Egg Carrying Attracts Enemies in a Cryptic Coreid Bug (Phyllomorpha laciniata )

In the golden egg bug (Phyllomorpha laciniata) eggs are laid mainly on the backs of conspecifics, and in many habitats eggs do not survive unless carried by bugs. Bugs are covered with small spines that may make them unpalatable. They are also cryptic, at least if not carrying eggs. We used domestic chicks as predators to examine if egg carrying influences susceptibility to avian predators. The special morphology of the bugs and/or possible chemical defense may make the bugs unattractive, as chicks that picked up bugs often rejected them. Eggs made bugs more attractive to chicks. The total number of attacks and the probability of being attacked at all increased significantly when bugs carried eggs. If mating with an egg-carrying bug, a female without eggs suffered as much as her egg-carrying partner when attacked. This study, together with previous results on ant predation, suggests that carrying eggs as well as mating with an egg-loadedbug are costly in terms of predation risk.     

Flexible oviposition behavior in the golden egg bug (Phyllomorpha laciniata) and its implications for offspring survival

Female golden egg bugs show unique flexibility in their oviposition behavior. Females can lay eggs on plants, where they are leftunattended, or on the back of conspecifics, where they remainuntil hatching. In this article we show that eggs have greatersurvival rates when carried by an adult than when laid on plants.The main causes of egg mortality are predators and a parasitoidwasp. Our results suggest that, although predation pressureis similar, fewer eggs are attacked by parasitoid wasps whencarried by an adult than when laid on plants. In addition,we show that, when given a choice, females prefer to lay eggson the backs of conspecifics than on plants. Thus, female ovipositionchoice is adaptive and minimizes individual offspring mortality.The factors that may maintain such behavioral variation in current populations are discussed.     

Copulation Is Not a Prerequisite to Male Reception of Eggs in the Golden Egg Bug Phyllomorpha laciniata (Coreidae; Heteroptera)

Females of the golden egg bug, Phyllomorpha laciniata, lay eggs on the backs of conspecifics. Male response to female oviposition attempts is either passive or resistant. Passive males remain still during oviposition, while resistant males move repeatedly and thereby delay or avoid being encumbered. We investigated how previous mating experience affected male acceptance of eggs. Males from two Spanish populations, from Andalusia and Catalonia, were allowed to mate with single females repeatedly. These experimental pairs were then presented with either a single, nonmating male or a pair in copula. We expected mating partners to resist oviposition attempts less strongly than nonmating males but no differences were detected. However, there was a significant difference in behavior exhibited by males from the two populations; males from Catalonia were relatively passive but males from Andalusia routinely resisted encumbrance. Predation pressure and the availability of receptive females may explain the observed differences between the populations.     

Adjustment of copula duration and ejaculate size according to the risk of sperm competition in the golden egg bug (Phyllomorpha laciniata)

Several hypotheses have been proposed to explain the adaptivesignificance of prolonged copulations in insects, which includemate guarding and sperm loading functions. We have exploredthe adaptive significance of the prolonged copulations in thegolden egg bug (copulations up to 50 h) and the effect of anincreased risk of sperm competition on ejaculate investment.Our data support predictions derived from sperm competitiontheory, which posits that males are expected to increase ejaculateexpenditure in response to an increased risk of sperm competition.Results show a combined response by males that has not beenpreviously described: males in the presence of rivals increasecopulation duration and the rate of sperm transfer. No relationshipwas found between male or female size and copulation durationor ejaculate size. Golden egg bug males transfer sperm slowlyand gradually throughout copulation; thus an increase in theamount of sperm transferred and the corresponding increase inthe male's numerical representation in the female's storageorgans could be particularly important in a system in whichso few sperm are transferred and in which so few sperm are storedby females. In addition, copulation duration may not only serveto increase the total amount of sperm transferred, but it mayalso increase the chances that the female will lay an egg soonafter copulation has ended. This could explain why males tendto accept eggs after copulation, since they could be maximizingthe chances that such eggs are fathered by them, and in thisway they would substantially increase the survival rates oftheir offspring because eggs laid on plants suffer high mortalityrates.     

