Feeding by marine fish larvae: developmental and functional responses

Synopsis The relationship between prey consumption rate and prey concentration (functional response), and its change with growth (developmental response) were examined in the laboratory for three species of marine fish larvae: bay anchovy Anchoa mitchilli (Engraulidae), sea bream Archosargus rhomboidalis (Sparidae) and lined sole Achirus lineatus (Soleidae). The major objective was to determine relative predatory abilities of the larvae by fitting feeding rate data to developmental and functional response models. Feeding success, prey capture success, attack rates, handling times and search rates were estimated. Prey consumption rates and attack rates of bay anchovy usually were highest, but at the lowest prey level (50 per liter) first-feeding sea bream larvae had the highest consumption rate. Sea bream could consume prey at near-maximum rates at prey levels lower than those required by the other species. As larvae grew, time searching per attack decreased rapidly for all species, especially at low prey levels. Handling time also decreased, but most rapidly for bay anchovy. Search rates were highest for bay anchovy and lowest for lined sole. Bay anchovy had the best apparent predation ability, but when previous results on larval growth rates, survival rates and growth efficiencies were considered, sea bream larvae were the most efficient predators and the least likely of the three species to be limited by low prey levels.     

Habitat use and feeding behavior of thirteen species of benthic stream fishes

Synopsis The densities, habitat use, and feeding behaviors of 13 fish species belonging to the benthic invertebrate-feeding guild were studied by snorkeling at three localities in the Little River of eastern Tennessee, U.S.A. Resource partitioning occurred by habitat, feeding behavior and time of activity. Differences were also found at the generic level.Cottus was a nocturnal feeder, whereasPercina andEtheostoma were, for the most part, diurnally active.Percina moved about rapidly and spent most of its time above the bottom. In contrast,Etheostoma varied considerably in the amount of time spent under cover, spent little time above the bottom, and exhibited low levels of swimming activity. Nearly all species sought cover at night, suggesting they may be particularly sensitive to predation at night. Species with small adult sizes (Etheostoma, Cottus andP. evides) were concentrated in shallow water habitats, whereas species with large adult sizes (Percina) were more abundant in deep water habitats. The habitat use data are consistent with the hypothesis that size-selective predation by centrarchid bass may cause smaller fish to avoid deep water areas. Large species should have a lower risk of predation due to their size and behavior.     

Feeding and fate of wild larval razorback sucker

The razorback sucker (Xyrauchen texanus) is disappearing throughout its native range in the Colorado River basin of western North America. The largest remaining wild population in Lake Mohave, Arizona-Nevada, has shown no recruitment since the 1950s. Although annual spawning is successful and larvae are seasonally abundant, no juveniles have been collected in recent decades. To evaluate the potential role of food availability in determining fate of larvae, fish and zooplankton samples were taken in 1985 from the reservoir and an adjacent, isolated backwater in which larvae were naturally produced. Food availability and primary dietary constituents were similar in both habitats. Reservoir larvae selectedBosmina spp. (Cladocera) and apparently avoided Copepoda, while larvae from the backwater selectedBosmina, but avoided Rotifera. Larvae from both places showed evidence of selection for certain sizes of zooplankters, but preferred sizes differed between habitats. These differences were neither attributable to larval size nor zooplankton community structure. Nutritional factors such as type, number, or size of available foods do not explain disappearance of larval razorback suckers from Lake Mohave, since larvae survive to far greater ages and size in the backwater. Predation by introduced fishes appears a significant cause of larval mortality.     

Intra- and inter-specific variation in buoyancy of some estuarine fishes

Synopsis Buoyancy was measured on eight species of estuarine fishes that were caught in 1 m depth or less. Mean buoyancies of the physoclists Fundulus heteroclitus, F. majalis, Cyprinodon variegatus and Leiostomus xanthurus were similar and ranged from –6.5 to –18.0 kiloPascals below atmospheric pressure at sea level. Menidia menidia and Pomatomus saltatrix measured –36.6 and –46.1 kPa, respectively. Two physostomes, Brevoortia tyrannus and Anchoa mitchilli, measured + 2.9 and –23.5 kPa, respectively, but the latter probably releases air when handled.The four most buoyant physoclist species live near the bottom in areas that receive daily tide induced currents. Negative buoyancy probably functions in them as in stream dwelling minnows and salmonids, which respond to currents by decreasing their buoyancy. The pronounced negative buoyancy of M. menidia may be a response to a preference for habitat where the currents are stronger, P. saltatrix, which can secrete gas into the swim bladder at the fastest rate known for any fish, combines high secretion (and resorption) rates with marked negative buoyancy. This enables it to quickly change depths over a wide vertical range, without overexpanding the swim bladder to cause positive buoyancy.     

