Title

ON THE SPECIES OF THE GENUS MARGARINOTUS
FROM JAPAN (COLEOPTERA : HISTERIDAE)

Author(s)OHARA, Masahiro

Citation

Insecta matsumurana. Series entomology. New series, 41: 1-50

Issue Date

1989-08

URL

http://hdl.handle.net/2115/9846

RightsTypebulletin

Additional
Information

Instructions for use

Hokkaido University Collection of Scholarly and Academic Papers : HUSCAPINSECTA MATSUMURANANEW SERIES 41

AUGUST

1989

ON THE SPECIES OF THE GENUS MARGARINOTUS FROM JAPAN(COLEOPTERA: HISTERIDAE)

By MASAHIRO OHARA

Abstract
OHARA, M. 1989. On the species of the genus Margarinotus from Japan (Coleoptera: Histeridae). Ins. matsum. n. s. 41: 1-50, 12 tabs., 21 figs.
The species of the genus Margarinotus Marseul from Japan are revised. A key and descriptions are given to ten species, of which, M. (Ptomister) yezoensis is new to science, and M. (P)cadaven'cola (Bickh.) and M. (P.) reichardti Kryzh. are newly recorded from Japan. M. (P.)
marginepunctatus is transferred from subgenus Promethister to subgenus Ptomister. The genitalia
of both the sexes, except for the female of M. (P.) reichardti, and several useful taxonomic features
are illustrated.

Author's address.
Sapporo, 060 Japan.

Entomological Institute, Faculty of Agriculture, Hokkaid6 University,

Contents
Introduction
3
Acknowledgements ..................................................................... . 3
Terminology ........................................................................... . 4
Systematics ........................................................................... . 6
Genus Margarinotus Marseul ......................................................... . 6
Key to the subgenera of Japanese Margarinotus ................................. . 7
Subgenus Ptomister Houlbert et Monnot ......................................... . 7
Key to the Japanese species of Ptomister ......................................... . 8
Species groups and species ..................................................... . 8
Boleti-group ............................................................... .
8
1. Margarinotus (Ptomister) boleti (Lewis) ................................... . 9
Weymarni-group ................................................................ 13
2. Margarinotus (Ptomister) cadavericola (Bickhardt) .......................... 13
3. Margarinotus (Ptomister) agnatus (Lewis) .................................. 15
4. Margarinotus (Ptomister) weymarni Wenzel ................................ 19
5. Margarinotus (Ptomister) reichardti Kryzhanovskij . . . . . . . . . . . . . . . . . . . . . . . . .. 22
6. Margarinotus (Ptomister) striola (C.R. Shalberg) ............................ 24
subspecies succicola (Thomson) . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 27
7. Margarinotus (Ptomister) yezoensis M. Ohara, sp. nov ....................... 27
Sutus-group . ..................................................................... 31
8. Margarinotus (Ptomister) marginepunctatus (Lewis) .......................... 31
9. Margarinotus (Ptomister) sutus (Lewis) .................................... 34
Subgenus Grammostethus Lewis .................................................. 36
10. Margarinotus (Grammostethus) niponicus (Lewis) ............................ 37
Notes on the phylogenetic relationship of some major groups within
the genus Margarinotus ........................................................ 41
References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 46
Appendix. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 49

INTRODUCTION

The genus Margarinotus was erected by Marseul (1853) with Hister scaber
Fabricius, 1787, as the type species. The genus was characterized by "configuration
singuliere du dos, OU les stries semblent remplacees par des rangees de tubercules
luisants et saiIIants, en guise de perles sur un fond obscur et rugueux" (Marseul,1853), and had long been limited to two species, the Palaearctic scaber F. and the
N earctic guttifer Horn.
Since Wenzel's (1944) study on the genus Margarinotus Mars. was published, it
has elicited much taxonomic discussion as to genera or subgenera which are related
to the genus Margarinotus Mars. (Kryzhanovskij, 1966; Mazur, 1972; Kryzhanovskij and Reichardt, 1976; Vienna, 1977 ; Olexa, 1982). This is due to the fact that heincluded in the genus the species of Paraiister, Grammostethus, Stenister, and a greatpart of the genus Hister by reason of similarities in the structure of the male
genitalia.
Nowadays many authors accept Margarinotus in Wenzel's sense as a good
genus. According to the recent world catalogue of Histeridae prepared by Mazur(1984b), the genus Margarinotus Mars. is divisible into 7 subgenera, Margarinotus
Marseul, 1853, Ptomister Houlbert et Monnot, 1923, Eucalohister Reitter, 1909,
Stenister Reichardt, 1926, Para lister Bickhardt, 1917, Grammostethus Lewis, 1906, andPromethister Kryzhanovskij, 1966, and includes 91 known species in the world.
However, phylogenetic relationships among the subgenera have not yet been revised
critically.
In this paper I recognize ten species of the genus Margarinotus Mars. fromJapan, which belong to two subgenera: Ptomister Houlbert et Monnot and Grammostethus Lewis. The subgenus Ptomister includes 9 of them; one of them is a new
species, yezoensis, and two, cadavericola and reichardti, are newly recorded. All
these species can be divided into three groups on the basis of the structure of themale- genitalia: 1) boleti-group, 2) weymarni-group, and 3) sutus-group. The
other subgenus Grammostethus is represented by one species, niponicus, in Japan.
At the conclusion, notes are given on the phylogenetic relationship among thesubgenera.
ACKNOWLEDGEMENTSFirst of all I would like to express my cordial thanks to the former Professor
Takehiko N akane of Kagoshima University, for his great encouragement to mystudy. Moreover, the present study is much based on his good and valued collection.
My thanks are also to Dr. O.L. Kryzhanovskij, Zoological Institute, Academy of
Sciences, Leningrad, USSR, for giving me the valuable information concerning M.
(P.) yezoensis. To Dr. M.E. Bacchus, British Museum (Natural History), lowe hiskind helping for the loan of Lewis's type specimens.
I am also indebted to the following entomologists for their kindness in offering
material used in this paper: Dr. K. Baba, Mr. S. Daido, Mr. H. Haga, Mr. M.
Hinakura, Mr. Y. Hirano, Mr. T. Ito, Mr. A. Iwasaki, Mr. S. Imasaka, Mr. M. Ishida,Dr. M. Kadosaki, Dr. P. Kanaar, Dr. M. Kiuchi, Mr. T. Kinoda, Dr. C. Kitamura, Mr.
S. Kondo, Mr. S. Kudo, Dr. Y. Kusui, Mr. S. Nomura, Mr. N. Nishikawa, Mr. M.
3

Nishikawa, Mr. M. Maegata, Mr. H. Makihara, Dr. S. Makino, Mr. K. Masumoto,
Mr. E. Matsui, Dr. S. Mazur, Mr. S. Mochizuki, Mr. M. Mori, Dr. S. Osawa, Mr. M.
SaitO, Mr. K. Sawada, Mr. H. Tanaka, Mr. Y. Takai, Dr. K. Tazoe, Mr. T. Tanabe,
Dr. A. Tishechkin, Mr. K. Tomiyama.
Last but not least my hearty thanks are also due to Prof. S. Takagi, Entomological Institute, Hokkaid6 University, for his kindness in reading the manuscriptand giving me his critical advices to the present study.
TERMINOLOGY

I mainly follow Wenzel and Dybas (1941) for the terminology of the striae andbody parts (Fig. I-A, B, C). Head with a frontal and a supraorbital stria.

Marginal pronotal
Outer lateral pronotal
Inner lateral pro notal

1

Oblique humeral

~::
3rd

1f-lf-HItH----4th

dorsalsMarg. epipieural---t-----'t-Mll
Marg. elytral

5thExternal subhumeral
Internal subhumeral

Lobe
Suture betw. prost. keel and lob.
Lat. margo prosternal
Prosternal keel

Lat. prosternal

Carina I

Post. margo profemoral -----it'"
Post. mesoeoxal~i7""""!'5J~;;::::::.-j\---- Marg. mesosternal
Metasternal-meseplmeral sut.

Lateral disk

Meso-metasternal sut.

Oblique metasternal

Metasternal-meteplsternal sut.

Metasternal longitudinal sut.
Intereoxal disk of 1st abdominal sternumIntereoxal disk

Fig.1. A: Dorsal view of a Margarinotus showing principal striae.
ditto. C: Ventral view, showing principal striae and parts.
4

B: Lateral view,

Pronotum with three striae which are named marginal, outer lateral, and innerlateral from outside to inside. Each elytron with eleven striae, which are named
marginal epipleural, marginal elytral, external and internal subhumeral, oblique
humeral, first to fifth dorsal, and sutural from outside to inside (the order of
numbering in Histeridae is opposite to that in most other Coleoptera).
Ventral side of prosternum with five striae, which are named marginal stria of
prosternal lobe, lateral prosternal stria, lateral marginal prosternal stria, sutural
stria between prosternal keel and lobe, and carinal stria.
Ventral side of meso- and metasternum with five striae, which are named
marginal mesosternal, meso-metasternal (suture), lateral metasternal (a-lateral striain Wenzel, 1944), oblique (b-lateral stria), and posterior mesocoxal stria (postcoxal).
Disk of metasternum is divided into two parts, which are named intercoxal andlateral. The intercoxal disk is divided on mid-line by a suture called metasternallongitudinal suture. A suture between metasternum and mesepimeron is named
metasternal-mesepimeral, and that between metasternum and metepisternum iscalled metasternal-metepisternal.
First abdominal sternum is divided into two parts, intercoxal and lateral disks.
A stria on each side of the intercoxal disk is named 1st abdominal stria. Ventral
side of profemur with two striae which are named anterior and posterior marginal... .

": .. .

I

,

r

A

"

:

J

&~4.~.
o~~~gOt

CoxlteParameres

Basal pieceE

StylusEjaculatory duct
Median armature, .........

k

Bursa copulatrlx----\\----I
Spermatheca

VaginaVulva"

2\\)/1.---Posterior apodemeMedian lobe

Fig. 2. A: Dorsal view of the aedeagus of the male genitalia of Margarinotus cadavericola, showing parts. B: Lateral view with median lobe extruded and median arma·
ture rotated upwards. C: Lateral view of the median lobe showing principal parts.
D: Dorsal view of the terminal female abdominal segments of M. agnatus showing
sclerotized principle structure. E: Lateral view (left side), ditto. F: Schematiclateral view (left side) of interior of terminal female abdominal segments. G:
Lateral view (left side) of bursa copulatrix and spermatheca of M. sln·ola.
5profemoral.
Parts of the male and female genitalia are named as in Fig. 2, A-G.
SYSTEMA TICS
GENUS MARGARINOTUS MARSEUL, 1853

Margarinotus Marseul, 1853: 549.
Type species: Hister scaber Fabricius, 1787: 32.

Wenzel's (1944) diagnosis. "Frontal and supraorbital striae present, the frontal
sometimes obsolescent. Anterior pronotal margin emarginate for the reception of
the head. One or two lateral pronotal striae present, the outer frequently abbreviated when present. External subhumeral stria nearly always complete, occasionallyabbreviated or obliterated; internal subhumeral stria absent, though a few punctures may occasionally be present apically. Dorsal striae well impressed, excepting
in scaber and guttijer. Prosternal keel seldom striate, the striae when present
usually emarginate, occasionally truncate (davisi) or nearly so. Protibiae with fourto sixteen denticles or teeth.
Aedeagus with a ring-shaped articulating basal piece; the rest of the tegmen
variable, either bifid apically on dorsal surface or tetrafid. Median lobe sclerotized,
subcylindrical, or spoon-shaped, with paired posterior apodemes; gonopore distal.
Median armature connected to the base of the median lobe by a sclerotized hinge orby a heavy membrane; median armature, when viewed form above, nearly always
forked apically, the forks sometimes fused."
The genus Margarinotus in Wenzel's sense may be recognized as a monophyletictaxon, because the species of the genus have the following characters in common
which can be interpreted as apomorphic.
1) Median lobe of the male genitalia thick, usually like a staff or spoon in
shape, with a sclerotized median armature. [ ~ This state is derivable from a thinand fiat median lobe without median armature. The supposed plesiomorphic stateis shared by other genera of the tribe Histerini, for example, Hister, Atholus,
Pactolinus and Merohister].
2) Sessil receptacles of the spermatheca of the female genitalia not slender,
usually sac-like and coiled at the base. [+- The receptacles slender, not coild at thebase.]
3) Profemoral posterior marginal stria not entire, usually present on apicalthird. [~The stria entire.]On the other hand, the following character of the genus may be plesiomorphic.
4) External subhumeral stria of each elytron entire. [ -> The stria interrupted
or abbreviated.]Distribution. Palaearctic, N earctic and Oriental Regions.
Remarks. At a glance the genus Margarinotus is very similar to other generaof the tribe Histerini, but it is a good taxon characterized by the apomorphic
characters as mentioned above. The genus may be recognized by the entire subhumeral stria of elytron, but the structure of the male genitalia affords more reliablecharacters for identification.
6

Key to the subgenera of Japanese Margarinotus
1(2)
2(1)

Pronotum with two lateral pronotal striae .
.. .. . . .. . ... . . . ..... . . .. . .. . ..... .. . . . . .. . subgenus Ptomister Houlbert et Monnot, 1923
Pronotum with one lateral pro notal stria ............. subgenus Grammostethus Lewis, 1906

Vienna (1977) gave a key of the genus Margarinotus in "New keys to the Italian
species of the tribe Histerini", but his key did not include the subgenus Margarinotus.
I add the subgenus Margarinotus to his key as follows:
1( 2)
2( 1)
3( 6)
4( 5)
5( 4)
6( 3)
7( 8)

8( 7)

9(10)
10( 9)
11(12)

12(11)

Elytra with shining and projecting tubercles......... subgenus Margarinotus Marseul
Elytra without tubercles, usually smooth and shining.
Pronotum with two lateral striae.
Meso· and meta tibiae broad, triangular, covered with long and yellow pubescence.
Epipleura of pronotum with sparse and short hairs..... subgenus Eucalohister Reitter
Meso· and metatibiae not broad, normal form without long yellow pubescence .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subgenus Ptomister Houlbert et Monnot
Pronotum with one lateral stria.
Pronotal marginal stria short, usually present on apical area, at most attaining to half
length of pronotum. Denticles of protibiae rather large. Body oval.
.................................................... subgenus Paralister Bickhardt
Pronotal marginal stria nearly entire, only abbreviated at the base. Protibiae withiarge denticles or numerous small ones. Body shape various, from oval to subcylin·
drical.
Body shape subcylindrical. All tibiae strongly dilated. Metafemur widened .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subgenus Stenister Reichardt
Body shape oval. Tibiae and metafemur normal.
Area between lateral stria and lateral margin of pronotum considerably swelled up.
Sutural stria of elytron almost entire. Prosternum without carinal stria .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . subgenus Promethister Kryzhanovskij
Area between lateral stria and lateral margin of pronotum flat. Sutural stria of
elytron shortened on basal half. Prosternum with carinal stria .
. . .. .. ... . . . . . . . ...... .. . .. . ... . . . .. . . . .. . . . . .. . subgenus Grammostethus Lewis

SUBGENUS PTOMISTER HOULBERT ET MONNOT, 1923

Type species: Hister merdarinus Hoffmann, 1803: 39.
Syn.: Eucalohister Cooman, 1947: 428, by Mazur, 1984: 164.

