Sex ratios in field populations of two parasitoids (Hymenoptera: Chalcidoidea) of Coccus hesperidum L. (Homoptera: Coccidae)

We tested several assumptions and predictions of host-quality-dependent sex allocation theory (Charnov et al. 1981) with data obtained for the parasitoid Metaphycus stanleyi Compere on its host, brown soft scale (Coccus hesperidum L.), in a California citrus grove and in the laboratory. Scales ceased growing after parasitization by M.?stanleyi. Thus, M. stanleyi may gauge host quality (=size) at oviposition. Host size positively influenced adult parasitoid size, and parasitoid size in turn influenced adult longevity of M. stanleyi. However, parasitoid fitness gains with host size and adult size were similar in males versus females. Sex allocation to individual hosts by M. stanleyi depended on host size; females consistently emerged from larger hosts than males. Host size was important in a relative sense; the mean host sizes of females versus males, and of solitary versus gregarious parasitoids varied with the available host size distribution. The offspring sex ratio of M. stanleyi reflected the available host size distribution; the sex ratio of emerging parasitoids varied with the available host size distribution. We did not detect a “critical host size” below which males emerged, and above which females emerged; rather, only females emerged from hosts in the upper size range, and a variable ratio of males and females emerged from hosts in the lower size range. We conclude that the sex ratio of field populations of M.?stanleyi is driven largely by the available size distribution of C. hesperidum. In addition, we tested predictions resulting from theoretical analyses of sex allocation in autoparasitoids with data obtained on Coccophagus semicircularis (Förster) parasitizing brown soft scale in the field. The sex ratio of C. semicircularis was consistently and strongly female biased (ca. 90% females). Based on available theoretical analyses, we suggest that this sex ratio pattern may have resulted from a very low encounter rate of secondary hosts coupled with a strong time limitation in C. semicircularis females. This explanation was the most plausible given constraints stemming from the detection of secondary hosts, their variable location within primary hosts, and their handling times. Finally, the size of hosts which yielded single versus multiple parasitoids, and the sizes of these parasitoids, were compared. These comparisons suggested that: (1) M. stanleyi females gauge host sizes precisely, and in terms of female offspring; thus a fitness penalty is not incurred by females which share a host, while males benefit from sharing a host, and; (2) instances where multiple C. semicircularis emerged from a single host were probably the result of parasitism by different females, or during different encounters by a single female.     

Preference and performance of the hyperparasitoid Syrphophagus aphidivorus (Hymenoptera: Encyrtidae): fitness consequences of selecting hosts in live aphids or aphid mummies

Abstract.  1. Theoretical models predict that ovipositional decisions of parasitoid females should lead to the selection of the most profitable host for parasitoid development. Most parasitoid species have evolved specific adaptations to exploit a single host stage. However, females of the aphid hyperparasitoid Syrphophagous aphidivorus (Mayr) (Hymenoptera: Encyrtidae) display a unique and atypical oviposition behaviour by attacking either primary parasitoid larvae in live aphids, or parasitoid pupae in dead, mummified aphids.
2. In the laboratory, the correlation between host suitability and host preference of S. aphidivorus on the host Aphidius nigripes Ashmead parasitising the aphid Macrosiphum euphorbiae (Thomas) was investigated.
3. The relative suitability of the two host stages was determined by measuring hyperparasitoid fitness parameters (survival, development time, fecundity, sex ratio, and adult size of progeny), and calculating the intrinsic rate of population increase ( r _m). Host preference by S. aphidivorus females and the influence of aphid defence behaviour on host selection was also examined.
4. Hyperparasitoid offspring performance was highest when developing from hosts in aphid mummies and females consistently preferred this host to hosts in parasitised aphids. Although aphid defensive behaviour may influence host selection, it was not a determining factor. Ecological and evolutionary processes that might have led to dual oviposition behaviour in S. aphidivorus are discussed.     

