Stimuli governing intraspecific egg predation in the flour beetles,Tribolium confusum andT. castaneum

Summary The present research has not identified all of the factors that interact to produce the observed egg cannibalism rates inTribolium. It has, however, shown that some physical and chemical properties of the egg and environment play an important role in determining cannibalistic activity in a population. Of general interest is the observation that cannibalism is not solely the result of the biological characteristics of the cannibal. While it is generally acknowledged that the quality and quantity of food may influence the intensity of cannibalism, it is not generally recognized that the environment can affect cannibalism in as complex a manner as is suggested by the current research. That two such closely related species, and even two life-stages of the same species, may respond so differently to the same environmental cues is impressive. Predicting the intensity and hence the importance of cannibalism in natural populations may be difficult without detailed empirical studies.     

The dynamics of laboratory populations ofCallosobruchus chinensis andC. Maculatus (Coleoptera: Bruchidae) in patchy environments

Summary Experiments are described showing the long-term dynamics of two species of bruchid beetles (Callosobruchus chinensis andC. maculatus) in arenas in which the resource of 50 black-eyed beans is divided between 5, 10 or 50 ‘patches’. Both species of adult beetles exhibit clumped distributions between patches. Within a patch there is a tendency for a density dependent reduction in (1) eggs laid per female, (2) the proportion of eggs hatching per bean (C. chinensis only) and (3) larval survival which is strongly overcompensating (particularly inC. maculatus). A discrete generation model is used as a framework to draw these results together and show how the different factors affecting natality and mortality can influence the population dynamics. Finally, the importance of the resource renewal interval in influencing the period of the population cycles is discussed.     

The effect of larval age range on survival of twoTribolium castaneum strains in mixed cultures, through pupal cannibalism

Summary The effect of larval age range on survival of twoTribolium castaneum strains in mixed cultures was studied in an experimental situation in which other selective forces were reduced to a minimum. Increasing the difference in larval age setween the strains by one day resulted in a reduction of survival of the faster-developing mutant strain, because more of its earlier pupae were cannibalized by the larvae. General aspects of the effects of pupal cannibalism by larvae on the outcome of intra-specific competition (sensu lato) inTribolium are discussed.     

A stochastic computer model for simulating population growth

Summary A model is described for investigating the interactions of age-specific birth and death rates, age distribution and density-governing factors determining the growth form of single-species populations. It employs Monte Carlo techniques to simulate the births and deaths of individuals while density-governing factors are represented by simple algebraic equations relating survival and fecundity to population density. In all respects the model’s behavior agrees with the results of more conventional mathematical approaches, including the logistic model andLotka’s Law, which predicts a relationship betwen age-specific rates, rate of increase and age distribution. Situations involving exponential growth, three different age-independent density functions affecting survival, three affecting fecundity and their nine combinations were tested. The one function meeting the assumptions of the logistic model produced a logistic growth curve embodying the correct values orr _m andK. The others generated sigmoid curves to which arbitrary logistic curves could be fitted with varying success. Because of populational time lags, two of the functions affecting fecundity produced overshoots and damped oscillations during the initial approach to the steady state. The general behavior of age-dependent density functions is briefly explored and a complex example is described that produces population fluctuations by an egg cannibalism mechanism similar to that found in the flour beetleTribolium. The model is free of inherent time lags found in other discrete time models yet these may be easily introduced. Because it manipulates separate individuals, the model may be combined readily with the Monte Carlo simulation models of population genetics to study eco-genetic phenomena.     

Damped oscillation of population density at equilibrium

Summary 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 ofC. maculatus. This difference seems to be due to the scramble type of competition that occurs in larval stage inC. maculatus, instead of in the egg stage asC. chinensis. Comparing with the oscillation from generation to generation obtained in the present experiment to that ofLucilia population found byNicholson, the oscillation inLucilia 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. Contribution from the Entomological Laboratory, Kyoto University, No. 404. Supported by a grant in aid for Scientific Research from the Ministry of Education.     

