Structures and Functions of the Endophallic Copulatory Tube in the Family Staphylinidae (Insecta: Coleoptera)

  • Shun-Ichiro Naomi


This paper deals with the structures and functions of the endophallic copulatory tube in Staphylinidae, which has been previously called “copulatory piece,” “endophallus,” “flagellum,” etc. in descriptive studies. First, the general morphology of the copulatory tube and the pseudocopulatory tube is discussed, together with the characterizations and terminology. Second, the distribution of the copulatory tubes in Staphylinidae is described. Third, the structures of the seven basic and some other unique forms of the copulatory tubes in Staphylinidae are described in detail, paying special attention to those of Steninae. Fourth, the functions of the copulatory tubes in Staphylinidae are described or hypothesized, paying special attention to those of Steninae. Namely, the function of the copulatory tube is simply a “spermatophore (or sperm) depositor” in some cases (e.g., some Stenus), whereas it has double function: a “spermatophore (or sperm) depositor” and an “extension tube (or sperm) guiding rod in some other cases (e.g., some Aleochara).



I would like to express my sincere gratitude to Dr. Alessandro Minelli (Padova University) and to Dr. Margaret Thayer (Field Museum) for their constructive and also critical comments on my earlier versions of the draft of this paper and to Dr. Yoko Matsumura (Hokkaido University) for her useful discussion with me about the morphology of the flagellum in Coleoptera. I sincerely thank Dr. Volker Puthz (Schlitz) for his kindness in sending me some valuable Stenus specimens for this study and also for informing me of some interesting forms of Stenus flagella, Dr. Alexey Shavrin (Daugavpils University) for letting me know valuable information on the copulatory tubes in Omaliinae, and Dr. Volker Assing (Hannover) for informing me of the matter with the paederine flagellum. My hearty thanks are due to Dr. Munetoshi Maruyama (Kyushu University Museum) for providing important information on the morphology of the coleopterous flagellum and to Dr. Hiroyuki Yoshitomi (Ehime University) for his advices for this study in various ways. Finally I sincerely thank the editors of this book for improving the manuscript of my paper in various ways.


  1. Abarbanell NR, Ashe JS (1989) Revision of the species of Pinophilus Gravenhorst (Coleoptera: Staphylinidae) of America north of Mexico. Field Zool 54:1–32Google Scholar
  2. Ahn KJ (1996) A review of Diaulos Casey (Coleoptera: Staphylinidae: Aleocharinae), with description of a new species and known larvae. Col Bull 50:270–290Google Scholar
  3. Ahn KJ (1997) A review of Liparocephalus Mäklin (Coleoptera: Staphylinidae: Aleocharinae), with descriptions of larvae. Pan-Pacif Ent 73:79–92Google Scholar
  4. Ahn KJ (2001) Phylogenetic relationships of the intertidal genus Halorhadinus Sawada and key to the genera of the Liparocephalini (Coleoptera: Staphylinidae: Aleocharinae). Insect Syst Evol 32:123–132CrossRefGoogle Scholar
  5. Ahn KJ, Ashe JS (1996) A revision of Rothium Moore and Legner (Coleoptera: Staphylinidae: Aleocharinae) with a discussion of its phylogenetic relationships. J Kansas Ent Soc 69:234–256Google Scholar
  6. Arai S, Nomura S (2003) Discovery of the tribe Odontalgini (Coleoptera: Staphylinidae; Pselaphinae) in Japan. Ent Sci 6:193–197CrossRefGoogle Scholar
  7. Ashe JS (1984) Generic revision of the subtribe Gyrophaenina (Coleoptera: Staphylinidae: Aleocharinae) with review of the described subgenera and major features of evolution. Quaest Ent 20:129–349Google Scholar
