www. europeanj ournaloftaxonomy. eu

2016 • Hsiao Y. et al.

http://dx.doi.org/10.5852/ejt.2016.170

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

Research article

urn:lsid:zoobank.org:pub:5203E7A5-1592-4FlD-9F3C-24E6791D9429

Taxonomic revision of the Lycocerus hanatanii species group

(Coleoptera, Cantharidae), with the description of

new species from Taiwan

Yun HSIAO >, Yuichi OKUSHIMA 2 , Ping-Shih YANG 3 & Chiun-Cheng KO 4 *

13 4 Department of Entomology, National Taiwan University, No. 27, Lane 113, Sec. 4,

Roosevelt Rd., Taipei 10617, Taiwan.

2 Kurashiki Museum of Natural History, Chuo 2-6-1, Kurashiki-shi,

Okayama Pref., 710-0046 Japan.

* Corresponding author: kocc2501 @ntu.edu.tw

1 Email: [email protected]

2 Email: [email protected]

3 Email: [email protected]

1 urn:lsid:zoobank.org:author:6E5C734A-1809-46Al-A595-5A0CB7B39BA0

2 urn:lsid:zoobank.org:author:BB590861-E5AE-438B-AC0E-A8724F8FAAB5

3 urn:lsid:zoobank.org: author: 0CD84731-DCC1-4A68-BE78-E543D35FA5A2

4 urn:lsid:zoobank.org:author:290CF231-2518-4C91-A2EA-C184C9728C7F

Abstract. The Lycocerus hanatanii species group is revised, with the addition of seven taxa:

L. araticollis (Fairmaire, 1897), L. nigripennis (Pic, 1938), L. griseopubens (Pie, 1928), L. yitingi

Hsiao & Okushima sp. nov., L. aurantiacus Hsiao & Okushima sp. nov., L. evangelium Hsiao &

Okushima sp. nov. and L. kintaroi Hsiao & Okushima sp. nov. Supplementary descriptions of the

males of L. araticollis and L. griseopubens are provided. Lycocerus nigripennis (Pic, 1938) and L.

pictus (Wittmer, 1983) are redescribed in detail. Each species is provided with photos or illustrations

of genitalia of both sexes and abdominal ventrite VII of the female if available. Distribution maps and

a key to the species of the L. hanatanii species group are presented. In addition, the monophyly of the

L. hanatanii species group is supported based on a morphological phylogenetic analysis.

Keywords. Soldier beetles, taxonomy, new taxa, classification, Asia.

Hsiao Y., Okushima Y., Yang P.-S. & Ko C.-C. 2016. Taxonomic revision of the Lycocerus hanatanii species group

(Coleoptera, Cantharidae), with the description of new species from Taiwan. European Journal of Taxonomy 170:

1-33. http://dx.doi.org/10.5852/ejt.2016.17Q

Introduction

The genus Lycocerus Gorham, 1889 is a highly species-rich taxon in Asia, with more than 300 species

and distributed in the Oriental and eastern Palaearctic regions (Kazantsev & Brancucci 2007). Okushima

(2005) revised the Japanese fauna of Lycocerus and proposed 9 species groups with 23 subgroups based

1

European Journal of Taxonomy 170 : 1-33 ( 2016 )

on morphological characters, including external morphology and the genital organs of both sexes.

Okushima & Brancucci (2008) and Yang et al. (2014) proposed two more species groups based on

species from Laos, Vietnam and China. All of the Japanese species and several species from China,

Taiwan, India and the Indo-China Peninsula were included in these defined species groups (Okushima

2005, 2007a, 2007b; Okushima & Brancucci 2008; Yang & Yang 2011; Okushima & Yang 2013; Yang

et al. 2014; Hsiao & Okushima 2015). However, there has been no systematic study to examine the

monophyly of each species group or to clarity their relationship using a phylogenetic analysis.

The Lycocerus hanatanii species group is characterized by the following combination of characters:

small-sized body; small and not very prominent eyes; knife-shaped and slender apical maxillary

palpomere; filiform antennae, not exceeding half of elytral length, without groove; subquadrate

pronotum; both sexes with simple tarsal claws; laterophyses of aedeagus very short, barely visible or

invisible in lateral view; long spermathecal duct; spermatheca provided with a spiral tube. So far only

three species, L. hanatanii (Okushima, 1996), L. ueharaensis (Okushima, 1996) and L. pictus (Wittmer,

1983), have been included (Okushima 2005).