The subgenus Ptomister was established by Houlbert and Monnot in 1923 as oneof the subgenera of the genus Hister Linnaeus. They simply gave a diagnosis of thesubgenus in their key as follows: "Cotes du thorax finement ponctues ou lisses endessous, sans poils dresses. Fossettes antenna ires distinctes./ Mesosternum
echancre en avant. Prosternum arrondi a la base, devant l'echancrure du mesosternum./ Elytres avec une strie subhumerale (interne ou externe)./ Vne strie subhumerale externe seulement, soit entiere, soit raccourcie en arriere./ Thorax avec
deux ou trois stries laterales; epipleures unistries (au moins chez des especes
franc;aises)."
Additional diagnosis. Body 4.0-8.5 mm in length, larger than in the othersubgenera. Frontal stria of head usually well impressed. Pronotum with marginal,
outer and inner lateral striae. Sometimes densely punctate inside inner lateral stria
7and between lateral stria and lateral margin of pronotum. External subhumeral
stria of elytron complete and well impressed. Internal one usually present on apicalhalf but rudimentary. Dorsal striae variable, but usually 1st to 3rd complete, 4th
shortened basally, and 5th and sutural strongly shortened. Propygidium andpygidium usually alutaceous in ground sculpture, with large and ocelloid punctures.
Punctation usually denser on pygidium than on propygidium. Prosternal keel withor without carinal stria. Epipleura of pronotum without yellow hair. Mesosternum with marginal stria. Metasternum with lateral and oblique striae. Lateralmetasternal disk usually with coarse punctation. Intercoxal disk smooth.
Protibiae with several large denticles or many small ones. All tibiae dilated
moderately, not strongly. Mesotibiae without yellow pubescence. Median lobe ofmale genitalia various in shape. The Japanese species of the subgenus Ptomisterare divided into at least three groups by the structure of the median lobe: boletigroup, weymarni-group and sutus-group (see description of each group). Spermatheca of female genitalia consisting of several receptacles, these are sac-shaped (notslender) and usually coiled at the base.
This subgenus includes 36 known species in the world. Most of the species are
distributed in the Palaearctic and N earctic Regions, and a few species are in the
Oriental Region.

Key to the Japanese species of Ptomister
1( 2)
2( 1)
3(14)
4(13)
5( 8)
6( 7)
7( 6)
8( 5)
9(10)
10( 9)
11(12)
12(11)
13( 4)
14( 3)
15(16)
16(15)

Inner lateral pronotal stria strongly undulate behind the eyes (Fig. 3-A).
.... .... ..... ...
. .................................... boleti (Lewis, 1884)
Inner lateral pronotal stria not strongly undulate behind the eyes.
Prosternal keel without carinal stria (sometimes striola (Fig. 10-E), weymarni and reichardti
with an obsolete one).
Body larger, 5.0-9.2 mm in length.
Lateral stria of metasternum united with the oblique stria of metasternum (Fig. 7-A).
Lateral disk of metasternum with long hairs (Fig. 4- J) ..... cadavericola (Bickhardt, 1920)
Lateral disk of metasternum without hair. ......
. ........... agnatus (Lewis, 1884)
Lateral stria of metasternum not united with the oblique stria of metasternum (Fig.7-E).
Lateral disk of metasternum without hair (Fig. 9-D). . . .. . reichardti Kryzhanovskij, 1976
Lateral disk of metasternum with long hair (Fig. 7-B).
Outer lateral pronotal stria extending beyond basal end of inner one (Fig.6-L).
.. . . . .. . . . .. . . . . .. . . .
. . . .... . . . . .
. .. weymarni Wenzel, 1944
Outer lateral pronotal stria not extending beyond basal end of inner one (Fig. 12-B).
..............
. ... striola (C.R. Sahlberg, 1819)
Body smaller, 4.0-4.5 mm in length.
. ............ . yezoensis M. Ohara, sp. nov.
Prosternal keel with carina I striae (Fig. 14-J, 0).
Inside the inner pronotal stria densely with large punctures (Fig. 12- B).
. .... .... .... ...... ....
.............
. .. marginepunctatus (Lewis, 1879)
Inside the inner pronotal stria without large punctures (Fig. 12-C) ..... sutus (Lewis, 1884)

SPECIES GROUPS AND SPECIESBoleti-groupl. boleti (Lewis)

Median lobe of the male genitalia not expanded on apical half, stick-like, and8

Fig. 3. A-B: lvl. (P.) boleti (Lewis). C-F: jVi. (P.) cadavericoia (Bickhardt). A, C:
Pronotum and left elytron. B, D: Pronotum. E, F: Left elytron.

without a denticle on dorsal mid-line. Long sclerotized projection developed on
basal third on dorsal surface of median lobe (Fig. 5- D). Median armature verysmall; posterior apodeme rather short and stout.
In external characters, prosternal inner lateral stria strongly undulate behindthe eyes.
Judging from Wenzel's (1960) description and figures, M. fenderi Wenzel from
Washington, North America, seems to be included in this group.
In ecology, M. boleti is fungus-dwelling species, and M. fenderi seems to be
associated with the nest of pocket gopher. Both species are not usual carriondwelling carnivorous histerids.
This group is distributed in the Far East, northern India and the Californian
subregion.

1.

Margarinotus (Ptomister) boleti (Lewis, 1884)

Hister boleti Lewis, 1884: 135 [Japan: ChGzenjiko; Kashiwagi in the Kii Peninsula] ; Lewis,
1902: 238 [noted]; Bickhardt, 1920: 97, 99 [key] ; Kamiya and Takagi, 1938: 30 [listed].
Hister (Hister) boleti: Bickhardt, 1910: 40 [catalogued] ; Bickhardt, 1917: 179 [catalogued].
Margarinotus boleti: Wenzel, 1944: 131 [listed, key].
Margarinotus (Margarinotus) boleti: Kryzhanovskij and Reichardt, 1976: 344 [noted, key].
Margarinotus (Ptomister) boleti: Hisamatsu and Kusui, 1984: 17 [noted] ; Hisamatsu, 1984:
23 [noted]; Mazur, 1984b: 165 [catalogued]; Hisamatsu, 1985: 227, pI. 4, f.12 [key, photo,
noted] .

Description.

cf1 ~. Body length, PPL (= length between anterior angles of9

Table 1.
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Biometric data for Margarinotus boleti (Lewis).
Male

Female1.63-1.88 (1.78± 0.029) 8
3.89-4.77 (4.34±0.078) 8
2.13-2.51 (2.40±0.048) 8
2.89-3.39 (3.29 ± 0.058) 8
4.52-5.71 (5.16±0.01l) 8
2.69-3.14 (3.01±0.052) 7
1.31-1.88 (1.54± 0.073) 7
1.44-1.88 (1.65±0.053) 8
1.31-1.63 (1.54±0.033) 8
1.44-1.82 (1.64±0.044) 8
1.82-2.32 (2.06±0.056) 8

1.57-1.88 (1.82± 0.056) 5
3.70-4.52 (4.30±0.137) 5
1.82-2.57 (2.32 ± 0.121) 5
2.88-3.51 (3.31 ± 0.099) 5
4.64-5.46 (5.20±0.129) 5
2.63-3.13 (2.94±0.075) 5
1.38-1.81 (1.53± 0.072) 5
1.57-1.88 (1.71±0.057) 5
1.51-1.63 (1.55±0.021) 5
1.44-1.69 (1.57±0.047) 5
1.69-2.20 (2.03±0.081) 5

Measurements in mm. APW - width between anterior angles ofpronotum; PPW - width between posterior angles of pronotum; PLlength of pronotum in middle; EL -length of elytron along sutural line ;EW - maximal width between outer margins of elytra; ProW - maxi·
mal width of propygidium; ProL -length of propygidium in mesial;
PyL -length of pygidium; PTL -length of protibia; MSTL -lengthof mesotibia; MTTL -length of metatibia. The table reads: range(mean±standard error) number of specimens measured.

pronotum and apex of pygidium) 6.0-8.1 mm, PEL (= length between anterior angles
of pronotum and apices of elytra) 5.3-7.0 mm. Width, 4.6-5.5 mm. Biometric data
are given in Table 1. Body oblong, oval, black and shining.
Frontal stria of head (Fig. 4-A, B, C) deep, complete (sometimes interrupted atanterior angle) and usually straight at middle.
Marginal pronotal stria (Fig. 3-A, B) interrupted behind head, but complete
laterally. Outer lateral stria abbreviated at basal sixth, hooked at basal end,
sometimes the apical end connected with inner lateral stria (in this case, inner one
frequently interrupted behind anterior angle of pronotum (Fig.3-B)). Inner lateral
stria often complete laterally and anteriorly, sometimes interrupted behind anterior
angle of pronotum, and strongly bent behind the eyes. Disk sparsely covered withmicroscopic punctures, and with a longitudinal puncture in antescutellar area.
Epipleural fossette sparsely and finely punctate. Marginal elytral stria absent.
Marginal epipleural stria complete, well impressed, with large punctures. Elytra(Fig.3-A) with a slight subapical impression. External subhumeral and Ist-3rd
dorsal striae complete. Internal subhumeral stria absent. Oblique humeral stria
represented on basal fourth. Fourth and 5th dorsal and sutural striae present on
apical third, 4th usually incomplete and with a short basal rudiment. All striae,
except the oblique humeral one, strongly crenated.
Pygidia feebly alutaceous. Propygidium (Fig. 18-A) coarsely and rather sparsely punctate and scattered throughout with minute punctures, the punctures becoming
finer along the margin. Propygidial disk with a strong and broad depression
posteriorly on each side. Pygidium (Fig. 18-B) coarsely and densely punctate, thepunctures deep in middle and fine along basal margin.
Prosternal lobe rounded at apex, coarsely punctate laterally, its marginal stria
10

1Pf.a~y~v: ' ~c
I

I

r~
H

Fig. 4. A-H: M. (P.) boleti (Lewis). I-M: M. (P.) cadavericola (Bickhardt). A-C, I:
Head, frontal view. D, J: Meso-, metasternum and 1st abdominal sternum. E:
Prosternal lobe and keel. F, K: Left protibia, dorsal view. G, L: ditto, ventral
view. H, M: Left pro femur, ventral view.

interrupted at middle. Prosternal keel (Fig. 4-E) with carinal striae between coxae.
Anterior margin of mesosternum feebly emarginate, its marginal stria complete
and strongly crenate. Meso-metasternal suture (Fig.4-D) complete, obtusely angulate at middle. Lateral metasternal stria extending obliquely and posteriorly.
Oblique stria extending inwardly from the middle of the metasternal-metepisternal
suture. Intercoxal disk of metasternum sparsely clothed with microscopic punctures. Lateral disk of metasternum coarsely and shallowly punctate, and scatteredwith fine punctures throughout, without hair.
Intercoxal disk of 1st abdominal sternum (Fig.4-D) striate on each side on
11/([=::~~~~4
G"@----,------------------

M
K

H:::",..

~~ \WL ~ N/'-.....

--------'"----Fig. 5. A-F: M. (P.) boleti (Lewis) [Nikko, HonshO], G-P: M. (P.) cadavericola (Bickhardt) [Owari, HonshO], A, G: Aedeagus of male genitalia, lateral view. B, H :ditto, dorsal view. C: Apex of aedeagus, ventral view. D, I, K: Median lobe, lateral view. E, J: ditto, dorsal view. L: Median armature, caudal view. M: ditto,
dorsal view. N: ditto, lateral view. F, 0, P: Spermatheca and' bursa copulatrix,
lateral view (right side). Scale: 1.0 mm.
12apical two-thirds.
Protibia (Fig. 4-F, G) with 5-6 denticles on outer margin and 2 denticles on
apical margin of ventral face. Profemur (Fig. 4- H) with posterior marginal stria on
apical fifth.
Male genitalia (Fig. 5-A, B, C, D, E): aedeagus with a ring-shaped articulated
basal piece. Median lobe slender, with short paired posterior apodemes. Medianarmature small (Fig. 5-D).
Female genitalia (Fig. 5-F): spermatheca consisting of 11 receptacles.
Specimens examined, 90"', 4~ and 25 exs. Hokkaido-2 exs., Nopporo, Ebetsu, 1l/viiij1973,
Y. Sakamoto leg. (Historical Museum of Hokkaido, coll.); 2 exs., Fukushima, Osima, 1l/viiij1976,
M. Kiuchi leg. HonsM - <Aomori-ken> 10"' and 1 ex., Shimokita, Yunomata, 2/viii/1956, S.
Ashida leg.; 1 ex., Towada, 3/viiij1954, A. Abe leg. <Miyagi-ken> 10"' and 1 ~, Natori, Akiho, 24/
vij1951, K. Nagayama leg. <Fukushima-ken> 4 exs., Nakanomoriyama, Tsushima, Namie,
Futaba, 14/ix/1974, K. Tazoe leg. <Tochigi-ken> 10"' and 2 exs., Nikko, Chuzenji, 22/viij1972, T.
N akane leg. <Tokyo-to> 1 ex., Takao-san, Tama, 5/viii/1961, S. Kondo leg. <Kanagawa-ken> 2~, Hakone, 2/viiij1970, K. Masumoto leg.; 1 ~, Mt. Tanzawa, 5/viiij1983, M. Nishikawa leg_; 1ex., Dodaira, Eastern Tanzawa, 22/viiij1982, M. Nishikawa leg. (Nagano-ken> 20"' and 1 ex.,Kiso, Komagatake, 4-5/viiij1946, S.Osawa leg. <N ara-ken> 10"' and 5 exs., Mt. Yoshino, 4/ix/
1952, K. Sakaguchi and K. Sawada leg. <Osaka-fu> 1 ex., N akanotani, Mt. Iwawaki, 17/viiij1948,
S. Ueno leg. <Fukui-ken> 10"', Karikomi-ike, Koike, Ono City, 2/viij1978, M. SaitO leg.
(Hiroshima-ken> 1 ex., Hikimi, Yoshiwamura, 7/viiij1983, T. Ito leg. Shikoku - (Tokushimaken> 1 ex., Higashi-iyayama, 8/viiij1961, M.Hirai leg. KyusM-<Miyazaki-ken> 10"', Takachihonomine, 21/viij1951, F_ Takahashi leg. <Kagoshima-ken> 10"', Kurino, Kirishima, 29/v /1983, S.
Daido leg.; 2 exs., ditto, 31/viiij1985, M. Ohara leg.

Distribution (Fig. 20). Japan (Hokkaid6, HonshU, Shikoku, KyOshU), Kuril IsIs.
(Dr. Kryzhanovskij, pers. com.); Taiwan. Newly recorded from Hokkaido andKuril IsIs.
Remarks. This species is easily distinguished from the other Japanese Margarinotus species by the strongly bent inner pronotallateral stria (Fig. 3-A). M. boleti
occurs commonly on fungi or in decaying trees.

Weymarni-group
2. cadavericola (Bickhardt)
3. agnatus (Lewis)
4. weymarni Wenzel

5. reichardti Kryzhanovskij
6. striola (C.R. Shalberg)
7. yezoensis M. Ohara, sp. nov.

Median lobe of the male genitalia expanded on apical half, spoon-like; posterior
apodeme short and stout; shape of median armature various. Apex of lobe feebly
bent backward, but not trilobate in dorsal view.
Judging from illustrations given by authors (see Appendix), the following species
seem to be placed in this group as understood above: brunneus (Fabricius, 1775),
distinctus (Erichson, 1834), fractifrons (Casey, 1893), multidens (Schmidt, 1889), rectus(Casey, 1916), and tristriatus Wenzel, 1944. This group is distributed in the Palaearctic and N earctic Regions.

2.

Margarinotus (Ptomister) cadavericola (Bickhardt, 1920)
Hister cadavericola Bickhardt, 1920: 99 [China: Fo-ken] ; Reichardt, 1930b: 46 [noted];13

Reichardt and Kryzhanovskij, 1964: 172 [China: Kuatun].
Margarinotus cadavericola: Wenzel, 1944: 126, 132 [listed, key].
Margarinotus (Margarinotus) cadavericola: Kryzhanovskij and Reichardt, 1976: 339, f. 645,657[key, noted, figured].
Margarinotus (Ptomister) cadavericola: Mazur, 1984b: 165 [catalogued].
Hister ussuriensis Reichardt, 1930: 285, pI. 8, f. 1 [key, figured] ; Kryzhanovskij and Reichardt,
1976: 339.
Margarinotus ussuriensis: Wenzel, 1944: 126, 132, pI. 8, f.l [key, figured].

