The eggshell of the almond wasp Eurytoma amygdali (Hymenoptera, Eurytomidae) - 1. Morphogenesis and fine structure of the eggshell layers

The almond wasp Eurytoma amvgdali (Hymenoptera: Eurytomidac) feeds and oviposits exclusively in almonds and therefore is characterized as an insect of economic importance. Its meroistic polytrophic ovaries include follicles with a tri-partitc configuration. The mature follicles exhibit two filaments occupying the two poles of the egg. One is the micropylar filament while the other might serve for respiration since it is likely that its flattened end layers remain outside the almond fruit. The eggshell is formed by aposition and the follicle cells, which surround the follicle until the end of oogenesis, may be responsible for protein synthesis and secretion which finally lead to the assembly of the eggshell. The eggshell comprises the thin vitelline membrane, possibly a 'wax' layer of waterproofing function, a transluscent layer which appears amorphous even at the end of choriogenesis, a granular layer, including large and small electron-dense granules, and finally a columnar layer very similar to layers found in other insect species of the same or different orders. Peroxidase is histochemicalLY found for the first time in an eggshell of the Hymenoptera order: the tranluscent layer in particular is positively stained (electron-dense). Two possible roles of this peroxidatic activity are discussed, first, in comparison to other fruit-infesting insects, we assume that elastic chorion is produced through the function of peroxidase induced bonds (resilin-type bonds), very important for avoiding premature breaking, while being oviposited through a narrow ovipositor. Second, referring to other studies, this layer can play a bactericidal role for additional embryonic itprotection.     

Studies of the behaviour, dispersal, and damage potential of the almond wasp, Eurytoma amygdali

Under laboratory conditions, females of Eurytoma amygdali were attracted to almond fruit, whereas males were not. Females changed their position twice as often as males, in the absence of fruit. Almond leaves had little, if any, attraction for females; moisture had none. Outdoors, free wasps were active in and around almond trees in sunny and in overcast weather, during the morning mainly on the eastern side, and in the afternoon mainly on the western side of trees. Males were often dancing horizontally in a plane parellel to the periphery of trees. Courting and mating were observed on fruits only. 527 wasps which emerged on three focus trees equally distributed in an otherwise uninfested orchard damaged an average of about 25.8 fruits per female. 70.9% of the damaged fruits were on the focus trees, 17.4% on trees directly facing them, and 12.8% on further removed trees. In some cases, trees bordering on open spaces were more heavily infested than those between them and the focus trees. No correlation was found between the number of infested fruits and fruit load of individual trees.

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Zusammenfassung In Käfigen im Laboratorium wurden Weibchen, Männchen jedoch nicht von Mandelfrüchten angezogen. Mandelblätter und Feuchtigkeit übten kaum bezw. keine Anziehungskraft auf Weibchen aus. In Abwesenheit von Früchten änderten Weibchen ihre Position ungefähr doppelt so oft wie Männchen. Frei lebende Wespen waren in sonnigem und bewölktem Wetter aktiv, morgens an der östlichen, nachmittags an der westlichen Seite der Bäume. Männchen tanzten oft nahe der Peripherie der Baumkronen. Vor dem Beginn der Aktivität am Morgen saßen Männchen und Weibchen auf der Oberseite der Blätter. Werben und Kopulation wurden auf Früchten beobachtet. In einer vorher nicht befallenen Mandelanlage wurden in den Kronen von drei gleichmäßig verteilten Bäumen insgesamt 527 Wespen zum Schlüpfen gebracht. Diese beschädigten im Durchschnitt pro Weibchen 25.8 Früchte. 70.9% der befallenen Früchte waren auf die 3 Focusbäume, 17.4% auf die Bäume, welche die Focusbäume direkt umgaben, die restlichen 12.8% auf die übrigen Bäume der Anlage verteilt. In einigen Fällen waren Bäume, die an offenes Feld grenzten, schwerer befallen als die Bäume zwischen ihnen und den Focusbäumen. Eine Korrelation zwischen Fruchtmenge und Befall pro Baum konnte nicht festgestellt werden.

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Contribution from The Volcani Center, Agricultural Research Organization, Bet Dagan, Israel. 1972 Series, No. 2219-E.     