In this study, we aim to revise the fauna of this species group, with the addition of three new members,

L. araticollis (Fairmaire, 1897), L. nigripennis (Pic, 1938) and L. griseopubens (Pie, 1928), and the

description of four new taxa, L. yitingi Hsiao & Okushima sp. nov., L. aurantiacus Hsiao & Okushima

sp. nov., L. evangelium Hsiao & Okushima sp. nov., and L. kintaroi Hsiao & Okushima sp. nov. We also

use a morphology-based phylogenetic approach to support our classification and confirm the monophyly

of the L. hanatanii species group.

Materials and methods

The materials examined in this paper are deposited in the following institutions and personal collection:

BMNH

CAS

CBM

EUMJ

KURA

MNHN

NHMB

NMNS

SDEI

TARI

YHC

Natural History Museum, London, UK

California Academy of Sciences, San Francisco, USA

Natural History Museum and Institute, Chiba, Japan

Ehime University Museum, Matsuyama, Japan

Kurashiki Museum of Natural History, Kurashiki, Japan

Museum national d’Histoire naturelle, Paris, France

Naturhistorisches Museum Basel, Switzerland

National Museum of Natural Science, Taichung, Taiwan

Senckenberg, Deutsches Entomologisches Institut, Muncheberg, Germany

Taiwan Agricultural Research Institute, Wufeng, Taichung, Taiwan

Y. Hsiao’s private collection

The terminology used in the descriptions and the methods employed follow those of Okushima (2005)

and Hsiao (2015), but Double Stain (BioQuip, USA) was used for the staining of female genitalia. The

distribution map was prepared using the SimpleMappr program (Shorthouse 2010), based on the label

data of examined material in this study. Some old names or supplementary information on localities are

updated in square brackets [ ] if necessary. The type specimen data are quoted verbatim: [p] indicates

that the following data are printed and [h] that they are handwritten. A double backslash (//) is used to

separate data from different labels and a single one (/) to separate data from different lines on the same

label. Abbreviations:

ex

ag

di

specimen/s

accessory gland

diverticulum

2

HSIAO Y. et al.. Revision of the Lycocerus hanatanii species group

dp =

dorsal plate

is =

inner sac

i P =

laterophyses

sd =

spermathecal duct

sp =

spermatheca

te =

tegmen

va =

vagina

vp =

ventral process

Following Okushima (2005), Kasantsev (1999) and our observations, 29 adult morphological characters

and 1 geographical character were used for the phylogenetic analysis as follows:

01 Body size: (0) small (5-7mm); (1) medium (8-10 mm ); (2) large (> 10 mm ).

02 Body form judged on the ratio of the length to the width at the widest part of the conjoint elytra:

(0) stout (about 2.2); (1) middle (about 2.5-3.0); slender (mainly greater than 3.0).

03 Antennal length in males: (0) exceeding Vi elytral length; (1) extending to Vi elytral length.

04 Ante nn al length in females: (0) extending to Vi of elytral length; (1) barely exceeding the humeri,

but not extending to Vi of elytral length.

05 Form of antennae: (0) filiform; (1) serrate.

06 Grooves on male antennal segments: (0) absent; (1) present.

07 Shape of terminal maxillary palpomere: (0) stout (Fig. 1A); (1) subtriangular (Fig. IB); (2) slender

(Fig. 1C).

08 Size of eyes: (0) large and prominent (Fig. ID); (1) small and not very prominent (Fig. IE).

09 Vertex: (0) flat and glabrous; (1) faintly hollowed and distinctly punctate.

10 Form of pronotum: (0) trapezoid (Fig. IF); (1) subquadrate (Fig. 1G); (2) flattened and rectangular

(Fig. 1H).

11 Greatest width of pronotum in male: (0) wider than head; (1) nearly as wide as head; (2) narrower

than head.

12 Angular convexity on pronotum: (0) distinct (Fig. 1G); (1) indistinct.

13 Form of elytra: (0) subparallel at the sides; (1) apically expanded; (2) moderately narrowed apically;

(3) strongly narrowed apically.

