Evolution of pseudo-arrhenotoky

In arrhenotokous arthropods, males arise from unfertilized eggs. Hence, by controlling the fertilization process mothers can adjust the sex ratio in their offspring. In pseudo-arrhenotokous phytoseiid mites, however, males are haploid, but arise from fertilized eggs. The haploid state is achieved through elimination of the paternal chromosome set during embryonic development. It is shown in this paper that phytoseiid females can control the sex ratio in their offspring and that this control seems as flexible as in arrhenotokous arthropods. As predicted by current evolutionary theory of sex allocation, sex ratios approached half males/half females under random mating, whereas a female bias was observed under sib-mating. The importance of these results for understanding the adaptive significance of pseudo-arrhenotoky is discussed. It is suggested that arrhenotoky is selected for when there is a substantial risk to the females of remaining unmated. When this risk of becoming a wall-flower is low, pseudo-arrhenotoky may evolve because it retains the possibility to reinstal lost genetic information in the maternally derived chromosome by using the paternal chromosome as a template for DNA-repair. The retention of the diploid state in males during embryonic development may thus have certain advantages. It is argued that pseudo-arrhenotoky may be an adaptive genetic system under certain conditions, and not an unstable system that readily reverts to diploidy or evolves towards arrhenotoky or thelytoky.     

Balancing in- and out-breeding by the predatory mite <Emphasis Type="Italic">Phytoseiulus persimilis</Emphasis>

In- and out-breeding depressions are commonly observed phenomena in sexually reproducing organisms with a patchy distribution pattern, and spatial segmentation and/or isolation of groups. At the genetic level, inbreeding depression is due to increased homozygosity, whereas outbreeding depression is due to inferior genetic compatibility of mates. Optimal outbreeding theory suggests that intermediate levels of mate relatedness should provide for the highest fitness gains. Here, we assessed the fitness consequences of genetic relatedness between mates in plant-inhabiting predatory mites Phytoseiulus persimilis, which are obligatory sexually reproducing but haplo-diploid. Both females and males arise from fertilized eggs but males lose the paternal chromosome set during embryogenesis, dubbed pseudo-arrhenotoky. Phytoseiulus persimilis are highly efficacious in reducing crop-damaging spider mite populations and widely used in biological control. Using iso-female lines of two populations, from Sicily and Greece, we assessed the fecundity of females, and sex ratio of their offspring, that mated with either a sibling, a male from the same population or a male from the other population. Additionally, we recorded mating latency and duration. Females mating with a male from the same population produced more eggs, with a lower female bias, over a longer time than females mating with a sibling or with a male from the other population. Mating latency was unaffected by mate relatedness; mating duration was disproportionally long in sibling couples, likely indicating female reluctance to mate and sub-optimal spermatophore transfer. Our study provides a rare example of in- and out-breeding depression in a haplo-diploid arthropod, supporting the optimal outbreeding theory.     

Foraging time and spatial patterns of predation in experimental populations

Summary Responses of the predaceous mites Phytoseiulus persimilis, Typhlodromus (=Metaseiulus) occidentalis, and Amblyseius andersoni to spatial variation in egg density of the phytophagous mite, Tetranychus urticae, were studied in the laboratory.The oligophagous predator P. persimilis showed initially a direct density dependent foraging time allocation and variation in foraging time increased with prey density. With changes in prey density due to predation, predator foraging rates (per hour) decreased with time and density dependent foraging gradually became density independence, because P. persimilis continued to respond to initial prey density, instead of the changing prey density and distribution. The consequent spatial pattern of predation by P. persimilis was density independent, although slopes of predation rate-prey density regressions increased with time.Compared with P. persimilis, the narrowly polyphagous predator T. occidentalis responded relatively slowly to the the presence or absence of prey eggs but not to prey density: the mean and variation of foraging time spent in patches with prey did not differ with prey density, but was significantly greater in patches with prey eggs than in patches without eggs. Prey density and distribution changed only slightly due to predation and overall foraging rates remained more or less constant. The consequent spatial pattern of predation by T. occidentalis was inversely density dependent. As with P. persimilis, slopes of predation rate-prey density regressions increased with time (i.e. the inverse density dependence in T. occidentalis became weaker through time).The broadly polyphagous predator A. andersoni showed density independent foraging time allocation with variation independent of prey density. With changes in prey density over time due to prey depletion, overall foraging rates decreased. The consequent spatial pattern of predation by A. andersoni also changed through time; it initially was inversely density dependent, but soon became density independent.Overall, P. persimilis and T. occidentalis spent more time in prey patches than A. andersoni, suggesting that A. andersoni tended to spend more time moving outside patches. The overall predation rates and searching efficiency were higher in P. persimilis than in A. andersoni and T. occidentalis. Predator reproduction was highest in P. persimilis, lower in T. occidentalis and the lowest A. andersoni.The differences in response to prey distribution among the three predaceous species probably reflect the evolution of these species in environments with different patterns of prey distribution. The degree of polyphagy is a major determinant of the aggregative response, but other attributes such as handling time are also important in other aspects of phytoseiid foraging behavior (e.g. searching efficiency or predation rate).     

