The influence of proline on diet selection: sex-specific feeding preferences by the grasshoppers Ageneotettix deorum and Phoetaliotes nebrascensis (Orthoptera: Acrididae)

Diet selection based on the level of proline in an insect's host plant has been observed for a number of phytophagous insects, but few studies have examined potential differences in feeding preferences between males and females. The level of proline among an insect's host plants, particularly in drought-stressed plants, can be highly variable and often is positively correlated with soluble nitrogen levels. Additionally, proline is known to participate in a number of physiological functions in insects. We tested the effect of proline as a feeding stimulant in reproductively active grasshoppers using the graminivorous Ageneotettix deorum and the generalist, but mostly graminivorous, Phoetaliotes nebrascensis. Feeding preference tests using diets with representative free amino acid and sucrose levels but varying proline levels (zero, normal and 3 x normal) were examined. The feeding preference exhibited by both species was sex-specific, although the sex-specific response was more pronounced in P. nebrascensis than in A. deorum. Females of both species displayed preferences for diets high in proline. Males of neither species exhibited a preference for proline when responses were averaged over all treatment levels. However, within specific treatment combinations, male A. deorum preferred diets with high proline over diets with zero proline. These results suggest that diet selection for specific nutrients may vary between males and females because of differences in their physiological status and, possibly, differences in the nutritional requirements associated with reproduction. These results also suggest that subtle shifts in the concentration of individual nutrients within an insect's host plant may greatly influence insect feeding patterns.     

Similarities,differences, and seasonal patterns in the diets ofEulemur rubriventer andEulemur fulvus rufus in the Ranomafana National Park,Madagascar

I studied the diets of two sympatric species ofEulemur (E. fulvus rufus andE. rubriventer) in the Ranomafana National Part in southeastern Madagascar from July 1988 through August 1989. Both species were highly frugivorous throughout the year and devoted similar amounts of time to feeding daily; the composition of their diets were similar. Three aspects of both lemur species' diets were correlated with seasonal food availability: changes in interspecific dietary overlap, the length of feeding bouts, and the amount of time devoted to feeding daily. Throughout the year neither species had a more diverse or more folivorous diet than its sympatric congener. However, there were several consistent year-round differences between their diets.E. f. rufus initiated more feeding bouts daily that were shorter in duration than those ofE. rubriventer. E. f. rufus also consistently exploited more unripe fruits and mature leaves thanE. rubriventer did throughout the study.E. f. rufus migrated from the study site during a period of fruit scarcity. These dietary patterns are discussed here in regard to feeding patterns observed in other sympatric congeneric primate species, whether these dietary differences can contribute to these two species' ability to coexist, and how differences in diet may be influenced by differences in social structure. I suggest that subtle, consistent differences in diet and seasonal dietary differences are sufficient to allow these two species to coexist.     

Temporal pattern of resource use and variation in diets of individual grasshoppers (Orthoptera: Acrididae)

Patterns of resource fidelity, switching, and variation in individual diet were examined in the polyphagous grasshopper Taeniopoda equesforaging in experimental cages over an 11-day period. Ten novel dicots were provided in the cages, but grasshoppers fed primarily on kale (Brassica oleraceavar. acephala),lobelia (Lobelia erinus),pansy (Viola x wittrockiana),and dry bermuda grass (Cynodon dactylon).Grasshoppers showed significant fidelity to a single plant within a meal but tended to eat less of the same plant in the next meal and, in the case of pansy, on the next day. Grasshoppers switched nonrandomly, suggesting that compensatory feeding on complementary resources may have occurred. There was no evidence that behavioral interactions among individuals increased switching rate or lowered fidelity. Overall patterns of resource use varied significantly among grasshoppers during the study, despite frequent switching among resources and a significant tendency to aggregate while feeding. All individuals were polyphagous but differed in relative consumption of available plants. The results suggest that individual grasshoppers express different feeding patterns that are consistent over time and that variation in diet among individual observed in the field may be more than simple sampling error.     

