Families and subfamilies of Coleoptera

(with selected genera, notes, references

and data on family-group names)

by J. F. Lawrence and A. F. Newton, Jr.

Division of Entomology, CSIRO, G.P.O. Box 1700, Canberra, ACT 2601,

Australia
Division of Insects, Field Museum of Natural History, Roosevelt Road at Lake
Shore Drive, Chicago, IL 60605, U.S.A.

Reprinted from:

Biology, Phylogeny, and Classification of Coleoptera

Papers Celebrating the 80th Birthday of Roy A. Crowson

Eds. J. Pakaluk and S.A. Slipinski

1995, Muzeum i Instytut Zoologii PAN, Warszawa

ISBN 83-85192-34-4
Families and subfamilies of Coleoptera
(with selected genera, notes, references
and data on family-group names)

by J. F. Lawrence and A. F. Newton, Jr.

Division of Entomology, CSIRO, G.P.O. Box 1700, Canberra, ACT 2601,

Australia
Division of Insects, Field Museum of Natural History, Roosevelt Road at Lake
Shore Drive, Chicago, IL 60605, U.S.A.

Reprinted from:

Biology, Phylogeny, and Classification of Coleoptera

Papers Celebrating the 80th Birthday of Roy A. Crowson

Eds. J. Pakaluk and S.A. Slipinski

1995, Muzeum i Instytut Zoologii PAN, Warszawa

ISBN 83-85192-34-4
Families and subfamilies of Coleoptera
(with selected genera, notes, references
and data on family-group names)

J. F. LAWRENCE
A. F. NEWTON, Jr.

Abstract. A complete, current, and annotated classification of Recent Coleoptera at

the subfamily level and above is presented, with complete or selected lists of genera
included in each group. Brief discussions summarize recent changes in composition
and placement of these higher groups, with extensive reference to studies providing
evidence for these changes as well as other recent systematic works and catalogs of
each group. The classification, summarized in a table, lists 166 families and 453
subfamilies placed in 4 suborders, the largest of which (Polyphaga) is divided into
5 series and 16 superfamilies. About 3300 representative genera are listed, some with
geographic distributions. No new taxa or names are proposed, but a few changes in
composition or rank of taxa are made.
The names of all recognized subfamilies, families and higher taxa were verified
and fully referenced. Name changes from common usage required by the current
Zoological Code are summarized in a table, with indication of those changes imple-
mented recently or here. Other nomenclatural problems are highlighted for further
study or possible conservation of well-known names.

Introduction

Although a general review of beetle classification was published by Lawrence

and Newton (1982), there are few complete lists of beetle families worldwide
published since the appearance of Crowson's major work (1955), the latest
ones probably being those of Crowson (1981), Lawrence (1982, 1991), Law-
rence and Britton (1991, 1994), and Paulian (1988); the last comprehensive list
including subfamilies is the much older Junk and Schenkling Coleopterorum
Catalogus, completed in 1940. Furthermore, there are no recent works dealing

1. Pakaluk and S.A. Slipiliski (eds.): Biology, Phylogeny, and Classification of Coleoptera:
Papers Celebrating the 80th Birthday of Roy A. Crowson.
© 1995, Muzeum i lnstytut Zoologii PAN, Warszawa. ISBN 83-85192-34-4.
780
with family limits and changes, except for brief family sections given in Law-
rence (1982) and the comments on relationships included in most Coleoptera
family treatments in Stehr's (1991) Immature Insects (Vol. 2). Finally, t?~re
has been no attempt to review and document the names of all beetle famthes
and subfamilies since the introduction of formal rules governing such names
into the International Code of Zoological Nomenclature (ICZN 1961).
The present work seeks to rectify this situation by attempting to give a com-
plete and up-to-date list of beetle suborders, series, superfamilies, families and
subfamilies, including common synonyms; partial or complete lists of included
genera, some with geographic distributions; notes on phylogenetic hyp~theses
and alternative schemes of classification; and lists of selected references mclud-
ing recent broad taxonomic reviews and catalogs. Also provided are complete
references for all recognized family and subfamily names and some wtdely
used synonyms, with discussions of nomenclatural problems at this level in-
cluding required name changes. Only those higher taxa containing Recent rep-
resentatives are included, but some extinct families are mentioned or discussed
in connection with certain suborders, series or superfamilies. An extensive list
of extinct families and subfamilies of Coleoptera is provided by Ponomarenko
(1995).
The depth of the bibliographic coverage varies from family to family, being
more extensive in some of the smaller and less well-known groups. It is by no
means complete anywhere, but is intended as a starting point for each group.
In general, major taxonomic papers on the taxon, and those with important
phylogenetic information are emphasized, but some biological papers are in-
cluded as well as faunal works with a broad coverage of taxa, and catalogs
more recent than the Coleopterorum Catalogus. Literature known to us through
mid-1994 is included.
In the classification used here (see synopsis in Table 1), higher taxa are
listed in a sequence inferred to be from less to more derived, with closely re-
lated taxa placed next to one another as far as possible. In general, phylo-
genetic relationships among beetle higher taxa are too inadequately known to
justify the use of a formal sequencing convention (e.g., Wiley 1:)79) that woul.d
relate the classification directly to a cladogram. Genera are hsted alphabeti-
cally; distributions are given for genera in the case of some small and geo-
graphically restricted groups, but the absence of distribution information does
not necessarily imply a worldwide distribution for a genus or higher group.
Several families, subfamilies, and genera of especially unclear relationships are
listed as incertae sedis at the end of the most appropriate series or family.
Although an attempt has been made to present some alternative views on the
classifications of various groups, the ideas presented and the classifications
chosen are those of the authors. JFL was responsible for the overall classifica-
tion and for comments dealing with phylogeny and classification, except for
781
Staphyliniformia (AFN); AFN was responsible for nomenclatural comments
and decisions.
General references on Coleoptera
These selected, mostly recent, works include taxonomically broad or complete
coverage of Coleoptera from various perspectives, including major books, cata-
logs, faunal works, and comparative morphological and ecological studies that
are generally not cited again under individual families. Multi-authored, multi-
year publication series are indicated by title only and do not appear in the Ref-
erences; those that are only partially completed for Coleoptera (in most cases,
active publication series) are indicated by a dollar sign ($).
CLASSIFICATION: Coleopterorum Catalog us (191 0-1940; last complete world
catalog of Coleoptera; the few subsequent supplements are listed under the
relevant family or subfamily), Crowson (1955, 1960, 1981), Genera Insecta-
rum$ (world keys to genera; very incomplete for Coleoptera), Lawrence
(1982), Lawrence and Britton (1991, 1994; Australian emphasis but world
view), Lawrence and Newton (1982), Lawrence, Slipinski and Pakaluk (1995;
most recent and complete historical review), Lawrence et al. in Stehr (1991;
comments on classification and biology for most families), Les Coleopteres du
Monde$ (large-format guides to a few popular groups only), Paulian (1949,
1988).
NOMENCLATURE: see following section.
LARVAE: Bertrand (1972), Boving and Craighead (1931), Costa et al. (1988),
Kawada (1960), Klausnitzer (1978), Lawrence et al. (1993), Lawrence et al.
in Stehr (1991), Morimoto and Hayashi (1986), Peterson (1960).
MORPHOLOGY: Browne and Scholtz (1994), Caveney (1986), Crowson (1938,
1944, 1981, 1984b), Dettner (1987), Forbes (1926), Hlavac (1973, 1975), Iablo-
koff-Khnzorian (1974, 1980), Jeannel (1955a), Jeanne} and Paulian (1944),
Kukalova-Peck and Lawrence (1993), Paulian (1988), Sharp and Muir (1912),
Smith and Virkki (1978), Stork (1980).
BIOLOGY AND ECOLOGY: Coiffait (1960), Crowson (1981, 1995), Evans
(1975), Kistner (1982), Lawrence et al. in Stehr (1991), Paulian (1988), Prins
(1984), Wheeler and Blackwell (1984), Wilding et al. (1989).
STORED PRODUCT PESTS: Booth et al. (1990), Bousquet (1990), Hinton
(1945), Lepesme (1945) .
PALEONTOLOGY: Arnoldi et al. (1977), Carpenter (1992), Crowson (1967b,
1975, 1981), Elias (1994), Poinar (1992), Ponomarenko (1969, 1985, 1995),
Ponomarenko and Ryvkin (1990), Spahr (1981).
 783
782
Table 1. Continued.
Table 1. Synopsis of families and subfamilies of Coleoptera.
STAPHYLINIDAE Latreille, 1802 OCHODAEIDAE Mulsant & Rey, 1871
GLYPHOLOMATINAE Jeanne!, 1962 OCHODAEINAE Mulsant & Rey, 1871
Order COLEOPfERA Linnaeus, 1758
Suborder POLYPHAGA Emery, 1886 MICROSILPHINAE Crowson, 1950 CHAETOCANTHINAE Scholtz in Scholtz et al.,
OMALIINAE MacLeay, 1825 1988
Suborder ARCHOSTEM ATA Kolbe, 1908
Series STAPHYLINIFORMIA Lameere, 1900 EMPELINAE Newton & Thayer, 1992 CERATOCANTHIDAE Cartwright & Gordon, 1971
PROTEININAE Erichsen, 1839 HYBOSORIDAE Erichson, 1847
OMMATIDAE Sharp & Muir, 1912
Superfamily HYDROPHILOIDEA Latreille, 1802 MICROPEPLINAE Leach, 1815 GLAPHYRIDAE MacLeay, 1819
TETRAPHALERINAE Crowson, 1962
HYDROPHILIDAE Latreille, 1802 NEOPHONINAE Fauvel, 1905 SCARABAEIDAE Latreille, 1802
OMMATINAE Sharp & Muir, 1912
HELOPHORINAE Leach, 1815 DASYCERINAE Reitter, 1887 APHODIINAE Leach, 1815
CROWSONIELLIDAE lablokoff-Khnzorian, 1983
EPIMETOPINAE Zaitzev, 1908 PROTOPSELAPHINAE Newton & Thayer, 1995 SCARABAEINAE Latreille, 1802
MICROMALTHIDAE Barber, 1913
GEORISSINAE Laporte, 1840 PSELAPHINAE Latreille, 1802 PACHYPODINAE Erichson, 1840
CUPEDIDAE Laporte, 1836
HYDROCHINAE C.G. Thomson, 1859 PHLOEOCHARINAE Erichson, 1839 ORPHNINAE Erichson, 1847
SPERCHEINAE Erichson, 1837 OLISTHAERINAE C.G. Thomson, 1858 ALLIDIOSTOMATINAE Arrow, 1940
Suborder MYXOPHAGA Crowson, 1955
HORELOPHINAE Hansen, 1991 TACHYPORINAE MacLeay, 1825 DYNAMOPODINAE Arrow, 1911
HYDROPHILINAE Latreille, 1802 TRICHOPHYINAE C.G. Thomson, 1858 ACLOPINAE Milne-Edwards, 1850
LEPICERIDAE Hinton, 1936 (1882)
SPHAERIDIINAE Latreille, 1802 HABROCERINAE Mulsant & Rey, 1877 EUCHIRINAE Hope, 1840
TORRIDINCOLIDAE Steffan, 1964
SPHAERITIDAE Shuckard, 1839 ALEOCHARINAE Fleming, 1821 PHAENOMERIDINAE Erichson, 1847
HYDROSCAPHIDAE LeConte, 1874
SYNTELIIDAE Lewis, 1882 TRIGONURINAE Reiche, 1865 MELOLONTHINAE MacLeay, 1819 !Leach in
MICROSPORIDAE Crotch, 1873
HISTERIDAE Gyllenhal, 1808 APATETICINAE Fauvel, 1895 Samouelle, 1819
NIPONIINAE Fowler, 1912 SCAPHIDIINAE Latreille, 1807 RUTELINAE MacLeay, 1819
Suborder ADEPHAGA Schellenberg, 1806
ABRAEINAE MacLeay, 1819 PIESTINAE Erichson, 1839 DYNASTINAE MacLeay, 1819
TRYPETICINAE Bickhardt, 1914 OSORIINAE Erichson, 1839 CETONIINAE Leach, 1815
GYRINIDAE Latreille, 1810
TRYPANAEINAE Marseul, 1857 OXYTELINAE Fleming, 1821
SPANGLEROGYRINAE Folkerts, 1979
SAPRININAE Blanchard, 1845 OXYPORINAE Fleming, 1821 Series ELATERIFORMIA Crowson, 1960
GYRININAE Latreille, 1810
DENDROPHILINAE Reitter, 1909 MEGALOPSIDIINAE Leng, 1920
HALIPLIDAE Aube, 1836
ONTHOPHILINAE MacLeay, 1819 STENINAE MacLeay, 1825 Superfamily SCIRTOIDEA Fleming, 1821
TRACHYPACHIDAE C.G. Thomson, 1857
TRIBALINAE Bickhardt, 1914 EUAESTHETINAE C.G. Thomson, 1859 DECLINIIDAE Nikitsky et al., 1994
NOTERIDAE C.G. Thomson, 1860
HISTERINAE Gyllenhal, 1808 SOLIERIINAE Newton & Thayer, 1992 EUCINETIDAE Lacordaire, 1857
PHREATODYTINAE Ueno, 1957
HETAERIINAE Marseul, 1857 LEPTOTYPHLINAE Fauvel, 1874 CLAMBIDAE Fischer, 1821
NOTERINAE C.G. Thomson, 1860
CHLAMYDOPSINAE Bickhardt, 1914 PSEUDOPSINAE Ganglbauer, 1895 CALYPTOMERINAE Crowson, 1955
AMPHIZOIDAE LeConte, 1853
PAEDERINAE Fleming, 1821 ACALYPTOMERINAE Crowson, 1979
HYGROBIIDAE Regimbart, 1878 (1837)
Superfamily STAPHYLINOIDEA Latreille, 1802 STAPHYLININAE Latreille, 1802 CLAMBINAE Fischer, 1821
DYTISCIDAE Leach, 1815
HYDRAENIDAE Mulsant, 1844 SCIRTIDAE Fleming, 1821
COPELATINAE Van den Branden, 1885
PROSTHETOPINAE Perkins in Perkins & Series SCARABAEIFORMIA Crowson, 1960
LACCOPHILINAE Gistel, 1856
Balfour-Browne, 1994 Superfamily DASCILLOIDEA Guerin-Meneville,
HYDROPORINAE Aube, 1836
HYDRAENINAE Mu1sant, 1844 Superfamily SCARABAEOIDEA Latreille, 1802 1843 (1834)
COLYMBETINAE Erichsen, 1837
OCHTHEBIINAE C.G. Thomson, 1859 LUCANIDAE Latreille, 1804 DASCILLIDAE Guerin-Meneville, 1843 (1834)
DYTISCINAE Leach, 1815
PTILIIDAE Erichson, 1845/Motschu1sky, 1845 AESALINAE MacLeay, 1819 DASCILLINAE Guerin-Meneville, 1843 (1834)
AUBEHYDRINAE Guignot, 1942
PTILIINAE Erichson, 1845/Motschulsky, 1845 NICAGINAE LeConte, 1861 KARUMIINAE Escalera, 1913
RHYSODIDAE Laporte, 1840
NANOSELLINAE Barber, 1924 SYNDESINAE MacLeay, 1819 RHIPICERIDAE Latreille, 1834
CARABIDAE Latreille, 1802
CEPHALOPLECTINAE Sharp, 1883 LAMPRIMINAE MacLeay, 1819
PAUSSINAE Latreille, 1807
ACROTRICHINAE Reitter, 1909 (1856) PENICHROLUCANINAE Arrow, 1950 Superfamily BUPRESTOIDEA Leach, 1815
GEHRINGIINAE Darlington, 1933
AGYRTIDAE C.G. Thomson, 1859 LUCANINAE Latreille, 1804 BUPRESTIDAE Leach, 1815
OMOPHRONINAE Bonelli, 1810
LEIODIDAE Fleming, 1821 PASSALIDAE Leach, 1815 SCHIZOPODINAE LeConte, 1861
CARABINAE Latreille, 1802
CAMIARINAE Jeanne!, 1911 AULACOCYCLINAE Kaup, 1868 JULODINAE Lacordaire, 1857
CICINDELINAE Latreille, 1802
CATOPOCERINAE Hatch, 1927 (1880) PASSALINAE Leach, 1815 BUPRESTINAE Leach, 1815
HILETINAE Schitidte, 1847
LEIODINAE Fleming, 1821 TROGIDAE MacLeay, 1819 AGRILINAE Laporte, 1835
LORICERINAE Bonelli, 1810
COLONINAE Hom, 1880 (1859) GLARESIDAE Semenov-Ttan-Shanskii &
ELAPHRINAE Latreille, 1802
CHOLEVINAE Kirby, 1837 Medvedev, 1932 Superfamily BYRRHOIDEA Latreille, 1804
MIGADOPINAE Chaudoir, 1861
PLATYPSYLLINAE Ritsema, 1869 PLEOCOMIDAE LeConte, 1861 BYRRHIDAE Latreille, 1804
SIAGONINAE Bonelli, 1813
SCYDMAENIDAE Leach, 1815 DIPHYLLOSTOMATIDAE Holloway, 1972 BYRRHINAE Latreille, 1804
SCARITINAE Bonelli, 1810
MASTIGINAE Fleming, 1821 GEOTRUPIDAE Latreille, 1802 SYNCALYPTINAE Mulsant & Rey, 1869
TRECHINAE Bonelli, 1810
SCYDMAENINAE Leach, 1815 BOLBOCERATINAE Mulsant, 1842 AMPHICYRTINAE LeConte, 1861
HARPALINAE Bonelli, 1810
SILPHIDAE Latreille, 1807 GEOTRUPINAE Latreille, 1802 ELMIDAE Curtis, 1830
lncertae sedis:
SILPHINAE Latreille, 1807 LETHRINAE Mulsant & Rey, 1871 LARAINAE LeConte, 1861
PSEUDOMORPHINAE Newman, 1842
NICROPHORINAE Kirby, 1837 BELOHINIDAE Paulian, 1959 ELMINAE Curtis, 1830
BRACHININAE Bonelli, 1810
784
Table I. Continued.
DRYOPIDAE Billberg, 1820 (1817)
LUTROCHIDAE Kasap & Crowson, 1975
LIMNICHIDAE Erichson, 1846
HYPHALINAE Britton, 1971
LIMNICHINAE Erichson, 1846
CEPHALOBYRRHINAE Champion, 1925
THAUMASTODINAE Champion, 1924
HETEROCERIDAE MacLeay, 1825
ELYTHOMERINAE Pacheco, 1964
HETEROCERINAE MacLeay, 1825
PSEPHENIDAE Lacordaire, 1854
EUBRIINAE Lacordaire, 1857
PSEPHENOIDINAE Hinton, 1939
EUBRIANACINAE Jacobson, 1913 (1880)
PSEPHENINAE Lacordaire, 1854
CNEOGLOSSIDAE Champion, 1897
PTILODACTYLIDAE Laporte, 1836
ANCHYTARSINAE Champion, 1897
CLADOTOMINAE Pic, 1914
APLOGLOSSINAE Champion, 1897
ARAEOPIDIINAE Lawrence, 1991
PTILODACTYLINAE Laporte, 1836
CHELONARIIDAE Blanchard, 1845
EULICHADIDAE Crowson, 1973
CALLIRHIPIDAE Emden, 1924
Superfamily ELATEROIDEA Leach, 1815
ARTEMATOPODIDAE Lacordaire, 1857
ELECTRIBIINAE Crowson, 1975
ARTEMATOPODINAE Lacordaire, 1857
BRACHYPSECTRIDAE LeConte & Horn, 1883
CEROPHYTIDAE Latreille, 1834
EUCNEMIDAE Eschscholtz, 1829
PEROTHOPINAE Lacordaire, 1857
PHYLLOCERINAE Reitter, 1905
PSEUDOMENINAE Muona, 1993
PALAEOXENINAE Muona, 1993
PHLEGONINAE Muona, 1993
MELASINAE Fleming, 1821
EUCNEMINAE Eschscholtz, 1829
MACRAULACINAE Fleutiaux, 1923
THROSCIDAE Laporte, 1840
ELATERIDAE Leach, 1815
CEBRIONINAE Latreille, 1802
TETRALOBINAE Laporte, 1840
THYLACOSTERNINAE Fleutiaux, 1920
LISSOMINAE Laporte, 1835
SEMIOTINAE Jacobson, 1913
PITYOBIINAE Hyslop, 1917
OXYNOPTERINAE Candeze, 1857
AGRYPNINAE Candeze, 1857 /Lacordaire, 1857
DENTICOLLINAE Stein & Weise, 1877 (1856)
NEGASTRIINAE Nakane & Kishii, 1956
DIMINAE Candeze, 1863
ELATERINAE Leach, 1815
CARDIOPHORINAE Candeze, 1859
HEMIOPINAE Fleutiaux, 1941
PHYSODACTYLINAE Lacordaire, 1857
Incertae sedis:
EUDICRONYCHINAE Girard, 1971
ANISCHIINAE Fleutiaux, 1936
SUBPROTELATERINAE Fleutiaux, 1920
PLASTOCERIDAE Crowson, 1972
DRILIDAE Blanchard, 1845
OMALISIDAE Lacordaire, 1857
LYCIDAE Laporte, 1836
LYCINAE Laporte, 1836
LEPTOLYCINAE Leng & Mutchler, 1922
ATELIINAE Kleine, 1928
METRIORRHYNCHINAE Kleine, 1926
EROTINAE LeConte, 1881
CALOCHROMINAE Lacordaire, 1857
TELEGEUSIDAE Leng, 1920
PHENGODIDAE LeConte, 1861
RHAGOPHTHALMINAE Olivier, 1907
PHENGODINAE LeConte, 1861
LAMPYRIDAE Latreille, 1817
PTEROTINAE LeConte, 1861
OTOTRETADRILINAE Crowson, 1972
CYPHONOCERINAE Crowson, 1972
OTOTRETINAE McDermott, 1964
AMYDETINAE Olivier, 1907
LAMPYRINAE Latreille, 1817
LUCIOLINAE Lacordaire, 1857
PHOTURINAE Lacordaire, 1857
OMETHIDAE LeConte, 1861
DRJLONIINAE Crowson, 1972
MATHETEINAE LeConte, 1881
OMETHINAE LeConte, 1861
CANTHARIDAE Imhoff, 1856 (1815)
CANTHARINAE Imhoff, 1856 (1815)
SJLINAE Mulsant, 1862
DYSMORPHOCERINAE Brancucci, 1980
MALTHININAE Kiesenwetter, 1852
CHAULIOGNATHINAE LeConte, 1861
ELATERIFORMIA Incertae Sedis
PODABROCEPHALIDAE Pic, 1930
RHINORHIPIDAE Lawrence, 1988
Series BOSTRICIDFORMIA Forbes, 1926
Superfamily DERODONTOIDEA LeConte, 1861
DERODONTIDAE LeConte, 1861
PELTASTICINAE LeConte, 1861
DERODONTINAE LeConte, 1861
LARICOBIINAE Mulsant & Rey, 1863-64
Superfamily BOSTRICHOIDEA Latreille, 1802
NOSODENDRIDAE Erichson, 1846
DERMESTIDAE Latreille, 1804
DERMESTINAE Latreille, 1804
Table 1. Continued.
MARIOUTINAE Jacobson, 1913
THORICTINAE Agassiz, 1846
ORPHILINAE LeConte, 1861
TRINODINAE Casey, 1900
THYLODRIINAE Semenov-Tian-Shanskij, 1913
ATTAGENINAE Laporte, 1840
MEGATOMINAE Leach, 1815
ENDECATOMIDAE LeConte, 1861
BOSTRICHIDAE Latreille, 1802
DYSIDINAE Lesne, 1921
POLYCAONINAE Lesne, 1896
BOSTRICHINAE Latreille, 1802
PSOINAE Blanchard, 1851
DINODERINAE C.G. Thomson, 1863
LYCTINAE Billberg, 1820
EUDERIINAE Lesne, 1934
ANOBIIDAE Fleming, 1821
EUCRADINAE LeConte, 1861
PTININAE Latreille, 1802
DRYOPHILINAE LeConte, 1861
ERNOBIINAE Pic, 1912
ANOBIINAE Fleming, 1821
PTILININAE Shuckard, 1840
ALVARENGANIELLINAE Viana & Martinez
XYLETININAE Gistel, 1856
DORCATOMINAE C.G. Thomson, 1859
MESOCOELOPODINAE Mulsant & Rey, 1864
BOSTRICHIFORMIA Incertae Sedis
JACOBSONIIDAE Heller, 1926
Series CUCUJIFORMIA Lameere, 1938
Superfamily LYMEXYLOIDEA Fleming, 1821
LYMEXYLIDAE Fleming, 1821
HYLECOETINAE Gistel, 1856
LYMEXYLINAE Fleming, 1821
MELITTOMMATINAE Wheeler, 1986
Superfamily CLEROIDEA Latreille, 1802
PHLOIOPHILIDAE Kiesenwetter, 1863
TROGOSSITIDAE Latreille, 1802
PROTOPELTINAE Crowson, 1966
LARINOTINAE S1ipiriski, 1992
PELTINAE Kirby, 1837
RENTONIINAE Crowson, 1966
DECAMERINAE Crowson, 1964
LOPHOCATERINAE Crowson, 1964
CALITINAE Reitter, 1922
EGOLIINAE Lacordaire, 1854
TROGOSSITINAE Latreille, 1802
CHAETOSOMATIDAE Crowson, 1952
CLERIDAE Latreille, 1802
THANEROCLERINAE Chapin, 1924
TILLINAE Leach, 1815
HYDNOCERINAE Spinola, 1844
785
CLERINAE Latreille, 1802
EPIPHLOEINAE Kuwert, 1893
ENOPLIINAE Gistel, 1856
TARSOSTENINAE Jacquelin du Val, 1860
KORYNETINAE Laporte, 1836
ACANTHOCNEMIDAE Crowson, 1964
PHYCOSECIDAE Crowson, 1952
PRIONOCERIDAE Lacordaire, 1857
MELYRIDAE Leach, 1815
MELYRINAE Leach, 1815
RHADALINAE LeConte, 1861
GIETELLINAE Constantin & Menier, 1987
DASYTINAE Laporte, 1840
MALACHIINAE Fleming, 1821
Superfamily CUCUJOIDEA Latreille, 1802
PROTOCUCUJIDAE Crowson, 1954
SPHINDIDAE Jacquelin du Val, 1860
PROTOSPHINDINAE Sen Gupta & Crowson,
1979
ODONTOSPHINDINAE Sen Gupta & Crowson,
1979
SPHINDIPHORINAE Sen Gupta & Crowson,
1979
SPHINDINAE Jacquelin du Val, 1860
BRACHYPTERIDAE Erichson, 1845
NITIDULIDAE Latreille, 1802
CALONECRINAE Kirejtshuk, 1982
CARPOPHILINAE Erichson, 1842
MELIGETHINAE C.G. Thomson, 1859
NITIDULINAE Latreille, 1802
CILLAEINAE Kirejtshuk & Audisio in
Kirejtshuk, 1986
CRYPTARCHINAE C.G. Thomson, 1859
CYBOCEPHALINAE Jacquelin du Val, 1858
SMICRIPIDAE Horn, 1879
MONOTOMIDAE Laporte, 1840
RHIZOPHAGINAE Redtenbacher, 1845
MONOTOMINAE Laporte, 1840
BOGANIIDAE Sen Gupta & Crowson, 1966
PARACUCUJJNAE EndrOdy-Younga & Crowson,
1986
BOGANIINAE Sen Gupta & Crowson, 1966
HELOTIDAE Reitter, 1876/Chapuis, 1876
PHLOEOSTICHIDAE Reitter, 1911
PHLOEOSTICHINAE Reitter, 1911
HYMAEINAE Sen Gupta & Crowson, 1966
AGAPYTHINAE Sen Gupta & Crowson, 1969
MYRABOLIINAE Lawrence & Britton, 1991
PRIASILPHINAE Crowson, 1973
TASMOSALPINGINAE Lawrence & Britton, -
1991
SILVANIDAE Kirby, 1837
BRONTINAE Erichson, 1845/Blanchard, 1845
SILVANINAE Kirby, 1837
PASSANDRIDAE Erichson, 1845/Blanchard, 1845
786
Table 1. Continued.
CUCUJIDAE Latreille, IS02
LAEMOPHLOEIDAE Ganglbauer, IS99
PROPALTICIDAE Crowson, 19S2
PHALACRIDAE Leach, ISIS
PHAENOCEPHALINAE Matthews, IS99
PHALACRINAE Leach, ISIS
HOBARTIIDAE Sen Gupta & Crowson, 1966
CAVOGNATHIDAE Sen Gupta & Crowson, 1966
CRYPTOPHAGIDAE Kirby, IS37
HYPOCOPRINAE Reitter, IS79
CRYPTOPHAGINAE Kirby, IS37
ATOMARIINAE LeConte, IS61
LAMINGTONIIDAE Sen Gupta & Crowson, 1969
LANGURIIDAE Crotch, IS73
XENOSCELINAE Ganglbauer, 1S99
SETARIOLINAE Crowson, 19S2
LANGURIINAE Crotch, IS73
CRYPTOPHILINAE Casey, 1900
TORAMINAE Sen Gupta, 1967
EROTYLIDAE Latreille, IS02
DACNINAE Gistel, ISS6
MEGALODACNINAE Sen Gupta, 1969
ENCAUSTINAE Crotch, 1S76/Chapuis, IS76
TRITOMINAE Curtis, IS34
EROTYLINAE Latreille, IS02
BYTURIDAE Jacquelin du Val, ISSS
PLATYDASCILLINAE Pic, 1914
BYTURINAE Jacquelin du Val, ISSS
BIPHYLLIDAE LeConte, IS61
BOTHRIDERIDAE Erichson, IS4S
TEREDINAE Seidlitz, 1S88
XYLARIOPHILINAE Pal & Lawrence, 19S6
ANOMMATINAE Ganglbauer, IS99
BOTHRIDERINAE Erichson, IS45
CERYLONIDAE Billberg, IS20
EUXESTINAE Grouvelle, !90S
LOEBLIORYLONINAE Slipinski, 1990
OSTOMOPSINAE Sen Gupta & Crowson, 1973
MURMIDIINAE Jacquelin du Val, 1S5S
CERYLONINAE Billberg, IS20
ALEXIIDAE Imhoff, !SS6
DISCOLOMATIDAE Horn, IS7S
NOTIOPHYGINAE Jacobson, 1915
DISCOLOMATINAE Horn, 1S7S
APHANOCEPHALINAE Grouvelle, 1912
CEPHALOPHANINAE John, 1954
PONDONATINAE John, 1954
ENDOMYCHIDAE Leach, 181S
MEROPHYSIINAE Seidlitz, 1872
HOLOPARAMECINAE Seidlitz, IS8S
LEIESTINAE C.G. Thomson, 1863
EUPSILOBIINAE Casey, IS95
ENDOMYCHINAE Leach, ISI5
EPIPOCINAE Gorham, 1873
LYCOPERDININAE Redtenbacher, 1S44
MYCETAEINAE Jacquelin du Val, 1S57
ANAMORPHINAE Strohecker, 19S3
PLEGANOPHORINAE Jacquelin du Val, ISSS
XENOMYCETINAE Strohecker in Arnett, 1962
COCCINELLIDAE Latreille, 1S07
STICHOLOTIDINAE Weise, 1901
COCCIDULINAE Mulsant, IS46
SCYMNINAE Mulsant, 1846
CHILOCORINAE Mulsant, 1S46
COCCINELLINAE Latreille, IS07
EPILACHNINAE Mulsant, 1S46
CORYLOPHIDAE LeConte, 1S52
PELTINODINAE Paulian, 19SO
CORYLOPHINAE LeConte, IS52
SERICODERINAE Matthews, ISSS
PARMULINAE Poey, ISS4
LATRIDIIDAE Erichson, IS42
LATRIDIINAE Erichson, IS42
CORTICARIINAE Curtis, IS29
Superfamily TENEBRIONOIDEA Latreille, IS02
MYCETOPHAGIDAE Leach, ISI5
ESARCINAE Reitter, 1882
MYCETOPHAGINAE Leach, ISI5
BERGININAE Leng, 1920
ARCHEOCRYPTICIDAE Kaszab, 1964
PTEROGENIIDAE Crowson, 1953
CIIDAE Leach in Samouelle, ISI9
SPHINDOCIINAE Lawrence, 1974
CIINAE Leach in Samouelle, ISI9
TETRATOMIDAE Billberg, 1820
PISENINAE Miyatake, 1960
TETRATOMINAE Billberg, 1820
PENTHINAE Lacordaire, IS59
MELANDRYIDAE Leach, ISI5
HALLOMENINAE Mulsant, 1856/Gistel, IS56
EUSTROPHINAE Gistel, 1856
MELANDRYINAE Leach, 1815
OSPHYINAE Mulsant, 1856 (1840)
MORDELLIDAE Latreille, 1S02
CTENIDIINAE Franciscolo, 1951
MORDELLINAE Latreille, 1802
RHIPIPHORIDAE Gemminger & Harold, 1870
(1853)
PELECOTOMINAE Seidlitz, 1875
MICHOLAEMINAE Viana, 1971
PTILOPHORINAE Gerstaecker, 1855
HEMIRHIPIDIINAE Heller, 1921
RHIPIDIINAE Gerstaecker, 1855
RHIPIPHORINAE Gemminger & Harold, 1870
(1853)
COLYDIIDAE Erichson, 1842
PYCNOMERINAE Erichson, 1845
COLYDIINAE Erichsen, 1842
MONOMMATIDAE Blanchard, 1845
ZOPHERIDAE Solier, 1S34
ZOPHERINAE Solier, 1834
Table 1. Continued.
USECHINAE Horn, 1867
ULODIDAE Pascoe, 1869
PERIMYLOPIDAE St. George, 1939
CHALCODRYIDAE Watt, 1974
TRACHELOSTENIDAE Lacordaire, 1859
TENEBRIONIDAE Latreille, 1802
LAGRIINAE Latreille, 182S (IS20)
PHRENAPATINAE Solier, 1834
ZOLODININAE Watt, 1974
PIMELIINAE Latreille, 1S02
TENEBRIONINAE Latreille, 1802
ALLECULINAE Laporte, 1840
DIAPERINAE Latreille, 1802
COELOMETOPINAE Lacordaire, 1859/Schaum,
!SS9
Incertae sedis:
COSSYPHODINAE Wasmann, 1899
PROSTOMIDAE C.G. Thomson, 1859
SYNCHROIDAE Lacordaire, 18S9
OEDEMERIDAE Latreille, 1810
CALOPODINAE Costa, 1852
OEDEMERINAE Latreille, 1810
STENOTRACHELIDAE C.G. Thomson, !SS9
STENOTRACHELINAE C.G. Thomson, IS59
NEMATOPLINAE LeConte, IS62
CEPHALOINAE LeConte, 1862
Incertae sedis:
STOLIINAE Nikitsky, 1985
MELOIDAE Gyllenhal, ISIO
ELETICINAE Wellman, 1910
MELOINAE Gyllenhal, lSIO
NEMOGNATHINAE Laporte, 1840
MYCTERIDAE Blanchard, IS45
MYCTERINAE Blanchard, 1845
LACCONOTINAE LeConte, 1862
HEMIPEPLINAE Lacordaire, 1854
BORIDAE C.G. Thomson, !S59
BORINAE C.G. Thomson, 1859
SYNERCTICINAE Lawrence & Pollock, 1994
TRICTENOTOMIDAE Blanchard, 1845
PYTHIDAE Solier, IS34
PYROCHROIDAE Latreille, 1807
TYDESSINAE Nikitsky, 19S6
PILIPALPINAE Abdullah, 1964
PEDILINAE Lacordaire, 1859
PYROCHROINAE Latreille, 1807
Incertae sedis:
AGNATHINAE Lacordaire, 1859
SALPINGIDAE Leach, 1815
OTHNIINAE LeConte, 1861
PROSTOMINIINAE Grouvelle, 1914
AGLENINAE Horn, 1878
INOPEPLINAE Grouvelle, 190S
SALPINGINAE Leach, 1815
AEGIALITINAE LeConte, 1S62
DACODERINAE LeConte, 1S62
787
ANTHICIDAE Latreille, 1819
EURYGENIINAE LeConte, IS62
LAGRIOIDINAE Abdullah & Abdullah, 196S
AFREMINAE Levey, 19SS
MACRATRIINAE LeConte, IS62
STEROPINAE Jacquelin du Val, IS63
ISCHALIINAE Blair, 1920
COPOBAENINAE Abdullah, 1969
LEMODINAE Lawrence & Britton, 1991
TOMODERINAE Bonadona, 1961
ANTHICINAE Latreille, ISI9
ADERIDAE Winkler, 1927
SCRAPTIIDAE Mulsant, 18S6/Gistel, 18S6
SCRAPTIINAE Mulsant, !S56/Gistel, !SS6
ANASPIDINAE Mulsant, 18S6
Superfamily CHRYSOMELOIDEA Latreille,
1S02
CERAMBYCIDAE Latreille, 1S02
VESPERINAE Mulsant, 1839
OXYPELTINAE Lacordaire, 1869
DISTENIINAE J. Thomson, 1860
ANOPLODERMATINAE Guerin-Meneville, 1840
PHILINAE J. Thomson, 1860
PARANDRINAE Blanchard, IS45
PRIONINAE Latreille, IS02
SPONDYLIDINAE Audinet-Serville, 1S32
APATOPHYSEINAE Lacordaire, 1869
NECYDALINAE Latreille, 1825
LEPTURINAE Latreille, 1802
CERAMBYCINAE Latreille, 1802
LAMIINAE Latreille, 1825
MEGALOPODIDAE Latreille, 1802
PALOPHAGINAE Kuschel & May, 1990
ZEUGOPHORINAE Biiving & Craighead, 1931
MEGALOPODINAE Latreille, 1802
ORSODACNIDAE C.G. Thomson, 1859
ORSODACNINAE C.G. Thomson, 1859
AULACOSCELIDINAE Chapuis, 1S74
CHRYSOMELIDAE Latreille, IS02
SAGRINAE Leach, 1815
BRUCHINAE Latreille, 1802
DONACIINAE Kirby, 1837
CRIOCERINAE Latreille, 1804
HISPINAE Gyllenhal, 1813
CHRYSOMELINAE Latreille, 1802
GALERUCINAE Latreille, 1802
LAMPROSOMATINAE Lacordaire, 1848
CRYPTOCEPHALINAE Gyllenhal, 1813
EUMOLPINAE Hope, 1840
Superfamily CURCULIONOIDEA Latreille, 1802
NEMONYCHIDAE Bedel, 1882
NEMONYCHINAE Bedel, 1882
RHINORHYNCHINAE Voss, 1922
DOYDIRHYNCHINAE Pierce, 1916
788 789

Table 1. Continued.
ANTHRIBIDAE Billberg, 1820
URODONTINAE C.G. Thomson, 1859
ANTHRIBINAE Billberg, 1820
CHORAGINAE Kirby, 1819
BELIDAE Schoenherr, 1826
BELINAE Schoenherr, 1826
OXYCORYNINAE Schoenherr, 1840
AGLYCYDERINAE Wollaston, 1864
ATTELABIDAE Billberg, 1820
RHYNCHITINAE Gistel, 1856
ATTELABINAE Billberg, 1820
BRENTIDAE Billberg, 1820
EURHYNCHINAE Lacordaire, 1863
CYLADINAE Schoenherr, 1823
North of Mexico$, Downie and Arnett (1994; final volumes in press), Hatch
(1953-71), Insects and Arachnids of Canada and Alaska$, White (1983).
BRENTINAE Billberg, 1820
ANTLIARHININAE Schoenherr, 1823 MEXICO, CENTRAL AND SOUTH AMERICA, WEST INDIES: Biologia Centrali-
NANOPHYINAE Gistel, 1856
Americana (Coleoptera completed in 18 parts, 1880--1911; see Godman 1915
APIONINAE Schoenherr, 1823
CARIDAE Thompson, 1992 for summary), Blackwelder (1944-57), Blackwelder and Arnett (1974-75),
ITHYCERIDAE Schoenherr, 1823 Costa Lima (1952-56), Costa et al. (1988), Elgueta and Arriagada (1989),
CURCULIONIDAE Latreille, 1802
Hogue (1993), Pefia (1987), Quintero and Aiello (1992), Spangler (1981,
BRACHYCERINAE Billberg, 1820
CURCULIONINAE Latreille, 1802 1982).
DRYOPHTHORINAE Schoenherr, 1825
COSSONINAE Schoenherr,, 1825 SUBANTARCTIC ISLANDS: Crafford et al. (1986), Gressitt (1964, 1970, 1971),
SCOLYTINAE Latreille, 1804 Robinson (1984).
PLATYPODINAE Shuckard, 1840

EUROPE: Die Kafer Mitteleuropas (only complete recent series, being up-
dated and extended to larvae), Fauna CSSR$, Fauna Entomologica Scandina-
vica$, Fauna Hungariae$, Fauna Iberica$, Fauna d'Italia$, Fauna Polski$,
Fauna Republicii Socialiste Romania$, Fauna SSSR$, Faune de France$,
Handbooks for the Identification of British Insects$, Harde (1984), Katalog
Fauny Polski$, Klausnitzer (1978), Klucze do Oznaczania Owad6w Polski
[Keys for the Identification of Polish Insects]$, Pope (1977), Silfverberg
(1992), Svensk Insektfauna$ .
ASIA: Check-list of Coleoptera of Japan$, Coleoptera of Japan in Color
(1984-86, 4 vols.), Fauna Japonica$, Fauna of British India$, Fauna of Saudi
Arabia$, Fauna SSSR$, Ler (1989, 1992).
AFRICA: Alfieri (1976), Faune de Madagascar$, Ferreira (1965-66, 1967),
Prins (1984), Scholtz and Holm (1985), South African Animal Life$.
ATLANTIC ISLANDS: Basilewski and Decelle (1972), Borges (1990), Geist-
hardt (1988), Wollaston (1864).
AUSTRALIA: Britton (1970), Lawrence and Britton (1991, 1994), Matthews
(1980--87), Moore (1980--92), Semmens et al. (1992), Zoological Catalogue of
Australia$.
NEW ZEALAND: Fauna of New Zealand$, Hudson (1934; including last com-
plete checklist), Kuschel (1990), Ramsay and Crosby (1992), Watt (1982).
NEW GUINEA: Gressitt and Hornabrook (1977), Gressitt and Szent-Ivany
(1968).
PACIFIC ISLANDS: Insects of Micronesia$, Nishida (1994), Peck and
Kukalova-Peck (1990).
CANADA AND UNITED STATES: Arnett (1963, 1985), Blackwelder and Arnett
(1974-75), Bousquet (1990, 1991), Catalog of the Coleoptera of America
Family-group names
A thorough summary of data on family-group names for all of Coleoptera does
not exist, but is essential if family and subfamily names are to become stabi-
lized and brought into compliance with current rules governing such names
since 1960 (ICZN 1961, 1985a). The treatments for many families in the last
complete world catalog of beetles, Junk and Schenkling's Coleopterorum Cata-
logus, gave no references at all for suprageneric taxa, or included references
to different taxonomic concepts of these higher taxa rather than nomenclatural
establishment of the names. Several authors of works covering all Coleoptera
made a special effort to cite author and year for all higher taxon names used,
notably Handlirsch (1925), Arnett (1963), authors ofthe Katalog Fauny Polski,
and Silfverberg (1992), but without direct references. With the exception of the
outdated but still useful Handlirsch classification, these works cover limited
regional faunas. Watt (1975) carefully reviewed the earliest (pre-1816) Coleop-
tera names. Detailed worldwide reviews of all family-group names have been
completed recently for terrestrial Adephaga (Madge 1989), Hydrophiloidea and
Staphylinoidea (Newton and Thayer 1992), and Cucujoidea (Pakaluk et al. in
press), or are included in catalogs or taxonomic treatments of several medium-
to small-sized families. For the remaining two-thirds of Coleoptera, though,
information on family-group names is unreliable or missing, potentially result-
ing in the use of incorrect names and later confusion when such names are
corrected.
The current rules (or Code) governing family-group names have been set by
the International Commission on Zoological Nomenclature (ICZN 1985a). The
most important provisions of the Code as they affect family-group names may
be summarized briefly as follows: Family-group names as defined and regu-
lated by the Code are those group names above the generic level up through
superfamily and based on a nominal type genus. The type genus must be an
available name (i.e., meet provisions of the Code specified in Articles 1b and
790
10-20) and must have been treated as the valid name for a genus inclu~ed in
the group when the group was proposed; it need n~t be the oldest ~a~e m the
group or be treated currently as a valid genus, but It may not be a ~umor ho~­
onym. The family-group name must have been proposed as a noun m the nomi-
native plural, provided with a latinized suffix, and clearly used as a_g~oup name
rather than as a plural term for the members of a genus; a non-latmized name
proposed before 1900 is available if it was later latinized and meets other con-
ditions (see (1) below). A name proposed after 1930 must have been accompa-
nied by a description or definition or a reference thereto, or be a replacement
name and after 1960 must not have been proposed conditionally. Each name
must be formed from the correct spelling and stem of the name of its type ge-
nus and have a standard ending determined by the rank at which it is used; a
name not so formed when proposed (including one based on an unjustified
emendation) is to be corrected but keeps its original author and date. All names
based on a given type genus are coordinate, i.e., take the author ~d.date of the
first available name based on that type genus, regardless of the ongmal or cur-
rent rank. With one exception (see (2) below), priority applies to application
of family-group names: the valid name for a taxon is the oldest available name
based on a generic name included in the taxon. Finally, the famil~-group name
must not be a homonym, a situation that can arise not only when Its type genus
is a homonym (leading to automatic rejection of both genus-group and fam~ly­
group names if the type genus is a junior homonym) but also ':hen. fam~ly­
group names have been based on different generic names havmg Ide~tlcal
stems (in which case the matter is to be referred to the ICZN for resolutwn).
The Code itself should be consulted for details, qualifications and examples
of these general provisions; it also includes an extensi':e glos~ary. ~or furt~er
discussion of general principles and specific problems m dealmg with family-
group names in beetles and other insects see Madge (1989), Michener (1986),
Newton and Thayer (1992), and Silfverberg (1990).
Our treatment of beetle family and subfamily names here is intended only
as a starting point toward documenting the correct names to use, by focussing
on those names commonly used at this level rather than all family-group names
in beetles. But we have gone beyond a simple compilation of names and dates
from the literature in several ways that are intended to document the names,
improve upon the accuracy of name use, and highlight problems in need of
further study. We proceeded as follows:
First, we compiled the earliest citations of author and year for all names
commonly used at the family and subfamily level from standard references
such as the Coleopterorum Catalogus, Genera Insectorum, Agassiz (1846a,
1846b), Handlirsch (1925), and the Zoological Record; from recent sources
known to provide original data on family-group names, including Arnett
(1963), Watt (1975), Abdullah (1975), Katalog Fauny Polski, Carpenter (1992),
and Silfverberg (1992); and from recent group-specific reviews of family-group
791
names, including Madge (1989), Newton and Thayer (1992), Nilsson et al.
(1989), and Pakaluk et al. (in press).
Second, we sought to confirm the earliest citation found for each name by
checking the original reference to be sure the name was actually made avail-
able there. In the case of citations that included author and year only, we lo-
cated references with the aid of the Index Litteraturae Entomologicae Series
I (through 1863, Hom and Schenkling 1928-29) and Series II (1864-1900,
Derksen and Scheiding 1963-72), and the Zoological Record (1864- present).
We entered data on the earliest use of each name, including original spelling,
rank, and type genus, in a computer database; references were added to a sec-
ond database. In the present work, we cite author/year/page number for all
names treated as valid in our list as well as those synonyms that have been in
recent use or are discussed for any reason, and include the full reference for
each such name in our bibliography. The page cited is of the main entry in the
text, unless the name appeared on an earlier page that was definitely published
prior to the main entry and was sufficient to validate the name. The only in-
stance in which we were unable to examine the original reference for a name
is indicated by including "[not seen]" after the citation in the text and after the
reference in the bibliography.
Third, we scanned each reference consulted in connection with a particular
name for reference to any earlier use of the name, for synonyms of it, and for
other names that might have been established in the same work but overlooked.
We also scanned many catalogs, other general references, and major reviews
or revisions for first use of names. In this manner, we discovered earlier estab-
lishment of many names generally attributed to later authors.
Fourth, we consulted standard nomenclatorial resources for potential prob-
lems with individual family-group names or their type genera. We scanned the
Official Lists of the International Commission on Zoological Nomenclature for
decisions that might affect family-group names in beetles. We also checked the
type genus of each family and subfamily name in Neave (1939-40, and supple-
ments to 1977) to detect possible problems with availability, original spelling,
or homonymy. Finally, we scanned the type genera for those of Greek origin
whose family-group name stems are often formed incorrectly, i.e. , not accord-
ing to grammatical rules given in the Code (ICZN 1985a) and discussed by
Steyskal (1980), Newton and Thayer (1992), and others. The few required
changes we found, indicated as correction of stem in Table 2, involve common
terminations derived from the Greek words for border (-lorna, stem -lomat-),
eye (-omma, stem -ommat-), foot (-pus, stem -pod-), and mouth (-stoma, stem
-stomat-).
Each of these kinds of checks turned up problems with certain names in
current use. In some cases, we have been able to resolve a problem after fur-
ther investigation. In other cases an apparent problem is indicated but left for
specialists in the group concerned to investigate and resolve. In either situation,
792
any necessary nomenclatural discussion is provided in the paragraphs marked
with a double asterisk (**). In a few cases where a name was first used inde-
pendently by two authors in the same year, and we could not determine the
relative or absolute dates, we cite both authors separated by a slash (/).
Even when the nomenclatural facts involved in a particular case are clear,
the choice of what name to use may not be. Judgement is often needed in three
areas:
(1) NON-LATINIZED NAMES (Article 11f(iii), ICZN 1985a). This Article al-
lows for availability of a non-latinized name proposed before 1900, provided
the name has been subsequently latinized and generally accepted as valid and
as dating from the original non-latinized form. Problems in implementing this
rule in Coleoptera were discussed by Madge (1989) and Newton and Thayer
(1992), who advocated that such names be accepted if they have been latinized
later and achieved wide use, without attempting to document "general accep-
tance" and citation of the non-latinized form. We have adopted this approach
in the present list, and have not differentiated between latinized versus non-
latinized original use. Certainly, numerous non-latinized names first used by
Lacordaire, Laporte, Mulsant, and other French-speaking authors have always
been accepted without question as dating from those authors, and we see no
reason not to accept all such names. (Care must be taken, however, to differen-
tiate between true group names and the often similar-looking terms used by
early French authors to refer to the members of a genus; several common cita-
tions for beetle family-group names are based on the latter.)
(2) SYNONYMY OF TYPE GENUS (Article 40b, ICZN 1985a). This Article pro-
vides for setting aside priority in one special circumstance. If a family-group
name was replaced before 1961 because its type genus was recognized as a
junior synonym, and the replacement family-group name won general accep-
tance, then the replacement name is to be maintained, but takes the date of
precedence of the replaced name. A name maintained by provision of this arti-
cle is to be cited with its own author and date, followed in parentheses by the
date of the older replaced name. Determining "general acceptance" of the re-
placement name is sometimes quite difficult, and in ambiguous cases we have
tended to follow priority rather than use Article 40b to conserve the replace-
ment name.
(3) ALTERATION OF WELL-KNOWN NAMES. Some changes required by priority
would result in the replacement of long-established and widely used names
with little-known ones (e.g., Sarotriidae over Colydiidae, Horiidae over
Meloidae, Carabidae: Graphipterinae over Harpalinae) or replacing the name
of a large group with that of a much smaller recently and/or doubtfully added
group (e.g., Cebrionidae over Elateridae, Ptinidae over Anobiidae, Tenebrion-
idae: Cossyphinae over Lagriinae). In other cases, correction of the family-
group name to reflect the correct original spelling of the type genus (e.g., Kryp-
793
tophagidae rather than Cryptophagidae) might cause confusion. We have been
especially reluctant to implement significant name changes given potential
changes to the Code now under consideration (ICZN 1990; a draft of a new
fourth edition of the Code is nearly complete but not yet available to us). In
those cases where we feel that specialists on the group concerned may choose
to petition the ICZN to conserve existing usage, or where specialists have indi-
cated their intention to do so, we point out the problem but maintain existing
usage. In still other cases (e.g., Cantharidae, Rhipiphoridae, Ipidae) the earliest
use of a well-known family-group name was based on a misidentified type
genus, but the name as currently used is based on the correctly identified type
genus; all such cases should be referred to the ICZN for permanent resolution
(Article 41).
Table 2 summarizes the more important changes from common usage of
family-group names in beetles that would result from implementing the current
Code (ICZN 1985a), indicating those that have not been implemented here or
elsewhere.
Problems associated with two early works in Coleoptera deserve special
note. Many names commonly attributed to "Leach 1817", specifically those
given the reference "Encycl. Brit." in Agassiz (1846a), were not used by Leach
but were actually first used by Fleming (1821). This mistaken attribution was
discussed at length and corrected by Newton and Thayer (1992); we note here
in further confirmation that the text in which these names appeared was re-
printed almost verbatim by Fleming (1822). The other problem area, still unre-
solved, concerns dozens of family names used in a checklist published by Gis-
tel (1856). In that widely overlooked (or ignored) work Gistel used many new
family-group names, and when these were based (explicitly or by implication)
on previously established type genera they are available and are cited by us
when appropriate. Many other family-group names, however, were based on
generic names that were also new, and which generally have been treated as
nomina nuda or ignored in the systematic literature (including Neave 1939-40).
These generic names had species listed under them, and when the species
names were also new (not attributed to an author) the whole set of names is
unavailable. Other new generic names, however, included available species
names of earlier authors, making these generic names (and the family-group
names based on them) available according to the current Code (ICZN 1985a).
The task of determining the availability and placement of the many generic and
family-group names involved would be difficult and tedious, and is likely to
result in the availability of several completely unused names that will upset
existing usage if strict priority is followed. The idea of seeking the rejection of
Gistel's (1856) entire work is tempting, but some generic names established
there are in current use, and a dozen family-group names for which we cite
Gistel (1856) as author would need to be reevaluated. Adequately resolving the
 Table 2. Summary of major changes from recent usage of family-group names in Coleoptera required by the Code (ICZN 1985a). See text discussion -.1
under each taxon for more detail. \0
~

Narne change required Reason Examples of recent

from: to: implementation
Sphaeriidae Microsporidae group name homonymy ICZN (1985b), Lawrence & Britton (1991, 1994)
Harpalinae Graphipterinae priority NOT IMPLEMENTED
Limulodinae Cephaloplectinae priority Newton & Thayer (1992)
Anisotomidae, -inae Leiodidae, -inae priority Peck (1990), Newton & Thayer (1992), Wheeler (1984)
Coloninae Koloninae original spelling of stem NOT IMPLEMENTED (ICZN appl. pending)
Catopinae, Cholevinae priority Peck (1990), Newton & Thayer (1992), Zwick (1979)
Leptodirinae
Leptininae Platypsyllinae priority Newton & Thayer (1992)
Clidicinae Mastiginae priority Newton & Thayer (1992)
Glypholominae Glypholomatinae correction of stem Newton & Thayer (1992, 1995)
Acanthoceridae Ceratocanthidae preoccupied type genus Cartwright & Gordon (1971), Paulian (1988), Scholtz (1990)
Allidiostominae Allidiostomatinae correction of stem present work
Dynamopinae Dynamopodinae correction of stem present work
Phaenomerinae Phaenomeridinae group name homonymy, and ICZN (1962)
correction of stem
Eucinetoidea Scirtoidea priority Lawrence & Britton (1994)
Helodidae Scirtidae priority/ unavailable type genus Pope (1976, 1977), Silfverberg (1992)
Julodinae Jalodinae original spelling of stem? NOT IMPLEMENTED
Larinae Larainae group name homonymy ICZN (1988), Lawrence & Britton (1991, 1994)
Callirhipidae Zenoidae priority NOT IMPLEMENTED
Artematopidae, Artematopodidae, -inae correction of stem Paulian (1988)
-inae
Eucnemidae Melasidae priority NOT IMPLEMENTED (ICZN appl. pending)
Elateroidea, -idae Cebrionoidea, -idae priority NOT IMPLEMENTED (ICZN appl. pending?)
Agrypninae Pangaurinae priority NOT IMPLEMENTED (ICZN appl. pending?)
Athoinae Denticollinae priority Stibick (1979)
Anthreninae Megatominae priority Lawrence & Britton (1994)
Anobiidae Ptinidae priority NOT IMPLEMENTED
Hedobiinae Eucradinae priority Katalog Fauny Polski 11

Table 2. Continued.

Name change required Reason Examples of recent

from: to: implementation
Tricoryninae Mesocoelopodinae priority present work
Melittomminae Melittommatinae correction of stem present work
Ostominae Peltinae priority Pope (1977), Silfverberg (1992), Pauiian (1988)
Phyllobaeninae Hydnocerinae priority present work
Epiphloeinae Ichneinae priority NOT IMPLEMENTED
Enopliinae Platynopterinae priority NOT IMPLEMENTED
Sphindidae, -inae Aspidiphoridae, -inae priority Pakaluk et al. (in press); NOT IMPLEMENTED here
Cateretidae/ Brachypteridae priority Pakaluk et al. (in press)
Kateretidae
Rhizophagidae Monotomidae priority Pakaluk et al. (in press), Lawrence & Britton (1994)
Rhizophaginae Ryzophaginae original spelling of stem NOT IMPLEMENTED (ICZN appl. pending)
Uleiotinae Brontinae priority Pakaluk et al. (in press), Lawrence & Britton (1994)
Cryptophagidae, Kryptophagidae, -inae original spelling of stem NOT IMPLEMENTED (ICZN appl. pending)
-inae
Triplacinae Tritominae priority Silfverberg (1992), Pakaluk et al. (in press)
Sphaerosomatidae Alexiidae priority [not required] Slipinski & Pakaluk (1992), Pakaluk et al. (in press)
Discolomidae, Discolomatidae, -inae correction of stem Pakaluk et al. (in press)
-inae
Stenotarsinae Epipocinae priority Pakaluk et al. (in press)
Eumorphinae Lycoperdininae priority Pakaluk et al. (in press)
M ychotheninae Anamorphinae priority Pakaluk et al. (in press)
Trochoideinae Pleganophorinae priority Pakaluk et al. (in press)
Saciinae Parmulinae priority Pakaluk et al. (in press)
Lathridiidae, Latridiidae, -inae original spelling of stem Silfverberg (1992), Pakaluk et al. (in press)
-inae
Latridiidae Corticariidae priority Pakaluk et al. (in press); NOT IMPLEMENTED here -.1
\0
VI
796 797

problems posed by Gistel's work is beyond the scope of our study, but we want
to call attention to the need for such resolution.
Names above the superfamily level (i.e., order, suborder, and series or infra-
order in Coleoptera) are not regulated by the Code. We have used names that
are widely accepted, including genus-based series names ending in "-iformia",
citing author and date of the first use of each name in more or less its present
form for a group more or less equivalent to the present group.
The limitations of the present treatment of family and subfamily names must
be emphasized. We estimate that more than 10,000 family-group names based
on distinct type genera have been proposed in Coleoptera, and perhaps half of
these are in current use at some level. A thorough review of all such names
would no doubt tum up earlier uses of some names cited here, earlier names
for some taxa used here, and other problems (e.g., misidentification of type
genera) that we could not easily detect in this review. Nevertheless, we feel
that the family and subfamily classification presented here provides an appro-
priate opportunity and format for a preliminary review of beetle family-group
names that may serve as a basis for further work.