Occurrence of protandry and a female-biased sex-ratio in a sponge-associated water mite (Acari: Unionicolidae)

The sex ratio of aUnionicola crassipes-type mite from a pond in Alberta, Canada, was found to be significantly female-biased at the time of adult emergence. Male mites emerged earlier, but were later surpassed in number by the females. This protandry seems more likely to be due to the smaller size of the males and hence their faster developmental rate than to any mating advatage the male mites gain by early emergence. It is possible that group selection acting on the productivity of mite colonies with different proportions of low-sex-ratio producers has selected for female-bias. Water mites are not known to exhibit any atypical modes of reproduction (e.g.arrhenotoky), nor is it known whether pre-adult maleU. crassipes have a greater mortality rate than females, so the mechanism behind the skewed ratio is not known.     

Evolutionary stable sex ratios with non‐facultative male‐eggs first sex allocation in fig wasps

Sex allocation theory has long generated insights into the nature of natural selection. Classical models have elucidated causal phenomena such as local mate competition and inbreeding on the degree of female bias exhibited by various invertebrates. Typically, these models assume mothers facultatively adjust sex allocation using predictive cues of future offspring mating conditions. Here we relax this assumption by developing a sex allocation model for haplodiploid mothers experiencing local mate competition that lay a fixed number of male eggs first. Female egg number is determined by remaining oviposition sites or remaining eggs of the mother, depending on which is exhausted first. Our model includes parameters for variation in foundress number, patch size, fecundity and offspring mortality that allow us to generate secondary sex ratio predictions based on specific parameterizations for natural populations. Simulations show that: 1) in line with classical models, factors that increase sib‐mating result in mothers laying relatively more female eggs; 2) high offspring mortality leads to relatively more males as fertilization insurance; 3) unlike classical model predictions, sub‐optimal predictions, such as more males than females are possible. In addition, our model provides the first quantitative predictions for the expected number of males and females in a patch where typically only one mother utilizes a given patch. We parameterized the model with data obtained from seven species of southern African fig wasps to predict expected means and variances for numbers of male and female offspring for typical numbers of mothers utilizing a patch. These predictions were compared to secondary sex ratio data from single foundress patches, the most commonly encountered situation for these species. Our predictions matched both the observed number and variance of male and female offspring with a high degree of accuracy suggesting that facultative adjustment is not required to produce evolutionary stable sex ratios.     

EVOLUTION OF HAPLODIPLOIDY IN DERMANYSSINE MITES (ACARI: MESOSTIGMATA)

