Awit Suwito

Taxonomy and phylogeny of Leptopilina species (Hymenoptera: Cynipoidea: Figitidae) attacking frugivorous drosophilid flies in Japan, with description of three new species

Despite the intensive use of the Leptopilina genus and its drosophilid hosts as model systems in the study of host–parasitoid interactions, the diversity and distribution of the species occurring in the Asian region remain elusive. Here we report the phylogeny of Japanese Leptopilina species attacking frugivorous drosophilid flies, based on COI, ITS1 and ITS2 sequences. Consistent with molecular data, hybridization experiments and morphological examination, five species were recorded in Japan: Leptopilina heterotoma, L. victoriae and three new species, two occurring in the Ryukyu archipelago, L. ryukyuensis and L. pacifica, and another species, L. japonica, distributed in Honshu and Hokkaido. Leptopilina japonica is further divided into two subspecies, L. j. japonica occurring in Japan, and L. j. formosana occurring in Taiwan. According to these results, we discuss the evolution, speciation and colonization history of Japanese Leptopilina species.

Diversity and abundance of frugivorous drosophilids and their parasitoids in Bogor, Indonesia

The diversity, abundance and association of frugivorous drosophilids and their parasitoids were studied in Bogor, Indonesia (tropical region), and compared with the results in Iriomote-jima (subtropical region) and Tokyo (temperate region). Among the adult drosophilid flies collected in traps baited with banana in wooded areas, the number of commonly observed frugivorous drosophilid species (i.e. species that occupied more than 0.5% of total drosophilid samples) was 10 in Bogor and nine in Iriomote-jima, more than in Tokyo (six species), probably reflecting the high diversity and abundance of fruits. The rate of parasitism was very high in Bogor, especially in species of the Drosophila ananassae and immigrans species groups. The diversity of parasitoids attacking frugivorous drosophilids was higher in Bogor and Iriomote-jima than in Tokyo, possibly because of the high species diversity of host drosophilids. Parasitoids generally showed wider latitudinal distributions than drosophilids. No remarkable difference was observed in the host range among tropical, subtropical and temperate parasitoids.

Genetic differentiation of Ganaspis brasiliensis (Hymenoptera: Figitidae) from East and Southeast Asia

Ganaspis brasiliensis (Ihering) (Hymenoptera: Figitidae: Eucoilinae) is a Drosophila parasitoid that has often been misidentified as G. xanthopoda (Ashmead) in recent studies. This study aims to clarify genetic differentiation of G. brasiliensis based on the nucleotide sequences of the mitochondrial cytochrome oxidase subunit 1 (CO1) gene and three nuclear DNA regions, the inter-transcribed spacers 1 and 2 (ITS1 and ITS2) and putative 60S ribosomal protein L37 (RpL37), as well as crossing experiments. Four lineages are recognized in individuals assigned as G. basiliensis by morphology, (1) individuals occurring in Japan and probably South Korea, (2) individuals from a small subtropical island of Japan, Iriomote-jima, (3) individuals from temperate lowlands of Japan and high altitude areas of Southeast Asia, and (4) individuals occurring widely in Asia, America, Hawaii and Africa. The first lineage is a specialist of Drosophila suzukii (Matsumura), a pest of fresh fruit, and also the fourth lineage has a capacity to parasitize this pest species. The first, third and fourth lineages occur sympatrically at least in Tokyo. The third and fourth lineages differed in mate choice and host use to some extent, but post-mating isolation between them was almost absent.

What Determines Host Acceptance and Suitability in Tropical Asian Drosophila Parasitoids

