Onshore–offshore variations of copepod community in northern South China Sea

Previous studies which have tested the feeding preferences of shredders for fungal species and the food quality of fungi used detritus uniformly colonized by a fungus, which is not the case for decaying leaves in streams. It is not known whether shredders in different development stages exhibit variations in feeding preference and larval performance. This study examined the feeding preferences and the growth of the third and the fifth instars of Pycnopsyche gentilis larvae using fungal-colonized patches and whole leaves, respectively, having different fungal species compositions (Alatospora acuminata, Anguillospora filiformis, Articulospora tetracladia, Tetrachaetum elegans, and all species combined). The aquatic hyphomycetes used were co-dominant on leaves in the stream inhabited by the caddisfly. During 14 d of feeding, the larvae of both instars did not show significant differences in feeding preferences for the patches growing on oak leaves, although the third instar larvae were slightly more selective than the fifth instar larvae. When fed with maple leaves for 18 d, larval growth rates, gross growth efficiencies, and survivorship were not significantly different among the fungal treatments. However, the larval growth of both instars fed with fungal-colonized leaves was always significantly greater than the growth of larvae fed with diets of uncolonized leaves. The third instar larvae grew faster than the fifth instar larvae, but the growth efficiencies of the two instars were similar. These results suggest that P. gentilis larvae exhibit less selectivity in their feeding than other caddisfly shredders that have been examined and that the dominant fungi colonizing leaves in their habitat are similar in palatability and food quality for this shredder. Handling editor: B. Oertli     

Condition and size of damselflies: a field study of food limitation

Summary Based on evidence from field manipulations, several authors have recently suggested that interference competition among larval odonates reduces individual growth rates and biomass by reducing foraging rates. This study was designed to test the effects of food shortage on condition (relative mass per unit head width) of larval Ischnura verticalis (Odonata: Coenagrionidae) under laboratory conditions and to use these results to estimate the degree of food shortage of larvae under naturally occurring field conditions. In the laboratory, there were marked differences in condition of larvae fed diets ranging from ad libitum feeding with worms to ad libitum feeding with Daphnia 1 day out of every 8. Condition of larvae collected from May through October from 17 different sites in southern Ontario indicated that, for most of the year, larvae had conditions similar to those fed ad libitum with Daphnia in the laboratory. There was no evidence that larval condition was related to population density. Condition of larvae in most sites during July was similar to that of larvae fed poor diets in the laboratory. It is unlikely that the low conditions were due to competition as there were no correlations with density across sites and population densities during July were at their lowest. Adult head widths showed a seasonal decline from mid June to the end of the flight season. There was no evidence that head widths were related to population density although there was some evidence that head widths of males were positively related to larval condition. My results do not support the hypothesis that competition is important in affecting foraging rates and subsequent development of larvae. Contrasts between my results and other studies may stem from difficulties with the interpretation of field experiments, that densities in my study may have been low due to fish predation, and/or that I. verticalis larvae are slow moving relative to other larvae and thus less likely to interact.     

EVOLUTIONARY LOSS OF LARVAL FEEDING: DEVELOPMENT,FORM AND FUNCTION IN A FACULTATIVELY FEEDING LARVA,BRISASTER LATIFRONS

Species with large eggs and nonfeeding larvae have evolved many times from ancestors with smaller eggs and feeding larvae in numerous groups of aquatic invertebrates and amphibians. This change in reproductive allocation and larval form is often accompanied by dramatic changes in development. Little is known of this transformation because the intermediate form (a facultatively feeding larva) is rare. Knowledge of facultatively feeding larvae may help explain the conditions under which nonfeeding larvae evolve. Two hypotheses concerning the evolutionary loss of larval feeding are as follows: (1) large eggs evolve before modifications in larval development, and (2) the intermediate form (facultatively feeding larva) is evolutionarily short-lived. I show that larvae of a heart urchin, Brisaster latifrons, are capable of feeding but do not require food to complete larval development. Food for larvae appears to have little effect on larval growth and development. The development, form, and suspension feeding mechanism of these larvae are similar to those of obligate-feeding larvae of other echinoids. Feeding rates of Brisaster larvae are similar to cooccurring, obligate-feeding echinoid larvae but are low relative to the large size of Brisaster larvae. The comparison shows that in Brisaster large egg size, independence from larval food, and relatively low feeding rate have evolved before the heterochronies and modified developmental mechanisms common in nonfeeding echinoid larvae. If it is general, the result suggests that hypotheses concerning the origin of nonfeeding larval development should be based on ecological factors that affect natural selection for large eggs, rather than on the evolution of heterochronies and developmental novelties in particular clades. I also discuss alternative hypotheses concerning the evolutionary persistence of facultative larval feeding as a reproductive strategy. These hypotheses could be tested against a phylogenetic hypothesis.     

