Contents

Editors’ preface to the 2nd edition of Volume 1 vii Rolf G. Beutel & Sergio A. Vanin Editors’ preface to the 1st edition of Volume 1 ix 6.2 Steffan, 1964 67 Contributors xv Rolf G. Beutel & Roberto Arce-Pérez 6.3 Sphaeriusidae Erichson, 1845 Rolf G. Beutel & Duane D. McKenna (Jäch 1999) (= Microsporidae) 70 1 Systematic position, basal branching pattern Sergio A. Vanin, Rolf G. Beutel & and early evolution 1 Roberto Arce-Pérez 6.4 Hydroscaphidae LeConte, John F. Lawrence 1874 71 2 Classification (families and subfamilies) 13

Rolf G. Beutel & Ignacio Ribera Duane D. McKenna 7 Schellenberg, 1806 77 3 Molecular systematics of Coleoptera 23 Rolf G. Beutel & Robert E. Roughley† 7.1 Gyrinidae Latreille, 1810 80 Rolf G. Beutel & John F. Lawrence Bernhard J. van Vondel 4 Coleoptera, morphology 35 7.2 Aubé, 1836 89

Konrad Dettner 5 Kolbe, 1908 41 7.3 Thomson, 1857 98 Thomas Hörnschemeyer Konrad Dettner Introduction and phylogeny 41 7.4 Amphizoidae LeConte, 1853 107 Thomas Hörnschemeyer Konrad Dettner 5.1 Crowsoniellidae Iablokoff- 7.5 Hygrobiidae Régimbart, 1879 112 Khnzorian, 1983 43 Michael Balke & Lars Hendrich Thomas Hörnschemeyer & Margarita Yavorskaya 7.6 Leach, 1815 118 5.2 Laporte, 1836 44 Rolf G. Beutel, Michael Balke & Ignacio Ribera Thomas Hörnschemeyer 7.7 Aspidytidae Ribera, Beutel, Balke & Vogler, 5.3 Micromalthidae Barber, 1913 48 2002 141 Thomas Hörnschemeyer & Rolf Georg Beutel Michael Balke, Yves Alarie & 5.4 Ommatidae Sharp & Muir, 1912 52 Rolf Georg Beutel Thomas Hörnschemeyer 7.8 Meruidae Spangler & Steiner, 5.5 Jurodidae Kirejtshuk, 2005 149 1999 (= Sikhotealiniidae) 57 Rolf G. Beutel & Erik Arndt 7.9  C. G. Thomson, 6 Crowson, 1955 63 1857 159

Rolf G. Beutel Erik Arndt, Rolf G. Beutel & Kipling Will Introduction, phylogeny 63 7.10 Carabidae Latreille, 1802 162 Roberto Arce-Pérez, José Luis Navarrete-Heredia & Rolf G. Beutel Rolf G. Beutel 6.1 Lepiceridae Hinton, 1936 7.11 Family Laporte, (= Cyathoceridae) 65 1840 (= Rhysodini s. Bell) 190 xii Contents

Rolf G. Beutel & Richard A. B. Leschen Alfred F. Newton 8 Emery, 1886 199 13.2 Synteliidae Lewis, 1882 277

Introduction, phylogeny 199 Peter W. Kovarik & Michael S. Caterino 13.3 Gyllenhal, 1808 281 Ladislav Bocak

9 Scirtiformia Fleming, 1821 201 14 Latreille, 1802 315 Introduction, phylogeny 201 Alfred F. Newton Introduction and phylogeny 315 10 Fleming, 1821 203 Manfred A. Jäch, Rolf G. Beutel, Juan A. Delgado & John F. Lawrence Juan A. Díaz 10.1 Decliniidae Nikitsky, Lawrence, 14.1 Mulsant, 1844 316 Kirejtshuk & Gratshev, 1994 203 W. Eugene Hall Richard A. B. Leschen 14.2 Erichson, 1845 345 10.2 Lacordaire, Alfred F. Newton 1857 206 14.3 Thomson, 1859 356 Richard A. B. Leschen Alfred F. Newton 10.3 Fischer von Waldheim, 14.4 Fleming, 1821 364 1821 211 Paweł Jałoszyński John F. Lawrence 14.5 Leach, 1815 376 10.4 Fleming, 1821 215 Derek S. Sikes 14.6 Latreille, 1807 386 Rolf G. Beutel & Richard A. B. Leschen 11 Lameere, Margaret K. Thayer 1900 227 14.7 Staphylinidae Latreille, 1802 394 Introduction, phylogeny 227 Clarke H. Scholtz & Vasily V. Grebennikov

