Interference competition in ant-lion (Macroleon quinquemaculatus) larvae

Abstract.

• 1 The effects of density, feeding regime, and body size on interference competition in the pit-digging larvae of the ant-lion Macroleon quinquemaculatus (Hagen) were investigated in laboratory experiments.
• 2 Competition had little effect on the pit size of winners but losers constructed much smaller pits than isolated larvae. Losers were less likely to dig or maintain pits and more likely to move than winners.
• 3 Competition was much stronger between well-fed larvae than between hungry ones, and well-fed competitors showed reduced growth rates. Well-fed larvae orientated themselves so that they could throw sand into their neighbour's pit whereas hungry larvae faced away from each other. Differences in hunger level reversed the competitive advantage of larger larvae only when individuals were of similar size.
• 4 Cannibalism was density-dependent and most frequent in hungry, similar-sized, larvae; the smaller larva was usually the victim.
• 5 Displays/challenges between larvae affected the distance between pits. Body size was the main determinant of contest outcome though pit ownership and hunger level also had an effect.

     

Behavioral Plasticity and Variation in Pit Construction of Antlion Larvae in Substrates with Different Particle Sizes

Behavioral plasticity allows animals to maximize their fitness in a variety of environmental conditions. Trap‐building predators represent case studies in such plasticity as the characteristics of their traps are dependent upon the substrate available. We investigated the effect of sand particle size on pit construction in antlions (Euroleon nostras), sand‐dwelling insect larvae that build pitfall traps to capture prey. The pit construction behavior of the species comprises six stages. When antlions were exposed to different sand particle sizes, their behavior differed in terms of the occurrence and duration of particular stages and in the frequency of jerks produced during sand tossing. Jerk frequency was negatively correlated with sand particle size and also changed during pit construction. Furthermore, at larger particle sizes, individuals occasionally constructed irregular traps with a figure of eight shape, and they crossed the center of the truncated cone during deepening. In the largest substrate, particle size of antlions did not construct pits. Our results demonstrate that variation in traps under differing environmental conditions stems directly from behavioral plasticity in this species.     

Substrate selection for pit making and oviposition in an antlion, Myrmeleon bore Tjeder, in terms of sand particle size

The larvae of the pit‐making antlion Myrmeleon bore Tjeder live in open sand in riverbeds with a substratum consisting of various particle sizes. We analyzed the spatial distribution of their pits in a sandy floodplain to determine their larval and adult responses to the heterogeneous substrate. The spatial distribution pattern of their pits had an aggregated distribution, and there was a significant positive correlation between pit density and the ratio of medium‐size sand particles to total weight of sand. We examined the size of sand particles selected in the larval pit‐building behavior and the oviposition behavior of the adult. Both larvae and adults selected medium‐size sand particles. The larvae of M. bore are relatively sedentary predators and rarely move great distances. Thus, the present results suggest that habitat selection by adult females is a major factor causing the aggregative distribution of the pits.     

Patterns of resource distribution among conspecific larvae in two fruit fly species: Anastrepha fraterculus and Ceratitis capitata (Diptera: Tephritidae)

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An experimental study on the foraging behavior of a pit-building antlion larva,Myrmeleon bore

Foraging behavior of a pit-building antlion larva, Myrmeleon boreTjeder was investigated experimentally to elucidate the relation between the feeding level and pit relocation.

1. In artificial sands constructed in the field the 3rd instar larvae of M. bore rarely changed the positions of their pits, though several antlions had moved actively until they constructed pits. The average feeding rate was 0.3 prey/day/pit, and about 60% of prey captured were ants.
2. To examine whether or not M. bore larvae concentrate into the area where they can capture more prey, 8 antlions were released into each of 6 boxes filled with sand. I divided the sand surface of each box into two half areas, then gave prey to the pits built in a half area and gave no prey to the pits built in the other half. During the 50-day observation period, nonfed antlions never moved into the area where prey were given.
3. The 3rd instar larvae were reared separately without food. Even under starved conditions they rarely relocated their pits until dealth. The average duration of survival period was 83.9 days.
4. The experimental results indicate that M. bore larvae adopt a tactic of sedentary ambushing. These larvae exhibit low movement rates which are independent of prey capture rates.

     

Density-dependent regulation of Aedes cantans (Diptera: Culicidae) in natural and artificial populations

Abstract.