  8. Ashe JS (1986) Phanerota cubensis and Phanerota brunnessa n. sp., with a key to the species of Phanerota occurring in Florida (Coleoptera: Staphylinidae). Florida Ent 69:236–245CrossRefGoogle Scholar
  9. Ashe JS (1992) Phylogeny and revision of genera of the subtribe Bolitocharina (Coleoptera: Staphylinidae: Aleocharinae). Univ Kansas Sci Bull 54:335–406CrossRefGoogle Scholar
  10. Ashe JS (2003) First record and four new species of Adelarthra Cameron 1920 from Australia, and a related new genus and species Notiomerinx zealandica, from New Zealand (Coleoptera: Staphylinidae: Aleocharinae). In: Cuccodoro G, Leschen RA (eds) Systematics of Coleoptera: Papers celebrating the retirement of Ivan Löbl. Memoirs on Entomology International 17:617–636Google Scholar
  11. Ashe JS, Newton AF Jr (1993) Larvae of Trichophya and phylogeny of the tachyporine group of subfamilies (Coleoptera, Staphylinidae) with a review, new species and characterization of the Tachyporinae. Syst Ent 18:267–286CrossRefGoogle Scholar
  12. Assing V (1999) A revision of Othius Stephens (Coleoptera, Staphylinidae). VIII. Further records, new species, and a new synonym. Linzer biol Beitr 31:661–691Google Scholar
  13. Assing V (2001) The first record of Dinusa Saulcy 1864 from Turkey (Coleoptera: Staphylinidae, Aleocharinae). Linzer biol Beitr 33:187–190Google Scholar
  14. Assing V (2002) On some micropterous species of Athetini from Nepal and China (Coleoptera: Staphylinidae, Aleocharinae). Linzer biol Beitr 34:953–969Google Scholar
  15. Assing V (2003) A revision of Zoosetha Mulsant & Rey and Poromniusa Ganglbauer. II. A new species from Spain and additional records (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa 310:1–8CrossRefGoogle Scholar
  16. Assing V (2006) A revision of the Palaearctic species of Orphnebius Motschulsky (Coleoptera: Staphylinidae: Paederinae). Ent Prob 36:1–26Google Scholar
  17. Assing V (2012) On the taxonomy and zoogeography of some Oxypoda species of the West Palaearctic region (Coleoptera: Staphylinidae: Aleocharinae). Linzer biol Beitr 44:365–399Google Scholar
  18. Assing V (2013) On the Palaearctic and Oriental species of Scymbalium and Micrillus (Coleoptera: Staphylinidae: Paederinae). Linzer biol Beitr 45:1479–1520Google Scholar
  19. Assing V (2014) A revision of the species of Pinobius MacLeay, 1871 of the Oriental, Palaearctic, and Australian Regions (Coleoptera: Staphylinidae: Paederinae). Kol Rund 84:115–191Google Scholar
  20. Assing V, Wunderle P (1995) A revision of the species of the subfamily Habrocerinae (Coleoptera: Staphylinidae) of the world. Rev Suisse Zool 102:307–359CrossRefGoogle Scholar
  21. Blackwelder R (1936) Morphology of the coleopterous family Staphylinidae. Smith misc coll 94:1–102Google Scholar
  22. Bordoni A (2002) Xantholinini della Regione Orientale (Coleoptera: Staphylinidae). Classificazione, filogenesi e revisione tassonomica. Museo Regionale di Scienze Naturali Monografie Turin 33:1–998Google Scholar
  23. Bordoni A (2011) Notes on Palaearctic Xantholinini. VI. New species and new records from Caucasus (Coleoptera: Staphylinidae). Fragm ent 43:41–56CrossRefGoogle Scholar
  24. Brunke AJ, Chatzimanolis S, Schillhammer H, Solodovnikov A (2016) Early evolution of the hyperdiverse rove beetle tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae) and a revision of its higher classification. Cladistics 32:427–451CrossRefGoogle Scholar