Fitness of two sibling species of Asobara (Braconidae:Alysiinae), larval parasitoids of Drosophilidae in different microhabitats

Abstract. 1. Two sibling species of larval endoparasitoids of Drosophilidae: Asobara tabida (Nees) and A.rufescens (Foerster) occur in the same macrohabitat, but inhabit different microhabitats. Each species is most attracted by odours of its own microhabitat.
2. In order to assess the adaptive value of the microhabitat preference we studied the survival of both parasitoids in the major host species that occur in these microhabitats.
3. Survival in the major host in the preferred microhabitat was shown to be 40% higher for A.tabida and 30% higher for A.rufescens when compared to survival in the major host in the non-preferred microhabitat.
4. Measurements of developmental rates, specific mortalities and dry weights of the parasitoids suggest that the differential survival is due to differences in synchronization with the hosts.
5. The possible evolutionary consequences of some biological characteristics in Asobara are discussed. Microhabitat selection, differential survival, development and mating behaviour are attributes likely to have played a role in the reduction of gene flow between populations of the ancestral species, either in primary or in secondary sympatry.     

Ant association facilitates the evolution of diet breadth in a lycaenid butterfly

The role of mutualistic interactions in adaptive diversification has not been thoroughly examined. Lycaenid butterflies provide excellent systems for exploring mutualistic interactions, as more than half of this family is known to use ants as a resource in interactions that range from parasitism to mutualism. We investigate the hypothesis that protection from predators offered to caterpillars by ants might facilitate host-range evolution. Specifically, experiments with the butterfly Lycaeides melissa investigated the role of ant association in the use of a novel host, alfalfa, Medicago sativa, which is a sub-optimal host for larval development. Survival on alfalfa is increased by the presence of ants, thus supporting the hypothesis that interaction with ants might be important for host-range evolution. Using a demographic model to explore ecological conditions associated with host-range expansion in L. melissa, we conclude that the presence of ants might be an essential component for populations persisting on the novel, sub-optimal host.     

Increased reproduction by pea aphids in the presence of secondary parasitoids

1. The influence of the presence of secondary parasitoids on aphid reproduction was tested in a field experiment. 2. Single pea aphids Acyrthosiphon pisum in clip cages on potted bean plants were exposed to a collection of secondary parasitoids enclosed within a small perforated bag. The aphids were exposed to volatile chemicals released by the secondary parasitoids but there was no physical contact. 3. The production of nymphs was recorded over an 11‐day period. Aphids produced significantly more offspring in the presence of secondary parasitoids than in the absence of secondary parasitoids (36.9 vs. 31.2). 4. This is the second reported occurrence of this phenomenon, and the reasons for its occurrence and its consequences for aphid population and community ecology are discussed.     

Body size variation in <Emphasis Type="Italic">Gryon gallardoi</Emphasis> related to age and size of the host

This study aimed to evaluate the influence of Spartocera dentiventris (Berg) (Hemiptera: Coreidae) egg quality (regarding age, size and superparasitism) on male and female body size of Gryon gallardoi Brèthes (Hymenoptera: Scelionidae). It was also analysed host size influence on offspring sex choice and female oviposition order (within egg groups). Groups of 12 eggs, 2, 3, 4, 5, 6, 7, 8, and 12 days old were individually exposed to a female parasitoid for 2 h, and the order of each egg parasitisation was recorded. Size of adult parasitoids (head width and tibia length of the second leg pair), and parasitised and superparasitised host eggs were measured. Sexual dimorphism related to size (SDs), was estimated through a model II linear regression. Females have a larger head width but a smaller tibia length. Parasitoid adult size is significantly related to host age and size. Within an egg group, females did not exhibit preference for any egg size category either to start oviposition or to oviposit male or female eggs. The body size of both sexes diminished with host aging. There was a positive response in both sexes to host size increasing. Estimated SDs diminished with host aging and increased with host volume augmentation. The results suggest that adult body size of G. gallardoi, as well as size differences between males and females, are strongly associated to host quality.     

Oviposition of resistant and susceptible strains of Drosophila melanogaster in the presence of deltamethrin

Oviposition of three strains of Drosophila melanogaster in the presence of deltamethrin was observed. These strains had different levels of physiological susceptibility to deltamethrin. Two-choice tests were conducted with couples of flies in petri-dish arenas containing two oviposition dishes. On the first day of the experiment, females were given a choice between a treated oviposition dish and an untreated control dish. On the second day of the experiment, two control oviposition dishes were given to females. Although individual females showed a tendency to aggregate their eggs in one of the dishes, control experiments demonstrated an overall equal distribution of eggs between the dishes. When one of the two oviposition dishes in the arena was treated with deltamethrin, the percentage of females ovipositing and the mean number of eggs laid by females were reduced, compared with control arenas. Females avoided the treated oviposition dish and laid significantly more eggs on the control dish. Furthermore, when the deltamethrin concentration was increased on the first day, female flies postponed their oviposition and laid significantly more eggs on the second day. The resistant strain, SR, demonstrated the same capacity to select the untreated site for oviposition as the susceptible strain, but it showed a smaller oviposition reduction and egg retention. The relationship between physiological and behavioural susceptibility to deltamethrin is discussed.     

