A comparison of the leaf shredding ability and influence on microbial films of surface and cave forms of <Emphasis Type="Italic">Gammarus minus</Emphasis> Say

Gammarus minus has both cave and surface forms that differ considerably in their morphology. We experimentally compared the ability of these two forms to consume leaves and influence the activity of the microbial films associated with leaves. Both forms of G. minus were effective leaf shredders, and were equally efficient at converting leaves to small particles. We found little relationship between amphipod size and the rate at which they shredded leaves. Both forms of G. minus stimulated the respiration rate of microbes associated with leaves by 32–52%. Cave forms of G. minus had a 15% greater stimulatory effect on microbial respiration. The stimulatory effect of G. minus on microbial respiration is evident in cave streams as well as in the laboratory. It appears that G. minus plays a particularly important role in cave systems by converting leaves to fine particles and stimulating microbial films which are important foods for many cave animals. Handling editor: D. Ryder     

Feeding rates,assimilation efficiencies and growth of two amphipod species on biodeposited material from zebra mussels

1. Accumulation of organic material by the zebra mussel Dreissena polymorpha is assumed to be the source of a biodeposition‐based food web. However, only little is known about the importance of the biodeposited material as a food source and its contribution to increased abundances of macroinvertebrates in the presence of D. polymorpha. 2. Feeding, assimilation and growth of the amphipods Gammarus roeselii and Dikerogammarus villosus on food sources directly and indirectly associated with D. polymorpha (biodeposited material and chironomids) and on conditioned alder leaves were measured. The stoichiometry of carbon, nitrogen and phosphorus of the diets was measured as an important determining factor of food quality. 3. Chironomids had the highest nitrogen and phosphorus contents, alder leaves were depleted in nitrogen and phosphorus, and the stoichiometry of biodeposited material was intermediate. 4. Both amphipod species had highest feeding rates and assimilation efficiencies on chironomids. Gammarus roeselii fed more on biodeposited material than on alder leaves, but assimilation efficiencies were similar; D. villosus also had similar feeding rates and assimilation efficiencies on the two diets. 5. Both amphipod species had highest growth rates on chironomids and lowest growth rates on alder leaves. Both grew at intermediate rates on biodeposited material of D. polymorpha. The growth rates of the amphipod species were related to food stoichiometry. Overall, the invasive D. villosus grew faster than the indigenous G. roeselii. 6. Food resources directly and indirectly associated with D. polymorpha are potential diets for amphipods, providing further evidence for a D. polymorpha biodeposition‐based food web.     

Consequences of the colonisation of leaves by fungi and oomycetes for leaf consumption by a gammarid shredder

1. Leaf litter breakdown by shredders in the field is affected by leaf toughness, nutritional value and the presence of secondary compounds such as polyphenols. However, experiments involving the use of single fungal strains have not supported the assumption that leaf parameters determine food selection by shredders perhaps because of a failure to test for high consumption prior to isolation of fungal strains, overrepresentation of hyphomycetes or the potential effects of accompanying bacteria. In this study, we used bacteria‐free, actively growing fungi and oomycetes isolated from conditioned leaf litter for which a shredder had already shown high consumption rates. 2. Black alder (Alnus glutinosa) leaf litter was exposed to the littoral zone of Lake Constance in autumn, and subsamples were analysed for leaf parameters and consumption by Gammarus roeselii under standard conditions at regular intervals. On dates with a high consumption rate of the exposed leaves, 14 single strains of fungi and oomycetes were isolated, freed of bacteria and grown on autoclaved leaves. 3. Six of eight measured leaf parameters of exposed leaves were significantly correlated with Gammarus consumption rates, with high colinearity among leaf parameters hampering the identification of causal relations between leaf parameters and feeding activity. 4. When single strains of fungi and oomycetes were grown on autoclaved leaf litter, toughness of colonised leaves was always lower than in the control and the content of protein, N and P were increased. There were pronounced strain‐specific effects on leaf parameters. Consumption rates also differed significantly, with nine of fourteen isolates consumed at higher rates than controls and none proving to be a deterrent. Protein and polyphenol content were significantly correlated with consumption rates. Oomycete‐colonised leaves were consumed at similar rates but were of lower food quality than fungi‐colonised leaves. 5. We argue that direct strain‐specific attractant or repellent effects of fungi and oomycetes on consumption by G. roeselii are not important. However, we found indirect strain‐specific role operating via effects on leaf parameters.     