Variation in dietary patterns among and within polyphagous grasshopper species (Orthoptera: Acrididae)

Individuals of two polyphagous grasshopper species,Melanoplus differentialis andSchistocerca albolineata, were reared under standardized conditions to eliminate prior experience as an influence on feeding behavior. Individuals were offered four plants in simultaneous choice tests from 8 to 14 times during development to adulthood. Individual insects varied in the preferred plant and the strength of preference, in the strength of the preference hierarchy among the four plants, and in breadth of feeding in choice tests. Average feeding diversity in choice tests increased significantly in later instars, but the sexes did not differ in feeding pattern. In both species a significant preference for one plant was generally but not invariably associated with a consistent preference hierarchy. Breadth of feeding in all tests was inversely related to the strength of preference and of the preference hierarchy. Although the two species differed in feeding diversity and in the relationship between preference and strength of preference hierarchy, evidence suggests that these differences were due to the specific plants used rather than to true interspecific differences. The existence of such variation under highly standardized rearing conditions suggests that genetic variation for these characteristics may exist in natural populations of polyphagous grasshoppers.     

Host plant predictability and the feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores

Summary Host plant preferences for 34 insect herbivore species are reported. Most polyphagous herbivores feeding on annuals, herbaceous perennials, and woody perennials show distinct preferences for the least abundant plant species among their various host plants. In addition, some populations of widely distributed polyphagous species are much more specialized in their diet than host plant lists alone would suggest. The high level of polyphagy on annuals and herbaceous perennials is suggested to be strongly influenced by the unpredictability of the host plant that is, in turn, controlled by environmental variability. Oligophagous herbivores preferred the least abundant woody perennials on the study sites. Ten of the 22 monophagous herbivores preferred the rarest of all the plant species on the same sites.     

Food intake, conversion efficiency, and feeding behaviour of tobacco hornworm caterpillars given artificial diet of varying nutrient and water content

ABSTRACT Fifth stadium tobacco hornworm caterpillars, Manduca sexta (L.), given artificial diet diluted to varying extents with either cellulose or water compensated for the food's reduced nutrient content by eating more of it. This compensation was, however, in most cases not sufficient to maintain normal growth rates. When the water content of the diet was reduced, the insects ate less than the usual fresh weight of food but maintained their intake of nutrients. Nevertheless, growth rate was impaired. The insects were better able to compensate for dilution of their food with water than with cellulose. The efficiency of conversion of ingested food (ECI) was decreased when the diet was adulterated with cellulose. At moderate dilution (50% nutrient) this was due mostly to decreased approximate digestibility (AD), but at greater dilution (25% and 10% nutrient content) the efficiency of conversion of digested food (ECD) was decreased. ECI was maintained when the water content of the diet was increased to give 50% nutrient concentration, but was decreased when water content was changed more radically (200%, 25% and 10% nutrient diets). This was due mostly to increased metabolic costs (decreased ECD) in all cases. The retention time of food in the gut was progressively decreased (i.e. speed of passage was increased) as nutrients were replaced by cellulose. By contrast, dilution of the diet with water resulted in only slight changes in retention time, except at extreme dilution (10% nutrient content) when retention time was reduced. Compensation of food intake was achieved by spending more (or less) time eating. Video analysis of feeding behaviour showed that there were significant changes in the length of feeding bouts and of interfeed gaps when caterpillars fed on diets of altered composition. For diets diluted with cellulose, changes in bout length and bout frequency contributed substantially to the increased time spent feeding on the adulterated food. For diets diluted with water, however, almost all of the compensatory change in behaviour was due to increased bout length, with bout frequency affected only slightly. This suggests that volumetric feedback contributes principally to the termination of feeding bouts in caterpillars, while nutrient flow may affect both the initiation and termination of feeding.     

Dietary mixing in grasshoppers: switching induced by nutritional imbalances in foods

Dietary mixing by nymphs of Schistocerca americana was studied in the laboratory using artificial diet cakes. Individuals were given either two different inadequate but complementary diet cakes or two adequate and identical ones. When unique flavors (coumarin or NHT) were added to the diet cakes, insects given the inadequate diet treatment switched between cakes more than insects exposed to the adequate diets. This was not the case when no identifying flavors were added.     