Adaptive superparasitism in solitary parasitoids: marking of parasitized hosts in relation to the pay-off from superparasitism

Abstract.

• 1 The pay-off from an egg laid in a parasitized host is an important parameter in models on adaptive superparasitism in solitary insect parasitoids.
• 2 For Leptopilina heterotoma, a parasitoid of larval Drosophila, the pay-off from a second egg laid in a host is 0.43 offspring when the interval between the two ovipositions is less than 3h. For longer intervals, this pay-off decreases to almost zero for an interval of 24 h.
• 3 When a female encountering a parasitized host is able to estimate the interval since the first oviposition, it is expected that she will take this into account in her host selection decisions. This is, however, not in the direct interest of the female that lays the first egg, and marks the host.
• 4 We studied whether superparasitism in hosts containing a young egg is more common than in hosts containing an older egg, when searching in a patch containing once-parasitized and unparasitized hosts.
• 5 The acceptance/encounter ratio of parasitized hosts increased for intervals longer than 6h, as predicted when the interests of the marking female and the longevity of the mark are taken into account.
• 6 Superparasitism occurred more often when parasitoids had previously searched a host patch 7 days before the experiment compared to when parasitoids had searched a patch 1 day before, a phenomenon predicted by dynamic optimal diet models.

     

Low parasitoid success on a myrmecophilous host is maintained in the absence of ants

Abstract.  1. Studies of Dinocampus coccinellae , a parasitoid of ladybird beetles, have generally shown congruence between field parasitism rates of different host species and parasitoid preference and/or host suitability in the laboratory, suggesting that host intrinsic factors rather than habitat-related extrinsic factors are of greatest importance in determining D. coccinellae occurrence.
2. The myrmecophilous Coccinella magnifica exhibits much lower D. coccinellae prevalence in the field than most other Coccinella species: it has been suggested that this is a manifestation of enemy-free space provided by the predatory Formica rufa group ants with which the C. magnifica occurs.
3. Coccinella magnifica collected at the same time and locality as parasitised Coccinella septempunctata were unparasitised by D. coccinellae . In the laboratory, in the absence of ants, although the parasitoid attacked C. magnifica as readily as C. septempunctata , C. magnifica was not parasitised successfully.
4. Such results are consistent with those from other ladybirds and C. magnifica does not now benefit directly from any putative D. coccinellae -free space provided by aggressive ants. Because its close relatives exhibit high levels of D. coccinellae parasitism, C. magnifica may be useful in determining some elements important in the evolution of host protection against parasitoid attack.     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non-significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

Genetic architecture of population differences in oviposition behaviour of the seed beetle Callosobruchus maculatus

Few studies have examined the genetic architecture of population differences in behaviour and its implications for population differentiation and adaptation. Even fewer have examined whether differences in genetic architecture depend on the environment in which organisms are reared or tested. We examined the genetic basis of differences in oviposition preference and egg dispersion between Asian (SI) and African (BF) populations of the seed beetle, Callosobruchus maculatus. We reared and tested females on each of two host legumes (cowpea and mung bean). The two populations differed in mean oviposition preference (BF females preferred cowpea seeds more strongly than did SI females) and egg dispersion (SI females distributed eggs more uniformly among seeds than did BF females). Observations of hybrid and backcross individuals indicated that only the population difference in oviposition preference could be explained by complete additivity, whereas substantial dominance and epistasis contributed to the differences in egg dispersion. Both rearing host and test host affected the relative magnitude of population differences in egg dispersion and the composite genetic effects. Our results thus demonstrate that the relative influence of epistasis and dominance on the behaviour of hybrids depends on the behaviour measured and that different aspects of insect oviposition are under different genetic control. In addition, the observed effect of rearing host and oviposition host on the relative importance of dominance and epistasis indicates that the genetic basis of population differences depends on the environment in which genes are expressed.     