Acknowledgements

"0
Since this family-subfamily list has been accumulating over a number of years,
1:-• 1:-•
§ beginning in the early 1970's, a large number of people have had input into it
a a » ;>, and any attempt to acknowledge them all individually would be futile. R.A.
~ £<ll a
»
aa
'+-< '+-< 1::
>,II)
1:: .....
0
Crowson has been a major source of ideas and inspiration, although he will not
0 0 a 0
a a<+-<
<ll

agree with portions of the classification presented. Curators of many of the

~
OJ) OJ) 0 0
1:: .s 0
..<:: ..<:: § world's major insect collections have provided material for study over the
~p.. ~
p..
'+-<
0 (I)
a
(1) ......
a ....\:!
<ll <ll 1::
0 o:l o:l years, and many specialists have commented on various beetle groups. We
oo~::o~::so
: : \:!·;:::o ·;:::o
.s ·;o·.::: 0000
(;3
§ 1::
(;3
·;::: ·;::: ·;::: ·;::: u ·;:::
·;::: ·;::: p.. ·;::: p.. thank all for their input and assistance.
<ll
o:l
II)
'So
·;::: · ~·8 8 .g .g
0 0 0 0
·;::: ·;::: ·;::: ·;::: ·;:::0 ·;:::0 8;::l ·;:::0 ;::l
0
0
·;::: We specifically thank the following individuals for comments on partial late
~ 0 oo..uo..o.. p..p..p..p.. P..P..bllP..6h p..
drafts or help with obscure literature: M.A. Alonso-Zarazaga, R.S. Anderson,
W.F. Barr, R. Bejsak, C.L. Bellamy, D.S. Chandler, R. deKeyzer, D.G. Furth,
L.H. Herman, H.F. Howden, M.A. Ivie, P.J. Johnson, A.G. Kirejtshuk, R.A.B.
Leschen, J.V. McHugh, D.R. Maddison, R. Miller, D.A. Pollock, R.D. Pope,
H. Sasaji, H. Silfverberg, S.A. Slipinski, J.B. Stribling, R.T. Thompson, B.
Valentine, T. Weir, and the late F.G. Werner. J. Pakaluk and M.K. Thayer
provided helpful comments on the entire work, and Thayer helped greatly with
final checking of text references against the bibliography.
Thanks are also given to the staff of the CSIRO Black Mountain Library,
Field Museum Library, and Museum of Comparative Zoology (Harvard Uni-
versity) Library for their help in obtaining references onsite or through interli-
brary loan, and to W. Dressler for a number of tasks. The scanning electron
micrographs used in Figs. 1-4 were produced by E. Seling and partly funded
798
by NSF grant BMS 7502606. The CSIRO Division of Entomology Photogra-
phy Unit (J. Green) and Graphics Unit (S. Smith, C. Hunt) prepared the plate.
Systematic list of families and subfamilies of Coleoptera
*Comments on classification, and general references.
**Comments on family-group names.
Order COLEOPTERA Linnaeus, 1758: 345
*The limits of the order Coleoptera based on Recent forms are generally agreed
upon, with one notable exception: the inclusion of the Strepsiptera. The phylo-
genetic relationships of the Strepsiptera have been disputed since the order was
first described by Kirby (see Lawrence, Slipinski and Pakaluk 1995). Many
authors, including Crowson (1955, 1960, 1981), consider the group to be either
the sister group of Coleoptera or a family (Stylopidae) within the order. A
sister group relationship between Strepsiptera and Coleoptera is often argued
on the basis of the following shared features: (1) presence of a gula, (2) free
prothorax and more or less closely associated meso- and metathorax, (3) use
of hind wings in flight, (4) the large metacoxae with little or no motility, (5)
the presence of a type of metendosternite similar to those of various beetle
groups but unknown outside the Coleoptera, (6) more extensive sclerotization
of abdominal sternites than tergites, (7) first-instar larva closely resembling
those of other endoparasitic beetles, such as Rhipiphoridae. Crowson (1960,
1981) also argued that the type of larval leg and absence of a notopleural suture
places the Strepsiptera (as a family Stylopidae) in the suborder Polyphaga, and
that the absence of functional spiracles on the freely exserted 8th abdominal
segment supports their placement in the series Cucujiformia, with a likely sister
group being the Lymexylidae.
Kinzelbach (1971a, 197lb), Kinzelbach and Pohl (1994), and Kristensen
(1981, 1991) dispute Crowson's claim that a gula is present in Strepsiptera and
stress those features of the order which appear to be more primitive than in
Coleoptera (and thus incompatible with the idea that Strepsiptera evolved
within a derived beetle group like the Cucujiformia): (1) several veins in the
forewing, (2) motility and lateral attachment of the forewing, (3) tegulae on
both fore and hind wings, (4) well-developed lOth abdominal segment, and (5)
incomplete holometabolous development (early appearance of compound eyes
and wing buds, incorporation of numerous organs unchanged into the adult, and
very short pupal stage). Kukalova-Peck and Lawrence (1993) found that the
hind wing venation supported a sister group relationship between Strepsiptera
and Coleoptera, but did not support the placement of the former within the
Cucujiformia.
799
Jeannel (1945) stressed the similarities between the strepsipteran thorax and
that of Mecoptera, in which the metathoracic postscutellum is fused to the first
abdominal tergite and separated from the scutellum by a membranous region.
He also noted similarities between the structure of the ninth abdominal segment
in Strepsiptera and primitive Hymenoptera, and referred to the strepsipteran
triungulinid larva as a chalcidoid planidium with legs. He concluded that Strep-
siptera may have evolved from an early hymenopteran lineage shortly after
their divergence from Paleozoic mecopteroid ancestors.
A very recent contribution to this subject was provided by Whiting and Whe-
eler (1994), who not only cited unpublished molecular sequence data (nuclear
18S ribosomal DNA) supporting a Strepsiptera-Diptera sister group relation-
ship, but also hypothesized that "a homeotic mutation resulting in ectopic ex-
pression of Ultrabithorax ... protein" transformed the second thoracic segment
into a metathorax and the third into a mesothorax in the ancestral strepsipteran.
Additional morphological support was also provided, including the presence in
all male Strepsiptera, Mecoptera and basal Diptera of an enlarged and ring-like
ninth abdominal segment.
Although the Adephaga have been recognized as a distinct group of beetles
since the time of Latreille (see Lawrence, Slipinski and Pakaluk 1995), the
Coleoptera was not divided into two suborders, Adephaga and Polyphaga, until
the late 19th Century (Emery 1886). Although proposed by Kolbe in 1908, the
Archostemata were not generally recognized until Forbes (1926) and Boving
and Craighead (1931) clearly defined the group as a third suborder of Coleo-
ptera. Myxophaga is an even more recent concept (Crowson 1955), which is
periodically reassessed (see below). The relationships of the four suborders
were first discussed by Crowson (1960), who proposed an evolutionary sce-
nario involving three original stocks: wood-boring Archostemata, predaceous
Adephaga, and a third retaining the original habits of a subcortical detritivore
or fungivore. Continuing this scenario, he further suggested that Myxophaga
split off this pre-polyphagan lineage after the reduction in the larval leg but
before the loss of the oblongum cell in the wing and the notopleural suture.
Klausnitzer (1975) presented a Hennigian argumentation scheme along similar
lines with Myxophaga and Polyphaga forming a monophyletic group (Hete-
rophaga) and these, in turn, forming a monophyletic group (Pantophaga) with
Adephaga.
Ponomarenko (1969, 1971, 1973) considered this problem of subordinal
relationships in a broader context, utilizing the rich collections of fossils from
various Russian deposits. Although he defined the Archostemata very broadly
to include all of Mesozoic fossils, rendering the concept of little use in a cladis-
tic sense, his ideas about the relationships among modern taxa and especially
the diversity among their Mesozoic precursors are enlightening. Ponomarenko
described two types of wing folding in beetles: the cupedoid type found in most
typical archostematan fossils, modern Cupedidae and Ommatidae, polyphagan-
800
like fossils in the Ademosynidae, and Polyphaga; and the schizophoroid type
found in the fossil taxa Schizophoridae and Catiniidae plus Adephaga, Myxo-
phaga and Micromalthidae. He agreed with Forbes (1926) that myxophagans
are m~ch closer to Adephaga than they are to Polyphaga. Kirejtshu~ (1992)
schematically illustrated the possible relationships between these fossll groups
and modem taxa. Kukalova-Peck and Lawrence (1993) also concluded that
Adephaga and Myxophaga were closely relat~d, but they .considered the micro-
malthid wing to be derivable from a more typical cuped01d type. These authors
also suggested that modern Archostemata, Adephaga and Myxophaga form a
monophyletic group, which is the sister group of t~e Polyphaga. If the pre~ence
of cervical sclerites is considered to be ancestral m Coleoptera, then theu ab-
sence in these three suborders is a synapomorphy.
A few more schemes have been proposed for the major breakdown of the
order, but most of these can hardly be taken seriously. King (1956) introduced
the suborder Apicalae to include only the genus Atractocerus (see Forbes 1956,
Selander 1959). Machatschke (1962) considered Archostemata to be a separate
order which with Coleoptera comprise a superorder Elytraria. Abdullah (197 4b)
used the term Entomophaga for Strepsiptera, assuming that the latter group
needed a new name if it were included within the order Coleoptera. Iablokoff-
Khnzorian (1983) added two suborders, Archaeogastra (for Hydraenoidea, Hy-
drophiloidea and Staphylinoidea) and Heterogastra (everything else), but ex-
cluded Myxophaga from his scheme. Ienistea (19~6) proposed more. than 80
orders for beetles, raising almost every supragenenc taxon to the family level.
Suborder ARCHOSTEMATA Kolbe, 1908: 246
*Although the Recent taxa comprising this suborder have been generally ~greed
upon at least since the publications of Forbes (1926) and Boving ~nd Craighead
(1931), Micromalthidae were still considered to be polyphagan m some more
recent works (see below). Ponomarenko (1969) defined Archostemata very
broadly, including in it all Permian taxa and those Triassic and later forms
possessing a suite of primitive features, .sue? a~ the well-developed ext~I?a~
propleuron and the metepisterna broadly Impmgi~g on the mesoco~al cavities,
however, he recognized the diversity of these fosstl forms and c~nstdered them
to belong to three distinct groups: the ancestral Tshekar:docoletdae, the cupe-
doid branch (Permocupedidae, Cupedidae, Taldycupedtdae, and Ademosyn-
idae ), and the schizophoroid branch (Asiocoleidae, Tricoleidae, Rhombocole-
idae, Schizophoridae, Catiniidae, and Micromalthidae). Crowson (1975) sug-
gested that the Lower and Upper Permian taxa be placed in the suborders Pro-
tocoleoptera and Archecoleoptera, respectively; and Kukalova-Peck (1991)
placed these forms in a separate order Protocoleoptera. Cr~"":'~on (19_75) also
pointed out that some Triassic fossils, particularly m the Catmudae, might rep-
801
resent the ancestors of modem Myxophaga. Atkins (1963), Monr6s and Monr6s
(1952), Vulcano and Pereira (1975).
OMMATIDAE Sharp and Muir, 1912: 521 (=Ommadidae; incl. Tetrapha-
leridae)
*Crowson (1976) placed Tetraphalerus in a separate family with there-
markable subterranean form Crowsoniella relicta Pace (see below). In addi-
tion to the characters cited by Crowson (1976), Omma and Tetraphalerus
share a distinctive feature of the apical maxillary palpomere, namely the
enclosure of specialized sensilla ("digitiform appendages" of Crowson 1976;
"chemosensory setae" of Watt 1974a) in a deep cavity (Figs. 1, 2) somewhat
similar to that in Micromalthus (Fig. 3) but lacking in Cupedidae (Fig. 4) or
Crowsoniellidae. A larva collected in soil near Perth, Western Australia, and
tentatively assigned to Omma sagitta Neboiss has most of the characteristics
of other known archostematan larvae, including the ligular sclerome, but
there are 4 distinct stemmata on each side, paired endocarinae, rather than
a median one, and no spine-like process on tergite 9. Crowson (1962), Law-
rence et al. (1993), Neboiss (1989).
TETRAPHALERINAE Crowson, 1962: 152
Tetraphalerus (South America).
OMMATINAE Sharp and Muir, 1912: 521
Omma (Australia).
CROWSONIELLIDAE Iablokoff-Khnzorian, 1983: 65
Crowsoniella (Italy).
*Iablokoff-Khnzorian was the first to make this family name available,
although Crowson (1976) stated that the genus Crowsoniella "will require
at least a tribe of its own". A relationship to Ommatidae is indicated by the
fusion of the labrum to the clypeus, laterally inserted antennae, contiguous
procoxae, simple tarsi and non-overlapping ventrites; however, all of these
features appear to be shared with Micromalthidae as well. The inclusion of
this genus in Tetraphaleridae was based in part on presence of dorsal anten-
na! grooves on the head in Crowsoniella and two Mesozoic Tetraphalerus;
however, these grooves are not obvious in figures of the latter (Ponomarenko
1969: 99, figs. g, d). Pace (1976).
MICROMALTHIDAE Barber, 1913a: 185
Micromalthus (eastern North America, introduced widely- Hong Kong,
Hawaii, South America, South Africa, etc.).
*The lack of sutures on the prothorax and an oblongum cell in the wing
have led some workers (Machatschke 1962, Arnett 1963, Klausnitzer 1975,
Hennig 1981) to place this highly specialized group in Polyphaga. Not only
802
Figures 1-4. Apical maxillary palpomeres in Archostemata, showing. location of speciali:zed digiti-
form sensilla. (1) Omma stanleyi Newman. (2) Tetraphalerus wagnen Waterhouse. (3) Mtcromaltus
debilis LeConte. (4) Priacma serrata (LeConte). Scale lines: 1, 2, 4 = 0.1 mm; 3 = 0.01 mm.
803
are the larval characters indisputably of the archostematan type (see Lawrence
1991), but the wing has a distinct medial hinge as in Archostemata, Adephaga
and Myxophaga (Emden 1932a; Kukalova-Peck and Lawrence 1993) and the
male genitalia resemble those of Cupedidae and Ommatidae in the shape of the
pregenital (9th) segment and the lack of a phallobase. The unusual life cycle
involving paedogenesis, as well as thelytokous, amphitokous and arrhenotokous
parthenogenesis, has been described by Barber (1913a, 1913b), Pringle (1938)
and Scott (1936, 1938, 1941). The species has been widely introduced via the
first-instar larvae, which occur in dead wood; such larvae are known from Cre-
taceous (Lebanon), Oligocene (Baltic), and Miocene (Mexico) amber (Rozen
1971; Crowson 1981; Lawrence unpublished).
CUPEDIDAE Laporte, 1836: 56 (=Cupesidae)
Adinolepis (Australia), Ascioplaga (New Caledonia), Cupes (North Amer-
ica), Distocupes (Australia), Paracupes (Brazil, Ecuador), Priacma
(western North America), Prolixocupes (western USA, southern South
America), Rhipsideigma (eastern Africa, Madagascar), and Tenomerga
(North America, eastern Asia, western Pacific, South Africa).
*Ponomarenko (1969) followed Crowson (1962) in placing Priacma in a
separate tribe from the remaining genera. Neboiss (1960, 1984) split the
genus Cupes into seven genera. Galian and Lawrence (1993) pointed out that
Distocupes varians (Lea) and Priacma serrata LeConte had follicular (rather
than tubular) testes and that the former had a male diploid chromosome
number of 19 (9 + X) and exhibits achiasmatic meiosis, previously recorded
only in Adephaga. The reports of tubular testes in the cupedid Prolixocupes
latreillei (Solier) and the ommatid Tetraphalerus wagneri Waterhouse (Vidal
Sarmiento 1969) need to be verified. Neboiss (1989).
Suborder MYXOPHAGA Crowson, 1955: 10
*Crowson proposed this suborder for four rather obscure taxa, previously
placed among the Hydrophiloidea (Lepicerus and Hydroscapha), Staphy-
linoidea (Sphaerius) and Clambidae (Calyptomerus), sharing a type of pro tho-
rax and hind wing characteristic of Archostemata and Adephaga combined with
a polyphagan type of larva (having a 5-segmented leg and asperate mandibular
mola). Forbes had previously (1926) placed these groups in Adephaga based
on wing venation and folding . Calyptomerus was later excluded from the sub-
order by Crowson (1960, 1968) and the genus was included in Clambidae in
EndrOdy-Younga's (1959) revision. The only myxophagan larva known to
Crowson was that of Hydroscapha (Boving 1914), but in 1964, Steffan de-
scribed both adult and larva of a new African genus and species (Torridincola
rhodesica) representing a new family of Myxophaga, and in 1966, Britton dis-
804
covered and described the larva of Sphaerius. Although these new larvae were
each of a distinctive type, they shared with the Hydroscapha larva the reduced
polyphagan leg and asperate mola and in addition had similar habits (aquatic
algal grazers) and abdominal tracheal gills (Hinton 1967b). In the years to fol-
low a number of new taxa were discovered, particularly in Torridincolidae,
which is now known to be the largest and most diverse of the myxophagan
families. The suborder is not universally recognized. Barlet (1972) concluded
on the basis of prothoracic structure that Hydroscapha should be placed in
Adephaga and Sphaerius in Polyphaga; in 1974 the same author suggested t~at
Hydroscapha represented a transition between Cupes and Adephaga, while
Torridincolidae represented a similar transition from Cupes to Polyphaga (see
Hlavac 1973, 1975). Ponomarenko (1969, 1973) noted the similarities between
extant myxophagans and members of the Mesozoic families Schizophoridae
and Catiniidae, and Lawrence and Newton (1982) suggested that these Recent
taxa could be isolated relicts of an early schizophoroid radiation.
LEPICERIDAE Hinton, 1936a: 473 (1882) (=Cyathoceridae)
Lepicerus (=Cyathocerus) (Mexico to northern South America).
*Reichardt (1973a, 1976d).
**Cyathoceridae Sharp (1882: 141) has priority over Lepiceridae which
was proposed to replace it because of the generic synonymy, but retention
of Lepiceridae has been justified under Art. 40b (Reichardt 1976d).
TORRIDINCOLIDAE Steffan, 1964: 199
Claudiella (Brazil), Delevea (South Africa and Japan), Hintonia (=Ptyo-
pteryx) (Brazil), lncoltorrida (Madagascar), Torridincola (southern
Africa), and Ytu (Brazil).
*Barlet (1974), Hinton (1969), Mesa and Fontanetti (1985), Reichardt
(1973a, 1976b), Reichardt and Costa (1967), Reichardt and Vanin (1976,
1977), Sat6 (1982), Spangler (1980b), Steffan (1964, 1973).
HYDROSCAPHIDAE LeConte, 1874: 45
Hydroscapha (North America, Eurasia, north Africa, Madagascar), Sca-
phydra (Brazil), and Yara (Panama, Brazil).
*Barlet (1972), Boving (1914), Orchymont (1945), Reichardt (1973a,
1973b, 1974), Reichardt and Hinton (1976).
MICROSPORIDAE Crotch, 1873b: 78/1873c: 27 (=Sphaeriidae)
Microsporus (=Sphaerius) (North and Central America, Eurasia, Madagas-
car and Australia).
*Barlet (1972), Britton (1966), ICZN (1985b), Reichardt (1973a).
**The family name Sphaeriidae Erichson (1845: 38) and its type genus
Sphaerius were rejected in favor of Microsporidae Reichardt (1976a: 204)
805
and Microsporus by Opinion 1331 (ICZN 1985b) to avoid confusion with
names in Mollusca. The name Microsporidae should actually be credited to
Crotch's (1873b, 1873c) much earlier use.
Suborder ADEPHAGA Schellenberg, 1806: 3
*The phylogenetic relationships of the few families of Adephaga probably have
been more intensively studied in recent years than those of Archostemata, My-
xophaga or any of the polyphagan superfamilies. Many of the recent phylo-
genetic hypotheses have resulted from or been part of a large series of compar-
ative morphological studies, leading to an expansion of the available suite of
taxonomic characters. These works, some of which have a functional emphasis,
include those dealing with the following adult systems: head - Belkaceme
(1991), Beutel (1986a, 1989a); mouthparts and feeding mechanisms- Acorn
and Ball (1991), Evans and Forsythe (1985), Forsythe (1982); antenna cleaner
- Hlavac (1971), Regenfuss (1975); thorax and locomotory mechanisms -
Baehr (1979a, 1979b), Belkaceme (1986, 1991), Bell (1967), Beutel (1986a,
1989b, 1990a, 1990b, 1992a), Beutel and Belkaceme (1986), Evans (1977,
1980, 1983, 1985), Hlavac (1973, 1975), Larsen (1966); hind wing- Balfour-
Browne (1943), Forbes (1926), Hammond (1979), Kukalova-Peck and Law-
rence (1993), Ward (1979); abdominal apex and ovipositor- Bils (1976), Bur-
meister (1976, 1980, 1990a, 1990b), Deuve (1988, 1993); abdominal defensive
glands- Dettner (1985), Forsyth (1968, 1970, 1972); defensive chemistry-
Aneshansley et al. (1983), Dettner (1979, 1985), Eisner et al. (1977), Moore
(1979); internal reproductive organs -Ali (1967), Smd (1981); alimentary
canal and nervous system- Ali (1967), Balfour-Browne (1944), Smd (1982);
and karyotype- Galian and Moore (1994), Serrano (1981); as well as those
dealing with larval morphology (especially chaetotaxy and head morphology):
Alarie (1991), Alarie and Harper (1990), Alarie et al. (1990), Arndt (1993),
Beutel (1986b, 1991, 1992b, 1992c, 1993a, 1993b), Bousquet and Goulet
(1984), Landry and Bousquet (1984), Nilsson (1987), Ruhnau (1986), Spangler
(1991), and Wolfe and Roughley (1985).
One of the more important advances in the understanding of adephagan phy-
logeny centers around the relationships of the aquatic families, which until
relatively recently were treated as a monophyletic assemblage - the Hydrade-
phaga. Crowson (1950, 1955) was apparently the first to recognize that the
vicariant genera Trachypachus and Systolosoma, placed by Jeanne} (1941) in
the section lsochaeta of Carabidae, had features of the antenna and metacoxa
which were more consistent with their inclusion in Hydradephaga as a separate
family. Crowson (1960) considered the Hydradephaga to have been derived
from the Geadephaga (i.e., terrestrial families of Adephaga) as a result of a
single invasion of the aquatic habitat, Gyrinidae to be a derived member of the
dytiscoid lineage, and Trachypachidae to be a remnant of the geadephagan
806
group which gave rise to Hydradephaga. Bell (1966) suggested that the Hydra-
dephaga represent three independent lineages, Haliplidae, Gyrinidae and the
dytiscoid complex, each representing a separate aquatic adaptation and each as
closely related to Trachypachidae as it is to the other lineages. He proposed the
term Glabricornia for a monophyletic group consisting of Trachypachidae plus
Hydradephaga. Ancestral Glabricornia, according to Bell, were derived from
terrestrial Isochaeta, from which the anisochaetous Carabidae had already sepa-
rated. Ponomarenko (1973, and in Arnoldi et al. 1977) also supported the three-
lineage hypothesis, but he maintained that these represented, not independent
aquatic derivatives from a terrestrial ancestor, but rather three different "paths
in the perfection of the aquatic mode of life already acquired by a common
ancestor somewhere close to the boundary of the Permian and Triassic." Gea-
dephaga were considered to be monophyletic, with a split between Trachypach-
idae and Carabidae occurring early in the Mesozoic . His conclusions were
based in large part on the existence in the Triassic of possible haliplid and
gyrinid ancestors (Triaplus and Triadogyrus, respectively), as well as a number
of early Mesozoic trachypachids (Eodromeinae) and carabids (Protorabinae).
A comparison of these three scenarios with two earlier ones is made by Bell
(1983). Nichols (1985) considered all Hydradephaga to have evolved from
within the Geadephaga, with the carabid tribe Omophronini as their sister
group. Kavanaugh (1986) proposed another scheme, in which halipli~s ~e the
sister group of the Geadephaga, Amphizoidae is the most basal family m the
dytiscoid lineage, and Gyrinidae are derived dytiscoids. Beutel and Roughley
(1988) proposed a phylogeny in which Gyrinidae are the sister group of other
Adephaga, but Haliplidae and other hydradephagans evolved from within the
Geadephaga. Burmeister (1990a) concluded that Noteridae and Haliplidae were
sister groups and that these formed a monophyletic group with Gyrinidae. In
Beutel's most recent scheme (1995) both Gyrinidae and Haliplidae are basal
to the lineage comprising Trachypachidae, the dytiscoids and the Geadephaga.
**According to Horn and Schenkling (1928-29), Schellenberg is the author
of the original text in which the name Adephaga first appeared; Clairville, au-
thor of the French translation appearing on facing pages of the same book, has
usually been credited with the name.
GYRINIDAE Latreille, 1810: 168
TRACHYPACHIDAE C.G. Thomson, 1857: 5 [not seen; after Madge 1989:
*Although Sharp (1899) and Sharp and Muir (1912) separated this family
from the remaining Adephaga on the basis of the primitive aedeagal struc-
ture, most authors have placed it at the end of the dytiscoid complex. Hatch
(1925) commented that "nothing is found in the latter (Gyrinidae) that cannot
be derived from the former (Dytiscidae)", and Crowson (1960) stated that
if the special features of gyrinids were discounted "there is really very little
to separate Gyrinidae from Noteridae or Dytiscidae." As indicated above,
recent studies of both recent and fossil Adephaga suggest that this group
807
may well occupy a basal position within the suborder. Beutel (1990a, 1995),
Beutel and Roughley (1988, 1994), Burmeister (1990a), Franciscolo (1979),
Ponomarenko in Arnoldi et al. (1977), Tranda (1972).
SPANGLEROGYRINAE Folkerts, 1979: 7
Spanglerogyrus (USA: Alabama).
*This remarkable genus possesses a number of primitive features mak-
ing ~t less likely that gyrinids evolved from dytiscoid ancestors and sup-
portmg the hypothesis that the family is the sister group of the remaining
Adephaga. Beutel (1989a, 1989b, 1990a), Burmeister (1990a), Folkerts
(1979), Lawrence and Newton (1982).
GYRININAE Latreille, 1810: 168 (incl. Orectochilinae)
Aulonogyrus, Dineutes, Enhydrus, Gyretes, Gyrinus, Heterogyrus, Macro-
gyrus, Orectochilus, Orectogyrus, etc.
*Generic phylogenies have been proposed by Beutel (1990a) and Beu-
tel and Roughley (1994).
HALIPLIDAE Aube, 1836: 15
Algophilus (southern Africa), Apteraliplus (western North America), Bry-
chius (Europe, USA: California), Haliplus (widespread), and Peltodytes
(North America, Eurasia, Madagascar).
*Haliplidae appear to represent an early adephagan offshoot, and their
feeding habits (algophagy) may reflect those of a a pre-adephagan ancestor
resembling present day Myxophaga. The suctorial mouthparts of the larva
could be interpreted as a secondary adaptation for algophagy in a preda-
tory ancestor, or a primary adaptation in an algophagous ancestor, which
served as a preadaptation for predation in the majority of Adephaga. These
relationships may be clarified after a study is completed of an undescribed
adephagan from South America, which has a number of haliplid-like fea-
tures but lacks the enlarged metacoxal plates characteristic of that group
(P.J. Spangler, unpublished). Belkaceme (1986), Beutel (1986b, 1995),
Beutel and Ruhnau (1990), Franciscolo (1979), Seeger (1971).
. **Brulle (1835) has been cited frequently for the family name, e.g., by
S1lfverberg (1992), but we can find no such name in that work.
468]
Systolosoma (Chile, Argentina) and Trachypachus (Holarctic).
*As discussed above, this taxon is generally considered to form a mo-
nophyletic group with members of the dytiscoid assemblage. Bell (1966,
1983), Beutel (1993a, 1995), Beutel and Belkaceme (1986), Lindroth
(1960), Roughley (1981).
808
NOTERIDAE C.G. Thomson, 1860: 34 (incl. Phreatodytidae) .
*This and the following three families are generally constdered to form
a monophyletic group, but many workers would include the Trachypach-
idae and/or Gyrinidae as well. Belkaceme (1991), Beutel (1995), Beutel
and Roughley (1987), Franciscolo (1979), Ueno (1957). . Vatellus, etc.
**Latreille (1825) has been cited in error as author of the famtly name.
PHREATODYTINAE Ueno, 1957: 251
Phreatodytes (Japan).
NOTERINAE C.G. Thomson, 1860: 34
Canthydrus, Hydrocanthus, Hydrocoptus, Noterus, Notomicrus, Prono-
terus, Suphis (=Colpius), Suphisellus, etc.
AMPHIZOIDAE LeConte, 1853: 228
Amphizoa (western North America and China).
*Beutel (1986a, 1991, 1995), Burmeister (1990b), Kavanaugh (1986),
Yu and Stork (1991).
HYGROBIIDAE Regimbart, 1878: 450 (1837) (=Pelobiidae)
Hygrobia (=Pelobius) (Australia and the Palearc~ic).
*Beutel (1986a, 1995), Britton (1981), Franctscolo (1979) ...
**Pelobiidae Erichson (1837: 182) has priority over Hygrobudae, but
the latter name is much more widely used and its retention can be justified
by Art. 40b. According to Neave (1939-40), the original spellings of the
type genera are Hygriobia and Paelobius; the former spellmg was emen-
ded to Hygrobia by the ICZN (1954).
DYTISCIDAE Leach, 1815: 84 . 19th Century classifications, but Redtenbacher ( 1886) showed that the hind
*Morphological studies by Beutel (1994, 1995), Bur~etster (1976),
Ruhnau and Brancucci ( 1984) and others have drawn attentiOn to problems
with the traditional subfamily classification of dytiscids, reflected by the
removal of Copelatini and Agabetini from Colymbetinae to form a basal
subfamily and a tribe of Laccophilinae, respectively. However, the formal
placement of other odd colymbetine genera such as Hydrotrupes and La~­
cetes, and relationships of Aubehydrinae, have not been reso!ved. Alarte
(1991), Alarie and Harper (1990), Alarie et al. (1990), Burmetster (1976),
Franciscolo (1979), Nilsson (1987), Nilsson et al. (1989), Ruhnau and
Brancucci (1984), Wolfe and Roughley (1985), Zaitsev (1953) ..
**"Degeer, 1774" has been cited in error as author of the famtly name.
COPELATINAE Van den Branden, 1885: 82
Agaporomorphus, Aglymbus, Copelatus, and Lacconectus.
LACCOPHILINAE Gistel, 1856: 354
Agabetes, Laccodytes, Laccophilus, Neptosternus, etc.
809
HYDROPORINAE Aube, 1836: 217
Bidessus, Carabhydrus, Celina, Clypeodytes, Coelambus, Derovatellus,
Desmopachria, Hydroporus, Hydrovatus, Hygrotus, Hyphydrus, Lac-
cornis, Methles, Pachydrus, Phreatodessus, Potamonectes, Siettitia,
*Wolfe (1985), Wolfe and Roughley (1990).
**Latreille (1825) has been cited as author of the subfamily name, but
we can find no family-group name in connection with the generic treat-
ment of Hydroporus there.
COLYMBETINAE Erichson, 1837: 149
Agabus, Anisomeria, Colymbetes, Coptotomus, Hydronebrius, Hydrotru-
pes, Ilybius, Lancetes, Matus, Platynectes, Rhantus, etc.
DYTISCINAE Leach, 1815: 84
Acilius, Cybister, Dytiscus, Eretes, Hydaticus, Hyderodes, Thermonetus,
etc.
AUBEHYDRINAE Guignot, 1942: 11
Notaticus (=Aubehydrus) (Brazil).
*This genus has been placed in a separate subfamily by Spangler
(1973), Franciscolo (1979) and Nilsson et al. (1989), but Notaticus was
originally placed with doubt in Hydaticini (Dytiscinae), and was listed
there by Wolfe (1985).
RHYSODIDAE Laporte, 1840a: 291
Clinidium, Dhysores, Kaveinga, Leoglymmius, Medisores, Omoglymmius,
Rhysodes, Rhyzodiastes, Sloanoglymmius, etc.
*This family was placed near Cucujidae or Colydiidae in almost all
wing was of the adephagan type and Ganglbauer (1892) placed it in his
Caraboidea. Crowson (1950) considered this family to be the sister group
of the remaining Adephaga, but in 1960 he noted that, if their special mod-
ifications were discounted, rhysodids have almost all of the features of
Jeannel's (1941) Simplicia (see under Carabidae below). Bell and Bell
(1962) considered this group to be a highly specialized offshoot of the
Carabidae, possibly related to Scaritini. Erwin and Sims (1984) and Erwin
(1985) considered the group to be related to psydrine and trechine Carab-
idae. Beutel (1990b, 1992a) found that thoracic structures did not support
a basal position of rhysodids within Adephaga and pointed to a possible
relationship with Scaritini, but the same author (1992b, 1993a, 1995) sug-
gested a carabid-rhysodid sister group relationship based primarily on
larval head characters. Bell (1970, 1994), Bell and Bell (1978, 1979, 1982,
1985, 1987, 1991, 1993), Burakowski (1975a), Hincks (1950).
810
CARABIDAE Latreille, 1802: 80 (incl. Cicindelidae, Paussidae, etc.)
*Since Bonelli (1810) first divided the family into three major groups,
Simplicimani, Integripennes and Truncatipennes, the higher classification
of Carabidae has been in a healthy state of flux (although the adjective
might be disputed by some). A detailed account of the history of carabid
classification is given by Ball (1979). Probably the most influential scheme
was that of Jeannel (1941, 1942), whose concepts are still used by most
carabid workers, although many of his phylogenetic hypotheses are no
longer supported. He divided the group into the Isochaeta (those with both
tibial spurs on the foreleg arising at the apex) and the Anisochaeta (those
with one of these spurs displaced by the development of an antennal
cleaner). The second group was subdivided into Simplicia (without visible
metepimera) and Limbata (those with visible metepimera). Finally, the
Limbata was divided into four groups (Scrobifera, Stylifera, Conchifera
and Balteifera) based on such features as the lateral closure of the meso-
coxal cavities, the presence or absence of a seta in the mandibular scrobe
and the structure of the aedeagus. Jeannel' s system of ranking taxa (almost
50 families in his 1949 version) was almost universally rejected by later
workers. Some of the more recent classifications of the group are given in
Ball (1960), Bell (1967), Bousquet and Larochelle (1993), Casale et al.
(1982), Crowson (1950, 1955), Darlington (1952-71), Erwin (1985, 1991),
Erwin and Sims (1984), Kryzhanovsky (1976), Lindroth (1945-49,
1961-69), and Reichardt (1977). The removal of Trachypachus and Sy-
stolosoma from the family is discussed above. The scheme given here is
based in part on recent morphological works by Deuve (1988, 1993) and
Beutel (1992a, 1992b, 1992c, 1993a, 1993b, 1995) (see Adephaga above).
Other references: Boer et al. (1986), Desender et al. (1994), Larochelle
(1990), Machado (1992), Noonan et al. (1992), Stork (1990), Thiele
(1977), Trautner and Geigenmtiller (1987).
**Madge (1989) thoroughly reviewed family-group names of carabids
and other terrestrial Adephaga but did not apply them to a classification.
PAUSSINAE Latreille, 1807a: 234
Arthropterites, Arthropterus, Cerapterus, Ceratoderus, Cicindis, Eopaus-
sus, Eustra, Heteropaussus, Hexaplatarthrus, Homopterus, Hylotorus,
Leleupaussus, M egalopaussus, M etrius, Mystropomus, Nototylus, Ozae-
na, Pachyteles, Paussus, Pentaplatarthrus, Physea, Platyrhopalus, Pro-
topaussus, Pseudozaena, etc.
*Beutel (1992c), Bousquet (1986), Darlington (1950), Deuve (1994),
Luna de Carvalho (1989, 1992), Stork (1985).
GEHRINGIINAE Darlington, 1933: 110
Gehringia (northwestern North America).
*Jeannel (1941) placed this unusual genus in his Isochaeta, but Bell
(1964) transferred it to the Anisochaeta based on a unique interpretation
811
of the protibial spurs disputed by Hlavac (1971). Lindroth (1960), Ball
(1960), and Kryzhanovsky (1976) included Gehringia in Trachypachini or
Trachypachinae; however, Bell (1966) pointed out that the condition of the
metacoxae in this genus is connected with the lateral displacement of the
legs and not the great enlargement of the coxae as in trachypachines .
Erwin and Sims (1984) placed the tribe Gehringiini in their supertribe
Psydritae, and Maddison (1985) noted similarities to Trechitae (both in
Trechinae below).
OMOPHRONINAE Bonelli, 1810: Tabula synoptic a
Omophron (Eurasia, Africa, North and Central America).
*On the basis of a cladistic analysis involving characters of the coxal
cavities, Nichols (1985) considered Omophron to be the sister group ofthe
Hydradephaga (including Trachypachidae) and derived both from a primi-
tive carabid stock, including Notiophilini, Nebriini and Carabini. Bils
(1976) came to a similar conclusion based on features of the female ab-
dominal apex. Landry and Bousquet (1984) .
CARABINAE Latreille, 1802: 80
Aplothorax, Calosoma, Carabus, Ceroglossus, Cychrus, Nebria, Notioka-
sis, Notiophilus, Opisthius, Pamborus, Scaphinotus, etc.
*Kavanaugh and Negre (1983), Moore (1966).
CICINDELINAE Latreille, 1802: 77
Amblycheila, Cicindela, Collyris, Ctenostoma, Distipsidera, Mantichora,
Megacephala, Omus, Pogonostoma, Pycnochila, Therates, Tricondyla,
etc.
*The genus Cicindela was broken up into 55 genera by Rivalier (1963).
Pearson (1988), Wiesner (1992).
HILETINAE Schibdte, 1847: 69
Eucamaragnathus (Asia, Africa, Madagascar, South America) and Hiletus
(Africa).
*Erwin and Stork (1985).
LORICERINAE Bonelli, 1810: Tabula synoptica
Loricera (North and Central America, Eurasia).
*Ball and Erwin (1969) .
ELAPHRINAE Latreille, 1802: 81
Blethisa, Diacheila, and Elaphrus (Holarctic).
MIGADOPINAE Chaudoir, 1861: 510
Amarotypus, Antarcticonomus, Migadops , Monolobus, Nebriosoma,
Stichonotus, etc. (Australia, New Zealand, southern South America) .
*Jeannel (1938).
SIAGONINAE Bonelli, 1813: 456
Enceladus (northern South America), Luperca (Africa, India), and Siagona
(Eurasia, Africa).
*Erwin (1978).
812
SCARITINAE Bonelli, 1810: Tabula synoptica
Carenum, Clivina, Conopterum, Dyscherus, Dyschirius, Euryscaphus, For-
cipator, Mouhotia, Pasimachus, Promecognathus, Salcedia, Scaraphi-
tes, Scarites, Storthodontus, etc.
*This corresponds to Jeannel's (1941) Scrobifera excluding Hiletinae.
Bousquet and Smetana (1986) could find no support for the inclusion of
Promecognathus based on larval characters. Moore and Lawrence (1994)
described a very different larval type in the Australian carenines, which
calls into question the monophyly of this subfamily. Baehr (1979a, 1980).
TRECHINAE Bonelli, 1810: Tabula synoptica
Apotomus, Barypus, Bembidion, Broscus, Creobius, Cymbionotum, Mecy-
clothorax, Melaenus, Merizodus, Nomius, Patrobus, Pericompsus, Peri-
leptus, Pogonus, Promecoderus, Psydrus, Stylulus, Tachys, Trechus,
Tropidopterus, etc.
*This corresponds to the Stylifera of Jeanne} (1941) and is combined
with the next group by some workers. Jeanne} (1926-30).
HARPALINAE Bonelli, 1810: Tabula synoptica
Acupalpus, Aephnidius, Agonica, Agonum, Agra, Amara, Amblystomus,
Anisodactylus, Anthia, Calathus, Callistus, Calophaena, Chlaenius,
Cnemalobus, Colliuris, Colpodes, Ctenodactyla, Cuneipectus, Drypta,
Feronia, Galerita, Geopinus, Glyptus, Graphipterus, Harpalus, Hellu-
odes, Helluo, Hexagonia, Lachnophorus, Lebia, Licinus, Loxandrus,
Masoreus, Morion, Mormolyce, Odacantha, Orthogonius, Panagaeus,
Pelecium, Pentagonica, Pericalus, Perigona, Platynus, Pterostichus,
Sarothrocrepis, Tetragonoderus, Thyreopterus, Zuphium, etc.
*This corresponds to the Conchifera of Jeanne} (1941). A number of
recent authors, such as Deuve (1988), place Pseudomorphinae here. Lieb-
herr and Ball (1990), Noonan (1973), Straneo and Ball (1989).
**Graphipterinae Latreille (1802: 83) has priority for the name of this
group, but Harpalinae is a well-known name that should probably be con-
served.
CARABIDAE incertae sedis:
PSEUDOMORPHINAE Newman, 1842: 365
Adelotopus and Cryptocephalomorpha (Oriental, Australian); Cainogenion,
Paussotropus, and Sphallomorpha (=Silphomor:pha) (Australian); and
Pseudomorpha (New World). ~ soniid~e are somewhat doubtfully associated with o~e of the larger ~ssemblages
*Although Notman (1925) considered pseudomorphineS'to form a dis-
tinct family, based on the unusual adult form, most workers (B\~el 1992a,
Bils 1976, Deuve 1988) have placed them among the derive Limbata
(Jeannel 1941). Moore (1974) placed them in a distinct subfami )(based
on the unique larvae of Sphallomorpha, which live in vertical burro\\zs and
have adaptations resembling those of tiger beetles. Erwin (1981) also dis-
cussed the distinctiveness of the group, based on a second myrmecophilous
\
813
Ia:val type found in Pseudomorpha and Adelotopus, and later (Erwin and
S1ms 1984, Erwin 1985) included them in his Division Loxomeriformes,
along with brachinines, paussines, scaritines, elaphrines and several other
relatively primitive taxa. Liebherr and Kavanaugh (1985) reported ovovivi-
parity in this group, which is known in no other Carabidae. Baehr (1992
1994). '
BRACHININAE Bonelli, 1810: Tabula synoptica
Aptinus, Brachinus, Crepidogaster, Mastax, Pheropsophus, Styphlomerus,
etc.
. *This subfamily was first placed in the Truncatipennes, the most de-
nved .group o~ carabids containing the Lebiini, but Jeanne! (1941, 1942)
c?~b1~e.d 1t.w1th Pseudo~orphinae in a distinct group Balteifera, based on
s1mllant1es m the abdommal apex and aedeagus. In his work on the female
abdominal apex, Bils (1976) also considered both brachinines and pseu-
domorphines to be among the derived Carabidae, but did not consider
them to be sister groups. The occurrence of a similar defensive mechanism
involving crepitation was considered by most carabid workers to have
evolved independently in the Paussinae and Brachininae. Moore and Wall-
bank (1968) found the chemical composition of the defensive secretions
to be similar in paussines and brachinines, but not in Pseudomorphinae;
they rejected Jeannel's inclusion of the last group in Balteifera, but con-
cl~d~d that ~ther I?orphological evidence did not support a paussine-bra-
chmme relat10nsh1p. Eisner and his colleagues (Eisner et al. 1977, Anes-
hansley et al. 1983) considered the similarities in crepitating mechanisms
and defensive chemicals to be sufficient to unite Paussinae and Brachin-
inae, and this idea was followed by Erwin and Sims (1984), Erwin (1985)
and Deuve (1988). In two recent papers dealing with the adult thorax
(1992a) and larval head (1993a), Beutel placed the brachinines among the
most derived carabid assemblage (Jeannel's Limbata), but did not relate
them to any particular group within this complex. Erwin (1970).
Suborder POLYPHAGA Emery, 1886: 654
*The relationships among the major polyphagan lineages are still not clearly
understood, and some groups, such as Scirtoidea Derodontidae and Jacob-
(Elatenformia or Bostrichiformia).
Series STAPHYLINIFORMIA Lameere 1900: 373
*This n~~ was first proposed for the present grou~ excluding Palpicomia
(Hydroph1hdae and Hydraenidae), but Forbes (1926) and Boving and Crai-
ghead (1931) provided strong evidence for including palpicoms and the name
has been uniformly applied to the larger group since Lameere (1938) and
814
Crowson (1960 and later). Thus delimited, staphyliniforms appear to be one of
the most basal groups of Polyphaga, with articulated larval urogomphi as one
of their few derived features (Crowson 1960, Lawrence and Newton 1982). The
main remaining question concerning the composition of this group is whether
scarabaeoids should be added, forming the traditional group Haplogastra (see
further discussion under Scarabaeiformia).
Within Staphyliniformia, four groups have been widely recognized as mono-
phyletic: Hydrophilidae (sensu lato, sometimes divided into six families); Hy-
draenidae; "Histeroidea" (Sphaeritidae, Synteliidae, and Histeridae); and Sta-
phylinoidea (with or without Hydraenidae). However, two mutually exclusive
interpretations of similarities among these groups have resulted in two alterna-
tives for combining them into superfamilies. The mainly aquatic groups (Hy-
drophilidae and Hydraenidae), historically associated as Palpicomia, are often
combined into a superfamily Hydrophiloidea because of unique aquatic adapta-
tions of adults and some other characters; the terrestrial groups Histeroidea and
Staphylinoidea (excluding Hydraenidae), which have little in common beyond
the general characters of the series, then form two other superfamilies (e.g.,
Crowson 1955, 1960). Alternatively, several suites of characters (highly derived
larval mouthparts and body form, wing venation/folding, genitalia and internal
organs) support a close relationship of Hydrophilidae and Histeroidea to the
exclusion of Hydraenidae, which have all derived characters of Staphylinoidea
and specifically share a number of unique features with Ptiliidae (e.g., Boving
and Craighead 1931, Forbes 1926, Lawrence and Newton 1982). This second
set of relationships supports the recognition of two monophyletic superfamilies,
Hydrophiloidea (Hydrophilidae plus Histeroidea) and Staphylinoidea (including
Hydraenidae). We continue to prefer this two-superfamily arrangement as best
supported by overall evidence, leaving open the question of the origin of
aquatic habits in staphyliniforms (independently derived in Hydrophilidae and
Hydraenidae, or ancestral for the series and independently lost in "histeroids"
and most staphylinoids?). Fossils clearly attributable to hydrophilids, hydra-
enids, staphylinids and probably agyrtids are known from the Jurassic and well
represented in Mesozoic deposits (e.g., Ponomarenko in Arnoldi et al. 1977,
Ponomarenko 1985, Ponomarenko and Ryvkin 1990, Ryvkin 1985).
"Histeroids", which should be recognizable from even fragmentary fossils, are
not known until the mid-Tertiary, although the Lower Cretaceous genus Mesy-
dra, placed in Hydrophilidae (Ponomarenko in Arnoldi et al. 1977), has charac-
ters of body form, abdomen, elytra, and position of wing hinge that are more
suggestive of synteliids.
Superfamily HYDROPHILOIDEA Latreille, 1802: 136
(incl. Histeroidea)
*Following Boving and Craighead (1931), Lawrence and Newton (1982) and
Newton and Thayer (1992), in contrast to Crowson (1955, 1974, 1981) and
''
815
others, Histeroidea are included here and Hydraenidae are placed in Staphy-
linoidea. Hansen (1995) accepted the latter action but recognized a more re-
stricted Hydrophiloidea including only Hydrophilidae (sensu lato ), with Spha-
eritidae through Histeridae placed in Histeroidea. However, if the characters
used to unite hydrophilids and histeroids to the exclusion of hydraenids are
accepted as synapomorphies (see under Staphyliniformia, above), there seems
to be no more justification for recognizing two superfamilies here than for
placing aquatic and terrestrial Adephaga or Scirtoidea in separate superfamilies.
The family and subfamily classification below follows Newton and Thayer
(1992), which in tum is based on works cited under each family.
HYDROPHILIDAE Latreille, 1802: 136 (incl. Georissidae=Georyssidae, Helo-
phoridae, Hydrochidae, Spercheidae, Sphaeridiidae)
*Hansen (1991b, 1995) thoroughly revised the generic and higher clas-
sification of this family and presented a phylogenetic analysis; his classifi-
cation is followed here except that he treated the first five subfamilies
below as separate families (see Newton and Thayer 1992). Bertrand
(1972), Hansen (1987), Ponomarenko (1985), Ponomarenko and Ryvkin
(1990), Smetana (1988b), Spangler (1991); a world catalog is in prepara-
tion by M. Hansen.
**"Degeer, 1774" has been cited in error as author of the family name.
HELOPHORINAE Leach, 1815: 95 (=Elophorinae)
Helophorus (=Elophorus) (Holarctic, Afrotropical).
*Angus (1992), Smetana (1985b).
**The original spelling of the type genus is Elophorus, but the much
better known spelling Helophorus has been conserved (ICZN 1993a).
EPIMETOPINAE Zaitzev, 1908: 353
Epimetopus (Neotropical, southwestern USA) and Eumetopus (Oriental,
Afrotropical).
*Costa et al. (1988).
GEORISSINAE Laporte, 1840b: 44 (=Georyssinae)
Georissus (worldwide).
*This genus was placed as a family near Elmidae or Byrrhidae until
Crowson (1950) transferred it to the vicinity of Hydrophilidae, a placement
strongly supported by larval structure (Emden 1956) and confirmed by
further analysis of adults (Hansen 1991b, Oliva 1992).
HYDROCHINAE C.G. Thomson, 1859: 15
Hydrochus (worldwide).
SPERCHEINAE Erichson, 1837: 193
Spercheus (widespread in Old World, and southern South America).
HORELOPHINAE Hansen, 1991b: 104
Horelophus (New Zealand).
816
HYDROPHILINAE Latreille, 1802: 136 (incl. Amphiopinae, Berosinae,
Hydrobiinae)
Acidocerus, Ametor, Amphiops, Anacaena, Berosus, Chaetarthria, Cym-
biodyta, Derallus, Enochrus, Helochares, Hydrobiomorpha, Hydrobius,
Hydrophilus (=Hydrous), Laccobius, Oocyclus, Paracymus, Sper-
chopsis, Sternolophus, Tropisternus, etc.
SPHAERIDIINAE Latreille, 1802: 135
Andotypus, Borborophorus, Cercyon, Coelostoma, Cryptopleurum, Cylo-
ma, Dactylosternum, Megasternum, Omicrus, Oosternum, Proto-
sternum, Rygmodus, Sphaeridium, Tormissus, etc.
*Bameul (1994), Hansen (1990), Smetana (1978).
SPHAERITIDAE Shuckard, 1839: 159
Sphaerites (western North America, Europe and China).
*Crowson ( 1974), Kryzhanovsky and Reichardt (197 6), Newton ( 1991 ),
Nikitsky (1976).
SYNTELIIDAE Lewis, 1882: 137
Syntelia (Mexico, Guatemala and eastern Asia).
*Kryzhanovsky and Reichardt (1976), Mamaev (1974), Newton (1991).
HISTERIDAE Gyllenhal, 1808: 74 (incl. Niponiidae)
*The classification of Mazur's (1984) world catalog (see Johnson et al.
1992 for corrections and index), which updates Wenzel's (1944) subfamily
system, is used here. Wenzel (1944) divided the subfamilies into two
groups, Saprinomorphae (Niponiinae through Saprininae, plus Chlamydo-
psinae) and Histeromorphae (Dendrophilinae through Hetaeriinae), each
recognized by Yelamos and Ferrer (1988) as a subfamily. However, the
monophyly of these groups and of several of the subfamilies remains un-
clear; dendrophilines and tribalines are probably para- or polyphyletic
assemblages of primitive taxa, and chlamydopsines are especially ill-de-
fined and of uncertain relationships. Larval structures including a unique
eversible maxilla closely link trypeticines and trypanaeines, but both
groups may belong in Abraeinae which also have an "extra" segment of
the labial and maxillary palps (Newton 1991). Bickhardt (1914, 1916-17),
Crowson (1974), Gorny (1983), KovarikandPassoa(1993), Kryzhanovsky
and Reichardt (1976), Vienna (1980).
NIPONIINAE Fowler, 1912: 93
Niponius (Oriental, Australia).
*Gardner (1935).
ABRAEINAE MacLeay, 1819: 25
Abraeus, Acritomorphus, Acritus, Aeletes, Chetabraeus, Halacritus, Plega-
derus, Teretriosoma, Teretrius, etc.
817
*Wenzel (1944).
TRYPETICINAE Bickhardt, 1914: 306
Trypetic_us (Oriental, New Guinea) and Pygocoelis and Trypobius (Afro-
tropical).
TRYPANAEINAE Marseul, 1857: 148 (=Tryponaeinae)
Coptotrophis, Trypanaeus, and Xylonaeus (Neotropical).
SAPRININAE Blanchard, 1845a: 276
Chalcionellus, Euspilotus, Geomysaprinus, Gnathoncus, Hypocacculus,
Hypocaccus, Myrmetes, Saprinus, Xerosaprinus, etc.
*Reichardt (1932).
DENDROPHILINAE Reitter, 1909: 288
Abraeomorphus, Anapleus, B.acanius, Carcinops, Dendrophilus, Para-
malus, Platylomalus, Xestipyge, etc.
ONTHOPHILINAE MacLeay, 1819: 25
Epiechinus (Eurasia, Africa, Australia), Onthophilus (Eurasia, North
America, Australia), etc.
TRIBALINAE Bickhardt, 1914: 307
Epierus, Idalia, Parepierus, Plagiogramma, Tribalus, etc.
HISTERINAE Gyllenhal, 1808: 74 (incl. Hololeptinae)
Atholus, Diabletes, Eblisia, Exosternus, Hister, Hololepta, Macrolister,
Margarinotus, Omalodes, Oxysternus, Pachycraerus, Phelister, Plaes-
ius, Platysoma, Spilodiscus, etc.
HETAERIINAE Marseul, 1857: 148
Hetaeriomorphus, Hetaerius, Mesynodites, Paratropus, Reninus, Sterno-
coelis, Terapus, etc.
*Helava et al. (1985).
CHLAMYDOPSINAE Bickhardt, 1914: 308
Chlamydopsis, Orectoscelis, Pheidoliphila, etc.
Superfamily STAPHYLINOIDEA Latreille, 1802: 124
*Several small families (Clambidae, Corylophidae, Hydroscaphidae, Micro-
sporidae) once included in Staphylinoidea have been justifiably removed to
other polyphagan series or to Myxophaga (e.g., Crowson 1955). The addition
ofHydraenidae (see under Staphyliniformia above) follows Boving and Craig-
head (1931), Lawrence and Newton (1982) and Hansen (1995). Lawrence and
Newton (1982) defined three monophyletic family clusters: Hydraenidae plus
Ptiliidae, Agyrtidae plus Leiodidae, and the remaining families dominated by
Staphylinidae. The family and subfamily classification below follows Newton
and Thayer (1992, 1995), which in tum are based on works cited under each
family. Leschen (1993), Newton (1984, 1985), Paulian (1941a).
818
HYDRAENIDAE Mulsant, 1844: 50 (=Limnebiidae)
*Perkins (1981) proposed a phylogeny for higher taxa and New World
genera of the family; most Old World genera have been assigned to one
of Perkins' two subfamilies (Hansen 1991a) or to a new subfamily Pro-
sthetopinae (Perkins and Balfour-Browne 1994). However, some odd gen-
era remain vaguely placed (incertae sedis, below) and there are und-
escribed genera in the southern hemisphere that do not fit the existing
classification well.
**Limnebiidae Mulsant (1844: 88) has equal priority for the family
name, but Hydraenidae is better known.
PROSTHETOPINAE Perkins in Perkins and Balfour-Browne, 1994: 7
Mesoceration, Nucleotops, Parasthetops, Prosthetops and Pterosthetops
(southern Africa), Protosthetops (Kenya), and Sicilicula (Madagascar,
Reunion).
HYDRAENINAE Mulsant, 1844: 50
Adelphydraena, Homalaena, Hydraena, Hydraenida, Laeliana, Limnebius,
Orchymontia, Podaena, etc.
*Ordish (1984), Perkins (1989).
OCHTHEBIINAE C.G. Thomson, 1859: 15
Gymnochthebius, Hughleechia, Meropathus, Micragasma, Neochthebius,
Ochthebius, and Tympanogaster.
*Hich (e.g., 1990).
HYDRAENIDAE incertae sedis:
Coelometopon, Decarthrocerus, Discozantaena and Parhydraena (Africa).
PTILIIDAE Erichsen, 1845: 15/Motschulsky, 1845: 504 (=Trichopterygidae;
incl. Cephaloplectidae, Limulodidae)
*Except for the addition of Cephaloplectidae (=Limulodidae; see Crow-
son 1981, Lawrence 1982, Lawrence and Newton 1982) the internal orga-
nization of the family used here is that discussed by Dybas (1976), who
indicated problems with these existing formal groups but did not propose
a new classification. Besuchet (1976), Besuchet and Sundt (1971), Bistrom
and Silfverberg 1979), Costa et al. (1988), Dybas (1990), C. Johnson
(1982, 1985).
**Heer (1843: 60) is usually cited as author of the family name (e.g.,
by Silfverberg 1992), but this name is unavailable (Ptilium was not treated
as a valid name). Trichopterygidae Erichsen (1845: 13) is based on a pre-
occupied generic name, Trichopteryx Kirby (not Hubner), now Acrotrichis.
PTILIINAE Erichsen, 1845: 15/Motschulsky, 1845: 504
Actidium, Cochliarion, Motschulskium, Nossidium, Ptenidium, Pteryx, Ptil-
ium, etc.
819
NANOSELLINAE Barber, 1924: 170
Cylindrosella, Mycophagus, Nanosella, Philagarica, Throscidium, etc.
CEPHALOPLECTINAE Sharp, 1883: 295 (=Limulodinae)
Cephaloplectus (Neotropical), Eulimulodes (Mexico), Limulodes and Para-
limulodes (New World), and Rodwayia (Australia).
*This group of specialized myrmecophiles was reviewed and redefined
by Seevers and Dybas (1943) to include Cephaloplectus and Eulimulodes
from Staphylinidae; they and Dybas (1976) treated it as a separate family,
Limulodidae.
**Cephaloplectinae has priority over Limulodinae Ganglbauer (1899:
297) for the name of this group.
ACROTRICHINAE Reitter, 1909: 272 (1856)
Acrotrichis, Nephanes, etc.
**Newton and Thayer (1992) used Art. 40b to maintain Acrotrichinae
over the unused name Cleopteriinae Gistel (1856: 360), based on Cleopter-
ium, a synonym of Acrotrichis.
AGYRTIDAE C.G. Thomson, 1859: 57 (=Silphidae, part; incl. Lyrosominae,
Pterolominae)
Agyrtes, Apteroloma, lpelates, Lyrosoma, and Pteroloma (Holarctic), Eca-
nus (northern Europe), and Necrophilus (Holarctic and New Zealand).
*This primitive group of genera showing relict distribution patterns was
previously treated as a subgroup of Silphidae (e.g., Arnett 1963, Madge
1980b), but lately recognized as a separate family more closely related to
leiodids than silphids (e.g., Anderson and Peck 1985, Lawrence 1982,
Lawrence and Newton 1982, Peck 1990, Schawaller 1991). Agyrtid larvae
are the most generalized known in Staphylinoidea (Newton 1991, Zwick
1981), and one of the earliest known staphylinoids, the early Jurassic ge-
nus Mesecanus (=Mesagyrtes), probably belongs here although originally
placed in Silphidae (Ponomarenko in Arnoldi et al. 1977, Ponomarenko
and Ryvkin 1990).
LEIODIDAE Fleming, 1821: 51 (=Anisotomidae, Liodidae; incl. Camiaridae,
Catopidae, Cholevidae, Colonidae, Leptinidae, Leptodiridae, Platypsyll-
idae, Sogdiidae)
*This family is used here in the broad sense of Crowson (1981), Law-
rence and Newton (1982), and Peck (1990) to include groups often placed
in up to five separate families (e.g., Jeanne! 1936, Perreau 1989), or in-
cluded in the old broad concept of Silphidae (e.g., Hatch 1928). Many
groups treated as tribes here are treated as subfamilies by some authors
(see discussion and tribal classification in Newton and Thayer 1992). A
defining synapomorphy for the family is the presence of complex antenna!
818
HYDRAENIDAE Mulsant, 1844: 50 (=Limnebiidae)
*Perkins (1981) proposed a phylogeny for higher taxa and New World
genera of the family; most Old World genera have been assigned to one
of Perkins' two subfamilies (Hansen 1991a) or to a new subfamily Pro-
sthetopinae (Perkins and Balfour-Browne 1994). However, some odd gen-
era remain vaguely placed (incertae sedis, below) and there are und-
escribed genera in the southern hemisphere that do not fit the existing
classification well.
**Limnebiidae Mulsant (1844: 88) has equal priority for the family
name, but Hydraenidae is better known.
PROSTHETOPINAE Perkins in Perkins and Balfour-Browne, 1994: 7
Mesoceration, Nucleotops, Parasthetops, Prosthetops and Pterosthetops
(southern Africa), Protosthetops (Kenya), and Sicilicula (Madagascar,
Reunion).
HYDRAENINAE Mulsant, 1844: 50
Adelphydraena, Homalaena, Hydraena, Hydraenida, Laeliana, Limnebius,
Orchymontia, Podaena, etc.
*Ordish (1984), Perkins (1989).
OCHTHEBIINAE C.G. Thomson, 1859: 15
Gymnochthebius, Hughleechia, Meropathus, Micragasma, Neochthebius,
Ochthebius, and Tympanogaster.
*Jach (e.g., 1990).
HYDRAENIDAE incertae sedis:
Coelometopon, Decarthrocerus, Discozantaena and Parhydraena (Africa).
I
PTILIIDAE Erichsen, 1845: 15/Motschulsky, 1845: 504 (=Trichopterygidta,
incl. Cephaloplectidae, Limulodidae)
*Except for the addition of Cephaloplectidae (=Limulodidae; see Cro I -
son 1981, Lawrence 1982, Lawrence and Newton 1982) the internal orga-
nization of the family used here is that discussed by Dybas (1976), ~o
indicated problems with these existing formal groups but did not prop se
a new classification. Besuchet (1976), Besuchet and Sundt (1971 ), Bistr ·m
and Silfverberg 1979), Costa et al. (1988), Dybas (1990), C. Johnson
(1982, 1985).
**Heer (1843: 60) is usually cited as author of the family name (e.g.,
by Silfverberg 1992), but this name is unavailable (Ptilium was not treated
as a valid name). Trichopterygidae Erichsen (1845: 13) is based on a pre-
occupied generic name, Trichopteryx Kirby (not Hubner), now Acrotrichis.
PTILIINAE Erichsen, 1845: 15/Motschulsky, 1845: 504
Actidium, Cochliarion, Motschulskium, Nossidium, Ptenidium, Pteryx, Ptil-
ium, etc.
819
NANOSELLINAE Barber, 1924: 170
Cylindrosella, Mycophagus, Nanosella, Philagarica, Throscidium, etc.
CEPHALOPLECTINAE Sharp, 1883: 295 (=Limulodinae)
Cephaloplectus (Neotropical), Eulimulodes (Mexico), Limulodes and Para-
limulodes (New World), and Rodwayia (Australia).
*This group of specialized myrmecophiles was reviewed and redefined
by Seevers and Dybas (1943) to include Cephaloplectus and Eulimulodes
from Staphylinidae; they and Dybas (1976) treated it as a separate family,
Limulodidae.
**Cephaloplectinae has priority over Limulodinae Ganglbauer (1899:
297) for the name of this group.
ACROTRICHINAE Reitter, 1909: 272 (1856)
Acrotrichis, Nephanes, etc.
**Newton and Thayer (1992) used Art. 40b to maintain Acrotrichinae
over the unused name Cleopteriinae Gistel (1856: 360), based on Cleopter-
ium, a synonym of Acrotrichis.
AGYRTIDAE C.G. Thomson, 1859: 57 (=Silphidae, part; incl. Lyrosominae,
Pterolominae)
Agyrtes, Apteroloma, /pelates, Lyrosoma, and Pteroloma (Holarctic), Eca-
nus (northern Europe), and Necrophilus (Holarctic and New Zealand).
*This primitive group of genera showing relict distribution patterns was
previously treated as a subgroup of Silphidae (e.g., Arnett 1963, Madge
1
1980b ), but lately recognized as a separate family more closely related to
leiodids than silphids (e.g., Anderson and Peck 1985, Lawrence 1982,
Lawrence and Newton 1982, Peck 1990, Schawaller 1991). Agyrtid larvae
are the most generalized known in Staphylinoidea (Newton 1991, Zwick
1981), and one of the earliest known staphylinoids, the early Jurassic ge-
nus Mesecanus (=Mesagyrtes), probably belongs here although originally
placed in Silphidae (Ponomarenko in Arnoldi et al. 1977, Ponomarenko
and Ryvkin 1990).
LEIODIDAE Fleming, 1821: 51 (=Anisotomidae, Liodidae; incl. Camiaridae,
Catopidae, Cholevidae, Colonidae, Leptinidae, Leptodiridae, Platypsyll-
idae, Sogdiidae)
*This family is used here in the broad sense of Crowson (1981), Law-
rence and Newton (1982), and Peck (1990) to include groups often placed
in up to five separate families (e.g., Jeannel 1936, Perreau 1989), or in-
cluded in the old broad concept of Silphidae (e.g., Hatch 1928). Many ·
groups treated as tribes here are treated as subfamilies by some authors
(see discussion and tribal classification in Newton and Thayer 1992). A
defining synapomorphy for the family is the presence of complex antenna!

