Inconstancy of Taylor'sb: Simulated sampling with different quadrat sizes and spatial distributions

Summary Taylor's power law,s ² =am ^b , provides a precise summary of the relationship between sample variance (s ² ) and sample mean (m) for many organisms. The coefficientb has been interpreted as an index of aggregation, with a characteristic value for a given species in a particular environment, and has been thought to be independent of the sample unit. Simulation studies were conducted that demonstrate that the value ofb may vary with the size of the sample unit in quadrat sampling, and this relationship, in turn, depends on the underlying spatial distribution of the population. For example, simulated populations with hierarchical aggregation on a large scale produced values ofb that increased with the size of the sample unit. In contrast, for a simulated population with randomly distributed clusters of individuals, the value ofb eventually decreased with increasing quadrat size, as sample counts became more uniform. A single value ofTaylor'sb, determined with a particular sample unit, provides neither a fixed index of aggregation nor a complete picture of a species' spatial distribution. Rather, it describes a consistent relationship between sample variance and sample mean over a range of densities, on a spatial scale related to the size of the sample unit. This relationship may reflect, but not uniquely define, density-dependent population and behavioral processes governing the spatial distribution of the organism. Interpretation ofTaylor'sb for a particular organism should be qualified by reference to the sample unit, and comparisons should not be made between cases in which different sample units were used. Whenever possible, a range of sample units should be used to provide information about the pattern of distribution of a population on various spatial scales.     

Natural control of populations of the southern green stink bug,Nezara viridula

Summary Based on the results of five-year study on theNezara population, it was concluded that the level of abundance of the insect was determined partly by the combination and relative adundance of the host plants and partly by the climatic factors. The endless increase of the population was checked by the deleterious effects associated with the overpopulation which occurred locally. The degree of such overpopulation was determined by the relative abundance of host plants to the current level of the population. Therefore, even when the general level ofNezara population seemed to be low, there still exists the possibility of overpopulation in some areas. Winter was the critical period which threatened the persistence ofNezara population. The survival of the hibernating adults was partly dependent on the physiological state of adults which was determined in the period of pre-hibernation. But when the severity of a winter exceeded the tolerance limit of the insect, the only safety-valve which prevented the elimination ofNezara was the heterogeneity of hibernacula. Therefore,Milne's theory (1957) was closest to the known facts.     

Elimination ofSitotroga cerealella (Olivier) bySitophilus zeamais Motschulsky from mixed laboratory cultures on maize

Summary This paper describes a series of experiments conducted to determine whySitophilus zeamais Mots. andSitotroga cerealella (Oliv.) could not survive together in maize cultures in the laboratory. The effect ofS. zeamais on different developmental stages ofS. cerealella was investigated. The presence of adultS. zeamais slightly affected moth copulation, egg laying and moth eggs in a mixed culture, but large numbers of developing moths inside maize grains were killed by the adult weevil through feeding on the grains. The major cause of elimination ofS. cerealella byS. zeamais from mixed cultures was therefore found to be damage to the immature moths in grain and such moth mortality increased as the developing moths became bigger in the grains. A weevil: grain ratio of approximately 1.4∶1 was found to be the critical weevil density at which the moth disappeared from the mixed cultures.     

Sibling and non-sibling cannibalism by larvae of a lady beetleHarmonia axyridisPallas (Coleoptera: Coccinellidae) in the field

Summary Sibling (cannibalism among siblings) and non-sibling cannibalism (cannibalism among non-siblings) were studied in a natural population of a lady beetle,Harmonia axyridis Pallas. Of all the eggs laid (n=2269), 24.76% (n=562) were killed by sibling cannibalism and 36.10% (n=819) were killed by non-sibling cannibalism. Sibling cannibalism occurred constantly and intensively in most egg batches throughout the entire oviposition period. On the other hand, non-sibling cannibalism was more intense in the middle and late oviposition periods, and when the egg batches were close to an aphid colony. This may be due to the high density ofH. axyridis larvae relative to aphid density in the middle and late oviposition periods and also the larvae searching intensively near an aphid colony.     

Functional response and searching efficiency inPseudogonatopus flavifemur Esaki and Hash. (Hymenoptera: Dryinidae), a parasite of rice planthoppers

Summary The functional response ofPseudogonatopus flavifemur E & H (Hym., Dryinidae) was investigated by offering hosts (brown planthopper) at densities ranging from 8 to 160 per cage. The response curve was found to be sigmoid, i. e.Holling's (1959) Type III curve. In experiments involving 310 hosts per cage distributed unevenly in 5 densities (10, 20, 40, 80 and 160 per hill), and a different female parasite density each time (viz. 1, 2, 4, 8 or 16 per cage), the behavioral response was described well by the “random predator equation” ofRoyama (1971) andRoger (1972), which is a convex exponential curve. The area of discovery (a) decreased with an increase in female parasite density (P), and the relationship was described by the equation: loga=−1.0099−0.3638 logP. There was an apparent increase in handling time per host as the number of female parasites increased. Superparasitism, a rare phenomenon under natural conditions, was often observed in the laboratory. The potential ofP. flavifemur as a biocontrol agent of the brown planthopper is discussed.     

