Title

A REVISION OF THE SUPERFAMILY HISTEROIDEA OF
JAPAN [COLEOPTERA]

Author(s)OHARA, Masahiro

Citation

Insecta matsumurana. Series entomology. New series, 51: 1283

Issue Date

1994-12URL

http://hdl.handle.net/2115/9872

RightsTypebulletin

Additional
Information

Instructions for use

Hokkaido University Collection of Scholarly and Academic Papers : HUSCAPINSECTA MATSUMURANANEW SERIES 51

DECEMBER 1994

A REVISION OF THE SUPERFAMILY HISTEROIDEAOF JAPAN [COLEOPTERA]
By MASAHIRO OHARA

Abstract
OHARA, M. 1994. A revision of the superfamily Histeroidea of Japan [Coleoptera]. Ins.
matsum. n. s. 51: 283 pp., 166 figs., 42 tabs.
The Japanese species of the superfamily Histeroidea are revised. A historical review shows that113 species of the superfamily, including 6 species newly described and 2 species newly recorded in thisrevision, are known to occur in the Japanese territory including small islands in the Pacific Ocean.
Morphology of external structures and their terms are presented. "Wenzel-Mazur System" is criticizedfrom a phylogenetic viewpoint, and another cladogram is constructed for the subfamilies of theHisteridae. The 6 new species are Hypocaccus akanensis, Platylomalus kusuii, Anap/eus nakanei,
A. nomurai, A. hagai, and Epierus uenoi. Gnathoncus communis (Marseul) and Chaetabraeus
cohaeres (Lewis) are newly recorded. Boreochlamydus ohtanii is transferred into the genus
Eucuritopsis.
Author's address. Otam Museum, Tronai 2-1-20, Otaru, 047 Japan.

Dissertation submitted to the Graduate School of Agriculture, Hokkaid6 University, in partial
fulfillment of the requirements for the degree of Doctor of Agriculture, 1994 [abridged for publication).
This study was supported in part by a Grant-in-Aid for Scientific Research from the Japan Ministry of
Education, Science and Culture, 1990 (No. 610950221833).

Contents
Introduction ..................................................................................................... 5
Methods ........................................................................................................... 5
Historical Review .............................................................................................. 9

1.
2.

3.
4.

5.
6.
6.1.

6.2.
6.3.
6.3.1.

6.3.2.
6.3.3.
6.3.4.

6.3.5.
6.3.6.

6.3.7.
6.3.8.

6.3.9.
6.3.10.

7.
8.

Morphology and Tenns ..................................................................................... 13
Phylogeny............ ..... .... ..... ......... ......... ......... ..... ..... ......... ............. ..... ..... ...... 53
Systematics .................................................................................................... 58
Family Sphaeritidae .......................................................................................... 58
Family Synteliidae ............................... ........................................................... 61
Family Histeridae ............................................................................................. 65
Subfamily Niponiinae ...................................................................................... 66
Subfamily Chlamydopsinae ................................................................................ 78
Subfamily Tribalinae ........................................................................................ 80
Subfamily Onthophilinae .................................................................................. 83
Subfamily Histerinae ......................................................................................... 89
Subfamily Hetaeriinae ...................................................................................... 143
Subfamily Dendrophilinae ................................................................................ 148
Subfamily Abraeinae ...................................................................................... 204
Subfamily Saprininae ...................................................................................... 214
Subfamily Trypeticinae .................................................................................... 267
Acknowledgments .......................................................................................... 272
References .................................................................................................... 273

Enumeration of genera and species
Sphaeritidae ......................................... 58
Sphaerites .... ...... ......... ..... ....... 58
poLitus Mannerheim .............. 59
Synteliidae .......................................... 61
SynteLia ................................. 61

Tribalinae ........................................ 80
Epierus ................................... 80

histeroides Lewis ................. 61
Histeridae ............................................ 65
Niponiinae ....................................... 66

OnthophiLus ........ .... ....... ......... 83
siLvae Lewis.................. ...... 83

Lucus Lewis............... .......... 80
uenoi M. Ohara, sp. nov ........ 80
Onthophilinae .................................. 83

aonoi M. Ohara et Nakane ...... 83

Niponius ...... .......... ................ 66

ostreatus Lewis .................... 84

furcatus Lewis..................... 66
impressicollis Lewis............. 68

niponensis Lewis ................. 84
ordinarius Lewis................... 84

itoi ChGj6 ........................... 71

flavicornis Lewis. ....... .......... 84

obtusiceps Lewis .................. 72

kamiyai Adachi .................... 85

osorioceps Lewis ........ .......... 74

Epiechinus ........ ...... ... ............. 85

Chlamydopsinae ............................... 78
Eucuritopsis ............................ 78
ohtanii (K. Sawada) ............... 78

arboreus (Lewis) ................... 85
Histerinae ........................................ 89
Exosternini .................................. 902

Binhister ...... .... ........... ... ......... 90
chujoi Cooman ..... ...... ......... 90

Atholus ................. ......... ....... 136
bimaculata (Linnaeus) .......... 137
coelestis (Marseul) .............. 137

Notodoma ........ ....................... 90
fungo rum Lewis................ ... 90

depistor (Marseul) ....... ........ 137

Hololeptini ................................... 91
Hololepta ................................ 91(Hololepta)
amurensis Rei tter ................. 91
depressa Lewis ..................... 91
higoniae Lewis .................... 91
Platysomatini ............................... 92
Platysoma .......... ..... ....... ...... ... 92(Platysoma)
celatum Lewis........ ............. 93
deplanatum (GyUenhal) .......... 93
lewisi Marseul ........ ... .......... 93
rasile Lewis............... .......... 94
takehikoi M. Ohara ............... 94
tsushimae M. Ohara .............. 94
vagans Lewis ......... .............. 94
unicum Bickhardt .................. 94(Platylister)
atratllnt (Erichson) ......... ....... 96
cambodjense (Marseui) ........... 99
horni (Bickhardt) .... .
101oberthuri (Cooman) .,. ... .... ... 103
pini (Lewis) ....................... 103(Cylister)
lineicollis (Marseul) ............ 105
elongatus (Thunberg) ** ....... 109(Apobletes)
schaumei Marseul ...
109
Eblisia ................................. 112
satZUlnae (Lewis) ................. 112
Histerini .................................... 112
Hister ................................... 112

duodecimstriatus
quatuordecimstriatus (Gyllenhal)............................ 137
pirithous (Marseul) .............. 138
Margarinotus .......................... 138(Kurilister)
karbatovi (Tishechkin) ......... 139(Ptomister)boleti groupboleti (Lewis) .................... 139
weymarni groupcadavericola (Bickhardt) ........ 139
agnatus (Lewis) .. ................ 140
weymarni Wenzel ................ 140
reichardt Kryzhanovskij ........ 141
striola striola (SaWberg) ....... 141
yezoellsis M. Ohara. ..... ... .... 141
sutus groupmarginepunctatus (Lewis) ..... 141
sutus (Lewis) ..................... 142(Grammostethus)
Ilipollicus (Lewis) ............... 142
Hetaeriinae ..................................... 143
Hetaerius ............................... 143
gratus Lewis ...................... 144
optatus Lewis ..................... 146

Dendrophilinae ................................ 148
Dendrophilini ............................. 149
Delldrophilus .......................... 149
xavieri Marseul ................... 149

Anapleini ................................... 152
Anapleus .... ...... ..... ......... .... ... 152

impunctatus Osawa ............. 114
japonicus Marseul ............... 114
congener Schmidt ... ............ 118

semen (Lewis) .................... 153
Ilakanei M. Ohara, sp. nov .... 159
Ilomurai M. Ohara, sp. nov ... 161

simplicisternus Lewis.... ...... 123

hagai M. Ohara, sp. nov ....... 162

unicolor leonhardi Bickhardt .. 126
concolor Lewis ................... 127

Bacaniini .................................... 166
Bacallius ................................ 166

Zabromorphus ............ ....... ..... 132(Mullerister)pUllctulatus (Wiedemann) ...... 132

Ilipollicus Lewis ................. 166

Merohister .. ...... ... ...... ....... ..... 135(Bacallius)

jekeli (Marseul) .. ... ............. 136
aino (Lewis) ....................... 136
uelloi M. Ohara... ..... ...... .... 136

mikado (Lewis) ................... 166

3

(Pycnacritus)

Paromalini .................................. 167
Carcinops .............................. 167

homoeopathicus Wollaston... 214
Saprininae ...................................... 214
Gnathoncus ............................ 215
nannetensis (Marseu!) .......... 215(Carcinops)
pumilio (Erichson) ......... ..... 167
Australomalus ........................ 171
montivagus (Lewis) ............. 171
Platylomalus ........ ..... ..... ....... 174
fujisanus (Lewis) ................ 174
mendicus (Lewis) ................ 177
niponensis (Lewis) . ... ...... .... 180
persimilis (Lewis) ......... ...... 184
viaticus (Lewis) .. ........ ........ 187
kusuii M. Ohara, sp. nov ..... 190
Pachylomalus ........................ 193

rotundatus (Kuge!ann) .......... 220
communis (Marseu!)* .......... 223
Saprinus ................................ 226(Sprinus)
cyaneus auricollis Marseu!.... 226
niponicus Dahlgren............. 229
pecuinus Marseu! .............. 232
planiusculus Motschu!sky .... 232
splendens (Paykull) ............. 236
Hypocacculus ......................... 240(Nessus)asticus (Lewis) ................... 240
Hypocaccus ............................ 243(Hypocaccus)
ainu (Lewis) ....................... 244
axeli Kryzhanovskij ............. 245
lewisii (Schmidt) ................ 248
sinae (Marseu!) ................... 250
subaenus (Schmidt) ........ ..... 252
akanensis M. Ohara, sp. nov.
............................ 255(Baeckmanniolus)(Canidius)
musculus (Marseu!) ............. 193
Eulomalus ............................. 196
lombokanus Cooman ........... 197
tardipes (Lewis) .................. 197
Paromalus ............................. 197(Paromalus)omineus Lewis ................... 197
parallelepipedus (Herbst) ....... 199
vernalis Lewis .................... 201
Abraeinae ....................................... 204
Abraeini ..................................... 204
Chaetabraeus .......................... 204
bonzicus (Marseu!) .............. 204
cohaeres (Lewis)* ................ 208
sp ...................................... 209
P!egaderini .................................. 211
Plegaderus ............................. 211(Plegaderus)
marseuli Reitter .................. 211
shikokensis Hisamatsu ......... 213
Acritini ...................................... 213
Acritus .................................. 214(Acritus)

varians varians (ScJnnidt).... 258
varians hatsune (Nakane) ...... 262
Eopachylopus ......................... 262
ripae (Lewis) .... .................. 264
Trypeticinae .................................... 267
Trypeticus ........ ..................... 267
fagi (Lewis) ....................... 267
vena tor (Lewis) ................... 272* New!y recorded from Japan.
** Introduced to Japan.

komai Lewis ...................... 214

4

1. INlRODUCTION

The superfamily Histeroidea is composed of 3 families, Sphaeritidae, Synteliidae
and Histeridae. The Sphaeritidae and the Synteliidae are smaIl 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 theworld. 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 hi sterid beetles have been
well known as predacious cadavericolous insects controlling populations of dipterous
flies and other insects (Summerlin et aI., 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 (Ohara andNakane, 1986, 1989; Ohara, 1986, 1989, 1991a, b, 1992a, b, c, 1993a). The present
review is therefore the first comprehensive study of the superfamily in Japan; 113
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 wereborrowed 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 Europe, and 100 from the rest of theworld. I tried to examine the type specimens of some Japanese species, especially ofproblematic species, and have succeeded to borrow the types of the following species:
Plegaderus shikokensis, Rypocaccus varians hatsune, Anapleus semen, A. japonicus,
and Rister 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
microscopes Olympus SO (magnification: up to x160) and Olympus SZH-131(magnification: up to x128). Some structures were also observed in a SEM (Hitachi S2000A).
Genitalia were removed from dried specimens, and then 1) heated in 10 % KOH
at 60°C for about 1 hour or more according to the size; 2) washed and dissected in 70%
ethyl alcohol (remaining muscles were removed), 3) transferred into lactic acidcontaining acid fuchsin and heated at 60°C for 3 hours, 4) transferred into a mixture ofglacial acetic acid 1 part and methyl salicylate 1 part and left there for 30 minutes, and5) 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 width5

(Fig. 2). Measurements of some body parts, with mean, ranges, standard error andsample 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:
PEL:
HOW:

length between anterior angles of pronotum and apex of pygidium.
length between anterior angles of pronotum and apices of elytra.
width between horns (projections) of epistoma.

B

~.
o

.

"

!I

~® t9

o

'c
\

~IVA SHIKOKU_.

. ",?---V. KYUSHU
~o

(/

c
",I

,

oj

1

',il

og

VI. NANSEI ISLES.

o·Fig. 1. A: Japan and adjacent regions. B: Japanese mainland and some associated islands. C: Nansei
Islands. For symbols see Table 1.

6

Table 1. Names of islands and regions in Japan under investigation. See Fig. 1.
No.

Main islands(put in brackets [ 1in Specimens examined).

I
Chishima Isles
II-I
Hokkaido
11-2
11-3
11-4
II-S
III-l-a Honshu
III -lob
III -l-c
III-l-dIII-l-e
III-l-f
III-l-g
III -l-h
III-l-iIII-l-jIII-l-k
III-I-1
III-l-m
III-l-n
III-l-oIII-l-p
III-l-q
III-l-r
III-l-s
III-l-t
III-l-u
III -l-v
I1I-l-w
I1I-l-x
I1I-I-y
III -l-z
lIl-l-aa
I1I-l-bb
III-l-cc
I1I-l-dd
III-l-ee
I1I-l-ff
III-l-gg
I1I-l-hh
I1I-2
1II-3

IlIA
III-S
IV-l-a Shikoku
IV-l-bIV-l-cIV-l-dV-I-a Kyushu
V-lob
V-l-c
V-I-d
V-l-e
V-l-f
V-log
V-2
V-3
V-4

Main islands are divided into
following islands or prefectures.

Etorofu Is., Kunashiri Is., Shikotan Is., Habomai Isles
properRishiri Is., Rebun Is. (located close to Wakkanai)
Teuri Is., Yagishiri Is. (located close to Haboro)
Okushiri Is. (located close to Setana)
Oshima Isles (located close to Matsumae)... _._ .. _.. __ .. _._._ .. _.. _.
Aomori-ken
Iwate-ken
Tohoku district
Miyagi-ken
Akita-ken
Yamagata-ken
Fukushima-ken
Ibaraki-ken
Tochigi-ken
Gumma-ken
Kanto district
Saitama-ken
Chiba-ken
Tokyo-toKanagawa-ken
Niigata-ken
Toyama-ken
Ishikawa-ken
Fukui-ken
Chubu district
Yamanashi-ken
Nagano-ken
Gifu-ken
Shizuoka-ken
Aichi-ken
Mie-ken
Shiga-ken
Kyoto-fu
Kinki district
6saka-fu
Hyogo-kenNara-ken
Wakayama-ken
Tottori-ken
Shimane-ken
Chugoku district
Okayama-ken
Hiroshima-ken
Yamaguchi-ken
Izu Isles
Sado Is., Awa-jima Is.
Hegura Is.
Oki Isles
Tokushima-ken
Kagawa-ken
Ehime-ken
Kochi-ken
Fukuoka-ken
Saga-ken
Nagasaki-ken
Kumamoto-ken
6ita-kenMiyazaki-ken
Kagoshima-ken
Tsushima Is.
Iki Is.
Goto Isles

7

Table 1. Continued.
Amakusa Isles
Koshiki Isles
Yaku-shima Is.
Tane-ga-shima Is.
Tokara Isles
Amami-Oshima Is.
Tokuno-shima Is.
Okinoerabu Is.
Yoron Is.
Okinawa-Hont6 Is.
Kume-jima Is.
Kerama Isles
Miyako Is.
Tarama Is.
Ishigaki Is.
Iriomote Is.
Hateruma Is.
Yonaguni Is.
Haha-jima Is., Chichi-jima Is.

V-5
V-6
VI-l-a Nansei Isles
VI-l-b
VI-I-cVI-l-dVI-l-eVI-l-fVI-l-gVI-l-hVI-l-iVI-l-jVI-l-k
VI-I-I
VI-I-m
VI-l-nVI-l-oVI-l-pVII
Ogasawara Isles

PPW
APW

PPL

EW
Fig. 2. Adult of the Histeridae in dorsal Oeft) and lateral (right) views, showing parts for biometricmeasurements.
HW:
APW:
PPW:
PL:
EL:
EW:
ProW:
ProL:
PyL:
P1L:
MSTL:
MTTL:

width of head.
width between anterior angles of pronotum.
width between posterior angles of pronotum.
length of pronotum along mid line.
length of e1ytron along sutural line.
maximal width between outer margins of e1ytra.
maximal width of propygidium.
length of propygidium.
length of pygidium.
length of protibia.
length of meso tibia.
length of metatibia.

Museum acronyms. Specimens were borrowed from the following institutions.
8

The people responsible for the loans are given after the names of the institutions.
Specimens without acronyms are in my collction, now deposited in EIHU.
BSM:
EIHU:
ELKU:

HFFP:
HOR:
IJ:
NA:
NHM:
NSMT:
TPM:

Bishop Museum, Honolulu; G. A. Samuelson.
Entomological Institute (now Laboratory of Systematic Entomology), Hokkaido
University, Sapporo; M. Suwa.
Entomological Laboratory, Kyushfi University, Fukuoka; O. Tadauchi.
Hokkaido Research Center, Forestry and Forest Products Research Institute; K.
MaetO.
Collection of Mr. S. Hori, Sapporo, Hokkaido.
Collection of Mr. K. Ijima, Shibecha, Hokkaido.
Collection of Dr. T. Nakane, Chiba.
Natural History Museum, London; E. De Boise.
Natural Science Museum, Tokyo; S.-I. Ueno and A. Shinohara.
Tochigi Prefectural Museum, Utsunomiya, Honshu; H. Higuchi and K. Satoh.

Survey areas. The present study covers the islands and regions shown in Table 1
and Figure 1. The Japanese suffixes -ken, -to, and -fu are applied to prefectures.
3. HISTORICAL REVIEWCLASSIFICATION OF THE FAMILY HISTERIDAE

In the following lines main 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.
1811.
1817.
1834.

1854.

1857.

LINNAEUS described the genus Hister for 6 new species.
PA YKULL proposed a new genus, Hololepta. He first classified the histerids into 2 genera,
Hister and Hololepta.
LEACH added 4 genera, Platysoma, Dendrophilus, Abraeus and Onthophilus.
ERlCHSON divided the family into 3 major groups, and 2 of them into 5 subgroups, as
follows:
I:
Hololepta, Phylloma, OxysternusII:A.
a) Plaesius, Placodes, Platysoma, Omalodes, Cypturusb) Hister, Hetaerius, Epierus, Tribarus
B.
Dendrophilus, Paromalus
Ill:A.
Saprinus, Pacylopus, Trypaneus
B.
Teretrius, Plaegacierus, Onthophilus, Abraeinae
He used the following character states in the classification: 1. head retractile or not (II,III) (I); 2. prostemallobe present or absent (II) (III); 3. club of antenna in repose is situated onanterior margin or on middle of surface (IIA) (IIB) of prostemum; 4. articulation of club ofantenna is distinct on both ventral and dorsal sides or not distinct on one or both sides (a) (b);5. antennae articulate on margin of front of head or on frontal disk (IlIA) (IIIB).
LACORDAlRE recognized 2 tribes and 2 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.
prostemum with prostemallobe or without it (i) (ii).
MARSEUL proposed a new classification of the family.
1:
HololeptiensIV:H€reriens
V:
SapriniensII:
Trypaneens
III:
Histeriens
VI:
Abreens
His classification consisted of 6 tribes. He used the following character states to divide

9the family into the tribes: 1. head retractile or not (I, II) (III, IV, V, VI); 2. mandible long or
short (I) (II); 3. prostemal lobe present or not (III, IV) (V, VI); 4. articulation of club ofantenna is distinct or not, and shape of the club is oval or cylindrical (III) (IV); 5. antennal
fossette situated on disk or margin of prostemum, and antennal furrow of head present under
the lateral margin of head or in an emargination on the anterior margin (V) (VI).
1899. GANGLBAUER almost agreed with Marseul (1857), but he did not distinguish between the
I-Iisterinae and the Hetaerinae and the latter was included in the former. He provided 4 tribes for
European histerids, and did not mention the tribe Trypanaeini.
1909. REITTER divided the family into 7 tribes.
Hetaerinini
I:
Hololeptini
V:
II:
Histerini
VI:
Saprini
Abraeini
Ill:
Paromalini
VII:IV:
Dendrophilini
1916-17. BICKHARD's classification:
1:
Hololeptinae
VIII:
Histerinae
1. T ri balini
II:
Trypanaeinae
II:
Trypeticinae
2. Platysomini
3. HisteriniIV:
Teretriinae
4. Exosternini
V:
Abareinae
VI:
IX:
Hetaeriinae
SaprininaeVII:
1. Hetaeriomorphini
Dendrophilinae
2. Hetaeriini
3. Cblamydopsini1941. REICHARDT's classification was as follows:
Saprininae
I:
Niponiinae
V:
Dendrophilinae
II:
Trypeticinae
VI:
Hololeptinae"II";
Trypanaeinae
VII:
III:
Teretriinae"VII":Histerinae
IV:
Abraeinae
X:
Hetaerinae
In this work the Trypanaeinae and the succeeding subfmilies were erroneously
numbered. He provided a key to the 10 subfamilies, which is quite artificial and does notrellect 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 aprimitive 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 regarded 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 Trypanaeinae. The 1st genus is allied to the Acritus-group;
Hololepta is rclated 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 betweenthem); the Trypanaeinae are close to the Trypeticinae. 3) The Saprininae are a 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 2 divisions, "Saprinomorphae" and "Histeromorphae", andchanged the ranks of some taxa as follows: 1) The Hololeptinae (sensu Reichardt, 1941) werereduced to a tribe in the Histerinae. 2) The Chlarnydopsinae were treated as a subfamily andplaced next to the Abraeinae. 3) The Teretriinae (sensu Reichardt, 1941) were ranked as a tribein the Abreinae. 4) The tribe Tribalini included in the Histerinae (Bickbardt, 1916) was risen inrank to the subfaJnily 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
II:
I-listeromorphae
I:
Cblamydopsinae
VI I:
Dendrophilinae
II:
Abraeinae
VIII:
Tribalinae
I:
Abraeini
IX:
Histerinae
101976.
1980.
1982.
1984.

1989.

1854.
1860.
1873.

1877.
1878
1879.
1882.
1884.

2:
PJegaderini
1:Histerini
3:
Acritomorphini
2:
Omalodini
4:
Acritini
3:
Platysomini
5:
Teretriini
4:
Hololeptini
III:
Niponiinae
5:
Exosternini
IV:
Trypanaeinae
X:
Hetaeriinae
V:
Trypeticinae
1:
Hetaeriomorphini
VI:
Sapriniae
2:
Hetaeriini
KRYZHANOVSKIJ and REICHARDT proposed a new taxon, the tribe Bacaniini.
VIENNA elevated the Onthophilina (Thomson, 1862) to the subfamily rank.
OLEXA proposed a new tribe, the Anapleini.
MAZUR mainly followed Wenzel. See also under Section 5 (phylogeny).
I:
Saprinimorphae
II:
Histeromorphae
I:
AbraeinaeVII:
Dendrophilinae
1.
Abraeini
1.
Dendrophilini
2.
PJegaderini
2.
Anapleini3.
Acritomorphini
3.
Bacaniini
4.
Acritini
4.
Paromalini
5.
Teretriini
VIII:
OnthophilinaeII:
Niponiinae
IX:
Tribalinae
Ill:Trypanaeinae
X:
Histerinae
1.
HisteriniIV:
Trypeticinae
V:
Saprininae
2.
Omalodini
VI:
Chlamydopsinae
3.
Platysomatini
4.
Hololeptini
5.
Exosternini
XI:
Hetaeriinae
HELAV A. Tribes in the Hetaerinae were not discriminated.
JAPANESE HIS1EROIDEAMARSEUL described a new species, Hister japonicus. ......................................
+1+0=1 *MOTSCHULSKY described a new species, Hister japan us, which was later synonymized under
Margarinotus striola. . ................................................................................... + 1+0=2
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.
H.
14-striatus(=Atholus
duodecimstriatus
jekeli(=Merohisterjekell),quatuordecimstriatus), H. punctulatus, H. cadaverinus (= Margarinotus weymari or one of
its allied species), H. navus (= Margarinotus niponicus), Carcinops pumilio, Saprinus
speciosus (= S. splendens), S. pecuinus, S. nitidulus (= S. planiusculus), S. sinae andSaprinus (Gnathoncus) rotundatus. . ............................................................. +7+11=20
REITTER described a new species, Plegaderus marseuli . ..................................... +1+0=21
HAROLD recorded a species, Onthophilus striatus . ............................................. +0+ 1=22
LEWIS described 4 new species, Hister marginepunctatus, Hister simplicisternus, Bacaniusniponicus, and Acritus komai. .. .................................................................. +4+0=26
LEWIS erected a new genus for a new species, Syntelia histeroides . ....................... +1+0=27
LEWIS described 21 new species, Hololepta depressa, H. parallela (nec Sturm, 1868),
Platysoma pini, P. vagans, P. rasile, P. celatum, Hister aino (=Merohister 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* ±A±B=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.

11and Onthophilus ostreatus. Onthophilus flavicornis is, however, the same with O.
striatus in Harold (1878). ....................................................................
+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 the new genus Niponius for 4 new species, impressicollis, furcatus,
obtusiceps, and osorioceps. ........................................................................ +4+0=55
1890. SCHMIDT described Saprinus lewisii, S. subaenus, and S. varians. Later, they weretransferred to Hypocaccus . ........................................................................... +3+0=58
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=69
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=69
1899. LEWIS described 3 new species, Platysoma satzumae, Hypocaccus ainu, and Paromalus
niponensis, the last for the misidentified P. complanatus in Lewis (1892), and synonymized
Hister japanus under H. striola. .................................................................... +2+0=71
1904. LEWIS transferred Hister congener into the new genus Pachylister.
1905. LEWIS transferred Tryponaeus vena tor andfagi into the correct genus Trypeticus.
1906. LEWIS recorded a species, Platysoma deplanatum. ............................................. +0+ 1=72
1906b. LEWIS described a new species, Platylister niponensis. ...................................... +1+0=73
1907. LEWIS transferred Paromalus parallelepipedus, P. omineus, and P. vernalis into the new
genus Microlomalrus.
1907c. LEWIS described a new species, Onthophilus niponensis . .................................... +1+0=74
1911. LEWIS described a new species, Hypocaccus asticus . .......................................... +1+0=75
1914. LEWIS described a new species, Hister togoii . ................................................... + 1+0=76
1918. BICKHARDT treated Hister aino as a synonym of jekeli. .................................... +0-1=75
1920. BICKHARDT trt:ated Hister togoii as a synonym of H. simplicisternus. ............ ..... +0-1=74
1920b. BICKHARDT treated Platysoma niponensis as a synonym of P. cambodjense.
1926. REICHARDT transferred Pachylopus ripae into the new genus Eopachylopus.
1930. ADACHI described a new species, Onthophilus kamiyai . ..................................... +1+0=75
1941. COOMAN described a new species, Binhister chujoi. .......................................... + 1+0=76
1944. WENZEL described a new species, Margarinotus weymami, 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=76
1948. COOMAN described a new species, Platysoma oberthuri, and a new genus Platylomalus,
and transferred Paromalus mendicus to it.
.................................................... +1+0=77
1952. GSA WA described a new species, Hister impunctatus and a new form, rujofasciatus, of
Eopachylopus ripae. .................................................................................. +1+0=78
1955. CHOlO described a new species, Niponius itoi . ................................................. + 1+0=79
1961. NAKANE described a new species, Anapleus japonicus . ....................................... +1+0=80
1962. DAHLGREN described a new species, Saprinus niponicus . ................................... + 1+0=81
1962. ADACHI and OHNO newly recorded a species, Spaherites politus. ........................ +0+1=82
1963. NAKANE newly recorded 2 species, Peranus bimaculatus and Platysoma unicum . .............. .
. ............................................................................................................... +0+2=84
1964. NAKANE transferred Triballus semen into the correct genus Anapleus.
1976. KRYZHANOVSKIl and REICHARDT described a new species, Hypocaccus axeli, transferred
Paromalus niponensis, P. jujisanus, and P. viaticus into the correct genus Platylomalus,
and newly recorded Hister unicolor opimus (= leonhardi). ............. ................. ..... +1+1=86
1977. NAKANE described a new subspecies, Hypocaccus varians hatsune, and newly
recorded Saprinus auricollis (=S. cyaneus auricollis) from Bonin Is. .................
+0+ 1=87
1981. MAZUR described a new genus Australomalus, and transferred Platylomalus montivagus toit.
121984.

MAZUR transferred Paromalus persimilis into the correct genus Platylomalus,
and Paromalus tardipes into Eulomalus. Anapleus japonicus is synonymized under A.
sem.en. ... ................................ ... ... ...... ..... ..... ........ ...... ...... .................. ..... +0-1=86
1984. HI SA MATSU and KUSUI newly recorded Atholus coelestis. ...... .......... ............. .... +0+ 1=87
1984. HISAMATSU newly recorded Platylomalus persimilis. .....................................
+0+1=88
1984. KUSUI recorded Cylister elongatus. Because this species is undoubtedly an introduced species,
it is not counted as a member of the fauna.
1985. HISAMATSU described a new species, Plegaderus shikokensis, and newly recorded Apobletes
tener. ........................................................................................................ +1+1=90
1985b. HISAMATSU newly recorded Gnathoncus nannetensis, Eulomalus lombokanus, Platylister
alralus and P. homi. .................... ........ ...... .. ........ ............ ........ ...... .. ......... +0+4=94
1986. OHARA revised the genus Platysoma from Japan, and added 2 new species, takehikoi andtsushimae . ................................................................................................ +2+0=96
1986. OHARA and NAKANE revised the genus Onlhophilus 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. OHARA and NAKANE redescribed 2 species of the tribe Exosternini, Notodoma fungorum andBinhister chujoi.
1989. OHARA reviewed the genus Margarinotus from Japan, described a new species, M.
(Ptomister) yezoensis, and newly recorded 2 species, cadavericola and reichardti. M. (P.)
marginepunctatus was transferred from subgenus Promethister to subgenus Ptomister. .......
............................................................................................................... +1+2=101
1991. OHARA revised the genus Hololepta from Japan.
1992. OHARA revised the genus Merohister from Japan, revitalized aino, and added a new species,
uenoi . ..................................................................................................... +1+1=103
1992c. OHARA revised the genus Atholus from Japan.
1993. OHARA recorded a species, Margarinotus (Kurilister) karhatovi. ....................... +0+ 1=104
1994. SAWADA described a new genus, Boreochlamydus, for a new species, ohtanii. ..... +1+0=105
1994. In this paper, I describe 6 new species, Hypocaccus akanensis, Anapleus nakanei, A.
nomurai, A. hagai, Pia tyloma Ius kusuii, Epierus uenoi, and newly record from Japan
Chaetabraeus cohaeres and Gnathoncus communis. I transfer Boreochlamydus ohtanii tothe genus Eucritopsis . ................................................................................ +6+2=113
In conclusion, 113 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 understudy. In the Histeroidea, many papers in English published after 1950 follow the styleand descriptive terms in Wenzel and Dybas (1941). The present paper also mainly
follows Wenzel and Dybas's system of terms and Naomi (1987 - 1990). Most of theterms used are illustrated in Figure 3.
HEAD

No detailed comparative morphological study of the head has been made formembers of the superfamily Histeroidea. Because in dried specimen the head is usually
retracted in the prothorax, many past authors, except Marseul (1853 - 1862), did notdescribe the details of the head sutures and mouthparts, which are observable on theventral side.
In this section the head sutures and various parts of the cranium are dealt with.

13

Situation and general appearance of head
Head is generally prognathous in the Coleoptera. In the families Sphaeritidae andSynteliidae and the subfamily Niponiinae of the Histeridae, the head is prognathous asin many other Coleoptera. But in other Histeridae the head is almost deflexed except inseveral members (e. g., Trypanaeinae and Hololepta). Usually the posterior 112 of thehead retracts into the anterior margin of the pronotum. The prognathous state in theseveral 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 1/2 of thehead is generally narrowed to form the frontoclypeal region.
Head is usually narrower than the prothorax except in the Synteliidae andNiponius, in which the head is as broad as the prothorax.

Marginal pronotal~

Outer lateral pronotal ~Inner lateral pronotal
Oblique humeral

dorsals
3rd
Su t ur a 1--I'ttII-t-i'--lH-,

IH-HtH---4thlitr-_!

Marg. epipleural--t-tr~iMarg. elylral _-+:::::17\11

5th

~

A

External subhumaralInternal subhumeral

Lobe
Suture betw. prost. keel and lobe

Prosternal keel
CarinalPost. margo profemoral - - c = Post. mesocoxal _ _ _~~J~~t3~tr--- Marg. mesosterna I

Lat.metasternal - - f - t - W = - I

Metasternal-meseplmeral sut.

Lateral disk

Meso-metasternal sut.

Oblique metasternal

Metasternal-meteplsternal sut.

Metasternal longitudinal sut.
I"tercoxal disk

Intercoxal disk of 1 st abdominal sternum

Fig. 3. A histerid beetle, Margarinotus, showing tenns of some striae and parts. A: Dorsal view. B:
Lateral view, C: Ventral view (after Ohara, 1989).
14

Frontoclypeal suture (clypeofrontal suture in Cook, 1943): This suture runs
arcuately between the anterior tentorial pits in Coleoptera, but the pits are obscure in theHisteroidea. The suture is present only in the Sphaeritidae; in Shpaerites the suture isvery 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 theventral side and terminates on each side anteriorly to the posterior articulation of themandible; 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 ashort one along the posterior margin of each eye (Fig. 60).
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 2 sutures, because the pit is absent in major groups and obscure in Spaherites andSyntelia (Fig. 40, 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 evencompletely to form a straight line. See also under Gula plate.
The postoccipital suture between the postocciput and postgena is variable in theHisteroidea, 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 usuallywell developed and oblong, the posterior margin is often inwardly arcuate. They areusually kidney-shaped or round in lateral view.
The eyes are, however, lost in a cave-inhabiting species, Geocollus caecus
Wenzel.
The ocelli 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 nearlysquare, 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 isshort 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 andcorrugated (Fig. 42A, 43A, 45A, 46A). Hornlike structure is developed in somespecies of the subfamily Tribalinae. The majority of the Onthophilinae have costae.
Frontal and supraorbital striae on the disk of epicanium are useful characters forclassifying the Histeroidea. The frontal stria usually appears in the Saprininae,
Trypeticinae, Oendrophilinae, 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 a15

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 areusually united with each other.

A

Midcarinal suture

F

Fig. 4. Head. A, B: Schematic. C, D, E: Sphaerites politus (Sphaeritidae). F, G, H: Syntelia
histeroides (Synteliidae). I, J, K: Dendrophilus xavieri. A, C, F, I: dorsal view. B, D, G, J:
ventral view. E, H, K: lateral view, left side.
16

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

Vertexr------*d-Gular plate~_ _~ ~____ Occiput

Postocciput

J

L

Fig. 5. Head. A, B: Schematic. C, D: Pachylomalus sp. (paromalini, Dendrophilinae, Histeridae). E,
F: Trypeticus fagi (Trypeticinae, Histeridae). G, H, I: Saprinus splendens (Saprininae,
Histeridae). J, K, L: Chaetabraeus bonzicus (Abraeinae, Histeridae). A, C, E, G, J: dorsal view.
B, D, F, H, K: ventral view. I, L: lateral view, left side.
17

Ventral plates conslstzng 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 is weakly convex or flat,
usually with small or large punctures. However, a modified structure, known asantennal furrow, is found in major groups.

Occipital suture~•••••••••••f<• • • • • • • • •:lFFig. 6. Head. A, B, C: Onthophilus ostreatus (Onthophilinae, Histeridae). D, E: Epierus comptus(Tribarinae, Histeridae). F, G, H: Merohister jekeli (Histerini, Histerinae, Histeridae). I, J, K:
Hololepta amurensis (Hololeptini, Histerinae, Histeridae). A, D, F, I: dorsal view. B, E, G, J:
ventral view. C, H, K: lateral view, left side.
18

Antennal furrows on the ventral plates are developed in the following groups:
Syntelia, Pachylomalus, Onthophilus, Hololepta (Fig. 4G, SO, 6B, J) and Niponius(Fig. 42B, 43B, 4SB, 46B); in the last taxon they are deeply foveate.
Gular plate: Gula is a median ventral plate of the head behind the posteriortentorial 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 eachother, 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, andform 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. 40, G, SO). The 2nd is present in the
Oendrophilinae and Saprininae (Fig. 4J, SH). The 3rd is found in the Trypeticinae,
Onthophilinae and Abraeinae (Fig. SF, L, 6B). The last type is the most derived stateand is found in the Tribalinae and Histerinae (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. It is situated behind occiput and separated from the latterby 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, andis 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 gularsuture, and is moderately narrow (Fig. SH). In the Histerinae, the postocciput isenlarged on basal 112 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, pedicel and flagellum. The baseof the antenna is attached to the ventral side of the anterolateral margin of the forehead
near the eye. Segments are numbered 11, except in Trypeticus... of which the antenna is
lO-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, butsometimes it is oval and broader in the Saprininae and Trypecitinae (Fig. 7H, 8A).
Aagellum is composed of the 3rd - lIth segments of the antenna. In Trypeticus, it is
the 3rd - 10th. In the Histeroidea, the apical 3 (usually 9th - lith) segments arecomprised in a "club" and the rest are called "funicle". The club is usually oval,
condensed and more or less densely covered with hairs; rarely it is cylindrical in the
Hetaeriinae. In the Saprininae, the club is with "Reichardt's organ" (De Marzo andVienna, 1982a, b) on the ventral surface, which is a particular sense apparatus (Fig. 71,
SA). The articulation of the club is variable from a complete to an obscure one. The
apical most segment of the funicle is usually expanded laterally to form a fringe.

19

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 incomplete. Here, I describe theirmorphologies based only on 2 species, Notodoma fungorum and Hololepta
amurensis.

Reichardt's organ... ~~~
Fig. 7. Antennae. A: Sphaerites politus (Spbaeritidae). B: Syntelia histeroides (Synteliidae). C:
Onthophilus ostreatus (Onthophilinae, Histeridae). D: Epierus comptus (fribarinae,
Histeridae). E: Chaetabraeus bonzicus (Abraeinae, Histeridae). F: Plegaderus marseuli(Abraeinae, Histeridae). G: Dendrophilus xavieri (Dendrophilinae, Histeridae). H: Trypeticus
fagi (frypeticinae, Histeridae). I: Saprinus splendens (Saprininae, Histeridae).
20

A

Fig. 8. Antennae. A: Gnathoneus nannetensis (Saprininae, Histeridae). B: Hister coneolor (Histerini,
Histerinae, Histeridae). C: Platysoma vagans (platysomatini, Histerinae, Histeridae).

E

MentumoFig. 9. Mouthparts. A - D: Notodoma fungorum (Exosternini, Histerinae). E - H: Hololepta
amurensis (Hololeptini, Histerinae). A, E: Labrum. B, F: Maxilla. C, G: Labium. D, H:
Right mandible, dorsal view.
21

Labrum (Fig. 9A, E) is an unpaired plate, situated before the clypeus and isseparated 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 Synteliidae, Sphaeritidae and other
Hi steridae.
Mandibles (Fig. 90, H) are in a pair and well developed in the Histeroidea. The
inner margin is usually provided with denticles. In Notodoma and Hololepta, there isno denticle.
Maxillae (Fig. 98, F) consist of cardines, stipites, palpifers, laciniae, subgaleae,
galeae and palpi all in pairs. The palpus is 4-segmented, but may appear 3-segmented
because of the minute 1st 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 pal pus
consists of 3 segments.
THoRAX
CervixThe cervical elements are much reduced and composed of 2 pairs of lateralsclerites 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 theanterior margin of the composite ventral plate of the prothorax by membrane. It issmall, longitudinal oblong or triangular. The 2nd (apical) sc1erite is much longer than
the 1st.

Prothorax (Fig. 10 - 12)
The prothorax is divided into pronotum and prosternum, forming dorsal andventral plates. The lateral side of pronotum is inflexed ventrally. The inflexed part isseparated from the dorsal part of the pronotum by a distinct notohypomeral ridge orsuture. Elements of the tergum of the prothorax are completely fused to form a wellsclerotized 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 112 members of the Histeroidea, and is called "prosternal lobe". The
furcasternum is a pair of semiglobulate plates, but difficult to observe in normal
condition, forming the anterior wall or bottom of the fore coxal cavity which isconcealed under the fore coxa.
General shape and modifications of pronotum: The pronotum is nearlyrectangular, usually transverse and broadest near the base, usually its sides being
convergent apically. The anterior foramen is small, but that of Syntelia is larger thanthe posterior foramen (Fig. lOC). 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. lOC). The pronotum issubquadrangular in shape in the Niponiinae (Fig. 450), Trypeticinae (Fig. 12F) and22

Plegaderini (Fig. llH). It is impressed by a marginal pronotal stria which runs along
the notohypomeroal ridge and anterior margin in all Histeroidea. In Plegaderus (Fig.
l24A), a transverse stria and longitudinal striae are present near the middle and on
lateral 1/3, 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 - 4 pairs of costae are present
on the pronotum in Onthophilus. A transverse stria is found along the posterior marginof the pronotum in Bacanius, Australomalus (Fig. 83A) and Acritus. A pair of
longitudinal impressions is present on medioposterior area in Pachylomalus (Fig.
lOlA).
Punctation of the pronotum is very various in the Histeroidea. The pronotum isshining and smooth in some species of Hister, Margarinotus, Hypocaccus and soon. 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 invarious combinations. The punctation is often coarser and denser on the lateral regionin Atholus, Merohister, some Hypocaccus, etc. A pair of large foveae are present on
the lateromedian sides in Niponius impressicoliis (Fig. 43D, E).
Hypomera: The hypomeron is strongly inflexed ventrally and extends from theanterior to the posterior margin of the pronotum. The posterior 113 of inner marginusually strongly projects mesially. The apex of the projection is united with theposterior apex of the furcasternum in Syntelia (Fig. 1OC). In all other members of theHisteroidea 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. lOG), the apical 113 of the hypomeron 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 ispresent just before the composite ventral plate in the Synteliidae, Tribarinae,
Onthophilinae, Histerinae, and Dendrophilinae. It is called "prosternal lobe" anddistinctly separated from the composite ventral plate by a suture between prosternal keel
and lobe. In other histeroids the presternum is reduced. The presternum is triangularand strongly acute in Syntelia (Fig. lOC)o There is a semicircular or obtuse triangularplate in the Histerinae, Dendrophilinae, and Tribalinae. In the Onthophilinae (Fig.
lOG), it is not projected anteriorly, being transverse quadrangular. Presternum isusually impressed along the anterior margin by a marginal stria of prosternallobe.
Composite ventral plate: The composite ventral plate is usually transverse andabout 1/3 as long as wide. It is very variable in shape in the Histeridae and useful andimportant 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 primitiveand is observed in Sphaerites (Fig. WE).
2. Plate flat, the median portion weakly convex. The medioposterior portion is
projected, but not produced as a process, being concealed under the fore coxae.
This state may be 2nd primitive and is observed in Syntelia (Fig. 1OC).
3. The median portion of the plate is elevated as a broad process, while themedioposterior portion is represented by the intercoxal process (prostemal keel
23

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 thepresternum, and the anterior margin of the united plate (presternum + composite
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/~QQsZ2!l.8222S:28~)«composite ventral plate(basisternum + preepisternum)

Fig. 10 Prothorax. A, B: Necrodes asiaticus (Silphidae). C, D: Syntelia histeroides (Synteliidae). E,
F: Sphaerites politus (Sphaeritidae). G, H: Onthophilus ostreatus (Onthophilinae, Histeridae).
A, C, E, G: ventral view. B, D, F, H: caudal view.

24ridge is formed by an obliquely running anterior margin of the presternum, andat least partly closed beneath by the presternum or the anterior margin of thecomposite ventral plate (prosternal alae). This state is observed in the Tribarinae

Presternum(Prosternal lobe)

Prosternalprocess

Suture between
prosternal keel and lobe

Fig.

11. Prothorax. A: Holoiepta amurensis (Hololeptini, Histerinae). B: Plaesius sp.
(Platysomatini, Histerinae). C, D, E: Margarinotus cadavericola (Histerini, Histerinae). F, G:
Dendrophilus xavieri (Dendrophilini, Dendrophilinae). H. I: Plegaderus marseuli (plegaderini.
Abraeinae). A, B. C. F. H: ventral view. D: frontal view. E, G. I: caudal view.

25

Furcasternum

Fig. 12. Prothorax. A, B: Saprinus sp/endens (Saprininae). C, D, E: Gnathoncus nannetensis(Saprininae). F, G, H: Trypeticus Jagi (Trypeticinae). A, C, F: ventral view. B, G: oblique
view. D: frontal view. E, H: caudal view.

and Histerinae (Fig. llA, B, C).
5. Similar to the preceeding type, but the anterior cavities are very deep andconnected with foveae of the hypomeron. In this state, the lateral margin of theplate is deeply interrupted transversely on each side by the cavity. This state isobserved in Onthophilus (Fig. lOG).
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 plateis produced to form the antennal cavity on each lateral portion. The cavity issituated in front of the procoxa, and is represented by a longitudinal deep26

furrow, of which the inner ridge is strongly costate and formed by alongitudinally running anterior margin of the presternum. This state is observedin the Dendrophilinae (Fig. IIF).
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 isobserved in the Saprininae (Fig. 12A, C), Trypeticinae (Fig. 12F) and
Abraeinae (Fig. 12H).
A pair of striae called lateral marginal pro sternal striae are present on the basal 112of 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 ofcarinal striae.
Furcasternum: The furcasternum is longitudinal and narrow in Syntelia, thebasal apex being united with the projection of hypomeron. In Sphaerites, it istransverse and broad. In Histeridae, the furcasternum forms the anterior wall of thefore coxal cavity and is composed of a pair of semi globular plates.
Fore coxal cavities: The fore coxal cavity is transverse, broad and situated on
the basal 112 of the prothorax. Its anterior wall is composed of the composite ventral
plate and the anterolateral part of the furcastemum, its outer lateral wall belongs to thehypomeron, and its inner lateral wall is the posterolateral part of furcasternum. The
posterior wall is usually partially closed by the hypomeral projection, but is absent in
Plegaderus (Fig. 12H). In Syntelia, the posterior wall is completely closed by thehypomeral projection, of which the inner apex is continuous with the intercoxal process(Fig. IOC).
Mesothorax
The mesothorax is composed of mesosternum, anepisterna, epimera, trochantins,prepectus and composite ventral plate. The mesosternum is much smaller than the proand metasternum. The anapleural suture is usually present between the anepisternum

Anaepipleural -\-~--"I'------\
sutureD

E

Fig. 13. Epimeron and anepimeron of the mesothorax. A: Hololepta amurensis (Hololeptini,
Histerinae). B: Onthophilus ostreatus (Onthophilinae). C: Saprinus splendens (Saprininae). D:
Chaetabraeus bonzicus (Abraeini, Abraeinae). E: Dendrophilus xavieri (Dendrophilini,
Dendrophilinae).
27

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

Q

Anterior
phragrnar)

A

Epimeron 2\.:

MesonotumBc
Mesonotum

Anepisternurn 2

Epirneron 2

CompOSite ventral plate

Hanging structure of
scutumScutum+scutellum

G

Fig. 14. Mesothorax. A. B. C: Syntelia histeroides (Synteliidae). D, E, F: Plaesius sp.
(PI atysomatini, Histerinae). G: Hololepta amurensis (Hololeptini, Histerinae). A, D:
Mesothorax, dorsal view. E: Ditto, frontal view. B, F, G: Mesonotum, dorsal view. C: Ditto,
lateral view.

28

usually triangular, but it is often invisible in minute histerids. The mesonotum iscomposed of anterior phragma, prescutum, scutum+scutellum and postnotum. The
anterior phragma is well developed in the Histeroidea, and is represented by a pair oflarge 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,

Anepislernurn 3

Epirneron 3

B

ElytronFig. 15. Meso- and metathoraces in ventral (A, B, D) and lateral (C) views. A: Sphaerites politus(Sphaeritidae). B, C: Syntelia histeroides (Synteliidae). D: Onthophilus ostreatus(Onthophilinae).

2939B), the plate is longitudinal triangular. In the Histeridae, the scutum is welldeveloped, 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 ventral view, because it is situated under the mid coxa. 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 ispresent as arcuate collars inside the anterior margin of the anepisternum In theHisteridae.
Composite ventral plate (Fig. 15, 16): The composite ventral plate of theHisteroidea represents 2 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), andmore primitive than Type 2. In Sphaerites, the ridge is quadrangular and highly
elevated.
Anepistemum 2
Epimeron 2

Anepisternum 3

Epimeron 3

\

I
/

'--//

,, .............

B
"

---"

"

Fig. 16. Meso- and metathoraces in ventral (A, B) and lateral (C) views. A: Saprinus splendells(Saprininae). B, C: Plaesius sp. (Platysomatini, Histerinae).

30

Anepisternum

Fig. 17. Anepistemum and epimeron of the metathorax. A: Saprinus splendens (Saprininae). B:
Dendrophilus xavier! (Dendrophilini, Dendrophilinae). C:
Onthophilus ostreatus(Onthophilinae). D: Hololepta amurensis (Hololeptini, Histerinae).

Type 2. The median portion of plate is strongly elevated, its anterior margin isvery thick. The portion is called "intercoxal disk of mesosternum". The anterolateralside 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. 150, 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 somemembers of the tribe Exosternini. The stria of the Saprininae is usually strongly crenate
and impressed along meso-metasternal suture, and called "meso-metasternal stria". Theintercoxal 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 anintercoxal 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 scleritein 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 issituated before the epimeron (Fig. 17). In the 1st 2 families they are long in the31

Furcal arm
Basal stalkFig. 18. Metendosternite in caudal view. A: Sphaerites politus (Sphaeritidae). B: Syntelia histeroides(Synteliidae). C: Hololepta amurensis (Hololeptini, Histerinae). D: Onthophilus ostreatus(Onthophilinae). E: Dendrophilus xavieri (DendrophiIini, Dendrophilinae).

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 withthe posterior margin of the epimeron of the mesothorax. 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 usuallytransverse, and represents 2 types.
Type 1. The anterior margin of the intercoxal process between the mid coxae isnarrower than the width of the midcoxal cavity. This type is found in the Synteliidae
and Sphaeritidae (Fig. 15A, B).
Type 2. The anterior margin ·of the intercoxal process between the mid coxae isbroader 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 thelateral area "lateral disk of metasternum". The plate is impressed by the lateralmetasternal and posterior mesocoxal striae.
The discriminalline (median longitudinal line or metasternal longitudinal suture)of the composite ventral plate is usually found in the Histeroidea; however, it is lost in
32

Sphaerites (Fig. 15A), and is obscure in Chaetabraeus.
The following modifications are found on the plate. A deep fovea is present
behind each lateral anterior angle in Epiechinus (Fig. 53A). The male of some speciesof Onthophilus is furnished with long hairs. A small tubercle is found on the mid line
in Acritus.
Positi;n of hind coxae: The hind coxae are set close to each other in the
Synteliidae and Sphaeritidae (Fig. 15A, B), while they are broadly separated from eachother in the Histeridae (Fig. 150, 16A, B). These states are in accord with thecondition of the composite ventral plate of the metathorax.

Metendosternite (Fig. 18)
The metendosternite is composed of an unpaired basal stalk and the anterior andfurcal 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 bybroad and subtransparent laminae (Fig. 18A). 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, 0, E), the basal stalk is very short and transverse, and the furcal arms are broadly
separated and extended obliquely.
Elytra (Fore wings)
The elytra are strongly sclerotized, usually black, sometimes maculate mediallywith 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 in the Synteliidae, Niponiinae andTrypeticinae. In the Sphaeritidae, the elytra are elongate oval.
The side margins of elytra are usually round and slightly convergent apically, andsometimes parallel. The anteromesial margin of the elytron fits to the lateral margin ofmesonotum. The apical margin is usually truncate, exposing the last 2 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 andsituation of the striae and costae are very useful for species identification, andterminology for these characters has been proposed (Wenzel, 1944; Arnett, 1962;
6hara, 1989). Basically the elytron has 11 striae, which are named from the outside to
inside as follows: marginal epipleural, marginal elytral, external and internal
subhumeral, oblique humeral, 1st - 5th 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 anddensely 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.
33

I-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 mostprimitive state, being found in the Dendrophilinae, Tribalinae, and somegroups of the Exostemini.
I-b. Fifth dorsal stria is absent. Ends of 4th and sutural striae are usuallyunited 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 Exostemini.
I-c. Interstice between 5th dorsal and the sutural striae not very broad. Basalends 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 somegroups 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 ofMargarinotus.

M

Fig. 19. Basal area of hind wing. Merohister jekeli (Histerini, Histerinae, Histeridae).

34

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

i?

.. ~

''\
A

eu

E
Fig. 20, Hind wings. A: Thanatophilus sp. (Silphidae). B: Spheridium scarabaeoides (Hydrophilidae).
C: Sphaerites politus (Sphaeritidae), D: Syntelia histeroides (Synteliidae), E: Onthophilus
ostreatus (Onthophilinae, Histeridae).

35

(Merohister jekeli, Hololepta amurensis, Epierus comptus, Plegaderus sp.,
Tryponaeus sp., Sphaerites glabratus, and Syntelia histeroides), Ohara and Nakane,

eu

B

c

Cu

o

E
Fig. 21. Hind wings. A: Epiechinus arboreus (OnthophiIinae). B: Epierus sp. (fribalinae). C:
Trypeticus fagi (frypeticinae). D: Niponius impressicollis (Niponiinae). E: Plegaderus
marseuli (Plegaderini. Abraeinae).

36

1989 (Notodoma fungorum and Binhister chujoi), and Ohara, 1991 (Hololepta
amurensis) .

A

M

cu

c

Fig. 22. Hind wings. A: Chaetabraeus bonzicus (Abraeini, Abraeinae). B: Saprinus sp. C:
Hypocaccus varians. D: Gnathoncus nannetensis. E: Eopachy/opus riape (B - E: Saprininae).

37

The hind wing is functional in all examined species, but reduction rarely occurs(Crowson, 1974). The wing is usually transverse oblong and hyaline, and the veins aredark brown to yellow.

BoFig. 23. Hind wings of the subfamily Dendrophilinae. A: Dendrophilus xavieri (Dendrophilini). B:
Anapleus nakanei (Anapleini). C: Carcinops pumilio (ParomaIini). D: Australomalus
montivagus (ParomaIini). E: Pachylomalus sp. (paromaIini).

38

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 adoptedA~~\"';)~"'Cu;";:';;::':~~);\~:M;'''')/»''::":.?:.,:.:~.:.~"!"'~.'......

A

R

M

c

eu ".

o

Fig. 24. Hind wings. A: Eulomalus sp. (Paromalini. Dendrophilinae). B: Bacanius sp. (Bacaniini.
Dendrophilinae). c: Hetaerius optatus (Heateriinae). D: Procoryhaeus sp. (Exosternini.
Histerinae). E: Notodomafungrum (Exostemini. Histerinae).

39are given below. In terminology I follow Snodgrass (1935) with somereference toMorimoto (1983) and Naomi (1989), who proposed terms on the basisof relationships
between the axillary sclerites and the veins (Fig. 19).

Cu

Fig. 25. Hind wings of some species of the subfamily Histerinae. A:
Binhister chujoi (Exosternini).
B: Hololepta amurensis (Hololeptini) (after Ohara. 1991a). C: Omalodes sp. (Omalodini). D:
Platysoma (Apobletes) sp. (platysomatini). E: Platysoma (Platylister) sp. (Platysomatini).
F: Platysoma (Cylister) lineicollis (platysomatini).
40

HP: Humeral plate.
C: Costa runs on anterior margin of the wing and is unbranched; connected at
the base with the anterodistal projection of the 1st axillary sclerite.

M

E
Fig. 26. Hind wings of some species of the subfamily Hi sterinae. A: Ebisia satzumae. B:
Platysoma (Platysoma) vagans. C: Atholus duodecimstriatus quatordecimstriatus. D:
Merohister jekeli. E: Hister concolor. (A - B: Platysomatini; C - E: Histerini).

41

Fig. 27. Hind wings of 3 speceis of the tribe Histerini of the subfamily Histerinae. A: Pachylisterchinensis. B: MargarinotWi niponicus. C: MargarinotWi agnatWi.

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 theHisteroidea.
1. A definite M-Cu loop is present in the middle area of the wing. This state isprimitive (Crowson, 1955), and is found in the Sphaeritidae, Synteliidae, allgenera 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. 26C - E, 27A - C);
the anal lobe is bilobulate (represented by 2 lobules).
3. Hind wings of the Saprininae and Dendrophilus are very similar, and share a
42

o
A.~

__==~~__----~~~B
CF/AB( .)

0.2

0.3

0.4

CE/AB( ©)

0.3

0.4

0.6

0.5

AC/AB( .)

0.2

0.3

0.4

OUT GROUP
1. Sphaeridium scarabaeoides (L.)
2. Thanatophilus sp.

0.5

@

•
•

i.

@

SHAERITIDAE
3. Sphaerites politus Mannerheim
SYNTELIIDAE
4. Syntelia histeroides Lew.

•

HISTERIDAE
5. Niponius impressicollis Lew.

-!

9.

:t~~r~~~~:-::-:~--:--:l;~~":~~:]-~~;f:~~~:-~-:...................................................................................................... 000 ... ·· ...................................... (................ ············000········································· ... .

10. Hololepta amurensis Reitt.
•
11. NotodomafungroumLew.
•
12. Binhister chujoi Cooman•
13. Procoryhaeus sp.
•
14. Dmalodessp.
•
15. Platysoma (Platysoma) vagans Lew.
•
16. P. (Apobletes) sp.
•
17. P. (Platylister) sp.
•
18. P. (Cylister) lineicollis ( M a r s . ) .
19. Eblisia satzumae ( L e w . ) .

11©

····~~·.··~h;,;;;~;;,;;;;;·;;;;;;;;~·:.:;··(~~:)······················

©.

i © •
i
©.

if)! 1•i©

•i if)!
i ©

•

1©

•

1©

20. Hister concolor L e w . .
21. Atholus l2-striatus ( G y l l . ) .
22. Margarinotus niponicus (Lew.)•
23. Margarinotus agnatus ( L e w . ) .
24. Pactolinus chinensis ( Q u e n s e l ) .
25. Merohister jekeli ( M a r s . ) .
26. Dendrophilus xo.vieri Mars.
27. Anapleus nakanei M. Ohara
28. Bacanius sp.
29. Carcinops pumilio (Er.)
30. Australomalus montivagus (Lew.)
31. Pachylomalus musculus (Mars.)
32. Eulomalus sp.

•

!

i

•

•

if)!
if)!

•
if)!.

©
if)!

©

•

•

•

©
©
© .

•

©.
•

•
•

•

................; .................................. ...··T·························-:--·························...........
~

34. Plegaderus marseuli Reitt. .
• :
:
©
:•
...................................................................................................... ..... ······································c··.. ············ ............ ........................................... ..
35. Saprinus niponicus Dahlgreni$>~
36. Hypocaccus varians (Schmidt)
i.
©
i
i •
37. Eopachylopus ripae (Lew.)
1 •
©1
1.
38. Gnathoncus nannetensis (Mars.)
i•
«lJ i
i•
~~:··~~;~;~~;f~~·;·~~~:····························· ··················;······r·········································~·······~················r··············· ...........................
~

~

....

Fig. 28. Ratios in the hind wings.
43

characteristic and unique anal vein which is situated close to the Pcu and with aclear curve at its middle (Fig. 22B - E, 23A).
In insect wings, some general trends correlated with the absolute body size havebeen 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. 2IE, 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 areprovided.
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 withthe 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.
Trochanters
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.
FemoraThe fore, mid and hind femora are baculiform and thick, often with setae on theanterior (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) ispresent 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.
TibiaeThe shape of the tibia is subject to various modifications, which may vaguely beclassified into the following 3 types.
1. The tibiae are slenderer than the femur and baculiform. Denticles on the marginof the ti biae are very small.
2. The tibiae are somewhat expanded apically, and wider than the corresponding
femur. The denticles are large and well developed.
3. The tibiae are strongly dilated, thus more or less round in outline. The denticles
are weakly developed.
44

The 1st type may be the most primitive and is found in the Sphaeritidae and somegroups of the Histeridae, that is, Onthophilinae, Niponiinae, Trypeticinae and some
genera of the Exosternini (Fig. 29A, G). The 2nd type is common in the Histeridae.

L
Fig. 29. Legs. A, B, C: Sphaerites politus (Sphaeritidae). D, E, F: Syntelia histeroides (Synteliidae).
G, H, I: Onthophilus ostreatus (Onthophilinae, Histeridae). J, K, L: Hololepta amurensis(Hololeptini, Histerinae). A, D, G, J: Fore legs. B, E, H, K: Mid legs. C, F, I, L: Hind legs.
45

The 3rd type is present in some species of the Hetaeriinae 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 tribes Dendrophilini

Fig. 30. Legs. A, B, C: Margarinotus cadavericola (Histerini, Histerinae, Histeridae). D, E, F:
Dendrophilus xavieri (Dendrophilini, Dendrophilinae). G, H, 1: Chaetabraeus bonzicus(Abraeini, Abraeinae). J, K, L: Saprinus splendens (Saprininae). A, D, G, J: Fore legs. B, E,
H, K: Mid legs. C, F, I, L: Hind legs.

46

and Anapleini. The ventral surface of the tibiae is densely covered with long hairs or
robust or long setae in some members of the Hypocaccus, Eopachylopus (Fig. 155)
and Philo this. This condition is often associated with their habitats in sand on beach.
Tarsi
The tarsus is composed of 5 tarsomeres in general, but in Acritus 4-segmented
in the hind leg, thus the tarsal formula is 5-5-4. The relative lengths of the tarsomeresare rather uniform, the 1st - 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
sc1erotized in the Histeridae. On the other hand, in the Sphaeritidae and Synteliidae, thedorsum 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 - 7th sterna areexposed and strongly sc1erotized. The 8th, 9th and 10th segments are almost telescoped
into the 7th.
Abdominal segments (Fig. 31)
Each of the 3rd - 7th terga is transversely rectangular in shape, and a transverse
tergal suture runs along the basal margin.

f
A
\
\

.. _--_ .....II
I

cFig. 31. Abdomen. A: Sphaerites poUtus (Sphaeritidae). B: Saprinus splendens (Saprininae). C:
Plaesius sp. (platysomatini, Histerinae).

47

The basal margin of the 3rd sternum is longitudinally carinated medially and thesides 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 1 or 2 pairs of
striae on each side of the median area. In taxonomic descriptions the disk of the 3rd isoften called" 1st abdominal sternum" and the striae" 1st abdominal striae", because thetrue 1st and 2nd disks are absent and the 3rd appears to be the 1st. The 4th - 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 forreceiving the posterior margin of the 7th tergite (pygidium).
Oenital segments
The 8th, 9th and 10th abdominal segments are called genital segments. Thesesegments 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 2 plates by a longitudinal mid line. A pair of short or longprocesses 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 2 plates bya longitudinal mid line, and each plate is elongated posteriorly and rounded apically in
the Sphaeritidae (Fig. 38), Carcinops (Fig. 96), some members of the tribe
PI atysomatini (Fig. 58, 59, 60) and Histerini. The ventral surface is strongly
sclerotized and densely covered with hairs in Saprinus (Fig. 137, 139, 140) and most
Saprininae. The 8th sternum is oblong in ventral view, and the caudal disk with around disk or Some small oblong disks in the tribe Paromarini (Fig. 99, 102, 105, 108,
1l0, 113, 115, 117, 120).
The 9th tergum is often divided longitudinally. This state is found in
Gnathoncus, Carcinops, Anapleus, Epiechinus and Platysoma (Fig. 132, 133,
134, 96, 87, 94, 54, 58, 59, 60, 62, 65). The anterolateral corner is projected. In
Trypeticus, the projection is very long (Fig. 165). The posterior margin is connectedwith the 10th tergum and nearly straight, but often emarginate arcuately or acutely forfitting 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 athin 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, 129F, 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. 870, 940). Platylomalus (Fig. 1l0F) and Dendrophilus (Fig.
82F). The lateral angles of the posterior apex are elongated posterolaterally, thus thesternum is T-shaped in the tribe Histerini and Platysomatini. In the Saprininae too, it isoften shaped like a T, the anterior apex often expanded (Fig. 1321, J, 1331, 1341, 1371,
1381,1391, 140H, 1480, 1490, 1500, 1510, 1550, 1570, 161H). The projectionsof the posterior apex are united together on dorsal side through membrame to form aring, into which the basal piece is fitted. In Merohister uenoi, the spicule forms asclerotized ring (Ohara, 1992b).
48

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

Male genitalia
Male genitalia consist of 3 parts: basal piece, a pairs of parameres, and medianlobe. Sharp and Muir (1912) first made comparative (study of Coleoptera. In
Hi steridae, the male genitalia are considered to be the most useful character for theclassification 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 themedian 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, 4OA, B, 87A, B, 89A, B, 94A, B). This condition
may be the most primitive in the Histeroidea. It is absent in some distant taxa: Acritus,
Bacanius and part of Hetaeriinae. The basal piece is variable in length in theHisteroidea. It is usually 112 as long as the paramere, but, in Niponius (Fig. 44A,
47A), members of the tribe Paromalini (Fig. 96A, 99A, 102A, 105A, 108A, 110A,
113A, 115A, 117A, 120A, 122A), Epierus (Fig. 51A), Trypeticus (Fig. 165A) andsome 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 andlobate structures in the Coleoptera. In the Histeroidea, the parameres are usually fusedbasally on the ventral side to form a tube structure. Their dorsal side is usuallyprovided 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 andalso separated in their apical halves ventrally; the median lobe is extruded from theventral side. In the majority of the Histeridae, the parameres are partly fused in thebasal 112 and moderately separated in the apical 112; the median lobe is extruded fromthe dorsal side. The median lobe is, however, extruded from the ventral side in
Hololepta and some members of the tribe Exosternini (Ohara 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 isstrongly emarginate; in members of the Paromalini, it is short.
Median lobe: The median lobe is usually slender and is surrounded by theparameres and the basal piece. It is various in shape, varying from a flat and weaklysclerotized 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. Thiscondition 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, Exostemini, Platysomatini and somemembers of the Histerini.

49

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: Rister, Atholus, Zabromorphus, Merohister, Margarinotus andso on (Ohara, 1989, 1992a, 1992b, 1992c, 1993).

L<
Fig. 32. Spermatheca and spermathecal glands of the female genitalia. A: schematic. B: Sphaerites
politus (Sphaeritidae). C: Syntelia histeroides (Synteliidae). D: Niponius impressicollis(Niponiinae, Histeridae). E: Trypeticus fagi (Trypeticinae). F: Chaetabraeus bonzicus(Abraeinae). G: Plegaderus marseuli (plegaderini, Abraeinae). H: Hypocaccus varians varians(Saprininae). I: Gnathoncus nannetensis (Saprininae). J: Carcinops pumilio (paromalini,
Dendrophilinae). K: Platylomalus niponensis (Platysomatini, Dendrophilinae). L: Eulomalus
sp. (paromalini, Dendrophilinae). M: Onthophilus ostreatus (Onthophilinae). N: Epierus sp.
(Tribalinae). sp: spermatheca; spg: spermathecal gland; bc: bursa copulatrix.

50

Fig. 33. Spennatheca and spennathecal glands of the female genitalia of the subfamily Histerinae. A:
Hololepta amurensis (Hololeptini) (after Ohara, 1991a). B: Platysoma sp. (platysomatini). C:
Omalodes sp. (Omalodini). D: Notodoma fungorum (Exosternini). E: Merohister aino. F:
Hister japonicus. G: Hister concolor. H: Hister congener. I: Atholus pirithous. J:
Margarinotus sp. K: Margarinotus sp. L: Margarinotus sp. (E - L: Histerini).

51

/)

\\~IA~~~,Histerini(Hister, ect.)

Yl+

~J

Histerini(Margarinotus)

Saprininae

Sphaeritidae
Synteliidae
Histeridae
Abraeinae(Chaetabraeus)
Niponiinae
Trypeticinae
Onthophilinae
Tribalinae
Oendrophilinae(Oendrophilus)
Histerinae(Exosternini,
Hololeptini,
Platysomatini)Abraeinae(Plegaderus)
Oendrophilinae(Carcinops)

1

iroPhmn"(Paromalini)

Fig. 34. Spermatheca and spennathecal gland: transformation series.

Female genitalia
Spennatheca: The spermatheca is the sperm receptacle of the female. De Marzoand Vienna (1982c) first made a careful morphological study of the organ in theHisteridae, and recognized 2 fundamental patterns: I) spermatheca including 1
receptacle, a more or less long duct and a distinct gland; II) spermatheca including 4 - 9
sessile receptacles, without a distinct spermathecal gland. The 2nd pattern is found in
the tribe Histerini, while the 1st 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 shaped like a globe, strongly sclerotized.
1. Without a 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 notcoiled.
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
52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 areaconnected with the duct of spermatheca and also around the area connected with thespermatheca! gland. Plegaderus has a pear-shaped spermatheca feebly invaginate
apically.
The 3rd type is found in Niponius (Fig. 320) and Pia tyloma Ius (Fig. 32K).
Probably this type has independently appeared in these genera.
The 4th type is found in the Dendrophilinae, with 1 internal projection in
Carcinops and 2 in Pia tyloma Ius (Fig. 321) and Eulomalus (Fig. 32L).
The 5th and 6th types are present only in the tribe Histerini, the former isobserved in Hister, Atholus, Merohister, Zabromorphus, and Pactolinus (Fig. 33E,
F, G, H, I), while the latter is found only in Margarinotus (Fig. 331, K, L).
The spermathecal gland is usually a small saccule, weakly sci erotized and with along straight or coiled spermathecal duct, but is sometimes absent.

5. PHYLOGENY
Phylogenetic Relationships of the Families
The superfamily Histeroidea was adumbrated by Sharp and Muir (1912), withthe remark that "four families Histeridae, Synteliidae, Sphaeritidae and Niponiidae areso closely related by the aedeagus, that they might form one family". The group was
adopted by most histerid systematists (Reichardt, 1941; Wenzel, 1944; Crowson, 1955;
1974; Kryzhanovskij and Reichardt, 1976; Mazur, 1984; Hisamatsu, 1985b).
The superfamily Histeroidea is included in the Staphyliniform lineage, which isoften divided into 3 superfamilies: the aquatic Hydrophiloidea and the terrestrial
Histeroidea and Staphylinoidea. The Histeroidea have been considered the most closely
related to the superfamily Hydrophiloidea by some authors (Boving and Craighead,
1931; Reichardt, 1941; Morimoto, 1986). On the contrary, Crowson (1955; 1974)
insisted that the Histeroidea are more closely allied to Staphylinoidea than to theHydrophiloidea. But recent works tend to refute Crowson's hypothesis. Discoveries oflarvae in the primitive hi steroid families Sphaeritidae and Synteliidae (Nikitsky, 1975;
Mamayev, 1976) have shown that these larvae share numerous specializations with
Histeridae and Hydrophilidae.
The Histeroidea and Hydrophiloidea 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 (pal pifer) of maxillary palp complete
and bearing articulated appendage, 4) tentoruium with posterior arm attached directly tohead, 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 instar without ecdysial line on head. Known pupae lack functional
spiracles on the first abdominal segment. Adult antennae are short, usually with asharply differentiated club composed of 3 densely pubescent segments; abdominal
segment 7 is invaginated at least ventrally; and the 7th or 7th and 8th spiracles areatrophied. Nearly all larvae are carnivorous with mouthparts adapted for extraoraI
digestion. Recently Lawrence and Newton (1982) have proposed to include theHisteroidea in the Hydrophiloidea (excluding Hydraenidae). According to them, there53

Table 2. Character states used in Wenzel-Mazur System and their presumed polarities.
No. Mnemonic
PsLo

Code

State

o

Prosternal lobe present (Histeromorphae).
Prosternal lobe absent (Saprinomorphae).
Antennal cavities absent.
Antennal cavities closed beneath by the prosternal alae.
Antennal cavities [if present] open beneath.
Antennal scape normal, the club usually a little longer than but
never 3 times as long as broad.
Antennal scape strongly angulate and usually greatly expanded,
the club greatly elongated, at least 3 times as long as broad.
Form round, oval or oblong-oval.
Form cylindrical, sometimes stoutly so.
Antennal cavities absent.
Antennal cavities situated in anterior prothoracic angles, atleast partly closed beneath by prosternal alae.
inanterior
prothoracic
Antennal
cavItIes
situatedangles, wholly closed beneath by prosternal alae.
Antennal cavities situated in front of procoxae, next to and nearly
always encroaching upon prosternal keel.
Antennal cavities situated in anterior
prothoracic angles, orin f ron t of procoxae.
Dorsal elytral striae rarely absent, if so, then at least
sutural stria present.
Dorsalelytral stria never present thoughrather vague
impression may be discernible.
Head not produced into 2 horns, though it may be produced as a longpointed rostrum in the female.
Head produced anteriorly into 2 horns of various length.
Mandibles moving on the same plane as the long axis of the head.
right angle to
Mandibles deflexed, moving on a plane at
the long axis of the head.
Head horizontal in repose.
Head vertical in repose.
Antennae consisting of 8 segments and a club.
Antennae consisting of 7 segments and a club.
ofAnterior
margin of prosternal alae without reception
antemnal funicle.
Anterior margins of prosternal alae with rather deep longitudinal incisions for the reception of the anternnal funicle
in repose.
of prosternalalae
at most with a very
Anterior margin
slight notch. Antennal cavities not defined.
Antennal scape normal, neither expanded nor strongly angulate.
Antennal scape expanded and strongly angulate.
Labrum wi th setigerous punctures
Labrum without setigerous punctures.
Antennal cavities absent.
Antennal cavities situated in anterior prothoracic angles, atleast partly closed beneath by prosternal alae.
Antennal
cavities completely
open, usuallysituated justanterior to procoxae at middle of hypomera.
Protibiae multidenticulate.
Protibiae oligodenticulate.
Dorsal surface without costae.
Dorsal surface with distinct costae.

1

2

AnCa

o
1
2

3

AnSc

o

4

Form

o

5

AnCa

o

1
1

2

3
4
6

ElSt

o

7

HeHn

o

8

Mand

o

1

1
9

HeSt

o

10

AnSe

o

11

PrAI

o

1

1

2
12

AnSc

o

13

LaSe

o

14

AnCa

o

1

1

1

2
15

PtDe

o

16

DoCo

o

1
1

are no sufficient differences between these taxa to divide them 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.

54

r - - - Saprinomorphae _ _ _--,

Nip.

Tpa.

Tpe.

Ahr.

Sap.

[hi.

r--Histeromorphae I
Onto

Tri.

Den.

His.

Het.

Fig. 35. Wenzel-Mazur System in a cladogram.

The superfamily Histeroidea consists of3 families, Sphaeritidae, Synteliidae andHisteridae, and is characterized in adult by the compact antennal club, very
prominent,acute mandibles, truncate elytra covering 6 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 a distinct family (Gardner, 1926; Nakane,
1963; Hisamatsu, 1985b). Since Reichardt (1941), however, the majority of thehisterid systematists have adopted a system consisting of the 3 families. The
phylogenetic relationships among these families have been written fragmentally bysome 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 classifications of the Histeridae (seeHistorical Review), but no cladograms have yet been published and no studies havebeen presented with specific intent to distinguish between apotypic and plesiotypiccharacter states.
55

Nip.

Chi.

Onto

Tri.

His.

Het.

Den.

"hr.

Sap.

Tpa.

Tpe.
AnSe 1

Fig. 36. Cladogram proposed in the present paper.

