Habitat use and diet of two stream gobies of the genus Rhinogobius in south-western Shikoku,Japan

The importance of interspecific competition to habitat use by two congeneric stream gobies, Rhinogobius sp. LD (large-dark type) and CB (cross-band type), was studied by: (i) examining differences in habitat use by each species along the course of the stream; and (ii) comparing microhabitat use and the diet of each species between in allopatry and in sympatry in tributaries of the Shimanto River, south-western Shikoku, Japan. Rhinogobius sp. LD mainly used riffles through the course of the stream. Although CB also mainly used riffles in the lower reaches where CB was numerically dominant, the greater use of riffles by CB was not observed in upper reaches where LD was dominant. Microhabitat analysis revealed that both LD and CB preferred lower bottom velocity in faster current in both allopatry and sympatry. In sympatry, however, LD used coarser substrate and faster current than CB, and both LD and CB used narrower ranges of microhabitat conditions than in allopatry. Dietary analysis indicated that both allopatric and sympatric LD preferred baetids, which are apt to drift, suggesting they adopted ambush foraging. Allopatric CB preferred leptophlebiids and chironomids, which are also apt to drift, whereas sympatric CB did not prefer such invertebrates. Overall results of this study suggest that LD and CB compete for better feeding habitats, CB shift their habitat use and foraging mode under the influence of LD, and current velocity and substrate coarseness are key factors in their habitat segregation.

     

Ambush Predation by the Ponerine Ant Ectatomma ruidum Roger (Formicidae) on a Sweat Bee Lasioglossum umbripenne (Halictidae), in Panama

An individual ambush predation is used by huntresses of the ponerine ant Ectatomma ruidum to capture halictid bees (Lasioglossum umbripenne) in the Panamanian mountains. Workers, which use this strategy and originated from a colony (A) situated within a nest aggregation of halictid bees, capture almost four times more prey than colony (B) with a foraging area which does not include this type of bee's nest. Forty-eight percent of the prey of colony (A) are halictid bees, demonstrating the local importance of this predatory strategy in E. ruidum. A close examination of the behavioral sequence of predation shows that ambush is successful in only 4.8% of cases, a very low success rate in comparison to other predatory strategies described in this species. Nevertheless, due to the high repetition (16.1 ± 5.9 times on average) of very short waiting phases (6.3 ± 1.9 s), the true success rate (i.e., according to the number of individual trips) can reach up to 80%. A review of ambush predation among ant species allows us to distinguish approaches between jumping, long stealth approaches, and true ambushes. Implications of learning and expectation processes are also discussed.     

Short-term cues used by foraging trout in a California stream

Stream salmonids choose foraging locations to maximize the energy benefit of foraging within the constraints of size-mediated dominance hierarchies and predation risk. But, because stream habitats are temporally variable, fish must use a search process to monitor changing habitat conditions as a means of locating potentially-better foraging locations. I explored the cues used by the cutthroat trout, Oncorhynchus clarki clarki, when searching for food at the pool scale by artificially increasing prey availability at different locations by using special feeders and by manipulating pool velocities. Behavior of individually marked fish was monitored from stream bank platforms under unmanipulated control conditions and under seven experimental sets of conditions involving different combinations of feeder location and velocity manipulation. Under natural conditions fish elected to forage in the deepest (>50 cm), fastest (0.10–0.25 m s⁻¹) locations and within 1 m of structure cover, but would readily move to shallower (<30 cm) water away from cover if velocities were manipulated to be highest there. Although fish did not locate feeders unless they were placed in high-velocity areas, when high velocity was provided fish would move into very shallow water (<20 cm) if prey were delivered there. Responses of individual trout to manipulations indicated that water velocity was the main physical cue used by fish to decide where to forage, and that fish could also learn about new food sources by observing conspecifics. Overall, results indicated fish were not “perfect searchers” that could quickly locate new food resources over short time scales, even when the new resources were within a few meters of the fish’s normal foraging location. When given the correct cues, however, fish could detect new food sources and defend them against subordinate fish. Movement of new fish into and out of the study pools during the ten-day observation period was common, consistent with the idea that trout used movement as a means of exploring and learning about habitat conditions at the reach scale.     

Competition between two congeneric stream gobies for habitat in southwestern Shikoku, Japan

Competition between two congeneric gobies, Rhinogobius sp. LD (large-dark type) and CB (cross-band type), for habitat was studied in a tributary of the Shimanto River, southwestern Shikoku, Japan. Habitat use by CB, measured by water depth, current velocity, and substrate, was compared between before and after the removal of LD. After the removal of LD, CB shifted their habitat use to coarser substrates than previously, suggesting a release from effects of LD. This result supports a hypothesis from our previous study that habitat partitioning between CB and LD is a consequence of interference effects of LD on CB.     

