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.     

Settling-site selection and survival of two scale insects,Ceroplastes rubens andC. ceriferus, on citrus trees

Summary We studied settling-site selection and the resulting survival of two sessile scale insects,Ceroplastes rubens andC. ceriferus, in the citrus tree,Citrus unshiu, in central Japan. C. rubens preferred 0-year-old twigs most as a settling-site; the density of nymphs settling on 0-year-old twigs was significantly higher than those on ≥1-year-old twigs, and few nymphs settled on ≥3-year-old twigs. The mean survival rates from settling until reproduction in the next year were significantly higher on more preferred twigs than on less preferred ones. InC. ceriferus, nymphs significantly preferred 1- and 2-year-old twigs to 0- and ≥3-year-old ones, and the mean survival rates on the more preferred 1- and 2-year-old twigs were significantly higher than those on less preferred ≥3-year-old twigs. However, the survival rate on less preferred 0-year-old twigs was slightly higher than those on 1- and 2-year-old ones. Thus, in both species of scale, it was the preferred twigs which were more profitable sites for survival after settling, except for less preferred 0-year-old twigs forC. ceriferus. In both scale species, most mortality was due to growth cessation, which is believed to be related to the twig quality as a food source. Predators and parasitoids were minor mortality factors. Both species showed constant survival rates until the density of settled nymphs exceeded double the “upper-limit” density, whereupon they decreased drastically. Nymphs ofC. rubens settling on twigs of high scale density showed a spacing-out distribution, those ofC. ceriferus did not. InC. rubens, an increase in preference for originally less profitable twigs at the later stage of the settling season was observed, but not inC. ceriferus. Accordingly, individuals ofC. rubens showed a stronger tendency to avoid conspecifics than didC. ceriferus. Although nymphs of the two scales clearly preferred more profitable sites, their settling-site selection did not agree with the predictions from the ideal free distribution theory (Fretwell and Lucas, 1970). The discrepancies were (1) frequent settling on less profitable sites at the early stage of the settling season, (2) insufficient utilization of the most profitable twigs, and (3) virtually 100% mortality on overcrowded twigs under conditions where unoccupied profitable twigs still remained. These discrepancies are thought due to the limited dispersal time of nymphs. Contribution to the ecological studies of scale insects 2.     

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.     

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.     

Life table analysis of the green rice leafhopper,Nephotettix virescens (distant) (Hemiptera: Cicadellidae), and efficient vector of rice tungro disease in asynchronous rice fields in Indonesia

Summary Population dynamics ofNephotettix virescens was studied in 17 paddy fields transplanted at intervals of about 1 month in 1988–1990. The adult density was highest either in the immigrant or the 1st generation and sharply decreased to the 2nd generation. The survival rate of the 1st generation was lowest in the transition season when areal population density increased. Key factor analysis revealed that the nymphal and adult mortality of the 1st generation (kn) was the principal source of population fluctuations. No significant correaltion was found between kn and natural enemy density, natural enemy density/healthy egg density, or the precipitation during the nymphal period. On these bases adult emigration was suspected to be the key factor. Areal population build-up ofN. virescens in the transition season was considered to occur as a result of increasing immigration to young stages of rice. Contribution from Indonesia-Japan Joint Program on Food Crop Protection (ATA 162), which was implemented by the Directorate of Food Crop Protection, Ministry of Agriculture, Indonesia and Japan International Cooperation Agency, Japan.     

Some beneficial effects of aggregation in young larvae ofPryeria sinica moore (Lepidoptera: Zygaenidae)

Summary The effects of group size on the survival and development of young larvae ofPryeria sinica Moore were investigated by laboratory and field experiments. Under laboratory conditions, about 20% of isolated larvae died of unsuccessful feeding in the first instar, however, larvae survived successfully in aggregations of four or more individuals. In the field, larvae emerge in early spring and wait for new leaves to open before feeding. In this period, the larger the group size of hatchlings the survival rate became higher. The nest-web spun by hatchlings was considered to play an important role in protecting them from desiccation. In the period that larvae began to feed on leaves, more than 36 larvae are necessary to aggregate for the successful establishment of feeding groups. The nest-web played an important role also in the establishment of feeding group. However, the natural group size of the first instar larvae was larger than the minimum group size to spin a sound nest-web in the field experiment. On the other hand, in later stage, larvae in a large group did not have an excess advantage in survival or developmental rate over larvae in a small group. It was found that the experiments on survival and developmental rates could not explain the reason that this species maintain large compact groups in the most part of larval period. The study was partially granted by the Ministry of Education (No. 374205).     

