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Velvet ants are characteristic wasps with extreme sexual dimorphism, with males usually being winged and females being wingless. They possess unique biological features resulting from flightless nature of females, which make them a useful model system in comparative biology. Most species are known as parasitoids of nest-provisioning wasps and bees. Given the high species richness and abundance, particularly in tropical or arid habitats, they potentially play an important role in natural and agricultural ecosystems. However, our knowledge on their systematics and natural history is fragmentary. Specifically, the East Asian velvet ants have been paid less attention comparing to other regions, despite of the potential diversity in this region. To fill these gaps, the present dissertation attempts to critically review the systematics of the East Asian velvet ants through examination of museum specimens and field surveys. Following a review of the velvet ant biology (Chapter 1), this dissertation includes: taxonomic review of the genus Eotrogaspidia Lelej, 1996 (Chapter 2), description a new genus previously confused with the genus Krombeinidia Lelej, 1996 (Chapter 3), taxonomic review of the genus Andreimyrme Lelej, 1995 based on the female morphology (Chapter 4), taxonomic review of Northeast Asian species of the genus Neotrogaspidia Lelej, 1996 (Chapter 5), taxonomic additions to the genus Mickelomyrme Lelej, 1995 (Chapter 6), faunal review and taxonomic additions to the velvet ants of Vietnam (Chapter 7) and Nepal (Chapter 8), taxonomic review and analysis of character displacement of the Far Eastern species of the genus Nemka Lelej, 1985 (Chapter 9), phylogenetic systematics of the East Asian genera of the tribe Smicromyrmini Bischoff, 1920 based on the male morphology (Chapter 10), and discussion on the utility of systematics within the context of comparative biology (Chapter 11). Based on over 3,500 specimens, I recognized many new taxa and taxonomic changes for East Asian velvet ants. Three new genera and 32 new species were described: Arkaditilla gen. nov. (type species: A. leleji sp. nov.), Chenmyrme gen. nov. (type species: Smicromyrme undata Chen, 1957), and Hadrodontilla gen. nov. (type species: Mutilla davidi André, 1898); Andreimyrme auricoma sp. nov., ♀ (Malaysia), An. laminatihumeralis sp. nov., ♀ (Indonesia, Malaysia), An. matsumotoi sp. nov., ♀ (Vietnam), An. naturalis sp. nov., ♀ (Indonesia), An. rong sp. nov., ♀ (Thailand, Vietnam), An. silvorientalis sp. nov., ♀ (Indonesia), An. takensis sp. nov., ♀ (Thailand), An. ursasolaris sp. nov., ♀ (Indonesia), An. yotoi sp. nov., ♀ (Laos, Vietnam), Ephucilla aequatoris sp. nov., ♂ (Indonesia), E. bicolor sp. nov., ♀ (Vietnam), E. bifida sp. nov., ♀ (Indonesia), E. okajimai sp. nov., ♀ (Thailand), E. retorta sp. nov., ♂ (Laos, Vietnam), Mickelomyrme carinata sp. nov., ♂ (Indonesia), Mi. cristaticauda sp. nov., ♂ (Vietnam), Mi. grandiguttata sp. nov., ♀ (Laos), Mi. recondita sp. nov., ♂ (Thailand), Mi. verticalis sp. nov., ♀ (Philippines), Mi. vietnaminix sp. nov., ♀ (Vietnam), Sinotilla complanata sp. nov., ♂ (Thailand), Si. unica sp. nov., ♂ (China, Vietnam), Smicromyrme (Smicromyrme) australis sp. nov., ♂ (Thailand) (Smicromyrmini), Arkaditilla frim sp. nov., ♂ (Malaysia), Ar. gibba sp. nov., ♂ (Indonesia), Ar. leleji sp. nov., ♂ (Indonesia), Eotrogaspidia adhabar sp. nov., ♂ (Nepal), Eo. buddha sp. nov., ♂ ((India, Nepal, Pakistan, Sri Lanka), Eo. hauseri sp. nov., ♂ (India, Nepal), Eo. melanopleura sp. nov., ♀ (China), (Trogaspidiini), Strangulotilla dubia sp. nov., ♂ (Nepal), and Zeugomutilla curvicarinata sp. nov., ♀ (Nepal) (Zeugomutillini). The following 36 new combinations and 28 new synonymies were proposed: An. borkenti (Williams, 2019), comb. nov. (from Smicromyrme), C. undata (Chen, 1957), comb. nov. (from Ephucilla), Ep. calacuasana (Tsuneki, 1993), comb. nov. (from Smicromyrme), Ep. cyaneiventris cyaneiventris (André, 1896), comb. nov. (from Sinotilla), Ep. cyaneiventris gobiana (Chen, 1957), comb. nov. (from Sinotilla), Ep. nigrothoracica (Okayasu, 2018), comb. nov. (from Sinotilla), Ep. petina (Mickel, 1937), comb. nov. (from Sinotilla), Ep. subparallela (Chen, 1957), comb. nov. (from