The first checklist of black fungus gnats (Diptera: Sciaridae) of Morocco

The first checklist of black fungus gnats is established; new faunistic records of Sciaridae are presented, providing a list of 10 genera and 55 species. Forty-eight species are newly listed for Morocco, increasing the total of Sciaridae known from Morocco to 69 species, belonging to 12 genera, of which six (Austrosciara Schmitz & Mjöberg, 1924, Bradysiopsis Tuomikoski, 1960, Epidapus Haliday, 1851, Lycoriella Frey, 1942, Pseudolycoriella Menzel & Mohrig, 1998 and Sciara Meigen, 1803) are newly reported for Moroccan fauna.     

Dark winged fungus gnats (Diptera: Sciaridae) collected from shiitake mushroom in Korea

Seven species of sciarid flies were collected in shiitake mushroom farm in Korea. Among them, Lycoriella ingenua (Dufour 1839) and Bradysia difformis Frey 1948 were dominant as possible pests of the shiitake mushroom because the larvae were found on both oak bed logs and in the artificial sawdust beds for shiitake cultivation. Five other species, which were collected in lower numbers, are reported for the first time in Korea: Bradysia longimentula (Sasakawa 1994), Bradysia trispinifera Mohrig & Krivosheina 1979, Leptosciarella (Leptospina) subdentata (Mohrig and Menzel 1992), Scatopsciara camptospina Mohrig and Mamaev 1990, and Xylosciara inornata Mohrig and Krivosheina 1979.     

Digestive system of larval black flies (Diptera: Simuliidae): structure and phylogeny

Abstract. Larval gut characters of 45 species of black flies, representing 17 subgenera and 15 genera, were studied for their utility in taxon diagnosis and phylogenetic inference. Phase-contrast and scanning electron microscopy revealed a wealth of new characters in the esophageal armature of the foregut. Larval gut morphology, particularly characters of the esophageal armature, varied significantly among taxa, permitting generic diagnoses. A cladogram inferred from 13 gut characters supported a sister-group relationship between Parasimulium and the remaining simuliids, but did not show a strong phylogenetic signal for the current, well-supported concept of tribal level monophyly.     

Molecular phylogeny of the fungus gnat family Mycetophilidae (Diptera,Mycetophiliformia)

Abstract A molecular phylogeny of the fungus gnat family Mycetophilidae based on the nuclear 18S, 28S, and the mitochondrial 16S rRNA genes is presented. The total alignment included 58 taxa and 1704 bp. The family was recovered as monophyletic in parsimony and Bayesian analyses. In the Bayesian analysis, Mycetophilinae and its two tribes, Mycetophilini and Exechiini, were monophyletic with good statistical support. The subfamily Mycomyinae was found consistently in a sister‐group relationship to Mycetophilinae. Gnoristinae was rendered paraphyletic, subtending Mycomyinae and Mycetophilinae. Within Gnoristinae, the genera Coelosia Winnertz, Boletina Staeger, Gnoriste Meigen group with Docosia Winnertz, usually considered to be a member of Leiinae. No support was found for the monophyly of the subfamilies Sciophilinae and Leiinae.     

Molecular phylogeny of the fungus gnat tribe Exechiini (Mycetophilidae, Diptera)

The phylogenetic relationships within the fungus gnat tribe Exechiini have been left unattended for many years. Recent studies have not shed much light on the intergeneric relationship within the tribe. Here the first attempt to resolve the phylogeny of the tribe Exechiini using molecular markers is presented. The nuclear 18S and the mitochondrial 16S, and cytochrome oxidase subunit I (COI) genes were successfully sequenced for 20 species representing 15 Exechiini genera and five outgroup genera. Bayesian, maximum parsimony and maximum likelihood analyses revealed basically congruent tree topologies and the monophyly of Exechiini, including the genus Cordyla , is confirmed. The molecular data corroborate previous morphological studies in several aspects. Cordyla is found in a basal clade together with Brachypeza , Pseudorymosia and Stigmatomeria . The splitting of the genera Allodiopsis s.l. and Brevicornu s.l. as well as the sistergroup relationship of Exechia and Exechiopsis is also supported. The limited phylogenetic information provided by morphological characters is mirrored in the limited resolution of the molecular markers used in this study. Short internal and long-terminal branches obtained may indicate a rapid radiation of the Exechiini genera during a short evolutionary period.     