Description. cf1 i'-. Body length, PPL 5.2-7.2 mm, PEL 4.7-6.5 mm. Width, 4.15.7 mm. Biometric data are given in Table 2. Body oblong-oval, black and shining.
Frontal stria of head (Fig. 4-1) complete and crenulate.
Marginal pronotal stria (Fig. 3-C, D) interrupted behind head and complete
laterally. Outer lateral stria complete and feebly crenulate. Inner lateral stria
complete, straight anteriorly, and strongly crenate. Disk of pronotum sparsely
covered with microscopic punctures, the punctures being dense and coarse between
two lateral striae, and with a longitudinal puncture in antescutellar area.
Elytra (Fig. 3-C) with a feeble subapical impression, and microscopically punctulate throughout. Epipleural fossette densely and coarsely punctate. Marginal
elytral stria absent. Marginal epipleural stria complete. External subhumeral
stria complete. Internal subhumeral stria represented on apical fourth by a row ofmoderate-sized punctures or absent. Oblique humeral stria distinctly present on
basal third. First-4th dorsal striae complete, but 4th stria shortly abbreviated atbase. Fifth dorsal stria present on apical fourth, and its basal rudiment represented
by a short arc which consists of a few (from 1 to 5) punctures or sometimes absent.
Sutural stria present on apical third, sometimes extending to middle. Sometimes
dorsal striae connected with neighboring striae (Fig. 3-E, F).
Pygidia (Fig. I8-C, D) alutaceous. Propygidium densely covered with ocelloid
punctures, and minutely punctulate among ocelloid ones. Disk of propygidium witha depression posteriorly on each side. Punctation of pygidium similar to that ofpropygidium, but denser than the latter.
Prosternal lobe broadly truncate at apex, its marginal stria interrupted atTable 2.

Biometric data for Margarinotus cadavericola (Bickhardt).

Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Male

Female1.56-2.01 (1.79±0.049) 10
3.51-4.83 (4.14±0.131) 10
1.81-3.07 (2.18±0.115) 10
2.51-3.39 (2.95 ±0.092) 10
4.14-5.65 (4.87±0.162) 10
2.51-3.45 (2.97± 0.098) 10
1.13-1.94 (1.41±0.084) 10
1.32-1.88 (1.57±0.07l) 10
1.32-1.82 (1.55±0.054) 10
1.38-1.88 (1.65±O.053) 10
1.69-2.51 (2.08±0.084) 10

1.88-2.13 (2.00±0.043) 6
4.39-4.89 (4.60±0.069) 6
2.07-2.51 (2.32±0.061) 6
3.14-3.58 (3.35±0.073) 6
5.08-5.65 (5.36±0.074) 6
3.01-3.45 (3.25±O.054) 6
1.38-1.76 (1.52 ± 0.048) 6
1.69-2.01 (1.88±0.039) 6
1.51-1.82 (1.60± 0.044) 6
1.57 -1.88 (1.69±O.042) 6
2.07 -2.45 (2.22 ± 0.048) 6

14

middle. Prosternal keel without carinal stria.
Anterior margin of mesosternum (Fig. 4- J) strongly emarginate at middle, its
marginal stria complete and crenate. Disk of mesosternum sparsely clothed withmicroscopic punctures, the punctures coarser laterally. Meso-metasternal suturecomplete, obtusely angulate at middle. Lateral stria of metasternum extending
posteriorly and obliquely, and united with the oblique stria which inwardly extends
from the middle of metasternal-metepisternal suture. Intercoxal disk of metasternum sparsely clothed with microscopic punctures, the punctures becoming coarser
laterally. Lateral disk of metasternum densely covered with large and shallow
punctures, and with long hairs.
Intercoxal disk of 1st abdominal sternum (Fig. 4-J) striate on each side, andcoarsely punctate laterally.
Protibia (Fig. 4-K, L) with 6 denticles on outer margin and 2-4 fine denticles on
apical margin of ventral face. Profemur (Fig. 4- M) with posterior marginal stria on
apical fourth.
Male genitalia: as illustrated in Fig.5-G, H, I, J, K, L, M, N.
Female genitalia: as shown in Fig. 5-0, P.
Specimens examined, 11 d". 7 Sf- and 26 exs. Hokkaid6 - 2d' and 1 Sf-, Sapporo, 3, 20/ix/1985,
20/ix/1986, M. Ohara leg.; 23 exs., Toyotaki, Sapporo, 3, 12/ix/1987, M. Ohara leg.; 2d', Miyatsu,
Okushiri Is., 3/ix/1986, Y. Kusui leg. Honshu - <Aichi·ken> Id', Higashiyama, Nagoya, 18/ix/
1948, T. Nakane leg.; 1 ex., ditto, 20/v/1948, T. Nakane leg.; 5d', I Sf- and 1 ex., ditto, vi/1948, S.
Osawa leg.; 1 ex., ditto, 27/vi/1948, S.Osawa leg.; 1 Sf-, Nagoya, Owari, 19/viii/1941, S.Osawa
leg.; 1 Sf-, Mt. Kong6, Owari, 2/vii/1941, S. Eda leg. <Nara-ken> 2 Sf-, Mt. Kasuga, 29/ix/1946, F.
Takahashi leg.; 1 Sf-, ditto, 22, 23/ix/1968, S. Kond6 leg. Kyushu - <Saga-ken> 1 d', Tashiro, nodate, Y. Miyake leg.

Distribution (Fig. 20). Japan (Hokkaid6, Honshu, Kyushfl); China; USSR
(Primorskij Kray, Kuril Isis.). Newly recorded from Japan.
Remarks. This species is closely related to M. agnatus, M. weymarni, M. striata
and M. reichardti, but it is distinguished from them by the characters given in the
key. These five species are difficult to distinguish in their external characters.
Japanese records of these species may probably contain errors and confusions andare in need of reexamination.
In Hokkaid6 this species appears in early autumn.

3.

Margarinatus (Ptamister) agnatus (Lewis, 1884)

Hister agnatus Lewis, 1884: 135 [Japan: Nikk6].
Hister (Hister) agnatus: Bickhardt, 1910: 40 [catalogued]; Bickhardt, 1917: 178[catalogued] ; Kamiya and Takagi, 1938: 30 [listed].
Margarinotus agnatus: Wenzel, 1971: 216.
Margarinotus (Margarinotus) agnatus: Kryzhanovskij and Reichardt, 1976: 338, f. 644 (not 647)
[key, noted].
Margan'notus (Ptomister) agnatus: Hisamatsu and Kusui, 1984: 16 [noted] ; Mazur, 1984b:
164 [catalogued] ; Hisamatsu, 1985: 227, pI. 41, f, 10 [key, photo, noted].
Margarinotus balloui Wenzel, 1944: 129, pI. 6, f. 2 [Japan: Nikk6].
Margan'notus ballowi [sic] : Kryzhanovskij and Reichardt, 1976: 338,
Hister balloui: Mazur, 1972: 140.

Description. c?' ~. Body length, PPL 5.4-8.3 mm, PEL 4.4-6.9 mm. Width, 4.25.3 mm. Biometric data are given in Table 3. Body oblong-oval, black and shining.
15

Table 3.
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Biometric data for Margarinotus agnatus (Lewis).
Male

Female1.57-1.88 (1.73±0.023) 12
3.64-4.08 (3.85±0.039) 12
1.88-2.07 (l.95±0.023) 12
2.45-3.01 (2.77 ±0.046) 12
4.20-4.89 (4.49±0.053) 12
2.38-3.07 (2.83±0.048) 12
1.19-1.38 (1.28 ± 0.020) 12
1.44-1.63 (1.55± 0.023) 12
1.19-1.51 (1.40±0.022) 12
1.38-1.63 (1.54± 0.023) 12
1.88-2.20 (2.01±0.026) 12

1.82-2.01 (1.89 ± 0.020) 12
3.89-4.52 (4.25±0.047) 12
1.94-2.38 (2.l9±0.033) 12
2.95-3.32 (3.l6± 0.033) 12
4.71-5.27 (5.02±0.049) 12
3.01-3.45 (3.23±0.036) 12
1.19-1.63 (1.41± 0.035) 12
1.63-2.01 (1.80±0.029) 12
1.32-1.57 (1.50± 0.022) 12
1.44-1.76 (1.60 ±0.025) 12
1.94-2.51 (2.l4± 0.042) 12

Frontal stria of head (Fig. 6-I, J, K) complete (sometimes interrupted at middle),
usually feebly and inwardly bent at middle.
Marginal pronotal stria (Fig. 6-A, F, G, H) interrupted behind head and complete
laterally. Outer lateral stria complete or interrupted, feebly crenulate and hooked
at apical end. Inner lateral stria complete, straight anteriorly, and strongly crenated. Disk of pronotum sparsely and finely punctate, the punctures becoming denser
laterally. Densely covered with large punctures between outer lateral stria andinner one, but the punctures variable, sometimes quite reduced. Not punctate
between margin of pronotum and outer lateral stria. Usually with a longitudinalpuncture in antescutellar area.
Elytra (Fig. 6-A, B, C, D, E) with a feeble subapical impression, and microscopically punctulate throughout. Epipleural fossette coarsely punctate. Marginal
elytral stria absent. Marginal epipleural stria complete. External subhumeral
stria complete. Internal subhumeral stria sometimes represented on apical fourth
by a row of punctures. First-4th dorsal striae usually complete, but 4th slightly
shortened at the base. Fifth dorsal stria present on apical third, and its basal
rudiment represented by a short arc. Oblique humeral stria present on basal third.
Sutural stria present on apical half.
Pygidia (Fig. 18-E, F) micro-alutaceous. Propygidium with an indistinct
impression on each side, densely covered with ocelloid punctures, and with a few finepunctures intermingled among ocelloid ones, the punctures becoming finer at posterior margin. Pygidial punctation similar to propygidial one, but rather dense.
Prosternal lobe rounded at apex, its marginal stria broadly interrupted at
middle, alutaceous in ground sculpture, and coarsely punctate laterally. Prosternal
keel rather broad, without carinal stria.
Anterior margin of mesosternum (Fig.7-A) deeply emarginate at middle, its
marginal stria complete. Disk of mesosternum sparsely clothed with microscopic
punctures. Meso-metasternal suture complete, angulate at middle. Lateral striaof metasternum extending obliquely and posteriorly, and united with the obliquestria which extends inwardly from the middle of the metasternal·metepipleural
suture. Lateral disk of metasternum densely covered with large, round and shallow
16

~Ccc
G

H

o
Fig. 6. A-K: M. (P.) agnatus (Lewis). L-O: M. (P.) weymarni Wenzel. A,L:
Pronotum and left elytron. F-H: Pronotum. B-E, M-O: Left elytron. I-K, P, Q:
Head, frontal view.

17

B

F

\/

--

: --,

G

n

~
D
Fig. 7. A-D: M. (P.) agnaius (Lewis). E-I: M. (P.) weymarni Wenzel. A, E: Meso-,
metasternum and 1st abdominal sternum. B, F, G: Left protibia, dorsal view. C, H :ditto, ventral view. D, I: Left profemur, ventral view.

punctures, without hair.
Intercoxal disk of 1st abdominal sternum (Fig.7-A) striate on each side, andmoderately punctate laterally.
Protibia (Fig. 7- B, C) with 5-7 denticles on outer margin, and 2 or 3 fine denticles
on apical margin. Profemur (Fig. 7-1) with posterior marginal stria very short,present only apically.
Male genitalia: as shown in Fig. 8-A, B, C.
Female genitalia: as in Fig. 8-D.
Specimens examined, 97 c?', 75 i'- and 114 exs. Hokkaid6 -1 ex., Akaiwa, Otaru, 7/viii/1986,
A. Iwasaki leg.; 7c?', 7 i'-, and 62 exs., N opporo, 14, 17,27 /vi/1986, 16,18,22, 29/v, 5,7, 12, 16, 19, 20,
23, 26/vi, 3, 4/vii, 20/viii/1987, M. Ohara leg.; 1 ex., Atsubetsu, Sapporo, 8/v /1987, M. Ohara leg. ;2 exs., }6zankei, Toyotaki, 10/v, 3/vi/1987, M. Ohara leg.; 1 ex., Mt. Hakken, Sapporo, 13/vi/
1977, N. Nishikawa leg.; 2 exs., Heiwa, Nishiku, Sapporo, 12/viii/1987, M. Ohara leg.; 1 ex., Oasa,
18

Sapporo, 12/ix/1987, M. 6hara leg.; 1 ex., Sapporo, 16/v/1974, M. Kiuchi leg.; 8 exs., Memuro, 24/
vi/1986, M.6hara leg.; 1 ex., Maruseppu, Kitami, 7/viii/1987, M.6hara leg.; 1d", Okushiri Is,
Miyatsu, 3/ix/1986, Y. Kusui leg. HonshO - <Fukushima-ken> 2 exs., Bunadaira, Hinoemata, 5/
vii/1987, K_ Haga leg_ (Tochigi-ken> 1 ex., Nikko, 12/v/1985, M. Kiuchi leg. <Tokyo·to> 2 st-,Takao-san, 30/v /1974, M. Ishida leg.; 1 ex., ditto, 4/vii/1971, K. Masumoto leg.; 1 ex., Kagenobu,
30/v/1970, M. Ishida leg.; 1 ex., Okutama, Odake, 7/vi/1980, K. Haga leg. <Kanagawa-ken> 1 ex.,6yama, 23/v/1969, M. Ishida leg. <Yamanashi-ken> 1 ex., Yatsugatake, 16/vii/1973, M. Ishidaleg. ; 13 exs., Mt. Daibosatsu, 3, 4/vii/1976, S. Kondo leg.; 1 ex., Hikawa-rindo, Daibosatsu, 29/vi/
1980, K. Haga leg. (Shizuoka-ken> 82 d" , 47st-, Nagatsuro, Itani, 6/v/1985, A. Yamamoto leg.
<Niigata-ken> 1 ex., Mt. Myoken, Sado, 21/viii/1985, K. Baba leg.; 1 ex., Shinhosan, Asahimura,Northern Echigo, 1/vi/1987, K. Baba leg. (Nagano-ken> 1 ex., Mt. Jonen, 900 m, 2/vi/1960, K.
Kamimura leg.; 2 st- and 1 ex., Kamikochi, 21-28/vii/1974, S.6sawa leg.; 1 ex., Jigokudani,
Yamanouchi, 29/iv /1983, M. Kiuchi leg. <Aichi-ken> 1st-, Higashiyama, Nagoya, vi/1948, S. 6sawaleg. <Kyoto-fu> 1st-, lex., Mizuho, Tozu-river, 10/v/1986, S.Nomura leg. <Hyogo-ken> 1st-,
Koya-ike, 25/x/1941, K. Sakaguchi leg. ; 3 exs., Taizan-ji, Kobe, 5/v /1987, T. Ito leg. <Tottoriken> lex., Yonago, 12/x/1987, M.6hara leg. Shikoku-(Ehime-ken> 3d" and 1st-, Omogo, 30/
vii/1986, T. Tanabe leg. KyOshO - <Miyazaki-ken> 1 d", Takachihonomine, 29/v /1983, M.6haraleg. ; 4 d" and llst-, ditto, 10/vii/1984, M.6hara leg.; 1 ex., ditto, 29/v /1983, S. Daid6 leg.; 2 exs.,
ditto, 19/vii/1983, S. Sameshima leg. (Kagoshima-ken> 1 ex., Kirishima, 19/vii/1984, T. Kinoda
leg.

