Jostein Kjærandsen

Stable structural color patterns displayed on transparent insect wings

Color patterns play central roles in the behavior of insects, and are important traits for taxonomic studies. Here we report striking and stable structural color patterns—wing interference patterns (WIPs)—in the transparent wings of small Hymenoptera and Diptera, patterns that have been largely overlooked by biologists. These extremely thin wings reflect vivid color patterns caused by thin film interference. The visibility of these patterns is affected by the way the insects display their wings against various backgrounds with different light properties. The specific color sequence displayed lacks pure red and matches the color vision of most insects, strongly suggesting that the biological significance of WIPs lies in visual signaling. Taxon-specific color patterns are formed by uneven membrane thickness, pigmentation, venation, and hair placement. The optically refracted pattern is also stabilized by microstructures of the wing such as membrane corrugations and spherical cell structures that reinforce the pattern and make it essentially noniridescent over a large range of light incidences. WIPs can be applied to map the micromorphology of wings through direct observation and are useful in several fields of biology. We demonstrate their usefulness as identification patterns to solve cases of cryptic species complexes in tiny parasitic wasps, and indicate their potentials for research on the genetic control of wing development through direct links between the transregulatory wing landscape and interference patterns we observe in Drosophila model species. Some species display sexually dimorphic WIPs, suggesting sexual selection as one of the driving forces for their evolution.

Sexual selection on wing interference patterns in Drosophila melanogaster.

Significance Recently, it was discovered that small insects like flies and wasps, with seemingly transparent wings, display vivid coloration against dark backgrounds because of so-called “wing interference patterns” (WIPs). It was proposed that such wing coloration could function in sexual selection and species recognition, but direct evidence for this hypothesis has been lacking. Here, to our knowledge we present the first experimental evidence that WIPs in male Drosophila melanogaster are targets of mate choice from females. Comparison of attractiveness of different male WIPs between black and white backgrounds revealed that two different color traits, saturation and hue, experience directional and stabilizing sexual selection, respectively. Our results suggest that vivid coloration in WIPs is a target of mate choice and might have evolved by sexual selection. Animals with color vision use color information in intra- and interspecific communication, which in turn may drive the evolution of conspicuous colored body traits via natural and sexual selection. A recent study found that the transparent wings of small flies and wasps in lower-reflectance light environments display vivid and stable structural color patterns, called “wing interference patterns” (WIPs). Such WIPs were hypothesized to function in sexual selection among small insects with wing displays, but this has not been experimentally verified. Here, to our knowledge we present the first experimental evidence that WIPs in males of Drosophila melanogaster are targets of mate choice from females, and that two different color traits—saturation and hue—experience directional and stabilizing sexual selection, respectively. Using isogenic lines from the D. melanogaster Genetic Reference Panel, we compare attractiveness of different male WIPs against black and white visual backgrounds. We show that males with more vivid wings are more attractive to females than are males with dull wings. Wings with a large magenta area (i.e., intermediate trait values) were also preferred over those with a large blue or yellow area. These experimental results add a visual element to the Drosophila mating array, integrating sexual selection with elements of genetics and evo-devo, potentially applicable to a wide array of small insects with hyaline wings. Our results further underscore that the mode of sexual selection on such visual signals can differ profoundly between different color components, in this case hue and saturation.

Review of fungus gnats in the genus Tarnania Tuomikoksi, with a phylogeny of the Rymosia s.l. genus group (Diptera : Mycetophilidae)

The four known species of Tarnania Tuomikoski are reviewed, viz. T dziedzickii (Edwards, 1941), T. fenestralis (Meigen, 1818), T. nemoralis (Edwards, 1941), and T. tarnanii (Dziedzicki, 1910). Males and females are re-described, photographed and their terminalia figured. An attempt is made to homologize and name specific substructures of the male gonostylus, widely recognizable within the tribe Exechiini. A phylogenetic analysis of the Tarnania species, including representatives of the six other genera of the Rymosia s. lat. genus group, strengthens the view that Tarnania is monophyletic. Tarnania is supported by six unambiguous character changes of which three are considered to be unique synapomorphies, viz. 1) lower parts of metepisternum and metepimeron covered with campaniform sensilla; 2) hind tibia with several curved bristles posterodorsally, placed irregularly in more than one row; and 3) anterior branch of male gonostylus forming an elongated, bifurcated lobe. The most parsimonious hypothesis for relationships among the species of Tarnania is ((T tarnanii + T nemoralis) + (T dziedzickii + T fenestralis)). The practice of regarding the genera of the Rymosia s. lat. genus group as an assemblage of plesiomorphic genera is also supported, the most parsimonious hypothesis for relationships among them being (Notolopha (Allodiopsis (Myrosia (Synplasta (Tarnania + (Rymosia + Pseudorymosia)))))). Tarnania is most diverse in northwestern Europe, with all four species known from Norway, Sweden, Great Britain, Belgium, Germany, Switzerland, and France. T dziedzickii extends its distribution southwards to include the North African and the Near East Regions. T. fenestralis is widely distributed in the western Palaearctic Region and extends its distribution eastwards to the eastern Palaearctic region. Only T. tarnanii displays a wide Holarctic distribution including most of Russia, Alaska, Canada and Greenland. (Less)

