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, 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.
Abstract. To understand the factors governing the diversity, abundance and host associations of parasitoids attacking frugivorous drosophilid flies on Iriomote-jima, a subtropical island of Japan, we monitored parasitism on several occasions over the period [2003][2004][2005][2006][2007][2008][2009] In addition, host acceptance and host suitability of the four most frequently recorded parasitoid species were studied in the laboratory. Most parasitoid and drosophilid species showed species-specific associations with more than one antagonist species, suggesting that they have been subjected to complex coevolutionary interactions. In addition, host range of most of the parasitoid species included one of the three major Drosophila species, suggesting that the abundance of potential hosts is one of the factors determining the evolution of parasitoid host use.
Local adaptation and ecological fitting are the major processes by which organisms colonize and persist in novel environments and form novel associations with unfamiliar species. In this study, we assessed the role of these two processes in the formation of associations between the parasitoid Leptopilina japonica and various host Drosophila species using laboratory populations from three different localities. Each L. japonica population was highly virulent to some Drosophila species from its original locality, meaning they used these Drosophila species as major hosts. However, major host species at a given locality were usually less successfully parasitized by parasitoid populations from different localities. These results indicate that the investigated parasitoid populations showed local adaptation in host use. We also observed cases in which unfamiliar Drosophila species were successfully parasitized or allopatric Drosophila populations were more successfully parasitized. These findings suggest that L. japonica can form novel associations with some Drosophila species in novel environments without undergoing adaptive changes, a phenomenon called ecological fitting. In this study, L. japonica also usually appeared to have oviposited well in D. melanogaster species group members irrespective of their suitability as hosts. This result suggests that the parasitoid makes oviposition decisions based on characteristics common to the drosophilids in this species group
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.
Island biotas provide a great opportunity to study not only the phylogeographic patterns of a group of species, but also to explore the differentiation in their coevolutionary interactions. Drosophila and their parasitoids are exemplary systems for studying complex interaction patterns. However, there is a lack of studies combining interaction-based and molecular marker-based methods. We applied an integrated approach combining phylogeography, interaction, and host-choice behavior studies, with the aim to understand how coevolutionary interactions evolve in Drosophila-parasitoid island populations. The study focused on the three most abundant Drosophila species in Ryukyu archipelago and Taiwan: D. albomicans, D. bipectinata, and D. takahashii, and the Drosophila-parasitoid Leptopilina ryukyuensis. We determined mitochondrial COI haplotypes for samples representing five island populations of Drosophila and four island populations of L. ryukyuensis. We additionally sequenced parts of the autosomal Gpdh for Drosophila samples, and the ITS2 for parasitoid samples. Phylogenetic and coalescent analyses were used to test for demographic events and to place them in a temporal framework. Geographical differences in Drosophila-parasitoid interactions were studied in host-acceptance, host-suitability, and host-choice experiments. All four species showed species-specific phylogeographic patterns. A general trend of the haplotype diversity increasing towards the south was observed. D. albomicans showed very high COI haplotype diversity, and had the most phylogeographically structured populations, with differentiation into the northern and the southern population-group, divided by the Kerama gap. Differentiation in host suitability was observed only between highly structured populations of D. albomicans, possibly facilitated by restricted gene flow. Differentiation in host-acceptance in D. takahashii, and host-acceptance and host-choice in L. ryukyuensis was found, despite there being no differentiation in these two species according to molecular markers. Host choice assays show that L. ryukyuensis populations that have had more time to coevolve adapt their behavior to exploit the most suitable host – D. albomicans. L. ryukyuensis parasitoids on border ranges may, on the other hand, benefit from broader host-acceptance, that may facilitate adaptation to uncertain and variable environments. All results indicate that Drosophila-parasitoid populations in the Ryukyu archipelago and Taiwan have different evolutionary trajectories, and coevolve in a dynamic, complex, and local-specific way.
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