820
"vesicles" or sensilla-filled iiwaginations in several preapical segments
(Angelini and De Marzo 1980). Newton (1984, 1985, 1991).
**Newton and Thayer (1992) reviewed problems with the use of the
name Anisotomidae Stephens (1828: 99) for the family and nominotypical
subfamily names.
CAMIARINAE Jeanne!, 1911: 192 (incl. Agyrtodinae)
Afropelates, Agyrtodes, Camiarus, Catopsolius, Dasypelates, Eublack-
burniella, Eupelates, Myrmecholeva, Neocamiarus, Neopelatops, Ragy-
todes, Sphaeropelatops, Zearagytodes, etc.
*This southern temperate group was redefined by Newton (1985) to
include Jeannel's (1936, 1962b, 1964b) groups Agyrtodini (from Cho-
levinae) and Neopelatopini (from Leiodinae) as well as Camiarinae (Je-
anne! 1958). It includes the least derived leiodids (e.g., Ragytodes is the
only leiodid with true ocelli, and all known larvae have 5 stemmata in
contrast to 3 or fewer in other leiodids) with species in New Zealand, Aus-
tralia, South Africa and southern South America; a new genus has been
discovered recently in New Guinea. Szymczakowski (1966), Zwick
(1979).
CATOPOCERINAE Hatch, 1927: 4 (1880) (incl. Glacicavicolinae)
Catopocerus (North America, eastern Siberia) and Glacicavicola (western
North America).
*This group of blind, flightless beetles was formerly recognized as two !'
subfamilies (Westcott 1968) but combined by Peck (1990). Catopocerus
is now known from eastern Russia (Perkovsky 1989), and an undescribed
genus and species from Chile probably belong in this subfamily (Newton!
1985). Peck (1975).
**Pinodytinae Horn (1880b: 248) has priority over Catopocerinae for
the name of this group but retention of the latter name has been justified
by Art. 40b (Newton and Thayer 1992).
LEIODINAE Fleming, 1821: 51 (=Anisotominae)
Agathidium, Aglyptinus, Anisotoma, Colenis, Colenisia, Cyrtusa, Dietta
(=Estadia), Hydnobius, Leiodes, Pseudoliodes, Scotocryptus, Sogda
(=Trichohydnobius), Triarthron, Zeadolopus, etc.
*This large group includes six tribes with disputed interrelationships
(Newton 1984, Wheeler 1984). Baranowski (1993), Costa et al. (1988),
Daffner (1983, 1989), Peck (1990), Wheeler (1979a, 1979b).
COLONINAE Horn, 1880b: 266 (1859)
Colon (worldwide) and Colonellus (Oriental).
*Szymczakowski (e.g., 1964b, 1969) has reviewed much of the world
fauna; a revision of Nearctic species is in preparation by K. Stephan and
S. Peck.
**The unused name Myloechinae C.G. Thomson (1859: 60) has prior-
ity over Coloninae for the name of this group but retention of the latter
821
name has been justified by Art. 40b (Newton and Thayer 1992). The origi-
nal spelling of the type genus is Kolon, but Silfverberg (1994b) has ap-
plied to the ICZN for conservation of the spelling Colon.
CHOLEVINAE Kirby, 1837: 108 (=Catopinae; incl. Anemadinae, Bathy-
sciinae, Eucatopinae, Leptodirinae, Nemadinae, Ptomaphaginae)
Anemadus, Antroherpon, Bathyscia, Bathysciola, Catops, Choleva, Disso-
chaetus, Eocatops, Eucatops, Leptodirus, Nargomorphus, Nemadus,
Oritocatops, Paracatops, Platycholeus, Prionochaeta, Pseudonemadus,
Ptomaphaginus, Ptomaphagus, Sciodrepoides, Speonomus, etc.
*The subfamily limits and internal classification of this large group are
not settled (see, e.g., Giachino and Vailati 1993, Newton and Thayer 1992,
Perreau 1989), although the group as a whole ranks among the best-known
staphylinoids. The primarily Palearctic and cave-dwelling Leptodirini
(=Bathysciinae) has been monographed by Jeanne! (1911, 1924), with
updated but divergent classifications presented by Casale et al. (1991),
Gueorguiev (1976), and Laneyrie (1967). Jeanne! (1936, 1962b, 1964b),
Peck (1973, 1990), Szymczakowski (1964a, 1966), Zwick (1979), Zwick
in Klausnitzer (1978).
**Cholevinae has priority over the other frequently-used names Catop-
inae Chaudoir (1845: 195), Leptodirinae Lacordaire (1854: 195) and Ba-
thysciinae Horn (1880b: 251) for the name of this subfamily, and Lepto-
dirini has priority over Bathysciini for the name of the largest tribe.
PLATYPSYLLINAE Ritsema, 1869: 38 (=Leptininae)
Leptinillus (North America), Leptinus and Platypsyllus (Holarctic), and
Silphopsyllus (Russia).
\ *These ectoparasites or inquilines of mammals were at one time treated
as the separate families Platypsyllidae (Platypsyllus) and Leptinidae (other
genera), but are clearly closely related to each other (Jeannel 1922) and to
leiodids. Besuchet (1980), Parks and Barnes (1955), Peck (1982), Wood
(1965).
\ **The name Leptininae is usually attributed to LeConte (1866b: 368)
and given priority over Platypsyllinae, but no family-group name based on
Leptinus was actually used until LeConte (1872: 802) (Newton and Thayer
1992).
SCYDMAENIDAE Leach, 1815: 92 (incl. Anisosphaeridae)
*Scydmaenids are an isolated family of uncertain relationships, some-
times placed near pselaphids (Boving and Craighead 1931) or leiodids
(Crowson 1955) but probably a derivative group of staphylinids (Lawrence
and Newton 1982, Newton and Thayer 1995). Plaumanniola, originally
placed in a new subfamily of Ptinidae (Plaumanniolinae Costa Lima 1962:
415), was shown to belong here by Lawrence and Reichardt (1966). The
subfamily and tribal classification is in need of revision; a world catalog
822
of the genera is in preparation by A. Newton and H. Franz. Brown and
Crowson (1980), De Marzo (1984), Franz (1967, 1975, 1980, 1985, 1986),
Leleup (1968), Newton (1991), Schmid (1988a, 1988b).
MASTIGINAE Fleming, 1821: 49 (=Clidicinae)
Ablepton, Clidicus, Leptomastax, Mastigus, Papusus, etc.
**Mastiginae has priority over the more widely used Clidicinae Casey
(1897: 541) for the name of this subfamily.
SCYDMAENINAE Leach, 1815: 92 (incl. Plaumanniolinae)
Ascydmus, Cephennium, Chevrolatia, Cyrtoscydmus, Euconnus, Eutheia,
Leptoscydmus, Lophioderus, Microscydmus, Neuraphes, Opresus, Plau-
manniola, Scydmaenus, Siamites, Stenichnus, Syndicus, Veraphis, etc.
SILPHIDAE Latreille, 1807a: 1
*This once vaguely defined group is now restricted to the larger carrion
and burying beetles (e.g., Anderson and Peck 1985, Peck 1990), after the
removal of some subfamilies or tribes to form a separate family Agyrtidae
(see above), and the removal of other groups to the families Leiodidae
(e.g., Bathysciinae, Estadiini) or Staphylinidae (e.g., Microsilphinae, Gly-
pholoma, Deinopteroloma) (Lawrence and Newton 1982, Newton 1985,
Smetana 1985a). In this restricted sense, silphids are clearly monophyletic
and closely allied to staphylinids (Lawrence and Newton 1982). Several
genera here treated as the staphylinid subfamilies Apateticinae and Trigo-
nurinae have been included in Silphidae by some authors (e.g., Madge
1980a, 1980b). Costa et al. (1988), Newton (1991), Peck and Anderson !
(1985), Peck and Kaulbars (1987), Portevin (1926), Schawaller (1987),,'
Young (1983).
SILPHLl\TAE Latreille, 1807a: 1 i
Aclypea (=Blitophaga) (Holarctic); Diamesus (eastern Asia, New Guinea:,
Australia); Heterosilpha (western North America); Necrodes, Necro-
phila and Oiceoptoma (Eurasia, North America); Oxelytrum (south-
western USA, Neotropical); Ptomaphila (Australia, New Guinea); Sil~
pha (Eurasia, Africa); and Thanatophilus (Eurasia, Africa, North Amer-
ica).
NICROPHORINAE Kirby, 1837: 95
Nicrophorus (=Necrophorus) (widespread except Afrotropical and Austra-
lian regions) and Ptomascopus (eastern Asia).
STAPHYLINIDAE Latreille, 1802: 124 (incl. Brathinidae, Dasyceridae, Em-
pelidae, Micropeplidae, Oxytelidae, Oxyporidae, Pselaphidae, Sca-
phidiidae, etc.)
*There is disagreement on the family limits as well as the subfamily
classification of this huge family (e.g., Coiffait 1972-84, Lawrence and
Newton 1982, Moore 1964, Naomi 1985, Newton and Thayer 1988, 1992,
823
1995, Tikhomirova 1973). Coiffait (1972-84), without discussion, and
Naomi (1985), on dubious phylogenetic grounds, broke the family into ten
and three families, respectively. Four groups often regarded as separate
families are treated as staphylinid subfamilies here because of strong evi-
dence relating them to certain staphylinid subgroups: Scaphidiidae (Crow-
son 1981, Kasule 1966, Lawrence and Newton 1982), and Dasyceridae,
Micropeplidae and Pselaphidae (Naomi 1985, Newton and Thayer 1995,
Thayer 1987). Silphidae and Scydmaenidae are also closely related to Sta-
phylinidae forming with them a probably monophyletic group, roughly
equivalent to the traditional "Brachelytra", defined by derived features of
wing folding pattern, aedeagal structure and larval structures (Lawrence
and Newton 1982). Tikhomirova and Melnikov (1970) demonstrated that
the articulated larval mala of paederines and staphylinines was a highly
derived condition in these groups and did not justify dividing staphylinids
(and related families) into two great groups as Paulian (1941a) had sug-
gested. The family was divided into four informal groups by Lawrence and
Newton (1982): Omaliine Group (Glypholomatinae through Pselaphinae );
Tachyporine Group (Phloeocharinae through Aleocharinae); Oxyteline
Group (Trigonurinae through Oxytelinae); and Staphylinine Group (Oxy-
porinae through Staphylininae, and possibly Scydmaenidae and Silphidae).
Newton and Thayer (1992) summarized the current classification of Sta-
phylinidae above the generic level, and Ashe and Newton (1993) and
Newton and Thayer (1995) completed phylogenetic analyses of the Tachy-
porine and Omaliine Groups, respectively. A world catalog is in prepara-
tion by L. Herman.
!
Blackwelder (1936, 1943, 1952), Cameron (1930-39), Coiffait and Safz
(1968), Dettner (1993), Frank (1991), Hammond (1984), Kasule (1966,
1968, 1970), Moore and Legner (1974, 1975, 1979), Naomi (1987-90),
Newton (1990a, 1991), Outerelo-Domfnguez and Gamarra-Hidalgo (1985),
Palm (1948-72), Ponomarenko and Ryvkin (1990), Pototskaya (1967),
Ryvkin (1985), Scheerpeltz (1972, 1974), Shibata (1976-85), Tikhomirova
(1968), Toppin Klausnitzer (1978), Welch (1993).
GLYPHOLOMATINAE Jeannel, 1962b: 482
Glypholoma (southeastern Australia, southern South America).
*Originally placed in its own tribe in Silphidae (J eannel 1962b), this
genus was moved to Omaliinae by Newton (1975), revised by Thayer and
Newton (1979), and elevated to subfamily rank by Newton and Thayer
(1994) based on its basal phylogenetic position within the Omaliine Group.
Glypholomatinae and the remaining nine subfamilies of the Omaliine
Group (except Micropeplinae and some Pselaphinae) have a unique and
complex defensive abdominal gland on sternum 8 (Dettner 1993, Newton
and Thayer 1995).
824
MICROSILPHINAE Crowson, 1950: 284
Microsilpha (=Micragyrtes,Acruliodema) (Australia, New Zealand, south-
ern South America).
*Originally placed as a subfamily of Silphidae (Crowson 1950), this
group was removed to the vicinity of Omaliinae by Lawrence and Newton
(1982), partially reviewed by Newton (1985), and its phylogenetic place-
ment discussed by Newton and Thayer (1995).
OMALIINAE MacLeay, 1825: 49 (incl. Brathinidae, Aphaenostemminae)
Acidota, Anthobium, Anthophagus, Aphaenostemmus, Boreaphilus, Brathi-
nus, Camioleum, Corneolabium, Coryphium, Crymus, Deinopteroloma,
Eusphalerum, Giulianium, Hadrognathus, Metacorneolabium, Micraly-
mma, Omalium, Phloeonomus, Tanyrhinus, Tetradelus, Trigonodemus,
etc.
*This and the preceding two subfamilies are among the most primitive
of all staphylinids, notable for the retention of a pair of ocelli on the head
and the temperate distribution of most genera. Brathinus, often placed in
a separate family Brathinidae LeConte (1861: 48), belongs here (Thayer
1985a), as does Deinopteroloma from Silphidae (Smetana 1985a). Newton
and Thayer (1995) attempted to resolve tribal relationships, adding Apha-
enostemminae Peyerimhoff (1914: 248) as a tribe including Giulianium
(from Phloeocharinae), and removing Glypholomatini. A world generic
revision of Omaliini is in preparation by M. K. Thayer, and revisions of
North American genera by J. M. Campbell. Steel (1970), Thayer (1985b),
Watanabe (1990), Zanetti (1987), Zerche (1990).
EMPELINAE Newton and Thayer, 1992: 25 (Abdullah, 1969b: 683, nomen
nudum)
Empelus (western North America).
*This odd genus with long elytra was considered a clambid until Crow-
son (1955) moved it to Leiodidae, and later (1960) characterized it as one
of the most primitive staphylinoids; it has also been treated as a separate
family (e.g., Abdullah 1969b, Crowson 1981). Newton and Thayer (1992,
1995) formally named the group and placed it near Omaliinae.
PROTEININAE Erichson, 1839a: 641 (=Pteroniinae; incl. Metopsiinae)
Alloproteinus, Anepius, Austrorhysus, Eupsorus, Megarthroides, Megar-
thrus, Metopsia, Nesoneus, Paranesoneus, Proteinus, and Silphotelus.
*Steel (1966) added Metopsiinae, described new larvae, proposed a
new tribal classification and revised the two south temperate tribes (Neso-
neini and Anepiini); Newton and Thayer (1995) proposed a tribal phylog-
eny with two new tribes for parts of Nesoneini. This and the following
five subfamilies have non-functional spiracles on the intermediate abdomi-
nal segments (4-6 or 3-6) of adults, a feature known elsewhere only
within Tachyporinae.
825
**The type genus Proteinus has been conserved in the present sense,
eliminating the need for the replacement names Pteronius and Pteroniinae
(ICZN 1969).
MICROPEPLINAE Leach, 1815: 90
Cerapeplus (Thailand, Borneo), Kalis sus (northwestern North America),
Micropeplus (Holarctic, Mexico), Peplomicrus (Neotropical, Afrotropi- \
cal), and Pseudokalissus (eastern Russia).
*This group is often treated as a distinct staphylinoid family (e.g., Lobi
and Burckhardt 1988a), in large part due to its odd larvae (Newton 1991),
but has adult and larval characteristics that clearly place it in the Omaliine
Group (Thayer 1987, Newton and Thayer 1995). Campbell (1968, later
supplements).
NEOPHONINAE Fauvel, 1905: 98
Neophonus (southern South America).
*The single isolated genus and species included here is a grazer of leaf-
surface microfungi (Thayer 1987).
DASYCERINAE Reitter, 1887: 8
Dasycerus (North America and Eurasia).
*This odd lathridiid-like genus was included in that family until Crow-
son (1955) moved it to Staphylinoidea as a distinct family. Adults have
abdominal glands and other features that clearly place them in the Omali-
ine Group (Thayer 1987, Newton and Thayer 1995). Lobi (1977b), New-
ton (1991).
PROTOPSELAPHINAE Newton and Thayer, 1995 (this volume)
Protopselaphus (Malaysia).
*A newly proposed subfamily for an isolated genus that is the sister
group to the former family Pselaphidae (Newton and Thayer 1995).
PSELAPHINAE Latreille, 1802: 239 (incl. Batrisinae, Bythininae, Clavi-
gerinae, Euplectinae, Faroninae, Goniacerinae, Mayetiinae)
Amaurops, Batrisodes, Batrisus, Brachygluta, Bryaxis, Bythinoplectus,
Bythinus, Claviger, Colilodion, Ctenistes, Cyathiger, Decarthron, Eupi-
nes, Euplectus, Faronus, Goniacerus, Hamotus, Jubus, Mayetia, Octo-
micrus (=Dimerus), Pselaphus, Pyxidicerus, Raffrayia, Reichenbachia,
Sagola, Trogaster, Tyrus, etc.
*Newton and Thayer (1995) reviewed conflicting opinions on the rela-
tionships of pselaphids, provided strong evidence nesting them within part
of the Omaliine Group of staphylinid subfamilies, and reduced the family
to a subfamily and the former pselaphid subfamilies to supertribes. The
last worldwide generic review (Raffray 1908) and keys to higher taxa
(Jeannel 1955b) are out of date but still useful; Newton and Chandler
(1989) provided a world generic checklist. Besuchet (1991), Chandler
(1990), Costa et al. (1988), Coulon (1989), De Marzo (1987), Grigarick
826
and Schuster (1980), Jeanne! (1950, 1959, 1962a, 1964a), Newton (1991),
Nomura (1991), Park (1942, 1952), Park et al. (1976).
PHLOEOCHARINAE Erichson, 1839a: 612
Charhyphus (North America to Guatemala, eastern Russia), Ecbletus (Pan-
ama, USA: California), Dytoscotes (western USA), Phloeocharis (Eu-
rope and USA: California), Phloeognathus (New Zealand), Pseudo-
phloeocharis (Australian region), and Vicelva (western USA).
*This small relict group was redefined by Herman ( 1972) to exclude
Olisthaerus (Olisthaerinae) and Derops (to Tachyporinae), but larval char-
acters of Olisthaerus are consistent with including this genus in Phloeo-
charinae (Ashe and Newton 1993). Giulianium, tentatively placed here by
Moore (1976), was moved to Omaliinae: Aphaenostemmini by Newton
and Thayer (1995).
OLISTHAERINAE C.G. Thomson, 1858: 38
Olisthaerus (Holarctic ).
TACHYPORINAE MacLeay, 1825: 49
Bolitobius, Bryoporus, Cilea, Coproporus (=Erchomus), Derops (=Par-
a/easter, Rimulincola), Ischnosoma, Leucotachinus, Lordithon, Megar-
thropsis, Mycetoporus, Sepedophilus (=Conosoma), Symmixus, Tachi-
nomorphus, Tachinus, Tachyporus, Vatesus, etc.
*Smetana (1983) confirmed the addition of Derops and provided a key
to most tribes of this subfamily. Several groups formerly included in Ta-
chyporinae were moved to Aleocharinae by Hammond (1975), but even
after such removals the subfamily is doubtfully monophyletic (Ashe and
Newton 1993). A generic revision and phylogenetic study are in prepara-
tion by J. M. Campbell, who has revised most North American genera in
a series of papers. Ullrich (1975).
**The name Tachyporinae has been conserved over the older name
Tachinidae Fleming (1821: 49) by Opinion 1743 (ICZN 1993c); the same
Opinion changed the correct spelling of Fleming's name to Tachinusidae
to avoid homonymy with the name Tachinidae in Diptera.
TRICHOPHYINAE C.G. Thomson, 1858: 30
Trichophya (Eurasia, North America, Mexico, Guatemala).
*Ashe and Newton (1993) provided evidence from larvae and adults
that this subfamily is the sister group of Habrocerinae.
HABROCERINAE Mulsant and Rey, 1877: 65
Habrocerus (Eurasia, North America, Brazil) and Nomimocerus (southern
South America, eastern Russia).
*The profoundly modified male abdominal apex and genitalia of hab-
rocerines have led some authors (e.g., Coiffait and Safz 1965) to recognize
them as a separate family, but Ashe and Newton (1993) demonstrated that
they are very closely related to trichophyines which have normal genitalia.
827
ALEOCHARINAE Fleming, 1821: 49 (incl. Cyphinae, Hypocyphtinae, Me-
soporinae, Mimanommatinae, Pseudoperinthinae, Pulicomorphinae,
Pygosteninae, Termitodiscinae, Trichopseniinae, Trilobitideinae, etc.)
Aleochara, Atheta, Bolitochara, Corotoca, Cypha (=Hypocyphtus), De ina-
psis, Dorylomimus, Ecitogaster, Falagria, Gymnusa, Gyrophaena, Ho-
malota, Hoplandria, Hygronoma, Leptusa, Lomechusa, Mesoporus,
Mimanomma, Mimeciton, Myllaena, Oligota, Oxypoda, Paraconosoma,
Phytosus, Placusa, Pseudoperinthus, Pulicomorpha, Pygostenus, Ter-
mitodiscus, Trichopsenius, Trilobitideus, Zyras, etc.
*This largest and taxonomically most difficult staphylinid subfamily
was defined by Hammond (1975) to include all those staphylinids that
share a uniquely derived complex aedeagal structure, including Tricho-
pseniinae and Hypocyphtinae, which have been kept as separate subfami-
lies by some authors (e.g., Pasteels and Kistner 1971, Seevers 1978). Mo-
nophyly of the subfamily is also supported by larval structure (Ashe and
Newton 1993) and for all but a few primitive groups by defensive glands
and chemistry (Steidle and Dettner 1993). Seevers (1978) attempted the
most recent internal classification of the subfamily in a review of the
North American genera, but the world tribal/subtribal classification re-
mains chaotic. Ashe (1984, 1986, 1992, 1994), Fenyes (1918-21), Frank
and Thomas (1984), Hoebeke (1985), Kistner (1958, 1982, and many oth-
ers with coworkers), Klimaszewski (1979), Pace (1989), Seevers (1957,
1965), Yosii and Sawada (1976).
TRIGONURINAE Reiche, 1865: 642
Trigonurus (southwestern Europe, Caucasus, western North America).
*This genus and Apateticinae traditionally have been included in the
subfamily Piestinae, but some authors (e.g., Madge 1980a, 1980b) have
placed them in Silphidae; Newton and Thayer (1992) treated both groups
as staphylinid subfamilies. Trigonurus may be the most primitive member
of the Oxyteline Group, with plesiomorphic larvae (Pototskaya 1976) com-
pared to those of the remaining subfamilies; three genera of Mesozoic age
have been compared to it (Ponomarenko and Ryvkin 1990, Tikhomirova
1968). Blackwelder (1941).
APATETICINAE Fauvel, 1895: 190
Apatetica and Nodynus (southeastern Asia).
*Apateticines have had a taxonomic history paralleling that of trigo-
nurines (see above), but larval characters (Fukuda in Kawada 1960) as
well as adult structure suggest they are primitive members of the Oxyteline
Group, possibly nearest to scaphidiines. Cameron (1930-39), Madge
(1980a).
828
SCAPHIDIINAE Latreille, 1807a: 3
Baeocera, Bironium (=Heteroscapha), Baeoceridium, Cerambyciscapha,
Cyparium, Diatelium, Scaphobaeocera, Scaphidium, Scaphisoma, Scap-
hium, Toxidium, etc.
*Scaphidiines traditionally have been treated as a separate family, or
even two families (Tamanini 1969), but we agree with Kasule (1966), who
corrected earlier larval misidentifications, and some later authors who treat
them as a staphylinid subfamily near piestines or apateticines. Leschen
(1993, including partial phylogeny), Lobi (1977a, 1990, 1992 and con-
tained references), Newton (1991).
PIESTINAE Erichson, 1839b: 31
Eupiestus and Piestoneus (southeastern Asia), Hypotelus and Piestus (Neo-
tropical, southern USA), Parasiagonum (New Zealand), Prognathoides
(Australia), and Siagonium (Holarctic).
*Only seven genera remain in this traditional staphylinid dumping
ground after removal of Trigonurus (Trigonurinae), Apatetica and Nodynus
(Apateticinae ), and various other groups or genera to Osoriinae (Blackwel-
der 1942), Phloeocharinae (Herman 1972), Pseudopsinae (Newton 1982a)
or Oxytelinae (Newton 1982b). Crowson (1995), Crowson and Ellis
(1969), Steel (1950a).
OSORIINAE Erichson, 1839b: 30 (incl. Eleusininae, Lispininae, Lepto-
chirinae)
Borolinus, Clavilispinus (=Paralispinus), Cylindropsis, Dirocephalus, Ele-
usis, Espeson, Geomitopsis, Glyptoma, Holotrochus, Leptochirus, Lispi-
nus, Mimogonus, Nacaeus, Neolosus (=Holosus), Osorius, Parosorius,
Priochirus, Renardia (=Eumalus), Thoracophorus, Zeoleusis, etc.
*This large subfamily, unusual among staphylinids in having an "un-
margined" abdomen, is used here in the expanded sense of Blackwelder
(1942) to include several groups formerly placed in Piestinae. A. Newton
is preparing a world review above the generic level. Coiffait (1979), Pagel
(1955, 1959), Greenslade (1971), McColl (1982), Newton (1990b), Steel
(1950b).
OXYTELINAE Fleming, 1821: 49
Anotylus, Aploderus, Apocellus, Bledius, Carpelimus (=Trogophloeus),
Coprophilus, Deleaster, Euphanias, Oxypius, Oxytelus, Platystethus,
Sartallus, Syntomium, Thinobius, Thinodromus, etc.
*Herman (1970) redefined the subfamily, with removal of several gen-
era to other subfamilies, as part of a complete generic revision and phylo-
genetic analysis. Newton (1982b) added Euphanias from Piestinae and
proposed a modified phylogeny for the group. Hammond (1976), Herman
(1986).
OXYPORINAE Fleming, 1821: 49
Oxyporus (Eurasia, North and Central America).
829
*Campbell (1969 and supplements).
MEGALOPSIDIINAE Leng, 1920: 98 (=Megalopininae, Stylopodinae)
Megalopinus (=Megalops, Megalopsidia, Stylopodus) (worldwide, espe-
cially in tropics).
*Benick (1951).
**Newton and Thayer (1992) reviewed the family-group name synon-
ymy.
STENINAE MacLeay, 1825: 49
Dianous (Holarctic, Oriental) and Stenus (worldwide).
*Puthz (1971, 1981, many other papers).
EUAESTHETINAE C.G. Thomson, 1859: 42
Alzadaesthetus, Austroesthetus, Coiffaitia, Edaphus, Euaesthetus, Fen-
deria, Nordenskioldia, Octavius, Protopristus, Stenaesthetus, Stictocra-
nius, Tamotus, etc.
*The tribal system proposed by Scheerpeltz (1974) did not include all
genera and needs review. Orousset (1988), Puthz (1974, 1978).
SOLIERIINAE Newton and Thayer, 1992: 27
Solierius (=Physognathus) (southern South America).
*An isolated pselaphine-like genus generally included in Omaliinae
(e.g., Coiffait and Saiz 1968), but removed to a separate subfamily of un-
certain relationships and later placed near Euaesthetinae by Newton and
Thayer (1992, 1995).
LEPTOTYPHLINAE Fauvel, 1874: 329
Cephalotyphlus, Chionotyphlus, Entomoculia, Leptotyphlus, M egatyphlus,
Metrotyphlus, Neotyphlus, etc.
*These minute, blind,.soil-dwelling species resemble euaesthetines, but
the articulated larval mala (A. Newton, unpublished) suggests closer kin-
ship to the following three subfamilies. Coiffait (1959, 1962, 1963, 1972-
84), Smetana (1986).
PSEUDOPSINAE Ganglbauer,,l895: 690
Pseudopsis (Holarctic, Neotropical, New Zealand) and Asemobius, Nano-
bius, and Zalobius (western North America).
*This subfamily was redefined by Newton (1982a) to include the last
3 genera, formerly placed in Oxytelinae or Piestinae; larval structures in-
cluding an articulated mala (Newton 1990a) indicate that pseudopsines are
related to Paederinae and Staphylininae rather than Oxytelinae (Herman
1975) or Phloeocharinae (Newton 1982a). Herman (1977), Zerche (1992).
PAEDERINAE Fleming, 1821: 49 (incl. Pinophilinae)
Achenium, Astenus, Cryptobium, Cylindroxystus, Dibelonetes, Dolicaon,
Domene, Echiaster, Eustilicus, Gnathymenus, Homaeotarsus, Hypero-
mma, Lathrobium, Lithocharis, Medon, Ochthephilum, Oedichirus,
Paederus, Palaminus, Pinophilus, Procirrus, Rugilus (=Stilicus), Scim-
balium, Scopaeus, Stilicopsis, Stiliderus, Sunius, etc.
830
*Divergent tribal/subtribal classifications have been proposed (e.g., by
Casey 1905, Blackwelder 1939, Pagel 1965) and not resolved. A world
tribal and generic revision is in preparation by L. Herman. Coiffait
(1972-84), Frank (1988), Herman (1981, 1991).
STAPHYLININAE Latreille, 1802: 124 (incl. Amblyopininae, Diochinae,
Platyprosopinae, Quediinae, Xantholininae, Xanthopyginae)
Amblyopinus, Anisolinus, Arrowinus, Atanygnathus (=Tanygnathus), Atre-
cus (=Baptolinus), Belonuchus, Cafius, Craspedomerus, Creophilus,
Diochus, Erichsonius, Gabrius, Glenus, Heterothops, Holisus, Leptaci-
nus, Neobisnius, Neohypnus, Ocypus, Ontholestes, Othius, Philonthus,
Platydracus, Platyprosopus, Quedius, Staphylinus, Thinopinus, Thyreo-
cephalus, Triacrus, Xantholinus, Xanthopygus, etc.
*The subfamily is used here in the broad sense (e.g., Arnett 1963,
Kasule 1970) to include Xantholininae of many authors and six subfami-
lies of Moore (1964). The presence of obtected pupae (unique among Sta-
phylinidae) and other larval and adult characters support monophyly of the
subfamily as a whole, but the tribal/subtribal classification needs revision,
especially for the groups near Staphylinini. Bordoni (1982), Coiffait (1972
-84), Seevers (1955), Smetana (1971, 1982, 1988a).
Series SCARABAEIFORMIA Crowson, 1960: 119
(=Lamellicornia)
*This series corresponds to the traditional Lamellicornia and not to Crowson's
series Scarabaeiformia, which also included Dascillidae (1960) and Rhipicer-
idae (1971a). Forbes (1926), Peyerimhoff (1933) and Jeanne} and Paulian
(1944) all united the Staphyliniformia and Scarabaeiformia, as here delimited,
into a group called the Haplogastra (originally coined by Kolbe 1908; see Law-
rence, Slipiri.ski and Pakaluk 1995), based mainly on a distinctive type of wing
venation and the presence of concealed lateral portions of the second abdomi-
nal sternite (absent in other adult polyphagans with the exception of some neo-
tenic groups). Boving and Craighead (1931) first suggested that the lamelli-
corns evolved from the Dascillidae, based on obvious larval features, such as
the well-developed head and antennae, large mandible with mol a and accessory
ventral process, free, articulated galea and cribriform spiracles. Crowson ( 1950)
recognized the Haplogastra, but in 1960 he combined the lamellicorns and
Dascillidae into a series Scarabaeiformia. Crowson (1971a) added the families
Rhipiceridae and Karumiidae to this series and noted the presence of a dascill-
oid-like dorsoventrally divided penis in the scarabaeoid genus Pleocoma, add-
ing at least one apparent adult synapomorphy to the list of larval similarities
between the two groups. Lawrence and Newton (1982) used the complex pro-
mesothoracic interlocking device and lack of a larval spiracular closing appara-
tus to argue for a link between Dascilloidea and typical Elateriformia; they
maintained that the larval similarities between dascillids and scarabaeoids are
831
either plesiomorphies or convergent adaptations for soil-dwelling, and that the
divided penis in dascilloids and Pleocoma may be homologous to that in Pse-
phenidae and some Scirtidae. Kukalova-Peck and Lawrence (1993) demon-
strated that Staphyliniformia and Scarabaeiformia share three major venational
synapomorphies not present in dascilloids or other members of the elateriform
lineage.
Superfamily SCARABAEOIDEA Latreille, 1802: 144
*A number of comparative morphological and phylogenetic studies in recent
years have led to the recognition of several scarabaeoid families in addition to
the six recognized by Crowson (1955). This has resulted mainly from the dis-
membering of the old Laparosticti (see Westwood 1845, 1852 and Janssens
1949), which was defined on the basis of primitive features, such as the loca-
tion of abdominal spiracles in pleural membrane. A general review of the sub-
ject is given by Scholtz (1990). Areekul (1957), Baraud (1985), Chalumeau
(1983), D'Hotman and Scholtz (1990), Rieke (1966, 1971), Hinton (1967a),
Howden (1982), Iablokoff-Khnzorian (1977b), Janssens (1946, 1949), Meine-
cke (1975), Moron (1984), Nel and De Villiers (1988), Nel and Scholtz (1990),
Paulian (1945, 1959-61), Paulian and Baraud (1982), Ritcher (1966, 1969a,
1969b, 1970), Ritcher and Baker (1974), Scholtz and Chown (1995, Woodruff
(1973), Yadav and Pillai (1979).
LUCANIDAE Latreille, 1804: 149
*Benesh (1960), Didier and Seguy (1953), Emden (1935), Holloway
(1960, 1968, 1969), Howden and Lawrence (1974), Kikuta (1986), Law-
rence (1981), Maes (1992), Mizunuma and Nagai (1994).
AESALINAE MacLeay, 1819: 102
Aesalus (Holarctic, Asia, Central America) and Lucanobium (northern
South America).
*Araya (1993), Howden and Lawrence (1974).
NICAGINAE LeConte, 1861: 130
Ceratognathus (Australia, New Zealand and Chile) and Nicagus (North
America and Japan).
*Kikuta (1986) placed Nicagus and Ceratognathus in the family Tro-
gidae on the basis of their non-geniculate antennae, presence of long setae
on the anterior part of the first antennomere, and the antenna} club consist-
ing of 3 coadapted lamellate segments. Martinez (1976).
SYNDESINAE MacLeay, 1819: 103 (incl. Sinodendrinae)
Ceruchus and Sinodendron (Holarctic ), and Syndesus (South America,
Australia, southwest Pacific).
LAMPRIMINAE MacLeay, 1819: 97
Dendroblax (New Zealand), Lamprima (Australia, New Guinea), Phala-
crognathus (Australia), and Streptocerus (Chile).

832
PENICHROLUCANINAE Arrow, 1950: 233
Penichrolucanus (southeast Asia, Melanesia) and Brasilucanus (South
America).
*Ratcliffe (1984).
LUCANINAE Latreille, 1804: 149 (incl. Chiasognathinae, Dorcinae, Figu-
linae, Odontolabinae, Platycerinae)
Chiasognathus, Colophon, Cyclommatus, Dendezia, Dorcus, Figulus, Lis-
sotes, Lucanus, Nigidius, Odontolabis, Pholidotus, Platycerus, Pro-
sopocoilus, Pseudodorcus, Rhaetulus, Rhyssonotus, Sclerostomus,
Scortizus, Serrognathus, etc.
*Moore (1978), Onore (1994), Scholtz and Endrody-Younga (1994).
PASSALIDAE Leach, 1815: 100
*Reyes-Castillo (1970), Schuster and Reyes-Castillo (1981).
AULACOCYCLINAE Kaup, 1868: 4
Aulacocyclus (Oriental, Australia) and Ceracupes, Comacupes, Cylin-
drocaulus, and Taeniocerus (Oriental).
PASSALINAE Leach, 1815: 100
Leptaulax, Mastochilus, Odontotaenius, Passalus, Paxillus, Popilius, Pro-
culus, Veturius, etc.
TROGIDAE MacLeay, 1819: 59
Omorgus (Africa, Australia, South America, southern North America and
Asia), Polynoncus (Neotropical) and Trox (Holarctic and Ethiopian).
*Baker (1968), Caveney and Scholtz (1993), Scholtz (1986, 1991),
Scholtz and Peck ( 1990), Scholtz et al. ( 1987 a).
GLARESIDAE Semenov-Tian-Shanskii and Medvedev, 1932: 337
Glaresis (all major continents except Australia).
*Scholtz et al. (1987b).
PLEOCOMIDAE LeConte, 1861: 128
Pleocoma (western North America).
*Paulian (1941b), Ritcher (1966).
DIPHYLLOSTOMATIDAE Holloway, 1972: 31
Diphyllostoma (western North America).
*Holloway (1972).
GEOTRUPIDAE Latreille, 1802: 142
BOLBOCERATINAE Mulsant, 1842: 347 (=Bolbocerinae; incl. Athyreinae)
Athyreus, Blackburnium, Bolbelasmus, Bolboceras, Bolbocerasoma, Ele-
phastomus, Eucanthus, Neoathyreus, etc.
833
*The structure of the axillary region of the hind wing suggests that this
taxon is more closely related to a group of families including Passalidae,
Lucanidae, Trogidae and Pleocomidae than it is to the next subfamily, and
thus should be accorded family rank (Browne 1993). Browne (1991) used
the same suite of characters to determine that Eucanthus is the sister group
of the remaining bolboceratines. Tribal and generic concepts have been
discussed by Howden (1992), Howden and Martfnez (1963) and Howden
and Cooper (1977). Krikken (1984a), Paulian (1939).
GEOTRUPINAE Latreille, 1802: 142 (incl. Taurocerastinae)
Frickius and Taurocerastes (southern South America), Ceratotrupes,
Chromogeotrupes, Geotrupes, Mycotrupes, Typhoeus, etc.
*Larval evidence provided by Howden (1982) and Howden and Peck
(1987) support the placement of both Taurocerastes and Frickius in this
subfamily. Zunino (1984a, 1984b).
LETHRINAE Mulsant and Rey, 1871: 417
Lethrus (Europe and Central Asia).
BELOHINIDAE Paulian, 1959: 40
Belohina (Madagascar).
*The single species on which this family is based is a wingless lapa-
rostict with 10-segmented antennae, a 3- segmented cupuliform club (with
the last two segments telescoped into the first) and concealed, membra-
nous labrum and partly membranous mandibles. Paulian and Lumaret
(1979).
OCHODAEIDAE Mulsant and Rey, 1871: 493
*Although Crowson (1981) tentatively included Belohina and the Tau-
rocerastinae in this family, the most recent revisions (Scholtz and Evans
1987, Scholtz et al. 1988) ignore these taxa, the latter of which is now
placed in Geotrupidae (see above). The only known larval ochodaeid is
that of Pseudochodaeus estriatus (Schaeffer) (Carlson and Ritcher 1974).
OCHODAEINAE Mulsant and Rey, 1871: 493
Codocera (Palearctic), Endognathus (Madagascar), Ochodaeus (wide-
spread, but not Australia), and Odontochodaeus (Madagascar).
CHAETOCANTHINAE Scholtz in Scholtz et al., 1988: 231
Chaetocanthus (southern Africa), Namibiotalpa (southern Africa), Pseu-
dochodaeus (western North America), and Synochodaeus (southern
Africa).
CERATOCANTHIDAE Cartwright and Gordon, 1971: 275 (=Acanthoceridae)
Ceratocanthus (=Acanthocerus), Cloeotus, Cyphopisthes, Madrasostes,
Philharmostes, etc.
834
*Ceratocanthids are usually distinguished from other scarabaeoids by
the ability of the adult to strongly deflect the head and pronotum and posi-
tion the highly modified legs in such a way as to form an .almost unmter-
rupted sphere; however, this compacti?n defense ~echan~sm also occurs
(to a lesser degree) in the related fam1ly Hybosondae (Lzparoch~us) and
is not developed in several termitophilous genera recently pla~ed m Cera-
tocanthidae by Howden and Gill (1988a, 1988b). Ceratocanth1ds are often
associated with termites, and the larvae of one species (Madrasostes kazu-
mai Ochi) is known to feed on a reddish-brown clay-like material associ-
ated with termite activity but possibly resulting from the action of a brown
rot fungus (Iwata et al. 1992). Paulian (1977, 1978). . *Paulian (1984), Paulian and Lumaret (1982).
**The name Ceratocanthidae was proposed to replace Acanthocendae
Lacordaire (1856: 155) which was based on a preoccupied generic name.
HYBOSORIDAE Erichson, 1847: 717
Anaides, Coelodes, Cryptogenius, Hybosorus, Liparochrus, Phaeochrous,
etc.
*Cryptogenius, formerly placed in Trogidae, was moved to this family
by Scholtz et al. (1987a). Allsopp (1984), Costa et al. (1988), Ide et al.
(1990), Kuijten (1985) .
GLAPHYRIDAE MacLeay, 1819: 76 (incl. Lichniidae)
Amphicoma,Anthypna, Cratoscelis, Dasychaeta, Glaphyrus, Lichnia, Lich-
nanthe, Toxocerus, etc.
*Machatschke (1959) divided this group into two families. Ritcher
(1966).
SCARABAEIDAE Latreille, 1802: 144 (incl. Aclopidae, Cetoniidae, etc.)
APHODIINAE Leach, 1815: 97 (incl. Aegialiinae, Aulonocneminae, Chiro-
ninae, Silluviinae, Termitotroginae)
Aegialia, Aphodius, Ataenius, Aulonocnemis, Chiron, Corythoderus, De-
marziella, Eremazus, Euparia, Oxyomus, Psammobius, Rhyparus, Rhys-
semus, Saprus, Silluvia, Termitotrox, Thinorycter, etc.
*Balthazar (1964), Cambefort (1987, 1991), Cornell (1967), Dellacasa
(1988), Gordon and Cartwright (1988), Hanski and Cambefort (1991),
Howden and Storey (1992), Jerath (1960), Stebnicka (1977, 1985, 1986).
SCARABAEINAE Latreille, 1802: 144 (incl. Coprinae)
Alloscelus, Ateuchus, Canthon, Copris, Coptodactylus, Deltochilum, Di-
chotomius, Eucranium, Eurysternus, Gymnopleurus, Heliocopris, Oni-
ticellus, Onitis, Onthophagus, Phanaeus, Pinotus, Scarabaeus, Sisy-
phus, etc.
*Balthazar (1963), Cambefort (1991), Edmonds (1972), Edmonds and
Halffter (1978), Ferreira (1968, 1972), Halffter and Edmonds (1982),
835
Hanski and Cambefort (1991), Howden and Young (1981), Matthews
(1972-76), Matthews and Stebnicka (1986), Zunino (1983).
PACHYPODINAE Erichson, 1840: 29
Pachypus (Europe).
. **The type genus, ~achypus Dejean, 1821, is preoccupied by Pachypus
Billberg, 1820 (Cetonunae) according to Neave (1939-40), but that name
is not in the Coleopterorum Catalogus or Agassiz (1846a, 1846b) and
seems to have been forgotten; further research is needed.
ORPHNINAE Erichson, 1847: 717
Aegidium, Chaetonyx, Orphnus, etc.
ALL~D!OSTOMAT~NAE Arrow, 1940: 16 (=Allidiostominae, Idiostominae)
Allzdzostoma (=Idzostoma) (South America).
**Idiostominae Arrow (1904: 747) is based on a preoccupied generic
name. The grammatically correct spelling of the subfamily name is Alli-
diostomatinae, not Allidiostominae as usually used.
DYNAMOPODINAE Arrow, 1911: 611 (=Dynamopinae)
Dynamopus (central Asia and northern Africa).
**The grammatically correct spelling of the subfamily name is Dy-
namopodinae, not Dynamopinae as usually used.
ACLOPINAE Milne-Edwards, 1850: 96
Aclopus (Brazil, Argentina), Neophaenognatha (Argentina), Phaenognatha
(Australia), and Xenaclopus (Borneo).
*Allsopp (1981, 1983).
EUCHIRINAE Hope, 1840b: 79
Cheirotonus (southeast Asia), Euchirus (Philippines, Indonesia), and Pro-
pomacrus (Greece to Iran, China).
*Lumaret and Tauzin (1992), Young (1989).
**The subfamily name has sometimes been erroneously attributed to
"Hope, 1837".
PHAENOMERIDINAE Erichson, 1847: 655 (=Phaenomerinae)
**The group name spelling Phaenomeridinae was adopted in Opinion
621 (ICZN 1962) as grammatically correct and to avoid homonymy with
a group name in Curculionidae based on Phaenomerus.
Oxychirus and Phaenomeris (Africa).
MELOLONTHINAE MacLeay, 1819: 79/Leach in Samouelle, 1819: 189
(incl. Hopliinae, Systellopodinae)
Ablabera, Automolus, Chasmatopterus, Dichelonyx, Diplotaxis, Heteronyx,
Hoplia, Liparetrus, Macrodactylus, Maechidius, Melolontha, Pachy-
dema, Phyllophaga, Phyllotocidium, Phyllotocus, Serica, Systellopus,
Xylonychus, etc.
*The tribal classification has been revised by Britton (1957, 1978,
1980, 1986, 1987, 1988, 1990).
836
RUTELINAE MacLeay, 1819: 69
Adoretus, Anomala, Anoplognathus, Bolax, Catalpa, Geniates, Pelidnota,
Phyllopertha, Plusiotis, Popilia, Repsimus, Rutela, Spodochlamys, etc.
*Machatschke ( 1972-7 4).
DYNASTINAE MacLeay, 1819: 64
Agaocephala, Cryptodus, Cyclocephala, Dynastes, Dyscinetus, Golofa,
Heteronychus, Hexodon, Ligyrus, Megasoma, Oryctes, Oryctoderus,
Pentodon, Phileurus, Strategus, Xyloryctes, etc.
*Endrodi (1985) included a key to world species with bibliographic
references to his earlier monographic volumes (beginning in 1966) and
other work on the subfamily.
CETONIINAE Leach, 1815: 99 (incl. Valginae, Trichiinae)
Cetonia, Coptomia, Cotinus, Cremastocheilus, Cryptodontes, Diaphonia,
Diplognatha, Euphoria, Goliathus, Gymnetis, Inca, Microvalgus,
Osmoderma, Phaedimus, Platygenia, Potosia, Schizorhina, Stenotarsia,
Trichiotinus, Trichius, Valgus, Xiphoscelis, etc.
*Holm and Marais (1992), Krikken (1984b), Marais and Holm (1992),
Miksic (1976-83).
Series ELATERIFORMIA Crowson, 1960: 120
(=Dascilliformia; incl. Eucinetiformia)
*This series is equivalent to the Dascilliformia as first defined by Crowson
(1950). The families Dascillidae and Rhipiceridae were later removed from this
series and placed with the lamellicoms in Scarabaeiformia (Crowson 1960,
1971a), and the Clambidae, Scirtidae (as Helodidae) and Eucinetidae were
transferred (as Eucinetoidea) to a new series Eucinetiformia (Crowson 1959,
1960, 1971b). The remaining dascilliform taxa were placed in a series Elateri-
formia (Crowson 1960), which has been more or less consistently applied to
this lineage for the past 35 years. Although we are defining the series in Crow-
son's original sense with Scirtoidea (=Eucinetoidea) and Dascilloidea included
as basal lineages, we prefer the term Elateriformia, because it has been consis-
tently used for most of the included taxa and because the term Dascilliformes
had already been used by Lameere (1938) for a group more equivalent to
Crowson's (1960) Scarabaeiformia. Lawrence and Newton (1982) applied the
term "elateriform lineage" to this group plus Scarabaeoidea, and Ponomarenko
and Zherikhin (1980) included the same taxa in their infraorder Elateromorpha.
Phylogenetic relationships among the taxa comprising Crowson's (1960)
original Elateriformia (Buprestoidea, Byrrhoidea and Elateroidea below) are
still not understood, although they have been reassessed on several occasions
over the past 20 years by Crowson (1972b, 1973b, 1975, 1978) and others
(Lawrence and Newton 1982, Lawrence 1988b, Lawrence and Britton 1991,
Lawrence, Nikitsky and Kirejtshuk 1995, Nikitsky et al. 1994). The group was
originally divided into six superfamilies: Byrrhoidea (Byrrhidae), Dryopoidea
837
(Psephenidae, Eurypogonidae, Ptilodactylidae, Chelonariidae, Heteroceridae,
Limnichidae, Dryopidae and Elmidae), Rhipiceroidea (Rhipiceridae and Calli-
rhipidae), Buprestoidea (Buprestidae), Elateroidea (Cebrionidae, Elateridae,
Throscidae [as Trixagidae], Cerophytidae, Perothopidae and Eucnemidae) and
Cantharoidea (Brachypsectridae, Omalisidae [as Homalisidae], Karumiidae,
Drilidae, Phengodidae, Telegeusidae, Lampyridae, Cantharidae and Lycidae).
Crowson (1971a) moved Rhipiceridae and Karumiidae out of the series and
into Dascilloidea (Scarabaeiformia) and later (1972b) added the families Cneo-
glossidae, Plastoceridae and Omethidae to the Cantharoidea. Crowson (1973b)
proposed a new superfamily Artematopoidea for the families Artematopodidae
(as Artematopidae, including Eurypogonidae), Callirhipidae and Brachypsectr-
idae. Kasap and Crowson (1975) added two more dryopoid families, Euli-
chadidae and Lutrochidae, but the latter was later considered as a subfamily of
Dryopidae (Crowson 1978).
Lawrence (1988b) redefined Elateriformia to exclude Scirtoidea (as Euci-
netoidea) and Scarabaeoidea and to include Dascilloidea (sensu Crowson
1971a). He also made the first attempt to subject this group to a cladistic analy-
sis. As a result of the analysis, he reorganized the elateriform families into four
superfamilies: Dascilloidea (Dascillidae, including Karumiidae, and Rhipi-
ceridae), Byrrhoidea (Buprestidae, Byrrhidae, Dryopidae, Lutrochidae, Elmidae,
Heteroceridae and Limnichidae), Psephenoidea (Psephenidae, Callirhipidae,
Eulichadidae, Cneoglossidae, Ptilodactylidae and Chelonariidae), and Elatero-
idea (Artematopodidae, Brachypsectridae and a new family, Rhinorhipidae,
plus the former Elateroidea and Cantharoidea). Lawrence and Britton (1991)
resurrected Buprestoidea and combined Psephenoidea and Byrrhoidea into a
single superfamily.
Lawrence, Nikitsky and Kirejtshuk (1995) analyzed a set of elateriform taxa
representing most of the above families plus several Scirtoidea using Lawren-
ce's (1988b) character set with some additions and modifications. The resulting
cladograms varied considerably with the outgroup used, but some clusters were
relatively robust. A broadly defined Elateroidea occurred in most of the dado-
grams, but always excluding Rhinorhipidae, which usually clustered with Das-
cilloidea. Byrrhoidea, in the sense of Lawrence (1988b) but without Bupre-
stidae, also occurred in some cladograms, but Psephenoidea was never sup-
ported, the taxa included by Lawrence either being attached to the base of Ela-
teroidea (Ptilodacty lidae, Chelonariidae, Eulichadidae, Callirhipidae) or forming
a separate clade near the base of the Elateriformia (Psephenidae, Cneogloss-
idae). Byrrhoidea, as delimited below, does not appear to be monophyletic, but
further changes in the classification are not warranted until stronger support can
be demonstrated for an alternative scheme.
838
Superfamily SCIRTOIDEA Fleming, 1821: 50
(=Eucinetoidea)
*Crowson (1960, 1971b) established the superfamily and series Eucinetoidea
and Eucinetiformia for three polyphagan families lacking the derived features
of Elateriformia, Staphyliniformia, Scarabaeiformia, Bostrichiformia and Cucu-
jiformia and sharing the following attributes: (1) aedeagus with parameres not
distinctly articulated, (2) 9th abdominal segment with pleurites fused together
in front oftergite (see below), (3) with characteristic metendosternite, (4) larva
with mandibular mala, (5) distinct but non-articulated larval galea and lacinia,
(6) larval spiracles with closing apparatus, (7) larva without urogomp~i, (8)
larval tergites and sternites similarly sclerotized. The second feature IS best
interpreted as the presence of a distinct tenth tergite and undivided ninth tergi~e
(Rieke 1971, Lawrence 1988b), which is plesiomorphous. The group, a~ on-
ginally defined, is also characterized by the relatively small pronotum, highly
reduced prosternum, and a compaction mechanism involving the opistho-
gnathous head resting against the procoxae or in some cases the meso- or meta-
thorax (Hlavac 1975). The head is either flattened beneath or concave between
a pair of sharp genal rid~es. The new family ~ecliniidae (Nikitsky e; al. ~9_94)
is based on female specimens only and thus Its effect on Crowson s ongmal
diagnosis of the group (the metendosternite being of the same type) is un-
known; however, Declinia has a well-developed prosternum, and the head,
though flattened beneath, is hypognathous and rests against the visible proster-
num.
** Scirtoidea Fleming, 1821 and Clamboidea Fischer, 1821 have priority over
Eucinetoidea Lacordaire, 1857 for the name of the superfamily. Fleming's
work dates from July, Fischer's has no exact date so takes the date 31 Decem-
ber (ICZN 1985a, Art. 21(c)).
DECLINIIDAE Nikitsky et al., 1994: 7
Declinia (Far Eastern Russia).
*These beetles, known only from females, look superficially like Scirt-
idae in their broad and somewhat flattened body and shortened pronotum;
both prosternum and propleurotrochantin, however, are well-developed and
broadly exposed as in Dryopidae and some other byrrhoids. Lawrence,
Nikitsky and Kirejtshuk (1995).
EUCINETIDAE Lacordaire, 1857: 281 (=Cryptomeridae)
Bisaya (Central Asia), Eucilodes (Eurasia), Eucinetus (=Apeosina, Crypto-
mera) (widespread), Euscaphurus (western North America), Jentozkus
(New World), Subulistomella (Japan), and Tohlezkus (Turkey and Fin-
land). .·
*This is a well-defined and more or less uniform group adults of which
have a streamlined, elliptical body with greatly enlarged, oblique meta-
839
coxal plates and an ability to jump. Sucking mouthparts appear to have
evolved on several occasions in the family. Sakai (1980), Vit (1977a,
1977b, 1985, 1990), Wheeler and Hoebeke (1984).
CLAMBIDAE Fischer, 1821: 52 (incl. Calyptomeridae)
*This family is relatively well defined, although Forbes (1926) and
Crowson (1955) excluded Calyptomerus. Acalyptomerus (Crowson 1979)
differs from most clambids in having a smaller head and less well devel-
oped compaction mechanism. EndrOdy-Younga (1959, 1960, 1961, 1981,
1990a, 1990b, 1993).
CALYPTOMERINAE Crowson, 1955: 11
Calyptomerus (Holarctic, introduced into Australia).
ACALYPTOMERINAE Crowson, 1979: 612
Acalyptomerus (Asia, Africa, Central and South America).
CLAMBINAE Fischer, 1821: 52
Clambus (cosmopolitan), Loricaster (North America and Europe), and
Sphaerothorax (Australia, New Zealand, Chile).
SCIRTIDAE Fleming, 1821: 50 (=Cyphonidae, Helodidae, Elodidae)
Amplectopus, Atopida, Byrrhopsis, Cyphanodes, Cyphanus, Cyphotelus,
Mesocyphon, and Veronatus (New Zealand); Macrocyphon, Macro-
dascillus, Macrohelodes, Peneveronatus, Pseudomicrocara, etc. (Aus-
tralia); Ora (New World); Cyphon, Elodes (=Helodes), Microcara,
Prionocyphon, and Scirtes (widespread); etc.
*This is a worldwide group with the greatest diversity in south temper-
ate regions. In the Junk Catalogue (Pic 1914) scirtid genera are placed in
the families Dascillidae (Cyphotelus, Veronatus, Cyphanus, Cyphanodes
and Atopida) and Helodidae (most of the genera, including Amplectopus).
Sharp (1878b) divided the diverse New Zealand fauna into two groups
(those with a distinct antennal fossa and those without or with a weak
fossa), and Pic (1914) placed Atopida in a separate tribe; otherwise there
has been no attempt to subdivide the family. Amplectopus is probably the
most distinctive genus. Sharp (1886) placed it in a broadly defined Dascill-
idae and considered it to be related to Chelonarium. Kasap and Crowson
(1975) placed the genus in Chelonariidae in a list of specimens examined,
and noted the fusion of the first three ventrites in both Chelonarium and
Amplectopus. Amplectopus superficially resembles chelonariids in the
highly compacted body and unusual antennae, but the male genitalia are
definitely of the distinctive scirtid type described in detail by Nyholm
(1969, 1972). The antennal modifications are entirely different in the two
groups. In Amplectopus antennomere 2 is small, subquadrate and heavily
sclerotized, 3 is longer, triangular, widened apically and lightly sclerotized,
4-11 are heavily sclerotized, 4 large and irregularly quadrate, 5 smaller
840
and subtriangular, 6 subquadrate, 7-10 strongly transverse and 11 about
as long as wide and rounded apically; in Chelonarium both 2 and 3 are
large and heavily sclerotized while the remainder are smaller and lightly
sclerotized, 2 is subquadrate and 3 elongate, 4 is very small and 5-11 are
elongate and slightly serrate. In Amplectopus the antennae are concealed
beneath the forelegs, while in Chelonarium antennomere 3 fits into the
mesosterna! cavity on top of the prosternal process, while 4--11 lie on the
metasternum. Amplectopus also resembles most scirtids and not chelo-
nariids in having strongly developed subantennal fossae and sharp sub-
genal ridges, non-exocone eyes, well-developed, abruptly curved mandi-
bles, basally prolonged trochanters, highly reduced prosternum, similar
wing venation and folding, and a pad beneath the penultimate tarsomere.
The differences between Amplectopus and the remaining Scirtidae may
justify the former's placement in a separate subfamily, but this should be
deferred until a more detailed comparative study can be carried out. Han-
nappe! and Paulus (1987, 1992), Klausnitzer (1974).
**Scirtidae has priority over Cyphonidae Stephens (1829a: 11) and
Elodidae Shuckard (1840: 41) for the name of the family; Helodidae
Agassiz (1846b: 176) is based on the unjustified emendation Helodes
which is also a junior homonym (Pope 1976).
Superfamily DASCILLOIDEA Guerin-Meneville, 1843: 193 (1834)
*This group, as here constituted, is that of Crowson (1971a), but its placement
in Elateriformia (and not Scarabaeiformia) is based on Lawrence (1988b). All
members of this group have soil-dwelling larvae and relatively short-lived,
surface-active adults. The grub-like detritivorous dascilline larvae probably
represents the ancestral type, from which the termitophilous karumiines and
ectoparasitic rhipicerids evolved.
**Rhipiceridae and Dascillidae both date from 1834, but the former name
may have priority for superfamily name since the relevant names were estab-
lished in the first and second numbers, respectively, of the same journal volume
(Annates de la Societe Entomologique de France, Vol. 3). In the absence of
information on the exact or relative publication dates of these two journal num-
bers, we continue use of Dascilloidea for the superfamily name.
DASCILLIDAE Guerin-Meneville, 1843: 193 (1834) (incl. Karumiidae)
*Many genera formerly placed here are now in the families Scirtidae,
Ptilodactylidae, Psephenidae, and Artematopodidae.
**"Guerin-Meneville, 1823" has been cited in error for the family
name, which was actually proposed in 1843 to replace Atopidae Laporte
(1834: 227) whenAtopa was placed as a synonym of Dascillus. We main-
tain Dascillidae over Atopidae by provision of Art. 40b.
841
DASCILLINAE Guerin-Meneville, 1843: 193 (1834) (=Atopinae, Cinna-
bariinae)
Coptocera (=Cinnabarium) (Asia and Africa); Dascillus (western North
America, West Indies, Eurasia, East Indies); Notodascillus (Australia);
Haematodes, Pseudolichas and Sinocaulus (Asia).
.KARUMIINAE Escalera, 1913: 320 (=Zarudniolidae)
Anorus (southwestern North America), Drilocephalus (Argentina), Esca-
lerina (Iran), Emmita and Genecerus (North Africa), Karumia (=Zarud-
niola) (Iran and Afghanistan), and Pleolobus (Chile).
*This group was described as a separate family and has been treated
as such by most authors (Arnett 1964, Crowson 1955, 1971a, Mandl1974,
Paulus 1972a, Solervicens 1991). It originally included a few highly modi-
~ed, sexu~lly d~morphic and apparently termitophilous groups occurring
m most arid regwns of the world and was placed among the Cantharoidea;
however, Crowson (1971 a) first recognized its dascilloid affinities and
added the generaAnorus, Emmita, Genecerus and Pleolobus from Dascill-
idae. When intermediate forms like Genecerus and Anorus are added to
this group, there is little to justify its separation as a distinct family. Imma-
ture stages have not been described, but a putative larva of Pleolobus from
Chile has all of the features of a typical dascilline, with short, acute uro-
gomphi like those of Dascillus cervinus (Linnaeus).
Paulus (1972a) included Cydistus in this group, but Crowson (1972b)
placed that genus in his expanded Phengodidae. Although Escalerina mi-
crocephala (Escalera), the type of the genus, probably belongs here, E.
serraticornis Paulus, transferred to Paulusiella by Mandl (1974), is proba-
bly a cebrionine elaterid, as are the other species of Paulusiella described
by Mandl (see below).
RHIPICERIDAE Latreille, 1834: 167 (=Sandalidae)
Arrhaphipterus (southern Europe, North Africa, Asia Minor), Chameor"
hipis (Africa), Polymerius (Chile), Rhipicera (Australia, New Caledo-
nia, South America), and Sandalus (North and South America, East
and South Africa, Asia).
*Th:re has been some confusion about this family name, since Boving
an~ Craighead (1931) used Rhipiceridae for the callirhipid Zenoa and San-
dahdae for Sandalus, a true rhipicerid. Also van Emden's works on "San-
dalidae" (1924b, 1931, etc.) deal mainly with Callirhipidae. Crowson in-
cluded both _gro~ps in Rhipiceroidea in 1955, but later (1971a, 1973b)
pla~ed .them m. di~fere~t superfamilies. Evidence for the inclusion of rhipi-
cends m Dascillmdea IS based almost entirely on adults but it appears to
be well founded. The only known rhipicerid larvae are those of a North
American Sandalus (Craighead 1921, Elzinga 1977), but both the triun-
842
gulin-like first instars and ectoparasitic late instars are highly specialized.
Lawrence (1991).
Superfamily BUPRESTOIDEA Leach, 1815: 85
*Lawrence (1988b) combined this with the following superfamily, and Crow-
son (1982) suggested that the closest affinities of Buprestidae are with the fam-
ilies Elmidae and Dryopidae.
BUPRESTIDAE Leach, 1815: 85
*Several attempts have been made in recent years to reassess the higher
classification of Buprestidae, and general summaries have been published
by Nelson (1982), Bellamy (1985, 1986) and Holynski (.1988, 1993}.
Toyama (1987) suggested combining the traditionally recogmzed subfami-
lies Chalcophorinae and Buprestinae, and Holynski (1988, 1993) recog-
nized only four main buprestid lineages. Bright (1987).
SCHIZOPODINAE LeConte, 1861: 176
Dystaxia, Glyptoscelimorpha (=Dystaxiella), and Schizopus (western North
America).
*The wing venation prompted Forbes (1942) to place schizopodines in
the family Dascillidae, and the peculiar larva was used by Rees (1941) to
give the taxon family status (see Lawrence 1991): ~elson and _Bellamy
(1991) consider the group to be sister to the remammg Buprestldae, but
this alone does not necessitate its elevation to family rank. Lawrence
(1991).
**The type genus Schizopus has been conserved over a homonym of
the same year (ICZN 1993b).
JULODINAE Lacordaire, 1857: 10
Aata (Asia Minor), Amblysterna (eastern Africa), Julodella (Asia Minor,
Africa), Julodis (Asia Minor, Africa), Neojulodis (Africa), and Sterno-
cera (India, southeast Asia, Asia Minor, Africa).
*Bily (1983), Holm (1979).
**According to Neave (1939-40), the original spelling of the type ge-
nus is Jalodis; this requires further research (see discussion of problems
with "Eschscholtz 1829" names in Newton and Thayer 1992: 25).
BUPRESTINAE Leach, 1815: 85 (incl. Acmaeoderinae, Chalcophorinae,
Chrysobothrinae, Mastogeniinae, Polycestinae, Sphenopterinae, Thri-
ncopyginae, etc.)
Acherusia, Acmaeodera, Acmaeoderoides, Actenodes, Agaeocera, Anth-
axia, Astraeus, Bubastes, Buprestis, Castiarina, Catoxantha, Chal-
cophora, Chrysobothris, Chrysodema, Cinyra, Conognatha, Cyphog~­
stra, Diadoxus, Dicerca, Epistomentis, Helferella, Hippomelas, Julodz-
morpha, Mastogenius (=Haplostethus), Melanophila, Melobasis, Nas-
cio, Nothomorpha, Polycesta, Polyctesis, Prospheres, Psiloptera, Pto-
843
sima, Sphenoptera, Stigmodera, Thrincopyge, Tylauchenia, Tyndaris,
Xyroscelis, etc.
*Bellamy (1986), Cobos (1980-81), Gardner (1990).
**Dates for recently added former subfamilies are Acmaeoderinae Ker-
remans (1893: 112), Chalcophorinae Lacordaire (1857: 14), Chryso-
bothrinae Laporte and Gory (1837: 1), Mastogeniinae LeConte and Hom
(1883: 199), Polycestinae Lacordaire (1857: 61), Sphenopterinae Lacor-
daire 1857: 68), and ThrincopyginaeLeConte (1861: 154). Haplostethinae
LeConte (1861: 155) has priority over Mastogeniinae, but the latter name
could be maintained by provision of Art. 40b. Chrysochroinae Laporte
(1835: 158) has priority over Chalcophorinae as well as the name of the
included tribe Catoxanthini Jacobson (1913: 778) (Chrysochroa is listed
as a subgenus of Catoxantha by Bellamy (1985)); retention of Catoxanth-
ini might possibly be justified by Art. 40b, but this name would then take
precedence over Chalcophorinae for the subfamily name.
AGRILINAE Laporte, 1835: 165 (incl. Cylindromorphinae, Galbellinae,
Trachyinae)
Agrilus, Brachys, Cisseis, Coroebus, Cylindromorphus, Ethon, Galbella,
Germarica, Habroloma, Meliboeus, Pachyschelus, Taphrocerus, Tra-
chys, etc.
*Bfly (1983),Volkovitsch and Hawkeswood (1990).
**Dates for recently added former subfamilies are Cylindromorphinae
Portevin (1931: 335), Galbellinae Cobos (1986: 71) (Bellamy 1985: 428,
nomen nudum), and Trachyinae Laporte (1835: 166).
Superfamily BYRRHOIDEA Latreille, 1804: 146
(incl. Dryopoidea, Psephenoidea)
*Crowson (1955, 1960, 1973b) restricted this group to the single family Byrrh-
idae, placing the remaining taxa below in the Dryopoidea. Lawrence (1988b)
united Byrrhidae with the next 5 families (Elmidae-Heteroceridae), but placed
the remaining groups (Psephenidae-Callirhipidae) in Psephenoidea. As men-
tioned above, some of the latter group of families may be more closely related
to Elateroidea (Lawrence, Nikitsky and Kirejtshuk 1995). Brown (1981b),
Crowson (1978), Hinton (1939a, 1955).
BYRRHIDAE Latreille, 1804: 146 (incl. Syncalyptidae)
*As noted by Watt (1971), this family is in drastic need of a world
revision based on adults and immature stages. Crowson (1955) placed the
syncalyptines with Pedilophorus in a subfamily Pedilophorinae. A number
of new genera remain to be described from the southern temperate regions.
El Moursy (1961), Johnson (1987, 1991), Paulus (1972c).
844 845

