Scanning behavior by larvae of the predacious diving beetle, <Emphasis Type="Italic">Thermonectus marmoratus</Emphasis> (Coleoptera: Dytiscidae) enlarges visual field prior to prey capture

Larvae of the predaceous diving beetle Thermonectus marmoratus bear six stemmata on each side of their head, two of which form relatively long tubes with linear retinas at their proximal ends. The physical organization of these eyes results in extremely narrow visual fields that extend only laterally in the horizontal body plane. There are other examples of animals possessing eyes with predominantly linear retinas, or with linear arrangements of specific receptor types. In these animals, the eyes, or parts of the eyes, are movable and perform scanning movements to increase the visual field. Based on anatomical data and observations of relatively transparent, immobilized young larvae, we report here that T. marmoratus larvae are incapable of moving their eyes or any part of their eyes within the head capsule. However, they do perform a series of bodily dorso-ventral pivots prior to prey capture, behaviorally extending the vertical visual field from 2° to up to 50°. Frame-by-frame analysis shows that such behavior is performed within a characteristic distance to the prey. These data provide first insights into the function of the very peculiar anatomical eye organization of T. marmoratus larvae.     

Feeding by larvae of Hypoatherina tropicalis (Pisces: Atherinidae) and its relation to prey availability in One Tree Lagoon,Great Barrier Reef,Australia

Synopsis Feeding of and food availability for larvae of Hypoatherina tropicalis were investigated in One Tree Lagoon, Great Barrier Reef, Australia, during November 1981 and January 1982. These surface-dwelling larvae and their microzooplankton prey were sampled as near to simultaneously as possible on 12 occasions during the daytime. Larvae of all sizes (5–17 mm SL) fed successfully over the observed range of mean prey densities (12–235 per liter), and the overall feeding incidence was 98.9%. Larger larvae consumed greater numbers and more categories of prey than did smaller larvae. Larvae selected copepods of all sizes, and nauplii, gastropods, bivalves, and foraminiferans that were greater than 75 ¢s mm in width. Tintinnids (mostly 37–74 µm in width) were generally avoided by larvae, but were occasionally important in the diets when they constituted more than 60% of the total available prey, regardless of the density of the selected prey categories. Larvae less than 14 mm SL ingested meroplankton (gastropods, bivalves, foraminiferans, and polychaetes) in direct relation to the densities available, and without regard to the densities of copepods available. However, the largest larvae (14–17 mm SL) ingested meroplankton in inverse relation to the density of copepods available, indicating that larvae consumed more meroplankton when the concentration of copepods was low. Such flexibility and opportunism in feeding behavior may increase the larvae's chances of obtaining adequate nutrition during periods of suboptimal feeding conditions.     

Effects of varying irradiance on feeding in larval weakfish (Cynoscion regalis)

Weakfish larvae, Cynoscion regalis (Bloch and Schneider), were used in laboratory experiments, during May and June 1991–1993, to examine the effects of varying irradiance levels on capture and ingestion of Zooplankton prey (rotifers). Treatments consisted of six different irradiance levels: no light, 5, 11, 15, 20, and 500 × 10¹² quanta·cm⁻²·s⁻¹. These levels are typical of the irradiance range found in a 10-m water column during the late-spring, weakfish spawning season in Delaware Bay. Early-stage larvae (8 days post-hatching) did not feed in total darkness, and there was no difference in the incidence of feeding among the other treatment groups. Similarly, late-stage larvae (13 days post-hatching) showed no significant difference between the incidence of feeding in darkness and at 5 × 10¹² quanta·cm⁻² s⁻¹, though feeding within these two intensities was significantly lower than feeding in the other light levels. Results of a subsequent experiment indicated that the ability to feed in total darkness may depend on the abundance of available prey. Scanning electron microscope analysis of preserved weakfish larvae showed that neuromasts were not fully developed until larvae had reached at least 12 days post-hatching, and that younger larvae had only lateral line pores along the body trunk. There were no neuromasts evident on the head region, regardless of age. Thus, neuromasts may be involved in the capture of prey in darkness.     

Formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma,larvae under laboratory conditions: role of visual and chemical stimuli

Synopsis Although planktonic marine fish larvae are often distributed in aggregations, the role of behavioral responses to environmental factors in these aggregations is not well understood. This work examines, under laboratory conditions, the influence of visual and chemical stimuli in the formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma, larvae. Larvae were exposed to a horizontal gradient of light (visual stimulus), prey scent (chemical stimuli: squid/copepod and rotifer) or prey density (visual & chemical stimuli: rotifers). While larvae did not respond to prey scent, they did respond to a gradient of light or prey, which resulted in the formation and maintenance of aggregations. Larvae moved into and remained in a zone of higher light intensity (0.56 versus 0.01 mol photons m^-2 s^-1). Once encountering a patch of prey, larvae remained aggregated within the patch to feed. In nature, movement of walleye pollock larvae in response to selected environmental factors (e.g., gravity, light, temperature, turbulence) may serendipitously bring them into contact with prey patches, where they then could remain to feed as long as light intensity remained at or above levels necessary for feeding.     

Prey selection and functional morphology through ontogeny of Amphiprion clarkii with a congeneric comparison

The relationship between the complexity of the feeding apparatus and prey selection through ontogeny was examined in Amphiprion clarkii larvae. Larvae were reared from 1 to 10 days post-hatch (dph) on a diet of rotifers, wild-caught plankton and newly hatched Artemia sp. nauplii. Results were compared with available data on the relationship between functional morphology and prey selection of Amphiprion frenatus to establish patterns of functional morphology and prey selection between the larvae of two species of coral-reef fishes. Larvae of both species exhibited an increase in selection of larger prey through ontogeny coincident with an increase in the complexity of the feeding apparatus. The first elements to ossify in larvae of both species were the pharyngeal teeth ( A. clarkii : 5 dph, near ± s.d. Standard length, L _S,4· 3 ± 0· 2 mm; A. frenatus : 5 dph, L _S5· 0 ± 0· 4 mm) which, in combination with the development of a more functional feeding apparatus, may have permitted larvae to better process new types and sizes of prey. Prey items, however, were selected differentially between the two fish species, which could not be fully explained by the functional state of the feeding apparatus. While prey selection is influenced by the functional state of the feeding apparatus, all aspects of larval fish biology (morphology, behaviour and physiology) should be considered.     

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.     

Visual and olfactory location of biotopes, prey patches, and individual prey by the ladybeetleChilocorus nigritus

Foraging behaviour of the predatorChilocorus nigritus (Fabricius) (Coleoptera: Coccinellidae) at the three spatial levels of biotope, prey patch and individual prey, was studied in the laboratory, and related to behaviour in the field. Vertically oriented parallel lines were more attractive than the same shapes in a horizontal position. A simulated horizon with a tree line was preferred to a simulated flat horizon. They were attracted to a tree image for the first 2 h of exposure, but were less attracted after longer exposure, possibly due to habituation. Leaf shape was recognised, and simple ovate leaves were preferred to compound bipinnate leaves and to squares. These responses were associated with biotope selection for feeding and aggregation at aestivation sites. The location of prey patches by adults involved prey odour but the location of such sites by larvae did not. Adults detected individual prey visually and olfactorily over short distances but physical contact with prey was required for detection by larvae. Location of individual prey and prey patches by adults and larvae was facilitated by alternation between intensive and extensive search. The differences in the ability of larvae and adults to locate prey, stem from the adults being the active locators of biotope and patch, whereas the comparatively immobile larvae depend on their parents’ ability for long-range location of prey. Two hypotheses concerning coccinellid foraging behaviour are proposed. Firstly, the duration of response to a visual cue is related to the distance over which such a cue may be perceived. It follows that habituation to closer range cues occurs more rapidly than to longer range cues. Secondly, visual cues used by adults at the different spatial levels of prey location, and the location of mates and aggregation sites, have the same or similar shape. These results also provide guidelines for orchard management to maximise the biocontrol value of this species.     

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).     

