Analysis of spatial association between two species based on the interspecies mean crowding

Summary and Conclusion The measurement of spatial association between two species is considered on the basis of interspecies mean crowding. Two indices of overlapping, γ andC _p, are derived as geometric and weighted arithmetic means of the same component ratios related to inter-and intraspecies mean crowdings. Both indices behave in a similar way, ranging from 1 when the distributions of two species are completely overlapped to 0 when they are completely exclusive with each other. The former is essentially identical with indices proposed byKuno (1968) andPianka (1973), and the latter is a modified form ofMorisita's (1959)C _σ index. Indices to measure the degree of spatial correlation between species, Ω andR _μ, are then derived for both kinds of overlapping indices, which vary from 1 in complete overlapping, through 0 in independent occurrence, to −1 in complete exclusion. Various kinds of interspecies association are analyzed using these indices and an extended form of the regression graph which provides a convenient way of indicating the spatial interrelation between two species as well as distribution patterns of respective species. The method presented in this paper may also be applicable to compare temporal distribution patterns between species, similarity between communities, etc. For such a wider application which includes continuous as well as discrete distributions, the interpretation of intra-and interspecies mean crowdings is not necessarily appropriate, and hence the concept of mean concentration with the symbols and for intraspecies relation and and for interspecies relation is suggested. This study was supported by Science Research Fund (No. 148041) from the Ministry of Education.     

The importance of non-linearity: A comment on the views of Taylor

Summary Taylor's (1984) view that his power law is better measurement of the distribution pattern of animals than 1/k, I _ρ and relation because almost all the data on means and variances of given species could be fitted by the power low, was critisized. Changes in values of 1/k, I _ρ and suggest the change of distribution pattern of the species, but the data can still be fitted by the power law because of the great linearizing power of log-log plots, and, using the power law only, we shall overlook the information on the meaningful changes in the distribution pattern.     

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.     

Analysis of contagiousness in the action of mortality factors on the western tent caterpillar population by using the \mathop m* - m\mathop m\limits^* - m relationshiprelationship

Summary The contagiousness in the operation of mortality processes on the colonies of the western tent caterpillar,Malacosoma californicum pluviale, was analyzed from two different aspects: successive changes in the frequency distribution of the number of surviving individuals per colony in the course of development, and the distribution pattern of the individuals killed by some biotic mortality factors. Also, for a tachinid parasite,Tachinomyia similis, the analysis was made on the egg-laying pattern on colonies as well as on individual larvae. The methods of these analyses were all based on the relation of mean crowding (m) on mean . A braconid parasite,Rogas sp., tended to kill few individuals together once it attacked a colony, but its effect on host colonies was rather equivalent to the random removal of individuals from all the colonies. Diseases in the late-stage larvae before cocooning was contagious in their action. Nuclear polyhedrosis virus seemed to have no basic contagiousness in its action, but it caused highly contagious distribution of deaths among the colonies when its average incidence was high. A spore-formingBacillus had a tendency to kill several individuals once it appeared in a colony, but the distribution of its incidence (no. of times it appeared per colony) was considered to be nearly at random. The female ofTachinomyia tended to lay more than one egg successively on the same colony. It also attacked individual larvae with a definite tendency for aggregation, which seemed to be resulted from the parasite’s preference to large hosts. When the number of eggs laid on prefered hosts exceeded a certain threshold, however, the fly seemed to change its attention to less attractive, smaller individuals.     

A new method of sequential sampling to classify populations relative to a critical density

Summary A method of sequential sampling for grading population level in relation to a critical density is proposed. The method is based on the relationship and can be used without restrictions on the distribution patterns. The formulae for simple random sampling as well as for two-stage sampling are given. This research was suported by science research fund from the Ministry of Education.     

Ecological studies of collembolan populations in a pine forest soil IV. Comparison of distribution patterns

Summary The distribution pattern of ten species of Collembola was studied during the four years period from July 1971 to May 1975 in a pine forest soil. The distribution patterns were analysed for two scales of distribution, i. e., the distribution over the plot of 10×10 m² and the micro-distribution within a block sample consisting of 36 contigious units each 2×2 cm² in area, by applying the -m regression method. The fundamental pattern which appeared was quite similar for the species examined and individuals were aggregated in response to the heterogeneity of habitat conditions. The causes of aggregations were discussed with regard to some environmental factors. The relative abundances of 10 species within the collembolan community was examined in relation to the habitat utilization and the relative abundance was not related to the degree of aggregation but rather to the area occupied by individuals. This suggests that the more numerically abundant species tend to occupy broader micro-habitat. Biological meaning of aggregation was discussed in connection with the population biology and community organization of collembola.     

