Estimating developmental and mortality rates and stage recruitment from insect stage-frequency data

A model for the analysis of insect stage-frequency data is developed which includes stage-specific variable developmental periods and stage-specific daily survival rates. The model can predict the development of an insect population through its developmental stages and consequently may form the basis for a simulation model of the population.     

Extensions to Kiritani and Nakasuji’s method for analysing insect stage-frequency data

Summary There is evidence that the simple method ofKiritani andNakasuji (1967) for the analysis of insect stage-frequency data produces estimates of stage-specific survival rates that compare well with the estimates from more complicated methods (Manly, 1975). However the method as originally proposed byKiritani andNakasuji cannot always be applied because it assumes that the insect population involved was sampled at regular intervals of time. Furthermore, parameters such as the durations of stages are not estimated. In this note modifications to the basic method are suggested with the idea of overcoming these limitations.     

Estimation of the individual number entering each development stage in an insect population

A method for estimating the number entering each development stage from data obtained by regular sampling through one generation of an insect population was described. This method is consisted of the following two procedures: The provisional estimates are calculated on the assumption that each stage has a common mortality in a sampling interval. Then these estimates are corrected on another assumption that the mortality is different in each stage but constant during a stage. The result of testing its validity with two laboratory populations of the common cabbage butterfly, Pieris rapae crucivora, showed the availability of the present method.     

A multiple regression method for analysing stage-frequency data

A linear regression method that allows survival rates to vary from stage to stage is described for the analysis of stage-frequency data. It has advantages over previously suggested methods since the calculations are not iterative, and it is not necessary to have independent estimates of stage durations, numbers entering stages, or the rate of entry to stage 1. Simulation is proposed to determine standard errors for estimates of population parameters, and to assess the goodness of fit of models.     

Note on a method for analyzing stage-frequency data

A new method for analyzing stage-frequency data is proposed which is based on the estimation of rates of transition between one stage and the next highest stage in one unit of time, and a unit time survival rate that is assumed to be constant. Once these estimates are calculated it becomes possible to also estimate the mean durations of stages, stage-specific survival rates, and numbers entering stages. An advantage of the method is that it can be applied with any distribution of entry times to stage 1, and any distribution of numbers in stages when sampling begins. Use of the method is illustrated on data from a copepod population in a Canadian lake.     

A method for the estimation of parameters for natural stage-structured populations

A relatively simple method is proposed for the estimation of parameters of stage-structured populations from sample data for situation where (a) unit time survival rates may vary with time, and (b) the distribution of entry times to stage 1 is too complicated to be fitted with a simple parametric model such as a normal or gamma distribution. The key aspects of this model are that the entry time distribution is approximated by an exponential function withp parameters, the unit time survival rates in stages are approximated by anr parameter exponential polynomial in the stage number, and the durations of stages are assumed to be the same for all individuals. The new method is applied to four Zooplankton data sets, with parametric bootstrapping used to assess the bias and variation in estimates. It is concluded that good estimates of demographic parameters from stagefrequency data from natural populations will usually only be possible if extra information such as the durations of stages is known.     

Differential mortality in Turkana agriculturalists and pastoralists

Nomadic pastoral populations appear to have much lower rates of growth than the otherwise very high growth rates now characteristic of populations in developing nations. Because dramatic declines in infant mortality have been a primary contributor to increased population growth rates in these countries, it has been assumed that nomadic pastoral populations are still characterized by high levels of mortality in the first few years of life. Few studies, however, have been undertaken to estimate demographic parameters for nomadic pastoral populations, and even fewer of a comparative nature have been undertaken to document the impact of subsistence strategy on demographic processes. This study compares indirect childhood mortality estimates for Turkana nomadic pastoralists with childhood mortality in a settled agricultural group within the same population and finds that pastoralists have substantially higher levels of mortality. Based on the childhood mortality estimates, model life tables are selected for pastoral and agricultural groups from which values for mean life expectancy and infant mortality are estimated and compared. Recent improvements in primary health care for the settled agricultural group are ruled out as being an important cause of their lower mortality levels, and some aspects of life-style associated with subsistence strategy are discussed as likely determinants of the mortality differences.     

