A Monte Carlo simulation model for studying evolution in age-structured populations

This model provides for any number of genotypes defined by age-specific survival and fecundity rates in a population with completely overlapping generations and growing under the control of density-governing functions affecting survival or fecundity. It is tested in situations involving two alleles at one locus. Nonselection populations at Hardy–Weinberg equilibrium obey the ecogenetic law; i.e., each genotype follows Lotka's law regarding rate of increase and stable age distribution as if it were an independent true-breeding population. Populations experiencing age- and density-independent selection approximate this situation, and the changes in gene frequency are predicted by relative fitnesses bases on λ, the finite rate of increase of the genotypes. Polymorphic gene equilibria occurring at steady-state population sizes are determined by fitnesses based on R, the net reproductive rate. In examples involving differences in generation time produced by age-dependent differences in fecundity, the allele associated with longer generation time may be favored or opposed by selection, depending on whether the density-governing factor controlling population size affects survival or fecundity. If such genotypes have similar R's, a genetic equilibrium may be established if the population is governed by a density function acting upon fecundity. Received: August 23, 1999 / Accepted: July 13, 2000     

Effect of pH on survival,reproduction, egg viability and growth rate of five closely related rotifer species

In the laboratory we examined the effect of pH (5–10 with one interval) on survival, reproduction, egg viability and growth rate (intrinsic growth rate—r _m and population growth rate—r) of five Brachionus rotifer species (B. calyciflorus, B. quadridentatus, B. urceolaris, B. patulus and B. angularis). The pH was shown to exert a major influence on egg viability and growth rate (r _m and r) for each species. The age-specific survivorship curves within a species were not significantly different at pH 6–10. The optimal pH for each species is near-neutral pH (pH 6–8), and the fecundity decreased as the pH deviated from these values. For each Brachionus species, there was no significant difference between age-specific fecundity curves at pH 7 and 8. At acid pH (pH 5 or 6) higher egg mortality was observed for each species. The r _m and population r of the five Brachionus species incubated at different pHs were significantly influenced by pH. The pH supporting the highest r _m or r was obtained at pH 6–8, but varied due to species. In this study B. urceolaris and B. patulus could tolerate a broad range of pH, while the populations of B. calyciflorus, B. quadridentatus and B. angulari declined at acid conditions.     

Effects of soybean proteinase inhibitor on development, survival and reproductive potential of the sugarcane borer, Diatraea saccharalis

One approach that can be employed in integrated pest management is the use of proteins with antinutritional effects on insect metabolism and development. The antimetabolic properties of soybean proteinase inhibitor (SPI) on growth of neonate larvae of the sugarcane borer, Diatraea saccharalis (Fabricius, 1794) (Lepidoptera: Crambidae) have been evaluated. When incorporated into an artificial diet at 0.5% (w/w), SPI retarded growth rate and development of larvae when compared with larvae fed on artificial diet alone. However, larval survival was not significantly affected. The purpose of our research was to calculate demographic statistics for the sugarcane borer reared on diet either with or without semi-purified extract of SPI. Net reproductive rate (R ₀), instantaneous rate of increase (r _m), combined age-specific survivorship (l _x) and age specific fecundity (m _x) provide information about population growth potential. These parameters were measured in order to determine the effects of the proteinase inhibitor on the insect's population dynamics. The observed differences would potentially translate into large reductions in population growth, indicating a potential value of using SPI for protecting sugarcane plants against damage by the sugarcane borer.     

Fecundity,reproductive rate,longevity, and intrinsic rate of increase of an aphidiid parasitoidLysiphlebia mirzai

A life-table was constructed for a little known aphidiid waspLysiphlebia mirzai, a parasitoid of cereal aphid,Rhopalosiphum maidis. The female parasitoid survived 6.4 ± 1.17 (SD) days and oviposited intensively 4.0 ± 0.47 days. The total fecundity rate, R_t, was 169.2 ± 6.94 mummies/female and net reproductive rate, R_o, was 92.70 female offspring/female. The intrinsic total fecundity rate, r_t, and intrinsic rate of natural increase, r_m, the finite rate of total fecundity, λ^t, and finite rate of increase, λ^m, was 0.27048, 0.24155, 1.31059 and 1.27322 respectively. The mean generation time (18.75 days) and doubling time of the population (2.87 days) was slightly higher than other aphidiids studied so far. The proportion of female progenies decreased significantly on the successive oviposition days.      

