Life cycles and host–parasitoid relationships of five species of Leucospis wasps in Argentina (Hymenoptera: Leucospidae)

The genus Leucospis Fabricius is comprised of parasitoid wasps relatively uncommon in nature. Their immature stages develop on aculeate Hymenoptera, in particular solitary bees, but hosts are known for only around 25 Leucospis species (about 20% of 115–120 world species), so the true relationship of this family with bees is largely unknown. Here we report on individuals of five species of Leucospis wasps which emerged from nests of different bee and wasp species during distinct sampling periods during a trap-nesting programme, in two contrasting areas: agro-ecosystems and natural habitats in the Pampean region. Some of these nests were parasitised by L. cayennensis Westwood, L. coxalis Kirby, L. egaia Walker, L. pulchriceps Cameron and L. signifera Bou?ek. Our results expand the available information of host species for L. cayennensis, L. coxalis, L. egaia and L. pulchriceps and represent the first record of hosts associated with L. signifera. The hosts were only bee species of the genera Centris, Tetrapedia (family Apidae) and Megachile (Megachilidae). These species were more abundant in the natural reserve than in agro-ecosystems, suggesting that anthropogenic activities could negatively affect their populations. Most nests were attacked in one (43.7%) or two (40.7%) cells, but the remaining nests (16.5%) had more (up to seven cells). However, the position of the attacked cells was variable, suggesting that females of Leucospis species oviposit in recently built cells, and that the hatching of larvae is delayed, or that the first larval stage waits until the host larva reaches a sufficient size to attack.     

The parasitoid complex and parasitoid‐induced mortality of spiders (Araneae) in a Central European woodland

The parasitoid complex of spiders was studied in three different sites of a forest and, for comparison, in open habitats in north‐western Lower Saxony (Germany). Qualitative data and, for four spider species, detailed quantitative rearing data were obtained. At least 25 parasitoids of spiders were recorded (23 species of Hymenoptera and two species of Diptera). External parasitoids of spiders were rare in the woodlands; only up to 1% of the spiders collected by hand searching and sweep netting had ectoparasitoids. During the investigation 23 parasitoid species were reared from spider egg masses, 10 of them occurring in woodlands. Two species were secondary parasitoids. Several parasitoid–host relationships were recorded for the first time. Egg masses of Floronia bucculenta were parasitized up to 5% by two species of the genus Gelis. Up to 10% of the egg masses of Linyphia hortensis were parasitized by Aclastus species. Egg masses of the spider genus Ero were parasitized by three species, causing an overall mortality of 40%. For egg masses of Agroeca spp. parasitization at some locations was up to 60%. An overview of the Central European parasitoid complex of spiders is provided.     

An overview of the mite fauna (Acari) associated with eumenine wasps (Hymenoptera: Vespidae) found in Brazilian collections

ABSTRACT

Mites can be associated with other organisms as parasites, commensals, phoretic, among others, such as some Winterschmidtiidae that present cooperative relationships with solitary wasps (Eumeninae). These wasps have one or more cavities in their body surface that are capable of carrying mites, called acarinaria. Studies carried out in the northern hemisphere suggest that these relationships are species-specific. However, in South America, there are few studies in this field. Aiming to recognize and increase our knowledge of the species of solitary wasps from Brazil that are associated with mites, we examined 61 wasps belonging to 29 species and four genera from Brazilian museum collections. All of the specimens studied presented at least one type of acarinarium in their bodies. There were mites in all specimens of wasps, but not all of them were associated with an acarinarium. The mites belong to 11 different genera: six in Winterschmidtiidae (possibly cooperation relationships); one in Oplitidae (phoresy); one in Erythraeidae (parasitism); two in Acaridae (phoresy); and one in Histiostomatidae (phoresy). Detrended correspondence analysis and indicator species analysis were conducted to test the preference of the mite genera for species of wasp and for site (regions of the wasp’s body). These tests were significant only for the mite genus Vespacarus preferring Parancistrocerus wasp species and the metasomal acarinaria. Some of the mites did not have a specific host, and some wasps carried more than one species of mite, differing from the specific interactions reported for the northern hemisphere.     