Comparison of current-velocity tolerance among six stream gobies of the genus Rhinogobius

Tolerance to current velocity was compared among six stream gobies of the genus Rhinogobius (cross-band, dark, large-dark, cobalt, and orange types, and R. flumineus). Each individual of the six gobies was exposed to incremental increases in current velocity within a laboratory flume. The current velocity at which each individual was dislodged was recorded and compared by species. Results from comparisons showed that the tolerance of gobies declined in the following order: cobalt, large-dark, dark, cross-band, R. flumineus, and orange type. This interspecific difference in velocity tolerance is consistent with their habitat segregation patterns as previously reported by field observations.     

Behavior and distribution of upstream-migrating juvenilerhinogobius sp. (The orange form)

Upstream-migration of juvenileRhinogobius sp. OR was studied along the Ado River, a tributary of Lake Biwa. Juvenile gobies, mainly 18–21 mm in standard length, migrated from the lake in a belt of about 3 m wide in lower reaches and about 10–20 cm wide in middler reaches. Their upstream-migration was restricted to places shallower than 20 cm with bottom current velocities less than 20 cm/sec. Individual migration speed varied between 4.2–8.9 m/min at places with bottom current velocity of 8.8–20.0 cm/sec, slowing down with increasing current velocity. Most active migration occurred between 14:00 and 16:00, number of upstream-migrating juveniles decreasing with distance from the river mouth. Riffles in the mid-and upper reaches, having few slow current areas, probably present barriers to upstream-migration of the juvenile gobies, although both adults and subadults of the species are distributed in more upper reaches.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Genetic variations estimated from PCR-RFLP analysis of two morphs of the freshwater goby <Emphasis Type="Italic">Rhinogobius</Emphasis> in the Lake Biwa water system

The freshwater goby Rhinogobius is the most abundant fish in the shore area of Lake Biwa, Japan. Recently, it has been reported that two morphs of Rhinogobius inhabit this lake. In this study, genetic variations in Rhinogobius sp. OR (Orange-type) and Rhinogobius sp. BW (Biwa-type) in the Lake Biwa water system have been investigated using polymerase chain reaction-based restriction fragment length polymorphism (PCR-RFLP) analysis of mitochondrial DNA, including the variable D-loop region. Samples of Rhinogobius sp. OR were collected from the middle sites of three rivers, two outlets, and two lakeshore sites, whereas samples of Rhinogobius sp. BW were taken from two lakeshore sites. Rhinogobius sp. OR and Rhinogobius sp. BW did not share any haplotypes, suggesting that PCR-RFLP analysis is effective for distinguishing between these species of goby. The haplotype diversity of Rhinogobius sp. OR (0.214–0.543) was lower than that of Rhinogobius sp. BW (0.543–0.682). There were no significant differences in haplotype frequencies between Rhinogobius sp. BW groups from the two localities. In addition, haplotype frequencies in Rhinogobius sp. OR did not differ significantly among samples from the middle sites of rivers, the outlets, and the shores. These results indicate that in Rhinogobius sp. OR there is frequent gene flow among fish inhabiting different sites, and that this species of goby consists of a single population throughout the Lake Biwa water system.     

Modelling the Effects of Current on Prey Acquisition in Planktivorous Fishes

Many planktivorous fishes forage in currents, where they actively maintain position and visually strike at current-entrained zooplankton. In general, the zooplankton are wafted by the foraging fish at a rate equivalent to the current velocity. From a fish's viewpoint the plankton approach either head-on or offset at varied distances from the fish's position. We present a model that describes the relative motion of particles as they approach and pass a foraging fish at different offset distances, and the rate of change in apparent size as they close on a fish. In addition, a series of experiments of fish feeding on plankton in a flume at increasing current velocities revealed that two basic tactics are utilized. At low current velocities (<10-14 cm s m 1), the fish swims toward the prey, whereas at higher current velocities the fish tends to fall back with the current to capture a prey item. The model and experimental results are discussed in terms of the visual problems associated with the detection and tracking of items in motion.     

