Antipredator Behavior of American Bullfrogs (Lithobates catesbeianus) in a Novel Environment

Invasive species capable of recognizing potential predators may have increased establishment rates in novel environments. Individuals may retain historical predator recognition and invoke innate responses in the presence of taxonomically or ecologically similar predators, generalize antipredator responses, or learn to avoid risky species in novel environments. Invasive amphibians in aquatic environments often use chemical cues to assess predation risk and learn to avoid novel predators via direct experience and/or associated chemical cues. Ontogeny may also influence recognition; experience with predators may need to occur at certain developmental stages for individuals to respond correctly. We tested predator recognition in invasive American bullfrog ( Lithobates catesbeianus) tadpoles that varied in experience with fish predators at the population and individual scale. We found that bullfrog tadpoles responded to a historical predator, largemouth bass ( Micropterus salmoides), only if the population was locally sympatric with largemouth bass. Individuals from a population that did not co‐occur with largemouth bass did not increase refuge use in response to either largemouth bass chemical cues alone or chemical cues with diet cues (largemouth bass fed bullfrog tadpoles). To test whether this behavioral response was generalized across fish predators, we exposed tadpoles to rainbow trout ( Oncorhynchus mykiss) and found that tadpoles could not recognize this novel predator regardless of co‐occurrence with other fish species. These results suggest that environment may be more important for predator recognition than evolutionary history for this invasive species, and individuals do not retain predator recognition or generalize across fish predators.     

Growth,survival, and foraging abilities of early life history stages of blue tilapia,Oreochromis aureus,and largemouth bass,Micropterus salmoides

Synopsis I compared growth, survival, and foraging abilities of early life history stages of blue tilapia,Oreochromis (=Sarotherodon = Tilapia) aureus, and largemouth bass,Micropterus salmoides, at equivalent prey (zooplankton) densities in the laboratory. Survival and growth of blue tilapia exceeded those of largemouth bass and, at comparable ages, blue tilapia were superior to largemouth bass in ability to prey on zooplankton; the enhanced survival and growth of blue tilapia appeared to result from their larger initial (egg) size. Predatory abilities of the species at equivalent sizes were similar. The success of blue tilapia in colonizing new systems and rapidly achieving high abundances in Florida may be a consequence of large initial size.     

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.     

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     

Gape Limitation, Prey Size Refuges and the Top–down Impacts of Piscivorous Largemouth Bass in Shallow Pond Ecosystems

Top–down control of phytoplankton biomass through piscivorous fish manipulation has been explored in numerous ecological and biomanipulation experiments. Piscivores are gape-limited predators and it is hypothesized that the distribution of gape sizes relative to distribution of body depths of prey fish may restrict piscivore effects cascading to plankton. We examined the top–down effects of piscivorous largemouth bass on nutrients, turbidity, phytoplankton, zooplankton and fish in ponds containing fish assemblages with species representing a range of body sizes and feeding habits (western mosquitofish, bluegill, channel catfish, gizzard shad and common carp). The experimental design consisted of three replicated treatments: fishless ponds (NF), fish community without largemouth bass (FC), and fish community with largemouth bass (FCB). Turbidity, chlorophyll a, cyclopoid copepodid and copepod nauplii densities were significantly greater in FC and FCB ponds than in NF ponds. However, these response variables were not significantly different in FC and FCB ponds. The biomass and density of shallow-bodied western mosquitofish were reduced and bluegill body depths shifted toward larger size classes in the presence of largemouth bass, but the biomass and density of all other fish species and of the total fish community were unaffected by the presence of largemouth bass. Our results show that top–down impacts of largemouth bass in ecosystems containing small- and deep-bodied fish species may be most intense at the top of the food web and alter the size distribution and species composition of the fish community. However, these top–down effects may not cascade to the level of the plankton when large-bodied benthivorous fish species are abundant.     