Egg distribution patterns in the almond seed wasp,Eurytoma amygdali

The egg distribution patterns ofEurytoma amygdali Enderlein (Hymenoptera, Eurytomidae), which oviposits singly in green, developing almonds, were studied in the laboratory and in the field. In the laboratory, individual females were caged with a number of almonds and the eggs deposited in each fruit were counted. In the field, eggs were censused in almonds of different varieties at regular intervals, over four seasons (1988–91). In the susceptible ‘Retsou’ variety, eggs were uniformly distributed among fruits, both in the laboratory and in the field, as long as the mean number of eggs per almond was ≤2.5. When the mean number of eggs per almond was higher the egg distributions were random. This suggests that, up to a certain level of infestation, females were able to assess egg load of fruits and oviposit in the less infested ones. The main factor enabling the females to discriminate and select the less infested almonds for oviposition is probably a host-marking pheromone. In 1990, the same patterns of egg distribution were observed in samples taken from 5 other almond varieties that are not as susceptible as Retsou and have thicker pericarp and harder endocarp. In 1991 however, when fruits developed more rapidly than in 1990, egg distributions in 3 of these 5 varieties (Truoito, Ai, and Marcona) were not uniform. Although the mean number of eggs per fruit was low (1.2–2.0), many fruits of these varieties contained no eggs. This suggests that, in some less susceptible varieties, egg distribution might also be affected by certain fruit parameters, unfavourable for oviposition, related to the pericarp thickness and endocarp hardness.     

A test of temporal and spatial density dependence in the parasitism rates of introduced parasitoids on host, the arrowhead scale (Unaspis yanonensis) in stable host–parasitoids system

Abstract: In 1980, two species of parasitoid wasps (Aphytis yanonensis DeBach et Rosen and Coccobius fulvus Compere et Annecke) were introduced to Japan from China as biological control agents to combat the arrowhead scale (Unaspis yanonensis Kuwana). These introductions represent one of the most successful projects in the history of biological control in Japan. To examine whether density dependent parasitism was inevitable for success of biological control, we tried to detect temporal and spatial density dependence in parasitism rates using time‐series data of scale density, as well as parasitism, over a 16‐year period. The work was conducted in a Satsuma mandarin orange (Citrus unshiu Marc.) orchard in which we previously demonstrated that the system appeared to have stabilized after a decline in scale density following the introduction of the parasitoids. Earlier work also indicated that C. fulvus contributes most to the reduction in, and the stability of, scale density. In this study, we examined: (1) the relationship, on a whole‐orchard basis, between scale density and the rates of parasitism by A. yanonensis, C. fulvus, and a combination of the two species; (2) whether parasitism was positively correlated to scale density on a single‐tree basis among generations and (3) whether spatial density dependence was detectable within generations on an individual‐tree basis. Parasitism by A. yanonensis was temporally density‐dependent on scale population density at the whole‐orchard level, while parasitism by C. fulvus was not. Parasitism by A. yanonensis or by C. fulvus was rarely positively correlated to scale density at the single‐tree level, and spatial density‐dependence was hardly detected at all at this level. Most analyses of combined parasitism rates were similar to rates of parasitism by C. fulvus alone. Contrary to conventional wisdom of biological control theory, this study demonstrates that density dependence is not necessarily detected, even in a system in which a natural enemy has long held pest density stable at low levels.     

Two new egg parasitoids (Hymenoptera: Encyrtidae) of the wood borer Agrianome spinicollis (Macleay) (Coleoptera: Cerambycidae), a pest of pecans in eastern Australia

Abstract Baeoanusia xanthopleuron sp. n. and Avetianella coombsi sp. n. (Hymenoptera: Encyrtidae) are described from New South Wales and Queensland. Photomicrographs are provided to illustrate morphological characters of both species, and their systematic position is discussed on the basis of their morphology and biology. Both species are egg parasitoids of the wood borer Agrianome spinicollis (Macleay), an important pest of pecans in eastern Australia.     

Host suitability of the New World stalkborer Diatraea considerata for three Old World Cotesia parasitoids

Biological control ofstalkboring Lepidoptera often has beensuccessful when the braconid parasitoids in thegenera Cotesia and Apanteles werethe natural enemies of choice. Constraints inusing these gregarious, koinobiont,endoparasitoids have included host suitability,especially as influenced by the host's immuneresponse. The suitability of a novel host, theNew World stalkborer Diatraeaconsiderata (Lepidoptera:Pyralidae), for parasitization by three OldWorld braconids, Cotesia chilonis, C. flavipes and C. sesamiae (Hymenoptera: Braconidae), wascompared to the suitability of another NewWorld novel host, Diatraea saccharalis. D. considerata was less suitable for allthree parasitoids than was D.saccharalis. The frequent occurrence ofparasitized D. considerata larvae thatdid not yield parasitoids, or pupate within anappropriate time interval, suggestedencapsulation of the parasitoid progeny, whichwas visible through the host cuticle. Giventhe suitability results, these threeparasitoids would not be appropriate candidatesfor use against D. considerata. Theresults also have important implications forthe narrow host range expressed by theseparasitoids.