Contest‐type competition between age classes in scramble‐type Callosobruchus maculatus (Coleoptera: Bruchidae)

We examined the effect of age differences on competition type in individuals of a scramble‐type strain of Callosobruchus maculatus (F.). When oviposition of two individuals on a bean was manipulated to introduce time intervals using two lines with different adult body colors, the frequency of two‐adult emergence decreased with the introduction of sequential oviposition. This result indicates that an age difference between two individuals induces contest competition. The frequency of adult emergence in older individuals decreased, whereas in younger individuals it increased with the introduction of sequential oviposition. Using a dissecting microscope, we observed that bodies of older individuals that died in the bean during the 4‐day oviposition interval were crushed at the pupal stage under the pupal chambers of younger individuals. These results show that an age difference between two larvae in a bean causes contest competition due to one‐sided interference by a younger individual during pupation of an older individual. Based on these experimental results, we discuss the ecological cause of contest competition and the population‐level consequences of identified interactions in scramble‐type C. maculatus.     

Variability of intra- and interspecific competitions of bamboo stump mosquito larvae over small and large spatial scales

We carried out field experiments to examine the variability of interspecific competition of mosquito larvae among microcosms in a bamboo grove (small spatial scale) and between bamboo groves at two sites, with single and multiple mosquito species (large spatial scale). Four types of microcosms that differed in capacity and litter input were set. In the hillside bamboo grove, where multiple species occurred, succession of the predominant species from Aedes albopictus to Tripteroides bambusa was observed in control microcosms from which no mosquito larvae were removed. Weekly removal of competitive species resulted in increased pupation of A. albopictus and adult body weight under both rich and poor resource conditions. In the late period of the experiments, the effect of competitor removal on pupation of A. albopictus was greater in deep containers that never dried than in shallow containers that were dried in the laboratory. The number of eggs showed a slight difference between competitor‐excluded and deep control microcosms. These results indicate that interspecific competition limits pupation of A. albopictus more strongly in deep containers than in shallow and drought‐prone containers.
Compared with the hillside site, the larval density of A. albopictus attained a higher density in the bamboo grove in the plain where no competitive species occurred, due to a higher oviposition rate. Lower rate of pupation and lower adult weight at the plain site than at the hillside site indicated that resource limitation was more severe at the plain site. Populations of A. albopictus at hillside and plain sites appeared to suffer from strong inter‐ and intraspecific competition, respectively. At the hillside site, the intensity of interspecific competition appeared to increase later in the breeding season, with a high larval density of T. bambusa. In contrast, at the plain site, intensity of intraspecific competition appeared to be reduced later in the breeding season with decreasing larval density of A. albopictus.     

Population dynamics ofTelenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas' disease vectors IX. Larval competition and population size regulation under laboratory conditions

A series of experiments were carried out with the endophagous egg parasite Telenomus fariai on its host Triatoma phyllosoma pallidipennis to determine the possible role of intraspecific competition by the parasite progeny in population regulation of the parasite. Eight parasite densities (1, 3, 5, 10, 20, 30, 40, and 50 individuals per vial) were used, and the design of sequentially sacrificed replicates applied. Survivorship curves for each density indicated smaller number of progeny per host at higher densities, and the shapes of the curves suggested a relatively early mortality process.Morris' linear regression technique for determining within-generation density-dependence was used, and the results showed that only larval mortality could be identified as density dependent. The same technique applied within the larval stage proved that only mortality of larvae in their second, third, and fourth day of development were responsible for population regulation. The applicability of the technique, as well as the relevance of the results for natural population, is discussed.     

Contest and scramble competitions in two bruchid species,Callosobruchus analis andC. phaseoli (Coleoptera: Bruchidae)

Larval competition between contest and scramble strategists was investigated using the two bruchid species, C. analis (contest species) and C. phaseoli (scramble species) with two different sized mung beans (large and small beans). In both sized beans, the adult emergences of each species dependen on total density of the initial larval densities of the two species and the ratio of the two densities. The emergence of one species was suppressed by the existence of the other species when the initial larval density per bean of the former species was less than that of the latter one. There were many cases in which both C. analis and C. phaseoli emerged from one bean in large beans, but such cases were quite rare in small beans. C. analis performed interference behavior only at late larval stages, whereas C. phaseoli was superior in exploitative competition all through their larval stages. These, combined with the niche segregation inside a bean, are throught to be the major factors of observed density- and frequency-dependent competition results. Based on the above experimental results, long-term competition results between the contest and scramble species were predicted.     

