Wetland species of aculeate Hymenoptera are poorly known, even though many of them may serve as diagnostic or flagship species in nature conservation. Here we examined 6,018 galls induced ≥1 year prior their collection by the chloropid flies Lipara spp. The galls were collected at 34 sites in Central Europe. We examined 1,389 nests (4,513 individuals) of nine species, part of which were parasitized by one dipteran and two chrysidid parasitoid species. We describe the nests of seven dominant species and larvae of four species (Pemphredon fabricii, Trypoxylon deceptorium, Hoplitis leucomelana and Hylaeus pectoralis) and two parasitoids (Trichrysis cyanea and Thyridanthrax fenestratus, both in nests of Pemphredon fabricii and Trypoxylon deceptorium). All the species, but H. pectoralis, preferred robust galls at very thin stalks (induced typically by Lipara lucens) over the narrow galls on thick stalks. The larvae of P. fabricii and T. deceptorium resembled strongly their sibling species (Pemphredon lethifer and Trypoxylon attenuatum sensu lato, respectively). The larvae of T. fenestratus showed features different from those previously described. By hatching set of another 10,583 galls induced by Lipara spp. ≥1 year prior their collection, we obtained 4,469 individuals of 14 nesting hymenopteran species, two cleptoparasites, three chrysidid and one dipteran parasitoid. Of these species, four new nesting species have been recorded for the first time in galls induced by Lipara spp.: Chelostoma campanularum, Heriades rubicola, Pseudoanthidium lituratum and Hylaeus incongruus. We also provide first records of their nest cleptoparasites Stelis breviuscula and Stelis ornatula, and the parasitoid Holopyga fastuosa generosa. Thyridanthrax fenestratus formed strong populations in nests of Pemphredon fabricii and Trypoxylon deceptorium, which are both newly recorded hosts for T. fenestratus. The descriptions provided here allow for the first time to identify the larvae of the most widespread central European aculeate hymenopteran reed gall specialists.
The ability of aculeate Hymenoptera to utilize wetlands is poorly understood, and descriptions of their nests and developmental stages are largely absent. Here we present results based on our survey of hymenopterans using galls induced by Lipara spp. flies on common reed Phragmites australis in the years 2015–2016. We studied 20,704 galls, of which 9,446 were longitudinally cut and the brood from them reared in the laboratory, while the remaining 11,258 galls reared in rearing bags also in laboratory conditions. We recorded eight species that were previously not known to nest in reed galls: cuckoo wasps Chrysis rutilans and Trichrysis pumilionis, solitary wasps Stenodynerus chevrieranus and Stenodynerus clypeopictus, and bees Pseudoanthidium tenellum, Stelis punctulatissima, Hylaeus communis and Hylaeus confusus. Forty five species of Hymenoptera: Aculeata are known to be associated with reed galls, of which 36 make their nests there, and the other are six parasitoids of the family Chrysididae and three cuckoo bees of the genus Stelis. Of these species, Pemphredon fabricii and in southern Europe also Heriades rubicola are very common in reed galls, followed by Hylaeus pectoralis and two species of the genus Trypoxylon. We also found new host-parasite associations: Chrysis angustula in nests of Pemphredon fabricii, Chrysis rutilans in nests of Stenodynerus clypeopictus, Trichrysis pumilionis in nests of Trypoxylon deceptorium, and Stelis breviuscula in nests of Heriades rubicola. We provide new descriptions of the nests of seven species nesting in reed galls and morphology of mature larvae of eight species nesting in reed galls and two parasitoids and one nest cleptoparasite. The larvae are usually very similar to those of related species but possess characteristics that make them easy to distinguish from related species. Our results show that common reeds are not only expansive and harmful, but very important for many insect species associated with habitats dominated by this plant species.
Gravel-sand river terraces were nearly eliminated from central European landscape by river channelization. Monotypic stands of common reed (Phragmites australis) growing on such terraces are often stressed by drought, which makes them vulnerable to Lipara spp. (Diptera: Chloropidae) gallmakers. Although Lipara are considered ecosystem engineers, only fragmentary information is available on the biology of their parasitoids and inquilines. We analyzed the assemblages of arthropods (Arachnida, Collembola, Dermaptera, Psocoptera, Thysanoptera, Hemiptera, Raphidioptera, Communicated by P. Ponel. Electronic supplementary materialThe online version of this article (Neuroptera, Coleoptera, Diptera, Lepidoptera and Hymenoptera) that emerged from 17,791 Lipara-induced galls collected in winter from 30 reed beds in the Czech Republic, 15 of which were situated at (post)industrial sites (gravel-sandpits, tailing ponds, limestone quarries, colliery dumps, and reclaimed lignite open-cast mines) and 15 were in nearnatural habitats (medieval fishponds, and river and stream floodplains). The Chao-1 estimator indicated 229.3 ± 18.1 species in reed galls at (post)industrial and 218.1 ± 23.6 species at near-natural sites, with the Sørensen index reaching only 0.58. We identified 18 red-listed species and four new species for the Czech Republic (Gasteruption phragmiticola, Echthrodelphax fairchildii, Haplogonatopus oratorius and Enclisis sp.), representing mostly obligate (64 %) or facultative (9 %) reed specialists. We propose that Lipara gallassociated assemblages undergo a long-term cyclic ecological succession. During first 10 years after reed bed formation, only Lipara spp. and several other species occur. During next decades, the reed beds host species-rich assemblages with numerous pioneer species (Singa nitidula, Polemochartus melas) that critically depend on presence of prior disturbances. Middle-aged reed beds (near medieval fishponds) are prevalently enriched in common species only (Oulema duftschmidi, Dimorphopterus spinolae). Habitats with the longest historical continuity (river floodplains) host again species-rich assemblages with several rare species that probably require long-term habitat continuity (Homalura tarsata, Hylaeus moricei). Landscape dynamics is thus critical for the persistence of a full spectrum of reed gall inquilines, with (post)industrials serving as the only refugia for pioneer species ousted from their key nesting habitats at once cyclically disturbed gravel-sand river terraces.
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