Pre-introductory risk assessment studies of Gonatocerus ashmeadi (Hymenoptera: Mymaridae) for use as a classical biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae) in the Society Islands of French Polynesia

Homalodisca vitripennis (Germar) (=H. coagulata [Say]) (Hemiptera: Cicadellidae) invaded French Polynesia in 1999. A classical biological control program against H. vitripennis was initiated in 2004 aiming to introduce the exotic egg parasitoid Gonatocerus ashmeadi (Girault) (Hymenoptera: Mymaridae) to the Society Islands archipelago. Prior to any release, two risks were assessed: (a) continued uncontrolled H. vitripennis spread and proliferation in French Polynesia, and (b) non-target impacts by G. ashmeadi on indigenous French Polynesian cicadellids. The primary risk of H. vitripennis is its potential to vector the lethal plant bacterium, Xylella fastidiosa. While the presence of X. fastidiosa in French Polynesia has not yet been demonstrated, the presence of uncontrolled H. vitripennis greatly elevates the risk of a disease outbreak and thus represents a major threat for numerous plant species. Assessing the risk of G. ashmeadi introduction for native cicadellids first required an inventory of the Cicadellidae of the Society Islands, resulting in at least 14 cicadellid species (nine not previously recorded). The risk to these species of attack by G. ashmeadi was assessed using four criteria: (1) their phylogenetic relationships to known hosts of G. ashmeadi, and their similarity in (2) body size, (3) egg laying biology, and (4) ecology. All indigenous cicadellid species found were considered to be at low risk of attack because they differed greatly from all known hosts for G. ashmeadi: (1) none of the indigenous species are in the tribe Proconiini, (2) all were very small and, when possible to determine, (3) lay tiny single eggs, which (4) are deposited on the undersides of leaves of trees. These results persuaded the French Polynesian Government that the benefits of establishing G. ashmeadi for H. vitripennis control outweighed the serious potential risks associated with either delaying release or not releasing G. ashmeadi in French Polynesia. Releases of G. ashmeadi in Tahiti began in May 2005.     

Determination of Homalodisca coagulata (Hemiptera: Cicadellidae) egg ages suitable for oviposition by Gonatocerus ashmeadi, Gonatocerus triguttatus, and Gonatocerus fasciatus (Hymenoptera: Mymaridae)

Homalodisca coagulata (Say) (Hemiptera: Cicadellidae) eggs 1–10 days of age were exposed to Gonatocerus ashmeadi Girault, Gonatocerus triguttatus Girault, and Gonatocerus fasciatus Girault (all Hymenoptera: Mymaridae) in no choice laboratory trials to investigate egg age utilization and to determine which egg ages are vulnerable to attack by these three parasitoids. The H. coagulata egg ages that were most suitable for oviposition by G. ashmeadi, G. triguttatus, and G. fasciatus were eggs 3, 4, and 2 days of age, respectively. Egg ages least suitable for parasitoid development were 6–10 days for G. ashmeadi (resulting in <50% parasitism), 1–2 and 7–10 days for G. triguttatus (resulting in <25% parasitism), and 3–10 days for G. fasciatus (resulting in <11% parasitism). Pooling parasitism data across all egg ages showed that parasitism by G. ashmeadi was 12.9 and 28.5% higher compared with G. triguttatus and G. fasciatus, respectively, and G. triguttatus resulted in 15.6% higher percentage parasitism compared with G. fasciatus. Egg age had a significant effect on the percentage of female G. ashmeadi offspring produced, but this was not significant for G. triguttatus, and low G. fasciatus parasitism prevented statistical analyses for comparisons. Results from tests where females were offered a choice for oviposition between eggs 1, 3, and 5 days of age demonstrated that G. ashmeadi and G. triguttatus showed no significant oviposition preference, while percentage parasitism by G. fasciatus was 29.4 and 7.4% higher when females were presented eggs 1 and 3 days of age, respectively, compared with eggs 5 days of age. Choice tests indicated that an overlap in egg age suitability for oviposition exists between G. ashmeadi, G. triguttatus, and G. fasciatus, and that interspecific competition for eggs 1, 2, and 3 days of age may occur in the field environment.     