Species richness and altitudinal variation in the aquatic metazoan community in bamboo phytotelmata from north Sulawesi

We compared the aquatic metazoan community structure in bamboo stumps between a lowland (Kosinggolan; 200 m a.s.l.) and a highland site (Moat; 1030–1050 m a.s.l.) in North Sulawesi. The lowland bamboo stumps harbored 38 taxa including 2 predators, and the highland stumps harbored 35 taxa including 2 predators. In total 45 taxa were recorded, including 3 predators. Dominant detritivores were Tipulidae, Scirtidae, Chironomidae, Culicidae and Ceratopogonidae. The sole dominant predators wereToxorhynchites mosquito larvae, which occurred in 67% and 28% of stumps at the lowland and the highland sites, respectively. Although the mean biomass per stump did not differ significantly between the sites, the mean number of species per stump was significantly smaller at the lowland site. In addition, the variation in species composition among stumps was greater at the lowland site than at the highland site. Among dominant taxonomic groups, the number of non-predatory culicid species per stump was smaller at the lowland site where their predator,Toxorhynchites, was more abundant, although both sites had the same number of culicid species. In the presence ofToxorhynchites, the density and biomass of other culicids per stump were reduced significantly. The difference in predator density might affect differences in the local-scale community structure of individual bamboo stumps.     

Natural mortality at different population densities of a pine shoot moth,Evetria cristata

Summary A field population ofEvetria cristata was studied in 10 plots in 1962 and in 6 plots in 1963. These plots were divided into 2 or 3 groups of different population levels of the shoot moth in respective years. The survival of the insect was then analysed in these different groups of plots. The survival rate ofE. cristata from eggs to adults in the first generation was found always higher in the group with low population density, which indicates the existence of some factors that affect the population more severely when the insect is more abundant.Lissonota evetriae andPediobius sp. seemed to have killed more proportion of the hosts where the shoot moth density was high. However, the total effect of the all natural enemies was not always great in the plots with high density of the moth. The survival of the second generation of the moth in 1963 was observed to be much higher at any population level than in the other generations.     

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.     

Intraspecies competition in a field population ofGregopimpla himalayensis (Hymenoptera: Ichneumonidae) parasitic onMalacosoma neustria testacea (Lepidoptera: Lasiocampidae)

Summary Intraspecies competition in a field population ofGregopimpla himalayensis (Hym.: Ichneumonidae) parasitic on the prepupae ofMalacosoma neustria testacea (Lep.: Lasiocampidae) was investigated. The parasite oviposits the sufficient number of progeny (5 individuals/0.1 g dry weight of host) to exhaust a single host in a single attack. However, at the intensity less than 22–26 individuals/0.1 g d.w. of host, all individuals can emerge, i.e. density-dependent mortality does not occur. Within this range of intensity, survival of parasite larvae is guaranteed by diminution in body size and decreasing sex ratio. In contrast, total biomass of parasites showed a peak at 5 individuals/0.1 g d.w. of host at which a single host is exhausted. Above the intensity of 22–26, extraordinary minute individuals appeared and they died before maturation. If intraspecies competition play a role in regulation ofG. himalayensis population in the field, the process is usually not through density-dependent mortality but through decreasing reproductive rate caused by decrease in the sex ratio, adult longevity and fecundity. Contribution Ser. 2, No. 275. Entomological Laboratory, Faculty of Agriculture, Kyushu University.     