Changes in taste receptor cell sensitivity in a polyphagous caterpillar reflect carbohydrate but not protein imbalance

Caterpillars of the polyphagous arctiid, Grammia geneura, have a single cell in the medial galeal sensillum that responds to some sugars and to some amino acids. After conditioning on artificial diet containing unbalanced amounts of carbohydrate and protein, the responses of this cell alter. After protein-biased food it increases slightly, but after carbohydrate-biased food it decreases. Responses to both sucrose and amino acids change in the same direction and the changes would not provide the information necessary to redress a shortage of protein. The lateral galeal sensillum contains one cell that responds to fructose and another responding to some amino acids. The responses of each of these cells in the lateral sensillum are not consistently affected by conditioning diets. After conditioning for 20 h on a protein- or carbohydrate-biased diet, the insects started to feed without delay if offered carbohydrate-biased diet, but only after a pause if given protein-biased diet. This occurred irrespective of the conditioning diet. The duration of the first feeding bout was also longer on carbohydrate-biased diet and the longest bouts followed protein-biased conditioning.     

Macrophytes in tropical shallow lakes: An important food item to benthic entomofauna or an underused resource?

Although macrophytes are known to increase benthic diversity in lakes, the importance of this resource as food for the insects living at the bottom of these ecosystems are still poorly understood. This study assessed the diets of benthic Chironomidae and Campsurus (Ephemeroptera) in two environments: a lake with macrophytes (M+) and another without macrophytes (M?). We expected a differential use of food resources in M+, where plant tissue is particularly important for the aquatic insects’ diet. The diet of 734 individuals from 16 taxa were analyzed. Contrary to expectations, benthic insects consumed low amounts of plant tissue. This finding led us to investigate whether the presence of macrophytes in lakes would indirectly contribute to the benthic insects’ feeding, as more food resources were explored in M+ and a spatial variation of resources intake was observed in this lake, in contrast to the homogeneous feeding in M?. We highlight that macrophytes were responsible for the organic matter build‐up in the sediment, especially at the lake region dominated by these plants, and contributed to increase the deposition of high‐quality amorphous organic matter, which favored taxa in M+ that fed exclusively on this item. The lower diversity of food items exploited in M?, and the Tanypus alga‐based diet in this lake, indicates the low quality of organic resources in its sediment. Although macrophytes were indirectly beneficial for benthic insects’ feeding, we found that this is not an attractive resource for prompt ingestion by most benthic taxa.     

Food mixing by generalist grasshoppers: Plant secondary compounds structure the pattern of feeding

Plant secondary compounds can prevent feeding by phytophagous insects or, if ingested, can be poisonous to them. Less attention has been paid to the additional effects they have on feeding behavior when they are only weakly deterrent or not deterrent at all. Experiments were carried out on the generalist grass-hopperSchistocerca americana. Individuals were presented either with two cakes of high-quality artificial food with a single deterrent compound added or with two cakes, each with a different added deterrent compound. The deterrents consisted of single plant secondary compounds that were either marginally or strongly deterrent. There were profound differences in feeding behavior between those individuals given identical and those given different cake types, including longer feeding bouts on single cakes when a choice of different cake types was available. The behavioral effects demonstrate that the presence of secondary compounds in one food can influence the patterns of feeding on other available foods and suggest that such chemicals could impact foraging activities in a complex manner.     

Host plant influence on chemical defense in conifer sawflies (Hymenoptera: Diprionidae)

Host diet affects the defensive efficacy of Neodiprion sawflies. In laboratory assays with wood ants (Formica obscuripes), secretions from larvae reared on Pinus banksiana were the most repellent, while those from P. resinosa feeders were the least so. This was explained diterpene resin acid content, but not total monoterpene content. The terpene content of regurgitant generally reflected dietary concentrations. Compounds were sequestered nonselectively by larvae. Host-based differences in defense persisted at the behavioral level. P. banksiana feeders regurgitated greater volumes of fluid and were less likely to be disabled or killed by ants in one-on-one interactions than were larvae fed on P. resinosa. The defensive advantages of host diet conflicted with developmental requirements. N. sertifer reared on P. banksiana (the best diet for defense in all cases) had lower cocoon weights (a correlate of fecundity) than those reared on other diets, and had prolonged larval development compared to insects fed P. sylvestris. No such tradeoff was detected in N. lecontei. Larvae of both species strongly preferred P. banksiana over P. resinosa in feeding choice assays.     