Two new species of Hymenoepimecis (Hymenoptera: Ichneumonidae: Pimplinae) with notes on their spider hosts and behaviour manipulation

The wasp genera included in the Polysphincta genus-group develop exclusively as koinobiont ectoparasitoids of spiders, an unusual pattern within Hymenoptera. All known species of Hymenoepimecis attack orb-web spiders and Hymenoepimecis argyraphaga was previously described inducing a sudden modification of its host's web-building behaviour. Information on parasitoid–host interactions and even on host identities for most members of the genus-group, however, remain scarce. In this study we describe two new species of Hymenoepimecis and present information on their hosts, spiders of the genera Leucauge (Tetragnathidae) and Manogea (Araneidae). Cocoon webs of both host species are distinct from their normal webs. The modified cocoon web of Leucauge roseosignata constructed under the influence of Hymenoepimecis japi sp. nov. is very similar to the cocoon web described for Leucauge argyra. It is composed of three axes, including several radial threads each, and a hub used by the larva to attach the cocoon's suspension line. Spiral viscous threads, present in normal webs and in webs of parasitized spiders carrying larvae in the initial instars, are absent in cocoon webs. In the cocoon web of Manogea porracea the horizontal sheet is poorly structured and the cocoon is attached far from the spider's normal resting position, in an area composed of a densely tangled structure.     

The occurrence and phylogenetic implications of the ovipositor clip within the Figitidae (Insecta: Hymenoptera: Cynipoidea)

The presence of the ovipositor clip is surveyed throughout Figitidae. This morphological structure is postulated to restrain the parasitoid host during oviposition. All Figitinae and Eucoilinae that attack semi‐concealed dipterous hosts were found to possess the clip. Figitids that attack fully concealed hosts all lacked the ovipositor clip. It is hypothesized here that two subfamilies, the Anacharitinae and Aspicerinae, attack fully exposed hosts yet lack the clip in order to quickly oviposit and prevent fighting with the host. Mapping of the presence/absence of the ovipositor clip on to two competing phylogenies suggests this structure evolved in parallel in Figitinae and Eucoilinae. The evolution of the ovipositor clip is probably responsible for the successful colonization of semi‐concealed dipterous larvae by Eucoilinae.     

The parasitoid complex attacking coexisting Spanish populations of Euphydryas aurinia and Euphydryas desfontainii (Lepidoptera: Nymphalidae,Melitaeini)

We report on a five‐year investigation of the complex of parasitoids associated with coexisting populations of Euphydryas aurinia and Euphydryas desfontainii at a Spanish site. A sample of over 7000 eggs, 1000 larvae and 200 pupae (the last of these in part having been experimentally placed) revealed the existence of no egg parasitoids, of three primary larval parasitoids and of five primary pupal parasitoids, plus 13 secondary and facultatively tertiary parasitoids associated with the cocoons of the main larval parasitoid, the specialist Cotesia sp. D. The most abundant secondary parasitoid, Neochrysocharis albiscapus, entirely escaped tertiary parasitism by being gregarious. The mortality induced by parasitoids on the butterfly populations, although heavy, was clearly partially minimized by secondary parasitoids acting on Cotesia sp. D. The striking coincidence of the parasitoid complexes attacking the two Euphydryas species suggests a good system for investigating apparent competition between hosts sharing common natural enemies.     

Host–parasite relationships and life cycles of cuckoo wasps in agro-ecosystems in Argentina (Hymenoptera: Chrysididae: Chrysidini)

Chrysididae is a diverse group of parasitoid/cleptoparasitic wasps; however, host–parasite relationships and life cycles of few species have been studied. Nests of different wasp and bee species were obtained during a trap-nesting programme, in the Pampean region. Some of these nests were parasitised by cuckoo wasps females of Caenochrysis taschenbergi (Mocsáry), Chrysis boutheryi (Brèthes), C. saltana Bohart, C. sp. 1 (ignita-group), C. sp. 2 (ignita-group), Neochrysis lecointei (Ducke), Pleurochrysis ancilla (Buysson) and P. lynchi (Bréthes). This paper reports new data about host–parasite relationships and life cycles for these species. Multiple parasites (from one species or from different families of insects) emerged from single cells of some parasitised nests: from each host cell parasitised by species of Pleurochrysis, two adults emerged successfully, information previously unknown for the genus; and in three cases of cells parasitised by C. boutheryi, two adults successfully emerged from a single cell; in two cases both individuals were chrysidine, and in third one was chrysidine and one Leucospis pulchripes (Leucospidae).     