• 1 Mortality and size variation (siphon length) in Aedes cantans larvae were examined in natural populations in northern England in 1989 and 1990.
• 2 Under crowded conditions, density-dependent competition led to reduction in the size of both larvae and adults and increased larval mortality.
• 3 Larvae were also maintained in cages in the field at different densities. Results paralleled those for the natural populations in the ponds; larvae maintained at high densities showed increased mortality and reduced size.
• 4 Possible density-dependent factors leading to mortality and size reduction include cannibalism and contact inhibition leading to food shortages.
• 5 The main density-independent factor contributing to larval mortality was habitat desiccation.

     

Effect of Population Density and Female Body Size on Number and Size of Offspring in a Species with Size‐Dependent Contests over Resources

When size‐dependent contests over resources influence reproductive success, the trade‐off between number and size of offspring depends on the frequency of contests. Under these circumstances, clutch size should decrease and offspring size should increase as contests become more frequent. We tested these predictions with the burying beetle Nicrophorus pustulatus through manipulation of rearing densities. Burying beetles reproduce on small vertebrate carcasses, a rare but high quality food source for the larvae. Large beetles are more likely to win contests over carcasses and gain exclusive access to a carcass. The winner of a contest kills eggs and larvae already present on a carcass. As a result of the rarity of carcasses, burying beetles are unlikely to breed more than once. As predicted, brood size of N. pustulatus decreased with increasing rearing density. Despite a negative correlation between brood size and larval mass, larval mass did not increase with increasing rearing density. This may be due to the special biology of N. pustulatus which can use snake eggs for reproduction. Potentially larger supply of resources and generally small population densities of N. pustulatus may weaken selection on body size and thus the correlation between brood size and larval mass. As size‐dependent constraints can limit reproductive phenotypes, we examined whether female size influenced reproductive phenotype. Small females produced larger broods with smaller, but more variable, offspring than large females. Mechanical constraints of egg size seem an unlikely explanation for the differences because burying beetles can compensate for small egg size through parental care. Energetic constraints may impact small females because body mass and brood size of small females decreased with increasing density. Yet, at all density levels small females produced larger, not smaller, broods than large females. The larger and more variable broods of small females seem to be in agreement with a bet‐hedging strategy.     

Larval development of two parasitic flies (Conopidae) in the common host Bombus pascuorum

Abstract.

• 1 Whenever parasitism by more than one female occurs, larvae of parasitoids not only have to resist host defence but also face competition with other (unrelated) larvae. Competition is particularly important in solitary parasitoids where only one larva is able to complete its development. Such a situation is found in Conopidae (Diptera) parasitizing adult bumble bees where larvae of two species of conopid flies, Sicus ferrugineus L. and Physocephala rufipes F. often compete within the common host Bombus pascuorum Scopoli. This study analysed the larval development of the two species and asks how competition among larvae may be regulated.
• 2 Parasitized workers of B.pascuorum were caught in the field and kept according to different experimental schedules in the laboratory. This provided stage-structured data for the temporal course of development of the parasitic larvae. For the analysis, a simulation model was constructed that estimated the duration of all parasitic stages (Manly, 1990, first method). In both species the egg stage was found to be approximately 2 days, first instar 3 days, second instar 4 days, and third instar 3 days. The total development time is an estimated 10.8 days from oviposition in S.ferrugineus and 11.4 days in P.rufipes. S.ferrugineus develops faster in the beginning, probably because of its larger egg size, whereas P.rufipes pupates at larger size. First-instar larvae of both species possess strong, pointed mandibles.
• 3 The success of conopid larvae seems only marginally affected by host defence, for a single larva per host almost always completes development. Under competition, however, mortality rate increases substantially, and most larvae die in their first instar. Moreover, they show signs of melanization. The estimates for developmental times and the patterns found in this study suggest that conopid larvae seem capable of physical attacks, particularly during the first instar, when elimination of competitors is most common, and that S.ferrugineus has a time advantage because of its faster early development. Because most studies have previously been carried out with hymenopteran parasitoids, this study provides new information about the other large group of parasitoid insects, the Diptera, and demonstrates convergent patterns.

     

Mechanisms of interference competition in the western tree-hole mosquito, Aedes sierrensis

Abstract.

• 1 Mechanisms of interference competition affecting Aedes sierrensis (Ludlow) were investigated in laboratory microcosms using reconstituted, natural tree-hole detritus as food and substrate.
• 2 Pupation success, larval development time, and pupal weight were all affected by larval density but not by surface area:volume ratio of the microcosm.
• 3 Mature fourth instars affected pupation success, pupal weight, and development time of developing cohorts separated from them by a 2 μm pore membrane impermeable to bacteria, indicating that chemical competition is proportional to density of at least older instars.
• 4 Cannibalism does not occur, regardless of the presence or absence of food or physical complexity.
• 5 Fourth instars inhibited each other's feeding at densities of 64 larvae/ml or greater. Feeding inhibition due to physical contact (encounter competition) was abated but not eliminated by increasing physical complexity of the microcosm by the addition of leaves.
• 6 Levels of detritus and larval density both affect weight of day-1 fourth instars. Resistance to encounter competition is proportional to fourth instar weight and weight-specific resistance is correlated with rank weight within a developing cohort.
• 7 At densities around the population equilibrium in nature, encounter competition should be taking place, especially in tree holes with few leaves or other large litter.