Time-specific life tables for the pea aphid,Acyrthosiphon pisum (Harris ), on alfalfa

Summary Time-specific life tables were constructed for three pea aphid,Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), populations using a modification ofHughes' analytical procedure. All populations were studied on second-growth alfalfa (mid-June to mid-July) in south central Wisconsin; data for two populations were collected during 1980, and data for the third population were collected during 1982. The intrinsic rate of increase (r _m) estimated on a physiological time (day-degree) scale under field conditions but in the absence of natural enemies, provided a reliable estimate of potential population growth rate and was used in preference toHughes' approach of estimating potential population growth rates directly from stage structure data. Emigration by adult alatae and fungal disease were the major sources ofA. pisum mortality in each of the three populations studied. These factors were most important because of their impact on reducing birth rates within the local population. Parasitism was never greater than 9 percent. Mortality attributable to predation ranged from 0.0 to about 30.0%; however, even at the highest predator densitiesA. pisum populations increased exponentially.     

Extraordinary effects of fertilization status on the reproduction of an arrhenotokous and sub-social spider mite (Acari: Tetranychidae)

Summary 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. This study was supported in part by Grant-in-Aid No. 61540468 from the Ministry of Education, Science and Culture, Japan.     

Population studies ofBrevicoryne brassicae (L.), its parasites and hyperparasites in England

Summary Studies on populations ofBrevicoryne brassicae (L.), its parasites and hyperparasites were carried out by actual counting in the sprouts field and by sticky and water traps.B. brassicae was found to be attacked by one primary parasite,Diaretiella rapae (McIntosh), which in turn is parasitized byAlloxysta brassicae (Ashm.),Asaphes vulgaris Walker,A. suspensus (Nees),Pachyneuron minutissimum (F?rster) andDendrocerus carpenterii (Curtis). The aphid population in the field was started by immigrant alates which were found flying too early to be synchronized with the sprouts plants. SimilarlyD. rapae was not synchronized with the aphids although many individuals could have been carried into a plot through parasitized immigrant alates, of which less than 30% were found parasitized. Because of high hyperparasitism (especially byA. brassicae)D. rapae was not able to maintain a high rate of parasitism to curb the aphid population growth. The maximum percentage mummies being 27.8%, while the maximum, percentage parasitism being 56.6% recorded only during the early 1974 season (mean=12.9%). The decline of aphid population from September onwards was largely due to the cold weather, Syrphid predation and occasionally fungal attack. The high rate of hyperparasitism byA. brassicae is attributed to its better synchronization withD. rapae. The mean percentage of parasite that emerged from mummies collected during 1973–74 wereD. rapae 31.3%,A. brassicae 64.3%,A. vulgaris andA. suspensus 4.3%,D. carpenterii 0.2% andP. minutissimum 0.1%.     

Studies on the basic components of the predation ofPhytoseiulus persimilis Athias-Henriot (Acarina: Phytoseiidae)

Summary The basic components of the predation ofPhytoseiulus persimilis Athias-Henriot feeding upon eggs ofTetranychus urticae Koch were studied in an open system where the predator could disperse freely. The type of the functional response of the predator to the density of its prey was the same as that studied so far in a closed system, i.e.,Holling's Type 2. The search rate of the predator, however, was much lower in comparison with the result from a closed system. The oviposition of the predator per day was only weakly related to prey densities higher than 10 per leaf disc. But the emigration rate was inversely dependent upon the initial prey density up to 60 per leaf disc. The searching behaviour of the predator was influenced by both the web density spun byT. urticae and the density of the prey: the predator searched for its prey intensively only after it had contact with web. Mutual interference was observed in prey consumption, but not in the emigration rate. The emigration rate was largely dependent upon the prey density available per predator.     