Adult beetles compensate for poor larval food conditions

Life history traits of herbivores are highly influenced by the quality of their hosts, i.e., the composition of primary and secondary plant metabolites. In holometabolous insects, larvae and adults may face different host plants, which differ in quality. It has been hypothesised that adult fitness is either highest when larval and adult environmental conditions match (environmental matching) or it may be mainly determined by optimal larval conditions (silver spoon effect). Alternatively, the adult stage may be most decisive for the actual fitness, independent of larval food exposure, due to adult compensation ability. To determine the influence of constant versus changing larval and adult host plant experiences on growth performance, fitness and feeding preferences, we carried out a match–mismatch experiment using the mustard leaf beetle, Phaedon cochleariae. Larvae and adults were either constantly reared on watercress (natural host) or cabbage (crop plant) or were switched after metamorphosis to the other host. Growth, reproductive traits and feeding preferences were determined repeatedly over lifetime and host plant quality traits analysed. Differences in the host quality led to differences in the development time and female reproduction. Egg numbers were significantly influenced by the host plant species experienced by the adults. Thus, adults were able to compensate for poor larval conditions. Likewise, the current host experience was most decisive for feeding preferences; in adult beetles a feeding preference was shaped regardless of the larval host plant. Larvae or adults reared on the more nutritious host, cabbage, showed a higher preference for this host. Hence, beetles most likely develop a preference when gaining a direct positive feedback in terms of an improved performance, whereby the current experience matters the most. Highly nutritious crop plants may be, in consequence, all the more exploited by potential pests that may show a high plasticity in reproduction and feeding preferences.     

The relationship between egg size and the duration of the facultative feeding period in marine invertebrate larvae

Feeding larvae of marine invertebrates fuel development from both endogenous egg energy and exogenous energy obtained from the planktonic environment. Although both sources of energy likely influence certain larval stages, only the effects of exogenous food have been well studied. Despite the lack of research on the effects of egg size on larval stages, investigators have hypothesized that egg size influences the duration of the facultative feeding stage—the stage in which larvae can feed but do not have to because development is still being fueled by egg energy. To test this hypothesis, we investigated six species of sand dollars with different sized eggs and quantified the duration of the larval facultative feeding period of each species by comparing when fed and starved larvae diverged in size. Regardless of whether phylogeny was taken into account, the duration of the facultative feeding period was positively correlated with egg size. We further determined that our conclusions were not sensitive to either our estimation of the duration of the facultative feeding period, or the branch lengths of the phylogeny we used. This relationship is likely a result of larger eggs being provisioned with more energy, and may affect how well larvae can cope with natural variability in food concentrations. Furthermore, our results support an assumption of a theoretical model developed to understand the evolution of different life-history strategies in marine invertebrate larvae, which suggests that this relationship has important evolutionary consequences.     

Optimal feeding strategy of the temperate herbivorous fish Aplodactylus punctatus: the effects of food availability on digestive and reproductive patterns

