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A REVISION OF THE JAPANESE EUMENIDAE
(HYMENOPTERA, VESPOIDEA)
YAMANE, Seiki
Insecta matsumurana. Series entomology. New series, 43: 1189
1990-08
http://hdl.handle.net/2115/9855
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Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP
INSECTA MATSUMURANA
NEW SERIES 43
AUGUST
1990
A REVISION OF THE JAPANESE EUMENIDAE
(HYMENOPTERA, VESPOIDEA)
By Sm::r YAMANE
Abstract
Yamane, Sk. 1990. A revision of the Japanese Eumenidae (Hymenoptera, Vespoidea). Ins. matsum. n. s.
43: 1-189, 7 tabs., 366 figs.
The Japanese forms of Eumenidae are revised, and 54 species (77 forms in total) are recorded. The
following five forms are described as new to science: Pararrhynchium oceanicum (Ogasawara Is.), P. o. miyanoi
(Ogasawara Islands), 5ymmorphus carinatus (Shikoku), 5. iwatai (Honshu), and 5. tsushimanus (Tsushima Is.).
New synonyms are: (5tenodynerus rufomaculatus kikaiensis Sk. Yamane) =5. rufomaculatus, (5. yambarah Sk.
Yamane et Gusenleitner) =5. kusigematii, (Rhynchium haemorrhoidale samurayi Giordani Soika) =R.
quinquecinctum fukaH, (Anterhynchium fIavomarginatum luctuosum Giordani Soika) =A. f. hanedai,
(Ancistrocerus yamanei Giordani Soika) =A. oviventris, and (Eumenes samurayi rufescens Giordani Soika) =E.
micado. Ancistrocerus parietinus is newly recorded from Japan and numerous new localities are presented for
the known species. The original description of Euodynerus bicingulatus Giordani Soika well agrees with small
specimens of male Anterhynchium flavomarginatum micado. The former is possibly a synonym of the latter.
The eumenid fauna of Japan is characterized by many Palearctic, some Oriental and some endemic
elements. But it lacks some of the genera widely distributed in the Palearctic region such as Pseudepipona,
Antepipona, etc. On Sado-ga-shima and other islands located near the Japanese mainlands, the fauna is rather
poor, with no endemic forms. Although the fauna is much poorer, the Izu Islands harbor one endemic form
at subspecies level, 5tenodynerus tokyanus fIavoscutellatus. Of the 13 eumenid forms 50 far known from the
Tsushima Islands, at least one is no doubt a Korean element (Eumenes punctatus), and two are apparently
endemic (Anterhynchium flavomarginatum tsushimarum and 5ymmorphus tsushimanus). Two of the five species
known to occur on the Ogasawara Islands are supposed to be native. Both are endemic and very peculiar in
morphology and color pattern, and are each differentiated into two subspecies occurring in the Haha-jima
and Chichi-jima group. The Ryukyu Eumenidae are composed of Palearctic, wide-ranging, endemic and
Oriental elements. The Northern RyukyU5 are dominated by Palearctic and wide-ranging elements, the
Central Ryukyus by wide-ranging and endemic elements, and the Southern Ryukyus by Oriental and wideranging elements. The largest gap in species composition is found between Amami-oshima and Yaku-shima.
Subspecies differentiation and regional convergence in color pattern among unrelated species are most
remarkable in the Ryukyil. Islands.
Author's address.
Department of Biology, Faculty of Science, Kagoshima University, Kagoshima, 890
Japan.
Part of a thesis submitted to the Hokkaido University in partial fulfillment of the requirements for
the degree of Doctor of Agriculture, 1990.
Contents
I. Introduction ................................................................................................................................................................. 3
Acknowledgements .............................................................................................................................................. 4
IT. Materials and survey areas .................................................................................................................................... 5
ITI. Family Eumenidae (Potter wasps) ...................................................................................................................... 11
1. Taxonomy ......................................................................................................................................................... 11
2. Biology .............................................................................................................................................................. 13
3. Eumenids as natural enemies of pests ......................................................................................................... 14
IV. Characters and their conditions ............................................................................ ;............................................ 15
V. Taxonomy of the Japanese Eumenidae ................................................................................................................ 20
Key to the Japanese genera of Eumenidae ...................................................................................................... 20
Genus Discoelius Latreille ................................................................................................................................... 22
Genus Stenodynerus Saussure ........................................................................................................................... 25
Genus Allodynerus Bliithgen .............................................................................................................................. 41
Genus Euodynerus Dalla Torre .......................................................................................................................... 46
Genus Rhynchium Spinola .................................................................................................................................. 63
Genus Anterhynchium Saussure ......................................................................................................................... 69
Genus Okinawepipona Sk. Yamane ............................................................ :....................................................... 83
Genus Pararrhynchium Saussure ........................................................................................................................ 87
Genus Orancistrocerus Vecht .............................................................................................................................. 95
Genus Andstrocerus Wesmael ............................................................................................................................ 98
Genus Symmorphus Wesmael ........................................................................................................................... 113
Genus "Pachymenes" Saussure .......................................................................................................................... 135
Genus Oreumenes Bequaert .............................................................................................................................. 137
Genus Eumenes Latreille ................................................................................................................................... 140
Genus Dellll Saussure ........................................................................................................................................ 154
Genus Pseumenes Giordani Soika .................................................................................................................... 159
Genus Pseudozumia Saussure ........................................................................................................................... 162
VI. Distribution pattern of the Eumenidae in some insular regions .................................................................. 164
1. Sado-ga-shima ............................................................................................................................................... 165
2. Izu Islands ...................................................................................................................................................... 166
3. Tsushima Islands ........................................................................................................................................... 167
4. Ogasawara Islands ........................................................................................................................................ 168
5. Ryflkyfl Islands .............................................................................................................................................. 169
References .................................................................................................................................................................. 177
Appendix .................................................................................................................................................................... 185
Index ............................................................................................................................................................................ 186
I. INrROOUCI10N
The Eumenidae are a large diplopterous family containing 177 extant genera
(Carpenter, 1986) and more than 2000 species in the world. They are all predators, hunting
lepidopterous or coleopterous larvae for their young (Iwata, 1971). They are principally
solitary wasps, even when gathering around suitable nesting sites. Some species of the
genus Zethus and a few other genera are known to be communal nesters, though it is not
certain whether they co-operate in provisioning. In still others, the mother wasp cares for
her young and progressively provisions larval cells with prey. In addition, in the
eumenids, in general, oviposition takes place prior to provisioning. All this shows that the
Eumenidae represent an incipient stage in the evolution of sociality, and they have been
paid increasing attention by insect sociologists.
Another important aspect in their biology is a potential role in controlling agricultural
and forest pests. Although at present we have only a few experiences in managing them
as useful predators (Takeshima, 1971; Lee, 1984), some eumenids may be involved in the
integrated control of leaf-eating insect pests such as microlepidopterous larvae.
Undoubtedly an essential step toward a comprehensive understanding of biology
and behavior of eumenid wasps is to establish a sound classification system of them. The
Japanese Eumenidae were first studied taxonomically by European entomologists such as
Smith (1852, 1873), Perez (1905), Schulthess (1908, 1934), Cameron (1911) and others. By
1931 Matsumura had recorded eleven species from Japan including the Ryukyus. In 1930s,
an important progress was made by Yasumatsu who studied material collected by Iwata
in the course of his behavioral studies of eumenids as well as material in the collection of
the Entomological Laboratory, Kyushu University. His treatments of species were,
however, often erroneous, chiefly because he did not examine type specimens deposited
in European institutes.
Since 1940s up to now, A. Giordani Soika of Italy has continued to study Japanese
eumenids on the basis of specimens sent from Japan. He has compared them with
abundant material gathered from all over the world and including type specimens.
However, his studies on Japanese Eumenidae have exclusively been based upon dead
material from restricted localities, so that he has sometimes failed in detecting sibling
species and in recognizing subspecies. Further, he has not revised the Japanese Eumenidae
as a whole.
The main purpose of the present study is to revise the eumenid fauna of Japan as
completely as possible, based upon all the efforts previously made by many authors and
upon my own study initiated in the early 1970s when I was at the Entomological Institute,
Hokkaido University as a postgraduate student. Revisions of some groups including
many new forms have already been published by me in co-operation with Mr. T. Tano and
others. In the present paper I will enumerate 54 species, of which four are new to science.
In order to facilitate future biological study, a summary of nesting behavior is given for
each species whenever information is available. This is based mainly on Iwata's
observations, but partly also on unpublished data collected by me or several other
hymenopterists. Because the scientific names adopted by Iwata in his papers are to be
critically revised, I have examined the Iwata collection preserved at Kobe University to
determine the correct names of the wasps studied by him.
Geographical distribution is discussed with special emphasis on small islands. Above
all, the Ryukyu Archipelago is very interesting because of its geographical position
3
(between the Kyushu mainland and Taiwan) and a large number of islands included.
Analyses are made regarding the relative abundance of Oriental and Palearctic elements,
area-species relation, competitive exclusion among species with similar nesting habits,
and regional convergence in color pattern among non-related species on some islands.
Acknowledgements. I wish to express my cordial thanks to Prof. Sadao Takagi
(Hokkaido Univ.), Prof. Toshio Nakashima (Hokkaido Univ.), Prof. Takehiko Nakane
(Chiba-shi), Prof. Mitsuru Hotta (Kagoshima Univ.) and Prof. Shoichi F. Sakagami
(Hokkaido Univ.) for their kind guidance and constant encouragements. Prof. S. Takagi,
Prof. Hans Mori (Hokkaido Univ.) and Dr. Tosio Kumata kindly read through earlier
drafts of the manuscript.
My hearty thanks should also be expressed to Dr. Masaaki Suwa (Hokkaido Univ.),
Prof. Setsuya Momoi (Kobe Univ.), Dr. Tikahiko Naito (Kobe Univ.), Dr. Yasuo Maeta
(Shimane Univ.), Prof. Yoshihiro Hirashima (Kyushu Univ.), Dr. Osamu Tadauchi (Kyushu
Univ.), Dr. Shuichi Ikudome (Kagoshima Women's Junior College) and Prof. Seiji Azuma
(Univ. Ryukyus) for permitting me to study valuable eumenid collections in their care and
their helpful suggestions.
My thanks are extended to Dr. A. Giordani Soika (Museo Civico Storia Nat., Venezia),
Prof. J. van der Vecht (Putten, Netherlands), Dr. J. Gusenleitner (Linz), Dr. N. V. Kurzenko
(Institute of Biology and Pedology, Vladivostok) and Dr. J. M. Carpenter (Harvard Univ.)
who provided me with valuable specimens for comparison and useful suggestions. Dr.
Giordani Soika, in particular, kindly determined Ancistrocerus specimens for me.
The following entomologists much helped me in proViding valuable materials, useful
information and suggestions: Shigeyuki Aoki (Rissho Univ.), Kintaro Baba (Niigata-ken),
Toyokazu Fujisawa, Haruo Fukuda (Kagoshima Pref. Mus.), Katsuo Goukon (Tohoku
Gakuin Univ.), Yoshito Haneda (Fukui-ken), Shiro Higashi (Kagoshima Univ.), Toru
Inaoka (Asahigawa-shi), Ryosuke Ishikawa (Tokyo Metropolitan Univ.), Hideo Itami
(Shibata-shi), Yosiaki Ito (Nagoya Univ.), Kazuaki Kamijo (Hokkaido Forest Exp. Sta.),
Teiji Kifune (Fukuoka Univ.), Jun-ichi Kojima (lbaraki Univ.), Utako Kurosu (Tokyo Noko
Univ.), Kanetosi Kusigemati (Kagoshima Univ.), Yoshihisa Kusui (Toyonaka-shi), Shun'ichi
Makino (Hokkaido Univ.), Taiji Moriyama (Kagoshima-shi), Tadao Murota (Fukui-ken),
Hirohiko Nagase (Kamakura-shi), Akira Nagatomi (Kagoshima Univ.), Koji Nakamine
(Kagoshima-shi), Yasuhiro Nakatani (Nara-ken), Junichi Nakayama (Sapporo-shi),
Toshiaki Nambu (Saitama-ken), Masahiro Ohara (Hokkaido Univ.), Ryohichi Ohgushi
(Kanazawa Univ.), Masayuki Onuma Ooetsu-shi), Isao Otsuka (Kumamoto-shi), Akio
Seino (Shibata-shi), Hirohisa Sud a (Sakura-shi), Tsukasa Sunose (Urawa Gakuin High
Schoo!), Yasushi Takai (Seki-shi), Yoshisuke Takeshima (Aomori-ken), Hideo Takahashi
(Hachijo-jima), Tadashi Tano (Fukui-ken), Mieko Tatsuno (Yokohama-shi), Mamoru
Terayama (Toh6 Women's Senior High Schoo!), Kiyonori Tomiyama (Tokyo Metropolitan
Univ.), Katsuji Tsuneki (Mishima-shi), Shoji Watahiki, Hirofumi Watanabe (Kagoshima
Univ.), Masateru Yamada (Aomori-ken), Kazuki Yamamuro (Tokyo-to), S6ichi Yamane
(Ibaraki Univ.), Seiei Yamauchi (Okinawa-ken) and Junichi Yukawa (Kagoshima Univ.).
Last but not least I thank Dr. Nobumitsu Kawakubo (Kagoshima Univ.) for his kind
help in editing the final draft of the manuscript.
4
II.
MATERIAlS AND SURVEY AREAS
The present study is mainly based upon my collection now at the Department of
Biology, Faculty of Science, Kagoshima University. The main part of the collection will
finally be deposited in the collection of the Entomological Institute, Faculty of Agriculture,
Hokkaido University. I have examined many valuable specimens preserved in the
following collections: the Iwata collection in the Entomological Laboratory, Faculty of
Agriculture, Kobe University, the Tano collection (Fukui-ken), the Ikudome collection,
Kagoshima Women's Junior College, the collection of the Entomological Institute,
Hokkaido University, and the collections of the Entomological Laboratory, Faculty of
Agriculture, Kyushu University, and University of the Ryukyus.
The names of some collectors are abbreviated as follows: AN (A. Nagatomi), HI (H.
Itami), KB (K. Baba), KT (K. Tomiyama), SI (S. Ikudome), SKY (Seiki Yamane), SY (SOichi
Yamane), TM (T. Murota), TN (T. Nambu), TT (T. Tano), YH (Y. Haneda), YM (Y. Maeta).
The present study covers the regions mentioned below. Honshu, Shikoku and
Kyushu together are called Japan proper, and these three islands and Hokkaido together
the Japanese mainlands (Figs. 1,2). The Japanese suffix -jima; -shima or -to usually but not
always means an island or islet. The suffixes -shoto, -gunto and -retto, which are applied
to archipelagos and island groups, are not used in this paper: for example, "Yaeyamashoto" is constantly replaced by the Yaeyama Islands or the Yaeyama group. Other suffixes
used are -ken, -to and -fu, all for prefectures.
1. Hokkaido (= Ezo, Yezo). 78,073 km 2 in area. Sapporo and its vicinity have most
intensively been surveyed. In other parts, especially northern and eastern parts, the
eumenid fauna is still only poorly known.
2. Islands close to Hokkaido. Munakata and S. Yamane (1970) studied vespid wasps of
Okushiri-to (143 km 2 ), and Munakata (1987) recorded two eumenid species from the
island. Other islands remain almost wholly unsurveyed. I have examined two species of
Ancistrocerus from Rebun-to (83 km2 ) located close to Wakkanai, the northernmost city in
Japan.
3. Honshu. The largest island of Japan, 227,414 km 2 in area. Some entomologists,
including non-professional, have accumulated eumenid collections in the following
regions: Aomori-ken (Yamada, 1983), Miyagi-ken (Goukon, 1983a,b), Niigata-ken (K.
Baba, A. Seino; see Yamane, 1982b), Chiba-ken (Suda, 1979), Saitama-ken (Nambu, 1978),
Ibaraki-ken (Hisamatsu et al., 1986), Nagano-ken (Y. Maeta), Fukui-ken (Haneda et al.,
1985), Kansai District (K. Iwata; Sato, 1963a,b, 1964), and Shimane-ken (Y. Maeta). They
reported local faunas in journals or have sent me representative specimens.
4. Sado-ga-shima and Awa-shima (Niigata-ken). These islands in the Japan Sea were
surveyed by members of the Essa Entomological Society, especially by Dr. K. Baba and Mr.
A. Seino. The eumenid fauna of Sado-ga-shima (857 km 2 ) is relatively well known
(Yamane, 1982b), whereas at present no information is available for Awa-shima (9.1 km 2).
5. Izu Islands (Tokyo-to). This island group is located south of the Kanto District
between 34°45'N and 29°25'N in the Pacific Ocean, and consists of some ten islets, of
which Izu-oshima is the largest (90.99 km 2 ) (Fig. 3B). There are brief papers reporting
insect faunas of these islands, but no complete list of aculeates is available. I have
examined some eumenid specimens through the courtesy of Prof. Y. Hirashima and Mr. H.
Takahashi.
6. Nanatsu-jima Islands (Ishikawa-ken). Located in the Japan Sea just north of the Noto
5
Okin o-tor i. shima
""'- TAIWAN
1
Fig. 1. Japa n and ad·Jacent regions.
6
Peninsula. The largest islet (Hegura-jirna) is 1.03 km 2 in area, and all the other islets of the
Nanatsu-jima group are less than 0.15 km 2. The wasp fauna of these islands was studied
by Ohgushi and Tokumoto (1986). I have examined the material used by them.
7. Oki Islands (Shimane-ken). This island group lies to the north of the Shimane
Peninsula in the Japan Sea, and comprises four main islands (13.64-244.31 km 2 ) and many
minute ones. I have examined a few eumenid specimens preserved in the collection of
Shimane University through the courtesy of Dr. Y. Maeta.
Q
Okushin
Oki Is.
,.0
Tsushima
IJ/IS.
P
Izu Is.
·04
Osumi
Is.
o,
.
300
km
2
Fig. 2. Japanese mainlands and some associated islands. 1, Hokkaido; 2, Aomori-ken; 3, Iwate-ken;
4, Miyagi-ken; 5, Akita-ken; 6, Yamagata-ken; 7, Fukushima-ken; 8, Ibaraki-ken; 9, Tochigi-ken;
10, Gumma-ken; 11, Saitama-ken; 12, Chiba-ken; 13, TokyO-to; 14, Kanagawa-ken; 15, Niigataken; 16, Toyama-ken; 17, Ishikawa-ken; 18, Fukui-ken; 19, Yamanashi-ken; 20, Nagano-ken; 21,
Gifu-ken; 22, Shizuoka-ken; 23, Aichi-ken; 24, Mie-ken; 25, Shiga-ken; 26, Kyoto-fu; 27, Osakafu; 28, Hyogo-ken; 29, Nara-ken; 30, Wakayama-ken; 31, Tottori-ken; 32, Shimane-ken; 33,
Okayama-ken; 34, Hiroshima-ken; 35, Yamaguchi-ken; 36, Tokushima-ken; 37, Kagawa-ken;
38, Ehime-ken; 39, Kochi-ken; 40, Fukuoka-ken; 41, Saga-ken; 42, Nagasaki-ken; 43,
Kumamoto-ken; 44, Oita-ken; 45, Miyazaki-ken; 46, Kagoshima-ken.
7
8. Other islllnds close to Honshu. Most of these islands remain unsurveyed for the
eumenid fauna. Iwata's collection (Kobe University) contains some specimens from Awajishima (593 km 2 ) located between Honshu and Shikoku. Okada (1981) recorded two
species of Eumenidae in the southern parts of this island. Yuki (1936) listed five eumenid
species from Iwai-jima (7.5 km2 ), an island in the Setonaikai Sea, where lie numerous
islands.
9. Shikoku. 18,256 km2 in area. I have examined many specimens collected by Mr. Y.
Sugihara and Dr. S. Ikudome mainly in Kochi-ken. Their materials may well represent the
eumenid fauna of this island (lkudome & Yamane, 1983). None of the islets located near
Shikoku has yet been surveyed for eumenids.
10. Kyushu. 36,554 km 2 in area. The following regions have been more or less
intensively surveyed: Fukuoka-ken (K. Yasumatsu), Nagasaki-ken (R. Ohgushi),
Kumamoto-ken (Otsuka, 1984), and Kagoshima-ken (Nagase, 1981, 1982).
11. Tsushima Islllnds (Nagasaki-ken). This interesting island group, located between
Korea and KyfIshu, has been paid special attention by biogeographers. Collections have
been made mainly on two large islands (Kami-agata, 253 km2; Shimo-agata, 445 km2 ),
and most of the numerous minute ones remain unsurveyed. Shirozu and Miyata (1976)
compiled a list of insects recorded. Through the courtesy of Dr. Y. Miyatake I could
examine the eumenid specimens mainly collected by the late Mr. Isamu Hiura and
preserved in the collection of the Osaka City Museum of Natural History. Mr. A. Seino
and Mr. K. Nakamine provided me with some interesting additional materials.
12. Gate) Islands (Nagasaki-ken). This island group lies in the west of northern
B
A
v7v
<f
Muko·
jima Is
.2
40°3
5"
06
Q
,~
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'/
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Izu
Is.
9·~
Haha·
8
jim~~
_ /" 1>"
Muko- .•..
jima
Haha·
jima Is
"10
3
Fig. 3. Ogasawara (Bonin) Islands (A) and Izu Islands (B). 1, Izu-Oshima; 2, To-shima; 3, Nii-jima;
4, Shikine-jima; 5, Kozu-jima; 6, Miyake-jima; 7, Mikura-jima; 8, Hachijo-jima; 9, Hachijokojima; 10, Ao-ga-shima.
8
Kytishu, and comprises numerous islands. Ejima et al. (1981) recorded four species of
Eumenidae. Mr. J. Nakayama sent me some specimens collected by him. However, most of
the islands of this archipelago remain unsurveyed.
13. Other Islands close to KyushU. Miyata et al. (1977) recorded four eumenid species
from Oki-no-shima (Fukuoka-ken) located in the Genkai-nada and at 77 km from
Fukuoka-shi. There are numerous islands belonging to Nagasaki-ken, most of which have
not been surveyed for the eumenid fauna. I have examined only a few specimens from the
Amakusa Islands (Kumamoto-ken), the Naga-shima Islands (Kagoshima-ken), the
Koshiki-jima Islands (Kagoshima-ken), and Akune-oshima (Kagoshima-ken). No
information is available about the eumenid fauna of the islets close to Oita-ken and
Miyazaki-ken.
14. Ryukyu Islands (Kagoshima-ken & Okinawa-ken). There are more than 200 islands
between the Kytishu mainland and Taiwan (Fig. 4; most of the small islets are omitted).
These islands, including the Dait6 and Senkaku groups, are called the Nansei Islands
(Nansei-shot6; South-western Islands). The Ryiikyti Islands here defined are those islands
that lie between Kytishii and Taiwan, excluding the Daito and Senkaku Islands. Synonyms
are the Ryukyus, the Ry.1kyu-retto, the Ryuky3. Archipelago and the Ryukyu Curve. In
older literature Rytikyii was often spelled as "Liukiu", "Loochoo", or "Riukiu". This long
archipelago (ca. 1200 km from north to south) comprises many island groups and
subgroups for which various nomenclatorial systems have been applied. In this paper I
use a slightly modified version of the system proposed by Mezaki (1980, 1983). Island
groups and main islands belonging to them are listed below (for island areas, see
Discussions on geographical distribution):
A. Northern Ryukyus (Kagoshima-ken)
Vji Islands: le-jima (= Iye-jima), Mukai-jima
Kusagaki Islands: Kami-no-shima, Shimo-no-shima
Mi-shima Islands (= Kuchi-no-mishima Is.): Take-shima, I6-jima (= Iwo-jima, Kikaiga-shima), Kuro-shima
Osumi Islands: Tane-ga-shima, Mage-shima, Yaku-shima, Kuchinoerabu-jima
Northern islets of Tokara Islands (Linshoten Is.): Kuchi-no-shima, Naka-no-shima,
Gaja-jima, Taira-jima, Suwanose-jima, Akuseki-jima
B. Central Ryukyus
Southern islets of Tokara Islands (Kagoshima-ken): Takara-jima, Kodakara-jima
Amami Islands (Kagoshima-ken): Amami-oshima (= Oshima), Kikai-jima,
Kakeroma-jima, Uke-shima, Yoro-shima, Tokuno-shima, Okinoerabu-jima (= Okierabujima), Yoron-to (Yoron-jima)
Okinawa Islands (Okinawa-ken): Iheya-jima, Izena-jima, Okinawa-jima (=
Okinawa-honto), Kouri-jima, Yagaji-jima, Sezoko-jima (=Sesoko-jima), Yokatsu group (Ikeshima, Miyagi-jima, Yabuchi-jima, Hamahiga-jima, Tsuken-jima, Henza-jima), Kudakajima, Kerama group (Mae-shima, Tokashiki-jima, Zamami-jima, Aka-shima, Kuba-jima),
Aguni-jima, Kume-jima
C. Southern Ryukyus (= Saki-shima Islands) (Okinawa-ken)
Miyako Islands: Miyako-jima, Ikema-jima, Kurima-jima, Irabu-jima, Shimoji-jima
Tarama Islands: Tarama-jima, Minna-jima
Yaeyama Islands (= Yayeyama Is.): Ishigaki-jima, Taketomi-jima, Kuro-shima,
Kayama-jima, Kohama-jima (= Kobama-jima), lriomote-jima, Yubu-jima, Hatoma-jima,
Vchibanare-jima, Sotobanare-jima, Hateruma-jima, Yonaguni-jima (= Yonakuni-jima)
9
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Fig. 4. Nansei Islands, comprising Ryilkyii Is., Daito Is. and Senkaku Is. Most of the small islets are
omitted. 1, Ie-jima; 2, Mukai-jima; 3, KUr<H;hima; 4, IO-jima; 5, Take-shima; 6, Tane-ga-shima;
7, Mage-shima; 8, Yaku-shima; 9, Kuchinoerabu-jima; 10, Kuchi-no-shima; 11, Naka-no-shima;
12, Suwanose-jima; 13, Akuseki-jima; 14, Takara-jima; 15, Kikai-jima; 16, Amami-&hima; 17,
Kakeroma-jima; 18, Uke-shima; 19, YOr<H;hima; 20, Tokuno-shima; 21, Okinoerabu-jima; 22,
Yoron-to; 23, Iheya-jima; 24, Izena-jima; 25, Okinawa-jim a; 26, Kouri-jima; 27, Yagaji-jima; 28,
Sezoko-jima; 29, Yabuchi-jima; 30, Hamahiga-jima; 31, Miyagi-jima; 32, Tokashiki-jima; 33,
Zamami-jima; 34, Aguni-jima; 35, Kume-jima; 36, Miyako-jima; 37, Irabu-jima; 38, Kurimajima; 39, Tarama-jima; 40, Minna-jima; 41, Ishigaki-jima; 42, Taketomi-jima; 43; Kuro-shima; 44,
Kohama-jima; 45, lriomote-jima; 46, Hatoma-jima; 47, Uchibanare-jima; 48, Hateruma-jima; 49,
Yonaguni-jima.
Intensive collections of insects have been made by Prof. S. Azuma and Mr. M. Kinjo of
the University of the Ryukyus, but their efforts have been restricted to a small number of
relatively large islands in Okinawa-ken. Recently, more than 50 islands have been
surveyed for the aculeate fauna by me and some other biologists (e.g., S. Ikudome, C.
Nozaka, T. Murota, K. Tomiyama, K. Nakamine, and Y. Kusui). Literature on the vespoid
fauna of the Ryukyus will be cited in Distribution pattern. The folloWing islands remain
almost wholly unsurveyed: uninhabited islets of the Tokara group such as Gaja-jima,
10
Kogaja-jima and Yokoate-jima, islets around Kakeroma-jima (Amami group) such as
Sukomobanare-jima and Eniyabanare-jima, many of the islets of the Okinawa group, and
some islets in the Southern Ryukyus such as Ogami-jima (Miyako group), Aragusuku
group, and Nakanougan-jima (Yaeyama group).
15. Senlazku Islands (Okinawa-ken). This island group is loclited to the north of the
Yaeyama group and consists of four islets (0.30-3.82 km2 ) and some rocks. Takara (1954)
and Ikehara and Shimojana (1971) reported the terrestrial animal fauna of this island
group, but their lists include no vespoid species.
.
16. Daifo Islands (= Oagari Is.) (Okinawa-ken). The Daito group comprises the
following isolated islands: Kitadaito-jima (= Kitaoagari-jima, North Borodino I., 12.58
km 2), Minamidaito-jima (= Minamioagari-jima, South Borodino I., 30.59 km 2 ), and
Okidaito-jima (= Okioagari-jima, Rasa I., 1.19 km2 ).It is separated from the islands of the
Ryukyus by the Ryukyu Trench (maximum depth: 7481 m) (Hanzawa, 1935). I have
examined eumenid specimens collected by Dr. S. Ikudome on Kita- and Minamidait6-jima
in 1987.
17. Ogasawara Islands (= Bonin Is.) (Tokyo-to). The Ogasawara group is volcanic in
origin (oceanic islands), and comprises three subgroups (Fig. 3A) and three isolated islets.
Aculeate fauna has been studied on a few larger islets (Chichi-jima, 23.9 km 2, & Hahajima, 21.2 km2 ). Nakane (1970) listed the insects known at that time, including three
eumenid species. I have examined some eumenids collected by several entomologists on
these two islets.
III.
FAMILY EUMENIDAE (POTIER WASPS)
1. Taxonomy
The Vespoidea are often called Diploptera because of the longitudinal folding of their
fore wings at rest, though this characteristic is absent in some groups and is not regarded
as part of the ground-plan of the Vespoidea (Carpenter, 1982). The outstanding
autapomorphies are the elongate 1st discoidal cell (discal cell in Carpenter, 1982) which is
at least the equal of submedian cell, spined parameres of male genitalia, and oviposition
prior to provisioning (Carpenter, 1982).
The classification system proposed by Richards (1%2) for the superfamily Vespoidea
has long been accepted by most of vespidologists. He recognized three subfamilies in the
family Eumenidae, viz., Raphiglossinae, Discoeliinae, and Eumeninae. Among them the
unnaturalness of the Discoeliinae (= Zethinae of Carpenter) was previously recognized to
some extent by Bohart and Stange (1965), and Carpenter (1982) conclusively showed the
paraphyly of the group. According to Carpenter the Raphiglossinae perhaps form a
monophyletic group, but do not merit taxonomic recognition, since a group composed of
the Eumeninae and Zethinae and lacking the Raphiglossinae is unnatural. He recognized
only one family (Vespidae) in the Vespoidea, and reduced the traditional Eumenidae to a
subfamily (Eumeninae) comprising the traditional Raphiglossinae, Zethinae, and
Eumeninae. His taxon Eumeninae is characterized by the following derived character
conditions:
Mandible: dypeus ratio 0.78-2.00; occipital carina forked, with a branch running to
the mandibular base and one running toward the hypostoma (either branch, and usually
the hypostomal, may be secondarily lost); parategula present; hind coxa with dorsal
11
carina; claws bifid; basal ring of male genitalia short, encompassing bases of parameres;
volsella with digitus, a broad pincerlike lobe; cuspis being a reduced scalelike lobe
attached mesally to parameres; larval labrum as wide as clypeus and narrowed where
these join, dorsally bisinuate and ventrally bilobed.
I agree with Carpenter in his view that at present no formal taxonomic recognition
should be made at suprageneric levels, because there has been made no detailed
phylogenetic analysis of the genera. However, in this paper Richards' view is adopted and
the eumenids are treated as a family, because I do not necessarily agree with the view held
by Carpenter that the classification system should be isometric with the cladogram.
Among the Japanese Vespoidea, the Eumenidae are readily distinguished from the
Vespidae Oapan is lacking in Masaridae) by the bifid claws, the presence of parategula
(process of mesoscutum sensu Richards), and the long mandibles usually crossing each
other.
Approximately 180 genera are currently recognized in the Eumenidae (Carpenter,
1986). No reliable figure is available for the number of world species of this large and
diverse group. Bequaert (1918) presented the following figures for various zoogeographical regions: 349 species in the Palearctic, 225 in the Nearctic, 369 in the Ethiopian,
549 in the Oriental and Australian, 607 in the Neotropical regions. Vecht and Fischer
(1972) recognized 702 Palearctic species in 56 genera; this figure is twice as big as that
given by Bequaert (1918) for the same region. According to Krombein (1979) 266 species in
23 genera occur in the Nearctic region. Taylor et al. (1985) listed more than 300 described
Australian species in 35 genera, many of them being endemic. Judging from these figures
the number of the world species may amount to 2500-3500. In Japan 54 species are known
to occur (this study).
Table 1. Chromosome numbers in Eumenidae·.
Ancistrocerus japanicus
Stenodynerus tokyanus 1)
Allodynerus mandschuricus2)
Euodynerus nipanicus3 )
Euod. foraminatus
Rhynchium quinquecinctum4)
Anterhynchium f/ilvomarginatum
Orancistrocerus drewseni
Ancistrocerus adiabatus
A. spilogas ter
A. simulator
A. tuberculiceps
Symmorphus apiciornatus 5)
S. {vveolatus 6)
Honshu, Jpn
ditto
ditto
ditto
Calif., USA
Iriomote I., Jpn
AmamiI.,Jpn
Honshu, Jpn
ditto
Calif., USA
ditto
ditto
ditto
Honshu, Jpn
ditto
n= 12
n= 18
n= 8
n= 5
n= 8
n= 15
n= 15
n= 6
n= 14
n= 6
n= 6
n= 7
n= 10
n= 4
n= 5
• Cited from Goodpasture (1974) for the N. American species, and from Nakatani (1988) for the Japanese ones. In Nakatani, the species 1) - 6) in this list are
inadequately referred to as: 1) Stenodynerus sp., 2) Stenodynerus frauenfeldi, 3)
Euodynerus notatus, 4) Rhynchium fukaii, 5) Symmorphus sp., and 6) S. captivus.
12
Karyological studies were made for some Nearctic and Japanese species by
Goodpasture (1974) and Nakatani (1988). Five out of the nine species studied by Nakatani
were unidentified or misidentified; corrected names for them are given in Table 1.
2. Biology
The Eumenidae are solitary wasps with some exceptions, in which female's caring for
the larvae (subsocial behavior) is observed. Among the subsocial species are Synagns spp.
(Bequaert, 1918), Calligaster cyanopterus (Williams, 1919), Antepipona tropicalis (Iwata, 1971),
Orancistrocerus drewseni (Iwata, 1972; Itino, 1980), Pararrhynchium ornatum ornatum (Iwata,
1938a, 1983, pp. 108-109), Paraleptomenes mephitis (Krombein, 1978), some species of Zethus
(Bohart & Stange, 1965), and probably Oreumenes decoratus (Tsuneki, 1980). Progressive
provisioning characterizes these subsocial species (Eickwort, 1981). The Japanese
population of the subsocial o. drewseni is very probably parthenogenetic, since no males
have been found in both field-captured adult and nest samples, while the Korean and
Taiwanese populations are biparental.
The eumenids build their nests at least partially with mud or plastic materials from
plants. Many of them use pre-existing hollow cavities for nests (tube-renters), some others
build their nests (often mud pots) above ground not using pre-existing tubes or holes
Table 2. Nesting types in the Japanese Eumenidae. The Japanese fauna
lacks burrowers.
Mud-daubers
OranCistrocerus drewseni'
Ancistrocerus japonicus'
A. melanocerus
A. oviventris'"
Eumenes rubronotatus
Oreumenes decoratus'
Delta esuriens
Delta [lavapidum'
Tube-renters
Disroelius japonicus'"
Stenodynerus frauenfeldi
Allodynerus delphinalis
A. mandschuricus
Euodynerus dantici
Euod. nipanicus
Euod. trilobusRhynchium quinquecinctum
Anterhynchium [lavomarginatum
Pararrhynchium ornatum
Orancistrocerus drewseni'
Ancistrocerus japonicus'
A. trifascia tus
A. parietinusA. nigricornisA. antilope'"'
Symmorphus foveolatus
S. decens
S. apiciornatus
S. mutinensis
S. c/iens
Pseumenes depressus"Pachymenes" yaeyamensis
Potter wasps
Eumenes fraterculus
E. rubrofemoratus
E. micado
Oreumenes decoratus'
Delta [lavapidum'
• Species with two nesting types .
•• Species constructing cell partitions made of plant leaves .
... Observed in Taiwan, China, or Europe by various authors.
13
(mud-daubers and potter wasps), and still others, not represented in Japan, dig either in
clayey ground or in walls (burrowers)(cf. Iwata, 1971; Spradbery, 1973) (fable 2). Muddaubers construct their nests on stones or walls of houses; the nests are broadly attached
to the substrates and not complete pots. The true potter wasps construct their pot nests on
plant stems, etc. These two types are, however, sometimes not cleiu-Iy distinguished. The
burrowers often make a chimney at the exit of the burrows with mud paste; The nesting
proceeds in the order: cell building - oviposition" hunting, with the only exception of the
Oriental genus PSeumenes in which the hunting is said to preCede OViposition (Piel, 1935).
In general female wasps hunt larvae of various lepidopteran families for their larvae.
Wasps of Raphiglossa, Psiloglossa and Symmorphus hunt for coleopteran larvae.
Biology of tube~nesting Ellmenidae in the USA was intensively studied by Krombein
(1967). Spradbery (1973) reviewed the natural history of eumenids, based mainly upon
British species. The comparative behavior of the eumenid wasps is reviewed by IWata
(1942, 1971, 1982), Malyshev (1968), Evans and West-Eberhard (1970), and Kurzenko
(1980).
3. Eumenids as natural enemies of pests
The members of the familly Eumenidae are predatory wasps and provide their nests
with caterpillars or other insect larvae. Therefore, they are at least potentially beneficial in
agriculture and forestry. However, in most countries of the world, they have not positively
been used as natural enemies of agricultural pests.
Some common Japanese species hunt for the mature larvae of leaf-rollering microlepidopterans of the families Tortricidae, Pyralidae and Gelechiidae. Among them Anterhynchium flavomarginatum micQdo and Rhynchium quinquecinctum fukaii prefer pyralid
larvae, while Euodynerus nipanicus (Odynerus quadrifasciatus sensu Iwata) predates chiefly
on Tortricidae. Mr. Yoshisuke Takeshima, a devoted farmer in Aomori-ken (northern
Honshu), set reed tubes for three eumenid species, A. flavomarginatum, Euod. nipanicus and
Discoelius japonicus, that were commonly seen in his apple orchard where damage by
tortricid larvae was serious. These tortricids (mainly Adoxophyes orana) not only feed on
young leaves that are rolled by them, but often damage the surface of the fruit itself. Of
the three eumenid species, Euod. nipanicus increased in number, and in 1970 the number of
tortricid larvae transported to nests of this species was estimated at 92,700 in the orchard
(ca. 1 hectar). The damage to the fruit was less extensive in the area provided with trap
nests than in the control. Unfortunately, his method has not been developed thereafter,
presumably because the management of wasps and timing of sprinkling insecticides are
burdensome.
In China, vespine and polistine wasps have been used to control agricultural pests
(caterpillars) (Lee, 1982, Lee et al., 1986). Lee et al. (1975) studied the biology of a
subtropical eumenid, Rhynchium quinquecinctum (R. brunneum), that predates on
lepidopterous pests in cotton and rice fields. They stated that in Zhenghai County one
female wasp captured 90-180 caterpillars per day. But the activity of this wasp did not
necessarily cover the effective time for controlling insect pests. Since this eumenid is
comparatively not vulnerable to pesticides, it is possible to use it together with the latter
(Lee et al., 1984). However, in general, the effectiveness of eumenid wasps as indigenous
biotic factors against pest insects is at present not accurately evaluated.
14
IV.
CHARACTERS AND 1HEIR CONDmONS
Numerous characters have been used in vespoid taxonomy and systematics.
Important characters are discussed in Bequaert (1918), Richards (1962) and others.
Carpenter and Cumming (1985) made a character analysis of the Nearctic Eumenidae, and
Nakamine (1987) discussed the polarity for characters with the Japanese species and
genera of Eumenidae. In this paper no attempt will be made to determine the polarity
(ancestral VS. derived). The characters of taxonomic importance used in this paper are
listed below with their observed conditions (states). No detailed study has been made on
the male genitalia. Here Yamane's (1982) and Nakamine's (1987) results are particularly
consulted, and only Japanese genera and species are referred to.
Head
Mandibular shape. The mandibles are relatively short and not crossing each other
(Discoelius, Fig. 20), or longer and decussate (other genera, e.g., Figs. 6, 337).
Cephalic fovea (Fig. 5, cej). There are often a pair of small pits (foveae) set in a slight
depression (da) on the female vertex near its occipital margin. The condition of the
depressed area and the distance between the pits vary according to genera and sometimes
even between congeneric species. Sometimes the depressed area bears only one pit, and in
several genera (Discoelius, Pseumenes, Oreumenes, Eumenes, Delta, and "Pachymenes") it
bears no pits. The state of the cephalic foveae is discussed at length by Cumming and
Leggett (1985).
Clypeal shape. Clypeus is higher than wide in many genera, while wider than high in
others such as Discoelius, Symmorphus and Allodynerus (Figs. 20, 62). It is apically
emarginate variously, and often more pronouncedly in the male.
Palpi. Maxillae and labium bear palpi. The number of segments is usually 6 in
maxillary and 4 in labial pal pi (Fig. 7), but reduced in some forms and most commonly to
5 and 3 (5 : 3 according to the palpal formula proposed by Carpenter & Cumming, 1985).
Among the Japanese genera this condition is seen only in Okinawepipona (Fig. 8;
Nakamine, 1987).
Terminal segment of male antenna. Usually the terminal (13th) segment of the male
antenna is recurved to form a hook. Only in Symmorphus and Oreumenes it is of normal
shape (but abnormal among the Eumenidae)(Figs. 236-245, 290). The size of the hook and
modification of segments that receive the hook are of taxonomic importance.
Alitrunk (mesosoma: thorax + propodeum)
Pits and punctures on the anterior face of pronotum. The anterior vertical face of
pronotum is generally smooth, shining, without hairs, but in some groups it has pits,
punctures or striae. For example, in Stenodynerus the area has a pair of median pits in
lower part, and some transverse striae just above the pits (Fig. 13). In "Pachymenes" the
striae are deep (Fig. 14). Distinct punctures are found in lateral parts of the area in
Euodynerus, "Pachymenes" and some species of Stenodynerus.
Pronotal carina (Figs. 9, 10). The presence of a pronotal carina constitutes part of the
ground-plan of the Eumenidae (Carpenter & Cumming, 1985). The carina is generally
complete, and often forms a translucent lamella. In Delta fiavopictum it is wholly lost, and
in some genera such as Ancistrocerus and Stenodynerus medially (dorsally) interrupted. It is
not interrupted but reduced in height in the middle in Okinawepipona, Euodynerus and
15
oao
0" po0
cet
¥
da
5
Figs. 5-8. Head of Eumenidae. 5,6, Stenodynerus ogasawaraensis. aD, anterior ocellus; cef, cephalic
fovea; el, cIypeus; da, depression for fovea; fl, antennal flagellum; fr, frons; OS, ocular sinus; pd,
pedicel; po, posterior ocelli; sc, scape. 7, 8, maxillary (A) and labial (B) palps of S.
ogasawaraensis (7) and Okinawepipona kog!mai (8).
Allodynerus (Nakamine, 1987).
Pretegulilr carina (Fig. 10). The pronotallobe (pT) is usually defined anteriorly by a
distinct carina (pc). It is found in most Japanese genera, but absent in Eumenes.
Epicnemial carina (Fig. 10). This carina (ec), separating epicnemium (ep) from the rest
of mesepisternum and commonly seen among eumenid genera, is absent in Ancistrocerus,
Pseumenes, Oreumenes, Eumenes, Delta, and Symmorphus foveoliltus.
Pleural and epipleural sutures (Fig. 10). Pleural suture (or sulcus, ps) divides the
mesopleuron into an anterior mesepisternum and posterior mesepimeron (me). Epipleural
suture (es) (=? dorsal mesepisternal groove of Richards, 1973) divides the mesepisternum
into dorsal (dms) and ventral mesepisternum (vms). These sutures vary in condition, from
a simple narrow furrow to a rather wi<;le one with many transverse carinae in it. They are
evanescent in "Pachymenes".
Notaulices (Fig. 9). These (nt) are a pair of furrows running on the mesoscutum (msc)
from near its posterior margin forward. Carpenter and Cumming (1985) states that this
character seems to be useful only at the specific level. Among Japanese genera, notaulices
are well developed in Discoelius and Symmorphus, and incomplete (seen only in posterior
part) in Pseudozumia, Orancistrocerus and Pararrhynchium. They may be called "prescutal"
16
pi
/
sc
ml
-n
;. ..\
n
v
ooo
13
12
Figs. 9-14. Alitrunk of Eumenidae. 9, Discoelius japonicus (-'f), from above (after Nakamine, 1987);
10, Rhynchium marginellum, profile (after Bequaert, 1918); 11, Stenodynerus chinensis, profile
(original); 12, Posterior portion of alitrunk of Rhynchium aestuans, from above (afetr Bequaert,
1918); 13, 14, anterior vertical face of pronotum in Stenodynerus frauenfeldi (13) (after Yamane,
1979) and "Pachymenes" yayeyamensis (14) (after Nakamine, 1987).
con, propodeal concavity; dlprp, dorsolateral part of propodeum; dms, dorsal
mesepistemum; dml, dorsal metapleuron; ec, epicnemial carina; ep, epicnemium; es, epipleural
suture; hml, horizontal part of metanotum; Iprp, lateral portion of propodeum; me, median
carina; me, mesepimeron; msc, mesoscutum; ml, metanotum; nl, notaulix; pc, pretegular carina;
pI, pronotaJ lobe; pml, posterior face of metanotum; ppl, parapsidal line; pre, pronotal carina;
prn, pronotum; prp, propodeum; prps, propodeal shelf; ps, pleural suture; pi, parategula; se,
scutellum; I, tegula; va, propodeal valvula; vms, ventral mesepisternum; vml, ventral
metapleuron. a-b, lateral ridge; b-c superior ridge; b-d, inferior ridge.
grooves when in the latter condition.
Parapsidallines (Fig. 9). Most eumenids have a second pair of mesoscutual marks, the
parapsidallines (pp/), lateral to the notaulices and extending anteriorly from the posterior
margin of mesoscutum (Gauld & Bolton, 1988). These lines are often called parapsidal
sulci or furrows, though on some occasions they are not furrows but carinae.
Tegula and parategula (= posttegula, mesoscutal hook) (Figs. 9, 10). Tegula (t) is
variable in shape and length. In some groups its posterior lobe exceeds the posterior end
of parategula (pt). The phylogenetic significance of the shape and development of tegula
17
is unknown, though this character is useful in recognizing the Japanese genera.
Metanotum (Figs. 9, 10). In some genera (Rhynchium, Anterhynchium, Euodynerus), the
posterior vertical face (pmt) is demarcated from dorsal horizontal part (hmt) by an angled
edge which is sometimes dentate or spinose (Fig. 79). Metanotum may be depressed
medially (lateral parts projecting) in Pararrhynchium and Rhynchium, or flat in most of the
other genera.
Propodeal concavity (Figs. 11, 12). Propodeum (morphologically first abdominal
segment) may have a median concavity (propodeal concavity; con) on posterior face.
There may be a narrow but distinct, horizontal part (called "propodeal shelf' (prps) by
Carpenter & Cumming, 1985) between metanotum and the concavity (e.g., some species of
Stenodynerus, and Symmorphus). The concavity, sometimes not distinctly concave but
rather flat, is divided by a median longitudinal furrow or a carina (the carina is often
running in the furrow). The conditions of the furrow and carina considerably vary.
Eumenes and Delta are completely lacking in the carina.
Ridges on propodeum (Figs. 10, 12). The posterior face of propodeum is more or less
angled on the lateral border, sometimes forming a ridge or carina; the upper and lower
17
19
DEC
"
Figs. 15, 16. Gaster of Symmorphus captivus (after Nakamine, 1987). If, longitudinal furrow on
tergite 2; tc, transverse carina.
Figs. 17-19. Wings of Stenodynerus frauenfeldi. Systems for veins and cells follow those by Das and
Gupta (1989).
Forewing veins. AB, costa; CD, subcosta; EFG, metacarpus; WKHYXZ, cubitus;
MNIT'F, radius; MN, abscissa 1 of radius; NT, abscissa 2; IT', abscissa 3; TF, abscissa 4;
PQRW, discoideus; RLO, subdiscoideus; CS, medius; UV, submedius; VW, brachius; BME,
stiga; SWD, basal vein; KN, intercubitus 1; XT, intercubitus 2; ZT, intercubitus 3; QH,
recurrent vein 1; YL, recurrent vein 2; PV, nervulus; QRW, postnervulus.
Forewing cells. MC, median; RC, radial; lCU, 1st cubital; 2CU, 2nd cubital; 3CU, 3rd
cubital; SMC, s~bmedian; IDC, 1st discoidal; 2DC, 2nd disroidal; 3DC, 3rd discoidal; IBC, 1st
branchial; 2BC, 2nd branchial; AC, anal; pst, parastigma; st, stigma.
Hindwing cells. MEC, mediellan; RC, radiellan; SEC, submediellan; CEC, cubitellan;
AEC, anellan; DEC, discoidellan; ai, anal lobe.
18
part of the ridge are called the superior (b-c) and inferior ridge (b-d), respectively. The
ridge may be toothed or spined. Well-developed ridges are called the propodeal crest,
which may bear distinct processes (upper teeth) (Figs. 78, 80; a) just behind the
metanotum or propodeal shelf. A ridge extending from the upper end of inferior ridge to
the anterolateral corner of propodeum is called lateral ridge (a-b) (Bequaert, 1918).
Apical part of propodeum. Each lateral half of the posterior face of propodeum may be
pointed, truncated or deeply emarginate at apex. The apical part bears a long spine in
"Pachymenes" (Fig. 287).
Mid-tibial spurs. Among the Japanese genera, Discoelius has two apical spurs on each
mid tibia (Fig. 25), while all the others have only one.
Gaster (meta soma)
In this paper gastral tergites and sternites are often called simply tergites and
sternites. Thus, "tergite 1" corresponds to abdominal tergite 2.
Shape of gastral segment 1. Petiolate condition occurs in Discoelius, Oreumenes,
Eumenes, Delta and Pseumenes (eg., Figs. 23, 291, 340). Segment 1 is longer than wide in
some Symmorphus, and 2/3 as wide as segment 2 in "Pachymen.es" (Fig. 287). In all the other
Japanese genera, it is as wide as segment 2 or only slightly narrower than the latter
(usually more than 4/5 as wide as the latter).
Transverse carina on tergite 1 (Figs. 15, 16). The carina (tc), separating the anterior
vertical face from posterior horizontal part of tergite 1, occurs in Symmorphus,
Orancistrocerus, Pararrhynchium, and Ancistrocerus, but is dorsally or in the middle
interrupted or evanescent in Orancistrocerus and Pararrhynchium. A much reduced
condition is observed in Okinawepipona in which the tergite bears a short carina on the
lateral face (Giordani Soika, 1986; Nakamine, 1987; Yamane, 1987).
Longitudinal furrow on tergite 1 (Fig. 16). The furrow (if> is rather wide and deep in
Symmorphus. It is slitlike and situated only apically in Pseumenes. Many other genera have
the horizontal part of the tergite 1 more or less depressed, but in Rhynchium, Anterhynchium and Euodynerus no trace of furrow or depression is found. There is a transverse
depression anterior to the thickened apical margin in Discoelius.
Apical margin of tergite 1. Condition of this part is quite variable, but is divided into
two principal types (Nakamine, 1987). Apical margin may be strongly thickened into a
cord like formation (Discoelius, Eumenes, Oreumenes)(tergite apically duplicate) or
thickened but less cord like (Symmorphus, "Pachymenes", Allodynerus), while in other genera
(e.g., Orancistrocerus, Pararrhynchium, Rhynchium, Anterhynchium, Euodynerus) it is in the
same plane as the disc of the tergite.
A narrow lamellate area occurs caudally on this tergite in some genera (Discoelius,
Eumenes, Symmorphus, Euodynerus, Stenodynerus). In the last genus this area is apically
dentate. A translucent area also occurs in Rhynchium and Anterhynchium.
Fore wing
Wing venation is rather stable among the Japanese Eumenidae. The condition in
parastigma, 2nd recurrent vein and postnervulus may vary among genera.
Parastigma. Parastigma (pst) is usually half or less as long as stigma (=pterostigma:
st). In Oreumenes, Delta, Pseudozumia, Rhynchium and Anterhynchium it is much longer,
often as long as stigma. In Pseumenes depress us parastigma is somewhat long.
19
Color pattern
Color pattern is especially useful in separating subspecies (geographical races). In the
Ryiikyii Islands some eumenids have several geographical races (color forms). Distinct
regional convergence in color pattern is seen on the Yaeyama group and Okinawa group
among species involving those of other families. Color pattern is also important at the
specific level. For example, two very closely related species of Allodynerus are nearly
constantly discriminated from each other solely by the color pattern.
In general the male wasps are more abundantly marked with light colors, and xanthic
forms tend to occur more often in the southern part of a given distribution range. But
exceptions to the latter rule are not rare, especially in insular regions. For example, a
remarkably melanic form of Anterhynchium flavomarginatum inhabits Tsushima situated
between Kyiishii and Korea. Forms occurring on Okinawa-jim a are usually darker in
maculation than those on Amami-Oshima situated northerly.
V.
TAXONOMY OF TIlE JAPANESE EUMENIDAE
Key to the Japanese genera of Eumenidae (applicable to the Japanese forms only)
This key is chiefly based upon Yamane (1982a) and Giordani Soika (1978). Useful
information can also be drawn from Kurzenko (1978), who studied the USSR genera of
Eumenidae.
1. Mid tibia apically with two spurs. Mandibles short, medially not crossing each other.
Gastral segment 1 slender, petiolate. Mesoscutum with developed notaulices. Tergite 1
with neither transverse carina nor longitudinal furrow. [Hokkaid6 - Kyiishii, N. & c.
Ryukyus1........................................................................................................................ Discoelius
- Mid tibia with only one apical spur.......................................................................................... 2
2. Tergite 1 with both a longitudinal furrow on the posterior horizontal part and a
transverse carina between the anterior vertical face and horizontal part. Gastral segment
1 more or less narrowed, sometimes longer than wide. Notaulices on mesoscutum
generally developed. Tergites 2-6(7) usually almost impunctate, shining. [Hokkaid6 Kyushiil ..................................................................................................................... Symmorphus
- Tergite 1 without longitudinal furrow; at most the corresponding part slightly
impressed. Gaster of variable shape ......................................................................................... 3
3. Gastral segment 1 slender, distinctly petiolate ....................................................................... 4
- Gastral segment 1 stumpy, always wider than long, more than half as wide as segment 2.
......................................................................................................................................................... 9
4. Mesoscutum with distinct prescutal grooves. Mesepistemum with epicnemial carina. [5.
Ryukyusl ................................................................................................................... Pseudozumia
- Prescutal grooves and epicnemial carina absent. .................................................................... 5
5. Propodeum with several pairs of carinae obliquely running downward from the midline. Mesoscutum, scutellum, metanotum without conspicuous punctures. [5. Ryukyusl
......................................................................................................................................... Pseumenes
- Propodeum without obliquely running carinae. Mesoscutum, scutellum, metanotum
densely punctate.......................................................................................................................... 6
6. Tergite 2 posteriorly with a well-defined lamellate area. Alitrunk subglobular. Tergite 1
20
usually densely punctate, posteriorly more or less thickened ............................................ 7
- Tergite 2 without lamella. Tergite 1 without distinct punctation, posteriorly not
thickened ...................................................................................................................................... 8
7. Female clypeus apically truncate. Lateral part of propodeum rather clearly demarcated
from posterior (dorsal) face. Terminal segment of male antenna very small, not
recurved. [Hokkaido - Kyushu, N. and S. Ryukyusl ............................................ Oreumenes
- Female clypeus apically more or less emarginate. Propodeum round on each side,
without an edge between lateral and posterior face. Terminal segment of male antenna
forming a recurved hook. [Hokkaido - Ky11shu, N. RyukyusJ ................................ Eumenes
8. Tergite 1 longer, much longer than alitrunk. Male terminal sternite without longitudinal
furrow. Metapleuron and lateral part of propodeum almost impunctate, shining. Body
markings vivid yellow. [C & S. RyukyusJ ....................................................... Delta (Phi auct.)
- Tergite 1 shorter, only slightly longer than alitrunk. Male terminal sternite with a
longitudinal furrow. Propodeum laterally with punctures, not shining. Body with
yellow and rufous markings. [C & S. Ryukyusl ............................................................. Delta
9. Gastral segment 1 rather slender (but not distinctly petiolate), two-thirds as wide as
segment 2. Tergite 2 "'lith a pair of yellow spots. Mesopleuron with the epipleural suture
quite evanescent. [C & S. Ryukyusl .................................................................... "Pachymenes"
- Gastral segment 1 wider, usually more than four-fifths as wide as segment 2. Tergite 2
only rarely with a pair of yellow spots (Stenodynerus kusigematii pachymenoides often has
such spots) .................................................................................................................................. 10
10. Tergite 1 with a transverse carina (sometimes medially obscure) between the anterior
vertical face and posterior horizontal part. .............................................................................11
- Tergite 1 without transverse carina (in Okinawepipona and some species of Stenodynerus
the carina absent, but the anterior face rather sharply demarcated from the dorsal part) .
....................................................................................................................................................... 13
11. Parastigma of fore wing more than half as long as stigma. Posterior half of mesoscutum and scutellum longitudinally and coarsely rugose rather than punctate. Propodeal crest not developed. Female clypeus wider than high, apically widely truncate.
Thorax entirely black. The male is not found in the Japanese population. [Honshu Kyushu, N. Ryukyusl ......................................................................................... Orancistrocerus
- Parastigma half or less as long as stigma. Mesoscutum and scutellum punctate rather
than rugose. Propodeal crest often developed ...................................................................... 12
12. Lateral face of propodeum strongly reticulate. Upper processes of propodeal crest
pointed, close to each other. Female clypeus higher than wide, widely and shallowly
emarginate at apex. Transverse carina on tergite 1 rather obscure in P. ishigakiense.
[Honshu -Kyushu, Ryukyusl ........................................................................... Pararrhynchium
- Lateral face of propodeum striate, not distinctly punctate. Upper processes of crest, if
any, blunt apically, widely separated from each other. Female clypeus wider than high,
narrowly truncate or emarginate. [Hokkaid6 - Kyushu, N. Ryukyusl .......... Ancistrocerus
13. Parastigma of fore wing much more than half as long as stigma, sometimes even as
long as stigma. Posterior end of tegula not reaching that of parategula. Tergite 1 without
lamella posteriorly..................................................................................................................... 14
- Parastigma half or less as long as stigma. Posterior end of tegula reaching or extending
beyond that of parategula ........................................................................................................ 15
14. Scutellum and posterior part of meso scutum finely and sparsely punctate, slightly
shining. Metanotum depressed medially (very weakly in the Japanese forms) and
21
bluntly projecting laterally. [Honshu - Kyiishui Ryukyusl ...................................Rhynchium
- Scutellum and posterior part of meso scutum strongly and densely punctate, dull.
Metanotum not projecting laterally. Propodeal crest well developed, with a pair of
upper processes. Tergites 3-6 basally with many very large punctures (visible only when
the segments are unusually extended). [Hokkaido - Kyiishu, Ryukyus, Ogasawaral .......
.... ........ ........ ... ..... ...... ..... ..... ... ..... ...... .......... ................ ................ .............. ............. ... Anterhynchium
15. Pronotal carina often interrupted medially (dorsally). Pronotum with a pair of pits in
the lower part of anterior vertical face, and sometimes with transverse striae above the
pits. Tegula wide. Gastral segment 1 distinctly narrower than segment 2. Body relatively
slender. [Hokkaido - Kyiishu; Ryukyus; Ogasawaral ....................................... Stenodynerus
- Pronotal carina not interrupted medially. Pronotum without pits and striae on the
vertical face ................................................................................................................................. 16
16. Tergites 1-5 each with a dark small spots in the brownish or yellowish apical band.
Propodeum without crest. Male antennal hook very small. Basal part of sternite 2 with
14-18 strong, longitudinal carinae. [e. Ryukyusl ............................................ Okinawepipona
- Tergites 1-5 without such dark spots ....................................................................................... 17
17. Tergite 1 with a posterior lamellate area. Pronotum with punctures on the lateral parts
of anterior vertical face. Clypeus ( 5j'. ;]') higher than wide, only rarely as high as wide or
slightly wider than high. Posterior end of tegula reaching or extending beyond that of
parategula. Propodeum often with a pair of upper processes. [Hokkaido - Kyushu;
Ryukyus; Ogasawaral ............................................................................................... Euodynerus
- Tergite 1 without posterior lamellate area. Pronotum without punctures on anterior
vertical face. Clypeus (5j'. ;]') much wider than high; male clypeus deeply incised
apically. Posterior end of tegula produced beyond that of parategula. Propodeum
without upper processes. [Hokkaido, Honshul ................................................... Allodynerus
Genus Discoelius Latreille
Discoelius Latreille, 1809, Gen. Crust. Ins. 4: 140 (subgenus of Eumenes Latreille)(type species: Vespa
zunalis Panzer, 1801, monotypic).
Tritodiscoelius Dalla Torre, 1904, Gen. Ins. 19: 18 (division of Discoelius)(type species: Vespa zonalis
Panzer, 1801, designated by Bequaert and Ruiz, 1942).
Japanese name: Futasuji-suzubachi Zoku.
Latreille's genus Discoelius was characterized by Saussure (1852) as follows. " Labium
moderately long, large; labial palp 4-segmented; segment 1 slightly widened apically.
Maxillary palp 6-segmented; segments 1-3 long (3)2>1), and the rest shorter; galea rather
short. Mandible obliquely truncate apically, wider at apex than at base. Antenna long,
club-shaped, inserted at the middle of the head. Head slightly concave posteriorly; the
sides of head not entirely occupied by the eyes; ocular sinus deep. Thorax long. Propodeal
groove well defined. Gaster with petiole (lst segment) which is strongly widened in the
middle then becomes oval; segment 2 bell-shaped, longer than wide. Fore leg with very
long, curved femur and much shorter tibia".
Saussure recognized three species groups (divisions I-III) in the genus, and Dalla
Torre (l904) named them Protodiscoelius (New World), Deuterodiscoelius (Tasmania), and
Tritodiscoelius (Palearctic). The last division, to which the Asian forms belong, was
22
characterized by Saussure as follows: "Galea short; maxillary paIp shorter than the basal
part of maxilla, rather thick. Petiole short, campanulate in the midst, strongly swollen
dorsally and rimmed posteriorly. Cubital cell 2 with distinct border to radial sector (offrant
un bord radial sensible)".
Later, Bequaert and Ruiz (1942) (after Carpenter, 1986) raised Dalla Torre's divisions
to genera, and Tritodiscoelius was synonymized with Discoelius. Only one species (D.
japonicus) is known to occur in Japan in spite of the fact that three other species have been
recorded from the Far East (Yasumatsu, 1934; Vecht and Fischer, 1972), D. manchurianus
Yasurnatsu from South Manchuria, D. zonalis (Panzer) from Ussuri and Sakhalin, and D.
esakii Yasurnatsu from Taiwan.
Discoelius japonicus Perez
(Figs. 20-26)
Discoelius japimicus Perez, 1905, Bull. Mus. Hist. Nat. Paris 1905(2): 85 (~)(type loc.: Japon central);
Yasumalsu, 1930, Mushi, 3(1): 35; Yano, 1932, Icon. Ins. Jpn. p. 312; Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat.
Ed. 3: 292, pI. 146, fig. 15.
Japanese name: Futasuji-suzubachi.
Diagnosis. Good descriptions were given for both the sexes by Yasumatsu (1930,
1934). Body length (h+th+t1+2): 12.0-15.0 mm in .<f., 7.5-12.0 mm in d'. Fore wing length:
9.5-14.0 mm in .<f., 7.5-9.5 mm in J'. In the female, the lamella of gastral tergite 2 is usually
developed and slightly reflex, but in the specimens from Hokkaido and Fukushima-ken
(Honshu) it is often very narrow and not reflex at all. Yellow and orange body markings
are relatively stable throughout the distribution range. In the specimens from Hokkaido,
however, these markings are generally reduced; sometimes clypeus is almost wholly black
and a spot just above the antennal socket is often lost.
The male is readily distinguished from the female by the following points as well as
by usual sexual characters: posterior face of propodeurn more strongly rugoso-reticulate
and rather shining; punctation on the sides of tergite 1 stronger; tergite 3 as well as 2 with
a developed lamella posteriorly that is distinctly reflex; yellow markings on mandible and
clypeus much larger, while antennal scape entirely blackish, and supra-antennal spot
always lost.
Material examined. Hokkaido: 1 ~, izumi, Niikappu, 3 viii 1974 (T. Sunose), 1 ~, Hakken-zan, Sapporo,
13 vii 1975 (H. Taoka), 1 ~, Shintoku, 16 vii 1977 (M. Kiuchi), 1 ~, Toyotaki, Sapporo, 28 viii 1977 (SKY), 1 ~,
Muine-yama, Sapporo, 9 viii 1978 (K. Kawamura), 1 ~,Nukabira, 19 vii 1979 (SKY), 1 ~, Kamui, Asahigawa,
19 vii 1979 (T. Inaoka), 1 ~,Heiwa-no-taki, Sapporo, 3 viii 1979 (K. Kawamura).
Honshu: Yamagata-ken - 1 ~, Nan'yO-shi, 11 vii 1976 (HI), 1 ~, Oguni, 1 viii 1976 (HI), 2 J' J', Nan'yOshi, 13 vii 1978 (HI), 2~ ~, 11-26 vii 1981 (HI), 1 J', Tamagawa, Oguni, 24 vi 1981 (KB); Fukushima-ken -1~,
Yunohana, Aizu, 21 vi 1977 (HI), 1 ~, same loc., 9 vi 1978 (HI), 1 ~, same loc., 23 vii 1978 (HI), 1 ~, Mishima,
24 vi 1980 (HI), 2~~, Tadami, 25 vi 1980 (HI), 2~~, Yunohana, Aizu, 27 vii 1980 (HI); Miyagi-ken -1~,
Rifu, 8 vi 1980 (K Goukon); Njigata-ken - 1 ~, Asahi-mura, 21 ix 1965 (KB), 9~ ~, Maki, 30 v - 3 vi 1973
(reared by YM), 1 ~, Shidai-hama, 10 vii 1974 (HI), 1 ~, Murakami, 21 vi 1976 (T. Sunose), 1 ~, Nagaoka, 10
vii 1976 (SKY), 1 ~,Kakuda-san, 8 vii 1976 (SKY), 1 ~,Senami, 29 ix 1979 (KB), 1 J', same loc., 5 ix 1981 (KB),
3~~, Sato, Suibara, 12-13 x 1981 (A. Seino); Tochigi-ken - 1~, Uisunomiya, 22 viii 1970 (T. Hasegawa);
Gumma-ken - 2 J' J', Hotaka-san, 8 viii 1976 (HI); Shizuolca-ken -1~, Hirano, Shizuoka, 18 ix 1977 (T. Hattori);
Nagano-ken - 1 ~ , 6machi, 5 vi 1982 (W. Miyata).
Kyiishii: Kagoshima-ken - 1 ~, Kagoshima-shi, 8 vi 1984 (5. Higashi).
23
o
o
0
.G22
.,.....j
~
....
20
,/'"
,/
FI,I>';,'
25
Figs. 20-25. Discoelius japonicus. 20, head in front (.!f. ); 21, terminal segments of male antenna; 22,
tegula and parategula; 23, gaster in profile (.!f.); 24, apical margin of gastral tergite 2 in profile
(d'); 25, apex of mid tibia (.!f.).
Distribution. Hokkaido; Honshu; Shikoku; Kyfishu; Osumi Is. (Tane-ga-shima; Yakushima); Amami Is. (Amami-Oshima). Korea (Kim, 1970); China (Lee, 19&5).
Taxonomic notes. This species is easily distinguished from all other Japanese eumenids
by having two apical spurs on the mid tibia. It disagrees with the diagnosis given by
Saussure (1&52) for Div. III of Discoelius in that maxillary palp is longer than the basal part
of maxilla and rather slender.
Biology. Masuda (1937) and Tsuneki (1970) gave detailed accounts of nesting biology
of D. japonicus in Japan. This species is unique among the Japanese Eumenidae in utilizing
plant leaf pieces as walls partitioning the nest into cells. According to Masuda, nests are
made in bamboo and reed tubes set up as trap nests, or, under natural conditions, in
tunnels in dead tree twigs bored by coleopterous larvae and in broken stems of herbs. The
female wasp first applies pellets of mesophyll mixed with her saliva as an adhesive to a
suitable part in the tube, then collects and carries a leaf piece into it and attaches it to the
applied adhesive. Subsequent leaf pieces are used one by one to complete a wall. In total
an average of 4.1 leaf pieces (range: 1-12) are needed for each wall. Wasps collect material
from broad-leaved trees such as Rosa spp. and Prunus avium. A nest consists of 4-11 cells,
of which 30-83 % are "empty" cells in which 2-9 leaf pieces are usually carried. These cells
seem to prevent brood cells from parasitism. An egg is hung in a cell from the ceiling by a
thread of approximately 1 mm long, then 2-10 (mean: 5.4) caterpillars of the moth families
Pyralidae and Tortricidae are carried in. The full-grown larva of wasp measures on
average 12.3 mm in the male and 14.5 mm in the female. The cocoon is pale brown in color
and cylindrical in shape. Although Yasumatsu (1930) mentions that male wasp must be
quite rare (his material contained no male), Masuda's (1937) data shows an approximately
1:1 sex ratio in at least collected nests that produced adult wasps. This species is
considered to be trivoltine in Yamanashi-ken, Honshu (Masuda, 1937).
24
Parasitoids: Chrysis (Hexachrysis) zetterstedii, C. daphne, C. ignita (Hymenoptera,
Chrysididae)(Baba, 1937; Tsuneki, 1973), Melittobia acasta (Hymenoptera, Eulophidae)
(Maeta & Yamane, 1974, referred to as M. japonica; Maeta, 1985), a bee fly (Diptera,
Bombyliidae)(Masuda, 1937), Amobia signata (Diptera, Sarcophagidae) and Macrosiagon sp.
(Coleoptera, Rhipiphoridae)(Yamane & Maeta, unpubl.).