LIFE-CYCLE COMPONENTS OF SELECTION IN ERIGERON ANNUUS: II. GENETIC VARIATION

Genetic variation for seedling and adult fitness components was measured under natural conditions to determine the relative importance of the seedling stage for lifetime fitness in Erigeron annuus. Variation in lifetime reproductive success can result from both the persistent effects of genetic variation expressed among seedlings and from variation in adult fitness components. Analysis of covariance was used to separate the stage specific from the cumulative effects of genetic variance expressed earlier in the life cycle. E. annuus produces seeds through apomixis, which allowed measurement of the fitness of replicate genotypes from germination through the entire life cycle. There were significant differences among genotypes for date of emergence, seedling size, survivorship and fecundity, but heritabilities were low, indicating slow response to selection. For all characters, environmental components of variance were one to two orders of magnitude larger than genetic variance components, resulting in broad sense heritabilities less than 0.1. For seedling size and fecundity, all of the genetic variance was in the form of genotype-environment interactions, often with large negative genetic correlations across environments. In contrast, genotypes differed in mean survivorship through one year, but there were no genotype-environment interactions for viability. Genetic differences in viability were primarily expressed as differences in overwinter survivorship. Genotype × environment interactions among sites and blocks were generated early in the life cycle while the genotype × environment interactions in response to competitive environment (open, annual cover, perennial cover) first appeared in adult fecundity. Genetic variation in lifetime fitness was not significant, despite a fourfold difference in mean fitness among genotypes.     

Effects of hydroxamic acids on the resistance of wheat to the aphid Sitobion avenae

Resistance to the aphid Sitobion avenae F. was assessed on six wheat cultivars of known concentrations of total hydroxamic acids. A range of values for intrinsic rate of increase (r_m) were obtained, 96% of the variation in which was explained by acid concentrations. Of the components of r_m, age-specific fecundity was the most significant factor contributing to resistance.     

Comparative life table analysis of Tetranychus urticae (Acari: Tetranychidae) on 14 soybean genotypes

Abstract The life table parameters of two‐spotted spider mite (TSSM), Tetranychus urticae Koch, on 14 soybean genotypes (Sari, Sahar, Tellar, Zane, Ks3494, L₁₇, Dpx, 032, 033, Clark, Hill, I27, Tms and Williams) were evaluated at 28 ± 1°C, 65%± 5% RH and a photoperiod of 16 : 8 h L : D. The survivorship data (l_x) of adult females were fitted to the Weibull frequency distribution model in all genotypes tested. Based on the Weibull parameters, the survival curve was type I on all genotypes, which indicated that the mortality mostly occurred in old individuals. Furthermore, two mathematical models (Analytis and Enkegaard) were fitted to age‐specific fecundity data (m_x). The highest intrinsic rate of increase (r_m) was recorded on L₁₇ (0.392) and the lowest values of this parameter were obtained on Tms (0.233), Hill (0.236), 032 (0.236), I27 (0.237) and Clark (0.240). In addition, net reproductive rate (R₀) and finite rate of increase (λ) of the TSSM had the highest value on L₁₇ as 45.521 and 1.475, respectively. The lowest values of these parameters were recorded on Tms as 12.149 and 1.258, respectively. Doubling time (DT) varied significantly on different genotypes and the shortest and longest values of this period were obtained on L₁₇ and 032, respectively. Our findings revealed that Tms, Hill, 032, I27 and Clark were less suitable genotypes, suggesting that they are more resistant to the TSSM than the other genotypes.     

Variation in selected life-history parameters of the parasitoid wasp,Aphidius ervi: Influence of host developmental stage

We determined the age-specific fecundity and survival of the solitary parasstoid wasp,Aphidius ervi Haliday (Hymenoptera: Aphidiidae), under constant laboratory conditions. Wasps were reared in each of the four nymphal instars of apterous pea aphids,Acyrthosiphon pisum (Harris) (Homoptera: Aphididae): L₁ (age 24h), L₂ (48 h), L₃ (72 h), and L₄ (120 h). Age-specific survival (l_x) and fecundity (m_x) differed between parasitoids developing in different aphid instars. The wasps’ life-time reproductive success, as indexed by the intrinsic rate of population increase (r_m), varied non-linearly with adult biomass and host size at parasitization. A close agreement between larval growth rates in different host instars and adult reproductive performance suggests that, inA. ervi, fitness correlates may be significantly influenced by larval ontogeny and trade-offs in resource allocation.     

