Researches on Population Ecology

The theories of the stochastic processes are applied to construct mathematical models for describing the processes of population change as an ever changing the distribution of individuals in a space. These models consist of two mathematical expressions which are named the spatial distribution probability function (Q n (t)) and the transition probability function (P i,n (t)), respectively. The former gives the spatial distribution at any future time. Given an actual spatial distribution at any time, the latter function converts it to the spatial distribution at any future time. According to these models, we discussed the time sequence of the mean crowding-mean density relation (Iwao andKuno, 1971) in some population processes such as mortality, birth, immigration, growth, and their combined processes.

Transformation is required to achieve homo-scedasticity when we perform ANOVA to test the effect of factors on population abundance. The effectiveness of transformations decreases when the data contain zeros. Especially, the logarithmic transformation or the Box–Cox transformation is not applicable in such a case. For the logarithmic transformation, 1 is traditionally added to avoid such problems. However, there is no concrete foundation as to why 1 is added rather than other constants, such as 0.5 or 2, although the result of ANOVA is much influenced by the added constant. In this paper, I suggest that 0.5 is preferable to 1 as an added constant, because a discrete distribution defined in {0, 1, 2, . . .} is approximately described by a corresponding continuous distribution defined in (0, ≧) if we add 0.5. Numerical investigation confirms this prediction.

Menida scotti (Puton) males have been shown to transfer secretions from their bulbus ejaculatorius and reservoir of ectodermal accessory gland to females by mating during hibernation. In the present study, the major components of the secretions were found to be proteins and lipids. To specify the female organ incorporating the male secretions, a radiotracer experiment in which the male secretions were labeled by [14C]valine was conducted in nine tissues of females collected in the fall and spring of the hibernation period. Relatively high radioactivities were detected in the haemolymph and the residual carcass (head, legs, air-sacs, exoskeleton, etc.) in the fall females, and in CO2 gas evolved and carcasses in the spring females. The radioactivities in the fat body were significantly higher in the fall mating females than in the spring mating females, and vice versa in the ovary. The radioactivities in six fractions (lipids, proteins, glycogen, sugars, free amino acids and the residues) were also assayed in the five organs of females that had a relatively high radioactivity. The highest radioactivity was detected in the protein fraction of the haemolymph in fall and spring females. There were significant differences in the radioactivities incorporated into the lipid fractions of the carcass between fall and spring females.

The advent of stable nitrogen isotope analysis in ecological research has at last enabled precise identification of trophic position and omnivory due to the differential enrichment of15N over14N with progressive assimilation up the foodweb. I compiled literature data on δ15N values in freshwater and marine foodwebs to search for qeneral patterns in omnivory, specifically the supposition originally proposed by Lindeman (1942) and most recently advanced by Peters (1977), that omnivory should increase with trophic height or position. Omnivory, measured as average intraspecific variability in δ15N, was indeed found to increase with trophic height, such that species at the top of foodwebs were comprised of animals relying, on average, on energy originating from a mixture of different trophic categories.

The dominance of the Pyroglyphidae in house dust has been influenced by a number of factors. The humidity in houses dropped recently to such an extent that only Pyroglyphidae can survive. When temperature and relative humidity are favorable forAcarus siro andGlycyphagus destructor as well asDermatophagoides pteronyssinus, Dermatophagoides diminished in numbers by competition. Introduction of the predatorCheyletus eruditus into this mixed breeds causes the extermination ofDermatophagoides. The vacuum cleaner trapped mites, butDermatophagoides was protected by its habit to hide in cracks and crevices. The use of insecticides has a more killing effect onAcarus andGlycyphagus than onDermatophagoides.

The number of potential annual generations of the rusty grain beetle,Cryptolestes ferrugineus, was simulated in wheat stored in granaries for all crop districts in the prairie provinces of Canada each year from 1952 to 1990 using a population dynamic model driven by ecological variables. Granary size was assumed to be 6 m in diameter. Historical data for temperatures at harvest and times when storage began were used in the simulation model. A second model, which predicted the rate of temperature change at the centre of a 6-m-diameter bulk of wheat, determined environmental parameters for the population dynamic model. (Grain moisture content was assumed constant at 14.5% wet mass basis.) The combined model shows that the initial storage temperature is the most important factor responsible for predicting the number of generations and levels of infestation ofC. ferrugineus. This finding was largely validated by historical grain storage and infestation data. For various years initial grain temperature ranged from 17.7 to 37.4 °C and harvest dates were between 1 August and 20 October. The number of generations annually in simulations based on field conditions ranged from 0.35 to 6.77 with a mean of 3.29. Three or more generations result in a severe infestation and every year at least three simulated generations were completed in some crop districts. In one year, at least three generations were completed in every crop district. Harvest temperature and date permit prediction of crop districts that will potentially have the largest populations of C.ferrugineus so that early monitoring of wheat for infestations can be targeted to areas most at risk.