Density-dependent foraging behaviour of sheep on alpine pastures: effects of scale

Foraging patterns of large herbivores may give important clues as to why their life history varies depending on population density. In this landscape-scale experiment, domestic sheep Ovis aries were kept at high (80 sheep km⁻²) and low (25 sheep km⁻²) population densities during summer in high mountain pastures in Hol, Norway. We predicted an increasing use of less preferred plant species or habitat types with increasing sheep population density. Foraging behaviour was investigated by direct observations of individually marked sheep on different spatial scales, and diet composition was also assessed with microhistological analysis of faecal samples from known individuals. We found that the effects of density on foraging behaviour depended on scale and were only detected at the scale of diet choice. Use of the common grass species Deschampsia flexuosa , which provided the bulk forage (10–65% of the diet), remained constant throughout the season at low densities, but increased significantly over time at high densities. On a coarser spatial scale, neither within vegetation type nor between vegetation types, selection was affected by density, but vegetation type selection differed depending on whether the sheep were grazing or resting. Our study provides evidence of density dependence in foraging behaviour, but only at the finest spatial scale (diet choice).     

Does prey community composition affect the way different behavioral types interact with their environment?

We examined how different exploratory behavioral types of largemouth bass responded to differing prey communities by determining effects on growth, survival and diet in experimental ponds. We found evidence that non-explorer largemouth bass target young-of-year bluegill early on in life, but bluegill were not an important diet item by late summer. The presence of young-of-year bluegill as prey does appear to affect the foraging strategy of the two exploring types differently. In the absence of small bluegill, both behavioral types feed primarily on benthic invertebrates and zooplankton. When small bluegill were present, we saw a shift away from zooplankton as prey for largemouth bass. However, that shift was toward more benthic invertebrates for non-exploring behavioral types and toward terrestrial insects for exploring behavioral types. Thus, it appears that prey community composition can have important effects on the way in which different behavioral types interact with their environment.     

Nonlinear effect of density on trophic niche width and between‐individual variation in diet in a neotropical cichlid

Density‐dependent dynamics have considerable effects in many ecological processes and patterns that characterize natural populations. In the present study, we aim at evaluating the effect of density on the diet width and between‐individual variability in nine different pike cichlid Crenicichla lepidota populations dwelling in floodplain lagoons. Our results indicated that low‐density populations exhibit small diet breadth as well as small between‐individual variability in their diet. In addition, these populations were characterized by a nested diet pattern, where the diet of specialist individuals represented a subset of the food items consumed by generalist individuals. Populations with intermediate densities had a larger populational diet breadth while the individual diet breadth remained unchanged. This pattern was due to an increase in between‐individual variability in diet, which consequently decreases the diet overlap among individuals and thus lower diet nestedness. Finally, under high densities, the niche width at the population level decreased because of lower between‐individual variability and higher diet overlap. Together, these results showed that niche width exhibits a non‐linear function with density. At first, an increase in density increased the niche width because of greater between‐individual diet variability. However, after a threshold density value, the effect was reversed, and the niche width decreased because of a higher diet overlap among individuals.     

The energetic impact of overwinter prey assemblages on age-0 largemouth bass, <Emphasis Type="Italic">Micropterus salmoides</Emphasis>

Synopsis We compared survival, growth, and swimming performance of two size classes of age-0 largemouth bass, Micropterus salmoides, in the spring after being fed diets of bluegill, Lepomis macrochirus, fathead minnows, Pimephales promelas, or invertebrate prey during the winter. Regardless of prey assemblage, survival was uniformly high and independent of size. Length, wet- and dry-mass, and condition was also similar among treatments for both size classes. However, variation in individual performance differed, with the lowest variability in growth occurring among small age-0 largemouth bass in the invertebrate only treatment. Absolute and length corrected swimming speeds of largemouth bass were highest for invertebrate prey assemblages, intermediate for fathead minnow prey, and lowest for bluegill prey. The patterns in growth and spring swimming performance likely reflect the varied nutritive quality of different prey, the ability of largemouth bass to capture different prey, and competition with the piscine prey.     