Towards a molecular phylogeny of the fungus gnat genus Boletina (Diptera: Mycetophilidae)

Martinsson, S., Kjærandsen, J. & Sundberg, P. (2011). Towards a molecular phylogeny of the fungus gnat genus Boletina (Diptera: Mycetophilidae). —Zoologica Scripta, 40, 272–281.

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.

Species assemblages and community structure of adult caddisflies along a headwater stream in southeastern Ghana (Insecta: Trichoptera)

Adult caddisflies were collected at 12 sites along a 5 km stretch of a forested headwater stream in southeastern Ghana in March and November 1993 and the catches are related to environmental variables. Some 34,000 specimens, belonging to 178 morphospecies in 43 genera and 11 families, were caught with Malaise traps and light traps. Many species demonstrate differences between sampling method, seasonal occurrence, and sex-ratios, with the light trap in the second wet sunny season (November) being most efficient with respect to the diversity of the catches. A Moran index analysis for global structures indicates a major transition in the fauna from the ravine with waterfalls to the forest, and secondly, a gradient through the forest. A Geary index analysis for local structures indicates further faunal turnovers in the lower reaches of the stream. Species-environment relationships were analyzed through correspondence analysis and co-inertia analysis, allowing ordination of both species and the environment into three zones. The first co-inertia axis reveals a transition between the waterfalls in the open ravine (sites 1–3), and the stream through the riverine forest (sites 4–12), while the second co-inertia axis reveals a gradient from the lotic stream in the moist semi-deciduous forest (sites 4–8) to more lentic stretches of the stream in a gradually more depleted forest (sites 9–12). The most important environmental factors related to the species transition are: riparian moss growth (r=0.94), leaves in the stream substratum (r=0.91), forest type (r=0.90), hygropetric surfaces (r=0.88), bedrock (r=0.87), lotic [riffle] (r=0.85) and lentic [pool] (r=0.85) stretches. The most important environmental factors related to the species gradient are: lotic [riffle] stretches (r=0.92), gravel (r=0.88) and leaves (r=0.85) in the stream substratum, forest type (r=0.81), and agricultural use (r=0.76). Three clusters representing three major environmental zones (zone I, II and III) along the stream are identified by projecting the average species positions on the co-inertia plane. By analyzing the Malaise trap samples significant indicator values are found for 29 species in zone I, 16 species in zone II, and 18 species in zone III. By analyzing the light trap samples significant indicator values are found for 17 species in zone I, 57 species in zone II, and 38 species in zone III. The high diversity of caddisflies in the sanctuary might be reflected both in the geological and climatological history of West Africa, as well as in the recognition of three major environmental zones, and the clean water with high diversity of stream microhabitats favorable for Trichoptera larvae. The closed forest seems to play an important role for the Trichoptera community, and the study demonstrates the importance of protecting forested headwater streams in order to maintain a sustainable aquatic biodiversity in tropical Africa.

A revision of the Afrotropical genus Dhatrichia (Trichoptera, Hydroptilidae)