Effect of humidity on the development and fecundity of <Emphasis Type="Italic">Ophraella communa</Emphasis> (Coleoptera: Chrysomelidae)

Ophraella communa, an unintentionally introduced leaf beetle in China, has good control efficiency on ragweed, Ambrosia artemisiifolia. Aspects of the climatic requirements for development, survival, longevity and fecundity of O. communa were studied under the conditions of constant temperature (25 ± 1°C), photoperiod of 14 L:10 D and three relative humidities (60%, 75% and 90% RHs). The results showed that the developmental periods of O. communa at different stages shortened along with the increasing relative humidity, except that of the pupal stage. Although no differences were observed in the pupal survival rate, ovipositional period, fecundity, longevity and adult female age-specific survivorship of O. communa under the three humidity conditions, the survival rates during the egg, larva and entire immature stage were significantly higher at 75% RH and 90% RH than at 60% RH. The innate rate of increase (r _m), net reproductive rate (R ₀), finite rate of increase (λ) reached the maximum at 75% RH, with values of 0.181, 1116.4 and 1.198, respectively. These results indicated that the optimum relative humidity for the development of O. communa ranged from 75% RH to 90% RH. Thus O. communa prefers moist microclimate habitats. Its population may expand rapidly during mid-May to late August in south, east and central China, when the humidity is relatively high.     

Interactions of two natural enemies of Tetranychus urticae,the fungal entomopathogen Beauveria bassiana and the predatory mite,Phytoseiulus persimilis

The effect of either untreated or treated adults of the spider mite, Tetranychus urticae Koch (Acari: Tetranychidae) by Beauveria bassiana (Bals.) Vuillemin (Ascomycota: Hypocreales) DEBI008 at 1×10⁶ (conidia/ml) was investigated on developmental stages and life table parameters of Phytoseiulus persimilis Athias-Henriot (Acari: Phytoseiidae) under laboratory conditions. Four time intervals (0, 24, 48 and 72 h post-inoculation of spider mites) were considered for studying the predator characteristics as different treatments. Duration of each life stage, longevity, reproduction rate, intrinsic rate of natural increase (r _m ), net reproductive rate (R ₀), mean generation time (T) and finite rate of increase (λ) of the P. persimilis were calculated on both untreated and B. bassiana treated spider mite adults. Data analysis showed that longevity and fecundity of predatory mites fed on untreated and treated mites (time interval 0) were higher in comparison with other time intervals after inoculation. The entomopathogenic fungus adversely affected longevity and fecundity of the predatory mite. Fertility life table parameters of predatory mites fed on T. urticae treated by B. bassiana at different time intervals showed that T, R ₀, λ and r _m are strongly affected by the fungus presence and these parameters had significant differences among time treatments. The least r _m value was observed in the time interval of 72 h post-inoculation. The fitness of T. urticae was affected by B. bassiana 24, 48 and 72 h post-inoculation of mite adults, and consequently it caused decreased longevity of P. persimilis and accordingly a decrease in the intrinsic rate of natural increase of the predator.     

Sublethal effects of spirodiclofen on life history and life-table parameters of two-spotted spider mite (<Emphasis Type="Italic">Tetranychus urticae</Emphasis>)