Principal component analysis of interrelations in stored-wheat ecosystems infested with multiple species of insects

Summary Principal component analysis (PCA) was used to determine the effects of infestation of bulk stored wheat by multiple species of insects at 30±2°C for 60 wk. Eight 204-liter drums containing wheat at 15.5% moisture content were used as three distinct man-made ecosystems: (a) Control system (2 drums), insect-free; (b) RST system (3 drums), artificially infested withRhyzopertha dominica (F.),Sitophilus oryzae (L.), andTribolium castaneum (Herbst); and (c) COT system (3 drums), infested withCryptolestes ferrugineus (Stephens),Oryzaephilus surinamensis (L.), andTribolium castaneum. The variables measured tri-weekly within each system included carbon dioxide, oxygen, temperature, grain moisture, seed damage, grain weight and volume, dust weight and volume, fat acidity values (FAV) of the wheat, seed germination, microflora includingAlternaria alternata (Fr.) Keissler,Aspergillus glaucus group,Aspergillus candidus Link, and bacteria, insects and the miteTarsonemus granarius Lindquist. PCA provided multivariate synopsis of the data quantifying several important relationships among the variables monitored. Tri-weekly and cumulative 60-wk analyses of each system showed that high bacterial counts were associated with high FAV levels;Tarsonemus numbers were positively related toAspergillus; Alternaria and seed germination were negatively related to FAV, bacteria and grain damage; and that the number of insects was related to the presence ofAspergillus and negatively related to the presence of bacteria. Seed germination andAlternaria infection often decrease rapidly presumably because of infection by fungi of theAspergillus glaucus group. The combined action ofR. dominica andAspergillus spp. enhanced seed damage and increased grain moisture content thus promoting bacterial growth which in turn inhibited insect and mold growth. Fat acidity values increased with time unless seed damage and bacterial infection were extensive as in the RST system. Contribution No. 903, Agriculture Canada, Research Station, 195 Dafoe Road, Winnipeg, Manitoba, R3T 2M9.     

Anopheles culicifacies giles: Adult ecological parameters measured in rural punjab province, pakistan using capture-mark-release-recapture and dissection methods, with comparative observations onAn. stephensi liston andAn. subpictus grassi

Summary Dispersal, immigration and emigration rates, horizontal and vertical survivorship and absolute population size were estimated for micropopulations ofAn. culicifacies, An. stephensi andAn. subpictus at a series of cattle sheds in rural Punjab Province, Pakistan, during November 1979 and May 1980 using capture-mark-release-recapture and dissection methods. Dispersal was temperature-related, with populations more vagile during May. Mean dispersal distance per individual was low for all species. More than 70% of all recaptures were taken at the point of release and the longest detected flight was 1250 meters. Horizontal survivorship was greater during November and was always less than vertical survivorship calculated from dissection agegrading data. Survivorship during the nulliparous period was greater than survivorship throughout total life, indicating the survivorship curve may be slightly sigmoid. Daily population sizes of endemic and immigrating females and males were calculated usingBailey's (1952) modification of the Lincoln Index, with the daily captures adjusted for immigration which was highest in May. Daily additions to the indoor resting population exclusive of immigrants were estimated using the method ofManly andParr (1968). The relationship of the present findings to malaria transmission and genetic control were discussed.     

A model for analyzing insect stage-frequency data when mortality varies with time

Summary A model is developed for the analysis of insect stage-frequency data which may be applied to populations with age-dependent mortality. The analysis of stage-frequency data is divided into two steps. In the first step, the number of different mortality rates and their values are estimated. The second step provides estimates of developmental rates and variances for each developmental stage and in addition provides estimates of the number of recruits to each stage. The model may be used both in analysis and prediction of insect stage frequencies. Hence, in addition to estimating developmental and mortality rates from stage-frequency data, it may also be used as a simulation model for an insect population. The model is applied to two populations ofHemileuca oliviae Cockerell, a lepidopterous pest of New Mexico grasslands. The model identifies, in the two populations, different mortality rates that are related to plant productivity.     

Age-specific survivorship analysis ofHeliothis spp. Populations on cotton

Summary Population dynamics ofHeliothis virescens (F.) andHeliothis zea (Boddie) (Lepidoptera: Noctuidae) eggs and larvae were studied for two years in a small plot of cotton,Gossypium hirsutum (L.). Due to morphological and ecological similarities, the pooledHeliothis population was considered for most of the analyses. Two generations ofHeliothis eggs and larvae were completed during each year. Stage recruitment was estimated for the eggs and larval instars 2–6, and recruitment variances were estimated by a Monte Carlo method. A modified form of the Weibull distribution was developed and used as a model to characterize survivorship curves for each of the fourHeliothis generations. A Type I survivorship curve (mortality rate increasing with age) was inferred for both Generation 1 (early season) data sets, whereas a Type II survivorship curve (mortality rate constant and thus independent of age) was inferred for both Generation 2 (late season) data sets. The shapes of the survivorship curves for the individualH. virescens andH. zea populations were inferred to be the same as those for the pooled populations. Analysis of the contributions of various factors toHeliothis stage-specific mortality indicated that natural enemies (predators and parasites) and the availability of food for larvae were responsible for between-generation differences in survivorship patterns.     

