A Revision of the Superfamily Histeroidea of Japan
(Coleoptera)
1

By Masahiro hara

Otaru Museum,
Ironai 2-1-20, Otaru, 047 Japan


Contents

1. Introduction ------------------------------------------------------------------------ 7
2. Methods ---------------------------------------------------------------------------- 7
3. Historical review ------------------------------------------------------------------ 10
4. Morphology and terms ----------------------------------------------------------- 17
5. Phylogeny -------------------------------------------------------------------------- 34
6. Systematics ------------------------------------------------------------------------- 38
6.1. Family Sphaeritidae --------------------------------------------------------------- 39
6.2. Family Synteliidae ----------------------------------------------------------------- 41
6.3. Family Histeridae ------------------------------------------------------------------ 45
6.3.1. Subfamily Niponiinae ------------------------------------------------------------- 46
6.3.2. Subfamily Chlamydopsinae ------------------------------------------------------ 56
6.3.3. Subfamily Tribalinae -------------------------------------------------------------- 57
6.3.4. Subfamily Onthophilinae --------------------------------------------------------- 60
6.3.5. Subfamily Histerinae -------------------------------------------------------------- 65
6.3.6. Subfamily Hetaeriinae ----------------------------------------------------------- 117
6.3.7. Subfamily Dendrophilinae ------------------------------------------------------ 126
6.3.8. Subfamily Abraeinae ------------------------------------------------------------- 174
6.3.9. Subfamily Saprininae ----------------------------------------------------------- 185
6.3.10. Subfamily Trypeticinae --------------------------------------------------------- 230
7. Acknowledgments --------------------------------------------------------------- 235
8. References ------------------------------------------------------------------------ 236

Enumeration of genera and species

Sphaeritidae ---------------------------------------------------------------------------------- 39
Sphaerites -------------------------------------------------------------- 39
politus Mannerheim --------------------------------------------- 40
Synteliidae ------------------------------------------------------------------------------------ 41
Syntelia ----------------------------------------------------------------- 42
histeroides Lewis ----------------------------------------------- 42
Histeridae ------------------------------------------------------------------------------------- 45
Niponiinae ----------------------------------------------------------------------------- 46
Niponius ---------------------------------------------------------------- 46
furcatus Lewis --------------------------------------------------- 47
impressicollis Lewis -------------------------------------------- 49
itoi Chj -------------------------------------------------------- 51
obtusiceps Lewis ------------------------------------------------ 52
osorioceps Lewis ----------------------------------------------- 53
Chlamydopsinae ---------------------------------------------------------------------- 56
Eucuritopsis ------------------------------------------------------------ 56
japonicus M. hara et Nakane, sp. nov. --------------------- 56
Tribalinae ------------------------------------------------------------------------------ 57
Epierus ------------------------------------------------------------------ 57
lucus Lewis ------------------------------------------------------ 58
uenoi M. hara, sp. nov. -------------------------------------- 58
Onthophilinae ------------------------------------------------------------------------- 60
Onthophilus ------------------------------------------------------------ 60
silvae Lewis ----------------------------------------------------- 60
aonoi M. hara et Nakane ------------------------------------- 61
ostreatus Lewis -------------------------------------------------- 61
niponensis Lewis ----------------------------------------------- 61
ordinarius Lewis ------------------------------------------------ 61
flavicornis Lewis ------------------------------------------------ 62
kamiyai Adachi -------------------------------------------------- 62
Epiechinus -------------------------------------------------------------- 62
arboreus (Lewis) ------------------------------------------------ 62
Histerinae ------------------------------------------------------------------------------ 65
Exosternini --------------------------------------------------------------------- 65
Binhister ---------------------------------------------------------------- 66
chujoi Cooman -------------------------------------------------- 66
Notodoma -------------------------------------------------------------- 66
fungorum Lewis ------------------------------------------------ 66
Hololeptini --------------------------------------------------------------------- 67
Hololepta --------------------------------------------------------------- 67
(Hololepta)
amurensis Reitter ------------------------------------------------ 67
depressa Lewis -------------------------------------------------- 68
higoniae Lewis -------------------------------------------------- 68
Platysomatini ------------------------------------------------------------------ 68
Platysoma -------------------------------------------------------------- 69
(Platysoma)
celatum Lewis --------------------------------------------------- 70
deplanatum (Gyllenhal) ---------------------------------------- 70
lewisi Marseul --------------------------------------------------- 71
rasile Lewis ------------------------------------------------------ 71
takehikoi M. hara --------------------------------------------- 71
tsushimae M. hara -------------------------------------------- 71
vagans Lewis ---------------------------------------------------- 72
unicum Bickhardt ----------------------------------------------- 72
(Platylister)
atratum (Erichson) ---------------------------------------------- 72
cambodjense (Marseul) ----------------------------------------- 75
horni (Bickhardt) ------------------------------------------------ 77
oberthuri (Cooman) --------------------------------------------- 80
pini (Lewis) ------------------------------------------------------ 80
(Cylister)
lineicollis (Marseul) -------------------------------------------- 82
elongatus (Thunberg)** --------------------------------------- 86
(Apobletes)
schaumei Marseul ---------------------------------------------- 86
Eblisia ------------------------------------------------------------------- 88
satzumae Lewis ------------------------------------------------- 88
Histerini ------------------------------------------------------------------------ 89
Hister ------------------------------------------------------------------- 89
impunctatus sawa --------------------------------------------- 90
japonicus Marseul ---------------------------------------------- 91
congener Schmidt ---------------------------------------------- 95
simplicisternus Lewis ----------------------------------------- 98
unicolor leonhardi Bickhardt ---------------------------------- 101
concolor Linne -------------------------------------------------- 102
Zabromorphus -------------------------------------------------------- 106
punctulatus (Wiedemann) ------------------------------------- 106
Merohister ------------------------------------------------------------- 108
jekeli (Marseul) ------------------------------------------------- 108
aino (Lewis) ----------------------------------------------------- 109
uenoi M. hara ------------------------------------------------- 109
Atholus ----------------------------------------------------------------- 109
bimaculata (Linne) ---------------------------------------------- 110
coelestis (Marseul) --------------------------------------------- 110
depistor (Marseul) ---------------------------------------------- 110
duodecimstriatus
quatuordecimstriatus
(Gyllenhal) ------------------------ 110
pirithous (Marseul) --------------------------------------------- 110
Margarinotus ---------------------------------------------------------- 111
(Kurilister)
karbatovi (Tishechkin) ----------------------------------------- 112
(Ptomister)
boleti group
boleti (Lewis) --------------------------------------------------- 112
weymarni group
cadavericola (Bickhardt) --------------------------------------- 112
agnatus (Lewis) ------------------------------------------------- 113
weymarni Wenzel ---------------------------------------------- 113
reichardt Kryzhanovsikj --------------------------------------- 114
striola striola (Sahlberg) --------------------------------------- 114
yezoensis M. hara -------------------------------------------- 115
sutus group
marginepunctatus (Lewis) ------------------------------------- 115
sutus (Lewis) --------------------------------------------------- 115
(Grammostethus)
niponicus (Lewis) ---------------------------------------------- 115
Hetaeriinae ---------------------------------------------------------------------------- 117
Hetaerius --------------------------------------------------------------- 117
gratus Lewis ---------------------------------------------------- 118
otaruensis M. hara, sp. nov. -------------------------------- 120
kubotai M. hara, sp. nov. ----------------------------------- 122
optatus Lewis --------------------------------------------------- 123
Dendrophilinae ----------------------------------------------------------------------- 126
Dendrophilini ------------------------------------------------------------------ 126
Dendrophilus ---------------------------------------------------------- 126
xavieri Marseul ------------------------------------------------- 127
Anapleini ----------------------------------------------------------------------- 129
Anapleus --------------------------------------------------------------- 130
semen (Lewis) -------------------------------------------------- 131
nakanei M. hara, sp. nov. ----------------------------------- 133
nomurai M. hara, sp. nov. ---------------------------------- 135
hagai M. hara, sp. nov. -------------------------------------- 136
Bacaniini ------------------------------------------------------------------------ 137
Bacanius --------------------------------------------------------------- 138
(Mullerister)
niponicus Lewis ------------------------------------------------ 138
(Bacanius)
mikado (Lewis) ------------------------------------------------- 139
Paromalini --------------------------------------------------------------------- 139
Carcinops -------------------------------------------------------------- 140
(Carcinops)
pumilio (Erichson) --------------------------------------------- 140
Australomalus --------------------------------------------------------- 145
montivagus (Lewis) -------------------------------------------- 145
Platylomalus ----------------------------------------------------------- 148
fujisanus (Lewis) ----------------------------------------------- 149
mendicus (Lewis) ---------------------------------------------- 151
niponensis (Lewis) --------------------------------------------- 154
persimilis (Lewis) ---------------------------------------------- 156
viaticus (Lewis) ------------------------------------------------- 159
kusuii M. hara, sp. nov. ------------------------------------- 161
Pachylomalus ---------------------------------------------------------- 163
musculus (Marseul) -------------------------------------------- 163
Eulomalus -------------------------------------------------------------- 166
lombokanus Cooman ------------------------------------------ 166
tardipes (Lewis) ------------------------------------------------ 167
Paromalus -------------------------------------------------------------- 167
omineus Lewis -------------------------------------------------- 168
parallelepipedus (Herbst) -------------------------------------- 169
vernalis Lewis -------------------------------------------------- 171
Abraeinae ----------------------------------------------------------------------------- 174
Abraeini ------------------------------------------------------------------------ 175
Chaetabraeus ---------------------------------------------------------- 175
bonzicus (Marseul) --------------------------------------------- 175
cohaeres (Lewis)* --------------------------------------------- 178
sp. ---------------------------------------------------------------- 179
Plegaderini --------------------------------------------------------------------- 180
Plegaderus ------------------------------------------------------------- 180
marseuli Reitter ------------------------------------------------- 180
shikokensis Hisamatsu ---------------------------------------- 182
Acritini ------------------------------------------------------------------------- 183
Acritus ------------------------------------------------------------------ 183
komai Lewis ---------------------------------------------------- 184
homoeopathicus Wollaston ------------------------------------ 184
Saprininae ---------------------------------------------------------------------------- 185
Gnathoncus ------------------------------------------------------------ 185
nannetensis (Marseul) ------------------------------------------ 186
rotundatus (Kugelann) ----------------------------------------- 190
communis (Marseul)* ------------------------------------------ 192
Saprinus ---------------------------------------------------------------- 195
(Sprinus)
cyaneus auricollis Marseul ------------------------------------ 196
niponicus Dahlgren --------------------------------------------- 199
pecuinus (Marseul) --------------------------------------------- 202
planiusculus Motschulsky ------------------------------------- 202
splendens (Paykull) -------------------------------------------- 205
Hypocacculus --------------------------------------------------------- 209
(Nessus)
asticus (Lewis) -------------------------------------------------- 209
Hypocaccus ------------------------------------------------------------ 211
(Hypocaccus)
ainu (Lewis) ----------------------------------------------------- 212
axeli Kryzhanovskij -------------------------------------------- 213
lewisii (Schmidt) ----------------------------------------------- 215
sinae (Marseul) ------------------------------------------------- 217
subaenus (Schmidt) -------------------------------------------- 220
akanensis M. hara, sp. nov. -------------------------------- 222
(Baeckmanniolus)
varians varians (Schmidt) ------------------------------------ 224
varians hatsune (Nakane) -------------------------------------- 227
Eopachylopus --------------------------------------------------------- 228
ripae (Lewis) ---------------------------------------------------- 228
Trypeticinae -------------------------------------------------------------------------- 230
Trypeticus -------------------------------------------------------------- 230
fagi (Lewis) ----------------------------------------------------- 231
venator (Lewis) ------------------------------------------------- 233
* Newly recorded from Japan.
** Introduced to Japan.


1. Introduction

The superfamily Histeroidea is composed of three families, Sphaeritidae, Synteliidae and Histeridae. The Sphaeritidae and the Synteliidae are small families, comprising 3 and 5 known species, respectively. The Histeridae or histerid beetles form a fairly large group, containing about 3800 described species, and are broadly distributed over the world. The vast majority of the beetles are predators and found in most kinds of decaying organic matter, such as droppings, decomposing bodies of animals, compost piles and other decaying plant materials. Some species of histerid beetles have been well known as predacious cadavericolous insects controlling populations of dipterous flies and other insects (Summerlin et al., 1981, 1982, 1984, 1989a, b; Bornemissza, 1968).
In spite of their importance in biological control in nature, the histerids had long been neglected in Japan except in catalogues and faunal insect lists until modern revisions of some tribes and genera started to appear in the late 1980s (hara and Nakane, 1986, 1989; hara, 1989, 1991a, b, 1992a, b, 1993a). The present review is therefore the first comprehensive study of the superfamily in Japan; 115 species are dealt with taxonomically, with scattered pieces of biological and distribution information summed up. The Japanese species have generally been difficult to identify. It is hoped that this review makes them easily identifiable and stimulates further studies on their habitats, life histories, distributions and applied importance.

2. Methods

General. This study is based mainly on dried specimens collected by me during 1980 - 1991 and by Dr. Takehiko Nakane; many further pinned specimens were borrowed from other institutions listed under Museum acronyms.
In this study were examined about 3,000 specimens from Japan and, in addition, about 300 specimens from Taiwan, 100 from Europa, and 100 from the rest of the world. I tried to examine the types of some Japanese species, especially of problematic species, and have succeeded to borrow the types of the following species: Plegaderus shikokensis, Hypocaccus vrians hatsune, Anapleus semen, A. japonicus, and Hister simplicisternus.
Specimens were collected under carrion or dung, or often in bait traps using chicken, or sometimes under bark.
General observations and dissection were carried out under the stereoscopic microscope Olympus SD (magnification: up to x160) and Olympus SZH-131 (magnification: up to x128). Some structures were also observed in a SEM (Hitachi S-2000A).
Genitalia were removed from dried specimens, and then 1) heated in 10 % KOH at 60C for about 1 hour or more according to the size; 2) washed and dissected in 70% ethyl alcohol (remaining muscles are removed), 3) transferred into lactic acid containing acid fuchsin and heated at 60C for 3 hours, 4) transferred into a mixture of glacial acetic acid 1 part and methyl salicylate 1 part and left there for 30 minutes, and 5) observed in a-terpineol in a small glass dish.
Description format. For each described species, bibliographic and nomenclatural information is given first. Description includes measurements of body length and width (Fig. 1). Measurements of some body parts, with mean, ranges, standard error and sample size, all in mm, are given in tables. Whenever possible, at least 20 specimens were measured for each sex. Abbreviations used in the measurements are as follows:
PPL: length between anterior angles of pronotum and apex of pygidium.
PEL: length between anterior angles of pronotum and apices of elytra.
HOW: width between hones (projections) of epistoma.
HW: width of head.
APW: width between anterior angles of pronotum.
PPW: width between posterior angles of pronotum.
PL: length of pronotum along mid line.
EL: length of elytron along sutural line.
EW: maximal width between outer margins of elytra.
ProW: maximal width of propygidium.
ProL: length of propygidium.
PyL: length of pygidium.
PTL: length of protibia.
MSTL: length of mesotibia.
MTTL: length of metatibia.
Museum acronyms. Specimens were borrowed from the following institutions. The people responsible for the loans are given after the names of the institutions.
BSM Bishop Museum, Honolulu; G. A. Samuelson.
EIHU Entomological Institute (now Laboratory of Systematic Entomology), Hokkaid University, Sapporo; M. Suwa.
ELKU Entomological Laboratory, Kysh University, Fukuoka; O. Tadauchi.
HFFP Hokkaid Research Center, Forestry and Forest Products Research Institute: K. Maet.
IJ Collection of Mr. K. Ijima, Shibecha, Hokkaid.
NA Collection of Dr. T. Nakane, Chiba.
NHM Natural History Museum, London; E. De Boise.
NSMT Natural Science Museum, Tky; S.-I. Uno and A. Shinohara.
Survey areas. The present study covers the islands and regions shown in Table 1 and Figure 2. The Japanese suffixes -ken, -to, and -fu are applied to prefectures.


Table 1. Names of Islands and regions in Japan under investigation.
See Fig. 2.
___________________________________________________________________
No. Main Islands, Islands or prefectures,
put in braces [ ] put in brackets < >
in Specimens examined. in Specimens examined.
___________________________________________________________________
I Chishima Isles. Etorofu Is., Kunashiri Is., Shikotan Is., Habomai Isles.
II-1 Hokkaid proper
II-2 Rishiri Is., Rebun Is. (located close to Wakkanai)
II-3 Teuri Is., Yagishiri Is. (located close to Haboro)
II-4 Okushiri Is. (located close to Setana)
II-5 Oshima Isles. (located close to Matsumae)
III-1-a Honsh Aomori-ken ...........................
III-1-b Iwate-ken
III-1-c Miyagi-ken Thoku district
III-1-d Akita-ken
III-1-e Yamagata-ken
III-1-f Fukushima-ken ............................
III-1-g Ibaraki-ken ............................
III-1-h Tochigi-ken
III-1-i Gumma-ken
III-1-j Saitama-ken Kant district
III-1-k Chiba-ken
III-1-l Tky-to
III-1-m Kanagawa-ken .............................
III-1-n Niigata-ken .............................
III-1-o Toyama-ken
III-1-p Ishikawa-ken
III-1-q Fukui-ken
III-1-r Yamanashi-ken Chbu district
III-1-s Nagano-ken
III-1-t Gifu-ken
III-1-u Shizuoka-ken
III-1-v Aichi-ken ............................
III-1-w Mie-ken ............................
III-1-x Shiga-ken
III-1-y Kyto-fu
III-1-z saka-fu Kinki district
III-1-aa Hygo-ken
III-1-bb Nara-ken
III-1-cc Wakayama-ken .............................
III-1-dd Tottori-ken .............................
III-1-ee Shimane-ken
III-1-ff Okayama-ken Chgoku district
III-1-gg Hiroshima-ken
III-1-hh Yamaguchi-ken .............................
III-2 Izu Isles.
III-3 Sado Is., Awa-jima Is.
III-4 Hegura Is.
III-5 Oki Isles.
IV-1-a Shikoku Tokushima-ken
IV-1-b Kagawa-ken
IV-1-c Ehime-ken
IV-1-d Kchi-ken
V-1-a Kysh Fukuoka-ken
V-1-b Saga-ken
V-1-c Nagasaki-ken
V-1-d Kumamoto-ken
V-1-e ita-ken
V-1-f Kysh Miyazaki-ken
V-1-g Kagoshima-ken
V-2 Tsushima Is.
V-3 Iki Is.
V-4 Got Isles.
V-5 Amakusa Isles.
V-6 Koshiki Isles.
VI-1-a Nansei Isles. Yaku-shima Is.
VI-1-b Tane-ga-shima Is.
VI-1-c Tokara Isles.
VI-1-d Amami-shima Is.
VI-1-e Tokuno-shima Is.
VI-1-f Okinoerabu Is.
VI-1-g Yoron Is.
VI-1-h Okinawa-Hont Is.
VI-1-i Kume-jima Is.
VI-1-j Kerama Isles.
VI-1-k Miyako Is.
VI-1-l Tarama Is.
VI-1-m Ishigaki Is.
VI-1-n Iriomote Is.
VI-1-o Hateruma Is.
VI-1-p Yonakuni Is.
VII Ogasawara Isles. Haha-jima Is., Chichi-jima Is.
___________________________________________________________________


3. Historical review

Classification of the family Histeridae
In the following lines main prominent studies in the higher classification of the family Histeridae are summarized. Detailed reviews are found in Marseul (1857), Kryzhanovskij and Reichardt (1976), and Vienna (1980).