The effects and limits of territoriality on population regulation in grey red-backed voles,Clethrionomys rufocanus bedfordiae

Summary The effects of breeding territoriality on the stability of grey red-backed vole (Clethrionomys rufocanus bedfordiae) populations were investigated on a control grid and a grid on which the voles were fed, in an outdoor enclosure in Hokkaido, Japan. Vole populations were monitored by live trapping from 1984 to 1986: (1) Population density was 2–7 times greater on the experimental grid to which food was added than on the control grid. Reproductive output was more closely associated with the difference in density between grids than survival or dispersal (immigration and emigration) rates. (2) The number of adult females and pregnancy rate of the experimental population were significantly greater than those of the control one. The difference in the number of adult females between the populations was greater than that in pregnancy rate. (3) The proportion of successful litters and the number of weanlings per litter were not significantly different between the control and experimental population. (4) Adult females held territories on both the control and experimental grid; they were spaced out more than would be expected from random occupation. The territories overlapped more on the experimental grid than on the control grid. (5) Mean territory size of adult females on the experimental grid was about half of that on the control grid. The territory size was correlated negatively with population density. (6) The proportion of trap sites that were used by adult females was significantly greater on the experimental grid than on the control grid. This suggests that adult females on the experimental grid used the area more extensively. This factor, in association with territory size and overlapping of territory, was also important in causing the difference in the number of adult females between the grids. (7) These results call into question the hypothesis that territoriality stabilizes the density in populations ofClethrionomys.     

On the nature of errors involved in estimating stage-specific survival rates by Southwood's method for a population with overlapping stages

Summary This paper has examined the effect of within-stage mortality on the estimation of stage-specific survival rates bySouthwood's (1978, p. 358) method. As pointed out bySouthwood, both the severity and timing of mortality affect the mean duration of a life stage, and consequently the estimate of the number of individuals entering that stage. Knowledge of the form of the survivorship curve permits correction of the estimate under certain circumstances. The use ofSouthwood's method with two overlapping stages having different rates and patterns of mortality leads to complex errors in the estimation of survival for the first stage. The nature of these errors is examined analytically and via a simulation model.Southwood's method is fairly robust, with moderate differences in mortality rates leading to acceptable errors in estimating survival for the first stage. When both the rate and pattern of mortality in both life stages are the same, then the survival estimate is made without error. Precise estimates of stage-specific survival will not usually be possible withSouthwood's method because of the errors introduced by the very parameters being measured. Direct measurement of mortality rates and survivorship patterns (seeSouthwood, 1978, p. 309) is strongly advised, at least in preliminary work.     

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.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Dynamics of buried seed population and seedling cohorts of two dominant weeds in a hill agroecosystem of the humid subtropics of India

Dynamics of the buried seeds and plant population of two dominant weeds, viz.,Emilia sonchifolia (Linn.) DC. andRichardsonia pilosa HBK were studied in the crop fields of Meghalaya, north-east India during radish and maize cropping and intervening fallow periods. The total buried seed population ofR. pilosa was always larger than that ofE. sonchifolia, but the germinable fraction was invariably greater in the latter. A major portion (39–41%) of the viable (germinable+dormant) seed population in both weeds was confined to the surface soil layer (0–5 cm). The viable seed population ofE. sonchifolia peaked during April, while that ofR. pilosa showed two peaks (during August and December). The survival pattern and half-lives of seedling cohorts showed, some differences in the two weed species, but both being summer annuals, their populations behaved in a similar manner by showing higher seedling recruitment (K) and survivorship (p) rates in the summer crop (maize) than in the winter crop (radish). However, the density of plants that could attain adulthood was significantly higher inE. sonchifolia thanR. pilosa which might have resulted in greater seed input of the former to the soil leading to its greater abundance in the crop fields. Supported by the University Grants Commission, New Delhi (Grants No. F. 3-37/87 SR II)     

Prey preference of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand)

Summary Functional responses of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand) attacking the rice brown planthopper,Nilaparvata lugens (St?l.), and the mirid predatorCyrtorhinus lividipennis Reuter were both those of Holling Type II. The attack rate was higher and handling time lower forC. lividipennis. However, when caged with the two prey, the wolf spider showed a significant preference forN. lugens at a lower prey proportion. Proportions of prey attacked were significantly different from the expected ratios of prey available as well as from the predicted preferences derived from the functional response parameters. As proportions ofN. lugens attacked changed from greater to less than expected as the proportions ofN. lugens available increased, a “reverse switch” behaviour seems to be evident.     