The role of ion regulation in the control of the distribution of Gammarus tigrinus (Sexton) in salt-polluted rivers

Fluctuating salinities at different sites on the German salt-polluted rivers Werra and Weser were compared with extracellular ion levels of specimens of Gammarus tigrinus (Sexton; Amphipoda, Crustacea), collected at the same sites. G. tigrinus regulated haemolymph concentrations of inorganic anions (Cl⁻, SO²⁻ ₄, PO³⁻ ₄) and cations (Na⁺, K⁺, Mg²⁺, Ca²⁺) during fluctuations of salt pollution in the upper Weser. This capacity to regulate varying levels of salt pollution in the upper Weser, correlated well with the distribution of the brackish amphipods in this river ecosystem. G. tigrinus tolerated periods of Na⁺ and Cl⁻ stress (>380 mmol l⁻¹) without compensating these maxima by regulating extracellular Na⁺ and Cl⁻. However, during such bursts of Na⁺ and Cl⁻ stress in Werra and Weser, the ability to regulate extracellular [K⁺] at river water K⁺ stress of ≥6.0 mmol l⁻¹ may explain why this brackish species has been more successful in these rivers than its competitors like Gammarus pulex. The present investigation demonstrates that the water salinity affects the [NO⁻ ₃] in the haemolymph of G. tigrinus. With increasing hypo-osmotic stress the animals accumulate increasing amounts of NO⁻ ₃. A simultaneous increase in stream water [NO⁻ ₃] causes an additional accumulation of NO⁻ ₃ in the haemolymph. The high extent of accumulation indicates that active ion transport systems may be involved. The accumulation of NO⁻ ₃ in the haemolymph has low physiological consequences to G. tigrinus, but when hypo-osmotically stressed under anoxic conditions, nitrite formed by the reduction of nitrate may have an adverse affect on the metabolism of G. tigrinus. Accepted: 4 October 1999     

Distribution and habitat utilization in two atyid shrimps (Crustacea: Decapoda) in rivers of Nuku-Hiva Island (French Polynesia)

The distribution of Atyoida pilipes (Newport, 1847) and Caridina weberi De Man, 1892 was studied at two spatial scales on Nuku-Hiva Island (Marquesas, French Polynesia). At the stream reach scale, the two species were present at 40 and 39 sites, respectively, out of a total of 49 sites sampled. These two taxa represented more than half of the invertebrates collected at 9 sites. There was no difference in distribution between the species at the reach scale. However, clear differences appeared between the two species at the microhabitat scale. A. pilipes was more abundant in lotic habitats, even waterfalls, while C. weberi was associated with accumulations of leaves in lentic habitats. A. pilipes, particularly males, were tolerant to parameters such as water velocity, leaves, and the presence of algae and rocks. C. weberi had a more restricted altitudinal distribution and were scarce upstream from large waterfalls. These differences were linked to differences in climbing ability rather than the effect of chemical parameters, such as oxygen content. Our results suggest that little competition for space or food resources occurs between the two species. Both are feeding on small detritus particles, but through differing mechanisms and in differing habitats.     

Production,nutrient dynamics and initial decomposition of floating leaves of Nymphaea alba L. and Nuphar lutea (L.) Sm. (Nymphaeaceae) in alkaline and acid waters

Production, turnover and nutrient dynamics of floating leaves of Nymphaea alba L. and Nuphar lutea (L.) Sm. were studied in four aquatic systems in The Netherlands, differing strongly in water quality. Production was 108–447 g AFDW.m⁻² for N. lutea and 319–348 g AFDW.m⁻² for N. alba. Turnover ranged from 3.6 to 4.4 without much difference between the sites and the species. During senescence 60–70% of the N and P from the leaves was resorbed by both of the plant species. The nutrient flow from the floating leaves into the detritus food chain differed considerably between the species and sites studied, mainly because of the differences in production. The data suggest that production is strongly influenced by the environment, whereas turnover and nutrient resorption during senescence seem to be plant characteristics. There was little difference in dynamics of the chlorophyll-α concentration in the leaves of the two species, irrespective of the growing site. Floating leaves of both species lost about 70% of their area due to fragmentation in the alkaline waters, whereas in the acid water very little fragmentation was observed.     

