State dependent superparasitism in a solitary parasitoid: egg load and survival

In solitary parasitoids, superparasitism (the allocation ofan egg to an already parasitized host) has a payoff, measuredin offspring produced and costs, measured in eggs and time invested.Solitary parasitoids that are capable of host discriminationmust adopt the strategy that ensures the best use of both theiregg load and available lifetime. In this paper, we develop astate-dependent model defining the optimal strategy of superparasitismfor a solitary parasitoid species with overlapping generations.The fitness measure we use is based on the growth rate of thenumber of genotype copies. The model predicts that the tendencyto superparasitize should increase as the egg load of the parasitoidincreases, or as its life expectancy decreases. The model alsopredicts that under particular conditions wasps should showpartial preferences for parasitized hosts. These predictionswere tested with the parasitoid Venturia canescens (Hymenoptera:Ichneumonidae). The tendency of the wasps to superparasitizein the presence of both healthy and parasitized hosts was correlatedto egg load and access to food before the experiment A complementaryexperiment, where parasitized hosts were given sequentiallyto parasitoids, showed that V. canescens exhibits partial preferencestoward superparasitism. These experimental results and a previouswork support the predictions of the model.     

The effect of egg load on superparasitism by the snowberry fly

In studies conducted with fruits of the host plant, Symphoricarpus albus (L.), we examine the influence of egg load on the oviposition behaviour of Rhagoletis zephyria Snow (Diptera:Tephritidae). By altering the availability of exogenous protein, three classes of females with progressively increasing egg loads were produced, while keeping confounding factors such as age, experience, and mating status constant. Flies from all three classes were randomly presented with either a pheromone marked fruit, or with an unmarked fruit. Results indicate that increased egg load led to a greater propensity to accept, or superparasitize, the pheromone marked fruit. Upon dissection it was revealed that females which superparasitized had a mean egg load of 19.5 eggs (n=22), while females which rejected marked fruit had a mean egg load of 13.5 eggs (n=26). These results are consistent with the theory of adaptive superparasitism.     

假眼小绿叶蝉卵的寄生蜂种类及种群动态

通过田间采梢、室内镜下查卵和饲养观察,初步获得假眼小绿叶蝉Empoasca vitis Gthe卵寄生蜂2种,分别隶属于三棒缨小蜂属Stethynium sp.和裂骨缨小蜂属Schizophragma sp.,其中三棒缨小蜂属Stethyniumsp.是优势种。2种缨小蜂的寄生动态调查结果显示,除8月和12月寄生率较低外,其它月份的自然寄生率都在30%以上,特别是10~11月寄生率最高达65%,对小绿叶蝉的虫口数量具有重要控制作用,值得进一步研究和保护利用。     

麦红吸浆虫及其卵寄生蜂混合种群空间格局

运用地统计学方法对不同时期麦红吸浆虫及其卵寄生蜂混合种群(宽腹姬小蜂和尖腹黑蜂)的空间格局进行了分析.结果表明:麦红吸浆虫休眠体的半变异函数的最优模型为球型,成虫的最优模型为球型-指数型,幼虫最优模型为线性有基台型,卵寄生蜂混合种群半变异函数的最优模型为球型-指数型.麦红吸浆虫休眠体、成虫羽化初期、成虫羽化高峰期、幼虫和卵寄生蜂混合种群的空间相关范围分别为53.6、190.6、154.1、4.2和280.3 m,空间变异强度分别为30.5%、95.6%、96.3%、14.9%和95.3%.采用普通克立格插值法模拟的空间分布模拟图可较好地从时间、空间两个角度直观地分析不同时期麦红吸浆虫及其卵寄生蜂混合种群的动态变化.     

Field studies on the behaviour of two egg parasitoids of the pine processionary mothThaumetopoea pityocampa

An investigation on the behaviour ofTetrastichus servadeii Domenichini (Hym.: Eulophidae) andOoencyrtus pityocampae Mercet (Hym.: Encyrtidae) was carried out in 2 stands ofPinus nigra Arnold in the Venitian Prealps (NE Italy) by quantitative observations in the field and by trapping with different baits. The results indicate that: 1)O.pityocampae is caught by traps baited with pityolure, the synthetic sex pheromone of the host, whileT. servadeii was never caught. 2) The host selection process ofT. servadeii seems to be associated with the host plant ofThaumetopoea pityocampa (Denis & Schiffermüller) and with the aggregative behaviour of the parasitoid females. 3) The frequency distribution ofT. servadeii on the egg masses follows the negative binomial distribution; the degree of aggregation decreases with the egg mass age. These results may be taken into account in the utilization of egg parasitoids for the biological control ofTh. pityocampa.      