Cyclicity and synchrony of historical outbreaks of the beech caterpillar,Quadricalcarifera punctatella (Motschulsky) in Japan

Historical records of the incidence of defoliation caused by the beech caterpillar,Quadricalcarifera punctatella (Motschulsky) in northern Japanese prefectures from 1910–1993 were used to characterize the cyclicity and synchrony of outbreaks. Cyclicity and synchrony were quantified using standard Box-Jenkins time series methods as well as spectral analysis and simple Markov models. Statistical analysis of these records indicated the presence of quasi-periodic behavior with 8–11 yr between outbreaks. Outbreaks tended to occur synchronously among different prefectures, though the onset of outbreaks was typically lagged. This study illustrates the use of specific statistical methods for characterizing cyclicity and synchrony from crude records of the presence/absence of outbreaks.     

The formal dynamics of controlled populations and the echo,the boom and the bust

This paper analyzes the pattern of fluctuations of births in an age-structured population whose growth is subject to environmental or economic constraint. It synthesizes the traditional demographic analysis of age-structured renewal with constant vital rates and the economic analysis which treats population change endogenously. When cohort fertility depends on relative cohort size, or when period fertility depends on labor force size, fluctuations of forty or more years replace the traditional "echo" or generation-length cycle. Twentieth-century U. S. fertility change agrees well with the theory, as the "Easterlin Hypothesis" suggests; but the period model fits better than the cohort model.     

Loss of genetic variability in a fragmented continuously distributed population

An individual-based simulation model was used to examine the effect of population subdivision, dispersal distance of offspring, and migration rates between subpopulations on genetic variability(H ₁ H _S andH _T ) in a continuously distributed population. Some difficulties with mathematical models of a continuously distributed population have been pointed out. The individual-based model can avoid these difficulties and can be used to examine genetic variability in a population within which individuals are distributed continuously and in which the dispersal of individuals is disturbed by geographical or artificial barriers. The present simulation showed that the pattern of decrease inH ₁ had three stages. During the first stage,H ₁ decreased at the rates predicted by Wright’s neighborhood size. During the second stage,H ₁ decreased more rapidly when the migration rate decreased, while during the third stage, it decreased less rapidly when the migration rate decreased. Increasing the number of subdivisions increased the rate of decrease after the 200th generation. The pattern of decrease inH _T was classified into 2 stages. During the first stage, the rates of decrease corresponded with those of a randomly mating population. During the second stage, a decrease in the migration rates of the subpopulations slowed the rate of decrease inH _T . A uniform spatial distribution and a reduced total dispersal distance of offspring causedH ₁ H _S , andH _T to decrease more rapidly. Habitat fragmentation in a continuously distributed population usually was detrimental to the genetic variability in the early generations. Other implications of the results for conservation are discussed.     

Persistence of a fig wasp population and evolution of dioecy in figs: a simulation study

The relationship between fig trees and their pollinator wasps is a well-known example of species-specific obligate mutualism. In this article we present a stochastic model of this mutualistic system, referring to data on a dioecious fig (Ficus schwarzii) in Borneo, and examine the conditions for the persistence of a wasp population for a given period. (1) When the average duration of the flowering interval of fig trees is short, even a small fig population can sustain a wasp population successfully. A population whose average period of flowering cycle is half that of another population can sustain a wasp population with a number of trees less than half of the other population. (2) The wasp survival rate (WSR) is higher when (a) the variation of the interval periods of fig flowering is smaller, (b) the fig population size is larger, and (c) figs can prolong their receptivity to wait for a wasp if no wasps are available. (3) WSR is predictable from the average proportion of the fig's receptive phases, in which wasps are available, to their total receptive phases. (4) The persistence period of a wasp population increases exponentially with the number of fig trees. Based on these results we propose a new hypothesis, as a possible scenario, on the evolution of dioecy from monoecy in Ficus. Received: November 13, 1998 / Accepted: July 14, 1999     