Intraspecific competition in ant-lion (Macroleon quinquemaculatus) larvae in the field

The number of larvae of the pit-digging ant-lion Macroleon quinquemaculatus at a food-poor site in Tanzania increased between May and October 1983 and then remained at high density until the following rainy season in March. Food availability was low and small larvae were most likely to suffer food shortage. Hunger level varied within and between instars: hunger was greatest after moults and only third-instar larvae weighing more than 100 mg were well fed. A simple model showed that exploitation competition was asymmetric and most likely to affect small larvae, by removing small prey. Exploitation was estimated to reduce food intake by 30–60%. Within-site differences in growth rate and size were associated with differences in larval density. Small larvae were much more likely to move than large larvae; the latter tended to occupy the periphery of the habitat. The reduction in food intake experienced by small larvae was a consequence of interference competition, which was the main factor operating at this site.     

Density and intercohort priority effects on larval <Emphasis Type="Italic">Salamandra salamandra</Emphasis> in temporary pools

Priority effects, i.e., effects of an early cohort on the performance of a later cohort, are generally studied between, and not within, species. The paucity of intraspecific assessments does not reflect a lack of ecological importance, but the technical problem associated with differentiating between conspecific cohorts. Here, we examine priority and density-dependent effects on larval Salamandra salamandra infraimmaculata. Larvae deposited by their mother early in the season have increased risk of desiccation, as rains at the beginning of the season are less frequent and unpredictable. However, breeding later may incur a high cost through conspecific priority effects, including cannibalism and competition. In an outdoor artificial pool experiment, we established densities of 0, 1, 2, 4 or 6 newly born larvae per pool (∼30 l), and 40 days later, added a second cohort of three newly born larvae to each pool. We differentiated between cohorts using natural individual-specific markings. For the early cohort, increasing density decreased survival and size at metamorphosis, and increased time to metamorphosis. For the late cohort, survival was 100% in pools without early-cohort larvae, but ranged between 13 and 33% in the presence of early-cohort larvae. Time to metamorphosis was significantly longer in the presence of low vs high densities of early-cohort larvae. Results suggest that early-cohort larvae are mainly subjected to exploitative competition and cannibalism mediated by food limitation, and that late-cohort larvae are subjected to cannibalism and interference due to size asymmetry between cohorts. The strong priority effects suggest that Salamandra females could increase their fitness by adjusting the number of larvae they deposit in specific pools to avoid cannibalism and intraspecific competition.     

Influence of the larval environment on performance and adult body size of the wood-boring beetle Phoracantha semipunctata

Large body size confers a reproductive advantage to adults of the wood‐boring beetle Phoracantha semipunctata (F.) (Cerambycidae: Cerambycinae: Phoracanthini). Larvae of this species feed subcortically in stressed and dying eucalypt trees and logs. We evaluated the influence of the larval environment on larval performance and adult body size by manipulating the post‐felling age of host logs (from freshly cut to 2‐weeks‐old) and the density of colonizing neonates (low density with minimal competition for resources vs. high density with intense competition). Adult beetles emerged in greater numbers from logs that had been subjected to the aging treatment which reduced bark moisture content and favored colonization by neonates. Survival was greatest in larger logs having lower densities of neonates, but was greatly diminished in all treatments by mortality during pupation. Development time varied from 2 months to more than a year and was shortest in smaller logs having high densities of larvae. The size of adult beetles emerging from a log was not influenced by larval density, but was positively correlated with the age of logs when the neonates colonized, and log size. These findings suggest that the optimal developmental conditions for P. semipunctata larvae, in terms of larval performance and adult body size, are available in large, aged host logs having low densities of larvae. Manipulation of the larval environment in this study resulted in a considerable variation in adult body size, but large individuals were relatively more common in the wild population that was the source of neonates for the experiment. Potential body size may have been constrained by our use of only one host species and a narrow range of log dimensions.     