Host-feeding and oviposition by parasitoids on hosts of different fitness value: Influences of egg load and encounter rate

A dynamic optimization model is presented for the decision to host feed or oviposit on hosts by female parasitoids. Optimal host utilization decisions are compared between two host types with different fitness payoffs for oviposition. The model predicts that hosts of higher fitness value should always be used for oviposition unless the egg load approaches zero. This general prediction is not influenced by levels of host availability or metabolic reserves, the age of the parasitoid, or the magnitude of the fitness difference. An egg-load threshold is predicted above which lower value hosts should be used for oviposition and below which they are used for host feeding. The position of this egg-load threshold is higher when the difference in fitness between host types is larger. The threshold is also higher when overall host availability is high or metabolic reserves for the production of new eggs are low. The threshold for oviposition on low-value hosts decreases to zero near the end of the parasitoid’s life. Under conditions where high-value hosts are rarely encountered compared to low-value hosts, the model predicts that lower value hosts should be accepted for oviposition at a lower egg-load threshold.     

A functional response model of a predator population foraging in a patchy habitat

1. Functional response models (e.g. Holling's disc equation) that do not take the spatial distributions of prey and predators into account are likely to produce biased estimates of predation rates. 2. To investigate the consequences of ignoring prey distribution and predator aggregation, a general analytical model of a predator population occupying a patchy environment with a single species of prey is developed. 3. The model includes the density and the spatial distribution of the prey population, the aggregative response of the predators and their mutual interference. 4. The model provides explicit solutions to a number of scenarios that can be independently combined: the prey has an even, random or clumped distribution, and the predators show a convex, sigmoid, linear or no aggregative response. 5. The model is parameterized with data from an acarine predator-prey system consisting of Phytoseiulus persimis and Tetranychus urticae inhabiting greenhouse cucumbers. 6. The model fits empirical data quite well and much better than if prey and predators were assumed to be evenly distributed among patches, or if the predators were distributed independently of the prey. 7. The analyses show that if the predators do not show an aggregative response it will always be an advantage to the prey to adopt a patchy distribution. On the other hand, if the predators are capable of responding to the distribution of prey, then it will be an advantage to the prey to be evenly distributed when its density is low and switch to a more patchy distribution when its density increases. The effect of mutual interference is negligible unless predator density is very high. 8. The model shows that prey patchiness and predator aggregation in combination can change the functional response at the population level from type II to type III, indicating that these factors may contribute to stabilization of predator-prey dynamics.     

Functional response and mutual interference of Peristenus spretus (Hymenoptera: Braconidae), a parasitoid of Apolygus lucorum (Heteroptera: Miridae)

Peristenus spretus Chen & van Achterberg (Hymenoptera: Braconidae) is a solitary endoparasitoid, which is considered for augmentative biological control of Apolygus lucorum Meyer-Dür (Heteroptera: Miridae) in Chinese cotton fields. Since the association of P. spretus with A. lucorum was only recently discovered, the biology of the parasitoid remains unknown. In order to understand its reproductive biology, the mutual interference and functional response of P. spretus were investigated by altering either the parasitoid or the host density while keeping the other constant. In both experiments, the effects of parasitoid and host densities on parasitism, superparasitism, progeny production and sex ratio were assessed. P. spretus exhibited a Holling type II functional response to changing host densities, indicating that parasitism increases with increasing host density until the parasitoid reaches its maximum reproductive capacity. The model suggested that a single P. spretus female could parasitise a maximum of 88 nymphs per day or four nymphs per hour. Increasing the wasp-nymph ratio from 1:10 to 1:80 significantly increased the offspring production more than fivefold from ±5.8 to ±35.6; further increasing the host densities (above 80 nymphs) did not significantly increase offspring production. Strong mutual interference of foraging P. spretus females occurred only at high parasitoid densities. Parasitoids foraging alone produced an average progeny of 33.4, whereas parasitoids foraging in groups of 16 produced only 2.6. The optimal wasp-nymph ratio for mass-rearing P. spretus is 4:100, given that resources of parasitoids and nymphs are unlimited.     

Status of biological control by egg parasitoids of Diaprepes abbreviatus (Coleoptera: Curculionidae) in citrus in Florida and Puerto Rico

Eggs of Diaprepes abbreviatus (L.) were routinely monitored in citrus groves at ten locations in Florida during 1997 and 1998 to study egg parasitism. One citrus location was studied in Puerto Rico. No native parasitoids were recovered from 1,337 D. abbreviatus egg masses studied in Florida citrus. In contrast, an average of 35.5% (range 12.5 to 68.8%) parasitism of egg masses was reported in Puerto Rico. The parasitoids Aprostocetus gala, Horismenus spp, and Quadrastichus haitiensis were recovered from the eggs of D. abbreviatusfstudied in Puerto Rico. The Horismenus parasitoids were suspected hyperparasitoids. Releases of the parasitoid Ceratogramma etiennei from Guadeloupe were initiated during 1998 at each of the Florida research sites. By the end of 1998, C. etiennei had been recovered from D. abbreviatus eggs at two of nine locations in Florida citrus. The parasitoid was recovered from 1 of 34 egg masses at one of these locations during the month of September and from 3 of 34 egg masses at the other location during the month of November. Whether or not C. etiennei establishes itself at one or more locations in Florida remains to be seen.     