Colony defense in some aphids is performed by sterile soldiers but in others by monomorphic larvae of a specific instar stage. This paper, focusing on a galling aphidHemipodaphis persimilis with monomorphic defensive first instars, examined the mechanism by which the proportion of defenders is regulated in the colonies. Demographic analyses showed that the ratios of first instars (defenders) were kept constantly high (58% on average) from mid June to late September. High proportions of first instars could be explained by consistently high birth rates (birth rate hypothesis) or by a prolonged duration of the first instar stage (instar span hypothesis). With the progress of colony age, the mature-embryo content of apterous adults, used as an index of the birth rate, decreased and the proportions of advanced instars increased. These results did not support the birth rate hypothesis. By contrast, calculation of a newly proposed index, the molting rate, showed that the duration of the first-instar stage was short in incipient galls but became longer with colony age. The duration of other instar stages was always kept short. These results corroborate the instar span hypothesis and suggest that the prolongation of the first-instar stage is an adaptive mechanism by which the defender ratio is kept high in mature colonies where the birth rate is declining. The frequency of aggressive behavior in first instars increased from incipient to mature galls. Seasonal changes in the instar span and aggressiveness of first instars suggest that inH. persimilis colonies there is a strategic shift from the reproductive to defensive phase with colony age.

In geological history, rapid speciation, called adaptive radiation, has occurred repeatedly. The origins of such newly developing taxa often evolved from the symbiosis of different species. Mutualistic symbioses are generally considered to evolve from parasitic relationships. As well as the previous model of host population with discrete generations, a differential equation model of host population with overlapping generations shows that vertical transmission, defined as the direct transfer of infection from a parent host to its progeny, is an important factor which can stimulate reduction of parasite virulence. Evolution of the vertical transmission rate from both points of view, the parasite and the host, is analyzed. There is a critical level of the rate, below which an evolutionary conflict arises (the parasite would want an increase in the rate while the host would not), and above which both species would correspond to increase the rate. Therefore, once the parasite dominates the evolutionary race so as to overcome this critical level, one-way evolution begins toward a highly mutualistic relationship with a high vertical transmission rate, possibly creating a new organism through symbiosis with perfect vertical transmission. Changes in other parameters may decrease the critical level, initiating one-way evolution. However, changes in traits, probably developed through a long interrelationship in parasitism, do not necessarily induce the evolution of mutualism. Establishment of the ability to make use of metabolic and digestive wastes from the partner certainly facilitates the evolution of mutualism, while improvements in reproductive efficiency of parasites and reduction of negative effects from exploitation in hosts on the contrary disturb mutualism.

A mathematical model is constructed to explain a density-dependent increase in the progeny sex ratios of gregarious parasitoids. In the model we considered non-cooperative game between females concerned with their own inclusive fitness. Equilibrium progeny sex ratios of the first and second females ovipositing on the same host are expressed in terms of the probability of double parasitism (p), the ratio of a male to a female in contribution to resource competition (α), the clutch size ratio between the two females (β), the crowding effect on female reproductive success (γ), and the inbreeding coefficient (f). Major predictions from the model are: 1) the progeny sex ratios of both the first and second females increase withp, 2) as β becomes smaller, the progeny sex ratios of the first females decrease, while those of the second females dramatically increases, 3) when a host is attacked by at most two wasps, the sex ratio of the total number of eggs laid on the host does not exceed 0.25. The effects of α and preferential death by female progeny in doubly parasitized hosts are considered as factors responsible for an excess number of males at emergence. Some possible modes of density-dependent increase in the sex ratios of the overall progeny populations is also discussed on the basis of the present model.

A simple differential equation model was developed to describe the competitive interaction that may occur between species through reproductive interference. The model has the form comparable to Volterra's competition equations, and the graphical analysis of the outcome of the two-species interaction based on its zero-growth isoclines proved that: (1) The possible outcome in this model, as in usual models of resource competition, is either stable coexistence of both species or gradual exclusion of one species by the other, depending critically upon the values of the activity overlapping coefficientc ij ; (2) but, for the samec ij -values, competitive exclusion is much more ready to occur here than in resource competition; (3) and moreover, the final result of the competition is always dependent on the initial-condition due to its non-linear isoclines, i.e., even under the parameter condition that generally allows both species to coexist, an extreme bias in intial density to one species can readily cause subsequent complete exclusion of its counterparts. Thus, it may follow that the reproductive interference is likely to be working in nature as an efficient mechanism to bring about habitat partitioning in either time or space between some closely related species in insect communities, even though they inhabit heterogeneous habitats where resource competition rarely occurs so that they could otherwise attain steady coexistence.

