http://dx.doi.org/10.5852/eit.2015.137

www. europeani ournaloftaxonomv. eu

2015 • Murao R., Tadauchi O. & Lee H.-S.

This work is licensed under a Creative Commons Attribution 3.0 License.

Monograph

urn:lsid:zoobank.org:pub:0CB83C33-E198-4B4F-A8F6-96C59B066112

Synopsis of Lasioglossum (Dialictus) Robertson, 1902 (Hymenoptera,

Apoidea, Halictidae) in Japan, the Korean Peninsula and Taiwan

Ryuki MURAO '•*, Osamu TADAUCHI 2 & Heung-Sik LEE 3

12 Department of Biological Science, Faculty of Sciences, Kyushu University,

Fukuoka, 812-8581 Japan.

3 Animal and Plant Quarantine Agency, Suwon, Korea.

* Corresponding author: [email protected] . [email protected]

1 urn:lsid:zoobank.org:author:7E665839-6675-4365-A7EA-169D4E4DA008

2 um:lsid:zoobank.org:author: 1A22CC5E-A7A0-449E-994D-D939EEF5BF42

3 um:lsid:zoobank.org:author:BEA91CE0-9306-4AC4-8E90-5FD07A69B7AE

Abstract. Twelve species in total are recognized for the subgenus Lasioglossum ( Dialictus ) Robertson,

1902 from Japan, the Korean Peninsula and Taiwan. Lasioglossum ( Dialictus ) alishanense sp. nov.

and L. ( D .) taiwanense sp. nov. from Taiwan, are described as new. Four species are recorded from

the following localities for the first time: L. (D.) miyabei Murao, Ebmer & Tadauchi, 2006 from the

Korean Peninsula (South Korea); L. (D.) ellipticeps (Bluthgen, 1923), L. (D.) virideglaucum Ebmer &

Salcagami, 1994 and L. (D.) viridellum (Cockerell, 1931) from South Korea. Lasioglossum ( Dialictus )

atroglaucum (Strand, 1913) is redescribed, L. (D.) ellipticeps , L. ( D .) problematicum (Bluthgen,

1923), L. ( D .) virideglaucum , and L. ( D .) viridellum are supplementary described. A key to species is

provided. Bionomical data, such as flight and flower records or habitat, are reported for some species.

The distributions of all species are mapped. DNA barcodes of L. (D.) miyabei , L. (. D .) pseudannulipes

(Bluthgen, 1925), and L. (D.) virideglaucum are provided.

Key words. Halictidae, Lasioglossum , Dialictus , Far East Asia.

Murao R., Tadauchi O. & Lee H.-S. 2015. Synopsis of Lasioglossum (. Dialictus ) Robertson, 1902 (Hymenoptera,

Apoidea, Halictidae) in Japan, the Korean Peninsula and Taiwan. European Journal of Taxonomy 137: 1-50. http://

dx.doi.org/10.5852.ejt.137

Introduction

The cosmopolitan bee genus Lasioglossum Curtis, 1833 (Halictidae, Halictinae) is a highly diverse

group and includes approximately 1800 species worldwide (Ascher & Pickering 2015). This group

contains diverse social structures (solitary, parasocial, and primitively eusocial, etc., reviewed in, e.g.,

Michener 1974; Yanega 1997), and are some of the most abundant bees in the warm temperate zone

and parts of the Neotropical Region (Salcagami et al. 1967; Salcagami & Fulcuda 1973; Yamauchi et al.

1974; Ilcudome 1992; Maeta et al. 2003; Goukon 2006; etc.). It is divided into two informal group on the

basis of the degree of strength of the second submarginal vein of the female fore wing (Michener 2000):

l)the Hemihalictus series (weak-veined Lasioglossum) includes all subgenera with a weak second

submarginal vein ( Acanthalictus Cockerell, 1924, Austrevylaeus Michener, 1965, Capalictus Pauly,

European Journal of Taxonomy 137 : 1-50 ( 2015 )

Gibbs & Kuhlmann, 2012, Dialictus Robertson, 1902, Dialictus sensu lato, Eickwortia McGinley,

1999, Evylaeus Robertson, 1902, Evylaeus sensu lato, Hemihalictus Cockerell, 1897, Paradialictus

Pauly, 1984b and Sphecodogastra Ashmead, 1899).

2) the Lasioglossum series (strong-veined Lasioglossum) includes all subgenera {Australictus

Michener, 1965, Callalictus Michener, 1965, Chilalictus Michener, 1965, Ctenonomia Cameron,

1903, Glossalictus Michener, 1965, Homalictus Cockerell, 1919, Ipomalictus Pauly, 1999,

Lasioglossum, Leuchalictus Warnclce, 1975, Papualictus Michener, 1980, Parachilalictus Pauly

et al. , 2013, Parasphecodes Smith, 1853, Pseudochilalictus Michener, 1965, Quasilictus Walker,

1986, Rubrihalictus Pauly, 1999 and Urohalictus Michener, 1980) with a strong second submarginal

vein (modified from Michener (2007), including the subgenera proposed or re-established in recent

phylogenetic and taxonomic studies (Gibbs et al. 2012, 2013; Pauly et al. 2012, 2013)).

Each series is monophyletic as supported by phylogenetic analyses based on molecular data (Danforth

et al. 2003; Gibbs et al. 2012). However, there is no agreement among taxonomists with regard to the

number and extent of the subgeneric classification of Lasioglossum and it is not necessarily applied

when classifying the male. The Palaearctic subgenera Acanthalictus, Evylaeus, Dialictus, Hemihalictus

and Sphecodogastra of the Hemihalictus series, e.g., have been treated as L. (Evylaeus ) (e.g., Ebmer

2002; Murao & Tadauchi 2007; Pesenko 2007a, as the genus Evylaeus). Recent phylogenetic studies

(Danforth 1999; Danforth et al. 2003; Gibbs et al. 2012, 2013) based on molecular data, however, do

not recognize L. {Evylaeus), used by most Palaearctic researchers, as a monophyletic group. They will

probably shed light on our understanding of the relationships among taxa, evolutionary patterns of

sociality and the strict delimitation of genera and subgenera.

In eastern Asia, taxonomic studies of the Lasioglossum group, with 217 species (Murao 2015), have

been provided from the following areas: East Palaearctic Region (Pesenko 2006); Mongolia (Ebmer

1982, 2005); Russian Far East (Ebmer 1996, 2006; Pesenko 2007b); China (Ebmer 1998, 2002); Japan

(e.g., Murao & Tadauchi 2007; Murao et al. 2009b); Korean Peninsula (Ebmer 1978; Murao et al. 2014).

However, the basic information on the Lasioglossum fauna in eastern Asia is still poor, particularly in

the Oriental Region.

Dialictus Robertson, 1902 is one of the subgenera in Lasioglossum. This name has been applied to the

majority of species with the metallic form, including the black one without complete lateral carina on the

posterior surface of the propodeum (referred to as black Dialictus, acarinate, noncarinate or carinaless

Evylaeus) in the Hemihalictus series (e.g., Michener 2000, 2007). On the other hand, Dialictus in the

Palaearctic Region have been treated as the green Lasioglossum {Evylaeus) (Ebmer 2002; Pesenko

2007a). In the recent treatment of Lasioglossum, as outlined above, mostly metallic species in the

Hemihalictus series are recognized as Dialictus (Gibbs et al. 2013). 420 L. {Dialictus) species are listed

in the world (Ascher & Pickering 2015, including Afrodialictus Pauly, 1984). Recently, L. {Dialictus)

species from Canada and part of North America were revised by Gibbs (2010,2011), who provided useful

identification keys and good photographs of many specimens, whereas for those of eastern Asia (Russian

Far East and eastern Siberia) only an identification key is available (Pesenko 2007b, as Evylaeus).

A further 10 species of Dialictus have been partially described or reported from Japan, the Korean

Peninsula and/or Taiwan by various researchers (Japan: Ebmer & Salcagami 1990; Ebmer et al. 1994;

Murao & Tadauchi 2008; Murao et al. 2006, 2009a; Usui et al. 1976. Korean Peninsula: Ebmer 1978.

Taiwan: Strand 1913). No revisional study has been conducted to date in these countries. In the course

of a collaborative study of the Asian Lasioglossum fauna, we examined the L. {Dialictus) specimens

collected from Japan, the Korean Peninsula and Taiwan. Through our examination, we recognized 12 L.

{Dialictus) species in total for these countries. Among them, two new species were found from Taiwan

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

and four known species were recorded from South Korea for the first time. In this paper, two new species

from Taiwan are described and the remainder are redescribed or additionally described, and drawings,

photographs and scanning electron micrographs for diagnostically important characters are provided.

DNA barcodes of some species and an illustrated key of all species are presented to facilitate species

identification in these countries.

Material and method

Collection

This study is based on the specimens deposited in the following institutions and personal collections,

which are referred to using the following abbreviations:

BPBM

cGou

cMur

ELKU

MNHAH

OLML

QIA

SDEI

SNU

SULE

ZISP

Maa’s collection in the Bernice P. Bishop Museum, Honolulu, Hawaii, USA

private collection of Mr. Katsuo Goukon, Miyagi, Japan

the first author’s private collection, Fukuoka, Japan

Entomological Laboratory, Faculty of Agriculture, Kyushu University, Fukuoka, Japan

the late Dr Shoichi F. Sakagami’s collection, deposited in the Museum of Nature and

Human Activities, Hyogo, Japan

Biologiezentrum/Oberosterreichisches Landesmuseum, Linz, Austria

Animal and Plant Quarantine Agency, Suwon, Korea

Senckenberg Deutsches Entomologisches Institut, Miincheberg, Germany

Insect Systematics Laboratory, Seoul National University, Seoul, Korea

Dr Yasuo Maeta’s collection, deposited in the Laboratory of Insect Ecology,

Faculty of Life and Environmental Science, Shimane University, Shimane, Japan

Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia

Terminology

Terminology and style used in the descriptions follow Murao et al. (2014). Abbreviations used in the

text are as follows:

AOD

CAL

CPL

FnL

FnW

IAD

IOD

LOD

MNL

MOD

antennocular distance (shortest distance between outer margin of antennal socket and inner

margin of compound eye)

body length (from antennal base to tip of pygidial plate)

clypealveolar distance (between lower margin of antennal socket and lower margin of

supraclypeus in frontal view)

clypeal length (between upper and lower margins of clypeus in frontal view)

eye length

eye width (maximum length and width of the compound eye)

nth antennal flagellomere

length of nth flagellomere (measured along the ventral surface)

width of nth flagellomere (measured from dorsal and ventral surfaces of flagellomere)

genal width (maximum width of the genal area when seen in lateral view)

head length (from top of vertex to lower margin of clypeus)

head width (between outer margins of compound eyes in frontal view)

interantennal distance (between inner margins of antennal socket)

interocellar distance (between lateral ocelli)

interspace between punctures (e.g., IS 0.5d means 1/2 of the diameter of a puncture)

lower interorbital distance

metanotal length

maximum interorbital distance

European Journal of Taxonomy 137 : 1-50 ( 2015 )

MPL

MsW

MtW

OCD

OOD

SCL

SPL

UOD

metapostnotal length

maximum mesosomal width

maximum metasomal width

ocelloccipital distance (shortest distance between margins of lateral ocellus and vertex when

seen in upper view)

ocellocular distance (shortest distance between lateral ocellus and inner margin of compound

eye)

punctures

mesoscutellar length

nth metasomal sternum

scape length (a straight line from base to tip of scape)

nth metasomal tergum

upper interorbital distance

wing length (length of fore wing from the apical point to the base including tegula)

Body measurements are given in ranges followed by the average and standard deviation.

Flower records

Records of flowers visited by each species are based on specimen label data. The scientific names of

flowering plants visited by bees are cited from Yonekura & Kajita (2003-).

DNA barcoding

DNA extraction and PCR were conducted at the Kyushu University Museum (Fukuoka,

Japan). DNA was extracted using a DNeasy Blood and Tissue Kit (Qiagen, Tokyo, Japan)

following the manufacturer’s instructions. A region of the cytochrome oxidase subunit

1 (COI) gene of mtDNA was amplified using polymerase chain reaction (PCR) with

the primers LCO1490 (5 -GGTCAACAAATCATAAAGATATTGG-3') and HC02198

(5 -TAAACTTCAGGGTGACCAAAAAATCA- 3') (Folmer et al. 1994). Each PCR mixture contained

2 pi of DNA template, 2.13 pi of 10x EX Taq Buffer (Takara bio), 0.75 pi of each primer (20 pM), 1.68

pi of dNTP (2.5 mM, Takara bio), and 0.25 pi of EX Taq DNA polymerase (Takara bio). MilliQ water

was added to reach a total reaction volume of 20 pi. PCR conditions were as follows: 94 °C for 1 min,

followed by 10 cycles of 94 °C for 1 min, 48 °C for 1 min 30 s, and 72 °C for 1 min 30 s, then 30 cycles

at 94 °C for 1 min, 52 °C for 1 min 30 s, 72 °C for 1 min 30 s, and a final 72 °C for 5 min. PCR products

were run on a 1% agarose gel for 20 min, and then purified with 2.6 pi of ExoSAP-IT (GE Healthcare

Life Sciences). DNA sequencing was outsourced to the FASMAC Co., Ltd. (Kanagawa, Japan). The

resulting sequences were aligned using the program MEGA (ver. 5; Tamura et al. 2011). The sequence

data reported in the present paper are deposited at the DNA Data Base of Japan (DDBJ).