The time and egg budget of Leptopilina clavipes, a parasitoid of larval Drosophila

Abstract.

• 1 For the understanding of the influence of natural selection on the persistence of host selection behaviour in populations of Drosophila parasitoids it is important to know whether parasitoids will become time- or egg-limited. We investigated whether the Drosophila parasitoid Leptopilina clavipes (Hartig) meets egg- or time-limited conditions in the field.
• 2 To this end the following aspects of the parasitoid's life were studied: egg load at emergence, travelling velocity between patches, patch residence times, oviposition rates and life expectancy. Together with the results from earlier studies on host and patch distributions, this formed the input of a ‘Monte Carlo’ simulation model, in which the life history of an individual parasitoid can be traced.
• 3 The simulations revealed that under the conditions found in the field 12.9% of the parasitoid population is egg-limited. The model was also run for a number of scenarios which reflect ‘good’ or ‘bad’ circumstances. In most cases a significant proportion of the parasitoid population proved to be egg-limited.
• 4 For the measurement of travelling velocities and patch residence times a marking method, especially applicable to small-sized parasitoids such as L.clavipes, is described. Marking did not affect survival, host habitat location or host detection rate. Parasitoids were found to be attracted to the odour of fruit-bodies of Phallus impudicus, the most important breeding substrate of their hosts.

     

The functional grazing response of a phytoplanktivorous fish Oreochromis mloticus to mixtures of toxic and non-toxicstrains of the cyanobacterium Microcystis aeruginosa

Small (10 g) tilapia ( Oreochromis niloticus ) were exposed to pure and mixed populations of toxic and non-toxic strains of the cyanobacterium Microcystis aeruginosa (100% toxic, 50% toxic, 25% toxic, 0% toxic) at two particle concentrations (1 × 10⁶ and 5 × 10^sparticles ml⁻¹). At both concentrations there was a progressive decrease in grazing rate as the percentage of toxic cells increased. Differences in opercular beat rates, and hence the volumes of water passed over the gills, were also recorded among treatments, opercular beat rates decreasing as the percentage of toxic cells increased. Although in all treatment groups with toxic cells present, the medium had detectable levels (>250 ng I⁻¹) of extracellular microcystin-LR toxin present, grazing was correlated with particle-bound rather than extracellular levels.     

Oviposition behaviour and distribution of egg masses of the European leaf roller,Archips rosanus (L.), on citrus trees

The citrus leaf roller (CLR), Archips rosanus, a threatening pest of leaves and fruitlets on fruit trees, which in recent years has led to considerable damage to different citrus trees in some parts of the Mazandaran province of Iran. Insect activity on citrus trees causes the growth of buds and leaves to stop and foliage to dry. Pest oviposition as egg masses occurs on the host tree’s trunk. Eggs spent a year in the hibernation stage and, in the following spring, hatched the first instar larvae that fed on the opened buds. The distributions of CLR egg masses, deposited in 2015–2016, were evaluated on the Thomson novel cultivar (Citrus sinensis L.) in the northern region of Iran. The results showed that the numbers of egg masses laid on the trees at different height ranges of 0–50, 50–100, 100–150 and 150–200 cm were, respectively, 38.65, 33.78, 18.54 and 9.03%. The mean number of A. rosanus eggs per mass, however, was not significant at different heights of the tree. Also, most of the egg masses were laid in the north and south directions in comparison with the west and east directions, with no significant difference in mean number in different geographical directions.     

Pollen as a source of food for egg parasitoids of the genus Trichogramma (Hymenoptera: Trichogrammatidae)