14 Costae on elytra: (0) distinct; (1) hardly recognizable.

15 Size of legs: (0) stout (Fig. II); (1) slender (Fig. 1J).

16 Form of male claws: (0) all claws simple; (1) each outer claw of fore and middle legs with tooth.

17 Form of female claws: (0) all claws simple; (1) each outer claw of fore and middle legs with tooth.

18 Dorsal plate of each paramere of aedeagus: (0) fused or nearly fused; (1) separated.

19 Inner surface of dorsal plate of aedeagus: (0) modified, with processes; (1) simple.

20 Laterophyses of aedeagus: (0) nearly fused; (1) widely separated on both sides of median lobe.

21 Laterophyses of aedeagus: (0) long, distinctly visible in lateral view; (1) short, barely visible or

invisible in lateral view.

22 Apical part of median lobe of aedeagus: (0) widened apically; (1) strongly narrowed apically; (2)

moderately narrowed apically.

23 Apical part of vagina: (0) broad; (1) extended apically as a thick duct.

24 Shape of diverticulum: (0) short, thick and rod-shaped; (1) long and thin; (2) short and thin.

25 Spermathecal duct: (0) absent; (1) present.

26 If spermathecal duct present, length of spermathecal duct: (0) short; (1) long.

27 Shape of spermatheca: (0) short, sac-shaped; (1) long and thin, provided with spiral tube(s).

28 If spermatheca long and thin: (0) provided with a spiral tube; (1) provided with two spiral tubes.

29 Shape of accessory gland: (0) simple (thin, filiform); (1) modified (abruptly expanded).

30 Distribution: (0) continental; (1) insular.

3

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Table 1. Data matrix.

x

Oj

05

•Hi

05

a

•Hi

"Hi

a-

<5

R

r

R

«H

R

»R

R

>5

SR

•Hi

<5

R

s

■H

R

05

<R

05

a

05

R

.05

5 Hi

R 2

R

.05

•Hi

05

R

a

R

Sh

R

&

R

•Hi

>5

R

s*

Sh

R

5h

R

H.

a-

>•

r

"Hi

05

R

§

1

a

R

•R

R

05

R

£

*4

ij ij ij ij ij ij ij

©

a

•S3

•

a

tfl

05

jR

R

05

R

•Hi

•5

•R

R

5h

t

6jO

•Hi

SR

R

*4

*4 *4 *4 kj

01

2

1

0

1

2

1

0

02

0

1

0

1

2

1

03

1

0

1

0

1

0

1

?

0

1

04

1

0

1

0

1

?

1

?

05

0

1

0

06

1

0

1

0

1

0

1

0

?

0

07

0

1

2

0

1

2

1

2

08

0

1

09

0

1

10

2

0

1

0

1

0

1

11

0

1

2

1

2

1

12

1

0

1

0

13

2

1

0

3

0

14

1

0

1

15

1

0

1

16

0

1

0

1

0

1

0

17

0

1

0

1

0

1

0

?

0

?

18

0

1

19

0

1

0

1

20

0

1

21

0

1

22

0

2

1

?

1

23

0

1

0

1

0

1

?

1

?

1

?

24

0

1

2

1

2

1

?

1

?

1

?

25

0

1

?

1

?

1

?

26

-

0

1

0

1

?

1

?

1

?

27

0

1

?

1

?

1

?

28

-

0

1

0

1

0

?

0

?

0

?

29

0

1

0

?

0

?

0

?

30

0

1

0

1

0

1

0

1

0

1

The ingroup taxa include 3 Lycocerus species previously attributed to the L. hanatanii Group in

Okushima (2005) and 6 species we herein attribute to this species group, L. rhagonychiformis (Wittmer,

1983) and one representative species of each defined species group, including L. maculicollis (Hope,

1831), L. suturellus suturellus (Motschulsky, 1860), L. vitellinus (Kiesenwetter, 1874), L. purpureus

Kazantsev, 2007, L. adusticollis (Kiesenwetter, 1874), L. yamatensis Okushima, 2005, L. watanabei

4

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

(Ishida & M. Sato, 1993), L. aegrotus (Kiesenwetter, 1874), L. michiakii Okushima & Brancucci, 2008

and L. sichuanus Y. Yang & X. Yang, 2014. The outgroup taxon is Themus ( Themus ) nobilis nobilis

(Gorham, 1889). The data matrix (Table 1) was coded and edited with WinClada ver. 10.00.08 (Nixon

2002). Inapplicable data were coded as and missing data were coded with “?”. Most characters

were based on our own observations. The data matrix was analyzed using NONA ver. 2.0 (Goloboff

1999) from WinClada. All characters were run equally weighted and not additive. The commands of

heuristic analysis, ‘hold 10000’, ‘hold/20’, ‘mult*50’ and ‘mult*max*’ were used to search for the most

parsimonious tree(s). In the resultant trees, the unsupported nodes were hard collapsed and only the

best trees were kept in WinClada. The characters and character states on the tree were optimized using

unambiguous character changes in WinClada.