Molecular phylogeny of fungus gnats (Diptera: Mycetophilidae) revisited: position of Manotinae,Metanepsiini, and other enigmatic taxa as inferred from multigene analysis

The phylogeny of selected genera from four subfamilies of fungus gnats (Diptera: Mycetophilidae) – Manotinae, Leiinae, Sciophilinae and Gnoristinae (including Metanepsiini) – is reconstructed based on the combined analysis of five mitochondrial (12S, 16S, COI, COII, cytB) and two nuclear (28S, ITS2) gene markers. Results of the different analyses all support Manotinae as a monophyletic group, with Leiinae as the sister group. Allactoneura DeMeijere is nested in the monophyletic and strongly supported clade of Leiinae. The tribe Metanepsiini is revealed as paraphyletic and the genera Metanepsia Edwards and Chalastonepsia Søli do not appear to be closely related. The genera Docosia Winnertz, Ectrepesthoneura Enderlein, Novakia Strobl and Syntemna Winnertz were placed with a group of genera included traditionally in the Gnoristinae. The monophyly of Dziedzickia Johannsen and Phthinia Winnertz is not supported. The genera of Sciophilinae (excluding Paratinia Mik but including Eudicrana Loew) form a monophyletic group in the Bayesian model.     

First fungus gnats of genus Manota Williston (Diptera: Mycetophilidae) from Japan

The genus Manota is recorded from Japan for the first time. Three new species, Manota satoyamanis, Manota indahae and Manota tunoae spp. nov., are described, based on specimens collected in an ecological sampling program of arthropods in the “satoyama” landscape of Ishikawa Prefecture. “Satoyama” represents the traditional rural landscape of Japan, which is characterized by a mosaic of secondary forests, plantations, ponds and rice paddy fields. The new species raise the number of Palearctic Manota species from five to eight.     

Molecular identification of dipteran pests (Diptera: Sciaroidea) from shiitake mushroom

On shiitake farms, mycophagous maggots can cause serious damage by preventing formation of the fruiting body. Recently, these pests have significantly reduced shiitake production in Korea. However, larvae and female adults cannot be identified due to their lack of morphological characteristics. Therefore, farmers and applied entomologists are unable to determine which species is the primary cause of the shiitake damage. In this study, mycophagous flies (colonized larvae) were collected from damaged shiitake farms and subsequently identified by matching identified males with the cytochrome c oxidase subunit I (COI) sequences from the larvae. Divergences of the COI sequences among the species discriminated the clusters clearly, and the mycophagous pests were identified as Camptomyia corticalis and C. heterobia. Interestingly, these two species coexisted under the bark of shiitake oak bed logs.     

Molecular phylogeny and host use evolution of the genus Exorista Meigen (Diptera: Tachinidae)

Members of the genus Exorista are parasitoids of a diverse array of insect hosts in the orders, Lepidoptera, Hymenoptera, Mantodea and Orthoptera. Phylogenetic relationships among subgenera and species of Exorista were inferred using four nuclear (Tpi, white, 18S and 28S) and four mitochondrial DNA (16S, 12S, ND5 and CO1) genes in maximum parsimony (MP), maximum likelihood (ML) and Bayesian Markov chain Monte Carlo (MCMC) analyses. Separate trees based on different sets of genes (mt DNA, nuclear, ribosomal, etc.) were compared and found to be nearly concordant. According to the molecular tree generated from the concatenated sequence data, the genus Exorista is paraphyletic. The phylogenetic analyses indicate the existence of two major clades of Exorista, including two genera Parasetigena and Phorocera. Morphological traits supporting clades indicated by molecular analyses within this genus are evaluated. Evolutionary patterns of the host use and host shifts are examined by optimizing host information using maximum likelihood on the molecular phylogeny. The ancestral host group of the tribe Exoristini (excluding Ctenophorinia and Phorinia) appears to be the order Lepidoptera, although hosts of some species are unknown. A major host shift to the Hymenoptera occurred in the clade of subgenus Adenia, and the ancestral state of subgenus Spixomyia is equivocal because there is little information available on the hosts in members of a subclade of this group (subclade A: Exorista hyalipennis group).     

Molecular phylogeny and timing of evolution of Anthomyza and related genera (Diptera: Anthomyzidae)