Distribution and diet of Suwannee bass and largemouth bass in the lower Santa Fe River,Florida

Synopsis Suwannee bass,Micropterus notius, and largemouth bass,Micropterus salmoides, were collected by electrofishing in six habitats in the lower Santa Fe River, Florida during May 1981–March 1982. Both bass species were collected concomitantly in all habitats and habitat segregation was not evident. Crayfish (Procambarus spp.) were the primary food of Suwannee bass. Fish were the primary food of largemouth bass, but crayfish were common in the diet of largemouth bass ≥300 mm long. Suwannee bass have a greater throat width and consumed longer and wider forage than did largemouth bass of equal length. Available evidence suggests that Suwannee bass exhibit a positive selection for crayfish and a diverse forage resource, including abundant crayfish, is necessary for a Suwannee bass to coexist with a largemouth bass. This is Journal Series Number 6034 of the Florida Agricultural Experiment Station.     

Response of littoral invertebrates to reduction of fish density: simultaneous experiments in ponds with different fish assemblages

1. We experimentally reduced densities of predatory fish in replicated 2 m² areas of the littoral zone in two ponds to test whether density and biomass of invertebrates would respond to release from fish predation. The ponds are of similar size and in close proximity, but support different fish assemblages: bluegills ( Lepomis macrochirus Rafinesque) and largemouth bass ( Micropterus salmoides (Lacepede)) in one pond, and bluespotted sunfish ( Enneacanthus gloriosus (Holbrook)) and chain pickerel ( Esox niger Lesueur) in the other. Fish densities were reduced to less than 15% of ambient levels in both experiments.
2. In the bluegill–bass pond, density and biomass of most invertebrate taxa and size classes were unaffected by the fish manipulation. Total invertebrate densities did not differ significantly between fish treatments, but total invertebrate biomass was significantly greater where fish density was reduced, averaging 30% higher over the course of the study. Likewise, manipulation of fish in the bluespotted sunfish–pickerel pond had few significant effects on individual taxa and size classes. There were no significant effects on total invertebrate abundance in the bluespotted sunfish–pickerel pond.
3. Our results provide direct experimental evidence consistent with the collective evidence from previous work, suggesting that the impact of fish predation on density and biomass of invertebrate prey in littoral habitats is variable, but generally weak. Invertebrates that coexist successfully with fish in littoral systems probably are adept at taking advantage of refugia offered by the structurally complex physical environment.     

Response of littoral invertebrates to reduction of fish density: simultaneous experiments in ponds with different fish assemblages

1. We experimentally reduced densities of predatory fish in replicated 2 m² areas of the littoral zone in two ponds to test whether density and biomass of invertebrates would respond to release from fish predation. The ponds are of similar size and in close proximity, but support different fish assemblages: bluegills ( Lepomis macrochirus Rafinesque) and largemouth bass ( Micropterus salmoides (Lacepede)) in one pond, and bluespotted sunfish ( Enneacanthus gloriosus (Holbrook)) and chain pickerel ( Esox niger Lesueur) in the other. Fish densities were reduced to less than 15% of ambient levels in both experiments.
2. In the bluegill–bass pond, density and biomass of most invertebrate taxa and size classes were unaffected by the fish manipulation. Total invertebrate densities did not differ significantly between fish treatments, but total invertebrate biomass was significantly greater where fish density was reduced, averaging 30% higher over the course of the study. Likewise, manipulation of fish in the bluespotted sunfish–pickerel pond had few significant effects on individual taxa and size classes. There were no significant effects on total invertebrate abundance in the bluespotted sunfish–pickerel pond.
3. Our results provide direct experimental evidence consistent with the collective evidence from previous work, suggesting that the impact of fish predation on density and biomass of invertebrate prey in littoral habitats is variable, but generally weak. Invertebrates that coexist successfully with fish in littoral systems probably are adept at taking advantage of refugia offered by the structurally complex physical environment.     

Do the effects of piscivorous largemouth bass cascade to the plankton?

