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.     

Damped oscillation of population density at equilibrium

It has been theoretically assumed that the population density at the equilibrium oscillates with damping from generation to generation. In the adult population of the southern cowpea weevil, Callosobruchus maculatus, it was exemplified. But, it was not so clear in the adult population of the azuki bean weevil, C. chinensis as seen in that of C. maculatus. This difference seems to be due to the scramble type of competition that occurs in larval stage in C. maculatus, instead of in the egg stage as C. chinensis. Comparing with the oscillation from generation to generation obtained in the present experiment to that of Lucilia population found byNicholson , the oscillation in Lucilia population is composed of the cycle in a generation and the descending phase of each cycle of it is not regulated density-dependently. The present result seems to be more appropriate for the demonstration of the theory of self-adjustment of population.     

Density dependence depends on scale; at larval resource patch and at whole population

A previous study (Tuda and Shimada, 1993) has shown that the equilibrium population size of the azuki bean beetle was lower at 32°C than at 30°C and that this difference was due to a reduced maximum population size of emerged progeny through inside-bean process. In this paper, these results were analyzed further on the scale of the individual bean where interaction among larvae took place. Per-bean numbers of deposited eggs, hatched eggs, and emerged adults have been recorded at seven different parental densities under the two temperature conditions. Three individual-bean-scale process hypotheses that may explain the reduced maximum emergence density on the whole population scale are suggested: (1) a lower maximum emergence per bean at 32°C than at 30°C, if the bean scale and the wholepopulation scale share the same density-dependent pattern in adult emergence, (2) a limited range of hatched egg number per bean at 32°C, resulting from the adult oviposition process outside beans, and (3) different patterns of density-dependent emergence between the two different scales. This study showed that the inside-bean pattern of responses on the bean scale was a simple saturated curve at 30°C, but one with a discontinuous decline at higher hatched egg densities at 32°C. On the contrary, during outside-bean process, the peak number of hatched eggs decreased on this scale as observed on the wholepopulation scale. I discuss why the extracted factor of inside-bean process on the whole-population in the previous study could not be applied to the bean-scale pattern.     

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae) II. Larval competition and interference mechanisms

Competition between contest and scramble strategists was examined using two strains of Callosobruchus maculatus, the contest strain (iQ) and the scramble strain (tQ). The direct larval interference experiment within each strain showed that the contest strain performed strong direct interference throughout its larval stage against the conspecific opponent(s). The scramble strain also performed its interference, but only during the 3rd and the 4th larval stages. The inter-strain larval competition experiments inside a large and small mung bean (Vigna radiata) showed that the contest strain was, in general, superior to the scramble strain in competition, but the competition results were density- and frequency-dependent. In the large beans, the proportion of adult emergence of the contest strain increased with the increase of its own initial density but decreased as the initial density of the scramble strain increased. The shape of the proportion of adult emergence became nonlinear in the small beans. The cause of the complexity of inter-strain competition results was discussed in the light of the difference in larval interference ability between the two strains.     

Studies on interspecies competition between the azuki bean weevil and the southern cowpea weevil

Experimental studies was made on the interspecies competition between the azuki bean weevil, Callosobruchus chinensis and the southern cowpea weevil, C. macultatus. And the following results were obtained.

1. The reverse result of competition between the two species observed under the air-tight condition and the air-free one. That is, the population of the azuki bean weevils destroyedby that of the southern cowpea weevils under the former condition and vice versa under the latter. It is thought that such a reversal is due to the difference of sensibility of each species to the air-tight condition.
2. Under the air-free condition, change of the time interval of food-supplying had no effect on the result of competition within the limits of this experiment. The population of the southern cowpea weevils was always destroyed by that of the azuki bean weevils. But there was a certain degree of correlation between the duration of co-existence and the interval of food-supplying.
3. From the results, it is possible to say that by changing the degree of air-tightness, or the time interval of food-supplying, the co-existence period and the final result of competition can be changed.
4. The different mechanism of competition between two species in the present experiment from the experiments ofUtida (1952) andYoshida (1957) were discussed.
5. A difference in the mechanism of interspecies competition and intraspecies competition was expected from the level of total population numbers of two species and the individual weight of C. chinensis.