Genus Stenodynerus Saussure
Stenodynerus Saussure, 1863, Mem. Soc. Phys. Hist. Nat. Genev. 17: 228 (as division of subgenus
Odynerus of genus Odynerus Latreille)(type species: Odynerus chinensis Saussure, 1863, designated by Bohart,
1939); Vecht, 1966, Entomol. Berichten, 26: 163 (as genus); Carpenter, 1986, Psyche, 93: 85 (as genus).
Nannodynerus Bliithgen, 1938, Konowia, 16: 281 (as subgenus of nEuodynerus B1iithgen n )(type species:
Uonotus leutonicus B1iithgen, 1937, original designation).
Japanese name: Chibi-dorobachi Zoku.
The wasps of this genus have a small and rather slender body, not exceeding 10 mm
Fig. 26. Distribution of Discoelius japonicus.
25
in total length in at least Japanese species. Gastral tergite 1 is distinctly narrower than
tergite 2, but not petiolate. Anterior vertical face of pronotum with a pair of pits in its
lower portion, and sometimes with transverse striae above the pits (Fig. 13). Cephalic
foveae are usually very small and inconspicuous, located just in front of a slightly curved
carina (Figs. 29-31). In frauenfeldi, rufomaculatus, chinensis and c/ypeopictus the foveae are
very close to each other and sometimes connected (Figs. 29,30). In ogasawaraensis they are
quite inconspicuous, usually not distinguished from nearby punctures. In tokyanus and
/cusigematii (Fig. 31) they are apart from each other as usual with many eumenids.
Maculation of tegula and parategula is often useful to distinguish between the species
(Figs. 48-57).
The genus is mainly Holarctic in distribution, with fewer species in Neotropical and
Oriental regions. Australia lacks this genus (Taylor et aI., 1985). The Palearctic species
were revised by Gusenleitner (1981), and 34 species were enumerated in five tentative
species groups. In Japan eight species occur and are tentatively divided into four species
groups (Yamane & Tano, 1987). Structural characteristics for the Japanese species are given
in Gusenleitner (1981) and Yamane and Gusenleitner (1982), and are accordingly not
repeated here.
Key to the Japanese species of Stenodynerus
1. Lateral sides of propodeum only finely punctate, not reticulate. Clypeus not distinctly
emarginate apically ( ~ ). Gastral tergites 1 and 2 rufous, with yellow apical bands ......... .
.................................................................................................. s. ogasawaraensis Yam. et Gusnlt.
- Lateral sides of propodeum coarsely punctate or reticulate at least partly. Clypeus more
or less emarginate apically ( ~). Tergite 2 always black, with a yellow or rufous apical
band ............................................................................................................................................... 2
2. Propodeum without horizontal space (propodeal shelO behind metanotum. Anterior
vertical face of tergite 1 clearly demarcated from the posterior horizontal part, almost
impunctate, and shining (in the male this demarcation is less distinct, and the vertical
face often punctate). Anterior vertical face of pronotum without transverse striae .
......................................................................................................................................................... 3
- Propodeum with a narrow but distinct shelf. Tergite 1 with round border between
anterior vertical face and posterior horizontal part; the former strongly punctate in its
upper portion. Anterior vertical face of pronotum with transverse striae in its upper
part ................................................................................................................................................ 4
3. Clypeus distinctly wider than high (~). Anterior vertical face of pronotum coarsely
punctate. Posterior horizontal part of pronotum and mesoscutum with large punctures
that are larger than interspaces ................................................................... s. tokyanus (Kost.)
- Clypeus as wide as long, or at most only slightly wider than long ( ~ ). Anterior vertical
face of pronotum with much finer and sparser punctation. Posterior horizontal part of
pronotum and mesoscutum with smaller punctures that are often as large as or smaller
than interspaces .......................................................................... s. kusigematii Yam. et Gusnlt.
4. Occipital carina bent at the level of clypeal base, forming a distinct angle there. Pronotal
carina distinct dorsally, forming a narrow transparent lamella, interrupted only in the
middle ........................................................................................................................................... 5
- Occipital carina more smoothly curved, without distinct angle. Pronotal carina less
26
developed, dorsally not forming a lamella .............................................................................. 6
5. Gastral tergite 1 largely rufous or orange. Propodeum without upper teeth behind
metanotum. Tergite 2 superficially punctate..................... S. rufomaculiltus Yam. et Gusnlt.
- Tergite 1 black, with a yellow apical band. Propodeum often with distinct teeth behind
metanotum. Tergite 2 coarsely punctate; punctures only slightly smaller than those on
tergite 1. ...................................................................................................... S. frauenfeldi (Sauss.)
6. Tibiae of all legs yellow ( .5f.). Clypeus with fine, sparse, superficial punctation (.5f.).
Terminal segment of male antenna large, seen from below, wide and parallel-sided; its
apical half yellow; segment 11 ventrally slightly concave to receive the terminal
segment. Innerorbit with a yellow stripe in its lower part (6'). Tegula marked with
yellow on anterior and posterior parts ................................................. S. clypeopictus (Kost.)
- Mid and hind tibiae brown or black ( .5f.). Clypeus with deep and dense punctures ( .5f.).
Terminal segment of male antenna much more slender, narrowed toward apex, wholly
blackish; ventral face of segment 11 not concave. Innerorbit without yellow stripe (6').
Tegula usually blackish brown, only rarely with a small spot on its anterior part .
......................................................................................................................... S. chinensis (Sauss.)
Frauenfeldi group
Stenodynerus frauenfeldi (Saussure)
(Figs. 27, 29, 32, 39, 43, 46-48)
Odynerus (Lionotus) frauenfeldi Saussure, 1867, Reise der Novara, Zool. 2(1) Hym. p. 15 (!f) (type loc.:
Shanghai, China); Yasumatsu, 1935, Kontyu, 9: 225-227 (part).
Odynerus (Hoplomerus) nigriclypeatus Sonan, 1930, Trans. Nat. Hist. Soc. Formosa, 20: 356 (!f }(type loc.:
Kagoshima, Kyushu).
Odynerus apiciornatus: Yano, 1932, Icon. Ins. Jpn. 1st ed. p. 309, fig. 600 (misidentification).
Stenodynerus frauenfeldi: Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat. Ed. 3: 297, pI. 149, fig. 7; Gusenleitner,
1981, Polsk. Pismo Entomol. 51: 287-289; Yamane and Gusenleitner,1982, Rep. Fac. Sci. Kagoshima Univ. 15:
113 (in key), 116-118.
Japanese names: Kiobi-chibi-dorobachi (Chibi-dorobachi; Hime-futaobi-dorobachi).
Diagnosis. Body length (h+th+t1 +2): 7.0-8.5 mm in .5f., 5.5-6.5 mm in 6'. Fore wing
length: 6.5-7.0 mm in .5f., 5.0-6.0 mm in 6'.
Female. Black, with the following parts yellow: a basal markings on c1ypeus (c1ypeus
usually wholly black), a round frontal spot, a basal marking on mandible (sometimes lost),
a minute spot on temple, a pair of markings on pronotum anteriorly, tegula extensively,
parategula, a spot on dorsal mesepisternum, metanotum largely, a pair of spots on
propodeum dorsolaterally, regular apical bands on gastral tergites 1 and 2, posterolateral
corners of sternite 2 (the sternite sometimes with a complete apical band), anterior face of
fore tibia.
Male. More extensively marked with yellow as follows: c1ypeus wholly, mandible
largely, frontal spot, antennal scape below, small spot on temple, an anterior band on
pronotum (laterally narrower than in .5f.), anterior faces of all tibiae, those of mid and hind
coxae (often reduced). Other markings on alitrunk and gaster as in .5f. (sternite 2 almost
always with an apical band).
Material examined. Hokkaido: 1 J'2!f!f, Sapporo, 5 vii 1954 (K. Kamijo), 1!f, Esashi, 9 ix 1958 (T.
27
Kumata), 2 d' d', Toyotaki, Sapporo, 29 viii 1979 (SKY), 1.!f , Misumai, Sapporo, 6 viii 1980 (5. Makino).
Honshu: lwate-ken - 1.!f, KUriyagawa, Morioka, 23 vi 1964 (YM), 2.!f .!f, JohOji, 21 viii 1982 (HI), 3 d' d'2
.!f.!f, Kanegasaki, 2 viii 1987 (SKY); Fukushima-ken - 1 d', Yunohana, 21 vi 1977 (HI); Niigata.Jcen - 2.!f.!f,
Muroya, Mikawa, 1 ix 1977 (HI), 6 d' d'1.!f, Sato, suibara, 15 vii 1979 (A. Seino), 1.!f, Senami, 2 vii 1983 (KB),
16', Shibata, 3 ix 1979 (HI), 1 d', Ishizumi, Asahi, 27 ix 1984 (KB); lbaraki-ken - 2 d' d', Tsuchiura, 29 viii 1976
(SKY); Nagano-ken - 1 d'1.!f, Minamiminowa, Ina, 6-8 vi 1962 (YM), 7 d' d'3.!f .!f, Habiro, Ina, 31 vii 1962
(yM), 2.!f .!f, Saku, 27-28 vii 1973 (YM), 1.!f, Todo, Otari, 26 viii 1982 (KB); Yamanashi-ken - 1 d'1.!f, Wadatoge, Kofu, 24 vii 1965 (K. Iwata); Ishikawa.Jcen -1.!f, Kaga, Haku-san, 17 vii 1953 (K. Nohara); Fukui-ken - 1.!f,
Ono, 10 vii 1971 (yH), 1 d', same loc., 15 ix 1971 (YH); Gifu-ken -1.!f, Horado, 7 viii 1982 (Y. Takai); Aichi-ken16', Irako, Atsumi Pen. 6 viii 1977 (Y. Takai); Wakayama-ken - 1.!f, Kiitanabe, 4 viii 1965 (K. Iwata); Hy6go-ken
-1 d', sasayama, Tamba, 24 vi 1950 (K. Iwata); Shimane-ken -1.!f, Dba, Matsue, 10 vii 1986 (N. SUgiura).
Nanatsu-jima Is. Oshikawa-ken): Hegura-jima - 2 d' d', 26 v 1983 (R. Ohgushi); Oshima - 4 d' tI.!f, 11 vi
1983 (R. Ohgushi).
Izu Is. (TokyO-to): Hachij6 jima - 2d' 6', Okagou - Fuji, 26 v 1964 (Y. Hirashima & M. Shiga), 1 d',
Kamogawa, Mitsune, 12 vii 1987 (H. Takahashi), 1.!f, Okagou - Eigou, 16 viii 1987 (H. Takahashi); Ao-gashima -1.!f, Yasundogou, 8 vii 1987 (H. Takahashi).
Shikoku: K6chi-ken - 1.!f, Monobe, Nankoku, 2 v 1974 (SI), 1.!f, Okoyama, Nankoku, 2 vii 1975 (sI);
Kagawa-ken - 4 6' 6', Miki, 23 vii 1963 (K. Iwata); Ehime.Jcen - 16', Matsuyama, 10 viii 1969 (HI); Tokushima-ken
-1.!f, Tsudayama, 22 viii 1964 (H. Iwasaki).
Kyushu: Fukuoka.Jcen - 1.!f, shingfl, Fukuoka-shi, 9 x 1983 (AN), 1.!f, Ushirog6chi, Kurogi, 27 vii 1983
(Y. Takai); Nagasaki.Jcen - 1 d', Haraguchi, Omura, 18 vi 1966 (R. Ohgushi), 1.!f, same loc., 24 viii 1966 (R.
Ohgushi); Kumamoto-ken - 2 d' d', Kagami, 5 v 1982 (M. Maeda), 5 d' d'2.!f .!f, same loc., 31 v 1982 (M. Maeda);
Kagoshima-ken - 4d' d'H, Ichiki, 11 vi 1984 (AN), 5d' 6'3.!f.!f, Jingfl, Kirishima, 3 vii 1981 (SKY), 1.!f,
Shiroyama, Kagoshima-shi, 16 x 1980 (51), 2.!f.!f, same loc., 30 vii 1981 (SKY), 56' d'2.!f.!f, Korimoto,
Kagoshima-shi, 6-7 viii 1981 (SKY), 2 d' 6', Ibusuki - Kiire, 13 x 1963 (H. Tanaka), 1 d', Izashiki, Sata, 3 viii
1963 (A. Tanaka).
Tsushima Is. (Nagasaki-ken): Kami.;zgata -1 d', Nii, Toyotama-son, 20 viii 1968 (I. Hiura); Shimo.;zgata-1
d', lzuhara, 27 vii 1986 (K. Nakamine).
Goto Is. (Nagasaki-ken): Aka-shima - 1 6'1.!f, 17 viii 1976 (J. Nakayama).
N. Ryukyus: Ie-jima (Vji Is.) -1 d', 11 v 1985 (H. Fukuda); Tane-ga-shima- 2d' d'1.!f, Kaminaka, 11-12 vii
1983 (SKY), 2 d' d'2.!f .!f, Hamada, 11 vii 1983 (SKY), 3 d' b.!f, same loc., 1-2 viii 1984 (SKY); Mage-shima- 3 d'
d', 22 vii 1984 (5. Watahiki); Yaku-shima - 36' d'1.!f, Miyanoura, 8-11 viii 1981 (SKY), 5 d' 6', Shitogo, 10 viii
1981 (SKY), 16', AmbO, 11 v 1981 (SKY), 1.!f, Kurio, 4 x 1981 (Y. Takai); Kuchinoerabu-jima-1.!f, Hommura, 15
vi 1989 (H. Watanabe).
C. Ryukyus: Amami-&hima -1 d', Naze, 20 vi 1987 (SKY), 6d' 6'1.!f, Setouchi, 22 vi 1987 (SKY), 2d' 6'1
.!f, Shinokawa, 23 vi 1987 (SKY).
Distribution. Hokkaido; Honshu; Nanatsu-jima Is. (Oshima; Hegura-jima); Izu Is.
(Hachijo-jima; Hachijo-kojima; Ao-ga-shima); Shikoku; Kyushu; Tsushima Is. (Kami-agata;
Shimo-agata); Goto Is. (Aka-shima); Osumi Is. (le-jima; Tane-ga-shima; Mage-shima;
Yaku-shima; Kuchinoerabu-jima); Amami Is. (Amami-oshima); Ogasawara Is. (introduced
?). China. Yasumatsu (1935d) recorded "Odynerus frauenfeldi" from Sado-ga-shima.
Taxonomic notes. Three species <frauenfeldi, chinensis and tokyanus) might have been
confused in Japan under the name Odynerus frauenfeldi or Stenodynerus frauenfeldi (e.g.,
Yasumatsu, 1935b). Further, the description of the female of Odynerus apiciornatus by Yano
(1932) well agrees with Stenodynerus frauenfeldi.
In a few female speciems from Kyiishu and the N. and C. Ryukyus, the scutellum and
mid tibia are marked with yellow. But the color pattern is rather stable throughout the
distribution range in Japan.
Yamane and Gusenleitner (1982) pointed out that abscissa 4 of radius is as long as or
slightly longer than abscissa 3 in S. frauenfeldi, but distinctly longer than the latter in S.
rufomaculatus. But, in some specimens (especially males) of frauenfeldi, abscissa 4 is much
28
29
0
()
0
...........
0
0
()
-.sv
31
30
0
0
C)
........
@:2S fila U
3
43
&a6 3;a? @3B
Figs. 27-42. Various characters in the Japanese Stenodynerus. 27,28, head in profile of frauenfeldi
(27) and chinensis (28), showing the condition of occipital carina; 29-31, ocellar region (5f-) of
frauenfeldi (29), rufomaculatus (30), kusigematii (31); 32-38, terminal segments of male antenna
(from below) in frauenfeldi (32), rufomaculatus (33), chinensis (34), clypeopictus (35), tokyanus (36),
kusigematii (37), ogasawaraensis (38); 39-42, penis of male genitalia in frauenfeldi (39),
rufomaculatus (40), chinensis (41), c/ypeopictus (42).
longer than 3, and in the female specimens of rufomaculatus from Yoron-to abscissa 4 is
often as long as 3. In some males of frauenfeldi abscissa 2 is almost lacking.
Biology. As mentioned above, in Japan two or three species have been confused
under the name frauenfeldi even by taxonomists. The biology of Japanese Stenodynerus has
been studied mainly by Iwata (1938b, 198Oa) and Tsuneki (1961, 1969). The material used
by Iwata (1938b) was identified by Yasumatsu, who erroneously believed that only one
species of this genus occurred in Japan. When I examined Iwata's collection at Kobe
University, I found that specimens of three Stenodynerus species (frauenfeldi, chinensis and
tokyanus) and Allodynerus mandschuricus were placed together under the head
identification label "Stenodynerus frauenfeldi". Although the photo of a nesting female in
Iwata (1982) is never of S. frauenfeldi (it is probably of s. tokyanus or even of A.
mandschuricus), the figure drawn by him (Iwata, 198Oa, p. 39) is correctly a female of s.
frauenfeldi. The female illustrated by Tsuneki (1961) is also no doubt S. frauenfeldi. The
following description may include information about the biology of two or three species.
Nests are constructed in pre-existing hollow tubes (3-4 mm in diam.), such as the pith
hollow of reed cane and the stems of bamboo-grass and flax. As these plants were
formerly utilized as blinds, fences and roofs, Stenodynerus females were abundant around
29
houses in rural regions. The nest consists of two or three brood cells (maximum: 6), and
usually two long empty cells, one in the back and the other in front. Cells are partitioned
with mud. Thirteen to 76 leaf-mining microlepidopterous larvae of the families
Lyonetiidae, Gracillariidae and Gelechiidae (4-5 mm long) are stored in each brood cell.
Hunting habits and prey species are given in Iwata (1963, 1980a). The egg is laid before
hunting, but Tsuneki (1969) suspected that the female wasp carried one or two prey
caterpillars into a cell prior to oviposition (behavior of the female within nest was not
observed). The color vision and figure discriminating capacity was intensively studied by
Tsuneki (1961).
Parasite: Pseudoxenos minor (Strepsiptera, Stylopidae) (Kifune & Maeta, 1987).
Parasitoids: a phorid fly (Diptera) mainly attacking prey?, and Chrysis viridula
(Hymenoptera, Chrysididae).
5tenodynerus rufomaculatus Sk. Yamane et Gusenleitner
(Figs. 30, 33, 40, 47, 49)
Stenodynerus rufCJmJlcuiatus Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ. 15: 118-119,
figs. 4, 8 (~ J') (type loc.: Okinawa-jim a).
Stenodynerus rufomaculatus kikaiensis: Yamane and Tano, 1987. Trans. Shikoku Entomol. Soc. 18: 328 (~
J')(type loc.: Kikai-jima, Amami Is). Syn. nov.
Japanese name: Akaobi-chibi-dorobachi.
Diagnosis. Body length (h+th+t1+2): 6.5-7.0 mm in ~, 5.5-7.0 mm in 6'. Fore wing
length: 6.0-6.5 mm in ~, 4.5-6.0 mm in 6'. Structurally similar to the preceding species,
but differs from the latter in the following points: abscissa 4 of radial vein of fore wing
usually distinctly longer than abscissa 3 (~ 6'), propodeum without distinct processes
behind metanotum ( ~ 6'), clypeus as wide as high ( 6').
In coloration, this species is easily distinguished by orange gastral markings. Orange
are: a large markings on propodeum (in 6' often reduced or lost), posterior horizontal part
of gastral tergite 1 extensively, sternite 1, a wide apical band on tergite 2, a narrower band
on sternite 2. Pair of large markings on pronotum yellow, often tinged with orange. The
following parts are yellow: clypeus largely (in ~ often black in peripheral and lower
parts; in 6' wholly yellow), frontal spot, mandible largely (6'; in ~ black with apical half
ferruginous), antennal scape below (6'), small spot on temple, tegula except for central
part and margins, parategula, a spot on dorsal mesepisternum (often lost in 6'),
metanotum, anterior face of fore tibia (~), anterior face of mid coxa (6'), apical part of
fore and mid femora (6'), tibiae of all legs extensively (6').
Material examined. C. Ryukyus: Kikai-jima - 5J' J'1 ~, Nakazato, 26-27 viii 1984 (S. Watahiki), 6 J' J',
same loc., 18 x 1987 (S1); Kakeroma-jima - 1 ~,Nishiamuro, 28 ix 1987 (SKY); Okinoerabu-jima - 1 J', Kimidome,
9 viii 1967 (T. Murota), 1 J', Shinjo - Tamina, 14 vii 1984 (Y. Takai); Varon-tO - 3J' J'3~~, Maenohama, 4 vi
1989 (SKY); OkinlZWll-jima -1 J', Oku, Kunigami, 4 viii 1982 (YH).
Distribution. Amami Is. (Kikai-jima; Kakeroma-jima; Okinoerabu-jima; Yoron-t6);
Okinawa Is. (Okinawa-jima; Kume-jima).
Taxonomic notes. Yamane and Tano (1987) regarded the population of Kikai-jima as a
subspecies (kikaiensis), but the present material shows that the supposed difference is not
constant. Up to the present this species has not been collected on Amami-6shima, where
the closely related S. frauenfeldi is relatively common.
30
Biology. On Yoron-t6 many female wasps were seen flying around limestones along
the coast, but nesting biology is not yet known.
Parasite: Pseudoxenos sp. (new record).
Stenodynerus chinensis simillimus Sk. Yamane et Gusenleitner
(Figs. 28, 34, 41, 44, 47, 50-52)
Odynerus (Lionotus) chinensis Saussure: Schulthess, 1934, Arb. Morph. Taxon. Entomol. 1: 91.
Odynerus (Uonotus) frauenfeldi: Yasumalsu, 1935, Kontyil, 9: 225-227 (part).
Stenodynerus chinensis simillimus Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ. 15:
119-121, fig. 14 (-'i'- d') (type loc.: Morioka, Honshu).
Japanese name: Kataguro-chibi-dorobachi.
Diagnosis. Body length (h+th+tl+2): 7.0-8.5 mm in ~, 5.5-6.5 mm in 6'. Fore wing
length: 6.5-7.0 mm in ~,5.0-6.0 mm in 6'.
Female. Black, with the following parts yellow: round frontal spot, a stripe on
mandibular base, minute spot on temple, a stripe on antennal scape below, a pair of
markings on pronotum anteriorly (often reduced or lost), parategula, metanotum
anteriorly, apical bands on gastral tergites 1 and 2 and stemite 2, anterior face of fore tibia.
Clypeus usually wholly black, but rarely with yellow spots in upper portion. Tegula
wholly blackish brown, only rarely with a yellow marking anteriorly.
Male. Similar to the female, but differs as follows: clypeus and mandible almost
wholly yellow, pronotal and metanotal markings much more reduced, anterior face of mid
coxa, tibiae of all legs extensively yellow, tarsi also marked with yellow.
Material examined. Hokkaido: l-'i'-, Jozankei, 24 vi 1951 (K. Kamijo), l-'i'-, Bibai, 15 viii 1976 (A. Seino), 1
-'i'-, same loc.,3 x 1976 (K. Kamijo), l-'i'-, Toyotaki, Sapporo, 26 viii 1979 (SKY).
Honshu: Aomori-lcen - 2 -'i'- -'i'-, Yunokawa, Shimokita, 23 vii 1965 (T. Naito); lwate-lcen - l-'i'-, Kuriyagawa,
Morioka,S ix 1968 (YM), 1 d'1-'i'-, JohOji, 21 viii 1982 (HI), 2d' d'1-'i'-, Kanegasaki, 2-3 viii 1987 (SKY), l-'i'-,
Oshuku, Shizukuishi, 7 viii 1987 (SKY); Yamagata-lcen -16', Chojuga-hara, Oguni, 22 vi 1979 (HI); Miyagi-lcen
- l-'i'-, Rifu, 14 ix 1980 (K. Goukon); Niigata-ken - l-'i'-, Nagaoka, 11 vii 1976 (S. Higuma), l-'i'-, Shibata, 28 viii
1976 (HI), l-'i'-, Muroya, Mikawa, 21 viii 1977 (HI), l-'i'-, Shidai-hama, 23 ix 1980 (HI), 2 d' d'3 -'i'- -'i'-, Senami, S18 vi 1983 (KE), 1 d', Kurokawa, 7 ix 1984 (KB), l-'i'-, Sarusawa, Asahi, 29 ix 1984 (KB); Ibaraki-lcen - l-'i'-, Mt.
Mamizo, 31 viii 1975 (SY), l-'i'-, Tsuchiura, 9 viii 1987 (SKY); Chiba-ken -1 d', Abiko, 10 v 1975 (SY); Naganoken - 1 d', Minamiminowa, Ina, 6 vi 1982 (YM); Gifu-lcen - 1-'i'-, Oyama, Seki, 2 x 1977 (Y. Takai); Fukui-ken - 1
-'i'-, Ono, 30 v 1973 (YH); Osaka-fu -1-'i'-, Kimi-toge, 15 vi 1975; Kyoto-fu -1 d', Shingoji, Takao, 27 vii 1965 (M.
Iwata); Shimane-ken -1-'i'-, Mt. Sambe, 28 viii 1982 (YM), l-'i'-, Mt. Hoshigami, 28 viii 1985 (M. Goubara).
Shikoku: K&hi-lcen -1-'i'-, Uranouchi, 17 viii 1933 (Y. Sugihara), 1 d', Monobe, Nankoku, 18 iv 1975 (SI),
2 d' d', Godaisan, K6chi-shi, 7 v 1955 (SI); Ehime-lcen - 1 d', Malsuyama, 10 viii 1969 (HI); Tokushima-lcen - 1-'i'-,
Tsudayama, Tokushima-shi,28 vii 1964 (H. Iwasaki).
Kyushu: Nagasaki-lcen - l-'i'-, Haraguchi, Omura, 27 viii 1966 (R. Ohgushi), 1-'i'-, same loc., 7 ix 1966 (R
Ohgushi); Kaga;hima-ken - 1 d', Iriki, 6 ix 1984 (AN), 1-\'-, Kurino, 18 v 1986 (K. Nakamine), 1 d', Shiroyama,
Kagoshima-shi, 30 vii 1981 (SKY), 1 d', KOrimoto, Kagoshima-shi, 17 viii 1981 (SKY), 2 d' d', Kanoya, 8-15 ix
1981 (SI), l-'i'-, Sata, 10 vi 1979 (H. Nagase), 2 d' d', Onakano, Sata, 10 x 1978 (H. Nagase), 3 d' d', Sata, 3-4 v
1988 (SKY).
Tsushima Is. (Nagasaki-ken): Kami-ilgata - l-'i'-, Sasuna, 17 ix 1977 (Tominaga), 4 d' d', Oboshi-yama, 24
vii 1979 (A. Seino); ShimO-1lgata - 2d' d', Izuhara, 27 vii 1986 (K. Nakamine).
Naga-shima Is. (Kagoshima-ken): Naga-shima - 3 d' 6'1-\'-, Shoura, 27 viii 1984 (SKY).
Koshiki-jima Is. (Kagoshima-ken): Kamika;hiki-jima - 1-\'-,6 ix 1984 (M. Maegata).
N. Ryukyus: Take-shima - l-'i'-, 3-5 vi 1982 (KT); Ii5-jima - 1 d', 1-3 vi 1982 (SKY), 2d' d', 19-22 v 1983
(SKY); KUrD-shima - 40d' d'15-'i'- -'i'-, Osato, 31 viii - 5 ix 1981 (SKY); Tane-ga-shima - 1 d', Hamada, 11 vii 1983
31
(SKY), 26' 6', Kaminaka, 11 vii 1983 (SKY), 16', Kamome, 5 viii 1986 (M. Tatsuno); Yaku-t;hima -1!f, Koseta,
30 x 1972 (K. Kusigemati), 86' 6'10!f !f, Miyanoura, 8-11 viii 1981 (SKY), 21 6' 6'2!f !f, Shitogo, 10 viii 1981
(SKY), 1!f, Onoaida,9 viii 1981 (SKY).
Distribution. Hokkaido; Honshu; Shikoku; Kyilshu; Tsushima Is. (Kami-agata; Shimoagata); Naga-shima Is. (Naga-shima); Koshiki-jima Is. (Kamikoshiki-jima); Osumi Is.
(Take-shima; Io-jima; Kuro-shima; Tane-ga-shima; Yaku-shima). Other subspecies are
known from northern China and Taiwan (Gusenleitner, 1981).
Biology. The female wasp hunts for tortricid larvae (Yukinari, pers. comm.). See also
under S. frauenfeldi.
Parasite: Pseudoxenos minor.
,
o
S.
frauenfeldi
43
,
o
44
C)
e S. tokyanus
.#
.. S. clypeopictus
0
IZU IS.
0
<J
0
·0
0
etokyanus
"frauenfeldi
.8
.,
.-
45
'I<
Figs. 43-46. Distribution of Stenodynerus species on the Japanese mainlands and Izu Islands.
32
46
Ogasawaraensis group
5tenodynerus ogasawaraensis Sk. Yamane et Gusenleitner
(Figs. 5-7, 38, 54)
Stenodynerus ogasawaraensis Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ. (Earth Sci.
BioI.) 15: 126-127 (.!f J')(type loc.: Chichi-jima, Ogasawara Is.).
Japanese name: Ogasawara-chibi-dorobachi.
Diagnosis. Body length (h+th+t1+2): 7.0-7.5 mm in ~, 5.0-5.5 mm in d'. Fore wing
length: 6.5-7.0 mm in ~, 5.0 mm in d'. This species is easily distinguished from the
Japanese congeners by the female clypeus not distinctly emarginate apically, very faint
pits on the anterior vertical face of pronotum, superficial punctation on mesopleuron and
propodeum laterally, and peculiar color pattern.
This species is endemic to the Ogasawara (Bonin) Islands, and at present two color
forms are known from Chichi-jima and Haha-jima. The phylogenetic position of this
species is not clear, but in structure this species has some character conditions shared by
the frauenfeldi group as follows: pronotal carina weak but clearly visible over whole length
except in the middle, propodeal shelf short but present, and anterior vertical face of
gastral tergite 1 not demarcated from the posterior horizontal portion. In this respect the
following female specimen from Chichi-jima (Omura, 16 v 1972, Y. Kusui leg.) is very
'-bv
,ok
.'
....
tf)
~O . ~,i
.
Q
()
/
Yakushima
1.1
"
• 0
Amami
Is.
~ ~
o~
.5.
chinensis
"S.
frauenfeldi
*5. rufomaculatuS
47
Fig. 47. Distribution of Stenodynerus species in the N. and C. Ryukyus.
33
interesting, because it retains some character conditions that are seen in S. frauenfeldi as
follows: punctation on meso soma especially on mesopleuron rather coarse, c1ypeus
apically emarginate, anterior vertical face of pronotum with weak but discernible
transverse striae in its upper portion, occipital carina with a distinct angle at some
distance from mandible; color pattern as in the typical nominate form of S. ogasawaraensis.
I consider this individual to represent an extereme variation of ogasawaraensis approaching
frauenfeldi in phenotype.
Nesting biology is not known.
Stenodynerus ogasawaraensis ogasawaraensis Sk. Yamane et Gusenleitner
Diagnosis. Female. Black; the following parts yellow: c1ypeus (sometimes with a
blackish marking at apex), a small basal marking on mandible, minute spot on temple,
antennal scape below (yellow often reduced or replaced by reddish brown), pronotum
largely, tegula largely, parategula, dorsal mesepisternum wholly, a longitudinal marking
on ventral mesepisternum (often reduced), a pair of very large markings on propodeum,
narrow apical band on gastral tergite I, wider apical bands on tergite 2 and sternite 2,
coxae of all legs, femora of all legs (hind femur with extensive black area), tibiae of all legs.
Yellow apical bands on tergites I, 2 and sternite 2 sometimes obscure. Mandible largely
brownish, slightly tinged with yellow. Gastral segments 1 and 2 wholly rufous except for
yellow apical bands. Tarsi of all legs blackish brown.
Male. Very similar to the female in coloration, differing therefrom as follows: c1ypeus
wholly yellow, mandible yellow except for teeth, antennal scape below extensively yellow,
yellow marking on ventral mesepisternum absent, lower half of lateral part of propodeum
rufous, gastral tergite 3 with a narrow yellow apical band.
Material examined. Ogasawara Is. (Tokyo-to): Chichi-jima - 1 Sf, Omura, 16 v 1972 (Y. Kusui), 1 Sf,
Tsutsuji-yama, 2 vi 1972 (Y. Kusui), 1 Sf, same loc., 12 vii 1972 (Y. Kusui), 1 6'1 if, 17 viii 1972 (YH)(including
the holotype female), 1 if, 9 vi 1974 (T. Nakane), 3if Sf, Chii6-san, 9-13 viii 1983 (TN); Haha-jima -1 if, 13-26
viii 1980 (H. Kurahashi).
Distribution. Ogasawara (Bonin) Is. (Chichi-jima; Haha-jima).
Stenodynerus ogasawaraensis rufoornatus Sk. Yamane
5tenodynerus ogasawaraensis rufoornatus: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 332
(~ 6')(type loc.: Haha-jima, Ogasawara Is.).
Diagnosis. Female. In structure and coloration similar to the nominotypical subspecies, but much more melanic especially in alitrunk and legs: c1yepus black below,
yellow spot on mandible much reduced, yellow pronotal band less extensive, pronotal
tubercle black, yellow marking on scutellum much reduced, yellow markings on fore and
mid femora less developed, hind femur and tibia wholly blackish brown.
Male. Similar to the female, with the following differences: c1ypeus wholly yellow,
scutellum without yellow marking, tegula divided by a wide blackish bar into upper and
lower portions, legs more extensively marked with yellow.
Material examined. Ogasawara Is. (TokyO-to): Haha-jima-l 6', Nagahama, 11 viii 1983 (TN) (holotype), 1
6'1 if, Kitakoo, 11 viii 1983 (TN); Muk6-jima - 16',23 v 1973 (Y. Kusui).
34
Distribution. Ogasawara (Bonin) Is. (Haha-jima; Muk6-jima).
Taxonomic notes. This fonn, restricted to the Haha-jima group, is not distinguished in
structure from the nominotypical form and may represent a geographical race or
subspecies. However, as already stated, one female of the nominotypical form was
captured from Haha-jima. If the occurrence of the nominotypical form on the island is not
accidental, the relationship of the two forms may be open to criticism.
Clypeopidus group
Stenodynerus clypeopidus (Kostylev)
(Figs. 35, 42, 45, 53)
Odynerus japonicus Matsumura, 1926, Ins. Matsum. 1: 37, pI. 3, fig. 12 (nee. 11) (t) (homonym of
Odynerus japonicus Schulthess, 1908)(type loc.: Nago,Okinawa-jima).
Odynerus clypeopictus Kostylev, 1940, Bull. Soc. Nat. Moscou, Sect. BioI. (N.S.), 49: 28 (S? )(type loc.:
5emiretchie, Kazakh, USSR).
Stenodynerus chinensis: Vecht and Fischer, 1972, Hym. Cat. (nov. ed.), p. 65 (from Okinawa);
Gusenleitner, 1981, Polsk. Pismo Entomol. 51: 289-291 (from Okinawa). .
Stenodynerus clypeopictus: Yamane and Gusenleitner, 1982, Rep. Fae. Sci. Kagoshima Univ. (Earth Sci.
Biol.), 15: 121-122, fig. 16.
Japanese name: Futokagi-chibi-dorobachi.
Diagnosis. Body length (h+th+t1+2): 7.0-7.5 mm in .!f, 5.0-6.0 mm in 6'. Fore wing
length: 5.5-6.0 mm in .!f, 4.0-5.0 mm in 6'. The female of this species is similar to that of S.
frauenfeldi in structure and coloration, but in the former the punctation on clypeus and
gastral tergites is much finer and the legs are more richly marked with yellow than in the
latter. The male is very distinctive in the last antennal segment (hook) not tapering
apically; antennal segment 11 ventrally slightly concave to receive the hook (Fig. 35).
Female. Black, with the following parts yellow: apical 1/3 to 1/2 of clypeus, frontal
spot, minute spot on temple, a triangular marking on mandibular base, antennal scape
below (sometimes reduced to lines), a pair of spots on pronotum anteriorly, outer margin
of tegula, dorsal mesepisternum largely, metanotum largely, narrow apical bands on
tergites 1 and 2, and also on sternite 2, apical part of fore (and sometimes also mid) femur,
tibiae of all legs extensively.
Male. Yellow markings on alitrunk and gaster as in .!f. Head and legs with yellow as
follows: clypeus largely, labrum wholly, frontal spot, minute spot on temple, a line on each
innerorbit, mandible largely, antennal scape below, anterior face of mid (often also fore
and hind) coxa, apical parts of all femora, tibiae and tarsi of all legs (apical parts of tarsi
including tarsal claws often brownish). Apical half of antennal hook yellowish or
ferruginous.
Material examined. Hokkaido: 2t t, Momiji-yama, Ishikari, 5 viii 1968 (TN).
Honshu: Yamagata-ken - 1 t, Nukumi, Oguni, 28 ix 1980 (HI); Niigata-ken - 3 t t, Fukushimagata,
Toyoura, 14 ix 1980 (HI); Nagano-ken - 1 t, Minamiminowa, Ina, 29 v 1963 (YM); Fukui-ken - IS?, Otto, 15 ix
1971 (YH), 1 d', same loc., 23 vii 1973 (YH).
Kyushu: Kagoshima-ken -IS?, Higashi-ichiki, 23 ix 1978 (51).
Tsushima Is. (Nagasaki-ken): Kami~gata -1 t, Nii, Toyotama-son, 20 viii 1968 (I. Hiura).
Naga-shima (Kagoshima-ken): IS?, Shoura, 27 viii 1984 (SKY).
c. Ryukyus: Okinawa-jima - 2 t t, viii 1905 (Kuroiwa}(type series of Odynerus japonicus Matsumura), 1
35
(/" Naha, 29 v 1984 (SKY), 1 (/" same loc., 12 iv 1985 (SKY).
Distribution. Hokkaid6; Honshu; Kyushu; Tsushima Is. (Kami-agata); Naga-shima;
Okinawa Is. (Okinawa-jima). Palearetic.
Taxonomic notes. I have compared the Japanese specimens with one male and one
female from France (sent by Dr. J. Gusenleitner). There are no remarkable difference in
color pattern between them. In the male specimens from Okinawa-jima the yellow
markings on pronotum are slightly larger and the anterior faces of fore and hind coxae are
more often marked with yellow than in those from the Japanese mainlands. However, all
these specimens cannot be divided into distinct forms. It seems that this species is little
variable in basic color pattern throughout its wide geographical range.
Biology. No information is available both in Japan and Europe.
Tokyanus group
Stenodynerus tokyanus (Kostylev)
(Figs. 36, 45, 46, 55-57)
Odynerus (Nannodynerus) tokyanus Kostylev, 1940, Bull. Soc. Nat. Moscou, Sect. BioI. (N.S.), 49 (5/6): 28
( ~ )(type loc.: Tokyo, Honshu).
Stenodynerus tokyanus: Vecht and Fischer, 1972, Hym. Cat. (nov. ed.), 8: 69; Gusenleitner, 1981, Polsk.
Pismo Entomol. 51: 291-294, fig. 67.
Japanese name: Munaguro-chibi-dorabachi.
Diagnosis. Body length (hHh+t1+2): 7.0-8.5 mm in -'f-, 6.5-7.5 mm in t. Fore wing
length: 7.0-8.0 mm in -'f-, 6.0-7.0 mm in t. This species is easily distinguished from the
congeners inhabiting the Japanese mainlands by the larger body, almost complete lack of
propodeal shelf, and well demarcated anterior vertical face of gastral tergite 1. The
anterior face of tergite has only a few scattered punctures and a vertical keel that is
somewhat obscure in t. Male antennal segments 8-11 very thick; segments 12 and 13
(hook) small; hook rather sharply pointed at apex (Fig. 36).
This species has a close relative, S. kusigematii, in the Ryukyus, but none out of Japan.
Gusenleitner (1981) placed S. tokyanus under "Gruppenzugeh6rigkeit nicht geklart". In
Japan two geographical races are recognized.
Nesting biology is not known.
Stenodynerus tokyanus tokyanus (Kostylev)
(Figs. 45, 55)
Stenodynerus tokyanus tokyanus: Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ. (Earth
Sci. Biol.), 15: 122-123, fig. 10.
Japanese name: Munaguro-chibi-dorobachi.
Diagnosis. Female. Black, with the following parts yellow: frontal spot, antennal scape
below (tinged with red), minute spot on temple, a pair of spots or lines on pronotum
anteriorly (often lost), parategula (often ferruginous), a large spot on dorsal
mesepisternum, metanotum (often wholly black), apical bands on gastral tergites 1, 2 and
36
~
53
Figs. 48-57. Color pattern of tegula and parategula in the Japanese Stenodynerus. 48, frauenfeldi; 49,
rufomaculatus; 50-52, chinensis simillimus; 53, clypeapictus; 54, ogasawaraensis; 55, 56, tokyanus
tokyanus; 57, t. flavostcutellatus.
sternite 2, anterior face of fore tibia. Apical 4 or 5 antennal segments below and apical 2/3
of mandible ferruginous. Tegula dark ferruginous. Legs largely blackish brown.
Male. In addition to the yellow markings seen in ~, the following parts yellow:
c1ypeus largely, mesal portion of mandible largely, anterior faces of mid and hind coxae
(sometimes wholly black), anterior face of mid tibia.
Material examined. Honshu: Miyagi-ken -IOf, Rifu, 8 x 1980 (K. Goukon); Niigata-ken - 20f Of, Mikunitege,6 ix 1981 (KE), 20f Of, Senami, 1-11 x 1984 (KE);Ishikawa-ken -IOf, Mt. Haku, Chugu Spa, 17 viii 1973 (1.
Togashi); Fukui-ken -1 Of, Nakajima, Ono, 24 vii 1956 (YH), IOf, Asahi, Ono, 10 viii 1957 (yH), 1 d', same loc.,
20 vi 1963 (yH), IOf, Shimojimmei, Ono, 12 ix 1963 (YH), IOf, Asahi, Ono, 4 viii 1966 (yH).
Shikoku: K8chi-ken -IOf, Engyoji, Kochi-shi, 19 ix 1933 (Y. Sugihara), 1 d', Kechi, 23 v 1935 (K Oike).
Kyushu: Nagasaki-ken - IOf , Haraguchi, Omura, 1 x 1967 (R. Ohgushi); Kagoshima-ken - 1 d', Takachihomine, Kirishima, 31 vii 1972 (K. Kusigemati).
Tsushima Is.: Shimo~gata - IOf , Shira-dake, 22 vii 1978 (K. Tani).
N. Ryukyus: Yaku-shima -1 Of, Kurio - Oko, 5 x 1981 (Y. Takai).
Distribution. Honshu; Shikoku; Kyushu; Tsushima Is. (Shimo-agata); Osumi Is. (Yakushima).
Stenodynerus tokyanus flavoscutellatus Sk. Yamane et Gusenleitner
(Figs. 46,57)
Stenodynerus tokyanus flavoscutellatus Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ.
d')(type loc.: HachijO-jima, Izu Is.).
Japanese name: Hachijo-chibi-dorobachi.
(Earth Sci. Bio!.), 15: 123 (Of
Diagnosis. Female. Black; the following parts yellow or orange yellow: a basal band on
c1ypeus, frontal spot, spot on temple, antennal scape below, a wide band on pronotum
anteriorly, anterior and posterior marking on tegula, parategula, dorsal mesepisternum
largely, a small spot on ventral mesepisternum (often lost), a pair of large spots on
scutellum, metanotum largely, inferior ridge of propodeum, a rather wide apical band on
gastral tergite 1 (the band with an anterior prong on each side), a slightly narrower apical
band on tergite 2, a still narrower band on sternite 2, lines on fore and mid femora
apically, outer faces of all tibiae extensively. Mandible dark ferruginous. 2-3 terminal
37
segments of antenna ferruginous below.
Male. Very similar to the female in coloration. Head more extensively marked with
yellow as follows: clypeus largely, mesal portion of mandible, a short line and a minute
spot along the inner margin of eye below. Scutellum almost wholly black.
ex.
Material examined. Izu Is. (Tokyo-to): Hachijo-jima - ISf, Okagou - Fuji, 26 v 1964
Hirashima & M.
Shiga)(holotype), 1 C!', Kamogawa, 27 v 1964
Hirashima & M. Shiga), ISf, Mitsune - Kantoyama, 30 v
1%4
Hirashima & M. Shiga), 2Sf Sf, Mitsune - Eigou, 24-30 viii 1987 (H. Takahashi).
ex.
ex.
Distribution. Izu Is. (Hachijo-jima).
Stenodynerus kusigematii Sk. Yamane et Gusenleitner
(Figs. 31, 37, 58-61)
Stenodynerus kusigematii Sk. Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ. (Earth Sci.
Bio!.) 15: 123-125 (Sf )(type loc.: Okinawa-jima); Yamane and Tano, 1987, Trans. Shikoku Entomo!' Soc. 18:
329-332, figs. 1-4.
Stenodynerus yambarah Yamane and Gusenleitner, 1982, Rep. Fac. Sci. Kagoshima Univ. (Earth Sci. BiD!.)
15: 125-126 (C!') (type loc.: Okinawa-jima). Syn. nov.
Japanese name: Minami-chibi-dorobachi.
Dingnosis. Body length (h+th+t1 +2): 7.0-7.5 mm in ~, 6.0-6.5 mm in J'. Fore wing
length: 6.0-7.0 mm in ~, 5.5-6.5 mm in J'. Structurally this species is distinguished from
the closely related S. tokyanus in the following points: clypeus nearly as wide as high,
punctation on the vertical face of pronotum much finer and sparser than in tokyanus,
punctures on the posterior horizontal part of pronotum and mesoscutum smaller and
often as large as or smaller than interspaces, abscissa 4 of fore wing much longer than
abscissa 5. In coloration all the subspecies so far known of this species are much more
richly marked with yellow or orange yellow.
This species is at present known only from the Central and Southern Ryukyus,
comprising several subspecies. It is not common on any island. Ishigaki-jima and
Iriomote-jima seem to lack it.
Biology is not known in any subspecies.
Key to the subspecies of Stenodynerus kusigerrultii ( ~ )
1. Gastral tergite 2 laterally with a pair of yellow spots. Ocular sinus extensively yellow.
Yonaguni-jima ................................................................................. subsp. pachymenoides Tano
- Gastral tergite 2 without such spots. Ocular sinus at most with a yellow spot ................. 2
2. Gastral sternites 1 and 2 extensively rufous. Tokuno-shima ....................................... .
.................................................................................................................. subsp. rufiventris Yam.
- Gastral sternites 1 and 2 black. .................................................................................................. 3
3. Clyepus black in lower 2/3. Ocular sinus without yellow spot. Scutellum and propodeum without yellow markings. Amami-Oshima ................................. subsp. tsunekii Tano
- Clypeus at most with a blackish bar in lower portion. Ocular sinus often with a small
yellow spot. Scutellum almost wholly yellow. Propodeum laterally with a pair of large
markings. Okinawa-jima ................................................... subsp. kusigematii Yam. et Gusnlt.
38
Stenodynerus kusigematii tsunekii Tano
(Fig. 58)
Stenodynerus kusigematii tsune1cii: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 329-330, fig.
1 (Sf }(type loc.: Amami-Oshima).
Diagnosis. Female. Black, the following parts yellow or orange yellow: clypeus in
upper 1/3, frontal spot, spot on temple, a line at mandibular base along mesal margin,
antennal scape below, a pair of large markings on pronotum anteriorly, tegula largely,
parategula, a large spot on dorsal mesepisternum, metanotum wholly, relatively wide
band on gastral tergite 1 (laterally dilated), slightly narrower apical band on tergite 2, still
narrower apical band on sternite 2, femora of all legs except in basal parts, tibiae of all
legs. Apical 3 segments of antenna below rufous. Apical half of mandible dark
ferruginous. Tarsi of all legs brownish.
Male unknown.
Materilll examined. C. Ryukyus: Amami~&hima -ISf, 26 vi 1961 (1(. Tsuneki) (holotype).
Distribution. Amami Is. (Amami-Oshima).
Stenodynerus kusigematii rufiventris Sk. Yamane
(Fig. 59)
Stenodynerus kusigematii rufiventns: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 330, fig. 2
( S? )(type loc.: Tokuno-shima, Amami Is).
Diagnosis. Female. Black, with yellow and rufous markings. Yellow are: clypeus (with
a brown stripe in its lower portion), a wedge-shaped marking on frons, spot on temple,
mandible largely (tinged with brown), antennal scape below, a pair of large markings on
pronotum anteriorly, tegula largely, parategula, dorsal mesepisternum wholly, metanotum
nearly wholly, a pair of large markings on propodeum laterally, wide apical band on
gastral tergite 1, slightly narrower apical band on tergite 2, still narrower and sinuated
apical band on sternite 2, anterior face of mid coxa, fore and mid femora, tibiae of all legs.
Several apical segments of antenna below ferruginous. Horizontal portion of tergite 1
except in the middle, sternites 1 and 2 rufous. Trochanters and tarsi of all legs brownish.
Male unknown.
Materilll examined. C. Ryukyus: Tokuno-shima - 1S?, Kametsu, 13 vii 1983 (T. Moriyama) (holotype).
Distribution. Amami Is. (Tokuno-shima).
Stenodynerus kusigematii subsp. ?
Stenodynerus kusigematii subsp.: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 330.
Diagnosis. Male. Black; the following parts orange yellow or yellow: clypeus wholly
(yellow), mandible (yellow) except for apex (ferruginous) and basal marking (black),
antenna I scape except for apical 1/3 of upper face black, frontal spot, small spot on
temple, wide anterior band on pronotum (medially narrowed), dorsal mesepisternum
largely, tegula, parategula, a transverse marking on scutellum, metanotum wholly, very
39
Figs. 58-61. Body color pattern in the four subspecies of Stenodynerus kusigematii (after Yamane &
Tano, 1987). 58, ssp. tsunekii (Amami-oshima); 59, ssp. rufiventris (Tokuno-shima); 60, ssp.
kusigematii (Okinawa-jim a); 61, ssp. pachymenoides (Yonaguni-jima).
wide apical band on gastral tergite 1, apical bands on tergite 2 and sternite 2, fore and mid
legs except for coxae, apical 1/3 of hind femur, hind tibia and tarsus wholly. Terminal
segments of antenna below ferruginous.
Female unknown.
Material examined. C. Ryukyus: Okinoerabu-jima - 1 C!', Oyama, 21 iii 1964 (T. !ida).
Stenodynerus kusigematii kusigematii Sk. Yamane et Gusenleitner
(Fig. 60)
Stenodynerus yambarah Yamane and Gusenleitner, 1982 [see under S. kusigematiil.
Japanese name: Minami-chibi-dorobachi.
Diagnosis. Female. Black, with the following parts yellow or lemon yellow: clypeus
(sometimes with a transverse black marking), wedge-shaped frontal marking, a relatively
large spot on temple, a minute spot on ocular sinus (often lost), mandible except for apical
part (brownish) and basal triangular marking (black), antennal scape except for apical 2/3
of its upper face, wide pronotal band medially narrowed, tegula largely, parategula,
dorsal mesepisternum wholly, scutellum and metanotum almost wholly, a pair of large
markings on propodeum laterally, wide apical band on tergite 1 (laterally much dilated),
apical band on tergite 2 (laterally slightly dilated), an incomplete apical band on tergite 3,
sternite 1 except medially, narrow apical band on sternite 2 (medially widened), an
incomplete apical band on sternite 3, anterior face of mid coxa, fore and mid femora, tibiae
of all legs. Apical 3 segments of antenna below dark orange. Tarsi of all legs blackish.
Male. The original description of the male of S. yambarah is reproduced below. Black,
with the following parts yellow: clypeus wholly, mandible largely, frontal marking (in one
specimen, produced below), minute spot on temple, antennal scape below, wide band on
40
pronotum anteriorly, a marking on dorsal mesepisternum, tegula posteriorly, parategula,
metanotum wholly, apical bands on gastral tergites I, 2 and sternite 2, incomplete band on
sternite 3 (in one specimen also on 4), anterior face of mid coxa, tibiae of all legs.
Material examined. C. Ryukyus: Okinawa-jima - 2<! <!, Hedo, Kunigarni, 5 iv 1979 (K. Ohara)(including
the holotype of S. yambarah), 1 ~ , Benoki, Kunigami, 7 iv 1979 (K. Kusigemati) (holotype), 1 ~, Oku,
Kunigarni,5 viii 1982 (YH), 1 ~, Hentona, 1 vi 1983 (AN), 1 ~, Yona, 2 vi 1983 (AN), 1 ~, Hentona, 5 x 1987
(AN).
Distribution. Okinawa Is. (northern parts of Okinawa-jima).
5tenodynerus kusigematii pachymenoides Tano
(Fig.61)
Stenodynerus kusigematii pachymenoides: Yamane and Tano, 1987, Trans. Shikoku Entomo!' Soc. 18: 330332, fig. 4 (~ <!) (type loc.: Yonaguni-jima, Yaeyama Is.).
Diagnosis. Female. In structure similar to the nominotypical subspecies, but the
anterior vertical face of pronotum virtually with no punctures except for the pair of pits
near neck.
Black (ground color of gastral tergites tinged with brown), with the following parts
vivid yellow: dypeus wholly, mandible largely, frontal marking produced so as to reach
upper margin of dypeus, ocular sinus, a long stripe behind eye, antennal scape below,
wide pronotal band medially narrowed, dorsal mesepisternum largely, a small spot or
spots on ventral mesepisternum, tegula except for center and margins, parategula,
scutellum and metanotum nearly wholly, a pair of large markings on propodeum
dorsolaterally, apical bands on gastral tergites 1-4 (that on tergite 1 with an anterior prong
on each side), a pair of spots on tergite 2, sinuated apical bands on sternites 2 and 3 (4), a
pair of spots on sternite 2, anterior face of fore and mid coxae, apical 2/3 of fore and mid
femora, tibiae of all legs (partly brownish). Antenna ferruginous below.
Male. Very similar to the female. The two specimens examined lack the small yellow
spot(s} on ventral mesepisternum, and yellow spots on sternite 2.
Material examined. S. Ryukyus: Yonaguni-jima - 2 <! <!1 ~, Kubura, 24 vii 1984 (c. Nozaka), 2~ ~ ,
Urabu, 10 viii 1985 (TM) (including holotype).
Distribution. Yaeyama Is. (Yonaguni-jima).
Genus Allodynerus Bliithgen
Al/odynerus BHithgen, 1938, Konowia 16 (1937): 280 (as subgenus of "Euodynerus B1iithgen")(type
species: Odynerus floricola Saussure, 1853, original designation); 1938, Dt. Ent. Zeitschr. 1938: 452, 458 (as
genus); 1953, Zoo!. Anz. 150: 50 (as genus); 1961, Faltenwespen Mitteleuropas, p. 69 (as genus, in key).
Japanese name: Kita-dorobachi Zoku.
The wasps of this genus are medium-sized, measuring 8-12 mm in total length. Head
and thorax bears golden pubescence. Clypeus is wider than high. The depression for
cephalic fovea (~ ) is large, kidney-shaped and posteriorly with a single fovea. Antennal
hook ( 6') is developed, reaching the apex or middle of segment 10 (Figs. 64, 65). Thorax is
relatively short; whole mesoscutum and scutellum are punctate. Tegula is narrow and
41
long, with its posterior lobe well developed, extending beyond the posterior end of
parategula (Figs. 66, 67). Posterior face of hind coxa posseses a keel which is obtuse in
profile. Propodeum has no shelf; superior carina of propodeum is only weakly developed
or almost absent. Gastral segment 1 is narrower than 2, tergite 1 being wider than long
and without transverse carina.
Only two species occur in Japan (Giordani Soika, 1986; Yamane & Tano, 1987), while
seven other species have been known from the Palearctic region (Vecht & Fischer, 1972).
Allodynerus delphinalis delphinalis (Giraud)
(Figs. 62-64, 66, 68-70, 72)
Odynerus (Leionotus) delphinRlis Giraud, 1866, Ann. Soc. Ent. Fr. (4)6: 464 (!f )(type loc.: Grenoble,
France).
Allodynerus delphina/is delphinalis: Bliithgen, 1953, Zool. Anz. 150: 53-54; 1961, Faltenwespen
Mitteleuropas, pp. 126-132; Yamane & Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 342-343, figs. 10, 11.
Allodynerus sp.: Yamane, 1982, Trans. Essa Entomol. Soc. 53: 5; Hisamatsu et al. 1986, Bull. Fac. Educ.
Ibaraki Univ. (Nat. Sci.) 35: 59.{j(), figs. 17,29,48,60, SO.
Japanese names: Atoboshi-kita-dorobachi (Kita-dorobachi).
Diagnosis. Female. Body length (h+th+t1+2): 7.0-10.0 mm. Fore wing length: 7.5-9.0
mm. Head distinctly wider than high (W /H = 1.17), densely punctate. Hairs on head
distinctly longer than those on thorax. Clypeus much wider than high, triangular,
anteriorly weakly emarginate (Fig. 62). Interantennal keel distinct. Ocellar triangle flat;
distance between posterior ocelli only slightly longer than ocello-ocular distance.
Depression for cephalic fovea large, wider than distance between posterior ocelli, with a
transverse carina just behind the fovea. Hairs on antennal scape microscopic. Thorax
densely punctate; anteroventral portion of mesepisternum partly impunctate; upper
metapleuron striate. Superior ridge of propodeum weak, but distinct edge present;
propodeal groove with a vertical carina; sides of propodeum only weakly reticulate in
upper portion. Gastral segments 3-6 much narrower than segment 2. Gastral tergites 1-3
densely punctate; punctation stronger on tergite 1 than on 2; apical margin of tergite 2
rarely slightly reflexed, often weakly denticulate; preapical depression on tergite 2 weak
(Fig. 68). Gastral sternite 1 reticulate; basal area of sternite 2 with ca. 15 vertical carinae;
punctation on this sternite similar to that on tergite 2; last tergite and sternite almost
impunctate.
Black. Yellow are: clypeus with a median black marking, a transverse frontal
marking, a stripe on inner orbit just above clypeus, a stripe behind eye, basal half of
mandible largely, scape below, an anterior pronotal band, a spot on dorsal mesepistemum,
an irregular marking on scutellum (often lost), metanotum wholly, tegula with a central
semi-transparent part, a large marking on each side of dorsal face of propodeum, an apical
bands on gastral tergites 1-5 (band on tergite 2 widest), apical spot on tergite 6, an apical
band on stemite 2, apical fourth to third of femora of all legs, tibiae and tarsi of all legs
nearly wholly (slightly tinged with brown). Antennal flagellum below ferruginous.
Male. Body length (h+th+t1+2): 6.0-7.5 mm. Fore wing length: 6.5-7.5 mm. Very
similar to the female in punctation and sculpture. Clypeus more deeply emarginate
anteriorly (Fig. 63). Distance between posterior ocelli distinctly longer than ocello-ocular
distance. Inner faces of antennal segments 10 and 11 with a concavity to receive recurved
42
segments 12 and 13. Marked with yellow more extensively than in the female. Oypeus
and mandible almost wholly yellow. Yellow stripe on inner orbit longer, extending into
ocular sinus. Mid coxa wholly yellow. Mesopleuron, metanotum, propodeum and last
tergite, however, almost always lacking yellow markings.
Material examined. Hokkaido: 1 d', Abashiri, 2 ix 1954 (K. Kamijo).
Honshu: lwate-ken - 1 d', Kuriyagawa, Morioka, 23 vi 1%9 (yM), 1 d', same loc., 24 vi 1973 (yM), 2 d' d',
Oshuku, Shizukuishi, 7 viii 1987 (SKY); Fukushima-ken - 1 !?-, Bantai-san, Aizu, 4 viii 1927 (S. Matsumura);
Niigata-ken - 2!?- !?-, Shidai-hama, 9 vii 1970 (HI), 1!?-, Nagaoka, 9 vii 1976 (SKY); Saitama-ken - 1 d',
Kodama, 25 vii 1964 (TN), 1!?-, same loc., 10 viii 1%9 (TN); Gumma-ken - 1 d', Buson-sanroku, 21 vii 1974
(HI); lbaraki-ken - 1 d'1!?-, Tsuchiura, 29 viii 1974 (SKY); Nagano-ken -1 d', Minamiminowa, Ina, 22 v 1962 (H.
Fujii), 1 d', same loc., 29 vii 1%2 (yM), 2d' d', Saku, 28 vii 1974 (YM); Fukui-ken - 1 d', Ono, 3 x 1964 (yH);
Ishikawa-ken - 1 d', Komaiko, 4 vii 1976 (H. Kurokawa); Shimane-ken -1 d', Nishikawatsu, Matsue, 15 viii 1985
(M. Goubara).
Distribution. Hokkaido; Honshu. Palearctic.
Taxonomic notes. This species is easily distinguished from all the other species of the
Japanese Eumenidae by the possession of both the black oval marking on dypeus and the
yellow round marking on tergite 6 in the female (Fig. 69). Although Bliithgen (1961) points
out that body markings in European populations greatly vary among specimens, the color
64
63
r;--1
65
Figs. 62-68. Morphological characters in the Japanese Allodynerus. 62-64, 66, 68, A. delphinalis; 65,
67, A. mandschuricus. 62, c1ypeus (!?- ); 63, ditto (d'); 64, 65, terminal segments of male antenna
(A, profile; B, segment 10 from below); 66, 67, tegula and parategula; 68, apical part of gastral
tergite 2 (!?-).
43
Figs. 69-71. Color pattern of the two Japanese species of Allodynerus (after Yamane & rano, 1987).
69, A. delphinaIis; 70, melanic individual of delphinalis; 71, A. mandschuricus.
pattern in the Japanese population is generally stable. However, in a rather melanic female
at hand the black clypeal marking is very large, widely connected with black basal and
apical margins (Fig. 70).
Biology. The nesting biology in Europe was studied by Enslin (1922). The following
description of the nesting behavior of this species is based upon unpublished data
amassed by Dr. K. Goukon at Rifu, Miyagi-ken, northern Honshu, during June/July, 1987.
This species nests in dead stems of Miscanthus sinensis growing in open lands. A few nests
were also found from stems of a bramble, Rubus palmatus var. coptophyllus, the upper
portion of which had been cut away in order to induce the small carpenter bee Ceratina
fflavipes to nest. The female wasps most probably take over abandoned nests of C. [lavipes,
but they may excavate by themselves the pith to enlarge the cell. The total number of
brood cells constructed in each tube nest varied from 1 to 5 (mean, 2.7; n=12). The first
(innermost) cells always contained a female young. A vestibular (empty) cell was
constructed in 7 of the 12 completed nests observed. Although the prey were not
identified even at family level, larvae of some microlepidopterous species were no doubt
involved. It is estimated that 4-8 caterpillars were provisioned for a male-producing cell
and 9-15 for a female-producing cell. In two nests the mother wasp was found in her
incompleted brood cell together with her medium-sized larva. This suggests the
occurrence of progressive provisioning in this species. At least two generations were
recognized in 1987.
Parasitoids: Melittobia sp. (Hymenoptera, Eulophidae) and two species of parasitic
flies.
44
Allodynerus mandschuricus Bliithgen
(Figs. 65, 67, 71, 72)
Allodynerus mandschuricus Bliithgen, 1953, Zoo!. Anz. 150 (5f) (type loc.: "Charbin", China): 57-58;
Giordani-Soika, 1970, Boll. Soc. Entomol. Ital. 102: 150-151 (d'); 1986, Boll. Mus. Civ. St. Nat. Venez. 35 (1984):
132; Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 343-344.
Japanese name: Ajia-kita-dorobachi.
Diagnosis. Femele. Body length (h+th+t1 +2): 8.5-9.0 mm. Fore wing length: 7.0-7.5
mm. This species differs from A. delphinalis in the following points. Head more round,
only slightly wider than high (W /H = 1.08). Clypeus slightly longer than in delphinalis.
Distance between the posterior ocelli longer than that between posterior ocellus and eye.
Hairs on antennal scape relatively long. Sides of propodeum distinctly reticulate in upper
portion; superior ridge absent. Punctation on gastral tergites 1 and 2 stronger than in
delphinalis. Gastral tergite 2 slightly reflexed but with smooth margin apically; preapical
• A. delphinalis
.. A. mandschur i cus
,.0
72
Fig. 72. Distribution of Allodynerus species in Japan.
45
depression distinct.
Black, marked with yellow as follows: two to four spots on clypeus, a very small
frontal spot, a short line along inner margin of eye below, a stripe on mandibular base, a
short basal line on antennal scape below, a small spot behind each eye, a pair of small
triangular markings anteriorly on pronotum, tegula except for margins and basal brown
part, a transverse marking on metanotum (sometimes lost), apical part of propodeum in
both sides, a medially narrowed apical band on gastral tergite I, a regular apical band on
tergite 2, a laterally dilated apical band on gastral sternite 2. Antennal flagellum below
slightly tinged with red. Legs largely blackish; anterior face of fore tibia orange yellow;
tarsi of all legs brownish.
Male. Body length (h+th+t1+2): ca. 7.5 mm. Fore wing length: ca. 6.5 mm. Clypeus
more deeply emarginate than in the female; but the emargination shallower than in the
male of A. delphinalis; punctation on clypeus much stronger than in delphinalis. Antennal
hook shorter than in A. delphinalis; its apex hardly reaching the apex of segment 10 which
is only weakly concave at apex (Fig. 65).
Black, with the following parts yellow: clypeus nearly wholly, labrum, mandible
except for periphery, stripe on antennal scape below, stripe on temple, short band on
pronotum anteriorly (medially interrupted), a pair of small markings on scutellum, a very
small marking on propodeum posteriorly, narrow bands on tergites 1 and 2, and sternite 2,
a small spot on anterior face of mid coxa, markings on anterior faces of fore and mid
tibiae. Antenna! flagellum wholly blackish. Tarsi of all legs blackish brown.
Material examined. Honshu: Fukui-/cen - 20f Of, Mt. Tokin, 23-24 vii 1974 (TM); Saitama-ken - lOf, Ogano,
7 vi 1970 (TN); Hyogo-ken - IOf, Sasayama, Tamba, 25 v 1952 (K. Nohara), lOf, same loc., 1 x 1961 (K.
Okamoto), IOf, same loc., 18 vi 1962 (K. Okamoto), IOf, same loc., 29 ix 1965 (K. Okamoto), IOf, Miki, 9 viii
1967 (K. Iwata), 1 d'IOf, Takasago, 1987 reared from a nest (Y. Nakatani); Shimane-ken - IOf, Kawatsu,
Matsue, 17 viii 1985 (N. Sugiura).
Distribution. Central and western Honshu. Korea (Giordani Soika, 1970); Manchuria.
Taxonomic notes. This rare species is in structure very similar to the preceding species.
The presence of relatively long hairs on antennal scape (in at least basal part) may be the
most reliable characteristic (in delphinalis microscopically pubescent). In coloration,
however, this species is very distinctive by its predominantly melanic body. Yellow
markings are much reduced: scutellum and mesopleuron never marked with yellow in the
female; gastral tergites 3-6 wholly black; legs largely blackish.
Biology. This species nests in bamboo tubes small in inner diameter (Nakatani, 1988).
One female specimen from Sasayama, Hyogo-ken, was placed under the identification
label "Stenodynerus frauenfeldi", and the female from Miki, Hyogo-ken, under "Ancistrocerus
japonicus", both in the collection of the Entomological Laboratory, Kobe University. These
arrangements may have been made by Dr. K. Iwata, but it seems that his published
observations on the biology of s. frauenfeldi and A. japonicus do not include Allodynerus
mandschuricus.
Genus Euodynerus Dalla Torre
Euodynerus Dalla Torre, 1904, Gen. Ins. 19: 38 (name for Odynerus, subgenus Leionotus, me Division,
Sect. II, Saussure, 1853, Et. Fam. Vesp. 1: 177)(type species: Vespa danlici Rossi, 1790); Vecht & Fischer, 1972,
Hym. Cat. (n. 00.) 8: 87 (as genus); Giordani Soika, 1978, Soc. Venez. Sc. Nat.-Lavori-, 3: 40 (as genus, in key).
Japanese name: Mikado-dorobachi Zoku.
46
Medium-sized wasps. Clypeus is variable in shape, apically truncated or shallowly
emarginate (Figs. 85-88). Anterior vertical face of pronotum has no transverse striae but
punctures laterally. Transverse pronotal carina is complete. Posterior pro notal lobe and
pretegular carina are well developed. Tegula is posteriorly produced as a triangular
process; posterior end of parategula does not extend beyond the posterior end of tegula.
Mesoscutum and scutellum are extensively punctate, without prescutal furrows.
Metanotum is not bidentate. Propodeum often bears upper spines just behind metanotum
(Fig. 78). Gastral segment 1 is, seen from above, wider than long, usually more than 4/5 as
wide as segment 2. Tergite 1 is lacking in both longitudinal groove and transverse carina,
posteriorly not swollen, and with an apical semitransparent Oamellate) area. According to
Bliithgen (1938b), the Palearctic species of this genus have the gastral tergite 2 not
provided with such an apical semitransparent part, but Euod. nipanicus is exceptional in
this respect (Figs. 98-100).
This genus comprises ca. 60 species in five subgenera in the Palearctic region (Vecht &
Fischer, 1972). In Japan four species have been recorded (Yamane & Tano, 1987). Giordani
Soika (1986) added one species, Euod. bicingulatus, from Hokkaido and Tokyo (?), but his
description best agrees with smaller specimens of male Anterhynchium flavomarginatum
micado as will be discussed below.
Key to the Japanese species of the genus Euodynerus
1. Vertex with distinct tubercles in ocellar region (one situated just behind anterior ocellus
and apically bifid; the others inside posterior ocelli). Posterolateral angles of propodeum
inconspicuous, not forming spines ........................................................... Euod. trilobus (Fab.)
- Vertex without tubercles in ocellar region. Posterolateral angles of propodeum projecting as upper spines behind metanotum ............................................................................ 2
2. Clypeus higher than wide, with many vertical carinae and a few punctures, apically
truncate ( ~ ). Posterior vertical face of metanotum shining below, coarsely denticulate in
uppermost part. Propodeum as seen from above only slightly narrowed toward
posterior margin. Metanotum wholly black. .......................................... Euod. dantici (Rossi)
- Clypeus as wide as or slightly wider than long, with relatively dense punctures over the
whole disc, without vertical carinae, distinctly emarginate at apex (~). Posterior vertical
face of metanotum not very shining below, finely denticulate in uppermost part.