Powdering of egg nests with brochosomes and related sexual dimorphism in leafhoppers (Hemiptera: Cicadellidae)

Leafhoppers from 16 New World genera currently placed in the tribes Proconiini (13) and Cicadellini (2) of the subfamily Cicadellinae, in the subfamily Phereurhininae (1), and one undescribed genus exhibit a suite of morphological specializations associated with a unique type of maternal care. The females produce a specific type of brochosomes (secretory particles manufactured in the Malpighian tubules of most leafhoppers and usually forming a non‐sex‐specific hydrophobic coating of the integument), which they store prior to oviposition on their forewings in the form of pellets and then apply as a powdery coating to the egg clusters placed under the epidermis of leaves or, in one genus, exposed. The function of the brochosome coating is unknown, but may include protection of eggs against parasitoids and pathogens, and facilitating gas exchange. Observations on several species from the genera Phera, Homalodisca, Oncometopia, Quichira, Cuerna, and the undescribed genus are reported, complementing scanty existing data on this behaviour. Its taxonomic occurrence is described for the first time based on study of collections, where ready‐to‐oviposit females are occasionally preserved with pellets of brochosomes on their forewings. A comprehensive survey of associated female‐specific structural modifications is presented based on examination of c. 80 predominantly Neotropical species. These modifications include (1) production of modified brochosomes (2) modified setation of the forewing area upon which pellets of such brochosomes are placed, and (3) elongation of the metathoracic tibial macrosetae, which transfer the brochosomes onto the egg nest. The traits are absent or rudimentary in males. The specialized brochosomes display remarkable interspecific diversity and can be used for taxonomic identification of eggs. Egg‐powdering apparently has evolved as a modification of the anointing and grooming behaviours involved in making coatings of integumental brochosomes in most leafhoppers. Morphological similarity, extending to characters not related to oviposition, suggests that all genera containing ‘powdering’ species may form a single lineage, but this hypothesis yet needs to be tested by phylogenetic analyses. Several such genera include species displaying powdering‐associated traits and species lacking these, suggesting multiple independent losses or origins of powdering. This peculiar kind of maternal care provides an interesting model for study of the evolution of novel functional complexes of traits. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140 , 353–381.     

Engineering an invasion: classical biological control of the glassy-winged sharpshooter, <Emphasis Type="Italic">Homalodisca vitripennis</Emphasis>, by the egg parasitoid <Emphasis Type="Italic">Gonatocerus ashmeadi</Emphasis> in Tahiti and Moorea,French Polynesia

The glassy-winged sharpshooter, Homalodisca vitripennis Germar (=H. coagulata Say) (Hemiptera: Cicadellidae), invaded Tahiti in 1999 and spread rapidly to the main island groups of French Polynesia becoming an important pest. It threatened agriculture, native biodiversity, and created serious social and recreational problems. Further, massive uncontrolled populations on Tahiti presented an elevated invasion threat to other South Pacific nations. In 2004, a classical biological control program against H. vitripennis was initiated in French Polynesia using the highly host-specific egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae). After risk assessment studies indicated an acceptably low level of risk to non-target species, 13,786 parasitoids were released at 27 sites in Tahiti between May and October 2005. Here we present the results of G. ashmeadi and H. vitripennis population surveys during the first year of their interaction in French Polynesia (until mid-May 2006). The impact of G. ashmeadi on H. vitripennis was extremely rapid and high. Parasitism of H. vitripennis egg masses by G. ashmeadi has averaged 80–100% in Tahiti since the introduction of the parasitoid, and populations of H. vitripennis nymphs and adults have decreased by more than 90% since December 2005. Populations of H. vitripennis have been successfully maintained at this low level for more than 1 year. The same results were obtained in nearby Moorea where the parasitoid was probably spread by the unregulated transport of plants infested with parasitized H. vitripennis eggs. Population monitoring continues in order to determine if a stable equilibrium between the pest and the parasitoid has been reached.     

Comparative assessments of Gonatocerus ashmeadi and the ‘new association’ parasitoid G. deleoni (Hymenoptera: Mymaridae) as biological control agents of Homalodisca vitripennis (Hemiptera: Cicadellidae)