Ecologists have hypothesized that an increase in the biomass of piscivorous fish in lakes will cause a decrease in populations of planktivorous fish, an increase in the size of herbivorous zooplankton and a decrease in the biomass of phytoplankton. Here we present an experimental test of whether the effects of largemouth bass (Micropterus salmoides) cascade to the planktivorous fish, zooplankton and phytoplankton of a 15-ha water storage reservoir. A pilot study indicated that the reservoir was eutrophic with dense populations of planktivorous fish dominated by threadfin shad (Dorosoma petenense). No piscovorous fish were present in the reservoir. We conducted a one-month mesocosm experiment using water and plankton from the reservoir showing that the presence of threadfin shad reduced large-sized zooplankton and increased the productivity and biomass of phytoplankton. To test whether the effects of piscivorous fish could cascade to the plankton, we assessed the effects of the addition of piscivorous largemouth bass on the planktivorous fish, zooplankton and biomass of phytoplankton of the reservoir by monitoring the reservoir during the year before and the two years after largemouth bass were stocked. In the second year after the addition of largemouth bass, the number of planktivorous fish decreased and the relative abundance of threadfin shad declined. Although the abundance of cladocerans increased after the addition of largemouth bass, the average size of zooplankton did not change. We did not detect changes in chlorophyll a, Secchi depth, or concentrations of total phosphorus and total nitrogen as a result of the addition of largemouth bass.     

Interactions between bluefish and striped bass: Behavior of bluefish under size- and number-impaired conditions and overlap in resource use

A decline in bluefish (Pomatomus saltatrix L.) recreational landings during the 1990s and the early 2000s led to multiple theories on the ultimate cause. One theory was that a large portion of the bluefish population moved offshore and was unavailable to nearshore recreational fishers; one reason given for the movement offshore was increased competition with striped bass (Morone saxatilis W.). We conducted laboratory experiments (feeding and non-feeding) to examine behavioral interactions between adult bluefish and sub-adult striped bass in a large (121,000 L) research aquarium. Additionally, we examined diet and habitat overlap of bluefish and striped bass from the fall and spring bottom trawl surveys conducted by the National Marine Fisheries Service. Observations of feeding trials for the following treatments were made: non-impaired (i.e., same number and size of bluefish and striped bass), size-impaired (i.e., large striped bass/small bluefish), number-impaired (i.e.,10 striped bass/3bluefish), and single-species controls. Within a species, there was no difference in a variety of behavioral measures (e.g., attack rate, capture success, ingestion rate, and activity) between mixed- and control treatments under non-impaired or size-impaired conditions. However, behavior of number-impaired bluefish differed from control and size-impaired fish suggesting that striped bass may have a negative influence on bluefish foraging when bluefish are “out-numbered”. Feeding had a strong effect on swimming speeds for both species. Diet and habitat overlap between bluefish and striped bass in continental shelf waters was low. Overall, foraging behavior in mixed-species treatments and field observations suggest no competitive interactions between adult bluefish and sub-adult striped bass.     

Above-ground competition does not alter biomass allocated to roots in Abutilon theophrasti

We tested whether plants allocate proportionately less biomass to roots in response to above-ground competition as predicted by optimal partitioning theory. Two population densities of Abutilon theophrasti were achieved by planting one individual per pot and varying spacing among pots so that plants in the two densities experienced the same soil volume but different degrees of canopy overlap. Density did not affect root:shoot ratio, the partitioning of biomass between fine roots and storage roots, fine root length, or root specific length. Plants growing in high density exhibited typical above-ground responses to neighbours, having higher ratios of stem to leaf biomass and greater leaf specific area than those growing in low density. Total root biomass and shoot biomass were highly correlated. However, storage root biomass was more strongly correlated with shoot biomass than was fine-root biomass. Fine-root length was correlated with above-ground biomass only for the small subcanopy plants in crowded populations. Because leaf surface area increased with biomass, the ratio between absorptive root surface area and transpirational leaf surface area declined with plant size, a relationship that could make larger plants more susceptible to drought. We conclude that A. theophrasti does not reallocate biomass from roots to shoots in response to above-ground competition even though much root biomass is apparently involved in storage and not in resource acquisition.     