Propodeum as seen from above distinctly narrowed toward posterior margin. Metanotum marked with yellow...................................................................................................... 3
3. Gastral sternite 2 basally with a longitudinal groove. Gastral tergite 2 usually with an
apical semitransparent part which is more conspicuous in d'. In profile head and
meso scutum bearing short, erect and uniform hairs as in a brush; hairs on head longer
than those on thorax. Clypeus as long as or slightly higher than wide (~ d'). Gaster
without long hairs ............................................................................... Euod. nipanicus (Schult.)
- Gastral sternite 2 basally without distinct groove. Gastral tergite 2 without apical
lamella. In profile head and mesoscutum bearing longer, disheveled hairs that are
uniform in length. Clypeus wider than high (~ d'). Gaster partly with long hairs .
............................................................................................................. Euod. quadrifasciatus (Fab.)
47
Euodynerus (Euodynerus) dantici (Rossi)
(Figs. 76, 79, 80, 88,90, 101, 102, 108, 110B)
Vesru dantici Rossi, 1790, Fauna Etrusca,2: 89, pI. 16, fig. 6 (Sf )(type loc.: Italia).
Euodynerus dantici: Vecht and Fischer, 1972, Hym. Cat. (n. ed.), 8: 88-89.
Japanese name: Kabaobi-dorobachi.
Diagnosis. Body length (h+th+t1+2): 10.5-11.5 mm in ~,and 7.5-10.0 mm in 6'. Fore
wing length: 9.5-12.0 mm in ~,and 8.5-10.0 mm in 6'. Structure as in the key.
This species was originally described from Italy, and is widely distributed from
Europe to eastern Asia. Anywhere in Japan it is less common than Euod. nipanicus, and
inhabits a smaller number of islands than does the latter. The northern part of the
Ryukyus (Osumi Is.; Mi-shima Is.; Tokara Is.) and possibly Sado-ga-shima, Tsushima Is.,
Shikoku and Amami-6shima lack the former (Figs. 108, 110B). In the Central and Southern
Ryukyus, this species is more common than in Japan proper, and small islands such as
Uke-shima (Amami Is.) and Hateruma-jima (Yaeyama Is.) are often inhabited by it (Fig.
110B). In this region the species is variable in color pattern, but the variation is transitional
from islet to islet so that the species cannot be divided into distinct geographical races
J'-"-
po
~-
75
~
b~
\
o-
__ \
84
\'
82
78
'9~"""':'::""
cor"""""""'';!.'
~I
B //
~,
Figs. 73-84. Morphological characters in the Japanese Euodynerus. 73, 74, Pubescence of head and
thorax in nipanicus (73) and 'fUildrifasciatus (74); 75, ocellar region in trilobus; 76, 77, terminal
segments of male antenna in dantici (76) and nipanicus (77A ,B); 78-82, metanotum and
propodeum (78, SO, posterior view; 79, 81, 82, profile) in nirunicus (78,79), dantici (80, 81) and
trilobus (82) (after Yamane, 1979); 83, 84, mid tarsus (6') of no/atus (European species) (83) and
nirunicus (84) (after Giordani Soika, 1986).
48
A
B
85
86- -____ B
___
B
87
B
Figs. 85-88. Clypeus of the four Japanese species of Euodynerus. 85, nipanicus; 86, quadrifasciataus;
trl, trilobus; 88, dantid. A, 6'; B, -\C.
t; 93fi94 i:95~96
'.
;,;....
-"\..
~.........
"',-
.... _ ' ....
""
Jt;-:-
100
Figs. 89-}()(J. Gastral characters in the Japanese Euodynerus (after Yamane, 1979). 89, gastral
stemite 2 of nipanicus, showing the basal furrow; 90-%, apical part of gastral tergite 2 (profile)
in dantid (90), quadrifasdatus (91), nipanicus flavicornis (92, 93), n. nipanicus from Honshu Kyushu (94, 95), n. nipanicus from Hokkaido (%); 97-100, first two tergites (from above) in
quadrifasciatus (97), nipanicus flavicornis (98), n. nipanicus from Honshu - Kyiishu (99), n.
nipanicus from Hokkaido (100).
49
(Yamane & Tano, 1987). I have recognized two principal subspecies in Japan, i.e., Euod. d.
violaceipennis (Japan proper) and Euod. d. nigrescens (c. and S. Ryukyus) (Figs. 101, 102).
The subspecies of eastern Asia were discussed at length by Giordani Soika (1986). This
species is widely distributed in the Palearctic region, and comprises more than ten
subspecies.
Key to the subspecies of Euod. dantici occurring in eastern Asia
1. Mesopleuron usually without yellow spot. Northern part of the Far East. ................. .
............................................................................................................. subsp. vio/aceipennis (G.S.)
- Mesopleuron always with a yellow spot under wing base ................................................... 2
2. Gastral tergites 1-3 in .'f. (also 4 in J') each with a yellow or orange yellow apical band.
Transbaikal. ..................................................................................... subsp. brachytomus (Kost.).
- Gastral tergites 1-4 in .'f. (also 5 in J') each with a yellow apical band. C. and S.
Ryukyus, Taiwan ............................................................................. subsp. nigrescens (Gusnlt.)
Euodynerus dantici violaceipennis Giordani Soika
(Figs. 101, 108)
Euodynerus dantici violaceipennis Giordani Soika, 1973, Boll. Mus. Civ. Stor. Nat. Venez. 24: 114 (S? t)
(type loc.: "Canton", China).
Odynerus dantici: Yasumatsu, 1950, Icon. Ins. Jpn. 2nd ed. p. 1455, fig. 4199; Ishikawa, 1965, Icon. Ins.
Jpn. Col. Nat. Ed. 3: 297, pI. 149, fig. 4.
Euodynerus dantici brachytomus: Vecht and Fischer, 1972, Hym. Cat. (n. 00.) 8: 88; Yamane, 1979, New
Entomol. 28: 8-9, figs. 3, 6, 9, 11; Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 333-334.
Japanese name: Kabaobi-dorobachi.
Diagnosis. Female. Black, marked with yellow or orange yellow as follows: clypeus
above (the mark with a triangular incision apically in the middle), a frontal spot extending
to the upper margin of dypeus, a narrow line in ocular sinus below, a stripe behind each
eye, a small basal spot on mandible, antennal scape below, a medially narrowed. anterior
band on pronotum, tegula with margins and center brown, parategula, a pair of spots on
scutellum (often lost), a very small spot under wing base (usually lost), a small spot on
posterolateral face of propodeum (sometimes lost), dorsal and lateral parts of gastral
tergite 1 except for median basal black area, a medially dilated apical band on tergite 2
(this tergite sometimes with lateral yellow markings which are often fused with the wide
apical band), a narrow apical band on tergite 3 (sometimes also on 4), posterolateral
corners of sternite 2, all legs except for coxae and the major part of femora. Mandible
ferruginous.
Male. Body marking paler than in the female, especially on head and thorax. Markings on sinus of eye and mandible slightly larger than in the female. Tergite 4 always with
an apical band. Posterolateral corners of sternite 3 also marked with yellow. Legs
extensively yellow.
Material examined. Honshu: lwate-lcen - 1 S?, Oshuku, Shizukuishi, 7 viii 1987 (SKY); Miyagi-lcen - 1 S?,
Okunikkawa, 19 viii 1974 (K. Goukon), 2d' d', Rifu, 6-13 vii 1980 (K. Goukon); Nagano-lcen -IS?, Ina, 14 ix
1961 (YM); Kyoto-fu -1 S?, Kurama, 6 viii 1980 (K. Goukon); Hyogo-ken - l t l S?, Sasayama, Tamba, 31 vii 1965
50
(K. Iwata); Osaka-fu-l-'i!-, Ikeda, 4 vii 1947 (K Iwata); Shimane-ken -l-'i!-, Kawatsu, Matsue, 12 vii 1986 (N.
Sugiura); 1 d', Matsue, 19 vii 1988 (SKY).
Kyushu: Fukuoka-ken -1 d', Karumachi, 5 vii 1958 (YM); Kagoshima-ken -1-'i!-. Kushikino, 14 viii 1978 (H.
Nagase);2d' d'l-'i!-, Taguchi, Kirishima,3 vii 1981 (SKY), l-'i!-, Kagoshima-shi, 7 viii 1981 (SKY).
Distribution. HonshU; Awaji-shima; KyushU. Korea; China.
Biology. Iwata (1938b) observed the nesting biology of this form in Honshu. The
female wasp nests in reed tubes used for mats and blinds, and prefers slender tubes (6-7
mm in inner diam.). The number of cells constructed per tube nest ranged from 1 to 9
(n=15, m=4). In every nest an empty cell was seen at the entrance. Nineteen to 50
caterpillars (Tortricidae ?) were stored in larval cells. Judging from the collection data this
form may be univoltine.
Euodynerus dantici nigrescens Gusenleitner
(Figs. 102, 110B)
Euodynerus dantici nigrescens Gusenleitner, 1979, Nachrichtbl. Bayer. Entomol. 28: 62 (-'i!- d')(type loc.:
Shihmen, N. Taiwan); Tano et aI. 1985, Hym. Comm. 23: 31; Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat.
Venez. 35: 133-134.
Euodynerus dantici Tlio/aceipennis: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 334-335.
Japanese name: Minami-kabaobi-dorobachi.
Diagnosis. Female. Black, more richly marked with yellow or orange yellow than in
the preceding subspecies as follows: c1ypeus wholly (rarely with a black apical mark),
basal marking on mandible, frontal marking extending to the upper margin of c1ypeus,
Figs. 101, 102. Body color pattern of Euodynerus dantici Tlio/aceipennis (101) and Euod. d. nigrescens
(H12).
51
ocular sinus largely, antennal scape below, relatively long stripe behind each eye, wide
pronotal band medially narrowed, spot under wing base (always present), tegula except
for margins and center, parategula, a pair of large spots on scutellum (never lost),
dorsolateral face of propodeum almost wholly. Gastral color pattern as in the subspecies
violaceipennis, but lateral projections of yellow apical band on tergite 2 always present,
tergite 4 almost aJways with a yellow apical band, and sternite 3 marked with yellow
posterolaterally. Coloration of legs as in the subspecies violaceipennis, but hind coxa
sometimes yellow anteriorly. Mandible sometimes extensively yellowish.
Male. Very similar in coloration to the female, but body markings much paler and
sometimes sulpher yellow. Mandible and legs more extensively, and dypeus always wholly yellow. Tergite 5 with apical band, and sternite 4 marked with yellow posterolaterally.
I have examined a paratype female of this form from Taiwan (sent by Dr. J.
Gusenleitner) and many males and females from Taiwan in my collection. No important
differnce has been found between the populations of the Ryukyus and Taiwan.
Mtiterial examined. C. Ryukyus: Uke-shima - 4 t J', 12 viii 1987 (M. Tatsuno); TokunCH;hilnll - 1 S?, Katoku,
7 ix 1983 (T. Moriyama), 1 S?, Kametoku, 13 vii 1984 (SKY); Okinoerabu-jima - 2J' J', Tokutoki, 4 viii 1972
(TN), 1 S?, China, 19 v 1981 (Kukidome &: Koya); 1 S?, Shinjo - Tamina, 14 vii 1981 (Y. Takai), 1 t2S? S?,
Wadomari, 15 vii 1984 (SKY); Yoron-tii - 1 J', 4 vi 1985 (SKY), 2 J' J', Chabana, 6 viii 1972 (TN); Okinawa-jima 1 S?, Naha, 9 viii 1972 (TN), 1 S?, Tanodake, 26 viii 1979 (H. Nagase), 1 S?, Hentona, 4 vii 1981 (AN), 1 J', same
loc.,3 vi 1983 (AN), 1 J', same loc., '1:1 vii 1987 (SKY); lzena-jima -1 t, Nakada, 25 vii 1%7 (T. Kifune); Kourijima -1 S?, 18 x 1988 (Y. Kusui); Miyagi-jima-l J', 31 vii 1971 (T. Kifune).
S. Ryukyus: Miyako-jima - 3 J' J', Sunayama, 2-4 viii 1984 (CN); lshigaki-jima - 1 J', Kabira, 4 vi 1983
(YH), 1 J', Kabira, 29 vii 1984 (CN), 1 J', Shinkawa, 19 vi 1986 (51), 1 J', Kabira, 21 vi 1986 (51), 7 J' J', Omotodake, 8 vii 1988 (K. Nakamine); Taketomi-jima - 2 J'1 S?, 8 vii 1988 (K. Nakamine); Hateruma-jima - 14 J' J',
30 vi -1 vii 1988 (SKY).
t
Distribution. Amami Is. (Uke-shima; Tokuno-shima; Okinoerabu-jima; Yoron-t6);
Okinawa Is. (Izena-jima; Okinawa-jima; Kouri-jima; Yagaji-jima; Miyagi-jima; Kume-jima);
Miyako Is. (Miyako-jima); Yaeyama Is. (lshigaki-jima; Taketomi-jima; Hateruma-jima).
Taiwan.
Euodynerus (Pareuodynerus) nipanicus (Schulthess)
(Figs. 73, 77, 78,80,84,85,89,92-100,103-106,107, 110A)
lionotus tomentosus var. nipanicus Schulthess, 1908, Mitt. Schweiz. Entomol. Ges. 11: 287-288 (t)(type
loc.: Japan).
Euodynerus (Pareuodynerus) nofJztus : Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 335-336.
Euodynerus (Pareuodynerus) notatus notatus: Vecht and Fischer, 1972, Hym. Cat. (n. 00.), 8: 95-96 (from
Japan).
Euodynerus (Pareuodynerus) nipanicus: Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 135-138.
Japanese name: Mikado-dorobachi.
Diagnosis. Body length (h+th+tl+2): 7.5-11.0 mm in ~, 6.5-10.5 mm in t. Fore wing
length: 9.5-10.5 mm in ~, 7.5-9.0 mm in t. Structure as in the key.
This species (nipanicus) has been treated by Japanese authors (e.g., Matsumura, 1911;
Yasumatsu, 1952) as Odynerus micado Kirsch or o. quadrifasciatus (Fabricius)(both agreeing
in the Japanese name applied, Mikado-dorobachi). But, as pointed out by Yamane (1979)
the form here is no doubt a dose relative of Euod. notatus, and Yamane and Tano (1987)
considered it to be a Japanese subspecies of the latter, which is Palearctic in distribution.
52
On the other hand, Giordani Soika (1986) separated it specifically from notatus, and
applied the name nipanicus to it. He enumerated the following differences: (1) in nipanicus
the punctation, especially on gaster, stronger and denser, (2) gastral tergite 2 more or less
strongly reflexed at apex to form a lamella, which is more developed in the male than in
the female, (3) sternite 2 basally more convexed (these three are useful to distinguish this
species from notatus and quadrifasciatus), (4) tenninal segment of mid tarsus of the male
much more slender than in notatus (Figs. 83, 84), (5) in both sexes, temple less developed
and seen from above shorter than the eye width, (6) digitus of male genitalia narrower,
elongate, more widely round at apex than in notatus. Giordani Soika's view that the form
is a distinct species is here adopted.
The species may widely occur in eastern Asia except in the tropics. At least some of
the previous records of Euod. notatus from the Far East may be referable to nipanicus
(Giordani Soika, 1986).
My comparison of nipanicus from the Japanese mainlands with "Odynerus flavolineatus"
from the Ryukyus (Matsumura, 1911) has revealed that these are geographical races of the
same species. I recognize three subspecies in Japan, one from the mainlands and the other
two from the Ryukyus. The Hokkaid6 population of nipanicus nipanicus, referred to as
Euod. notatus pubescens by Yamane and Tano (1987), slightly differs in structure and color
pattern from the form inhabiting Honshu, Shikoku and Kyushu. However, in this paper
both the forms are tentatively treated under the same subspecies, because the subspecies
name to be applied to the Hokkaido form may be fixed only after comparing the Japanese
forms with those from Sakhalin and eastern Siberia.
Key to the Japanese subspecies of Euodynerus nipanicus
1. Clypeus usually wholly black, rarely yellow at base (~). Scutellum often lacking
yellow markings (~ 6'). Dorsolateral face of propodeum and gastral sternite 2 rarely
with large spots ( ~ 6'). Legs largely black or brownish ............. subsp. nipanicus (Schult.)
- Clypeus largely yellow, often laterally or apically with black markings (~). Scutellum
always with yellow markings (~ 6'). Dorsolateral face of propodeum always and
gastral sternite 2 sometimes with large yellow spots (~ 6'). Legs extensively marked
with yellow ( ~ ) ........................................................................................................................... 2
2. Upper tooth of superior ridge of propodeum always yellow; this yellow part is united
with a large yellow marking, which spreads over the dorsolateral face of propodeum ( ~
6'). Clypeus often with lateral black or brownish markings ( ~ )... subsp. flavicornis Yam.
- Upper tooth of superior ridge of propodeum usually black; when yellow it is separated
from the yellow marking of propodeum (~ 6'). Clypeus sometimes with black or
brownish markings apically and less often laterally { ~) .............. subsp. ryukyuensis Tano
Euodynerus nipanicus nipanicus (Schulthess)
(Figs. 94-%, 99, 103, 104, 107, 110A)
Odynerus micado Kirsch: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 682, pI. 39, fig. 14 (subgenus
Leionotus); 1930, Ill. Thous. Ins. Jpn. 2: 13-14, pI. 2, fig. 14; Yano, 1932, Icon. Ins. Jpn. 1st ed. p. 308.
Odynerus micado Kriechb.: Matsumura, 1931. 6000 Ill. Ins. Jpn.-Emp. p. 16, no. 78.
53
Odynerus (Lionotus) flaviclypeatus Sonan, 1930, Trans. Nat. Hist. Soc. Formosa, 20: 355 (J')(type loc.:
Kagoshima, Kyushu).
Rhynchium satsumanus (!) Son an, 1930, Trans. Nat. Hist. Soc. Formosa, 20: 356
(~Htype loc.:
Kagoshima, Kyushu).
Odynerus nigripes: Yasumatsu, 1935, Kontyu, 9: 223-225.
Odynerus quadrifasciatus: Yasumatsu, 1950, Icon. Ins. Jpn. 2nd ed. p. 1456, fig. 4202; Ishikawa, 1965,
Icon. Ins. Jpn. Col. Nat. Ed. 3: 297, pI. 149, fig. 3.
Euodynerus notatus ssp. 2, 3: Yamane, 1979, New Entomol. 28: 10-11, figs. 18-20,23,24 (in key).
Euodynerus notatus nipanicus: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 337-338, fig. 7.
Euodynerus notatus pubescens: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 337, fig. 6.
Japanese name: Mikado-dorobachi.
Diagnosis. Female. Black, with the following parts yellow: a basal marking on clypeus
(usually absent), an interantennal spot, basal triangular marking on mandible, small spot
on temple, a medially narrowed band on the horizontal part of pronotum anteriorly,
tegula and parategula largely, a spot on dorsal mesepisternum, irregular marking(s) on
scutellum (often lost), metanotum largely, irregular marking(s) on dorsolateral face of
propodeum (often lost), propodeal valve, apical bands on gastral tergites 1-3 (sometimes
also 4), a medially widely interrupted apical band on sternite 2. Legs blackish brown; mid
and hind tibiae often irregularly marked with yellow.
Male. More extensively marked with yellow. Clypeus almost wholly, antennal scape
below, anterior face of mid and hind coxae, apical 1/3 - 2/3 of all femora below, outer
faces of all tibiae yellow. Gastral tergites 1-5 usually with apical bands; sternite 2 with a
pair of irregular yellow markings and a complete apical band; posterolateral corners of
sternite 3 often with yellow spots. All the tarsi more or less yellOWish.
In the specimens from Hokkaido, apical lamella of tergite 2 less developed (Fig. 100);
in the male the lamella not strongly reflexed (Fig. 96). Yellow markings less extensive:
clypeus, mesepisternum, scutellum and propodeum almost always without yellow
markings <Fig. 103), and apical band on sternite 2 of the male often medially interrupted.
The description of the "female" of Odynerus micado by Matsumura (1911, 1931) well
Figs. 103-106. Body color pattern of Euodynerus nipanicus (after Yamane & Tano, 1987). 103, ssp.
nipanicus from Hokkaido; 104, ssp. nipanicus from Honshu - Kyushu; 105, ssp. jlavicornis; 106,
ssp. ryukyuensis.
54
agrees with the male sex.
Material examined. Hokkaido: 1 t, Sapporo, 25 viii 1935 (Y. Sugihara), 2t t, same loc., 4-10 vii 1954 (K.
Kamijo), 1!f, same loc., 14 vii 1955 (K. Kamijo), 1!f, Miyanomori, Sapporo, 5 vii 1961 (SKY), 1!f, Obihiro, 3
viii 1975 (M. Usui), 1!f, Sapporo, 9 viii 1976 (T. Sunose).
Honshu: lwate-ken - 1-'f, Kuriyagawa, Morioka, 16 vi 1964 (YM), 1-'f, same Ioc., 25 viii 1968 (YM), 2 t t
1!f, JoMji, 21 viii 1%2 (HI); Fukushima-ken - 1!f, Yunohana, 27 vii 1980 (HI); NiiglltJl-lcen - 1 t2!f !f,
Nagaoka, 22-23 viii 1951 (Yamazaki), 1 tl!f, same loc., 29 viii 1952 (yamazaki), 1 t, Fukushima-gata, 10 ix
1977 (HI), 1!f, same loc., 9 ix 1979 (HI), 1 t, Shidaihama, 20 viii 1977 (HI), 1!f, same loc., 11 ix 1981 (HI),2t
t, Shibata, 21 ix 1978 (HI), 1 t, Kami-ishikawa, Shibata, 12 viii 1979 (HI), 1 t, Senami, 31 v 1983 (KB), 1 t,
same loc., 13 vi 1984 (KB), 1!f, same loc. 25, ix 1984 (KB); Nllgllno-ken -1 tl!f, Habiro, Ina, 1 ix 1%1 (YM), 1
t, Minamiminowa, Ina,25 v 1%2 (YM), 4t t, same loc, 8-17 vi 1962 (YM), 1 t, Niiyama, Ina, 24 vii 1%2
(yM), 2t t, Habiro, Ina, 27-31 vii 1962 (YM), 2t tl!f, same Ioc., 4-10 viii 1%2 (YM), 4t t, same loc., 1731 viii 1%2 (YM), 2 t tl!f, 1-12 ix 1%2 (YM); Ishikawa-ken - 1 t, Nakashima, 26 vii 1948 (I. Togashi); Aichiken - 1!f, Irako, Atsumi Pen. 6 viii 1977 (Y. Takai); Kyo/o-fu -1 t, Saga, 27 v 1955 (K. Iwata), 1!f, Gosho, 18 vii
1955 (K Iwata); Wakllyama-ken - 4t t, Kiitanabe, 4 viii 1965 (K. Iwata); Hyogo-ken - 2!f!f, Sasayama, Tamba,
17 x 1952 (K. Iwata); Shimane-lcen- 1 t, Mt. Makuragi, 31 v 1983 (yM), 1 t, Nishikawatsu, Matsue, 22 vii 1985
(M. Goubara).
Oki Is. (Shimane-ken): NishinlHlhima -1!f, Urago, 11 v 1982 (YM).
Shikoku: KOchi-ken - 1 t, Monobe, Nankoku, 2 v 1974 (SI), 1!f, same loc., 23 vi 1975 (SI); Kagawa-ken - 1
tl-'f, Oiwagaike, 26 vii 1964 (K. Iwata).
Kyushu: Fulcuoka-ken - 1 J', Harumachi, Fukuoka-shi, 5 vii 1958 (YM), 1 J', Hakozaki, Fukuoka-shi, 14
ix 1958 (YM), 2t tl!f, same loc., 5-23 vii 1959 (YM), 1 tl!f, Wajiro, Kasuya, 15 ix 1959 (YM); Nagasaki-ken 1 t, Haraguchi, Omura, 16 viii 1%7 (R. Ohgushi), 1!f, same loc., 27 viii 1%7 (R. Ohgushi); Kag06hima-ken 10 t t, Korimoto, Kagoshima-shi, 6-7 viii 1981 (SKY), 1 t, same loc., 26 viii 1981 (SKY), 1 t, Meiwa,
Kagoshima-shi, 15 viii 1981 (SKY), 2 t t, Shiroyama, Kagoshima-shi, 30 vii 1981 (SKY), 1 t, Ichiki, 8 v 1984
(AN), 3t t, Kanoya, 30 viii - 8 ix 1981 (SI), 1 t, Sata, 10 vi 1979 (H. Nagase), 1 t, Onakano, Sata, 10 x 1978
(H. Nagase), 1!f, Makurazaki, 8 ix 1984 (AN).
Tsushima Is. (Nagasaki-ken): Kami-1lgatJl-l t, Nii, Toyotama-son, 19 viii 1968 (I. Hiura).
Island dose to Kagoshima-ken-hondo: Alcune-6shima - 1 t, 5 viii 1983 (SKY).
N. Ryukyus: Tane-ga-shima - 1 t, Hamada, 11 vii 1983 (SKY), 3 t t, Kaminaka, 12 vii 1983 (SKY), 1!f,
Hamada, 2 viii 1984 (SKY), It, Ikeno, 21 vii 1984 (S. Watahiki), 1!f, Kamome, 5 viii 1986 (M. Tatsuno);
Mage-shima - 4 t t, 22 vii 1984 (5. Watahiki); Yaku-shima -1 t, Miyanoura, 10 v 1981 (SKY), 11 t t2-'f, same
loc., 8-10 viii 1981 (SKY), 1-'f, Onoaida, 9 viii 1981 (SKY), 1 t, Ambo, 11 v 1981 (SKY), 2t t, Shitogo, 10 viii
1981 (SKY), 4 t t, Miyanoura, 26-28 v 1982 (SI), 1-'f, Onoaida, 29 vi 1982 (SI), 1 t, same loc., 27 vii 1982 (SI),
2t t, Isoo, 30 vii 1988 (SKY); Kuchinoerabu-jima - 1 t, 16-17 v 1982 (SKY), 1!f, Hommura, 18 v 1989 (H.
Watanabe).
Distribution. Hokkaido; Honshu; Iwai-jima; Oki Is. (Nishino-shima); Shikoku;
Kytishu; Tsushima Is. (Kami-agata; Shimo-agata); Amakusa Is. (Shimo-jima); Akuneoshima; Osumi Is. (Tane-ga-shima; Mage-shima; Yaku-shima; Kuchinoerabu-jima);
Ogasawara Is. (introduced ?).
Biology. Iwata (1938b, 1979) intensively studied the biology of this form (referred to as
Odynerus nigripes). This wasp is bivoltine and a tube-renter, usually nesting in bamboo or
reed tubes placed horizontally above the ground. Nests are also made in tubes lying on
the ground, those placed vertically, small holes in the stone, deserted mud cells of Eumenes
fraterculus on stones and of Scelifron madraspatanum, and empty cocoons of Monema
flavescens (Lepidoptera, Heterogeneidae). The number of brood cells constructed per nest
ranged from 1 to 8 (n=50, m=4 [26%]). Empty cells are often constructed between brood
cells as well as at the entrance, and are usually smaller than the brood cells in length. The
wasp sometimes arranges the brood and empty cells alternately. This behavior is
exceptional among the Japanese Eumenidae. The cell partitions are made of mud, and not
water-proofed, measuring 1 to 3 mm in thickness. The outermost wall is much thicker (555
10 mm thick) and water-proofed. The female wasp hunts larvae of Microlepidoptera,
especially of Tortricidae. This species is used to control leaf-eating pests (Tortricidae) of
apple trees (Takeshima, 1971; Iwata, 1979; see Chapter 1). The female does not guards her
nest, and quickly provisions and closes cells one after another.
Parasite: Pseudoxenos iwatai (Strepsiptera, Stylopidae). Parasitoids: a phorid, a
tachinid (Diptera), Chrysis ignita, C. viridula(?) (Hymenoptera, Chrysididae), Macrosiagon
nasuta (Coleoptera, Rhipiphoridae), and Argyromoeba distigma.
Euodynerus nipanicus flavicornis Sk. Yamane
(Figs. 92, 93, 98, 105, 110A)
Odynerus flavolineatus Smith: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 111, no. 681, pI. 39, fig. 13;
Matsumura and Uchida, Ins. Matsum. 1: 36; Kuroiwa, 1926, Trans. Nat. Hist. Soc. Formosa, 16: 140;
Matsumura, 1930, Ill. Thous. Ins. Jpn. 2: 13, p1.2, fig 13; 1931, 6000 Ill. Ins. Jpn.-Emp. p. 16, no. 77.
(Misidentification.)
Euodynerus no/atus flavicarnis: Yamane & Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 339, fig. 8 (Sf d')
(type Ioc.: Okinawa-jim a).
Japanese name: Kisuji-dorobachi.
Diagnosis. Very distinctive in coloration from the nominotypical subspecies of Japan
proper. Apicallamellate area of gastral tergite 2 in the male very narrow (Figs. 92,93).
Female. Black, marked with yellow as follows: clypeus largely, frontal marking, a
stripe at the base of mandible, antennal scape below, stripe behind each eye, wide
pro notal band medially narrowed, pro notal tubercle partly, tegula except for margins and
central brown spot, parategula, metanotum including posterior vertical face above, spot
under wing base, dorsolateral face of propodeum usually wholly; a very wide band on
gastral tergite 1 laterally much dilated, a wide, laterally dilated apical band on tergite 2,
narrow apical bands on tergites 3 and 4, posterolateral corners of gastral sternite 2
(sternite 2 often with a yellow, narrow apical band), mid and hind coxae partly, apical 1/3
to 1/2 of all femora, tibiae of all legs. Clypeus often with lateral black areas, and rarely
with a black or brown mark apically. Mandible brownish. Superior ridge of propodeum
(especially upper tooth) yellow or orange yellow; the upper tooth involved in the large
yellow marking on propodeal dorsum. Yellow of femora and tibiae tinged with brown;
tarsi dark brown.
Male. Similar to the female, more extensively marked with yellow. Clypeus entirely
yellow; frontal yellow mark often with a projection which reaches the upper margin of
clypeus; legs largely yellow.
Material examined. C. Ryukyus: Kikai-jima -1 Sf, viii 1959 (E. Nitta), 1 d'1 Sf, 5 viii 1961 (H. Tanaka), 1 d'1
Sf, 26-27 viii 1984 (S. Watahiki); Amami-&hima - 1 Sf, 20 viii 1943 (HI07), 1 d'1 Sf, Shimmura, Sumiyo, 11 vii
1981 (yH), 1 d', Gusuku, 4 v 1981 (T. Fujisawa), 2d' d'3Sf Sf, Koniya, 22 vi 1987 (SKY), 1 d', Sumiyo, 22 vi
1987 (SKY), 1 d', Nishinakama, 25 vii 1987 (AN), 1 d', Koniya, 14 viii 1987 (M. Tatsuno), 2 Sf Sf, Koniya, 20 x
1987 (SI); Kakeroma-jima -1 d', Osai, 27 ix 1987 (SKY), 4d' d'1 Sf, Nishiamuro, 28 ix 1987 (SKY); Uke-shima - 26'
6'3 Sf Sf, 12 viii 1987 (M. Tatsuno); OIcinoerabu-jima-lSf, 17 vii 1984 (M. Maegata); Yoron-tO- ISf, 6 vi 1985
(SKY); lzena-jima - 1 d', Nakada, 25 vi 1967 (T. Kifune); Okinawa-jima-l 6', Hyakuna, 1 vii 1981 (AN), 7 d' 6'1
Sf, Hentona, 1-6 vi 1983 (AN), 1 Sf, same loc., 27 vii 1987 (SKY); Sezoko-jima - 1 d', 28 v -1 vi 1982 (y. Ikimori
& Y. Ohira); Tokashiki-jima -1 Sf, 11 x 1988 (SKY); Miyagi-jima -1 d', 31 vii 1971 (T. Kifune).
Distribution. Amami Is. (Kikai-jima; Amami-6shima, Kakeroma-jima; Uke-shima;
Tokuno-shima; Okinoerabu-jima; Yoron-to; Okinawa Is. (lzena-jima; Okinawa-jima;
56
Sezoko-jima; Yagaji-jima; Miyagi-jima; Tokashiki-jima; Kume-jima).
Biology. No information is available.
Euodynerus nipanicus ryukyuensis Tano
(Figs. 106, 1l0A)
Odynerus f1avalineatus: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 111, no. 681, pI. 39, fig. 13;
Matsumura and Uchida, Ins. Matsum. 1: 36; Matsumura, 1931, 6000 III. Ins. Jpn.-Emp. p. 16, no. 77.
Euodynerus naliltus ryukyuensis: Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 339-340, fig. 9
(!f J')(type loc.: Ishigaki-jima, Yaeyama Is.).
Japanese name: Yaeyama-mikado-dorobachi.
Diagnosis. Female. Similar to the subspecies fiavicornis, but distinguished from the
latter by the following points: apical portion of clypeus more often marked with brown
(lateral parts also often marked with black as in [lavicornis), interocular area above with a
small yellow spot close to each eye, frontal marking often with a narrow projection which
reaches the upper margin of c1yepus, yellow marking of dorsolateral face of propodeum
less developed and not involving the upper tooth of superior ridge (Fig. 106).
Male. Lacking interocular spots. Distinguished from the preceding subspecies by the
less developed yellow marking on the dorsolateral face of propodeum.
Material examined. S. Ryukyus: Miyako-jima - 5 J' J'1!f, Ueji, 28 ix 1988 (51); Tarama-jima - 1!f, 18 vii
1987 (SKY), 3 J' J'1!f !f, 29 vi 1988 (SKY); Ishigaki-jima - 2 J' J'1!f, Shinkawa, 19 vi 1986 (51), 1!f, Kabira, 22
vii 1987 (SKY), 1!f, Shiraho, 26 vi 1987 (SKY), 5 J' J'1!f, Omoto-dake, 8 vii 1988 (K. Nakamine); Kohama-jima
-1!f, 25 vii 1987 (SKY); Iriomote-jima - 2!f!f, Toyohara, 30 vii 1983 (AN), 1 J', Otomi, 29 vii 1983 (AN), 1 J'1
!f, Ohara, 30 vii 1983 (AN), 1!f, same loc., 25 vii 1983 (A. Matsumoto), 4J' J', 17-18 viii 1983 (S.F. Sakagami
& YM), 1 J', Otomi, 25 vii 1985 (AN), 1 J', Uehara, 2 vii 1988 (SKY), 1 J'1!f, Ohara, 20 vi 1988 (K. Nakamine);
Hateruma-jima -1 J'1!f, 1 vii 1988 (SKY); Yonaguni-jima - 9 J' J'3!f !f, Sonai, 5 vii 1988 (SKY).
Distribution. Miyako Is. (Miyako-jima); Tarama Is. (Tarama-jima); Yaeyama Is.
Oshigaki-jima; Kohama-jima; Iriomote-jima; Hateruma-jima; Yonaguni-jima).
Biology. On Tarama-jima (June 30 1988) I observed some female wasps nesting in
bamboo tubes placed horizontally as supports for melon vines at about 1.5 m above
ground.
Euodynerus (Pareuodynerus) quadrifasciatus (Fabricius)
(Figs. 74, 86, 91, 109)
Vespz quadrifasciata Fabricius, 1793, Entomol. Syst. 2: 266 (J')(type loc.: Denmark).
Odynerus (Uonotus) toment06us Thomson, 1870, Opusc Entomol. 2: 86.
?Odynerus (Lionotus) quadrifasciatus Herr.-Schaff.: Schulthess, 1934, Arb. Morph. Taxon. Entomol. 1: 75
(in key).
Japanese name: Nise-mikado-dorobachi.
Diagnosis. Body length (h+th+tl +2): 8.5-10.0 mm in -'f-, 6.0-7.5 mm in d'. Fore wing
length: 8.5-10.0 mm in -'f-, ca. 9 mm in d'. Structural ~haracters as in the key.
This species has long been confused with Euod. nipanicus (=Euod. notatus sensu
Yamane & Tano, 1987) under the Japanese name "Mikado-dorobachi". Two subspecies are
known in Japan, and both are relatively rare.
57
Euod.
nipanicus
,
o
Euod.
dantici
108
• Euod. quadrifasciatus eburnus
lIfEuod. q. atripes
eEuod. trilobus
Figs. 107-109. Distribution of the four Euodynerus species on the Japanese mainlands.
Information about the nesting biology of this species in Japan is not available.
According to Nielsen (1932) and Bliithgen (1961), in northern Europe, the species nests in
pre-existing burrows in wood. Brood cells are provisioned with larvae of tortricid moths.
Euodynerus quadrifasciatus eburnus Sk. Yamane
(Fig. 109)
Euodynerus quadrifasciatus subsp.: Yamane, 1979, New EntomoI. 28: 10 (in key), figs 5, 15,21.
Euodynerus quadrifasdatus eburnus: Yamane and Tano, 1987, Trans. Shikoku EntomoI. Soc. 18: 340-341
(~ J')(type loc.: Sapporo, Hokkaido).
58
Diilgnosis. Female. Black, marked with ivory-white and pale yellow as follows: frontal
mark, a small spot on mandibular base, small spot behind each eye, narrow pronotal band
interrupted medially, anterior and posterior spots on tegula, dorsal face of metanotum,
narrow regular bands on gastral tergites 1-4, posterolateral corners of gastral sternite 2.
Antenna and legs almost wholly black; tarsi tinged with reddish brown.
Male. Very similar to the female. In addition to the markings mentioned for the
female, the following parts are ivory-white or pale yellow: c1ypeus wholly, antennal scape
below, mandible largely, an apical band on tergite 5, posterolateral corners of sternite 3,
anterior faces of mid and hind coxae and mid femur, and outer faces of all tibiae.
This form differs from the nominotypical subspecies by the tegula and metanotum
always marked with pale color, and the almost wholly black legs as well as ivory-white
body markings.
Material examined. Hokkaid6: 1 J', Toyotaki, Sapporo, 2 vii 1972 (SKY), 1 ~, same loc., 2 vii 1980 (S.
Makino) (holotype), 1 J', Iwaobetsu, Shiretoko, 10 vii 1975 (T. Matsumura).
Distribution. Hokkaido.
Euodynerus quadrifasciatus atripes Giordani Soika
(Fig. 109)
Euodynerus ifuadrifasciatus atripes Giordani Soika, 1976, Ann. Hist.-Nat. Mus. Nat. Hung. 68: 292 (~)
(type loc.: Ryang-gang, N. Korea); Yamane and Tano, 1987, Trans. Shikoku Entomol. Soc. 18: 341 (from Japan
proper).
Diilgnosis. Female. Much as in the preceding subspecies except that body markings
are yellow or orange yellow. Other minor differences are: pro notal band in each side
posteriorly dilated (in eburnus posterior margin straight); anterior spot on tegula usually
much smaller than posterior one, sometimes even lost.
Male. Distinguished from the preceding subspecies by the darker coloration and the
following variations: scutellum sometimes with a pair of small yellow spots, sternite 2
rarely with a yellow apical band in addition to lateral markings.
Material examined. Honshu: Aomori..Jcen - 1 J', Hirosaki, 5 vi 1983 (M. Yamada); Gumma-ken - 1 J',
Onioshidashi, Tsumagoi, 19 vii 1%7 (T. & H. Suda); Nagano-ken - 1 ~, Mt. Yatsugatake, Chino, 3 vii 1963
(yM).
Distribution. Central and northern Honshu. Korea.
Taxonomic notes. This form was originally described by Giordani Soika (1976) based
upon a single female from Korea, but his description was so brief that the present
treatment of the Japanese population should remain tentative.
Euodynerus (Pareuodynerus) bicingulatus Giordani Soika
Euodynerus (Pareuodynerus) bicingu/atus Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 138139, fig. 41 (J')(type loc.: Tokyo (?».
Diilgnosis. Giordani Soika's original description will be reproduced here (translation
byrne).
[Male. Similar to E. notatus (Jur.), from which the present species is distinguished by
the following characters: Antenna much longer; segments 5-11 all longer than wide.
59
Temple less developed, seen from above much shorter than the superior lobe of eye.
Pronotal carina not angulate on the dorsolateral parts. Parategula well developed, long,
finger-shaped; its apex extending beyond the apex of tegula. Superior carina of
propodeum less developed; instead, lateral carina much more developed than in notatus,
forming a large black lamella which, connected with the inferior carina, forms a robust
denticle that is flattened triangle in shape. Tergite 2 not reflexed at apex. Sternite 2 convex
at base, slightly convex posteriorly, without median longitudinal furrow at base.
Punctation on the first two tergites coarse and dense, with interspaces in average
smaller than punctures; near the apex of tergite 2 punctures denser but not larger. On
tergites 3-4 and 5 punctation densest, considerably coarser than on tergite 2. Tergite 6 with
punctures of the same size but rather sparse. Sternite 2 laterally with large and dense
punctures; punctures much smaller and sparser in the middle.
Black, with the tegula and legs reddish brown or blackish brown. Yellow are: clypeus,
antennal scape below, a rhombic frontal marking, a narrow line on temple, pro notal band
medially narrowed, a marking on mesepisternum, a line on tegula, parategula, small
irregular spots on scutellum, a very narrow line on metanotum, apices of femora,
markings on fore and mid tibiae, apical band on tergite 1 that is narrow and regular, apical
band on tergite 2 that is regular and slightly wider, markings at posterolateral corners of
sternite 2. Wings diffusely brownish.
Body length (to the posterior margin of tergite 2): n mm.
Female unknown.]
Distribution. Hokkaid6 (?); Honshu (f6ky6)(?).
Taxonomic notes. I have never seen any Euodynerus specimen that agrees with Giordani
Soika's description of Euod. bicingulatus. On the other hand, small male specimens of
Anterhynchium flavomarginatum micado well agree with his description, with minor
disagreements in the condition of punctation. In this paper I include this "species" in this
genus tentatively, but not in the key to the Japanese species.
Euodynerus trilobus (Fabricius)
(Figs. 75, 87, 109, nOC)
VesPl triloba Fabricius, 1787, Mant. Ins. 1: 290 (type loc.: China (?».
Odynerus trilobus: Schulthess, 1934, Arb. Morph. Taxon. Entomol. 1: 74 (in key); Sonan, 1938b, Trans.
Nat. Hist. Soc. Fonnosa, 28: 79-80; Yasumatsu, 1950, Icon. Ins. Jpn. 2nd. ed. p. 1456, fig. 4200.
Odynerus emma 5<man, 1929, Trans. Nat. Hist. Soc. Fonnosa, 19: 535 (J')(type loc.: Penghu 1. (= Hokoto), Taiwan).
Euodynerus trilobus: Vecht and Fischer, 1972, Hym. Cat. (n. ed.), 8: 101; Yamane and Tano, 1987, Trans.
Shikoku Entomol. Soc. 18: 341-342.
Japanese names: JUji-dorobachi (Emma-hime-dorobachi; Emma-dorobachi).
Diagnosis. Body length (h+th+t1+2): 7.5-10.5 mm in !f, 6.5-8.0 mm in J'. Fore wing
length: 9.5-10.0 mm in !f, 6.5-9.0 nun in J'. Structural characters as in the key.
Female. Black, marked with yellow or orange yellow as follows: a basal bar on
clypeus, frontal spot, small spot behind each eye, a small marking on mandibular base,
antennal scape below, a relatively wide band (medially interrupted) anteriorly on
pronotum, tegula except for margins and central spot, parategula, a relatively large spot
under wing base, scutellum partly (seen only in the specimens from Yaeyama Is.), dorsal
60
•
e'~
~PI
J
C
c6
If
I'
iF
,,0
c6
•
r
.,
l)
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.
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. .
'
~
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.
~"
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~
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.~
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.
i~
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de
~e
o~
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S
r"iu~'
Figs. 110. Distribution of Euodynerus nipanicus (A; three subspecies), Euod. dantici (B) and Euod.
trilobus (C) in the Ryukyus.
110
face of metanotum, dorsolateral face of propodeum partly, laterally dilated band on
gastral tergite 1 (the band in the middle slightly incised), slightly sinuated apical bands on
tergites 2-5, posterolateral corners of gastral sternites 2-4, apical portion of mid and/or
hind femora apically, anterior face of mid coxa (often lost).
Male. Differs from the female as follows: clypeus and mandible entirely yellow;
frontal spot connected with clypeus by interantennal yellow mark; ocular sinus sometimes
with a yellow stripe in inferior part, gastral tergite 2 with isolated yellow spots laterally;
apical band on tergite 2 laterally widened; tergite 6 often with a yellow apical band;
sternite 2 with yellow lateral markings which are very large; sternite 2 with a narrow,
yellow apical band; sternites 3-5 with posterolateral corners yellow; anterior faces of all
legs almost wholly, fore and mid femora below, outer faces of tarsi largely yellow (other
parts of legs brownish).
Material examined. Honshu: Wakayama-ken - 3 t t, Kozanji, Tanabe, 4 viii 1965 (K Iwata).
Island close to Kagoshima-ken-hondo: Akune-6shima - 6 t t, 5 viii 1983 (SKY).
N. Ryukyus: Kuro-shima- 1~, Osato, 4 ix 1981 (SKY); Yaku~hima - 2tt, Koseta, 17 vii 1972 (K.
Kusigemati), 5t t, Shitogo, 10 viii 1981 (SKY).
C. Ryukyus: Takara-jima - 3 t 6'3~ ~, 16-19 vii 1964 (A. Tanaka), 3 t t, 29-30 v 1982 (KT), 5 t t, 29 vii
1986 (51), 1 t, 16 viii 1988 (SKY); Yoron-tO - 2 t t1 ~, 4-6 vi 1985 (SKY); lzena-jima -1 ~, Nakada, 1 viii 1967
(T. Kifune); Okinawa-jima -1 ~, Toyogusuku, 4 viii 1976 (5. Yamauchi);
S. Ryukyus: IshigaJci-jima -1 t, 19 iv 1981 (T. Fujisawa); lriomote-jima 1 t, Ohara, 15 v 1981 (AN), 1~,
Toyohara, 20 v 1981 (AN), 1 t, Otomi, 25 vii 1985 (AN), 1~, Toyohara, 26 vii 1985 (AN), 1~, Funaura, 22 vi
1986 (51), 1 ~, Uehara, 2 vii 1988 (SKY); Yonaguni-jima - 1 t, 22-24 vii 1983 (H. Kodama), 2 t t, Sonai, 5 vii
1988 (SKY).
Daito Is.: KitadaitO -jima -18t t6~~, 19 viii 1987 (51); Minamidait6-jima- 2t t1 ~,20 viii 1987 (51).
Distribution. Honshu; Shikoku(?); Kyushu; Akune-6shima; Osumi Is. (Kuro-shima;
Yaku-shima); Tokara Is. (Takara-jima); Amami Is. (Amami-Oshima; Okinoerabu-jima;
Yoron-to); Okinawa Is. (lzena-jima; Okinawa-jima; Yagaji-jima; Kume-jima); Miyako Is.
(Miyako-jima); Yaeyama Is. (lshigaki-jima; Iriomote-jima; Yonaguni-jima); Daito Is.
(Kitadaito-jima; Minamidaito-jima). Taiwan; continental China; Africa.
Taxonomic notes. Although widely distributed in the Palearctic region and northern
part of the Oriental region, this species has not been divided into geographical races
(Vecht & Fischer, 1972). In at least Japan, the color pattern is relatively stable throughout
its range. The slight geographical variation is set off by intra populational one. This species
is widely distributed in the southern parts of Japan, but seems to be very rare in Honshu
and Shikoku.
Biology. Most of the known localities in Japan are restricted to the coast (d. Nagase,
1981). Iwata (1939b) observed females flying around driftwood thrown up on the eastern
shore of southern Taiwan (referred to as Pseudoepipona trilobus). Wasps were nesting close
to each other in burrows (3-5 cm deep, 6 mm in diam.) dug in the wood. Two or three
brood cells and an empty cell were constructed in each burrow. Cell partition was made of
red clay mixed with sand, 0.5-1 mm in thickness, and very fragile; it is hardened by
cocoon lining. Prey: larvae of Tortricidae(?).
Parasitoid: Hexachrysis sp. (Hymenoptera, Chrysididae).
62
Genus Ryhnchium Spinola
Rygchium Spinola, 1806, Ins. Ligur. 1: 84 (type species: Rygchium europaeum Spinola, 1806 (= Vespa
oculilta Fabricius, 1781»; Saussure, 1852, Et. Fam. Vesp. 1: 101.
Rhynchium: Vecht, 1963, Zool. Verh. 60: 109.
Japanese names: O-dorobachi Zoku (Fukai-dorobachi Zoku).
This genus is an Old World group, comprising relatively large-sized species mainly
distributed in Middle East and Africa. A brief diagnosis is given in the key to the Japanese
genera. Vecht (1963) mentioned that in the wasps of this genus the scutellum and posterior
part of mesoscutum are smooth and at most finely and sparsely punctate. In the Japanese
populations of R. quinquecinctum (especially of the mainlands), these parts are often
distinctly punctate. This fact and his misunderstanding of Vecht's (1963) paper led Sato
(1963) to conclude that the single Japanese species is a member of Anterhynchium.
Two other forms, very closely related to quinquecinctum (Fabricius, 1787), are known
from eastern Asia. R. brunneum (Fabricius, 1793) was described from Bengal, and has often
been considered to be a synonym or a geographical race of quinquecinctum. However,
Giordani Soika (1986) and Gusenleitner (1988) treated both forms as separete species.
Giordani Soika stated, "la descrizione del quinquecinctum corrisponde bene a molti
esemplari della China, mentre gli esemplari a me noti dell'India, Burma e Tonkino hanno
la colorazione del brunneum". On the other hand, these two forms have sometimes been
treated as synonyms or subspecies of R. hilemorrhoida1e (Fabricius, 1775) (e.g., Dalla Torre,
1904), but at least quinquecinctum is specifically separated from haemorrhoida1e as follows:
Scutellum and posterior part of mesoscutum finely and sparsely punctate (.'f. 6').
Clypeus with fine white hairs only ( 6'). Narrow longitudinal depression just adjacent
to the outer side of each posterior ocellus distinct ( .'f. 6') .......... R. quinquecinctum (Fab.)
Scutellum in anterior 2/3 and posterior part of mesoscutum almost impunctate (.'f. 6').
Clypeus with dark bristles in addition to fine hairs ( 6'). Longitudinal depression near
each posterior ocellus obscure or absent ( .'f. 6') ............................ R. haemorrhoidl:l1e (Fab.)
Rhynchium quinquecinctum (Fabricius)
(Figs. 111-117, 147)
Vespa quinquecincta Fabricius, 1787, Mant. Ins. 1: 288 (type loc.: China (1».
Rhynchium haemorrhoidale var. quinquecinctum: Dalla Torre, 1894, Cat. Hym. 9: 45; 1904, Gen. Ins. 19: 34;
Liu, 1936, Peking Nat. Hist. Bull. 11: 109-110; Sonan, 1937, Trans. Nat. Hist. Soc. Formosa, Z7: 107-109.
Rhynchium quinquecinctum: Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S. Pac. 4: 122123; Lee, 1985, Econ. Ins. Fauna China, 30: 142-143, pI. 9, fig. 14.
Japanese names: Fukai-O-dorobachi (Fukai-dorobachi).
Diagnosis. Female. Body length (h+th+t1+2): 12.5-17.0 mm. Fore wing length: 13.015.5 mm. Head subcircular, moderately punctate on vertex and gena, and strongly
punctate on frons above supradypeal area that is almost impunctate. Supradypeal area
with a distinct interantennal vertical keel, which is distichous above. Clypeus pyriform,
moderately punctate, with the apex narrowly truncate (Fig. 111); spaces between
punctures forming vertical carinae in the middle of the disc. Ocellar region with a narrow
longitudinal depression just adjacent to the outer side of each posterior ocellus.
63
111
10"
115
113
116
Figs. 111-116. Rhynchium quinquecindum. 111, female head (frontal view); 112, male c1ypeus; 113,
terminal segments of male antenna (from below); 114-116, body color pattern in ssp. nambui
(114), ssp. murotai (115), ssp. fukaii (116).
Depression for the cephalic foveae large, distinct, with dense hairs and a longitudinal
median keel, and posteriorly well defined by an elevated rim. Anterior vertical face of
pronotum smooth, shining; posterior horizontal part and lower lateral part densely
punctate; a distinct depression just anteriorly to the posterior pronotai lobe. Mesoscutum
strongly and densely punctate in anterior 2/3, finely and sparsely punctate in posterior
1/3; scutellum finely and sparsely punctate. Mesopleuron more coarsely punctate,
somewhat reticulate except for epicnemium, which is almost impunctate and well
demarcated by epicnemial carina. Metanotum very densely punctate, medially weakly
depressed, with blunt lateral projections; posterior vertical face almost impunctate in
lower 2/3, dull. Metapleuron transversely striate in upper 1/4 and lower 1/2. Propodeum
without a shelf, with spined inferior ridges; dorsal face reticulate; lateral face striate;
concavity with a V-shaped median carina, a triangular area just below metanotum and
many striae, and shining. Gastral segments punctate much more finely than alitrunk.
Gastral sternite 1 strongly punctate and transversely carinate; sternite 2 distinctly
64
punctate.
Male. Body length (h+th+t1+2): 9.5-14.5 mm. Fore wing length: 10.0-12.0 mm. Similar
to the female. Clypeus much higher than wide, very superficially punctate, with short fine
hairs, shallowly emarginate at apex (Fig. 112). Antennal segment 12 very small; the apex
of antennal hook (seg. 13) reaching the apex of segment 10 (Fig. 113). Mid femur distinctly
emarginate at base.
The population of the Japanese mainlands has often been dealt with as a distinct
species, R. fulmii Cameron, by authors (e.g., Vecht & Fischer, 1972), but I have concluded
that all the Japanese forms are geographical races of R. quinquednctum (Yamane & Tano,
1983).
Key to the Japanese forms of Rhynchium quinquednctum
1. Alitrunk marked with yellow. Apical bands on gastral tergites yellow or orange yellow,
not tinged with brown. Metanotum constantly marked with yellow........................... .
....................................................... ................................................................. subsp. nambui Yam.
- Alitrunk marked with dark orange yellow to ferruginous. Apical bands on gastral
tergites dark orange yellow, sometimes tinged with brown. Metanotum very often
almost wholly black. ................................................................................................................... 2
2. Scutellum usually dark orange yellow in posterior 1/2 - 2/3. In the male the tibiae of all
legs with yellow markings on outer faces (outer face of fore tibia often extensively
yellow); hind tibia not blackish ................................................................ subsp. murotai Tano
- Scutellar markings much reduced, often lost. Male tibiae usually not marked with yellow; hind tibia often wholly blackish ......................................................... subsp. fulmii Cam.
Rhynchium quinquednctum fukaii Cameron
(Figs. 116, 117)
Rhynchium fu1caii Cameron, 1911, Entomologist, 44: 2i37 «(1) (type loc.: Japan); Yano, 1932, Icon. Ins. Jpn.
p. 307; Yasumatsu, 1938, Ins. Jpn. III. Icon. Col. Nat. Dep. p. 359; Giordani Soika, 1976, Ann. Hist.-Nat. Mus.
Nat. Hung. 68: 292.
Rhynchium haemorrhoidale F.: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 112, no. 684, pI. 39, fig. 16;
1930, lll. Thous. Ins. Jpn. 2: 14-15, pI. 2, fig. 16 (part); 1931, 6000 TIl. Ins. Jpn.-Emp. p. 16, no. SO.
Odynerus (Rhynchium) haemorrhoidalis var. Fukaii: Giordani Soika, 1941, Boll. Soc. Venez. Stor. Nat. 2:
261.
Rhygchium haemorrhoidale fukaii: Yano, 1950, Icon. Ins. Jpn. 2nd ed. p. 1455; Ishikawa, 1965, Icon. Ins.
Jpn. Col. Nat. Ed. 3: 297, pI. 149, fig. 5 (Rhynchium).
Rhynchium haemorrhoidale (E) ssp. samurayi Giordani Soika, 1973, Boll. Mus. Civ. Venez. 24: 119-120 (~)
(type loc.: Tokyo, Honshu). Syn. nov.
Japanese names: Fukai-6-dorobachi (Fukai-<iorobachi).
Diagnosis. Female. Black, with yellow and (dark) orange yellow markings. Yellow are
dypeus (centrally darker), a line on inner orbit below, a transverse marking on frons and
antennal scape below. Other markings are darker, sometimes reddish or brownish: a line
on inner orbit above and genal band (complete) connected with it, a pair of curved lines in
ocellar region, a transverse marking across vertex (interrupted medially at cephalic
65
foveae), upper portion of the anterior vertical face of pronotum, posterior horizontal part
of pronotum largely, a large spot under wing base, tegula posteriorly, posttegula,
scutellum posteriorly (usually lost), irregular markings on metanotum (reddish, often
lost), irregular markings on propodeum (reddish), a narrow apical band on gastral tergite
1, wider ones on tergites 2~, narrow and medially widely interrupted apical bands on
sternites 2-4, sternite 5 largely. Antenna except for scape below, mandible largely, apical
2/3 of fore femur, fore tibia, fore tarsus, mid femur in apical 1/2 below are dark orange or
rufous.
Male. Similar to the female, but differs in the following points: clypeus wholly lemon
yellow; mandible often with a yellow triangular marking; mid and hind tarsi sometimes
marked with reddish brown.
Material examined. Honshu: Niigabl-ken - 15f, "Echigo", 16 vii 1935 (Nohira), 15f, "Echigo", 3 viii 1940, 1
6', "Echigo", 10 ix 1940 (Nohira), 15f, Nagaoka, 26 viii 1951 (yamazaki), 2 5f 5f, Niigata-shi, 'Z7 viii 1963 (YM);
Ibaraki-ken - 15f, Tsuchiura, 9 viii 1987 (SKY); Nagano-ken - 16', Nagano-shi, 15 vii 1%8 (T. Kitamura);
Wakayama-ken - 26' 6', K6zanji, Kiitanabe, 4 viii 1965 (K. Iwata); Hyiigo-ken - 26' 6', Sasayama, Tamba, 30 vii
1964 (K. Iwata); Okayama-ken - 16', Ukai-gawa, 25 viii 1960 (K. Iwata); Shimane-lcen - 16', Kawatsu, Matsue, 6
viii 1985 (N. Sugiura).
Shikoku: Kiichi-ken -1 6', Monobe, Nankoku, 9 vii 1975 (51).
KyushU: Nagasaki-ken -15f, Haraguchi, Omura, 18 vii 1967 (R. Ohgushi); Kag05hima-ken -1 6', Onejime,
Sata,3 ix 1978 (H. Nagase), 26' 6', Korimoto, Kagoshima-shi, ~7 viii 1981 "(SKY), 16', same loc., 26 viii 1981
(SKY).
Distribution. HonshU; Awaji-shima; Shikoku; Kyushu. Korea (Giordani Soika, 1973);
E. Siberia and N. China (Kurzenko, 1984b).
Biology. Nesting biology of this form was studied in Osaka-fu, Honshu, by Iwata
(1938b, 1980c). Nests are made in bamboo and reed tubes. One to seven (usually 1-3)
brood cells are constructed per nest, together with 1-5 empty cells. Nesting proceeds
quickly, two or three cells being completed in a day. Entrance plug is rough, not waterresistant. Caterpillars of Pyralidae (more than 2 cm long) are most preferred, and those of
Tortricidae are only occasionally hunted. A cell is provisioned with three to ten (usually 67) caterpillars.
Parasite: Pseudoxenos iwatai (Strepsiptera, Stylopidae). Parasitoids: Amobia distorta
(Diptera, Sarcophagidae), and a phorid (Diptera).
Rhynchium quinquecinctum murotai rano
(Figs. 115, 147)
Rhynchium fukaii: Yano, 1932, Icon. Ins. Jpn. p. 307; Yasumatsu, 1938, Ins. Jpn. HI. Icon. Col. Nat. Dep. p.
359.
Rhynchium haemorrkoidale: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 112, pI. 39, fig. 16; Matsumura
and Uchida, 1926, Ins. Matsum. 1: 37; 1930, Ill. Thous. Ins. Jpn. 2: 14-15, pI. 2, fig. 16; 1931, 6000 Ill. Ins. Jpn.Emp. p. 16, no. SO.
Rhynchium quinquecinctum murobli: Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S. Pac.
4: 124, fig. 12 (5f 6') (type loc.: Amami-&hima); Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 72-
73.
Japanese names: Murota-o-dorobachi (Murota-dorobachi).
Diagnosis. Similar to the preceding subspecies. Scutellum always marked with dark
orange or reddish brown, while metanotum only rarely with markings of these colors. In
66
the male, tibiae and tarsi of all legs testaceous or rufous; tibiae very often extensively
yellowish on outer faces.
Material examined. N. Ryukyus: Naka-no-shima -1.'f, 16 viii 1943 (Hl00).
C. Ryukyus: Takara-jima - 2 (f' (f', 16 viii 1988 (SKY); Kikai-jima -1 (f'1.'f, 18 x 1987 (SI); Amami-Oshima - 1
.'f, Santaro Pass, 23 vii 1967 (TM), 1 (f', Sumiyo, 29 vii 1973 (K. Sakamoto), 1 (f'1.'f, Sumiyo, 11-13 vii 1981
(yH), 2(f' (f', Koniya, 22 vi 1987 (SKY), 1.'f, Nishinakama, 25 vii 1987 (AN), 2(f' (f', Uttabaru, Kasari, 17-18 x
1987 (SI), 2 (f' (f'1.'f, Shimmura, Sumiyo, 20-21 x 1987 (SI); Kakeroma-jima - 1.'f, 23 ix 1984 (AN), 1.'f, Shadon,
24 ix 1984 (AN), 1 (f'1.'f, Kanyu, 25 ix 1984 (AN), 4(f' (f'4.'f.'f, Osai, 26-27 ix 1987 (SKY), 1.'f, NishiamuJ"O, 28
ix 1987 (SKY); Yoro-shima -1 (f', 13 viii 1987 (M. Tatsuno); Uke-shima - 2.'f.'f, 12 viii 1987 (M. Tatsuno); Tokuno-
_.j
\ .. ,;"_."
l~.:
-..'
R. quinquecinctum
P.
ornatum
118
,
o
~..•
o.
drelVseni
119
Figs. 117-119. Distribution of Rhynchium quinquecinctum, Pararrhynchium ornatum, and
Orancistrocerus drewseni on the Japanese mainlands.
67
shima - 10", Kametsu, 13 vii 1980 (Y. Takai), 1-'?, Mikyo-dake, 6 ix 1983 (M. Ohara), 10", Kametoku, 13 vii
1984 (SKY); OkintmJa-jima - 30" O"8-'?.Sf, "Okinawa" (Sakaguchi), 30" <!'3.Sf.Sf, "Okinawa", 1923 (Sakaguchi), 2
0" 0", Sashiki, 1 viii 1976 (S. Yamauchi), 10", Toyogusuku, 4 viii 1976 (S. Yamauchi), 10"3 -'? -,?, Naha, 29 ix
1977 (SY), 20" O"I-'?, Kudeken, 30 ix 1977 (SY); 40" O"I.Sf, Nakagusuku,3 x 1977 (SY), 10", Ogimi, 26 viii 1979
(H. Nagase), 1 0", Hyakuna, 1 vii 1981 (AN), 3 J' J', Hentona, 1-5 vi 1983 (AN), 1 J', Sate, 5 vi 1983 (AN), 3 0"
J'2.Sf -,?, Yona, 18 vii 1984 (SKY), 3 J' J', Hentona, 27-28 vii 1987 (SKY), 1-'?, Yona, 6 x 1987 (AN), 1.Sf, Nago,
3 x 1987 (AN); Kouri-jima - 2J' J'5-,? -,?, 18 x 1988 (Y. Kusui); Hamahiga-jima - 3J' J', 22 x 1988 (Y. Kusui);
Sezoko-jima -1-'?, 28 v -1 vi 1982 (Y. lkimori & Y. Ohira); Tokashiki-jima - 6J' J'3-,? -,?, 11 x 1988 (SKY).
Distribution. Tokara Is. (Naka-no-shima; Takara-jima); Amami Is. (Kikai-jima;
Amami-Oshima; Kakeroma-jima; Yoro-shima; Uke-shima; Tokuno-shima; Okinoerabujima; Yoron-to); Okinawa Is. (Okinawa-jima; Kouri-jima; Yagaji-jima; Sezoko-jima;
Hamahiga-jima; Tokashiki-jima; Kume-jima).
Biology. Kinjo (in Azuma et al. 1987, p. 188) mentioned nesting sites of this species in
the Ryukyus (see below).
Parasite: Pseudoxenos iwatai (Maeta, 1980; Kifune & Yamane, 1985).
Rhynchium quinquecindum nambui Sk. Yamane
(Figs. 114, 147)
Rhynchium fukaii: Yano, 1932, Icon. Ins. Jpn. p. 307; Yasumatsu, 1938, Ins. Jpn. III. Icon. Col. Nat. Dep. p.
359.
Rhynchium haemorrhoidale: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 112, pI. 39, fig. 16; Matsumura
and Uchida, 1926, Ins. Matsum. 1: 37; Matsumura, 1931,6000 Ill. Ins. Jpn.-Emp. p. 16, no. SO.
Rhynchium quinquedndum nambui: Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S. Pac.
4: 123 (-'? J')(type loc.: Iriomote-jima, Yaeyama Is.).
Anterhynchium Jlaoomarginatum umenoi: Kinjo, 1987, in S. Azuma ed., Field GUide-book to the Insects of
Okinawa, 3: 188 (misidentification).
Japanese names: Nambu-O-dorobachi (Nambu-dorobachi).
Diagnosis. The pattern of body markings is similar to that of the subspecies murotai,
but the markings are much paler, especially on head and thorax. Female mandible usually
with a basal yellow marking. Scutellum and metanotum constantly marked with yellow.
Propodeum often with orange or reddish brown markings.
Material examined. S. Ryukyus: Miyako-jima - 2 -'? .Sf, Hirara, 18 vii 1987 (SKY), 1-'?, Hirara, 28 vi 1988
(SKY); lrabu-jima - 1 J', Sarahama, 28 iv 1981 (T. Fujisawa); Tarama-jima - 2 J' J'1-'?, 19-20 vii 1987 (SKY), 10",
28 vi 1988 (SKY); Minna-jima - 2J' J', 29 vi 1988 (SKY); Ishigaki-jima -1 J', vii 1922 (S. Hirayama), 1-'?, viii
1922 (S. Hirayama), 1-'?, 11 vii 1973 (H. Takizawa), 2 J' 0", Ishigaki-shi, 25 iii - 1 iv 1973 O. Nakayama), 2 -'? .Sf,
same loc., 10 x 1977 (SY), 1!f, Yonehara, 16 iv 1981 (KB), 2J' J'1-'?, Ishigaki-shi, 18 iv 1981 (T. Fujisawa), 1 J'1
.Sf, Ibaruma, 14 x 1987 (AN), 2J' J', Omoto-dake, 3 vii 1988 (K. Nakamine), 30" J', Banna-dake, 3-8 vii 1988
(K. Nakamine), 1 J', Banna-dake, 4 vii 1988 (SKY); Taketomi-jima - 5J' 0",8 vii 1988 (K. Nakamine); Kuroshima - 2J' J', 23 vii 1987 (SKY); lriomote-jima - 2J' J', 17 iv 1962 (G. Kuno), 1-'?, Komi, 11 viii 1972 (TN), 1 J'1
-,?, Funaura, 5-9 x 1977 (SY), 1-'?, Urauchigawa, 6 x 1977 (SY), 1 J', 16 iv 1978 (K. Ohara), 2J' J', Sonai, 20 iv
1981 (T. Fujisawa), 2J' J', atomi, 17 v 1981 (AN), 1-'?, Ohara, 24 v 1981 (AN), 2.Sf -,?, Ohara, 7 viii 1981 (AN),
1.Sf, atomi, 28 iv 1982 (AN), 1-'?, Ohara, 28 iv 1982 (AN), 2J' 0", Toyohara, 29 iv 1982 (AN), 2J' 0", Otomi, 29
vii 1983 (AN), 1.Sf, Komi, 31 vii 1983 (AN), 1-'?, Toyohara, 26 vii 1985 (AN), 1-'?, atomi, 'Z1 vii 1985 (AN), 3 0"
J', Amitori, 30 vii -1 viii 1985 (AN), 2J' J', Ohara, 20 vi 1988 (K. Nakamine), 1-'?, Uehara, 2 vii 1988 (SKY);
Uchibanare-jima - 2-'? -,?, 3-4 viii 1988 (Y. Fujii); Hateruma-jima - 10 J' J'1-'?, 1-2 vii 1988 (SKY), 2.Sf -'?, Naishi, 2
x 1988 (SI); Yonaguni-jima -1-'?, Hikawa, 26 viii 1978 (TN), 1 J', Sonai, 5 vii 1988 (SKY).
Distribution. Miyako Is. (Miyako-jima; Irabu-jima); Tarama Is. (Tarama-jima; Minnajima); Yaeyama Is. (lshigaki-jima; Taketomi-jima; Kuro-shima; Iriomote-jima; Uchibanare68
jima; Hatoma-jima; Hateruma-jima; Yonaguni-jima).
Biology. Kinjo (in Azuma, 1987) states that in the Ryukyus this species nests in
hollowed plant stems, abandoned mud nests of other wasps or tunnels dug in plant
material (referred to as Anterhynchium fIavomargilUltum).
Parasite: Pseudoxenos iwatai (Maeta, 1980; Kifune & Yamane, 1985).
Genus Anterhynchium Saussure
Anterhynchium Saussure, 1863, Mem. Soc. Phys. Hist. Nat. Geneve, 17: 205 (name for Rygchium or
Rhynchium, I Division of Saussure, 1852, 1855)(type species: Rygchium synagroides Saussure, 1852, designated
by Vecht, 1%3); Vecht, 1963, Zool. Verh., 60: 58 (in key), 73, 74; Vecht and Fischer, 1971, Hym. Cat. (nov. ed.),
8: 105.
Japanese name: Futaobi-dorobachi Zoku.
A good diagnosis for this genus was given by Vecht (1963), who divided it into three
subgenera, Dirhynchium Vecht, Anterhynchium Saussure, and Epiodynerus Giordani Soika.
The two Japanese species <flavomarginatum and melanopterum) belong to Dirhynchium,
which is characterized by the dense transverse striation on the narrow basal part of gastral
sternite 1 and very coarse punctation on the basal part of gastral tergites 3 to 5 (the latter
condition can be seen when the gaster is unusually extended). In northeastern Asia three
very closely related species are known to occur, for which a key will be given below.
Key to the species of Anterhynchium in northeastern Asia
1. DepreSSion bearing cephalic foveae obscure ( 5f.). Aedeagal shaft only slightly narrower
than aedeagal tip (t). Oypeus wider than high ( 5f. ). Scutellum and metanotum almost
always without orange markings ( 5f. t) . ............................................. A. melanopterum Yam.