ABSTRACT For successful parasitism, parasitoid females must oviposit and the progeny must develop in individual hosts. Here, we investigated the determinants of host acceptance for oviposition and host suitability for larval development of Drosophila parasitoids from Bogor and Kota Kinabalu (≈1,800 km northeast of Bogor), Indonesia, in tropical Asia. Asobara pleuralis (Ashmead) from both localities oviposited frequently (>60%) in all of the drosophilid species tested, except the strain from Kota Kinabalu oviposited rarely (10%) in Drosophila eugracilis Bock & Wheeler. Leptopilina victoriae Nordlander from both localities only oviposited frequently (>77%) in species from the Drosophila melanogaster species group except D. eugracilis (<3.7%), whereas Leptopilina pacifica Novković & Kimura from Bogor oviposited frequently (>85%) only in species from the Drosophila immigrons species group. Thus, host acceptance appeared to be affected by host taxonomy, at least in Leptopilina species. Host suitability varied considerably, even among closely related drosophilid species, which suggests that the host suitability is at least in part independent of host taxonomy and that it has been determined via parasitoid-host coevolutionary interactions (i.e., arms race). Host acceptance did not always coincide with host suitability, i.e., parasitoids sometimes oviposited in unsuitable host species. Geographic origin strongly affected the host acceptance and suitability in the A. pleuralis—D. eugracilis parasitoid—host pair, whereas it only weakly affected the acceptability and suitability in other parasitoid—host combinations.

Colocasiomyia (Diptera: Drosophilidae) revised phylogenetically, with a new species group having peculiar lifecycles on monsteroid (Araceae) host plants.

The phylogeny of Colocasiomyia (Drosophilidae) is analysed using data for 70 morphological characters, many of which are re‐evaluated from or added to those used previously, for an expanded taxon sample of 24 Colocasiomyia ingroup species. A special focus is put on three species, of which two have remained unresolved for their relationships to other Colocasiomyia species, and the other is a newly discovered species. The analysis results in a single, most parsimonious cladogram, in which a clade comprising the three focal species is recognized along with other clades recovered for the known species groups of Colocasiomyia. Based on this, a new species group—the gigantea group—is established, including Colocasiomyia gigantea (Okada), C. rhaphidophorae Gao & Toda, n.sp. and C. scindapsae Fartyal & Toda, n.sp. These species of the gigantea group breed on inflorescences/infructescences of the subfamily Monsteroideae (Araceae) exceptionally among Colocasiomyia species, most of which use plants of the subfamily Aroideae as their hosts. Colocasiomyia gigantea uses Epipremnum pinnatum (L.) Engler, C. rhaphidophorae uses Rhaphidophora hookeri Schott and C. scindapsae uses Scindapsus coriaceus Engler as their hosts. The host plants of the gigantea group are epiphytes and differ in the structure of spadix and the fruiting process from those of the Aroideae. To understand how the species of the gigantea group adapt to properties of their host plants, their reproductive ecology—most intensively that of C. gigantea—is investigated. The lifecycle of C. gigantea is characterized by its relatively slow embryonic development (taking approximately 6 days), the very long duration of the full‐grown first instar within the egg capsule (approximately three months) until dehiscence of host infructescence, and its relatively fast larval and pupal development (taking approximately 11 or 12 days). Some morphological adaptations and the reproductive strategy in terms of ‘egg size vs. number’ trade‐off are discussed in relation to their reproductive habits and peculiar lifecycles.

Altitudinal patterns of abundances and parasitism in frugivorous drosophilids in west Java, Indonesia

Abundances and parasitism rates of frugivorous drosophilid flies were studied in three localities at altitudes of 250–1350 m in and near Bogor, Indonesia. The drosophilid and parasitoid species were classified into four groups: low-altitude species, high-altitude species, species abundant at a mid-altitude location and species occurring rather evenly from low to high locations. The ananassae and immigrans species groups were major drosophilids collected. All species of the ananassae species group were more abundant at lower altitudes, and the parasitism rate in this species group decreased with increasing altitude. Thus, the host abundance seems to affect the parasitism rate. On the other hand, the rate of parasitism in the immigrans species group showed no apparent relation with altitude or density, possibly due to the fact that species of this species group varied in altitudinal distribution. It is also suggested that the diversity of drosophilid species affects the composition of parasitoid species.

