Aggregative oviposition varies with density in processionary moths—Implications for insect outbreak propensity

1. In gregarious insects, groups commonly originate from females laying eggs in masses and feeding groups are established as soon as larvae hatch. Some group-living insect species may aggregate beyond the individual parent level, such that offspring from two or more egg masses develop within a common resource.
2. Here we show that aggregative oviposition can vary with population density at oviposition and possibly be an important factor in outbreak dynamics of phytophagous insects.
3. We analysed density data with respect to egg mass aggregation for two species of pine processionary moths, Thaumetopoea pinivora (in Sweden 2005–2019) and T. pityocampa (in Spain 1973–1991). Both species lay their eggs in egg masses and feed in groups. During the study periods, insect population density for both species varied by at least an order of magnitude.
4. The two species showed strikingly similar patterns of egg mass aggregation. Egg masses were overdispersed at high population density, with few trees showing a high load of egg masses.
5. Our data suggest that aggregative oviposition can be important in explaining the previously documented higher propensity for outbreaks in insects laying eggs in clusters, compared with those laying individual eggs.

     

Caddisfly egg mass morphology mediates egg predation: potential costs to individuals and populations

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Life history of a dipteran predator (Scathophagidae: Acanthocnema) of insect egg masses in a northern California stream

1. Little is known about the predators of insect eggs in fresh waters. This study describes aspects of the life history of a scathophagid fly (Acanthocnema sp.), whose larvae are predators of aquatic insect eggs. 2. Because the Acanthocnema predator oviposits its eggs on the surface of aquatic insect egg masses, all insect egg masses were collected regularly within a 200‐m reach of Redwood Creek (California, U.S.A.) between September 2003 and June 2007. Acanthocnema predators were found predominantly within egg masses of the caddifly Neophylax rickeri (Trichoptera: Uenoidae). 3. There was a mean of 0.25 Acanthocnema individuals per N. rickeri egg mass (n = 2367 egg masses). In general, N. rickeri egg masses were more commonly found clustered in aggregations (93.7%) than singly (6.3%), and Acanthonema were found more often within the aggregations of N. rickeri (98.7%) compared to singly laid egg masses (1.3%). 4. The duration of the Acanthocnema predator life stages was: egg 2.9 ± 0.8 (mean ± SD) days, larva 15.6 ± 10.2 days, pupa 80.3 ± 24.9 days and adult 7.2 ± 4.8 days. The short duration of the Acanthocnema egg stage (1–7 days) compared to that of its prey N. rickeri (2–4 weeks) raises the probability that the undeveloped eggs of N. rickeri would be available to the young predators upon hatching. Egg consumption of N. rickeri eggs by Acanthocnema averaged 262.6 eggs per larval period. 5. Acanthocnema had a bivoltine life cycle in which the first generation fed exclusively on N. rickeri egg masses in the winter and the second generation fed on the egg masses of several species, including other Trichoptera (Brachycentridae) and Diptera (Ceratopogonidae, Chironomidae) in the spring. These findings suggest differing feeding strategies by the first and second generations of Acanthocnema in response to the seasonal availability of prey species. This type of autecological information is important for understanding mechanisms of community interactions.     

Environmental cues or conspecific attraction as causes for egg mass aggregation in hydrobiosid caddisflies

The distribution of egg masses throughout the environment can strongly influence the population dynamics of aquatic insects. Many species lay eggs in aggregations and most eggs will subsequently hatch from only a few locations—knowing how and why these aggregations arise is therefore needed to understand the population dynamics of these species. The hydrobiosid caddisfly Ulmerochorema rubiconum lays eggs in large aggregations on the undersides of emergent rocks in streams. Our aim was to test whether females oviposit in response to conspecific egg masses or the environmental characteristics of rocks by manipulating the number and age of egg masses initially present on rocks and monitoring the accumulation of new masses. Our first experiment used rocks that had recently been used for oviposition and initial egg masses encompassed a range of ages; our second used rocks with no history of oviposition and initial masses of uniform age. Females did not respond to conspecific eggs of any age, as removing initial egg masses had no impact on the number of new masses laid in either experiment. There was a strong positive relationship between the number of initial and new masses, regardless of whether initial egg masses were removed, at the scale of whole rocks and for rock microhabitats. The results suggest that females select oviposition sites based primarily on physical characteristics of emergent rocks. We demonstrate for the first time that the spatial arrangement of egg masses may be consistent and predictable through time, with females showing a high degree of fidelity for particular rocks and microhabitats.     