Visual acuity and feeding in larval Premnas biaculeatus

Ontogenetic change in the visual acuity of Premnas biaculeatus larvae was determined both behaviourally and anatomically. Visual acuity improved substantially between early feeding (day 3 post-hatch) and the pre-settlement (day 10 post-hatch) larvae but, at both ages, the anatomically-measured visual acuity was greater than that determined behaviourally. It appears that estimated anatomical visual acuity values substantially over-estimate the functional visual acuity realized under normal conditions. The distribution of the reactive angles indicated that most frequently prey within 0 to 9° of the longitudinal larval axis elicited a feeding response at both larval ages. This suggests that stereoscopic vision is used extensively during feeding in this species. The prey capture success with rotifers ranged from 96% at 3 days post-hatch to 100% at 10 days post-hatch. These values differ markedly from previous studies on temperate species and highlight the well developed abilities of larval P. biaculeatus at a given     

Vulnerability of larval white perch,Morone americana,to fish predation

Synopsis The vulnerability of white perch, Morone americana, larvae to yearling bluegill, Lepomis macrochira, predators was examined in relation to larval size, nutritional condition and relative abundance of alternative prey. Short-term (15 min) predation trials were conducted in 381 tanks in the laboratory. Larval vulnerability was measured as the proportion of larvae killed and the proportion of successful attacks per predator in each 15 min trial. No significant differences in vulnerability were apparent among larvae regardless of feeding history at sizes < 6 mm SL. At larval lengths > 6 mm SL, size of larvae was the crucial determinant of their vulnerability. Percentage of larvae killed in 15 min was nearly 100% at sizes < 6 mm SL, decreased to 30% at a length of 12.0 mm SL and dropped to 18% at 14.0 mm SL. Larvae initially feeding at low food levels for 2–4 d exhibited decreased growth of 13–25% over the first 3 wks of life, and simulations based on laboratory results indicated that these growth deficits could result in 5- to 68-fold decreases in survivorship at 38 days after hatching (DAH). The relative abundance of alternative prey also had a pronounced effect on mortality of larvae. A 10-fold increase in alternative prey (Daphnia magna) abundance decreased bluegill predation rates on white perch larvae by 10–20%, while a 100-fold increase in Daphnia density decreased larval mortality by 75–90%.     

Observations on the larval development of Mystus macvopterus (Bleeker): Bagridae

From June to July 1988, larvae of Mystus macropterus (Bleeker) were obtained by artificial propagation of spawners collected from the Jialing River, China. Larvae grown in water temperatures ranging from 26 to 29°C were fully developed at approximately 20 days. The newly-hatched larvae measured 6.0–7.5 mm t.l. , exceeding the dimensions of any known newly-hatched larvae of freshwater catfishes indigenous to China. Five days after hatching, when the larvae were 11.4 mm t.l. and the prolarval stage was complete, exogenous feeding commenced. Twelve days after hatching, at 18.4 mm t.l. , the yolk sac disappeared, organogenesis was almost complete and the juvenile period began. At 20 days post-hatching, at 23.0 mm t.l. , the lateral line system had formed and the juveniles resembled the adults with respect to all external features, which signalled the end of thejuvenile period. Comparison of the larval development in eight species ofeconomic freshwater catfishes shows that the most significant aspect of the larval development of M. macropterus is the sustained (6–7 days) mixed nourishment period and the early differentiation of organs, which enhances the survival rate of the larvae.     

Laboratory culture of the Nudibranch Tritonia diomedea bergh (Tritoniidae: Opisthobranchia) and some aspects of its behavioral development