Host Preference and Progeny Sex Ratio in a Solitary Koinobiont Mealybug Endoparasitoid, Anagyrus Pseudococci (Girault), in Response to its Host Stage

Anagyrus pseudococci (Girault) , a koinobiont endoparasitoid , is a potential biological control agent of the citrus mealybug Planococcus citri (Risso) . This paper examines the effect of host stage of the mealybugs on the parasitoid preference , development , survival and sex ratio of the offspring . Host susceptibility and preference tests showed that the parasitoid had a significant preference for the older host instars . Percentage parasitization increased 4.8 - fold in the adult stage , compared with the second - instar host , although the parasitoid developed and emerged successfully from all stages . Mean developmental time for male parasitoids was faster than for the females . There was some evidence of delayed development of males in young hosts . Higher percentage emergence of parasitoid was observed from older mealybugs . A. pseudococci exhibited maternal adjustment of sex ratio as a function of host size with an increased proportion of females with the increasing host size . Females maximized their fitness by adjusting offspring sex ratio in response to host size without evaluating the future host quality . This information is important for mass rearing of A. pseudococci and for evaluating it in a biological control programme for P. citri.     

The reproductive strategy of the gregarious parasitoid,Pteromalus puparum (Hymenoptera: Pteromalidae)

Summary Host size of Pteromalus puparum, a gregarious pupal parasitoid, shows a wide inter- and intraspecific variation. Experiments were made to study the regulation of the number and sex ratio of progeny per host by the parasitoid. The parasitoid could discriminate inter- and intraspecific size differences of the host and regulate the number of eggs according to the host size when a single female attacked the host. The sex ratio of progeny (proportion males) was about 0.1. The number of progeny laid by the female agreed with the energetically most efficient number og eggs in order to maximize total weight of progeny per host but not with the reproductively most efficient number of eggs to maximize the total fecundity of the progeny. The parasitoid laid smaller number of eggs in a half buried host, but the number was much larger than a half of those in a fully exposed host. When more than one female attacked a single host, the number and sex ratio of progeny per host increased with the number of females attacking the host, but the number of progeny per female decreased. The change of the sex ratio agreed with the prediction of the local mate competition model.     

Effect of expected offspring survival probability on host selection in a solitary parasitoid

Optimal Foraging Theory predicts that parasitoid females should optimize their host selection to maximize their lifetime fitness gain and parasitize the most profitable hosts. In particular, in solitary parasitoids, females should avoid superparasitism, at least when sufficient unparasitized hosts are available. However, when unparasitized hosts are scarce, they should prefer, among already parasitized hosts, those that provide the best survival probability to their progeny, which depends on the age and the developmental stage of the first parasitoid. To test this hypothesis in a solitary ectoparasitoid, Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae), we first assessed the survival probability of a second parasitoid according to the time elapsed since initial parasitism. We then analyzed the female selection behavior in patches containing a mixture of hosts parasitized over various time intervals. Our results showed that the older the opponent larva was, the lower the survival probability of the second parasitoid was. However, when the first individual had reached the prepupal stage, both individuals could complete their development. At this stage, the survival probability of the second parasitoid was surprisingly high but such individuals were reduced in size. Our study also showed that host acceptance by females was strongly correlated with the survival probability of their progeny when the first parasitoid was from 0 to less than 10 days‐old. When the first parasitoid had reached the prepupal stage, females usually rejected these hosts, although the survival probability of the offspring was quite high. This discordance between female host selection behavior and progeny survival probability is discussed.     

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.     

Sex ratio regulation in Coccygomimus turionella Linnaeus (Hymenoptera: Ichneumonidae) and its ecological implications

Abstract. 1. Sex regulation and its relevance to the ecology of a k-selected pupal parasitoid was investigated in Coccygomimus turionellae.
2. The ratio varied with host size, females predominating in large hosts.
3. Egg transfer experiments and comparative mortality rates among the progeny of virgin and inseminated females demonstrated that the phenomenon was due to parental behaviour.
4. The study of the mechanism confirmed Aubert's hypothesis of host size estimation with the addition that the host had to be exposed from the poles and had to contain the proper host kairomones.
5. Host size was highly correlated to parasitoid fecundity via influences on size and longevity but not the number of ovarioles per female or daily egg production.
6. Host encounters are more limiting than egg production and host size had no effect on host acceptance.
7. Sex regulation allows a maximal host encounter rate at the same time that it maximizes the reproductive potential of female progeny.
8. Low host density increased the production of female progeny resulting in more offspring searching for the limited resource. Concealed hosts and a high host density resulted in a shift towards a Fisher 1:1 sex ratio.     