Population momentum: A wider definition

Summary

Keyfitz has derived an elegant formula for estimating the ultimate size of an initially stable, growing population that abruptly reduces its fertility to replacement level. Reduction of fertility is achieved by the rather unrealistic device of dividing the original age schedule nffertility rates by the net reproduction rate. Only the inertia of the age distribution is thus accounted for, but not that of the fertility schedule. The key idea of an abrupt imposition of a fixed regimen capable in the long run of generating zero population growth may be retained, but the regimen made more realistic. By elaborating the population setting, such disparate ZPG regimens as reduction of marital fertility by contraception, delayed and/or less universal marriage, raised mortality risks, or permanent net out-migration may be formulated. Convergence of the populaton to stationarity becomes a two-phase process: a primary adjustment period of changing fertility rates followed by a period of age adjustment.

The present paper treats what happens when a fixed ZPG sterilization regimen, defined by a minimum age of sterilization γ and constant continuous risk φ of sterilization among unsterilized wives aged γ to β, is imposed abruptly (or else progressively over an interval T) upon an initially stable, growing population. Additional sources of residual growth are: (1) the nine-month lag in sterilization effect owing to pregnancy: (2) the more youthful pattern of child-bearing under sterilization: (3) the extra adjustment period (of length β-γ-0.75) of changing fertility rates; and (4) any delays in exposing elements of the population to the sterilization regimen.

Two questions are pursued. First, how important are the additional sources of residual growth? Secondly, how do their relative sizes vary as a function of the characteristics of the initial population?     

Modelling and analyzing density-dependent population processes: Comparison between wild and laboratory strains of the bean weevil,Callosobruchus chinensis (L.)

Three models were constructed for analyzing the population characteristics ofC. chinensis on stored beans; model A describing the whole reproductive process with a single equation, model B describing the three age-specific processes (oviposition, egg survival and larval survival) with separate equations, and model C which describes all these processes not for the whole habitat but for the individual beans comprizing it. The logit equation was employed here as a common basis to describe the density-response relationship involved. All three models showed very good fit to the experimental data obtained for both laboratory and wild strains of the weevil. The parameter values characterizing the population dynamics were, however, widely different between the two strains; the laboratory one which had been reared for some 500 generations showed significantly higher reproductive capacity, less sensitive and gentler response to crowding in both adult and egg stages, and more uniform egg distribution among individual beans, as compared with the wild strain newly introduced. Sensitivity analyses using these models suggested that these changes in population characteristics have been attained by the process of domestication or adaptation to stable laboratory conditions through a long period of time. This process seemed in effect to have optimized the population's performances in the laboratory environment. Evolutionary significance of such optimization was discussed with reference to the selection pressure which may have acted upon individuals.     

Toward a general analysis of endogenous Easterlin cycles

Easterlin believed that there were two features associated with the birth cycles he observed: the cycles were related to the labor market, and they might be self-generating. This paper tries to set up a model that contains both of these two features. We suppose that the welfare of various age-specific cohorts are determined by their respective marginal productivity, and that the underlying technology which puts together labor force of various age-specific cohorts can be characterized by a general production function. Under these weak assumptions, we show that the well-analyzed cohort and period models along the lines of Lee (1974) are restricted versions of our general setting. Given that both the cohort model and the period model were rejected by statistical tests, we adopt the coefficient values obtained from the estimation of the unrestricted version to perform the bifurcation analysis. We go beyond the previous study which focused upon the possible existence of limit cycles, and show that the U. S. fertility limit cycle solution is unstable. Therefore the population trajectory will never converge to that limit cycle.We thank Professors Kenneth Wachter, Shripad Tuljapurkar, and two anonymous referees for their valuable help, comments and suggestions.     