Application of diet theory reveals context-dependent foraging preferences in an herbivorous coral reef fish

Dietary preferences of grazers can drive spatial variability in top-down control of autotroph communities, because diet composition may depend on the relative availability of autotroph species. On Caribbean coral reefs, parrotfish grazing is important in limiting macroalgae, but parrotfish dietary preferences are poorly understood. We applied diet-switching analysis to quantify the foraging preferences of the redband parrotfish (Sparisoma aurofrenatum). At 12 Caribbean reefs, we observed 293 redband parrotfish in 5-min feeding bouts and quantified relative benthic algal cover using quadrats. The primary diet items were macroalgal turfs, Halimeda spp., and foliose macroalgae (primarily Dictyota spp. and Lobophora spp.). When each resource was evaluated independently, there were only weak relationships between resource cover and foraging effort (number of bites taken). Electivity for each resource also showed no pattern, varying from positive (preference for the resource) to negative (avoidance) across sites. However, a diet-switching analysis consisting of pairwise comparisons of relative cover and relative foraging effort revealed clearer patterns: parrotfish (a) preferred Halimeda and macroalgal turfs equally, and those two resources were highly substitutable; (b) preferred Halimeda to foliose macroalgae, but those two resources were complementary; and (c) also preferred turf to foliose macroalgae, and those resources were also complementary. Thus parrotfish grazing rates depend on relative, not absolute, abundance of macroalgal types, due to differences in substitutability among resources. Application of similar analyses may help predict potential changes in foraging effort of benthic grazers over spatial gradients that could inform expectations for reef recovery following the protection of herbivore populations.     

Insectivory by Gorilla gorilla gorilla in Southeast Cameroon

Our study extends quantitative analyses of insect-eating by gorillas (Gorilla gorilla gorilla) to Cameroon. During a 2-mo period (May–June 2001), we recorded and analyzed feeding traces on plants and insects and in gorilla feces. We found 180 feeding traces, 17% of which involved insectivory. Seventy-eight percent of the fecal samples (n = 36) contained insects. Ants were found in 61% of the samples, termites in 39%, while 56% of the samples contained remains of other insects. We added 14 new species to the known insect diet of western lowland gorillas. Overall, social insects are predominant. The choice of prey by the Ntonga gorillas gives new clues for the existence of cross-cultural differences among gorilla populations. A comparison of the overall frequency of insectivory with those at other sites in Central Africa indicate a possible effect of forest disturbance on the insectivorous behavior of gorillas. The study suggests the existence of temporal variation in ant- and termite-eating by gorillas.     

The effects of a multispecies sequential diet on the growth and survival of a tropical polyphagous caterpillar

Hylesia lineata Druce (Lepidoptera: Saturniidae) caterpillars are highly polyphagous at the individual level. I examined the effects of a multispecies sequential diet on the performance of the larvae. The experiment included three natural hosts respectively —Casearia corymbosa HBK,Thouinia paucidentata Radlk., andErythroxylum havanense Jacq — used by ovipositing females in proportion as expected by their abundance;C. corymbosa andE. havanense representing the most and least abundant hosts respectively. All possible two-host and single-host diets were tested. Larval survival to the pupal stage (as protected by a field cage) did not differ among all diets. Also, for females, regardless of the identity of the initial host, the final weight and the developmental time depended entirely on the nature of the final host. Male performance, in contrast, only differed in terms of developmental time but again the initial host did not affect the final result. It was concluded that a host switch early in the development of the caterpillars does not affect further growth on the other host plants. Caterpillars that finished their development onE. havanense reached small sizes but females devoted a larger proportion of their resources to egg biomass; as a result female fertility was similar among females regardless of their feeding history. I concluded that the caterpillar's abilities and the female compensatory flexibility may oppose selection for a strong preference hierarchy in the female ovipositing behavior.     