Miguel Archangelsky, Rolf Georg Beutel & 15 Latreille, 1802 443

Albrecht Komarek Introduction 443

12 Latreille, 1802 231 15.1 Lucanidae Latreille, 1802 465

Introduction 231 15.2 Leach, 1815 468

12.1 Latreille, 1802 238 15.3 MacLeay, 1819 470 12.2 Helophoridae Leach, 1815 254 15.4 Glaresid ae Semenov-Tian-Shanskij &

12.3 Epimetopidae Zaitzev, 1908 257 Medvedev, 1932 471

12.4 Georissidae Laporte, 1840 259 15.5 Pleocomidae LeConte, 1861 472

12.5 Hydrochidae C.G. Thomson, 1859 262 15.6 Bolboceratid ae Mulsant, 1842 473

12.6 Spercheidae Erichson, 1837 264 15.7 Diphyllostom atidae Holloway, 1972 475 15.8 Latreille, 1802 476 15.9 Belohinidae Paulian, 1959 477

13 Histeroidea Gyllenhal, 1808 273 15.10 Mulsant & Rey, 1871 478

Rolf G. Beutel 15.11 Erichson, 1847 480 Introduction, phylogeny 273 15.11.1 Ceratocanthinae Cartwright & Gordon, 1971 480 Alfred F. Newton 15.11.2 Hybosorinae Erichson, 1847 483

13.1 Sphaeritidae Shuckard, 1839 274 15.12 MacLeay, 1819 484 Contents xiii

15.13 Family 486 Crystal A. Maier, John F. Lawrence & 15.13.1 Aphodiinae Leach, 1815 487 Richard A. B. Leschen 15.13.2 Subfamily Scarabaeinae Latreille, 19.1 Latreille, 1804 554 1802 490 Ján Kodada, Manfred A. Jäch & Fedor Čiampor Jr. 15.13.3 Pachypodinae Erichson, 1840 494 19.2 Curtis, 1830 561 15.13.4 Orphninae Erichson, 1847 494 15.13.5 Allidiostomatinae Arrow, Ján Kodada, Manfred A. Jäch & Fedor Čiampor Jr. 1940 495 19.3 Bilberg, 1820 15.13.6 Dynamopodinae Arrow, 1911 496 (1817) 590 15.13.7 Aclopinae Milne-Edwards, 1850 496 Sergio Ide, Cleide Costa & Sergio Antonio Vanin 15.13.8 Subfamily Euchirinae Hope, 19.4 Lutrochidae Kasap & Crowson, 1840 496 1975 602 15.13.9 Phaenomeridinae Erichson, 1847 497 15.13.10 MacLeay, 1819 497 Carles Hernando & Ignacio Ribera 15.13.11 Rutelinae MacLeay, 1819 499 19.5 Limnichidae Erichson, 1846 605 15.13.12 Dynastinae MacLeay, 1819 501 Sergio A. Vanin, Cleide Costa, Sergio Ide & 15.13.13 Cetoniinae Leach, 1815 Rolf G. Beutel (excluding Valginae) 503 19.6 MacLeay, 1825 612 15.13.14 Valginae Mulsant, 1842 504 Chi-Feng Lee, Manfred A. Jäch & Rolf G. Beutel 19.7 Psephenidae Lacordaire, Rolf G. Beutel, Richard A. B. Leschen, Ladislav Bocak & 1854 615 Milada Bocakova 16 Crowson, 1960 527 Cleide Costa, Sergio Antonio Vanin & Sergio Ide Introduction, phylogeny 527 19.8 Cneoglossidae Guérin-Méneville, 1843 629

John F. Lawrence John F. Lawrence 17 Guérin-Méneville, 1843 531 19.9 Laporte, 17.1 Dascillidae Guérin-Méneville, 1843 531 1836 632 17.2 Rhipiceridae Latreille, 1834 537 Rolf G. Beutel & Richard A. B. Leschen 19.10 Blanchard, 1845 639 Charles L. Bellamy† & Mark G. Volkovitsh Michael A. Ivie 18 Crowson, 1955 543 19.11 Eulichadidae Crowson, 1973 644 18.1 Leach, 1815 543 John F. Lawrence 19 Latreille, 1804 553 19.12 Callirhipidae Emden, 1924 648

Rolf G. Beutel Introduction, phylogeny 553 Taxonomic Index 655