In this study we examined differences in feeding behevior of populations of the marine temperate herbivorous fish Aplodactylus punctatus, in three different localities off the Chilean coast, which differ qualitatively and quantitatively in food availability. We test whether food selection follows optimal foraging strategics, whether there is any modification of the fishes' digestive tracts in relation to their diets, and whether differences in diet quality affect the allocation of energy into reproduction and maintenance in these populations. Samples of this fish and of the understore algal assemblages were taken seasonally from May 1989 to February 1990. For each population we analyzed dietary composition, weight of the digestive tract and of the food content, the condition factor (K), and the gonadosomatic index (GSI). Our results showed that the diet observed in the three populations closely resembled the differences in macro-algal abundance and composition among the three localities studied. Local differences in diet quality were inversely related to the amount of food consumption and size of the digestive tract, suggesting that under differential conditions of food availability A. punctatus is able to compensate for variations in food quality through a flexibility in its digestive strategies. The nutritional status (K) of individuals, and their reproductive pattern (GSI) were directly related to diet quality. These results indicate that although A. punctatus is able to adjust its digestive processes to different algal food regimes, the digestive modifications observed in food-poor environments are not sufficient to compensate for the lack of food and allow fish to reach the nutritional status and reproductive output reached in a food-rich environment. This study represents the first natural experiment demonstrating a direct relationship among food availability, feeding patterns, digestive processes, and reproductive effort.     

Density‐dependent natural selection in Drosophila: correlations between feeding rate,development time and viability

We have previously hypothesized that density‐dependent natural selection is responsible for a genetic polymorphism in crowded cultures of Drosophila. This genetic polymorphism entails two alternative phenotypes for dealing with crowded Drosophila larval cultures. The first phenotype is associated with rapid development, fast larval feeding rates but reduced absolute viability, especially in the presence of nitrogenous wastes like ammonia. The second phenotype has associated with it the opposite set of traits, slow development, slow feeding rates and higher viability. We suggested that these traits are associated due to genetic correlations and that an important selective agent in crowded larval cultures was high levels of ammonia. To test this hypothesis we have examined viability and larval feeding rates in populations kept at low larval densities but selected directly for (i) rapid egg‐to‐adult development, (ii) tolerance of ammonia in the larval environment and (iii) tolerance of urea in the larval environment. Consistent with our hypothesis we found that (i) larvae selected for rapid development exhibited increased feeding rates, and decreased viability in food laced with ammonia or urea relative to controls, and (ii) larvae selected to tolerate either ammonia or urea in their larval environment show reduced feeding rates but elevated survival in toxin‐laced food relative to controls. It would appear that development time and larval feeding rate are important characters for larvae adapting to crowded cultures. The correlated fitness effects of these characters provide important insights into the nature of density‐dependent natural selection.     

Inspection and feeding of larvae by worker honey bees (Hymenoptera: Apidae): Effect of starvation and food quantity

Honey bee larvae are frequently inspected and, sometimes, provided with food by adult workers, but the stimuli that elicit the important task of food provisioning have never been investigated. Larvae with their food experimentally deprived received more frequent inspection and feeding visits from nurse bees than normally fed larvae, suggesting that there could be a hunger signal. Food-deprived larvae with artificially supplied larval food received the same rate of feeding visits from nurse bees as did normally fed larvae but still received more inspection visits. These results suggest that stimuli eliciting feeding are different from those for inspection. They also support the hypothesis that worker bees deposit food in a larval cell only when the quantity of food is below a certain minimum threshold that is perceived during larval inspections. A model is presented regarding the stimuli from larvae that result in worker feeding behavior.     

Effects of food,water depth,and temperature on diving activity of larval Anopheles gambiae sensu stricto: evidence for diving to forage

Anopheles gambiae larvae have frequently been observed to dive, but the ecology of this behavior has not been extensively examined. We manipulated food level, water depth, and temperature for individually‐reared larvae and observed diving activity. Larvae dived more often under low food, which suggests that they dive to forage. There was only weak evidence for effects of water depth or temperature on diving. Experimental results are discussed in the context of energy budgets. Understanding larval ecology of this species is important for predicting how it will respond to environmental change. Further study is needed to assess the role that larval diving plays in both feeding ecology and thermal regulation of this and other medically important species.     