Since Wenzel (1944) proposed a new system, most authors (Hatch, 1962;
Mazur, 1981; Vienna, 1980; Yelamos, 1985 and Hisamatsu, 1985b) have followed
him. Mazur (1984a) 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 apotypic-plesiotypic
relationships.
The subfamilies are so diverse and distinct that it has been difficult to findadditional characters which can be used to construct morphoclines through them. In thissection, 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 in Arnett, 1962) and Mazur (1984)
did not distinguish between primitive and derived states of the adopted characters.
Hence details of the phylogenetic relationships 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 theWenzel-Mazur System.
The character states are given in Table 2. They have been picked up from the 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 the56Table. 3. Data matrix for subfamilies of Histeridae.
No.
0
9
7

8
5
11
3
4
6
10
13
16

MsWi
HeSt
HeHn
Mand
AnCa
PsLo**PrA!
AnSe**
Forrn**
ElSt
AnSe
LaSe
DoCo

Ane
0
0
0
0
0
0
0
0
Oil
0
0
0
0

Nip
1
0
1
0
0
0
0
1
0
0
0
0

Chl
1
0
0
2
0
1
0
0
0
0
0

Ont
1
0
0
2
0
0
0
0
0
0
0
1

Tri
1
1
0
0
1
0
0
0
0
0
0
0
0

His
1

Het
1

Den
1

Abr
1

Sap
1

0
0
1
0
0
0
0
0
0

0
0
1
0
0

0
0
3
0
1
0
0
0
0
0
0

0
0
4

0
0
3

0
0
1
0
0
0

1
0
0
0
0
0
0

0

0
0
0
0
0

Tpa
1
0*
0
0
3

Tpe

0
0
3

2
0

0

0
0
0
0

0
1
0
0

1
1

1

Ane: Synteliidae and Sphaeritidae; Nip: Niponiinae; Chl: Chlarnydopsinae; Ont: Onthophilinae; Tri:
Tribalinae; His: Histerinae; Het: Hetaeriinae; Den: Dendrophilinae; Abr: Abraeinae; Sap: Saprininae;
Tpa: Trypanaeinae; Tpe: Trypetieinae.
* Reversal; ** Parallelism.
No. 0, MsWi, code 0: mesosternum between mesoeoxae narrower than mesoeoxal cavity; code 1:
mesosternum between mesocoxae broader than the mesocoxal cavity.

table. The adopted out-groups are Synteliidae (Syntelia histeroides) and Sphaeritidae(Sphaerites politus). However, Character 2,5 and 14 could not be determined as to thepolarities of their states, though these are arranged with code numbers. Character 5 has
5 states (Code 0 - 4), which may not be in a linear relationship. Character 1, 6, 10 and
13 include negative or reductive states, which are interpreted as derived.
The diagram thus constructed is shown in Fig. 35. It includes 5 cases ofhomoplasy, 3 of reversal and 2 of parallelism.
However, I do not think that 2 of the indicated cases 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 caseof AnCa may not represent real parallelism. The derived state of AnSc is apparently
associated with the myrmecophilous habit in the Chlamydopsinae and Hetaeriinae.
Proposed cladogram. In Section 4 (Morphology and Terms) I examined andcompared many external characters, but I could not find any additional characters usefulin phylogeny reconstruction at the subfamily level except the mesosternal width.
In Table 3, the mesosternal width and the characters mentioned in Table 2 arearranged 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 isconstructed (Fig. 36). The new cladogram is characterized by the erection of 3 majorgroups instead of 2 (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 2 supposed reversal states in Fig. 35 are cleared in the new cladogram.
2) Wenzel (1944) recognized the 2 major groups on the basis of Character PsLo.
However, the prosternal lobe is present (PsLo: 0) in the Niponiinae, which were57

erroneously included in the Saprinomorphae by Wenzel. Moreover, Character AnCa ismore weighed than PsLo in the new cladogram; the Dendrophilinae, therefore, are not ahisteromorph but a saprinomorph subfamily. In the new cladogram it is postulated thatthe 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 theHisteromorphae, 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 toconform 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 beplaced next to the Onthophilinae and the Tribalinae.
4) The parallel evolution of Character Form (Code 1) may be associated withliving in the gallery of scolid beetles, the concerned histerids preying on them.
In the Saprininae and Dendrophilinae the anal vein of the hind wing is distinctly
curved at basal 113. This may be a common derived state in these subfamilies,
supporting the new c1adogram. (In the Abraeinae, Trypanaeinae and Trypeticinae theveins of the hind wing are strongly reduced in association with their small body size.)
These 2 subfamilies also agree in having a large and triangular gular plate.
6. SYSTEMATICS

Key to the families of the Histeroidea
1(4)

2(3)
3(2)
4(1)

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

6. 1. FAMILY SPHAERITIDAE THOMSON, 1862
GENUS SPHAERITES DUFTSCHMIDT, 1805
Spaherites Duftschmidt, 1805: 205 [type species: Hister glabratus Fabricius, 1792: 73]; Lacordaire,
1854: 212; Thomson, 1862: 23; Reitter, 1885: 88; 1909: 246; Ganglbauer, 1899: 415;
Jakobson, 1915: 289; Reichardt, 1941: 5; Arnett, 1962: 289; Witzgall, 1971: 189; Hisamatsu,
1985b: 219, pI. 40, no. 1.
Sarapus Fisher-Waldheim, 1821: 39 (nom. nud.).

A good description is given by Arnett (1962).
Nikitsky (1976) described the larva of Sphaerites glabratus.

58

A

Fig. 37. Sphaerites politus Mannerheim. A: Adult, dorsal view. B: Pygidium, dorsal view. C: Right
protibia, dorsal view. D: Ditto, ventral view.

Sphaerites poUtus Mannerheim, 1846

(Fig. 37, 48, 41)
Spaerites politus Mannerheim, 1846: 514; Kryzhanovskij and Reichardt, 1976: 22; Adachi and Ohno,
1962: 149; Nakane, 1963: 67, pI. 34, no. 1; Matsumoto, 1981: 6 [Mt. Hidaka]; Yasuda, 1982:
76 [Mt. Daisetsu]; Hisamatsu, 1985b: 219, pI. 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 comer 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 112, and nearly straight on basal 112; posterior margin strongly arcuate
outwards in median 1/3; marginal stria complete laterally, its apical end curved behindapical comer, extended inwards, and attaining to near the lateral 113 of pronotum, thestria somewhat distantly present from the pronotal margin; surface sparsely andcoarsely punctate laterally, and with other fine punctures intermingled, the finepunctures progressively finer inwardly; on median area of the surface clothed withmicroscopic punctures.
Epipleura narrow. Marginal epipleural stria complete and carinate. Marginal59

elytral stria strongly carinate and complete. Area between the margin of epipleura andthe marginal elytral stria shining and sparsely clothed with several microscopic
punctures. Elytra (Fig. 37A) each with 10 nearly complete rows of coarse and round
punctures, which are separated by 1.5 (1 - 2) times the diameter but irregularly
scattered on apical 116; 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 comer, curved before the corner and shortly extended
inwards; surface of disk densely covered with coarse punctures which are a littlecoarser than in elytral rows; other fine punctures intermingled among them, becoming
sparser apically.
Antenna! 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; medianarea 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, thepunctures as large as those of lateral disk of metasternum; on area along the posterior
margin transverse stria present, but broadly interrupted on median 1/3, the median endof the stria densely with coarse punctures.
Protibia (Fig. 37D, C) slender, not expanded, usually with 7 denticIes on outermargm.
Male genitalia as shown in Fig. 38A - F.
Female genitalia: spermatheca as shown in Fig. 380.
Specimens examined. [Hokkaido] 1if' 3 !f !f ,lex., Mt. Kurodake (Alt. 1900m), Mt. Daisetsu,
18/vii/1981, N. Yasuda (NA); 7 exs., Ditto, 23/vii/1981, N. Yasuda; 4 exs., Ditto, 28/viiI1982, N.
Yasuda; 2 exs., Akadake, 16-23/vii/1975, M. Suwa, M. Furukawa, M. Kiuchi and T. Sunose (ElHU).
Rishiri Is.: 1 ex., Rishiri Is., 24/vii/1963, Y. Shibata (NSMT); 1 ex., Oniwaki trail, Mt. Rishiri,
7/viii/1992, M. Sato.

Distribution (Fig. 41). Japan (Hokkaid6, Rishiri Is.); North America.
Table 4. Biometric data of Sphaerites poUtus Mannerheim and Syntelia histeroides Lewis.

APW
PPW
PL
EL
EW
PyW
PyL
PTL
MSTL
MITL

Sphaerites politus

Syntelia histeroides

1.31-1.63 (1.47±0.03) 9
2.38-2.81 (2.65±0.05) 9
1.25-1.56 (1.48±0.04) 9
2.75-3.75 (3.41±0.09) 9
2.94-3.56 (3.28±0.07) 9
1.63-2.00 (1.85±0.04) 9
0.88-1.25 (1.05±0.05) 9
1.13-1.50 (1.33±0.04) 9
1.25-1.56 (1.35±0.03) 9
1.50-1.94 (1.72±0.05) 9

3.00-4.38 (3.64±0.08) 20
2.50-3.75 (3. 15±0.07) 20
4.88-6.63 (5.84±0.1O) 20
3.25-4.75 (4.09±0.08) 20
2.00-3.13 (2.61±0.05) 20
2.13-3.13 (2.61±0.05) 20
1.75-2.50 (2.19±0.05) 20
1.88-2.50 (2.11±0.04) 20
1.88-2.75 (2.30±0.05) 20

60

,0.5 "''''

Fig. 38. Sphaerites poUtus Mannerheim. A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view. G: Spermatheca.

Remarks. Sphaerites politus can rather easily be recognized by the characteristics
given in the key to the families.
Little is known about the habitat of this species. This species occurs at highaltitudes. Yasuda (1982, 1988) collected it by using pit-fall traps in the forest of Pinus
pumila at alt. 1800 - 1980 m on Mt. Kurodake, Daisetsu, Hokkaido.
6. 2. FAMILY SYNTEI.1IDAE LEWIS, 1882
GENUS SYNTEUA WESTWOOD, 1864
Syntelia Westwood, 1864: 11 [type species: Syntelia indica Westwood, 1864]; Reitter, 1876: 18;
Sharp, 1891: 439, t. 14, fig. 1, Ib; 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 Hokkaido, Kumakawa in Higo,
Sado]; Sharp and Muir, 1912: 511, t. 55, fig. 77,438; Ganglbauer, 1899: 415; Jakobson,
1915: 869; Hetschko, 1926: 13; Matsumura, 1931: 16; Yuasa, 1930: 257; Kryzhanovskij andReichardt, 1976: 414; Nakane, 1963: 67, pI. 34, no. 2; Hisamatsu, 1985b: 219, pI. 40, no. 2.

61

Japanese name: Emma-mushi-modoki.

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 4. 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 withmicroscopic 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 alongposterior 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 thanthose 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 about 7 (5 - 10) 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 114; its basal and apical ends extended inwardly along basal and apical margins ofelytra, and the ends united with basal and apical end of sutural stria respectively; area
between marginal epipleural and elytral striae with 3 rows of setiferous punctures on~~",!i\I}J~ '.
•

"','.

I,'

:;: :.:

(",

7\··
S .'

··········,··,'.·····················r.·······,···. . . . . . ..

'/
,',:

o

A
Fig. 39. Syntelia histeroides Lewis. A: Adult, dorsal view. B: Scutellum, dorsal view. C: Pygidium,dorsal view. D: Right protibia, dorsal view. E: Ditto, ventral view.
62dorsal 114, the setae being long and stout. External subhumeral stria (Fig. 39A)
complete and densely covered with coarse punctures. Internal subhumeral stria usuallypresent on basal 113, but absent on humeral area (nearly basal 1/6). First dorsal stria
nearly complete, but abbreviated on basal 116 (humeral area) and apical 114. Second
dorsal stria complete, usually abbreviated on apical 1/6. Third dorsal stria present on
apical 112, but shortened on apical 116, and short rudiments present at base and on
median 113 of apical 112. Fourth dorsal stria absent, sometimes with a short rudiment
which consists of 2 or 3 punctures occurring on apical 116. Fifth dorsal stria present on
about median 113, but sometimes shortened apically, and a short rudiment present on
apical 115. 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 thedepression 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 113
of head on mid line.
Anterior margin of prosternal lobe outwardly cuspidate at middle, and with long
hairs; disk of the lobe impunctate. Prosternal process divided into anterior and posterior
parts by procoxae; apical 2/3 of the anterior part elevated and with smooth disk, and thebasal 1/3 triangular, its basal margins strongly convergent along procoxae, the apicalmargin of process with 2 or 3 long setiferous punctures; the posterior part short, oblong
and im punctate.
Mesosternum moderately convex on median area; surface densely covered withcoarse punctures, the punctures becoming denser medially, producing many rugae; areaof these rugae clothed with other dense and fine microscopic rugae; area before
posterior comer and between mesocoxae impunctate.
Meso-metasternal suture shortly present. Metasternum deeply with an arcuate
depression between mesocoxae; intercoxal disk shallow and longitudinally sulcate on
longitudinal mid line, and impunctate; lateral disk densely covered with coarse andshallow 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.
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 2 denticles on apical 113
of outer margin.
Male genitalia as shown in Fig. 40A - G.
Female genitalia: spermatheca as shown in Fig. 40H.
Specimens examined. [Hokkaid6] 8 exs., Nopporo, 19,20,22, 26/vi, 24/vii/1987, M. Ohara; 1ex., Ditto, 30/vi/1987, S. Kudo; 10 exs., Eniwa, lzari-gawa, 16/vi/1986, M. Mori; 1 ex., Maruyama,
Sapporo, 3/vii/198S, M. Ohara; 1 ex., Ditto, 22/vi/1940, B. Nishio; 1 ex., Mt. Moiwa, Sapporo,
S/vii/1978, T. Fujisawa; 2 exs., Ditto, IS/viIl904, no collector's name (NSMT); 1 ex., Heiwa-noTaki, Sapporo, 2/viiil 1977, T. Fujisawa; 2 exs., Ditto, l/vi/1966, N. Ueda; 1 ~ , Sapporo, 31/v/1976,
M. Furukawa; S exs., Hakkenzan, Jozankei, 7/vi1l976, 16/vi/1982, M. Mori; 3 exs., Ditto, 11,24/vi/1978, T. Fujisawa; 1 ex., Misumai, Sapporo, 10/vi/1979, A. Shinohara (NSMT); 3 exs.,
63

E

..~
... .
f'

//

(

:=. . .'./.

~. . . . . . . . .•. .=~. . . . . . . . . . . . ......

/

~

'

-"

,

."".. F..

O.5mm I

Fig. 40. Syntelia histeroides Lewis. A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view. G: Spermatheca.
JOzankei, 17/vi, 13/vii/1985, M. Ohara; 1 ex., Shikotsu-ko, 6/viii/1971, Y. Doi (NSMT); 1 ex.,
Enshurin, Tomakomai, 3/viii/1974, A. Kashizaki; 1 ex., Ditto, 18-30/viii/1978, H. Detani; 2 exs.,
Okusawa, Otaru, 29/vii/1955, Y. Watanabe; 5 exs., Mt. Yotei, Hirafu, Kutchan, 15-16, 22/vii/1991,
M. Ohara; 1 ex., Daisengen, 17/vii/1973, T. Ichikawa.
[Honshu] Miyagi-ken: 1 ex., Moriyasu, Akiu-mura, 24/vi/1951, no collector's name (NSMT).
Tochigi-ken: 1 ex., ChUzenji, 14/viii/1917, E. Gallois (NSMT). Gumma-ken: 1 ex., Hoshi-Onsen,
6/vi/1953, Y. Kurosawa (NSMT). Saitama-ken: 1 ex., Oku-Chichibu, 24/vii/1940, K. Kojima(NSMT). Nagano-ken: 1 ex., Mt. Yatsu-ga-take, 25/vii/1957, S. Nomura (NSMT); 1 ex., Mt. Chausudake, Yokokubo-zawa, 1600 m, S. Jap. Alps, 24/viiI1940, S. Ueno (NSMT).

Distribution (Fig. 41). Japan (Hokkaido, Honshu, Kyushu).
Remarks. Syntelia histeroides is a distinctive species within the superfamily inits shape and rather large size. It cannot be confused with any other species of thehisterids.
This species frequents sap of elm and is sometimes found under the bark. Larva
is found under bark.

64

./~

Sphaerites politusV.j-<Syntelia histeroides

Fig. 41. Collection sites of Sphaerites politus and Syntelia histeroides in Japan.
6.3. FAMILY HrSTERIDAE QYllENHAL, 1808

Key to the Japanese subfamilies of Histeridae
Prosternum without antennal grooves or cavities. Ventral side of head with large foveae forreception of antennae. Mandibles vertically connected with head ............. Subfamily Niponiinae
2(1)Prosternum with antennal grooves or cavities. Ventral side of head without foveae (except inTribes Hololeptini and Histerini, which have shallow and narrow longitudinal grooves).
Mandibles porrect, horizontally connected with head.
3(12) Antennal grooves or cavities on prostemum transverse, occurring on anterior side, and usuallyclosed beneath by the pro sternal alae.
4(9) Labrum with a few setiferous punctures.
5(6)Lateral sides of elytra strongly elevated. Prostemallobe absent, not distinctly separated fromprocess by prosternal suture ..................................................... Subfamily ChJamydopsinae
6(5)Lateral sides of elytra not elevated. Prosternallobe 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) Prostemal lobe present.............................................................. Subfamily Dendrophilinae
14(13) Prosternallobe 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.
1(2)

65

19(20) AntelUlae consisting of 8 segments and a club. Head horizontal in repose. Anterior margin of
prosternal alae at most only slightly notched. Antennal cavities not distinct. ........................ ..
................................................................. ............................... Subfarnily Trypanaeinae
20(19) AntelUlae consisting of 7 segments and a club. Head vertical in repose. Anterior margin of
pro sternal alae with rather deep longitudinal incisions for reception of the antelUlal funicles inrepose ....................................................................................... Subfamily Trypeticinae

6.3. 1. SUBFAMILY NIPONIINAE FOWLER, 1912
GENUS NIPONlUS LEWIS, 1885

Niponius Lewis, 1885b: 333 [type species: Niponius irnpressicollis Lewis, 1885b: 333, designatedby Gardner, 1935: 4]; Marseul, 1888: 148; Jakobson, 1911: 638, 642; Gardner, 1926: 194;
1935: 2; Reichardt, 1926a: 274; 1941: 66; Schenkling, 1931: 1; Kryzhanovskij and Reichardt,
1976: 76.

Gardner (1935) gave a detailed description of the genus.
Gardner (1930) and Hayashi (1986) described the larva of Niponius spp.
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. Fore foveae on
propygidium .................................................................... 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. Propygidium without fovea ................................... ..
...... .... ............ ............ ...... ...... ...... ........................ ........ ....... N. obtusiceps Lewis, 1885
4(3)Projection of epistoma long and slender. Propygidium with several excavations.
5(6) Projections of epistoma divergent anteriorly. Surface of elytra wholly with rows of punctures.
Propygidium with 4 foveae. AntelUlal grooves very deep under eyes, and other longitudinal
foveae present on basal 112 of head .............................................. Nfurcatus 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 - 4 foveae. AntelUlal grooves confined
to under eyes, without longitudinal deep foveae on basal 112 of head.................................. ..
......................................................................................... N. osoriocepus Lewis, 1885
Another species, Niponius itoi, has been recorded from Japan.

Niponius Jurcatus Lewi s, 1885
(Fig. 42, 48)
Niponius furcatus Lewis, 1885b: 333 [Japan, Yuyama]; Gardner, 1926: 3; 1935: 5 [catalogued];Kryzhanovskij and Reichardt, 1976: 77 [key]; Hisamatsu, 1985b: 219, pI. 40, fig. 4.
Japanese name: Fusru-tsuno-hoso-emma-musru.

Description. Biometric data are given in Table 5. 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), their apices strongly carinate and dorsal surface with 2 transverse carinae.
Head densely covered with moderate punctures and with several weak rugae on base of
the projection, the punctures becoming coarser on basal 2/3, where they are separated
by about 2.5 (2 - 3) times their diameter.
Pronotum completely with marginal stria on lateral side, the stria well carinate;66

surface densely covered with punctures, the punctures various in size from fine to large(the large punctures are 5 times as large as the fine ones) and becoming finer aroundmargin.
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 115 of elytron. All dorsal striae absent, butthere 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 4 large, longitudinal oblong, shallow andtransversely placed foveae, sometimes these foveae fused; surface sparsely covered
with punctures, which are as coarse as elytral ones and separated by about 2.5 (2 - 3)times their diameter. Pygidium (Fig. 42D) with a large, round and shallow fovea
behind each basal comer; inside of the fovea sparsely covered with coarse punctures;
surface coarsely punctate, the punctures separated by their own diameter and becoming
finer and denser apically and laterally.
Antennal grooves deep under the eyes, and deep and longitudinal foveae on basal
112 of underside of head (Fig. 42B). Mid line strongly impressed on apical 112, but not
impressed near the apical margin of the head. Surface of underside of head evenly
scattered with fine punctures on apical 112; on basal 112 very sparsely clothed with finepunctures, the punctures becoming coarser and denser near the basal margin.

Fig. 42. Niponius furcatus Lewis. A: Head, dorsal view. B: Ditto, ventral view. C: Pro-, meso- andmetasternum, ventral view. D: Propygidium and pygidium, dorsal view. E: Head, lateral view.

67

Prosternal lobe (Fig. 42C) transverse and narrow, its anterior margin denselywith hairs. Prosternal keel narrow, with carinal stria on basal 3/4, the striae deeply
impressed and slightly convergent basally; surface of keel evenly and finely punctate.
Mesosternum longitudinally sulcate on mid line; surface sparsely covered withmoderate punctures which are separated by about 2.5 (2 - 3) times their diameter, andwith large punctures near basal margin. Meso-metasternal suture well impressed andcomplete. 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 andshallow punctures on basal 1/3, the punctures becoming finer and sparser posteriorly.
Intercoxal disk of 1st abdominal sternum striate on basal 2/3 on each lateral side,
and evenly and moderately punctate, the punctures separated by about 2.5 (2 - 3) times
their diameter.
Protibia slender and with 2 denticles on outer margin.
Specimens examined. [Kyfrshu] Kagoshima-ken: 1!f, Kirishima-Jingu, 9/vi/1981, T. & T.
Nakane (NA).
[Nansei Isles] Tolruno-shima Is.: 1!f ,Mikyo, 25/vii/1963, Y. Kurosawa (NA).
[Kochi-ken, Shikoku?]: 1 ex., Wada, Taisho-mura, 4/x/1936, no collecters name, collectedfrom Tsuga sieboldi. (NSMT: Kono collection, No. NSMT-C-23669).

Distribution (Fig. 48). Japan (Shikoku, KyQshQ, Amami-Oshima Is., Tokunoshima Is.).
Remarks. Niponius Jurcatus is characterized by the narrow body, the diverse
projections of the epistoma and the 4 foveae on the propygidium.
Niponius impressicollis Lewis, 1885
(Fig. 43, 44, 48)
Niponius impressicollis Lewis, 1885b: 333 [Japan, Yuyama, in Higo to Junsai in Yezo (= Kyushu to
Hokkaido)]; Gardner, 1926: 2; Reichardt, 1929: 274, 273; Gardner, 1935: 5; Reichardt, 1941:
69; Nakane, 1963: 67, pI. 34, fig. 3; Kryzhanovskij and Reichardt, 1976: 77 [key, description];
Hisamatsu, 1985b: 219, pI. 40, fig. 3.

Japanese name: Hoso-e=a-mushi.
Table 5. Biometric data of Niponius furcatus Lewis and Niponius impressicollis Lewis.
N.furcatus

HOW
HW
PW
PL

EL
EW
ProW
ProL
PyL
PTL
MSTL

MTTL

0.44-0.48
0.73-0.76
0.79-0.85
1.14
1.14-1.24
0.85-0.89
0.50
0.26
0.44-0.48
0.56-0.57
0.48-0.50
0.47-0.50

N. impressicollis(0.45±0.01O) 2(0.75±0.010) 2(0.82±0.021) 2(1.14)2(1.l9±0.036) 2(0.87±0.016) 2
(0.50
)2
(0.26
)2
(0.45±0.01O) 2(0.56±0.005) 2(0.49±0.005) 2(0.48±0.010) 2

68

0.57-0.70 (0.65±0.021) 5
1.09-1.25 (1.19±0.026) 5
1.30-1.56 (1.46±0.042) 5
1.27-1.66 (1.51±0.067) 5
1.61-1.93 (1.77±0.061) 5
1.40-1.66 (1.54±0.051) 5
0.83-0.94 (0.88±0.021) 5
0.57-0.42 (0.48±0.028) 4
0.73-0.57 (0.64±0.023) 5
1.01-0.88 (0.96±0.025) 4
0.83-0.78 (0.79±0.011) 4
0.96-0.78 (0.88±0.033) 4

Fig. 43. Niponius impressicollis Lewis. A: Head, dorsal view. B: Ditto, ventral view. C: Ditto,
frontal view. D: Pronotum, dorsal view. E: Ditto, dorso-oblique view. F: Pro-, meso- andmetasternum, ventral view. G: Propygidium and pygidium, dorsal view.

Description. Biometric data are given in Table 5. Body cylindrical, moderately
stout. Cuticle shining, black; tarsi; antennae and projection of epistoma dark
rufopiceous.
Projections of epistoma (Fig. 43A, C) moderately stout and long, their apices
strongly carinate and dorsal surface with 2 or 3 transverse carinae. Head densely withtransverse rugae on apical 113; median 113 of the surface sparsely covered withmoderate punctures, the punctures separated by about 3.5 (2 - 5) times their diameter,with other fine punctures intermingled; basal 113 finely and sparsely punctate;69

depression on mid line deeply present on apical 112.
Pronotum completely with marginal stria on lateral side, the stria strongly
carinate; surface with an excavation (Fig. 430, E) on each mediolateral side, andsparsely 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 ofwhich the apical end attains to the apical 115; area between epipleural margin and elytral
marginal stria impunctate and extremely finely strigate. First dorsal stria present on
basal 1/4 and strongly carinate, its basal end deeply excavate. Second - 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 2/3.
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 112, and densely on
apical 112, the punctures becoming coarser on apical 1/3, and in lateral area thesepunctures somewhat much denser.
Propygidium (Fig. 430) with 4 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 denselyintermingled among the coarse ones. Pygidium (Fig. 430) with a large, round, andsomewhat deep fovea behind each basal comer; inside of the fovea sparsely andmoderately punctate; surface coarsely punctate on median area, the punctures separated
by 3 - 4 times their diameter and becoming finer anteriorly, and other fine punctures
sparsely intermingled among coarse ones, the fine ones becoming denser apically andlaterally.
Antennal grooves (Fig. 438) deep under eyes. Mid line of underside of head
deeply excavate, its basal end beyond transverse line. Surface of underside of headshining, sparsely and finely punctate on apical 112, and evenly clothed with finepunctures which are separated by about 2.5 (2 - 3) times their diameter on basal 112.
Prosternal lobe (Fig. 43F) transverse and narrow, its anterior margin denselywith hairs. Prosternal keel completely with 2 carinal striae, of which the basal ends areunited with each other in an arch; surface of keel evenly punctate as basal 112 of head.
Mesosternum longitudinally and shallowly sulcate on median 113; lateral 1/3
evenly punctate as basal 112 of head and extremely finely strigate; sometimes lateral
marginal stria present on basal 112 at each anterior corner. Meso-metasternal suture wellimpressed and complete. Metasternum longitudinally and shallowly sulcate on mid line;
lateral metasternal striae strongly carinate and divergent posteriorly. Intercoxal disk ofmetasternum sparsely and coarsely punctate, the punctures twice as coarse as
mesosternal punctures, and other fine ones sparsely intermingled among them. Lateraldisk sparsely covered with large punctures, the punctures becoming finer apically.
Intercoxal disk of 1st abdominal sternum deeply striate on basal 3/4 on eachlateral side, and punctate as intercoxal disk of metasternum, but the punctation is muchdenser.
Protibia slender and with 2 dentides on outer margin.
Male genitalia as shown in Fig. 44.

70

~-------",--1~... .

C2f:.~~.~ ~.-.~ ~.- ~:r:~.i3~.~~.~
B~..

=:::::=... ...-

O.Smm

I

F
Fig_ 44. Niponius impressicollis Lewis A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view.
Specimens examined. [Honshu] Fukushima-ken: 2 exs., Yunohana, Tateiwa V., Minami-Aizu,
19/vi11947, Y. Kurosawa (NA); 4 exs., Onozawa, Egawa, Minami-Aizu, 17/viI1950, Y. Kurosawa(NSMT). Mie-ken: 2 exs., Mimune, l8-211vi/1955, H. Ohira (NA). Hiroshima-ken: 20'0',
Sandankyo, Aki, 7-8/viii/1966, S. Osawa (NA).

Distribution (Fig. 48). Japan (Hokkaido, Honshu, Shikoku, Kyushu); 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 Chujo, 1955
Niponius itoi Chfijo, 1955: 57 [Japan, Adera, Okuwa-mura, Nishichikuma-gun, Nagano pref.,Honshu]; Hisamatsu, 1985b: 220 [=? Niponius osorioceps Lewis].
Japanese name: Ito-hoso-emma-mushi.
Specimens examined. No specimens of the species have been available for my study.

Distribution. Japan (Honshu).
Remarks. Judging from the original description this species is apparently similar
to N. osorioceps, but it is distinguished from the latter by the smaller body size. I havebeen unable to trace it in the Chujo collection. Accoding to Hisamatsu (l985b), thisspecies is probably a synonym of N. osorioceps.
71

Niponius obtusiceps Lewis, 1885
(Fig. 45, 48)
Niponius obtusiceps Lewis, 1885b: 334 [Japan, Oyayama near Kumamoto in Higo, and Ishikari riverin Yezo in 1883]; Gardner, 1926: 3; Reichardt, 1929: 275; Gardner, 1935: 5, t. 1, fig. 2;
Reichardt, 1941: 69; Kryzhanovskij and Reichardt, 1976: 78; Hisamatsu, 1985b: 220 [key].
Japanese name: Tsuno-buto-hoso-emma-mushi.

Description. Body cylindrical, moderately stout. Cuticle shining, black; tarsi,
antennae and projections of epistoma dark rufopiceous.
Projections of epistoma (Fig. 45 A, C) stout and long, their dorsal surface with 2transverse carinae. Apical 113 of head densely covered with fine punctures and weaktransverse rugae, the punctures becoming sparser and coarser posteriorly; median 113 of
the surface coarsely punctate, the punctures being separated by about 1.5 (1 - 2) times
their diameter; basal 113 with much smaller punctures than on median 113, the punctures
being separated by 4 (3 - 5) times their diameter; depression on mid line deep on apical
1/4.
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 ofwhich 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 andtheir punctures becoming finer on apical 113; intervals among the rows impunctate on
basal 2/3, but sometimes sparsely with several fine punctures, which are as large as the
pronotal fine ones, separated by about 1.5 (1 - 2) times the diameter and become denser
and finer posteriorly. Surface of elytra weakly convex on humeral area, and with adeeply 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 2/3 on lateral area; area inside fovea sparsely andfinely punctate; surface between foveae sparsely covered with coarse punctures andintermingled fine ones among the coarse ones; fine punctures present also on apical 113
and along the margin.
Antennal grooves divided into 2 parts; a deep furrow under eyes, and an oblong
fovea on basal 112 of underside of head (Fig. 45B); surface of the groove densely
clothed with fine punctures on apical 112, the punctures much sparser on basal 112.
Prosternal lobe (Fig. 45E) transverse and narrow, its anterior margin densely
furnished with hairs. Prosternal keel completely with 2 carinal striae, of which the basal
ends are united with each other in an arch; surface of keel sparsely clothed with finepunctures.
Mesosternum longitudinally and shallowly sulcate on median 113; area of lateral
113 sparsely clothed with several fine punctures. Meso-metasternal suture absent.
Metasternum longitudinally, broadly and shallowly sulcate on mid line; lateral72

metasternal stria strongly carinate, its apical end extending posteriorly, and attaining tonear apical 114 of metasternum. Intercoxal disk of metasternum sparsely covered withcoarse and longitudinal oblong punctures. Lateral disk densely covered with coarse andround 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, thestriae 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 3 denticles on outer margin.
Specimens examined. [HonshU] Tokyo-to: 1 ex., Mt. Takao, 19/ii/1981, K. Horie. Fukui-ken:
1 ~,Ikegahara, Ono-shi, 20/vi/1988, M. Saito.

Distribution (Fig. 48). Japan (Honshu).
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 collectedunder bark. Hisamatsu (1985b) noted that this species was found in tunnels made bycossonine beetles, Curculionidae.

Fig. 45. Niponius obtusiceps Lewis. A: Head, dorsal view. B: Ditto, ventral view. C: Ditto, lateralview. D: Pronotum, dorsal view. E: Pro-, meso- and metasternum, ventral view. F:
Propygidium and pygidium, dorsal view.

73

Niponius osorioceps Lewis, 1885
(Fig. 46, 47, 48)
Niponius osorioceps Lewis, 1885b: 333 [Japan, Higo, Yuyama and Konose], pI. 8, fig. 12-14;
Gardner, 1935: 3; Nakane, 1963: 67, pI. 34, fig. 4 [noted, photo]; Hisamatsu, 1985b: 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.

Description. Biometric data are given in Table 6. Body cylindrical, moderately
stout. Cuticle shining, black; tarsi, antennae and projections of epistoma dark
rufopiceous.
Projections of epistoma (Fig. 46A, C, D) moderately stout and long, their apices
strongly carinate and dorsal surface with 2 transverse carinae. Apical 1/3 of headdensely covered with transverse rugae; median 1/3 evenly and coarsely punctate, thepunctures being separated by their own diameter; basal 113 densely covered with coarse
punctures, which are a little coarser than the median ones; depression on mid line
present and shallow on apical 113.
Pronotum completely with marginal stria on lateral side, the stria strongly
carinate; surface covered with coarser punctures which are as coarse as those on thebasal 113 of the head, the punctures being separated by about 1.5 (1 - 2) times thediameter.
Epipleura of elytra completely with marginal elytral stria, which is strongly
carinate and of which the apical end attains to near the apical 115; area between
epipleural margin and elytral marginal stria 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 irregularlyintermingled 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 somewhat clear row of coarse and deep punctures. Surface of
elytra with a deep and transverse excavation on near the basal margin.
Propygidium (Fig. 46E) with 2 large, transverse oblong, or 4 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 behindeach basal corner; area inside foveae sparsely covered with moderate punctures; surface
between foveae coarsely punctate, the punctures being separated by about 2.5 (2 - 3)times their diameter, and other moderate punctures sparsely intermingled among thecoarse ones, all these punctures becoming finer and denser apically and laterally.
Antennal grooves (Fig. 46B) deep under eyes. Surface of underside of headsparsely clothed with fine punctures, the punctures separated by about 3 (2 - 4) times
their diameter and becoming denser basally and laterally.
Prosternal lobe (Fig. 46F) transverse and narrow, its anterior margin denselywith hairs. Prosternal keel completely with 2 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
74longitudinally, shallowly and completely sulcate on mid line; lateral metasternal stria
strongly carinate and divergent posteriorly, its apical end extending to near the hindcoxa. Intercoxal disk of metasternum sparsely clothed with fine punctures, thepunctures being separated by about 3.5 (2 - 5) times their diameter, and becoming
denser and a little coarser on apical 114. Lateral disk irregularly scattered with large,
round and deep punctures on basal 113, the punctures becoming finer apically and along
the metasternallateral stria.

Fig. 46. Niponius osorioceps Lewis. A: Head, dorsal view. B: Ditto, ventral view. C: Ditto, lateralview. D: Ditto, frontal view. E: Propygidium and pygidium, dorsal view. F: Pronotum, dorsal
view.

75

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 114 of intercoxal disk of metasternum
and separated by about 1.5 times the diameter; surface among the punctures extremely
finely strigate.
Protibia slender and with 3 denticIes on outer margin.
Male genitalia as shown in Fig. 47.
Specimens examined. [Hokkaid6] 2 exs., Gamushi, 101 vi 1956, A. Nobuchi.

Table 6. Biometric data of Niponius osorioceps Lewis.

HOW

0.58-0.50
0.96-0.75
1.15-0.92
1.29-1.02
1.61-1.20
1.24-0.96
0.75-0.53
0.41-0.26
0.61-0.44
0.73-0.61
0.70-0.53
0.76-0.58

HW
PW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL

MTTL

O.5mm

(0.53±O.006) 18(0.89±0.012) 19(l.04±O.015) 18(1.l5±0.017) 19(1.43±0.024) 19(l.l4±O.019) 18(0.65±0.013) 18(0.35±O.009) 17(0.53±O.013) 17(0.67±0.012) 15(0.64±0.013) 15(0.68±0.012) 17J

Fig. 47. Niponius osorioceps Lewis. A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view.
76[Honshu] Fukushima-ken: 1!f and 7 exs., Yunohana, Tateiwa, Minami-Aizu, 15/vi, 1820/vii/1947, Y. Kurosawa (NA); 2 exs., Ditto, 9/vii/ 1948, T. Nakane (NA); 1 ex., Onozawa, Egawa,
Minami-Aizu, 2/viI1951, Y. Kurosawa (NSMT); 1 ex., Ditto, 17/vi/1950, Y. Kurosawa (NSMT).
Yamagata-ken: 1 ex., Owisawa, 28/vi/1960, Y. Kurosawa (NSMT); 1 ex., Oishi, Oguni, 9/vi/1969, K.
Shirahata (NSMT). Tochigi-ken: 1 ex., Utsunomiya, 6/v/1968, H. Takizawa. Saitama-ken: 1 ex.,
Chichibu, Musashi, 17/vi/1913, H. Takabayashi (NA). Tokyo-to: 2 exs., Hikagesawa, Mt. Takao-san,
4/v/1975, K. Kawada; 1 ex., Ditto, 211vi/1963, S. Nomura (NSMT); 1 ex., Hikawa, Okutama,
13/v/1951, K. Umeya (NSMT). Kanagawa-ken: 11 exs., Mt. Oyama, 5/v/1989, H. Takizawa; I ex.,
Soun-zan, 29/v/1988, H. Takizawa. Nagano-ken: 1 ex., Yatsu-ga-take, 14/vii/1951, 1. Suzuki(NSMT). Shizuoka-ken: I ex., lzu, Inatori, 6/vI1973, H. Takizawa; 1 ex., Mt. Amagi-san, 5/v/1956,
S. Tsuyuki (NSMT). Niigata-ken: 1 ex., Kurokawa, northern Echigo, 9/vi/1963, K. Baba. Gifu-ken: 1ex., Nara-Toge, 4/vii/1953, H. Torigai (NA). Shiga-ken: 1 ex., Bomura, 15/vi/1957, T. Nakane (NA).
Kyoto-fu: 1 ex., Serio, 29/v/1954, T. Nakane (NA). Tottori-ken: I ex., Daisen, 9/vii/1951, N. Yato
(NA).
[Kyushu] Kagoshima-ken: 1 ex., lriki, Aira-gun, IO/V/ 1984, T. & T. Nakane (NA).

Distribution (Fig. 48). Japan (Hokkaid6, Honshu, Kyiishu); Ussuriyskiy Kray.
Remarks. Niponius osorioceps is characterized by the long and slender
projections of the epistoma and the presence of 2 - 4 foveae on the propygidium.
This species is known as a predator of the bark beetle Phloeosinus perlatus
Chapuis, Scolytidae.

Niponius furcatus: •Niponius impressicollis: _Niponius obtusiceps: ...

Niponius osorioceps

Fig. 48. Collection site of Niponius spp. in Japan.

77

6.3. 2. SUBFAMILY CHLAMYDOPSINAE BICKHARDT, 1917
GENUS EuCURITOPSIS SILVESTRI, 1926
Eucuritopsis Silvestri, 1926: 265 [type species, Eucuritopsis mirabilis Silvestri, 1926: 268,
originally designated].
Boreochlamydus K. Sawada, 1994: 357 [type species, Boreochlamydus ohtanii K. Sawada, 1994:359], new synonym.

Description. Characterized by unique shape of pronotum and elytra. Medio-apicalportion of pronotum strongly elevated. Elytral lateral ridge strongly elevated andinterrupted transversely medially; the inside of the interrupted portion furnished denselywith waxed hairs.
Comments. The genus is a small one containing 2 species, E. mirabilis Silvestri,
1926 from Taiwan and E. ohtanii (K. Sawada, 1994) from Japan. Species of thisgenus occur in the nest of ants. E. mirabilis is collected from the nest of Formica legi.
Sawada (1994) described the new genus Boreochlamydus. But he did notcompare it with Eucuritopsis.
Eucuritopsis ohtanii (K. Sawada, 1994).
(Fig. 49)
Eucuritopsis sp.: Nishikawa and Maruyama, 1993: 230 [Zama, Kanagawa, Honshu].
Boreochlamydus ohtanii K. Sawada, 1994: 359 [Mt. Sobo, Oita, Kyushu; Mt. Gozaisyo, Mie,
Honshu].
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=l; female:F, n=I): 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 withcoarse, round punctures. Labrum semicircular. Mandible short and robust. Scape ofantenna large, thick and triangular in frontal view, and coarsely and densely punctate.
Pronotal sides (Fig. 49B) feebly convergent apically, the basal 114 slightly
emarginate to fit fore femur 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 finepunctures 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 apicalend of the sutural stria. Elytral marginal stria absent. Elytron (Fig. 49B) with lateral
margin raised into a narrow, prominent ridge on lateral 113; the ridge narrowlyinterrupted medially; furnished with waxed hairs inside the median interrupted portion;
surface of the basal 112 of the ridge with a narrow elevated carina; mediobasal 112 of the78

Fig. 49. Eucuritopsis ohtanii (K. Sawada). A: Head, forotal view. B: Pronotum and elytra, dorsal
view. C: Propygidium and pygidium.

elytral disk depressed, shining and sparsely furnished with short and thick hairs basally;
apical 112 of the disk convex and evenly covered with coarse, setiferous punctures,
which are separated by about their own diameter, except inside the apical 112 of thelateral elevated ridge and in a longitudinal band narrowly distant from the suture, thepunctured area extending basally narrowly along the suture and reaching to basal 114.
Sutural stria complete and closely impressed along the suture.
Propygidium feebly convex, densely covered with large, round and shallow
punctures which are separated by about 0.6 (0.3 - 1) times the diameter, and sparselyintermingled with fine and setiferous punctures. Pygidium (Fig. 49C) has punctation
simiar to that of the propygidium.
Anterior margin of prosternum feebly sinuate, the median portion slightly andoutwardly arcuate. Disk of prosternum convex medially, densely covered with coarse,
round and setiferous punctures which are separated by 0.3 times their diameter andsparsely 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 about 3 (1 - 5) times their diameter and with short setae. Lateraldisk separated from basal 113 by post-mesocoxal stria; the basal area deeply excavate to
fit mesotibia, impunctate and shining; the apical area has punctation similar to that of theintercoxal disk, but the punctation is much sparser and coarser, except on a triangulararea before latero-apical angle, and with an oblique stria apically which extends inwardly
and posteriorly from the apical 113 of the metasternal-metepisternal suture to the apicalend of the metasternal suture.
Intercoxal disk of 1st abdominal sternum sparsely covered with coarse, roundand setiferous punctures which are separated by about twice their diameter and become
sparser medially, and evenly intermingled with other fine punctures; anterior margin79

sparser medially, and evenly intermingled with other fine punctures; anterior margin
broadly and strongly carinate, the lateral end of the carina extending posteriorly andobliquely, and at apical 113 strongly bent outwardly. Lateral disk separated from basal
2/3 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. Outermargins of meso- and metatibiae strongly angulate at basal 113.
Specimens examined. [Honshu] 18', Sayama-ko, lruma, Saitama-ken, 5/x/1980, T. Syooda,
collected from the nest of PheidoLe ferridoa determined by Dr. Y. Ito (NA). 1 ~, Susono-shi,
Shizuoka-ken, 29/ix/1988, M. Mori, collected with bait trap.
Distribution. Japan (Honshu; Kyushu).
Remarks. Eucuritopsis ohtanii is easily recognizable by the character states givenin the key and description; it cannot be confused with any other Japanese species of thefamily. Nishikawa and Maruyama (1993) recorded a host ant, Phaidole fervida F.
Smith, of this species.
6.3.3. SUBFAMILY TRIBAllNAE 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; Ganglbauer, 1899: 370; Jakobson, 1911: 640, 646;
Bickhardt, 1917: 122; Arnett, 1962: 376, 381; Witzgall, 1971: 180; Kryzhanovskij andReichardt, 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.
Specimens examined. No specimens of the species have been available for my study.
Distribution. Japan (Honshu).

Epierus uenoi M. Ohara, 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, blackand shining; femora, tibiae, tarsi, antennae and mouthparts castaneous.
Frontal stria (Fig. SOB) absent anteriorly, but rudimentarily present laterally on
basal 113; surface of head depressed medially and elevated obliquely above antennal
cavity; punctation of surface fine and evenly separated by about twice their diameter.
Labrum with 2 long setae. Club of antenna without segmentation, its apical margin
truncate.
Pronotal sides (Fig. 50A) strongly convergent apically; marginal stria complete
laterally, but entirely absent anteriorly; disk finely punctate, the punctures being
separated by about 3 (2 - 4) times their diameter and becoming a little coarser laterally;80basal margin of pronotum obtusely angulate at middle; broadly even medially.
Epipleural marginal stria complete, sparsely crenate, feebly carinate, and runninga little distant from the margin. Narrow band between the margin and epipleuralmarginal stria with fine punctures in a row. Elytral marginal stria deeply impressed on
apical 112 and sparsely crenate. Epipleura sparsely and somewhat coarsely punctate andeven. Subhumeral stria absent. Oblique humeral stria finely impressed on basal 114.
First dorsal stria deeply impressed on basal 2/3, sparsely and coarsely crenate. Secondand 3rd dorsal striae present on basal 3/4, 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 116.
Sutural stria shortly present on median 113 and lightly impressed. Interstices among
dorsal elytral striae sparsely clothed with fine punctures, which are separated by about
4.S (4 - S) times their diameter; on median 113 and apical liS the punctures becomingcoarser and denser and being separated by about 2.5 (2 - 3) times their diameter except
on a narrow sutural band, which is densely and finely punctate, and an extreme apicalband, which is impunctate.
Propygidium (Fig. SOC) with coarse punctures which are separated by about I.S
times the diameter and become finer around margin. Pygidium evenly covered withcoarse and round punctures which are separated by about their own diameter.
Anterior margin of prosternal lobe (Fig. SOD) nearly straight medially, its
marginal stria clearly impressed and carinate; apical 116 of the lobe transversely
elevated; disk sparsely clothed with fine punctures. Prosternal keel feebly convex on
apical 112 and even on basal 112; 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 marginof the keel broadlyemarginate.

Fig. 50. Epierus uenoi M. Ohara, sp. nov. A: Pronotum and left elytron. B: Head, frontal view. C:
Propygidium and pygidium. D: Prostemum, mesosternum, metasternum and 1st abdominal
sternum. E: Protibia, dorsal view. F: Ditto, ventral view.

81Anterior margin of mesosternum (Fig. 50D) round, its marginal stria nearlycomplete laterally and broadly interrupted medially on anterior margin; disk sparsely
clothed with fine punctures which are a little finer than the pro notal ones and separated
by about 2.5 (2 - 3) times their diameter. Meso-metasternal suture strongly andsparsely crenate, carinate and arcuate anteriorly. Punctation of intercoxal disk ofmetasternum similar to that of the mesosternum, the punctures becoming coarser andsparser laterally. Mid line of the disk weakly elevated on median 115. Lateralmetasternal stria extending obliquely and posteriorly on basal 2/3. Lateral disk ofmetasternum covered with large and round punctures which are separated by 0.75 times
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 about 5 (4 - 6) times their diameter; 1st abdominal stria
complete on each lateral side.
Protibia (Fig. 50E, F) slender and multidentate (with 11 small denticles on outermargin).
Male genitalia as shown in Fig. 5l.
Specimens examined. Holotype, d", Tsunagu, Amami -Oshima Is., Nansei Isles, 2/iiil 1989, T.
Ueno (EIHU).

Distribution. Japan (Nansei Isles: Amami-Oshima Is.).
Etymology. This species is named in honor of Mr. Teruhisa Ueno, who gave me
the opportunity to study the specimen.
Remarks. Epierus uenoi can easily be distinguished from E. lucus by thestriation of the elytra, and by its limited distribution (Amami-Oshima, Nansei Isles).

\~~- --- ----------.,.,,~~~~

B

~

E~
/,j!Y; F

0.25 mm I.y."':

Fig. 51. Epierus uenoi M. Ohara, sp. nov. A: Aedeagus, dorsal view. B: Ditto, lateral view. C: MaIe
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitaIia, 9th and10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view.

826.3.4. SUBFAMILY ONlHOPHIUNAE THOMSON, 1862
Key to the Japanese genus of the subfamily OnthophiJinae
1(2)
2(1)

Dorsal surface without hair .............................................. Genus Onthophilus Leach, 1817
Dorsal surface with short and stout hairs .............................. Genus Epiechinus Lewis, 1891
GENUS ONTIIOPHILUS LEACH, 1817

Onthophilus Leach, 1817: 77 [type species: Hister striatus Forster, 1771]; 6hara and Nakane, 1986:
3.
Scolytus O. Muller, 1776: 22 [nee 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 by 6hara and Nakane(1986). Some specimens examined thereafter are added below.
Key to the Japanese species of the genus Onthophilus.
1( 2)
2( 1)
3( 8)
4( 5)

Pronotum with 8 costae. Body length 2.50 - 2.65 mm .................... O. silvae Lewis, 1884
Pronotum with 4 or 6 costae.
EC3 (elytral costa 3) ending just caudad of a deep transverse fossa situated within front marginof elytron.
PC3 (pronotal costa 3) present on apical and basal 112 of pronotum, and strongly developed.

5( 4)Body length, 3.60 - 4.68 mm .................................................... O. ostreatus Lewis, 1879
~ absent, or present only on basal 112 of pronotum.

6( 7)~ present. Body length, 2.50 - 3.02 mm ...............................

7( 6)~ absent. Body length, 3.28 - 3.38 mm .................... O. aonoi M. 6hara et Nakane, 1986

O. niponensis Lewis, 1907

8( 3)EC3 entire upto front margin of elytron.
9(10) Body longer than 2.5 (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: 6hara and Nakane, 1986: 5.

Japanese name: Shinano-sesuji -emma-mushi.
Specimens examined [additional records]. [Hokkaido] 2 exs., Yahata, Kutchan, 20, 30/viii/1993,
M. 6hara, collected near the nest of ant, Lasius (Dendrolasius) fulginosus.
[Honshu] Kanagawa-ken: 1 ex., Daiyfi-zan, Hakone, 14/viii/1982, Y. Hirano (NSMT).

Distribution. Japan (Hokkaid6, Honshu).
Onthophilus aonoi M. 6hara et Nakane, 1986
Onthophilus aonoi 6hara and Nakane, 1986: 6; Hirano, 1988: 41 [Kanagawa, HonshU].

Japanese name: Nise-ko-sesuji-emma-mushi.

Distribution. Japan (Honshu).
83

Onthophilus ostreatus Lewis, 1879
Onthophilus ostreatus: Ohara and Nakane, 1986: 5.

Japanese name: O-sesuji-e=a-mushi.
Specimens examined [additional records]. [Honshu] Ibaraki-ken: 1 ex., Tadobe, Sakuragawa,Niiharu-gun, 3/xi/1984, K. Haga. Tokyo-to: 1 ex., Nakano, 20/xi/1923, no collector's name (NSMT);
1 ex., Tokyo, no data and no collector's name (NSMT); 5 exs., Tokyo, 25/x/1936, T. Adachi (NSMT).
Kanagawa-ken: 6 exs., Mizonokuchi, no date and no collector's name (NSMT). Hyogo-ken: 1 ex.,
Harada, Kobe, 19/xi/1914, no collector's name (BSM).

Distribution. Japan (Honshu, Shikoku, KyOshu); Continental China; Taiwan.
Onthophilus niponensis Lewis, 1907
Onthophilus niponensis: Ohara and Nakane, 1986: 6.

Japanese name: Ko-sesuji-e=a-mushi.
Specimens examined [additional records]. [Honshu] Fukushima-ken: 6 exs., Wakamatsu,
19/iv/1947, Y. Kurosawa (NSMT). Tokyo-to: 1 ex., Komazawa, 4/vii/1931, K. Seiki (NSMT). Gifuken: 1 ex., Nannou, ll/iv/1981, T. Nohira (refer to Takai, 1989).

Distribution. Japan (Honshu, Kyushu).
Onthophilus ordinarius Lewis, 1879
Onthophilus ordinarius: Ohara and Nakane, 1986: 9.

Japanese name: Ezo-sesuji-e=a-mushi.
Specimens examined [additional records]. [Hokkaido] 10 exs., Nopporo, 5/vi/1987, M. Ohara; 5exs., Ditto, 12/vi/1987, M. Ohara; 1 ex., Ditto, 18/vii/1987, M. Ohara; 2 exs., Sapporo, 10/vi/1940,
Y. Nishijima (NSMT).

Distribution. Japan (Hokkaid6); Russia (Vladivostok, Baikal Sea, Novosibirsk,
Irkutsk, Ussuri).
Onthophilus flavicornis Lewis, 1884
Onthophilusj1avicornis: Ohara and Nakane, 1986: 9.

Japanese name: Kinoko-sesuji-e=a-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Sapporo, 26/vi/1939, Y. Nishijima(NSMT).
[Honshu] Ibaraki-ken: 1 ex., Kamisaki, Sakura-mura, 4/v/1985, K. Haga. Tochigi-ken: 1 ex.,
Nikko, ll/v/1985, M. Kiuchi. Saitama-ken: 1 ex., 20/iv/1968, H. Takizawa. Tokyo-to: 1 ex.,
Kakinoki-zaka, Meguro, 5/iv/1947, H. Watanabe (NSMT). Nagano-ken: 4 exs., Jigoku-dani,
Yamanouchi, 29/iv/1983, M. Kiuchi; 1 ex., Shiga-Kogen, 7/v/1986, H. Tanaka.

Distribution. Japan (Hokkaid6, Honshu, Shikoku, Kyushu).

84

Ontlwphilus kamiyai Adachi, 1930
Onthophilus kamiyai: Ohara and Nakane, 1986: 9.

Japanese name: Nise-kinoko-sesuji-enuna-mushi.

Distribution. Japan (HonshU).
GENUS EPIECHINUS LEWIS, 1891
Epiechinus Lewis, 1891: 320 [type species: Onthophilus costipennis Fahreaus in Boheman, 1851:
549; originally designated]; 1892c: 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: Honshu: Nara].
Epiechinus arboreus: Lewis, 1891: 320; Hisamatsu, 1985b: 226, pI. 4, f. l.
Scolytus arboreus: Jakobson, 1911: 652.

Japanese name: Chibi-ke-sesuji-enuna-mushi.

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 7.
Frontal lateral sides (Fig. 52D) convergent apically on basal 112 and strongly
emarginate behind antennal sockets; anterior margin roundly arcuate. Margin of thefrontal disk strongly carinate. Disk with 5 longitudinal costae; the median costa present
on basal 112; the lateral costae convergent apically through behind the antennal sockets
and united with each other anteriorly; 5 crossing costae present anteriorly between thelateral costa and the anterior marginal carina; the mediolateral costae shortly present on
basal 114. Surface of the disk sparsely furnished with minute spines.
Pronotal sides (Fig. 52A) feebly convergent forward on basal 3/4, thence
strongly convergent apically, and strongly carinate and densely furnished with minutespines. Anterior margin of pronotum broadly emarginate, the median portion nearlystraight. Disk with 6 costae (Fig. 52E); the median 4 costae present on apical 113 andfeebly elevated; the lateral costae strongly elevated, complete and interrupted at apical
114; sides of these costae furnished with minute spines. Surface of the disk sparsely
covered with coarse and round punctures which are separated by about 1.5 (1 - 3)times the diameter and have minute setae in a trapezoid area on the mediobasal 2/3; otherarea, 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 112; surface with 2 punctured rows,
the punctures quadrate, large and shallow, the inner row complete and the outer(running close to the margin) shortly present on media-apical 114. Disk of elytron (Fig.
52A) with 5 spinal costae (excluding the costa on elytral margin); the 1st - 4th costae(counted from outside to inside) strongly elevated, complete and furnished with minute85

Fig. 52. Epiechinus arboreus (Lewis). A: Adult, dorsal view. B: Ditto, covered with mudlike scales,
dorsal view. C: Ditto, ventral view. D: Head, frontal view. E: PronotuID, dorsal view. F: Left
elytron, dorsal view.

86spines 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 2 punctured rows, the punctures round, large, deep,and 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 amongcoarse ones; large, round and shallow punctures present along the anterior margin.
Pygidium (Fig. 530) 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 atransverse punctured row on apical 115, 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 striae strongly carinate and divergent apically.

Fig. 53. Epiechinus arboreus (Lewis). A: Prosternum and mesosternum, ventral view. B: Fovea ofmesosternum. C: Intercoxal disk of 1st abdominal sternum, ventral view. D: Pygidium, caudal
view.

87

F

G

0.25 mm!

E

Fig. 54. Epiechinus arboreus (Lewis). A: Aedeagus, dorsal view. B: Ditto, lateral view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9thsternum (spicule), dorsal view. F: Male genitalia, 9th and 10th tergite, dorsal view. F: Ditto,
lateral view.
Table 7. Biometric data of Epiechinus arboreus (Lewis).

APW
PPW
PL
EL

EW
ProW
ProL
PyL
PTL
MSTL

MTIL

0.55-0.67
1.06-1.30
0.59-0.71
0.95-1.18
1.36-1.62
0.51-0.67
0.16-0.26
0.39-0.47
0.47-0.63
0.37-0.53
0.39-0.57(0.62±0.01) 7(1.21±0.03) 7(0.65±0.02) 7(l.l0±0.03) 7(1.51±0.03) 7(0.60±0.02) 7(0.22±0.01) 7(0.44±0.01) 7(0.56±0.02) 7(0.49±0.02) 7(0.53±0.02) 7

Anterior margin of mesosternum (Fig. 53A) sinuate, the median portion
produced to fit the prostemal emargination; disk short and deeply excavate laterally(Fig. 538); posterior margin angulate obtusely at middle; surface sparsely covered withfine spiral punctures, deeply excavate laterobasally and at apical 1/3 on lateral side, thelaterobasal excavation becoming broader posteriorly and inwardly. Lateral metasternal
stria carinate, extending posteriorly obliquely and angulate outwardly at basal 115.
88

Lateral disk with 2 deep excavations (Fig. 53C), the basal one present behind themesocoxa and the apical one transverse and present on apical 112; remains of the basal
112 covered with large round and shallow punctures.
Intercoxal disk of 1st abdominal sternum (Fig. 53C) short, with 2 excavations;the basal one deeply present on each lateral area, and the apical one shallowly on eachlatero-apical angle.
Protibia narrow, its outer margin with 10 setae and angulate medially; interspace
between apical 2nd and 3rd setae broad. Mesotibia with 12 setae on outer margin.
Metatibia with 13 setae on outer margin.
Male genitalia as shown in Fig. 54.
Specimens examined. [HonshU] Nara-ken: 1 ex., Kasuga-yama, 8/i/1954-, M. Otake (NA); 3
exs., Ditto, 19/vi, 7, 14/vii/1957, T. Shibata (NA); 2 exs., Ditto, 13/vi/1958, T. Nakane (NA); 1 ex.,
18/iv/1965, M. Gota (NA); 8 exs., Ditto, 26/xi/1958, T. Shibata (NSMT); 1 ex., Nara, 28/iii/1959,
no collector's name (NSMT).

Distribution. Japan (Honshu).
Remarks. Epiechinus arboreus is easily distinguished from other Japanese
species of the subfamily Onthophilinae by its small size and the presence of minutespines on the dorsal surface in combination with the distinctly developed elytral costae.
Little is known about the habitat of this species. Lewis (1884) noted that thisspecies was residing in galleries of wood-borers, probably Tomicus.
6.3. 5. SUBFAMILY HISTERINAE GYLLENHAL, 1808

Key to the Japanese tribes and genera of the subfamily Histerinae
1(6)
2(3)

Tarsal groove of protibia S-shaped.
Head porrect, horizontal in repose .................................... Tribe Hololeptini Lacordaire, 1854.................................................................................... Genus Hololepta Paykull, 1811
3(2) Head vertical in repose ................................................ TribePlatysomatini Bickhardt, 1917
4(5)Body oblong-oval. .. ........................................................ Genus Platysoma Leach, 1817
5(4)Body oval.
.. ..................................................................... Genus Eblisia Lewis, 1889
6(1) Tarsal groove of protibia straight.
7(10) Anterior margin of mesosternum bisinuate with a more or less distinct median projection whichfits into basal margin of prostemurn ................................. Tribe Exosternini Bickhardt, 1917
8(9) Body red. ................................................................... Genus Notodoma Lacordaire, 18549(8) Body darkish bronze. ....................................................... Genus Binhister Cooman, 1934
10(7) Anterior margin of mesosternum straight or truncate or emarginate, not bisinuate. .. .............. ..
.......... ............ ................ ...... ...................... ................... Tribe Histerini Gyllenhal, 1808
11(18) Anterior margin of mesosternum truncate or emargiuate at middle (in Hister simplicisternus,
the anterior margin straight; but simplicisternus is detected by having 2 pronotal lateral striae
and by the short and deeply impressed external subhumeral stria on the eIytron).
12(17) External subhumeral stria of eIytron not completed.
13(16) Pronotum with 2 lateral striae.
14(15) Pronotum and elytra without large and deep punctures. .............. Genus Hister Linnaeus, 1758
15(14) Pronotum and elytra densely covered with large and deep punctures ..................................... ..
................................................................................ Genus Zabromorphus Lewis, 1906
16(13) Pronotum with 11ateral stria............................................. Genus Merohister Reitter, 1908
17(12) External subhumeral stria of e1ytron complete ................. Genus Margarinotus Marseul, 1853
18(11) Anterior margin of mesosternum straight or feebly arcuate outwardly, sometimes feebly
emarginate at middle, but marginal stria of mesosternum not emarginate............................. ..
.................................................................................... Genus Atholus Thomson, 1859

89

TRIBE EXOSTERNINI BICKHARDT, 1917

This tribe has already been revised by Ohara and Nakane (1989). I add somespecimens examined thereafter.
GENUS BINHISTER COOMAN, 1934
Binhister Cooman, 1934: 122 [type species: Binhister barbarus Cooman,designated]; Mazur, 1984: 288; Ohara and Nakane, 1989: 290.

1934, originallyBinhister chujoi Cooman, 1941
Binhister chujoi Cooman, 1941: 332; Ohara and Nakane, 1989: 290.

Japanese name: Chujo-chibi-emma-mushi.
Specimens examined [additional records]. [Honshu] Tokyo-to: 3 exs., Mt. Takao, 211vi/1963,
S. Nomura (NSMT). Nagano-ken: 1 ex., Nojiri, Gaijin-mura, 25/viii/1941, T. Nakane (refer to Osawa
and Nakane (1951». Kyoto-fu: 1 ex., Kyoto University, 3/vii/1951, F. Takahashi (NA); 1 ex., Mt.
Daihi, 30/v/1959, Y. Hama (NSMT).

Distribution. Japan (Honshu).
GENUS NOTODOMA LACORDAlRE, 1854
Notodoma Lacordaire, 1854: 266; Ohara and Nakane, 1989: 285.

NotodomaJungorum Lewis, 1884
Notodomafungorum Lewis, 1884: 136; Ohara and Nakane, 1989: 286.

Japanese name: Kinoko-aka-maru-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Sapporo, 7/vii/1955, T. Kumata(ElHU); 1 ex., Maruyama, Sapporo, 291v/1991, K. Miyashita; 8 exs., Toya-Naka-no-shima, Abuta,
17/vii/1993, S. Hori (HOR).
[Honshu] Iwate-ken: 3 exs., Matsuo, 17-19Ivii/1954, S. Takagi (EIHU). Fukushima-ken: 3exs., Yunohana, Tateiwa, Minami-Aizu, 211vi/1947, 7/vii/1948, Y. Kurosawa (NSMT); 1 ex.,
Yunohana, Ouchi, Narahara, Minami-Aizu, 13/vi/1948, Y. Kurosawa (NSMT). Tokyo-to: 24 exs.,
Takao, 29/v/1968, 3/vii/1969, H. Takizawa; 2 exs., Ditto, 21/vi/1963, S. Nomura (NSMT); 1 ex.,
Mt. Kuwanori, 28/v/1968, H. Takizawa; 2 exs., Nippara, 26/vii/1966, Y. Kurosawa (NSMT).
Yamanashi-ken: 1 ex., Torii-toge, 101vii/1938, Imazaki (NSMT). Nagano-ken: 2 exs., Kutsukake,
Shinano, 7/viii/1941, T. Niimura (NSMT); 1 ex., Amori, Nagano-shi, 24/vii/1979, T. Fujisawa.
Hyogo-ken: 2 exs., Kirihata, Takarazuka, 7/viIl992, T. Ito. Tottori-ken: 2 exs., Mt. Daisen,
14/vii/1947, H. Kono (NSMf).
[Kyushu] Tsushima Is.: 3 exs., Meboro, Mt. Mitake, 15-18/vii/1968, S. Miyamoto and A.
Nakanishi (NSMT); 2 exs., Mine, 101 vii 1969, T. Nakane (NSMT).
[Nansei Isles] Yaku-shima Is.: 8 exs., Yaku-Sugi land, 3/vii/1975, Y. Kurosawa (NSMT); 2
exs., Shiratani, Yaku-shima, 201vi/1975, Y. Kurosawa (NSMT); 1 ex., Ambo, 211vi/1975, Y.
Kurosawa (NSMT).

Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu, Izu Isles, Tsushima
Isles, Nansei Isles); Taiwan.

90

TRIBE HOLOLEPTINI LACORDAIRE, 1854
GENUS HOLOLEPfA PAYKULL, 1811
Hololepta Paykull, 1811: 101; Ohara, 1991a: 102.

The Japanese species of the genus HoLoLepta have already been revised by Ohara(1991a, b). I add some specimens examined thereafter.
SUBGENUS HOLOLEPTA PAYKULL, 1811
Key to the Japanese species of the subgenus HoLoLepta
1(4)
2(3)
3(2)
4(1)

Propygidium without stria.
Anterior margin of prostemal lobe outwardly arcuate. Body (head excluded) larger, 7.3 - 8.7
mm. Large fovea present on pronotal anterior angle in male .......... H. amurensis Reitter, 1879
Anterior margin of prostemallobe feebly emarginate. Pronotum without foveae. Body smaller,
5.7 -7.0 mm.
. ................................................................ H. depressa Lewis, 1884
Propygidium with a stria on each side .......................................... H. higoniae Lewis, 1894

HoLoLepta (HoLoLepta) amurensis Reitter, 1897
Hololepta amurensis Reitter, 1879: 213.
Hololepta (Hololepta) amurensis: Ohara, 1991a: 103.

Japanese name: O-hirata-emma-mushi.
Specimens examined [additional records]. (lIokkaid6] 8 exs., Mt. Y6tei, Hirafu, Kutchan, 15-16,
22/vii/1991, M. Ohara; 1 ex., Asahi-Tenb6dai, Otaru, 211v/1991, M. Ohara.

Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu, Izu Isles, Tsushima
Isles, Yaku-shima Is.); East Siberia; Primorskij Kray; Korea; Taiwan; Continental
China.

HoLoLepta (HoLoLepta) depressa Lewis, 1884
Hololepta depressa Lewis, 1884: 132.
Hololepta (Hololepta)depressa: Ohara, 1991b: 235.

Japanese name: Hirata-emma-mushi.
Specimens examined [additional records]. (lIokkaid6] 4 exs., Mt. Y6tei, Hirafu, Kutchan, 15-16,
22/vii/1991, 26/v/1992, M. Ohara.

(lIonshfi] Tochigi-ken: 1 ex., Aoyagi, Kuriyama, 16/vi/1979, S. Takahashi (TPM); 1 ex.,
Yunishi-Kawahira-Sawa..Kuriyama, 7/viil1991 , K. Suzuki (TPM).

Distribution. Japan (Hokkaido, Honshu, Shikoku, KyushG); Taiwan.

HoLoLepta (HoLoLepta) higoniae Lewis, 1894
Hololepta parallela Lewis, 1884: 132 (nec G. Koch, 1868).
Hololepta higoniae Lewis, 1894: 174; Miwa, 1938: 84,85; Kurosawa, 1952; 24 (lIonshfiJ
Hololepta (Hololepta) higoniae: Ohara, 1991b: 238; Kurosawa, 1992: 7.

91

Japanese name: Higo-hirata-emma-mushi.

Distribution. Japan (Honshu, KyOshu); Laos; Viet-Nam; Taiwan; Continental
China (Yunnan).
TRIBEPLA1YSOMATINI BICKHARDT, 1917
GENUS PLATY SOMA LEACH, 1817
Platysoma Leach, 1817: 77 [type species: Hister depressus Fabricius, 1787: 32, designated byWestwood, 1840: 22].
Abbotia Leach, 1830: 155, synonymized by Waterhouse, 1868: 168. Without type species.

Part of this genus, including subgenus Platysoma, has already been revised in
Ohara (1986). I renew the key to the Japanese species and add some specimens to thespecies treated at that time.
Key to the Japanese subgenera of the genus Platysoma
1(2)
2(1)
3(4)
4(3)
5(6)
6(5)

Body strongly depressed. Interspace between coxae broad, its width the same as length ofmetatibia.................................................................. Subgenus Apobletes Marseul, 1860
Body moderately convex. Interspace between coxae not broad, its width the same as 113 lengthof metatibia.
Body slender, and narrower than length of sutural line of elytra. . ........................................ .
................................................................................. Subgenus Cylister Cooman, 1941
Body oval or oblong-oval. Body as wide as length of sutural line of elytra.
Margin of pygidium strongly carinate. ............................. Subgenus Platylister Lewis, 1892
Margin of pygidium not carinate ..................................... Subgenus Plutysumu Leach, 1817
SUBGENUS PLATY SOMA LEACH, 1817

Key to the Japanese species of the subgenus Platysoma
1(4)
2(3)

Pronotum smooth, impunctate (or at most microscopically punctulate).
Elytra rugulose on apical 113; dorsal striae 1-3 complete, 4th abbreviated on basal 1/3, 5th andsutural present on apical 112; pygidium with a deep fovea on each side at base; body length 3.3- 3.65 mm. .............................................................
P. (P.) tsushimae M. Ohara, 1986
3(2) Elytra not rugulose; dorsal striae 1 - 3 complete, 4th apical, 5th abbreviated on basal 1/3,
sutural stria wanting; pygidium without fovea; body length 3.05 -3.8 mm .......................... .
................................................................... ............... P. (P.) unicum Bickhardt, 1912
4(1) Pronotum punctate, especially densely on lateral area.
5(8) Body length 4.2 - 5.3 mm; prostemal process with carinal striae; lateral pronotal stria rather
distant from the side margin.
6(7)Sutural stria on basal 112, consisting of moderately coarse punctures; dorsal striae 1 - 4complete, 5th obsolescent at base; body length 4.2 - 5.3 mm ............................................ .
................................................................... ................... P. (P.) lewisi Marseul, 1873
7(6)Sutural stria absent; dorsal striae 1 - 3 complete, 4th and 5th found on apical 112; body length
5.0 mm. ........................................................................... P. (P.) vagans Lewis, 1884
8(5)Body length 2.13 - 3.8 mm; prostemal process without carinal striae; lateral pronotal stria
rather close to the margin.
9(12) Dorsal striae 1 - 4 complete (in P. celatum Lew. rarely abbreviated on the base).
10(11) Pygidial punctures almost as large as the coarse propygidial ones; metatibia without a spine, orwith very fine spinules on outer margin; lateral pronotal stria complete; body length 2.13 2.95 mm. ..................................................... ................... P. (P.) celatum Lewis, 188492

11(10) Pygidium more closely punctate than propygidium; metatibia always with a large spine onouter margin at apical 2/3; lateral pronotal stria narrowly interrupted behind each eye; body
P. (P.) deplanatum (Gyllenhal, 1808)
length 2.9 - 3.4 mm. ............................................
12(9) Dorsal striae 1 - 3 complete.
13(14) Sutural stria is present on apical area; lateral pronotal stria narrowly interrupted behind each eye;P. (P.) takehikoi M. Ohara, 1986
body length 3.25 - 3.8 mm. ............................. ...........
14(13) Sutural stria is absent; lateral pronotal stria complete; body length 2.4 - 3.1 mm .................. .
P. (P.) rasile Lewis, 1884.. ........ .......... ...... ...... ............ .......... ........ ...... ............ .........

Platysoma (Platysoma) celatum Lewis, 1884
Platysoma celatum Lewis, 1884: 134; Ohara, 1986: 99.
Japanese name: Hime-naga-emma-mushi.
Specimens examined [additional records]. [HonshU] Yamagata-ken: 1 ex., Tama-gawa, 7/v/1975,
Y. Kurosawa (NSMT). Fukushima-ken: 1 ex., Yunohana, Tateiwa, Minami-Aizu, 211vii/1947, Y.
Kurosawa (NSMT); 1 ex., Mt. Senaka-Aburi, Higashi-yama, Kita-Aizu, 13/v/1951 , Y. Kurosawa(NSMT); 1 ex., Suimon, Asahida, Minami-Aizu, 4/vi11949, Y. Kurosawa (NSMT); 1 ex., Jabami,
Nagaino, Onuma, 25/vi/1949, K. Nagayama (NSMT). Tochigi-ken: 1 ex., Yokokawa, Fujiwara-ch6,
24/ix/1989, H. Yoshimoto (TPM); 1 ex., Ohira-yama, 27/v/1989, H. Yoshitomi (TPM); 3 exs., Kajiga-sawa, Ujiie-cho, 27/iv/1988, K. Satoh (TPM); 1 ex., Tsuruta-Numa, Utsunomiya, ll/vi/1991, K.
Satoh (TPM). Niigata-ken: 1 ex., Echigo-Koma-ga-take, 13-14/ix/1970, H. Takizawa. Nagano-ken: 2exs., Mt. Yatsu-ga-take, 4/x/1941, M. Hanao (NSMT). Okayama-ken: 1 ex., Mimasaka, vii/1912, J.
E. A. Lewis (BSM).
[Kyushu] Fukuoka-ken: 2 exs., Mt. Shoji, 28/ivI1964, 16/vi/1964, S. Nakao (NA).

Distribution. Japan (Hokkaido, Shikoku, Kyushu); Continental China.
Platysoma (Platysoma) deplanatum (Gyllenhal, 1808)
Hister deplanatum Gyllenhal, 1808: 85; Lewis, 1906a: 340.
Platysoma deplanatum: Ohara, 1986: 100.
Japanese name: Kita-naga-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Rubeshibe, 30/v/1980, H. Torikura;
1 ex., Oku-Shibetsu, Kamikawa, Teshio, 29/vi/1949, T. Hasegawa (NSMT); 4 exs., Raibetsu,
Shibecha, Kushiro, 6/vii/1972, 12/vii/1973, 151vii11975, K. Ijima (lJ); 3 exs., Futatsu-yama,
Shibecha, Kushiro, 12/vi/1956, 24/vi/1973, 12/vi/1975, K. Ijima (11); 1 ex., Rururan, Sugano,
Shibecha, Kushiro, 24/viil 1992, K. Ijima (11); 1 ex., Kami-Ashoro, Ashoro-cho, 4/viil 1993, K. Ito.

Distribution. Japan (Hokkaid6); Europe; Siberia; Mongolia; Korea.
Platysoma (Platysoma) lewisi Marseul, 1873
Platysoma lewisi Marseul, 1873: 222; Ohara, 1986: 97.
Japanese name: O-naga-emma-mushi.
Specimens examined [additional records]. [Honshu] Osaka-fu: 2 exs., Minoo, 151vi/1958, Y.
Hama (NSMT). Hyogo-ken: 1 ex., Kobe, vi/1912, J. E. A. Lewis (BSM).
[KyfishU] Fukuoka-ken: 1 ex., Mt. Shoji, 13/vii/1963, S. Nakao (NA). Kagoshima-ken: 2 exs.,
Takachiho-gawara, Kirishima, 5/vi/1985, Y. Takai.

93

Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu, Tsushima Isles);Continental China.
Platysoma (Platysoma) rasile Lewis, 1884
Platysoma rasile Lewis, 1884: 134; Ohara, 1986: 104.

Japanese name: Nise-hime-naga-ennna-mushi.
Specimens examined [additional records]. [Honshu] Tochigi-ken: 1 ex., o chi ai , Kaminagamachi, Kurino-cho, 10/vii/1991, K. Satoh (TPM). Chiba-ken: 1 ex., Mt. Kiyosumi, 24/v/1976, M.
Tomokuni (NSMT). Tokyo-to: 1 ex., Aihara, 16/v/1987, H. Takizawa; 1 ex., Kinuta, Setagaya,
30/iv/1967, Y. Kurosawa(NSMT).
[Kyfishfi] Fukuoka-ken: 1 ex., Mt. Shoji, 30/v/1971, S. Nakao (NA).

Distribution. Japan (Honshu, Shikoku, Kyushu, Tsushima Isles, Nansei Isles).
Platysoma (Platysoma) takehikoi M. Ohara, 1986
Platysoma takehikoi M. Ohara, 1986: 102.

