New damselfly hosts and species identification of an aquatic parasitoid Hydrophylita emporos (Hymenoptera: Trichogrammatidae) in Taiwan

ABSTRACT

The host–parasitoid relationship and species identity of aquatic parasitoids of two damselflies Coeliccia cyanomelas (Platycnemididae) and Psolodesmus mandarinus dorothea (Calopterygidae) from Fushan and Lienhuachih in Taiwan were studied using morphological characters and DNA barcoding sequences. The parasitoids reared from the damselflies’ eggs, and the field-collected parasitoids, were morphologically identified as Hydrophylita emporos (Trichogrammatidae), a recently described parasitoid of the damselfly P. m. mandarinus from Northern Taiwan. The CO1 (cytochrome c oxidase I) gene tree supported the identification as H. emporos, as well as all parasitoid samples from C. cyanomelas, P. m. dorothea and P. m. mandarinus. The sampled H. emporos populations did not differ genetically despite their different host associations. However, some genetic differences were found between H. emporos populations from Northern and Central Taiwan, indicating that the dispersal of H. emporos may be limited by geographical distances. Our results suggest that H. emporos can parasitise not only closely related sister subspecies, P. m. mandarinus and P. m. dorothea, but also phylogenetically distant species of another damselfly family, C. cyanomelas. This is the first record of multiple damselfly hosts for the aquatic parasitoid genus Hydrophylita. This finding implies that the host range of H. emporos and congeneric species may be broader than previously thought.     

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.     

Aphidiinae parasitoids (Hymenoptera: Braconidae) of Macrosiphoniella aphids (Hemiptera: Aphididae) in the western Palaearctic region

The parasitoids (Hymenoptera: Braconidae: Aphidiinae) of Macrosiphoniella del Guercio (Hemiptera: Aphididae) from the western Palaearctic region are reviewed and keyed. The host associations and distribution of each species are summarized. Aphidius stigmaticus sp. nov. reared from Macrosiphoniella tanacetaria on Tanacetum polycephalum, which is another member of the Macrosiphoniella aphid parasitoid guild from high-mountain habitats in western Iran, is described in the present contribution. Some taxonomic problems within Macrosiphoniella parasitoid guilds are discussed as well.     

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.     

Ecological notes of the alien species Godiva quadricolor (Gastropoda: Nudibranchia) occurring in Faro Lake (Italy)

The first record from Sicily of the introduced facelinid nudibranch Godiva quadricolor allowed the detection of trophic relationships with the polycerid Polycera hedgpethi, another non-native nudibranch, and with two bryozoan species, namely the naturalized Cheilostomatida Bugula neritina and the cryptogenic Ctenostomatida Amathia verticillata. The settlement of both nudibranchs was presumably promoted by a trophic shift of P. hedgpethi from the natural prey B. neritina towards the largely available and not exploited A. verticillata. This short food web, without evident links with native fauna and having G. quadricolor as the top predator, is described. A DNA barcoding approach was used to confirm the identity of this facelinid species and to explore the possible genetic divergence occurring among the samples analysed.     

Systematics and fossil evidence of host-parasitoid relationships of Calamiscus Borgmeier (Diptera: Phoridae)

The life history and hitherto unknown males of Calamiscus cryptopalpis are described. A second species of this genus, C. inflativenis n. sp. is described from Costa Rica. The species C. cryptopalpis is a gregarious parasitoid of injured stingless bees in Costa Rica. The co-occurrence of a specimen of the genus Calamiscus in a piece of amber with the oldest known fossil bee is hypothesized to be the most ancient host-parasitoid relationship between a pair of insect species documented thus far. The otherwise rarely-seen fly was probably lured to the amber site by the alarm pheromones of the bee, and then trapped in the sticky tree resins.     

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.     

Generalist hymenopteran parasitoids of the leaf-miner Chromatomyia fuscula (Zett.) (Dipt.: Agromyzidae)

In southeastern Norway, by the last week of June (1992), 48%of Chromatomyia fuscula emerged, based on the number of mines collected from Hordeum vulgare leaves. Among the parasitoids Diglyphus begini was the dominant species, but Cirrospilus vittatus and Neochrysocharis aratus also emerged in significant numbers. During June and July, N. aratus was the dominant species in the 1st and 2nd leaves, but D. begini was dominant in the 4th to 6th leaves. C. vittatus and Cyrtogaster vulgaris affected all larvae of the leaf-miner fly, living on various leaves. In central Norway, 48% of C. fuscula also emerged. C. vulgaris and D. begini were the dominant parasitoids and C. vittatus was also found. C. fuscula has no specialist parasitoids. Adults of the generalist parasitoids of the leafminers always search for different hosts. In investigation of the parasitoid complex of a lepidopteran leaf-miner, Phyllocnistis labyrinthella (living in Populus tremula leaves at the edge of the examined barley field), three shared parasitoids (C. vittatus, Pnigalio soemius and Pnigalio pectinicornis) were found. The complex network of host plant-phytophagous insect-insect parasitoid-facultative hyperparasitoid relationship is discussed.     