Assessment of fitness costs of resistance against the parasitoid Leptopilina victoriae in Drosophila bipectinata

How insects evolve resistance or counter-resistance against antagonists is a basic issue in the study of host-parasitoid coevolution. One of the factors that affect their coevolution is fitness costs of resistance and counter-resistance. Here, we assess fitness costs of resistance against the parasitoid Leptopilina victoriae in Drosophila bipectinata on the basis of selection experiments. We made a base population by mixing three geographic fly populations that differed in resistance. The established base population was divided into four populations, two for selection of resistance against a L. victoriae population and two for control. Resistance increased rapidly in response to selection and reached a very high level within four generations in the selected populations, while resistance of the control populations remained low during 20 generations. High resistance of the selected populations was maintained at least for 10 generations after selection was stopped. Both selected populations had lower female longevity than the control populations, and at least one of the selected populations had shorter thorax length, lower female desiccation tolerance and adult heat tolerance than both or either of the control populations. On the other hand, the selected populations had higher male starvation tolerance and longevity than the control populations. There were no significant differences in resistance against another population of L. victoriae and two other parasitoid species between the selected and control populations. These results suggest that the resistance against the L. victoriae population in D. bipectinata may incur some but not so high costs and act parasitoid-species- and/or parasitoid-population-specifically.

Diapause and cold tolerance in Asian species of the parasitoid Leptopilina (Hymenoptera: Figitidae)

Diapause and cold tolerance are essential for temperate insects to pass the winter, with the mechanisms controlling these two traits varying considerably among insects. In the present study, diapause and cold tolerance are compared among three Leptopilina species: Leptopilina japonica Novković & Kimura, Leptopilina victoriae Nordlander and Leptopilina ryukyuensis Novković & Kimura, all larval parasitoids of frugivorous drosophilid flies, with the aim of understanding their climatic adaptations. The first species is divided into the temperate (Leptopilina japonica japonica) and subtropical subspecies (Leptopilina japonica formosana), and the latter two species are distributed in the tropical and subtropical regions. The temperate subspecies of L. japonica enters prepupal diapause at low temperatures (15 or 18 °C), irrespective of photoperiod, and some individuals enter diapause when exposed to 0 °C for 1 or 2 day(s) or when placed at low humidity. Leptopilina victoriae also shows signs of diapause initiation at 15 °C, although L. ryukyuensis and L. j. formosana from the subtropical regions do not. Preimaginal viability at low temperature (13, 14 or 15 °C) is usually lower in L. victoriae from the tropical regions compared with L. japonica or L. ryukyuensis from the temperate or subtropical regions. Diapausing prepupae of the temperate subspecies appear to be cold tolerant. However, the cold tolerance of nondiapausing prepupae, pupae and adult females varies little among the tropical, subtropical and temperate species or subspecies, and adult males of the temperate subspecies of L. japonica are less cold tolerant than those of the tropical or subtropical species or subspecies. Cold tolerance may be unnecessary, except for diapausing individuals of the temperate species, because nondiapausing individuals appear in warmer seasons.

Discovery of the Drosophila (Drosophila) robusta species group (Diptera, Drosophilidae) from Southeast Asian tropics, with the descriptions of six new species

Six new species of the Drosophila robusta species group are described from Southeast Asian Islands. Kalimantan and Sunda Islands lie east of Bali, from which Drosophila barobusta sp.nov and D. uncinata sp.nov belong to the lacertosa subgroup, and D. sungaicola sp.nov, D. baliensis sp.nov, D. hitam sp.nov and D. subaquatica sp.nov to the okadai subgroup. The robusta group from Southeast Asian tropics exclusively inhabits streamsides in mountainous highlands with an elevation of more than 600 m from the sea level.

A New Species Group in The Genus Dichaetophora, with Descriptions of Six New Species from The Oriental Region (Diptera, Drosophilidae)

Abstract The genus Dichaetophora Duda comprises 61 described species classified into four species groups: agbo, tenuicauda, acutissima and sinensis. This genus is distributed exclusively in the Old World, and is rich in species in the tropical and subtropical areas of the Oriental, Australasian, and Afrotropical regions. In this paper, a new species group, the trilobita group, is established for six new species discovered from the Oriental region. The delimitation of these species is firstly performed in light of morphology and further with the aid of DNA sequences of the mitochondrial COI and COII (cytochrome c oxydase, subunits I and II, respectively) genes, considering also their respective geographical origins. Then, the new species (trilobita Yang & Gao, sp. n., heterochroma Yang & Gao, sp. n., flatosternata Yang & Gao, sp. n., borneoensis Yang & Gao, sp. n., javaensis Yang & Gao, sp. n., and sumatraensis Yang & Gao, sp. n.) are described, and a key, based on not only morphological but also molecular information, is provided.