Occurrence of different development time patterns induced by photoperiod in Anagrus atomus (Hymenoptera: Mymaridae), an egg parasitoid of Empoasca vitis (Homoptera: Cicadellidae)

The wasp Anagrus atomus L. parasitizes eggs of the cicadellid Empoasca vitis (Göthe) on grapevines during the vegetative season. In vineyards, abundant parasitoid emergence occurs in the autumn as a result of a developmental delay in part of the A. atomus population. In this context, physiological times required by the parasitoid to develop in E. vitis eggs are recorded during the summer. In early summer, all individuals develop fully, with physiological times in agreement with those reported in the literature. From early August onwards, an increasing proportion of parasitoids show a delay in development involving pre‐imaginal stages. Under transmission microscopy, larvae of A. atomus inside the eggs of E. vitis can be clearly differentiated as ‘light’ and ‘dark’ types, which are associated with normal and delayed development, respectively. The decreasing photoperiod as autumn approaches appears to promote the retardation of growth. This phenomenon can be considered a typical case of risk spreading because, when the availability of E. vitis eggs in vineyards is high (i.e. in early summer), A. atomus shows (i) only a normal developmental pattern that allows a faster population increase, whereas (ii) when E. vitis egg availability begins to be scarce (i.e. from mid summer), an increasing proportion of individuals having delayed development provides better fitness because it allows the parasitoid to synchronize its life history with leafhopper species that lay overwintering eggs on plants other than grapevines.     

Superparasitism and mutual interference in the egg parasitoid Anagrus delicatus (Hymenoptera: Mymaridae)

Abstract.

• 1 In nature, interference among Anagrus delicatus (Hymenoptera: Mymaridae) parasitoids reduced the per-capita number of hosts parasitized. Interference increased with parasitoid density.
• 2 Anagrus delicatus did not avoid parasitizing hosts that had recently been parasitized by conspecific wasps. Evidence indicated that this superparasitism was largely a random process, increasing with the ratio of parasitized to unparasitized hosts.
• 3 Individual parasitoid efficiency, the number of hosts killed per wasp per unit time, decreased with increasing wasp density. This occurred whether wasps searched the patch together (simultaneously) or one by one (sequentially), and was the result of an increase in time spent superparasitizing hosts at higher wasp density. This is known as indirect mutual interference.
• 4 Increasing numbers of parasitoids together on the same patch caused a significant decline in the rate and per-capita number of hosts parasitized. However, there was not a correspondent decline in searching efficiency with increasing wasp density (i.e. no direct mutual interference).
• 5 These forms of parasitoid density dependence should contribute to the stability of the host—parasitoid interaction.

     

Influence of adult diet on fitness and reproductive traits of the egg parasitoid Anagrus virlai (Hymenoptera: Mymaridae), a potential biocontrol agent against the corn leafhopper

Dalbulus maidis is the most important leafhopper pest of maize in the Americas. Anagrus virlai is an egg parasitoid commonly associated with the corn leafhopper. We evaluated whether the performance of A. virlai is dependent on different diets provided during 24 hr or throughout adult female lifetime. Additionally, functional response of A. virlai on D. maidis eggs using maize leaves containing honeydew plus honey was described. A. virlai is a mostly pro-ovigenic autogenic species whose females are able to parasitize eggs immediately after emergence. We found that wasps oviposit the majority of their eggs in the first day of their adult life. Realized lifetime fecundity and lifetime potential fecundity did not differ significantly among treatments, but longevity and egg production increased when honey was added to diet. Data were consistent with an intermediate functional response between type II and III, but closer to type II, indicating a high parasitism rate at low host densities and a decrease in the oviposition rate at high host densities, due to a possible egg limitation. Our results suggest that carbohydrate food sources (honeydew and honey) might not be the factor limiting reproductive success during the first 24 hr. Food supply, however, might influence egg maturation and survivorship of wasps, thus potentially enhancing biological pest control when hosts are scarce in the course of the first few days of adulthood.     