On the parasitoid community of Aulacidea laurae (Hym., Cynipidae), with description of a new species of Adontomerus (Hym., Torymidae)

An account is given of parasitoids (Hym., Chalcidoidea) reared from Spanish galls of Aulacidea laurae (Hymenoptera, Cynipidae) which develop in achenes of Scorzonera laciniata (Asteraceae). The parasitoid community includes Adontomerus brevicaudatus (Hym., Torymidae) which is described as new. Final instar larva of the new species is also described. The generic placement of A. brevicaudatus and the composition of the parasitoid community are discussed.     

Identification of Megaselia scalaris (Loew, 1866) (Diptera: Phoridae) in mummified human body from Itacambira (MG), Brazil,using scanning electron microscopy and cuticular hydrocarbons

Insect puparia were found adhered to the ribs and other tissues in the abdominal cavity of a natural male mummy found in Itacambira (State of Minas Gerais, Brazil) dating to the Colonial Period. They were collected for identification by scanning electron microscopy, and for comparison of several morphological features with those described in the literature. The puparia were found open and dorsoventrally flattened, making it difficult to visualize the dorsal projections. The tegument is covered by tapered spines and contains rows of small tubercles on the dorsal and lateral regions of the puparium. The posterior spiracle consists of four parallel openings arranged in pairs. These results are indicative that the specimens belong to the species Megaselia scalaris (Loew, 1866) (Diptera: Phoridae). Additionally, cuticular hydrocarbons of the puparia were analysed by gas chromatography coupled to mass spectrometry and compared with the profile of M. scalaris reared in the laboratory.     

The preimaginal stages of Minotetrastichus frontalis (Nees) and Chrysocharis laomedon (Walker) (Hymenoptera: Eulophidae), parasitoids associated with Phyllonorycter issikii (Kumata) (Lepidoptera,Gracillariidae)

The biology and morphology of the early larval instars of Minotetrastichus frontalis (Nees) are redescribed and the morphology of the preimaginal stages of Chrysocharis laomedon (Walker) (Hymenoptera: Eulophidae) are described in detail for the first time. Both species are larval-pupal parasitoids of Phyllonorycter issikii (Lepidoptera, Gracillariidae), which is associated with Tilia sp. The female of M. frontalis lays her egg beside the host larva and the newly hatched first instar larva moves and has to find the host larva. The larva of P. issikii tries to continue its feeding but never pupates after paralysis. The female of C. laomedon lays its egg inside the cuticle of the larva of P. issikii, but the newly hatched parasitoid larva vacates the host larva and develops externally on its surface as an ectoparasitoid. Such behaviour might be facilitated by the constant temperature and humidity inside the mine. Some peculiarities of parasitoid–host relationships are described and discussed.     

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.     

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.     

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.     

Meteorus acerbiavorus sp. nov. (Hymenoptera,Braconidae), a gregarious parasitoid of Acerbia alpina (Quensel) (Lepidoptera,Arctiidae) in North Finland

The new gregarious parasitoid Meteorus acerbiavorus sp. nov. (Hymenoptera, Braconidae) was reared from the cocoons of Acerbia alpina (Quensel) (Lepidoptera, Arctiidae) in north-western Finnish Lapland. This species belongs to Meteorus rubens (Nees) species group and differs from the most related M. rubens in the following features: the eyes densely setose; the median lobe of the mesoscutum, scutellum, mesopleuron, and the hind coxa entirely or at least partly rugulose-granulate or rugose-areolate and sometimes with granulation; the ovipositor subapically with distinct dorsal node; the ventral borders of the first metasomal tergum weakly separated by narrow space in its basal half; the colour of the body and legs mostly or entirely dark; the fore wing more or less darkened. Phylogenetic relationships among several Meteorus species close to M. rubens including new M. acerbiavorus were investigated based on DNA sequence fragments of the mitochondrial COI and the nuclear 28S rDNA genes. The discussions on the species groups of Meteorus, on distribution of Acerbia alpina in the Holarctic and on its known parasitoids are presented.     

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).     