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.     

A test ofJackson's method for separating death and emigration with mark-recapture data

Summary Jackson's method for separating death and emigration has been studied by computer simulation. In general this method seems to be quite effective in producing nearly unbiased survival estimates. However, the correction can cause a substantial increase in the variance of these estimates.     

Group predatory behavior by the assassin bugAgriosphodrus dohrniSignoret (Hemiptera: Reduviidae)

Summary Nymphs ofAgriosphodrus dohrni Signoret (Reduviidae) have a strong gregariousness and show group predatory behavior. This study was conducted to clarify adaptive significance of group predation of this species, including laboratory observations and 6-year field surveys. In the laboratory, observations on both solitary and group attacking against armyworms were made at varying prey size classes to compare the capture success rate by solitary predators with that by groups. The efficiency in capturing the prey was significantly higher in group attacking at any prey size class compared. Data obtained from the field surveys indicated the tnedency for searching nymphs to feed in group and to increase the number of predators feeding per prey item with increasing prey size. Average sizes of prey captured were also larger in group feeding throughout the nymphal stage. In particular, it was remarkable that, when prey were “creeping” types, the upper size limit of prey eaten was dramatically increased.     

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.     

Mortality process in relation to aggregation in the southern green stink bug

Summary A simple experiment of simulation was done to analyze the natural mortality process of young larval colonies and egg masses of the southern green stink bug. In this experiment, a degree of contagiousness was allowed in regard to the action of a mortality factor, and was defined as the mean number killed per a colony or an egg mass by the mortality factor within a unit time and the number killed per a colony was assumed to follow the Poisson series with the mean . Thus each component of the Poisson series was opposed to each colony or egg mass which was taken at random from 162 egg masses, 135 and 117 colonies of the first and the second instar larvae, respectively. It was revealed that mortality factors in the field did not act with a small degree of contagiosness, e. g. , on all colonies or egg masses, but acted with a large degree of contagiousness, e. g. , on some of the colonies or egg masses. Thus differential survival somewhat in all or none way occurred among the insect colonies irrespective of their initial sizes. These results were well explained by taking actual mortality factors into account.     

A population study ofStenocranus minutus (Fab. ) (Hemiptera: Delphacidae)

Summary The seasonal and annual fluctuations in a population of the delphacid,Stenocranus minutus (Fab.) onDactylis glomerata L. were studied from March 1968 to September 1970. The study involved the comparison of several sampling methods to estimate the egg, nymphal and adult numbers and an investigation into causes of mortality within each stage. A population budget was made for the years 1969 and 1970.     

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.     

Overwintering ecology of two social halictine bees,Lasioglossum duplex andL. problematicum

Summary Overwintering of two social halictine bees,Lasioglossum duplex (Dalla Torre) andL. problematicum (Blüthgen), was studied. InL. duplex many females stay near the old nests, each female preparing a hibernaculum separated from the natal nest. InL. problematicum most females overwinter communally within the natal nest. This difference in overwintering habits relates to the social structure in the next spring.L. duplex is nearly always solitary in spring although later becoming eusocial, whereas many nests ofL. problematicum are polygynous, beginning in early spring.L. duplex overwinters much deeper ( ) in the soil than doesL. problematicum ( ), but both species are safe from drops in soil temperature, which is above 0°C even in midwinter, and the bees' supercooling points are lower than −6°C. This cold resistance and the storage of sufficient food in the crop are reflected in the winter survival, which is much higher (79%) than the rate of successful nests in the spring active phase (25%). Bionomics of the eusocial halictine beeLasioglossum duplex VII. Contribution No. 2654 from the Inst. Low Temp. Sci.     