     

Systems analysis of density-dependent population processes in the azuki bean weevil,Callosobruchus chinensis

Systems analysis of density-dependent population processes was conducted in an experimental population of the azuki bean weevil,Callosobruchus chinensis. Density-dependent population change was formulated for two separate processes; adult production of eggs and hatching of the eggs outside the beans, and survival of the first instar larvae inside the beans and their emergence as new adults. The formulation of each process was based on a common basic equation modified from the logistic-difference equation. Parametric values were estimated from experimental data onC. chinensis in a laboratory population. Simulations of the model showed good fits not only to observed population changes both outside and inside the beans, but also to the observed reproduction curve reflecting the population change as a whole within one generation in an experimental population. Furthermore, population dynamics from generation to generation were simulated from the model reproduction curve. The simulation showed a slight dumped oscillation and converged to a carrying capacity, which fitted well with the experimental population dynamics.     

Competitive exclusion between contest and scramble strategists in <Emphasis Type="Italic">Callosobruchus</Emphasis> seed–beetle modeling

Interspecific competition between an intermediate contest strain of Callosobruchus maculatus and a scramble strain of C. chinensis was investigated on two types of resource beans: the azuki (Vigna anguralis, small seed) and the black-eye bean (Vigna unguiculata, large seed). We conducted both single-generation competition experiments and multiple-generation experiments and then analyzed the system with a discrete version of the Lotka–Volterra competition model. The estimated competition coefficients showed the competitive dominance of the C. maculatus strain over the C. chinensis strain in both types of bean. Zero-growth isocline analysis based on the predicted model showed that an overall dominance of the contest species, C. maculatus, in azuki beans, whereas the outcome of competition was dependent on the initial population size of each species in the black-eye beans. In the multiple-generation experiments, C. maculatus was the successful competitor irrespective of the ratio of the two beans, whereas C. chinensis overcame C. maculatus in one of five replicates with the black-eye beans system. When we incorporated the cost of scramble competition, or density-dependent reduction in body mass into the predicted model, the model alteration decreased the survival region of C. chinensis in phase space on the black-eye bean. Thus, the competitive equations, which do not consider the density-dependent body size reduction, tend to overestimate the advantages of scramble strategists. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Cause of egg mortality in azuki bean weevils (Callosobruchus chinensis): physical or chemical?

The cause of high egg mortality at high adult density in azuki bean weevil is still controversial. It may be physical, as a result of adult trampling, or chemical, as a result of substance(s) secreted by adults and coated on the bean surface. In this study, I first constructed simple mathematical models based on each mechanism, which could predict the change of egg hatchability oviposited on different dates at different adult densities. I then devised an experimental method that could identify the fate of eggs oviposited on different dates. The experimental results showed that egg mortality was most likely caused by adult trampling. An erratum to this article can be found at     

Evolution of contest competition and its effect on host–parasitoid dynamics

In experimental populations of the cowpea bean weevil Callosobruchus maculatus (Coleoptera: Bruchidae) and a parasitic wasp Heterospilus prosopidis (Hymenoptera: Braconidae), large changes in the abundances and the fluctuations of both species occurred after approximately 20 generations. In this paper, we examine the hypothesis that this observed change in the dynamics may have been caused by an evolutionary shift in the mode of competition among the bean weevils. A Nicholson-Bailey type model is developed using parameters measured from the experiments. The host larvae can differ in the type of competitive behaviour that they exhibit, which can be either of a contest type or of a scramble type. If a bean contains one or more larvae of the contest type, only one of these will survive and any scramble-type larvae in the bean will be killed. If no contest-type larvae are present within a bean, multiple individuals of the scramble type can emerge from a single bea n. The model assumes many genotypes, differing in the fraction of offspring of the two types. If a high per capita resource availability is maintained, then the scramble type is selected for, but if resources are limited, then the contest type is selected for. The host population at the start of the experiment, taken from a stock culture, was composed mostly of the scramble type. The model is successful in explaining the initial quick increase in the host's abundance, followed by the evolutionary increase in the fraction of the contest type among hosts, resulting in the more stable population dynamics of the host–parasitoid system, as observed in the experiments. However, it predicts a parasitoid abundance much higher than that observed. We discuss alternative hypotheses to explain the observed evolutionary shift in the population dynamics. We also examine the effect of the difference in size of the beans in the stock culture and those used in the experiments.     

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae)

Larval competition curves and resource sharing patterns of 5 strains of Callosobruchus maculatus (iQ, yQ, aaQ, wQ, and tQ) were examined. Offspring emergences as a function of the initial larval density were recorded to construct competition curves. Elytron length of emerged adults was used as the indicator of resource sharing patterns among competing larvae inside a bean. In the large beans, strain iQ showed a saturated competition curve and tQ strain showed a humped curve. Competition curves of the other 3 strains (yQ, aaQ, and wQ) were between those two extremes. In the small beans, strains iQ and tQ also showed a saturated and a humped competition curves, respectively, whereas the competition curves of the 3 intermediate scramble strains could not be distinguished from that of the iQ strain. Thus, the classification based on competition curves was sensitive to the resource condition (bean size). In both the large and the small beans, the elytron lengths of iQ strain remained constant irrespective of initial larval density. On the contrary, the elytron lengths of the 4 other strains decreased monotonically with higher initial larval density. Thus, the judgment based on the resource sharing pattern was shown to be robust. Only iQ strain should be designated as a contest type, and the remaining strains as scaramble types. Contest and scramble types in C. maculatus were also compared with those observed in C. analis and C. phaseoli using competition curves, resource sharing patterns, and other physiological characters.     

