Effect of host diet and adult parasitoid diet on egg load dynamics and egg size of braconid parasitoids attacking Anastrepha ludens

The quantity and quality of host nutrients can affect fitness‐related traits in hymenopteran parasitoids, including oogenesis. The present study tested the prediction that a high host quality will influence oogenesis‐related traits positively in synovigenic parasitoids, and that a high‐quality adult parasitoid diet can positively affect the same parameters, potentially compensating for development on low‐quality hosts. Four braconid parasitoid species with contrasting life histories are reared on a low‐quality diet [Anastrepha ludens Loew (Diptera: Tephritidae) larvae reared on mango] or a high‐quality (artificial) diet. Adult parasitoids are provided with a high‐quality (honey ad libitum), moderate‐quality (honey every other day) or low‐quality (guava pulp) diet. Generalist species that encounter high variation in host quality naturally are predicted to be more flexible in dealing with nutrient shortfalls than specialist species. By contrast to the predictions, low‐quality hosts yield parasitoids with higher egg loads in two species: Opius hirtus Fisher and Diachasmimorpha longicaudata Ashmead. However, as predicted, a high‐quality adult diet exerts a positive effect on egg load (Utetes anastrephae Viereck), egg size (Doryctobracon crawfordi Viereck) and egg maturation rate (D. longicaudata, O. hirtus and U. anastrephae). The generalist D. longicaudata varies in egg load and maturation rate depending on host quality and adult diet, respectively. Evidence of the combined effect of both factors on parasitoid fertility is presented for the specialist O. hirtus. The theoretical and practical implications of these findings are discussed.     

Influence of initial egg density and host size on the development of the gregarious parasitoid <Emphasis Type="Italic">Bracon hebetor</Emphasis> on three different host species

Bracon hebetor (Say) (Hymenoptera : Braconidae) is a gregarious parasitoid that attacks a variety of important lepidopterous pests of stored product and in the field. In this study the effect of host species, size and larval competition on parasitoid size, survival and development were investigated. In laboratory studies, wasp eggs at a range of densities, were placed on larvae of different weight of three Lepidoptera host species namely Adoxophyes orana (Fischer von Röslerstamm, Tortricidae), Plodia interpunctella (Hubner, Pyralidae) and, Lobesia botrana(Dennis & Schiffermueller, Tortricidae). On A. orana survival of immature parasitoids was very low at all densities and different host weights. On L. botrana survival progressively reduced as egg density increased at both host weights examined for this host. Survival on P. interpunctella was significantly affected by egg density but not by host weight. Initial egg density had a significant effect on the size of emerging adults from each rearing host. Smaller adult parasitoids emerged as egg density per larva increased. Larval host weight of P. interpunctella and A. orana had a significant effect on the size of emerging adult parasitoids mainly at the higher egg densities used in these experiments. The above results of host quality on fitness of parasitoid are discussed.     

Gamete number and size correlate with adult size in the egg parasitoid Trichogramma euproctidis

In parasitoids, the size of the adult is influenced by the size and quality of the host in which it develops. Body size is generally positively correlated with several adult fitness proxies (fecundity, longevity, and mating capacity). The initial resources available to an individual can influence gamete production (sperm and oocytes), and the number and quality of gametes produced directly influence the expected fitness of both males and females. Gamete production in relation to adult body size was quantified in Trichogramma euproctidis (Girault) (Hymenoptera: Trichogrammatidae), a short‐lived egg parasitoid of lepidopteran species. To avoid host quality variation, male and female parasitoids of different body sizes were produced using superparasitism by allowing mated and virgin female parasitoids to oviposit on Trichoplusia ni Hübner (Lepidoptera: Noctuidae) eggs. Seminal vesicles and ovaries of their offspring were dissected to count oocytes and to measure sperm length and oocytes volume. Tibia length was also measured to estimate body size. The number of oocytes, volume of oocytes, maternal investment index [= (number of oocytes × mean volume of oocytes)/10 000] and sperm length were all significantly positively correlated to body size. These results show that initial resources acquired during larval stage induce phenotypic plasticity in gamete production in both male and female T. euproctidis. Whereas number of sperm and oocytes can influence the fitness of males and females through increased mating capacity and fecundity, variation in gamete size (sperm length and oocyte volume) could also affect the fitness of an individual through sperm and larval competition.     

