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.     

Studies on the spatial distribution pattern of larvae of the mosquito,Anopheles sinensis, in rice fields

Summary The spatial distribution patterns of the population ofAnopheles sinensis larvae were studied in the rice field area in the suburb of Urawa city in Japan, during the summer seasons in 1973 and 1974. The distribution pattern of the larval population within the field, analysed by the m−m regression method, indicated that the basic component of larval distribution was not a group of individuals but a single individual and such components were distributed contagiously over the field. This basic pattern did not change significantly according to developmental stage, census date or field. Therefore, we could describe the distribution pattern of the population in a rice field by the single linear regression, x=0.021+1.339x(r²−0.912). Also, the relation for the whole population in the field area including the five fields could be shown by the linear regression, x=0.049+1.749x(r²−0.959). The value of α remained to be nearly equal to zero, but the value of β became larger than the value for the single-field relation. Such a change in distribution pattern seemed to reflect the greater heterogeneity in conditions among the fields than within individual field. Using the information on the distribution patterns mentioned above, some considerations were given on the sampling plans for mosquito larvae, including samplesize determination and application of sequential methods to estimate population size as well as to classify population level.     

Relationship between surplus energy and body weight in fish populations

Summary This paper theoretically analyses the relationship between surplus energy, which is available for either somatic growth or reproduction, and body weight. From the data of metabolism and growth of the biwamasu,Oncorhynchus rhodurus, obtained by Miura et al., a Bernoulli's differential equation is induced to represent the relationship between body weight and the sum of surplus energy and active metabolic rate. Solving this equation gives the amount of surplus energy,f(W_x), as follows:f(W_x) = (αW _x ^1−γ +β^1−γ)^1/(1−γ)−W_x, in which α, β and γ are constants andW _x is body weight at agex. The function is applied to ten fish populations and consequently it is found to be useful for a wider age range and a wider variety of fishes than the conventional function.     

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.     

A probabilistic interpretation of the United Nations’ 1995–2005 population projections

I develop probabilistic interpretations for the United Nations’ 10-year population forecasts by comparing 1995 projections for 212 countries to the population sizes reported for 2005. Errors in the estimation of the intrinsic rate of increase, presumably caused by erroneous assumptions about birth, death and/or immigration rates, appear to be more consequential than errors based on inaccurate estimation of the starting, or ‘jump-off’, population size. For only about 20% of the countries did the ‘actual’ 2005 population size fall between the United Nations’ low- and high-variant projections. I propose prediction intervals for country-specific population sizes 10 years in the future of the form [ N_i^￠ (t+10) / k ,  k ·N_i^￠ (t+10) ],[ N_i^{\prime} (t+10) / k , \, k \cdot N_i^{\prime} (t+10) ], where N _i ′(t + 10) is the medium-variant prediction for year t + 10 made in year t, and k is a number that varies with starting population size. Based on the 1995–2005 United Nations’ data, values of k giving 95% coverage range from 1.11 for countries with a population on the order of 10⁹, to 1.45 for countries with a population of 10⁵.     

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.     

Yearly variation in recruitment and its effect on population dynamics inYoldia notabilis (Mollusca: Bivalvia), analyzed using projection matrix model

Summary Long-term variation in recruitment was estimated by constructing projection matrices for a marine bivalve,Yoldia notabilis, at two stations in Otsuchi Bay, northeastern Japan, and the effects of its variation on population dynamics were examined using a simple matrix model. The matrix model was developed from the Leslie matrix, in which the population growth rate λ was expressed as a function of recruitment rater ₀. The equilibrium recruitment rater _s, or the recruitment rate required to maintain population at constant size (λ=1), was expressed by the reciprocal of the reproductive value of a newly recruited individual. The estimates ofr _s for the field population were lower at the shallower station than at the deeper station, reflecting higher survivorship and fecundity. Past recruitment rate estimated both by the field samplings for 3 years and by the back-calculation from the current age structure for over 10 years showed large yearly variation, ranging between 0 and 58.6×10⁻⁴. The estimates were larger thanr _s, and hence, large enough to increase population size (λ>1) only in approximately one-third of the estimated years. This suggests that the population has been maintained by occasional successful recruitment occurring once every few years.     