Distribution (Fig. 20). Japan (Hokkaid6, Honshfi, Shikoku, Kyfishfi);
Himalaya; North India. New to Hokkaid6.
Remarks. This species occurs commonly in animal carrion. Hisamatsu (1985)
reported that this species is distributed in western Japan, but I found it from
Hokkaid6 and examined many specimens from eastern Japan.
The male genitalia shown by Kryzhanovskij and Reichardt (1976) as of thisspecies actually belong to weymarni.

4.

Margarinotus (Ptomister) weymarni Wenzel, 1944

Margarinotus weymarni Wenzel, 1944: 127, pI. 4, f. 2, pI. 6, f.3 [North Manchuria: Erhtaohotze, Kirin, and Japan: Masike, Nikko] : Wenzel, 1971: 216; Nakane, 1963: 70, pI. 35, f.19 [photo,
noted].
Margarinotus (Margarinotus) weymarni: Kryzhanovskij and Reichardt, 1976: 338, f.647 (not644), 666 [noted].

Table 4.
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Biometric data for Margarinotus weymarni Wenzel.
Male

Female1.57-2.01 (1.83±0.025) 20
3.07 -4.58 (3.99 ± 0.075) 20
1.76-2.51 (2.08±0.043) 20
2.32-3.51 (3.02±0.058) 20
3.70-5.27 (4.65±O.083) 20
2.32-3.39 (3.00±O.065) 20
0.94-1.51 (1.27±0.030) 20
1.19-1.88 (1.62±0.034) 20
1.13-1.63 (1.50 ± 0.030) 20
1.32-1.88 (1.60 ± 0.043) 18
1.57-2.32 (1.99±0.038) 20

1.82-2.20 (1.98 ± 0.029) 16
3.76-4.96 (4.41 ±0.079) 16
1.82-2.70 (2.22±0.053) 16
3.01-4.02 (3.46±0.065) 16
4.45-5.77 (5.19±0.083) 16
3.01-3.76 (3.34±0.053) 16
1.19-1.63 (1.38±0.030) 16
1.63-2.13 (1.90±0.034) 16
1.44-1.88 (1.61 ± 0.029) 16
1.38-2.01 (1.66±0.041) 16
1.88-2.57 (2.17± 0.044) 16

19

Margarinotus (Ptomister) weymarni: Hisamatsu and Kusui, 1984: 16 [listed, noted] ; Mazur,
1984b: 169 [catalogued] ; Hisamatsu, 1985: 227, pI. 4, f.10 [key, photo].
Hister weymarni: Mazur, 1972 : 140.

Description. c?'~. Body length, PPL 5.0-7.9 mm, PEL 4.3-6.5 mm. Width,3.75.S mm. Biometric data are given in Table 4. Body oblong-oval, black and shining.
Frontal stria of head (Fig. 6-P, Q) complete and crenulate.
Marginal pronotal stria (Fig.6-L) interrupted behind head, complete laterally.
Outer lateral stria complete. Inner lateral stria complete and strongly crenate,sometimes abbreviated at basal fourth. Inner lateral stria not ending beyond thebasal end of outer one. Disk of pronotum sparsely covered with microscopic
punctures, the punctures being coarse between two lateral striae, and usually with alongitudinal puncture at antescutellar area.
Elytra (Fig. 6- L, M, N, 0) with a feeble subapical impression. Epipleural fossette densely and coarsely punctate. Marginal elytral stria absent. Marginal
epipleural stria complete. External subhumeral stria complete. Internal subhumeral stria indistinct, represented on apical third by a row of punctures. Oblique
humeral stria present on basal third. First-4th dorsal striae complete but 4th stria
shortly abbreviated at base. Fifth dorsal stria usually present on apical third or alittle less, its basal rudiment represented by an arc. Sutural stria usually present on
apical third, sometimes extending to middle.
Pygidia (Fig. IS-G, H) alutaceous. Propygidium densely covered with large andocelloid punctures, and minute punctures intermingled among the punctures. Diskof propygidium depressed near each posterior angle. Punctation of pygidium similar to that of propygidium, which is somewhat denser.
Prosternal lobe rounded at apex, its marginal stria complete, or sometimes
interrupted at middle. Disk of prosternal lobe alutaceous, and coarsely punctate
laterally. Prosternal keel without carinal stria.
Anterior margin of mesosternum (Fig. 7-E) strongly emarginate at middle, its
marginal stria complete and crenate. Disk of mesosternum sparsely clothed withmicroscopic punctures, the punctures dense and coarse laterally. Meso-metasternal
suture complete and obtusely angulate at middle. Lateral stria of metasternum
extending posteriorly to near, but not united with, the oblique stria which extends
inwardly from the middle of metasternal-metepisternal suture. Intercoxal disk ofmetasternum sparsely clothed with microscopic punctures, the punctures coarse
along the oblique stria of metasternum. Lateral disk of metasternum densely
covered with round, shallow and large punctures, and with long hairs.
Intercoxal disk of 1st abdominal sternum (Fig.7-E) striate on each side, anddensely and moderately punctate laterally.
Protibia (Fig. 7-F, G, H) with 5-6 denticles on outer margin, and 2 denticles on
apical margin. Profemur (Fig. 7-1) with posterior marginal stria short, present
apically.
Male genitalia: as in Fig. S-E, F, G.
Female genitalia: as in Fig. S- H, I.
Specimens examined, 18d", 15 ~ and 41 exs. Hokkaid6 -1 ex., Otaru, 10/vii/1985, M. Ohara
leg. ; 2 exs. ditto, vii/1985, Y. Kusui leg.; 2 exs., Zenibaka, Otaru, 19/vi/1979, N. Nishikawa leg.; 1ex., Tsukisappu, Sapporo, 26/viii/1986, S. Makino leg.; 2d", 4 exs., Sapporo, 7, 16, 23/v, 19/vi/1985,
M. Ohara leg.; 1 ex, ditto, 16/v /1974, M. Kiuchi leg. ; 3d", 3 ~, 9 exs., Nopporo, 12,14,17,27 /vi/
20

G

Fig. 8. A-D: M. (P.) agnatus (Lewis) [Kagoshima, Kyftshft]. E-H: M. (P.) weymarni
Wenzel [Sapporo, Hokkaid6]. A, E: Aedeagus of male genitalia, lateral view. B,
F: ditto, dorsal view. C, G: Median lobe, lateral view. D, I, H : Spermatheca andbursa copulatrix, lateral view (right side). Scale: 1.0 mm.

1986, 7,10, 16/v, 5, 23/vi, 3,28/vii/1987, M. Ohara leg.; 1 ex., Atsubetu, Sapporo, 8/v/1987, M.
Ohara leg.; 2 exs., Toyotaki, Sapporo, 10/v /1987, M. Ohara leg.; 1 ex., Mt. Hakken, Sapporo, 13/
vi/1977, N. Nishikawa leg.; 40", Maruyama, Sapporo, 10/vii/1942, N. Aoyama leg.; 1 ex., ditto, 5/v /1973, S. Kud6 leg.; 10", 1 ex., Memuro, 24/vi/1986, M. Ohara leg.; 1 ~, Daisetsu, 7/vi/1971, M.
Sako leg.; 1 ex., Tomakomai, 30/v/1976, A. Kashizaki leg.; 10", ditto, 11/vi/1988, M. Hinakuraleg. Honshft- <Yamagata· ken> 10", 1 ~, Sakata, 9, 11/vii/1948, K. Shirah at a leg. <Fukushima·ken> 1 ex., Wakamatsu, 1/vi/1947, M.W. leg.; 1 ex., Oyama, Wakamatsu, 6/v/1938, Y. Kurosawa
leg. <Ibaraki·ken> 1 ~, Mito, Hitachi, 16/iv /1938, S. Eda leg. <Chiba·ken> 10", Oami, 3/iv /1982,
21

M. Ohara leg. ; 2 exs., Abiko, Tonegawa-dote, 24/x/1971, M. and K. Ishida leg. <Tokyo-to> 1 ex.,
Setagaya, 18/ix/1949, Ohira leg. <Kanagawa-ken> 1 f, Kannonzaki, 16/x/1965, K. Masumotoleg.; 1 ex., Kamigo, 6/iv/1974, K. Masumoto leg. <Niigata-ken> 1 ex., Itoigawa, Southern Echigo,
9/x/1956, K. Baba leg.; 1 ex., Mt. Kinpoku, Sado, 27 /v /1936 M. Kaneko leg.; 1 ex., Senami,Northern Echigo, l/v /1985, K. Baba leg. <N agano-ken> 10", 1 f, Kamikochi, 7/viii/1938, K.
Sakaguchi leg.; 1 ex., ditto, 21-28/vii/1947, S. Osawa leg. <Gifu-ken> 10", Ogaki, Mino, 10/v /1946,
Y. Moriya leg. <Aichi-ken> 1 ex., Obata, Owari, 30/v /1947, S. Osawa leg.; 10", 1 f Shikatsumura,
Owari, 12/iv/1946, S. Osawa leg.; 1 f, Higashiyama, Owari, 20/v /1946, T. N akane leg. <Mie-ken>
If, Tsu, Ise, 23/iii/1939, K. Sakaguchi leg. <Osaka-fu> 10", Ikuecho, 2/vi/1935, no collector's
name; 1 ex., N ishinariku, 15/v/1950. K. Sawada leg.; 1 f, 2 exs., ditto, 3,4/viii/1949, K. Sawada
leg. <Hyogo-ken> 1 ex., Mayasan, i8/ix/1937, no collector's name; 2f, Motoyamamura, 14/v/
1938, 16/x/1941, K. Sakaguchi leg. <Wakayama-ken> 10", Kii-tomitagawa, 29/iii/1937, no collector's name; 1 ex., Minabe, 17/iii/1943, K. Sakaguchi leg. KyQshQ - <Kagoshima-ken> 1 f,
Shigeta, Kamitakakumacho, Kanoya, 22/v/1983, Y. Takai leg. USSR, Sakhalin-lei", If,
Konuma, 3/vii/1946, Y. Nishio leg.

Distribution (Fig. 20). Japan (Hokkaid6, HonshO, Shikoku, KyOshO); Northeast China (Manchuria); USSR (Khabarovskij and Primorskij Kray).

5.

Margarinotus (Ptomister) reichardti Kryzhanovskij, 1976

Margarinotus (Margarinotus) reichardti Kryzhanovskii in Kryzhanovskii and Reichardt, 1976:
339 [Primorskii Kray].
Margarinotus (Ptomister) reichardti: Mazur, 1984b: 168 [catalogued].
Hister parilis Reichardt in Kryzhanovskii and Reichardt, 1976: 341 (nom. nud.).

Description. J'I. Boby length, PPL 4.9-5.4 mm, PEL 5.5-6.1 mm. Width, 4.3-5.0
Biometric data are given in Table 5. Body oblong-oval, black and shining.
Head (Fig. 9- B) even, sparsely and finely punctate. Frontal stria complete,
feebly and inwardly bent or straight at middle. Labrum transverse oblong.
Mandible strongly developed and finely punctate.
Marginal pronotal stria (Fig. 9- A) broadly interrupted behind head and complete
laterally. Outer pronotal stria complete, but not attaining basal margin, and feebly
crenate. Inner lateral stria complete, nearly straight anteriorly and strongly
crenate. Disk of pronotum sparsely and finely punctate. A longitudinal puncturemm.

Table 5. Biometric data for Margarinotus reichardti Kryzhanovskii.
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Male
1.57 -1.76 (1.66± 0.067) 2
3.64-4.08 (3.86±0.155) 2
1.94-2.45 (2.20±0.018) 2
2.51-2.82 (2.67 ± 0.111) 2
4.33-4.96 (4.64±0.222) 2
2.57-2.89 (2.73±0.1ll) 2
1.25-1.44 (1.35± 0.067) 2
1.32-1.63 (l.47±0.111) 2
1.32-1.44 (1.38±0.044) 2
1.32-1.63 (1.47±0.1l1) 2
1.88-2.01 (1.94± 0.044) 2
22

Fig. 9. M. (P.) reichardti Kryzhanovskij [Fukushima, HonshO]' A: Pronotum and left
elytron. B: Head, frontal view. c: Prosternallobe and keel. D: Meso·, metaster·
num and 1st abdominal sternum. E: Right profemur, ventral view. F: Right
protibia, ventral view. G: ditto, dorsal view. H: Aedeagus of male genitalia,
lateral view. I: ditto, dorsal view. J: Median lobe, dorsal view. K: ditto, lateralview. Scale: 1.0 mm.

present in antescutellar area.
Elytra (Fig.9-A) with a subapical impression, and sparsely covered withmicroscopic punctures. Epipleural fossette densely and coarsely punctate. Marginal epipleural stria complete. Marginal elytral stria absent. External subhumeral and 1st-4th dorsal striae complete, but 2nd and 4th slightly shortened at the base.
Internal subhumeral stria absent. Fifth dorsal stria present on apical third, its
23apical end united with apical end of 4th. Basal rudiment of 5th represented by ashort arc on basal third. Sutural stria present on apical two·thirds. Oblique
humeral stria present on basal third. All striae (except oblique humeral stria)
strongly crenate. Anterior margin of elytra strongly crenate.
Pygidia (Fig. IS-I, J) micro·alutaceous. Propygidium feebly impressed on eachside moderately covered with ocelloid puncture, and with a few fine puncturesintermingled between ocelloid ones. Pygidial punctation similar to propygidial one,but rather more densely punctate.
Prosternal lobe (Fig. 9-C) rounded at apex, its marginal stria interrupted at
middle, and its disk coarsely punctate, the punctures becoming larger laterally.
Prosternal keel with obsolete carinal stria on basal area.
Anterior margin of mesosternum (Fig. 9-D) feebly emarginate at middle, its
marginal stria complete. Disk of mesosternum sparsely covered with microscopic
punctures throughout, and coarsely punctate along the inner side of marginal stria.
Inside apical angles coarsely punctate. Meso·metasternal suture complete, an·gulate at middle and slightly crenate. Lateral stria of metasternum extending
obliquely and posteriorly, and not united with the oblique stria which extends
inwardly from the middle of the metasternal-metepipleural suture. Intercoxal diskof metasternum sparsely clothed with microscopic punctures throughout and coarse
punctures along the lateral stria. Lateral disk of metasternum densely covered
with large, round and shallow punctures, without hair.
Intercoxal disk of 1st abdominal sternum (Fig.9-D) striate on each side, andcoarsely punctate laterally.
Protibia (Fig. 9-F, G) with 7 denticles on lateral margin and 2 denticles on apicalmargin. Base of dentides weakly projected. Distance between denticles narrow.
Profemur (Fig. 9-E) with posterior marginal stria very short, present apically.
Male genitalia: as in Fig. 9- H, I, J. K.
Specimens examined, 20". HonshO-(Fukushima·ken> 10", Tsuchiyu, Azumayama, Fuku·
shima City, 21/vi/1979, K. Tazoe leg.; <Iwate-ken> 10", Kazubezawa, Shimohei, 20/vi/1983, A.
Izumi leg.

Distribution (Fig. 20). Japan (HonshU); USSR (Primorskij and southern
Khabarovskij Kray). Newly recorded from Japan.
Remarks. This species is distinguished from M. weymarni, M. agnatus, M.
striola, and M. cadavericola by the more numerous denticles of protibiae (S-9 dentides) and by the narrower distance between the denticles. In the latter four denticles
are 4 to 7 in number (Kryzhanovskij and Reichardt, 1976). For correct
identifications, however, the structure of the male genitalia should be examined.

6.