Field population biology of citrus whitefly, Dialeurodes citri (Ashmead) (Heteroptera: Aleyrodidae), on oranges in California

Populations of Dialeurodes citri (Ashmead) on oranges were studied in two orange orchards in southern California, USA, to quantify developmental times and mortalities affecting this species. Four populations were followed in detail for three generations over 2 years, by monitoring the development and survival of 200 individuals in each population. Overall, preimaginal mortality among the generation studied varied from 56 to 94%. No evidence of disease or parasitism was present in the populations, and mortality was likely due to generalist predators. Weather records for the period were typical for the region, and periods of rainfall or elevated temperatures did not show any correlation with periods of mortality in the populations. Overall, the populations have high survival rates for whitefly populations, and this high survival is likely a major contributing factor to their pest status in California orchards. Addition of natural enemies to reduce the survival rates of these populations is proposed as the next course of action in suppressing populations below damaging levels.     

Further improvements to a method for analysing stage-frequency data

An iterative procedure for correcting stage-frequency data is described to allow for situations where the period during which a population is sampled begins after some individuals have entered stage 2 or ends before all individuals are dead. The reason for correcting data in this way is to enableKiritani andNakasuji's method for estimating stage-specific survival rates, with extensions proposed byManly (1976, 1977), to be used to analyse the data. The proposed procedure is illustrated on data obtained by sampling a population of the grasshopper Chorthippus brunneus passing through four instar stages to reach the adult stage.     

Impact of developmental variance on behavior of models for insect populations I. Models for populations with unrestricted growth

The concept of developmental variance is discussed with reference to its use in models for insect populations. When included in a model, developmental variance is typically used to describe the variation of developmental periods among individuals. However, its presence in a model can also have indirect impact on survival and fertility schedules. This impact can lead to significant changes in population growth rates and generation times. These relationships between developmental variance and population growth in models are quantified and discussed.     

The relationship between conidial dose,moulting and insect developmental stage on the susceptibility of cotton aphid,Aphis gossypii,to conidia of Lecanicillium attenuatum,an entomopathogenic fungus

The cotton aphid is one of the most serious pests of greenhouse vegetable crops worldwide. It is difficult to control because field populations usually include simultaneously several insect developmental stages. The current research evaluated an isolate (CS625) of Lecanicillium attenuatum, a fungal pathogen of aphids, as to its virulence against different developmental stages of cotton aphid, Aphis gossypii. The influence on mortality of several other factors also was examined: (a) insect moulting, (b) the number of conidia attached to insect cuticles and (c) germination rates of conidia on cuticles of aphids at various developmental stages. Mortality of cotton aphids treated with L. attenuatum conidia varied according to the developmental stage of the host, i.e. the LT_50s with third-instar nymphs and adults was shorter than with first-instar nymphs. The number of spores attached to the surface of first-instar nymphs was approximately one-half of that on third-instar nymphs and adults. Also, the level of spore germination on the surface of first-instar nymphs was lower than on the surface of other stages of the aphid. After moulting, the numbers of conidia attached to new insect cuticles were less than on exuviae. These results suggest that early nymphal stages of cotton aphids may escape fungal disease due, at least in part, to a combination of three factors: low numbers of conidia attached to their cuticles; low levels of conidial germination and rapid ecdyses, which removed conidia before their germ tubes penetrated the host hemolymph.     

Do Insect Populations Die at Constant Rates as They Become Older? Contrasting Demographic Failure Kinetics with Respect to Temperature According to the Weibull Model

Temperature implies contrasting biological causes of demographic aging in poikilotherms. In this work, we used the reliability theory to describe the consistency of mortality with age in moth populations and to show that differentiation in hazard rates is related to extrinsic environmental causes such as temperature. Moreover, experiments that manipulate extrinsic mortality were used to distinguish temperature-related death rates and the pertinence of the Weibull aging model. The Newton-Raphson optimization method was applied to calculate parameters for small samples of ages at death by estimating the maximum likelihoods surfaces using scored gradient vectors and the Hessian matrix. The study reveals for the first time that the Weibull function is able to describe contrasting biological causes of demographic aging for moth populations maintained at different temperature regimes. We demonstrate that at favourable conditions the insect death rate accelerates as age advances, in contrast to the extreme temperatures in which each individual drifts toward death in a linear fashion and has a constant chance of passing away. Moreover, slope of hazard rates shifts towards a constant initial rate which is a pattern demonstrated by systems which are not wearing out (e.g. non-aging) since the failure, or death, is a random event independent of time. This finding may appear surprising, because, traditionally, it was mostly thought as rule that in aging population force of mortality increases exponentially until all individuals have died. Moreover, in relation to other studies, we have not observed any typical decelerating aging patterns at late life (mortality leveling-off), but rather, accelerated hazard rates at optimum temperatures and a stabilized increase at the extremes.In most cases, the increase in aging-related mortality was simulated reasonably well according to the Weibull survivorship model that is applied. Moreover, semi log- probability hazard rate model illustrations and maximum likelihoods may be usefully in defining periods of mortality leveling off and provide clear evidence that environmental variability may affect parameter estimates and insect population failure rate. From a reliability theory standpoint, failure rates vary according to a linear function of age at the extremes indicating that the life system (i.e., population) is able to eliminate earlier failure and/or to keep later failure rates constant. The applied model was able to identify the major correlates of extended longevity and to suggest new ideas for using demographic concepts in both basic and applied population biology and aging.     