Haplodiploidy, a widespread phenomenon in which males are haploid and females are diploid, can be caused by a number of different underlying genetic systems. In the most common of these, arrhenotoky, males arise from unfertilized eggs, whereas females arise from fertilized eggs. In another system, pseudoarrhenotoky, males arise from fertilized eggs, but they eliminate the paternal genome at some point prior to spermatogenesis, with the consequence that they do not pass this genome to their offspring. In 1931 Schrader and Hughes-Schrader suggested that arrhenotoky arises through a series of stages involving pseudoarrhenotokous systems such as those found in many scale insects (Homoptera: Coccoidea), however, their hypothesis has been largely ignored. We have used a phylogenetic analysis of 751 base pairs of 28S rDNA from a group of mites (Mesostigmata: Dermanyssina) that contains arrhenotokous, pseudoarrhenotokous, and ancestrally diplodiploid members to test this hypothesis. Neighbor-joining, maximum-parsimony, and maximum-likelihood methods all indicate that the arrhenotokous members of this group form a clade that arose from a pseudoarrhenotokous ancestor, rather than directly from a diplodiploid one. This provides unequivocal support for the hypothesis of Schrader and Hughes-Schrader. The wider implications of this result for the evolution of uniparental genetic systems are discussed.     

Reproductive tables of predatory phytoseiid mites (<Emphasis Type="Italic">Phytoseiulus persimilis,Galendromus occidentalis</Emphasis>, and <Emphasis Type="Italic">Neoseiulus cucumeris</Emphasis>)

Feeding of predatory mites (Phytoseiulus persimilis, Galendromus occidentalis, and Neoseiulus cucumeris) on different life stages of Tetranychus atlanticus under optimal conditions was studied. Daily and total consumption of prey by predators and selection of prey of different life stages were studied for 5 days and for the entire feeding period. Average daily food consumption [number of individuals] for the entire life period of mature mite females constituted 0.43 females + 5.0 [nymphs and males] + 3.4 eggs of Tetranychus atlanticus in P. persimilis; 0.12 females + 3.70 [nymphs and males] + 3.10 eggs in G. occidentalis; and 0.19 females + 4.10 [nymphs and males] + 3.50 eggs in N. cucumeris. During the entire period of feeding, P. persimilis preferred large individuals and at the postembryonic stages selected prey to a greater extent than G. occidentalis and N. cucumeris (61.8 and 55.1%, respectively). The use of a 5-day express-method is possible for estimation of some biological characteristics of phytoseiids that previously consumed the same food for a long period. In other cases, analysis of characteristics for the entire life period is necessary.     

Insemination Capacity and Dispersal in Relation to Sex Allocation Decisions in Goniozus legneri (Hymenoptera: Bethylidae): Why Are There More Males in Larger Broods?

Models considering sex ratio optima under single foundress strict local mate competition predict that female bias will be reduced by stochasticity in sex allocation, developmental mortality of males and limited insemination capacity of males. In all three cases the number of males per brood is expected to increase with brood size. Sex ratio optima may also be less female biased when several mothers contribute offspring to local mating groups or if non‐local mating occurs between members of different broods; again more males are expected in larger broods. In the parasitoid wasp Goniozus legneri (Hymenoptera: Bethylidae), sex allocation has only a small stochastic component, developmental mortality is low and non‐siblings are unlikely to develop in the same brood. However, the number of males per brood increases with the size of the brood (produced by a single mother). We investigated the further possibilities of limited insemination capacity and non‐local mating using a naturalistic experimental protocol. We found that limited insemination capacity is an unlikely general explanation for the increase in number of males with brood size. All males and females dispersed from both mixed and single sex broods. Although most females in mixed sex broods mated prior to dispersal, these data suggest that non‐local mating is possible, for instance via male immigration to broods containing virgin females. This may influence sex ratio optima and account for the trend in male number.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Constrained sex allocation in a parasitoid due to variation in male quality

The theory of constrained sex allocation posits that when a fraction of females in a haplodiploid population go unmated and thus produce only male offspring, mated females will evolve to lay a female-biased sex ratio. I examined evidence for constrained sex ratio evolution in the parasitic hymenopteran Uscana semifumipennis. Mated females in the laboratory produced more female-biased sex ratios than the sex ratio of adults hatching from field-collected eggs, consistent with constrained sex allocation theory. However, the male with whom a female mated affected her offspring sex ratio, even when sperm was successfully transferred, suggesting that constrained sex ratios can occur even in populations where all females succeed in mating. A positive relationship between sex ratio and fecundity indicates that females may become sperm-limited. Variation among males occurred even at low fecundity, however, suggesting that other factors may also be involved. Further, a quantitative genetic experiment found significant additive genetic variance in the population for the sex ratio of offspring produced by females. This has only rarely been demonstrated in a natural population of parasitoids, but is a necessary condition for sex ratio evolution. Finally, matings with larger males produced more female-biased offspring sex-ratios, suggesting positive selection on male size. Because the great majority of parasitic hymenoptera are monandrous, the finding of natural variation among males in their capacity to fertilize offspring, even after mating successfully, suggests that females may often be constrained in the sex allocation by inadequate number or quality of sperm transferred.     