Evidence for embryo–embryo interactions in controlling hatching time and synchronization in the brown marmorated stink bug,Halyomorpha halys (Hemiptera: Pentatomidae)

The mechanisms controlling egg diapause and circadian rhythms of hatching activity have been extensively studied in insects. However, relatively little attention has been paid to the mechanisms controlling synchronized hatching from an egg mass. In this study, we examined the possible involvement of embryo–embryo interaction in controlling hatching time in Halyomorpha halys (Stål). Eggs tended to hatch earlier as the egg mass size increased. Egg separation and clumping of separated eggs at various times showed that hatching synchrony was largely determined shortly before hatching. However, whether eggs were kept in a mass or separated until several hours before hatching also influenced the hatching time, indicating the presence of embryo–embryo interactions. Eggs derived from different masses and kept in physical contact with one another hatched synchronously if their ages were within approximately 8 h. In this case, both younger and older eggs advanced only in hatching time, in contrast to a case of locusts reported by others. Eggs separated by more than 7 mm hatched as synchronously as those kept in a mass when glued to the same substrate, suggesting an important role of the egg substrate in transmitting the vibrational hatching signals to neighboring sibling eggs to synchronize hatching.     

Egg mass destroying behaviour of the female giant water bugLethocerus deyrollei Vuillefroy (Heteroptera: Belostomatidae)

After being laid on emergent aquatic vegetation, the egg masses ofLethocerus deyrollei are brooded by the male. In laboratory studies, females were observed to destroy egg masses and ingest the fluid of eggs. Brooding males fought with these attackers at first, but then gave up the defense and mated with them. After destroying the egg masses, females laid new ones at the same sites on the same night or the following one, and males brooded the new offspring. The population density of this species is very low. Most males engage in brooding and cease to become acceptable mates after most of the females lay the first egg masses. It is costly for females to search out free males in the aquatic vegetation. By destroying egg masses, however, individual females can gain new mates with little exertion and can make them brood their own offspring. Furthermore, the survival of their own offspring increases with the elimination of their competitors.     

Protective function of maternal care against egg predators in the shield bug Sastragala esakii (Hemiptera: Acanthosomatidae)

Females of many acanthosomatid shield bugs attend eggs and young nymphs by covering them with their bodies. Although such form of maternal care has been suggested to have evolved independently in four genera, Elasmucha, Sastragala, Acanthosoma and Sinopla, previous studies exploring its adaptive function have solely focused on species of Elasmucha. This study pioneered an experimental examination of maternal care in the Japanese species Sastragala esakii. Field experiments demonstrated that unattended egg masses suffered intense predation, whereas egg masses attended by their mothers were rarely preyed upon. The ant Crematogaster sp. was the most common egg predator, while two other insect species were also observed to prey on eggs. The exclusion of ant workers and other walking predators from accessing egg masses through the utilization of a sticky trap resulted in a remarkably improved egg survival rate in orphaned egg masses, suggesting that, at least in our study site, maternal care of S. esakii primarily functions to protect eggs from walking predators. Laboratory observations revealed that egg protection against predators was achieved through at least two distinctive defensive behaviors shown by egg-tending females: (i) tilting their bodies when approached by predators; and (ii) fanning their wings when in contact with predators. However, females displayed a limited response to predators approaching from the posterior direction, suggesting that visual cues play a significant role in predator recognition. These results indicate a similarity in the maternal care functions between Elasmucha and Sastragala, and suggest the parallel evolution of female defensive behaviors.     

Oviposition strategies employed by the western spruce budworm: tests of predictions from the phylogenetic constraints hypothesis

1 Predictions from the Phylogenetic Constraints Hypothesis were tested for the first time in an eruptive forest Lepidopteran species, the western spruce budworm. 2 In previous work, we established that western spruce budworm females exhibit oviposition preferences with regard to tree age, tree vigour and host species. However, there was no evidence to support a link between oviposition preference and larval performance, which supports the Phylogenetic Constraints Hypothesis. 3 Our preference data led us to test whether female budworms use oviposition strategies to select the sites where they lay their egg masses. Our experiments were designed to make direct comparisons between latent and eruptive insect herbivores with respect to two oviposition behaviours: egg retention and avoidance of conspecifics. This type of research has not previously been conducted on any eruptive forest Lepidopteran. 4 Female budworms retained eggs instead of laying them on less preferred hosts in two of three experiments, but the percentage of eggs they retained was significantly less compared to latent insect herbivores. 5 In addition, female budworms actively avoided oviposition in areas with the highest density of conspecific egg masses, but they laid egg masses in all the other locations provided. This contrasts with the pattern seen in latent insect herbivores, which consistently avoid laying their eggs near any sites already used by conspecifics. 6 Our research indicates that there are extreme differences between latent and eruptive insect herbivores with respect to egg retention and avoidance of conspecifics, thus supporting the Phylogenetic Constraints Hypothesis.     

