A comparative study of the searching efficiencies of a parasite and a hyperparasite

Summary The searching efficiencies of a primary parasite (Diaeretiella rapae (McIntosh)) and a hyperparasite (Alloxysta brassicae (Ash.)) were investigated and compared. In both species, at all parasite densities, there was a curvilinear relationship (P<0.001) between the number of hosts parasitised and the host density. A linear regression (loga=logQ−m logP) was fitted for log area of discovery against log parasite density (P<0.001). The area of discovery for its immediate (i.e. primary) host (viz.Diaeretiella for the hyperparasite and aphid forDiaeretiella) is lower in the hyperparasite than in the primary parasite. InDiaeretialla both the searching efficiency and the mutual interference constant increased (but not significantly,P>0.05) in the presence of its males.     

Spatial pattern of pine wilt disease caused byBursaphelenchus xylophilus (Nematoda: Aphelenchoididae) within aPinus thunbergii stand

Summary To understand the mechanism of spread of pine wilt disease caused by the pinewood nematode,Bursaphelenchus xylophilus, which is vectored by a cerambycid,Monochamus alternatus, the spatial distribution of trees weakened by the nematode was examined within aPinus thunbergii stand from June to October for 4 years. The weakened trees were distributed in a clumped pattern in 1980 and 1981, at an early stage of infestation. In many cases, they showed a double-clumped pattern. The degree of aggregation was higher in June or July than after August. They were uniformly distributed in June or July 1982 and in June 1983 whereas they showed a double-clumped pattern after August. The trees were frequently weakened in June or July when they were near the trees weakened during the previous year. At quadrat sizes of more than 25 m², spatial overlapping was pronounced between trees weakened during June–July of the current year and those weakened in the previous year. The seasonal changes in spatial distribution of weakened trees were explained by the interaction amongM. alternatus, B. xylophilus andPinus trees.     

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.     

Distribution patterns of and sampling plans forTetranychus urticae Koch (Acarina: Tetranychidae) on roses

Summary 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.     

Temporal/spatial structure and the dynamical property of laboratory host-parasitoid systems

The effects of spatial structure in terms of local capacity, or the maximum number of larvae surviving competition at resource patches, and temporal structure in terms of the period vulnerable to parasitoid attack in host populations on the persistence of host-parasitoid systems were quantitatively evaluated by laboratory experiments and well-parameterized model analyses. One of two bruchid beetles,Callosobruchus maculatus andC. phaseoli, were used as a host with Heterospilus prosopidis used as the parasitoid.C. maculatus, in which few larvae survive competition to become adults in each bean, andC. phaseoli, in which many larvae become adults in each bean, along with two kinds of beans, the mung and the azuki, were combined to construct four (2×2) resource-herbivorous host-parasitoid systems that differed in local capacity and vulnerable period. The mung-C. maculatus system with the parasitoid was the most persistent, i.e., took the longest time for extinction of either the host or parasitoid to occur. Since this resource-herbivorous host combination exhibited the lowest local capacity and the shortest vulnerable period, these two conditions possibly promoted the persistence of the system. A model incorporating the host population structure supported the observed persistence. Furthermore, the possible contribution of the timing of density-dependent competition of the host on the host-parasitoid persistence is predicted.     

Measurement of non-randomness in spatial distributions

Summary The measurement of departure from randomness in spatial distributions has widespread application in ecological work. Several “indices of non-randomness” are compared with regard to their dependence on sample number, sample size and density. Criteria for the best choice of index for specific situations are discussed. A new coefficientC _x is proposed for use with positively contagious distributions and tests of significance are given. WhenC _x and another index (S ²/m−1) are used for positive and negative contagion respectively, values ranging from −1 through 0 (random) to +1 are obtained, regardless of sample number, sample size or density.     