Results

Phylogenetic analysis of the L. hanatani i species group

The heuristic analysis resulted in 10 most parsimonious trees (L 63; Cl 61; RI 80) and their strict

consensus tree (L 72; Cl 54; RI 72) is used in the discussion (Fig. 9). The monophyly of the L. hanatanii

species group is supported by several synapomorphies: relatively short antennae, slender apical maxillary

palpomere and very short laterophyses of the aedeagus. Lycocerus rhagonychiformis (Wittmer, 1983)

is presented as the sister group of the L. hanatanii species group, which shows high morphological

similarity to it. However, the relationships between the clade L. hanatanii Group + L. rhagonychiformis

and the other species groups are presented as an unresolved bush. Hence, the phylogenetic position of

this clade remains unknown.

Taxonomy

Key to the species of the L. hanatanii species group

1. Coloration: mostly yellowish or blackish, with wide yellowish stripe on head, pronotum and elytra,

respectively .2

- Coloration: mostly blackish; pronotum completely black or fringed with yellowish margins or

orange to yellowish brown, with black marking in middle; elytra completely black, or with small

and slender yellowish brown stripes on humeri .5

2. Head black, with frons yellowish; pronotum yellowish brown, with a large black marking in middle;

aedeagus: dorsal plate of each paramere nearly touching on inner margin (Fig. IK) .3

- Head dark yellow, with a black marking on the vertex; pronotum dark yellow, with a longitudinal

black marking in middle, elytra completely pale yellow or fuscous, with pale yellow longitudinal

stripe on each elytron; aedeagus: dorsal plate of each paramere concave on inner margin

(Fig. 1L).4

3. Aedeagus: ventral process expanded apically, curved inwards; dorsal plate with a rounded apex;

laterophyse acute and barely visible in lateral view . L. hanatanii (Okushima, 1996)

- Aedeagus: ventral process almost straight and slightly curved inwards; dorsal plate with a pointed

apex; laterophyse stout and invisible in lateral view. L. ueharaensis (Okushima, 1996)

4. Elytra completely pale yellow; aedeagus: laterophyse with pointed apex (Fig. 3B).

. L. pictus (Wittmer, 1983)

- Elytra fuscous, with wide pale yellow longitudinal stripe on each elytron; aedeagus: laterophyse

with truncated apex (Fig. 3E) . L. yitingi Hsiao & Okushima sp. nov.

5. Aedeagus: dorsal plate of each paramere nearly touching on inner margin (Fig. IK) .6

— Aedeagus: dorsal plate of each paramere concave on inner margin (Fig. 1L) .9

5

European Journal of Taxonomy 170 : 1-33 ( 2016 )

6. Pronotum completely black; aedeagus: apical margin of dorsal plate truncated (Fig. 4B)

. L. araticollis (Fairmaire, 1897)

- Pronotum completely black or orange, with a black marking in middle; aedeagus: apical margin of

dorsal plate rounded (Fig. 4C, G, J).7

7. Coloration of pronotum varying from completely black or orange, with a black marking in middle;

aedeagus: dorsal plate longer than ventral process. L. nigripennis (Pic, 1938)

- Coloration of pronotum uniform, orange, with a black marking in middle; aedeagus: dorsal plate

nearly as long as ventral process .8

8. Surface of body lusterless; aedeagus: ventral process moderately expanded apically, curved

inwards. L. griseopubens (Pie, 1928)

- Surface of body semilustrous; aedeagus: ventral process strongly expanded apically, slightly curved

inwards. L. aurantiacus Hsiao & Okushima sp. nov.

9. Body and legs nearly black; aedeagus: apex of laterophyse more slender and acute (Fig. 5B)

. L. evangelium Hsiao & Okushima sp. nov.

- Pronotum black, fringed with yellowish margins; aedeagus: apex of laterophyse far from apical

margin of dorsal plate in lateral view (Fig. 5E). L. kintaroi Hsiao & Okushima sp. nov.

Fig. 1 . Morphological characters used for the phylogenetic analysis and key. A-C. Terminal maxillary

palpomere. D-E. Eyes. F-H. Pronotum. I-J. Leg. K-L. Inner margin of dorsal plate of aedeagus.