Laboratory bioassays were conducted to evaluate the effects of spirodiclofen on life history and life-table parameters of two-spotted spider mite (Tetranychus urticae Koch) females treated at pre-ovipositional period with a series of acaricide concentrations starting with the concentration discriminative for eggs and immatures i.e. the lowest concentration that causes 100% mortality of those stages. After a 24 h exposure, the proportion of females that survived treatments was 0.71 (6 mg/l), 0.51 (12 mg/l), 0.41 (24 mg/l), 0.30 (48 mg/l) and 0.25 (96 mg/l). At the end of the trial, the survival rate of females treated with the lowest concentration was significantly lower than the survival rate of untreated females but it remained above that of females treated with higher concentrations. Total fecundity/fertility significantly decreased as concentrations of spirodiclofen increased. Viable eggs were laid by females treated with 6, 12 and 24 mg/l, and total fertility was reduced by 42, 84 and 97%, respectively. Compared with control, the gross fecundity/fertility of the treated females was significantly reduced throughout the trial, except in females treated with 6 mg/l. All concentrations caused a significant reduction in the net fecundity/fertility throughout the trial. The females treated with 12 mg/l had significantly reduced net reproductive rate (R ₀ = 6.45), compared to females treated with 6 mg/l (R ₀ = 23.35) and to untreated females (R ₀ = 28.92); there was no significant difference between the last two treatments. The intrinsic rate of increase (r _m ) and finite rate of increase (λ) were significantly reduced in treated females (r _m = 0.141, λ = 1.156 and r _m = 0.214, λ = 1.232; 12 and 6 mg/l, respectively), compared to control (r _m = 0.251, λ = 1.276). The reduction was significantly greater in females treated with the highest concentration. As a result of the lowered r _m , the doubling time in treated females was significantly extended. Sublethal effects of spirodiclofen and its impact on T. urticae management are discussed.     

Reproduction,adult survival and intrinsic rate of growth ofUrolepis rufipes [Hymenoptera: Pteromalidae], a pupal parasitoid of house flies,Musca domestica

Urolepis rufipes Ashmead, a recently discovered parasitoid of house flies at New York dairies, was reared at 15, 20, 25, 30 and 34°C to measure daily fertility, fecundity and adult survivorship. Little reproduction occurred at 15°C, and only a few ♀ successfully emerged at 34°C. The intrinsic rate of growth was fastest at 30°C (0.282 ♀/day), but fecundity was highest at 25°C (165.5 hosts attacked, producing 124.5 progeny). Some reproductive statistics at 25°C were: net reproductive rate (R₀=72.1 ♀/♀, generation time =18.7 days, intrinsic rate of increase (r_m)=0.228, finite rate of increase (λ)=1.26, daily birth rate =0.302, daily death rate =0.021 and Fisher's reproductive value =418. Sex ratio (average =75.9%) did not vary significantly with temperature (between 20–30°C) nor with mother's age.      

GENETIC CONSTRAINTS TO LIFE-HISTORY EVOLUTION IN THE PITCHER-PLANT MOSQUITO,WYEOMYIA SMITHII

Life-history theory relies heavily on the hypothesis that genetic tradeoffs among the components of fitness constrain their independent evolution and joint maximization. Herein we show that selection on preadult development time in the pitcher-plant mosquito, Wyeomyia smithii, leads to a correlated response in cohort mean generation time but no correlated response in survivorship, fecundity, or cohort replacement rate. Lines selected for fast development achieve a higher capacity for increase (r_c) than lines selected for slow development, independently of larval density. These results imply that tradeoffs due to underlying antagonistic pleiotropy affecting growth, development, survivorship, and reproduction are not necessary constraints to life-history evolution. Previous work with W. smithii has shown a positive genetic correlation between development time and a general, genetically coordinated diapause syndrome. We propose that the observed nontradeoffs among the components of r_c may be subsumed into an even more fundamental tradeoff between performance during the summer generations and synchronization of development and reproduction with the changing seasons. Consequently, critical tests of genetic tradeoffs as a constraint to the independent evolution or simultaneous optimization of fitness components may need to consider the seasonal context.     

r‐ and K‐selection in fluctuating populations is determined by the evolutionary trade‐off between two fitness measures: Growth rate and lifetime reproductive success

In a stable environment, evolution maximizes growth rates in populations that are not density regulated and the carrying capacity in the case of density regulation. In a fluctuating environment, evolution maximizes a function of growth rate, carrying capacity and environmental variance, tending to r‐selection and K‐selection under large and small environmental noise, respectively. Here we analyze a model in which birth and death rates depend on density through the same function but with independent strength of density dependence. As a special case, both functions may be linear, corresponding to logistic dynamics. It is shown that evolution maximizes a function of the deterministic growth rate r₀ and the lifetime reproductive success (LRS) R₀, both defined at small densities, as well as the environmental variance. Under large noise this function is dominated by r₀ and average lifetimes are small, whereas R₀ dominates and lifetimes are larger under small noise. Thus, K‐selection is closely linked to selection for large R₀ so that evolution tends to maximize LRS in a stable environment. Consequently, different quantities (r₀ and R₀) tend to be maximized at low and high densities, respectively, favoring density‐dependent changes in the optimal life history.     