The species-area relation II. A second model for continuous sampling

Summary A second mathematical model describing the species-area relation was proposed for continuous expanding of sample area. This model is expressed asS=λ ln(1+x/E) whereS is the number of species occurring in an areax, and λ andE are the constants termedspecific diversity andelemental area respectively. As a result of testing the validity of the model for several sets of data, it was shown that the above equation would provide an adequate fit to a group of species belonging to a single synusia which exists in an open habitat. The ecological implications of parameters involved were discussed and the characteristic area presented previously (Kobayashi, 1974) was defined in terms ofE. The relation between results obtained by discrete sampling and continuous sampling was examined and the possibility of converting one to another was suggested. Contribution from the Laboratory of Applied Zoology, Yamagata University, No. 79.     

Coexistence and weak amensalism of congeneric gall-forming aphids on the Japanese elm

Closely related species of gall-forming aphids are often associated with a single host species. SixTetraneura species coexist on the Japanese elm,Ulmus davidiana, in Sapporo, northern Japan. This paper describes the probabilities of coexistence on macro- and microgeographic scales (i.e., on host trees and host leaves) and examines whether coexistence with conspecific or heterospecific galls on leaves or shoots has any effect on the fecundity of each aphid species using multiple regression. A Monte Carlo simulation showed that the frequency distribution of the numbers of species on individual host trees differed significantly from that expected from the null model. There were significantly positive or negative associations between species.Tetraneura radicicola andsorini always coexisted with other species on trees they infested. Multiple regression revealed that the coexistence of conspecific or heterospecific galls on individual leaves had no influence on the fecundity ofradicicola andsorini, but had a negative influence on that of sp. O. On average,radicicola andsorini produced a smaller number of offspring in galls than sp. O, and obviously consumed only a small part of resources available on the galled leaves. Evidence available suggests that although amensalism does arise between sp. O and other species, its influence in not so strong as to exclude sp. O competitively from theTetraneura community.     

Binomial sampling plans for estimatingBemisia tabaci populations in cantaloupes

Early season infestations of the sweet potato whitefly,Bemisia tabaci (Gennadius), on cantaloupes,Cucumis melo L., were determined by counts of the number of adults per leaf in fields near Yuma, Arizona. We used these data to develop binomial sampling plans based on the relationship between mean densities of whiteflies per leaf,m, and proportion of leaves infested with more thanI whiteflies,P _I, according to the empirical model lnm=a′+b′ ln[−ln(1−P _I)]. The models were developed for the presence-absence approach (I=0) and for a cutoff value of three whiteflies per leaf (I=3). Four independent data sets were used to evaluate the models. Both methods yielded reliable predictions at low infestation levels, but some of the higherm values were overestimated. As the tentative economic threshold forB. tabaci is three adults per leaf, which corresponds to lowP _I values, results of the binomial sampling were satisfactory for pest management purposes.     

Extinction of the trochid gastropod,Umbonium (Suchium) moniliferum (Lamarck), and associated species on an intertidal sandflat

On an intertidal sandflat in western Kyushu, Japan in 1979, the trochid gastropod,Umbonium (Suchium) moniliferum (Lamarck), and the thalassinidean ghost shrimp,Callianassa japonica Ortmann, densely inhabited the lower and upper zones, respectvely.Callianassa japonica subsequently expanded its distribution range considerably, having occupied almost the entire sandflat by 1983. Concurrently, theU. moniliferum population gradually declined, becoming extinct in 1986. Furthermore, the populations of 9 species associated withU. moniliferum (predators, an ectoparasite, and subsequent inhabitants of emptyU. moniliferum shells) had disappeared by the end of 1992. The extinction processes ofU. moniliferum and the two other most numerically dominant species, the ectoparasitic gastropod,Odostomia sp., and the hermit crab,Diogenes nitidimanus Terao, were described in detail. It is believed that the bioturbation of sediments byC. japonica was responsible for the extinctions, possible mechanisms involved being discussed. This is the first documented record of the extinction of a large part of a macrobenthic assemblage centering on a filter-feedingUmbonium species which is characteristic of the benthic communities on many intertidal sandflats extending from Japanese to Southeast-Asian waters.     