The value of corn pollen as a food source for Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) was assessed in the laboratory by testing the effect of different pollen diets on the longevity, capacity of parasitism and other life-table parameters. T. brassicae females that fed on corn pollen and water (wet filter paper dusted with pollen) lived significantly longer (4.97 days) than females that fed on water alone (2.67 days), but significantly shorter than those feeding on honey alone (8.37 days) or corn pollen and honey (8.23 days), respectively, in the absence of host eggs. The presence of European corn borer eggs (Ostrinia nubilalis Huebner, Lepidoptera: Pyralidae) significantly increased longevity. With host eggs available, female T. brassicae that fed on corn pollen and water lived significantly longer (4.9 days) than on water alone (2.60 days), but significantly shorter than on honey alone (12.33 days) or corn pollen and honey (12.17 days), respectively. The cumulative lifetime fecundity was 82.53 offspring pupae when fed on pollen and water, which was significantly higher than 61.70 for water alone but lower than feeding on honey alone (95.70) or corn pollen and honey (99.97), respectively. Compared to water alone, corn pollen and water increased the net reproduction rate (R₀), intrinsic rate of increase (r_m), and finite rate of increase (λ), and reduced the doubling time (T₂), but had no effects on the emergence rate, sex ratio, and mean generation time (T_c). In another set of experiments, 13 out of 30 females reduced the number of pollen grains offered to them. The average number of pollen grains utilized per female in this 48-h test 2.46±0.40 (mean±SE). Hence, T. brassicae used corn pollen as food that significantly increased longevity and fecundity of the tested females. Pollen and honey supplied a complete diet for T. brassicae, compared to pollen and water or water alone.     

Prey: predator ratio dependence in the functional response of a freshwater amphipod

1. First known for their shredding activity, freshwater amphipods also behave as active predators with consequences for prey population regulation and amphipod coexistence in the context of biological invasions. 2. A way to quantify predation is to determine the average consumption rate per predator, also known as its functional response (FR). 3. Although amphipods are gregarious and can display social interactions that can alter per capita consumption rates, previous studies using the FR approach to investigate amphipod predation ignored such potential mutual interference because they did not consider variations in predator density. 4. We investigated the FR of Echinogammarus berilloni feeding on dipteran larvae with joint variations in prey and predator densities. This bivariate experimental design allowed us to estimate interference and to compare the fits of the three main classes of theoretical FR models, in which the predation rate is a function of prey density alone (prey‐dependent models), of both prey and predator densities (predator‐dependent models) or of the prey‐to‐predator ratio (ratio‐dependent models). 5. The Arditi–Ginzburg ratio‐dependent FR model provided the best representation of the FR of E. berilloni, whose predation rate showed a decelerating rise to a horizontal asymptote as prey abundance increased. 6. Ratio dependence means that mutual interference between amphipods leads to prey sharing. Mutual interference is likely to vary between amphipod species, depending on their level of aggressiveness.     

苜蓿叶象甲卵块在苜蓿植株上垂直分布的研究

苜蓿叶象甲卵块分布在苜蓿植株层次之间,差异显著;在不同田块之间,差异不显著。分布区间随着生育期的变化而变化,在分枝期,主要集中在0～20cm,区间估计为[0．6524,0．7876];在开花期,主要在0～60cm,区间估计为[0．8427,1．000]。苜蓿叶象甲每个卵块的卵粒平均数为8．4粒,卵块与卵粒数存在着显著的线性关系：y＝0．0069＋8．6594xr＝0．8422**     

Pollinator functional response and plant population dynamics: Pollinators as a limiting resource

Summary During recent years, much work has focused on which factors limit the reproductive success in plants. Several studies show a strong influence of either resource limitation, pollen limitation or a combined effect of both. The theoretical arguments for resource limitation are abundant, but there has been very little work done concerning the effect of pollinator availability. In this paper we construct a model to study how the reproductive success in plants is influenced by the foraging behaviour of the pollinators. The pollinator population is assumed to have a constant population density. A functional response function for the pollinators is derived. It is similar to a Holling type II functional response. It is shown that, since the pollinators are regulated by factors not included in the model and their capability to pollinate is limited by the functional response, this is sufficient for regulating the plant population. There also exists a threshold condition for the persistence of the plant population that depended on the search rate of the pollinators and the demographic parameters of the plant population. If this threshold condition is not satisfied the plant population cannot persist and will become extinct. If the condition is satisfied the plant population grows until it is limited at the equilibrium mentioned above.     

长柄俑小蜂寄生橘小实蝇蛹的功能反应

长柄俑小蜂寄生橘小实蝇蛹的功能反应符合功能反应模型HollingⅡ型,其方程为:Na=1.5324TrN0/(1 1.5324ThN0)。每只寄生蜂在24 h内最大寄生量为17.57只橘小实蝇蛹。寄生1只寄主蛹所需的时间为1.37 h;瞬时攻击率(功能系数)为0.6526;长柄俑小蜂自身密度对寄生产生干扰效应,用Hassell-Varley模型可表示为:α=0.1085P-0.1929。     

七星瓢虫、大草蛉对桃粉蚜捕食功能研究

经测定,七星瓢虫成虫、4龄幼虫和大草蛉3龄幼虫对桃粉蚜的捕食功能反应属于Holling—Ⅱ型圆盘方程式,两种天敌对桃粉蚜具有较强的捕食能力。