An energy budget for Simocephalus vetulus (O. F. Muller) (Crustacea : Cladocera)

We discuss the energetics of a cladoceran, Simocephalus vetulus at different temperatures (8.0 ± 1.0, 15.0 ± 1.0, 21.0 ± 1.0 and 28.0 ± 1.0 °C) and food (Chlamydomonas sp.) concentrations (25 × 10³, 50 × 10³, 75 × 10³ and 100 × 10³ cells ml⁻¹). Increase in temperature accelerated ingestion and, to some extent, oxygen consumption. The study revealed a high reproduction efficiency in S. vetulus. Net growth efficiency (ECI) was higher (13.17–41.18%) in pre-adults than in adults (2.71–8.40%). The assimilated energy (A) increased with increasing food concentration at all temperatures. Assimilation efficiency (AD) decreased with increasing food concentrations. The energy used for growth (P) was nearly constant at all food levels because the egested energy increased and assimilation efficiency decreased as food concentration increased.     

Feeding Patterns and Dietary Profile of Nocturnal Southern Woolly Lemurs (Avahi meridionalis) in Southeast Madagascar

Folivory has evolved in all primate radiations but the relative importance of resource quantity and nutritional quality of food is controversial. To understand how food abundance and different nutrients affect a folivorous diet, we investigated food composition of the leaf-eating lemur Avahi meridionalis. From June to December 2004, we conducted 26 nights of focal observation (256.1 h) in Sainte Luce pluvial littoral forest (southeast Madagascar) and recorded feeding behavior of 4 radiocollared individuals. Within the subjects’ home ranges, we recorded vegetation characteristics (morphospecies, phenology, diameter at breast height) and sampled food and nonfood items for chemical analyses. A. meridionalis did not eat fruit but only leaves and flowers and did not base their choice on food abundance. Adult leaves eaten were higher in easily soluble protein than adult leaves that were not consumed. The subjects ate young leaves and flowers as soon as they became available. These young leaves contained the same concentrations of soluble protein, higher concentrations of crude protein, and lower concentrations of acid detergent fiber and sugar than mature food leaves. A. meridionalis ate leaves with condensed tannins, alkaloids, and intermediate concentrations of polyphenolics. Contrary to previous studies that considered Avahi spp. a specialist, A. meridionalis acted as leaf-eating generalists with moderate selectivity, based on nutritional quality and tolerance of a wide array of plant secondary metabolites. This illustrates the dietary flexibility within a single genus of primates that seems to be driven by environmental constraints rather than morphological or physiological adaptations.     

Competition between perch (Perca fluviatilis) and ruffe (Gymnocephalus cernuus): the advantage of turning night into day

1. The outcome of interspecific competition for food resources depends both on the competitors’ sensory abilities and on environmental conditions. In laboratory experiments we tested the influence of daylight and darkness on feeding behaviour and specific growth rate (SGR) of two species with different sensory abilities. 2. We used perch (Perca fluviatilis) as a visually orientated, and ruffe (Gymnocephalus cernuus) as a mechano‐sensory oriented predator and tested their growth rates and behaviour under conditions of interspecific and intraspecific competition. Three different foraging conditions were used: food supplied (i) only during the day, (ii) only during the night or (iii) during both day and night. 3. In perch neither SGR nor feeding behaviour were influenced substantially by interspecific competition during daylight. During darkness their foraging behaviour changed markedly and their access to the food source as well as their SGR were negatively affected by the presence of ruffe. 4. Ruffe's foraging behaviour did not change during either day or night with interspecific competition. During the night ruffe's SGR was higher with interspecific competition, probably because of a release from intraspecific competition and the competitive inferiority of perch during the night. 5. Because of its seonsory abilities ruffe feeds predominantly at night, thereby reducing competitive interference from perch.     

Resource use by howler monkeys (Alouatta palliata) in the rain forest of Los Tuxtlas,Veracruz, Mexico

The feeding preferences of howler monkeys at their northernmost distribution in the Neotropics are reported for an annual cycle. A remarkable selectivity for 27 species representing 15 families was observed. The Moraceae and Lauraceae plant families were the most important in the diet. The howlers spent an almost equal proportion of their feeding time eating leaves and fruit, and displayed a marked preference for young leaves and mature fruit. The consumption of different plant parts was markedly seasonal and the howlers’ ranging behavior was closely associated with the availability of young leaves and mature fruit. Their home range was unusually large (ca. 60 ha) for howlers and the food species exploited occur at very low densities (93%, ≤ 4 ind/ha). They chose food items richer in protein and energy. Alkaloid compounds, present in some of the leaves, play a secondary role in their dietary selectivity.     