Relationship between female body size and demographic parameters inBactrocera Malaysian A (Diptera: Tephritidae)

Summary The effect of body size, as measured by the head width, of the femaleBactrocera sp. Malaysian A (kept separately in sexual pairs) on the demographic parameters was investigated in the laboratory under ambient conditions of 28–30°C, 78–85% RH and natural photoperiod. Body size was shown to influence significantly all the demographic parameters. The expectation of life of females at eclosion from pupae was respectively for head widths of 1.6, 1.8, 1.9, 2.0 and 2.1 mm: 76.2, 73.4, 73.8, 102.4 and 115.2 days. The mean number of eggs laid per female in its life time was respectively: 86.4±48.7, 181.8±56.1, 229.7±72.6, 364.3±69.4 and 477.5±109.3 which was significantly different from one another (F=3.73,P<0.05) especially the two smaller sizes from the two larger sizes. The regression line for total eggs laid (Y) against head width (X) wasY=785.2X−1208.7 (R ²=0.35,P<0.001). The net reproductive rate (R ₀) was respectively 15.8, 34.0, 43.5, 66.9 and 88.8 eggs, while the intrinsic rate of increase (r) was respectivley 0.0435, 0.0538, 0.0670, 0.0665 and 0.0711. The results confirm that for mass rearing purposes, larger females which produce more offspring are to be preferred.     

Spatial pattern analysis of the incidence of aster yellows disease in lettuce

The degree of aggregation of lettuce plants infected by aster yellows phytoplasma (AYP) was investigated in 12 fields from three experiments. Position of diseased and healthy plants was mapped in a 6–9×12-m section of each field; for most analyses, fields were divided into 10-plant quadrats. Mean disease incidence (p) ranged from 0.01 to 0.30. The frequency of diseased plants was described by the beta-binomial distribution, with an index of aggregation (θ) ranging from 0 to 0.17, positively correlated withp, and generally increasing over time within a field. Distance-class analysis revealed a core-cluster size of only a few plants. However, spatial autocorrelations ofp between quadrats were not significant, indicating that the scale of spatial pattern was small, generally less than 10 plants. An overall measure of aggregation was given by the slope parameter of the binary form of the power law, in which the log of the calculated variance is regressed on the log of the theoretical variance for a binomial distribution. The slope was 1.18 and significantly different from 1. Results for this “simple-interest” disease are interpreted in relation to the persistent transmission of AYP by its aster leafhopper vector.     

The role of vole populations in prevalence of the parasite (Echinococcus multilocularis) in foxes

Effects of population fluctuation of the gray-sided vole(Clethrionomys rufocanus) on the prevalence (infection rates) of the parasiteEchinococcus multilocularis in red fox(Vulpes vulpes) populations was investigated from 1985 to 1992 in eastern Hokkaido (Abashiri, Nemuro, and Kushiro area), Japan. This parasite needs two hosts to complete its life cycle; the gray-sided vole as its intermediate host and the red fox as its final host. We found that: (1) Infection rates in foxes depended on the current-year abundance of voles in all three study areas, particularly in Abashiri. (2) In addition to this direct density-dependence, delayed density-dependence between the infection rate and the prior-year abundance of voles was detected in Nemuro and in Kushiro. (3) The regional differences in density-dependence pattern were related to regional differences in the winter food habits of red foxes: in Abashiri the proportion of voles in the fox’s diet greatly decreases in winter, while the proportion remains high in winter in Nemuro and in Kushiro, probably because of shallower snowpack. These results suggest that infection rates in foxes in Abashiri were less influenced by the prior-year prevalence, since the infection cycle might be interrupted in winter, when voles became less important in fox’s diet. In contrast, the state of the prevalence may carry over from year to year in Nemuro and in Kushiro, because red foxes continue to eat a considerable amount of voles throughout year. The regionally contrasted results for the relationship between infection rate in foxes and vole abundance were parallel to the regional difference in fluctuation pattern of vole populations, which are highly variable in Abashiri area, but less variable in Kushiro-Nemuro area. Drastic change in vole populations appears to affect the host-parasite system.     