  25. Clarke DJ, Grebennikov VV (2009) Monophyly of Euaesthetinae (Coleoptera: Staphylinidae): phylogenetic evidence from adults and larvae, review of austral genera, and new larval description. Syst Ent 34:346–397CrossRefGoogle Scholar
  26. Coiffait H (1972) Coléoptères Staphylinidae de la région Paléarctique occidentale. I. Suppl Nouv Rev Ent 2:1–651Google Scholar
  27. Coiffait H (1974) Coléoptères Staphylinidae de la région Paléarctique occidentale. III. Suppl Nouv Rev Ent 4:1–593Google Scholar
  28. Crowson RA (1981) The biology of the Coleoptera. Academic Press, LondonGoogle Scholar
  29. Frisch J (1997) A revision of some West Palaearctic species of Scopaeus Erichson (Coleoptera, Staphylinidae, Paederinae). Rev Suisse Zool 104:523–557CrossRefGoogle Scholar
  30. Frisch J (1998) A revision of some West Palaearctic species of Scopaeus Erichson (Coleoptera, Staphylinidae, Paederinae). Rev Suisse Zool 105:89–124CrossRefGoogle Scholar
  31. Frisch J (1999) A revision of the Scopaeus debilis species group, with description of a new species from Madagascar (Coleoptera, Staphylinidae, Paederinae). Rev Suisse Zool 106:361–383CrossRefGoogle Scholar
  32. Frisch J (2003) A revision of the Scopaeus laevigatus species group, with descriptions of ten new species from the East Palaearctic, the Oriental and the Australian regions (Coleoptera, Staphylinidae, Paederinae). In: Cuccodoro G, Leschen RA (eds) Systematics of Coleoptera: Papers celebrating the retirement of Ivan Löbl. Memoirs on Entomology International 17:649–726Google Scholar
  33. Gack C, Peschke K (1994) Spermathecal morphology, sperm transfer and a novel mechanism of sperm displacement in the rove beetle, Aleochara curtula (Coleoptera, Staphylinidae). Zoomorphology 114:227–237CrossRefGoogle Scholar
  34. Gack C, Peschke K (2005) ‘Shouldering’ exaggerated genitalia: a unique behavioural adaptation for the retraction of the elongate intromittent organ by the male rove beetle (Aleochara tristis Gravenhorst). Biol J Linn Soc 84:307–312CrossRefGoogle Scholar
  35. Gusarov V (2003) Revision of some types of North American aleocharines (Coleoptera: Staphylinidae: Aleocharinae), with synonymic notes. Zootaxa 353:1–134CrossRefGoogle Scholar
  36. Hammond PM (1973) Notes on British Staphylinidae 3. The British species of Sepedophilus Gistel (Conosomus auctt.) Ent Mon Mag 108:130–165Google Scholar
  37. Hemming BS (2003) Insect development and evolution. Cornell University Press, New YorkGoogle Scholar
  38. Herman LH (1972) A revision of the rove-beetle genus Charhyphus (Coleoptera, Staphylinidae, Phloeocharinae). Am Mus Nov 2496:1–16Google Scholar
  39. Herman LH (1975) Revision and phylogeny of the monogenetic subfamily Pseudopsinae for the world (Staphylinidae, Coleoptera). Bull Am Nat Hist 155:245–317Google Scholar
  40. Herman LH (1981) Revision of the subtribe Dolicaonina of the New World, with discussions of phylogeny and the old world genera (Staphylinidae, Paederinae). Bull Am Mus Nat Hist 167:327–520Google Scholar
  41. Hlaváč P, Nomura S (2001) A taxonomic revision of Tyrini of the Oriental region 1. Paralasinus (Coleoptera, Staphylinidae, Pselaphinae), a new genus of Tyrina from Indochina. Elytra 29:163–174Google Scholar
  42. Hoebeke ER, Ashe JS (1994) New species of Autalia Leach 1819 from the Neotropics (Coleoptera Staphylinidae Aleocharinae), with new distributional data for A. phricotrichosa Hoebeke 1988 and key to the Neotropical species of Autalia. Tropical Zool 7:191–208CrossRefGoogle Scholar
  43. Hoshina H, Sugaya H (2003) New records of the genera Scaphobaeocera and Scaphoxium (Coleoptera: Staphylinidae) from the Ryukyus, Japan, with descriptions of two new species. Ent Rev Jpn 58:35–41Google Scholar