Amphipod (Gammarus minus) responses to predators and predator impact on amphipod density

Recent theoretical work suggests that predator impact on local prey density will be the result of interactions between prey emigration responses to predators and predator consumption of prey. Whether prey increase or decrease their movement rates in response to predators will greatly influence the impact that predators have on prey density. In stream systems the type of predator, benthic versus water-column, is expected to influence whether prey increase or decrease their movement rates. Experiments were conducted to examine the response of amphipods (Gammarus minus) to benthic and water-column predators and to examine the interplay between amphipod response to predators and predator consumption of prey in determining prey density. Amphipods did not respond to nor were they consumed by the benthic predator. Thus, this predator had no impact on amphipod density. In contrast, amphipods did respond to two species of water-column predators (the predatory fish bluegills, Lepomis macrochirus, and striped shiners, Luxilus chrysocephalus) by decreasing their activity rates. This response led to similar positive effects on amphipod density at night by both species of predatory fish. However, striped shiners did not consume many amphipods, suggesting their impact on the whole amphipod “population” was zero. In contrast, bluegills consumed a significant number of amphipods, and thus had a negative impact on the amphipod “population”. These results lend support to theoretical work which suggests that prey behavioral responses to predators can mask the true impact that predators have on prey populations when experiments are conducted at small scales. Received: 21 March 1997 / Accepted: 15 December 1997     

Effects of pH,humic substances and animal interactions on survival and physiological status of Asellus aquaticus L. and Gammarus pulex (L.)

Summary The mortality and physiological status (body water content) of Asellus aquaticus (Isopoda) and Gammarus pulex (Amphipoda) were measured after 25 days exposure in 20 natural streams with a pH range of 4.3–7.5 and a colour range of 8–280 mg Pt L^–1. In addition, the effects of keeping the animals as single species or together were studied. The response of Gammarus to low pH was an increased mortality and lower physiological status of surviving individuals in streams with a pH lower than 6.0. In Asellus the physiological status was correlated with pH, while the mortality was not pH dependent. The effects of humus on the physiological status of Asellus was significant when fitted to a second order polynomial function. The influence of humus can, however, be regarded as small relative to pH. The interactions between the species could be described as asymmetric under optimal conditions of high pH and low humus concentrations, where the presence of Gammarus decreased the survival and physiological status of Asellus. Acid stress did not seem to reverse the direction of this asymmetry, but the presence of Gammarus improved the physiological status of Asellus at pH lower than 6.0. Since the presence of Asellus did not increase the mortality or decrease the physiological status of Gammarus, this could be explained by Asellus feeding on Gammarus that died from physiological stress solely. This mechanism suggests that food quality, and thus effects of diffuse competition, can be important for the ability to withstand acid stress. The results, though, give no support for the hypothesis that competition from Asellus is important for the disappearance of Gammarus during the acidification of streams.     

Importance of fungi in the diet of Gammarus pulex and Asellus aquaticus

An important component of the interaction between macroinvertebrates and leaf litter in streams in the extent to which consumers can differentiate between undecomposed and decomposing leaves. The detritivores Gammarus pulex and Asellus aquaticus fed preferentially on conditioned rather on unconditioned leaf material. Growth in A. aquaticus was significantly reduced when unconditioned leaves were provided, but in G. pulex no significant effect of conditioning on growth was observed. The capacity of G. pulex to tolerate reductions in food quality seems to be a consequence of a compensatory system in which respiration rates change to compensate for reductions in food quality. In this way a constant growth rate is maintained. Increases in ingestion rates to compensate for low quality food were not observed.     

Further laboratory studies on the interspecific competition between Helisoma duryi (Wetherby) and the intermediate hosts of schistosoma mansoni sambon: Biomphalaria alexandrina (Ehrenberg) and B. camerunensis (Boettger)

Biological control of schistosomiasis by means of introduction of a competitor, Helisoma duryi, of the intermediate host snails has been proposed. In the present laboratory studies the competitive relationships between H. duryi and two Biomphalaria species were investigated both when direct interference between the two competing species was possible and when not. A reduction of growth and reproduction of Biomphalaria was found when direct interference was involved but the effect of competition was lessened when the two species were separated by a mesh.The data showed that despite continuous presence of food in the aquaria food competition was involved in the direct interference between the two species, while the role of chemical interactions could not be evaluated conclusively from the present experiments.The role of food is discussed.     