1758. Linn described the genus Hister for 6 new species.
1811. Paykull proposed a new genus, Hololepta. He firstly classified the histerids into two genera, Hister and Hololepta.
1817. Leach added four genera, Platysoma, Dendrophilus, Abraeus and Onthophilus.
1834. Erichson divided the family into 3 major groups, and 2 of them into 5 subgroups, as follows:
I: Hololepta, Phylloma, Oxysternus
II: A. a) Plaesius, Placodes, Platysoma, Omalodes, Cypturus
b) Hister, Hetaerius, Epierus, Tribarus
B. Dendrophilus, Paromalus
III: A. Saprinus, Pacylopus, Trypaneus
B. Teretrius, Plaegaderus, Onthophilus,
Abraeinae
He used the following character states in the classification: 1. head retractile or not (II, III) (I); 2. prosternal lobe present or absent (II) (III); 3. club of antenna in repose is situated on anterior margin (IIA) or on middle of surface (IIB) of prosternum; 4. articulation of club of antenna is distinct on both ventral and dorsal sides (a) or not distinct on one or both sides (b); 5. antennae articulate on margin of front of head (IIIA) or on frontal disk (IIIB).
1854. Lacordaire recognized two tribes and two subtribes.
I: Hololeptides
II: Histerides
(i) Histerides
(ii) Saprinides
His system is based on the following character states: 1. head retractile or not (II) (I); 2. prosternum with prosternal lobe or without it (i) (ii).
1857. Marseul proposed a new classification of the family.
I: Hololeptiens
II: Trypanens
III: Histriens
IV: Htriens
V: Sapriniens
VI: Abrens
His classification consisted of 6 tribes. He used the following characters to divide the family into the tribes: 1. head retractile or not (I, II) (III, IV, V, VI); 2. mandible long or short (I) (II); 3. prosternal lobe present or not (III, IV) (V, VI); 4. articulation of club of antenna is clearly distinct or not, and shape of the club is oval or cylindrical (III) (IV); 5. antennal fossette situated on disk or margin of prosternum, and antennal furrow of head present under the lateral margin of head or in emargination on the anterior margin (V) (VI).
1899. Ganglbauer almost agreed with Marseul (1857), but he did not distinguish between the Histerinae and the Hetaerinae and the latter was included in the former. He provided four tribes for European histerids, and did not mention the tribe Trypanaeini.
1909. Reitter divided th family into 7 tribes.
I: Hololeptini
II: Histerini
III: Paromalini
IV: Dendrophilini
V: Hetaerinini
VI: Saprini
VII: Abraeini
1916-17. Bickhardt's classification.
I: Hololeptinae
II: Trypanaeinae
III: Trypeticinae
IV: Teretriinae
V: Abareinae
VI: Saprininae
VII: Dendrophilinae
VIII: Histerinae
1. Tribalini
2. Platysomini
3. Histerini
4. Exosternini
IX: Hetaeriinae
1. Hetaeriomorphini
2. Hetaeriini
3. Chlamydopsini
1941. Reichardt's classification was as follows:
I: Niponiinae
II: Trypeticinae
"II": Trypaeinae
III: Teretriinae
IV: Abraeinae
V: Saprininae
VI: Dendrophilinae
VII: Hololeptinae
"VII": Histerinae
"X": Hetaerinae
In this work the Trypaeinae, Histerinae and Hetaerinae are erroneously numbered. He provided a key to the 10 subfamilies, which is quite artificial and does not reflect his classification. In his "Klassifikation" (p. 346) the following notes are given: 1) the genus Niponius is a member of the family Histeridae, because the paramera of the male genitalia are free and similar to those of the Trypeticinae and Trypaeinae, representing a primitive condition [in my observation, however, the paramera of these taxa are not free]. 2) The retractile or non-retractile condition of the head has been traditionally regard as a primarily important character state in the family, but Reichardt did not adopt this view. According to him the retractile head originated independently in some groups, e. g., in Spelaeacritus, Hololepta and the Trypaeinae. The first genus is allied to the Acritus-group; Hololepta is related to the Histerinae (therefore, the subfamily Histerinae was placed next to the Hololeptinae; in fact, the male genitalia and the hind wing vein are very similar between them); the Trypaeinae close to the Trypeticinae. 3) The Saprininae is highly specialized group and allied to the Abraeinae and Dendrophilinae based on many characters (however, no detailed explanation was given by him).
1944. Wenzel proposed a new system, which remarkably differs from the traditional classification. He divided the Histeridae into two divisions, "Saprinomorphae" and "Histeromorphae", and changed the ranks of some taxa as follows: 1) The Hololeptinae (sensu Reichardt, 1941) was reduced to a tribe in the Histerinae. 2) The Chlamydopsinae was treated as a subfamily and placed next to the Abraeinae. 3) The Teretriinae (sensu Reichardt, 1941) is ranked as a tribe in the Abreinae. 4) The tribe Tribalini included in the Histerinae (Bickhardt, 1916) was risen in rank to the subfamily Tribalinae. His system has been adopted by most recent authors. See also Section 5 (Phylogeny) for a detailed discussion. His system is summarized as follows:
I: Saprinomorphae
I: Chlamydopsinae
II: Abraeinae
1: Abraeini
2: Plegaderini
3: Acritomorphini
4: Acritini
5: Teretriini
III: Niponiinae
IV: Trypanaeinae
V: Trypeticinae
VI: Sapriniae
II: Histeromorphae
VII: Dendrophilinae
VIII: Tribalinae
IX: Histerinae
1: Histerini
2: Omalodini
3: Platysomini
4: Hololeptini
5: Exosternini
X: Hetaeriinae
1: Hetaeriomorphini
2: Hetaeriini
1976. Kryzhanovskij and Reichardt proposed a new taxon, the tribe Bacaniini.
1980. Vienna elevated the Onthophilina (Thomson, 1862) to the subfamily rank.
1982. Olexa proposed a new tribe, the Anapleini.
1984. Mazur is mainly followed Wenzel. See also in the Section 5 (Phylogeney).
I: Saprinimorphae
I: Abraeinae
1. Abrareini
2. Plegaderini
3. Acritomorphini
4. Acritini
5. Teretriini
II: Niponiinae
III: Trypanaeinae
IV: Trypeticinae
V: Saprininae
VI: Chlamydopsinae
II: Histeromorphae
VII: Dendrophilinae
1. Dendrophilini
2. Anapleini
3. Bacaniini
4. Paromalini
VIII: Onthophilinae
IX: Tribalinae
X: Histerinae
1. Histerini
2. Omalodini
3. Platysomatini
4. Hololeptini
5. Exosternini
XI: Hetaeriinae
1989. Helava. Tribes in the Hetaerinae were not discriminated.


Japanese Histeroidea
1854. Marseul described a new species, Hister japonicus. .......... +1+0=12
1860. Motschulsky described a new species, Hister japanus, which was later synonymized under Margarinotus striola. ........................... +1+0=2
1873. Marseul described 7 new species, Platysoma lewisi, P. lineicolle, Hister pirithous, H. depistor, Dendrophilus xavieri, Paromalus musculus (=Pachylomalus musculus), and Abraeus bonzicus (=Chaetabraeus bonzicus), and newly recorded from Japan H. jekeli, H. 14-striatus (=Atholus duodecimstriatus quatuordecimstriatus), H. punctulatus, H. cadaverinus (= Margarinotus weymari or its allied species), H. navus (= Margarinotus niponicus), Carcinops pumilio, Saprinus speciosus (= S. splendens), S. pecuinus, S. nitidulus (= S. planusculus), S. sinae and Saprinus (Gnathoncus) rotundatus. ............................... +7+11=20
1877. Reitter described a new species, Plegaderus marseuli. .......... +1+0=21
1878 Harold recorded a species, Onthophilus striatus. ................ +0+1=22
1879. Lewis described four new species, Hister marginepunctatus, Hister simplicisternus, Bacanius niponicus, and Acritus komai. ....... +4+0=26
1882. Lewis erected a new genus for a new species, Syntelia histeroides. ........................................................................... +1+0=27
1884. Lewis described twenty-one new species, Hololepta depressa, H. parallela (nec Sturm, 1868), Platysoma pini, P. vagans, P. rasile, P. celatum, Hister aino, H. concolor, H. boleti, H. agnatus, H. sutus, Epielus lucus, Notodoma fungorum, Hetaerius gratus, H. optatus, Triballus semen, Triponaeus fagi, T. venator, Onthophilus flavicornis, O. silvae, and O. arboreus (=Epechinus arboreus), and newly recorded Hololepta amurensis and Onthophilus ostreatus. Onthophilus flavicornis is, however, the same with O. striatus in Harold (1879). .................. ...................................................................... +21(+2-1)=49
1885. Schmidt described a new species, Hister congener. ............. +1+0=50
1885. Lewis described a new species, Pachylopus ripae. .............. +1+0=51
1885. Lewis described a new genus Niponius for four new species, impressicollis, furcatus, obtusiceps, and osorioceps. ............ +4+0=55
1888. Lewis described a new species, Paromalus persimilis. .......... +1+0=56
1890. Schmidt described Saprinus lewisii, S. subaenus, and S. varians. Later, they were transferred to Hypocaccus. ....................... +3+0=59
1892. Lewis described Paromalus mendicus, P. viaticus, P. fujisanus, P. vernalis, P. tardipes, P. omineus, P. montivagus, Acritus shogunus (=A. homoeopathicus), and Abraeus mikado, and recorded Paromalus parallelepipedus and P. complanatus; the last was, however, corrected later by himself (see 1899). ........................................... +9+2=70
1893. Schmidt transferred Abraeus mikado into the correct genus Bacanius.
1894. Lewis proposed a new name, higoniae, for his Hololepta parallela.
1895. Lewis described a new species, Hister niponicus, which was recorded as H. navus by Marseul (1873). ..................................... +1-1=70
1899. Lewis described three new species, Platysoma satzumae, Hypocaccus aniu, and Paromalus niponensis, the last for the misidentified P. complanatus in Lewis (1892), and synonymized Hister japanus under H. striola. ............................................................... +2-1=72
1904. Lewis transferred Hister congener into the new genus Pachylister.
1905. Lewis transferred Tryponaeus venator and fagi into the correct genus Trypeticus.
1906. Lewis described a new species, Platylister niponensis. ......... +1+0=73
1906b. Lewis recorded a species, Platysoma deplanatum. .............. +0+1=74
1907. Lewis transferred Paromalus parallelepipedus, P. omineus, and P. vernalis into the new genus Microlomalrus.
1907b. Lewis described a new species, Onthophilus niponensis. ....... +1+0=75
1911. Lewis described a new species, Hypocaccus asticus. ........... +1+0=76
1914. Lewis described a new species, Hister togoii. .................... +1+0=77
1918. Bickhardt treated Hister aino as a synonym of jekeli. .......... +0-1=76
1920. Bickhardt treated Hister togoii as a synonym of H. simplicisternus. ............................................................................. +0-1=75
1920. Bickhardt treated Platysoma niponensis as a synonym of P. cambojense.
1926. Reichardt transferred Pachylopus ripae into the new genus Eopachylopus.
1930. Adachi described a new species, Onthophilus kamiyai. ......... +1+0=76
1941. Cooman described a new species, Binhister chujoi. ............. +1+0=77
1944. Wenzel described a new species, Margarinotus weymarni, which may be the same with Hister cadaverinus in Marseul (1873), and transferred Hister agnatus, H. marginepunctatus, H. niponicus, H. sutus, H. striola, and H. boleti into the new genus Margarinotus. ................... +1-1=77
1948. Cooman described the new genus Platylomalus, and transferred Paromalus mendicus to it.
1948. Cooman described a new species, Platysoma oberthuri. ........ +1+0=78
1952. ssawa described a new species, Hister impunctatus and a new form, rufofasciatus, of Eopachylopus ripae. ............................. +1+0=79
1955. Chjj described a new species, Niponius itoi. ................... +1+0=80
1961. Nakane described a new species, Anapleus japonicus. .......... +1+0=81
1962. Dahlgren described a new species, Saprinus niponicus. ........ +1+0=82
1962. Adachi and Ohno newly recorded a species, Spaherites politus. ............................................................................ +0+1=83
1963. Nakane newly recorded two species, Peranus bimaculatus and Platysoma unicum. .................................................... +0+2=85
1964. Nakane transferred Triballus semen into the correct genus Anapleus.
1976. Kryzhanovskij and Reichardt described a new species, Hypocaccus axeli, transferred Paromalus niponensis, P. fujisanus, and P. viaticus into the correct genus Platylomalus, and newly recorded Hister unicolor opimus (= leonhardi). ................................................ +1+1=87
1977. Nakane described a new subspecies, Hypocaccus varians hatsune and newly recorded Saprinus auricollis (=S. cyaneus auricollis) from Bonin Is. ........................................................................ +0+1=88
1984. Mazur described a new genus, Australomalus, and transferred Platylomalus montivagus to it.
1984. Mazur transferred Paromalus persimilis into the correct genus Platylomalus, and Paromalus tardipes into Eulomalus. Anapleus japonicus is synonymized under A. semen. ........................ +0-1=87
1984. Hisamatsu and Kusui newly recorded Atholus coelestis. .... +0+1=88
1984. Kusui recorded Cylister elongatus. Because this species is undoubtly an introduced species, it is excluded from the fauna.
1985. Hisamatsu described a new species, Plegaderus shikokensis, and newly recorded Apobletes tener. ............................................. +1+1=90
1985. Hisamatsu newly recorded Gnathoncus nannetensis, Eulomalus lombokanus, Platylister atratus and P. horni. ..................... +0+4=94
1986. hhara revised the genus Platysoma from Japan, and added two new species, takehikoi and tsushimae. .................................. +2+0=96
1986. hhara and Nakane revised the genus Onthophilus from Japan, and added a new species, aonoi, and newly recorded ordinatus. ............... ............................................................................ +1+1=98
1987. Mazur synonymized Acritus shogunus under Acritus (Pacnacritus) homoeopathicus.
1989. hhara and Nakane redescribed two species of the tribe Exosternini, Notodoma fungorum and Binhister chujoi.
1989. hhara reviewed the genus Margarinotus from Japan, described a new species, M. (Ptomister) yezoensis, and newly recorded two species, cadavericola and reichardti. M. (P.) marginepunctatus was transferred form subgenus Promethister to subgenus Ptomister. ........... +1+2=101
1991. hhara revised the genus Hololepta from Japan.
1992. hhara revised the genus Merohister from Japan, revitalized aino, and added a new species, uenoi. ........................................ +1+1=103
1992. hhara revised the genus Atholus from Japan.
1993. hhara recorded a species, Margatinotus (Ptomister) karbatovi. .......................................................................... +0+1=104
1994. In this paper, I describe 9 new species, Hypocaccus akanensis, Eucuritopsis japonicus, Anapleus nakanei, A. nomurai, A. hagai, Platylomalus kusuii, Epierus uenoi, Hetaerius otaruensis, H. kubotai and newly recorded from Japan Chaetabraeus cohaeres and Gnathoncus communis. ........................................................... +9+2=115
In conclusion, 115 species of the Histeroidea are now known to occur in Japan.


4. Morphology and terms

Because of the diversity in external structure of Coleoptera, descriptive terms often need to be clarified for each superfamily, family, or even lower category under study. In the Histeroidea, many papers in English published after 1950 follow the style and descriptive terms in Wenzel and Dybas (1936). The present paper also mainly follows Wenzel and Dybas's system of terms. Most of the terms used are illustrated in Figure 3.

Head
No detailed comparative morphological study of the head has been made for members of the superfamily Histeroidea. Because in dried specimen the head is usually retracted in the prothorax, many past authors, except Marseul (1853 - 1862), did not describe the details of the head sutures and mouthparts, which are observable on the ventral side.
In this section the head sutures and various parts of the cranium are dealt with.

Situation and general appearance of head
Head is generally prognathous in the Coleoptera. In the families Sphaeritidae and Synteliidae and the subfamily Niponiinae of the Histeridae, the head is prognathous as in many other Coleoptera. But in other Histeridae the head is almost deflexed except in several members (e. g., Trypanaeinae and Hololepta). Usually the posterior half of the head retracts into the anterior margin of the pronotum. The states of the prognathous in the several members is probably secondary in association with a changed living style.

Head in dorsal view is short, broad, convex, and oval or oblong-oval in general. In Trypeticus (Fig. 5E, F) the head shows a triangular shape in dorsal view, the lateral margins being convergent apically. In the majority of Histeridae, the anterior half of the head is generally narrowed to form the frontoclypeal region.
Head is usually narrower than the prothorax except in the Synteliidae and Niponius, in which the head is as broad as the prothorax.

Head suture (Fig. 4 - 6)
In the Histeroidea, the following sutures are observed on the head (Fig. 4A, B): frontoclypeal, hypostomal, occipital and postoccipital suture; midcarinal suture (cornal suture of Blackwelder, 1936) absent.

Frontoclypeal suture (clypeofrontal suture in Cook, 1943)
This suture runs arcuately between the anterior tentorial pits in Coleoptera, but the pits are obscure in the Histeroidea. The suture is present only in the Sphaeritidae; in Shpaerites the suture is very distinct and gently arcuate (Fig. 4C).

Occipital suture (infraorbital ridge of Smetana, 1971; premandibular suture of Cook, 1943)
This suture crosses dorsally the hind part of the head, continues to the ventral side and terminates on each side anteriorly to the posterior articulation of the mandible; it demarcates the vertex from the occipital region in generalized Coleoptera, for example, a group of Adephaga (Cook, 1943) and the Staphilinidae (Naomi, 1987). In the Histeroidea, the suture is lost in the majority, and is found only in Epielus as a short one along the posterior margin of each eye (Fig. 6D).

Hypostomal and postoccipital sutures
The hypostomal and postoccipital sutures are united with each other at the posterior tentorial pit to form a line in general Coleoptera. In the Histeroidea, however, it is usually difficult to discriminate between the two sutures, because the pit is absent in major groups and obscure in Spaherites and Syntelia (Fig. 4D, G). Part of the postoccipital suture between the gula and postgena is usually called "gular suture".
The hypostomal suture is distinct and found in the majority of the Histeroidea.
The gular sutures are variable in the Histeroidea: sometimes they are separated, running parallel to each other, but in other species they are fused partly or even completely to form a straight line. See also under Gula plate.
The postoccipital suture between the postocciput and postgena is variable in the Histeroidea, and is discussed under Postocciput.

Eyes and ocelli (Fig. 4 - 6)
The eyes are present in a pair and located laterally in general. They are usually well developed and oblong, the posterior margin is often inwardly arcuate. They are usually kidney-shaped or round in lateral view.
The eyes are, however, lost in a cave-inhabiting species, Geocollus caecus Wenzel.
The occeli are absent in the Histeroidea.

Cranium (Fig. 4 - 6)
Cranium is usually composed of the frontoclypeal region, vertex, postgena, ventral plate of the parietals, postocciput and gula in the Histeroidea.

Frontoclypeal region
In the majority of the Histeroidea, the region is nearly square, its lateral margins being often gradually convergent apically, or parallel. This is not ture of the Histerinae (Fig. 6F, I) and Paromalini (Fig. 5C), in which the region is short with the anterior margin gently arcuate.

Epicranial modification
Epicranium is usually weakly convex or subflat. In the Niponiinae, there are projected structures in a pair, which are well developed and corrugated (Fig. 42A, 43A, 45A, 46A). Hornlike structure is developed in some species of the subfamily Tribalinae. The majority of the Onthophilinae have costae.
Frontal and supraorbital striae on the disk of epicarum are useful character for classifying the Histeroidea. The frontal stria usually appears in the Saprininae, Tryptecitinae, Dendrophilinae, and Histerinae; it is variable from a feebly impressed to a deep one, often interrupted at anterior middle or latero-apical angles, and also from a gently arcuate to a hexagonal line. The supraorbital stria is found in the Saprininae (Fig. 5G). The posterior end of the frontal and the lateral end of the surpraorbital are usually united with each other.

Occiput
Occiput is equal to the so-called "neck" in Coleoptera. In the Histeroidea the occipital constriction is absent. Only in Trypeticus the portion is separated by a distinct ridge (Fig. 5E, F).

Ventral plates consisting of postgenae and ventral parts of parietals
The postgena and ventral part of the parietal on one side are usually fused into a plate because of the absence of the occipital suture. The plate are weakly convex or flat, usually with small or large punctures. However, a modified structure, known as antennal furrow, is found in major groups.
Antennal furrows on the ventral plates are developed in the following groups: Syntelia, Pachylomalus, Onthophilus, Hololepta (Fig. 4G, 5D, 6B, J) and Niponius (Fig. 42B, 43B, 45B, 46B); in the last taxon they are deeply foveate.