Deme formation in gall-making aphidsAdelges japonicus (Homoptera: Adelgidae)

The deme formation hypothesis in herbivorous insects states that herbivores differentiate genetically into small demes, each specialized in the idiosyncratic traits of individual plants. We examined this hypothesis in gall-forming aphids,Adelges japonicus, by reciprocal transfers of aphids among three susceptible clones ofPicea jezoensis standing in proximity. Performance of aphids transferred to their natal clone was compared with that of aphids transferred to other novel clones. ANCOVA showed no significant donor (source of aphids) by receptor (recipient of aphids) interactions for both total aphid performance on the host tree over a life cycle and its three subdivisions(q ₁,q ₂ andq ₃). In addition, the total performance on the natal clone was lowest among those in the three clones, which was explicitly against the prediction from the local adaptation. In contrast, there were significant receptor effects onq ₁ (fecundity of gallicolae multiplied by survivorship of fundatrices while feeding on needles) andq ₃ (fecundity of fundatrices multiplied by survivorship of gallicolae before emergence from the galls), suggesting genetic differences in resistance among clones. The two-fold difference inq ₃ among clones was due to different gall size, which was highly correlated with the number of gallicolae that emerged from the gall. The two-fold difference inq ₁ seemed to be caused by different survivorship of larval fundatrices while feeding on needles.     

Niche modification and stability of competitive systems. II. Persistence of interspecific competitive systems with parasitoid wasps

Summary Effects of niche shift in ecological time scale on the population dynamics of competing species were studied in the experimental populations of two parasitoid wasp species,Anisopteromalus calandrae andHeterospilus prosopidis (both are solitary parasites), on a host, the azuki bean weevil,Callosobruchus chinensis. Four resource conditions were set up with combination of kind of bean (azuki or black eye), and host distribution (uniform or clumped). In each resource condition, four developmental stages of hosts were provided as a resource spectrum for parasitoid wasps. Population dynamics of the two wasp populations were investigated in each resource condition in Multi-Generation Competitive Systems (MGCS), in which fresh hosts of four developmental stages were periodically introduced and were parasitized competitively by the two wasp species. Competitive coexistence of both wasps occurred in the azuki-clumped condition, where the peaks of the resource utilization curves separated in the two species; pupae inA. calandrae and the early or late fourth instar inH. prosopidis, A. calandrae was eliminated in the azuki-uniform condition andH. prosopidis went extinct in two black eye conditions irrespective of host distributions. The degrees of overlap of the resource utilization patterns of the two wasp species during population dynamics were not significantly different among resource conditions irrespective of the results of coexistence or extinction. Even in the azuki-clumped condition, however, extinction ofA. calandrae was observed when resource partitioning could not be realized with only the late fourth instar larvae available to wasps. Further analytical experiments showed that parasitizing ability ofA. calandrae increased with host density per bean with azuki beans, butA. calandrae could express higher parasitizing ability with black eye beans thanH. prosopidis irrespective of host density per bean. The flexibility in parasitizing ability byA. calandrae for various host stages under different resource conditions was thought to be the major factor in determining the competitive coexistence or the extinction of either species under different resource conditions. The present experiments also suggested that different second-best host stages between competitors could be a major contributing factor to competitive coexistence.     

The outbreak mechanisms of the green rice leafhopper,Nephotettix cincticeps Uhler, in northern Japan

Intraspecific regulatory processes keep the population ofNephotettix cincticeps stable at a low density in southern Japan. In northern Japan, however, the yearly population density of the insect fluctuates violently, and large outbreaks occasionally take place. To clarify the difference in the population dynamics between the two regions, we analyzed light-trap and sweep-net sampling records from prefectural and national agricultural experimental stations. The survival rate of the overwintering population decreased with increases in the period of continuous snow cover (PCSC) in the north, and initial population densities in the years of long PCSC were too low for populations to reach equilibrium density by the end of the active breeding season. This made yearly population fluctuations in the north much larger than in the south. The equilibrium density in the north was higher than in the south. The higher equilibrium density presumably permits the higher population density and larger yearly population fluctuations in the north. A major factor responsible for the difference in equilibrium densities between the two regions is the difference in heading dates of the host plant (rice). Qualitative differences among rice plant varieties, and among biotypes ofN. cincticeps, may also be important.     

Optimal Treatment Schedule in Insect Pest Control in Viticulture

A model for the control of the European grapevine moth Lobesia botrana includes two control methods: insecticides and mating disruption. It yields the combination and schedule of application that minimize cost and losses due to the pest. A simulation is presented for an experimental situation.     