The statistical analysis of insect phenology

We introduce two simple methods for the statistical comparison of the temporal pattern of life-cycle events between two populations. The methods are based on a translation of stage-frequency data into individual 'times in stage'. For example, if the stage-k individuals in a set of samples consist of three individuals counted at time t(1) and two counted at time t(2), the observed times in stage k would be (t(1), t(1), t(1), t(2), t(2)). Times in stage then can be compared between two populations by performing stage-specific t-tests or by testing for equality of regression lines of time versus stage between the two populations. Simulations show that our methods perform at close to the nominal level, have good power against a range of alternatives, and have much better operating characteristics than a widely-used phenology model from the literature.     

Optimization and accuracy of faecal pellet count estimates of population size: The case of European rabbits in extensive breeding nuclei

Cost-effective indices to estimate relative abundances of species are crucial for their management and conservation. As an example, population indices are needed to monitor extensive breeding nuclei used for translocating wild rabbit populations. Based on counts of faecal pellets from four high density rabbit nuclei in southwest Spain: (i) we assess the accuracy of this population index in high-density populations; and (ii) present a simulation approach to evaluate how the reduction of the counting effort affects accuracy of the population estimates. Our findings suggest that this method provides a valid estimate in high-density rabbit populations, and, notably, estimates would have not substantially changed after a reduction of 45–65% in the number of counted plots depending on variation on rabbit density between nuclei. We provide a framework that managers and other scientists could use to improve data collection of pellet counts in order to optimize their chances of detecting relative abundance estimates of rabbits. In addition, we present a R function to implement our approach that can easily be applied in a variety of monitoring programmes for other species based on count data. This will likely help to reduce field-effort in different studies without compromising population indices estimates.     

Estimation of survival rate and extinction probability for stage-structured populations with overlapping life stages

The development of methods providing reliable estimates of demographic parameters (e.g., survival rates, fecundity) for wild populations is essential to better understand the ecology and conservation requirements of individual species. A number of methods exist for estimating the demographics of stage-structured populations, but inherent mathematical complexity often limits their uptake by conservation practitioners. Estimating survival rates for pond-breeding amphibians is further complicated by their complex migratory and reproductive behaviours, often resulting in nonobservable states and successive cohorts of eggs and tadpoles. Here we used comprehensive data on 11 distinct breeding toad populations (Bufo calamita) to clarify and assess the suitability of a relatively simple method [the Kiritani–Nakasuji–Manly (KNM) method] to estimate the survival rates of stage-structured populations with overlapping life stages. The study shows that the KNM method is robust and provides realistic estimates of amphibian egg and larval survival rates for species in which breeding can occur as a single pulse or over a period of several weeks. The study also provides estimates of fecundity for seven distinct toad populations and indicates that it is essential to use reliable estimates of fecundity to limit the risk of under- or overestimating the survival rates when using the KNM method. Survival and fecundity rates for B. calamita populations were then used to define population matrices and make a limited exploration of their growth and viability. The findings of the study recently led to the implementation of practical conservation measures at the sites where populations were most vulnerable to extinction.     

Population fluctuations of the diamondback moth,Plutella xylostella (L.) on cabbages inBacillus thuringiensis sprayed and non sprayed plots and factors affecting within-generation survival of immatures

Summary The population fluctuations and within-generation survival of immatures stages of the diamondback moth,Plutella xylostella on summer cabbages, were examined in unsprayed plots in 1984 and 1985 and in plots sprayed with two formulations ofBacillus thuringiensis in 1985. There was two distincy generations per crop and no noticeable difference in population fluctuations were observed in unsprayed plots between years. There was distinct difference between unsprayed and sprayed plots in 1985, from the larval period onwards. However, the survivorship patterns in all plots in both years were a Type 2 based onDeevey's (1947) classification. Life table studies showed that there was essentially no difference in the mortality agents acting on each of the stages except for numerical differences in the within-generation mortality rates. The major mortality rates during the egg stage were the parasitoid,Trichogramma spp. and unknown factors including rainfall; in the larval 1 stage was unknown mortality; in the larval 2 stage was the parasitoid,Apanteles plutellae and during the pupal stage was unknown mortality and parasitoid,Diadromus subtilicornis. Adult mortality was determined for generation 2. It was relatively higher in the unsprayed plots compared to theB. thuringiensis sprayed plots. The contribution of abiotic factors such as rainfall and temperature, and biotic factors such as parasitoids and predators in determining within-generation population levels and the fluctuation of populations on cabbage were discussed.     