Genetic variation in pupation height in a population of Drosophila simulans

Preadult mortality of Drosophila simulans in the laboratory is influenced by larval behavior during pupation site choice. The possibility that larvae select a suitable place to pupate may have profound repercussions on their darwinian fitnesses. Twenty-three isofemale lines of a population of D. simulans were analyzed twice for pupation preference, the first time inmediately after being captured, and again after a year of laboratory culture. Pupation height per vial, pupation height on the vial walls, percentage of pupae either in food or on the wall, and egg-to-pupa viability were estimated. A great intrapopulational variation was noticed with an important genetic component for pupation site choice. This suggests a great larval adaptative potential in the population for selecting and using heterogeneous habitats. Time seems not to have modified the population in both the mean values and the phenotypic and genetic variances. Larval viability, however, increased with time. Pupation site choice is discussed on the basis of two independent sets of genes.     

DIRECTIONAL AND STABILIZING DENSITY-DEPENDENT NATURAL SELECTION FOR PUPATION HEIGHT IN DROSOPHILA MELANOGASTER

Six populations of Drosophila melanogaster have been kept at extreme population densities, three high and three low, for 175 generations. Larvae from the high density populations pupate 50%-100% higher than larvae from the low density populations. At high larval test densities there is both a directional and a stabilizing component to selection, with viabilities ranging from 0.14 to 0.992, depending on the choice of pupation site. The directional component is stronger on the populations which have evolved at low densities, while the stabilizing component is stronger on the populations which have evolved at high densities. There is no indication that the evolution of this trait, in response to density, has altered its phenotypic plasticity.     

Responses of Mamestra brassicae (Lepidoptera: Noctuidae) to crowding: interactions with disease resistance,colour phase and growth

This study examines phenotypic plasticity in relation to rearing density in larvae of the moth, Mamestra brassicae. Larval phase, growth rate, weight at moulting and susceptibility to disease were quantified when reared at five densities. Larvae develop more quickly, but attain a smaller size and are more susceptible to disease, when reared at high than at intermediate densities. They also exhibit a higher degree of melanisation than larvae reared at intermediate densities, or singly. A review of the literature suggests that a switch to a rapidly developing dark phase at high densities is a widespread phenomenon within the Lepidoptera. Rapid development at the expense of attaining a large size, and increased melanisation, are interpreted as adaptive responses to reach pupation before food supplies are depleted, as is likely when larval density is high. High susceptibility to viral infection at high density may be a result of physiological stress associated with rapid development, or due to a shift in allocation of resources from resistance to development: larvae that developed quickly were more susceptible to infection. Larvae reared singly appeared to be less fit than larvae reared at intermediate densities: they exhibited many of the characteristics of larvae reared at high density, particularly low weight, a right-hand skew in their weight frequency distribution, and high susceptibility to disease. I hypothesise that expression of resistance may be phenotypically plastic with regard to environment. Contact with other larvae may, up to a point, stimulate both growth and resistance to infection, for the risk of infection will increase with the density of conspecifics.     

Onion maggot feeding and development on heterogeneous sections of the onion bulb

When given a choice of feeding sites in laboratory arenas, Delia antiqua (Meigen) larvae exhibited a 3: 1 bias for internal vs outer sections of onion bulbs and distributed themselves non-randomly among identical preferred onion sections. Larval clustering did not appear to be correlated with increased fitness in the laboratory; larval development was identical across the range of densities from 1 to 50 larvae per onion. Larvae feeding on preferred internal sections developed 14% faster, were 38% heavier upon pupation, and were two times more likely to survive to the adult stage than larvae feeding on outer sections. D. antiqua larval food preference was therefore positively correlated with increased fitness.     

The role of abiotic factors in the pupation of Thaumatotibia leucotreta Meyrick (Lepidoptera: Tortricidae) in the soil

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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.     

Spatial and temporal population differences in male density and condition in the American rubyspot, Hetaerina americana (Insecta: Calopterygidae)

Increased resource availability should favor higher animal density. It may also affect sex ratio, the male condition, and mating competition over access to females, although the direction of these variables is not straightforward to predict. Using a non-experimental approach, we carried out preliminary research using the territorial American rubyspot (Hetaerina americana) by comparing two spatially separated populations and the same population in two different seasons (each comparison with varying population densities). We first compared the sex ratio by counting males (using two categories, territorial and non-territorial) and females; relative foraging time (as an indicator of how much feeding resources each site provides); wing spot size (a sexual ornament), body size and immune melanization response (these two variables were used to assess male condition); and fighting time and contest number (to assess competition). For the seasonal comparison we used a third population in which we only compared male spot size and two indicators of condition and immune response, phenoloxidase (PO) and nitric oxide (NO) activity. The high-density population had higher values of non-territorial males (but similar sex ratio), relative foraging time, contest time and number, wing spot (but similar body size) and melanization response than the low-density population. According to season, at high density, males had higher values of wing spots, PO, and NO. Our results suggest that in a population where animals have more dietary resources, males reach a better condition despite the competition being more intense.     