Altitudinal patterns of abundances and parasitism in frugivorous drosophilids in west Java,Indonesia

Abundances and parasitism rates of frugivorous drosophilid flies were studied in three localities at altitudes of 250–1350 m in and near Bogor, Indonesia. The drosophilid and parasitoid species were classified into four groups: low-altitude species, high-altitude species, species abundant at a mid-altitude location and species occurring rather evenly from low to high locations. The ananassae and immigrans species groups were major drosophilids collected. All species of the ananassae species group were more abundant at lower altitudes, and the parasitism rate in this species group decreased with increasing altitude. Thus, the host abundance seems to affect the parasitism rate. On the other hand, the rate of parasitism in the immigrans species group showed no apparent relation with altitude or density, possibly due to the fact that species of this species group varied in altitudinal distribution. It is also suggested that the diversity of drosophilid species affects the composition of parasitoid species.     

Brood parasitism and nest survival of Brown-headed Cowbird hosts at high-elevation riparian sites in the eastern Sierra Nevada, California

Deciduous riparian ecosystems in the western United States provide habitat for a higher density of breeding birds than reported for any other avian habitat type and provide habitat for more breeding bird species than adjacent uplands. On the east slope of the Sierra Nevada, riparian ecosystems make up Molothrus ater )—an obligate brood parasite that forages on bare ground and feedlots but typically commutes to distinct shrubland or woodland habitats for breeding. We examined nest survival, brood parasitism, breeding phenology, and causes of nest failure for birds at North Lake and Rock Creek: 2 high-elevation (>2500 m) riparian breeding habitats adjacent to recreational development and within cowbird commuting distance to additional potential foraging sites. Nest survival tended to be higher for host species at Rock Creek than for those at North Lake, but parasitism rates were not significantly different between plots. Of 21 open-cup nesting species, 12 were parasitized. We found the highest rate of parasitism (92%) for Warbling Vireos ( Vireo gilvus ) at North Lake, and parasitism contributed to lower total nest survival there (14%). For nearly all species, parasitized nests were less successful and produced fewer young than nonparasitized nests. However, predation was the leading cause of complete nest failure across all species and contributed to the lowest total nest survival estimates for Western Wood-Pewees ( Contopus sordidulus, 11%) and Dusky Flycatchers ( Empidonax oberholseri, 15%) at North Lake and for Dark-eyed Juncos ( Junco hyemalis, 15%) at Rock Creek. Nest survival was relatively high for Western Wood-Pewees (41%) at Rock Creek and for Yellow Warblers ( Dendroica petechia, 47%) at North Lake. We noted whether the arrival of pack animals at pack-station corrals contributed to variation in cowbird numbers at corrals or in parasitism rates at the 2 sites. Cowbirds occupied corrals before and after pack-stock arrival, and most host clutches were completed prior to pack-stock arrival at nearby corrals, suggesting that the presence of pack animals did not directly affect cowbird host species.     

Satellite flies (Leucophora personata,Diptera: Anthomyiidae) and other dipteran parasites of the communal bee Andrena agilissima (Hymenoptera: Andrenidae) on the island of Elba,Italy

