The food and feeding relationships of larval and 0+ year juvenile fishes in lowland rivers and connected waterbodies. I. Ontogenetic shifts and interspecific diet similarity

Many fish species are zooplanktivorous at the onset of exogenous feeding, but distinct shifts in their foraging biology may occur with development. This study investigated the food and feeding relationships of the larvae and 0+ year juveniles of 13 fish species in 'main river', 'marina' and 'pond' macrohabitats in the lower River Trent, England. Young larvae had narrow diet spectra, with feeding diversity and the importance of species-specific traits increasing with development. Two main ontogenetic shifts in diet were differentiated, corresponding with the transition from finfold to finformed larvae, and from the larval to the juvenile period. Dietary overlap was generally greatest among ecologically similar species, with competition and resource partitioning most likely where prey availability was lowest. The greater availability of zooplankton in connected waterbodies compared with main river channels provides suitable prey for a wide range of fish species, and may enhance recruitment success.     

Feeding ecology of the Iberian cyprinid Barbus bocagei Steindachner, 1865 in a lowland river

Patterns of food resource utilization by Barbus bocagei in a lowland river in the central part of the Iberian Peninsula are presented and discussed. Focal points include feeding activity, diet composition and seasonal and ontogenetic diet shifts. Striking features of overall feeding pattern of barbel are its continuous feeding activity, bottom habits, high consumption of detritus and dipteran larvae and narrow diet breadth. Seasonal changes in food resource availability elicited changes in food utilization patterns, revealing a trend to more generalist feeding during spring. Barbel increase consumption of plant material and large prey during ontogeny, but overlap in prey resource utilization between size classes is high. The versatility of barbel foraging behaviour may be an important feature in its exploitation of Iberian lotic systems.     

Diet changes in prey size selectivity in larvae of perch <Emphasis Type="Italic">Perca fluviatilis</Emphasis>

A hypothesis on change in prey size selectivity in relation to illumination level was tested on the basis of data on weight and size composition of the content of the digestive tract of larvae of perch Perca fluviatilis and on zooplankton in the layer 0–6 m (Wallersee Lake, Austria). Larvae foraging in the twilight-night period had almost two times more food in the intestine than those foraging in the daytime. The size composition of perch larvae and concentration and size composition of zooplankton hardly differed in the daytime and twilight-night samples. For the first time, it is shown on field material that more intensive feeding of larvae at twilight is related to selection of significantly larger prey than in daytime feeding. In the day-time, the larvae consumed more prey but their maximum size did not exceed 0.6 of the diameter of mouth opening of the fish; at twilight it was over 0.8. In case of feeding on so large prey, not only the weight of the consumed feed increases but the time used for capture and swallowing also considerably increases. The larvae which in the period of investigation did not yet form schools, which perform the principal defensive function, were especially vulnerable for predators feeding on relatively large prey. The decrease of the part of small-sized prey at twilight is not related to their lesser availability due to a low illumination. It is assumed that feeding on energetically more valuable but less available prey is shifted to the period of low illumination when the larvae are less exposed to predation risk. The obtained results are discussed from positions of the triotroph concept (Manteifel’, 1961).     

Diet and feeding habits of monkey goby (Neogobius fluviatilis) in a newly invaded area

The monkey goby (Neogobius fluviatilis) is one of the Ponto-Caspian species that in recent decades made substantial East-to-West invasions in European inland waters. Now the species is present in the Danube as far as its Slovakian section. In the last decade the species also invaded the Western Bug and Vistula rivers, in the latter it is numerous nearly as far as its delta. The purpose of our study was to define the diet spectrum, feeding preferences, spatial and size related changes in diet and diurnal feeding activity of this goby. Another goal was to predict which groups of native prey would be most affected by the presence of this exotic predator in newly invaded areas. The study was carried out at three sampling locations in the Vistula River system. The food spectrum of the monkey goby was broad. The fish consumed insect larvae and pupae, crustaceans, annelids, gastropods and fish. Chironomid larvae were a prevalent food category in all sampling sites, followed by amphipod crustaceans at one site and by trichopteran larvae and chironomid pupae at another. At the third site, there was no distinct subdominant food category. According to the values of the Ivlev’s selectivity index, the preferred food category were chironomid larvae. No significant differences in diet were found over the 24-h cycle. There was no variation among different fish size groups. From our studies and from the available literature data it can be concluded that the species, due to its ability to use locally available food resources, displays a generalistic and highly flexible feeding strategy.     