Comparison of dispersal ability between the green leafhoppers,Nephotettix virescens andN. cincticeps (hemiptera: Cicadellidae) by tethered flight

Summary Dispersal propensity of green leafhoppers was compared between a tropical species,Nephotettix virescens, and a temperate species,N. cincticeps. The flight ability was measured with tethered flight technique under laboratory conditions (25°C, 16L-8D). The pre-flight period was shorter and the flight duration was longer inN. virescens than inN. cincticeps in both sexes. No significant correlations were found between the flight activity and morphometric characters for either of the two species. The results suggested that dispersal propensity ofN. virescens is higher than that ofN. cincticeps.     

Crucial factors determining the spatio-temporal distribution patterns of threeAthalia sawflies feeding on common cruciferous plants

Summary ThreeAthalia sawflies,A. japonica, A. rosae andA. infumata, feeding on cruciferous plants, coexist in Japan. However, it is not known what ecological strategies they use and what environmental factors are crucial to such strategies. I attempted to explain these questions by examining the relationship between the spatio-temporal distribution patterns of threeAthalia sawflies and their habitats in three districts (Lowland, Intermediate and Mountain). The three sawflies have different spatio-temporal distribution patterns, though they usually used common cruciferous plants.A. japonica was abundant in spring and autumn but disappeared during summer in all the districts. In the Lowland, populations ofA. rosae andA. infumata, like that ofA. japonica, crashed in summer. HoweverA. rosae occurred mainly in summer in the Intermediate and Mountain. AlthoughA. infumata occurred in the same seasons asA. rosae in all districts, population levels ofA. infumata were always lower than those ofA. rosae. The crucial factors controlling their population patterns were the availability of host plants and temperature. Population crashes ofA. rosae andA. infumata were due to food depletion, and those ofA. japonica were due to heat stress. On the other hand, their population patterns may be interpreted as phenological synchronization with their original host plants, though they all existed on common cruciferous plants. The three sawflies may have evolved different strategies to escape from unfavorable habitat conditions. Such strategies are speculated to be summer diapause inA. japonica, long distance migration inA. rosae, and local dispersal inA. infumata.     

The spatial distribution pattern of the brown planthopperNilaparvata lugens St?l (Homoptera: Delphacidae) in west Java, Indonesia

Summary Spatial distribution pattern of the brown planthopper (BPH) was analyzed at 9 experimental fields in the northern part of West Java during two consecutive rice cropping seasons, i.e., wet and dry seasons. The population of each developmental stage and wing form of BPH at each location showed consistent departure from the random (Poisson) distribution, the variances of the densities in most cases exceeding their means. Namely, the distribution pattern of BPH per hill of rice plant was found to have a general tendency to be aggregated or contagious and to fit fairly well to the negative binomial model. The tendency for aggregation was further confirmed by both the β-values of -m regression being larger than unity and theC _A -values being larger than zero for each developmental stage. Although significant variations in the distribution pattern as measured by β- orC _A -value were observed between different developmental stages, between wing forms and among locations, the degree of aggregation for a given developmental stage at each experimental field remained fairly stable throughout the crop period, despite wide temporal changes in population density. Possible factors to explain these characteristics of the spatial distribution pattern of the BPH in West Java were discussed with reference to the process generating it.     

Comparative studies on the seasonal occurrence, nymphal development and food menu in two giant water bugs,Diplonychus japonicus vuillefroy andDiplonychus major esaki (Hemiptera: Belostomatidae)

Summary We compared the seasonal changes in population density and nymphal development at different water temperatures and under different food conditions between two giant water bugs,Diplonychus japonicus andD. major, in Okayama, Japan. D. japonicus produced 1–2 generations a year, whileD. major was strictly inivoltine. The developmental velocity was higher inD. japonicus than inD. major. The thermal constant ofD. japonicus was less than that ofD. major. These results suggest thatD. japonicus is adapted, to higher water temperature thanD. major. In the field,D. japonicus preyed predominately on Lymnaeidae and Physidae snails, whileD. major preyed on aquatic insects such as dragonfly nymphs.