Development of the solitary endoparasitoid Microplitis demolitor: host quality does not increase with host age and size

Abstract.  1. Many studies examining the relationship between host size, an index of host quality, and parasitoid fitness use development time and/or adult parasitoid size as currencies of fitness, while ignoring pre-adult mortality. Because the physiological suitability of the host may vary in different stages, sizes, or ages of hosts, a misleading picture of host quality may therefore be obtained in cases where fitness is based on only one or two developmental traits.
2. The development of the solitary koinobiont endoparasitoid Microplitis demolitor is examined in different larval age-classes of its host the soybean looper Pseudoplusia includens . Hosts were parasitised on days 1–8 after hatching from the egg, and development time, adult body size, and mortality of the parasitoid were compared.
3. A comparison of larval growth trajectories (using dry body mass) of M. demolitor revealed that parasitoid larvae attained over twice as much body mass in old hosts than in younger hosts. Similarly, adult parasitoid size at eclosion generally increased with host size, although parasitoids developing in smaller hosts lost a much lower proportion of mass between pupation and eclosion.
4. Overall egg-to-adult development was most rapid in intermediate-aged hosts, and longer in hosts at opposite ends of the age continuum. Moreover, parasitoid mortality varied non-linearly with host stage, and was generally higher in very young and older hosts.
5. Based on these results and other empirical data for koinobionts, it is argued that fitness functions in this group of parasitoids are not simply a positive function of host size or age, but instead may be distinctly dome-shaped, both patterns reflecting the degree of physiological and nutritional compatibility between the two organisms.     

Influence of egg load and host size on host-feeding behaviour of the parasitoid Aphytis lingnanensis

Abstract.

• 1 Direct behavioural assays were used to investigate the influences of host size and parasitoid egg load on the decision to host feed versus oviposit made by the parasitoid Aphytis lingnanensis Compere. Egg load was manipulated without concurrent influences on the history of host contact by exploiting size-related variation in fecundity and by holding parasitoids at different temperatures to vary the rate of oocyte maturation.
• 2 Host feeding comprised a series of feeding bouts, separated by renewed probing of the scale insect body. Successive feeding bouts were progressively shorter, suggesting that hosts represent ‘patches’ yielding resources at a decelerating rate.
• 3 Parasitoids were significantly more likely to host feed on smaller hosts and oviposit on larger hosts.
• 4 Neither egg load nor the treatment variables (parasitoid size and holding temperature) exerted significant influences on the decision to host feed versus oviposit on second instar (low quality) hosts.
• 5 The failure to observe an effect of egg load on host-feeding decisions was not simply a reflection of the parasitoids being entirely insensitive to egg load; significant effects of egg load on parasitoid search intensity and clutch size decisions were observed.
• 6 Parasitoids developing on second instar (low quality) hosts experienced high levels of mortality late during development and yielded very small adults.
• 7 The discord between these experimental results and predictions regarding the importance of egg load underscores the need for additional work on the proximate basis for host-feeding decisions and the nutritional ecology of insect parasitoids.

     

Age-dependent clutch size in a koinobiont parasitoid

Abstract.  1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.
2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals.
3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.
4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time.
5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.
6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.
7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.     

Host egg pigmentation protects developing parasitoids from ultraviolet radiation

Parasites rely on their hosts not only for nutrition and reproduction, but also for protection against natural enemies and adverse climatic conditions. In host‐parasite interactions, protective characteristics of both players are important to consider regarding damaging effects of environmental hazards. While ultraviolet radiation (UVR) is pervasive and harmful to organisms in general, its impact on parasite fitness remains understudied. Moreover, studies that do examine the effects of UV exposure on parasitic organisms tend to neglect host traits, which may vary inter‐ or intra‐specifically and thus confer different levels of environmental protection. We examined in the laboratory whether the UV‐protective value of host egg pigmentation could also benefit parasitoids, using the egg parasitoid Telenomus podisi and the predatory stink bug Podisus maculiventris. This host species lays eggs of variable pigmentation levels from light to dark grey, an adaptation protecting its own embryos from UVR. We showed that higher levels of host egg pigmentation protect parasitoids subjected to a developmental exposure to UVR, increasing emergence rates by up to 86% and reducing development time by up to 4%. This protective effect of host pigmentation was context‐dependent, being less pronounced at low UVR intensity and towards the end of parasitoid development. Parasitoids that emerged from darker‐coloured eggs exposed to UVR were of slightly larger size than those developing in light‐coloured eggs, but other fitness‐related traits (fecundity, longevity, sex ratio) were unaffected. This study provides the first experimental evidence that host pigmentation can increase host suitability for parasitic organisms, and emphasizes the importance of considering trait variation in interacting species when investigating the susceptibility of ecological communities to important abiotic environmental factors.     

