foraging performance and predator evasiveness in different lake habitats.     

Diel movement of smallmouth yellowfish Labeobarbus aeneus in the Vaal River,South Africa

Diel movements of Orange–Vaal smallmouth yellowfish Labeobarbus aeneus (Burchell, 1822) in the Vaal River, South Africa, were determined by externally attaching radio transmitters to 11 adult fish and manually tracking them between March and May 2012. Twenty-four radio telemetry monitoring surveys produced 2 304 diel tracks. At night, yellowfish displayed a preference for slow shallow (<0.3?m s?1, <0.5?m) and fast shallow habitats (>0.3?m s?1, <0.3?m), whereas by day they avoided these habitats, preferring fast deep areas (>0.3?m s?1, >0.3?m). The average total distance of 272?m moved per 24-hour period was three times greater than the diel range, and the average maximum displacement per minute was significantly higher in daytime (4?m) than at night (1.5?m). These findings suggest that L. aeneus is active primarily during the day in fast-flowing, deeper waters, and relatively inactive at night, when it occupies shallower habitats. This behaviour should be further explored to identify causal mechanisms underlying the diel habitat shifts in this species such as water temperature, foraging tactics and/or predator avoidance.     

Hunting behavior of Florida largemouth bass, Micropterus salmoides floridanus, in a channelized river

The distribution and behavior of Florida largemouth bass, Micropterus salmoides floridanus, and their main prey (sunfish, genus Lepomis, and the cichlid Tilapia mariae) were studied in southern Florida to determine how fish behave in the simplified habitats found in channelized rivers. Time budgets were constructed from focal animal observations on 69 bass. Patterns of behavior associated with hunting were performed during a significantly higher proportion of the time when bass were in vegetated habitats. Scan samples of the behavior of 236 observed bass revealed that hunting was more common in areas of high structural complexity. Only 38% of observed bass were solitary, with the majority occurring in groups with either conspecifics or in mixed-species groups with similar sized bluegill sunfish, Lepomis macrochirus. Largemouth bass (n=1014) and sunfish (n=1372) were significantly more abundant in areas with vegetation and were almost entirely absent from the water column in the center of the canal. All species of fish avoided the water column, where currents were swift and no cover was available. The structure of the habitat appears to be important in the way largemouth bass organize their activity patterns. This suggests that habitat availability in channelized rivers significantly influences important behaviors such as hunting, thus potentially altering energy budgets and population dynamics of both predator and prey.     

Antipredator behaviour of 0+ year Perca fluviatilis: effect of vegetation density and turbidity

In this study, the combined influence of vegetation density and water turbidity on habitat utilization of a prey fish, 0+ year perch Perca fluviatilis , under predation risk (pike, Esox lucius ) was investigated. The vegetated habitat was overall preferred over the open habitat in the presence of a predator. The level of turbidity, and to a lesser extent vegetation density, however, influenced the response of 0+ year perch. The use of the vegetated habitat was lower in very turbid than in clear and turbid conditions, suggesting reduced antipredator behaviour in very turbid water. The effect of vegetation density on antipredator behaviour was only present in clear water, where the use of a structural refuge decreased with increasing vegetation density. No such effect was observed in turbid and very turbid water. The results showed that the structuring role of vegetation or habitat complexity may diminish with increased turbidity. The observed masking effect of turbidity suggests that predator‐prey interactions in vegetated habitats are more complex than what has generally been thought.     

Fish stocking creates an ecological trap for an avian predator via effects on prey availability