Phylogenetic hypotheses of the relationships of Diptera: Anthomyzidae (61 taxa) are discussed with special reference to the genera Fungomyza Rohá?ek, 1999, Anthomyza Fallén, 1810, Epischnomyia Rohá?ek, 2006 and Arganthomyza Rohá?ek, 2009 based on the analysis of 7 combined mitochondrial + nuclear gene markers in comparison with results of the most recent cladistic analysis of morphological characters. The majority of revealed inter‐ and intrageneric relationships of these genera in both analyses were largely congruent except for the topology of Fungomyza and Arganthomyza being equivocal because of different topologies in phylograms generated by alternative molecular methods and Epischnomyia forming a branch within Anthomyza in the molecular data hypothesis. The formerly unsettled Anthomyza drachma proved to be the sister species of the A. umbrosa group while the affinity of A. flavosterna to the A. bellatrix group has not been confirmed. The first phylogenetic hypothesis of Anthomyzinae to include timing of the nodes of divergence is presented. The origin of the subfamily is dated into the Eocene, ca 37.5 (30.6 – 45.3) MYA, and agrees with the age of the oldest known fossils from Baltic amber. The analysed members of the Anthomyza group of genera were found to have already evolved in the upper Oligocene. The divergences of the Palaearctic and Nearctic sister species in Arganthomyza and Anthomyza were found to occur at several different times during the Neogene (upper Miocene) to lower Quaternary (Pleistocene), from about 7 to 0.7 MYA. These were likely the result of fragmentation of widespread (Holarctic or Sino‐Japanese–Nearctic) ranges of ancestral taxa by vicariance events that were, in turn, caused by multiple interruptions of Beringia Land Bridges or cooling of climate and seem to be consistent with the times of multiple disjunctions of the flora.     

Pollination by fungus gnats (Diptera: Mycetophilidae) and self-recognition sites in Tolmiea menziesii (Saxifragaceae)

Coleeae (Bignoniaceae) are a tribe almost entirely restricted to Madagascar. Coleeae have previously been placed in neotropical Crescentieae due to species with indehiscent fruits, a character otherwise unusual in Bignoniaceae. A phylogeny based on three chloroplast regions (ndhF, trnT-L spacer, trnL-F spacer) identifies a monophyletic Coleeae that is endemic to Madagascar and surrounding islands of the Indian Ocean (Seychelles, Comores and Mascarenes). African Kigelia is not a member of Coleeae, rather it is more closely related to a subset of African and Southeast Asian species of Tecomeae. The molecular phylogeny indicates that indehiscent fruit have arisen repeatedly in Bignoniaceae: in Coleeae, Kigelia and Crescentieae. The characteristic fleshy fruits of species of Coleeae likely arose autochthonously in Madagascar. Within Coleeae Colea and Ophiocolea are sisters, Phyllarthron is sister to Colea + Ophiocolea, and Rhodocolea is sister to the rest of the tribe.     

Characteristics of the sperm flagellum in fungus gnats (Insecta,Diptera, Mycetophiloidea)

Spermatozoa from several members of the closely related Mycetophilidae and Keroplatidae were examined by electron microscopy using a fixative that contains glutaraldehyde and tannic acid, followed by a post-fixative that consists of uranyl acetate rather than osmium tetroxide. With this fixative, the detailed architecture of the flagellar axoneme and its various microtubules could be resolved. The so-called accessory tubules, which surround the central 9+2 unit of the sperm axoneme, were found to have 16 protofilaments in several examined Mycetophiloidea, but in no other Diptera. As 16 is the common number in holometabolic insects, it is presumably the plesiomorphic condition in Diptera. Other fungus gnats have accessory tubules with 15 or 14 protofilaments. The intertubular material situated between the accessory tubules is smaller in the examined members of the Mycetophilidae than in the Keroplatidae. The acrosome consists of an apical vesicle, which in one species, Macrorhyncha ancae, has three microtubular doublets in its anterior part and two large and three small extensions which extend posteriorly along the sperm axoneme.     

Studies of some fungus gnats (Diptera: Mycetophilidae) including nine additions to the British list

Abstract A key is provided to the eight species of Anatella recorded in Britain, including three species newly recorded ( A.dampfi Landrock; A.gibba Winnertz; A.lenis Dziedzicki). A further six species are newly recorded from Britain ( Exechia sororcula Lackschewitz; Exechiopsis dumitrescae (Burghele-Balacesco); Allodia ( Brachycampta ) angulata Lundström; Mycetophila autumnalis Lundström; M.lubomirskii Dziedzicki; M.lunata Meigen); these and several other species are redescribed. The following nomenclatural changes are proposed. Exechia bicincta (Staeger) (= spinosa Bukowski, syn.n.); Mycetophila confusa Dziedzicki (= affluctata Edwards, syn.n.); Mycetophila dziedzickii nom.n. (= obscura Dziedzicki nec Walker). Female characters of seventeen of the twenty-two British species of Exechia and eight species of Mycetophila are discussed and illustrations provided.     