     

Interspecific competition among larvae ofHemipyrellia ligurriens (Calliphoridae) andBoettcherisca formosensis (Sarcophagidae) (Diptera)

1. The effects of larval rearing density and species relative proportions on life-history parameters of two necrophagous Diptera, Hemipyrellia ligurriens (Wiedemann) (Calliphoridae) and Boettcherisca formosensis Kirner and Lopes (Sarcophagidae), were investigated in mixed cultures. Larval rearing density had a significant effect on larval to adult survivorship, duration of immature development, adult size and relative performance (measured by the composite index of performance, r′) of both species. However, species relative proportions affected adult size of both flies and the duration of immature development of B. formosensis only.
2. B. formosensis had a higher survivorship than H. ligurriens in all mixed cultures and showed a similar survivorship pattern to that in pure cultures. By contrast, survivorship of H. ligurriens was lower in mixed than in pure cultures.
3. H. ligurriens adults reared from mixed cultures were smaller than those from pure cultures of comparable density, but B. formosensis adults from pure and mixed cultures were of similar size.
4. The results suggest that competition between B. formosensis and H. ligurriens larvae was asymmetric and the former was the superior competior.
5. At low larval densities in mixed cultures, the presence of H. ligurriens enhanced the performance (as measured by r′) of B. formosensis, a consequence of suspected interspecific facilitation of larval growth.

     

Disadvantages of living in a populous neighborhood for sit-and-wait predators: Competition for space reduces pit-trap size in antlion larvae

The study of how trap design responds to biotic and abiotic conditions can help to understand the selective forces affecting the foraging of trap-building organisms. We experimentally tested whether pit design can be modified by intraspecific competition for space in larvae of Myrmeleon crudelis, a common sit-and-wait predator that digs conical pit traps in the soil to capture walking arthropods. In a tropical forest in Costa Rica, we measured pit dimensions, larval body size, and the level of competition (i.e., density of neighboring traps) in 40 antlion larvae. These larvae were then taken to the laboratory and allowed to build new traps in individual containers. We measured within-individual changes in the size of traps in the field and in the laboratory, and related these to the level of competition experienced in the field. Larvae with relatively high levels of competition in the field showed a greater increase in the size of their pits in the laboratory. This change was independent of larval size. Larvae with none or few neighbors in the field showed little change in their pit sizes, whereas those with higher competition levels increased their diameter and depth up to 1,400% and 1,000%, respectively. Our results demonstrate that, at least in high-density aggregations, pit design is restricted by competition in addition to the constraints imposed by body size. This work suggests that biotic interactions can play a role in the design of extended phenotypes in sit-and-wait predators that live in dense aggregations.     

The effect of sublethal neogregarine infections in the spruce needleminer, Epinotia tedella (Lepidoptera: Tortricidae)

Abstract.

• 1 A hitherto unknown factor causing cyclic reduction in population fertility of Epirzotia tedella (Cl.) was identified as sublethal infections of the neogregarine Mattesia sp.
• 2 Infection probably takes place when larvae enter hibernation, but does not develop into spore production until the formation of pupae and adult moths.
• 3 At the individual level, the sublethal neogregarine infection causes a slight delay in adult emergence, a decreased adult lifespan, a suppression of egg development, and thus a reduction in fertility.
• 4 At the population level, fertility reduction has a delayed density dependent component and, through correlation, Mattesia is assumed to be causative. Since fertility reduction is a key-factor in E.tedella, Mattesia may even be a dominant factor in the dynamics of this species.
• 5 Mattesia infections were found in no other dominant insect species at the locations, and consequently the interaction between E.tedella and Mattesia sp. seems to be specific. Such delayed density dependent interaction can create host oscillations, as seen in many forest insects. However, host-parasitoid interaction alone leads to oscillations with the observed period of 6–8 years length, and the neogregarine infections apparently act in parallel with the parasitoids and enhance the amplitude of the oscillations.