In anthropogenic landscapes animals may be lured into low‐quality habitats where they survive or reproduce poorly (‘ecological traps’). I investigated breeding habitat selection in relation to intra‐seasonal changes in food availability and reproductive output in red‐necked grebes Podiceps grisegena, a size‐limited predator, of common carp Cyprinus carpio ponds. Carp farms constitute highly heterogeneous habitat mosaics due to separate stocking of different age/size fish. Pond features significant for grebe settling decisions, i.e. hydroperiod and emergent vegetation cover, had no obvious effects on prey abundance for chicks and on fledging success. Breeding grebes avoided ponds containing fish too large for them to ingest but exhibited little preference between ponds with medium‐sized one‐year‐old carp that could be exploited by pre‐laying birds, and ponds designated for young‐of‐the‐year carp, where only invertebrates and amphibians were available as prey in early spring. Red‐necked grebes settling on ponds with medium‐sized fish failed to predict future shifts in interactions with carp stocks; carp exceeded the prey‐size threshold of chicks and adversely affected their non‐fish prey levels. The resulting food shortage led to severe egg‐to‐fledging mortality rates compared to fishless ponds or those containing young‐of‐the‐year fish. This study shows that waterbirds vulnerable to competition from fish can risk maladaptive habitat selection due to unrecognised spatial and temporal variation in food resources caused by fish stocking practices. Ecological traps created by perturbations to trophic interactions may be common but difficult to detect because altered dynamics of trophic resources can affect wildlife indirectly. As with other types of ecological traps, manipulation of habitat features identified as attractive cues for settling animals, but not related to critical food resources, may help to reduce perceptual pitfalls. For example, wetland management to mitigate trap effects driven by commercially stocked fish should preserve abundant emergent vegetation in habitats with weak fish impact and extend their hydroperiod.     

Changes in aquatic vegetation and fish communities following 5 years of sustained low water levels in coastal marshes of eastern Georgian Bay,Lake Huron

Aquatic vegetation in the relatively pristine coastal wetlands of eastern Georgian Bay provides critical habitat for a diverse fish community. Declining water levels in Lake Huron over the past decade, however, have altered the wetland plant assemblages in favour of terrestrial (emergent and meadow) taxa and have thus reduced or eliminated this important ecosystem service. In this study, we compared IKONOS satellite images for two regions of eastern Georgian Bay (acquired in 2002 and 2008) to determine significant changes in cover of four distinct wetland vegetation groups [meadow (M), emergent (E), high‐density floating (HD) and low‐density floating (LD)] over the 6 years. While LD decreased significantly (mean ?2995.4 m²), M and HD increased significantly (mean +2020.9 m² and +2312.6 m², respectively) between 2002 and 2008. Small patches of LD had been replaced by larger patches of HD. These results show that sustained low water levels have led to an increasingly homogeneous habitat and an overall net loss of fish habitat. A comparison of the fish communities sampled between 2003 and 2005 with those sampled in 2009 revealed that there was a significant decline in species richness. The remaining fish communities were also more homogeneous. We suggest that the observed changes in the wetland plant community due to prolonged low water levels may have resulted in significant changes in the fish communities of coastal wetlands in eastern Georgian Bay.     

Effects of turbidity and an invasive waterweed on predation by introduced largemouth bass

Anthropogenic activities lead to changes in characteristics of aquatic ecosystems, including alteration of turbidity and addition of invasive species. In this study, we tested how changes in turbidity and the recent invasion of an aquatic macrophyte, Egeria densa, may have changed the predation pressure by introduced largemouth bass on juvenile striped bass and delta smelt, two species that have seen a drastic decline in recent decades in the Sacramento-San Joaquin Delta. In a series of mesocosm experiments, we showed that increases in vegetation density decreased the predation success of largemouth bass. When placed in an environment with both open water and vegetated areas, and given a choice to forage on prey associated with either of these habitats, largemouth bass preyed mainly on open water species as opposed to vegetation-associated species, such as juvenile largemouth bass, bluegill or red swamp crayfish. Finally, we showed that turbidity served as cover to open water species and increased the survival of delta smelt, an endemic species at risk. We also found that such open water prey tend not to seek refuge in the vegetation cover, even in the presence of an imminent predation threat. These results provide the beginning of a mechanistic framework to explain how decreases in turbidity and increases in vegetation cover correlate with a decline of open water species in the Sacramento-San Joaquin Delta.     

Fish reduce habitat coupling by a waterbird: evidence from combined stable isotope and conventional dietary approaches