Diet composition of an invasive population of Lithobates catesbeianus (American Bullfrog) from Argentina

The American bullfrog Lithobates catesbeianus has been introduced around the world, with invasive populations reported from almost all South American countries. A population of this species was introduced in the Calingasta department of San Juan province, which is an arid environment in western Argentina. This work provides information on the dietary composition of an invasive population of L. catesbeianus, and compares the degree of dietary overlap between adults and juveniles. Stomach contents of 169 bullfrogs (82 adults and 87 juveniles) were analysed. Adults consumed 40 prey taxa and Hymenoptera (Insecta) was the most numerous prey item (41.8%), followed by Araneae (13.6%) and Aeglidae (13.4%). Juveniles consumed 29 prey taxa and Hymenoptera constituted the highest percentage in prey number (77.2%). The trophic overlap niche index at the same level shows a value of 0.64 overlap in dietary community between adults and juveniles of this bullfrog. Aeglidae was volumetrically the most important trophic item (25.4%), followed by Anura (25.02%). Our results showed that cannibalism in bullfrogs is more common than the consumption of native anurans, coinciding with that reported in other populations of introduced bullfrogs. The high similarity in the diets of both size classes and the association between the size of the predator and prey suggest that the impact caused by bullfrogs throughout their ontogeny is high and probably has an impact on their prey. Freshwater crabs are the main items in the diet of Lithobates catesbeianus in other introduced populations and are usually the most conspicuous at our study site. The crabs in freshwater ecosystems are part of the lowest trophic level in the food chain. The major threats to the southern region’s freshwater crabs include deforestation, farming and exotic species. Lithobates catesbeianus has a generalist diet and high overlap between adults and juveniles.     

Host ranges of parasitoids (Hymenoptera: Braconidae and Ichneumonidae) reared from Epermenia chaerophyllella (Goeze) (Lepidoptera: Epermeniidae) in Britain,with description of a new species of Triclistus (Ichneumonidae)

The complex of parasitoids attacking larval stages of Epermenia chaerophyllella in Britain is outlined, and the host range of each of the species recorded (three braconids and four ichneumonids) is discussed. ‘Ecological’ factors, such as the behavioural similarity of unrelated hosts during the vulnerable parts of their life histories, seem to have been important in the evolution of the host associations of several of these parasitoids, even though they are koinobionts. A new species, Triclistus epermeniae Shaw and Aeschlimann, is described.     

The Mollissima subgroup of fannia Desvoidy,with the description of a new species from Burma and a revised key to species (Diptera: Muscidae)

Synopsis

A comparison of the type-species of the genus Coelomyia Haliday (C. mollissima Haliday) with that of the genus Fannia Desvoidy (F. scalaris Fabricius) shows that these two genera cannot be maintained as equivalent phylogenetie taxa. Coelomyia is synonymized with Fannia and its species placed in Fannia, four new combinations thus being established. The mollissima subgroup is defined and its affinities shown, a new species is described, and a revised key to species is given.     

Biology and defensive secretion of myrmecophilous Thiasophila spp. (Coleoptera: Staphylinidae: Aleocharinae) associated with the Formica rufa species group

Various aspects of the biology of the myrmecophilous Thiasophila angulata and T. szujeckii (Staphylinidae; Aleocharinae), mainly associated with the Formica rufa species group, are studied. The composition of defensive gland secretions from adults of T. angulata has been analysed. The complete development (egg-L_1-3-pupa-adult form), including adult overwintering, of both species takes place solely in ant nests. Under laboratory conditions, this lasts for about 22 days in T. angulata, and the period elapsing between the old and new generations of insects is approximately two months. The mean fecundity of adults of this species (28 eggs) was determined, as well as the duration of its reproductive period (mean 28.5 days), and lifespan (mean 67 days). In natural conditions T. angulata, found within nests of Formica polyctena and F. rufa, produces three generations, and the sibling species T. szujeckii one generation per year. These two species differ in their phenologies and abundance dynamics of adults and larvae, which is linked to the size and temperature regime of the host nest. The results of this study uphold the recent separation of T. angulata and T. szujeckii based on morphological features of adults and selected molecular markers. Adults and larvae of T. angulata forage on both live and dead food items accumulated by the host, as well as on the host’s eggs and larvae. Both the beetle larvae and adults remain unmolested among the host workers. The adults use their defensive gland secretions, which contains substantial quantities of toxic quinones, when necessary. According to the current categories of myrmecophiles, T. angulata (and by analogy, T. szujeckii as well) should be classified as a species wholly integrated with the host.