Comparative material examined

IRAN: Lasioglossum ( Dialictus ) aeratum (Kirby, 1802): 1 c?, Kuhha-ye, Tu-Chal, N Iran, alt. 2000-

2500 m, 18-19 Jul. 1970 (C. Elburz, MNHAH).

IRAN: Lasioglossum (. Dialictus ) algirum (Bluthgen, 1923): 1 S, Kopet Dagh, 30 km N Quchan,

Pafihohe, alt. 2000 m, 18 Jul. 1977 (J. Gusenleitner, MNHAH).

IRAN: Lasioglossum ( Dialictus ) annulipes (Morawitz, 1876): 1 Kopet Dagh, 30 km N Quchan,

Pafihohe, alt. 2000 m, 18 Jul. 1977 (J. Gusenleitner, MNHAH).

AUSTRIA: Lasioglossum ( Dialictus ) leucopus (Kirby, 1802): 1 Karnten, Farchtnersee, 23 Aug. 1983

(J. Gusenleitner, MNHAH).

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

CHINA: Lasioglossum ( Dialictus ) sanitarium (Bluthgen, 1926): 15 5 $, 4 SS, Fukien, Chungan, Bohea

Hill, 6 May 1938 (T.C. Maa, BPBM).

CHINA: Lasioglossum ( Dialictus ) sichuanense Fan & Ebmer, 1992: 1 § (paratype), Omei, Sichuan

(label in Chinese), 10 May 1964 (MNHAH); 1 S, Lijiang, alt. 1800 m, Yunnan Prov., 26°53’ N 100°18’

E, 23 Jun-21 Jul. 1992 (S. Becvar, cMur).

CHINA: Lasioglossum (Dialictus) xizangense Fan & Ebmer, 1992: 4 $ Fukien, Chungan, Bohea Hill,

May 1938 (T.C. Maa, BPBM); 1 S (paratype), Motuo, Xizang (label in Chinese), alt. 1300 m, 27 Nov.

1982 (Han Yin-heng, MNHAH).

Results

Class Hexapoda Blainville, 1816

Order Hymenoptera Linnaeus, 1758

Superfamily Apoidea Latreille, 1802

Epifamily Anthophila Latreille, 1804

Family Halictidae Thomson, 1869

Subfamily Halictinae Thomson, 1869

Tribe Halictini Thomson, 1869

Genus Lasioglossum Curtis, 1833

Lasioglossum (Dialictus ) Robertson, 1902.

Paralictus Robertson, 1901: 229. Type species: Halictus cephalicus Robertson, 1892 (preoccupied by

Morawitz 1873 = Halictus cephalotes Dalla Torre, 1896), by original designation.

Dialictus Robertson, 1902: 48. Type species: Halictus anomalus Robertson, 1892, by original designation

and monotypy.

Chloralictus Robertson, 1902: 248. Type species: Halictus cressonii Robertson, 1890, by original

designation.

Halictus (Gastrolictus ) Ducke, 1902: 102. Type species: Halictus osmioides Ducke, 1902, by monotypy.

Halictomorpha Schrottky, 1911: 81. Type species: Halictomorphaphaedra Schrottky, 1911, by original

designation.

Rhynchalictus Moure, 1947: 5. Type species: Rhynchalictus rostratus Moure, 1947, by original

designation.

Halictus (Smeathhalictus) Warncke, 1975: 88. Type species: Melitta smeathmanella Kirby, 1802, by

original designation.

Habralictellus Moure & Hard, 1982: 46. Type species: Halictus auratus Ashmead, 1900, by original

designation.

Lasioglossum (Afrodialictus) Pauly, 1984: 142. Type species: Halictus bellulus Vachal, 1909, by original

designation.

Evylaeus (Viridihalictus ) Pesenko, 2007a: 26. Type species: Halictus viridis Brulle, 1840, by original

designation.

Evylaeus (Glauchalictus ) Pesenko, 2007a: 26. Type species: Halictusproblematicus Bluthgen, 1923, by

original designation.

Evylaeus (Virenshalictus ) Pesenko, 2007a: 26. Type species: Halictus virens Erichson, 1835, by original

designation.

Evylaeus (Loethalictus ) Pesenko, 2007a: 26. Type species: Halictus loetus Brulle, 1840, by original

designation.

Evylaeus (Aerathalictus) Pesenko, 2007a: 27. Type species: Melitta aerata Kirby, 1802, by original

designation.

European Journal of Taxonomy 137 : 1-50 ( 2015 )

Diagnosis

This subgenus is usually separated from the other subgenera of Lasioglossum occurring in eastern Asia

by having a combination of the head and mesosoma with brilliant or dull green-metallic luster in both

sexes and the female fore wing with a weak second submarginal vein. In eastern Asia, the female of the

Fig. 1. A-E. Lasioglossum ( Dialictus ) viridellum (Cockerell, 1931). A-B. Head in frontal view.

C-D. Labrum. E. Mesepisternum. A, C. ?. B, D-E. $.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

apristum group belonging to L. (Sphecodogastra) (corresponding to the carinate L. (Evylaeus ) in the

dissenting classification (Ebmer 2002; Pesenko 2007a, as the genus Evylaeus )) has the same character

states as L. ( Dialictus ). However, the members of this group can easily be separated from L. ( Dialictus)

in having the mesepisternum coarse reticulate-rugulose, the posterior surface of the propodeum with

strong complete carina, and the inner hind tibial spur finely serrate.

Comments

As stated above, the green L. (Evylaeus ) used by most Palaearctic researchers corresponds to L. ( Dialictus )

in the recent classification. Pesenko (2007a) re-classified Evylaeus sensu lato into 29 subgenera for the

Palaearctic Region. In his systematic study, 6 of the subgenera (Aerathalictus Pesenko, Glauchalictus

Pesenko, Loethalictus Pesenko, 2007, Smeathhalictus Wamcke, 1975, Viridihalictus Pesenko, 2007 and

Virenshalictus Pesenko, 2007) correspond to L. ( Dialictus ) as shown above in the synonymy list (synonymies

by Gibbs 2010). These subgenera sensu Pesenko (2007a) are treated as a species group in this paper.

The leucopus group

Species included in eastern Asia:

L. aeratum (Kirby, 1802)

L. algirum (Bliithgen, 1923)

L. angaricum (Cockerell, 1937)

L. leucopus (Kirby, 1802)

L. meakanense Murao & Tadauchi, 2009

L. pseudannulipes (Bluthgen, 1925)

L. viridellum (Cockerell, 1931)

Among the members of this group, both L. meakanense and L. pseudannulipes are known from Japan,

and L. viridellum from the Korean Peninsula.

Diagnosis

Species of the leucopus group may be strictly characterized by a combination of the following character

states: 1) male F2 short, approximately 1.2-1.5 x as long as FI; 2) mesepisternum reticulate-punctate on

lower area in both sexes (Fig. IE); 3) male metasomal sterna not modified (apical margin straight); 4)

ventral retrorse lobe of male genitalia with moderately long setae only apically and distinct lineolation

on surface (Fig. 2D, F-I).

The diagnostic characters of this group are also shown in Ebmer & Sakagami (1990, as the leucopus

group) and Pesenko (2007a, as subgenus Aerathalictus Pesenko, 2007).

Distribution

Palaearctic to northern Oriental Region.

Comments

This group is assumed to be monophyletic, based on the ventral retrorse lobe of male genitalia with

moderately long setae only apically and the distinct lineolation on the surface. This character state may

be unique within the subgenus. A comprehensive comparative study of the ventral retrorse lobe has not

been performed for the subgenus, but this state may be an autapomorphy for the leucopus group.

European Journal of Taxonomy 137 : 1-50 ( 2015 )

Fig. 2. — A-E. Lasioglossum (Dialictus) viridellum (Cockerell, 1931), S. —F. L. ( D .) aeratum (Kirby,

1802), S. — G. L. (D.) aligrum (Bliithgen, 1923), S. — H. L. (D.) annulipes (Morawitz, 1876), $.

— I. L. (D.) leucopus (Kirby, 1802), f. A. Genitalia in ventral view. B. Genitalia in dorsal view. C.

Genitalia in lateral view. D, F-I. Ventral retrorse lobe of genitalia. E. S7-S8. Scale bars: A-C, E = 0.2

mm, D, F-I = 0.1 mm.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Lasioglossum ( Dialictus ) pseudannulipes (Bluthgen, 1925)

Figs 3A, 25B

Halictus pseudannulipes Bluthgen, 1925: 128 (holotype: Museum fur Naturkunde Humboldt an der

Universitat zu Berlin, Germany, $; type locality: Kwangtung Prov. (Guangdong Prov.), China).

Halictus pseudannulipes - Hirashima 1957: 16.

Lasioglossum ( Evylaeus) algirum pseudannulipes - Ebmer 1978: 313. — Ebmer & Sakagami 1990:

835-837.

Lasioglossum (Evylaeus) pseudannulipes -Murao et al. 2009a: 166-171 (c?)-

Diagnosis

This species is similar to Lasioglossum angaricum (Cockerell, 1937) and L. viridellum (Cockerell, 1931)

from eastern Asia in the leucopus group. It is separated from L. angaricum by the female T1 usually

being without lineolation (sometimes T1 basally lineolate, but weaker than in L. angaricum) and the

male sterna with sparse uniform hairs. In contrast, in L. angaricum the female T1 has distinct lineolation

and the male S3-S5 is medially bare, laterally with tufts of long dense erect hairs (Pesenko 2007b). For

the differences between females of this species and L. viridellum , see Key.

Material examined

JAPAN: Kyushu: 1 §, Matoishi wilderness, Aso-machi, Aso-shi, Kumamoto Pref., 37°27 , 15” N

128° V 10” E, 14 Aug. 2010 (R. Murao, cMur); 8 §§, Aso-shi, Kumamoto Pref., 33°0 , 16.115” N

131°8’5.658” E, 21 Jul. 2013 (R. Murao, cMur).

Distribution

China (Guangdong Prov.), Japan (Hokkaido, Honshu, and Kyushu).

Flight records

Female: April to October.

Male: June to October.

Flower records

Thirty-two species in 6 families were reported as floral hosts in Japan by Goubara et al. (2004).

Habitat

This species was collected from grassland in the Kumamoto Pref., Kyushu, western Japan. One of the

collecting sites there is shown in Fig. 23B.

DNA barcodes

The COI gene sequences are deposited as DNA barcodes of L. (D.) pseudannulipes in the DDBJ under

accession numbers LC027534 and LC027535. These numbers are also available in GenBank.

Lasioglossum (Dialictus) viridellum (Cockerell, 1931)

Figs 1, 2A-E, 3B

Halictus (Chloralictus) viridellus Cockerell, 1931: 14 (holotype: American Museum of Natural History,

New York, USA, §; type locality: Shanghai, China).

Halictus (Chloralictus) viridellus - Hirashima 1957: 24.

European Journal of Taxonomy 137: 1-50 (2015)

Lasioglossum (Evylaeus) viridellum -Ebmer 1978: 312-313 (c?)- —Ebmer 1996: 283-284. —Ebmer

2006: 568.

Evylaeus ( Aerathalictus ) viridellus - Pesenko, 2007a: 27. —Pesenko 2007b: 85 (in key), 97 (in key), 114.

Diagnosis

This species is closely similar to Lasioglossum angaricum (Cockerell, 1937) and L. pseudannulipes

(Bluthgen, 1925), as stated above. It is separated from L. angaricum by the disc of male sterna having

sparse and uniform hairs. In contrast, in L. angaricum , the male S3-S5 are medially bare, laterally with

tufts of long dense erect hairs (Pesenko 2007b).

Material examined

SOUTH KOREA: Gangwon-do: 2 § §, 2 Hong Cheon, 5-6 Jun. 1984 (K. Morimoto, ELKU); Jeju-

do, Jeju Is.: 11 $ $, Sanisu-dong Namjeju-gun, 23 Apr. 1997 (O. Tadauchi, ELKU); 1 §, Pijarim Forests

Pulccheju-gun, 24 Apr. 1997 (O. Tadauchi, ELKU); 2 5?, Kwangpyong-ri Namjeju-gun, 23 Apr. 1997

(O. Tadauchi, ELKU).

RUSSIA: 1 c?, Primorsky Cray, 20 km SE of Spassk forest (label in Russian), 17 Jul. 1995 (Belokobylsky

leg., ZISP).

Additional description

LABRUM(Fig. 1C-D). Basal area approximately 2.0 x as wide as long in female; basal elevation moderately

developed in both sexes, depressed centrally in male; distal process of female slender, nearly as long

as basal area, and without lateral projection, that of male absent; keel of distal process narrow, apically

pointed; labral fimbria acutely pointed at apex in both sexes. S7-S8 (Fig. 2E): S7 with moderately long,

apex exceeding S8; S8 without median process.

Male genitalia (Fig. 2A-D). Gonobase flat at bottom, ventral arms connected with each other at upper

ends; gonocoxite smooth, gently sloped in lateral view, inner and outer dorsal margins nearly parallel;

gonostylus truncate apically in lateral view, located at top of gonocoxite, with sparse short hairs; ventral

retrorse lobe long, reaching gonobase, with long and dense stetae apically (remaining parts with sparse

short hairs) and distinct lineolation; penis valve higher than gonocoxite, without cleft on top.

Distribution

Russian Far East (Siberia, Khabarovsk, and Primorsky), China (Heilongjiang Prov., Shanghai), the

Korean Peninsula (north, new record for south).

Flight records

Female: April to September.

Male: July to September.