BYRRHINAE Latreille, 1804: 146 . LUTROCHIDAE Kasap and Crowson, 1975: 442
Byrrhus, Cytilus, Liochoria, Lioligus, Lioon, Morychus, Pedzlophorus, Lutrochus (North and South America).
Simplocaria, etc. *Crowson (1978) treated this group as a subfamily of Dryopidae.
SYNCALYPTINAE Mulsant and Rey, 1869: 31 . Brown (1991), Brown and Murvosh (1970), Hinton (1939c).
Chaetophora (=Syncalypta), Curimopsis, Microchaetes (Australia), and **Kasap and Crowson (1975: 442) were apparently the first to use the
Sierraclava (western North America). family-group name and provide enough comparative notes to make it
*C. Johnson (1978), P.J. Johnson (1982). available, but they did not indicate the name as new or formally describe
AMPHICYRTINAE LeConte, 1861: 111 the group.
Amphicyrta (western North America).
LIMNICHIDAE Erichson, 1846: 465
ELMIDAE Curtis, 1830: Plate 294 (text) (=Elminthidae, Helminthidae, El- *Brown (1991), Hinton (1939b, 1939c), Paulus (1970b), Sate (1966),
mididae, Limniidae) Wooldridge (1975, 1986).
*Hinton (1940), Olmi (1976), Steffan (1961). HYPHALINAE Britton, 1971: 88
**Jach (1994) noted that the older name Limniidae Steph~ns (1828: Hyphalus (Australia and New Zealand).
104) is unavailable (Art. llf(i)1), and has propo~ed conservatl?n of the *Britton (1971, 1977).
usual spelling of the family name, Elmidae. Brulle ( 1835: 314) IS usually LIMNICHINAE Erichson, 1846: 465 (incl. Bothriophorinae)
cited as author of Elmidae. Byrrhinus, Limnichites, Limnichoderus, Limnichus, Paralimnichus, Pelo-
LARAINAE LeConte, 1861: 116 (=Larinae) chares, Physemus, etc.
Hydora, Lara (western North America), Ovolara (Australia), Phanocerus, *Most limnichids are included in this subfamily. Kasap and Crowson
Potamophilus, etc. . (1975) placed the genus Paralimnichus in Chelonariidae but Wooldridge
*Brown (1981a), Spangler and Brown (1981), Spangler and Santlago- (1983) retained it in Limnichidae and we agree. The distinctive divided
Fragoso (1992). . . . eye and tarsal lamellae distinguish Paralimnichus from all other limni-
**The original group name Larinae was emended to Lar~mae m Opm- chids; however, most of its features place it in this subfamily. Lawrence
ion 1515 (ICZN 1988) to avoid confusion with the bird family name Lar- (1987a), Wooldridge (1977a, 1977b, 1980, 1981a, 1982, 1987).
idae. CEPHALOBYRRHINAE Champion, 1925: 174
ELMINAE Curtis, 1830: Plate 294 (text) Cephalobyrrhus (Asia), Par'athroscinus (Asia), and Throscinus (New
Ancyronyx, Austrolimnius, Elmis, Macronychus, Stenelmis, etc. World).
*Hinton (1939c, 1968, 1971). *Wooldridge (1981b, 1984).
THAUMASTODINAE Champion, 1924a: 25
DRYOPIDAE Billberg, 1820: 38 (1817) (=Parnidae; incl. Chiloeidae) Acontosceles and Pseudeucinetus (=Thaumastodus) (Indo-Australian), and
Dryops (=Parnus), Elmoparnus, Geoparnus, Helichus: Oreoparnus, Pelo- Martinius and Mexico (Neotropical).
nomus, Protoparnus, Sostea, Sosteamorphus (=Chzloea), Stygoparnus, *Spilman (1959, 1972), Wooldridge (1988).
etc. .
*Dryopids were often lumped with Elmidae an_d Lutrochidae in earlier HETEROCERIDAE MacLeay, 1825: 34
classifications. Dryopids occur in leaf litter and sml throughout most parts *Charpentier (1968), Pacheco (1964).
of the world (but not Australia); however, some (e.g., Helichus) are aq- **"Latreille, 1810" has been listed in error for the family name (e.g.,
uatic as adults and at least one (Stygoparnus) occurs in subterranean wa- by Handlirsch 1925).
ters (Barr and Spangler 1992). Besuchet (1978), Ballow (1936), Brow~ ELYTHOMERINAE Pacheco, 1964: 120
(1970, 1991), Dajoz (1973), Deleve (1965), Hinton (193~b, 1937), Olm1 Elythomerus (Australia).
(1976), Perkins and Spangler (1985), Steffan (1961), U~nch (1987a). HETEROCERINAE MacLeay, 1825: 34
**We maintain the family name Dryopidae over Parmdae Leach ( 1817: Auglyes, Heterocerus, Lanternarius, Micilus, Tropicus, etc.
88) by provision of Art. 40b.
846
PSEPHENIDAE Lacordaire, 1854: 497 (incl. Eubriidae, Psephenoididae)
*Eubriinae and Eubrianacinae were placed in Dascillidae by Arnett
(1963); Bertrand (1972) treated Eubriinae and Psephe~oidinae as separate
families and included Eubrianacinae in Dascillidae. Hmton (1955, 1966).
**The older name Eurypalpidae LeConte (1852a: 41) is based on a
preoccupied generic name (Eurypalpus Dejean, not Macquart).
EUBRIINAE Lacordaire, 1857: 283
Afroeubria (Africa), Dicranopselaphus (Neotropical), Ectopria (New
World), Eubria (Eurasia), Schinostethus (Asia), Sclerocyphon (Austra-
lia), Tychepsephus (Chile), etc.
*Davis (1986), Lee and Yang (1993).
PSEPHENOIDINAE Hinton, 1939a: 167
Psephenoides (Asia).
EUBRIANACINAE Jacobson, 1913: 726 (1880) (=Placonychinae)
Eubrianax (=Placonycha) (all continents except Australia, mainly Eur-
asia). . . . ,.
**Placonychinae Horn ( 1880a: 11 0) has pnonty over Eubnanac.I~ae for
the name of this group, but we maintain the latter name by provlSlon of
Art. 40b.
PSEPHENINAE Lacordaire, 1854: 497
Mataeopsephus (Palearctic, Oriental), Pheneps (~eotrop~cal), Psephenops
(Neotropical), Psephenus (Nearctic, Neotropical, Onental), etc.
CNEOGLOSSIDAE Champion, 1897: 594
Cneoglossa (Neotropical). . *Thi~ gr?up is based mainly on the distinctive larval respiratory sys-
*Crowson (1972b) placed this genus in the cantharoid complex (Plasto-
ceridae to Cantharidae below), but lack of functional 8th spiracles, to-
gether with the type of wing venation suggest a position n~ar P~ilodactyl­
idae or Psephenidae. Brown (1981 b) included Cneoglossa m a hst of pse-
phenid genera. Immature stages are unknown.
PTILODACTYLIDAE Laporte, 1836: 21
*This is a large and diverse family, most of wh.ich hav.e be~n p~aced m
Dascillidae at one time or another. The supragenenc classificatiOn IS badly
in need of revision, and the first subfamily is almost certainly paraph~letic,
as suggested by a preliminary cladistic analysis based on adult attributes
(Stribling 1986b). .
**Stribling's (1986b) thesis includes names of new higher taxa but was
not set up to meet publication criteria of the Code.
ANCHYTARSINAE Champion, 1897: 593
Anchycteis (northwestern North America, Japan), Anchytarsus (=Tetra-
glossa) (New World), Byrrocryptus (Australia, New Zealand), Daemon
847
(Madagascar), Epilichas (Asia), Octoglossa (Neotropical), Pseudoepi-
lichas (Japan), etc.
*This group contains those taxa which lack the special features of the
oth~r subfamilies and would probably be dismembered if the family were
subjected to a thorough phylogenetic analysis. Those genera for which
lar:~e are known, including Anchytarsus, Anchycteis, Byrrocryptus and
E_ptlzchas, do possess one apparent synapomorphy in the longitudinal divi-
siOn of the expanded submentum into three parts (Lawrence and Stribling
1992). Stribling (1986a).
**Epilichadinae Lawrence and Stribling (1992: 19, 25) is a nomen
nudum.
CLADOTOMINAE Pic, 1914: 45
Austrolichas (Australia), Cladotoma (South America), Paralichas (=Od-
ontonyx) (Holarctic), Hovactyla (Madagascar), etc.
. ~La~rence and Stribling (1992) defined this group on the basis of a
dtstmctlve larval type bearing a spiracular siphon and a combination of
adult features, including a transverse labrum, inflated pronotal disc, com-
plete lateral pronotal carinae and a reduced wedge cell in the hind wing.
At least some of these features suggest a relationship to the Chelonariidae.
APLOGLOSSINAE Champion, 1897: 623 (=Haploglossinae)
Aploglossa (=Haploglossa) (Neotropical).
ARAEOPIDIINAE Lawrence, 1991: 250
Araeopidius (western North America).
ter.n, whtch mvolves plastron plates associated with all but the last pair of
spiracles (Lawrence 1991 ).
**The name Araeopidiinae was made available by Lawrence (1991:
250, ~91, 394~ b~ including the larva of Araeopidius in a key and in com-
parati~e descn~tlve notes.' ~?d by indicating thatAraeopidius was the only
genus mcluded m Araeoptdunae; the subfamily was not formally described
or indicated as new.
. PTILODACTYLINAE Laporte, 1836: 21
Chelonariomorphus (South America), Falsotherius (Southeast Asia East
Indies), Stirophora (Neotropical), Pherocladus (East Indies, Fiji), Lach-
~od~ctyla (New World), Ptilodactyla (widespread), etc.
Thts .group is charact.erized by the concealment of the protrochantin
by postenor late~al ~xtenswns of the prosternum, which meet the pronotal
(hypomeral) projectiOns. Larvae are also of a distinctive type, which are
more clearly terrestrial in habits than those of the other subfamilies.
CHELONARIIDAE Blanchard, 1845b: 70
Brounia (New Zealand), Chelonarium (tropicopolitan), and Pseudo-
chelonarium (southeast Asia).
848
*The New Zealand Brounia thoracica Sharp (1878a) has pectinate
antennae and resembles some members of the Callirhipidae more than it
does species of Chelonarium or Pseudochelonarium; however, the serrate
tibiae, tarsal structure, wing venation and male genitalia are definitely of
the chelonariid type. Mequignon (1934, 1935), Paulus (1970a), Spangler
(1980a).
EULICHADIDAE Crowson, 1973b: 237 (=Lichadidae)
Eulichas (=Lichas) (Asia) and Stenocolus (western North America).
*Forbes (1926) proposed Lichadidae for the generaLichas (=Eulichas),
Callirhipis, and Zenoa, based on their distinctive wing structure. The
unassociated larva of Eulichas was illustrated by Boving and Craighead
(1931) who considered it to be an aberrant ptilodactylid. Crowson (1960)
included Eulichas in his discussion of Dryopoidea, and later (Crowson
1973b) gave distinguishing characters of Eulichadidae and discussed its
similarities to both Ptilodactylidae and Callirhipidae. Kasap and Crowson
(1975), Lawrence (1991). ' the presence of only four Malpighian tubules, absence of a mandibular mola
**Crowson (1973b: 237) apparently was the first to use the name Euli-
chadidae, as an implied replacement name for Lichadidae Forbes (1926:
102) which was based on the preoccupied generic name Lichas; Lichad-
idae and the synonymy of Lichas with Eulichas had been mentioned ear-
lier in the same work (Crowson 1973b: 235).
CALLIRHIPIDAE Emden, 1924b: 87
Callirhipis, Celladonia, Ennometes, Horatocera, Zenoa, etc. . the well-developed and freely articulated basal abdominal segments are the
*This and the preceding family appear to share a number of attributes
with the previous groups (especially Ptilodactylidae and Chelonariidae)
and others with the superfamily Elateroidea. Callirhipids were traditionally
included in Rhipiceridae (Sandalidae of Emden 1924b, 1931, 1932b), but
Forbes ( 1926) considered the wing structure to be very similar to that
found in eulichadids (as Lichadidae), and Boving and Craighead (1931)
considered callirhipid larvae to be separable at the family level from those
of Sandalus and included them in the superfamily Elateroidea. Crowson
(1950) recognized two families, Rhipiceridae and Callirhipidae, within a
superfamily Rhipiceroidea, but later (Crowson 1971 a) placed Rhipiceridae
in the superfamily Dascilloidea (with Karumiidae and Dascillidae) and
moved Callirhipidae to the superfamily Artematopoidea (with Artemato-
podidae and Brachypsectridae) (Crowson 1973b). More recently they have
been placed near Eulichadidae, either in Dryopoidea (Lawrence and ~ew­
ton 1982), Psephenoidea (Lawrence 1988b) or an expanded Byrrh01dea
(Lawrence and Britton 1991, 1994). In cladograms produced by Lawrence,
Nikitsky and Kirejtshuk (1995), Eulichadidae and Callirhipidae usually
849
cluster with ptilodactylids and chelonariids at the base of the Elateroidea
as delimited below.
**"Lacordaire, 1857'' has been cited by Arnett (1963) and others for
the family name, but we can find no family-group name in association
with the treatment of Callirhipis by Lacordaire (1857: 249, 568), or any
other use earlier than Emden (1924b: 87). Zenoidae LeConte (1866a: 50)
then has priority for the name of this group, but further investigation is
needed; if necessary, we recommend application be made to the ICZN to
conserve the well-known name Callirhipidae over LeConte's unused name.
Superfamily ELATEROIDEA Leach, 1815: 85
(incl. Cantharoidea, Artematopodoidea)
*The monophyly of Artematopodoidea + Elateroidea + Cantharoidea was sug-
gested by Crowson (1972b, 1973b) and Lawrence and Newton (1982) and the
lumping of the three superfamilies (with the exception of Cneoglossidae, see
above) was formally proposed by Lawrence (1988b). Synapomorphies include
and transverse metasternal suture, presence of a single pair of large stemma in
the larva (absent in some) and the derived type of larval feeding mechanism
involving extraoral digestion and the reduction, loss, and/or fusion of various
structures, such as the labrum, mandibular mola, maxillary lobes, and maxillo-
labial sclerites. The connation of four ventrites is usually considered to be part
of the elateroid groundplan, but this character breaks down in the soft-bodied,
"malacoderm" forms comprising the old Cantharoidea, where it is assumed that
result of neoteny. The Cantharoidea are generally considered to be a monophy-
letic group, and published phylogenies (Crowson 1972b, Pototskaya 1983)
make this assumption; however, it is difficult to find distinguishing features
which are not either plesiomorphic or connected with the "malacoderm" facies.
The difficulties associated with placing highly derived, lightly sclerotized males
is demonstrated by the previous placement of Karumiidae in Cantharoidea and
the inclusion in that family of the elaterid genus Paulusiella (Crowson 1955,
Paulus 1972a, Mandl 1974).
**Cebrionoidea has priority over Elateroidea for the superfamily name, and
was actually used by Zherikhin in Arnoldi et al. (1977) and Ponomarenko and
Ryvkin (1990), but we maintain Elateroidea here (see under Elateridae).
ARTEMATOPODIDAE Lacordaire, 1857: 260 (=Artematopidae; incl. Eury-
pogonidae)
*Members of this group were included in Dascillidae in older classifi-
cations, but Crowson moved them to Dryopoidea (1955) and then included
them in a separate superfamily, along with Callirhipidae and Brachypsec-
tridae (Crowson 1973b).
850
**The grammatically correct spellings of the family and nominotypical
subfamily names are Artematopodidae and -inae (e.g., Paulian 1988),
rather than the more common Artematopidae and -inae.
ELECTRffiiiNAE Crowson, 1975: 77
Electribius (Mexico and Central America; Oligocene of northern Europe).
*This group was originally described from Baltic amber (Crowson
1973b), but is now known from the Recent fauna of Mesoamerica
(Lawrence 1995a).
**Crowson (1975: 77) referred to a group "Electropogonini" including
"Electropogon", but this generic name is clearly a lapsus calami for Elec-
tribius Crowson, which is correctly named in the referred-to Fig. 30 of
Crowson (1975) and in his earlier work (Crowson 1973b) cited in the
same sentence. This lapsus itself does not prevent use of the tribal name
after correction to "Electribiini", if the family-group name is otherwise
available. The group was not described by Crowson (1975) but he referred
there to his earlier description of the genus Electribius (Crowson 1973b)
and (supposedly) the tribe Electribiini. Although he did not actually' name
a tribe in 1973, his referral to the key constitutes a sufficient indication to
validate the name, provided the characterization of the genus in the key is
also taken as the tribal characterization (since Electribius was the only
included genus). The group was formally described by Lawrence (1995a).
ARTEMATOPODINAE Lacordaire, 1857: 260 (incl. Ctesibiinae, Eury-
pogoninae, Macropogoninae)
Allopogonia (California), Artematopus (Central and South America), Car-
cinognathus (South America), Ctesibius (Mexico and Central America),
Eurypogon (incl. Pseudodactylus) (North America, Italy, Japan), and
Macropogon (North America, Asia).
*Lawrence (1995a) included Ctesibiinae Crowson (1973b: 228) and
Macropogoninae LeConte (1861: 178) (=Eurypogoninae Boving and
Craighead 1931: 45) in this subfamily. Costa et al. (1985), Cooper (1991).
BRACHYPSECTRIDAE LeConte and Hom, 1883: 170
Brachypsectra (southwestern North America, southern India, Malaysia,
northern Australia).
*This group has been included in Dascilloidea, Artematopodoidea,
Elateroidea and Cantharoidea (Blair 1930, Crowson 1973b, Kasap and
Crowson 1975, Lawrence 1988b). Species of Brachypsectra have been
described from southwestern North America, India and Malaysia, but an
undescribed species is known from northern Australia (Lawrence and Brit-
ton 1991, 1994) and there is a Miocene record of a larva in amber from
the Dominican Republic (Poinar 1992).
851
CEROPHYTIDAE Latreille, 1834: 133
Cerophytum (Holarctic and Neotropical).
*The genus Cerophytum is usually considered to be the sister group of
those taxa comprising the elateroid complex (Eucnemidae, Throscidae and
Elateridae below) because of the well-developed propleurocoxal articula-
tion, which is reduced in the other three f~ilies; the same character
prompted Hlavac (1975) to place the family along with Artematopodidae
and Brachypsectridae in a redefined Artematopodoidea. The clicking
mechanism of cerophytids has been said to differ from those of other ela-
teroids in that it involves the hind legs (Crowson 1951), but a detailed
description of the mechanism by Buysson (1910) showed that it is of the
same type as that described for Elateridae (Evans 1972). The questionable
Cerophytum larva described and figured by van Emden (1932b) is actually
a dryopid. The true larva of Cerophytum elateroides Latreille was
described by Mamaev (1978) and Lawrence (1991) on the basis of the
same specimens collected in the Voronezh Region of Russia. Lawrence
noted that the spiracles described by Mamaev were actually paired glands,
the real spiracles being biforous with a closing apparatus. The genus Anis-
chia, often included here, is tentatively placed in Elateridae. Golbach
(1983), Soares and Peracchi (1964).
EUCNEMIDAE Eschscholtz, 1829: 10 (=Melasidae; incl. Perothopidae, Phyl-
loceridae)
*Muona (1993b) suggested that the taxa included in this and the next
two families represent two distinct lineages as follows : (Phyllocerinae +
Eucnemidae s. str.) and (Perothopinae + Throscidae + Lissominae + Thy-
lacosteminae + Elateridae). If such a scheme is accepted, then Eucnemidae
as delimited here would not be monophyletic. The subfamily classification
used here is according to Muona (1993b). Burakowski (1989), Cobos
(1965), Fleutiaux (1920, 1921, 1947a), Mamaev (1976b), Muona (1987,
1991a, 1991b, 1993a).
**Melasidae Fleming (1821: 49) has priority for the name of the fam-
ily, but Muona (1993b) has applied for conservation of Eucnemidae over
Melasidae (Case 2938).
PEROTHOPINAE Lacordaire, 1857: 128
Perothops (North America).
PHYLLOCERINAE Reitter, 1905: 2
Anelastes (North America, Asia) and Phyllocerus (Europe and Africa).
*Ghilarov (1979).
PSEUDOMENINAE Muona, 1993b: 41 (1991a: 167, nomen nudum)
Pseudomenes (=Eumenes) (Australia) and Schizophyllus (North America).
PALAEOXENINAE Muona, 1993b: 42 (1991a: 168, nomen nudum)
Palaeoxenus (North America).
852
PHLEGONINAE Muona, 1993b: 42 (1991a: 168, nomen nudum)
Phlegon (South America).
MELASINAE Fleming, 1821: 49
Arrhipis, Compsocnemis, Hylis, Hylochares, Isorhipis, Melasis, Micro-
rhagus, Xylophilus, etc.
EUCNEMINAE Eschscholtz, 1829: 10
Arisus, Dendrocharis, Dyscharachthis, Entomosatopus, Eucnemis, Galb-
ites, Phaenocerus, etc.
MACRAULACINAE Fleutiaux, 1923: 304
Anelastidius, Dromaeolus, Echthrogaster, Euryptychus, Fornax, Hemio-
psida, Macraulacus, Orodotes, etc.
THROSCIDAE Laporte, 1840a: 228 (=Trixagidae)
Aulonothroscus (widespread), Pactopus (western North America); Pater-
gus (Indo-Australian region), and Trixagus (=Throscus) (widespread).
*This group excludes Lissominae and Thylacosterninae, which are
tentatively placed in Elateridae. Kistner and Abdel-Galil (1986) included
the genus Neocrowsonia in this family, but there is no clear-cut evidence
from their paper that that unusual termitophile belongs here. Although the
authors placed it in Throscinae, it is likely that they intended to place it in
Lissominae, based on the tarsal lamellae. The latter group is included
among the Elateridae below. Burakowski (1975b), Coffin (1993), Cobos
(1961, 1967), Muona (1993a).
**We follow priority in the use of Throscidae (and -inae) over Trixa-
gidae Gistel ( 1856: 361 ), since the latter name has not been accepted wide-
ly enough to justify its maintenance over Throscidae under Art. 40b.
ELATERIDAE Leach, 1815: 85 (incl. Cebrionidae, Dicronychidae, Lissom-
idae)
*The classification of Elateridae is currently in a state of flux, and the
subfamilies listed are variously combined by different authors. The treat-
ment of the major elaterid subfamilies below follows Calder et al. (1993).
Burakowski (1973, 1975b), Costa and Casari-Chen (1993), Crowson
(1961a), Dolin (1975, 1978a, 1978b), Emden (1932b), Fleutiaux (1947b),
Gur'yeva (1969, 1974), Hyslop (1917, 1923), Laurent (1961, 1966), Naka-
ne and Kishii (1956), Ohira (1962), Stibick (1979), Zacharuk (1962).
**Cebrionidae Latreille (1802: 97) has priority for the family name, but
we maintain Elateridae here pending resolution of an application to the
ICZN being prepared by P.J. Johnson (in litteris) proposing precedence of
Elateridae (and hence Elateroidea) over Cebrionidae (-oidea).
CEBRIONINAE Latreille, 1802: 97 (incl. Aplastinae, Cebriognathinae, Plas-
tocerinae of LeConte not Crowson)
853
Aplastus (western North America), Cebrio (Europe, North Africa, Central
Asia), Cebriognathus (North Africa), Cebriorhipis (Asia), Euthysanius
(western North America), Octinodes (=Plastocerus of LeConte, not
Schaum) (North and South America), Paulusiella (Iran), Scaptolenus
(Nearctic, Neotropical), Selonodon (North America), Stenocebrio
(Chile), etc.
*This group is usually considered to be a distinct family. Some of the
North American genera included were once placed in a family Plastocer-
idae LeConte (1861: 172); this group was based on a misidentified type
genus (Plastocerus of LeConte, not Schaum) and is not the same as Plasto-
ceridae Crowson listed below. The genus Paulusiella, placed in Ka-
rumiidae by Paulus (1972a) and Mandl (1974), probably belongs here. It
is possible that the group is not monophyletic, since soft-bodied, loosely-
built males with pectinate antennae may have evolved more than once.
Hyslop (1923), Paulus (1981), Solervicens (1988).
TETRALOBINAE Laporte, 1840a: 230
Neotetralobus (Africa), Piezophyllus (Madagascar), Pseudalaus (Africa),
Pseudotetralobus (Australia, New Guinea), and Tetralobus (Africa,
southeast Asia).
*Costa et al. (1992), Laurent (1967).
THYLACOSTERNINAE Fleutiaux, 1920: 94 (=Balginae, Soleniscinae)
Balgus, Cussolenis (=Soleniscus), Pterotarsus (=Lissothyreus), Thylaco-
sternus, etc.
*Emden (1932b), Fleutiaux (1920, 1926), Gardner (1936).
LISSOMINAE Laporte, 1835: 178 (=Drapetinae; incl. Oestodinae, Prot-
elaterinae)
Anaspasis (Chile). Austrelater (Australia), Drapetes (widespread), Hypo-
chaetes (South America), Lissomus (widespread), Oestodes (North
America), Paradrapetes (South America), Protelater (New Zealand),
and Sphaenelater (New Zealand).
*Some members of this group (e.g., Drapetes and Lissomus) are often
placed in the family Throscidae. Burakowski (1973, 1975b), Calder et al.
(1993), Cobos (1967).
SEMIOTINAE Jacobson, 1913: 736
Oistus, Semiotinus, and Semiotus.
*Costa (1972), Golbach (1970).
PITYOBIINAE Hyslop, 1917: 262
Metablax (New Zealand), Parablax (Australia), Pityobius (North Amer-
ica), Tibionema (Chile), etc.
*Calder (1976, 1992), Ulrich (1987b).
OXYNOPTERINAE Candeze, 1857: 355
Campsosternus, Oxynopterus, Pectocera, etc.

854
AGRYPNINAE Candeze, 1857: 17/Lacordaire, 1857: 138 (incl. Adelo-
cerinae, Cavicoxumidae, Campyloxeninae, Hemirhipinae, Pyrophor-
inae, etc.)
Adelocera (=Cavicoxum), Agrypnus, Alaus, Anthracalaus, Aphileus, Cam-
pyloxenus, Chalcolepidius, Conoderus, Hemirhipus, Lacon, Lanelater,
Octocryptus, Pachyderes, Paracalais, Platycrepidius, Pyrophorus,
Tetrigus, Thoramus, etc.
*Calder (1990a), Costa (1975, 1992), Golbach (1976), Hayek (1973,
1979). . .
**Pangauridae Gistel (1856: 366) (Pangaura=Lacon) has pnonty for
the subfamily name but has never been used since its proposal; P.J. John-
son (in litteris) is submitting an application to the ICZN to conserve Agry-
pninae over Pangaurinae. . porte (now Eudicronychus), not Dicronychus Eschscholtz in Brulle (now
DENTICOLLINAE Stein and Weise, 1877: 96 (1856) (=Athomae, Cam-
pylinae; incl. Crepidomeninae, Hypnoidinae, Hypolithinae, Melanact-
inae, etc.)
Athous, Crepidomenus, Ctenicera, Denticollis (=Campylus), Hemicrepi-
dius, Hypnoidus, Hypolithus, Limonius, Melanactes, Pleonomus, etc.
*Calder (1986), Glen (1950), Stibick (1976).
**Campylinae Gistel (1856: 366) has priority over Athoinae Candeze
(1859: 4) and most other names for this subfamily, but Campylus has Ion~
been synonymized under Denticollis and the subfamily name based on It
replaced by Denticollinae Stein and Weise, 1877 (1856) by Art. .40b
(Stibick 1979). At least Prosteminae Gistel (1856: 367) and Melanactmae
Candeze (1857: 182) also predate Athoinae.
NEGASTRIINAE Nakane and Kishii, 1956: 203
Negastrius, Oedostethus, Quasimus, etc.
*Stibick (1971).
DIMINAE Candeze, 1863: 237
Dima, etc.
ELATERINAE Leach, 1815: 85 (incl. Adrastinae, Melanotinae, etc.)
Adrastus, Agriotes, Ampedus, Anchastus, Dicrepidius, Elater, Glyph~nyx,
Ischiodontus, Megapenthes, Melanotus, Melanoxanthus, Physorhznus,
Sericus, etc.
*Hayek (1990).
CARDIOPHORINAE Candeze, 1859: 4
Cardiophorus, Cardiorhinus, Horistonotus, Hypodesus, Nyctor, Para-
cardiophorus, etc.
HEMIOPINAE Fleutiaux, 1941: 31 (=Hemiopsinae)
Hemiops, Parhemiops, etc. . .
**Hemiopinae is the original and correct spellmg for the subfamily
name.
855
PHYSODACTYLINAE Lacordaire, 1857: 237
Physodactylus, etc ..
*Fleutiaux ( 1892).
ELATERIDAE incertae sedis:
EUDICRONYCHINAE Girard, 1971 : 645 (=Dicronychinae)
Anisomerus, Eudicronychus, and Tarsalgus.
*Schwarz (1897) first named this group as an elaterid subfamily and
later (1907) as a distinct family of Elateroidea. Dolin (1975) considered
the genera Eudicronychus and Anisomerus to be "outside the system" (of
Elateridae), having a unique type of aedeagus and wing venation similar
to that in the tribe Dicrepidiini (Elaterinae).
**The name Eudicronychinae replaced Dicronychinae Schwarz (1897:
11), which was based on a misidentified type genus, Dicronychus of La-
a subgenus of Cardiophorus) .
ANISCHIINAE Fleutiaux, 1936: 292
Anischia.
. *The g~nus Anischia Fleutiaux (1896) was first included by its author
m Eucnemidae (along with Cerophytum) and later (1936) moved to Ela-
teridae and placed in a separate subfamily. It is usually included in the
family ~erophytidae (Sc~enkling 1928), but the only feature uniting the
genus with Cerophytum IS the reduction of the metacoxal plates.
SUBPROTELATERINAE Fleutiaux, 1920: 99
Subprotelater.
*Fleutiaux (1920) placed this group in the family Eucnemidae, but it
has been excluded from that family and also from Throscidae based on its
well-~cler?tized, exposed labrum combined with the freely movable 5th
ventnte With exposed membrane at its base (Cobos 1961, Muona 1993b).
PLASTOCERIDAE Crowson, 1972b: 44 (not LeConte, 1861: 172)
Plastocerus (=Ceroplastus) (Asia Minor, southeast Asia) .
*This family was proposed by Crowson (1972b) for Plastocerus an-
gulosus (Germar) from Asia Minor and does not include those North
American species once included in Plastocerus but now placed in Octi-
~odes. The Southeast Asian species P. thoracicus Fleutiaux (1918) is also
mcluded here.
**Pl~stocerus as used here and by Crowson (1972b) refers to the origi-
nal restncted sense of Schaum (1852: 49); the genus is attributed toLe-
Conte's later and broader use in Neave (1939-40), Coleopterorum Cata-
logus and Genera Insectorum. Plastoceridae LeConte (1861: 172) is based
on LeCo~te' s generic concept, not Schaum's (see Elateridae: Cebrioninae).
Any family-group name based on Ceroplastus prior to Crowson (1972b)
would have priority for the family name, but we have found none.
856
DRILIDAE Blanchard, 1845b: 53
Drilus (Europe and Africa), Pseudeuanoma (eastern Europe, Asia Minor),
and Selasia (Africa).
*Most of the East Asian taxa once included in this family are now
placed in Lampyridae. Wittmer (1944).
OMALISIDAE Lacordaire, 1857: 303 (=Omalysidae, Homalisidae)
Omalisus (=Omalysus, Homalisus) and Thilmanus (southern Europe).
*Crowson (1972b) tentatively transferred Thilmanus to Lycidae and
Pseudeuanoma to Drilidae, but the former was returned to Omalisidae by
Bocak and Bocakova (1990). The Oligocene family Berendtimiridae
(Winkler 1987) is said to have the most features in common with Omali-
sidae. Burakowski (1988).
**The type genus, Omalisus Geoffroy, 1762, has been conserved by the
ICZN (1994a).
LYCIDAE Laporte, 1836: 25
*Thilmanus is now placed in Omalisidae (Bocak and Bocakova 1990).
Pristolycus was placed in a tribe Pristolycini of the family Lycidae (Win-
kler 1952), but has since been transferred to Lampyridae (McDermott
1964, 1966; Crowson 1972b). Crowson (1972b) also transferred Platero-
drilus from Drilidae to this family. One unanswered question is whether
those Asian taxa with distinctive "trilobite" larvae and completely larvi-
form females (lacking ovipositor and free tarsomeres, and with fewer than
four antennomeres), as described by Gravely (1915), Mjoberg (1925) and
Rosenberg (1943), form a monophyletic group, possibly the sister group
of the remaining lycids, as implied by Mjoberg (1925) and Crowson
(1972b). Adult males of two lycid species have been found copulating
with larviform females of the trilobite type: Lypropaeus biguttatus West-
wood and Duliticola paradoxa Mjbberg. Crowson (1972b) suggested,
based on adult similarities, that the genus Platerodrilus might also belong
to this group. Duliticola was not mentioned in the most recent suprage-
neric revision of this group (Bocak and Bocakova 1990); these authors
included Lypropaeus in the subfamily Leptolycinae, along with members
of the Leptolycini and Dexorini. Nakane (1969).
LYCINAE Laporte, 1836: 25
Caenia, Calopteron, Celetes, Dilophotes, Lycostomus, Lycus, Macrolycus,
Thonalmus, etc.
LEPTOLYCINAE Leng and Mutchler, 1922: 430 (incl. Duliticolinae?)
Dexoris, Leptolycus, Lypropaeus, etc.
*Includes at least some large "trilobite" larvae from Southeast Asia.
**Duliticolinae Mjoberg (1925: 140), based on one genus with a "trilo-
bite" larva (Duliticola), may belong here.
857
ATELIINAE Kleine, 1928: 222
Atelius and Scarellus.
METRIORRHYNCHINAE Kleine, 1926: 97
Cautires, Cladophorus, Conderis, Dilolycus, Flabellotrichalus, Hemi-
conderis, Metriorrhynchus, Trichalus, Xylobanellus, Xylobanus, etc.
**According to Neave (1939-40), the original spelling of the type ge-
nus is Metriorhynchus Guerin-Meneville, 1838, with Metriorrhynchus
Gemminger and Harold, 1869, as an unjustified emendation; the original
name (dated 1838 by Neave, 1830 or 1833 elsewhere) may be preoccupied
by the same name of Meyer, 1830. Further study is needed.
EROTINAE LeConte, 1881: 23
Dictyoptera, Dihammatus, Ditoneces, Eros, Libnetis, Lyponia, Plateros,
Platycis, Stadenus, Taphes, etc.
CALOCHROMINAE Lacordaire, 1857: 301
Calochromus, Dumbrelia, Lucaina, Lygistopterus, etc.
TELEGEUSIDAE Leng, 1920: 152
Pseudotelegeusis (Trinidad and Ecuador) and Telegeusis (southern Ari-
zona, northern Mexico and Panama).
*A group of doubtful affinities known only from males. Barber (1913b,
1952) considered Telegeusis to be close to Atractocerus in the Lymexyl-
idae, based mainly on the structure of the aedeagus; however, Crowson
( 1972b) placed it near Phengodidae and hypothesized that Barber's ( 1908)
luminous larva from Guatemala could be an adult female or large female
larva belonging to this genus. Allen and Hutton (1970), Wittmer (1976b).
PHENGODIDAE LeConte, 1861: 185
*As here constituted Phengodidae contains two very distinct groups,
Rhagophthalminae, which are restricted to Asia, and Phengodinae, which
occur only in the New World.
RHAGOPHTHALMINAE Olivier, 1907: 63
Cydistus, Dioptoma, Diplocladon, Dodecatoma, Falsophrixothrix, Mimo-
chotrya, Ochotrya, Rhagophthalmus.
*Olivier (1907) proposed the family Rhagophthalmidae for the Asian
genera Rhagophthalmus, Dioptoma and Ochotrya, and Pic (1937) added
Mimochotrya from Java. Gorham (1883) noted the resemblance of the
luminous males of Diplocladon and Dodecatoma to Dioptoma but did not
remove them from Drilidae, where they were also placed in Wittmer's
(1944) catalogue. McDermott (1964, 1966) considered the rhagophthal-
mids to be a subfamily of Lampyridae. Crowson (1972b) transferred Rha-
gophthalmus and Dioptoma to the family Phengodidae, along with Diplo-
cladon, Cydistus and Falsophrixothrix; his inclusion ofthese taxa in Phen-
godidae was based in large part on the occurrence of phengodid-like larvae
858
and larviform females in Rhagophthalmus, Dioptoma and Diplocladon. Raj
(1957) reported on the larva of an Indian Rhagophthalmus, which is simi-
lar to those of Phengodinae in lacking a basal mandibular brush and hav-
ing a well-developed, terminal lOth segment, apparently without holdfast
organs. Green (1913) described the larviform female of a Sri Lankan Dio-
ptoma as having two types of photogenic organs, a series of lateral spots
on the abdomen plus a larger ventral organ on the penultimate segment.
Neither author described the female legs or antennae, but females of In-
dian Dioptoma in the National Museum of Natural History, Washington,
D.C., are wingless and grub-like but with adult antennae and legs, like
those of Drilidae and some Lampyridae. Wittmer and Ohba (1994) con-
firmed the existence of a similar type of female in a Japanese species of
Rhagophthalmus. Diplocladon does have true larviform females with larval
antennae and legs; the larvae and adult females, which are called "star
worms", have a number of spot-like photogenic organs like those of typi-
cal phengodids (Harvey 1952). The adult males resemble those of ~ther
rhagophthalmines in having 12-segmented antennae and large, postenorly
emarginate eyes, but the emarginations are much shallower. The larvae
and females also have a series of large, disk-like organs, which do not
appear to correspond to light organs, on the head and abdominal sternites
2-9. Although the posterior eye emargination cannot be seen in Westw-
ood's (1843) figure, Dodecatoma bicolor has a similar head with 12-se-
gmented antennae and appears to belong to this group. Cydistus was
placed in the family Karumiidae by Paulus (1972a).
**Rhagophthalminae was credited to "Olivier, 1902" in McDermott
(1966) but we can find no evidence of the publication that McDermott
referred to.
PHENGODINAE LeConte, 1861: 185 (incl. Mastinocerinae, Pseudo-
phengodidae)
Cenophengus, Distremocephalus, Euryopa, Mastinocerus, Neophengus,
Oxymastinocerus, Penicillophorus, Phengodes, Phrixothrix, Pseudo-
phengodes, Ptorthodius, Zarhipis, etc.
*This group is restricted to the New World with the greatest diversity
in tropical America. Pic's (1930: 313) Pseudophengodidae was proposed
for Pseudophengodes because of the presence of photogenic organs in the
adult. The larva provisionally namedAstraptor illuminator Murray (1868)
probably belongs to one of the many genera of Mastinocerini (Barber
1908). Paulus (1975), Tiemann (1967), Wittmer (1963, 1976a).
**Phengodinae has priority over Mastinocerinae LeConte (1881: 40).
859
LAMPYRIDAE Latreille, 1817: 236
*This family was expanded by Crowson (1972b) to include several taxa
formerly considered to be Drilidae. Larvae are known only for the larger
subfamilies. McDermott (1964, 1966), Wittmer (1944).
**"Leach, 1815" was cited erroneously by Arnett (1963) for the family
name.
PTEROTINAE LeConte, 1861: 185
Pterotus (California).
*The larva of Pterotus obscuripennis LeConte has both the characteris-
tic anal holdfast (or grooming) organ consisting of a number of asperate
tubes and light organs on abdominal segments 7 and 8. The female is lar-
viform with at least 6 antennomeres and 4 tarsomeres and reduced oviposi-
tor (Dean 1979).
OTOTRETADRILINAE Crowson, 1972b: 55
Ototretadrilus (Asia).
CYPHONOCERINAE Crowson, 1972b: 55
Cyphonocerus (Asia) and Pollaclasis (North America).
OTOTRETINAE McDermott, 1964: 47
Brachylampis (California), Harmatelia, Lamellipalpus, Ototreta, Steno-
cladius, etc.
AMYDETINAE Olivier, 1907: 48
Amydetes, Cladodes, Ethra, Psilocladus, Vesta, etc.
LAMPYRINAE Latreille, 1817: 236
Cratomorphus, Dadophora, Diaphanes, Lamprigera, Lamprocera, Lam-
pyris, Lucidota, Lucio, Microphotus, Nyctophila, Phosphaenus, Pho-
tinus, Pleotomus, Pristolycus, Pyractomena, Pyractonema, etc.
*Pristolycus, originally described as a lycid and later placed in a lycid
tribe Pristolycini Winkler (1952), has been placed in Lampyridae by
Crowson (1972b) and Bocak and Bocakova (1990) and in Lampyrinae by
McDermott (1964).
LUCIOLINAE Lacordaire, 1857: 333
Bourgeoisia, Colphotia, Curtos, Luciola, Pyrophanes, Pteroptyx, etc.
PHOTURINAE Lacordaire, 1857: 338
Bicellonycha, Photuris, Pyrogaster, etc.
OMETHIDAE LeConte, 1861: 187
*This family was established by Crowson (1972b) for various taxa
previously placed in Cantharidae and Lampyridae.
DRILONIINAE Crowson, 1972b: 58
Drilonius (Asia).
MATHETEINAE LeConte, 1881: 29
Ginglymocladus and Matheteus (California).
860
OMETHINAE LeConte, 1861: 187
Blatchleya (eastern North America), Omethes (=Elianus) (North America
and Japan), Symphyomethes and Troglomethes (western North Amer-
ica).
*Fender (1975), Wittmer (1970).
CANTHARIDAE Imhoff, 1856: 69 (1815) (=Telephoridae; incl. Chaulio-
gnathidae)
*A new subfamily classification was proposed by Brancucci (1980).
Delkeskamp (1977-78), Verhoeff (1917, 1923b).
**Watt (1975) pointed out that Cantharidae of Latreille (1802: 185)
and most other authors of early to mid-Nineteenth Century was based on
a misidentified type genus (now Lytta, in Meloidae). Cantharidae appar-
ently was first used correctly for the present family by Imhoff (1856: 69).
This case should be referred to the ICZN for a decision (Art. 41). In the
meantime, we date Cantharidae from its first correct use (Imhoff 1856).
Telephoridae Leach (1815: 85) then has priority for the family and subor-
dinate family-group names, but retention of Cantharidae can be justified
by Art. 40b since Telephorus has long been treated as a junior synonym
of Cantharis.
CANTHARINAE Imhoff, 1856: 69 (1815)
Cantharis (=Telephorus), Podabrus, Rhagonycha, etc.
SILINAE Mulsant, 1862: 342
Discodon, Polemius, Silis, Sphaerarthrum, ?Tytthonyx, etc.
DYSMORPHOCERINAE Brancucci, 1980: 292
Asilis, Dysmorphocerus, Heteromastix, Oontelus, etc.
MALTHININAE Kiesenwetter, 1852: 239
Frostia, Malchinus, Malthinus, Malthodes, etc.
CHAULIOGNATHINAE LeConte, 1861: 186
Belotus, Chauliognathus, Ichthyurus, Trypherus, etc.
*Miskimen (1961) elevated this group to family rank, but this was dis-
puted by Magis and Wittmer (1974) and others.
ELATERIFORMIA Incertae Sedis
PODABROCEPHALIDAE Pic, 1930: 314
Podabrocephalus (southern India).
*Pic (1930) based this family on the genus Podabrocephalus (Pic
1913), a curious, soft-bodied beetle with a long head, strong genal ridges,
sickle-shaped, ventrally curved mandibles and conical, projecting pro-
coxae. The genus is likely to belong to one of the elateriform families, but
a more detailed study is required.
861
RHINORHIPIDAE Lawrence, 1988b: 3
Rhinorhipus (eastern Australia).
*This monotypic family is known from adults only, and its phyloge-
netic relationships are in doubt. In cladograms produced by Lawrence
(1988b), the taxon was basal to Elateroidea (as delimited above), Pse-
phenoidea (families Psephenidae through Eulichadidae above), or a clade
combining these two groups. In cladograms produced by Lawrence,
Nikitsky and Kirejtshuk (1995) the taxon usually formed part of a clade
containing Dascilloidea and Buprestidae. It is hoped that the discovery of
the immature stages may shed some light on its phylogenetic position.
Series BOSTRICHIFORMIA Forbes, 1926: 95
*This series was proposed by Crowson (1955, 1960) for a small group of fami-
lies lacking the special features of Haplogastra or Dascilliformia, and sharing
some but not all of the synapomorphies of Cucujiformia with which he earlier
placed them (Crowson 1950). These taxa were usually associated with Teredilia
(Bostrichidae, Anobiidae ), Brachymera (Dermestidae, Nosodendridae) or Clavi-
cornia (Derodontidae, Jacobsoniidae) in earlier classifications. Forbes (1926)
had used the same term for a group characterized by a primitive wing type and
including most of these taxa plus a number of the more primitive groups of
Elateriformia (as defined above), such as Scirtidae, Dascillidae and Artemato-
podidae. The series is likely to be paraphyletic, since there are no clear-cut
synapomorphies shared by all of its members, and Jacobsoniidae are tentatively
included here only because they lack at least two important synapomorphies of
Cucujiformia (open rhabdom [Caveney 1986] and loss of functional spiracles
on abdominal segment 8).
Superfamily DERODONTOIDEA LeConte, 1861: 100
DERODONTIDAE LeConte, 1861: 100 (incl. Laricobiidae, Peltasticidae)
*Although this family was included in Dermestoidea by Crowson
(1955), it has become evident in recent years that derodontids occupy a
unique place among the most primitive groups of Polyphaga. Fukuda
(1963), Lawrence (1985), Lawrence and Hlavac (1979), Nikitsky (1987).
PELTASTICINAE LeConte, 1861: 88
Peltastica (eastern Asia and western North America).
DERODONTINAE LeConte, 1861: 100
Derodontus (North America and Eurasia).
LARICOBIINAE Mulsant and Rey, 1863-64: 128
Laricobius (Eurasia and North America) and Nothoderodontus (New Zea-
land, Australia and Chile).
862
Superfamily BOSTRICHOIDEA Latreille, 1802: 202
(incl. Dermestoidea)
*This group combines the Dermestoidea and Bostrichoidea of Crowson (1959,
1961 b), with the exclusion of Derodontidae. Nosodendridae is doubtfully in-
cluded, since it has free Malpighian tubules, lacking the special type of crypto-
nephridism characterizing the remaining bostrichoids. Ivie (1985).
NOSODENDRIDAE Erichson, 1846: 465
Nosodendron (all continents, East Indies, Melanesia, New Zealand).
*Nosodendrids were considered to be Byrrhoidea in older classifica-
tions, and they remain in this position in a few relatively recent works
(Britton 1970, Endrody-Younga 1989). Boving and Craighead (1931)
placed the family in Dascilloidea, along with Scirtidae, Dascillidae and
Heteroceridae. Like Derodontidae, its connection with the bostrichoid
complex is tenuous. Ivie (1985) considered Orphilus to belong to this fam-
ily, but we have retained it in Dermestidae. Costa et al. (1986), Reichardt
(1976c).
DERMESTIDAE Latreille, 1804: 146 (incl. Thorictidae, Thylodriidae)
*Beal (1961), Hinton (1945), Mroczkowski (1968), Zhantiev (1976).
DERMESTINAE Latreille, 1804: 146
Dermestes (cosmopolitan) and Montandonia (eastern Europe).
*Dermestinae, Marioutinae and Thorictinae share at least one derived
feature which is apparently unknown in the other dermestid subfamilies.
In all three groups, there is a secondary ball and socket joint formed by a
mesal impression near the apex of the mesocoxa and a small lateral projec-
tion near the apex of the metasternal intercoxal process. Crowson (1992)
pointed out a similar type of joint in some groups of Chrysomelidae (Eu-
molpinae and related groups). Although marioutine larvae are unknown,
those of dermestines and thorictines are the only dermestid larvae with
urogomphi, and, if the median ocellus of adults is considered to be a
groundplan feature of the family, then its absence in these three groups
may also be synapomorphous.
MARIOUTINAE Jacobson, 1913: 832 (incl. Rhopalosilphinae)
Mariouta (Central Asia and North Africa) and Rhopalosilpha (Iran).
*Arrow (1929a), Zhantiev (1976).
THORICTINAE Agassiz, 1846a: 162 (incl. Thaumaphrastinae)
Thorictodes (=Thaumaphrastus) (widespread) and Thorictus (southern
Europe and Asia Minor to South Africa).
*Anderson (1949), Emden (1924a, 1951), John (1965), Reichensperger
(1926).
ORPHILINAE LeConte, 1861: 109
Orphilus (North America and Mediterranean).
863
.*Ivie (1985) placed Orphilus in Nosodendridae, but the larvae de-
scnbed by ~aulian (1943) and Beal (1985) have a number of features in
common With those of other Dermestidae.
TRINODINAE Casey, 1900: 163
Apsectus (North and Central America), Evorinea (Asia and the Pacific)
T:ichelodes .(Australia and New Zealand), Trinodes (Palearctic, Ethio~
pian and Onental), etc.
*Peacock (1978).
THYLOD~IIN~E Semenov-Tian-Shanskij, 1913: 498 (=Thylodriadinae)
Thylodrzas (widespread).
~Franciscolo (1975) placed this genus in its own family and super-
family.
ATTAGENIN~E Laporte, 1840b: 35 (incl. Egidyellinae?)
Attagenus (widespread), Novelsis (North America), and probably Egidyella
(southeastern Europe, Central Asia).
*Mroczkowski ( 1968) included Egidyella in the subfamily Egidyellinae
Semeno:-Tian-Shanskij (1915: 15), but Zhantiev (1976) placed the genus
here. Reitter (1899).
MEGATOMINAE Leach, 1815: 94 (incl. Anthreninae)
Anthrenocerus, Anthrenus, Megatoma, Neoanthrenus, Orphinus, Thau-
maglossa, Trogoderma, etc.
**Megatominae has priority over Anthreninae Gistel (1856: 363).
ENDECATOMIDAE LeConte, 1861: 207
Endecatomus (North America, Japan and Europe).
*The genus Endecatomus was included in the family Ciidae by early
workers, but was .moved to Bostrichidae by LeConte (1861). Crowson
(1961 b) first descnb~d the larva and proposed a sister group relationship
betv.:een Ende~atomidae and the remaining bostrichoids as one of two
possible evolutiOnary scenarios. Kompantzev (1978), Lawrence (1991).
BOSTR~CHIDAE L~treille, 1.802: .202. (=Bostrychidae; incl. Lyctidae, Psoidae)
. ~he su~famlly classificatiOn Is that of Ivie (1985), except that Anobi-
I~aeIS considered a distinct family. Anderson (1939), Crowson (1961 b),
Fisher (1950), Lesne (1896-1909, 1911).
DYSIDINAE Lesne, 1921: 286 (=Apoleoninae)
Apoleon (Asia) and Dysides (South America).
POLYCAONINAE Lesne, 1896: 111
Melalgus. (mainly Neotropical and Asia) and Polycaon (North and South
Amenca).
BOSTRICHINAE Latreille, 1802: 202
Apate, Bo.st:ich~s, Bostrychopsis, Dinapate, Enneadesmus, Lichenophanes,
Scrobtcza, Smoxylon, Xylopertha, Xylothrips, etc.
864
PSOINAE Blanchard, 1851: 434
Chilenius (Chile), Psoa (Europe and North America), Stenomera (Asia
Minor), etc.
DINODERINAE C.G. Thomson, 1863: 201
Dinoderus, Rhyzopertha, etc.
LYCTINAE Billberg, 1820: 48
Lyctoxylon, Lyctus, Minthea, Tristaria, Trogoxylon, etc. . . . .
*Although this group is usually considered to be a distmct family m
general texts, it is more closely related to Dinoderinae than to other mem-
bers of the family. Gerberg (1957).
EUDERIINAE Lesne, 1934: 392
Euderia (New Zealand).
*Lesne (1934) considered this genus and not Endecatomus to be the
most primitive living bostrichid.
ANOBIIDAE Fleming, 1821: 50 (incl. Ectrephidae, Gnostidae, Ptinidae)
*According to !vie (1985), this group should be a subfamily ofBostri-
chidae. Boving (1954), Ford (1970), White (1971a, 1971b, 1974a, 1974b,
1982).
**Ptinidae Latreille (1802: 112) has priority over Anobiidae if the two
families are combined as done here.
EUCRADINAE LeConte, 1861: 202 (=Hedobiinae)
Clada, Eucrada, Hedobia, Ptilineurus, Ptinomorphus, etc.
*White (1974a) placed Eucrada in Dryophilinae. Espafiol (1970),
Viedma (1973, 1977).
**Eucradinae has priority over Hedobiinae Mulsant and Rey (1868 : 23)
for the name of this group if Eucrada is included here.
PTININAE Latreille, 1802: 112 (incl. Gibbiinae)
Casopus, Diplocotes, Ectrephes, Fabrasia, Gibbium, Gnostus, Gynopterus,
Maheoptinus, Mezium, Neoptinus, Niptus, Ptinus, Sphaericus, Trigona-
genius, Xylodes, etc.
*This group is often given family rank. Belles (1981, 1985, 1991a,
1991b), Belles and Lawrence (1990), Hall and Howe (1953), Hinton
(1941a), Lawrence and Reichardt (1969).
DRYOPHILINAE LeConte, 1861: 205
Dryophilodes, Dryophilus, Grynobius, etc.
ERNOBIINAE Pic, 1912: 12 (incl. Cerocosminae)
Cerocosmus, Ernobius, Ochina, Ozognathus, Xestobium, etc.
*Crowson (1967a), White (1974a).
**Cerocosminae Pic (1912: 45) (=Cosmocerinae Solier, 1849: 476,
based on a preoccupied generic name) was placed here by White (1974a).
865
ANOBIINAE Fleming, 1821: 50
Anobium, Gastrallus, Hadrobregmus, Macranobium, Stegobium, Xyleto-
bius, etc.
PTILININAE Shuckard, 1840: 45
Ptilinus, etc.
ALVARENGANIELLINAE Viana and Martinez, 1971: 121
Alvarenganiella (Brazil).
XYLETININAE Gistel, 1856: 368
Deroptilinus, Lasioderma, Leanobium, Pronus, Vrilletta, Xyletinus, etc.
DORCATOMINAE C.G. Thomson, 1859: 90
Caeno.cara, Calymmaderus, Dicoelocephalus, Dorcatoma, Eutylistus, Pe-
talzum, Protheca, Stagetus, etc.
*Espafiol (1977a, 1977b) continued to recognize this subfamily in a
broader sense, including the following group as one of eight sections (Sec-
cion Tricorynus).
**The origina! spelling of the type genus is Dorkatoma, but Silfverberg
(1994b) has applied to the ICZN for conservation of the spelling Dorca-
toma.
MESOCOELOPODINAE Mulsant and Rey, 1864: 311 (=Tricoryninae)
Cryptorama, Deltocryptus, Mesocoelopus, Rhamna, Secretipes, Tricorynus,
etc.
*White (1971b) hypothesized that the highly derived, compacted "dor-
catomines" represented two independent lineages, the present group being
more closely related to Xyletininae and the true dorcatomines to Anobi-
inae.
**Mesocoelopodinae, as well as Mesothinae Portevin (1931: 489), has
priority over Tricoryninae White (1971b: 1301) for the name of this sub-
family (M.A. Alonso-Zarazaga in litteris).
BOSTRICHIFORMIA Incertae Sedis
JACOBSONIIDAE Heller, 1926: 127 (=Sarothriidae; incl. Derolathrinae)
Derolathr~s (tropical and subtropical regions including Hawaii and many
other Islands), Saphophagus (New Zealand), and Sarothrias (=la-
cobsonium) (Indo-Australian to Fiji).
*A small family of uncertain position. Although Crowson (1959, 1960)
placed the group in Dermestoidea (along with Derodontidae Nosodendr-
idae ~nd Dermestidae), he .noted similarities to Staphylinoidea in wing
venatiOn and the larval maxilla of Saphophagus. Its exclusion from Cucu-
jifor~ia is based. partly on the relatively complete abdominal apex with
functiOnal 8th spiracles. Sen Gupta (1979) included the Derolathrinae in
the cucujoid family Merophysiidae (see Endomychidae below). Lobi and
Burckhardt (1988b).