Wolbachia bacteria are obligatory intracellular parasites of arthropods and have been detected in about 70 species of parasitic wasps and three parasitoid flies. Wolbachia are transmitted cytoplasmically (maternally) and modify host reproduction in different ways to enhance their own transmission: parthenogenesis induction (PI), cytoplasmic incompatibility (CI), or feminization (F) of genetic males. Only PI and CI are known in parasitoids. PI-Wolbachia cause thelytoky in otherwise arrhenotokous parasitoids by generating diploid (rather than haploid) unfertilized wasp eggs. CI-Wolbachia cause incompatibility of crosses between infected males and uninfected females because the paternally derived chromosomes fail to decondense and are destroyed after syngamy. More complex situations arise when hosts harbor multiple infections, which can lead to bidirectional incompatibility and may be involved in parasitoid speciation. The relative fitness of infected and uninfected hosts is important to the population dynamics of Wolbachia, and more data are needed. Evolutionary conflict should be common between host genes, Wolbachia genes, and other "selfish" genetic elements. Wolbachia-specific PCR primers are now available for several genes with different rates of evolution. These primers will permit rapid screening in future studies of spatial and temporal patterns of single and multiple infection. Molecular phylogenies show that CI- and PI-Wolbachia do not form discrete clades. In combination with experimental transfection data, this result suggests that host reproductive alterations depend on the interaction between attributes of both Wolbachia and host. Moreover, Wolbachia isolates from closely related hosts do not usually cluster together, and phylogenies suggest that Wolbachia may have radiated after their arthropod hosts. Both results support considerable horizontal transmission of Wolbachia between host species over evolutionary time. Natural horizontal transmisson between parasitoids and their hosts, or with entomoparasitic nematodes or ectoparasitic mites, remains a tantalizing but equivocal possibility.

vigintioctomaculata-complex (Hv-complex), a series of closely related phytophagous ladybirds, has attracted much attention in Japan to the process of speeiation and comparative demography because of its remarkable variations in both external morphology and biology (KATAKURA, 1981; NAKAMU~, 1983). The complex includes a wide variety of groups from pest species (E. vigintioctomaculata, or Evm) to non-pest species feeding on wild plants such as thistles (E. pustulosa and E. niponica) and blue cohosh (E. ya~eutomii). The large 28-spotted lady beetle, E. vigintioctomaculata, a serious pest of solanaceous crops like potato and eggplant in the northern part of Japan. Many studies have been conducted on the ecology of Evm: TAKAHASHI (1932), IWAO (1954), YASUE (1963) and SIMBO (1978) on geographical distribution; KATAKUI~a~ (1976, 1982) on phenology; IWATA (1953) and IWAO (1956, 1968) on spatial distribution pattern on food plants; IWATA (1955) and IWAO et al. (1963) on adult dispersal. Review articles are available on food preference (K_nTAKURA et al., 1977; also see Hosnlga~wA, 1983a, b) and reproductive isolation mechanisms (KAT~U~ and NA~NO, 1986). However, no detailed study has been done on the population dynamics of Evm except IwAo's excellent field research in 1971. In 1981 and 1982, a field population of Evm in Kanazawa, Japan was studied by marking-recapture of adults and construction of a life table. NAKAMURA (1983) summarized the results of this study, comparing the demographic traits of Evm with those of closely related non-pest species and other pest epilachnine species of E. vigintioctopunctata. Without repeating the general discussion previously presented, this article documents in detail adult population parameters such as daily survival rate, longevity, sex ratio and multiplication rate per generation and constructs a life table.

The effects on productivity by the addition of various numbers of radiated adult males or females to populations of two densities were determined for two species of flour beetles. Modifications in productivity after exposure to gamma radiation from60Co included: 1) females ofT. confusum were sterilized by an exposure between 2620 to 5230 R. 2) females ofT. castaneum were not sterilized by 10, 200 R. 3) males were not sterilized by 10, 200 R. 4) sterile females may serve as depositories for normal sperm thereby reducing the productivity. Addition of such females to populations of flour beetles did not reduce the numbers of progeny. 5) since sterile females did not suppress population development, and since they destroy grain and cereal products, they should be eliminated from control programs featuring the sterile insect release technique. 6) population density decreased the numbers of progeny. The use of the sterile insect release technique to control insects of stored grain products does not seem likely with present day sanitary standards since large numbers of insects need be added to effect control.

This study examines the role of learning and memory in the butterflyPieris rapae crucivora Boisduval during foraging for flowers. In an outdoor cage with 6 flower species,P. rapae showed various visiting patterns: some visited only one species, while others visited several species in a day. The foraging process for flowers ofErigeron annuus (L.) Pers. could be divided into two successive steps: (1) landing on the nectaring caputs, and (2) finding the source of nectar in the caput. Butterflies learned to proceed through the two steps more efficiently with successive attempts: they gradually decreased landings on nectarless caputs and probings on the nectarless petals of ligulate flowers respectively. As a result, handling time per unit caputs became shorter, and apparent rewards per unit time, i.e. the efficiency of collecting nectar, increased. In addition, once learned,P. rapae could remember a rewarding flower color for 3 days, which was not interfered with by learning another flower color. This indicates thatP. rapae keeps memory for a period longer than 3 days, and that they can remember at least two flower species as suitable flower resources. Furthermore, data indicated that they sometimes can apply the foraging skills obtained on other flower species to a novel one. These abilities could enable butterflies to easily switch flower species, or to enhance labile preference. It has been known thatP. rapae also shows flower constancy, which may be due to memory constraints. Therefore, they may appropriately use two foraging tactics: visit consistency and labile preference, to get enough nectar according to their circumstances.