Egg age preference, competitive ability, and behavior of Gonatocerus deleoni (‘new association’ parasitoid) and G. ashmeadi (the co-evolved dominant parasitoid in California) were investigated in the laboratory to determine whether one species exhibited competitive superiority. When searching concurrently for Homalodisca vitripennis egg masses, G. ashmeadi consistently outperformed G. deleoni by parasitizing 59–89% more eggs under three different experimental systems in the laboratory with varying host densities, egg ages, and exposure times. G. ashmeadi parasitism in control vials containing one parasitoid ranged from 58 to 86% and was up to 28% higher at egg ages 1 and 3 days compared with 5 days. G. deleoni in control vials parasitized on average 7% of H. vitripennis egg masses in 1 h regardless of egg age. G. deleoni failed to parasitize H. vitripennis egg masses in 15 min when caged alone or in competition with G. ashmeadi. In a combined species treatment, parasitism by G. ashmeadi was 11% higher when H. vitripennis eggs were exposed for 5 days compared with 24 h. Conversely parasitism by G. deleoni was 5% lower for this comparison. G. ashmeadi had a significantly female biased sex ratio for all three experimental designs, whereas, G. deleoni offspring sex ratio was not significantly greater than 50%. In comparison to G. ashmeadi, behavioral observations indicated that G. deleoni was absent from host egg infested leaves 53% more frequently and it oviposited 66% less frequently. No incidences of females aggressively chasing competitors off H. vitripennis egg masses were recorded during this study.     

Comparative assessments of Gonatocerus ashmeadi and the ‘new association’ parasitoid Gonatocerus tuberculifemur (Hymenoptera: Mymaridae) as biological control agents of Homalodisca vitripennis (Hemiptera: Cicadellidae)

Egg age preference, competitive ability, and behavior of Gonatocerus tuberculifemur (‘new association’ parasitoid) and Gonatocerus ashmeadi (‘old association’ parasitoid) were investigated in the laboratory to determine if one species exhibited competitive superiority. When searching concurrently for Homalodisca vitripennis egg masses, G. ashmeadi consistently outperformed G. tuberculifemur by parasitizing 25–53% more eggs under three different experimental systems in the laboratory with varying host densities, egg ages, and exposure times. G. ashmeadi parasitism in control vials containing one parasitoid ranged from 81–97% across all egg ages. G. tuberculifemur in control vials parasitized 60–66% of eggs 1 and 3 days old, and just 18% of eggs 5 days old. G. ashmeadi produced 5–16% more female offspring than G. tuberculifemur for all experimental conditions. In comparison to G. ashmeadi, G. tuberculifemur was observed off leaves with host eggs 20% more frequently and it oviposited 15% less frequently. G. ashmeadi and G. tuberculifemur when confined together allocated ∼1% of behaviors to antennating or aggressively chasing competitors off egg masses, and up to 2% of behaviors to antennating host egg masses and/or ovipositing into eggs from the opposite side of the leaf. These latter behaviors did not occur when parasitoids were confined alone with host eggs.     

Maximum realised lifetime parasitism and occurrence of time limitation in Gonatocerus ashmeadi (Hymenoptera: Mymaridae) foraging in citrus orchards

The relationship between body size (hind tibia length), <12 h egg load, wing wear and parasitoid age was used to estimate realised lifetime parasitism of recently dead Gonatocerus ashmeadi collected in a citrus orchard. Under prevailing field conditions and methodology assumptions, it was estimated that female G. ashmeadi lived on average for 183 ± 17 degree-days, parasitised a total of 87 ± 9 Homalodisca vitripennis eggs, and died with 34 ± 5 eggs remaining in the ovaries. Only 17% of dead G. ashmeadi died with no mature eggs suggesting that 83% of G. ashmeadi were not egg limited at time of death. Estimates of realised lifetime parasitism for female G. ashmeadi under prevailing field conditions in July and August in a southern California citrus orchard indicated that time of year had a significant effect on reproductive output. Additionally, live G. ashmeadi captured daily during June through August 2006 had body size, egg load and wing wear recorded to detect possible monthly changes in parasitoid age and egg load. Foraging G. ashmeadi captured alive in June were older and oviposited more eggs in the field compared with August. Only 0.5% of live G. ashmeadi were captured with no mature eggs in their ovaries indicating that the vast majority of live G. ashmeadi were not egg limited.     

Sex ratio dynamics of Gonatocerus ashmeadi,a parasitoid of the glassy‐winged sharpshooter in California

We examined whether Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), a quasi‐gregarious egg parasitoid of Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), produces precise sex ratios under a field setting. Under laboratory conditions, previous studies have shown that G. ashmeadi exhibits strongly female‐biased sex ratios with low variance in the number of males produced per host. Field‐collected G. ashmeadi tend to produce much less female‐biased sex ratios with high variance in male numbers. We found significant positive effects of proportion parasitism and host density on sex ratio. Proportion parasitism also had a positive effect on sex ratio variance. The findings of this study are discussed in the context of theoretical predictions.     