Biology of Coeloides sordidator (Hymenoptera: Braconidae), a Possible Candidate for Introduction Against Pissodes strobi (Coleoptera: Curculionidae) in North America

Various aspects of the biology of Coeloides sordidator, an ectoparasitoid of pine weevils of the genus Pissodes, were studied. When reared in the laboratory on P. castaneus, females preferred to oviposit on third and fourth instar larvae in galleries, but ovipositions were also observed on second instars and prepupae. Pupae were not accepted as hosts. At 23°C and a 16 h day length, pre-imaginal development lasted 15-20 days. For each parasitoid population tested, between 8 and 28% of the individuals diapaused as prepupae in cocoons. The rate of diapause was not influenced by the photoperiod during larval development. In contrast, diapause was induced by a short day photoperiod on parents before and during oviposition and by low temperatures during the larval stage. Diapause termination was achieved by cold storage at 2°C for at least 8 weeks. Rearing of C. sordidator on P. castaneus under natural conditions showed that in the Swiss Jura it is mainly univoltine, with less than 10% of the parasitoids emerging for a second generation. When C. sordidator females were offered healthy P. castaneus larvae and larvae parasitized by the braconid endoparasitoid Eubazus semirugosus, there was no evidence that they were able to discriminate between healthy and parasitized hosts. The potential of C. sordidator as a biological control agent against the North American species P. strobi is discussed.     

Bionomics of Opius bellus (Hymenoptera: Braconidae), an endoparasitoid of Anastrepha fraterculus (Diptera: Tephritidae) in fruit-growing areas of Northwestern Argentina

Opius bellus is a neotropical larval-prepupal parasitoid known to attack the pestiferous fruit fly, Anastrepha fraterculus. Due to interest in the use of native parasitoids in forthcoming fruit fly biocontrol programmes in Argentina, O. bellus was colonised for the first time using laboratory-reared A. fraterculus larvae. A series of experiments were conducted to (1) best achieve an efficient parasitoid rearing by determining optimal larval host age, host:parasitoid ratio and host exposure time and (2) assess their potential as biological control agents by determining reproductive parameters. The most productive exposure regimen was: 7–9 d-old (early and middle third-instars) A. fraterculus larvae for 4 h at a 4:1 host:parasitoid ratio; this array of factors was sufficient to achieve the highest average adult emergence (48%) and an offspring sex ratio at equitable proportion. Increasing both host:parasitoid ratio further than 4:1 and the host exposure time beyond 4 h did not significantly enhance parasitoid female offspring yield. Females produced eggs for 29.5 ± 1.4 days. At 32 days of age, 50% of the females were still alive. The majority of the progeny were produced by females between 20 and 24 d-old. At 26°C, gross fecundity rate, net reproductive rate, intrinsic rate of increase and mean generation time were 20.7 ± 4.2 offspring/female, 9.6 ± 2.5 females/newborn females, 0.06 ± 0.01 females/female/day and 8.4 ± 0.2 days, respectively. The long lifespan and reproductive parameters suggest that this parasitoid species has suitable attributes for mass-rearing.     

寄主大小及寄生顺序对蝇蛹佣小蜂寄生策略的影响

寄主大小模型认为寄生蜂后代性比与寄主大小相关,寄生蜂倾向于在大寄主上产出更多雌性后代,在小寄主上产出更多雄性后代.探讨了以家蝇蛹为寄主时,蝇蛹佣小蜂后代产量和性比变化;单次寄生情况下,寄主大小及寄生顺序对寄生蜂后代性比等影响.结果表明,蝇蛹佣小蜂的产卵期为(8.93±3.34)d,单头雌蜂能产雌性后代(34.11±16.34)头和雄性后代(11.04±8.87)头,且雄性百分比为0.24±0.11.随成蜂日龄的增大,寄生蜂产生雄性后代的比率显著增加.蝇蛹佣小蜂在寄生家蝇蛹时,会优先选择寄生个体较大的蛹;在单次寄生的情况下,蝇蛹佣小蜂倾向于在较大的家蝇蛹内产出更多的雌性后代.     