Effect of temperature on the termination of diapause in the univoltine almond seed wasp Eurytoma amygdali

Diapausing larvae of Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) were collected in early August and late September. They were subjected to various photoperiod and temperature regimens for up to 20 weeks, then kept at L16:D8 and 19 °C for another 14 to 26 weeks for diapause to be terminated and pupation to take place. Photoperiod did not affect diapause completion. It was confirmed that the two morphologically distinct diapause stages have different temperature requirements for their completion. The first diapause stage was completed synchronously at temperatures between 16 and 19 °C. A higher temperature of 26 °C delayed diapause development. The second stage required lower temperatures between 4 and 10 °C. Spontaneous termination of diapause was observed at constant 19 °C. When applied to the first diapause stage for 20 weeks, low temperatures made the larvae refractory to subsequent intermediate temperatures. The first stage was thus maintained until a higher temperature of 26 °C made the larvae regain their ability to respond to the intermediate temperatures and complete this stage. Larvae grown in Retsou almonds had a higher diapause intensity than larvae grown in Truoito almonds. The results suggest that, in nature, the high temperatures of late summer and early autumn are likely to maintain the first diapause stage. Subsequently, the less warm temperatures of autumn allow the completion of the first stage by late autumn, and the low temperatures of late autumn and of winter allow the completion of the second diapause stage by mid winter.     

Host discrimination and evidence for a host marking pheromone in the almond seed wasp, Eurytoma amygdali

Laboratory experiments and observations on the oviposition behaviour of the almond seed wasp Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) revealed that the females of this species deposit a host-marking pheromone, immediately after an oviposition, by dragging the tip of their abdomen on the fruit surface. This pheromone enables them to discriminate between the infested and uninfested fruit and to select for oviposition the latter. Its primary function is apparently the prevention of repeated ovipositions in already infested fruit, thus contributing to the optimal utilisation of the available resources for larval development. The responses of individual females to different treatments of almonds, in a series of two-choice tests, revealed that the pheromone can be perceived by the females on direct contact and, when at high concentrations, also olfactorily from a short distance. The pheromone was present inside the abdomen and thorax of females but not of males, and, although water soluble, could not be entirely removed from heavily infested almonds when rinsed with water. Direct observations revealed that after an average of 3.7 successive visits to pheromone-bearing almonds, females were induced to walk or, most often, fly away from the experimental set-up. This suggests that the pheromone may also contribute to the dispersion of the wasps.

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Résumé Des expériences de laboratoire et des observations concernant le comportement de ponte de Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) ont montré que, immédiatement après la ponte, les femelles déposent une phéromone de marquage de l'hôte en trainant le bout de leur abdomen sur la surface de l'amande. Cette phéromone les rends capable de distinguer les fruits infectés des non-infectés dt de sélectionner pour la ponte les derniers. Apparemment, la fonction principale de cette phéromone est la prévention de la répétition des ovipositions dans les fruits déjà infectés et la répartition uniforme des oeufs dans les amandes, contribuant ainsi à la meilleure utilisation des ressources disponsibles pour le développement des larves. Des expériences de deux choix entre des fruits de différents traitements ont montré que la phéromone pouvait être perçue par les femelles par le direct contact et, quand elle était à hautes concentrations, par olfaction d'une courte distance. La phéromone était présente dans l'abdomen et dans le thorax des femelles, et bien qu'elle soit soluble à l'eau, elle ne pouvait pas s'éloigner entièrement par lavage des amandes sérieusement infectées à l'eau. Les observations ont démontré qu'après un numéro de 3.7 visites successives sur des amandes portant de la phéromone les femelles s'éloignaient du lieu de ponte en marchant ou, le plus souvant, en s'envolant, ce qui suggère que la phéromone contribue à la dispersion des femelles.

     

Dispersal of the thistle gallfly Urophora cardui and its endoparasitoid Eurytoma serraiulae (Hymenoptera: Eurytomidae)

Abstract. 1. The dispersal of the gall-forming tephritid Urophora cardui L. (Diptera: Tephritidae) and its endoparasitoid Eurytoma serratulae Latr. (Hymenoptera: Eurytomidae) from an experimental colony were investigated over a period of 4 years.
2. The dispersal rate for both fly and endoparasitoid was one to two magnitudes higher than in previous studies, which found 100 m per generation maximally. The dispersal rate of the endoparasitoid was not less than that of its host. Most newly founded colonies were small and had high extinction rates.
3. The viability of colonies of U. cardui did not correlate with the distance from the dispersal centre.
4. The dispersal rates observed are high enough to allow a multiple spread and extinction in mid-Europe in the post-pleistocene period.     

