Systematics and evolution of predatory flower flies (Diptera: Syrphidae) based on exon-capture sequencing

Flower flies (Diptera: Syrphidae) are one of the most species-rich dipteran families and provide important ecosystem services such as pollination, biological control of pests, recycling of organic matter and redistributions of essential nutrients. Flower fly adults generally feed on pollen and nectar, but their larval feeding habits are strikingly diverse. In the present study, high-throughput sequencing was used to capture and enrich phylogenetically and evolutionary informative exonic regions. With the help of the baitfisher software, we developed a new bait kit (SYRPHIDAE1.0) to target 1945 CDS regions belonging to 1312 orthologous genes. This new bait kit was successfully used to exon capture the targeted loci in 121 flower fly species across the different subfamilies of Syrphidae. We analysed different amino acid and nucleotide data sets (1302 loci and 154 loci) with maximum likelihood and multispecies coalescent models. Our analyses yielded highly supported similar topologies, although the degree of the SRH (global stationarity, reversibility and homogeneity) conditions varied greatly between amino acid and nucleotide data sets. The sisterhood of subfamilies Pipizinae and Syrphinae is supported in all our analyses, confirming a common origin of taxa feeding on soft-bodied arthropods. Based on our results, we define Syrphini stat.rev. to include the genera Toxomerus and Paragus. Our divergence estimate analyses with beast inferred the origin of the Syrphidae in the Lower Cretaceous (125.5–98.5 Ma) and the diversification of predatory flower flies around the K–Pg boundary (70.61–54.4 Ma), coinciding with the rise and diversification of their prey.     

Results of the global conservation assessment of the freshwater crabs (Brachyura,Pseudothelphusidae and Trichodactylidae): The Neotropical region,with an update on diversity

The freshwater crabs of the Neotropics comprise 311 species in two families (Pseudothelphusidae and Trichodactylidae) and one or both of these families are found in all of the countries in the Neotropical region (except for Chile and some of the Caribbean islands). Colombia (102 species, 81% endemic) and Mexico (67 species, 95% endemic) are the biodiversity hotspots of freshwater crab species richness and country-level endemism for this region. The results of the IUCN Red List conservation assessments show that 34% of pseudothelphusids and 10% of trichodactylids have an elevated risk of extinction, 29% of pseudothelphusids and 75% of trichodactylids are not at-risk (Least Concern), and although none are actually extinct, 56% of pseudothelphusids and 17% of trichodactylids are too poorly known to assess (Data Deficient). Colombia (14 species), Venezuela (7 species), Mexico (6 species), and Ecuador (5 species) are the countries with the highest number of threatened species of Neotropical freshwater crabs. The majority of threatened species are restricted-range semiterrestrial endemics living in habitats subjected to deforestation, alteration of drainage patterns, and pollution. This underlines the need to prioritize and develop conservation measures before species decline to levels from which they cannot recover. These results represent a baseline that can be used to design strategies to save threatened Neotropical species of freshwater crabs.     

Hoverflies (Diptera,Syrphidae) in Agrolandscapes of St. Petersburg and Leningrad Province

A total of 57 species from 32 genera of hoverflies (Diptera, Syrphidae) were found in the agricultural landscapes of St. Petersburg and Leningrad Province. One species, Cheilosia vulpina (Mg.), was recorded in the region for the first time. The largest numbers of species occurred at forest and field edges covered with wild grasses and shrubs, while 21 species of hoverflies were recorded in crops. The species most commonly found in crops were representatives of the genera Melanostoma, Sphaerophoria, and Platycheirus whose larvae feed on aphids.     

Cover Caption

An adult flower fly (Syrphidae) rests on a flower of Erigeron annuus (Asteraceae) growing on a rice farm in eastern China. Plants of non‐crop species can be important in providing nectar and pollen to natural enemies of crop pests. Whilst the larvae of flower flies prey on pests such as aphids, the adults are non‐predacious and females need pollen in order to produce eggs. See pages 1–12. Photo by Ping‐Yang Zhu.     