Japanese name: Nakane-naga-ennna-mushi.
Specimens examined [additional records]. [Nansei Isles] Ishigaki Is.: 1 ex., Banna-dake, Ishigaki,
311x/1974, K. Tazoe. Iriomote Is.: 1 ex., Ohara, 12/iii/1990, Y. Okushima.

Distribution. Japan (Nansei Isles).
Platysoma (Platysoma) tsushimae M. Ohara, 1986
Platysoma tsushimae M. Ohara, 1986: 94.

Japanese name: Tsushima-naga-ennna-mushi.

Distribution. Japan (Tsushima Isles).
Platysoma (Platysoma) vagans Lewis, 1884
Platysoma vagans Lewis, 1884: 133; Ohara, 1986: 97.

Japanese name: Heri-naga-ennna-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., 13-no-sawa, Mitsumata, KamiShihoro, 23/viii/1992, K. Miyashita; 1 ex., Nopporo, Ebetsu, 19/vi/1987, M. Ohara.
[Shikoku] Kochi-ken: 1 ex., Tebako, 2S/viil 1962, D. Motomura (NSMT).

Distribution. Japan (Hokkaido, Honshil).
Platysoma (Platysoma) unicum Bickhardt, 1912
Platysoma unicum Bickhardt, 1912: 124; Ohara, 1986: 96.

Japanese name: Tokara-naga-ennna-mushi.
94

Distribution. Japan (Nansei Isles); Taiwan.
SUBGENUS PLATYLISTER LEWIS, 1892
Platylister Lewis, 1892d: 103 [type species: Platysoma ovatum Erichson, 1834: 108, originallydesignated].

Key to the Japanese species of the subgenus Platylisler
1(4)
2(3)
3(2)
4(1)
5(6)
6(5)

Front of head strongly concave.
First - 4th dorsal elytral striae complete. Body small, 4.00 - 4.75 rum ................................ .
...... ...... .... .... ........ .... ........ .... ...... .... ...... ........ ...... .......... ....... P. (P.) pini Lewis, 1884
First - 3rd dorsal striae complete. Body large, 4.50 - 6.06 mm .......................................... .
.......... ........ ...... .................. ............ ...... ........ ......... P. (P.) cambodjense Marseul, 1864
Front of head flat or weakly concave.
Fourth dorsal e1ytral stria longer than or as long as 5th .......... . P. (P.) horni Bickhardt, 1913
Fourth dorsal elytral stria shorter than 5th......................... . P. (P.) atratum Erichson, 1834

Fig. 55. Platysoma (Platylister) spp. Pronotum and left e1ytron. A: P. (P.) atratum Erichson. B:
P. (P.) combodjense Marseu!. C: P. (P.) horni Bickhardt. D: P. (P.) pini Lewis.
95

Platysoma (Platylister) atratum Erichson, 1834
(Fig. 55,56,57,58)
Hister paral/elipipedus Dejean, 1821: 48 (nom. nud.), given as synonym by Marseul, 1857: 472.
Platysoma atratum Erichson, 1834: 110.
Platysoma paral/elepipedum (sic): Dejean, 1837: 143.
Platylister atratus: Lewis, 1905b: 13; Hisamatsu, 1985b: 229; Sawada, 1988: 40 [Miyake-jima, Izu
Isles].
Platysoma buteanum Desbordes, 1922: 7, synonymized by Cooman, 1948: 132.
Platysoma (Platylister) atratum: Mazur, 1984: 228.

Japanese name: Indo-naga-emma-mushi.

Description. Body somewhat convex, oblong, black and shining; tibiae darkbrown; tarsi and antennae reddish brown. Body length, PPL, 4.69 - 6.25 mm (5.62 ±
0.11, n=14), PEL, 4.06 - 5.56 mm (4.99 ± 0.10, n=14), Width, 2.69 - 3.56 mm(3.16 ± 0.07, n=14). Biometric data are given in Table 8.

B

{

V

~

Fig. 56. Platysoma (Platylister) spp. Prosternum, mesosternum, metasternum and 1st abdominal
sternum. A: P. (P.) atratum Erichson. B: P. (P.) combodjense Marseul. C: P. (P.)
horni Bickhardt. D: P. (P.) pini Lewis.

96Surface of head (Fig. 57A) even but feebly depressed on apical 112; frontal stria
usually interrupted at each apical angle behind eye, and well impressed and carinate;disk evenly clothed with fine punctures which are separated by about 3 times theirdiameter. Labrum transverse oblong, its anterior margin strongly emarginate at middle.
Mandibles stout, short, and with longitudinal impression on dorsal surface.
Pronotal sides (Fig. 55A) parallel on basal 113, convergent forward on apical 2/3,
then strongly convergent on apical 116; emarginated portion of anterior margin bisinuatewith distinctly acute angle at middle (Fig. 55A). Marginal pronotal stria complete
laterally and weakly impressed, its apical end curved at apical angle and shortly
extending inwardly. Pronotallateral stria deeply impressed and complete laterally, andnarrowly interrupted at middle on anterior portion; disk densely clothed with finepunctures which are denser than those of the head.
Epipleural and elytral marginal striae weakly impressed and complete, the apicalend of the elytral stria extending along apical margin of elytron to middle. Epipleuraeven, and sparsely clothed with fine punctures. Subhumeral stria absent. Oblique
humeral stria (Fig. 55A) present on basal 1/3. First - 3rd elytral dorsal striae complete
and well impressed, the 3rd weakly bent inwardly on basal 112. Fourth dorsal stria
short and present on apical 116. Fifth often reduced basally, but usually present on
apical 113. Disk of elytra evenly and densely covered with fine punctures which are as
dense as on the pronotum.

........

~
[

J \ .

1

~

Fig. 57. Platysoma (Platylister) spp. A, D, G, J: Heads, frontal view. B, E, H, K: Propygidia andpygidia. C, F, I, L: Protibiae, dorsal view (left) and ventral view (right). A - C: P. (P.)
atratum Erichson. D - F: P. (P.) cambodjense MarseuL G-I: P. (P.) horni Bickhardt. J L: P. (P.) pini Lewis.
97Propygidium (Fig. 57B) irregularly covered with large punctures, which become
finer along the posterior margin and on a narrow longitudinal median band; interspaceamong the large punctures evenly and densely covered with fine punctures. Surface of
propygidium even. Pygidium densely covered with large punctures which are separated
by 0.3 times their diameter; interspace among the large punctures sparsely clothed withfine punctures; narrow band along posterior margin feebly elevated.
Anterior margin of prosternal lobe (Fig. 56A) round; marginal stria of lobe
deeply impressed, its posterior end strongly curved inwardly; disk flat, densely covered
with coarse punctures which are separated by their own diameter. Prosternal keel
narrow and without carinal stria; disk evenly covered with coarse punctures; basal apex
round. Lateral stria and lateral marginal stria deeply impressed and carinate.
Anterior margin of mesosternum strongly emarginate at middle; marginal stria
well impressed and complete, rarely interrupted at middle of anterior portion; disk
densely and finely punctate. Meso-metasternal suture weakly impressed. Punctation ofintercoxal disk of metasternum slightly finer than mesosternal ones. Lateral metasternal
stria deeply impressed, extending obliquely and posteriorly, its apical end attaining nearmetacoxa. Lateral disk densely covered with large and shallow punctures, which areusually fused with each other on apical 112.
Intercoxal disk of 1st abdominal sternum with punctation similar to that ofintercoxal disk of metasternum; 2 lateral striae present on each lateral side.

/~_..r.

r

\
I:

,

I

':~' ................-'1>. . . .• "--- , •.••.' '.: ... : •.
,{if _ 'I\,,; "'.

-'"....----,

o~~"Qo ~?" ,!, ~i'--;':";
0.<: ::o):~~/':~:/:'". :. ........
.....

o

~:

0

~.

~ii/~
E

F
•....

~ .. '~

0.5 mmFig. 58 Platysoma (Platylister) atratum Erichson. A: Aedeagus, dorsal view. B: Ditto, lateral view.
C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male
genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view.

98Table 8. Biometric data of Platysoma (Platylister) atratum Erichson and
P. (Platylister) cambodjense Marseul.

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

P. atratum

P. cambodjense

1.31-1.81 (1.52±0.03) 14
2.50-3.44 (3.05±0.06) 14
1.50-2.19 (1.90±0.04) 14
2.25-3.00 (2.74±0.05) 14
2.69-3.56 (3. 16±0.07) 14
1.69-2.25 (1.97±0.04) 14
0.44-0.69 (0.59±0.02) 14
0.63-0.94 (0.80±0.03) 14
0.94-1.25 (1.17±0.03) 14
0.81-1.19 (1.03±0.03) 14
1.00-1.31 (1.21±0.03) 14

1.50-2.00 (1.74±0.05) 8
2.75-3.63 (3.20±0.08) 8
1.63-2.06 (1.88±0.05) 8
2.31-3.00 (2.72±0.07) 8
2.81-3.75 (3.33±0.09) 8
1.56-2.19 (1.88±0.06) 8
0.44-0.56 (0.52±0.02) 8
0.63-0.94 (0.83±0.03) 8
1.00-1.50 (1.23±0.05) 8
0.94-1.25 (1.13±0.04) 8
1.19-1.69 (1.41±0.06) 8

Protibia (Fig. 57C) with 4 denticies on outer margin. Mesotibia with 4 denticies,
the median 2 denticies large. Metatibia with 4 denticies, the basal one very small.
Male genitalia as shown in Fig. 58.
Specimens examined. [Kyfishfr] Kagoshima-ken: 10" 1 ~, Sata-misaki, Sata, 3/v/1981, Y.
Takai; 1 ex, Ditto, 5/v/1985, Y. Takai; 1 ~ , Sata, lIviil1984, S. Masuda.
[Nansei Isles] Yaku-shima Is.: 10", Miya-no-ura, 28/vii/1969, H. Nara (NA). Tokara-Nakanoshima Is.: 1 ex., Nakano-shima, 22/viiI1986, H. Fujita (NA). Amami-Oshima Is.: 2~ ~ , Naze,
27/v/1978,H. Makihara; 1 ex., Kinsakubaru, 10/ix/1983, M. Ohara; 10", Shinokawa, Il1v/1953, T.
Shiraki (NA). Tokuno-shima Is.: 20"0' , Yonama, 12/iv/1964, R. Kawai. Okinawa-Ronta Is.: 10",
Yona, 30Ivi/1976, H. Makihara; 10", Kudeken, Chinen-mura, 22/v/1974, H. Makihara. Takekomi Is.:
1 ex., 19Iiii/v983, T. and T. Nakane (NA).
[faiwan] 1 ~ , Nanzankei, 30/iv/1965, T. Shirozu (NA).

Distribution. Japan (Izu Isles; Nansei Isles); India; Burma; Viet-Nam; Laos;
Taiwan; East China.
Remarks. Platysoma atratum superficially resembles P. horni; however, theformer can easily be distinguished by the different striation of the elytra and the pronotal
stria which is interrupted behind the head.
This species frequently occurs in decaying pineapple and banana; it was
sometimes collected also under bark and in rotting wood, and, according to Sawada(1988), under decaying Taro, Colocasia esculenta.
Platysoma (Platylister) carnbodjense Marseul, 1864
(Fig. 55,56,57,59)
Platysoma cambodjense Marseul, 1864a: 300 [Cambodia].
Platylister cambodjensis : Lewis, 1905b: 13.
Platylister niponensis Lewis, 1906b: 398 [Japan: Kioto (=Kyato), but this locality is corrected to true
locality "Kyfishfr" in Lewis, 1907]; 1907b: 98 [Kyfrshfr], synonymized by Bickhardt, 1920b:
61.
Platysoma (Platylister) cambodjense: Mazur, 1984: 228.
Japanese name: Oni-naga-emma-mushi.

Description. Body somewhat convex, oblong, black and shining; tibiae, tarsi,
99

antennae and mouthparts dark brown. Body length, PPL, 4.50 - 6.06 mm (5.34 ±
0.14, n=8), PEL, 4.19 - 5.44 mm (4.88 ± 0.13, n=8), Width, 2.81 - 3.75 mm (3.33± 0.09, n=8). Biometric data are given in Table 8.
Head (Fig. 570) broad, and deeply and broadly excavate; frontal stria weakly
impressed, complete and sinuate behind antennal cavity; disk evenly clothed with finepunctures which are separated by 2 - 4 times their diameter. Labrum transverse oblong,its anterior margin feebly emarginate inwardly. Mandible stout, well developed andwith a denticle on inner margin.
Pronotal sides (Fig. 55B) convergent forward slightly on basal 2/3 and thenstrongly. Apical angles acute. Marginal pronotal stria complete laterally and its apicalend united with lateral pronotal stria. Lateral pronotal stria strongly carinate andcomplete. Disk of pronotum densely covered with fine punctures which are separated
by about 4 times their diameter. Antescutellar area with a longitudinal puncture which islightly impressed.
Epipleura of elytra shining and scattered with microscopic punctures. Marginal
epipleural stria well impressed and usually complete, sometimes interrupted at middle.
Marginal elytral stria deeply impressed, carinate and complete. Interval between thesestriae with a short stria medially. Subhumeral stria absent. Oblique humeral stria (Fig.
55B) lightly impressed on basal 113. First - 3rd dorsal striae complete, and deeply andbroadly impressed. Fourth dorsal stria present on apical 1/2. Fifth dorsal on apical 113,
its basal 112 often represented by several punctures. Sutural stria represented by several
punctures and present on medio-apical 115. Surface of elytra sparsely clothed withmicroscopic punctures.
Propygidium (Fig. 57E) densely covered with large, deep and longitudinal
oblong punctures which are absent along margin. Pygidium densely covered with large,
deep and hexagonal punctures which are twice as large as the punctures of the
propygidium; surface feebly convex medially and posterior margin strongly elevated.
Anterior margin of prostemallobe (Fig. 56B) broadly truncate. Marginal stria oflobe well impressed, narrowly interrupted at middle, and its posterior end strongly bent
inwardly. Disk of lobe irregularly and finely punctate, the punctures being separated byabout 3 (1 - 5) times the diameter. Prosternal keel broad, without carinal stria;
punctation of surface similar to that of the lobe. Lateral and lateral marginal striae deeply
impressed and carinate.
Anterior margin of mesosternum (Fig. 56B) strongly emarginate medially;
marginal stria complete and carinate laterally and broadly interrupted anteriorly; disk
densely and moderately punctate, the punctures being separated by about their own
diameter. Meso-metasternal suture lightly impressed and obtusely angulate at middle.
Punctation of intercoxal disk of metasternum similar to that of mesosternum. Lateralmetastemal stria deeply impressed, extending obliquely and posteriorly, its apical end
attaining at apical 116 of metepisternal suture. Lateral disk densely covered with large,
shallow and round punctures, which are usually fused with each other medio-apically.
Intercoxal disk of 1st abdominal sternum with punctation similar to that of theintercoxal disk of the metasternum; 1 lateral stria present on each lateral side; the lateraldisk densely with longitudinal impressions.
Protibia (Fig. 57F) with 4 large denticles on outer margin; interval between 2
median denticles broad. Mesotibia with 4 dentic\es. Metatibia with 3 dentic\es.
Male genitalia as shown in Fig. 59.

100

0.5 mm,Fig. 59 Platysoma (Platylister) cambodjense Marseu!. A: Aedeagus, dorsal view. B: Ditto, lateralview. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Male genitalia, 9th and 10th
tergites, dorsal view. E: Male genitalia, 9th sternum (spicule), ventral view.
Specimens examined. [Nansei Isles] Yaku-shima Is.: Id", 15/vii/1988, K. Nakamine; 1 d",Miyanoura, 13/vii/1970, M. Ishida; 1 ex., Kosugi-dani, 14/vii/1970, T. Watanabe (NSMT); 2 exs.,
Okawa, 22/vii/1975, M. Akiyama (NSMT); 1 ex., Ditto, 28/vii/1975, H. Akiyama; 1 ex., Shiratani,
21/vii/1975, H. Akiyama; 1 ~, Kurio, 6/viii/1958, Y. Asano. Tokara-Nakano-shima Is.: 1 ex.,
Nakano-shima, 26/iv/1975, H. lrie (NSMT); Id", 1 ex., Ditto, 25/vi/1972, 24/vi/1973, H. Makihara.
Amami-Oshimals.: 1 ex., Ikari, 29!v!1960, T. Shibata (NSMT); 1d", Kinsakubaru, 30/vi/1984, M.
Ohara. Okinawa-Honta Is.: 1 ex., Y ona, Kunigami, 13/vil 1973, Y. Kurosawa (NSMT).

Distribution. Japan (KyOshO, Nansei Isles); Cambodia; Burma; Taiwan; North
India; Viet-Nam; Laos; Thailand; Bhutan.
Remarks. Platysoma cambodjense is a relatively large species, but rare small
specimens of this species resemble P. pini; however, they can easily be recognized
mainly by the striation of the elytra (see also the key and the description).
Little is known about the habitat of this species. It frequently occurs under bark.
Platysoma (Platylister) horni Bickhardt, 1913
(Fig. 55, 56, 57)
Platysoma horni Bickhardt, 1913: 169.
Platylister horni: Lewis, 1915: 55.
Platysoma (Platylister) horni: Mazur, 1984: 230 [catalogued].

Japanese name: Horn-naga-e=a-mushi.

Description. Body somewhat oblong, black and shining; tarsi and antennae
reddish brown. Body length, PPL, 4.63 - 4.81 mm (4.72 ± 0.07, n=2), PEL, 4.064.44 mm (4.25 ± 0.13, n=2), Width, 2.63 - 2.81 mm (2.72 ± 0.07, n=2). Biometricdata are given in Table 9.

101

Table 9. Biometric data of Platysoma (Platylister) horni Bickhardt and P. (Platylister) pini Lewis.

APW
PPW

PL

EL
EW
ProW

ProL
PyL
PTL
MSTL
MEL

P. horni

P.pini1.19-1.31 (1.25±0.04) 2
2.50-2.75 (2.63±0.09) 2
1.50-1.63 (1.56±0.04) 2
2.31-2.56 (2.44±0.09) 2
2.63-2.81 (2.72±0.07) 2
1.56-1.75 (1.66±0.07) 20.44
20.75-0.81 (0.78±0.02) 20.88
20.81-0.94 (0.88±0.04 2
1.00-1.06 (1.03±0.02) 2

1.31-1.50 (1.41±0.02) 17
2.38-2.75 (2.57±0.03) 17
1.38-1.63 (1.50±0.02) 17
2.00-2.44 (2.22±0.03) 17
2.44-2.75 (2.60±0.02) 17
1.38-1.75 (1.54±0.03) 17
0.38-0.56 (0.44±0.01) 17
0.63-0.81 (0.70±0.01) 17
0.81-1.13 (1.02±0.02) 16
0.81-1.19 (0.93±0.02) 16
1.00-1.25 (1.15±0.02) 16

Frontal stria of head (Fig. 570) deeply impressed, carinate and complete, thelateral portions convergent basally; the disk feebly depressed on apical 112 and evenly
with fine punctures which are separated by about 4 times their diameter. Labrumtransverse, its anterior margin feebly curved inwardly.
Pronotal sides (Fig. 55C) nearly straight and weakly convergent forward on
basal 3/4, thence strongly arcuate and convergent apically. Apical angles acute.
Marginal pronotal stria complete laterally. Pranoial lateral stria complete andsubcarinate, the lateral portion and median part of the anterior portion distant from the
pro notal margin. Disk densely clothed with fine punctures which are separated by 2 - 3times their diameter.
Epipleural marginal stria weakly impressed and complete. Elytral marginal stria
straight and deeply carinate. Oblique humeral stria (Fig. 55C) present on basal 113.
First - 3rd dorsal striae deeply impressed and complete, the 2nd basal end being
somewhat shortened. Fourth stria impressed on apical 112. Fifth dorsal stria present on
apical 1/3, its basal end often abbreviated. Surface of elytra with punctation similar tothat of the pronotum.
Propygidium (Fig. 55H) irregularly covered with coarse, round and deeppunctures which are separated by 0.5 times their diameter; interspace among the coarse
punctures finely and sparsely punctate, the punctures becoming denser along theposterior margin. Pygidium densely covered with coarse, round and deep punctures
which become somewhat larger than those of the propygidium and are separated by 0.3
times their diameter; interspace among the coarse punctures and a narrow band along
the posterior margin scattered with fine punctures, the band feebly elevated.
Prosternallobe (Fig. 56C) broad and even; apical margin truncate; marginal stria
well impressed and somewhat distant from the margin, its posterior ends convergent
inwardly; disk irregularly and finely punctate, the punctures being separated by about 3(1 - 5) times the diameter. Prosternal keel even; disk finely clothed with punctures
which become sparser than those of the lobe; lateral stria and lateral marginal stria
deeply impressed and carinate.
Anterior margin of mesosternum (Fig. 56C) feebly emarginate medially; marginalstria complete laterally and narrowly interrupted at middle on anterior portion; disk
evenly clothed with fine punctures which are separated by about 3 times their diameter.
Meso-metasternal suture absent. Punctation of intercoxal disk of metasternum similar to102that of the mesosternum; lateral metasternal stria deeply impressed, extending obliquelyand posteriorly, its apical end irregularly curved and attaining near the metacoxa. Lateralmetasternal disk with oblique, oval and shallow punctures on medio-basal 112,
elosewhere with semicircular, large and shallow punctures, which become smaller
apically, with moderate punctures intermingled.
Intercoxal disk of 1st abdominal sternum with punctation similar to that of themesosternum; 2 lateral striae present on each side, the outer one abbreviated on basal
113.
Protibia (Fig. 571) with 4 denticles on outer margin; interval between 2nd and 3rd
denticles broad. Mesotibia with 4 denticles. Metatibia with 3 denticles.
Specimens examined. [Nansei Isles] Ishigaki Is.: 10"', xi-xiii 1952, G. E. Bohart (BSM); 1 ex.,
Banna-dake, 51 viii 1974, H. Takizawa.

Distribution. Japan (Nansei Isles: Ishigaki Is.); Taiwan.
Remarks. Platysoma horni is easily recognized by the striation of the elytra.
Platysoma (Platylister) oberthuri Cooman, 1948
Platysoma oberthuri Cooman, 1948: 132; Mazur, 1984: 230 [catalogued].
Platysoma mimicum Desbordes, 1922: 235 (nec Lewis, 1914a: 239), correctly detected by Cooman,
1948: 132.
Platysoma (Platylister) oberthuri: Mazur, 1984: 231.
Japanese name: Amami-naga-emma-mushi.
Specimens examined. No specimen has been available for my study.

Distribution. Japan (Nansei Isles: Amami-Oshima Is.).
Platysoma (Platylister) pini Lewis, 1884
(Fig. 55,56,57,60)
Platysoma pini Lewis, 1884: 133 [Japan].
Platylister pini: Hisamatsu, 1985b: 229, pi. 41, no. 29.
Platysoma (Platylister) pini: Mazur, 1984: 231.
Japanese name: Matsu-naga-emma-mushi.

Description. Body oblong-oval, black and shining; tibiae, tarsi, mouthparts andantennae rufopiceous. Body length, PPL, 4.00 - 4.75 mm (4.36 ± 0.05, n=17), PEL,
3.63 - 4.31 mm (4.03 ± 0.04, n=17). Width 2.44 - 2.75 mm (2.60 ± 0.02, n=l7).
Biometric data are given in Table 9.
Frontal stria of head (Fig. 571) feebly carinate and complete; the disk strongly
depressed on apical 112 and densely covered with fine punctures which are separated byabout 1.5 (1 - 2) times the diameter. Labrum transverse, its anterior margin feebly
emarginate inwardly.
Pronotallateral sides (Fig. 55 D) convergent forward weakly on basal 5/6 andthen strongly and arcuately. Apical angles acute. Marginal pronotal stria complete
laterally. Pronotallateral stria complete and strongly carinate, the lateral portion broadly
distant from the margin and the anterior portion densely carinate. Disk densely andfinely punctate, the punctures being separated by 2 - 5 times their diameter.
103

Epipleural marginal stria deeply impressed and carinate, and present on basal 112.
Elytral marginal stria complete, deeply impressed and carinate, the apical end extending
along anterior margin and attaining to apical end of the 2nd dorsal stria. Another stria
complete and deeply carinate between epipleural and elytral marginal striae. Oblique
humeral stria (Fig. 55D) present on basal 114. First - 4th dorsal elytral striae complete
and deeply impressed, the basal portion of 4th being a little shortened. Fifth dorsal stria
abbreviated on basal 116. Sutural stria represented by several coarse punctures on
medio-apical 114. Disk densely and finely punctate, the punctures being much sparser
than those of the pronotum.
Propygidium (Fig. 57K) irregularly covered with ocelloid, large, oblong andshallow punctures; interspace among the ocelloid ones sparsely clothed with finepunctures. Pygidium densely covered with ocelloid, large, round and shallow
punctures, which are separated by about 0.5 times their diameter; interspace among theocelloid ones sparsely and finely punctate; narrow area along the posterior margin
strongly elevated and sparsely and finely punctate.
Anterior margin of prostemal lobe (Fig. 56D) truncate, its marginal stria wellimpressed and complete, the posterior end extending inwardly and attaining to suture
between prosternal keel and lobe; disk evenly covered with fine punctures which areseparated by about 4 times their diameter. Prostemal keel broad, even and sparsely andfinely punctate; lateral stria and lateral marginal stria deeply impressed and carinate.
Anterior margin of mesosternum (Fig. 56D) feebly emarginate medially; marginalstria complete laterally and very weakly impressed medially on anterior portion;
punctation of disk similar to that of the prosternal keel. Meso-metasternal suture lightly,///~~"'~" ~-==\/,'""-,,"'?-"- - (

"'./)

.,

c

F
0.5 mm,Fig. 60. Platysoma (Platylister) pini Lewis. A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia,
9th and 10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view.

104impressed. Punctation of intercoxal disk of metasternum similar to that of themesosternum; lateral metasternal stria deeply impressed, extending obliquely andposteriorly; lateral metasternal disk densely covered with large and shallow punctures,
which are often fused together.
Intercoxal disk of 1st abdominal sternum with punctation similar to that of themesosternum; 1st abdominal stria completely present, and another fragmentary stria
impressed on apical 112.
Protibia (Fig. 57L) with 5 denticles on outer margin. Mesotibia with 4 denticles
on outer margin, the apical 2 set close near the apical angle of tibia. Metatibia with 3
denticles on outer margin, the apical 2 present on the apical angle of tibia.
Male genitalia as shown in Fig. 60.
Specimens examined. [Hokkaido] 1 ex., Mt. Kariba, Shimamaki, 6-8/viiiI1988, M. Ohara, T.
Tanabe and K. Sayama.
[Honshu] Aomori-ken: 1 ex., Rangan-no-mori, Nishi-Tsugaru, 24/vii/1977, A. Abe (NA).
Fuknshima-ken: 3 exs., Karasugawa-rindo, Moniwa, Iizaka-cho, 16/vi, 6, 22/vii/1976, K. Tazoe.
Miyagi-ken: 3 exs., Akiu, Moriyasu, 24/vi/1951, no collector's name (NSMT). Tochigi-ken: 1 ex.,
Kawamata, Kuriyama-mura, 14/vii/1990, K. Satoh (TPM). Tokyo-to: 1 ex., Takao-san, 27/vi/1971,K.
Masumoto. Gifu-ken: 1 ex., Hokiwaki, Mino-shi, 23/vi/1979, Y. Takai. Kyoto-fu: 2 exs., Mt. Daihi,
15/vi/1941, K. Sakaguchi (NA). Osaka-fu: 2 exs., Katsuno-ji, Mino, 2/viii/1949, K. Sawada (NA).
Hyogo-ken: 1 ex., Hataganaru, 24, 25/v/ 1955, T. Nakane (NA).
[KyushU] Miyazaki-ken: 1 ex., Takachiho-no-mine, 1O/vii/1984, M. Ohara.
Distribution. Japan (Hokkaid6, Honshu, Shikoku, Kyushfi); Taiwan. New to
Hokkaid6.
Remarks. Platysoma pini resembles P. cambodjense, but differs from the latter
externally by the smaller average size and the different striation of the elytra (see thedescription).
Little is known about the habitat of this species. P. pini is frequently collectedunder the bark of the pine trees.
SUBGENUS CYLISTER COOMAN, 1941
Cylister Cooman, 1941: 307 [type species: Hister elongatus Olivier, 1789: 16, nec Thunberg, 1787:33, originally designated].

Platysoma (Cylister) lineicollis Marseul, 1873
(Fig. 61, 62, 63)
Platysoma lineicollis Marseul, 1873: 223 [Kiu-Siu et Niphon].
Clistosoma lineicolle: Lewis, 1905b: 18.
Cylister lineicolle: Cooman, 1948: 309.

Japanese name: Naga-emma-mushi.
Description. Body oblong, black and shining; femora, tibiae, tarsi and antennae
rufopiceous. PPL, 2.84 - 3.50 mm (3.28 ± 0.04, n=20); PEL, 2.52 - 3.17 mm (2.97± 0.04, n=20). Width, 1.34 - 1.64 mm (1.55 ± 0.02, n=20). Biometric data are given
in Table 10.
Frontal stria of head (Fig. 61A) deeply impressed, carinate and complete. Disk ofhead depressed on apical 112 and sparsely covered with coarse punctures which are105

separated by 3 - 4 times their diameter; interspace among the coarse punctures evenly
and finely punctate; these coarse and fine punctures becoming denser and finer apically.
Labrum transverse, its anterior margin feebly curved inwards.
Pronotal sides (Fig. 618) straight on basal nine-tenths, thence strongly andarcuately convergent forward. Apical angles acute. Marginal pronotal stria complete
laterally. Pronotallateral stria carinate and complete, the anterior portion nearly straight,
interrupted behind eyes, and sometimes interrupted at middle; the lateral end of the
portion often extending obliquely and posteriorly. Disk evenly covered with coarse
punctures which become coarser and somewhat more oblong on lateromedian 1/6 andfiner narrowly along lateral stria; median 1/3 with fine punctures intermingled.
Epipleura even, not concave. Epipleural marginal stria lightly impressed,
complete and broadly distant from outer margin. Elytral marginal stria complete anddeeply impressed. Subhumeral striae absent. Oblique humeral stria (Fig. 618) present
on basal 113. First - 4th dorsal striae deeply impressed and complete. Fifth dorsal stria
present on apical 2/5, its basal portion usually represented by several punctures. Sutural
stria present on apical 112, its apical portion somewhat shortened. Surface of elytra
evenly clothed with fine punctures which are separated by about 3 times their diameter;
apical transversal band sparsely and coarsely punctate.
Propygidium (Fig. 61C) irregularly covered with coarse, round and shallow
punctures; interspace among the coarse punctures finely and sparsely punctate.
Pygidium densely covered with coarse, round and somewhat deep punctures on basal
112, which are separated by 0.5 - 1 times the diameter; interspace among the coarse
ones and apical 112 of disk clothed with fine punctures which are separated by 4 - 6times their diameter.

"-~{~Q\jVI\':.:.'.;< .:) ~P, ,- - - \

I

,

--

I

\, ~

,
,,
,

I

o

Fig. 61. Platysoma (Cylister) lineicollis Marseul. A: Head, dorsal view. B: Pronotum and left
elytron. C: Propygidium and pygidium. D: Prostemum, mesosternum, metasternum and 1st
abdominal sternum. E: Protibia, dorsal view. F: Ditto, ventral view.
106

Table 10. Biometric data of Platysoma (Cylister) lineicollis Marseul
and P. (Apobletes) shaumei Marseu!.

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL

MTIL

P. (Cylister) lineicollis

P. (Apobletes) shaumei

0.82-1.01 (0.95±0.01) 20
1.34-1.64 (1.52±0.02) 20
0.98-1.24 (1.15±0.01) 20
1.34-1.83 (1.62±0.03) 20
1.34-1.64 (1.55±0.02) 20
0.82-1.11 (1.00±0.02) 20
0.33-0.52 (0.44±0.01) 20
0.46-0.62 (0.53±0.01) 20
0.59-0.72 (0.64±0.01) 20
0.46-0.69 (0.60±0.01) 20
0.62-0.78 (0.70±0.01) 20

0.95-1.14 (1.01±0.04) 4
1.86-2.26 (2.04±0.08) 4
0.92-1.14 (1.02±0.04) 4
1.50-1.80 (1.67±0.06) 4
1.90-2.32 (2.08±0.08) 4
1.11-1.31 (1.19±0.04) 4
0.36-0.43 (0.39±0.02) 4
0.46-0.49 (0.47±0.01) 4
0.59-0.72 (0.63±0.03) 4
0.49-0.62 (0.55±0.03) 4
0.62-0.78 (0.70±0.03) 4

Prosternallobe (Fig. 61D) broad and even; its apical margin round; marginal stria
carinate and impressed, its posterior ends inwardly bent and convergent medially; disk
densely covered with moderate punctures which are separated by their own diameter.
Prosternal keel narrow and elevated, its top even; carinal striae wanting.
Anterior margin of mesosternum (Fig. 61D) broadly and deeply emarginate;marginal stria complete and strongly carinate; disk shining, and sparsely and finely
punctate. Meso-metasternal suture lightly impressed. Punctation of intercoxal disk ofmetasternum similar to that of the mesosternum; lateral metasternaI stria deeply
impressed, extending posteriorly and obliquely, its apical end attaining to near themetacoxal cavity; another lateral stria (post-mesocoxal stria) present outside the lateral
stria; lateral disk densely covered with large, round and shallow punctures and withmoderate ones intermingled, these punctures becoming finer apically.
Punctation of intercoxal disk of 1st abdominal sternum similar to that of theintercoxal disk of the mesosternum; 2 lateral striae present, the outer stria abbreviatedon basal 112.
Protibia (Fig. 61E, F) with 4 denticles on outer margin. Mesotibia with 3
denticles on outer margin, the apical most one bearing 2 spinules. Metatibia with 2
denticIes, the apical most one with 2 spinules.
Male genitalia as shown in Fig. 62.
Specimens examined. [HonshU] Iwate-ken: 1 ex., Matsuo, 17-19/vii/1954, S. Takagi (ElHU).
Fukushima-ken: 1 ex., Oyama, Wakamatsu, 14/v/1947, no collector's name. (NA); 4 exs., Mt. Aoki,
northern Aizu, lIxi/1948, K. Nagayama (NA); 3 exs., Kawanuma, Nozawa, 211iii/1949, K. Nagayama(NA); 2 exs., Akaidani, 5/v/1949, K. Nagayama (NA); 3 exs., Yanaizu, Onuma, 18/iv/1948, K.
Nagayama (NA); 2 exs., Niizuru, Onuma, 311iv/1948, 6/iv/1949, K. Nagayama (NA); 1 ex., Takizawa
Pass., Ikki, Wakamatsu, 14/iii/1948, Y. Kurosawa (NSMT); 1 ex., Yunohana, Tateiwa, Minami-Aizu,
6/v/1949, Y. Kurosawa (NSMT); 1 ex., Oyama, Wakamatsu, 14/v/1947, Y. Kurosawa (NSMT).
Tokyo-to: 1 ex., Mt. Takao, 26/vi/1960, S. Nomura (NSMT). Niigata-ken: 15 exs., Kurokawa,northern Echigo, 211i, 2/iv, 20/v, 24/ix/1956, lIxi/1957, 28/xii/1958, K. Baba (NA); 2 exs., Ditto,lIvi/1958, H. Koike (NA). Sado Is.: 1 ex., Takaehi Y., 27/viiI1947, M. Iga (NSMT). Yamanashi-ken:
5 exs., Kiyosato, no date, Y. Kishii (NA); 1 ex., Agematsu, Mt. Koma, 12/vi/1949, A. Nagatomi.
(NA). Gifu-ken: 1 ex., Seki-shi, 13/xii/1976, Y. Takai; 1 ex., Yoro-koen, Yoro, 6/v/1979, Y. Takai; 2exs., Tachibana, Mino-shi, 16/iii/1979, Y. Takai. Shizuoka-ken: 1 ex., Mt. Amagi, Izu, 30Iiii/1952,
J. Aoki (NA). Aiehi-ken: 4 exs., Nagoya, 4/v/1944, S. Osawa (NA); 1 ex., Nagoya, Owari, 4/v/1944,
S. Osawa (NA); 2 exs., Okehazama, i/1946, M. Yasuda (NA). Osaka-fu: 1 ex., Hirano, 22/viii/1950,
107

, ..-,-);,.;..;!~""

.,;'

"

."

'~~{~--~

D,,~
0.25 mmFig. 62. Platysoma (Cylister) lineicollis Marseul. A: Aedeagus, dorsal view. B: Ditto, lateral view,
C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view, E: Male
genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view,Fig.63. Collection sites of Platysoma (Cylister) lineicollis Marseul in Japan,
F. Takahashi (NA); 3 exs., Minoo, 30/iv/2605 (=1945), no collector's name. (NA). Kyoto-fu: 1 ex.,
Sugi-toge, 4/v/1983, H. Ashida, Hyogo-ken: 2 exs., Mikage, Kobe, 5, 6/i/1940, no collector's name.
(NA); 1 ex" Kobe, 27/iii/1930, J, E A Lewis (BSM); 1 ex., Motoyama-mura, near Kobe, 27/x/1941,
K. Suzuki (NA); 1 ex" Rokko, 17/iii/1935, T Nakane (NA). Wakayama-ken: 1 ex., Kii-Oshima, 2124/v/1964, T. Kumata (EIHU); 1 ex., Mt. Koya, Kii, 4/viii/1940, T. Okada (NA). Okayama-ken: 1108

ex., Tengu-yama, Tamano, 13/viii/1975, T. Aono (NA). Tottori-ken: 1 ex., Daisen, 18/vii/1952, T.
Nakane (NA); 1 ex., Ditto, 18/x11987, H. Daido.
[Shikoku] Ehime-ken: 2 exs., Omogo, 9/xi/1953, T. Nakane (NA). Kochi-ken: 1 ex.,Mikazuki, Tosa, 10/iv/1938, M. Okamoto (NA).
[Kyfishu] Fukuoka-ken: 1 ex., Moji, 29/v/1947, C. Watanabe (EIHU); 2 exs., Ditto,
27/vii11936, M. Uno (NA); 2 exs., Shimo, Hirokawa, no date, Y. Miyake (NA); 1 ex., Hakozaki,
27/iii/1945, Y. Kurosawa (NA); 1 ex., Ditto (NSMT); 2 exs., Togamori, 28/v/1956, N. Gyotoku(NA); 1 ex., Homan-zan, 2/iv/1980, S. Ohmomo (NSMT); 1 ex., Wakasugi, 29/iii11933, T. Shirozu(NSMT); 1 ex., Mt. Shoji, 19/iv/1964, S. Nakao (NA). Miyazaki-ken: 1 ex., Shiiba-mura,l/viii/1970, A. Nagai (NA). Kagoshima-ken: 2 exs., Kaimon-cha, 4/xi/1980, Y. Takai.
Distribution (Fig. 63). Japan (Honshu, Shikoku, Kyushu); Taiwan.
Remarks. Being the only representative of the subgenus Cylister in Japan, C.
lineicollis can easily be recognized by the subgeneric characteristics.
This species occurs under the bark of pine trees.

Platysoma (Cylister) elongatum (Thunberg, 1787)
Hister elongatus Thunberg, 1787: 33.
Hister oblongus Fabricius, 1792: 75 [South Sweden]; Thunberg, 1794: 63, synonymized by Kannar,
1979: 25.
Platysoma oblongus: Leach, 1817: 79.
Ahbotia georgiana Leach, 1830: 157, synonymized by Waterhouse, 1868: 168.
Cylistosoma oblongum: Lewis, 1905b: 18.
Cylister oblongum: Kryzhanovskij, 1965: 103.
Cylister oblongus: Kusui, 1984: 7.

Kusui (1984) recorded a specimen collected from an imported larch wood at
Naoetsu harbor, J6etsu, Niigata-ken, Honshu, Japan.
Distribution. Europe; Siberia; Mongolia; Amurskiy Kray.
SUBGENUS ApOBLETES MARSEUL, 1860
Apobletes Marseul, 1860: 852 [type species: Apobletes tener Marseul, 1860: 857, designated by
Bickhardt, 1917: 136].

Platysoma (Apobletes) schaumei (Marseul, 1860)(Fig. 64, 65)
Apobletes schaumei Marseul, 1860: 857 [Burma].
Apobletes tener Marseul, 1860: 859; Hisamatsu, 1985b: 229 [Japan: Nansei Isles], synonymized by
Reichardt, 1932: 114.
Apobletes schaumi (sic):- Marseul, 1864a: 290.
Apobletes schaumi var. tener: Cooman, 1932b: 99.
Platysoma (Apobletes) schaumei: Mazur, 1984: 235.

Japanese name: Hirata-naga-emrna-mushi.
Description. Body length, PPL, 3.37 - 4.12 mm (3.74 ± 0.15, n=4); PEL, 2.84
- 3.40 mm (3.11 ± 0.10, n=4). Width, 1.90 - 2.32 mm (2.08 ± 0.08, n=4). Body
strongly depressed (Fig. 64A), oblong-oval, shining and black; [omora, tibiae and tarsi
reddish-brown. Biometric data are given in Table 10.
109

Fig. 64. Platysoma (Apobletes) schaumei Marseul. A: Adult, lateral view. B: Head, frontal view. C:
Pronotum and left eIytron. D: Prosternum, mesosternum, metasternum and 1st abdominal
sternum. E: Propygidium and pygidium. F: Protibia, dorsal view. G: Ditto, ventral view.

Frontal stria (Fig. 64B) usually narrowly interrupted at each apical angle,
sometimes complete, the anterior portion nearly straight and the lateral portion regularly
curved. Disk of head excavate on apical 1/2 and evenly covered with fine punctures
which are separated by about twice their diameter; interspace among the fine punctures
evenly clothed with microscopic punctures. Labrum transverse oblong, its anterior
margin emarginate medially. Mandible short, stout, with a large denticle on inner side.
Pronotal sides (Fig. 64C) convergent forward feebly, arcuately on basal 5/6 andthen strongly arcuately. Marginal stria complete and carinate laterally. Pronotal lateral
stria complete and narrowly interrupted at middle on anterior portion. Disk of pronotum
sparsely clothed with microscopic punctures which are separated by 3 - 4 times theirdiameter and become larger on the lateral 116; the large punctures dense on lateral 114 ofbasal margin.
Epipleural and elytral marginal striae deeply carinate and complete. Epipleura
somewhat excavate medially. Subhumeral stria deeply impressed on basal 112. Oblique
humeral stria (Fig. 64C) present on basal 113. First and 2nd dorsal striae complete andsparsely crenate; 3rd dorsal stria broadly interrupted medio-apically, the interrupted
portion 112 as long as elytron; 4th shortly present on apex; 5th and sutural striae absent.

110

Disk of elytra evenly clothed with microscopic punctures which are separated by about
5 times their diameter and become larger along apical margin.
Propygidium (Fig. 64E) with large and ocelloid punctures on anterolateral area,
the punctures being separated by 0.3 times their diameter and becoming sparser
medially; with microscopic punctures sparsely intenningled; surface feebly convex onlateral 114. Pygidium coarsely and deeply punctate, the punctures irregularly separated
by 0.3 - 1 times the diameter; surface feebly depressed on each angle.
Prosternallobe (Fig. 64D) broad and even, its anterior margin round; marginalstria complete, the posterior end inwardly extending on basal 112; disk of lobe sparsely
clothed with microscopic punctures, and on lateral 114 of disk coarsely and densely
punctate. Prosternal keel even and broad, without carinal stria; lateral stria and lateral
marginal prosternal stria clearly impressed, the fonner complete and the latter shortened
anteriorly; disk of keel with punctation similar to that of the lobe; posterior margin
nearly straight.
Anterior margin of mesosternum feebly bisinuate (Fig. 64D), its marginal stria
straight along the anterior margin, paralleled laterally, shortly interrupted on each basal
angle, and narrowly interrupted medially; punctation of disk similar to that of prosternalkeel. Meso-metasternal suture complete, obtusely angulate at middle. Intercoxal disk ofmetasternum broad, even and clothed with microscopic punctures which are finer andsparser than those of the prosternal keel. Lateral metasternal stria extending obliquely.......

E

o
<::.

o

0

0

0'"
<> • •,)':;>

00

o

F

~

~
H

0.5 mm!

Fig. 65. Platysoma (Apobletes) schaumei Marseul. A: Aedeagus, dorsal view. B: Ditto, lateralview. C: Male genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. D: Ditto,
lateral view. E: Male genitalia, 8th tergite. F: Ditto, lateral view. G: Male genitalia, 8thsternum, ventral view. H: Ditto, lateral view.

111and posteriorly, and nearly complete but sometimes interrupted at apical 113, the apicalend attaining near the metacoxa. Posterior mesocoxal stria wanting. Lateral metastemal
disk densely covered with large, shallow and semi-circular punctures which arerounded basally; interspace among the large punctures intermingled with fine punctures.
Intercoxal disk of 1st abdominal sternum sparsely clothed with microscopic
punctures which are separated by 3 - 5 times their diameter; with 2 striae on each side,
the outer stria present on apical 112. Lateral disk with 8 - 9 longitudinal and large
punctures. Lateral area of 2nd - 4th abdominal sterna coarsely and densely punctate.
Protibia (Fig. 64F, G) with 4 denticles on outer margin; interval between the 2nd
and 3rd denticles broad. Mesotibia without denticle, but the apex expanded outwardly
and with 2 spines.
Male genitalia as shown in Fig. 65.
Specimens examined. [Nansei Isles] Okinawa-Honto Is.:' 2d" d" 1!f, Yona, 30/vi/1976, H.
Makihara

Distribution. Japan (Nansei Isles); Burma; Oriental Region.
Remarks. The only Japanese representative of the subgenus Apobletes, A.
shaumei can easily be recognized by the subgeneric characteristics.
Little is known about habitat of this species. It occurs under bark.
GENUS EBUSIA LEWIS, 1889
Eblisia Lewis, 1889: 280 [type species: Phelister lunaticus Marseul, 1864a: 316].
Eurylister Bickhardt, 1920: 213, synonymized by Mazur, 1990: 748.
Chronus Lewis, 1914b: 285, synonymized by Mazur, 1990: 748.

Eblisia satzumae (Lewis, 1899)
Platysoma satzumae Lewis, 1899: 8; Ohara, 1986: 94.
Eblisia satzumae: Ohara, 1993c: 5.

Japanese name: Satsuma-naga-emma-mushi.

Distribution. Japan (KyOshO, Nansei Isles).
TRIBE HrslERiNI GYLLENHAL, 1808
GENUS HrSlER LINNAEUS, 1758
Hister Linnaeus 1758: 358 [type species: Hister unicolor Linnaeus, 1758: 358, designated byWestwood, 1840: 22].
Histeranus Rafinesque, 1815: 112 (emend.).
Humister Houlbert et Monnot, 1923: 23, synonymized by Vienna, 1980: 257. Without type species.
Rhabdister Houlbert et Monnot, 1923: 35 (emend.).
Spilister Houlbert et Monnot, 1923: 22, synonymized by Cooman, 1947: 428. [type species: Hister
quadrimaculata Linnaeus, 1758: 358].

Diagnosis. Body oval or oblong-oval, somewhat convex, usually shining andblack, but rarely elytra with red maculation. Anterior margin of labrum usually nearlystraight, but rarely projecting slightly and outwardly. Frontal stria of head usually entire
and carinate, sometimes interrupted at middle of anterior portion. Pronotum with 1 or 2112

lateral striae which are usually crenulate; outer lateral stria usually complete on side,sometimes reduced posteriorly, and broadly interrupted behind head; inner lateral stria
usually entire. Marginal stria complete on side, and shortly or broadly interrupted
behind head; area inside apical angles without excavation or coarse punctation.
Epipleura of pronotum even or concave, often furnished with hairs. Antennal cavity not
deep, at most moderately deep. Elytron completely with 3 or 4 dorsal striae, other striae
usually reduced, rarely 2 or 5 striae complete. Degree of development of subhumeral
striae variable; external subhumeral stria usually present, not entire, and sometimes
absent, the stria usually deeply impressed, rarely formed as an arc and strongly
crenulate; internal subhumeral stria usually impressed, but not complete. Epipleuron of
e1ytron more or less concave, smooth and rarely punctate. Propygidium and pygidiumwith variable punctures, which are useful for species recognition. Anterior margin ofmesosternum usually clearly emarginate, rarely nearly straight; its marginal stria usuallywell impressed and complete. Protibia moderately expanded; 3 - 5 denticles on outermargin. Outer margin of meso- and metatibiae with 2 rows of setae. Aedeagus of male
genitalia slender cylindrical, usually depressed, and slit on midline of dorsal surface; theapical portion of dorsal surface often with small lobes along midline; median lobe
without sclerotized median armature. Spermatheca of female genitalia consisting ofseveral (4 - 8) receptacles on vagina, the receptacles slender, not coiled; bursa
copulatrix usually sclerotized with no function as a sack.
Remarks. The genus Hister Linnaeus contains about 240 nominal species
worldwide. In Japan 6 species are recorded. As the genus Hister is the oldest erectedof the family, many related genera have been separated from the Hister; for example,
Athlous Thomson, 1859, Macrohister Lewis, 1904, Margarinotus Marseul, 1853,
Merohister Reitter, 1909, and Pachylister Lewis, 1904. The genus, however, stillcontains a number of species, having no unique and clearly discriminating character
states, that is, it is a large complex genus. Kryzhanovskij and Reichardt (1976)
mentioned the necessity of separating the Hister complex into several subgroups.
Olexa (1982) has recently discussed the systematics of the genus and its allied genera
occurring in the Palaearctic Region. He divided the genus into 2 subgroups based on
the state of profemoral stria. But phylogenetic survey of the genus, as regards the world
species, is not satisfactory and needs detailed reexaminations.
Key to the Japanese species of the genus Hister
1(2)
2(1)
3(6)
4(5)
5(4)
6(3)
7(8)

Subhumeral stria absent ....................................................... H. impunctatus Osawa, 1952
Subhumeral stria present.
Ventral surface of profemur with a nearly complete stria.
First and 2nd dorsal striae of elytra complete............................. H.japonicus Marseul, 1854
First - 3rd dorsal striae of elytra complete ................................. H. congener Schmidt, 1885
Ventral surface of profemur with a stria shortly present on apical 113.
Pronotal anterior margin emarginate, the median portion of the emargination outwardly arcuate.
............. .......................................... ............................. H. simplicisternus Lewis, 1879
8(7)Pronotal anterior margin emarginate, the median portion not outwardly arcuate.
9(10) Pygidium with fine punctures which are separated by their own diameter.............................. .
............................................................................ H. unicolor leonhardi Bici,hardt, 1910
10(9) Pygidium densely with coarse punctures which are separated by 0.3 times their diameter to theirown diameter. ......................................................................... H. concolor Lewis, 1884

113

Hister impunctatus Osawa, 1952
Hister (Hister) impunctatus Osawa, 1952: 6 [Japan: My6ken, Osaka, Honshu].
Hister impunctatus: Mazur, 1984: 190 [catalogued].

Japanese name: Tsuya-emma-mushi.
Specimens examined. No specimens of the species have been available for my study.

Distribution. Japan (Honshu).
Hister japonicus Marseul, 1854
(Figs. 66, 67, 70 and 71)
Hister japonicus Marseul, 1854: 201, t. 6, f. 22 [Japon]; Marseul, 1873: 220 [Niphon et Kiu-siu (=Kyushu, Japan); commun dans Ie fumier].
Hister (Hister) japonicus: Bickhardt, 1918: 229 [China]; Desbordes, 1919: 387 [Japon; Chine; Indochine; Tonkin].

Japanese name: Yamato-emma-mushi.

Description. Male and female. Body length, PPL, male, 8.74 - 10.17 mm,
female, 9.50 - 11.59 mm, PEL, male, 7.89 - 8.74 mm, female, 7.70 - 9.41 mm.
Width, male, 6.65 - 7.51 mm, female, 6.56 - 8.17 mm. Biometric data are given in
Table 11. Body oblong-oval, black and shining; tibiae, tarsi and antennae reddish
brown.
Frontal stria of head (Fig. 66A) deeply impressed, carinate and usually complete,
the anterior portion usually straight, but sometimes narrowly interrupted and feebly andinwardly arcuate at middle; surface even, sparsely and finely punctate. Labrum short,transverse oblong.
Pro notal sides (Fig. 66B) rather strongly convergent forward on basal ninetenths. Apical angles round. Marginal pronotal stria complete laterally and broadlyinterrupted behind head. Outer lateral pro notal stria clearly impressed, carinate and its
length variable, usually complete, sometimes abbreviated on basal 1/2. Inner lateralpronotal stria complete, the anterior portion regularly and coarsely crenate. Disk of
pronotum sparsely covered with fine punctures which are separated by 10 times theirdiameter or more, and wholly covered with coriaceous ground sculptures; lateral 112 ofnarrow band along posterior margin coarsely punctate. Antescutellar area with a shortlongitudinal puncture.
Epipleural fossette of elytra feebly excavate and sparsely and coarsely punctate.
Marginal epipleural stria lightly impressed on apical 1/2. Marginal elytral stria carinate
and complete. Oblique humeral stria (Fig. 66B) lightly impressed on basal 1/2. External
subhumeral stria absent. Internal subhumeral stria deeply impressed and present on
apical 112. First and 2nd dorsal striae complete, well impressed, the basal 1/3 inwardlybent. Third dorsal stria present on basal 112, and its apical rudiment present on apical
1/8. Fourth and 5th dorsal striae usually short, occurring apically, sometimes wanting.
Sutural stria absent. Disk of elytra wholly covered with coriaceous ground sculpturesand sparsely and finely punctate.

114

Table 11. Biometric data for Hister japonicus Marseul.

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PIL
MSTL
MITL

Male

Female2.66-2.95 (2.80±0.023) 20
6.08-6.84 (6.39±0.049) 20
2.76-3.23 (2.99±0.030) 20
4.09-4.75 (4.44±0.038) 20
6.65-7.51 (7.03±0.053) 20
3.71-4.66 (4. 19±0.048) 20
1.38-1.71 (1.53±0.022) 20
1.71-2.28 (1.94±0.033) 20
1.90-2.47 (2. 19±0.027) 20
1.90-2.28 (2.05±0.024) 20
2.38-2.95 (2.74±0.030) 20

2.57-3.33 (2.90±0.036) 20
5.89-7.41 (6.75±0.075) 20
2.71-3.42 (3. 14±0.036) 20
4.09-5.23 (4.68±0.051) 20
6.56-8.17 (7.52±0.081) 20
3.90-5.13 (4.55±0.064) 20
1.33-1.81 (1.65±0.030) 20
1.85-2.28 (2.07±0.025) 20
2.09-2.76 (2.29±0.035) 20
2.00-2.38 (2. 14±0.024) 20
2.57-3.04 (2.84±0.025) 20~lFig. 66. Hister japonicus Marseul. A: Head, frontal view. B: Pronotum and left elytron, dorsal view.
C: Pro-, meso-, metasterna and 1st abdominal sternum. D: Protibia, dorsal view. E: Ditto,
ventral view. F: Profemur, ventral view. G: Female genitalia, spermatheca and bursa copulatrix,
lateral view (left side).

115

Propygidium with a slight depression on each side, and sparsely covered withcoarse, round and rather deep punctures which are unevenly separated by 1 - 10 times
their diameter; area between the coarse punctures sparsely scattered with fine punctures,and wholly covered with alutaceous ground sculptures. Punctation of pygidium similar
to that of propygidium, but much denser and coarser.
Anterior margin of prosternal lobe (Fig. 66C) regularly round; marginal stria
deeply impressed, carinate and complete; another short stria present on lateral area
between margin and the marginal stria; disk densely and coarsely punctate, thepunctures becoming finer and sparser medially. Suture between prosternal keel and lobecF'--_ _-----', 0.5 mm

DFig. 67. Hister japonicus Marseul. Male genitalia. A: Aedeagus, dorsal view. B: Ditto, lateral view.
C: 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: 9th and 10th tergites and 9thsternum (spicule), dorsal view. F: Ditto, lateral view.

116interrupted medially. Surface of prostemal keel even, impunctate, but lateral descendingarea on anterior 112 coarsely and densely punctate; carinal stria wanting; posterior
margin of keel outwardly arcuate. Descending lateral stria deeply impressed andcomplete. Epipleura of prothorax densely furnished with long hairs.
Anterior margin of mesosternum deeply emarginate medially; marginal stria
complete and deeply impressed; a short stria present behind antero-lateral angle on eachside; disk wholly covered with coriaceous ground sculpture, and sparsely and finely
punctate. Meso-metasternal suture lightly impressed and feebly angulate at middle.Postmesocoxal stria of metasternum extended along posterior margin of mesocoxa, andbecoming a little more distant from the margin outwards. Lateral metasternal stria
deeply impressed, strongly carinate, obliquely and posteriorly extending, and united
with an oblique stria which is strongly carinate and inwardly extends from the middle ofmetasterno-metepisternal suture. Intercoxal disk of metasternum punctate like that ofmesosternum. Lateral disk densely covered with large, round, shallow and setiferous
punctures on basal 112, the hairs very long; the apical 112 of disk sparsely covered withmoderate punctures which are separated by 5 times their diameter or more.
Intercoxal disk of 1st abdominal sternum wholly covered with coriaceous ground
sculptures, and sparsely clothed with microscopic punctures; lateral stria deeply
impressed and complete on each side.
Protibia with 4 denticles on outer margin, the apical 2 appressed together on
apical angle; 4 small denticles on apical margin of ventral surface. Profemoral stria
nearly complete, usually shortened to basal 1/8, and broadest distant from posterior
margin at middle to apical 1/3. Mesotibia with 3 rows of strong spines on outer margin;
the median one composed of 14 spines; dorsal surface evenly with long hairs. Metatibia
with 3 rows of strong spines. Meso- and metatibiae somewhat dilated.
Male genitalia as shown in Fig. 67.
Female genitalia as shown in Fig. 660.
Specimens examined. [Honshu] Yamagata-ken: 1 ex., Togo-mura, Kita-maruyama-gun,
18/vi/1944. K. Shirahata (NA). Fukushima-ken: 2 exs., Yunohana, Tateiwa, Minami-Aizu. 16,
9/viii/1949, K. Nagayama (NA); 10" 1 ~ ,Ditto, 1O/vi/1950, T. Nakane (NA); 1 ex., Ditto, 8/vii/ 1948,
Y. Kurosawa (NSMT); 2 exs., Numayama, Narahama, 6/vi/1948, Y. Kurosawa (NA); 1 ex., Oyama,
Monden, Wakamatsu, ll1v/1984, Y. Kurosawa (NA); 1 ex., Aizu-Wakamatsu, 9/vi11951, Y. Kurosawa(NSMT); 1 ex., Fukiya-yama, Wakamatsu, 8/vi/1947, Y. Kurosawa (NA); 1 ex., Senaka-aburi,
Higashi-yama, Kita-Aizu, 13/vi/1948, K. Ikarashi (NSMT). Ibaraki-ken: 1 ex., Yoshida, HigashiIbaraki, 19/v/1935, T. Tani (NSMT). Tochigi-ken: 3 exs., Kawaraya, Utsunomiya-shi, 23/vi/1981, K.
Aida, collected from a dead crow (TPM). Chiba-ken: 10" 1 ~ , Oami, 22/vi/ 1984, M. Ohara; 1 ex., Mt.
Kiyosumi, 24/v/1976, M. Tomokuni (NSMT); 1 ~ , Kashiwa, 21/v/1985, N. Kobaysahi. Tokyo-to:
1 ~ , Tokyo, no date and no collector's name (EIHU). Kanagawa-ken: 1 ex., Mt. Oyama, 5/v/1975, H.
Takizawa; 1 ex., Yamakita, 20/ix/1968, M. Ishida; 1 ~ , Matsuda-yama, Tanzawa, 14/ix/1972, M.
Ishida; 10", Akuwa-cho, Yokohama, 27/v/1984, K. Wada; 1 ex, Yokono, Hadano City, 2/vi/1989, Y.
Okushima. Niigata-ken: 10", Y oshigahira, northern Echigo, 25/vi/1954, K. Baba; 10", Sarusawa,northern Echigo, 26/ix/1985, K. Baba. Yamanashi-ken: 1 ex., Koike, 3l1vi/1967, T. Kocha (EIHU); 1
ex., Masutomi, 12/vi/2603 (=1943), no collector's name (NSMT). Nagano-ken: 1 ex., Shimashima,
22/vii/1924, no collector's name. (NSMT); 1 ex., Fujimi-dai, 27/viii/1943, S. Osawa (NA); 1 ~,Nojiri, 21/viii/1943, T. Nakane (NA); 1 ex., Fukushima, Kiso, 23/vii/1947, T. Nakane (NA); 10", 5exs., Kami-Kochi, Shinano, 21-8/vii/1947, S. Osawa. Shizuoka-ken: 1 ~,Gotemba, 1O/viiiI1934, K.
Nakamura (NA); 1 ~, Kida, 6/vil1951, K. Kojima (NA). Aichi-ken: 1 ~, 3 exs., Higashi-yama,
Nagoya, 10/ix/1946, vi/1948, S. Osawa (NA). Mie-ken: 1 ex., Yunoyama, Ise, 31/v/1948, S. Osawa(NA); 1 ~ ,Osugi Valley, 5/vi/1952, T. Horio (NA). Shiga-ken: 1 ex., Hiei-zan, Otsu, 10/x/1989, T.
Ito. Kyoto-fu: 30"0" ,lex., Mizuho, Tozu River, 1O/v/1986, S. Nomura (NA). Osaka-fu: 10" 1 ~,117

Iwawaki-san, southern Osaka, 21vii11980, T. Ito; Id" 1!f, 1 ex., Minoa, 241v11936, K. Sakaguchi(NA); Id", Kongo-san, Chihaya-Akasaka, Kawachi, Il1vi11988, T. Ito. Hyogo-ken: 1 ex., Nose,
201ix11945, F. Takahashi (NA); 10", Nitani-bokujo, Taka, 16IixI1984, T. Ito. Nara-ken: 1 d" 1 ~,Nara, 31viii11946, S. Osawa (NA); 1 ex., Kasuga, 1OIvi/1950, N. Y. (NA). Wakayama-ken: Id" 1 ~,Shingfi, southern Kii, 7/v, Il1viiI1946, S. Osawa (NA); 2 exs., Kushimoto, 241v, 101vi11957, C.
Morishima (NSMT); 1 ~ , Kawayu, 24/viiI1946, S. Osawa. Okayama-ken: 10", Gimba, 291vii11949,
T. Okutani; 1 ex., Mimasaka, viI1913, J. E. A. Lewis (BSM); Id", Yuge, Kumenan, 151vii11989, M.
Hayashi. Oki Is.: 1!f., Saigo-cho, 13/x/1987, T. Kumata. Izu Isles.: 1 ex., near Yakeyama, K6zushima Is., 13lv/1983, T. Niisato and M. Nishimura (NSMT).
[Kyfishfi] Fukuoka-ken: 1 ex., Fukuchi-yama, 4/viiiI1951, F. Takahashi (NA); 1 ex., Ditto,
31IvI1946, R. Matsuda (NSMT); 1 ex., Ditto, 31/v/1946, R. Matsuda (NA); 1 ex., Tashiro,
1OIvi/1952, Y. Miyake (NA); 1 ex., Mt. Hiko-san, Buzen, 27/vii/1947, A. Yamaguchi (NA). Sagaken: 10", Hokuzan-damu, 101viii11973, H. Makihara. Nagasaki-ken: 1 ex., Mt. Unzen, Alt. 2000 ft.,
22Iv/l928, J. E. A. Lewis (BSM). Kumamoto-ken: 1 ex., Touchi, Itsuki, lIvii/1983, M. Maeda.
Miyazaki-ken: 1 ex., Ornata, 15/vii/1954, K. Nohara; 10", Mi-ike, 3/v/1983, S. Mizoguchi.
Kagoshima-ken: 1 ex., Sata, 18/v/1980, M. Miura; 1 d" 1!f, Iriki-bokujo, 20/v/1984, M. Ohara; 1 ex.,Izashiki, Odomari, 21/v/1955, K. Sawada (NA); 1 ex., Imuta, II/vi 1984, T. and T. Nakane (NA); 1 ~,Yae-yama,Koriyama,6/vii/1983, S. Sameshima; 1 ex., Kushikino, 24/v/1982, M. Ohara. Tsushima
Is.: 1!f., Sasuno, 30/v/1968, K. Suga (NA); 10", Hitakatsu, 7/ix/1964, T. Nakane (NA); 5 exs., NitaMeboro, 9/vi/1968, Y. Kurosawa (NSMT); 1 ex., Oboshi-yama, 13/vi/1968, Y. Kurosawa (NSMT); 5
exs., Meboro, 17/vii/1969, Y. Kurosawa (NSMT); 1 ex., Meboro, Mt. Mitake, 15-18/vii/1968, S.
Miyamoto and A. Nakanishi (NSMT); 1 ~, Aoumi, 24/viiI1989, F. Hirakawa.

Distribution (Fig. 71), Japan (Honshu, Shikoku, Kyushu, Tsushima Is" Izu
Isles, Oki Is,); Amurskij Kray; Korea; China; Viet-Nam.
Remarks. This species is allied to Rister congener, but separated from the latterby the striation of elytra, by having 2 spines on the apical most denticle of the protibia,
and by having sparsely furnished hairs on the hypomeron of the pronotum.
Rister congener Schmidt, 1885
(Figs. 68,69,70 and 71)
Hister congener Schmidt, 1885a: 242 [Japonia]; Nakane, 1%3: 70; Kryzhanovskij and Reichardt,
1976: 312; Mazur, 1984: 186 [catalogued]; Hisamatsu, 1985b: 226, pI. 41, f. 5 [noted, photo].
Hister (Paclzylister) congener: Bickhardt, 191Oa: 38 [catalogued].
Hister (Hister) congener: Bickhardt, 1917: 180 [catalogued]; Karniya and Takagi, 1938: 30 [listed].
Paclzylistercongener: Lewis, 1904: 146.

Japancse name: Ibushi-emma-mushi.

Description. Male and female. Body length, PPL, 9.52 - 10.48 mm, PEL, 8.25
- 9.52 mm. Width, 6.83 -7.94 mm. Biometric data are given in Table 12. Body large,
oblong-oval, black and shining; tibiae, tarsi and antennae dark reddish brown.
Frontal stria of head (Fig. 68A) complete, well impressed, carinate and densely
crenate, the anterior portion straight, sometimes narrowly interrupted at middle; disk ofhead even, sparsely clothed with fine punctures which are separated by 3 - 10 times
their diameter, other microscopic punctures intermingled with fine ones. Labrumtransversely oblong, the anterior margin feebly and outwardly angulate. Mandibles welldeveloped.
Pronotal sides (Fig. 68B) nearly parallel on basal 112, and gradually and feebly
convergent on apical 112, the apical ninth especially strongly convergent. Apical angles

118

Fig. 68. Hister congener Schmidt. A: Head, frontal view. B: Pronotum and left e1ytron, dorsal view.
C: Pro-, Meso-, metasterna and 1st abdominal sternum. D: Protibia, dorsal view. E: Ditto,
ventral view. F: Profemur, ventral view. G: Female genitalia, spermatheca and bursa copulatrix,
lateral view Oeft side).
Table 12. Biometric data for Hister congener (Schmidt).

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

3.10-3.57 (3.31±0.06)
6.35-7.30 (6.88±0.12)
2.94-3.49 (3.32±0.08)
4.44-5.08 (4.76±0.09)
6.83-7.94 (7.45±0.15)
4.44-4.92 (4.60±0.07)
2.06-2.54 (2.28±0.06)
2.38-2.86 (2.63±0.06)
2.38-2.86 (2.63±0.06)
2.22-2.70 (2.46±0.06)
3.17-3.49 (3.41±0.O5)

119

6
6
6
6
6
6
6
6
6
6
6

round. Margin of pronotum, except basal margin, densely covered with long hairs.
Marginal pronotal stria complete laterally and broadly interrupted behind head. Outerlateral pronotal stria completely impressed laterally. Inner lateral stria complete anddensely and coarsely crenate. Disk of pronotum sparsely clothed with fine punctures
which are separated by 3 - 5 times their diameter, and microscopic puncturesintermingled among the fine ones. Antescutellar area with a large longitudinal
impression.
Epipleura of elytra feebly excavated. Marginal epipleural stria well impressed on
apical 2/3. Marginal elytral stria strongly carinate and complete. Subhumeral stria (Fig.
68B) stout and shallowly impressed on apical 2/3. Oblique humeral stria lightly
impressed on basal 113. First - 3rd dorsal striae completely impressed and sparsely andcoarsely crenate. Fourth dorsal stria present on apical 1/2 or more, the basal portion
usually represented by coarse punctures. Fifth dorsal stria present on apical 113. Sutural
stria variable, usually present on apical 113, sometimes on apical 112, or absent. Disk of.... : .... : ..
.
.'. '....
........ :',:.
,.'

'

o

11

mm

Fig. 69. Hister congener Schmidt. A: Aedeagus, dorsal view. B: Ditto, lateral view. C: 8th tergite andsternum, dorsal view. D: Ditto, lateral view. E: 9th and 10th tergites and 9th sternum (spicule),
dorsal view. F: Ditto, lateral view.

120Fig. 70. Propygidia (A, C) and pygidia (E, D). A, C: Hister japonicus Marseul. B, D: Histercongener Schmidt.

...

\

\

Hister congener

Fig. 71. Collection sites of Hister japonicus Marseul and Hister congener Schmidt in Japan

elytra evenly covered with fine punctures which are separated by about 4 times theirdiameter.
Propygidium (Fig. 70B) densely covered with large, round and shallow
punctures which are separated by their own diameter; interspace between the large
punctures covered with moderate-sized punctures which are separated by about 1.5 (1 121

2) times the diameter; these punctures becoming finer around margin; disk wholly
clothed with alutaceous ground sculptures. Punctation of pygidium (Fig. 70D) similar
to that of propygidium but much denser; the punctures becoming minute at apex.
Anterior margin of prosternal lobe (Fig. 68C) narrowly truncate apically, its
marginal stria complete and carinate; disk of lobe densely and coarsely punctate, thepunctures becoming finer and sparser medially. Prosternal keel without carinal stria, its
disk even on basal 112; the disk sparsely and finely punctate; lateral descending area on
apical 1/2 densely covered with coarse punctures. Lateral descending stria of keel
complete and carinate.
Anterior margin of mesosternum (Fig. 68C) narrowly and strongly emarginate
medially; marginal stria complete, sparsely and coarsely crenate; another short stria
present behind each antero-lateral angle; disk sparsely clothed with microscopic
punctures except on the area outside the marginal stria, the area being coarsely anddensely punctate. Meso-metasternal suture lightly but distinctly impressed and strongly
angulate at middle. Post-mesocoxal stria of metasternum impressed along posterior
margin of mesocoxa and becoming more distant from the margin laterally. Lateralmetasternal stria well impressed and extending posteriorly and obliquely, the apical end
attaining near apical 113 of metasternal disk. Intercoxal disk of metasternum similar tothat of mesosternum. Lateral disk densely covered with coarse, shallow and setiferous
punctures, and transversely elevated at middle.
Intercoxal disk of 1st abdominal sternum (Fig. 68C) sparsely covered with finepunctures which are separated by 2 - 5 times their diameter; the lateral stria wellimpressed, carinate and shortly abbreviated basally.
Protibia (Fig. 68D, E) with 3 denticles on outer margin, the apical one very large.
Meso- and metatibiae with 2 rows each consisting of about 18 spines which are stout
and long. Profemoral stria nearly complete, shortened on basal 118.
Male genitalia as shown in Fig. 69.
Female genitalia as shown in Fig. 680.
Specimens examined. [Honshu] Aomori-ken: 1 ex., Shiriya, 5/viii/1958, A. Abe (NA).
Yamagata-ken: 1 ex., Yonczawa, 16/v/1944, Y. Kurosawa (NSMT); Id", Ditto, 17/v/191O, S.
Matsumura (EIHU). Fukushima-ken: 1 ex., Tokusa, Tateiwa, 26/v/1991, K. Haga; Id", Yunohana,
Tateiwa, Minami-Aizu, 15/vI1948, W. Hoshi; 1 ex., Ditto, l/v/1948 (NSMT); 1 ex., Odayama,
Monden, Kita-Aizu, 26/vi/1946, Y. Ikarashi (NSwIT); 1 ex., Akai, Kinato, Kita-Aizu, 7/vi/1948, Y.
Kurosawa (NA). Nagano-ken: Id", Hokushin-bokujo, Sugadaira, 10/viii/1963, S. Kondo; Id",
Uminokuchi-bokuj6, Yatsugatake, 18/v/1969, M. Kiuchi; 1 ~, Kuzukawa-Heraka, 4/v/1954, A.
Abe(NA). Hyogo-ken: 1 ex., Mt. Rokko, Kobe, vii/1928, E. R. Dickover (BSM); 1 ex., Ditto,
16/vi11929, E. R. Dickovcr (BSM); 1 ex., Ditto, 6/vii/1928, F. C. Hadden (BSM); 1 ex., Kobe,
6/vii/1930, E. R. Dickover (F. C. Hadden collection), Hister congener Schmidt, H. Desbordes det.,
1932 (BSM). Tottori-ken: 1 ex., Hyonosen, 12/vi/1961, K. Tsuji (NSMT).
[Korea] 1 ex., Mt. Kongo, 28/vii/l924, K. Sato (NSMT).

Distribution (Fig. 71). Japan (Honshu); Primorskij Kray; Korea; Taiwan;
northern China.
Remarks. Lewis (1904) treated this species as a member of the genus
Pachylister on the basis of the following states interpreted by him as generic; protibiae
with 3 large denticles on outer margin, and anterior margin of labrum outwardly
projecting. But since Bickhardt (1917) this species has been treated as a member ofHister, because the anterior margin of the labrum of this species is very slightly
projecting. These generic characteristics are unclear and problematical. This species isallied to Hister japonicus (see H. japonicus).
122Hister simplicisternus Lewis, 1879
(Figs. 72, 73, 75, 76)Hister simplicisternus Lewis, 1879: 461 [Japan: Higo, where it is common]; Olexa, 1982b: 198 [key,figures]; Mazur, 1984: 197 [catalogued]; Hisamatsu, 1985b: 227, pI. 41, no. 8 [noted, photo].
Hister simplicisternis (sic): Kryzhanovskij and Reichardt, 1976: 313 [description, figures].
Hister (Hister) simplicisternus: Bickhardt, 191Oa: 49 [catalogued]; Bickhardt, 1917: 186 [catalogued].
Hister togoii Lewis, 1914b: 286 [Tsushima Is.], synonymized by Bickhardt, 1920: 30.
Hister togoi (sic): Bickhardt, 1917: 187 [catalogued]; Bickhardt, 1920a: 30.

Japanese name: Hime-tsuya-emma-mushi.

Description. Male and female. Body length, PPL, male, 4.09 - 5.57 mm,
female, 4.62 - 6.14 mm, PEL, male, 3.43 - 4.43 mm, female, 3.86 - 4.95 mm.
Width, male, 3.05 - 4.00 mm, female, 3.33 - 4,17 mm. Biometric data are given in
Table 13. Body oval, rather convex, black and shining; tibiae, tarsi and antennae
rufopiceous.
Frontal stria of head (Fig. 72A) deeply impressed and carinate, the anterior
portion broadly straight. Disk of head feebly depressed inside the stria which issparsely clothed with moderate punctures separated by about 3 times their diameter.
Pronotal sides (Fig. 72B) arcuate and strongly convergent forward. Apical angle
round. Anterior margin of pronotum broadly emarginate, the median portion ofemargination feebly arcuate outwardly. Marginal pronotal stria complete laterally andbroadly interrupted behind head. Outer lateral pronotal stria abbreviated on basal 113;
feebly and sparsely crenate. Inner lateral pronotal stria abbreviated on basal 116, anddensely and strongly crenate. Disk of pronotum sparsely clothed with fine punctures
which are separated by 4 - lO times their diameter; a narrow band along posterior
margin covered wi th large and round punctures.
Epipleuron of elytron (Fig. 72B) broad and even, and with 3 striae; the outermost
stria clearly impressed on apical 112; median one (epipleural marginal stria) nearlycomplete and carinate, but shortly reduced basally; inner one (marginal elytral stria)
complete and strongly carinate. Lateral margin of elytron strongly carinate. External
subhumeral stria shortly present medially as an arc, and deeply impressed. Oblique
humeral stria lightly impressed on basal 1/3. First - 3rd dorsal striae complete, deeply
impressed and sparsely and coarsely carinate. Fourth dorsal stria present on apical 114,
with a short rudiment basally. Fifth dorsal stria present on apical 118 but indistinct.
Sutural stria present on apical 112. Disk sparsely clothed with fine punctures which areseparated by about lO times their diameter.
Propygidium (Fig. 75A) unevenly and sparsely covered with large, round anddeep punctures which are separated by 0.5 - 2 times their diameter; surface between
large punctures sparsely and finely punctate; area between these punctures withalutaceous ground sculptures. Punctation of pygidium (Fig. 75E) similar to, but muchfiner than, that of propygidium, and the punctures becoming finer apically.
Anterior margin of pronotal lobe (Fig. 72C) round, its marginal stria deeply
impressed, complete and carinate; disk densely and coarsely punctate, the punctures
separated by about their own diameter and becoming sparser basally. Prosternal keelrather broad, even, without carinal striae; disk coarsely punctate on basal 112, thepunctures separated by about 1.5 (1 - 2) times their diameter; basal margin of keel
nearly straight. Descending lateral stria deeply impressed, carinate and complete.
123

Table 13. Biometric data for Hister simplicisternus Lewis.
Female

Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

1.00-1.43 (1.28±0.02) 20
2.9S-3.71 (3.37±0.OS) 20
I.S2-1.9S (1.70±0.02) 20
2.0S-2.67 (2.36±0.04) 20
3.33-4.17 (3.84±0.06) 20
2.09-2.71 (2.39±0.04) 20
0.71-1.0S (0.9S±0.02) 20
1.0S-1.38 (1.23±0.02) 20
1.0S-1.43 (1.19±0.02) 20
1.09-1.38 (1.24±0.02) 20
1.38-1.71 (1.S7±0.OS) 20

1.07-1.43 (1.24±0.08) 20
2.S7-3.S2 (3.09±0.OS) 20
1.29-1.76 (1.S6±0.03) 20
1.81-2.38 (2.14±0.03) 20
3.0S-4.00 (3.S6±0.06) 20
1.81-2.48 (2. 14±0.04) 20
0.71-1.0S (0.89±0.04) 20
0.90-1.29 (1.13±0.02) 20
0.90-1.24 (1.09±0.02) 20
0.90-1.24 (1.12±0.02) 20
1.24-1.71 (1.48±0.03) 20

Fig. 72. Hister simplicisternus Lewis. A: Head, frontal view. B: Pronotum and left eIytron, dorsal
view. C: Pro-, meso-, metasterna and 1st abdominal sternum. D: Protibia, dorsal view. E:
Ditto, ventral view. F: Profemur, ventral view. G: Female genitalia, spermatheca and bursa
copulatrix, lateral view (left side).