     

Substrate particle size affects pit building decision and pit size in the antlion larvae Euroleon nostras (Neuroptera: Myrmeleontidae)

Abstract. The larvae of the antlion Euroleon nostras are pit-builders, constructing pitfall traps in loose sand. The number of pits and the pit diameter are recorded when larvae are kept in substrates with different particle sizes. The most convenient pit-building sand fractions are two fractions with fine sand (≤ 0.23 mm; 0.23–0.54 mm). The largest pits are constructed in sand with a particle size of 0.23–0.54 mm. In this sand fraction, larvae of all three instars most readily build pits. No pits are constructed in sand with a particle size greater than 1.54 mm. First- and second-instar larvae avoid building pits in substrates of particle size 1–1.54 mm, but third-instar larvae construct pits in this sand fraction. It is assumed that the antlion is capable of distinguishing between substrate types and this hypothesis is tested by giving larvae the choice of building a pit in one of four particle-size fractions. Larvae of all three instars prefer to build pits in the fraction with a particle size of 0.23–0.54 mm. Only third-instar larvae build pits in all four fractions, but only occasionally in the coarser fraction.     

Larval cannibalism during the late developmental stages of a facultatively gregarious encyrtid endoparasitoid

• 1 The larvae of many gregarious parasitoid species are usually non‐aggressive when they develop in or on a host, but those of Metaphycus flavus are one of the few exceptions known. Herein we describe their aggressive behaviour and the conditions under which it occurs, using observations in which larval development and physical conflict within parasitised and superparasitised hosts were mapped daily.
• 2 Metaphycus flavus larvae often engaged in physical conflict that resulted in consumption of the losing larvae (= cannibalism ) in superparasitised hosts, whereas such conflict and consumption occurred rarely when a single brood developed in a host.
• 3 Cannibalism among M. flavus larvae only occurred after the host resources had become scarce. Typically it occurred after the sixth day of development (fourth‐instar larvae) when the larvae in a clutch had separated from their aeroscopic plate and were freed of their attachment to the host's cuticle.
• 4 Female larvae in the initial clutch appeared more aggressive than male larvae when a second clutch was allocated 4 h after the first clutch. The probability of a larva being attacked and consumed by a brood mate increased as the number of larvae increased in the host. This partial tolerance might allow the members of the initial brood to defend themselves from offspring of a superparasitising female (= competitors ). Such post‐ovipositional regulation of brood size might be interpreted as high‐density intolerance among female offspring.

     

Studies on the interference among larvae of the citrus leaf miner,Phyllocnistis citrella stainton (lepidoptera: Phyllocnistidae)

Observation of the citrus leaf miner, Phyllocnistis citrellaStainton suggested that contacts among larvae frequently occur and either of two larvae encountered is sometimes killed by the interference. A model to describe the competitive interactions in the process of interference was constracted with this species. The distribution of hatchling on an area was considered in the model. The model was based on that of encounters in randomly moving particles in two dimensions. First the time when first death occurs was calculated and n hatchlings were reduced to n−1. Further calculation was made to obtain the time that next death occurs. Such calculation was repeated until the number of larvae was reduced to 1 or the area of mine attained to the completed size. By the simulation based on the data observed on the citrus leaf miner, the numbers of survivors were calculated at different hatchling densities and on different leaves in size. Based on this simulation analysis it was suggested that the following characteristics observed in the present studies enable the maximul number of individuals to become adults.

1. When two hatchlings occur on a leaf, they tend to be separated by the midrib. This implies that female moths tend to reject to lay eggs on the area which has already contained the egg.
2. The larva sometimes avoids the contact immediately before it occurs.
3. The larval mortality caused by a contact is reduced with the larval growth. Since the frequency of contacts increases as the mine is enlarged, the change to high survival prevents the larva from high mortality caused by the increase of contacts.
4. The larval survival does not decrease in spite of the reduction of the area available for mining.

     

The effect of population density on the reproduction ofTrichogramma japonicum Ashmead (Hymenoptera: Trichogrammatidae)

When a fixed number of the hosts, the eggs of the almond moth were exposed experimentally to various numbers of the parasites, Trichogramma japonicum, the following changes were observed with increasing parasite density:

1. The percentage of parasitism rises and approaches to 100 with gradually diminishing rate.
2. The number of parasite progeny increases and reaches a maximum, then decreases gradually.
3. The number of eggs laid per parasite female decreases gradually.
4. The proportion of hyperparasitized hosts progressively rises. The frequency distribution of parasite eggs in a host is of an intermediate type between random and uniform.
5. The competition among parasite larvae becomes severe. The progressive rise in mortality, the declining percentage of females in progeny and the emergence of stunted adults at the higher densities are observed.