Wall-making behavior as a proximate mechanism to generate variation in larval competition in Callosobruchus maculatus (Coleoptera: Bruchidae)

Variation from contest to scramble in larval competition types was observed among laboratory lines derived from a geographic strain of Callosobruchus maculatus. In contest competition, only one adult can emerge from a small bean because the successful larva monopolizes resources. In scramble competition, however, multiple adults can emerge from the bean because larvae share resources. To explain the variation in competition types, we used six lines of the geographic strain to test the hypothesis that the larval competition type is determined by the larval behavior of building walls, which prevent larvae from interfering with each other, allowing multiple adults to emerge from a single bean. We also investigated the proportions of wall-making in contest-scramble hybrid lines to test whether the formation of a wall structure was genetically determined. Results support our hypothesis that wall-making behavior determines the type of larval competition within a geographic strain, and that the behavior is genetically determined. Scramble-type lines exhibited higher frequencies of wall-making than contest-type lines when two larvae of the same line infested a bean. Larval competition type and the tendency towards wall formation in contest-scramble hybrid lines ranged intermediate of parental lines. We concluded that the variation in larval competition type is determined by the variation in larval wall-making behavior among laboratory lines derived from the geographic strain. We will discuss the evolution of scramble-type larvae in C. maculatus based on our results.     

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.     

Larval competition causes the difference in male ejaculate expenditure in Callosobruchus maculatus

The seed beetle Callosobruchus maculatus larvae exhibit two types of resource competition: scramble, in which a resource is shared, and contest, in which the resource is monopolized. This difference in larval behavior results in different adult densities. Under contest competition, adult density remains constant regardless of larval density, but under scramble competition, adult density increases with larval density. This in turn affects mating frequency during adulthood, and thus, the intensity of sexual selection operating on males. In this study, we examined the relationship between larval competition types and male reproductive investment in mating. We assessed the male ejaculate expenditure per mating across geographic strains of C. maculatus. The male investment (ejaculate expenditure) increased with the degree of scramble competition and decreased with the degree of contest competition. We therefore suggest that males experience different selective pressures depending on the type of larval competition: scramble type males are selected for increased reproductive investment.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

Contest competition inDrosophila subobscura

The role of larval intraspecific competition in laboratory populations ofDrosophila subobscura was investigated. Mortality is density-independent during the first 3 days after hatching but becomes density dependent as development proceeds to pupation. Although total biomass per patch was independent of initial egg density, competition betweenDrosophila larvae leads to the formation of smaller pupae. This resulted in a population that was dominated by suppressed individuals. Development rate ofD. subobscura larvae was not affected by high larval densities. Smaller pupae give rise to females with fewer eggs in their ovarioles. A simple simulation model, predicting the effects of intraspecific competition on the fecundity of the nextDrosophila generation is described.     

Population regulation of a mosquitoArmigeres theobaldi with description of the animal fauna in zingiberaceous inflorescences

1. Inflorescences of some Curcuma and Zingiber (Zingiberaceae) in tropical Asia provide an unique aquatic habitat being discrete, small, and made of numerous smaller compartments (the bracts).
2. The aquatic community in inflorescences of Curcuma in northern Thailand was composed of immature Diptera, of which the biting midge Dasyhelea and the mosquito Armigeres theobaldi were the commonest. No important competitors, predators or pathogenic parasites for the mosquito were confirmed.
3. Inter-inflorescence distribution of the mosquito was contagious. Within each inflorescence, the fourth-instar larvae or pupae usually occupied bracts singly.
4. The k-value analysis detected density-dependent mortality due to contest competition in the mosquito larvae.
5. Variations in the larval and pupal mosquito size were density-independent and remarkably small as compared with size variations known for other mosquitoes.
6. These population attributes (large density-dependent mortality with density-independent, minimally variable individual size) appear unique among mosquitoes, arising from conspecific killing, efficient foraging (inter-bract movement by crawl and single occupation of bracts), and availability of host plant tissues as supplementary food.
7. A simple population model suggested that a small proportion of adult females lay eggs.