On the mathematical basis of the variance-mean power relationship

The mathematical basis of a widely-known variance-mean power relationship of ecological populations was examined. It is shown that the log variance (S ²)—log mean, (m) plot is virtually delimited by two lines logS ²=logn+2 logm and logS ²=logm, thus increasing the chance that a linear regression line can be successfully fitted, without a profoundly behavioural background. This makes difficult the task of interpreting a successful fit of the power law regression and its parameterb in a biologically meaningful manner. In comparison with the power law regression, Iwao'sm ^*-m regression is structurally less constrained, i.e. has a wider spatial region in which data points can scatter. This suggests that a comparison between the two methods in terms of how good a fit is achieved for a particular data set is largely meaningless, since the power law regression may inherently produce a better fit due to its constrained spatial entity. Furthermore, it could be argued that a successful fit in Iwao's method, when found, is less taxed with mathematical arterfacts and perhaps more clearly linked to some biological mechanisms underlying spatial dispersion of populations.     

On the mode of selection inEscherichia coli

Summary Competition experiments between a lac⁺ prototroph and a lac⁻ auxotroph ofEscherichia coli were conducted by serial transfer procedure on four levels of sugar concentration. Changes of the relative frequencies of both genotypes were followed and the relative fitness was estimated. Fitness was proved to be density-dependent. No explicit dependence of fitness upon genotype frequencies could be detected by the simple method ofKosuda (1981).     

Concepts of aggregation and their quantification: A critical review with some new results and applications

Summary Various indices of spatial patterns based on plot counts are reviewed for theoretical population models appropriate to ecological studies. It is seen that many of the indices proposed in the literature are essentially equivalent to either the index ω= σ²/μ or to the index γ= (σ²-μ)/μ², thus providing a variety of motiviations and interpretations of these two indices as measures of spatial patterns. A vector approach to measuring spatial patterns which suggests both a unifying relationship between these indices and an extension of them is proposed. This leads to an interpretation of the measures of spatial patterns in terms of the transition probabilities of a pure birth process. Part of a Doctoral Dissertation of the junior author at the Pennsylvania State University.     

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

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

Analyses of spatial patterns and population processes of clonal plants

The nonrandom spatial structure of terrestrial plants is formed by ecological interactions and reproduction with a limited dispersal range, and in turn this may strongly affect population dynamics and population genetics. The traditional method of modelling in population ecology is either to neglect spatial pattern (e.g. in transition matrix models) or to do straightforward computer simulation. We review here three analytical mothods to deal with plant populations in a lattice-structured habitat, which propagate both by seeds that scatter over the whole habitat and by vegetative reproduction (producing runners, rhizomes, etc.) to neighboring vacant sites. [1]Dynamics of global and local densities: Dynamical equations of population density considering nearest-neighbor correlation (spatial clumping) are developed as the joint dynamics of global average density and local density (comparable to mean crowding) based onpair approximation. If there is a linear trade-off between seed production and vegetative reproduction, the equilibrium abundance of the population may be maximized by engaging both means of reproduction. This result is accurately predicted by the pair approximation method, but not by mean-field approximation (neglect of spatial structure). [2]Cluster size distributions: Using global and local densities obtained by pair approximation, we predicted cluster size distribution, i.e. the number of clusters of occupied sites of various sizes. [3]Clonal identity probability decreasing with distance: Multi-locus measurement of allozymes or other neutral molecular markers tells us whether or not a given pair of individuals belong to the same clone. From the pattern of clonal identity probability decreasing with the distance between ramets, we can estimate the relative importance of two modes of reproduction: vegetative propagation and sexual seed production.     

Sex change and spatial distribution pattern in an intertidal holothurian Polycheira rufescens in the reproductive season

The patterns of sex change and spatial distribution in an intertidal holothurian, Polycheira rufescens (Chiridotidae; Echinodermata), were investigated on a stony beach in Amakusa, western Kyushu. Field caging experiments revealed that some individuals of P. rufescens underwent a sequential sex change from male to female via hermaphrodite stages and back to male again within a single reproductive season. The sex ratio of the population gradually changed from male dominance at first to equal proportions of males and females as the reproductive season progressed. Toward the end of the reproductive season, immature or spent individuals increased in proportion. Stone size appeared to be an important factor affecting the occurrence of P. rufescens individuals on a stony beach. Analysis of spatial distribution by means of Morisita's index of dispersion and nearest neighbor distances indicated that (i) males showed a slightly stronger tendency to aggregate than females, while females had a tendency toward uniform distribution; (ii) females tended to attract males, as shown by relatively short female-to-male distances; and (iii) there was a substantial variation in male-to-female distances, such that some males were positioned close to females while others were not. It may be suggested that spatial distribution of P. rufescens individuals during the reproductive period is partly dictated by the differential needs of individuals of different sexual states. Received: January 25, 1999 / Accepted: June 10, 1999     

