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Acari : Mesostigmata) Associated with Dung Beetles Title (Coleoptera: Scarabaeidae) in Kalimantan, Indonesia

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Acari : Mesostigmata) Associated with Dung Beetles Title (Coleoptera: Scarabaeidae) in Kalimantan, Indonesia Taxonomy and Biogeography of the Family Macrochelidae (Acari : Mesostigmata) Associated with Dung Beetles Title (Coleoptera: Scarabaeidae) in Kalimantan, Indonesia Author(s) DWIBADRA, Dhian Citation 北海道大学. 博士(農学) 甲第11830号 Issue Date 2015-03-25 DOI 10.14943/doctoral.k11830 Doc URL http://hdl.handle.net/2115/60923 Type theses (doctoral) File Information Dhian_Dwibadra.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP Taxonomy and Biogeography of the Family Macrochelidae (Acari: Mesostigmata) Associated with Dung Beetles (Coleoptera: Scarabaeidae) in Kalimantan, Indonesia (インドネシア・カリマンタンにおける食糞性コガネムシ科甲虫に便乗するハエ ダニ科（ダニ亜綱：トゲダニ目）の分類および生物地理学的研究) Hokkaido University Graduate School of Agriculture Division of Environmental Resources Doctor Course Dhian Dwibadra Contents Abstract .......................................................................................................................... 1 Preface ........................................................................................................................... 4 I. Description of Phoretic Macrochelidae Associated with Dung Beetles in Kalimantan, Indonesia 1. Introduction .................................................................................................. 6 2. Materials and Methods ................................................................................ 8 3. General Morphology and Terminology ....................................................... 9 4. Remarks on the Description ......................................................................... 17 5. Description and Remarks ............................................................................. 19 Family Macrochelidae Vitzthum, 1930 ........................................................... 19 Genus Glyptholaspis Filipponi and Pegazzano, 1960 ..................................... 19 Glyptholaspis asperrima (Berlese, 1905) .............................................. 20 Glyptholaspis confusa (Foà, 1900) ...................................................... 21 Genus Holostaspella Berlese, 1903 ................................................................... 22 Holostaspella bifoliata (Trägårdh, 1952) ............................................... 23 Holostaspella katakurai Hartini and Takaku, 2003 ............................... 24 Holostaspella moderata Berlese, 1921 .................................................. 25 Genus Macrocheles Latreille, 1829 ................................................................... 26 Macrocheles baramensis Evans and Hyatt, 1963 .................................. 27 Macrocheles kalimantanensis Hartini and Takaku, 2003 ...................... 28 Macrocheles lumbiensis sp. nov. ............................................................ 31 Macrocheles dayaci Dwibadra and Takaku, 2014 ................................. 33 Macrocheles oigru Walter and Krantz, 1986 ......................................... 34 Macrocheles sp. aff. glaber (Mülller, 1860) .......................................... 35 Macrocheles hallidayi Walter and krantz, 1986 .................................... 36 Macrocheles kraepelini (Berlese, 1904) ................................................ 38 Macrocheles pumilus Hartini, Dwibadra and Takaku, 2009 .................. 39 Macrocheles riparius Dwibadra and Takaku, 2014 ............................... 40 Macrocheles dispar (Berlese, 1910) ...................................................... 42 Macrocheles entetiensis Hartini and Takaku, 2005 ............................... 43 Macrocheles jabarensis Hartini and Takaku, 2003 ................................ 44 Macrocheles nidus Hartini and Takaku, 2013 ........................................ 46 Macrocheles persimilis Hartini, Dwibadra and Takaku, 2007 ............... 47 Macrocheles krantzi Evans and Hyatt, 1963 ......................................... 48 Macrocheles sumbaensis Hartini and Takaku, 2005 .............................. 48 Macrocheles muscaedomesticae (Scopoli, 1772) .................................. 49 Macrocheles merdarius (Belese, 1889) ................................................. 50 Macrocheles mammifer Berlese, 1918 ................................................... 51 Macrocheles plumosus Evans and Hyatt, 1963 ..................................... 52 Macrocheles sukaramiensis Takaku, 2001............................................. 52 Macrocheles wainensis Dwibadra and Takaku, 2014 ............................ 53 Macrocheles lorensis sp. nov. ................................................................ 55 Genus Neopodocinum Oudemans, 1902 ............................................................ 