The effect of host size on the sex ratio of Syngaster lepidus, a parasitoid of Eucalyptus longhorned borers (Phoracantha spp.)

The solitary larval ectoparasitoid, Syngaster lepidus Brullé, parasitizes the cryptic larvae of two wood-boring beetles, Phoracantha recurva Newman and Phoracantha semipunctata F. The objective of this study was to determine how the female parasitoids allocated the sex of progeny when presented with larval hosts of uniform size classes. Host size was directly correlated with age of the Phoracantha larval hosts. Groups of Phoracantha larvae of a single age class (2-, 3-, 4-, or 5-week-old) were exposed to parasitoids, and sex ratios of the resulting parasitoid progeny from each host age class were determined. A significant relationship was observed among the sizes of P. recurva and P. semipunctata hosts and the sex ratio of emerging parasitoids. Parasitized 2-week-old beetle larvae of both Phoracantha spp. produced only male S. lepidus progeny, whereas older larval hosts produced increasing proportions of female parasitoids (up to 80% females from 5-week-old hosts). Two-week-old Phoracantha larvae of both species produced fewer parasitoids than host larvae 3–5-week-old. The size of parasitoid progeny consistently increased with host larval age (size), and female parasitoids were larger than males across all host size classes. Male S. lepidus developed in approximately 25 days from 2-week-old hosts, and 19–21 days in 3–5-week-old hosts. Female S. lepidus developed in 22–25 days, with developmental time increasing with host size.     

Income resources and reproductive opportunities change life history traits and the egg/time limitation trade‐off in a synovigenic parasitoid

1. Synovigenic parasitoids emerging with no or only a few mature oocytes could not rely on only capital resources, but also need to acquire income resources. Income resources in nature can either contribute to egg maturation as a food resource and/or create unpredictability in realised reproductive opportunities for synovigenic parasitoids. Therefore, we hypothesised such resources could affect life history traits and the risks of egg/time limitation in synovigenic parasitoids. 2. Using the Ovigeny Index, we investigated the effects of various host availability levels (unavailable, limited, and unlimited availability) and non‐host foods (water and honey) on life history traits and on the occurrence of egg/time limitation in Eretmocerus hayati, a predominant parasitoid on Bemisia tabaci. 3. The Ovigeny Index of Er. hayati was 0.28, which suggested it was a typical synovigenic species. Both host availability levels and non‐food type had major effects on life history traits of this parasitoid, but the availability of hosts for both feeding and reproduction was the key factor. Meanwhile, egg/time limitation was encountered by all wasps and its intensity varied with host availability levels. 4. Our results confirmed that the income resource and reproductive opportunity played a central role in shaping the life history and risks of egg/time limitation of a synovigenic parasitoid.     

The developmental strategies and related profitability of an idiobiont ectoparasitoid Sclerodermus pupariae vary with host size

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Increasing host age does not have the expected negative effects on the fitness parameters of an egg parasitoid

Host age is an important determinant of host acceptance and suitability for egg parasitoids. As host embryonic development advances, the quality of resources available to the parasitoid offspring typically declines, usually resulting in reduced acceptance levels by foraging females and lower offspring fitness. We examined the ability of the parasitoid Telenomus podisi Ashmead (Hymenoptera: Scelionidae) to parasitize and develop in Podisus maculiventris (Say) (Hemiptera: Pentatomidae) eggs of different ages. In laboratory experiments, we measured the effect of host age (6, 24, 48, 72, 96, or 120 h old) on parasitism rate and offspring fitness parameters such as survival, development time, sex ratio, and size. Contrary to our expectations, parasitism rate did not differ between host age treatments, nor did sex ratio allocation, offspring size, or the fecundity of newly emerged female offspring. However, parasitoid offspring had a longer development time with increasing host age. This trend was stronger for males than for females, which we suggest could reduce the degree of protandry among offspring emerging from older host eggs, thus increasing the rate of virginity upon leaving the emergence patch and resulting in more frequent off‐patch mating by female offspring in nature. Overall, our results suggest that all stages of P. maculiventris embryonic development are suitable for acceptance and development of T. podisi. Unlike most species of egg parasitoids, T. podisi has evolved mechanisms to utilize host resources, regardless of host developmental stage, with relatively minor fitness consequences.     