Tritonia diomedea Bergh was reared from oviposition, through metamorphosis to reproductive maturity in the laboratory. The larvae of T. diomedea are planktotrophic and undergo considerable shell growth (from 144.6–329 μ average maximum shell length). Metamorphosis does not require induction, but there may be a preference to metamorphose in the presence of the probable adult prey, a small Virgularia sp. Larvae in cultures fed no food, Dunaliella tertiolecta Butcher Isochrysis galbana Parke, or Monochrysis lutheri Droop did not achieve metamorphic competence at near ambient sea-water temperatures (11.9±1.3 and 13.0±0.8°C). Larvae from cultures fed Monochrysis at room temperature (20.8±1.5°C) or fed a 1 : 1 mixture of Isochrysis and Monochrysis at near ambient sea-water temperatures did metamorphose. Even so, only those larvae fed the 1 : 1 mixture survived more than a few days following metamorphosis. Adult behavioral patterns developed gradually, feeding being first observed at 5 days, swimming in response to NaCl crystals at about 60 days, copulation at about 272 days, and oviposition at about 277 days after metamorphosis. Growth rates were determined for field collected Tritonia diomedea; smaller animals gained and lost weight relatively faster than larger animals.     

Trophic ecology of Atlantic bluefin tuna Thunnus thynnus larvae

Feeding intensity, diet composition, selectivity, energy ingestion and dietary niche breadth of larval Atlantic bluefin tuna Thunnus thynnus were studied on the eastern (Mediterranean) spawning grounds of the species. Larval T. thynnus were collected in the Balearic Archipelago (north-west Mediterranean Sea) during 2004 and 2005 using surveys specific for larval scombrids. Larvae between 2·6 and 8·7 mm standard length (L(S) ) are diurnal feeders, and 94% of the guts collected during daylight hours were full. The mean ±s.d. number of prey per gut was 7·1 ± 5·7, with mean ±s.d. ranging from 3·0 ± 1·6 in the smallest T. thynnus larvae to 11·1 ± 5·8 in 5·0-6·0 mm L(S) larvae. Up to 21 prey were found in a single larval gut (5·0-6·0 mm L(S) ) at the end of the day. Larvae progressively selected larger prey and exhibited increased carbon content concurrent with preflexion development of feeding and locomotory structures. Larvae of 5·0-6·0 mm L(S) exhibited positive selection of cladocerans over other prey (Chesson's index), whereas copepod nauplii dominated the diets of earlier stages. The dietary niche breadth measured increased initially but decreased at c. 5·5 mm L(S) . Appendicularians were found in the diet of larger larval sizes, but no piscivory was observed. Results are discussed in light of the sparse existing data for larval T. thynnus and other larval tuna species.     

Vertical migrations of herring,Clupea harengus,larvae in relation to light and prey distribution

Synopsis The influence of light and prey abundance on the vertical distribution of herring larvae was evaluated by three investigations made under calm weather conditions in the North Sea off the Scottish coast. The investigations took place at different time after hatching and the vertical distributions of three size groups of larvae (mean sizes 8,15 and 19 mm) were related to time of day and the vertical distribution of copepods. No migratory behaviour of copepods was observed but their vertical distribution differed between investigations. In the investigation on intermediate sized larvae, copepod density peaked at the pycnocline (40 m). Larvae concentrated at this depth at noon. At dawn and dusk larvae migrated towards the surface and the vertical distributions fluctuated semidielly. In the two other investigations, copepods were homogeneously distributed in the water column and after migration towards the surface at dawn larvae stayed in the upper water column during the day. The observations suggest that the daytime vertical distribution of larvae in calm weather is mainly determined by feeding conditions: the larvae move to depths were light is sufficient for feeding, and refinement within that zone is made according to a compromise between optimal light conditions for feeding and optimal prey densities.     

Feeding of larval blue whiting and Atlantic mackerel: a comparison of foraging strategies

Fish larvae employ different feeding strategies depending on area and season of spawning and hatching of larvae. Feeding and growth of larvae of blue whiting Micromesistius poutassou and mackerel Scomber scombrus from Porcupine Bank and the Celtic Shelf Break, west of Ireland, were compared based on prey concentrations in the environment and larval feeding behaviour. Both species were adapted to different environmental conditions. The mesopelagic blue whiting spawned in oceanic water that was well mixed. It was characterized by low production and low prey densities with minimum prey densities <1.0 organism 1⁻¹. Larvae of the Atlantic mackerel hatched later in the season in more productive water that was well stratified. Prey densities in the mackerel environment reached up to 1001⁻¹. Blue whiting larvae displayed a rather random distribution in the water column. Mackerel larvae <7 mm standard length ( L _s) were concentrated above the thermocline, while larvae >7 mm traversed the thermocline into deeper layers. Mackerel larvae >5mm L _s displayed marked cannibalism, exceeding 70%. Daily ration calculated on the basis of gut contents was rather low in both species: between 2.6 and 5.0% in blue whiting, but only 0.6 to 5.4% in mackerel. The results are discussed in relation to the respective environment both species encounter during their early larval life.     