Short-distance dispersal behavior and establishment of the parasitoid Gonatocerus ashmeadi (Hymenoptera: Mymaridae) in Tahiti: Implications for its use as a biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae)

The egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), was introduced into French Polynesia as a biological control agent to control the invasive plant feeding pest Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). The short-distance dispersal of G. ashmeadi was monitored as part of the biological control program. G. ashmeadi showed exponential dispersal capacity with 47 m/day being a minimum estimate of its natural rate of spread at high host densities (>150 nymphs per minute of sweep net sampling) in urbanized areas at sea level, which were characterized by a high diversity of exotic ornamental plants. This rate of spread contrasted starkly with almost nonexistent establishment and dispersal where host densities were very low (<2 nymphs per minute of sweep net sampling) at high elevation (800 m) with relatively undisturbed native vegetation. Survey results across different altitudes revealed an effect of vegetative diversity and host density on the measurable mobility and establishment of G. ashmeadi. In contrast, no significant influence of wind direction was found on G. ashmeadi dispersal rate or direction. Survey results for G. ashmeadi from French Polynesia suggest that the best release establishment strategies for classical biological control of H. vitripennis are: (1) many small releases where host density is high, or (2) larger and fewer releases where host densities are low.     

Use of life table statistics and degree-day values to predict the invasion success of Gonatocerus ashmeadi (Hymenoptera: Mymaridae), an egg parasitoid of Homalodisca coagulata (Hemiptera: Cicadellidae), in California

Life table statistics and degree-day requirements for Gonatocerus ashmeadi Girault, a parasitoid of the glassy-winged sharpshooter Homalodisca coagulata (Say), were used to estimate the number of expected parasitoid generations in California (USA). Between two to 51 and one to 37 generations per year were estimated across different climatic regions in California, using life table and degree-day models, respectively. Temperature-based values for net reproductive rate, R_o, were estimated in California using a laboratory-derived equation and ranged from zero to approximately 48 and analyses indicate that a minimum of eight generations are required each year to sustain a population increase of G. ashmeadi. Long-term weather data from 381 weather stations across California were used with an Inverse-Distance Weighting algorithm to map temperature-based estimations for the entire state of California. This Geographic Information Systems model was used to determine number of G. ashmeadi generations based on day-degree accumulation, T_c, and R_o. GIS mapping indicated that Californian counties in the north, central west coast, central west and Sierra Nevada regions may be climatic conditions unfavorable for supporting the permanent establishment of invading populations of G. ashmeadi should H. coagulata successfully establish year-round populations in these areas. Southern counties in California that experience warmer year round temperatures and support year round populations of H. coagulata, appear to be conducive to the establishment of permanent populations of G. ashmeadi. The mechanisms facilitating G. ashmeadi invasion and the implications of these temperature-based estimates for biological control of H. coagulata are discussed.     

Documentation of brochosomes within Hemiptera, with emphasis on Heteroptera (Insecta)

Brochosomes, small secretory particles often found in abundance on the integument of leafhoppers, are currently considered a unique feature of the family Cicadellidae. The present work documents, for the first time, brochosomes in two other groups of Hemiptera, i.e. Psylloidea and Heteroptera. In Heteroptera the occurrence of relatively large amounts of brochosomes on various body parts is documented in several species of different families. Morphological structures of the legs and the setation are illustrated in detail and compared to those of Cicadellidae. The results are based on scanning electron microscopy (SEM) examination of museum specimens. The possible function of brochosomes within Hemiptera is briefly discussed.     

Species identity of geographically distinct populations of the glassy-winged sharpshooter parasitoid Gonatocerus ashmeadi: morphology, DNA sequences, and reproductive compatibility