Process of Egg Formation in the Female Body Cavity and Fertilization in Male Eggs of Phytoseiulus Persimilis (Acari: Phytoseiidae)

The process of egg formation in the body cavity of a phytoseiid mite, Phytoseiulus persimilis, was observed to examine fertilization of male eggs. After insemination, one of the ova at the periphery of the ovary began to expand, taking up yolk. Two pronuclei appeared in the expanded egg, located dorsally in the ovary, and yolk granules were formed gradually. After the egg became filled with yolk granules the two pronuclei fused. The egg moved via the narrow entrance at the ventral region into the oviduct, where the eggshell was formed. When the eggshell was complete, and while embryogenesis proceeded, the egg was deposited. In the meantime some ova began to expand sequentially and two joining pronuclei appeared in expanding eggs. The joining pronuclei in the first egg proved male diploidy. This is additional evidence of pseudo-arrhenotoky in this phytoseiid mite species, since the first eggs developed into males.     

Some Aspects of Natural Selection in Arrhenotokous Populations

SYNOPSIS. Obligate arrhenotoky is known to have evolved onlyeight times, once each in the rotifer order Monogonanta, thearachnid order Acarina, the insect orders Hymenoptera and Thysanoptera,and in two families of Homoptera (Coccidae and Aleyrodidae)and Coleoptera (Scolytidae and Micromalthidae). In this paper it is argued that in male haploids the populationgenetic parameter which is most nearly analogous to the averagefitness in ordinary diploids is the geometric mean of the averagefitness in the two sexes. The geometric mean fitness is maximizedby selection in a random mating population, provided one takesinto account the fact that the system of inheritance imposesconstraints on the gene frequencies in males and females. Furthermore,under sufficiently small selection intensities, the rate ofchange in geometric mean fitness is approximately equal to theadditive genetic variance infitness, where the additive varianceis defined as the variance due to the regression on a modelof gene action which assumes no dominance in females and completedosage compensation in males. Some problems concerning the sex ratio in male haploids whenthere is extreme inbreeding are also discussed.     

Sex ratio bias in the dung beetle <Emphasis Type="Italic">Onthophagus taurus</Emphasis>: adaptive allocation or sex-specific offspring mortality?

Sex allocation theory predicts that females should adjust the sex of their offspring when the fitness returns of one sex are higher than the other. However, biased sex ratios may also arise if mortality differs between the sexes. Here, we examine whether offspring sex ratio bias in the dung beetle, Onthophagus taurus, represents adaptive sex allocation by females or is due to sex-specific mortality. First, we re-analyze an existing data set to show that females produce an excess of daughters when mating to smaller, less attractive males and near equal sex ratio with large, more attractive males. We show, that this results from females adjusting larval provisions after mating to males of variable attractiveness which in turn influences the likelihood that sons die during development. Second, we conduct a manipulative experiment varying the quantity and quality of larval provisions and show that the mortality of sons increased when larval provisions were reduced. Collectively, our work demonstrates that offspring mortality is contingent on the amount of resources provisioned by females and that sons have greater nutritional demands than daughters during development, leading to higher mortality. Our results therefore demonstrate the importance of considering sex-specific offspring mortality in studies of sex ratio evolution.     