Flower records

In South Korea, it was collected on the flowers of Potentilla hebiichigo Yonek. & H. Ohashi (Rosaceae)

and Brassica sp. (Brassicaceae).

Comments

Male specimens from Primorsky and South Korea are studied in the present paper, including characters

of the labrum and genitalia. Through this examination, we have concluded that the males of both this

species and L. pseudannulipes cannot be clearly separated. A female specimen of this species examined

in this study shows weak lineolation on T1 basally (female T1 basally usually with strong lineolation

as in Fig. 25A). The character state of this female specimen overlaps with some female specimens

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

of L. pseudannulipes (see the description of Murao et al. 2009a). Lasioglossum viridellum and L.

pseudannulipes could possibly represent the same species. It is difficult to resolve this, because we

could not examine enough material of L. viridellum in this study. This problem needs to be addressed by

including DNA analysis in a future study.

Lasioglossum ( Dialictus ) meakanense Murao & Tadauchi, 2009

Figs 3A, 26A, 28B

Lasioglossum (. Evylaeus ) meakanensis Murao & Tadauchi, 2009a (erroneous spelling): 113-117

(holotype: ELKU, S', type locality: Hokkaido, Japan).

Diagnosis

This species is separated from related species by the combination of the following characters: female

supraclypeus with sparse PP (IS = 1—4.7), male labrum without basal elevation, and male S4-S5

(particularly S5) laterally with hair tufts that are longer than the surrounding hairs.

Distribution

Japan (Hokkaido). This species has only been collected from Mt. Meakan-dake in the eastern part of

Hokkaido.

Flight records

Both sexes: August.

Flower records

Campanula lasiocarpa Cham. (Campanulaceae).

Fig. 3. Distribution maps. A. Lasioglossum ( Dialictus ) pseudannulipes (Bluthgen, 1925) and L. (D.)

meakanense Murao & Tadauchi, 2009 in Japan (•: L. pseudannulipes. u: L. meakanense). B. Lasio¬

glossum ( Dialictus ) viridellum (Cockerell, 1931) in the Korean Peninsula.

European Journal of Taxonomy 137: 1-50 (2015)

The atroglaucum group

Species included:

L. alishanense Murao sp. nov.

L. atroglaucum (Strand, 1913)

*L. callophrys Ebmer, 2002

*L. kintonense (Bliithgen, 1926)

*L. mystaphium Ebmer, 2002

L. negoroi Murao & Tadauchi, 2008

L. problematicum (Bliithgen, 1923)

L. taiwanense Murao sp. nov.

L. sanitarium (Bliithgen, 1926)

*L. sauterum Fan & Ebmer, 1992

L. sichuanense Fan & Ebmer, 1992

*L. subversicolum Fan & Ebmer, 1992

*L. versicolum Fan & Ebmer, 1992

L. virideglaucum Ebmer & Sakagami, 1994

L. yamanei Murao, Ebmer & Tadauchi, 2006

The members of this group marked with an asterisk are tentatively included because the male has not yet

been described. Seven species in total are known from the study area: three from Japan, one from Japan

and the Korean Peninsula, and three from Taiwan.

Diagnosis

Species of the atroglaucum group may be strictly characterized by a combination of the following

character states: 1) male F2 approximately 1.7-2.3 x as long as FI; 2) mesepisternum punctate over

entire surface in both sexes (e.g., Fig. 6E); 3) male S4 or S5 modified, or with unique hair tufts as in

Figs 4, 13-14; and 4) penis valve of male genitalia with high or low cleft on top (Figs 7C, 9C, 11C, 16C,

18C). The diagnostic characters of this group are also shown in Ebmer (2002) and Pesenko (2007a, as

subgenus Glauchalictus Pesenko, 2007).

In addition, this group can be classified into two subgroups. The atroglaucum subgroup (L. atroglaucum ,

L. negoroi , L. taiwanense sp. nov., L. virideglaucum , and L. yamanei) is characterized by a combination of

the male S5 with apical depression and without brush-like hairs (Fig. 5B, E, H, K, N), and the gonostylus

of male genitalia without setae on the inner surface. The problematicum subgroup ( L. alishanense sp.

nov., L. problematicum , L. sanitarium , and L. sichuanense ) is characterized by the male S5 with brush¬

like hairs along apical margin (Figs 13, 14B, E, H) and the gonostylus of male genitalia with dense short

setae on the inner surface (Fig. 16D). The latter subgroup can be regarded as a monophyletic group by

the characters of the male S5 and genitalia. These character states are unique in the subgenus and are

considered to be autapomorphies.

Distribution

This group is distributed from the Far East to southern Asia. It is diverse in the northern Oriental Region.

Comments

Lasioglossum miyabei Murao, Ebmer & Tadauchi, 2006, L. pronotale Ebmer, 2002, and L. xizangense

Fan & Ebmer, 1992 are included as members of the atroglaucum group sensu Ebmer (2002) and Murao

et al. (2006). These three species, however, do not share character states 3) and 4). They are considered

to constitute another, separate group of its own.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

The atroglaucum subgroup

Lasioglossum ( Dialictus ) atroglaucum (Strand, 1913)

Figs 4A, 5A-C, 6, 7, 12A

Halictus atroglaucus Strand, 1913: 170 (holotype: SDEI, type locality: Suisharyo, Formosa).

Halictus atroglaucus - Hirashima 1957: 5.

Lasioglossum ( Evylaeus) atroglaucum - Ebmer 2002: 860-861.

Diagnosis

This species is easily separated from other members of the atroglaucum subgroup by the male S4 having

dense and curly plumose hairs (Fig. 4A) and by the shape of the gonostylus (Fig. 1C).

Material examined

Holotype

TAIWAN: $ (Code No. DEIGISHyml0517), Suisharyo, 11 Oct. (Sauterleg., SDEI).

Additional material

TAIWAN: 1 Ya Kou, alt. 2800 m, Kao Hsiung Hsien, S-Taiwan, 1 Aug. 1986 (K. Baba, ELKU);

3 SS, Yu Shih, (alt. 1900 m), Nao Tow Hsien, M-Taiwan, 4 Jul. 1986 (K. Baba, ELKU); 1 S, Hehuanxi,

23 Jun. 1976 (H. Makihara, ELKU).

Measurements (n= 5, unit mm )

BL = 5.57-7.29 (6.17 ±0.66), WL = 4.86-5.29 (5.06 ±0.22), HL = 1.29-1.42 (1.37 ±0.05), HW = 1.35-

1.39 (1.38 ±0.01), IOD = 0.24-0.29 (0.26 ±0.02), OOD = 0.23-0.26 (0.24 ±0.01), OCD = 0.16-0.19

(0.18 ±0.01, n= 4), UOD = 0.84-0.87 (0.85 ±0.02), MOD = 0.97-1.00 (0.99 ±0.02), LOD = 0.50-0.58

(0.55 ±0.03), IAD = 0.19-0.23 (0.21 ±0.01), AOD = 0.16-0.18 (0.16 ±0.01), CAL = 0.23-0.27 (0.25

±0.02), CPL = 0.34-0.37 (0.35 ±0.01), EL = 1.00-1.10 (1.06 ±0.04), EW = 0.45-0.50 (0.49 ±0.02), GW

= 0.25-0.35 (0.30 ±0.04), SPL = 0.31-0.34 (0.32 ±0.01), F1L = 0.13-0.16 (0.14 ±0.01), F2L = 0.27-

0.29 (0.28 ±0.01), F3L = 0.27-0.32 (0.31 ±0.02), F2W = 0.13-0.16 (0.15 ±0.01), MsW = 1.50-1.60

(1.55 ±0.06, n= 4), SCL = 0.34-0.38 (0.36 ±0.02), MNL = 0.18-0.23 (0.20 ±0.02), MPL = 0.30-0.34

(0.33 ±0.02), MtW = 1.00-1.15 (1.07 ±0.07).

Redescription

Male

Coloration. Body black except for the following parts: head dark green and mesosoma metallic green;

clypeus yellow on lower half; mandible reddish brown apically; flagellum blackish brown ventrally;

tegula blackish brown translucent; tibial spur yellow; metasomal terga narrowly brown translucent

apically. Wings transparent, veins and stigma brown.

Pubescence. Body hairs whitish, and covered with erect and sparse fine, branched hairs except for the

following parts: lower paraocular area with sparse tomentose; lower clypeus with sparse simple hairs;

tibial and tarsal hairs nearly simple; disc of metasomal terga with sparse and simple short hairs; disc

of S2-S3 apically with moderately dense fine, branched hairs; disc of S4 apically with dense and curly

plumose hairs (Fig. 4A); disc of S5 with sparse short and simple hairs, mixed with sparse fine, branched

hairs laterally; disc of S6 with sparse short and simple hairs.

Head. As long as wide; HW:HL = 1:0.99. Vertex rounded in frontal view. MOD: UOD UOD =1:0.86:0.56.

IOD:OOD:OCD = 1:0.90:0.67. IAD:AOD = 1:0.76. Ocellocular and paraocular areas and frons dull,

European Journal of Taxonomy 137: 1-50 (2015)

with reticulate PP. Supraclypeal area weakly convex, dull, with sparse PP, IS distinctly tessellate (IS

= 1—4). CPL:CAL = 1:0.72. Clypeus nearly flat, with sparse PP, IS weakly tessellate (IS = 1-4.5).

EW:GW = 1:0.61. Genal area with straight ridges and distinct tessellation. Malar space linear. Occiput

not carinate. Postgena wealdy tessellate. Mandible edentate. Labrum (Fig. 6C) without basal elevation

and distal process. Antenna long, reaching metasoma. F2L:F2W = 1:0.54; flagellum nearly flattened

ventrally.

Thorax. Dorsolateral angle of pronotum obtuse; lateral surface without ridges; lateral lobe rounded.

Tegula ovoid, nearly smooth. Mesoscutum (Fig. 6D) with moderately dense PP on marginal area.

Fig. 4. Hairs onmetasomal sterna in species of the atroglaucum subgroup, f. A. Lasioglossum (Dialictus)

atroglaucum (Strand, 1913). B. Lasioglossum (Dialictus) virideglaucum Ebmer & Sakagami, 1994.

C. Lasioglossum (Dialictus)yamanei Murao etal ., 2006. D. Lasioglossum (Dialictus) negoroi Murao &

Tadauchi, 2008, holotype. E. Lasioglossum (Dialictus) taiwanense Murao sp. nov., holotype.

MURAO R. et al ., Dialictus of Japan, the Korean Peninsula, and Taiwan

Fig. 5. S4-S6 in species of the atroglaucum subgroup, S- —A-C. Lasioglossum ( Dialictus) atroglaucum

(Strand, 1913). — D-F. Lasioglossum (. Dialictus) virideglaucum Ebmer & Sakagami, 1994. — G-I.

Lasioglossum (Dialictus) yamanei Murao et al., 2006 — J-L. Lasioglossum (Dialictus) negoroi Murao

& Tadauchi, 2008, paratype. — M-O. Lasioglossum (Dialictus) taiwanense Murao sp. nov., paratype.

A, D, G, J, M: S4. B, E, H, K, N: S5. C, F, I, L, O: S6. Dotted area on S5 shows the apical depression.

Scale bars: 0.2 mm.

European Journal of Taxonomy 137: 1-50 (2015)

gradually sparse toward anterior to medial areas; IS weakly tessellate anteriorly, but nearly smooth

medially and posteriorly (IS = 0.5-2 in marginal area, 1.5—4.5 in anterior to medial areas); parapsidal line

a narrow groove. Mesoscutellum with dense PP over entire surface, IS smooth (IS = 1-2). Metanotum

weakly rugulose. Mesepisternum (Fig. 6E) shiny, with sparse PP over entire surface, IS smooth (IS

= 1.5-6). SCL:MNL:MPL = 1:0.55:0.91. Propodeum: metapostnotum (Fig. 6F) gently inclined, with

longitudinal ridges not reaching posterior margin; junction between metapostnotum and posterior surface

Fig. 6. Lasioglossum ( Dialictus ) atroglaucum (Strand, 1913), $. A. General habitus. B. Head in frontal

view. C. Labrum. D. Mesoscutum. E. Mesepisternum. F. Metapostnotum.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

smooth, not carinate; lateral surface weakly rugulose; posterior surface with lateral carina on lower half,

without oblique carina. Fore trochanter narrow, longer than wide. Hind tibia without basitibial plate.

Hind basitarsus slender, approximately 4.6 x as long as wide. Inner hind tibial spur finely serrate. Fore

wing with three submarginal cells.

Abdomen. Discs of T1-T3 with sparse fine PP. T1 without lineolation. Lineolation on T2 basally, on

T3-T5 over entire surface. Apical margin of S4-S6 nearly straight (Fig. 5A-C). S5 with shallow apical

depression (Fig. 5B). S7-S8 (Fig. 7E): median process of S7 triangular, apex not exceeding S8; S8

without median process.

Genitalia (Fig. 7A-D). Gonobase flat at bottom, ventral arms not connected with each other at upper

ends; gonocoxite smooth, gently sloped in lateral view, inner dorsal margin angulate at the approximately

basal half; gonostylus small and thin, bud-like in lateral view, located on top of gonocoxite, and with

Fig. 7. Lasioglossum ( Dialictus) atroglaucum (Strand, 1913), S- A. Genitalia in ventral view. B. Geni¬

talia in dorsal view. C. Genitalia in lateral view. D. Ventral retrorse lobe of genitalia. E. S7-S8. Scale

bars: A-C, E = 0.2 mm, D = 0.1 mm.