Prey selection by young trout fry (Salmo trutta)

Factors influencing the choice of invertebrate food by early feeding trout are sought, special attention being given to the dimensions of the prey. The stomach contents of fish of similar age feeding on lake plankton and drifting stream fauna are compared. Little consistent preference was displayed for the smaller planktonic organisms but prey size was of importance in the distribution of stream benthos among predators of different lengths. The width of the prey was of greater significance to selection than the mean volume or the area of the largest surface of the food items. When the fish were in obvious competition for food the correlation between predator length and all prey dimensions improved.     

Size-biased predation by the gila topminnow poeciliopsis occidentalis (Baird and Girard)

The objective of this study is to document under both field and laboratory conditions the phenomenon of size-biased feeding by the Gila topminnow, Poeciliopsis occidentalis, and to shed light on the mechanisms responsible for this process. Adult female Gila topminnows feed in Monkey Spring, Santa Cruz County, Arizona, on a variety of food items, but Hyalella azteca is an important element of its invertebrate diet. The size of the amphipods ingested by female topminnows is a function of the size of the fish, with larger fish taking larger prey. There is also a strong preference for topminnows to select large amphipods from within the size limits imposed by the width of the mouth.The correlation between fish size and prey size was also seen in the laboratory, utilizing Daphnia similis and food pellets as prey. The tendency to select large prey from a range of available sizes was prominent with both Daphnia and pellets. The feeding period in the laboratory is characterized by two distinct phases. During the first phase, feeding is rapid and nonselective. As the feeding progresses, the rate of feeding drops dramatically and selectivity increases.More complex laboratory feeding experiments gave equivocal results. When fish were presented pellets of two sizes at different densities and at different ratios of small to large, size biased feeding was not clearly evident. Both predator and prey densities, agonistic behavior, and reaction distance should be explored more thoroughly as next steps in attempts to mimic feeding behavior in nature.Extracted from a Ph. D. thesis written by the junior author     

The autecology of the chub, Squalius cephalus (L.), of the River Lugg and the Afon Llynfi

(1) Seasonal variation in the intensity of feeding activity was demonstrated in two chub populations. Feeding activity was higher in summer and more pronounced in the young age-classes. The intensity of feeding activity was significantly correlated with temperature. (2) The diet of the total chub populations embraced a wide range of food items but vegetable material predominated. Substrate material was frequently encountered which suggested considerable feeding on the river bed. Fish were the most important animal food, by volume, in the River Lugg but Trichoptera larvae were most important in the Afon Llynfi. No remains of salmon parr or trout were found in chub stomachs but grayling were eaten. The most important prey fish, by volume, was the eel but cyprinids, especially minnows, were most important by frequency of occurrence. (3) Seasonal variation in the composition of the diet could be related to availability of food organisms but some selection was shown. Vegetable foods were most important in summer; animal foods, Trichoptera larvae in particular, were important in winter. (4) Analysis of the variation in the composition of the diet with age revealed several trends, notably a decrease in the importance of plant material, aquatic and aerial insects with age and a corresponding increase in the importance of larger prey foods such as fish, frogs and crayfish. (5) The commonly held view that chub are deleterious to salmonid stocks was examined. The absence of direct predation and the small likelihood of direct competition suggests that this view is incorrect.     