Gonatocerus ashmeadi is a common and seemingly widespread egg parasitoid of Homalodisca coagulata, the glassy-winged sharpshooter (GWSS). Location records for G. ashmeadi indicate its natural range to be the southeastern USA and northeastern Mexico (which coincides with the presumed native range of GWSS), and possibly southern and central California (CA) (the adventive range of GWSS). The purpose of our work was to determine whether G. ashmeadi in the USA and northeastern Mexico is one species or a complex of reproductively incompatible sibling species. We used three approaches to determine the species identity of different G. ashmeadi populations: (1) reassessment of key morphological features using scanning electron microscopy (SEM) to determine if subtle morphological differences exist between G. ashmeadi populations which could indicate species differences; (2) to determine if molecular differences exist between G. ashmeadi populations collected from different regions by comparing mitochondrial and ribosomal DNA sequences; (3) mating compatibility studies to determine if different populations of G. ashmeadi are reproductively isolated, or if mating occurs, whether offspring are viable thereby defining species groups on the basis of successful interbreeding. Results from these three areas (morphology, DNA sequences, and reproductive compatibility) have been evaluated collectively; leading us to the conclusion that G. ashmeadi as it is currently viewed is a valid species and not an aggregate of morphologically indistinguishable cryptic species.     

Secretion of brochosomes during the ontogenesis of a leafhopper, Oncometopia orbona (F.) (Insecta, Homoptera, Cicadellidae)

Secretion of brochosomes, intricately structured symmetrical protein-lipid particles produced in the specialized Malpighian tubules of the Cicadellidae, was studied during the ontogenesis of Oncometopia orbona (F.). Unlike most other cicadellids, O. orbona displays sexual dimorphism in the production of brochosomes. The Malpighian tubule secretory cells of nymphs, males and young females produce spherical brochosomes of 0.3-1.4 microm in diameter, extruded after molts through the hindgut and spread across the integument as a coating. In females with developing eggs the same cells switch to producing rod-like brochosomes of 3.5-11.0 microm in length, which are used to powder the oviposition sites. The transition involves a dramatic change in morphogenesis of the secretory particles. Both types of brochosomes derive from homogenous condensing Golgi granules and acquire their definitive structure as a result of growth, differentiation of the contents into a wall and a core, and development of a lattice of surface invaginations. Final stages of this process take place in Golgi-derived vacuoles. In the development of rod-like brochosomes the condensing granules coalesce into masses several times larger than nascent spherical type. These giant granules flatten, then become doughnut-shaped and finally break open into a pair of linear particles. Whether the chemical composition differs between the two types of brochosomes remains unknown.     

Egg maturation,oosorption, and wing wear in Gonatocerus ashmeadi (Hymenoptera: Mymaridae), an egg parasitoid of the glassy-winged sharpshooter,Homalodisca vitripennis (Hemiptera: Cicadellidae)

Egg maturation and oosorption in Gonatocerus ashmeadi were investigated in the laboratory and the relationship between hind tibia length (HTL) and <12 h egg load, and wing wear and parasitoid age were determined. G. ashmeadi given access to honey-water and hosts, on average, matured 77 eggs in excess of those they were born with. The number of mature eggs in female G. ashmeadi provided honey-water with no hosts significantly declined after 163 degree-days eggs, while the number of ‘dissolved’ eggs (partially disintegrated mature eggs) increased by nine eggs after 163 degree-days. These results are consistent with oosorption. There was a significant positive correlation between HTL and <12 h egg load. The ovigeny index (the number of mature eggs at female emergence divided by potential lifetime fecundity) for G. ashmeadi was calculated as 0.22 indicating that this parasitoid is a syn-ovigenic species when studied under laboratory conditions. There was a significant positive correlation between wing wear (measured as the number of broken setae per wing) and parasitoid age in the laboratory. The practical implications of these results for G. ashmeadi on the biological control of Homalodisca vitripennis are discussed.     

Parasitization by Edovum puttleri (Hymenoptera: Eulophidae) in relation to host density in the field

Abstract.

• 1 The relationship between parasitization by Edovum puttleri Grissell and density of eggs of the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae), was studied on two spatial scales (eggs mass and 6 m² cage).
• 2 For both scales, rates of parasitism were generally inversely related to host density for periods ranging from 2 to 8 days after parasitoid release. Thereafter, parasitism became independent of host density.
• 3 The initial inverse-density relationship and subsequent shift to density independence may result from several factors: (1) ambient temperatures, (2) the parasitoid's limited egg production, (3) differential times of exposure of egg masses to parasitoids, and/or (4) the parasitoid's patterns of host feeding and oviposition.
• 4 Although overall levels of parasitism were relatively low, total mortality of L.decemlineata eggs (including nonviable and cannibalized eggs, and those killed by parasitoid feeding) in parasitized egg masses was consistently high (?70–90%).