Margarinotus (Ptomister) striola (C.R. Sahlberg, lS19)

Hister striota C.R. Sahlberg, 1819: 25 [North Finland] ; Lewis, 1899: 17; Reitter, 1909: 283;
Auzat, 1914: 171, f. 1,2,7; Horion, 1949: 358.
Hister (Hister) striota: Bickhardt, 1910: 50 [catalogued] ; Bickhardt, 1917: 186 [catalogued] ;Bickhardt, 1920: 99.
Margarinotus striota: Wenzel, 1944: 126 [listed].
Margarinotus (Margarinotus) striota: Kryzhanovskij and Reichardt, 1976: 342, f. 676-679, 682[key, figured].
Margarinotus (Ptomister) striota: Hisamatsu and Kusui, 1984: 17 [noted] ; Mazur, 1984b: 16824

[catalogued] ; Hisamatsu, 1985: 227 [key].
Hister eschscholtzii Dejean, 1837: 140 (nom. nud.).
Hister eschscholtzii: Marseul, 1854: 282, t. viii, f. 81.
Hister carbonarius ; Dejean, 1837: 140.
Hister japanus Motschulsky, 1860: 13; Marseul, 1862: 700; Lewis, 1884: 135 [Yokohama,
Honshu; misidentification?] ; Lewis, 1895: 188.
Margarinotus striolides Wenzel, 1944: 129, pI. 7, f.l [Tunkun, Sajan Mountains, Siberia] ;Wenzel, 1971: 216.
Hister striolides: Mazur, 1970: 59; Mazur, 1972: 140.

Description. cf1 ~. Body length, PPL 5.5-8.3 mm, PEL 4.7-6.6 mm. Width,3.8-4.7
Biometric data are given in Table 6. Body oblong, oval, black and shining.
Frontal stria of head (Fig. lO-C, D) deep, complete and inwardly angulate at
middle, sometimes interrupted at middle.
Marginal pronotal stria (Fig. IO-A) broadly interrupted behind head and com·plete laterally. Outer lateral stria abbreviated basally in various degrees, ending
usually at basal two· thirds and sometimes at apical third or near posterior marginof pronotum, but not extending beyond end of inner lateral stria. Inner lateral stria
complete and crenulate. Disk of pronotum sparsely covered with microscopic
punctures, and usually with a longitudinal large puncture in antescutellar area.
Epipleural fossette of elytra densely and coarsely punctate. Marginal elytral
stria absent. Marginal epipleural stria complete. Elytron with two slight depres·sions at base of 3rd dorsal and oblique humeral striae. External subhumeral stria
complete. Internal subhumeral stria (Fig.IO-A) absent or obscurely indicated on
apical third. First-3rd dorsal striae complete. Fourth dorsal stria variously abo
breviated on basal sixth to third. Fifth dorsal stria present on apical third.
Sutural stria present on apical half, sometimes on apical third. All striae, except
the oblique humeral one, feebly crenulate.
Pygidia (Fig. 19-A, B) alutaceous. Propygidium with an indistinct depression
at each side, densely covered with coarse punctures and minute punctures inter·mingled. Punctation of pygidium similar to propygidial one.
Prosternallobe truncate at apex, sparsely and moderately punctate, its marginalmm.

Table 6.

Biometric data for Margarinotus striola striola (C.R. Shalberg).

Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Male

Female1.57-1.88 (1.72±0.022) 19
3.26-4.14 (3.81±0.057) 19
1.51-2.07 (1.88±0.032) 19
2.51-3.14 (2.90±0.041) 19
3.83-4.71 (4.42±0.063) 19
2.26-2.95 (2.68± 0.054) 19
0.94-1.32 (1.15±0.025) 19
1.25-1.57 (1.44 ±0.019) 19
1.19-1.51 (1.36±0.018) 19
1.13-1.69 (1.43 ± 0.031) 19
1.57-2.38 (1.88±0.042) 19

1.69-2.01 (l.88±0.022) 13
3.70-4.45 (4.23±0.055) 13
1.82-2.20 (2.08±0.025) 13
2.95-3.58 (3.31±0.044) 13
4.39-5.21 (4.99±0.063) 13
2.70-3.32 (3.13±0.046) 13
1.19-1.44 (1.33±0.020) 13
1.44-1.82 (1.69± 0.027) 13
1.38-1.63 (1.49 ± 0.020) 13
1.44-1.76 (1.61± 0.024) 13
1.88-2.26 (2.07± 0.027) 13

25

D

G ~

E

J

"--_ _ _ _ _ _ _ ~ I

h

'. .-

...
.

(j

0',

~.'.

L

.

.

~/~'."...
F;:---------Fig. 10. A-I: M. (P.) striata (C.R. Shalberg) [Nopporo, Hokkaid6]' A: Pronotum andleft elytron. B: Pronotum. C-D: Head, frontal view. E: Prosternal lobe andkeel. F: Meso-, metasternum and 1st abdominal sternum. G: Left protibia, dorsal
view. H: ditto, ventral view_ I: Left pro femur, ventral view. J-L: M. (P.) striala
succicala (Thomson) [Byelorussia, USSR] J: Median lobe of male genitalia, lateralview. K: Median armature, lateral view. L: ditto [Netherland]. Scale: 1.0 mm.

stria complete (Fig. 10-E). Prosternal keel sparsely and moderately punctate, witha large puncture on each side at basal third of prosternum, and sometimes with(usually without) carinal stria between coxae.
Anterior margin of mesosternum (Fig.lO-F) strongly emarginate at middle,
straight laterally, its marginal stria complete. Disk of mesosternum sparsely andfinely punctate, the punctures becoming larger laterally, and with a large fovea
26

behind each anterior angle. Meso-metasternal suture complete, obtusely angulate
at middle. Lateral metasternal stria not united with the oblique stria of metasternum which extends inwardly from the middle of the metasternal-metepisternal
suture. Intercoxal disk of metasternum sparsely and finely punctate, the punctures
becoming larger laterally. Lateral disk densely covered with large, round andshallow punctures, and with hairs.
Intercoxal disk of 1st abdominal sternum (Fig. 10-F) completely striate on eachside, and largely punctate laterally.
Protibia (Fig. 10-G, H) with 8-9 denticles on outer margin, and 3 small denticles
on apical margin of ventral face. Profemur (Fig. 10-I) with posterior marginal stria
on apical fourth.
Male genitalia: as in Fig.ll-A, B, C.
Female genitalia: as in Fig. 15-A.
Specimens examined, 170", 8 ~ and 27 exs. Hokkaid6 -10", Mt. Kariba, 14/vi/1986, S.
Nomura leg.; 10", and 2~, Sapporo, no date, S. Matsumura leg.; (Hokkaid6 University collec·tion); 1 ex., ditto, 27/vi/19l3, S.Issiki leg. (Hokkaid6 University collection); 150", 6~, 26 exs.,
Nopporo, 12, 14/vi/1986, 8,10,11; 16, 22, 29/v, 5,7,12, 19/vi/1987, M. Clhara leg.

Distribution (Fig. 20). Japan (Hokkaid6); North Europe; Siberia; North
Manchuria; Korea.
Remarks. This species is distributed over the Palaearctic region. As far as I
know, however, this species is limited to Hokkaid6 in Japan. Lewis (1884) reported
this species as Hister japanus from Yokohama, HonshO, but what he recorded might
be another species (see under M. cadavericola).
Subspecies Margarinotus (Ptomister) striola succicola (Thomson, 1862)
Hister succicala Thomson, 1862: 224 [South Sweden].
Hister striala succicala: Mazur, 1973: 59.
Margarinatus succicala: Vienna, 1977: 40.
Margarinatus striala: Wenzel, 1944: 126, pI. 7, f.2.
Margarinatus (Margarinatus) striala succicala: Kryzhanovskij and Reichardt, 1976: 342 (succi·
claa [sic] in key, p. 332).
Hister striala vaT. gavayi Auzat, 1920: 4.

The subspecies succicola is distinguished from the subspecies striola by the shapeof the median armature of male genitalia (Fig. 10-J, K, L). Biometric data are given
in Table 7.
Specimens examined, 60", 5~. Europe-20", 1~, Netherland, Rijssen (Ov.), Hall. Schwarz·
wald, 27/vi/1974, A. van Assen leg.; 40", 4~, USSR, Byelorussia, Minsk reg., 25 Km, W·Myadel,
12/v, 1/vi/1985, A. Tishechkin leg.

Distribution.
7.

Central and South Europe, Central Asia.

Margarinotus (Ptomister) yezoensis M. Ohara, sp. nov.
Margarinatus (Ptamister) sp. : Clhara, 1988: 63 [noted].

Description. cl'.5f.. Body length, PPL 4.0-4.5 mm (4.0 mm in holotype), PEL
3.72-4.0 mm (3.72 mm in holotype). Width, 3.07-3.46 mm (3.11 mm in holotype).
Biometric data are given in Table 8. Body oval, black and shining. Funicle andclub of antennae, tarsi and maxilla reddish brown.
27

Table 7.

Biometric data for Margarinotus striota succicola (Thomson).

Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Male

Female1.57-1.76 (1.67±0.029) 5
3.39-3.95 (3.71±0.084) 5
1.69-1.88 (1.81± 0.031) 5
2.63-3.01 (2.86±0.065) 5
3.95-4.67 (4.36±0.114) 5
2.32-2.95 (2.66± 0.104) 5
1.00-1.25 (1.12 ±0.041) 5
1.31-1.51 (1.41±0.029) 5
1.19-1.44 (l.30± 0.037) 5
1.19-1.44 (1.31± 0.040) 5
1.57-1.88 (1.76±0.053) 5

1.63-1.88 (1.77 ± 0.045) 5
3.70-4.33 (4.05±0.119) 5
1.76-2.13 (1.98±0.065) 5
2.82-3.32 (3.11±0.086) 5
4.45-5.08 (4.79±0.1l6) 5
2.63-3.20 (3.01 ± 0.089) 5
1.10-1.38 (1.27 ±0.047) 5
1.51-1.82 (1.66±0.052) 5
1.25-1.51 (1.38± 0.040) 5
1.25-1.57 (1.43±0.057) 5
1.69-2.01 (l.89±0.062) 5

Head (Fig. 12- E) fiat, sparsely and finely punctate. Frontal stria well impressed, complete, crenate and shortly straight at middle. Labrum transversely oblong,densely and finely punctulate. Mandible strongly developed, and densely covered
with fine punctures. Inner side of mandible concave on basal half to fit labrum.
Marginal pronotal stria (Fig. 12-A) usually broadly interrupted behind head bya space variable from 0.05-0.78 mm (in holotype, 0.05 mm), and complete laterally.
Outer lateral stria complete. Inner one complete and strongly crenate. Disk of
pronotum distinctly and finely punctate throughout, sometimes with coarse punctures in lateral area along inner lateral stria (Fig. 12-D), and usually with a longitudinal puncture in antescutellar area.
Elytra (Fig. 12-A) with a feeble subapical impression, and sparsely and finely
punctate. Epipleural fossette with a few fine punctures. Marginal elytral stria
absent. Marginal epipleural stria complete. External subhumeral stria complete.

Table 8.
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Biometric data for Margarinotus yezoensis M. Ohara.
Male

Female1.10-1.22 (1.17± 0.021) 4
2.64-2.76 (2.67±0.022) 4
1.30-1.54 (1.42 ± 0.042) 4
1.93-2.13 (2.05±0.037) 4
3.07 -3.31 (3.17± 0.045) 4
1.77-2.05 (1.90±0.051) 4
0.71-0.83 (0.78±0.026) 4
1.02-1.18 (1.10± 0.028) 4
0.83-1.02 (0.93 ± 0.035) 4
0.91-1.06 (0.96±0.030) 4
1.22-1.38 (1.28±0.030) 4

1.14-1.26 (1.20 ±0.022) 4
2.68-2.95 (2.81 ± 0.056) 4
1.42-1.65 (1.50± 0.048) 4
1.97 -2.17 (2.09±0.042) 4
3.11-3.46 (3.29 ±0.072) 4
1.85-2.09 (l.99±0.043) 4
0.83-0.94 (0.88±0.021) 4
1.14-1.30 (1.20±0.030) 4
0.91-1.02 (0.95 ± 0.021) 4
0.94-1.02 (0.98±0.014) 4
1.26-1.38 (1.33± 0.021) 4

28

Inner subhumeral stria absent. Oblique humeral stria present on basal third.
First-3rd dorsal striae complete. Fourth dorsal stria shortly abbreviated at base.
Fifth dorsal stria present on apical half, its basal rudiment represented by a long arc
on basal fifth. Sutural stria abbreviated at basal third.
Scutellum an equilateral triangle.
Propygidium (Fig. 19-C) finely depressed at sides, without alutaceous ground
sculpture, and densely covered with large, deep and ocelloid punctures, the punctures
becoming smaller at apex. Fine punctures sparsely scattered between the ocelloid
punctures. Punctation of pygidium (Fig. 19-D) similar to propygidial one.
Prosternal lobe rounded at apex, its marginal stria complete. Disk of lobe
alutaceous, and coarsely punctate laterally. Prosternal keel rather broad, without
carinal stria.
Anterior margin of mesosternum (Fig. 14-E) feebly emarginate at middle, its
marginal stria complete. Disk of mesosternum sparsely clothed with microscopic
punctures. Meso-metasternal suture complete, and obtusely angulate at middle.
Lateral stria of metasternum extending posteriorly and obliquely for about half thelength of the metasternum. Oblique stria of metasternum extending inwardly fromthe middle of metasternal-metepisternal suture. Intercoxal disk of metasternum
sparsely clothed with microscopic punctures. Lateral disk densely covered withlarge and shallow punctures, and with short hairs.
Intercoxal disk (Fig. 14-E) of 1st abdominal sternum striate on each side, andcoarsely punctate along latero-posterior margin.
Protibia (Fig. 14-A, B, C) with a minute denticle on apical margin, and 6-7
denticles on outer margin, the basal three minute. Ventral surface of profemur(Fig. 14-D) sparsely and coarsely punctate, and posterior marginal stria present on
apical fourth.
Male genitalia: as in Fig.ll-D, E, F, G.
Female genitalia: as in Fig. 15- B.
Specimens examined. Holotype- ei", Nopporo, Hokkaido, Japan, 31/vii/1986, M. Ohara leg.
Para types - 5ei", 3 -'f- and 4 exs. Hokkaido - 4ei" and 1-'f-, N opporo, 31/vii/1986, M. Ohara leg.; 1ex., ditto, 20/viiij1987, M. Ohara leg.; 1 ex., Sapporo, vii/1943, N. Aoyama leg. Honshi1<Kanagawa-ken> 1 ex., Asase, Tanzawa, 7/v/1972, Y. Hirano leg. <Niigata-ken> lei", 1 ex.,YGkyG-zan, Nagaoka, 15/ix/1985, M. Nishikawa leg.; 1-'f-, ditto, 7/x/1985, K. Baba leg. <Fukuiken> 1-'f-, Takefu City, 5/v /1936, K. Hatsuda leg.

Distribution (Fig. 21). Japan (Hokkaido, HonsM).
Remarks. M. (P.) yezoensis is closely related to M. (P.) distinctus (Erichson)
and M. (P.) wenzelianus Kryzhanovskij. It may be distinguished from the latter
two by the following characters: 1) Aedeagus is quite different in structure; 2)
Mandibles are flattened (convex in distinctus); 3) Fifth dorsal and sutural striae of
elytron are long and have a curved basal rudiment (short and without such arudiment in the latter two species); 4) Punctation of pygidium is somewhat larger(Dr. O.L. Kryzhanovskij, pers. com.).
Among the Japanese species of Margarinotus, this new species is similar to M.
sutus in body size and general external characters. However, it is more closely
related to M. weymarni, M. striola, M. cadavericola and M. agnatus in regard to themale genitalia. The new species is distinguished from M. sutus by the 5th elytral
dorsal stria having a basal rudiment. The striation of this species is similar to that
29

B

E~

M

o __ ~~~:J----+-=__

L
Fig.l1. A-C: M. (P.) striata (C.R. Shalberg) [Nopporo, Hokkaid6]. D-G: M. (P.)
yezaensis M. Ohara [Nopporo, Hokkaid6]. H- J: M. (P.) marginepunctatus (Lewis)[Osaka, Honshfi]. K-N: M. (P.) sutus (Lewis) [Fukushima, HonshU]' A, D, H, K:
Aedeagus of male genitalia, lateral view. B, E, I, L: ditto, dorsal view. C, F, J, M :Median lobe, lateral view. G, N : ditto, dorsal view. Scale: 1.0 mm.