Diet niche relationships among North American grassland and shrubsteppe birds

Summary We consider the dietary relationships of the numerically dominant breeding bird species in four North American grassland/shrubsteppe habitats, sampled over 2–3 consecutive years. Overall, the diets of these species contained primarily insects: orthopterans comprised 29% of the diet biomass, coleopterans 24%, and lepidopteran larvae 23%, while seeds contributed 15% of the average diet. These diets varied substantially, however, and we evaluated several aspects of this variation. Intersexual differences in diets within a species were few, despite the occurrence of significant sexual size dimorphism in several species. For many species, however, there were substantial shifts in dietary composition between years at a given location; overall, the average between-year similarity of species' dietary composition was 70%. Different species exhibited rather different diet patterns. Horned Larks were relatively omnivorous, had broad diet composition niches, and varied considerably in diets between different locations. Meadowlarks were also broad-niched and geographically variable in their diets, but were the most highly carnivorous of the species we considered. Dietary niche breadths of Grasshopper Sparrows were intermediate, but diet composition was rather stable, both between years and between locations. Chestnut-collared Longspurs exhibited narrow diet niches, but substantial annual variation: each year this species apparently exploited a different but limited set of prey types rather heavily. Larger avian predators generally consumed a broader array of functional groups of prey, but did not differ in the taxonomic variety of their diets from small birds. Variation in diet composition between individuals within local populations was considerable; in most species, an individual contained on the average 30–40% of the prey taxa represented in entire population smaples.Patterns of dietary overlap among species were quite inconsistent from year to year at most locations, although at the shrubsteppe site overlap among all species present was consistently quite high. Relatively few cooccurring species pairs exhibited low diet overlap. The degree of diet niche overlap was unrelated to body size differences of the birds, despite as much as six-fold differences in weight among some coexisting species. Relationships of the bird species on another dimension of the trophic niche, prey size, also differed substantially between sites and years. The ranking of co-occurring species by the mean sizes of the prey they consumed generally did not parallel their rankings by body sizes, and in some cases the smallest and the largest species present ate prey of similar sizes. At the shrubsteppe site, all the breeding species exhibited quite similar frequency distributions of prey sizes in their diets.As species number and diversity increased in the breeding avifaunas, diet niche breadths generally decreased, species packing by prey size decreased, and diet composition niche overlap remained relatively unchanged. These trends are in at least partial agreement with predictions of diffuse competition theory, but the patterns were derived from broad inter-site comparisons of overall site averages, and the relationships generally did not hold within local assemblages of species. In general, our attempts to match values of dietary niche features with site characteristics failed to demonstrate close agreement with the predictions of prevailing ecological theory based upon assumptions of resource limitation and competition. Instead, our findings seem generally most consistent with the suggestion that food is not normally limiting to bird populations in these systems, and individuals and populations are exploiting the food resources in an opportunistic fashion, which leads to considerable individual, between-year, and between-location variation in diet compositions and interspecific overlaps.Our attempts to discern clear relationships that accord with theoretical expectations in these avian assemblages are thwarted by our lack of detailed information on the resource base and by the lack of clear tests that will separate alternative hypotheses of community organization and structuring. We suggest that these complications may compromise the findings of many community studies.     

Spatial and seasonal variability in the trophic role of aquatic insects: An assessment of functional feeding group applicability

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Implications of small scale variation in ecological conditions for the diet and density of red colobus monkeys

In this 3-year investigation we documented patterns of density, diet, and activity of red colobus monkeys (Procolobus tephrosceles) in six areas in or near Kibale National Park, Uganda and related these patterns to availability of food resources. There were large differences in the density and behavior of the red colobus among the sites. For example, the red colobus at one site with a diverse plant community of more than 61 tree species, had a diet that included at least 42 species. In contrast, at a second site red colobus spent 92% of their feeding time eating from one species that dominated the tree community. The density of important red colobus food trees varied among sites from 32 trees/ha to 204 trees/ha, and red colobus density ranged from 0.70 groups/km² to 7.41 groups/km². Among sites, red colobus density was related to the cumulative DBH of important food trees, when one apparently anomalous site was excluded, and populations with more plant species in their diets tended to be those that were found at higher densities. Activity budgets of the red colobus populations varied markedly among sites. For example, feeding time ranged among sites from 29 to 55%, and traveling varied from 5 to 20%. When faced with increased foraging demands, red colobus reduced the time spent resting, while the time spent socializing remained fairly constant. Comparative socioecological studies typically contrast species separated by large geographical distances to ensure there is sufficient variation in the environment to detect behavioral responses. The marked differences in ecological conditions and red colobus behavior we documented over short geographical distances, suggests that small-scale contrasts are a useful tool to examine ecological determinants of behavior and community structure.     