European Journal of Taxonomy 137: 1-50 (2015)

sparse short and a few relatively long hairs; ventral retrorse lobe tongue-like, moderately long but not

reaching gonobase, with dense short setae; penis valve higher than gonocoxite, with low cleft on top.

Female

Unknown.

Distribution

Taiwan.

Flight period

Male: late June to early August.

Flower records

Not recorded.

Lasioglossum ( Dialictus ) virideglaucum Ebmer & Sakagami, 1994

Figs 4B, 5D-F, 8, 9, 12B-C, 24B

Lasioglossum (. Evylaeus) virideglaucum Ebmer & Sakagami in Ebmer et al. , 1994: 27-31 (holotype:

Andreas Werner Ebmer’s collection, Linz, Austria, 3', type locality: Heishu, Yunnan Prov., China).

Lasioglossum {Evylaeus) virideglaucum - Ebmer 2002: 864-865. — Ebmer 2006: 564. — Murao et al.

2006: 50.

Evylaeus ( Glauchalictus ) virideglaucus - Pesenko 2007b: 85 (in key), 99 (in key), 115. — Pesenko

2007a: 26.

Diagnosis

This species is separated from the other members of the atroglaucum subgroup by a combination of the

following characters: disc of male S4 with short and moderately dense hairs (Fig. 4B); male S6 with a

pair of thin hair tufts (Fig. 4B); and the gonostylus of male genitalia nearly rounded on top (Fig. 9C).

Material examined

Paratypes

JAPAN: 2 S3, Kosugidani-Hananoego, Yaku Is., Kagoshima Pref., 24 Jul. 1952 (Takeya & Hirashima,

MNHAH).

Additional material

JAPAN: Honshu: 1 S , Kuzukawa (Hiraka), Aomori Pref., 26 Aug. 1983 (M. Yamada, MNHAH); 4 S3,

Ashiro, Iwate Pref., 19 Sep. 1976 (Y. Maeta, ELKU); 1 3, Takeshikakami-honiri, Ueda-shi, Nagano

Pref., 18 Sep. 2011 (R. Murao, cMur); 1 3, Arimine, Oyama-machi, Toyama Pref., 6 Oct. 1993 (H.

Negoro, MNHAH); 3 33, Masutomi, Yamanashi Pref., 3 Sep. 1975 (J. Emoto, ELKU). Kyushu: 1 3,

Mt. Hiko-san, Fukuoka Pref., 3 Oct. 1965 (S. Ibe, ELKU); Mt. Kujyu, alt. 1600-1700 m, Oita Pref.,

8 ??, 3 33, 19 Sep. 1972 (ELKU), 143 33, 29 Aug. 1978 (ELKU); 1 ?, Mt. Taisen, Takeda-shi,

Oita Pref., 7-9 Jun. 1978 (ELKU); 2 Matoishi, Aso-machi, Aso-gun, Kumamoto Pref., 12 Sep.

2004 (T. Sugimoto, ELKU); 1 3, Sobozan (Bungo), 9 Sep. 1933 (K. Yasumatsu, ELKU); 1 $, Mt.

Sobo, Golcasho, Talcachiho, Miyazaki Pref., 27 Apr. 2003 (O. Tadauchi, ELKU); 2 33, Kosugidani-

Hananoego, Yaku Is., Kagoshima Pref., 24 Jul. 1952 (Takeya & Hirashima, ELKU).

SOUTH KOREA: Gangwon-do: 2 33, Mt. Gariwang-san, Jeongseon-gun, 30 Jul. 2013 (R. Murao,

cMur); Bukdaesa temple, Mt. Odae-san, Jinbumyeon, Pyeongchang-gun, 3 33, 3 Sep. 1998 (H.S. Lee,

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

QIA), and 15 S3, 2 Sep. 1999 (H.S. Lee, QIA); Mt. Jeombong-san, Jindongri, Inje-gun, 4 S3, 21 Sep.

1998 (H.T. Kim, SNU), 3 33, 28 Aug. 1999 (H.T. Kim, SNU). Jeju-do: 1 3, Yonsil, Mt. Halla-san, Jeju

Is., 27 Jul. 1990 (O. Tadauchi, EFKU).

Additional description

Labrum (Fig. 8C-D). Basal area approximately 2.6 x as wide as long in female, 3 x in male; basal

elevation of female moderately developed, that of male absent; distal process of female slender, nearly

as long as basal area, and without lateral projection, that of male absent; keel of distal process narrow,

apically pointed in female; labral fimbria acutely pointed at apex in both sexes.

Sterna. S4 and S6 normal shaped, not modified. S7-S8 (Fig. 9E): S7 with long, apically rounded median

process, apex exceeding S8; S8 without median process.

Male genitalia (Fig. 9A-D). Gonobase nearly flat at bottom, ventral arms connected with each other

at upper ends; gonocoxite smooth, gently sloped in lateral view, and inner dorsal margin angulate at

approximately basal 1/3; gonostylus large, located at top of gonocoxite, with sparse short and moderately

long hairs; ventral retrorse lobe moderately long but not reaching gonobase, with moderately dense short

setae ventrally; penis valve higher than gonocoxite, with low cleft on top.

Fig. 8. Lasioglossum ( Dialictus ) virideglaucum Ebmer & Sakagami, 1994. A-B. Head in frontal

view. C-D. Fabrum. A, C. §. B, D. 3-

European Journal of Taxonomy 137: 1-50 (2015)

Distribution

Russian Far East (Primorsky, Khabrovsk), the Korean Peninsula (new record), Japan (Honshu, Shikoku,

Kyushu, Yaku Is.), China (Sichuan, Shanxi, Yunnan Provs.). While conducting our study, the male of

this species was found from several localities in South Korea (the female cannot be clearly separated

from that of some related species such as L. problematicum). On the Korean Peninsula, Lasioglossum

problematicum has been recorded from North Korea based on female specimens (Ebmer 1978). Pesenko

(2007b) regarded the continental records of L. problematicum as L. virideglaucum. Our results support

Pesenko’s opinion. It might be prudent to remove L. problematicum from the Korean fauna.

Fig. 9. Lasioglossum ( Dialictus) virideglaucum Ebmer & Sakagami, 1994, f. A. Genitalia in ventral

view. B. Genitalia in dorsal view. C. Genitalia in lateral view. D. Ventral retrorse lobe of genitalia.

E. S7-S8. Scale bars: A-C, E = 0.2 mm, D = 0.1 mm.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Flight period

Female: April to September in Kyushu, Japan.

Male: August to October.

The flight record of the female is based on specimens collected from Kyushu, Japan, because one of the

related species, L. problematicum , does not sympatrically inhabit this area.

Flower records

The specimens examined in this paper were collected on the flowers of Aster var. ovatus (Franch. & Sav.)

Soejima & Mot. Ito (Asteraceae), Beilis perennis T. (Asteraceae), Brassica napus L. (Brassicaceae),

Cimicifuga simplex (DC.) Wormsk. ex Turcz (Ranunculaceae), Isodon inflexus (Thunb.) Kudo

(Lamiaceae), Persicaria sp. (Polygonaceae), Senecio cannabifolius Tess. (Asteraceae), and Solidago

virgaurea L. subsp. asiatica (Nakai ex H. Hara) Kitam. ex H. Hara var. asiatica Nakai ex H. Hara

(Asteraceae). The other floral records in Japan belong to 12 species in 4 families (Negoro 2012).

Habitat

This species has been collected in mountainous areas in Japan and South Korea. The collecting sites are

shown in Fig. 23A, C (A = Nagano Pref., Japan; C = Mt. Gariwangsan, South Korea).

DNA barcodes

The COI gene sequences are deposited as DNA barcodes of L. ( D .) virideglaucum in the DDBJ under

accession numbers LC027537 and LC027538. These numbers are also available in GenBank.

Lasioglossum {Dialictus) yamanei Murao, Ebmer & Tadauchi, 2006

Figs 4C, 5G-I, 12D

Lasioglossum {Evylaeus) yamanei Murao, Ebmer & Tadauchi, 2006: 46—49 (holotype: ELKU, $; type

locality: Ibaraki Pref., Japan).

Evylaeus (Glauchalictus) yamanei -Pesenko 2007a: 26.

Diagnosis

This species is separated from the other members of the atroglaucum subgroup by a combination of the

following characters: disc of male S4 with sparse hairs (Fig. 4C); male S5 deeply H-shaped apically

(Figs 4C, 5H); and gonostylus of male genitalia large, obliquely truncated in lateral view (Murao et al.

2006, Fig. 11B). It may be closely related to L. taiwanense Murao sp. nov. from Taiwan in sharing the

male S5 without a groove, the apical margin of male S5 incurved, and the ventral retrorse lobe of male

genitalia short.

Material examined

JAPAN: Honshu: 1 §, Ohokawara, Aomori Pref., 28 Jun. 1992 (K. Goukon, cGou); 2 §§, Tomiya-

cho, Miyagi Pref., 19 Apr. 2009 (K. Goukon, cGou); 1 §, Nishikawa-cho, YamagataPref., 15 Jul. 2001

(K. Goukon, cGou); 2 Ogawa, Saitama Pref., 15 Sep. 1971 (T. Nambu, ELKU); 1 $, Mt. Mitake,

Tokyo Pref., 12 Sep. 1964 (H. Suda, ELKU). Shikoku: 1 <S, Monobe village, Kochi Pref., 23 Sep. 1971

(S. Ikudome, ELKU). Kyushu: 1 $, Mizunashi (Chikuzen), Fukuoka Pref., 15 Sep. 1935 (K. Yasumatsu,

ELKU); 7 $$, Mt. Wakasugi, Fukuoka Pref., 4 Nov. 1973 (Y. Yoneda, ELKU); 1 $, Mt. Hiko, Fukuoka

Pref., 16 Aug. 1973 (Y. Yoneda, ELKU); 2 §§, 1 (?, Matoishi wilderness, Aso-shi, Kumamoto Pref.,

32°55 , 46.305” N 130°58’11.555” E, 5 Aug. 2013 (Y. & R. Murao, cMur); 1 $, Mt. Sobo, Nishiusuki,

Miyazaki Pref., 29 Jun. 1985 (K. Kusumoto, ELKU).

European Journal of Taxonomy 137: 1-50 (2015)

Distribution

Japan (Honshu, Shikoku, Kyushu), China (Zhejiang Prov.).

Flight period

Female: April to October.

Male: June to November.

Flower records

Twelve species in 5 families were reported as the floral records in Japan by Goubara et al. (2004,

as Lasioglossum sp. 3), 15 species in 9 families by Murao et al. (2006), and 9 species in 7 families

by Negoro (2012). The total number of species is 36 in 13 families as follows. Apiaceae: Angelica

polymorpha Maxim.; A. spp.; Osmorhiza aristata (Thunb.) Rydb. Asteraceae: Aster ageratoides Turcz.

var. ageratoides ; A. glehnii F.Schmidt var. glehnii ; Erigeron annuus (L.) Pers.; Ixeridium dentatum

(Thunb.) Tzvelev subsp. dentatum ; Lactuca indica L.; Solidago virgaurea T. subsp. asiatica (Nakai

ex H. Hara) Kitam. ex H. Hara var. asiatica Nakai ex H. Hara; Solidago virgaurea L. subsp. leiocarpa

(Benth.) Hulten; Taraxacum officinale Weber ex F.H. Wigg. Brassicaceae: Brassica rapa T. var. oleifera

DC.; Capsella bursa-pastoris (L.) Medik.; Cardamine leucantha (Tausch) O.E. Schulz; Eutrema tenue

(Miq.) Makino. Caryophyllaceae: Stellaria sessiliflora Y. Yabe. Ericaceae: Elliottiapaniculata (Siebold

& Zucc.) Hook.f. f. albiflora (Y. Kimura) Yonek. Geraniaceae: Geranium sp.; G. thunbergii Siebold

ex Lindl. et Paxton. Lamiaceae: Mosla dianthera (Buch.-Ham. ex Roxb.) Maxim. Liliaceae: Allium

thunbergii G. Don. Polygonaceae: Persicaria thunbergii (Siebold et Zucc.) H. Gross. Ranunculaceae:

Ranunculusjaponicus Thunb. Rosaceae: Geum japonicum Thunb.; Potentillafragarioides L. var. major

Maxim.; P freyniana Bornm.; P. hebiichigo Yonek. & H. Ohashi; Rubuspalmatus Thunb.; R. parvifolius

L. Salicaceae: Salix spp. Violaceae: Viola obtusa Makino; V rostrata Pursh.

Habitat

This species has been collected in mountain areas in Japan. One of the collecting sites in Japan

(Kumamoto Pref., Kyushu) is shown in Fig. 23B.

Lasioglossum ( Dialictus ) negoroi Murao & Tadauchi, 2008

Figs 4D, 5J-L, 12E

Lasioglossum (. Evylaeus) negoroi Murao & Tadauchi, 2008: 37^10 (holotype: ELKU, type locality:

Toyama Pref., Japan).

Diagnosis

This species is separated from other members of the atroglaucum subgroup by a combination of the

following characters: male S4 basally with sparse and moderately long erect hairs (Murao & Tadauchi

2008; Fig. 2H); male S5 with a linear shallow groove (Fig. 5K); male S6 with a pair of thin hair tufts;

and gonostylus of male genitalia truncated on top (Murao & Tadauchi 2008; Fig. 2B-C).

Material examined

Holotype

JAPAN: S (Code No. ELKU3261), Kobo, Tateyama, Toyama Pref., Honshu, 10 Sep. 1997 (H. Negoro,

ELKU).