In Lepidoptera, females that produce only female progeny have been found in wild populations of at least 12 species. In some species, recoveries, where abnormal females return to normal females, have been observed. A mathematical model of the population dynamics with recovery was developed to identify the conditions for realizing the persistence of abnormal females. Analysis indicated normal and abnormal females coexist and reach an equilibrium state at certain recovery rate values. The equilibrium values of normal and abnormal females were determined. When a population was in equilibrium it was shown that the ratio of normal to abnormal females and the sex ratio after reproduction are always functions of the recovery rate and the proportion of female offspring from an abnormal female to that from a normal female. Using the simulation it was found that, even when a population fluctuates under variable environmental conditions, the two ratios mentioned above reach equilibrium. Equilibrium relationships were applied to published data, and it was concluded that recovery from abnormal to normal females explains the persistence of abnormal females in some species of Lepidoptera. The model developed in this paper can also be used for analysing the persistence of abnormal females of other insect species.

1. Almost all the models so far presented assume that predators are omniscient in the sense that they always have complete information about the spatial distribution of prey abundance and its change over time. But this type of model cannot cover the situation where the prey abundance in each patch changes over time due to factors other than predation. The model with a data window and absolute criterion (SAC) here enables us to treat such situations. 2. The strategy of non-omniscient predators can be generally devided into four procedures; collection of information, its memorization, decision of tactics and its execution. SAC involves only two tactics; to stay another time period in the patch the predator is staying presently or to move to another patch chosen at random. The choice of either one of the two tactics is made by comparing the profitability of the current patch estimated by the data window with a pre-determined absolute criterion. 3. Three changing patterns of prey abundance are considered. In the most general pattern good patches have a higher mean profitability than poor patches, but the profitability changes cyclically in each of patches. 4. There are only two possibilities for an optimal strategy; the “patch choice strategy” in which once the predator has taken a good patch, it tries to stay there even when the state becomes poor, and the ‘state choice strategy” in which the predator seeks for only good states in good patches. The condition for which either of the two foraging strategies is superior to the other is specified analytically.

The attraction range of a human bait forAedes albopictus (Diptera, Culicidae) adults and its absolute density in the range were estimated by a new removal method which is applicable to populations with immigrants. In this model, the number of mosquitoes removed during thekth collection unity k, is expressed as yk = (x0 - \fracba)(1 - s)sk - 1 + by_k = (x_0 - \frac{b}{a})(1 - s)s^{k - 1} + b wherex 0 is the initial number of mosquitoes in the attraction range,a is an index directly proportional to collection efficiency,b is the number of immgirants per collection unit ands(=e−a) is the survival rate of mosquitoes in the attraction range per collection unit. From the result of successive two collections with various interval distances, it was considered thatAedes albopictus adults are attracted to a human bait from the distance of 4–5 m in the bamboo forest on the calm day. The applicability of this method for the population census of forest mosquitoes was discussed.

A mathematical model is proposed to describe the relationship between the abundance and the rank of species in order from the most abundant to the least in a community in an open habitat. This model is derived as a corollary of a species-area equation (Kobayashi, 1975) which could be expected in the case where the individuals of each species are uniformly distributed over a habitat area. Numerical simulation reveals that a rank-abundance curve for a universe results in different species-area or species-individual curves according to the spatial distribution of individuals, and that the relative abundance of each species in a sample varies with sample size unless the spatial distribution of individuals is uniform. A species-individual curve obtained bySanders’s (1968) rarefaction method agrees with that observed actually only for the spatially uniform distribution. Change in the pattern of rank-abundance curve with species diversity and with sample size is discussed in relation to the present model.

1. Comparison of relative abundance of each two successive life history stages was made for studying population dynamics of ayu,Plecoglossus altivelis, in Lake Biwa. 2. Number of eggs spawned is proportional primarily to the size of biomass of spawner population although spawning success plainly depends on presence or absence of heavy rain in the spawning period. 3. Although density of fry tends to be proportional to that of egg, yearly fluctuation of the fry mortality may be caused by changes in density of food organisms but is not in compensatory way. 4. From the fry stage to the juvenile one the rate of mortality and/or of growth seem to fluctuate in compensatory way which is supposed to be related to a shortage of food. 5. Yearly fluctuation of mortality of adult stage is caused by an artificial factor as changes of fishing intensity. 6. Necessity of studying actual events at each stage of the life history process separately in relation with environmental conditions is discussed through the result of the present analysis.

Relative abundance of natural food resource, leaves of wild rose, ofArge migrinodosa was evaluated by comparing the food quantity which the larvae composing a colony utilized throughout the larval stage with that which is available in the habitat. In this connection, behavioral pattern of larval feeding was also observed under natural conditions, and its ecological significance was discussed. It was quantitatively clarified that the larvae faced the severe shortage of food and had to disperse to other shoots with the expense of probably high risk. Under pressures of such insufficient supply of food, the larvae follow the most efficient pattern of feeding behavior that they move to the tip of shoot at first and feed regularly downward thereafter. Though individual larvae behaved completely independently of each other, the aggregation appeared to be formed and maintained because all the larvae from the same egg mass follow this fixed feeding pattern.