Foraging modes of stream benthic fishes in relation to their predation effects on local prey density

Habitat use and foraging behavior of two benthic insectivorous gobies, Rhinogobius sp. CO (cobalt type) and Rhinogobius sp. DA (dark type), were examined in relation to their predation effects on local prey density in a small coastal stream in southwestern Shikoku, Japan. Correlations among the foraging range, frequency of foraging attempts and current velocity indicated that individuals using fast-current habitats had small foraging ranges and infrequently made foraging attempts while those in slow currents frequently foraged over large areas. The former and the latter were recognized as ambush and wandering foragers, respectively. Interspecific comparisons of habitat use, foraging behavior and prey preference suggested that Rhinogobius sp. CO selectively forage mobile prey by ambushing in fast currents, whereas Rhinogobius sp. DA randomly forage available prey by wandering in slow-current habitats. A cage experiment was conducted to assess prey immigration rate and the degree of predation effects on local prey density in relation to current velocity. The results of the experiment support, at least in part, our initial predictions: (1) prey immigration rates increase with current velocity and (2) the effects of fish predation on local prey density are reduced as current velocity increases. Overall results illustrated a link between the foraging modes of the stream gobies and their predation effects on local prey density: fish adopt ambush foraging in fast currents, where the decrease in prey density tends to be less, whereas fish actively forage over large areas in slow currents, where the decrease in prey is relatively large.     

Occupation of submerged aquatic vegetation by fishes: testing the roles of food and refuge

Summary We conducted a series of field experiments to examine the roles of refuge and food availability in explaining the distribution and abundance of fish in tidal freshwater marsh creeks. Two hypotheses were tested: (1) relative predation pressure is less in SAV than in unvegetated areas and (2) fish food availability is greater in SAV than in nearby unvegetated areas. Tethering experiments using mummichogs (Fundulus heteroclitus) in vegetated and unvegetated areas revealed that relative predation pressure was significantly less in areas with SAV. Banded killifish (Fundulus diaphanus) maintained in vegetated enclosures consumed prey associated with SAV, whereas those held in unvegetated pens had empty stomachs. No differences were found in the number of prey eaten by bluespotted sunfish (Enneacanthus gloriosus) or mummichogs when confined in vegetated or unvegetated enclosures. However, larger prey were consumed by bluespotted sunfish and mummichogs maintained in vegetated enclosures. These data suggest that foraging profitability is significantly enhanced by feeding in the SAV. Submerged plant beds in tidal freshwater marsh creeks not only afford protection from predators, but also provide a rich foraging habitat. By foraging in SAV, fish consume larger prey and may have higher growth rates, lower mortality, and higher fecundity.     

Changing handling times during feeding and consequences for prey size selection of 0+ zooplanktivorous fish

The interrelationship of fish size, prey size and handling time within a 15-min feeding period was studied in three size groups of 0 + roach, Rutilus rutilus, and bleak, Alburnus alburnus. Four size classes of cladoceran prey were used to measure changes in feeding rate and handling time from initial rapid feeding to sustained feeding. Observed differences in increase of handling time between prey size classes led to a change in the prey profitability ranking of those size classes within the first 2 min of the experiments. A 1-min feeding period is interpreted as reflecting an intermediate motivational status between extreme hunger and satiation. The use of average handling times for this period revealed a substantial change in prey profitability estimates compared to previous studies which used handling times based on short-term (a few seconds up to 1 min) feeding. It is not the largest prey items a fish can handle and swallow that are most profitable, but prey of intermediate size. By this approach a closer fit between expectations derived from optimal foraging theory and empirical data on prey size selection of 0 + zooplanktivorous fish is qualitatively achieved. Optimal prey size was found to be close the mouth gape width in small fish of 15 mm standard length, decreasing to 50% of mouth gape width in fish of 40 mm standard length.     