Paratype

JAPAN: 1 (?, Koganeyu, Minami-ku, Sapporo-shi, Hokkaido (label in Japanese), 18 Sep. 2004 (K.

Kinota, ELKU).

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Additional material

JAPAN: Honshu: 1 3, Nikko-Yumoto (Shimozulce), 15 Aug. 1937 (T. Esaki & K. Yasumatsu, ELKU);

Mt. Zao, MiyagiPref., 1 3, 4 Sep. 1980 (K. Goukon, cGou), 1 3, 1 Sep. 1982 (K. Goukon, cGou); 1 3,

Mt. Zao, alt. 1300 m, Miyagi Pref., 28 Sep. 1996 (K. Goukon, cGou).

Female

Unknown.

Distribution

Japan (Hokkaido, northern and central Honshu).

Flight period

Male: August to October.

Flower records

Araliaceae: Aralia continentalis Kirag.. Asteraceae: Aster microcephalus (Miq.) Franch. & Sav. var.

ovatus (Franch. & Sav.) Soejima & Mot. Ito; Solidago virgaurea L.

Lasioglossum (Dialictus) taiwanense Murao sp. nov.

um:lsid:zoobank.org:act:C164E332-E591-4C50-A4BD-9E4C208276B8

Figs 4E, 5M-0, 10, 11, 12A

Diagnosis

This species is separated from other members of the atroglaucum subgroup by a combination of the

following characters: male head wider than long; disc of male S4 with sparse and short hairs (Fig. 4E);

male S5 gently incurved apically (Fig. 5N); and the shape of gonostylus of male genitalia (Fig. 11C-D).

Etymology

The specific name is derived from the type country, Taiwan.

Material examined

Holotype

TAIWAN: 3, Hehuanxi, Songguangang, Nantou Pref., 24 Jun. 1976 (H. Makihara, EFKU).

Paratypes

TAIWAN: 4 33, Meishan-Tinchi (alt. 800-2300 m), Kao Hsiung Hsien, S-Taiwan, 29 Jun. 1986 (K.

Baba, EFKU); 2 33, Pi Fu Chih, alt. 2500 m, Nan Tow Hsien, M-Taiwan, 5 Aug. 1986 (K. Baba,

EFKU); 1 3, Tung Chih, alt. 1600 m, near Fiu Km, S-Taiwan, 30 Sep. 1986 (K. Baba, EFKU); 8 33,

Hehuanxi, Songguangang, Nantou Pref., 24 Jun. 1976 (H. Makihara, EFKU); 3 33, Hehuanxi, 23 Jun.

1976 (H. Makihara, EFKU); 2 33, Sung Kang, alt. 2000-2300 m, Nantou Hsien, 29 Jun. 1965 (S.

Kimoto, EFKU).

Measurements (n = 10, unit mm )

BE = 5.29-6.00 (5.65 ±0.32), WE = 4.43-5.14 (4.91 ±0.25), HE = 1.26-1.40 (1.32 ±0.06), HW =

1.32-1.45 (1.37 ±0.05), IOD = 0.19-0.26 (0.24 ±0.02, n= 9), OOD = 0.24-0.28 (0.27 ±0.01), OCD =

0.16-0.19 (0.18 ±0.02, n= 7), UOD = 0.84-0.90 (0.87 ±0.03), MOD = 0.92-1.03 (0.97 ±0.04), FOD =

0.55-0.61 (0.59 ±0.02), IAD = 0.23-0.26 (0.24 ±0.01), AOD = 0.16-0.19 (0.17 ±0.01), CAE = 0.19-

0.28 (0.25 ±0.02), CPF = 0.31-0.35 (0.33 ±0.01), EE = 0.95-1.05 (1.00 ±0.04), EW = 0.45-0.50 (0.47

±0.02), GW = 0.20-0.35 (0.28 ±0.04), SPL = 0.29-0.32 (0.31 ±0.01), F1L = 0.13-0.15 (0.14 ±0.01),

European Journal of Taxonomy 137: 1-50 (2015)

F2L = 0.23-0.27 (0.24 ±0.01), F3L = 0.24-0.27 (0.26 ±0.01), F2W = 0.15-0.16 (0.15 ±0.01, n= 9),

MsW = 1.30-1.50 (1.44 ±0.07), SCL = 0.30-0.35 (0.33 ±0.02), MNL = 0.15-0.20 (0.17 ±0.02), MPL =

0.28-0.33 (0.31 ±0.02), MtW= 1.00-1.18 (1.07 ±0.06).

Fig. 10. Lasioglossum ( Dialictus) taiwanense Murao sp. nov., $. A. Lateral habitus. B. Head in frontal

view. C. Labrum. D. Mesoscutum. E. Mesepisternum. F. Metapostnotum. A-B, E-F. Holotype. C-D.

Paratype.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Description

Male

Coloration. Body black except for the following parts: head dark green and mesosoma metallic green;

clypeus yellow on lower half or lower 1/3; mandible reddish brown apically; flagellum blackish brown

or brown ventrally; tegula blackish brown translucent; tibial spur yellow; metasomal terga narrowly

brown translucent apically. Wings transparent, veins and stigma brown.

Pubescence. Body hairs whitish, and covered with erect and sparse fine, branched hairs except for the

following parts: lower clypeus with sparse simple hairs; tibial and tarsal hairs nearly simple; disc of

metasomal terga with sparse and simple short hairs. Metasomal sterna with sparse and short hairs,

without special hair tufts.

Head. Wider than long; HW:HL = 1:0.96. Vertex rounded in frontal view. MOD:UOD:LOD =

1:0.90:0.61. IOD:OOD:OCD = 1:1.11:0.76. IAD:AOD = 1:0.71. Ocellocular areas, paraocular areas

and frons dull, with reticulate PP. Supraclypeal area weakly convex, dull, with moderately dense PP,

Fig. 11. Lasioglossum (. Dialictus ) taiwanense Murao sp. nov., paratype, $. A. Genitalia in ventral view.

B. Genitalia in dorsal view. C. Genitalia in lateral view. D. Gonostylus in dorsal view. E. Ventral retrorse

lobe of genitalia. F. S7-S8. Scale bars: A-C, F = 0.2 mm, D-E = 0.1 mm.

European Journal of Taxonomy 137: 1-50 (2015)

IS distinctly tessellate (IS = 0.5-2). CPL:CAL = 1:0.77. Clypeus nearly flat, its sculptures similar to

supraclypeal area, IS = 1-3.5. EW:GW = 1:0.59. Genal area with straight ridges and distinct tessellation.

Malar space linear. Occiput not carinate. Postgena weakly tessellate. Mandible edentate. Labrum (Fig.

10C) without basal elevation and distal process. Antenna long, reaching metasoma. F2L:F2W = 1:0.63;

flagellum nearly flattened on lower side.

Thorax. Dorsolateral angle of pronotum obtuse; lateral surface without ridges; lateral lobe rounded.

Mesoscutum (Fig. 10D) with dense PP on marginal area, gradually sparse anterior to medial areas; IS

weakly tessellate (IS = 0.5-1.5 on marginal area, IS= 0.5-3.7 on anterior to medial areas); parapsidal

line a narrow groove. Mesoscutellum with dense PP (IS = 0.5-1.5) over entire surface, IS smooth.

Metanotum weakly rugulose. Mesepisternum (Fig. 10E) with sparse PP over entire surface, IS smooth

(IS = 1-6). SCF:MNF:MPF = 1:0.53:0.95. Propodeum: metapostnotum (Fig. 10F) gently inclined,

with longitudinal ridges on basal half, with distinct tessellation on apical half; the junction between

metapostnotum and posterior surface weakly tessellate, not carinate; lateral surface weakly rugulose and

distinctly tessellate; posterior surface with lateral carina on lower half. Fore trochanter narrow, longer

than wide. Hind tibia without basitibial plate. Hind basitarsus slender, approximately 4.7 x as long as

wide. Inner hind tibial spur finely serrate.

Abdomen. Discs of T1-T3 with sparse fine PP. T1 smooth except for punctures. T2 basally and apically

lineolate, T3-T5 over entire surface. S4 and S6 normal shaped, not modified. S5 gently incurved apically

(Fig. 5N). S7-S8 (Fig. 1 IF): S7 with triangular median process, apex not exceeding S8; S8 medially

weakly projecting, but not forming median process.

Genitalia (Fig. 11A-E). Gonobase short, flat at bottom, ventral arms connected with each other at upper

ends; gonocoxite smooth, gently sloped in lateral view, and inner dorsal margin angulate approximately

at basal Vy gonostylus small, truncated apically in lateral view, located on top of gonocoxite, and with

sparse short and a few long hairs; ventral retrorse lobe leaf-shaped, short, with moderately dense short

setae ventrally; penis valve with low cleft on top.

Female

Unknown.

Variation

The postgena nearly smooth in five paratypes (holotype and rest of paratypes weakly tessellate).

Distribution

Taiwan.

Flight period

Male: June to September.

Flower records

Not recorded.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

The problematicum subgroup

Lasioglossum {Dialictus) problematicum (Bluthgen, 1923)

Figs 13A, 14A-C, 15, 16, 19A, 27B

Halictus problematicus Bluthgen, 1923: 331-332 (lectotype: Institute of Systematic and Experimental

Zoology, Polish Academy of Sciences, Krakow, Poland, $; type locality: unknown (Siberia?); designated

by Pesenko, 2007b: 115).

Halictus problematicus - Hirashima 1957: 15.

Lasioglossum ( Evylaeus ) problematicum - Ebmer 1978: 311-312. — Ebmer 1996: 282; Ebmer 2002:

865-867 (c?). — Ebmer 2006: 563. — Murao et al. 2006: 50.

Evylaeus ( Glauchalictus ) problematicus - Pesenko 2007a: 26. — Pesenko 2007b: 85 (in key), 95 (in

key), 115.

Fig. 12. Distribution maps. A. Lasioglossum (. Dialictus ) atroglaucum (Strand, 1913) and L. (D.)

taiwanense Murao sp. nov. B-C. Lasioglossum ( Dialictus ) virideglaucum Ebmer & Sakagami, 1994

(B in the Korean Peninsula, C in Japan). D. Lasioglossum {Dialictus) yamanei Murao et al., 2006.

E. Lasioglossum ( Dialictus ) negoroi Murao & Tadauchi, 2008.

European Journal of Taxonomy 137 : 1-50 ( 2015 )

Diagnosis

This species is closely related to Lasioglossum alishanense Murao sp. nov., L. sanitarium (Bliithgen,

1926) and L. sichuanense Fan & Ebmer, 1992 from the northern Oriental Region, as stated above.

For the differences between this species and L. alishanense , see the Key. It is also separated from the

remaining two species by having the disc of the male S5 without an apical depression (Fig. 14B). The

female cannot be clearly separated from the related species, as stated above.

Material examined

JAPAN: Hokkaido: 3 $$, Rausu-seseki, Shiretoko, 16 Jun. 1984 (O. Tadauchi, ELKU); 3 33, Rebun

Is., Rebun-no-oka, 7-8 Sep. 2005 (K. Goukon, cGou); 27 S3, Kabuka, Rebun Is., 19 Aug. 1984 (O.

Tadauchi, ELKU); 1 3, Air Port Wakkanai, 16 Aug. 1984 (O. Tadauchi, ELKU); 2 33, Botanical

Garden, Sapporo, 3 Aug. 1959 (S. F. Sakagami & H. Fukuda, MNHAH); 1 3, Hamakoshimizu, 9 Aug.

1967 (MNHAN); 10 33, Kami-Otoineppu, Nakagawa Exp. For., 14 Sep. 1969 (S. F. Sakagami & H.

Fukuda, MNHAH); 1 3, Kitamoshiri, Hole. Uryu Exp. For., 14 Sep. 1969 (S. F. Sakagami & H. Fukuda,

MNHAH); 2 33, Kussharo Lake, Teshikaga, 21 Jul. 1984 (O. Tadauchi, ELKU), 30 Aug. 1984 (O.

Tadauchi, ELKU); 2 33, Nishiashoro, 9-10 Aug. 1955 (S. F. Sakagami & H. Fukuda, MNHAH); 1 3,

Nissho Pass, Hidaka, 22 Sep. 1984 (O. Tadauchi, ELKU); 1 3, Cape Nosappu, Nemuro, 13 Sep. 1984

(O. Tadauchi, ELKU); 1 3, Nukabira, 10 Aug. 1965 (Y. Hirashima, ELKU); Sapporo, 1 3, 17 Aug.

1958, 4 33, 8 Sep. 1958 (S. Ueda, ELKU); 1 3, Shari Pass, Shari, 28 Jul. 1984 (O. Tadauchi, ELKU);

1 3, Shimamaki, Shiribetsu, 2 Sep. 1984 (O. Tadauchi, ELKU); 6 33, Shotoshibetsu, 20 Jul. 1984, 9 &

Fig. 13. Hairs on metasomal sterna in species of the problematicum subgroup, 3 ■ A. Lasioglossum

(. Dialictus ) problematicum (Bliithgen, 1923). B. Lasioglossum ( Dialictus ) sanitarium (Bliithgen, 1926).

C. Lasioglossum ( Dialictus ) sichuanense Fan & Ebmer, 1992. D. Lasioglossum (. Dialictus ) alishanense

Murao sp. nov., holotype.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

26 Aug. 1984 (O. Tadauchi, ELKU); 29 33, Sugatami, 6 Sep. 1967 (MNHAH); 2 S3, Tenneru, north

Kushiro, 1968 (E. Ohtsuka, MNHAH); 4 S3, Tokachimitsumata, Kamishihoro, 27 Aug. 1984, 11 Sep.