A test of the preference–performance hypothesis with stream insects: selective oviposition affects the hatching success of caddisfly eggs

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Variations in fecundity over catchment scales: Implications for caddisfly populations spanning a thermal gradient

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Significant mortality of eggs and young larvae of two pine processionary moth species due to the entomopathogenic fungus Metarhizium brunneum

Bioassays were conducted to determine the susceptibility of egg masses and young larvae of two pine processionary moth species, Thaumetopoea pityocampa and Thaumetopoea wilkinsoni, to two strains (ARSEF4556, V275) of the entomopathogenic fungus Metarhizium brunneum. Mortality of treated eggs by both strains ranged from 96% to 99% but not all of this was caused by M. brunneum since control groups also experienced egg mortality due to saprophytic fungi. Still, larvae hatched in the laboratory from eggs treated with M. brunneum were all killed by this fungus, acquiring M. brunneum conidia, whereas larval mortality was 0% in the control groups. Young larvae of both pine processionary moth species were also highly susceptible to ARSEF4556 and V275 with larval mortality ranging between 94% and 100%, 8 days post-inoculation, with the vast majority of larvae being killed within the first 2–4 days. Larval mortality was dose dependent. Results were consistent across the two pine processionary moth species, showing that the pathogenicity of M. brunneum to both eggs and young larvae might be promising for biological control of these insect pests. The study also showed that non-target parasitoids of pine processionary moth eggs were also susceptible to M. brunneum. Further work is required to understand and reduce the M. brunneum effect on non-target insects.     

Egg and time limitation mediate an egg protection strategy

The number of mature eggs remaining in the ovaries and the time left for oviposition determine the reproductive decisions of the hyperdiverse guild of insects that require discrete and potentially limiting resources for oviposition (such as seeds, fruits or other insects). A female may run out of eggs before all available oviposition sites are used (egg limitation), or die before using all of her eggs (time limitation). Females are predicted to change clutch size depending on whether eggs or time is the limiting resource. We extend this framework and ask whether the same constraints influence a strategy in which females modify eggs into protective shields. In response to egg parasitism cues, female seed beetles (Mimosestes amicus) lay eggs in vertical groups of 2–4, modifying the top 1–3 eggs into shields in order to protect the bottom egg from attack by parasitoids. We made contrasting predictions of how egg and time limitation would influence egg size and the incidence and level of egg protection. By varying access to seed pods, we manipulated the number of remaining eggs a female had at the time she received a parasitism cue. Although egg size was not affected, our results confirm that egg‐limited females protected fewer eggs and time‐limited females protected more eggs. Female body size explained the number of eggs in a stack rather than host deprivation or the timing of parasitoid exposure. Our results clearly show that host availability relative to female age influences the incidence of egg protection in M. amicus. Furthermore, our study represents a novel use of life history theory to explain patterns in an unusual but compelling defensive behaviour.     

EGG ACTIVATION AND PARTHENOGENETIC REPRODUCTION IN INSECTS

1. Many insects reproduce by parthenogenesis. In one of the largest orders of the animal kingdom, the Hymenoptera, most of its members reproduce by arrhenotokous parthenogenesis. Egg activation in parthenogenetic animals obviously cannot be caused by fertilization of the egg. The question of what initiates egg development in parthenogenetically reproducing animals has been studied for a few insect species and is discussed in this article. 2. The grasshopper Melanoplus differentialis is one of several Orthoptera displaying accidental parthenogenesis. In this species, egg laying provides the stimulus to the completion of meiosis and start of embryonic development in unfertilized and probably also in fertilized eggs. The same holds true for the dipteran insect Drosophila melanogaster which exhibits rudimentary parthenogenesis, and for D. mercatorum showing accidental parthenogenesis. The precise way in which oviposition affects the egg is unknown. 3. The stick insect Carausius morosus reproduces by obligatory thelytoky. The triggering factor for removal of the meiotic block and initiation of embryonic development is oxygen from the air which penetrates to the egg through the micropyle immediately after oviposition. The oviposition act itself is not necessary for activation of the egg. 4. Comparative studies of the different types of oogenesis in the dipteran insect Heteropeza pygmaea show that in paedogenetically developing follicles meiotic arrest in prophase is of very short duration and a meiotic block at the end of oogenesis is absent. It is suggested that in this case triggering events for egg development are dispensable. On the other hand, under certain experimental conditions a meiotic block can be established in some of these follicles. 5. Investigations on the Ichneumonid wasp Pimpla turionellae have shown that unfertilized, male-determined eggs - and most likely also fertilized, femaledetermined eggs - are activated by mechanical stress exerted on the eggs during natural or imitated oviposition. This mechanical stress, in addition, activates a streaming system which is independent of meiotic completion and nuclear multiplication. Egg activation by egg distortion is also found in the Pteromalid species Nasonia vitripennis and occurs presumably in many other Hymenoptera. 6. Carausius morosus, Pimpla turionellae and Nasonia vitripennis are species with parthenogenetic reproduction for which the natural factors responsible for the initiation of egg development have been identified. The cases of Pimpla turionellae and Nasonia vitripennis are of particular interest because of the feasibility of artificially imitating the natural activating mechanism. 7. It is concluded that apart from fertilization various events at oviposition may trigger egg development. In addition, the occurrence of rudimentary parthenogenesis in many sexually reproducing animal species suggests that sperm entry and fertilization may frequently be necessary for the continuation of egg development rather than for its initiation.     