Differential zooplankton feeding behaviors,selectivities, and community impacts of two planktivorous fishes

Synopsis We examined the feeding behaviors and selectives of two common planktivorous fishes, pumpkinseeds Lepomis gibbosa and fathead minnows Pimephales promelas in the laboratory. Ingestion rates for both pumpkinseeds and fathead minnows feeding on zooplankton increased as a function of fish length. Pumpkinseeds fed on zooplankton strictly as particulate feeders, with preferences increasing as a function of zooplankton body size regardless of taxonomic identity. Preferences were highest for large Daphnia, intermediate for intermediate-sized copepods, and lowest for small Ceriodaphnia. Fathead minnows displayed the ability to use both particulate-feeding and filter-feeding behaviors. Differential preferences tended to reflect both zooplankton size and taxon, being highest for large, slow-swimming Daphnia, intermediate for small Ceriodaphnia, and lowest for faster-swimming copepods. These differences in prey capture behaviors and preferences of the two fishes are reflected in the zooplankton taxonomic composition of small ponds containing each fish type. The crustacean zooplankton assemblages in ponds containing both pumpkinseeds and fathead minnows were dominated by copepods. Cladocerans were rare. In ponds containing pumpkinseeds, but no fathead minnows, cladocerans were abundant, generally accounting for up to 80% of total crustacean zooplankton biomass. These results suggest that the type of planktivore, and not simply the presence or abundance of planktivores in a system, can determine zooplankton community structure.     

Effect of prey size on attack components of the functional response by Notonecta undulata

The number of encounters per prey, the proportion of encounters resulting in attacks, and the proportion of attacks that were successful were observed while fourth-instar Notonecta undulata nymphs preyed on smaller N. undulata nymphs. While encounters per prey and proportion of encounters resulting in attacks increased with prey size, the proportion of attacks that were successful decreased. The increase in encounter rate per prey was due in part to an increase in the predator's reactive distance to prey as prey size increased. While none of the attack parameters varied significantly with prey density, logarithmic regression of the number of encounters per unit search time on prey density suggested that prey density tends to have a positive effect on encounters per first-instar prey but a negative effect on encounters per second-instar prey. A functional response model is presented that incorporates components of the predator's attack rate as exponential functions of prey density and allows for effects of the time the predator may spend evaluating prey encountered but not attacked and time spent attacking prey not captured. Estimates of the attack parameters derived from the experimental data are used in the model to generate functional response curves for fourth-instar N. undulata preying on first- or second-instar conspecifics. The predicted curve for second-instar prey is typical type II but the curve for firstinstar prey is slightly positively density dependent at low prey densities, i.e., type III.     

Resource utilization of sympatric and experimentally allopatric cutthroat trout and Dolly Varden charr

Summary Resource utilization by cutthroat trout (CT) and Dolly Varden charr (DV) was studied 8 years after experimental transfers from sympatry had established reproducing allopatric populations in two nearby fishless lakes. Allopatric DV significantly increased their utilization of shallow-dwelling zoobenthos, and increased their vertical distribution in comparison to that in sympatry. In contrast, allopatric CT showed little change in the proportions of major prey types utilized, and, if anything, restricted their vertical distribution in comparison to that in sympatry. The results can be explained by the hypothesis that the resource use of DV is strongly influenced by interspecific competition from CT, whereas CT largely remains unaffected by this interaction. An alternative hypothesis, that lake differences can explain the differences in resource use between sympatry and allopatry, was evaluated by comparing food resource availability and other biotic and abiotic characteristics of the three study lakes. None of these could account for the shift in resource use by DV between sympatry and allopatry, but lake differences may explain why allopatric CT showed a restricted habitat use in comparison with their sympatric donor stock. The results of this whole-lake transfer experiment are consistent with earlier reported field and laboratory studies, and suggest that the aggressive dominance of CT is the most important mechanism by which DV are displaced from littoral and near-surface habitats in sympathy with CT.     