Melon aphid performance on chrysanthemum as mediated by cultivar, and differential levels of fertilization and irrigation

Population parameters of the melon aphid, Aphis gossypii Glover, were compared for insects grown on six cultivars of chrysanthemum, Dendranthema grandiflora Tzvelev ('Fontana', 'Iridon', 'Pink Lady', 'Splendor', 'White Diamond', and 'White View Time') fertilized with 3 different levels of nitrogen (80, 160 and 240 mg N/l) and 2 levels of irrigation (high 300 ml and low 210 ml every other day). Fecundity, longevity, and survivorship of aphids were significantly affected by the cultivar treatment, but were not affected by irrigation or fertilizer treatments or by any combination of interactions among cultivar, fertilizer and irrigation. Significantly fewer aphids survived on the cultivar 'Pink Lady' (populations reduced by 20.8%) than any other cultivar examined. Intrinsic rates of increase r _m , finite rates of increase (R ₀), and population doubling times (DT) for aphids were different among fertilizer levels and cultivars, but were not different among irrigation levels. There were no significant fertilizer by irrigation interaction effects upon r _m , R ₀, or DT. The highest rate of fertilizer applied (240 mg N/liter) adversely affected the intrinsic rate of increase of melon aphids; however, aphid intrinsic rate of increase was weakly correlated with foliar soluble protein levels (r=0.989, P=0.0954). Foliar soluble protein levels were not associated with fertilization treatment. Melon aphids perform consistently better on the cultivars 'White Diamond', 'Fontana', and 'Splendor', relative to those aphids growing on 'Iridon', 'Pink Lady', or 'White View Time'. In general, melon aphids performed best on 'White Diamond' and poorest on 'Pink Lady'.     

Mhc‐linked survival and lifetime reproductive success in a wild population of great tits

Major histocompatibility complex (Mhc) genes are frequently used as a model for adaptive genetic diversity. Although associations between Mhc and disease resistance are frequently documented, little is known about the fitness consequences of Mhc variation in wild populations. Further, most work to date has involved testing associations between Mhc genotypes and fitness components. However, the functional diversity of the Mhc, and hence the mechanism by which selection on Mhc acts, depends on how genotypes map to the functional properties of Mhc molecules. Here, we test three hypotheses that relate Mhc diversity to fitness: (i) the maximal diversity hypothesis, (ii) the optimal diversity hypothesis and (iii) effect of specific Mhc types. We combine mark–recapture methods with analysis of long‐term breeding data to investigate the effects of Mhc class I functional diversity (Mhc supertypes) on individual fitness in a wild great tit (Parus major) population. We found that the presence of three different Mhc supertypes was associated with three different components of individual fitness: survival, annual recruitment and lifetime reproductive success (LRS). Great tits possessing Mhc supertype 3 experienced higher survival rates than those that did not, whereas individuals with Mhc supertype 6 experienced higher LRS and were more likely to recruit offspring each year. Conversely, great tits that possessed Mhc supertype 5 had reduced LRS. We found no evidence for a selective advantage of Mhc diversity, in terms of either maximal or optimal supertype diversity. Our results support the suggestion that specific Mhc types are an important determinant of individual fitness.     

Fitness comparison between clones differing in their ability to produce sexuals in the aphid Thopalosiphum padi

This study aimed at evaluating the intrinsic rate of increase (r _m ) of clones of the cereal aphid Rhopalosiphum padi (L.) that differ in their ability to produce sexuals. The value of r _m was measured for wingless parthenogenetic females in experiments conducted at two temperatures (15 °C and 20 °C). We studied six holocyclic clones, five androcyclic clones and five anholocyclic clones and showed that life-cycle has no significant effect on the age at first reproduction, on fecundity nor on longevity. As a result there is no influence on (summer) fitness: on the one hand, holocyclic clones do not compensate their lower fitness in mild winters by a higher fitness during the summer. On the other hand, anholocyclic and androcyclic clones do not seem to obtain any advantage from living on a more restricted range of hosts: there appears to be no cost of polyphenism for holocyclic clones.     