124A---(;//--':~~,~'-

...

----

......'- .. :....
... ".
,'-. . . 'IO.5mm.

'..

.......

/-.-----

-

,',

0

':'<,'~

F

Fig. 73. Hister simplicisternus Lewis. Male genitalia. A: Aedeagus, dorsal view. B: Ditto, lateralview. C: 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: 9th and 10th tergites,
dorsal view. F: Ditto, lateral view. G: 9th sternum (spicule), ventral view.

Anterior margin of mesosternum (Fig. 72C) nearly straight, feebly emarginate
medially; marginal stria deeply impressed, strongly carinate, sparsely and coarsely
carinate, and complete; 2 short striae present behind the anterolateral angle on each side,
the posterior stria united with the marginal stria; disk sparsely and finely punctate, thepunctures separated by 10 times their diameter; another microscopic puncturesintermingled with them. Meso-metasternal suture distinctly impressed, and angulate at
middle. Post-mesocoxal stria of metasternum deeply impressed and carinate, andbecoming more distant from posterior margin of mesocoxa on lateral side; the outer endof stria attaining near the middle of metasternal-mesepimeral suture. Lateral metasternal
stria extending obliquely and posteriorly, beginning from lateral 114 of mesometasternal suture, deeply impressed and carinate, and not united with oblique stria
which is strongly carinate and inwardly extends from the middle of metasternalmetepisternal suture. Intercoxal disk of metasternum sparsely and finely punctate, thepunctures separated by about 8 times their diameter. Lateral disk densely covered withlarge, round, shallow and setiferous punctures, the punctures becoming finer apically;hairs of the punctures long.
Intercoxal disk of 1st abdominal sternum (Fig. 72C) sparsely and finely punctate,
the punctures coarser before apical angles; lateral stria complete, densely and coarsely
crenate.
Protibia with 5 denticles on outer margin. Profemur with femoral stria shortly
present on apical 114.
Profemur with femoral stria shortly present on apical 1/4.
Male genitalia as shown in Fig. 73.
Female genitalia as shown in Fig. 720.
125Specimens examined. [Honshu] Aomori-ken: 2 exs., Higashi-dori, Shiriya-zaki, 6lvi,
261vii11987, H. Saito; 2 exs., Shiriya, 51viii11958, A. Abe (NA). Iwate-ken: 4 exs., Kuriya-gawa,
201v11974, H. Hayakawa (NA). Miyagi-ken: 1 ex., Mt. Saho, Sendai, 21vi11951, K. Nagayama (NA).
Yamagata-ken: 1 ex., Sakata, lIviiI1948, T. Nakane (NA). Fukushima-ken: 1 ex., Wakamatsu,
29IviI1948, T. Nakane (NA); 3 exs., Ditto, 211v11949, Y. Kurosawa (NSMT); 1 ex., Hinoemata,Aizu, 11viii11947, K. Nagayama (NA); 1 ex., Nagurazaka, Asahi-dake, Minami-Aizu, lIviiil1949, K.
Nagayama (NSMT); 1 ex., Aizu-Wakamatsu, 141vi11950, Y. Kurosawa (NSMT); 1 ex., Yunohana,
Tateiwa, Minami-Aizu, 1OIviiI1948, Y. Kurosawa (NSMT); 3 exs., Oyama, Monden, Kita-Aizu,
211v11949, Y. Kurosawa (NSMT); 1 ex., Tsuchiyu, no date, S. Omomo. Ibaraki-ken: 1 ex., Yatabe,
51vii11986, M. Kiuchi. Tochigi-ken: 3 exs., Kawaraya-machi, Utsunomiya-shi, 261vi, 31vii11981, K.
Aida, collected from a dead crow (TPM); 1 ex., Chikushi-yama, Karasuyama-machi, 221vi11987, K.
Satoh(TPM). Chiba-ken: 1d" 1!f, 1 ex., Oami, 19, 221vi11984, M. Ohara; 1d", Abiko, Tone-gawa,
51v11973, M. and K. Ishida; 1 ex., Narita, 91vi11914, E. Gallois (ElHU). Tokyo-to: 8 exs., Tokyo, 29,
30, 311v, 31vi11914, E. Gallois (ElHU); 1 ex., Komaba, Tokyo, 241v11914, S. Hirayama (EIHU); 1
ex., Tokyo, no date, S. Matsumura (EIHU); 1 ex., Kinuta, Setagaya, 151v11988, Y. Kurosawa(NSMT); 1 ex., Rokugien, 91v11953, Y. A. (NSMT); 1d", 1 ex., Ogikubo, 5lvii, 301viii11966, M.
Ishida; 1 ex., Aoyama, 27IvI1946, T. Nakane (NA); 1 ex., Kawajiri, 23IviI1940, T. Okada (NA);
2!f!f, 3 exs., Toshima-ku, 191v11963, S. Kondo; 2!f!f , Fuchu, 111vi11968, M. Ishida; 3 exs.,
Sayama, lllv11957, M. Suwa. Kanagawa-ken: 3d"d", 2 exs., Akuwa-cho, Yokohama, 71vi11983, 25,271v, 12, 15IviI1984, K. Wada; 1d"1!f, Fuchinobe, Sagamihara, 71v11973, M. Ishida; Id"1!f,Daiyu-zan, 291v11982, M. Nishikawa. Niigata-ken: 1 ex., Maki, Matsumura, northern Echigo,
71vii11987, K. Baba. Toyama-ken: 1 ex., Tateyama-machi, 211viiI1954, S. Takagi (ElHU).
Yamanashi-ken: 1 ex., Nirasaki, 181v11986, N. Kobayashi. Nagano-ken: 1!f, Koma-ga-take, Kiso, 45IviiiI1946, S. Osawa (NA); 1d", Nojiri, Toyo-eiwa, 211vii11941, T. Nakane (NA); 1 ex., Fujimi,
201vii11956, S. Nomura (NSMT); 1 ex., Sanjiro, Utsugushi-ga-hara, 51viiil 1951, S. Ueno (NA). Gifuken: 2!f!f , Ogaki, Mino, 101 vi 1946, Y. Moriya (NA). Shizuoka-ken: 1 ex., Kida, 61vi/1951, K.
Kojima (NA); 1 if-, Asagiri-Kogen, Fuji, 800 m, 23IviiiI1972, M. Ishida; 1 ex., Yawatano, Ito,
15IviI1977, M. Ishida. Aichi-ken: 4d"d" 1!f , 4 exs., Higashi-yama, Nagoya, 23, 27IviI1948, S.
Osawa (NA); Id" 2!f !f , 2 exs., Shikatsu-mura, Owari, 191viii11945, 271vi, 16IviiiI1946, S. Osawa(NA); Id" 1!f, Nagoya, 101v, 41viii11947, S. Osawa (NA). Mie-ken: 1 ex., Osugi, 121vi11952, T.
Horino (NA); 1 ex., Osugi-dani, Ise, 101vi11952, Takeuchi (NA). Kyoto-fu: 1 ex., Kyoto, 1956, T.
Nakane (NA). Osaka-fu: 1 ex., Myoken, 221vi11952, F. Takahashi (NA); 1!f , Iwawaki-san, southern
Osaka, 21vii11988, T. Ito. Wakayama-ken: 1!f, Shingu, Kii, v11946, S. Osawa (NA); Id", Nachi, Kii,
16IviiI1946, S. Osawa (NA). Hyogo-ken: 1 ex., Motoyama-mura, Kobe, 141v11938, K. Sakaguchi(NA); 1 ex., Kobe, 301vii11916, J. E. A. Lewis (BSM); 1 ex., Harada, Kobe, 15IviiiI1915, nocollector's name (BSM); 1 ex., Ditto, 201vi11912, J. E. A. Lewis (BSM); 1!f, Mitani-bokujo, Taka,
161ix11984, T. Ito.
[Kyushu] Fukuoka-ken: 1 ex., Fukuchi-yama, 41viii11951, F. Takahashi (NA). Kumamoto-ken:
1!f ,Komezuka, Aso, 71vi11981, M. Ohara; 1 d" 1!f ,Mt. Maru, Takarnori, Aso, 6lvi/ 1986, E. Matsui;Id", Nabeno-daira, Aso, 5/viI1986, E. Matsui. Oita-ken: 1!f, Kuju, 61v/1981, K. Sakai. Kagoshimaken: 1!f, Kurino, Kirishima, 251vi11983, K. Sugawara; 1!f ,Ditto, l/vi/ 1981, M. Ohara; 1 d", Ditto,
3/ixI1990, M. Ohara; Id", Iriki, 201v11984, M. Ohara.

Distribution (Fig. 76). Japan (Honshu, Shikoku, Kyushu); Korea.
Remarks. This species is easily distinguished from all the other Japanese speciesof the tribe Histerini by the deeply impressed external subhumeral stria and by thefeeble and outward projection on the anterior margin of pronotum.

Hister unicolor leonhardi Bickhardt, 1910
(Figs. 75)
Hister leonhardi Bickhardt, 191Ob: 180 [East Siberia].
Hister unicolor abo leonhardi: Reichardt, 1938: 236.

126

Hister unicolor opimus Reichardt in Kryzhanovskij and Reichardt, 1976: 316 [USSR: Zabaikal'e andPrimorskij; Japan: Unzen], synonymized by Mazur, 1984: 199.
Hister unicolor leonhardi: Mazur, 1984: 199.

Japanese name: Nise-kuro-emma-mushi.

Distribution. Japan (KyiishG: Shimabara); East Siberia; Primorskij Kray;
Mongolia; Korea; North-eastern China.
Hister concolor Lewis, 1884
(Figs. 74, 75, 76)Hister conc%r Lewis, 1884: 135 [Japan: Ishikari river, Hokkaido].
Hister (Hister) conc%r. Bickhardt, 191Oa: 42 [Japan]; Bickhardt, 1917: 197; Karniya and Takagi,
1938: 30; Osawa, 1952: 6.
Hister conc%r: Kryzhanovskij and Reichardt, 1976: 316; Hisamatsu and Kusui, 1984: 16 [noted];
Hisamatsu, 1985b: 227, pl. 41, f. 7 [noted, photo]; Ohara and Li, 1988: 96 [China: Liaoning].

Japanese name: Kuro-emma-mushi.

Description. Male and female. Body length, PPL, male, 6.31 - 8.00 mm,
female, 6.44 - 8.69 mm, PEL, male, 5.83 - 7.00 mm, female, 5.63 - 7.50 mm.
Width, male, 4.94 - 6.19 mm, female, 5.00 - 6.63 mm. Biometric data are given in
Table 14. Body oval and black; spines of tibiae, tarsi, antennae and mouthparts except
mandibles reddish brown.
Frontal stria of head (Fig. 74A) carinate and usually interrupted at middle,
sometimes complete, the anterior portion feebly and inwardly arcuate and densely andcoarsely crenate; disk evenly and finely punctate, the punctures separated by 3 - 4 times
their diameter, and interspace between the fine punctures sparsely and microscopicallypunctate. Labrum transversely oblong.
Pronotal sides (Fig. 75B) regularly arcuate, strongly convergent forward on
apical 2/3 and convergent posteriorly on basal 113. Apical angles angulate. Marginal
pronotal stria clearly impressed and complete laterally, and broadly interrupted behind
head. Outer lateral pronotal stria deeply impressed, crenate and abbreviated on basal 113- 112. Inner lateral stria deeply impressed, abbreviated on basal 113, and the anterior
portion complete and densely and coarsely crenate. Disk of pronotum sparsely clothedwith microscopic punctures which are separated by 5 times their diameter or more, andwholly covered with light coriaceous ground sculptures; lateral 113 of a narrow band
along the posterior margin coarsely and densely punctate. Antescutellar area with ashort longitudinal puncture.
Epipleural fossette of elytra feebly excavated. Marginal epipleural stria slightly
carinate and present on apical 112. Marginal elytral stria deeply impressed, carinate andcomplete. External subhumeral stria (Fig. 74B) present on medio-apical 114 in an arc,and broadly and shallowly impressed and crenate. Internal subhumeral stria broadly andshallowly impressed on apical 112, and densely and coarsely crenate. Oblique humeral
stria lightly impressed on basal 112. First - 3rd dorsal striae complete, broadly andshallowly impressed and densely crenate. Fourth dorsal stria usually present on apical
112, sometimes on apical 2/3. Fifth dorsal stria variable in length, usually present on
apical 114, sometimes on apical 113 or 115. Sutural stria variable, usually present
127Table 14. Biometric data for Hister cone%r Lewis.
Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Female1.69-2.06 (1.88±0.1O) 20
4.25-5.38 (4.77±0.08) 20
1.75-2.50 (2. 17±0.04) 20
3.06-3.88 (3.44±0.06) 20
4.94-6.19 (5.60±0.09) 20
2.94-3.75 (3.33±0.05) 20
1.06-1.44 (1.25±0.02) 20
1.50-1.88 (1.64±0.03) 20
1.31-1.81 (1.56±0.03) 20
1.31-1.88 (1.63±0.03) 20
1.69-2.50 (2.08±0.04) 20

1.69-2.19 (2.00±0.03) 20
4.19-5.69 (5.18±0.09) 20
1.81-2.63 (2.33±0.05) 20
3.06-4.13 (3.72±0.07) 20
5.00-6.63 (6. 1O±0. 10) 20
2.94-4.13 (3.70±0.08) 20
1.19-1.63 (1.40±0.03) 20
1.38-2.06 (1.85±0.04) 20
1.44-1.88 (1.69±0.03) 20
1.31-2.00 (1.76±0.04) 20
1.88-2.63 (2.31±0.05) 20

apically, a little longer than 5th. Disk of elytron wholly covered with coriaceous ground
sculptures and sparsely clothed with fine punctures which are separated by 5 - 10 times
their diameter, the punctures becoming coarser on apical 1/6.
Propygidium (Fig. 75D) densely covered with large, round and shallow
punctures, which are separated by 0.75 (0.5 - 1) times the diameter and becoming finer
around margin; interspace between the large punctures intermingled with moderatesized punctures which are as large as those of apical 116 of elytron; surface, except for
punctures, wholly covered with alutaceous ground sculptures. Punctation of pygidium(Fig. 75H) similar to that of propygidium, but much denser than the latter, thepunctures separated by 0.3 - 0.5 times their diameter.
Anterior margin of prosternal lobe (Fig. 74C) round; marginal stria broadlyinterrupted at middle; disk sparsely and coarsely punctate, the punctures separated byabout 4 times their diameter and becoming finer medially. Prosternal keel even andsomewhat broad, and without carinal stria; the posterior margin arcuate outwards; disk
evenly covered with fine punctures which are separated by about 3 times their diameteron median area; lateral descending area with coarse and dense punctures. Descending
lateral stria strongly carinate and complete.
Anterior margin of mesosternum (Fig. 74C) strongly emarginate at middle;
marginal stria complete and carinate; a short and broad stria present behind each anterolateral angie; disk sparsely and finely punctate, the punctures separated by about 3 - 5times their diameter and becoming coarser laterally. Meso-metasternal suture clearly
impressed and angulate at middle. Post-mesocoxal stria of metasternum extending alongposterior margin of mesocoxa, and becoming a little more distant from the margin, theouter end attaining at middle of metasternal-mesepimeral suture. Lateral metasternal stria
deeply impressed, strongly carinate, obliquely extending posteriorly, and united withoblique stria, which is strongly carinate and inwardly extends from the middle ofmetasternal-metepisternal suture. Punctation of intercoxal disk similar to that ofmesosternum, the punctures becoming somewhat coarser laterally. Lateral disk densely
covered with large, deep and round punctures which are separated by 0.3 - 0.5 times
their diameter; interspace between large punctures usually coarsely punctate; thesepunctures becoming finer inwardly.

128

Fig. 74. Hister concolor Lewis. A: Head, frontal view. B: Pronotum and left elytron, dorsal view. C:
Pro-, meso-, metastema and 1st abdominal sternum. D: Protibia, dorsal view. E: Ditto, ventral
view. F: Profemur, ventral view. G: Female genitalia, spermatheca and bursa copulatrix, lateralview (left side).

Intercoxal disk of 1st abdominal sternum evenly covered with moderate-sizedpunctures which are separated by about 3 times their diameter, the punctures becomingcoarser laterally; lateral stria deeply impressed, carinate and complete.
Protibia (Fig. 740, E) with 3 or 4 denticles on outer margin, and with 3 or 4 on
apical margin. Meso- and metatibiae with 2 rows each consisting of 6 - 8 stout spines.
Profemoral stria (Fig. 74F) present on apical 113 - 112.
Male genitalia as Fig. 76.
Female genitalia as Fig. 740.
Specimens examined. [Hokkaido] 4 exs., Hamayftchi, Wakkanai, 23/viii/1993, S. Hori (HOR);
1 ex., Sakanoshita, Wakkanai, 10/viiil 1974, Y. Kusui; 1 ex., Horonobe, 30/vii/1988, Y. Ishihama; 7exs., Futatsu-yama, Shibecha, Kushiro, 20/ix/1955, 15/x/1970, 26/ix/1974, 24/vi/1979, 15/viii/1981,
K. Ijima (11); 1 ex., Ikutahara, near Abashiri, 7/vii/1987, M. Ohara; 1 ex., Kenebetsu, Nemuro,
211vii/1973, M. Nishimura; 1 ex., Gensei-kaen, Abashiri, 19/vii/1986, M. Ohara; 1 ex., NakaTombetsu-cho, Kami-Tombetsu, 27/vii/1987, K. Haga; 2 exs., Kawakami, Rikubetsu, 311vii11988, K.
Haga; 5 exs., Onne-shisui-damu, Minami-Kitoushi, Ashoro, 9/vii/1991, K. Haga; 1 ex., Satsunai,
8/viii/1968, H. Torikura; 20'0' 4~ ~ ,lex., Hitsuji-ga-oka, Sapporo, 20/vii/1985, M. Ohara; 10',
Moiwa, Sapporo,4/vi/1953, T. Kumata(EIHU); 1~, 1 ex., Sapporo, 9/vi/1975,M. Kiuchi; 2!j?!j?, 1
ex., Ditto, 19/v, 71vi11952, K. Houma (EIHU); 7 exs., Nopporo, 5, 121vi, Ilviiil1987, M. Ohara; 1

129

Fig. 75. Propygidia (A, B, C, D) and pygidia (E, F, G, H). A, E: H. simplicisternus Lewis. B, F: H.
unicolor leonhardi Bickhardt. C, G: H. u. unicolor Linnaeus. D, H: H. concolor Lewis.

ex., Ishikari, 7/viiiI1978, T. Fujisawa (EIHU); 1 ex., Asari, Otaru, 18/vi/1978, T. Fujisawa (EIHU);
1 ~, 1 ex., Rokushinai, Shakotan, 22, 23/ixI1985, M. Mori; 10", 1 ex., Otobe, 15, 22/ixI198S, M.
Mori; 1 ~ , 2 exs., Ditto, 2/viii/1973, T. Kumata (EIHU); 1 ex., Tomakomai, 8/vi/1986, M. Mori;
20"0" 3 ~ ~ , Bibi, 8/vi/1986, M. Mori; 3 exs., Iwanai, 9-10/vii/1990, M. Ohara; 1 ex., Toyohama,
Shimamaki, 8/vii/1988, M. Ohara, M. Sato et N. Ishihama; 3 exs., Kuromatsunai, 6, 25/vi/1986, M.
Mori; 2 exs., Ditto, 8/viii/1988, M. Ohara; 1 ex., Kita-Hiyama, 28/vI1988, M. Ohara. Rishiri Is.:10" 2 ~ ~ , Oshidomari, 4/viil1974, Y. Kusui.
[HonshU] Aomori-ken: 1 ex., Shiriya, 5/viii/1958, A. Abe (NA); 1 ~, Heraka, Towada,
27/vii/1958, A. Abe (NA); 1 ex., Onabe, Towada, 27/vii/1958, A. Abe (NA); 1 ex., Shichinobe,
17/vi/1974, H. Hayakawa (NA); 10"2~ ~ , Ohata, Shimokita, 26/viilI983, S. Yamauchi; 3 exs.,
Mutsu, Nakotani, 19/vi/1987, H. Saito; 1 ~, Ditto, K. Shimoyama. Iwate-ken: 2 exs., Morioka,
18/vi/1944, no collector's name (NA); 1 ex., Nishine, 28/vi/1974, H. Hayakawa (NA); 10", 1 ex.,
Kuriya-gawa, 20/v/1974, H. Hayakawa (NA). Fukushima-ken: 1 ex., Hinoemata, Minami-Aizu,
15/vii/1965, K. Tazoe. Tochigi-ken: 4 exs., Nikko, 23/viii/1958, M. Suwa; 20"0" , 3 exs., Ozan
River., 5/viii, l/ixI1981, H. Tanaka; 1 ex., Akada, Nishinasuno-machi, 8/viii/1991, S. Ohmomo

130

.. ~

..

D
I

0.5 mmFig. 76. Hister concolor Lewis. Male genitalia. A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: 9th and 10th tergites and 9thsternum (spicule), dorsal view. F: Ditto, lateral view.

Fig. 77. Collection sites of Hister simplicisternus Lewis and H. concolor Lewis in Japan.

131

(TPM); 1 ex., Duns, Shikamihira, 27/vii/1985, T. Kumazawa (TPM); 1 ex., Kotoku, Nikko-shi,
9/x/1972, T. Miyamoto (TPM); 11 exs., Shiobara-machi, 29/viii/1979, T. Tanaka (TPM); 2 exs.,
Ozasa-bokujo, Imaichi-shi, 25/viii/1991, S. Ohmomo (TPM); 3 exs., Ozasa, Imaichi-shi, 4/x/1981, K.
Aida, collected from cow dung (TPM); 5 exs., Ozasa, Imaichi-shi, 4/x/1981, T. Miyamoto (TPM); 1
ex., Ditto, 25/vii/1981, K. Aida (TPM). Tokyo-to: 1!f, Futatsu-yama, Oku-Chichibu, 2/ix/1939, K.
Kojima (NA). Kanagwa-ken: 1 ex., Kashiwabara, Tanzawa, 13/viii/1920, Y. Yamamoto (NA). Niigataken: 1!f, Kurokawa, Echigo, 1O/viii/1957, K. Baba. Nagano-ken: lei" 1!f, 2 exs., Togakushi,
27/vii/1978, T. Fujisawa (EIHU); 1 ex., Sugadaira, 24/vii/1978, T. Fujisawa (EIHU); 1 ex.,
Hokushin-bokujo, Sugadaira, 15/viii/1963, S. Kondo; 1 ex., Mt. Yatsu, 20/vii/1951, no collector's
name (NSMT); 1 ex., Inako-bokujo, Mt. Yatsu, 26/vii/2602 (=1942), no collector's name (NSMT);
lei", 2 exs., Tateshina, 15/viii/1966, K. Masumoto; 3ei"ei" 5!f!f , 2 exs., Kita-Karuizawa, 242S/viiiI1937, K. Sakaguchi (NA); 1 ex., Karuizawa, 11-15/viii/1944, H. Inoue (NA); lei"l!f,Kiyosato, 22, 23/vii11981, M. Ohara; lei", Saku, Meshimori-yama, 3/x/1973, M. Ishida (NA);lei"2!f!f ,Ogawa River, Kawakami ViII., 17/viii/1985, H. Tanaka; 3 exs., Ditto, 211ix/1985, K.
Haga; 1 ex., Hirasawa-bokujo, Meshimori-yama, 26/viii/1984, K. Haga; lei", 2 exs., Nobeyama,
Minami-Saku, 18/viii/ 1972, M. Kiuchi; 1 ex., Asama-Kogen, 1O/viiiI1967, M. Ishida. Gifu-ken: lei",
1 ex., Hirayu, 23-24/vii/1946, M. Hayashi. Kyoto-fu: 1 ex., Kashiwabara, Tamba, 13/viii/1950, Y.
Yamamoto (NA). Hyogo-ken: 4 exs., Mitani-bokujo, Taka, 16/ix/1984, T. Ito.

Distribution (Fig. 77). Japan (Chishima Isles, Hokkaid6, Honshu); Continental
China.
Remarks. This species is closely related to Hister unicolor, but distinguished by
the dense and coarse punctation of the pygidium. It usually occurs in fresh cow dung,and sometimes under the carrion.
GENUS ZABROMORPHUS LEWIS, 1906
Zabromorphus Lewis, 1906a: 399 [type species: Rister pachysomus Ancey, 1882: 55. Originallydesignated].

This genus is closely related to the genus Hister Linnaeus in the characteristics
of paramera, median lobe and spermatheca, with 21 species in the world.
Zabromorphus punctulatus (Wiedemann, 1819)(Fig. 78, 79)
Rister punctulatus Wiedemann, 1819: 162 [Java]; Marseul, 1854: 256, t. 7, f. 60; 1873: 220 [noted,
Japan: Kyushu]; Nakane, 1981: 9 [listed].
Hister (Hister) punctulatus: Bickhardt, 1910a: 47 [catalogued]; Desbordes, 1919: 389 [Jnde, Indechine, lie Quelparert, Java].
Hister (Zabromorphus) punctulatus: Bickhardt, 1917: 177 [catalogued]; Kamiya and Takagi, 1938:
29 [listed].
Zabromorphus punctulatus: Mazur, 1984: 166 [noted of subsp. salebrosus]; Mazur, 1984: 204[catalogued]; Hisamatsu, 1985b: 227, pI. 41, f. 9 [noted, photo].

Japanese name: Arame-emma-mushi.

Description based on a specimen collected from Owari (=Nagoya), Honshu.
Male. Body length, PPL, 6.84 mm, PEL, 5.70 mm. Width, 4.28 mm. Body oblong,
black and shining.

132

F~
Fig. 78. Zabromorphus punctulatus (Wiedemann). A: Head, frontal view. B: Pronotum and left
elytron, dorsal view. C: Pro-, meso-, metastema and 1st abdominal sternum. D: Protibia, dorsal
view. E: Ditto, ventral view. F: Profemur, ventral view. G: Propygidium and pygidium.

Frontal stria of head (Fig. 78A) complete. Disk within the stria densely andcoarsely punctate. Clypeus densely covered with coarse punctures. Mandible welldeveloped, its surface feebly concave.
Pronotal marginal stria (Fig. 78B) interrupted behind head, complete laterally.
Outer lateral stria complete, close to lateral margin. Inner lateral stria complete, strongly
and inwardly sinuate at apical 113 on lateral side, close to the margin behind head.
Punctation of surface strong, dense and deep, lacking on posteromedian area, thepunctures occurring anteriorly on the median area dense, round and deep, those on
lateral area dense, continued with each other and rugose. Antescutellar area of disk witha longitudinal puncture.
Epipleural fossette of elytra densely punctate. Marginal epipleural stria present on
apical 2/3. Marginal elytral stria complete. External subhumeral stria (Fig. 78B)
consisting of deep punctures and present on basal 112. Internal subhumeral stria deep,
present on apical 2/3. Oblique stria indistinctly impressed on basal 113. First - 4th
dorsal striae complete, the 1st and 2nd strongly sinuate at basal 113. Fifth dorsal stria
present on apical 113. Sutural stria present on apical 114. Disk of elytra densely covered
with large and deep punctures inside internal subhumeral stria, except on basal 2/3
between 4th dorsal and suture, a basal humeral area, a median area between 1st and 2nd
dorsal striae, basal and apical small areas between the 2nd and 3rd, and an apical area
between the 4th and 5th. These impunctated areas often reduced or broadened.
Propygidium (Fig. 780) densely covered with coarse punctures, which areabsent in the middle of the apical area. Pygidium densely covered with large and round133

punctures, except on area along the lateroposterior margin and on an oblong apical areaon the meson, which are impunctate.
Prosterna! lobe (Fig. 78C) round anteriorly, its marginal stria complete.
Prosternal keel rather narrow, without carina! stria. Epipleura of prothorax with long
hairs.

G,1 mm

Fig. 79. Zabrorrwrphus punctulatus (Wiedemann). Male genitalia. A, E: Aedeagus, dorsal view. B, F:
Ditto, lateral view. C: Median lobe, dorsal view. D: Ditto, lateral view. G: 8th tergite, dorsal
view. H: Ditto, lateral view. I: 8th sternum, dorsal view. J: Ditto, lateral view. K: 9th and 10th
tergites and 9th sternum (spicule), dorsal view. L: Ditto, lateral view. A - D: Specimen
collected in Aichi-ken, Japan. E - L: in Philipines.

134

Table. 15. Biometric data of Zabromorphus punctulatus (Weidennann).

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MIlL
PEL
PPL

Owari

OsakaA. P.

Taiwan (n=2)

Philippines

2.00
4.66
2.85
4.28
5.04
2.95
1.33
1.71
1.90
1.71
2.38
6.65
7.60

1.62
3.99
2.38
3.33
4.28
2.66
1.05
1.43
1.90
1.43
1.71
5.70
6.84

2.09
4.92-5.23
3.33
3.33-3.61
5.23-5.61
3.33-3.52
1.43
1.90
1.90-2.09
1.71
2.19-2.38
7.13-7.41
8.27-8.65

1.90
4.37
2.85
4.66
4.66
2.95
1.24
1.62
1.81
1.71
2.19
6.27
6.94

Anterior margin of mesosternum (Fig. 78C) feebly emarginate at middle.
Marginal stria of mesosternum complete. A short stria impressed behind eachanterolateral angle. Disk of mesosternum sparsely and coarsely punctate laterally.
Meso-metasternal stria complete, strongly angulate at middle. Lateral metasternal stria
extending posteriorly, united with an oblique stria which inwardly extends from the
middle of the metasternal-metepisternal suture. Lateral disk of metasternum sparselywith large and round punctures, and with short hairs. Intercoxal disk of 1st abdominal
sternum completely striate on each side.
Protibia (Fig. 78E, F) with 4 teeth on outer margin, the apical 2 large, and apicalmargin without teeth. Dorsal surface of protibia with a stria along the base of teeth, theoutside stria densely punctate. Profemoral stria (Fig. 78F) complete and carinate.
Male genitalia as shown in Fig. 79.
Specimens examined. [Honshu] Aichi-ken: 10", Shikatsu-mura, Owari, 27/vi/1948, S. Osawa(NA).Osaka-fu: 1 ex., Osaka Air Port, Osaka, 6/vi/1991, Y. Kusui (introduced ?).
[Taiwan] 1 ~, 1 ex., Taizhong, 5/iv/1907, S. Matsumura (EIHU).
[philippines] Luzon Is.: 10", Asin hot spring, Luzon, 18/viii/1976, K. Matsuda.

Distribution. Japan (Honshu, Kyushu); Taiwan; Indochina; Java, Philippines.
New to the Philippines.
Remarks. This species is easily recognized by having a dense and coarse
punctation on the elytra and pronotum, and by the dorsal surface of the protibia with ashort stria on the base of teeth. The specimen collected in Osaka Air Port may be an
introduced individual, because it is small in size (Table 15), strongly metallic blue andslightly different in the punctation of the pronotum; the punctational pattern is very
similar to that of specimens from Taiwan and the Philippines.
GENUS MEROHISTER REnTER, 1909
Merohister Reitter, 1909: 282; Ohara, 1992a: 378.

This genus has already been revised in Ohara (1992a, b). Herein I give additional
records to Merohister jekeli and M. aino.

135

1(2)
2(1)
3(4)
4(3)

Key to the Japanese species of the genus Merohister
Head with a few coarse punctures just behind frontal stria anteriorly ...... M. aino (Lewis, 1884)
Head without coarse punctures within frontal stria.
Meso- and metatibiae normal, not dilated. Punctation of pronotum usually present on apical 112of lateral area. Paramera of male genitalia slender. ........................ M. jekeli (Marseul, 1857)
Meso- and metatibiae broadly dilated. Punctation of pronotum present on whole lateral area.
Paramera of male genitalia stout ................................................ M. uenoi M. Ohara, 1992

Merohister jekeli (Marseul, 1857)
Merohister jekeli: Ohara, 1992a: 375.

Japanese name: Emma-mushi.
Specimens examined [additional records]. [I-lokkaido] 6 exs., Toya-Naka-no-shima, Abuta,
29/vii/1993, S. Hori (HOR).

[I-lonshu] Tochigi-ken: 1 ex., Yunishi-gawa, Il1viiill980, M. Inaizumi (TPM); 1 ex.,
Kawamata, Kuriyama-mura, 7/viii/1978, T. Tanaka (TPM); 10 exs., Hikida, Kanuma-shi, 3, 5,
16/vi/1984, M. Takaoka (TPM); 1 ex., Aoyagi, Kuriyama, 16/vi/1979, S. Takahashi (TPM); 1 ex.,
Mashiko, 3/viii/1979, M. Inaizumi (TPM); 1 ex., Kawaraya, Utsunomiya-shi, 20/viii/1981, M.
Miyamoto (TPM); 1 ex., Ditto, 25/vil1981, K. Aida, collected by a dead crow (TPM); 1 ex., Tomita,
Ashikaga-shi, 4/xl 1981, H. Ohkawa (TPM). Kanagawa-ken: 1 ex., Yokono, Hadano City, 2/vi/1989,
Y. Okushima.
[Kyushu] Nagasaki-ken: 1 ex., Kabashima Is., Nishisonoki, 13/vi/1992, Y. Kusui.
Distribution. Japan (Chishima Isles, Hokkaido, HonshU, Shikoku, KyushU,
Sado Is., Izu Isles, Tsushima Isles, Nansei Isles); Sakhalin; Primorskij Kray; Korea;
Taiwan (proper, Lanyu Is.); Continental China; Philippines; India.

Merohister aino (Lewis, 1884)
Merohister aino: Ohara, 1992a: 385.

Japanese name: Ainu-emma-mushi.
Specimens examined [additional records]. [I-lonshu] Tochigi-ken: 1 ex., Yunishi-gawa,
25/vi/1980, T. Tanaka (TPM).

Distribution. Japan (Hokkaido, Honshu).

Merohister uenoi M. Ohara, 1992
Merohister uenoi M. Ohara, 1992b: 495.

Japanese name: Amami-emma-mushi.
Distribution. Japan (Nansei Isles: Amami-Oshima Is.).

GENUS ATHOLUS THOMSON, 1859
This genus has already been revised in Ohara (1992c, 1993a). I gIve some
additional records.

136

Key to the Japanese species of the genus Atholus
1(2)
2(1)
3(4)
4(3)
5(6)
6(5)
7(8)
8(7)

Elytra red maculate ........................................................ A. bimaculatus (Linnaeus, 1758)
Elytra entirely black.
Lateral pronotal stria present on apical 112. Apical end of 3rd elytral dorsal stria strongly bent
inwards .............................................................................. A. coe/estis (Marseul, 1857)
Lateral pronotal stria nearly entire. Third elytral dorsal stria normal.
Fifth elytral dorsal stria present on apical 1/2. ......................... A. pirithous (Marseul, 1873)
Fifth e1ytral dorsal stria nearly complete.
Lateral disk of metasternum with long hairs. Anterior margin of prosternal lobe narrowly
truncate on medium. Punctation of propygidium even ................. A. depistor (Marseul, 1873)
Lateral disk of metasternum without hairs. Anterior margin of prosternallobe round. Punctationof propygidium becoming coarser basally. . ................................................................ .
............................................... A. duadecimstriatus quatuordecimstriatus (GyUenhal, 1808)

Atholus bimaculatus (Linnaeus, 1758)
Atho/us bimaculatus: Ohara, 1992c: 169.

Japanese name: Aka-mon-emma-mushi.

Distribution. Japan (Oki Is., KyOshO, Iki Is., Tsushima Isles); Europe; Holarctic
Region; Argentina (introduced); Chad (introduced); India; Tenasserim.
Atholus coelesis (Marseul, 1857)
Atha/us coe/estis: Ohara, 1992c: 173.

Japanese name: Sujimagari-emma-mushi.
Specimens examined [additional records]. [Nansei Isles] Yonaguni Is.: 1 ex., Mt. Urabu-dake,
19/ii/1990, Y. Okushima.

Distribution. Japan (Nansei Isles); Taiwan; Continental China; India; Sri Lanka;
Indonesia (Java, Celeves).
Atholus depistor (Marseul, 1873)
Athalus depistor: Ohara, 1992c: 176.

Japanese name: Munakubo-emma-mushi.

Distribution. Japan (Hokkaid6, Honshu, Shikoku, KyOshO, Nansei Isles,
Ogasawara Isles); Taiwan; southeastern China; Korea; Siberia; Primorskij Kray.
Atholus duodecimstriatus quatuordecimstriatus (Gyllenhal, 1808)
Atholus duodecimstriatus quatardecimstriatus: Ohara, 1993a: 135.

Japanese name: Ko-tsuya-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Hamayiichi, Wakkanai,
23/viii/1993, S. Hori (HOR); 1 ex., Futatsu-yama, Shibecha, Kushiro, 6/vi/ 1974, K. Ijima (11); 1 ex.,137

Tokachi-gawa-kak6, Otsu, Toyohoro-ch6, 27/v/1990, J. Mochizuki; 2 exs., Sapporo, 9/vii/1985, M.
Ohara.
[Honshu] Tochigi-ken: 4 exs., Ozawa-bokuj6, Imaichi-shi, 25/vii/1981, K. Aida (rPM); 1 ex.,
Kawaraya-machi, Utsunominya-shi, 20/vii/1981, K. Aida (rPM).

Distribution. Japan (Hokkaido, HonshO, Shikoku, KyOshO, Tsushima Isles,
Nansei Isles); North Europe and Central Europe (at high altitudes); Siberia; Mongolia;
Continental China; Taiwan.

A tho Ius pirithous (MarseuI, 1873)
Atholus pirithous: Ohara, 1993a: 141.

Japanese name: Tsuya-maru-emma-mushi.
Specimens examined [additional records]. [Hokkaid6] 4 exs., Futatsu-yama, Shibecha, Kushiro,
15/ix/1955, 12/vii/1975, 28/vii/1991, K. Ijima (11); 3 exs., Sapporo, 7/vi/1985, M. Ohara.
[Honshu] Tochigi-ken: 1 ex., Kawaraya-machi, Utsunomiya-shi, 20/vii/1981, K. Aida (rPM);
1 ex., Kawaraya-machi, Utsunomiya-shi, 3/vii/1981, K. Aida, collected from a dead crow (rPM); 3
exs., Turuta-Numa, Utsunomiya-shi, 13-14/v/1991, no collector's name (rPM); 1 ex., Tomita,
Ashikaga-shi, 4/xl 1981, H. Ohkawa (TPM).
[Kyushu] Got6 Isles.: 1 ex., Uki Is., 15/v/1993, Y. Kusui.

Distribution. Japan (Hokkaido, HonshO, Shikoku, Kyfishfi, Tsushima Isles,
Goto Isles, Nansei Isles); Taiwan; Continental China; Korea; Primorskij Kray; VietNam.
GENUS MARGARINOTUS MARSEUL, 1853
Margarinotus Marseul, 1853: 549; Wenzel, 1944: 123; Ohara, 1989: 6.

This genus has already been revised in Ohara (1989). Herein I add somespecimens newly examined and renew the key to the Japanese species thereafter.
Key to the Japanese species of the genus Margarinotus
1(2)Lateral sides of pronotum strongly depressed. Intercoxal disk of metasternum wholly covered
with coarse punctures .............................................. Subgenus Kurilister Tishechkin, 1992................................................................... ....... M. (K.) kurbatovi (rishechkin, 1992)
2(1)Lateral sides of pronotum not depressed. Intercoxal disk of metasternum not punctate (rarely
punctate, with punctures only along lateral metasternal stria).
2(19) Pronotum with 2 lateral pronotal striae........... Subgenus Ptomister Houlber et Monnot, 1923
3(4) Inner lateral pronotal stria strongly undulate behind the eyes. .... M. (P.) boleti (Lewis, 1884)
4(3)Inner lateral pronotal stria not strongl y undulate behind eyes.
5(16) Prosternal keel without carinal stria (sometimes with an obsolete one in stria/a, weymarni andreichardti, but these species are densely covered with moderate-sized punctures on pygidium).
6(15) Body larger, 5.0 - 9.2 mm in length.
7(10) Lateral stria of metasternum united with oblique stria of metasternum.
8(9)Lateral disk of metasternum wi th long hairs. ............. M. (P.) cadavericala (Bickhardt, 1920)9(8)Lateral disk of metasternum without hairs. ...................... .... M. (P.) agnatus (Lewis, 1884)
10(7) Lateral stria of metasternum not united with oblique stria of metasternum.
11(12) Lateral disk of metasternum without hairs. ..............
M. (P.) reichardti Kryzhanovskij, 1976
12(11) Lateral disk of metasternum with long hairs.
13(14) Outer lateral pronotal stria extending beyond basal end of inner one.................................... .
................................................................................. M. (P.) weymarniWenzel, 1944
14(13) Outer lateral pronotal stria not extending beyond basal end of inner one ................................ .
........................................................................... M. (P.) striota (c. R. Sahlberg, 1819)
138

15(6) Body smaller, 4.0 - 4.5 mm in length. ............ ............. M. (P.) yezoensis M. Ohara, 1989
16(5) Prostemal keel with carinal striae. Pygidium sparsely covered with coarse punctures.
17(18) Inside the inner pronotai stria densely with large punctures................................................ .
.. .... .......... ........ ...... .... ...... ...... .... ...... .... .........
M. (P.) marginepunctatus (Lewis, 1879)
18(17) Inside the inner pronotal stria without large punctures. ............. M. (P.) sutus (Lewis, 1884)
Subgenus Grammostethus Lewis, 1906
19(2) Pronotum with 1 lateral pronotal stria. ..................
................................................................................... M. (G.) niponicus (Lewis, 1884)

Margarinotus (Kurilister) kurbatovi (Tishechkin, 1992)
Kurilister kurbatovi Tishechkin, 1992: 328 [Kuriles, SW Kunashiri Is.].
Margarinotus (Kurilister) kurbatovi: Ohara, 1993b: 3.

Japanese name: Hirata-kaku-hime-emma-mushi.

Distribution. Japan (Chishima Isles, Hokkaid6, Honshu, Shikoku).
Margarinotus (Ptomister) boleti (Lewis, 1884)
Margarinotus (Ptomister) boleti: Ohara, 1989: 9.

Japanese name: Kinoko-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Futatsu-yama, Shibecha, Kushiro,
10/ix/1976, K. Ijima (IJ); 10" 1 ~ ,Kariba-yama, Osima Peninsula, 19-23/vii/1972, no collector's name

(ElHU).
[Honshu] Fukushima-ken: 1 ex., Futamata-zawa, Yumoto, Iwase, 15/viii/1950, Y. Kurosawa(NSMT); 1 ex., Nagarazawa, Asahida, Minami-Aizu, l/viii/1949, Y. Kurosawa (NSMT); 1 ex.,
Akaiwa, Fukushima-shi, 25/vi/1960, K. Tazoe. Tochigi-ken: 1 ex., Chugushi, Nikko, 9-11/viii/2603(=1903), T. Adachi (NSMT); 1 ex., Kotoku, Nikko, 8/vii/1967, J. Aoki (NSMT); 1 ex., Yunishigawa, Kuriyama-mura, Shioya-gun, 15/vi/1980, M. Inaizumi (TPM); 1 ex., Ditto, Il/viii/1980, S.
Sugimoto (TPM); 1 ex., Yokokawa, Fujiwara-machi, Sbioya-gun, 16/vi/1985, T. Miyamoto (TPM); 1
ex., Yunago-sawa, Yunishi-gawa, Kuriyama-mura, Shioya-gun, 8/viiI1991, K. Satoh (TPM); 3 exs.,
Minamikata, Yamizo-yama, Kurobane-machi, 12/viii/1984, M. Takaoka (TPM). Nagano-ken: 1 ex.,Kami-Kochi, Tokusawa, 25/viii/1967, H. Higuchi. Gifu-ken: 1 ex., Oshirakawa, Shirakawa-mura,
26/vii/1987, Y. Takai; 1 ex., Kammuri-yama-toge, Tokuyama-mura, Gifu, 28/vii11985, Y. Takai.
[Kyushu] Kagoshima-ken: 1 ex., Onami-ike, Kirishima, 5/iv/1985, H. Shimozono.

Distribution. Japan (Chishima Isles, Hokkaid6, Honshu, Shikoku, Kyushu);Taiwan.
Margarinotus (Ptomister) cadavericola (Bickhardt, 1920)
Margarinotus (Ptomister) cadavericola: Ohara, 1989: 13.

Japanese name: Osawa-hime-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Harutori, Kusbiro-shi, 23/x/1983,
E. Osaki; 1 ex., Seiryu-Basbi, Bisei-gawa-Joryu, Memuro, 2/ix/1990, K. Haga; 1 ex., Sapporo,vi 1904, no collector's name.; 1 ex., Ditto, 20/v/1955, T. Kumata (ElHU); 1 ex., Hirafu side, alt. 1000
- 1400 m, Mt. Yotei, Kutchan, 30/viii/1993, M. Ohara; 7 exs., TOya-Naka-no-shima, Abuta, 17,29/vii, 28/viii/1993; S. Hori (HOR).
[Honshu] Aomori-ken: 1 ex., Takada, Aomori-shi, 14/vii/1986, H. Saito. Shizuoka-ken: 3 exs.,
Fujinomiya-shi, 2/viii/1978, S. Mochizuki; 1 ex., Koizumi, Fujinomiya-shi, l/xi/1978, S.
Mochizuki.
139

Distribution. Japan (Chishima Isles, Hokkaido, Honshu, Kyushu); Primorskij
Kray; Continental China.
Margarinotus (Ptomister) agnatus (Lewis, 1884)
Margarinotus (Ptomister) agnatus: Ohara, 1989: 15.

Japanese name: Nise-hime-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Futatsu-yama, Shibecha, Kushiro,
26/vi/1988, K. Ijima (IJ); 2d"d" ,4 exs., Sapporo, 20/v/1955, T. Kumata (EIHU); 2 exs., Maruyama,
Sapporo, 8/vi/1954, T. Kumata (EIHU); 1 ex., Moiwa, Sapporo, 4/vi/1905, no collector's name.
(EIHU); 5 exs., Nopporo, 24/vi, 1, 211viil 1988, M. Ohara; 2 exs., Hirafu side, alt. 350 m, Mt. Y6tei,
Kutchan, 20/viii/1993, M. Ohara; 2 exs., Ditto, alt. 350 - 600 m, 30/viii/1993, M. Ohara; 1 ex., Mt.
Hakodate, 26/viiil 1951 , K. Honma (EIHU); 20 exs., Toyo-Naka-no-shi ma, A buta, 26/v, 12/vi, 17, 28,
29/vii, 28/ix/1993, S. Hori (HOR).
[Honshu] Fukushima-ken: 1 ex., Fukiya-yama, Monden, Kita-Aizu, 211v/1948, K. Hayashi(NSMT); 1 ex., Nittahara-Chippu-Kojo, Tateiwa-mura, I/vi/1993, K. Haga. Tochigi-ken: 2 exs.,
Nikko, 14/viil1943, no collector's name (NSMT); 1 ex., Awanono-cho, 14/v/1989, H. Yoshitomi(TPM); 1 ex., Senjugahara, Nikko-shi, 211vi/1985, K. Niimi (TPM); 4 exs., Hikita, Kanuma-shi,
21/v/1984, M. Takaoka (TPM). Tokyo-to: 1 ex., Takao-san, 23/iv/1969, H. Takizawa. Shizuoka-ken:
4 exs., Subaru Line, Ni-gome, Fujinomiya-shi, 2/viii/1978, S. Mochizuki.
[Shikoku] Tokushima-ken: 1 ex., Okunoi, Yamagawa, 16/iv/1986, S. Mano.
[Kyushu] Nagasaki-ken: 1 ex., Kagasahima Is., Nishisonoki, 13/v/1992, Y. Kusui.

Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu); Himalaya; North
India; Korea.
Margarinotus (Ptomister) weymarni Wenzel, 1944
Margarinotus (Ptomister) weymarni: Ohara, 1989: 19.

Japanese name: Hime-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 3 exs., Futatsu-yama, Shibecha, Kushiro,
10/vi/1975, 9/v/1990, K. Ijima (IJ); 1 ex., Oku-Shibetsu, Kamikawa, Teshio, 16/v11947, T. Hasegawa(NSMT); 1 ex., Abashiri, 211iv/1950, T. Kumata (EIHU); 1 ex., Sapporo, 30/v/1987, M. Ohara; 2exs., Ditto, 6/viiil?, 6Iix/?, no collector's name (EIHU); 2 exs., 1 !j?, Sapporo, no date, S. Matsumura(EIHU); 1d", 1 ex., Ditto, no date, Ishida; 1 ex., Ditto, 16/v/1955, S. Takagi (EIHU); 1 ex., Moiwa,
Sapporo, 19/iv/1965, T. Kocha (EIHU); 1 ex., Maruyama, Sapporo, 23/ix/1953, S. Takagi (EJHU); 2
exs., Nopporo, 24/vi, 211vii/1988, M. Ohara; 1 ex., Menashibetsu-gawa, Shizunai, 5/vii/1993, S.
Hori (HOR); 1 ex., Mt. Kariba, Shimamaki, 6-8/viiiI1988, M. Ohara, T. Tanabe and K. Sayama; 1ex., Kita-Hiyama, 28/v/1988, M. Ohara.
[Honshu] Aomori-ken: 1 ex., Aomori, viI1904, no collector's name; 1 ex., Minami-Tsugaru,
19/vi/1952, K. Shimoyama; l!j? , Hachinohe, 10/vi/1934, S. Fukuda (Fukuda Collection). Fukushimaken: 1 ex., Tokusa, Tateiwa, 26/v/1991, K. Haga. Yamagata-ken: 4 exs., Yonezawa, 12/iv/1941,
16/v/1944, Y. Kurosawa (NSMT); 1 ex., Ditto, 3/v/1944, A. Suzuki. Fukushima-ken: 1 ex.,
Yunohana, Tateiwa, Minami-Aizu, 8/viil1948, Y. Kurosawa (NSMT); 1 ex., Aizu-Wakamatsu,
Fukushima, 15/v/1949, no collector's name (NSMT); 5 exs., Wakamatsu, 13/iv/1946, 5/v11948, K.
Nagayama (NSMT). Tochigi-ken: 1 ex., Miyuki, Utsunomiya-shi, lIvi/1977, T. Kumazawa (TPM); 1
ex., Hikita, Kanuma-shi, 10/vi/1984, M. Takaoka (TPM). Tokyo-to: 1 ex., Shimonoge, Tama-gawa,28/iv/1929, no collector's name; 2 exs., Arakawa, 13/iii/1960, S. Fukumoto; 1 ex., Ushigome,
20/iv/1988, T. Wakamori (NSMT). Kanagawa-ken: 2 exs., Hatano, 15/iii/1974, 6/vi/1985, H.
Takizawa; 1 ex., Yokohama, vi/1923, J. F. Illingworth (BSM). Nagano-ken: l!j?, Ulsugushi-ga-hara,140

24lviiI1952, K. Nishikawa (NSMT). Shizuoka-ken: 15 exs., Hamaoka-cho, 2, 151v, 311x, 3, 7,
301xi11982, 28liii, 241 vi 1983, 2liv, ll1xiI1984, S. Mochizuki. Toyama-ken: 1 ex., Toyama,
201vi11955, S. Takagi (ElHU).

Distribution. Japan (Hokkaido, Honshu, Shikoku, Kyushu); Khabarovskij andPrimorskij Kray; North-eastern China (Manchuria).
Margarinotus (Ptomister) reichardti Kryzhanovskij, 1976
Margarinatus (Ptamister) reichardti: Ohara, 1989: 22.

Japanese name: Tohoku-hime-emma-mushi.

Distribution. Japan (Honshu); Taiwan; Continental China; Korea; Primorskij andsouthern Khabarovskij Kray.
Margarinotus (Ptomister) striola (c. R. Sahlberg, 1819)
Margarinatus (Ptamister) striala: Ohara, 1989: 24.

Japanese name: Ezo-hime-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 1 ex., Futatsu-yama, Shibecha, Kushiro,
25/v/1972, 10/vi/1975, 10/v/1982, K. Ijima (IJ); 2d"'d'" 1 Sf, 5 exs., Moiwa, 211iv, Il1vi/1905, nocollector's name (EIHU); 12 exs., Nopporo, 29/v/1987, 7/vii/1988, M. Ohara; 1 ex., Jozankei,
5/vii/1910, S. Matsumura (EIHU); 1 ex., Menashibetu-gawa, Shizunai, 5/vii/1993, S. Hori (BOR); 2
exs., Hirafu side, alt. 350 m, Mt. Yotei, Kutchan, 20/viii/1993, M. Ohara; 1 ex., Ditto, alt. 350 - 600

m, 30/viii/1993, M. Ohara.

Distribution. Japan (Hokkaido); North Europe; Siberia; Korea; North-eastern
China (Manchuria)
Margarinotus (Ptomister) yezoensis M. Ohara, 1989
Margarinatus (Ptomister) yezoensis: Ohara, 1989: 27.

Japanese name: Nopporo-hime-emma-mushi.
Specimens examined [additional records]. [Honshu] Tokyo-to: 1 ex., Ushigome, 4/vi/1933, T.
Wakamori (NSMT). Nagano-ken: 1 ex., Shimashima-dani, 4/vi/1940, T. Nimura (NSMT).

Distribution. Japan (Hokkaido, Honshu).
Margarinotus (Ptomister) marginepunctatus (Lewis, 1884)
Margarinatus (Ptomister) marginepundatus: Ohara, 1989: 31.

Japanese name: Heriten-emma-mushi.
Specimens examined [additional records). [Honshu] Fukushima-ken: 1 ex., Kuromori, Ota,
Kita-Aizu, 28/iv/1951, Y. Kurosawa (NSMT). Yamanashi-ken: 1 ex., Senyo-ga-taki, ShOsen-kyo,
KOfu-shi, ll1iv/1992, K. Haga. Nagano-ken: 1 ex., Kita Alps, Nanakura, 14/vii11967, K. Tazoe.

Distribution. Japan (Hokkaido, Honshu, Shikoku).

141

Margarinotus (Ptomister) sutus (Lewis, 1884)
Margarinotus sutus: Wenzel, 1944: 126.
Margarinotus (Ptomister) sutus: Ohara, 1989: 34.

Japanese name: Hakone-emma-mushi.
Specimens examined [additional records]. [HonshU] Aomori-ken: 1 ex., Kuroishi, Okawara,
17/v/1992,H. Saito. Yamagata-ken: 1 ex., Shirabutakayu, 17/vi/1980, S. Ohmomo. Fukushima-ken:
1 ex., Aizu-Wakamatsu, 2/v/1948, Y. Kurosawa (NSMT); 1 ex., Jabami, Nagaino, Onuma,
25/vi/1941, Y. Kurosawa (NSMT); 1 ex., Senaka-aburi, Higashi-yama, 28/v/1947, Y. Kurosawa(NSMT); 1 ex., Naka-Moniwa, 15/v/1973, K. Tazoe. Tochigi-ken: 1 ex., Miyuki, Utsunorniya-shi,
29/v/1976, T. Kumazawa (TPM); 1 ex., Moro-yama, Kanuma-shi, 12/v/1983, M. Takaoka, collectedfrom a fungus, Armillaria sp. (TPM). Tokyo-to: 2 exs., Arakawa, 13/iii!1960, S. Fukumoto(NSMT). Shiga-ken: 5 exs., lbuki-yama, 1O/vi/1990, H. Takizawa.

Distribution. Japan (Honshu, Shikoku, Kyushu); Korea.
Margarinotus (Grammostethus) niponicus (Lewis, 1895)
Margarinotus (Grammostethus) niponicus: Ohara, 1989: 37.

Japanese name: Ko-emma-mushi.
Specimens examined [additional records]. [Hokkaido] 2 exs., Shirikoma-dake, Naka-Tombetsu,
22/viii/1993, S. Hori (HOR); 3 exs., Raibetsu, Shibecha, Kushiro, 27/vi/1975, 25/vii/1983,
3/vii11985, K. ljima (11); 3 exs., Futatsu-yama, Shibecha, Kushiro, 17/vi/1972, 6/vii/1974,
8/vii/1984, K. Ijima (I1); 2 exs., Okushihoro, Teshio, 14/vi/1951, T. Hasegawa (NSMT); 1 ex.,

Motokamabetsu, Kita-Moshiri, 9/vii/1987, T. N.; 5 exs., Kenebetsu, Nemuro, 2/viii/1973, T. Kumata(EIHU); 1 ex., Onne-shisui-damu, Minami-Kitoushi, Ashoro, 1l/vii/1992, K. Haga; 1 ex.,
Kitanosilwa, Sapporo, 18/vii, lIviii/1993, S. Hori (HOR); 1 ex., Moiwa, Sapporo, ll/vi/1904, nocollector's name (EIHU); 2 exs., Maruyama, Sapporo, 26/vi/1913, S. Matsumura (EIHU); 1 ex., Ditto,
13/vi/1939, Y. Nishijima (EIHU); 1 ex., Ditto, 10/vi/1954, M. Konishi (EIHU); 1 ex., Sapporo, nodate and no collector's name (EIHU); 1 ex., Ditto, no date, S. Matsumura (EIHU); 1 ex., Ditto, 1903,
no collector's name (EIHU); 2 exs., Ditto, 20/iv/1925, 25/v/1942, H. Kono (EIHU); 3d"d" 2~ ~ ,9exs., Ditto, 22/vi/1985, 4!vi/1987, M. Ohara; 19 exs., Hokkaido Univ., Sapporo, 27/vi/1985, M.
Ohara; 1 ex., Mt. Joten, Sapporo, 6/vi/1978, T. Fujisawa (EIHU); 5 exs., Misumai, Sapporo, 9,12/vii/1991, K. Sayama; 2 exs., Jozankei, 27/vi/1954, K. Honma (EIHU); 1 ex., Ditto, 27/vi/1954,
M. Konishi (EIHU); 1 ex., Nopporo, 17/vi/1936, S. Hirai; 1 ex., Ditto, 29/v/1987, M. Ohara; 1 ex.,
Naganuma, 20/v/1991, A. Iwasaki; 3 exs., Shikotsu-ko, 6/vii/1988, M. Ohara; 7 exs., Hobetsu,
23/vi/1985, M. Ohara; 6 exs., Bibi, Chitose, 4-18/vi/1993, S. Hori (HOR); 1 ex., Meguro, Erimo,
23/vi, 71vii11993, S. Hori (HOR); 1 ex., Mt. Yotei, Kutchan, 291vii11989, N. Yasuda; 16 exs., Hirafu
side, alt. 350 m, Mt. Yotei, Kutchan, 20/viiiI1993, M. Ohara; 6 exs., Ditto, alt. 350 - 600 m,
30/viii/1993, M. Ohara; 3 exs., Sorioi-yama, Rusutsu, 17-27/vi/1991, S. Hori (HOR); 9 exs., ToyaNaka-no-shima, Abuta, 28/vi, 17, 29/vii/1993, S. Hori (HOR); 1 ex., Kariba-yama, Oshima Penins.,
19-23/vii/1972, H. U. Ent. Inst. (EIHU).
[Honshu] Aomori-ken: 1 ex., Tsuta-onsen, 14/vii/1954, T. Kumata (ElHU); 1 ex., Ditto,
25/viiI1954, J. Aoki (NSMT); 1 ex., Minami-Tsugaru, 18/x/1953, K. Shimoyama (NA). Yamagataken: 1 ex., Yonezawa, 25/v/1944, Y. Kurosawa (NSMT). Fukushima-ken: 1 ex., Wakamatsu,
30/vi/1949, Y. Kurosawa (NSMT); 1 ex., Senaka-aburi, Higashi-yama, Kita-Aizu, 13/vi/1948, K.
Ikarashi (NSMT); 3 exs., Futamata-zawa, Yumoto, Iwase, 16/vii/1949, Y. Kurosawa (NSMT); 5 exs.,
Oyama, Monden, Kita-Aizu, 211v/1949, Y. Kurosawa (NA); 2 exs., Yunohana, Tateiwa, Minami-Aizu,
5-8/viii/1983, Y. Kurosawa and S. Okajima; 1 ex., Moniwa, Masuzawa, 3/vii/1970, K. Tazoe; 1 ex.,
Nanairi, Hinoemata, 22/vii/1990, N. M. Ibaraki-ken: 8 exs., Ami-machi, 16/vi/1985, K. Akiyama.
Saitama-ken: 1 ex., Mt. Tatekawa, 10/vi/1979, H. Takizawa. Tochigi-ken: 2 exs., Chuzenji,
19/viii/1915, E. Gallois (EIHU); 3 exs., Nikko, 17/vii/1957, Y. Kurosawa (NSMT); 1 ex., Chugushi,
142

Nikko, 9-11/viii/2603 (=1943), T. Adachi (NSMT); 1 ex., Yamizo-yama, Tanakura-machi,
25/vii/1985, K. Tsurumi (TPM); 2 exs., Shimo-Toshimo-machi, Utsunomiya-shi, 24/iv, 5/viii/1992,
K. Satoh (TPM); 1 ex., Shimo-Togami-machi, Utsunomiya-shi, 1O/vi/1990, K. Satoh (TPM); 3 exs.,
Futaba-cho, Utsunomiya-shi, 18, 29/viiiI1972, T. Miyamoto (TPM); 1 ex., Kawaraya, Utsunomiyashi, 25/vi/ 1981, K. Aida, collected from a dead crow (TPM); 1 ex., Ditto, 23/vi/ 1981, K. Aida (TPM);
1 ex., Higashi-mine, Utsunomiya-shi, 13/viii/1979, M. Inaizumi (TPM); 2 exs., Miyuki,
Utsunomiya-shi, 22/v/1976, lIvi/1977, T. Kumazawa (TPM); 3 exs., Tsuruta-Numa, Utsunomiya-shi,lO-ll/vi, 22-23/vii/1991, no collector's name (TPM); 12 exs., Akada, Nishi-Nasuno-machi, 22,
30/v/1990, 8/viii/1991, S. Ohmomo (TPM); 3 exs., Nogi-Jinja, Nishi-Nasuno-machi, 1O/viii/1984,
M. Takada, collected from a fungus, Psathyrella candolleana (TPM); 17 exs., Hikida, Kanuma-shi,
211v, 16/vi/1984, M. Takaoka (TPM); 16 exs., Ditto, /1984, M. Takaoka (TPM); 2 exs., Moro-yama,
Kanuma-shi, 12/v/1984, M. Takaoka, collected from a fungus, Armillaria sp. (TPM); 2 exs., Ditto,
12/v/1985, M. Takaoka, collected from a fungus, yeast plant on a tree, Styrax japonica (TPM); 16
exs., Takadate-yama, Mashiko-cho, 24/vii/1984, M. Takaoka, collected from a fungus, Tylopilus
reofelleus (TPM); 1 ex., Kuriyama-mura, Shioya-gun, 27/v/1989, Omoda and Okawa (TPM); 1 ex.,
Yunishi-gawa, Kuriyama-mura, Shioya-gun, 15/vii/1980, S. Takahashi (TPM); 1 ex., Tashiro-yama,
Kuriyama-mura, Shioya-gun, 25/viii/1990, K. Watanabe (TPM); 1 ex., Yokokawa, Fujiwara-machi,
Shioya-gun, 23/vi/1985, K. Satoh (TPM); 1 ex., Ditto, 16/vi/1985, T. Miyamoto (TPM); 1 ex.,
Kokufu, Tochigi, 15/vii/1985, H. Yoshitomi (TPM); 3 exs., Azusa-machi, 30/vii/1988, H. Yoshitomi(TPM); 1 ex., Ohira-yama, Tochigi, 2/vii/1989, H. Yoshitomi (TPM); 6 exs., Sochi-Shikenjo, NishiNasuno-cho, 22/v/1990, S. Ohrnomo (TPM). Chiba-ken: 1 ex., Sarashina, Chiba-shi, 7/v/1987, J.
Mochizuki. Tokyo-to: 1 ex., Toshima-ku, 19/v/1963, S. Kondo; 2 exs., Aihara, 16/v/1987, H.
Takizawa; I ex., Tokyo, v/1913, F. Muir (BSM); 2 exs., Mt. Takao-san, Il1v/1957, Y. Kurosawa(NSMT); 1 ex., Ditto, lIvi/1924, no collector's name (NSMT); 2 exs., Ditto, 211vi/1963, S. Nomura(NSMT); 4 exs., Tokyo, 13, 27/v, 13/viiI1914, E. Gallois; 1 ex., Kunitachi, 25/iv/1954, S. Nomura(NSMT); 3 exs., Kinuta, Setagaya, 16/vi/1987, 5/v/1980, 19/iv/1987, Y. Kurosawa (NSMT).
Kanagawa-ken: 2 exs., Yokohama, vi/1923, J. F. Illingworth (BSM); 1 ex., Ditto, vi/1941, K. Sato;
30 exs., Akuwa-cho, Yokohama, 7/vi11983, 25/v, 12/vi/1984, K. Wada; 1 ex., Mt. Oyama, 5/v/1989,
H. Takizawa; 3 exs., Fudakake, Tanzawa, ll/v/1980, H. Tanaka; 9 exs., Kurokawa-Eki, Tarna-ku,
Kawasaki, 26/v, 6/vi/1987, J. Mochizuki. Toyama-ken: 1 ex., Kanayama, Izumi, Toyama, 8/v/1983,
K. Sakai. Yamanashi-ken: 1 ex., Mt. Daibosatsu, 12-13/vii/1969, H. Takizawa; 2 exs., Akeno,
25/viii/1985, N. Kobayashi (NSMT). Nagano-ken: 1 ex., Sugadaira, 24/vii/1978, T. Fujisawa (ElHU);
1 ex., Yatsu-ga-take, 4/x/1941, M. Hanano (NSMT); I ex., Mt. Nyugasa, 3/viiiI1960, S. Nomura; 1ex., Shirahone Spa., 17/vii/1988, K. Tazoe. Shizuoka-ken: 1 ex., Hamaishi, 24/iv/1977, H. Takizawa;
2 exs., Amagi-san, lzu, 17/vii/1955, S. Tsuyuki (NSMT). Nara-ken: 5 exs., Hasedera, Sakurai-shi,
3/v/1985, T. Ito; 1 ex., Kojin, Nosegawa, Yoshino, Nara, 13/vii/1985, T. Ito; 3 exs., Nara-koen, Nara,
3/v/1987, K. Haga, collected from dead deer. Wakayama-ken: 3 exs., Ryumon-yama, Kokawa,
211iv/1985, T. Ito. Hyogo-ken: 1 ex., Saijo, Himeji, 15/v/ 1991, Y. Kusui.
[Shikoku] Ehime-ken: 2 exs., Mt. Ishizuchi, 8/viii11987, K. Matsuda.
[Kyushu] Fukuoka-ken: 2 exs., Tachibana-yama, 17/vii/1988, S. Nomura. Saga-ken: 2 exs.,
Sashi, Karatsu, 16/vii/1950, T. Shibuya (NSMT). Kumamoto-ken: 3 exs., Ichifusa-yama, 12/viii/1988, S. Nomura; 107 exs., Kawaguchi, Sakamoto-mura, 15/vi/1985, T. Tanabe. Kagoshimaken: I ex., Chi ran, 13/viiiI1963, Y. Kurosawa (NSMT); 1 ex., Odomari, Sata, Osumi-hanto,
2/v/1992, K. Haga. Tsushima Is.: 1 ex., Nita, Meboro, 9/vi/1968, Y. Kurosawa (NSMT).

Distribution. Japan (Hokkaido, HonshU, Shikoku, KyushU, Tsushima Isles,
Nansei Isles); Taiwan; Continental China; Korea; Khabarovskij and Primorskij Kray.
6. 3. 6. SUBFAMILY HETAERIINAE MARSEUL, 1857
GENUS HETAERlUS ERICHSON, 1834
Hetaerius Erichson, 1834: 156 [type species: Hister quadratus Kugelann, 1794: 519, designated bymonotypy].
Haeterius (sic): Dejean, 1837: 143.

143

Key to the Japanese species of the genus Hetaerius
1(2)
2(1)

Body red or reddish brown. Pronotallateral area without rugae .............. H. gratus Lewis, 1884
Body black. Pronotallateral area densely with rugae ......................... H. optatus Lewis, 1884

Hetaerius gratus Lewis, 1884
(Fig. 80A, B)
Hetaerius gratus Lewis, 1884: 137 [midway between Shimonosuwa and Wada-toge]; Hasegawa andKubota, 1944: 84 [Yokohama]; Tanoguchi and Koshio, 1979: 8; Kurosawa, 1976: 11;
Hisamatsu, 1985b: 230.
Japanese name: Aka-arizuka-emma-mushi.

Description. Body oval, red brown, shining and furnished with yellow, long
hairs on dorsal surface. Body length, PEL, 1.53 - 1.76 mm (1.67 ± 0.04, n=5), PPL,
1.69 - 2.08 mm (1.91 ± 0.06, n=5). Width, 1.41 - 1.53 mm (1.48 ± 0.02, n=5).
Biometric data are given in Table 16.
Frontal disk flat, feebly concave apically, evenly covered with punctures which
are separated by about 3 (2 - 4) times their diameter. In femlae, the disk covered with
setiferous punctures. Lateral margin strongly carinate from occiput to front margin ofclypeus, the carinae feebly convergent forward before antennal sockets, thence strongly
convergent apically, on clypeus nearly parallel and feebly arcuate inwardly, andseparated on front. Labrum rectangular, its anterior margin cuspidate at middle.
Mandible short and robust.
Pronotal sides (Fig. 80A, B) medially arcuate inwardly, convergent forward,
furnished with long robust hairs, and feebly marginate and carinate laterally, the carinae
running inwardly on basal 113; anterior angles rather broadly and obliquely truncate;anterior margin strongly emarginate, its marginal stria impressed at a little distance fromthe margin on each lateral 113. Pronotallateral stria strongly elevated, complete laterally,reaching nearly to anterior margin and not crenate, behind anterior margin feebly
impressed on lateral 113. Disk separated from lateral 115 by the pronotal lateral stria;
lateral area convex, evenly covered with fine, punctures, which are separated by 4 times
their diameter; a dark brown bulla present on basal 114 of the lateral area; median area
convex medially, feebly depressed along the lateral stria, and sparsely covered with finepunctures which are separated by about 3 times their diameter. In female, the punctures
setiferous, the setae rather long (Fig. 80B).
Epipleura feebly concave medially. Epipleural marginal stria impressed on
median 113. Elytral marginal stria (Fig. 80A, B) complete and feebly carinate. External
subhumeral stria strongly carinate, sinuate and complete, the apical end extending
across elytral apex to medio-apical angles of elytra. Internal subhumeral stria carinate
and abbreviated on apical 114. First - 2nd dorsal striae carinate and complete, apical endof the 1st continuing to the extended external subhumeral stria. Third dorsal elytral stria
carinate and shortened on apical 1/4. Fourth, 5th and sutural striae absent. Surface of
eiytral disk evenly covered with fine punctures which are separated by about 3 times
their diameter, except inside the dorsal striae narrowly. In female, the punctures longsetiferous (Fig. 80B).
Propygidium broad, flat, sparsely and shallowly punctate on basal 113, thepunctures united with each other to form rugae, and sparsely furnished with long hairs.
144

Pygidium sparsely, shallowly and finely punctate, the punctures being separated byabout 3 times their diameter.

Fig. 80. Hetaerius spp. A: H. gratus Lewis, male. B: Ditto, female. C: H. optatus Lewis, male. D:
Ditto, sex undetennined.
145

Anterior margin of pronotallobe straight, its marginal stria complete; disk short,elevated, with a longitudinal impression. Suture between the lobe and the keel clearly
impressed, but narrowly interrupted medially, each section of the suture united withapical end of the lateral prosternal stria. Prosternal keel convex medially; carinal striae
convergent apically, joined in front and extending anteriorly as a carina; disk flat andsparsely and finely punctate; posterior margin broadly and roundly emarginate. Lateral
marginal striae strongly carinate and convergent apically.
Anterior margin of mesosternum produced in front to fit prosternal emargination.
Marginal stria absent. Disk elevated medially on basal 1/3 along the anterior margin,
and with deep fovea laterally; surface microscopically and sparsely punctate. Intercoxal
disk of metasternum convex medially, strongly depressed behind anterolateral angles,
and its punctation similar to that of the mesosternum. Two lateral metasternal striae
present on each side; the inner outwardly arcuate, carinate, the apical end reaching near
the inner angle of the metacoxa; the outer sinuate, lightly carinate, and extending
obliquely outwardly, the apical end reaching near the middle of basal margin of themetacoxa. Post-mesocoxal stria lightly carinate, extending posteriorly on basal 112, thenstrongly bent outwardly and reaching middle of the lateral disk. Lateral disk densely
covered with transverse oblong and coarse punctures, which are often fused with eachother.
Intercoxal disk of 1st abdominal sternum has punctation similar to that of themetasternum; 2 lateral striae present on each side, these striae sinuate, carinate andcomplete. Lateral disk coarsely punctate.
Protibia dilated, oblong and with 16 setae on outer margin. Mesotibia with 16
setae on outer margin. Metatibia with 8 setae on outer margin.
Specimens examined. [Hokkaido] 2ei' ei' 1!f , Okusawa, Otaru, 1, 2/viiil 1993, M. Ohara.
[Honshu] Kanagawa-ken: 1 ex., Kamiyama, Hakone, 9/v/1983, Y. Hirano (Hirano collection); 2
exs., Nishizawa, Tanzawa, 3/v/1983, Y. Hirano (Hirano collection); 1 ex., Shiozawa, Tanzawa,
9/vl 1983, Y. Hirano (Hirano collection). Nagano-ken: 1 ex., Kiso-Ontake-san, Hinagura-Hongo-mura,
19/vi/1952, S. Ueno (NA); 1 ex., Nimeki-daira, 22/v/1983, Y. Hirano (Hirano collection).

Distribution. Japan (Hokkaido, Honshu; KyQshu). New to Hokkaido.
Remarks. Hetaerius gratus resembles H. optatus; however, it is easily
distinguished by the color and the smooth lateral pronotal sides.
Biology of this species has been noted and recorded by some workers, whorecorded the following ants as hosts: Formica exsecta fukaii Wheeler (Tanoguchi &Koshio, 1979), F. japonica Motschulsky (Hasegawa & Kubota, 1944; Kurosawa,1976), F. sanguinea Latreille and F. sp. (Hirano, 1984).
Hetaerius optatus Lewis, 1884

(Fig. 80C, D)
Hetaerius optatus Lewis, 1884: 137 .

Japanese name: Kuro-arizuka-emma-mushi.