Aquatic consumers can function as habitat couplers by using allochthonous subsidies of prey that migrate across ecosystem boundaries. We examined the relative use of allochthonous (invertebrates—terrestrial or living on littoral vegetation; immigrating amphibians) versus autochthonous (aquatic invertebrates, fish) resources by the red-necked grebe Podiceps grisegena, a generalist predator, on fishless ponds versus ponds stocked with common carp Cyprinus carpio. We combined conventional methods of diet estimation with stable carbon and nitrogen analyses of egg yolks and putative prey of grebes. Prelaying grebes were observed to take mainly adult amphibians on fishless ponds and fish on stocked ponds. Alimentary tract analyses gave more weight to invertebrate prey, especially leaf beetles Donaciinae, apparently picked off water or emergent plants. Bayesian isotopic mixing models did not reveal predominance of a single food source but indicated that in the presence of fish grebes received relatively less energy for egg formation from amphibians and leaf beetles. Overall, our results show that grebes relied more on allochthonous resources (range of means 50–97 % of the biomass contribution estimated by different assessment methods) in the absence than in the presence of fish (8–23 %). We suggest that habitat coupling by waterbirds may be controlled by fish, which can suppress external prey subsidies, apart from being an attractive food for piscivorous birds.     

Effects of Submerged Vegetation on Water Clarity Across Climates

A positive feedback between submerged vegetation and water clarity forms the backbone of the alternative state theory in shallow lakes. The water clearing effect of aquatic vegetation may be caused by different physical, chemical, and biological mechanisms and has been studied mainly in temperate lakes. Recent work suggests differences in biotic interactions between (sub)tropical and cooler lakes might result in a less pronounced clearing effect in the (sub)tropics. To assess whether the effect of submerged vegetation changes with climate, we sampled 83 lakes over a gradient ranging from the tundra to the tropics in South America. Judged from a comparison of water clarity inside and outside vegetation beds, the vegetation appeared to have a similar positive effect on the water clarity across all climatic regions studied. However, the local clearing effect of vegetation decreased steeply with the contribution of humic substances to the underwater light attenuation. Looking at turbidity on a whole-lake scale, results were more difficult to interpret. Although lakes with abundant vegetation (>30%) were generally clear, sparsely vegetated lakes differed widely in clarity. Overall, the effect of vegetation on water clarity in our lakes appears to be smaller than that found in various Northern hemisphere studies. This might be explained by differences in fish communities and their relation to vegetation. For instance, unlike in Northern hemisphere studies, we find no clear relation between vegetation coverage and fish abundance or their diet preference. High densities of omnivorous fish and coinciding low grazing pressures on phytoplankton in the (sub)tropics may, furthermore, weaken the effect of vegetation on water clarity.     

Influence of intra‐ and interspecific variation in predator–prey body size ratios on trophic interaction strengths

1. Predation is a pervasive force that structures food webs and directly influences ecosystem functioning. The relative body sizes of predators and prey may be an important determinant of interaction strengths. However, studies quantifying the combined influence of intra‐ and interspecific variation in predator–prey body size ratios are lacking.
2. We use a comparative functional response approach to examine interaction strengths between three size classes of invasive bluegill and largemouth bass toward three scaled size classes of their tilapia prey. We then quantify the influence of intra‐ and interspecific predator–prey body mass ratios on the scaling of attack rates and handling times.
3. Type II functional responses were displayed by both predators across all predator and prey size classes. Largemouth bass consumed more than bluegill at small and intermediate predator size classes, while large predators of both species were more similar. Small prey were most vulnerable overall; however, differential attack rates among prey were emergent across predator sizes. For both bluegill and largemouth bass, small predators exhibited higher attack rates toward small and intermediate prey sizes, while larger predators exhibited greater attack rates toward large prey. Conversely, handling times increased with prey size, with small bluegill exhibiting particularly low feeding rates toward medium–large prey types. Attack rates for both predators peaked unimodally at intermediate predator–prey body mass ratios, while handling times generally shortened across increasing body mass ratios.
4. We thus demonstrate effects of body size ratios on predator–prey interaction strengths between key fish species, with attack rates and handling times dependent on the relative sizes of predator–prey participants.
5. Considerations for intra‐ and interspecific body size ratio effects are critical for predicting the strengths of interactions within ecosystems and may drive differential ecological impacts among invasive species as size ratios shift.