Gular plate
Gula is a median ventral plate of the head behind the posterior tentorial pit, but it is not separated from the submentum by a suture. In the Histeroidea, the gula is important for phylogenetic study, and is classified into the following types on the basis of the shape.
1. Gula is trapezoid. The gular sutures are distinct in a pair, not fused with each other, and are convergent apically.
2. Gula is a small triangular plate. The gular sutures are fused on anterior part.
3. Gula is reduced, being represented by a tiny triangular or transverse quadrangular plate. The gular sutures are almost fused with each other, and are forming a straight line.
4. Gula is absent. The gular sutures are completely fused, forming a straight line.
The 1st type is the most primitive state and is found in the Spheritidae, Synteliidae and Pachylomalus (Fig. 4D, G, 5D). The 2nd is present in the Dendrophilinae and Saprininae (Fig. 4J, 5H). The 3rd is found in the Trypeticinae, Onthophilinae and Abraeinae (Fig. 5F, L, 6B). The last type is the most derived state and is found in the Tribalinae and Histeridae (Fig. 6E, G, J). It is difficult to determine which of the 2nd and 3rd is more primitive.

Postocciput
Postocciput is a ring-shaped structure of the cranium and is usually reddish or yellowish brown in general. It is situated behind occiput and is separated from the latter by the postoccipital suture.
The postocciput is very important for phylogenetic study in the Histeroidea. In the most primitive state the postocciput is represented by a narrow and small plate and is situated along the lateral margin of the cavity. This state is found in the majority of the Histeroidea. In the Saprininae, the plate is elongate anteriorly along the gular suture, and is moderately narrow (Fig. 5H). In the Histerinae, the postocciput is enlarged on basal half of the ventral plate (Fig. 6G, J). This is the most derived state.

Antennae (Fig. 7 - 8)
Antennae are clavate and composed of the scape, pedicels and flagella. The base of the antenna is attached to the ventral side of the anterolateral margin of the forehead near the eye. Segments are numbered 11, except in Trypteticus, of which the antennae are 10-segmented (Fig. 7H). Scape is robust and geniculately connected with the pedicel. In the Hetaeriinae and the Chlamydopsinae, the scape is expanded and strongly angulate. Pedicel is usually short, small, and is usually narrower than the scape, but sometimes it is oval and broader in the Saprininae and Trypecitinae (Fig. 7H, 8A). Flagella are the 3rd to 11th segments of the antenna. In Trypeticus, they are the 3rd to 10th. In the Histeroidea, the apical three (usually 9th to 11th) are comprised in a "club" and the rest are called "funicle". The club is usually oval, condensed and more or less densely covered with hair; rarely it is cylindrical in the Heaterininae. In the Saprininae, the club is with "Reichardt's organ" (De Marzo and Vienna, 1982a, b) on the ventral surface, which is a particular sense apparatus (Fig. 7I, 8A). The articulation of the club is variable from a complete to an obscure one. The apicalmost segment of the funicle is usually expanded laterally to form a fringe.

Mouthparts (Fig. 9)
The mouthparts are composed of labrum, mandibles, maxillae, labium and their appendages. In the Histeroidea, the morphology of the mouthparts has not been sufficiently studied. My comparative studies are still imcomplate. Here, I describe their morphologies based only on two species, Notodoma fungorum and Hololepta amurensis.
Labrum (Fig. 9A, E) is an unpaired plate, situated before the clypeus and is separated from the latter by a distinct suture at the adult stage. It is transverse oblong, and the anterior margin is arcuate in Notodoma. The anterior margin is emarginate at middle and the lateral margins are also emarginate anteriorly in Hololepta. The lateral margin is furnished with hairs. In the Histerinae the surface of the labrum is not setiferous, while with a pair of setae in the Synteliinae, Sphaeritidae and other Histeridae.
Mandibles (Fig. 9D, H) are in a pair and well developed in the Histeroidea. The inner margin is usually provided with denticles. In Notodoma and Hololepta, there is no denticle.
Maxillae (Fig. 9B, F) consists of cardines, stipites, palpifers, laciniae, subgaleae, galeae and palpi all in pairs. The palpi is four-segmented, but may appear three-segmented because of the minute first segment. The galea is long in Hololepta.
Labium (Fig. 9C, G) is composed of the submentum, mentum, prementum, ligula and a pair of labial palpi. Mentum is nearly rectangurlar in Notodoma, while in Hololepta the anterior margin is strongly and acutely emarginate. Labial palpus consists of three segments.


Thorax

Cervix
The cervical elements are much reduced and composed of two pairs of lateral sclerites in the Histeroidea. They are situated ventrolaterally on the membrane between the cranium and the prothorax. The 1st (basal) sclerite is connected with the side of the anterior margin of the composite ventral plate of the prothorax by membrane. It is small, longitudinal oblong or triangular. The 2nd (apical) sclerite is much longer than the 1st.

Prothorax (Fig. 10 - 12)
The prothorax is separated into pronotum and prosternum in dorsal and ventral plates. The lateral side of pronoum is inflexed ventrally. The inflexed part is separated from the dorsal part of the pronotum by a distinct notohypomeral ridge or suture. Elements of the tergum of the prothorax are completely fused to form a well-sclerotized plate. Line of demarcation between the hypomeron and the composite ventral plate is tentatively called "tergopleural suture" (Naomi, 1988). The presternum is present in about half members of the Histeroidea, and is called "prosternal lobe". The furcasternum is a pair of semiglobulate plates, but they are difficult to observe in normal condition, forming the anterior wall or bottom of the fore coxal cavity which is concealed under the fore coxa.

General shape and modifications of pronotum
The pronotum is nearly rectangular, usually transverse and broadest near the base, its sides usually being convergent apically. The anterior foramen is small, but that of Syntelia is larger than the posterior foramen (Fig. 10C). The posterior margin of the pronotum is obtusely angulate at middle or gently arcuate. The surface is often punctate, striate, and/or costate.
The following modifications are observed on the pronotum in the Histeroidea. The pronotal sides are convergent basally in Syntelia (Fig. 10C). It is subquadrangular in shape in the Niponiinae (Fig. 45D), Trypteticinae (Fig. 12F) and Plegaderini (Fig. 11H). The pronotum is impressed by a marginal pronotal stria which runs along the notonypomeroal ridge and anterior margin in all Histeroidea. In Plegaderus (Fig. 113A), a transverse stria and longitudinal striae are present near the middle and on lateral third, respectively. There are other striae, an outer lateral pronotal stria and an inner lateral pronotal stria, in the subfamily Histerinae. The former is usually represented only on lateral portion, while the latter is complete on lateral and anterior portion except for an interruption behind the head. One to four pairs of costae are present on the pronotum in Onthophilus. A transverse stria is found along the posterior margin of the pronotum in Bacanius, Australomalus (Fig. 83A) and Acritus. A pair of longitudinal impressions is present on medioposterior area in Pachylomalus (Fig. 101A).
Punctation of the pronotum is very various in the Histeroidea. The pronotum is shining and smooth in some species of Hister, Margarinotus, Hypocaccus and so on. In most species of the Histeroidea, the punctation covers the whole surface of the pronotum, the punctures being fine to coarse, and dense to sparse, sometimes in various combinations of them. The punctation is often coarser and denser on the lateral region in Atholus, Merohister, some Hypocaccus, etc. A pair of large foveae are present at the lateromedian sides in Niponius impressicollis (Fig. 43D, E).

Hypomera
The hypomeron is strongly inflexed ventrally and extends from the anterior to the posterior margin of the pronotum. The posterior third of inner margin is usually strongly projects mesially. The apex of the projection is united with the posterior apex of the furcasternum in Syntelia (Fig. 10C). In all other members of the Histeroidea its apex is pointed and not united with the furcasternum. The notohypomeral ridge is distinct in all groups. The tergopleural suture is distinct in the majority of the Histeroidea, but is indistinct or completely lost in Trypeticus (Fig. 12F). In Onthophilus (Fig. 10G), the apical third of the hypomera is provided with an antennal cavity which is at least partly closed beneath by lateral region of composed ventral plate (prosternal alae).

Presternum
A transverse triangular, subquadrangular, or semicircular plate is present just before the composite ventral plate in the Synteliidae, Tribarinae, Onthophilinae, Histerinae, and Dendrohpilinae. It is called "prosternal lobe" and distinctly separated from the composite ventral plate by a suture between prosternal keel and lobe. In other histeroids the presternum is reduced. The prestenum is triangular and strongly acute in Syntelia (Fig. 10C). There is a semicircular or obtuse triangular plate in the Histerinae, Dendrophilinae, and Tribainae. In the Onthophilinae (Fig. 10G), it is not projected anteriorly, being transverse quadrangular. Presternum is usually impressed along the anterior margin by a marginal stria of prosternal lobe.

Composite ventral plate
The composite ventral plate is usually transverse and about 1/3 as long as wide. It is very variable in shape in the Histeridae and useful and important for phylogenetic study as shown below:
1. Plate flat, the median portion weakly convex. The medioposterior portion is projected as a narrow intercoxal process. This state may be the most primitive and is observed in Sphaerites (Fig. 10E).
2. Plate flat, the median portion weakly convex. The medioposterior portion projected, but not produced as a process, being concealed under the fore coxae. This state may be second primitive and is observed in Syntelia (Fig. 10C).
3. The median portion of the plate is elevated as a broad process, while the medioposterior portion is represented by the intercoxal process (prosternal keel or process). The anterior margin of the plate is not produced into the antennal cavity on each side. The medio-apical portion of the plate is connected with the presternum, and the anterior margin of the united plate (presternum + composed ventral plate) is nearly straight. This state is observed in the Niponiinae (Fig. 42C, 43F, 45E, 46F).
4. As in 3, but the anterior margin of the plate is produced into the antennal cavity on each lateral portion. The cavity is a transverse fovea, of which the posterior ridge is formed by an oblique running anterior margin of the presternum, and at least partly closed beneath by the presternum or the anterior margin of the composed ventral plate (prosternal alae). This state is observed in the Tribarinae and Histerinae (Fig. 11A, B, C).
5. Similar to the preceeding type, but the anterior cavities are very deep and connected with foveae of the hypomeron. In this state, the lateral margin of the plate is deeply interrupted transversely on each side by the cavity. This state is observed in Onthophilus (Fig. 10G).
6. The median portion of the plate is elevated as a broad process, and the basal portion is represented by the intercoxal process. The anterior margin of the plate is produced to form the antennal cavity on each lateral portion. The cavity is situated in front of the procoxa, and is represented by a longitudinal deep furrow, of which the inner ridge is strongly costate and formed by a longitudinally running anterior margin of the presternum. This state is observed in the Dendrophilinae (Fig. 11F).
7. Very similar to the 6th type. The median portion of the plate is strongly elevated as a process or a narrow keel. The presternum is absent. This state is observed in the Saprininae (Fig. 12A, C), Trypeticinae (Fig. 12F) and Abrareinae (Fig. 12H).
A pair of striae called lateral marginal prosternal striae are present on the basal half of the median area of the composite ventral plate along the median elevated process. The intercoxal process (prosternal process or keel) is often impressed by a pair of carinal striae.

Furcasternum
The furcasternum is longitudinal and narrow in Syntelia, the basal apex being united with the projections of hypomeron. In Sphaerites, it is transverse and broad. In Histeridae, the furcasternum forms the anterior wall of the fore coxal cavity and is composed of a pair of semiglobular plates.

Fore coxal cavities
The fore coxal cavity is transverse, broad and situated on the basal half of the prothorax. Its anterior wall is composed of the composite ventral plate and the anterolateral part of the furcasternum, its outer lateral wall belongs to the hypomeron, and its inner lateral wall is the posterolateral part of furcasternum. The posterior wall is usually partially closed by the hyopomeral projection, but is absent in Plegaderus (Fig. 12H). In Syntelia, the posterior wall is completely closed by the hypomeral projection, of which the inner apex is continuous with the intercoxal process (Fig. 10C).

Mesothorax
The mesothorax is composed of mesosternum, anepisterna, epimera, trochantins, prepectus and composite ventral plate. The mesosternum is much smaller than the pro- and metasternum. The anapleural suture is usually present between the anepisternum and the composite ventral plate. The trochantin is difficult to recognize as a distinct sclerite in the mesothorax. The presternum is not found in the Histeroidea.

Mesonotum
The mesonotum is usually called scutellum and its parts are almost concealed under the elytra. The exposed part of the mesothorax in dorsal view is usually triangular, but it is often invisible in minute histerids. The mesonotum is composed of anterior phragma, prescutum, scutum+scutellum and postnotum. The anterior phragma is well developed in the Histeroidea, and is represented by a pair of large deflexed lateral plates. The scutum and scutellum are amalgamated into a triangular plate in various degree in the Brachelytra. In Syntelia (Fig. 14A, B, C, 39B), the plate is longitudinal triangular. In the Histeridae, the scutum is well developed, lobulate, represented by a pair of lateral hanging structures (Fig. 14F, G).

Anepisterna and epimera (Fig. 13, 14)
The anepisterna and epimera are distinctly separated, and recognized in ventral view in the Synteliidae and Sphaeritidae. These plates are amalgamated in the Histeridae (Fig. 13). The anepisternum is usually not recognized in dorsal view, because it is situated under the mid coxae. The pleural suture is often absent in the Histeridae (e.g. Saprinus, Hololepta, and Plaesius).

Prepectus
The prepectus is absent in the Synteliidae and Sphaeritidae. It is present as arcuate collars inside the anterior margin of the anepisterna in the Histeridae.

Composite ventral plate (Fig. 15, 16)
The composite ventral plates of the Histeroidea representes two types.
Type 1. The plate is rather even, its anterior margin is thin. The sides of the plate are convergent apically. The median longitudinal ridge is developed on the posterior area. This type is present in the Synteliidae and Sphaeritidae (Fig. 15A, B, C), and more primitive than the type 2. In Sphaerites, the ridge is quadrangular and highly elevated.
Type 2. The median portion of plate is strongly elevated, its anterior margin is very thick. The portion is called "intercoxal disk of mesosternum". The anterolateral side is connected with the anepisternum on the anterior side. The posterior margin of the median portion is straight or angulate, not produced into a longitudinal ridge. This type is found in all members of the Histeridae (Fig. 15D, 16A, B).
Marginal mesosternal stria is present on the intercoxal disk of mesosternum. Another transverse stria is often found on median area in the Saprininae and some members of the tribe Exosternini. The stria of the Saprininae is usually strongly crenate and impressed along meso-metasternal suture, and called "meso-metasternal stria". The intercoxal disk has a transverse and subrectangular impression on the median area in the Paromalini.

Mesocoxal cavities (Fig. 15, 16)
The mesocoxal cavities are variable in accord with the types of the composite ventral plate. The cavities are narrowly separated by an intercoxal ridge, and closed anteriorly by the posterior margin of the plate in Type 1. On the other hand, the cavities are broadly separated by the "intercoxal disk of mesosternum", and open anteriorly in Type 2.

Metathorax (Fig. 15, 16, 17)
The metathorax is composed of metanotum, anepisterna, katepisterna, epimera and composite ventral plate. The trochantin is difficult to recognize as a distinct sclerite in the metathorax.

Metanotum
The metanotum is composed of acrotergite, anterior phragma, prescutum, scutum, scutellum, postnotum and posterior phragma. The antecostal suture is distinct in Onthophilus. The plate before the antecosta is acrotergite. The acrotergite projects anteriorly and deeply emarginate at the middle in Onthophilus, while it is elevated as costa at the middle in Saprinus.

Anepisterna and epimera (Fig. 17)
The anepisternum and epimeron are longitudinal quadrangular in shape. They are situated nearly parallel to each other in the Synteliidae and Sphaeritidae (Fig. 15A, C), while in the Histeridae the anepisternum is situated before the epimeron (Fig. 17). In the first two families they are long in the exposed area, while in the last they are rather short. The anepisternum is usually exposed completely. The anterior margin of the anepisternum is usually connected with the posterior margin of the epimeron of the mesothrax. It is not connected with the epimeron in Chaetabraeus, of which the anepisternum is very short and situated next to the hind coxa. The epimeron is more or less covered by the elytral epipleuron, its dorsal part is membranous and separated from the lateral part by a suture which may occur secondarily.

Composite ventral plate (Fig. 15, 16)
The composite ventral plate is usually transverse, and represents two types.
Type 1. The anterior margin of the intercoxal process between the mid coxae is narrower than the width of the midcoxal cavity. This type is found in the Synteliidae and Sphaeritidae (Fig. 15A, B).
Type 2. The anerior margin of the intercoxal process between the mid coxae is broader than the width of the midcoxal cavity. This type is found in the Histeridae (Fig. 15D, 16A, B). The median area is called "intercoxal disk of metasternum" and the lateral area "lateral disk of metasternum". The plate is impressed by the lateral metastenal and posterior mesocoxal striae.
The discriminal line (median longitudinal line or metasternal longitudinal suture) of the composite ventral plate is usually found in the Histeroidea; however, it is lost in Sphaerites (Fig. 15A), and is invisible in Chaetabraeus.
The following modifications are found on the plate. A deep fovea present behind each lateral anterior angle in Epiechinus (Fig. 53A). The male of some species of Onthophilus is furnished with a long hair. A small tubercle is found on the mid line in Acritus.

Position of hind coxae
The hind coxae are continuous to each other in the Synteliidae and Sphaeritidae (Fig. 15A, B), while they are broadly separated from each other in the Histeridae (Fig. 15D, 16A, B). These states are in accord with the condition of the composite ventral plate of the metathorax.

Metendosternite (Fig. 18)
The metendosternite is composed of an unpaired basal stalk and the anterior and furcal arms in pairs. The anterior arms are rudimentary in the Sphaeritidae and lost in the other groups of the Histeroidea. In Sphaerites, the metendosternite is Y-shaped, with the anterior arms short and the furcal arms extended obliquely and strengthened by broad and subtransparent laminae (Fig. 17A). In Syntelia, it is similar to that of Sphaerites, but the anterior arms are absent and the furcal arms are shortly developed mediolaterally (Fig. 17B). In the Histeridae, the metendosternite is U-shaped (Fig. 17C, D, E), the basal stalk is very short and transverse, and the furcal arms are broadly separated and extended obliquely.

Elytra (Fore wing)
The elytra are strongly sclerotized, usually black, sometimes maculate medianly with red or yellow, and having many striae, costae or tubercles, which are useful for species identification.
The elytra are usually oval in the majority of the Histeridae, but variable in length and width. They are elongate quadrangular elytra are present in the Synteliidae, Niponiinae and Trypeticinae. In the Sphaeritidae, the elytra are elongate oval.
The side margins of elytra are usually round and slightly convergent apically, and sometimes parallel. The anteromesial margin of the elytron fits to the lateral margin of mesonotum. The apical margin is usually truncate, exposing the last two abdominal tergites, except in Bacanius and related genera and also Sphaerites. The sutural margins of elytra always meet along the median line, forming a straight elytral suture. The cross section of the elytral suture is illustrated in Vienna (1980).
The elytra are variously striate and punctate on the upper surface. The length and situation of the striae and costae are very useful for species identification, and terminology for these characters has been proposed (Wenzel, 1944; Arnett, 1962; hhara, 1989). Basically the elytron has eleven striae, which are named from the outside to inside as follows: marginal epipleural, marginal elytral, external and internal subhumeral, oblique humeral, first to tenth dorsal, and sutural. The order of numbering in the Histeridae is opposite to that in most other Coleoptera, because the striae are frequently reduced from the inside in the Histeridae.
In the Sphaeritidae, the elytra are longitudinal rectangular, the outer lateral sides being moderately arcuate. The surface is completely striate and the striae are finely and densely punctate.
In the Synteliidae, the elytra are longitudinal rectangular. The surface is deeply striate, and the striae are crenate and often interrupted.
In the Histeridae, the elytral upper surface represents the following types.
1. The surface with striae.
1-a. All striae are nearly straight. Ends of 4th or 5th dorsal and sutural striae are not united with each other. Probably this type represents the most primitive state, being found in the Dendrophilinae, Tribalinae, and some groups of the Exosternini.
1-b. Fifth dorsal stria is absent. Ends of 4th and sutural striae are usually united with each other by an arc in basal area of elytron. Interstice between 4th dorsal and sutural striae very broad. This state is observed in Saprininae and some groups of Exosternini.
1-c. Interstice between 5th dorsal and the sutural striae not very broad. Basal ends of 5th dorsal and sutural striae are sometimes united with each other. This state is observed in the Histerinae.
2. The surface of elytra without stria or costa, only punctate variously. This type is often found in the small-sized histerids (e.g., Bacanius, Acritus, and some groups of the Paromalini).
3. The surface with strong costae. This type is found in the Onthophilinae.
4. The surface with many tubercles. A rare type, being limited to some species of Margarinotus.