- Cephalic foveae situated in a large, posteriorly well defined depression ( 5f.). Aedeagal
shaft much thinner (t). Shape of clypeus variable. Scutellum and/or metanotum often
marked with yellow or orange .................................................................................................. 2
2. Clypeus nearly as wide as high ( 5f. ); its apical emargination rather wide and deep, and
lateral teeth divergent ( 5f. t) . ............................................................. A. flavopunctatum (Sm.)
- Clypeus higher than wide (5f.); its apical emargination narrower and shallower (5f. t) .
................................................................................................................ A. flavomargilUltum (Sm.)
Anterhynchium fIavomarginatum (Smith)
(Figs. 120,122,124,126-142,144-146)
Rhynchium fiavo-margiMtum Smith, 1852, Trans. Entomol. Soc. Lond. (2)2: 35 (ct)(type loc.: China).
Odynerus nigrifrons Smith, 1857, Cat. Hym. Brit. Mus. 5: 62 (~ )(type loc.: N. China).
Anterhynchium flavomarginatum: Vecht, 1963, Zoo!. Verh. 60: 79; Yamane and Tano, 1983, Mem.
Kagoshima Univ. Res. Center S. Pac.,4: 124,125.
Japanese name: O-futaobi-dorabachi.
Diagnosis. Body length (h+th+t1+2) 12.5 mm in 5f., 9.0-12.5 mm in
69
t.
Fore wing
122Q}
sO:
121
\ i\;!
A
f1I
123~
:. .:.*
~.
/
b .~
....
120
125
124
Figs. 120-125. Morphological characters in the Japanese Anterhynchium (after Yamane, 1981). 120,
122,124, A. flavomarginatum; 121, 123, 125, A. me/anopterum. 120,121, dypeus (A, ~; B, 6'); 122,
123, segments 10-13 of male antenna (original); 124, 125, apical part of penis.
length 12.0-14.5 mm in ~, 8.5-11.5 mm in t. This is a polytypic species comprising 13
subspecies and widely distributed in southern and eastern Asia. Up to now eight
subspecies have been described and recorded from Japan (Yamane, 1981; Yamane & Tano,
1983). Most of them, known from the Ryflkyfl islands, have a more or less wide range of
variation in color pattern. Although in some cases the subspecies may be recognized only
statistically, this treatment is useful in discussing the faunal relations among islands.
The ground color of this species is black, sometimes partly replaced with brown or
reddish brown. Body markings vary from sulpher yellow through yellow and orange
yellow to reddish brown in color. Generally the markings on the head and alitrunk are
paler than those on the gaster. Main markings are explained below to facilitate the
description of the geographical races.
1. Clypeus: ground color much reduced in ~ and almost lost in t ; in ~ the black
portions will be called black clypeal markings for convenience (in tsushimarum only basal
part pale yellow); the black markings (sometimes replaced with brown) are generally
developed in hanedai, amamense and insulicola, much reduced in umenoi and procella, but
vary in development even in a single population.
2. Mandibular stripe: only seen in t of umenoi.
3. Frontal marking: situated just above the antennal insertion, rarely connected with
the supradypeal marking, when the latter is present.
4. Supradypeal marking: usually very small and often lost.
5. Stripe on each ocular sinus along the lower inner margin of eye: always present in
70
6' except in tsushimarum, but variable in size; in
~ only seen in umenoi.
6. Stripe on antennal scape below: seen in all the forms, and variable in color.
7. Genal band (umenOl), genal stripe or spot (other forms): usually upper genal stripe
alone is seen; lower spot is seen only in ~ of procella.
8. Pronotal marking: large and variable in size.
9. Pair of median stripes on mesoscutum: sometimes weakly developed, but usually
absent.
10. Lateral stripe on meso scutum along the base of each tegula: usually undeveloped,
most strongly developed in ~ of micado and procella.
11. Marking on tegula: variable in shape.
12. Parategula: usually wholly yellow, in a few forms the yellow extending forward
as the lateral stripe of mesoscutum.
13. Mesopleural spot under each wing base: usually present, but often lost in ~ of
hanedai.
14. Marking on scutellum: always seen, sometimes divided into two.
15. Stripe on metanotum (postscutellum).
16. Propodeal spots: variable in size and shape, and completely lost in some forms.
17. Apical bands on gastral tergites 1-5 (or 1-6 in 6'): bands on first two tergites are
always complete except in tsushimarum in which bands are nearly wholly lost, but those
on the subsequent tergites may be replaced with brown or dark reddish brown, or lost.
18. Spots on stemite 2 (also 3 in 6'), and apical bands on stemites 3-5 (or -6 in 6'):
these markings are usually replaced with brown or lost.
19. Lateral spots on male mid and hind coxae: in some forms completely lost; other
parts of legs in 6' are also marked with yellow, but their position and size are highly
variable even in a single population.
Key to the Japanese subspecies of A. flavomarginatum
1. Body almost entirely black. Yellowish color confined to the basal half of c\ypeus
(cIypeus entirely yellow in 6'), and to frontal spot (in a few specimens pronotum and
gaster with minute pale markings). Wings blackish. Tsushima ........................................... .
........................................................................................................ subsp. tsushimarum (Yasum.)
- Body marked with yellow or orange; at least gastral tergites 1 and 2 with the complete.
apical bands .................................................................................................................................. 2
2. Ocular sinus with yellow stripe ( ~ ). Mandible with yellow stripe ( 6'). Antennal scape
almost entirely yellow (6'). Genal band developed ( ~ 6'). Yaeyama Is ............................... .
.......................................................... ........................................................ subsp. umenoi (Yasum.)
- Ocular sinus without yellow stripe ( ~ ). Mandible without yellow stripe ( 6'). Antennal
scape brownish or blackish above (6'). Gena with upper stripe (~ 6').............................. 3
3. Antennal flagellum brownish or ferruginous even on upper face (especially in ~;
darker in 6'). Mesopleural spot often lost ( ~). Propodeum without yellow markings
(~; 6' often with such markings). Body markings darker, partly brownish (~;
somewhat paler in 6'). Okinawa Is ......................................................... subsp. hanedai Tano
- Antennal flagellum black above ( ~ 6'). Mesopleural spot present ( ~ ). Propodeum often
with yellow markings dorsolaterally ( ~ 6'). Body markings yellow or orange yellow
(~; paler in 6') ........................................................................................................................... 4
71
4. Only tergites 1 and 2 with distinct yellow apical bands; other tergites completely black
or with dark or blackish brown bands which are inconspicuous ( !f. 6'). Last tergite and
sternite almost entirely black ( !f. ). Antennal pedicel usually black above ( 6'). Hokkaido,
Honshu, Awaji-shima, Shikoku, KyfIshu, etc ..................................... subsp. micado (Kirsch)
- Apical bands on gastral tergites 3-5 (or -6 in 6') also yellow or orange yellow; (orange)
yellow sometimes replaced with brown or reddish brown, but the bands usually clearly
discernible ( !f. 6'). Last tergite and sternite sometimes ferruginous, at least partly ( !f. ),
Antennal pedicel often ferruginous above (6') ..................................................................... 5
5. Markings on head and thorax sulpher yellow ( !f. 6'). Yellow bands on tergites usually
complete ( !f. 6'); that on tergite 1 distinctly widened laterally, often with lateral prongs
anteriorly (!f. 6'). Akuseki-jima; Takara-jima ..................................... subsp. sulphreum Yam.
- Markings on head and thorax yellow or orange yellow (at most pale yellow in 6').
Apical bands on tergites 3-5 (-6 in 6') often brownish or ferruginous (!f. 6') .................... 6
6. Last tergite almost entirely ferruginous ( !f. ). Orange yellow apical bands on tergites 3-5
usually complete (!f.). Antennal pedicel above usually black (6'). Mid and hind coxae
usually without yellow lateral spots (6'). Kuchi-no-shima; Naka-no-shima; Suwanosejima ............................................................................................................ subsp. insulicoLl Yam.
- Last tergite black, at least partly (!f.). Apical bands on tergites 3-5 often replaced with
brown or reddish brown (!f.). Antennal pedicel above often ferruginous (6'). Mid and
hind coxae often with yellow lateral spots (6') ...................................................................... 7
7. Yellow apical bands on tergite 1 wider, widened laterally, often with lateral prongs
anteriorly (!f. 6'). Gena without lower yellow spot (!f.). Yellow lateral stripes on
mesoscutum much reduced or lost ( !f. d'). Black clypeal markings often developed ( !f.).
Amami-6shima, Okinoerabu-jima, etc ................................................ subsp. arruzmense Tano
- Yellow apical band on tergite 1 narrower, narrowed laterally (!f. d'). Gena often with
lower yellow or orange spot in addition to the upper stripe (!f.; not seen in 6').
Mesoscutum with lateral yellow stripes (usually in !f., sometimes in 6'). Black clypeal
markings much reduced ( !f. ). Mi-shima Is.; Vji Is ................................ subsp. procelLl Yam.
Anterhynchium [lavorruzrginatum micado (Kirsch)
(Figs. 133, 142, 144, 145, 148)
Rhynchium ardens Smith, 1873, Trans. R. Entomol. Soc. Lond. 1873: 1% (-'f )(type loc.: Nagasaki,
KyUshU)(primary homonym of Rh. ardens Walker, 1871).
Odynerus (Leionotus) micado Kirsch, 1878, Mitt. Zool. Mus. Dresden, 3: 380 (-'f )(type loc.: Japan).
Rhygchium (!) micado: Yasumatsu, 1950, Icon. Ins. Jpn. (2nd ed.), p. 1455, fig. 4197.
Rhynchium japonicum Dalla Torre, 1894, Cat. Hym. 9: 46 (new name for Rh. ardens Smith); Yano, 1932,
Icon. Ins. Jpn. (lst ed.), p. 306, fig. 594; Yasumatsu, 1938, Ins. Jpn. m. Icon. Col. Nat. Dep., p. 360, pI. 161, fig.
631-632.
Rhynchium mandarineum Sauss.: Matsumura, 1911, Thous. Ins. Jpn. Supple 3, p. 113, no. 685
(mantlarinum i); 1930,
Thous. Ins. Jpn. 2: 15, pI. 2, fig. 17; 1931, 6000 Ill. Ins. Jpn. p. 16, no. 81.
Rhynchium varipes Perez, 1905, Bull. Mus. Hist. Nat. Paris, 11: 25, 85 (-'f )(type loc.: Yokohama, HonshU).
Anterhynchium f/a"Domarginatum micado: Vecht, 1963, Zool. Verh. 60: 77 (in key), 80; Ishikawa, 1%5, Icon.
Ins. Jpn. Col. Nat. Ed. 3: 297, pI. 149, fig. 6; Yamane, 1981, Trans. Shikoku Entomol. Soc. 15: 225 (in key);
Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S. Pac., 4: 126 (in key), 131, figs. 21, 30.
m.
Japanese names: O-futaobi-dorobachi (Futo-futaobi-dorobachi).
Diagnosis Female. Body markings are yellow or orange yellow. Yellowish are: clypeus
72
with the apical margin brown and the lateral and lower margins narrowly black, frontal
spot, stripe behind eye, antennal scape below, dorsal part of pronotum except posterior
1/3, median stripes on mesoscutum (rarely), parategula, lateral stripes on mesoscutum
(rarely complete), anterior and inner margins of tegula, scutellum largely, mesopleural
spot, metanotum largely, irregular marking(s) on each side of dorsal part of propodeum,
apical bands on gastral tergites 1 and 2 (rarely also on tergite 3), marking on each posterolateral corner of sternite 2. Apical margins of tergites 3-5 usually black or dark
ferruginous; only very rarely tergite 3 with a yellowish apical band. Antenna black; scape
above and flagellum below brownish. Mandible ferruginous. All the legs ferruginous to
dark brown. Wings brownish hyaline.
Male. Very similar to the female in coloration, but differs from the latter in the
following details: c1ypeus without black rims; ocular sinus usually with a short yellow
stripe; yellow markings on scutellum and metanotum often reduced to a varying extent
(completely lost in one specimen from Kagoshima); outer faces of fore and mid femora
and tibiae with yellow apical spots (these spots sometimes very large, and in such cases
femur and tibia of hind leg also extensively yellowish).
Material examined. HonshU: Yamagata-ken - 1 ~, Nan'yo~hi, 20 vii 1980; Niigata-ken - 1 d', 5enami, 30 vi
1979 (KB), 26' d', same loc., 20 vi 1981 (KB), 11 d' d'4~ ~,same loc., 16-23 vii 1981 (KB), 2d' d', same loc., 28
viii 1981 (KB), 1 d'5~~, same loc., 3-11 ix 1981 (KB), 1~, same loc., 25 vii 1983 (KB), 1 d', same loc., 30 vi
1984 (KB), 1 ~,same loc., 26 ix 1984 (KB), 1 d', Kurokawa, 27 vii 1979 (KB), 1~, same loc., 25 ix 1981 (KB),1
~, same loc., 18 x 1984 (KB), 1 ~, Shibata, 12 viii 1979 (HI), 3 d' d', Suibara, 30 v - 3 vi 1978 (A. Seino), 1 ~ ,
Nagaoka, 29 viii 1952 (H. Kozaki); Nagano-ken -1 d', Habiro, Ina, 14 viii 1962 (YM), 1 d', Amori, Nagano-shi,
24 viii 1979 (T. Fujisawa); Tochigi-ken - 1 ~, Shimozuke, Otawara, 10 viii 1949 (Usuba); Chiba-ken - 1 d'1 ~,
Yokoto,3 ix 1981 (J. Tsukahara), 1 ~,Shirako, Oami, 31 vii 1983 (M. Ohara); Saitama-ken-l d'1 ~,Ageo, 31 vii
1980 (T. Sunose), 4d' d', same loc., 23 vii 1983 (T. Sunose); Ibaraki-ken - 2d' d', Tsuchiura, 29 viii 1976 (SKY);
Gifu-ken - 1 d', Hong5-chO, Seki, 19 viii 1976 (Y. Takai), 1 d'1!f, same loc., 31 vii - 5 viii 1982 (Y. Takai), 1 d',
Horado, 7 viii 1982 (Y. Takai); Wakayama-ken - 3 ~ ~, Koza, 25 ix 1974 (5. Takagi); Ky6to-fu - 3 ~ ~, Gosho, 12
vii 1955 (K. Iwata); Hy6go-ken - 1 ~, Sasayama, Tamba, 31 vii 1953 (K. Iwata), 17 d' d', same loc., 2~ vi 1964
(H. Katayama), 1 d', same loc., 5 vii 1967 (K. Iwata), 1 ~, same loc., 19 x 1966 (H. Katayama); Awaji-shima - 3
~ ~, 18-19 viii 1954 (T. Morl); Osaka-fu - 1 6', Iwawaki, Kawachi, 6 vi 1964 (H. Katayama); Shimane-ken - 1 d',
Mt. Dake, Matsue, 15 vii 1985 (N. Sugiura), 1 ~, Mt. Makuragi, 28 viii 1985 (M. Goubara).
Figs. 126-133. Color pattern of female clypeus in the Ryilkyil subspecies of Anterhynchium
flavomarginatum (after Yamane & Tano, 1983). 126, ssp. umenoi; 127, 128, ssp. hanedai; 129, ssp.
amamense; 130, ssp. sulphreum; 131, ssp. insulicola; 132, ssp. procel/a; 133, ssp. micado.
73
Shikoku: KOch i-ken -1 d', Asakura, Kiichi-shi, 5 vi 1974 (51), I5/-, Okoyama, Nankoku, 17 ix 1976 (51);
KagaUIQ-ken -15/-, Yashima, 14 viii 1976 (KB).
Kyushu: Fukudca-ken - 25/- 5/-, Kurogi, 27-28 vii 1983 (Y. Takai); Kumamoto-ken -1 d', Kagami, 31 v 1982
(M. Maeda), 15/-, same loc., 17 vii 1982 (M. Maeda), 15/-, same loc., 30 vii 1982 (M. Maeda); Nagasaki-ken -15/-,
Haraguchi, Omura, 17 ix 1967 (R. Ohgushi); Oita-ien -1 d', Saeki, 23 vii 1954 (K. Iwata); Miyazaki-ken -I5/-,
Ornata, 15 vii 1954 (K. Nohara), Id', Uchiumi, 20 vii 1954 (K. Yoshida), I5/-, Maruno, 21 vii 1954 (5.
Kawamura); Kagoshima-ien - 7 d' d'15/-, Taguchi, Kirishima,3 vii 1981 (SKY), I5/-, JingO, Kirishima, 23 vii
1981 (SKY), 1 d', Shiroyama, Kagoshima-shi, 5 vi 1981 (K. Sakamoto), 15d' d'55/- 5/-, same loc., 4-10 vii 1981
(SKY), 4 d' d'25/- 5/-, same loc., 30 vii 1981 (SKY), 15/-, same loc., 6 ix 1982 (SKY), 23 d' d'45/-!f, Korimoto,
Kagoshima-shi, 4-7 viii 1981 (SKY), 15/-, same loc., 13 vii 1981 (M. Maeda), 5 d' d'15/-, same loc., 17 viii 1981
(SKY), 4 d' d'15/-, same loc., 22-26 viii 1981 (SKY), 7 d' d'25/- 5/-, Meiwa, Kagoshima-shi, 15 viii 1981 (SKY), 25/5/-, Sakamoto, Kagoshima-shi, 20-24 ix 1981 (5. Higashi), 15/-, same loc., viii 1982 (5. Higashi), 15/-, Kekura,
Kagoshima-shi,2O ix 1981 (SKY), 15/-, nr Yamakawa, 6 viii 1984 (M. Maegata), 15/-, Kaimon-dake, 26 viii 1986
(SKY), I5/-, Takakuma (800 m alt.), 31 vii 1983 (KD, 2d' d'25/- 5/-, Koyama, 24 v 1979 (emerged from a nest; H.
Nagase), 1 d', Sata-<ho, 10 vi 1979 (H. Nagase).
Goto Is. (Nagasaki-ken): Alaa-shima - 25/- 5/-,17 viii 1976 (J. Nakayama).
Islands close to Kagoshima-ken-hondo: Naga-shima - 2 d' d', Shoura, 27 viii 1984 (SKY); Take-shima - 3 d'
d', 28 viii 1984 (SKY); Kamikoshiki-jima - 25/- 5/-, Sa to, 5-6 ix 1984 (M. Maegata); Shimokoshiki-jima - 5d' d',
Kashima,5 viii 1986 (51).
N. Ryukyus: Tane-ga-shima - 15/-, Ikeno, 21 vii 1984 (5. Watahiki), 15/-, Urata, 24 vii 1984 (5. Watahikj), 1
5/-, Hamada, 2 viii 1984 (SKY), 15/-, Kamome, 5 viii 1986 (M. Tatsuno); Mage-shima - 1 d', 22 vii 1984 (5.
Watahiki); Yaku-shima - 1!f, Kurio, 2 xi 1975 (F. Komaj), 4 d' J'45/- 5/-, Miyanoura, 8-11 viii 1981 (SKY), 1 d',
same loc., 28 vi 1982 (51), 15/-, same loc., 24 viii 1982 (51), 2 J' d'15/-, Kusugawa, 9 viii 1981 (SKY), 1 d'15/-,
Onoaida, 27-29 vii 1982 (51); Kuchinoerabu-jima- 25/- 5/-, Hommura,20 vii 1989 (SKY).
Distribution. Hokkaido (5. & c. part, Endou, 1977); Honshu; Sado-ga-shima; Awajishima; Izu Is. (To-shima); Shikoku; Kyushu; Goto Is. (Aka-shima); Chikuzen-okino-shima;
Naga-shima Is. (Naga-shima; Take-shima); Koshiki-jima Is. (Kami- & Shimo-koshiki-jima);
Osumi Is. (Tane-ga-shima; Mage-shima; Yaku-shima; Kuchinoerabu-jima).
Taxonomic notes. According to Smith's description, Rhynchium ardens (=Rh. japonicum
D.T.) has a yellow fascia on gastral tergite 3. Vecht (1%3) mentioned that japonicum may
therefore prove to be sub specifically different from micado. Since specimens with an apical
band on tergite 3 do occur (though rarely) in the population of KyfIshu, I consider such
specimens to represent variations within the subspecies micado. Vecht (1963) recorded a
female of "typical" micado from Korea, but Korean specimens (subsp. koreanum Sk.
Yamane) differs from micado by the darker mandible, much reduced pro notal markings,
wholly black scutellum, etc. (Yamane, 1981). Many specimens collected on Yaku-shima are
intermediates between the subspecies micado and procel/a, and a few are typical procel/a.
Since Yaku-shima is connected with Kyushu by liners, there may be some possibility that
micado-type individuals have invaded from Kyushu and mixed with indigenous
individuals belonging to procel/a.
Biology. This form is one of the most common eumenid wasps in Japan. Its biology
has been studied by Iwata (1938b, Rhygchium japonicum), Tsuneki (1970) and ltino (1986a,
b). The female wasps construct their nests in bamboo and reed tubes, in tunnels in wood
bored by coleopterans, and sometimes in old mud cells of other wasps such as Oreumenes
decoratus, Orancistrocerus drewseni and Scelifron madraspatanum (Iwata, 1980c; Onuma,
1989c). A nest comprises 1-9 brood and some empty cells (Iwata, 1938b). Itino (1986a)
reported an average of 2.2 cells per nest (241 trap nests were examined) in Kyoto. One
female may make several nests. The cells are separated with mud partitions and each cell
is mass-provisioned with 3-21 (mean: ca. 8) larvae of Microlepidoptera (Pyralidae and
Tortricidae) as prey. Maternal care for young is generally not observed. Univoltine in
74
Figs. 134-142. Color pattern of female gaster in the Ryfikyil subspecies of Anterhynchium
flavomarginatum. 134, ssp. umenoi; 135, ssp. hanedai; 136-138, ssp. amamense; 139, ssp. sulphreum;
140, ssp. insulicola; 141, ssp. procella; 142, ssp. micado.
Kyoto (ltino, 1986b), but the very long flying period (early June to mid-October) suggests
the occurrence of two generations per year in at least southern parts of Japan.
Parasite: Pseudoxenos iwatai (Strepsiptera, Stylopidae). Parasitoids: Macrosiagon nasuta
(Coleoptera, Rhipiphoridae), Megaselia sp. (Diptera, Phoridae), Amobia distorta (Dipt.,
Sarcophagidae), a chrysidid wasp, Acroricnus ambulator, Campoplex sp. (Hym., Ichneumonidae)(ltino, 1986a), Nematopodius sp. (Hym., Ichneumonidae), and Melittobia acasta
(Hym., Eulophidae) (Yamane & Maeta, unpub!.).
Anterhynchium flavomarginatum tsushimarum (Yasumatsu)
(Fig. 144)
Rhynchium flavopunctatum Smith forma tsushimarum Yasumatsu, 1935, Mushi 8: 86 (!f )(type loc.:
Shimo-agata, Tsushima Is.).
Anterhynchium (Dirhynchium) flavomarginatum tsushimarum: Vecht, 1963, Zoo!. Verh. 60: 71 (in key);
Ishikawa, 1970, Mem. Nat. Sci. Mus. Tokyo 3: 269-271, pI. 22; Yamane, 1981, Trans. Shikoku Entomol. Soc. 15:
223-224 (in key).
Diagnosis. Female. Body almost wholly black. The following parts are yellow: large
basal mark of cIypeus, frontal spot, a pair of short lines on the horizontal part of pronotum
anteriorly (often lost), medially widely interrupted apical bands on gastral tergites 1 and 2
(these bands are very narrow and often lost). Mandibles, antennal scape below, femora of
all legs apically, tibiae and tarsi of all legs partly dark ferruginous. Mandibles and tibiae
often wholly black. Wings blackish hyaline.
Male. Very similar to the female in coloration. Clypeus almost wholly and antennal
scape below yellow. In one specimen the yellow pronotal marking and apical bands on
tergites 1 and 2 are rather developed.
75
Material examined. Tsushima Is. (Nagasaki-ken): 1!?-, Sumo, 9 x 1959 (YM); Kami-<lgata - 1!?-, Uchiyama
(140 m alt.), 1 (J', Oboshi-yama, 24 vii 1979 (A. Seino); Shimo-agata - 1!?-, Kashi - Shiradake, 22 vii 1978 (K.
Katsura), lex, Izuhara, 11 viii 1978 (K. Haruyama), 2(J' (J'2!?-!?-, Tsutsu, 22-23 viii 1979 (I. Hiura).
Distribution. Tsushima Is. (Kami-agata; Shimo-agata).
Taxonomic notes. I have examined one female specimen collected in Hakozaki,
Fukuoka-shi, KyUshu (19 vii 1959, Y. Murakami leg.; parasitized by a female Pseudoxenos),
with a color pattern very similar to the present subspecies. In this specimen the mandibles
are much paler in color, and the basal marking on the clypeus is very small. This wasp
may have been introduced from Tsushima to Kyushii as suggested by Ishikawa (1970).
Yasumatsu (1935a) mentioned that on the Tsushima islands there occurs also the
typical form of A. flavopunctatum with a yellow fascia on two basal tergites of gaster each. I
have not examined such specimens from Tsushima. The form mentioned by Yasumatsu
seems most probably A. melanopterum, which is sympatric with A. flavomarginatum
tsushimarum.
According to Ishikawa (1970) the Korean population of this species belongs to A. f.
micado, which is widely distributed in the Japanese mainlands, but, in my view, it is a
distinct subspecies closely related to the nominate subspecies occurring in China (Yamane,
1981).
Anterhynchium flavomarginatum procella Sk. Yamane
(Figs. 132, 141, 148)
Anterhynchium flavomarginatum procellll: Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S.
Pac. 4: 131, figs. 20, 29 {!?- (J'){type loc.: Kuro-shima, Osumi Is.).
Diagnosis. This subspecies is characterized by the following combination of character
states. Black clypeal markings much reduced (-'f-); gena often with a lower yellow (or
reddish) spot or line in addition to the upper one ( -'f- ); mesoscutum with a yellow stripe
on each side along the whole length of the base of tegula (usually in -'f-, sometimes in (f);
propodeum with yellow markings on the dorsolateral face (-'f- (f); yellow apical band on
gastral tergite 1 narrow, laterally narrowed (-'f- (f); gastral tergites 3-5 usually with yellow
apical bands which are sometimes replaced with reddish brown (-'f- (f); last tergite and
sternite black (-'f-); antennal pedicel above often tinged with reddish brown «(f).
Material examined. N. Ryukyus: Take-shima - 28!?- !?-, 29-30 viii 1983 (KT); Io-jima - 5 (J' (J'2!?- !?-, 1-3 vi
1982 (SKY), 3 (J' (J', 7 viii 1982 (KT); Kur~hima - 36 (J' (J'32!?- -\'- (incl. holotype and paratypes), Osato, 30 viii 5 ix 1981 (SKY); Ie-jima -1!?-, 6-9 x 1981 (SKY); Mukai-jima -l-\'-, 9 x 1983 (M. Ohara).
Distribution. Mi-shima Is. (Take-shima; I6-jima; Kuro-shima); Uji Is. (le-jima; Mukaijima).
Taxonomic notes. This form is closely allied to the subsp. micado. The lower genal spot
(line) in the female is peculiar to this form, but is not seen in all the specimens. The gastral
tergite 3 almost always possesses a yellow apical band (sometimes interrupted medially),
but this band is also seen in micado though rarely. Many specimens collected on Yakushima are intermediates between these two forms as discussed above. Although one male
and one female specimen collected on August 4 1916 from Tane-ga-shima are rather
procella-typed (see Yamane & Tano, 1983), recently collected specimens all belong to the
micado type (also see Ogata & Nagase, 1987).
The following specimens from the Ogasawara Is. are more or less procella-typed. All
76
of them have the gastral tergite 3 with a rather distinct apical band that is yellowish at
least partly. One female from Chichi-jima has a trace of reddish lower genal line on each
side. It is not known whether the Ogasawara population of this species is native or has
recently been introduced.
Ogasawara (Bonin) Is.: Orichi-jima -1 J'1Sf, 16-17 viii 1972 (YH), 2Sf Sf, Futago, 9 viii 1983 (TN); Hahajima -1 J', Kitakoo,11 viii 1983 (TN), 1Sf, Okikoo, 12 viii 1983 (TN).
Biology. Parasite: Pseudoxenos iwatai (Kifune & Yamane, 1985).
Anterhynchium flavomarginatum insulicola Sk. Yamane
(Figs. 131, 140, 148)
Anterhynchium fiavomarginatum insulicola: Yamane and Tano, 1983, Mem. Kagoshima Univ. Rec. Center
S. Pac. 4: 130, figs. 19, 28 (Sf J') (type loc.: Kuchi-no-shima, Tokara Is.).
Ditlgnosis. Similar to the subsp. amamense from Amami-oshima and Okinoerabu-jima.
Blackish and brownish markings on c1ypeus rather developed (.'f-). Antennal flagellum
below blackish (.'f-); antennal pedicel above black (d'). Apical bands on gastral tergites 1-5
(6) usually orange yellow, rarely replaced with reddish brown ( .'f- d'); the band on tergite 1
relatively narrow, not distinctly widened laterally (.'f- d'); last tergite and sternite
ferruginous (.'f-). Mid and/or hind coxae wholly black (d'). In the population of
Suwanose-jima, yellow propodeal markings well developed.
Material examined. N. Ryukyus : Kuchi-no-5hima - 4 J' J'3 Sf Sf, 28 vii - 1 viii 1981 (KT), 39 J' J'8Sf Sf (ind.
holotype and paratypes), 24-27 vi 1982 (KT); Naka-no-5hima - 1Sf, 21 viii 1943, 1 J'8 Sf Sf, Satomura, 13-19 vii
1982 (Y. Takaj), 2 Sf Sf, Funakura, 14 vii 1982 (Y. Takai); Suwanose-jima - 8 J' J'9 Sf Sf, 30 vii -1 viii 1985 (S1).
Distribution. Tokara Is. (Kuchi-no-shima; Naka-no-shima; Suwanose-jima).
Biology. Parasite: Pseudoxenos iwatai (Kifune & Yamane, 1985).
Anterhynchium flavomarginatum sulphreum Sk. Yamane
(Figs. 130, 139, 148)
Anterhynchium fiavomarginatum sulphreum: Yamane and Tano, 1983, Mem. Kagoshima Univ. Rec. Center
S. Pac. 4: 129-130, figs. 18,27 (Sf J')(type loc.: Akuseki-jima, Tokara Is.).
Ditlgnosis. This subspecies is unique in its sulpher yellow body markings, especially
on head and thorax (apical bands on gastral tergites often slightly tinged with orange).
The following character states are also useful in separating it from the other forms. Black
clypea1 markings moderately developed (.'f- ); frons sometimes with a suprac1ypea1 yellow
spot (d'); antennal flagellum below often ferruginous (.'f-); yellow markings on scutellum,
metanotum and dorsal face of propodeum well developed ( .'f- d'); gastral tergites 1 and 2
with apical bands distinctly widened laterally (that on tergite 1 often with lateral prongs
anteriorly)(.'f- d'); apical bands on tergites 3-5(6) usually not replaced with brown (.'f- d').
Material examined. N. Ryukyus: Akuseki-jima -12J' J'8Sf Sf (ind. holotype and paratypes), 7-10 viii 1981
(KT),3SfSf,25vii 1982 (M.Ohara); 1J'5SfSf, 3-8 vii 1984 (KT),6J'J',2 viii 1985 (SI),7Sf Sf,18-19viii 1987
(T. Moriyama).
C. Ryukyus: Takara-jima - 1Sf , 29 vii 1986 (SI).
Distribution. Tokara Is. (Akuseki-jima; Takara-jima).
Biology. Parasite: Pseudoxenos iwatai (Kifune & Yamane, 1985).
77
tc~)
•
00 Izu Is.
o
Q.
145
Ogasawara
Is.
.~
u
.~.
. .. ,:~f~=
146
....""
~, .'
emi CBdo
*tsushimBrum
144
Figs. 143-146. Distribution of Anterhynchium melanapterum (143) and A. fozvomarginatum (144-146;
two subspecies) on the Japanese mainlands, Izu Islands and Ogasawara Islands.
Anterhynchium flavomarginatum amamense Tano
(Figs. 129, 136-138, 148)
Anterhynchium flavomarginatum amamense: Yamane and Tano, 1983, Mem. Kagoshima Univ. Rec. Center
S. Pac. 4: 128-129, figs. 17, 24-26 (.'f J')(type loc.: Amami-oshima).
Diagnosis. Female. Body markings are in pattern similar to those of the subspecies
hanedai, but much paler in color. Black (brown) clypeal markings generally developed, but
variable in size and shape. Antennal flagellum above entirely blackish, rarely tinged with
reddish brown, but sometimes brownish below. Mesopleural spots never lost. Yellow
markings on scutellum and metanotum rather developed. Propodeal spots always
present, sometimes much developed. Yellow apical band on tergite 1 sometimes slightly
widened laterally; that on tergite 2 usually distinctly widened laterally; last tergite and
sternite usually blackish, at least partly. Yellow bands on tergites 3-5 very often replaced
with reddish or dark brown. Legs ferruginous; femora usually blackish basally.
Male. Body markings paler than in the female. Similar to the male of the subspecies
hanedai, but differs in the following points: body markings paler (yellow; only rarely
orangish), antennal flagellum blackish (segments 7-9 below and last two segments wholly
ferruginous), propodeum almost always with yellow spots, yellow band on tergite 1 much
wider and often with anterior prongs on each side, apical bands on tergites 3-6 very often
reduced or replaced with brown.
Material examined. C. Ryukyus: Amami-ilshima - 14J' J'4.'f.'f, Shimmura, 16 vii -1 viii 1967 (TM), 1 J'1
.'f, Gusuku, 21 vii 1%7 (TM); 1 J', Santaro Pass, 23 vii 1967 (TM), 2.'f .'f, Kasari, 17-21 vii 1%7 (TM), 1.'f,
Sato, 14 vii 1967 (TM), 1.'f, Asani, 13 vii 1967 (TM), 1.'f, Tomori, 16 vii 1%7 (TM), 1.'f, Uken, 28 vii 1972 (TN)
(holotype), 1 J', Yuwan, 27 vii 1972 (TN), 1.'f, Naze, 27 vii 1972 (TN), 1 J', Yuwan, 6 vii 1980 (H. Nagase), 1
J', Eboshi-dake, 6 vii 1980 (H. Nagase), 1 J'4.'f.'f, Naze, 10-18 vii 1981 (yH), 1 J'2.'f.'f, Kasari, 7 vii 1981
(yH), 1 J', Setouchi, 16 vii 1981 (YH), 1.'f, Sumiyo, 13 vi 1981 (YH), 2 J' J', Akaogi, 30 vi 1984 (M. Ohara), 1
J', Naze, 20 vi 1987 (SKY), 1 J', same loc., 10 vii 1987 (AN), 33' 3', Sumiyo, 22 vi 1987 (SKY), 1 J'1.'f, Koniya,
78
22 vi 1987 (SKY), 1 ~, Setouchi, 23 vi 1987 (M. Tatsuno), 3 ~ ~, Nishinakama, 25 vi 1987 (SKY), 1 6'1 ~, Naze,
14 vii 1988 (AN), 1 ~, Nishinakama, 15 vii 1988 (AN); Kakeroma-jima - 2~ ~, 23-24 ix 1984 (AN); Yoro-shima 2C!' C!'2~~, 13 viii 1987 (M. Tatsuno); Tokuno-shima - 2C!' C!'3~~, Kametoku, 13 vii 1984 (SKY); Okinoerabujima - 1 C!', Tokutoki, 4 viii 1972 (TN), 1 ~, Uchigusuku, 3 viii 1972 (TN), 2 ~ ~, China, 19 v 1981 (KT,
Kukidome& Koya), lC!'1~,Shinjo-Oyama, 15-16 vii 1981 (Y. Takai), lC!'2~~, 17-18 vii 1984(M. Maegata).
Distribution. Amami Is. (Amami-oshima; Kakeroma-jima; Yoro-shima; Tokunoshima; Okinoerabu-jima). Kikai-jima seems to lack this species.
Taxonomic notes. The color pattern of the specimens from Okinoerabu-jima is rather
variable. Although most of them fall into the range of variation seen on Amami-oshima,
the two females from China and the male from Shinjo - Oyama agree well with the subsp.
micado in many details.
Biology. Parasite: Pseudoxenos iwatai from Amami-Oshima and Tokuno-shima.
Population of Anterhynchium flavomarginatum from Yoron-to
Yoron-to is a small flat island situated between Okinoerabu-jima and Okinawa-jima.
All the specimens examined from this island, collected between 1983 and 1986, cannot be
separated from A. f. micado occurring in the Japanese mainlands. Yellow apical bands are
restricted to gastral tergites 1 and 2. Tergite 3 rarely possesses a rufous band, but usually
tergites 3-6(7) are wholly black. It is possible, therefore, that the individuals from Yoron-to
and some from Okinoerabu-jima are descendants of immigrants from Kyushu (subsp.
micado), which invaded through human activities (Yamane & Tano, 1983).
Material examined. 1 C!'1 ~, Chabana, 6 viii 1972 (TN), 1 ~, 22 v 1983 (E. Matsui), 1 ~, Gusuku, 22 v
1983 (M. Ohara), 4 C!' C!'9~ ~, 4-6 vi 1985 (SKY), 6 C!' C!' 5~ ~, 24-28 v 1986 (SKY).
Biology. In early June, 1985, a female wasp was observed nesting in a horizontal
tunnel in a wooden pillar of a cabin in Akasaki.
Parasite: Pseudo xenos iwatai (new record for this island).
Anterhynchium flavomarginatum hanedai Tano
(Figs. 127,128,135,148)
Anterhynchium flavomarginatum hanedai: Yamane and Tano, 1983, Mem. Kagoshima Univ. Rec. Center S.
Pac. 4: 128, figs. 16, 23 (~ C!') (type loc.: Okinawa-jima); Giordani Soika, 1986, Boll. Mus. Civ. St. Nat. Venez.
35: 73.
Anterhynchium flavomarginatum luctuosum Giordani Soika, 1986, Boll. Mus. Civ. St. Nat. Venez. 35: 74
(~)(type loc.: Sezoko-jima, Okinawa Is.). Syn. nov.
Rhynchium mandarineum Saussure: Matsumura and Uchida, 1926, Ins. Matsum. 1: 37-38.
Anterhynchium flavomarginatum umenoi: Yamane, 1981, Trans. Shikoku Entomol. Soc. 15: 224-225 (in key,
part).
Japanese name: Okinawa-futaobi-dorobachi.
Diagnosis. Female. Body markings yellow, orange yellow and brown. Clypeus
yellow, with lateral black or brown markings which are variable in size and shape. Frontal
spot yellow, tinged with brown. Upper genal stripe very small and orange yellow.
Depression for cephalic foveae on vertex with dense reddish brown pubescence. Antennal
scape brownish above, yellowish below; flagellum blackish brown to reddish brown even
on the upper face. Horizontal face of pronotum yellow (sometimes orangish or
ferruginous); its lower and posterior portions black. Mesopleural spot yellow to
79
ferruginous, often lost. Scutellum and metanotum with orangish or ferruginous markings
which are variable in size (often reduced). Propodeum always without yellowish
markings. Apical bands on gastral tergites 1-5 usually dark orange yellow (those on
tergites 3-5 rarely ferruginous). Last tergite light brown to ferruginous. Sternite 2 with an
orange yellow apical band which is widely interrupted medially; subsequent sternites
often with dark ferruginous, narrow, apical bands; last sternite ferruginous. Legs almost
entirely ferruginous, but coxae, and sometimes trochanters and femora basally blackish;
without yellow spots.
Male. Similar to the female in color pattern, but body markings paler. Antennal scape
brown or blackish above, yellow below; flagellum ferruginous (sometimes much darker
above); hook slightly yellowish. Propodeum sometimes with yellow spots. Orange yellow
apical band on tergite 1 sometimes widened laterally, rarely with lateral prongs anteriorly;
last tergite usually blackish. All the apical bands on tergites yellowish (not ferruginous).
Sternites 2-6 with yellow or orange apical bands (often interrupted medially). Legs light
brown to ferruginous, extensively marked with yellow.
Material examined. C. Ryukyus: OkintrWa-jima - 1 J'1 Sf, "Okinawa" (5. 5akaguchi), 1 J', "Okinawa", viii
1905 (Kuroiwa), 1 Sf, Shun, 11 vii 1968 (S. Hashimoto); 1 J', Naha, 9 viii 1972 (TN), 4J' J'1 Sf, Nakagusuku, 3
x 1977 (5Y), 1 J', Ogumi-son, 26 viii 1979 (H. Nagase), 1 J', Kunigami, 3-4 viii 1980 (KB), 1 J', Hyakuna, 1 vii
1981 (AN), 2 J' J', Yona, 3 vii 1981 (AN), 2J' J', Hentona, 4 vii 1981 (AN), 1 Sf, Oku, 5 vii 1981 (AN), 12 J' J'5
Sf Sf (inc!. ho!otype), Tobaru, Kunigami, 3-9 vii 1982 (YH), 3Sf Sf, Yabu, Nago, 30 vi 1982 (YH), 1 Sf,
NagojOshi,29 vi 1982 (YH), 4 J' J', Okuma, Kunigami, 6-7 vii 1982 (yH), 1 J', Beshi, Motobu, 1 vii 1982 (yH),
1 J'1 Sf, Yona, 2 vi 1983 (AN), 17 J' J'3Sf Sf, Hentona, 1-6 vi 1983 (AN), 1 J', Yona, 18 vii 1984 (SKY); Sezokojima - 3 Sf Sf, 28 v -1 vi 1982 (Y. Ikimon & Y. Ohira); Hamahiga-jima - , 1 Sf, 22 x 1988 (Y. Kusui).
Distribution. Okinawa Is. (Okinawa-jima; Sezoko-jima; Hamahiga-jima; Kume-jima).
Taxonomic notes. This subspecies is characterized by the ferruginous flagellum
(especially of the female), and the darkest body markings in the female among the
RyiikyCl forms. However, there are a few specimens with lighter body markings and they
are indistinguishable from the subsp. amamense. Tano (1987) mentioned that female
specimens from Kume-jima often had a brownish marking on vertex posteriorly, and that
in the male markings on scutellum and metanotum tended to be much reduced.
Biology. Parasite: Pseudoxenos iwatai from Okinawa-jima and Sezoko-jima (Maeta,
1980; Kifune & Yamane, 1985).
Anterhynchium flavomarginatum umenoi (Yasumatsu)
(Figs. 126, 134, 148)
Rhynchium umenoi Yasumatsu, 1933, Annot. Zoo!. Jpn. 14: 262-265,271, fig. 3 (Sf- )(type loc.: Ishigakijima, Yaeyama Is.).
Anterhynchium jlavomarginatum umenoi: Vecht, 1963, Zoo!. Verh. 60: 81; Yamane, 1981, Trans. Shikoku
Entomo!. Soc. 15: 224-225 (in key); Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S. Pac. 4: 126128, figs. 14, 22.
Japanese name: Yaeyama-futaobi-dorobachi.
Diagnosis. Female. The specimens examined well agree with the original description
by Yasumatsu (1933). Body markings are yellow, but sometimes orangish; the type
specimen which was said to have orange yellow markings may represent an extreme
variation. Pronotum is black in its posterior portion of the horizontal face. Mesoscutum
sometimes possesses very faint median stripes. Propodeum almost always lacks yellow
80
markings.
Male. Yellow markings rich; the markings paler than in the female, and only rarely
orangish. Mandible with a yellow stripe. Supraclypeal spot present, often connected with
frontal mark. Antennal scape entirely yellow; pedicel brownish; flagellar segment 1 below
and some of the terminal segments below ferruginous. Gena yellow except for posterior
portion. Tegula yellow except for central ferruginous part. Apical bands on gastral tergites
1-6 rather wide (last tergite entirely brownish but sometimes apically yellowish); apical
bands on sternites 2-6 narrow and yellow (last stemite light brown). Tergites 5 and 6, and
sternites 5 and 6 sometimes brownish basally. Legs light brown, with yellow markings on
femora and tibiae.
Material examined. S. Ryukyus: Miyako-jima - 1~, Hirara, 28 vi 1988 (SKY); Ishigaki-jima - 26'6',
Nagurahama,31 vii 1969 (TI), 26' 6', Takeda, 13 viii 1972 (TN), 1!f, Omoto-dake, 30 viii 1978 (1<. Hara), 1~,
Yonehara,28 viii 1978 (TN), 1~, Banna-dake, 29 viii 1978 (TN), 36'6', Kabira,22 vii 1987 (SKY), 1 6'1 ~
Shiraho, 26 vii 1987 (SKY), 1!f, Kabira, 20 v 1988 (K. Nakamine), 16', Omoto-dake, 11 vi 1988 (K.
Nakamine), 76' 6'3 ~ ~, same loc., 3-8 vii 1988 (1<. Nakamine), 56' 6'2!f !f, Banna-dake, 3-8 vii 1988 (K.
Nakamine), 106' t2~!f, same loc., 4 vii 1988 (SKY). 16'. same loc.,3 x 1988 (51); Taketomi-jima- 16'.25 vii
1987 (SKY). 16'.8 vii 1988 (1<. Nakamine); Kuro-shima - 8 6' 6'1 !f. 23-24 vii 1987 (SKY); Kohama-jima - 36'6'1
-'f. 25 vii 1987 (SKY); lriol1UJte-jima - 66' 6'4-'f -'f. Funaura, 9 x 1977 (SY), 1-'f, Hoshidate, 8 x 1977 (SY), 56' 6'
6!f !f, Komi, 11-24 viii 1978 (TN), 2!f -'f, Kanebire-taki, 23 viii 1978 (TN), 1-'f, Funaura, 22 viii 1978 (TN), 1
-'f, Ohara, 19 viii 1978 (TN), 1 6'2!f -'f, same loc., 20-24 v 1981 (AN), 26' 6'1-'f, Toyohara, 20 v 1981 (AN),l
6', Sonai, 20 iv 1981 (T. Fujisawa), 1-'f, Ohara, 2 v 1982 (AN), 1 6', Otomi, 29 vii 1983 (AN), 1-'f ,Ohara,29 vii
•
0
'.
V~
!}
•• 5
~.~
•
8
148
Figs. 147, 148. Distribution of Rhynchium quinquecinctum (147; two subspecies) and Anterhynchium
flavomarginatum (148) in the Ryukyus. 1, ssp. micado; 2, ssp. procel/a; 3, ssp. insulicola; 4, ssp.
sulphreum; 5, ssp. amamense; 6, population of Yoron-to; 7, ssp. hanedai; 8, ssp. umenoi.
81
1983 (AN),2d'd', Toyohara, 30 vii 1983 (AN), 4d'd'3'f- 'f-, Komi, 29-31 vii 1983 (AN), 2d'd'1'f-, Otomi,25
vii - 3 viii 1985 (AN), 3'f- 'f-, Amitori, 30 vii - 1 viii 1985 (AN), 1 'f-, Toyohara, 26 vii 1985 (AN), 2 d' d'1 'f-,
same loc., 11 x 1987 (AN), 2 d' d', Ohara, 13 x 1987 (AN), 1 d'2'f- 'f-, Komi, 3 vii 1988 (SKY), 1 d', Uehara, 2 vii
1988 (SKY), 1 'f-, Mihara, 4 xii 1988 (SKY); Hateruma-jima - 3d' d', 1 vii 1988 (SKY); 1 d'1 'f-, Naishi, 2 x 1988
(51).
Distribution. Miyako Is. (Miyako-jima); Yaeyama Is. (lshigaki-jima; Taketomi-jima;
Kuro-shima; Kohama-jima; Iriomote-jima; Hatoma-jima; Hateruma-jima; Yonaguni-jima
after Tano, 1985).
Taxonomic notes. The male of this form is readily separated from that of any other
form of this species in Japan by the entirely yellow scape, well developed yellow genal
band, and the yellow stripe on mandible. We could not find any remarkable difference in
color pattern among the populations of the islands. However, the female specimen
(parasitized by a female Pseudo xenos) from Miyako-jima is very peculiar in having
antennae nearly wholly orange yellow, and the depression (in which lie the cephalic
foveae) on vertex orange yellow. It is unceratin whether this color pattern is normal in the
population of the island. Vecht (1963) mentioned that the Taiwanese form well agreed
with Yasumatsu's description of this form, but in fact these two are not alike at all.
Biology. On December 41988, on Iriomote-jima, a female wasp was observed nesting
in a bamboo tube that was used as part of a chair at a bus stop:
Parasite: Pseudoxenos iwatai from Miyako-jima (new record), Ishigaki-jima (new
record), and lriomote-jima (Kifune & Yamane, 1985).
Anterhynchium melanopterum Sk. Yamane
(Figs. 121, 123, 125, 143)
Anterhynchium me/anopterum Yamane, 1981, Trans. Shikoku Entomol. Soc. 15: 221, 222, figs. 1, 3 ('f- d')
(type loc.: Kagoshima, Kyilshil).
Japanese name: Haguro-futaobi-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 12.5-16.0 mm. Head with strong and
dense punctures that are somewhat reticulate. Hairy depression (with cephalic foveae) on
the vertex obscure or almost invisible (the depression is distinct in A. flavomarginatum).
Clypeus slightly wider than long, strongly punctate, only narrowly and shallowly
emarginate at apex; apical teeth blunt. Keel between the antennae sharp. Hairs on frons
and ocellar region not very uniform in length and shape. Thorax strongly and densely
punctate or reticulate, with weak transverse striae on the anterolateral part of pronotum;
anterior fourth of mesepisternum and lower half of dorsal metapleuron almost
impunctate. Posterior edge of metanotum spinose. Propodeum with the posterior face
distinctly striate; upper projections distinct and sharp behind the metanotum;
posterolateral angles developed. Below metanotum a large triangular impunctate area,
between the lateral halves of the posterior face of propodeum. Lateral sides of propodeum
transversely striate, partly reticulate. Gastral tergites 1 and 2 finely and somewhat densely
punctate; punctures on tergites 3-5 slightly larger than those on the preceding ones,
especially near the apex. Narrow basal part of sternite 1 finely striate; posterior triangular
part with rather coarse striation. Sternites 2-5 punctate as in tergites; last sternite nearly
impunctate.
Black, with orange yellow markings. Orange yellow are: a large basal triangular
82
marking on clypeus, a transverse spot between antennae, scape below, a small spot behind
each eye, a transverse band on pronotum anteriorly (medially interrupted), apical bands
on gastral tergites 1 and 2. Tarsi of all legs somewhat brownish. Wings infuscated, darker
than in A. flavomarginatum micado (blackish vs. brownish).
Male. Body length (h+th+t1+2): 9.5-10.5 mm. Similar to the female. Clypeus entirely
yellow, longer than wide; clypeal ratio (alb; Fig. 121): ca. 2.9 (in flilvomarginatum 3.13-3.44);
apex more widely emarginate than in the female. Hairs on head and thorax disheveled,
long; those on clypeus much shorter. Aedeagus of male genitalia with a weak subapical
constriction; shaft thicker than in A. flilvomarginatum.
Material examined. Honshu: Niigata-ken - 1.'f, Nagaoka, 9 vii 1976 (SKY); TOkyi3-to - 1.'f, Akabane, 23 ix
1937 (N. Kumazawa).
Shikoku: K6chi-ken - 1.'f, Kochi, 13 ix 1956 0. Minamikawa), 2//, Saigyoji, 2 viii 19590.
Minamikawa), 1.'f, Godaisan, Kochi-shi, 1 x 1975 (SI); Kagawa-ken - 1/, Kotohira, 22 viii 19590.
Minamikawa).
Kyushu: Fukuoka-ken - 1.'f, Koshii, 29 ix 1959 (YM)(gaster lost), 1.'f, Hakozaki, 7 vii 1959 (Y. Miyatake)
(gaster lost); Miyamki-ken -1.'f. Ornata, 15 vii 1954 (K. Nohara); Kag06hima-ken -1.'f (holotype), Kagoshimashi, 14 ix 1980 (H. Nagase).
Tsushima Is. (Nagasaki-ken): 1/, Sumo, 23 vii 1978 (K. Tani); ShimCHgata - 1.'f, Tatsura-yama, 15 x
1979 (T. Kumata), 1 /, izuhara, 27 vii 1986 (K Nakamine).
Distribution. Honshu; Shikoku; Kyushu;Tsushima Is.
Taxonomic notes. This species is closely related to A. flavomarginatum, but distinguished from the latter by the wider clypeus (5?- t), obscure depression for cephalic
foveae on vertex ( 5?-), and the thicker aedeagal shaft (t). In color pattern this species is
quite similar to Orancistrocerus drewseni drewseni, while easily distinguished from A.
flavomarginatum micado, the latter two forms being widely sympatric with this species on
the mainlands of Japan. Another unnamed subspecies of A. melanopterum occurs in
northeastern China (I have examined some specimens deposited in the Zoological
Museum, Leiden). One male specimen from Tsushima has an exceptionally short clypeus,
condition being comparable to that in A. flavopunctatum of the continental Asia. I
tentatively consider this condition to be included within the range of variation in
melanopterum.
Biology. Nothing is known about the nesting behavior of this species.
Parasite: Pseudo xenos iwatai (Strepsiptera, Stylopidae) (new host record) from
Akabane, Tokyo.
Genus Okinawepipona Sk. Yamane
Okinawepipona Yamane, 1987, Mem. Kagoshima Univ. Res. Center S. Pac. 8: 52-53 (type species:
Anterhynchium (Epiodynerus) kogimai Giordani Soika, monotypic).
Japanese name: Okinawa-dorobachi Zoku.
Diagnosis. Clypeus moderately convex, as high as wide, narrowly emarginate at apex
in both sexes. Male mandible not deeply emarginate on inner side. Maxillary palp 5segmented; labial palp 3-segmented (Fig. 8). Scutellum flat; metanotum medially not
concave. Tegula not extending beyond the apex of parategula (Fig. 150). Propodeum
without lateral crest. Anterior vertical face of gastral tergite 1 almost impunctate, clearly
separable from the posterior horizontal part, but not by a carina; laterally the tergite
83
divided by a sharp carina into upper and lower part. Gastral segment 1 slightly narrower
than segment 2. Tergites 1-5 each with a dark apical spot medially where the integument is
weakly impressed. Narrow basal part of sternite 1 irregularly striate posteriorly, not
shining. Basal part of sternite 2 with 14-18 strong, longitudinal carinae (Fig. 152).
Parastigma of fore wing short, less than half as long as the stigma.
This genus was first mentioned in the key to the Japanese genera of Eumenidae by
Yamane (1982)(referred to as "Okinawa-dorobachi-zoku"). Although there are few
autapomorphies, the genus is distinguished from the related genera by the characteristics
mentioned above, especially by the reduced numbers of segments of maxillary and labial
pal pi and short para stigma. In the latter condition, this genus is similar to Pararrhynchium
rather than to Anterhynchium. Thus Giordani Soika's (1986) assignment of his new species
kogimai to the subgenus Epiodynerus of the genus Anterhynchium is not supported.
Okinawepipona kogimai (Giordani Soika)
(Figs. 8, 149-162, 182)
Anterhynchium (Epiodynerus) kogimai Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 74-76 (!f
J')(type loc.: Sezoko-jima, Okinawa Is.).
Okinawepipona kogimai: Yamane, 1987, Mem. Kagoshima Univ. Res. Center S. Pac. 8: 53-54, figs. 1-4.
Japanese name: Okinawa-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 11.0-12.5 mm. Fore wing length: 11.012.0 mm. Head subcircular, densely punctate. Supraclypeal area with a vertical carina
between the antennal sockets; above the carina interantennal region with a narrow vertical
groove. Occipital carina complete, but dorsally somewhat weak. Ocellar triangle flat;
distance between the posterior ocelli slightly longer than or as long as the distance
between posterior ocellus and eye. Depression for cephalic foveae not conspicuous. Gena
moderately developed, in profile narrower than the eye. Mandible pointed at apex; apical
part often heavily worn; inner edge with four round teeth. Thorax somewhat elongate,
densely punctate. Pronotal carina almost complete, only slightly interrupted dorsally.
Anterior vertical part of pronotum virtually without punctures. Posterior 1/3 of
mesoscutum and whole scutellum nearly flat. Dorsal mesepisternum below impunctate.
Punctation sparse on metapleuron. Metanotum slightly convex. Propodeum rather
developed; basal triangular area with a median groove, at the lower end of which a
median carina starts; dorsolateral face of propodeum with rugose punctation; posterior
face with striae; lateral face above with punctures and below with weak striae. Punctation
on gastral tergites much weaker than on thorax and propodeum except on the dorsal face
of tergite 1 where the punctures are larger and dense but shallower than on thorax.
Punctation on gastral sternites 2-6 still finer than on corresponding tergites. Hind coxa
with a distinct carina, which is elevated to form a triangular process (Fig. 151).
Male. Body length (t+th+t1+2): 9.0-10.0 mm. Fore wing length: 9.0-10.5 mm. Similar to
the female. Apical emargination of clypeus deeper than in the female. Antennal segments
12 and 13 very small (Fig. 149).
This species is peculiar to the Central Ryukyus (subspecies nagasei & kogimai) and
Taiwan (subsp. taiwana) (Figs. 153-162). No information is available regarding the nesting
biology of this species.
84
Okinawepipona kogimai nagasei Sk. Yamane
(Figs. 153, 154, 158,159, 182)
Okinawepipona kogimlli nagasei Yamane, 1987, Mem. Kagoshima Univ. Res. Center S. Pac. 8: 54-55, figs. 5,
6, 10, 11 (6'!f){type loc.: Amami-ashima).
Diagnosis. Female. Black with yellow and orange yellow markings as follows: clypeus
wholly (with brownish lower rim), mandible except for apex and inner teeth, frontal
marking which is connected by a narrow line to the yellow of clypeus, a vertical line
below anterior ocellus, a stripe along inner edge of eye from the upper margin of clypeus
to the top of eye, gena along outer edge of eye, antennal scape below, posterior horizontal
portion of pronotum extensively, a large spot under wing base, tegula (with a central spot
and margins brownish), a pair of short lines on mesoscutum, a line on mesoscutum along
the inner edge of tegula, parategula, scutellum and metanotum largely, apical bands on
gastral tergites 1-5 (widest on tergite 2 and very narrow on tergites 3-5), an apical band on
gastral sternite 2. Scape above and pedicel brown; flagellum blackish brown above and
orange below. Legs testaceous; all the coxae black; basal 1/3 of fore and mid femora, and
ventral face of hind femur dark.
Male. In coloration very similar to the female, but yellow marking along inner edge of
eye not reaching the top of eye. Frons without yellow line just below anterior ocellus.
Genal band reduced to a short line. Gastral tergites 6 and 7 wholly black. Legs yellow.
Material examined. C. Ryukyus: Takara-jima - 1!f, 2-11 viii 1985 (M. Kawana); Amami-Oshimll - 2!f!f,
Nishinakama, 25-26 vi 1961 (K. Tsuneki); 2!f !f, Gusuku, 1 vii 1961 (K. Tsuneki); 1!f, Akaogi, 7 vii 1961 (K.
Tsuneki); 1!f, Kachiura, 3 vii 1961 (K. Tsuneki); 1!f, Santaro Pass, 23 vii 1967 (TM); 1!f, Shimmura, 1 viii
1969 (TM); 16' (holotype), Kasari, 20 v 1979 (H. Nagase), 1 6'1!f, Yuwan-dake, 20 vi 1983 (K. NichO), 2!f !f,
Sumiyo, 22 vi 1987 (SKY), 1 6', Shinokawa, 23 vi 1987 (SKY).
Distribution. Tokara Is. (Takara-jima); Amami Is. (Amami-Oshima).
Figs. 149-152. Okinawepipona kogimai (after Yamane, 1987). 149, terminal segments of male antenna;
150, tegula and parategula (PT); 151, hind coxa, showing posterior process (PP); 152, gastral
stemites 1 and 2 (BS2, basal part of stemite 2).
85
Figs. 153-157. Facial color pattern in the three subspecies of Okinawepipona kogimai (after Yamane,
1987). 153, subsp. nagasei 6'; 154, ditto !iC; 155, subsp. kogimai 6'; 156, ditto !iC; 157, subsp.
taiwana 6'.
Okinawepipona kogirnai kogirnai (Giordani Soika)
(Figs. 155, 156, 160, 161, 182)
Okinawepipona kogimai kogimai: Yamane, 1987, Mem. Kagoshima Univ. Res. Center S. Pac. 8: 55-56, figs.
7,8,12,13.
Diagnosis. Female. Similar to the preceding subspecies, but body markings much
darker, especially on frons, vertex, thorax and gastral tergites 1 and 2. Head extensively
orange or rufous; a narrow space around antennal sockets, a U-shaped area on vertex
including ocellar region, and markings on gena posteriorly and temple black. Propodeum
occasionally with reddish spots. Apical band on gastral tergite 2 wide; basal black area
being less than 1/3 as wide as the total length of the tergite (in subsp. nagasei apical bands
always less than half as wide as the length of the tergite). Fore and mid femora almost
wholly testaceous.
Male. Orange and yellowish markings much less developed than in the female.
Pattern of the body markings similar to.that in the subsp. nagasei except that markings are
much darker in this form.
Material examined. C. Ryukyus: Okinawa-jima - 1!iC, Shuri, 6 vi 1970 (M. Kinjo); 16', 15 v 1973 (M.
Kinjo); 16', Kudeken, 15 v 1973 (M. Kinjo); l!iC, Nago, 4-5 v 1976 (H. Takizawa); 1!iC, Motobu, 29 v 1981 (K.
Kojima); 2!iC !iC, Yona, 17 vii 1984 (SKY); Sezoko-jima -1!iC, 9 v 1982 (J. Kojima)(paratype), l!iC, 28 v -1 vi 1982
(Y. Ikimori & Y. Ohira).
Distribution. Okinawa Is. (Okinawa-jima; Sezoko-jima).
86
Genus Pararrhynchium Saussure
Pararrnynchium Saussure, 1855, Et. Fam. Vesp. vol. 3, p. 173 (division of genus Rhynchium Spinola)(type
species: Rhynchium ornatum Smith, 1852, monotypic); Dalla Torre, 1894, Cat. Hym. vol. 9, p. 42 (Parrhynchium
!); Vecht, 1963, Zool. Verh. 60: 58 (in key), 94; Vecht and Fischer, 1972, Hym. Cat. pars 8, p. 107; Carpenter,
1986, Psyche 93: 79.
Prorhynchium Saussure, 1855, Et. Fam. Vesp. vol. 3, p. 174 (division of genus Rhynchium Spinola)(type
species: Rhynchium smithii Saussure, 1855, monotypic).
Japanese names: Kabafu-dorobachi Zoku (Kabafusuji-dorobachi Zoku).
This genus is characterized by the following characteristics (Vecht, 1963). Labrum of
female with very short hairs, at most with a few longer hairs at its anterior margin. Apical
margin of clypeus wide, more or less emarginate. Concavity of propodeum margined by a
crest, which is incised dorsally in the middle, and is separated from the metanotum by a
short horizontal area (propodeal shelf). Gastral tergite 1 with a transverse ridge separating
the anterior vertical face from the posterior horizontal part. Parastigma short, distinctly
less than half as long as the stigma. Cubital cell 3 separated from the apex of the radial cell
by a long distance, much more than half the width of the latter cell.
Recurrent vein 2 of fore wing attaches to the cubital vein at the point relatively close
to intercubital vein 2, and in the Japanese population of P. ornatum the former is very close
to the latter or these two are even directly connected. These conditions disagree with
Vecht's view that in this genus the recurrent vein 2 is never very close to intercubital vein.
The females of all the Japanese species have a long, medially flattened clypeus which is
widely truncate and shallowly emarginate at apex. P. oceanicum, described below as a
new species, is unusual in that both the epicnemial carina and propodeal shelf are
completely lost and body sculpture is very superficial.
Figs. 158-162. Body color pattern in the three subspecies of Okinawepipona kogimai (after Yamane,
1987). 158, subsp. nagasei .!f.; 159, ditto (1'; 160, subsp. kogimai .!f.; 161, ditto (1'; 162, subsp.
laiwana (1'.
87
Key to the Japanese spcies
1. Propodeum without shelf; crest undeveloped. Epicnemial carina absent. Male
unknown ..................................................................................................... P. oceanicum sp. nov.
- Propodeum with a distinct shelf between the crest and metanotum. Epicnemial carina
present ........................................................................................................................................... 2
2. Body markings yellowish. Head and alitrunk extensively marked with yellow. Female
clypeus with the apical margin lamellate. Occipital carina in profile somewhat strongly
bent at 3/5 from the top of head. Metanotum without distinct longitudinal furrow. Basal
lobe of aedeagus relatively thick, distinctly shorter than aedeagal apodeme; each lobe of
ventral process of aedeagus not strongly curved ............................. P. ishigakiense (Yasum.)
- Body markings orange. Head and alitrunk almost wholly black or with a few markings.
Apical margin of female clypeus not lamellate, black. Occipital carina in profile evenly
arched so that the gena gradually narrowed below. Metanotum various .......................... 3
3. Female clypeus above orange. Frontal spot present. Female antennal scape below
orange. Flattened part of female c1ypeus less sharply defined, not shining, with sparse
medium-sized punctures. Metanotum usually with a deep median furrow. Mesoscutum
nearly as long as wide. Basal lobe of aedeagus slender, as long as or only slightly shorter
than aedegal apodeme; each lobe of the ventral process of aedeagus strongly curved .
............................................................................ ............................... P. ornatum ornatum (Smith)
- Female c1ypeus entirely black. Frontal spot absent or much reduced. Female antennal
scape below ferruginous. Alitrunk entirely black. Gastral tergites 3-6 wholly black.
Flattened part of c1ypeus well defined, shining, with sparse fine punctures. Median
furrow of metanotum indistinct. Mesoscutum longer than wide (50:43). Male unknown .
.................................................................................................................. P. tsunekii Tano et Yam.
Pararrhynchium ishigakiense (Yasumatsu)
(Figs. 163, 165, 168, 170, 173, 182)
Ancislrocerus ishigakiensis Yasumatsu, 1933, Annot. Zoo!. Jpn. 14: 260-262,271, figs. 1,2 (~)(type loc.:
Ishigaki-jima, Yaeyama Is.); 1935, Ibid. 15: 38, figs. IB, C (J').
Pararmyncium ishigakiense: Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center s. Pac. 4: 120,
figs. 1,3,6,9.
Japanese name: Ishigaki-kabafu-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 11.5-12.0 mm. Fore wing length: 11.012.5 mm. Head nearly circular, strongly punctate. Cephalic foveae small, situated colse to
each other, the area being slightly elevated. Ocellar triangle flat. Gena, seen from above,
slightly wider than eye. Interantennal keel distinct. Clypeus higher than wide, sparsely
punctate, apically broadly truncate (emargination quite shallow); median flattened
portion relatively well defined; apical margin lamellate (Fig. 165). Occipital carina in
profile somewhat strongly bent at 3/5 from the top of head (Fig. 163). Pronotum very
densely and strongly punctate, somewhat reticulate. Pretegular carina developed.
Epicnemium with a row of large punctures along the anterior margin, with much smaller
punctures near the sterno pleural groove. Other parts of mesopleuron densely and
strongly punctate. Mesoscutum strongly punctate, somewhat reticulate; prescutal grooves
88
not conspicuous. Scutellum punctate, with a narrow median groove; just in front of
metanotum there is a wide transverse groove with many short carinae. Metanotum
weakly punctate. Metapleuron and lateral side of propodeum punctate and often also
striate; punctures much smaller than on propodeum. Dorsal face of propodeum strongly
and densely punctate, somewhat reticulate; propodeal concavity much wider than high,
dull, with obliquely running indistinct carinae; inferior median carina about half the
height of concavity; superior carinae apically forming a pair of relatively sharp teeth
behind metanotum (Fig. 170). Upper half of the anterior vertical face of gastral tergite 1
punctate, with a median longitudinal carina; transverse ridge not sharp. Gastral segment 1
distinctly narrower than segment 2. Gastral tergites finely punctate; tergite 2 lamellate
apically; tergites 1-5 each with a preapical drop-shaped dark spot. Sternite 2 finely
punctate; punctation on other sternites much weaker. Para stigma less than half the
pterostigma.
Black, with the following parts yellow: c1ypeus wholly, a wide band along the inner
margin of eye from c1ypeus beyond the upper margin of eye, a heart-shaped frontal mark,
a rhombic marking above it, gena wholly, postgena, a triangle basal marking on mandible
(other parts of mandible yellowish brown), antennal scape (pedicel and basal part of
flagellar segment 1 orange), anterior vertical portion of pronotum above, dorsal part of
pronotum largely, a U-shaped marking on mesoscutum, dorsal mesepisternum, a stripe on
ventral mesepisternum (often lost in the populations of Iriomote-jima and Yonaguni-jima),
tegula except for central and peripheral transparent parts, parategula (often wholly black),
scutellum, metanotum, wide apical bands on tergites 1-5, a pair of large spots on tergite 2
which are laterally connected with the wide apical band, narrower apical bands on
sternites 2-5. Segment 6 wholly yellow or ocherous, tinged with brown. All the legs
extensively yellowish; mid and hind coxae and femora often brownish). Wings yellowinfumated.
Male. Body length (h+th+t1+2): 9.0-11.0 mm. Fore wing length: 7.5-9.5 mm. Very
similar to the female in stucture and coloration. Clypeus approximately as high as wide;
apical emargination deeper than in the female; lateral angles sharp; median flattened area
ill defined. Apex of antenna! hook reaching the apical margin of segment 10 (Fig. 168).
Area behind the ocelli less elevated and strongly punctate. Gena less developed, in profile
much narrower than eye. Facial yellow markings much reduced, never connected with the
yellow parts of inner orbits. Mesoscutal marking also much reduced, never forming a U.
Material examined. S. Ryukyus: Ishigaki-jima - 76' 6'25f 5f, Ishigaki-5hi, 25 iii -1 iv 1977 (J. Nakayama),
1 6'15f, same loc., 18-19 iv 1981 (T. Fujisawa), 15f, Banna-dake, 4 vii 1988 (SKY); Iriomote-jima - 1!f,
Shirahama, 18 viii 1971 (5. Yamauchi), 16', Funaura, 9 x 1977 (SY), 1 6'45f 5f, Otomi, 15-19 v 1981 (AN), 16',
Toyohara, 26 vii 1985 (AN); Yonaguni-jima -1 6'15f, 24-25 iv 1988 (yM), 3 6' 6'2!f !f, Sonai, 5 vii 1988 (SKY).