Description. Body oval, black, shining and sparsely furnished with long hairs on
dorsal surface; legs, mouthparts, antennae, margins of pronotum and of elytra darkbrown. Body length, PEL, 2.01- 2.14 mm (2.09 ± 0.03, n=3), PPL, 2.18 - 2.27 mm(2.22 ± 0.02, n=3). Width, 1.57 - 1.64 mm (1.81 ± 0.02, n=3). Biometric data are
given in Table 16.
146

Frontal disk flat, densely and coarsely punctate, and sparsely furnished with long
hairs; lateral margin strongly carinate from occiput to posterior angles of clypeus, thecarinae nearly parallel before antennal socket, thence strongly convergent apically,
absent on clypeus, their apical ends separated anteriorly. Labrum rectangular, its
anterior margin broadly emarginate. Mandible short and robust.
Pronotal sides (Fig. 80C) inwardly arcuate medially, convergent forward,
furnished with long and robust hairs and completely and lightly carinate, on basal 113
the carinae running inwardly; anterior angles rather broadly and obliquely truncate;anterior margin strongly emarginate. Pronotal lateral stria not clear, represented byseveral sinuated carinae on lateral 115; these carinae set close, variously fused andinterruputed. A lightly impressed stria behind lateral 1/4 of the anterior emarginatedportion. Disk sparsely furnished with long hairs, separated from lateral 115 by thelateral carinae; the lateral area convex, with various carinae, sparsely and coarsely
punctate, and with a bulla on basal 1/3; the median area convex medially, depressed
laterally, densely covered with large, ocelloid and setiferous punctures which areseparated by about 2 (1 - 3) times the diameter and become denser anteriorly, the setae
long and castaneous, and intermingled with coarse punctures.
Epipleura flat, not concave. Epipleural marginal stria slightly carinate on apical
1/2. Elytral marginal stria sinuate, slightly carinate and complete, the apical end
extending across elytral apex to medio-apical angle of elytron and united with the apicalend of the sutural stria. External subhumeral stria (Fig. 80C) sinuate basally, carinate
slightly and complete. Internal subhumeral stria shortened on apical 114. First - 3rd
dorsal elytral striae slightly carinate and complete. Fourth and 5th dorsal elytral striae
absent. Sixth dorsal elytral stria shortened on basal 114 and not attaining to the posterior
margin. Sutural stria complete, close to the sutural margin. Surface of elytral disk
covered with large, ocelloid and setiferous punctures which are separated by about 2.5(1 - 5) times the diameter, the setae long and yellow, and intermingled with coarse andround punctures which are separated by about 2.5 (1 - 5) times the diameter.
Propygidium broad, convex, densely covered with variously sized, setiferous,
round and shallow punctures. Pygidium sparsely covered with round, coarse andsetiferous punctures which are separated by about 2.5 (1 - 4) times the diameter.
Anterior margin of pro sternal lobe truncate, its marginal stria lightly carinate andinterrupted medially; disk densely and coarseiy punctate. Suture between the keel andthe lobe clearly impressed. Prosternal keel flat; carinal striae present on basal 112, nearlyparallel, strongly carinate and feebly sinuate; disk densely covered with coarse
punctures; posterior margin broadly and roundly emarginate. Lateral marginal striae
carinate and convergent basally.
Anterior margin of mesosternum feebly produced anteriorly to fit prosternal
emargination. Marginal stria strongly carinate, a little distant from the margin; disk
short, sparsely covered with shortly setiferous punctures and with a deep fovea
laterally. Intercoxal disk of metasternum convex medially, strongly depressed onanterolateral angles, and evenly covered with shortly setiferous punctures. Two lateralmetasternal striae present, the inner strongly carinate, arcuate inwards and shortened on
apical 1/5, the outer carinate, extending posteriorly outwardly and reaching near the
middle of metasternal-metepistenal suture. Post-mesocoxal stria slightly carinate,
extending posteriorly and outwardly and reaching near the middle of the lateral disk.
Lateral disk densely covered with transverse, oblong, large and shallow punctures
147

Table 16. Biometric data of Hetaerius gratus Lewis and H. optatus Lewis

APW

PPW
PL
EL
EW
ProW
ProL
PyL
PTL

MSTL
MTTL

H. gratus

H. optatus

0.49-0.55 (0.52±0.01) 5
1.22-1.33 (1.27±0.02) 5
0.55-0.63 (0.60±0.01) 5
0.86-1.13 (0.99±0.04) 5
1.14-1.53 (1.48±0.02) 5
0.71-0.82 (0.75±0.02) 5
0.42-0.47 (0.45±0.01) 5
0.43-0.67 (0.50±0.04) 5
0.51-0.59 (0.54±0.02) 5
0.51-0.71 (0.63±0.03) 5
0.67-0.82 (0.72±0.02) 5

0.57-0.59
1.31-1.35
0.70-0.76
1.22-1.24
1.57-1.64
0.87-0.92
0.52-0.57
0.55-0.63
0.65-0.70
0.87
0.92-0.96(0.57±0.01) 3(1.33±0.01) 3(0.73±0.02) 3(1.23±0.01) 3(1.61±0.02) 3(0.90±0.01) 3(0.54±0.01) 3(0.58±0.02) 3(0.67±0.01) 3
3(0.94±0.01) 3

which are often fused with each other.
Intercoxal disk of 1st abdominal sternum has punctuation similar to that of themesosternum; 2 lateral striae present on each side, strongly sinuate basally andcomplete. Lateral disk evenly covered with oblong, large and shallow punctures.
Protibia dilated, oblong and with 8 robust setae on outer margin. Mesotibia andmetatibia furnished with long hairs on outer margin, their outer margin strongly arcuate
outwardly.
Specimens examined. [Hokkaido] 1 ex., Nopporo, 19/vi/1987, M. Ohara; 1if, Maruyama,
Sapporo, 23/v/1951, T. Kumata (EIHU); 3 exs., Toya-Naka-no-shima, Abuta, 12-29/vi, 28/vi17/viil1993, S. Hori (HOR).
[Honshu] Akita-ken: 1 ex., Chokai-san, 30/v/1939, Suzuki (NA). Kanagawa-ken: 1 ex.
Manazuru, 22/v/1955, M. Kubota, collected from the nest of Apherwgaster famelica. Nara-ken: 1 ~,Muroo, l/viii/1989, M. Mori.
Distribution. Japan (Hokkaido, Honshu, Shikoku).
Remarks. Hetaerius optatus resembles H. gratus; but they can easily be
distinguished by the color and the dense rugae on the lateral pronotal sides.
The specimen (Fig. 800) from Manazuru, Kanagawa-ken differs from the otherexamined specimens in the dense short hairs and the absence of yellow and long hairson the dorsal surface. The possibility can not be eliminated that it should be referred toan undescribed species. Further material is necessary for a precise comparison.
Little is known about the habitat of this species. Nishikawa (1987) noted that H.
optatus was found in the nest of an ant, Lasiusfuliginosus (Latreille).
6.3. 7. SUBFAMILY DENDROPHILINAE REITTER, 1909

1(2)
2(1)
3(6)
4(5)
5(4)

Key to the Japanese tribes of the subfamily Dendrophilinae
Outer margin of protibia thick, with 2 edges bearing short setae, without dentic1es and spinula.
Groove between the edges inset with protarsus ........................... Tribe Anapleini Olexa, 1982
Outer margin of protibia thin, single-edged, the edge usually with small dentic1es, and with asmall or strong hooked spinula. Protarsus put on dorsal surface of protibia.
Epistoma narrow, its lateral margins weakly convergent apically. Front without stria. Body
round or oval, and usually convex. Basal piece of male aedeagus usually short.
Elytral disk with normal dorsal striae, which are well developed and parallel. Spinula of protibia
large............................................................................ Tribe Dendrophilini Reitter, 1909
Elytral disk punctate, without dorsal stria except for vague rudiments, these usually present
148

6(3)

basally and oblique. Spinula of protibia small. .............................................................. ..
............................................................ Tribe Bacaniini Kryzhanovskij and Reichardt, 1976
Epistoma broad, trapezoid; frontal stria well developed and completely impressed behind labrum(sometimes interrupted anteriorly, ex. Eulomalus). Body oblong-oval, sometimes oval, andmoderately convex. Basal piece of male aedeagus long, usually 3 times as long as parameres .
.......................................................................................Tribe Paromalini Reitter, 1909

TRIBE DENDROPHIUNI RElITER, 1909
GENUS DENDROPHILUS LEACH, 1817
Dendrophilus Leach, 1817: 77 [type species: Hister punctatus Herbst, 1792: 41, designated bymonotypy].

SUBGENUS DENDROPHILUS LEACH, 1817

Dendrophilus (Dendrophilus) xavieri Marseul, 1873
(Fig. 81, 82,83)
Dendrophilus xavieri Marseul, 1873: 221, 226 [Japan]; Jakobson, 1911: 647; Hinton, 1945: 317, 333[genitalia]; Hisamatsu, 1985b: 224, pI. 40, no. 23.
Dendrophilus (Dendrophilus) xavieri: Kryzhanovskij and Reichardt, 1976: 256; Mazur, 1984: 115.
Dendrophilus califomicus Ross, 1937: 68, synonymized by Hatch, 1962: 268.
Dendrophilus sexstriatus Hatch, 1938: 18, synonymized by Hinton, 1945: 334 .

. ""'/-aalCoo~~'x.....

~·i~tif.1

p~~~;o,/{:;go~4 ~':':':'...

·oooo~·o"oo~:o: ':." . .

F.

.::~ ... ~.O."o :.: ',':, :: .:.
l~--'---,...(',Pi '\

B~

Fig. 81. Dendrophilus xavieri Marseul. A: Pronotum and left elytron. B: Prosternum and meso- andmetasterna. C: Head. D: Protibia, dorsal view. E: Ditto, ventral view. F: Mesotibia, dorsal
view. G: Metatibia, dorsal view.
149Dendrophilus punctatus Ross, 1940: 107, synonymized by Hatch, 1962: 268.
Dendrophilus punctatus var. sexstriatus: Hatch and McGrath, 1941: 55.

Japanese name: O-mame-emma-mushi.

Description. Male and female. Body length, PPL, 2.78 - 3.30 mm (3.01 ± 0.03,n=20), PEL, 2.62 - 2.98 mm (2.78 ± 0.03, n=20). Width, 1.93 - 2.26 mm (2.09 ±0.02, n=20). Biometric data are given in Table 17. Body oblong, oval. Cuticle blackand shining; tarsi and antennae dark rufopiceous.
Head (Fig. 8IC) even, densely and coarsely punctate, the punctures being
separated by about 1.5 (1 - 2) times the diameter; other punctures fine, intermingledwith the coarse ones, sparser medially; frontal stria absent. Mandible short and stout;
labrum with setiferous punctures.
Pronotal sides (Fig. 8IA) strongly convergent forward; disk sparsely covered
with coarse punctures which are separated by 2 - 5 times their diameter and become
denser laterally, with fine punctures intermingled. Marginal pronotal stria complete andfeebly crenulate.
Marginal epipleural stria complete, strongly carinate, and feebly sinuate on apical112. Marginal elytral stria deeply impressed and complete. Epipleura densely covered
with coarse punctures which become sparser and finer on the inner 112. Disk of elytra
densely covered with coarse punctures on apical 112, which are separated by about theirown diameter and often united with each other to form rugae on the apical 113; on basal
112, interstices among dorsal striae sparsely covered with moderate or fine punctures,
which are irregularly separated by about 2.5 (1 - 5) times the diameter; extreme apicalband impunctate. Subhumeral stria absent, but sometimes present on median 113.
Oblique humeral stria indistinctly present on basal 113. First dorsal stria nearly complete
and deeply impressed, but its apical 113 obsolete. Second and 5th dorsal striae present
on basal 2/3, sometimes nearly complete. Third and 4th dorsal striae present on basal
112. These dorsal striae strongly impressed and crenate basally and progressively
obsolete apically. Sutural stria present on basal 2/3 but its basal 113 indistinctly
impressed, and its median 113 crenated.
Propygidium short, with 2 or 3 transverse rows consisting of coarse punctures
along posterior margin. Pygidium densely covered with coarse and ocelloid punctures
which are separated by 0.5 - 1 times the diameter.
Anterior margin of prosternal lobe (Fig. 81B) round and shortly marginate on
median portion; sides marginate completely; disk of lobe densely and coarsely punctate.
Prosternal process somewhat sparsely punctate. Carinal striae complete, strongly
carinate and not united with each other at their posterior terminal ends. Descending
lateral stria complete and carinate.
Anterior margin of mesosternum deeply emarginate in median portion.
Mesosternum short, sparsely covered with coarse punctures, which are separated byabout their own diameter, and strongly marginate laterally. Meso-metasternal sutureindistinctly impressed and complete, and distantly with a strongly crenated stria on
mesosternum. Intercoxal disk of metasternum even, and sparsely and finely punctate,
the punctures coarse along lateral metasternal stria and along posterior margin. Lateraldisk of metasternum densely covered with large and shallow punctures. Mesocoxal stria
complete along mesocoxa. Lateral metasternal stria carinate and extending posteriorly
and obliquely, its apical end reaching about 1/2 to apical 113 of metasternal150

metepisternal suture.
Intercoxal disk of 1st abdominal sternum completely striate on each side, andsparsely and coarsely punctate laterally.
Protibia (Fig. 81E, D) dilated, with 12 denticles on outer margin.
Male genitalia as shown in Fig. 82.
Specimens examined. [Hokkaido] 1 ex., Teshio, 18/vii/1969, no collector's name; 11 exs.,
Otaru, iv/1985, Y. Kusui; 1 ex., Tokiwa, Sapporo, 10/vii/1985, M. Ohara; 2 exs., Bibi, 28/vi/1987,
M. Ohara.
[Honshu] Miyagi-ken: 1 ex., Oshiro-yama, Sendai, 27/iv/1953,K. Nagayama. Yamagata-ken: 1ex., Yubama-Onsen, 26/vii/1951, K. Nagayama (NSMT). Fukushima-ken: 1 ex., Moriai, Fukushimashi, 1/vii/1960, K. Tazoe. Tokyo-to: 3 exs., Setagaya, 18/ix/1949, Ohira (NA). Kanagawa-ken: 1 ex.,
Hatano, 7/viii/1972,H. Takizawa; 1 ex., Yokohama, 1/vi/1941,K. Sato (NSMT). Nagano-ken: 1 ex.,
Kiso-Koma-Kogen, 13/vi/1993, K. Tazoe (Tazoe coli.). Wakayama-ken: 1 ex., Igumbashi, Kinokawa,
Table 17. Biometric data of Dendrophilus xavieri Marseu\.

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

0.75-0.92 (0.83± 0.01) 20
1.54-1.80 (1.66± 0.02) 20
0.75-0.92 (0.83± 0.01) 20
1.80-2.09 (1.95± 0.02) 20
1.93-2.26 (2.09± 0.02) 20
1.05-1.24 (1.14± 0.01) 20
0.16-0.23 (0.20±0.005) 20
0.72-0.92 (0.83± 0.01) 20
0.62-0.75 (0.70± 0.01) 20
0.65-0.85 (0.77± 0.01) 20
0.82-0.98 (0.90± 0.01) 20

oFig. 82. Dendrophilus xavieri Marseu\. A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Apex of 8th sternum,
ventral view. F: Male genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. G:
Ditto,lateral view.
151

Dendrophilus xavieriFig. 83. Collection sites of Dendrophilus xavieri Marseul in Japan.
7/xi/1953, K. K. (NA). Hyogo-ken: 1 ex., Goten-yama, Takarazuka, 25/ix/1948, M. Goto (NA).
Okayama-ken: 1 ex., Saeki, 22/vi/1949, K. Kurosa (NA).
[Shikoku] Kagawa-ken: 2 exs., Toyonaka-cho, Samp6-gun, 4/v/1980, S. Ohmomo (NSMT).
[Kyushu] Fukuoka-ken: 1 ex., Mt. Adachi, 5/iii/1963, S. Nakao (NA). Nagasaki-ken: 1 ex.,
Shimabara, 18-19/iv/1952, K. Oyama (NA); 22 exs., Kono-ura, Hirado-shi, 16/ix/1979, A. Oda.

Miyazaki-ken: 2 exs., Miyazaki, 13/x!'!, no collector's name. Kagoshima-ken: 1 ex., Kurino,
Kirishima, 23/vil 1985, H. Daid6; 1 ex., Takachiho-gawara, Kirishima, ll1iiil 1980, T. Kinoda.
Distribution (Fig. 83). Japan (Hokkaid6, Honshu, Shikoku, KyQshQ, Izu Isles);Taiwan; East Siberia; introduced to England and North America (British Colombia,
Washington, Oregon).
Remarks. Dendrophilus xavieri superficially resembles Gnathoncus spp.;
however, it can easily be distinguished by the presence of the prostemal lobe. This
species is well characterized among the Japanese histerids.
Dendrophilus xavieri occurs in many kinds of rotting organic matter, such ascompost piles, hen and pig manure, carrion, etc., and also found in nests of ants andbirds. Hinton (1945) recorded the species on pumice in warehouse and in rotting grainin the basement of a flour-mill. Hisamatsu (l985b) recorded it from a nest of rats.
TRIBE ANAPLEINI OLEXA, 1982
GENUS ANAPLEUS HORN, 1873
Anapleus Hom, 1873: 311 [type species: Bacanius marginatus J. L. LeConte, 1853: 292, designatedby monotypy]; Bickhardt, 1913: 175; 1916-17: 63,73; Reichardt, 1941: 84, 126; Arnett, 1962:
375,380; Kryzhanovskij and Reichardt, 1976: 260; Mazur, 1984: 116.
Sphaeroderma J. L. LeConte, 1861: 76, nec Stephens, 1831: 328, synonymized by Mazur, 1984: 116[type species: Bacanius marginatus J. L. LeConte, 1853: 292].
Ahraeodes Reitter, 1886: 272, synonymized by Bickhardt, 1913: 175 [type species: Abraeus raddei
Reitter, 1877: 292]; Jakobson, 1911: 642,653.
Tadzhister Kryzhanovskij, 1966: 55, synonymized by Kryzhanovskij and Reichardt, 1976: 260 [typespecies: Tadzhister graci/ipes Kryzhanovskij, 1966: 55].
Diagnosis. Body small, about 2.5 mm long; oval, convex, usually dark brown or
black; tibiae, tarsi, antennae and mouthparts rufopiceous. Frontal stria of head absent;152surface obliquely convex in front of eyes and feebly depressed on longitudinal medianline. Labrum transverse, with a pair (or more) of setiferous punctures. Mandibles shortand stout. Antennae long; scape oblong and stout; pedicel somewhat long and stout;
club consisting of apical 3 flagella, of which the sutures are distinct. Pronotum
transverse, its sides usually strongly convergent apically; lateral and anterior margin
narrowly marginate; apical angles acute and strongly depressed. Elytra usually coarsely
punctate, sometimes strigate apically; dorsal stria absent; apex of elytra truncate.
Propygidium transverse and nearly vertical. Pygidium curved downwards.
Prosternallobe quadrate and broad; suture between lobe and process indistinct; antennal
cavity longitudinal, deep and situated along prosternal lobe and process; carinal striae
deeply impressed. Mesosternum short and transverse. Meso-metasternal suture distinct
and crenate. Metasternum coarsely and densely punctate. Protibia narrow, dilated
medially, without large denticles, usually dentate-serrate with very small denticles.
Meso- and metatibiae narrow and long.
Key to the Japanese species of the genus Anapleus
1(2)
2(1)
3(4)
4(3)
5(6)
6(5)

Pygidium with a depressed area near tip ................................... A. hagai M. Ohara, sp. nov.
Pygidium without depressed area
Punctation of elytra dense; on humeral area of elytra, the punctures being separated by 0.3 times
their diameter ........................................................................... A. semen (Lewis, 1884)
Punctation of elytra sparse; on humeral area of elytra, the punctures being separated by 1 - 2times the diameter.
Punctation of metasternum with uniform punctures ................ A. nakanei M. Ohara, sp. nov.
Punctation of metasternum with large and fine punctures ........ A. nomurai M. Ohara, sp. nov.

Anapleus semen (Lewis, 1884)(Fig. 84,85,86,87)
Triballus semen Lewis, 1884: 137 [Japan: Honshu].
Anapleus semen: Nakane, 1963: 68; Mazur, 1984: 117.
Anapleus japonicus Nakane, 1961: 4 [Rausu, Hokkaido]; Kryzhanovskij and Reichardt, 1976: 262;
Olexa, 1982: 44.

Japanese name: Tsubu-emma-mushi.

Description. Body oval, moderately convex, black and shining; tarsi andantennae rufopiceous. Body length, PPL, 2.05 - 2.23 mm (2.14 ± 0.03, n=5), PEL,
1.95 - 2.11 mm (2.03 ± 0.03, n=5). Width, 1.46 - 1.85 mm (1.67 ± 0.05, n=6).
Biometric data are given in Table 18.
Head (Fig. 85C) densely, coarsely and deeply punctate. Epistoma convex. Frontdepressed between eyes, without stria.
Pronotal sides (Fig. 85A) strongly convergent apically. Disk of pronotum
densely covered with punctures, which are as large as those of head and are separated
by 0.75 (0.5 - 1) times the diameter, and with a shallow depression behind each eye.
Marginal stria strongly carinate and complete, the anterior portion densely crenate.
Epipleural marginal stria complete and densely crenate. Elytral marginal stria
somewhat carinate, complete and sparsely crenate, its apical end extending along theapex of elytra and attaining to near the suture. There is a narrow impunctate bandoutside the epipleural marginal stria. Area between epipleural and elytral marginal striae
153with 2 or 3 rows of coarse punctures. Lateral sides of elytra elevated. Surface of elytra(Fig. 86A, B) densely covered with coarse punctures, which are a little larger thanthose of the pronotum and are separated by 0.3 - 0.5 times their diameter; finepunctures narrowly along suture; basal 112 of elytra densely with longitudinal rugae;
extreme apex impunctate. Dorsal stria absent.
Propygidium densely covered with somewhat deep punctures, which are a little
finer than the elytral ones and separated by about 112 their diameter. Pygidium (Fig.
860) sparsely punctate, the punctures being separated by their own diameter and aslarge as those of propygidium.
Anterior margin of prosternal lobe (Fig. 86C) straight medially and slightly
marginate and carinate; sides strongly carinate and marginate; posterior angles deeply
excavate, the excavations are situated just before ends of carinal striae of prosternal
process. Disks of the lobe and process densely covered with coarse, deep puncturesand sometimes intermingled with ocelloid ones, these coarse punctures being separated
by about 112 their diameter and becoming sparser on a narrow basal band. Carinal striae
completely impressed.
Anterior margin of mesosternum (Fig. 86C) nearly straight; disk of mesosternum
short (width: length = 4 : 1), and coarsely and sparsely covered with ocelloid andshallow punctures, which become finer medially; lateral side completely marginate.
Meso-metasternal suture complete and strongly carinate. Mesosternum feebly convex
on median area, and covered with deep, round and large punctures, which are separated
by about 1/2 their diameter, the punctures becoming sparser on median area. Lateralmetastemal stria extending obliquely and posteriorly, reaching near middle ofmetasternal-metepisternal suture. Intercoxal disk of 1st abdominal sternum densely
covered with punctures, which are a little finer than those of the metasternum,

Fig. 84. A: Anap/eus semen (Lewis), Holotype. B: Anap/eusjaponicus Nakane, Holotype (NA).
154

Fig. 85. Anapleus semen (Lewis), adult. A: Dorsal view. B: Ventral view. C: Head, frontal view. D:
Meso- and metasterna. E: Lateral side of metastenum. F: Punctation of middle area of intercoxal
disk of metasternum.

155

Table 18. Biometric data for Anapleus semen Lewis

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Holotype

Holotype of
A. japonicus

0.43
1.14
0.51
1.26
1.46
0.67
0.20

0.63
1.40
0.59
1.48
1.69
0.79
0.22
0.45
0.61
0.69
0.75

male and female

0.43-0.63 (0.58±0.03) 6
1.14-1.42 (1.33±0.04) 6
0.51-0.67 (0.59±0.02) 6
1.26-1.48 (1.40±0.03) 6
1.46-1.85 (1.67±0.05) 6
0.67-0.79 (0.75±0.02) 6
0.20-0.28 (0.23±0.01) 6
0.39-0.45 (0.42±0.01) 5
0.53-0.67 (0.61±0.02) 5
0.59-0.69 (0.63±0.02) 5
0.65-0.75 (0.70±0.02) 5

Fig. 86. Anapleus semen (Lewis), adult. A: Elytra around scutellum, dorsal view. B: Punctation of
medio-basal area of elytron. C: Prostemum, ventral view. D: Pygidium.

156

~

B

0.25 mmo

I

Fig. 87. Anapleus semen (Lewis). A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto,lateral view. E: Male genitalia, 9th and10th tergites, dorsal view. F: Ditto,lateral view. G: Ninth sternum (spicule), dorsal view.

longitudinal oblong, separated by 0.3 times their diameter laterally, and become finerand sparser on the median area; its sides completely striate and strongly carinate.
Protibia dilated, its outer margin angulate about middle.
Male genitalia as shown in Fig. 87.
Specimens examined. Holotype of Tribal/us semen Lewis, 1884, labelled "Type","Miyanoshita, Japan, 1880" and "Tribal/us semen Lewis" (The Natural History Museum, London).
Holotype of Anapleus japonicus Nakane, labelled "Holotype", "Rausu, Shiretoko (Hokkaid6)
8/vii/1958, T. Nakane." and "Anapleusjaponicus, Det. T. Nakane, 1961" (NA).
[Hokkaid6] Id" 1 Sf, H6heiky6, J6zankei, 16/vii/1988, M. Ohara.
[Honshu] Tochigi-ken: 1 ex.,Nikko,ll-13/vi/1967,H. Takizawa. Tokyo-to: 1 ex., Takao-san,
27/vi/1970, H. Takizawa.

Distribution. Japan (Hokkaido, Honshu).
Remarks. Anapleus semen is a relatively large species; it is similar to A.
nakanei and A. nomurai; however, the rather larger body and the dense punctation on
the elytra will distinguish it from them.
This species is found on logs.

157

Fig. 88. Anapleus nakanei M. Ohara, sp. nov., adult. A: Dorsal view.B: Elytra around scutellum. C:
Punctation of medio-basal area of elytron. D: Prosternum, ventral view. E: Lateral side ofmetasternum. F: Meso- and metaslerna, ventral view.

158

Table 19. Biometric data of Anapleus nakanei M. Ohara and A. nomurai M. Ohara.

APW
PPW
PL
EL
EW

ProW
ProL
PyL
PTL
MSTL

MTIL

A. nakanei

A. nomurai

0.49-0.57 (0.54±0.01) 6
1.14-1.26 (1.20±0.02) 6
0.45-0.55 (0.51±0.01) 6
1.18-1.32 (1.24±0.02) 6
1.42-1.52 (1.46±0.02) 6
0.61-0.71 (0.65±0.01) 6
0.18-0.22 (0.20±0.01) 6
0.33-0.39 (0.38±0.01) 6
0.55-0.59 (0.56±0.01) 6
0.53-0.57 (0.56±0.01) 6
0.59-0.67 (0.62±0.01) 6

0.53-0.59 (0.56±0.02) 2
1.18-1.28 (1.23±0.03) 2
0.57-0.61 (0.59±0.0l) 2
1.22-1.26 (1.24±0.01) 2
1.44-1.56 (1.50±0.04) 2
0.67-0.71 (0.69±0.01) 22
0.24(0.24)
0.33-0.35 (0.34±0.01) 2
0.51-0.55 (0.53±0.01) 22
0.51(0.51)
0.51-0.59 (0.55±0.03) 2

F
O.25mmI

Fig. 89. Anapleus nakanei M. Ohara, sp. nov. A: Aedeagus, lateral view. B: Ditto, dorsal view. C:
Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia,
9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: Ninth sternum (spicule), dorsal
view.

Anapleus nakanei M. Ohara, sp. nov.
(Fig. 88, 89)
Japanese name: Nara-tsubu-emma-mushi.

Description. Male and female. Body weakly convex, oval, dark brown andshining; tarsi and antennae light brown. Body length, PPL, 1.89 - 2.07 mm (1.99 ±159

0.03, n=6), PEL, 1.73 - 1.87 mm (1.81 ± 0.02, n=6). Width, 1.42 - 1.52 mm (1.46± 0.02, n=6). Biometric data are given in Table 19.
Head densely and deeply punctate. Epistoma convex. Front feebly depressed
between eyes, without stria.
Pronotum (Fig. 88A) strongly convergent apically on sides. Disk sparsely
covered with moderate and shallow punctures, which are separated by about 1.5 (1- 2)
times the diameter, the punctures becoming denser on lateral sides where they areseparated by 0.75 times the diameter, and with a shallow depression behind each eye.
Marginal stria weakly crenate and complete.
Epipleural marginal stria complete with dense punctures. Elytral marginal stria
entirely and deeply impressed, and separately punctate. Impunctate narrowly along
epipleural marginal stria. Area between epipleural and elytral marginal striae with 1 or 2
rows of moderate punctures. Lateral side of elytron feebly elevated, and with 2 rows ofsparse and moderate punctures. Surface of e1ytra (Fig. 88B, C) sparsely and coarsely

Fig. 90. Anapleus nomurai M. Ohara, sp. nov., adult. A: Dorsal view. B: Head, frontal view. C:
Elytra around scutellum. D: Punctation of medio-basal area of elytra.

160

punctate, the punctures a little coarser than those of the pronotum, shallow andseparated by about 1.5 (1 - 2) times the diameter; impunctate narrowly on basal 112
near lateral 113 - 112 of elytron, the impunctate area slightly striate; finely punctate in anarrow band along suture; apical 114 of elytra densely covered with shallow rugae;
extreme apex impunctate. Oblique humeral stria present on basal 113. All dorsal striae
obsolescent, but the 1st (or 2nd) stria entirely and feebly impressed, and the 2nd (or3rd) indistinctly on basal 112.
Propygidium densely covered with coarse and deep punctures, which areseparated by 112 their diameter. Pygidium coarsely and sparsely punctate, the punctures
being separated by about 1.5 (1- 2) times the diameter, but becoming denser apically.
Anterior margin of prostemallobe (Fig. 88D) straight and indistinctly margined.
Disk of lobe densely covered with coarse punctures. Prostemal process nearlyquadrate, its disk slightly convex, densely covered with coarse punctures except on aband along basal margin, and with complete carinal striae.
Anterior margin of mesosternum nearly straight (Fig. 88F). Mesosternum short(width: length = 16 : 5), densely and coarsely punctate. Lateral side of mesosternum(Fig. 88E) deeply marginate. Meso-metasternal suture with strongly crenate stria. Diskof metasternum convex medially, and coarsely with shallow punctures, which areseparated by 0.5 - 1 times the diameter, the punctures becoming finer on median area.
Lateral metastemal stria carinate, extending obliquely and posteriorly, reaching near
middle of metastemal-metepistemal suture.
Intercoxal disk of 1st abdominal sternum completely striate on each side, andfinely and densely punctate, the punctures becoming coarser towards lateral and anterior
margms.
Protibia dilated, its outer margin arcuate.
Male genitalia as shown in Fig. 89.
Specimens examined. Ho)otype, if, Yoshino-yama, Nara-ken, Honshu, Japan, 27/v/1985, S.
Nomura (EUill).
Paratypes. [Honshu] Fukushima-ken: 2 exs., Kaimon-rinda, Towada, Kami-Ogawa, Ogawamachi, Iwaki-shi, 19/vii/1993, A. Matsushima (1 ex., in Matsushima Collection). Nara-ken: 2d"d",
Yoshino-yama, 27/v/1985, S. Nomura. Kyato-fu: 1d", Kurama, 17/vi/1954, T. Nakane (NA); 1!j1.,
Kunogahata, 8/vi/1954, T. Nakane (NA). [Kyushu] Fukuoka-ken: 1d", Mt. ShOji, 28/iv/1964, S.
Nakano (NA). Gita-ken: 1 ex., Saeki, no date, K. Kurosa (NA).

Distribution. Japan (Honshu, KyushQ).
Remarks. Anapleus nakanei is very similar to A. nomurai, but differs from thelatter by the punctation of the pronotum, elytra and meso- and metastema (see thedescription of A. nomurai).
Anapleus nomurai M. Ohara, sp. nov.
(Fig. 90,91)
Japanese name: Okayama-tsubu-emma-mushi.

Almost agrees with Anapleus nakanei, but differs by the following characters:
punctation on pronotum and elytra deeper (Fig. 90C, D) than in nakanei; lateral sidesof elytra not elevated; punctation of intercoxal disk of meso- and metasternal and 1st
abdominal sternum sparse, coarse and very deep, with fine punctures intermingled(Fig. 9IB, C).
161

Fig. 91. Anapleus nomurai M. Ohara, sp. nov. A: Prosternum, ventral view. B: Meso- andmetasterna. C: Lateral side of metasternum. D: Propygidum and pygidium, caudal view.

Body length, PPL, 2.09 - 2.13 mm (2.11 ± 0.01, n=2), PEL, 1.89 - 12.93 mm(1.91 ± 0.01, n=2). Width, 1.44 - 1.56 mm (1.50 ± 0.04, n=2). Biometric data are
given in Table 19.
Specimens examined. Ho!otype, Sf, Yoshino-yama, Nara-ken, HonshU, Japan, 27/v/1985, S.
Nomura (ElliU).
Paratype. [Okayama-ken] Sf, Ikura-nimi-shi, 211v/1985, S. Nomura.

Distribution. Japan (Honshu).
Remarks. This species is strikingly similar to A. nakanei, but can be
distinguished easily by having 2 kinds of punctures on the meso- and metasterna.
This species seems to live in litter.
Anapleus hagai M. Ohara, sp. nov.
(Fig. 92, 93, 94)
Japanese name: O-tsubu-e=a-mushi.

162

Description. Female. Body oval, convex, black and shining; tarsi, tarsus andantennae light brown. Body length, PPL, 2.45 - 2.84 mm (2.65 ± 0.07, n=4), PEL,
2.29 - 2.62 mm (2.49 ± 0.06, n=4). Width, 1.96 - 2.09 mm (2.03 ± 0.02, n=4).
Biometric data are given in Table 20.
Head (Fig. 92B) densely and coarsely punctate. Epistoma convex. Frontdepressed between eyes, without striae.
Pronotum strongly convergent anteriorly on sides. Disk densely covered withcoarse and deep punctures, which are separated by about 0.3 times their diameter.
Marginal stria weakly carinate and complete.
Epipleural marginal stria complete and strongly crenate. Elytral marginal stria
carinate, complete and crenate. Impunctate narrowly along epipleural marginal stria.
Area between these striae with 2 or 3 rows of coarse and dense punctures. Surface of
elytra (Fig. 92C, D) sparsely covered with large, shallow, and ocelloid punctures,
which are separated by about 1.5 (l - 2) times the diameter medially, the punctures
becoming denser and a little finer laterally; dense fine punctures narrowly along thesutural line; apical 115 of elytra with shallow longitudinal rugae; extreme apex

Fig. 92. Anapleus hagai M. Ohara, sp. nov., adult. A: Dorsal view. B: Head, frontal view. C: Elytra
around scutellum. D: Punctation of medio-basal area of elytron.

163

Fig. 93. Anapleus hagai M. Ohara, sp. nov., adult. A: Prosternum, ventral view. B: Punctation ofintercoxal disk of metasternum. C: Meso- and metasterna. D: Protibia, ventral view.

Table 20. Biometric data of Anapleus hagai M. Ohara.
APW
PPW
PL

EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

0.69-0.78
1.50-1.67
0.82-0.92
1.57-1.70
1.96-2.09
0.92-1.08
0.29-0.39
0.46-0.59
0.65-0.78
0.69-0.78
0.82-0.85(0.74±0.02) 4(1.61±0.03) 4(0.86±0.02) 4(1.65±0.03) 4(2.03±0.02) 4(1.01±0.03) 4(0.35±0.02) 4(0.54±0.02) 4(0.74±0.02) 4(0.73±0.02) 3(0.83±0.01) 4

impunctate. Dorsal striae obsolescent, but the 1st dorsal stria slightly impressed on
mediobasal 114, and the 2nd slightly on about the basal 112.
Propygidium densely covered with moderate and deep punctures, which are aslarge as those on the elytral apices. Pygidium moderately punctate, the punctures164becoming denser and finer towards apical margin. Disk of pygidium convex and feebly
depressed behind tip.
Anterior margin of prosternallobe (Fig. 93A) straight, and indistinctly margined.
Prosternal process nearly quadrate, its disk coarsely punctate, sometimes with finepunctures intermingled, and with complete carinal striae (Fig. 93A).
Anterior margin of mesosternum feebly and outwardly arcuate in median portion.
Mesosternum short (width: length = 24 : 7), densely and coarsely punctate, with finepunctures intermingled. Lateral sides distinctly margined. Meso-metasternal suture withstrongly crenated stria. Mesosternum even, densely covered with large and deeppunctures, which are separated by 0.5 - 1 times the diameter, with fine puncturesintermingled (Fig. 938). Lateral metasternal stria extending obliquely and posteriorly,
reaching near middle of metasternal-metepisternal suture. Intercoxal disk of 1st
abdominal sternum completely striate on each side, densely covered with moderatepunctures, which are coarser along the anterior margin.
Protibia (Fig. 930) dilated, its outer margin with an obtuse angle at middle.
Male genitalia as shown in Fig. 94.
Specimens examined. Holotype,Sf, Ko-tombetsu, Naka-tombetsu, Hokkaid6, Japan,

25/vii/1987, K. Haga (ElHU).

0.25 mm, A-8

':

"

..

"

....

~
o ,'-------'
Fig. 94. Anap/eus hagai M. Ohara, sp. nov. A: Aedeagus, lateral view. B: Ditto, dorsal view. C:
Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia,
9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: 9th sternum (spicule), dorsal view.

165Paratypes. [Hokkaido] 1!f, Shintoku, 19/viii/1989, K. Haga; 1!f, Mt. Yotei, Kutchan,
29/vii/1989, N. Yasuda. [Honshu] Aomori-ken: Id", Oirase-gawa, Fukura-machi, I/viii/1988, S.
Yamauchi.

Distribution. Japan (Hokkaid6, Hoshfi).
Remarks. Anapleus hagai is easy to recognize by its large size and the depressed
pygidium.
TRIBE BACANIINI KRYZHANOVSKIJ AND REICHARDT, 1976

The Japanese species included in the tribe Bacaniini have not been sufficiently
studied until now. Probably 5 or more species occur in Japan. In this papaer, theknown Japanese species are only listed.
GENUS BACANIUS J. L. LECONTE, 1853
Bacanius J. L. LeConte, 1853: 291 [type species: Bacanius tantillus J. L. LeConte, 1853: 291,
designated by Bickhardt, 1916: 71]; iacordaire, 1854: 271; Marseu!, 1856: 576; Schmidt,
1885a: 332; Reitter, 1886: 273; 1909: 297; GangJbauer, 1899: 409; Cooman, 1936: 136;
Reichardt, 1941: 84, 117; Wenzel and Dybas, 1941: 438; Wenzel, 1944: 54,95-101; Arnett,
1962: 375,380; WitzgalJ, 1971: 175; Kryzhanovskij and Reichardt, 1976: 267; Mazur, 1984:
118.

SUBGENUS MULLERISTER COOMAN, 1936
Mullerister Cooman, 1936: 137 [type species: Bacanius tonkinensis Cooman, 1936: 135, originallydesignated]; Mazur, 1984: 119.
Muellerister Kryzhanovskij and Reichardt, 1976: 270 (unjust. emend.).
Neobacanius G. MUller (part.), 1925: 20; Reichardt, 1941: 118,121.

Bacanius (Mullerister) niponicus Lewis, 1879
Bacanius niponicus Lewis, 1879: 461 [Japan: Nagasaki].
Bacanius (Mullerister) niponicus: Mazur, 1984: 119; Hisamatsu, 1985b: 224, pI. 40, no. 22.
Bacanius (Muellerister) niponensis (sic): Kryzhanovskij et Reichardt, 1976: 268 [key].

Japanese name: Aka-tsubu-emma-mushi.

Distribution. Japan; Taiwan.
SUBGENUS BACANIUS 1. L. LECONTE, 1853

Bacanius (Bacanius) mikado (Lewis, 1892)
Abraeus mikaM Lewis, 1892b: 356 [Japan].
Bacanius mikado: Schmidt, 1893b: 238.
Bacanius (Bacanius) mikaM: Kryzhanovskij and Reichardt,
Hisamatsu, 1985b: 224, pI. 40, no. 21.

Japanese name: Ko-aka-tsubu-emma-mushi.

166

1976:

269; Mazur, 1984:

122;

Distribution. Japan (Honshu, Shikoku, Kyushu).
TRIBE PAROMAUNI REITIER, 1909

Key to the Japanese genera of the tribe Paromalini
1(2)
2(1)
3(4)

Elytral disk with nonnal dorsal striae................................. Genus Carcinops Marseul, 1855
Elytral disk punctate, without nonnal dorsal striae except for vague rudiments.
Propygidium with a transverse line. Metasternal lateral stria not extended posteriorly. Alltibiae dilated. . ......................................................... Genus Pachylomalus Schmidt, 1879
4(3) Transverse line of pygidium not visible. Metastemal lateral stria extended posteriorly. Protibia
dilated, but others narrow.
5(6) Prostemal keel without carinal stria.................................. Genus Paromalus Erichson, 1834
6(5) Prostemal keel with carinal stria.
7(10) Frontal stria complete.
8(9) Basal margin of pronotum without an irregular row of large punctures. Body oblong-oval, andless convex. .. ......................................................... Genus Platylomalus Cooman, 1948
9(8) Basal margin of pronotum with an irregular row of large punctures. Body oval, and convex .
................................................................ ................. GenusAustralomalus Mazur, 1981
10(7) Frontal stria interrupted behind labrum .............................. Genus Eulomalus Cooman, 1937
GENUS CARCINOPS MARSEUL, 1855
Carcinops Marseul, 1855: 83 [type species: Dendrophilus quatuordecimstriatus Stephens, 1835: 412,
designated by Bickhardt, 1917: 113]; Schmidt, 1885b: 283; Ganglbauer, 1899: 373; Reitter,
1909: 287; Jakobson, 1911: 640,647; Bickhardt, 1916-17: 108, 112-114; 1921: 149; Arnett,
1962: 375,380; Witzgall, 1971: 177; Kryzhanovskij and Reichardt, 1976: 263; Mazur, 1984:
127.

SUBGENUS CARCINOPS MARSEUL, 1855

Carcinops (Carcinops) pumilio (Erichson, 1834)(Fig. 95, 96, 97)
Paromalus pumilio Erichson, 1834: 169 [Spain, Egypt, North America]; Wollaston, 1854: 213;
Redtenbacher, 1858: 313.
Dendrophilus quattuordecimstriatus Stephens, 1835: 412, synonymized by Gemminger and Harold,
1868: 777.
Paromalus quattuordecimstrataus: Hom, 1873: 308.
Dendrophilus pumilio: Dejean, 1837: 143.
Hister nanus J. E. LeConte, 1845: 61, synonymized by Gemminger and Harold, 1868: 777.
Dendrophilus nanus Dejean, 1837: 143 [nom. nud.].
Phelister nanus: Marseul, 1857: 476.
Carcinops quattuordecimstriatus: Wollaston, 1864: 166; Kolbe, 1909: 21; Scott, 1913: 225;
Bickhardt, 1917: 114; Hinton, 1945: 334.
Carcinops quatuordecimstriatus: Gangl bauer, 1899: 374; Reitter, 1909: 287; Jakobson, 1911: 647;
Auzat, 1916-25: 106; Horion, 1949: 347.
Epierus krujanensis Mader, 1921: 181, synonymized by Mader, 1927: 193.
Carcinops pumilio: Marseul, 1855: 91; 1873: 221 [Japan, Nangasaki]; Seidlitz, 1891: 216; Kolbe,
1909: 21; Kuhnt, 1913: 371; Wenzel, 1955: 608,617; Witzgall, 1971: 177; Kryzhanovskij andReichardt, 1976: 264; Hisamatsu, 1985b: 224, pI. 40, no. 24.

Japanese name: Kuro-chibi-emma-mushi.

167

A good description is given by Hinton (1949) under the name Carcinops
quattuordecimstriatus.

Description. Body length, PPL, 1.98 - 2.57 mm (2.39 ± 0.03, n=20), PEL,
1.76 - 2.40 mm (2.19 ± 0.03, n=20). Width, 1.30 - 1.74 mm (1.60 ± 0.02, n=20).
Biometric data are given in Table 21. Marginal elytral stria well impressed andcomplete. Marginal epipleural stria complete. Epipleura with another stria between
marginal elytral and epipleural striae, which is strongly crenate with coarse puncturesand complete. In Japanese specimens, the apical non-striate belt of elytra only with finepunctures, not covered with coarse ones. Apical 112 of 1st and 2nd intervals of dorsal
striae and apical 113 of the 3rd - 5th usually covered with coarse and fine punctures.
Prostemal lobe densely and coarsely punctate, the punctures becoming sparser
mediobasally. Laterobasal angles of the lobe deeply margined, and lateral 113 of suture
between keel and lobe strongly and deeply with a longitudinal impression, which

Fig. 95. Carcinops (Carcinops) pumilio (Erichson). A: Adult, dorsal view. B: Ditto, ventral view.
C: Head, frontal view. D: Propygidium and pygidium. E: Protibia, dorsal view.

1680.25mmFig. 96. Carcinops (Carcinops) pumilio (Erichson). A: Aedeagus, lateral view. B: Ditto, dorsal
view. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Ditto,
ventral view. F: Male genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. G:
Ditto, lateral view.

Fig. 97. Collection sites of Carcinops (Carcinops) pumilio (Erichson) in Japan.
169

Table 21. Biometric data of Carcinops pumilio (Erichson) and Australomalus montivagus (Lewis).

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MITL

C.pumilio

A. montivagus

0.54-0.74 (0.68±0.01) 20
1.15-1.49 (1.39±0.02) 20
0.56-0.78 (0.73±0.01) 20
1.18-1.49 (1.39±0.02) 20
1.30-1.74 (1.60±0.02) 20
0.78-0.96 (0.85±0.01) 20
0.27-0.42 (0.36±0.01) 20
0.34-0.49 (0.44±0.01) 20
0.51-0.69 (0.58±0.01) 19
0.54-0.69 (0.59±0.01) 19
0.51-0.74 (0.68±0.01) 20

0.88-1.10 (0.99±0.01) 16
1.88-2.38 (2. 11±0.04) 16
1.00-1.25 (1.17±0.02) 16
2.07-2.57 (2.29±0.04) 16
2.13-2.95 (2.51±0.05) 16
1.07-1.57 (1.24±0.03) 16
0.38-0.56 (0.42±0.01) 16
0.47-0.75 (0.67±0.02) 16
0.75-0.94 (0.86±0.02) 16
0.69-0.88 (0.81±0.02) 16
0.82-1.07 (0.95±0.02) 16

extends obliquely and posteriorly on prosternum. Protibia (Fig. 95E) with 2 large
denticles on apical 112 and 3 or 4 small ones on basal 112 of outer margin, the interval
between large ones wide, being about 113 as long as protibia.
Male genitalia as shown in Fig. 96.
Specimens examined. [Hokkaido] 1 ex., Kun'neppu, 16/ix/1989, H. Torikura; 4~ ~ , Bibi,
28/vi/1987, M. Ohara; 1 ex., Naganuma, 4/viii/1989, A. Iwasaki; 2 exs., Hitsuji-ga-oka, Sapporo,
20/vii/1985, M. Ohara; 1 ex., Sapporo, 27/vii/1985, M. Suwa; 1 ex., Ditto, 2/vii/1987, S. Kondo; 1ex., Hakken-zan, Sapporo, 13/vii/1985, M. Ohara; 16 ~ ~ , Jozankei, 27, 30/viii, l/x/1985, 1986, M.
Ohara; 6 exs., Misumai, Sapporo, 14/vii/1986, M. Ohara; 1 ex., Nakagoya, Ishikari-Tobetsu,
8/vii/ 1990, S. Shiyake; 1 ~ , 2 exs., Zenibako, 30/viii/1985, M. Ohara; 1 ex., Ditto, 19/viI1979, N.
Nishikawa.
[Honshu] Ibaraki-ken: 1 ex., Tsukuba-Gakuen-Toshi, 3/v/1979, M. Ohara. Tochigi-ken: 1 ex.,
Miyuki, Utsunomiya-shi, Il1v/I980, T. Kumazawa (TPM). Tokyo-to: 1 ex., Mt. Jimba, 16/viI1967,
H. Takizawa; 1 ex., Kinuta, 28/vi/I964, Y. Kurosawa (NSMf); 3 exs., Tokyo, 24/v11971 , nocollector's name (NSMf); 1 ex., Kunitachi, 3/vii/1958, S. Nomura (NSMf). Kanagawa-ken: 1 ex.,
Fudakake, Tanzawa, IS/vii, 1971, M. Ishida; 31 exs., Hatano, 5/vi/1971, 7/viii/1972, 22/vi/1974,
I6/ix/1976, 28/vI1977, 24/vi/1984, 7/vii/1985, H. Takizawa; 2 exs., Izu, Daruma-yama, 25/viiI1966,
M. Ishida. Gumma-ken: 1 ex., Osawa, 15/vi/1963, S. Kondo. Nagano-ken: 1 ex., Shimashima,
Shinano, 29/viiI1948, S. Osawa. Niigata-ken: 2 exs., Senami, northern Echigo, 14/ix/1986, K. Baba;
1 ex., Kurokawa, Echigo, 1O/viii/1958, K. Baba; 1 ex., Mizutanizawa, Mt. Mino-ji, northern Echigo,
7/vi/1958, K. Baba. Aichi-ken: 2 exs., Shikatsu-mura, Owari, 25/xl 1945, 27/iv/1946, S. Osawa (NA);
1 ex., Meito-ku, Nagoya, 13/ix/1986, T. Tanabe; 1 ex., Higashiyama, Nagoya, Owari, 13/v/1949, T.
Nakane (NA); 1 ex., Nagoya, 13/vi/1947, S. Osawa (NA); 1 ex., Ditto, 30/ix/1945, T. Nakane (NA); 4
exs., Ditto, viii/1950, no collector's name. Gifu-ken: 1 ex., Mino-shi, 19/v/1979, Y. Takai. Osaka-fu:
2 exs., Nishinari, 12/vi/1949, 17/vl 1951, K. Sawada (NA). Nara-ken: 1 ex., Kasuga, 22-23/ix/1968, S.
Kondo & T. Okutani. Fukui-ken: 1 ex., Inumi, Oi, 9/v/1987, T. Ito. Shiga-ken: 1 ex., Omi-hachiman,
IS/xl 1990, S. Shiyake. Tottori-ken: 15 exs., Daisen, 12/x/1987, M. Ohara.
[Shikoku] Kagawa-ken: 1 ex., Toyonaka-cho, Sampo-gun, 4/vl 1980, S. Ohmomo (NSMf).
[Kyushu] Fukuoka-ken: 1 ex., Hakozaki, 3/vii/1946, Y. Kurosawa (NA); 1 ex., Mt. Adachi,
311v/1969, S. Nakao (NA). Kumamoto-ken: 60 exs., Yokoshima-chO, Tamana-shi, 9/vi/1986, E.
Matsui; 3 exs., Ue-mura, Kuma-gun, 6/vi/ 1986, E. Matsui. Miyazaki-ken: 1 ex., Takachiho-no-mine,
9/vii/1984, S. Masuda; 99 exs., Ditto, 19, 27/vii/1983, 10/vii/1984, M. Ohara; 6 exs., Ebino-kogen,29/iv/1986, E. Matsui. Kagoshima-ken: 3 exs., Toso, Kagoshima-shi, 7/iv/1984, M. Ohara; 1 ex.,
Korimoto, Kagoshima-shi, 16/vi/1981, Y. Takai; 4 exs., ChOjo-bokujo, Kushikino, 20/v/1984, M.
Ohara; 51 exs., Iriki-bokujo, 17/iii/1985, K. Tomiyama; 1 ex., Kirishima, 29/vii/1971, T. Nakane(NA); 15 exs., Kaimon, Satsuma, lIvi/1983, K. Tomiyama (NA); 14 exs., Kamijo-Kogen, Kihokucho, 26/v/1985, Y. Harada; 1 ex., Chosa, 22/v/1984, M. Ohara.
[Nansei Isles] Okinawa-Honto Is.: 1 ex., Nago sea park, 22/iiil 1983, T. & T. Nakane (NA). IO170

tori-shima Is.: 5 exs., 128°15',27°50', 13/vii/1991, Y. Kusui.
[Ogasawara Isles] Haha-jima Is.: 1 ex., Kita-mura, vii-ix/1976, Y. Yoshida (NA).

Distribution (Fig. 97). Japan (Hokkaido, Honshu, Shikoku, Kyushu, Izu Isles,
Nansei Isles, Ogasawara Isles); nearly cosmopolitan.
Remarks. Carcinops pumilio is easily recognized by the characteristics given in
the key and description: it cannot be confused with any other Japanese species of thetribe Paromalini.
The species occurs on various decaying organic matters, such as compost piles,
cow, pig and hen manure, donkey and human excrement, carrion, under bark, etc.
Hinton (1945) summed up the habitats of this species and also recorded it in waste
grain in the store house.
Jones (1929) noted that this species is an intermediate host of Hymenolepis
carioca.
GENUS AUSTRALOMALUS MAZUR, 1981
Australomalus Mazur, 1981b: 379 [type species: AustralomalusfiJianus Mazur, 1981b: 379].

Australomalus montivagus (Lewis, 1892)(Fig. 98,99, 100)
Paromalus montivagus Lewis, 1892a: 37 [Japan: Fujisan].
Platylomalus montivagus: Kryzhanovskij and Reichardt, 1976: 275; Hisamatsu, 1985b: 224, pI. 40,
no. 26.
Australomalus montivagus: Mazur, 1984: 133.

Japanese name: Maru-chibi-hirata-emma-mushi.

Description. Male and female. Body length, PPL, 3.57 - 3.70 mm (3.62 ± o.m,n=4), PEL, 3.39 - 3.51 mm (3.43 ± 0.03, n=4). Width, 2.13 - 2.95 mm (2.51 ±
0.05, n=16). Biometric data are given in Table 21. Body oval, black and shining; tarsi,
mouthparts and antennae reddish brown.
Head sparsely covered with coarse punctures, which are separated by 2 or 3
times their diameter, the punctures becoming a little finer anteriorly; interspace among
the punctures with alutaceous microsculptures.
Pronotum (Fig. 98A) sparsely covered with coarse, somewhat longitudinal
oblong, and shallow punctures, which are separated by 2 - 4 times their diameter, andwith other fine punctures intermingled; interspace among these punctures withalutaceous microsculptures.
Epipleural marginal stria fine and complete. Elytral marginal stria (Fig. 98A) (inoriginal description, epipleural stria) strongly impressed and carinate. Epipleura
impunctate, with alutaceous microsculptures. Oblique humeral stria present on basal
1/4.

Propygidium densely covered with large, round and shallow punctures, which
are unevenly separated by 0.3 - 1 times the diameter. The punctation is not similar tothat of the pronotum in disagreement with the original description.
The male and female are confused in the original description; in reality, thepygidium is sparsely clothed with fine punctures and with alutaceous microsculptures in
the male; in the female it has a coarse vermicular sculpture (Fig. 98C).
171

";J. _ : ··'Y".:.
.
~

>1'
':.,

..•.....

•

..

"'iii

•.. 'i'C

Fig. 98. Australomalus montivagus (Lewis). A: Pronotum and left elytron, dorsal view. B:
Prosternum, mesosternm, metasternum and 1st abdominal sternum. C: Pygidium.

Fig. 99. Australomalus montivagus (Lewis). A: Aedeagus, lateral view. B: Ditto, dorsal view. C:
Male genitalia, 8th tergite, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and 10th
tergites, dorsal view. F: Ditto, lateral view. G: Male genitalia, 8th sternum, dorsal view. H:
Ditto, lateral view. I: Ditto, caudal view.

Anterior margin of prostemaI lobe (Fig. 98B) truncate, its marginal stria complete
and inwardly curved basally; lateral side of the lobe with another short strongly carinate
margin; disk of the lobe evenly and finely punctate, the punctures being separated byabout twice their diameter and becoming finer and sparser around margin. ProstemaI172

Australomalus montivagus

Fig. 100. Collection sites of Australomalus montivagus (Lewis) in Japan.

process completely with carinal striae, which are strongly carinate and nearly parallel,and of which the basal ends are usually not jointed with each other along the basal
margin of the lobe.
Anterior margin of mesosternum (Fig. 98B) deeply emarginate at middle, its
marginal stria carinate and nearly complete, being shortly interrupted at middle; disk
sparsely covered with fine punctures. Meso-metasternal suture strongly carinate,complete and widely sinuous. Disk of metasternum sparsely covered with large, oblongand shallow punctures, which are separated by 2 - 5 times their diameter, and become
finer medially; area along mid line finely punctate, sometimes densely punctate;interspace among punctures with alutaceous microsculpture; surface feebly depressed
mediobasally. Lateral metasternal stria carinate, extending obliquely and posteriorly,
attaining to near basal 113 of metasternal-metepisternal suture.
Intercoxal disk of 1st abdominal sternum sparsely covered with coarse punctures,
which are separated by 2 - 5 times their diameter.
Protibia dilated, with 5 or 6 denticIes on outer margin.
Male genitalia as shown in Fig. 99.
Specimens examined. [Hokkaido] 1 ~ , Nopporo, 301v11987, S. Kudo; 10"', Ditto, 211v11985,
M. Ohara; 1 ~,Maruyama, 28liiiJ?, no collector's name (EIHU).
[HonshU] Miyagi-ken: 10"', Moriyasu, Akiu-mum, 241vi11951, no collector's name (NSMf).
Fukushima-ken: 10"', Hirasawa-rindo, Hinoemata-mura, 211v11988, H. Hag a; 10'" Iff, Hizumari,Tateiwa, Minami-Aizu, 81v11949, Y. Kurosawa (NSMT); 10"', Yunohana, Tateiwa, Minami-Aizu,
71v11949, Y. Kurosawa (NSMf); 1 ex., Otakiyama, Ornata, Onuma, 151vii11950, Y. Kurosaw(NSMf). Yamagata-ken: Iff, Tokuami, Kitaoguni, Oguni, 21vi11957, I. K. (NSMf). Tochigi-ken: 1ex., Usukubo-Daira, Oku-Kinu, Kuriyama-mura, 171v11989, K. Satoh (TPM). Tokyo-to: Iff, Nippara,
Okutama, 121iv11983, M. Ishida. Kanagawa-ken: 10"', Owaki-dani, Hakone, 191ix11976, Y. Hirano.
Shizuoka-ken: 10"', Mt. Amagi, 271v11956, H. Hayashi. Niigata-ken: 1!j1., Sasagamine, MyokoKogen, 291viii11932, Y. Kusui. Gifu-ken: 20"'0'" Iff, Karnmuri-yama-toge, Tokuyama, lIvi11986, Y.
Takai; 10"', Ditto, 11 v11986, Y. Takai; 10"', Ditto, 101x11986, Y. Takai. Nara-ken: 10"', Kasuga,
91iv11950, S. Ueno; 20"'0'" Iff, Ditto, 201v11951, K. Sawada. Kyoto-fu: 10"', Kibune, 91iii11950, A.
Nobuchi; 10'" 1 ~,Ditto, 91vi11953, A. Nobuchi. Wakayama-ken: Iff, Mt. Gomanodan, 21viii11957,
H. Kono (Osaka Mus.).
[Kyilshil] Fukuoka-ken: 2 exs., Mt. Adachi, 71vii11982, 28IivI1964, S. Nakao (NA). Oita-ken:173

1 ex., Mt. Kuju, 20/v/1979, S. Nakao (NA). Miyazaki-ken: 2 exs., Mt. Sobo, 1O-12/v/1969, H.
Takizawa.

Distribution (Fig. 100). Japan (Hokkaid6, Honshu, Shikoku, Kyushil). Newly
recorded from Hokkaido and Kyushu.
Remarks. Australomalus montivagus is quite peculiar by its very broadly oval
and convex form and the transverse punctured row on the posterior margin of the
pronotum; it cannot be confused with any other species of the tribe.
No details are known about habitat requirements of this species. Some specimens
were collected in rotting woods.
GENUS PLATYLOMALUS COOMAN, 1948
Platylomalus Cooman, 1948: 134 [type species: Hister complanatus Panzer, 1797].
Paromalus Lewis, 1907a: 316, nec Erichson, 1834: 167, synonymized by Cooman, 1948: 134.

Key to the Japanese species of the genus Pia tyloma Ius
1(4)
2(3)
3(2)
4(1)
5(6)
6(5)

Apical end of marginal elytral stria extending to suture of elytron, and curved at medio-posterior
angle of elytron, then extending anteriorly.
Disk of mesosternum with a transverse stria, which is trapezoid. Pygidium of female deeply
engraved with furrows ............................................................. P.fujisanus (Lewis, 1892)
Disk of mesosternum without a transverse stria. pygidium of female densely punctate at apex,
not engraved with furrows ......................................................... P. viaticus (Lewis, 1892)
Apical end of marginal elytral stria not attaining to suture of eiytron.
Elytra coarsely punctate, the area along oblique humeral depression with large punctures. Diskof mesosternum without a transverse stria. .................... .......... P. kusuii M. Ohara, sp. nov.
Elytra rather evenly punctate, without large punctures. Disk of mesosternum with a transverse
stria. (In P. mendicus, the transverse stria sometimes absent.)

spine. Pygidium offemale engraved with a large and ocelloid sculpture ............................... ..
.......................................................................................... P. persimilis (Lewis, 1888)
8(7)Disk of mesosternum with a transverse stria, which is smoothly curved and sometimes
interrupted. Mesotibia with 4 spines. Metatibia with 2 spines.
9(10) Body length 3.0 - 3.5 mm. Marginal stria of pronotum nearly complete anteriorly. Pygidium of
female engraved with irregular furrows except on its margin .......... P. niponicus (Lewis, 1899)
10(9) Body length 1.8 - 2.3 mm. Marginal stria of pronotum broadly interrupted at anterior middle.
Pygidium of female engraved with a pair of curved sculptures. ....... P. mendicus (Lewis, 1892)

PlatylomalusfuJisanus (Lewis, 1892)(Fig. 101, 102)
Paromalusfujisanus Lewis, 1892a: 34 [Japan: Honshu]; Cooman, 1937: 160.
Platylomalus fujisanus: Hisamatsu, 1968: 31 [Shikoku]; 1985b: 224 [photo]; Kryzhanovskij andReichardt, 1976: 276; Mazur, 1984: 134.

Japanese name: Fuji-chibi-hirata-emma-mushi.

Description. Male and female. Body length, PPL, male, 1.62 - 1.97 mm (1.82 ±0.03, n=13), female, 1.56 - 2.01 mm (1.76 ± 0.04, n=13), PEL, male, 1.52 - 1.85
mm (1.70 ± 0.03, n=13), 1.50 - 1.77 mm (1.64 ± 0.02, n=13). Width, male, 1.00 1.18 mm (1.10 ± 0.02, n=13), female, 0.99 - 1.14 mm (1.00 ± 0.01, n=13).
Biometric data are given in Table 22. Body oblong, black and shining; femora, tarsi,
mouthparts and antennae red brown.
174

3
3

•
c

Fig. 101. Platylomalusfujisanus (Lewis). A: Pronotum and left elytron, dorsal view. B: Prosternum,
mesostemm, metasternum and 1st abdominal sternum. C: Pygidium.

O.25mm,Fig. 102. Platylomalusfujisanus (Lewis). A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Apex
of aedeagus, caudal view. D: Male genitalia, 8th tergite, dorsal view. E: Ditto, lateral view. F:
Male genitalia, 9th and 10th tergites, dorsal view. G: Ditto,lateral view. H: Male genitalia, 8thsternum, dorsal view. I: Ditto,lateral view. J: Ditto, caudal view.
175

Frontal stria of head weakly carinate and complete. Disk of head sparsely clothedwith fine and deep punctures, which are separated by 2 - 4 times their diameter andbecome denser apically.
Pronotal sides (Fig. lOlA) evenly and rather strongly convergent forward.
Surface of pronotum weakly depressed behind eyes. Marginal stria weakly carinate andcomplete laterally and apically. Disk of pronotum evenly and moderately punctate
except near the apical angles and behind eyes, the impunctate areas being clothed withmicroscopic punctures only; the moderate-sized punctures becoming denser and fineron lateral 113 and becoming coarser along basal margin, with the microscopic puncturesintermingled. Antescutellar area with a shallow and longitudinal depression.
Epipleural marginal stria present on apical 1/2 and well impressed. Marginal
elytral stria deeply impressed and complete, its apical end extended along apical marginof elytron, attaining to sutural angle, then inwardly bent, and shortly extended
anteriorly. Epipleura impunctate, rarely clothed with microscopic punctures, and withcoarse rugae medially. Disk of elytra evenly and coarsely punctate except on a narrow
band along the suture, which is clothed with microscopic punctures and feebly elevated,and on extreme apical band; the coarse punctures somewhat longitudinal and separated
by 1 - 3 times the diameter, becoming coarser around scutellum and near obliquedepression, their interspaces with microscopic punctures intermingled. Dorsal stria
absent, but oblique and shallow depressions present on laterobasal 112.
Propygidium sparsely cov red with coarse, shallow and round punctures, whichbecome sparser and finer laterally d basally. Pygidium (Fig. 101C) evenly covered
with fine punctures, which are separate by twice their diameter; some oblong, shallow
and large punctures present near the bas I margin. In female, whole surface except for anarrow area along margin evenly coverttd with vermicular sculptures (Fig. 101C).
Anterior margin of prosternallobe (Fig. 101B) truncate anteriorly; marginal striaof lobe absent; basal angles strongly carinate and its insides deeply depressed; disk
irregularly scattered with moderate or coarse and deep punctures, which are separated
by 2 - 5 times their diameter. Prosternal process completely with carinal striae, which
are well impressed and feebly divergent apically; disk sparsely and finely punctate.
Lateral descending stria short and strongly carinate.
Anterior margin of mesosternum strongly emarginate at middle; marginal stria
absent anteriorly, but complete and well impressed laterally, its anterior end strongly
bent inwards and posterior end continued to lateral metasternal stria; disk with
punctation similar to that of prosternal process; transverse stria well impressed, straighton middle part, and obliquely and posteriorly arcuate on each side, usually not angulate(sometimes angulate). Meso-metasternal suture indistinct. Intercoxal disk ofmetasternum evenly covered with longitudinal and moderate punctures, which areseparated by about 1.5 (1 - 2) times the diameter, and become finer medially andcoarser laterally. Lateral disk covered with few large punctures around margin. Lateralmetasternal stria well impressed, extending obliquely and posteriorly and attaining tonear apical 113.
Intercoxal disk of 1st abdominal sternum evenly covered with oblong and coarse
punctures, which become finer medially and are separated by 2 - 4 times their diameter.
First abdominal stria complete and well impressed.
Protibia with 4 denticles on outer margin. Mesotibia with 3 spinules on outermargin. Metatibia with 2 spinules near the apex.
Male genitalia as shown in Fig. 102.
176

Table 22. Biometric data of Platylamalus fujisanus (Lewis).
Female

Male
APW
PPW
PL

EL
EW
ProW
ProL
PyL

PTL
MSTL

MITL

0.47-0.57 (0.53± 0.01) 13
0.95-1.12 (1.02± 0.02) 13
0.51-0.67 (0.59± 0.01) 13
0.89-1.10 (1.02± 0.02) 13
1.00-1.18 (1.l0± 0.02) 13
0.47-0.59 (0.53± 0.01) 13
0.12-0.20 (0.18± 0.01) 13
0.22-0.32 (0.26± 0.01) 13
0.35-0.43 (0.40± 0.01) 13
0.33-0.43 (0.38± 0.01) 13
0.45-0.53 (0.50±0.005) 13

0.47-0.55 (0.51± 0.01) 13
0.91-1.06 (0.99± 0.01) 13
0.55-0.65 (0.59± 0.01) 13
0.85-1.12 (1.00± 0.02) 13
0.99-1.14 (1.08± 0.01) 13
0.49-0.59 (0.55± 0.01) 13
0.16-0.55 (0.38± 0.05) 13
0.24-0.32 (0.27± 0.01) 13
0.35-0.45 (0.40± 0.01) 13
0.33-0.39 (0.37±0.005) 12
0.39-0.45 (0.42± 0.01) 13

Specimens examined. [Honshu] Aomori-ken: 1 ~, Towada, 16-19/vi/1968, H. Takizawa.
Nagano-ken: 1 ex., Togakushi, 9/viii/1963, Y. Kishi; 1 ex., Agematsu, Koma-ga-take, 12/vi/1949, S.
Osawa and T. Nakane. Gifu-ken: 2<1'<1' 3 ~ if , Kammuri-yama-t6ge, Tokuyama-mura, 1O/x/1986,
26/xi/1988, Y. Takai. Mie-ken: 1 ex., Okimi Pass, 12/vi/1952, H. Ishida. Wakayama-ken: Gomadan,
27/viii/1967, M. Yoshida. Hy6go-ken: 2 exs., Hataganaru, 24, 26/vi/1955, T. Nakane; 1<1',
Hy6nosen, 27/iv/1986, T. Ito. Tottori-ken: 1 ex., Daisen, 261iv/1942, K. Sakaguchi; 1 ex., Ditto,
6/vi/1955, K. Sawada; 1 ex., Ditto, Il1vii/1948, M. Iga; 3 exs., Ditto, H6ki, 800 m, 4/viii/I942, M.
Azuma. Sado Is.: 10", Tatara-mine, 28/vii/ 1947, M. Iga.
[Kyushu] Fukuoka-ken: 2<1' <1' 1 if , Mt. Hiko-san, 23/x/ 1985, M. Ohara. Kumamoto-ken: 1 ~ ,Shiraga-dake, 13/v/ 1984, M. Ohara. Miyazaki-ken: 1 ex., Mt. Sobo, 1O-12/v/1967, H. Takizawa.

Distribution. Japan (Honshu, Kyushu). New to KyushG.
Remarks. Platylomlaus fujisanus resembles P. viaticus superficially, but they
can readily be distinguished by several characteristics given in the key and description.
Little is known about the habitat of this species. Some specimens were collectedunder bark.
Pia tyloma Ius mendicus (Lewis, 1892)(Fig. 103, 104, 105)Paramalus mendicus Lewis, 1892a: 33 [Japan]; Lewis, 1899: 21 [Java]; Cooman, 1937: 138[Shanghai, Indochina].
Platylamalus mendicus: Cooman, 1948: 138; Mazur, 1984: 135; Hisamatsu, 1985b: 225 [photo].

Japanese name: Hime-chibi-hirata-emma-mushi.

Description. Male and female. Body length, PPL, 1.87 - 2.23 mm (2.05 ± 0.02,n=20), PEL, 1.73 - 2.03 mm (1.90 ± 0.02, n=20). Width, 1.14 - 1.34 mm (1.26 ±
0.01, n=20). Biometric data are given in Table 23. Body oblong, dark brown or blackand shining; femora, tibiae, mouthparts and antennae rufopiceous.
Frontal stria (Fig. 104C) of head well impressed and complete. Disk sparsely
covered with moderate punctures which are separated by 2 - 4 times their diameter andbecome a little denser apically; area between the punctures shining.
Pronotal sides (Fig. I03A, 104A) feebly arcuate and convergent forward on
basal 2/3, then strongly convergent apically. Emarginated portion of anterior margin of
pronotum feebly bisinuate. Marginal stria complete laterally, but broadly interrupted
177

::~:'. %8000;.··110

.:

,

3
3

A
Fig. 103 Platylomalus mendicus (Lewis). A: Pronotum and left elytron, dorsal view. B: Prosternum,
mesostemm, metasternum and 1st abdominal sternum. C: Pygidium of the specimens from
Fukushima pref., HonshU. D: Ditto, from Okinawa, Ryukyu Isles.

Fig. 104. Platylomalus mendicus (Lewis). A: Adult, dorsal view. B: Ditto, ventral VIew. C: Head,frontal view. D: Protibia, dorsal view. E: Prosternum, ventral view.

178

anteriorly behind neck. Disk sparsely and evenly clothed with fine punctures, withmicroscopic punctures intermingled and becoming denser on median 113 of pronotum.
A small puncture present on antescutellar area somewhat distant from margin.
Epipleural marginal stria weakly impressed on apical 112. Elytral marginal stria
deeply impressed and complete, its apical end extended to lateral 2/3 of posterior marginof elytron. Epipleura even and sparsely clothed with microscopic punctures. Disk of
elytra (Fig. 103A) sparsely and coarsely punctate except on a narrow band along
suture, which is sparsely clothed with microscopic punctures and feebly elevated; on
extreme apical band, the punctures becoming finer mediobasally, and coarser near
oblique depression. Dorsal stria absent, but oblique depression present on lateral 112 on
basal 113.
Propygidium sparsely covered with coarse, shallow and round punctures, which
are separated by 1 - 4 times the diameter. Pygidium (Fig. 103C, D) sparsely and finely
punctate, the punctures becoming coarser near basal margin. In female, a pair of curved
sculptures present on apical 1/2 as shown in Fig. l03D. These female sculptures arefound in the specimens from Okinawa, Ryilkyu Isles, but not in the specimens from
Fukushima-ken, Honshu.
Anterior margin of prostemal lobe (Fig. l03B) truncated; anterior angles of thelobe round; marginal stria present at lateral 113 on basal 112, extending obliquely andinwardly; lateral margin of lobe strongly carinate and its inside deeply depressed near
the posterior angles; disk irregularly scattered with fine and deep punctures, which are

A

B

O.5mm

D

~
::
-~
~.:
. . .C . . . . . .

'<

O.5mm

J

G

if.l
"if

=j

~

..
........ ~H

Fig. 105. Platylomalus mendicus (Lewis). A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and 10th
tergites, dorsal view. F: Ditto, lateral view. G: Male genitalia, 8th sternum, dorsal view. H:
Ditto, lateral view. I: Ditto, caudal view.
179

Table 23. Biometric data of Platylomalus mendicus (Lewis).
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

0.53-0.63
1.04-1.26
0.59-0.71
1.02-1.30
1.14-1.34
0.57-0.69
0.20-0.28
0.20-0.30
0.41-0.47
0.30-0.39
0.37-0.49(0.60± 0.01) 20(1.16± 0.01) 20(0.66± 0.01) 20(1.13± 0.01) 20(1.26± 0.01) 20(0.63± 0.01) 20(0.23±0.004) 20(0.24± 0.01)20(0.44±0.004) 20(0.36± 0.01) 20(0.42± 0.01) 20

separated by 1 - 4 times the diameter. Prosternal process completely with carinal striae,
which are divergent anteriorly and posteriorly, narrowest at basal 2/3, and not meetingat their ends. Lateral descending stria short, complete and strongly carinate.
Anterior margin of mesosternum (Fig. 1mB) feebly emarginate at middle; its
marginal stria absent anteriorly, complete laterally; basal end of the marginal stria bent
inwards and apical end continued to lateral metasternal stria; punctation of mesosterna!
disk similar to that of prosternal process; transverse stria weakly impressed, usuallyinterrupted at middle (in the original description, the stria is absent). Meso-metasternal
suture finely depressed. Intercoxal disk of metasternum sparsely clothed with finepunctures which are separated by 3 - 8 times their diameter. Lateral disk sparsely
covered with large, round and shallow punctures, which become finer apically. Lateralmetasternal stria well impressed and extended obliquely and posteriorly, its apical end
attaining to near metacoxal cavity. Post-mesocoxa! stria extended along mesocoxal
cavity and its outer end strongly bent posteriorly.
Intercoxal disk of 1st abdominal sternum feebly depressed on lateral sides, andwith punctation similar to metasternal ones. First abdominal stria complete and wellimpressed.
Protibia with 4 dentic1es on outer margin. Metatibia with 4 spinules on outermargin. Metatibia with 2 spinules near the apex.
Male genitalia as shown in Fig. 105.
Specimens examined. [Honshu] Fukushima-ken: 13 exs., Kita-Aizu, Kuwabara, 15/iv/1950, T.
Nakane (NA); Id", 12 exs., Kami-miyori, Kita-Aizu, 16/iv/1950, K. Nagayama (NA); 3 exs.,
Kawatani, Higashi-yama, Kita-Aizu, 24/x/1948, Y. Kurosawa (NA).
[Nansei Isles] Okinawa-Honto Is. : 11 exs., Sueyoshi-koen, Naha, 311v/1988, T. Veno.
Ishigaki Is.: 1 ex., Kabira, 25/v/1988, H. Daido.

Distribution. Japan (Honshu, Shikoku, Kyushu, Oki Is., Nansei Isles); Russia(Primorskiy Kray); China; Viet-Nam; India; Taiwan; Indonesia.
Remarks. Platylomalus mendicus resemlbes P. persimilis; however, thedifferent shape of the mesosternal transverse stria and the sculptures of the female
pygidium will distinguish it from P. persimilis.
Platylomalus niponensis (Lewis, 1899)(Fig. 106, 107, 108)
Paromalus complanatus: Lewis, 1892: 33, nec Panzer, 1797, corrected by Lewis, 1899: 21.
Paromalus niponensis Lewis, 1899: 21 [Japan]; Cooman, 1937: 142.