56 Neopodocinum bosschai (Oudemans, 1901).......................................... 57 Neopodocinum halimunense Hartini and Takaku, 2003 ........................ 59 Neopodocinum kalimantanense Hartini and Takaku, 2004 ................... 60 Neopodocinum maius Berlese, 1911 ...................................................... 61 Neopodocinum subjaspersi Hartini and Takaku, 2003........................... 63 Neopodocinum murungensis sp. nov. ..................................................... 64 6. Key to species (female) of the Family Macrochelidae in Kalimantan ........ 68 7. Phoretic Association between Macrochelidae and Dung Beetles ................ 73 8. Summary ...................................................................................................... 80 II. Biogeography of the Macrochelid Mites in Southeast Asia, especially Indonesia 1. Introduction ................................................................................................ 82 2. Materials and Methods .............................................................................. 84 3. Results .......................................................................................................... 87 4. Discussion .................................................................................................... 89 5. Summary ........................................................................................................ 93 Acknowledgements ...................................................................................................... 95 References .................................................................................................................... 96 Appendix ..................................................................................................................... 114 Tables and Figures Abstract As a result of faunal survey on macrochelid mites in Kalimantan, Indonesia, 35 species of four genera belonging to the family Macrochelidae (Acari: Mesostigmata) associated with dung beetles (Scarabaeidae) were recorded and described. Three new species were described for the first time in the present thesis (Macrocheles lumbiensis, M. lorensis, and Neopodocinum murungensis), three species were already described and published by author during the study (M. dayaci, M. riparius, and M. wainensis), 16 species have already known previously, and 13 species recorded from Kalimantan for the first time. A key to 35 species (female) of the family in Kalimantan was provided. Based on morphological characters, especially dorsal and sternal characters, three species of the genus Holostaspella were classified into 3 species groups (bifoliata, mirabilis, and sculpta species group). Nineteen species of the genus Macrocheles maybe classified to the four species groups (glaber species group, muscaedomesticae group, subbadius group, and krantzi group) and seven species complexes. Five complexes are belonging to glaber group, i.e., glaber complex (including Macrocheles sp. aff. glaber (Müller, 1860), M. oigru Walter and Krantz, 1986, and M. dayaci Dwibadra and Takaku, 2014), kraepelini complex (including M. kraepelini (Berlese, 1905) and M. hallidayi Walter and Krantz, 1986), scutatus complex (M. dispar (Berlese, 1910), M. entetiensis Hartini and Takaku, 2005, M. jabarensis Hartini and Takaku, 2003, M. nidus Hartini and Takaku, 2013, and M. persimilis Hartini, Dwibadra and Takaku, 2007), pumilus complex (including M. pumilus Hartini, Dwibadra and Takaku, 2009 and M. riparius Dwibadra and Takaku, 2014), baramensis complex (including M. baramensis, M. kalimantanensis, and M. lumbiensis). One species complex, merdarius complex, is included in subbadius group. The krantzi species group and pumilus and baramensis 1 complexes belonging to glaber species group were established and their diagnoses were defined here. Macrochelids of Kalimantan (including Malaysian part) were found to be phoretic on 9 genera of scarab beetles Aphodius, Caccobius, Catharsius, Copris, Onitis, Oniticellus, Onthophagus, Paragymnopleurus, and Sisyphus (Coleoptera: Scarabaeidae), one genus of geotrupid beetle Bolbochromus (Geotrupidae), and one genus of scavenger scarabaeid beetle Phaeocroops (Hybosoridae). Twenty two species of mites were collected from the genus Catharsius, 19 spp. from Onthophagus and 13 spp. from Paragymnopleurus. These results suggest that larger sized genus or diversified genus of beetle tend to harbor large number of species of macrochelid mites. Most of macrochelid mites were associated with several genera of beetles (e.g., Macrocheles hallidayi were found on 5 genera of beetles, M. kalimantanensis on 7 genera of beetles), and macrochelids associated with dung beetles in Kalimantan did not show preference to particular species of dung beetles. In this study, five groups of geographical region based
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