1984 (O. Tadauchi, ELKU); 6 33, Rubetu, Etorofii Is., 31 Aug. 1940 (S. Kuwayama & Y. Sugihara,

ELKU); 13 S3, Sibetoru, Etorofii Is., 6-7 Sep. 1940 (S. Kuwayama & Y. Sugihara, ELKU). Honshu:

4 33, Kuzukawa (Hiraka), Aomori Pref., 20 & 26 Aug. 1983 (M. Yamada, MNHAH); 1 3, Minami-

Hakkoda (Hiraka), Aomori Pref., 17 Aug. 1984 (M. Yamada, MNHAH); 6 33, Mt. Iwaki, Aomori

Pref., 7 & 21 Sep. 1980, 29 Aug. 1981 (M. Yamada, MNHAH); 4 33, Sulcayu (Towadako), Aomori

Pref., 2 Aug. 1987 (M. Yamada, MNHAH); 1 3, Takinosawa, Aomori Pref., 4 Sep. 1982 (M. Yamada,

MNHAH); 1 3, Hachimantai, Akita Pref., 28 Jul. 1961 (M. Shiga, ELKU); 6 33, Mt. Akitakoma, Akita

Pref., 28 Aug. 1972 (M. Honda, ELKU); 1 3, Mt. Chokai-san, Akita Pref., 27 Aug. 1972 (M. Honda,

ELKU); 1 3, Fukenoyu spa, Kazono city, Akita Pref., 2 Aug. 1974 (T. & H. Suda, ELKU); 2 33, Mt.

Zao, Shibalcusadaira, Miyagi Pref., 27 Aug. 2004 (K. Goulcon, cGou); 1 3, Mt. Zao, alt. 1300 m, Miyagi

Pref., 28 Sep. 1996 (K. Goulcon, cGou); 2 33, Mt. Zao, Miyagi Pref., 31 Aug. 1979 (K. Goulcon,

cGou), 24 Aug. 1979 (Y. Maeta, ELKU); 15 33, Kazawa, Nagano Pref., 17 Aug. 1972 (Y. Yoshiyasu,

ELKU); 2 33, Shiga Highlands, Nagano Pref., 16 Aug. 1972 (Y. Yoshiyasu, ELKU); 2 33, Sugadaira

Highlands, Sanada-machi, Nagano Pref., 22 Aug. 2002 (K. Mitai, cMur); 2 33, Asama, Karuizawa,

Nagano Pref., 23 Aug. 1967 (T. & H. Suda, ELKU); 1 3, Shirakaba Lake, Chino city, Nagano Pref., 7

Sep. 1966 (T. & H. Suda, ELKU); 1 3, Arimine, Oyama-machi, Toyama Pref., 11 Aug. 1994 (H. Negoro,

Fig. 14. S4-S6 in species of the problematicum subgroup, 3 ■ A—C. Lasioglossum (. Dialictus )

problematicum (Bliithgen, 1923). D-F. Lasioglossum (Dialictus) sanitarium (Bliithgen, 1926).

G—I. Lasioglossum ( Dialictus) alishanense Murao sp. nov., paratype. A, D, G: S4. B, E, H: S5. C, F, I.

S6: Dotted area on S5 shows the apical depression. Scale bars: 0.2 mm.

European Journal of Taxonomy 137: 1-50 (2015)

MNHAH); 39 SS, Migatahara, Mt. Tate-yama, Toyama Pref., 25 Aug. 2004 (T. Sugimoto, cMur); 1

Mt. Daicho, Katsuyama-shi, Toyama Pref., 27 Aug. 2004 (T. Sugimoto, cMur); 1 S, Sen-ninmori, near

Mt. Ikenodaira, north Alps, Toyama Pref., 19 Aug. 2005 (H. Hirano, cMur); 1 Tateyama-Migatahara,

Toyama Pref., 16 Oct. 1995 (H. Negoro, MNHAH); 2 $$, Mt. Haku-san, Ishikawa Pref., 6 Aug. 1970

(K. Kanmiya, ELKU), 9 Aug. 1974 (I. Togashi, ELKU); 1 Mt. Haku-san, alt. 1750-1500 m, Ishikawa

Pref., 22 Aug. 1988 (I. Togashi, ELKU).

Additional description

Labrum (Fig. 15C-D). Basal area approximately 2.3 x as wide as long in female, 3 x in male; basal

elevation of female moderately developed, that of male absent; distal process of female slender, nearly

as long as basal area, and without lateral projection, that of male absent; keel of distal process narrow,

apically pointed; labral fimbria acutely pointed at apex in both sexes.

Sterna. S4 medially with shallow groove on apical half; apical margin nearly straight. S5 without apical

depression, medially weakly swollen and apico-medially cleaved; apical margin gently incurved. S6

normally shaped, not modified. S7-S8 (Fig. 16F): S7 with elongate, apically rounded median process,

apex exceeding S8; S8 without median process.

Male genitalia (Fig. 16A-E). Gonobase nearly flat at bottom, ventral arms connected with each other

at upper ends; gonocoxite smooth, gently sloped in lateral view, and inner dorsal margin gently angulate

Fig. 15. Lasioglossum ( Dialictus ) problematicum (Bliithgen, 1923). A-B. Head. C-D. Labrum. A,

C. ?.B, D.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

at approximately basal 1/3; gonostylus large, located at top of gonocoxite, with dense short setae on

inner surface; ventral retrorse lobe moderately long but not reaching gonobase, with dense short setae

ventrally and relatively long blunt setae laterally; penis valve higher than gonocoxite, with high cleft on

top.

Distribution

Russian Far East (Sakhalin) and Japan (Hokkaido, northern to central Honshu). Ebmer (1978, 2006)

recorded this species from Primorsky and North Korea, respectively, based on female specimens. These

records are not included in this paper, because the female cannot be clearly separated from that of some

of the related species.

Fig. 16. Lasioglossum (. Dialictus ) problematicum (Bluthgen, 1923), $. A. Genitalia in ventral view.

B. Genitalia in dorsal view. C. Genitalia in lateral view. D. Gonostylus in inner-lateral view. E. Ventral

retrorse lobe of genitalia. F. S7-S8. Scale bars: A-C, F = 0.2 mm, D-E = 0.1 mm.

European Journal of Taxonomy 137: 1-50 (2015)

Flight period

Female: May to September in Hokkaido, Japan.

Male: August to October.

The flight record of females is based on specimens collected from the eastern part of Hokkaido, Japan.

This record can be considered accurate, because one of the related species, L. virideglaucum Ebmer &

Salcagami, does not sympatrically inhabit this area.

Flower records

In Japan this species visits the following 19 species in 6 families: Apiaceae: Angelica ursina (Rupr.)

Maxim. AstQracQSLQ: Anaphalismargaritacea(L.)Benth. &Hook. f. subsp. margaritacea;AsterglehniiV.

Schmidt; Cirsium setosum (Willd.) M. Bieb.; Crepidiastrum denticulatum (Houtt.) J.H. Pak & Kawano;

Erigeron annuus (L.) Pers.; Hieracium umbellatum L.; Picris hieracioides subsp. japonica (Thunb.)

Krylov; Rudbeckia laciniata L.; Senecio pseudoarnica Less.; Solidago altissima L S. virgaurea L.

subsp. asiatica (Nakai ex H. Hara) Kitam. ex H. Hara var. asiatica Nakai ex H. Hara; S. virgaulea

L. subsp. leiocarpa (Benth.) Hulten; Sonchus brachyotus DC.; Taraxacum sp. Brassicaceae: Barbarea

orthoceras Ledeb. Geraniaceae: Geranium thunbergii Siebold ex Lindl. & Paxton. Lamiaceae: Isodon

sp. Rosaceae: Sorbaria sorbifolia (L.) A. Braun.

Biological reference

According to Salcagami & Kuribayashi (1979) and Sakagami et al. (1984), some biological information

is available. Nest place and structure: the female prefers to make a nest at the forest edge; the nest

structure is the Ilia type of Sakagami & Michener (1962). Social structure and life cycle: the brooding

period of the overwintered generation is from early May to late July, in Sapporo city, Hokkaido; most

of the overwintering females stay communally in the natal nests; the nest are communally reused in

the next brooding season, hence many colonies are semisocial or sometimes delayed eusocial from the

beginning of spring activities; females living in the same nests are mostly of the same generation and

seem to be sisters in most cases, although some nests are inhabited by a two-year-old female and her

daughters; all cohabiting females are inseminated, but only one of them has fully developed ovaries at

the peak of brooding season, the others with undeveloped ovaries participate in foraging.

Lasioglossum ( Dialictus ) alishanense Murao sp. nov.

urn:lsid:zoobank.org:act:34A726F0-E3B3-4E3F-B6B8-lllE08F19BD2

Figs. 13D, 14G-I, 17, 18, 19B

Diagnosis

This species may be closely related to Lasioglossum problematicum , L. sanitarium , and L. sichuanense,

as stated above. For the differences between this species and L. problematicum , see the Key. It is

separated from the two other related species by the disc of the male S4 having short and sparse hairs

(Fig. 13D), the male S5 without an apical depression (Fig. 14H), and the shape of gonostylus of male

genitalia (Fig. 18C).

Etymology

The specific name is derived from the type locality, Mt. Alishan in Taiwan.

Material examined

Holotype

TAIWAN: Chiayi-Hsien Alishan, 27 Jul. 1966 (T. Tano, ELKU).

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Paratypes

TAIWAN: 1 3, Taiheizan (Taihoku-shu), Toganoo-Minamotojobu-Kussha, 22 Jul. 1932 (T. Esaki,

ELKU); 1 3, Chiayi-Hsien Shihtzulu, 29 Jul. 1966 (T. Tano, ELKU); 3 33, Chiayi-Hsien Alishan, 27

Jul. 1966 (T. Tano, ELKU); 1 3, Fenchifu, alt. 1405 m, 1 Sep. 1979 (Y. Hirashima, ELKU).

Measurements (n = 6, unit mm)

BL = 5.86-6.86 (6.38 ±0.43), WL = 5.43-6.00 (5.79 ±0.22), HL = 1.39-1.58 (1.51 ±0.07), HW = 1.42-

1.58 (1.52 ±0.06), IOD = 0.23-0.24 (0.23 ±0.01), OOD = 0.27-0.35 (0.31 ±0.03), OCD = 0.23-0.24

(0.24 ±0.01), UOD = 0.90-1.03 (0.98 ±0.05), MOD = 1.02-1.13 (1.08 ±0.05), LOD = 0.61-0.74 (0.69

±0.05), IAD = 0.24-0.31 (0.27 ±0.03), AOD = 0.16-0.21 (0.19 ±0.02), CAL = 0.24-0.27 (0.25 ±0.01),

CPL = 0.35-0.39 (0.37 ±0.02), EL = 0.98-1.10 (1.03 ±0.05), EW = 0.50-0.53 (0.50 ±0.01), GW =

0.28-0.35 (0.33 ±0.03), SPL = 0.32-0.39 (0.36 ±0.02), F1L = 0.15-0.16 (0.15 ±0.01), F2L = 0.27-0.29

(0.28 ±0.01), F3L = 0.29-0.32 (0.30 ±0.01), F2W = 0.16-0.18 (0.16 ±0.01), MsW = 1.55-1.70 (1.60

±0.06), SCL = 0.35 (0.35 ±0.00), MNL = 0.18-0.20 (0.19 ±0.01), MPL = 0.31-0.38 (0.34 ±0.03), MtW

= 1.15-1.40(1.29 ±0.09).

Description

Male

Coloration. Body black, except for the following parts: head dark green and mesosoma metallic green;

clypeus yellow on lower half; mandible reddish brown apically; flagellum blackish brown ventrally;

tegula dark brownish translucent; tibial spur yellow; metasomal terga narrowly dark brownish translucent

apically. Wings transparent, veins and stigma brown.

Pubescence. Body hairs whitish, and covered with elect and sparse fine, branched hairs except for the

following parts: lower paraocular area with sparse tomentose; lower clypeus with sparse simple hairs;

tibial and tarsal hairs nearly simple; disc of metasomal terga with sparse and simple short hairs. S2-S4

medioapically with sparse erect hairs; in addition, S4 lateroapically with sparse short hairs. S5 with

dense, brush-like, and brownish hairs along apical margin.

Head. As long as wide; HW:HL =1:0.99. Vertex rounded in frontal view. MOD: UOD UOD =1:0.90:0.64.

IOD:OOD:OCD = 1:1.33:1.01. IAD:AOD = 1:0.70. Ocellocular areas, paraocular areas and frons dull,

with reticulate PP. Supraclypeal area weakly convex, dull, with sparse PP, IS distinctly tessellate (IS =

0.5-3). CPL:CAL = 1:0.68. Clypeus nearly flat, its sculptures similar to supraclypeal area, IS = 1-4.

EW:GW = 1:0.64. Malar space linear. Occiput not carinte. Postgena nearly smooth. Mandible edentate.

Labrum (Fig. 17C) without basal elevation and distal process. Antenna long, reaching metasoma.

F2L:F2W = 1:0.57; flagellum nearly flattened ventrally.

Thorax. Dorsolateral angle of pronotum obtuse; lateral surface without ridges; lateral lobe rounded.