Searching and Oviposition Behavior of Anagrus atomus L. (Hymenoptera: Mymaridae) on Four Host Plants of Its Host, the Green Leafhopper Empoasca decipiens Paoli (Homoptera: Cicadellidae)

Anagrus atomus L. is an important egg parasitoid of the green leafhopper Empoasca decipiens Paoli. In this study the ability of the parasitoid to locate and parasitize its host was investigated on four host plants, i.e., broad beans (Vicia faba L.), sweet pepper (Capsicum annuum L.), cucumber (Cucumis sativus L.), and French beans (Phaseolus vulgaris L.). For each plant species, the behavior of the parasitoid was observed on E. decipiens infested and noninfested plants. Searching and oviposition behavior were characterized by drumming, probing, and resting. Parasitoids spent significantly less time on non-infested than infested plants, 274.5 and 875.7 s, respectively, and no probing behavior was observed on non-infested plants. Frequency of resting behavior was significantly greater on non-infested than on infested plants. Total foraging time was significantly longer on infested than on non-infested plants, indicating that A. atomus females can efficiently discriminate between leaves with and without infestation. Parasitism of A. atomus was influenced by parasitoid density, with the highest parasitism rate (64.0%) obtained at a density of 10 A. atomus females/0.1356 m² but the number of parasitized eggs per female and the searching efficiency decreased with increasing parasitoid density.     

Life-table study of Anagrus atomus, an egg parasitoid of the green leafhopper Empoasca decipiens, at four different temperatures

The objective of our study was to assess thepotential of the egg parasitoid Anagrusatomus L. (Hymenoptera: Mymaridae) for controlof the greenhouse leafhopper Empoascadecipiens Paoli (Homoptera: Cicadellidae). Theegg-adult development time, survivorship andreproduction of A. atomus were evaluatedat four constant temperatures (16, 20, 24 and28°C). Developmental time ranged from33.6 days at 16°C to 13.3 days at 28°C. Based on a linear regression ofdevelopment rate on temperature the lowerthreshold was estimated at 8.39°C. Anagrus atomus required 263.2 degree-days tocomplete its development from egg to adult. Theegg-adult survival rate and the sex ratio weresignificantly lower at 28°C than at theother three temperatures tested. The intrinsicrate of increase (r _m) variedsignificantly between all four temperatures.The potential of A. atomus to attackdifferent host ages was additionallyinvestigated. Host eggs were parasitizedthroughout their development but rate ofparasitism was reduced in host eggs older thansix days. The number of eggs parasitized waspositively density dependent but the rate ofparasitism decreased with increasing hostdensity. A maximum rate of parasitism of 62.5%was recorded. The potential impact of the eggparasitoid on the population dynamics of E. decipiens is discussed.     

Reproductive and developmental biology of Gonatocerus ashmeadi (Hymenoptera: Mymaridae), an egg parasitoid of Homalodisca coagulata (Hemiptera: Cicadellidae)

The reproductive and developmental biology of Gonatocerus ashmeadi Girault, a parasitoid of the glassy-winged sharpshooter Homalodisca coagulata (Say), was determined at five constant temperatures in the laboratory: 15; 20; 25; 30; 33 °C. At 30 °C, G. ashmeadi maintained the highest successful parasitism rates with 46.1% of parasitoid larvae surviving to adulthood. Lifetime fecundity was greatest at 25 °C and fell sharply as temperature either increased or decreased around 25 °C. Temperature had no effect on sex ratio of parasitoid offspring. Mean adult longevity was inversely related to temperature with a maximum of 20 days at 15 °C to a minimum of eight days at 33 °C. Developmental rates increased nonlinearly with increasing temperatures. Developmental rate data were fitted with the modified Logan model for oviposition to adult development times across each of the five experimental temperatures to determine optimal and upper lethal temperature thresholds. The lower developmental threshold estimated by the Logan model and linear regression were 1.10 and 7.16 °C, respectively. Linear regression of developmental rate for temperatures 15–30 °C indicated that 222 degree-days were required above a minimum threshold of 7.16 °C to complete development. A temperature of 37.6 °C was determined to be the upper development threshold with optimal development occurring at 30.5 °C. Demographic parameters were calculated and pseudo-replicates for intrinsic rate of increase (r_m), net reproductive rates (R_o), generation time (T_c), population doubling time (T_d), and finite rate of increase (λ) were generated using the bootstrap method. Mean bootstrap estimates of demographic parameters were compared across temperatures using ANOVA and nonlinear regression.     