Distribution. S. Ryukyus (lshigaki-jima; Iriomote-jima; Yonaguni-jima).
Biology. No information is available.
Pararrhynchium ornatum ornatum (Smith)
(Figs. 118, 164, 167, 169, 172, 174, 182)
Rhynchium ornatum Smith, 1852, Trans. Entomol. Soc. Lond. (2)2: 36 (5f )(type loc.: Tein-tung, China);
Yano, 1932, Icon. Ins. Jpn. p. 308, f. 598.
Odynerus ornatus: Schulthess, 1934, Arb. Morph. Tax. Entomol. 1: 70 (in key); Yasumatsu, 1938, Ins. Jpn.
Dl. Icon. p. 359, f. 628-1 (subgenus Ancistrocerus).
89
Odynerus shinto Schulthess, 1908, Mitt. Schweiz. Entomol. Ges. 11: 286 (5f )(type loc.: Yokohama,
HonshO).
Ancistrocerus ornatus: Yasumalsu, 1950, Icon. Ins. Jpn. 2nd Ed. p. 1457, f. 4204.
Pararrhynchium ornatum ornatum: Vecht, 1963, Zool. Verh. 60: 95; Yamane and Tano, 1983, Mem.
Kagoshima Univ. Res. Center S. Pac. 4: 121-122, ff. 2, 5, 8, 10.
Paranhynchium ornatum: Ishikawa, 1965, Icon. Ins. Jpn. Col.Nat. Ed. 3: 297, pI. 149, f. 2.
Japanese names: Nami-kabafu-dorobachi (kabafu-suji-dorobachi).
Diagnosis. Female. Body length (h+th+t1+2): 12.0-14.0 mm. Fore wing length: 11.012.0 mm. Head slightly wider than high, strongly punctate. Gena, seen from above, as
wide as eye. Oypeus slightly higher than wide, relatively finely and sparsely punctate;
median flattened portion ill defined (Fig. 167). Interantennal carina blunt. Cephalic foveae
very small, situated in a shallow depression in a slightly elevated part which bears dense
pubescence. Occipital carina not strongly bent so that gena gradually narrowed toward
the mandibular base (Fig. 164). Anterior vertical face of pronotum shining, laterally with
dense punctures and finely punctate near upper margin. Dorsal part of pronotum strongly
punctate; pretegular carina developed. Mesoscutum densely and strongly punctate;
median area with sparse punctation; prescutal furrows distinct. Scutellum strongly
punctate, with a median longitudinal furrow, anteriorly with a transverse furrow which
has 8-9 short carinae. Epicnemial carina complete; epicnemium with dense punctures
below, with a row of larger punctures along anterior margin. Mesepisternum and
mesepimeron rugosely punctate; on dorsal mesepisternum spaces between punctures
forming irregular carinae. Metanotum punctate, depressed posteriorly; anterior margin
with a median notch. Metapleuron superficially striate, and relatively finely punctate.
Propodeum with large punctures on dorsal and lateral portions; spaces between
punctures often running as irregular carinae; propodeal concavity with the inferior
median carina that is short and only 1/3 as long as the height of concavity; crest
developed, with apical teeth that are less pointed than in P. ishigakiense (Fig. 172).
Punctation on gaster very similar to that in P. ishigakiense. Recurrent vein 2 of fore wing
often very close to or even connected with intercubital vein 3.
Black, with the following parts orange yellow or orange: transverse frontal marking,
upper half of clypeus, small triangular marking at the base of mandible, antennal scape
below, a pair of markings in the dorsal part of pronotum anteriorly (these extend slightly
into the anterior vertical face of pronotum), posterior horizontal part of gastral tergite 1
(medially deeply incised), a wide apical band on tergite 2 (often widely incised in the
middle), narrow apical bands on tergites 3 and 4 (sometimes reduced or lost) and very
rarely on tergite 5, stripe on the anterior face of fore tibia.
In the material from Kyushu and Tane-ga-shima, the dypeus is often largely orange.
Male. Clypeus nearly as high as wide, widely emarginate at apex, without defined
flattened area. Tergites 2 and 3 with small lateral yellow markings at apex. Tibiae of all the
legs usually with yellow markings.
Material eXilmined. HonshO: Miyagi.Jcen - 15f, Tsuchitoi, Sendai, 9 viii 1977 (K. Goukon); Niigata-ken - 1
if, "Echigo", 26 vii 1938 (Nohira), 2d' d', "Echigo", 9-21 viii 1948 (Nohira), 1 d', Fukushima-gata, Toyosaka, 12
viii 1978 (HI), 15f, Shibata, 3 ix 1979 (HI); Tokyo-to - 1 d'25f 5f, Ota-ku, 26 viii 1971 (M. Furukawa); Saitamaken - 2 d' d', Omiya, emerged from a nest on 10 v 1976 (collected by YM in 1975 & reared by SKY); lbaraki.Jcen
- 15f, Tsuchiura, 29 viii 1976 (SKY); Kyoto-fu - 15f, Gosho, 11 vii 1955 (K. Iwata); Osaka-fu - 1 d', Iwawaki,
Kawachi, 6 vi 1964 (H. Katayama); Hyogo-ken - 15f, Ogi-no-sen, Tajima, 15 vii 1963 (H. Yuasa), 25f 5f,
Sasayama, Tamba, 24 viii 1952 (K. Iwata), 1 d', same loc., 2 vi 1964 (H. Katayama); Okayama-ken - lif,
Kanagawa, 9 vi 1960 (K. Iwata); Shimane-ken - 15f, Kawatsu, Matsue, 16 vi 1985 (N. Sugiura), 15f, HonjO,
90
164
~""""165
169
"
(1
11
12
166
167
:
0
o{;'
LJ
'8 "'A
171
Figs. 163-174. Morphological characters in the Japanese species of Pararrhynchium (168,169,
original; others after Yamane & Tano, 1983). Head in profile of ishigaJciense (163) and ornatum
(164); female cIypeus Oaterallobes omitted) of ishigakiense (165), tsunekii (166) and ornatum
(167); terminal segments of male antenna of ishigakiense (168) and ornatum (169); propodeum
(posterior view) of ishigakiense (170), tsunekii (171) and ornatum (172); aedeagus of male
genitalia (dorsal view) of ishigakiense (173) and ornatum (174)(A, apodeme; B, basal lobe; C,
ventral view of ventral process).
91
Malsue,24 vii 1986 (N. Sugiura).
Awaji-shima: 2.!f .!f, 18 viii 1954 (T. Mori).
Shikoku: Ehime-ken - 1.!f, Matsuyama, 10 viii 1969 (HI); Tokushima-ken - 1.!f, Mt. Tsudayama,
Tokushima-shi,28 vii 1964 (H. Iwasaki); K6chi-ken - 1 d', Asakura, K6chi-shi, 5 vi 1974 (51).
Kyushu: Fukuoka-ken - 1.!f, Kurogi, 1 viii 1983 (Y. Takai); Nagasaki-ken - 1.!f, Haraguchi, Omura, 4 ix
1966 (R. Ohgushi), 1.!f, same loc., 18 viii 1967 (R. Ohgushi); 1.!f, Kawadana, 6 viii 1976 (J. Nakayama);
Kagoshima-ken - 1.!f, Onejime, Osumi, 3 ix 1978 (H. Nagase), 1.!f, Meiwa, Kagoshima-shi, 6-17 viii 1981
(SKY).
Tsushima Is. (Nagasaki-ken): Shimo-agata -1.!f, Kusudama-jinja, Tsulsu, 23 viii 1979 (I. Hiura).
Goto Is. (Nagasaki-ken): Nakadori-jima - 1.!f, 9 viii 1976 (J. Nakayama).
Island close to Kagoshima-ken-hondo: Akune-i!shima -1.!f, 5 viii 1983 (SKY).
N. Ryukyus: Tane-ga-shima -1.!f, Shimama, 6 viii 1916 (H120), 1 d', Nishi-no-{)mote, 9 viii 1916 (HU7), 1
.!f, 20 vii 1961 (AN). (Ogata & Nagase, 1987, recorded this species from Nakatan~hO of this island based on
material collected between 1982 and 1986.)
Distribution. Honshu (rare in northern part); Awaji-shima; Iwai-jima; Shikoku;
Kyiishu; Tsushima Is. (Shimo-agata); Goto Is. (Nakadori-jima); Akune-6shima; Osumi Is.
(Tane-ga-shima). China.
Biology. This species in Japan constructs its nests in reed or bamboo tubes. The nesting
proceeds very slowly because the female wasp does not close a cell until her larva
becomes large (Iwata, 1938a; referred to as Ancistrocerus ornatus). It took, for example, 23
days to complete a nest finally consisting of five larval cells and one empty cell. Each cell
might have been completed in 4 or 5 days. The female wasp was observed to guard the
cell in which young larva was growing for the greater part of daytime and through night.
By the time when the last cell was closed with mud the wasplings in the first three cells
had finished to spin their cocoons (Iwata, 1938a). Apparently this species provisions
progressively, thus representing a subsocial stage (Iwata, 1971). In total 45 prey caterpillars
were brought into a cell for which a continuous observation was made (Iwata, 1938a).
Prey consists of the larvae of several species of Microlepidoptera.
Parasitoids: a chalcid wasp, and Macrosiagon nasuta (Coleoptera, Rhipiphoridae)
(Iwata, 1938a).
Pararrhynchium tsunekii Tano et Sk. Yamane
(Figs. 166, 171, 182)
Pararrhynchium Isunekii: Yamane and Tano, 1983, Mem. Kagoshima Univ. Res. Center S. Pac., 4: 120-121,
fig. 4, 7 (.!f) (type loc.: Amami-Oshima).
Japanese name: Amami-kabafu-dorobachi.
Female. Similar to P. ornatum ornatum but differs in the following details: clypeus
more deeply emarginate at apex; flattened part of clypeus more clearly defined (Fig. 166)
and more finely punctate; thorax more slender (mesoscutum longer than wide); upper
teeth of superior carina of propodeum rather sharp (Fig. 171); metanotum without distinct
median furrow; punctures on gastral tergites much finer.
Black, with the following parts yellow or orange: basal triangular marking on
mandible, frontal spot (lost in one specimen), very small spot behind eye, vertical face of
gastral tergite 1 largely, other parts of tergite 1 almost wholly, tergite 2 laterally and
apically, two large basal markings on sternite 2, spot at each postero-Iateral angle of
sternite 2. Mandible dark ferruginous. Antennal scape below ferruginous. Tibiae and tarsi
92
tinged with red; apical segments of tarsi brownish.
Male unknown.
Material examined. C. Ryukyus: Amami-ilshima - 1-'?-, Shimmura, 28 vii 1967 (TM)(ho\otype), 1-'?-,
SantarO Pass, 23 vii 1967 (TM)(paratype).
Distribution. Amami Is. (Amami-Oshima).
Biology. No information is available.
Pararrhynchium oceanicum Sk. Yamane, sp. nov.
(Figs. 175-178)
Japanese name: Tsuya-kabafu-dorobachi.
Diagnosis. Female. Head slightly wider than high, rather finely and sparsely punctate;
punctures smaller than inters paces even on the gena. Clypeus higher than wide, almost
impunctate and shining, apically moderately emarginate (Fig. 175). Interantennal keel
blunt; frons shining, with a depression just above the keel. Vertex elevated behind
posterior ocelli; cephalic foveae situated in a depressed region between posterior ocelli
and the elevated part; the depression dull and with a few distinct punctures; the narrow
space between the elevated part and occipital carina irregularly punctate (Fig. 177).
OCcipital carina less developed than in P. ornatum, not strongly bent so that gena
gradually narrowed toward the base of mandible. Postgena very finely and sparsely
punctate. Mandibles each with four blunt teeth and rather sharply pointed apical one.
Maxillary palp 6-segmented; labial palp with 4 segments, of which the terminal one is the
shortest. Anterior vertical face of pronotum shining; punctation on it similar to that in P.
ornatum but finer and sparser. Dorsal and lateral parts of pronotum relatively densely
punctate but never reticulate. Mesoscutum shining, with micropunctures all over the disc
and larger punctures in the anterior half of the disc; prescutal furro,w distinct, measuring
about half the length of the disc. Posterior end of tegula slightly exceeding that of
posttegula. Scutellum very finely and sparsely punctate, shining, with a faint median
longitudinal line, and with a distinct transverse furrow with more than ten short carinae
on the border of mesoscutum. Mesopleuron without epicnemial carina; epicnemium
almost impunctate. Mesepisternum with relatively weak and sparse punctures;
mesepimeron superficially punctate and striate. Metanotum without distinct median
furrow; punctures stronger than on scutellum. Metapleuron almost impunctate, with
feeble carinae in upper portion. Propodeum without shelf; dorsal face relatively strongly
and densely punctate; lateral part and concavity without distinct punctures or carinae,
and dull; crest undeveloped; inferior median carina half as long as the height of concavity
(Fig. 178). Gastral segment 1 narrower than segment 2. Anterior vertical face of tergite 1
very weakly punctate, with a median longitudinal carina; transverse carina separating the
anterior face from posterior horizontal part weak and blunt; horizontal part relatively
finely punctate (interspaces usually larger than punctures), with a shallow median
depression near apical margin. Punctation on tergite 2 sparser than on tergite 1; other
tergites with only micropunctures. Gastral sternite 1 without distinct punctures or carinae.
Stemite 2 very weakly punctate; narrow basal area with strong longitudinal carinae; other
sternites with only micro punctures.
Black, with yellow and rufous markings (Fig. 176). Yellow are: c1ypeal base widely,
93
frontal marking connected by a bar with the yellow part of clypeus, a stripe on antennal
scape below, a marking on temple, triangular basal marking on mandible, anterior vertical
face of pronotum largely, a pair of large markings on pronotum dorsally, a pair of spots on
mesoscutum, scutellum largely, tegula basally, dorsal mesepisternum, spot on ventral
mesepisternum, metanotum largely, dorsal part of propodeum, lateral parts of propodeal
concavity, apical portion of the lateral part of propodeum (all these propodeal markings
are connected), narrow apical bands on tergites 1-4, those on sternites 2-4, coxae except for
posterior faces of all legs, femora and tibiae extensively. Mandible ferruginous. First two
gastral segments largely rufous. Tarsi of all legs dark brown. Wings brownish-infumated.
Holotype. Sf-, Fukiage-dani, Chichi-jima, Ogasawara Is., 18 vi 1972 C'f. Kusui).
Paratype. Sf-, Tsutsuji-yama, Chichi-jima, Ogasawara Is., 12 vii 1972 C'f. Kusui).
Distribution. Ogasawara (Bonin) Is. (Chichi-jima).
Biology. No information is available.
Pararrhynchium oceanicum miyanoi Sk. Yamane, ssp. nov.
Diagnosis. Female. Structure as in the nominotypical form. Black, with yellow and
rufous markings. Yellow are: upper lateral part of clypeus, triangular marking on
mandibular base, small interantennal marking (lost in one specimen), transverse frontal
marking, spot on temple, horizontal part of pronotum anteriorly (the marking extends
onto the anterior vertical face of pronotum), relatively large spot under wing base, tegula
except for center and margins, two large spots on scutellum, two transverse spots on
metanotum, dorsal part of propodeum extensively, apical band on gastral tergite 1 (not
clearly demarcated anteriorly), those on tergite 2 and sternite 2, irregular spots in apical
portion of tergite 3 and sternites 3 and 4, anterior face of coxae below of all legs, apical half
~178
:-:
o
o
0
Figs. 175-178. Pararrhynchium oceanicum sp. nov. (!f.). 175, head in frontal view; 176, body color
pattern; 177, median part of vertex; 178, propodeum in posterior view.
94
of fore and mid femora, outer face of tibiae of all legs. Gastral segments 1 and 2 almost
wholly rufous. Line on antennal scape below dirty yellow.
Similar to the nominotypical form, but differs therefrom in the following points:
yellow pronotal marking less extensive, propodeal concavity wholly black, rufous
markings on gastral segments 1 and 2 darker, leggs much less extensively marked with
yellow, and wings blackish-infumated. This form quite resembles the Haha-jima form of
Stenodynerus ogasawaraensis in color pattern.
Holotype., Kitakou, Haha-jima, Ogasawara Is., 21 v 1989, S. Miyano leg.
Paratypes. 1, Omotobashi, Haha-jima, 24 v 1989 (5. Miyano), 1 ,same loc. 10 xii 1989,
from a nest (5. Miyano).
Distribution. Ogasawara (Bonin) Is. (Haha-jima).
Biology. This form may be a tube renter.
Genus Orancistrocerus Vecht
Orancistrocerus Vecht, 1963, Zoo\. Verh. 60: 99-101 (type species: Odynerus drewseni Saussure, 1857,
original designation).
Japanese names: Entotsu-dorobachi Zoku (O-kabafusuji-dorobachi Zoku; O-kabafudorobachi Zoku).
This genus has few autapomorphies, and is separated from the related genera by the
combination of the following characteristics: cubital (submarginal) cell 3 further away
from the apex of the radial (marginal) cell; gastral tergite 1 with a distinct transverse ridge
separating the anterior vertical face from the posterior horizontal part; anterior margin of
clypeus wide, more or less emarginate; labrum of female rather densely covered with long
stiff hairs (Fig. 179); parastigma of fore wing at least half as long as stigma; propodeum
without a shelf; propodeal declivity not margined with a crest (Vecht, 1%3).
Only four species have been known from eastern Asia. In Japan, a single species, O.
drewseni, is widely distributed chiefly on the mainlands.
Orancistrocerus drewseni drewseni (Saussure)
(Figs. 119, 179-181)
Odynerus drewseni Saussure, 1857, Ann. Soc. Entomo!' Fr. (3)5: 318 (type loc.: China (?».
Rhynchium flavomarginatum: Matsumura, 1911, Thous. Ins. Jpn. Supp!. 3, p. 112, pI. 39, fig. 15; 1930, III.
Thous. Ins. Jpn. 2, p. 14, p!.2, fig. 15; 1931,6000 II\, Ins. Jpn.-Emp. p. 16, no. 79 (misidentification).
Odynerus (Ancistrocerus) fukaianus Schulthess, 1913, Ark. Zoo\. Stockholm, 8: 4, 9, fig. 8 (group
Euancistrocerus)(!f-) (type loc.: Harima, Honshu); 1934, Arb. Morph. Tax. Entomo\. 1: 71 (in key; fukayanus !);
Yasumatsu, 1938, Ins. Jpn. Ill. Icon. Co\. Nat. Dep. p. 359, fig. 628.
Ancistrocerus fukaianus: Sonan, 1938, Trans. Nat. Hist. Soc. Formosa,28: 79; Giordani Soika, 1941, Boll.
Soc. Venez. Stor. Nat. 2: 238; Yasumatsu, 1950, Icon. Ins. Jpn. 2nd 00. p. 1457.
Orancistrocerus drewseni: Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat. Ed. 3, p. 297, p!. 149, fig. 1.
Orancistrocerus drewseni drewseni: Vecht, 1963, Zoo!. Verh. 60: 102-103; Yamane and Tano, 1983, Mem.
Kagoshima Univ. Res. Center S. Pac. 4: 122.
Japanese names: Entotsu-dorobachi (O-kabafusuji-dorobachi; 6-kabafu-dorobachi).
Diagnosis. Female. Body length (h+th+t1+2): 12.5-16.5 mm. Fore wing length: 12.5-14.0
95
",\
'\
,
"
181
Figs. 179-181. Orancistrocerus drewseni. 179, female c1ypeus and labrum; 180, male c1ypeus; 181,
terminal segments of male antenna. 180 and 181 are based upon a Korean specimen.
mm. Head subcircular, densely punctate. Clypeus slightly wider than high, medially
striate and laterally punctate; apical margin wide, shallowly emarginate (Fig. 179).
Interantennal keel blunt. Depression for cephalic foveae ill defined; area around the
depression impunctate; distance between the foveae as long as that between the posterior
ocelli. Anterior vertical face of pro not urn smooth, shining, with a few punctures; posterior
horizontal part densely punctate. Mesoscutum densely punctate; punctures running into
longitudinal striae. Scutellum more coarsely sculptured than mesoscutum. Mesopleuron
with large punctures densely, somewhat reticulate; epicnemium well defined by
epicnemial carina, smooth but not shining. Metanotum sloping posteriorly, with large
punctures. Metapleuron with a few punctures. Propodeum without shelf; concavity
shallow, with a median carina which is interrupted at some distance from the tip, with illdefined large punctures; dorsolateral face with large punctures; lateral side above with
punctures and below with transverse carinae. Gastral segment 1 narrower than segment 2.
Anterior vertical face of tergite 1 with small, sparse punctures; tergites 1, 3-5 strongly and
densely punctate; punctation on tergite 2 much finer except in basal and apical narrow
zones; tergite 6 without large punctures. Narrow basal part of sternite 1 impunctate;
posterior part weakly striate. Sternite 2 anteriorly with strong short carinae; posterior
portion somewhat shining, sparsely punctate; punctures coarser near its base. Punctures
on other sternites much finer.
Black, with the following parts orange or orange yellow: cIypeus, a transverse frontal
mark with a median projection toward cIypeus, antennal scape below, a basal triangular
marking on mandible, relatively wide apical bands on tergites 1 and 2. Mandible largely,
antennal flagellum below, upper faces of mid and hind femora apically brownish or
rufous. Wings dark brown.
Male. Not examined.
Material examined. Honshu: Yamagata-ken -1!f, Oguni, 1 viii 1976 (HI), 1!f, Nan'yo, 16 vii 1976 (HI), 1
!f, same loc., 26 vii 1981 (HI); Niigata-ken -1!f, Shibata, 18 vi 1967 (HI), 3!f !f, Nagaoka, 10 vii 1976 (SKY), 3
96
!f.!f., Shibata, 7 vii 1976 (SKY), 5!f. !f., Akadani, Shibata, 3 vii 1977 (A. Seino & H. Koike), 1 !f., Takane, 2 vii
1978 (HI), 3!f. !f., Iwakuzure, 17 ix 1981 (KB), 6!f. !f., Senami, 11-30 vii 1981 (KB), 1 !f., same loc., 11 ix 1981
(KB), 1 !f., same loc., 27 vi 1983 (KB), 1 !f., same loc., 10 vii 1984 (KB); Tochigi-ken - 1 !f., Utsunomiya, 2 viii
1970 (T. Hasegawa); Hyogo-ken - 1 !f., Sasayama, Tamba, 7 viii 1963 (K. Iwata), 1 !f., same loc., 13 viii 1965 (H.
Katayama); Osaka-fu - 5!f.!f., Iwawaki, Kawachi, 6-10 vi 1964 (H. Katayama); Kyoto-fu -l-'f-, Minna-ji, 20 vi
1958 (K. Iwata); Wakayama-ken - 1 !f., Koyasan, 7 viii 1955 (K. Iwata); Shimane-ken - 1 !f., Sugata, Matsue, 9 viii
1985 (N. Sugiura), 1 !f., Hikimi, 8 xi 1987 (N. Sugiura).
Sado-ga-shima: 1 !f., Tagirisu, Mano, 23 ix 1981 (KB).
Awaji-shima: 1 !f., 10 viii 1954 (T. Mori).
Shikoku: Tokushima-ken - 2!f. !f., Tsudayama, Tokushima-shi, 22-24 viii 1964 (H. Iwasaki).
Ky11shii: Fukuoka-ken - 1 !f., Kurogi, 28 vii 1983 (Y. Takai), 2!f. !f., same loc., 30 viii 1984 (Y. Takai);
Miyazaki-ken - 1 !f., Hinokage, Nishiusuki, 5 vi 1987 (K. Suzuki); Kagoshima-ken - 1 !f., Takachiho-no-mine,
Kirishima, 8 vi 1973 (K. Ohara), 1 !f., Shiroyama, Kagoshima-shi, 10 vii 1981 (SKY).
Goto Is. (Nagasaki-ken): Nakadori-jima -1!f., 11 viii 1976 (J. Nakayama).
Tsushima Is. (Nagasaki-ken): Shimo-agata - 2!f. !f., Izuhara, 26 vii 1986 (K. Nakamine).
N. Ryukyus: Tane-ga-shima - 1 !f., Nishi-no~mote, 13 v 1981 (SKY), 1!f., same loc., 4 viii 1983; Yakushima- 1 !f., Miyanoura, 11 viii 1981 (SKY).
Distribution. Honshu; Sado-ga-shima; Awaji-shima; Shikoku; Kyushu; Coto Is.
(Nakadori-jima; Fukue-jima; Naru-jima); Tushima Is. (Shimo-agata); Amakusa Is. (Kamishima); Osumi Is. <Tane-ga-shima; Yaku-shima). China.
Taxonomic notes. Judging from the figures the females of "Rhynchium flavomarginatum
Sm." in Matsumura (1911, 1930, 1931) apparently have pale markings on alitrunk and
gastral tergites 3-6 (?). However, Matsumura did not refer to these markings at all in the
text: "Body black, densely punctate, with sparse short hairs. Clypeus, marking between
antennal bases, scape below, apical markings on 1st and 2nd abdominal segments orange
yellow...." (translation by me). The figures seem incorrect.
Biology. The nesting habits and life history of this form were studied by Iwata (1938a;
*
.. P. ornatum
P. tsuneki i
• P. ishigakiense
.& o. kogimai nagasei
• O. kogimai kogimai
...-
182
Figs. 182. Distribution of Okinawepipona and Pararrhynchium species in the Ryukyus.
97
referred to as Ancistrocerus fukaianus). Additional data are given in a text written in
Japanese (Iwata, 1983). A series of beautiful photos of nesting females and nests are given
in Iwata (1982). Itino (1986a) made a comparative study on the population ecological
aspects of this form and Anterhynchium f1o.vomarginatum micado. Iwata (1939b) studied the
Taiwanese subspecies ingens (Schulthess) that had a similar nesting behavior. O. aterrimus
erythropus (Bingham) also shows a similar pattern of nesting behavior (Iwata, 1964).
The Japanese population of this form is very probably parthenogenetic, because only
females are collected in the field and from nests. In the Korean population (also belonging
to the subsp. drewsem) and the Taiwanese subspecies ingens the male is commonly found.
Female wasps build their gourd-shaped nests of clay in cavities of wood, depressions
on rocks and stones, bamboo tubes, etc. It was once observed that a nest was built on a
rootlet of a plant hanging from a clayey cliff (Iwata, 1938a). When bamboo tubes with a
small diameter are used, they are simply partitioned into cells by clayey wall. The nest is
provided with a delicate chimney (ca. 10 mm diam.) of variable length which is built
downward from the mouth of nest and removed after the completion of nest. A nest
contains 1-4 (rarely 5) cells that are linearly arranged; each cell is supplied with 5-16
caterpillars before the hatching of wasp egg, and 24-46 (m=36) in total. These caterpillars
are progressively provided by the mother wasp with the growth of her young. When the
nest is disturbed, the mother wasp may abandon the prey in the cell under provisioning
and make empty cells to hide the inner larval cells already completed. The female may lay
a maximum of six eggs. Two generations occur in a year. According to ltino (1966c) adult
daughter wasps generally do not disperse, sometimes reutilize mother nests or construct
their nests on those of preceding generations. As a result nest aggregations are not rare.
Prey: Larvae of Pyralidae, Gelechiidae, Tortricidae, Noctuidae (Lepidoptera) (Iwata, 1938a,
1983).
Parasite: Pseudoxenos iwatai (Strepsiptera, Stylopidae) (Iwata, 1938a, 1983).
Parasitoids: Anthrax distigma ? (Diptera, Bombyliidae), Amobia distorta (Sarcophagidae),
Megaselia sp. (Phoridae), Acroricnus ambulator (Hymenoptera, Ichneumonidae), a
chrysidid, and Macrosiagon nasuta (Coleoptera, Rhipiphoridae) (Iwata, 1938a; Itino, 1986a).
Genus Ancistrocerus Wesmael
Andstrocerus Wesmael, 1836, Bull. Acad. R. Belg. 3: 45 (subgenus of Odynerus Latreille)(type species:
Vespi pirietum Linnaeus, deSignated by Giraud, 1879); Bequaert, 1925, Trans. Amer. Entomol. Soc. 51: 85 (as
subgenus of Andstrocerus Wesmael, s. lat.); Carpenter, 1986, Psyche, 93: 64 (as genus); Bliithgen, 1961,
Faltenwespen Mitteleuropas, p. 151 (as genus).
Euancistrocerus Dalla Torre, 1904, Gen. Ins. 19: 36 (new name).
Japanese name: Suji-dorobachi Zoku.
Diagnoses for this taxon have been given by various authors, though that in the key
to the European eumenid genera by Bliithgen (1961) may be most useful.
Head, thorax, propodeum, gastral tergite 1 (+ 2), and stemites 1 and 2 generally with
abundant long hairs. Head with a rounded depression (Scheitelgriibe) for cephalic foveae
on the vertex in the female (rarely evanescent). Supraclypeal area with a rather deep pit
just below each antenna I socket. Front corners of pronotum more or less angled, often
sharply pointed in the male (Figs. 192-198). Notaulices absent. Prescutal groove usually
98
inconspicuous. Epicnemial carina absent. Metanotum posteriorly not spinose. Tegula
relatively slender, produced and more or less tapering caudally (Fig. 191), and not
strongly punctate. Propodeum with no or at most a narrow shelf (in at least Japanese
species), with a series of sharp ridges enclosing the posterior face (propodeal concavity)
that is flat rather than concave, but the superior ridges sometimes much reduced. Lateral
ridges and/or posterolateral angles of propodeum sometimes developed. Punctation on
tergites 1 and 2 generally weak. The anterior vertical face of tergite 1 clearly separated
from the horizontal disc by a sharp transverse carina; tergite 1 without longitudinal
furrow. Tergite 2 without yellow lateral spots. Gastral color pattern is relatively stable
within a species and is useful in separating species (Figs. 214-220).
Earlier authors (e.g., Saussure, 1852-8; Bequaert, 1918, 1925) lumped almost all the
species of "Odynerus" with a transverse carina on tergite 1 into a group (Ancistrocerus).
Bequaert (1925), however, wrote: "Ancistrocerus as at present understood is evidently
polyphyletic ..... The transverse carina is evidently a structure aquired independently in
several lines of descendent among the Eumenidae". In the present paper I treat Bequaert's
subdivision Ancistrocerus (proper) as a genus. This genus is widely distributed in the
Palearctic and Nearctic regions, fewer number of species being found also in the Old and
New World tropics. One Japanese species (A. anti/ope) is also found in both Europe and
North America, showing a Holarctic distribution (cf. Krombein, 1979). Useful keys are
available for the European species (e.g., Bliithgen, 1961; Yeo & Corbet, 1983).
Key to the Japanese species of Ancistrocerus
1. Body hairs very long; hairs on vertex distinctly longer than the distance between the
posterior ocelli; all the tergites but the last one with long hairs; female antennal scape
with many long hairs especially near its base. Male mandible with a large concavity in
the middle on inner margin. Metanotum elevated above a level with the posterior
portion of scutellum. Last tergite of the female with a yellow spot. Tegula marked with
yellow................................................................................................. A. oviventris oviventris (L.)
- Body hairs shorter; hairs on the vertex as long as or shorter than the distance between
the posterior ocelli; hairs on tergites 2-4(5) very short; female antennal scape without
long hairs (if present, then few in number and inconspicuous). Metanotum not elevated,
nearly at the same level as scutellum ....................................................................................... 2
2. Metapleuron and lateral part of propodeum in the lower half smooth, with glaring
luster. Posterior face of propodeum not striate, polished. Lateral ridge of propodeum in
the male well developed, forming a flat wing. Terminal segment of male antenna thick
and short, the apex not reaching the middle of segment 11. Yellow pronotal band widely
interrupted medially. Female clypeus wholly black. .................. A. anti/ope anti/ope (Panz.)
- Metapleuron and lateral part of propodeum in the lower half finely striate, wrinkled, or
micropunctured, without glaring luster. Posterior face of propodeum often striate.
Lateral ridge of propodeum not forming a distinct wing. Terminal segment of male
antenna larger; the apex extending beyond the middle of segment 11 (if shorter, then it
is slender and apically pointed) ................................................................................................ 3
3. Gastral tergites 1-4 each with a yellow apical band; last tergite without yellow spot in
the female. Superior ridge of female propodeum much reduced, often almost absent.
Female clypeus with a pair of upper yellow spots. In the male fore and mid femora
99
extensively yellow on the anterior face. Sternite 2 distinctly angulate near the base .
............................................................................... ........... ............................. A. nigricornis (Curt.)
- Tergites differently marked with yellow. Superior ridge of female propodeum usually
distinct. In the male fore and mid femur usually blackish, at most marked with yellow
near its apex ................................................................................................................................. 4
4. Female dypeus with two pairs of yellow spots, one near the base and the other near the
apex. Tergites 1-4 with yellow apical bands; last tergite with a yellow spot in the female.
Female tegula with anterior and posterior yellow spots. The male is not known in Japan .
.......................................... ............................................. ... .................................... A. parietinus (L.)
- Female dypeus differently colored. At most tergites 1-3 with a yellow band. Tegula in
both sexes wholly blackish brown ............................................................................................ 5
5. Only tergites 1 and 2 with yellow apical bands. Female dypeus with a pair of yellow
spots near its apex. Antennal flagellum in the female ferruginous below. Terminal
segment of male antenna small; the apex not reaching the base of segment 11 .
....................................................................... ............. ........................ ........... A. japonicus (Schult.)
- Tergites 1-3 with yellow apical bands (the band on tergite 3 sometimes reduced or absent
in the female). Female dypeus usually wholly black. Antennal flagellum in the female
blackish below, at most brownish apically. Terminal segment of male antenna longer; the
apex almost reaching the base of segment 11 .......................................................................... 6
6. Body slender; in the female tergite 1 (as measured from the transverse carina to the
apical margin) 5/3 times as wide as long. Female scutellum wholly black. In the male,
antennal scape usually largely black; legs often without yellow markings, at most with
them on the anterior face of fore and mid tibiae ................ A. trifasciatus shibuyai (Yasum.)
- Body more stumpy; in the female, tergite 1 two times as wide as long. Female scutellum
almost always with yellow spots. In the male, antennal scape usually yellow below; legs
more extensively yellow........: .................................................................. A. melanocerus (D.T.)
Ancistrocerus japonicus (Schulthess)
(Figs. 183, 192, 201, 202, 214, 221)
Odynerus (Euandstrocerus) japonicus Schulthess, 1908, Mitt. Schweiz. Entomol. Ges. 11: 285 (~) (type
loc.: Yokohama, Honshu); 1934, Arb. Morph. Taxon. Entomol. 1: 73 (in key).
Andstrocerus japonicus: Vecht and Fischer, 1972, Hym. Cat. (n. ed.) 8: 113; Giordani Soika, 1986, Boll.
Mus. Civ. Stor. Nat. Venez. 35: 150-152.
Japanese names: Yamato-suji-dorobachi (Kiobi-suji-dorobachi).
Diagnosis. Female. Body length (h+th+tl+2): 9.0-11.5 mm. Forewing length: 8.5-11.0
mm. Head slightly wider than high, densely punctate. Clypeus wider than high, apically
shallowly emarginate; lateral angles of the emargination round (Fig. 201); punctation
rather superficial. Frontal keel distinct. Depression for cephalic foveae weakly defined;
foveae close to each other. Temple, as seen from above, slightly wider than the superior
lobe of eye. Distance between posterior ocelli approximately as long as distance between
posterior ocellus and eye. Occiput sparsely punctate. Antenna relatively short; flagellum
gradually widened toward apex until segment 8. Thorax coarsely and densely punctate.
Each side of the anterior vertical face of pronotum with a transverse furrow; pronotal
carina weak in dorsal part. Prescutal grooves only basally visible, not conspicuous;
mesoscutum with a shallow longitudinal furrow on each side at a short distance from
100
6}87 ~8
Figs. 183-200. Various characters in the Japanese Ancistrocerus. 183-188, segments 10-13 of male
antenna in japonicus (183), trifasciatus (184), melanocerus (185), nigricornis (186), oviventris (187)
and anti/ope (188); 189, scape, pedicel and flagellar segment 1 of female antenna of oviventris;
190, male mandible of oviventris; 191, tegula and parategula of nigricornis; 192-198, right front
comer of pronotum in japonicus 3'(192), trifasciatus 3'(193), me/anocerus 3'(194), nigricornis .'f
(195) and 3'(96), oviventris 3'(197) and antilope 3'(198); 199, 200, gastral segments 1 and 2
(profile) of trifasciatus (199) and nigricornis (200).
tegula; punctation on epicnemium fine and sparse; scutellum without longitudinal furrow.
Metanotum without horizontal part, nearly directly sloping to propodeum; metapleuron
without distinct punctures. Propodeal shelf very narrow, with a median pit; propodeal
concavity nearly wholly enclosed by ridges, obliquely and shallowly striate, with a
complete vertical carina; superior ridge sometimes reduced to some extent in its lower
101
'v--J
205
202
201
,
/
\
206
/
209
208
,-----/
,
/
/
210
212
\--J
\
213
'-.
VV
203
'v---J
204
~
Figs. 201-213. Clypeus of Japanese Ancistrocerus species. 201, japonicus !f; 202, ditto d'; 203,
trifasciatus !f; 204, ditto 6'; 205, melanocerus !f; 206, ditto 6'; 2rJ7, parietinus !f; 208, nigricornis
!f; 209, ditto 6'; 210, oviventris !f; 211, ditto d'; 212, antilope !f; 213, ditto d'.
part; dorsolateral part of propodeum with large punctures; lateral ridge almost absent;
lateral face indistinctly striate. Anterior vertical face of gastral tergite 1 only superficially
and sparsely punctate in its upper part; punctation on posterior horizontal part much
denser. Tergite 2 much more finely and sparsely punctate than tergite 1. Apical half of
tergites 3-5 rather strongly punctate; last tergite almost impunctate. Apical part of stemites
2-5 rather strongly punctate; last sternite impunctate.
Black; the following parts yellow or orange yellow: a pair of spots on clypeus apically,
a frontal spot, a small spot between antennal socket and eye, a very short line on temple, a
triangular basal marking on mandible, antennal scape below, pronotal band medially
interrupted and not extending to lateral face of pronotum, a narrow and regular apical
102
band on tergite 1, a wider apical band on tergite 2 (Fig. 214), posterolateral corners of
sternite 2, anterior and apical part of fore femur, anterior face of fore tibia. Mandible
blackish brown, apically rufous. Antennal flagellum below ferruginous.
Male. Body length (h+th+t1+2): 7.5-8.0 mm. Fore wing length: 7.0-7.5 mm. Similar to
the female in structure and color pattern. Differs from the latter in the following details:
head distinctly wider than high; clypeus nearly as wide as high, anteriorly more deeply
emarginate; lateral angles of the emargination more sharply pointed (Fig. 202); clypeus
nearly wholly yellow; mandible more extensively yellow; yellow spots on frons, between
antennal socket and eye, and on temple much reduced and sometimes lost; pronotal band
much reduced; anterior face of mid tibia often marked with yellow. Terminal segment of
antenna very small; the apex not reaching the base of segment 11 (Fig. 183).
Material examined. Honshu: lwate-ken -1!?-, 27 vi 1944 (Ogasawara), 1!?-, Tamayama, 1968-69, reared
from nest (SKY); Niigata-ken - 1!?-, Senami, 9 vi 1979 (KB), 1!?-, Shibata, 30 vi 198? (A. Seino); Tokyo-to - 1!?-,
Okutama,23 ix 1978, 2!?-!?-, same loc., iv-v 1978, reared from nest (Y. Tanaka), 1 d'1!?-, Nishitama, iv-v 1978,
reared from nest (Y. Tanaka); Nagano-ken -1!?-, Takase-dani, Omachi, 20 vii 1982 cYV. Miyata); Yamanashi-ken 1 d'1!?-, Shioyama, iv-v 1978, reared from nest (Y. Tanaka); Fukui-ken - 1!?-, Ashuwa, 17 v 1959 (TT), 1 d',
Otani, 6 viii 1%4 (TT), 1!?-, Koike, Ono,22 ix 1968 (YH), 1!?-, Asahimaesaka, 30 viii 1979 (TT), 1!?-, Nagadani, Natasho, 2S vii 1980 (TM); Ishikawa-ken - 1 d', Koyajiri-sekkei, Mt. Hakusan, 17 vii 1984 (1. Togashi);
Gifu-ken -1!?-, Kamiaizu, Kaminoho, MUgi, 5 viii 1981 (Y. Takai).
Sado-ga-shima: 1 d', 1987-88, reared from a nest (T. Onuma).
Shikoku: Kiichi-ken - 2!?-!?-, Engyoji, Kochi-shi, 31 v 1931 (Y. Sugihara), 1 d', Kodakasa-yama, 21 v 1936
(H. Okamoto).
Kyushu: Kagoshima-ken - 1!?-, Saruga-jo, Tarumizu, v 1980, reared from nest (79T-6) (H. Nagase), 1!?-,
Takakuma, 19 vi 1981 (KT).
Figs. 214-220. Body color pattern in the Japanese Ancistrocerus species (!?-). 214, japonicus; 215,
trifasciatus; 216, melanocerus; 217, parietinus; 218, nigricornis; 219, oviventris; 220, antilope.
103
N. Ryukyus: Yaku-shima -1!f, Ishizuka, 30 vi 1965 (1<. Hashimoto), It, Kosugidani, 20 v 1968 (A.
Mori), 1!f ,Onoaida, 23 iv 1982 (SI).
Distribution. Honshu; Sado-ga-shima; Shikoku; Kyushu; Shika-no-shima (Fukuokaken); Osumi Is. (Yaku-shima).
Biology. According to Iwata (1938a), this species nests on depressed rocks or
inscriptive stones, whose surfaces are always vertical and facing south or east. A
completed nest studied by Iwata was constructed in an inscription deeply incised in
granite, and contained eight cells attached to each other in a mass; the entire surface was
thickly covered with "crepissage". Each cell is a cylindrical and elongate mud pot.
However, I found a 1-celled nest made in a reed tube in Iwate-ken. In this case the cell was
not a pot, but with a mud entrance plug (cell wall). Similar nests were found also on Sadoga-shima (Onuma, 1989a) and in Kagoshima-ken (Nagase, pers. comm.). Onuma
mentioned that the number of cells per tube nest ranged between 1 and 16 (usually
between 2 and 6). Thus, this species seems to be either a tube-renter or a mud-dauber
(sensu Spradbery, 1973). The brood cell is provided with six to seven lepidopterous
caterpillars. The wasp has at least two generations a year in the Kansai District (Honshu),
and overwinters as prepupal stage in the nest.
Parasitoids: an ichneumon wasp (Hymenoptera) and Chrysis sp. (Hymenoptera,
Chrysididae).
Ancistrocerus trifasciatus shibuyai (Yasumatsu)
(Figs. 184, 193, 199, 203, 204, 215, 222)
Odynerus (Ancistrocerus) shibuyai Yasumalsu, 1938, Mushi, 11: 83 (t!f )(type loc.: Osaka, Honshu);
Vecht and Fischer, 1972, Hym. Cat. (n. ed.) 8: 117.
Ancistrocerus trifosciatus shibuyai: Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 147-148.
Japanese name: Shibuya-suji-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 9.0-10.5 mm. Fore wing length: 8.0-9.5
mm. Head as wide as high, densely punctate. Clypeus wider than high, distinctly
punctate and somewhat carinate, apically very narrowly and shallowly emarginate; lateral
teeth of the apical emargination blunt (Fig. 203). Temple moderately developed, as seen
from above as wide as superior lobe of eye. Punctation on gena and vertex finer than on
frons. Depression for cephalic foveae ill defined; foveae very small and close to each other.
Anterior ocellus slightly larger than posterior one; distance between the posterior ocelli as
long as that between posterior ocellus and eye. Thorax rather depressed dorsoventrally,
densely punctate. Anterior vertical face of pronotum with a transverse furrow on each
side. Prescutal groove evanescent; epicnemium almost impunctate; punctation on
scutellum sparser than on meso scutum. Metanotum gently sloping toward propodeum;
metapleuron without punctures. Propodeum with a narrow shelf that is sloping and has a
median pit. Propodeal concavity wholly enclosed by ridges, superficially striate, with a
complete median carina. Dorsal face of propodeum with large punctures; lateral ridge not
developed; lateral face finely and superficially striate. Punctation on gastral tergites finer
than in A. japonicus, especially on tergite 2. Tergites 3-5 and sternites 3-5 with relatively
large punctures.
Black, with the following parts yellow: small triangular marking on mandible basally,
104
frontal spot, minute spot between antennal socket and eye, very minute spot on temple,
apex of antennal scape below, narrow and short band on pronotum anteriorly, apical
bands on tergites 1 and 2 (rarely also on 3) (Fig. 215), a complete apical band on sternite 2,
anterior faces of fore and mid tibiae (the latter often wholly black). Mandible black, with
brown teeth. Antennal flagellum almost wholly black.
Male. Body length (h+th+t1+2): 6.5-8.0 mm. Forewing length: 6.5-7.5 mm. Differs
from the female in the following details: head wider than high; clypeus nearly as wide as
high; front corners of pronotum distinctly produced (Fig. 193); mandible and clypeus
yellow except periphery; 3-4 terminal segments of antenna partly rufous; yellow markings
on head and thorax much reduced, sometimes partly lost. Antenna! hook larger than in
the preceding species; the apex almost reaching the base of segment 11 (Fig. 184).
Material eXilmined. Hokkaide: 1!j'., loco not stated, 4-10 viii 1928 (Uchida & Keno), 1!j'., Shikaribetsu, 24
viii 1934 (Uchida), 1!j'., Sapporo, 25 viii 1935 (Y. Sugihara), 1!j'., Tengu-<lake, 21 vii 1935 (S. Sasaki), 1 C!',
Moiwa-yama,25 v 1946 (S.F. Sakagami), 1 C!', Moiwashita, Sapporo, 11 ix 1957 (TN), 1 C!', SOunkye, 4 viii 1959
,
o
A. japonicus
221
l"./
\ ..•
•
A. trifasciatus
222
A. melanocerus
223
Figs. 221-223. Distribution of Japanese Ancistrocerus 0>.
105
(H.D.), 1 ~, Miyanomori, Sapporo, vi 1%1 (yamane), 1 ~, same loc., 12 vi 1%2 (T. Kubo), 1 ~, same loc., 13
vi 1%2 (SKY), 3 ~ ~ , same loc., 8 vii 1965 (SY), 1 ~, Teine-yama, 25 vii 1965 (SKY), 1 ~ , Kozawa, Shiribetsu, 2
viii 1968 (TN), 1 ~,Jozankei, vi 1971 (5. Aoki), 1 ~, Bibai, 23 viii 1971 (K. Kamijo), 1 ~, Bibai, 17 vi 1972 (M.
Taira), 1 (J', Shikotsu-ko, 17 vii 1977 (SKY), 1 ~, Aizankei, 2-4 viii 1977 (TM), 1 ~, Muine-yama, 20 viii 1977
(SKY), 1 ~, Bibai, 13 vi 1978 (K Kamijo), 1 ~,Asahigawa, 6 viii 1979 (T. Inaoka).
Rebun-to: 1 (J', 26 vii 1951 (M. Konishi).
Honshu: ADmori-ken -1 ~,Sukayu, 18 viii 1968 (TN), 1 ~, Iwaki--san, 8 viii 1982 (M. Yamada); Iwate-ken
- 1 ~,Ashiro, 6 ix 1979 (YM), 1 ~, Mt. Hayachine, 31 vii 1979 (YM); Miyagi-ken - 1 ~, Mt. Zao, 27 vii 1979 (T.
Nishioka), 1 (J', same loc., 2 ix 1979 (K. Goukon), 1 (J', same loc., 29 v 1982 (K. Goukon); Niigata-ken - 1 ~,
Mikuni Pass, 6 ix 1981 (KB); Gumma-ken - 2~ ~, Osawa, Nikko, 16 vii 1978 (HI), 1 (J', Kensetsu-daira,
Okunikko, 31 vii 1982 (HI); Saitama-ken - 1 ~, Karisaka, 26 vii 1974 (K. Hara); Tochigi-ken - 1 ~, Chuzenji, 31
vii 1915 (E. Gallois); Nagano-ken - 1 (J', Mt. Tenguhara, 12 ix 1982 (HI); Yamanashi-ken - 1 (J'2~ ~, Shiroyama,
iv - v 1978, reared from nest (Y. Tanaka); Fukui-ken - 1 (J', Ono, 23 ix 1974 (yH), 1 ~, same loc., 25 viii 1976
(yH), 1 ~, Koike, 29 viii 1978 (H. Kurokawa), 16', Kanakusa-dake, Ikeda, 13 ix 1981 (H. Kurokawa); Naraken - 1 ~,Obako-dake, 22 vii 1957 (K. Iwata).
Distribution. Hokkaido; Rebun-to; Honshu (central and northern parts). The nominotypical subspecies is widely distributed in the Palearctic region, from Europe in the west
to Sakhalin and Kamchatka in the east (Vecht & Fischer, 1972).
Biology. According to Shibuya (1938) this form generally constructs one-celled nests
in dry reed tubes; two-celled nests are rather rare. Caterpillars of leaf-rollers (microlepidopterous larvae) are stored in brood cells. In one case eleven caterpillars (135 mg in
total) were found from in a cell (May 26 1934). The wasp constructs a double entrance
plug; the inner wall is relatively thick, dirty black in color, and probably made of mud and
sand grains mixed with wasp's saliva, while the outer one is made of yellowish, semitransparent, sticky resin that is finally plastered with a few sand grains or fibers of
weather-bitten wood on its outer surface. Though adult wasps are collected from May to
September (see Material examined above), this form is univoltine and overwinters at
prepupal stage in Kyoto, Honshu (Shibuya, 1938). The biology of the nominotypical
subspecies was studied by Enslin (1921), and summarized in Bliithgen (1961).
Parasitoids: Chrysis (Tetrachrysis) galloisi (?) (Hymenoptera, Chrysididae) and a
tachinid fly (Diptera) (Shibuya, 1938).
Ancistrocerus melanocerus (Dalla Torre)
(Figs. 185, 194,205,206,216,223)
Anc:istrocerus nigricornis Morawitz, 1889, Hor. Soc. Entomol. Ross. 23: 161 «J')(type loc.: Kansu, China)
(homonym of Ancistrocerus nigricornis (CurtiS, 1791)).
Odynerus melanocerus Dalla Torre, 1894, Cat. Hym. 9: 78 (new name).
Ancistrocerus melanocerus: Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 149-150.
Ancistrocerus densepilocellus Cameron, 1911, Entomologist, 44: 288 «J')(type loc.: Japan); Sato, 1963,
Nature Study, 9: 5 (in key), figs. 36,37, 39-41; Vecht and Fischer, 1972, Hym. Cat. (n. ed.), 8: 111.
Odynerus (Ancistrocerus, Euancistrocerus) densepilosellus: Schulthess, 1934, Arb. Morph. Taxon. Entomol.
1: 73-74 (in key).
Japanese names: Kebuka-suji-dorobachi (Ko-suji-dorobachi; Miyama-suji-dorobachi).
Diagnosis. Body length (h+th+tl+2): 6.5-10.5 mm in ~, 5.5-7.5 mm in t. Fore wing
length: 7.0-10.0 mm in ~,6.0-8.5 mm in 6'. Closely related to A. trifasciatus shibuyai, from
which the present species is distinguished by the following points: body more stumpy;
gastral tergite 1 (measured from the transverse carina to the apical margin) 2 times as
106
wide as long (in trifasciatus shibuyai 1 2/3 times as wide as long); punctation on tergites 1
and 2 coarser; propodeal shelf narrower, often virtually absent. In the female, scutellum
with two yellow spots (in the specimens from Hokkaid6, the spots often reduced or lost);
basal marking on mandible much reduced or lost; tergite 3 with a yellow apical band (in
the specimens from Hokkaid6 the band is often reduced or lost) (Fig. 216); anterior face of
fore femur apically and that of fore tibia rufous (not yellowish). In the male, apical margin
of c1ypeus much more deeply incised (but the condition varies among individuals) (Fig.
206); scape below extensively yellow; tergite 4 and sternite 3 often with apical bands; hind
tibia usually extensively yellow.
This species varies in size, structure and color pattern among individuals. Superior
carina is often not developed. Female c1ypeus has rarely two or four yellow spots.
Anterior emargination of male c1ypeus considerably variable in shape and depth, but
constantly deeper than in trifasciatus shibuyai.
Material examined. Hokkaido: 1 e!, Sapporo, 7 vi 1938 (Y. Sugihara), 1 e!, Ashiribetsu, 19 ix 1965
(Tamura), 1-'?-, Bibai, 20 vii 1972 (M. Taira), 1 e!, Kiyokawa, Kamikawa-gun, 8 vii 1977 (Nosaka), 3 e! e!,
Muine-yama (Horainuma - Hiitte), 8 viii 1978 (SKY), 2 C!' C!' , Muine-yama (Hiitte), 9 viii 1978 Tsujii), 2 e!
C!', Muine-yama (top, 1400 m alt.), 8 viii 1978 (SKY), 6e! C!'3-l'- -l'-, Teine-yama (1000 m alt.), 13 viii 1978 (SKY),
Ie!, Soranuma-dake, 20 viii 1978 (T. Fujisawa), 2e!e!1-l'-, Asahigawa, 25-26 v 1979 (T. Inaoka), 1-l'-,
Nukabira, 19 vii 1979 (SKY), 1-l'-, Misumai, Sapporo, 11 viii 1984 (S. Makino), 3-l'- -l'-, Tomakomai, 19 ix 1984
(I. Tateyama).
Rebun-to: 1-l'-, 25 viii 1951 (M. Konishi).
Honshu: Aomori-ken -1 e!4-l'- -l'-, Otake, HakkOda-san, 21 viii 1983 (M. Yamada), 1 C!', Sukayu, 11 ix 1983
(M. Yamada); Fukushima-ken - 4 C!' e!3 -l'- -l'-, Azuma-yama (1400 m alt.), 5 ix 1981 (HI); Miyagi-ken - 1-l'-,
Aobayama, 8 v 1981 (K. Goukon), 1 C!', Mt. Zao, 4 vii 1979 (K. Kojima & T. Nishioka), 1 C!'1-l'-, same loc., 31
a.
,
o
eA. parietinus
.A. oviventris
224
eA. nigricornis
.A. antilope
225
Figs. 224, 225. Distribution of Japanese Ancistrocerus (II).
107
viii - 2 ix 1979 (K Goukon), 1 d', same loc., 4 ix 1980 (K Goukon); Niigata-ken -1 d', Myoko, 24 viii 1962 (HI),
1 d', same loc., 8 viii 1967 (HI), 1 d', Mikuni Pass, 6 ix 1981 (1<B), 1 -,?-, Ojiya, 18 ix 1982 (KB), 1 d', Seki Spa, 22
ix 1984 (1<B); Saitama-ken -1 d', Kumotori-rindo, 27 v 1972 (TN), 1-'?-, Chichibu, 24 vii 1977 (TN); Gumma-ken 1 d', Nikko, viii 1912 (5. Matsumura), 1-'?-, Osawa, Nikko, 16 vii 1978 (HI), 1-'?-, Suganuma, Okunikko, 5 vii
1980 (HI); Nagano-ken - 2 d' d', Todai, Ina, 29 iv 1963 (YM), 1 d', same loc., 17 x 1977 (5. Aoki), 1-'?-, Tomono,
Saku,3 ix 1977 (M. IGuchi); Ishikawa-ken -1J', Haku-san, 1 viii 1960 (TT), ISf, same loc., 8 ix 1978 (1.
Togashi), 1 Sf, same loc., 24 viii 1984 (I. Togashi); Fukui-ken- 1 Sf, Arashi, 18 ix 1977 (H. Kurokawa), 1 d',
Koike, 12 viii 1979 (H. Kurokawa).
Shikoku: Kikhi-ken -1 Sf, Kodakasa-yama, K6chi-shi, 17 vii 1930 (Y. Sugihara), 1!j?, same loc., 12 iv 1938
(Y. Sugihara), 1 d'1!j?, Hirooka, 22 iv 1934 (H. Okamoto). Ehime-ken - 1 J', Ishizuchi-yama, 23 vii 1935 (H.
Okamoto), 6 J' J', Kamega-mori, 17 vii 1933 (Y. Sugihara); Tokushima-ken- 1 J', Tsurugi-san, 24 vii 1970 (TT).
Distribution. Hokkaido; Rebun-to; Honshu; Shikoku; Kyushu. Korea; Mongolia.
Biology. Iwata (1938a) observed two nests in Kamikochi (1500-1600 m in alt.), Naganoken (referred to as A. densepilosellus). Recently, Goukon (1983c, 1984, pers. comm.) made
more intensive surveys on the biology of this species in Miyagi-ken, Tohoku District.
According to them, this species is bivoltine, and nests in crevices of wood and inscriptions
of gravestones. In Miyagi-ken the female wasp of the fall (2nd) generation (late August to
mid November) constructs cask-shaped mud cells in a mass in a depression. One to nine
cells (n=96, m=2.8) are constructed, and are finally covered with mud (Goukon, 1983).
Prey menu consists mainly of gelechiid larvae (Lepidoptera).
Parasitoids: an ichneumon wasp (Hymenoptera) and Chrysis ignita-group (Hymenoptera, Chrysididae) (Goukon, 1983c).
Ancistrocerus parietinus (Linnaeus)
(Figs. 207, 217, 224)
Vespa parietina Linnaeus, 1761, Fauna Sued. Ed. 2: 418 (type loc.: "einheimisch"[Swedenl; type
destroyed).
Ancistrocerus parietinus: Thomson, 1874, Opusc. Entomol. 2: 70; B1iithgen, 1961, Faltenwespen
Mitteleuropas, p. 154 (in key), 173-176; Vecht and Fischer, 1972, Hym. Cat. (n. ed.) 8: 115-116.
Japanese name: Yotsuboshi-suji-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 9.0-10.5 mm. Forewing length: 9.0-10.0
mm. Structurally similar to the preceding three species, especially to A. japonicus.
Punctation on head and thorax finer than in japonicus. The apical emargination of clypeus
slightly narrower; lateral teeth slightly more produced (Fig. 207). In color pattern,
however, this species is very distinctive (Figs. 207, 217).
Black, with the following parts yellow: two pairs of spots on clypeus, triangular basal
marking on mandible, frontal spots, short line on temple, antennal scape below, medially
narrowed pro notal band, anterior and posterior spots on tegula, two spots (always
isolated) on scutellum, regular apical bands on gastral tergites 1-5 (band on tergite 5
shortest, sometimes lost), a spot on tergite 6, a complete band on sternite 2, posterolateral
corners of stemite 3 (this sternite rarely possesses an apical band), apical half of fore tibia
anteriorly, tibiae of all legs extensively. Antennal flagellum below ferruginous. Tarsi of all
legs more or less brownish.
Male unknown in Japan.
Material examined. Hokkaido: 1!j?, Hattaribetsu, 2 vii 1965, 1!j?, Vryil, 5 vii 1935(?) (Okada), 2 -'?- !j?,
Nukabira, 14 vii 1959 (TN), 1-'?-, S6unkyo, 27 vii 1971 (TN), 1-'?-, Higashiyama, Furano, 24 vi 1974 (SKY), 1!j?,
108
Osashima (date & collector not indicated).
Distribution. Hokkaido. C. Asia to Europe.
Taxonomic notes. This may be the first record of this species from eastern Asia.
Giordani Soika (1982) pointed out that A. mongolicus (Kostylev) from MongOlia and Korea
may belong to the A. parietinus group.
Biology. Nesting behavior is not known in Japan. Bliithgen (1961) gives a brief
account of the biology of this species in Europe. The female wasp nests in reed tubes and
man-made concavities around houses. Caterpillars of microlepidopterans, and also larvae
of Chrysomelidae (Coleoptera) are hunted as prey.
Ancistrocerus nigricornis (Curtis)
(Figs. 186, 191, 195, 200, 208, 209, 218, 225)
Odynerus nigricarnis Curtis, 1826, Brit. Entomol. 3: 137b, no. 8 (type loc.: England).
Odyneru.s callosus Thomson, 1870, Opusc. Entomol. 2: 87 (type loc.: Sweden); Yasumatsu, 1938, Ins.
Matsum. 13: 15 (from Sakhalin).
Andstrocerus nigricornis: Vecht and Fischer, 1972. Hym. Cat. (n. ed.) 8: 114; Giordani Soika, 1986, Boll.
Mus. Civ. Stor. Nat. Venez. 35: 152, 153.
Japanese name: Ezo-suji-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 8.0-9.5 mm. Forewing length: 7.0-10.0
mm. Head as wide as high, densely punctate. Clypeus wider than high, strongly punctate;
punctures tend to run into striae; apical emargination very shallow and its lateral teeth
round (Fig. 208). Frontal keel rather sharp. Depression for cephalic foveae ill defined;
foveae very small, close to each other, with reddish short hairs. Anterior ocellus as large as
posterior ocellus. Temple rather developed, as seen from above wider than the superior
lobe of eye. Thorax densely punctate. Pronotal carina well developed, but evanescent on
the dorsal part of pronotum; frontal corners of pronotum distinctly produced. Prescutal
groove ill defined; epicnemium with sparse but distinct punctures. Metanotum with a
narrow anterior horizontal part, then sloping toward propodeum; metapleuron above
superficially punctate, below impunctate and not shining. Propodeum without shelf;
posterior face (concavity) obliquely striate, with a vertical carina; superior ridge often
reduced or virtually absent; posterolateral corners produced into processes; dorsolateral
face with large punctures; lateral face above with slightly smaller punctures. Punctation
on gastral tergites 1-5 much finer than on thorax, but coarser than in japonicus, trifasciatus
and melanocerus, especially on tergites 1 and 2. Sternite 2 distinctly angulate near the base
(Fig. 200), sparsely punctate; apical halves of sternites 3-5 also punctate. Last tergite and
sternite impunctate.
Black, with the following parts yellow: two pairs of spots on clypeus (one located in
upper portion, the other near the apex; the latter very often lost, and clypeus rarely wholly
black)(Fig. 208), frontal spot, antenna I scape below, spot on mandibular base (often lost),
small spot on temple, medially narrowed pronotal band that does not extend to the lateral
part of pronotum, anterior and posterior spots on tegula, small spot on dorsal
mesepisternum, a pair of small spots on scutellum, apical band on gastral tergite 1 (often
laterally dilated), wider and regular apical band on tergite 2, narrower apical bands on
tergites 3 and 4 (Fig. 218), sinuated apical bands on sternites 2 and 3, posterolateral
corners of sternite 4 (often lost), apical half of fore femur anteriorly, tibiae of all legs
109
extensively. Mandible blackish brown, apically rufous. Antennal flagellum ferruginous
below. Coxae and femora largely black; tarsi brownish, sometimes partly tinged with
yellow.
Male. Body length (h+th+t1+2): 6.0-7.0 mm. Forewing length: 6.0-7.5 mm. Differs
from the female in the following points: clypeus slightly higher than wide (when
excluding lateral lobes)(Fig. 209), wholly yellow; front corners of pronotum more acutely
produced (Figs. 195 vs. 196); supraclypeal area with a yellow line along eye; mandible
largely yellow; scutellum wholly black; fore and mid coxae and fore and mid femora
yellow on anterior face; tarsi extensively yellowish. Antennal hook medium-sized,
apically rather pointed (Fig. 186).
Material examined: Hokkaido: 1 ~, Maruyama, Sapporo, 7 v 1906, 1 ~, Sapporo, 26 v 1908 (5.
Matsumura), 1 ~, Jozankei, 18 v 1910 (5. Matsumura), 1 ~, Sapporo, 25 v 1910 (5. Matsumura), 1 ~, Jozankei,
18 v 1910 (5. Matsumura), 1!f-, Moiwa, Sapporo, 13 vi 1911 (5. Matsumura), 1!f-, same loc., 20 vi 1912 (5.
Matsumura), 1~, Jozankei, 3 vii 1912 (5. Matsumura), 16', Sapporo, 14 x 1917 (5. Matsumura), 1 6', Tokachidake, 1 viii 1939 (T. Sawamoto), 1!f-, Jozankei, 5 vi 1938 (Y. Sugihara), 1!f-, Sapporo, 25 viii 1935 (Y.
Suiihara), H-, Maruyama, Sapporo, 15 ix 1929 (c. Watanabe), H-, same loc., 18 v 1930 (T. Uchida), 2 ~ ~,
Uryu, 5 vii 1935 (Okada), 1!f-, Sapporo, 21 vii 1943 (S.F. Sakagami), 1!f-, same loc., 4 v 1947 (S.F. Sakagami), 1
~, Maruyama, Sapporo, 25 ix 1955 (K. Kamijo), 1~, Jugoshima, Sapporo, 29 x 1962 (SKY), 1!f-, Miyanomori,
Sapporo, 30 v 1965 (SY), 1~, Heiwa, Sapporo, 27 v 1967 (K. Hoshii), 1~, Bibai, 29 v 1972 (M. Taira), 16',
Jozankei, 9 ix 1972 (YH), 1~, Hoheikyo, Sapporo, 23 v 1973 (T. Okazawa), 2~ -,?-, Hakken-zan, Sapporo, 11 v
1975 (H. Taoka), 1-'?-, same loc., 23 v 1976 (H. Taoka), 1-'?-, Moiwa, Sapporo, 10 v 1977 (T. Kumata), 1-'?-, same
loc.,8 vi 1979 (1. Yasui), 2!f-!f-, Koshunai, Bibai, 6--8 vi 1978 (SKY), 2!f-!f-, Mizuho, Higashi-asahigawa, 15-19
v 1978 (SKY), 1-'?-, Moiwa, Sapporo, 23 iv 1978 (M. Morl), 2-'?-~, Toyotaki, Sapporo, 24 v 1978 (SKY), 1-'?-,
Hakken-zan, Sapporo, 24 vi 1978 (SKY), 46'6'2!f-~, Heiwano-taki, Sapporo, 23 ix 1978 (SKY), 3!f--'?-,
Takisato, 10 v 1979 (H. Fukuda), 1!f-, Toyotaki, Sapporo, 30 v 1979 (T. Fujisawa), 2!f- -,?-, Asahigawa, 3-4 vi
1979 (T. Inaoka), 3-'?- ~, Toyotaki, Sapporo, 30 v - 8 vi 1979 (SKY), 1!f-, Kariba--dake, 14 vii 1979 (T. Fujisawa),
1-'?-, Furano, 13 vi 1979 (H. Fukuda), 1!f-, Otarunai, vii 1979 (T. Fujisawa), 1-'?-, Asahigaoka, Sapporo, 17 ix
1979 (SKY), 1-'?-, Toyotaki, Sapporo, 30 v 1980 (SKY), 1-'?-, same loc., 21 vi 1980 (5. Makino), 1-,?-, Misumai,
Sapporo, 23 viii 1983 (5. Makino), 26' 6', same loc., 11 ix 1983 (5. Makino), 1!f-, Toyotaki, Sapporo, 7 vi 1986
(5. Makino), 2-'?- -,?-, Nishino, Sapporo, 5 vii 1986 (U. Kurosu), 1-'?-, Maruyama, Sapporo, 11 vi 1986 (M. Sato).
Honshu: Aomori-ken -1-'?-, Kuroishi, 3 v 1935 (I. Tateyama), 1-'?-, Yamagata, 31 v 1936 (I. Tateyama).
Distribution. Hokkaid6; HonshU (T6hoku District). Sakhalin; Europe.
Biology. No information is available for the Japanese population. According to the
collection data, this species seems to have at least two generations, but males are collected
only in August - October.
In central Europe, the second generation females of this species are known to
hibernate and appear in early spring next year, then prepare their nests that produce both
sexes in June to mid-July (lst generation); the females of the first generation again build
nests that produce both sexes (mid-August to September), then inseminated females
overwinter. The female wasp nests in pre-existing cavities in wood, stones and metal, and
hollow plant stems (Nielsen, 1932, referred to as A. caliosus; summarized in Bliithgen,
1961).
Ancistrocerus oviventris oviventris (Wesmael)
(Figs. 187, 189, 190, 197, 210,211,219,224)
Odynerus oviventris Wesmael, 1836, Bull. Acad. R. Belg. 3: 45, pI. 2, fig. 1 (!f- 6')(type loc.: Bruxelles,
Belgium).
Ancistrocerus oviventris oviventris: B1iithgen, 1961, Faltenwespen Mitteleuropas, pp. 179-180; Vecht and
110
Fischer, 1972, Hym. Cat. (n. ed.) 8: 115.
Andstrocerus yamanei Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venezia, 35: 14S-149, fig. 45 (.!f)
(type loc.: Tokyo). Syn. nov.
Japanese name: Kenaga-suji-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 9.5-10.0 mm. Forewing length: 9.0-10.0
mm. Head and ali trunk with long and dense hairs; hairs on vertex distinctly longer than
the distance between the posterior ocelli; antennal scape, especially near the base, with
many long hairs (Fig. 189); gastral tergites 1-5 with long hairs. Head slightly wider than
high. Punctures on frons and eye sinus relatively small, but very dense; punctation on
gena much sparser. Clypeus much wider than high, with sparse punctation, somewhat
shining, apically narrowly but relatively deeply emarginate; lateral teeth of this
emargination round (Fig. 210). Distance between the posterior ocelli distinctly shorter
than that between posterior ocellus and eye. Depression for cephalic foveae ill defined;
foveae very small and close to each other. Temple well developed, as seen fom above,
much wider than the superior lobe of eye. Thorax stumpy, densely punctate. Punctation
on lateral face of pronotum irregular in shape, and spaces between punctures running into
irregular carinae in lower portion of this part. Prescutal grooves almost invisible.
Punctures on mesopleuron larger than on mesoscutum, ill defined; interspaces running
into carinae. Metanotum with a flat anterior part that is slightly elevated above the level of
the posterior part of scutellum; metapleuron posteriorly not demarcated by a suture from
propodeum, superficially striate above, smooth but dull below. Propodeum without shelf,
with a vertical part between metanotum and superior ridge. Propodeal concavity very
weakly and obliquely carinate, with a complete median keel; superior ridge often much
reduced, especially in mesal part; inferior ridge developed, with distinct posterolateral
angle. Dorsolateral part of propodeum without distinct punctures; lateral ridge not
developed; lateral face almost impunctate and dull, in upper portion with weak carinae.
Punctures on gastral tergite 1 relatively large, but ill defined and interspaces often larger
than punctures; punctation on other tergites much finer and sparser. Punctation on
sternite 2 fine and sparse; that on the apical half of other sternites slightly coarser.