866
**Jacobsoniidae has priority over the replacement name Sarothriidae
Crowson (1955: 75) (which in our view has not become sufficiently estab-
lished to justify maintaining it over J acobsoniidae under Art. 40b) and
Derolathrinae Sen Gupta (1979: 692).
Series CUCUJIFORMIA Lameere, 1938: 358
*This group is used here in the sense of Crowson (1955) to include cleroids,
lymexyloids, cucujoids, tenebrionoids, chrysomeloids and curculio~oids. Lame-
ere (1938) used a variant term for the last four groups only, wh1le Crowson
(1950) initially included the present Bostrichiformia in his Cucujiformia. This
series is defined by having cryptonephridic Malpighian tubules of the normal
type (except where secondarily lost as in the chrysomelid genus Donacia), an
acone ommatidium with open rhabdom, no functional spiracles on abdominal
segment 8, metendostemite of the hylecoetoid type, metacoxae not excavate,
and aedeagus of the sheath, cucujoid, or pseudotrilobed type.
Superfamily LYMEXYLOIDEA Fleming, 1821: 49
*Crowson (1981) placed the Strepsiptera (as family Stylopidae) in this super-
family (see Order Coleoptera above).
LYMEXYLIDAE Fleming, 1821: 49 (=Lymexylonidae; incl. Atractoceridae)
*Barber (1913b, 1952) consideredAtractocerus to be related to Telege-
usis, and Arnett (1963) included both genera in Telegeusidae. The genera
of Lymexylidae were revised by Wheeler (1986), who excluded Hylecoe-
topsis (Lane 1955) (tentatively moved to Cleridae) and Heteromeroxylon
(Rhipiphoridae). Kurosawa (1985) split the genus Atractocerus into five
genera.
**Fischer (1821: 37) also established a family-group name "Lyme-
xyla", but in the absence of evidence of an exact publication date for
Fischer's work Fleming's name of July 1821 has priority; both have prior-
ity over Atractoceridae Laporte (1840a: 290).
HYLECOETINAE Gistel, 1856: 384
Elateroides (=Hylecoetus) (North America, Eurasia).
LYMEXYLINAE Fleming, 1821: 49
Atractocerus (pantropical), Lymexylon (Eurasia), etc.
MELITTOMMATINAE Wheeler, 1986: 160 (=Melittomminae)
Australymexylon (Australia), Melittomma (North America, Neotropical,
Old World tropics, Madagascar, Australia), Melittommopsis (South
America), and Promelittomma (Madagascar, Seychelles).
**The grammatically correct spelling of the subfamily name is Melit-
tommatinae, not Melittomminae as originally proposed.
867
Superfamily CLEROIDEA Latreille, 1802: 110
*Crowson ( 1955, 1964b) was the first one to clearly define this superfamily in
the modem sense, removing the Dermestidae, Passandridae, Bothrideridae and
Ciidae, which had been included by Boving and Craighead (1931). Typical
cleroids are predacious with a mandibular mola absent in all stages, and larvae
usually have basal mandibular processes collectively referred to as the lacinia
mobilis and often a pedunculate seta on the mala. However, basal, fungivorous
members of the superfamily share many features with primitive Cucujoidea.
PHLOIOPHILIDAE Kiesenwetter, 1863: 666 (=Phloeophilidae)
Phloiophilus (Europe).
*The genus Phloiophilus was included among the clavicoms or mala-
coderms in older classifications, and in the Junk Catalogue (Pic 1926) it
was placed in a distinct family "Phloeophilidae" (see below), along with
Acanthocnemus and Xerasia (see Acanthocnemidae and Byturidae below).
Crowson (1964b) limited the family to this one genus, which he consid-
ered to be a basal member of the Cleroidea. Crowson (1964c).
**The family name was originally and has been commonly misspelled
Phloeophilidae, based on Phloeophilus Agassiz, 1846, an unjustified emen-
dation of Phloiophilus Stephens, 1830. These names should not be con-
fused with the anthribid group name Phloeophilini Lacordaire ( 1866: 517),
based on Phloeophilus Schoenherr, 1833.
TROGOSSITIDAE Latreille, 1802: 110 (incl. Lophocateridae, Ostomidae,
Peltidae, Temnochilidae)
*As here constituted, this family exhibits considerable structural diver-
sity, and it has been split into three different families. Crowson (1955)
originally divided the group along traditional lines into two subfamilies:
o.ne incl~ding predacious species, adults of which are narrow and parallel-
Sided With closed procoxal cavities, and another including mainly fungi-
vorous species, which are broader with open procoxal cavities. In a later,
more detailed study of the group, Crowson (1964b, 1966b, 1970) found
the variation to be much more complex. In the first paper, he divided it
into two families based mainly on the presence of a distinct mola or laci-
nia mobilis, plus a hooked laciniar spine, in the adult (Peltidae), or their
~bsence (Trogossitidae). In this scheme, Peltidae included Egoliinae, tradi-
twna~ly placed among the trogossitines, and Trogossitidae included Lopho-
catennae, usually treated as peltines. Within the first family, Decamerinae
included the North American Eronyxa, and Peltinae included the Holarctic
Ostoma, Thymalus and Calitys, plus the New Zealand Protopeltis. Crow-
son later (1966b) proposed a new subfamily, Rentoniinae, for several small
clambid-like forms from the Southern Hemisphere, and in 1970 he revised
the family again, proposing a new peltine subfamily for Protopeltis, ele-
868
vating Lophocaterinae to family rank, and moving both Calit:Ys (as Calit-
inae) and Egoliinae to Trogossitidae. Because of the uncertamty of some
of these placements and new ipput on i~I?~~ur~ stages, some authors (Bar-
ron 1971, Lawrence and Britton 1991, Shpmski1992) have returned to. the
broader family concept which is followed here. Barron (1975), Reitter
(1876).
PROTOPELTINAE Crowson, 1966b: 120
Protopeltis (New Zealand).
LARINOTINAE Slipinski, 1992: 443 . .
Colydiopeltis (Thailand) and Larinotus and Parapel~ts (Ap~t~~ha~.
*The inclusion here of Colydiopeltis and Parapeltts by Shpmski (1992)
is somewhat weakly founded, being based mainly on the absence ?f ~ibial
spurs. Crowson's (1964b) putative larva .of Acant~ocne_~u~- m?ncans
(Hope) almost certainly belongs to Parapeltts australlcum Shpmski, based
on association with adults from both Mt. Coottha and Mt. Cook, Queens-
land. This larva has a number of features in common with that described
by Crowson (1964b) for Protopeltis.
PELTINAE Kirby, 1837: 104 (=Ostominae) .
Ostoma (Holarctic), Peltis (Palearctic), and Thymalus (Holarctlc).
**Peltinae of Latreille (1807a: 8; 1825) did not include Peltis as a vahd
genus and thus is unavailable (Art. 11f(i) 1). The name .was mad~ a~ailable
at least by Kirby (1837: 104) if not earlier, and so still has pnonty over
Ostominae Reitter (1882: 146). The type genus, Peltis Kugelann, 1792, has
been conserved in the present sense by the ICZN (1994a).
RENTONIINAE Crowson, 1966b: 120
Australiodes, Rentonellum, Rentonidium and Rentonium (New Zealand),
Parentonium (Australia, New Zealand), etc. . THANEROCLERINAE Chapin, 1924: 251
*Crowson (1966b, 1970) combined Australiodes (originally descnbed
as a leiodid by Endrody-Younga 1960) with several new ge~era from Aus-
tralia and New Zealand to form a new taxon related to Peltmae and Pro~o­
peltinae. Adults are characterized ?Y th~ir small siz~ .and unusual clambid-
like body. The group also occurs m Chile and additional genera and spe-
cies remain to be described.
DECAMERINAE Crowson, 1964b: 287
Antixoon (Central America), Decamerus and Diontolobus (South Amer-
ica).
LOPHOCATERINAE Crowson, 1964b: 297
Ancyrona (widespread), Eronyxa (western North America!, Grynocharis
(Holarctic), Grynoma (New Zealand), Lophocateres (widespre~d), Ly-
coptis (North America), Neaspis (Australia), Peltonyxa (Australia), etc.
*Barron (1971) retained this group within Peltinae. Tait et al. (1990)
transferred Eronyxa from Decamerinae to this group based on the newly
discovered larva of Eronyxa expansus Van Dyke. Barron (1975).
869
CALITINAE Reitter, 1922: 66
Calitys (North America, Europe, South Africa).
*Crowson (1970) moved this genus from Peltinae to this subfamily.
EGOLIINAE Lacordaire, 1854: 334
Acalanthis (Chile), Egolia . (Australia), Necrobiopsis (Australia) Par-
acalanthis (Australia), etc. '
, . ~~ro~son' s (1970) .larval information on this group was supported by
Shpmski (1992), who Illustrated an Egolia larva from Tasmania.
TROGOSSITINAE Latreille, 1802: 159 (=Temnochilinae Tenebroidinae
~~~~ ' '
Acrops (Asia),Airora (New World), Lepidopteryx (=Leperina) (Asia, New
Zealand, New Caledonia, Australia, Pacific), Nemosoma (New World,
Europ~, No~h Africa), Phanodesta (Chile), Temnoscheila (=Te-
mnochtla) (widespread), Tenebroides (=Trogosita, Trogossita) (wide-
spread), etc.
CHAETOSOMATIDAE Crowson, 1952: 67
Chaetosoma and Chaetosomodes (New Zealand), Malgassochaetus and
. Somatochaetus (Madagascar).
*Ekis and Menier (1980) and Menier and Ekis (1982) reviewed this
group and added taxa from Madagascar.
CLERIDAE Latreille, 1802: 110 (incl. Corynetidae, Korynetidae)
*Barr (1962), Boving and Champlain (1920), Chapin (1924 ), Corporaal
(1950), Crowson (1964b, 1972a), Ekis and Gupta (1971) Kolibac (1987
1989), Winkler (1982, 1989). ' '
lsoclerus (=Ababa) (No~th America, Mexico, Brazil, southeast Asia), Neo-
clerus (southeast Asia), Thaneroclerus (Asia, one widespread) Zeno-
dosus (North America), etc. '
*Kolibac (1992) reviewed the group, treating it as a separate family.
Corporaal (1939), Foster (1976).
TILLINAE Leach, 1815: 87
Cylidrus, Cymatodera, Monophylla, Tillus, etc.
HYDN?CERINAE Spinola, 1844: Tableau generique .. p. 3 (=Phylloba-
enmae)
lsohydnocera, Lemidia, Phyllobaenus (=Hydnocera), etc.
**Hydn?cerinae has priority for the name of this group. Furthermore,
Phyllobaenmae of Lacordaire (1857: 466) and all other authors until
Wolc~tt (1944: 121) is based on a misidentified type genus (Phyllobaenus
?f Spmola, 1844, not D.ejean, 1837; now Phlogistosternus of Epiphloe-
~na~). Use ~fPhyllobaenmae Wolcott over Hydnocerinae could perhaps be
JUstified usmg Art. 40b, but continued use of Hydnocerinae by Crowson
 871
870
. dubious and in any event any legiti- on the pronotal hypomera; the function of these organs is unknown. The
( 1955) and other authors makes thldiS . a' decision by the ICZN (Art. larva tentatively attributed to this species by Crowson (1970) is probably
h 11 0 b ninae wou reqmre d
mate use of P Y ae · 'ty here and use Hy nocer- that of Parapeltis australicum Slipinski (see above).
41). We feel it is much better to follow pnon
**A subfamily name "Acanthocneminae Herm., 1912-20", based on
inae. ·
'11 1802· 110 (incl. Priocennae, D'Ierops inae , Clero- the preoccupied generic name Acanthocnemis Blanchard (not Hawle and
CLERINAE Latrei e, · Corda) and therefore unavailable, has been used in Diptera-Asilidae (Han-
piestinae) . . E clerus Priocera, Sedlacekvia, Thana- dlirsch 1925).
Cleropiestus, Clerus, Dzeropszs, no '
simus, Trichode~, etc. . 7 and Cleropiestinae Winkler.c_t~80: PHYCOSECIDAE Crowson, 1952: 125
*Dieropsinae Wmkler (196~. ?\i989) Menier (1981) also cntlcized Phycosecis (Australia, New Zealand, New Caledonia, New Hebrides).
437) were placed here by Ko~Ib~c ae Th~ anomalous Australian gen~s *This group was placed by Crowson (1952) in Cucujoidea but later
Winkler's concept of. Cleropie~~ tder (1989) to be doubtfully placed m (1964b) transferred to Cleroidea. This was based in part on a larval speci-
Sedlacekvia was considered by m men, the identity of which has been confirmed by rearing.
Cleridae.
EPIPHLOEINAE Kuw.ert, 1893: 492 d Neichnea (North America), Phlogi- PRIONOCERIDAE Lacordaire, 1857: 411
Epiphloeus (Neotropical), Ichnea an Jdgia and Prionocerus (Asia and Africa), and Lobonyx (southern Palear-
stosternus (New Wor~d) •. etc. I h belongs here rather than in Eno- ctic).
*W. Barr (in litteris) mdicates c nea
*Champion (1919) suggested that this group be treated as a separate
pliinae. " .. , S . la (1841 : 75) has priority for the name family of Malacodermata (see Lawrence, Slipinski and Pakaluk 1995).
**The name IchnOides pmo h' me may never have been Majer (1987) claimed that prionocerids are more closely related to Cler-
of this group if Ichnea is included, but t IS na
idae than to Melyridae. Gardner (1929).
. . d and seems to have been forgotten.
1atlmze **A group name "Prionocerites" was used by Laporte (1840a: 285) but
ENOPLIINAE Gistel, 1856: 36~ d ) Chariessa (New World), did not include the genus Prionocerus, which he placed in "Telephorites"
Enoplium (Eurasia, North Afnca,. Ma Nagas~o~ld) Tenerus (Old World), (Laporte 1840a: 275); the group did include Priocera, indicating that the
E Asia Mmor, ew '
Orthopleura ( urope, Pl optera (Neotropical), etc. name "Prionocerites" was undoubtedly a lapsus calami for "Priocerites",
Phymatophaea (New Z~al~?d);, S a::la (1844: Tableau generique .. p. a name already established by Laporte (1836: 33) for the same group of
**The name "PlatynopterOldes p 1857) has priority for the name.of genera including Priocera (now Cleridae: Clerinae).
4) (latinized by Chenu and De~me~ef~rgotten since Lacordaire (1857) Ig-
this group, but seems to h~ve e MELYRIDAE Leach, 1815: 87 (incl. Dasytidae, Malachiidae, Rhadalidae, Car-
nored it in favor of Enopl~mae. 1860· 198 phuridae)
TARSOSTENINAE Jacquelin du Va~A t iia) Paratillus (Australia, intro. *Constantin (1990), Majer (1986, 1987, 1990).
Blackburniella and TcUarss:)te~::~ars:S~::us (cosmopolitan). MELYRINAE Leach, 1815: 87
Europe, western • . Cerallus (eastern Europe, Central Asia, Asia Minor), Chalchas, Astylus
4
KORYNETINAE Laporte, 1836. 3 b. ( osmopolitan), Lebasiella (New and Arthrobrachus (Neotropical), Melyris and Falsomelyris (Europe,
Korynetes (widespread), Necr~ ~~f ~ a) and Pylus (Australia). Asia Minor, Africa), Melyrodes (New World), etc.
World), Opetiopalpus (Eurasia, nc '
RHADALINAE LeConte, 1861: 194 (=Aplocneminae, Haplocneminae)
Anthriboclerus (Seychelles), Aplocnemus (Palearctic, Asia Minor, Africa,
ACANTHOCNEMIDAE Crowson, 1964bd: 3 ~d7 1 ) Philippines), Eucymbolus (Central America), Hemipleurus (Borneo,
alia introduce WI e Y · · d d
Acanthocnemus (Aus t ~ . ' e) is native to Australia but mtro uce Sarawak), Malthacodes (=Donaldia) (Africa, India, islands of Indian
*Acanthocnemus nzgrzcans (H?P f. d Europe (Champion 1922). Ocean), Microjulistus (Mediterranean, Africa, Central Asia), Peleco-
to New Caledonia and parts of A~, lA .r~ca : d was illustrated under the phora (Mauritius, Reunion), Rhadalus (New World), Semijulistus (=Eu-
The genus is often included in e yn ~e . Hope by Britton (1970). relymis (North America, Japan, Turkestan), Trichoceble (Eurasia), etc.
. Dasytes fusczpennzs • ..
name of a jumor synonym, . f partially enclosed cavities *Peacock (1987) reviewed this subfamily. Vinson (1957).
A unique feature of this group IS the presence o
872
GIETELLINAE Constantin and Menier, 1987: 62
Gietella (Canary Islands). . h
*According to Constantin and Menie~ (1987, 1990), this groups ares
features with both Rhadalinae and Dasytmae.
DASYTINAE Laporte, 1840a: 280 . .
Amauronia, Amecocerus, Danacaea, Dasytes, Dolzchosoma, Emcopus,
Trichochrous, etc.
MALACHIINAE Fleming, 1821: 50
Attalus, Axinotarsus, Carphurus, Collops, Ebaeus, Endeodes, He/co gaster,
Laius, Malachius, Troglops, etc.
*Evers (1968).
Superfamily CUCUJOIDEA Latreille, 1802: 210
(=Clavicornia) . .
*Pakaluk et al. (in press) summarized the supragener~c cl~ssi~catiOn and
family-group nomenclature for all families. Their ~lasslficatiOn ~s ~ollowe~
below except for the retention of the well-known family.na~es Sphmdid~e an
Latridlidae (instead of the older Aspidiphoridae ~d Co:ticamdae,..respecu:ely).
S'l· . , ki and Pakaluk (1992) reviewed the classificatiOn of the cerylomd se-
Ipms
ries" of families (Bothrideridae to Latn'd"d b 1 )
11 ae e ow ·
PROTOCUCUJIDAE Crowson, 1954: 60 (1952: 121, nomen nu~um)
Ericmodes (=Protocucujus) (Australia, southern Sou.th Amenca) ..
*Slipinski and Pakaluk (in press) revised the family and descnbed the
first larvae.
SPHINDIDAE Jacquelin du Val, 1860: 224 (incl. Aspidiphoridae)
*Sphindidae is generally considered to be the Sister group of Proto-
cucujidae (Sen Gupta and Crowson 1979!. A ~lad~ gram of the genera wa~
iven by McHugh (1993) and his classificatiOn Is followed here. Bura
~owski
and Slipinski (1987), McHugh (1990), McHugh and Wheeler
(19~!~he section of Jacquelin du Val's "1859-63" work in which Sphind-
idae is named has been dated as 1860 (Silfverberg 1992, and ~.D. P?pe
. l'tteris to J Pakaluk). The oldest name for the family and nommotypic~
~:b~amily is fuen Coniporidae C.G. Thomson (1859: ~0}, based onC~m­
porus (proposed for and an objective synonym of Aspz.dzphorus). Compo-
rus was soon sunk under Aspidiphorus and the family-group name re-
placed with Aspidiphoridae Kiesenwetter (1877: 198). The latter name can
be retained by Art. 40b, as Aspidiphoridae Kiesenwett~r, 1877 (1859),
which then has priority over Sphind.idae;.Pak~luk et al. (m press) accord-
ingly use Aspidiphoridae and Aspidiphonnae m place of the better-known
873
names Sphindidae and Sphindinae. The original spelling of Aspidiphorus
is Arpidiphorus; an application to the ICZN to conserve Aspidiphorus has
been submitted by J. McHugh.
PROTOSPHINDINAE Sen Gupta and Crowson, 1979: 181
Protosphindus (Chile).
ODONTOSPHINDINAE Sen Gupta and Crowson, 1979: 180
Odontosphindus (Holarctic ).
SPHINDIPHORINAE Sen Gupta and Crowson, 1979: 180
Sphindiphorus (southern Africa).
SPHINDINAE Jacquelin du Val, 1860: 224 (incl. Aspidiphorinae, Eury-
sphindinae)
Aspidiphorus (Eurasia, Africa, Australia), Carinisphindus (Central Amer-
ica, West Indies), Eurysphindus (North to South America), Genisphi-
ndus (Neotropical), Notosphindus (Australia), Sphindophorus (southern
Africa), and Sphindus (North America, Eurasia, Africa, Madagascar,
Neotropical).
BRACHYPTERIDAE Erichson, 1845: 125 (=Cateretidae, Kateretidae)
Amartus (California, Central Asia, Asia Minor), Brachyleptus (Central
Asia, eastern Europe, North Africa), Brachypterolus (Holarctic), Bra-
chypterus (North America, Eurasia, Africa), Heterhelus (Holarctic),
Jelinekia (Australia), Kateretes (Holarctic), Neobrachypterus (South
America), Notobrachypterus (Australia), etc.
*Although Verhoeff (1923a) separated Brachypteridae from Nitidulidae
on the basis of larval features, the former is usually considered to be a
subfamily of the latter. Kirejtshuk (1986b) made a case for elevating the
group to family rank. Audisio (1993).
**The oldest name cited for this group is "Cateretes" Erichson (1843:
227), but in that work Erichson merely appropriated the generic name
Cateretes (unjustified emendation of Kateretes) as a group name for the
genera Cercus and Brachypterus; he did not recognize a valid genus Cate-
retes in this group, so this group name is unavailable (Art. llf(i) 1). Later,
Erichson ( 1845: 125) replaced the name "Cateretes" with "Brachypterinae"
for the group including the same two genera. The name Kateretidae was
apparently first made available by Ganglbauer (1899: 447) (as Cateretini,
including Cateretes), but Brachypteridae Erichson has priority.
NITIDULIDAE Latreille, 1802: 132 (incl. Cybocephalidae)
*The subfamily classification is after Kirejtshuk (1986a and unpub-
lished). Audisio (1993), Boving and Rozen (1962), Gillogly (1965), Haya
shi (1978), Jelfnek (1975), Kirejtshuk (1982, 1986b, 1986c, 1990), Kirej -
tshuk and Lawrence (1992), Parsons (1943, 1972).
874
CALONECRINAE Kirejtshuk, 1982: 117
Calonecrus (southeast Asia).
CARPOPHILINAE Erichson, 1842: 148
Amphicrossus, Amystrops, Carpophilus, Epuraea, Mystrops, Platychoro-
psis, Taenioncus, Trimenus, Urophorus, etc.
MELIGETHINAE C.G. Thomson, 1859: 67
Meligethes, Metapria, Microporum, Pria, etc.
NITIDULINAE Latreille, 1802: 132
Aethina, Camptodes, Cychramptodes, Cychramus, Cyllodes, Lasiodactylus,
Lawrencerosus, Lobiopa, Nitidula, Omosita, Orvoenia, Pallodes, Poca-
dius, Psilopyga, Soronia, Thalycra, etc.
CILLAEINAE Kirejtshuk and Audisio in Kirejtshuk, 1986a: 219
Brachypeplus, Cillaeus, Colopterus, Conotelus, Nesopeplus, Platynema,
etc.
CRYPTARCHINAE C.G. Thomson, 1859: 69
Arhina, Cryptarcha, Eucalosphaera, Glischrochilus, Pityophagus, etc.
CYBOCEPHALINAE Jacquelin du Val, 1858: 151
Cybocephalus, Pycnocephalus, etc.
SMICRIPIDAE Hom, 1879: 325
Smicrips (North to South America). . .. .
*This group is sometimes considered to be a subfamily of Nitlduhdae.
MONOTOMIDAE Laporte, 1840b: 377 (incl. Rhizophagidae)
*The classification here follows Sen Gupta (1988). Lawrence (1991),
Pakaluk and Slipinski (1993).
**Monotomidae has priority over Rhizophagidae for the family name.
Chaudoir (1845: 209, sometimes cited erroneously as 1842) is usually
cited as author of the family name. Neave (1939-40) lists a "Monotoma
Panzer, 1792" (Colydiidae) as well as Monotoma Herbst, 1793 (present
sense); this needs further investigation.
RHIZOPHAGINAE Redtenbacher, 1845: 125
Rhizophagus (North America, Eurasia). . **Lawrence and Britton (1991) characterized the genus Myrabolia in
**The original spelling of the type genus is Ryzophagus, app~ied to the
family name by Rfha (1989), but Silfverberg (1994b) has ~pphe~ to the
ICZN for conservation of the spelling Rhizophagus and fixatiOn of Its type
species. . . .
MONOTOMINAE Laporte, 1840b: 377 (incl. Lenacmae, Thwmnae)
Bactridium (New World, New Guinea, Sumatra), Europs (Neotropical, Old
World tropics), Hesperobaenus (North America, Pacific), Lenax (New
Zealand), Mimemodes (Asia, New Guinea, Australia), Monotorr:a
Herbst (cosmopolitan), Monotomopsis (Asia, Australia), Shoguna (Asia,
Australia, Madagascar), Thione (Neotropical), etc.
875
BOGANIIDAE Sen Gupta and Crowson, 1966: 63
*Crowson (1990), EndrOdy-Younga and Crowson (1986), Sen Gupta
and Crowson (1966) .
PARACUCUJINAE EndrOdy-Younga and Crowson, 1986: 255
Metacucujus (South Africa) and Paracucujus (Australia).
*EndrOdy-Younga (1991).
BOGANIINAE Sen Gupta and Crowson, 1966: 63
Afroboganium (South Africa), Athertonium and Boganium (Australia).
*Crowson (1990), Lawrence et al. (1993)
HELOTIDAE Reitter, 1876: 5/Chapuis, 1876: 15
Helota (Asia and Africa).
*Fukuda (1943), Olliff (1883).
**"Gorham, 1874" has been cited for the family name; family status
was implied but no family-group name used by Gorham (1874).
PHLOEOSTICHIDAE Reitter, 1911: 48
*This group contains a diverse assemblage of primitive cucujoids, and
each of the six subfamilies may be elevated to family rank by Crowson
and Slipinski (manuscript in preparation). Crowson (1973a), Lawrence
(1988a), Lawrence and Britton (1991), Sen Gupta and Crowson (1969b).
PHLOEOSTICHINAE Reitter, 1911: 48
Phloeostichus (Europe).
*Weise (1897).
HYMAEINAE Sen Gupta and Crowson, 1966: 77
Hymaea (Australia) and Rhopalobrachium (Chile, Australia).
*Lawrence (1995b).
AGAPYTHINAE Sen Gupta and Crowson, 1969b: 579
Agapytho (New Zealand).
MYRABOLIINAE Lawrence and Britton, 1991: 650
Myrabolia (Australia).
*Lawrence (1988a).
their family key and in the text, in the latter case assigning the genus to a
subfamily Myraboliinae. The subfamily was not characterized as such or
indicated as new, but the subfamily name can be considered available if
it is assumed that Myrabolia is the only included genus, and hence that the
generic and subfamily characterizations coincide.
PRIASILPHINAE Crowson, 1973a: 56
Priasilpha (New Zealand) and Priastichus (Australia).
*Crowson (1973a).
TASMOSALPINGINAE Lawrence and Britton, 1991: 651
Tasmosalpingus (Australia).
876
*Lawrence (1988a).
**Lawrence and Britton (1991) characterized the genus Tasmosal-
pingus in their family key and in the text, in the lat~er case assigning the
genus to a subfamily Tasmosalpinginae. The subf~muly was not charact~r­
ized as such or indicated as new, but the subfamily name can be consid-
ered available if it is assumed that Tasmosalpingus is t?e ?nly in~lu?ed
genus, and hence that the generic and subfamily charactenzatwns comcide.
SILVANIDAE Kirby, 1837: 110
*Boving (1921), Thomas (1988, 1993). . .
BRONTINAE Erichson, 1845: 309/Blanchard, 1845b: 134 (=Ulewtmae,
Hyliotinae; incl. Cryptamorphinae, Psammoecinae)
Brontopriscus, Cryptamorpha, Dendrophagus, Psammoecus, Telephanus,
Uleiota (=Brontes, Hyliotes), etc.
* Uleiota Dendrophagus and related genera are often included in the
family Cuc~jidae. Crowson (1973a), Crowson and Ellis (1969), Thomas
(1984a, 1992), Pal (1985), Pal et al. (1985). . (1960).
**Brontinae has priority over Uleiotinae Gistel (1856: 382) and ~yho­
tinae Reitter (1879: 80), and in our opinion neither ?f the latter n~es h~s
become accepted widely enough to justify its retentiOn over Brontmae v~a
Art. 40b. It is not clear if the relative priority of the 1845 works by ~n­
chson and Blanchard can be established, but Erichson is the usual cited
author of Brontinae and his name is latinized.
SILVANINAE Kirby, 1837: 110 . *Matthews (1899) excluded Phaenocephalus from the Corylophidae,
Ahasverus, Cathartus, Monanus, Oryzaephilus, Silvanolomus, Szlvanoprus,
Silvanus, etc.
*Grouvelle (1913), Halstead (1973), Pal and Sen Gupta (1985).
PASSANDRIDAE Erichson, 1845: 304/Blanchard, 1845b: 134 (=Catogeni~ae)
Ancistria (Asia, Africa, Australia), Aulonosoma (=Laen:otmetus) (wide-
spread), Catogenus (=Scalidia) (North to So.uth A~enca).' and Passan-
dra (=Hectarthrum) (Mexico to South Amenca, Asia, Afnca, Madagas-
car, Australia). , John (1954) (see further discussion under Discolomatidae).
*This group is sometim~s included in Cucujida~ ..slipinski ( 1983, 1987,
1989) and Burckhardt and Slipinski (1991) are revismg the group. Thomas
(1988, 1993). . brus, Parasemus, Phalacropsis, Phalacrus, Stilbus, Tolyphus, etc.
**See under Brontinae regarding the 1845 works ofEnchson and Blan-
chard.
CUCUJIDAE Latreille, 1802: 210 . .
Cucujus (North America, Eurasia), Palaestes .(Neotropica~), Pedzacus
(North and Central America, Eurasia, Australia), and Platlsus (Austra-
lia).
877
LAEMOPHLOEIDAE Ganglbauer, 1899: 606
Brontolaemus, Carinophloeus, Cryptolestes, Laemophloeus, Lathropus,
Microlaemus, Narthecius, Placonotus, Rhabdophloeus, Rhinomalus,
etc.
*This group, often placed in Cucujidae, includes all of the smaller,
flattened species with submarginal carinae on the pronotum. Crowson and
Sen Gupta (1969), Iablokoff-Khnzorian (1977a-78), Lefkovitch (1959,
1961, 1962), Thomas (1984b, 1984c, 1988, 1993).
PROPALTICIDAE Crowson, 1952: 125
Discogenia (Africa) and Propalticus (Asia, Africa, Australia, Pacific).
*The larval description in Crowson and Sen Gupta (1969) is apparently
based on a misidentification, and the family relationships discussed in that
paper were influenced by that association. Crowson (in litteris) believes
that Propalticidae should be included in Laemophloeidae, and another
larva associated with adult Propalticus supports this placement. John
PHALACRIDAE Leach, 1815: 116 (incl. Phaenocephalidae)
*Blackburn (1895), Emden (1928), Guillebeau (1894), Steiner (1984).
PHAENOCEPHALINAE Matthews, 1899: 205
Phaenocephalus (Japan, southeast Asia), Phalacrinus (Australia), and
probably Sphaerostilbus (India).
where it was placed by its author (Wollaston 1873-74), and proposed a
new family for it. Lawrence (1982) and Pakaluk (1991) placed the genus
in Phalacridae, and the latter discussed the similarities of this genus to the
Australian Phalacrinus. Champion (1924c) noted the similarities of Pha-
lacrinus to the Indian genus Sphaerostilbus (Champion 1924b), which may
also belong to this group. Paulian (1950).
**The name Aphaenocephalidae Ganglbauer ( 1903: 316) is an apparent
laps us calami for Phaenocephalidae, not Aphanocephalidae as assumed by
PHALACRINAE Leach, 1815: 116
Acylomus, Biophytus, Eustilbus, Litochropus, Litochrus, Ochrolitus, Oli-
PHALACRIDAE incertae sedis:
Cyclaxyra (New Zealand).
*The placement of this genus is under review by Crowson and Slipin-
ski.
HOBARTIIDAE Sen Gupta and Crowson, 1966: 71
Hobartius (Australia, South America) and Hydnobioides (Australia).
878
CAVOGNATHIDAE Sen Gupta and Crowson, 1966: 66
Cavognatha and Taphropiestes (Australia and southern South America),
Neocercus (New Zealand), and Zeonidicola (New Zealand).
*South temperate group associated with the nests of birds. Crowson
(1965, 1973a), Sen Gupta and Crowson (1969b), Watt (1980).
CRYPTOPHAGIDAE Kirby, 1837: 111 (incl. Catopochrotidae, Hypocopridae)
*Bousquet (1989), Crowson (1980), Hinton (1945).
**The original spelling of the type genus is Kryptophagus, but Silfver-
berg (1994b) has applied to the ICZN for conservation of the spelling
Cryptophagus .
HYPOCOPRINAE Reitter, 1879: 81
Hypocoprus (Europe, Central Asia, North America).
CRYPTOPHAGINAE Kirby, 1837: 111 (incl. Telmatophilinae)
Antherophagus, Caenoscelis, Catopochrotus, Chiliotis, Cryptophagus,
Cryptosomatula, Emphylus, Henoticus, Henotiderus , Mnioticus, Para-
mecosoma, Picrotus, Pictorus, Pteryngium, Spaniophaenus, Sternodea ,
Telmatophilus, Thortus, etc.
*Coombs and Woodroffe (1962), Kieseritzky and Reichardt (1936),
Leschen and Lawrence (1991), Lyubarsky (1989, 1992), Reitter (1889),
Scott (1935).
ATOMARIINAE LeConte, 1861: 99 (incl. Ephisteminae)
Atomaria, Anchicera, Ephistemus, Salltius, etc.
CRYPTOPHAGIDAE incertae sedis:
Alfieriella (=Cyprogenia) (southern Europe, central Asia), ?Amydropa (Chi-
le).
*Ratti (1976), Wittmer (1935).
**Crowson (1980: 282) discussed a subfamily Alfieriellinae including
these genera, but he used the subfamily name conditionally and without
description so it must be considered a nomen nudum.
LAMINGTONIIDAE Sen Gupta and Crowson, 1969a: 125
Lamingtonium (Australia).
*Lawrence et al. (1993), Sen Gupta and Crowson (1969a).
LANGURIIDAE Crotch, 1873a: 184 (incl. Cryptophilidae)
*As presently defined, this family includes a number of genera for-
merly placed in Cryptophagidae, as well as Cryptophilus, which has been
included in Erotylidae (Chujo 1969). Lawrence (1991), Lawrence and
Vaurie (1983), Martinez and Barrera (1966), Rymer Roberts (1939, 1958),
Sen Gupta and Crowson (1969a, 1971).
**Arnett (1963) cites "Wiedeman, 1823" for the family name but this
refers to a generic treatment (J. Pakaluk, in litteris).
879
XENOSCELINAE Ganglbauer, 1899: 649 (=Loberinae; incl. Pharaxono-
thinae)
Bolerus, Hapalips, Loberonotha, Loberus, Pharaxonotha, Telmatoscius
;enocryptus, Xenoscelis, Zavaljus (=Eicolyctus), etc. '
Lyubarsky (1994), Sen Gupta (1968b) Sen Gupta and Crowson
(1967, 1969a). '
SETAR!OLINAE ~rowson, 1952: 127 (=Setariinae)
Setarwla (=Setana) (southern Europe).
*Falcoz (1921).
**Setariini Casey (1900: 77) is based on a preoccupied generic name
LANGURIINAE Crotch, 1873a: 184 (=Cladoxeninae) .
AcroP_teroxys, A~adastus, Cladoxena, Crotchia, Dasydactylus, Languria,
Mlcrolangurza, Platoberus, Thallisella, etc.
*Sen Gupta (1968a), Villiers (1943, 1946, 1961).
CRYPTOPHILINAE Casey, 1900: 77
Coelocryptus, Cryptophilus, and Xenoscelinus.
*Larvae of Cryptophilinae described by Sen Gupta and Crowson
(1971) were base? on misid~ntifications (see Pal and Lawrence 1986).
Crows~n_(1981) listed a fam1ly Cryptophilidae including Toraminae and
Propalt1c1dae (see above) with this group. Sasaji (1989a).
TORAMINAE Sen Gupta, 1967: 167
Atomarops, Empocryptus, Loberoschema, and Toramus .
*Sen Gupta (1967).
EROTYLIDAE Latreille, 1802: 233 (incl. Dacnidae)
*Boyle (1956), ChUjo (1969), ChUjo and ChUjo (1988-90), Delkes-
kamp (1981), Iablokoff-Khnzorian (1975), Lawrence (1988a) RymerR 0 b-
erts (1939, 1958). '
DACNINAE Gistel, 1856: 360
Cnecosa, Cryptodacne, Dacne, Microsternus Thallis etc
**Th ' ' .
e older n~~e En.gidae MacLeay (1825: 40) as originally proposed
was based on. a misidentified type genus, Engis of MacLeay (=Encaustes,
no~ Encaustmae), not Engis P~ykull (=Dacne). Any use of MacLeay's
fami.ly-~roup na~e would reqmre a proposal to the ICZN to resolve its
~P?hcatwn, but smce the name has not been used for more than a century
It Is probably best forgotten.
MEGALODACNINAE Sen Gupta, 1969: 100
Episcapha, Episcaphula, Megalodacne, etc.
ENCAUSTINAE Crotch, 1876: 476/Chapuis, 1876: 46
Aulacochilus, Encaustes, Micrencaustes, etc.
*Lawrence (1988a).

880
TRITOMINAE Curtis, 1834: Plate 498 (text) (=Triplacinae)
Jschyrus, Mycotretus, Pselaphacus, Renania, Triplax, Tritoma (=Cyrto-
triplax), etc. .
**Tritominae, based on Tritoma of Fabricius (not Geoffroy, m Myceto-
phagidae), has priority over Triplacinae Imhoff (1856: 144) for the name
of the subfamily. Tritomidae of Crotch (1873b: 78, 1873c: 42) was based
on Tritoma Geoffroy and although used for a time in place of the name
Mycetophagidae (e.g., by Casey 1900: 128) has never come into general
use. The ICZN (1994a) has conserved Tritoma Fabricius, thus also con-
serving Curtis's Tritominae.
EROTYLINAE Latreille, 1802: 233
Brachysphaenus, Cypherotylus, Erotylus, Homoeotelus, etc.
BYTURIDAE Jacquelin du Val, 1858: 211
*This and the following family have been considered as sister groups
since the study ofFalcoz (1926), but their position within the superf~ily
is uncertain. Biphyllids are usually placed near Languriidae or Erot~hdae
(Arrow 1929b, Rymer Roberts 1958), and Byturidae were once co~s1der~d
to be related to Dermestidae. Crowson (1960) suggested an affimty w1th
the Tenebrionoidea, which is supported by the single pretarsal seta in the
larva; however, the aedeagus is of a type found in Cleroidea, which also
have a single seta. Abdullah and Abdullah (1966), Barber (1942), Law-
rence (1991), Springer and Goodrich (1983, 1987, 1990, 1994).
PLATYDASCILLINAE Pic, 1914: 20
Bispinatus (Vietnam), Dascillocyphon (Malaysia, Ind?nesia), Platy-
dascillus (Malaysia, Sumatra), and Remigera (Malaysm).
BYTURINAE Jacquelin du Val, 1858: 211 . .
Byturodes (China, India), Byturus (North America, ~uras1a), and Xerasza
(=Byturellus, Satorystia) (western North Amenca, Central Europe,
Middle East, Japan).
BIPHYLLIDAE LeConte, 1861: 105 (=Diphyllidae)
Althaesia, Anchorius, Biphyllus (=Diphyllus), Diplocoelus, Gonicoelus,
etc.
*Lawrence (1991), Goodrich and Springer (1992) . . MURMIDIINAE Jacquelin du Val, 1858: 227
**Silfverberg (1992) cited "Sharp 1900 (1861)" for the famll~ name,
evidently referring to Biphyllidae Sharp (1900: 604) and assummg th~t
Biphyllus Dejean and Diphyllus Berthold (on whic~ LeConte based h1s
family-group name) are independent names. Followmg Nea~e (1939-40)
we consider Diphyllus to be an unjustified emendation of Bzphyll~s; Le-
Conte is then the author of the family-group name, not Sharp as c1ted by
Goodrich and Springer (1992: 362).
881
BOTHRIDERIDAE Erichson, 1845: 287 (incl. Anommatidae)
*Craighead (1920), J?ajoz (1980), Heinze (1943), Lawrence (1991), Pal
and Lawrence (1986), Slipinski and Pakaluk (1992), Stephan (1989).
TEREDINAE Seidlitz, 1888: 60
Oxylaemus, Sosylopsis, Sysolus, Teredolaemus, Teredus, etc.
*Slipinski and Pal (1985).
XYLARIOPHILINAE Pal and Lawrence, 1986: 197
Xylariophilus (India, southeast Asia, East Indies, Melanesia, Australia).
*Pal and Lawrence (1986).
ANOMMATINAE Ganglbauer, 1899: 894
Abromus (Europe) and Anommatus (widespread).
*This group is sometimes included in Cerylonidae. Dajoz (1968, 1977).
BOTHRIDERINAE Erichson, 1845: 287 (incl. Deretaphrinae)
Antibothrus, Ascetoderes, Bothrideres, Dastarcus, Deretaphrus, Erotyla-
thris, Lithophorus, Machlotes, Ogmoderes, Prolyctus, Pseudobothride-
res, Sosylus, etc.
*Roberts (1980), Slipinski et al. (1989).
CERYLONIDAE Billberg, 1820: 47 (=Cerylidae; incl. Aculagnathidae, Dolo-
sidae, Euxestidae, Murmidiidae)
*World generic review published by Slipinski (1990). Dajoz (1980),
Heinze (1944), Lawrence and Stephan (1975), Sen Gupta and Crowson
(1973), Slipinski (1988).
EUXESTINAE Grouvelle, 1908: 452 (incl. Metaceryloninae)
Cycloxenus, Euxestoxenus, Euxestus, Hypodacne, Hypodacnella, Meta-
eery/on, Pachyochthes, and Tachyoryctidium.
LOEBLIORYLONINAE Slipinski, 1990: 81
Loebliorylon (Asia).
**With reference to Slipinski's (1990) discussion of the etymology of
the name Cerylonidae, the name of this subfamily should be Loeblio-
ryloninae, not Loebliorylinae as proposed.
OSTOMOPSINAE Sen Gupta and Crowson, 1973: 400
Ostomopsis (Neotropical, Indo-Australian, New Caledonia).
*Lawrence and Stephan (1975).
Botrodus, Murmidius, and Mychocerinus.
*Hinton (1942).
CERYLONINAE Billberg, 1820: 47 (incl. Lapethinae)
Axiocerylon, Cautomus (=Aculagnathus), Cerylon, Glyptolopus, Mycho -
cerus (=Lapethus), Philothermopsis, Philothermus, Pleosoma, Thyro-
derus (=Dolosus), etc.
*Besuchet (1972), Dajoz (1963), Slipinski (1984).
882
**Silfverberg (1994a) has applied to the ICZN for conservation of Ce-
rylon in the sense used here.
ALEXIIDAE Imhoff, 1856: 151 (=Sphaerosomatidae, Sphaerosomidae)
Sphaerosoma (=Alexia) (Eurasia).
*Alexiids have often been included in the family Endomychidae. Sli-
pinski and Pakaluk (1992).
**Alexiidae has priority over Sphaerosomatidae Ganglbauer (1899:
913); the latter widely used name could be conserved over the former by
use of Art. 40b, but Pakaluk et al. (in press) have chosen not to do this.
DISCOLOMATIDAE Horn, 1878: 557 (=Discolomidae, Notiophygidae)
*Fukuda (1969), John (1954, 1959, 1967).
**The grammatically correct spellings of the family and nominotypical
subfamily names are Discolomatidae and -inae, respectively, not Discolo-
midae and -inae.
NOTIOPHYGINAE Jacobson, 1915: 920
Dystheamon (Madagascar), Holophygus (Central and South America),
Parmaschema (southeast Asia, central Africa), and Notiophygus,
Pachyplacus, and Praviclava (southern Africa).
DISCOLOMATINAE Horn, 1878: 557
Cassidoloma (Africa) and Discoloma (Neotropical).
APHANOCEPHALINAE Grouvelle, 1912: 198
Aphanocephalus (eastern Asia, Pacific and Indian Oceans, central Africa,
Australia), Fallia (central and South America, Hawaii), Parafallia and
Profallia (southeast Asia), and Solitarius (Brazil).
**John (1954) quoted Matthews (1899) and Ganglbauer (1903) as us-
ing "Aphanocephalidae" for a group including Aphanocephalus, but both
authors used "Pseudocorylophidae" (not a genus-based name) for this
group; Ganglbauer (1903: 316) also used a name "Aphaenocephalidae" as
an apparent lapsus calami for Phaenocephalidae Matthews (1899). The
first use of a name based on Aphanocephalus was apparently by Grouvelle
(1912).
CEPHALOPHANINAE John, 1954: 19
Cephalophanus (southeast Asia).
PONDONATINAE John, 1954: 19
Katoporus (central Africa) and Pondonatus (South Africa).
ENDOMYCHIDAE Leach, 1815: 116 (incl. Merophysiidae, Mycetaeidae)
*Lawrence (1982, 1991) and Slipinski (1990) added the holoparame-
cines and merophysiines, which were included in Merophysiidae by Crow-
son (1955); Slipinski and Pakaluk (1992) followed this but stated that En-
883
domychidae was probably not a monophyletic group. Sasaji (1978b,
1987a, 1991), Strohecker (1953, 1956, 1986).
MEROPHYSIINAE Seidlitz, 1872: 39
Coluocera, Evolocera, Merophysia, etc.
*This subfamily is often combined with the next one, and both were
included in a family Merophysiidae by Crowson (1955). Sen Gupta ( 1979)
added the genus Derolathrus (see Jacobsoniidae above) to this complex (as
a subfamily of Merophysiidae), and Sasaji (1987a) treated this as a tribe
of Merophysiinae. Silvestri (1912), Sasaji (1991).
HOLOPARAMECINAE Seidlitz, 1888: 61
Holoparamecus, Lycoperdinella, etc.
LEIESTINAE C.G. Thomson, 1863: 306
Leiestes, Phymaphora, and Rhanidea.
EUPSILOBIINAE Casey, 1895: 454 (=Eidoreinae; incl. Cerasommatidiidae)
Cerasommatidia (South America), Chileolobius (South America), Eidoreus
(=Eupsilobius) (Neotropical, Indo-Australian, Pacific), lbicarella
(South America), and Microxenus (South Africa).
*Sen Gupta and Crowson (1973) synonymized Eupsilobius Casey
(1895) (placed in Colydiidae: Murmidiinae) with Eidoreus Sharp (1885)
(placed in Erotylidae) and suggested that this taxon (for which Casey had
proposed the name Eupsilobiini) might represent a distinct subfamily of
Endomychidae. This was followed by Strohecker (1986), Lawrence (1991)
and Sasaji (1986, 1987a, as Eidoreinae; 1991 as Eupsilobiinae). Pakaluk
and Slipinski (1990) revised the group, adding Microxenus and two new
genera. Brethes (1925) placed the Brazilian genus Cerasommatidia in a
new family considered to have attributes of both Endomychidae and Coc-
cinellidae. Costa Lima (1952-56) listed Cerasommatidiidae as a synonym
of Coccinellidae, but Pakaluk et al. (in press) transferred the genus to Eu-
psilobiinae and suggested a relationship to lbicarella.
**Eupsilobiinae has priority over Cerasommatidiidae Brethes (1925:
199) and Eidoreinae Sasaji (1986: 235).
ENDOMYCHINAE Leach, 1815: 116
Bolbomorphus, Cyclotoma, Endomychus, Eucteanus, and Meilichius.
EPIPOCINAE Gorham, 1873: 20 (=Stenotarsinae)
Archipines, Danae, Epipocus, Periptyctus, Saula, Stenotarsus, etc.
*Sasaji (1978a).
' **Epipocinae has priority over the widely used Stenotarsinae Chapuis
(1876: 125) for the name of this group.
LYCOPERDININAE Redtenbacher, 1844: 120 (=Eumorphinae)
Amphix (=Corynomalus), Ancylopus, Dapsa, Encymon, Eumorphus, Indal-
mus, Lycoperdina, Mycetina, Trycherus, etc.
**Lycoperdininae has priority over the widely used Eumorphinae Gis-
tel (1856: 382) for the name of this group.
 885
884
CHILOCORINAE Mulsant, 1846: 166
MYCETAEINAE Jacquelin du Val, 1857: 102 Aspidimerus, Brumus, Chilocorus, Exochomus, Platynaspis, Telsimia, etc.
Mycetaea (Europe, southern Africa, introduced elsew~ere) . *Chapin (1965), Korschefsky (1934), Sasaji (1992),
ANAMORPHINAE Strohecker, 1953: 15 (=Mychothe~mae) COCCINELLINAE Latreille, 1807b: 70
Anamorphus, Bystus, Idiophyes, Mychothenus:. Symbwtes, etc. Adalia, Adonia, Alesia, Anatis, Anisosticta, Bulaea, Coccinella, Coelo-
*This was considered as a family by SasaJl (1987a). Pakaluk (1986), phora, Cydonia, Discotoma, Halyzia, Hippodamia, Micraspis, Myzia,
Sasaji (1978b, 1990). . .. . Psyllobora, Seladia, Singhikalia, Synonycha, Thea, Tytthaspis, etc.
**Anamorphinae has priority over Mychothemnae SasaJ1.(1~78b. 8). *Iablokoff-Khnzorian (1982), Pope (1989).
PLEGANOPHORINAE Jacquelin du Val, 1858: 186 (=Trochmdemae) EPILACHNINAE Mulsant, 1846: 190
Dadophorus, Pleganophorus, and Trochoideus . Cynegetis, Epilachna, Epiverta, Eremochilus, Mada, Subcoccinella, etc.
*Kemner (1924). . . . *Gordon (1975).
**Pleganophorinae has priority over the wtdely used Trochmdemae
Chapuis (1876: 146) for the name of this group. CORYLOPHIDAE LeConte, 1852b: 141 (=Orthoperidae)
XENOMYCETINAE Strohecker in Arnett, 1962: 802 *Lawrence (1991), Matthews (1899), Paulian (1950).
Xenomycetes (western North America). **Corylophidae has priority over Orthoperidae Jacquelin du Val (1857:
*Johnson (1986). 100) for the name of the family and nominotypical subfamily.
ENDOMYCHIDAE incertae sedis: PELTINODINAE Paulian, 1950: 19
Agaricophilus (eastern Europe) . Peltinodes (North Africa).
*Lawrence (1991), Mamaev (1977). , CORYLOPHINAE LeConte, 1852b: 141 (incl. Orthoperinae)
**Agaricophilinae Lawrence (1991: 484) and Slipinski and Pakaluk Aenigmaticum, Corylophodes, Corylophus, Ectinocephalus, Foadia, Ho-
(1992: 81) is a nomen nudum. lopsis, Hoplicnema, Hyplathrinus, Lewisium, Orthoperus, Priamima,
Rypobius, Teplinus, etc.
COCCINELLIDAE Latreille, 1807b: 70 (incl. Epilachnidae) *Paulian (1950), Pakaluk (1985a, 1985b, 1985c, 1987), Pakaluk and
*Belicek (1976), Boving (1917), Chazeau et al. (1990), Gordon (1985), Lawrence (1986).
Hodek (1973), Kamiya (1965), Klausnitzer (1970), Phuoc and Stehr SERICODERINAE Matthews, 1888: 117
(1974), Pope (1979), Sasaji (1968, 1971). . Anisomeristes, Sericoderus, etc.
STICHOLOTIDINAE Weise, 1901: 430 (=Phannae) PARMULINAE Poey 1854: 323 (=Arthrolipinae, Saciinae)
Carinodula, Catana, Cephaloscymnus, Coccidophilus, Coelopterus, Del- Alloparmulus, Arthrolips, Molamba, Sacium, etc.
phastus, Ghanius, Jauravia, Microweisea, .Nipus, Pharoscymnus, Se- *Paulian (1950) revised the African genera in this group and later
rangium, Shirozuella, Sticholotis, Sukunahzkona, etc. . (1962) applied some of these generic concepts to the Asian fauna.
*Fi.irsch (1985), Gordon (1977, 1991), Gordon and Almetda (1991), **The well-known name Saciinae Matthews (1888: 104) is predated by
Gordon et al. (1989), Sasaji (1967, 1992). . Parmulinae Poey (1854: 323), proposed as a replacement name for Clypea-
**Pharinae Casey (1899: 110)/Ganglbauer (1899: 973) 1s based on a strinae Redtenbacher (1845: 122). Clypeastrinae was based on the preoccu-
preoccupied generic name, Pharus Mulsant (now Pharoscymnus). pied generic name Clypeaster Dejean (not Lamarck), replaced in Poey
COCCIDULINAE Mulsant, 1846: 266 (=Tetrabrachinae) . (1854) by Parmulus Gundlach; however, the oldest replacement name for
Azya, Coccidula, Exoplectra, Monocoryn.a, Nov.ius, Oryssomus, Rhyzobzus, Clypeaster Dejean is Clypastraea Haldeman, 1842, which has priority over
Rodolia, Sumnius, Tetrabrachys (=Lzthophzlus), etc. not only the later replacement names Aspidocha Gistel, 1848 and Par-
*Miyatake (1988), Sasaji (1989b). mulus Gundlach, 1854 but also the subjective synonym Sacium LeConte,
SCYMNINAE Mulsant, 1846: 210 . 1852, the name in current use for this genus. Pakaluk et al. (in press) used
Brachiacantha, Bucolus, Cranophorus, Cryptognatha, Dw~us, Hypera- Parmulinae in place of Saciinae but did not address the generic synonymy.
spis, Nephus, Ortalia, Pentilia, Poria, Scymnus, Selvadzus, Stethorus,
Zilus (=Scymnillus), etc.
*Gordon (1976, 1985), Pope and Lawrence (1990).

886
LATRIDIIDAE Erichson, 1842: 122 (=Lathridii~ae)
*Belon (1902), Dajoz (1962, 1971), Hmton (1941b, 1945), Rucker
(1978, 1985), Watt (1969). . ...
**Corticariidae has priority over the umversally used Lathndndae for
the family name. The original spelling of the type genus ?f the latter name
is Latridius; this name and Latridiidae were used by S1lfverberg (1992)
and Pakaluk et al. (in press) and are used here.
LATRIDIINAE Erichson, 1842: 122 . . . Guadeloupe?).
Adistemia, Aridius, Cartodere, Dienerella, Enicmus, Latrzdzus (=Lathrz-
dius), Lithostygnus, Metophthalmus, etc.
*Andrews (1976b ), Sen Gupta (1976).
CORTICARIINAE Curtis, 1829: Plate 283 (text)
Bicava, Caserus, Corticaria, Cortinicara, Fuchsina, Melanophthalma,
Migneauxia, etc. . . .
*Caserus was included in the mycetophag1d subfam1ly Esarc1~a~ by
Dajoz (1969), but it clearly belongs in Latridiidae and represents a dlstmc-
tive type of corticariine. Andrews (1976a), Johnson (1975).
Superfamily TENEBRIONOIDEA Latreille, 1802: 165
(=Heteromera) .
*Crowson ( 1955, 1981) considered this to be a section within the prev10us
superfamily. The limits of this superfamily are generally agreed up~n, although
Mycetophagidae, Colydiidae and the tenebri~nid ~enus Myrmechzxenus were
included in Clavicornia in many older classifications. Crowson (1960) sug-
gested that Biphyllidae and Byturidae could be included in J:Ieter~mera base.d
on aedeagal structure and the oblique trochanterofemoral aruculauon, and th1s
wed by Abdullah (1973, 1974a, 1976) and Lawrence (1977); however,
was f ollo · C · 'd
a much stronger case can be made for their inclusion m ucUJOl ea.
MYCETOPHAGIDAE Leach, 1815: 110
*Hayashi (1971 ), Lawrence ( 1977, 1987b), Leschen and Lawrence
(1991), Miyatake (1957, 1959), Nikitsky (1988b, 1989b), Parsons (1975),
Reitter (1885).
**The name Tritomidae Crotch (1873b: 78, 1873c: 42), used by .a few
authors for this family, is based on a misidentified type genus (see discus-
sion under Erotylidae: Tritominae).
ESARCINAE Reitter, 1882: 133
Esarcus (southern Europe, North Africa). . .
*The genus Pseudesarcus (Champion 1913) belongs to the tenebnomd
subfamily Lagriinae. Dajoz (1964).
887
MYCETOPHAGINAE Leach, 1815: 110
Catopius, Eulagius, Filicivora, Litargops, Litargus, Mycetophagus, Pseud-
ochrodes, Pseudotriphyllus, Triphyllina, Triphyllus, Typhaea, Typha-
eola, etc.
BERGININAE Leng, 1920: 246
Berginus (Holarctic ).
MYCETOPHAGIDAE incertae sedis:
Nesolathrus (Seychelles, Christmas 1., Australia, Society Is., Austral Is.,
*This genus was originally placed in the Latridiidae and compared to
Holoparamecus (see Endomychidae) and Derolathrus (see Jacobsoniidae)
(Scott 1922). It appears to belong to Mycetophagidae, based on features
of both adults and associated larvae from northern Australia and Christmas
Island; however, its position within the family is uncertain.
ARCHEOCRYPTICIDAE Kaszab, 1964: 361
Archeocrypticus (Chile and Argentina, New Zealand, Australia), Enne-
boeus (=Uloporus) (North to South America, Australia), Pseudenne-
boeus (Brazil), Sivacrypticus (Asia, Africa, Australia), and Austra-
lenneboeus, Enneboeopsis, Falsoplatydema, Gondwanenneboeus, No-
thenneboeus and Wattianus (Australia).
*This group was included in Tenebrionidae by Kaszab (1964, 1981,
1984), but recognized as distinct family by Watt (1974b). Lawrence (1977,
1994b), Triplehorn and Wheeler (1979).
PTEROGENIIDAE Crowson, 1953: 47
Histanocerus (S~utheast Asia, East Indies, New Guinea), Katagenius
(southern lnd1a), Kryptogenius (southern India, Greater Sunda Is.), Pte-
rogenius (Sri Lanka), and Tychogenius (Borneo).
*Lawrence (1977), Burckhardt and Lobi (1992).
CIIDAE Leach in Samouelle, 1819: 206 (=Cisidae)
*Lawrence (1971, 1974a, 1987c), Miyatake (1954).
SPHINDOCIINAE Lawrence, 1974a: 24
Sphindocis (California).