  44. Hwang WS, Ahn KJ (2000) Taxonomy of the Korean Oxyporinae (Insecta, Coleoptera, Staphylinidae). Korean J Syst Zool 16:191–202Google Scholar
  45. Irmler U (1981) Descriptions of new neotropical Holotrochus and a key to the species of the genus (Coleoptera: Staphylinidae). Col Bull 35:379–397Google Scholar
  46. Irmler U (1985) Neue Arten der Gattungen Aueucamptus und Thoracophorus (Col., Staphylinidae) aus der Neotropis. Ent Blät 81:41–58Google Scholar
  47. Irmler U (2005) New neotropical species of the genus Holotrochus (Coleoptera: Staphylinidae: Osoriinae). Sci Pap Nat Hist Mus Univ Kan 36:1–18Google Scholar
  48. Jałoszyński P (2010) Cephenniini of the Philippines. Part 3. New species of Cephennomicrus Reitter and Hlavaciellus Jałoszyński from Palawan (Coleoptera: Staphylinidae: Scydmaeninae). Genus 21:13–20Google Scholar
  49. Jałoszyński P, Nomura S (2009) The Cephenniini (Coleoptera, Scydmaenidae) of Vietnam. Bull Natl Mus Sci Ser A 35:167–225Google Scholar
  50. Jeannel R (1959) Révision des Psélaphides de l’Afrique intertropicale. Ann Mus r Congo Belge Tervuren Ser 8 Sci Zool 75:1–742Google Scholar
  51. Kanao T, Maruyama M, Hashim R (2011) A new genus of Compactopediina (Coleoptera: Staphylinidae) associated with Longipeditermes (Isoptera: Termitidae) in Peninsular Malaysia with a key to and phylogenetic analysis of all genera of the subtribe. Insect Syst Evol 42:349–364CrossRefGoogle Scholar
  52. Kim HJ, Ahn KJ (2000) A taxonomic review of the genus Tachinus Gravenhorst in Korea (Insecta: Coleoptera: Staphylinidae). Korean J Env Biol 18:125–131Google Scholar
  53. Kishimoto T (2000) A new Trigonurus (Coleoptera: Staphylinidae, Trigonurinae) discovered in Sichuan, China. Elytra 28:305–309Google Scholar
  54. Klimaszewski J (1984) A revision of the genus Aleochara Gravenhorst of America north of Mexico (Coleoptera: Staphylinidae, Aleocharinae). Mem Ent Soc Can 129:1–211Google Scholar
  55. Kocian M (1996) A revision of Western Palearctic species of the genus Ischnosoma Stephens (Coleoptera: Staphylinidae, Tachyporinae). Acta Univ Carol Biol 40:241–299Google Scholar
  56. Lawrence JF, Britton EB (1994) Australian beetles. Melbourne University Press, CarltonGoogle Scholar
  57. Leschen RAB, Löbl I (2005) Phylogeny and classification of Scaphisomatini (Staphylinidae: Scaphidiinae) with notes on mycophagy, termitophyly, and functional morphology. Col Soc Monogr 3:1–63Google Scholar
  58. Löbl I (2002) New species of Scaphisoma Leach (Coleoptera: Staphylinidae: Scaphidiinae) from Mt. Wilhelm, Papua New Guinea. Acta Zool Hung 48:181–189Google Scholar
  59. Löbl I (2012) On Taiwanese species of Baeocera Erichson (Coleoptera: Staphylinidae: Scaphidiinae). Zool Stud 51:118–130Google Scholar
  60. Löbl I (2015) On the Scaphidiinae (Coleoptera: Staphylinidae) of the Lesser Sunda Islands. Rev Suisse Zool 122:75–112Google Scholar
  61. Löbl I, Burckhardt D (1988) Cerapeplus gen. n. and the classification of micropeplids (Coleoptera: Micropeplidae). Syst Ent 13:57–66CrossRefGoogle Scholar
  62. Löbl I, Calame FG (1996) Taxonomy and phylogeny of the Dasycerinae (Coleoptera: Staphylinidae). J Nat Hist 30:247–291CrossRefGoogle Scholar
  63. Löbl I, Mattila J (2010) Euplectus lapponicus (Coleoptera: Staphylinidae: Pselaphinae), a new species from boreal Finland. Ent Fenn 21:181–186Google Scholar
  64. Makranczy G, Schülke M (2001) Typenstudien an den mitteleuropäischen Vertreten der Artengruppe des Thinobius linearis Kraatz, 1857 (Coleoptera, Staphylinidae, Oxytelinae). Ent Blät 97:185–193Google Scholar