Habitat requirements of weasels <Emphasis Type="Italic">Mustela nivalis</Emphasis> constrain their impact on prey populations in complex ecosystems of the temperate zone

Differences in habitat use by prey and predator may lead to a shift of occupied niches and affect dynamics of their populations. The weasel Mustela nivalis specializes in hunting rodents, therefore habitat preferences of this predator may have important consequences for the population dynamics of its prey. We investigated habitat selection by weasels in the Bia?owie?a Forest in different seasons at the landscape and local scales, and evaluated possible consequences for the population dynamics of their prey. At the landscape scale, weasels preferred open habitats (both dry and wet) and avoided forest. In open areas they selected habitats with higher prey abundance, except during the low-density phase of the vole cycle, when the distribution of these predators was more uniform. Also in winter, the distribution of weasels at the landscape scale was proportional to available resources. In summer, within open dry and wet habitats, weasels preferred areas characterised by dense vegetation, but avoided poor plant cover. In winter, weasels used wet open areas proportionally to availability of habitats when hunting, but in contrast to summer, they rested only in habitats characterized by a lower water level, which offered better thermal conditions. At the local scale, the abundance of voles was a less important factor affecting the distribution of these predators. Although we were not able to provide direct evidence for the existence of refuges for voles, our results show that they may be located within habitat patches, where availability of dense plant cover and physiological constraints limit the activity of weasels. Our results indicate that in complex ecosystems of the temperate zone, characterized by a mosaic pattern of vegetation types and habitat specific dynamics of rodents, impact of weasels on prey populations might be limited.     

Foraging behaviour in fishes: perspectives on variance

Synopsis The positive relationship between size of prey and frequency of ingestion by predators has been a focal point of investigations in foraging ecology. Field studies compare the frequency distribution of prey sizes in the predator's gut with that in the environment. Laboratory and field (enclosure) studies are based upon comparison of the frequency distributions of prey sizes in controlled environments, before and after the introduction of a predator. Optimal caloric return for foraging effort (i.e. the theory of optimal foraging) has been widely used as a guiding principle in attempts to explain what a fish consumes. There is a body of information, however, which seems to indicate that the perceptual potentialities and cognitive abilities of a predator can account for both the direction of the prey size versus ingestion frequency relationship and the variance surrounding it. Part of this variance may be evidence of systematic ambiguity, a property of cognitive skills causing predators to respond to the same stimulus in different ways and to different stimuli in the same way. More extensive examination of cognitive skills (minimally defined as learning, remembering and forgetting) in fish may permit causal interpretations (immediate and ultimate) of variance in predatory skills. In such a paradigm of foraging behaviour, environmental stimulus is not taken as the predator's object of response (percept); a cognitive representation connects mind to stimulus and this is the criterion for the act of perception. Cognition, here considered as a formal system which acts upon representations, connects mind to response and thus to adaptation. Studies of the relationships among rates of learning, long and short-term memory, rates of forgetting, prey behavior, size and population turnover rates, lateralization of brain functions, diel fluctuations in predator activity levels and sleep, experience, and critical periods in the development of the predator's nervous system should be examined in relation to foraging behaviour.     

Ecological and morphological shifts in Lake Michigan fishes: glimpses of the ghost of competition past

Synopsis Both historical patterns and recent evidence of resource partitioning and complementarity within the Lake Michigan fish community provide circumstantial evidence for interspecific competition. But competition is difficult to document in the field without controlled experimentation. In Lake Michigan, controlled experiments on competition within the fish community are nearly impossible, but we still need to understand the interactions among the dominant fishes. For this purpose, I have relied upon hypothesis-based field observation, natural experiments in the field and designed laboratory experiments to evaluate competitive interactions. Resource use patterns and trophic morphology of the bloater, Coregonus hoyi, a native cisco, from samples taken before alewife, Alosa pseudoharengus, became abundant (1960) were compared to more recent data (1979–80). After the alewife density increase, bloaters had significantly fewer and shorter gill rakers. This suggests a morphological shift toward greater benthic foraging efficiency in response to high abundances of an efficient pelagic planktivore, alewife. Resource use comparisons suggested that bloaters now shift from pelagic zooplanktivory to benthic habitats and diets at least two years earlier in their life history than they did before alewife became abundant. This evidence, albeit not experimental, provides strong support for the importance of competition in the structure of the current Lake Michigan fish community.In Lake Michigan, seasonal thermal habitat compression can pack fish into a narrow thermal zone across the lake bottom, leading to increased habitat overlap, reduced prey availability and fish diets containing fewer and smaller prey. Thermal habitat compression, which can occur intermittently through the season, may create competitive bottlenecks which help maintain the observed resource partitioning among these fishes.     