In connection with both the nature of the parasitizing behaviour of adult and that of the competition among larvae, the nature of the density effect on the parasite population was discussed.     

Interspecific interference and the functional response of four species of carnivorous stoneflies

1. A previous study compared the functional responses to their prey and intraspecific interference in mature larvae of Perlodes microcephalus, Isoperla grammatica, Dinocras cephalotes and Perla bipunctata. The present study extends this work by assessing interspecific interference between pairs of these species in equal numbers (one, two or three larvae per species) to provide total predator densities of two, four or six larvae. Baetis larvae as prey were replaced as they were eaten, and their density per predator was varied between 20 and 200 larvae. 2. The number of prey eaten by each competing species increased curvilinearly with prey density, the relationship being well described by a Type II model. Of the two constants in the model, handling time varied considerably between species, mean values being shortest for Perlodes, slightly higher for Isoperla, and much higher for Dinocras and Perla. It was not affected significantly either by predator density or the identity of the competing species. 3. Attack rate also varied between species and decreased with predator density. This decrease was slight for Perlodes, and also for Dinocras and Perla in competition with Isoperla. The decrease in Dinocras and Perla was similar to that for intraspecific interference. 4. The decrease in attack rate was described by a convex curve for Perlodes with the other three species and for Dinocras/Perla with Isoperla, but by a concave curve (negative power function) for Isoperla competing with the other three species, and for both Dinocras and Perla in competition with Perlodes. Prey consumption also decreased with predator density, the severity of competition with different species reflecting that for attack rate. 5. A comparison with previous results for intraspecific interference showed that the latter was dominant for Perlodes in all contests and for Dinocras or Perla competing with Isoperla, whilst interspecific interference dominated for Isoperla in all contests and for Dinocras and Perla competing with Perlodes. Both types of interference were applicable to competition between Dinocras and Perla. Isoperla was the least, and Perlodes the most, aggressive of the four species with Dinocras and Perla intermediate.     

Effect of population density on reproduction of two sympatric dung beetle species,Aphodius haroldianus andA. Elegans (Coleoptera: Scarabaeidae)

The effect of population density on reproduction as well as the oviposition modes of two sympatric dung beetle species, Aphodius haroldianus (a low fecundity/high parental effort species) and A. elegans (a high fecundity/low parental effort species) was studied at a pasture in central Japan from 1982 to 1986.

1. The adult population density of A. haroldianus was high (>40 per dung pat). The density of A. elegans was low (<5 per dung pat). Oviposition of A. haroldianus was suppressed above the density of about 10 adults of conspecifics but not affected by the density of other species. The intra- and interspecific density effects on oviposition were not detected in A. elegans.
2. The number of A. haroldianus adults per pat decreased with dung age, while that of A. elegans increased until sixth day after deposition and then decreased. The number of eggs laid per pat was not different between 1-day-and 3-day-old pats for A. haroldianus. However, more eggs of A. elegans were found in 3-day-old pats than in 1-day-old ones.
3. In the both species, the amplitude of population fluctuations was not remarkable; the maximum/minimum ratio for five years being 2.3 for A. haroldianus and 2.9 for A. elegans. Different density dependent processes were suggested to function for the small fluctuation of population density between the two species, i.e. intraspecific density effect on oviposition for A. haroldianus and contest type competition for food among larvae of A. elegans.

     

Local Competition Between Foraging Relatives: Growth and Survival of Bruchid Beetle Larvae

Kin selection theory states that when resources are limited and all else is equal, individuals will direct competition away from kin. However, when competition between relatives is completely local, as is the case in granivorous insects whose larval stages spend their lives within a single seed, this can reduce or even negate the kin-selected benefits. Instead, an increase in competition may have the same detrimental effects on individuals that forage with kin as those that forage with non-kin. In a factorial experiment we assessed the effects of relatedness and competition over food on the survival and on fitness-related traits of the bruchid beetle Callosobruchus maculatus. Relatedness of competitors did not affect the survival of larvae. Larval survival substantially decreased with increasing larval density, and we found evidence that beetles maturing at a larger size were more adversely affected by competition, resulting in lower survival rates. Furthermore, females showed a reduction in their growth rate with increasing larval density, emerging smaller after the same development time. Males increased their growth rate, emerging earlier but at a similar size when food was more limited. Our results add to the growing number of studies that fail to show a relationship between relatedness and a reduction in competition between relatives in closed systems, and emphasize the importance of the scale at which competition between relatives occurs.     

Exogenous and endogenous factors acting on the spatial distribution of a chrysomelid in extensively managed blueberry fields

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