  65. Maruyama M (2000) A revision of the myrmecophilous genus Aspidobactrus (Coleoptera: Staphylinidae: Aleocharinae). Sociobiology 35:149–173Google Scholar
  66. Maruyama M (2004a) Redescription of the genus Creochara (Coleoptera: Staphylinidae: Aleocharinae: Aleocharini) and its systematic position. Can Ent 136:621–637CrossRefGoogle Scholar
  67. Maruyama M (2004b) Taxonomy of the myrmecophilous genus Myrmecopella Kistner with descriptions of seven new species (Coleoptera: Staphylinidae: Aleocharinae). Kol Rund 74:201–219Google Scholar
  68. Maruyama M (2006) Revision of the Palearctic species of the myrmecophilous genus Pella (Coleoptera: Staphylinidae: Aleocharinae). Nat Sci Mus Monogr Tokyo 32:1–207Google Scholar
  69. Maruyama M (2008) Giraffaenictus eguchii (Coleoptera, Staphylinidae, Aleocharinae), a new genus and species of fully myrmecophile from a colony of Aenictus binghami (Hymenoptera, Formicidae, Aenictinae) in Vietnam. Esakia 48:51–56Google Scholar
  70. Maruyama M, Hlavác P (2003) A new genus and species of myrmecophilous Aleocharini from Morocco (Coleoptera, Staphylinidae, Aleocharinae), associated with Aphaenogaster (Hymenoptera: Formicidae: Myrmicinae). Ann Soc ent Fr (new ser) 39:225–228CrossRefGoogle Scholar
  71. Maruyama M, Klimaszewski J (2004) A new species of the myrmecophilous genus Goniusa (Coleoptera, Staphylinidae, Aleocharinae) from Canada. Elytra 32:315–320Google Scholar
  72. Maruyama M, Matsumoto T, Itioka T (2011) Rove beetles (Coleoptera: Staphylinidae) associated with Aenictus laeviceps (Hymenoptera: Formicidae) in Sarawak: strict host specificity, and first myrmecoid Aleocharini. Zootaxa 3102:1–26Google Scholar
  73. Maruyama M, Sugaya H (2002) A new species of Tetrasticta (Coleoptera, Staphylinidae, Aleocharinae) from Japan and Taiwan. Jpn J Syst Ent 8:17–21Google Scholar
  74. Matsumura Y, Yoshizawa K (2012) Homology of the internal sac components in the leaf beetle subfamily Criocerinae and evolutionary novelties related to the extremely elongated flagellum. J Morph 273:507–518PubMedCrossRefGoogle Scholar
  75. Mckenna DD, Farrell BD, Caterino MS, Farnum CW, Hawks DC, Maddison DR, Seago AE, Short AEZ, Newton AF, Thayer MK (2015) Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Syst Ent 40:35–60CrossRefGoogle Scholar
  76. Naomi S (1986) Taxonomic study on the subfamily Osoriinae (Coleoptera, Oxytelidae) from Japan, I. Elytra 14:33–42Google Scholar
  77. Naomi S (1990) Comparative morphology of the Staphylinidae and the allied groups (Coleoptera, Staphylinoidea) XI. Abdominal glands, male genitalia and female spermatheca. Jpn J Ent 58:16–23Google Scholar
  78. Naomi S (1996) Revision of the genus Lispinus Erichson and Neolosus Blackwelder (Coleoptera: Staphylinidae) from Japan. Jpn J Ent 64:762–771Google Scholar
  79. Naomi S (1997a) A revision of the genus Nacaeus Blackwelder (Coleoptera: Staphylinidae) from Japan. Jpn J Ent 65:127–142Google Scholar
  80. Naomi S (1997b) Taxonomic study on Stenus wasabi Hromádka and its allied species (Coleoptera; Staphylinidae; Steninae), with discussion on the internal structure of the aedeagus. Jpn J Ent 65:600–611Google Scholar
  81. Naomi S (2006a) Taxonomic revision of the genus Stenus Latreille, 1797 (Coleoptera, Staphylinidae, Steninae) of Japan: species group of S. indubius Sharp. Jpn J Syst Ent 12:39–120Google Scholar
  82. Naomi S (2006b) Taxonomic revision of the genus Stenus Latreille, 1797 (Coleoptera, Staphylinidae, Steninae) of Japan: species group of S. (Hypostenus) rufescens Sharp. Nat Hist Res Chiba Spec Issue 9:1–81Google Scholar