The Association Between Perceived Spirituality, Religiosity, and Life Satisfaction: The Mediating Role of Self-Rated Health

This study examined the mediating role of self-perceived health between perceived spirituality, religiosity, and life satisfaction among a stratified, random sample of college students, while controlling for gender. Although both models displayed excellent fit criteria, the perceived spirituality and life satisfaction model was fully mediated by self-perceived health χ² (n=459, 4) = 1.64, p=0.80, CFI =0.99, TLI=0.99), and the perceived religiosity and life satisfaction model was partially mediated by self-perceived health χ² (n=459, 10) = 22.29, p=0.01 CFI = 0.99, TLI = 0.99). Both models were equal for men and women. Students who describe themselves as spiritual (or religious) are likely to report greater self-perceived health and greater self-perceived health likely influences life satisfaction for both men and women. Results preliminarily support the contention that life satisfaction is related to differing reported health status, whether physical or mental, and that life satisfaction may be influenced by religiosity and spirituality engagement. Implications for colleges and universities are discussed.     

Inter- and intra-population variations in diapause attribute of the two-spotted spider mite,Tetranychus urticae Koch, in Japan

Summary Populations of the two-spotted spider mite,Tetranychus urticae Koch collected from various localities and from various host plants in Japan showed wide variations in diapause attribute. Diapause percentages at 18°C/9L15D varied from nearly 100% in the north to 0% in the south-west. At intermediate latitudes the mites showed wide inter-population variations. Populations on herbaceous hosts in vinyl- or glass-houses gave significantly lower incidence of diapause than those on roses and deciduous fruit trees. Presence of winter hosts and better host quality under protected environments seemed to favour non-diapausing mites. The temperature threshold for diapause expression also varied widely among local populations. Northern populations consistently had higher and less variable thresholds than populations at intermediate latitudes with thresholds between 15 and 18°C. Inbred lines derived from a population in Kyoto exhibited a wide variation in diapause percentage at 18°C. These results show that diapause inT. urticae is a quantitative threshold trait and that populations in central Japan consist of a variety of genotypes with different diapause traits. This might provide a genetic source for adaptation to local and temporal variations in environmental conditions.     

Accumulation of phyllodulcin in sweet-leaf plants ofHydrangea serrata and its neutrality in the defence against a specialist leafmining herbivore

Among wild plants ofHydrangea serrata (Hydrangeaceae) in Japan, there are sweet plants whose leave contain a kind of isocoumarin, phyllodulcin, which happens to be 350 times as sweet as sucrose to the human tongue. In a primary beech forest in Ashu, Kyoto, the spatial distribution of sweet plants and temporal and the spatial distribution of phyllodulcin within and among plants were investigated using a high performance liquid chromatograph. The distribution of sweet plants was confined within a valley and was parapatric with non-sweet plants. A plant's characteristic phyllodulcin accumulation did not change, even when transplanted into the different habitats. The phyllodulcin content of the sweet plants varied greatly among plants, and the population mean peaked in July when the plants flowered. Within a plant, phyllodulcin content was elevated by partial defoliation. We examined the possible effect of phyllodulcin on herbivory by a specialist leafmining herbivore,Antispila hydrangifoliella (Lepidoptera: Heliozelidae). We transplanted sweet and nonsweet plants reciprocally between their original habitats, excluded attacks by parasitoids, and compared performance of the leafminer. Leafminer colonization and larval survivorship on transplanted andin situ plants was not significantly different between sites. The fact that accumulation of phyllodulcin did not augment a defensive function, at least against herbivory by the leafminer, and the sporadic distribution of phyllodulcin-accumulating plants, suggest that the genotypes synthesizing phyllodulcin emerged independently at separate localities by mutation, and that the genotypes are almost adaptively neutral in defence against the specialist herbivore.     