Solitary and presocial aculueate Hymenoptera are parasitized by a range of dipteran species in the families Anthomyiidae, Bombyliidae, Conopidae, Phoridae, and Sarcophagidae that are likely to impact on their hosts. We undertook a study over several years of a univoltine and communal bee, Andrena agilissima, and its main dipteran parasites, in particular the satellite fly Leucophora personata (Diptera: Anthomyiidae). Behavioural and ecological data were collected from one nesting aggregation of the host bee on the island of Elba, Italy, from 1993 to 2003, and from a foraging site of the bee, ca 5?km from the nesting aggregation. Other Diptera associated with A. agilissima at the field site were the bee fly Bombylius fimbriatus (Bombyliidae), the conopid fly Zodion cinereum (Conopidae), and the scuttle fly Megaselia andrenae (Phoridae). The phenology of the Diptera broadly overlapped with that of their host across the season of activity (end of April and all of May). Diurnal activity patterns differed slightly; L. personata in particular was active at the host's nesting site before A. agilissima. Female satellite flies also showed a range of behaviours in gaining entry to a host nest. We summarize published data on this and other Leucophora species that parasitize Andrena host bees. Host bees returning to their nests occasionally undertook zig‐zag flight manoeuvres if followed by a satellite fly that were generally successful in evading the fly. Satellite flies that entered a nest, presumably to oviposit, were less likely to remain therein if another host bee entered the same nest, suggesting that one advantage to communal nesting for this host is a reduction in brood cell parasitism by L. personata. We provide the first clear evidence for parasitism by a Zodion of any Andrena host. Both L. personata and M. andrenae concentrated their parasitic activities in the zone of the host nesting aggregation with highest nest densities. Three of the Diptera, L. personata, B. fimbriatus, and Z. cinereum, seemed to have extremely low rates of parasitism whilst that of M. andrenae appeared low. Though they have refined parasitic behaviour that allows them to gain entry into host nests (L. personata, B. fimbriatus, and M. andrenae) or to parasitize adults (Z. cinereum), these parasites seem not to impact upon the dynamics of the host A. agilissima at the nesting aggregation, and the host possesses traits to reduce parasitism.     

Structure of insect community in the fungus Inonotus radiatus in riparian boreal forests

Basidiomes of polypore fungi host many insects. Yet systematic information about insect assemblages from most fungal species is lacking. We studied the insect community associated with the wood-decaying fungus Inonotus radiatus (Sowerby: Fr.) P. Karst. (Hymenochaetales). More specifically, we studied the effect of successional stage and weight of basidiomes, as well as shore exposition (north or south), on species richness and composition, as well as occurrence and abundance of the most abundant fungivores. Basidiomes were collected from riparian forests at five lakes in Sweden. Insects were reared out from the basidiomes in the laboratory. A total of 5645 adult insect individuals of 117 taxa were obtained. Among these, 2782 specimens of 36 taxa use Inonotus radiatus basidiomes as breeding habitat. Eight species of parasitic wasps were new to Sweden. The most abundant fungivore was Ennearthron cornutum (Ciidae), which is a generalist breeding in many polypore species. Based on our material and literature, the melandryid beetles Abdera affinis and A. flexuosa seem to be specialists on the order Hymenochaetales. Other frequent fungivores were Dorcatoma dresdensis (Ptinidae), and the lepidopterans Archinemapogon yildizae, Nemapogon cloacellus and N. picarellus (Tineidae). The parasitoid community associated with the tineid moths was similar to the community associated with the other polypore species in the order Polyporales. In contrast, the parasitoids associated with Dorcatoma (and/or Abdera) were different from the other Polyporales species, suggesting that the fungal host species is more important for these parasitoid species than the beetle host species itself. The most abundant and frequent parasitoids were the braconids Diospilus dispar and Colastes fritzeni, which both parasitise Dorcatoma. Species richness was significantly smaller in fresh than in more decayed basidiomes, but species composition did not differ. There was no difference in species composition or richness between north and south shorelines.     

Distinct parasitoid communities associated with host races of the leaf-mining moth Acrocercops transecta on distantly related host plants (Juglandaceae and Ericaceae)

Differences in host-plant species may affect the structure of parasitoid communities of phytophagous insects. Phytophagous insect species often consist of host races, and comparisons between host races enable us to infer the effects of host-plant differences on parasitoid communities. Here, we compare the parasitoid fauna of two host races of the leaf-mining moth Acrocercops transecta, which are associated with either Juglandaceae or Lyonia ovalifolia (Ericaceae). Field collection and subsequent laboratory rearing revealed a significantly higher parasitization ratio in the Juglandaceae race than in the Lyonia race. In the Juglandaceae race the majority of parasitoids were Braconidae, whereas in the Lyonia race the majority were Eulophidae. Furthermore, even within the same host race the most abundant parasitoid species differed between populations with different host associations. Hence, the present results strongly indicate that different host-plant taxa affect the structure of parasitoid communities even within a single phytophagous insect species.     