Field populations of blue alfalfa aphid and pea aphid on alfalfa were sampled during 1985 and 1986 to determine the association of co-occurrence, interspecific interactions and comparative temporal variations in the spatial dispersion patterns of these species in Oklahoma. Relative abundance of these species is discussed in the light of above analyses.Cole's coefficient revealed a high degree of association between these species in terms of their occurrence on the same alfalfa stems in the field. Regression analyses indicated that the species populations tended to increase in concert on the same stems without evidence of competitive displacement. Spatial dispersion patterns of both species were highly aggregated at low population densities early in the season. Over time, both species tended to disperse and became less aggregated as numbers increased. It was concluded that magnitude of interspecific interactions between the blue alfalfa aphid and the pea aphid were not of a nature that they could be termed as competing species. On the contrary, a concept of an “ecospecies” is proposed for practical applications such as sampling plans and economic threshold determinations.

1. This report demonstrates the well-defined bathymetric distribution of relative abundance of groundfishes in relation to the distribution pattern conjectured from the records of catch along a row of setline with remarkably large variation in the settled depth (from shallower than 350 m to deeper than 700 m) observed around 580 18′ N and 1750 28′ W on Aug. 24, 1962. 2. The results of the analyses are depicted in Fig. 1. It is found that halibut, true cod andSebastodes sp. (or Menuke in Japanese) show sharp or abrupt decrease in the relative abundance in accordance with depth whereas sablefish and rattail increase sharply. Arrow toothed halibut and ray do not show any clear bathymetric change in relative abundance and Pacific ocean perch shows decrease in the density in accordance with the disparity in the depth from the 500 m isobath. 3. The patterns conjectured from this example are compared with either those shown in other reports, the results of the observations on other examples or general view of our fishermen. And many proofs to the high reliability and wide universality of the conjectured pattern of the first four species are found out. But, for the last three species, the present results differ more or less from the general view of our fishermen and the observations on other examples. 4. For the distribution pattern, most of groundfishes incline to show chance distribution in the shallower part of the continental slope (350 m to 700 m deep) and only the species showing sharp decrease in density with increase in the depth are distributed contagiously on the shallower bottom (not deeper than 450 m).

The mating system of the winter cherry bug,Acanthocoris sordidus, was analyzed precisely. As a result, it was found that male adults of this species establish a small territory for mating on the stem of host plant which harbors females. These males abandoned their territories soon after the disappearance of monopolized females. Thus it was confirned that the cue for the territorial establishment of males is the presence of females per se on the host plant. Moreover, most aggregations of adults observed on the host plant contained only a single male. This one-male unit in the mating was named a harem. Harem holding males were usually big in body-size and had a high chance of copulations. The defence behaviors of harem holding males, the mating disparity among males, and the oviposition habit of females in relation to the mating system, were observed. The results obtained were discussed in relation to the sexual selection theories.

The winter cherry bug,Acanthocoris sordidus Thunberg, lives in aggregation especially in their early larval instars. Using the 1st-instar larvae of this species, the author tried to clarify both the processes and the mechanisms of the breakup and later re-formation of colony in relation to the defence against their enemies. The results obtained were summarized as follows.(1) In the field population, there is a high possibility of dispersal of the 1st-instar larvae from a colony possibly through the disturbance by some predators but they can re-form a colony with each other or join, with colonies of different instar larvae. (2) The individuals in a colony immediately disperse through the attack of predatory coccinellid beetle,Harmonica axyridis but tend to re-form a colony in a short time. (3) The breakup of colony is caused by the secretion from the attacked individual. (4) The formation of colony is attributed to the habit closely related with the senses of smell and/or contact. From these results, it was concluded that the dispersal of 1st-instar larvae from a colony, followed by the re-formation of a colony, is an an adaptive behaviour to escape from the attack by their predators.

The mating system of a subsocial spider mite,Schizotetranychus miscanthi Saitō, which is closely related toSchizotetranychus longus Saitō (the long seta form ofSchizotetranychus celarius (Banks) is a synonym of the latter) was studied in comparison with that ofS. longus. Comparisons between nesting patterns of the two related species,S. miscanthi andS. longus revealed a difference in distribution of males among nests. Although more than one male sometimes occurred in the large nests ofS. miscanthi, most nests were occupied only by a single male. On the other hand, many nests ofS. longus included several males. Behavioral experiments revealed that the male and females ofS. miscanthi which cohabited in a nest defended their offspring from phytoseiid predators. Observations and a census of the nesting pattern in a wild population indicate that this is the second example of biparental defense and of a subsocial life-pattern in spider mites. Differences in mating systems were experimentally demonstrated in the two species. Only a single male ofS. miscanthi survived in a nest, as a result of highly aggressive male-male combat, while two males ofS. longus cohabited in a nest. The mating system of the former species is thus considered as harem polygyny, while that of the latter as scramble type polygyny. Furthermore, observations by video recording and scanning electron microscopy showed that the winning male in the male-male combat inS. miscanthi often preyed on the loser, suggesting cannibalism among them.