Influence of flowering cover crops on Anagrus parasitoids (Hymenoptera: Mymaridae) and Erythroneura leafhoppers (Homoptera: Cicadellidae) in New York vineyards

Abstract 1 We tested the hypothesis that providing nectar‐producing cover crops will enhance the biological control of grape leafhoppers (Erythroneura spp.) by Anagrus wasps in commercial vineyards in New York, U.S.A. 2 We established three cover crops between vine rows in a commercial vineyard: buckwheat (Fagopyrum esculentum (Moench)), clover (Trifolium repens L.) and mowed sod (Dactylis glomerata L.). 3 There was no effect of cover crop on adult Anagrus in 1996, whereas in 1997 adults were more abundant within edge vines with buckwheat compared to vines with clover or sod; adults were more abundant at the vineyard edge, especially early in the season. 4 Parasitism of ‘sentinel’ leafhopper eggs was higher on vines with buckwheat compared to parasitism on vines with clover or sod in 1996; a similar, non‐significant trend, was observed in 1997. 5 Neither the abundance nor the distribution of leafhoppers was influenced by cover crops, although in 1997 there was a trend toward greater numbers of nymphs on edge vines with buckwheat. 6 In a cage experiment, parasitism by Anagrus of leafhopper eggs on grapes was greater when adults had access to flowering buckwheat rather than buckwheat without flowers. 7 In a laboratory study, longevity of female Anagrus was increased when provided with honey or sugar water compared to water only or nothing. 8 Our results suggest that parasitism of grape leafhoppers by Anagrus may be enhanced by providing floral resources within vineyards in New York, although it is unclear whether this will produce meaningful reductions in pest abundance.     

The British species of Gonatocerus Nees (Hymenoptera: Mymaridae), egg parasitoids of Homoptera

Abstract. The thirteen British species of Gonatocerus Nees are characterized by means of a key and short diagnostic notes based on the females, with notes on their hosts and distribution. The species-groups proposed by Debauche are discussed; two species, Gonatocerus rogersi and G.minor, are described as new, and eighteen specific synonymies are proposed.     

Chromosomal polymorphism in an Australian leafhopper (Homoptera: Cicadellidae)

Variation in chromosome number from 2n=9 to 2n=7 was found in a species of leafhopper, Alodeltocephalus draba. It is argued that this variation is due to chromosome fusion and that A. draba is presently evolving to a lower number. The process of reduction is more advanced in some populations than in others.Seconded from: Department of Biology, University of Oregon, Eugene, Oregon, U.S.A.     

Some British species of Anagrus (Hymenoptera: Mymaridae)

Five species of Anagrus , egg parasites of various leafhopper species (Delphacidae and Cicadellidae, Honioptera), were bred for 3–13 generations in laboratory conditions. Biological and niorphological characteristics as well as attempts at crossbreeding were used to recognise the species Anagrus ensijer Debauche and four new. species. A. ensifer occurred in two varieties. The four new species and the two varieties of A. ensifer are described, and a key is given to both sexes.     

Evaluation of the Entomopathogenic Fungi Metarhizium anisopliae and Paecilomyces fumosoroseus (Deuteromycotina: Hyphomycetes) for Control of the Green Leafhopper Empoasca decipiens (Homoptera: Cicadellidae) and Potential Side Effects on the Egg Parasitoid Anagrus atomus (Hymenoptera: Mymaridae)