A biological assessment of Apophua simplicipes (Hymenoptera: Ichneumonidae) as a parasitoid of the oblique banded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

Apophua simplicipes (Cresson) (Hymenoptera: Ichneumonidae) is a common parasitoid of the oblique banded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) in organically managed apple orchards in the southern interior of British Columbia, Canada. The biological characteristics of a laboratory colony of A. simplicipes were studied. When held at 15 and 25°C and provided with honey water, individual females survived an average of 60.6±6.1 and 29.8±4.7 days, oviposited 196.7±50.7 and 326.6±51.3 eggs and parasitized a total of 163.4±40.4 and 229.4±35.8 hosts, respectively. Females oviposited into first through fourth instar oblique banded leafrollers, with significantly more parasitism occurring in the first two instars compared to the third and fourth instars. No parasitoid larvae survived past the first larval stage in parasitized fourth instar hosts. Apophua simplicipes did not parasitize larvae of three-lined leafroller, Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) which is sympatric with oblique banded leafrollers in orchards in the southern interior of British Columbia. Female predation and host feeding from wounds on early instars of both leafroller species was observed under laboratory conditions. In addition, early instar hosts exited diet feeding sites in response to the probing activity of the ovipositing wasps. A similar escape reaction in the orchard may cause a leafroller larva to move away from its feeding site, making it more vulnerable to predation or movement off the tree. Apophua simplicipes larvae emerged from fifth and sixth instar hosts. Parasitized oblique banded leafroller hosts consumed significantly less meridic diet than unparasitized female larvae from fifth instar through to parasitoid emergence or leafroller pupation. Our laboratory results suggest that A. simplicipes may reduce field populations of oblique banded leafroller and decrease pest feeding damage.     

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

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Latent infection: a low temperature survival strategy in steinernematid nematodes

1. Entomopathogenic nematodes penetrate and kill Galleria mellonella within 48 h at optimal temperatures.

2. Low temperature induces infection latency, preventing host death until optimal conditions resume.

3. Infected Galleria survived 25 days at 5°C. On transfer to 25°C, 100% and 12.5% of Steinernema carpocapsae and Steinernema riobravis infected larvae died within 72 h.

4. Infective juvenile penetration decreased with decreasing temperature; declining from 49.7 and 49.3 nematodes/host at 25°C to 6.3 and 0.25 nematodes/host at 5°C for S. carpocapsae and S. riobravis, respectively.

5. Latent infection occurs, albeit infrequently, due to low host penetration at low temperature.

Effects of host age and host availability on developmental period,adult size,sex ratio,longevity and fecundity inLariophagus distinguendus Förster (Hymenoptera: Pteromalidae)

The effects of host age on parasitoid reproductive capacity are studied using the pteromalid parasitoid Lariophagus distinguendusFörster and its bruchid hosts, Callosobruchus chinensis (L.) and C. maculatus (F.). A series of experiments were performed to investigate relationships between age and size of host parasitized and the developmental period of pre-imaginal progeny, sex ratio, female size, longevity, fecundity and oviposition rate. There was no effect of host size on preimaginal parasitoid developmental period. Sex ratio varied from less than 5% females from young (small) hosts to 60% females from mature (large) hosts. Adult size, female longevity, fecundity, and oviposition rate were also positively related to host age. Females provided mature hosts lived longer than those provided either young hosts or no hosts, possibly because of an increased ability to host-feed from the larger hosts. The implications of these findings to parasitoid population reproductive capacity and host-parasitoid synchrony are discussed.     

Reproductive strategies under multiparasitism in natural populations of the parasitoid wasp Nasonia (Hymenoptera)

Parasitoid Nasonia wasps adjust their progeny sex ratio to the presence of conspecifics to optimize their fitness. Another trait under female control is the induction of offspring diapause. We analysed progeny sex ratios and the proportion of diapausing offspring of individual Nasonia females in host patches parasitized by two species, Nasonia vitripennis and Nasonia giraulti, in North American field populations using microsatellite fingerprinting. Both Nasonia species produced similar sex ratios on hosts that were co‐parasitized by their own species as by the other species, indicating that females do not distinguish between con‐ and heterospecific clutches. The sex ratios of the diapause and adult fractions of mixed broods from single females were not correlated. We found further indications that N. vitripennis females take the emergence time of the offspring into account in their sex allocation. The reproductive strategies of Nasonia under multiparasitism are largely adaptive, but also partially constrained by information.     

Host-size-dependent feeding behaviour and progeny sex ratio of Anisopteromalus calandrae (Hym., Pteromalidae)

Host-size related feeding and oviposition behaviour, and allocation of progeny sex by Anisopteromalus calandrae (Howard) were tested on Sitophilus oryzae L. The parasitoid showed a host-size-dependent partition of feeding and oviposition behaviour, preferring small hosts for feeding, but large hosts for oviposition. Neither the mutual interference nor the host density showed any effect on the behaviour of the parasitoid. Allocation of progeny sex by the female parasitoid appeared to be based more likely on absolute than on relative host size encountered. A model for the progeny sex ratio was constructed based on: (1) ovipositional preference of the parasitoid on large hosts; (2) feeding preferentially on small hosts; and (3) host-size-related regulation of progeny sex ratio. The progeny sex ratio of the parasitoid predicted by the model was in close agreement with the observed value.     