Behavioural and life history effects of predator diet cues during ontogeny in damselfly larvae

A central issue in predator–prey interactions is how predator associated chemical cues affect the behaviour and life history of prey. In this study, we investigated how growth and behaviour during ontogeny of a damselfly larva (Coenagrion hastulatum) in high and low food environments was affected by the diet of a predator (Aeshna juncea). We reared larvae in three different predator treatments; no predator, predator feeding on conspecifics and predator feeding on heterospecifics. We found that, independent of food availability, larvae displayed the strongest anti-predator behaviours where predators consumed prey conspecifics. Interestingly, the effect of predator diet on prey activity was only present early in ontogeny, whereas late in ontogeny no difference in prey activity between treatments could be found. In contrast, the significant effect of predator diet on prey spatial distribution was unaffected by time. Larval size was affected by both food availability and predator diet. Larvae reared in the high food treatment grew larger than larvae in the low food treatment. Mean larval size was smallest in the treatment where predators consumed prey conspecifics, intermediate where predators consumed heterospecifics and largest in the treatment without predators. The difference in mean larval size between treatments is probably an effect of reduced larval feeding, due to behavioural responses to chemical cues associated with predator diet. Our study suggests that anti-predator responses can be specific for certain stages in ontogeny. This finding shows the importance of considering where in its ontogeny a study organism is before results are interpreted and generalisations are made. Furthermore, this finding accentuates the importance of long-term studies and may have implications for how results generated by short-term studies can be used.     

Trypsin activity and physiological aspects in larval rearing of European sea bass (Dicentrarchus labrax) using live prey and compound diets

The aim of this study was to evaluate the potential of a compound diet as a live prey substitute for feeding European sea bass larvae ( Dicentrarchus labrax L.). The effect of a commercial diet (Nippai ML feed) and live prey ( Artemia nauplii) on tryptic enzyme activity, protein content, growth (standard length) and survival rates of sea bass larvae were tested during a 27-day rearing experiment. Sea bass larvae were divided into two groups. The live food group (control group) was fed exclusively on newly hatched Artemia nauplii (Inve AF grade), the test group was fed exclusively with the compound diet from day 15 onwards. As trypsin has been demonstrated to be a useful indicator for evaluating digestibility of food and the nutritional condition of fish larvae, individual tryptic enzyme activity was determined in both feeding groups. Larvae older than 14 days after hatching and fed on live food showed a significantly higher tryptic enzyme activity than larvae fed the compound diet. A similar relationship between tryptic activity and standard length in both test groups was detected only in small larvae (standard length < 7 mm). The usefulness of proteolytic enzyme activity measurements in larval fish research, as well as its use in aquaculture nutrition research, was confirmed. Protein content, increase in length and survival rates of the sea bass larvae were additionally determined in order to evaluate an influence on the diet. The protein content of larvae fed the Artemia nauplii was higher and the growth of larvae fed the compound diet was reduced. Larval mortality was not affected by the diet given.     

Effects of season and body-size on the distribution and diet of the Iberian chub Leuciscus pyrenaicus in a lowland catchment

Seasonal and ontogenetic variations in space and food use by Leuciscus pyrenaicus were analysed in a Portuguese lowland catchment. Large fish occurred mostly in deep permanent-flowing sites whereas small fish occurred mostly in the shallowest intermittent-flowing site. No seasonal or size-related changes in feeding intensity were found, but the diet changed both across seasons and throughout ontogeny. The diet was dominated by aquatic prey over all seasons, but during winter and summer more plant material and terrestrial prey, respectively, were eaten. Throughout ontogeny fish shifted from soft-bodied to hard-shelled prey and decreased animal prey breadth. Mean prey size increased with fish size but the prey size spectrum was more variable for medium-sized fish than for either small or large fish. It is suggested that: (i) large fish avoid shallow drying areas owing to the risk of mortality, either by thermal and respiratory stresses or increased predation by mammals and birds; (ii) seasonal changes in diet are a response to differences in prey availability; and (iii) morphological constraints, prey handling costs and habitat partitioning are responsible for size-related changes in diet.     