Ecological aspects of ontogenetic shifts in prey size utilization in the bay goby (Pisces: Gobiidae)

Summary Two age classes (0+ to 3+ and 4+ to 7+) of bay gobies (Lepidogobius lepidus Girard) differed in the sizes of whole prey (except polychaetes) recovered from the digestive tract. Although older fish consumed greater amounts of larger prey they did not capture larger individuals of a given prey type for seven of nine prey classes. The remaining two, harpacticoid copepods and ostracods contribute minimally to ontogenetic differences. The switch to larger prey appears to either decrease exposure to predation through a reduction in foraging time or increase energy intake. This is facilitated through older fishes greater size and/or superiority in intraspecific competition. These differences can probably be viewed as adaptations to selective pressures imposed by fluctuating food resources on predation levels in bays and estuaries.     

Efficiency evaluation of two competing foraging modes under different conditions

Various foraging modes are employed by predators in nature, ranging from ambush to active predation. Although the foraging mode may be limited by physiological constraints, other factors, such as prey behavior and distribution, may come into play. Using a simulation model, we tested to what extent the relative success of an ambush and an active predator changes as a function of the relative velocity and movement directionality of prey and active predator. In accordance with previous studies, we found that when both active predator and prey use nondirectional movement, the active mode is advantageous. However, as movement becomes more directional, this advantage diminishes gradually to 0. Previous theoretical studies assumed that animal movement is nondirectional; however, recent field observations show that in fact animal movement usually has some component of directionality. We therefore suggest that our simulation is a better predictor of encounter rates than previous studies. Furthermore, we show that as long as the active predator cannot move faster than its prey, it has little or no advantage over the ambush predator. However, as the active predator's velocity increases, its advantage increases sharply.     

Interactions of multiple predators with different foraging modes in an aquatic food web

Top predators can have different foraging modes that may alter their interactions and effects on food webs. Interactions between predators may be non-additive resulting from facilitation or interference, whereas their combined effects on a shared prey may result in emergent effects that are risk enhanced or risk reduced. To test the importance of multiple predators with different foraging modes, we examined the interaction between a cruising predator (largemouth bass, Micropterus salmoides) and an ambush predator (muskellunge, Esox masquinongy) foraging on a shared prey (bluegill sunfish, Lepomis macrochirus) with strong anti-predator defense behaviors. Additive and substitution designs were used to compare individual to combined predator treatments in experimental ponds. The multiple predator interaction facilitated growth of the cruising predator in the combined predator treatments, whereas predator species had substitutable effects on the growth of the ambush predator. The combined predator treatments created an emergent effect on the prey; however, the direction was dependent on the experimental design. The additive design found a risk-reducing effect, whereas the substitution design found a risk-enhancing effect for prey fish. Indirect effects from the predators weakly extended to lower trophic levels (i.e., zooplankton community). Our results highlight the need to consider differences in foraging mode of top predators, interactions between predators, and emergent effects on prey to understand food webs.     

Utilizing stomach content and faecal DNA analysis techniques to assess the feeding behaviour of largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus

In this study, the feeding behaviour of the non‐native invasive predatory fishes largemouth bass Micropterus salmoides and bluegill Lepomis macrochirus was studied in the Ezura River, a northern tributary of Lake Biwa, Japan. Prey composition was estimated based on visual examination of stomach contents and faecal DNA analysis to determine feeding habits of these predatory fishes. Stomach content analysis showed that native fishes (e.g. ayu Plecoglossus altivelis and gobies Rhinogobius spp.) and shrimps (e.g. Palaemon paucidens) were the major prey items for M. salmoides, while snails, larval Chironomidae and submerged macrophytes were the dominant prey items of L. macrochirus. Micropterus salmoides tended to select larger fish in the case of crucian carp Carassius spp., but smaller fishes in the case of P. altivelis and Rhinogobius spp. Faecal DNA analyses revealed prey compositions similar to those identified in predator stomach contents, and identified additional prey species not detected in stomach content inspection. This study demonstrated that both stomach content inspection and DNA‐based analysis bear several inherent shortcomings and advantages. The former method is straightforward, although identification of species can be inaccurate or impossible, whereas the latter method allows for accurate species identification, but cannot distinguish prey size or stage. Hence, integration of morphology‐based and DNA‐based methods can provide more reliable estimates of foraging habits of predatory fishes.     