Mating Behavior and Reproductive Potential in the Turnip Moth Agrotis segetum (Lepidoptera: Noctuidae)

We investigated the lifetime mating potential and the reproductive behavior of male and female turnip moths Agrotis segetum (Schiff.) under field and laboratory conditions. The sex ratio was 1 : 1 in a lab-reared population as well as in two wild populations. Males were capable of mating repetitively a relatively large number of times (mean of 6.7 ± 2.7 matings) when given access to new virgin females throughout their lifetimes. Females seldom mated more than once (mean ± 1.3 ± 0.6 matings), indicating a male-biased operational sex ratio. The mean potential lifetime mating was five times higher in males, while the coefficient of variance was lower in males. There was no differences in longevity between animals that were allowed to mate and animals not allowed to mate, indicating no direct costs or benefits of mating in physiological terms. In males, the number of matings was positively correlated with longevity, but this was not the case in females. Nor was there a correlation between the number of female matings and the number of fertilized eggs. There was a negative correlation between the number of eggs fertilized and the number of times males had previously mated, indicating that male ejaculates were limited. Male spermatophore size also decreased with number of achieved matings. Laboratory-reared females attracted males in the field throughout their lifetimes, with a peak at 3–7 days of age. Wild males, allowed to choose between pairs of caged females in the field, were attracted in equal numbers to females of different ages. Females did not show any mate-rejection behavior in the field. They mated with the first male that courted them. No incidence of mate replacement by males arriving later to already courted females were recorded.     

Comparative toxicity of some acaricides to the predatory mite, Phytoseiulus persimilis and the twospotted spider mite, Tetranychus urticae

The relative toxicity of someacaricides to the predatory mite, Phytoseiulus persimilis and the twospottedspider mite, Tetranychus urticae (Acari: Phytoseiidae, Tetranychidae) wasevaluated in laboratory. Five of theacaricides tested, including bifenazate,acequinocyl, chlorfenapyr, flufenoxuron andfenbutatin oxide, were much less toxic to adultfemales and immatures of P. persimilisthan to those of T. urticae, and adultfemale predators treated with these fiveacaricides produced 84±96% as many eggs as didcontrol females. Etoxazole did not seriouslyaffect the survival and reproduction of adultfemale predators but caused high mortalityrates in eggs and larvae of P.persimilis. Milbemectin and fenazaquin werevery toxic to adult females and immatures ofP. persimilis. Adult female predatorssurvived on a diet of spider mites treated withbifenazate, acequinocyl, chlorfenapyr,flufenoxuron and fenbutatin oxide, and theirfecundity, prey consumption and the sex ratioof the progeny were not substantially affected. Based on the results, bifenazate, acequinocyl,chlorfenapyr, flufenoxuron and fenbutatin oxideappeared to be the promising candidates for usein integrated mite management programs whereP. persimilis is the major naturalenemy.     

INEXPLICABLY FEMALE‐BIASED SEX RATIOS IN MELITTOBIA WASPS

The sex ratio behavior of parasitoid wasps in the genus Melittobia is scandalous. In contrast to the prediction of Hamilton's local mate competition theory, and the behavior of numerous other species, their extremely female‐biased sex ratios (1–5% males) change little in response to the number of females that lay eggs on a patch. We examined the mating structure and fitness consequences of adjusting the sex ratio in M. australica and found that (1) the rate of inbreeding did not differ from that expected with random mating within each patch; (2) the fitness of females that produced less female‐biased sex ratios (10 or 20% males) was greater than that of females who produced the sex ratio normally observed in M. australica. These results suggest that neither assortative mating nor asymmetrical competition between males can explain the extreme sex ratios. More generally, the finding that the sex ratios produced by females led to a decrease in their fitness suggests that the existing theory fails to capture a key aspect of the natural history of Melittobia, and emphasizes the importance of examining the fitness consequences of different sex ratio strategies, not only whether observed sex ratios correlate with theoretical predictions.     