Feeding habits and diet overlap of marine fish larvae from the peri-Antarctic Magellan region

The Magellan region is a unique peri-Antarctic ecosystem due to its geographical position. However, the knowledge about the distribution and feeding ecology of fish larvae is scarce. Since this area is characterized by low phytoplankton biomass, we hypothesize that marine fish larvae display different foraging tactics in order to reduce diet overlap. During austral spring 2009–2010, two oceanographic cruises were carried out along southern Patagonia (50–56°S). Larval fish distribution and feeding of the two most widely distributed species were studied, the smelt Bathylagichthys parini (Bathylagidae) and black southern cod Patagonotothen tessellata (Nototheniidae). Larvae of B. parini showed a lower increase in the mouth gape at size, primarily feeding during daytime (higher feeding incidence during the day) mostly on nonmotile prey (invertebrate and copepod eggs, appendicularian fecal pellets, diatoms). They showed no increase in feeding success (number, total volume of prey per gut and prey width) with increasing larval size, and the niche breadth was independent of larval size. Larvae of P. tessellata showed a large mouth gape at size, which may partially explain the predation on motile prey like large calanoid copepods (C. simillimus) and copepodites. They are nocturnal feeders (higher feeding incidence during night) and are exclusively carnivorous, feeding on larger prey as the larvae grow. Nonetheless, niche breadth was independent of larval size. Diet overlap was important only in individuals with smaller mouth gape (<890 μm) and diminished as larvae (and correspondingly their jaw) grow. In conclusion, in the peri-Antarctic Magellan region, fish larvae of two species display different foraging tactics, reducing their trophic overlap throughout their development.     

Modulation in Feeding Kinematics and Motor Pattern of the Nurse Shark <Emphasis Type="Italic">Ginglymostoma cirratum</Emphasis>

Synopsis Studies of feeding in bony fishes have almost universally demonstrated the ability of individuals to modulate their method of capture in response to differing stimuli. Preliminary evidence indicates that morphologically specialized inertial suction feeding sharks are the most likely fishes to lack inherent modulatory ability. We examined the ability of the nurse shark, Ginglymostoma cirratum, to modulate its feeding behavior based on different food types and sizes. G. cirratum is an inertial suction feeding fish that is apparently stereotyped in its food capture behavior. Electromyography showed no statistical difference between feeding motor patterns based on food type (squid or fish) or size (gape width or twice gape width), although there were slight inter-individual differences in the onset of muscle firing for some muscles. Kinematic analysis showed a statistical difference in variables associated with durations for different food types, with the durations for all variables being faster for squid bites than fish bites, but no difference based on the size of the food item. This apparent lack of modulation may be associated with specialization of the morphology and behavior of G. cirratum for obligate suction prey capture. This functional specialization constrains the method in which G. cirratum captures prey but does not appear to result in dietary specialization. An unusual post capture spit-suck manipulation allows this shark to handle and ingest large prey.     

Effect of Diet History on Prey and Pollen Food Choice by Two Lady Beetle Species

Mixed diets of prey and plant-provided foods, such as pollen, have been shown to benefit a wide range of arthropods. However, diet shifting between these two very different food sources remains poorly understood. We hypothesized that previous diet should influence subsequent time allocation between prey and plant food types; to reach a balanced diet, consumers are expected to allocate more time to resources previously lacking in their diet. We tested this hypothesis by observing the foraging choices of larvae of two omnivorous coccinellid species: Coccinella septempunctata L. and Hippodamia variegata (Goeze) (Coleoptera: Coccinellidae), following a diet history of prey, pollen or a mixed diet of both food types. Results showed an asymmetrical tendency of C. septempunctata larvae to complement their previous diet with unfamiliar food: larvae allocated more time to pollen feeding, but not to prey, when each of the foods was previously absent from their diet. Study results have important implications for the use of plant-provided food supplements to enhance biological control by these omnivorous consumers.     

