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Epyris extraneus Bridwell (Bethylidae), a fossorial wasp that preys on the larva of the tenebrionid beetle, Gonocephalum seriatum (Boisduval)

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... Flat wasps are larval parasitoids of holometabolan insect immatures (mostly coleopteran and lepidopteran larvae) and their adults are mostly smaller than their future offspring's hosts (Gauld and Bolton 1988), which the females paralyse with their venom sting (Powell 1938;Finlayson 1950;Schaefer 1962;Evans 1964;Kühne and Becker 1974;Gordh and Medved 1986;Griffiths and Godfray 1988;Abraham et al. 1990;Howard et al. 1998). Since the host immatures often occur in more cryptic or concealed habitats, like soil, stems, wood or seeds (Evans 1964;Gauld and Bolton 1988;Howard and Flinn 1990), flat wasp adults may show additional adaptations for entering these habitats (Williams 1919;Gordh and Medved 1986), such as fossorial (digging, burrowing) forelegs and reduced wings (Evans 1964). Some flat wasps even exhibit subsocial behaviours (Evans 1964), additional (to parasitoidism) maternal care (Casale 1991;Hu et al. 2012;Yang et al. 2012;Tang et al. 2014) and many engage in prey carriage and some also in a sort of nest building (Finlayson 1950;Evans 1964;Rubink and Evans 1979;Howard et al. 1998; for review of prey carriage in wasps in general see e.g. ...
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Parasitism, a malignant form of symbiosis, wherein one partner, the parasite, derives benefits to the detriment of another, the host, is a widespread phenomenon. Parasitism sensu lato is understood here to include many phenomena, like parasitoidism, kleptoparasitism, phoresy and obligate parasitism. Insecta has many in-groups that have evolved a parasitic life-style; one of the largest in-groups of these is probably the group of Hymenoptera. Bethylidae, the group of flat wasps, is a smaller in-group of Aculeata, the group of hymenopterans with venom stings; representatives of Bethylidae are parasitic. They are more specifically larval ectoparasitoids, meaning that their immature stages are externally developing parasites that kill their host organism at pupation (end of interaction). They mostly parasitise immature representatives of Coleoptera and Lepidoptera. Female flat wasps search for a host for their progeny, paralyse it with their venom sting and then oviposit onto it. Herein we describe one of the oldest findings of parasitic interactions of parasitoid wasps with their progenies’ hosts, specifically a flat wasp female grasping and (potentially) stinging a beetle immature in Cretaceous Kachin (Myanmar) amber (ca. 100 million years old). This finding indicates that this type of parasitic interaction existed since the Cretaceous, temporally close to the earliest findings of representatives of Bethylidae.
... The females are predators. They prepare simple nests on the ground and drag small larvae into it (Williams, 1919;Rubink & Evans, 1979). This behaviour is very unusual within the bethylids. ...
... Apparently Chrysidinae display antennae with three sensilla, often located on a distinct papilla (Tormos et al. 1998. 4) Mandibles of mature dryinid larvae are generally falciform or securiform without additional teeth, whereas tridentate mandibles, as in Ampulicomorpha, have often been described in other Chrysidoidea families, where different types of pluridentate mandibles have been observed (Williams 1919;Asís et al. 1994;Tormos et al. 1996Tormos et al. , 1997Tormos et al. , 1998Tormos et al. , 1999Tormos et al. , 2009) but simple mandibles are apparently quite rare. 5) The complete absence of the galea, as in Dryinidae + Embolemidae, has also been observed in Bethylidae and in Caenochrysis doriae (Gribodo, 1874) (Chrysididae: Chrysidinae) . ...
Article
The mature larva of Ampulicomorpha schajovskoyi De Santis & Vidal Sarmiento, 1977, is described and figured for the first time. Larval characters of Dryinidae and Embolemidae are discussed in regard to possible synapomorphies of each family and of both families together (Dryinidae + Embolemidae) as monophyletic groups. Some larval characters are compared with the corresponding conditions in other Chrysidoidea families.