Mixed artificial diets enhance the developmental and reproductive performance of the edible grasshopper, <Emphasis Type="Italic">Ruspolia differens</Emphasis> (Orthoptera: Tettigoniidae)

Diet mixing is a common feeding habit among polyphagous insect herbivores and is believed to be advantageous for performance-related factors like growth, survival and oviposition. However, relatively little is known about the influence of artificial diet or their mixtures on the performance of edible insects. We examined the effects of artificial diet mixtures on the developmental and reproductive performance (survival, developmental time, fresh adult weight and female fecundity) of an edible grasshopper, Ruspolia differens (Orthoptera: Tettigoniidae). We raised individuals from eggs and reared newly hatched nymphs to adult stage on six different dietary treatments consisting of a single diet, and mixtures of two, three, five, six and eight artificial diets. More diversified diets resulted in shorter development time and greater adult fresh weight and female fecundity compared to the single diet or less diversified diets. Even with slight diet diversification, survival to adult stage was greatly improved. Overall, these results highlight the potential of diet mixtures in achieving maximum adult weights and female fecundity and shortening development time, information which could be used when designing mass-rearing programs for this edible grasshopper.     

Nicotine moderates the effects of macronutrient balance on nutrient intake by parasitized <Emphasis Type="Italic">Manduca sexta</Emphasis> L

Effects of dietary nicotine and macronutrient ratio on M. sexta larvae were examined. Larvae were fed a carbohydrate-biased, protein-biased or diet having equal amounts of casein and sucrose, with and without nicotine. Without nicotine, larvae displayed compensatory feeding on the low protein diet, but despite consuming more, grew least on this diet. Nicotine at 0.5% had no effect on nutrient consumption. Nicotine at 1.0 and 2.0% reduced overall consumption and thereby also reduced nicotine consumption. Larvae parasitized by C congregata displayed reduced nutrient intake and growth on all diets. Parasitized larvae responded to 1% nicotine similarly to unparasitized larvae. At 0.5% nicotine, they displayed reduced consumption on all diets, possibly due to altered chemoreceptor sensitivity to nicotine. When offered a choice of two diets having different macronutrient ratios, one with and the other without 0.1% nicotine, all larvae preferred the diet lacking nicotine and failed to regulate nutrient intake such that the nutrient intake target, a ratio of nutrients supporting optimal growth, was achieved. Parasitized larvae consumed less nicotine on a fresh weight basis than unparasitized insects, suggesting that the feeding response of parasitized larvae to nicotine minimizes the exposure of nicotine to developing parasites.     

Diversity and abundance of insect herbivores foraging on seedlings in a rainforest in Guyana

1. Free-living insect herbivores foraging on 10 000 tagged seedlings representing five species of common rainforest trees were surveyed monthly for more than 1 year in an unlogged forest plot of 1 km² in Guyana. 2. Overall, 9056 insect specimens were collected. Most were sap-sucking insects, which represented at least 244 species belonging to 25 families. Leaf-chewing insects included at least 101 species belonging to 16 families. Herbivore densities were among the lowest densities reported in tropical rainforests to date: 2.4 individuals per square metre of foliage. 3. Insect host specificity was assessed by calculating Lloyd’s index of patchiness from distributional records and considering feeding records in captivity and in situ. Generalists represented 84 and 78% of sap-sucking species and individuals, and 75 and 42% of leaf-chewing species and individuals. In particular, several species of polyphagous xylem-feeding Cicadellinae were strikingly abundant on all hosts. 4. The high incidence of generalist insects suggests that the Janzen–Connell model, explaining rates of attack on seedlings as a density-dependent process resulting from contagion of specialist insects from parent trees, is unlikely to be valid in this study system. 5. Given the rarity of flushing events for the seedlings during the study period, the low insect densities, and the high proportion of generalists, the data also suggest that seedlings may represent a poor resource for free-living insect herbivores in rainforests.