Competitive food exploitation of smelt Osmerus eperlanus by great crested grebes Podiceps cristatus and perch Perca fluviatilis at Lake IJsselmeer,The Netherlands

The impact of predation by an avian predator (great crested grebe Podiceps cristatus) and a predatory fish (perch Perca fluviatilis) on a common resource, the small planktivorous fish smelt (Osmerus eperlanus) was studied in Lake IJsselmeer, The Netherlands, in July–October 1985–1988. At this time of the year the grebes are in simultaneous wing-moult and therefore temporarily flightless. Four factors limit the predation pressure by grebes: (1) an underwater visibility threshold of c. 40 cm Secchi depth is the lower limit at which large numbers attend the moulting site, (2) a lower density threshold of exploitable smelt biomass of c. 30 kg/ha determines the actual fishing areas, (3) a fish size threshold of 6.5 cm total length further limits prey availability, and (4) the vertical movements of the prey impose a strong (diel) time constraint on the birds, allowing only crepuscular foraging. Perch, the other main predator in this system, is less restricted in its foraging. Perch were found to be in direct competition with the grebes, preying heavily on the juvenile smelt. Over the 4 years of study 90% of all predation from 15 July to 15 October (72–233 kg/ha) was due to perch. The high level of predation by birds and fish was only possible due to a continuous immigration of smelt into the area (1.45 kg/ha per day). In three of the four years, however, the combined predation by fish and birds surpassed the immigration rate of the prey, which led to a strong reduction in smelt stocks in the study area. A conceptual model is developed to describe the different sets of constraints on the grebes' foraging. First, the state of eutrophication in relation to the weather condition determines the degree of algal blooms, and thus underwater visibility, in late summer. This is the major factor governing the numbers of grebes on the moulting area. Second, the size of the population of predatory fish determines the overall food availability (biomass and size distribution of smelt). These factors are partly interconnected and related to human action (pollution, fisheries). It is suggested that, despite deteriorating visibility conditions, the largest moulting site for grebes in Europe at Lake IJsselmeer exists because the stocks of predatory fish are kept low by overfishing.     

Variation in male mating success in the paternal nest brooder, Rhinogobius goby, along a river course

Rhinogobius sp. CB (cross band type) is an amphidromous freshwater goby which receives paternal nest care under stones on river beds. This goby is known to migrate to the upper reaches of rivers as it grows. In the present study, the relationships among male mating success of Rhinogobius sp. CB, male body size, the size frequency distribution of river bed stones and the presence of the sympatric goby Tridentiger brevispinis (which uses nest sites similar to those of Rhinogobius sp. CB) were investigated along the course of the Aizu River, Wakayama Prefecture, Japan. In the lower reach, where T. brevispinis was present and large stones were scarce, the sizes of the nest stones and the egg clusters of male Rhinogobius sp. CB were smaller than those of T. brevispinis. In the middle and the upper reaches, where T. brevispinis was absent and large stones were abundant, males of Rhinogobius sp. CB used larger nest stones than in the lower reach. In both gobies, there were positive correlations among male body size, nest size and egg cluster size in all reaches. Nest-choice experiments carried out in the laboratory, with or without a rival, showed that males of both gobies preferentially used large nest sites and that large males had an advantage in male-male competition for large nest sites. These results indicate that large male body size can increase male mating success and that male mating success increases in the upper reach in this goby. The migration pattern of Rhinogobius sp. CB was also discussed in relation to these findings.     

Influence of Predator Presence and Prey Density on Behavior and Growth of Damselfly Larvae (Ischnura elegans) (Odonata: Zygoptera)

Foraging behavior is often determined by the conflicting benefits of energy gain and the risk of mortality from predation or other causes. Theory predicts that animals should have lower activity levels when either the risk of predation or the availability of resources in the environment is high. We investigated the adjustment of the behavior of I. elegans larvae to predator presence (Anax imperator) and prey density (Daphnia sp.) and their interaction in a completely crossed factorial experiment in the lab and the effect of behavior on growth. The foraging activity of the I. elegans larvae was significantly reduced in the presence of a free-swimming predator but not a caged predator. Abdominal movements were significantly reduced at a low prey density. Growth was significantly reduced by the presence of a free swimming predator and low prey densities. These results provide evidence that these damselfly larvae adjust their behavior to the presence of predators to increase their survival at the expense of reduced growth and development.     

Linking piscivory to spatial–temporal distributions of pelagic prey fishes with a visual foraging model

A visual foraging model (VFM) used light-dependent reaction distance and capture success functions to link observed prey fish abundance and distribution to predation rates and the foraging performance of piscivorous cutthroat trout Oncorhynchus clarki in Lake Washington (WA, U.S.A.). Total prey density did not correlate with predation potential estimated by the foraging model for cutthroat trout because prey were rarely distributed in optically favourable conditions for detection. Predictions of the depth-specific distribution and timing of cutthroat trout foraging were qualitatively similar to diel stomach fullness patterns observed in field samples. Nocturnal foraging accounted for 34–64% of all prey fish consumption in simulations for 2002 and 2003. Urban light contamination increased the access of nocturnally foraging cutthroat trout to vertically migrating prey fishes. These results suggest that VFMs are useful tools for converting observed prey fish density into predictions of predator consumptions and behavioural responses of predators to environmental change.     