Sinotilla), H. davidi (André, 1898), comb. nov. (from Andreimyrme), H. pekiniana (André, 1905), comb. nov. (from Sinotilla), Mi. atrata (Mickel, 1934), comb. nov. (from Smicromyrme), Mi. autonoe (Mickel, 1934), comb. nov. (from Smicromyrme), Mi. caecina (Cameron, 1903), comb. nov. (from Smicromyrme), Mi. meator (Mickel, 1935), comb. nov. (from Smicromyrme), Mi. lavinia (Mickel, 1934), comb. nov. (from Smicromyrme), Mi. palacala (Tsuneki, 1993), comb. nov. (from Smicromyrme), Promecilla annexa (Cameron, 1909), comb. nov. (from Andreimyrme), Si. dardanus (Smith, 1857), comb. nov. (from Smicromyrme), Si. drola drola (Zavattari, 1913), comb. nov. (from Ephucilla), Si. drola lodra (Chen, 1957), comb. nov. (from Ephucilla), Si. ilerda (Cameron, 1902), comb. nov. (from Smicromyrme), Si. nitela (Mickel, 1934), comb. nov. (from Andreimyrme), Si. scitula (Mickel, 1935), comb. nov. (from Smicromyrme), Si. sparsilis (Mickel, 1934), comb. nov. (from Smicromyrme), Si. viriata (Mickel, 1934), comb. nov. (from Andreimyrme), Si. volupia (Mickel, 1935), comb. nov. (from Andreimyrme) (Smicromyrmini), Ar. bagrada (Cameron, 1902), comb. nov. (from Krombeinidia), Ar. depressicornis (Mickel, 1935), comb. nov. (from Krombeinidia), Ar. nallinia (Zavattari, 1914), comb. nov. (from Krombeinida), Eo. lena (Cameron, 1899), comb. nov. (from Trogaspidia), Eo. rubripes (André, 1901), comb. nov. (from Trogaspidia), Eo. saussurei (Lelej, 2005), comb. nov. (from Trogaspidia), Neotrogaspidia circumcincta (André, 1896), comb. nov. (from Orientidia), N. elpinice (Mickel, 1933), comb. nov. (from Trogaspidia), N. recticarinata (Chen, 1957), comb. nov. (from Trogaspidia), and Vanhartenidia ekka (Nurse, 1902), comb. nov. (from Eotrogaspidia) (Trogaspidiini); C. undata (= Sm. yuliana Tsuneki, 1972, syn. nov.), Ep. naja (Zavattari, 1913) (= Sm. thalia Mickel, 1933, syn. nov.), Ep. petina (Mickel, 1937) (= Sm. spinicauda Chen, 1957, syn. nov., Si. lambirensis Lelej, 1996, syn. nov., Si. belokobylskiji Lelej, 1995, syn. nov., Si. calopoda Okayasu, 2017, syn. nov., and Si. ohbayashii Okayasu, 2017, syn. nov.), Mi. athalia (Pagden, 1949) (= Sm. bicristata Chen, 1957, syn. nov.), Mi. bakeri (Mickel, 1934) (= Sm. tanoi Tsuneki, 1972, syn. nov.), Mi. caecina (= Sm. punctinotus Mickel, 1935, syn. nov.), Mi. norna (Zavattari, 1913) (= Sm. norna abnorma Chen, 1957, syn. nov.), Mi. zebina (Smith, 1860) (= Sm. aborlana aborlana Tsuneki, 1993, syn. nov. and Sm. aborlana zamboangae Tsuneki, 1993, syn. nov.), Nemka taiwanensis (= Sm. curvisquamata Chen, 1957, syn. nov. and Sm. ilanica Tsuneki, 1972, syn. nov.), P. decora (Smith, 1879) (= Sm. contractula Chen, 1957, syn. nov.), Si. dardanus (= Sm. dardanus salacia Mickel, 1935, syn. nov., Sm. gribodoana Invrea, 1943, syn. nov., and Sm. cyaneiventris boheana Chen, 1957, syn. nov.), Si. drola drola (= Sm. hombuceiana Tsuneki, 1982, syn. nov.), Si. nitela (= Sm. aponis Tsuneki, 1993, syn. nov.), and Si. viriata (= Sm. mindanaonis Tsuneki, 1993, syn. nov.) (Smicromyrmini), Eo. auroguttata (Smith, 1855) (= T. vallicola Tsuneki, 1993, syn. nov.), Neo. elpinice (= T. aestivalis Hammer, 1949, syn. nov. and T. maritima Chen, 1957, syn. nov.), Neo. circumcincta (= T. yuliensis Tsuneki, 1972, syn. nov. and K. agricola Williams, 2019, syn. nov.), and T. mackieae (Cockerell, 1928) (= T. pacifica Tsuneki, 1972, syn. nov.) (Trogaspidiini). Two species were resurrected as full species: Neo. elpinice, stat. resurr. from the synonym of Neo. pustulata and T. acidalia (Cameron, 1897), stat. resurr. from the synonym of T. villosa (Fabricius, 1775). The type species of Sinotilla is replaced by Si. dardanus under Article 70.3.2 of the International Code of Zoological Nomenclature 4th edition. Systematic and phylogenetic results of this study suggest that: 1) the diversity of East Asian velvet ants is much higher than currently recognized, but sampling efforts are inadequate and biased; 2) the generic assignment of many previously described species is inappropriate and should be critically revised based on reexamination of voucher specimens; and 3) the East Asian genera of the Smicromyrmini are defined only by a unique combination of character states due to high level of homoplasy.
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