Individual repeatability in laying behaviour does not support the migratory carry‐over effect hypothesis of egg‐size dimorphism in Eudyptes penguins

Penguins of the genus Eudyptes are unique among birds in that their first‐laid A‐egg is 54–85% the mass of their second‐laid B‐egg. Although the degree of intra‐clutch egg‐size dimorphism varies greatly among the seven species of the genus, obligate brood reduction is typical of each, with most fledged chicks resulting from the larger B‐egg. Many authors have speculated upon why Eudyptes penguins have evolved and maintained a highly dimorphic 2‐egg clutch, and why it is the first‐laid egg that is so much smaller than the second, but only recently has a testable, proximate mechanism been proposed. In most species of Eudyptes penguins females appear to initiate egg‐formation at sea during return migration to breeding colonies. In macaroni penguins E. chrysolophus, females with a shorter pre‐laying interval ashore (and thus presumably greater overlap between migration and egg‐formation) lay more dimorphic eggs, suggesting a physiological conflict may constrain growth of the earlier‐initiated A‐egg. This migratory carry‐over effect hypothesis (MCEH) was tested in eastern rockhopper penguins E. chrysocome filholi on Campbell Island, New Zealand, by recording the arrival and lay dates, body sizes, and egg masses of transponder‐tagged females over two years. Females with longer pre‐laying intervals laid less dimorphic clutches, as predicted by the MCEH. However, repeated measures of individual females revealed that within‐individual variation in egg‐size dimorphism between years was unrelated to within‐individual variation in pre‐laying interval. Egg masses, and to a lesser extent egg‐size dimorphism, were highly repeatable traits related to body size and body mass. These results and a detailed consideration of the MCEH suggest that egg‐size dimorphism in Eudyptes penguins is unlikely to be caused by a migratory carry‐over effect.     

Termite queens have disproportionately more DNA in their fat body cells: reproductive division of labor and endoreduplication

Increases in DNA content caused by endoreduplication are widely observed in the metabolically active tissues of plants and animals. During egg production, insect females synthesize very large amounts of vitellogenin in their fat bodies, and female fat bodies of some insects become polyploid to accelerate vitellogenin production. Social insects have developed reproductive division of labor, wherein queens lay most of the eggs while other individuals have reduced fertility and undertake tasks required for maintaining the colony. Therefore, only queens are engaged in vitellogenin synthesis for egg production in social insects. Here, we show that termite queens have disproportionately more DNA in their fat body cells. Our DNA content analysis using flow cytometry demonstrated that more cells contained 4C‐DNA than 2C‐DNA in the fat bodies of Reticulitermes speratus queens. This high level of endoreduplication was not found in the fat body cells of other castes. This caste‐dependent doubling of DNA content in fat body cells suggests that termites exploit endoreduplication to boost egg production, in conjunction with the development of reproductive division of labor. This study highlights nuclear polyploidization as an adaptive strategy in social insects.     

Silent assassins: predation of native New Zealand trichopteran eggs by non-native freshwater gastropods

Aquatic insects that oviposit on rocks may evolve strategies to mitigate loss of eggs to opportunistic predation by common native grazers, but such strategies may be ineffective against non-native grazers. We tested whether the eggs of common New Zealand caddisfly families (Hydrobiosidae and Hydropsychidae) that oviposit on rocks were more susceptible to predation by native or non-native snails. The native snail Potamopyrgus antipodarum (Grey, 1843) and non-native snails Physa acuta Draparnaud, 1805 and Pseudosuccinea columella (Say, 1817) were presented with caddisfly egg masses and the number of individual eggs consumed was recorded after 24 and 48 hours. The larger Pseudosuccinea readily consumed the eggs, especially spumaline-encapsulated Hydrobiosidae eggs. Physa consumed very few Hydrobiosidae eggs and no Hydropsychidae eggs, whereas Potamopyrgus did not consume eggs of either family. The trichopteran egg masses tested did not succumb to predation by the native Potamopyrgus but are highly vulnerable to predation by a larger non-native snail.     