30

of M. weymarni, but the two species can be separated from each other by the bodysize and the punctation of the pygidium.
The holotype of this species is deposited in the Entomological Institute, Facultyof Agriculture, Hokkaid6 University, Sapporo, and one paratype in the collection of
Zoological Institute, Academy of Sciences, Leningrad, USSR, in that of Hokkaid6
National Agricultural Experiment Station, Sapporo and also in that of Dr. T.
Nakane.

Sutus-group
8. marginepunctatus (Lewis)

9.

sutus (Lewis)

Median lobe of the male genitalia not expanded on apical half, stick-like, and its
dorsal mid-line with a large denticle on apical third; median armature usuallyarrowhead-shaped in lateral view (Fig. 11-], M, N).
No other species of this group are known. Species of the subgenus Eucalohisterare very similar to this group in regard to the male genitalia (see Phylogenetic
notes).
Distribution: Japan and the Far East.

8.

Margarinotus (Ptomister) marginepunctatus (Lewis, 1884)

Hister marginepunctatus Lewis, 1884: 461 [Japan: Hakodate].
Hister marginipunctatus [sic] : Lewis, 1905: 29; Lewis, 1909: 297 [ecological notes].
Hister (Hister) marginipunctatus [sic]: Bickhardt, 1910: 45 [catalogued]; Kamiya and Takagi, 1938: 31 [listed].
Hister (Grammostethus) marginipunctatus [sic] : Bickhardt, 1917: 191 [catalogued].
Margarinotus marginipunctatus [sic] : Wenzel, 1944: 126 [listed].
Margarinotus marginepunctatus: Hisamatsu and KU8ui, 1984: 17 [noted].
Margarinotus (Ptomister) marginepunctatus: Hisamatsu, 1985: 227, pI. 4, f.14 [key, photo].
Margarinotus (Promethister) marginepunctatus: Mazur, 1984b: 177 [catalogued].

Table 9. Biometric data for Margarinotus
marginepunctatus (Lewis).
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Female1.26-1.46 (1.36± 0.070) 2
2.83-2.99 (2.91±0.056) 2
1.54-1.61 (1.57 ± 0.028) 2
2.17-2.36 (2.26±0.070) 2
3.27 -3.54 (3.41 ±0.097) 2
1.97-2.28 (2.l3±0.1l1) 2
0.83-0.94 (0.89±0.042) 2
1.14-1.22 (1.18 ± 0.028) 2
1.14-1.14 (1.14±0.000) 2
1.18-1.26 (1.22 ± 0.028) 2
1.42-1.61 (1.52±0.070) 2
31

Description. cf'I 'f.. Body length, PPL 4.3-4.8 mm. PEL 3.95 mm. Width, 3.33.5 mm. Biometric data are given in Table 9. Body oblong· oval, moderately convex. Body black and shining, but legs, maxillary palpi and funicle brownish red.
Frontal stria of head (Fig. 12-F) complete, sometimes narrowly interrupted at
middle. Head sparsely and finely punctate.
Marginal pronotal stria (Fig. 12-B) interrupted behind head, and abbreviated on
basal two-thirds or half laterally. Outer lateral stria complete. Inner lateral stria
complete, strongly crenate. Disk of pronotum densely and coarsely punctate inside
the inner lateral stria, and wholly covered with microscopic punctures.
Epipleural fossette with some large punctures, especially densely along outermargin. Marginal epipleural stria complete, continuously with large punctures.
Marginal elytral stria absent. Elytra (Fig. 12- B) with a slight subapical impression.
External subhumeral and 1st-3rd dorsal striae complete, well impressed. Internal
subhumeral stria absent. Oblique humeral stria present on basal fourth. Fourth
dorsal stria complete, sometimes abbreviated at base (in holotype, complete). Fifth
dorsal and sutural striae present on apical half.
Propygidium (Fig. 19-E) densely covered with coarse and shallow punctures, anda few minute punctures intermingled, the punctures finer along margin. Punctationof pygidium (Fig. 19-F) similar to propygidial one, minute at apex.
Prosternal lobe (Fig. 14- J) rounded at apex, densely and finely punctate, thepunctures becoming coarser laterally, its marginal stria complete and deeply impressed. Prosternal keel with long carinal stria, which are well impressed between theprocoxae.

Fig. 12. A, D, E: M. (P.) yezoensis M. Ohara. B, F: M. (P.) marginepunctatus (Lewis).
C, G: M. (P.) sutus (Lewis). A-C: Pronotum and left elytron. D: Pronotum. EG: Head, frontal view.

32

Anterior margin of mesosternum (Fig. 14-1) feebly emarginate at middle, its
marginal stria complete. Disk of mesosternum punctulate finely and sparsely.
Meso-metasternal suture complete and well impressed. Lateral metasternal stria
hooked at apical end, extending obliquely and posteriorly, not united with the obliquestria which extends obliquely and medially from two-thirds of the metasternalmetepisternal suture. Intercoxal disk of metasternum smooth and finely punctate(in holotype with a large excavation on lateral side). Lateral disk of metasternum
covered with large, round and shallow punctures, and without hair.
Intercoxal disk of the 1st abdominal sternum (Fig. l4-I) finely and sparsely
punctulate, strongly striate on each side.
Protibia (Fig. 14-F, G) with 7 denticles on outer margin, its apical margin
straight. Ventral surface of profemur (Fig. 14-H) densely with round and large
punctures. Posterior marginal stria short and present apically.
Male genitalia: basal piece: paramera =1: 5.5. Medial lobe with 1 denticle on
dorsal mid-line (Fig. 11- H, I, J).
Female genitalia: spermatheca divided into 8 receptacles, of which the basal
two are small (Fig. 15-D).
Specimens examined, 2ef' and 2 f. Holotype (Fig. 13-A, B) - ef', Hakodate, Hokkaid6 [Lewis
collection, British Museum (N atural History), No. 1926-369]. Honsht1- <Aomori-ken> If, Mutsu
City, Osorezan, 27/viiij1983, S. Yamauchi leg. <Gifu-ken> If, Takayama City, ll/v/1968, T.
Nohira leg. <Osaka-fu> lef', Minomo, 13/iv/1919, no collector's name [Kyushu University collection].

Distribution (Fig. 20). Japan (Hokkaid6, HonshU, Shikoku).
Remarks. This species is included in the subgenus Promethister Kryzhanovskij,
1966, by Mazur (l984b). I agree with Hisamatsu (1985) in his opinion that it shouldbe transferred to the subgenus Ptomister Houblert and Monnot, 1922, because it has
the following characters: (1) club of antenna clearly segmented and without long
hair; (2) pronotum with two lateral striae; (3) male genitalia of this species aredifferent from those of M. (Promethister) prometheus (Kryzhanovskij) and M. (Pro.)
marginatus (Erichson) and rather similar to those of the subgenus Eucalohister
Reitter (and also to those of sutus). This species and M. sutus differ from thesubgenus Eucalohister in lacking long yellow pubescence on dorsal side of the mesoand metatibiae (see Phylogenetic notes).
This species is easily distinguished from all the other Japanese species of theFig. 13. M. (P.) marginepunctatus (Lewis).
B: Labels of Holotype.

A: Holotype (British Museum), dorsal view.
33

tribe Histerini by having coarse punctation inside the inner lateral pro notal stria.
9.

Margarinotus (Ptomister) sutus (Lewis, 1884)

Rister sutus Lewis, 1884: 136 [Japan: Kiga, near Miyanoshita] ; Mazur, 1970: 59 [Korea].
Rister (Rister) sutus: Bickhardt, 1910: 50 [catalogued] ; Bickhardt, 1917: 186 [catalogued] ;Kamiya and Takagi, 1938: 30 [listed].
Margarinotus (Margarinotus) sutus : Kryzhanovskij and Reichardt, 1976: 344 [key, noted].
Margarinotus (Ptomister) sutus : Hisamatsu and Kusui, 1984: 16 [listed] : Mazur, 1984b: 168[catalogued] ; Hisamatsu, 1985: 227, pI. 4, f. 13 [key, noted, photo].

Description. cf'I ~. Body length, PPL 3.75-5.2 mm, PEL 3.2-4.3 mm. Width,
2.7-3.6 mm. Biometric data are given in Table 10. Body oblong, oval, black andshining.
Frontal stria of head (Fig. 12-G) complete, well impressed and straight.
Marginal pronotal stria (Fig. 12-C) broadly interrupted behind head, complete
laterally. Outer lateral stria complete. Inner lateral stria complete, feebly sinuate
at middle laterally. Disk of pronotum sparsely covered with microscopic punctures(rarely with moderate punctures inside the inner lateral pronotal stria).
Epipleural fossette of elytra densely and largely punctate. Marginal elytral
stria absent. Marginal epipleural stria complete, with large punctures. Elytra
sparsely clothed with microscopic punctures. External subhumeral (Fig. 12-C) andIst-3rd dorsal striae complete. Internal subhumeral stria absent. Fourth dorsal
stria abbreviated on basal fourth. Fifth shorter than sutural, and present on apicalfourth. Sutural stria present on apical fourth. Oblique humeral stria present on
basal third.
Pygidia (Fig. 19-G, H) feebly alutaceous. Propygidium densely covered withocelloid punctures, and minute punctures scattered between ocelloid ones. Punctation of pygidium similar to propygidial one, this becoming finer apically.
Prosternal lobe (Fig.14-0) rounded at apex, alutaceous, and sparsely andmoderately punctate, its marginal stria complete. Pro sternum with alutaceous
sculpture on apical half, sparsely punctate on basal half, and a large puncture atTable 10.
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Biometric data for Margarinotus sutus (Lewis).
Male

Female1.10-1.26 (1.19 ± 0.023) 5
2.36-2.80 (2.57±0.066) 5
1.22-1.50 (1.38 ± 0.043) 5
1.85-2.20 (2.06±0.062) 5
2.72-3.19 (2.94±0.079) 5
1.38-1.93 (1.69 ± 0.080) 5
0.63-0.87 (0.74±0.036) 5
0.94-1.14 (1.02 ±0.030) 5
0.94-1.10 (1.02 ±0.029) 5
0.98-1.10 (1.03± 0.021) 5
1.14-1.46 (1.31±0.045) 5

1.22-1.41 (1.33±0.030) 5
2.72-3.11 (2.92±0.061) 5
1.38-1.65 (1.53± 0.044) 5
2.24-2.48 (2.37± 0.038) 5
3.15-3.58 (3.35±0.068) 5
1.85-2.20 (2.01±0.053) 5
0.83-0.98 (0.91±0.029) 5
1.14-1.38 (1.24± 0.035) 5
0.98-1.18 (1.11±0.030) 5
1.06-1.18 (1.13±0.020) 5
1.38-1.50 (1.44±0.024) 5

34

or·

00

•

. •• . 0 0 .'

00

.

"

"0"

I

Fig. 14. A-E: M. (P.) yezoensis M. Ohara. F-]: M. (P.) marginepunctatus (Lewis). K0: M. (P.) sutus (Lewis). A, S, F, K: Left protibia, dorsal view. C, G, L: ditto,
ventral view. D, H, M: Left pro femur, ventral view. E, I, N: Meso·, metasternumand 1st abdominal sternum. ],0: Prosternal lobe and keel.

middle of prosternal keel on each side. Carinal stria present on basal half of keel
on each side.
Anterior margin of mesosternum (Fig. 14-N) feebly emarginate, its marginalstria complete and crenate. Area between the marginal stria and antero-lateralangle shortly striate on each side. Meso-metasternal suture complete, obtusely
angulate at middle. Lateral metasternal stria running along meso-metasternal
suture, extending posteriorly and obliquely, its apical end not united (sometimes
united) with the oblique stria, which extends inwardly from the middle of themetasternal-metepisternal suture. Intercoxal disk of metasternum sparsely cov35

ered with microscopic punctures. Lateral disk densely covered with large, roundand shallow punctures, without hair.
Intercoxal disk of 1st abdominal sternum (Fig. 14-N) completely striate on eachside, and punctate along lateral and posterior margin.
Protibia with 7-10 denticles (Fig. 14-K, L) externally, its apical angle bearing
three denticles. Ventral surface of pro femur (Fig. 14-M) alutaceous and coarsely
punctate. Posterior marginal femoral stria short and present apically.
Male genitalia: median lobe with 1 denticle on dorsal mid line (Fig.ll-K, L, M,
N).

Female genitalia: spermatheca divided into 5 receptacles (Fig. 15-C).
Specimens examined, 40", 4~ and 5 exs. HonshO-<Fukushima·ken> 10", 1~ and 3 exs, Mt.
Monden, Odayama, 15/v/1947, K. Nagayama leg.; 10", ditto, 8/vi/1947, K. Nagayama leg.; 10"
and 1~, Nr.Wakamatsu, Ikki, 1l/v/1947, Y.Kurosawa leg.; 1 ex., Nuruyu, Azumayama, 23/v/
1961, K. Tazoe leg.; 1~, Noji, Azumayama, 8/v/1963, K. Tazoe leg. <Tochigi·ken> 10", Mokk6,
Nikk6, 17/vii/1981, H. Tanaka leg. <T6ky6·to) 1~, Suginami, 16/iv/1982, M. Ishida leg. KyO·
sM - <Kumamoto· ken> 1 ex., Kurokawa, 2/iii/1983, H. Aramaki leg.

Distribution (Fig. 21). Japan (HonshU, Shikoku, KytishU); Korea.
Remarks. This species is closely related to M. marginepunctatus in the median
lobe of male genitalia (see M. marginepunctatus).
SUBGENUS GRAMMOSTETHUS LEWIS, 1906
Type species: Hister ruficornis Grimm, 1852: 222.
Syn.: Coprister Houlbert et Monnot, 1923: 23 (part), by Cooman, 1947: 428.