Hind wing (Fig. 19 - 27)
Studies of the hind wing are very poor in the Histeroidea, a few studies having been made by Forbes, 1922 (Hister inaequalis), Kryzhanovskij and Reichardt, 1976 (Merohister jekeli, Hololepta amurensis, Epierus comptus, Plegaderus sp. Tryponaeus sp. Sphaerites glabratus, and Syntelia histeroides), hhara and Nakane, 1989 (Notodoma fungorum and Binhister chujoi), and hhara, 1991 (Hololepta amurensis).
The hind wing are functional in all examined species, but reduction rarely occurs (Crowson, 1974). The wing is usually transverse oblong and hyaline, and the veins are dark brown to yellow.
The veins of the hind wing in the Histeroidea are frequently and variously reduced. It is difficult to determine the homology of the veins with those of general insects. Veins recognizable in the Histeridae and their names and abbreviations adopted are given below. In terminology I follow Snoddgrass (1935) with some reference to Morimoto (1983) and Naomi (1989), who proposed terms on the basis of relationships between the axillary sclerites and the veins (Fig. 19).
HP: Humeral plate.
C: Costa runs on anterior margin of the wing and is unbranched; connected at the base with the antero-distally projection of the 1st axillary sclerite.
Sc: Subcosta arises from the 1st axillary sclerite and is unbranched.
R: Radius usually arises from the 2nd axillary sclerite in general insects. In the Histeridae, the base of radius is reduced and not attached to the 2nd axillary.
M: Media is connected with the median plate.
Cu: Cubitus is also connected with the median plate
Pcu: Post-cubitus is divided from near the base of the cubitus.
A: Anal vein is connected with the 3rd axillary sclerite.
There is no peculiarity in the other veins and axillary plates. No study has been made on folding patterns in repose.
Following notes are probably important for the phylogenetic study of the Histeroidea.
1. A definite M-Cu loop is present in the middle area of the wing. This state is primitive (Crowson, 1955), and is found in the Sphaeritidae, Synteliidae, all genera of Saprininae and the following genera of the Histeridae: Onthophilus, Dendrophilus, Anapleus, Epiechilus, Epierus (Fig. 20C, D, E, 21A, B, 22B, C, D, E, 23A, B).
2. Wings of all the observed members of the tribe Histerini are fairly uniform in venation and shape; the M-Cu loop is situated basally (Fig. 25A - E, 26A - C); the anal lobe is bilobulate (represented by two lobules).
3. Hind wings of the Saprininae and Dendrophilus are very similar, and share a characteristic and unique anal vein which is situated close to the Pcu and with a clear curve at its middle (Fig. 22B - E, 23A).
In insect wings, some general trends correlated with the absolute body size have been known. In histerids the following generalizations are recognized.
4. The veins are strongly reduced in the majority of small-sized histerids (e.g., Bacanius, Platylomalus, Chaetabraeus, Plegaderus, etc.), which are only about 1 mm long (Fig. 21E, 22A, 24B).
5. Large histerids (e.g., Merohister and Pactolinus, about 20 mm long) have slender and oblong wings. In small histerids the wings are oval, except in Bacanius, which has slender wings.
In Fig. 28, ratios of lengths between the several points on the hind wing are provided.


Legs
The legs (Fig. 29, 30) are composed of coxae, trochanters, femora, tibiae, tarsi and pretarsi (including claws) in the Histeridae.

Coxae
Fore coxae
The fore coxa is subcylindrical and elongate. A deep and narrow furrow is present on apical area in Syntelia (Fig. 29D), apparently in association with the closely set fore coxal cavities.

Mid coxae
The mid coxa is spherical and ovate in ventral view.

Hind coxae
The hind coxa is transverse triangular and subflat in the Synteliidae and Sphaeritidae, while in the Histeridae it is small and thick, and triangular in ventral view. The former type represents a more primitive state than the latter.

Trochanter
The fore trochanter is small and subconical, and subtriangular in ventral view. The mid and hind trochanters are similar to it. Usually with setae in the posterior margin, but often without them in the hind trochanter.

Femora
The fore, mid and hind femora are baculiform and thick, often with setae on the anteior (fore femur) or posterior margin (mid and hind femora). There are robust truncate setae on the fore femur of Saprinus. A longitudinal stria (profemoral stria) is present along the posterior margin of the fore femur on the ventral side in the tribe Histerini; in the tribe Platysomatini, the ventral surface is corrugated.

Tibiae
The shape of the tibia is subject to various modifications, which may vaguely be classified into the following 3 types.
1. The tibiae is slenderer than the femur and baculiform. Denticles on the margin of the tibiae are very small.
2. The tibiae is somewhat expanded apically, and wider than the femur. The denticles are large and well developed.
3. The tibiae is strongly dilated, thus more or less round in outline. The denticles are weakly developed.
The 1st type may be the most primitive and found in the Sphaeritidae and some groups of the Histeridae, that is, Onthophilinae, Niponinae, Trypeticinae and some genera of the Exosternini (Fig. 29A, G). The 2nd type is common in the Histeridae. The 3rd type is present in some species of the Hetaeriniae and Dendrophilus (Fig. 30D, E, F). The last type is often found in the members of myrmecophilous histerids.
The tarsal groove is present on the dorsal surface of the fore tibia in the Synteliidae and Histeridae. In the Hololeptini and the Platysomatini, the groove is very deep and shaped like an S. It is situated on the apical margin in the tribe Dendrophilinae and Anapleini. The ventral surface of the tibiae is densely covered with robust or long hairs or setae in some members of the Hypocaccus, Eopachylopus (Fig. 155) and Philothis. This condition is often associated with their habitats in sand on beach.

Tarsi
The tarsus is composed of five tarsomeres in general, but in Acritus, four-segmented in the hind leg, thus the tarsal formula is 5-5-4. The relative lengths of the tarsomeres are rather uniform, the 1st to 4th tarsomeres being short and the 5th long. In Saprinus, the mid tarsus shows a sexual dimorphism, being furnished with long setae in the male.

Abdomen
General structure
The abdomen is fundamentally composed of 10 segments in the Histeroidea. The elytra are truncate, and the 6th and 7th abdominal terga are exposed and strongly sclerotized in the Histeridae. On the other hand, in the Sphaeritidae and Synteliidae, the dorsum of the abdomen is almost covered by elytra and only the 7th abdominal tergite is exposed. The 1st and 2nd sterna are completely lost. The 3rd to 7th sterna are exposed and strongly sclerotized. The 8th, 9th and 10th segments are almost telescoped into the 7th.

Abdominal segments (Fig. 31)
Each of the 3rd to 7th terga is transversely rectangular in shape, and a transverse tergal suture runs along the basal margin.
The basal margin of the 3rd sternum is longitudinally carinated medially and the sides are shallowly concave for receiving hind coxa in the Synteliidae and Sphaeritidae. In the Histeridae, the carina is not present and the sides of the basal margin are roundly and deeply emarginate for receiving hind coxae, the emarginate portions being broadly separated from each other. The disk of the 3rd sternum is usually with one or two pairs of striae on each side of the median area. In taxonomic descriptions the disk of the 3rd is often called "1st abdominal sternum" and the striae "1st abdominal striae", because the true 1st and 2nd disks are absent and the 3rd appear to be the 1st. The 4th to 7th sterna are convex and a little broader than the terga, and each of them is transversely trapezoidal in shape in ventral view. The lateral sides of sterna are usually convergent posteriorly. The posterior margin of the 7th sternum is arcuately emarginate for receiving the posterior margin of the 7th tergite (pygidium).

Genital segments
The 8th, 9th and 10th abdominal segments are called genital segments. These segments are not exposed, and weakly sclerotized and pigmented.

Male
The 8th tergite is usually rectangular to subrectangular in shape in dorsal view and often divided into two plates by a longitudinal mid line. A pair of short or long processes are usually present at the anterolateral corners of the tergite. The 8th sternum is more variously modified than the tergum. For example, it is divided into two plates by a longitudinal mid line, and each plate is elongated posteriorly and rounded apically in the Sphaeritidae (Fig. 38), Carcinops (Fig. 92), some members of the tribe Platysomatini (Fig. 58, 59, 60) and Histerini. The ventral surface is strongly sclerotized and densely covered with hairs in Saprinus (Fig. 133, 135, 136) and most Saprininae. The 8th sternum is oblong in ventral view, and the caudal disk with a round disk or some small oblong disks in the tribe Paromarini (Fig. 95, 98, 101, 104, 106, 109, 111, 113, 116).
The 9th tergum is often divided longitudinally. This state is found in Gnathoncus, Carcinops, Anapleus, Epiechinus and Platysoma (Fig. 126, 129, 130, 92, 83, 90, 54, 58, 59, 60, 62, 65). The anterolateral corner is projected. In Trypeticus, the projection is very long (Fig. 160). The posterior margin is connected with the 10th tergum and nearly straight, but often emarginate arcuately or acutely for fitting the anterior margin of the 10th. The 10th tergum is present in the Histeroidea, but, in my examination, not in Trypeticus.
The 9th sternum is composed of an unpaired plate, and is called "spicule". It is a thin plate, longer than broad, and may be longitudinal and oblong in its primitive state as represented by the Sphaeritidae and Synteliidae, Plegaderus, and Niponiinae (Fig. 38E, 40E, 114F, 44E, 47E). The anterior apex is strongly sclerotized in a V in Epierus (Fig. 51E) and Niponius. In most Histeridae, the 9th sternum is modified variously. The posterior apex is expanded and produced into a triangular disk in Anapleus (Fig. 83G, 90G), Platylomalus (Fig. 106F) and Dendrophilus (Fig. 79F). The lateral angles of the posterior apex are elongated posterolaterally, thus the sternum is T-shaped in the tribe Histerini and Platysomatini. In the Saprininae too, it is often shaped like a T, the anterior apex often expanded (Fig. 128I, J, 129I, 130I, 133I, J, 134I, 135I, 135H, 143G, 144G, 145G, 146G, 150G,152G, 156H). The projections of the posterior apex are united together on dorsal side to form a ring, into which the basal piece is fitted.

Female
The female genital segments have not been studied enough to make a revision. They are out of scope in this study.

Male genitalia
Male genitalia consist of 3 parts: basal piece, a pairs of parameres, and median lobe. Sharp (1800) firstly made comparative study of coleoptera. In Histeridae, the male genitalia are considered to be the most useful character for the classification at species level, because they are usually different in shape from species to species and morphologically stable within a species.

Basal piece
The basal piece is usually a ring surrounding the basal part of the median lobe, and the posterior margin is connected with basal margin of paramere. In Syntelia, Sphaerites and Anapleus, it is an incomplete ring, being not closed on dorsal or lateral side (Fig. 38A, B, 40A, B, 83A, B, 85A, B, 90A, B). This condition may be the most primitive in the Histeroidea. It is absent in some unrelated taxa: Acritus, Bacanius and part of Hetaeriinae. The basal piece is variable in length in the Histeroidea. It is usually half as long as the paramere, but, in Niponius (Fig. 44A, 47A), members of the tribe Paromalini (Fig. 92A, 95A, 98A, 101A, 104A, 106A, 109A, 111A, 113A, 116A, 118A), Epierus (Fig. 51A), Trypeticus (Fig. 151A) and some members of the tribe Platysomatini (Fig. 65A), it is as long as or much longer than the paramere.

Parameres
The parameres are principally represented by a pair of slender and lobate structures in the Coleoptera. In the Histeroidea, the parameres are usually fused basally on the ventral side to form a tube structure. Their dorsal side is usually provided with a slit. The basal margin of the parameres is connected with the posterior margin of the basal piece. The most primitive state of the parameres is found in Sphaerites and Syntelia (Fig. 38A, 40A): the parameres are not fused dorsally and also separated in their apical halves ventrally; the median lobe is extruded from the ventral side. In the majority of the Histeridae, the parameres are partly fused in the basal half and moderately separated in the apical half; the median lobe is extruded from the dorsal side. The median lobe is, however, extruded from the ventral side in Hololepta and members of the tribe Exosternini (hhara and Nakane, 1989). The parameres are various in shape and length: they are usually slender and convergent apically; in Platysoma (Fig. 58A, 59A, 60A), the apical part is lobate, elongate apically and convergent apically; especially in Apobletes (Fig. 65A) the lateral side is strongly emarginate; in members of the Paromalini, it is short.

Median lobe
The median lobe is usually slender and is surrounded by the parameres and the basal piece. It is various in shape, varying from a flat and weakly sclerotized lobe to a strongly sclerotized one with armature. Their shapes may tentatively be classified into the following types.
1. A weakly sclerotized and flat lobe is found in Syntelia, Sphaerites, Niponius, Trypeticus, Epierus and most members of the Saprininae. This condition is the most primitive state.
2. The lobe is moderately sclerotized and flat, with a slender process on the basal angle. This condition is found in the following taxa: Onthophilus, Epiechinus, most members of the Dendrophilinae, Exosternini, Platysomatini and some members of the Histerini.
3. The lobe is strongly sclerotized and provided with armatures, a condition probably derived from Type 2. This is found in some members of the tribe Histerini: Hister, Atholus, Zabromorphus, Merohister, Margarinotus and so on (hhara, 1989, 1992a, 1992b, 1992c, 1993).

Female genitalia
Spermatheca
The spermatheca is the sperm receptacle of the female. De Marzo and Vienna (1982) first made a careful morphological study of the organ in the Histeridae, and recognized two fundamental patterns: I) spermatheca including 1 receptacle, a more or less long duct and a distinct gland; II) spermatheca including 4 to 9 sessile receptacles, without distinct spermathecal gland. The second pattern is found in the tribe Histerini, while the first widely in other groups. In this study, I have also recognized both the patterns, which may be sudivided into the following 6 types:
I). Spermatheca shape like a globe, strongly sclerotized.
1. Without projection or invagination.
2. With a developed invagination.
3. With a developed external projection.
4. With developed internal projections.
II). Spermatheca consisting of several small sacs each with slender tube basally.
5. The sacs are attached to the wall of the vagina, the tubes being not coiled.
6. The sacs are attached to the wall of the bursa copulatrix, usually small and globe-shaped; the basal tubular part is often coiled.
The 1st type is probably the most primitive state, being common in various groups.
The 2nd type is found in the Saprininae (Fig. 32H, I) and Plegaderus (Fig. 32G). The spermatheca of the Saprininae is strongly vaginate around the area connected with the duct of spermatheca and also around the area connected with the spermathecal gland. Plegaderus has a pear-shaped spermatheca feebly invaginate apically.
The 3rd type is found in Niponius (Fig. 32D) and Platylomalus (Fig. 32K). Probably this type has independently appeared in these genera.
The 4th type is found in the Dendrophilinae, with one internal projection in Carcinops and two in Platylomalus (Fig. 32J) and Eulomalus (Fig. 32L).
The 5th and 6th types are present only in the tribe Histerini, the former is observed in Hister, Atholus, Merohister, Zabromorphus, and Pactolinus (Fig. 33E, F, G, H, I), while the latter is found only in Margarinotus (Fig. 33J, K, L).
The spermathecal gland is usually a small saccule, weakly sclerotized and with a long straight or coiled spermathecal duct, but is sometimes absent.


5. Pylogeny

Phylogenetic Relationships of the Families
The superfamily Histeroidea was adumbrated by Sharp and Muir (1912), with the remark that "four families Histeridae, Synteliidae, Sphaeritidae and Niponiidae are so closely related by the aedeagus, that they might form one family". The group was adopted by a majority of the histerid systematists (Reichardt, 1941; Wenzel, 1944; Crowson, 1955; 1974; Kryzhanovskij and Reichardt, 1976; Mazur, 1984; Hisamatsu, 1985).
The superfamily Histeroidea is included in the Staphyliniform lineage, which is often divided into three superfamilies: the aquatic Hydrophiloidea, and the terrestrial Histeroidea and Staphylinoidea. The Histeroidea have been considered the most closely related to the superfamily Hydophiloidea by some authors (Bvving and Craighead, 1931; Reichardt, 1941; Morimoto, 1986). On the contrary Crowson (1955; 1974) insisted that the Histeroidea are more closely allied to Stahpylinoidea than to the Hydrophyloidea. But recent works tend to refute Crowson's hypothesis. Discoveries of larvae of the primitive histeroid families Sphaeritidae and Synteliidae (Nikitsky, 1975; Mamayev, 1976) have shown that these larvae share numerous specializations common to Hiseridae and Hydrophilidae.
The Histeroidea and Hydrohiloidea are united by the following derived larval character states: 1) labrum fused to head capsule and without tormae , 2) mandible falcate with reduced molar lobe, 3) basal segment (palpifer) of maxillary palp complete and bearing articulated appendage, 4) tentoruium with posterior arm attached directly to head, with a short bridge attached well above venter of head, 5) spiracle biforous with an elateroid molting process, 6) abdomen largely membranous with scattered small sclerites, and 7) final inster without ecdysial line on head. Known pupae lack functional spiracles on the first abdominal segment. Adult antennae are short, usually with a sharply differentiated club composed of three densely pubescent segments; abdominal segment seven is invaginated at least ventrally; and the seventh or seventh and eighth spiracles are atrophied. Nearly all larvae are carnivorous with mouthparts adapted for extraoral digestion. Recently Lawrence and Newton (1982) have proposed to include the Histeroidea in the Hydorphiloidea (excluding Hydraenidae). According to them there are no sufficient differences between these taxon to divided into the superfamilies. In this study, however, I adopt the superfamily Histeroidea as a good taxon because of sufficient differences in life habit between these groups.
The superfamily Histeroidea consists of three families, Sphaeritidae, Synteliidae and Histeridae, and is characterized in adult by the compact antennal club, very prominent, acute mandibles, truncate elytra covering six abdominal segments at most, ovipositor with scoop-like gonocoxites bearing mesal styli, and carnivorous feeding habit. There is a general agreement on the monophyly of this group and on the derived position of the highly compact Histeridae.
Some authors treated the Niponiinae as distinct family (Gardner, 1926; Nakane, 1963; Hisamatsu, 1985). Since Reichardt (1941), however, the majority of the histerid systematists has adopted a system consisting of the three families. Studies of the phylogenetic relationships among these families have been written fragmentally by some authors. Crowson (1955) stated that the families Sphaeritidae and Synteliidae are more primitive than the Histeridae.

Phylogenetic Relationships of the Subfamilies
Many authors have proposed infrafamilial classification of the Histeridae (see Historical review), but no cladograms have yet been published and no studies have been presented with specific intent to distinguish between apotypic and plesiotypic character states.
Since Wenzel (1944) proposed a new system, most authors (Hatch, 1962; Mazur, 1981; Vienna, 1981; Yllamos, 1985 and Hisamatsu, 1985) have followed him.
Mazur (1984) refined the Wenzel's system in his world catalogue. This revised system, here called "Wenzel-Mazur System", adopts traditional and important character states for constructing phylogeny without evaluating their atypic-plesiotypic relationships.
The subfamilies are so diverse and distinct, that it has been difficult to find other characters which can be used to construct morphoclines through them. In this section, the character states used in the Wenzel-Mazur System are evaluated first. Then I propose a new cladogram for the subfamilies of the Histeridae.

Wenzel and Mazur System
Wenzel (1944, 1962) and Mazur (1984) did not distinguish between primitive and derived states of their characters. Hence details of the phylogenetic relationship within the family Histeridae are not clear in their system. I have tried to determine polarities (primitive-derived relationships) for their character states, and then applied them to the key branching diagram of the Wenzel-Mazur System.
The character states are given in Table 2. They have been picked up from key in Wenzel (1944 in Arnett, 1962) and also from a diagnosis given by Olexa (1982). Out-group comparisons have been made to determine polarities between paired states, which are given 0 (primitive) and 1 (derived), sometimes also 2 (more derived), in the table. The out-groups adopted Synteliidae (Syntelia histeroides) and Sphaeritidae (Sphaerites politus). However, character 2, 5 and 14 could not be determined as to the polarities of their states, though these are arranged with code number. Character 5 has five states (Code 0 - 4), which may not be in a linear relationship. Character 1, 6, 10 and 13 includes negative or reductive states, which are interpreted as derived.
The diagram thus constructed is shown in Fig. 35. It includes five cases of homoplasty, three of reversal, and two of parallelism.
However, I do not think that two of the indicated case of reversal are real: Character 2AnCa and 9HeSt in the Niponiinae. The other indicated case is 9HeSt in the Tryponaeinae. Parallelism is indicated for 2 and 14AnCa and 3 and 12AnSc. The case of AnCa may not represent real parallelism. The derived state of AnSc is apparently associated with the myrmecophilous habit in the Chlamydopsinae and Hetaerinae.