Black; yellow are: two pairs of spots on dypeus (upper two are large and lower two
small; all are lost in the specimens from Hokkaido), triangular marking on mandibular
base, minute spot on each temple, frontal spot, narrow pronotal band (only slightly
dilated laterally), an outer band on tegula, a transverse marking on scutellum (narrowly
interrupted in the middle), regular apical bands on tergites 1-5 (Fig. 219), apical margins of
sternites 1 and 2, posterolateral corners of sternites 3 and 4 (in the specimens from
Honshu, also sternite 3 with apical band), spot on tergite 6, apical part of fore femur
anteriorly, tibiae of all legs extensively. Antenna almost wholly black. Tarsi of all legs
brownish.
Male. Body length (h+th+t1+2): 7.5-9.0 mm. Fore wing length: 7.0-8.5 mm. Much as in
the female from which it differs in the following points: clypeus very deeply emarginate at
apex; lateral teeth of the emargination long and sharp (Fig. 211); inner margin of mandible
with a large concavity in the middle (Fig. 190); front corner of pronotum produced into a
process (Fig. 197); dypeus, labrum and mandible largely yellow; antennal scape below
yellow; frontal spot smaller; scutellum without yellow; last tergite without yellow spot;
apical parts of femora of all legs, tibiae of all legs almost wholly yellow; tarsi also marked
with yellow; 2-3 terminal segments of antenna ferruginous. Antennal hook with the apex
111
almost reaching the base of segment 11 (Fig. 187).
Material examined. Hokkaido: 1 ~,Sapporo (date and collector not indicated), 1 ~, Jozankei, 24 vi 1951
(K. Kamijo), 1~, Hakken-zan, Sapporo, 24 vi 1978 (SKY), 1 d', Toyotaki, Sapporo, 30 v 1979 (SKY), 1 d', same
loc., 7 vi 1986 (5. Makino).
Honshu: Tokyo-to - 1 ~, Kobotoke, Minamitama, 29 iv 1950 (K. Harada) (holotype of A. yamanei);
Saitama-lcen - 1 ~, Kodama, 22 iv 1965 (TN), 2 ~ ~, same loc., 8 v 1965 (TN).
Distribution. Hokkaid6; Honshu (Kant6 District). Eastern Asia to central and northern
Europe.
Taxonomic notes. This rare species is easily distinguished from other Japanese
congeners by the long body hairs, widely and deeply emarginate clypeal apex in the male,
median concavity on inner margin of male mandible, etc. There are constant differences in
color pattern between the populations of Hokkaid6 and Honshu. In the specimens from
Hokkaid6, clypeus wholly black and sternite 3 without yellow apical band, while those
from Honshu have the clypeus with four yellow markings and the sternite 3 with a yellow
apical band.
I have examined the holotype ( -'f ) of A. yamanei through the courtesy of Dr. Giordani
Soika. It was compared with A. parietinus by him to show constant differences (Giordani
Soika, 1986), but actually it well agrees with A. oviventris.
Biology. Nothing is known of the nesting biology of this species in Japan. In Europe,
this wasp is univoltine and constructs its mud nests on depressions of stones, etc.; the nest
often consists of more than ten cells in a mass (Nielsen, 1932). The female wasp hunts for
caterpillars of Microlepidoptera; some authors reported the larvae of snout beetles and
leaf beetles as prey of this wasp (see Bliithgen, 1961).
Ancistrocerus antilope antilope (Panzer)
(Figs. 188, 198, 212, 213, 220, 225)
Vespz antilape Panzer, 1798, Fauna Ins. Germ. 5(53): 9, tav. 9 (~)(type loc.: Austria).
Ancistrocerus antilope: BHithgen, 1943, Stet!. Ent. Ztg. 104: 156; Yasumatsu, 1938, Ins. Matsum. 13: 15;
Vecht and Fischer, 1972, Hym. Cat. (n. ed.) 8: 108-109; Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez.
35: 152.
Japanese name: Tsuya-suji-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 10.5-11.5 mm. Fore wing length: 11.0-11.5
mm. Head nearly as wide as high, densely punctate on frons and sinus of eye; punctation
on gena sparser. Clypeus much wider than high, narrowly and shallowly emarginate at
apex (Fig. 212), sparsely punctate. Distance between the posterior ocelli shorter than that
between posterior ocellus and eye. Temple rather developed, much wider than the
superior lobe of eye. Depression for cephalic foveae large but ill defined; foveae small,
close to each other. Thorax moderately punctate; spaces between punctures shining.
Prescutal grooves visible. Punctation on ventral mesepisternum sparse; dorsal
mesepisternum posteriorly and mesepimeron irregularly striate. Metapleuron not clearly
separated from propodeum by a suture, almost impunctate, with glaring luster.
Propodeum virtually without shelf. Propodeal concavity irregularly and obliquely striate,
with a median keel; superior ridge present but less developed than inferior ridge.
Dorsolateral face of propodeum irregularly reticulate; lateral ridge present; lateral face
smooth and shining below, irregularly striate above. Horizontal part of gastral tergite 1
112
moderately punctate; tergite 2 with very fine punctures apically; tergites 3-5 with slightly
larger and ill-defined punctures in apical half; tergite 6 with a preapical depression, with
ill-defined punctures around it. Punctation on sternites finer than on tergites.
Black, the following parts yellow: a pair of longitudinal markings on clypeus near its
base (in the specimens from Hokkaid6, these are much reduced), small frontal spot,
minute spot on temple, basal marking on mandible, antennal scape below, a pair of
triangle markings on pronotum anteriorly, regular bands on gastral tergites 1-3 and
sternite 2, small spot on last tergite (Fig. 220), posterolateral corners of sternite 3, apical
part of fore femur anteriorly; anterior faces of tibiae of all legs. Tarsi reddish or blackish
brown. Antennal flagellum below ferruginous.
Male. Body length (h+th+t1 +2): ca. 10 mm. Fore wing length: ca. 10 mm. The male
differs from the female as follows: head wider than high; clypeus nearly as wide as high;
front corners of pronotum bluntly produced (Fig. 198); mesopleuron much more coarsely
punctate; scutellum with a longitudinal median groove; lateral ridge of propodeum much
more developed so as to form a wing; clypeus and mandible largely yellow; frontal spot
absent; antennal flagellum orange yellow below; tergite 4 with a narrow apical band;
sternite 3 with an apical band; mid and hind coxae and mid femur with yellow; tibiae
more extensively yellow. Antennal hook very small; its apex not reaching the middle of
segment 11 (Fig. 188).
Material examined. Hokkaido: 1 ~, Moiwa, Sapporo, 23 vi 1932 (T. Uchida).
Honshu: Fukui-ken -1 6'2 ~ ~, Ashuma, Fukui~hi, 10-21 v 1959 (IT).
Distribution. Hokkaid6; Honshu. Holarctic. Among the Eumenidae, this is the only
Japanese species that is Holarctic in distribution.
Biology. Nesting biology is not known in Japan. In Europe and North America, this
wasp is univoltine and nests in deserted mud cells of other wasps, tunnels dug in wood
by larvae of longicorn beetles and wood wasps, and also in reed tubes (Bliithgen, 1961).
This species has a quite wide prey range: caterpillars of lepidopterous families
Oecophoridae, Gelechiidae, Olethreutidae, Tortricidae, Pyralidae, Noctuidae (Krombein,
1979), and Hesperidae (Bliithgen, 1961); larvae of Nematus ericksoni (Hymenoptera,
Tenthredinidae) (Ashmead, 1894); larvae of leaf beetles (Coleoptera, Chrysomelidae) (in
Finland; cf. Bliithgen, 1961). Krombein listed parasitoids found in nests of this species in
North America.
Genus Symmorphus Wesmael
Symmorphus Wesmael, 1836, Bull. Acad. R. Belg. 3: 45 (subgenus of Odynerus Latreille) (type species:
Odynerus elegans Wesmael, 1833, deSignated by Richards, 1935); Dalla Torre, 1904, Gen. Ins. 19: 35, 36
(subgenus of Odynerus Latreille).
Protodynerus Saussure, 1855, Et. Fam. Vesp. 3: 184, 186,352 (new name for Symmorphus Wesmael).
Japanese names: Hamushi-dorobachi Zoku (Hoso-dorobachi Zoku).
Body usually slender; but some large species such as S. captivus and others resemble
wasps of Ancistrocerus in body shape. Female clypeus usually wider than high, apically
narrowly emarginate (in s. decens widely truncate), with more or less acute lateral teeth.
Male antenna with the terminal segment of normal shape (not forming a recurved hook).
Thorax usually duplipunctate, i.e., with macropunctures of normal type and micropunctures. Mesoscutum with distinct notaulices. Epicnemial carina usually present (absent in
113
S. foveolatus). Propodeum often with a shelf behind metanotum; the shelf medially with a
pit of varying size. Gastral segment 1 distinctly narrower than segment 2, but not petiolate. Tergite 1 strongly punctate on posterior horizontal disc, with a transverse carina at
the basal border of the disc; the disc also with a longitudinal furrow; anterior face of the
tergite sometimes with a short median carina. Tergites 2-6(7) usually impunctate or only
weakly punctate.
Most species of this genus hunt chrysomelid larvae (Coleoptera) for their young. But
at least three species (including one Japanese species) hunt only larvae of Curculionidae,
and some species only small lepidopterous larvae. The Nearctic S. canadensis (Sauss.)
hunts larvae of various coleopteran families (Iwata, 1971). The biology of the Japanese
species should be entirely resurveyed, because the taxonomy of these wasps had been
quite confused and only two species had been recognized until Giordani Soika (1975)
published a revision.
Key to the Japanese species
An excellent key to the Japanese species was given by Giordani Soika (1975) who
recognized seven species (the key was translated into Japanese by Tsuneki, 1976). Tsuneki
(1977, 1986) added two new species and one new subspecies of S. mutinensis to the fauna
of Japan, and I have found three additional forms which will be described as new species
below. In Japan this genus contains several closely related species which are very difficult
to separate. Here, a practical key using mainly color pattern is presented. It is strongly
advised to use this key together with Giordani Soika's that refers more frequently to
structural characters but does not include all the forms dealt with in the present paper. In
the following key the punctation concerns macropunctation alone.
1. Epicnemial carina absent. Gastral tergite I, seen from above, longer than wide at its
apical margin .................................................................................................... S. foveolatus Gus.
- Epicnemial carina present. Gastral tergite I, seen from above, as long as or shorter than
wide ............................................................................................................................................... 2
2. Clypeus widely truncate apically. Mesopleuron densely and uniformly punctate.
Tergites 1,2, (3), 4, 5 each with a yellow apical band (.!f.) ........................... S. decens (Kost.)
- Clypeus narrowly emarginate apically. Punctation on mesopleuron various, but in most
species weaker and sparser. Tergite 5 without yellow band ( .<f- )......................................... 3
3. Large species; h+th+t1+2: 12-13 mm in .<f-,9-10 mm in t. Body shape of Ancistrocerustype ............................................................................................................................................... 4
- Smaller species; h+th+t1+2: less than 9 mm in .<f-, usually less than 7 mm in t. Body
much more slender. Female clypeus wholly black. Ventral mesepisternum impunctate or
only weakly and sparsely punctate (rather strongly punctate in S. iwatai) ......................... 5
4. Clypeus with a basal yellow marking (.<f-). Antennal scape and flagellum wholly
blackish ( .!f. )....................................................................................................... S. captivus (Sm.)
- Clypeus wholly black ( .<f-). Antennal scape and flagellum ferruginous yellow below
(.<f-) ............................................................................................................... S. sounkionis Tsuneki
5. Antenna 13-segmented. Gaster with 7 visible segments............... Males ........................... 6
- Antenna 12-segmented. Gaster with 6 visible segments ............... Females ...................... 13
6. Antennal scape with a yellow spot or stripe. If wholly black, apical segments of
114
antenna below largely ferruginous ........................................................................................... 7
- Antennal scape wholly blackish ................................................................................................ 9
7. Clypeus basally yellow. Pronotum anteriorly with a pair of large triangular yellow
markings. Tergite 3 with a yellow apical band. Apical lamella of tergite 1 medially
produced backward ............................................................................... s. tsushimanus sp. nov.
- Clypeus nearly wholly yellow. Pronotum wholly black, at most with a pair of very
minute orange spots. Tergite 3 without yellow band. Apical lamella of tergite 1 normal.
......................................................................................................................................................... 8
8. Apical segments of antenna much thickened; undersides of these segments extensively
ferruginous. Mandible largely yellow................................................... s. apiciomatus (Cam.)
- Apical segments of antenna moderately thickened, with ferruginous stripes below.
Mandible at most with a small yellow marking ..................................... S. carinatus sp. nov.
9. Clypeus nearly wholly yellow. Pronotum with a pair of yellow spots. Metapleuron and
lateral face of propodeum finely striate ........................................................ S. iwatai sp. nov.
- Clypeus wholly black or with an irregular yellow marking. Pronotum without yellow
marking. Metapleuron and lateral face of propodeum more coarsely striate or strongly
punctate ...................................................................................................................................... 10
10. Mesopleuron strongly punctate. Mesepimeron reticulate. Scutellum with large
macropunctures .......................................................................................... S. mutinensis (Bald.)
- Mesopleuron shining, at most finely and sparsely punctate. Mesepimeron not reticulate.
Punctures on scutellum much smaller................................................................................... 11
11. Posterior part of metanotum transversely striate or corrugated. Transverse carina on
tergite 1 not incised medially. Mesoscutum very finely and sparsely punctate. Propodeal
pit very large .............................................................................................. S. mizuhonis Tsuneki
- Posterior part of metanotum not striate. Transverse carina on tergite 1 incised medially.
....................................................................................................................................................... 12
12. Mesoscutum and metanotum rather strongly punctate. Dorsal mesepisternum with
punctures. Propodeal pit very small ..................................................................... S. diens G.S.
- Mesoscutum and metanotum with much finer punctation. Dorsal mesepisternum
impunctate. Propodeal pit large ...................................................................... S. ishikawai G.S.
13. Cephalic foveae as large as ocelli. Mesoscutum with very sparse and superficial
punctation. Mesosoma wholly black. Tergites 1 and 2 with yellow apical bands .
............................................................................................................................... S. ishikawai G.S.
- Cephalic foveae distinctly smaller than ocelli. ...................................................................... 14
14. Propodeal shelf very narrow and indistinct; median pit inconspicuous. Metapleuron,
and lateral and posterior face of propodeum finely striate, usually without luster.
Posterior half of metanotum dull. Mesosoma wholly black. ............. S. apiciomatus (Cam.)
- Propodeum with a distinct (but sometimes slightly sloping) shelf, which has usually a
large median pit (in dims the pit is often a narrow slit) ...................................................... 15
15. Dorsal mesepisternum with large punctures. Mesepimeron strongly punctate or
reticulate. Mesoscutum often with a pair of yellow spots ................... S. mutinensis (Bald.)
- Dorsal mesepisternum and mesepimeron at most with small punctures sparsely.
Mesoscutum always wholly black. ......................................................................................... 16
16. Posterior part of mesopleuron, metapleuron wholly, and lateral face of propodeum
microscopically striate or shagreened, dull. Pronotum anteriorly with a pair of yellow
spots. Anterior vertical face of tergite 1 with a strong vertical carina ................................... .
.......................................................................................................................... S. carinatus sp. nov.
115
- Posterior part of mesopleuron, metapleuron and lateral face of propodeum at least
partly shining; striation, if any, much coarser. Pronotum entirely black. Anterior vertical
face of tergite 1 without carina ................................................................................................ 17
17. Median pit on propodeal shelf large and round. Cephalic foveae very small. Dorsal
mesepisternum often with a yellow spot. Transverse carina on tergite 1 not incised
medially. Tergite 4 without yellow apical band .................................... S. mizuhonis Tsuneki
- Median pit small and often narrow. Cephalic foveae larger, more than half as large as
posterior ocelli in diameter. Thorax usually wholly black. Transverse carina on tergite 1
medially incised. Tergite 4 often with an incomplete yellow apical band ....... S. dims G.S.
Symmorphus foveolatus Gussakowskij
(Figs. 226, 227, 236, 246, 256, 277)
Symmorphus faveolatus Gussakowskij, 1933, Ark. Zoo!. 24 A (10): 55 (i'-) (type loc.: Ussuri); Giordani
Soika, 1963, Boll. Soc. Entomo!' Ita!. 93: 124; 1!175, Boll. Mus. Civ. Venez. 27: 149 (in key), 153; Vecht and
Fischer, 1972, Hym. Cat. (n. ed.), 8: 121.
Odynerus (Symmorphus) CIlptivus: Yasumatsu, 1938, Fukuoka Hakub. Zasshi, 2: 11, tab. 3, figs. 1-5
(part?); 1950, Icon. Ins. Jpn. 2nd ed. p. 1457, fig. 4203 (part?) (misidentification).
Japanese names: Haranaga-hamushi-dorobachi (Sumisu-hamushi-dorobachi).
Dwgnosis. Female. Body length (h+th+t1+2): 7.0-10.5 mm. Fore wing length: 7.5-10.0
mm. Head nearly circular in frontal view; macro punctures on frons not very dense,
smaller than interspaces; punctation on gena and vertex much finer and sparser. Cephalic
foveae slightly smaller than posterior ocelli, each situated in a depression; the depressions
not connected to each other by a furrow. Clypeus distinctly wider than high, very finely
punctate, narrowly and shallowly emarginate at apex (Fig. 226). Interantennal area
roundly swollen, with a Y-shaped carina which is often obscure. Distance between the
posterior ocelli slightly shorter than that between posterior ocellus and eye (9:10).
Flagellar segment 1 of antenna slightly longer than wide at its apex. Thorax with
macro punctures that are usually as large as or smaller than interspaces which are shining.
Front corners of pronotum round. Mesoscutum with complete notaulices; parapsidallines
very weak; median scuta I line visible only in anterior third. Scutellum with a weak
median furrow. Mesopleuron without epicnemial carina (this condition is seen only in this
species among the Japanese species). Metanotum with large macropunctures, impunctate
and dull in its posterior 1/ 4 to 1/3; metapleuron above with a few macropunctures, below
micropunctate and without macropunctures. Dorsolateral parts of propodeum with large
and dense macropunctures; propodeal shelf narrow, sloping posteriorly, with a large
median pit (Fig. 256); propodeal concavity with fine and dense striae running obliquely,
and with a complete median vertical carina. Lateral face of propodeum macro punctate
(punctation often superficial); spaces between punctures micropunctate and with
inconspicuous striae. Superior and inferior ridges of propodeum poorly developed, often
almost absent. Gastral tergite 1 seen from above longer than wide at its apex, gradually
widened toward apex, strongly punctate, apically slightly duplicate, with a preapical
swelling; each lateral end of the swelling bearing a short carina running toward the base
of the tergite but not connected with the transverse carina, which is not sharply defined
and not incised in the middle; anterior face of tergite 1 coarsely and irregularly punctate.
Tergite 2 shining, weakly punctate at base; other tergites almost impunctate. Sternite 1
116
235
227
234
230
\
I
'\
I
/
,/
232
Figs. 226-235. Clypeus of Japanese Symmorphus. 226, foveolatus !f; 227, ditto t; 228, deeens !f; 229,
ditto t; 230, captivus !f; 231, ditto t; 232, carinatus !f; 233, ditto 6'; 234, iwatai t; 235,
tsushimanus
t.
irregularly and strongly reticulate; sternites 2-6 almost impunctate.
Black, with the following parts yellow: two small frontal spots (sometimes connected
to form a single spot, or completely lost), a minute spot on temple, an apical band on
tergite 1 medially dilated and at the middle incised, apical bands on tergites 2, 4 and
sternite 2 (the band on sternite 2 sinuated or medially interrupted), anterior face of fore
(rarely also mid) tibia. Mandible ferruginous in apical 1/3. Antennal flagellum below and
legs slightly brownish.
Male. Body length (h+th+t1+2): 7.5-8.0 mm. Fore wing length: 7.5-8.0 mm. Clypeus
more deeply emarginate at apex than in the female (Fig. 227; lateral angles of the
emargination acute), almost wholly yellow. Yellow frontal marking completely lost. Dorsal
117
mesepisternum sometimes with a small yellow spot. Apical band on tergite 1 narrower
and more regular than in the female; tergite 5 also with an apical band. Tibiae of all legs
extensively yellowish. Flagellum almost wholly black, without distinct tyloids; last
segment longer than wide at its base, as long as the preceding segment (Fig. 236).
Material examined. Hokkaido: 1 (1', Garugawa, 22 vi 1922 (H. Kono), 1 (1', same loc., 8 vi 1925 (H. Kono),
1 ~, Sapporo, 1 vii 1955 (K. Kamijo), 1 ~, same loc., 3 vii 1965 (T. Kocha), 1 ~, Gamushi, 12 vii 1958, 1 ~,
Otarunai (Helvetien Hiitte), 26 vii 1965 (SY), 1 ~, Shikotsu-ko, 17 vii 1977 (SKY), 1 ~, Mt. Teine (7-800 m alt.),
13 viii 1978 (SKY).
Honshu: ADmori-ken - 1 ~, Osore-zan, 12 viii 1979 (K. Goukon); Yamagata-ken - 1 ~, Oguni, 25 vi 1980
(HI), 1 ~, Nan'yo, 13 vii 1980 (HI); Miyagi-ken - 1 (1', Rifu, 29 v 1983 (K. Goukon), 1 (1', TagajO, 4 vi 1983 (K
Goukon); Niigata-ken -1 ~, Numa, Sekikawa, 10 vii 1966 (HI), 6~ ~, Shibata, 12 vii 1981 (HI); Nagano-ken -1
~, Komaga-take, Ina, 10 viii 1%2 (K Oshima); Saitama-ken -1 ~, Izuga-dake, 3 vii 1984 (5. Aoki); Tokyo-to-l
(1' (5. Hirayama); Fukui-ken -1 ~, Hatogayu, 15 vii 1956 (K Iwata).
Shikoku: K6chi-ken - 1 ~, Kodakasa-yama, K6chi-shi, 6 v 1930 (Y. Sugihara), 1 ~, same loc., 5 vii 1931 (Y.
Sugihara), 1 ~, Mt. Washio, 14 v 1933 (Y. Sugihara).
Distribution. Hokkaid6; Honshu; Shikoku; Kyushu. E. Siberia. Yasumatsu (1938a)
recorded "Odynerus captivus" also from Korea.
Taxonomic notes. Yasumatsu's (1938a) material contained very small males with fore
wing 6.0 mm in length. I have never seen such a small specimen of this species. It is quite
possible that there were included males of some other species in his material.
Biology. Iwata (1938b) reported the biology of "Odynerus captivus". Yasumatsu's
(1938a) material contained several specimens of "0. captivus" collected by Iwata in Settsu
(Osaka). The wasp studied by Iwata and identified by K. Yasumatsu was most probably S.
foveolatus as pointed out by Giordani Soika (1975). Here I summarize Iwata's account on
the biology of "0. captivus". The wasp flies from May to July, and has only one generation
in a year. The female nests in wheat straws and slender tubes of bamboo and reed which
are bundled to make roofs and blinds. Rarely deserted beetle burrows in wood may be
utilized. The nest consists of 1 to 6 brood cells (usually 1-3; the cell measures 21 to 45 mm
in length and 3 to 6 mm in width), and 1 to 3 empty cells. Cell partitions (0.5-1.0 mm thick)
and entrance plug (2-5 mm thick) are made of mud. Brood cells are each provisioned with
4-31 beetle larvae (4-8 mm long); the number of beetle larvae stored in a cell depends on
their body size. In Osaka the full-grown larvae of two chrysomelids, Gastroidea atrocyanea
on Rumex and Plagiodera versicola on Salix, are mainly captured. Additional accounts on
the nesting biology of this species are given in Iwata (1978b) and Tsuneki (1973a).
Parasitoids: Chrysis galloisi and C. sarafschana rubripyga (Hymenoptera, Chrysididae)
(Tsuneki, 1973a).
Symmorphus decens (Kostylev)
(Figs. 228,229,237,247,257,278)
Odynerus deeens Kostylev, 1940, Bull. Soc. Nat. Moscou, Sect. Bio!. NS, 49 (5/6): 40
(~)
(type loc.:
Minoo, Osaka).
Odynerus apiciornatus: Yasumatsu, 1938, Fukuoka Hakub. Zasshi, 2: 111, 114, tab. 3, figs. 6-9.
Symmorphus decens: Giordani Soika, 1975, Boll. Mus. Civ. Venezia, 27: 154-156, figs. 12, 13.
Japanese names: Kuchibiro-hamushi-dorobachi CYamato-hamushi-dorobachi).
Diagnosis. Female. Body length (h+th+t1+2): 7.0-9.5 mm. Fore wing length: 8.5-10.5
mm. Head wider than high, densely macropunctate; punctures relatively small. Cephalic
118
foveae quite small, each located in an elliptical depression; the depressions not connected
to each other. Oypeus much wider than high, apically very broadly truncate; each lateral
end of the truncated margin with a small but distinct denticle (Fig. 228). Distance between
the posterior ocelli as long as ocello-ocular distance. Flagellar segment 1 more than 1.5
times as long as wide at its apex. Thorax very densely macro punctate except on pro notal
tubercle and in anterior and posterior parts of mesopleuron. Front comers of pronotum
round. Notaulices complete, but narrow; parapsidal lines still narrower; median scutal
line visible only in anterior third. Epicnemial carina visible only in its lower part. Macropunctation on metanotum coarser than on scutellum; posterior portion of metanotum
with vertical carinae; metapleuron impunctate, finely striate above. Propodeal shelf
almost absent, with a pit just behind metanotum (Fig. 257). Dorsal face of propodeum
coarsely punctate or reticulate; concavity with a complete median carina, without striae;
inferior ridge visible; superior ridge almost absent; lateral portion of propodeum
macro punctate and striate above. Gastral tergite 1 nearly as long as wide at its apex;
anterior face with a few ill-defined macropunctures, without vertical carina; transverse
carina rather sharply defined, not incised in the middle; posterior horizontal part
distinctly punctate. Tergites 2-6 finely and superficially punctate. Sternite 1 with a strong
median carina and irregularly running carinae; punctation on sternites much finer than on
tergites.
Black, with the following parts yellow: a spot on the base of antennal scape anteriorly
(often reduced), minute spot on temple, a pair of small markings on pronotum anteriorly,
apical bands on gastral tergites 1, 2, 4,5 (band on tergite 2 widest), apical band on sternite
2 medially interrupted, and anterior face of fore tibia.
Male. Body length (h+th+tl+2): 6.0-8.5 mm. Fore wing length: 7.0-8.0 mm. Similar to
&B arB
239
243
E8JJ
240
Figs. 236-245. Apical segments of male antenna in Japanese Symmorphus. 236, foveolatus; 237,
decens; 238, captivus; 239, apiciornatus; 240, carinatus; 241, iwatai; 242, tsushimanus; 243,
mutinensis; 244, mizuhonis; 245, diens.
119
the female in structure and coloration, with the following differences: clypeus largely and
anterior face of mandible in basal 2/3 yellow; yellow marking on antennal scape larger; 45 terminal segments of antenna more or less extensively orange yellow; pro notal spots
often lost; tergite 6 with an apical band; sternite 3 sometimes with yellow spots apically;
fore and mid tibia anteriorly and tarsi of all legs extensively yellowish. Antennal segments
10-13 with ill-defined tyloids; last segment small, as long as wide at its base and shorter
than segment 12 (Fig. 237).
Material examined. Honshu: Tokyo-to -1 J', Kobotoke-toge, 22 v 1946 (S.F. Sakagami); Ishikawa-ken -1 J',
Yoshioka, Kamachi, 14 v 1974 G. Togashi); Osaka-fu -1 J', Minoo, 27 v 1934 (Y. Sugihara); Wakaya7TllJ-ken - 4 J'
J', Amami-toge, 24 iv 1964 (H. Katayama); Hy6go-ken -1-'? ,Sasayama, Tamba, 15 v 1964 (K. Iwata).
Shikoku: K6chi-ken - 1-'?, Engyoji, 15 iv 1930 (Y. Sugihara), 4-'? -'f, Kodakasa-yama. KOchi-shi. 6 v 1930
(Y. Sugihara), 2-'f -'f, same loc., 30 iv 1931 (Y. Sugihara), 1-'f, Yamada, 9 iv 1932 (Y. Sugihara), 1 J', Godai-san,
Kochi-shi, 7 v 1975 (51),1 J', same loc., 6 v 1976 (51).
Distribution. Honshu; Shikoku; Kyushu (Giordani Soika, 1975; Nagase, 1981). E.
Siberia.
Taxonomic notes. I have examined one male and one female from Primorskij, eastern
Siberia (sent by Dr. Kurzenko), which principally agree with S. decens. In the male,
however, the truncated apical margin of clypeus is narrower than in the Japanese
specimens. The female possesses two yellow spots on frons, and the yellow marking on
antennal scape and paired pro notal markings are larger.
Biology. Iwata (1938b) observed the nesting behavior of this species (referred to as
Odynerus apidornatus) in Settsu, Honshu in May and June (1929-32). One or rarely two
brood cells, and one to three empty cells are constructed in wheat straws used as roofs of
cottages. Most nests have an empty cell just inside the entrance plug. Cell partitions (0.5-1
mm thick) and entrance plug (2 mm thick) are made of fine earth or red clay. The brood
cell is provisioned with 4-11 (usually 5-8) mature larvae of a chrysomelid, Phytodecta
rubripennis; the prey measures 5-8 mm in length.
Parasitoid: Chrysis japonicus was seen haunting colonial nesting sites of S. decens
(Iwata, 1938b).
Symmorphus captivus (Smith)
(Figs. 230, 231, 238, 248, 258, 279)
Odynerus captivus Smith, 1873, Trans. R. Entomol. Soc. London, 1873: 197 (-'f )(type loc.: "Hiogo",
Japan); Schulthess, 1934, Arb. Morph. Taxon. Entomol. 1: 66; Giordani Soika, 1941. Boll. Soc. Venez. Stor. Nat.
2:232.
Symmorphus captivus: Giordani Soika, 1975, Boll. Mus. Civ. Stor. Nat. Venez. 27: 150 (in key), 151-153,
figs. 1,7; Tsuneki, 1986, Spec. Publ. Jpn. Hymen. Assoc. 32: 22 (J').
Japanese name: O-hamushi-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): ca. 12.5 mm. Fore wing length: ca. 12.0
mm. Head rather densely punctate on frons and ocular sinus, finely punctate in ocellar
region, and sparsely punctate on gena and vertex. Cephalic foveae only slightly smaller
than posterior ocelli, deep, each situated in a transverse depression; the depressions
connected to each other by a furrow. Clypeus relatively finely and sparsely punctate;
apical emargination relatively wide and its lateral teeth rather acute (Fig. 230). Flagellar
segment 1 of antenna ca. 1.6 times as long as wide at its apex. Distance between the
120
~
246
~249
metanotum
\\ \\
247
\\
250
256~
pit
253
262
248
~251
257 ~
~
258
\\ \\ \\
252
Shelf/~
254
259
~
~
263
264
265
~
~
260
~
261
]J:n::J;£
266~
Figs. 246-255. Front comer (right) of male pronotum in Japanese Symmorphus. 246, foveolatus; 247,
decens; 248, captivus; 249, apiciornatus; 250, carinatus; 251, iwatai; 252, tsushimanus; 253,
mutinensis; 254, mizuhonis; 255, diens.
Figs. 256-266. Shelf and pit of propodeum in Japanese Symmorphus. 256, foveolatus; 257, decens;
258, captivus; 259, apiciornatus, 260, carinatus; 261, iwatai; 262, tsushimanus; 263, mutinensis; 264,
mizuhonis; 265, diens; 266, ishikawai. In foveolatus and deeens the shelf is virtually lacking (in the
figures the area is not defined posteriorly).
posterior ocelli only slightly shorter than ocello-ocular distance (14:15). Pronoturn rather
coarsely punctate; pronotal carina widely interrupted in the middle; front corners of
pronotum not produced. Mesoscutum much finely but densely macropunctate in its
anterior half, but with only a few macropunctures and with numerous micropunctures in
its posterior half; notaulices narrow but complete; parapsidallines obscure; median scutal
line visible only in anterior third. Epicnemial carina strong; epicnemium and dorsal
mesepisternum sparsely macropunctate; ventral mesepisternum and mesepimeron
densely macro punctate but punctures rather superficial. Metanotum coarsely macropunctate in anterior half; posterior half with weak vertical carinae; metapleuron with
microscopic transverse striae, with irregular macropunctures or reticula in its lower
portion, and with a deep pit just above the base of hind leg. Propodeum with a shelf that
has a large median pit with several carinae on its bottom (Fig. 258). Dorsal and lateral
parts of propodeum coarsely punctate. Propodeal concavity not sharply demarcated by
inferior and superior ridges (the former ridge relatively well developed), shining, with
irregular and indistinct carinae over the surface, and with a complete and high median
carina. Anterior vertical part of gastral tergite 1 with ill-defined macropunctures, without
distinct vertical carina; transverse carina shallowly incised in the middle; lateral part with
irregularly shaped large punctures; posterior horizontal part with much smaller and
dense punctures. Tergite 2 finely punctate; punctures tending to become larger toward
base; tergites 3-5 superficially punctate in their apical portion. Punctation on sternites 2-5
121
~cr;~Q
~ ~~ ~- ~~ ~ ~~~
~
267
"268
n 270
269
o
n 273
~~276
.. ,-
......... . .
n
274
275
Figs. 267-275. Anterior face of gastral segment 1 in Japanese Symmorphus. 267, apidornatus 5f-; 268,
ditto r1'; 269, carinatus 5f- r1'; 270, iwatai r1'; 271, tsushimanus r1'; 272, mutinensis 5f- r1'; 273,
mizuhonis 5f- r1'; 274, cliens 5f- r1'; 275, ishikawai 5f-.
Figs. 276. Apical part of gastral tergite 1 of S. tsushimanus.
much finer than on corresponding tergites.
Body black, with the following parts yellow: a basal marking of variable size on
clypeus, frontal spot (slightly tinged with orange), minute spot on temple, a pair of spots
on pronotum anteriorly, a spot on dorsal mesepisternum (often lost), apical bands on
gastral tergites 1, 2, 4 (band on t2 widest and that on t4 narrowest), a medially narrowed
apical band on sternite 2, anterior faces of fore tibia and tarsus. Mandible ferruginous
apically. Legs tinged with brown, especially on tarsi.
Male. Body length (h+th+t1+2): 8.5-9.5 mm. Fore wing length: 8.0-8.5 mm. The male
differs from the female in the following details (also see Tsuneki, 1986): clypeus only
slightly wider than high (Fig. 231), much more finely punctate, largely yellow; front
corners of pronotum roundly produced (Fig. 248); mid tibia and tarsus also with yellow
markings. Antennal segments 9-13 with distinct narrow tyloids that are yellowish; last
segment slightly longer than wide at its base and nearly as long as segment 12 (Fig. 238).
Material examined. Honshu: Aomori-len - 15f-, Zatoishi, Hirosaki, 20 vii 1973 (5. Takahashi); Yamagata-ken
- 15f-, Ichino-taki (500 m alt.), Yamagata-shi, 23 vii 1973 (5. Takahashi); Tochigi-ken -15f-, Senjugahara, Nikko,
2 viii 1971 (HI); Yamanashi-len - 2r1' r1'15f-, Shioyama, iv-v 1978, bred from a nest by Y. Tanaka.
Shikoku: K6chi-ken - 15f-, Hirooka (H. Okamoto); Kagatm-ken - 15f-, Hirai-ch6, Sanuki, 30 v 1948 (K
Iwata).
Distribution. Honshu; Shikoku. Korea. Smith (1873) mentioned "Hakodadi" as a
locality of this species, but it is not certain whether "Hakodadi" denotes Hakodate
(southern Hokkaid6) or not.
122
Biology. I have received 2 males and 1 female of this species bred from a nest by Mr. Y.
Tanaka, who, however, has not published his nest data. This species may be a tube-renter.
"Odynerus (Symmorphus) captivus Smith" reported for nesting biology by Iwata (1938b,
1971) was most probably S. foveoliltus as suggested by Giordani Soika (1975).
Symmorphus sounkionis Tsuneki
(Fig. 279)
Symmorphus sounkionis Tsuneki, 1986, Spec. Pub!. Jpn. Hym. Assoc. 32: 22, fig. 65
(~)
(type loc.:
SOunkyo, Hokkaido).
Japanese name: Ezo-hamushi-dorobachi.
Diagnosis. Female. I have not examined any specimen of this species. Here, Tsuneki's
(1986) original desciption is reproduced. "Closely related to S. captivus (Smith) and
similarly a large-sized species, about 14 mm in length, but can be separated therefrom by
the following differences: 1. c1ypeus without yellow mark, Gf [gastral tergitel 3 apically
yellow banded, but 4 without the band, mark on pronotum pointed at antero-Iateral
corners and mandible with a yellow mark at base on outer side; further GS 2 broadly,
thoroughly banded at apex (in captivus medianly narrowed and interrupted); 2. antennal
scape and flagellum ferruginous yellow beneath; 3. apical margin of c1ypeus narrower,
with emargination much weaker; 4. longitudinal furrow of mesopleuron narrower, with
lower margin more distinctly outlined and sectioned foveae more distinct; 5. side of
propodeum almost without distinct longitudinal striae, only very sparsely, obscurely
striate, with intervals faintly microstriolate, outward expansion at dorsal and posterior
margins stronger, with punctures much less marked; 6. posterior aspect of propodeum
obliquely, regularly, much more strongly and coarsely striate; 7. punctures on GT 1 much
finer; 8. punctures at base and apex of mesoscutum much less different in size, surface
very closely, sub reticulately covered with large, uniform punctures, everywhere not
bipunctate; 9. clypeus not bipunctate, anteriorly longitudinally rugosely and closely,
posteriorly sparsely punctate; 10. frons more distinctly and closely rugoso-reticu)atepunctate; 11. postscutellum and its posterior excavation somewhat different in structure
from those of captivus".
Male. Unknown.
Distribution. Hokkaid6. This species is known only from the holotype collected at
S6unky6 at the foot of Mts. Taisetsu.
Symmorphus apiciomatus (Cameron)
(Figs. 239, 249, 259, 267, 280)
Ancistrocerus apiciornatus Cameron, 1911, EntomolOgist, 44: 288 (~ J')(type loc.: Japan).
Odynerus apiciornatus: 5chulthess, 1934, Arb. Morph. Taxon. Entomol. 1: 74; Giordani Soika, 1941, Boll.
Soc. Venez. Star. Nat. 2: 232.
Symmorphus apiciornatus: Vecht and Fischer, 1971, Hym. Cat. (n. 00.) 8: 119; Giordani Soika, 1975, Boll.
Mus. Civ. Venez. 27: 150 (in key), 156-158, figs. 4, 9.
Japanese name: Saij6-hamushi-dorobachi.
123
Diagnosis. Female. Body length (h+th+t1+2): 6.5-8.0 mm. Fore wing length: 7.0-8.0
mm. Head subcircular, relatively finely macropunctate; punctation on frons much denser
than in other parts; frontal keel rather sharp; temple as seen from above wider than eye;
cephalic foveae slightly smaller than posterior ocelli, not placed in depressions. Clypeus
wider than high, very finely punctate, shallowly emarginate at apex. Distance between the
posterior ocelli only slightly shorter than that between posterior ocellus and eye (11:12).
Flagellar segments short; segment 1 (antennal seg. 3) as long as wide at its apex;
subsequent segments wider than long. Pronotum moderately punctate, with a deep
furrow in front of pro notal tubercle; front comers of pronotum round. Mesoscutum with
relatively small macropunctures and numerous micropunctures that are not conspicuous;
notaulices complete; parapsidallines weak. Mesopleuron shining, with fine and sparse
S.
277
foveolatus
,
o
~~
S. decens
278
es.
.5.
captivus
sounkionis
'.
279
Figs. 271-279. Distribution of Japanese Symmorphus (I).
124
macropunctation (posterior portion of ventral mesepisternum dull); epicnemial carina
developed in its upper portion alone; scutellum nearly as wide as long, flat over the disc,
with a median longitudinal furrow; macropunctation on scutellum fine and sparse.
Mctanotum more coarsely punctate anteriorly; dull posteriorly. Metapleuron very finely
and transversely striate above, dull below. Propodeal shelf very narrow; median pit small
(Fig. 259); dorsal face irregularly reticulate; lateral face very finely striate, without distinct
punctures; propodeal concavity not sharply demarcated by ridges, with fine striae that
run obliquely; and with a complete median carina. Anterior face of gastral tergite 1 with a
weak vertical median carina; transverse carina well developed, rather deeply incised in
the middle (Fig. 267); posterior horizontal part coarsely punctate, apically slightly
duplicate. Tergite 2 very faintly punctate at base; other tergites virtually without
punctures; disc of sternite 2 very finely punctate at base; sternites 3-6 almost impunctate.
Black, with the following parts yellow: minute spot on temple, apical bands on
tergites 1, 2 and 4 (the last often reduced or lost), and posterolateral spots on sternite 2, an
irregular marking on the anterior face of fore tibia (often lost).
Male. Body length (h+th+tl +2): 5.5-6.0 mm. Fore wing length: 5.0-5.5 mm. Differs
from the female in the following points: c1ypeus as wide as high, more narrowly
emarginate at apex, largely yellow; mesal part of mandible extensively yellow; anterior
face of antennal scape with a large yellow marking; 3 terminal segments of antenna
ferruginous below; distance between posterior ocelli distinctly longer than that between
posterior ocellus and eye; median incision of transverse carina on tergite 1 much
shallower than in the female (Fig. 268); tergite 5 sometimes with a yellow apical band;
tibiae and tarsi of all legs extensively yellow. Antennal segments 8-12 very thick; segments
10-13 each with a trace of tyloid; last segment very small, as wide as or wider than long at
its base (Fig. 239).
Milterial examined. Hokkaido: 16', Tokachi-dake (1000-2000 malt.), 4-6 vii 1938 (T. Sawamoto & H.
Takahasi).
Honshu: Miyagi-ken - IOf, TagajO, 4 vi 1983 (K. Goukon); Fukushima-ken - 56' 6'1 Of , Yunohana, Aizu, 9
vi 1978 (HI); Nagano-ken -1 Of, Niiyama, Ina, 20 v 1962 (YM), 1Of, Takato, Ina, 19 v 1963 (YM); Gifu-ken - 20f
Of, Yoro Park, Yom-gun, 6 v 1979 (y. Takai); Ky6to-fu - IOf, Iwakura, 10 v 1985 (T. Ichino); Ishikawa-ken -1 Of ,
Sodani, Tsurugi, 23 v 1985 G. Toga~hi); Shimane-ken -1 d', Mt. Makuragi, 25 iv 1983 (yM).
Sado-ga-shima (Niigata-ken): 1 d', Ogi, 18 v 1975 (A. Seino), 2 d' d'IOf, bred from a nest by T. Dnuma
(1987-88).
Shikoku: K6chi-ken - IOf, Kodakasa-yama, K6chi-shi, 23 vi 1930 (y. Sugihara), IOf, same loc., 7 v 1931
(y. Sugihara), IOf, Susaki, 21 v 1933 (Y. Sugihara).
Distribution. Hokkaid6; Honshu; Sado-ga-shima; Shikoku; KyUshu.
Biology. This species nests in Miscanthus stems, and hunt for mature larvae or
prepupae of curculionids, Rhyncaenus galloisi and R. takabayasii, for the young (Nakatani,
pers. comm.).
Parasitoid: Melittobia acasta (Hymenoptera, Eulophidae) (Maeta, 1985).
Symmorphus carinatus Sk. Yamane, spec. nov.
(Figs. 232, 233, 240, 250, 269, 281)
Japanese name: Tsuyakeshi-hamushi-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 7.5 mm; fore wing length: 7.0 mm in the
125
holotype. Head wider than high, relatively densely punctate on frons and ocular sinus
(punctures not well defined), much finely and sparsely punctate on gena and vertex and
in ocellar region. Frontal carina sharp. Clypeus distinctly wider than high, very finely
punctate above and more strongly punctate below; apical emargination shallow (Fig. 232).
Cephalic foveae small, bearing brownish hairs. Distance between the posterior ocelli as
long as that between posterior ocellus and eye. Antenna very short and thick; flagellar
segment 1 thick, as long as wide at its apex. Pronotum coarsely macropunctate or
reticulate in the lateral portion, much finely punctate along anterior margin of
mesoscutum. Mesoscutum moderately macropunctate (punctures not clearly defined),
with many micropunctures; notaulices complete; parapsidal lines obscure. Mesopleuron
with sparse macropunctures; epicnemial carina complete; posterior portion of ventral
mesepisternum shagreened; scutellum slightly wider than long, with a wide, ill-defined
median furrow, moderately punctate. Metanotum coarsely macropunctate in anterior half
and dull posteriorly; metapleuron wholly microscopically striate or shagreened.
Propodeum with a distinct shelf which has a large median pit (Fig. 260); dorsal part of
propodeum irregularly reticulate; lateral face much as metapleuron, with a few punctures
above; propodeal concavity obliquely rugulose, somewhat shining, with a complete
median carina. Anterior vertical face of gastral tergite 1 with a strong median vertical
carina; transverse carina not incised in the middle (Fig. 269); posterior horizontal part of
the tergite with large punctures which are ill defined, with a median furrow which is
shallow and indistinct. Tergite 2 finely punctate at base; other tergites almost impunctate.
Disc of sternite 2 very finely punctate at base.
Black; the following parts yellow: a pair of spots on frons, spot on temple, a pair of
spots on pronotum anteriorly, a relatively large spot on dorsal mesepisternum, apical
bands on tergites 1, 2, 4 (the last interrupted medially), an incomplete apical band on
sternite 2, fore and mid tibiae on anterior face extensively. Legs somewhat brownish.
Male. Body length (h+th+t1+2) 6.5 mm, fore wing length 6.5 mm in the paratype.
Similar to the female but different therefrom in the following points: clypeus largely
yellow, antennal scape with a yellow stripe below, 3 terminal sgments of antenna with
ferruginous stripes below, mandible with a yellow mark in mesal portion, distance
between the posterior ocelli longer than that between posterior ocellus and eye, front
corners of pronotum acute to some extent (Fig. 250), median pit of propodeal shelf with a
distinct longitudinal carina on the bottom, fore and mid tibiae and tarsi of all legs more
extensively yellow. Flagellar segments of antenna rather thick; antennal segments 11-13
each with a trace of tyloid; last segment small, wider than long at its base (Fig. 240).
Holotype. 5?-, Kodakasa-yama, K6chi-shi, K6chi-ken, Shikoku, 7 v 1931 (Y. Sugihara).
Para type. 6', Engyoji, K6chi-shi, 19 iv 1931 (Y. Sugihara).
Distribution. Shikoku.
Taxonomic notes. This species resembles S. apiciornatus in the sculpture of metapleuron
and of the lateral face of propodeum, but is distinguished from the latter by the following
aspects: propodeum with a wider shelf which has a large median pit, transverse carina on
tergite 1 not interrupted medially, and alitrunk with yellow markings.
I have examined one male specimen from Honshu (Mt. Hayachine, Iwate-ken, 1 viii
1965, A. Nakanishi leg.) which largely agrees with the paratype male but differs in the
following details: striae on metapleuron and the lateral face of propodeum coarser,
posterior horizontal part of tergite 1 reticulate rather than punctate, and yellow markings
on mesosoma much reduced.
126
Symmorphus iwatai Sk. Yamane, spec. nov.
(Figs. 234, 241, 251, 261, 270, 281)
Japanese name: Iwata-hamushi-dorobachi.
Diagnosis. Male. Body length (h+th+t1+2): 7.5 mm; fore wing length: 8.0 mm in the
holotype. Head distinctly wider than high; width: height (from top to lower margin of
clypeus) = 73 : 67. Punctures on frons and ocular sinus dense and not uniform in shape
and size; region between the antennae raised to form "tubercle"; frontal keel rather sharp;
anterior tentorial pit deep. Punctation on gena and vertex and in the area surrounded by
the ocelli much sparser. Clypeus slightly wider than high, very finely punctate, apically
282
Figs. 280-282. Distribution of Japanese Symmorphus
127
aI).
rather deeply emarginate, with acute lateral teeth (Fig. 234). Distance between the
posterior ocelli longer than ocello-ocular distance (12:10). Temple as seen from above
slightly narrower than eye. Flagellar segments of antenna moderately thick; antennal
segments 9-13 with rather distinct tyloids; last segment slightly longer than wide at its
base, nearly as long as segment 12 (Fig. 241). Pronotum macropunctate in dorsal part
(punctures not uniform), strongly and transversely striate on lateral face; pronotal carina
developed but much reduced medially; front corners of pronotum slightly angulate (Fig.
251). Mesoscutum with coarse punctation mainly in anterior half; micropunctures more
numerous in posterior portion; notaulices well defined and relatively deep; parapsidal
lines carinate. Epicnemial carina well developed, extending from the lower margin of
dorsal mesepisternum to the base of mid leg; epicnemium with a few macro punctures
below; dorsal mesepisternum finely and very sparsely macropunctate; ventral
mesepisternum and mesepimeron rather strongly punctate; the area surrounded by
mesepisternum and mesepimeron broadly concave; posterior portion of ventral
mesepisternum shagreened. Scutellum with large, irregular punctures; median furrow
indistinct. Metanotum anteriorly coarsely sculptured, posteriorly dull; metapleuron striate
above, shagreened below. Propodeal shelf present, with a distinct median pit; the bottom
of the pit sloping and with a vertical carina (Fig. 261). Dorsal face of propodeum coarsely
punctate or reticulate; lateral face finely striate. Propodeal concavity not sharply
demarcated by ridges, superficially striate, with a complete median carina. Anterior face
of gastral tergite 1 with ill-defined, large punctures, without median vertical carina.
Transverse carina of tergite 1 interrupted in the middle (Fig. 270), laterally not connected
with the carinae that extend from the preapical swelling with a yellow apical band.
Posterior horizontal part of tergite 1 with coarse punctures which are variable in shape
and size, slightly duplicate at apex. Tergite 2 finely punctate at base; tergites 3-5 slightly
more strongly punctate near apex. Disc of sternite 2 punctate at base; punctation on
sternites 3-5 much finer than on corresponding tergites.
Black, with the following parts yellow: c1ypeus almost wholly, spot on temple, a pair
of triangle markings on propodeum anteriorly, apical band on tergite I, wider band on
tergite 2, apical band on sternite 2 which is dilated laterally and narrowly interrupted at
middle, apex of fore femur, tibiae and tarsi of all legs extensively. Tyloids of apical 4
segments of antenna ferruginous.
Female unknown.
Holotype. 6', Johoku, Sasayama, Hyogo-ken, Honshu, 8 v 1957 (K. Iwata).
Distribution. Honshu.
Biology. On the data label, Iwata wrote "Yanagihamushi" [Chrysomela vigintipunctata,
Chrysomelidael, though it is not certain whether this specimen was bred from a nest by
Iwata.
Symmorphus tsushimanus Sk. Yamane, spec. nov.
(Figs. 235, 242, 252, 262,271,276,281)
Japanese name: Tsushima-hamushi-dorobachi.
Diagnosis. Male. Body length (h+th+t1+2): 6.5 mm, fore wing length: 7.0 mm in the
holotype. Head wider than high, with medium-sized punctures that are smaller between
128
the posterior ocelli and sparse on the vertex and gena, and with erect hairs which are
longer than hairs on thorax and gaster. Clypeus wider than high, finely punctate above
and more strongly but sparsely punctate below, anteriorly very shallowly emarginate (Fig.
235). Temple seen from above as wide as eye. Antenna club-shaped, with thickened
flagellum; scape as long as pedicel + flageller segment 1; antennal segments 10-13 with illdefined tyloids; tyloids on the last two segments wide; last segment as long as wide at its
base (Fig. 242). Pronotum with medium-sized macropunctures on dorsal part, macropunctate and striate on lateral face; pro notal carina sharp; front corners only slightly
produced (Fig. 252). Mesoscutum rather strongly macro punctate; spaces between
punctures generally larger than punctures and with numerous micropunctures;
macropunctation much denser near the base of mesoscutum; notaulices complete;
parapsidallines slightly raised; median scutalline visible only in anterior third. Scutellum
sparsely macropunctate except in the posterior region, with a median furrow.
Epicnemium very weakly macropunctate, almost impunctate above; dorsal
mesepisternum macropunctate only in posterior part; ventral mesepisternum sparsely
macropunctate; macropunctation on mesepimeron denser than on mesepisternum;
epipleural suture wide, with several transverse carinae; posterior part of ventral
mescpisternum shagreened. Metanotum coarsely punctate in anterior half and shagreened
in posterior part; metapleuron finely striate. Propodeum with a narrow but distinct shelf
which has a deep median pit (Fig. 262); the shelf and dorsal face of propodeum coarsely
macro punctate or reticulate; propodeal concavity microstriate over the surface and its
median carina distinct. Lateral face of propodeum striate, with large punctures posteriorly.
Gastral segment I, seen from above, as long as wide. Anterior face of gastral tergite 1 with
ill-defined macro punctures laterally, without vertical carina; transverse carina narrowly
and shallowly incised in the middle (Fig. 271); dorsal horizontal part strongly punctate;
posterior lamella medially produced backward (Fig. 276; this condition is peculiar to this
species among the Japanese species). Sternite 1 also strongly punctate. Other tergites and
sternites much more finely punctate. Anterior faces of coxae and femora below with silver
hairs.
Black, with yellow and orange yellow markings as follows: a transverse spot between
antennae, upper half of clypeus, a large marking on antennal scape below, apical 3 or 4
antennal segments below, small spot on temple, a pair of triangular marking on pronotum
anteriorly, spot on dorsal mesepisternum, a pair of spots on scutellum, apical band on
'.'-"
'-
....
283
Fig. 283. Distribution of Japanese Symmorphus (III).
129
tergite 1 (medially widened with anterior incision), wide apical band on tergite 2
somewhat sinuate, a medially interrupted narrow apical band on tergite 3, narrow but
complete apical bands on tergites 4 and 5, medially interrupted apical band on sternite 2,
apices of fore and mid femur, anterior face of fore tibia, segment 1 of fore tarsus, irregular
markings on the outer face of mid tibia. Legs blackish brown, marked with yellow as
mentioned above.
Holotype. (/" top of Ariake-san (558 malt.), Tsushima Is., 4 vii 1975 (Y. Miyatake).
Deposited in the Collection of Osaka City Museum of Natural History.
Distribution. Tsushima Is. (Shimo-agata).
Symmorphus mutinensis (Baldini)
(Figs. 243, 253, 263, 272, 281)
Odynerus (Protodynerus) sinuaius var. mutinensis Baldini, 1894, Atti Soc. Nat. Modena, (3)13: 78, pI. 3,
fig. 6 (type loc.: environs of Modena, Italy?).
Odynerus (Symmorphus) mutinensis: B1iithgen, 1961, Faltenwespen Mitteleuropas, p. 188 (in key), 197,
198.
Symmorphus mutinensis: Vecht and Fischer, 1972, Hym. Cat. (n. ed.) 8: 122.
Japanese name: Mitsuten-hamushi-dorobachi.
Diagnosis (based upon subsp. auster G. S.). Female. Body length (h+th+t1+2): 6.5-7.5
mm. Fore wing length: 6.5-7.0 mm. Head subcircular, strongly and irregularly punctate on
frons and ocular sinus, finely and sparsely punctate on vertex and gena. Frontal carina
sharp. Clypeus much wider than high, strongly convex, finely punctate above, strongly
and sparsely punctate below, relatively shallowly emarginate at apex; lateral teeth of the
emargination blunt. Cephalic foveae half as large as posterior ocelli in diameter. Distance
between the posterior ocelli as long as or slightly longer than oceno-ocular distance.
Flagellar segment 1 of antenna 1.2 times as long as wide at its apex. Pronotum irregularly
macro punctate; spaces between punctures with numerous micropunctures along posterior
margin of pronotum; depression just in front of pro notal tubercle very large, with sharp
carinae on its bottom. Mesoscutum distinctly duplipunctate; macropunctation denser in
anterior portion; micropunctation dense and very conspicuous over the disc; notaulices
complete, deep near the base of disc; parapsidallines carinate; median scutalline running
from anterior margin to basal margin of the disc, forming a narrow groove in basal half.
Epicnemial carina well developed, sometimes extending onto dorsal mesepisternum;
epicnemium above almost impunctate; dorsal and ventral mesepisternum with large and
sparse punctures; mesepimeron densely macro punctate or even reticulate; posterior part
of ventral mesepisternum shagreened, slightly shining. Scutellum with a few large
punctures and numerous micropunctures, with wide median furrow; narrow posterior
zone with many short carinae. Metanotum coarsely and irregularly macropunctate; its
posterior zone smooth, somewhat shining; metapleuron finely striate above and coarsely
striate in anterior portion of lower half. Propodeal shelf distinct, with a large median pit
(Fig. 263). Dorsal face of propodeum including the shelf coarsely punctate or reticulate;
lateral face strongly punctate in upper and posterior portion, finely striate toward the
border with metanotum. Propodeal concavity defined only below, with superficial
micro striation over its surface, with a complete median carina. Anterior face of gastral
tergite 1 without distinct punctures and median carina; transverse carina not interrupted
130
or only shallowly incised in the middle (Fig. 272), extending backward from both lateral
ends to be connected with preapical swelling; posterior horizontal part coarsely punctate,
with inconspicuous micropunctures. Tergite 2 with superficial punctures at base; other
tergites almost impunctate. Basal half of the disc of sternite 2 distinctly punctate; stemites
3-6 almost impunctate.
Male. Body length (h+th+t1 +2): 5.5-6.5 mm. Fore wing length: 6.0-6.5 mm. Sculpture
as in the female. Front corners of pronotum more produced (Fig. 253). Median scutal line
obscure. Macropunctation on meso scutum much stonger and denser. Antennal segments
10-13 with ill-defined tyloids; last segment slightly longer than wide at its base and nearly
as long as segment 12 (Fig. 243).
This species is widely distributed in the Palearctic region. The nesting biology of this
species was studied by Enslin (1921) and J0rgensen (1942) in Europe (referred to as S.
sinuatus Fabricius). Female wasps construct their nests in bramble straws and thatch. The
cell partitions are made of clay. The number of cells constructed per nest ranges between 4
and 8 in the nests made in bramble straws, and between 1 and 6 (mostly 1 and 2) in those
made in thatch. Larval cells are provisioned with chrysomelid larvae. Enslin (1921)
reported that larvae of Microlepidoptera were found in nests of this species, though this
has not been confirmed by other authors.
In Japan two subspecies, auster Giordani Soika and yezoanus Tsuneki, are known to
occur; both are not common in any locality.
Symmorphus mutinensis auster Giordani Soika
(Fig. 281)
Symmorphus mutinensis auster Giordani Soika, 1975, Boll. Mus. Civ. Stor. Nat. Venez. 27: 1.50 (in key),
(~) (type loc.: Mt. Hayachine, Iwate-ken); Tsuneki, 1977, Spec. Publ. Jpn. Hym. Assoc. 5: 16 (d').
160,161
Japanese name: Mitsuten-hamushi-dorobachi.
Diagnosis. Female. Black, with the following parts yellow or orange yellow: transverse
frontal marking, minute spot on temple, a pair of spots on scutellum (sometimes lost),
apical band on gastral tergite 1 (medially widened and anteriorly incised at middle), a
sinuated apical band on tergite 2, much narrower bands on tergite 4 and sternite 2,
anterior face of fore tibia, mid and sometimes hind tibiae partIy. Legs black, but tinged
with brown especially on tibiae and tarsi.
Male. Body markings paler than in the female. Mandible and/or clypeus often with
irregular yellow markings. Tyloids on 4 terminal segments of antenna ferruginous.
Scutellum always wholly black. Tarsi of all legs at least partly yellowish.
Material examined. Honshu: Aomori-ken - 1 d', Yamagata, 16 vi 1936 (I. Tateyama), 3 d' d', Zatoishi,
Hirosaki,29 vi 1983 (M. Yamada); [wate-ken - 2d' d', Kadoma-guchi, Mt. Hayachine,28 vii - 1 viii 1965 (A.
Nakanishi); Miyagi-ken - 1 d', Mt. Zao, 19 vi 1983 (K Goukon); Nagano-ken - 1 ~, "Shinshu", 4 viii 1929 (Y.
Ota); Fukui-ken -1 (/" Nakamura, Ono, 12 vi 1971 (TT), 1~, Itoshiro Riv. 25 vii 1973 (TT), 1 d', Arashiguchi Ono, 24 vi 1980 (H. Kurokawa), 1 ~, Nagatani - Obama, 25 vii 1980 (H. Kurokawa); Gifu-ken - 1 ~, Itoshiro,
Shiratori,21 ix 1986 (Y. Takail.
Distribution. HonshiL
Biology. In Niigata-ken, this form nests in bamboo tubes. One to five brood cells are
constructed in a tube nest. The prey is not identified (T. Onuma, pers. comm.).
131
Symmorphus mutinensis yezoanus Tsuneki
(Fig. 281)
Symmorphus mutinensis yezoanus Tsuneki, 1977, Spec. Pub\. Jpn. Hym. Assoc. 5: 16, 17 (J' -'f )(type loc.:
Sapporo, Hokkaido).
This subspecies is said to be distinguished from ssp. auster by several aspects of
punctation (Tsuneki, 1977). It has been known from Hokkaid6 and the northernmost part
of Honshu where it may occur together with ssp. auster. As Tsuneki mentioned, further
study with ample material is necessary to solve the subspecies problem in this species in
Japan.
Symmorphus mizuhonis Tsuneki
(Figs. 244, 254, 264, 273, 282)
Symmorphus mizuhonis Tsuneki, 1977, Spec. Pub\. Jpn. Hym. Assoc. 5: 17-20 (J' -'f )(type loc.: Okunasu,
Tochigi-ken, Honshu).
Japanese name: Mizuho-hamushi-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 7.0-8.5 mm. Fore wing length: 6.5-8.0
mm. Head strongly punctate on frons and ocular sinus, more finely and sparsely on gena
and vertex. Clypeus finely and sparsely macropunctate, much less swollen than in S.
mutinensis, relatively widely emarginate at apex. Cephalic foveae very small, usually less
than half as large as posterior ocelli in diameter, situated more distantly from posterior
ocelli than in S. mutinensis. Distance between the posterior ocelli distinctly shorter than
ocello-ocular distance (8:10). Flagellar segment 1 of antenna slightly longer than wide at
its apex. Pronotum moderately macropunctate in dorsal part, much more densely on
lateral face (spaces between punctures tend to form carinae); pro notal carina well
developed to form lamella in the lateral face of pronotum, but almost absent dorsally;
front corners of pronotum, seen from above, rather strongly produced as in S. mutinensis.
Mesoscutum densely micro punctate over the disc, sparsely with small macropunctures;
notaulices evanescent in anterior half except for anterior excavation; parapsidal lines
inconspicuous; median scutal line undeveloped, not forming a furrow near the base of
disc. Mesopleuron with a few macropunctures; epicnemial carina complete but not
extending onto dorsal mesepisternum; posterior portion of ventral mesepisternum
shagreened. Scutellum smooth, with a few small macropunctures, with a wide median
furrow, and posteriorly shagreened. Metanotum irregularly sculptured anteriorly,
somewhat shining; posterior half shagreened; metapleuron finely striate and partly
shagreened. Propodeum with relatively wide shelf which has a very large median pit (Fig.
264); dorsal face of propodeum including the shelf coarsely punctate or reticulate; lateral
face striate below, and with large punctures above. Propodeal concavity well demarcated
in lower half, with a complete median vertical carina which is connected with the ridge
enCircling the median pit; surface of concavity irregularly striate. Anterior vertical face of
gastral tergite 1 with a few ill-defined punctures, with a trace of vertical carina; transverse
carina not incised medially (Fig. 273); posterior horizontal part of the tergite weakly
punctate, with a short median carina at base. Tergite 2 finely punctate (punctures become
132
stronger and denser toward the base); punctation on other tergites and sternites 2-6 much
finer.
Black, with the following parts yellow: two contiguous markings on frons, minute
spot on temple, medially widened apical band on tergite 1 (anteriorly incised in the shape
of a triangle), relatively regular band on tergite 2, narrower and sinuated band on sternite
2. All the legs wholly blackish. Wings rather strongly clouded.
Male. Body length (h+th+t1 +2): 6.0-7.5 mm. Fore wing length: 6.0-7.5 mm. Similar to
the female in structure and coloration. Clypeus and/or mandible often marked with
yellow. Distance between the posterior ocelli slightly longer than ocello-ocular distance.
Front corners of pronotum more pronounced than in the female (Fig. 254). Anterior face of
fore tibia marked with yellow. Antennal segments 11-13 with tyloids; tyloid on segment 11
evanescent; last segment small, shorter than segment 12 (Fig. 244); two or three apical
segments sometimes extensively ferruginous.
Material examined. Honshu: Akita-ken - 2 d' d'J!?-, Mt. Nyuto, 6 viii 1983 (M. Yamada); lwate-ken - 1-'?,
Ashiro, 19 ix 1976 (YM); Miyagi-ken - 4 d' d'1-'?, Kamoshika Spa, Mt. Zao, 25 vi 1979 (TN), 1-'?, Mt. Za6, 22
viii 1979 (K. Kojima & T. Nishida), 1-'?, same loc., 31 viii 1979 (K. Goukon); Niigata-ken - 1 d', Yuzawa, 8 viii
1966 (HI); Tochigi-hn - 1;1, Okunasu, 26 vii 1970 (TN); Gumma-ken - 1;1, Buson-5anroku, 21 vii 1974 (HI);
Saitama-ken -1 d'1-'?, Karisaka-toge, Otaki, 29 vii 1983 (TN); Nagano-ken - 4d' d', Mt. Komaga-take, Ina, 19 vii
1962 (K Oshima); lshikawa-ken - 1-'?, Haku-5an, 2 viii 1962 (T. Naito); Gifu-ken - 1-'?, Migimata-dani (15002000 m alt.), Mts. Hida, 7-8 viii 1987 (Y. Takai).
Distribution. Honshu (mountainous regions).
Symmorphus cliens Giordani Soika
(Fig. 245,255,265,274,283)
Symmorphus diens Giordani Soika, 1975, Boll. Mus. Civ. Stor. Nat. Venez. 27: 150 (in key), 158, 159, figs.
5,10 (-'? d') (type loc.: Mt. Hayachine, Iwate-ken, Honshu).
Japanese name: Katatoge-hamushi-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 5.5-7.0 mm. Fore wing length: 5.5-6.5
mm. Head subcircular; punctation on head similar to that in mizuhonis, but slightly
stronger on gena. Frontal keel sharp. Clypeus distinctly wider than high, more swollen
than in mizuhonis, sparsely macropunctate and micro punctate over the disc; apical margin
shallowly emarginate. Cephalic foveae relatively large, more than half the diameter of
posterior ocelli. Distance between the posterior ocelli slightly longer than ocello-ocular
distance. Flagellar segment 1 of antenna small, only slightly longer than pedicel, nearly as
long as wide at its apex. Pronotum densely macropunctate; spaces between punctures
tend to run into carinae in lower portion of lateral face; pro notal carina not distinctly
lamellate on the lateral face of pronotum, weakened dorsally; front corners of pronotum
somewhat angulate but less pronounced than in mizuhonis. Mesoscutum much more
strongly macropunctate over the disc; micropunctures less conspicuous than in mutinensis;
notaulices complete but narrow and shallow in its anterior half; parapsidal lines weak;
median scutal line absent in the basal half of the disc. Epicnemial carina complete, not
extending onto dorsal mesepisternum; epicnemium and dorsal mesepisternum almost
impunctate; ventral mesepisternum with a few macropunctures, posteriorly shagreened;
mesepimeron sparsely macropunctate. Scutellum sparsely macropunctate, posteriorly
with short carinae; micropunctation inconspicuous. Metanotum irregularly sculptured
133
anteriorly, smooth and shining posteriorly; metapleuron striate above, smooth below.
Propodeal shelf relatively narrow; median pit distinctly smaller than in mizuhonis, often a
narrow slit (Fig. 265). Dorsal face of propodeum including the shelf coarsely punctate or
reticulate; lateral face striate, with some large punctures above. Propodeal concavity well
defined below, lustrous, with superficial striae over the surface and with a complete
median carina. Anterior face of gastral tergite 1 with ill-defined punctures; vertical carina
almost invisible; transverse carina roundly incised in the middle (Fig. 274); posterior
horizontal part more weakly and sparsely punctate than in mizuhonis, without median
carina at base. Tergite 2 basally sparsely punctate; punctation very weak in apical 1/2.
Punctation on other tergites finer. Sternite 2 rather strongly punctate in the basal half of
the disc; sternite 2 in apical part and other sternites much more finely punctate.
Black; the following parts yellow: two contiguous markings on frons (rarely lost),
medially widened apical band on tergite 1 (anteriorly incised at the middle), slightly
sinuated band on tergite 2, an incomplete narrow band on tergite 4 (sometimes lost),
sinuated band on sternite 2, anterior face of fore tibia. Terminal segment of antenna
ferruginous below. Legs blackish brown.
Male. Body length (h+th+t1+2): 5.5-6.5 mm. Fore wing length: 5.5-6.0 mm. Clypeus
more finely punctate than in the female, with a large irregular yellow marking. Mandible
yellow-maculated. Frontal mark always absent. Apical band on tergite 4 often much
reduced or lost. Fore and mid tibiae extensively yellow. Antennal segments 11-13 with
rather distinct tyloids; last segment very small (Fig. 245) and wholly orange in color;
segments 11 and 12 often partly orange.
Material examined. Hokkaido: 2~ ~,Shikotsu-ko, 17 vii 1977 (SKY).
Honshu: Aomori-ken - 4~ ~, Hirosaki, 5 vii 1976 (SKY), 1 ~, SOma, 31 vii 1982 (M. Yamada), 1 ~,
Ikarigaseki, 14 viii 1982 (M. Yamada), 2~ ~,Mt. Bonju, Namioka, 2 vii 1983 (M. Yamada), 2~ ~,Kuroishi, 9
vii 1983 (M. Yamada), 1 ~, same loc., 27 viii 1983 (M. Yamada); Akita..Jcen - 1 ~, Mt. Nyuto, 5 viii 1983 (M.
Yamada); Fukushima..Jcen - 1 t, Yunohana, 24 vi 1977 (H. Koike); Niigata..Jcen - 1 t, Gimpei-zan, bred from a
nest by T. Onuma (1987-88); Gumma-ken - 7 t t, Hoshi Spa, 24 vi 1984 (A. Seino); Saitama-ken - 1 ~, Izugadake,3 vii 1984 (S. Aoki); Nagano-ken -1 ~,Mt. Nyugasa, Ina, 17 vii 1962 (YM).