... Estes generos nao ficaram agrupados, tendo se ligados em um nivel de similaridade de 0,60, que foi baixo quando comparado com os dados da analise de agrupamento da armadilha Janela (Fig. 5). Os Epyrinae, incluindo algumas especies de Holepyris, sao parasit6ides de larvas de Coleoptera de graos armazenados, mic6fagos e de casca de arvores (EVANS 1964(EVANS , 1969(EVANS , 1977WILLIANS 1919). Estes generos ficaram juntos na analise de agrupamento da armadilha Malaise (Fig. 4) bem como na analise de agrupamento da armadilha ...
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Bethylidae specimens from the Reserve were studied in its ecological and faunistic aspects. The material was collected by Malaise and Window traps simultaneously in ten different areas of the Reserve during four years. The total number of genera and specimens were analyzed. Indices of diversity and evenness were used for characterizing the community ecology. Clustering analysis of localities and genera were provided. Nine genera of Bethylidae were found in the Reserve, being Pseudisobrachium Kieffer, 1904 and Apenesia Westwood, 1874 the most common ones. Window trap was more efficient than Malaise trap in terms of genus diversity.
... The larvae of solitary bethylid species do not appear to differ morphologically P. A. Luft, unpublished manuscript). However, it is possible that the fitness accrued from large size could de-from gregarious forms and are nonmotile (e.g., Williams 1919; Maneval 1930; Bognár 1957; Abraham et al. 1990). pend on host size across species. ...
Article
Parent-offspring conflict over clutch size may lead to siblicidal behavior between juveniles. In parasitoid wasps, selection for siblicide in small broods is predicted to produce a dearth of gregarious broods with few eggs. Here we document the clutch size distribution in the Bethylidae, a large family of aculeate parasitoids. Small gregarious clutches are the most common. Further data suggest that the most common gregarious clutches in the parasitoid Hymenoptera as a whole contain only a few eggs. Across bethylid species, both clutch size and wasp size increase with host size. Within genera clutch size is more closely related to host size, but between genera or larger clades wasp size is more closely related to host size. The volume of the emerging wasp brood does not depend on whether a species lays single- or multiple-egg clutches once host size is taken into account. These data suggest that clutch size in bethylid wasps is best described by traditional optimality models and that siblicide plays little role in this and possibly other families. We propose several ecological reasons for the rarity of siblicide in bethylids: ectoparasitism, idiobiosis, and a suite of characteristics associated with high within-brood relatedness.
Article
Soil-dwelling tenebrionid larvae have developed in three evolutionary lines and are pests in all faunal regions. Description of and keys to most of the important species are available. However, revisional studies, on both adult and larval characters, are needed in many groups. The basic number of non-sex chromosomes appears to be 18, but the number and type of sex chromosomes are variable. Host ranges of both larvae and adults are usually extensive but usually only include plants. Development rates are related mainly to temperature, with life-cycles of one, two or three years. Larval development periods are rather long, and adults are long-lived. The number of instars is high, often above ten and frequently variable within a species. Sampling is best carried out by direct counts and pitfall trapping. In the species studied, mating follows a regular pattern and pheromones are involved. Species show definite temperature, moisture and soil-type preferences and these influence both spatial and temporal distribution. Recorded parasitoids include Diptera (mainly tachinids), Hymenoptera, mites, bacteria and fungi. Birds, carabids and a variety of other vertebrates are the main predators, but abdominal secretions and postural mechanisms provide some defence. Cultural control was formerly widely practised but has given way to chemical control.
Article
Parasitoid wasps of the family Bethylidae lay groups of eggs on the larvae of beetles and moths. The larvae develop together and pupate in the vicinity of the host. On hatching, the majority of copulations are between siblings leading to a high degree of local mate competition. Sex ratio theory predicts that as the number of individuals developing on a host increases, the population sex ratio will become progressively more female biassed. A comparative study of sex ratios across the family supports this prediction.
1. The wasp Megascolia flavifrons stings larvae of the stag beetle Oryctes nasicornis on the ventral side of all segments, except the last three, which do not contain nerve ganglia. 2. Experiments indicate a central rather than a peripheral action of the venom. 3. From pharmacological analysis it is concluded that the venom does not contain cholinergic or serotonergic activity, but contains histamine- and bradykinin-like substances. 4. The presence of histamine was confirmed by a radioenzymatic method.
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