Food selectivity and diel vertical distribution of Chaoborus edulis (Diptera, Chaoboridae) in Lake Malawi

1. Diel diet and vertical distribution patterns of the larval instars of Chaoborus edulis were studied in deep water near the central part of Lake Malawi, Africa.
2. First instar larvae contained very little food in their crops and probably depended on reserves from the egg. Second, third and fourth instars fed on zooplankton and were size-selective in their feeding. The mean size of prey eaten by the three instars was significantly different from each other, with larger instars feeding on larger prey. Smallest available prey was selected against and the upper size of prey was probably constrained by larval gape. Nauplii were not found in any of several thousand larvae examined. Phytoplankton did not form a significant part of the diet.
3. There was a progressive and related increase in diel periodicity in feeding and vertical migrations of successive instar stages. Fourth instars migrated particularly large distances. Such migrations removed them from their zooplankton food supply but avoided predators. A refuge from predators is probably found in or near the permanent zero oxygen boundary, at depths greater than 200 m.     

Diet overlaps and feeding relationships between the year classes in the yellow perch (Perca flavescens)

Synopsis A comparative study of the diets of five year classes of perch in Lake Opinicon showed Year class 0 to be mainly Cladocera feeders, the Year class 11 to take a diversified range of insect larvae, whereas from Year class V onwards Anisoptera nymphs, decapods, and fish dominated the diet. Sizes of the dominant food items changed strikingly with growth, prey weights of 0.1, 0.5–1.0, and 100–250 mg predominating in these three age groupings. There were marked month to month changes in types and proportions of different prey consumed.Overall and month to month feeding overlaps between consecutive year classes were considerable, most values ranging from 0.68–1.0 by the Levins formula. However, diet overlap dropped sharply with increasing age gap.Month to month dietary changes and year class differences were interpreted relative to benthic invertebrate abundance cycles. Months of increased diet overlap were characteristically those when favored prey types peaked, thereby simultaneously attracting several year classes to them. Hence increased diet. overlap is not necessarily an indicator of increased intraspecific competition.Habitat differences between the year classes of perch in Lake Opinicon were partial and concerned mainly the smaller, and larger, fish relative to those in the middle size range.     

Diel-feeding activity in early summer of racer goby Neogobius gymnotrachelus (Gobiidae): a new invader in the Baltic basin

The first recording of the Ponto‐Caspian racer goby in Poland was during 1995 in the River Bug (River Vistula system). Within 5 years, the species had spread to the downstream section of the Vistula. One of the potential impacts of invasive species on native fauna is competition for food. Therefore, the diel patterns in diet composition and gut fullness coefficient (FC) of racer goby were examined at one study site in the W?oc?awski Reservoir (lower River Vistula), during May 2003. An average of 20 individuals were examined each 4 h over one 24‐h period (125 fish in total). The proportion of main food items and diet width did not differ among three size‐groups (43–59, 60–79 and 80–97 mm total length), and the relative biomass ratio of main food categories did not differ over the diel cycle. Amphipods constituted 11–70% of total gut content biomass and were found on average in 84% of analysed alimentary tracts. The second prey types were chironomid larvae (16–63% of total food biomass; frequency occurrence: 61–91%), and to a lesser extent chironomid pupa, ceratopogonid larvae, oligochaets, dipteran imagines and copepods, with fish larvae found in the gut of eight gobies. Gut fullness coefficient (FC) differed significantly over the 24‐h period, with the highest value at night and in early morning. In conclusion, racer goby forages mainly on benthos and has a nocturnal‐feeding activity. No significant diet overlap was found between racer goby and native percids, i.e. Eurasian perch Perca fluviatilis and ruffe Gymnocephalus cernuus.     