Mesoscutum (Fig. 17D) with dense PP on marginal area, gradually sparse toward anterior to medial areas;

IS weakly tessellate (IS = 0.5-1.5 on marginal area, 0.5-2.5 on anterior to medial area); parapsidal line

a narrow groove. Mesoscutellum with dense PP over entire surface, IS smooth (IS = 1-2). Metanotum

distinctly tessellate, not coarsely rugulose. Mesepistemum (Fig. 17E) with sparse PP over entire surface,

IS smooth (IS= 1-6.5). SCL:MNL:MPL = 1:0.54:0.98. Propodeum: metapostnotum (Fig. 17F) gently

inclined, with nearly longitudinal ridges not attaining posterior margin; junction between propodeal

dorsum and shield smooth, not carinate; lateral surface weakly rugulose and distinctly tessellate; posterior

surface weakly tessellate, with lateral carina on lower 1/3 and without oblique carina. Fore trochanter

narrow, longer than wide. Hind tibia without basitibial plate. Hind basitarsus slender, approximately 4 x

as long as wide. Inner hind tibial spur finely serrate. Fore wing with three submarginal cells.

European Journal of Taxonomy 137: 1-50 (2015)

Abdomen. Discs of T1-T3 with sparse fine PP. T1-T2 smooth except for punctures. T3- 5 with weak

lineolation over entire surface. Apical margin of S4-S5 incurved (Fig. 14G, H); shape of S5 as in L.

problematicum. S6 normally shaped, not modified. S7-S8 (Fig. 18E): S7 with narrow median process,

apex as high as top of S8; S8 without median process.

Fig. 17. Lasioglossum ( Dialictus) alishanense Murao sp. nov., $. A. Lateral habitus. B. Head in frontal

view. C. Labrum. D. Mesoscutum. E. Mesepisternum. F. Metapostnotum. A-B, E-F. Holotype. C-D.

Paratype.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Genitalia (Fig. 18A-D). Gonobase flat at bottom, ventral arms not connected with each other at

upper ends; gonocoxite smooth, gently sloped in lateral view, and inner dorsal margin gently angulate

approximately at basal 1/3; gonostylus large, similar to that of L. problematicum in shape, located on

top of gonocoxite, and with dense setae on inner surface; ventral retrorse lobe tongue-like, moderately

long but not reaching gonobase, with dense short setae ventrally and relatively long blunt setae laterally;

penis valve with high cleft on top.

Female

Unknown.

Distribution

Taiwan.

Fig. 18. Lasioglossum (. Dialictus ) alishanense Murao sp. nov., paratype, S- A. Genitalia in ventral view.

B. Genitalia in dorsal view. C. Genitalia in lateral view. D. Ventral retrorse lobe of genitalia. E. S7-S8.

Scale bars: A-C, E = 0.2 mm, D = 0.1 mm.

European Journal of Taxonomy 137: 1-50 (2015)

Flight period

Male: late July to September.

Flower record

Not recorded.

The morio group

Species included in eastern Asia:

L. briseis Ebmer, 2005

L. circe Ebmer, 1982

L. ellipticeps (Bluthgen, 1923)

L. eomontanus Ebmer, 2006

L. lambatum Fan & Ebmer, 1992

L. mows Ebmer, 2002

L. spinosum Ebmer, 1982

Only one species, listed below, is known to occur on the Korean Peninsula.

Diagnosis and comments

This group in the present paper corresponds to Evylaeus (Smeathhalictus) Warncke sensu Pesenko

(2007a). According to Pesenko’s key (2007a), this group is separated from the other Palaearctic groups

by a combination of the following character states: 1) body small, body length = 4.0-6.5 mm in both

sexes, 2) head usually longer than wide in both sexes, 3) male antenna long, reaching propodeum, 4)

Fig. 19. Distribution maps. A. Lasioglossum ( Dialictus ) problematicum (Bluthgen, 1923) in Japan.

B. Lasioglossum ( Dialictus) alishanense Murao sp. nov.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

male F2 1.5-1.7 x as long as its diameter, 5) female mesocutum and mesopleuron punctate, 6) length

of metapostnotum as long as or longer than mesoscutellum in both sexes, 7) female metapostnotum

striate, more or less shiny, 8) posterior surface of propodeum usually with complete lateral carina in

both sexes, 9) male metasoma elongate, 10) female T1 apically with fine lineolate or shagreened, 11)

T2-T4 with apical fimbriae or developed lateral spots in both sexes, 12) male metasomal sterna not

modified, 13) gonostylus of male genitalia small, rounded triangular, trapeziform or elongate elliptical,

and not narrowed at base and 14) ventral retrorse lobe of male genitalia rounded, elliptical, sometimes

triangular. However, it is very difficult or impossible to separate from all other group based on these

character states. This group needs a revision of the diagnostic characters through detailed morphological

and phylogenetic studies.

Distribution

Palaearctic to northern Oriental Region. It is diverse in the western Palaearctic Region.

Lasioglossum {Dialictus) ellipticeps (Bluthgen, 1923)

Figs 20, 21, 22A, 26B

Halictus ellipticeps Bluthgen, 1923: 254 (holotype: Museum far Naturkunde, Humboldt Universitat zu

Berlin, Germany, §; type locality: Amur, Russia).

Halictus permicus Bluthgen, 1923: 330-331 (lectotype: Institute of Systematic and Experimental

Zoology, Polish Academy of Sciences, Krakow, Poland, type locality: “Perm”, designated by

Pesenko, 2007b: 117). Synonymized by Ebmer, 1978.

Halictus ( Chloralictus) mayacensis Cockerell, 1924: 582 (syntypes: U. S. National Museum of Natural

History, Smithsonian Institution, Washington, DC, USA, §; type locality: Primorsky, Russia); Synonymy

by Bluthgen 1931.

Halictus ellipticeps - Hirashima 1957: 7-8.

Lasioglossum ( Evylaeus ) ellipticeps Ebmer 1978: 310-311. —Ebmer 1982: 215-216. —Ebmer 1996:

282-283. — Ebmer 2005: 375. — Ebmer 2006: 565.

Evylaeus ( Smeathhalictus ) ellipticeps Pesenko 2007a: 28. —Pesenko 2007b: 117.

Diagnosis

This species is similar to Lasioglossum briseis Ebmer and L. eomontanus Ebmer from Far East Asia

(Pesenko 2007b). For the differences between these species, see Pesenko (2007b).

Material examined

SOUTH KOREA: Gangwon-do: 2 $ $, Doam Myon Pyongchang-gun, 20 Aug. 2001 (Y. Maeta, SULE);

1 5, Mt. Hambaek-san, 14 Aug. 1999 (S. Park, SNU). Jeollabuk-do: JeongLyongChy, SanNaeMeon,

NamWeon-gun, 14 14 May 1991 (O. Tadauchi, ELKU), 1 §, 14 Jul. 1991 (K. Kanmiya, ELKU).

Ulleung-do: 18 ?$, 1 Chusan, 11 Aug. 1999 (H. Kim, SNU); 3 ??, 2 $$, Jukdo, 12 Aug. 1999 (H.

Kim, SNU); 2 §§, Naesujeon, 12 Aug. 1999 (H. Kim, SNU); 1 §, 2 SS, Naribunji, 11 Aug. 1999 (H.

Kim, SNU); 2 c?c?, Seoginbong, Dodong, 10 Aug. 1999 (H. Kim, SNU). Gyeongsangnam-do: 1 Sam-

Jeong Ri, 14 Jul. 1991 (K. Kanmiya, ELKU). Jeollanam-do: 3 $$, Jongseokdae, Mt. Jiri-san, 29 Jul.

1999 (H. Kim, SNU); 1 $, 1 $, Nogodan, Mt. Jiri-san, Gurye-gun, 29 Jul. 1999 (H. Kim, SNU). Jeju-

do: Yonsil, Mt Halla-san, 50 $?, 24 Jul. 1990 (O. Tadauchi, ELKU), 12 $$, 2 27 Jul. 1990 (O.

Tadauchi, ELKU); 13 ? $, 3 <$<$, Mt Halla-san, alt. 1400-1600 m, 25 Jul. 1990 (O. Tadauchi, ELKU);

1 5, Ryuzinkaku, alt. 1600 m, Mt. Hanna, 16 Jul. 1968 (T. Shirozu, ELKU).

European Journal of Taxonomy 137: 1-50 (2015)

Additional description

Labrum (Fig. 20C-D). Basal area approximately 2.2 x as wide as long in female; basal elevation

moderately developed in female, absent in male; distal process of female slender, nearly as long as basal

Fig. 20. Lasioglossum (. Dialictus ) ellipticeps (Bliithgen, 1923). A-B. Head. C-D. Labrum. E. Mese-

pisternum. F. Hairs on metasomal sterna. A, C, E: §. B, D, F: f.

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

area, and without lateral projection, that of male absent; keel of distal process narrow, apically bluntly

pointed; labral fimbria acutely pointed at apex in both sexes.

Sterna. S7-S8 (Fig. 2 IE): S7 with short, apically rounded median process; median process of S8 slightly

present, round or truncate apically, with sparse short hairs.

Male genitalia (Fig. 21A-D). Gonobase nearly flat at bottom, ventral arms connected with each other

at upper ends; gonocoxite smooth, gently sloping in lateral view, inner and outer dorsal margins nearly

parallel; gonostylus small, bud-like in lateral view, located at top of gonocoxite, with sparse short hairs;

ventral retrorse lobe tongue-like, long, apex reaching gonobase, with short hairs ventrally, apical hairs

longer than around one; penis valve higher than gonocoxite, without cleft on top.

Distribution

Russian Far East (Yakutia, Khabarovsk and Primorsk Terr., Irkutsk Prov.), Mongolia and the Korean

Peninsula (north, new record for south).

Fig. 21. Lasioglossum ( Dialictus ) ellipticeps (Bluthgen, 1923), S- A. Genitalia in ventral view. B. Geni¬

talia in dorsal view. C. Genitalia in lateral view. D. Ventral retrorse lobe of genitalia. E. S7-S8. Scale

bars: A-C, E = 0.2 mm, D = 0.1 mm.

European Journal of Taxonomy 137: 1-50 (2015)

Flight records

Female: May to August.

Male: July.

Flower records

In South Korea, this species visits the following 10 species in 6 families. Asteraceae: Erigeron annuus

(L.) Pers., Lactuca indicaL. Balsaminaceae: Impatiens balsaminaL., I. noli-tangere L. Campanulaceae:

Campanula takeshimana Nalcai, Codonopsis lanceolata (Siebold & Zucc.) Trautv., Platycodon

grandiflorus (Jacq.) A. DC. Geraniaceae: Geranium sp. Lamiaceae: Agastache rugosa (Fisch. & C.A.

Mey.) Kuntze. Lythraceae: Lythrum anceps (Koehne) Makino.

Incertae sedis

Lasioglossum ( Dialictus ) miyabei Murao, Ebmer & Tadauchi, 2006

Figs 22B-C, 23D, 28A

Lasioglossum {Evylaeus) miyabei Murao, Ebmer & Tadauchi, 2006: 42—46 (holotype: ELKU, $; type

locality: Hokkaido, Japan).

Lasioglossum (Evylaeus) miyabei - Ebmer 2006: 564-565.

Evylaeus (Glauchalictus) miyabei - Pesenko 2007a: 26. — Pesenko 2007b: 85 (in key), 99 (in key),

114-115.

Diagnosis

This species is separated from other eastern Asian species of Lasioglossum (Dialictus) by a combination

of the following character states: 1) head wider than long in both sexes (HL/HW ratio = 0.91-0.94 in

female, = 0.94-0.98 in male); 2) metepisternum punctate in both sexes; 3) female metasomal terga dimly

Fig. 22. Distribution maps. A. Lasioglossum (Dialictus) ellipticeps (Bluthgen, 1923) in the Korean

Peninsula. B-C. Lasioglossum (Dialictus) miyabei Murao et al. , 2006 (B in the Korean Peninsula, C in

Japan).

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

shiny; 4) male metasomal sterna not modified and 5) and the structure of the male genitalia (Murao et

al. 2006; Fig. 9A-E).

Material examined

JAPAN: Honshu: 2 $ 13 $$, Ashiro, Iwate Pref., 19 Sep. 1976 (Y. Maeta, ELKU); 2 5 5, Nishikawa-

cho, Yamagata Pref., 3 Jun. 2001 (K. Goukon, cGou); 1 $, Mt. Zao, Yokokawa-rindo, Miyagi Pref.,

21 Aug. 2004 (K. Goukon, cGou); 1 $, Agano, Saitama Pref., 23 Oct. 1971 (T. Nambu, ELKU); 1 $,

Sengataki, Karuizawa, Nagano Pref., 27 Sep. 1972 (H. Suda, ELKU); 1 $, Mt. Haku-san, Ishikawa

Pref., 30 Jul. 1962 (Y. Haneda, ELKU).

SOUTH KOREA: Gangwon-do: 1 5, Ohsaenk-ri, Yangyang-gun, 17 Apr. 1994 (O. Tadauchi, ELKU);

1 5, Taewallyong, Myongju-gun, 19 Apr. 1994 (O. Tadauchi, ELKU); 3 5?> Mt. Gariwang-san,

Jeongseon-gun, 30 Jul. 2013 (R. Murao, cMur); 3 $$, Mt. Gariwang-san, alt. 1100 m, 37°27 , 15” N

128°E10”E, 10 Jun. 2013 (O. Tadauchi, ELKU); 4 $ $, Mt. Odae-san, alt. 900m, 37°47 , 0 ,, N 128°32U9

E, 9 Jun. 2013 (O. Tadauchi, ELKU); Mt. Odae-san, 1 $, 10 May 1996 (H.S. Lee, SNU), 1 $, 24 May

2002 (H.S. Lee, QIA); 1 5, Odaesanjang, Mt. Odae-san, Jinbueup, Pyengchang-gun, 24 Apr. 1997 (H.S.