Molecular phylogeny of the fungus gnat family Diadocidiidae and its position within the infraorder Bibionomorpha (Diptera)

The molecular phylogeny of the family Diadocidiidae (Diptera: Sciaroidea) is reconstructed based on the combined analysis of four mitochondrial (12S, 16S, COI, cytB) and two nuclear (28S, ITS2) gene markers. All the analyses strongly support Diadocidiidae as a monophyletic group. Genus Diadocidia Ruthe, 1831 includes monophyletic subgenera Diadocidia s. str. and Taidocidia Papp and ?ev?ík (Acta Zoologica Academiae Scientiarum Hungaricae, 51 , 2005b, 329). The monophyly of Adidocidia La?tovka & Matile, 1972 was not confirmed. The position of Diadocidiidae and relationships of the families within the infraorder Bibionomorpha are demonstrated in the analyses based on three gene markers (28S, 12S and 16S). The Bayesian and maximum likelihood analyses of 10 families of Bibionomorpha revealed Sciaridae as the closest relative of Diadocidiidae. Most of the currently recognised extant families of Bibionomorpha proved to be monophyletic. The family Keroplatidae revealed as paraphyletic, with the genera of Macrocerinae being more related to Cecidomyiidae, but the support is low.     

Molecular phylogeny of black flies (Diptera: Simuliidae) from Thailand, using ITS2 rDNA

The sequences of the second internal transcribed spacer (ITS2) of ribosomal DNA (rDNA) were determined for 40 black fly species from Thailand, belonging to 4 subgenera of the genus Simulium, namely Gomphostilbia (12 species), Nevermannia (5 species), Montisimulium (1 species), Simulium sensu stricto (21 species), and an unknown subgenus with one species (Simulium baimaii). The length of the ITS2 ranged from 247 to 308 bp. All black fly species had high AT content, ranging from 71 to 83.8%. Intraindividual variation (clonal variation) occurred in 13 species, ranging from 0.3 to 1.1%. Large intrapopulation and interpopulation heterogeneities exist in S. feuerboni from the same and different locations in Doi Inthanon National Park, northern Thailand. Phylogenetic relationships among 40 black fly species were examined using PAUP (version 4.0b10) and MrBAYS (version 3.0B4). The topology of the trees revealed two major monophyletic clades. The subgenus Simulium and Simulium baimaii were placed in the first monophyletic clade, whereas the subgenera Nevermannia + Montisimulium were placed as the sister group to the subgenus Gomphostilbia in the second monophyletic clade. Our results suggest that S. baimaii belongs to the malyschevi-group or variegatum-group in the subgenus Simulium. The molecular phylogeny generally agrees with existing morphology-based phylogenies.     

有瓣蝇类分类、系统发育及演化

有瓣蝇类(Calyptratae)隶属于昆虫纲(Insecta)四大超适应辐射类群之一的双翅目(Diptera),占双翅目已知物种多样性的近20%。有瓣蝇类分布广泛,生物学习性极为多样,在维系生态系统稳定中发挥着重要作用,是媒介、法医、传粉和天敌昆虫学研究领域的热点类群,也是探究双翅目系统演化及其成功适应辐射的关键类群。为了还原有瓣蝇类的演化历史,许多著名昆虫学者先后对该类昆虫开展过不同层面的研究。有瓣蝇类的单系性得到了普遍支持,并被分为3个总科——虱蝇总科(Hippoboscoidea)、蝇总科(Muscoidea)和狂蝇总科(Oestroidea),其中单系的狂蝇总科与多系的蝇总科聚为一支,再与虱蝇总科成为姐妹群。在科级阶元水平,蝠蝇科(Streblidae)(虱蝇总科)、花蝇科(Anthomyiidae)(蝇总科)、丽蝇科(Calliphoridae)(狂蝇总科)、邻寄蝇科(Rhinophoridae)(狂蝇总科)等类群的单系性仍有待验证,且新的科仍在不断被建立［如粉蝇科(Polleniidae)、乌鲁鲁蝇科(Ulurumyiidae)］,因此,有瓣蝇类科级系统发育关系仍不十分明晰。已有研究对虱蝇总科虱蝇科(Hippoboscidae)、蝠蝇科、蛛蝇科(Nycteribiidae),蝇总科蝇科(Muscidae)、粪蝇科(Scathophagidae),狂蝇总科麻蝇科(Sarcophagidae)、狂蝇科(Oestridae)胃蝇亚科(Gasterophilinae)的演化历史进行研究,明确了起源与扩散、寄主转移、取食策略等关键生物学习性的演化历史。但由于部分关键类群生活史信息的缺失,以及尚未有效解决的系统发育关系,有瓣蝇类演化历史仍有许多待解之谜。本文综述了有瓣蝇类分类、系统发育及演化研究进展,是在系统学研究进入系统发育基因组学时代后对该类群相关研究进展的首次全面总结。     