Experimental observations on the arrenotokous reproductive patterns of two spider mite species (Acari: Tetranychidae), the long-seta form ofSchizotetranychus celarius (Banks) andTetranychus urticae Koch, revealed that reproduction of unfertilized females of the former is very differnt from that of the latter. Unfertilized females ofS. celarius, which has a subsocial life, laid a few eggs and then became inactive. In contrast, the fecundity of unfertilizedT. urticae females was only slightly reduced as compared with fertilized females. Mother-son matings may thus sometimes occur in naturalS. celarius populations. A two-year field survey revealed that, in the absence of males, overwintering females ofS. celarius occasionally remain unfertilized until early spring. Furthermore, nest foundation observed in late spring indicated that most of the season's first nests were founded by single females. These two sets of observations strongly suggest that motherson mating takes places in nature, corresponding to the reproductive trait seen in the experiment. Mother-son mating inevitably increases the relatedness between nest members. The estimated father's relatedness to its offspring is extraordinarily high under such condition. The possibility that kin-selection in the long seta-form ofS. celarius led to subsociality, especially paternal care, is suggested.

Presentation d'un modele mathematique simulant la dynamique de populations interactives d'un acarien predateur (P.p.) et de sa proie (T.u.) et developpe pour determiner quels sont les traits du systeme predateur-proie qui ont l'effet le plus marque sur le comportement de ce systeme. Etudes de validation

The numerical response of adult femaleT. pyri feeding on different levels of ERM larvae, and at a range of temperatures was examined. The duration of the pre-oviposition period decreased as larval consumption increased, and the rate of oviposition was linearly related to the rate of consumption of larvae. Mathematical models were used to describe these relationships in terms of temperature and consumption of ERM larvae. The results were discussed in relation to the nature of theT. pyri/ERM interaction in New Zealand apple orchards.

This study examines the responses of the predatory mite,Phytoseiulus persimilis, to the density and distribution of its prey,Tetranychus urticae. It is divided into three parts. Firstly, the functional responses of protonymph, deutonymph and adult females towards different prey stages are displayed. The great majority of the responses are of the type II form, and the variations in the values of attack ratea′ and handling timeT h are discussed. Experiments are then described in which individual protonymph, deutonymph and adult female predators are presented with varying ratios of two prey age-classes (eggs and deutonymphs, and larvae and deutonymphs). Any observed preference for one of the prey stages is discussed in relation to the predicted preference on the basis of the separate functional response experiments. Finally, the response of different densities of adult female predators to a non-random distribution of deutonymph prey on bean leaflets is examined. The predators show a clear tendency to aggregate on the leaflets of high prey density, counteracted to some extent by interference increasing the probability of dispersal to other leaflets.

Life history and resource utilization pattern were compared between two closely related mite species of the genusTetranychus.1. Tetranychus urticae developed more quickly and had a higher oviposition rate thanT. kanzawai. Consequently, the potential for population increase was greater forT. urticae than forT. kanzawai but the difference was slight. 2. T. urticae was more tolerant of a deteriorating food resource and/or over-crowding.T. kanzawai responded more sensitively to food deterioration and dispersed more quickly. 3. There was a considerable difference between injury patterns by the two species.T. kanzawai damaged host plants more severely, causing earlier and more extensive defoliation of the plants. 4. The maximum population size achieved byT. urticae was 2 to 3 times greater than that ofT. kanzawai on potted host plant. This showed that the former could utilize a food resource much more efficiently.

The distribution ofPanonychus ulmi (Koch) andTetranychus urticae Koch as observed in Swiss apple orchards was analyzed in a multi-stage sampling procedure. Based on the variance-mean relationship (Taylor, 1961) a transformation of mite counts was made. Major sources of variation were found in between-tree and between-crown level differences. In usingBliss andOwen's (1958) method the negative binomial model was concluded to be appropriate for the description of theP. ulmi but not for theT. urticae distribution. HenceCroft's et al. (1976) sampling statistics was recalculated forP. ulmi only. The mean crowding index (Lloyd, 1967) was linearly related (Iwao, 1968) to the mean forP. ulmi and to a less extent forT. urticae that was found in much lower density range. Based on the mean crowding statistics a sampling program afterKuno, (1976) is proposed for both species. Some differences between the sampling plans and their applicability are discussed.

Home areas of hosts are, in effect the gardens in which parasite populations grow. The parasites flourish in response to the kind and amount of host activity modified, of course, by the habitat, and become microcommunities supported by their host and its artifacts. When scatter diagrams of points at which hosts have been observed are divided by median major and minor axes, and when the resulting four quarters, designated A, B, C, D, corresponding to concentration of points in the quarters, the relative concentration usually is: A>B>C>D. The concentration of observed points also is generally greatest nearest the major axis and frequently, but not always, near the median center. Nests of hosts, hence nest-parasites, appear generally to be near the center. For the host examined in detail here, the New Guinea coarse-haired rat, mostLeptotrombidium deliensis appear to have been acquired in the AB sector of the host's home areas but a greater diversity of chiggers is acquired in the CD sector. It is postulated that, ifL. akamushi had been present, it would have occurred in the CD sector where, though the observed concentration of host activity was least, the mixed grassforb and shrub habitat would have been more suitable than in the AB sector.Gahrliepia pingue, C. ewingi, Schoengastia blestowei andS. schueffneri, appear also to be more abundant in the CD sector than in the AB sector. The chiggers themselves have home clusters, or lairs, and adjacent areas of activity (home areas) somewhat resembling those of the hosts.