Trophic ecology of the Southern Grey Shrike (Lanius meridionalis) in insular environments: the influence of altitude and seasonality

The seasonal diet and prey selection of the Southern Grey Shrike (Lanius meridionalis) was studied in two different insular habitats: shrub environments of the Canary Islands in coastal and high mountain zones. We measured, in each season, food availability and prey size in order to determine prey size selection of shrikes along an altitudinal gradient. Moreover, we compared the diet patterns observed with those documented on the continent, to determine if Southern Grey Shrikes in the islands’ high mountain zone (which has a continental climate) showed seasonal diet variation similar to those in northern continental areas. We analysed a total of 1,139 shrike pellets collected in 1 year and identified 10,179 prey items. Numerically arthropods (91%), and in terms of biomass lizards (70%) were the main prey consumed by the shrikes. The proportions of the main prey items differed significantly between seasons and habitats. Diet in the coastal areas was less variable than in the high mountain zone. The greater seasonal climatic variation in the high mountain zone was associated with diet patterns similar to those found in some northern continental areas, such as the Iberian Peninsula and southern France. Finally, shrikes selected the largest prey in the high mountain habitat. This suggests that foraging behaviour in this species is related to climatic conditions, as the biggest and most profitable prey were consumed in the most harsh habitats.     

Optimal foraging and feeding mode shifts in fishes

Synopsis Most optimal foraging models for fishes are based on particulate feeding behavior. But many obligate planktivores also filter zooplankton. I suggest that feeding mode shifts (e.g. from particulate feeding to filtering) may be predictable from the costs and benefits of foraging in various modes. Quantitative examples of feeding mode shifts in three species of fishes (northern anchovy, pacific mackerel and alewife) from 3 different families support this hypothesis. Feeding mode shifts seem to depend on relative profitability of each mode, but improvements in model predictions will need to include the effects of spatial and temporal patchiness on encounter rates of prey of various sizes.     

Ichthyofaunal utilization of newly-created versus natural salt marsh creeks in Mission Bay,CA

Loss of wetland habitat has proceeded at an alarmingrate in southern California, and increasingly marshrestoration and creation are being used to mitigatethese losses. As part of an effort to evaluatefunctional equivalence of created systems, theichthyofaunal assemblages in a created and adjacentnatural marsh in Mission Bay, San Diego, Californiawere compared. Fishes trapped in both marshes includedFundulus parvipinnis, Gillichthysmirabilis, Acanthogobius flavimanus, Ctenogobius sagittula, Atherinops affinis, andMugil cephalus. Fundulus parvipinniswasnumerically dominant in both systems, representing onaverage 69% of all fishes trapped in the createdmarsh and 65% of all fishes trapped in the naturalmarsh. Gillichthys mirabiliswas the second-mostabundant species, representing on average 31% of allfishes trapped in the created marsh and 28% of allfishes trapped in the natural marsh. Species richnessand dominance measures were similar between the twosystems, while abundances were higher in the naturalrelative to the created marsh. The size-structure ofF. parvipinnisand G. mirabilisdifferedbetween the created and natural marsh creeks, with thecreated marsh populations being skewed towards largersize classes. These size differences are believed toarise from differences in creek morphology between thecreated and natural systems, and potentially affectboth predators and prey of these species in the marsh.Mark-release-recapture revealed considerable marshfidelity, with as many as 35% of the F.parvipinnistagged in a marsh being recovered one daylater in the same marsh. Stable isotope analyses ofF. parvipinnisrevealed similar ¹⁵Nand ³⁴S values between marshes; howeverthere was a consistent enrichment in ¹³C (>3per mil) in tissues of F. parvipinnisfrom thecreated marsh, supporting the high marsh fidelitysuggested by tagging results. This first publisheddocumentation of the Mission Bay marsh resident fishessuggests that the created marsh ichthyofaunalassemblage was distinct in density and size structurefrom the adjacent natural marsh, and provides lessonsfor future restoration efforts.     