The relationship among host plant species,egg clutch size,and level of parasitism for the sharpshooter Tapajosa rubromarginata

The sharpshooter Tapajosa rubromarginata (Signoret) (Hemiptera: Cicadellidae, Proconiini), a vector of the bacterium Xylella fastidiosa Wells et al. (Xanthomonadaceae) that causes citrus variegated chlorosis, has more than 30 reported host plant species. The fitness of a phytophagous insect is determined by the host plant suitability, plant resistance, and the natural enemies. The aim of this study was to: (1) identify plant species utilized as oviposition substrate by T. rubromarginata in the field; (2) establish the relationship between plants and clutch size; (3) establish the relationship among host plants, clutch size, and level of parasitism; and (4) establish variations in parasitoid composition and abundance in the various host plants. Egg masses of the sharpshooter were surveyed on plants reported as hosts, or those that were abundant in the study site. The number of eggs of the sharpshooter and emerged parasitoids were recorded for all the collected masses. We found egg masses of T. rubromarginata on 12 out of 21 plant species sampled. The size of the egg masses was greatly influenced by the type of leaf venation and to a lesser extent by the plant species. Parasitism rates were influenced by both leaf venation and host plant. Trichogrammatidae species were mostly associated with egg masses in plants with parallel-veined leaves, whereas Mymaridae attacked masses laid in reticular-veined leaves. The choice between a good host plant, but heavily attacked by parasitoids, and the host plants that are less suitable for nymphs but less frequently attacked by natural enemies, was a trade-off for T. rubromarginata females to increase their fitness. We conclude that the host plant utilization by T. rubromarginata females in the field could be influenced by leaf structure and the strategy to avoid parasitism by selecting plants that were less attractive for parasitoids.     

Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom‐up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host‐plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host‐plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.     

How two different host species influence the performance of a gregarious parasitoid: host size is not equal to host quality

1. Hyssopus pallidus Askew (Hymenoptera, Eulophidae) is a gregarious ectoparasitoid of the two tortricid moths species Cydia molesta Busck and C. pomonella L. (Lepidoptera, Tortricidae). It paralyses and parasitizes different larval instars of both species inside the apple fruit, which leads to the death of the caterpillar. 2. We assessed the influence of host species characteristics and host food on the performance of the parasitoid female in terms of clutch size decisions and fitness of the F(1) generation. 3. A comparison of clutch size revealed that female parasitoids deposited similar numbers of eggs on the comparatively smaller C. molesta hosts as on the larger C. pomonella hosts. The number of parasitoid offspring produced per weight unit of host larva was significantly higher in C. molesta than in C. pomonella, which is contrary to the general prediction that smaller hosts yield less parasitoid offspring. However, the sex ratio was not influenced by host species that differed considerably in size. 4. Despite the fact that less host resources were available per parasitoid larva feeding on C. molesta caterpillars, the mean weight of emerging female wasps was higher in the parasitoids reared on C. molesta. Furthermore, longevity of these female wasps was neither influenced by host species nor by the food their host had consumed. In addition we did not find a positive relationship between adult female weight and longevity. 5. Parasitoid females proved to be able to assess accurately the nutritional quality of an encountered host and adjust clutch size accordingly. These findings indicate that host size is not equal to host quality. Thus host size is not the only parameter to explain the nutritional quality of a given host and to predict fitness gain in the subsequent generation.     

Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

The effect of host size on quality attributes of the egg parasitoid,Trichogramma pretiosum

In a study of the quality ofTrichogramma pretiosum Riley (Hymenoptera: Trichogrammidae), we compared female wasps emerging from natural hosts, parasitized in the laboratory or the field with those emerging from factitious hosts used for commercial mass production. Females from the natural hosts were larger, more fecund, and longer lived than those from the factitious hosts. Compared to small females, large female wasps are substantially more fecund when honey (carbohydrate) is available but marginally more fecund when honey is unavailable. The size of a femaleT. pretiosum depends on two factors: the size of the host egg from which it emerges even when the wasp was gregarious, and the number of conspecifics that emerge with it. The similarities in the size distribution of female wasps emerging from natural hosts, in conjunction with the mechanism by whichTrichogramma measure host size and allocate eggs accordingly, suggests the hypothesis that size related components of fitness in femaleT. pretiosum are under strong selection in the field.     