180

Platylomalus niponensis: Kryzhanovskij and Reichardt, 1976: 274; Mazur, 1984: 225; Hisamatsu,
1985b: 225 [photo].
Japanese name: O-chibi-hirata-emma-mushi.

Description. Male, Female. Body oblong, black and shining; tarsi and antennae
rufopiceous. Body length, PPL, male, 2.52 - 3.17 mm (2.98 ± 0.04, n=17), female,
2.65 - 3.01 mm (2.82 ± 0.04, n=9), PEL, male, 2.19 - 2.91 mm (2.68 ±0.04, n=17), female, 2.45 - 2.68 mm (2.54 ± 0.03, n=8). Width, male, 1.47 - 1.90
mm (1.76 ± 0.02, n=19), female 1.60 - 1.77 mm (1.68 ± 0.02, n=9). Biometric data
are given in Table 24.
Head (Fig. 107C) evenly covered with moderate punctures which are separated
by twice their diameter. Frontal stria well impressed and complete.
Sides of pronotum (Fig. 106A) arcuate, slightly convergent anteriorly on basal
3/4, thence strongly arcuate and convergent apically. Marginal stria nearly complete, but
a little broken at middle. Disk of pronotum sparsely clothed with fine punctures which
are separated by 4 - 6 times their diameter and become coarser on lateral 114.
Antescutellar area distinctly with a short longitudinal impression.
Marginal epipleural stria feebly impressed on apical 112. Marginal elytral stria
strongly carinate and complete, its apical end extending along the apical margin of
elytron, and attaining to near lateral 2/3. Epipleura shining and microscopicallypunctate. Oblique humeral stria present on basal 114. First dorsal stria present on
median 1/3, weakly impressed. Another oblique stria deeply impressed on laterobasal
112. Disk of elytra sparsely covered with coarse and shallow punctures which areseparated by 1 - 4 times the diameter and become finer on mediobasal 112 and along thesuture.

!
!

Fig. 106. Platylomalus niponensis (Lewis). A: Pronotum and left eIytron, dorsal vIew. B:
Prosternum, mesostemm, metasternum and 1st abdominal sternum. C: Pygidium.

181

Propygidium sparsely covered with moderate punctures, with fine puncturesintermingled. The male and female are confused in the original description; in reality,
the pygidium (Fig. 107C) is densely and finely punctate in the male; in female, it has ananastomosed sculpture medially (Fig. 107D).
Anterior margin of prostemal lobe (Fig. 106B) truncate and rather broad;marginal stria of the lobe absent anteriorly, but distinctly present on lateral 113; anothershort marginal stria on each lateral side and strongly carinate at basal angles of the lobe;
disk sparsely and finely punctate, the punctures being separated by 2 - 5 times theirdiameter. Prostemal process with complete carinal striae; disk of the process sparsely
and finely punctate; descending lateral stria short, strongly carinate.

Fig. 107. Platylomalus niponensis (Lewis). A: Adult, dorsal view. B: Ditto, ventral view. C: Head,frontal view. D: Propygidium and pygidium. E: Pro-, meso- and metasterna, ventral view. F:
Protibia, dorsal view.

182

A

(

f

2~··

E

O.5mm

Fig. 108. Platylomalus niponensis (Lewis). A: Aedeagus, lateral view. B: Ditto, dorsal view. C:
Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite, dorsal view. E: Ditto, lateralview. F: Male genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. G: Ditto,
lateral view. H: Male genitalia, 8th sternum, lateral view. I: Ditto, lateral view. J: Ditto, caudal
view.

Table 24. Biometric data of Platyiomalus niponensis (Lewis).
Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Female0.69-0.88 (0.83±0.01) 19
1.34-1.77 (1.66±0.02) 19
0.78-1.01 (0.93±0.01) 19
1.37-1.73 (1.60±0.02) 19
1.47-1.90 (1.76±0.02) 19
0.72-1.01 (0.88±0.01) 19
0.23-0.39 (0.32±0.01) 19
0.33-0.49 (0.40±0.01) 19
0.52-0.72 (0.65±0.01) 19
0.49-0.69 (0.59±0.01) 19
0.59-0.78 (0.69±0.01) 19

0.75-0.82 (0.79±0.01) 9
1.50-1.64 (1.57±0.01) 9
0.78-0.92 (0.86±0.02) 8
1.44-1.60 (1.54±0.02) 8
1.60-1.77 (1.68±0.02) 9
0.82-1.01 (0.90±0.02) 8
0.29-0.33 (0.31±0.01) 8
0.29-0.43 (0.36±0.02) 8
0.56-0.62 (0.59±0.01) 9
0.46-0.62 (0.54±0.01) 9
0.56-0.69 (0.62±0.02) 9

183

Anterior margin of mesosternum deeply emarginate at middle, its marginal stria
complete and carinate laterally, absent anteriorly; basal angles rectangular; disk evenly
and sparsely clothed with fine punctures, which are separated by about 3 times theirdiameter, and indistinctly with a biarcuate transverse line (Fig. 106B). Mesometasternal suture slightly impressed and broadly sinuous. Intercoxal disk ofmetasternum with punctures similar to those of metasternum. Lateral metasternal stria
strongly carinate, extending obliquely and posteriorly on basal 2/3, thence runningalong the metasternal-metepisternal suture, and attaining near the latero-posterior angle.
Intercoxal disk of 1st abdominal sternum sparsely and finely punctate, andcompletely striate on lateral side, the stria strongly carinate.
Protibia (Fig. l07F) with 6 denticles on outer margin. Mesotibia with 4 spinules
on outer margin. Metatibia with 2 spinules near the apex.
Male genitalia as shown in Fig. 108.
Specimens examined. [Hokkaido] 20"'0'" 3!f !f , Maruyama, Sapporo, 25/v, vi/1989, S.
Shiyake; 40"'0'" 2!f !f , Ditto, 10, Il1v/1989, M. Ohara; 1 d", 1 ex., Ditto, 5/v/1973, S. Kudo; 2d"d" ,Mt. Yotei, Hirafu, Kutchan, 15-16, 22/vii/1991, M. Ohara.
[Honshu] Fukushima-ken: 1 ex., Naka-Moiwa, Azuma-yama, 13/iv/1966, K. Tazoe; 3 exs.,
Kami-miyori, Kita-Aizu, 16/iv/1950, K. Nagayama (NA); 5 exs., Kuwabara, Kita-Aizu, 15/iv/1950,
Y. Kurosawa (NA); 40"'0'" 4!f !f , Ditto, (NSMT). TOkyo-to: 2 exs., Takao, 14/vi11930, S. Asahina.
Nagano-ken: lei", Sanosaka, Hakuba ViiI., 8/v11983, M. Kiuchi.

Distribution. Japan (Hokkaid6, Honshu); Taiwan.
Remarks. Platylomalus niponicus is a relatively large species; it is similar to P.
persimilis and P. mendicus, but can be distinguished by the different shape of themesosternal transverse stria and the larger size.
Little is known about the habitat of this species. Some specimens were collectedunder bark.
Platylomalus persimilis (Lewis, 1888)(Fig. 109, 110)
Paromnlus persimilis Lewis, 1888: 641 [TenasseriumJ; Cooman, 1937, 133 [Tonkin].
Platylomalus persimilis: Mazur, 1984: 136; Hisamatsu, 1985b: 224 [photo].

Japanese name: Kuro-chibi-hirata-emma-mushi.

Table 25. Biometric data of Playlomalus persimilis (Lewis).
Male
APW
PPW
PL

EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Female\0.63-0.69 (0.66± 0.01) 9
1.08-1.28 (1.20± 0.02) 9
0.64-0.76 (0.71± 0.02) 9
1.08-1.27 (1.20± 0.02) 9
1.15-1.39 (1.28± 0.02) 9
0.59-0.78 (0.72± 0.02) 9
0.20-0.29 (0.25± 0.01) 9
0.34-0.41 (0.36± 0.01) 9
0 .. 44-0.52 (0.48±0.01) 9
0.37-0.49 (0.43± 0.01) 9
0.47-0.59 (0.52± 0.01) 9

0.54-0.68 (0.60± 0.02) 6
0.98-1.27 (1.10± 0.04) 6
0.56-0.73 (0.65± 0.02) 6
0.98-1.27 (1.09± 0.04) 6
1.03-1.37 (1.19± 0.05) 6
0.61-0.71 (O.66± 0.02) 6
0.15-0.27 (0.22± 0.01) 6
0.24-0.29 (0.26± 0.01) 6
0.37-0.49 (0.43± 0.02) 6
0.34-0.44 (0.39± 0.02) 6
0.42-0.51 (0.46± 0.02) 6

184

Description. Body oblong, black and shining; femora, tarsi and antennae clear
red brown. Body length, PPL, male, 2.03 - 2.50 mm (2.31 ± 0.06, n=9), female,
1.76 - 2.35 mm (2.05 ± 0.08, n=6); PEL, male, 1.79 - 2.25 mm (2.04 ± 0.05, n=9),female, 1.64 - 2.11 mm (1.87 ± 0.08, n=6). Width, male, 1.15 -1.39 mm (1.28± 0.02, n=9), female, 1.03 - 1.37 mm (1.19 ± 0.05, n=6). Biometric data are
given in Table 25.
Frontal stria of head clearly impressed and complete; disk of head evenly andfinely punctate, the punctures being separated by about twice their diameter.
Pronotal sides (Fig. 109A) slightly arcuate and convergent forward. Apicalangles acute. Emarginated portion of apical margin bisinuated behind neck. Marginalstria complete laterally and rather broadly interrupted medially on anterior margin. Diskof pronotum evenly clothed with fine and deep punctures, which are separated by 3 - 5times their diameter.
Epipleural marginal stria present on apical 112. Elytral marginal stria deeply
impressed, densely crenate and complete, its apical end extending along the apicalmargin of elytron and attaining to lateral 112. Epipleura impunctate and shining. Disk of
elytra sparsely covered with coarse and deep punctures; the punctures 3 times as coarse
as those of pronotum, separated by 2 - 5 times their diameter and becoming coarser
laterobasally, and becoming finer narrowly along the suture and on extreme apicalband.
Propygidium sparsely and coarsely punctate, the punctures becoming finer anddenser basally. Pygidium (Fig. 109C) convex, and sparsely clothed with fine punctures
which are separated by about 3 times their diameter in the male. In female, pygidiumwith a large ocelloid sculpture, which is very deep, usually with several short foveae
around.
Anterior margin of prostemal lobe (Fig. 109B) truncate, its median portion
broadly straight. Marginal stria of lobe absent anteriorly and present on basal 112 oflateral 1/3; another short marginal stria present on each posterior angle and strongly
carinate. Disk of the lobe sparsely covered with fine punctures which are separated by 2
- 4 times their diameter. Prostemal process completely with carinal striae, the striae

~2
~
.. :

.......;.;... \ ...... .

C

Fig. 109. Pia tyloma Ius persimilis (Lewis). A: Pronotum and left e1ytron, dorsal view. B: Prosternum,
mesostemm, metasternum and 1st abdominal sternum. C: Pygidium.

185

A

c~~~) 7--"~~?~=~~~==~~:::~:::-2-~=~~~
B

D

O.5mm lF

Fig. 110. Pia tyloma Ius persimilis (Lewis) A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and 10th
tergites and 9th sternum (spicule), dorsal view. F: Ditto, lateral view. G: Male genitalia, 8thsternum, lateral view. H: Ditto, caudal view.

being well impressed; disk sparsely and finely punctate. Descending lateral stria shortand strongly carinate.
Anterior margin of mesosternum strongly emarginate medially, its marginal stria
absent anteriorly and complete and well impressed laterally, the basal end of the stria
rectangular inwards, and the apical end continued to the metasternal lateral stria; disk of
mesosternum densely clothed with fine punctures except for an inside area surrounded
by a transverse stria and a meso-metasternal suture; transverse stria (Fig. 109B) rather
strongly carinate. Meso-metasternal suture slightly impressed and complete. Intercoxal
disk of metasternum sparsely clothed with fine punctures which are separated by 4 - 6times their diameter and become sparser laterally. Lateral disk of metasternum sparsely
covered with coarse punctures and with postcoxal stria which runs along the mesocoxal
cavity and strongly sinuates near the lateral end. Lateral metasternal stria deeply
impressed, extending posteriorly and obliquely on basal 2/3, thence running along
suture between metasternum and metepistenum, and its apical end attaining to nearmetacoxa and curved inwardly.
Intercoxal disk of 1st abdominal sternum sparsely and finely punctate and 1st
abdominal stria present on each lateral side.
Protibia with 5 denticles on outer margin. Mesotibia with 4 spinules on outermargin. Metatibia with 1 spinule near apex.
Male genitalia as shown in Fig. 110.
Specimens examined. (Nansei Isles] Tokara-Nakano-shima Is.: 4ci'ci' 2~ ~ , Nakano-shima Is.,
Tokara Isles, 18/vii/1982, M. Ohara. Yoron Is.: 1 ~ , Yoron Is., 5/ix/1988, Y. Kusui. Okinawa-Hont6
186

Is.: 10", Yona, 13-15/vi/1970, H. Makihara. Kerama Is.: 10" 1!f., Aka, Kerama, 2/viii/1987, Y.
Kusui. Ishigaki Is.: 10", Mt. Banna-dake, 20/ii/1982, K. Sawada; 1 ex., Ditto, 311x/1974, K. Tazoe.

Distribution. Japan (Nansei Isles); Tenasserim; Viet-Nam.
Remarks. Platylomalus persimilis is very similar to P. mendicus, but differs by
the black body, the trapezoid transverse stria of the mesosternum and the mesotibia
bearing 1 spine.

Platylomalus viaticus Lewis, 1892
(Fig. 111, 112, 113)
Paromalus viaticus Lewis, 1892a: 33 [Japan: Nikko, Oyama, Kashiwagi, Nara, Kumamoto, andYuyama, chiefly in the warmer districts of the islands].
Platylomalus viaticus: Kryzhanovskij and Reichardt, 1976: 275; Mazur, 1984: 136; Hisamatsu,
1985b: 224 [photo].
Japanese name: Tsuya-chibi-hirata-emma-mushi.

Description. Male and female. Body oblong, black and shining; femora, tarsi,
mouthparts and antennae rufopiceous. Body length, PPL, male, 1.64 - 1.91 mm (1.79± 0.03, n=9), female, 1.49 - 1.77 mm (1.63 ± 0.06, n=5); PEL, male, 1.52 - 1.72
mm (1.64 ± 0.02, n=9), female, 1.45 - 1.64 mm (1.54 ± 0.03, n=4). Width, male,
0.95 - 1.13 mm (1.05 ± 0.02, n=9), female, 0.95 - 1.08 mm (1.02 ± 0.03, n=5).
Biometric data are given in Table 26.
Head (Fig. 112D) densely covered with coarse punctures which are separated by
0.5 - 1 times the diameter, with microscopic punctures intermingled. Frontal stria
complete and feebly carinate.
Pronotal sides (Fig. lIlA) slightly arcuate and convergent apically. Emarginatedportion of apical margin bisinuate behind neck. Marginal stria complete laterally andanteriorly. Disk of pronotum densely covered with coarse and somewhat oblong
punctures which are somewhat coarser than those of the head. Antescutellar area
without a large puncture.

..

~

: ..

3
3

.:::.\ ~ ~:.;...:
A
Fig. 111. Platylomalus viaticus (Lewis). A: Pronotum and left eiytron, dorsal view. B: Prosternum,
mesostemm, metasternum and 1st abdominal sternum. C: Pygidium.
187

Fig. 112. Platylomalus viaticus (Lewis). A: Adult, dorsal view. B: Ditto, ventral view. C:
Prostemurn, ventral view. D: Head, frontal view. E: Protibia, dorsal view. F: Ditto, ventral
VIew.

Epipleural stria weakly impressed on apical 112. Marginal elytral stria deeply
impressed and complete, coarsely punctate inside; the apical end of the stria extending
along apical margin of elytron, through apical sutural angle, and shortly extending
anteriorly. All dorsal striae indistinct, but feeble oblique depressions present onlaterobasal 112 of elytron. Punctation of elytra (Fig. lIlA) similar to pronotal one, butmuch coarser and sparser except on a narrow band along suture and on extreme apex ofelytra, the punctures becoming finer mediobasally and coarser around the obliquedepression.
Propygidium densely and moderately punctate, the punctures becoming sparser
and finer laterally. Pygidium evenly clothed with fine punctures which are separated by
3 - 5 times their diameter and become a little denser on apical 1/3. In female, thepunctation of pygidium becoming much denser on apical 1/3 and sometimes with atransverse depression near apex.
Anterior margin of prostemal lobe (Fig. I11B, 112C) truncate; marginal stria
188

shortly impressed only near posterior angle and carinate; disk sparsely covered withmoderate punctures, which are separated by 2 - 5 times their diameter. Prosternal
process with complete carinal striae; disk scattered with moderate punctures which areseparated by 2 - 10 times their diameter. Descending lateral stria short, complete andstrongly carinate.
Anterior margin of mesosternum (Fig. 111B) deeply emarginate at middle, its
marginal stria absent anteriorly, but complete and well impressed laterally; apical end of
the marginal stria continued to basal end of lateral metasternal stria; disk sparsely
clothed with fine punctures which are separated by 3 times their diameter; transverse
stria absent. Intercoxal disk of metasternum similar to punctation of mesosternum, thepunctures becoming a little coarser laterally; lateral stria well impressed, extending
obliquely and posteriorly, running along mesosternal-metepisternal suture on apical 112,
its apical end attaining near metacoxal cavity; lateral disk of metasternum sparsely andcoarsely punctate. Postcoxal stria extending along mesocoxal cavity, its outer end
strongly bent posteriorly.
Intercoxal disk of 1st abdominal sternum sparsely and moderately punctate; 1stabdominal stria straight, well impressed and complete.
Protibia (Fig. 112E, F) with 4 denticles on outer margin. Mesotibia with 3
spinules on outer margin. Metatibia with a spinule near apex.
Male genitalia: as shown in Fig. 113.

A

O.25mmo~...... fb\
L:i\~............... ' .. :. H
O.25rrun I

Fig. 113. Platylomalus viaticus (Lewis). A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and 10th
tergites, lateral view. F: Ditto, dorsal view. G: Ma1e genitalia, 8th sternum, dorsal view. H:
Ditto, lateral view. I: Ditto, caudal view.

189

Table 26. Biometric data of Platylamalus viaticus (Lewis).
Female

Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

0.46-0.54 (0.50± 0.02) 5
0.81-1.00 (0.90± 0.03) 5
0.47-0.56 (0.53± 0.01) 5
0.86-0.98 (0.91± 0.02) 5
0.95-1.08 (1.02± 0.03) 5
0.42-0.59 (0.50± 0.03) 5
0.14-0.15 (0. 14±0.004) 5
0.17-0.25 (0.21± 0.01) 5
0.34-0.37 (0.36± 0.01) 5
0.29-0.34 (0.31± 0.01) 5
0.35-0.42 (0.40± 0.01) 5

0.47-0.57 (0.53± 0.01) 9
0.90-1.01 (0.97± 0.02) 9
0.51-0.61 (0.56± 0.01) 9
0.91-1.08 (0.99± 0.02) 9
0.95-1.13 (1.05± 0.02) 9
0.46-0.56 (0.50± 0.01) 9
0.10-0.15 (0. 13±0.003) 9
0.17-0.27 (0.23± 0.01) 9
0.35-0.44 (0.38± 0.01) 9
0.29-0.42 (0.34± 0.01) 9
0.37-0.42 (0.40± 0.01) 8

Specimens examined. [Honshu] Aomori-ken: 1 ex., Mt. Osore, Shimokita, 31/vii/1956, T.
Nakane (NA). Fukushima-ken: 1<1", Araumi, Minami-Aizu, 9/v/1948, K. Nagayama (NA); 1 ex.,
Ashinohara, Egawa, Minami-Aizu, 18/v/1948, Y. Kurosawa (NA); 1 ex., Yunohana, Tateiwa,
Minami-Aizu, 9/viii/1948, Y. Kurosawa (NA). 1 Jf, nr. Oyama, nr. Wakamatsu, Aizu, 14/v/1947,
Y.Kurosawa (NA); 1 <1", Suimon, Omaki, Asahida V., Minami-Aizu, 20/v/1948, Y. Kurosawa (NA).
Saitama-ken: 1 ex., Yanagi-Koya, Chichibu, 22/viil 1933, S. Asahina (NA). Nara-ken: 3 exs., Kasuga,
Nara, 19/vi/1957, 13/vi/1958, T. Nakane (NA); 1 ex., Ditto, 30/iv/1950, N. Yato (NA); 1<1", Nara
Park, Nara-shi, 7/x/1987, K. Sawada. Hyogo-ken: 1<1", Hataganaru, 24/v/1955, T. Nakane (NA).
[Izu Isles] Hachijo-jima Is.: 1 ex., Suigenchi, Hachijo-jima, 26/vi/1958, K. Mizusawa (NA).
[Kytlshu] Fukuoka-ken: 1<1" 1 Jf ,Mt. Hiko-san, 23/x/1985, M. Ohara.
[faiwan] 2 exs., Karapin, near Mt. Ari [now Ali Shan], lIv/1941, Y. Yano (NA).

Distribution. Japan (Honshu, Kyushu); Taiwan; Russia (Chabarovskiy Kray).
New to Kyushu.
Remarks. Platylonwlus viaticus can easily be distinguished from P. fujisanusby the absence of a transverse stria on the mesosternum and the absence of furrows on
the female pygidium. Lewis (1892) described sexual dimorphism in the state of thepygidium, but he confused the sexes.
Platylomalus kusuii M. Ohara, sp. nov.
(Fig. 114, 115)
Japanese name: Ogasawara-chibi-hirata-emma-mushi.

Description. Body oblong, black and shining; femora, tarsi, mouthparts andantennae red brown. Body length, PPL, female, 2.09 - 2.60 mm (2.34 ± 0.08, n=6);
PEL, male, 1.99 mm, female, 1.89 - 2.36 mm (2.11 ± 0.06, n=7). Width, male, 1.26mm, female, 1.20 - 1.50 mm (1.35 ± 0.04, n=7). Biometric data are given in Table 27.
Frontal stria of head clearly impressed and complete; disk of head sparsely andfinely punctate, the punctures being separated by 2 - 5 times their diameter.
Pronotal sides (Fig. 114A) slightly arcuate and convergent to apical 114, thence
strongly convergent apically. Emarginated portion of apical margin bisinuate behindneck. Marginal pronotal stria complete, but shortly interrupted at middle on anterior
margin. Disk of pronotum densely clothed with microscopic punctures, which become
sparser and coarser on lateral 113 and are separated by 2 - 3 times their diameter.
Antiscutellum area with a large puncture.
190

..................

~

3

3

~
···········.·.·······/>0..... .

!

...
~
/7~.....
.
,..'\:!f5Z/9) ............••. \
c···
:~.;:

..

:::.

B

0(.' ...
.. ,

I

~

Fig. 114. Platylomalus kusuii sp. nov. A: Pronotum and left elytron, dorsal view. B: Prosternum,
mesostemm, metasternum and 1st abdominal sternum. C: Pygidium.

Epipleural marginal stria present on apical 2/3. Elytral marginal stria deeply
impressed, coarsely crenate and complete, its apical end extending along the apicalmargin of elytron and attaining to lateral 3/4. Epipleura shining and sparsely clothedwith fine punctures. Disk of elytra (Fig. 114A) sparsely covered with moderatepunctures; the punctures separated by 2 or 3 times their diameter, becoming denser andfiner on apical 112 narrowly along the suture, and becoming coarser broadly on apical
113 of elytron and narrowly around an oblique depression which is present on thelaterobasal 1/2 of the elytron; the punctation around the oblique depression very large,
round and deep; extreme apical band impunctate. Dorsal striae of elytron absent, but arudimentary impression (2nd dorsal stria ?) present on apical 2/3 of lateral 113 andanother one (3rd 7) on basal 112 of lateral 112.
Propygidium evenly and coarsely punctate, the punctures being separated byabout 1.5 (1 - 2) times the diameter. Pygidium (Fig. 114C) densely clothed with finepunctures which are separated by about 1.5 (1 - 2) times the diameter. In female,pygidium with punctation similar to that of the male and also with an irregular and liner
sculpture (Fig. 114C).
Anterior margin of prosternallobe (Fig. 114B) truncate; marginal stria of the lobe
absent anteriorly and present laterally on median 113; disk evenly covered with deep andmoderate punctures which are separated by about 1.5 (1 - 2) times the diameter.
Prosternal process completely with carinal striae, which are well impressed and feebly
sinuate at middle, sometimes reduced basally; disk of the process sparsely clothed withfine punctures which are separated by 2 - 4 times their diameter. Descending lateral
stria short and strongly carinate; outside area of the stria deeply depressed.
Anterior margin of mesosternum (Fig. 114B) feebly emarginate medially, its
marginal stria absent anteriorly, and complete and well impressed laterally; the basal endof the stria hooked inward; the apical end continued to metasternal stria; disk of themetasternum sparsely with fine punctures which are separated by 3 - 5 times theirdiameter, and with microscopic punctures sparsely intermingled. Meso-metasternal191

suture weakly impressed and feebly arcuate. Punctation of intercoxal disk ofmetasternum a little denser and finer than mesosternal ones, and the fine punctures
being usually represented in pairs; lateral disk sparsely covered with large, round andshallow punctures, which become smaller and sparser; lateral metasternal stria deeply
impressed, extending obliquely and outwardly, and reaching near the posterior coxal
cavity; postmesocoxal stria well impressed, extending along basal margin and its lateralend strongly bent posteriorly.
Punctation of 1st abdominal sternum similar to that of metasternum, thepunctures becoming coarser before apical angles; 1st abdominal stria complete on eachlateral side.
Protibia with 4 denticles on outer margin. Metatibia with 5 spinules on outermargin, but the basal most very small. Metatibia with 2 spinules near apex.

----- -

A

/
/

/
/

/

/
/

K
~s::sz::,>=::,.:::::::::::::::::::~._o,:."'~.. .... :~"":"":"

0.25 mm , G-K

0.25 mm!

A-F,L

Fig. 115. Platylomalus kusuii sp. nov. A - K: Male genitalia. A: Aedeagus, lateral view. B: Ditto,
dorsal view. C: Apex of aedeagus, caudal view. D: 8th sternum, dorsal view. E: Ditto, lateralview. F: Ditto, caudal view. G: 8th tergite, dorsal view. H: Ditto, lateral view. 1: 9th and 10th
tergites, dorsal view. J: Ditto, lateral view. K: 9th sternum (spicule), dorsal view. L: Female
genitalia, spermatheca.

192

Table 27. Biometric data of Platylomalus kusuii M. Ohara, sp. nov.
Male (n=l)
APW
PPW
PL
EL
EW
ProW
ProL
PyL

PTL
MSTL

MTIL

0.61
1.24
0.75
1.14
1.26
0.63
0.26
0.32
0.47
0.43
0.47

Female0.59-0.71 (0.66± 0.02) 7
1.16-1.42 (1.28± 0.04) 7
0.63-0.89 (0.76± 0.04) 7
1.10-1.36 (1.23± 0.04) 7
1.20-1.50 (1.35± 0.04) 7
0.61-0.75 (0.68± 0.02) 6
0.22-0.30 (0.25± 0.01) 6
0.22-0.32 (0.28± 0.01) 6
0.43-0.53 (0.49± 0.01) 7
0.33-0.47 (0.43± 0.02) 7
0.45-0.57 (0.52± 0.02) 7

Male genitalia: as shown in Fig. 115A - K.
Female genitalia: as shown in Fig. 115L
Specimens examined. Holotype, 0', Sakai-ura, Chichi-jima Is., Ogasawara Isles, 8/vi/1972, Y.
Kusui (ElHU).
Paratypes. [Ogasawara Isles] 1 ~, Nishi-ura, Haha-jima Is., 15/vi/1972, Y. Kusui; 1ei"3 ~ ~ ,and 4 exs., Nakano-taira, Haha-jima Is., 29/vi/1976, Y. Kurosawa (NSMT); 1 ex., Bonin Is., no date,
M. Nishihara; 1 ex., Bonin Is., vil1969, T. Nakane (NA); I ~ , Wai beach, Haha-jima Is., 22/v/1989,
S. Miyano (Collection of Natural History Museum and Institute, Chiba)

Distribution. Japan (Ogasawara Isles).
Remarks. Platylomalus kusuii can easily be distinguished by the characteristic
large punctation on the humeral area of the elytra and the absence of a transverse striaon the mesosternum, in combination with its limited distribution (Ogasawara Isles).
GENUS PACHYLOMALUS SCHMIDT,1879

Pachylomalus Schmidt, 1897: 295 [type species: Paromalus leo Marseul, 1879: 272, designated by
Bickhardt, 1917: 115]; Bickhardt, 1917: 115.

SUBGENUS CANIDIUS COOMAN,1941
Canidius Cooman, 1941: 292 [type species: Pachylomalus opulentus Cooman, 1932: 65, originallydesignated].

Pachylomalus (Canidius) musculus (Marseul, 1873)(Fig. 116,117,118)
Paromalus musculus Marseul, 1873: 221, 225 [Niphon, Nangasaki]; Lewis, 1892a: 37 [Nara;
Kiushu]; 1915: 56 [Formosa].
Pachylomalus musculus: Bickhardt, 1917: 116; Hisamatsu, 1985b: 224, pi. 40, no. 25.
Pachylomalus (Canidius) musculus: Cooman, 1941: 298, fig. 6; Hisamatsu, 1968: 31 [Shikoku,
RyUkyfi].

Pachylomalus inopsCooman, 1933: 195, synonymized by Cooman, 1935: 217.
Japanese name: Hasuji-chibi-hirata-e=a-mushi.

193

.:<. :.:.: : \.:..-./.\. . :.~.::>.:::: .

.... .

.:'::::.':.:~>:~~: :.;} .•.... '\:i~;·:::~;·.....-).~____,10'I. :::" :..:.::::..,:~ >:: (':::.:/>:?')::/.):
A . ::.. :.. :.::.;':":;':.':.

.~ )
' . '

.......... :.

"."D·····/

rfB'-----~>;,~.i'i.::ii': (~,/ I~G.'.: ......

J

y

.....

Fig. 116. Pachylomalus (Canidius) musculus (Marseul). A: Pronotum and left elytron, dorsal view.
B: Prosternum, mesosternum, metasternum and 1st abdominal sternum. C: Propygidium andpygidium, male. D: Pygidium, female, caudal (left) and lateral views.

Description. Body length, PPL, 2.08 - 2.57 mm (2.33 ± 0.04, n=20), PEL,
1.79 - 2.38 mm (2.13 ± 0.04, n=20). Width, 1.27 - 1.72 mm (1.52 ± 0.03, n=20).
Biological data are given in Table 28. Body oblong-oval, black or dark brown andshining; tibiae, mouthparts and antennae rufopiceous.
Frontal stria of head weakly carinate and complete; disk sparsely clothed withfine punctures which are separated by 3 - 4 times their diameter; interspace among thepunctures shining.
Pronotal sides (Fig. 116A) arcuate and strongly convergent apically; anterior
angles acute. Emarginated portion of anterior margin bisinuate. Marginal stria complete,
weakly impressed. Disk of pronotum convex, sparsely clothed with fine punctures
similar to the pronotal ones; a pair of divergent striae present on mediobasal 113, area
between the striae feebly depressed. Basal margin of pronotum sparsely covered withlarge punctures, but absent medially.
Epipleural marginal stria weakly impressed and interrupted on mediobasal 114,
sometimes complete. Epipleura shining, sparsely clothed with microscopic punctures.
Elytral marginal stria deeply impressed and complete, its apical end extending along
apical margin of elytron, attaining to sutural angle, then bent inward, shortly extending
anteriorly. Disk of elytra evenly covered with coarse punctures which are separated by
3 - 4 times their diameter. Extreme apical band impunctate. Oblique humeral stria
slightly impressed on basal 1/3.
Propygidium (Fig. 116C) distinctly with a transverse stria which is wellimpressed; the lateral end of the stria not attaining to lateral margin; disk sparsely andfinely punctate, the punctures being separated by 3 - 4 times their diameter. Pygidium(Fig. 116C, D) with punctation similar to that of propygidium. In female, punctation of
pygidium as in male; surface with a large oval depression on apical 112.
Anterior margin of prostemal lobe (Fig. 117B) truncate, its marginal stria wellimpressed and strongly and inwardly bent laterally; basal angle shortly carinate, of
194which the inside area is strongly and obliquely excavated; disk of lobe covered withalutaceous ground sculpture and microscopic punctures which are separated by nearly 3
times their diameter. Prosternal keel narrow, its top even; disk impunctate, shining;carinal striae well impressed, divergent basally, and its apical ends united with eachother in an arc. Descending lateral stria deeply impressed.
Anterior margin of mesosternum strongly and deeply emarginate medially,
marginal stria absent; basal angles depressed; disk of mesosternum shining and sparsely~,-;;~:;~:~--~f:"--~··~
O.25mmr/;:;>""';~"~:.; . ;.'/~~~;~.<-~~-,"';.:"

o
"

~
,
J

'
tI2/
!

... / (,/ :!
. . / . . \ / /' H

O.5mrn,Fig. 117, Pachylomalus (Canidius) musculus (Marseul). A: Aedeagus, lateral view. B: Ditto, dorsal
view. C: Male genitalia, 8th tergite, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th
and 10th tergites, lateral view. F: Ditto, dorsal view. G: Male genitalia, 8th sternum, dorsal
view. H: Ditto, lateral view. I: Ditto, caudal view. J: Male genitalia, 9th sternum (spicule),
dorsal view.

Fig. 118. Collection sites of Pachylomalus (Canidius) mususculus (Marseul) in Japan.
195

Table 28. Biometric data of Pachylomalus musculus (Marseul).

APW
PPW
PL
EL
EW

ProW
ProL
PyL
P1L
MSTL

MTTL

0.56-0.71
1.20-1.52
0.66-0.83
1.10-1.47
1.27-1.72
0.61-0.83
0.25-0.29
0.27-0.44
0.42-0.56
0.42-0.54
0.44-0.61(0.66± 0.01) 20(1.35± 0.02) 20(0.76± 0.01) 20(1.27± 0.02) 20(1.52± 0.03) 20(0.73± 0.01) 20(0.27±0.004) 20(0.36± 0.01) 20(O.50± 0.01) 20(0.47± 0.01) 19(0.54± 0.01) 20

clothed with microscopic punctures. Meso-metasternal suture strongly carinate andanteriorly arcuate medially, its lateral end continued to lateral metasternal stria. Disk ofmetasternum strongly shining, and evenly and finely punctate, the punctures being
separated by 4 - 5 times their diameter and becoming sparser laterally. Lateralmetasternal stria strongly carinate, impressed along mesocoxal cavity, its lateral end
attaining to basal 113 of metepisternal-metasternal suture. A deep fovea present nearlateral 1/2 of the lateral metasternal stria. Lateral disk of metasternum with transverse,
alutaceous and microscopic ground sculptures and fine punctures. Mesocoxal stria wellimpressed, extending along the mesocoxal cavity and posteriorly bent on lateral 114 oflateral metasternal disk.
Intercoxal disk of 1st abdominal sternum sparsely and microscopically punctate,with 2 striae on each lateral side.
Protibia dilated, with 4 denticles on outer margin. Meso- and metatibiae
somewhat dilated.
Male genitalia as shown in Fig. 117.
Specimens examined. [Honshu] Saitama-ken: 10"', Urawa, 1O/vi/1954, no collector's name(NA). Nara-ken: 10"', Kasuga, 19/i/1958, T. Shibata (NA).
[Shikoku] Tokushima-ken: 1!f, Kuhina-mura, 20/vii/1953, M. Hirai (NA). Ehime-ken: 1!f,Shiroyama, Matsuyama, 24/v/1949, T. Yano (NA); 1 ex., Kurokawa, Ishizuchi, 2/vi/1952, N. Yato
(NA).
[Kyushu] Fukuoka-ken: 470"'0'" 35!f!f , Mt. Tachibana, Fukuoka, 24/ix, 23/x/1983, 24/v,
6/vi, 25/vii/1984, S. Nomura; 120"'0'" 6!f!f, Kashigu, 8, 15/ix/1983, S. Nomura; 2 exs., Kane-noMisaki, Fukuoka, l/xi/1978, Y. Takakura. Nagasaki-ken: 240"'0'" 6!f!f ,Suwa Shrine, Nagasaki-shi,
9/vi/1983, S. Nomura. Gita-ken: 1!f, Saeki, 20/viil 1953 , K. Kurosa (NA).

Distribution (Fig. 118). Japan (Honshu, Shikoku, Kyushu, Tokara Isles);Taiwan; Burma; Viet-Nam.
Remarks. Being the only Japanese representative of the genus, P. musculus caneasily be recognized by the generic characteristics.
Little is known about the habitat of this spcies. Some specimens were collected in
the litter of forests.
GENUS EULOMALUS COOMAN, 1937
Eulomalus Cooman, 1937: 97 [type species: Eulomalus vermicipygus Cooman, 1937: 98, originallydesignated].

196

The Japanese species included in the genus Eulomalus have not been sufficiently
studied. Probalby 3 or more species occur in Japan. In this papaer, the known Japanese
species are only listed.
Eulomalus lombokanus Cooman, 1937
Eulomalus lombokanus Cooman, 1937: 104 [Lombok]; Mazur, 1984: 137; Hisamatsu, 1985b: 225[photo].

Japanese name: Mizo-chibi-hirata-emma-mushi.

Distribution. Japan (Nansei Isles); Lombok Is.
Eulomalus tardipes (Lewis, 1892)
Paromoalus tardipes Lewis, 1892: 35 [Japan: Honshu].
Eulomalus tardipes: Mazur, 1984: 138; Hisamatsu, 1985b: 225 [photo].

Japanese name: Chairo-chibi-hirata-emma-mushi.

Distribution. Japan (Nansei Isles); Lombok Is.
GENUS PAROMALUS ERICHSON, 1834
Paromalus Erichson, 1834: 167 [type species: Hister jlavicomis Herbst, 1792: 40, designated byWestwood, 1840: 22].
Microlomalus Lewis, 1907a: 318, syononymized by Cooman, 1948: 133. [type species: Histerflavicornis Herbst, 1792: 40].

Key to the Japanese species of the genus Paromalus
1(2)Apical end of marginal elytral stria extending to suture of elytron, and curved at medio-posterior
angle of elytron, then extending anteriorly ........................ P. parallelepipedus (Herbst, 1792)
2(1)Apical end of marginal elytral stria not attaining to medio-apical angle of elytron ................. ..
............................................................................................... P. vernalis Lewis, 1982
Another species, Paromalus omineus Lewis, 1892, occurs in Japan.
SUBGENUS PAROMALUS ERICHSON, 1834

Paromalus (Paromalus) omineus Lewis, 1892
Paromalus omineus Lewis, 1892a: 36 [Japan: Ominesan]; Mazur, 1984: 139; Hisamatsu, 1985b: 225.
Micromalus omineus: Lewis, 1907a: 318

Japanese name: Aka-chibi-hirata-emma-mushi.
Specimens examined. No specimen is availabe in this study.

Distribution. Japan (Honshu); Bhutan?

197

I
I

Fig. 119. Paromnlus parallelepipedus (Herbst). A: Pronotum and left eJytron, dorsal view. B:
Prosternum, mesosternum, metasternum and 1st abdominal sternum. C: Pygidium.

A
O.5mm

~n

~------~--~-~-~-~-~-~-~~~===--------

D

O.5mm

Fig. 120. Paromalus parallelepipedus (Herbst). A: Aedeagus, lateral view. B: Ditto, dorsal view. C:
Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite, dorsal view. E: Ditto, lateralview. F: Male genitalia, 9th and 10th tergites, lateral view. G: Ditto, dorsal view. H: Male
genitalia, 8th sternum, dorsal view. I: Ditto, lateral view. J: Ditto, caudal view.
198

Paromalus (Paromalus) parallelepipedus (Herbst, 1792)(Fig. 119, 120)
Hister parallelepipedus Herbst, 1792: 37.
Hister pusillus Kugelann, 1792: 305, synonymized by Kugelann, 1794: 518.
Hister picipes: Sturm, 1805: 248 [emend.].
Microlomalus parallelepipedus: Lewis, 1907a: 318.
Paromalus parallelepipedus: Lewis, 1892a: 36 [Sapporo, Miyanoshita, Nikko, Nishimura]; Mazur,

1984: 139; Hisamatsu, 1985b: 225 [photo].
Japanese name: Hoso-chibi -hirata-emma-mushi.

Description. Male and female. Body length, PPL, male, 1.99 - 2.36 mm (2.13 ±0.03, n=16), female, 1.85 - 2.23 mm (2.06 ± 0.03, n=20), PEL, male, 1.81 - 2.13
mm (1.98 ± 0.02, n=16); female, 1.73 - 2.13 mm (1.94 ± 0 02, n=20). Width, male,
1.08 -1.24 mm (1.16 ± 0.01, n=16), female, 0.99 - 1.26 mm (1.14 ± 0.02, n=20).
Biometric data are given in Table 29. Body oblong, black and shining; tibiae,
mouthparts and antennae rufopiceous.
Frontal stria of head impressed and complete, its lateral portions parallel. Disk
feebly depressed on apical 112, and evenly covered with moderate punctures which areseparated by 1- 3 times the diameter, with microscopic punctures intermingled.
Pronotal sides (Fig. 119A) feebly arcuate and convergent forward on basal 3/4,
then strongly convergent apically. Emarginated portion of anterior margin of pronotum
bisinuate. Marginal stria well impressed and complete laterally and anteriorly. Disk
evenly covered with moderate punctures which are separated by 2 - 4 times theirdiameter or more; interspace among the moderate ones sparsely clothed withmicroscopic punctures. Antescutellar area without a large puncture, but feebly
depressed.
Epipleural marginal stria weakly impressed on apical 112. Marginal elytral stria
strongly carinate and complete, its apical end extending along posterior margin,attaining to sutural angle, then curved and a little extending basally. Epipleura sparsely
clothed with microscopic punctures. Disk of elytra sparsely covered with coarse
punctures which are separated by 2 - 5 times their diameter, except on the following
areas: a narrow band along suture which is clothed with microscopic punctures andfeebly elevated, a narrow area along the basal margin and an extreme apical band which
are impunctate; surface between the coarse punctures sometimes clothed withmicroscopic punctures. Dorsal stria absent, but a weak oblique depression present onlaterobasal 112.
Propygidium densely covered with coarse, round and shallow punctures, whichbecome finer and sparser around margin; surface between the punctures with weaklytransverse alutaceous microsculpture. Pygidium (Fig. 119C) evenly clothed with finepunctures which are separated by about 1.5 (1- 2) times the diameter; surface between
the fine punctures intermingled with microscopic punctures. In female, punctures ofpygidium similar to those of male, and a deep sculpture present as shown in fig. 119C.
Anterior margin of prosternal lobe (Fig. 119B) truncate medially; the anterior
angles of the lobe broadly round; marginal stria absent anteriorly, weakly impressed
laterally on basal 112; inside of basal angle deeply excavate; disk sparsely covered withmoderate punctures which are separated by 1 - 5 times the diameter. Punctation of
prosternal process similar to that of the lobe. Descending lateral stria strongly carinate
199

Table 29. Biometric data of Paromalus parallelepipedus Herbst.
Female

Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSIL
MrTI,

0.51-0.59 (0.56±0.005) 16
1.00-1.14 (1.07± 0.01) 16
0.59-0.73 (0.68± 0.01) 16
1.12-1.38 (l.23± 0.02) 16
1.08-1.24 (1.16± 0.01) 16
0.55-0.63 (0.59± 0.01) 16
0.22-0.28 (0.25± 0.01) 16
0.32-0.39 (0.37± 0.01) 16
0.39-0.49 (0.45± 0.01) 16
0.39-0.49 (0.44± 0.01) 16
0.45-0.53 (0.50±0.005) 16

0.51-0.61 (0.55± 0.01) 20
0.95-1.14 (1.06± 0.01) 20
0.61-0.73 (0.67± 0.01) 20
1.02-1.36 (1.21± 0.02) 20
0.99-1.26 (1.14± 0.02) 20
0.49-0.63 (0.58± 0.01) 20
0.20-0.30 (0.24±0.005) 20
0.24-0.39 (0.32± 0.01) 20
0.37-0.47 (0.44± 0.01) 20
0.35-0.47 (0.41± 0.01) 20
0.39-0.53 (0.47± 0.01) 20

and complete.
Anterior margin of mesosternum (Fig. 1198) strongly emarginate medially;
marginal stria absent anteriorly and deeply impressed and complete laterally, its
posterior end continued to basal end of lateral metasternal stria; disk sparsely covered
with moderate punctures, which are separated by 2 - 5 times their diameter, except on
area surrounded by transverse stria and meso-metasternal suture; transverse stria wellimpressed, acutely angulate on each lateral 112 and arcuate medially. Meso-metastemal
suture indistinct. Intercoxal disk of metasternum sparsely and coarsely punctate, thepunctures becoming finer medially, being separated by 2 - 3 times their diameter;surface between coarse punctures intermingled with fine punctures. Lateral disk
scattered with moderate punctures which are separated by 2 - 5 times their diameter ormore. Lateral metastemal stria strongly carinate, extending obliquely and posteriorly,
and attaining to the middle of metastemal-metepisternal suture. Post-mesocoxal stria
extending along mesocoxal cavity, its outer end strongly bent posteriorly.
Punctation of intercoxal disk of 1st abdominal sternum similar to that of sides ofmetastemal intercoxal disk; 1st abdominal stria complete and well impressed.
Protibia with 3 denticles on outer margin. Mesotibia with 4 spinules on outermargin, the basal one small. Metatibia with 1 spinule near the apex.
Male genitalia as shown in Fig. 120
Specimens examined. [Hokkaido] 4 exs., Kami-Shihoro, 17-24/v, 19-23/vi/1989, K. Haga; 12
exs., Nopporo, 5,19,20,22,26, 30/vi/1987, M. Ohara; 12 exs., Ditto, 19/v/1936, M. Inoue (HFFP);
1 ex., Kitanoue, 23/viii/1949, M. Inoue, collected form Picea glehnii. (HFFP); 21 exs., Kamui-dake,
Jozankei, Il1vil 1990, M. Ohara; 4 exs., Abuta, 4/vi/1951, M. Inoue, collected from Picea jezoensis.
(HFFP); 3 exs., Nanae-cho, 19/vii/1948, M. Inoue (HFFP).
[Honshu] Iwate-ken: 2 exs., Numakunai, 25/vi/1948, M. Inoue, red pine (HFFP); 1 ex.,
Mizusawa, 28/vi/1948, M. Inoue (HFFP). Fukushima-ken: 6 exs., Yunohana, Tateiwa, Minami-Aizu,
6/v/1949, Y. Kurosawa (NSMT); 1 ex., Wakamatsu, 30/iv/1949, T. Yoneyama (NSMT). Tokyo-to: 1ex., Omaro, Inagi, 10/iv/1988, K. Sawada; 4 exs., Akaidani, 5/vl 1949, K. Nagayama (NA).

Distribution. Japan (Hokkaido, Honshu); Europe; Crimea; Caucasus; Siberia.
Remarks. Small specimens of Paromalus parallelepipedus resemble P.
omineus, but can be distinguished by the different shape of the transverse mesosternal
stria and the body color.
This species is frequently collected on Abies sachalinensis in Hokkaido. Mazur(1981) recorded it from the following trees: Pinus silvestris, Pice a abies, Abies alba,
200

Quercus, Fagus silvatica, Salix, and Tomicus piniperda.
Paromalus (Paromalus) vernalis Lewis, 1892
(Fig. 121, 122)
Paromalus vernalis Lewis, 1892a: 35 [Japan; Nara, Oyayama, Yuyama]; Hisamatsu, 1968: 31[Ryfikyii]; 1985b: 225 [photo]; Mazur, 1984: 134.
Microlomalus vernalis: Lewis, 1907a: 318.

Japanese name: Ko-chibi-hirata-e=a-mushi.

Description. Male and female. Body length, PPL, male, 1.42 - 1.64 mm (1.51 ±
0.03, n=6), female, 1.39 - 1.66 mm (1.53 ± 0.02, n=l1), PEL, 1.32 - 1.54 mm (1.41± 0.03, n=7), female, 1.23 - 1.61 mm (1.44 ± 0.03, n=l1). Width, male, 0.83 0.98 mm (0.88 ± 0.02, n=7), female, 0.76 - 0.98 mm (0.89 ± 0.21, n=11). Biometricdata are given in Table 30. Body black, sometimes reddish brown and shining; femora,
tibiae, mouthparts and antennae rufopiceous.
Frontal stria of head deeply impressed, its sides feebly divergent anteriorly; disk
sparsely covered with moderate and deep punctures which are separated by 4 - 6 times
their diameter and become denser apically; surface between the moderate ones rarelyintermingled with microscopic punctures.
Pronotal sides (Fig. 121A) feebly arcuate and convergent forward. Marginal stria
distinctly carinate and complete anteriorly and laterally. Disk of pronotum evenly
covered with coarse punctures which are separated by about 2 (1- 3) times the diameterand become finer medio-anteriorly, with other microscopic punctures sometimes
intermingled. AntcscutcIlar area without a large depression.
Epipleural marginal stria weakly impressed on apical 112. Elytral marginal stria
deeply impressed and sparsely and coarsely carinate, its apical end extending along

Fig. 121. Paromalus vernalis Lewis. A: Pronotum and left elytron, dorsal view. B: Prosternum,
mesosternum, metasternum and 1st abdominal sternum. C: Pygidium.

201

apical margin, attaining to near the middle of apical margin of elytron. Dorsal stria
absent, but an oblique depression shortly present at lateral 112 on basal 113. Disk of
elytra evenly covered with coarse, round and shallow punctures except on the following
areas: a narrow band along suture which is impunctate and feebly elevated, a narrow
band along basal margin, and an extreme apical band; the coarse punctures becomingcoarser near the oblique depression and gradually sparser apically.
Propygidium densely covered with coarse, round, shallow punctures andintermingled with fine punctures, and finely punctate narrowly along the margin.
Pygidium evenly covered with fine punctures which are separated by about 2 (1 - 3)times the diameter. In female, punctation of the pygidium as in male and a deep
sculpture present as shown Fig. 121C; the sculpture consisting of 2 parts, a transverse
deep part basally and a circular one on apical 2/3 area, inside the circular one deeply andirregularly rugose.
Anterior margin of prosternal lobe (Fig. 121B) nearly round; marginal stria
sometimes present anteriorly and impressed deeply on laterobasal 1/2; disk irregularly
scattered with fine punctures. Punctation of prosternal process similar to that of theprosternallobe. Lateral descending stria strongly carinate, short and complete.
Anterior margin of mesosternum (Fig. 121B) strongly emarginate medially.
Marginal stria absent anteriorly, and complete and carinate laterally. Punctation of disk
similar to that of the prosternal process; impunctate and depressed behind transverse
stria and narrowly along lateral marginal stria. Meso-metasternal suture absent.
Intercoxal disk of metasternum sparsely covered with coarse and oblong punctures
which are separated by 2 - 3 times their diameter and become finer medially and coarser
on lateral 113, the punctures on lateral third 3 times as large as those of the median 113.
Lateral metasternal stria strongly carinate, extending obliquely and posteriorly, andattaining to the middle of metasternal-metepisternal suture. Lateral disk sparsely covered
with coarse punctures which are smaller than those of sides of intercoxal metasternal
disk. Post-mesocoxal stria extending along mesocoxal cavity, its outer end strongly
bent posteriorly.
Intercoxal disk of 1st abdominal sternum sparsely and coarsely punctate, thepunctures becoming finer medially; 1st abdominal stria completely present on eachlateral side.
Protibia with 4 denticles on outer margin. Mesotibia with 3 spinules on outermargin. Metatibia with a spinule near the apex.
Male genitalia as shown in Fig. 122.
Specimens examined. [Honshu] Fukushima-ken: 1 ex., Niizuru, Onuma, 6/iv/1949, K.
Nagayama (NA). Nara-ken: Id", Nara Park, Nara, 7/x1l987, K. Sawada (NA); 1 ex., Mt. Kasuga,
26/ix/1948, M. Golo (NA); 1 ex., Ditto, 9/v/1952, K. Sawada (NA).
[Shikoku] Kochi-ken: 1 ex., Yakyo, 4/ixl 1945, K. Sakaguchi, collected under bark (NA).
[Kyfishu] Miyazaki-ken: 1 ex., Koshima, 24/vii/1953, M. Yoshida (NA). Kagoshima-ken: 1!f,Kurino, 3/iil 1985, M. Ohara; 3 exs., Sata, 9/iiil 1964, no collector's name (NA).
[Nansei Isles] Tokara-Nakano-shima Is.: 1 ex., Nakano-shima, 12/vi/1953, S. Ueno (NA); Id",
3 exs., Ditto, 25/v, 9/vi/1953, T. Nakane (NA); 1 ex., Ditto, 5/vi/1953, S. Miyamoto (NA); 1!f,
Funakura, Mitake, Nakano-shima, 18/viii/1982, Y. Takai. Tokara-Takara-jima Is.: 4 exs., Takara-jima,
6/vii/1953, T. Nakane. Amami-Oshima Is.: 1!f, Kinsakubaru, Naze, 30/vi/1984, M. Ohara. Ishigaki
Is.: 1!f, Mt. Banna, Ishigaki, I/iv/1976, N. Nishikawa.

202

......

/_.

(

A

~.

c

D

F

~I

O.25mm ,Fig. 122. Paromalus vernalis Lewis. A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and 10th
tergites and 9th sternum (spicule), lateral view. F: Ditto, dorsal view. G: Male genitalia, 8thsternum, dorsal view. H: Ditto, lateral view. I: Ditto, caudal view.
Table 30. Biometric data of Paromalus vema lis Lewis.
Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Female0.39-0.47 (0.42±0.01) 11
0.68-0.90 (0.80±0.02) 11
0.42-0.52 (0.49±0.01) 11
0.76-1.00 (0.87±0.02) 11
0.76-0.98 (0.89±0.02) 11
0.35-0.46 (0.41±0.01) 11
0.12-0.19 (0.16±0.01) 11
0.19-0.25 (0.22±0.01) 11
0.27-0.34 (0.31±0.01) 11
0.27-0.34 (0.30±0.01) 11
0.29-0.37 (0.33±0.01) 10

0.41-0.44 (0.42±0.01) 7
0.76-0.88 (0.79±0.02) 7
0.42-0.49 (0.48±0.01) 7
0.83-1.01 (0.92±0.02) 7
0.83-0.98 (0.88±0.02) 7
0.37-0.46 (0.41±0.01) 6
0.12-0.17 (0.15±0.01) 6
0.24-0.27 (0.25±0.01) 6
0.27-0.34 (0.32±0.01) 7
0.25-0.32 (0.29±0.01) 7
0.30-0.37 (0.34±0.01) 6

Distribution. Japan (Honshu, Shikoku, Kyushu, Nansei Isles); Taiwan; Russia(Primorskiy Kray).
Remarks. Paromalus vernalis resembles P. parallelepipedus, but isdistinguishable by the marginal elytral stria not attaining the medio-apical angle of theelytron (see the key and description).

2036.3.8 SUBFAMILY ABRAEINAE MARSEUL, 1857
Key to the Japanese tribes and genera of the subfamily Abraeinae
1(2)
2(1)
3(4)

4(3)

Posterior tarsus consisting of 4 tarsomeres. , .... ,."""""" .. ,...... " ..Tribe Acritini Wenzel, 1944." .. ".".".".""""."."."" . .,,,,,.,,,,.,,.,,.,,,,,,,,,,,,,,.,,,,,,,,, Genus Acritus J, L, LeConte, 1853
Posterior tarsus consisting of 5 tarsomeres,
Pronotum with a longitudinal depression laterally and with a transverse line on middle or near
middle. Prostemal process with an X-shaped excavation, Mesosternum and metasternum fused .
. " .................................... " ........................................... Tribe Plegaderini Portevin, 1929..................... " ........................................................... Genus Plegaderus Erichson, 1834
Pronotum smooth, without line, Pro sternal process with no excavation. Mesosternum andmetasternum separated by meso-metasternal suture, """""""",,,Tribe Abraeini Portevin, 1929... ", ....... " .... " ...... " .... " ............................................ Genus Chaetabraeus Portevin, 1929

TRiBE ABRAEINI MARSEUL, 1857
GENUS CHAETABRAEUS PORTEVIN, 1929
Chetabraeus Portevin, 1929: 624 [type species; Hister globulus Creutzer, 1799: 83, designated bymonotypy].
Chaetabraeus: Mazur, 1989: 31, emendation for Chetabraeus Portevin, 1929.
Chartabraeus: Witzgall, 1971: 162 [error].

Recently, Mazur (1989) has studied this genus. In his opInIOn this genus ischaracterized by the following characters: (1) long and pentagonal propygidium, (2)strongly convex dorsum and usually brownish black or black coloration (3) living in thedung of various mammals, on carrion or in rotting vegetation.
Key to the Japanese species of the genus Chaetabraeus
1(2)
2(1)

3(4)
4(3)

Disk of prosternum with a deep fovea on each side, "."""."""" C. bonzicus (Marseul, 1873)
Disk of prosternum without deep fovea on each side.
Body smaller; 1.37 - 1.64 mm, black with red luster; legs reddish brown. """""" .. """"" .. "...................... " ................................................................... C. co haeres (Lewis, 1898)
Body larger; 1.84 rum, black with green luster; legs yollowish brown. """"""",,""'" C. sp.

Chaetabraeus bonzicus (Marseul, 1873)(Fig. 123, 124, 125, 126, 127)
Abraeus bonzicus Marseul, 1873: 221, 226 [Japan: Nangasaki (=Nagasaki, Kyilshil)]; Lewis, 1898:
181 [Otaru, Yezo (= Hokkaido)]; Jakobson, 1911: 653; Bickhardt, 1916-17: 74, t. 3, fig. 23;
Cooman, 1937: 38; Reichardt, 1941: 130,131, fig. 59.
Chetabraeus bonzicus: Mazur, 1984: 8.
Chaetabraeus bonzicus: Mazur, 1989: 32.
Abraeus (Chetabraeus) bonzicus: Kryzhanovskij and Reichardt, 1976: 96.

Japanese name: Kuro-tsubu-emma-mushi.

Description. Biometric data are given in Table 31. Body oval, strongly convex
and black; tarsi and antennae rufopiceous. Body length, PPL, 1.84 - 2.25 mm (2.08 ±0.03, n=20), PEL, 1.74 - 2.21 mm (1.97 ± 0.03, n=20). Width, 1.62 - 1.94
mm (1.78 ± 0.02, n=20).
204

Head densely covered with coarse and longitudinal oblong punctures, andsparsely with stout and erect setae, which are about 113 as long as the transverse
diameter of the eye; surface with a transverse carina behind each antennal cavity (Fig.
124F).

Anterior margin of pronotum emarginate, the median portion nearly straight;
lateral sides evenly arcuate and weakly convergent to apices on basal 2/3, thence
strongly arcuate and convergent to the apices. Disk of pronotum densely covered withcoarse and round punctures, the punctures being as large as those of the head andseparated by 0.3 - 0.5 times their diameter, becoming denser and a little finer apically,and sparsely with erect setae which are as large as those of the head.
Epipleura of elytra without stria. All dorsal striae absent. Disk of elytra with
punctation and erect setae, the punctures being similar to, but a little denser than,pronotal ones, usually fused to form longitudinal rugae on lateral and apical area, andbecoming finer on area along the sutural line.
Propygidium pentagonal, its disk with punctation and setae similar to those of
pronotum, the coarse punctures being separated by 0.3 - 0.5 times their diameter.
Pygidium longitudinal oblong, its punctation similar to propygidial one but somewhat
sparser and shallower.
Prosternal process nearly quadrangular; its anterior angles extended lateroanteriorly, acute, and with a deep fovea (Fig. 12SA); anterior margin completely
impressed, with a transverse depression a little distant form the margin; disk of theprocess sparsely covered with coarse punctures which are separated by 0.5 - 1 times
the diameter and as coarse as elytral ones, with other moderate punctures intermingledwith them.
Mesosternum narrow and transverse; anterior margin straight medially andcurved laterally along the posterior margin of procoxa; the anterior and lateral sides
distinctly carinate and marginate; disk of mesosternum sparsely covered with coarse
punctures which are a little coarser than elytral ones and separated by 0.3 - 1 times thediameter, and fine punctures sparsely intermingled among the coarse ones. Meso-

Fig. 123. Adults in dorsal view. A: Chaetabraeus bonzicus (Marseul). B: C. cohaeres (Lewis.)

205

Fig. 124. A - E: Chaetabraeus cohaeres (Lewis). F: C. bonzicus (Marseul). A: Adult, dorsal view. B:
Surface of elytron. C: Adult, ventral view. D: pygidium. E: Right protibia, dorsal view. F:
Head, frontal view.

206metasternal suture distinctly impressed and strongly crenate. Metasternum with shortlateral stria, which is obliquely and posteriorly extended, its apical end attaining to thebasal 113 of metasternum; intercoxal disk evenly covered with coarse, shallow andround punctures which are a little coarser than those of the mesosternum and separated
by 0.5 - 1 times the diameter, other fine punctures being intermingled among the coarse
ones; longitudinal mid line clearly and deeply impressed; lateral disk densely covered
with large, round and somewhat deep punctures, fine punctures being intermingled on
apical 2/3 (Fig. 12SA).
Intercoxal disk of 1st abdominal sternum narrow and transverse, with punctation
similar to that of mesosternum.
Protibia slender, its outer margin with many small setae.
Male genitalia as shown in Fig. 126.

I
Fig. 125. Prosternal process and meso- and metasternum. A: Chaetabraeus bonzicus (Marseul). B: C.
cohaeres (Lewis).

A~~;=:'::;:-:;::':::';::':':;;"~:~:"':;;;'~"

O.25rnm,~~~,~,,~~~~~~:~o;i~~Fig. 126. Chaetabraeus bonzicus (Marseul). A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and10th tergites and 9th sternum (spicule), dorsal view. F: Spermatheca.
207

Table 31. Biometric data of Chaetabraeus banzicus (Marseul) and C. cahaeres (Lewis).

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

C. banzicus

C. cahaeres

0.61-0.76 (0.68±0.01) 20
1.47-1.79 (1.61±0.02) 20
0.61-0.96 (0.82±0.02) 20
0.91-1.45 (1.18±0.03) 20
1.62-1.94 (1.78±0.02) 20
0.71-1.00 (0.85±0.02) 20
0.47-0.56 (0.53±0.01) 20
0.44-0.66 (0.56±0.01) 20
0.49-0.61 (0.58±0.01) 20
0.47-0.59 (0.53±0.01) 20

0.47-0.59 (0.53± 0.01) 28
1.15-1.40 (1.25± 0.01) 28
0.51-0.69 (0.61± 0.01) 28
1.25-1.49 (1.39± 0.01) 28
0.76-1.03 (0.91± 0.01) 28
0.54-0.74 (0.66± 0.01) 28
0.37-0.51 (0.43± 0.01) 28
0.37-0.49 (0.41± 0.01) 25
0.37-0.44 (0.42±0.005) 27
0.32-0.44 (0.40± 0.01) 26
0.37-0.49 (0.42±0.004) 26

0.54-0.66 (0.60±0.01) 20

Female genitalia: spermatheca as shown in Fig. 126F.
Specimens examined. [Hokkaido] 2 exs., Sapporo, 25/iii, 23/v/1985, M. Ohara; 1 ex., Hitsujiga-oka, Sapporo, 20/vii/1985, M. Ohara; 1 ex., Jozankei, 29/vi/1985, M. Ohara; 1 ex., Nopporo,
15/ix/1989, M. Ohara.
[Honshu] Fukushima-ken: 1 ex., Arai, Mizubayashi, 12/xi/1975, K. Tazoe; 3 exs., Wakamatsu,
29/vi/1948, Y. Kurosawa (NSMT); 1 ex., Ditto, 5/vi/1949, T. Yoneyama (NSMT). Ibaraki-ken: 3exs., Yatabe, 5/vi/1986, M. Kiuchi; 1 ex., Kamisakai, Sakura-mura, 4/v11985, H. Haga. Chiba-ken: 1ex., Oarni, 19/vi/1984, M. Ohara. Tokyo-to: 2 exs., Aihara, 16/v/1987, H. Takizawa; 1 ex.,
Suginami, 14/iv/1979, M. Ohara. Nagano-ken: 4 exs., Kakeyu, Maruko, Chiisagata, 3/iii/1969, M.
Kiuchi. Hyogo-ken: 4 exs., Kobe, 25/vi, 28/viil 1930, no colIector's name (BSM).
[Kyfishu] Kagoshima-ken: 1 ex., Shimadomari, Sata, 4/v/1992, K. Haga.

Distribution (Fig. 127). Japan (Hokkaido, Honshu, Shikoku, Kyushu);
Prymorskij Kray.
Remarks. Chaetabraeus bonzicus is very similar to C. cohaeres; however, it
differs from the latter by the large size and the deep foveae on the prosternal process.
See also the description of C. cohaeres.
This species occurs in many kinds of rotting organic substances, in manure,animal dropping, carrion, etc.
Chaetabraeus cohaeres (Lewis, 1898), sp. rev.
(Fig. 123, 124, 125, 127)
Abraeus cahaeres Lewis, 1898: 181 [Hongkong]; Desbordes, 1919: 409.
Abraeus banzicus: Bickhardt, 1913: 177 [partim, Taiwan].
Chaetabraeus cahaeres: Mazur, 1989: 32.

Japanese name: Ko-kuro-tsubu-emma-mushi.

Description. Body length, PPL, 1.37 - 1.64 mm (1.54 ± 0.01, n=28), PEL,
1.35 - 1.62 mm (1.50 ± 0.01, n=28), Width, 0.76 - 1.13 mm (0.92 ± 0.01, n=29).
Extremely similar to C. bonzicus but differs as follows: smaller in size (Table31); punctation of propygidium a little sparser (Fig. 1240); anterior angles of prosternal
process without deep fovea, only weakly depressed; lateral disk of metasternum
densely covered with coarser punctures, with no fine punctures intermingled;
longitudinal mid line of metasternum present, but obscure, not strongly impressed; male
genitalia not expanded at caudal apex.
208

·' ~, • Chaetabraeus bonzicus. /~ .... Chaetabraeus cohaeres/..,~~-./'---.[

/

~

/

"

'/ " ,''

Fig, 127. Collection sites of Chaetabraeus bonzicus (Marseul) and C. cohaeres (Lewis) in Japan.
Specimens examined. [Nansei Isles] Yaku-shima Is.: 21 exs., 2/iv/1985, M. Kiuchi. TokaraNakano-shima Is.: I ex., 14/vii/1982, M. Ohara. Tokara-Takara-jima Is.: 11 exs., 18/iii/1986, E.
Matsui. Iriomote Is.: 1 ex., Otomi, 17/iii/1985, S. Masuda; 3 exs., Ditto, K. Nakamine. Yonaguni Is.:2 exs., Urabu-take, 211II/1993, Y. Hirano.
[Taiwan] Nantou pref.: 40 exs., Songkang, alt. 2000 m., 3 I1iii, 2, 5, 14/iv/1986, M. Ohara; 1ex., Nanshanxi, 281xii11974, K. Masumoto. Pingtung pref.: 28 exs., Sizhongchi, 111v11986, M.
Ohara. Lanyu Is. (Orchid Is.): 22 exs., 18-22IivI1986, M. Ohara.

Distribution (Fig. 127). Japan (Nansei Isles); Taiwan; China (Hongkong). New
to Japan.
Remarks. Bickhardt (1913) erroneously synonymized this species with C.
bonzicus. It is distinct from the latter, being distingushied by the characteristics given
above.
Chaetabraeus sp.
In general appearance similar to bonzicus and cohaeres, but distinguishable by
the following characteristics: fovea of prosternum absent; punctation of intercoxal diskof metasternum much denser than in bonzicus; body with green luster; body nearly aslarge as bonzicus and distinctly larger than cohaeres. Biometric data are as follows:
PPL 1.84, PEL 1.79, APW 0.59, PPW 1.47, EL 1.62, EW 1.13, ProW 0.69, ProL
0.51, PyL 0.47, PTL 0.49, MSTL 0.47, MTTL 0.49. At present it is uncertain to me
whether this form is a good species, because it is represented by a single specimen.
Specimen examined. [Nansei Isles] Iriomote Is.: 1 ex., Iriomote-jima, 221iii11982, M. Kiuchi.

209ventral view. C: Head. D:
Fig. 128. Plegaderus marseuli Reitter. A: Adult, dorsal view. B: Ditto,
view. F: Protibia, dorsal
ventralnum,
mesosterandum
Propygidium and pygidium. E: Prostern
view.
210

TRIBE PLEGADERINl PORTEVIN, 1929
GENUS PLEGADERUS ERICHSON, 1834
Plegaderus Eri chs on, 1834: 203 [type species Hister caesus Herbst, 1792: 40, designated byThomson, 1859: 75]; Lacordaire, 1854: 278; Marseul, 1856: 256; Schmidt, 1885a: 284;
Ganglbauer, 1899: 397; Reitter, 1909: 204; Jakobson, 1911: 83,97; Arnett, 1962: 373,379;
Witzgall, 1971: 160; Kryzhanovskij and Reichardt, 1976: 85; Mazur, 1973: 19; 1981a: 55;
1984: 12; Vienna, 1980:73.
SUBGENUS PLEGADERUS ERICHSON, 1834

Plegaderus (Plegaderus) marseuli Reitter, 1877
(Fig. 128, 129)
Plegaderus marseuli Reitter, 1877: 371 [Japan].
Plegaderus (Plegaerus) marseuli: Nakane, 1963: 67; Kryzhanovskij and Reichrdt, 1976: 86 [key];

Mazur, 1984: 13; Hisamatsu, 1985b: 222.
Japanese name: Mune-mizo-tsubu-emma-mushi.

Description. Body oblong-oval. Cuticles shining and black; tibiae, tarsi andantennae rufopiceous. Body length, PPL, 1.40 - 1.52 mm (1.47 ± 0.02, n=8), PEL,
1.35 - 1.49 mm (1.43 ± 0.01, n=8), Width, 0.93 - 1.03 mm (0.99 ± 0.01, n=8).
Biometric data are given in Table 32.
Head (Fig. 128C) with a convex area behind each antenna! cavity; surface
unevenly covered with coarse and setiferous punctures which are separated by 1 - 3times the diameter, the punctures becoming denser on epistoma.

I

O.lmm,

F........... .

~
p?

'

.

£ ---.'.--------.:- \"-.
E··· ..········· .........
"':: :

Fig. 129. Plegaderus marseuli Reitter. A: Aedeagus, lateral view. B: Ditto, dorsal view. C: Apex of
aedeagus, ventral view. D: Male genitalia, 8th tergite and sternum, dorsal view. E: Ditto, lateralview. F: Male genitalia, 9th and 10th tergites and 9th sternum (spicule), dorsal view. G: Ditto,
lateral view. H: Spermatheca and spermathecal gland.
211

Table 32. Biometric data of Plegaderus marseuli Reitter.

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

0.46-0.51
0.83-0.93
0.57-0.66
0.74-0.90
0.93-1.03
0.44-0.56
0.10-0.15
0.29-0.34
0.34-0.39
0.34-0.41
0.39-0.44(0.48± 0.01)(0.88± 0.01)(0.62± 0.01)(0.81± 0.02)(0.99± 0.01)(0.49± 0.01)(0.14± 0.01)(0.31± 0.01)(0.37±0.004)(0.36± 0.01)(0.41± 0.01)

8
8
8
8
8
8
8
8
8
8
8

Pronotum strongly convex; anterior margin broadly emarginate, with a complete
marginal stria; sides divergent posteriorly on basal 3/4, and strongly convergent
anteriorly on apical 114, its lateral marginal stria clearly impressed along the margin andnearly complete, but not attaining to the apical angle. Surface of pronotum divided into
2, anterior and posterior, areas by a transverse line at apical 113; the anterior area with alongitudinal convex area on each lateral side, the inside margin of the convex area
complete and distinct, and the disk of the convex area densely and coarsely punctate; theremain of the anterior area evenly, rather sparsely covered with coarse punctures which
are separated by about 1.5 (1 - 2) times the diameter and as coarse as those of head;along the convex area there is a narrow impunctate area inside; the posterior area
laterally with a longitudinal convex area which is, however, not distinct, being only
weakly convex, and is sparsely and finely punctate; the remain of the posterior area
sparsely clothed with fine punctures which are separated by 1 - 4 times the diameter.
Epipleura not concave. Marginal elytral stria entirely impressed but obscure on
apical 112. A narrow area along epipleural margin impunctate. No dorsal striae present,
but oblique humeral stria deeply impressed on basal 114 and another short stria(longitudinal depression) present near the middle of anterior margin of elytron. Surfaceof elytra densely covered with somewhat longitudinal oblong and coarser punctures
which are a little coarser than those of head and separated by 0.5 - 1 times the diameter,
the punctures becoming sparser around scutellum; a narrow band along sutural line andextreme apex impunctate.
Propygidium (Fig. 1280) densely covered with coarse, round and deeppunctures which are separated by 0.5 - 1 times the diameter, and with a transverse bandof moderate punctures along the posterior margin. Pygidium (Fig. 1280) somewhat
sparsely covered with coarse, round and deep punctures which are separated by about
1.5 (1 - 2) times the diameter.
Prostemal process with an X-shaped excavation (Fig. 128E), which is present
near basal 1/3 on the process and densely covered with long hairs; anterior margincompletely with marginal stria, the lateral end of the stria continued to apex of outermargin of the X-shaped excavation; lateral side of the process completely marginate;disk of apical 112 of the process strongly convex, and sparsely covered with setiferous
fine punctures which are separated by 2 - 3 times their diameter; lateral disk impunctate.
Antennal groove deeply excavate, round and present on each lateromedian 116.
212

Meso-metasternal suture absent. Mesosternum and metasternum fused. Intercoxal
disk of meso-metasterna shallowly sulcate on longitudinal midline, which is 113 as wide
as anterior margin of mesosternum, the sulcus becoming narrower posteriorly; surfaceof disk sparsely and coarsely punctate, the punctures as coarse as elytral ones andseparated by 1 - 4 times the diameter. Two lateral metasternal striae present; internal
one strongly carinate basally, and its apical end extended posteriorly and obliquely,
attaining to the apical margin of metacoxae; external one (probably homologous with theposterior mesocoxal stria in Wenzel and Dybas, 1941) weakly carinate, its apical end
attaining to the posterior comer of metasternum. Surface between these 2 lateralmetasternal striae sparsely with fine punctures. Lateral disk of metasternum sparsely
covered with several large, round, and deep punctures. Metepisternum with a row
consisting of large and dense punctures.
Intercoxal disk of 1st abdominal sternum striate on each side, the stria carinate,
its apical end attaining to the apical 113; surface densely and coarsely punctate, thepunctures separated by about their own diameter and as coarse as those of intercoxal
disk of metasternum.
Protibia (Fig. 128F) slender, but progressively broader on apical 112, with 6
denticles on outer margin. Metatibia with 4 denticles on outer margin.
Male genitalia as shown in Fig. 129A - O.
Female genitalia: spermatheca as shown in Fig. 129H.
Specimens examined. [Hokkaid6] 1 ex., Kami-Shihoro, 9/vi/1989, H. Haga; 10 exs., Kamuidake, J6zankei, ll/vi/1990, M. Ohara; 1 ex., H6heiky6, J6zankei, 16/vii11988, S. Kud6; 1 ex., Ditto,
16/vii/1988, M. Ohara; 1 ex., J6zankei, 25/v/1990, M. Ohara.
[Honshu] Gifu-ken: 1 ex., Kawayu, Hida, 31/vii/1954, T. Nakane (NA). Wakayama-ken: 6 exs.,
Mt. Rokk6, Naga-gun, 4/i/1950, M. Aoyagi (NA); 1 ex., Ditto (NSMT). Okayama-ken: 10" 1!f, 1
ex., Okayama, 23/vi/1948, M. Inoue (HFFP).

Distribution. Japan (Hokkaid6, Honshu, Shikoku, KyOshO).
Remarks. Plegadeus marseuli is similar to P. shikokensis, but differs by thelarge size, the fine punctation of the surface and the divided convex area on the lateralpronotal area.
This species was found under the bark of Pinus in galleries of bark beetles.
Plegaderus (Plegaderus) shikokensis Hisamatsu, 1985
Plegaderus (Plegaderus) shikokensis Hi samatsu , 1985a: 5 [Japan: Mt. Ishizuchi, Ehime, Shikoku];

Hisamatsu, 1985b: 222, pI. 40, fig. 8 [noted, photo]; Hirano, 1986: 42 [HonshU].
Japanese name: Chairo-mune-mizo-tsubu-emma-mushi.
Specimens examined. No specimen was available in this study.

Distribution. Japan (Honshu, Shikoku).
TRIBE ACRITINI WENZEL, 1944

The Japanese species included in the tribe Acritini have not been sufficiently
studied until now. Probably 4 or more species occur in Japan. In this papaer, theknown Japanese species are only listed.