Two multivariate statistical procedures were used to determine the basic trends of morphologic and geographic variations between males of a common stored-grain mite,Glycyphagus destructor (Schrank) collected from Canada and Japan. All analyses were carried out on physical measurements of 25 morphological features. Three principal component analyses bases on the Canadian (50 specimens), Japanese (50 specimens) and the combined populations from the 2 countries (100 specimens) revealed that the first component, accounting for 40% of the variability in all 3 solutions represented a measure of the morphologic dimension of the body. The second component, which explained over 18% of the variability, is a measure of the leg dimension. Smaller clusters of variates characteristic of the population from each county were also evident. Discriminant analysis, performed with the Canadian and Japanese populations, identified the variates that differed between the 2 populations and provided an approximate appraisal of interrelations. The general conclusion based on these analyses is that the Canadian and the Japanese populations are morphologically distinct. The difference is most evident in the diameters of genu 2, lengths of the sensory seta WI, lengths of the body, and the distances between the vertical external setae.

Dispersal behaviour was studied on a predacious phytoseiid mite,Phytoseiulus persimilis Athias-Henriot in response to the density of its prey,Tetranychus kanzawai Kishida. And the effect of the change in the rate of successful dispersal of the predators among patches was tested on the persistence of the predator-prey system. The results of the study are summarized as follows:1. With a severe decline in prey density available per individual predator, the predators exhibited a marked behavioural change and dispersed to other areas.

Reproductive compatibility was studied among populations of different types of the citrus red mite,Panonychus citri (McGregor), i.e., the diapausing type from pear (DP), the non-diapausing type from citrus (C) and that from pear (NP). Copulation was also observed between mates of different types (DP and C). Only in crosses between C ♀ and DP ♂, was copulation occasionally broken off prematurely; duration of copulation varied considerably between pairs and the average duration was much shorter than that between DP ♀ and C ♂ and than that in crosses between mates of the same types. No F1 adult females were produced at all from crosses between DP and C, showing that there was a complete reproductive isolation between the two types. There was a significant reciprocal difference in the egg hatchability and survival rate of immatures in the F1 progeny; mortality in the eggs and that in the immature stage were significantly higher in crosses between DP ♀ and C ♂, as compared to those in the reciprocal cross. This suggested that fertilization may have occurred in the former cross, whereas it seemed that eggs were not fertilized in crosses between the C ♀ and DP ♂. The NP was compatible with the C, whereas it was completely incompatible with the DP. Thus, there was a complete reproductive incompatibility between the diapausing and non-diapausing type ofP. citri.

I sampledTetranychus urticae Koch (Acarina: Tetranychidae) regularly from four rose gardens in Kyoto and Nara Prefectures in 1988–1990. When mite density was low,T. urticae showed an uneven vertical distribution, being more abundant in the lower third and absent from the upper third of the plants. Mite density was less variable within than between plants, suggesting that a sampling plan which includes more leaves from different plants rather than from different levels of a plant is favourable. Spatial distribution ofT. urticae was nonrandom and followed the negative binomial distribution. In addition, both the Taylor’s power law and the Iwao’s patchiness regression described the distribution well. An empty-sample method for estimating mite density from the proportion of empty sampling units was developed. Sampling plans for determining the sample size required to reach a predetermined precision level, based on this method and by directly counting the mites, were designed. The counting method was more accurate than the empty-sample method. However, when the time factor was taken into account, the latter was more favourable, because it was faster than the former at a density range of 1.5–300 mites leaf−1, which was most commonly encountered in the field.

WhenPhytoseiulus persimilis was reared withTetranychus urticae, infesting roses propagated in a greenhouse at controlled daily temperatures of 24°C (12 hrs) and 18°C (12 hrs), prey numbers fluctuated with peaks of increasing amplitude. Differential dispersal of prey and predator species was one factor contributing to the inability of the natural enemy to control the pest population.

An acarine predator-prey system in a circular stepping-stone environment was described with a simulation model to elucidate the factors responsible for persistence of the system. The main assumptions in this model are: (1) The prey are inevitably eliminated in patches in which predators exist. (2) The density of prey declines and becomes extinct by plant defoliation due to feeding by prey. In this regard this model is different from the models which mimickedHuffaker's (1958) experiments and assumed stable plant-prey relations. Analyses showed that the critical factor in persistence of the predator-prey system was the plant-prey relations, at any combination of other parameters involved in the model. The predator-prey system did not persist long under the unstable relationship of prey and plant. Otherwise the system persisted longer especially when I used a larger number of patches, a larger amount of plant in each patch, and long-distance-migrations of the prey. In particular, frequent emigration of the prey regardless of plant conditions was most effective.

Population dynamics and aggregation patterns of nine kinds of stored-grain mites were studied in two 7.5 tonne lots of hulled (cv. Random) and hulless (cv. Terra) oat cultivars with 12–14% moisture content stored in two wooden bins in Manitoba, Canada during 1978–84. Random oats harbored more mites than Terra oats.Lepidoglyphus destructor Schrank was the most common granivorous mite andCheyletus eruditus Shrank the most common predatory mite. Ecological data on a tydeid mite,Paratriophtydeus coineaui André, are presented for the first time. Because they were more abundant in Terra oats with higher fat acidity values (FAV) than Random oats with lower FAVs,L. destructor andTarsonemus granarius Lindquist could be used as bioindicators of spoilage of stored oats. All species analyzed showed some significant difference in their abundance at different depths in the grain bulk; some species showed depthxtime interaction. Aggregation patterns indicated most mite species had overdispersed (clumped) distribution.T. granarius, andBlattisocius keegani Fox-Androlaelaps casalis Berlese had a distinct aggregation pattern in each oat cultivar.