Oviposition Behavior and Conspecific Host Discrimination in Diachasmimorpha longicaudata (Hymenoptera: Braconidae), a Fruit Fly Parasitoid

We studied conspecific host discrimination and oviposition behavior of Diaschasmimorpha longicaudata (Ashmead), in third instar Anastrepha ludens (Loew) under laboratory conditions. The complete process of oviposition in D. longicaudata required an average of 29±11.7 s (Mean±SE), during which time the female remained completely immobile. This contrasts with attempts to oviposit for lesser durations (2-4 s), during which the female constantly moved her antennae and abdomen. In order to determine the host discrimination ability of this species (i.e., the capacity to distinguish between parasitized and non-parasitized hosts), third-instar A. ludens non-parasitized or parasitized 24 h earlier, were exposed simultaneously to individual female wasps with or without previous oviposition experience. An identical test was performed using larvae parasitized 48 h earlier. Both types of females showed a similar behavioral pattern of oviposition, but with significant differences in the number of eggs laid in parasitized and non-parasitized larvae. Experienced females showed reduced incidence of oviposition in parasitized larvae as well as a greater number of probes that did not lead to parasitism (rejection), although this difference was only significant in the 24-h test. This suggests that the host discrimination capacity of this parasitoid is innate and that previous oviposition experience increases the discrimination ability of females.     

Effect of host size on male fitness in the parasitoid wasp Dinarmus basalis

The effect of host size on male fitness was tested in the parasitoid wasp Dinarmus basalis (Hymenoptera, Pteromalidae) using hosts of different fresh weight. Fitness was measured as the sperm stock in seminal vesicles, and the ability to access females in single or competition situations. Both body size and sperm in seminal vesicles increased with host fresh weight. Males from small hosts had a reduced size and sperm stock compared to those from larger hosts. In single situations, males from both small and large hosts had similar reproductive capacities, whereas in multiple mating or competition situations, males from small hosts were at a disadvantage, inseminating fewer females and copulating less frequently. However, females did not appear to choose between males, and no effect on sperm stored in the spermatheca was observed. Being small does not prevent a D. basalis male mating and producing progeny in single situations, although more offspring could be expected from larger males because of their better competitive abilities.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

The biology of Diadromus collaris (Hymenoptera: Ichneumonidae), a pupal parasitoid of Plutella xylostella (Lepidoptera: Plutellidae), and its interactions with Oomyzus sokolowskii (Hymenoptera: Eulophidae)

The ichneumonid Diadromus collaris (Gravenhorst) has been recorded in many parts of the world as an important parasitoid of the diamondback moth, Plutella xylostella (Linnaeus), a serious pest of brassica vegetable crops worldwide. Some aspects of its biology and its interactions with Oomyzus sokolowskii (Kurdjumov), another major parasitoid of the same pest, were studied in the laboratory. At 25 degrees C, female wasps did not have mature eggs in their ovaries until about 12 h after emergence. Both males and females mated successfully 24-48 h after emergence, and females started to oviposit one to two days after emergence. Unmated females produced male progeny only; mated females produced progeny of both sexes. The development rate of the parasitoid increased linearly with temperature from 15 to 30 degrees C, with an estimated low temperature threshold of 7.4 degrees C and a thermal constant of 225.1 day-degrees for development from egg to adulthood. Rates of survival from larva to adulthood were about 90% between 20 and 28 degrees C and decreased as temperature decreased or increased. No immatures survived to adulthood at 35 degrees C. When provided with honey solution, the females lived on average 8.3, 11.5 and 7.0 days, and parasitized 26, 44 and 46 host pupae at 20, 25 and 30 degrees C, respectively. Female wasps could be stored at 15 degrees C for up to four weeks without detrimental effects on reproduction. Females of D. collaris attacked host pupae already parasitized by O. sokolowskii, inserting their ovipositor into the hosts at a similar frequency as into unparasitized host pupae, but they did not lay eggs inside the hosts.