Alpine newts (Triturus alpestris) as top predators in a high-altitude karst lake: daily food consumption and impact on the copepod Arctodiaptomus alpinus

1. Population dynamics and feeding ecology of adult and larval alpine newts (Triturus alpestris, Laurenti) were investigated in a high-altitude karts lake to estimate their feeding pressure on the copepod Arctodiaptomus alpinuf (Imhof). Estimates of population size for reproducing adults ranged from 666 to 864 individuals in the lake during July and August. Total abundance of larvae before the onset of ice cover varied considerably between 4400 and 25400 individuals in different years. 2. Arctodiaptomus alpinus was an important prey item for adult and larval alpine newts. During the second half of their aquatic period, adult newts moved to deeper water where the copepod reached its highest densities near the sediment. Adults and larvae exhibited no periodic feeding pattern. The feeding rhythm was more synchronized among the larvae than among the adults. 3. Daily food consumption, estimated using the Elliott & Persson (1978) model, reached 4–21 mg dry biomass in adults. The daily ration of larvae was about 7% of body dry weight in the temperature range 6-11°C. Compared to published estimates of daily food consumption in salmonid fishes, the feeding pressure of newts appears low.     

Effects of food availability on larval development in the slipper limpet Crepidula onyx (Sowerby)

Effects of food availability on the larval survival and development of Crepidula onyx were studied in four experiments by feeding the larvae with different concentrations of the chrysophyte Isochrysis galbana and by starving the larvae for different periods of time. Food concentration had a clear impact on the survival, growth and development time of C. onyx veligers. Larval development occurred only at 10⁴ cells ml⁻¹ and higher algal concentrations. No shell increment was detected in the veligers cultured for 12 days at 10² cells ml⁻¹I. galbana or the blank control. At 10³ cells ml⁻¹, there was only a slight increase in shell length over 12 days. At 10⁴ cells ml⁻¹, about 40% of the larvae became competent in 18 days. At 10⁵ and 10⁶ cells ml⁻¹, more than 90% of the larvae reached competence in 7 days. Initial starvation negatively affected the larval development, but the sensitivity differed among parameters measured on day 5: lower survivorship was detected only for larvae that had suffered 3 days or longer initial starvation, whereas one-day initial starvation caused shorter shells and lower percentage of competent larvae. Three days of continuous feeding was required for 50% of the larvae to reach competence. After feeding for 3 days, most larvae could become competent to metamorphose even under starvation. The time of starvation was also critical: larvae that suffered 1-day food deprivation in the first 2 days of larval release had shorter shells and lowered percent competent larvae than those that suffered the same length of food deprivation in later stages of development. Our study thus indicates that both food concentration and short-term starvation have detrimental effects on the larval development of this species, and that once the larva has consumed certain amount of food, starvation may induce metamorphosis.     

Changes in spatial distribution pattern during the larval stage of the fall webworm,Hyphantria cunea drury (Lepidoptera: Arctiidae)

Summary The changes in spatial distribution pattern during larval stage of the fall webworm,Hyphantria cunea were quantitatively investigated in the field experimental populations. The female adult deposits eggs as a cluster and the hatchlings make a compact colonial-web. In this period, the all-or-none type mortality which is characteristic in gregarious insect species was occasionary recognized before spinning a compact colonial-web. Once making a compact colonial-web, the larvae feed the leaves in the colonial-web up to about 5th instar. In this period, the movement of larvae occurred due to the local food shortage in a colonial-web and the expansion of colonial-web. As the larvae developed, the colonial-web was separated into several small groups. These larvae began to disperse about 5th instar. In this period, the local food shortage seems to be an important trigger for the larval dispersal. The mean concentration of larvae on leaves abruptly decreased, and finally the larvae became solitary at the 6th or 7th instars. The dispersal process in later larval stage is not necessarily due to the complete food shortage. The dispersal prior to the occurrence of food shortage may be a safety mechanism to protect the larvae from the food shortage.     

Age differences in food intake and dietary selection of wild male Japanese macaques

Age differences in food intake and dietary selection were studied for 8 months among wild male Japanese macaques (Macaca fuscata) under non-predatory situations. Juveniles' feeding time was longer than adult males' in some months, in particular in mating seasons, but did not differ in the other months. Juveniles' feeding speed was slower than that of adult males. However, the age difference in average feeding speed was smaller (90%) than that in expected daily energy expenditure (62–58%). The extent of age difference in feeding speed varied with the food type: the difference was large for fibrous foods, but small for fruits or seeds. As a consequence of the age differences in time spent feeding and feeding speed, the age difference in daily food intake was smaller than expected from metabolic demands. Thus, the hypothesis that juveniles are more vulnerable to starvation than adults was not supported among male Japanese macaques in predator-free Yakushima. Juveniles ate more animal matter, while adult males ate more fibrous foods.     