     

Analysis of the Anointing and Grooming Behavior of Several Adult Insects in Typhlocybinae (Hemiptera: Cicadellidae)

Anointing and grooming behaviors of the adult of Singapora shinshana (Matsumura) (Erythroneurini), Empoascanara (Empoascanara) mai Dworakowska (Erythroneurini), Austroasca vittata (Lethierry) (Empoascini), and Limassolla diospyri Chou et Ma (Zyginellini) were recorded and analysed. Through observation and analysis of their anointing and grooming, we found that there is a preparation stage before anointing behavior in typhlocybines. Anointing and grooming are repeated several times and the numbers and interval times differ from species to species among the Typhlocybinae. Ultramorphological examination of brochosomes and the morphology of the brochosome field, metathoracic tibiae and the structures on the head, face, mesothorax and pygofer of E. mai were done using a scanning electron microscope. Brochosomes (IBS) on the cuticle of typhlocybine adults were spherical visible above these integument. Characteristics of anointing and grooming behavior, the provide evidence that the brochosome field may function as a storage site in preparation for the next grooming session.     

Biology and host specificity of <Emphasis Type="Italic">Gonatocerus deleoni</Emphasis> (Hymenoptera: Mymaridae), a potential biocontrol agent of <Emphasis Type="Italic">Homalodisca vitripennis</Emphasis> (Hemiptera: Cicadellidae) in California,USA

The host-specificity and biological traits of Gonatocerus deleoni Triapitsyn, Logarzo & Virla (Hymenoptera: Mymaridae), a potential candidate for biological control of the glassy-winged sharpshooter (GWSS), Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae), were determined under laboratory conditions. G. deleoni is a solitary egg parasitoid native to Argentina, originally reared from sentinel eggs of Tapajosa rubromarginata (Signoret) (Cicadellidae). With GWSS as a fictitious host, G. deleoni’s average development time from oviposition to adult emergence was 18.8 ± 1.4 days, with males developing faster than females (18.0 ± 1.3 days, males; 19.0 ± 1.3 days, females). The average parasitism rate on 1–8-day-old eggs was 45.7% but this was significantly affected by the age of the egg, ranging from 1.4% to 69.9% (egg ages 8 and 3, respectively). The average sex ratio was 0.34 (percent males) and sex ratio was not significantly affected by egg age. G. deleoni was able to develop in eggs of GWSS and Homalodisca liturata Ball (both in the tribe Proconiini), but was unable to develop on eggs of Graphocephala atropunctata (Signoret) (different tribe, same subfamily) or Erythroneura elegantula Osborn (different subfamily).     

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.     

Cold storage effects on maternal and progeny quality of Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae)

This study determined the effects of cold storage on the survival, development and reproduction of the mymarid wasp, Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae). Following storage of the immature parasitoids within host eggs of the glassy-winged sharpshooter, Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae) under a daily fluctuating temperature for up to 50 d, the quality of the parental and F₁ generations was assessed by examining several reproductive and developmental parameters indicating fitness. Immature wasps were stored for 20 d within the host without reducing their subsequent survival, development or progeny fitness parameters. After 30 d of storage, survival declined, post-storage developmental time was extended, and the fecundity of the adult females decreased. Storage for 40 d severely damaged G. ashmeadi, because it not only yielded a 12% survival rate, 44% female sterility and increased the proportion of progeny males by 155%, but it also reduced parasitism and fecundity by 70% and 73%, respectively. No wasps emerged after 50 d of storage. Cold storage affected the emergence pattern of the parental but not the F₁ and F₂ generations. Parental emergence was extended and the pattern displayed two additional peaks after the initial onset. Analysis of several demographic parameters for the parental and F₁ generations further confirmed that the quality of the adult parents declined after they had been stored as immatures for 30 d. The detrimental effects caused by cold storage of the parental generation do not extend to the F₁ generation. Our results indicate that short-term cold storage of G. ashmeadi within its host could be used for maintaining and accumulating these parasitoids during mass propagation for release in a control program.     

Egg mass destroying and guarding behaviour of the giant water bug,Lethocerus deyrollei Vuillefroy (Heteroptera: Belostomatidae)

Egg mass destroying behaviour of femaleLethocerus deyrollei and male defending behaviour were observed in a densely vegetated concrete pond. In 3 of 9 cases of male-female interactions, the males successfully guarded their egg masses. In 4 of the 6 remaining cases in which the egg masses were destroyed by the female, females successfully took over the mate and laid their own egg masses the same night. These results suggest that male brooding guards the egg mass from females, as well as supplying eggs with water.