Reproductive biology of Epidendrum tridactylum (Orchidaceae: Epidendroideae): a reward-producing species and its deceptive flowers

Epidendrum L. (Epidendroideae, Laeliinae) is one of the largest orchid genera distributed throughout tropical America. Based on data on reproductive phenology, floral morphology, labellar histochemistry, pollinators, pollination mechanisms and breeding system, this study explored the reproductive biology of Epidendrum tridactylum Lindl. growing in the state of São Paulo, southeastern Brazil. Epidendrum tridactylum produces an average of 87 rewardless flowers with a longitudinal protuberance on the median portion of its labellum, whose epidermal cells produce a pleasant citric fragrance. This fragrance attracts both male and female flies of families Calliphoridae, Sarcophagidae and Syrphidae that end up drinking the extrafloral nectar produced at the base of the bracts. Pollinaria are removed when the fly proboscis contacts the viscidium in search of nectar. After removing the pollinarium, the fly gets trapped in the flower, which frightens it, thus inhibiting any possible intent to immediately visit another flower and, at least presumably, reducing the possibility of geitonogamy. Since E. tridactylum is self-incompatible, the positive consequence of such behavior is a reduction of pollen loss. The fruits obtained through experimental cross-pollinations present highly variable percentages of potentially viable seeds. This is the first comprehensive study on the reproductive biology of a myophilous Epidendrum and a pioneer report of effective pollinators picking extrafloral nectar from Orchidaceae.     

A morphological and phylogenetic analysis of <Emphasis Type="Italic">Ornithodoros marinkellei</Emphasis> (Acari: Argasidae), with additional notes on habitat and host usage

Ornithodoros marinkellei was described from larvae collected on Pteronotus spp. bats in Colombia and Panama. More recently, this tick was reported in the Brazilian Amazon. Because some morphometric differences were observed between O. marinkellei larvae from Colombia and Brazil, it was proposed that further investigations were needed to assess whether the differences could be attributed to intra- or inter-specific polymorphism. Herein, we collected O. marinkellei specimens in the type locality of Colombia, in Brazil, and in a new locality in Nicaragua, expanding the distribution of the species to Nicaragua. Morphometric analysis of larvae and adults, corroborated by a principal component analysis (PCA), indicated that the Brazilian specimens were larger than specimens from Colombia and Nicaragua. Phylogenetic analysis inferred from the mitochondrial 16S rRNA gene showed ticks from Colombia and Nicaragua more genetically related than any of them with ticks from Brazil, although ticks from the three countries grouped in a clade sister to a major clade containing sequences of various Neotropical Ornithodoros species. We concluded that ticks identified as O. marinkellei from Colombia, Nicaragua, and Brazil represent the same taxon, and that the genetic and morphological differences between them are likely to have a geographical bias. We redescribed the nymph of O. marinkellei, which has a vestigial hypostome, probably incompatible with blood feeding. We also report human infestation by O. marinkellei adults. As all reports of O. marinkellei adults have been from hot caves (temperature?>?35 °C), this abiotic condition could be a limiting factor for the occurrence of this tick species.     

The value of insect pollination to yield of oilseed rape (Brassica rapa) in Bangladesh

Brassica rapa L., is a crop grown globally and studies have indicated that insect pollination can improve yields. However, the importance of insect pollination in this crop depends on cultivar reproductive biology, insect pollinator species and their abundances. In Bangladesh, the acreage of B. rapa for oilseed production has been expanding, but little is known about whether insects contribute to yield improvements. Using the commonly grown variety Tori-7, we found that plants with inflorescences left exposed to flower visiting insects (body width > 1.5 mm) had a 30.8% greater seed yield compared to those where insect flower visitors were excluded. Of 794 insect flower visiting individuals recorded from observation and trapping surveys conducted across four separate fields, Apis bees (four spp.) were the most abundant (66.1%), followed by flies Musca domestica (14.7%), Sepsis fulgens (6.3%) and hoverflies (3.0%; 3 spp; Syrphidae). Other flower visitors included ants, wasps, beetles, butterflies and moths. For the cultivar assessed, we calculated the economic value of oilseed rape was $US 87.5 million per annum in Bangladesh, of which the economic value of insect pollination was $US 26.92 million per annum. Thus, one in every four dollars earned by our Bangladeshi growers resulted from insect crop pollination. This demonstrates to growers the need to promote and protect insect pollination to optimize their economic returns.     