Relationship between gape size and feeding selectivity of fish larvae from a Neotropical reservoir

Larvae feeding selectivity of Iheringichthys labrosus , Hypophthalmus edentatus and Plagioscion squamosissimus was assessed, examining the role of mouth gape in prey selection. Fish larvae were sampled in the Itaipu Reservoir (Brazil–Paraguay). Iheringichthys labrosus and H. edentatus larvae, with small and similar gape sizes, exhibited slightly different diets; I. labrosus preferred cladocerans ( Bosmina hagmanni , Bosmina huauriensis and Bosminopsis deitersi ) and the rotifer Brachionus calyciflorus . Hypophthalmus edentatus , however, primarily ingested the cladocerans B. hagmanni , Ceriodaphnia cornuta , Daphnia gessneri and Diaphanosoma spinulosum . Plagioscion squamosissimus , with a greater gape size, preferred Calanoida. The mechanistic processes that determine food selectivity of fish larvae in temperate aquatic systems were similar in the Neotropical system. The trophic spectrum of these species is characterized by small- to intermediate-sized prey. Plagioscion squamosissimus larvae, which have larger mouths, exploit primarily larger prey differing from the most abundant species or size classes; consequently, their diet is quite different from I. labrosus larvae and modestly similar to H. edentatus larvae, opportunistic feeders that they eat more abundant prey.     

The influence of prey size and abundance, and individual phenotype on prey choice by the three-spined stickleback, Gasterosteus aculeatm L.

Prey selection behaviour of three-spined sticklebacks, Gasterosteus aculeatus L., was studied in two experiments. Where possible, the experimental apparatus satisfied the assumptions of the simplest optimal diet model (the basic prey model); prey were presented sequentially, the fish could not search for and handle prey at the same time, and net energy gain, handling time and encounter rate were fixed. Experiment 1 presented fish with a range of Asellus sizes so that pursuit ( p ) and handling ( h ) time could be related to prey size. Published energy values of Asellus together with pursuit and handling times were used to calculate E /( p+h ) for Asellus measuring 3,4,5,6,7 and 9 mm. Pursuit times did not differ with prey size but handling times did. E /( p+h ) was very variable particularly at the larger prey sizes. Experiment 2 presented fish with two sequences of prey differing in the encounter rate with the most profitable prey sizes. Fish did not select the diet predicted by the basic prey model tending to always ignore the largest prey even when net energy gain would have been maximized by including them in the diet. Further analysis showed that the probability of a prey size being taken was a function of prey size, fish stomach fullness and encounter rate. It is concluded that the basic prey model is too simple to capture the behaviour of the fish. One of its main faults is that the changing state of the fish through the feeding bout is ignored.     

Non-selective predator – the versatile diet of Amur sleeper (Perccottus glenii Dybowski, 1877) in the Vistula River (Poland), a newly invaded ecosystem

The Amur sleeper diet spectrum was investigated in addition to its spatial, seasonal and size-related changes, in order to predict which groups of native prey would be most affected by this exotic predator that has rapidly invaded many Eastern and Central European inland waters in recent decades. In total, 527 Perccottus glenii individuals were collected in four sites in the Włocławski Reservoir (the Vistula River, Baltic basin, Poland). Altogether 50 food categories were identified, including crustaceans, insects, molluscs, annelids, araneids, fishes and amphibians. Most of them were associated with aquatic vegetation, indicating that the Amur sleeper collects its prey from the submerged plants rather than directly from the bottom sediments. Amphipods and chironomid larvae, supplemented by zygopteran larvae and molluscs, composed the main forage base in all studied sites, although their importance was varied. The diet composition differed among P. glenii size-groups. Significant variation was found in numeric abundance of amphipods, chironomid larvae, fish and zygopteran larvae (Kruskal–Wallis P < 0.01). Seasonal changes in the diet were also observed whereby in spring and summer, dipteran larvae and amphipods were prevalent food items; the importance of molluscs and zygopteran larvae increased considerably in September; and fish showed the highest occurrence in June, July and August. The broad diet spectrum of the Amur sleeper indicates that it is a non-selective, opportunistic predator and that several taxonomic groups of native hydrofauna as well as macroinvetebrates and fish may be affected by its presence. The highly flexible feeding strategy undoubtedly favours Amur sleeper expansion in invaded watersheds.