Comparison of male adult population densities of the oriental and artocarpus fruit flies,Dacus spp. (Diptera: Tephritidae), in two nearby villages in Penang, Malaysia

Summary The populations of native male adult oriental fruit flyDacus dorsalis (Hendel) and artocarpus fruit flyD. umbrosus (F.) in two selected site (BU and SD) were estimated weekly by the capture-recapture technique using live traps baited with methyl eugenol. In BU where many varieties of fruit trees were grown, the estimated population densities ofD. dorsalis were between 980 and 3100 male flies per ha between May and July, 1984. During the same period, in SD where there were fewer number and varieties of fruit trees, the estimated population densities were between 300 and 1000 flies per ha. The estimated population densities ofD. umbrosus over the same period were between 570 and 1290 flies per ha in BU; and between 5 and 95 flies per ha in SD. Of a total 6828 markedD. dorsalis flies released only one fly (released 6 weeks earlier in BU) was caught in a different site.     

Population regulation of the euonymus gall midgeMasakimyia pustulaeYukawa andSunose (Diptera: Cecidomyiidae) by hymenopterous parasitoids

Summary Yearly population fluctuations ofM. pustulae were investigated at 19 sites in Kyushu. In sites where a platygastrid is the only parasitoid of the midge, the percentage parasitism was very low in the incipient stage of the outbreak of the midge populations. After the number of midges reached a peak, the midge populations declined as the percentage parasitism increased, and then the outbreak ceased. On the other hand, in several populations no outbreak was found and the percentage parasitism was constantly at a high level. Therefore, the immediate cause for the outbreak seemed to be a decline of the percentage parasitism. Like the midge, the platygastrid has one generation each year, and its females also emerge in spring to deposit their eggs within host eggs. The decline of the percentage parasitism seemed to be mainly affected by the time lag between emergence periods ofM. pustulae and the platygastrid. In the midge populations parasitized by both the platygastrid and a eulophid (Chrysonotomyia sp.), an extinction of the population was observed, resulting from parasitism by the latter,Chrysonotomyia sp. is polyphagous and multivoltine, and is a late parasitoid, as discussed byAskew (1975). When the density of the midges is very low, the platygastrid may leave the host eggs unparasitized, whileChrysonotomyia sp. may not, because the mature galls are conspicuous.     

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.     

Geographical variation of elytral spot patterns in the phytophagous ladybird,Epilachna vigintioctopunctata (Coleoptera: Coccinellidae) in the Province of Sumatera Barat, Indonesia

Summary Geographical variation in the elytral spot patterns of a phytophagous ladybird,Epilachna vigintioctopunctata, was studied in the Province of Sumatera Barat, Indonesia. Populations ofE. vigintioctopunctata were divided into four major groups (I–IV) by the incidence of spot pattern variations. Group I, occurring in the coastal plains and inland lowlands, and Group IV, confined to the highlands, were the extremes of the spot pattern variations, the latter had many more non-persistent spots and confluences with larger body size and advanced melanism than the former. These two groups were connected with each other via the intermediate groups. A positive relationship was detected between the elevations of sample sites and the average number of non-persistent spots per elytron. Consequently, present results favor the view that the two previously recognized forms ofE. vigintioctopunctata (formae A and B inKatakura et al., 1988) represent a complicated intraspecific variation rather than two distinct sibling species. Elytral spot pattern variations were not different between the sexes or between the beetles collected from different kinds of host plants. Contribution No. 30 of Sumatra Nature Study (Entomology). Supported in part by grants from Japan Society for the Promotion Science for JSPS-DGHE Scientific Cooperation (1980, 1982) and for the RONPAKU program (toI. Abbas, 1984) and Grants-in-Aid for Overseas Scientific survey from the Ministry of Education, Science and Culture of Japan (Nos. 56041027 and 58041030).     

Interspecific competition between two species of Trichogrammatidae III The analytical study on the competition

Summary The experiment was carried out in order to analyze the results of interspecific competition which were mentioned in the previous reports (Sasaba 1964, 1965). The following definite tendencies were obtained and an account was given with regard to the process of competition. (1)T. minutum had a higher reproductive rate thanT. japonicum at a lower density of the parent wasps. (2) The parasitic efficiency ofT. minutum was high as compared withT. japonicum. (3)T. japonicum was consistently superior toT. minutum in the interspecific competition at larval stage. Contribution from the Entomological Laboratory, College of Agriculture, Kyoto University, No. 384.