Body size and the timing of egg production in parasitoid wasps

In insects several key fitness-related variables are positively correlated with intraspecific variation in body size, but little is known about size-related variation in the timing of egg production within species. Female insects are known to vary in the degree to which they concentrate egg production into the early part of life. This variation has been quantified as the ovigeny index, defined as the proportion of the maximum potential lifetime complement of eggs that is mature following emergence from the pupa. We tested the hypothesis that the timing of egg production depends both on body size and on host availability, by means of a dynamic programming model that predicted optimal resource allocation to reproduction and survival together with the resulting ovigeny index, in non-feeding synovigenic parasitoids of different sizes. As body size increases, the proportionate increase in resource allocation to initial egg load is less than the proportionate increase in allocation to lifetime fecundity and potential life span, leading to a deferred investment in reproduction as shown by a decrease in ovigeny index. High habitat quality and high habitat stochasticity in reproductive opportunities have a significant effect on the optimal allocation of resources to reproduction and survival, and thus select for early reproduction, i.e. an increased ovigeny index. The ovigeny concept – ovigeny index together with its life-history correlates – enables understanding of the general occurrence of size-related deferment of reproductive investment in parasitoid wasps and also helps explain a significant part of the considerable life-history variation found among such insects.     

The relationship between host selection behaviour and offspring fitness in a koinobiont parasitoid

1. When host quality varies, optimal foraging theory assumes that parasitic wasps select hosts in a manner that increases their individual fitness. In koinobiont parasitoids, where the hosts continue developing for a certain period of time after parasitisation, host selection may not reflect current host quality but may be based on an assessment of future growth rates and resources available for the developing larvae. 2. When presented with hosts of uniform quality, the koinobiont parasitoid Leptomastix dactylopii exhibits a characteristic host‐selection behaviour: some hosts are accepted for oviposition on first encounter, while others are rejected several times before an egg is laid in them, a behaviour that is commonly associated with a changing host acceptance threshold during the course of a foraging bout. 3. The fitness of the offspring that emerged from hosts accepted immediately upon encounter was compared with the fitness of offspring emerged from hosts rejected several times before being accepted for oviposition. 4. The pattern of host acceptance and rejection was not related to any of the measured fitness parameters of the offspring emerging from these hosts (development time, size at emergence, sex ratio at emergence, and female offspring egg load). 5. While complex post facto adaptive explanations can be devised to explain the nature of such a time and energy consuming host selection process, it is suggested that physiological constraints on egg production or oviposition may provide an alternative, purely mechanistic, explanation for the results obtained.     

Flexible larval development and the timing of destructive feeding by a solitary endoparasitoid: an optimal foraging problem in evolutionary perspective

1. In studying the evolution of life-history strategies in parasitoids, considerable attention has been paid to the relationship between host quality and parasitoid fitness. Various workers have reported that host quality influences parasitoid size, development time, and survival. Because body size is frequently correlated with fecundity, longevity, and host-finding ability in parasitoids, this parameter is usually considered to be the main target of selection. 2. In koinobiont parasitoids that consume the entire host before pupation, adult parasitoid size and development time are often strongly correlated with host size at the time when it is developmentally arrested through destructive feeding by the parasitoid larva. 3. Here, a mathematical model is proposed to describe the larval feeding behaviour of the solitary koinobiont endoparasitoid Venturia canescens in four larval stadia of its host Plodia interpunctella. In particular, the model describes how adult size, represented by an exponential growth rate, and development time are traded off when the parasitoid develops in nutritionally suboptimal second stadium hosts. 4. Using a graphical model, the different conditions faced by V. canescens during development in various host species of greatly differing mass are illustrated. 5. It is argued that the relative importance of size and development time on parasitoid fitness is determined by ecological and biological characteristics of both host and parasitoid, and it is suggested that there may be correlations between life-history traits and host-utilisation strategies among koinobionts.