Ithomiini Butterflies (Lepidoptera: Nymphalidae) of Antioquia, Colombia

Colombia is one of the most biodiverse countries on the planet. However, economic and scientific investment in completing inventories of its biodiversity has been relatively poor in comparison with other Neotropical countries. Butterflies are the best studied group of invertebrates, with the highest proportion of known to expected species. More than 3,200 species of butterflies have been recorded in Colombia, although the study of the still many unexplored areas will presumably increase this number. This work provides a list of Ithomiini butterflies collected in the department of Antioquia and estimates the total number of species present, based on revision of entomological collections, records in the literature and field work performed between 2003 and 2011. The list includes 99 species and 32 genera, representing 27% of all Ithomiini species. We report 50 species of Ithomiini not formerly listed from Antioquia, and found the highest diversity of ithomiine species to be at middle elevations (900–1,800 m). The mean value of the Chao2 estimator for number of species in Antioquia is 115 species, which is close to a predicted total of 109 based on known distributions of other Ithomiini not yet recorded from the department. Nine species are potentially of particular conservation importance because of their restricted distributions, and we present range maps for each species. We also highlight areas in Antioquia with a lack of biodiversity knowledge to be targeted in future studies. This paper contributes to mapping the distribution of the Lepidoptera of Antioquia department in particular and of Colombia in general.     

Comparative morphology of immature stages of four species of Chinavia (Hemiptera,Pentatomidae), with a key to the species of Rio Grande do Sul,Brazil

Chinavia Orian (1965) is one of the most diverse genera of Pentatomidae, distributed in the Afrotropical, Neotropical and Nearctic Regions. Thirty-two species are recorded for Brazil, some of them having potential economic impact because they are found on crops and referred to as pests. The morphology of the five nymphal instars of Chinavia armigera (Stål, 1859), Chinavia aseada (Rolston, 1983), Chinavia brasicola (Rolston, 1983) and Chinavia runaspis (Dallas, 1851) are described here. Through a comparative study, identification keys were developed to allow an early identification of the 12 Chinavia species of Rio Grande do Sul.     

Review of the fauna of the beetle families Scydmaenidae and Pselaphidae (Coleoptera, Staphylinoidea) of Chuvashia

A review of the coleopteran families Scydmaenidae (18 species) and Pselaphidae (23 species) of the Chuvash Republic is presented based on examination of the material collected in 1975?C2011. 18 species and 7 genera of Scydmaenidae and 21 species of 13 genera of Pselaphidae are recorded from this territory for the first time. Euplectus bescidicus Reitt. (Pselaphidae) is for the first time recorded from Russia.     

Ectomycorrhizal fungal sporocarp diversity and discovery of new taxa in Dicymbe monodominant forests of the Guiana Shield