     

Factors affecting species number and density of dabbling duck guilds in North Europe

We addressed how species number and pair density in guilds of co-existing species is related to habitat structure, and to the abundance and diversity of food resources using the assemblage of seven species of dabbling ducks (genus Anas ) breeding in 60 lakes distributed over six regions in temperate north Europe
Partial correlation and multiple regression revealed that species richness was best predicted by habitat structural diversity as indexed by a principal component analysis based on 18 vegetation and lake characteristics, and by the abundance of aquatic and emergent prey We found no effect of lake size or prey size diversity on species richness Pair density was correlated with the percentage of shoreline with horsetails ( Equisetum ), by habitat structural diversity and by the abundance of emergent invertebrate prey Neither prey size diversity nor abundance of aquatic prey correlated with pair density Species richness and pair density in North European duck guilds vary both with habitat structure and prey availability     

A comparison of prey capture kinematics in hatchery and wild Micropterus salmoides floridanus: effects of ontogeny and experience

Analysis of high‐speed videography demonstrated that juvenile wild Florida largemouth bass Micropterus salmoides floridanus captured live prey with very rapid movements and large excursions. Hatchery fish of the same age, raised on pelleted feed, however, used slower kinematics with smaller excursions, yielding strikes with a higher degree of 'suction'. Capture events of hatchery Florida largemouth bass fed live prey for the first time were characterized by movements that had smaller excursion measurements than wild fish and resulted in a decreased level of capture success. After five exposures to elusive mosquitofish Gambusia holbrooki , hatchery Florida largemouth bass adapted their behaviour to capture prey at the kinematic level of wild fish.     

Factors affecting the habitat selection of tench in a shallow eutrophic lake

The activity patterns and habitat selection of tench in a shallow eutrophic lake were determined by radio telemetry. Environmental variables, including prey availability, depth and vegetational composition were quantified in order to explain observed patterns of distribution. Tench were generally only active at night, foraging on benthic animal prey, particularly chironomid larvae. Feeding fish clearly selected for particular locations although only weak associations with depth and densities of preferred prey were found. In addition, it appears that fish spent a considerable time searching for prey and travelling relatively large distances in the process, with the result that only relatively few prey were ingested during the course of a night. During daylight, fish were almost completely inactive, resting together in favoured locations and displaying a strong association for the littoral emergent vascular plant, Typha angustifolia . This may be because Typha usually grows in relatively deep water and forms stands of relatively widely spaced stems, thus allowing such large fish to penetrate deeply into cover. Management of eutrophic waters to encourage tench should take this habitat preference into account.     

Ontogenetic changes in foraging tactics of the piscivorous cornetfish <Emphasis Type="Italic">Fistularia commersonii</Emphasis>

Foraging behaviors of the piscivorous cornetfish Fistularia commersonii were observed at shallow reefs in Kuchierabu-jima Island, southern Japan. This fish foraged on two types of prey fishes: one was reef fish that typically dwell on or near substrata (e.g., Tripterygiidae and Labridae), and the other was pelagic fish that shoal in the water column (e.g., Clupeidae and Carangidae). The prey sizes, prey types and foraging behaviors changed as the predator size increased. Prey sizes were largely limited by gape size of the cornetfish, and small predators consumed small prey. The small cornetfish (10–30 cm in total length) fed only on reef fish captured after stalking (where the fish slowly approaches the prey and then suddenly attacks). The stalking was done either solitarily or in foraging association with conspecifics. Large fish (30–120 cm) fed on both types of fishes by stalking and/or chasing (where the fish chases the prey using its high mobility and attacks), either solitarily or in foraging association with con- or heterospecifics. Thus, chasing was only performed by the large cornetfish against pelagic prey fish in associative foraging with other con- and heterospecific predators. As their body sizes increased, F. commersonii began to show a diversification of foraging behaviors, which was strongly related not only to the habitat types and anti-predatory behaviors of the prey fishes but also to associative foraging with con- or heterospecifics, which improves their foraging success.     

Predation of Japanese dace, Tribolodon hakonensis, by largemouth bass, Micropterus salmoides, in experimental aquaria

 To test the size range of prey fish that largemouth bass, Micropterus salmoides, can successfully consume, live Japanese dace, Tribolodon hakonensis, were given as prey fish to individual largemouth bass in aquaria. The ratio of maximum standard length (SL) of the Japanese dace consumed by largemouth bass was 46–69% of bass SL. The maximum length of Japanese dace consumed did not differ significantly between largemouth bass and smallmouth bass (M. dolomieu) previously studied, although largemouth bass have relatively larger mouth sizes than smallmouth bass. Largemouth bass occasionally injured and killed Japanese dace larger than the limit that could be consumed.