  83. Naomi S (2010) Descriptions of two new species of the genus Stenus Latreille (Coleoptera: Staphylinidae: Steninae) from Chubu District, Central Japan. Nat Hist Res Chiba 11:35–40Google Scholar
  84. Naomi S, Irmler U (2012) First record of the genus Allotrochus Fagel, 1955 (Coleoptera: Staphylinidae: Osoriinae) from the Oriental region, with description of the new species A. takahashii from Ishigaki Is., Nansei Islands, Japan. Biogeography 14:105–110Google Scholar
  85. Naomi S, Maruyama M (1997) Revision of the genus Sepedophilus Gistel (Coleoptera, Staphylinidae, Tachyporinae) from Japan: species group of S. glabratus and S. exiguus. Ent Rev Jpn 52:61–71Google Scholar
  86. Naomi S, Maruyama M (1998) A revision of the genus Sepedophilus Gistel (Coleoptera, Staphylinidae, Tachyporinae) from Japan: species group of S. pedicularius. Jpn J Syst Ent 4:51–75Google Scholar
  87. Naomi SI, Nomura S, Puthz V (2017) The subfamily Steninae Macleay, 1825 (Coleoptera: Staphylinidae) of Japan. Part 1. Dianous and Stenus (S. comma group to S. guttalis group). Nat Sci Mus Monogr Tokyo 46:1–339Google Scholar
  88. Nichols SW (1989) The Torre-Bueno glossary of entomology. New York Entomological Society, New YorkGoogle Scholar
  89. Nomura S (1991) Systematic study on the genus Batrisoplisus and its allied genera from Japan (Coleoptera, Pselaphidae). Esakia 30:1–462Google Scholar
  90. Nomura S (2001a) A taxonomic revision of the Japanese species of the genus Pselaphogenius (Coleoptera, Staphylinidae, Pselaphinae). Part 3, Species from central Honshu. Mem Natn Sci Mus Tokyo 37:279–291Google Scholar
  91. Nomura S (2001b) Descriptions of two new species of the Clavigerine genus Articerodes (Coleoptera, Staphylinidae, Pselaphinae) from the Ogasawara Islans, Japan. Elytra 29:343–351Google Scholar
  92. Nomura S (2010) A taxonomic revision of Asian genera and species of the tribe Dimerini (Coleoptera, Staphylinidae, Pselaphinae). Jpn J syst Ent 16:221–247Google Scholar
  93. Nomura S, Sakcoowong W, Chanpaisaeng J (2011) Three new species of the genus Megatyrus (Coleoptera, Staphylinidae, Pselaphinae) from Thailand. Spec Publ Jpn Soc Scarab 1:121–129Google Scholar
  94. Ogawa R, Löbl I (2016) A review of the genus Xotidium Löbl, 1992 (Coleoptera, Staphylinidae, Scaphidiinae), with descriptions of five new species. Dt Ent Zeit 63:155–169CrossRefGoogle Scholar
  95. Pace R (1999) Nuove specie del genere Leptusa Kraatz raccolte da Manfred Kahlen. Veröff Tiroler Land Ferd 79:207–214Google Scholar
  96. Pace R (2009) Aleocharinae del cile raccolte dal Dr. P. M.Giachino (Coleoptera, Staphylinidae). Lavori Soc Ven Sc Nat 34:25–32Google Scholar
  97. Pace R (2010) Thamiaraeini, Lomechusini, Oxypodini, Hoplandriini e Aleocharini di Taiwan (Coleoptera, Staphylinidae). Boll Mus Civ Stor Nat Verona 34:19–54Google Scholar
  98. Peschke K (1978) Funktionsmorphologische Untersuchungen zur Kopulation von Aleochara curtula Goeze (Coleoptera, Staphylinidae). Zoomorphologie 89:157–184CrossRefGoogle Scholar
  99. Puthz V (1973) On some Neotropical Euaesthetinae (Coleoptera, Staphylinidae). Stud Neotr Fauna 8:51–73CrossRefGoogle Scholar
  100. Puthz V (1974) Bemerkungen über die Gattung Edaphus Motschulsky und über Kistnerische Edaphus-Arten nebst neuen und alten afrikanischen Euaesthetinen. Rev Zool Afr 88:749–770Google Scholar
  101. Puthz V (1976) Zwei neue Gattungssynonyme und ein Nomen novum bei den Euaesthetinen (Coleoptera: Staphylinidae). Ent Scand 7:239–240CrossRefGoogle Scholar