Patterns of sex ratio, virginity and developmental mortality in gregarious parasitoids

Theory considering sex ratio optima under ‘strict local mate competition with offspring groups produced by a single foundress’ makes a suite of predictions, one of which is a mean female bias. Treating individual offspring as discrete units, theory further predicts sex ratios to have low variance (precise sex ratio) and to equal the reciprocal of clutch size (one male per clutch). The maternal decision may be complicated by imperfect control of sex allocation, limited insemination capacity of sons and offspring developmental mortality: each can lead to virgin daughters (with zero fitness) and consequently select for less biased sex ratios. When clutches are small and/or developmental mortality is common, appreciable proportions of virgins are expected, even when control of sex allocation is perfect and the mating capacity of males is unlimited. This suite of predictions has been only partially tested. We provide further tests by examining sex ratios and developmental mortalities within and across species of locally mating parasitoids. We find a wide range of mean developmental mortalities (6–67%), but mortality distributions are consistendy overdispersed (have greater than binomial variance) and sexually differential mortality appears to be absent. Sex ratios are female biased and have low variance, but are not perfectly precise and variance is increased by mortality within species and (equivocally) across species. Sex ratios less biased than the reciprocal of clutch size are observed; probably due to a maternal response to developmental mortality in one species, and to limited insemination capacity in others. Cross species comparisons indicate that mean proportions of mortality and virginity are positively correlated. Virginity is more prevalent than predicted among species with higher mortalities but not among lower mortality species. Predicted relationships between virginity and clutch size are supported in species with lower mortalities but only partially supported when mortality rates are higher.     

Canalization of body size matters for lifetime reproductive success of male predatory mites (Acari: Phytoseiidae)

The adaptive canalization hypothesis predicts that highly fitness‐relevant traits are canalized via past selection, resulting in low phenotypic plasticity and high robustness to environmental stress. Accordingly, we hypothesized that the level of phenotypic plasticity of male body size of the predatory mites Phytoseiulus persimilis (low plasticity) and Neoseiulus californicus (high plasticity) reflects the effects of body size variation on fitness, especially male lifetime reproductive success (LRS). We first generated small and standard‐sized males of P. persimilis and N. californicus by rearing them to adulthood under limited and ample prey supply, respectively. Then, adult small and standard‐sized males were provided with surplus virgin females throughout life to assess their mating and reproductive traits. Small male body size did not affect male longevity or the number of fertilized females but reduced male LRS of P. persimilis but not N. californicus. Proximately, the lower LRS of small than standard‐sized P. persimilis males correlated with shorter mating durations, probably decreasing the amount of transferred sperm. Ultimately, we suggest that male body size is more strongly canalized in P. persimilis than N. californicus because deviation from standard body size has larger detrimental fitness effects in P. persimilis than N. californicus. © 2014 The Authors. Biological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 889–899.     

Mating system and sex allocation in the gregarious parasitoid Cotesia glomerata

The gregarious parasitoid Cotesia glomerata (L.) is often presumed to possess the characteristic attributes of a species that manifests local mate competition (LMC), as it commonly produces female-biased broods. However, our field surveys of sex ratio and laboratory observations of adult behaviour showed that this species is subject to partial local mate competition caused by natal dispersal. On average, 30% of males left their natal patch before mating, with the proportion of dispersing males increasing with an increase in the patch's sex ratio (i.e. proportion of males). Over 50% of females left their natal patch before mating, and only 27.5% of females mated with males emerging from the same natal patch. Although females showed no preference between males that were and were not their siblings, broods from females that mated with siblings had a significantly higher mean brood sex ratio (0.56) than broods from females that mated with nonsiblings (0.39). Furthermore, brood sex ratios increased as inbreeding was intensified over four generations. A field population of this wasp had a mean brood sex ratio of 0.35 over 3 years, which conformed well to the evolutionarily stable strategy sex ratio (r=0.34) predicted by Taylor's partial sibmating model for haplodiploid species. These results suggest that the sex allocation strategy of C. glomerata is based on both partial local mate competition in males and inbreeding avoidance in females. In turn, this mating system plays a role in the evolution of natal dispersal behaviour in this species.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Photographic sampling: a photographic sampling method for mites on plants

A photographic sampling method for mites on plants was evaluated using Tetranychus urticae and Phytoseiulus persimilis on pepper plants. It was found to be 92% accurate for T. urticae eggs and 98% accurate for P. persimilis eggs at densities up to 45 eggs per cm² for T. urticae, and up to 3 eggs per cm² for P. persimilis. The motiles of the two species were not confused, nor were they confused with exuviae or other matter.