Resistance and susceptibility of some wheat lines to bird cherry-oat aphid <Emphasis Type="Italic">Rhopalosiphum padi</Emphasis> (Hemiptera: Aphididae)

Bird cherry-oat aphid, Rhopalosiphum padi (L.), is one of the most serious pests of cereals, with an almost worldwide distribution. A primary screening test was carried out to gauge the resistance or susceptibility of 40 wheat lines to R. padi, and follow-up experiments were conducted to determine the development and fecundity rates of R. padi on eight lines. The forty wheat lines examined were classified into two major classes and four subclasses: A (e.g., ERWYT 87-1) and B (e.g., ERWYT 87-20, ERWYT 87-11), with the highest average number of aphids 14 days after infestation, subclass C (e.g., ERWYT 88-8, ERWYT 87-6, ERWYT 87-4), with the lowest average number of aphids, and subclass D (e.g., ERWYT 88-12, ERWYT 88-13), with intermediate status. Aphid nymph developmental time and survival both differed among the wheat lines. Differences were also observed in the fecundity of R. padi, the intrinsic rate of increase (r _m), and other parameters among the wheat lines tested. The highest and lowest values of r _m were obtained for ERWYT 87-1 and ERWYT 88-8 (0.36 and 0.26 nymphs/female/day, respectively). Three lines (ERWYT 87-4, ERWYT 87-6, ERWYT 88-8) were relatively resistant to R. padi, which could prove useful in the development of IPM programs for this aphid in wheat fields.     

Resistance in tomato to the greenhouse whitefly: analysis of population dynamics

Life history parameters including longevity, developmental time, and reproduction were determined for whiteflies, Trialeurodes vaporariorum (Westwood), kept in clip-on cages on susceptible parent, Lycopersicon esculentum, resistant parent, L. hirsutum glabratum, and intermediate breeding lines of tomato. Using the Lewontin triangular reproductive function, the intrinsic rate of increase (r_m) was calculated for each genotype. This is an elegant tool for detecting resistance, as it incorporates all salient factors of life history. Comparing r_m to the other parameters measured, selection criteria were chosen. Developmental time is not a suitable selection criterion as it varies very little even between the most susceptible and the most resistant genotypes. Additionally, developmental time of T. vaporariorum offspring was found to be highly significantly correlated to parental age on all tomato genotypes except L. hirsutum glabratum. Total reproduction, truncated population counts, and longevity appear good criteria for selection. This test, focusing on antibiotic factors, shows large differences between the resistant and susceptible parent genotypes, but much smaller differences between the breeding lines and the susceptible parent. Earlier screenings relying on a variety of tests appear to have maintained antixenotic rather than antibiotic properties in the breeding lines.     

Antibiosis resistance to cereal leaf beetle,Oulema melanopus (L.) (Coleoptera: Chrysomelidae), in Central Asian wheat germplasm

The cereal leaf beetle, Oulema melanopus (Coleoptera: Chrysomelidae), is a new invasive pest of cereals in western Canada. Host plant resistance is a potential option in the integrated management of this insect. Wheat (Triticum aestivum L.) genotypes are known to possess antixenosis against O. melanopus; however, mechanisms involving antibiosis are relatively less explored. We present results of an investigation exploring antibiotic resistance in wheat germplasm of Central Asian origin with putative resistance to O. melanopus. Our laboratory assessment of development and survivorship of O. melanopus on test genotypes indicated antibiotic properties in four of the six genotypes evaluated. Antibiosis effects were reflected through lower survivorship, extended developmental periods and low adult fitness. However, two genotypes were excellent hosts, and beetles had higher survivorship and fitness on them. The performance of O. melanopus on these two genotypes was comparable to that on the susceptible genotype CDC GO included as a check. The genotypes with antibiotic resistance identified by this research can now serve as sources of genetic resistance for breeding cultivars adapted to western Canadian conditions.