The efficacy of Metarhizium anisopliae strain Ma43 and Paecilomyces fumosoroseus strain Pfr12 (both Deuteromycotina: Hyphomycetes) against adults of Empoasca decipiens (Homoptera: Cicadellidae) and potential side effects on the egg parasitoid Anagrus atomus (Hymenoptera: Mymaridae), were investigated in greenhouse cage and laboratory experiments. Treating leafhopper-infested faba bean plants at a dose rate of 1×10⁷ conidia mL^-1 resulted in up to 97% mortality 7 days after application and a 100% infection rate. Experiments on the residual effects revealed a significant decrease in adult E. decipiens mortality with increasing time from application to insect release. The decrease in mortality over time corresponded well with data from conidia germination tests. The germination of conidia on agar medium after washing them from the surafce of sprayed plants declined significantly from 95 and 96% immediately after application for M. anisopliae Ma43 and P. fumosoroseus Pfr12, respectively, to 29 and 27% 5 days later. Experiments on potential side effects of the entomopathogenic fungi on A. atomus showed that the tested isolates had no influence on adult emergence and longevity; however, the rates of parasitism were significantly reduced by the fungal treatments. The latter might be due to either density effects and/or could indicate that A. atomus avoids fungal-treated plants. However, the parasitoid is substantially less susceptible to the fungal strains tested than the host itself.     

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.     

First host record for Anteon pilicorne (Ogloblin) (Hymenoptera: Dryinidae), a parasitoid of Cicadellidae, including the corn leafhopper (Hemiptera: Cicadellidae)

For the first time the dryinid wasp Anteon pilicorne (Ogloblin) is recorded as a parasitoid of two Macrostelini leafhoppers: Balclutha rosea (Scott) and the corn leafhopper Dalbulus maidis (DeLong & Wolcott). New distributional records are presented.     

榆树隐毡蚧各龄若虫和雄成虫形态及其生物学记述(同翅目: 蚧总科: 毡蚧科)

榆树隐毡蚧 Cryptococcus ulmi Tang et Hao是近年来在北京和山西中部发现的一种榆树新害虫。本文首次详细记述和图示了该虫幼期各虫态和雄成虫的形态特征 ;简要报道了其生物学特性 :一年发生 1代 ,以受精雌成虫在树干裂缝内越冬 ,卵胎生 ,每雌产仔 2 3～ 2 5头 ,平均值 35.2头。     

An integrative approach to species discrimination in the Anagrus atomus group sensu stricto (Hymenoptera: Mymaridae), with a description of a new species

Egg parasitoids of the genus Anagrus Haliday (Hymenoptera: Mymaridae) are natural enemies of many pests around the world. We used an integrative approach to characterize some species belonging to the Anagrus atomus group, using specimens reared from leafhoppers infesting some Lamiaceae. Starting from morphological identifications based on available keys, we carried out a multi-locus genetic characterization using phylogenetic and species delimitation analyses, and integrated it with biological and morphometric evidence. This approach revealed the existence of a new species, A. nepetellae sp. nov., here described along with its phenological traits. The approach also allowed us to characterize the expected taxonomic stability of other putative species within the group. We propose two new species groups called the atomus group sensu stricto and the vilis group, with the goal of streamlining the taxonomy of the atomus group. We also present a key for the identification of females of the European species of the atomus group sensu stricto. Some specimens show heterozygosity in 28S-D2 sequences, suggesting hybridization between A. atomus and A. nepetellae, which, if it is common between other species as well, could partly explain the taxonomic problems in the genus Anagrus. DNA sequencing of specimens reared by the same biofactory at different times suggests involuntary contamination leading to the displacement of a laboratory strain of A. atomus by A. nepetellae, which may have resulted from undetected partial reproductive compatibility.http://zoobank.org/urn:lsid:zoobank.org:pub:DFCD7654-5AA5-4E4F-AE48-24377BAACEFE     

Egg parasitoids (Hymenoptera: Mymaridae and Trichogrammatidae) of okra leafhopper,Amrasca biguttula (Ishida) (Hemiptera: Cicadellidae), in southern Japan