Field and laboratory observations on the substrates of the mushroom fungus gnat Lycoriella auripila (Diptera: Sciaridae)

The movements of Psila rosae larvae were studied under field and laboratory conditions. In the field, larvae moved up to 60 cm along and between carrot rows, although more larvae moved during the first generation than the second. Sudden increases in soil moisture, and not its prevailing level, stimulated the larvae to move from carrots into the soil surrounding the roots. Up to 50% of the larvae that left their mines in carrots following a sudden increase in soil moisture did not return to the same mine. Evacuation of mines by larvae appears to be a reflex response to the flooding of their mine and might be connected with the production of toxic conditions within the mine following flooding.     

Quantitative sampling method for fungus gnat (Diptera: Sciaridae) eggs in soilless growing media

A simple method is described for separating fungus gnat eggs from soilless growing media. The flotation/extraction method primarily involves the use of a MgSO4 solution (density 1.065 g cm(-3)) and a series of sieving procedures. Bradysia sp. nr. coprophila Lintner eggs were collected in an isolation chamber containing adult fungus gnats. Three soilless growing media were used: Metro-Mix 560 with Scott's Coir, Sunshine LC1 Mix, and Universal SB 300 Mix. Each growing medium was inoculated with 100 fungus gnat eggs. Growing medium samples were processed using separatory funnels and a MgSO4 solution. Eggs were washed through a 250-microm sieve and collected on a 53-microm sieve, and then the eggs were rinsed into glass petri dishes. The sieving process separated fungus gnat eggs from the larger growing medium particulates (perlite, bark, and peat), providing a sample consisting of water, eggs, and fine growing medium particulates (peat). Using a vortex for 60 s increased the number of fungus gnat eggs recovered (37.4 +/- 3.7-67.1 +/- 1.9 [mean +/- SEM]) for all three growing media. The mean number of fungus gnat eggs recovered from the three soilless growing media was significantly different, with the growing medium containing coir (Metro-Mix 560) having the highest recovery rate of 61.0 +/- 2.1. For all three growing media, there was > 50% recovery of fungus gnat eggs. In addition to fungus gnat eggs, this method also may prove useful for quantitative recovery of shore fly (Scatella sp.) eggs and the eggs of other arthropods from soilless growing media.     

Molecular phylogeny of the fungus gnat subfamilies Gnoristinae and Mycomyinae,and their position within Mycetophilidae (Diptera)

The phylogeny of the fungus gnat family Mycetophilidae (Diptera) is reconstructed with a focus on the species‐rich and taxonomically difficult subfamilies Gnoristinae and Mycomyinae. The multigene phylogenetic analyses are based on five nuclear (18S, 28S, CAD, MCS, ITS2) and four mitochondrial (12S, 16S, COI, CytB) gene markers. The analyses strongly support the monophyly of Mycetophilidae and the subfamilies Manotinae, Sciophilinae, Leiinae, and Mycomyinae, although Gnoristinae is paraphyletic with respect to Mycetophilinae. All the genera and groups of genera included are supported as monophyletic, except for Acomoptera Vockeroth, Boletina Staeger, Dziedzickia Johannsen, Ectrepesthoneura Enderlein, and Neoempheria Osten Sacken. Ancestral character state reconstructions were applied to two morphological features present in Gnoristinae and Mycomyinae (i.e. presence of setae on wing membrane and wing vein R₄) in order to assess their evolution. The wing vein R₄ appears as an unstable character, spread throughout different clades. A dated phylogeny of the family Mycetophilidae showed that most of the subfamilies of Mycetophilidae originated and diversified during the Cretaceous. The youngest subfamilies, originated in the Paleogene, appear to be Mycomyinae and Mycetophilinae.     

Ingestion of Pythium spp. by larvae of the fungus gnat Bradysia impatiens (Diptera:Sciaridae)

Mycelium, oospores and zoospore cysts of Pythium spp. were fed to larvae of the fungus gnat Bradysia impatiens. The fungal structures were all ingested and provided a complete nutritional source for the insect's development from egg to adult. Mycelium seemed the major source of food as only empty fragments were found either in the digestive tract or in larval faeces. Oospores appeared intact and were viable both during passage through the tract and when expelled. Germination of oospores was normal. Most encysted zoospores also survived passage through the gut, although some appeared damaged. After transferring recently-fed larvae to a new food source, oospores were still detected in the digestive tract 48 h later. These results show that the larval stage of B. impatiens may serve as an important vector for Pythium spp.