Proposed cladogram
In Section 4 (Morphology and structure and terms), I examined and compared many external characters, but I could not fine any additional states useful in phylogeny reconstruction at the subfamily level except the mesosternal width.

Table 2. Character states used in Wenzel-Mazur System and their presumed polarities.
____________________________________________________________________
No. Mnemonic Code State
____________________________________________________________________
1 PsLo 0 Prosternal lobe present (Histeromorphae).
1 Prosternal lobe absent (Saprinomorphae).
2 AnCa 0 Antennal cavities absent.
1 Antennal cavities closed beneath by the prosternal alae.
2 Antennal cavities [if present] open beneath.
3 AnSc 0 Antennal scape normal, the club usually a little, but never three times, large than broad.
1 Antennal scape strongly angulate and usually greatly expanded, the club greatly elongated, at last three times as long as broad.
4 Form 0 Form round, oval or oblong-oval.
1 Form cylindrical, sometimes stoutly so.
5 AnCa 0 Antennal cavities absent.
1 Antennal cavities situated in anterior prothoracic angles, at least partly closed beneath by prosternal alae.
2 Antennal cavities situated in anterior prothoracic angles, wholly closed beneath by prosternal alae.
3 Antennal cavities situated in front of procoxae, next to and nearly always encroaching upon prosternal keel.
4 Antennal cavities situated in anterior prothoracic angles, or in front of procoxae.
6 ElSt 0 Dorsal elytral striae rarely absent, if so, then at least sutural stria present.
1 Dorsal elytral stria never present through rather vague impression may be evident.
7 HeHn 0 Head not produced into two horns, though it may be produced as a long pointed rostrum in the female.
1 Head produced anterior into two horns of variable length.
8 Mand 0 Mandibles moving in the same plane as the long axis of the head.
1 Mandibles deflexed, moving in a plane at right angles to the long axis of the head.
9 HeSt 0 Head horizontal in repose.
1 Head vertical in repose.
10 AnSe 0 Antennal consisting of eight segments and a club.
1 Antennal consisting of seven segments and a club.
11 PrAl 0 Anterior margin of prosternal alae without reception of anternnal funicle.
_____________________________________________________________________

Table 2. Continued.
_____________________________________________________________________
1 Anterior margins of prosternal alae with rather deep longitudinal incisions for the reception of the anternnal funicle in repose.
2 Anterior margin of prosternal alae at most with only very slight notch. Antennal cavities not defined.
12 AnSc 0 Antennal scape normal, neither expanded nor strongly angulate.
1 Antennal scape expanded and strongly angulate.
13 LaSe 0 Labrum with setigerous punctures
1 Labrum without setigerous punctures.
14 AnCa 0 Antennal cavities absent.
1 Antennal cavities situated in anterior prothoracic angles, at least partly closed beneath by prosternal alae.
2 Antennal cavities completely open, usually situated just anterior to procoxae at middle of hypomera.
15 PtDe 0 Protibiae multidenticulate.
1 Protibiae oligodenticulate.
16 DoCo 0 Dorsal surface normal, without costae.
1 Dorsal surface with distinct costae.
_____________________________________________________________________

In Table 3, the mesosternal width and the characters mentioned in Table 2 are arranged against subfamilies, and their states are indicated by code numbers, which are the same as those given in Table 2 (for codes for the mesosternal width see under Table 3). Character 2, 5 and 14 in Table 2 are represented by Character 5, and Character 3 and 12 by Character 3, in Table 3. Based on this data matrix, a cladogram is constructed (Fig. 36). The new cladogram is characterized by the erection of three major groups instead of two (the Saprinomorphae and the Histeromorphae) in the Wenzel-Mazur System. It is also remarkable in the following interpretations:
1) The Niponiinae are the most primitive subfamily (Character HeSt: Code 0). The two supposed reversal states in Fig. 35 are cleared in the new cladogram.
2) Wenzel (1944) recognized the two major groups on the basis of Character PsLo. However, the prosternal lobe is present (PsLo: 0) in the Niponiinae, which were erroneously included in the Saprinomorphae by Wenzel. Moreover, Character AnCa is more weighed than PsLo in the new cladogram; the Dendrophilinae, therefore, are not a histeromorph but a saprinomorph subfamily. In the new cladogram it is postulated that the prosternal lobe was lost in the common ancestral taxon of the Abraeinae, Saprininae, Trypanaeinae and Trypeticinae, and the supposed parallel evolution of 14AnCa Code 3 and 2AnCa 3 is not accepted. After all, the Saprinomorphae and the Histeromorphae, as composed in the Wenzel-Mazur System, are rejected.
3) The Chlamydopsinae are highly adapted to myrmecophilous habit. Their prosternal structures are diversified and sometimes reduced, and it is difficult to conform them to a transformation series. However, their antennal cavities are closed beneath by the prosternal alae, and there is no distinct prosternal keel. They should be placed next to the Onthophilinae and the Tribalinae.
4) The parallel evolution of Character Form (Code 1) may be associated with living in the gallery of scolid beetles, the concerned histerid preying on them.
In the Saprininae and Dendrophilinae the anal vein of the hind wing is distinctly curved at basal one-third. This may be a common derived state in these subfamilies, supporting the new cladogram. (In the Abraeinae, Trypanaeinae and Trypeticinae the veins of the hind wing are strongly reduced in association with their small body size.) These two subfamilies also agree in having a large and triangular gular plate.

Table. 3. Data matrix.
_____________________________________________________________________
No. Anc Nip Chl Ont Tri His Het Den Abr Sap Tpa Tpe
0 MsWi 0 1 1 1 1 1 1 1 1 1 1 1
9 HeSt 0 0 1 1 1 1 1 1 1 1 0* 1
7 HeHn 0 1 0 0 0 0 0 0 0 0 0 0
8 Mand 0 1 0 0 0 0 0 0 0 0 0 0
5 AnCa 0 0 2 2 1 1 1 3 4 3 3 3
1 PsLo** 0 0 1 0 0 0 0 0 1 1 1 1
11 PrAl 0 0 0 0 0 0 0 1 1 1 2 1
3 AnSc** 0 0 1 0 0 0 1 0 0 0 0 0
4 Form** 0/1 1 0 0 0 0 0 0 0 0 1 1
6 ElSt 0 0 0 0 0 0 0 0 1 0 0 0
10 AnSe 0 0 0 0 0 0 0 0 0 0 0 1
13 LaSe 0 0 0 0 0 1 0 0 0 0 0 0
16 DoCo 0 0 0 1 0 0 0 0 0 0 0 0
_____________________________________________________________________
Anc: Synteliidae and Sphaeritidae; Nip: Niponiinae; Chl: Chlamydopsinae; Ont: Onthophilinae; Tri: Tribalinae; His: Histerinae; Het: Hetaeriinae; Den: Dendrophilinae; Abr: Abraeinae; Sap: Saprininae; Tpa: Trypanaeinae; Tpe: Trypeticinae.
* Reversal; ** Parallelism.
No. 0, MsWi, code 0: mesosternum between mesocoxae narrower than mesocoxal cavity; code 1: mesosternum between mesocoxae broader than the mesocoxal cavity.


6. Systematics

Key to the families of the Histeroidea

1(4) Hind coxae not separated from each other. Anterior margin of intercoxal disk of mesosternum between mid coxae is narrower than the width of the mid coxal cavity. Mid coxa closed anteriorly by mesothorax. Only last abdominal tergite exposed.
2(3) Body oblong-oval. Prosternal process short. Striae of elytra finely punctate, not deeply impressed. ........................................... Family Sphaeritidae
3(2) Body oblong. Prosternal keel almost absent, but a tubercle present posteriorly between fore coxae. Striae of elytra deeply impressed. ...... Family Synteliidae
4(1) Hind coxae broadly separated from each other. Anterior margin of intercoxal disk of mesostrernum between mid coxae is broader than the width of the mid coxal cavity. Mid coxae broadly separated by mesosternal intercoxal disk and opened anteriorly. Anepisterna of mesothorax concealed under mid coxa, not visible in ventral view. Last 2 abdominal tergites exposed. ... Family Histeridae

6. 1. Family Sphaeritidae (Heer, 1841)

Thomson, 1862: 23 [type genus: Sphaerites Duftschmid, 1805]; Ganglauber, 1899: 412; Ganglauber, 1903: 305; Jakobson, 1915: 658, 869; Schenkling, 1913: 1-2; Reichardt, 1941: 1-10; Crowson, 1955: 13, 24; 1974: 133; Arnett, 1962: 385; Adachi and Ohno, 1962: 148; Witzgall, 1971: 189.
English name: the false clown beetles.

Genus Sphaerites Duftschmid, 1805

Diagnosis by Arnett (1962). "These beetles superficially resemble a hister, but somewhat loosely articulated, not compact, legs not tightly co-apted to body; they lack the strongly geniculate antennae and the external teeth on the tibiae. The elytra, although still truncate, cover all but the last abdominal tergite, and more loosely cover the abdomen.
Shape oval, convex; size about 6 mm. in length; color dark greenish-metallic; vestiture absent except on the tarsi, antennae, and around the mouthparts.
Head much narrower than the pronotum, deflexed, surface punctate. Antennae eleven-segmented, strongly capitate, the club three-segmented, not strongly geniculate; inserted under a slight frontal ridge close to the margin of the eyes and the base of the mandibles. Labrum very short, transverse; mandibles very large, curved apices very acute, almost hook-like with stout tooth; maxillary palpus four-segmented, the first segment very small, the remaining segments stout, the apical segment the largest, cylindrical, somewhat depressed; labial palpus three-segmented, the segments stout, cylindrical. Eyes lateral, moderate, oval.
Prothorax freely movable; pronotum much broader than the head; anterior border emarginate for the insertion of the head; sides evenly acute, broader posteriorly; posterior border sinuate, sides and anterior border margined; surface punctate more so laterally; pleural region broad, explanate; procoxal cavities narrowly open. Mesosternum narrow; metasternum broad. Legs with front trochantins exposed; anterior coxae transverse; contiguous, prominent; middle coxae small, globular; hind coxae transverse; trochanters small; femora swollen; anterior tibia with external spines, but without teeth. Tarsal formula 5-5-5, the segments slender, fringed with numerous setae; claws prominent with a bisetose empodium between. Scutellum moderate, triangular. Elytra convex, truncate, exposing only the pygidium; striae represented by a row of punctations; epipleural fold moderate. Wing venation with m-cu loop relatively large and distinct, radial cell distinct, stigma reduced.
Abdomen with five visible sternites, surface slightly punctate. Penis thin; parameres nearly completely fused, large; pars basalis small, asymmetrical, forming a band. Female genitalia undescribed."
Larvae: Nikitsky (1976) described the larva of Sphaerites glabratus.

Sphaerites politus Mannerheim, 1846
(Fig. 37, 48, 41)

Spaerites politus Mannerheim, 1846: 514; Kryzhanovskij and Reichardt, 1976: 22; Adachi and Ohno, 1962: 149; Nakane, 1963: 67, pl. 34, no. 1; Matsumoto, 1981: 6 [Mt. Hidaka]; Yasuda, 1982: 76 [Mt. Daisetsu]; Hisamatsu, 1985: 219, pl. 40, no. 1; Yasuda, 1988: 25.
Japanese name: Emma-mushi-damashi.

Description. Body length, PPL, 4.56 - 5.63 mm (5.14 0.12, n=9), PEL, 4.38 - 5.31 mm (4.99 0.10, n=9). Width, 2.94 - 3.56 mm (3.28 0.07, n=9). Body oblong (Fig. 37A). Cuticle shining and bronze; tarsi and antennae dark rufopiceous. Biometric data are given in Table 4.
Anterior margin of head narrowly marginate laterally between basal corner of epistoma and eye just behind antennal cavity. Surface of head coarsely and densely punctate, the punctures becoming finer and sparser on central area and epistoma.
Anterior margin of pronotum (Fig. 37A) evenly emarginate; lateral sides weakly arcuate on apical half, and nearly straight on basal half; posterior margin strongly arcuate outwards in median third; marginal stria complete laterally, its apical end curved behind apical corner, extended inwards, and attaining to near the lateral third of pronotum, the stria somewhat distantly present from the pronotal margin; surface sparsely and coarsely punctate laterally, and with other fine punctures intermingled, the fine punctures progressively finer inwardly; on median area of the surface clothed with microscopic punctures.
Epipleura narrow. Marginal epipleural stria complete and carinate. Marginal elytral stria strongly carinate and complete. Area between the margin of epipleura and the marginal elytral stria shining and sparsely clothed with several microscopic punctures. Elytra (Fig. 37A) with 10 rows which are nearly complete and consist of coarse and round serial punctures, which are separated by their own diameter to twice the diameter but irregularly scattered on apical sixth; intervals among the rows sparsely clothed with microscopic punctures.
Pygidium (Fig. 37B) with marginal stria along lateroposterior margin, the basal end not attaining to the basal corner, curved before the corner and shortly extended inwards; surface of disk densely covered with coarse punctures which are a little coarser than in elytral rows; other fine punctures intermingled among them, becoming sparser apically.
Antennal grooves present on apical area of underside of head, the grooves running obliquely from apex of eyes to middle of head.
Anterior margin of prosternum evenly emarginate, and with long hairs; median area of disk densely covered with coarse setiferous punctures, the setae being somewhat long.
Mesosternum small and quadrangular, its disk densely covered with coarse punctures. Meso-metasternal suture distinctly present. Anterior margin of metasternum behind mesocoxae strongly carinate and marginate; intercoxal disk sparsely covered with fine punctures, the punctures becoming coarser laterally; lateral disk sparsely covered with large, round and shallow punctures. Metepisternum densely punctate, the punctures as large as those of lateral disk of metasternum; on area along the posterior margin transverse stria present, but broadly interrupted on median third, the median end of the stria densely with coarse punctures.
Protibia (Fig. 37D, C) slender, not expanded, usually with 7 denticles on outer margin.
Male genitalia as shown in Fig. 38A - F.
Female genitalia: spermatheca as shown in Fig. 38G.
Specimens examined. [Hokkaid]] 13, 1 ex., Mt. Kurodake (Alt. 1900m), Mt. Daisetsu, 18/vii/1981, N. Yasuda leg. (NA); 7 exs., Ditto, 23/vii/1981, N. Yasuda leg.; 4 exs., Ditto, 28/vii/1982, N. Yasuda leg.; 2 exs., Akadake, 16-23/vii/1975, M. Suwa, M. Furukawa, M. Kiuchi and T. Sunose leg. <Rishiri Is.> 1 ex., Rishiri Is., 24/vii/1963, Y. Shibata leg. (NSMT); 1 ex., Oniwaki trail, Mt. Rishiri, 7/viii/1992, M. Sat leg.
Distribution (Fig. 41). Japan (Hokkaid, Rishiri Is.); North America.
Remarks. Sphaerites politus can rather easily be recognized by the characters given in the key to the families.
Little is known about the habitat of this species. This species occurs at high altitudes. Yasuda (1982, 1988) collected it by using pit-fall traps in the forest of Pinus pumila at alt. 1800 m to 1980 m on Mt. Kurodake, Daisetsu, Hokkaid.

6. 2. Family Synteliidae Lewis, 1882

Synteliidae Lewis, 1882: 137-138 [type genus: Syntelia Westwood, 1864]; Sharp, 1891: 438; 1899: 229; Ganglbauer, 1899: 415; Ganglbauer, 1903: 282, 305; Kolbe, 1901: 134-135; 1908: 121, 122; Jakobson, 1915: 658, 869; Handliksch, 1923: 527, 596; Hetschko, 1926: 13; Yuasa, 1930: 253; Crowson, 1955: 24, 25.

Table 4. Biometric data of Sphaerites politus Mannerheim.
-----------------------------------------------------------------------
APW 1.31-1.63 (1.470.03) 9
PPW 2.38-2.81 (2.650.05) 9
PL 1.25-1.56 (1.480.04) 9
EL 2.75-3.75 (3.410.09) 9
EW 2.94-3.56 (3.280.07) 9
PyW 1.63-2.00 (1.850.04) 9
PyL 0.88-1.25 (1.050.05) 9
PTL 1.13-1.50 (1.330.04) 9
MSTL 1.25-1.56 (1.350.03) 9
MTTL 1.50-1.94 (1.720.05) 9
-------------------------------------------------------------------------

Genus Syntelia Westwood, 1864

Syntelia Westwood, 1864: 11 [type species: Syntelia indiea Westwood, 1864]; Reitter, 1876: 18; Sharp, 1891: 439, t. 14, fig. 1, 1b; Kolbe, 1901: 108; Jakobson, 1915: 869; Hetschko, 1926: 13; Yuasa, 1930: 253.

Syntelia histeroides Lewis, 1882
(Fig. 39, 40, 41)

Syntelia histeroides Lewis, 1882: 137 [Japan: Yashiu, Southern Hokkaid, Kumakawa in Higo, Sado]; Waterhouse, 1882-1890: t. 133, fig. 8; Redtenbacher, 1886: 216, t. 19, fig. 122; Sharp and Muir, 1912: 511, t. 55, fig. 77, 438; Sharp, 1899: 229; Ganglbauer, 1899: 415; Jakobson, 1915: 869; Hetschko, 1926: 13; Mastumura, 1931: 16; Yuasa, 1930: 257; Kryzhanovskij and Reichardt, 1976: 414; Nakane, 1963: 67, pl. 34, no. 2; Hisamatsu, 1985: 219, pl. 40, no. 2.
Japanese name: Emma-mushi-modoki.