Life-history styles of fishes in South African estuaries

Synopsis Fish utilizing South African estuaries may be divided into two major groups according to the location of their spawning sites. The marine group comprises large species which spawn at sea, enter estuaries mainly as juveniles, and return to the sea prior to attaining sexual maturity. The estuarine group is dominated by small species which have the ability to complete their life cycle within the estuarine environment. They tend to produce relatively few, demersal eggs, or exhibit parental care, which facilitates the retention of eggs and young within the estuary, whereas the marine group release large numbers of small pelagic eggs during spawning and exhibit no parental care. This is contrary to the theory that estuaries (unpredictable environments) should favour altricial life-history styles and the marine inshore zone (a more predictable environment) should favour precocial styles. However, if the total ichthyofauna of South African estuaries is considered, then altricial species predominate. The fact that both altricial and precocial traits are well represented within the overall estuarine fish community suggests that the various taxa have adapted their life-history styles, in different ways, to ensure the utilization of abundant food resources available within these fluctuating systems. A detailed comparison of the life-history styles of the estuarine teleostGilchristella aestuaria and the marine fishMugil cephalus is used to illustrate the contrasting manner in which these two species have succeeded in exploiting South African estuaries.     

Differential host utilization by two parasites in a population of ponderosa pine

We compared phloem characteristics of individual Pinus ponderosa attacked by the dwarf-mistletoe Arceuthobium vaginatum (Viscaceae) or by the beetle Dendroctonus ponderosae (Scolytidae) or by neither species. We quantified total nonstructural carbohydrates and a broad range of chemical elements for these three categories of trees. There were significant differences between trees parasitized by Arceuthobium, trees parasitized by Dendroctonus, and non-infected trees. Discriminant function analysis of trees attacked by either Arceuthobium or Dendroctonus correctly predicted group membership for 59 of 60 trees tested. Some of the differences detected may be induced, but many probably are not. Given that the accumulation of certain elements and compounds is under genetic control, and that both parasites often cause severe reductions in fitness, including death of their host, our results suggest that the two parasites may generate diversifying selection in ponderosa pine populations.     

Foraging behaviors and the interaction of alewife,Alosa pseudoharengus,and bloater,Coregonus hoyi

Synopsis Alewife, Alosa pseudoharengus, and bloater, Coregonus hoyi, are common planktivores in Lake Michigan. Both alewife and bloater use a variety of feeding modes. Alewives can filter, gulp and particulate feed; bloaters can only gulp and particulate feed. We examined handling time per prey and probability of capture for alewife and bloater particulate feeding on Mysis relicta. Using these estimates and available data for filtering alewives, cost curves were derived for alewife and bloater particulate feeding and for alewife using all three modes of feeding. Alewives filter small prey relative to their own body size and particulate feed on larger prey. Feeding mode appears to be dependent on prey size and density and shifts in feeding mode are apparently based on maximizing biomass eaten per time. The ability to filter confers a competitive advantage on alewife when small prey are abundant as they were in the mid 1960s in Lake Michigan. If the zooplankton are large, bloater young-of-year do not suffer this relative disadvantage. In fact, large bloaters can consume prey on the bottom not available to alewife. This shifting competitive balance may explain, in part, the observed dynamics of alewife and bloater.     

Trophic relationships between primary producers and fish yields in Lake Balaton

Relationships between chlorophyll a content of the water, the shoreline-length: water area ratio and the annual total fish yield as catch per unit effort (CUE: kg ha^–1 100 h^–1 as annual mean values) have been calculated by multivariable regression. The determination coefficient (r ² = 0.913) showed a significant dependence of fish yield on morphometry of different lake areas. Accordingly, fish carrying capacity of the open water areas, calculated from chlorophyll a content and S/A, ranged from 12 to 34%, but that of the littoral zone between 66 and 88%. These findings have also been supported by echo-sounding records of the horizontal distribution of fish.Bream (Abramis brama L.) contributes the majority (70–80%) of fish stock and yield. Its food mainly consists of zooplankton and benthic invertebrates in ratios that are widely variable with season and depend on the age of fish. Average daily food consumption of individuals (age group 3 + and over) varies between 2 and 5 g. Bream consumes two- to three-times more food in the SW basin than in the NE one. This means that the present stocks inhabiting areas from NE to SW consume annually 13249–20085 t yr^–1 of food. According to estimated calorific values, the annual energy consumption of local populations along the longitudinal axis of the lake varies between 93 and 141 kJ m^–2 yr^–1. The efficiency of energy transfer from primary producers to fish is low and varies from 0.04 to 0.1%.