Phylogenetic evaluation of the tropical Camptotypus genus-group (Hymenoptera: Ichneumonidae), with a key to the world genera

The phylogeny of the disjunct pantropical Camptotypus genus-group (Hymenoptera, Ichneumonidae, Pimplinae) was reconstructed using 40 morphological characters.The monophyly of many established genera (Parvipimpla, Clydonium, Odontopimpla) was found to be supported, but Hemipimpla was found to nest within Camptotypus and has thus been treated as a junior synonym of it (syn. n.). A group of undescribed Neotropical species were found to form a separate monophyletic group, herein referred to as Genus A. Species of the Neotropical genus Zonopimpla formed a basal paraphyletic grade which could not satisfactorily be resolved into discrete monophyletic units. Rather than subsuming all the more derived genera within one large group, or attempting to erect numerous new and probably ephemeral genera (for isolated species of Zonopimpla), we propose the retention of this grade-group as a separate genus at present. The host utilization patterns of the genus-group, and the biogeography of the genus-group are discussed. The group is hypothesized as having originated and radiated in the Neotropics, as ectoparasitoids of weakly concealed lepidopterous hosts. One lineage, Clydonium, has become specialized at attacking hosts in woody galls where it utilizes coleopterous as well as lepidopterous hosts. The Old World genera (Camptotypus and Parvipimpla) comprise a monophyletic group and are suggested to have originated from a single dispersal event across the widening Atlantic. No evidence for either a southern or northern circum-polar dispersal was observed in this entirely tropical genus-group. One lineage in the Afrotropical region has switched from attacking lepidopterous hosts, to attacking pre-pupal and pupal polistine vespids.     

A new genus and new species (Orthoptera: Acrididae: Gomphocerinae) from south-east Xizang,China

In 2007, a 3-year survey was started in northeastern Italy to assess the influence of agricultural management on parasitoid composition and parasitism rates of parasitoids attacking larvae and pupae of Syrphidae (Diptera). Two types of management were considered – intensive (including wheat and maize monoculture with very low or null vegetation diversity) and extensive (with different types of habitats, characterized by a higher vegetation complexity). In habitats simplified by intensive agriculture, and particularly in maize fields, parasitism rates of syrphid larvae were higher than recorded in more diversified habitats. A substantial difference in the composition of the parasitoid complex was also found. In extensive and wheat sites, Diplazontinae (Braconidae), highly specialized to Syrphidae, were the dominant parasitoids, whereas in maize fields Pachyneuron (Pteromalidae) species, known as generalists, accounted for more than 80% of total parasitoids. In the habitats where Pachyneuron spp. were dominant, parasitism rates were very high. Conversely, in habitats where Diplazontinae were more abundant, the parasitism rates were very low. The higher rate of parasitism of syrphid larvae and pupae recorded in intensive monocultures could have a large impact on the development and control of aphid populations in such crops.     

Community composition of laelapine mites (Acari: Laelapidae) associated with the nests and fur of Cerradomys subflavus (Wagner, 1842)

Knowledge about the role of the nest environment in the life cycle of laelapine mites associated with neotropical small mammals is unclear because nests have rarely been collected. Here we use infestation data to make predictions about host–mite ecology and examine the nests of the host to assess the validity of these predictions. Spool-and-line method was used to track Cerradomys subflavus nests. We found that: the structure of mite populations infesting the body of C. subflavus, was strongly biased toward the reproductive female stage; recaptured hosts were re-infested with laelapine mites at different rates; and female hosts were not more highly infested than males. We found strong evidence of nidicoly with both, Gigantolaelaps vitzthumi and Androlaelaps fahrenholzi. The G. vitzthumi was more abundant in the nests of female rodents and with both these mite species, we encountered males and immature mites in the nests, although not in the host fur.     