  102. Puthz V (1977) Die Gattung Octavius Fauvel (Coleoptera: Staphylinidae) weltweit verbreitet! Opusc Zool Budap 14:105–126Google Scholar
  103. Puthz V (1978) Bemerkungen über Schatzmayrina Koch (Coleoptera, Staphylinidae). Ent Blät 74:47–51Google Scholar
  104. Puthz V (1979) Eine neue Euaesthetinengattung aus dem Himalaya (Coleoptera, Staphylinidae). Reichenbachia 17:9–14Google Scholar
  105. Puthz V (1985a) Neue und alte neotropische Edaphus-Arten aus dem Genfer Museum (Coleoptera, Staphylinidae). Rev Suisse Zool 92:351–357CrossRefGoogle Scholar
  106. Puthz V (1985b) Neue Octavius-Arten aus Nordindien und aus dem Himalaya-Gebiet (Coleoptera: Staphylinidae). Revue suisse Zool 92:697–705CrossRefGoogle Scholar
  107. Puthz V (1986a) Eine neue Art der Gattung Edaphosoma Scheerpeltz. Reichenbachia 23:135–138Google Scholar
  108. Puthz V (1986b) Fünf neue Edaphus-Arten aus Afrika (Staphylinidae, Coleoptera). Philippia 5:311–317Google Scholar
  109. Puthz V (1986c) Neue Arten der Gattung Octavius Fauvel, 1872, aus Südafrika nebst einer Bestimmungstabelle der in der Äthiopis lebenden Gattungsvertreter (Coleoptera: Staphylinidae). Ann Trans Mus 34:185–201Google Scholar
  110. Puthz V (1988a) Three new Stenaesthetus species from Ethiopia (Insecta, Coleoptera: Staphylinidae). Reichenbachia 26:9–13Google Scholar
  111. Puthz V (1988b) Die Steninen von Borneo II (Coleoptera, Staphylinidae). Neue und alte Arten, vorwiegend aus den Ausbeuten des Genfer Naturhistorischen Museums. Rev Suisse Zool 95:631–679CrossRefGoogle Scholar
  112. Puthz V (1989a) Zwei neue Euaesthetinen von der Elfenbeinküste (Coleoptera, Staphylinidae). Ent Blät 85:75–77Google Scholar
  113. Puthz V (1989b) The male of Stictocranius mariae (Hatch) (Coleoptera, Staphylinidae). Ent Blät 85:165–166Google Scholar
  114. Puthz V (1989c) Eine neue Octavius -Art aus Sumatra (Coleoptera, Staphylinidae). Rev Suisse Zool 96:115–116CrossRefGoogle Scholar
  115. Puthz V (1992) Neue und alte afrikanische Edaphus-Arten (Insecta, Coleoptera, Staphylinidae: Euaesthetinae). Ent Abhand 55:13–33Google Scholar
  116. Puthz V (1998) Neuweltliche Euaesthetus-Arten (Staphylinidae, Coleoptera). Phillipia 8:223–244Google Scholar
  117. Puthz V (2000a) The genus Dianous Leach in China (Coleoptera, Staphylinidae). Rev Suisse Zool 107:419–559CrossRefGoogle Scholar
  118. Puthz V (2000b) Neue und alte neotropische Arten der Gattung Stenus Latreille (Staphylinidae, Coleoptera). Philippia 9:165–213Google Scholar
  119. Puthz V (2000c) Aptere Stenus-Arten aus den mexikanischen Hochgebirgen (Coleoptera: Staphylinidae). Mitt internat-entomol Ver 25:103–127Google Scholar
  120. Puthz V (2002) Revision der Gattung Tamotus Schaufuss, 1872 (Coleoptera, Staphylinidae). Z Arb Gem Öst Ent 54:115–125Google Scholar
  121. Puthz V (2003a) Revision of the genus Fenderia Hatch, 1957 (Coleoptera, Staphylinidae). In: Cuccodoro G, Leschen RA (eds) Systematics of Coleoptera: Papers celebrating the retirement of Ivan Löbl. Memoirs on Entomology International 17:637–648Google Scholar
  122. Puthz V (2003b) Neue und alte Arten der Gattung Stenus Latreille aus China (Insecta: Coleoptera: Staphylinidae: Steninae). Ent Abhand 60:139–159Google Scholar
  123. Puthz V (2008a) Stenus Latreille und die segenreiche Himmelstochter (Coleoptera, Staphylinidae). Linzer biol Beitr 40:137–230Google Scholar
  124. Puthz V (2008b) Kiwiaesthetus, a new genus of Euaesthetinae from New Zealand (Coleoptera: Staphylinidae). Z Arb Gem Öst Ent 60:59–69Google Scholar
  125. Puthz V (2010) Die Gattung Edaphosoma Scheerpeltz, 1976 in China (Coleoptera, Staphylinidae). Ent Blät 106:289–306Google Scholar