6

HSIAO Y. et al.. Revision of the Lycocerus hanatanii species group

Class Hexapoda Blainville, 1816

Order Coleoptera Linnaeus, 1758

Superfamily Elateroidea Leach, 1815

Family Cantharidae Imli off 1856

Genus Lycocerus Gorham, 1889

Lycocerus hanatanii Group

Lycocerus hanatanii (Okushima, 1996)

Athemellus hanatanii Okushima, 1996: 118, figs 6-10.

Athemus (Athemellus) hanatanii - Takahashi 1998: 43.

Lycocerus hanatanii - Okushima 2005: 242, figs llr, 13u, 15r, 60, 131, 174. —Kazantsev & Brancucci

2007: 250.

Material examined

Holotype

JAPAN: 3, Mt. Omoto-dake, Ishigaki-jima Is., Ryukyus, 1 Mar. 1990, Y. Okushima leg. (KURA).

Other material

JAPAN: 1 3, 1 5 , Omoto-dake, Ishigaki-jima, Okinawa, Ryukyu, 9 Mar. 1999, T. Kuriharaleg. (KURA);

1 3, 1 5 , same locality, 12 Mar. 2002, T. Kurihara leg. (KURA); 15 S3, 7 55 , same locality, 21 Feb.

2003, T. Nalcata leg. (13 S3, 5 5 5 in KURA; 2 33, 2 5 5 in YHC); 2 33, Aira-gawa, Iriomote-jima,

Okinawa, Ryukyu, 14 Mar. 1999, T. Kuriharaleg. (KURA).

Description

See Okushima (1996, 2005).

Distribution

Japan (Okinawa: Iriomote Island and Ishigaki Island).

Lycocerus ueharaensis (Okushima, 1996)

Athemellus ueharaensis Okushima, 1996: 121, figs 11-14.

Athemus (Athemellus) ueharaensis - Takahashi 1998: 43.

Lycocerus ueharaensis - Okushima 2005: 245, figs 61, 132, 175. —Kazantsev & Brancucci 2007: 254.

Material examined

Holotype

JAPAN: 3, Mt. Uehara-yama, Iriomote-jima Is., Ryukyus, 11 Mar. 1995, Y. Okushima leg. (KURA).

Other material

JAPAN: 1 3, Nakasuji, Ishigaki-jima, Okinawa, Ryukyu, 6 Mar. 1999, T. Kurihara leg. (KURA); 10

33, Yanbare, Ishigaki-jima, Okinawa, Ryukyu, 3 Feb. 2003, T. Nakata leg. (8 33 in KURA; 2 33 in

YHC).

Description

See Okushima (1996, 2005).

7

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Distribution

Japan (Okinawa: Iriomote Island and Ishigaki Island).

Lycocerus pictus (Wittmer, 1983)

Figs 2A-B, 3A-C, 6A-B, 7

Athemelluspictus Wittmer, 1983: 170.

Lycocerus pictus - Okushima 2005: 242. — Okushima 2008: 282, fig. 26. — Kazantsev & Brancucci

2007: 253. —Sato etal. 2015: 51.

Diagnosis

This species resembles L. yitingi Hsiao & Okushima sp. nov.; the differential diagnosis is given in

Table 2.

Material examined

Holotype

TAIWAN: 3, Nan shan chi, Nantou Hsien, 9-11 Mar. 1980, T. Shimomura leg. (NHMB).

Paratypes

TAIWAN: 10 ex, same data as for holotype (NHMB).

Other material

TAIWAN: 15 33, 7 5 5> Nanshanchi, Nantou, 27 Mar. 1981, H. Yoshimoto leg. (KURA); 1 §, Hsileng,

Taouan, 26 Apr. 1982, N. Obayashi leg. (KURA); 13 33, 5 $$, Nanshanchi, Nantou, 13 Apr. 1985,

Y. Kusakabe leg. (KURA); 1 §, Kukuan, Taichung, 11-12 May 1986, C.-S. Tin leg. (NMNS); 4 33,

3 55, Nanshanchi, Nantou, 5 Apr. 1988, C. Tou leg. (KURA); 1 3, Baling, Taoyuan, 5 Apr. 1991, Y.