Original description. "Black, shining: head with a few large scattered punctures; thorax, disc smooth with some deep punctures at the sides, lateral edge and base emarginate, elytra smooth with deep punctate striae, four dorsal more or less broken, one outer and one sutural complete, the last continuous, running round both the apex and base of elytra and joining the outer elytral margin. The pygidium is evenly and coarsely punctured, convex in the middle, with lateral depressions deepening and ending before the base. Beneath, the segments of the abdomen are sparsely punctured in the middle, more thickly at the sides, mesosternm behind the middle coxae smooth in the medial region, and in front of coxae thickly and somewhat strigosely punctate.
This species differs much from S. indica, the chief points of variance being colour, punctuation of pygidium, the more convex and quadrate thorax, and the deep irregular elytral striae. The spines on the tibiae correspond in both species. The elongation of the thorax in Westwood's figure of indica is somewhat exaggerated. Length 6. 1/2 - 7. 1/2 lines. "

Description. Body length (length between apex of head and apex of pygidium), 11.00 - 16.00 mm (13.60 0.31, n=20). Width, 3.25 - 4.75 mm (4.09 0.08, n=20). Biometric data are given in Table 5. Body (Fig. 39A) cylindrical, somewhat stout. Cuticle shining, black; tarsi and antennae dark rufopiceous.
Head (Fig. 39A) coarsely punctate on a transverse narrow band between eyes and near around eyes, the punctures irregularly scattered and sometimes present posteriorly; surface among the coarse punctures sparsely and evenly clothed with microscopic punctures.
Anterior margin of pronotum (Fig. 39A) densely with short hairs and its median portion slightly arcuate outwards. Sides of pronotum clearly convergent posteriorly, carinate and marginate, the marginal stria curved behind anterior angles and shortly extended inwards, and the posterior ends of the marginal stria extending inwards along posterior margin and united with each other. Surface of pronotum coarsely and densely punctate on a narrow area along the marginal stria, the punctures a little coarser than those of head and several ones scatteringly present on broad lateral area; remain of surface of disk smooth, evenly clothed with microscopic punctures, the punctures being separated by five to ten times their diameter.
Scutellum (Fig. 39B) longitudinally oblong.
Epipleura of elytra narrow. Marginal epipleural stria feebly carinate and complete. Marginal elytral stria complete, strongly carinate and crenate, and inwardly sinuate on basal fourth; its basal and apical ends extended inwardly along basal and apical margins of elytra, and the ends united with basal and apical end of sutural stria respectively; area between marginal epipleural and elytral striae with three rows of setiferous punctures on dorsal fourth, the setae being long and stout. External subhumeral stria (Fig. 39A) complete and densely covered with coarse punctures. Internal subhumeral stria usually present on basal third, but absent on humeral area (nearly basal sixth). First dorsal stria nearly complete, but abbreviated on basal sixth (humeral area) and apical fourth. Second dorsal stria complete, usually abbreviated on apical sixth. Third dorsal stria present on apical half, but shortened on apical sixth, and short rudiments present at base and on median third of apical half. Fourth dorsal stria absent, sometimes with a short rudiment which consists of two or three punctures occurring on apical sixth. Fifth dorsal stria present on about median third, but sometimes shortened apically, and a short rudiment present on apical fifth. Sutural stria complete. All striae deeply impressed and coarsely punctate. Intervals among the striae sparsely clothed with microscopic punctures.
Pygidium (Fig. 39C) deeply depressed on each lateral side, the basal end of the depression not attaining to the base; its sides strongly carinate; surface densely covered with coarse and deep punctures, and other fine punctures intermingled among coarse ones, these punctures becoming denser and finer basally.
Antennal grooves present on anterior area of underside of head, the grooves strongly carinate basally and running obliquely from anterior margin of eye to apical third of head on mid line.
Anterior margin of prosternal lobe outwardly acute at middle, and with long hair; disk of the lobe impunctate. Prosternal process divided into anterior and posterior parts by procoxae; apical two-thirds of the anterior part elevated and with smooth disk, and the basal third triangular, its basal margins strongly convergent along procoxae, the apical margin of process with two or three long setiferous punctures; the posterior part short, oblong and impunctate.
Mesosternum moderately convex on median area; surface densely covered with coarse punctures, the punctures becoming denser medially, producing many rugae; area of these rugae clothed with other dense and fine microscopic rugae; area before posterior corner and between mesocoxae impunctate.
Meso-metasternal suture shortly present. Metasternum deeply with arcuate depression between mesocoxae; intercoxal disk shallow and logitudinally sulcate on longitudinal mid line, and impunctate; lateral disk densely covered with coarse and shallow punctures which are shallower, sparser and finer posteriorly. Metepisternum somewhat sparsely covered with coarse, transversely oblong and shallow punctures which are a little coarser than those of lateral disk of metasternum.
Table 5. Biometeric data of Syntelia histeroides Lewis.
-----------------------------------------------------------------------
APW 3.00-4.38 (3.640.08) 20
PL 2.50-3.75 (3.150.07) 20
EL 4.88-6.63 (5.840.10) 20
EW 3.25-4.75 (4.090.08) 20
PyW 2.00-3.13 (2.610.05) 20
PyL 2.13-3.13 (2.610.05) 20
PTL 1.75-2.50 (2.190.05) 20
MSTL 1.88-2.50 (2.110.04) 20
MTTL 1.88-2.75 (2.300.05) 20
-------------------------------------------------------------------------

First abdominal sternum densely and coarsely punctate except on medioposterior area. Second - 5th abdominal sterna densely and coarsely punctate laterally.
Protibia (Fig. 39E, D) with 5 denticles on outer margin. Mesotibia with 4 denticles on outer margin, the basal one small. Metatibia with two denticles on apical third of outer margin.
Male genitalia as shown in Fig. 40A - G.
Female genitalia: spermatheca as shown in Fig. 40H.
Specimens examined. [Hokkaid]] 8 exs., Nopporo, 19, 20, 22, 26/vi, 24/vii/1987, M. hhara leg.; 1 ex., Ditto, 30/vi/1987, S. Kud leg.; 10 exs., Eniwa, Isarigawa, 16/vi/1986, M. Mori leg.; 1 ex., Maruyama, Sapporo, 3/vii/1985, M. hhara leg.; 1 ex., Ditto, 22/vi/1940, B. Nishio leg.; 1 ex., Mt. Moiwa, Sapporo, 5/vii/1978, T. Fujisawa leg.; 2 exs., Ditto, 15/vi/1904, no collector's name (NSMT); 1 ex., Heiwa-no-taki, Sapporo, 2/viii/1977, T. Fujisawa leg.; 2 exs., Ditto, 1/vi/1966, N. Ueda leg.; 1, Sapporo, 31/v/1976, M. Furukawa leg.; 5 exs., Hakkenzan, Jzzankei, 7/vi/1976, 16/vi/1982, M. Mori leg.; 3 exs., Ditto, 11, 24/vi/1978, T. Fujisawa leg.; 1 ex., Misumai, Sapporo, 10/vi/1979, A. Shinohara leg. (NSMT); 3 exs., Jzzankei, 17/vi, 13/vii/1985, M. hhara leg.; 1 ex., Shikotu-ko, 6/viii/1971, Y. Doi leg. (NSMT); 1 ex., Enshrrin, Tomakomai, 3/viii/1974, A. Kashizaki leg.; 1 ex., Ditto, 18-30/viii/1978, H. Detani leg.; 2 exs., Okusawa, Otaru, 29/vii/1955, Y. Watanabe leg.; 5 exs., Mt. Yotei, Hirafu, Kutchan, 15-16, 22/vii/1991, M. hhara leg.; 1 exs., Daisengen, 17/vii/1973, T. Ichikawa leg.
[Honsh]] <Miyagi-ken> 1 ex., Moriyasu, Akiu-mura, 24/vi/1951, no collector's name (NSMT). <Tochigi-ken> 1 ex., Chzzenji, 14/viii/1917, E. Gallois leg. (NSMT). <Gumma-ken> 1 ex., Hsshi-onsen, 6/vi/1953, Y. Kurosawa leg. (NSMT). <Saitama-ken> 1 ex., Oku-Chichibu, 24/vii/1940, K. Kojima leg. (NSMT). <Nagano-ken> 1 ex., Mt. Yatsu-ga-take, 25/vii/1957, S. Nomura leg. (NSMT); 1 ex., Mt. Chausu-dake, Yokokubo-zawa, 1600m, S. Jap. Alps, 24/vii/1940, S. Unno leg. (NSMT).
Distribution (Fig. 41). Japan (Hokkaid, Honsh, Kyssh).
Remarks. Syntelia histeroides is a distinctive species within the superfamily due to its shape and rather large size. It cannot be confused with any other species of the histerids.
This species frequents in sap of elm and under bark. Larva is found under bark.


6. 3. Family Histeridae Gyllenhal, 1808

Key to the Japanese subfamilies of Histeridae

1(2) Prosternum without antennal grooves or cavities. Ventral side of head with large foveae for reception of antennae. Mandible vertically connected with head. ........................................................... Subfamily Niponiinae
2(1) Prosternum with antennal grooves or cavities. Ventral side of head without foveae (except in Tribes Hololeptini and Histerini, which have shallow and narrow longitudinal grooves). Mandibles porrect, horizontally connected with head.
3(12) Antennal grooves or cavities on prosternum transverse, occurring on anterior side, and usually closed beneath by the prosternal alae.
4(9) Lbrum with a few setiferous punctures.
5(6) Lateral sides of elytra strongly elevated. Prosternal lobe absent, not distinctly separated by prosternal suture. ....................... Subfamily Chlamydopsinae
6(5) Lateral sides of elytra not elevated. Prosternal lobe present.
7(8) Elytra without costa, usually with normal striae or punctures. .................... ................................................................... Subfamily Tribalinae
8(7) Elytra with costae. ........................................ Subfamily Onthophilinae
9(4) Labrum without setae.
10(11) Antennal scape expanded and strongly angulate. ......... Subfamily Hetaeriinae
11(10) Antennal scape normal, neither expanded nor strongly angulate. ................. ................................................................... Subfamily Histerinae
12(3) Antennal grooves on prosternum longitudinal, usually situated next to prosternal keel, and open beneath.
13(14) Prosternal lobe present. ................................. Subfamily Dendrophilinae
14(13) Prosternal lobe absent.
15(18) Body oval or oblong-oval.
16(17) Dorsal elytral striae absent though sometimes represented by rather vague impressions. .................................................... Subfamily Abraeinae
17(16) Dorsal elytral striae present. ................................. Subfamily Saprininae
18(15) Body cylindrical.
19(20) Antennae consisting of eight segments and a club. Head horizontal in repose. Anterior margin of prosternal alae at most with only very slightly notched. Antennal cavities not distinct. ............................. Subfamily Trypanaeinae
20(19) Antennae consisting of seven segments and a club. Head vertical in repose. Anterior margin of prosternal alae with rather deep longitudinal incisions for reception of the antennal funicles in repose. ............. Subfamily Trypeticinae

6. 3. 1. Subfamily Niponiinae Fowler, 1912

Genus Niponius Lewis, 1885

Niponius Lewis, 1885: 333 [type species: Niponius impressicollis Lewis, 1885b: 333. Designated by Gandner, 1935: 4]; Marseul, 1888: 1489; Jakobson, 1911: 638, 642; Gardner, 1926: 194; 1935: 2; Reichardt, 1926: 274; 1941: 66; Schenkling, 1931: 1; Kryzhanovskij and Reichardt, 1976: 76.
<Larva> Gardner, 1930: 15, 17, t.1, fig. 1; Hayashi, 1986: pl. 10.

Gardner's (1935) description. "Cylindrical, slender to moderately stout, shining. Head exserted, nearly as wide and thick at base as, and subequal in length to, prothorax; produced anteriorly into two horn of variable length, each horn carinate at apex and usually with one or two postapical carinae; a groove passing back just below each eye, receives the antennal scape when at rest. Eyes small, entire, finely facetted, well removed from anterior margin of prothorax. Antenna geniculate, the scape about as long as remaining segments together; funicle with six segments, the first angled, longer than wide, remaining segments transverse, gradually increasing in size towards the club; club compact, flattened, more or less circular, apparently four-segmented, the three sutures pale, curved, each with a fringe or setae; basal segment of club large, the others strongly transverse; in addition a fourth row of setae, very near the apex, might be interpreted as defining an extremely narrow fifth segment of the club. Labrum small, vertical, triangular. Mandibles stout, ventrally directed, inwardly curved, each with two small teeth near the middle. Maxillae with lacinia rather extending far beyond lacinia, slender, tapering to apex, densely ciliate. Maxillary palps four-segmented, the apical segment about equal in length to the three preceding segments together; the palp is borne on a segment-like palpifer which is distinctly longer than wide. Submentum a sclerotized transverse plate, the anterior margin with a rather wide emargination at the base of which are two teeth or angulations of variable size. Labial palp with three segments, the apical as long as the other two together; each palp at the apex of a parallel-sided palpiger which incurves and approaches its fellow on the buccal surface of submentum; attached to the palpigers are the widely separated membranous, ciliate paraglossae; ligula not distinct.
Prothorax margined, cylindrical, anterior margin straight with no projection over ventral part of head; slightly transverse to longer than wide; coxal cavities very narrowly closed posteriorly, separated from one another by a prosternal process which bears two longitudinal grooves or striae. Mesosternum very short, with median channel. Metasternum large, with median channel; posterior coxal cavities moderately widely separated. Elytra truncate, abbreviated, exposing two posterior abdominal terga, loner than prothorax and usually longer than their combined width; often with lines of punctures giving the appearance of more or less distinct striae. Pygidium and propygidium usually with two to four foveae. Abdomen with five apparent sterna. Legs with slender five segmented tarsi, the apical segment very long; tibiae with two apical spurs, one much larger than the other and especially strong on anterior tibiae. Anterior leg with transverse coxa; tibia with outer margin rather strongly expanded on distal half, usually with two teeth, each borne in a socket, one near apex and one near middle; with a longitudinal groove for reception of tarsus. Mesothoracic tibia less strongly expanded, with three socketted teeth on outer margin, two near apex and one nearer the middle. Posterior tibia slightly and gradually widened from base to apex, with a pair of subapical socketted teeth and sometimes a minute tooth near middle. Male genitalia as described by Sharp and Muir (loc. cit.): (( Median lobe tubular, slender and long; lateral lobes longer than median lobes and enveloping them. Basal piece forming a long tube, constricted near its base and bent. Internal sac undifferentiated))".

Key to the Japanese species of the genus Niponius

1 (2) Body 4.8 - 5.5 mm long. Pronotum with a deep excavation on each side. Foveae on propygidium four. .................... N. impressicollis Lewis, 1885
2(1) Body 3.5 - 4.7 mm long. Pronotum with no deep excavation.
3(4) Projection of epistoma short and stout (Fig. 45A). Propygidium without fovea. ............................................................ N. obtusiceps Lewis, 1885
4(3) Projection of epistoma long and slender (Fig. 42A, 46A). Propygidium with several excavations.
5(6) Projections of epistoma divergent anteriorly. Surface of elytra wholly with rows of punctures. Propygidium with 4 foveae. Antennal grooves very deep under the eyes, and other longitudinal foveae present on basal half of head ..... ............................................................... N. furcatus Lewis, 1885
6(5) Projections of epistoma parallel to each other. A row of punctures on elytral surface present along sutural line, others not clear. Propygidium with 2 to 4 foveae. Antennal grooves confined to under the eyes, without longitudinal deep foveae on basal half of head. ................. N. osoriocepus Lewis, 1885

Niponius furcatus Lewis, 1885
(Fig. 42, 48)

Niponius furcatus Lewis, 1885b: 333 [Japan, Yuyama]; Gardner,1926: 3 1935: 5 [catalogued]; Kryzhanovskij and Reichardt, 1976: 77 [key]; Hisamatsu, 1985: 219, pl. 40, fig. 4.
Japanese name: Fushi-tsuno-hoso-emma-mushi.

Original description. "Cylindricus, angustatus, niger, punctatus, antennis, pedibusque piceis, thorace grosse punctato, sine fovea, propygidio quadrisulcato pygidio biimpresso. Long. 4.1/4 mm.
The narrow form of this species, the divergent direction of the projection on the epistoma, and the different form of the abdominal sulci separate it form the others."

Description. Biometric data are given in Table 6. Body cylindrical, slender. Cuticle shining, black; tarsi, antennae and projection of epistoma dark rufopiceous.
Projections of epistoma moderately stout, long and divergent anteriorly (Fig. 42A, E), its apex strongly carinate and dorsal surface with two transverse carinae. Head densely covered with moderate punctures and with several weakly rugae on base of the projection, the punctures becoming coarser on basal two-thirds, where they are separated by two to three times their diameter.
Pronotum completely with marginal stria on lateral side, the stria well carinate; surface densely covered with punctures, the punctures various in size from fine to large (the large punctures are five times as large as the fine ones) and becoming finer around margin.
Marginal elytral stria on epipleura of elytra complete and carinate, its basal end extending inwardly along anterior margin of elytron and united with basal end of sutural stria, its apical end attaining to the apical fifth of elytron. All dorsal striae absent, but there are rows of coarse and dense punctures; intervals among the rows sparsely covered with coarse punctures, the punctures becoming denser apically.
Propygidium (Fig. 42D) with four large, longitudinal oblong, shallow and transversely placed foveae, sometimes these foveae fused; surface sparsely covered with punctures, which are as coarse as elytral ones and separated by two to three times their diameter. Pygidium (Fig. 42D) with a large, round and shallow fovea behind each basal corner; inside of the fovea sparsely covered with coarse punctures; surface coarsely punctate, the punctures separated by their diameter and becoming finer and denser apically and laterally.
Antennal grooves deep under the eyes, and deep and longitudinal foveae on basal half of underside of head (Fig. 42B). Mid line strongly impressed on apical half, but not impressed near the apical margin of the head. Surface of underside of head evenly scattered with fine punctures on apical half; on basal half very sparsely clothed with fine punctures, the punctures becoming coarser and denser near the basal margin.
Prosternal lobe (Fig. 42C) transverse and narrow, its anterior margin densely with hair. Prosternal keel narrow, with carinal stria on basal three-fourths, the striae deeply impressed and slightly convergent basally; surface of keel evenly and finely punctate.
Mesosternum longitudinally sulcate on mid line; surface sparsely covered with moderate punctures which are separated by two to three times their diameter, and with large punctures near basal margin. Meso-metasternal suture well impressed and complete. Metasternum shallowly sulcate on mid line; lateral metasternal striae strongly carinate and divergent posteriorly. Intercoxal disk of metasternum sparsely covered with moderate sized punctures. Lateral disk densely covered with large, round and shallow punctures on basal third, the punctures becoming finer and sparser posteriorly.
Intercoxal disk of 1st abdominal sternum striate on basal two-thirds on each lateral side, and evenly and moderately punctate, the punctures separated by two to three times their diameter.
Protibia slender and with two denticles on outer margin.
Specimens examined. [Kyssh]] <Kagoshima-ken> 1, Kirishima-Jing, 9/vi/1981, T. & T. Nakane leg. (NA).
[Nansei Isles.] <Tokuno-shima Is.> 1, Miky, 25/vii/1963, Y. Kurosawa leg.
<? Shikoku, Kcchi-ken> 1 ex., Wada, Taish-mura, 4/x/1936, no collecters name, collected from Tsuga sieboldi. (NSMT: Knno collection, No. NSMT-C-23669).
Table 6. Biometric data of Niponius furcatus Lewis.
------------------------------------------------------------------
HOW 0.44-0.48 (0.450.010) 2
HW 0.73-0.76 (0.750.010) 2
PW 0.79-0.85 (0.820.021) 2
PL 1.14 (1.14 ) 2
EL 1.14-1.24 (1.190.036) 2
EW 0.85-0.89 (0.870.016) 2
ProW 0.50 (0.50 ) 2
ProL 0.26 (0.26 ) 2
PyL 0.44-0.48 (0.450.010) 2
PTL 0.56-0.57 (0.560.005) 2
MSTL 0.48-0.50 (0.490.005) 2
MTTL 0.47-0.50 (0.480.010) 2
----------------------------------------------------------------------

Distribution (Fig. 48). Japan (Shikoku, Kyssh, Amami-sshima Is., Tokuno-shima Is.).
Remarks. Niponius furcatus is characterized by the narrow body, the diverse projections of the epistoma and the four foveae on the propygidium.

Niponius impressicolis Lewis, 1885
(Fig. 43, 44, 48)

Nioponius impressicolis Lewis, 1885b: 333 [Japan, Yuyama, in Higo to Junsai in Yezo (= Kyssh to Hokkaid)]; Gardner, 1926: 2; Reichardt, 1929b: 274, 273; Gardner, 1935: 5; Reichardt, 1941: 69; Nakane, 1963: 67, pl. 34, fig. 3; Kryzhanovskij and Reichardt, 1976: 77 [key, description]; Hisamatsu, 1985: 219, pl. 40, fig. 3.
Japanese name: Hoso-emma-mushi.

Original description. "Cylindricus, sat elongatus, robustus, niger, punctatus. Antennis pedibusque, nigro-piceis, clava tarsisque rufis; epistomo cornuto, piceo, transversim bicarinato, thorace grosse punctato, utrinque profunde impresso. Elytris striis 1 validis et caeteris punctiformibus vel obsoletis. Propygidio quadri foveolate. Pygidio profunde bisulcato. Long. 5 - 5.1/2 mm."

Description. Biometric data are given in Table 7. Body cylindrical, moderately stout. Cuticle shining, black; tarsi, antennae and projection of epistoma dark rufopiceous.
Projection of epistoma (Fig. 43A, C) moderately stout and long, its apex strongly carinate and dorsal surface with two or three transverse carinae. Head densely with transverse rugae on apical third; median third of the surface sparsely covered with moderate punctures, the punctures separated by two to five times their diameter, with other fine punctures intermingled; basal third finely and sparsely punctate; depression on mid line deeply present on apical half.
Pronotum completely with marginal stria on lateral side, the stria strongly carinate; surface with an excavation (Fig. 43D, E) on each mediolateral side, and sparsely covered with large, round and deep punctures which are irregularly scattered, and with other fine punctures sparsely intermingled among the large ones.
Epipleura of elytra completely with marginal elytral stria, which is carinate and of which the apical end attains to the apical fifth; area between epipleural margin and elytral marginal stria impunctate and extremely finely strigate. First dorsal stria present on basal fourth and strongly carinate, its basal end deeply excavate. Second to 5th dorsal striae almost obsolete, being represented by rows of coarse punctures. Sutural stria strongly impressed and represented by a row of longitudinal coarse punctures on basal two-thirds. Surface of elytra somewhat convex on humeral area, and deeply and transversely excavated near the basal margin; represented by rows of coarse punctures; intervals among the rows sparsely clothed with fine punctures on basal half, and densely on apical half, the punctures becoming coarser on apical third, and in lateral area these punctures somewhat much denser.
Propygidium (Fig. 43G) with four large, round, shallow and transversely placed foveae which become shallower posteriorly; surface sparsely covered with coarse punctures, which are as coarse as elytral ones, and other fine punctures densely intermingled among the coarse ones. Pygidium (Fig. 43G) with a large, round, and somewhat deep fovea behind each basal corner; inside of the fovea sparsely and moderately punctate; surface coarsely punctate on median area, the punctures separated by three to four times their diameter and becoming finer anteriorly, and other fine punctures sparsely intermingled among coarse ones, the fine ones becoming denser apically and laterally.
Antennal grooves (Fig. 43B) deep under eyes. Mid line of underside of head deeply excavate, its basal end beyond transverse line. Surface of underside of head shining, sparsely and finely punctate on apical half, and evenly clothed with fine punctures which are separated by two to three times their diameter on basal half.