Lee, QIA). 14 55> Mt. Seolcbyung-san, Imgye-ri, Imgyemyeon, Jeongseon-gun, 22 May 2002 (Y. Lee

& J. Yeo, QIA); Baekdamsa temple, Mt. Seolak-san, Sokchosi, 1 5, 13 May 2010 (J. Jeong, QIA), 4

55,6 Sep. 2010 (H.S. Lee, QIA); 1 5, Hwacheondong, Imgyemyeon, Jeongseon-gun, 24 Apr. 1997 (J.

Son, QIA); 1 5, Hwang-i-ri, Seomyeon, Yangyang-gun, alt. 256 m, 37°56 , 25.8” N 128°3U19.7” E, 29

Fig. 23. Habitats in Japan and South Korea. A. Takeshikakami-honiri, Ueda-shi, Nagano Pref., Japan.

B. Matoishi wilderness, Aso-shi, Kumamoto Pref., Japan. C. Mt. Gariwangsan, Gangwon-do, South

Korea. D. Lasioglossum (. Dialictus ) miyabei Murao et al., 2006, 5 on flowers of Patrinia scabiosifolia

Fisch. ex Trevir. in South Korea.

European Journal of Taxonomy 137: 1-50 (2015)

Fig. 24. Photographs for keys: T1 of female. A. Lasioglossum ( Dialictus ) viridellum (Cockerell, 1931).

B. L. (D.) virideglaucum Ebmer & Salcagami, 1994.

Fig. 25. Photographs for keys: T1 of female. A. Lasioglossum ( Dialictus ) viridellum (Cockerell, 1931).

B. L. ( D .) pseudannulipes (Bltithgen, 1925).

Fig. 26. Photographs for keys: PP on supraclypeal area of female. A. Lasioglossum (. Dialictus )

meakanense Murao & Tadauchi, 2009. B. L. ( D .) ellipticeps (Bluthgen, 1923).

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

Apr. 2007 (H.S. Lee, QIA); Mt. Jeombong-san, 1 28 Aug. 1999 (H. Kim, SNU), 1 $, 29 Aug. 1999

(H. Kim, SNU). Jeollabuk-do: 1 §, Jeonglyongchy, Sannaemeon, Namweon-gun, 14 May 1991 (T.

Saigusa, ELKU); 1 $, Sanlyong-ri, Sannao-myon, Namweon-gun, 29 Apr. 1992 (O. Tadauchi, ELKU).

Gyeongsangnam-do: 1 §, Sam Jeong Li Ma Cheong Meon, Hamyang-gun, 15 May 1991 (T. Saigusa,

ELKU). Jeollanam-do: 1 §, Nogodan, Mt. Jiri-san, Gurye-gun, 26 May 1997 (H.S. Lee, QIA).

Distribution

Russian Far East (Primorsky), the Korean Peninsula (new record) and Japan (Hokkaido, Honshu,

Shikoku, Kyushu, northern Ryukyus).

Flight period

Female: April to October.

Male: July to October.

Fig. 27. Photographs for keys: labrum of male. A. Lasioglossum ( Dialictus ) viridellum (Cockerell,

1931). B. L. (D.) problematicum (Bluthgen, 1923).

Fig. 28. Photographs for keys: hairs on metasomal sterna of male. A. Lasioglossum ( Dialictus ) miyabei

Murao et al., 2006. B. L. (D.) meakanense Murao & Tadauchi, 2009.

European Journal of Taxonomy 137: 1-50 (2015)

Flower records

The floral records in Japan are reported as 36 species in 13 families by Murao et al. (2006). The specimens

from South Korea were collected on the flowers of Aster ageratoides Turcz. (Asteraceae), Potentilla

verticillaris Stephan ex Willd. (Rosaceae), Rhododendron mucronulatum Turcz. (Ericaceae), Salix sp.

(Salicaceae), Sisymbrium luteum (Maxim.) O.E. Schulz (Brassicaceae), and Syringa sp. (Oleaceae).

Habitat

This species has been collected in mountain areas in both Japan and South Korea. One of the collecting

sites (Mt. Gariwangsan) in South Korea is shown in Fig. 23C.

DNA barcodes

The COI gene sequences are deposited as DNA barcodes of L. (D.) miyabei in the DDBJ under accession

number LC027536. This number is also available in GenBank.

Comments

This species belongs to the atroglaucum group in the dissenting systematic study (Murao et al. 2006;

Pesenlco 2007a, as subgenus Glauchalictus ). It differs, however, considerably with regard to the male

character states of the atroglaucum group shown in the present study in having the metasomal sterna

without unique hair tufts (Fig. 28A), S4 or S5 not modified (Fig. 28A), and the penis valve of male

genitalia without a cleft on top. The systematic status of this species could not be decided in the present

study. It will need to be correctly determined using phylogenetic analysis in a future study.

Key to female Lasioglossum (Dialictus) of Japan , the Korean Peninsula and Taiwan:

1. T1 basally covered with thin, appressed hairs (Fig. 24A) .2

- T1 basally with sparse and erect hairs (Fig. 24B) .3

2. T1 usually with strong lineolation on basal to medial area (Fig. 25A).

. Lasioglossum ( Dialictus ) viridellum (Cockerell, 1931)

- T1 without lineolation or with weak lineolation on basal area (Fig. 25B) .

. L. (D.)pseudannulipes { Bliithgen, 1925)

3. Head wider than long (HF/HW ratio 0.91-0.94, n = 10). L. ( D .) miyabei Murao et al., 2006

- Head nearly as long as wide or longer than wide (HL/HW ratio 0.98-1.12, n = 30) .4

4. Mesepisternum reticulate punctate on lower area (Fig. 20E) .5

- Mesepisternum sparsely shallow punctate on lower area (e.g.. Fig. 6E) .6

5. Supraclypeus sparsely punctate as in Fig. 26A (IS = 1—4.7) .

. L. (D.) meakanense Murao & Tadauchi, 2009

- Supraclypeus densely punctate as in Fig. 26B (IS = 1-2) . L. (D.) ellipticeps (Bltithgen, 1923)

6. Mesoscutum posteriorly nearly smooth. L. (Dfyamanei Murao etal., 2006

- Mesoscutum posteriorly wealdy or strongly tessellate .

. L. (D. ) problematicum (B1 uthgen, 1923)and F. (D.)virideglaucum Ebmer&Sakagami, 1994

Key to male Lasioglossum (Dialictus) of Japan , the Korean Peninsula , and Taiwan:

1. Clypeus without yellow spot; labrum with basal elevation depressed centrally (Fig. 27A)

. L. (D.) pseudannulipes (Bltithgen, 1923) and L. ( D .) viridellum (Cockerell, 1931)

- Clypeus yellow on lower half; labrum without basal elevation (Fig. 27B) .2

MURAO R. et al., Dialictus of Japan, the Korean Peninsula, and Taiwan

2. Hind tibia yellow basally and apically, tarsi yellow .3

- Legs black, without yellow coloration.4

3. Head wider than long (FFL/FtW ratio 0.94-0.98, n = 10); metasomal sterna with uniform and sparse

hair, without hair tuft (Fig. 28A). L. (D.) miyabei Murao et al., 2006

- Head longer than wide (HL/HW ratio 1.07-1.12, n = 5); S5 laterally with moderately long hair tufts

(Fig. 28B) . L. ( D .) meakanense Murao & Tadauchi, 2009

4. S5 flat on whole surface, without apical depression.5

- S5 with shallow or deep apical depression (Fig. 5B, E, H, K, N).7

5. S4-S5 flat, laterally with moderately long hair tufts (Fig. 20F); penis valve of genitalia without cleft

on top (Fig. 21C) . L. ( D .) ellipticeps (Bluthgen, 1923)

- S4-S5 weakly swollen medially; S5 with brush-like hairs along apical margin (Figs 13, 14B, H); penis

valve of genitalia with cleft on top (e.g., Fig. 16C) .6

6. Apical margin of S4 straight (Fig. 14A); gonostylus wider than that of L. alishanense (Fig. 16C)

. L. (D.)problematicum (Bluthgen, 1923)

- Apical margin of S4 incurved (Fig. 14G); gonostylus narrower than that of L. problematicum (Fig.

18C) . L. ( D .) alishanense Murao sp. nov.

7. Apical margin of S5 incurved .8

- Apical margin of S5 nearly straight.9

8. Clypeus dull, with strong tessellation; S5 gently incurved (Fig. 5N); gonostylus of genitalia small (Fig.

11C) . L. ( D .) taiwanense Murao sp. nov.

- Clypeus more shiny, with weak tessellation; S5 deeply incurved (F ig. 5H); gonostylus of genitalia large

(Murao et al. 2006: fig. 11) . L. (D.) yamanei Murao et al., 2006

9. S5 with a longitudinal, shallow groove (Fig. 5K). L.(D.) negoroi Murao & Tadauchi, 2008

- S5 without groove .10

10. Hair pattern of S4 as in Fig. 4B; S6 with a distinct pair of hair tufts (Fig. 4B); gonostylus of

genitalia large (Fig. 9C). L. (D.) virideglaucum Ebmer & Sakagami, 1994

Hair pattern of S4 as in Fig. 4A; S6 with sparse hairs (Fig. 4A); gonostylus of genitalia small and

bud-like (Fig. 7C) . L. ( D .) atroglaucum (Strand, 1913)

A checklist of Lasioglossum (Dialictus) in Japan , the Korean Peninsula and Taiwan:

The leucopus group:

Lasioglossum (Dialictus) pseudannulipes (Bluthgen, 1925): Japan

Lasioglossum (Dialictus) viridellum (Cockerell, 1931): Korean Peninsula

Lasioglossum (Dialictus) meakanense Murao & Tadauchi, 2009: Japan

The atroglaucum group:

The atroglaucum subgroup:

Lasioglossum (Dialictus) atroglaucum (Strand, 1913): Taiwan

Lasioglossum (Dialictus) virideglaucum Ebmer & Sakagami, 1994: Japan, Korean Peninsula

Lasioglossum (Dialictus) yamanei Murao, Ebmer & Tadauchi, 2006: Japan

Lasioglossum (Dialictus) negoroi Murao & Tadauchi, 2008: Japan

Lasioglossum (Dialictus) taiwanense Murao sp. nov.: Taiwan

European Journal of Taxonomy 137: 1-50 (2015)

The problematicum subgroup:

Lasioglossum ( Dialictus ) problematicum (Bliithgen, 1923): Japan

Lasioglossum ( Dialictus ) alishanense Murao sp. nov.: Taiwan

The morio group:

Lasioglossum ( Dialictus) ellipticeps (Bluthgen, 1923): Korean Peninsula

Incertae sedis:

Lasioglossum ( Dialictus ) miyabei Murao, Ebmer & Tadauchi, 2006: Japan, Korean Peninsula

Discussion

Twelve species in total are recognized in the countries studied. All species, except for Lasioglossum

( Dialictus ) ellipticeps and L. (D.) virideglaucum, are endemic in Far East Asia. The number of species

per country is as follows: 7 in Japan, 4 in the Korean Peninsula and 3 in Taiwan. We could not examine

enough material of L. (Dialictus) from either the Korean Peninsula (particularly, from the sample of

North Korea) or Taiwan in the present study to definitively survey the fauna. The number of species

in both countries will likely increase, resulting in additional taxonomic studies of the L. ( Dialictus )

fauna. In the present study, all L. (Dialictus ) species from Taiwan are endemic. The other subgenera of

Taiwanese Lasioglossum may include many endemic species according to the first author’s preliminary

examination of Taiwanese Lasioglossum specimens (Murao unpublished). In eastern Asia, Taiwan (as a

consequence of the isolation from or land-bridge formation with the continent) and the Ryukyu Islands

(Japan, with its many endemic Lasioglossum species, see Murao 2012; Murao & Tadauchi 2006, 2009b;

Murao et al. 2009b, 2010) would be important areas to consider in order to investigate the allopatric

speciation of Lasioglossum.

Acknowledgements

We would like to express our thanks to Dr. Layne Julian Westover (ELKU) for his kindness in brushing

up an early draft. Ryuki Murao would especially like to thank Dr. Yulia Astafurova (ZISP), Mr. Fritz

Gusenleitner (OLML), Mr. Maximilian Schwarz (Linz, Austria), Mr. Andreas Werner Ebmer (Puchenau,

Austria), Dr. Andreas Taeger (SDEI), Mr. Andrew Liston (SDEI) and Dr. Frank Koch (Museum far

Naturkunde an der Humboldt Universitat zu Berlin, Germany) for their help in my examining their

Palaearctic and Oriental Lasioglossum collections. He also thanks Prof. Emeritus Dr. Yasuo Maeta

(Shimane, Japan), Mr. Katsuo Goukon (Miyagi, Japan), and Dr. Ryoichi Miyanaga (Shimane University,

Japan) for loan of valuable specimens and literature. This research was supported by the JSPS Institutional

Program for Young Researcher Overseas Visits (to Murao), the Environment Research and Technology

Development Fund (S-9-2(8)) of the Ministry of the Environment, and the Grant-in-Aid for Scientific

Research (B) from the Japan Society for the Promotion of Science (No. 24405016) (Head Researcher:

Osamu Tadauchi).

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Manuscript received: 11 March 2015

Manuscript accepted: 3 June 2015

Published on: 14 September 2015

Topic editor: Koen Martens

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, De nm ark