Homeostatic responses in the gut of Porcellio scaber (Isopoda: Oniscidea) optimize litter degradation

In order to examine the influence of differences in food conditions on gut characteristics in Porcellio scaber, pH-manipulated and microbially inoculated leaf litter from three different tree species were offered. Microbial activity was clearly influenced by the pH levels of the leaves. Analyses of the pH levels in the gut indicated the ability of P. scaber to buffer the pH value in the intestinal tract to about 5.5–6.0 in the anterior hindgut, and to about 6.0–6.5 in the posterior hindgut. The pH levels of the gut sections remained in this range, within a range of food pH from 4.0 to 7.5, no matter what kind of leaves the animals were fed. Homeostatic responses to changes in food pH guarantee optimized digestion of leaf litter. However, when the pH level of the litter dropped below 3.5, P. scaber was not able to maintain the pH conditions in the gut. Furthermore, microorganisms colonizing the litter biased the pH level in the anterior hindgut where digestive processes mainly take place. These results indicate a decline of litter quality with regard to the nutrition of terrestrial isopods, caused by acidification and consequently reduced microbial activity. Accepted: 19 July 1997     

A laboratory study on feeding plasticity of the shredder <Emphasis Type="Italic">Sericostoma vittatum</Emphasis> Rambur (Sericostomatidae)

Since litter input and availability of leaves in many streams is highly seasonal in Portugal, we investigated whether Sericostoma vittatum, a typical shredder, was able to grow using alternative food sources. To test this hypothesis we fed S. vittatum with Alnus glutinosa (alder, CPOM, coarse particulate organic matter), leaf powder from A. glutinosa and Acacia dealbata and FPOM (fine particulate organic matter) from a 5th and a >6th order river, the macrophyte Myriophyllum aquaticum and biofilm. Growth in S. vittatum was significantly influenced by the food item given (ANOVA, P = 0.0082). The food item promoting the highest growth was A. glutinosa, in the form of FPOM (6.48% day⁻¹) and CPOM (4.24% day⁻¹); all other forms of FPOM and biofilm provided relatively low growth rates (0.77–1.77% day⁻¹). The macrophyte M. aquaticum was also used as food source by S. vittatum and promoted intermediate growth (1.96% day⁻¹). Neither nitrogen, phosphorus nor caloric content was correlated with growth. However, since higher growth was achieved with alder, in the form of CPOM and FPOM, we concluded that the chemical content of food was more important for S. vittatum than the physical form of such food. This may partially explain why shredders are able to survive when leaves are scarce in streams. Handling editor: K. Martens     

Host-herbivore studies of Stenoscepa sp. （Orthoptera：Pyrgomorphidae）, a high-Ni herbivore of the South African Ni hyperaccumulator Berkheya coddii （Asteraceae）

Nymphs of Stenoscepa sp. feed on leaves of the Ni hyperaccumulator Berkheya coddii at serpentine sites in Mpumalanga Province, South Africa. These sites contain Ni hyperaccumulators, Ni accumulators, and plants with Ni concentrations in the normal range. We conducted studies to： （i） determine the whole-body metal concentration of nymphs （including those starved to empty their guts）; （ii） compare Stenoscepa sp. nymphs against other grasshoppers in the same habitat for whole-body metal concentrations; and （iii）compare the suitability of Ni hyperaccumulator and Ni accumulator plants as food sources for Stenoscepa sp. and other grasshoppers. Stenoscepa nymphs had extremely high whole-body Ni concentrations （3 500μg Ni/g）. This was partly due to food in the gut, as starved insects contained less Ni （950 pg Ni/g）. Stenoscepa nymphs survived significantly better than other grasshoppers collected from either a serpentine or a non-serpentine site when offered high-Ni plants as food. In a host preference test among four Berkheya species （two Ni hyperaccumulators and two Ni accumulators）, Stenoscepa sp, preferred leaves of the Ni hyperaccumulator species. A preference experiment using leaves of three Senecio species （of which one species, Senecio coronatus, was represented by both a Ni hyperaccumulator and a Ni accumulator population） showed that Stenoscepa sp. preferred Ni accumulator Senecio coronatus leaves to all other choices. We conclude that Stenoscepa sp. is extremely Ni-tolerant. Stenoscepa sp. nymphs prefer leaves of hyperaccumulator Berkheya species, but elevated Ni concentration alone does not determine their food preference. We suggest that the extremely high whole-body Ni concentration of Stenoscepa nymphs may affect food web relationships in these serpentine communities.     