Monitoring populations of the almond seed wasp, Eurytoma amygdali, with sex pheromone traps and other means, and optimal timing of chemical control

During the four years 1986–1989 we monitored an adult male population of the almond seed wasp, Eurytoma amygdali Enderlein, in the region of Thessaloniki, using sex pheromone traps, each containing 25 living virgin females. The emergence of adults from infested almonds of the previous year was also monitored in cages in the field, as well as the progress of fruit infestation and egg hatching in the almonds of the Retsou variety. Finally, two chemical control experiments were done against the newly hatched larvae to determine the optimal time for chemical control.During the 4-year study the start of adult captures occurred between April 13 and April 24, and the maximum between April 17 and May 11. In most cases, adult emergence from infested almonds in field cages began with a delay of 2 to 14 days in relation to the start of trap captures, and was affected by factors such as the date of cage installation. In all years most eggs hatched within 12–18 days of oviposition.A treatment with phosphamidon at a concentration of 0.06%, applied to tree branches of the Retsou variety with a hand-sprayer on May 23, 1987, when 54.1% of the eggs had hatched, was very effective (final infestation in July was 4.7% as compared to 89.1% of the untreated control). Three treatments with phosphamidon 0.06%, applied each on different tree branches, on May 14, 20, and 25, 1988, when the percentage of egg hatching was 7.8%, 65.3%, and 93.5%, respectively, were all very effective (final infestation in July was 1.6%, 3.6%, and 4.2% respectively, as compared to 93.6% of the untreated control). However, the presence of mines in the perisperm and outer endosperm, caused by the feeding larvae before they were killed, was more conspicuous in the almonds treated when the percentage of egg hatching was high.Our data suggest that the almond seed wasp can be effectively controlled with a single treatment of a systemic insecticide against the newly hatched larvae, preferably applied when the percentage of egg hatching is low to medium (between 10% and 50%). In the area of Thessaloniki, during 1986–89, 10% and 50% of egg hatching occurred about 21 and 27 days respectively after the beginning of male adult captures in pheromone traps.     

Biology and behavior of the seed borer wasp Bephratelloides cubensis ashmead (Hymenoptera: Eurytomidae)

The soursop Annona muricata is an important fruit for national market, and for exportation, but the crop is affected by pests and diseases. The seed borer wasp Bephratelloides cubensis Ashmead is the pest that produces the highest damage to the crop in Mexico. Sixty percent of damaged fruits and 5-50 seeds per fruit have been registered, with 25% reduction in yield. In Nayarit, Mexico, 100% of damaged fruits were recorded. In this State, an experiment with soursop was conducted to study the life cycle under field conditions and to determine diurnal behavior of the female of B. cubensis. The highest activity of the wasp was observed between 12:00h and 13:00h (35oC, 54% RH and 409.34 luxes). Females oviposited in fruits with a diameter of 3.1-7.6 cm. Larvae of B. cubensis developed five instars, adults survived no longer than 22 days, and female survived longer than males; they lived 22 and 15 days, respectively. Life cycle of B. cubensis varied from 69 to 122 days.     

New data on trophic associations and distribution of the eurytomid genera Eurytoma Illiger and Systole Walker (Hymenoptera, Chalcidoidea: Eurytomidae)

New data on the distribution and trophic associations are given for three eurytomid species: Eurytoma infracta Mayr, Eu. salvicola Zerova, and Systole (Trichosystole) cuspidata Zerova. Eurytoma infracta and S. (T.) cuspidata were reared in Portugal, the first species from Cynipid galls on Salvia sclareoides Brot. (Lamiaceae), the second, from seeds of the same plant. Eurytoma salvicola Zerova was found for the first time in France, having being reared there from galls of Oxyacina tibialis R.-D. (Diptera, Tephritidae) on Rosmarinus officinalis L.     

Biological species problems in some Tetramesa (Hymenoptera, Eurytomidae)

Using horizontal starch gel electrophoresis, ten enzyme systems were surveyed in ten populations of Tetramesa reared from eight species of grass collected from different sites in Britain. Eight species of Tetramesa were recognized on the basis of the number and relative mobilities of electromorph bands for larvae, pupae and adults. No differences were found between populations of T. hyalipennis (Walk.) reared from Elymus repens (L.) and E.farctus (Viv.) or between populations of T. eximia (Gir.) reared from Calamagrostis epigejos (L.) Roth and Ammophila arenaria (L.) Link. Experiments on mate-choice and host preference also supported the electrophoretic results for populations of T. hyalipennis and T. eximia.     