Table 7. Biometric data of Niponius impressicollis Lewis.
-------------------------------------------------------------------
HOW 0.57-0.70 (0.650.021) 5
HW 1.09-1.25 (1.190.026) 5
PW 1.30-1.56 (1.460.042) 5
PL 1.27-1.66 (1.510.067) 5
EL 1.61-1.93 (1.770.061) 5
EW 1.40-1.66 (1.540.051) 5
ProW 0.83-0.94 (0.880.021) 5
ProL 0.57-0.42 (0.480.028) 4
PyL 0.73-0.57 (0.640.023) 5
PTL 1.01-0.88 (0.960.025) 4
MSTL 0.83-0.78 (0.790.011) 4
MTTL 0.96-0.78 (0.880.033) 4
---------------------------------------------------------------------

Prosternal lobe (Fig. 43F) transverse and narrow, its anterior margin densely with hair. Prosternal keel completely with two carinal striae, of which the basal ends are united with each other in an arch; surface of keel evenly punctate as basal half of head.
Mesosternum longitudinally and shallowly sulcate on median third; lateral third evenly punctate as basal half of head and extremely finely strigate; sometimes lateral marginal stria present on basal half at each anterior corner. Meso-metasternal suture well impressed and complete. Metasternum longitudinally and shallowly sulcate on mid line; lateral metasternal striae strongly carinate and divergent posteriorly. Intercoxal disk of metasternum sparsely and coarsely punctate, the punctures twice as coarse as mesosternal punctures, and other fine ones sparsely intermingled among them. Lateral disk sparsely covered with large punctures, the punctures becoming finer apically.
Intercoxal disk of 1st abdominal sternum deeply striate on basal three-fourths on each lateral side, and punctate as intercoxal disk of metasternum, but the punctation is much denser.
Protibia slender and with two denticles on outer margin.
Male genitalia as shown in Fig. 44.
Specimens examined. [Honsh]] <Fukushima-ken> 2 exs., Yunohana, Tateiwa V. Minami-Aizu, 19/vi/1947, Y. Kurosawa leg.; 4 exs., Onozawa, Egawa, Minami-Aizu, 17/vi/1950, Y. Kurosawa leg. (NSMT). <Mie-ken> 2 exs., Mimune, 18-21/vi/1955, H. hhira leg. <Hiroshima-ken> 2, Sandanky, Aki, 7-8/viii/1966, S. ssawa leg.
Distribution (Fig. 48). Japan (Hokkaid, Honsh, Shikoku, Kyssh); Taiwan; Ussuriyskiy Kray; China.
Remarks. Niponius impressicollis is the largest species of the genus in Japan. It can easily be recognized by its size and the presence of excavations on the pronotum.
This species is found from the galleries of a scolid beetle, Hylesinus striatus Egg. on ash, Fraxinus mandchuria (Kryzhanovskij and Reichardt, 1976).

Niponius itoi Chjj, 1955

Niponius itoi Chjj, 1955: 57 [Japan, Adera, kkuwa-mura, Nishichikuma-gun, Nagano pref., Honsh]]; Hisamatsu, 1985: 220 [=? Niponius osorioceps Lewis].
Japanese name: It-hoso-emma-mushi.

Specimens examined. No specimens of the species have been available for my study.
Distribution. Japan (Honsh).
Remarks. Judging from the original description this species is apparently similar to N. osorioceps, but it is distinguished from latter by the smaller body size. I have been unable to trace it in the Chjj collection. Accoding to Hisamatsu (1985) this species is probably a synonym of N. osorioceps.

Niponius obtusiceps Lewis, 1885
(Fig. 45, 48)

Niponius obtusiceps Lewis, 1885b: 334 [Japan, Oyayama near Kumamoto in Higo, and Ishikari river in Yezo in 1883]; Gardner, 1926: 3; Reichardt, 1929b: 275; Gardner, 1935: 5, t. 1, fig. 2; Reichardt, 1941: 69; Kryzhanovskij and Reichardt, 1976: 78; Hisamatsu, 1985: 220 [key].
Japanese name: Tsuno-buto-hoso-emma-mushi.

Original description. "Cylindricus, elongatus, brunneus, punctatus. Antennis pedibusque fufis, epistomo obtuso subfurcato, thorace grosse necnon minute punctato. Elytris fasciis transversis medio nigris. Propygidio punctato, pygidio bisulcato. Long. 4 mm."

Description. Body cylindrical, moderately stout. Cuticle shining, black; tarsi, antennae and projection of epistoma dark rufopiceous.
Projection of epistoma (Fig. 45 A, C) stout and long, its dorsal surface with two transverse carinae. Apical third of head densely covered with fine punctures and weakly transverse rugae, the punctures becoming sparser and coarser posteriorly; median third of the surface coarsely punctate, the punctures being separated by their own diameter to twice the diameter; basal third with much smaller punctures than on median third, the punctures being separated by three to five times their diameter; depression on mid line deep on apical fourth.
Pronotum completely with marginal stria on lateral side, the stria strongly carinate; surface irregularly scattered with deep, large and somewhat longitudinal oblong punctures (Fig. 45D) and other fine punctures intermingled among the large ones.
Epipleura of elytra completely with marginal elytral stria, which is carinate and of which the apical end attains to near the posterior margin of elytron; area between epipleural margin and elytral marginal stria impunctate and extremely finely strigate. All dorsal and sutural striae represented by rows of large punctures, the rows obsolete and their punctures becoming finer on apical third; intervals among the rows impunctate on basal two-thirds, but sometimes sparsely with several fine punctures, which are as large as the pronotal fine ones, separated by their own diameter to twice the diameter and become denser and finer posteriorly. Surface of elytra with weakly convex on humeral area, and deeply transverse excavation on near the basal margin.
Propygidium (Fig. 45F) without large foveae, and sparsely covered with coarse punctures which are twice as large as fine punctures of elytra, and other fine punctures densely intermingled among the coarse ones. Pygidium (Fig. 45F) with large, semicircular and deep foveae on basal two-thirds on lateral area; area inside fovea sparsely and finely punctate; surface between foveae sparsely covered with coarse punctures and intermingled fine ones among the coarse ones; fine punctures present also on apical third and along the margin.
Antennal grooves divided two parts; a deep furrow under eyes, and an oblong fovea on basal half of underside of head (Fig. 45B); surface of the groove densely clothed with fine punctures on apical half, the punctures much sparser on basal half.
Prosternal lobe (Fig. 45E) transverse and narrow, its anterior margin densely furnished with hair. Prosternal keel completely with two carinal striae, of which the basal ends are united with each other in an arch; surface of keel sparsely clothed with fine punctures.
Mesosternum longitudinally and shallowly sulcate on median third; area of lateral third sparsely clothed with several fine punctures. Meso-metasternal suture absent. Metasternum longitudinally, broadly and shallowly sulcate on mid line; lateral metasternal stria strongly carinate, its apical end extending posteriorly, and attaining to near apical fourth of metasternum. Intercoxal disk of metasternum sparsely covered with coarse and longitudinal oblong punctures. Lateral disk densely covered with coarse and round punctures, which are a little coarser than those of intercoxal disk, the punctures becoming sparser along the lateral metasternal stria.
Intercoxal disk of 1st abdominal sternum deeply striate on each lateral side, the striae nearly complete, but not attaining to the apical margin; surface densely and finely punctate, the punctures progressively sparser and finer posteriorly.
Protibia slender and with three denticles on outer margin.
Specimens examined. [Honsh]] <Tkky-to> 1 ex., Mt. Takao, 19/ii/1981, K. Horie leg. <Fukui-ken> 1, Ikegahara, nno-shi, 20/vi/1988, M. Sait leg.
Distribution (Fig. 48). Japan (Honsh).
Remarks. Niponius obtusiceps can readily be recognized by the absence of fovea on the propygidium and the short and stout projections of the epistoma.
No details are known about the habitat of this species. Specimens were collected under the bark. Hisamatsu (1985) noted that this species was found in tunnels made by cossonine beetles, Curculionidae.

Niponius osorioceps Lewis, 1885
(Fig. 46, 47, 48)

Niponius osorioceps Lewis, 1885b: 333 [Japan, Higo, Yuyama and Konose], pl. 8, fig. 12-14; Gardner, 1935: 3; Nakane, 1963: 67, pl. 34, fig. 4 [noted, photo]; Hisamatsu, 1985: 219, 220, pl.40, fig. 5.
Niponius osoriiceps (sic): Jakobson, 1911: 642; Reichardt, 1929: 274; Kryzhanovskij and Reichardt, 1976: 78.
Japanese name: Hime-hoso-emma-mushi.

Original description. "Cylindricus, elongatus, niger, punctatus, pedibus nigro-brunneis, epistomo piceo transversim tricarinato, thorace grosse punctato utrinque obsolete impresso. Elytris striis N. impressicollis simillimis. Propygidio pygidioque profunde bifoveolatis. Long. 4.1/2 mm."

Description. Biometric data are given in Table 8. Body cylindrical, moderately stout. Cuticle shining, black; tarsi, antennae and projection of epistoma dark rufopiceous.
Projection of epistoma (Fig. 46A, C, D) moderately stout and long, its apex strongly carinate and dorsal surface with two transverse carinae. Apical third of head densely covered with transverse rugae; median third evenly and coarsely punctate, the punctures being separated by their diameter; basal third densely covered with coarse punctures, which are a little coarser than the median ones; depression on mid line present and shallow on apical third.
Pronotum completely with marginal stria on lateral side, the stria strongly carinate; surface covered with coarser punctures which are as coarse as those on the basal third of the head, the punctures being separated by their own diameter to twice the diameter.
Epipleura of elytra completely with marginal elytral stria, which is strongly carinate and of which the apical end attains to near the apical fifth; area between epipleural margin and elytral marginal striae impunctate and extremely finely strigate. All dorsal striae represented by rows of moderate punctures, the rows obscure and not clear, and their punctures a little finer than the pronotal ones and irregularly intermingled with punctures of intervals; intervals among the rows sparsely covered with moderate punctures, the punctures becoming denser laterally and apically. Sutural stria represented by a row of coarse and deep punctures, the row being somewhat clear. Surface of elytra with a deeply transverse excavation on near the basal margin.
Propygidium (Fig. 46E) with two large, transverse oblong, or four longitudinal oblong, and shallow foveae which become shallower posteriorly; surface densely covered with moderate punctures, the punctures becoming sparser on area between the foveae. Pygidium (Fig. 46E) with large, longitudinal oblong and deep foveae behind each basal corner; area inside foveae sparsely covered with moderate punctures; surface between foveae coarsely punctate, the punctures being separated by two to three times their diameter, and other moderate punctures sparsely intermingled among the coarse ones, these punctures becoming finer and denser apically and laterally.
Antennal grooves (Fig. 46B) deep under eyes. Surface of underside of head sparsely clothed with fine punctures, the punctures separated by two to four times their diameter and becoming denser basally and laterally.
Prosternal lobe (Fig. 46F) transverse and narrow, its anterior margin densely with hair. Prosternal keel completely with two carinal striae, which are strongly carinate and slightly convergent basally.
Mesosternum longitudinally and shallowly sulcate on mid line, its disk sparsely and finely punctate. Meso-metasternal suture clearly impressed. Metasternum longitudinally, shallowly and completely sulcate on mid line; lateral metasternal striae strongly carinate and divergent posteriorly, its apical end extending to near the hind coxa. Intercoxal disk of metasternum sparsely clothed with fine punctures, the punctures being separated by two to five times their diameter, and becoming denser and a little coarser on apical fourth. Lateral disk irregularly scattered with large, round and deep punctures on basal third, the punctures becoming finer apically and along the metasternal lateral stria.
Intercoxal disk of 1st abdominal sternum deeply striate on lateral sides, the striae nearly complete and strongly divergent posteriorly; surface densely and moderately punctate, the punctures as large as those on apical fourth of intercoxal disk of metasternum and separated by their own diameter to twice the diameter; surface among the punctures extremely finely strigate.
Table 8. Biometric data of Niponius osorioceps Lewis.
---------------------------------------------------------
HOW 0.58-0.50 (0.530.006) 18
HW 0.96-0.75 (0.890.012) 19
PW 1.15-0.92 (1.040.015) 18
PL 1.29-1.02 (1.150.017) 19
EL 1.61-1.20 (1.430.024) 19
EW 1.24-0.96 (1.140.019) 18
ProW 0.75-0.53 (0.650.013) 18
ProL 0.41-0.26 (0.350.009) 17
PyL 0.61-0.44 (0.530.013) 17
PTL 0.73-0.61 (0.670.012) 15
MSTL 0.70-0.53 (0.640.013) 15
MTTL 0.76-0.58 (0.680.012) 17
---------------------------------------------------------------

Protibia slender and with three denticles on outer margin.
Male genitalia as shown in Fig. 47.
Specimens examined. [Hokkaid]] 2 exs., Gamushi, 10/v/1956, A. Nobuchi leg.
[Honsh]] <Fukushima-ken> 1 and 7 exs., Yunohana, Tateiwa, Minami-Aizu, 15/vi, 18-20/vii/1947, Y. Kurosawa leg.; 2 exs., Ditto, 9/vii/1948, T. Nakane leg.; 1 ex., Onozawa, Egawa, Minami-Aizu, 2/vi/1951, Y. Kurosawa leg. (NSMT); 1 ex., Ditto, 17/vi/1950, Y. Kurosawa leg. (NSMT). <Yamagata-ken> 1 ex., wwisawa, 28/vi/1960, Y. Kurosawa leg. (NSMT); 1 ex., iishi, Oguni, 9/vi/1969, K. Shirahata leg. (NSMT). <Gumma-ken> 1 ex., Utsunomiya, 6/v/1968, H. Takizawa leg. <Saitama-ken> 1 ex., Chichibu, Musashi, 17/vi/1913, H. Takabayashi leg. <Tkky-to> 2 exs., Hikagesawa, Mt. Takao-san, 4/v/1975, K. Kawada leg.; 1 ex., Hikawa, Okutama, 13/v/1951, K. Umeya leg. (NSMT); 1 ex., Mt. Takao, 21/vi/1963, S. Nomura leg. (NSMT). <Kanagawa-ken> 11 exs., Mt. yyama, 5/v/1989, H. Takizawa leg.; 1 ex., Suun-zan, 29/v/1988, H. Takizawa leg. <Nagano-ken> 1 ex., Yatsu-ga-take, 14/vii/1951, J. Suzuki leg. (NSMT). <Shizuoka-ken> 1 ex., Izu, Inatori, 6/v/1973, H. Takizawa leg.; 1 ex., Mt. Amagi-san, 5/v/1956, S. Tsuyuki leg. (NSMT). <Niigata-ken> 1 ex., Kurokawa, northen Echigo, 9/vi/1963, K. Baba leg. <Gifu-ken> 1 ex., Naratoge, 4/vii/1953, H. Torigai leg. <Shiga-ken> 1 ex., Bomura, 15/vi/1957, T. Nakane leg. <Kytto-fu> 1 ex., Serio, 29/v/1954, T. Nakane leg. <Tottori-ken> 1 ex., Daisen, 9/vii/1951, N. Yato leg.
[Kyssh]] <Kagoshima-ken> 1 ex., Iriki, Aira-gun, 10/v/1984, T. & T. Nakane leg. (NA).
Distribution (Fig. 48). Japan (Hokkaid, Honsh, Kyssh); Ussuriyskiy Kray.
Remarks. Niponius osorioceps is characterized by the long and slender projections of the epistoma and the presence of two to four foveae on the propygidium.
This species is known as a predator of the bark beetle Phloeosinus perlatus Chapuis, Scolytidae.

6. 3. 2. Subfamily Chlamydopsinae Bickhardt, 1917

Genus Eucuritopsis Silvestri, 1926

Description. Characterized by unique shape of pronotum and elytra. Medio-apical portion of pronotum strongly elevate. Elytral lateral ridge strongly elevate and interrupted transversely medially; the inside of the interrpted portion furnished densely with waxed hairs.
Comments. The genus is a fairly small one containing two species, E. mirabilis Silvestri, 1926 from Taiwan and E. japonicus sp. nov. from Japan. Species of this genus occur in the nest of ants. E. mirabilis is collected from the nest of Formica legi.

Eucuritopsis japonicus M. hhara et Nakane, sp. nov.
(Fig. 49)

Japanese name: Arinosu-kobu-emma-mushi.

Description. Body oblong, reddish brown, shining and furnished with yellow hairs. Body length, width and biometric data are as follows (male: M, n=1; female: F, n=1): PPL, M 1.84, F 2.06; PEL, M 1.79, F 1.88; APW, M 0.69, F 0.71; PPW, M 0.86, F 0.91, PL, M 0.74, F 0.66, EL, M 1.10, F 1.23, EW, M 1.37, F 1.39, ProW, M 0.66, F 0.71, ProL, M 0.34, F 0.39, PyL, M 0.34, F 0.39, PTL, M 0.54, F 0.49, MSTL, M 0.51, F 0.46, MTTL, M 0.54, F 0.49.
Frontal lateral margins (Fig. 49A) parallel and carinate; disk densely covered with coarse, round punctures. Labrum semicircular. Mandible short and robust. Scape of antenna large, thick and triangular in frontal view, and coarsely and densely punctate.
Pronotal sides (Fig. 49B) feebly convergent apically, the basal fourth slightly emarginate to fit forefemur in repose; anterior angle strongly emarginate; disk convex, the medio-apical area strongly elevated and on the mid line feebly depressed; surface densely covered with round, large and deep punctures, sparsely intermingled with fine punctures among the large ones, and densely furnished with yellow long hairs; posterior margin angulate at middle.
Epipleura flat. Epipleural marginal stria strongly sinuate medially, the apical end extending across elytral apex to medio-apical angle of elytron and united with the apical end of the sutural stria. Elytral marginal stria absent. Elytron (Fig. 49B) with lateral margin raised into a narrow, prominent ridge on lateral third; the ridge narrowly interrupted medially; furnished with waxed hairs inside the median interrupted portion; surface of the basal half of the ridge with a narrow elevated carina; mediobasal half of the elytral disk depressed, shining and sparsely furnished with short and thick hairs basally; apical half of the disk convex and evenly covered with coarse, setiferous punctures, which are separated by about their own diameter, except inside the apical half of the lateral elevated ridge and in a longitudinal band narrowly distant from the suture, the punctured area extending basally narrowly along the suture and reaching to basal fourth. Sutural stria complete and closely impressed along the suture.
Propygidium feebly convex, densely covered with large, round and shallow punctures which are separated by their own diameter to one-third the diameter, and sparsely intermingled with fine and setiferous punctures. Pygidium (Fig. 49C) has a punctation simiar to that of the propygidium.
Anterior margin of prosternum feebly sinuate, the median portion slightly and outwardly arcuate. Disk of prosternum convex medially, densely covered with coarse, round and setiferous punctures which are separated by one-third their diameter and sparsely intermingled with fine punctures. Prosternal process elevated, its posterior margin truncate. Lateral sides of the process and anterior margins of the procoxae marginate and carinate.
Mesosternum short and coarsely punctate, its anterior margin broadly emarginate. Intercoxal disk of metasternum sparsely and coarsely punctate, the punctures being variously separated by one to five times their diameter and with short setae. Lateral disk separated from basal one-third by post-mesocoxal stria; the basal area deeply excavate to fit mesotibia, impunctate and shining; the apical area has a punctation similar to that of the intercoxal disk, but the punctation is much sparser and coarser, except on a triangular area before latero-apical angle, and with an oblique stria apically which extends inwardly and posteriorly from the apical third of the metasternal-metepisternal suture to the apical end of the metasternal suture.
Intercoxal disk of 1st abdominal sternum sparsely covered with coarse, round and setiferous punctures which are separated by about twice their diameter and become sparser medially, and evenly intermingled with other fine punctures; anterior margin broadly and strongly carinate, the lateral end of the carina extending posteriorly and obliquely, and at apical third strongly bent outwardly. Lateral disk separated from basal two-thirds by the carina; the basal area deeply excavate to fit metatibia; the apical area impunctate and short.
Protibia narrow, its outer margin angulate medially and without denticles. Outer margins of meso- and metatibiae strongly angulate at basal third.
Specimens examined. Holotype, 1, Sayama-ko, Iruma, Saitama-ken, 5/x/1980, T. Syooda leg. collected from the nest of Pheidole ferridoa. (NA). Paratype, 1, Susono-shi, Shizuoka-ken, 29/ix/1988, M. Mori leg. collected by bait trap.
Distribution. Japan (Honsh).
Remarks. Eucuritopsis japonicus is easy to recognize by the characters given in the key and description; it cannot be confused with any other Japanese species of the family.
This species was found in the nest of an ant, Pheidole ferridoa Smith (Determined by Dr. Y. It), and also was collected by a bait trap.