Bottom-up derivation of population models for competition involving multiple resources

This paper provides first-principles derivations of population models for competition involving multiple resources with different competition types, based on resource partitioning between individuals. The following two cases are investigated. The first is the case in which the resource competed for and its competition type change depending on life stages from scramble to contest competition, or from contest to scramble competition. The second is the case in which individuals compete for two resources simultaneously with scramble and contest types, respectively. In both cases, population models are derived analytically, and in particular, the Hassell model is derived in the second case. The nature of reproduction curves and the stability properties of three population models derived are compared with each other. These models provide three representative models for competition involving both scramble and contest types.     

The development of SSR markers by a new method in plants and their application to gene flow studies in azuki bean [<Emphasis Type="Italic">Vigna angularis</Emphasis> (Willd.) Ohwi &; Ohashi]

To gain a better understanding of wild and weedy azuki population structures in relation to the cultigens we have developed simple sequence repeat (SSR) markers based on a new methodology for plant material. In the azuki bean genome, the number of (AG)_n and (AC)_n motif loci per haploid genome has been estimated to be 3,500 and 2,100, respectively, indicating that (AG)_n motifs are a rich source of markers. We constructed a (AG)_n-SSR-enriched library in azuki bean in order to obtain a comprehensive range of SSR markers efficiently. The method applied in this study resulted in a 116-fold enrichment over the non-enriched genomic library, with a high percentage (98%) of successful single-locus amplification by the primer pairs designed. Consequently, this method can be applied to construct SSR-enriched libraries suitable for large-scale sequencing. We obtained 255 unique sequences from an (AG)_n-enriched library for azuki bean. Fifty primer pairs were designed and screened against five populations of wild azuki bean. Among these five populations, one population from Bato town, Tochigi prefecture, Japan, showed greater polymorphism using these primers than the others and was therefore chosen for the in-depth study. The genotypes of 20 individuals were investigated using eight of the SSR primers developed. The genetic relationships among individuals revealed a complex spatial pattern of population structure. Although azuki bean is considered to be a predominantly self-pollinating species, 3 of the 20 individuals tested in the population showed heterozygous genotypes, indicating outcrossing. Allele size and DNA sequence in each of the 20 individuals were compared with those of landraces and released cultivars of azuki bean. Plants in part of the population had many alleles of the same size and with the same sequence as those in cultivated azuki bean, suggesting that gene flow from the cultigen to wild plants has occurred in this population. Unintentional transgene escape from azuki could therefore occur when transgenic azuki is grown in areas where its wild and weedy relatives occur. The approach used here could be applied to biosafety monitoring of transgenic azuki bean.Communicated by C. MöllersX.W. Wang and A. Kaga contributed equally to the results presented in this paper.     

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.     

Biological control of indianmeal moth and rice weevil by parasitoids with reference to the intraspecific competition pattern

Biological control of rice weevil, Sitophilus oryzae (L.), and Indianmeal moth, Plodia interpunctella (Hübner), by their parasitoids Anisopteromalus calandrae (Howard) and Bracon hebetor Say was examined while considering the intraspecific competition pattern of the pests. In both experimental and simulation studies, A. calandrae was shown to suppress the rice weevil population, a contest type competitor, regardless of the parasitoid/weevil ratios tested. In contrast, B. hebetor only significantly suppressed the Indianmeal moth, a scramble type competitor, when the parasitoid/moth ratio was >0.05. At ratios lower than 0.05, the role of B. hebetor was negligible, and the correlation coefficients between the number of moths that had emerged and the parasitoid/moth ratio was estimated to be 0.07. The control efficiency of the two parasitoids with respect to the parasitoid/host ratio was estimated using a ratio-response model. To suppress the weevil density to a level that was only 10% of the current density, the ratio was estimated to be 0.02, whereas this value was 0.14 for the Indianmeal moth. However, for the continuous suppression of the Indianmeal moth, periodic and iterative introduction of B. hebetor was required.     