Feeding habits of larval Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret (Israel) II. Experimental study

Interactions between the larvae of Mirogrex terraesanctae (Steinitz, 1952) in Lake Kinneret, Israel, and their zooplankton prey were studied experimentally. Prey species preference and size selectivities were measured. Larvae were hatched in the lab from eggs collected in the field, and fed different food items in various concentrations. The food items included lake zooplankton, algae, and commercial pellets. It was shown that small, first feeding larvae (7–8.5 mm SL) prefer small bodied zooplankters (< 180 µ). The effect of these food sources on larval growth was measured. It was found that larval Mirogrex grew at a higher rate when fed zooplankton prey sized from 63 µ–250 µ. Food items smaller than 63 µ, larger than 250 µ and Scenedesmus sp., produced less than optimal growth rates. The importance of Mirogrex feeding habits and their potential influence on the Kinneret ecosystem is considered.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Turbidity enhances feeding abilities of larval Pacific herring,Clupea harengus pallasi

Fishes inhabiting estuaries, rivers, and embayments are subject to turbid conditions. Larvae of many fishes utilize estuaries as nursery areas. For visual plankton feeders such as larval fishes, turbidity may reduce search and reaction distances, resulting in lowered feeding abilities. In this study feeding Pacific herring larvae, Clupea harengus pallasi, were exposed to suspensions of estuarine sediment and Mount Saint Helens volcanic ash at concentrations ranging from 0 mg · l^–1 to 8 000 mg · l^–1. In all experiments, maximum feeding incidence and intensity occurred at levels of suspension of either 500 mg · l^–1 or 1000 mg · l^–1 with values significantly greater than controls (0 mg · l^–1). Feeding decreased at greater concentrations. The suspensions may enhance feeding by providing visual contrast of prey items on the small perceptive scale used by the larvae. Larval residence in turbid environments such as estuaries may serve to reduce predation from larger, visual planktivores, while searching ability in the small larval perceptive field is not decreased.     

Gut evacuation of walleye pollock larvae in response to feeding conditions

Gut residence times of first-feeding walleye pollock larvae were measured at 6°C in continuous and discontinuous feeding regimes. Larvae fed tagged copepod prey evacuated their guts more quickly in continuous feeding as compared to the discontinuous feeding treatment. The mean gut residence time was estimated to be 5.0 h for larvae feeding continuously. Gut evacuation by larvae fed tagged prey and then isolated without food (discontinuous treatment) was slower and more variable, with an estimated gut residence time exceeding 8.0 h. Field and laboratory observations suggest that the larval fish gut may be modeled as an intermittent plug-flow reactor (PFR) in response to diel feeding patterns. The cyclical nature of gut dynamics has implications for gut content analyses and the estimation of daily food rations.     

Avoidance of unpalatable prey by predaceous diving beetle larvae: The role of hunger level and experience (Coleoptera: Dytiscidae)

The effects of hunger and experience on the avoidance of unpalatable prey were examined. Larvae of the predaceous diving beetle,Dytiscus verticalis, after feeding on one of two feeding regimes, were offered palatable and unpalatable prey during a series of trials. Consumption of palatable prey (calf heart) was not affected by hunger or experience. Avoidance of unpalatable prey (tails of the red-spotted newtNotophthalmus viridescens) decreased with increased hunger, with hungrier predators sampling unpalatable prey more often. Although the overall level of avoidance varied across trials, the degree of difference in avoidance between predators at different hunger levels remained relatively constant across trials. Beetle larvae responded to an increase in their feeding regime within 36 h (after two trials), by becoming more selective. In a separate experiment, hunger level affected the number of beetle larvae that seized unpalatable newt tails, but experience did not.