Ectomycorrhizal (ECM) fungi historically were considered poorly represented in Neotropical forests but in the central Guiana Shield substantial areas are dominated by leguminous ECM trees. In the Upper Potaro Basin of Western Guyana, ECM fungi were sampled for 7?years during the rainy seasons of 2000–2008 in three 1-ha plots in primary monodominant forests of the ECM canopy tree Dicymbe corymbosa (Fabaceae subfam. Caesalpinioideae). Over the plot sampling period sporocarps of 126 species of putative or confirmed ECM fungi were recovered. These taxa represented 13 families and 25 genera of primarily Agaricomycetes, but also Ascomycota (Elaphomycetaceae), the majority of which are new to science. Russulaceae contained the most species (20 Russula; 9 Lactarius), followed by Boletaceae (8 genera, 25 spp.), Clavulinaceae (17 Clavulina), and Amanitaceae (16 Amanita). An additional 46 species of ECM fungi were collected in forests of the Upper Potaro Basin outside the study plots between 2000 and 2010, bringing the regional number of ECM species known from sporocarps to 172. This is the first long-term ECM macrofungal dataset from an ECM-dominated Neotropical forest, and sporocarp diversity is comparable to that recorded for ECM-diverse temperate and boreal forests. While a species accumulation curve indicated that ECM sporocarp diversity was not fully recovered inside of the plots,?~80% of the total species were recovered in the first year. Sequence data from ECM roots have confirmed the ECM status of 56 taxa represented by corresponding sporocarp data. However,?>50% of ECM fungal species from roots remain undiscovered as sporocarps, leading to a conservative estimate of?>?250 ECM species at the Potaro site. Dicymbe forests in Guyana are a hotspot for ECM fungal diversity in the Neotropics.     

Annotated catalog,biological notes and diagnoses of the flower bugs (Heteroptera: Anthocoridae sensu lato) in the Korean Peninsula

The flower bugs or minute pirate bugs, Anthocoridae sensu lato, are represented by 12 genera and 24 species in the Korean Peninsula. The present catalog includes three newly recorded species to the fauna: Anthocoris confusus Reuter, Bilia japonica Carayon et Miyamoto, and Montandoniola pictipennis (Esaki). Distributional data and bibliographical references of each taxon are included. Biological notes (habitats, prey types, etc.) and diagnoses based on the local materials of the Korean Peninsula are also presented for the field of applied entomology.     

Pterandra pyroidea: a case of pollination shift within neotropical Malpighiaceae

Background and Aims

Most Neotropical species of Malpighiaceae produce floral fatty oils in calyx glands to attract pollinating oil-collecting bees, which depend on this resource for reproduction. This specialized type of pollination system tends to be lost in members of the family that occur outside the geographic distribution (e.g. Africa) of Neotropical oil-collecting bees. This study focused on the pollination ecology, chemical ecology and reproductive biology of an oil flower species, Pterandra pyroidea (Malpighiaceae) from the Brazilian Cerrado. Populations of this species consist of plants with oil-secreting (glandular) flowers, plants with non-oil-secreting flowers (eglandular) or a mix of both plant types. This study specifically aims to clarify the role of eglandular morphs in this species.

Methods

Data on pollinators were recorded by in situ observations. Breeding system experiments were conducted by isolating inflorescences and by enzymatic reactions. Floral resources, pollen and floral oils offered by this species were analysed by staining and a combination of various spectroscopic methods.

Key Results

Eglandular flowers of P. pyroidea do not act as mimics of their oil-producing conspecifics to attract pollinators. Instead, both oil-producing and oil-free flowers depend on pollen-collecting bees for reproduction, and their main pollinators are bumble-bees. Floral oils produced by glandular flowers are less complex than those described in closely related genera.

Conclusions

Eglandular flowers represent a shift in the pollination system in which oil is being lost and pollen is becoming the main reward of P. pyroidea flowers. Pollination shifts of this kind have hitherto not been demonstrated empirically within Neotropical Malpighiaceae and this species exhibits an unusual transition from a specialized towards a generalized pollination system in an area considered the hotspot of oil-collecting bee diversity in the Neotropics. Transitions of this type provide an opportunity to study ongoing evolutionary mechanisms that promote the persistence of species previously involved in specialized mutualistic relationships.     

Comparison of the pollen content on the body and in the gut of hoverflies (Diptera,Syrphidae)

We analyzed the pollen content in the gut and on the body of hoverflies (Diptera: Syrphidae), mostly from three genera (Eristalis, Syrphus s. l., and Sphaerophoria) that visited flowers of two jointly flowering species of Apiaceae with indistinguishable pollen (Anthriscus sylvestris and Aegopodium podagraria) and of Bunias orientalis (Brassicaceae). Pollen compositions depended on the plant from which the flies were collected rather than on the insect genus. At the same time, individual preferences of the studied flies to food plant species were confirmed. The pollen compositions in the gut and on the body were weakly correlated. Two possible reasons for such a weak correlation are discussed: individual differences in the fly feeding habits and different time of pollen preservation in the gut and on the body.     