Attachment site selection of Hydrachna and Eylais (Acari: Hydrachnellae) watermite larvae infecting Corixidae (Hemiptera: Heteroptera)

The sites of attachment of four larval watermite species were recorded from four corixid species in Lough Corrib, Ireland. Parasitic watermites exhibited preferences in respect of attachment sites on the corixid hosts and these patterns were consistent for all host species examined. Hydrachna cruenta utilized areas of the host which were in direct contact with the water, the distribution of larvae reflecting the general behaviour and microdistribution of the hosts. Hydrachna conjecta infected the ventral hemielytra with a significant asymmetrical distribution on the right hemielytron, resulting from the folding pattern of the hemielytra and the larval route of entry to the subhemielytral air space. Eylais species attached to the abdominal terga of the host, E. infundibulifera infecting the anterior two terga and E. discreta terga three and four. Cymatia bonsdorffi was not parasitized by E. infundibulifera and on this host the majority of E. discreta attached to the second abdominal tergite. Eylais species infecting small corixid hosts may avoid interference competition with H. conjecta by attaching to the left side of the abdomen. Attachment site selection may contribute towards alleviation of intraspecific and interspecific interference competition for larval growing space.     

Anatomy,reach and classification of the parasitoid complex of a common British moth,Anthophila fabriciana (L.) (Choreutidae)

The small plurivoltine moth Anthophila fabriciana is widespread and often abundant in Britain wherever its main larval foodplant, stinging nettle, occurs. It overwinters as a larva (first generation) then has one or more partly overlapping summer broods (notionally second generation). A total of 5017 larvae were collected and reared from widely distributed populations in Britain, and the resulting 2167 host mortalities due to parasitoids were assessed. Small collections of pupae were also made. Altogether 25 parasitoid species, including secondary parasitoids, were found. Larval parasitism was heaviest in the second generation. In each generation there was a dominant parasitoid that was absent from the other. Summary information on the developmental biology and host range (expressed quantitatively and resulting from very broad rearing programmes) for each parasitoid is given. They are classed as ‘absolute specialists’, ‘taxon oligospecialists’, ‘niche oligospecialists’, ‘niche generalists’, ‘casuals’ and ‘strays’. Both kinds of oligospecialists can be ‘paraspecialists’ if only one potential host occurs locally. Although constructing a quantitative food web is not appropriate, providing both source web and sink web data in quantitative form enables the parasitoid complex to be understood in the wider context of the ecosystem, necessary for both biodiversity and nature conservation interests. In Appendix 1, parasitoids reared from other European Choreutidae are listed quantitatively.     

A phytophagous braconid,Allorhogas conostegia sp. nov. (Hymenoptera: Braconidae), in the fruits of Conostegia xalapensis (Bonpl.) D. Don (Melastomataceae)

Before 1989 all braconid wasps were thought to be parasitoids, but in that year the first phytophagous species was reported. Subsequently, a few other examples of phytophagy have been discovered, most of which are species of Allorhogas in the subfamily Doryctinae. Until now, all demonstrated examples of phytophagy in this genus have been as gall inducers in the fruits of Fabaceae. Here we describe a new species from Costa Rica, Allorhogas conostegia Marsh and Shaw, and provide evidence that it forms galls in the fruits of Conostegia xalapensis (Melastomataceae). We also provide information on the phenology of the plant and of the galls and the effects of the galls on the host plant, and we discuss the potential species richness of Allorhogas in the Neotropics.     

Reproductive biology and strategies of nine meloid beetles from Central Europe (Coleoptera: Meloidae)

The reproductive biology of meloid species from Central Europe is investigated by means of laboratory breeding experiments. They show that the total reproductive potential of meloids, with up to 40,000 eggs, largely exceeds prior data. Furthermore, the number of laid eggs strongly relates to the way by which the triungulins find their host. Clutch size correlates significantly with the size of the beetle, while egg size is independent of this. Both clutch size and egg size decrease with each oviposition. For some species, reproductive data are used to demonstrate how they have adapted to their habitat and to point out existing trade‐offs. Based on the obtained results and on an evaluation of the relevant literature, three types of reproductive strategy can be distinguished within the meloids: (1) very high reproductive rates of open field species with phoretic larvae; (2) average reproductive rates of forest populations of species with phoretic larvae and of species the larvae of which search the nests of their hosts actively; and (3) small to average reproductive rates of those species depositing their clutches in the immediate proximity of their hosts' nests.