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.     

Individual Growth and Foraging Responses of age-0 Largemouth Bass to Mixed Prey Assemblages during Winter

We conducted an outdoor pool experiment at a mid-temperate latitude (Ohio, 40°N) to determine how commonly occurring prey assemblages affect individual foraging and growth of individually marked, age-0 largemouth bass during winter. The treatments were low prey, bluegill prey only, macroinvertebrates only, and bluegill plus macroinvertebrates. Across all treatments, growth in mass (g) was unrelated to body size. Conversely, small individuals lost more energy (kJ) than large counterparts in all but the macroinvertebrate-only treatment. With low prey, overall growth of largemouth bass was negative, with losses varying among individuals by 30% and 60% for mass and energy content, respectively. Counterparts in bluegill-only pools also consistently lost mass and energy, with less variability (15% mass; 30% energy). In the macroinvertebrate-only treatment, 31% of individuals gained mass, reflecting the greatest range in mass (100%) and energy (60%) change. With macroinvertebrates plus bluegill, overall growth was generally negative, with intermediate variance among individuals. Variation in growth among individuals typically increased with the frequency that prey occurred in diets during sampling. Apparently, some individuals were inactive, foraged infrequently, and consistently lost intermediate quantities of mass and energy. Others were active and foraged with variable success. Because activity and growth vary among individuals as a function of prey composition during winter, prey assemblages during this season will affect patterns of first-year survival and cohort strength.     

Determination of Foraging Thresholds and Effects of Application on Energetic Carrying Capacity for Waterfowl

Energetic carrying capacity of habitats for wildlife is a fundamental concept used to better understand population ecology and prioritize conservation efforts. However, carrying capacity can be difficult to estimate accurately and simplified models often depend on many assumptions and few estimated parameters. We demonstrate the complex nature of parameterizing energetic carrying capacity models and use an experimental approach to describe a necessary parameter, a foraging threshold (i.e., density of food at which animals no longer can efficiently forage and acquire energy), for a guild of migratory birds. We created foraging patches with different fixed prey densities and monitored the numerical and behavioral responses of waterfowl (Anatidae) and depletion of foods during winter. Dabbling ducks (Anatini) fed extensively in plots and all initial densities of supplemented seed were rapidly reduced to 10 kg/ha and other natural seeds and tubers combined to 170 kg/ha, despite different starting densities. However, ducks did not abandon or stop foraging in wetlands when seed reduction ceased approximately two weeks into the winter-long experiment nor did they consistently distribute according to ideal-free predictions during this period. Dabbling duck use of experimental plots was not related to initial seed density, and residual seed and tuber densities varied among plant taxa and wetlands but not plots. Herein, we reached several conclusions: 1) foraging effort and numerical responses of dabbling ducks in winter were likely influenced by factors other than total food densities (e.g., predation risk, opportunity costs, forager condition), 2) foraging thresholds may vary among foraging locations, and 3) the numerical response of dabbling ducks may be an inconsistent predictor of habitat quality relative to seed and tuber density. We describe implications on habitat conservation objectives of using different foraging thresholds in energetic carrying capacity models and suggest scientists reevaluate assumptions of these models used to guide habitat conservation.     

Parental care and the ontogeny of predator-avoidance in two species of centrarchid fish

Stamps (1978) suggested that the type and extent of parental care can influence the behavioural ontogeny of the offspring. Largemouth bass (Micropterus salmoides) males guard the fry for up to a month after they leave the nest while rock bass (Ambloplites rupestris) males guard the fry only until they leave the nest. In laboratory experiments, naive largemouth fry displayed reduced predator-avoidance responses to large and small predators during their first 3 weeks of free-swimming. Rock bass fry, on the other hand, avoided predators throughout the study period. From 6 weeks (largemouth) and 5 weeks (rock bass) of age the fry showed a significantly greater response to the large predator than the small predator. The responses of laboratory-reared largemouth bass fry to the small predator changed from avoidance to association as the fry grew. The size of the fry relative to predator size was found to be important.