*Lawrence (1974a, 1974b).
. **This subfamily was described as new in Lawrence (1974b: 9, pub-
hshed 28 June), but already had been made available in Lawrence (1974a:
24, published 22 January).
CIINAE Leach in Samouelle, 1819: 206 (incl. Orophiinae, Octotemninae)
Ceracis, Cis, Ennearthron, Euxestocis, Falsocis, Hadraule, Malacocis,
Nipponocis, Octotemnus (=Orophius), Orthocis, Porculus, Rhopalodon-
tus, Scolytocis, Xylographus, etc.
888
TETRATOMIDAE Billberg, 1820: 34
*This family may well be paraphyletic and its relationships to Myceto-
phagidae, on the one hand, and Melandryi~ae, on the.?t?er, are still.some-
what uncertain. The Penthinae were combmed by Bovmg and Cratghead
(1931) with eustrophine Melandryidae (see below) and dacnine e.rotylids
(see above) to form a family Dacnidae, based ?n larval adaptat10~s for
feeding on soft fungi. Crowson (1966a) and y1edma (1971) co.nstdered
Eustrophinae to be annectant between Tetratomtdae.and Me.l~drytdae, and
Hayashi (1975) placed Holostrophus in Tetratomtdae. Nildts~y (~989a)
placed the melandryid subfamilies Hallomeninae and Eustrophmae m Tet-
ratomidae. Crowson (1964a), Nikitsky (1988a, 1989b).
PISENINAE Miyatake, 1960: 124
Pisenus (eastern North America), Notopisenus (Chile), and Triphyllia
(=Eupisenus) (western North America, Caucasus).
*Hayashi (1972), Miyatake (1960), Nikitsky (1992b).
TETRATOMINAE Billberg, 1820: 34
Falsoxanthalia (China), Incolia (North America), and Tetratoma (=Ab-
strulia) (North America, Eurasia).
*Crowson (1964a).
PENTHINAE Lacordaire, 1859: 456
Penthe (North America, Asia).
*Hayashi (1972).
MELANDRYIDAE Leach, 1815: 104 (=Serropalpidae)
*As mentioned above, Hallomeninae and Eustrophinae are sometimes
included in the previous family. The Melandryinae appears to be a mono-
phyletic group, most members of which have larvae ~hich inhabit hard
substrates, such as bracket fungi or wood. Burakowsk1 (1995), Crowson
(1966a), Hayashi (1975), Lawrence (1991), Nikitsky (1989a), Nikitsky and
Belov (1982a, 1982b), Viedma (1966, 1971).
HALLOMENINAE Mulsant, 1856: 232/Gistel, 1856: 384
Hallomenus (North America, Eurasia) and Mycetoma (Eurasia).
EUSTROPHINAE Gistel, 1856: 382
Eustrophinus, Eustrophopsis, Eustrophus, Holostrophus, Synstrophus, etc.
MELANDRYINAE Leach, 1815: 104 (incl. Dircaeinae, Hypulinae)
Abdera, Callidircaea, Ctenoplectron, Dircaea, Hylobia, Hypulus, Lederia,
Mecorchesia, Melandrya, Mystes, Neorchesia, Orchesia, Phrygano-
philus, Serropalpus, Talayra, Xylita, Zilora, etc.
OSPHYINAE Mulsant, 1856: 300 (1840) (=Nothinae)
Conopalpus, Osphya (=Nothus), etc.
**Nothinae Shuckard (1840: 51) has priority over Osphyinae for the
subfamily name, but we maintain the latter well-known name by provision
of Art. 40b.
889
MORDELLIDAE Latreille, 1802: 183
*The subfamily Anaspidinae, considered to be mordellids in older clas-
sifications, are now included in Scraptiidae. Batten (1990), Ermisch
(1950), Franciscolo (1952, 1957, 1965, 1967, 1980), Liljeblad (1945),
Odnosum (1991).
CTENIDIINAE Franciscolo, 1951: 56
Ctenidia (South Africa).
MORDELLINAE Latreille, 1802: 183
Conalia, Glipa, Hoshihananomia, Mordella, Mordellistena, Stenalia, To-
moxia, etc.
RHIPIPHORIDAE Gemminger and Harold, 1870: 2117 (1853)
*Besuchet (1956), Selander (1957).
**Selander (1957) discussed the problem of misidentification of Rhipi-
phorus by many 19th-Century authors, but apparently made no formal
application to the ICZN to resolve the problem. Laporte (1840b: 261),
Costa.(1853), Gerstaecker (1855) and other commonly cited early users of
a family-group name based on Rhipiphorus used the generic name in the
sense of Fabricius, 1792 (now Metoecus), not Bose, 1791 (=Myodites).
Met~ecus and Rhif!i!'hor~s ar~ now included in different tribes of Rhipi-
phonnae, so the mlSldentlficatlon affects tribal names, and the matter must
be referred to the ICZN for a decision (Art. 41). In the meantime Gem-
minger and Harold (1870: 2117) were apparently the first to use a family-
group name correctly based on Rhipiphorus Bose, so we date the name
from then. Myoditini Costa (1853: 2) is then the oldest available name for
the family and nominotypical subfamily and tribe, but since Myodites has
long b~~n. synonym.ize~ under !?hipiphorus Bose and Rhipiphorini replaced
Myodttlm we can JUstify retaming Rhipiphorini via Art. 40b. According
to Neave (1939-40), the original spelling of Rhipiphorus is Ripiphorus·
further research is needed. '
PELECOTOMINAE Seidlitz, 1875: 104
Pelecotoma (North America, Europe) and Trigonodera (North and South
A!llerica, Eurasia, Africa, Madagascar, New Guinea, Australia), etc.
*Svacha (1994).
MICHOLAEMINAE Viana, 1971: 69
Ancholaemus and Micholaemus (South America), and Clinops (Asia Mi-
nor, South Africa).
*Viana (1971).
PT~OPHORINAE Gerstaecker, 1855: 9 (=Evaniocerinae)
Ptzlophorus (=Evaniocerus) (Eurasia, Africa, Australia) and Toposcopus
(western North America).
HEMIRHIPIDIINAE Heller, 1921: 168 (=Nephritinae)
Nephrites (=Hemirhipidius) and Sitarida (Australia).
890
*Riek (1973), Toyama (1986), Toyama and Hatayama (1985). . *Lawrence (1994a) suggested that this family forms a monophyletic
**Nephritinae Selander (1957: 101) was prop?sed to r~p.lace Hem~­
rhipidiinae because of the generic synon~my, but. m ?ur opi~I?n ~ephri­
tinae has not become sufficiently established to JUStify retammg It over
Hemirhipidiinae via Art. 40b.
RHIPIDIINAE Gerstaecker, 1855: 23
Rhipidioides (Australia), Rhipidius (Eurasia, Africa), etc.
*Riek (1955).
**According to Neave (1939-40), the original spelling of the type ge-
nus is Ripidius; further research is needed.
RHIPIPHORINAE Gemminger and Harold, 1870: 2117 (1853)
Macrosiagon and Rhipiphorus (widespread), and Metoecus (Europe).
*Linsley et al. (1952).
COLYDIIDAE Erichson, 1842: 213 (incl. Adimeridae, Monoedidae) . America and Asia), Phloeodes (California, Mexico), Scoriaderma
*Preliminary studies (Lawrence 1994a) have shown t?at Pycnom.e n?ae
are more closely related to Monommatidae and Zophenda~ (as delimited
below) than they are to Colydiinae, but this needs confirmatiOn. B~rakow­
ski and Slipiri.ski (1986), Craighead (1920), Dajoz (1977), Hayashi (1972),
Ivie and Slipiri.ski (1990), Law~ence (1980), Nikitsky a~d Belov (1980),
Pope (1961 ), Slipiri.ski (1985), Slipiri.ski and Burakowski (1988), Stephan
(1989). . .
**Sarrotriini Billberg (1820: 9) has priority over Orthocenm Blanch~rd
(1845b: 29) for the name of one tribe and for the family name. R.ete~t10n
of Orthocerini over Sarrotriini can be justified by Art. 40b, but m ~Ither
case this name would date from 1820. Silfverberg (1994a) has apphed to
the ICZN for conservation of Colydiidae over these names and over Cery-
lonidae Billberg (1820) (when the latter family is included here) .
PYCNOMERINAE Erichson, 1845: 290
Dechomus, Pycnomerus, and Pycnomerodes.
COLYDIINAE Erichson, 1842: 213
Acropis, Afrorthocerus, Aprostoma, Aulonium, Bitoma, Cicones, Colydium,
Coxelus, Endestes, Enhypnon, Gempylodes, Langelandia, Lasconotus,
Monoedus (=Adimerus), Namunaria, Nematidium, Neotrichus, Or:ho-
cerus (=Sarrotrium), Phreatus, Plagiope, Pristoderus, Pseudaulomum,
Rhagodera, Rhopalocerus, Rytinotus, Synchita, Tarphius, Todima, Tra-
chypholis, etc.
MONOMMATIDAE Blanchard, 1845b: 16 (=Monommidae)
Aspathenes (Neotropical), Hyporhagus (New World), lnscutomonomma
(Africa), Monomma (Asia, Africa, Madagascar), etc.
891
group with Zopheridae (as delimited below) and pycnomerine Colydiidae.
Freude (1955a, 1955b, 1957, 1958).
**The grammatically correct spelling of the family name is Monom-
matidae (e.g., Costa Lima 1952-56), which has been used occasionally in
place of the better-known spelling Monommidae.
ZOPHERIDAE Solier, 1834: 505 (incl. Merycidae)
*As here understood, this family excludes the Ulodinae (see Ulodidae
below) and the genera Melytra and Parahelops (see Perimylopidae below).
Doyen and Lawrence (1979), Lawrence (1994a).
ZOPHERINAE Solier, 1834: 505 (incl. Nosoderminae)
Meralius (Cuba, Venezuela), Noserinus (Brazil), Noserosus (North Amer-
ica), Nosoderma (North and Central America), Phellopsis (North
(Central Africa, Comoros), Zopherosis (Australia), and Zopherus
(North and Central America).
USECHINAE Hom, 1867: 294
Usechus (western North America and Japan) and Usechimorpha (western
North America).
*Blaisdell (1929), Kamiya (1963).
ZOPHERIDAE incertae sedis:
Cotulades, Docalis, and Latometus (Australia).
*These three genera were placed in Zopheridae by Watt (197 4b) and
Doyen and Lawrence (1979), but their position within the family is not
settled (Lawrence 1994a).
ULODIDAE Pascoe, 1869: 31 (incl. Merycidae)
Arthopus (New Zealand), Brouniphylax (New Zealand), Dipsaconia (Aus-
tralia), Exohadrus (New Zealand), Ganyme (Australia), Meryx (Austra-
lia), Notocerastes (Australia), Phaennis (Australia), Pte rode res (Chile),
Syrphetodes (New Zealand), Trachyderas (Chile), Trachyderastes (New
Caledonia), Ulodes (Australia), etc.
*Crowson (1953: 43) placed Meryx in a new family Merycidae, but
Watt (1974b) suggested that the genus be placed in Zopheridae, in spite
of the 4-4-4 tarsi. Doyen and Lawrence (1979) expanded Ulodinae to
include not only Meryx but a number of other Southern Hemisphere taxa
formerly placed in Tenebrionidae. Lawrence (1994a) elevated this taxon
to family rank and gave a key to world genera.
PERIMYLOPIDAE St. George, 1939: 212
Chanopterus (southern South America), Darwinella (Falkland Is.), Hydro-
median (southern South America, South Georgia), Melytra (Tasmania),
892
Parahelops (southern South America and Falkland Is.), Perimylops
(South Georgia), and Sirrhas (Tasmania).
*Lawrence (1994a) transferred Sirrhas from Chalcodryidae and Mely-
tra and Parahelops from Zopheridae to this family. Doyen and Lawrence
(1979), St. George (1939), Watt (1967b, 1970, 1974a). . chosen not to replace the well-known name Lagriinae with the older name
**The name Parahelopinae of Watt (1974b: 424), Elgueta and Arna-
gada (1989: 37), and Lawrence and Britton (1991: 665) is a nomen nudum.
CHALCODRYIDAE Watt, 1974a: 24
Chalcodrya, Philpottia, and Onysius (New Zealand).
*Watt (1974a) also included the Tasmanian Sirrhas, but Lawrence
(1994a) transferred that genus to Perimylopidae.
TRACHELOSTENIDAE Lacordaire, 1859: 567
Trachelostenus (Chile).
*This group was formerly included in Lagriidae (Borchmann 1936,
Seidlitz 1916) or tentatively Pythidae (Watt 1974b); it was given family
status by Watt (1987). Some features of both adult and putative larvae
suggest a possible relationship to the Tasmanian tenebrionid genus Leaus
(Matthews and Lawrence 1992). If further study supports a sister group
relationship between these two taxa, then Trachelostenus will have to be
moved to Tenebrionidae or Leaus to Trachelostenidae.
TENEBRIONIDAE Latreille, 1802: 165 (incl. Alleculidae, Cossyphodidae,
Lagriidae, Nilionidae, Petriidae, Rhysopaussidae, Tentyriidae)
*The subfamily classification is based mainly on that of Doyen et al.
(1990), except that Zolodininae is given subfamily rank, as proposed by
Watt (1974b). Doyen (1972), Doyen and Lawrence (1979), Doyen and
Tschinkel (1982), Hayashi (1964, 1966), Medvedev (1977, 1990), Nikitsky
(1983), Steiner (1995), Tschinkel and Doyen (1980), Watt (1967a, 1992).
**Further work is needed in relating early family-group names to the
subfamilies recognized here. Lagriinae, Alleculinae and Coelometopinae
are definitely predated by other names, probably several each in the last
two cases. There are many Solier 1834-38 names of uncertain placement.
LAGRIINAE Latreille, 1825: 381 (1820) (incl. Cossyphinae, Pycnocerinae)
Adelium, Adelonia, Anaedus, Arthromacra, Belopus, Brycopia, Cardia-
thorax, Chaetillus, Cossyphus, Cymbeba, Ecnolagria, Goniadera, Isopt-
eron, Laena, Lagria, Lorelus, Luprops, Mitua, Paratenetus, Pheloneis,
Phobelius, Pycnocerus, Rhypasma, Seirotrana , Sphargeris, Statira,
Zeadelium, etc.
*Borchmann (1936), Merkl (1987).
893
**Cossyphinae Latreille (1802: 164) and Lachninae Billberg (1820: 34)
have priority over Lagriinae. Lachna, type species of the latter name, is a
long-forgotten junior objective synonym of Lagria, allowing retention of
Lagriinae over Lachninae by use of Art. 40b (Silfverberg 1992). We have
Cossyphinae because the latter group has been merged only recently and
somewhat tentatively with lagriines.
PHRENAPATINAE Solier, 1834: 488
Archaeoglenes, Clamoris, Delognatha, Dioedus, Peneta, Phrenapates,
Picnotagalus, Scolytocaulus, Tagalinus, Zypoetes, etc.
*Doyen and Lawrence (1979).
**Solier (1834: 488) actually used the names "Phrepatides" and Phre-
pates, the latter name evidently a misspelling of Phrenapates Gray, 1832
(Neave 1939-40).
ZOLODININAE Watt, 1974b: 401
Tanylypa (Tasmania) and Zolodinus (New Zealand).
*Doyen et al. (1990) included this group within Pimeliinae; Doyen
(1994) used it as an outgroup in his analysis of selected pimeliine taxa.
PIMELIINAE Latreille, 1802: 166 (=Tentyriinae)
Adesmia, Akis, Anepsius, Araeoschizus, As ida, Cnemeplatia, Coelus, Cryp-
toglossa, Elenophorus, Epitragus, Eurychora, Falsomycterus, Lachno-
gya, Nyctoporus, Pimelia, Platyope, Psammetichus, Salax, Sepidium,
Stenosis, Tentyria, Trimytis, Typhlusechus, Vacronus, etc.
*Doyen (1994).
TENEBRIONINAE Latreille, 1802: 165
Acropteron, Alphitobius, Amarygmus, Ammophorus, Blaps, Blapstinus,
Bolitotherus, Callismilax, Centronopus, Cyphaleus, Eleodes, Gonoce-
phalum, He lea, Heterocheira, Leichenum, Meneristes, Neatus, Nyctozo-
ilus, Palorus, Rhipidandrus, Rhysopaussus, Tauroceras, Tenebrio, Tita-
ena, Toxicum, Tribolium, Uloma, etc.
*Doyen (1985, 1989), Matthews (1992, 1993). The genus Leaus
Matthews and Lawrence (1992) is tentatively included here.
ALLECULINAE Laporte, 1840b: 242 (=Cistelinae; incl. Omophlinae)
Allecula, Chromomoea, Cteniopus, Eucaliga, Hymenorus, Lobopoda, Ly-
stronychus, Mycetochara, Omophlus, Petria, Prionychus, Tanychilus,
Xystropus, etc.
**The older name Cistelinae Latreille (1802: 188) was based on a mis-
identified type genus (Cistela of Fabricius, not Geoffroy or Schaeffer) and
is thus unavailable; Cistela Geoffroy has also been suppressed (ICZN
1994a). At least Cteniopodinae Solier (1835: 245) and Xystropodinae
~oli~r (1835: 229) have priori~y for the name of this subfamily. ("Cteniop-
mae was used over Omophlmae for a subgroup in Silfverberg (1992)).
894
DIAPERINAE Latreille, 1802: 161
Alphitophagus, Corticeus, Crypticus, Diaperis, Gnatocerus, Hyocis, Leio-
chrodes, Menimus, Myrmechixenus, Neomida, Nilio, Pentaphyllus, Pha-
leria, Platydema, Spiloscapha, Trachyscelis, Tyrtaeus, Ulomoides, etc.
*Doyen (1984).
COELOMETOPINAE Lacordaire, 1859: 358/Schaum, 1859: 71
Alobates, Apterotheca, Bradymerus, Chariotheca, Cnodalon, Coelocnemis,
Coelometopus, Cuphotes, Cyrtosoma, Derosphaerus, lphthiminus, Oea-
tus, Promethis, Strongylium, Talanus, Upis, Zophophilus, etc.
*Doyen (1989), Matthews and Doyen (1989).
**At least Stenochiinae Kirby (1837: 238) (based on Stenochia Kirby,
1819, =Strongylium) and Cnodalinae Gistel (1856: 382) have priority over
Coelometopinae for the subfamily name. Also, according to Neave
(1939-40) Strongylium Kirby, 1819 (not Ditmar, 1809, Protozoa) is preoc-
cupied and the generic and tribal names will need to be replaced; further
research is needed.
TENEBRIONIDAE incertae sedis:
COSSYPHODINAE Wasmann, 1899: 161 (incl. Cossyphoditinae, Para-
melloninae)
Cossyphodes (Africa, Madeira, Canary Is.), Cossyphodinus (India), Cossy-
phodites (Africa), Esemephe (Peru), Paramellon (India), Paramellops
(South Africa), etc.
*This group was placed in the Tenebrionidae by Crowson (1955) and
Watt (1974b), but its position in the family is still uncertain. Andreae
(1961), Steiner (1980).
PROSTOMIDAE C.G. Thomson, 1859: 84
Dryocora (New Zealand, Australia) and Prostomis (widespread).
*Prostomids were considered to be Cucujidae until Boving (1921) ele-
vated the group to family rank based on larval characters, a move later
supported by genitalic features (Wilson 1930). Crowson (1968) placed the
family in Heteromera near Colydiidae. Hayashi (1969), Schawaller (1993).
SYNCHROIDAE Lacordaire, 1859: 544
Synchroa (North America, Asia) and Mallodrya (North America).
*This small group is often included in Melandryidae, but Boving and
Craighead (1931) and Crowson (1966a) noted the larval similarities to
Zopheridae and Stenotrachelidae (as Cephaloidae) and recognized the
group as a distinct family. Hayashi (1975) placed Synchroa in Cephaloidae
based on larval characters.
895
OEDEMERIDAE Latreille, 1810: 216
*Arnett 1950, 1951, 1961), Hudson (1975), Rozen (1960), Svihla
(1986).
CALOPODINAE Costa, 1852: 4
Calopus (North America, Eurasia), Sparedropsis (Asia), and Sparedrus
(Europe, North and Central America).
OEDEMERINAE Latreille, 1810: 216 (incl. Nacerdinae)
Agasma, Alloxantha, Asclera, Chrysanthia, Copidita, Ditylus, Eobia, Isch-
nomera, Nacerdes, Oedemera, Oncomera, Oxacis, Pseudolycus, Sessi-
nia, Stenostoma, Xanthochroa, etc.
STENOTRACHELIDAE C.G. Thomson, 1859: 124 (=Cephaloidae)
*Hayashi (1963) placed Stenocephaloon in Melandryidae, but he later
(1975) returned it to this family. Mamaev (1973) suggested that the first
three subfamilies each be given family rank. Arnett (1953).
**Stenotrachelidae has priority over Cephaloidae for the name of the
family, and was used by Silfverberg (1990, 1992).
STENOTRACHELINAE C.G. Thomson, 1859: 124
Anelpistus (Nearctic), Scotodes (Palearctic), Stenocephaloon (Japan), and
Stenotrachelus (boreal Holarctic ).
*Mank (1942), Saalas (1913).
NEMATOPLINAE LeConte, 1862: 264
Nematoplus (Holarctic).
*Abdullah (1965), Mamaev (1973).
CEPHALOINAE LeConte, 1862: 259
Cephaloon (Holarctic ).
STENOTRACHELIDAE incertae sedis:
STOLIINAE Nikitsky, 1985: 32
Stolius (Japan).
*This genus has usually been placed in Melandryidae, but was trans-
ferred by Nikitsky (1985, 1992a) to Cephaloidae and made the type of a
new subfamily. Sasaji (1987b ).
MELOIDAE Gyllenhal, 1810: 481 (incl. Tetraonychidae)
*Bologna (1991), Gupta (1966, 1971a, 1971b, 1978), Kaszab (1969),
MacSwain (1956), Selander (1964, 1966a, 1991).
**Horiidae Latreille (1802: 182) has priority for the family name as
not.ed but not implemented by Watt (1975) and Selander (1991); an appli-
catiOn to the ICZ.N to conserve Meloidae has just been submitted (see
under Nemognathmae below). The name Cantharidae Latreille (1802: 185)
also .refers to this family but is based on a misidentified type genus, Can-
tharzs of Muller (now Lytta), not Linnaeus; see Watt (1975) and discussion
above under Cantharidae.
896
ELETICINAE Wellman, 1910: 221
Anthicoxenus, Ceriselma, Eletica, Eospasta, !selma, Morphozonitis (=Ert-
liana), Protomeloe, Spastica, Xenospasta, etc.
*Abdullah (1964b), Kaszab (1966), Selander (1966a, 1966b, 1967),
Werner (1974).
MELOINAE Gyllenhal, 1810: 481 (incl. Lyttinae)
Cerocoma, Cysteodemus, Epicauta, Epispasta, Eupompha, Linsleya, Ly-
dus, Lytta, Megetra, Meloe, Mylabris, Oenas, Prolytta, Pyrota, Wagne-
ronota, etc.
*Pinto (1984), Selander (1964).
**Conservation of the genus Mylabris and tribal name Mylabrini Bill-
berg (1820: 36) as used here (rather than in Bruchidae) has been proposed
by Borowiec (1988).
NEMOGNATHINAE Laporte, 1840b: 280 (=Horiinae, Zonitidinae)
Apalus, Cissites, Gnathium, Horia, Hornia, Meloetyphlus, Nemognatha,
Palaestra, Sitaris, Stenodera, Synhoria, Tetraonyx, Tricrania, Zonitis,
etc.
*Cros (1924, 1927), Parker and Boving (1924).
**Horiinae Latreille (1802: 182) has priority over the well-known name
Nemognathinae as well as Zonitidinae Mulsant (1857: 164) for the sub-
family name, and was used by Selander (1991). However, M.A. Bo.logna
and J.D. Pinto have applied to the ICZN (Case 2924) for conservatiOn of
Nemognathinae over Horiinae or Zonitidinae for the subfamily name, and
for Meloidae over Horiidae for the family name.
MYCTERIDAE Blanchard, 1845b: 97 (incl. Hemipeplidae)
*Mycterids have been placed in Pythidae or Melandryidae by many
workers but Crowson and Viedma (1964) made a case for treating the
group a; a separate family and adding the hemipeplines, which had previ-
ously been placed in Cucujidae. Iablokoff-Khnzorian (1985b), Lawrence
(1991) . .
**The older name Rhinomaceridae Fleming (1821: 50) is based on
Rhinomacer of Fabricius (now Mycterus), a misidentification or junior
homonym of Rhinomacer Geoffroy (see under Nemonychidae), and is
therefore unavailable. The ICZN ( 1994a) has effectively suppressed Rhino-
macer in all senses.
MYCTERINAE Blanchard, 1845b: 97
Mycterus (North America, Eurasia) and Mycteromimus (Seychelles).
LACCONOTINAE LeConte, 1862: 254
Conomorphus (Neotropical), Eurypinus (Africa), Eurypus (South Amer-
ica), Lacconotus (North America), Loboglossa (Chile, Australia), Tri-
chosalpingus (Australia), etc.
*Costa and Vanin (1984).
897
**The forgotten name Eurypinae J. Thomson (1860a: 63) (proposed in
Cleridae!) has priority over Lacconotinae for the subfamily name.
HEMIPEPLINAE Lacordaire, 1854: 404
Hemipeplus (widespread) and Holopeplus (West Indies).
*Thomas (1985), Thomas and Woodruff (1986).
BORIDAE C.G. Thomson, 1859: 117
*Although LeConte (1862) recognized the close relationship of Boros
and Lecontia, the latter was retained in Pythidae by most authors until
Young (1985a) transferred it to this family on the basis of larval and adult
features . Synercticus had also been placed in Pythidae based on a larval
misidentification (Crowson 1968, Lawrence 1982, Lawrence and Britton
1991). Lawrence and Pollock (1994) transferred Synercticus to Boridae
and removed Osphyoplesius from Boridae, placing the latter tentatively in
Pythidae (see below). lablokoff-Khnzorian (1985b), Lawrence and Britton
(1994), St. George (1931), Spilman (1952).
BORINAE C.G. Thomson, 1859: 117
Boros (North America, Eurasia) and Lecontia (North America).
SYNERCTICINAE Lawrence and Pollock, 1994: 36
Synercticus (Australia, New Guinea).
TRICTENOTOMIDAE Blanchard, 1845b: 137
Autocrates and Trictenotoma (Asia).
*These large beetles superficially resemble prionine cerambycids.
Crowson (1955) included Trimitomerus in this family, and Watt (1987)
placed Ischyomius here (see below). Gahan (1908), Pollock and Lawrence
(1995).
PYTHIDAE Solier, 1834: 496
Anaplopus (Australia), Priognathus (North America), Pytho (Holarctic),
and Sphalma (North America).
*Young (1976a) added Sphalma and (1985a) removed Lecontia to Bo-
ridae. Anaplopus has been placed in Pedilidae and Tenebrionidae, but both
adult and larval characters confirm its placement in Pythidae (Lawrence
1987b, Pollock and Lawrence 1995). Abdullah (1966a), Hayashi (1969),
Iablokoff-Khnzorian (1985b), Pollock (1991).
PYTHIDAE incertae sedis:
Ischyomius (Panama, South America).
*Ischyomius, type genus of Ischyomiinae Champion (1886: 258), was
placed in Trictenotomidae by Watt (1987) on the basis of adult features
only; it has also been included in Melandryidae. Pollock and Lawrence
(1995) retained the genus in Pythidae pending a future revision.
898
Trimitomerus (USA: Arizona).
*Crowson (1980) suggested that Trimitomerus plus Trictenotoma (Tric-
tenotomidae) could constitute a subfamily of Pythidae. Pollock and
Lawrence (1995) tentatively retained Trimitomerus in Pythidae pending
a study of the larva. Spilman (1952).
Osphyoplesius (southeastern Europe).
*This genus was placed by Reitter (1917) in a tribe Osphyoplesiini
within the tenebrionid subfamily Borinae (now Boridae). Lawrence and
Pollock (1994) and Pollock and Lawrence (1995) tentatively placed it
in Pythidae.
PYROCHROIDAE Latreille, 1807a: 199 (incl. Cononotidae, Pedilidae, Pili-
palpidae)
*Iablokoff-Khnzorian (1985b), Pollock (1994).
TYDESSINAE Nikitsky, 1986: 1188
Tydessa (eastern Asia, western North America).
*The genus Tydessa was placed by Peacock (1982) in Techmessinae
and considered to be closely related to /ncollogenius. Nikitsky (1986) de-
scribed the larva of T lewisi (Pic) and placed the genus in a new tribe of
his family Pilipalpidae. Pollock (1992a) described a North American spe-
cies of Tydessa in Pilipalpinae, and more recently (Pollock 1994) pub-
lished a cladogram in which Tydessinae is the sister group of the remain-
ing Pyrochroidae, and Pilipalpinae the sister group of Pedilinae plus Pyro-
chroinae.
PILIPALPINAE Abdullah, 1964a: 4 (=Techmessinae)
Cycloderus (Chile), Exocalopus (New Zealand), Incollogenius (Madagas-
car), Morpholycus (Australia), Paromarteon (Australia), Pilipalpus
(Chile), Techmessa (New Zealand), Techmessodes (New Zealand), and
Temnopalpus (Australia).
*Although Crowson (1955) included Techmessa and its allies in Pyro-
chroidae, Lawrence (1982, 1987b) and Watt (1987) excluded them from
Pyrochroidae as then defined, placing them in Pythidae, and Nikitsky
( 1986) treated the group as a separate family. The Australian forms were
also included in Pythidae by Britton (1970) and Lawrence and Britton
( 1991, 1994). With the inclusion of Pedilinae in Pyrochroidae and the
discovery of transitional larval features in some Pedilus (Young and Pol-
lock 1991), the balance of evidence favored a return to Pyrochroidae, and
Pollock (1994) has provided strong evidence for this move. A detailed
review with descriptions of new genera has been submitted for publication
by Pollock. Abdullah (1964a), Paulus (1972b), Pollock (1992a, 1992b).
PEDILINAE Lacordaire, 1859: 574
Anisotria (western North America) and Pedilus (Holarctic).
899
. *Pedil~s has often been placed in Anthicidae or in a separate family
Wtth certam groups (e.g., Eurygeniinae) now placed in Anthicidae. Young
(1~84) and Yo~~g and Pollock (1991) expanded the limits ofPyrochroidae
to mclude Pedtlmae, based in part on larval characters described and dis-
cussed by Mamaev (1976a), Doyen (1979), Young and Pollock (1991) and
Wharton (1979).
**"Fisch~r, 1822" was cited by Paulian (1949) for the family-group
name, but thts apparently refers to Fischer's description of the genus Pe-
dilus that year.
PYROCHROINAE Latreille, 1807a: 199
Dendroides (North America, Japan), Neopyrochroa (North America),
Pseudopyrochroa (Asia), Pyrochroa (Europe, Asia Minor), Schizotus
(Holarctic), etc.
*!he Neotropical genus Pogonoceromorphus, included in the Pyro-
chrom~e ?Y Vulcan? and Pe~eira (1972), almost certainly belongs to the
tenebnomd subfamtly Lagnmae. Blair (1914), Dettner (1984) Young ''I
(1975). ' I
PYROCHROIDAE incertae sedis:
AGNATHINAE Lacordaire, 1859: 531 (=Cononotinae)
Agnathus (Europe) and Cononotus (western North America).
*Mamaev (1976a) and Doyen (1979) considered Agnathus and Con-
onotus, respectively, to be close to Pedilus on the basis of larval features.
Pollock (1994) excluded Agnathinae from his pyrochroid cladogram al-
though he stated that this group possibly belongs in Pyrochroidae.
** Agnathinae has priority over Cononotinae LeConte (1862: 256) for
the subfamily name.
SALPINGIDAELeach, 1815: 106 (incl. Aegialitidae, Dacoderidae, Elacatidae,
Eurystethidae, Inopeplidae, Othniidae, Tretothoracidae)
*The Salpingidae is used here in a broad sense to include some taxa
usually P!aced in separate families. The relationships within the group and
between tt and other tenebrionoid families are not well understood. Iablo-
koff- Khnzorian (1985b), Lawrence (1982), Sasaji (1988) Spilman (1952
1954). ' '
OTHNIINAE LeConte, 1861: 102 (=Elacatinae)
Elacatis (=Othnius) (widespread).
*This is treated as a distinct family by many workers, and Crowson
(19;;) co~~ined it w~th .the following subfamily. Hayashi (1969).
.. Othnnnae has pnonty over Elacatinae Chapin (1923: 83), and in our
op1~10n the latter name has not been accepted widely enough to justify
retaming it over Othniinae via Art. 40b.
900
PROSTOMINIINAE Grouvelle, 1914: 152
Anepsicus, Eurycratus, Holosternus, lpsimorpha, Ocholissa, Prostominia,
Serrotibia, Szekessya, Trogocryptoides, Trogocryptus, etc.
*Prostominiines have been included in a number of different families,
including Cryptophagidae, Tenebrionidae, Prostomidae and Colydiidae.
Crowson (1955, 1968), Grouvelle (1914), Lawrence (1982), Sasaji (1988),
Scott (1926).
**The name Trogocryptinae of Crowson (1953: 51) and Lawrence
(1977: 43; 1980: 307) is apparently a nomen nudum.
AGLENINAE Hom, 1878: 573
Aglenus (widespread).
*This genus is sometimes included in the family Colydiidae, but ap-
pears to be closely related to prostominiines (Lawrence 1980).
INOPEPLINAE Grouvelle, 1908: 461
Aciphus (South America), Diagrypnodes (New Zealand), Inopeplus (wide-
spread), and Uruminopeplus (Japan).
*This is given family rank by many authors. A revision is now in prog-
ress by S.A. Slipinski and J.P. Lawrence, and preliminary studies indicate
that Inopeplus is a composite genus and that Diagrypnodes and Aciphus
should be transferred to another subfamily. Hayashi (1969), Olliff (1884),
Sasaji (1988), Sato and Hatta (1988), Waterhouse (1876).
SALPINGINAE Leach, 1815: 106 (incl. Lissodeminae)
Istrisia (Japan), Lanthanus (Neotropical), Lissodema, Notosalpingus (Aus-
tralia), Platamops (Neotropical), Poophylax (Falkland Is.), Rhinosimus,
Sphaeriestes, etc.
*Blair (1919).
AEGIALITINAE LeConte, 1862: 241 (=Eurystethinae)
Aegialites (=Eurystethes) (western North America, Siberia, Japan) and
Antarcticodomus (Campbell and Auckland Is.) .
*Spilman (1967).
**Eurystethinae Seidlitz (1916: 127) was an unnecessary replacement
name for Aegialitinae (Spilman 1954).
DACODERINAE LeConte, 1862: 216
Dacoderus (western North America) and Tretothorax (Australia).
*Watt (1967b).
ANTHICIDAE Latreille, 1819: 363
*This family is in need of major revision on a world level. Some of the
subfamilies listed below have been included in other families, such as
Pedilidae (see Pyrochroidae). Abdullah (1969a), Bonadona (1974), Buccia-
relli (1980), Uhmann (1976, 1978).
**Anthicus was not among the listed genera in Latreille's (1819, 1825)
treatment of this group, but the genus was not explicitly treated as a syn-
901
onym sow~ consider Latreille's name as available. Anthicidae has priority
over N~tox1dae Stephens (1829b: 254) for the family and nominotypical
subfamily names. "Notoxii" Sturm (1826: 38) is based on a misidentified
type genus, Notoxus of Fabricius (now Opilo, in Cleridae), not Notoxus
Geoffroy; the latter name has been conserved by the ICZN (1994a).
EURYGENIINAE LeConte, 1862: 264
Diacalla, Duboisius, Egestria, Egestriomima, Eurygenius, Ictystygna, Mas-
to remus, Pergetus, Rectocomus, Stereopalpus, etc.
*Some eurygeniine genera with externally closed procoxal cavities
(e . g., .lctistygna,.. Egestriomima) have been included in Lagriidae (Tene-
bnomdae: Lagnmae) by some workers (Armstrong 1948, Borchmann
1936). Champion (1916d).
LAGRIOIDINAE Abdullah and Abdullah, 1968: 73
Lagrioida (Australia, New Zealand, southern South America).
*Abdullah and Abdullah (1968), Costa et al. (in press).
AFREMINAE Levey, 1985: 420
Afremus (southern Africa).
*Levey (1985) compared this genus with members of the Eurygeniinae
and Copobaeninae.
MACRATRIINAE LeConte, 1862: 265
Macratria.
*Champion (1916b).
STEROPINAE Jacquelin du Val, 1863: 365
Steropes (Eurasia).
*Abdullah (1966b).
ISCHALIINAE Blair, 1920: 134
Ischalia (North America and Asia).
*Young (1985b) described the larva and moved this group to Anthi-
cidae from Pyrochroidae. Blair (1920), Nikitsky and Egorov (1992),
Paulus (1972b ).
COPOBAENINAE Abdullah, 1969a: 334
Copobaenus (Chile).
LEMODINAE Lawrence and Britton, 1991: 603 (Lawrence, 1977: 43, no-
men nudum)
Lemodes, Lemodinus and Trichananca (Australia), Lagriomorpha (New
Guinea, Moluccas), Cotes (New Zealand), etc.
*Lemodes was included in the family Pyrochroidae by most early
workers, and Blackburn (1899) noted the similarities between the genus
and Trichananca, suggesting that the latter be transferred from Oedeme-
ridae, where it was originally placed, to Pyrochroidae. Blair (1913) pointed
out the obvious affinities of both genera to Anthicidae, but placed them at
the end of Pyrochroidae in his volume of the Junk Catalogue (Blair 1928).
Although listed as "aberrant pyrochroids" by Lawrence (1977), Lemodes

902
and related taxa had been placed in Anthicidae by Britton (1970) and were
again by Lawrence and Britton (1991, 1994); Young (1978). transferred
Lagriomorpha to the same family. Although Lawrenc~ and Bntt~n (1991)
inadvertently validated the suprageneric name Lemodmae assummg a pre-
vious usage (see below), a case can be made for recognizing a group (di~­
tinct from Tomoderinae) to which these relatively large Southern Hemi-
sphere anthicids belong. Although they exhibit considerable variation,
lemodines may be distinguished from tomoderines by the subacute or nar-
rowly rounded intercoxal process on the first ventrite and by having the
sutures between the mesosternum and mesepisterna meet at a point on the
anterior edge of the meso thorax. In Tomoderinae, the abdominal intercoxal
process is truncate and the sutures dividing the mesosternum fr~m the
mesepisterna meet well behind the anterior edge, so that the mesep1sterna
are broadly contiguous anteriorly.
**The subfamily name Lemodinae was made available by Lawrence
and Britton (1991: 603, in larval key); Lawrence (1977: 43) and Young
(1985b: 205) used the subfamily name but gave no differential characters
to validate it.
TOMODERINAE Bonadona, 1961: 11
Elgonidium, Pseudotomoderus, Rimaderus, Tomoderus, etc.
*Bonadona (1961, 1978), Heberdey (1937), Werner (1958).
ANTHICINAE Latreille, 1819: 363 (=Notoxinae)
Acanthinus, Amblyderus, Anthicoxenus, Anthicus, Aulacoderus, Chilean-
thicus, Endomia, Formicomus, Jschyropalpus, Mecynotarsus, Microho-
ria, Notoxus, Omonadus, Sapintus, etc.
*Kitayama (1982), Werner (1966).
ADERIDAE Winkler, 1927: 831 (=Euglenidae, Euglenesidae, Hylophilidae,
Xylophilidae)
Aderus,Axylophilus, Cnopus, Emelinus, Escalerosia, Euglenes, Ganascus,
Gonzalosia, Hintonosia, Megaxenus, Olotelus, Phytobaenus, Pseuda-
nanca, Pseudanidorus, Pseudolotelus, Scraptogetus, Syzeton, Syzeto-
ninus, etc.
*Although the classification of Baguena Corella (1948) is based on t~e
world fauna, it is not particularly useful. The distinguishing features of h1s
Phytobaenini are not clear-cut, and the remaining tribes are separated b~
secondary sexual characters only. Werner (1990) divided the North Amen-
can species into two unnamed subfamilies based on aedeagal characters.
Baguena (1962), Casey (1895), Champion (1916a), Hayashi (1972), Law-
rence et al. (1990), Pic (1903, 1905), Watt (1987), Werner (1990).
**The names Xylophilidae Shuckard (1840: 47) and Hylophilidae Pic
(1900: 754) are based on preoccupied generic names and are thus unavail-
able. Euglenidae Seidlitz (1875: 106) has priority over Aderidae for the
903
fa~ily and nominotypical subfamily names, and also over Euglenidae
Stem, 1878 (based on Euglena) in Protozoa. However, in Opinion 1549
(ICZN 1989b) Euglenidae Seidlitz was emended to Euglenesidae to re-
move the family-name homonymy, and Aderidae was given precedence
over Euglenesidae whenever these two names are considered synonyms.
SCRAPTIIDAE Mulsant, 1856: 441/Gistel, 1856: 384 (incl. Anaspididae)
*Franciscolo (1964, 1972), Watt (1987).
SCRAPTIINAE Mulsant, 1856: 441/Gistel, 1856: 384
Biphida, Canifa, Nothotelus, Pectotoma, Phytilea, Scraptia, Tolmetes, Tro-
tomma, Trotommidea, Xylophilostenus, etc.
*Champion (1916c), Fairmaire (1900), Svacha (1995), Young (1976b).
ANASPIDINAE Mulsant, 1856: 389
Allopoda, Anaspimorda, Anaspis, Diclidia, Pentaria, etc.
*The anaspidine scraptiids were placed in Mordellidae in older classifi-
cations. Liljeblad (1945).
TENEBRIONOIDEA Incertae Sedis
Polypria (North to South America).
*This genus was provisionally included by Champion (1889) in a
broadly-defined Melandryidae and placed near Eurypus (now Mycteridae).
Spilman (1952) excluded Polypria from the mycterid group on the basis
of its genitalia (which he did not describe) and "temporarily" transferred
it to Melandryidae. Larvae are unknown.
Troctontus (Africa).
. *Silvestri (1920) described this genus on the basis of larvae occurring
m the nests of a termite (Microcerotermes), and Lawrence et al. (1990)
suggested that it could be an aderid related to Megaxenus. Grasse and
Lesperon (1938). I
i
v Superfamily CHRYSOMELOIDEA Latreille, 1802: 220
*Sv~cha and Dani~evsky (1987-89) place the first family below in a super- ,I
family Cerambycotdea; they also consider each of the first four cerambycid !I
subfamilies to be of family rank. Chen (1973, 1985), Iablokoff-Khnzorian
I.
(1985a), Kasap (1979), Lee (1993), Mann and Crowson (1983a, 1983c, 1983d, II
1984), Medvedev (1968, 1971), Napp (1994), Reid (1995), Schmitt (1992,
I
1994), Schmitt et al. (1982), Stork (1980), Suzuki (1988, 1989, 1994).
CERAMBYCIDAE Latreille, 1802: 211 (incl. Anoplodermatidae, Disteniidae,
Hypocephalidae, Oxypeltidae, Parandridae, Spondylidae, Vesperidae,
etc.)
*Crowson (1981) combined the first three of these subfamilies plus
Philinae in a family Disteniidae, but if this group is to be recognized at all,
904
905

then Mulsant' s Vesperidae is the oldest name. The firs tv four subfamilies and in litteris) has noted that the family-group name Spondylinae is preoc-
below were excluded from the family Cerambycidae by Svacha and Dani- cupied by Spondylidae Gray, 1826 (based on Spondylus in Mollusca), and
levsky (1987-89) because their larvae lack a gular or hypostomal area has proposed use of the more grammatically correct spelling Spondylid-
between the ventral mouthparts and the prothorax, a structure they consid- inae to avoid this homonymy.
ered to be a synapomorphy uniting the remaining subfamilies of Ceramby- APATOPHYSEINAE Lacordaire, 1869b: 234
cidae proper (see below). One possible synapomorphy uniting the first Apatophysis (Asia, North Africa).
four subfamilies is the very broad larval tentorial bridge, which was con- NECYDALINAE Latreille, 1825: 401
sidered by Svacha and Danilevsky to represent a "similar and independent Callisphyris, Hephaestion, Necydalis, Ulochaetes, etc.
response to general need for greater ridigity of the cranium." The place- LEPTURINAE Latreille, 1802: 218
ment of Philinae is uncertain since their larvae are unknown. Cherapanov Acmaeops, Centrodera, Gaurotes, Grammoptera, Leptura, Rhagium, Ste-
(1979-85), Duffy (1953, 1960), Fragoso (1985a, 1985b), Gahan (1906), nocorus, etc.
Gressitt et al. (1970), Linsley (1959, 1961-64), Linsley and Chefl!sak CERAMBYCINAE Latreille, 1802: 211
(1972-84), Manne and Giesbert (1994), Napp (1994), Saalas (1936), Sva-
Callidium, Cerambyx, Ceresium, Clytus, Molorchus, Phoracantha, Sten-
cha and Danilevsky (1987-89), Veiga Ferreira (1966-69), Villiers (1978), oderus, Strongylurus, etc.
Villiers and Quentin (1975). LAMIINAE Latreille, 1825: 401
VESPERINAE Mulsant, 1839: 214
Acanthocinus,Ancita, Dorcadion, Dorcaschema, Enicodes, Lamia, Mono-
Vesperus (southern Europe, northern Africa). chamus, Oncideres, Tetraopes, etc.
OXYPELTINAE Lacordaire, 1869a: 461 *Breuning (1950, 1958-69).
Cheloderus and Oxypeltus (southern South America).
*Duffy (1960), Fragoso (1985b). MEGALOPODIDAE Latreille, 1802: 227
DISTENIINAE J. Thomson, 1860b: 181
*Baccetti and Daccordi ( 198 8), Iablokoff-Khnzorian ( 1966-67), Jali vet
Cyrtonops (Asia), Distenia (North to South America, Asia), Noemia (Mad- (1957b, 1988), Kuschel and May (1990), Mann and Crowson (1983a,
agascar, Asia), etc.
198~c, 1984), Petitpierre (1988), Petitpierre et al. (1988), Reid (1995),
*Linsley (1961-64).
Santiago-Blay (1994), Seeno and Wilcox (1982), Steinhausen and Lochs
ANOPLODERMATINAE Guerin-Meneville, 1840: 276 (incl. Hypo- (1985).
cephalidae) PALOPHAGINAE Kuschel and May, 1990: 699
Anoploderma, Hypocephalus, Migdolus, etc. Cucujopsis and Palophagus (Australia).
*Dias (1984-87). *Kuschel and May (1990).
** Anoplodermatinae has priority over Hypocephalinae Blanchard ZEUGOPHORINAE Boving and Craighead, 1931: 63
(1845b: 135) for the subfamily name. Zeugophora (Europe, Asia, Africa, Australia), etc.
PHILINAE J. Thomson, 1860b: 297 *Jolivet (1957a), Lee (1990a), Reid (1989).
Doesus (Asia, Africa) and Philus (Asia). MEGALOPODINAE Latreille, 1802: 227
PARANDRINAE Blanchard, 1845b: 134
Agathomerus, Ateledera, Macrolopha, Mastostethus, Megalopus, Tem-
Erichsonia (Mexico, Central America) and Parandra (widespread). naspis, etc.
PRIONINAE Latreille, 1802: 212 *Monr6s (1955a), Yu and Yang (1994).
Archetypus, Callipogon, Derancistrus, Ergates, Eurynassa, Macrotoma,
Prionus, Stenodontes, Xixuthrus, etc. ORSODACNIDAE C.G. Thomson, 1859: 154
SPONDYLIDINAE Audinet-Serville, 1832: 122 (=Aseminae, Spondylinae)
*Iablokoff-Khnzorian (1966-67), Jolivet (1957b, 1988), Kusch~l and
Arhopalus, Asemum, Atimia, Megasemum, Michthisoma, Pectoctenus, Sa-
May (1990), Mann and Crowson (1983a, 1983c, 1983d, 1984), Monr6s
phanus, Spondylis, Tetropium, etc.
(1960a), Petitpierre et al. (1988), Reid (1995), Seeno and Wilcox (1982),
**Spondylidinae has priority over Aseminae J. Thomson, 1860b: 259 Steinhausen and Lochs (1985).
for the name of this subfamily, as Svacha and Danilevsky (1987-89) noted ORSODACNINAE C.G. Thomson, 1859: 154
in Part II of their work after using Aseminae in Part I. Silfverberg (1992 Orsodacne (Holarctic).
906
*Cox (1981), Jolivet (1957a), Mann and Crowson (1981).
AULACOSCELIDINAE Chapuis, 1874: 54
Aulacoscelis (North and Central America) and Janbechynea (New World).
*Monr6s (1953, 1954, 1960a).
CHRYSOMELIDAE Latreille, 1802: 220 (incl. Bruchidae, Sagridae, etc.)
*Two major changes have occurred in this family concept as a result
of broadly based phylogenetic studies: (1) the inclusion of the Bruchidae,
which was suggested many years ago by Boving and Craighead (1931)
and supported by Crowson (1960) and Monr6s (1960a), and (2) the re-
moval of those taxa presently included in Megalopodidae and Orsodacn-
idae above, a more recent move supported by Kuschel and May (1990)
and Reid (1995) and to some extent by Suzuki (1988, 1994). Baccetti and
Daccordi (1988), Crowson (1992), Deroe and Pasteels (1982), Gressitt and
Kimoto (1961-63), Hsiao (1994), Iablokoff-Khnzorian (1966-67), Jolivet
(1957b, 1959, 1988), Jolivet et al. (1988, 1994), Kasap and Crowson
(1985), Lopatin (1977), Mann and Crowson (1983a, 1983c, 1983d, 1984),
Monr6s (1960a), Petitpierre (1988), Petitpierre et al. (1988), Santiago-Blay
(1994), Seeno and Wilcox (1982), Steinhausen and Lochs (1985).
SAGRINAE Leach, 1815: 113
Atalasis (Argentina), Carpophagus, Diaphanops, Mecynodera, Polyoptilus,
etc. (Australia), Megamerus (Australia, Madagascar, Brazil), and Sagra
(Africa, Asia, Australia).
*Crowson (1946), Mann and Crowson (1992), Monr6s (1955b, 1958b).
BRUCHINAE Latreille, 1802: 192
Acanthoscelides, Amblycerus, Bruchidius, Bruchus, Callosobruchus, Cary-
oborus, Caryodon, Caryopemon, Eubaptus, Kytorhinus, Megacerus,
Pachymerus, Rhaebus, Zabrotes, etc.
*Borowiec (1987), Kingsolver and Pfaffenberger (1980), Nilsson and
Johnson (1993), Schmitt (1989), Southgate (1979), Udayagiri and Wadhi
(1989).
**Borowiec (1988) submitted a proposal to the ICZN for conservation
of Bruchus in the present sense (Linnaeus, 1767, not Muller, 1764); if
approved, this will also conserve the name Bruchidae Latreille.
DONACIINAE Kirby, 1837: 222
Donacia, Macroplea, Plateumaris, etc.
*Askevold (1990), Jolivet (1970), Lee (1991), Mann and Crowson
(1983b), Reid (1993).
CRIOCERINAE Latreille, 1804: 159
Crioceris, Lema, Lilioceris, Oulema, Stethopachys, etc.
*Schmitt (1985a, 1985b, 1988).
907
HISPINAE Gyllenhal, 1813: 448 (incl. Cassidinae)
Alurnus, Aproida, Arescus, Aspidimorpha, Cassida, Cephaloleia, Chale-
pus, Cryptonychus, Dorynota, Hispa, Notosacantha, Oediopalpa, Phys-
onota, Polychalca, Uroplata, etc.
*Borowiec (1995), Chen (1973), Samuelson (1989), Uhmann
(1957-64).
CHRYSOMELINAE Latreille, 1802: 220
Calligrapha, Calomela, Carystea, Chalcolampra, Chrysolina, Chrysomela,
Dicranosterna, Entomoscelis, Gonioctena, Leptinotarsa, Paropsis, Pha-
edon, Plagiodera, Timarcha, Zygogramma, etc.
*Kimoto (1962).
GALERUCINAE Latreille, 1802: 228 (incl. Alticinae)
Altica, Aulacophora, Blepharida, Chaetocnema, Crepidodera, Diabrotica,
Galeruca, Gale rue ella, Longitarsus, Microdonacia, Mono lepta, Oedio-
nychus, Oides, Orthaltica, Phyllotreta, Poneridia, Psylliodes, Pyrrh-
alta, etc.
*Furth (1988), Furth and Suzuki (1992), Jolivet (1968), Monr6s
(1958c), Reid (1992), Scherer (1988), Wilcox (1965, 1971-75).
LAMPROSOMATINAE Lacordaire, 1848: 559 (incl. Sphaerocharitinae)
Lamprosoma, Neochlamys, Oomorphus, Sphaerocharis, etc.
*Monr6s (1949, 1959, 1960b).
CRYPTOCEPHALINAE Gyllenhal, 1813: 582 (incl. Clytrinae, Chlamisinae)
Aporocera, Bahia, Cadmus, Chlamisus, Clytra, Cryptocephalus, Diachus,
Ditropidus, Fulcidax, Leasia, Megalostomis, Pachybrachis, Saxinis,
Stylosomus, etc.
*Kasap and Crowson (1976), Monr6s (1951), Reid (1990).
EUMOLPINAE Hope, 1840a: 162 (incl. Megascelidinae, Synetinae)
Adoxus, Chalcophana, Chrysochus, Chrysodina, Colaspis, Colaspoides,
Colasposoma, Edusella, Eumolpus, Fidia, Glyptoscelis, Mariamela,
Megascelis, Metachroma, Nodina, Nodonota, Odontionopa, Rhyparida,
Scelidonta, Spilopyra, Syneta, Thricolema, Typophorus, etc.
*Bechyne and Bechyne (1969), Kurcheva (1967), Lee (1990b), Mann
and Crowson (1981), Monr6s (1958a).
Superfamily CURCULIONOIDEA Latreille, 1802: 195
(=Rhynchophora)
*Three major works on the classification of curculionoids (Kuschel in press;
Thompson 1992; Zimmerman 1991, 1992, 1993, 1994a, 1994b) have appeared
within the last four years, and a fourth classification based on wing venation
is included in this volume (Zherikhin and Gratshev 1995). The classification
used here is based mainly on that of Kuschel (in press). Exceptions include the
recognition of Ithyceridae and Caridae as distinct families. Bright (1993), Cal-
der (1989, 1990b), Emden (1938, 1952), Kasap and Crowson (1977), May
908
(1993, 1994), Morimoto (1962a, 1962b, 1976), Morimoto and Lee (1987),
Zherikhin and Egorov (1990), Zherikhin and Gratshev (1994).
**A thorough review of family- and genus-group names in Curculionoidea
has been nearly completed (M.A. Alonso-Zarazaga, in litteris ).
NEMONYCHIDAE Bedel, 1882: 16 (=Rhinomaceridae)
*Although most workers since Crowson (1955) recognize this group as
a separate family, which is the sister group of the remaining Curcul-
ionoidea, Zherikhin and Gratshev (1995) suggest that Nemonychidae form
a monophyletic group with Anthribidae based on characters of the hind
wing. Crowson (1985) elevated Nemonychinae to family rank. Anderson
(1947), Kuschel (1959, 1983, 1989, 1993, 1994), Ter-Minassian (1984).
**Rhinomaceridae Schoenherr (1823: 1136) is the oldest name that has
been used for this family, but the type genus Rhinomacer and family-
group names based on it have been used in more than one sense (see under
Mycteridae). The ICZN (1994a) has effectively suppressed Rhinomacer in
all senses.
NEMONYCHINAE Bedel, 1882: 16
Nemonyx (Eurasia).
RHINORHYNCHINAE Voss, 1922: 17
Atopomacer (western U.S., Mexico), Basiliorhinus (Australia), Bunyaeus
(Australia), Mecomacer (Chile, Argentina), Notomacer (Australia, New
Caledonia), Pagomacer (Australia), Rhinorhynchus (New Zealand),
Rhynchitoplesius (Brazil), etc.
DOYDIRHYNCHINAE Pierce, 1916: 463 (=Cimberidinae, Neocimbe-
ridinae)
Cimberis (=Neocimberis) (North America, Eurasia), Doydirhynchus (Eu-
rope, Asia Minor, North Africa), Lecontellus (western North America),
Pityomacer (western North America), etc.
ANTHRIBIDAE Billberg, 1820: 39 (=Platystomidae, Platostomatidae, Platy-
rhinidae; incl. Bruchelidae, Urodontidae)
*Anderson (1947), Holloway (1982), Valentine (1960), Zimmerman
(1994a).
** Anthribidae has been given precedence over Choragidae for the fam-
ily name (ICZN 1994b).
URODONTINAE C.G. Thomson, 1859: 128 (=Bruchelinae)
Bruchela (=Urodon) (western Palearctic), Cercomorphus (southern Eu-
rope, North Africa), Urodontus (southern Africa), Urodoplatus (South
Africa), etc.
*This group was given family rank by Crowson (1984a) and Louw
(1993), but is usually considered to be a subfamily of Anthribidae. Prisnii
(1991) .
909
**The name Urodontinae (based on Urodon Schoenherr, not Urodontus
Louw) has priority over Bruchelinae Pierce (1916: 463) for the name of
the subfamily, and in our opinion the latter name is not sufficiently estab-
lished to justify conserving it via Art. 40b.
ANTHRIBINAE Billberg, 1820: 39 (incl. Brachytarsinae)
Allandrus, Anthribus, Basitropis, Discotenes, Eugonus, Euparius, Gymno-
gnathus, Ormiscus, Phloeotragus, Piesocorynus, Platyrhinus, Platys-
tomos, Tropideres, etc.
*Zherikhin and Gratshev (1995) consider Anthribinae to be para-
phyletic based on hind wing characters.
CHORAGINAE Kirby, 1819: 447 (=Araeocerinae)
Araecerus (=Araeocerus), Choragus, etc.
**Choraginae has priority over Araeocerinae Lacordaire (1866: 588),
based on the unjustified emendation Araeocerus Schoenherr, 1839 which
is also a preoccupied name.
BELIDAE Schoenherr, 1826: 73
*Each of the included subfamilies has been given family rank by most
authors, but May (1994) and Kuschel (in press) provided reasonably strong