Analysis and simulation modelling of population dynamics and bioenergetics ofCryptolestes ferrugineus (Coleoptera: Cucujidae) in stored wheat

Summary The consequences of infestation of stored wheat by the rusty grain beetle,Cryptolestes ferrugineus (Stephens) was determined for 222 d at 30°C in 70-1 drums containing wheat at 13.5% moisture content. Temperature, grain moisture, seed damage, germination and weight, dust weight, fat acidity values (FAV), published data on growth, reproduction, survival and cannibalism rates and energy budget were used to develop a computer simulation model to simulate the population dynamics ofC. ferrugineus at 30°C. In the insect-free control system, the fungi,Alternaria alternata decreased,Aspergillus glaucus group andPenicillium spp. increased, probably causing a rise in FAV of the grain. In the insect-infested system,C. ferrugineus could only eat the wheat germ of kernels that had a broken bran layer; 35.7% of the wheat germ or 914.6 J per 100 kernels was consumed. Within two generations after initial introduction,C. ferrugineus reached a peak in numbers and biomass polluting the ecosystem with excreta and remains, and accelerating the deteriorative process observed in the insect-free control system by increasing respiration temperature, FAV and reducing grain germination. After 87 d, the insect population declined to low levels. The simulation model provided a close match between the observed and predicted numbers of insect life stages and bioenergetic variables during the insect population growth phase. Simulation trials suggested that cannibalism of larger compared with smaller immature stages would be more wasteful of developmental time and energy, reducing the number of individuals reaching reproductive age, and that density-dependent fecundity was probably not an important regulatory mechanism ofC. ferrugineus population dynamics in this study. Contribution No. 1314 from Agriculture Canada Research Station, Winnipeg, Manitoba, R3T 2M9, Canada     

Continuous population estimates forDendroctonus frontalis Zimm. (Coleoptera: Scolytidae) occurring in infestations

Summary Infestations ofDendroctonus frontalis Zimm. are often observed to enlarge continuously by the colonization of new hosts in a pattern similar to a forest fire. This pattern of infestation growth presents unique problems in quantitatively estimating populations ofD. frontalis. Beetle populations on each infested tree in an infestation go through five processes: attack, oviposition, reemergence, survivorship, and emergence. These processes, which have been described mathematically in the literature, each take several days for completion. In order to follow the distribution and abundance ofD. frontalis throughout the course of development of a spot, we need a daily estimate of the number of beetles involved in each process on every tree. Since it is not practical to sample each tree daily, we developed a procedure whereby quantitative estimation procedures for within-tree populations were used in combination with the mathematical models for the life processes to produce a daily record of the number of adults successfully attacking trees, the number of eggs oviposited, the number of beetles reemerging, number of beetles surviving within the trees, and the number of beetles emerging. These daily estimates were then summarized for all trees in the spot for the duration of the infestation. The daily record of populations ofD. frontalis, used with information on infestation geometry, were suggested to be of value in describing and elucidating several important facets of population dynamics including dispersal patterns within infestations, between tree beetle loss (mortality), and time lags among the various population processes. The information reported can be used to develop simulation models of population dynamics or to validate existing models. Texas Agric. Experiment Stn. TA No. 14689.     

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.     

Application of a marking-and-recapture method for the analysis of larval-adult populations of an insect,Nezara viridula (Hemiptera: Pentatomidae)

Summary Difficulty arises in applying marking-and-recapture methods to insects when the probability of recapture of marked individuals is changed with advancing age, either due to detachment of the mark by moulting (in the case of larvae) or to changes in their survival rate or their behaviour. A modification of the re-recapture method (Leslie et al., 1953) has been devised to analyze the capture-recapture data of the 5th-instar larvae and adults ofNezara viridula L. Estimation of the rate of moulting to the adult stage is made with the aid of additional information on larval survival. Migration rates of the larvae between the two halves of the census field is estimated byIwao's (1963) method. Through these analyses, the dynamic feature of the population during transition from the 5th instar to, adult is revealed. Several problems involved in the application of marking-and-recapture methods to insect populations are discussed. Contribution from the Entomological Laboratory, Kyoto University No. 392.