  126. Puthz V (2011a) Afrikanische Stenaesthetus-Arten (Coleoptera: Euaesthetinae). Linzer biol Beitr 43:793–812Google Scholar
  127. Puthz V (2011b) Neue und alte Euaesthetinen (Coleoptera: Staphylinidae). Z Arb Gem Öst Ent 63:13–31Google Scholar
  128. Puthz V (2012a) Über die neuweltlichen Megalopinus-Arten (Coleoptera: Staphylinidae). Linzer biol Beitr 44:613–834Google Scholar
  129. Puthz V (2012b) Zwei neue Arten der Gattung Nothoesthetus Saíz aus Chile (Coleoptera: Staphylinidae). Mitt int ent Ver Frankfurt a M 37:159–162Google Scholar
  130. Puthz V (2012c) Über die Megalopinus-Arten der Orientalis (Coleoptera: Staphylinidae). Linzer biol Beitr 44:1373–1430Google Scholar
  131. Puthz V (2013) Übersicht über die orientalischen Arten der Gattung Stenus Latreille 1797 (Coleoptera, Staphylinidae). Linzer biol Beitr 45:1279–1470Google Scholar
  132. Puthz V (2014) Nordamerikanische Arten der Gattung Euaesthetus Gravenhorst (Coleoptera, Staphylinidae). Linzer biol Beitr 46:845–876Google Scholar
  133. Sharp D, Muir F (1912) The comparative anatomy of the male genital tube in Coleoptera. Trans Ent Soc Lond 1912:477–642Google Scholar
  134. Shavrin A (2012) On the taxonomic placement of Geodromicus pusillus Coiffait, 1983 (Coleoptera: Staphylinidae: Omaliinae). Zootaxa 3552:66–68Google Scholar
  135. Shimada T (2002) A new Aploderus (Coleoptera, Staphylinidae, Oxytelinae) discovered in central Japan. Spec Bull Jpn Soc Coleopt 5:233–239Google Scholar
  136. Smetana A (1973) Die Leptusa-Arten der Tschechoslovakischen Republik einschließlich Karpatorußlands (Col., Staphylinidae). Stutt Beitr Naturk Ser A Biol 255:1–46Google Scholar
  137. Snodgrass RE (1935) Principles of insect morphology. McGraw-Hill, New YorkGoogle Scholar
  138. Steel WO (1964) Insects of Campbell Island. Coleoptera: Staphylinidae. Pac Ins Monogr 7:340–375Google Scholar
  139. Steel WO (1966) A revision of the staphylinid subfamily Proteininae (Coleoptera) I. Trans R Ent Soc Lond 118:285–311CrossRefGoogle Scholar
  140. Thayer MK (1987) Biology and phylogenetic relationships of Neophonus bruchi, an anomalous south Andean staphylinid (Coleoptera). Syst Ent 12:389–404CrossRefGoogle Scholar
  141. Thayer MK (1997) Proglypholoma aenigma gen. et sp. nov., Glypholoma spp. nov. and new records, and a phylogenetic analysis of Glypholomatinae (Coleoptera: Staphylinidae). Ann Zool Warszawa 47:157–174Google Scholar
  142. Thayer MK, Newton AF Jr (1978) Revision of the south temperate genus Glypholoma Jeannel, with four new species (Coleoptera: Staphylinidae: Omaliinae). Psyche 85:25–63CrossRefGoogle Scholar
  143. Williams SA (1976) The genus Oligota (Coleoptera, Staphylinidae) in New Zealand. New Zeal J Zool 3:247–255CrossRefGoogle Scholar
  144. Williams SA (1978) The genus Oligota Mannerheim (Col., Staphylinidae) in the Ethiopian region. Ent Mon Mag 114:177–190Google Scholar
  145. Wunderle P, Assing V (2000) A revision of the European species of Hygropetrophila Bernhauer (Coleoptera, Staphylinidae, Aleocharinae). Ent Blät 96:31–40Google Scholar
  146. Yamamoto S, Maruyama M (2012) Revision of the seashore-dwelling subgenera Emplenota Casey and Triochara Bernhauer (Coleoptera: Staphylinidae: genus Aleochara) from Japan. Zootaxa 3517:1–52Google Scholar
  147. Zanetti A (2014) Taxonomic revision of North American Eusphalerum Kraatz, 1857 (Coleoptera, Staphylinidae, Omaliinae). Ins Mundi 379:1–80Google Scholar

Copyright information

© Crown 2018

Authors and Affiliations

  1. 1.Natural History Museum and InstituteChibaJapan

Personalised recommendations