support from both larvae and adults for a monophyletic group combining
these three taxa.
BELINAE Schoenherr, 1826: 73 (incl. Pachyurinae)
Agathinus, Belus, Dicordylus, Homalocerus, Pachyura, Rhinotia, etc.
*Kuschel (1959), Vanin (1976), Zimmerman (1994a).
OXYCORYNINAE Schoenherr, 1840: 581 (incl. Allocoryninae)
Afrocorynus, Hydnorobius, Metrioxena, Oxycorynus, Oxycraspedus, Rho-
palotria (=Allocorynus), etc.
*Kuschel (1959), Mufiiz and Barrera (1971), Voss (1957).
**Oxycoryninae has priority over Allocoryninae Sharp (1890: 45) for
the subfamily name.
AGLYCYDERINAE Wollaston, 1864: 384 (=Proterhininae)
Aglycyderes (Canary Is.), Aralius (=Platycephalus) (New Caledonia, New
Zealand), and Proterhinus (Hawaii, Fiji, Polynesia).
*Aralius is a new name proposed by Kuschel (1990) for the preoccu-
pied Platycephalus. Anderson (1941), Paulian (1944), Zimmerman and
Perrault (1989).
**Aglycyderinae has priority over Proterhininae Sharp (1899: 298) for
the name of the subfamily.
ATTELABIDAE Billberg, 1820: 39 (incl. Apoderidae, Pterocolidae, Rhyn-
chitidae)
910
*Voss (1965) placed the genus Camarotus in this family, but Vanin
and Reichardt (1974) transferred it to Curculionidae, as was suggested by
Crowson (1955) . Lee and Morimoto (1988a, 1988b), Vogt (1992) .
RHYNCHITINAE Gistel, 1856: 374 (=Pterocolinae)
Auletes, Auletobius, Byctiscus, Deporaus, Eugnamptus, Involvulus, Minu-
rus, Pterocolus, Rhynchites, etc.
*Zherikhin and Gratshev (1995) consider this taxon to be paraphyletic
based on hind wing characters.
**Rhynchitinae has priority over Pterocolinae Lacordaire (1866: 190).
ATTELABINAE Billberg, 1820: 39 (incl. Apoderinae)
Apoderus, Archolabus, Attelabus, Euops, Euscelus, Pilolabus, etc.
BRENTIDAE Billberg, 1820: 40 (=Brenthidae; incl. Apionidae, Cyladidae,
Eurhynchidae)
*Kuschel (1992, in press) recognized four subfamilies: Carinae (see
below), Cyladinae, Brentinae (including Eurhynchinae and Antliarhininae)
and Apioninae (including Nanophyinae) . Morimoto (1976), Sanborne
(1981), Zimmerman (1994a).
**According to Alonso-Zarazaga (in litteris) Brentus Fabricius (present
sense) is preoccupied by Brentus Panzer; the latter name is not in Neave
(1939-40). He has applied to ICZN for conservation of Brentus Fabricius
(Case 2887).
EURHYNCHINAE Lacordaire, 1863: 527
Aporhina, Eurhynchus, etc.
**Eurhynchinae and its type genus Eurhynchus Kirby were recently
conserved by Opinion 1352 (ICZN 1985c).
CYLADINAE Schoenherr, 1823: 1137
Cylas, Lispotherium, and Myrmacicelus.
BRENTINAE Billberg, 1820: 40
Amorphocephalus, Arhenodes, Brentus, Calodromus, Cerobates, Cordus,
Cyphagogus, Eupsalis, Nemocephalus, Taphroderes, Trachelizus,
Ulocerus, etc.
ANTLIARHININAE Schoenherr, 1823: 1137 (incl. Tanainae)
Antliarhis (=Antliarhinus, Antliarrhinus), Platymerus, and Tanaos.
**The name of the type genus is generally given asAntliarhinus, based
on Schoenherr' s (1823) misspelling of the type genus Antliarhis Billberg
(Alonso-Zarazaga, in litteris), or Antliarrhinus based on later misspellings
of Schoenherr's names.
NANOPHYINAE Gistel, 1856: 370
Nanophyes, etc.
*Alonso-Zarazaga (1989).
**Nanophyes has been conserved over the older name Nanodes by
Opinion 1526 (ICZN 1989a).
911
APIONINAE Schoenherr, 1823: 1136
Apion, Cybebus, Piezotrachelus, Podapion, etc.
*Alonso-Zarazaga (1990), Kissinger (1968) .
CARIDAE Thompson, 1992: 882 (Zimmerman, 1991: 112, not nomenclatu-
rally available)
Car and Carodes (Australia), Caenominurus and Chilecar (southern South
America).
*This group was included by most workers in the Attelabidae, and
Crowson (1955) considered species of Car to be the most primitive living
members of that family. Thompson (1992) moved Car to the family Be-
lidae and established a new subfamily Carinae. Kuschel (1992: 196) also
proposed a subfamily Carinae later that year, but included it in the family
Brentidae, adding two South American genera. Zimmerman (1994a: 499)
elevated the taxon to family rank, based in part on the unique larva de-
scribed by May (1994). May pointed out, however, that the larva has a
frontocranial bridge, shared only by Ithycerus and true Curculionidae.
Zherikhin and Gratshev (1995) also place the group in Belidae and suggest
a close relationship to the extinct Eccoptarthrini Arnoldi (Arnoldi et al.
1977: 169); combining these two groups would necessitate adopting the
latter, senior name for the group.
ITHYCERIDAE Schoenherr, 1823: 1136
Ithycerus (North America) .
*May (1993) included this family in the subfamily Brachycerinae of
the Curculionidae, based on the presence of a frontocranial bridge (sepa-
rating the frontal sutures from the mandibular membrane) on the larval
head capsule, presence of mandibular setae in the pupa, and the habit of
ovipositing in the soil. The larva resembles those of Brentidae and some
other more primitive weevil families in the presence of well-developed
thoracic legs, three stemmata, four labral setae and four Malpighian tu-
bules. Sanborne (1981) gave evidence supporting a relationship to Brent-
idae, and Zherikhin and Gratsev (1995) noted similarities to Belidae in the
hind wing. Lawrence (1991), Morimoto (1976).
CURCULIONIDAELatreille, 1802: 195 (incl. Calendridae, Cossonidae, Rhyn-
chophoridae, Scolytidae, etc.)
*The classification used here is that of Kuschel (in press), except for
the placement of Ithycerus in a separate family. Zherikhin and Gratshev
(1995) split the group into three families: Brachyceridae, Barididae and
Curculionidae. Other subfamily arrangements may be found in Thompson
(1992) and Zimmerman (1991, 1992, 1993, 1994a, 1994b). Kissinger
912
(1964), O'Brien and Wibmer (1982, 1984), Wibmer and O'Brien (1986,
1989).
BRACHYCERINAE Billberg, 1820: 39 (incl. Alophinae, Amycterinae, Ater-
pinae, Brachyderinae, Byrsopinae, Cryptolarynginae, Cylindrorhininae,
Desmidophorinae, Diabathrariinae, Dinomorphinae, Ectemnorhininae, Enti-
minae, Eremninae, Gonipterinae, Hipporhininae, Hyperinae, Leptopiinae,
Otiorhynchinae, Pachyrhynchinae, Rhadinosominae, Rhyparosominae,
Rhytirhininae, Somatodinae, Tanyrhynchinae, Thecesteminae, Ulomas-
cinae, Viticinae)
Alophus, Amycterus, Brachycerus, Dinomorphus, Entimus, Gonipterus,
Leptopius, Mandalotus, Myllocerus, Naupactus, Otiorhynchus, Panto-
morus, Polydrosus, Rhadinosomus, Sitona, Tanymecus, Tanyrhynchus,
etc.
CURCULIONINAE Latreille, 1802: 195 (incl. Anoplinae, Anthonominae,
Bagoinae, Baridinae, Camarotinae, Campyloscelinae, Ceratopodinae, Ceu-
torhynchinae, Cioninae, Cleoninae, Cryptorhynchinae, Erirhininae, Eu-
gnominae, Gymnetrinae, Lixinae, Magdalininae, Molytinae, Nerthopinae,
Otidocephalinae, Prionomerinae, Pyropinae, Raymondionyminae, Rhyn-
chaeninae, Tachygoninae, Trigonocolinae, Tychiinae, Zygopinae)
Anthonomus, Bagous, Baris, Campyloscelis, Ceutorhynchus, Cleonus, Co-
notrachelus, Cryptorhynchus, Curculio, Gymnetron, Hylobius, Lixus,
Magdalis, Rhynchaenus, Tachygonus, Tychius, Zygops, etc.
*Kuschel (1987), Lyal (1993), Papp (1979).
DRYOPHTHORINAE Schoenherr, 1825: 588 (=Rhynchophorinae, Calan-
drinae, Calendrinae)
Cryptoderma, Dryophthorus, Orthognathus, Rhynchophorus, Sipalinus
(=Sipalus), Sitophilus, Sphenophorus, Stromboscerus, etc.
*Zimmerman (1993).
**References to early use of Rhynchophorinae by Schellenberg/Clai-
rville, 1798 ([not seen], see Watt 1975), Latreille (1804, 1807, 1817),
Leach (1815), and Billberg (1820) refer to use of "Rhynchophori" or a
variant term for all weevils, and not (apparently) to a genus-based name;
even if the cited authors intended this term as a genus-based name, none
of them treated Rhynchophorus as a valid genus so the group name is un-
available. Schoenherr (1823, 1825, 1826) treatedRhynchophorus as a valid
genus but did not use a group name based on it until 1833; Rhyncho-
phorinae should therefore be attributed to Schoenherr (1833: 26) as noted
by Thompson (1992). The names Calandra and Calandrinae Billberg
( 1820: 40) were rejected in Opinion 572 (ICZN 1959), leaving Dryophtho-
rinae Schoenherr (1825: 588) as the oldest available name for this group
(Zherikhin and Egorov 1990, Thompson 1992, Alonso-Zarazaga in litte-
ris). According to Alonso-Zarazaga (in litteris) the original spellings Rhyn-
chophorus and Rynchophorus were used by Herbst.
913
COSSONINAE Schoenherr, 1825: 587
Araucarius, Cossonus, Hexarthrum, Pentarthruni, Protoplatypus, Rhyn-
colus, Stenotrupis, etc.
*Several workers (Lekander 1968, Viedma 1963) have emphasized the
difficulties in separating cossonine larvae from those of scolytines, and
Kuschel (1966) and May ( 1967) have shown that members of the cosson-
ine tribe Araucariini are somewhat intermediate between the two groups
in both adult and larval characters.
SCOLYTINAE Latreille, 1804: 157 (incl. Hylesininae, Ipinae, Tomicinae,
Coptonotinae, etc.)
Acacicis, Bothrosternus, Cactopinus, Carphodicticus, Coccotrypes, Copto-
notus, Corthylus, Cryphalus, Crypturgus, Dendroctonus, Diamerus,
Dryocoetes, Hylastes, Hylesinus, Hylurgops, Hyorrhynchus, Hypo-
horus, Hypothenemus, Ips, Micracis, Phloeosinus, Phloeotribus,
Phrixosoma, Polygraphus, Scolytodes, Scolytoplatypus, Scolytus, Tomi-
cus, Xyleborus, Xyloctonus, etc.
*This and the following subfamily were treated as separate families
until Crowson (1955, 1960) included them as subfamilies of Curculion-
idae. Wood (1973) presented arguments based on adult features for consid-
ering scoltyids and platypodids as a separate lineage from curculionids
proper. His agruments have been criticized by Kuschel (in press), who also
pointed out that Wood's "most primitive platypodid", Protoplatypus, be-
longs in the Cossoninae, and his "most primitive hylesinine scolytid", Pro-
tohylastes, is related to the cryptorhynchine genus Psepholax (Curcu-
lioninae above). Bright (1976), Wood (1978, 1982, 1986), Wood and
Bright (1987-93).
**The older name Ipidae of Latreille (1802: 131) (not 1807 as given
by Watt (1975) and Silfverberg (1992)) was based on Ips of Olivier (now
what?), not Ips DeGeer of Scolytinae; Ipini Bedel (1888: 395) is the first
correct use of a family-group name based on the latter genus (Wood
1986). This problem needs more work; Ipinae in Nitidulidae is yet another
concept.
PLATYPODINAE Shuckard, 1840: 64 (incl. Chapuisiinae, Crossotarsinae,
Diaporinae, Periommatinae, Platytarsilinae)
Austroplatypus, Crossotarsus, Diapus, Mecopelmus, Notoplatypus, Per-
iommatus, Platypus, Platytarsulus, Schedlarius (=Chapuisia), Tesse-
rocerus, etc.
*Browne (1972), Kuschel (1994b), Schedl (1972), Wood and Bright
(1987-93).
914
References
Abdullah, M. 1964a. A revision of the genus Pilipalpus (Coleoptera, Anthicidae: Pedilinae).
Beitriige zur Entomologie, 14: 3-9.
Abdullah, M. 1964b. Protomeloe argentinensis, a new genus and species of Meloidae (Coleo-
ptera), with remarks on the significance of cantharidin and the phylogeny of the families
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I,,,
1
~~ 1
•! I

i
·~
1006
INDEX
[note: this index is not a part of original publication; it was only included in reprints]
Zimmerman, E.C. 1994b. Australian Weevils (Coleoptera: Curculionoidea). Vol. 2, Brentidae,
Eurhynchidae, Apionidae, and a Chapter on Immature States by Brenda May. CSIRO, East
Melbourne, x + 755 pp.
Zimmerman, E.C. and Perrault, G.H. 1989. Aglycyderidae of the Society Islands (Coleoptera:
Aata 842 Acropteron 893
Curculionoidea). Bishop Museum Occasional Papers, 29: 151-173. .
Ababa 869 Acropteroxys 879
Zunino, M. 1983. Essai preliminaire sur !'evolution des armures genetales des Sc~r~bae1~ae,
Abdera 888 Acrotrichinae 782, 819
par rapport a Ia taxonomie du groupe et a !'evolution du comportement de mdlficatwn.
Ablabera 835 Acrotrichis 818, 819
Bulletin de la Societe Entomologique de France, 88: 531-542.
Ablepton 822 Acruliodema 824
Zunino, M. 1984a. Sistematica generica dei Geotrupinae (Coleoptera, Scarabaeoidea: Geotru-
Abraeinae 782, 816 Actenodes 842
pidae), filogenesi della sottofamiglia e considerazioni biogeografiche. Bollettino del Museo
Abraeomorphus 817 Actidium 818
Regionale di Scienze Naturali, Torino, 2: 9-162. . . . Abraeus 816
Zunino, M. 1984b. Analisi sistematica e zoogeografica della sottofam1gha Tauroc~rastma~ Aculagnathidae 881
Abromus 881 Aculagnathus 881
Germain (Coleoptera, Scarabaeoidea: Geotrupidae). Bollettino del Museo Regwnale d1
Abstrulia 888 Acupalpus 812
Scienze Naturali, Torino, 2: 445-464.
Acacicis 913 Acylomus 877
Zwick, P. 1979. Contributions to the Knowledge of Australian Cholevidae (Catoptidae auct.:
Acalanthis 869 Adalia 885
Coleoptera). Australian Journal of Zoology (Supplementary Series), 70: 1-56._
Acalyptomerinae 783, 839 Adelium 892
Zwick, P. 1981. Die Jugendstadien des Kafer Necrophilus subterraneus (Col. Silph. Agyr-
Acalyptomerus 839 Adelocera 854
tinae). Beitriige zur Naturkunde Osthessens, 17: 133-140.
Acanthinus 902 Adelocerinae 854
Acanthoceridae 794, 833, 834 Adelonia 892
Acanthocerus 833 Adelotopus 812, 813
Acanthocinus 905 Adelphydraena 818
Acanthocnemidae 785, 867, 870 Ademosynidae 800
Acanthocneminae 871 Adephaga 782, 789, 799, 800, 803-807,
Acanthocnemis 871 809, 810, 815, 915, 917, 919-921,
Acanthocnemus 867, 868, 870, 928 923, 926, 935, 939, 954, 956, 966,
Acanthocnemus nigricans 868, 870, 974, 975, 982, 992, 1000
928 Aderidae 787, 796, 902, 903, 917, 947,
Acanthoscelides 906 952, 962, 1001
Achenium 829 Aderus 902
Acherusia 842 Adesmia 893
Acidocerus 816 Adimeridae 890
Acidota 824 Adimerus 890
Acilius 809 Adinolepis 803
Aciphus 900 Adistemia 886
Aclopidae 834 Adonia 885
Aclopinae 783, 835, 915 Adoretus 836
Aclopus 835 Adoxus 907
Aclypea 822 Adrastinae 854
Acmaeodera 842 Adrastus 854
Acmaeoderinae 842, 843 Aegialia 834, 930
Acmaeoderoides 842 Aegialiinae 834
Acmaeops 905 Aegialites 900
Acontosceles 845 Aegialitidae 899
Acritomorphus 816 Aegialitinae 787, 900, 993
Acritus 816 Aegidium 835
Acropis 890 Aeletes 816
Acrops 869 Aenigmaticum 885, 976
[2] [3]

Aephnidius 812 Alaus 854 Ametor 816 Anelpistus 895, 967

Aesalinae 783, 831, 950, 968 Aleochara 827 Ammophorus 893 Anemadinae 821, 943
Aesalus 831, 916 Aleocharinae 783, 823, 826, 916, 917, Amorphocephalus 910 Anemadus 821
Aethina 874, 957 940,942,949,958,971,989,994 Ampedus 854 Anepiini 824
Afreminae 787, 901, 964 Alesia 885 Amphicoma 834 Anepius 824
Afremus 901 Alexia 882 Amphicrossus 874, 977 Anepsicus 900
Afroboganium 875 Alexiidae 786, 795, 882 Amphicyrta 844 Anepsius 893
Afrocorynus 909 Alfieriella 878, 982 Amphicyrtinae 783, 844 Anischia 85 1, 855
Afroeubria 846 Alfieriellinae 878 Amphiopinae 816 Anischiinae 784, 855 l'
Afropelates 820 Algophilus 807 Amphiops 816 Anisochaeta 810 i'
Afrorthocerus 890 Allandrus 909 Amphix 883 Anisodacty lus 812 I.
,J
Agabetes 808 Allecula 893 Amphizoa 808, 921 Anisolinus 830 l,j·
,,
Alleculidae 892 Amphizoidae 782, 806, 808, 921, 926, Anisomeria 809 I
Agabetini 808
Agabus 809 Alleculinae 787, 796, 892, 893 941, 956, 998, 1005 Anisomeristes 885
Agaeocera 842 Allidiostoma 835 Amplectopus 839, 840 Anisomerus 855
Agaocephala 836 Allidiostomatinae 794, 835 Amycterinae 912, 1005 Anisosphaeridae 821
Agaporomorphus 808 Allidiostominae 794, 835 Amycterus 912 Anisosticta 885
Agapythinae 785, 875 Allocoryninae 909 Amydetes 859 Anisotoma 820, 1001
Agapytho 875 Allocorynus 909 Amydetinae 784, 859 Anisotomidae 794, 819, 820
Agaricophilinae 884 Alloparmulus 885 Amydropa 878 Anisotominae 820
Agaricophilus 884, 967 Allopoda 903 Amystrops 874 Anisotria 898, 1004
Agasma 895 Allopogonia 850 Anacaena 816 Anobiidae 785, 792, 794, 861, 863,
Agathidium 820 Alloproteinus 824 Anadastus 879 864, 924, 939, 980, 999, 1001,
I
Agathinus 909 Alloscelus 834 Anaedus 892 1002 I
Alloxantha 895 Anobiinae 785, 865 ,i,J
Agathomerus 905 Anaides 834
Agleninae 787, 900 Alobates 894 Anamorphinae 786, 795, 884 Anobium 865
Aglenus 900 Alophinae 912, 937 Anamorphus 884 Anomala 836
Aglycyderes 909 Alophus 912 Anapleus 817 Anommatidae 881, 926, 934
Aglycyderinae 788, 909 Alphitobius 893 Anaplopus 897, 914, 980 Anommatinae 786, 881
Aglymbus 808 Alphitophagus 894 Anaspasis 853 Anommatus 881, 934
Aglyptinus 820 Althaesia 880 Anaspididae 903 Anoplinae 912
Agnathinae 787, 796, 899 Altica 907 Anaspidinae 787, 889, 903, 941 Anoploderma 904
Agnathus 899, 967 Alticinae 907, 942, 987 Anaspimorda 903 Anoplodermatidae 903
Agonica 812 Alumus 907 Anaspis 903 Anoplodermatinae 787, 904, 935
Agonum 812 Alvarenganiella 865 Anatis 885 Anoplognathus 836
Agra 812 Alzadaesthetus 829 Anchastus 854 Anorus 841
Agrilinae 783, 843 Amara 812 Anchicera 878 Anotylus 828, 946
Agrilus 843, 1000 Amarotypus 811 Ancholaemus 889 Antarcticodomus 900
Agriotes 854 Amartus 873 Anchorius 880 Antarcticonomus 811
Agrypninae 784, 794, 854, 854, 947 Amarygmus 893 Anchycteis 846, 847 Anthaxia 842
Agrypnus 854 Amauronia 872 Anchytarsinae 784, 846 Antherophagus 878
Agyrtes 819 Amaurops 825 Anchytarsus 846, 847, 995 Anthia 812
Agyrtidae 782, 817, 819, 822, 915, Amblycerus 906 Ancistria 876 Anthicidae 787, 899-902, 914, 923,
973,979,986 Amblycheila 811 Ancita 905 925, 928, 931, 947, 958, 964, 980,
Agyrtodes 820 Amblyderus 902 Ancylopus 883 998, 1001, 1004
Agyrtodinae 820 Amblyopininae 830 Ancyrona 868 Anthicinae 787, 902
Agyrtodini 820 Amblyopinus 830 Ancyronyx 844 Anthicoxenus 896, 902
Ahasverus 876 Amblystema 842 Andotypus 816 Anthicus 900, 902
Airora 869 Amblystomus 812 Anelastes 851 Anthobium 824
Akis 893 Amecocerus 872 Anelastidius 852 Anthonorninae 912
.I
i
.I
[4] [5]
Anthonomus 912 Apoderinae 910 Arthopus 891 Atopinae 841
Anthophagus 824 Apoderus 910 Arthrobrachus 871 Atopomacer 908
Anthracalaus 854, 926 Apoleon 863 Arthrolipinae 885 Atractoceridae 866
Anthreninae 794, 863 Apoleoninae 863 Arthrolips 885 Atractocerus 800, 857, 866, 941, 957,
Anthrenocerus 863 Aporhina 910 Arthromacra 892 989
Anthrenus 863 Aporocera 907 Arthropterites 810 Atrecus 830
Anthribidae 788, 908, 915, 950, 952, Apotomus 812 Arthropterus 810 Attageninae 785, 863
998, 1005 Aproida 907 Ascetoderes 881 Attagenus 863
Anthribinae 788, 909 Aprostoma 890 Ascioplaga 803 Attalus 872
Anthriboclerus 871 Apsectus 863 Asclera 895 Attelabidae 788, 909, 911, 963, 999,
Anthribus 909 Apteraliplus 807 Ascydmus 822 1000, 1005
Anthypna 834 Apteroloma 819 Aseminae 796, 904 Attelabinae 788, 910, 963
Antibothrus 881 Apterotheca 894 Asemobius 829, 948 Attelabus 910
Antixoon 868 Aptinus 813 Asemum 904 Aubehydrinae 782, 808, 809, 993
Antliarhininae 788, 910 Araecerus 909 Asida 893 Aubehydrus 809
Antliarhinus 91 0 Araeocerinae 909 Asilis 860 Auglyes 845
Antliarhis 910 Araeocerus 909 Asiocoleidae 800 Aulacochilus 879
Antliarrhinus 910 Araeopidiinae 784, 847 Aspathenes 890 Aulacocyclinae 783, 832
Antroherpon 821 Araeopidius 847 Aspidimerus 885 Aulacocyclus 832
Apalus 896 Araeoschizus 893 Aspidimorpha 907 Aulacoderus 902
Apate 863 Aralius 909 Aspidiphoridae 795, 872 Aulacophora 907
Apatetica 827, 828, 966 Araucariini 913, 968 Aspidiphorinae 872, 873 Aulacoscelidinae 787, 906
Apateticinae 783, 822, 827, 828 Araucarius 913 Aspidiphorus 872, 873 Aulacoscelis 906
Apatophyseinae 787, 905 Archaeogastra 800 Aspidocha 885 Auletes 910
Apatophysis 905 Archaeoglenes 893 Astenus 829 Auletobius 910
Apeosina 838 Archecoleoptera 800 Astraeus 842 Aulonium 890
Aphaenocephalidae 877, 882 Archeocrypticidae 962, 998 Astraptor illuminator 858, 972 Aulonocneminae 834, 994
Aphaenostemminae 824 Archeocrypticus 887 Astylus 871 Aulonocnemis 834
Aphaenostemmus 824 Archetypus 904 Ataenius 834 Aulonogyrus 807
Aphanocephalidae 877, 882 Archipines 883 Atalasis 906, 970 Aulonosoma 876, 926
Aphanocephalinae 786, 882 Archolabus 910 Atanygnathus 830 Aulonothroscus 852
Aphanocephalus 882, 942, 945 Archostemata 782, 799, 800, 802, 803, Ateledera 905 Australenneboeus 887
Aphileus 854 805,935,943,954,973,976,980, Ateliinae 784, 857 Australiodes 868
Aphodiinae 783, 834, 930, 944, 951, 999 Atelius 857 Australymexylon 866
954, 968, 994, 1003 Arescus 907 Aterpinae 912 Austrelater 853, 927
Aphodius 834 Arhenodes 91 0 Ateuchus 834 Austroesthetus 829
Apicalae 800 Arhina 874 Athertonium 875 Austrolichas 847
Apion 911, 958 Arhopalus 904 Atheta 827, 1004 Austrolimnius 844, 949
Apionidae 910, 915, 971, 1006 Aridius 886 Athoinae 794, 854 Austroplatypus 913
Apioninae 788, 910, 911, 915, 958 Arisus 852 Atholus 817 Austrorhysus 824
Aplastinae 852 Arpidiphorus 873 Athous 854, 984 Autocrates 897
Aplastus 853 Arrhaphipterus 841 Athyreinae 832 Automolus 835
Aplocneminae 871, 978 Arrhipis 852 Athyreus 832 Axinotarsus 872
Aplocnemus 871 Arrowinus 830 Atimia 904 Axiocerylon 881
Aploderus 828 Artematopidae 794, 837, 849, 850, 930 Atomaria 878 Axylophilus 902
Aploglossa 847 Artematopodidae 784, 794, 837, 840, Atomariinae 786, 878 · Azya 884
Aplog1ossinae 784, 847 848-851, 861, 962 Atomarops 879 Babia 907
Aplothorax 811 Artematopodinae 784, 850 Atopa 840 Bacanius 817
Apocellus 828 Artematopodoidea 849-851 Atopida 839 Bactridium 874
Apoderidae 909 Artematopus 850 Atopidae 840 Baeocera 828
[6] [7]
Baeoceridium 828 Blatchleya 860 Brachygluta 825 Buprestinae 783, 842
Bagoinae 912 Bledius 828, 948 Brachylampis 859 Buprestis 842, 982
Bagous 912 Blepharida 907 Brachyleptus 873 Buprestoidea 783, 836, 837, 842
Balginae 853, 943 Blethisa 811 Brachymera 861 Byctiscus 910
Balgus 853 Blitophaga 822 Brachypeplus 874 Byrrhidae 783, 815, 836, 837, 843,
Balteifera 810, 813 Boganiidae 785, 875, 933, 938, 990 Brachypsectra 850, 922 936, 955, 978, 1000
Baptolinus 830 Boganiinae 785, 875 Brachypsectridae 784, 837, 848-851 Byrrhinae 783, 844
Barididae 911 Boganium 875 Brachypteridae 785, 795, 873 Byrrhinus 845, 1004
Baridinae 912 Bolax 836 Brachypterinae 873 Byrrhoidea 783, 836, 837, 843, 848,
Baris 912 Bolbelasmus 832 Brachypterolus 873 862, 936, 937
Barypus 812 Bolboceras 832 Brachypterus 873 Byrrhopsis 839
Basiliorhinus 908 Bolbocerasoma 832 Brachys 843 Byrrhus 844
Basitropis 909 Bolboceratinae 783, 832, 925 Brachysphaenus 880 Byrrocryptus 846, 847
Bathyscia 821 Bolbocerinae 832 Brachytarsinae 909 Byrsopinae 912
Bathysciinae 821, 822, 927, 945, 953, Bolbomorphus 883 Bradymerus 894 Bystus 884
960 Bolerus 879 Brasilucanus 832 Bythininae 825
Bathysciini 821 Bolitobius 826 Brathinidae 822, 824 Bythinoplectus 825
Bathysciola 821 Bolitochara 827 Brathinus 824, 996 Bythinus 825
Batrisinae 825 Bolitotherus 893 Brenthidae 910, 988 Byturellus 880
Batrisodes 825 Borborophorus 816 Brentidae 788, 910, 911, 971, 1006 Byturidae 786, 867, 880, 886, 914,
Batrisus 825 Boreaphilus 824 Brentinae 788, 910 993, 994
Belidae 788, 909, 911, 959, 998 Boridae 787, 897, 898, 962, 994, 1004 Brentus 910 Byturinae 786, 880
Belinae 788, 909 Borinae 787, 897, 898 Brontes 876 Byturodes 880
Belohina 833 Borolinus 828 Brontinae 785, 795, 876 Byturus 880
Belohinidae 783, 833, 978 Boros 897, 994 Brontolaemus 877 Cactopinus 913
Belonuchus 830 Bostrichidae 785, 861, 863, 864, 915, Brontopriscus 876 Cadmus 907
Belopus 892 940, 988 Broscus 812 Caenia 856
Belotus 860 Bostrichiformia 784, 785, 813, 838, Brounia 847, 848 Caenocara 865
Belus 909 861, 865, 866, 949, 953 Brounia Thoracica 848 Caenominurus 911
Bembidion 812 Bostrichinae 785, 863 Brouniphylax 891 Caenoscelis 878
Berendtimiridae 856, 1002 Bostrichoidea 784, 862 Bruchela 908, 933, 981 Cafius 830
Bergininae 786, 887 Bostrichus 863 Bruchelidae 908, 981 Cainogenion 812
Berginus 887 Bostrychidae 863, 932, 958 Bruchelinae 908, 909 Calandra 912, 952
Berosinae 816 Bostrychopsis 863 Bruchidae 896, 906, 923, 956, 957, Calandrinae 912
Berosus 816 Bothrideres 881 975,987,993,998 Calathus 812
Bicava 886 Bothrideridae 786, 867, 872, 881, 926, Bruchidius 906 Calendridae 911
Bicellonycha 859 931, 976, 992, 995 Bruchinae 787, 906 Calendrinae 912
Bidessus 809 Bothriderinae 786, 881, 992 Bruchus 906, 923 Calitinae 785, 868, 869
Biophytus 877 Bothriophorinae 845 Brumus 885 Calitys 867-869
Biphida 903 Bothrostemus 913 Bryaxis 825 Callidircaea 888
Biphyllidae 786, 880, 886, 914, 944 Botrodus 881 Brychius 807 Callidium 905
Biphyllus 880 Bourgeoisia 859 Brycopia 892 Calligrapha 907
Bironium 828 Brachelytra 823 Bryoporus 826 Callipogon 904
Bisaya 838 Brachiacantha 884 Bubastes 842 Callirhipidae 784, 794, 837, 841, 843,
Bispinatus 880 Brachininae 782, 813, 935 Bucolus 884 848, 849
Bitoma 890, 933 Brachinus 813 Bulaea 885 Callirhipis 848, 849
Blackbumiella 870 Brachyceridae 911 Bunyaeus 908 Callismilax 893
Blackbumium 832 Brachycerinae 788, 911, 912 Buprestidae 783, 837, 842, 861, 920, Callisphyris 905
Blaps 893 Brachycerus 912 922, 924, 929, 933, 943, 950, 973, Callistus 812
Blapstinus 893 Brachyderinae 912 998, 1000 Callosobruchus 906
[8] [9]
Calochrominae 784, 857 Carcinops 817 Cavicoxumidae 854 Ceratopodinae 912
Calochromus 857 Cardiophorinae 784, 854 Cavognatha 878 Ceratotrupes 833
Calodromus 910 Cardiophorus 854, 855 Cavognathidae 786, 878, 932, 990, Cercomorphus 908
Calomela 907 Cardiorhinus 854 1000 Cercus 873
Calonecrinae 785, 874 Cardiothorax 892 Cebrio 853 Cercyon 816
Calonecrus 874, 957 Carenum 812, 971 Cebriognathinae 852 Ceresium 905
Calophaena 812 Caridae 788, 907, 911 Cebriognathus 853 Ceriselma 896
Calopodinae 787, 895 Carinisphindus 873, 969 Cebrionidae 792, 837, 852, 978, 992 Cerobates 910
Calopteron 856 Carinodula 884 Cebrioninae 784, 852, 855 Cerocoma 896
Calopus 895 Carinophloeus 877, 933 Cebrionoidea 794, 849 Cerocosminae 864
Calosoma 811 Carodes 911 Cebriorhipis 853 Cerocosmus 864, 932
Calymmaderus 865 Carpelimus 828 Celetes 856 Ceroglossus 811
Calyptomeridae 839 Carphodicticus 913 Celina 809 Cerophytidae 784, 837, 851, 855, 944,
Calyptomerinae 783, 839 Carphuridae 871, 939 Celladonia 848 967, 992
Calyptomerus 803, 839, 937 Carphurus 872 Cenophengus 858 Cerophytum 851, 855, 937, 967
Camarotinae 912, 998, 1000 Carpophagus 906 Centrodera 905 Cerophytum Elateroides 851, 967
Camarotus 910 Carpophilinae 785, 874, 957 Centronopus 893 Ceroplastus 855
Camiaridae 819, 954 Carpophilus 874 Cephalobyrrhinae 784, 845, 1004 Ceruchus 831
Camiarinae 782, 820 Cartodere 886 Cephalobyrrhus 845 Cerylidae 881, 926
Camiarus 820 Caryoborus 906 Cephaloidae 796, 894, 895, 914, 916, Cerylon 881, 882, 991
Camioleum 824 Caryodon 906 947, 974 Cerylonidae 786, 881, 890, 934, 963,
Campsosternus 853 Caryopemon 906 Cephaloinae 787, 895 967, 974, 990-992
Camptodes 874 Carystea 907 Cephaloleia 907· Ceryloninae 786, 881
Campylinae 854 Caserus 886, 934 Cephaloon 895 Cetonia 836
Campy loscelinae 912 Casopus 864 Cephalophaninae 786, 882 Cetoniidae 834, 959
Campyloscelis 912 Cassida 907 Cephalophanus 882 Cetoniinae 783, 835, 836, 968, 969
Campyloxeninae 854, 930 Cassidinae 907 Cephaloplectidae 818 Ceutorhynchinae 912
Campyloxenus 854 Cassidoloma 882 Cephaloplectinae 782, 794, 819, 989 Ceutorhynchus 912
Campylus 854 Castiarina 842 Cephaloplectus 819 Chaetarthria 816
Canifa 903 Catana 884 Cephaloscymnus 884 Chaetillus 892
Cantharidae 784, 793, 837, 846, 859, Cateretes 873 Cephalotyphlus 829 Chaetocanthinae 783, 833
860, 895, 924, 935, 964, 967, Cateretidae 795, 873 Cephennium 822 Chaetocanthus 833
1001, 1003 Cathartus 876 Ceracis 887 Chaetocnema 907
Cantharinae 784, 860 Catiniidae 800, 804 Ceracupes 832 Chaetonyx 835
Cantharis 860, 895 Catogenidae 876 Cerallus 871 Chaetophora 844
Cantharoidea 837, 841, 849, 850, 915, Catogenus 876, 991 Cerambycidae 787, 903, 904, 929, 936, Chaetosoma 869
932,967,969,978,981,1002 Catopidae 819, 945, 954, 960, 996 941,943,965,970,973,999 Chaetosomatidae 785, 869, 936, 969
Canthon 834 Catopinae 794, 821, 979 Cerambycinae 787, 905 Chaetosomodes 869
Canthydrus 808 Catopius 887 Cerambyciscapha 828 Chalchas 871
Car 804, 850, 871, 876, 898, 911 Catopocerinae 782, 820 Cerambycoidea 903, 996 Chalcionellus 817, 982
Carabhydrus 809 Catopocerus 820, 979 Cerambyx 905, 941 Chalcodrya 892
Carabidae 782, 792, 805, 806, 809, Catopochrotidae 878, 957 Cerapeplus 825, 965 Chalcodryidae 787, 892, 962, 1000
810, 812, 813, 914-921, 923, 924, Catopochrotus 878 Cerapterus 810 Chalcolampra 907
927,934,938,939,941,943, 956," Catops 821 Cerasommatidia 883 Chalcolepidius 854
959-961, 965, 966, 970, 971, 975, Catopsolius 820 Cerasommatidiidae 883 Chalcophana 907
983, 988, 995, 998 Catoxantha 842, 843 Ceratocanthidae 783, 794, 833, 834, Chalcophora 842
Carabinae 782, 811, 927 Catoxanthini 843 953,978 Chalcophorinae 842, 843
Carabini 811, 938 Cautires 857 Ceratocanthus 833 Chalepus 907
Carabus 811 Cautomus 881 Ceratoderus 81 0 Chameorhipis 841
Carcinognathus 850 Cavicoxum 854 Ceratognathus 831 Chanopterus 891
[10] [11]

Chapuisia 913 Chrysodema 842 Clavicornia 861, 872, 886, 914, 920, Codocera 833
Chapuisiinae 913 Chrysodina 907 933,937,955,963,969,981,986, Coelambus 809
Charhyphus 826, 948 Chrysolina 907 990, 995, 998, 1003 Coelocnemis 894
Chariessa 870 Chrysomela 907, 933 Claviger 825 Coelocryptus 879
Chariotheca 894 Chrysomelidae 787, 862, 906, 917, Clavigerinae 825, 920 Coelodes 834
Chasmatopterus 835 923,931,933,935,942,945,951, Clavilispinus 828 Coelometopinae 787, 796, 892, 894
Chauliognathidae 860 955, 956, 959, 963-965, 967-970, Cleoninae 912 Coelometopon 818
Chauliognathinae 784, 860, 967 979, 983, 985, 987, 989, 996, 998, Cleonus 912 Coelometopus 894
Chauliognathus 860 1002, 1005 Cleopteriinae 819 Coelophora 885
Cheirotonus 835 Chrysomelinae 787, 907, 957 Cleopterium 819 Coelopterus 884
Cheloderus 904 Chrysomeloidea 787, 903, 929, 955, Cleridae 785, 866, 869-871, 897, 901, Coelostoma 816
Chelonariidae 784, 837, 839, 845, 847, 956,967,973,983,987 918,924,928,930,932,936,941, Coelus 893
848, 978, 993 Cicindela 811, 984 958, 969, 1002 Coiffaitia 829
Chelonariomorphus 847 Cicindelidae 810, 941, 998, 1002 Clerinae 785, 870, 871 Colaspis 907
Chelonarium 839, 840, 847, 848, 969 Cicindelinae 782, 811, 961, 965 Cleroidea 785, 867, 871, 880, 918, 930, Colaspoides 907
Chetabraeus 816 Cicindis 810 932, 936, 958, 969, 992 Colasposoma 907
Chevrolatia 822 Cicones 890 Cleropiestinae 870, 969, 1002 Colenis 820
Chiasognathinae 832 Ciidae 786, 863, 867, 887, 957, 961, Cleropiestus 870 Colenisia 820
Chiasognathus 832 962, 969 Clerus 869, 870 Coleoptera 779-782, 788, 789,
Chileanthicus 902 Ciinae 786, 887 Clidicinae 794, 822 792-794, 797-800, 866, 914-1006
Chilecar 911 Cilea 826 Clidicus 822 Colilodion 825
Chilenius 864 Cillaeinae 785, 874, 957 Clinidium 809, 919 Colliuris 812
Chileolobius 883 Cillaeus 874 Clinops 889 Collops 872
Chiliotis 878 Cimberidinae 908 Clivina 812 Collyris 811
Chilocorinae 786, 885 Cimberis 908 Cloeotus 833 Colon 820, 821, 991, 996
Chilocorus 885 Cinnabariinae 841 Clypastraea 885 Colonellus 820
Chiloea 844, 934 Cinnabarium 841 Clypeaster 885 Colonidae 819, 996
Chiloeidae 844, 934 Cinyra 842 Clypeastrinae 885 Coloninae 782, 794, 820
Chionotyphlus 829, 992 Cioninae 912 Clypeodytes 809 Colophon 832, 988
Chiron 834 Cis 887 Clytra 907 Colopterus 874
Chironinae 834 Cisidae 887 Clytrinae 907 Colphotia 859
Chlaenius 812 Cisseis 843 Clytus 905 Colpius 808
Chlarnisinae 907 Cissites 896, 931 Cnecosa 879 Colpodes 812
Chlarnisus 907 Cistela 893
Cistelinae 893
Cnemalobus 812
Cnemeplatia 893
Coluocera 883
Colydiidae 786, 792, 796, 809, 874,
~
Chlamydopsinae 782, 816, 817
Chlamydopsis 817 Clada 864, 999 Cneoglossa 846 883,886,890,894,900,915,926, ~
Choleva 821 Cladodes 859 Cneoglossidae 784, 837, 846, 849, 932 931, 934, 945, 947-950, 953, 954, ~
II
Cholevidae 819, 927, 943, 979, 1006 Cladophorus 857 Cnodalinae 894 961, 974, 981, 983, 984, 991, 992,
Cholevinae 782, 794, 820, 821 C1adotoma 847 Cnodalon 894 995
Choragidae 908, 952 Cladotominae 784, 847 Cnopus 902 Colydiinae 786, 890
Choraginae 788, 909 Cladoxena 879 Coccidophilus 884 Colydiopeltis 868

~
Choragus 909 Cladoxeninae 879, 990 Coccidula 884 Colydium 890
Chromogeotrupes 833 Clambidae 783, 803, 817, 836, 839, Coccidulinae 786, 884 Colymbetes 809
Chromomoea 893 933,937,938 Coccinella 885, 929 Colymbetinae 782, 808, 809
Chrysanthia 895 Clambinae 783, 839 Coccinellidae 786, 883, 884, 919, 927, Comacupes 832
Chrysobothrinae 842, 843 Clamboidea 838 928, 944, 949, 951, 956, 958, 963, Compsocnemis 852
Chrysobothris 842 Clambus 839 970,980,981,985,986 Conalia 889
Chrysochroa 843 Clamoris 893 Coccinellinae 786, 885, 981 Conchifera 810, 812
Chrysochroinae 843 Claudiella 804 Coccotrypes 913 Conderis 857
Chrysochus 907 Cochliarion 818 Coniporidae 872
 [13]
[12]
Cossoninae 788, 913, 968, 971 Cryptoglossa 893 Cuphotes 894
Coniporus 872 Cryptognatha 884 Curculio 912, 971, 975
Conoderus 854 Cossonus 913
Cryptolarynginae 912 Curculionidae 788, 835, 910, 911, 913,
Conognatha 842 Cossyphinae 792, 796, 892, 893
Cryptolestes 877 924,958,959,966,968,975,977,
Conomorphus 896 Cossyphodes 894
Cryptomera 838 991 , 998-1000, 1002, 1005
Cononotidae 898 Cossyphodidae 892, 915
Cryptomeridae 838 Curculioninae 788, 912, 913
Cononotinae 796, 899 Cossyphodinae 787, 894
Cryptonychus 907 Curculionoidea 787, 907, 908, 915,
Cononotus 899, 936 Cossyphodinus 894
Cryptophagidae 786, 793, 795, 878, 924, 926, 933, 956, 959, 960, 966,
Conopalpus 888 Cossyphodites 894
900, 923, 927, 930, 945, 964, 966, 968, 971 , 975, 997, 1002, 1005,
Conopterum 812 Cossyphoditinae 894
968,988,991 1006
Conosoma 826 Cossyphus 892
Cryptophaginae 786, 878, 923 , 939 Curimopsis 844
Conotelus 874 Cotalpa 836
Cryptophagus 878, 991 Curtos 859
Conotrachelus 912 Cotes 901
Cryptophilidae 878, 879 Cussolenis 853, 940
Copelatinae 782, 808 Cotinus 836
Cryptophilinae 786, 879, 986 Cyathiger 825
Copelatini 808 Cotulades 891
Cryptophilus 878, 879 Cyathoceridae 804, 948, 990, 1005
Copelatus 808 Coxelus 890
Cryptopleurum 816 Cyathocerus 804
Copidita 895 Cranophorus 884
Cryptorama 865 Cybebus 911
Copobaeninae 787, 901 Craspedomerus 830
Cryptorhynchinae 912, 966, 977 Cybister 809
Copobaenus 901 Cratomorphus 859
Cryptorhynchus 912 Cybocephalidae 873
Coprinae 834, 940 Cratoscelis 834
Cryptosomatula 878 Cybocephalinae 785, 874
Copris 834 Cremastocheilus 836
Crypturgus 913 Cybocephalus 874
Coprophilus 828 Creobius 812
Ctenicera 854 Cychramptodes 874
Coproporus 826 Creophilus 830
Ctenidia 889, 941 Cychramus 874
Coptocera 841 Crepidodera 907
Ctenidiinae 786, 889 Cychrus 811
Coptodactylus 834 Crepidogaster 813
Cteniopinae 893 Cyclaxyra 877
Coptornia 836 Crepidomeninae 854, 926
Cteniopodinae 796, 893 Cyclocephala 836
Coptonotinae 913 Crepidomenus 854
Criocerinae 787, 906, 987 Cteniopus 893 Cycloderus 898, 980
Coptonotus 913 Ctenistes 825 Cyclommatus 832
Coptotomus 809 Crioceris 906
Crossotarsinae 913 Ctenodactyla 812 Cyclotoma 883
Coptotrophis 817 Ctenoplectron 888 Cycloxenus 881
Cordus 910 Crossotarsus 913
Ctenostoma 811 Cydistus 841, 857, 858
Comeolabium 824 Crotchia 879
Crowsoniella 801, 933, 976 Ctesibiinae 850 Cydonia 885
Coroebus 843 Ctesibius 850, 962 Cyladidae 910
Corotoca 827 Crowsoniella Relicta 801, 976
Crowsoniellidae 782, 801 Cucujidae 786, 809, 872, 876, 877, Cyladinae 788, 910
Corthylus 913 894, 896, 924, 933, 945, 951, 964, Cylas 910
Corticaria 886 Crymus 824
975, 983, 988, 991, 997, 1002, Cylidrus 869
Corticariidae 795, 872, 886 Cryphalus 913
Cryptamorpha 876 1003 Cylindrocaulus 832
Corticariinae 786, 886, 955 Cucujiformia 785, 798, 838, 861, 865, Cylindromorphinae 843
Corticeus 894 Cryptamorphinae 876 :··
Cryptarcha 874 866,949,1001 Cylindromorphus 843
Cortinicara 886, 955 Cucujoidea 785, 789, 867, 871, 872, Cylindropsis 828
Corylophidae 786, 817, 877, 885, 927, Cryptarchinae 785, 874
Crypticus 894 886, 914, 926, 934, 938, 947, 951, Cylindrorhininae 912
968, 976-978 962, 967, 969, 976, 982, 986, 991, Cylindrosella 819
Corylophinae 786, 885 Cryptobium 829
Cryptocephalinae 787, 907, 983 992 Cylindroxystus 829
Corylophodes 885 Cucujopsis 905 Cyllodes 874
Corylophus 885 Cryptocephalomorpha 812
Cryptocephalus 907 Cucujus 876 Cyloma 816
Corynetidae 869 Cuneipectus 812 Cymatodera 869
Corynomalus 883 Cryptodacne 879
Cryptoderma 912 Cupedidae 782, 799-801, 803, 917, Cymbeba 892
Coryphium 824 932, 970, 973 Cymbiodyta 816
Corythoderus 834 Cryptodontes 836
Cryptodus 836 Cupes 803, 804, 973 Cymbionotum 812
Cosmocerinae 864 Cupesidae 803, 999, 1000 Cynegetis 885
Cossonidae 911 Cryptogenius 834, 952, 988
 [15]
[14]
Dasycerinae 783, 825 Derodontus 861 Dilolycus 857
Cyparium 828 Dasycerus 825, 965 Derolathrinae 865, 866 Dilophotes 856
Cypha 827 Derolathrus 865, 883, 887 Dima 854
Dasychaeta 834
Cyphagogus 910 Derops 826, 992 Dimerus 825
Dasydactylus 879
Cyphaleus 893 Deroptilinus 865 Diminae 784, 854
Dasypelates 820
Cyphanodes 839 Derosphaerus 894 Dinapate 863
Dasytes 870, 872
Cyphanus 839 Dasytes Fuscipennis 870 Derovatellus 809 Dineutes 807
Cypherotylus 880 Desmidophorinae 912 Dinoderinae 785, 864
Dasytidae 871, 999
Cyphinae 827 Desmopachria 809 Dinoderus 864
Dasytinae 785, 872
Cyphogastra 842 Decamerinae 785, 867, 868 Dexorini 856 Dinomorphinae 912
Cyphon 839 Dexoris 856 Dinomorphus 912
Decamerus 868
Cyphonidae 839, 840 Dhysores 809 Diochinae 830
Decarthrocerus 818
Cyphonocerinae 784, 859 Diabathrariinae 912 Diochus 830
Decarthron 825
Cyphonocerus 859 Diabletes 817 Dioedus 893
Dechomus 890
Cyphopisthes 833 Diabrotica 907 Diomus 884, 944
Declinia 838
Cyphotelus 839 Decliniidae 783, 838, 962, 974 Diacalla 901 Diontolobus 868
Cyprogenia 878, 982 Diacheila 811 Dioptoma 857, 858
Deinopsis 827
Cyrtonops 904 Deinopteroloma 822, 824, 992 Diachus 907 Diphyllidae 880
Cyrtoscydmus 822 Diadoxus 842 Diphyllostoma 832, 950
Deleaster 828
Cyrtosoma 894 Diagrypnodes 900 Diphyllus 880, 939
Delevea 804
Cyrtotriplax 880 Diamerus 913 Diplocladon 857, 858
Delognatha 893
Cyrtusa 820, 934 Diamesus 822 Diplocoelus 880, 939
Delphastus 884
Cysteodemus 896 Dianous 829, 982 Diplocotes 864
Deltochilum 834
Cytilus 844 Diaperinae 787, 894 Diplognatha 836
Deltocryptus 865
Dacne 879 Demarziella 834, 968 Diaperis 894 Diplotaxis 835
Dacnidae 879, 888 Diaphanes 859 Dipsaconia 891
Dendezia 832
Dacninae 786, 879 Diaphanops 906 Diptera 799, 826, 871, 952, 979
Dendroblax 831
Dacoderidae 899, 1000 Diaphonia 836 Dircaea 888
Dendrocharis 852
Dacoderinae 787, 900 Diaporinae 913 Dircaeinae 888
Dendroctonus 913
Dacoderus 900 Diapus 913 Dirocephalus 828
Dendroides 899
Dactylostemum 816 Diatelium 828 Discodon 860
Dendrophagus 876, 933
Dadophora 859 Dendrophilinae 782, 816, 817 Dibelonetes 829 Discogenia 877
Dadophorus 884 Dicerca 842 Discoloma 882
Dendrophilus 817
Daemon 846 Denticollinae 784, 794, 854, 930, 961 Dichelonyx 835 Discolomatidae 786, 795, 877, 882
Danacaea 872 Denticollis 854, 961 Dichotomius 834 Discolomatinae 786, 882
Danae 883 Diclidia 903 Discolomidae 795, 882, 954, 955
Deporaus 910
Dapsa 883 Dicoelocephalus 865 Discotenes 909
Derallus 816
Darwinella 891 Dicordylus 909 Discotoma 885
Dascillidae 783, 830, 836, 837, Derancistrus 904
Deretaphrinae 881 Dicranopselaphus 846 Discozantaena 818
839-842, 846, 848, 849, 861, 862, Dicranostema 907 Dissochaetus 821
Deretaphrus 881
928,941,980,990 Dicrepidiini 855 Distenia 904
Dermestes 862
Dascilliformia 836, 861 Dermestidae 784, 861-863, 865, 867, Dicrepidius 854 Disteniidae 903, 970
Dascillinae 783, 841 880, 915, 919, 927, 971, 977, 978, Dicronychidae 852, 988 Disteniinae 787, 904
Dascillocyphon 880 Dicronychus 855, 988 Distipsidera 811
983,988,1005
Dascilloidea 783, 830, 836, 837, 840, Dictyoptera 857 Distocupes 803
841,848,850,861,862,932,956, Dermestinae 784, 862
Dermestoidea 861, 862, 865, 931 Dienerella 886 Distocupes Varians 803
967 Derodontidae 784, 813, 861, 862, 865, Dieropsinae 870, 1002 Distremocephalus 858
Dascillus 839-841, 880, 943 Dieropsis 870, 1002 Ditoneces 857
942, 962, 974
Dascillus Cervinus 841, 943 Dietta 820 Ditropidus 907
Derodontinae 784, 861
Dastarcus 881 Dihammatus 857 Ditylus 895
Derodontoidea 784, 861
Dasyceridae 822, 823, 965
 [17]
[16]
Elacatis 899 Emphylus 878
Dumbrelia 857 Elaphrinae 782, 811 Empocryptus 879
Docalis 891 Dynamopinae 794, 835, 916
Dodecatoma 857, 858, 1001 Elaphrus 811 Encaustes 879
Dynamopodinae 783, 794, 835 Elater 854, 947 Encaustinae 786, 879
Dodecatoma Bicolor 858
Dynamopus 835 Elateridae 784, 792, 837, 849, 851 , Enceladus 811, 938
Doesus 904 Dynastes 836
Dolicaon 829 852,855,926,927,930,932,935, Encymon 883
Dynastinae 783, 836, 937 939, 940, 943, 944, 946, 947, 951, Endecatomidae 785, 863
Dolichosoma 872 Dyscharachthis 852
Dolosidae 881 961,972,975,984,995,998,1005 Endecatomus 863, 864, 932, 958
Dyscherus 812 Elateriformia 783, 784, 813, 830 Endeodes 872
Dolosus 881, 934
Dyschirius 812 836-838, 840, 860, 861 956 962, Endestes 890
Domene 829 Dyscinetus 836 963 ' ' '
Donacia 866, 906, 983 Endognathus 833
Dysides 863 Elaterinae 784, 853-855 Endomia 902
Donaciinae 787, 906, 917, 955, 964,
Dysidinae 785, 863 Elateroidea 784, 794, 836, 837 843 Endomychidae 786, 865, 882-884 887
967,983
Donaldia 871
Dysmorphocerinae 784, 860 6
848-850, 852, 855, 861 92 930' 955, 976, 983, 986, 995, 996, '
Dysmorphocerus 860 943 ' ' ' Endomychinae 786, 883
Dorcadion 905
Dystaxia 842 Elateroides 851, 866, 967 Endomychus 883
Dorcaschema 905
Dystaxiella 842 Elateromorpha 836 Engidae 879
Dorcatoma 865, 991
Dystheamon 882 Electribiinae 784, 850 Engis 879
Dorcatominae 785, 865, 939, 1002 Dytiscidae 782, 806, 808, 915, 921,
Dorcinae 832 Electribiini 850 Enhydrus 807
926,941,945,974,975,982,985, Electribius 850, 962 Enhypnon 890
Dorcus 832 993,998,1003,1005
Dorkatoma 865 Electropogon 850 Enicmus 886
Dytiscinae 782, 809 Electropogonini 850 Enicodes 905
Dorylomimus 827
Dytiscus 809 Elenophorus 893 Enicopus 872
Dorynota 907 Dytoscotes 826
Doydirhynchinae 787, 908 Eleodes 893 Enneadesmus 863
Ebaeus 872 Elephastomus 832 Ennearthron 887
Doydirhynchus 908
Eblisia 817 Eletica 896 Enneboeopsis 887
Drapetes 853, 925
Drilidae 784, 837, 856-859, 944, 1003 Ecanus 819 Eleticinae 787, 896, 989 Enneboeus 887
Ecbletus 826 Eleusininae 828 Ennometes 848
Drilocephalus 841 ,
Eccoptarthrini 911 Eleusis 828 Enochrus 816
Driloniinae 784, 859
Echiaster 829 Elgonidium 902 Enoclerus 870
Drilonius 859 Echthrogaster 852
Drilus 856 Elianus 860 Enopliinae 785, 795, 870
Ecitogaster 827 Elmidae 783, 815, 837, 842-844 925 Enoplium 870
Dromaeolus 852
Ecnolagria 892 949, 953, 993 ' ' Entiminae 912
Dryocoetes 913 Ectemnorhininae 912
Dryocora 894 Elmididae 844 Entimus 912
Ectinocephalus 885 Elminae 783, 844 Entomoculia 829
Dryophilinae 785, 864
Ectopria 846 Elminthidae 844, 949, 975 Entomophaga 800, 914
Dryophilodes 864
Ectrephes 864 Elmis 844, 933, 953 Entomosatopus 852
Dryophilus 864 Ectrephidae 864
Dryophthorinae 788, 796, 912 Elmopamus 844 Entomoscelis 907
Edaphus 829, 981 Elodes 839 Eobia 895
Dryophthorus 912
Edusella 907 Elodidae 839, 840 Eocatops 821
Dryopidae 784, 837, 838, 842, 844,
845,918,923,925,934,948,969, Egestria 901 Elophorinae 815 Eodromeinae 806
Egestriomima 901 Elophorus 815 Eopaussus 810
975,979,998,1005
Egidyella 863 Elythomerinae 784, 845 Eospasta 896
Dryopoidea 836, 843, 848, 849, 925,
Egidyellinae 863 Elythomerus 845 Ephisteminae 878
933, 934, 948, 949, 978, 994, 995,
Egolia 869 Elytraria 800 Ephistemus 878
1004 Egoliinae 785, 867-869
Dryops 844, 948 Emelinus 902 Epicauta 896
Eicolyctus 879, 990 Emmita 841 Epiechinus 817
Drypta 812 Eidoreinae 883
Duboisius 901 Empelidae 822 Epierus 817
Eidoreus 883, 986 Empelinae 783, 824 Epilachna 885
Duliticola 856 Elacatidae 899, 928
Duliticola Paradoxa 856 Empelus 824 Epilachnidae 884
Elacatinae 899
Duliticolinae 856
[18] [19]

Epilachninae 786, 885, 944 Ethra 859 Eumalus 828 Euscaphurus 838
Epilichadinae 847 Euaesthetinae 783, 829, 976, 981, 982 Eumenes 851 Euscelus 910
Epilichas 847, 847 Euaesthetus 829 Eumetopus 815 Eusphalerum 824
Epimetopinae 782, 815 Eubaptus 906 Eumolpinae 787, 862, 907, 959 Euspilotus 817
Epimetopus 815 Eublackbumiella 820 Eumolpus 907 Eustilbus 877
Epiphloeinae 785, 795, 869, 870 Eubria 846 Eumorphinae 795, 883 Eustilicus 829
Epiphloeus 870 Eubrianacinae 784, 846 Eumorphus 883 Eustra 810
Epipocinae 786, 795, 883 Eubrianax 846 Euops 910 Eustrophinae 786, 888
Epipocus 883 Eubriidae 846 Euparia 834 Eustrophinus 888, 999
Episcapha 879 Eubriinae 784, 846, 963 Euparius 909 Eustrophopsis 888
Episcaphula 879 Eucaliga 893 Eupelates 820 Eustrophus 888
Epispasta 896 Eucalosphaera 874 Euphanias 828 Eutheia 822
Epistomentis 842 Eucamaragnathus 811 Euphoria 836 Euthysanius 853
Epitragus 893 Eucanthus 832, 833, 925, 950 Eupiestus 828 Eutylistus 865
Epiverta 885 Eucatopinae 821 Eupines 825 Euxestidae 881
Epuraea 874 Eucatops 821 Eupisenus 888 Euxestinae 786, 881
Erchomus 826 Euchirinae 783, 835, 1005 Euplectinae 825 Euxestocis 887
Eremazus 834 Euchirus 835 Euplectus 825 Euxestoxenus 881
Erernninae 912 Eucilodes 838 Eupompha 896 Euxestus 881
Eremochilus 885 Eucinetidae 783, 836, 838, 980, 985, Eupsalis 910 Evaniocerinae 889
Eretes 809 999 Eupsilobiinae 786, 883, 976 Evaniocerus 889
Ergates 904 Eucinetiformia 836, 838 Eupsilobius 883 Evolocera 883
Erichsonia 904 Eucinetoidea 794, 836-838, 937, 938, Eupsorus 824 Evorinea 863
Erichsonius 830 999, 1001 Eurelymis 871 Exocalopus 898
Ericmodes 872 Eucinetus 838, 999, 1001 Eurhynchidae 910, 1006 Exochomus 885
Erirhininae 912 Eucnemidae 784, 794, 837, 851, 855, Eurhynchinae 788, 910 Exohadrus 891
Emobiinae 785, 864 926, 929, 967, 972 Eurhynchus 910 Exoplectra 884
Emobius 864 Eucneminae 784, 852 Europs 874 Exostemus 817
Eronyxa 867, 868, 996 Eucnemis 852 Eurychora 893 Fabrasia 864
Eronyxa Expansus 868, 996 Euconnus 822 Eurycratus 900 Falagria 827
Eros 857 Eucrada 864 Eurygeniinae 787, 899, 901, 914 FaBia 882
Erotinae 784, 857 Eucradinae 785, 795, 864 Eurygenius 901 Falsocis 887
Erotylathris 881 Eucranium 834 Eurynassa 904 Falsomelyris 871
Erotylidae 786, 878-880, 883, 886, Eucteanus 883 Euryopa 858 Falsomycterus 893
916, 924, 929, 931, 935, 951, 985, Eucymbolus 871 Eurypalpidae 846 Falsophrixothrix 857
990 Euderia 864, 932 Eurypalpus 846 Falsoplatydema 887
Erotylinae 786, 880 Euderiinae 785, 864 Eurypinae 796, 897 Falsotherius 847
Erotylus 880 Eudicronychinae 784, 855 Eurypinus 896 Falsoxanthalia 888
Ertliana 896 Eudicronychus 855 Eurypogon 850 Faroninae 825, 931
Esarcinae 786, 886 Euglena 903 Eurypogonidae 837, 849, 930 Faronus 825
Esarcus 886, 934 Euglenes 902 Eurypogoninae 850 Fenderia 829
Escalerina 841, 978 Euglenesidae 902, 903, 952 Euryptychus 852 Feronia 812
Escalerina microcephala 841 Euglenidae 796, 902, 903, 917, 952 Eurypus 896, 903 Fidia 907
Escalerina serraticornis 978 Eugnamptus 910 Euryscaphus 812 Figulinae 832
Escalerosia 902 Eugnominae 912 Eurysphindinae 873 Figulus 832
Esemephe 894 Eugonus 909 Eurysphindus 873, 969 Filicivora 887
Espeson 828 Eulagius 887 Eurystemus 834 Flabellotrichalus 857
Estadia 820 Eulichadidae 784, 837, 848, 861 Eurystethes 900 Foadia 885
Estadiini 822 Eulichas 848 Eurystethidae 899 Forcipator 812
Ethon 843, 1000 Eulimulodes 819 Eurystethinae 900 Formicomus 902
[20] [21]

Fornax 852 Glaphyridae 783, 834 Gyrinidae 782, 805, 806, 808, 921, Herniconderis 857
Frickius 833, 950 Glaphyrus 834 926,940,941,947,961,998,1005 Hernicrepidius 854
Glaresidae 783, 832, 988 Gyrininae 782, 807 Herniopinae 784, 854
Frostia 860
Fuchsina 886, 915 Glaresis 832, 988 Gyrinus 807 Herniops 854
Fulcidax 907 Glenus 830 Gyrophaena 827 Herniopsida 852
Gabrius 830 Glipa 889 Habrocerinae 783, 826 Herniopsinae 854
Galbella 843 Glischrochilus 874 Habrocerus 826 Hemipeplidae 896
Glypholoma 822, 823, 973, 996 Habroloma 843 Hernipeplinae 787, 897
Galbellinae 843
Glypholomatinae 783, 794, 823 Hadraule 887 Hernipeplus 897, 997
Galbites 852
Galerita 812 Glypholomatini 824 Hadrobregmus 865 Hemipleurus 871
Galeruca 907 Glyphonyx 854 Hadrognathus 824 Hernirhipidiinae 786, 889, 890, 998
Galerucella 907 Glyptolopus 881 Haematodes 841 Hemirhipidius 889
Galerucinae 787, 907, 983, 1002 Glyptoma 828 Halacritus 816 Hemirhipinae 854, 944
Glyptoscelimorpha 842 Haliplidae 782, 806, 807, 919-921, Hemirhipus 854
Ganascus 902
Ganyme 891 Glyptoscelis 907 941, 988, 993, 998, 1005 Henoticus 878
Gastrallus 865 Glyptus 812 Haliplus 807, 919, 920 Henotiderus 878
Gaurotes 905 Gnathium 896 Hallomeninae 786, 888 Hephaestion 905
Geadephaga 805, 806, 924 Gnathoncus 817 Hallomenus 888 Hesperobaenus 874
Gehringia 810, 811, 919, 965, 966 Gnathymenus 829 Halyzia 885 Hetaeriinae 782, 816, 817, 948
Gehringiinae 782, 810 Gnatocerus 894 Hamotus 825 Hetaeriomorphus 817
Gehringiini 811 Gnostidae 864 Hapalips 879 Hetaerius 817
Gempylodes 890 Gnostus 864 Haplocnemlnae 871 Heterhelus 873
Genecerus 841 Goliathus 836 Haplogastra 814, 830, 861, 977 Heteroceridae 784, 837, 843, 845, 862,
Geniates 836 Golofa 836 Haploglossa 847 928, 976, 1005
Genisphindus 873 Gondwanenneboeus 887 Haploglossinae 847 Heterocerinae 784, 845
Geornitopsis 828 Goniacerinae 825 Haplostethinae 843 Heterocerus 845
Geomysaprinus 817 Goniacerus 825 Haplostethus 842 Heterocheira 893
Geoparnus 844 Goniadera 892 Harmatelia 859 Heterogastra 800, 977
Geopinus 812 Gonicoelus 880 Harpalinae 782, 792, 794, 812 Heterogyrus 807
Georissidae 815 Gonioctena 907 Harpalus 812 Heteromastix 860
Georissinae 782, 815 Gonipterinae 912 Hectarthrum 876 Heteromera 886, 894, 914, 919, 922,
Georissus 815 Gonipterus 912 Hedobia 864 923,928,933,936,941,961,967,
Georyssidae 815, 993, 1005 Gonocephalum 893 Hedobiinae 795, 864, 939, 999 974, 978, 980, 995, 998, 1000,
Geory ssinae 815 Gonzalosia 902 Helcogaster 872 1004
Geotrupes 833 Grammoptera 905 Helea 893 Heteromeroxylon 866
Geotrupidae 783, 832, 833, 925, 959, Graphipterinae 792, 794, 812 Helferella 842 Heteronychus 836
