Kobe University Repository : Thesis Systematic study of the subfamily Acaenitinae (Hymenoptera, 学位論文題目 Ichneumonidae) based on morphological and molecular characters(形 Title 態および分子形質を用いたケンオナガヒメバチ亜科（ハチ目、ヒメバ チ科）の分類学的研究) 氏名 Ito, Masato Author 専攻分野 博士（農学） Degree 学位授与の日付 2017-03-25 Date of Degree 公開日 2019-03-25 Date of Publication 資源タイプ Thesis or Dissertation / 学位論文 Resource Type 報告番号 甲第6948号 Report Number 権利 Rights JaLCDOI URL http://www.lib.kobe-u.ac.jp/handle_kernel/D1006948 ※当コンテンツは神戸大学の学術成果です。無断複製・不正使用等を禁じます。著作権法で認められている範囲内で、適切にご利用ください。 PDF issue: 2019-04-18 Doctoral Dissertation Systematic study of the subfamily Acaenitinae (Hymenoptera, Ichneumonidae) based on morphological and molecular characters Masato ITO Laboratory of Insect Biodiversity and Ecosystem Science, Graduate School of Agricultural Science, Kobe University February, 2017 [Unpublished for the purposes of Zoological Nomenclature (Art. 8.2, ICZN)] Doctoral Dissertation Systematic study of the subfamily Acaenitinae (Hymenoptera, Ichneumonidae) based on morphological and molecular characters 形態および分子形質を用いたケンオナガヒメバチ亜科 （ハチ目、ヒメバチ科）の分類学的研究 Masato ITO 伊藤 誠人 Laboratory of Insect Biodiversity and Ecosystem Science, Graduate School of Agricultural Science, Kobe University February, 2017 Table of Contents Chapter 1 ―Introduction.......……………………....……………………....…..………1 1. 1 Family Ichneumonidae.....…………………....……………………. ... ....…….2 1. 2 Subfamily Acaenitinae.....……………………………. ... ..……………………2 1. 3 Aims of this study.....…………………………………………………………..4 References………………………...………………………………………………….4 Chapter 2 ― Systematics of the Subfamily Acaenitinae from Japan……………….7 2. 1 Introduction……………………………………………………………………8 2. 2 Materials and methods…………………………………………………………8 2. 3 Revision of Japanese Acaenitinae…………………………………………….10 References……………………………………………………………...……………91 Chapter 3 ― Molecular Phylogeny of the Subfamily Acaenitinae………...………….97 3. 1 Introduction………………………………………………………..………….98 3. 2 Materials and methods………………………………………….…………….98 3. 3 Phylogenetic analyses based on COI sequence………………….……………99 3. 4 Phylogenetic analyses based on 28S sequence……………………..………..100 3. 5 Discussion……………………………………………………….………100 References………………………………………………………………………….101 Chapter 4 ― Intraspecific Body Color Variation………………………………….....103 4. 1 Introduction………………………………………………………………….104 4. 2 Materials and methods…………………………………….…………………104 4. 3 Result………………………………………………………………………105 4. 4 Discussion……………………………………………………………………105 References………………………………………………………………………….106 Chapter 5 ―General Discussion…………………………………………………..108 5. 1 Species diversity of Acaenitinae from Japan………………………………...109 5. 2 Usability of molecular approaches in parasitoid taxonomy……………….109 5. 3 Intraspecific body color variation in Spilopteron and other Acaenitinae i species……………………..………………………………………………109 5. 4 Conclusion…………………………………………………………………110 References……………………………………………………………………….….110 Summary……………………………………………………………………………112 Acknowledgements…………………………………………………………………116 Figures……………………………………………………………………………….117 Tables………………………………………………………………………………..164 Appendix……………………………………………………………………………168 Some parts of this thesis have been published as follows: Chapter2 Masato ITO, Kyohei WATANABE & Kaoru MAETO: Spilopteron luteum Uchida and S. mucronatus Lee (Hymenoptera, Ichneumonidae, Acaenitinae) new to Japan. Japanese Journal of Systematic Entomology, 18(2): 443-446, 2012. Masato ITO, Kyohei WATANABE & Kaoru MAETO: A review of Metachorischizus unicolor Uchida, 1928, with the first description of males (Hymenoptera, Ichneumonidae, Acaenitinae). Japanese Journal of Systematic Entomology, 19(1): 181-185, 2013. Masato ITO & Kaoru MAETO: Revision of the genus Yamatarotes Uchida (Hymenoptera, Ichneumonidae, Acaenitinae) from Japan, based on morphological and molecular evidence. Japanese Journal of Systematic Entomology, 20(1): 107-113, 2014. ii Masato ITO, Kyohei WATANABE & Kaoru MAETO: Revision of the genus Arotes Gravenhorst (Hymenoptera: Ichneumonidae: Acaenitinae) from Japan. Zootaxa, 3893(2): 196-208, 2014. Masato ITO & Kaoru MAETO: Phaenolobus koreanus Uchida (Hymenoptera, Ichneumonidae, Acaenitinae) new to Japan. Japanese Journal of Systematic Entomology, 20(2): 353-355, 2014. Masato ITO & Kaoru MAETO: Revision of the genus Jezarotes Uchida (Hymenoptera: Ichneumonidae: Acaenitinae), with the description of a new species from Laos. Zootaxa, 3946 (3): 416–426, 2015. Masato ITO & Kaoru MAETO: Two species of the genus Coleocentrus (Hymenoptera: Ichneumonidae: Acaenitinae) new to Japan. Japanese Journal of Systematic Entomology, 21(1): 87-89, 2015. Masato ITO, Kyohei WATANABE & Kaoru MAETO: Molecular evidence resolving the confusion of two ichneumonid species of Spilopteron (Hymenoptera) caused by marked geographical colour variation. European