The subgenus Grammostethus was established by Lewis in 1906 as a genus, with
Hister ruficornis Grimm designated as the type. Later, Bickhardt (1910) treated theIf'

I ,', ,.I! r

A

~

\ i'

B

~

(~

C ~Id

~

7r jl," (Fig, 15, A: M, (P,) striota (C.R, Shalberg) [Nopporo, Hokkaid6]. B: M, (P,) yezoensis
M,6hara [Nopporo, Hokkaid6]' C: M. (P,) sutus [Fukushima, HonshO]' D: M.
(P ,) marginepunctatus (Lewis) [Aomori, HonshO]. A-D: Spermatheca and bursa
copulatrix, lateral view (right side), Scale: 0,5 mm.
36

genus as a subgenus under the genus Hister Linnaeus. However, Lewis (1915) did
not follow him.
In 1944, Wenzel expanded the genus Margarinotus Mars. to include Grammostethus as a synonym. Kryzhanovskij and Reichardt (1976) revised the species ofMargarinotus occurring in the USSR, and recognized Grammostethus as a subgenus
under Margarinotus. Up to now, many authors have followed them.
Lewis's (1906) description. "Body oval or shortly oval, convex; head, funiculusof the antenna not widening out behind, the club is similar to that figured for H
merdarius in Marseul's monograph, labrum length and breadth nearly equal, mandibles somewhat long and convex above, frontal stria well marked, semicircular or
bowed in outline; thorax with one lateral stria strong and complete, usually somewhat sinuous, antenna I fossa in the anterior angle, almost circular and open to view
below; elytra, fourth dorsal stria with a detached basal appendage; prosternum
bistriate; mesosternum, anterior edge feebly sinuous or nearly truncate; anterior
tibia with many small denticles."Additional description: Body small, 3.0-6.0 mm in length, usually convex andshining. Pronotum with complete marginal stria and 1 (inner) lateral stria. External subhumeral stria of elytron complete. Internal one absent. First-4th dorsal
usually complete, and 5th and sutural striae shortened basally. Usually 5th dorsal
stria with basal rudiment in an arc. Propygidium and pygidium alutaceous, andcovered with coarse and ocelloid punctures. Prosternal keel usually with carinal
striae. Epipleura of pronotum without yellow hair. Protibiae usually with manysmall denticles and distance between denticles narrow. Meso- and metatibiae
without yellow pubescence on dorsal surface. Profemoral posterior marginal stria
present on apical sixth. Spermatheca of female genitalia consisting of several
receptacles, the receptacles sac-shaped (not slender).
Two types of the male genitalia have been recognized. 1) When median lobe is
extruded, apical fourth of tegmen is bent downwards; apical half of median lobe
expanded and spoon-like. This type is represented by ruficornis Grimm and curvicollis Bickhardt. 2) Apical part of tegmen not bent downwards; median lobe
slender. This type is found in niponicus Lewis. The genitalia of the other speciesof this subgenus have not been studied.
This subgenus is composed of 17 known species, and most of them are distributed in the Oriental Region and a few in the Palaearctic Region.
10.

Margarinotus (Grammostethus) niponicus (Lewis, 1895)

Hister navus: Marseul, 1873: 224 [recorded from Nagasaki, KyOshO, Japan].
Hister nanus [sic] : Reitter, 1879: 209 [eastern Siberia].
Hister niponicus Lewis, 1895: 188 [Japan: "found in all the islands"]; Lewis, 1900: 282[synonymy noted].
Grammostethus niponicus: Lewis, 1906: 401; Lewis, 1915: 55 [Taiwan, Arisan].
Hister (Grammostethus) niponicus: Bickhardt, 1910: 52 [catalogued]; Bickhardt, 1913: 172[Taiwan: Hoozan] ; Bickhardt, 1917; 191 [catalogued]; Desbordes, 1919: 396; Kamiya and
Takagi, 1938: 31 [listed] ; Osawa and Nakane, 1951: 6, f. 6 [described, illustrated].
Margarinotus niponicus: Wenzel, 1944: 126 [listed] ; Nakane, 1963: 70, pI. 35, f.14 [photo,
noted].
Margarinotus (Grammostethus) niponicus : Kryzhanovskij and Reichardt, 1976: 367, f.757-762[noted, key, illustrated]; Nakane, 1981: 10 [listed]; Hisamatsu and Kusui, 1984: 16 [listed,37

noted] ; Mazur, 1984b: 176 [catalogued] ; Hisamatsu, 1985: 227, pI. 4, f. 15 [key, noted, photo].

Description. d'~. Body length, PPL 3.1-5.0 mm, PEL 2.9-4.35 mm. Width,
2.5-3.7 mm. Biometric data are given in Table 11. Body oblong-oval, black andshining.
Frontal stria of head (Fig. 16-B, C, D) complete and sometimes inwardly bent orinterrupted at middle.
Marginal pronotal stria (Fig. 16-A) broadly interrupted behind head, and complete laterally. Lateral pronotal stria complete, crenulate, and with a feeble angulation behind the eyes. Disk of pronotum clothed with microscopic punctures, andwith a longitudinal puncture in antescutellar area.
Subapical impression of elytra (Fig. 16-A, E, F, G) feeble. Disk of elytra
sparsely clothed with microscopic punctures. Epipleural fossette densely andcoarsely punctate. Marginal elytral stria absent. Marginal epipleural stria complete, and densely punctate. External subhumeral and 1st-4th dorsal striae complete. Fifth dorsal stria a little shorter than the sutural, and its basal rudiment
represented by a short arc. Sutural stria present on apical half. Oblique humeral
stria present on basal third. All striae, except the oblique humeral one, strongly
crenate.
Pygidia (Fig. 19-1, J) alutaceous. Propygidium with an indistinct depression oneach posterior side, and its punctures coarse, deep, separated by 0.5-1.0 times theirdiameter, and minute punctures intermingled with them. Punctures on pygidium
denser than those on propygidium.
Prosternal lobe (Fig. 16-L, M, N) rounded at apex, its marginal stria complete,of which the basal end deeply and largely foveate. Prosternal keel rather broad,with carinal stria on each side, the basal ends of the striae being connected with eachother along the basal margin of prosternum (often interrupted).
Anterior margin of mesosternum (Fig. 16-H) slightly emarginate at middle, its
marginal stria complete and feebly crenulated. Disk of mesosternum sparsely
clothed with microscopic punctures. Meso-metasternal suture complete and subcarinate. Lateral stria of metasternum crenate, extending posteriorly to near, but
Table 11. Biometric data for Margarinotus (Grammostethus) niponicus(Lewis).
Part measured
APW
PPW
PL
EL
EW
ProW
ProL
PyL
PTL
MSTL
MTTL

Male

Female0.96-1.30 (1.14±0.010) 50
2.09-2.72 (2.47 ±0.021) 50
1.06-1.40 (1.25± 0.012) 50
1.46-2.13 (1.81±0.017) 50
2.48-3.28 (2.91± 0.025) 50
1.38-1.89 (1.70±0.016) 50
0.63-0.87 (0.76±0.009) 50
0.71-1.08 (0.92±0.010) 50
0.75-1.02 (0.89±0.009) 50
0.75-1.06 (0.92± 0.011) 50
0.94-1.38 (1.17±0.012) 50

1.02-1.38 (1.22± 0.009) 50
2.32-3.07 (2.71 ±0.026) 50
1.18-1.65 (1.40± 0.014) 50
1.73-2.28 (2.02±0.019) 50
2.80-3.66 (3.24± 0.030) 50
1.61-2.20 (1.91 ± 0.020) 50
0.71-1.14 (0.87±0.011) 50
0.87 -1.18 (1.03± 0.010) 50
0.79-1.10 (0.96±0.009) 50
0.83-1.18 (0.99± 0.011) 50
1.10-1.42 (1.27±0.012) 50

38

Fig. 16. M. (C.) niponicus (Lewis). A: Pronotum and left elytron. B-D: Head, frontal
view. E-G: Left elytron. H: Meso-, metasternum and 1st abdominal sternum. I:
Left protibia, dorsal view. J: ditto, ventral view. K: Left profemur, ventral view.
L-M: Prosternallobe and keel. N: ditto, lateral view. 0: Spermatheca and bursa
copulatrix, lateral view (right side) [Iwate, HonshO]. Scale: 0.5 mm.

not united with, the oblique stria which extends inwardly from the middle of themetasternal-metepisternal suture. Lateral disk of metasternum covered withcoarse and round punctures, and with short hairs_
1ntercoxal disk of 1st abdominal sternum (Fig. 16-H) completely striate on eachside.
Protibia (Fig. 16-1, J) with 7-8 dentic1es on external dorsal edge, and with 10
small dentic1es on external ventral edge. Profemur (Fig. 16-K) with posterior
marginal femoral stria, the stria short, present on apical area_
39

Male genitalia: as in Fig. 17-A, B, C, D, E, F.
Female genitalia: as in Fig. 16-0.
Specimens examined, 7 rJ', 4 ~ and 142 exs. Hokkaido -1 ex., Maruyama, Sapporo, 16/vii/
1942, N. Aoyama leg.; 1 ex., Usubetsu, 22/vii/1954, K. Sawada leg.; 1 ex., Horokanai, Moshiri, 9/
viii/1983, T. Hosokawa leg. ; 2 ~ and 28 exs., Nopporo, 17, 27/vi/1986, M. Ohara leg.; 1rJ', 4 exs.,
lozankei, 7/vi/1985, 13, 16/vii/1986, M. Ohara leg.; 1 ex., Tomakomai, 15/viii/1978, no collector's
name; 1 ex., Zenibako, 8/vii/1977, N. Nishikawa leg.; 1rJ', 1 ex., Chimikeppu lake, Tsubetsu, 6/viii/1987, M. Ohara leg.; 1rJ', Horonai, Okushiri Is., 18/vii/1986, Y. Kusui leg.: 1rJ', Inaho, OkushiriIs., 12/viii/1986, Y. Kusui leg.; 1 ex., Yagishiri Is., 21/vi/1985, Y. Kusui leg. HonshO - <Aomori·ken> 1 ex., Tuta'onsen, Towada, 25/vii/1954, J. Aoki leg. <Iwate·ken> 3rJ', 1 ~, 10 exs., Kuriyag·
awa, 20/v/1974, H. Hayakawa leg.; 4 exs., Nishine, 28/vi, 12/ix/1974, no collector's name.
<Fukushima·ken> 1 ex., Mt. Odayama, Wakamatsu, 12/v/1948, Y. Kurosawa leg.; 1 ex., Innai,
Higashiyama, Kita-aizu, 17/vii/1947, Y. Kurosawa leg.; <Tochigi·ken) 27 exs., Nikko, Kiyotaki,
15/v, 10/vii, 10/viii/1977, no collecter's name; 1 ex., Nikko, Tyugusi, 9-11/viii/?, T. Adachi leg.
<Chiba·ken> 5 exs., Oami, 22/vi/1984, M. Ohara leg. <Tokyo·to) 1 ex., Hongo, 29/v /1946, T.
N akane leg.; 3 exs., Aoyama, 27 /v /1946, 29/iv, 6/viii/1955, T. N akane leg.; 1 ex., Tokyo, 18/iv/
1952, H. Kajimura leg. <Kanagawa·ken> 1rJ', 1 ex., Sagami·oyama, 13/v/1967, 5/v/1968, K.
Masumoto leg. <Shizuoka·ken) 1 ex., Gotenba, 21/viii/1934, K. Nakamura leg. <Nagano·ken> 1
ex., Shimashima, 25, 26/vii/1946, T. Nakane leg.; 1 ex., Komanoyu, 25/vii/1947, T. Nakane leg.; 2exs., Noziri, 19/vii/1941, T. Nakane leg.; 1 ex., Komagatake, Kiso, 4, 5/viii/1946, S. Osawa leg.; 1ex., Kamikochi, 21-28/vii/1947, S. Osawa leg. <Gifu·ken> 1 ex., Hirayu, 24/vii/1946, M. Hayashi
leg. <Mie·ken> 1 ex., Yonoyama, Ise, 1/vi/1947, S. Osawa leg.; 1 ex., ditto, 3/vi/1948, S. Tabuchileg.; 1 ex., Hayakawa, 27-30/vii/1953, H.Ohira leg. <Kyoto·fu) 1 ex., Kurama, 1l/vii/1956, T.
N akane leg.; 1 ex., Mt. Daihi, 15/vi/1941, K. Sakaguchi leg.; 1 ex., Miyama, 4/vi/1984, T. Hosok·
awa leg. <N ara·ken) 1 ex., Kasuga, 23/vi/1946, F. Takahashi leg.; 1 ex., Yoshino, v /1939, K.
Kurosa leg. <Hyogo·ken> 1 ex., Hataganaru, Tazima, 12/v/1954, T. Okutani leg. <Wakayama·AE~"."""""",~~
Fig.I7. M. (G.) niponicus (Lewis) [Iwate, HonshQ]. A: Male genitalia, lateral view.
B: ditto, dorsal view. C: Apex of genitalia, oblique view. D: ditto, in median lobe
extruded, lateral view. E: Median lobe, dorsal view. F: ditto, lateral view.
Scale: 0.5 mm.
40

ken> 1 ex., N ishinokawa, 2/viij1957, M. Yoshida leg.; 1 c?', Singu, Southern Kii, 18/viij1946, S.
Osawa leg.; 1 ex., Nachi, Southern Kii, 16/viij1946, S. Osawa leg. <Tottori-ken> 2 exs., Mt.
KyOshO, 7/vij1984, S. Nomura leg. Shikoku - <Kochi-ken> 1 ex., Kodagasayama, Tosa, 12/iv/
1496, S. Nakamura leg. KyOshO - <Fukuoka-ken> 2 exs., Fukuchi-yama, 4/viiij1951, F. Takahashi leg.; 2 exs., Mt. Hiko, 18/v/1955, K. Sawada leg. <Saga-ken> 2 exs., Nakahara, 12/v/1952, 23/
iv/1950, Y.Miyake leg.; 1 !f., lex., Shimo, Hirokawa, 16, 18/vij1954, V_Miyake leg.; 2 exs.,
Tashiro, 12/v /1952, Y. Miyake leg.; 1 ex., N akamachi, 21/vij1952, A. Yamaguchi leg.; 1 ex.,
Ogashirayama, 7/v /1984, H. Yamaguchi leg. <Kumamoto-ken> 7 exs., Sakamoto, Kawaguchi, 15/
vij1985, T. Tanabe leg. <Kagoshima-ken> 1 ex., Kurino, Kirishima, 31/viij1985, M. Ohara leg.; 1ex., Takachihonomine, Kirishima, vij1967, H_ Makihara leg.; 1 ex., Kirishima, 25/viij1971, T.
Nakane leg.; 1 ex., Shiroyama, 19/vij1951, F. Takahashi leg.; 1 c?', 1 ex., Toso, Kagoshima City,
20/iv /1984, M. Ohara leg.; 1 ex., Yoshino, Kagoshima City, 6/v /1985, E. Matsui leg.; 1 ex.,Takakuma-yama, 5/v/1980, T. Kinoda leg. Nansei IsIs. -1 ex., Okinoerabu-jima, 19/v/1981, K.
Tomiyama leg.

Distribution (Fig. 21). Japan (Hokkaid6, HonshO, Shikoku, KyOshO, N ansei
IsIs.); Taiwan; Continental China; Korea; USSR (Khabarovskij and PrimorskijKray)_
Remarks. This species is very common in J apan_ It usually occurs on carcasses, sometimes on sap, fungi or decaying vegetable matter, and is rarely found indung.

NOTES ON THE PHYLOGENETIC RELATIONSHIP OF SOME MAJORGROUPS WITHIN THE GENUS MARGARINOTUS

The genus Margarinotus may be a monophyletic group and a good taxon.
However, the relationship of the species composing it is still uncertain, and thesubgenera as currently understood appear problematical so far as the male genitalia
examined (see Appendix) are concerned_ The Japanese species of Ptomister can bedivided into three groups on the basis of the male genitalia. Moreover, judging fromfigures of Kryzhanovskij and Reichardt (1976), Margarinotus koltzei (Schmit) and M.
hailar Wenzel seem to represent another type of the male genitalia and to form afourth group, the koltzei-group*. These groups are, in regard to the male genitalia,
often more closely related to other subgenera than to each other. For example, thesutus-group is apperently related to the subgenus Eucalohister Reitter, sharing withthe latter the apomorphic character that dorsal mid-line of the median lobe of themale genitalia is armed with a large denticle (fig. 11-J, M, N .)_ The koltzei-group is
closely related to the subgenus Paralister Bickhardt. They agree in the uni- or
trilobate apex of the median lobe of the male genitalia, a character also possibly
apomorphic_ I think that the male genitalia are very useful in distinguishing closely
related species because they might have evolved rapidly and divergently. Yet, on
the other hand, the fundamental structure of the male genitalia may reflect
phylogenetic relationships among species groups.
Relationships of the subgenera and groups, based on the male genitalia, areshown in Table 12_ This table suggests that the characters which have been used to• Koltzei-group. Median lobe of male genitalia not expanded on apical half, stick-like_
Apex of lobe bent backwards and uni- or trilobate in dorsal view. This group consists ofkoltzei (Schm.) and hailar Wenzel, and is distributed in northern China and eastern USSR.