The okra leafhopper, Amrasca (Sundapteryx) biguttula (Ishida) (Hemiptera: Cicadellidae), is an economically important pest of okra, Abelmoschus esculentus (L.) Moench (Malvaceae), in Japan. During 2019 and 2022 surveys for its egg parasitoids (Hymenoptera) were conducted on Kyushu and Ryukyu islands in southern Japan. Five species of Mymaridae, Anagrus (Anagrus)?atomus (L.), A. (A.) japonicus Sahad, A. (A.) turpanicus Triapitsyn and Hu, Arescon enocki (Subba Rao and Kaur), and Stethynium empoascae Subba Rao, as well as one of Trichogrammatidae, Pseudoligosita nephotetticum (Mani), were recorded from eggs of A. biguttula on okra in Japan. Of these, A. (A.) turpanicus, identified using a combination of molecular and morphological methods, is newly recorded from Japan; A. (A.)?atomus, A. (A.) turpanicus and P. nephotetticum are for the first time reported from eggs of okra leafhopper.     

Biology and morphology of the immature stages of Hermeuptychia atalanta (Lepidoptera: Nymphalidae: Satyrinae)

This paper describes the morphology of the immature stages (egg, larva and pupa) and larval behavior of the butterfly Hermeuptychia atalanta, a common and widespread species of satyrinae in South America. Eggs are white and spherical, laid singly on leaves of grasses. The first instar has a light-green body and dark-brown head; all remaining instars present body and head capsule light green. Larvae have no spines, and general morphology is similar to other species of the subtribe Euptychiina.     

Oviposition behavior of Anagrus nigriventris,an egg parasitoid of beet leafhopper, Circulifer tenellus

Oviposition behavior of Anagrus nigriventris,a parasitoid of the eggs of the beetleafhopper,Circulifer tenellus, wasdescribed on sugar beet. Ovipositor probes ofrelatively long duration were accompanied by a periodof abdominal vibrations toward the end of the probe. These periods began as high frequency, low amplitudevibrations in the abdomen, and climaxed as a tremblingof the whole body in an up and down direction.Oviposition by the parasitoid was associated with theoccurrence of abdominal vibrations during a probe andwith probe durations 50 s; however thepresence/absence of abdominal vibrations during aprobe was considered a more reliable indicator ofoviposition than was a threshold probe duration of 50s. The majority of probes (63% of all probes) were inempty plant tissue, lasted 15 s, and were notassociated with abdominal vibrations. While probeswithout abdominal vibrations at host egg sites (14%of all probes) lasted an average of 48 s, probes withabdominal vibrations at host egg sites (23% of allprobes) lasted an average of 102 s, and all were 50s. It is hypothesized that the relative frequency ofthe 3 types of probes could be affected by the degreeof wasp experience, by the level of host cues on thesubstrate, and by the relative importance of ovipositor probing (vs. antennation) in the hostdetection process. Data from the study suggest thatA. nigriventris is able to discriminate againstpreviously attacked host eggs.     

Local augmentation efficiency of Anaphes nitens (Hymenoptera,Mymaridae), the egg parasitoid of Gonipterus platensis (Coleoptera,Curculionidae)

The efficiency of local augmentation releases of the egg parasitoid Anaphes nitens to control the Eucalyptus snout‐beetle Gonipterus platensis was tested in Eucalyptus globulus plantations in Galicia (NW Spain). On May–June 2006, at two localities of Pontevedra province, the release of host egg capsules parasitized by A. nitens at a potential rate of 300 adults/ha was compared with a release density of 900 adults/ha, and a control group of eucalypts not subjected to augmentation. Parasitism rate after 1–2 weeks did not significantly increase over the control plots at both localities. The high release rate did not ensure a higher crop protection and therefore could be not economically justified. On March–April 2017, at four localities of Pontevedra province, the test was replicated by releasing 300 parasitoids/ha. Parasitism level did significantly increase over the control just in one locality. Augmentation of A. nitens at small scale generally failed to achieve a higher protection from the pest, possibly due to the large extension of the E. globulus plantations, the magnitude of the G. platensis population and the fluctuations of the established parasitoid population, whose density is in turn affected by host egg availability and density‐dependent dispersal.