Five previously described species of Dhatrichia Mosely, 1948 are recognized: D. inasa Mosely, 1948 from Yemen; D. bipunctata Statzner, 1977 from Zaire; D. lerabae (Gibon, Guenda & Coulibaly, 1994) comb. n. from Burkina Faso and Ghana; and D. cinyra Wells & Andersen, 1995 and D. divergenta Wells & Andersen, 1995 from Tanzania. A sixth species D. feredougoubae Gibon, 1987 from the Ivory Coast and Ghana is transferred to Microptila Ris, 1897 comb. n. In addition, nine new species are described and illustrated as males: D. ankasaensis sp. n., D. botiensis sp. n., D. hunukani sp. n., D. minuta sp. n., D. paraminuta sp. n. and D. wliensis sp. n. from Ghana, D. anderseni sp. n. from Tanzania, and D. madagascarensis sp. n. and D. giboni sp. n. from Madagascar. Females are associated, described and illustrated for all species, except D. inasa and D. giboni. The larvae and pupae of D. ankasaensis, D. hunukani, D. lerabae, D. minuta and D. wliensis are described and illustrated as the first known immatures of the genus. Keys to known larvae, pupae, males and females are provided. A phylogenetic analysis of the genus revealed four species groups. The most plesiomorphous taxa are West African, followed by East African and Malagasian taxa. A sister group relationship between the Eburneo–Ghanean and the Sudanian biogeographical regions is encountered twice. Among several possible sister taxa, the sister group turned out to be Kumanskiella Harris & Flint, 1992 and Microptila Ris, 1897 in part, combined. Implications for generic classification and biogeography are outlined. A morphometric principal component analysis revealed good separation of the sexes by the number and shape of antennal segments, and by eye size. Other measures are strongly dependent on overall size, and show best separation of females between species and species groups. A functional fit between male inferior appendages and secondary sexual characters in female sternite VIII is demonstrated for all species with associated females.

Katatopygia gen. n., a monophyletic branch segregated from Boletina (Diptera, Mycetophilidae)

Abstract The genus Katatopygia gen. n. is proposed for the Boletina erythropyga/punctus-group that was first introduced by Garrett (1924, 1925) and currently comprises eight described species. Molecular studies have strongly indicated that this group forms a monophyletic sister-group to a clade consisting of all other Boletina, Coelosia and Gnoriste, and its monophyly is supported by morphological data as well. The new genus includes the following species: Katatopygia antoma (Garrett, 1924), comb. n., Katatopygia antica (Garrett, 1924), comb. n., Katatopygia erythropyga (Holmgren, 1883), comb. n.,Katatopygia hissarica (Zaitzev & Polevoi, 2002), comb. n., Katatopygia magna (Garrett, 1925), comb. n., Katatopygia laticauda (Saigusa, 1968), comb. n., Katatopygia neoerythropyga (Zaitzev & Polevoi, 2002), comb. n. andKatatopygia sahlbergi (Lundström, 1906), comb. n., all transferred from Boletina. Katatopygia sahlbergi is found to be a senior synonym of Boletina punctus Garrett, 1925, syn. n. A phylogeny based on morphological data and using parsimony analysis yielded four most parsimonious trees where the new genus is retrieved as monophyletic with high support. Katatopygia neoerythropyga is found to be the sister-taxon to all other species that form two clades, one with Katatopygia sahlbergi-like species and one with Katatopygia erythropyga-like species. A key to males of Katatopygia is provided.

Review of the European Greenomyia Brunetti (Diptera, Mycetophilidae) with new descriptions of females

Abstract The females of the four continental Greenomyia Brunetti species in Europe are associated with the males, diagnosed and keyed, providing the first association and description of the females of Greenomyia baikalica Zaitzev, 1994 and Greenomyia stackelbergi Zaitzev, 1982. Colour photographs of their habitus and line drawings of their female terminalia are provided. Greenomyia mongolica Laštovka & Matile, 1974 is found to be a senior synonym of Greenomyia theresae Matile, 2002. syn. n. The diagnostic characters used to distinguish between Greenomyia and Neoclastobasis Ostroverkhova in keys did not hold up to a closer scrutiny and leave the status of Neoclastobasis as separate genus questionable.

Six new species of Afrotropical Allodia (Diptera: Mycetophilidae): DNA barcodes indicate recent diversification with a single origin

Only one species of the genus Allodia has been previously recorded from the Afrotropical region, Allodia (Brachycampta) flavorufa Matile, 1978. Six new species are described here, all representing the nominotypical subgenus, Allodia s.s. The new species are described from material collected in different mountainous areas in south and east Africa; A. jaschhofi sp. nov., A. karkloofensis sp. nov., A. drakensbergensis sp. nov., A. nyeriensis sp. nov., A. mazumbaiensis sp. nov. and A. keurbosensis sp. nov. The species are morphologically very similar, and can only be separated based on minor differences in wing venation and characters of the male terminalia. The genetic differences between the species in the DNA barcode region (CO1), however, support delimitation. The origin and distribution of these Afrotropical taxa, in relation to each other and to their Holarctic relatives, is discussed.