213

GENUS ACRITUS J. L. LECONTE, 1853
Acritus J. L. LeConte, 1853: 288 [type species: Rister nigricornis Hoffmann, 1803: 127, designatedby Thomson, 1859: 75]; Lacordaire, 1854: 281; Marseul, 1856: 595; Sclunidt, 1885b: 284;
Reitter, 1886: 278; Ganglbauer, 1899: 405; Reitter, 1909: 294; Jakobson, 1911: 642, 653;
Bickhardt, 1916-17: 63,76; Reichardt, 1941: 84,138 (part); Wenzel, 1944: 58.

SUBGENUS ACRITUS J. L. LECONTE

Acritus (Acritus) komai Lewis, 1879
Acritus komai Lewis, 1879: 461 [Japan: Nagasaki]; Reichardt, 1941: 141, 147.
Acritus insularis Sharp in Blackburn and Sharp, 1885: 129, synonymized by Kryzhanovskij andReichardt, 1976: 105.
Acritus volitans Fall, 1901: 238, synonymized by Kryzhanovskij and Reichardt, 1976: 105.
Acritus optatus Cooman, 1947: 423, synonymized by Kryzhanovskij and Reichardt, 1976: 105.
Acritus oregonensis Hatch, 1962: 256, synonymized by Kryzhanovskij and Reichardt, 1976: 105.

Japanese name: Arame-yofushi-emma-mushi.

Distribution. Japan (KyfishQ, Nansei Isles); China; North Africa; France;
Mariana Isles; Hawaii; Oregon, South California.
SUBGENUS PYCNACRITUS CASEY, 1916
Pycnacritus Casey, 1916: 252 [type species: Acritus discus LeConte, 1853: 289]; Mazur, 1987: 31.

Acritus (Pycnacritus) homoeopathicus Wollaston, 1857
Acritus homoeopathicus Wollaston, 1857: 77 [Madeira]; Reitter, 1909: 297; Jakobson, 1911: 653;
Reichardt, 1941: 141,148, fig. 63c, 64c; Horion, 1949: 331; Halstead, 1963: 11; Witzgall,
1971: 164.
Acritus rhenanus Fuss, 1868: 356; Schmidt, 1885b: 324; Gangl bauer, 1899: 408, synonymized by
Sclunidt, 1895: 179.
Acritus shogunus Lewis, 1892b: 357 [Japan]; Reichardt, 1941: 141, 148, synonymized by Mazur,
1987: 31.

Japanese name: Kuro-yofushi-emma-mushi.

Distribution. Japan (Hokkaido); Madeira; North Africa; Central and South
Europe; Caucasus; USSR (Chabarovskij Kray).
6. 3. 9. SUBFAMILY SAPRININAE LACORDAlRE, 1854

Key to the Japanese genera of the subfamily Saprininae
1(2)
2(1)
3(4)
4(3)
5(8)

Head with frontal stria ........................................ Genus Gnathoncus Jacquelin-Duval, 1858
Head without frontal stria.
Surface of elytra impunctate. Prostemal keel strongly narrow ............................................. .
................................................................................. Genus Eopachylopus Lewis, 1926
Surface of elytra punctate posteriorly. Prosternal keel broad and flat at top.
Prostemal keel with a pair of small foveae (pre-apical prostemal foveae in Wenzel, 1962) on
apical 113. Body length, 2.0 - 4.0 mm.

214

6(7)
7(6)

8(5)

Frontal disk of head with 1 or 2 transverse rows or irregular transverse rugae; its anteriortransverse margin strongly impressed and straight ............ Genus Hypocaccus Thomson, 1867
Frontal disk of head usually with light punctation or weak rugae, without strongly transverse
rows; its anterior transverse margin lightly impressed, usually partly arcuate, and sometimes
interrupted .............................................................. Genus Hypocacculus Bickhardt, 1916
Prostemal keel without small foveae. Body length, 4.6 - 8.6 mm ...................................... .
.................................................................................... Genus Saprinus Erichson, 1834

GENUS GNATHONCUS JACQUEUN-DUYAL, 1858
Gnathoncus Jacquelin-Duval, 1858: 112 [type species: Hister rotundatus Kugelann, 1792: 304,
designated by Thomson, 1859: 75]; Thomson, 1867: 391; Reitter, 1896: 306; Ganglbauer,
1899: 378; Reitter, 1904: 35; 1909: 290; Jakobson, 1911: 641, 648; Bickhardt, 1916-17: 104;
Auzat, 1917: 206; Horion, 1936: 238; Reichardt, 1941: 154, 157; Stochmann, 1957: 67-76;
Arnett, 1962: 374,380; Halstead, 1963: 9, 11-13; Witzgall, 1971: 166; Kryzhanovskij andReichardt, 1976: 113; Yienna, 1980: 117; Mazur, 1973: 27; 1981: 87; 1984: 103; Yelamos,
1990: 78.

Key to the Japanese species of the genus Gnathoncus
1(2)
2(1)
3(4)

4(3)

Punctures of pygidium round. Apical area of prostemal process with a deep excavation.
.................................................................................... G. nannetensis (Marseul, 1862)
Punctures of pygidium transversely oblong. Apical area of prostemal process without
excavation.
Protibia with outer margin deeply emarginate between denticles. Sutural stria of elytra present
on basal 114. Posterior margin of 10th genital tergite of male genitalia truncate .................... .
.. ... .............. .............. ............ .......... ........ ...... ...... ......... G. rotundatus (Kugel ann, 1792)
Protibia with outer margin only feebly emarginate between denticles. Sutural stria of elytra
rudimentary on basal 116. Posterior margin of 10th genital tergite bifid ............................... .
...................................................................................... G. communis (Marseul, 1862)

Gnathoncus nannetensis (Marseul, 1862)(Fig. 130, 131,132,135)
Saprinus nannetensis Marseul, 1862: 499, fig. 2 [lye groupe]; Reitter, 1896: 308; Auzat, 1917: 208;
Reichardt, 1941: 160, 162; Hinton, 1945: 316, 325; Stockmann, 1957: 60, 73; Witzgall,
1971: 166.
Gnathoncusrotundatus: Thomson, 1862: 242necKugelann, 1792, corrected by Reichardt, 1941: 162;
Schmidt, 1885: 317; Ganglbauer, 1899: 379; Reitter, 1904: 36; 1909: 291; RecIaire et van derWiel, 1936: 234; Horion, 1949: 45.
Hister rotundatus var. "a": Hoffmann, 1803: 87, synonymiZed by Reichardt, 1941: 162.
Gnathoncus urganensis Reitter, 1896: 307; 1904: 36, synonymized by Bickhardt, 1916: 105.
Gnathoncus rotundatus var. suturalis: Ganglbauer, 1899: 380, synonymized by Bickhardt, 1916: 105.
Gnathoncus rotundatus var. nannetensis: Roubal et Labler, 1933: 30.
Gnathoncus nannetensis: Gemrninger et Harold, 1868: 792; Kryzhanovskij and Reichardt, 1976: 117;
Mazur, 1980: 91; 1984: 105; Yienna, 1980: 121; Hisamatsu, 1985b: 222; Yelamos, 1990: 79.

Japanese name: O-maru-mame-emma-mushi.

Description. Body length, PPL, male, 2.33 - 3.43 mm (2.73 ± 0.06, n=20),
female, 2.77 - 3.48 mm (3.10 ± 0.04, n=20), PEL, male, 2.21 - 3.31 mm(2.57 ± 0.06, n=20), female, 2.47 - 3.14 mm (2.89 ± 0.04, n=20). Width, male, 1.72
- 2.47 mm (2.04 ± 0.04, n=20), female, 2.13 - 2.50 mm (2.31 ± 0.02, n=20).
Biometric dala are given in Table 33. Body broadly oval and strongly convex. Cuticle
shining and black; antennae dark rufopiceous with club rufoteslaceous; legs dark
215

Table 33. Biometric data of Gnathoncus nannetensis (Marseul).
Female

Male
APW

PPW
PL
EL
EW
ProW
ProL

PyL
PTL
MSTL

MTIL

0.66-0.93 (0.77±0.01) 20
1.52-2.11 (1.78±0.03) 20
0.81-1.08 (0.91±0.01) 20
1.27-1.91 (1.58±0.03) 20
1.72-2.47 (2.04±0.04) 20
0.98-1.35 (1. 11±0.02) 20
0.17-0.25 (0.20±0.01) 20
0.74-1.13 (0.92±0.02) 20
0.59-0.78 (0.65±0.01) 20
0.61-0.78 (0.71±0.01) 20
0.71-0.93 (0.81±0.01) 20

0.81-0.93 (0.87±0.01) 20
1.86-2.21 (2.01±0.02) 20
0.88-1.15 (1.00±0.02) 20
1.57-1.94 (1.77±0.02) 20
2.13-2.50 (2.31±0.02) 20
1.10-1.47 (1.29±0.02) 20
0.17-0.29 (0.23±0.01) 20
0.86-1.13 (1.04±0.02) 20
0.64-0.81 (0.71±0.01) 20
0.69-0.86 (0.77±0.01) 20
0.78-1.03 (0.91±0.02) 20

reddish brown.
Front sparsely clothed with deep and moderate punctures.
Pronotal sides (Fig. 130A) feebly arcuate and convergent forward on basal 2/3,
thence strongly convergent apically. Apical angles obtuse. Marginal stria carinate andcomplete. Middle of disk moderately punctate, the punctures being separated by 2 - 4times their diameter, becoming coarser and denser towards sides; base with a single or
double transverse rows of punctures, which are nearly as coarse as those on sides.
Interspace among these punctures smooth and with occasional microscopic punctures.
Epipleura of elytra not concave. Marginal epipleural stria finely impressed andcomplete. Another epipleural stria complete, carinate and well impressed between themarginal epipleural and elytral striae. Marginal elytral stria complete, carinate and its
apical end extending across elytral apex to medio-apical angle of elytron, then a little
extending basally. Surface of epipleura between the marginal epipleural stria andanother epileural one smooth, and with several fine punctures medially; surface
between marginal elytral stria and another epipleural one with a row of coarse
punctures. Internal subhumeral stria present on medio-apical 114. Oblique humeral stria
present on basal 113 and usually crossed by a number of fine oblique rugae so that it
appears to be irregular. First dorsal stria deeply impressed and nearly complete. Second
and 4th striae deeply impressed and present on basal 112. Third stria a little longer thanthe 2nd. Fifth stria only represented by a short, curved, transverse line at base. Sutural
stria confined to basal region, usually present on basal 116. Surface of apical 112 - 3/5
of elytra covered with deep oval punctures, which are about as coarse as those on theextreme base of the pronotumand separated by about 1.5 (1 - 2) times the diameter; surface of basal 112 - 2/5 slightly
more finely and more sparsely punctate than middle of pronotal disk; extreme apex ofelytra (around apical stria) impunctate. Interspace among all discal elytral puncturessmooth or nearl y so.
Propygidium densely with coarse punctures on apical 112, which are as large as
those occurring anteriorly to the apical elytral stria, and more finely and more sparsely
punctate on basal 112. Interspace among the punctures with lightly impressed, fine andirregularly transverse lines, these lines heavily impressed on basal 112. Pygidium
coarsely and sparsely covered with round and umbilicate punctures, which are216

Fig. 130. Gnatlwncus spp. A - C: Pronotum and left elytron. D - F: Left protibia, dorsal view. A,
D: G. nannetensis (Marseul). E, E: G. rotundatus (Kugelann). C, F: G. communis(Marseul).

separated by about 1.5 (1 - 2) times the diameter; on apex those punctures are muchfiner.
Anterior margin of median portion of prosternum (Fig. 131A) round. Ventral
disk of keel even, somewhat broad, and sparsely and moderately punctate. Carina!
striae complete, carinate and sinuate, the anterior apex of striae attaining to a large
preapica! fovea. Descending lateral stria shortly present on media! 1/3 of prosternum,
its medio-apical end attaining to apical 113 of keel.
Anterior margin of mesosternum bisinuate with a feeble median projection.
Marginal stria of mesosternum complete and subcarinate. Disk sparsely and coarsely
217

:3
:3

Fig. 131. Gnathoncus spp. Prosternum, mesosternum, metasternum and 1st abdominal sternum. A:
G. nannetensis (Marseul). B: G. rotundatus (Kugelann). C: G. communis (Marseul).

covered with transverse and setiferous punctures, and interspace among punctures
sparsely with microscopic punctures. Meso-metasternal suture indistinctly impressed
and accompanied by a coarsely crenate line. Median area of intercoxal disk ofmetasternum sparsely clothed with fine and setiferous punctures, the punctures being
separated by 4 - 7 times their diameter; anterior comer sparsely with large andsetiferous punctures, the punctures being sparser and finer on mediolateral area;
posterior comer evenly covered with coarse punctures, which are separated by about
twice their diameter. Lateral metasternal stria carinate and deeply impressed on lateral
2/3 of the intercoxal disk, the area inside the stria bearing coarse punctures. Lateral disk
densely covered with large, round, shallow and setiferous punctures. Metepistemum
more densely covered with large punctures than those of lateral disk.
Intercoxal disk of 1st abdominal sternum completely striate, the stria carinate,running along post-coxal cavity on basal 112 and straight towards posterior margin on
apical 112. Surface of disk sparsely and coarsely punctate, the punctures being finer on
median area.
Protibia (Fig. 1300) with 9 spinules on outer margin.
Male genitalia as shown in Fig. 132.
218

Fig. 132. Gnathoncus nannetensis (Marseul) A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite and sternum, dorsal view. E: Apexof 8th sternum, ventral view. F: Eighth tergite and sternum, lateral view. G: Male genitalia, 9th
and 10th tergites, dorsal view. H: Ditto, lateral view. I: Ninth sternum (spicule), dorsal view. J:
Ditto, lateral view.
Specimens examined. [Hokkaido] 3 exs., Futatsu-yama, Shibeeha, Kushiro, 3/v/1985, K. Ijima(1J); 1 ex., Bekkambe-dai, Shibecha, Kushiro, 26/v/ 1971, K. Ijima (1J); 1 ex., Usubetsu, Taisetsu-eho,
22/vii/1954, K. Sawada (NA); 1 ex., Kuroishi-daira, Kami-Shihoro, 11/iv/1990, K. Haga; 1 ex.,
Nopporo, 1O/v/1987, M. Ohara; 59 exs., Ashoro, 30/v/1989, Y. Hayashi, N. Kuhara and O. Suda
(from a nest of an owl, Ketupa blakistoni).
[Honshu] Toehigi-ken: 1 ex., Miyuki, Utsunomiya-shi, 18/iii/1982, T. Kumazawa (TPM).
Kanagawa-ken: 6 exs., Bodai, Tanzawa, 8/iv/1973, Y. Hirano (NA); 3 exs., Ditto, 3/iii/1974, Y.
Hirano (NA); 1 ex., Hatano, Kanagawa, lIix/1978, H. Takizawa; 2 exs., Zukko-yama, Hatano,4/vil1989, A. Yamagami and K. Udagawa; 1 ex., Tanzawa-Oyama, 14/iv/1973, K. Masumoto.
Yamanashi-ken: 1 ex., Narusawa, Fuji, 311v/1983, M. Abe (NA). Aiehi-ken: 1 ex., Okehazama,
Owari, i/1946, M. Yasuda (NA). Osaka-fu: 2 exs., Nishinari-ku, 2/vii/1949, K. Sawada (NA). Tottoriken: 1 ex., Takenouehi, 4/iv/1987, H. Daido.
lKyftshu] Fukuoka-ken: 10"', Mt. Adachi, 25/v/1978, S. Nakao (NA); 1 ex., Matsunoe, Moji,
4/v/1936, S. Watanabe (NA). Kumamoto-ken: 1 ex., Uemura, Kuma-gun, 6/viI1986, K. Kimura(NA); 16 exs., Yokoshima, Tamana, 9/vi/1986, E. Matsui. Miyazaki-ken: 1 ex., Nobeoka, 7/iv/1982,
T. Kinoda; 12 exs., Furushiro-eho, Nobeoka, 4/vi/1985, T. Kinoda.
[Sakhalin] 1 ex., Muka, 20/vii/1944, Y. Nishijima (ElHU).

Distribution (Fig. 135). Japan (Hokkaido, Honshu, Shikoku, Kyushu); nearly
the whole Palearctic Region. New to Hokkaido.
Remarks. Gnathoncus nannetensis is a relatively large species; it is similar to219

G. rotundatus and G. communis, but distinguished by the different shape of thepunctures on the pygidium and by the presence of an apical excavation on theprosternal keel (see the key and description).
This cadavericole and coprophile species frequents a wide range of habitats; it
occurs on decomposing bodies of chicken, on dung of pig and cow, and so on. It isalso found in nests of birds, the fish owl Ketupa blakistoni (Seebohm) and theswallow Hirundo rustica Linnaeus, for example. Mazur (1981) recorded this species
from the nests of the following birds and animals: carrion craw, Corvus corax
Linnaeus; jackdaw, Corvus monedula (Linnaeus); starling, Sturnus vulgaris
Linnaeus; swift, Apus apus (Linnaeus); blue tit, Parus caeruleus Linnaeus; roughlegged buzzard, Buteo lagopus (Brunn); wood pigeon, Columba oenas Linnaeus;
badger, Meles meles Linnaeus, and Laetiporus sulphureus (Bull. et Fr.).
Gnathoncus rotundatus (Kugelann, 1792)(Fig. 130, 131, 133, 135)
Hister rotundatus Kugelann, 1792: 304.
Saprinus (Gnathoncus) rotundatus: Marseul, 1873: 221 [New to Japan; Nangasaki].
Hister nanus Scriba, 1790: 73, synonymized by Hoffmann, 1803: 87, the synonymy verified by
Mazur, 1984: 105.
Gnathoncus nanus: Reichardt, 1941: 191; Mazur, 1973: 27; Kryzhanovskij and Reichardt, 1976: 116;
Vienna, 1980: 120; Hisamatsu, 1985b: 222.
Gnathoncus rotundatus: Jacquelin-Duval, 1858: 112; Mazur, 1980: 88; 1984: 105.
For detailed synonymy, see Mazur (1984: 105).

Japanese name: Maru-mame-emma-mushi.

Description. Body length, PPL, 1.91 - 2.45 mm (2.15 ± 0.04, n=17), PEL,
1.84 - 2.35 mm (2.05 ± 0.04, n=17). Width, 1.42 - 1.76 mm (1.58 ± 0.02, n=17).
Biometric data are given in Table 34. Body broadly oval and feebly depressed
medially. Cuticle shining and black; antennae, mandibles, tibiae dark rufopiceous, andclub of antennae rufotestaceous.
Front sparsely covered with deep and moderate punctures, interspace among thepunctures sparsely clothed with fine punctures.
Pronotal sides (Fig. BOB) feebly arcuate and convergent forward on basal 3/4,
thence strongly convergent apically. Apical angles obtuse. Marginal stria feebly carinate
and complete. Disk sparsely and evenly covered with coarse, round and deeppunctures, which become coarser on the basal 1/2; interspace among the punctures
sparsely and finely punctate. Antescutellar area feebly depressed and coarsely andsomewhat densely punctate.
Epipleura of eiytra not concave. Marginal epipleural stria finely and clearly
impressed and complete. Another epipleural stria complete, carinate and clearly
impressed between marginal epipleural and elytral striae. Marginal elytral stria complete
and carinate, its basal end inwardly bent, and apical end extending across elytral apex
to medio-apical angle of elytron, then a little extending anteriorly. Surface of epipleuraand marginal elytral striae with a row of coarse punctures. Internal subhumeral stria
rudimentary on basal 113 and usually crossed by a number of fine, oblique rugae. First
dorsal stria (Fig. BOB) deeply impressed on basal 3/4. Second to 4th dorsal striae
220c

0.5 mm~52=··'=.~
Fig. 133. Gnathoncus roturuiatus (Kugel ann) A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite and sternum, dorsal view. E: Apexof 8th sternum, ventral view. F: Eighth tergite and sternum, lateral view. G: Male genitalia, 9th
and 10th tergites, dorsal view. H: Ditto, lateral view. I: Ninth sternum (spicule), dorsal view.

Table 34. Biometric data of Gnathoncus roturuiatus (Kugel ann) and G. communis (Marseul).

APW
PPW
PL
EL
EW

ProW
ProL
PyL
PTL
MSTL

MTTL

G. rotundatus

G. communis

0.51-0.74 (0.63± 0.01) 17
1.23-1.52 (l.36± 0.02) 17
0.64-0.83 (0.75± 0.01) 17
1.15-1.42 (1.30± 0.02) 17
1.42-l.76 (1.58± 0.02) 17
0.78-0.98 (0.90± 0.01) 17
0.15-0.25 (0.19± 0.01) 17
0.74-0.88 (0.79± 0.01) 17
0.49-0.56 (0.53±0.005) 17
0.54-0.64 (0.59± 0.01) 17
0.61-0.76 (0.69± 0.01) 17

0.64-0.88 (0.72±0.02) 10
1.35-1.62 (1.49±0.02) 10
0.74-0.91 (0.80±0.02) 10
1.23-1.64 (1.36±0.04) 10
1.47-1.89 (1.69±0.04) 10
0.81-1.05 (0.94±0.02) 10
0.15-0.25 (0.20±0.01) 10
0.64-0.93 (0.78±0.03) 10
0.47-0.64 (0.54±0.02) 10
0.54-0.69 (0.61±0.01) 10
0.66-0.78 (0.73±0.01) 10

221deeply impressed, crenate and present on basal 112, but usually a little extending
beyond the half. Fifth stria represented only by a short, curved, transverse line at base.
Sutural stria usually present on basal 114, sometimes shorter than basal 116. Surface ofapical 112 - 2/3 of elytra covered with longitudinal, oblong, deep punctures, which areabout as coarse as those of the extreme base of the pronotum and separated by 2 - 3times their diameter; surface of basal 112 - 113 finely and sparsely punctate, thepunctures being separated by 4 - 7 times their diameter; apical transverse region(around apical stria) densely and longitudinally rugate; extreme apex of elytra
impunctate. Interspace among discal elytral punctures smooth and rarely
microscopically punctate.
Propygidium densely covered with coarse and transverse oblong punctures on
apical 112, which are as large as those occurring posteriorly to apical elytral stria, andmore finely and sparsely punctate on basal 112. Interspace among the punctures withlightly impressed, fine and irregularly transverse lines, these lines being heavily
impressed on basal 112. Pygidium coarsely and densely covered with transverse oblong
punctures, the punctures being separated by their longer diameter to twice the diameterand their posterior margin usually emarginate; on apex those punctures are much finer.
Anterior margin of median portion of prosternum (Fig. BIB) round. Ventral
disk of keel even, broad and sparsely and finely punctate; the punctures being separated
by about 3 - 7 times their diameter; ground surface densely covered with microscopic
and longitudinal rugae except medially. Carinal stria complete, carinate and sinuate.
Keel without preapical fovea. Descending lateral stria short and present on median 1/3
of prosternum, its apical end attaining to medio-apical 113 of keel.
Anterior margin of mesosternum feebly bisinuate with a slight projection.
Marginal stria of mesosternum strongly carinate and complete. Disk sparsely covered
with moderate and round punctures which are separated by 4 - 6 times their diameter;interspace among the punctures sparsely and finely punctate. Meso-metasternal suturestrongly impressed and densely crenate, usually the lateral 116 reduced. Intercoxal diskof metasternum sparsely clothed with fine punctures which are separated by 5 - 10times their diameter and become coarser in the lateral and posterior areas along themarginal suture or lateral stria. Lateral metasternaI stria carinate and deeply impressedon lateral 3/4 of the intercoxal disk. Lateral disk densely covered with large, round andsetiferous punctures, which become denser posteriorly. Metepisternum more densely
covered with large punctures than on lateral disk.
Intercoxal disk of 1st abdominal sternum almost completely striate, the stria not
attaining the posterior margin, deeply impressed and carinate. Surface of disk evenly
covered with coarse longitudinal oblong and deep punctures; interspace among thepunctures sparsely clothed with microscopic punctures.
Protibia (Fig. 130E) with 8 denticles on outer margin, the interspace between theapical 3th and 4th denticles strongly emarginate.
Male genitalia as shown in Fig. 133.
Specimens examined. [Hokkaido] 1 ex., Futatsu-yama, Shibecba, Kushiro, 16/vl 1986, K. Ijima(IJ); 10 exs., Shimoboro, Tsurui, Kushiro moor, l/vi11992, M. Obara; 9 exs., Ditto, 9/vi/1992, M.
Obara; 1 ex., Kushiro Zoo, Yamabana, Kushiro, 9/vil 1992, M. Obara.
[Honsbu] Osaka-fu: 1 ex., Nisbinari, 15/v/1950, K. Sawada (NA).

Distribution (Fig. 135). Japan (Hokkaido, Honshu); nearly whole Holarctic
Region, Taiwan, South Africa.
Remarks. Gnathoncus rotundatus is quite similar to G. communis in all
222

external characters examined, and the differences are very subtle; the shape of the
protibia and the male genitalia should be used to distinguish these 2 species positively.
Gnathoncus rotundatus is coprophile and known from a wide range of habitats.
Mazur (1981) recorded this species from the nests of the following birds and mammals:
owl, Strix Linnaeus; hoopoe, Upupa epops Linnaeus; hooded-crow, Corvus corone
cornu Linnaeus; starling, Sturnus vulgaris Linnaeus; squirrel, Sciurus vulgaris
Linnaeus; rabbit, Oryctolagus cuniculus (Linnaeus), and hamster, Cricetus cricetus(Linnaeus).

Gnathoncus communis (Marseul, 1862)(Fig. 130, 131, 134, 135)
Saprinus communis Marseul, 1862: 501, synonymized with Gnatlwncus roturuiatus by Horn, 1873:
314, but revived by Mazur, 1990: 744.
Gnathoncus communis: Mazur, 1990,744 [Australia].
Gnathoncus schmidti Reitter, 1894: 239; Stockmann, 1957: 75, fig. 4, 9, 14, 19,24; Halstead, 1963:
13, fig. 27, 27b-d; Vienna, 1980: 123, fig. 45c, i, 46c, synonymized by Mazur, 1990: 744.
Gnatlwncus nidicola Joy, 1907: 133, tab., V, fig. 2; Auzat, 1917: 208; Reclaire et van der Wie1,
1936: 208, synonymized with G. schmidti by Stockmann, 1957: 73.
Gnatlwncus nidicola var. auzati Pic, 1918: 9.
Gnatlwncus punctor: Reichardt, 1941: 164, fig. 75A, 77B, nee Reitter, 1896:307; Hinton, 1945: 326,
fig. 32,34,36,41,42,45 [male genitalia], corrected by Stockmann,1957: 73.
Gnatlwncus idiopygus Casey, 1916: 255, synonymized by Mazur, 1990: 744.

Japanese name: Nise-maru-mame-ernrna-mushi.

Description. Body length, PPL, 2.21 - 2.57 mm (2.34 ± 0.04, n=10), PEL,
2.01 - 2.40 mm (2.19 ± 0.03, n=lO). Width, 1.47 - 1.89 mm (1.69 ± 0.04, n=lO).
Biometric data are given in Table 34. Body broadly oval and strongly convex. Cuticle
shining and black, but a few specimens very dark rufopiceous; antennae dark
rufopiceous with club rufotestaceous; legs dark reddish brown.
Front of head sparsely with fine and transverse punctures, and with round
punctures posteriorly.
Pronotal sides (Fig. 130C) feebly arcuate and convergent forward on basal 2/3,
thence feebly convergent apically. Apical angles obtuse. Marginal stria carinate andcomplete, its anterior portion weakly carinate and indistinct. Middle of disk finely
punctate, the punctures being separated by 3 - 5 times their diameter; towards sides the
punctures become coarser and denser; base sparsely and coarsely punctate, thepunctures being nearly as coarse as those of sides. Interspace among puncturessmoother and with occasional microscopic punctures.
Epipleura of elytra not concave. Marginal epipleural stria finely impressed andcomplete. Another epipleural stria complete and subcarinate between the marginal
epipleural and elytral striae. Marginal elytral stria complete and carinate; its apical end
extending across elytral apex to medio-apical angle of elytron and then a little extending
basally. Surface between marginal epipleural stria and another epipleural one with 2
rows of longitudinal-oblong and coarse punctures. Subhumeral stria present on median
113. Oblique humeral stria (Fig. 130C) present on basal 113, and usually crossed by anumber of fine, oblique rugae so that it appears to be irregular. First and 3rd dorsal
striae deeply impressed on basal 2/3 and carinate. Second and 4th dorsal striae a little
223

B

Fig. 134. Gnathoncus communis (Marseul). A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite and sternum, dorsal view. E: Apexof 8th sternum, ventral view. F: Eighth tergite and sternum, lateral view. G: Male genitalia, 9th
and 10th tergites, dorsal view. H: Ditto, lateral view. I: Ninth sternum (spicule), dorsal view.

shorter than the 3rd. Fifth and sutural striae only represented by a short, curved,
transverse line at base. Surface of apical 1/2 of elytra covered with coarse, ovalpunctures which are about as coarse as those of the extreme base of the pronotum andseparated by about 1.5 (1 - 2) times the diameter; surface of basal 112 slightly more
finely and distinctly more separately punctate than middle of pronotal disk; extreme
apex of elytra (around apical stria) impunctate. Interspace among all discal e1ytral
punctures smooth or nearly so.
Propygidium densely covered with coarse, round punctures on apical 112 which
are as large as those occurring anteriorly to apical elytral stria and finely and sparsely
punctate on basal 112. Interspace among punctures on basal 112 with lightly impressed,
fine and irregularly transverse lines. Pygidium densely and coarsely covered withtransverse-oblong and umbilicate punctures, which become denser apically and roundon the mediobasal region.
Anterior margin of median portion of prostemum (Fig. 131C) round. Ventral
disk of keel even, sparsely covered with moderate and longitudinal-oblong punctures.
Carinal striae complete, strongly carinate and sinuate, the anterior ends united with eachother in an arch. Preapical fovea absent. Descending lateral stria shortly present on
median 1/3, its apical end attaining to apical 113 of keel.

2247,,;/• Gnathoncus nannetensis
DGnathoncus rotundatus
V Gnathoncus communis
Fig. 135. Collection sites of Gnathoncus spp. in Japan.

Anterior margin of mesosternum bisinuate with a feeble median projection.
Marginal stria of mesosternum complete and subcarinate. Disk sparsely covered withlarge and round punctures, which are separated by 2 - 3 times their diameter. Mesometasternal suture slightly impressed and accompanied by a coarsely and sparsely
crenate line. Median area of intercoxal disk finely punctate, the punctures being
sepamted by 3 - 4 times their diameter; lateral and posterior 114 sparsely covered withlarge, round and setiferous punctures, these being a little larger than those of
mesosternal disk; interspace among large punctures sparsely clothed with finepunctures. Lateral metasternal stria strongly carinate, extending obliquely andposteriorly, and usually present on basal 2/3. Lateral disk densely covered with large,
round and setiferous punctures, which are somewhat larger than those of the intercoxal
disk. Metepisternum densely punctate, the punctures being as large as those of lateral
disk.
Intercoxal disk of 1st abdominal sternum completely striate. Surface of disk
sparsely and finely punctate on median area, and coarsely punctate laterally, thepunctures on lateral areas being smaller than the punctures of mesosternal disk.
Protibia (Fig. 130F) with 7 spinules on outer margin, the apical-outer angle
bearing 3 large spinules, and only feebly emarginate between spinules.
Male genitalia as shown in Fig. 134.
Specimens examined. [Honshu] Tokyo-to: 10 exs., Setagaya, 18/ix/1949, Ohira (NA).
[Kyfishfi] Kagoshima-ken: lei", Uenosono, Kagoshima, 2/xi/1982, T. & T. Nakane (NA).

Distribution (Fig. 135). Japan (HonshU, KyilshO); North America; Europe;
Caucasus; Central Asia; Sakhalin; Egypt; Australia. New to Japan.
Remarks. Gnathoncus communis is quite similar to G. rotundatus in all
external characters, but the 2 species can positively be distinguished by the shape of the
protibia and the male genitalia.
Little is known about the habitat of this species in Japan. Mazur (1981) recorded
G. communis in the nests of the following birds: carrion crow, Corvus corax
225Linnaeus; jackdaw, Corvus monedula (Linnaeus); tawny owl, Strix aluco Linnaeus
and goshawk, Accipiter gentilis (Linnaeus).
GENUS SAPRINUS ERICHSON, 1834
Saprinus Erichson, 1834: 172 [type species: Hister nitidulus Fabricius, 1801: 85, designated byWestwood, 1840: 22].
Plesioprinus Houlbert et Monnot, 1923: 72 (nom. nud.), synonymized by Cooman, 1947: 428.

Key to the Japanese species of the genus Saprinus
1(4)
2(3)

Basal 112 of interval between 1st and 4th dorsal elytral striae coarsely punctate.
Pygidium wholly punctate, without impunctate band on longitudinal mid line. Mesosternum
coarsely punctate....................................................... S. cyaneus auricollis Marseul, 1855
3(2) Pygidium with an impunctate band on longitudinal mid line. Mesosternum without coarse
punctures. ...................................................................... S. splendens (Paykull, 1811)
4(1)Basal1J2 of interval between 1st and 4th dorsal elytral striae smooth, at least finely punctate.
5(6) Third elytral dorsal stria short, usually 112 as long as 4th or 2nd stria. .. ............................. ..
............................................................................... S. planiusculus Motschulsky, 1849
6(5) Third elytral dorsal stria as long as 4th or 2nd stria ................... S. niponicus Dahlgren, 1962
Another species, Saprinus pecuinus, has been recorded from Japan.

SUBGENUS SAPRINUS ERICHSON, 1834

Saprinus (Saprinus) cyaneus auricollis Marseul, 1855
(Fig. 136, 137)
Saprinus auricollis Marseul, 1855: 390 [lIes Philippines]; Dahlgren 1971: 49; Nakane, 1977: 148[Ogasawara Isles].
Saprinus cyaneocupreus Marseul, 1864: 337, synonymized by Dahlgren, 1967: 220.

Japanese name: Ko-ruri-emma-mushi.

Description. Body length and biometric data are as follows: PPL 4.95, PEL
3.86, APW 1.33, PPW 3.47, PL 1.52, EL 2.28, EW 3.24, ProW 2.00, ProL 0.57,
PyL 1.52, PTL 1.19, MSTL 1.19, MTTL 1.62. Body broadly oval and strongly
convex. Cuticle strongly shining with purple metallic lustre on pronotum and darkgreen metallic lustre on elytra; funicle of antennae and tarsi dark rufopiceous; tibiae
black.
Frontal stria of head complete and carinate. Supraorbital stria complete andevenly arcuate. Disk of front sparsely covered with moderate punctures, which areseparated by 2 - 3 times their diameter; interspace among the punctures withmicroscopic punctures. Epistoma sparsely punctate, the punctures being a little coarser
than those of frontal disk. Labrum broadly depressed anteriorly.
Pronotal sides (Fig. 136) evenly arcuate and feebly convergent forward on basal
3/4, thence strongly arcuate and convergent apically. Apical angles obtusely angulate.
Marginal stria carinate and complete. Disk densely covered with large, round, and deeppunctures on longitudinal area along the sides, the punctures becoming finer on basal
114; interspace among the large punctures smooth, occasionally clothed withmicroscopic punctures; narrow lateral band along the side with dense and finepunctures; base with 2 or 3 rows of large, deep and longitudinal punctures, the rows
226

cyaneausauricollis
splendens

niponicus

planiusculus

2 mm I

Fig. 136. Saprinus spp., dorsal view of pronotum and left e1ytron, and pygidia.

absent on median fifth of posterior margin of pronotum, and becoming finer laterally;
elsewhere sparsely clothed with microscopic punctures.
Epipleura of elytra even; surface between epipleural and elytral marginal striae
smooth, shining, with a row consisting of sparse and fine punctures; finely punctate
inside marginal elytral stria, the punctures being separated by 2 - 3 times theirdiameter. Marginal epipleural stria lightly carinate and complete. Marginal elytrai stria
heavily carinate, complete and densely with coarse punctures; its apical end extending
inwards, reaching lateral 114 of posterior margin of elytron. External subhumeral stria
present on basal 114. Internal subhumeral stria nearly complete and densely crenate.
Oblique humeral stria present on basal 1/3 and its apical portion fused with the basal
227portion (nearly on basal 113 of elytron) of internal subhumeral stria, so that they appear
to be a single stria. First to 4th dorsal striae present on about basal 112 and densely
crenate with coarse punctures; 2nd a little longer than the 1st; 3rd a little shorter than the
1st; 4th as long as the 1st. Fifth dorsal stria absent. Sutural stria strongly carinate,sparsely crenate and abbreviated on basal 114. Surface of elytra, excepting flanks,
mediobasal area and extreme apices (around apical stria) densely covered with coarse
punctures, which are 112 as coarse as those of the longitudinal lateral area of the
pronotum, becoming finer apically, and separated by about 1.5 (1 - 2) times thedimeter; interspace among the punctures occasionally with fine punctures; flanks evenly
covered with fine punctures which are separated by 2 or 3 times their diameter;
mediobasal area, that is, basal 112 of elytra inside 4th dorsal stria sparsely clothed withmicroscopic punctures; area inside the apical stria densely covered with fine punctures.
Propygidium densely covered with coarse, round and shallow punctures, which
are as coarse as the coarse elytral punctures; interspace among the punctures withlightly impressed, fine and irregularly transverse alutaceous microsculptures, andsometimes with fine punctures. Pygidium (Fig. 136) sparsely with deep punctures,
which are as coarse as the propygidial ones; interspace among the punctures withalutaceous microsculpture basally, but elsewhere smooth and sometimes with finepunctures.

i\

C

C:J
( - ..

-~----~

I
10.5 mmFig. 137. Saprinus (Saprinus) cyaneus auricollis Marseul. A: Aedeagus, dorsal view. B: Ditto,
lateral view. C: Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite and sternum,
dorsal view. E: Apex of 8th sternum, ventral view. F: Eighth tergite and sternum, lateral view.
G: Male genitalia, 9th and 10th tergites, dorsal view. H: Ditto, lateral view. I: Ninth sternum(spicule), lateral view. J: Ditto, dorsal view.

228

Anterior margin of median portion of prosternum nearly straight. Ventral disk ofkeel even, but slightly descending on apical 116; surface of disk sparsely with finepunctures. Anterior marginal stria strongly carinate, nearly straight, but feebly andinwardly arcuate medially. Carinal striae strongly carinate and complete. Descending
lateral stria strongly carinate, its apical end attaining to apical 116 of keel.
Anterior margin of mesosternum deeply emarginate medially. Marginal stria
complete and strongly carinate. Disk sparsely with coarse punctures and other finepunctures intermingled among coarse ones. Meso-metasternal suture lightly impressed,
and with a line which is densely and coarsely crenate. Intercoxal disk of metasternum
sparsely clothed with microscopic punctures, these becoming coarse basally; atransverse apical band densely with coarse punctures. Lateral metasternal stria strongly
carinate, extending obliquely and posteriorly, and present on basal 3/4. Lateral disk
densely covered with large, round and shallow punctures, these progressively
becoming sparser and finer apically. Metepisternum broad, and densely covered withlarge, shallow and longitudinal punctures.
Intercoxal disk of 1st abdominal sternum sparsely with microscopic punctures
except along anterior margin and behind anterior comer, these parts being densely andcoarsely punctate; also with an apical transverse row consisting of moderate punctures.
Lateral stria of the disk strongly carinate basally and complete.
Protibia with 9 spinules on outer margin, the apical 3 and basal 2 small.
Male genitalia as shown in Fig. 137.
Specimens examined. [Ogasawara Isles] 1ci' , Bonin Is:, viii 1969 (NA).

Distribution. Japan (Ogasawara Isles); Philippines; Bali; New Guinea; Volcano
Is.; Buru Is.
Remarks. Saprinus cyaneus auricollis can easily be recognized by the color of
the body, which is purple on the pronotum and green on the elytra.
Saprinus (Saprinus) niponicus Dahlgren, 1962
(Fig. 136, 138, 141)
Saprinus niponicus Dahlgren, 1962: 245 [Japan: Nagano (holotype and a paratype); Hachinohe;
Aomori; Kiso-Fukushima; Hirayu; Gifu; Hiogo; Nagasaki]; Mazur, 1970: 52; 1984: 55;
Kryzhanovskij and Reichardt, 1976: 158; Hisamatsu, 1985b: 223.
Japanese name: Nise-dogane-emma-mushi.

Description. Body length, PPL, male, 4.05 - 5.71 mm (4.92 ± 0.09, n=20) ,female, 4.52 - 6.66 mm (5.89 ± 0.12, n=20), PEL, male, 3.33 - 4.81 mm (4.33 ±
0.08, n=20), female, 3.90 - 5.33 mm (4.82 ± 0.07, n=20). Width, male, 3.00 - 4.24
mm (3.80 ± 0.07, n=20), female, 3.38 - 4.71 mm (4.28 ± 0.07, n=20). Biometricdata are given in Table 35. Body broadly oval and strongly convex. Cuticle verystrongly shining with brassy lustre and black; tarsi and funicle of antennae dark
rufopiceous.
Frontal stria of head weakly carinate, complete, sometimes interrupted medially.
Supraorbital stria well impressed and complete. Disk of front sparsely covered withmoderate punctures, the punctures much sparser on basal 1/2; interspace among thesepunctures smooth or nearly so. Epistoma more densely punctate than on anterior areaof front. Labrum deeply depressed medially.
229

D.·• • ·.·.··,~=;·~·:;jB

0.5 mm I

Fig. 138. Saprinus (Saprinus) niponicus Dahlgren. A: Aedeagus, dorsal view. B: Ditto, lateral view.
C: Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite and sternum, dorsal view. E:
Apex of 8th sternum, ventral view. F: Eighth tergite and sternum, lateral view. G: Male
genitalia, 9th and 10th tergites, dorsal view. H: Ditto, lateral view. I: Ninth sternum (spicule),
dorsal view.

Pronotal sides (Fig. 136) evenly arcuate and convergent forward on basal 5/6,
thence strongly convergent apically. Apical angles obtuse. Marginal stria lightly
carinate and complete. Disk smooth, but densely, coarsely and deeply punctate along
the sides except on basal 114, sparsely clothed with fine punctures in a narrow bandoutside the coarsely punctate area; base with 1 or 2 irregular rows of large punctures,the rows absent on lateral 116 and the punctures sparse on median 113; elsewhere
extremely finely, sparsely punctate; area behind eyes feebly depressed.
Epipleura of elytra even; surface between elytral and epipleural marginal striae
with 2 rows consisting of moderate and sparse punctures; area inside elytral marginalstria sparsely and moderately punctate, the punctures being denser on apical 113.
Marginal epipleural stria complete and distinctly carinate. Marginal elytral stria strongly
carinate, area inside it being crenate with coarse punctures; its apical end extending
along the posterior margin of elytron to medio-apical angle of elytron and then a little
extending basally. External subhumeral stria confined basally. Internal subhumeral stria
present on median 113, but frequently abbreviated basally and sometimes absent.
Oblique humeral stria distinctly impressed on basal 113. First to 4th dorsal striae
densely crenate with coarse punctures, deeply impressed, and present on basal 2/3.
Fifth dorsal stria absent. Sutural stria shortly represented medially by a row of about 4- 7 coarse punctures. Surface of apical 113 of elytra covered with coarse, oval
230

punctures, which are finer than those on the longitudinal area along the pronotal sides,
and separated by about 1.5 (1 - 2) times the diameter, the punctured area usually
expanded basally to nearly sutural area; interval between 1st and 2nd dorsal striae
usually with several longitudinal and short lines; surface of basal 2/3 smooth, sparsely
clothed with microscopic punctures; extreme apex of elytra (around apical stria)
sparsely and finely punctate; interspace among all discal punctures smooth,
occasionally clothed with microscopic punctures.
Propygidium densely covered with coarse punctures, which are as coarse as
those of the apical 113 of the elytra but somewhat shallower, and become finer on thebasal 113; interspace among punctures occasionally and finely punctate, and with lightly
impressed, fine and irregularly alutaceous microsculptures. Pygidium (Fig. 136)
densely covered with punctures, which are a little coarser than the propygidial ones andprogressively become finer apically.
Anterior margin of median portion of prosternum straight. Ventral disk of keel
even, sparsely clothed with microscopic punctures, which are coarser laterally.
Anterior marginal stria of keel deeply impressed, carinate and complete. Carinal striae
complete, divergent anteriorly and posteriorly, and nearly straight on apical 1/3.
Descending lateral stria strongly carinate, its apical end attaining to apical 113 of carinal
stria.
Anterior margin of mesosternum broadly and feebly emarginate medially.
Marginal stria complete and strongly carinate. Disk sparsely clothed with coarse
punctures, sometimes the punctures sparser and finer medially. Meso-metasternal
suture lightly impressed, with a strongly crenate line. Intercoxal disk of metasternum
feebly convex, smooth, but with coarse and dense punctures along the lateral stria, andwith coarse and dense punctures in a transverse apical band. Lateral metasternal stria
strongly carinate, extending obliquely and posteriorly, and usually present on basal
112. Lateral metasternal disk densely covered with round, shallow, large and setiferous
punctures, which become smaller on the apical 113, with fine punctures intermingled.
Metepisternum more densely punctate than those of lateral disk of metasternum, thepunctures becoming sparser on apical 112.
Intercoxal disk of 1st abdominal sternum with strongly carinate lateral stria on
basal 2/3; sparsely clothed with fine punctures on medio-apical area except for a narrow
band along posterior margin with a row of coarse punctures; elsewhere densely andcoarsely punctate.
Protibia with 13 spines on outer margin, the apical 2 and basal 3 small.
Male genitalia as shown in Fig. 138.
Specimens examined. [Hokkaido] 3 exs., Futatsu-yama, Shibecha, Kushiro, lIviii/1954,
17/vi/1972, 151viii11987, K. Ijima (IJ); 1 ex., Shiriuchi, Oshima, lllviiil1976, T. Kumata et al.

(EIHU); 3 exs., Shibetsu, 241vii/1966, H. Takizawa; 2 exs., Kamui-kotan, near Asahikawa,
7/viii/1987, M. Ohara; 1 ex., Mt. Yilbari, 101viii/1966, H. Takizawa; 15 exs., Hokkaido Univ.,
Sapporo, 7, 27/vi/1985, M. Ohara; 8 exs., Misumai, Sapporo, 10 & 12/vii/1991, K. Sayama; 3 exs.,
Toyotaki, Sapporo, 3/ix/1987, M. Ohara; 1 ex., Hirafu side, alt. 350 m, Mt. Yotei, Kutchan,

201viii11993, M. Ohara; 3 exs., Toya-Naka-no-shima, Abuta, 29/vii-1Iviii/1993, S. Hori (HOR).
[Honshu] Aomori-ken: Paratype, 1d", Hachinohe, 8/vi11946, A. Fukuda (NA). Fukushima-ken:
1 ex., Yunohana, Tateiwa, Minami-Aizu, 8IviiI1948, Y. Kurosawa (NSMT). Gumma-ken: 1 ex.,
Sarugakyo, 6/vi/1966, M. Ishida. Tokyo-to: 2d"d" 1 Sf, Tokyo, 19/viii/1972, no collector's name; 2exs., Kunitachi, 211vi11963, S. Nomura (NSMT). Kanagawa-ken: 1 ex., Tozuka, Yokohama,
211vI1972, no collector's name. Yamanashi-ken: 1 ex., Mt. Daibosatsu, 221vii11963, T. Okumura; 1231

Table 35. Biometric data of Saprinus niponicus Dahlgren.
Female

Male
APW

PPW
PL

EL
EW

ProW
ProL
PyL
PTL
MSTL
MTTL

1.09-1.52 (1.35±0.02) 20
2.76-3.95 (3.51±0.07) 20
1.29-2.00 (1.64±0.04) 20
2.00-2.86 (2.60±0.05) 20
3.00-4.24 (3.80±0.07) 20
1.81-2.67 (2.34±0.05) 20
0.33-0.52 (0.45±0.01) 20
1.43-1.90 (1.68±0.03) 20
1.09-1.48 (1.35±0.02) 20
1.24-1.57 (1.42±0.02) 20
1.33-1.90 (1.67±0.03) 20

1.24-1.67 (1.49±0.02) 20
2.95-4.33 (3.88±0.07) 20
1.33-2.14 (1.83±0.04) 20
2.38-3.05 (2.87±0.04) 20
3.38-4.71 (4.28±0.07) 20
2.00-3.09 (2.58±0.05) 20
0.43-0.62 (0.48±0.01) 20
1.43-2.33 (2.09±0.04) 20
1.00-1.67 (1.45±0.03) 20
1.05-1.71 (1.50±0.03) 20
1.14-1.95 (1.79±0.04) 20

ex., Sagashiro Spa., 28-30/vii/1961, S. Nomura (NSMT). Nagano-ken: 1d" , Fukushima, Kiso, 1112/vi/1946, T. Nakane (NA); 2 exs., Kami-Kachi, Shinano, 21-28/vii/1947, S. Osawa (NA); 4 exs.,
Koma-ga-take, Kiso, 4, 5/viii/1946, S. Osawa (NA). Gifu-ken: Paratypes, Id" 1 ~, Hirayu, Hida,
24/vii/1946, M. Hayashi (NA).
[Kyilshil] Miyazaki-ken: 2 exs., Koiwaya, Kitagawa-cha, 14/v/1993, T. Kinoda.

Distribution (Fig. 141). Japan (Hokkaido, HonshU, Shikoku, Kyushu);Primorskij Kray; Korea.
Remarks. Saprinus niponicus resembles S. planiusculus; however, the long
3rd dorsal elytral stria and the shape of the 8th sternum of the male genitalia of S.
niponicus will distinguish it from S. planiusculus.
This species is cadavericolous.
Saprinus (Saprinus) pecuinus Marseul, 1855
Saprinus pecuinus Marseu!, 1855: 391; 1873: 221 [Kagoshima, Kiu-Siu, Japon]; Mazur, 1984: 56.

Specimens examined. No material of this species has been available.

Distribution. China; Japan (?).
Remarks. Marseul's record (1873) remains in doubt. I have had no opportunity
to check his specimens.
Saprinus (Saprinus) ploniusculus Motschulsky, 1849
(Fig. 136, 139, 141)
Saprinus planiusculus Motschulsky, 1849: 97 [Kirgiz SSR]; Mazur, 1984: 56.
Saprinus cuspidatus Ihssen, 1949: 183, synonymized by Kryzhanovskij, 1972: 435.

Japanese name: Dagane-emma-mushi.

Description. Body length, PPL, male, 3.95 - 5.66 mm (4.72 ± 0.08, n=20),
female, 4.09 - 6.33 mm (5.30 ± 0.12, n=20), PEL, male, 3.52 - 4.71 mm(4.20 ± 0.06, n=20), female, 3.81 - 5.09 mm (4.47 ± 0.08, n=20). Width, male,
3.14 - 4.19 mm (3.64 ± 0.05, n=20), female, 3.28 - 4.57 mm (3.94 ± 0.07, n=20).
Biometric data are given in Table 36. Body broadly oval and strongly convex. Cuticle
2320.5 mmI

Fig.139. Saprinus planiusculus Motschulsky. A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
Apex of aedeagus, caudal view. D: Male genitalia, 8th tergite and sternum, dorsal view. E:
Ditto, lateral view. F: Apex of 8th sternum, ventral view. G: Male genitalia, 9th and 10th
tergites, dorsal view. H: Ditto, lateral view. I: Ninth sternum (spicule), dorsal view.

very strongly shining with brassy lustre, black; tarsi and funicle of antennae dark
rufopiceous.
Frontal stria of head weakly carinate and complete. Supraorbital stria wellimpressed and complete. Disk of front densely covered with moderate punctures,
which become sparser on the basal 112; interspace among these punctures usuallysmooth, occasionally clothed with fine punctures. Epistoma media-apically more
densely punctate than the disk. Labrum deeply depressed medially.
Pronotal sides (Fig. 136) evenly arcuate and convergent forward on basal 5/6,
thence strongly convergent apically. Apical angles obtusely angulate. Marginal stria
carinate and complete. Disk smooth, but densely, coarsely and deeply punctate along
the sides except on basal 114, sparsely clothed with fine punctures in a narrow bandoutside the coarsely punctate area; base with 2 or 3 irregular rows of large punctures
except on lateral 7th and a narrow antescutellar area; elsewhere extremely finely,sparsely punctate; area behind eyes feebly depressed.
Epipleura of elytra sparsely with moderate punctures, which become denser on
the apical 113. Marginal epipleural stria complete and finely impressed. Marginal elytral
stria lightly carinate and complete, its apical end extending along the posterior marginof elytron to media-apical angle of elytron and then bending basally and running for ashort distance. External subhumeral stria confined basally. Internal subhumeral stria
present on median 1/3, with coarse punctures, and sometimes its basal end united withthe apical end of oblique humeral stria. Oblique humeral stria (Fig. 136) distinctly
impressed on basal 113. First, 2nd and 4th dorsal striae present on about basal 112, andcoarsely and sparsely crenate; 2nd usually a little longer than the 1st. Third dorsal stria
233

short, present on basal 5th, and coarsely crenate. Fifth dorsal stria wanting. Sutura!
stria shortly represented by a row of 4 - 5 coarse punctures medially. Surface of apical
112 or 113 of elytra covered with coarse and oval punctures, which are finer than thosein the area along the pronotal sides, and usually separated by about 1.5 (1 - 2) times
the diameter, but sometimes sparser; the punctate area usually expanding basally onto
sutural area and also onto interval between 2nd and 4th dorsa! striae; interval between
1st and 2nd dorsal striae usually with several longitudinal and short lines; surface ofbasal 112 smooth, sparsely clothed with microscopic punctures; extreme apex of elytra(around apical stria) sparsely and finely punctate; interspace among all discal puncturesof elytra smooth, occasionally clothed with microscopic punctures.
Propygidium densely covered with coarse punctures, which are a little coarser
than the elytral coarse punctures and become finer on basal 112; interspace among thepunctures occasionally finely punctate and with lightly impressed, fine and irregularly
alutaceous microsculptures. Pygidium densely covered with punctures, which areslightly coarser than those of the propygidium and progressively become finer on theapical 113 and nearly absent at the apex.
Anterior margin of median portion of prosternum straight. Ventral disk of keel
even, sparsely clothed with microscopic punctures, which are coarser laterally.
Anterior marginal stria of keel deeply impressed, carinate, and complete. Carinal stria
complete, divergent anteriorly and posteriorly, and nearly straight on apical 1/3.
Descending lateral stria strongly carinate, its apical end attaining to apical 113 of carina!
stria.
Anterior margin of mesosternum broadly and feebly emarginate medially.
Marginal stria complete, and strongly carinate. Disk sparsely clothed with microscopic
punctures, which become coarser laterally. Meso-metasternal suture lightly impressed,
and accompanied with a strongly crenate line. Intercoxal disk of metasternum feebly
convex, sparsely clothed with microscopic punctures, which are coarser along lateralmetasternal stria and in a transverse apical band. Latera! metasternal stria strongly
carinate, extending obliquely and posteriorly, and present usually on basal 112. Lateralmetasternal disk densely covered with round, shallow, large and setiferous punctures,
which become smaller on the apical 113, with fine punctures intermingled.
Metepisternum more densely punctate than those of lateral disk of metasternum, thepunctures becoming sparser on apical 112.
Intercoxal disk of 1st abdominal sternum with strongly carinate lateral stria on
basal 3/4; sparsely and microscopically punctate on median area; lateral area along
lateral stria sparsely and coarsely punctate, with fine punctures intermingled.
Protibia with 13 spines on outer margin, the apical 2 and basal 3 small.
Male genitalia as shown in Fig. 139.
Specimens examined. [Hokkaid6] 1 ex., Shibecha, Kushiro, 16/vi/1955, K. Ijima (D); 1 ex.,
Kamui-kotan, Asahikawa, 20/viil 1987, M. Ohara; 10", Ombetsu, 25/vil 1985, M. Ohara; 1 ex., o taru ,Shiribeshi, I/viil1943, T. Hasegawa; 3 exs., Ishikari-hama, 7/viii/1978, T. Fujisawa; 2 exs.,
Zenibako, 27/vi/1985, 19/vil 1985, M. Ohara; 5 exs., Sapporo, 19, 27/vi/1985, 9/vii11987, M. Ohara;
1 ex., Maru-yama, Sapporo, 23/vi/1979, N. Nishikawa; 1 ex., Ditto, 2/vi/1951, T. Kumata (ElHU); 1
ex., 16zankei, Sapporo, 8/vii11907, S. Matsumura (ElHU); 1 ex., Misumai, Sapporo, 27/vi11990, M.
Ohara; 18 exs., Ditto, 9,10 & 18/vii/1991, K. Sayama; 13 exs., Nopporo, 12/vi/1986, M. Ohara; 9exs., Kamayausu, Sapporo, 20/vii/1987, M. Hinakura; 10" 1 i?- , Hakodate, 19/vi/1954, T. Nakane(Saprinus cuspidatus, Dahlgren det.) (NA); 1 ex., Hakodate, 17/vi/1954, T. Nakane (NA); 1 ex., Mt.
Hakodate, 26/viii11951, K. Honma (NA); 2 exs., Yagishiri Is., 211vi/1985, Y. Kusui.
[Honshu] Iwate-ken: 1 ex., Yashikidai, 19/vii/1954, T. Kumata (ElHU). Yamagata-ken: 12234

exs., Yonezawa, 16/v/1944, Y. Kurosawa (NSMf). Fukushima-ken: 1 ex., Wakamatsu, 29/vi/1949,
T. Nakane (NA); 2 exs., Mt. Koma-ga-take, southern Aizu, 10/viii/1949, K. Nagayama (NSMT); 2
exs., Wakamatsu, 14/v/1947, Y. Kurosawa (NSMT); 1 ex., Fukushima-shi, 9/vii11965, K. Tazoe.
Tochigi-ken: 1 ex., Kawaraya, Utsunomiya, 23/vi/1981, K. Aida, collected from a dead crow (TPM); 6
exs., Tsuruta-numa, Utsunomiya-shi, 22-23/vii/1991, no collector's name (TPM). Chiba-ken: 2 exs.,
Cape Futtu, 2/vii/1961, M. Yamazaki (NA); 1 ex., Oami, 19/vi11984, M. Ohara; 2 exs., Ichikawa,
27/iv11958, M. Suwa. Tokyo-to: 2 exs., Ogikubo, 16/vi/1963, 211iv11968, M. Ishida; 1 ex.,
Kumegawa, Higashi-mura-yama, 26/iv/1964, M. Ishida; 1 ex., Setagaya, 18/ix/1949, Ohira (NA); 1
ex., Takamizu-yama, 2/vi/1935, T. Nakane (NA); 1 ~ ,lex., Hongo, Musashi, v/1947, T. Nakane(Saprinus cuspidatus, Dahlgren det.) (NA); 2 exs., Aoyama, 26/viil 1943,25/vl 1948, T. Nakane (NA);
1 ex., Komazawa, 23/v/2603 (= 1943), Watanabe (NSMf); 1 ex., Kinuta, 16/v/1965, Y. Kurosawa(NSMf); 1 ex., Ueno-koen, lIvii/1934, H. Uchida (NSMf); 1 ex., Hatagaya, 111v/1924, nocollector's name. Yamanashi-ken: 1 ex., Masutomi, 12/vi/2603 (= 1943), no collector's name
(NSMf); 1 ex., Sagashiro Spa., 28-30/vii/1961, S. Nomura (NSMT). Niigata-ken: 1 ex., Niigata-shi,
26/vi/1933, K. Baba. Nagano-ken: 1 ex, Fukushima, Kiso, 11-12/vi/l940, T. Nakane (Saprinus
cuspidatus, Dahlgren det.) (NA); 1 ex., Kami-Kochi, Shinano, 21-28/vii/1947, S. Osawa (NA); 3
exs., Sunama, Nojiri, 211viii/1943, T. Nakane (NA); 1 ex., Gaijin-mura, Nojiri, 19/viii/1941, T.
Nakane (NA). Aichi-ken: 3 exs., Obata, Owari, 15/vi/1941, 30/v/1947, S. Osawa (NA); 1 ex., Miya,
Mikawa, 1-2/ix/1947, S. Osawa (NA); 1 ex., Shiadeki-machi, Nagoya, lIvi/1941, S. Osawa (NA).
Gifu-ken: 3 exs., Hirayu, 24/vii/1946, M. Hayashi. Hyogo-ken: 1 ex., Motoyama-mura, near Kobe,
14/vl 1938, K. Sakaguchi (NA); 1 ex., Kobe, 22/vi/1930, no collector's name (BSM). Tottori-ken: 2exs., Daisen, 27/v/1935, M. Hamano (NSMf).
[Kyfishu] Fukuoka-ken: 1 ex., Hakozaki, 24/vil 1946, F. Nakasuji (NA).

Distribution (Fig. 141). Japan (Hokkaid6, Honshu, Shikoku, Kyushu); northern
Palearctic Region; Europe; Siberia; China; Korea; Canary and Azores Is.; North Africa;
Asia Minor; Viet-Nam (?)
Remarks. Saprinus planiusculus resembles Saprinus niponicus, with which it
is sympatric to a great extent; however, it can be distinguished from the latter by theshortened 3rd dorsal elytral stria and the shape of the 8th sternum of the male genitalia.
This species occurs on dead animal bodies.
Marseul (1873: 221) recorded Saprinus nitidulus from "Japan; Shimabara (KiuSiu) et Hiogo (Niphon)". Although I have not examined Marseul's specimens, they
may be planiusculus or niponicus.

Table 36. Biometric data of Saprinus planiusculus Motschulsky.
Male
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Female1.19-1.43 (1.32±0.01) 20
2.81-3.81 (3.36±0.05) 20
1.33-1.86 (1.58±0.03) 20
2.28-2.86 (2.57±0.03) 20
3.14-4.19 (3.64±0.05) 20
2.05-3.05 (2.29±0.04) 20
0.48-0.71 (0.58±0.02) 20
1.29-1.81 (1.54±0.03) 20
1.14-1.52 (1.34±0.02) 20
1.29-1.76 (1.46±0.02) 20
1.48-2.00 (1.68±0.03) 20

235

1.24-1.57 (1.42±0.02) 20
3.'05-4.00 (3.61±0.06) 20
1.38-1.86 (1.68±0.03) 20
2.24-3.19 (2.70±0.06) 20
3.28-4.57 (3.94±0.07) 20
2.00-3.05 (2.42±0.05) 20
0.43-0.62 (0.54±0.01) 20
1.52-2.14 (1.81±0.03) 20
1.14-1.67 (1.37±0.03) 20
1.19-1.67 (1.42±0.03) 20
1.38-1.95 (1.68±0.04) 20

Saprinus (Saprinus) splendens (PaykulI, 1811)(Fig. 136,140,141)
Hister splendens Paykull, 1811: 53 [Capland].
Hister elegans Paykull, 1811; 57, synonymized by G. Muller, 1938: 165.
Hister speciosus: Dejean, 1821: 48 [nom. nud.].
Saprinus speciosus Erichson, 1834: 170; Marseul, 1873: 221 [Japan; Nangasaki et Hiogo],synonymized by Dahlgren, 1967: 214.
Hister cyaneus: Boisduval, 1835: 147, correctly identified by Mazur, 1976: 717.
Hister capicola Dejean, 1837: 141 [nom. nud.], synonymized by Gemminger and Harold, 1868: 791.
Saprinus viridicupreus Blanchard, 1853: 57, synonymized by Blackburn, 1903: 107.
Saprinus advena Marseul, 1855: 335, synonymized by Gemminger and Harold, 1868: 789.
Saprinus ovalis Marseul, 1855: 382, synonymized by Dahlgren, 1967: 214.
Saprinus elegans: Marseul, 1855: 383.
Saprinus ater MacLeay, 1864: 118, synonymized by Blackburn, 1903: 104.
Saprinus rasselas Marseul, 1855: 379, synonymized by Bickhardt, 1921: 117.
Saprinus splendens elegans: G. Muller, 1938: 165.

Japanese name: Ruri-emma-mushi.

Description. Body length, PPL, male, 5.05 - 6.85 mm (6.12 ± 0.13, n=20),
female, 6.14 - 7.66 mm (7.07 ± 0.10, n=20), PEL, male, 4.28 - 5.43 mm(4.88 ± 0.07, n=20), female, 4.86 - 5.90 mm (5.52 ± 0.07, n=20). Width, male, 3.76
- 4.76 mm (4.33 ± 0.06, n=20), female, 4.33 - 5.24 mm (4.80 ± 0.06, n=20).
Biometric data are given in Table 37. Body broadly oval and strongly convex. Cuticle
very strongly shining with dark blue metallic lustre; elytra with light blue metallic
lustre; funicle of antennae and tarsi dark rufopiceous; tibiae black.

F

0.5 mill,

Fig. 140. Saprinus splendens (paykull). A: Aedeagus, dorsal view. B: Ditto, lateral view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Apex of 8th sternum, ventral view. E: Eighth
tergite and sternum, lateral view. F: Male genitalia, 9th and 10th tergites, dorsal view. G: Ditto,
lateral view. H: Ninth sternum (spicule), dorsal view.
236

Frontal stria of head carinate, complete, its sides straight. Supraorbital stria
complete and well impressed. Disk of front moderately punctate, the punctures being
separated by 0.5 - 1 times the diameter on apical half, becoming sparser and finer onbasal 1/2 and coarser laterally; sometimes with short lines medially, which aredivergent apically, and with a deep and large puncture on basal 113 on mid line.
Epistoma more sparsely punctate than the disk medio-apically. Labrum broadly anddeeply depressed medially.
Pronotal sides (Fig. 136) evenly arcuate and convergent forward on basal 5/6,
thence strongly convergent apically. Apical angles obtuse. Marginal stria lightly
carinate and abbreviated on basal 1/6. Disk smooth, densely, largely and deeply
punctate along the sides, the punctate area narrowest at middle, attaining to basal
margin, and the punctures much coarser behind anterior comer; base with a row ofcoarse punctures, but the punctate area broadly interrupted on median 1/3 of pronotal
posterior margin; elsewhere microscopically, sparsely punctate; area behind eyes
strongly depressed.
Epipleura of elytra even; surface between epipleural and elytral marginal striae
with 3 or 4 rows of moderate punctures, the rows being often abbreviated on basal 1/3;
area inside elytral marginal stria sparsely clothed with microscopic punctures, which
are finer on the apical 1/3. Marginal epipleural stria lightly but distinctly carinate andcomplete. Marginal elytral stria strongly carinate, complete, its apical end extending
along the posterior margin of elytron and united with the apical end of sutural stria.
External subhumeral stria (Fig. 136) confined to a basal area. Internal subhumeral stria
absent. Oblique humeral stria well impressed and present on basal 112. First to 3rd
dorsal striae densely crenate with coarse punctures, strongly impressed and present on
about basal 1/2; 2nd a little longer than the 3rd; 1st a little longer than the 2nd. Fourth
dorsal stria short, usually present on mediobasal 114. Fifth dorsal stria absent. Sutural
stria distinctly carinate, sparsely crenate and abbreviated on basal 116. Surface of elytra
densely covered with large, round, and deep punctures, which are as coarse as those of
the extreme base of the pronotum, except on flanks, mediobasal 112, and extreme apex
(around apical stria); elsewhere sparsely and microscopically punctate; interval between
1st and 2nd dorsal striae with several longitudinal rugae.
Propygidium short; disk densely covered with punctures, which are a littlecoarser than the elytral coarse punctures and sparser and finer on the basal 1/2.
Pygidium (Fig. 136) densely covered with large punctures, which are about 3 times ascoarse as the elytral coarse punctures but finer on the apical 1/3; usually with acomplete or nearly complete, narrow, median longitudinal line which is impunctate.
Anterior margin of median portion of prosternum nearly straight and shortly
marginate on median 113. Ventral disk of keel descending anteriorly and even on basal
114; its disk finely covered with punctures, which are separated by 2 - 3 times theirdiameter; interspace among these punctures with fine and alutaceous microsculpture.
Anterior marginal stria of keel strongly carinate and complete. Carinal stria weakly
carinate and strongly divergent posteriorly. Descending lateral stria complete andstrongly carinate, its apical end attaining to apical 1/6 of keel.
Anterior margin of mesosternum slightly emarginate medially, its marginal stria
well carinate and complete. Disk sparsely clothed with fine punctures, which are a littlecoarser laterally. Meso-metasternal suture well impressed, complete, angulate at
middle, and without a crenate line. Intercoxal disk of metasternum feebly convex, but
strongly depressed along longitudinal sutural line in male, while feebly depressed in237

Table 37. Biometric data of Saprinus splendens (paykull).
Male
APW
PPW
PL
EL

EW
ProW
ProL
PyL
PTL
MSTL

MTTL

Female1.24-1.62 (1.49± 0.02) 20
3.47-4.83 (4.01± 0.06) 20
1.71-2.09 (1.91± 0.02) 20
2.38-3.24 (2.90± 0.05) 20
3.76-4.76 (4.33± 0.06) 20
2.19-2.95 (2.67± 0.05) 20
0.29-0.62 (0.49± 0.02) 20
1.67-2.19 (1.93± 0.03) 20
1.29-1.81 (1.54± 0.03) 20
1.33-1.81 (1.64± 0.03) 20
1.57-2.09 (1.87± 0.03) 20

1.38-1.76 (1.60± 0.02) 20
4.05-4.81 (4.46± 0.05) 20
1.81-2.33 (2.16± 0.03) 20
2.86-3.47 (3.19± 0.04) 20
4.33-5.24 (4.80± 0.06) 20
2.62-3.24 (2.96± 0.04) 20
0.38-0.62 (0.48± 0.01) 20
2.00-2.62 (2.26± 0.03) 20
1.43-1.81 (1.64± 0.03) 20
1.43-2.00 (1.71± 0.03) 20
1.76-2.19 (2.03± 0.02) 20

female; median area sparsely clothed with fine punctures, which are separated by 3 - 8times their diameter; lateral area along the lateral stria sparsely covered with coarse,
round and deep punctures, and with other fine ones intermingled; similarly punctate inan apical transverse band, the punctures becoming sparser and finer on median 1/3;
interspace among punctures on lateral area and apical transverse band with finely
impressed, alutaceous micro-sculpture. Lateral metasternal stria strongly carinate,
extending obliquely and posteriorly, and present on basal 2/3. Lateral metasternal disk
densely covered with large, round, deep punctures, and with fine ones intermingled;
interspace among punctures with lightly impressed, irregular, and transverse alutaceous
microsculpture. Metepisternum densely covered with large, deep, and longitudinalpunctures, and with fine punctures intermingled; interspace among these puncturessmooth and shining.
Intercoxal disk of 1st abdominal sternum sparsely clothed with fine punctures,
which become coarser laterally; lateral stria well carinate and abbreviated on apical 1/6.
Protibia with 11 spinules on outer margin, the apical 3 and basal 2 small.
Male genitalia as shown in Fig. 140.
Description of larva: Hayashi (1986: pI. 11).
Specimens examined. [Honshu] Yamagata-ken: 2 exs., Sakata, 27/vii11956, K. Shirahata (NA).
Chiba-ken: 1 ex., Oami, 22/vi/1984, M. Ohara; 2 exs., Okitsu, 1913, F. Muir (BSM). Tokyo-to: 1ex., Inogashira, 2/viiil 1952, no collector's name. Kanagawa-ken: 1 ex., Kamigo, Yokohama, viiI1977,
K. Masumoto; 1 ex., Yokohama, vi/1923, J. F. Illingworth (BSM); 2 exs., Hiratsuka, vii/1958, S.
Nomura (NSMT). Niigata-ken: 2 exs., Senami, northern Echigo, 29/vii/1984, viii/1984, K. Baba.
Shizuoka-ken: 17 exs., Toi, Tagata, vi-ix/1987, S. Mochizuki; 1 ex., Gotemba, 17/viii/1933, K.
Nakamura; 1 ex., Miho, 10/vi/1932, no collector's name (NSMT); 1 ex., Ditto, 20/x/1927, K. Sato
(NSMT). Aichi-ken: 2 exs., Obata, Owari, 20, 30/v/1947, S. Osawa (NA); 1 ex., Nagoya, Owari,
1/viii11947, S. Osawa (NA); 1 ex., Miya, Mikawa, 1-2/ix/1947, S. Osawa (NA). Hyogo-ken: 2 exs.,
Uzaki, 20/vi/1935, S. Eda (NA); 1 ex., Kobe, 23/vi/1930, no collector's name (BSM). Wakayama-ken:
1 ex., Kawayu, Hongu, southern Kii, 24/vii/1946, S. Osawa (NA); 7 exs., Kushimoto, 30/vi/1957,
C. Morishima (NSMT). Osaka-fu: 1 ex., Mino, 2/viii1l947, F. Takahashi (NA); 1 ex., Senriyama,
28/v1l949, K. Sawada (NA). Okayama-ken: 1 ex., Kamba, 19/vIl940, T. Okutani (NSMT).
[Kyushu] Fukuoka-ken: 1 ex., Tatara, 10/v/1945, Y. Kurosawa; 2 exs., Ditto (NSMT); 1 ex.,
Hakozaki, 15/v/1964, F. Nakasuji (NA). Saga-ken: 1 ex., Chika, 30/ix/1952, H. Yamaguchi (NA).
Kagoshima-ken: 1 ex., Kushikino, 28/v/1982, M. Maegata; 1 ex., Midori-ga-oka, Kagoshima-shi,
9/vi/1982, N. Mizoguchi; 1 ex., Sakura-jima, 20/vii/1951, F. Takahashi (NA).
[Nansei Isles] Yaku-shimaIs.: 1 ex., 7-13/vii/1945, R. Matsuda (NA); 1 ex., Kosugi-dani, 7-238

10/vi/1965, T. Kumata (ElHU); 1 ex., Shiratani, 211vi/1975, Y. Kurosawa (NSMT). Amami-Oshima
Is.: 1 ex., Naze, 19/v/1960, T. Shibata (NSMT). Ishigaki Is.: 2 exs., Ishigaki, 26/iii/1984, Yoshida.

Distribution (Fig. 141). Japan (Hokkaido, HonshU, Shikoku, KyOshO, Nansei
Isles); tropical Africa; Arabia; Kashmir; Afghanistan, Oriental Region; Australia.
Remarks. Saprinus splendens can easily be recognized by the metallic blue bodyand the pygidium with an impunctate area on the mid line.
The biology of S. splendens, including the immature stages, was described byNakasuji (1965) and Mochizuki (1980, 1985).

/
Saprinus splendens//

Fig. 141. Collection sites of Saprinus spp. in Japan.

239

':<

/"""'/ .

GENUS HYPOCACCULUS BICKHARDT, 1916

Hypocacculus Bickhardt, 1916-17: 82,95 [type species: Saprinus metallescens Erichson, 1834: 192,
originally designated]; Reichardt, 1926b: 14; 1932: 28, 96; 1941: 156, 280; Peyerimhoff,1936: 228; Kryzhanovskij and Reichardt, 1976: 201; Mazur, 1984: 83.
SUBGENUS NESSUS REICHARDT, 1932

Nessus Reichardt, 1932: 32 [type species: Saprinus rubripes Erichson, 1834: 193. originallydesignated]; 1941: 293; Kryzhanovskij and Reichardt, 1976: 207; Mazur, 1984: 86.

Hypocacculus (Nessus) asticus (Lewis, 1911)(Fig. 142, 143, 144)
Hypocaccus asticus Lewis, 1911: 89 [Japan: Enoshima).
Hypocacculus asticus: Bickhardt, 1916: 96.
Hypocacculus (Nessus) asticus: Mazur, 1984: 83.
Japanese name: Hime-hamabe-emma-mushi.

Description. Body length, PPL, 2.16 - 2.39 mm (2.26 ± 0.06, n=3); PEL, 1.83
- 2.06 mm (1.95 ± 0.05, n=3). Width, 1.60 - 1.80 mm (1.70 ± 0.05, n=3). Body
oval and strongly convex. Biometric data are given in Table 38. Cuticle very strongly
shining and black or nearly so with a strong bronze or brassy lustre, but in an observed

.'.'

.-

::.,.".

tj:)C;;;;. . .•:
',','

3
3

Fig. 142. Hypocacclus (Nessus) asticus Lewis. A: Pronotum and elytra, the right elytron withnormal striation and the left with strongly reduced striation. B: Prosternum and meso- andmetasterna. C: Head.

240

!