Responses of native egg parasitoids to the invasive seed bug Leptoglossus occidentalis

1. The Western conifer seed bug Leptoglossus occidentalis, a native insect of North America, was accidentally introduced in Europe in the late 1990s. Since then, it has spread rapidly. Biological control could provide an efficient management option but natural enemies of the pest have been poorly examined in Europe.
2. In this study, we exposed sentinel egg masses and collected naturally laid egg masses in southern France in 2016 and 2017, to identify the egg parasitoids of L. occidentalis and investigate their potentials.
3. Three egg parasitoids were detected: Anastatus bifasciatus, Ooencyrtus pityocampae and Ooencyrtus telenomicida. The overall parasitism was low compared to that observed in the native range with 6.4% of all eggs being parasitized, while 17.1% of egg masses carried at least one parasitized egg. The total number of parasitized egg masses was similar between parasitoid species, but the mean number of parasitized eggs per egg mass was highest for A. bifasciatus (5.57 vs. 1.25 for Ooencyrtus spp.).
4. Sentinel egg masses underestimated the parasitism compared to natural egg masses (respectively, 1.42% and 7.71%).
5. Our results suggested that the three generalist parasitoids detected can respond in a Leptoglossus egg density-dependent manner, but this requires further investigations.

     

Patch-exploitation patterns in an egg parasite of elm leaf beetle

Eggs of elm beetle,Pyrrhalta luteola (Muller), normally occur in masses which can be viewed as discrete host patches in space and time. Analysis of>800 egg masses from 3 field sites in northern California revealed that the number of eggs/mass varied from 2 to>40 and that the imported egg parasiteTetrastichus gallerucae (Fonsc.) exploited a relatively large proported of the egg masses at certain times. The spatial relationship between parasitization and number of eggs/egg mass was assessed in 2 ways-i.e., for exploited masses only and for all masses combined (exploited+nonexploited). Percent parasitization was density independent in most cases for both data sets. It is suggested that both methods of analysis can provide patterns which are relevant to biological control of insect pests.      

Do reproductive activities compromise immunological competence as measured by phenoloxidase activity? Field and experimental manipulation in females of two damselfly species

Recent studies of female insects indicate that reproductive activities, such as mating and oviposition, can impair immune ability. Using the two tropical damselfly species Argia anceps Garrison and Hetaerina americana (Fabricius), egg production and phenoloxidase (PO) activity, a key enzyme in insect immunity, are measured in mating, ovipositing and perching females in December and March. Perching females of both species have fewer eggs compared with mating and ovipositing females, which suggests that perching females are not engaged in reproduction. There is seasonal variation in egg number for the three categories in H. americana but not in A. anceps, which can be interpreted in terms of adaptive changes in egg production depending on female–male interactions in the former species but not in the latter species. There is no difference in PO activity among mating, ovipositing or perching females within either species, although measurements in December and March indicate distinct seasonal changes. Juvenile Hormone (JH) is known to reduce the effectiveness of the immune system by favouring the use of resources for reproduction. A possible role for JH is examined in H. americana, using the JH analogue methoprene to manipulate hormone activity, revealing that PO activity is reduced in methoprene‐treated H. americana females. Thus, although the results of the present study are indicative of possible hormone‐driven changes in PO, there is not necessarily a down‐regulation of immune function (as determined by PO activity) during mating or oviposition. The results complement some recent studies countering the idea that reproductive activities reduce the immune ability in insects.     

Clutch size in parasitoids: the egg production rate as a constraint

Summary In many species of insect parasitoids, adult females mature eggs as they search their environment for hosts. In such species, the number of mature eggs, at the point of finding a host, is a function of the interhost time and the rate of egg maturation. Assuming that interhost search times are variable, we use a version of the marginal value theorem to derive a decision rule for optimizing the time spent exploiting individual hosts; this indirectly determines clutch size. We find that a threshold search time exists above which a female should simply lay her currently mature eggs and depart from the host. However, when the search time has been less than the threshold, a female should oviposit, but then remain on the host to mature and lay additional eggs, until the threshold time is reached.