Distribution. Hokkaid6; Honshu.
Biology. This species constructs its nests in bamboo tubes (3-4 mm in diam.). The
nesting was observed between June and October in a mountainous region of Niigata-ken
(Onuma, 1989b).
Symmorphus ishikawai Giordani Soika
(Figs. 266,275,283)
Symmorphus ishikawai Giordani Soika, 1975, Boll. Mus. Civ. Stor. Nat. Venez. 27: 151 (in key), 159, 160
(t ~ )(type loc.: Yokoo (1600 m alt.), Nagano-ken).
Japanese name: Soten-hamushi-dorobachi.
Diagnosis. Female. Body length (h+th+t1+2): 7.0-8.0 mm. Fore wing length: 7.0-7.5
mm. Head higher than wide, densely macro punctate on frons and ocular sinus;
punctation on these parts slightly sparser and much finer than in mizuhonis and cliens;
gena finely and vertex very weakly macropunctate. Clypeus wider than high, shallowly
emarginate apically, with relatively large micropunctures densely over the disc; apical part
with a few ill-defined macropunctures. Cephalic foveae very large, as large as posterior
134
ocelli. Distance between the posterior ocelli as long as ocello-ocular distance. Temple as
seen from above slightly wider than eye. Occipital carina relatively gradually curved, not
forming a distinct angle at some distance from mandibular base. Flagellar segment 1 of
antenna only slightly longer than wide at its apex. Pro no tum dorsally weakly
macropunctate (punctures ill defined), in lateral face more strongly punctate (spaces
between punctures running into carinae); front corners of pronotum less produced than in
mizuhonis and cliens. Mesoscutum finely micropunctate, with sparse, small
macropunctures that are ill defined; notaulices complete, rather deep over its length;
parapsidallines weak; median scutalline weak, but extending toward the base of disc as a
shallow, weak furrow. Epicnemial carina developed but not extending onto dorsal
mesepisternum; epicnemium above and dorsal mesepisternum impunctate; ventral
mesepisternum and mesepimeron sparsely and finely macropunctate; posterior portion of
dorsal mesepisternum shagreened but somewhat lustrous. Scutellum finely
micropunctate; posterior zone with irregular carinulae. Metanotum almost smooth
posteriorly; metapleuron strongly striate above, rather smooth below. The condition of
propodeal shelf and dorsolateral portion of propodeum as in diens, but the median pit
slightly larger (Fig. 266). Lateral face of propodeum striate below, reticulate above.
Propodeal concavity relatively well defined by ridges, with almost smooth surface;
median carina complete. Anterior vertical face of gastral tergite 1 without vertical carina;
transverse carina very shallowly incised in the middle (Fig. 275); posterior horizontal part
without basal carina, superficially and sparsely macropunctate. Tergite 2 and sternite 2
finely and sparsely macropunctate at base. Other tergites and sternites almost impunctate.
Black; the following parts yellow or orange yellow: two contiguous frontal spots,
minute spot on temple, apical bands on tergites 1 and 2, apical band on sternite 2 medially
widely interrupted. Legs wholly blackish brown.
Male. According to Giordani Soika (1975), clypeus apically with a large yellow
marking, frons without marking, antennal segments 10-13 below with ferruginous
markings, anterior faces of fore and mid tibiae yellow, and fore femur apically yellow.
Material examined. Honshu: Nagano-ken - 2 ~ ~, Yokoo (1600 malt.) nr Kamik&hi (type loc.), 30 vii 1957
(R. Ishikawa).
Distribution. Honshu (Nagano-ken)
Biology. No information is available.
Genus "Pachymenes" Saussure
Pachymenes Saussure, 1852, Et. Fam. Vesp. 1: 73-77 (type species: Pachymenes sericea Saussure, 1852,
designated by Ashmead, 1902); 1855, Et. Fam. Vesp. 3: 152, 153; Bequaert, 1918, Bull. Amer. Mus. Nat. Hist.
39:87.
Japanese name: Tokkuribachi-modoki Zoku.
This genus is a mixture of species with the following character conditions, and is
distributed in the warmer regions of the world (Bequaert, 1918): "clypeus pyriform,
truncate or bidentate at its apex; mandibles lengthened, crossing each other in an X,
dentate or lobate along their inner margin; mouth parts as in Eumenes, Odynerus, and
Nortonia; shape of the abdomen intermediate between that of Eumenes and that of
Odynerus subgenus Rhygchium; its first segment funnel-shaped, subcampanulate; the first
tergite without any trace of transverse carina; the abdomen as a rule depressed".
135
According to Vecht (pers. comm., 1981) the Oriental "Pachymenes" species differ from
the true Pachymenes of South America, and he is going to describe a new genus for the
Oriental forms (see also Giordani Soika, 1986).
"Pachymenes" yayeyamensis (Matsumura)
(Figs. 284-287, 35SC, 361)
Odynerus yayeyamensis Matsumura, 1926, Ins. Matsum. 1: 36, pI. 3, fig. 11 (nee 12) (J')(type Ioc.:
Okinawa-honto ?).
"Pachymenes" yayeyamensis (Mats.): Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 67-71, figs.
5-9.
Odynerus hOkolt5ensis Sonan, 1929, Trans. Nat. Hist. Soc. Formosa, 19: 534, 535 (Sf J')(type loc.: "Hokoto", Taiwan).
Pachymenes fragilis (Sm.): Yasumatsu, 1938, Trans. Nat. Hist. Soc. Formosa, 28: 446-447; Azuma and
Kinjo, 1987, Cheek-list Ins. Okinawa, p. 315.
Odynerus (Lionolus) fragi/is Sm.: Schulthess, 1934, Arb. Morph. Taxon. Entomol. 1: 94 (in key; from
Taiwan).
Japanese name: Hime-tokkuribachi-modoki.
For diagnosis see Giordani Soika (1986).
Material examined. C. Ryukyus: Yoron-tO - ISf, 4 vi 1985 (SKY), 1 J', 24-8 v 1986 (SKY); "Okinawa" - 1 J'
(S. Sakaguchi).
S. Ryukyus: "Yaeyama" - 1 J', "viii?) 0'7 (SK)" (type); Miyako-jima -lSf , Bora, 27 iv 1981 (T. Fujisawa), 2 J'
J', Bora, 17 vii 1987 (SKY); Tarama-jima - 9J' J'8Sf Sf, 18-19 vii 1987 (SKY); Minna-jima - 3Sf Sf, 29 vi 1988
(SKY); Takelomi-jima - 6 J' J'2 Sf Sf, 24-25 vii 1987 (SKY); Kohama-jima -1 J', 25 vii 1987 (SKY); Kuro-shima - 6 J'
J'ISf, 23-24 vii 1987 (SKY); Iriomole-jima -1 J', 17 iv 1962 (G. Kuno), ISf, 16 iv 1978 (K. Ohara), 2 J' J', Ohara,
23-24 v 1981 (AN), ISf, same Ioc., 1 v 1982 (AN), 1 J', Toyohara, 29 iv 1982 (AN), 2J' J', Ohara, 25 vii 1982
(A. Matsumoto), 1 J'ISf, Funaura, 25 vii 1982 (A. Matsumoto), 2J' J'lSf, Ohara, 29-30 vii 1983 (AN), 1 J',
Amitori, 5 viii 1983 (AN), 2J' J', Otomi, 25 vii 1985 (AN), 1 J'lSf, 30 vii - 1 viii 1985 (AN), ISf, Otomi, 12 x
1987 (AN), 2J' J', 13 x 1987 (AN); Haleruma-jima - 4J' J'4Sf Sf, 30 vi - 1 vii 1988 (SKY); Yonaguni-jima - 1 J',
22-24 vii 1983 (H. Kodama), 7 J' J'ISf, Sonai, 5 vii 1988 (SKY).
Distribution. Amami Is. (Yoron-to); Okinawa Is. (Okinawa-jima ?); Miyako Is.
(Miyako-jima); Tarama Is. (Tarama-jima; Minna-jima); Yaeyama Is. (lshigaki-jima after
Azuma & Kinjo, 1987; Taketomi-jima; Kohama-jima; Kuro-shima; Iriomote-jima;
Hateruma-jima; Yonaguni-jima). Taiwan.
Taxonomic notes. A species of "Pachymenes" common and widely distributed in
Oriental region has been treated as Odynerus fragilis Smith (1857) or "Pachymenes" fragilis
(Smith). Vecht (pers. comm., 1981) has examined the type of Odynerus troglodytes Saussure
(1855) and confirmed the identity of fragilis with troglodytes. The former, therefore, must be
a junior synonym of the latter. The species troglodytes was originally described from
Senegal (Saussure, 1855, p. 249), but Vecht doubts Saussure's statement about the type
locality. Meade-Waldo (1914) in his revision of the Ethiopian Odynerus listed O. troglodytes
under "species unknown to the author".
Recent studies by Giordani Soika (1986) on the "P". troglodytes group revealed that at
least four species are involved in it. According to him "P." yayeyamensis belongs to this
species group and is distinguished from troglodytes by the following male characters:
clypeus more narrowly emarginate apically, with punctures much larger and denser;
tergite 1 wider, with sparser punctures; tergite 2 and sternite 2 with larger and denser
punctures. My study based upon more extensive material from various parts of the
136
Figs. 284-287. "Pachymenes" yayeyamensis. 284, facial color pattern
apical segments of male antenna; 287, body color pattern (~).
(~);
285, male c\ypeus; 286,
Ryukyu. Islands supports his conclusion, and has revealed that the Taiwanese form which
Seman (1929) named Odynerus hOkotOensis belongs to this species.
According to Matsumura and Uchida (1926), the type locality of this species is
Okinawa-honto (=Okinawa-jima), which does not belong to the Yaeyama group. The type
specimen (J', in ColI. Entomol. Inst. Hokkaido Univ.) bears four labels: (1) on upper side,
"Japan Matsumura" in print, and "SK" [So Kiyamu?] in handwriting; on underside,
"Yaeyama" in Chinese characters, and "vii[?] 0'7", both in handwriting, (2) "Odynerus n. sp.
yayeyamensis in handwriting, and "det. Matsumura" in print, (3) red label, "Type
Matsumura" in print, and (4) "Pachymenes fragilis (Smith) det. K. Yasumatsu, 1938" in
handwriting. Another male specimen, believed to be the one in the type-series, bears two
labels: (1) "56" in handwriting, and (2) "Okinawa S. Sakaguchi" in print. In the early 20th,
to many people on the Japanese mainlands, "Okinawa" meant a district covering the
Okinawa Is., Miyako Is., and Yaeyama Is., today called together "Okinawa-ken". I have not
yet examined any specimen of this species from Okinawa-jima, though the species
actually extends its range further north onto Yoron-to. Thus, at present it is reasonable to
regard one of the islands of the Yaeyama group as the type locality of this species.
Biology. This species nests in bamboo tubes (Y. Nakatani, pers. comm.).
Genus Oreumenes Bequaert
Oreumenes Bequaert, 1926, Ann. S. Afr. Mus. 23: 488 (subgenus of Eumenes Latreille)(type species:
(Eumenes harmandi Perez.) =E. decaratus Smith, original designation); Vecht and Fischer, 1972, Hym. Cat. (n.
ed.) 8: 134 (as subgenus).
Japanese name: Suzubachi Zoku.
137
Bequaert (1926) gave a diagnosis for his new subgenus Oreumenes as follows:
"Medium-sized species; meso pleura without anterior epicnemial or posterior carina.
Mandibles of the usual Eumenes type. Vertex without hairy fovea in female. Antennae 12segmented in female; 13-segmented in male, the last segment short, of normal shape, not
forming a recurved hook. Hind margin of second abdominal tergite slightly duplicate." In
addition to the peculiar condition (Fig. 290) in the terminal segment of male antenna
among the species with a petiolated gastral segment 1, Oreumenes differs from the species
of Eumenes (of at least Japan) in the following points: pretegular carina present, female
clypeus straightly truncate apically (emarginate in the male) (Figs. 288, 289), the lateral
part of pronotum separated from the dorsal part by a carina which branches off from
pro notal carina, and the lateral part of propodeum rather clearly separated from the
posterior face by an edge. Giordani Soika (1941), Yamane (1982) and Carpenter (1986)
treated Oreumenes as a genus. Only one species (0. decoratus) occurs in Japan, Korea,
Taiwan and continental China (Tosawa, 1934; Vecht & Fischer, 1972).
In the Japanese species of Oreumenes, Delta, and Pseudozumia, the abscissa 3 of radial
vein of forewing is distinctly longer than abscissa 4, and cubital cell 3 is convex anteriorly.
Oreumenes decoratus (Smith)
(Figs.2~291,327)
Eumenes decoratus Smith, 1852, Trans. R. Entomol. Soc. Lond. (2)2: 36 (Sf )(type loc.: Teitung, N. China);
Esaki et al. 1938, Icon. Ins. Jpn. p. 360, fig. 631(1); Yano, 1950, Icon. Ins. Jpn. 2nd ed. p. 1454 (no. 41%);
Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat. Ed. 3: 292, pI. 246, fig. 20.
Eumenes jtzponicus Saussure, 1858, Rev. Mag. Zool. (2)10: 164 (Sf )(type loc.: Japan); Matsumura, 1911,
Thous. Ins. Jpn. Suppl. 3: 108-109, pI. 39, fig. 9; 1930, TIl. Thous. Ins. Jpn. 2: 11, pl.2, fig. 9; 1931,6000 III. Ins.
Jpn. p. 15 (no. 73).
Eumenes harmandi Perez, 1905, Bull. Mus. Hist. Nat. Paris, 11: 84-85 (Sf J')(type loc.: central Japan);
Yasumatsu, 1933, Mushi, 6: 31.
Japanese name: Suzubachi.
Diagnosis. Female and male. This species is the largest of the eumenids known from
Japan proper. Body length (h+th+t1+2): 21-23 mm in !f., 15.5-18.5 mm in 6'. Fore wing
length: 18.5-20.5 mm in !f., 15.5-16.5 mm in 6'. For the structural characteristics, see key to
the Japanese genera and the diagnosis for the genus. The color pattern is relatively stable
in the Japanese population. Black, with the following parts orange yellow: clypeus, a
longitudinal mark between antel'\nae (reaching downward to the upper margin of
clypeus, and often pOinted above), a line on the inner orbit below, a very narrow line
behind eye, basal half of antennal scape below, pronotum (posterior and lower lateral
portion black), tegula with a central transparent spot, parategula, a small spot under wing
base (often lost), scutellar crest, metanotum posteriorly, an irregular marking along the
lateral ridge of propodeum, propodeal orifice and valvae, a medially deeply incised apical
band on tergite 1, a wide regular band on tergite 2 apically, a narrower apical band on
sternite 2. Antenna (at least 2-3 apical segments) below, mandible in apical half, and legs
extensively brownish or ferruginous in both sexes.
Material examined. Hokkaido: 2J' J', Hakken-zan, Sapporo, 14 vii 1978 (T. Fujisawa).
Honshu: lwate-ken - 1 J', JohOji, 21 viii 1982 (HI), 1 Sf, Kanegasaki, 2 viii 1987 (SKY); Niigata-ken - 1 Sf,
Tsubame Spa, 15 viii 1981 (KB), IJ', Iwakuzure, 17 ix 1981 (KB); lbaraki-ken - ISf, Tsuchiura, 9 viii 1987
138
(SKY); Nagano-ken - H-, Amori, Nagano-shi, 24 viii 1979 (H. Fujisawa); Gifu-ken - 1 d', Hongo, Seki, 5 viii
1982 (Y. Takai), 2 d' d', same loc., 26 viii 1982 (Y. Takai).
Sado-ga-shima: 2 d' d'1-!f, Ogi, 13 vii 1980 (A. Seino), 1 d', Tagirisu, Mano, 23 ix 1981 (KB).
Kytlshtl: Nagasaki-ken - 1-!f, Haraguchi, Omura, 17 viii 1967 (R. Ohgushi), 1-!f, same loc., 3 ix 1967 (R
Ohgushi); Kagoshima-ken -1 d', Koyama, 2 vii 1978 (H. Nagase), 2 d' d'1-!f, Iriki, 3-6 ix 1984 (AN).
Tsushima Is.: Kami-agata -1-!f, TembOdai, 19 x 1979 (T. Kumata); Shimo"1lgata -1 d'1-!f, Izuhara (140-180
m alt.), 11 viii 1978 (K. Harusawa), 11 d' d'3-!f -!f, same loc., 27 vii 1986 (K. Nakamine), 1-!f, Tsutsu, 23 vi 1974
Hiura), 1-!f, Tatsura-yama, 29 ix 1977, 1-!f, same loc., 23 vii 1978 (0. Tominaga).
N. Ryukyus: Tane-ga-shima - 1 d'1-!f, 2-4 viii 1916 (H96, H97), 1-!f, Nakatane, 4 viii 1983 (SKY), 1-!f,
Utara,6 viii 1986 (M. Tatsuno); Yaku-shima - 2-!f -!f, Kurio, 2 xi 1975 (F. Komal), 1 d'2-!f -!f, Miyanoura, 8-11 viii
1981 (SKY), 1 d', Onoaida, 9 viii 1981 (SKY), 1 d', Miyanoura, 5 viii 1986 (SKY); Kuchinoerabu-jima - 1 d',
Hommura,21 vii 1989 (SKY).
S. Ryukyus: Ishigaki-jima - 1-!f , Shinkawa, 7 viii 1978 (M. Terayama).
a.
Distribution. Hokkaid6; Okushiri-t6 (Munakata, 1987); Honshu; Sado-ga-shima;
Shikoku; Kyushu; Tsushima Is. (Kami- & Shimo-agata); Got6 Is. (Fukue-jima); Chikuzenokino-shima; Osumi Is. (Tane-ga-shima; Yaku-shima; Kuchinoerabu-jima); Yaeyama Is.
(Ishigaki-jima). Korea; Taiwan; continental China.
Taxonomic notes. Tosawa (1934) stated that the specimens from Taiwan, Korea and
continental China examined by him agreed in coloration with Smith's original description
of Eumenes decoratus, while those from Japan proper with Perez's original description of E.
fu1rmandi. I have also found some differences between the Japanese specimens and Smith's
description. For example, Smith (1852) mentioned: "... its apical margin above [apical
margin of tergite 1], as well as those of the second segment, broadly, and the following
segments narrowly, orange yellow" (the holotype of E. decoratus is a female labelled
merely "N. China", though Smith gave Tein Tung as the type locality - Bequaert, 1928).
Lee's (1985) description agrees in these characters (p. 93). The Japanese specimens,
however, sometimes have a very narrow brownish part, but not a distinct orange band, on
Figs. 288-291. Oreumenes decoratus. 288, head in frontal view (-!f); 289, male dypeus; 290, apical
segments of male antenna; 291, gaster from above( -!f ).
139
each of tergites 3-5 apically. Furthermore, in the Chinese specimens the pronotum seems
nearly wholly orange except on the anterior vertical face, while in the Japanese ones it is
marked with orange only in the anterior 2/3 of dorsal part. If these differences are
constant, the Japanese population may deserve sub specific rank (in this case the name
japonicus Saussure will be applied).
I have examined one female from Ishigaki-jima, Southern Ryukyus, kindly sent by
Mr. M. Terayama. The specimen well agrees in coloration with those from the Japanese
mainlands, but distinctly differs from those of Taiwan. No additional specimen has been
collected on this island.
Biology. This species builds its mud nests on stones, wooden buildings, slender twigs
of living trees, and tree trunks (Iwata, 1953). Several pots, quartered- or hemi-spherical in
shape, are made in a cluster and the whole surface of the cluster is later covered with a
thick coating of mud paste. The female wasp hunts for the larvae, relatively large in size,
of geometrid moths. Tsuneki's (1980, 1982) continual observation of a second-generation
female revealed that she attended her nest for more than one month, constructing a total
of nine pots. Both mass and progressive provisioning were observed, and the first
offspring emerged before the mother left the nest. These facts strongly suggest that this
species is subsocial, though the social relation between the mother and her offspring is
unclear. This species is bivoltine in at least central and southwestern Japan (Tsuneki, 1980).
Additional observations were made by Katayama (1935, 1936) and Tsuneki (1978). A series
of photos given by Iwata et a1. (1982) illustrate the nest-building behavior.
Parasite: Pseudoxenos iwatai (Strepsiptera, Stylopidae). Parasitoids: Macrosiagon
nasuta, M. iwatai (Coleoptera, Rhipiphoridae), Acroricnus ambulator (Hymenoptera,
Ichneumonidae), Stilbum cyanurum (Chrysididae), Amobia signata (Diptera, Sarcophagidae)
and a species of Phoridae.
Genus Eumenes Latreille
Eumenes Latreille, 1802, Hist. Nat. Crust. Ins. 3: 360 (type species: Vespa coarctaia Linne, designated by
Latreille, 1810).
Alpha Saussure, 1855, Et. Fam. Vesp. 3: 128, 137, pI. 7 (name for Eumenes, "Ier Division" of Saussure,
1852, p. 28) (Type species: Vespa coarcta/a Linne, designated by Bequaert, 1926).
Japanese name: Tokkuribachi Zoku.
This genus is closely related to Oreumenes and Delta (including Phi of authors), with
which it shares the following character conditions: cephalic foveae absent, epicnemial
carina absent, tegula short, parategula produced posteriorly beyond tegula, gastral tergite
1 petiolate, fore leg with a well-defined carina on its outer face, propodeal orifice with a
pair of processes which can be seen when the gastral petiole is removed (Nakamine, 1987).
The following characteristics are also seen in Eumenes: clypeus more or less emarginate at
base and apex, male antenna with apically pointed hook (terminal segment), mesosoma
strongly convex and subglobular, pretegular carina absent, propodeum without inferior
and superior ridges, tergite 2 with a well-defined lamellate area at apical margin.
The genus is distributed all over the world. The Japanese forms have been studied by
Tosawa (1934), Sonan (1939), Giordani Soika (1941, 1973, 1986), and Yamane (1977a, b).
Five species (6 forms) are known to occur in Japan, one species (pundatus) being confined
to the Tsushima Islands. Most species are confined to the Japanese mainlands and some
140
relatively large islands. No species are found in the Central and Southern Ryukyus despite
the fact that four species occur in Taiwan. The following key is chiefly based upon
Giordani Soika (1941), Sato (1964), and Yamane (1977a).
Key to the Japanese forms of Eumenes
1. Gastral tergite 1 relatively stumpy, seen from above two times as long as wide (at the
widest part). Tergite 2 with a yellow spot on each lateral side. Apical half of femora,
tibiae, and tersi of all legs yellow or yellowish brown. Antennal hook of male recurved
to approach the base of segment 10.......................................................................................... 2
- Gastral tergite 1 relatively slender, seen from above more than two times as long as
wide. Tergite 2 usually without yellow spots (if spots present, female clypeus wholly
black). Female legs largely black or blackish brown. In the male, femora below, tibiae
and tarsi often yellowish, but femora above darker. Antennal hook smaller; its apex not
exceeding beyond the middle of segment 10.......................................................................... 3
2. Larger species; body length (h+th+t1+2) more than 12 mm. Female clypeus as wide as
high. Gastral sternites 4-6 with long, dense hairs in 6'. Hind trochanter and femur
below with long hairs in 6' ........................................................................... E. fraterculus D.T.
- Smaller species; body length (h+th+t1 +2) less than 12 mm. Female clypeus higher than
wide. Sternites 4-6 with much shorter hairs, and long hairs restricted to posterior
margins in 6'. Hind trochanter and femur below with inconspicuous hairs in 6' .
..................................................................................................................... E. rubrofemoratus G.S.
3. Base of tergite 2 dorsally swollen; in profile, tergite usually meets at a right angle with
sternite. Female clypeus wholly black; only rarely with a yellow marking. Male clypeus
with long hairs, not shining. Male antennal segment 10 in profile wider than long;
antennal hook recurved to reach segment 10 ........................................................................... 4
- Base of tergite 2 dorsally not swollen; in profile, the angle formed by tergite and sternite
acute. Female clypeus with large yellow markings. Male clypeus with much shorter
hairs, shining. Male antennal segment 10 in profile as long as wide; hook small, not
reaching segment 10 ...................................................... E. micado Cam. (=E. samuray Schult.)
4. Tergite 2 less elongate, less gibbous at base, and without yellow spots ............................. .
........................................................................................................ E. rubronotatus Per............... .5
- Tergite 2 more elongate, more gibbous at base, and with a yellow spot on each lateral
side .................................................................................................. E. pundatus pundatus Sauss.
5. Tergite 2 with dense and coarse punctation: in medio-apical part spaces between
punctures usually smaller than puncture diameter. Honshu - Kyushu and some
associated islands ..................................................................................... E. r. rubronotatus Per.
- Tergite 2 with sparser and finer punctation: in medio-apical part spaces between
punctures much larger than puncture diameter. Hokkaido ................ E. r. aquilonius Yam.
Key to the nests of Japanese Oreumenes and Eumenes excluding E. pundatus (adopted from
Iwata, 1971, translated by Smithsonian Institution, with slight modifications)
1. A few pots are made in a cluster and the whole surface of the cluster is later covered
with a thick coating of mud paste ............................................................................................. 2
141
- Each pot is prepared separately and is not coated with thick mud paste............................ 3
2. Each pot is large and quartered- or hemi-spherical in shape and the long axis is horizontal. ................................................................................................................................ O. decoratus
- Each pot is either hemispherical or subspherical and medium in size, with the mouth of
the pot near the center of the spherical surface.................................................. E. fraterculus
3. The pot is always attached to the surface (flat or depressed). The hemispherical shape is
the basic type and the pot is later reinforced and camouflaged with bast as the coating
material. ................................................................................................................ E. rubronotatus
- The pots may also be attached to a linear substratum ............................................................ 4
4. The pot is placed on a flat surface or on a linear substratum. The pot is oval and
medium in size and the collar is large ....................................................................... E. micado
- The pot is attached only to a linear substratum, elliptical and small in size ........................ .
.,............................................................................................................ ,.............. E. rubrofemoratus
Eumenes fraterculus Dalla Torre
(Figs. 292, 293, 302, 307, 311, 313, 324, 329, 333)
Eumenes fraterna Smith (nee Say, 1824), 1873, Trans. R. Entomol. Soc. Lond. 1873: 195 (!j'- )(type loc.:
"Hiogo", Honshu).
Eumenes fratercula Dalla Torre, 1894, Cat. Hym. p. 24 (new name for E. fraterna Smith); 1904, Gen. Ins.
19: 22 (in list); Tosawa, 1934, Trans. Kansai Entomol. Soc. 5: 13, fig. 1; Yano, 1950, Icon. Ins. Jpn. p. 1454;
Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat. Ed. 3: 292, pI. 146, figs. 16, 17.
Eumenes fraterculus Dalla Torre: Giordani Soika, 1941, Boll. Soc. Venez. Stor. Nat. 2: 134 (in key), 136,
figs. 1(1),3; Yamane, 1977, New Entomol. 26: 14-15, figs. 1,5,9,13,15,20.
Eumenes pomiformis F.: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 110 (no. 680), pI. 39, fig. 12; 1931,
6000 Ill. Ins. Jpn. p. 16 (no. 76) (misidentification).
Japanese name: Kiboshi-tokkuribachi.
Diagnosis. Female. Body length (h+th+t1+2): 12.0-16.0 mm. Fore wing length: 10.013.0 mm. Head wider than high, densely punctate, with long hairs on frons and vertex.
Clypeus nearly as wide as high (Fig. 292), weakly punctate. Thorax and propodeaum
densely punctate, with long hairs; punctation on scutellum and metanotum coarse.
Gastral tergite 1 densely punctate; punctation finer than on thorax. Tergite 2 densely
punctate; punctures become coarser toward apex. Hairs on gastral tergite 1 long; those on
other segments much shorter.
Black, with the following parts yellow: a wide basal band on dypeus (Fig. 292), an
interantennal mark not reaching the base of dypeus (Fig. 307), antennal scape below, a
band on pronotum anteriorly, a spot under wing base, tegula with a brownish median
part, a pair of spots on scutellum, metanotum largely, a spot on each side of propodeum
near base, a pair of small spots on tergite 1 (often lost), a pair of larger spots on tergite 2,
apical bands on tergites 1-4 (band on t2 widest) and stemite 2, and legs extensively.
Male. Body length (h+th+t1+2): 10.5-14.0 mm. Fore wing length: 9.0-11.0 mm. Similar
to the female in both structure and coloration. Clypeus distinctly higher than wide (Fig.
293), basally with relatively long hairs. Gastral stemites 4-6 with long hairs densely (Fig.
324). Trochanter and femur of hind leg below with long hairs (Fig. 311). Clypeus usually
wholly yellow. Interantennal bar usually reaching the base of clypeus. Antennal hook
large, almost reaching the base of segment 10 (Fig. 302).
142
297
298
Figs. 292-301. Clypeus of Japanese Eumenes. 292, fraterculus "?-; 293, ditto d'; 294, rubrofemDratus
"?-; 295, ditto d'; 296, rubronotatus -\,-; 297, ditto d'; 298, punctatus "?-; 299, ditto d'; 300, micado
"?- ; 301, ditto d'.
Material examined. Honshu: Niigata-ken - 2"?- -\,-, Fukushima-gata, 2 x 1977 (HI), 1"?-, same loc., 21 ix
1980 (HI); Fukui-ken -1"?-, 5anri-hama, 22 v 1971 (IT), ld'l"?-, Ono, 27 vi 1973 (TT); Hy6go-ken - 1"?-,
Sasayama, Tamba, 21 vii 1952 (K. Iwata), 1 d', Miki, 1 v 1968 (1<. Iwata).
Shikoku: K6chi-ken -1 d', Okoyama, Nankoku, 2 vii 1975 (51), 1"?-, Godaisan, Kfx:hi-shi, 27 v 1976 (51).
Kyushu: Nagasaki-ken - 1"?-, Haraguchi, Omura, 18 v 1967 (R. Ohgushi); Kagoshima-ken - 1"?-, Kanoya, 8
ix 1981 (51),1"?-, Ibusuki, 16 x 1988 (SKY).
Distribution. Honshu; Shikoku; KyushU; Goto Is. (Fukue-jima). E. Siberia (Kurzenko,
1984a).
Biology. This species is widely distributed on the mainlands of Japan except
Hokkaido. Though it was common when Iwata (1953) studied its biology, recent collection
efforts show that it has become much rarer probably due to the deterioration in rural
environments. This species is bivoltine and builds mud nests among the foliage of trees or
143
bushes, attaching them to plant stems, in summer, but it prefers sunny stone surfaces in
late fall (Fig. 333). A few pots are made in a cluster and the whole surface of the cluster is
later covered with a thick coating of mud paste; each pot is either hemispherical or
subspherical, with the mouth near the center of the spherical surface. Four to 16
caterpillars chiefly of noctuid moths are stored in a cell (Iwata, 1953, 1978a, 1980b; also see
Masuda, 1941). Nesting behavior is illustrated by color photos in Iwata et al. (1982).
Parasitoids: Acroricnus ambulator (Hymenoptera, Ichneumonidae), Chrysis apicata
(Hymenoptera, Chrysididae), Amobia signata (Diptera, Sarcophagidae), Macrosiagon nasuta
and M. iwatai (Coleoptera, Rhipiphoridae). For the biology of M. nasuta, see also Iwata
(1939c).
Eumenes rubrofemoratus Giordani Soika
(Figs. 294, 295, 303, 312, 325, 330, 334)
"Eumenes rubrofemoratus Perez": Tosawa, 1934, Trans. Kansai Entomol. Soc. 5: 4 (key), 7, pI. 1, fig. 7
Oapsus for E. rubronotatus Perez).
Eumenes rubrofemoratus (Tos.) n. sp.: Giordani Soika, 1941, Boll. Soc. Venez. Stor. Nat. 2: 135, 145, fig. 1
(9,10) (Sf 3')(type loc.: Japan).
Eumenes rubrofemoratus Tosawa: Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat. Ed. 3: 292.
Eumenes rubrofemoratus Giordani Soika: Vecht and Fischer, 1972, Hym. Cat. (n. ed.) 8: 132; Yamane,
1977, New Entomol. 26: 15-16, figs. 2, 6,14,21; Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35: 156.
Japanese name: Kiashi-tokkuribachi.
Diagnosis. Female and male. Body length (hHh+t1+2): 10.0-11.5 mm in .<f.,9.0-1O.0
mm in d'. Fore wing length: 9.0-9.5 mm in .<f., ca. 8 mm in d'. Similar to E. fraterculus, but
differs from the latter in the following points: much smaller, body length less than 12 mm;
female clypeus higher than wide (Fig. 294); emarginated part of male clypeus slightly
wider than the base of clypeus (Fig. 295); antennal hook relatively short, with short hairs
on its inner face (Fig. 303); gastral sternites 4-6 of the male with much shorter hairs, and
long hairs confined to posterior margins (Fig. 325); hind trochanter and femur of the male
below with inconspicuous hairs (Fig. 312); female antenna below sometimes extensively
yellowish brown or ferruginous; yellow spots on scutellum always lost.
Material examined. Honshu: lwate-ken - 1 Sf, Takizawa, 20 ix 1979 (YM); Niigata-ken - 1 Sf, Shidai-hama,
29 ix 1974 (HI); Fukui-ken -13', Sanri-hama, 30 v 1973 (TT), 1Sf, same loc., 20 vi 1976 (H. Kurokawa);
Yamaguchi-ken - 1 Sf, BMu, 29 vi 1976 (reared by S. Nagai); Hyogo-ken - 1 Sf, Sasayama, Tamba, 21 ix 1952 (K
Iwata); Okayama-ken -1 (f', Nonoguchi, 16 vii 1960 (R. Momoi).
Kyflshfl: Nagasaki-ken - 13', Haraguchi, Omura, 9 x 1966 (R. Ohgushi), 1 Sf, same loc., 5 xi 1966 (R.
Ohgushi).
Distribution. Honshu; Shikoku; Kyushu.
Biology. Like the preceding species, E. rubrofemoratus was common in rural areas of
southwestern Japan during the first half of 19OOs. Now it may be the rarest of the five
Japanese Eumenes species, though rather common in the Kanto District, Honshu
(Hisamatsu et al., 1986). Tsuneki (1929) and Iwata (1953) studied the biology of this
species. Two generations occur in a year. The female wasp builds her nest on dry stems of
herbs (mainly Rumex) in the grass from late spring to summer, while on stems or blades of
small gramineous plants quite near the ground in late fall (Fig. 334). The elliptical pots are
prepared separately, not coated with thick mud paste, and always attached to a linear
substrate. Many horny projections are occasionally built on mud surface. Larvae of
144
various moths (Geometridae; Pyralidae; Olethreutidae) are captured for the young.
Parasite: Pseudoxenos iwatai (Strepsiptera, Stylopidae). Parasitoids: Acroricnus
ambulator, Chrysis cyanurum, Amobia signata and Macrosiagon nasuta.
Eumenes rubronotatus Perez
(Figs. 296, 297, 304, 30B, 309, 314, 316, 319-321, 32B, 335)
Eumenes rubronotatus Perez, 1905, Bull. Mus. Hist. Nat. Paris, 11: 85 (J')(type loc.: Yokohama, Honshu).
Eumenes dimidiatidypeus Giordani Soika, 1973, Boll. Mus. Civ. Stor. Nat. Venez. 24: 127-129 (!f- )(Type
loc.: Tokyo).
Japanese names: Mumon-tokkuribachi (Samurai-tokkuribachi).
Diagnosis. Body length (h+th+t1 +2): 10.0-13.0 mm in ~, 8.0-11.0 mm in 6'. Fore wing
length: 10.0-11.5 mm in ~,B.0-9.0 mm in 6'. The following character conditions separate
this species from the other Japanese congeners: head distinctly wider than high; female
clypeus higher than wide, without yellow marking (Fig. 296); male clypeus with relatively
long hairs, punctate, not shining; antenna! hook medium in size, reaching the middle of
segment 10 (Fig. 304); in profile the tergite meets sternite at a right angle at the base of
gastral segment 2 (Fig. 316); tergites 1 and 2 coarsely and densely punctate; tergite 2
usually without yellow spots; sternite 2 finely and sparsely punctate; on other tergites and
sternites punctation much finer.
This is the most common species of the Japanese Eumenes, and occurs even on some
small islands such as Kuro-shima and Kuchinoerabu-jima in the N. Ryukyus.
Eumenes rubronotatus rubronotatus Perez
(Figs. 297, 30B, 309, 320, 321, 32B, 335)
Eumenes rubronotatus; Giordani Soika, 1941, Boll. Soc. Venez. Stor. Nat. 2: 132 (in key), 144, figs 1(8, 10),
2(6); Yamane, 1977, New Entomol. 26: 16-17, figs. 3, 7, 10, 16, 18; Giordani Soika, 1986, Boll. Mus. Civ. Stor.
Nat. Venez. 35: 156.
Eumenes architectus Smith: Tosawa, 1934, Trans. Kansai Entomol. Soc. 5: 6-7, fig. 8 (misidentification).
Eumenes samuray Schulthess: Ishikawa, 1965, Icon. Ins. Jpn. Col. Nat. Ed. 3: 292 (no. 19), pI. 146, fig. 9
(misidentification).
Japanese name: Mumon-tokkuribachi.
Diagnosis. Body black, with the following parts yellow: male clypeus almost wholly
(Fig. 297; sometimes with irregular black markings; female c1ypeus usually wholly black,
but very rarely with yellow spots), interantennal bar approaching close to the base of
clypeus in the female (Fig. 309), a line behind eye, anterior band on pronotum (much
reduced in the male), spot under wing base, a spot on tegula posteriorly, metanotum (in
the male often wholly black), a narrow apical band on tergite 1 (medially concave), a
relatively wide band on tergite 2 (medially narrowed), a narrow apical band on stemite 2.
Scutellum never marked with yellow. Tergite 2 very rarely with a yellow spot on each
side.
Material examined. Honshu: Iwate-ken - 1!f-, Kuriyagawa, Morioka, 7 ix 1970 (yM), 1 J', Ashiro, 22 ix
1974 (yM), 1!f-, Takizawa, 26 ix 1976 (yM & T. Matsumura), 1!f-, Oshuku, Shizukuishi, 7 viii 1987 (SKY);
145
308 /'.
0(1)0
AJ\
304
0 '"",I') (0"
\
310~
305
O\,T)O
309~
Figs. 302-306. Segments 10-13 of male antenna in Japanese Eumenes (305, original; others, after
Yamane, 1977a). 302, fraterculus; 303, rubrofemoratus; 304, rubronotatus; 305, punctatus; 306,
micado.
Figs. 307-310. Interantennal marking (after Yamane, 1977a). 307, fraterculus ~; 308, rubronotatus
J'; 309, ditto ~ ;310, micado ~.
Figs. 311, 312. Hind trochanter and femur (J') (after Yamane, 1977a). 311, fraterculus; 312,
rubrofemora tus.
Yamagata-ken - 1 J', Chojagahara, Oguni, 22 vi 1979; Fukushima-ken - 1 ~, Mishima, 24 vi 1980 (HI); Niigataken - 1 J', Muroya, Mikawa, 21 viii 1977 (HI), 1 J', Kami-ishikawa, Shibata, 12 viii 1979 (HI), 1 J'1 ~, Shiori
Pass (1000 m alt.), Komaga-take, 1 ix 1979 (HI), 1 ~,Senami, 3 vii 1980 (KB), 1 J', same loc., 20 vi 1981 (KB),1
~,same loc., 17 vii 1981 (KB), 1 J', same loc., 30 vii 1981 (KB), 1 ~,same loc., 28 viii 1981 (KB), 1 J', same
loc., 3 ix 1981 (KB), 1 J'1 ~, Iwakuzure, 17 ix 1981 (KB); Ibaraki.Jcen - 1 J', Tsuchiura, 9 viii 1987 (SKY);
Saitama-ken - 1 J', Ageo, 23 vii 1983 (T. Sunose); Nagano-ken - 1 ~, Yokoyama, Ina, 17 ix 1%1 (YM), 1 ~,
Habiro, Ina, 27 viii 1962 (yM), 1 t, Yokoyama, Ina, 31 viii 1962 (YM); Gifu-ken - 1 d'3 ~ ~, Horado, 7 viii 1982
(Y. Takai), 1 ~,Hong<'Kho, Seki, 11 viii 1982 (Y. Takai), 1 ~, Tonohora, Seki, 2 viii 1982 (Y. Takai); Aichi.Jcen - 5
~ ~, Jokoji, 3 x 1976 (yM); Wakayama.Jcen - 1 J', Koza, 25 ix 1974 (5. Takagi); Shimane.Jcen - 1 ~, Sakane,
Izumo, 7x 1976 (T. Sunose); Tottori-ken -1~, Mt. Daisen, 31 x 1%1 (H. Fujii).
Sado-ga-shima: 1 ~, Kobayashi, 23-5 x 1976 (A. Seino).
Oki Is.: Nishi-no-ilhima -1 ~,Urago, 11 v 1982 (YM).
Shikoku: KOchi.Jcen - 1 J', Monobe, Nankoku, 9 v 1974 (51), 1 ~, Godaisan, K6chi-shi, 7 v 1975 (SI), 1 J',
same loc., 22 v 1975 (51), 1 ~, Monobe, Nankoku, 24 vii 1975 (51).
Kyushu: Fukuoka.Jcen - 2~ ~, Hakozaki, Fukuoka-shi, 18 ix 1958 (yM), 1 d'1~, same loc., 2 vii 1959
(yM), 1 J', Kashii, 16 ix 1959 (YM), 1 ~, Mii-machi, Kurume, 2 vi 1960 (YM); Nagasaki.Jcen - 1 ~, Haraguchi,
Omura, 5 vi 1966 (R. Ohgushi), 1 J', same loc., 26 viii 1966 (R. Ohgushi); Kumamoto-ken - 1 ~, Toyo-mura, 8
viii 1983 (M. Maeda); Kagoshima.Jcen - 1 ~,Osaki, Kagoshima-shi, 4 x 1977 (H. Nagase), 1 d', Uchinoura, 16 x
1977 (H. Nagase), 1 J', Koyama, 2 vii 1978 (H. Nagase), 1 J', Shiroyama, Kagoshima-shi, 4 vii 1981 (SKY),1
146
313
...........
··~~~~.-. . t· . '"'-. .-·/
315
320
~321
325
Figs. 313-315. Gastral tergite 1 from above in Japanese Eumenes. 313, fraterculus; 314, rubronotatus;
315, micado.
Figs. 316-318. Gastral segment 2 in profile (316, original; others, after Yamane 1977a). 316,
rubronotatus; 317, punctatus; 318, micado.
Figs. 319-323. Apical part of gastral tergite 2 from above (after Yamane, 1977a,b). 319, rubronotatus
aquilonius; 320, r. rubronotatus (typical); 321, ditto (rare condition); 322, micado (samuray-type);
323, ditto (micado-type).
Figs. 324-326. Male gastral sternites in profile (after Yamane, 1977a). 324, fraterculus; 325,
rubrofemoratus; 326, mica do.
147
d', same loc., 10 vii 1981 (SKY), 1!f, Kirishima-jingil, 23 vii 1981 (SKY), 4 d' d', Shiroyama, Kagoshima~hi, 30
vii 1981 (SKY), 7 d' d'1!f, Korimoto, Kagoshima-shi, 7 viii 1981 (SKY), 1 d', same loc., 17 viii 1981 (SKY), 1 d',
Meiwa, Kagoshima-shi, 15 viii 1981 (SKY), 2d' d', Irino, Ei, 11 ix 1983 (M. Ohara), 2d' d', Haruyama,
Kagoshima-shi, 11 ix 1983 (M. Ohara), 1!f, Nagashiro-bokujO, 20 v 1984 (M. Ohara), 1 d', Eboshi-dake, 25 vii
1984 (AN), 1!f , Takeyama nr Yamakawa, 6 viii 1984 (M. Maegata), 1 !f, Iriki, 5 ix 1984 (AN), 1 d', Kaimondake, 26 viii 1986 (SKY), 1!f, Shiroyama, Kagoshima-shi, 20 ix 1987 (SKY).
Islands located close to Kagashima-ken-hondo: Akune-lishima - 3!f !f, 5 viii 1983 (SKY); Naga-shima - 4
d' d', 27 viii 1984 (SKY); Take-shima (nr Naga-shima) -1 d', 28 viii 1984 (SKY); Kamikoshiki-jima -1 d', 5 ix 1984
(M. Maegata).
N. Ryukyus: Kuro-shima - 246' d'11!f !f, asato, 29 viii-4 ix 1981 (SKY); Tane-ga-shima - 1!f, Hirayama, 9
viii 1916 (H114), 1!f, Nishino-omote, 31 vii 1982 (KT), 2 d' d'2!f !f, Hamada, 1-2 viii 1984 (SKY), 16', Ikeno,
21 vii 1984 (5. Watahiki); Mage-shima -1 d', 22 vii 1984 (5. Watahiki); Yaku-shima - 1 d'4!f!f, Miyanoura, 8-11
viii 1981 (SKY), 2 d' 6', Kusugawa, 9 viii 1981 (SKY), 1!f, Onoaida, 9 viii 1981 (SKY), 1!f, Shitogo, 10 viii 1981
(SKY), 3 d' d'1!f, Onoaida (40-200 m alt.), 27-9 vi 1982 (51), 1 d'1!f, Miyanoura (O~ m alt.), 26-8 vi 1982 (51);
Kuchinoerabu-jima -1!f, Hommura, 18 v 1989 (H. Watanabe); 1 d'1!f, Shin-dake, 21 vii 1979 (H. Watanabe).
Distribution. Honshu; Sado-ga-shima; Oki Is. (Nishino-shima); Shikoku; Kyushu;
Goshoura-jima; Akune-Oshima; Naga-shima; Take-shima (nr Naga-shima); Kamikoshikijima; Osumi Is. (Kuro-shima; Tane-ga-shima; Mage-shima; Yaku-shima; Kuchinoerabujima). Korea (new record); China (Kwantung).
Biology. The biology of this form has been studied by Iwata (1953) and reviewed in
Iwata (1971, 1978a, 1980b). Unfortunately Iwata (1953, 1971) used the incorrect name (E.
samuray Schulthess) for this species, and Vecht and Fischer (1972) cited his record without
correction. His misapplication may have been based upon Yasumatsu's misidentification.
Later, Iwata (1980b) used the correct Japanese name "Mumon-tokkuribachi", with no
scientific name, instead of "Samurai-tokkuribachi" erroneously applied in his earlier
publications.
This species flies from early May to mid October, and is probably bivoltine. Female
wasps build their nests on the stone surface, especially in depressions (Fig. 335). The pots,
hemispherical in shape, are prepared separately and not coated with thick mud paste. It is
later reinforced and camouflaged with bast as the coating material. The greater part of the
prey consists of larvae of Gelasma illiturata Walker (Geometridae); when they are scarce,
caterpillars of other geometrids are also hunted.
Parasitoids: Acroricnus ambulator, Chrysis sp. and Amobia signata.
Eumenes rubronotatus aquilonius Sk. Yamane
(Figs. 319, 328)
Eumenes rubronotatu5 aquilonius Yamane, 1977, New Entomol. 26: 59-61, figs. 1,2, 5,8 (!f d')(type loc.:
Sapporo, Hokkaido).
Diagnosis. This subspecies differs from the nominotypical subspecies in the following
points: body slightly smaller; gastral tergite 2 very feebly and sparsely punctate, polished;
spaces between punctures distinctly larger than puncture diameter (Fig. 319); sternite 2
almost impunctate; male dypeus black, with an irregular yellow marking; anterior band
on pronotum reduced, sometimes lost.
Material examined. Hokkaido: 1!f, Sapporo, 22 vi 1924 (H. Kono), 1 d'1!f, Jozankei, Sapporo, 16 vii
1926 (5. Matsumura) (holotype), 1!f, Maruyama, Sapporo, 19 vii 1928 (T. Uchida), 1!f, Sapporo, 25 viii 1935
(y. Sugihara), 1 d', same loc., 21 vi 1953 (C. Watanabe), 1 d', Otarunai near Sapporo, 14 ix 1968 (SY), 1!f,
Heiwa, Sapporo, 26 viii 1974 (SKY).
148
\..~-
.
o.
decoratus
327
OE. rubronotatus aquilonius
r. rubronotatus
:If E. punctatus
.E.
Figs. 327,328. Distribution of Oreumenes decoratus and two Eumenes species in Japan.
Distribution. Hokkaido.
Biology. No information is available. The collection records suggest that this form is
bivoltine.
149
Eumenes punctatus punctatus Saussure
(Figs. 298, 299, 305, 317, 328)
Eumenes punctatus Saussure, 1852, Et. Fam. Vesp. 1: 37 (!f J')(type loc.: China); Giordani Soika, 1941,
Boll. Soc. Venez. Stor. Nat. 2: 132 (in key), 142-143, figs. 1(7), 2(5); 1976, Ann. Hist.-Nat. Mus. Hung. 68: 295
(from Korea).
Eumenes architectus Smith: Yasumatsu, 1936, Insects of Jehol, 7 (Eumenidae): 1-3, 10-12, pI. 1, fig. 1
(from Manchoukuo; misidentification).
Eumenes pundatus punctatus Saussure: Yamane, 1977, New Entomol. 26: 61 (from Korea).
Japanese name: Tairiku-tokkuribachi.
"
):-:~.
.• .J
E.
fraterculus
329
,
o
E. rubrofemoratus
330
E. micado
331
Figs. 329-331. Distribution of three Eumenes species in Japan.
150
@}>
332
•.,.'1
.
.
•
335
~~~©
336
334
Figs. 332-336. Nests of Japanese Oreumtmes and Eumenes (after Iwata, 1980c). 332, O. decoratus; 333,
E. fraterculus (A,B, summer nest; C, fall nest); 334, E. rubrofemoratus (A, summer nest; B, fall
nest); 335, E. rubronotalus (left, pot made in a stone depression; right, section of a nest); 336, E.
micado.
Diagnosis. Female and male. Body length (hHh+t1+2): 11.0-13.0 mm in if., 10.0-11.0
mm in d'. Fore wing length: 10.0-11.5 mm in if., 8.5-9.0 mm in d'. Good descriptions are
given in Yasumatsu (1936) and Giordani Soika (1941). Here, important character
conditions useful in separating this form from the closely related E. rubronotatus will be
given. Head, thorax and propodeum much more finely and densely punctate; spaces
between punctures not forming carinae. Clypeus more shallowly emarginate at base and
more deeply at apex in both sexes (Figs. 298, 299). Punctation on gastral tergites 1 and 2
also much finer. Propodeal groove less developed, almost absent near the base. Tergite 2
more swollen dorsally, preapical depression more pronounced (Fig. 317).
Tergite 2 almost always with a yellow marking on each side (Fig. 317) (Korean
specimens often lacking the spots). Female legs black, but orange or ferruginous parts
more extensive: fore femur apically, outer face of fore tibia in apical 1/3, mid and hind
tibiae in basal half marked with these colors. Fore and mid tibiae of male legs yellowish
on outer face; tarsi extensively ferruginous.
Material examined. Tsushima Is.: Kami-1lgata - 2 Sf- Sf-, Sasuna - Nembutsu-zaka, 22 x 1975 (I. Hiura), 1 d',
Sasuna, 17 ix 1977 (Tominaga), 2 Sf- Sf-, Shiohama, Nii, 24 viii 1979 (I. Hiura), 1 Sf-, Oboshi-yama, 24 viii 1979
(A. Seino); Shimo-1lgata - 1 d'1 Sf-, Tsutsu, 6 viii 1967 (yH), 1 Sf-, Banshoin, Izuhara, 18 viii 1968 (K. Tani), 1 d',
151
Tsutsu-misaki,22 viii 1979 (I. Hiura).
Distribution. Tsushima Is. (Kami-agata; Shimo-agata). Korea; China. "E. fratercula"
recorded from Izuhara, Asamo and Uchiyama (Tsushima Is.) by Tano (1966) is E. punctatus
(I have examined the female from Tano's collection).
Taxonomic notes. In Japan this form may easily be distinguished from the allopatric E.
r. rubronotatus according to localities. In Korea, however, where these species seem
completely sympatric, separation of them is rather difficult. E. pundatus often lacks yellow
gastral spots, and punctation is intermediate in size and condition between the Japanese
populations of E. pundatus and E. rubronotatus. The following character conditions may be
useful in separating E. punctatus from E. rubronotatus in Korea: propodeal groove
shallower, almost absent near the base of propodeurn (in rubronotatus the groove is more
distinct and discernible over the whole length of propodeum); tegula yellow with the
basal area blackish and with a brownish median spot (in rubronotatus the tegula almost
wholly blackish); metanotum usually with a yellow band (in rubronotatus the band very
often lost); tibiae of all legs in the female usually brownish (in rubronotatus legs darker);
tibiae of all legs in the male extensively yellow (in rubronotatus legs much darker, at least
hind tibia wholly blackish); male clypeus relatively elongate.
Biology. Nothing is known in Japan.
Eumenes micado Cameron
(Figs. 300, 301, 306, 310, 315, 318, 322, 323, 326, 331, 336)
Eumenes micado Cameron, 1904, Entomologist 37: 35 (!f) (type loc.: "Sharo-kowa", Japan); Bequaert,
1928, Ann. Mag. Nat. Hist. 10(2): 161; Tosawa, 1934, Trans. Kansai Entomol. Soc. 5: 5 (in key), 11, pI. 2, fig. 2
(E. mikado (!»; Giordani Soika, 1941, Boll. Soc. Venez. Stor. Nat. 2: 135 (in key), 137, figs. 1(2),2(1,8); Yano,
1950, Icon. Ins. Jpn. 2nd ed. p. 1454 (no. 4195); Ishikawa, 1%5, Icon. Ins. Jpn. Col. Nat. Ed. 3: 292, pI. 146, fig.
18; Yamane, 1977, New Entomol. 26: 49, figs. 4, 8, 12, 17, 19,22,24.
Eumenes samuray Schulthess, 1~, Mitt. Schweiz. Entomol. Ges. 11: 284 (6'!f )(type loc.: Yokohama (?);
Nagasaki); Giordani Sojka, 1941, Boll. Soc. Venez. Stor. Nat. 2: 135 (samurayi (!), in key), 137, fig. 2(7);
Yamane, 1977, New Entomol. 26: 49, fig. 23.
Eumenes samurayi (1) YUfescens Giordani Soika, 1973, Boll. Mus. Civ. Stor. Nat. Venezia, 24: 127 (6')(type
loc.: Tokyo). Syn. nov.
Japanese names: Mikado-tokkuribachi (Samurai-tokkuribachi).
Diagnosis. Female. Body length (h+th+t1+2): 13.0-15.0 mm. Fore wing length: 10.512.0 mm. Head distinctly wider than high, densely punctate; punctation on gena sparser.
Hairs on frons and vertex long. Clypeus higher than wide (Fig. 300), with shorter hairs;
punctures sparser and shallower than those on frons. Alitrunk longer than wide, densely
punctate, with long hairs. Metapleuron almost impunctate. Propodeum with shallow
vertical groove. Gastral petiole coarsely and densely punctate, nearly as long as tergite 2.
In profile the angle formed by tergite and sternite at the base of segment 2 less than 90"
(Fig. 318) . Punctation on tergite 2 much finer than on petiole (in the spring specimens
punctation still finer and sparser, and in the Hokkaido population even the summer
individuals show such a tendency). Sternite 2 shining, with very sparse, superficial
punctures. Other tergites and sternites with micropunctures alone.
Black, with the following parts yellow or orange: paired elongate markings at the
base of clypeus (sometimes clypeus basally extensively yellow), a pair of smaller spots
152
below the elongate markings (often lost) (Fig. 300), interantennal bar (Fig. 310), narrow
short line behind eye, pronotum anteriorly, a pair of comma-shaped markings on
mesoscutum anteriorly (often lost), a large spot under wing base, tegula with a median
blackish spot, a pair of square markings on scutellum, meta not urn largely, a pair of
irregular markings on propodeum (often lost), narrow apical band on tergite 1, relatively
wide apical band on tergite 2 (medially narrowed), a small spot on each side of tergite 2
(only rarely seen), narrow apical band on sternite 2, outer face of mid coxa, apical part of
femora of all legs. Two or three apical segments of antenna below ferruginous. Tibiae and
tarsi of all legs ferruginous to a varying degree.
Male. Length (h+th+t1+2) 11.5-14.0 mm. Fore wing length 9.5-11.0 mm. Much as in
the female. Antennal hook ferruginous in color, small, not reaching the apex of segment 10
(Fig. 306). Clypeus wholly yellow. Antennal scape (at least basal part) below yellow.
Mesoscutum usually wholly black. Yellow markings on scutellum much reduced, often
lost. Propodeum usually without markings. Apical band on sternite 2 wider than in the
female, with a pair of distinct projections. Legs more extensively marked with yellow.
Material examined. Hokkaido: 1 d', Maruyama, Sapporo, 29 viii 1926 (T. Uchida).
HonshU: lwate-ken - 15f-, Kuriyagawa, Morioka, 25 viii 1969 {yM), 2 d' d', Kanegasaki, 2 viii 1987 (SKY);
Yamagata-ken -1 d', Nukumi-daira, Oguni, 6 ix 1982 (KB); Miyagi-ken -1 d', Rifu, 31 viii 1980 (1(. Goukon), 1
!f, same loc., 14 ix 1980 (K. Goukon), 1 d', Aoba-yama, 5 ix 1980 (1(. Goukon); Fukushima-ken -1 d', Kashima,
28 ix 1973 (yM); Niigata-ken -1!f, "Echigo", 9 vii 1933 (Nohira), 1!f, Shiori-toge (1000 m alt.), Komaga-take, 1
ix 1979 (HI), 1 d', Senami, 13 viii 1981 (KB), 1 d', same loc., 5 ix 1981 (l(B), 1 d', Iwakuzure, 17 ix 1981 (l(B),2
d' d', Gozu-san, Suibara, 20 ix 1981 (HI); Saitama-ken - 1 !f, Ageo, 31 vii 1980 (f. Sunose), 1 d'2!f !f, same loc.,
23 vii 1983 (f. Sunose); Chiba-ken - 2d' d', Yokodo, 3 ix 1981 (J. Tsukahara); Ibaraki-ken -1!f, Tsuchiura, 9 viii
1987 (SKY); Nagano-ken - 2 d' d', Minamiminowa, Ina, 25 v 1962 (YM), 2d' d', same loc., 19-21 viii 1961 (YM),
5d' d', same loc., 25 viii - 2 ix 1961 (YM), 3 d' d', same loc., 5-8 ix 1%1 (YM), 4 d' d', same loc., 11-14 ix 1%1
(YM), 1 d', 29 ix 1961 (yM); Gifu-ken - 1!f, Katachi, Mino, 7 viii 1982 (Y. Takai), 15f-, Hongo, Seki, 11 viii 1982
(Y. Takai), Id', Otogari, Mino, 14 ix 1986 (Y. Takai); Nara-ken - Id', Dorokawa, 29 vii 1955 (0. Sato);
Wakayama-ken - 1!f, Torokyo, 31 viii 1951 (0. Tsuzimoto).
Sado-ga-shima: 1 d', Mt. Myoken, 12 viii 1979 (N. Kato), 2d' d', Tagirisu, Mano, 23 ix 1981 (KB).
Shikoku: K6chi-ken -1!f, Kojigamori, 12 vi 1960 (0. Sato), 1!f, Okoyama, Nankoku, 17 ix 1976 (51).
Kyushu: Fukuoka-ken - 5d'd', Kurogi, 28-29 vii 1983 (Y. Takai), 1!f, same loc., 30 viii 1984 (Y. Takai);
Nagasaki-ken -1 d'15f-, Haraguchi, Omura, 18-27 viii 1%7 (R. Ohgushi); Kumamolcrken -1 d', ToYO-mura, 8 viii
1983 (M. Maeda); Kagoshima-ken - 5d'd'1!f, Korimoto, Kagoshima-shi, 6-7 viii 1981 (SKY), 25f-5f-, Meiwa,
Kagoshima-shi, 15 viii 1981 (SKY), 3 d' d'15f-, Korimoto, Kagoshima-shi, 22-26 viii 1981 (SKY), 1 d'15f-,
Haruyama, 11 ix 1983 (M. Ohara), 1 d', Eboshi-dake, 25 vii 1984 (AN), 1 d', Iriki, 5 ix 1984 (AN), 1 d'15f-,
Kaimon-dake, 26 viii 1986 (SKY).
Islands located close to Kagoshima-ken-hondo: Naga-shima - 3d' d', Shoura, 27 viii 1984 (SKY); Takeshima (nr Naga-shima) - 2 d' d', 28 viii 1984 (SKY).
N. Ryukyus: Tane-ga-5hima - 1 d', Ikeno, 21 vii 1984 (S. Watahiki); 1-'?-, Hamada, 2 viii 1984 (SKY) Yakushima - 2d' d', Miyanoura, 8 viii 1981 (SKY), 2d' d', Onoaida, 9 viii 1981 (SKY), 1 d', Shitogo, 10 viii 1981
(SKY), 2 d' d', Miyanoura, 5 viii 1986 (SKY).
Distribution. Hokkaidf>; Honshu; Sado-ga-shima; Awaji-shima; Shikoku; Kyushu;
Tsushima Is.; Gotf> Is. (Fukue-jima; Naru-jima); Naga-shima; Take-shima (nr Naga-shima);
Osumi Is. (Tane-ga-shima; Yaku-shima).
Taxonomic notes. There have been recognized two forms within this species in Japan.
Giordani Soika (1941) called one of them "micado", in which the punctation on gastral
tergite 2 is quite fine and superficial (Fig. 323) and the yellow body markings are less
extensive. In the other, called "samuray", the punctation on tergite 2 coarser (Fig. 322) and
the body is more extensively marked with yellow. Giordani Soika treated these two forms
as distinct species, and his view has been followed by Sato (1964), Vecht and Fischer (1972)
153
and Yamane (1977a). Cameron (1904), however, mentioned in his original description of E.
micado "the dilated part [of the first segment] strongly and closely punctured .... ; the second
segment closely and much more finely punctured", and "a large irregular mark, broader
than long and with irregular edges, on the sides of the metanotum [=propodeum]". These
conditions are those more frequently met in "samuray" sensu Giordani Soika. Bequaert
(1928) wrote: "The holotype of E. micado is a female from Japan, which does not differ from
the description of E. samuray, nor from two specimens in the British Museum Collection
determined as E. samuray by A. v. Schulthess. The two specimens, a female (received from
A. v. Schulthess as the "type") and a male (labelled "cotype") from Japan, are, I presume,
paratypes of E. samuray". Tosawa (1934), Iwata (1953) and Ishikawa (1965) correctly
recognized only one species (micado) (note that Iwata's and Ishikawa's "samuray" no doubt
correspond to E. rubronotatus as discussed earlier).
Suda (1979) studied the variation in structure and coloration with many specimens of
"micado" and "samuray" from the Kant6 District, Honshu. He concluded that "micado" was
represented by spring individuals while "samuray" by summer I faIl individuals. In
mountainous regions both the "micado"-type and "samuray"-type wasps were collected in
summer. After careful considerations on variation, distribution and life cycle, he suspected
that "micado" and "samuray" are spring and summer generations of one and the same
species (micado). In this paper I have followed his reasoning.
Biology. This common species flies from May to October, and is probably bivoltine.
The biology of this species was studied by Iwata (1953, 1978a, 1980c). It nests on the
surface of wooden buildings as well as on slender twigs of various kinds of shrubs or
trees, but most often on slender rootlets under overhanging cliffs. The mud pot, oval and
provided with a large collar, is prepared separately and is not coated with thick mud paste
(Fig. 336). Larvae of various moth families are hunted (Iwata, 1978a).
Parasitoid: Acroricnus ambulator.
Genus Delta Saussure
Delta Saussure, 1855, Et. Fam. Vesp. 3: 130, 132, 143 (as division of genus Eumenes Latreille; "Eumenes,
lIe et me Divisions" in Saussure, 1852) (type species: Vespa maxiIlosa DeGeer, 1775 (=Delta emarginatum (L.»,
designated by Bequaert, 1926); Dalla Torre, 1904, Gen. Ins. 19: 20 (as group of genus Eumenes).
Phi Saussure, 1855, Et. Fam. Vesp. 3: 132, 145 (as division of Eumenes Latreille; "Eumenes lye Division"
in Saussure, 1852) (type species: Vespa arenata Fabricius, 1775, deSignated by Bequaert, 1926); Dalla Torre,
1904, Gen. Ins. 19: 20 (as group of Eumenes Latreille).
Japanese names: Nettai-suzubachi Zoku (Kurosuji-suzubachi Zoku; Haranagasuzubachi Zoku).
Japanese authors have dealt with the species of Delta (including Phi of authors) as
members of the genus Eumenes (e.g., Matsumura, 1911; Tosawa, 1934; Iwata, 1939a). Here,
I follow Giordani Soika (1972) who regarded some of Saussure's (1855) divisions as
genera. Diagnosis for Delta given by Saussure and Dalla Torre (1904) is reproduced below
with slight modifications.
Clypeus much higher than wide. Mandible very long, somewhat curved; teeth absent
or inconspicuous. Galea 1.5 times as long as the basal part of maxilla; segments of
maxillary palp not gradually becoming short (segment 2»1; 3=4+5+6); segment 1
154
widened in the middle; segment 2 curved and apically widened. Labium with a straight,
very long, deeply bifid tongue; paraglossa "fadlich"; segment 2 of labial palp longer than
segment 1. Eye large, swollen, narrowly notched. Terminal segments of male antenna very
long, recurved to lie on some preceding segments. Thorax [=alitrunk) short. gaster pearshaped, depressed. Gastral petiole slender, a little longer than the thorax, elongate pearshaped, posteriorly widened, often in the middle slightly swollen or almost linear, often
with a pair of dentides at about middle length, with venter invisible, and slightly,
impressed. Segment 2 bell-shaped, posteriorly narrowed, widest at 1/3 length from the
base or in the middle, never compressed laterally, but slightly depressed dorso-ventrally.
Phi Saussure has been treated by some authors as a distinct genus (e.g., GiordaniSoika, 1972), while Vecht (1981) and Carpenter (1986) synonymized it with Delta.
Saussure's diagnosis for his I~ Division (= Phi) is as follows: "Labium and maxillae as in
lIe Division [now Delta); mandibles very long; [gastral) petiole linear, very long (1.5 times
as long as thorax [alitrunkl, or more), without obvious dentides; abdomen [gaster)
compressed, higher than wide; segment 2 bell-shaped, basally narrowed so as to form a
short stalk toward the petiole, and slightly narrower posteriorly than at the middle."
According to Giordani Soika (1972) Phi is distinguished from Delta by the following
points: tergite 1 much longer (Fig. 349), subcylindrical and more or less strongly curved;
male terminal sternite without longitudinal furrow which is seen in all the members of
Delta (s. str.). In spite of this fact, in this paper I will tentatively combine these two groups
into the genus Delta. Furthermore, since the name Phi is preoccupied by a subgenus of a
New World polistine genus (Mischocyttarus), we cannot use this name in Eumenidae (cf.
Carpenter & Day, 1988).
Delta has been treated as neuter (Vecht & Fischer, 1972; Gusenleitner, 1987) or as
masculine (Giordani Soika, 1972; Vecht, 1981, pers. comm.). Here I follow the most recent
work by Giordani Soika (1986) where it is treated as neuter.
This genus is widely distributed in southern Palearctic, Oriental, Australian and
Ethiopian regions.
Delta esuriens okinawae Giordani Soika
(Figs. 337-340,342,355)
Eumenes campaniformis var. gracilis Saussure: Sonan, 1938, Trans. Nat. Hist. Soc. Formosa, 28: 7So79.
Eumenes esuriens Fabricius: Matsumura, 1911, Thous. Ins. Jpn. Supp!. 3: 109, pI. 40, fig. 10; Matsumura
and Uchida, 1926, Ins. Matsum. 1: 35; Matsumura, 1930, III. Thous. Ins. Jpn. 2: 11, pI. 2, fig. 10; 1931, 6000 1lI.
Ins. Jpn.-Emp. p. 15 (no. 74); Tosawa, 1934, Trans. Shikoku Entomol. Soc. 5: 13-14, fig. 9.
Delta campaniforme dcinawae Giordani Soika, 1986, Boll. Mus. Civ. Stor. Nat. Venez. 35 (1984): 76-77 (.!f
J')(type loc.: Ie-jima, Okinawa Is.).
Delta esuriens (Fabricius): Azuma and Kinjo, 1987, Check-list Ins. Okinawa, p. 315.
Japanese name: Kurosuji-suzubachi.
Diagnosis. Female and male. Body length (h+th+t1+2): 17.5-19.0 mm in if, 13.5-16.5
mm in J'. Fore wing length: 14.5-15.0 mm in if, 11.0-13.5 mm in J'. Structurally as in the
nominotypical subspecies. Antennal hook (J') large, received by a concavity of segment
10 (Fig. 339).
Head black, with the following parts yellow (Fig. 337): cJypeus, a large interanten-nal
marking, a wide marking on inner orbit which fills up the lower 2/3 of ocular sinus, space
155
337
342
349
"
j
/
i
\.
"'.
....,{,.,
(~ ..J
t .. '?
Figs. 337-342. Delta esuriens okinawae. 337, head in frontal view (~); 338, male c1ypeus; 339,
terminal segments of male antenna; 340, body color pattern (ssp. okinawae); 341, ditto (ssp.
esuriens); 342, gastral segment 2 in profile.
Figs. 343-349. Delta flavopictum form05anum. 343, head in frontal view (~ ); 344, 345, color pattern of
female c\ypeus; 346, male c\ypeus; 347, terminal segments of male antenna; 348, gastral
segment 2 in profile; 349, tergite 1 (petiole) from above.
156
between c1ypeus and antennal socket (this yellow part often connected with the
interantennal marking), genal band (more developed in !f.). Mandible brownish. Antenna
ferruginous, darker in apical segments; scape below yellowish, above blackish or
ferruginous. Thorax black, with the following parts yellow: pronotum wholly, a large
marking on mesopleuron, apical 1/3 of tegula, parategula, metanotum wholly.