Journal of Entomology, 112 (3): 543-556, 2015. Masato ITO & Kaoru MAETO: Revision of Ishigakia Uchida (Hymenoptera: Ichneumonidae: Acaenitinae) from Japan, with a new species having a close relative in South Africa. Zootaxa, 4136: 174–180, 2016. Masato ITO & Kaoru MAETO: Two species of the genus Yezoceryx (Hymenoptera: Ichneumonidae: Acaenitinae) new to Japan. Japanese Journal of Systematic Entomology, 22: 37–38, 2016. iii Masato ITO & Kaoru MAETO: Revision of the genus Spilopteron Townes (Hymenoptera: Ichneumonidae: Acaenitinae) from Japan. European Journal of Taxonomy, in press. Chapter4 Masato ITO, Kyohei WATANABE & Kaoru MAETO: Molecular evidence resolving the confusion of two ichneumonid species of Spilopteron (Hymenoptera) caused by marked geographical colour variation. European Journal of Entomology, 112 (3): 543-556, 2015. iv Chapter 1 ― Introduction 1 1. 1 Family Ichneumonidae The family Ichneumonidae of the order Hymenoptera (Insecta) contains many current and many more potential biocontrol agents against pest insects in agriculture and forestry (e.g., Neumann & Morey, 1984; Philip et al., 1993). This is one of the largest taxa of all animal groups, and includes more species than the entire subphylum Vertebrata and more than any other insect family with the possible exception of the coleopterous family Curculionidae (Townes, 1969). According to Yu et al. (2012), 1,579 genera and 24,281 species of Ichneumonidae (including fossil species) have been described from all over the world, but these are thought to be a small proportion of the actual numbers. Townes (1969) calculated that there were 60,000 species of ichneumonids in the world, but most current researchers have considered this as an underestimate. For example, Gauld (2002) estimates the number of ichneumonid species to be more than 100,000. Ichneumonids are distributed across the whole world except for the polar regions, and are commonly found in a wide variety of environments such as waterside vegetation, deserts, tropical rainforests, temperate regions, grasslands, and even in suburban gardens (Gauld, 1991). The classification of this parasitoid family has long been based on morphological characteristics such as body proportion, surface structure, wing venation, hair distribution, and so forth. The use of body color has also been common for species identification, because of the ease of using this characteristic and the fact that it is reliable in some instances (Quicke et al., 2006). However, there is often large intraspecific variation in body color as well as in other morphological characters due to both genetic and environmental factors (Abe et al., 2013). Thanks to recent developments in molecular taxonomy, mitochondrial and nuclear DNA sequencing have increasingly been used for the species-level classification of parasitoid wasps (e.g., Quicke et al., 2006; Rugman-Jones et al., 2009; Stigenberg et al., 2011). Accurate biological classification using both morphological and molecular data is now possible and practical, but many parasitoid groups are still far from seeing the benefit of molecular taxonomy. 1. 2 Subfamily Acaenitinae The subfamily Acaenitinae is a small taxa of the family Ichneumonidae, and 27 genera and about 275 species have been recorded from around the world, of which 82 are known from the Palearctic region, 21 from the Nearctic region, 77 from the Oriental 2 region, 56 from the Ethiopian region, five from the Australian region, and three from the Neotropical region (Yu et al. 2012). 1. 2. 1 Phylogeny This subfamily is defined by having the following four shared-derived character-states: clypeus with a rounded declivity at midpoint, propodeal spiracle elongate, metasomal tergite 2 with anterolateral carinae, and female hypopygium large and triangular (Wahl & Gauld, 1998). The subfamily Acaenitinae had been divided into two tribes: Acaenitini and Coleocentrini. Acaenitini had the following five character-states in common: fore and mid tarsal claws with an acute accessory tooth (without it in Coleocentrini), frons with a median vertical carina (without it in Coleocentrini), forewing vein 3rs-m absent (present in Coleocentrini), teeth on apex of lower valve of ovipositor reduced or absent (well developed in Coleocentrini), and fifth hind tarsomere longer than the second one (as long as second one in Coleocentrini) (Townes, 1971). However, Wahl and Gauld (1998) reported that the above characters of Coleocentrini were plesiomorphic with respect to the characters used to define Acaenitini based on their morphological phylogenetic analysis, and now these tribes are usually not accepted. Nevertheless, molecular phylogenetic relationships among genera of the subfamily Acaenitinae remain unresolved. 1. 2. 2 Bionomics Ichneumonids