Pugs of the tribe Eupitheciini (Lepidoptera,Geometridae): the history of study and distribution over biogeographic regions

The World fauna of the tribe Eupitheciini is the most species-rich in the family Geometridae. This tribe includes about 1900 species (almost 3000 species-group names) from 47 genera; about one third of the genera (15) are monotypic. The generic diversity of Eupitheciini is the highest in the Australian (38 genera, 11 of them endemic) and Oriental regions (32 genera, 4 endemic) and the lowest in the Neotropical Region (possibly one genus only). The faunas of different biogeographic regions can be arranged in following order by their species richness: the Palaearctic (487 species), Oriental (397), Neotropical (346), Australian (251), Afrotropical (198), and Nearctic Regions (166 species). Eupithecia is the most species-rich genus in the family Geometridae and the entire order Lepidoptera, and one of the largest genera in the whole World fauna of insects. The greatest number of species of this genus is recorded in the Palaearctic Region (466 species), where Eupithecia accounts for about 95% of the tribe Eupitheciini. The mainland of the Oriental Region (especially the Himalayas) is also very species-rich; however the proportion of the Eupithecia representatives decreases towards Malaysia, Sundaland, and the Australian Region (about 2% of the tribe). The Eupitheciini faunas have the greatest similarity at the generic level between the Oriental and Australian Regions (the Jaccard and Sørensen coefficient values being 0.62 and 0.77, respectively). The Palaearctic fauna is more similar to the Afrotropical and Oriental faunas at the genus-group level. On the whole, the fauna of the Nearctic Region is similar to that the West Palaearctic, with the exception of the fact that representatives of the genera Gymnoscelis and Chloroclystis are absent in North America, although two endemic genera Nasusina and Prorella are present. At the genus-group level, the Nearctic fauna of Eupitheciini is more similar to the Neotropical (the Jaccard and Sørensen coefficients 0.20 and 0.33, respectively) than to the Palaearctic fauna (0.17 and 0.29). The number of synonymies is very high in the tribe Eupitheciini because of the homogeneity of this group, whose species are difficult to identify without the use of elaborate anatomical techniques. Modern revisions, catalogues, surveys, and atlases on Eupitheciini are absent for many countries and large geographic regions. Revisions of pugs of the tribe Eupitheciini for some biogeographic regions are extremely difficult because of fragmentation of entomological collections including the type specimens of many species-group taxa. A large fraction of synonyms is characteristic of parts of the World with the best known faunas: Europe (64% of synonyms) and North America (39%). On the contrary, the lowest levels of synonymy are typical of the less known faunas of the regions situated at the equatorial latitudes, namely the Neotropical (9%) and Afrotropical (8%) ones.     

Almeidea A. St.-Hil. Belongs to Conchocarpus J.C. Mikan (Galipeinae,Rutaceae): Evidence from Morphological and Molecular Data,with a First Analysis of Subtribe Galipeinae