Predicting the Competitive Regime of Female <Emphasis Type="Italic">Colobus vellerosus</Emphasis> from the Distribution of Food Resources

We examined the spatial and temporal distribution of the foods of ursine colobus (Colobus vellerosus) at Boabeng-Fiema, Ghana as a means to predict the monopolizablity and usurpability of their food resources. Recent evidence suggests that food may not be limiting for folivorous primates, and that male sexual coercion may be a more important influence on folivore social organization. To address the question, we collected focal data on the feeding behavior of adult females and males over 11 mo (September 2000-August 2001) on 2 groups: WW (n = 31–33 individuals) and B (n = 8–16 individuals). We also conducted phenological monitoring and a tree survey of the two-group home ranges to establish food availability and distribution. We used 2 behavioral or organism-defined indicators of feeding behavior to assess potential resource contestability: food site residence time and distance moved between food sites. The colobus fed on a high diversity of species, most of their food trees were not clumped in distribution, within-tree interfood distances were short, and food trees were large. The only condition associated with the potential for monopolization was low food tree density. However, low food tree density may be offset by the colobus’ use of large trees. Taken together, the ecological and behavioral indicators suggest the food resources of Colobus vellerosus had a low potential for monopolization. Our results also indicate mature leaves had the longest food site residence time, which may suggest they should be the most usurpable plant part, though their presumed low quality and high abundance probably counteracted the effect. The pattern implied the potential for direct feeding competition among Colobus vellerosus at Boabeng-Fiema was low and agonistic interactions over food are not expected. Instead, a group size effect on feeding efficiency should be a more predominant influence on feeding efficiency, if food is limiting for the species.     

Gut passage times in Gammarus pulex (Crustacea,Amphipoda) and aspects of summer feeding in a stony stream

Gut passage times in Gammarus pulex (Crustacea, Amphipoda) were calculated by counting and measuring voided faecal pellets. Following feeding on elm leaves conditioned in a stream, or summer fine detritus from the same source, calculated gut passage times were usually short, about 2 h or less at 15 °C. Using harder oak and beech leaves, not previously conditioned in the stream, apparent gut passage times were much longer, 8–48 h, but since intermittent feeding occurred these figures are not comparable to the others.In connection with summer feeding of the animal in the stream, the microbiological status of the fine detritus there was studied. It contained fungal hyphae, which were mostly empty, viable fungal spores of terrestrial derivation, and bacteria, all largely carried on fragments of vascular plant tissues. In experiments, such fungal spores (of Aureobasidium pullulans and of Mucor sp.), incorporated into fine detritus, were not digested by the animal. The bacterium Bacillus cereus was present in the fine detritus and when the latter was consumed it survived passage through the gut of the animal. Using B. cereus as a proportion marker it was concluded that other unicellular bacteria in the fine detritus were not digested either. In other experiments, not immediately related to the summer season, nutrient extraction by the animal from hyphae of the fungus Nectria lugdunensis was examined; it probably occurs through sub-microscopic pores connecting the cells. This mode of extraction may apply when the animal consumes vascular plant tissues.     

Resource exploitation of larvae ofGastrophysa atrocyanea Motschulsky andgalerucella vittaticollis Baly (Coleoptera: Chrisomelidae) under a limited resource condition

Resource exploitation by and intraspecific competition in larvae of Gastrophysa atrocyanea and Galerucella vittaticollis were investigated in field and laboratory experiments. Larvae of both species frequently suffered from food shortages in the field. When G. atrocyanea larvae suffered from a food shortage, severe intraspecific competition occurred because of lack of predation and parasitism. This exploitive competition was caused by a local food shortage of the host plant. Individuals survived by fast exploitation when food became abundant (contest type competition). the G. atrocyanea larvae were wasteful of the food resource, and no mechanism by which to economize on the utilization of the resource was acquired because of their exploitation of the abundant resource. In contrast, the G. vittaticollis population probably is regulated by extrinsic factors such as predation and parasitism. Those larvae grew into smaller adults than those of G. atrocyanea under a food shortage, so that their wasted food consumption was lower than that of G. atrocyanea. Although intraspecific competition was similar to that for G. atrocyanea, it was not as severe. The food for G. vittaticollis was apt to be appropriated by other wasteful exploitators such as G. atrocyanea, which was superior in resource exploitation; therefore G. vittaticollis frequently suffered a food shortage. Consequently selection in relation to tolerance to starvation became more acute for G. vittaticollis than for G. atrocyanea, and individuals of G. vittaticollis that could endure starvation better may have been selected.