A review of the Palaearctic species of the genus Eurytoma, belonging to the E. robusta species-group (Hymenoptera, Eurytomidae), with description of two new species

A review of Palaearctic species of the Eurytoma robusta group is given. This group includes 46 species. A key to these species is provided. Two new species, Eurytoma ibaraca Zerova, sp. n. from Japan and E. clarissae Zerova, sp. n. from Israel, are described. E. ibaraca is close to E. armenica Zer., but differs from it in the longer metasoma of the female, longer funicular segments, longer postmarginal vein, and sharp prominence on the fore coxa. E. clarissae is similar to E. asiatica Zer., but differs from it in the bulging fore coxa and longer postmarginal vein. The type material is deposited in the collection of the Shmalhausen Zoological Institute, National Ukrainian Academy of Sciences (Kiev).     

Species of the genus Nikanoria related to Nikanoria mongolica (Hymenoptera, Eurytomidae)

A complex of the species related to Nikanoria mongolica Zerova is discussed. This complex includes a new subspecies (N. mongolica punctulata subsp. n.) and two new species (N. kazakhstanica sp. n. and N. sugonjaevi sp. n.). Data on the distribution and trophic associations of these species are provided. The type material is deposited in the collection of the Schmalhauzen Institute of Zoology, National Ukrainian Academy of Sciences, Kiev.     

The structure of the female reproductive system in the Eurytomidae (Chalcidoidea: Hymenoptera)

An account is given of the structure of the gaster and mechanism of the ovipositor in the females of the Eurytomidae. The histology and the possible functions of the components of the reproductive systems are described and its bearing on their phylogenetic relationships of the group discussed.  

Summary:

The female gastral sclerites and ovipositor mechanism in the Eurytomidae have a number of distinctive characters. Within the family there is a progressive evolution of a coiling of the ovipositor associated with accommodating a longer shaft within the abdomen. Each ovary has three ovarioles and the shape of the egg is constant. There are two pairs of colleteral glands, a well developed "venom" system and a distinctive spermatheca.
The family has affinities with the Torymidae and may be close to the ancestral chalcid stock.     

Relationships among the tiphiid wasp subfamilies (Hymenoptera)

Abstract. The phylogenetic relationships among the subfamilies of Tiphiidae are discussed, with detailed analysis of characters, and presentation of a phylogenetic tree. The tribe Diammini Turner is elevated to subfamily status. A key to the subfamilies is included, and a brief synopsis of each subfamily is given.     

Revision and phylogeny of the European species of the Eurytoma morio species group (Hymenoptera: Eurytomidae), parasitoids of bark and wood boring beetles

A revision of Eurytoma (Hymenoptera, Eurytomidae) species belonging to the morio group is proposed. Species discrimination is based on morphological and, partly, on molecular data, including the barcoding fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene and the D2 expansion region of the 28S ribosomal gene. Morphological and molecular phylogenetic analyses largely support the morphological evidence. E urytoma cristata Delvare sp. nov. , E urytoma saliphila Delvare sp. nov. , and E urytoma sylviae Delvare sp. nov. are described from France, E urytoma ithma Delvare sp. nov. is described from France and Italy, and E urytoma gatesi Delvare sp. nov. is described from North America and France. Decatomidea polygraphi Ashmead, 1894 and Ipideurytoma spessivtsevi Bou?ek & Novicky, 1954 are synonymized with Eurytoma afra Boheman, 1836. Eurytoma auricoma Mayr, 1878 is removed from synonymy with Eurytoma arctica Thomson, 1875 and is synonymized with Eurytoma maura Boheman, 1836. Eurytoma eccoptogastri Ratzeburg, 1844, Eurytoma flavoscapularis Ratzeburg, 1844, Eurytoma flavovaria Ratzeburg, 1844, and Eurytoma masii var. flavonigra Russo, 1938 are synonymized with Eurytoma morio Boheman, 1836. Eurytoma masii Russo, 1925 and Eurytoma kemalpasensis Narendran, Tezcan & Civelek, 1995 are synonymized with Eurytoma striolata Ratzeburg, 1848. Eurytoma melanoneura Walker, 1871 and E. masii are removed from synonymy with E. morio. Lectotypes are designated for E. afra, E. auricoma, E. masii, Decatoma aloisifilippoi Russo, 1938, and E. masii var. flavonigra. © 2014 The Linnean Society of London     