Subtribe Galipeinae (tribe Galipeeae, subfamily Rutoideae) is the most diverse group of Neotropical Rutaceae, with 28 genera and approximately 130 species. One of its genera is Almeidea, whose species are morphologically similar to those of the genus Conchocarpus. Species of Almeidea occur in the Atlantic Rain Forest of Eastern Brazil, with one species (Almeidea rubra) also present in Bolivia. The objective of this study was to perform a phylogenetic analysis of Almeidea, using a broader sampling of Galipeinae and other Neotropical Rutaceae, the first such study focused on this subtribe. To achieve this objective, morphological data and molecular data from the nuclear markers ITS-1 and ITS-2 and the plastid markers trnL-trnF and rps16 were obtained. Representatives of eight genera of Galipeinae and three genera of Pilocarpinae (included also in Galipeeae) and Hortia (closely related to Galipeeae) were used. Five species of Almeidea and seven of Conchocarpus were included, given the morphological proximity between these two genera. Individual (for each molecular marker) and combined phylogenetic analyses were made, using parsimony and Bayesian inference as optimization criteria. Results showed Galipeinae as monophyletic, with the species of Almeidea also monophyletic (supported by the presence of pantocolporate pollen) and nested in a clade with a group of species of Conchocarpus, a non-monophyletic group. Additionally, C. concinnus appeared in a group with Andreadoxa, Erythrochiton, and Neoraputia, other members of Galipeinae. As a result, Conchocarpus would be monophyletic only with the exclusion of a group of species related to C. concinnus and with the inclusion of all species of Almeidea with the group of species of Conchocarpus that includes its type species, C. macrophyllus. Thus, species of Almeidea are transferred to Conchocarpus, and the new combinations are made here.     

Weberiella De Carlo, 1966 (Insecta: Heteroptera: Belostomatidae) revisited: Redescription with a key to the genera of Belostomatidae and considerations on back-brooding behaviour

Males of some subgroups of Belostomatidae brood the eggs attached to their backs, whereas the eggs are attached to the vegetation by females in others. Male brood care is obligatory in the belostomatine species of Abedus Stål, 1862, Belostoma Latreille, 1807, Diplonychus Laporte, 1833, Hydrocyrius Spinola, 1850, and Limnogeton Mayr, 1853. Recent investigations into relationships among Neotropical Belostomatinae have led authors to recognize a clade Belostomatinae, which is mainly characterized by back-brooding behaviour. It is likely that Weberiella belongs to this clade. Males of the only described species W. rhomboides (Menke, 1965) are reported as carrying eggs on their back for the first time here. Since this species was described based on a single female specimen from French Guiana, W. rhomboides is redescribed based on specimens from Amazonas, Mato Grosso, Rondônia, and Roraima States, Brazil. Its distribution in Brazil is mapped. The scale-like abdomen is a presumably autapomorphic condition not found in the other belostomatid species. A key to the genera of Belostomatidae including Lethocerinae and Horvathiniinae is provided. Even though a formal parsimony analysis is not presented here, a placement of W. rhomboides in Belostomatinae is tentatively suggested based on the back-brooding behaviour of males as a shared apomorphy. Convergent evolution cannot be excluded as suitable vegetation for depositing eggs is not available in the specific habitat (kinon) of W. rhomboides.     

The systematic value of flower structure in Crotalaria and related genera of the tribe Crotalarieae (Fabaceae)

Flowers in the tribe Crotalarieae of the family Fabaceae are generally adapted to bee pollination mechanisms. Molecular systematics have recently provided a major step towards a profound insight into generic relationships, thereby creating the opportunity to re-evaluate the taxonomic and functional significance of flower structure in the tribe, with emphasis on the large genus Crotalaria. A representative sample of flowers from 211 species was dissected to record morphological character states. These data were supplemented from the literature to allow for generalizations for the tribe as a whole. Six structural–functional flower types were identified: (1) pump; (2) gullet; (3) hugging; (4) saddle; (5) tunnel and (6) brush (saddle and tunnel types described here for the first time). Crotalaria uniquely has the brush type, characterized by a rostrate keel, highly dimorphic anthers, stylar trichomes and elaborate callosities on the standard petal. Remarkably, Crotalaria and Bolusia are the only genera of the tribe Crotalarieae with callosities present in all of the species. In other genera, callosities are generally absent or infrequent. Trends towards specialization of pollination syndromes are apparent as assemblages of apomorphic states that co-occur in what we refer to here as “specialized flowers”; individual characters are labile or non-homologous (e.g. callosities) and diagnostically less valuable. Unique combinations of flower characters are often useful to support current generic concepts in Crotalarieae.