Rotifers as food in aquaculture

The rotifer Brachionus plicatilis (O.F. Muller) can be mass cultivated in large quantities and is an important live feed in aquaculture. This rotifer is commonly offered to larvae during the first 7–30 days of exogenous feeding. Variation in prey density affects larval fish feeding rates, rations, activity, evacuation time, growth rates and growth efficiencies. B. plicatilis can be supplied at the food concentrations required for meeting larval metabolic demands and yielding high survival rates. Live food may enhance the digestive processes of larval predators. A large range of genetically distinct B. plicatilis strains with a wide range of body size permit larval rearing of many fish species. Larvae are first fed on a small strain of rotifers, and as larvae increase in size, a larger strain of rotifers is introduced. Rotifers are regarded as living food capsules for transferring nutrients to fish larvae. These nutrients include highly unsaturated fatty acids (mainly 20: 5 n–3 and 22: 6 n–3) essential for survival of marine fish larvae. In addition, rotifers treated with antibiotics may promote higher survival rates. The possibility of preserving live rotifers at low temperatures or through their resting eggs has been investigated.     

Diel food utilization by woundfin,Plagopterus argentissimus,in Virgin River,Arizona

Synopsis Woundfin, Plagopterus argentissimus, fed predominantly on simuliid larvae during the day, and shifted to the larger Hydropsyche spp. at night. Ephemeropteran larvae were eaten nearly uniformly throughout the 24 hour period. Mean weight of food consumed varied from 20–40 mg per individual (0.7–2.5% body weight) throughout the 24 h period. Our estimated daily ration of about 8% body weight, at temperatures that varied between 15–25°C, is based upon our measure of mean gut contents over a 24h period, combined with intestinal evacuation rates suggested for other cyprinids in the literature. The relatively continuous feeding pattern suggests a foraging strategy that would minimize competition with other species for food during June, a period of minimum annual food abundance in Virgin River.     

The feeding ecology of the pit-making ant lion larva,Myrmeleon bore: Feeding rate and species composition of prey in a habitat

The prey species composition and feeding rate of the pit-making ant lion larva,Myrmeleon bore Tjeder, which inhabits open sandy areas, were examined. Not less than 30 prey species, most of which were ants, were collected during a research period of 1.5 years. First instar larvae most often (81.1%) captured ants. Although 3rd instar larvae captured larger-sized prey than individuals of any other instar, they also captured small prey. The feeding rate of 3rd instar larvae was estimated by using the frequency of observed predation (FOP; (no. of ant lions handling a prey)/(total no. of pits observed)), the prey-handling time and the rhythm of daily foraging activity. FOP ofM. bore larvae was constant on the whole from spring to autumn. It was estimated that each captured 1.25 prey per day on average during this period. This estimate, however, was the feeding rate for days on which there was no rain. Assuming that the larvae cannot capture prey due to pit destruction when there is more than 10 mm of rainfall per day, the figure was reduced to 1.03 prey/day. The estimated feeding rate was evaluated with reference to larval foraging behavior.     

Influence of larval food consumption on the fecundity of the lacewingChrysoperla carnea

The effects of food consumption on larval growth and development and adult fecundity of the common green lacewing,Chrysoperla carnea (Stephens), were studied on two populations of larvae derived from either a laboratory colony or from field-collected adults. The number of eggs of the Mediterranean flour moth,Anagasta kuehniella (Zeller), provided to individual lacewing larvae was varied to produce three food-supply treatments: low, intermediate, and high. Food-supply was found to influence larval growth and development and adult fecundity. Lacewing larvae provided with an overabundance of moth eggs developed faster than larvae provided with fewer moth eggs than they could have consumed. Adult females that developed from the high feeding treatment had a substantially shorter preoviposition period, a later decline in egg deposition, and a significantly higher fecundity than adults arising from the other feeding classes. Unrestricted feeding by adult lacewings on an artificial diet did not compensate for prior low feeding regimes. The overall performance of the lacewings derived from the laboratory colony was substantially poorer than that of the lacewings derived from field-collected adults. This effect was accentuated when the larvae were given a low food-supply. This overall decrease in vigor is attributed to inbreeding of the laboratory culture over a one year period. These finds are relevant to mass rearing programs for this biological control agent.     