An index of spatial distribution

Summary The index , which takes the value of −1 for a perfectly regular distribution, +1 for a highly aggregated distribution and 0 for the distribution implied by the theoretical varianceσ ², is proposed and a significance table provided.     

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.     

Larval growth dynamics ofHemipyrellia ligurriens (Calliphoridae) andBoettcherisca formosensis (Sarcophagidae) in crowded and uncrowded cultures

Summary Larval growth patterns ofHemipyrellia ligurriens (Calliphoridae) andBoettcherisca formosensis (Sacophagidae) in crowded and uncrowded cultures were compared. Growth of the larvae followed a sigmoid curve. The highest larval growth rates were 0.33 and 0.36 mg h⁻¹ for uncrowded and crowdedH. ligurriens respectively. The corresponding figures were 2.38 and 1.23 mg h⁻¹ forB. formosensis. Larvae of both species attained maximum weight earlier in crowded cultures than in uncrowded cultures, although the final weights attained in crowded cultures were less. The earlier period of most rapid growth in both species was interpreted as a result of intraspecific facilitation at higher larval densities. UncrowdedB. formosensis had a shorter larval to pupal development and an earlier period of most rapid growth than uncrowdedH. ligurriens, suggesting the former may be superior in exploiting carcasses with limiting food.     

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.     

Survival Convergence and the Preceding Mortality Crossover for Two Population Subgroups

In this paper, we present and develop the argument that if the survival functions for two population subgroups converge in later life, a mortality crossover must precede the occurrence of this convergence. Specifically, two survival curves, S ₁(x) and S ₂(x), associated with two distinct population subgroups, G₁ and G₂, tend to converge before all members die out, as often observed and anticipated. This convergence leads to an increased mortality acceleration for the “advantaged” group, and eventually fosters the occurrence of a mortality crossover. We present a mathematical proof for this relationship and offer several explanations for the mechanisms involved in the process of survival convergence and the preceding mortality crossover. This new presentation demonstrates that mortality crossover is a highly observable demographic event given the trend of survival convergence in later life.     

Loss of genetic variability in a fragmented continuously distributed population

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

Adult population parameters and life tables of an epilachnine beetle (Coleoptera: Coccinellidae) feeding on bitter cucumber in Sumatra

Summary The population dynamics of an epilachnine beetle, which is closely related toEpilachna sparsa Dieke (henceforth called “sp. C”) and feeds on bitter cucumberMomordica charantia, was studied by mark-recapture of adults and the construction of life tables. The study was repeated three times, i.e., March–May, July–September and October–December in 1982, in Padang, Sumatra, Indonesia. After the establishment of the host plants, adults of “sp. C” soon colonized, and each study period ended in the death of the plants due to defoliation by the larvae and adults. The estimated mean length of residence of adults ranged from 6–11 days, but this was probably much shorter than the actual longevity, because the adults were so active that they flew away, or dropped off the plants, when they were approached or slightly disturbed. Life tables indicated that egg mortality ranged from 17.8–53.9%, and a parasitic waspTetrastichus sp. B made up 41.1–64.2% of egg mortality. Two wasps,Tetrastichus sp. C andPediobius foveolatus killed 1.2–19.4% (7.6–100%)^* of 4th instars and only the latter species attacked the pupae, killing 24.6–59.1% (45.1–72.4%). Parasitism and starvation by overcrowding contributed most to the total mortality from egg to adult emergence, which ranged from 89.4–99.5%. “Sp. C” had a higher diversity and level of parasitism than the Japanese species,E. vigintioctopunctata. The high dispersal power of “sp. C”, coupled with the prolongedl _x−m_x schedules shown under laboratory conditions, was advantageous for exploiting the food plant which was available throughout the year, but was rather patchily distributed in space.