41

Fig. 18. Propygidium (A, C, E, G, 1) and pygidium (B, D, F, H, J) of Margarinotus spp. AB: M. (P.) boleti (Lewis). C-D: M. (P.) cadavericola (Bickhardt). E- F: M. (P.)
agnatus (Lewis). G-H: M. (P.) weymarni Wenzel. I-J: M. (P.) reichardti
Kryzhanovskij.

42

Fig. 19. Propygidium (A, C, E, G, 1) and pygidium (B, D, F, H,]) of Margan·notus spp. AB: M. (P.) striola (C.R. Schalberg). C-D: M. (P.) yezoensis M. Ohara. E-F: M.
(P.) marginepunctatus (Lewis). G- H: M. (P.) sutus (Lewis). 1- J: M. (P.) niponicus(Lewis).

43

Fig. 20. Geographic distribution of Margarinotus spp.

44

Fig. 21.

Geographic distribution of Margarinotus spp.

Table 12. Types of the male genitalia of the genus Margarinotus.
Types of male genitalia

Subgenera or groups

Spoon-shaped

Stick-shaped

Weymami-group*
Margarinotus
A group of Grammostethus(including ru/icomis)
KOltzei-grou p*

Apex of median lobe
trilobate[ Paralister

Mid-line of median lobe
with a large denticle

EUCalohister[ Sutus-group*Median lobe without lobe
and denticle

Boleti-group *Another group of Grammostethus(including niponicus)
Stenister
Promethister* Groups of the subgenus Ptomister.

45

form the subgenera do not reflect phylogenetic relationships, involving parallelism
and even convergence, and that the present classification within the genus Margarinotus is not natural. I may conclude, therefore, that (1) the subgenus Ptomister isa polyphyletic group, and (2) the koltzei-group should be united with the subgenus
Paralisler, and the sutus-group with the subgenus Eucalohister, to form a monophyletic taxon respectively. These conclusions, however, should be tested withfurther material.

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Cooman, A., 1947. Coleopteres Histeridae d'Extreme-Orient. Not. Ent. Chin., 11: 421-428.
'Dejean, P. F., 1837. Catalogue des Coleopteres de la collection. Troisieme edition, revue,
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Desbordes, H., 1919. Contribution a la connaissance des Histeridae. 4e memoire. Etude desHisteridae de !'Indo-Chine (Tonkin, Laos, Siam, Annam, Cambodge, Cochinchine).
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Fabricius, ]. Ch., 1787. Mantissa Insectorum sistens eorum species nuper detect as adiectis
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Flensburgi et Lipsiae, I-II, Hafniae, I: xx + 348 pp., II: 382 pp.
'Grimm, B., 1852. Hister ruficornis n. sp. Ent. Ztg., 13: 221-223.
Hatch. M.H .. 1962. The beetles of the Pacific Northwest, Part III. Pselaphidae & Diversicornia 1. Univ. Wash. Pub I. BioI.. 16 (1961): ix + 503 pp.
Hisamatsu, S .• 1984. Histeridae beetles collected by Dr. K. Baba in Niigata Prefecture.
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Essa ent. Soc. N iigata, 57: 15-18 (in Japanese).
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*Horion. A .. 1949. Faunistik der mitteleuropaischen Kafer. Frankfurt a: MB 11, Hist.:
317 -369.
Houlbert. c.. and E. Monnot. 1922-1923. Faune entomologique armoricaine. Coleopteres.
42 e Famille. Histerides (Escarbots). Bull. Soc. Sci. Med. Ouest.. 31-32: sep. pp. 175.1922: 1-16,1923: 17-75.
Kamiya. K .. and S. Takagi. 1938. A list of Japanese Histeridae. Scient. Agr., 19 (1): 21-32.
Kryzhanovskij. O.L., 1966. Novye vidy zhukov-gisterid (Coleoptera, Histeridae) fauny SSSR.
Trudy Zool. Inst. Akad. N auk SSSR, 37: 51-59.
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Lewis, G., 1884. On some Histeridae new to the Japanese fauna, and notes of others. Ann.
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Mag., (2) 6 (31): 186-189.
- - - 1899. On new species of Histeridae and notices of others. Ann. Mag. Nat. Hist., (7)4: 1-29- - - 1900. On new species of Histeridae and notices of others. Ann. Mag. Nat. Hist., (7)6: 265-290.
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10: 223-239.
- - - 1905. A systematic catalogue of Histeridae. London, vi+81 pp.
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18: 397-403.
- - - 1915. On new species of Histeridae and notices of others. Ann. Mag. Nat. Hist., (8)16: 54-56.
Marseul, S.A., 1853. Essai monographique sur la famille des Histerides, comprenant ladescription et la figure au trait des genres et des especes, leur distribution methodique,avec un resume de leurs moeurs et de leur anatomie. Ann. Soc. Ent. France, (3) 1 :131-160, 177-294, 447-553.
- - - 1854. Essai monographique sur la famille des Histerides (suite). Ann. Soc. Ent.
France., (3) 2: 161-311,525-592,671-707.
- - - 1873. Coleopteres du Japon recueillis par M. Georges Lewis. Enumeration desHisterides et des Heteromeres avec la description des especes nouvelles. Ann. Soc.
Ent. France, (5) 3: 219-230.
Mazur, S., 1970. Contribution to the knowledge of Histeridae of Korea (Coleoptera).
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- - - 1972. Systematic and synonymic notes upon certain species of Histeridae(Coleoptera). Pol. Pismo Ent., 42: 137-143.
- - - 1973. Spaheritidae i Gniliki - Histeridae. In Klucze do oznaczania owad6w
Polski, xix, 11-12, Warszawa, 74 pp.
- - - 1984. Description of a new Margarinotus species with additional notes about twohisterids from Nepal (Col., Histeridae). Rev. Suisse Zool., 91: 163-167.
- - - 1984b. A world catalouge of Histeridae. Pol. Pismo Ent., 54: 1-379.
Motschulsky, V., 1860. Insectes du Japon. Etud. Ent. 9: 4-39.
Nakane, T., 1963. Histeridae. In Nakane, T., et al. (eds.), Iconographia Insectorum
Japonicorum Colore naturali edita, II (Coleoptera): 67-70, pis. 34-35 (in Japanese).
- - - 1981. Histeroid beetles hitherto recorded from Kyushu, Japan. [Nagasaki-ken
seibutsu-gaku gakkai-shiJ, 21: 7-10 (in Japanese).
Ohara, M., 1988. Some records of Histerid beetles from Niigata Prefecture. Trans. Essa
ent. Soc. Niigata, 65: 63-64 (in Japanese).
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Systematik und Bionomie der Histerini (Coleoptera, Histeridae). Acta Ent. Bohemoslov., 79: 196-206.
Osawa, Sh., and T. N akane, 1951. Studies on Japanese Histeridae (1) - Histeridae of N oziri- (N. Shinano, Japan). Bull. Takarazuka Insectaurium, 79: 1-10 (in Japanese).
Reichardt, A.N., 1926. Zametki 0 palearkticheskich Histeridae (Coleoptera). I, Rusk. Ent.
Obozr., 20: 269-274- - - 1930. De Histeridis (Coleoptera) novis faunae palearcticae. Annu. Mus. Zool., 30(1929): 285-304.
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Reichardt, A.N., and O.L. Kryzhanovskij, 1964. On the fauna of the Histerid-beetles(Coleoptera) of South-eastern China. Ent. Obozr., 43 (1): 170-174.
Reitter, E., 1879. Verzeichniss der von H. Christoph in Ost-Sibirien gesammelten Clavicornier, etc. Dtsch. Ent. Z., 23: 209-226.
47

- - - 1909. Fauna Germanica. Die Ktlfer des Deutschen Reiches. II Band, Schr. Dtsch.
Lehrerver. Naturk., 24, 392 pp.
'Sahlberg, e.R., 1819. Insecta Fennica, dissertationibus academicis, a. 1817-1834 editis,
enumerata. Pars I: a. 2-4. Helsingforsiae, viii +519 pp. [pp.9-56 were edited in
1819J.
Sharp, D., and F. Muir, 1912. The comparative anatomy of the male genital tube in
Coleoptera. Trans. ent. Soc. London, 1912 (III): 477-642, pis. XLII-LXXVIII.
'Thomson, e.G., 1862. Skandinaviens Coleoptera, synoptiskt bearbetade. IV, Lund, 269 pp.
Vienna, P., 1977. Nuove tabelle per la determinazione delle specie Italiana della tribu'
Histerini (Coleoptera, Histeridae). Soc. Ven. Sci. Nat. Lavori, 2: 35-42.
Wenzel, R.L., 1944. On the classification of the histerid beetles. Fieldiana, zool., 28: 51-151.
- - - 1960. Three new histerid beetles from the Pacific Northwest, with records andsynonymies of additional species (Coleoptera, Histeridae). Fieldiana, zool., 39 (41) :447-463.
- - - 1971. Zoogeography of the Holarctic genus Margarinotus (Col., Histeridae). XIIIIntern. Congr. of Entom., Proceedings, I: 215-216.
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, Original not seen.
48

ApPENDIX

A list of species of the genus Margarinotus of which the male genitalia have been studied
in the present paper ("Ohara, 1989" in the list) and by other authors.
References

SpeciesMargarinotus(Margarinotus)
gutti/er Hom(Ptomister)<Boleti-group>
boleti (Lew.)
fenderi Wenz.
<Weymarni-group>
agnatus (Lew.)

brunneus (F.)

cadavericola (Bickh.)

distinctus (EL)fractifrons (Csy.)
multidens (Schm.)
rectus (Csy.)
reichardti Kryzh.
striala (C.R. SahIb.)

ssp. succicala (Thomson)

tristriatus Wenz.
weymarni Wenz.

yezaensis M. Ohara<Kaltzei-group>
hailar Wenz.
kaltzei (Schm.)<Sutus-group>
marginepunctatus (Lew.)
sutus (Lew.)

Wenzel, 1944 : pI. 6, f. 1.

Ohara, 1989: 12, f.5.
Wenzel, 1960: 461, f. D, E.
Wenzel, 1944: pI. 6, f. 2, as ballaui Wenz.
Kryzhanovskij and Reichardt, 1976: 333, f. 644.
Ohara, 1989: 17, f.6.
Sharp and Muir, 1912: f. 79, 79a, as cadaverinus Hoffm.
Wenzel, 1944: pI. 6, f. 4, as ditto.
Kryzhanovskij and Reichardt, 1976: 333, f. 643, as ditto.
Wenzel, 1944: pI. 8, f. 1, as ussuriensis Reich.
Kryzhanovskij and Reichardt, 1976: 333, f. 645.
Ohara, 1989: 12, f.5.
Kryzhanovskij and Reichardt, 1976: 333, f. 650.
Hatch, 1962: pI. 56, f.8.
Kryzhanovskij and Reichardt, 1976: 333, f. 649.
Hatch, 1962: pI. 56, f.9.
Kryzhanovskij and Reichardt, 1976: 333, f. 646.
Ohara, 1989: 23, f.9.
Wenzel, 1944: pI. 7, f.1, as strialides Wenz.
Kryzhanovskij and Reichardt, 1976: 343, f. 676-9.
Ohara, 1989: 30, f. 11.
Wenzel, 1944: pI. 7, f.2, as striata c.R. Shalb.
Kryzhanovskij and Reichardt, 1976: 333, f. 648; 343, f.674-5.
Ohara, 1989: 26, f. 10.
Wenzel, 1944: pI. 7, f. 3, 4.
Kryzhanovskij and Reichardt, 1976: 349, f.690.
Wenzel, 1944: pI. 6, f.3.
Kryzhanovskij and Reichardt, 1976: 333, f. 647.
Ohara, 1989: 21, f. 8.
Ohara, 1989: 30, f. 11.
Kryzhanovskij and Reichardt, 1976: 349, f.694-5.
Kryzhanovskij and Reichardt, 1976: 349, f.691-3.
Ohara, 1989: 30, f. 11.
Ohara, 1989: 30, f. 11.
49

Species

References<?>
kathmandu Mazur

Mazur, 1984: 164, f.3-5.

merdarius (Hoffm.)
terricola (Germ.)
wenzelianus Kryzh.
(Eucalohister)
bipustulatus (Schrank)

Kryzhanovskij and Reichardt, 1976: 349, f.688.
Kryzhanovskij and Reichardt, 1976: 349, f. 689.
Kryzhanovskij and Reichardt, 1976: 416, f. 4.

binotatus (EL)gratiosus (Mnnh.)
kurdistanus (Mars.)
mirabilis (Khnz.)
solskyi (Schm.)(Stenister)
bickhardti (Rtt.)graecus (Brulle)
kabakovi Kryzh.
obscurus (Kugel ann)
(Paralister)
carbonarius (Ill.)ignobilis (Mars.)
koenigi (Schm.)
laevi/ossa (Schm.)
lecontei Wenz.
neglectus (Germ.)
oblongulus (Schm.)
punctiventer (Mars.)
purpurascens (Hbst.)
silantjevi (SchiL)
uncostn'atus (Mars.)
ventralis (Mars.)(Grammostethus )niponicus (Lew.)
ruficomis (Grimm)

Kryzhanovskij and Reichardt, 1976: 374, f.785-7.
Kryzhanovskij and Reichardt, 1976: 375, f. 788.
Kryzhanovskij and Reichardt, 1976: 375, f. 790-l.
Kryzhanovskij and Reichardt, 1976: 375, f. 792-3.
Kryzhanovskij and Reichardt, 1976: 375, f. 794-5.
Kryzhanovskij and Reichardt, 1976: 375, f. 789.
Kryzhanovskij and Reichardt, 1976: 371, f. 773-4.
Kryzhanovskij and Reichardt, 1976: 371, f. 775-7.
Kryzhanovskij, 1980: 143, f. 3.
Kryzhanovskij and Reichardt, 1976: 371, f. 770-2., as stercorarius(Hoffm.).
Kryzhanovskij and Reichardt, 1976: 357, f.725-6.
Kryzhanovskij and Reichardt, 1976: 357, f.731-2.
Wenzel, 1944: pI. 8, f.2.
Kryzhanovskij and Reichardt, 1976: 357, f. 723-4; 361, f. 738-9.
Kryzhanovskij and Reichardt, 1976: 361, f. 740-l.
Wenzel, 1944: pI. 8, f.3.
Kryzhanovskij and Reichardt, 1976: 361, f.735.
Kryzhanovskij and Reichardt, 1976: 361, f.742.
Kryzhanovskij and Reichardt, 1976: 357, f.728., as stigmosus(Mars.).
Kryzhanovskij and Reichardt, 1976: 361, f.736-7.
Kryzhanovskij and Reichardt, 1976: 357, f. 727.
Kryzhanovskij and Reichardt, 1976: 357, f. 729-30.
Kryzhanovskij and Reichardt, 1976: 357, f. 733-4.
Kryzhanovskij and Reichardt, 1976: 368, f.757-60.
Ohara, 1989: 40, f.17.
Kryzhanovskij and Reichardt, 1976: 368, f. 751-4.

(Promethister)con/usus Wenz.
marginatus (EL)marginicollis (J.E. LeConte)
prometheus (Kryzh.)(Species incertae sedis)/elipae (Lew.)

Wenzel, 1944: pI. 9, f. 3.
Kryzhanovskij and Reichardt, 1976: 365, f.749.
Wenzel, 1944: pI. 9, f. l.
Kryzhanovskij and Reichardt, 1976: 365, f.748.
Wenzel, 1944: pI. 9, f.2.

50