Structure and ultrastructure of the spermatozoa of Bephratelloides pomorum (Fabricius) (Hymenoptera: Eurytomidae)

The spermatozoa of Bephratelloides pomorum are very long and fine. Each spermatozoon measures about 620 μm in length by 0.38 μm in diameter and, when seen under the light microscope, appears to be wavy along its entire length. The head, which is approximately 105 μm, comprises a small acrosome and a nucleus. The acrosome is made up of a cone-shaped acrosomal vesicle surrounding the perforatorium and the anterior end of the nucleus. Innumerable filaments radiate from it. The perforatorium has a diameter equal to that of the nucleus at their junction, where it fits with a concave base onto the rounded nuclear tip. The nucleus is helicoidal and completely filled with homogeneous compact chromatin. It is attached to the tail by a very long and quite electron-dense centriolar adjunct that extends anteriorly from the centriole in a spiral around the nucleus for approximately 8.5 μm. The tail consists of an axoneme with the 9+9+2 microtubule arrangement pitched in a long helix, as well as a pair of spiraling mitochondrial derivatives (with regularly arranged cristae) that coil around the axoneme, and two small accessory bodies. As well as the spiraling of the nucleus, mitochondrial derivatives and axonemal microtubules, the sperm of B. pomorum present other very different morphological features. These features include the acrosome and centriolar adjunct, both of which differentiate the spermatozoa from the majority of sperm found in other Hymenoptera. In addition these structural variations demonstrate that the sperm of chalcidoids provide characteristics that can certainly prove useful for future phylogenetic analysis at the subfamily level and, possibly, the genus too.     

Emergence patterns of adult citrus gall wasp,Bruchophagus fellis (Hymenoptera: Eurytomidae), and its key parasitoids in southern Australia

Emergence of adult citrus gall wasp (CGW) Bruchophagus fellis (Girault) (Hymenoptera: Eurytomidae) in citrus orchards in far western New South Wales, Australia, was monitored with sticky traps for three seasons (2010 to 2012 inclusive). Emergence started from early October to early November, peaked from late October to mid-November, and was mostly finished by mid–late November. Emergence timing was mainly influenced by year and site, with the largest differences observed between years. Most emergence (90%) took place during an average period of only 19 days each season (range of 11–28 days across all trap types, sites and seasons). The role of temperature in emergence timing was investigated using a degree-day (DD) model. DD accumulated since 1 April using a lower threshold temperature of 15°C and an upper threshold temperature of 35°C or 40°C gave the best predictions of median emergence dates in the three years. The required DDs to achieve 5, 50, and 95% emergence were 336, 403 and 447 DD, respectively. The maximum difference between predicted and observed median dates for 2010–2012 was only four days. CGW adult emergence in future years can be predicted using these DD parameters and a combination of observed and average historical temperature data for the target site. Effective prediction of emergence peaks will allow the timing of pesticide applications to be optimised whilst providing protection for establishing parasitoids which emerge from the galls 2–3 weeks after citrus gall wasp.     

Phylogeny of the parasitic wasp subfamily Charipinae (Hymenoptera, Cynipoidea, Figitidae)

The Charipinae are a major group of hyperparasitoids of Hemiptera. Here, we present the first cladistic analysis of this subfamily's internal relationships, based on 96 morphological characters of adults. The data matrix was analysed using uniformly weighted parsimony. The effects of using alternative weighting schemes were explored by performing additional searches employing implied weights criteria. One of the caveats of implied weights analysis is that it lacks an objective criterion for selecting the value of the concavity function. In the present study, differential weighting was used to explore the sensitivity of our results to the alternative assumptions made in the analysis and to select one of the most parsimonious trees under equal weights, which we regard as being the hypothesis that minimizes the amount of ad hoc assumptions. The validity of the two existing tribes and the monophyly of all the genera of Charipinae were tested, in particular the cosmopolitan and highly species-rich Alloxysta and Phaenoglyphis , which appear repeatedly in ecological and biochemical studies of host–parasitoid associations. The evolution of several major characters and the relationships between genera are discussed. On the basis of the phylogenetic results, we discuss a number of taxonomic issues. A new classification of the subfamily is proposed in which no tribes are maintained, Carvercharips is synonymyzed with Alloxysta , and the creation of a new genus from Nepal is justified. Our analysis points to the need for a world revision of the basal genus Phaenoglyphis , which is shown as paraphyletic.