Interrelations among acarine, fungal, and environmental components of bulk grain ecosystems were determined by canonical correlation analyses. Twenty-seven variables were measured monthly in samples collected from 2 identical grain bulks in a granary in Winnipeg during the years 1959–67. The relationships between 9 kinds of arthropods and 6 environ mental variables, and between the same arthropods and 12 kinds of actinomycetes and fungi were examined. The maximum canonical correlation between arthropods and environmental factors was 0.35, and between arthropods and microorganisms was 0.28; both are highly significant (p<0.001). In the first analysis correlations of the variables with the canonical variates revealed that correlations of the variables with the canonical variates revealed that sampling location, depth, and temperature are the primary environmental antecedents involved, and the criterion is primarily composed of mitesTarsonemus spp.,Tydeus interruptus and the psocid,Lepinotus reticulatus. In the second analysis the fungiNigrospora sphaerica, Aspergillus spp., andCochliobolus sativus are involved with the mitesCheyletus eruditus andAcarus siro. Generally, the results of these analyses complement the findings of factor and regression analyses of the same data reported earlier.

1. Laboratory experiments on the interaction between phytophagous tetranychid mites and predacious phytoseiid mites in multi-patch systems showed that it was difficult to obtain a continued interaction between them when prey heavily over-exploit their host plants. The experiments suggested that the stability conditions concluded from theMaynard Smith model which assumed a stable relationship between prey and resource would be of secondary importance, if the interactive behaviour between prey and resource was unstable. 2. An acarine predator-prey system was theoretically analyzed by simulations on the basis of the structure of theMaynard Smith model, but by introducing modifications to the model, considering several properties characteristic to the prey and predacious mites. The simulations showed that if there was resource over-exploitation by prey, a variety of patterns could be generated in the dynamic behaviour between prey and resource, and depending on the interactive pattern, the predator-prey system displayed a stable behaviour in some cases but in others it became unstable. To achieve a stable interaction, it was important to lower the rate of local prey extinction and this could be achieved either by predation which prevented the prey from over-exploiting the resource or by lowering the rate of successful migration of the prey.

The population dynamics of the housefly,Musca domestica, on patchy and unstable habitats consisting of refuse was investigated at a waste disposal site by using sticky flypaper and mark-release-recapture technique (Jolly-Seber's method). The newly disposed garbage was favorable for breeding of the flies for about one month after being disposed, while a mixture of garbage and ash from incinerated refuse was less favorable. On the garbage under favorable conditions, the rates of population increase was 1.25–2.82 per day, and approximately 1300–1500 flies were produced per square meter within the available period of one month. The rapid decrease in the fly density was observed just after the appearance of high density peaks. The mark-release-recapture study suggested that this rapid decrease would be mainly due to the density-dependent emigration of adult flies from the patchy habitats. The emigration was also activated when the time after garbage disposition became long.

The accuracy and precision of the topological mapping procedure for estimating within-tree populations of bark beetles was investigated for a variety of different sampling conditions. Simulation techniques were used to define the mean and standard deviation of proportional errors encountered in estimation under different sampling intensities. The number of samples collected at a particular height and the vertical spacing between heights were varied. Information presented should aid in developing sampling plans for studies of bark beetle populations and will permit the reexamination and/or recovery of historical data sets on bark beetle populations.

Acoustic signals are part of the specific mate recognition system of planthoppers. The genetic control of acoustic signal characters was studied in the planthopperRibautodelphax imitans. Artificial selection for interpulse interval in the female call revealed a large additive genetic component for this polygenic character. Other female call characters showed a correlated response. Some male call characters also appeared to be genetically correlated with the female character selected for, despite the rather different structure of male and female calls. Parent-offspring regression provided significant heritability estimates for those male call characters that also responded to artificial selection in the female call, one of which appeared to be influenced by sex-linked genes. It is argued that the differentiation of this mate recognition system in planthopper populations and species could be the result of founder effects, enabled by the genetic plasticity of the call characters and the existence of a wing length dimorphism in these animals.

Spatial relationships of mate acquisition probability for individuals of both sexes of a gregariously-mating coreid bug,Colpula lativentris, were studied in relation to aggregation size. Operational sex ratio was always strongly male biased. Mate acquisition probability of females was rather constant and independent of aggregation size, as predicted by an ideal free distribution. Moreover laboratory experiments showed that both multiple mating and rearing density little affected female fecundity, suggesting ideal free distribution of females in terms of reproductive success. On the other hand, mate acquisition probability of males was higher in larger aggregations, where more receptive females were available. This male discrepancy from an ideal free distribution was similar to the patterns predicted by an ideal free distribution under perceptual constraints (Abrahams, 1986), but not by that under unequal competitive ability.