977, 1006 Graphipterus 812 Helichus 844, 998 Heteronyx 835
Geotrupinae 783, 833, 950, 951, 1003, Grynobius 864 Heliocopris 834 Heteropaussus 810
1006 Grynocharis 868 Helluo 812 Heterophaga 799
Germarica 843 Grynoma 868 Helluodes 812 Heteroscapha 828
Ghanius 884 Gymnetis 836 . Helminthidae 844 Heterosilpha 822
Gibbiinae 864, 920 Gymnetrinae 912 Helochares 816 Heterothops 830
Gibbium 864 Gymnetron 912 Helodes 839, 840, 958 Hexagonia 812
Gietella 872, 930 Gymnochthebius 818 Helodidae 794, 836, 839, 840, 946, Hexaplatarthrus 810
Gietellinae 785, 872, 930 Gymnognathus 909 958,975,980,981 Hexarthrum 913
Ginglymocladus 859 Gymnopleurus 834 Helophoridae 815 Hexodon 836
Giulianium 824, 826, 971 Gymnusa 827 Helophorinae 782, 815, 916, 992 Hiletinae 782, 811, 812
Glabricornia 806 Gynopterus 864 Helophorus 815, 952 Hiletus 811
Glacicavicola 820 Gyretes 807 Helota 875, 942, 944, 975 Hintonia 804
Glacicavicolinae 820 Helotidae 785, 875 Hintonosia 902
[22] [23]
Hippodamia 885 Hughleechia 818 Hylecoetopsis 866 Ichnea 870
Hippomelas 842 Hybosoridae 783, 834, 952, 988 Hylecoetus 866 Ichnoldes 870
Hipporhininae 912 Hybosorus 834 Hylesininae 913 Ichthyurus 860
Hispa 907 Hydaticini 809 Hylesinus 913 lctistygna 901
Hispinae 787, 907, 923, 998 Hydaticus 809 Hyliotes 876 lctystygna 901
Histanocerus 887 Hyderodes 809 Hyliotinae 876 ldgia 871, 928
Rister 817 Hydnobioides 877 Hylis 852 ldiophyes 884
Histeridae 782, 814-816,921,943,944, Hydnobius 820 Hylobia 888 Idiostoma 835
948, 955, 958, 959, 967, 968, 973, Hydnocera 869 Hylobius 912 Idiostominae 835
982, 999, 1004 Hydnocerinae785, 795,869,870,1003 Hylochares 852 Idolia 817
Histerinae 782, 817 Hydnorobius 909 Hylophilidae 902, 917, 980 Ilybius 809
Histeroidea 814, 815, 920, 933, 959, Hydora 844, 993 Hylotorus 810 Inca 836
974 Hydradephaga 805, 806, 811, 917, Hylurgops 913 Incolia 888
Histeromorphae 816 919-921, 926, 935, 974, 985 Hymaea 875 Incollogenius 898
Hobartiidae 786, 877 Hydraena 818 Hymaeinae 785, 875, 962 Incoltorrida 804
Hobartius 877 Hydraenida 818 Hymenoptera 799, 976 lndalmus 883
Holisus 830 Hydraenidae 782, 813-815, 817, 818, Hymenorus 893 Inopeplidae 899, 986
Hololepta 817 946,953,976,979,993 Hyocis 894 Inopeplinae 787, 900
Hololeptinae 817 Hydraeninae 782, 818 Hyorrhynchus 913 Inopeplus 900
Holoparamecinae 786, 883 Hydrobiinae 816 Hyperaspis 884, 924 Inscutomonomma 890
Holoparamecus 883, 887 Hydrobiomorpha 816 Hyperinae 912 Integripennes 810
Holopeplus 897 Hydrobius 816 Hyperomma 829 lnvolvulus 910
Holophygus 882 Hydrocanthus 808 Hyphalinae 784, 845 Ipelates 819
Holopsis 885 Hydrochidae 815 Hyphalus 845 Iphthiminus 894
Holosternus 900 Hydrochinae 782, 815 Hyphydrus 809 Ipidae 793, 913
Holostrophus 888 Hydrochus 815 Hyplathrinus 885, 976 Ipinae 913
Holosus 828 Hydrocoptus 808 Hypnoidinae 854, 995 Ipini 913
Holotrochus 828 Hydromedion 891 Hypnoidus 854 Ips 913
Homaeotarsus 829 Hydronebrius 809 Hypoborus 913 Ipsimorpha 900 I
Homalaena 818 Hydrophilidae 782, 813-815, 916, 918, Hypocacculus 817, 982 Isch.alia 901, 922, 1004
Homalisidae 856, 926 946,975,992,993,1005 Hypocaccus 817 Ischaliinae 787, 901, 1004

I
Homalisus 856 Hydrophilinae 782, 816 Hypocephalidae 903, 904 Ischiodontus 854
Homalocerus 909 Hydrophiloidea 782, 789, 800, 803, Hypocephalus 904 Ischnomera 895
Homalota 827 814,815,946,947,958,973 Hypochaetes 853 Ischnosoma 826
Homoeotelus 880 Hydrophilus 816 Hypocopridae 878 Ischyomiinae 897
Homopterus 810 Hydroporinae 782, 809, 915, 1003 Hypocoprinae 786, 878
Hypocoprus 878
Ischyomius 897 J
Hoplandria 827 Hydroporus 809 Ischyropalpus 902
I

Hoplia 835 Hydroscapha 803, 804, 924, 975 Hypocyphtinae 827, 827 Ischyrus 880
I
Hoplicnema 885, 976
Hopliinae 835
Horatocera 848
Hydroscaphidae 782, 804, 817, 982,
983
Hydrotrupes 808, 809, 921
Hypocyphtus 827
Hypodacne 881
Hypodacnella 881
!selma 896
Isochaeta 805, 806, 810, 919
Isoclerus 869
~I
'
'I
1

~
Horelophinae 782, 815 Hydrous 816 Hypodesus 854 Isohydnocera 869
Horelophus 815 Hydrovatus 809 Hypolithinae 854 Isopteron 892
Hypolithus 854
Horia 896, 931
Horiidae 792, 796, 895, 896
Hygriobia 808, 951
Hygrobia 808, 920, 951 Hyporhagus 890
Isorhipis 852
Istrisia 900
i
~
Horiinae 796, 896 Hygrobiidae 782, 808, 925, 941, 1005 Hypotelus 828 Ithyceridae 788, 907, 911
Horistonotus 854 Hygronoma 827 Hypothenemus 913 Ithycerus 911, 985
Hornia 896 Hygrotus 809 Hypulinae 888
Hoshihananomia 889 Hylastes 913, 964 Hypulus 888
Jacobsoniidae 785,813, 861, 865, 866,
883, 887, 948, 965
II
Hovactyla 847 Hylecoetinae 785, 866 Ibicarella 883 Jacobsonium 865 i j~
!I

~
I
[24] [25]

Jalodis 842 Lagrioida 901, 914, 931 Latridiidae 786, 795, 872, 886, 887 Leptopiinae 912
Janbechynea 906 Lagrioidinae 787, 901 Latridiinae 786, 886 Leptopius 912
Jauravia 884 Lagriomorpha 901, 902, 1004 Latridius 886 Leptoscydmus 822
Jelinekia 873 Laius 872 Lawrencerosus 874 Leptotyphlinae 783, 829
Jentozkus 838 Lamellicornia 830, 916, 918, 937, 948, Leanobium 865 Leptotyphlus 829
Jubus 825 951,966,969, 981 Leasia 907, 970 Leptura 905
Julodella 842 Lamellipalpus 859 Leaus 892, 893 Lepturinae 787, 905
Julodimorpha 842 Lamia 905 Lebasiella 870 Leptusa 827, 976
Julodinae 783, 794, 842 Lamiinae 787, 905 Lebia 812 Lethrinae 783, 833
Julodis 842, 922 Lamingtoniidae 786, 878 Lecontellus 908 Lethrus 833
Kalissus 825 Lamingtonium 878 Lecontia 897, 1004 Leucotachinus 826
Karumia 841 Lamprigera 859 Lederia 888, 974 Lewisium 885
Karumiidae 830, 837, 840, 848, 849, Lamprima 831 Leichenum 893 Libnetis 857
853,858,916,932,967, 978,993 Lampriminae 783 , 831 Leiestes 883 Lichadidae 848
Karumiinae 783, 841 Lamprocera 859 Leiestinae 786, 883 Lichas 847, 848
Katagenius 887 Lamprosoma 907, 970 Leiochrodes 894 Lichenophanes 863
Kateretes 873 Lamprosomatinae 787, 907, 970 Leiodes 820, 918 Lichnanthe 834
Kateretidae 795,873,917 Lampyridae 784, 837, 856-859, 934, Leiodidae 782, 794, 817, 819, 822, Lichnia 834
Katoporus 882 963, 964, 968, 975 824,918,934,979,1001 Lichniidae 834
Kaveinga 809 Lampyrinae 784, 859 Leiodinae 782, 820 Licinus 812
Kolon 821 Lampyris 859 Leleupaussus 810 Ligyrus 836
Korynetes 870 Lancetes 808, 809, 985 Lema 906 Lilioceris 906
Korynetidae 869 Lanelater 854 Lemidia 869 Limbata 810, 812, 813
Korynetinae 785, 870 Langelandia 890, 934 Lemodes 901, 922 Limnebiidae 818
Kryptogenius 887 Languria 879, 931 Lemodinae 787, 901 , 902 Limnebius 818
Kryptophagus 878 Languriidae 786, 878, 880, 963, 966, Lemodinus 901 Limnichidae 784, 837, 845, 924, 925,
Kytorhinus 906 986, 989, 990 Lenacinae 874 949,978,993,1003,1004
Laccobius 816 Languriinae 786, 879 Lenax 874 Limnichinae 784, 845, 1004
Laccodytes 808 Lanternarius 845 Leoglymmius 809 Limnichites 845, 1004
Lacconectus 808 Lanthanus 900 Leperina 869 Limnichoderus 845, 1004
Lacconotinae 787, 796, 896, 897, 930 Lapethinae 881 Lepiceridae 782, 804, 948, 983 Limnichus 845, 978
Lacconotus 896 Lapethus 881, 991 Lepicerus 803, 804, 948 Limniidae 844
Laccophilinae 782, 808 Lara 844 Lepidopteryx 869 Limonius 854
Laccophilus 808 Larainae 783, 794, 844, 993 Leptacinus 830 Limulodes 819
Laccornis 809, 1003 Laricobiidae 861 Leptaulax 832 Limulodidae 818, 819, 936, 989
Lachna 893 Laricobiinae 784, 861 Leptinidae 819, 821, 920, 977, 979, Limulodinae 794, 819, 989
Lachninae 893, 893 Laricobius 861 1003 Linsleya 896
Lachnodactyla 847 Laridae 844, 952 Leptinillus 821, 977, 1003 Liochoria 844
Lachnogya 893 Larinae 794, 844, 925 Leptininae 794, 821, 953 Liodidae 819, 954
Lachnophorus 812 Larinotinae 785, 868, 992 Leptinotarsa 907 · Lioligus 844
Lacon 854 Larinotus 868 Leptinus 821, 920, 953, 979 Lioon 844
Laeliana 818 Lasconotus 890 Leptochirinae 828 Liparetrus 835, 925
Laemophloeidae 786, 877, 997 Lasiodactylus 874 Leptochirus 828 Liparochrus 834
Laemophloeus 877, 964 Lasioderma 865 Leptodiridae 819 Lispininae 828
Laemotmetus 876 Lathridiidae 795, 886, 915, 920, 934, Leptodirinae 794, 821 Lispinus 828
Laena 892 949,955,983,985,988,990,1000 Leptodirini 821 Lispotherium 910
Lagria 892, 893 Lathridius 886 Leptodirus 821 Lissodema 900
Lagriidae 892, 901, 923, 928, 947 Lathrobium 829 Leptolycinae 784, 856 Lissodeminae 900
Lagriinae 787,792,796, 886, 892, 893, Lathropus 877 Leptolycus 856 Lissomidae 852, 925, 926
899, 901 Latometus 891 Leptomastax 822 Lissominae 784, 851-853
[26] [27]
Lissomus 853 Lutrochidae 784, 837, 844, 845, 925 Mada 885 Mecynotarsus 902
Lissotes 832 Lutrochus 845, 925 Madrasostes 833, 834, 953 Medisores 809
Lissothyreus 853 Lycidae 784, 837, 856, 923, 958, 964, Madrasostes kazumai 953 Medon 829
Litargops 887, 970 972, 1002 Maechidius 835 Megacephala 811
Litargus 887, 970 Lycinae 784, 856 Magdalininae 912 Megacerus 906
Lithocharis 829 Lycoperdina 883 Magdalis 912 Megalodacne 879
Lithophilus 884 Lycoperdinella 883 Maheoptinus 864, 920 Megalodacninae 786, 879
Lithophorus 881 Lycoperdininae 786, 795, 883 Malachiidae 871 Megalopaussus 810
Lithostygnus 886 Lycoptis 868, 918 Malachiinae 785, 872 Megalopininae 829
Litochropus 877 Lycostomus 856 Malachius 872 Megalopinus 829
Litochrus 877 Lyctidae 863, 943, 983, 988 Malacocis 887 Megalopodidae 787, 905, 906, 960
Lixinae 912 Lyctinae 785, 864 Malacodermata 871, 937, 1003 Megalopodinae 787, 905, 983, 1005
Lixus 912 Lyctoxylon 864 Malchinus 860 Megalops 829
Loberinae 879 Lyctus 864, 991 Malgassochaetus 869, 936 Megalopsidia 829
Loberonotha 879 Lycus 856 Mallodrya 894 Megalopsidiinae 783, 829, 920
Loberoschema 879 Lygistopterus 857 Malthacodes 871 Megalopus 905, 970
Loberus 879 Lymexylidae 785, 798, 857, 866, 918, Malthininae 784, 860 Megalostomis 907
Lobiopa 874 1001 Malthinus 860 Megamerus 906
Loboglossa 896 Lymexylinae 785, 866 Malthodes 860 Megapenthes 854
Lobonyx 871 Lymexyloidea 785, 866 Mandalotus 912 Megarthroides 824
Lobopoda 893 Lymexylon 866 Mantichora 811 Megarthropsis 826
Loebliorylinae 881 Lymexylonidae 866, 959, 960, 988 Margarinotus 817 Megarthrus 824
Loebliorylon 881 Lyponia 857 Mariamela 907 Megascelidinae 907
Loeblioryloninae 786, 881 Lypropaeus 856 Mariouta 862 Megascelis 907, 919
Lomechusa 827 Lypropaeus Biguttatus 856 Marioutinae 785, 862 Megasemum 904
Longitarsus 907 Lyrosoma 819 Martinius 845, 1004 Megasoma 836
Lophioderus 822 Lyrosominae 819 Masoreus 812 Megastemum 816
Lophocateres 868 Lystronychus 893 Mastax 813 Megatoma 863
Lophocateridae 867 Lytta 860, 895, 896, 945 Mastiginae 782, 794, 822 Megatominae 785, 794, 863
Lophocaterinae 785, 867, 868 Lyttinae 896 Mastigus 822, 934 Megatyphlus 829
Lordithon 826 Machlotes 881 Mastinocerinae 858 Megaxenus 902, 903
Lorelus 892 Macranobium 865 Mastinocerini 858 Megetra 896
Loricaster 839 Macratria 901 Mastinocerus 858 Mei1ichius 883
Loricera 811 Macratriinae 787, 901 Mastochilus 832 Melaenus 812
Loricerinae 782, 811 Macraulacinae 784, 852 Mastogeniinae 842, 843 Melalgus 863
Loxandrus 812 Macraulacus 852 Mastogenius 842 Melanactes 854
Loxomeriformes 813 Macrocyphon 839 Mastoremus 901 Melanactinae 854
Lucaina 857 Macrodactylus 835 Mastostethus 905 Melandrya 888
Lucanidae 783, 831, 833, 916, 949, Macrodascillus 839 Mataeopsephus 846 Melandryidae 786, 888, 894-897, 903,
950, 957, 961, 967, 968, 971, 975, Macrogyrus 807 Matheteinae 784, 859 926, 932, 947, 967, 974, 999
982,984,988 Macrohelodes 839 Matheteus 859 Melandryinae 786, 888
Lucaninae 783, 832 Macrolister 817 Matus 809, 1003 Melanophila 842
Lucanobium 831 Macrolopha 905 Mayetia 825 Melanophthalma 886
Lucanus 832 Macrolycus 856 Mayetiinae 825 Melanotinae 854
Lucidota 859 Macronychus 844 Mecomacer 908 Melanotus 854, 947
Lucio 859 Macroplea 906 Mecopelmus 913 Melanoxanthus 854
Luciola 859 Macropogon 850 Mecoptera 799, 979 Melasidae 794, 851, 940, 987
Luciolinae 784, 859 Macropogoninae 850 Mecorchesia 888 Melasinae 784, 852
Luperca 811 Macrosiagon 890 Mecyclothorax 812 Melasis 852
Luprops 892 Macrotoma 904 -' Mecynodera 906 Meliboeus 843
 [29]
[28]
Microsternus 879 Murrnidiinae 786, 881, 883
Meligethes 874 Metoecus 889, 890 Microvalgus 836 Murrnidius 881, 949
Meligethinae 785, 874, 954 Metophthalrnus 886, 915 Microweisea 884 Mycetaea 884
Melittornrna 866, 943, 960 Metopsia 824 Microxenus 883 Mycetaeidae 882, 983
Melittornrnatinae 785, 795, 866 Metopsiinae 824 Migadopinae 782, 811 Mycetaeinae 786, 884
Melittornrninae 795, 866 Metriorhynchus 857 Migadops 811 Mycetina 883
Melittornrnopsis 866 Metriorrhynchus 857 Migdolus 904 Mycetochara 893
Melobasis 842 Metrioxena 909 Migneauxia 886 Mycetoma 888, 926
Meloe 896 Metrius 810, 923 Mirnanornrna 827 Mycetophagidae 786, 880, 886-888,
Meloetyphlus 896 Metrotyphlus 829 Mirnanornrnatinae 827 934, 947, 964, 970, 974, 977, 988
Meloidae 787, 792, 796, 860, 895, 896, Mexico 789, 803, 804, 816, 819, 825, Mirneciton 827 Mycetophaginae 786, 887
914, 923, 945, 946, 956, 966, 980, 826,845,850,857,869,876,891, Mirnernodes 87 4 Mycetophagus 887
989, 1001 904, 908, 916, 918, 922,924, 931, Mirnochotrya 857 Mycetoporus 826
Meloinae 787, 896 944, 946, 949, 958, 963-965, 968, Mirnogonus 828 Mychocerinus 881
Melolontha 835 971, 976, 985, 992, 993, 995-998, Minthea 864 Mychocerus 881
Melolonthinae 783, 835, 924, 925 1002, 1004 Minurus 910 Mychotheninae 795, 884
Melyridae 785, 870, 871, 930, 957, Meziurn 864 Mitua 892 Mychothenus 884
967, 978 Micholaerninae 786, 889, 999 Mnioticus 878 Mycophagus 819
Melyrinae 785, 871 Micholaernus 889, 999 Molarnba 885 Mycotretus 880
Melyris 871 Michthisorna 904 Molorchus 905 Mycotrupes 833
Melyrodes 871 Micilus 845 Molytinae 912, 959 Mycteridae 787, 896, 903, 908, 930,
Melytra 891, 892 Micracis 913 Monanus 876 997
Meneristes 893 Micragasrna 818 Monocharnus 905 Mycterinae 787, 896
Menirnus 894 Micragyrtes 824 Monocoryna 884, 970, 986 Mycterornirnus 896
Meralius 891 Micralyrnrna 824 Monoedidae 890 Mycterus 896, 933
Merizodus 812 Micraspis 885 Monoedus 890 Mylabrini 896
Meropathus 818 Micrencaustes 879 Monolepta 907 Mylabris 896, 923
Merophysia 883 Microcara 839 Monolobus 811 Myllaena 827
Merophysiidae 865, 882, 883, 990 Microceroterrnes 903 Monornrna 890 Myllocerus 912
Merophysiinae 786, 883 Microchaetes 844 Monornrnatidae 786, 796, 890, 891 Myloechinae 820
Merycidae 891, 914 Microdonacia 907, 955, 970, 983 Monornrnidae 796, 890, 891, 942 Myodites 889
Meryx 891 Microhoria 902 Monophy lla 869 Myrabolia 875
Mesagyrtes 819 Microjulistus 871 Monotorna 874 Myraboliinae 785, 875
Mesecanus 819 Microlaernus 877 Monotornidae 785, 795, 874 Myrrnacicelus 910
Mesoceration 818 Microlanguria 879 Monotorninae 785, 874 Myrrnechixenus 886, 894, 974
Mesocoelopodinae 785, 795, 865 Micrornalthidae 782, 800, 801, 918, Monotornopsis 874 Myrrnecholeva 820
Mesocoelopus 865 985 Montandonia 862 M yrrnetes 817
Mesocyphon 839 Micrornalthus 801, 918, 937, 981, 985, · Mordella 889 Mystes 888
Mesoporinae 827 988 Mordellidae 786, 889, 903, 919, 941, Mystropornus 810
Mesoporus 827 Micropeplidae 822, 823, 965 942, 965, 975 Mystrops 874
Mesothinae 865 Micropeplinae 783, 823, 825, 927 Mordellinae 786, 889 Myxophaga 782, 799-801, 803, 805,
Mesydra 814 Micropeplus 825 Mordellistena 889 807, 817, 918, 949, 969, 982, 983
Mesynodites 817 Microphotus 859 Marion 812 Myzia 885
Metablax 853, 926 Microporurn 87 4 Morrnolyce 812 Nacaeus 828
Metacerylon 881 Microrhagus 852 Morpholycus 898 Nacerdes 895
Metaceryloninae 881 Microscydrnus 822 Morphozonitis 896 Nacerdinae 895
Metachrorna 907 Microsilpha 824 Morychus 844 Narnibiotalpa 833
Metacorneolabiurn 824, 996 Microsilphinae 783, 822, 824 Motschulskiurn 818 Narnunaria 890
Metacucujus 875 Microsporidae 782,794,804,805,817, Mouhotia 812 Nanobius 829, 948
Metapria 874 952 Murrnidiidae 881 Nanodes 910
Methles 809 Microsporus 804, 805
[30]
[31]
Nanophyes 910, 952 Neohypnus 830 Nomius 812 Ochodaeinae 783, 833, 927, 1003
Nanophyinae 788, 910, 915 Neojulodis 842 Nordenskioldia 829 Ochodaeus 833
Nanosella 819 Neolosus 828 Noserinus 891 Ocholissa 900
Nanosellinae 782, 819 Neomida 894 Noserosus 891 Ochotrya 857
Nargomorphus 821 Neopelatopini 820 Nosodendridae 784, 861-863, 865,930, Ochrolitus 877
Narthecius 877 Neopelatops 820 937, 983 Ochthebiinae 782, 818
Nascio 842 Neophaenognatha 835, 915 Nosodendron 862 Ochthebius 818, 953
Naupactus 912 Neophengus 858 Nosoderma 891 Ochthephilum 829
Neaspis 868 Neophoninae 783, 825 Nosoderminae 891 Octavius 829
Neatus 893 Neophonus 825, 996 Nossidium 818 Octinodes 853, 855
Nebria 811 Neoptinus 864, 920 Notaticus 809, 993 Octocryptus 854
Nebriini 811 Neopyrochroa 899 Noteridae 782, 806,808, 919, 941, 993 Octoglossa 847
Nebriosoma 811 Neorchesia 888 Noterinae 782, 808 Octomicrus 825
Necrobia 870 Neotetralobus 853 Noterus 808, 919 Octoternninae 887
Necrobiopsis 869 Neotrichus 890 Nothenneboeus 887 Octoternnus 887
Necrodes 822 Neotyphlus 829 Nothinae 888 Ocypus 830
Necrophila 822 Nephanes 819 Nothoderodontus 861, 962 Odacantha 812
Necrophilus 819, 1006 Nephrites 889, 989 Nothomorpha 842 Odontionopa 907
Necrophorus 822 Nephritinae 889, 890, 984 Nothotelus 903 Odontochodaeus 833
Necydalinae 787, 905 Nephus 884, 944 Nothus 888 Odontolabinae 832
Necydalis 905 Neptostemus 808 Notiokasis 811 Odontolabis 832
Negastriinae 784, 854, 995 Nerthopinae 912 Notiophilini 811, 938 Odontonyx 847
Negastrius 854 Nesolathrus 887 Notiophilus 811 Odontosphindinae 785, 873
Neichnea 870 Nesoneini 824 Notiophygidae 882, 942, 954, 955 Odontosphindus 873
Nemadinae 821, 996 Nesoneus 824 Notiophyginae 786, 882 Odontotaenius 832
Nemadus 821 Nesopeplus 874 Notiophygus .882 Oeatus 894
Nematidium 890 Neuraphes 822 Notobrachypterus 873 Oedemera 895
Nematoplinae 787, 895 Nicaginae 783, 831 Notocerastes 891 Oedemeridae 787, 895, 901, 916, 943,
Nematoplus 895, 914, 967 Nicagus 831 Notodascillus 841 951, 985, 996
Nemocephalus 910 Nicrophorinae 782, 822 Notomacer 908 Oedemerinae 787, 895
Nemognatha 896 Nicrophorus 822 Notomicrus 808, 921 Oedichirus 829
Nemognathinae 787, 796, 895, 896 Nigidius 832 Notopisenus 888 Oedionychus 907
Nemonychidae 787, 896, 908, 933, Nilio 894 Notoplatypus 913 Oediopalpa 907
959, 996 Nilionidae 892 Notosacantha 907 Oedostethus 854
Nemonychinae 787, 908 Niponiidae 816 Notosalpingus 900 Oenas 896
Nemonyx 908, 933 Niponiinae 782, 816, 943 Notosphindus 873, 969 Oestodes 853
Nemosoma 869 Niponius 816 Nototylus 810 Oestodinae 853
Nemosominae 869 Nipponocis 887 Notoxidae 901 Ogmoderes 881
Neoanthrenus 863 Niptus 864 Notoxii 901 Oiceoptoma 822
Neoathyreus 832 Nipus 884 Notoxinae 902 Oides 907
Neobisnius 830 Nitidula 874 Notoxus 901, 902 Oistus 853
Neobrachypterus 873 Nitidulidae 785, 873, 874, 913, 917, Novelsis 863 Olibrus 877
Neocamiarus 820 924, 943, 945, 947, 950, 954, 957, Novius 884 Oligota 827
Neocercus 878 958, 977 Nucleotops 818 Olisthaerinae 783, 826
Neochlamys 907 Nitidulinae 785, 874, 943 Nyctophila 859 Olisthaerus 826
Neochthebius 818 Nodina 907 Nyctoporus 893 O!otelus 902
Neocimberidinae 908 Nodonota 907 Nyctor 854 Omaliinae 783, 823, 824, 826, 829,
Neocimberis 908 Nodynus 827, 828 Nyctozoilus 893 992, 994, 996, I 000, I 005
Neoclerus 869 Noemia 904 Ochina 864 Omaliine Group 823, 825, 974
Neocrowsonia 852 Nomimocerus 826 Ochodaeidae 783, 833, 988 Omalisidae 784, 837, 856
[32] [33]
Omalisus 856, 926 Ormiscus 909
Ototretinae 784, 859 Palaeoxeninae 784, 851
Omalium 824 Orodotes 852
Oulema 906 Palaeoxenus 851
Omalodes 817 Orophiinae 887
Ovolara 844 Palaestes 876
Omalysidae 856 Orophius 887
Oxacis 895 Palaestra 896
Omalysus 856 Orphilinae 785, 862, 977
Oxelytrum 822 Palaminus 829
Omethes 860 Orphilus 862, 863, 919
Oxychirus 835 Pallodes 874
Omethidae 784, 837, 859, 932, 940 Orphinus 863
Oxycoryninae 788, 909, 972 Palophaginae 787, 905, 960
Omethinae 784, 860, 940 Orphninae 783, 835
Oxycorynus 909 Palophagus 905
Omicrus 816 Orphnus 835
Oxycraspedus 909 Palorus 893
Omma 791, 801, 802 Orsodacne 905, 931, 967
Oxylaemus 881 Palpicornia 813, 814, 920, 981
Omma sagitta 801 Orsodacnidae 787, 905, 906
Oxymastinocerus 858 Pamborus 811, 970
Ommadidae 801 Orsodacninae 787, 905, 955
Oxynopterinae 784, 853 Panagaeus 812
Ommatidae 782, 799, 801, 803, 973 Ortalia 884
Oxynopterus 853 Pangaura 854
Ommatinae 782, 801 Orthaltica 907
Oxyomus 834 Pangauridae 854
Omoglymmius 809, 919-921 Orthocerus 890
Oxypeltidae 903 Pantomorus 912
Omonadus 902 Orthocis 887
Oxypeltinae 787, 904, 941 Pantophaga 799
Omophlinae 893 Orthognathus 912
Oxypeltus 904 Papusus 822
Omophlus 893 Orthogonius 812
Oxypius 828 Parablax 853, 926
Omophron 811, 960, 974 Orthoperidae 885
Oxypoda 827 Paracalais 854
Omophroninae 782, 811 Orthoperinae 885
Oxyporidae 822 Paracalanthis 869
Omophronini 806 Orthoperus 885
Oxyporinae 783, 823, 828, 927 Paracardiophorus 854
Omorgus 832, 988 Orthopleura 870
Oxyporus 828 Paracatops 821
Omosita 874 Orvoenia 874
Oxysternus 817 Paraconosoma 827
Omus 811 Oryctes 836
Oxytelidae 822, 929 Paracucujinae 785, 875
Oncideres 905 Oryctoderus 836
Oxytelinae 783, 823, 828, 829, 948, Paracucujus 875
Oncomera 895 Oryssomus 884
973 Paracupes 803
Oniticellus 834 Oryzaephilus 876
Oxyteline Group 823, 827 Paracymus 816
Onitis 834 Osmoderma 836
Oxytelus 828 Paradrapetes 853
Ontholestes 830 Osoriinae 783, 828, 922, 929, 939, 968,
Ozaena 810 Parafallia 882
Onthophagus 834 973
Ozognathus 864 Parahelopinae 892
Onthophilinae 782, 817 Osorius 828
Pachybrachis 907 Parahelops 891, 892
Onthophilus 817, 958 Osphya 888
Pachycraerus 817 Paraleaster 826
Onysius 892 Osphyinae 786, 888
Pachydema 835 Paralichas 847
Oocyclus 816 Osphyoplesiini 898
Pachyderes 854 Paralimnichus 845, 1004
Oomorphus 907, 956 Osphyoplesius 897, 898
Pachydrus 809 Paralimulodes 819
Oontelus 860 Ostoma 867, 868
Pachymerus 906 Paralispinus 828
Oosternum 816 Ostomidae 867, 984
Pachyochthes 881 Paramecosoma 878
Opetiopalpus 870 Ostominae 795, 868
Pachyplacus 882 Paramellon 894
Opilo 901 Ostomopsinae 786, 881
Pachypodinae 783, 835 Paramelloninae 894
Opisthius 811, 965 Ostomopsis 881
Pachypus 835 Paramellops 894
Opresus 822 Othius 830
Pachyrhynchinae 912 Parandra 904
Ora 839 Othniidae 899
Pachyschelus 843 Parandridae 903
Orchesia 888 Othniinae 787, 899
Pachyteles 810 Parandrinae 787, 904, 999
Orchymontia 818 Othnius 899
Pachyura 909 Paranesoneus 824
Orectochilinae 807 Otidocephalinae 912
Pachyurinae 909 Parapeltis 868, 871
Orectochilus 807 Otiorhynchinae 912
Pactopus 852 Parapeltis Australicum 868, 871
Orectogyrus 807 Otiorhynchus 912
Paederinae 783, 829, 939, 948, 956 Parasemus 877
Orectoscelis 817 Ototreta 859
Paederus 829, 942 Parasiagonum 828
Oreoparnus 844 Ototretadrilinae 784, 859
Paelobius 808 Parasthetops 818
Oritocatops 821 Ototretadrilus 859
Pagomacer 908 Paratenetus 892
[34] [35]

Parathroscinus 845, 1004 Pelonomus 844 Phaenocerus 852 Phloeophilidae 867, 980
Paratillus 870 Peltastica 861, 942 Phaenognatha 835, 915 Phloeophilini 867
Paratropus 817 Peltasticidae 861 Phaenomeridinae 783, 794, 835 Phloeophilus 867
Parentonium 868 Peltasticinae 784, 861 Phaenomerinae 794, 835 Phloeosinus 913
Parepierus 817 Peltidae 867, 932 Phaenomeris 835 Phloeostichidae 785, 875, 932, 962,
Parhemiops 854 Peltinae 785, 795, 867-869 Phaenomerus 835 990
Parhydraena 818 Peltinodes 885 Phaeochrous 834 Phloeostichinae 785, 875
Parmaschema 882 Peltinodinae 786, 885 Phalacridae 786, 877, 945, 976, 988 Phloeostichus 875
Parmulinae 786, 795, 885 Peltis 868 Phalacrinae 786, 877 Phloeotragus 909
Parmulus 885 Peltodytes 807 Phalacrinus 877 Phloeotribus 913
Parnidae 844, 963 Peltonyxa 868 Phalacrognathus 831 Phlogistostemus 869, 870
Parnus 844 Peneta 893 Phalacropsis 877 Phloiophilidae 785, 867, 932
Paromalus 817 Peneveronatus 839 Phalacrus 877, 937 Phloiophilus 867, 932
Paromarteon 898 Penichrolucaninae 982 Phaleria 894 Phobelius 892
Paropsis 907 Penichrolucanus 832 Phanaeus 834 Pholidotus 832
Parosorius 828 Penicillophorus 858, 978 Phanocerus 844 Phoracantha 905
Pasimachus 812 Pentagonica 812 Phanodesta 869 Phosphaenus 859
Passalidae 783, 832, 833, 916, 978, Pentaphyllus 894 Pharaxonotha 879 Photinus 859
984, 988 Pentaplatarthrus 810 Pharaxonothinae 879 Photurinae 784, 859
Passalinae 783, 832 Pentaria 903 Pharinae 884 Photuris 859
Passalus 832 Pentarthrum 913 Pharoscymnus 884 Phreatodessus 809
Passandra 876, 991 Penthe 888 Pharos 884 Phreatodytes 808
Passandridae 785, 867, 876, 926, 991, Penthinae 786, 888 Pheidoliphila 817 Phreatodytidae 808
997 Pentilia 884 Phelister 817 Phreatodytinae 782, 808
Patrobus 812 Pentodon 836 Phellopsis 891 Phreatus 890
Paulusiella 841, 849, 853 Peplomicrus 825 Pheloneis 892 Phrenapates 893
Paussidae 810, 941 Pergetus 901 Pheneps 846 Phrenapatinae 787, 893
Paussinae 782, 810, 813, 927, 966 Pericalus 812 Phengodes 858 Phrepates 893
Paussotropus 812 Pericompsus 812 Phengodidae 784, 837, 841, 857, 858, Phrepatides 893
Paussus 810 Perigona 812 978, 997, 1003 Phrixosoma 913
Paxillus 832 Perileptus 812 Phengodinae 784, 857, 858 Phrixothrix 858
Pectocera 853 Perimylopidae 787, 891, 892, 962,994, Pherocladus 847 Phryganophilus 888
Pectoctenus 904 1000 Pheropsophus 813 Phycosecidae 785, 871
Pectotoma 903, 1004 Perimylops 892, 994 Philagarica 819 Phycosecis 871
Pediacus 876 Periommatinae 913 Phileurus 836 Phyllobaeninae 795, 869, 870, 1003
Pedilidae 897, 898, 900, 928, 980, Periommatus 913 Philharmostes 833 Phyllobaenus 869
1001 Periptyctus 883 Philinae 787, 903, 904 Phylloceridae 851
Pedilinae 787, 898, 899, 914, 1004, Permocupedidae 800 Philonthus 830 Phyllocerinae 784, 851
1005 Perothopidae 837, 851 Philothermopsis 881 Phyllocerus 851, 943
Pedilophorinae 843 Perothopinae 784, 851 Philothermus 881 Phyllopertha 836
Pedilophorus 843, 844 Perothops 851 Philpottia 892 Phyllophaga 835
Pedilus 898, 899, 1001, 1005 Petalium 865 Philus 888, 904 Phyllotocidium 835
Pelecium 812 Petria 893 Phlegon 852 Phyllotocus 835
Pelecophora 871, 999 Petriidae 892 Phlegoninae 784, 852 Phy llotreta 907
Pelecotoma 889, 996 Phaedimus 836 Phloeocharinae 783, 823, 824, 826, Phymaphora 883
Pelecotominae 786, 889 Phaedon 907 828, 829, 948 Phyrrtatophaea 870
Pelidnota 836 Phaennis 891 Phloeocharis 826 Physea 810
Pelobiidae 808, 998 Phaenocephalidae 877, 882, 976 Phloeodes 891 Physemus 845
Pelobius 808 Phaenocephalinae 786, 877 Phloeognathus 826 Physodactylinae 784, 855
Pelochares 845, 978 Phaenocephalus 877, 976 Phloeonomus 824 Physodactylus 855
[36]
[37]
Physognathus 829 Platostomatidae 908
Platycephalus 909 Pocadius 874 Prioninae 787, 904, 999
Physonota 907
Platycerinae 832 Podabrocephalus 860 Prionoceridae 785, 871, 980
Physorhinus 854
Platycerus 832 Podabrus 860 Prionocerites 871
Phytilea 903
Podaena 818 Prionocerus 871, 928
Phytobaenus 902 Platycholeus 821
Podapion 911 Prionochaeta 821
Phytosus 827 Platychoropsis 874
Platycis 857 Pogonoceromorphus 899 Prionocyphon 839
Picnotagalus 893
Pogonostoma 811 Prionomerinae 912
Picrotus 878 Platycrepidius 854
Platydascillinae 786, 880, 994 Pogonus 812 Prionus 904
Pictorus 878
Platydascillus 880 Polemius 860 Prionychus 893
Piesocorynus 909
Piestinae 783, 827-829, 870, 948, 981 Platydema 894 Pollaclasis 859 Pristoderus 890
Platydracus 830 Polycaon 863 Pristolycini 856, 859
Piestoneus 828
Platygenia 836 Polycaoninae 785, 863 Pristolycus 856, 859
Piestus 828
Platylomalus 817 Polycesta 842 Procirrus 829
Piezophyllus 853
Piezotrachelus 911 Platymerus 91 0 Polycestinae 842, 843, 929 Proculus 832
Platynaspis 885, 958 Polychalca 907 Profallia 882
Pilipalpidae 898, 974
Pilipalpinae 787, 898, 980 Platynectes 809 Polyctesis 842 Prognathoides 828
Platynema 874 Polydrosus 912 Prolixocupes 803
Pilipalpus 898, 914
Platynoptera 870 Polygraphus 913 Prolixocupes latreillei 803
Pilolabus 910
Platynopteroi'des 870 Polymerius 841 Prolyctus 881
Pimelia 893
Platynus 812 Polynoncus 832, 988 Prolytta 896
Pimeliinae 787, 893
Polyoptilus 906 Promecoderus 812
Pinodytinae 820 Platyope 893
Platypodinae 788, 913, 925 Polyphaga 779, 782, 798-801, 804, Promecognathus 812, 924
Pinophilinae 829
Platyprosopinae 830 813, 814, 861, 917, 932, 936, 939, Promelittomma 866
Pinophilus 829
Platyprosopus 830 956, 962, 974, 975, 978, 981 Promethis 894
Pinotus 834
Platypsyllidae 819, 821 , 963 Polypria 903 Pronoterus 808
Piseninae 786, 888
Platypsyllinae 782, 794, 821 Pondonatinae 786, 882 Pronus 865
Pisenus 888, 970
Platypsyllus 821, 953, 1003 Pondonatus 882 Propalticidae 786, 877, 879, 933, 955
Pityobiinae 784, 853, 926, 998
Poneridia 907 Propalticus 877, 933, 988
Pityobius 853 Platypus 913
Poophylax 900 Propomacrus 835, 966
Pityomacer 908 Platyrhinidae 908
Platyrhinus 909 Popilia 836 Prosopocoilus 832
Pityophagus 874
Popilius 832 Prospheres 842
Placonotus 877 Platyrhopalus 81 0
Platysoma 817 Porculus 887 Prosteminae 854
Placonycha 846
Platystethus 828 Poria 884 Prosthetopinae 782, 818
Placonychinae 846
Platystomidae 908 Potamonectes 809 Prosthetops 818
Placusa 827
Potamophilus 844 Prostomidae 787, 894, 900, 986
Plaesius 817 Platystomos 909
Platytarsilinae 913 Potergus 852, 929 Prostominia 900
Plagiodera 907
Platytarsulus 913 Potosia 836 Prostominiinae 787, 900
Plagiogramma 817
Plaumanniola 821, 822, 962 Praviclava 882 Prostomis 894, 986
Plagiope 890
Plaumanniolinae 821, 822, 930 Pria 874 Proteininae 783, 824, 956, 994
Plastoceridae 784, 837, 846, 853, 855,
Priacma 802, 803 Proteinus 824, 952
951 Plegaderus 816
Priacma Serrata 802, 803 Protelater 853
Plastocerinae 852 Pleganophorinae 786, 795, 884
Priamima 885, 976 Protelaterinae 853
Plastocerus 853, 855 Pleganophorus 884
Priasilpha 875 Proterhininae 909
Plastocerus Angulosus 855 Pleocoma 830-832, 977
Priasilphinae 785, 875 Proterhinus 909, 915
Platamops 900 Pleocomidae 783, 832, 833
Priastichus 875 Protheca 865
Platerodrilus 856 Pleolobus 841, 993
Priocera 870, 871 Protocoleoptera 800
Plateros 857 Pleonomus 854
Priocerinae 870 Protocucujidae 785, 872, 992
Plateumaris 906 Pleosoma 881
Priocerites 871 Protocucujus 872
Platisus 876 Pleotomus 859
Priochirus 828, 945 Protohylastes 913
Platoberus 879 Plusiotis 836
Priognathus 897 Protomeloe 896, 914, 989
[38] [39]

Protopamus 844, 948 Pseudolichas 841 Pterotinae 784, 859 Pyrochroinae 787, 898, 899
Protopaussus 810 Pseudoliodes 820 Pterotus 859, 934 Pyrogaster 859
Protopeltinae 785, 868 Pseudolotelus 902 Pterotus Obscuripennis 859, 934 Pyrophanes 859
Protopeltis 867, 868 Pseudolycus 895 Pteryngium 878 Pyrophorinae 854, 930
Protoplatypus 913 Pseudomenes 851 Pteryx 804, 818 Pyrophorus 854
Protopristus 829 Pseudomeninae 784, 851 Ptiliidae 782, 814, 817, 818, 918, 920, Pyropinae 912
Protopselaphinae 783, 825, 974 Pseudomicrocara 839 922, 936, 955, 989 Pyrota 896, 989
Protopselaphus 825, 974 Pseudomorpha 812, 813, 965 Ptiliinae 782, 818 Pyrrhalta 907
Protorabinae 806 Pseudomorphinae 782, 812, 813, 917, Ptilineurus 864 Pythidae 787, 892, 896-898, 922, 951,
Protosphindinae 785, 873 935, 970 Ptilininae 785, 865 980, 1000
Protosphindus 873, 926 Pseudonemadus 821 Ptilinus 865 Pytho 897, 980
Protostemum 816 Pseudoperinthinae 827 Ptilium 818 Pyxidicerus 825
Protosthetops 818 Pseudoperinthus 827 Ptilodactyla 847 Quasimus 854
Psammetichus 893 Pseudophengodes 858 Ptilodactylidae 784, 837, 840, 846, Quediinae 830
Psammobius 834 Pseudophengodidae 858 848, 963, 995 Quedius 830
Psammoecinae 876 Pseudophloeocharis 826 Ptilodactylinae 784, 847 Raffrayia 825
Psammoecus 876, 976 Pseudopsinae 783, 828, 829, 948, 973, Ptilophorinae 786, 889 Ragytodes 820
Pselaphacus 880 1005 Ptilophorus 889 Raymondionyminae 912
Pselaphidae 822, 823, 825, 920, 928, Pseudopsis 829, 1005 Ptinidae 792, 794, 821, 864, 920, 930, Rectocomus 901
931, 943, 945, 954, 973-975, 977, Pseudopyrochroa 899 946,949,962,1000 Reichenbachia 825, 978
982 Pseudotelegeusis 857 Ptininae 785, 864 Remigera 880
Pselaphinae 783, 823, 825 Pseudotetralobus 853 Ptinomorphus 864 Renania 880
Pselaphus 825 Pseudotomoderus 902 Ptinus 864, 920, 923 Renardia 828
Psephenidae 784, 831, 837, 840, 843, Pseudotriphyllus 887, 970 Ptomaphaginae 821 Reninus 817
846,861,925,934,949,963 Pseudozaena 81 0 Ptomaphaginus 821 Rentonellum 868
Psepheninae 784, 846 Psilocladus 859 Ptomaphagus 821, 979 Rentonidium 868
Psephenoidea 837, 843, 848, 861 Psiloptera 842 Ptomaphila 822 Rentoniinae 785, 867, 868
Psephenoides 846 Psilopyga 874 Ptomascopus 822 Rentonium 868
Psephenoididae 846 Psoa 864 Ptorthodius 858 Repsimus 836
Psephenoidinae 784, 846 Psoidae 863 Ptosima 843 Rhabdophloeus 877
Psephenops 846 Psoinae 785, 864 Ptyopteryx 804, 983 Rhadalidae 871, 980
Psephenus 846 Psydritae 811 Pulicomorpha 827 Rhadalinae 785, 871, 872, 978
Psepholax 913 Psydius 812 Pulicomorphinae 827 Rhadalus 871
Pseudalaus 853 Psylliodes 907 Pycnocephalus 874 Rhadinosominae 912
Pseudananca 902 Psyllobora 885 Pycnocerinae 892 Rhadinosomus 912
Pseudanidorus 902 Ptenidium 818 Pycnocerus 892 Rhaebus 906, 957
Pseudaulonium 890 Pterocolidae 909 Pycnochila 811 Rhaetulus 832
Pseudenneboeus 887 Pterocolinae 910 Pycnomerinae 786, 890 Rhagium 905
Pseudesarcus 886 Pterocolus 910 Pycnomerodes 890 Rhagodera 890
Pseudeuanoma 856 Pteroderes 891 Pycnomerus 890 Rhagonycha 860
Pseudeucinetus 845 Pterogeniidae 786, 887, 926, 961 Pygocoelis 817 Rhagophthalminae 784, 857, 858
Pseudobothrideres 881 Pterogenius 887 Pygosteninae 827 Rhagophthalmus 857, 858
Pseudochelonarium 847, 848, 969, 978 Pteroloma 819 Pygostenus 827 Rhamna 865
Pseudochodaeus 833, 927 Pterolominae 819 Pylus 870 Rhanidea 883
Pseudochodaeus Estriatus 833, 927 Pteroniinae 824, 825 Pyractomena 859 Rhantus 809
Pseudochrodes 887 Pteronius 825 Pyractonema 859 Rhinomacer 896, 908
Pseudocory1ophidae 882 Pteroptyx 859 Pyrochroa 899 Rhinomaceridae 896, 908
Pseudodactylus 850 Pterosthetops 818 Pyrochroidae 787, 898-901, 914, 922, Rhinomalus 877
Pseudodorcus 832 Pterostichus 812 935, 947, 978, 980, 1000, 1004, Rhinorhipidae 784, 837, 861, 962
Pseudokalissus 825 Pterotarsus 853 1005 Rhinorhipus 861
[40] [41]
Rhinorhynchinae 787, 908 Rhyzobius 884 Scaphisoma 828 Scotocryptus 820
Rhinorhynchus 908 Rhyzodiastes 809, 919 Scaphium 828 Scotodes 895
Rhinosimus 900 Rhyzopertha 864 Scaphobaeocera 828 Scraptia 903, 996
Rhinotia 909 Rimaderus 902 Scaphydra 804, 982 Scraptiidae 787, 889, 903, 941, 996,
Rhipicera 841 Rimulincola 826, 992 Scaptolenus 853 1000, 1004
Rhipiceridae 783, 830, 836, 837, 840, Ripidius 890 Scarabaeidae 783, 834, 915-918, 924, Scraptiinae 787, 903
841, 848, 932, 988 Ripiphorus 889 925, 927, 936, 944, 950, 951, 953, Scraptogetus 902
Rhipidandrus 893 Rodolia 884 954, 959, 968, 977, 978, 984, 989, Scrobicia 863
Rhipidiinae 786, 796, 890 Rodwayia 819 994, 1000, 1003-1005 Scrobifera 810, 812
Rhipidioides 890 Rugilus 829 Scarabaeiformia 783, 814, 830, 831, Scydmaenidae 782, 821, 823, 934, 943,
Rhipidius 890, 920 Rutela 836 836-838, 840 962,987,988
Rhipiphoridae 786, 793, 796, 798, 866, Rutelinae 783, 836, 966 Scarabaeinae 783, 834, 930, 936, 940, Scydmaeninae 782, 822
889, 920, 965, 984, 989, 996, 998, Rygmodus 816 946, 951, 968, 1003, 1006 Scydmaenus 822
999 Rynchophorus 912 Scarabaeoidea 783, 831, 836, 837, 918, Scymnillus 884
Rhipiphorinae 786, 889, 890 Rypobius 885 925, 934, 935, 949, 950, 966, 973, Scymninae 786, 884
Rhipiphorus 889, 890, 965 Rytinotus 890 978, 984, 987, 988, 1006 Scymnus 884, 944
Rhipsideigma 803 Ryzophagus 874, 984 Scarabaeus 834 Secretipes 865
Rhizophagidae 795, 874, 976, 990 Saciinae 795, 885, 978 Scaraphites 812 Sedlacekvia 870, 1002
Rhizophaginae 785, 795, 874 Sacium 885 Scarellus 857 Seirotrana 892
Rhizophagus 874, 991 Sagola 825 Scarites 812, 917 Seladia 885
Rhomboco1eidae 800 Sagra 906 Scaritinae 782, 812 Selasia 856
Rhopa1obrachium 875, 962 Sagridae 906 Scaritini 809, 917 Selonodon 853
Rhopa1ocerus 890, 992 Sagrinae 787, 906, 931, 968, 970 Scelidonta 907 Selvadius 884
Rhopalodontus 887 Salax 893 Schedlarius 913 Semijulistus 871
Rhopalosilpha 862 Salcedia 812 Schinostethus 846 Semiotinae 784, 853, 930, 944
Rhopalosilphinae 862 Salltius 878 Schizophoridae 800, 804 Semiotinus 853
Rhopalotria 909, 972 Salpingidae 787, 899, 974, 993, 1004 Schizophyllus 851 Semiotus 853, 930
Rhynchaeninae 912 Salpinginae 787, 900 Schizopodinae 783, 842 Sepedophilus 826
Rhynchaenus 912 Sandalidae 841, 848, 937 Schizopus 842, 952, 982 Sepidium 893
Rhynchites 910 Sandalus 841,848,931,937 Schizorhina 836 Serangium 884
Rhynchitidae 909 Saphanus 904 Schizotus 899 Serica 835
Rhynchitinae 788, 910, 963, 1000 Saphophagus 865 Scimbalium 829 Sericoderinae 786, 885
Rhynchitoplesius 908 Sapintus 902 Sciodrepoides 821 Sericoderus 885
Rhynchophora 907, 937, 980, 996 Saprininae 782, 816, 817, 982 Scirtes 839 Sericus 854
Rhynchophoridae 911, 1005 Saprinomorphae 816 Scirtidae 783, 794, 831, 836, 838-840, Serrognathus 832
Rhynchophorinae 796, 912 Saprinus 817 861, 862, 981 Serropalpidae 888
Rhynchophorus 912 Saprus 834 Scirtoidea 783, 794, 813, 815, 836-838, Serropalpus 888
Rhyncolus 913 Sarothrias 865, 965 962 Serrotibia 900
Rhyparida 907 Sarothriidae 865, 866 Sclerocyphon 846 Sessinia 895
Rhyparosominae 912 Sarothrocrepis 812 Sclerostomus 832 Setaria 879, 939
Rhyparus 834 Sarrotrium 890 Scolytidae 911, 924, 971, 1003 Setariinae 879
Rhypasma 892 Sartallus 828 Scolytinae 788, 913 Setariola 879
Rhysodes 809, 926 Satorystia 880 Scolytocaulus 893 Setariolinae 786, 879
Rhysodidae 782, 809, 919-921, 924, Saula 883, 985 Scolytocis 887 Shirozuella 884
926, 945, 948, 983, 984 Saxinis 907 Scolytodes 913 Shoguna 874
Rhysopaussidae 892 Scalidia 876 Scolytoplatypus 913 Siagona 811
Rhysopaussus 893 Scaphidiidae 822, 823, 965, 988 Scolytus 913 Siagoninae 782, 811, 938
Rhyssemus 834 Scaphidiinae 783, 828 Scopaeus 829 Siagonium 828
Rhyssonotus 832 Scaphidium 828 Scoriaderma 891 Siamites 822
Rhytirhininae 912 Scaphinotus 811 Scortizus 832 Sicilicula 818
[42] [43]
Sierraclava 844 Sosteamorphus 844
Spilopyra 907 Stenotrachelinae 787, 895
Siettitia 809 Sosylopsis 881
Spiloscapha 894 Stenotrachelus 895, 985
Silinae 784, 860 Sosylus 881, 984
Spodochlamys 836 Stenotrupis 913
Silis 860 Spanglerogyrinae 782, 807
Spondylidae 903, 905 Stenus 829, 981, 982
Silluvia 834 Spanglerogyrus 807, 921, 926, 940
Spondylidinae 787, 796, 904, 905 Stereopalpus 901
Silluviinae 834 Spaniophaenus 878
Spondylinae 796, 904, 905 Stemocera 842
Silpha 822 Sparedropsis 895
Spondylis 904 Stemocoelis 817
Silphidae 782, 819, 822-824, 827, 915, Sparedrus 895
Stadenus 857 Stemodea 878
943, 947, 950, 953, 954, 966, 973, Spastica 896
Stagetus 865 Stemolophus 816
978,979,986,992,1005 Speonomus 821
Staphylinidae 783,817, 819, 822, 823, Steropes 901, 914
Silphinae 782, 822 Spercheidae 815
830,916,917,922,923,927,929, Steropinae 787, 901, 914
Silphomorpha 812, 917 Spercheinae 782, 815
933, 935, 939, 940, 942, 943, 945, Stethopachys 906
Silphopsyllus 821 Spercheus 815
946, 948, 949, 956, 958, 968, Stethorus 884
Silphotelus 824 Sperchopsis 816
971-974, 976, 977, 981, 982, Sticholotidinae 786, 884, 944
Silvanidae 785, 876, 946, 976, 997 Sphaenelater 853
985-987, 989-992, 994, 996-998, Sticholotis 884
Silvaninae 785, 876 Sphaerarthrum 860
1001, 1004, 1005 Stichonotus 811
Silvanolomus 876 Sphaericus 864
Staphyliniformia 781, 782, 813-815, Stictocranius 829
Silvanoprus 876 Sphaeridiidae 815
817,830,831,838,973,975 Stigmodera 843
Silvanus 876, 946 Sphaeridiinae 782, 816, 946, 992
Staphylininae 783, 823, 829, 830, 956 Stilbus 877
Simplicia 809, 810 Sphaeridium 816
Staphylinine Group 823 Stilicopsis 829
Simplicimani 810 Sphaeriestes 900
Staphylinini 830 Stilicus 829
Simplocaria 844 Sphaeriidae 794, 804, 924, 952, 968,
Staphylinoidea 782, 789, 800, 803, Stiliderus 829
Singhikalia 885 982
814, 815, 817, 819, 825, 865, 920, Stirophora 847
Sinocaulus 841 Sphaerites 816, 933, 965, 974
929, 936, 942, 943, 964, 972, 973, Stoliinae 787, 895
Sinodendrinae 831 Sphaeritidae 782, 814-816, 959, 973
977, 987 Stolius 895
Sinodendron 831, 949 Sphaerius 803, 804, 924
Staphylinus 830 Storthodontus 812
Sinoxylon 863 Sphaerocharis 907
Statira 892 Strategus 836
Sipalinus 912 Sphaerocharitinae 907
Stegobium 865 Strepsiptera 798-800, 866, 914, 957,
Sipalus 912 Sphaeropelatops 820
Stenaesthetus 829 981
Sirrhas 892, 962 Sphaerosoma 882
Sphaerosomatidae 795, 882 Stenalia 889 Streptocerus 831
Sisyphus 834
Stenelmis 844 Stromboscerus 912
Sitarida 889 Sphaerosomidae 882
Sphaerostilbus 877 Stenichnus 822 Strongylium 894
Sitaris 896
Steninae 783, 829, 956 Strongylurus 905
Sitona 912 Sphaerothorax 839
Stenocebrio 853, 992 Stygoparnus 844, 918
Sitophilus 912 Sphallomorpha 812, 917, 970
Stenocephaloon 895 Stylifera 810, 812
Sivacrypticus 887 Sphalma 897, 1004
Stenochia 894 Stylopidae 798, 866
Sloanoglymmius 809 Sphargeris 892
Stenochiinae 796, 894 Stylopodinae 829
Smicripidae 785, 874 Sphenophorus 912
Stenocladius 859 Stylopodus 829
Smicrips 874 Sphenoptera 843
Stenocolus 848 Stylosomus 907
Sogda 820 Sphenopterinae 842, 843
Sphindidae 785, 795, 872, 873, 926, Stenocorus 905 Stylulus 812
Sogdiidae 819
Stenodera 896 Styphlomerus 813
Soleniscinae 853 969,983,988,990
Stenoderus 905 Subcoccinella 885
Soleniscus 853 Sphindinae 785, 873
Sphindiphorinae 785, 873 Stenodontes 904 Subprotelater 855
Solieriinae 783, 829
Sphindiphorus 873 Stenomera 864 Subulistomella 838
Solierius 829
Stenosis 893 Sukunahikona 884
Solitarius 882 Sphindociinae 786, 887
Sphindocis 887, 961 Stenostoma 895 Sumnius 884
Somatochaetus 869
Stenotarsia 836 Sunius 829
Somatodinae 912 Sphindophorus 873
Stenotarsinae 795, 883 Suphis 808
Soronia 874 Sphindus 873
Stenotarsus 883 Suphisellus 808
Sostea 844 Spilodiscus 817
Stenotrachelidae 787, 796, 894, 895 Symbiotes 884
 [45]
[44] '"

Tanygnathus 830 Tenerus 870 Thea 885

Symmixus 826
Tanylypa 893 Tenomerga 803 Thecesterninae 912
Symphyomethes 860
Tanymecus 912 Tentyria 893 Therates 811
Syncalypta 844
Syncalyptidae 843, 936 Tanyrhinus 824 Tentyriidae 892, 936 Thermonetus 809
Tanyrhynchinae 912 Tentyriinae 893 Thilmanus 856
Syncalyptinae 783, 844
Tanyrhynchus 912 Teplinus 885 Thinobius 828
Synchita 890
Taphes 857 Terapus 817 Thinodromus 828
Synchroa 894
Taphrocerus 843 Teredilia 861 Thinopinus 830
Synchroidae 787, 894
Taphroderes 910 Teredinae 786, 881 Thinorycter 834
Syndesinae 783, 831
Taphropiestes 878 Teredolaemus 881 Thione 874
Syndesus 831, 949
Tarphius 890 Teredus 881 Thioninae 874
Syndicus 822
Tarsalgus 855, 988 Teretriosoma 816 Thonalmus 856
Synercticinae 787, 897
Tarsosteninae 785, 870 Teretrius 816 Thoracophorus 828
Synercticus 897, 962
Tarsostenodes 870 Termitodiscinae 827 Thoramus 854, 930
Syneta 907, 959, 963, 967
Tarsostenus 870 Termitodiscus 827 Thorictidae 862, 937, 955
Synetinae 907, 963
Tasmosalpinginae 785, 875, 876 Termitotroginae 834 Thorictinae 785, 862
Synhoria 896
• Tasmosalpingus 875, 876 Termitotrox 834 Thorictodes 862, 937
Synochodaeus 833
Tauroceras 893 Tesserocerus 913 Thorictus 862, 955, 983
Synonycha 885
Taurocerastes 833, 951 Tetrabrachinae 884 Thortus 878
Synstrophus 888
Taurocerastinae 833, 1006 Tetrabrachys 884 Thricolema 907
Syntelia 816, 965, 967
Synteliidae 782, 814, 816, 959, 965, Techmessa 898 Tetradelus 824 Thrincopyge 843
967 Techmessinae 898 Tetraglossa 846 Thrincopyginae 842, 843
Syntornium 828 Techmessodes 898 Tetragonoderus 812 Throscidae 784, 837, 851-853, 855,
Syrphetodes 891 Telegeusidae 784, 837, 857, 866, 915, Tetralobinae 784, 853, 961 926, 929, 958, 972, 988
Sysolus 881, 992 1003 Tetralobus 853 Throscidium 819
Systellopodinae 835 Telegeusis 857, 866, 918 Tetraonychidae 895 Throscinae 852
Systellopus 835 Telephanus 876, 997 Tetraonyx 896 Throscinus 845, 1004
Systolosoma 805, 807, 810 Telephoridae 860, 964 Tetraopes 905 Throscus 852, 929
Syzeton 902 Telephorites 871 Tetraphaleridae 801, 976 Thylacosteminae 784, 851 -853
Syzetoninus 902 Telephorus 860 Tetraphalerinae 782, 801 Thylacostemus 853
Szekessya 900 Telmatophilinae 878 Tetraphalerus 801-803 Thylodriadinae 863
Tachinomorphus 826 Telmatophilus 878 Tetraphalerus Wagneri 802, 803 Thylodrias 863, 941
Tachinus 826, 998 Telmatoscius 879 Tetratoma 888 Thylodriidae 862
Tachygoninae 912 Telsimia 885 Tetratomidae 786, 888, 932, 947, 970, Thylodriinae 785, 863
Tachygonus 912 Temnaspis 905, 1005 974 Thymalus 867, 868
Tachyoryctidium 881 Temnochila 869 Tetratominae 786, 888 Thyreocephalus 830
Tachyporinae 783, 824, 826 Temnochilidae 867 Tetrigus 854 Thyreopterus 812
Tachyporine Group 823, 917 Temnochilinae 869 Tetropium 904 Thyroderus 881
Tachyporus 826 Temnopalpus 898 Thallis 879 Tibionema 853
Tachys 812 Temnoscheila 869 Thallisella 879 Tillinae 785, 869
Taeniocerus 832 Tenebrio 893, 956, 1000 Thalycra 874 Tillus 869
Taenioncus 874 Tenebrionidae 787, 792, 887, 891, 892, Thanasimus 870 Timarcha 907
Tagalinus 893 894,897,900,901,914,928,936, Thanatophilus 822 Titaena 893
Talanus 894 943, 947, 956, 968, 969, 977, 984, Thaneroclerinae 785, 869, 930, 941 Todima 890
Talayra 888 994, 995, 1000 Thaneroclerus 869 Tohlezkus 838
Taldycupedidae 800 Tenebrioninae 787, 893 Thaumaglossa 863 Tolmetes 903
Tamotus 829 Tenebrionoidea 786, 880, 886, 903, Thaumaphrastinae 862 Tolyphus 877
Tanainae 910 931,936,962,980,1000 Thaumaphrastus 862, 915, 937 Tomicinae 913
Tanaos 910 Tenebroides 869 Thaumastodinae 784, 845, 993, 1004 Tomicus 913
Tanychilus 893 Tenebroidinae 869 Thaumastodus 845 Tomoderinae 787, 902
[46] [47]
Tomoderus 902, 947, 1001 Trichopseniinae 827, 977 Trogossitidae 785, 867, 868, 992 Urodon 908, 909, 933
Tomoxia 889 Trichopsenius 827 Trogossitinae 785, 869 Urodontidae 908, 966
Toposcopus 889 Trichopterygidae 818 Trogoxylon 864 Urodontinae 788, 908, 909
Toraminae 786, 879 Trichopteryx 818, 947 Tropicus 845 Urodontus 908, 909
Toramus 879, 989 Trichosalpingus 896 Tropideres 909 Urodoplatus 908
Tormissus 816 Tricoleidae 800 Tropidopterus 812 Urophorus 874
Torridincola 803, 804 Tricondyla 811 Tropistemus 816 Uroplata 907
Torridincola rhodesica 803 Tricoryninae 795, 865, 1002 Trotomma 903 Uruminopeplus 900
Torridincolidae 782, 804, 969, 982, Tricorynus 865 Trotommidea 903 Usechimorpha 891
983, 986, 993, 994 Tricrania 896, 977 Trox 832 Usechinae 787, 891
Toxicum 893 Trictenotoma 897, 898, 943 Truncatipennes 810, 813 Usechus 891, 950, 955
Toxidium 828 Trictenotomidae 787, 897, 898 Trycherus 883 Vacronus 893
Toxocerus 834 Trigonocolinae 912 Trypanaeinae 782, 817 Valginae 836
Trachelizus 910 Trigonodemus 824 Trypanaeus 817 Valgus 836
Trachelostenidae 787, 892 Trigonodera 889 Trypeticinae 782, 817 Vatellus 809
Trachelostenus 892 Trigonogenius 864 Trypeticus 817 Vatesus 826
Trachyderas 891 Trigonurinae 783, 822, 823, 827, 828 Trypherus 860 Veraphis 822
Trachyderastes 891 Trigonurus827, 828,922,981,983 Trypobius 817 Veronatus 839
Trachyinae 843 Trilobitideinae 827 Tryponaeinae 817 Vesperidae 903, 904
Trachypachidae 782, 805-808, 811, Trilobitideus 827 Tshekardocoleidae 800 Vesperinae 787, 904
919, 921, 924, 985 Trimenus 874 Tychepsephus 846 Vesperus 904
Trachypachini 811 Trimitomerus 897, 898 Tychiinae 912 Vesta 859
Trachypachus 805, 807, 810, 919, 965 Trimytis 893 Tychius 912 Veturius 832
Trachypholis 890 Trinodes 863 Tychogenius 887 Vicelva 826
Trachys 843 Trinodinae 785, 863 Tydessa 898, 978, 980 Viticinae 912
Trachyscelis 894 Triphyllia 888, 974 Tydessinae 787, 898 Vrilletta 865
Trechinae 782, 811, 812, 954 Triphyllina 887 Tylauchenia 843 Wagneronota 896
Trechitae 811 Triphyllus 887, 970 Tympanogaster 818 Wattianus 887
Trechus 812 Triplacinae 795, 880 Tyndaris 843 Xanthochroa 895
Tretothoracidae 899 Triplax 880 Typhaea 887 Xantholininae 830, 923, 992
Tretothorax 900 Tristaria 864 Typhaeola 887 Xantholinus 830
Triacrus 830 Tritoma 880, 933 Typhlusechus 893 Xanthopyginae 830
Triadogyrus 806 Tritornidae 880, 886, 927, 983 Typhoeus 833 Xanthopygus 830
Triaplus 806 Tritorninae 786, 795, 880, 886 Typophorus 907 Xenaclopus 835
Triarthron 820 Trixagidae 852 Tyrtaeus 894 Xenocryptus 879
Tribalinae 782, 817 Trixagus 852, 926 Tyrus 825 Xenomycetes 884, 955
Tribalus 817 Trochoideinae 795, 884 Tytthaspis 885 Xenomycetinae 786, 884
Tribolium 893 Trochoideus 884, 956 Tytthonyx 860 Xenoscelinae 786, 879
Trichalus 857 Troctontus 903, 944 Uleiota 876, 976 Xenoscelinus 879, 986
Trichananca 901 Trogaster 825 Uleiotinae 795, 876 Xenoscelis 879
Trichelodes 863, 978 Trogidae 783, 831-834, 927, 987, 988 Ulocerus 910 Xenospasta 896
Trichiinae 836 Troglomethes 860 Ulochaetes 905 Xerasia 867, 880
Trichiotinus 836 Troglops 872 Ulodes 891 Xerosaprinus 817
Trichius 836 Trogocryptinae 900 Ulodidae 787, 891, 962 Xestipyge 817
Trichoceble 871 Trogocryptoides 900 Uloma 893 Xestobium 864
Trichochrous 872 Trogocryptus 900 Ulomascinae 912 Xiphoscelis 836
Trichodes 870 Trogoderma 863 Ulomoides 894 Xixuthrus 904
Trichohydnobius 820 Trogophloeus 828 Uloporus 887, 998 Xylariophilinae 786, 881
Trichophya 826, 917 Trogosita 869 Ultrabithorax 799 Xylariophilus 881
Trichophyinae 783, 826, 917 Trogossita 869 Upis 894, 985 Xyleborus 913
[48]

Xyletininae 785, 865 Zeadolopus 820

Xyletinus 865 Zearagytodes 820
Xyletobius 865 Zenoa 841, 848
Xylita 888 Zenodosus 869
Xylobanellus 857 Zenoidae 794, 849
Xylobanus 857 Zeoleusis 828
Xyloctonus 913 Zeonidicola 878, 1000
Xylodes 864 Zeugophora 905, 963
Xylographus 887 Zeugophorinae 787, 905, 963
Xylonaeus 817 Zilora 888
Xylonychus 835 Zilus 884
Xylopertha 863 Zolodininae 787, 892, 893
Xylophilidae 902, 914, 928 Zolodinus 893
Xylophilostenus 903 Zonitidinae 896
Xylophilus 852 Zonitis 896
Xyloryctes 836 Zopheridae 786, 890-892, 894, 936,
Xylothrips 863 962
Xyroscelis 843 Zopherinae 786, 891
Xystropodinae 796, 893 Zopherosis 891
Xystropus 893 Zopherus 891
Yara 804 Zophophilus 894
Ytu 804, 969, 983, 993 Zuphium 812
Zabrotes 906 Zygogramma 907
Zalobius 829, 948 Zygopinae 912
Zarhipis 858, 997 Zygops 912
Zarudniola 841 Zypoetes 893
Zarudniolidae 841 Zyras 827
Zavaljus 879
Zeadelium 892