0.5 mmFig. 143. Hypocacclus (Nessus) asticus Lewis. A: Aedeagus, dorsal view. B: Ditto, lateral view. C:
Male genitalia, 9th and 10th tergites, dorsal view. D: Ditto, lateral view. E: Male genitalia, 8th
tergite and sternum, dorsal view. F: Ditto, lateral view. G: Ninth sternum (spicule), dorsal
view.

specimen dark rufopiceous, lacking the metallic lustre; antennae dark rufopiceous withclub rufotestaceous; legs dark reddish brown.
Frontal stria of head (Fig. 142C) complete and strongly carinate, straightanteriorly, and its sides parallel to each other and straight. Disk of front wholly withirregular and shallow rugae as shown in Fig. 142C. Epistoma scabrous.
Pronotal sides (Fig. 142A) feebly arcuate and not strongly convergent forward
on basal 5/6, thence strongly arcuate and convergent apically. Apical angles round.
Disk densely covered with coarse and round punctures except on mediobasal area, thepunctures becoming deeper on longitudinal lateral area along the sides; base with 2 or 3
rows of coarse punctures, occasionally with fine punctures intermingled; sparsely
clothed with microscopic punctures narrowly along the sides and on mediobasal areas.
Epipleura of elytra smooth, sparsely clothed with microscopic punctures.
Marginal epipleural stria complete and distinctly carinate. Marginal elytral stria strongly
carinate, complete, and extending along posterior margin of elytra, its apical end
attaining to near lateral 113 of elytron. External subhumeral stria present on basal 114.
Internal subhumeral stria present on median 113 and densely with moderate punctures.
Oblique humeral stria distinctly impressed and present on basal 113. First and 2nd
dorsal striae present on basal 2/3; the basal end of the 2nd extending inwards. Third
dorsal stria a little shorter than the 2nd. Fourth dorsal stria present on basal 112, its
basal end united with the end of sutural stria in an arch. First - 4th dorsal striae densely
crenate with moderate punctures. Sutural stria complete and sparsely crenate withmoderate punctures. Surface of apical 112 of elytra densely covered with coarse, round,and shallow punctures, which are as coarse as the pronotal coarse punctures and241separated by about 1.5 (1 - 2) times the dimeter; basal 112, flanks and extreme apex ofelytra impunctate.
Propygidium densely covered with coarse punctures, which are a little coarser
than the elytral punctures and become finer anteriorly; interspace among punctures withlightly impressed, transverse alutaceous microsculpture on basal 112. Pygidium
sparsely covered with coarse and transverse punctures, which are a little finer thanthose of the propygidium and separated by 2 - 3 times their diameter.
Anterior margin of median portion of prosternum (Fig. 142B) round. Preapicalfoveae distinct and large. Keel moderately compressed, its apex narrowly truncate.
Descending lateral striae strongly carinate and convergent to apex. Carinal striae nearlyparallel and slightly divergent basally on apical 112 and strongly so on basal 114. Lateralside of keel coarsely punctate laterally, and with alutaceous microsculpture.
Anterior margin of mesosternum (Fig. 142B) feebly emarginate, its marginalstria distinctly carinate and shortly interrupted medially. Disk very sparsely andmoderately punctate. Meso-metasternal suture lightly impressed, and sparsely carinate
with large punctures. Intercoxal disk of metasternum sparsely clothed with microscopic
punctures, which are occasionally coarser medio-apically. Lateral stria strongly
carinate, extending obliquely and posteriorly, and reaching near hind coxae. Lateraldisk sparsely covered with coarse, round, shallow and setiferous punctures, which aretwice as coarse as the propygidial punctures, separated by their own diameter, andbecome sparser and finer apically. Metepistemum coarsely punctate on basal 2/3 andsmooth on apical 113.
Intercoxal disk of 1st abdominal sternum (Fig. 142B) completely striate laterally,and sparsely and microscopically punctate except for a transverse apical band which ismoderately punctate.
Protibia with 7 spines on outer margin, the apical 4 large, and strongly dentate.
Male genitalia as shown in Fig. 143.
Specimens examined. [Honshu] Kanagawa-ken: 3cl'd" 1!f, 1 ex., Yui-ga-hama, Kamakura,
22/iv/1988, M. Tao; Wakayama-ken: 1 ex., Wakayama-shi, 10/ii/1957, M. Yoshida (NA).

[Kyushu] Nagasaki-ken: 1 ex., Maehama, Kazusa-chO, 28/iii/1979, S. Imasaka.

Distribution (Fig. 144). Japan (Honshu, Shikoku, Kyushu).
Remarks. Hypocacculus asticus can easily be recognized by the rugae on thefront, the rather small size, the striation of the elytron and the punctation of the
pronotum and elytra.

Hypocacculus asticus~/

Fig. 144. Collection sites of Hypocacculus (Nessus) asticus Lewis in Japan.
242

Table 38. Biometric data of Hypocacculus asticus (Lewis) and Hypocaccus lewisii (Schmidt).

APW
PPW
PL
EL
EW
ProW
ProL
PYL
PTL
MSTL
MTTL

Hypocacculus asticus

Hypocaccus lewisii

0.69-0.72 (0.71±0.01) 3
1.47-1.64 (1.56±0.04) 3
0.69-0.82 (0.74±0.03) 3
1.18-1.24 (1.20±0.02) 3
1.60-1.80 (1.70±0.05) 3
0.98-1.01 (0.99±0.01) 3
0.23-0.33 (0.28±0.02) 3
0.95-1.19 (1.06±0.06) 3
0.59-0.65 (0.63±0.02) 3
0.56-0.62 (0.60±0.02) 3
0.65-0.75 (0.71±0.02) 3

0.75-1.01 (0.89±0.01) 20
1.57-2.13 (1.88±0.03) 20
0.75-1.08 (0.93±0.02) 20
1.24-1.80 (1.56±0.03) 20
1.67-2.39 (2.07±0.04) 20
0.98-1.37 (1.22±0.02) 20
0.23-0.43 (0.32±0.01) 20
0.86-1.43 (1.24±0.03) 20
0.62-0.98 (0.74±0.02) 20
0.65-0.88 (0.74±0.02) 20
0.72-0.95 (0.88±0.02) 20

According to Ohara and Imasaka (1994), this species occurs on the root ofClystegia soldanella Roem. et Shult.

GENUS HYPOCACCUS THOMSON, 1867
Hypocaccus Thomson, 1867: 400 [type species: Hister quadristriatus Hoffmann, 1803: 85, nec
Thunberg, 1794: 65, designated by Lewis, 1899: 3J; Schmidt, 1885b: 302; Ganglbauer, 1899:
382; Lewis, 1899: 3, fig. 6, 7; Reitter, 1909: 291; Bickhardt, 1916-17: 82-98; Reichardt,
1926b: 14; 1941: 156,305; Peyerimhoff, 1936: 226; Arnett, 1962: 374, 380; Dahlgren, 1969:
64-66; Kryzhanovskij and Reichardt, 1976: 217; Mazur, 1984: 92.
Rhytidoprinus Houlbert et Monuot, 1923: 46, synonymized by Cooman, 1947: 428 [type species notdesignated].

Key to the Japanese species of the genus Hypocaccus
1(2)Pronotum smooth, at least finely punctate laterally. Meso-metasternal suture without crenate
line............................................................... Subgenus Baeckmaninolus Reichardt, 1926.......................................................................................
H. varians (Schmidt, 1890)
2(1) Pronotum coarsely punctate. Meso-metasternal suture with strongly crenate line................... .
.......................................................................... Subgenus Hypocaccus Thomson, 1867
3(4) Pygidium with impunctate area, which is divided by coarsely punctate mid line..................... .
....................................................................................... H. axeli Kryzhanovskij, 1976
4(3) Pygidium evenly punctate wholly, or the punctures becoming finer apically, mid line without
coarse punctures.
5(6) Frontal disk of head densely covered with coarse rugae. Mesosternal marginal stria interrupted at
middle. .. ....... ...... ....... ........ .................. ............................ H. subaenus (Schmidt, 1890)
6(5) Frontal disk of head with 1 or 2 transverse rows. Mesosternal marginal stria complete.
7(10) Surface of elytra smooth on basal 112.
8(9) Sutural elytral stria united with marginal elytral stria. ................... H. sinae (Schmidt, 1890)
9(8) Sutural elytral stria not united with marginal stria ............... H. akanensis M. Ohara, sp. nov.
10(7) Surface of elytra coarsely punctate on basal 1/2. ......................... H. lewisii (Schmidt, 1890)
Another species, H. ainu, has been described from Japan.

243

SUBGENUS HYPOCACCUS THOMSON, 1867

Hypocaccus (Hypocaccus) ainu Lewis, 1899
Hypocaccus ainu Lewis, 1899: 24 [Japan: Yezo, Ishikari river]; Reichardt, 1941: 308,322.
Hypocaccus (? Hypocaccus) ainu: Kryzhanovskij and Reichardt, 1976: 229.

Japanese name: Ainu-hamabe-emma-mushi.

o
00

,0

OfggooDooo

ooooooooo~oo 00°

00
00000000°0°0°0°0 0 0 0 0

Fig. 145. Hypocaccus spp. Pronotum and left elytron. A: H. axeli Kryzhanovskij. B: H. subaenus(Schmidt). C: H. sinae (Marseul). D: H. lewisii (Schmidt).

244

Specimens examined. No material of this species has been available for my study.

Distribution. Japan (Hokkaido).
Hypocaccus (Hypocaccus) axeli Kryzhanovskij, 1976
(Fig. 145, 146, 147, 148, 149)
Hypocaccus (Hypocaccus) axeli Kryzhanovskij in Kryzhanovskij and Reichardt, 1976:[Primorskij Kray; Japan: Harima (= Hyogo»).

223

Japanese name: Arame-hamabe-emma-mushi.

Description. Body length and biometric data as follows; PPL 3.68, PEL 3.36,
APW 0.99, PW 2.57, PL 1.15, EL 2.11, EW 2.76, ProW 1.65, ProL 0.66, PyL
1.12, PTL 0.95, MSTL 0.99, MTTL 1.15. Biometric data are given in Table 38. Body
broadly oval and moderately convex. Cuticle shining and black or bronze with brassy
lustre; antennae dark piceous with club black; legs dark brown.
Frontal stria of head (Fig. 147A) nearly complete, sometimes interrupted behindbases of antennae. Anterior area of frontal disk with 1 or 2 rugae which are arcuate andsometimes irregularly interrupted. Supraorbital stria complete. Epistoma clearly striate
on lateral sides, the stria feebly carinate.
Pronotal sides (Fig. 145A) feebly arcuate and convergent on basal 4/5, thence
straight and strongly convergent apically. Apical angles obtuse. Marginal stria strongly
carinate and complete. Disk densely and coarsely punctate except on a mediobasal
triangular area, a small mediolateral area and a narrow band inside the marginal stria,
the coarse punctures not rugose behind anterior comer; base with 2 or 3 irregular rowsof coarse punctures, the rows broadest on antescutellar area; elsewhere smooth.
Epipleura of elytra even, and sparsely and finely punctate. Marginal epipleuralstria clearly and strongly carinate, and densely with coarse punctures, its apical end
extending inwards, reaching to lateral 113 of posterior margin of elytron. Internal
subhumeral stria deeply impressed, strongly crenate and present on median 113.
Oblique humeral stria present on basal 113. First dorsal stria deeply impressed, andnearly complete, its apical portion sinuate. Second dorsal stria present on basal 2/3.
Third and 4th dorsal striae present on basal 112; basal end of 4th united with the basal
end of sutural stria. First to 4th dorsal striae densely crenate with coarse punctures.
Sutural stria deeply impressed and complete. Surface of elytra densely and coarsely
punctate except on flanks, extreme apex of elytra, a narrow band between sutural
striae, and a medio-basal area inside 4th dorsal striae and occupying basal 113 of elytra;
the punctures as coarse as those of the extreme base of the pronotum and separated by
0.5 times their diameter, usually appearing to be longitudinal rugae; elsewhere sparsely
clothed with microscopic punctures.
Propygidium evenly covered with coarse punctures, which are a little coarser
than the elytral ones and separated by about their own diameter; interspace among thepunctures occasionally finely punctate. Pygidium (Fig. 148E) densely covered withcoarse, round and deep punctures, which are a little coarser than the propygidial ones,
along lateral margin and on mid line on apical 113 of pygidium, with a heart-shaped
impunctate area on apical 113.
Anterior margin of median portion of prostemum (Fig. 146A) obtuse-acute.

245

Fig. 146. Hypocaccus spp. Prosternum, mesosternum, metasternum and 1st abdominal sternum. A:
H. axeli Kryzhanovskij. B: H. subaenus (Schmidt). C: H. sinae (Marseul). D: H. lewisii(Schmidt).

Preapical foveae present. Keel moderately compressed, its anterior apex narrowly
truncate. Descending lateral stria strongly carinate and convergent to apex. Carinal stria
present on basal 3/4 and distinctly impressed, its basal 112 gradually divergent basally.
Lateral side coarsely punctate medially, and with alutaceous ground microsculptures.
Anterior margin of mesosternum feebly emarginate medially, its marginal stria
heavily carinate and complete. Disk smooth. Meso-metasternal suture finely impressed,
and with sparsely and coarsely crenate line. Intercoxal disk of metasternum smooth
except in a transverse apical band, which is sparsely and finely punctate. Lateralmetasternal stria well impressed, extending obliquely and posteriorly, reaching near246

hind coxae. Lateral disk covered with large, shallow and round punctures, which aretwice as coarse as pygidial punctures, separated by 0.5 times their diameter and become
sparser apically; interspace among the punctures with fine punctures and alutaceous
microsculpture. Metepisternum densely covered with coarse punctures, which are alittle finer than those of the lateral disk of the metasternum; interspace among thepunctures with moderate punctures.
Intercoxal disk of 1st abdominal sternum completely striate on each side, andmoderately punctate along the lateral stria, behind anterior corner and in a transverse
apical band.

~

A~
... ,

c~

Fig. 147. Hypocaccus spp. Head (A - D) and pygidium (E). A, E: H. axeli Kryzhanovskij. B: H.
subaenus (Schmidt). C: H. sinae (Marseul). D: H. lewisii (Schmidt) .

.........
,0.5 mm.......... '" : ·0"·: ..

...

D,0.5 mmFig. 148. Hypocaccus (Hypocaccus) axeli Kryzhanovskij. A: Aedeagus, dorsal view. B: Ditto,
lateral view. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E:
Male genitalia, 9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: Ninth sternum(spicule), dorsal view.
247Protibia with 9 spinules on outer margin, the apical 2 small.
Male genitalia as shown in Fig. 148.
Specimens examined. (Hokkaido] 1d", Akan, Kushiro, 5/viii/1918, M. Suzuki, (ELKU).
(Honshu] Aomori-ken: 1d", Akaishi River, Ajigasawa-machi, 7/vi/1987, H. Sawada. Naganoken: 1d", Kami-Kochi, Pv. Shinano, 27/iii/1918, M. Suzuki (ELKU). Gifu-ken: 1d", Nagara, Pv.
Mino, 19/vii/1919, M. Suzuki (ELKU).

Distribution (Fig. 152). Japan (Hokkaid6, Honshu). New to Hokkaid6.
Remarks. Hypocaccus axeli is easily recognized by the rather large size, thepygidium with an impunctate area medially, and the punctation of the pronotum and theelytra.
Little is known about the habitat of this species. It apparently lives on the pebbled
beach of mountain rivers.
Hypocaccus (Hypocaccus) lewisii (Schmidt, 1890)(Fig. 145, 146, 147, 149, 152)
Saprinus lewisii Schmidt, 1890: 53 [Japan: Hakodate).
Hypocaccus lewisii: Reichardt, 1926b: 273; 1941: 308,320, fig. 167.
Hypocaccus (Hypocaccus) lewisii: Kryzhanovskij and Reichardt, 1976: 226.

Japanese name: Karakane-hamabe-emma-mushi.

Description. Body length, PPL, 2.26 - 3.17 mm (2.81 ± 0.05, n=20), PEL,
2.13 - 2.94 mm (2.55 ± 0.04, n=20). Width, 1.67 - 2.39 mm (2.07 ± 0.04, n=20).
Body oval and moderately convex. Biometric data are given in Table 38. Cuticle
shining and black or nearly so with a strong bronzy, blue or brassy lustre; antennae
dark rufopiceous with club rufotestaceous; legs dark reddish brown.
Frontal stria of head (Fig. 1470) complete, well impressed and straight on eachside. Supraorbital stria well impressed. Disk of front with a transverse ruga, andusually densely and strongly rugose in central area, and impunctate posteriorly.
Epistoma scabrous.
Pronotal side (Fig. 1450) feebly arcuate and not strongly convergent on basal
5/6, thence strongly arcuate and convergent apically. Marginal stria complete andsubcarinate. Disk coarsely and densely punctate, the punctures becoming more rugose
laterally, and impunctate mediobasally; basal margin of pronotum with 2 or 3 rows ofcoarse punctures, and with fine ones intermingled, the coarse punctures becoming
larger in antescutellar area and lateromedian 115 of pronotum, and finer laterally.
Epipleura of elytra smooth and even. Marginal epipleural stria complete, finely
impressed. Marginal elytral stria (Fig. 1450) well impressed, carinate, and complete,
its apical end continuing across elytral apex and united with the apical end of sutural
stria at medio-apical angle of elytron. External subhumeral stria absent. Internal
subhumeral stria shortly present on medial 113. Oblique humeral stria well impressedon basal 113. First dorsal stria unclearly present on basal 112, usually consisting ofcoarse punctures in a row, close to and parallel with the oblique humeral stria. Second,
3rd and 4th dorsal striae indistinguishable from elytral punctures, the 2nd sometimes
represented by a ruga on basal 112; the basal end united with the 4th stria in an arch.
Sutural stria deeply impressed and complete, the apical end united with marginal elytral
stria; the basal end united with the 4th stria, in an arch. Surface of elytra densely
covered with coarse and somewhat longitudinal punctures except on mediobasal area,248

i,- - -- - ----

./:>
Q:J
_/

; - - n - ___ n _ n--§j-----

s;::;:: ~;:__________ n

.

~_-

c
o=~
=-~
B

'

Fig. 149. Hypocaccus (Hypocaccus) lewisii (Schmidt). A: Aedeagus, dorsal view. B: Ditto, lateralview. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male
genitalia, 9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: Ninth sternum (spicule),
dorsal view.

basal 116 of interstice between 2nd and 3rd dorsal striae and flanks of elytra; elsewheresparsely covered with microscopic punctures.
Propygidium densely and moderately punctate, the punctures becoming finer andsparser basally. Pygidium densely and moderately punctate, the punctures finer near
apex.
Anterior margin of median portion of prosternum (Fig. 1460) weakly obtuseangulate. Preapical foveae distinctly present. Keel somewhat broad, moderately
compressed, its anterior apex acute, rarely truncate. Descending lateral striae strongly
carinate and convergent apically. Carinal striae present on basal 2/3, parallel to eachother on median 113, and divergent basally, rarely their apical ends divergent, attaining
to the descending lateral striae. Lateral sides of keel sparsely with moderate punctures.
Anterior margin of mesosternum feebly emarginate, its marginal stria carinate andcomplete. Disk smooth, sparsely with microscopic punctures. Meso-metasternal suture
lightly impressed, accompanied by a coarsely crenated parallel line. Intercoxal disk ofmetasternum smooth except in a transverse apical band, which is sparsely and finely
punctate. Lateral metasternal stria well impressed, extending obliquely and posteriorly,
and reaching near hind coxae. Lateral disk evenly covered with coarse and setiferous
punctures, impunctate along the lateral metasternal stria. Metepisternum densely andcoarsel y punctate.
Intercoxal disk of 1st abdominal sternum completely striate on each side, andsparsely covered with fine punctures along anterior margin.
Protibia with 10 spinules on outer margin, its apical-outer angles bearing 3 large
spinules.
Male genitalia as shown in Fig. 149.
Specimens examined. [Hokkaido] 1 ex., Obira, Rumoi, 28/vii/1985, M. Ohara; 1 ex., Shari249

ch6, 14/x/1987, H. Torikura; 2 ex., Zenibako, Otaru, 30/v/1977, 23/vi/1977, N. Nishikawa; 1 ex.,
Ishikari, 24/vi/1987, A. Iwasaki; 1 ex., Raisatsu, Ishikari, 24/vii-7/viii/1993, S. Hori (HOR); 7 exs.,
Shiranuka, 25/vi/1985, M. Ohara; 12 exs., Toyokita-kaigan, Urahoro, 24/vi-7/vii/1993, S. Hori
(HOR); 1 ex., Bansei-kaigan, Taiki, 7-8/vii/1993, S. Hori (HOR); 2 exs., Koj6-hama, Shiraoi,
30/vii/1953, M. Otake; 2 exs., Yokoma, Suttu, ll/vii/1979, T. Fujisawa (BlHU); 1 ex., Abashiri,l/viii/1954, T. Nakane (NA).; 1 ex., Hakodate, 17/vii/1954, T. Nakane (NA); 1 ex., Ditto,
15/vii/1952, K. Honma (BlHU); 1 ex., Nanae-hama, Hakodate, 17/viii/1977, Y. Kusui; 1 ex., Ditto,
5/iv/1978, N. Nishikawa; 1 ex., Nezaki-ch6, Hakodate-shi, 15/v/1978, H. Sawada; 1 ex., Ditto,
19/vii/1952, K. Honma (EIHU). Okushiri Is.: 3 exs., Aonae, 28/vi/1986, Y. Kusui.
[Honshu] Aomori-ken: 1 ex., Nakanosawa, Mutsu, 8/vi/1989, H. Saito. Fukushima-ken: 2exs., Fujima-kaigan, Iwaki, 25/vi/1988, K. Tazoe.
[Kyfishu] Fukuoka-ken: 1 ex., Aino-shima, 22/v/1983, S. Nakao.
[?] 1 ex., Imahama, 23/v/1948, no collector's name.

Distribution (Fig. 152). Japan (Hokkaid6, HonshO, KyOshO). New to KyOshO.
Remarks. Hypocaccus lewisii resembles H. axeli, but differs by the rather
dense punctation on the pronotum and the pygidium without an impunctate area.
Hypocaccus lewisii occurs mainly on sandy beaches. It is occasionally foundalso on the sandy beach of mountain rivers (Kurosawa, 1988).
Hypocaccus (Hypocaccus) sinae (Marseul, 1862)(Fig. 145, 146, 147, 150, 152)
Saprinus sinae Marseul, 1862: 496, fig. 52 [China: Shang-Hail; 1873: 221 [Japon, Hiogo];Blackburne, 1903: 106; Jakobson, 1911: 651.
Hypocaccus sinae: Desbordes, 1919: 415; Reichardt, 1941: 307, 312, fig. 158.
Hypocaccus (Hypocaccus) sinae: Kryzhanovskij and Reichardt, 1976: 226.

QJL:_-_-_--~::::_:~-_~-_-_-_ -_n:~~
--A~.
_,
<S)1C
.
..

..

~:----:-.::-:::
.. ..

_

_---,

'

B

\,

C

'.

"',

.1.· .

o(. .).. <.. .
. ".:" ... .

0.5 mm,G·····

Fig. 150. Hypocaccus (Hypocaccus) sinae (Marseul). A: Aedeagus, dorsal view. B: Ditto, lateralview. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male
genitalia, 9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: Ninth sternum (spicule),
dorsal view.
250

Japanese name: Nise-hamabe-emma-mushi.

Description. Body length, PPL, 2.39 - 3.34 mm (2.90 ± 0.06, n=20), PEL,
2.22 - 2.94 mm (2.58 ± 0.05, n=20). Width, 1.86 - 2.45 mm (2.15 ± 0.03, n=20).
Biometric data are given in Table 39. Body oval and moderately convex. Cuticle verystrongly shining and black or nearly so with strong bronzy or brassy lustre; antennae
dark rufopiceous with club rufotestaceous; legs dark reddish brown.
Frontal stria of head (Fig. 147C) carinate, complete and straight on each side.
Supraorbital stria well impressed. Disk of front impunctate, with 2 or 3 transverse
rugae, which are sometimes interrupted. Epistoma scabrous.
Pronotal sides (Fig. 145C) rather feebly arcuate and not strongly convergent on
basal 4/5, thence strongly arcuate and convergent apically. Apical angles somewhat
round. Marginal stria complete and carinate. Disk smooth, its sides strongly, densely
and moderately punctate anteriorly, the punctate area sometimes becoming broader
basally and the punctures coarser and more rugose in the region of the anterior angles.
Basal margin of pronotum with 1 or 2 rows of large punctures, which, however,
become finer laterally.
Epipleura of elytra impunctate and even. Marginal epipleural stria complete.
Marginal elytral stria deeply impressed and complete, its apical end extending along theposterior margin of elytron and united with the apical end of sutural stria at medioapical angle of elytron. External subhumeral stria absent. Internal subhumeral stria
shortly present on medial 113. Oblique humeral stria well impressed on basal 113 - 112,
close to and parallel with the 1st dorsal stria. First dorsal stria deeply impressed,
crenate and present on basal 2/3. Second, 3rd and 4th dorsal striae well impressed andcoarsely punctate; these striae a little longer than the 1st. Sutural stria represented by afine, crenulate, complete line, the apical end united with marginal elytral stria and thebasal end united with the 4th in an arch. Surface of elytra smooth, impunctate on basal
112, coarsely and densely punctate on apical 112, the punctures being separated by 0.5- 1 times the diameter; extreme apex of elytra (around apical stria) and flanks
impunctate.
Propygidium densely and moderately punctate, the punctures being separated byslightly more than their own diameter. Pygidium sparsely and moderately punctate, thepunctures becoming denser towards basal angles, and finer and sparser apically; apexof pygidium impunctate.
Anterior margin of median portion of prostemum (Fig. 146C) round. Preapicalfoveae distinctly present. Keel moderately compressed, its anterior apex narrowly
truncate. Descending lateral striae carinate and convergent to apex. Carinal striae
usually present on basal 3/4, and usually parallel apically and divergent basally,sometimes their apical ends united with each other at middle of keel. Lateral side of keel
sparsely (sometimes densely) covered with coarse punctures on apical 112, with finepunctures intermingled.
Anterior margin of mesosternum feebly emarginate, its marginal stria carinate andcomplete. Disk smooth, sparsely clothed with microscopic punctures. Mesometasternal suture finely impressed, accompanied with a coarsely crenated parallel line.
Intercoxal disk of metasternum smooth except in a transverse apical band, which issparsely and coarsely punctate. Lateral metasternal stria well impressed, extending
obliquely and posteriorly, reaching near hind coxa. Lateral disk sparsely covered withlarge punctures, with fine ones intermingled. Disk of metepisternum densely covered
251

Table 39. Biometric data of Hypocaccus sinae (Marseul) and H. subaenus (Schmidt).

APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MITL

H. sinae

H. subaenus

0.75-0.98 (0.86±0.01) 20
1.57-2.22 (1.95±0.04) 20
0.85-1.14 (1.04±0.02) 20
1.31-1.83 (1.54±0.03) 20
1.86-2.45 (2.15±0.03) 20
1.14-1.57 (1.33±0.02) 20
0.33-0.46 (0.39±0.01) 20
1.14-1.62 (1.30±0.03) 20
0.65-0.88 (0.76±0.01) 20
0.62-0.92 (0.74±0.02) 20
0.69-0.98 (0.82±0.02) 20

0.62-0.78 (0.71±0.01) 20
1.50-1.96 (1.69±0.02) 20
0.75-0.95 (0.85±0.01) 20
1.28-1.70 (1.42±0.02) 20
1.70-2.22 (1.92±0.03) 20
0.92-1.41 (1.13±0.03) 20
0.26-0.39 (0.31±0.01) 20
1.00-1.33 (1.14±0.02) 20
0.59-0.75 (0.66±0.01) 20
0.56-0.75 (0.67±0.01) 20
0.65-0.88 (0.78±0.01) 20

with moderate punctures.
Intercoxal disk of 1st abdominal sternum completely striate on each side, andmoderately punctate along the apical and basal margins, the punctures becoming denser
at anterior comers.
Protibia with 6 large spinules on outer margin.
Male genitalia as shown in Fig. 150.
Specimens examined. [Hokkaido] 1 ex., Sakanoshita, Wakkanai, 1O/viii/1974, Y. Kusui; 2exs., Teshio-gawa, Teshio, 9-22/vii/1993, S. Hori (HOR); 6 exs., Kitahama, Abashiri, 1O/vii/1958,
T. Nakane (NA); 2 exs., Toen-kaigan, Taiki, 23/vi-7/vii/1993, S. Hori (HOR); 7 exs., Zenibako,
24/vi/1954, T. Kumata (ElHU); 1 ex., Ditto, 21/vi/1985, M. Ohara; 1 ex., Momigi-yama, Ishikari,l/vi/1974, S. Kudo. Okushiri Is.: 2c?'c?' ,lex., Aonae, 28/vi/1986, Y. Kusui.
[Honshu] Yamagata-ken: 1 ex., Miyaumi, Yamagata, 28/v/1953, K. Shirahara (NA). Ibarakiken: 2 exs., Tone River, Kawachi vill., 1O/x/1985, M. Kiuchi. Chiba-ken: 2 exs., Iwai, Boso-hanto,
23/viiil1966, Y. Kusui; 2 exs., Okitsu, v/1913, F. Muir (BSM). Niigata-ken: 12 exs., Niigata-shi,
5/v/1933, K. Baba (NA). Ishikawa-ken: 4 exs., Uchida-sakyu, 31/viiiI1968, H. Ebihara. Osaka-fu: 3exs., Yodo River, 31/x/1954, T. Shibata (NA). Hyogo-ken: 2 exs., Hiraiso, 14/viii/1954, T. Shibata(NA). Wakayama-ken: 1 ex., Minabe, 17/iii/1942, K. Sakaguchi (NA).
[Kyushu] Fukuoka-ken: 1 ex., Hakozaki, l/ix/1964, F. Nakasuji (NA). Kagoshima-ken: 1 ex.,
Fukiage, 29/vii/ 1984, M. Ohara; 2 exs., Uchinoura, Hirosegawa-kaigan, 5/v/1991, K. Haga.

Distribution (Fig. 152). Japan (Hokkaido, Honshu, Shikoku, Kyushu).
Remarks. Hypocaccus sinae superficially resembles H. (Baeckmanniolus)
varians, but can be distinguished from the latter by the characteristics given in the key.
This species occurs on sandy beaches. It is occasionally found also on the sandy
beach of mountain rivers (Kurosawa, 1988).
Hypocaccus (Hypocaccus) subaenus (Schmidt, 1890)(Fig. 145, 146, 147, 151, 152)
Saprinus subaenus Schmidt, 1890: 53 [Japan: Nikko].
Hypocaccus subaenus: Schleicher, 1930: 137; Reichardt, 1941: 308,321.
Hypocaccus (? Hypocaccus) subaenus: Kryzhanovskij and Reichardt, 1976: 229.

Japanese name: Yarna-hamabe-emma-mushi.

Description. Body length, PPL, 2.22 - 2.91 mm (2.48 ± 0.04, n==20), PEL,
252

2.03 - 2.62 mm (2.27 ± 0.03, n=20). Width, 1.70 - 2.22 mm (1.92 ± 0.03, n=20).
Biometric data are given in Table 39. Body oval and moderately convex. Cuticle
shining and black or nearly so, with bronzy, blue or brassy lustre; antennae dark
rufopiceous with club rufotestaceous; legs dark reddish brown.
Frontal stria of head (Fig. 147B) complete, carinate and straight on each side.
Supraorbital stria complete. Disk of front scabrous on apical 2/3 and sometimes
wholly. Epistoma scabrous.
Pronotal sides (Fig. 145B) feebly arcuate and convergent on basal 4/5, thenstrongly arcuate and convergent apically. Apical angles round. Marginal stria carinate
and complete. Disk coarsely and densely punctate except mediobasally, the punctures
becoming more rugose on lateral region; base with 2 or 3 transverse rows of coarse
punctures with fine ones intermingled, the rows broadest at middle and becoming
progressively narrower laterally.
Epipleura of elytra sparsely clothed with fine punctures. Marginal epipleural stria
complete and feebly carinate. Marginal elytral stria complete and well carinate, its apicalend extending near to lateral 1/6 of elytron. External subhumeral stria absent. Internal
subhumeral stria shortly present on median 113. Oblique humeral stria well impressedon basal 113. First dorsal stria deeply impressed and crenate on basal 4/5. Second
dorsal stria present on basal 3/5. Third dorsal stria a little longer than the 2nd. Fourth
dorsal stria present on basal 112. Sutural stria represented by a fine complete line; thebasal end united with the 4th stria in an arch. Surface of elytra smooth on basal 112,and densely covered with coarse punctures on apical 112 and sometimes also on basal

Fig. 151. Hypocaccus (Hypocaccus) subaenus (Schmidt). A: Aedeagus, dorsal view. B: Ditto, lateralview. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateral view. E: Male
genitalia, 9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: Ninth sternum (spicule),
dorsal view.

253Hypocaccus axeli,..

.. Hypocaccus sinae

~~

~

..

,g

", Hypocaccus subaenus

Fig. 152. Collection sites of Hypocaccus spp. in Japan.

area between 4th dorsal and marginal elytral striae, the punctures often rugose; extreme
apex of elytra (around apical stria) and flanks impunctate, but sometimes flanks
coarsely punctate.
Propygidium densely covered with coarse punctures, which become finer and254sparser towards base. Pygidium convex, and densely and moderately punctate, thepunctures sparser apically and medially.
Anterior margin of median portion of prosternum (Fig. 146B) obtusely angulate
or round. Preapical foveae distinctly present. Keel somewhat broad, and moderately
compressed medially, the apex round. Descending lateral striae complete, carinate, andconvergent at apex. Carinal striae present on basal 2/3, convergent apically andsometimes united with each other, rarely completely present, its apical end attaining to
the apex of keel. Lateral side of keel sparsely with moderate punctures, with finepunctures intermingled.
Anterior margin of mesosternum feebly emarginate medially, its marginal stria
well impressed on each side behind angle, usually interrupted in median 113, rarely
complete. Disk transverse (19 : 5), evenly clothed with fine or moderate punctures.
Meso-metasternal suture finely impressed, accompanied with a coarsely crenatedparallel line. Intercoxal disk of metasternum sparsely clothed with fine punctures, andwith a transverse apical band of sparse and moderate punctures. Lateral metasternal
stria well impressed, carinate, extending obliquely and posteriorly, and reaching nearhind coxa. Lateral disk densely and largely punctate, the punctures becoming sparser
apically. Metepisternum densely covered with large punctures.
Intercoxal disk of 1st abdominal sternum completely and crenatel y striate on eachside, and moderately punctate along posterior margin and inside the lateral striae.
Protibia with 8 spinules on outer margin.
Male genitalia as shown in Fig. 151.
Specimens examined. [Hokkaid6] 8 exs., Mt. Obira-yama, 29/v, 6-8/viii/1988, M. Ohara, fromdung of bear; 56 exs., Toyotaki, J6zankei, 4, 6/vi, 14/viiI1986, l/v, 3, 18/vi/1987, M. Ohara.
[Honshu] Kyoto-fu: 1 ex., Kurama, 4/vi/1933, T. Kimura (NA).

Distribution (Fig. 152). Japan (Hokkaid6, HonshU).
Remarks. Hypocaccus subaenus can easily be distinguished by the front of thehead with coarse rugae, the dense punctation of the pronotum, the mesosternum withcoarse punctures and the interrupted mesosternal marginal stria.
Hypocaccus subaenus usually occurs on the sandy beach of mountain rivers.
Some specimens were collected under dung of bear, Ursus arctos Linnaeus, on thebeach of a river.
Hypocaccus (Hypocaccus) akanensis M. Ohara, sp. nov.
(Fig. 153, 154, 155)
Japanese name: Akan-hamabe-emma-mushi.

Description. Body length, PPL, 2.13, PEL, 1.90. Width, 1.52. Biometric data
as follows: APW 0.67, PPW 1.43, PL 0.76, EW 1.52, ProW 0.88, ProL 0.23, PTL
0.53, MSTL 0.55, MTTL 0.61. Body oval, convex, not shining and black; legs andantennae dark reddish brown.
Frontal stria of head (Fig. 153B) rudimentary behind apical angle (but it seems
that the state of the only specimen available is not complete, with dermal structures
defaced to some extent). Disk of front with rudimentary transverse rugae.
Pronotal sides (Fig. 153A, 154) convergent forward on basal 116, thence
strongly arcuate and convergent apically. Apical angles round. Marginal stria carinate
255and complete laterally, the apical portion broadly interrupted behind head. Disk
coarsely and densely punctate, the punctures becoming more rugose in lateral region
and absent mediobasally; base with 2 or 3 transverse rows of large and oblong
punctures, the rows being broadest at middle and becoming narrower laterally.

J
Fig. 153. Hypocaccus (Hypocaccus) akanensis M. Ohara, sp. nov. A: Pronotum and left elytron. B:
Head, frontal view. C: Pro sternum, mesosternum, metasternum and 1st abdominal sternum.

Fig. 154. Hypocaccus (Hypocaccus) akanensis M. Ohara, sp. nov. A: Holotype, dorsal veiw. B:
Left antero-Iateral area of pronotum, dorsal view.

256

Marginal epipleural stria complete and clearly impressed. Marginal elytral stria
complete, broadly impressed in basal 113, the apical end extending inwards, reaching tonear lateral 114 of posterior margin of elytron. External subhumeral stria absent.
Internal subhumeral stria impressed on median 113. Oblique humeral stria present on
basal 113, the area around the stria with several short rugae. First dorsal elytral stria
completely present, but shortly reduced on apical 118. Second to 4th dorsal striae wellimpressed on basal 112, the basal end of 4th extending inwards and united with thebase of sutural stria. Sutural stria completely impressed. Surface of elytra smooth onbasal 112, and densely covered with coarse and deep punctures, which are often united
to form longitudinal rugae; extreme apex of elytron and flank sparsely and finely
punctate.
Propygidium densely covered with coarse and round punctures, which become
finer and sparser towards the base, but mostly are separated by their own diameter.
Pygidium densely covered with coarse punctures, which are a little smaller than thoseof the propygidiurn and separated by 0.3 - 0.5 times their diameter; on the apex the
punctures becoming finer and sparser.
Anterior margin of median portion of prosternum (Fig. 153C) weakly obtuseangulate. Preapical foveae deeply and distinctly present. Keel narrow, strongly
compressed, its anterior apex truncate. Descending lateral striae carinate and convergent
to apex, but their apical portions nearly parallel. Carinal striae completely impressed,
divergent apically and basally, the anterior ends united with descending lateral striae.
Lateral side of keel coarsely punctate, the punctures being separated by about l.5 (1 2) times the dimeter.

~~~lo~.5~m~m~

B____________________~

.. '

,

0.

Fig. 155. Hypocaccus (Hypocaccus) akanensis M. Ohara, sp. nov. A: Aedeagus, dorsal view. B:
Ditto, lateral view. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateralview. E: Male genitalia, 9th and lOth tergites, dorsal view. F: Ditto, lateral view. G: Ninthsternum (spicule), dorsal view.

257

Anterior margin of mesosternum feebly emarginate, its marginal stria carinate andcomplete. Disk of mesosternum smooth. Meso-metasternal suture lightly impressed,
accompanied with a coarsely crenated parallel line. Intercoxal disk of metasternum
smooth. Lateral metasternal stria well impressed and carinate, extending obliquely andposteriorly and reaching near hind coxa. Lateral disk densely covered with coarse,
round and shallow punctures on basal 112, the punctures gradually becoming smallerand sparser; impunctate along the lateral stria and on posterior 112 of the disk.
Metepisternum densely and coarsely punctate.
Intercoxal disk of 1st abdominal sternum smooth, and completely striate on eachside.
Protibia with 6 spinules on outer margin, its apical outer angle bearing 2 large
denticles.
'
Male genitalia as shown in Fig. 155.
Specimens examined. Holotype, d', Mount Akan, Kushiro, Hokkaid6, 5/viii/1918, M. Suzuki(ELKU).

Distribution. Japan (Hokkaido).
Remarks. The aedeagus of this species is similar in shape to that of Hypocaccus(Hypocaccus) speculum (Schmidt) (Kryzhanovskij and Reichardt, 1976: 225, fig.
443), but the punctuation of elytra is quite different.
Hypocaccus akanensis resembles superficially H. sinae but can be
distinguished by the smaller body and the sutural stria being not united with marginal
elytral stria.

SUBGENUS BAECKMANNIOLUS REICHARDT, 1926
Baecknumniolus Reichardt, 1926b: 14 [type species: Hister dimidiatus Illiger, 1807: 41, originallydesignated].
Pachylopus Zimmermann in J. L. LeConte, 1869: 253 (part.), synonymized by Reichardt, 1926b: 14.

Diagnosis. Pronotum smooth, or a little covered with punctures behind eachanterior angle. Frontal disk of head smooth, sometimes irregularly rugose in apicalarea.

Hypocaccus (Baechnanniolus) varians varians (Schmidt, 1890)(Fig. 156,157,158)
Saprinus varians Schmidt, 1890: 55 [Japan; China]; Lewis, 1910: 58; Desbordes, 1919: 416.
Hypocaccus varians: Reichardt, 1926b: 15; Reichardt, 1941: 323,326.
Hypocaccus (Baeckmanniolus) varians: Kryzhanovskij and Reichardt, 1976: 229.

Japanese name: Hamabe-emma-mushi.

Description. Body length, PPL, 2,45 - 4.19 mm (3.44 ± 0.10, n=20), PEL,
2.22 - 3.60 mm (3.11 ± 0.08, n=20). Width, 1.90 - 3.20 mm (2.56 ± 0.07, n=20).
Biometric data are given in Table 40. Body oval and moderately convex. Cuticle verystrongly shining and black or nearly so with strong bronzy, or brassy lustre; antennae
dark rufopiceous with club rufotestaceous; legs dark reddish brown.
Frontal stria (Fig. 156C) carinate, complete and straight on each side.
258

Supraorbital stria well impressed. Disk of front with only 1 ruga, sometimes with 2 or
3 short ones. Epistoma scabrous.
Pronotal sides (Fig. 156A) rather feebly arcuate and feebly convergent on basal
4/5, thence strongly arcuate and convergent apically. Apical angles round. Marginalstria carinate and complete. Disk smooth, rarely with several moderate punctures inanterior angle. Basal margin of pronotum with 1 row of moderate punctures, whichbecome finer near the middle and on the lateral 116.
Epipleura of elytra sparsely clothed with microscopic punctures and even.
Marginal epipleural stria complete and finely impressed. Marginal elytral stria carinate,complete and densely punctate, its apical end extending along the posterior margin of
elytron, and united with the apical end of sutural stria at medio-apical angle of elytron.
External subhumeral stria absent. Internal subhumeral stria shortly present on median
113. Oblique humeral stria present on basal 113. First dorsal stria present on basal 112
and well impressed. Second dorsal stria a little longer than the 1st, and densely crenate
apically. Third dorsal stria about as long as the 2nd, and densely crenate. Fourth dorsal
stria shortly present on median area, composed of 5 or 6 punctures. Sutural stria
present on apical 2/3, finely impressed, its apical end united with the marginal elytral
stria. Mediobasal area with a short arch consisting of the ends of 4th dorsal and sutural
striae, sometimes the arch extending apically, rarely continued to the basal end of
sutural stria. Surface of elytra smooth on basal 112; on apical 112 moderately andsparsely punctate except on extreme apex (around apical stria), the punctures being
separated by about 1.5 (1 - 2) times the dimeter.
Propygidium densely and coarsely punctate. Pygidium (Fig. 1560) densely andcoarsely punctate on basal 2/3, the apical 113 being sparsely punctate along margin andmid line, with 2 convex and impunctate areas on both sides.

~J
~~-~

a/L-}
D
Fig. 156. Hypocaccus (Baeckmanniolus) varians varians (Schmidt). A: Pronotum and left elytron.
B: Prosternum and meso- and metasterna. C: Head. D: Pygidium.

259

0.5 mm ,B...

--~- ~-~:.::.-----,0.5 mmFig. 157. Hypocacaccus (Baeckmanniolus) varians varians (Schmidt). A: Aedeagus, dorsal view. B:
Ditto, lateral view. C: Male genitalia, 8th tergite and sternum, dorsal view. D: Ditto, lateralview. E: Male genitalia, 9th and 10th tergites, dorsal view. F: Ditto, lateral view. G: Ninthsternum (spicule), dorsal view.

Anterior margin of median portion of prosternum (Fig. 145B) obtuse-angulate.
Preapical foveae distinctly present. Keel moderately compressed, its apex round.
Descending lateral stria carinate and convergent to apex. Carinal striae present on basal
112 and convergent apically. Lateral sides of keel impunctate.
Anterior margin of mesosternum feebly emarginate, its marginal stria carinate andusually complete, sometimes interrupted at middle. Disk smooth. Meso-metasternal
suture finely impressed, without a crenate parallel line. Intercoxal disk of metasternum
smooth, sparsely clothed with microscopic punctures except for a transverse apicalband of sparse and fine punctures. Lateral metasternal stria well impressed, carinate,
extending obliquely and posteriorly, reaching near hind coxa. Lateral disk evenly andcoarsely punctate, the punctures becoming sparser apically. Metepisternum densely
covered with moderate punctures.
Intercoxal disk of 1st abdominal sternum completely striate on each side, andfinely punctate along posterior margin.
Protibia with 7 spinules on outer margin.
Male genitalia as shown in Fig. 157.
Specimens examined. [Hokkaido] 1 ex., Daisetsu, 7/viii/1954, T. Nakane (NA); 1 ex.,
Kitahama, Abashiri, 1O/vii/1958, T. Nakane (NA); 2 exs., Toyokita-kaigan, Urahoro, 24/vi7/vii/1993, S. Hori (HOR); 1 ex., Ishikari-Kako, 19/ix/1987, M. Hinakura; 6 exs., Ishikari,lIvii/1976, M. Kiuchi; 5 exs., Raisatsu, Ishikari, 9/vi/1993, S. Hori (HOR); 1 ex., Maruyama,
Sapporo, 23/vi1l979, N. Nishikawa; 2 exs., Toyotaki, Sapporo, 14/vi/1987, M. Ohara; 1 ex.,
Zenibako, Otaru, 24/vi/1954, T. Kumata (ElHU); 38 exs., Ditto, 23, 30/v, 8/vii/1977, 1O/v/1978,
19/vi/1979,N. Nishikawa; 31 exs.,Ditto,14/v,30/viii/1985,M. Ohara; 2 exs., Kojo-hama, Shiraoi,
30/vii/1953, M. Otake (NA); 1 ex., Iwanai, 13/viii/1946, Y. Nishijima; 3 exs., Shiriuchi, Oshima,
8/vii/1976, T. Kumata (ElHU); 10 exs., Nezaki-kaigan, Hakodate, 19/vii/1952, K. Honma (NA); 9
exs., Hakodate, 19/vii/1954, T. Nakane (NA); 25 exs., Ditto, 19/vii/1954, K. Sawada (NA); 2 exs.,
260Ditto, 19/vii/1954, K. Sawada (NA); 4 exs., Ditto, 19/viii/1952, K. Honma (ElHU). Okushiri Is.:
1 ~, Aonae, 28/vi/1986, Y. Kusui.
[Honshu] Yamagata-ken: 1 ex., Fukura, 11/vi/1972, H. Takizawa. Fukushima-ken: 1 ex.,Nashidaira, Fukushima-shi, 3/vii/1988, K. Tazoe; 11 exs., Fujima-kaigan, Iwaki, 25/vi/1988, K.
Tazoe. Chiba-ken: 130 exs., Oami, 19/vi/1984, 20/v/1986, lIiv/1987, M. Ohara; 5 exs, Shirako,
18/v/1984, M. Ohara; 1 ex., Katsuura, 3/viii/1963, M. Ishida; 1 ex., Iwai, Boso-hanto, 23/viii/1966,
Y. Kusui; 2 exs., Narashino, 18/viI1963, S. Kondo; 1 ex., Cape Futtu, 2/vii/1961, H. Yamazaki(NA); 4 exs., Ichinomiya, Kazusa, 6/vi/1936, T. Nakane (NA). Kanagawa-ken: 2 exs., Aburatsubo,
111v/1959, S. Kondo; 2 exs., Koshigoe, 3/viii/1944, Y. Kurosawa (NA); 5 exs., Kamakura-koko-eki,
Shichiri-hama, Kamakura, 20/iv/1991, K. Haga. Niigata-ken: 1 ex., Niigata-shi, 5/v/1933, K. Baba
(NA). Shizuoka-ken: 1 ex., Heda, Mihama-misaki, Izu-hanto, 7/vi/1974, M. Ishida. Aichi-ken: 35
exs., Miya, Mikawa, 1-2/ix/1947, S. Osawa (NA). Wakayama-ken: 4 ex., Nachi, Kii, 19/vii/1946, 26,
29/iv/1947, S. Osawa (NA); 1 ex., Ditto, 28/iii/1953, K. Sawada (NA); 3 exs., Shingu, southern Kii,
10/vii/1946, S. Osawa (NA); 1 ex., Minabe, sea shore, 2/vi/1966, M. Yoshida (NA). Osaka-fu: 1 ex.,Takaishino-hama, 1O/ix/1966, no collector's name. Ishikawa-ken: 10 exs., Uchinada-sakyfi,
31/viii/1968, H. Ebihara (NA). Hyogo-ken: 1 ex., Hiraiso, 14/viii/1954, T. Shibata (NA); 2 exs.,
Akashi, 8/v/1954, H. Ishida (NA); 3 exs., Mikage-kaigan, 22/vii/1938, K. Sakaguchi (NA). Okayamaken: 2 exs., Ushimado, Oku, 20/viii/1975, T. Aono (NA). Izu Isles: 2 exs., Shikine-jima, 1941, K.
Oyama (NA).
[Shikoku] Tokushima-ken: 1 ex., Kiki, Yuki-cho, 4/v/1986, S. Mano; 2 exs., Anan,
Kitanawaki-kaigan, lIvi11986, S. Mano.
[Kyushil] Fukuoka-ken: 2 exs., Satsuki-matsubara, 8/v/1983, S. Nakao (NA); 5 exs.,
Ainoshima, 22/v/1983, S. Nakao (NA). Nagasaki-ken: 2 exs., Kuchinotsu, Shimabara, 20/ix/1977, S.
Imasaka; 3 exs., Noda-hama, Kazusa-cho, 30Iiii/1985, S. Imasaka; 1 ex., Ditto, 5/iv/1977, S.
Imasaka. Kagoshima-ken: 26 exs., Chosa, 14/vi/1983, 22/v/1984, M. Ohara; 24 exs., Fukiage,
29/vii/1984, M. Ohara; 1 ex., Sata-misaki, 29/v/1971, K. Tazoe. Tsushima Is.: 8 exs., Kuro-shima,
4/viii/1959, K. Tsukamoto (NA).
[Nansei Isles] Tane-ga-shima Is.: 15f, 59 exs., Hamada, 31Ivii/1984, Y. Harada. AmamiOshima Is.: 69 exs., Akaogi, 2/vii/1984, M. Ohara. Tokuno-shima Is.: 2 exs., Kametsu, 24/iv/1954,
T. Kumata (EIHU). Yoron Is.: 1 ex., 27/iii/1967, T. Matsumoto. Ishigaki Is.: 1 ex., Botanical Garden,
Banna, 12/vi/1974, H. Makihara.

Distribution (Fig. 158). Japan (Hokkaid6, Honshu, Shikoku, Kyushu); China;
Taiwan; Sakhalin; Viet-Nam; Philippines; Sri Lanka; Solomon Is.; Austmlia.
Remarks. Hypocaccus varians resembles H. sinae superficially; however, it
can easily be distinguished from the latter by the mesosternum without carinal line, theelytral sutuml stria usually absent on the basal 112 and the apical 112 of the pygidium
impunctate.
Table 40. Biometric data of Hypocaccus v. varians (Schmidt) and H. v. hatsune (Nakane).

APW
PPW
PL
EL

EW
ProW
ProL
PyL
PTL
MSTL

MTTL

H. v. varians

H. v. hatsune

0.72-1.21 (1.02±0.03) 20
1.67-2.71 (2.31±O.06) 20
0.88-1.37 (1.12±0.03) 20
1.41-2.22 (1.89±0.05) 20
1.90-3.20 (2.56±0.07) 20
1.21-1.90 (1.53±0.04) 20
0.36-0.59 (0.46±0.01) 20
1.14-1.81 (1.48±0.04) 20
0.65-1.01 (0.88±0.02) 20
0.62-1.05 (0.87±0.02) 20
0.65-1.18 (1.01±0.03) 20

0.62-0.98 (0.87±O.03) 13
1.54-2.26 (1.98±0.07) 13
0.85-1.21 (1.06±0.03) 13
1.18-1.93 (1.61±0.07) 13
1.67-2.45 (2.14±0.08) 13
1.01-1.57 (1.27±0.05) 13
0.26-0.43 (0.36±0.01) 13
1.05-1.67 (1.32±0.05) 13
0.59-0.88 (0.77±0.03) 13
0.56-0.88 (0.74±0.03) 13
0.65-1.08 (0.90±0.03) 13

261

;.' /,/ <'~ ".',
l~ . '< "".~~/ ~'v/ Hypocaccus varians varians
I

/,

'"

7

Fig. 158. Collection sites of Hypocaccus (Baeckmanniolus) varians varians (Schmidt) in Japan.

Hypocaccus varians is a very common species. It lives on sandy beaches andoccurs in various decaying animal matters, such as carrion of birds, dog, turtle andfish.
Hypocaccus (Baeckmanniolus) varians hatsune (Nakane, 1977)
Baeckmanniolus varians hatsune Nakane, 1977: 148, 157 [Hatsune-ura, Chichi-jima, Ogasawara].

Japanese name: Ogasawara-hamabe-emma-mushi.

Additional description. Body length, PPL, 2.45 - 3.83 mm (3.06 ± 0.12,n=13), PEL, 1.99 - 3.20 mm (2.76 ± 0.11, n=13). Width, 1.67 - 2.45 mm (2.14 ±
0.08, n=13). Biometric data are given in Table 40. Punctation of pygidium represented
only on basal 113 - 112.
Specimens examined. Holotype, Hatsune-ura, Chichi-jima, Bonin Is. (Ogasawara Is.),5/vi/1975, K Ichikawa (NSMT); Paratypes, 13 exs., Ditto (NA); 31 exs., Ditto (NSMT).

Distribution. Japan (Ogasawara Isles).
GENUS EoPACHYLOPUS REICHARDT, 1926
Eopachylopus Reichardt, 1926b: 14 [type species: Pachylopus ripae Lewis, 1885a: 469, originallydesignated].

262

Fig. 159. Eopachylopus ripae (Lewis). A: Adult, dorsal view. B: Ditto, ventral view. C: Head andpronotum, oblique frontal view. D: Antenna. E: Prostemum, ventral view. F: Scape of antenna.

263

Eopachylopus ripae (Lewis, 188S)(Fig. IS9, 160, 161, 162)
Pachylopus ripae Lewis, 1885a: 469 [Enoshima and Hakodate, in Japan].
Eopachylopus ripae: Reichardt, 1926b: 14.
Eopachylopus ripae abo tsherskii Reichardt, 1941: 328.
Neopachylopus ripae f. rufofasciatus Osawa, 1952: 4.

Japanese name: Tsuya-hamabe-emma-mushi.

Description. Body length, PPL, 2.09 - 3.34 mm (2.61 ± 0.06, n=20), PEL,
1.96 - 2.91 mm (2.40 ± O.OS, n=20). Width, 1.64 - 2.49 mm (1.98 ± 0.04, n=20).
Biometric data are given in Table 41. Body oval and strongly convex. Cuticle shining
and black wholly or with a red spot on each elytron, rest of body black; antennae dark
rufopiceous with club rufotestaceous; legs dark reddish brown.
Frontal stria of head (Fig. IS9C) lightly carinate anteriorly and absent laterally.
Disk smooth, sparsely clothed with microscopic punctures. Epistoma smooth.
Pronotal sides (Fig. IS9A) nearly straight and weakly convergent on basal S/6,
thence strongly arcuate and convergent apically. Apical angles round. Marginal stria
complete and feebly carinate basally. Disk smooth and impunctate.
Marginal epipleural stria weakly carinate and complete. Marginal elytral stria
strongly carinate and complete, its apical end extending across elytral apex and united
with apical end of sutural stria. Surface of epipleura between marginal epipleural andelytral striae smooth and shining on basal 112; on apical 112 sparsely with shallow andtransverse impressions. Subhumeral stria absent. Oblique humeral stria sharply
impressed on basal 112. First, 2nd and 3rd dorsal striae well impressed, beginningfrom basal 116; 1st shortly present, ending at basal 113; 2nd ending at basal 112; 3rd
ending at basal 2/3. Fourth dorsal stria confined to basal region, usually represented byan arch. Sutural stria deeply impressed, nearly complete, beginning at basal fifth and its
apical end united with end of marginal elytral stria. Surface of elytra smooth, sparselywith microscopic punctures.
Propygidium sparsely covered with fine punctures, which are coarser and denser
laterally. Interspace among punctures with lightly impressed, fine and irregularlytransverse alutaceous microsculpture, which are heavily impressed on basal 1/2.
Pygidium sparsely clothed with microscopic punctures, and shining. Interspace among
punctures with more lightly impressed irregularly transverse microsculptures than on
propygidium.
Anterior margin of median portion of prosternum (Fig. IS9E) obtuse-angulate.
Preapical fovea weakly impressed on lateral side of keel at apical 116. Keel strongly
compressed, on basal 113 represented by a triangular disk. Carinal striae of keel
distinctly and shortly impressed between precoxae, convergent anteriorly and united
with each other at apex. Descending lateral striae shortly present and convergent
apically, their apical ends attaining at middle of keel.
Anterior margin of mesosternum strongly emarginate medially, its marginal stria
well impressed, complete and carinate. Disk smooth, sparsely with microscopic
punctures. Meso-metasternal suture sparsely and coarsely crenate. Intercoxal disk ofmetasternum smooth, sparsely clothed with microscopic punctures; feebly depressedon anteromedian area, and strongly depressed on nearly apical 112 along longitudinal264

Fig. 160. Eopachylopus ripae (Lewis). A: Protibia, dorsal view. B: Tarsus of fore leg. C: Protibia,
ventral view. D: Mesotibia, ventral view. E: Metatibia, ventral view. F: Setae of metatibia.

265

suture. Lateral metastemal stria well impressed and carinate, extending obliquely andposteriorly, reaching near hind coxae. Lateral disk densely covered with large, shallow
and setiferous punctures. Interspace among punctures with fine, irregular andtransverse alutaceous microsculptures. Posterior mesocoxal stria well impressed,
carinate, and extending along posterior margin of mesocoxal cavity. Disk ofmetepisternum densely covered with large punctures.
Intercoxal disk of 1st abdominal sternum broadly depressed mediobasally andsparsely covered with moderate punctures before posterior corner. Lateral stria of thedisk complete, well impressed and carinate.

---:;;;?
O.25mmIo 25mmFig. 161. Eopachylopus ripae (Lewis) A: Aedeagus, dorsal view. B: Ditto, lateral view. C: Male
genitalia, 8th tergite and sternum, dorsal view. D: Apex of 8th sternum, ventral view. E: Eighth
tergite and sternum, lateral view. F: Male genitalia, 9th and 10th tergites, dorsal view. G: Ditto,
lateral view. H: Ninth sternum (spicule), dorsal view.

Eopachylopus ripae

Fig. 162. Collection sites of Eopachylopus ripae in Japan.

266

Table 41. Biometric data of Eopachylopus ripae (Lewis).
APW
PPW
PL
EL
E\V

ProW
ProL
PyL
PTL
MSTL
M1TL

0.72-1.11 (0.88±0.02) 20
1.50-1.99 (1.81±0.03) 20
0.72-1.05 (0.85±0.02) 20
1.18-1.83 (1.52±0.04) 20
1.64-2.49 (1.98±0.04) 20
0.95-1.47 (1.14±0.03) 20
0.20-0.39 (0.25±0.01) 20
0.86-1.48 (1.17±0.04) 20
0.59-0.92 (0.75±0.02) 20
0.52-0.92 (0.72±0.02) 20
0.65-1.08 (0.85±0.02) 20

Protibia (Fig. 160A, B, C) with 10 spines on outer margin, the 3 spines on
apical 112 being clavate. Ventral sides of meso- and metatibiae with many stout spines.
Male genitalia as shown in Fig. 16l.
Specimens examined. [Hokkaid6] 2 exs., Muroran, 25/vi/1960, T. Yamashita.
[Honshu] Fukushima-ken: 1 ex., Fujima-kaigan, Iwaki, 25/vi/1988, K. Tazoe. Chiba-ken: 7exs., Oami, 19/vi/1984, l/v/1990, M. Ohara. Aichi-ken: 1 ex., Miya, Mikawa, 1-2/ix/1947, S. Osawa(NA). Mie-ken: 5 exs., Akogi-ga-ura, Mie Coast, 30/vi/1957, M. Got6 (NA and NSMT); 1 ex., Tsukaigan, 30/v/1957, Z. Naruse (NSMT). Wakayama-ken: 32 exs., Nachi, southern Kii, 26/iv/1947, S.
Osawa (NA); 2 exs., Ditto, 19/vii/1946, S. Osawa (NA); 14 exs., Ditto, 29/iv/1947, S. Osawa (NA).
Ishikawa-ken: 7 exs., Uchinada-sakyu, 311viii/1968, H. Ebihara.
[Kyfishu] Fukuoka-ken: 1 ex., Satsuki-Matsubara, 8/v/1983, S. Nakao (NA); 1 ex.,
Ainoshima, 22/vI1983, S. Nakao (NA).
[Nansei Isles] Tane-ga-shima Is.: 2 exs., Hamada, 31/vii/1984, Y. Harada.

Distribution (Fig. 162). Japan (Hokkaid6, HonshQ, Shikoku, KyQshQ, Nansei
Isles).
Remarks. Eopachylopus ripae is easily recognized by the character states givenin the key and description; it is hardly confused with any other species of the tribe.
The species occurs under decaying bodies of fish on sandy beaches.
6.3. 10. SUBFAMILY TRYPETICINAE BICKHARDT, 1913
GENUS TRYPETICUS MARSEUL, 1864
Trypeticus Marseul, 1864: 281 [type species: Trypeticus gilolous Marseul, 1864: 283, designated by
Bickhardt, 1916: 53]; Schmidt, 1893a: 15; Jakobson, 1911: 638,642; Bickhardt, 1916-17: 53;
Reichardt, 1941: 73; Kryzhanovskij and Reichardt, 1976: 79.

Key to the Japanese species of the genus Trypeticus
1(2)
2(1)

Body shining. Pronotum without lateral stria ..................................... T.fagi (Lewis, 1884)
Body filiform. Pronotum with a complete lateral stria..................... T. venator (Lewis, 1884)

Trypeticusfagi (Lewis, 1884)(Fig. 163,164,165,166)
Tryponaeusfagi Lewis, 1884: 138 [Japan, no further locality].
Trypeticusfagi: Lewis, 1905b: 7; 1912: 251 [male and female genitalia illustrated].
267

Japanese name: Tsutsu-emma-mushi.

Description. Male: Body length, PPL, male, 3.11 - 3.50 (3.36 ± 0.04, n=10)mm, female, 3.11- 3.60 mm (3.42 ± 0.06, n=7), PEL, male, 2.84 - 3.11 mm (2.99± 0.03, n=lO), female, 2.94 - 3.14 mm (3.07 ± 0.02, n=7). Width, male, 1.31 1.34 mm (1.31 ± 0.001, n=lO), female, 1.24 - 1.31 mm (1.28 ± 0.01, n=7).
Biometric data are given in Table 42. Body cylindrical, moderately stout. Cuticle
shining, black; tibiae, tarsi, and antennae dark rufopiceous.
Head without frontal stria (Fig. 164C, D); surface slightly depressed on medianarea, and sparsely clothed with fine punctures which are separated by 2 - 5 times theirdiameter and become coarser posteriorly; surface among the punctures extremely finely
covered with alutaceous ground sculpture.
Anterior margin of pronotum shallowly emarginate on median 2/3; sides parallel(Fig. 163) and straight; marginal stria complete and carinate laterally, and , on anterior
margin, absent behind head; surface of disk densely and coarsely punctate, but not on
longitudinal mid line, the punctures becoming denser on apical 114 and on area along
lateral margin; surface among coarse punctures usually impunctate and shining, butsometimes covered with fine punctures, which are 113 as large as the coarse punctures.
Epipleura broad, not concave. Marginal epipleura and elytral stria absent.
Narrow band along epipleural margin impunctate, but extremely finely striate. Sutural
stria completely impressed. Surface of elytra irregularly scattered with coarse punctures
which are somewhat smaller than those of the pronotum and usually separated by 2 - 5times their diameter, the punctures becoming coarser on basal margin and denser along
the sutural line and on lateral area along epipi eura; a small oblong area on humeral
region impunctate; extreme apex of elytra impunctate.
Disk of propygidium (Fig. 164) evenly covered with round and shallow
punctures, which are a little coarser than the elytral ones and separated by about theirown diameter, and with other fine punctures intermingled. Punctation of pygidium

Fig. 163. TrypeticusJagi (Lewis). A: Adult, dorsal view.

268

Fig. 164. Trypeticus fagi (Lewis). A, B, E: Female. C, D: Male. A, C: Head and pronotum, dorsal
view. B, D: Head, frontal view. E: Propygidium and pygidium, dorsal view.

similar to that of propygidium, but much sparser, the punctures becoming finer
apically; tip of pygidium clearly acute.
Prosternal process longitudinal quadrangular; anterior margin straight; marginalstria of the process nearly complete and somewhat carinate; disk densely covered withcoarse and shallow punctures and intermingled with fine punctures among the coarse
ones; surface among these punctures finely clothed with alutaceous ground sculpture.
Median 113 of anterior margin of mesosternum nearly straight; lateral 113
inwardly curved, fitting to posterior margin of procoxa. Lateral sides of mesosternum
nearly parallel and straight on basal 112 and divergent apically on apical 112; each lateralside marginate and deeply carinate; punctation of disk similar to that of prosternum.
269

Meso-metastemal suture impressed and obtuse-angulate (about 120°) at middle.
Metasternum clearly depressed on longitudinal mid line; lateral metasternal striae
slightly divergent posteriorly and impressed on basal 2/3. Intercoxal disk ofmetasternum sparsely covered with coarse, somewhat longitudinally oblong andshallow punctures which are separated by 1 - 3 times the dimeter; surface amongcoarse punctures sparsely and finely punctate except on a broad band along longitudinal
mid line. Lateral disk of metasternum sparsely and coarsely punctate, the punctures
much sparser than those of the intercoxal disk. Metepisternum with 2 rows of dense
and coarse punctures.
lntercoxal disk of 1st abdominal sternum densely covered with coarse punctures
which are a little smaller than those of the metasternal intercoxal disk, the punctures
becoming sparser on basal 112.
Protibia slender, with 5 large denticles on outer margin. Mesotibia with 5
denticles on outer margin.
Female. In general appearance similar to the male except in the following
character states: head (Fig. 164A, B) with clear frontal marginal stria which is strongly
carinate; pronotum distinctly with short costa on apical 114 on longitudinal mid line,
and area around the costa densely clothed with longitudinal short rugae and finealutaceous ground microsculpture where it is lusterless; pronotum evenly punctate
wholly, without impunctate area on longitudinal mid line; punctation of pygidium muchdenser than that of the male; tip of pygidium round, not acute; prostemal process
without marginal stria.
Male genitalia as shown in Fig. 165.
Specimens examined. [Honshu] Aomori-ken: 1 Sf, no date, A. Abe; Id", Oirase, Nishigun,
Fukaura-machi, 23/v/1976, A. Abe; 1d", Yachi Spa., Towada, 26/vii/1952, Y. Kurosawa.
Fukushima-ken: 1d", Yunohana, Minami-Aizu, 20/vii11963, K. Tazoe; 1 Sf, Nagasaka, Iide-san,
14/vi/1970, K. Tazoe; 1d", Hinoemata, 30/v/1982, H. Makihara; 1 Sf, Shin-machi, Tateiwa, MinamiAizu, 18/vi/1947, Y. Kurosawa. Saitarna-ken: 1d", Yanagi-koya, Chi chi bu, 22/vii/1933, nocollector's name. Gifu-ken: Id",Idani, Kawai, Hida, Slvi/1955, H. Torigai; 2d"d", Ditto, lIvi/1957,
T. Nakane. Mie-ken: Id", Mie Univ. Forest., Ichishi-gun, 11/vi/1956, H. Ichihashi; 1 Sf, Ditto,
12/vi/1956, H. Ichihashi. Kyoto-fu: 1 Sf, Ashiil, vi/1951, A. N.
[Shikoku] Tokushima-ken: 1 d" , Ashu, 27/vii/1940, M. Tokunaga (Host: Aesculus turbinata
Blume). Ehime-ken: 1 Sf , Omogo, Il/viil 1956, Z. Naruse and M. Sato.

Table 42. Biometric data of Trypeticus fagi (Lewis).
Male
PW
PL

EL
EW
ProW
ProL
PyL

PTL
MSTL
MTTL

Female1.21-1.31 (1.27± 0.01) 11
1.44-1.57 (1.49± 0.01) 11
1.31-1.54 (1.46± 0.02) 11
1.31-1.34 (1.31±0.001) 10
0.72-0.85 (0.79± 0.01) 11
0.29-0.36 (0.33± 0.01) 11
0.49-0.59 (0.52± 0.01) 11
0.52-0.65 (0.62± 0.01) 11
0.49-0.65 (0.59± 0.02) 11
0.59-0.69 (0.63± 0.01) 11

270

1.24-1.31 (1.27±0.01) 7
1.47-1.64 (1.57±0.02) 7
1.34-1.50 (1.43±0.02) 7
1.24-1.31 (1.28±0.01) 7
0.69-0.82 (0.75±0.02) 7
0.26-0.33 (0.30±0.01) 7
0.49-0.65 (0.57±0.02) 7
0.52-0.62 (0.58±0.01) 7
0.52-0.65 (0.58±0.02) 7
0.59-0.65 (0.62±0.01) 7--A~0=.2~5-,m",m-"----_ _ _ _ _ _ _ _----,,

0.25 mm

I

A.G

H

Fig. 165. Trypeticusfagi (Lewis) A: Aedeagus, dorsal view. B: Ditto, lateral view. C: Male genitalia,
8th tergite and sternum (?), dorsal view. D: Ditto, lateral view. E: Male genitalia, 9th and 10th(?) tergites. F: Ninth sternum (spicule), dorsal view. G: Ditto, lateral view. H: Male genitalia,
dorsal view.

Trypeticus fagi

Fig. 166. Collection sites of Trypeticusfagi (Lewis) in Japan.

271

[?] 1d" 15j? ,Sakashimo, Rora, 13/vi/1954, H. Torigai.

Distribution (Fig. 166). Japan (HonshO, Shikoku, KyOshO, Izu Isles, Yakushima Is.).
Trypeticus venator (Lewis, 1884)
Tryponaeus vena tor Lewis, 1884: 138 ["South Japan, at Yuyama and Konose"].
Trypeticus venator: Lewis, 1905b: 8.

Japanese name: Roso-tsutsu-emma-mushi.
Specimens examined. No material of this species has been available for my study.

Distribution. Japan (HonshO, KyOshO); Taiwan.
7. ACKNOWLEDGMENTS

Numerous individuals and institutions have aided this study, and I extend myhearty thanks to all of them.
First and foremost, I wish to express my thanks to Prof. Sadao Takagi,
Laboratory of Systematic Entomology, Faculty of Agriculture, Hokkaido University,
for his kind guidance and constant encouragements.
Special thanks are due to Dr. Takehiko Nakane (Chiba-shi), for his great
encouragement. Moreover, the present study is partly based on his good and valued
collection.
I thank Prof. Hisashi Abe, Dr. Toshio Kumata, Dr. Masaaki Suwa, Dr. Shinichi Akimoto (Hokkaido University), Dr. Shun-Ichi Ueno (NSMT), Dr. Sadanari
Hisamatsu (Matsuyama-shi), Mr. Yukiatsu Hiroya and Mr. Hiroshige Kurihara (Otaru
Museum) for their kind guidance and encouragement.
My thanks are extended to many histeridologists who provided me with valuablespecimens for comparison and also with literature; Dr. Gerardo Arriagada Santic(Santiago, Chile), Dr. Yves Gorny (Fontenay-sous-bois, France), Dr. Oleg L.
Kryzhanovskij (St. Petersburg, Russia), Dr. Aldo Olexa (Prague, Czech), Dr. Piet
Kanaar (Leiderdorp, Netherland), Dr. Oldrich Kapler (Prague, Czech), Dr. Alexey
Tishechkin (Minsk, Russia), Dr. Alfred F. Newton (Field Museum of Nat. Hist.,
Chicago, USA), Dr. Slawomir Mazur (Warszawa, Poland), Dr. J. W. Summerlin(Texas, USA), Dr. Pierpalo Vienna (Venezia, Italy), and Dr. Tomas Yelamos(Barcelona, Spain).
I am thankful to Mr. Kayo Akiyama (Y okohama-shi), late Dr. Kintaro Baba
(Kurokawa-cho), Ms. E. De Boise (NHM), Mr. Toshitsugu Endo (Edogawa-ku), Mr.
Kaoru Haga (Kawasaki-shi), Dr. Masami Hayashi (Saitama Univ.), Dr. Hiromichi
Higuchi and Mr. Koichi Satoh (TPM), Mr. Yoshiyuki Hirano (Odawara-shi), Mr.
Shigehisa Hori (Sapporo-shi), Mr. Shoichi Imasaka (Shimabara-shi), Mr. Msaaki
Ishida (Suginami-ku), Mr. Kazuo Ijima (Shibecha-cho), Dr. Shin-ichi Kudo and Mr.
Nobuo Ishihama (Hokkaido Univ.), Messrs. Hidenori Torikura and Akeo Iwasaki(Hokkaido Plant Protection Office), Mr. Takeshi Ito (Nishinomiya-shi), Dr. Makoto
Kiuchi (Tsukuba-shi), Mr. Shigeyuki Kondo (Urayasu-shi), Dr. Yoshihisa Kusui(Nagasaki-shi), Mr. Masahiro Saito (Mikuni-cho), Dr. Mitsuhiro Maegata (Kushikinoshi), Dr. Kaoru Maeto (HFFP), Dr. Hiroshi Makihara (T6hoku Research Center,
Forestry and Forest Products Research Inst.), Dr. Shun'ichi Makino (KyOshO
272

Research Center, Forestry and Forest Products Research Inst.), Dr. Kimio Masumoto(Otsuma Women's Univ.), Mr. Eishi Matsui (Amakusa-shi), Dr. Yoshikazu Miyake(Tama-shi), Mr. Masato Mori (Nishinomiya-shi), Dr. Shinya Miyano and Dr. ShunIchiro Naomi (Nat. Hist. Museum of Chiba), Mr. Masaaki Nishikawa (Zama-shi), Mr.
Naomi Nishikawa (Sapporo-shi), Mr. Yilichi Okushima (Kurashiki Museum of Nat.
Hist.), Dr. Masahiro Sakai (Ehime Univ.), Dr. O. A. Samuelson (BSM), Mr.
Kazuhiro Sawada (Setagaya-ku), Mr. Masahiko Sato (Rishiri Twon Musuem), Dr.
Akihiko Shinohara (NSMT), Mr. Shigehiko Shiyake (Osaka Museum of Nat.
Hist.), Dr. Osamu Tadauchi, Dr. Shilhei Nomura and Mr. Teruhisa Ueno (Kyilshu
Univ.), Mr. Yasushi Takai (Seki-shi), Mr. Haruo Takizawa (Biological ResearchCenter, Japan Tabacco Inc.), Dr. Tsutomu Tanabe (Tokushima Prefectural Museum),
Mr. Minoru Tao (Yokohama-shi), Dr. Ky6ji Tazoe (Kannami-ch6), Dr. Seiki Yamane(Kagashima Univ.) and Mr. Nobuki Yasuda (S6unky6 Museum of Nat. Hist.); allthese people have kindly provided specimens from their collections, useful informationand suggestions. Without their help the data upon which this revision is based would
be much less complete. Many other individuals and institutions also have provided
specimens for this study.
Fund for this study was partly donated by the Japan Society for Promotion of
Science for Japanese Junior Scientists.
8. REFERENCES
Adachi, T., 1930. On the Japanese species of the genera Onthophilus and Hololepta (Coleoptera.
Histeridae), with description of a new species. Kontyfi, 4: 247-252.
Adachi, T. & M. Ohno, 1962. Discovery of the family Sphaeritidae (Coleoptera) in Japan. Zoological
Magazine, 71(4): 148-151.
Ancey, C. F., 1882. Contributions a Ie faune de l'Afrique orientale. Descriptions de Coleopteres
nouveaux. Natural. (paris). (4) 2: 54-55.
Arnett, R. H., 1962. The beetles of the United States. Washington, 1112 pp.
Auzat, (V.,) 1925. Travaux scientifiques de l'Armee d'Orient (1916-1918). CoIeopteres Histerides. Bull.
Mus. Hist. Nat., 31: 74-76.
Bickhardt, H., 191Oa. Histeridae. In Junk, W. & S. Schenkling (eds.). Coleopt. Cat., pars 24: 137 pp,
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