Mesoscutum often marked with reddish brown basally; scutellum wholly and upper 2/3
of tegula ferruginous (tegula sometimes with a black spot at base); mesopleuron and
metapleuron also sometimes with ferruginous markings. Propodeum brown or
ferruginous, posteriorly with a large black basal marking in J'; posterior face above and
below with yellow markings. Legs brown to ferruginous, with yellow markings on fore
femur and tibiae of all legs; femora and tibiae extensively blackish and tarsi almost wholly
blackish in J'. Gastral petiole ferruginous; preapical band (incised medially) yellow; a
wide band just before the yellow band and basal part of petiole black. Gastral tergite 2
yellow in apical 1/2; basal 2/5 ferruginous; space between the yellow and ferruginous
area black, forming a distinct band; ferruginous area often with an inconspicuous black
marking at base. Subsequent tergites yellow, but when the segments are extended the
basal black parts are visible. Gastral sternites 1 and 2 extensively ferruginous; sternite 2
with a yellow apical band; subsequent sternites yellow with basal part ferruginous.
In the nominotypical subspecies from India (Fig. 341), yellow markings are more
abundant especially on head, and alitrunk is extensively ferruginous. One female
specimen from Thailand was intermediate between the nominotypical form and okinawae.
Materials examined. C. Ryukyus: Okinoerabu-jimll - 1 !?-, 17 vii 1984 (M. Maegata); Yoron-tO - 2 J' J', 4 vi
1985 (SKY); Okinawa-jimll -1 J', Naha, 29 ix 1977 (SY), 2!?- !?-, Kudeken, 30 ix 1977 (SY), 1 J', Nakagusuku, 3 x
1977 (SY), 1 J', Ogimi-son, 26 viii 1979 (H. Nagase), 1!?-, T6baru, Kunigami, 3 vii 1982 (yH), 2 J' J', Hentona,
27 vii 1987 (SKY), 1 J', Nago, 3 x 1987 (A. Nagatomi), 2!?- !?-, Ginoza-son, 4 x 1987 (AN); Kouri-jimll - 2!?- !?-, 18
x 1988 (Y. Kusui); Yabuchi-jimll - 1 J', 22 x 1988 (Y. Kusui); Hamahiga-jimll - 4J' J'1!?-, 22 x 1988 (Y. Kusui);
Tokashiki-jimll - 4 J' J'1!?-, 11 x 1988 (SKY).
S. Ryukyus: Miyako-jima - 5J' J', Gusukube, 17 vii 1987 (SKY), 1 J', Hirara, 18 vii 1987 (SKY); Taramajimll -1 J', 19 vii 1987 (SKY); Ishigaki-jima - 1!?-, Ishigaki-shi, 10 x 1977 (SY), 1 J', Kabira, 22 vii 1987 (SKY), 2J'
3', Shiraho, 26 vii 1987 (SKY); Taketomi-jimll - 3 J' J'2!?- !?-, 24-25 vii 1987 (SKY); Kohamll-jimll -13',25 vii 1987
(SKY); lriomote-jimll - 9J' J'4!?- !?-, Funaura, 5-9 x 1977 (SY), 1!?-, Gtomi, 25 vii 1985 (AN), 1!?-, Toyohara, 11 x
1987 (AN); Haterumll-jimll -1!?-, 1 vii 1988 (SKY); Yonaguni-jimll - 1 J', Sonai, 5 vii 1988 (SKY).
Distribution. Amami Is. (Okinoerabu-jima; Yoron-to); Okinawa Is. (Okinawa-jima; lejima; Kouri-jima; Yagaji-jima; Sezoko-jima; Yabuchi-jima; Hamahiga-jima; Tokashiki-jima;
Kume-jima); Miyako Is. (Miyako-jima); Tarama Is. (Tarama-jima); Yayeyama Is. (Ishigakijima; Taketomi-jima; Kohama-jima; Iriomote-jima; Hateruma-jima; Yonaguni-jima).
Taiwan. The nominotypical form ranges from Senegal in the west through Persia to Malay
Archipelago in the east (Bequaert, 1918).
Taxonomic notes. The form esuriens, originally described from India, is structurally
very similar to D. campaniforme (Fabricius), and has often been treated as a subspecies of
the latter (Gusenleitner, 1987) or of "Eumenes" caffer (Linne) (Bequaert, 1918). Esuriens and
campaniforme are, however, distict species (Vecht, pers. comm., 1981): esuriens has fine
short bristles on inner surface of the terminal segment of male antenna (Fig. 339), while
campaniforme lacks them. In coloration constant differences exist as follows: in
campaniforme scutellum and sides of propodeum yellow, and gastral tergite 2 basally with
two large yellow spots. I have found no intermediate specimen even in the series
containing both the forms from the same or adjacent localities.
Biology. This species builds its pot nests on the depressed part of rock surface. The
157
pot is hemispherical, and its bottom is widely attached to the substratum. An excel-lent
photo was given by Takara and Azuma (1973; Eumenes esuriens). Lepidopterous larvae are
hunted for the young. Males wait females around flowers, and the copulation occurs there
(Yamamuro, 1988).
Delta fUzvopidum formosanum (Zimmermann)
(Figs. 343-349,355)
Eume7les arcuata formosana Zimmermann, 1931, Z. Morph. Oekol. liere, 22: 206, fig. 26 (no. 5) (~ J')
(type loc.: Pilan, Taiwan).
Eumenes arcuata Fabricius: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 109-110, pI. 39, fig. 11;
Matsumura and Uchida, 1926, Ins. Matsum. 1: 35-36; Matsumura, 1930, 1lI. Thous. Ins. Jpn. 2: 12, pI. 2, fig.
11); 1931, 6000 TIL Ins. Jpn.-Emp. p. 75-76; Tosawa, 1934. Trans. Kansai Entomol. Soc. 5: 15, fig. 4.
Eumenes f/aTJopictus formosanus Zimmermann: Vecht, 1959, Zool. Verh. 41: 39.
Phi arcuata (!) Fabricius: Azuma and Kinjo, 1987, Check-list Ins. Okinawa, p. 315.
Japanese name: Haranaga-suzubachi.
Diagnosis. Body length (hHh+t1+2): 21.5-25.0 mm in ~, 17.5-20.5 mm in d'. Fore
wing length 18.5-21.0 mm in ~, 14.5-18.0 mm in d'. Structure as in the nominotypical
form for which a good description was given by Vecht (1959). In profile, gastral tergite 2
more produced below than in D. esuriens (Fig. 348 vs. 342). Antennal hook (d') strongly
curved, without hairs on inner face (Fig. 347).
In color pattern the Japanese form is very similar to the Taiwanese one, but ocular
sinus is more extensively yellow (Fig. 343), clypeal black marking in the female is
generally larger, and fore femur constantly has a yellow line on its outer face. Although
color pattern is generally stable in Japan, the black marking on the female clypeus may
vary among individuals (Figs. 343-345).
Material examined. S. Ryukyus: Ishigaki-jima - 1 ~, vii 1922 (S. Hirayama), 1 ~, Banna-dake, 12 vii 1973
(H. Takizawa), 1~, Kabira, 22 vii 1987 (SKY), 7 J' J'10~ ~,Banna-ilake, 3-8 vii 1988 (K Nakamine); Iriomotejima - 2~ ~, Komi, 24-26 x 1973 (M. Owada), 6J' J'2~ ~, Funaura, 5-9 x 1977 (SY), 1 ~, Ohara, 23 v 1981
(AN), 1 ~, Komi, 17 v 1981 (AN), 1 ~, Urauchi, 22 x 1981 (T. Moriyama), 2 J' J', Ohara, 6 i 1982 (AN), 1.!f.,
Komi,31 vii 1983 (AN), 1 ~, Ohara, 29 vii 1983 (AN), 1.!f., Uehara, 21 xi 1983 (T. Moriyama), 1 ~, Toyohara,
11 x 1987 (AN), 1 J'2~ ~, Mihara, 4 xii 1988 (SKY); Yubu-jima -1 ~,4 xii 1988 (SKY).
Distribution. Okinawa Is. (Okinawa-jima ?); Miyako Is. (Miyako-jima); Yaeyama Is.
(lshigaki-jima; Iriomote-jima; Yubu-jima). Taiwan. I have not seen any specimen from
Okinawa-jima and Miyako-jima.
Biology. The nesting behavior of this form was intensively studied by Iwata (1939a) in
Taiwan. Nests are constructed on the flat surface of stone gates or on tree twigs. The nest
consists of several quatered-spherical mud pots that are finally covered by mud (secondary coating) into a single cluster. Iwata et al. (1982) present a series of beautiful photos of
a nest, though unfortunately the locality is not stated. Sonan (1927) observed a female
wasp hunting the larva of a tortricid, Homona menciana, in Taiwan, while Iwata (1939a)
stated that most of the prey insects stored in brood cells were larvae of Geometridae and
Noctuidae.
158
Genus Pseumenes Giordani Soika
Pseumenes Giordani Soika, 1935, Ann. Mus. Civ. Stor. Nat. Genova, 57: 145 (as subgenus of Pareumenes
Saussure)(type species: Eumenes eximius Smith, 1861, original designation).
Japanese name: Kagimon-tokkuribachi Zoku.
Pseumenes and its allied Old World groups had been placed together under
Montezumia and as divisions or subgenera within the latter until Vecht (1963) raised these
to genera and created another genus, Coeleumenes. Currently the name Montezumia is used
only for New World species.
Head elongate in frontal view, higher than wide in .!f. Mandible relatively short (Fig.
350). Thorax depressed, wider than high, oval as seen from above. Mesepisternum
without epicnemial carina. Disk of metanotum very slightly convex, well defined laterally.
Propodeum gradually sloping from base to apex; the dorsal (posterior) portion sharply
demarcated from each lateral portion by a carina which is produced into a distinct process
or teeth at the apex of propodeum; dorsal portion basally with a longitudinal slit, from
which runs a median carina to the apex. Gastral segment 1 distinctly petiolate (Fig. 353).
Gastral stemite 1 with long and narrow anterior part which is weakly striate and fused
with the tergite; posterior part short, triangular, not striate (Fig. 354). Outer face of hind
tibia not spinose.
Pseumenes depressus depressus (Sa us sure)
(Figs. 350-355, 361)
Eumenes depressus Saussure, 1855, Et. Fam. Vesp. 3: 135 (!f J')(type loc.: les Indes orientale); Dalla
Torre, 1894, Cat. Hym. 9: 22; Tosawa, 1934, Trans. Kansai Entomol. Soc. 5: 6 (depressa).
Pareumenes depressa: Dalla Torre, 1904, Gen. Ins. 19: 19; Vecht, 1937, Treubia, 16: 273, fig. 3b (depressus: in
subgenus Pseumenes); Tosawa, 1936, Kansai Koncho. Zasshi, 4: 46; Sonan, 1938, Trans. Nat. Hist. Soc.
Formosa, 28: 77-78.
Pareumenes quadrispinosus Saussure: Liu, 1941, Notes Entomol. Chin. 8(6): 256 (in key), 280
[misidentificationl.
Pseumenes depressus depressus: Vecht, 1963, Zool. Vern. 60: 25-26.
Pseumenes depresus (!) (Saussure): Azuma and Kinjo, 1987, Check-list Ins. Okinawa, p. 315.
Japanese name: Kagimon-tokkuribachi.
Diagnosis. Female. Body length (h+th+t1+2): 16.0-19.0 mm. Fore wing length: 16.018.0 mm. Head with vertex fairly swollen (Fig. 350); upper portion of frons and anterior
portion of vertex densely punctate; the punctures not well defined. Ocellar triangle flat;
distance between posterior ocelli much longer than that between posterior and anterior
ocellus. Clypeus as wide as high, weakly punctate, apically narrowly emarginate. Labial
palp 4-segmented; segment 1 longest and apically widened. Pronotum and mesoscutum
except for posterior 1/5 punctate; other parts of thorax impunctate or only very weakly
punctate, and somewhat shining. Propodeum laterally weakly striate, and dorsally
weakly punctate and shining; propodeal groove with obliquely running striae. Each
lateral half of pronotum emarginate at apex (Fig. 353). Gastral segment 1 with parallel
sides in anterior half, and gradually widened toward apex, with a preapicallongitudinal
furrow. Tergite 2 approximately as long as wide, densely but weakly punctate laterally; on
159
tergites 3-6 punctation quite inconspicuous. Slender anterior part of sternite 1 very weakly
striate; punctation on sternites 2-5 very weak; terminal sternite almost impunctate.
Parastigma of fore wing more than half as long as stigma.
Black, with the following parts yellow: c1ypeus wholly (sometimes apically black), a
marking along inner side of eye extending onto the whole ocular sinus, a long bar on frons
starting at just below anterior ocellus and reaching c1ypeus, complete genal band,
antennal scape below (flagellum below ferruginous), pronotum wholly, a large spot on
mesopleuron above, a pair of fishhook-shaped marks on mesoscutum, tegula with a
brownish central spot, parategula, a pair of spots on scutellum, a pair of large markings on
dorsal face of propodeum each with a median black spot which is often connected to the
lateral black area, a pair of small spots on gastral petiole at 2/5 length from apex, a narrow
apical band on tergite 1 which is medially and laterally interrupted and extending from
each lateral end toward base for some distance, a pair of large basal spots and a wide
apical band on tergite 2 (the latter medially incised), apical bands on tergites 3-5 (apical
bands 2-5 each containing a brownish, small median spot), an irregular marking or
markings on sternite 2, fore femur below, outer face of tibiae of all legs.
Male. Body length (h+th+t1+2): 12.0-14.0 mm. Fore wing length: 11.0-11.5 mm. Very
similar to the female in structure and coloration. Body more slender. Head nearly circular.
Figs. 350-354. Pseumenes depress us. 350, head in frontal view (-'?-); 351, male c1ypeus; 352, terminal
segments of male antenna; 353, body color pattern; 354, gastral sternite 1.
160
• 0
!
-5f
. -, (?
-f}jJ
:.. IJ
D. esurie ns
"------"----355
Fig. 355. Distribution of Pseudozumia indosinensis, Pseumenes depressus, "Pachymenes" yayeyamensis,
Delta esuriense and D. flavapicutum in Japan.
Clypeus higher than wide (Fig. 351). Two apical segments of antenna very small, recurved
to reach barely the base of segment 11 (Fig. 352). Mandible with a basal yellow mark. Legs
more extensively marked with yellow.
Material examined. S. Ryukyus: Ishigaki-jima - 13', 11 vii 1973 (H. Takizawa), 13', 16 viii 1983 (S.F.
Sakagami & YM), 13',22 vii 1987 (SKY), 23'6', Banna-dake, 4 vii 1988 (SKY), 23' 3', same loc., B vii 19BB (K
Nakamine), 13', Omoto-dake, B vii 1988 (K Nakamine); Kohama-jima -1~, 25 vii 1987 (SKY); lriomote-jima -1
~, Ohara, 15 v 1981 (AN), 13', Otomi, 29 vii 1983 (AN), 13'1 ~, Komi, 29-31 vii 1983 (AN); Hateruma-jima - 2
3' 3', 2 vii 1988 (SKY).
Distribution. Miyako Is. (Miyako-jima); Yaeyama Is. (lshigaki-jima; Kohama-jima;
Iriomote-jima; Hateruma-jima). Taiwan; Thailand; Indo-China; Malaya; S. China; India
(West Bengal).
Biology. There is little information available about the bionomics of this species in
Japan. According to the classical work by Piel (1935, summarized in Liu 1941; in both,
referred to as Pareumenes quadrispinosus) in southern China, this species nests in long
internodes of dry bamboo and makes a small funnel-shaped hole on the side of bamboo
near the upper node; provisioning and oViposition are done through this hole. This
species is ethologically unique in that hunting caterpillars precedes oviposition (in all the
other groups of Eumenidae the order is reversed) and that eggs are directly glued to the
inner surface of the cell wall (without a suspensory thread). Iwata (1976, 1980b) reviewed
the biology of this species adding his own observations on Hainan Island and in Thailand,
but did not confirm Piel's observation regarding the hunting-oviposition sequence.
Yamamuro (1985) observed male behavior in mid-July on Iriomote-jima, the S.
161
Ryukyus. Males flew between 9:30-15:00 (peak: 11:00-13:00) around flowers of Vitex
rotundifolia etc., patrolling a certain area including flowers along a regular route. Although
female wasps visited flowers in the male-patrolled area, no copulation was observed.
Genus Pseudozumia Saussure
Pseudozumia Saussure, 1875, Smiths. Misc. Coli. No. 254(1), p. 128 (division of Monlezumia Saussure)
(type species: Monlezumia indica Saussure, 1855); Bequaert, 1921, Rev. Zool. Afr. 9: 235-251 (as subgenus of
Monlezumia); Giordani Soika, 1941, Boll. Soc. Venez. Stor. Nat. 2: 33 (as genus); Vecht, 1963, Zool. Verh. 60: 16
(in key), 41-42 (as genus).
Japanese name: Koshibuto-suzubachi Zoku.
This genus is similar to the genus Pseumenes, but differs therefrom in the following
points: head and thorax rather strongly punctate; mesoscutum with prescutal grooves (fig.
359); mesopleuron with epicnemial carina; gastral segment 1 much shorter (less than twice
longer than wide at apex of the tergite); sternite 1 irregularly rugose.
Bequaert's (1921) Pseudozumia obviously included species of Pareumenes, Coeleumenes,
Nortozumia and Pseudozumia (sensu Vecht), and Giordani Soika's (1941) concept included
some species of Coeleumenes. This genus is mainly Oriental in distribution, being known
from Celebes, Java, Sumatra, Borneo, Malaya, Sikkim, southern China, the Philippines,
Taiwan, and the Ryukyus.
Pseudozumia indosinensis Giordani Soika
(Figs. 355, 356-360,361)
Pseudozumia indosinensis Giordani Soika, 1960, Boll. Mus. Civ. Venez. 11: 93-94 (!f. J')(type loc.: Runji A.
Tal, Sikkim).
Monlezumia (Pseudozumia) indica Saussure: Sonan, 1937, Trans. Nat. Hist. Soc. Formosa, 27: 14-15;
Sonan, 1938, Arb. Morph. Taxon. Entomol. 5: 69.
Monleozumia (!) indica: Azuma and Kinjo, 1987, Check-list Ins. Okinawa, p. 315 (Okinawa Is.)
Monolezumia (!) indica: Matsumura, 1911, Thous. Ins. Jpn. Suppl. 3: 113, pI. 39, fig. 18; 1930, III. Thous.
Ins. Jpn. 2: 15-16, pI. 2, fig. 18; 1931,6000 III. Ins. Jpn.-Emp. p. 17.
Japanese name: Koshibuto-suzubachi.
Diagnosis. Female. Body length (h+th+t1+2): 16.5-19.0 mm. Fore wing length: 16.019.5 mm. Head subcircular (Fig. 356), densely punctate especially on frons and vertex.
Clypeus slightly higher than wide, relatively widely truncate at apex, almost flat in apical
half; the flat part with a few punctures and irregularly striate; punctation in basal half fine
and dense. Supra-antennal area with very minute punctures. Interantennal keel sharp.
Depression for cephlic foveae well defined, with a median keel. Occipital carina
developed only laterally. Anterior vertical face of pronotum with a few minute punctures;
posterior horizontal portion strongly and densely punctate; posterior pronotallobe with a
few punctures. Mesoscutum strongly and densely punctate; the punctation sparser near
apex and in posterior portion; spaces between punctures and bottoms of large punctures
micropunctate. Prescutal grooves deep, with transverse carinae. Scutellum finely and
sparsely punctate; spaces between punctures micropunctate. Furrow between meso162
scutum and scutellum deep in the middle. Tegula sparsely punctate at base; remainder
rnicropunctate. Epicnernium almost irnpunctate except for lower posterior portion;
mesepisternum and mesepimeron strongly and densely punctate and partly reticulate.
Punctures on metanotum large, but sparse. Metapleuron with a few punctures.
Propodeum without shelf, posteriorly not concave, with a deep median groove; a carina
starting from the lower end of the groove; posterior face of propodeum with dense
punctures of variable size; each side of the posterior face deeply emarginate at apex (Fig.
359); lateral part with large punctures above and superficially striate below. Gastral
segment 1 subpetiolate, shorter than segment 2. Tergite 1 moderately punctate, carinate in
the middle; tergite 2 more finely punctate especially in apical 1/3; tergites 3 and 4 with
small macropunctures; tergites 5 and 6 micropunctate; tergites 2 and 3 apically with a
distinct lamellate area. Sternite 1 irregularly rugose; other sternites more finely punctate
than corresponding tergites.
Body almost wholly black. Yellow are: a median marking on cIypeus, a transverse
frontal mark, antennal scape below, a small basal spot on mandible, a pair of very small
spots on the anterior vertical face of pronotum above, apex of each side of dorsal face of
propodeum. Antennal flagellum below rufous. Wings blackish.
Male. Body length (h+th+t1 +2): 11.0-11.5 mm. Fore wing length: 11.5-12.0 mm. Similar
360
Figs. 356-360. Pseudozumia indosinensis. 356, head in frontal view (~); 357, male c1ypeus; 358,
terminal segments of male antenna; 359, body from above; 360, gastral tergites 2-4 in profile
( d').
163
to the female in structure and coloration. Clypeus more elongate and distinctly emarginate
apically (Fig. 357). Antennal hook not large, just reaching the apex of segment 10 (Fig.
358). In one specimen, propodeum posteriorly with a small medial concavity. Gastral
tergites 2-4 with lamellate areas at apex (Fig. 360). Mandible without yellow spot.
Material examined. S. Ryukyus: Ishigaki-jima - 2 d' d'3Sf- Sf-, Banna-dake, 3-8 vii 1988 (K. Nakamine);
Iriomote-jima - 1 Sf- , Komi, 28 vii 1985 (AN).
Distribution. Okinawa Is. (Okinawa-jima after Azuma & Kinjo, 1987) (?); Yaeyama Is.
(lshigaki-jima; Iriomote-jima). Taiwan; continental China; Sikkim. I have not seen any
specimen from Okinawa-jima.
Taxonomic notes. This species is readily distinguished from the other congeners by the
well-developed apical lamellae on gastral tergites 2 and 3 in the female and 2-4 in the
male. According to the original description by Giordani Soika (1960), mid and hind
femora are rufous or ferruginous. In the Japanese specimens, however, all femora are
wholly blackish.
Biology. Nothing is known of nesting behavior.
VI.
DISTRIBUl10N PATIERN OF EUMENIDAE IN SOME ISLAND GROUPS IN JAPAN
Japan is an island country, comprising the four main islands Hokkaido, Honshu,
Shikoku, and Kyushu and some island groups with many islands, most of which are very
small. The Japanese mainlands repeatedly became part of the Asian continent during the
glacial ages so that their fauna and flora are expected to be essentially very similar to
those of the Korean Peninsula, northern China and eastern Siberia. The number of species
of a given taxon inhabiting Japanese mainlands should be in general smaller than that
inhabiting northeastern part of continental Asia, mainly owing to a much smaller area (d.
MacArthur & Wilson, 1967). In fact Japan lacks such genera as Pseudepipona and Antepipona that are widely distributed in eastern Asia. However, in some taxonomic groups, we
have species endemic to Japan. They may have evolved during interglacial periods when
the Japanese mainlands were isolated from the continent by the Japan Sea, and have
already completed speciation.
At present, out of the 54 Japanese eumenid species ten are known only from the
Japanese Archipelago (mainlands and nearby islands), six are endemic to the Ryukyus or
Ogasawara Islands, and one is endemic to the Tsushima Islands. The others are found also
in Korea, Taiwan, continental China, eastern Siberia or Europe, though the Japanese
populations are often differentiated at subspecies level. Most (7 spp.) of the species known
only from the Japanese Archipelago belong to the genus Symmorphus, which is one of the
most poorly studied genera in the Far East. It is possible that some of the Japanese species
also occur on the continent. Furthermore, another species (Euodynerus bicingulatus
Giordani Soika) is a doubtful taxon as discussed before. The remaining two (Stenodynerus
tokyanus and Ancistrocerus japonicus) are apparently peculiar to Japan, but should be
subjected to further faunistic surveys on the continent.
The area-species relation for the mainlands, Sado-ga-shima, Awaji-shima and Yakushima is shown in Table 3. The largest species number (36) is found on Honshu, which has
the largest area and covers the longest distance from north to south. On the second largest
island Hokkaido, however, only 22 species are known to occur. This may be partly due to
the insufficient survey, but mainly caused by the fact that most of the warm-temperate
164
Table 3. Number of eumenid species on the Japanese mainlands and some relatively large islands."
Genera
Hokkaid6
(78073)
Discoelius
Stenodynerus
Allodynerus
Euodynerus
Rhynchium
Anterhynchium
Pararrhynchium
Oranostrocerus
Ancis trocerus
Symmorphus
Oreumenes
Eumenes
Total
1
3
1
2
Honshii
(227414)
Sado I.
(857)
4
1
Shikoku
Kyiishii
Yakul.
(593)
(18256)
(36554)
(501)
1
1
1
3
4
3
2
2
2
3
1
1
1
2
1
2
4
1
1
Awaji I.
2
1
1
1
3
1
1
1
2
9
1
4
1
2
5
2
1
4
4
22
36
10
6
23
23
6
5
1
1
6
1
1
1
1
2
3
1
2
12
" Island areas are given in parentheses in km 2.
species do not extend northward beyond the Tsugaru Strait. The species number (23) for
Shikoku is the same as that for Kyushu that is approximately twice as large as Shikoku in
area. The main reason for this might be ascribed to relative collection efforts on these two
islands. Mountainous regions of Shikoku were intensively surveyed by Mr. Y. Sugihara in
the early 1900s, whereas no such effort has been made on Kyilshu. Furthermore, there is
no subtropical eumenids that have their northern limits of distribution in Kyushu; in other
hymenopteran groups subtropical forms often have their northern limits in southern
Kyushu and contribute to enriching the fauna of Kyilshu (e.g., Odontomachus monticola in
Formicidae and Scolia kuroiwae in Scoliidae).
In short, the eumenid fauna of Japan is characterized by many Palearctic (especially
eastern Palearctic) elements, some Oriental species and some endemic ones in the
Ogasawara or Ryukyu Islands. Eventually only a few will prove to be endemic to the
Japanese Archipelago. The genus Ancistrocerus is notable in that most species found in
Japan have quite wide distribution ranges and have not undergone subspeciation in
Japan. A. antilope is the only species with a Holaretic distribution among the Japanese
Eumenidae.
In the following lines I will discuss the distribution pattern of eumenids in some
island groups that have been relatively well studied.
1. Sado-ga-shima (Niigata-ken)
The eumenid fauna of this relatively large island quite resembles that of the opposite
part of the mainland of Honshu (Niigata-ken-hondo), but much poorer than the latter in
species number. Only eight species (Stenodynerus sp., Anterhynchium flavomarginatum,
Orancistrocerus drewseni, Symmorphus foveolatus, S. decens, S. apiciornatus, Oreumenes
decoratus and Eumenes micado) had been known by 1982 (Baba, 1935; Yasumatsu, 1935d;
Yamane, 1982b; Onuma, 1989a), and the present study has added two (Ancistrocerus
japonicus and Eumenes rubronotatus). Thus, only ten (43.5 %) of the 23 species inhabiting
165
the whole Niigata-ken are known from Sado-ga-shima. The percentage is even small in
comparison with social wasps (Vespidae) (13 out of 22 spp., 59.1 %) (note that Sado-gashima is the only non-mainland island in Japan where Vespa crabro occurs). Though
further collection efforts will add some species, the known faunal composition and its
characteristics in comparison with Niigata-ken-hondo can be explained by the geological
history of the island and the current biogeographical theory. This island may have become
close to (but not connected with) the mainland during the last glaciation (Wiirm).
Imadate and Kaneda (1984) stated that out of the 17 Protura species found in the
whole Niigata-ken, 13 (76.5 %) inhabit also Sado-ga-shima. Although the island has
neither endemic species nor subspecies, the Protura fauna is rather rich in species number.
Further, in different localities, different sets of Protura species tend to occur. This means
that Sado-ga-shima is large enough to harbor these small insects, by partitioning the
island into numerous habitats. Higuma (1987) gave other examples. Forty eight (48.5 %)
out of the 99 Odonata species inhabiting Niigata-ken-hondo have been recorded from
Sado-ga-shima. For resident butterflies a similar figure (71 out of 125: 56.8 %) is given. The
cicada fauna of this island is much richer: 10 (83.3 %) out of the 12 species inhabiting
Niigata-ken-hondo have been recorded. In these three groups there are virtually no
species that occur on Sado-ga-shima but not on Niigata-ken-hondo. Thus, the insect fauna
of Sado-ga-shima is essentially the same as that of Niigata-ken-hondo, but more or less
poorer than the latter.
2. Izu Islands <Tokyo-to)
This island group consists of some ten small islands lying just south of the Kanto
Table 4. Vespoid wasps recorded from the Izu Islands.
Island
Area
(km 2 )
Distance
from Honshu
References
Polistes hebracus 1)
P. h. nigruotum2 )
Kamiya, 1931
29
Polistes fadwiga~)
Rhynchium japonicum 4)
Kamiya, 1934
35
Seguro-ashinagabachi5)
Umeya, 1956a
90.99
22km
To-shima
4.19
Nii-jima
23.42
Izu-Oshima
Species recorded
Polistes yolwhama/»
Vespa trapica pulchra
Shikine-jima
40
No species recorded
Kozu-jima
18.58
3.82
47
Seguro-ashinagabachi5)
Ki-ashinagabachi6)
Miyake-jima
55.14
75
No species recorded
No species recorded
Mikura-jima
19.69
100
Hachijo-jima
69.17
178
Stenodynerus tolcyanus
Umeya, 1956b
S. frauenfeldi
Yamane & Gusenleitner,
1982
Hachijo-kojima
3.22
178
Odynerus frauenfeldP)
Sawada & Watanabe, 1959
Aoga-shima
5.23
246
Stenodynerus frauenfeldi
Present study
1) =?Polistes rothneyi; 2) =?P. jadwigae; 3) =P. jadwigae; 4) =Anterhynchium flavomarginatum;
5) =Polistes jadwigae; 6) =P. rothneyi; 7) =Stenodynerus frauenfeldi.
166
District (Honshu) between 30 0 24'N and 34°4S'N. Regarding the Vespoidea, up to now
only three eumenid, two polistine and one vespine species have been recorded from these
islands (Table 4). No relict or endemic species has been known in these groups. The
population of Stenodynerus tokyanus is differentiated into a distinct geographical race
endemic to Hachij6-jima. It is not clear whether all of the other species recorded are native
or not, because hibernating queens (social wasps) and prepupae (eumenids) can be
transported via human activities over a long distance. However, the fact that the queen of
Vespa tropica generally overwinters in soil or decaying stumps (Matsuura, 1966) strongly
suggests that there has been few chances for this species to be introduced by man. If so,
the sole food of its young, the polistines, must have been also native. On the other hand, it
is often thought that two eumenid species, Anterhynchium flavomarginatum and
Stenodynerus frauenfeldi, are apt to be transported with bamboo or reed tubes in which they
nest. Data pertaining to this possibility are, however, at present not available.
Kurosawa (1978) mentioned that the insect fauna of the Izu Islands is quite poor and
principally the same as that of Japan proper in species composition, with a few endemic
species such as Lucanus gamunus (Coleoptera, Lucanidae). This is also true of the vespoid
fauna; only approximately 10 % (with no endemic) of the species known from lowland
Honshu occur on the islands (present study). Kurosawa ascribed such features to the
relative closeness of this archipelago to the Honshu mainland and the occurrence of a
peninsula called Paleo-Izu Peninsula during the Pleistocene. Ikeda (1984), on geological
basis, criticized the latter aspect of Kurosawa's argument, stating that the Paleo-Izu
Penninsula if any occurred 3 million years ago, and that during the Quaternary many
islands in this region repeatedly appeared from and sank into the sea by volcanic
activities. Sunose (1981) also doubted the Pleistocene connections of the Izu group with
Honshu mainly based upon the distribution pattern of a gall-midge, Masakimyia pustulae.
Aside from the controversy, considering the sizes of the extant islands they must possess
the nature of oceanic islands. In this respect, the fact that the only eumenids inhabiting
there are species of Anterhynchium and Stenodynerus is of special interest, because they are
often principal components of eumenid faunas on small islands in temperate Japan.
3. Tsushima Islands (Nagasaki-ken)
Two main islands of Tsushima (Kami-agata and Shimo-agata) have been relatively
intensively surveyed for vespoid fauna. The most interesting aspect in biogeography of
this island group is the mixed occurrence of Korean (continental) and Japanese elements.
Also interesting is the occurrence of endemic species and subspecies.
In the present study has been recognized a total of 13 eumenid species from this
island group (Stenodynerus frauenfeldi, S. chinensis, S. clypeopictus, Euodynerus nipanicus,
Anterhynchium flavomarginatum, A. melanopterum, Pararrhynchium ornatum, Orancistrocerus
drewseni, Symmorphus tsushimanus, Eumenes punctatus, E. micado and Oreumenes decoratus).
This figure corresponds to ca. 62 % of the total number of species inhabiting the Kyflshu
mainland. Of these only one species (Symmorphus tsushimanus) is at present considered to
be peculiar to Tsushima, all the others inhabiting also continental Asia, while the Japanese
mainlands lack one of them, Eumenes punctatus.
Among the Vespoidea, Vespa tropica esakii Sonan and Anterhynchium flavomarginatum
tsushimarum are no doubt endemic to Tsushima (Ishikawa, 1970). V. t. esakii is very
peculiar in having the last gastral tergite almost completely yellow. In almost all the other
subspecies of this species, the last two tergites are extensively black (a condition also seen
167
in V. affinis). Over the greater part of its distribution range, V. tropica seems to mimic V.
affinis that is widely sympatric with the former (Yamane, unpubl.). On the Japanese
mainlands V. tropica has not undergone a drastic transformation in color pattern despite of
the fact that V. affinis is lacking there (Matsuura & Yamane, 1984), while on Tsushima it
has lost the black maculation on the last tergite. This condition may be explained by
supposing a simple genetic drift in the small population, but actually it can be mimicking
V. mandarinia japonica and V. analis insularis, both also occurring on Tsushima. On the
other hand, Anterhynehium flavomarginatum tsushimarum is quite unique in possessing an
almost wholly black body, while there is known no candidate model of this wasp.
Ishikawa (1970) mentioned that the subspecies micado occurs in Korea as well as on the
Japanese mainlands, but actually the Korean population belongs to another subspecies, A.
f. koreanum (Yamane, 1981), which is much more melanic than micado. In this respect
tsushimarum seems to have originated from more probably a koreanum-type than mieadotype population (or individual).
Shirozu (1980) pointed out that the insect fauna of Tsushima is a mixture of
subtropical and boreal elements in addition to the basic species composition common to
warm-temperate Japan. This is not true of the vespoid fauna at least. No subtropical
species such as Delta spp., "Paehymenes" yayeyamensis and Okinawepipona kogimai occur, and
the north-temperate and boreal genus Ancistroeerus is completely lacking.
4. Ogasawara Islands <Tokyo-to)
Up to now the following five eumenid species have been recorded from this archipelago: Stenodynerus ogasawaraensis, S. frauenfeldi, Euodynerus nipanieus, Anterhynehium
flavomarginatum and Pararrhynehium oeeanicum. Most records came from the two main
islands, Chichi-jima and Haha-jima; other islands have been very insufficiently surveyed.
Of the five species two (40 %; S. ogasawaraensis and P. oceanicum) are endemic to the
archipelago, and are each differentiated into two geographical races there.The two species
are no doubt involved in Mullerian mimicry. The Ogasawara populations of the other
three species are very similar to those of Honshu in color pattern, though the specimens of
A. flavomarginatum from Chichi-jima and Haha-jima are, in some respects, of subspecies
procella-type (proeella inhabits the Mi-shima Is., the N. Ryukyus).
The two endemic species are very peculiar in structure and color pattern, apparently
having no close relatives in Japan proper and the South Pacific. They possess some
tendencies in common: mesosoma is extensively marked with yellow, gastral tergites 1
and 2 are wholly rufous except for apical yellow bands, and the punctation on mesosoma
and gaster is very weak. These conditions are quite exceptional in the genera Stenodynerus
and Pararrhynehium. The derivation of these species is at present unknown (but see under
S.ogasawaraensis).
The percentage endemism for the whole insects occurring in the Ogasawara group is
rather high: 185 (29.8 %) out of the 620 recorded species are endemic (Habu, 1986).
Endemism is especially outstanding in Diptera, Hymenoptera, Coleoptera, Hemiptera and
Odonata. Among bees, 9 out of the ten recorded species are peculiar to this island group
(Nakane, 1970; Ikudome, 1989). The remaining one, Apis mellifera, was no doubt
introduced by man for apiculture. The three eumenids that are not endemic might have
been introduced through human activities. Unlike the honeybee, these wasps are
generally not considered to be beneficial insects in Japan so that it is not likely that they
have been introduced intentionally. But their immatures are apt to be transported by ship
168
Rp.f.
Ropalidia fasciata
Del ta esuriens
"Pachymenes n yayeyamensis
Vespa affinis
D.e.
Pc.y.
V.a.
D.f.
Delta flavopictum
Ps.d.
Pseumenes depressus
Pz.i.
Pseudozumia indosinensis
. "0
Q
Rp/
D.e",
PC.Y._ ---__ ~
-
...... c?
!
-¥
(I
~ '"
\,
V
<J " , -
,
\
. ., (J6
361
Fig. 361. Northern limits of distributions of some Oriental species of Vespoidea.
together with reed and bamboo tubes in which their nests are constructed. A similar
reasoning may apply to Polistes jadwigae (referred to as P. fadwigae in Nakane, 1970), whose
overwintering new queens are easily transported by ship together with loaded goods.
Kurosawa (1976) mentions that approximately 30 % of the beetles recorded from
Ogasawara are endemic and that more than 30 % may have been accidentally introduced
by man. Among indigenous species, those living in the dead wood are dominant. He
argued that these beetles might have migrated on driftwood from variou; sources. Among
Aculeata, Xylocopa (Koptortosoma) ogasawarensis is a candidate with this dispersing
mechanism.
5. Ryilkyil Islands (Kagoshima-ken and Okinawa-ken)
5.1. Relative abundance of Palearctic and Oriental species. Twenty five species are known
from this archipelago situated in the warm-temperate and subtropical zones (Table 5). On
the biogeographical basis, they are sorted into four categories as follows:
A. Palearctic species. Species that are found on the Japanese mainlands but not in
Taiwan and other parts of Southeast Asia. They, therefore, have the southern limits of their
ranges somewhere in the Ryilkyil Archipelago in at least Japan. The following eight
species are included in this category: Discoelius japonicus, Stenodynerus frauenfeldi, S.
chinensis, S. tokyanus, Euodynerus nipanicus, Ancistrocerus japonicus, Eumenes rubronotatus
and E. micado. Of these, Euod. nipanicus has a close relative in Taiwan (Yamane &
Gusenleitner, unpubl.), and might eventually prove to be con specific with the latter.
B. Wide-ranging species. The species of this category generally occur almost
throughout the archipelago, and also in both Japan proper and Taiwan. This category
169
corresponds to Ikehara's (1966) "whole-area-type" applied to the termites of the Ryukyus.
Seven species are included: Euodynerus dantici, Euod. trilobus, Rhynchium quinquecinctum,
Anterhynchium flavomarginatum, Orancistrocerus drewseni, Pararrhynchium ornatum and
Oreumenes decoratus. Judging from the color pattern, the Ryukyil populations of some
species (0. drewseni, P. ornatum and Oe. decoratus) are considered to have been derived
from the forms of Japan proper. Solely females have been collected for o. drewseni and this
fact supports the above suggestions, since the Taiwanese form of this species adopts
sexual reproduction. On the other hand, the populations of Euod. dantici in the Central and
Southern Ryukyus are not distinguishable from the form of Taiwan in color pattern.
However, all these species are tentatively included in this category.
C. Endemic species. This category includes species that are found only within the
Ryukyil Archipelago. The following four species are known: Stenodynerus rufomaculatus, s.
kusigematii, Pararrhynchium tsunekii and P. ishigakiense. The two species of Pararrhynchium
are closely related to P. ornatum and may have originated from the latter. However, the
situation is somewhat complicated, because P. ornatum is found in both Japan proper and
Taiwan. At present the dispersal route or routes of the supposed ancestors of P. tsunekii
and P. ishigakiense into the Ryukyus cannot be determined. On the other hand, the two
Stenodynerus species (rufomaculatus and kusigematii) have the closest relatives <frauenfeldi
and tokyanus, respectively) in Japan proper, but no relatives in Taiwan.
D. Oriental species. This category, corresponding to Ikehara's (1966) "south-areatype", includes species that occur in Taiwan or widely in Southeast Asia and have the
northern limits of their distribution in the Ryukyus (Fig. 361). The following six species
may belong to this category: Okinawepipona kogimai, "Pachymenes" yayeyamensis, Delta
esuriens, D. flavopictum, Pseumenes depressus and Pseudozumia indosinensis. O. kogimai and
"P." yayeyamensis are at present known only from the Ryukyus and Taiwan; the others
have much wider ranges in Southeast Asia.
The relative abundance (no. species in percentage) of these four categories in the
Ryukyus is shown in Fig. 362. In the S. Ryukyus, the proportion of Oriental species is
;;;
"
«l
,~
'M
,-
.'"" ., :;;
,~rO
M
«l-
,~
«l
",5
~,~
H'M
0-
«l«l
;;;
;:;
N
N
e;
-
..
C «l
«l
.,-i.,..j
"5
,~
",'M
'M
«l
5
,~
'"
rO
N
I
0
~ 5
,~
0",
0<
~
00
, :;;
'" 5
«lI
,~
5
«l
5
,~
«l",
5 ~
'" <0
I
~
«l
5
",~
«l",
><~
I
«l
~
C
,~
«l",
0<
~
362
Fig. 362 Proportions of species of the four categories on some large islands in the Ryukyus. P,
Palearctic species; W, Wide-ranging species; E, endemic species; 0, Oriental species (see text).
Island codes are in parentheses.
170
Table 5. Distribution of eumenids in the Ryukyu and Daito Islands
Code' DJ SF SR SC ST SK ED EN ET RQ AF OK PO PI PI OaD AeJ PeY OeD EmR EmM DlE DIF PsD PzI" Code
1
2
+
+
4
5
6 +
8 +
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
10
11
+
12
13
14
+
+
+
19
20
21
22
24
25
26
+
+
+
+
28
29
30
31
+
36
+
+
+
+
+
+
+
+
42
+
43
44
45
46
48
49
+
+
+
39
40
41
+
4
5
+
+
+
+
+
52
6
5
+
+
+
+
+
+
+
+
+
10
11
12
13
14
IS
16
17
18
+
+
+
19
+
+
+
+
+
20
21
22
24
25
26
27
28
29
+
+
30
+
+
+
+
+
+
32
35
36
+
+
+
31
+
+
+
+
+
+
+
+
+
+ +
+
+
+
+
+
+
+
+
+
+
6
+
+
+
+
51
Total
2
+++++++
27
32
35
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+ + + +
+
+
+ +
+ +
+
+ +
+ + +
+ +
++++++
+ + + + +
+ + +
+
15
16 +
17
18
1
+
+
+
+
+
+
+
+
+ +
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
39
40
41
+
42
+
+
43
44
45
+
+
+
+
14 25 17 27 33
+
+
+
+
+
+
46
+
48
49
+
+
51
52
4
1
3
2
13
4
5
2
17
2
'Island codes as in Fig. 4 and Table 7.
"'DJ, Discoelius japonicus; SF, Stenodynerus frauenfeldi; SR, S. rufomaculatus; SC, S. chinensis; ST, S.
tokyanus; SK, S. kusigematii; ED, Euodynerus dantici; EN, E. nipanicus; ET, E. trilobus; RQ,
Rhynchium quinquecinctum; AF, Anterhynchium flavomarginatum; OK, Okinawepipona kogimai;
PO, Pararrhynchium ornatum; PT, P. tsunekii; PI, P. ishigakiense; OaD, Orancistrocerus drewseni;
AcJ, Ancistrocerus japonicus; PcY, "Pachymenes yayeyamensis"; OeD, Oreumenes decoratus; EuR,
Eumenes rubronotatus; EuM, E. micado; DIE, Delta esuriens; DlF, D. flavopictum; PsD, Pseumenes
depress us; PzI, Pseudozumia indosinensis.
171
Table 6. Subspecies differentiation in the three subregions of the Ryukyu Archipelago.
Eumenidae
A
B
N. Ryukyus
C. Ryukyus
S.Ryukyus
1
5
3
9
4
4
Total
Vespinae·
A
B
A
3
3
2
0
4
3
2
2
8
6
11
B··
6
• After Yamane (1987, 1988).
.. A: Number of species that have subspecies confined to the Ryukyus.
B: Number of their Ryukyu subspecies altogether found in each subregion.
highest, while that of Palearctic species is lowest. The Central Ryukyus are characterized
by the high proportion of wide-ranging species and the relatively high proportion of
endemic species. This agrees with the general expectation that the islands that are very
remote from the source areas mostly consists of specialized species (or relicts) and species
with a great ability to disperse. In the N. Ryukyus the Palearctic elements are dominant
and no Oriental and endemic species found.
The largest gap in species composition is found between Amami-Oshima and Yakushima, where the border of the Oriental and Palearctic regions is said to exist. This border,
originally recognized for land vertebrates and called Watase's line, is often not significant
for winged insects (lkehara, 1966). However, there is generally a profound gap in species
composition somewhere between these two islands even in insects (e. g., Terayama, 1982),
though the precise position of Watase's line is controversial and may depend on animal
groups. More important is whether this gap has been caused by geological history (early
establishment of the Tokara Strait) or by climatic factors. This question may be answered
only after the distribution of species (or groups) concerned is carefully studied in
continental China.
5.2. Subspecies differentiation. Five of the 25 species are more or less differentiated into
subspecies (geographical races) in the Ryukyu Archipelago. The number of subspecies per
species ranges from two to seven in the following order: Rhynchium quinquecindum and
Okinawepipona kogimai (2) - Euodynerus nipanicus (3) - Stenodynerus kusigematii (4) Anterhynchium f1avomarginatum (7). No Oriental species has subspecies peculiar to the
archipelago, while the wide-ranging and endemic types have an equal number (2) of
species that are differentiated into subspecies in the archipelago. The number of such
species and the total number of their subspecies endemic to the Ryukyus are both largest
in the C. Ryukyus (Table 6). This means again a longer history of isolation of the
populations in the C. Ryukyus mainly through distance effect.
The geographical borders of subspecies often lie between the three major subregions
(for example, in Euod. nipanicus and R. quinquecinctum). In other cases, however, distinct
subspecies occur on one or a few islands, some of which are very small (A. f1avomarginatum and S. kusigematil). According to Kimoto (1979) some chrysomelid beetles in the
Ryukyus also have distinct color forms on small islands such as Naka-no-shima, Okinoerabu-jima, Miyako-jima and Yonaguni-jima. He argues that these differentiations
occurred rapidly by genetic drift in small populations after the defaunation during the
RyukyfI period, when many low islands were submerged, followed by recolonization.
5.3. Area-species relation. The relationships between island areas and species numbers
for the Ryukyu eumenids and vespoids as a whole are shown in Table 7 and Figs. 363-366.
172
Table 7. Species numbers of vespoid groups in the Ryukyu and Daito Islands.
Code
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
24
25
26
27
28
29
30
31
32
35
36
39
40
41
42
43
44
45
46
48
49
51
52
Islands
Area
(km 2)
0.61
1.75
Kuro~hima
15.69
11.71
Io-jima
Take-shima
4.18
Tane-ga~hima
447.42
Mage~hima
8.40
Yaku~hima
500.59
Kuchinoerabu-jima 38.04
Kuchi-no-shima
13.25
Naka-no-shima
27.50
Suwanose-jima
22.32
Akuseki-jima
7.03
Takara-jima
5.94
Kikai-jima
55.71
Amami-oshima
718.74
Kakeroma-jima
75.15
Uke~hima
13.70
Yoro~hima
9.48
Tokuno~hima
248.11
Okinoerabu-jima 94.54
Yo ron-to
20.82
Izena-jima
13.87
Okinawa-jima
1182.52
Kouri-jima
2.99
Yagaji-jima
7.79
Sezoko-jima
3.43
Yabuchi-jima
0.62
Hamahiga-jima
2.01
Miyagi-jima
5.57
Tokashiki-jima
15.75
Kume-jima
58.50
Miyako-jima
159.00
Tarama-jima
19.88
Minna-jima
2.51
Ishigaki-jima
223.41
Taketomi-jima
5.53
Kuro~hima
10.04
Kohama-jima
8.26
Iriomote-jima
287.66
Hatoma-jima
1.00
Hateruma-jima
12.68
Yonaguni-jima
28.38
Kitadaito-jima
12.58
Minamidaito-jima 30.59
Ie-jima
Mukai-jima
Highest Eumenidae
Total
Vespidae
point(m)
Polistinae Vespinae
95.0
324.6
621.9
703.7
219.9
282.3
71.1
1935.3
657.0
628.3
979.0
799.0
584.0
291.9
224.0
694.4
329.0
398.4
297.0
644.8
246.0
97.1
119.9
498.0
107.4
55.2
40.8
42.4
78.7
121.4
227.3
326.0
114.6
34.4
8.0
525.8
20.5
14.0
99.4
469.7
33.8
59.5
231.2
74.6
75.8
2
1
4
2
2
10
4
12
7
1
2
6
3
2
3
1
4
3
9
4
3
2
5
7
6
3
10
2
5
5
1
3
2
3
7
9
4
2
11
5
3
5
10
3
7
8
1
1
3
5
2
3
2
3
3
2
2
5
3
3
2
1
4
2
2
4
3
2
2
5
2
2
2
4
2
2
3
1
1
3
1
1
5
4
6
2
2
3
2
1
2
3
2
7
3
3
19
4
24
12
3
7
1
2
5
6
17
8
6
5
10
10
8
7
16
5
8
7
2
7
4
5
11
1
1
3
1
1
1
3
1
13
7
5
19
8
6
8
17
5
10
12
Code
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
24
25
26
27
28
29
30
31
32
35
36
39
40
41
42
43
44
45
46
48
49
51
52
'Island codes correspond to those in Fig. 4.
-Eumenids are recorded also from Irabu-jima and Yubu-jima, but they are omitted because the
faunal survey is not sufficient.
Probably owing to insufficient surveys and reflecting variation in topography and
remoteness among the islands plotted, the points are widely scattered around the
regression line, but there is a general tendency common to both Eumenidae and
Vespoidea.
173
Since MacArthur and Wilson (1967) reformulated the area-species relation in the light
of new biogeographical theory, many archipelagos of the world have been resurveyed
with this view point. Several animal groups have been analyzed in the Ryukyu
Archipelago regarding the area-species relation (e.g., chrysomelid beetles by Kimoto, 1982;
land snails by Tomiyama, 1983, and Kurozumi, 1984; ants by Terayama & Yamane, 1984).
The results of all these studies fairly well fitted Arrhenius' equation, S=cAz .
Kimoto (1982) made a further analysis, comparing the area-species relation between
•
•
•
•
•
•
•
Ul
"'
"'""
H
U
Ul
•
""'0
0z
•
•
•• • • ••
S = 1.90AO.27
R2= 0.647
363
10
S = 2.46AO.27
R2= 0.740
gJ
5
H
U
"'""
S
1.89 AO.23
R2= 0.604
Ul
o""'
o
z
,
,,
.- .-
.-
,
NR
S = 1.45AO.33
R2= 0.811
5
10
AREA
I N
50
K M2
100
1000
364
Fig. 363. Area-species relation for the Ryfikyfi Eumenidae. Islands listed in Table 5 are plotted, but
most islets of the Tokara (10-13) and Daito (51, 52) groups are omitted because they are not
sufficiently surveyed.
Fig. 364. Area-species relation for the Ryukyfi Eumenidae. Regression lines are separately drawn
for N., C. and S. Ryukyus (NR, CR and SR respectively). 6, Tane-ga-shima; 8, Yaku-shima; 16,
Amami-<ishima; 25, Okinawa-jima; 36, Miyako-jima; 41, Ishigaki-jima; 45, Iriomote-jima.
174
the South Kurile Islands (situated around 44 ON) and the Ryukyu Islands. He discovered
that the number of chrysomelid species in the Ryukyus was constantly larger than that of
Southern Kuriles. In general, the number of insect species in relation with the area may be
larger in warmer regions. In this respect the regression lines in Figs. 364 and 366 are quite
interesting.
The fact that the largest species number is found in the S. Ryukyus seems to be
consistent with Kimoto's argument. The species numbers for the C. Ryukyus, however, are
not definitely larger than those for the N. Ryukyus. The large islands in the C. Ryukyus
(Okinawa-jima; Amami-Oshima) hold smaller species numbers than do slightly smaller
•
I
Ul
r.l
H
U
r.l
p..
Ul
~
o
o
z
•
•
S = 3.34 AO.26
R2= 0.711
365
20
S = 4.57Ao.24
R2= 0.850
Ul
10
r.l
H
U
r.l
~
S
3.67 AO.21
R2= 0.716
/
p..
Ul
/////'
5
0
az
/'/
/'\NR
S = 2.13AO.37
R2= 0.859
5
10
ARE A
INK M2
50
100
1000
366
Figs. 365. Area-species relation for the Ryfikyfi Vespoidea.
Figs. 366. Area-species relation for the Ryfikyfi Vespoidea. Regression lines are separately drawn
for N., C. and s. Ryukyus. For abbreviations and island code see Fig. 364.
175
islands in the N. Ryukyus (Yaku-shima; Tane-ga-shima). As a result the curves for these
two regions cross between 10-40 km 2. The geological history and location of these island
groups may be responsible for this. First, Yaku-shima and Tane-ga-shima are typical
continental islands and relatively close to the colonizing source (Kyushu), while some
small islands in this region such as islands of Mi-shima and Uji groups, remote from
colonizing source, are of volcanic origin or have long been isolated. All this no doubt
contributes to the larger values of z, or slope of curve <0.33 for Eumenidae; 0.36 for
Vespoidea).
On the other hand, the Central Ryukyus are less easily accessible by the immigrants,
so the equilibrium species number is expected to be relatively low. However, the small
islands in this island group are not so remote from the source regions (in this case
Okinawa-jima and Amami-oshima) or from each other (or at least clustered by means of
stepping stones). As MacArthur and Wilson (1967) pointed out, under these situations z
may be decreased, the area-species curve having a low slope. The low values of z for the S.
Ryukyus (0.26 for Eumenidae; 0.23 for Vespoidea) can be explained in a similar way.
However, although such analyses are not made, land snails (Tomiyama, 1983) and ants
(Terayama & Yamane, 1984) do not seem consistent with the present results.
5.4. Geographical replacement. The small islands are generally poor in eumenid fauna
as discussed above, and counterparts of supposed competing species are often lacking
(here, species with similar nesting behaviors are supposed to be competing).
In the N. Ryukyus two species Cfrauenfeldi and chinensis) of Stenodynerus are common,
but coexist only on larger islands (more than 30 km 2 in area). The Uji and Mi-shima
groups completely lack S. frauenfeldi, while on Mage-shima only S. chinensis has been
collected (Fig. 47). In this connection, however, other small islands in the other parts of
Japan have not yet been carefully surveyed. In the C. Ryukyus also, S. frauenfeldi is paired
with another closely related species, rufomaculatus, and they are completely allopatric. On
even large islands such as Amami-oshima and Okinawa-jima one of them is absent.
However, my observations on Yoron-to and Amami-6shima suggest that rufomaculatus
cannot be an all-out competitor with frauenfeldi, because the former is mostly confined to
the coast, while the latter is collected also inland. The above case of supposed geographical replacement, therefore, is open to criticism.
In the N. Ryukyus, Rhynchium quinquecinctum has been collected only on Naka-noshima of the Tokara group. Although extending as north as Niigata-ken, Honshu, this
species prefers warmer regions and is generally much rarer than Anterhynchium
fIavomarginatum, a supposed competitor, in any part of Japan proper, while it is quite
common in Southeast Asia including Taiwan. In general, on islands rare species are
expected to go extinct more easily than common species even without any competitive
encounter. Further, most of the small islands in the C. and S. Ryukyus harbor both R.
quinquecinctum and A. fIavomarginatum, thus the possibility of geographical replacement
by means of competitive exclusion is unplausible in this case. Euodynerus dantici is also
completely lacking on the islands of the N. Ryukyus. This species has a similar
distribution pattern as R. quinquecinctum, being much rarer than the supposed competitor
(Euod. nipanicus) in temperate Japan, while common in the S. Ryukyus and Taiwan.
Although competition might be partly responSible for the lack of R. quinquecinctum and
Euod. dantici in the N. Ryukyus, an alternative explanation (e.g., extinction by chance)
cannot be excluded.
A case of supposed geographical replacement has been reported for large carpenter
176
bees (Xylocopa) (Yamane et aI., 1983; Yamane, 1986). The four Ryukyu forms are very
closely related and completely allopatric, but distribution pattern is not of a typical
chessboard (cf. MacArthur, 1972). If these forms are distinct species (not subspecies of the
same species), the complete allopatric pattern is best explained by competitive exclusion,
though geological, topographical and climatic factors also might have played roles in the
formation of the present distribution pattern. Yamane (1986) suggested a chessboard
pattern for two social wasps of the subgenus Megapolistes in the Ryukyus. In this case the
two species (jadwigae and rothneyl) are similarly common on the Japanese mainlands and
similarly uncommon on Taiwan. After all, competitive exclusion cannot be excluded for
the lack of counterparts on many islands in the various parts of the Ryukyus (14 out of the
26 islands studied).
5.5. Regional convergence. On the Okinawa Islands, C. Ryukyus, several aculeate
species have exceptionally dark body markings. For example, Polistes rothneyi, usually
characterized by bright yellow markings in Japan and Taiwan, is often mistaken on
Okinawa-jima by its extremely dark body for P. jadwigae, which is usually darker than P.
rothneyi, even by professional entomologists. The Okinawa form of P. japonicus closely
parallels P. rothneyi. In Eumenidae, Anterhynchium flavomarginatum, Rhynchium
quinquecinctum and Okinawepipona kogimai are characterized by dark body markings
compared with the forms of other parts of Japan. On the other hand much smaller species
such as 5tenodynerus spp. and some Oriental elements, that are supposed to have
immigrated recently, are not affected in color pattern at all.
The forms of the S. Ryukyus of the species above mentioned are without exception
marked with bright yellow (0. kogimai does not occur in the S. Ryukyus). Furthermore,
body markings themselves are much more extensive so that sometimes the wasps look
entirely yellow. Some sphecids and pompilids are also extensively yellow-maculated.
The forms of these vespoid species in the N. Ryukyus, Amami Islands and Taiwan
are, roughly speaking, intermediate in coloration between those of the Okinawa group
and the S. Ryukyus. All this is not consistent with a general view that forms of warmer
regions have paler or brighter body markings. Apart from the real cause, the regional
convergence illusterated above is best explained by Mullerian mimicry as suggested by
Yamane (1988).
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184
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APPENDIX
After the completion of the main part of this paper, J. M. Cumming (Biosystematics
Research Centre, Agriculture Canada) published a world revision of the genus
Symmorphus (Cumming, 1989, Mem. Entomol. Soc. Can., 148: 1-168). Since I have not had
enough time to examine all his results, only his views concerning the Japanese forms are
summarized below.
New synonymy is as follows: (S. mutinensis auster C.S. and S. mutinensis yezoanus
Tsuneki) = S. bifasciatus (Linnaeus); (S. ishikawai C.S.) = S. lucens (Kostylev). S. sounkionis
Tsuneki is transferred as AncistToceTus sounkionis (Tsuneki) comb. nov.
185
INDEX TO GENERA AND SUBGENERA
Italicized names refer to junior synonyms or names incorrectly applied to the Japanese forms. Boldface
numbers refer to the page with the generic description.
Okinawepipona
15,19,22,83
Orands trocerus
13,16,19,21,95
Oreumenes
13,15,21,137,140
15,16,19,20,22,29,41
Alpha
140
16,19,21,88,90,98,123,164,168
Ancistrocerus
13,164
Antepipona
14,18-20,22,69,83,168
Anterhynchium
A1lodynerus
Delta
"Pachymenes"
15,19,21,135
Paraleptomenes
13
13,16,19,21,84,87,168,170
Pararrhynchium
Pareumenes
159,161,162
Pareuodynerus
52,57,59
Parrhynchium
87
69,83,84
Euancistrocerus
95,98,100,106
15,16,19,21,137,138,140,154
Eumenes
Euodynerus
14,15,18,19,22,41,46
Phi
154,155
Prorhynchium
87
Pseudepipona
164
Protodiscoelius
22
Protodynerus
130
Pseudozumia
16,19,20,162
Pseumenes
15,16,19,20,159
14
Psiloglossa
13
Calligaster
Coeleumenes
159,162
15,16,19,21,140,154
Deuterodiscoelius
22
69
Dirhynchium
15,19,20,22
Discoelius
Epiodynerus
Hoplomerus
Leionotus
l.ionotus
42,46,53,72
25,52,54,57,136
Mischocyttarus
Monteozumia
Montezumia
Monotezumia
155
162
159,162
162
Nannodynerus
Nortozumia
25
162
Odynerus
Raphiglossa
Rhygchium
Rhynchium
Rygchium
27
Stenodynerus
Symmorphus
Synagris
13
15,18,19,22,25,167,176
14-16,18-20,113,164
Tritodiscoelius
23
Vespl
Zethus
14,27,42,46,95,98,113
14
65
14,18,19,22,63,69,87
63,69
22,46,63,98,140,154
13
INDEX TO SPECIES AND SUBSPECIES
Boldface numbers refer to the page with species or subspecies deSCription.
arcuata, Phi
158
aterrinus, Orancistrocerus
98
59
atripes, Euodynerus
auster,Symmorphus
131
adiabatus, Ancistrocerus
12
amamense, Anterhynchium
70,72,77,78
antilope, Ancistrocerus
99,112,165
112
antilope, Vespa
apiciomatus, Ancistrocerus
118
apiciomatus, Odynerus
27,28,118,120,123
apiciornatus,Symmorphus
115,123,126,165
aquilonius, Eumenes
141,148
architectus, Eumenes
145,150
72
ardens, Rhynchium
arcuata, Eumenes
158
bicingulatus, Euodynerus
bifasciatus,Symmorphus
brachytomus, Euodynerus
brunneum, Rhynchium
caffer, Eumenes
186
157
59,164
185
50
14,63
callosus, Odynerus
109,110
campaniforme, Delta
155
campaniformis, Eumenes
155
canadensis,Symmorphus
114
captivus, Odynerus
116,118,120,123
captivus,Symmorphus
113,114,120,123
carinatus,Symmorphus
115,125
chinensis, Odynerus
25,31
26,27,31,35,167,169,
chinensis,Stenodynerus
176
diens,Symmorphus
115,116,133
dypeopictus, Odynerus
35
26,27,35,167
dypeopictus,Stenodynerus
coarctata, Vespa
140
cyanopterus, Calli gaster
13
flavopictum, Delta
15,158,170
flavopictus, Eumenes
158
flavopunctatum, Anterhynchium
69,83
flavopunctatum, Rhynchium
75
flavoscutellatus,Stenodynerus
37
floricola, Odynerus
41
foraminatus, Euodynerus
12
formosana, Eumenes
158
formosanum, Delta
158
formosanus, Eumenes
158
foveolatus,Symmorphus
16,114,116,165
fragilis, Odynerus
136
fragilis, Pachymenes
136
fratercula, Eumenes
142
fraterculus, Eumenes
141,142
fraterna, Eumenes
142
27,31
frauenfeldi, Odynerus
26,27,46,167-169,
frauenfeldi,Stenodynerus
176
fukaianus, Ancistrocerus
95,98
95
fukaianus, Odynerus
Fukaii, Odynerus
65
65
fukaii, Rhygchium
fukaii, Rhynchium
14,65,66,68
fukayanus, Odynerus
95
dantici, Euodynerus
47,48,170,176
dantici,Odynerus
50
dantici, Vespa
46,48
decens, Odynerus
118
113,114,118
decens, Symmorphus
decoratus, Eurhenes
137,138
decoratus,Oreumenes
13,74,138,165,167,
170
delphinalis, Allodynerus
42,45,46
42
delphinalis, Odynerus
densepilocellus, Ancistrocerus
106,108
106
densepilocellus, Odynerus
depressa, Pareumenes
159
depress us, Eumenes
159
depressus, Pseumenes
19,159,170
depresus, Pseumenes
159
dimidiatidypeus, Eumenes
145
95
drewseni, Odynerus
drewseni, Orancistrocerus
13,74,83,95,165,
167,170
ebumus, Euodynerus
58
elegans, Odynerus
113
emarginatum, Delta
154
60
emma, Odynerus
erythropus, Orancistrocerus
23
esakii, Discoelius
esuriens, Delta
155,170
esuriens, Eumenes
155
europaeum, Rygchium
63
eximius, Eumenes
159
gracilis, Eumenes
155
haemorrhoidale, Rhynchium
63-68
65
haemorrhoidalis, Odynerus
hanedai, Anterhynchium
70,71,79
harmandi, Eumenes
137,138
h6koti>ensis, Odynerus
136,137
indica, Montezumia
162
indosinensis, Pseudozumia
162,170
ingens,Orancistrocerus
98
70,72,77
insulicola, Anterhynchium
88,170
ishigakiense, Pararrhynchium
ishigakiensis, Ancis trocerus
88
ishikawai,Symmorphus
115,134
iwatai,Symmorphus
115,127
98
japonicum, Rhygchium
74
japonicum, Rhynchium
72,74
japonicus, Ancistrocerus
46,100,164,169
japonicus, Discoelius
23,169
japonicus, Eumenes
138
japonicus Matsum., Odynerus
35
japonicus Schult., Odynerus
35,100
flavidypeatus,Odynerus
54
flavicornis, EUodynerus
53,56,57
flavolineatus,Odynerus
53,56,57
flavomarginatum, Anterhynchium
14,20,60,
69,83,165,168,170-172
flavomarginatum, Rhynchium
69,95
kikaiensis, Stenodynerus
kogimai, Anterhynchium
kOgimai, Okinawepipona
187
30
83
84,86,168,170,
koreanum, Anterhynchium
kusigematii,Stenodynerus
lucens,Symmorphus
185
luctuosum, Anterhynchium
pachymenoides, Stenodynerus
38,41
parietina, Vespa
108
100,108,112
parietinus, Ancistrocerus
98
parietum, Vespa
142
pomiformis, Eumenes
71,72,748,76,168
procella, Anterhynchium
53,54
pubescens, Euodynerus
140,141,150,167
punetatus, Eumenes
172,177
74,168
26,38,40,170,
172
79
23
manchurianus, Discoelius
72,79
mandarineum, Rhynchium
mandarinum, Rhynchium
72
29,45
mandschuricus, Allodynerus
maxillosa, Vespa
154
melanocerus, Ancistrocerus
100,106
106
melanocerus, Odynerus
melanopterum, Anterhynchium
69,82,167
13
mephitis, Paraleptomenes
micado, Anterhynchium
14,60,71,72,76,79,
83,98
114,115,121,131,133,134
micado, Eumenes
52,53,72
micado, Odynerus
72
micado, Rhygchium
152
mikado, Eumenes
94
miyanoi, Pararrhynchium
115,116,132
mizuhonis,Symmorphus
109
mongolicus, Ancistrocerus
murotai, Rhynchium
65,66
mutinensis, Odynerus
130
mutinensis, Symmorphus
115,,130-132
quadrifasciata, Vespa
57
quadrifasciatus, Euodynerus
quadrifasciatus, Odynerus
quadrispinosus, Pareumenes
63
quinquecincta, Vespa
quinquecinctum, Rhynehium
47,57-59
14,52,54,57
159,161
14,63-68,
170-172,176
141,142,144
rubrofemoratus G.5., Eumenes
144
rubrofemoratus Per., Eumenes
144
rubrofemoratus Tos., Eumenes
rubronotatus, Eumenes
141,142,145,151,
152,154,165,169
152
rufeseens, Eumenes
rufiventris,Stenodynerus
38,39
rufomaculatus,Stenodynerus
26-28,30,170,
176
34
rufoomatus, Stenodynerus
53,57
ryukyuensis, Euodynerus
samuray, Eumenes
142,145,148,152-154
samurayi, Eumenes
152
63
samurayi, Rhynehium
54
satsumanus, Rhynehium
serieea, Paehymenes
135
100,104,106,107
shibuyai, Ancistrocerus
shibuyai, Odynerus
104
shinto, Odynerus
90
simillimus, Stenodynerus
31
Simulator, Ancistrocerus
12
sinuatus, Odynerus
130
smithii, Rhynchium
87
sounkionis, Ancistrocerus
185
114,123
sounkionis, Symmorphus
12
spilogaster, Ancistrocerus
72,77
sulphreum, Anterhynchium
synagroides, Rygehium
69
85
nagasei,Okinawepipona
65,68
nambui, Rhynchium
27
nigric!ypeatus,Odynerus
nigricomis, Ancistrocerus
100,106,109
109
nigricomis,Odynerus
69
nigrifrons, Anterhynchium
SO,51
nigrescens, Euodynerus
nigripes, Odynerus
54
nipanicus, Euodynerus
14,17,52,53,167-169,
172
nipanicus, Lionotus
52
notatus, Euodynerus
52,54,56
oceanicum, Pararrhynchium
88,93,94,168
63
oculata, Vespa
ogasawaraensis,Stenodynerus
26,33,34,168
okinawae, Delta
155
omatum, Pararrhynchium
13,88,89,92,93,
167,170
omatum, Rhynchium
87,89
omatus, Ancistrocerus
90,92
omatus, Odynerus
89
oviventris, Ancistrocerus
99,110
110
oviventris, Odynerus
teutonicus, Lionotus
25
tokyanus, Odynerus
36
26,28,36,38,164
tokyanus, Stenodynerus
167,169
52,57
tomentosus, Lionotus
100,104
trifasciatus, Ancistrocerus
60
triloba, Vespa
188
trilobus, Euodynerus
47,60,170
trilobus, Odynerus
60
trilobus, Pseudoepipona
62
troglodytes, "Pachymenes"
136
13
tropicalis, Antepipona
88,92,170
tsunekii, Pararrhynchium
38,39
tsunekii,Stenodynerus
115,128,167
tsushimanus, Symmorphus
71,75,167
tsushimarum, Anterhynchium
75
tsushimarum, Rhynchium
12
tuberculicpes, Andstrocerus
umenoi, Anterhynchium
umenoi, Rhynchium
80
varipes, Rhynchium
72
violaceipennis, Euodynerus
50,51
yambarah,Stenodynerus
38
yamanei, Ancistrocerus
111,112
yayeyamensis, Odynerus
136
yayeyamensis, "Pachymenes"
136,168,170
yezoanus, Symmorphus
132
23
zonalis, Discoelius
zonalis, Vespa
22
70,71,79,80
189