Foraging time and spatial patterns of predation in experimental populations

Summary Responses of the predaceous mites Phytoseiulus persimilis, Typhlodromus (=Metaseiulus) occidentalis, and Amblyseius andersoni to spatial variation in egg density of the phytophagous mite, Tetranychus urticae, were studied in the laboratory.The oligophagous predator P. persimilis showed initially a direct density dependent foraging time allocation and variation in foraging time increased with prey density. With changes in prey density due to predation, predator foraging rates (per hour) decreased with time and density dependent foraging gradually became density independence, because P. persimilis continued to respond to initial prey density, instead of the changing prey density and distribution. The consequent spatial pattern of predation by P. persimilis was density independent, although slopes of predation rate-prey density regressions increased with time.Compared with P. persimilis, the narrowly polyphagous predator T. occidentalis responded relatively slowly to the the presence or absence of prey eggs but not to prey density: the mean and variation of foraging time spent in patches with prey did not differ with prey density, but was significantly greater in patches with prey eggs than in patches without eggs. Prey density and distribution changed only slightly due to predation and overall foraging rates remained more or less constant. The consequent spatial pattern of predation by T. occidentalis was inversely density dependent. As with P. persimilis, slopes of predation rate-prey density regressions increased with time (i.e. the inverse density dependence in T. occidentalis became weaker through time).The broadly polyphagous predator A. andersoni showed density independent foraging time allocation with variation independent of prey density. With changes in prey density over time due to prey depletion, overall foraging rates decreased. The consequent spatial pattern of predation by A. andersoni also changed through time; it initially was inversely density dependent, but soon became density independent.Overall, P. persimilis and T. occidentalis spent more time in prey patches than A. andersoni, suggesting that A. andersoni tended to spend more time moving outside patches. The overall predation rates and searching efficiency were higher in P. persimilis than in A. andersoni and T. occidentalis. Predator reproduction was highest in P. persimilis, lower in T. occidentalis and the lowest A. andersoni.The differences in response to prey distribution among the three predaceous species probably reflect the evolution of these species in environments with different patterns of prey distribution. The degree of polyphagy is a major determinant of the aggregative response, but other attributes such as handling time are also important in other aspects of phytoseiid foraging behavior (e.g. searching efficiency or predation rate).     

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: selective feeding of fish and predation effects on the zooplankton communities

Bag-type enclosures (75 m³) with bottom sheets and tube-type enclosures (105 m³) open to the bottom sediment were stocked with exotic whitefish (Coregonus lavaretus maraena) to study their predation effects on the plankton community. The fish fed mainly on adult chironomids during the period of their emergence (earlier part of the experimental period). Thereafter, the food preference was shifted to larvae of chironomids and crustacean zooplankters. The predation effects on the plankton community were not evident in the bag-type enclosures where zooplankton densities were consistently low. The fish reduced the crustacean populations composed ofBosmina fatalis, B. longirostris andCyclops vicinus in the tube-type enclosures where the prey density was high (above ca. 50 individuals 1⁻¹). The results suggested that the intensity of predation depended on the prey density. Rotifers increased in the fish enclosure, probably becauseCoregonus reduced the predation pressure byCyclops vicinus on rotifers and allowed the latter to increase. In the fish enclosures, no marked changes in species composition were observed. Zooplankton predated by the fish seemed to be distributed near the walls of the enclosures. Problems of enclosure experiments for examining the effects of fish predation on pelagic zooplankton communities are discussed.     

Variation in foraging success among predators and its implications for population dynamics

The effects of the expected predation rate on population dynamics have been studied intensively, but little is known about the effects of predation rate variability (i.e., predator individuals having variable foraging success) on population dynamics. In this study, variation in foraging success among predators was quantified by observing the predation of the wolf spider Pardosa pseudoannulata on the cricket Gryllus bimaculatus in the laboratory. A population model was then developed, and the effect of foraging variability on predator–prey dynamics was examined by incorporating levels of variation comparable to those quantified in the experiment. The variability in the foraging success among spiders was greater than would be expected by chance (i.e., the random allocation of prey to predators). The foraging variation was density‐dependent; it became higher as the predator density increased. A population model that incorporates foraging variation shows that the variation influences population dynamics by affecting the numerical response of predators. In particular, the variation induces negative density‐dependent effects among predators and stabilizes predator–prey dynamics.