The fitting of the logistic equation to the rate of increase of population density

Though there are many problems on the usefulness of the logistic curve, it may be necessary to examine before discussing these problems whether or not the actual data fit to the theoretical values. It has been clarified in this paper that the relation between the population density and its rate of increase per individual described by the differential equation (1) is represented by a straight line on a finite difference diagram on which N_i+1−N_i/N_i values are plotted against N_i+1. Utilizing this linear relation we may examine the fittness of the logistic curve to the actual data and when it is fitted we may estimate the parameters of the logistic equation by (5) and (6). The result of the application of this method to the experimental populations of azuki bean weevil indicates that the relation between parent and progeny densities fits well to the logistic type as has been proved byFujita andUtida (1953) who utilized the linear reltion between 1/R+2σ and parent density where R is the apparent rate of reproduction and σ is a constant dependent primarily upon the length of adult life (0≦σ≦1).     

Larval competition within seeds: From the behaviour process to the ecological outcome in the seed beetle Callosobruchus maculatus

Abstract Environmental conditions experienced by organisms during development can have profound impacts on adult fitness and behaviour. Internally feeding larvae unable to leave the seed selected by their mother face limitations of resource suitability and competition. The host seed may guide the larval behaviour within the seed leading to differential intensity of competition and determining its process and outcome, which varies in strains of the legume seed beetle Callosobruchus maculatus (Coleoptera: Bruchidae). However, the intensity, process and outcome of larval competition in different hosts have yet to be simultaneously considered, the objective of the present study. Here we assessed the intensity, process and outcome of intrastrain larval competition as related to host type, and how they are interrelated. Larval competition was faced with two distinct strategies – scramble and contest competition depending on the insect strain and host seed species. The intensity of competition did not show any straight link with the process and outcome of competition. Only a single strain showed a contest competition process with likely interference between larvae, while the four other strains studied showed the process of scramble competition. The process of scramble competition, however, led to variable outcomes in mung beans based on larval competition curves. Such differences were not apparent on cowpea seeds and either the plateau or the peak expected on the larval fitness curves were not reached preventing the distinction of the competition outcome, a likely consequence of the egg laying behaviour of these strains limiting the maximum number of eggs laid per seed. Seed host species rather than seed size are the likely cause of the differences observed from the initial expectation. The strain showing the process of contest competition increased larval fitness with density of larvae emerged per seed regardless of the host species, an unexpected outcome based on theoretical models. In this case the egg laying behaviour of the adult female is probably the main fitness determinant of its progeny.     

Sterilizing effect of apholate and metepa on adults of the azuki bean weevil,Callosobruchus chinensis L.

Summary The sterilizing effects of apholate and metepa on adults of the azuki bean weevil have been compared. Statistical analysis of the data shows that the physiological effect of apholate on the hatching and emergence of the offspring of the treated weevil was about two times stronger than that of metepa.

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Zusammenfassung Die sterilisierenden Wirkungen von Apholat und Metepa auf erwachsene Kundekäfer wurden verglichen. Die statistische Analyse der erhaltenen Daten zeigt, daß die physiologische Wirkung von Apholat auf das Schlüpfen und Auftreten der Nachkommen der behandelten Käfer etwa zweimal stärker war als bei Metepa.

     

Studies on the interspecies competition between the azuki bean weevil and the southern cowpea weevil IV. Competition between strains

This study analyzes results of interspecies competition between the azuki bean weevil, Callosobruchus chinensis and the southern cowpea weevil, C. maculatus using the four geographical strains of each species.

1. Fifteen combinations of one strain from each species were set up to compete. The strains of C. chinensis, rather than of C. maculatus, characteristically determine the eventual outcomes of competition. Some strains of C. chinensis (jC and kC) invariably won every encounter with C. maculatus. Another strain (nC) always lost its encounters, while one other strain (iC) won some encounters (:cQ and :cQ) and lost others (:tQ and :bQ). For any combination the competition results were deterministic and no reversal in outcome was observed among the replicates.
2. The weakness of strain nC in competition is explained by its characteristics in single species populations, different greatly from other strains of C. chinensis, although it had the similar primary (individual) characteristics as other strains of C. chinensis. Strain iC had the shortest adult longevity in the four strains of C. chinensis although it behaved similar to strain jC at the single species population level. Especially under the food supply schedule of the present experiment the shortness of adult longevity might be the important factor determining the outcome.
3. The relationship of the eight strains based on the individual and single species population characteristics, and that based on the competition results were very similar. Even when the competition results could not be explained by any one or two of these characteristics alone, the outcome would be largely decided by the overall biotic profile made by several characteristics.

     

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.