Spatial variation in female fertility related to interactions with flower consumers and pathogens in a forest metapopulation ofPrimula sieboldii

Antagonistic biological interactions with flower consumers and pathogens may influence reproductive success of flowering plants, affecting population dynamics and natural selection for floral traits. However, ecological and evolutionary consequences of the interactions may depend on both spatial and temporal patterns of the interactions. In a forest metapopulation ofPrimula sieboldii E. Morren, an endangered clonal plant species, we measured between-subpopulation patterns of seed sets and interactions with an influential flower consumer, a rove beetle,Eusphalerum bosatsu Watanabe, and a specific smut fungal pathogen,Urocystis tranzschelina (Lavrov) Zundel (Ustilaginales), for three years. Mean female fertility (seed set per flower) for individual subpopulations fluctuated moderately among years but was highly variable within each year among the five subpopulations studied. In two subpopulations, the impact ofEusphalerum beetle, was sufficiently large to result in almost complete failure in seed production over eight years including the three study and five previous preliminary observation years. In the two other subpopulations, seed set failure was caused by infection by the smut fungus. Infected capsules which constitute 10–30% of the capsules produced in the subpopulations were filled with ustilospores instead of seeds. In the subpopulation that escaped flower damage byEusphalerum beetles and smut fungal infection, seed sets of both pin and thrum flowers were much higher than in the other subpopulations. The spatial restriction of individual antagonistic agents to a part of subpopulations suggest that dispersal of the agents, as well as the mode of spatial subdivision of the plant population would be important for determining the overall effects of antagonistic interactions on plant performances at the metapopulation level.     

A distance measure of spatial pattern in a biological population

Summary Suppose thatn individuals locate independently and randomly on a segment of line of finite length (habitat). Let the theoretical and observed ranges of the sites of the individuals on the segment be μ _n-1 andr _n-1, respectively. Then, the degree of dispersion of the individual sites is measured by the ratio, T _n =n _n-1/μ _n-1, as follows: A random spatial pattern forI _r−1 =1 An aggregated spatial pattern for 0≤I _r <1 A uniform spatial pattern for (n+1)/(n−1)≥I _r >1. Another method was derived. Let the probability that an observed range is less thanr _n−1 beI _p , under the hypothesis of a Beta distribution. Then indicates A random spatial pattern forI _p =1/2 An aggregated spatial pattern forI _p <1/2 A uniform spatial pattern forI _p >1/2. The first index can be used for comparing populations having the same number of individuals, whereas the second one can be used for comparing populations with different numbers of individuals.     

Evaluation of statistical precision and design of efficient sampling for the population estimation based on frequency of occurrence

Summary The binomial sampling to estimate population density of an organism based simply upon the frequency of its occurrence among sampled quadrats is a labour-saving technique which is potentially useful for small animals like insects and has actually been applied occasionally to studies of their populations. The present study provides a theoretical basis for this convenient technique, which makes it statistically reliable and tolerable for consistent use in intensive as well as preliminary population censuses. Firs, the magnitude of sampling error in relation to sample size is formulated mathematically for the estimate to be obtained by this indirect method of census, using either of the two popular models relating frequency of occurrence (p) to mean density (m), i.e. the negative binomial model,p=1−(1+m/k) ^−k, and the empirical model,p=1−exp(−am ^b). Then, the equations to calculate sample size and census cost that are necessary to attain a given desired level of precision in the estimation are derived for both models. A notable feature of the relationship of necessary sample size (or census cost) to mean density in the frequency method, in constrast to that in the ordinary census, is that it shows a concave curve which tends to rise sharply not only towards lower but also towards higher levels of density. These theoretical results make it also possible to design sequential estimation procedures based on this convenient census technique, which may enable us with the least necessary cost to get a series of population estimates with the desired precision level. Examples are presented to explain how to apply these programs to acutal censuses in the field.