An instantaneous coalescent method insensitive to population structure

Conventional coalescent inferences of population history make the critical assumption that the population under examination is panmictic. However, most populations are structured. This complicates the prevailing coalescent analyses and sometimes leads to inaccurate estimates. To develop a coalescent method unhampered by population structure, we perform two analyses. First, we demonstrate that the coalescent probability of two randomly sampled alleles from the immediate preceding generation(one generation back)is independent of population structure. Second, motivated by this finding, we propose a new coalescent method: i-coalescent analysis. The i-coalescent analysis computes the instantaneous coalescent rate by using a phylogenetic tree of sampled alleles. Using simulated data, we broadly demonstrate the capability of i-coalescent analysis to accurately reconstruct population size dynamics of highly structured populations, although we find this method often requires larger sample sizes for structured populations than for panmictic populations. Overall, our results indicate i-coalescent analysis to be a useful tool, especially for the inference of population histories with intractable structure such as the developmental history of cell populations in the organs of complex organisms.     

Model life table for captive chimpanzees

Mortality statistics from three captive populations of chimpanzees (Pan troglodytes) were combined to generate standard model life tables for each sex in this species. The model is compared to an estimate of survivorship of a group of wild animals, and is applied to an incomplete data set to illustrate how the model may be used to extend estimates of mortality statistics to missing older ages. © 1995 Wiley-Liss, Inc.     

Coral reef fish populations can persist without immigration

Determining the conditions under which populations may persist requires accurate estimates of demographic parameters, including immigration, local reproductive success, and mortality rates. In marine populations, empirical estimates of these parameters are rare, due at least in part to the pelagic dispersal stage common to most marine organisms. Here, we evaluate population persistence and turnover for a population of orange clownfish, Amphiprion percula, at Kimbe Island in Papua New Guinea. All fish in the population were sampled and genotyped on five occasions at 2-year intervals spanning eight years. The genetic data enabled estimates of reproductive success retained in the same population (reproductive success to self-recruitment), reproductive success exported to other subpopulations (reproductive success to local connectivity), and immigration and mortality rates of sub-adults and adults. Approximately 50% of the recruits were assigned to parents from the Kimbe Island population and this was stable through the sampling period. Stability in the proportion of local and immigrant settlers is likely due to: low annual mortality rates and stable egg production rates, and the short larval stages and sensory capacities of reef fish larvae. Biannual mortality rates ranged from 0.09 to 0.55 and varied significantly spatially. We used these data to parametrize a model that estimated the probability of the Kimbe Island population persisting in the absence of immigration. The Kimbe Island population was found to persist without significant immigration. Model results suggest the island population persists because the largest of the subpopulations are maintained due to having low mortality and high self-recruitment rates. Our results enable managers to appropriately target and scale actions to maximize persistence likelihood as disturbance frequencies increase.     

Post-settlement emigration affects mortality estimates for two Bahamian wrasses

Understanding the dynamics of open marine populations is inherently complex, and this complexity has led to decades of debate regarding the relative importance of pre- versus post-settlement processes in structuring these populations. Movement between patches may be an important modifier of patterns established at settlement, yet local immigration and emigration have received less attention than other demographic rates. I examined loss rates from tagged populations of juvenile wrasses (yellowhead wrasse Halichoeres garnoti and bluehead wrasse Thalassoma bifasciatum) at two sites in the Bahamas. Assuming that all losses were due solely to mortality would have significantly underestimated survivorship of yellowhead wrasse by 29% and bluehead wrasse by 14%. On average, per capita mortality and emigration rates were higher for yellowhead than bluehead wrasse, but neither demographic rate differed between sites for either species. With respect to within-species density, bluehead wrasse mortality was density-dependent at the patch reef site, but mortality rates of yellowhead wrasse were consistently density-independent. Evaluating the effects of between-species density, yellowhead wrasse mortality increased with a decrease in bluehead wrasse density, but this effect was limited to the patch reef site. Emigration rates were not a function of either within-species or between-species density, but instead varied inversely with isolation distance. Numerous previous studies of coral-reef fish, conducted on patch reefs separated by only a few meters of sand and often using untagged fish, may have confounded losses due to emigration with those due to mortality. A better understanding of the factors affecting emigration in marine fishes is important to their effective management using spatial tools such as marine protected areas.