Effects of host and parasitoid age on search behaviour and oviposition rates inLariophagus distinguendusF?rster (Hymenoptera: Pteromalidae)

Summary Oviposition rates and related behaviours were quantified forLariophagus distinguendus F?rster attackingCalosobruchus chinensis (L.) andC. maculatus (F.). Oviposition rates varied with parasitoid age; parasitoids aged 1–7 days laid approximately twice as many eggs per day as those aged 8–14 days. Similar differences were noted in search rates and handling times; younger parasitoids had higher attack rates and lower handling times than older parasitoids. Search rates and handling times also varied with the host stage available for attack. Search rates were higher and handling times were lower on larger stages. The results are discussed with reference to their impact on the dynamical behavior of insect parasitoid-host populations.     

Reproductive compatibility between populations of the citrus red mite,Panonychus citri (McGregor ) (Acarina: Tetranychidae)

Summary 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. This work was supported in part by Grant-in-Aid No. 60560049 from the Ministry of Education, Science and Culture, Japan.     

Effects of gall volume on survival and fecundity of gall-making aphidsAdelges japonicus (Homoptera: Adelgidae)

Summary The number of larvae reaching maturity within the gall ofAdelges japonicus was positively related to gall volume, and the relationship between the number of mature larvae and gall volume did not change with different densities of colonized larvae. The population changes in the bud galls ofA. japonicus were surveyed by collecting the galls, which did not suffer predation or parasitism within the galls, from young artificial plantations ofPicea jezoensis over two years. In the year when the density of colonized larvae was high, they suffered a 42% mortality within the galls, whereas mortality was nearly zero in the low density year. The numbers of larvae per gall were positively correlated with gall volume. The regression lines of the number of colonized larvae on gall volume did not differ significantly in the regression coefficients between the two years, whereas the intercept was significantly higher in the year when the density of colonized larvae was higher. However, different within-gall mortality nullified this difference, and the regression lines of the number of mature larvae on gall volume had no significant difference both in the regression coefficients and the intercepts. This suggests that the number of mature larvae per gall was limited by available resources within the gall which were positively related to gall volume. In 25% of the galls in which mature larvae inhabited, the space within the galls were completely filled by the larvae, indicating that space was one of the limiting resources. Gall volume also affected the number of adults that emerged from the gall and the potential number of their progeny.     

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.     

Distribution ofMogannia minutamatsumura (Homoptera: Cicadidae) in the Island of Okinawa: Expansion of distribution range during five years

Summary A five-year study on the microgeographical distribution pattern of a sugarcane cicada,Mogannia minuta (=iwasakii), in the Island of Okinawa revealed that the trapezoid type distribution was an unique feature of the distribution of this species. The rate of expansion of the distribution range was remarkably small despite the continuous occurrence of sugarcane plants. These facts can be explained well by the theory of escape. That is, the epidemic cicada populations should be free from predation pressure in the infested area, but those in peripheral zones could be eradicated by predators. The possible role of regulatory factor(s) which suppress further increase in the density in the infested area was suggested.     

Biology ofHyphantria cuneaDrury (Lepidoptera: arctiidae) in Japan. V. Preliminary life tables and mortality data in urban areas

Summary Studies on the population dynamics of the fall webworm,Hyphantria cunea have been carried out at three survey stations and along selected roads in the urban area of Tokyo since 1966. Twelve survivorship curves obtained during two years and 8 life tables show that the mortality rate in early developmental stages of the fall webworm is remarkably low as compared with that of other lepidopterous defoliaters and the mortality rate in later developmental stages is compensatory high. The low mortality rate in early stages is considered to be due to the protective role of the nest-web and the lack of egg and larval parasites. All but one parasitic species emerge from prepupae and pupae. Spiders living in the nest-web of the fall webworm play an important role in reducing the number of young larvae. Direct observations and caging experiments showed that relatively high mortality during later larval stages is mainly due to predation by birds (in the first generation) and wasps (in the second generation). The generation mortality in the survey stations always exceeded the level where the population is kept at the steady state, and the outbreak of this moth is considered to be continued by the immigration of adults from large trees growing in gardens on which the larvae can escape from predation pressure. Contributions from JIBP-PT No. 51. A part of this study was supported by the special project research, ‘Studies on the dynamic status of biosphere’, sponsored by the Ministry of Education.