6. 3. 3. Subfamily Tribalinae Bickhardt, 1917

Genus Epierus Erichson, 1834

Epierus Erichson, 1834: 158 [type species: Hister fulvicornis Fabricius, 1801: 90. Designated by Bickhardt, 1917: 123]; Marseul, 1854: 671; Schmidt, 1885: 272; Ganglbauer, 1899: 370; Jakobson, 1911: 640, 646; Bickhardt, 1917: 122; Arnett, 1962: 376, 381; Witgall, 1971: 180; Kryzhanovskij and Reichardt, 1976: 280; Vienna, 1980: 230; Mazur, 1984: 150.
Epierus lucus Lewis, 1884

Epierus lucus Lewis, 1884: 136 [Japan: Kasuga no Miya, Nara]; Bickhardt, 1917: 124; Mazur, 1984: 152.

Original description. "Ovalis, parum convexus, niger, nitidus; antennis rufis, pedibus piceis; pronoto punctulato; elytris striis 5 dorsalibus et suturali integris; propygidio pygidioque dense punctatis. L. 2.1/2 mill.
Larger and more ovate than E. comptus; the head is very finely punctured, with a transverse stria between the eyes. The thorax is clearly punctate, and rather thickly so at the base in front of the scutellum; the interstices of the elytral striae are all finely punctured.
The type of this species came from a rotten tree in the ground of Kasuga no Miya, at Nara, June 1881, and I believe all the species of this genus are of arboreal halbits. I have only one specimen."
Remarks. No specimens of the species have been available for my study.
Distribution. Japan (Honsh).

Epierus uenoi M. hhara, sp. nov.
(Fig. 50, 51)

Japanese name: Amami-ana-aki-emma-mushi.

Description. Body length, width and biometric data are as follows: PPL 1.83, PEL 1.71. Width 1.32, APW 0.44, PPW 1.18, PL 0.56, EL 1.10, EW 1.32, ProW 0.61, ProL 0.20, PyL 0.34, PTL 0.47, MSTL 0.39, MTTL 0.42. Body oval, black and shining; femora, tibiae, tarsi, antennae and mouthparts castaneous.
Frontal stria (Fig. 50B) absent anteriorly, but rudimentarily present laterally on basal third; surface of head depressed medially and elevated obliquely above antennal cavity; punctation of surface fine and evenly separated by about twice their diameters. Labrum with two long setae. Club of antenna without segmentation, its apical margin truncate.
Pronotal sides (Fig. 50A) strongly convergent to apices; marginal stria complete laterally, but entirely absent anteriorly; disk finely punctate, the punctures being separated by two to four times their diameter and becoming a little coarser laterally; basal margin of pronotum obtusely angulate at middle; broadly even medially.
Epipleural marginal stria complete, sparsely crenate, feebly carinate, and running a little distant from the margin. Narrow band between the margin and epipleural marginal stria with fine punctures in a row. Elytral marginal stria deeply impressed on apical half and sparsely crenate. Epipleura sparsely and somewhat coarsely punctate and even. Subhumeral stria absent. Oblique humeral stria finely impressed on basal fourth. First dorsal stria deeply impressed on basal two-thirds, sparsely and coarsely crenate. Second and 3rd dorsal striae present on basal three-fourths, deeply and broadly impressed, and coarsely crenate. Fourth dorsal stria a little shorter than the 3rd, lightly impressed, and sparsely crenate. Fifth dorsal stria absent or sometime rudimentarily present on medio-basal sixth. Sutural stria shortly present on median third and lightly impressed. Interstices among dorsal elytral striae sparsely clothed with fine punctures, which are separated by about four to five times their diameter; on median third and apical fifth the punctures becoming coarser and denser and being separated by two to three times their diameter except on a narrow sutural band, where the punctures are densely and finely punctate, and an extreme apical band, which is impunctate.
Propygidium (Fig. 50C) with coarse punctures which are separated by their own diameter to twice the diameter and become finer around margin. Pygidium evenly covered with coarse and round punctures which are separated by about their diameter.
Anterior margin of prosternal lobe (Fig. 50D) nearly straight medially, its marginal stria clearly impressed and carinate; apical sixth of the lobe transversely elevated; disk sparsely clothed with fine punctures. Prosternal keel feebly convex on apical half and even on basal half; carinal striae distinctly impressed, and divergent apically and basally; punctation of disk of keel similar to that of the prosternal lobe. Descending lateral striae well impressed, complete and divergent apically. Basal margin of the keel broadly emarginate.
Anterior margin of mesosternum (Fig. 50D) round, its marginal stria nearly complete laterally and broadly interrupted medially on anterior margin; disk sparsely clothed with fine punctures which are a little finer than the pronotal ones and separated by two to three times their diameter. Meso-metasternal suture strongly and sparsely crenate, carinate and arcuate anteriorly. Punctation of intercoxal disk of metasternum similar to that of the mesosternum, the punctures becoming coarser and sparser laterally. Longitudinal mid line of the disk weakly elevated on median fifth. Lateral metasternal stria extending obliquely and posteriorly on basal two-thirds. Lateral disk of metasternum covered with large and round punctures which are separated by their own diameter to half the diameter. Mesocoxal stria extending obliquely and posteriorly, its lateral end not attaining to the lateral margin of metasternum.
Intercoxal disk of 1st abdominal sternum sparsely clothed with fine punctures which are separated by four to six times their diameters; 1st abdominal stria complete on each lateral side.
Protibia (Fig. 50E, F) slender and multidentate (with 11 small denticles on outer margin).
Male genitalia as shown in Fig. 51.
Specimens examined. [Nansei Isles.] <Amami-sshima Is.> 1 ex., Tsunagu, 2/iii/1989, T. Ueno leg.
Distribution. Japan (Nansei Isles: Amami-sshima Is.)
Etymology. This species is named in honor of Mr. Teruhisa Ueno, who gave me the opportunity to study the spceimen.
Remarks. Epierus uenoi can easily be distinguished from E. lucus by the striation of the elytra, in combination with its limited distribution (Amami-sshima, Nansei Isls.).

6. 3. 4. Subfamily Onthophilinae Thomson, 1862

Key to the Japanese genus of the subfamily Onthophilinae

1(2) Dorsal surface without hair. .................... Genus Onthophilus Leach, 1817
2(1) Dorsal surface with short and stout hair. ....... Genus Epiechinus Lewis, 1891

Genus Onthophilus Leach, 1817

Onthophilus Leach, 1817: 77 [type species: Hister striatus Forster, 1771]; hhara and Nakane, 1986: 3.
Scolytus O. Mlller, 1776: 22 [nec. Geoffroy, 1762: 309], synonymized by Jakobson, 1911: 641.
Orthophilus: Westwood, 1840: 22 [misspelled], corrected by Westwood, 1840: 157.

Japanese species of this genus were already revised in hhara and Nakane (1986). Herein I add some specimens examined.

Key to the Japanese species of the genus Onthophilus.

1( 2) Pronotum with 8 costae. Body length 2.50 - 2.65 mm. ............................ .................................................................. O. silvae Lewis, 1884
2( 1) Pronotum with 4 or 6 costae.
3( 8) EC3 (elytral costa 3) ending just caudad of a deep transverse fossa situated within the front margin of the elytron.
4( 5) PC3 (pronotal costa 3) present on apical and basal half of the pronotum, and strongly developed. Body length, 3.60 - 4.68 mm. ................................ ............................................................. O. ostreatus Lewis, 1879
5( 4) PC3 absent, or present only on basal half of the pronotum.
6( 7) PC3 present. Body length, 2.50 - 3.02 mm. ....... O. niponensis Lewis, 1907
7( 6) PC3 absent. Body length, 3.28 - 3.38 mm. ......................................... ................................................. O. aonoi M. hhara et Nakane, 1986
8( 3) EC3 entire upto the front margin of elytron.
9(10) Body longer than 2.5 mm (2.86 - 3.54) mm. Elytral costae sometimes interrupted. .................................................... O. ordinarius Lewis, 1879
10( 9) Body shorter than 2.5 mm. Elytral costae clear and complete.
11(12) Propygidium with 3 costae. Body length, 1.82 - 2.34 mm. ....................... ........................................................... O. flavicornis Lewis, 1884
12(11) Propygidium with 1 costa. Body length 2 mm (after Adachi, 1930). ............ .............................................................. O. kamiyai Adachi, 1930

Onthophilus silvae Lewis, 1884

Onthophilus silvae: hhara and Nakane, 1986: 5.
Japanise name: Sinano-sesuji-emma-mushi.

Specimens examined [additional records]. [Hokkaid]] 2 exs., Yahata, Kutchan, 20, 30/viii/1993, M. hhara leg. collected near the nest of ant, Lasius (Dendrolasius) fulginosus.
[Honsh]] <Kanagawa-ken> 1 ex., Daiy-zan, Hakone, 14/viii/1982, Y. Hirano leg. (NSMT).
Distribution. Japan (Hokkaid, Honsh).

Onthophilus aonoi M. hhara et Nakane, 1986

Onthophilus aonoi hhara and Nakane, 1986: 6; Hirano, 1988: 41 [Kanagawa, Honsh]].
Japanese name: Nise-ko-sesuji-emma-mushi.

Distribution. Japan (Honsh).

Onthophilus ostreatus Lewis, 1879

Onthophilus ostreatus: hhara and Nakane, 1986: 5.
Japanese name: -sesuji-emma-mushi.

Specimens examined [additional records]. [Honsh]] <Ibaraki-ken> 1 ex., Tadobe, Sakuragawa, Niiharu-gun, 3/xi/1984, K. Haga leg. <Tkky-to> 1 ex., Nakano, 20/xi/1923, no collector's name (NSMT); 1 ex., Tkky, no data and collector's name (NSMT); 5 exs., Tkky, 25/x/1936, T. Adachi leg. (NSMT). <Kanagawa-ken> 6 exs., Mizonokuchi, no date and collector's name (NSMT). <Hyggo-ken> 1 ex., Harada, Kbbe, 19/xi/1914, no collector's name (BSM).
Distribution. Japan (Honsh, Shikoku, Kyssh); Continental China; Taiwan.

Onthophilus niponensis Lewis, 1907

Onthophilus niponensis: hhara and Nakane, 1986: 6.
Japanese name: Ko-sesuji-emma-mushi.

Specimens examined [additional records]. [Honsh]] <Fukushima-ken> 6 exs., Wakamatsu, 19/iv/1947, Y. Kurosawa leg. (NSMT). <Tkky-to> 1 ex., Komazawa, 4/vii/1931, K. Seiki leg. (NSMT). <Gifu-ken> 1 ex., Nannou, 11/iv/1981, T. Nohira leg. (refer to Takai, 1989).
Distribution. Japan (Honsh, Kyssh).

Onthophilus ordinarius Lewis, 1879

Onthophilus ordinarius: hhara and Nakane, 1986: 9.
Japanese name: Ezo-sesuji-emma-mushi.

Specimens examined [additional records]. [Hokkaid]] 10 exs., Nopporo, 5/vi/1987, M. hhara leg.; 5 exs., Ditto, 12/vi/1987, M. hhara leg.; 1 ex., Ditto, 18/vii/1987, M. hhara leg.; 2 exs., Sapporo, 10/vi/1940, Y. Nishijima leg. (NSMT).
Distribution. Japan (Hokkaid); Russia (Vladivostok, Baikal Sea, Novosibirsk, Irkutsk, Ussuri).

Onthophilus flavicornis Lewis, 1884

Onthophilus flavicornis: hhara and Nakane, 1986: 9.
Japanese name: Kinoko-sesuji-emma-mushi.

Specimens examined [additional records]. [Hokkaid]] 1 ex., Sapporo, 26/vi/1939, Y. Nishijima leg. (NSMT).
[Honsh]] <Ibaraki-ken> 1 ex., Kamisaki, Sakura-mura, 4/v/1985, K. Haga leg. <Tochigi-ken> 1 ex., Nikk, 11/v/1985, M. Kiuchi leg. <Saitama-ken> 1 ex., 20/iv/1968, H. Takizawa leg. <Tkky-to> 1 ex., Kakinokizaka, Meguro, 5/iv/1947, H. Watanabe leg. (NSMT). <Nagano-ken> 4 exs., Jigokudani, Yamanouchi, 29/iv/1983, M. Kiuchi leg.; 1 ex., Shiga-kggen, 7/v/1986, H. Tanaka leg.
Distribution. Japan (Hokkaid, Honsh, Shikoku, Kyssh).

Onthophilus kamiyai Adachi, 1930

Onthophilus kamiyai: hhara and Nakane, 1986: 9.
Japanese name: Nise-kinoko-sesuji-emma-mushi.

Distribution. Japan (Honsh).

Genus Epiechinus Lewis, 1891

Epiechinus Lewis, 1891d: 320 [type species: Onthophilus costipennis Fahreaus in Boheman, 1851: 549. Originally designated]; 1892: 232; Bickhardt, 1916-17: 66; 1921: 80; Desbordes, 1919: 408; Reichardt, 1941: 94; Kryzhanovskij and Reichardt, 1976: 290; Mazur, 1984: 146.

Epiechinus arboreus (Lewis, 1884)
(Fig. 52, 53, 54)

Onthophilus arboreus Lewis, 1884: 139 [Japan: Honsh: Nara].
Epiechinus arboreus: Lewis, 1891: 320; Hisamatsu, 1985: 226, pl. 4, f. 1.
Scolytus arboreus: Jakobson, 1911: 652.
Japanese name: Chibi-ke-sesuji-emma-mushi.

Original description. "Orbicularis, niger, hispidus, opacus; antennis clava ferruginea. O. hispido proxime affinis, sed minor. L. 1.1/2 mill."
Description. .Body oval, dark brown, furnished with minute spines and usually covered with mudlike scales. Body length, PPL, 1.58 - 1.99 mm (1.82 0.04, n=7), PEL, 1.56 - 1.91 mm (1.78 0.04, n=7). Width, 1.36 - 1.62 mm (1.51 0.03, n=7). Biometric data are given in Table 9.
Frontal lateral sides (Fig. 52D) convergent apically on basal half and strongly emarginate behind antennal sockets; anterior margin roundly arcuate. Margin of the frontal disk strongly carinate. Disk with five longitudinal costae; the median costa present on basal half; the lateral costae convergent apically through behind the antennal sockets and united with each other anteriorly; five cross costae present anteriorly between the lateral costa and the anterior marginal carina; the mediolateral costae shortly present on basal fourth. Surface of the disk sparsely furnished with minute spines.
Pronotal sides (Fig. 52A) feebly convergent forward on basal three-fourths, thence strongly convergent apically, and strongly carinate and densely furnished with minute spines. Anterior margin of pronotum broadly emarginate, the median portion nearly straight. Disk with six costae (Fig. 52E); the median four costae present on apical third and feebly elevated; the lateral costae strongly elevated, complete and interrupted at apical fourth; sides of these costae furnished with minute spines. Surface of the disk sparsely covered with coarse and round punctures which are separated by their own diameter to thrice the diameter and have minute setae in a trapezoid area on the mediobasal two-thirds; other area, except on costae, impunctate and shining.
Epipleura without stria, with a costa on margin of elytron, the costa sinuate at middle and furnished with minute spines on basal half; surface with two punctured rows, the punctures quadrate, large and shallow, the inner row complete and the outer (running close to the margin) shortly present on medio-apical fourth. Disk of elytron (Fig. 52A) with five spinal costae (excluding the costa on elytral margin); the 1st to 4th costae (naming from outside to inside) strongly elevated, complete and furnished with minute spine on their lateral sides; the sutural costa (closely present along suture) slightly elevated and furnished with minute spines only on the outer lateral side; interspace (Fig. 52F) between these costae with two punctured rows, the punctures round, large, deep and being separated by about their own diameter.
Propygidium short, depressed laterobasally and irregularly covered with coarse, round, shallow and spiral punctures, and intermingled with fine punctures among coarse ones; large, round and shallow punctures present along the anterior margin. Pygidium (Fig. 53D) irregularly covered with various sized and large punctures which are densely present medially, and other fine punctures intermingled.
Anterior margin of prosternal lobe (Fig. 52C, 53A) nearly straight; disk with a transverse punctured row on apical fifth, and sparsely with fine spines. Suture between the lobe and the keel clearly impressed. Prosternal keel feebly depressed, its carinal striae strongly elevated, convergent apically; posterior margin broadly and roundly emarginate. Lateral prosternal stria strongly carinate and divergent apically.
Anterior margin of mesosternum (Fig. 53A) sinuate, the median portion produced to fit the prosternal emargination; disk short and deeply excavate laterally (Fig. 53B); posterior margin angulate obtusely at middle; surface sparsely covered with fine spiral punctures, deeply excavate laterobasally and at apical third on lateral side, the laterobasal excavation becoming broader posteriorly and inwardly. Lateral metasternal stria carinate, extending posteriorly obliquely and angulate outwardly at basal fifth. Lateral disk with two deep excavations (Fig. 53C), the basal one present behind the mesocoxa and the apical one transverse and present on apical half; remains of the basal half covered with large round and shallow punctures.
Intercoxal disk of 1st abdominal sternum (Fig. 53C) short, with two excavations; the basal one deeply present on each lateral area, and the apical one shallowly on each latero-apical angles.
Protibia narrow, its outer margin with ten setae and angulate medially; interspace between apical 2nd and 3rd setae broad. Mesotibia with twelve setae on outer margin. Metatibia with thirteen setae on outer margin.
Male genitalia as shown in Fig. 54.
Specimens examined. [Honsh]] <Nara-ken> 1 ex., Kasuga-yama, 8/i/1954, M. ttake leg. (NA); 3 exs., Ditto, 19/vi, 7, 14/vii/1957, T. Shibata leg. (NA); 2 exs., Ditto, 13/vi/1958, T. Nakane leg. (NA); 1 ex., 18/iv/1965, M. Got leg. (NA); 8 exs., Ditto, 26/xi/1958, T. Shibata leg. (NSMT); 1 ex., Nara, 28/iii/1959, no collector's name (NSMT).
Distribution. Japan (Honsh).
Remarks. Epiechinus arboreus is easy to be distinguished from other Japanese species of the subfamily Onthophilinae by its small size and the presence of minute spines on the dorsal surface in combination with the distinctly developed elytral costae.
Little is known about the habitats of this species. Lewis (1884) noted that this species was residing in galleries of wood-borers, probably Tomicus.

Table 9. Biometric data of Epiechinus arboreus (Lewis).
------------------------------------------------------------------------------------------------------
APW 0.55-0.67 (0.620.01) 7
PPW 1.06-1.30 (1.210.03) 7
PL 0.59-0.71 (0.650.02) 7
EL 0.95-1.18 (1.100.03) 7
EW 1.36-1.62 (1.510.03) 7
ProW 0.51-0.67 (0.600.02) 7
ProL 0.16-0.26 (0.220.01) 7
PyL 0.39-0.47 (0.440.01) 7
PTL 0.47-0.63 (0.560.02) 7
MSTL 0.37-0.53 (0.490.02) 7
MTTL 0.39-0.57 (0.530.02) 7
-------------------------------------------------------------------------------------------
1 This study was supported in part by a Grant-in-Aid for Scientific Research from the Japan Ministry of Education, Science and Culture in 1990 (No. 610950221833).
2** AB=C. +A: Number of new species described; -A: no. of sp. synonymized; +B: no. of sp. newly recorded; -B: no. of sp. deleted; C: sum.


references