Extrinsic vs. intrinsic food shortage and the strength of feeding links: effects of density and food availability on feeding rate of Hyphydrus ovatus

Summary We use field and laboratory experiments to determine whether Hyphydrus ovatus, a predatory aquatic beetle, is food limited, and whether any food shortage results from depletion of prey by these predators (intrinsic food shortage) or is independent of predation by these beetles (extrinsic food shortage). In the laboratory, differences in feeding rate influence body fat content, thus making fat content a useful index of recent feeding history. H. ovatus collected during the breeding season have fat contents significantly greater than those of H. ovatus starved for 25 days, but not significantly different from those of H. ovatus fed ad libitum for 25 days, indicating that natural feeding rates are near the maximum possible. H. ovatus confined at a density 60 times greater than natural show reduced fat content and feeding rate relative to natural, indicating that at very high densities H. ovatus is capable of depleting its prey. Addition of supplemental natural prey (primarily Cladocera) to experimental enclosures resulted in an order of magnitude increase in prey availability, and a significant increase in fat content and feeding rate of confined H. ovatus. Adults of this species do not appear to be food limited during the breeding season, and extraordinarily high densities of adults seem to be necessary to produce intrinsic food shortage. These results suggest that feeding links between H. ovatus an its principal prey do not have major effects on population dynamics under typical field conditions, and call into question the assumption that closely coupled predator-prey interactions are the sole explanation for observed food-web patterns.     

Temporal and developmental aspects of diet-induced food preferences in larvae of the tobacco hornworm, Manduca sexta

Diet‐induced changes in food preferences in an oligophagous caterpillar were studied in order to characterize the conditions under which this induction occurs. The time course of acquisition and extinction of induced food preferences by larvae of the tobacco hornworm, Manduca sexta (Johan.) (Lepidoptera, Sphingidae), was examined by varying the food and duration of larval food experience. Larvae were given feeding experience with the host plant tomato, Lycopersicon esculentum (Mill.) (Solanaceae), or the acceptable non‐host plant cowpea, Vigna unguiculata (Walp.) (Fabaceae), or switched from one to the other during different instars. Food preferences for V. unguiculata by the fourth or fifth instar larvae were measured individually in two‐choice tests with discs of V. unguiculata and moist filter paper. The acquisition or extinction of an induced food preference for V. unguiculata was indicated by larvae preferring V. unguiculata to filter paper or the reverse, respectively. Results showed that: (1) the period required for the acquisition of induced preference for V. unguiculata can be short (36 h), (2) food experience in either the third or fourth instar period is sufficient, and (3) the most recent feeding experience appears to be important. In contrast, the period required for extinction of induced preference for V. unguiculata appeared to be longer (1–3 instars), and both the most recent feeding experience and total duration of larval experience with the inducing food seem to play a role. Experience is not restricted to a particular instar period to acquire or extinguish such an induced food preference. The induced food preference for V. unguiculata was not very rigid and could be reversed by having one instar of feeding experience on L. esculentum. The findings indicate that diet‐induced food preferences in M. sexta contains elements of habituation and associative learning, but do not support food imprinting and induction of oligophagy.     

Group feeding as a strategy for exploiting food resources in the burnet moth Pryeria sinica

Summary Pryeria sinica (Zygaenidae) larvae feed on young and growing leaves of Euonymus japonicus in groups. The larvae often defoliate their host plant. Hence, the larvae are occasionally subject to serious shortage in food resources. We hypothesize that larval aggregation is an adaptation for the economical utilization of limited food resources. To test this hypothesis, the patterns of resource utilization were studied on larvae settled on shoots of host plant in various group sizes. The amount of food resources in a shoot was affected by the following three factors; (1) shoot growth, (2) food consumption of larvae, and (3) inhibition of growth or degeneration of shoot by larval activity on it. These factors were measured and the efficiency in resource utilization was compared among groups of varied sizes. The loss in resources caused by the third factor was found to be decreased with group size, i.e., the larger the group size, the larvae utilized the food resources more economically. The advantages of group feeding in survival and reproduction of this species were discussed using a simple graphical model.