Okushima leg. (KURA); 1 §, Shitzutou, Nantou, 9 Apr. 1991, Y. Okushima leg. (KURA); 28 33, 37

55, Nanshanchi, Nantou, 10 Apr. 1991, Y. Okushima leg. (KURA); 1 3, 3 55 > Kukuan, Taichung, 11

Apr. 1991, Y. Okushima leg. (KURA); 1 5, Shitzutou, Nantou, 12 Apr. 1991, Y. Okushimaleg. (KURA);

1 3, Nanshanchi, Nantou, 12 Apr. 1991, K. Ta leg. (KURA); 4 33, 10 5 5 » Nanshanchi, Nantou, 3 Apr.

1993, K. Okajima & T. Nonaka leg. (KURA); 4 33, Nanshanchi, Nantou, 6 Apr. 1993, T. Nonaka leg.

(KURA); 3 55, Wushe, Nantou, 28-29 Apr. 1993, C.-S. Lin & W.-T. Yang leg. (NMNS); 4 33, 14

55 , Lushan Hot Spring, Nantou, 24 Apr. 1994, T. Kishimoto leg. (KURA); 1 3, Gauvaw [=Luofu],

380 m, Taovuan, 25 Apr. 1996, L.-J. Wang leg. (KURA); 4 33, 3 55» Nanshanchi, Nantou, 21 Mar.

1998, F. Hayashi leg. (KURA); 3 S3, Chunyang, Nantou, 12 Mar-9 Apr. 2002, C.-S. Lin & W.-T. Yang

leg. (NMNS); 1 S, 1 5> Nanshanshi, Nantou, 1 Apr. 2012, Y.-T. Wang leg. (YHC); 1 5, Puli, Nantou,

Apr. 2014, C.-H. Wu leg. (YHC); 2 S3, Chungyang, Nantou, 4 May 2014, Y. Hsiao leg. (YHC); 1 5,

Simashuan Forest Trail, 500m, Miaoli, Y. Hsiao leg. (YHC).

Redescription

Coloration

Eyes black. Head dark yellow, with black marking on vertex. Antennae dark yellow or black, with

first two or three segments and basal parts of remaining segments dark yellow. Pronotum dark yellow,

with black marking in medio-longitudinal part. Elytra completely pale yellow or slightly dark in

narrow suture. Legs dark yellow, with apical parts of femora, tibiae and tarsi blackish. Prosternum dark

yellow, mesoventrites, metaventrites and abdomen black, with margin of abdominal ventrites and entire

abdominal ventrite VII yellowish brown. Body closely covered with fine yellowish pubescence; anterior

margin of clypeus fringed with pale bristles; antennae, elytra and legs with some yellowish bristles

intermingled with primary pubescence.

8

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

Male (Fig. 2A)

Head. Nearly as long as wide; vertex faintly hollowed, depressed along apical margin of clypeus and in

lateral areas before eyes; surface densely and finely punctate and semilustrous; anterior margin of clypeus

Fig. 2. Habitus, dorsal view. A-B. Lycoceruspictus (Wittmer, 1983). C-E. L. yitingi Hsiao & Okushima

sp. nov. F-G. L. araticollis (Fairmaire, 1897). H-I. L. griseopubens (Pie, 1928). J-M. L. nigripennis

(Pic, 1938). N-O. L. aurantiacus Hsiao & Okushima sp. nov. P-Q. L. evangelium Hsiao & Okushima

sp. nov. R. L. kintaroi Hsiao & Okushima sp. nov. C, F, H, J, N, P, R. Name-bearing types. D, O, Q.

Paratypes. A-B, E, G, I, K-M. Non-type specimens. A, C, F-G, I, K-L, N, P, R. Male. B, D-E, H, J, M,

O, Q. Female. Scale bars: 1.0 mm.

9

European Journal of Taxonomy 170 : 1-33 ( 2016 )

arcuate and faintly indented in middle; eyes not so large, globular and slightly prominent, ratio of eye

diameter to interocular space 1:4.4; terminal labial palpomeres rounded ax-shaped; terminal maxillary

palpomeres rounded ax-shaped; antennae filiform, extending to elytral midlength; antennomere I clavate,

II short and a little expanded apicad. III to XI subcylindrical, all antennomeres without groove, ratio of

ante nn omere lengths: 13:10:13:15:16:16:14:14:13.5:13 :14.

Pronotum. Subquadrate, slightly narrower than head, nearly as long as wide; anterior and posterior

margins moderately arcuate; lateral margins feebly sinuate, slightly diverging posteriorly; anterior

angles rounded; posterior angles obtuse; dorsum convex in postero-lateral and hollowed in antero-lateral

areas; medio-longitudinal groove distinct in posterior half; surface smooth and semilustrous. Scutellum

triangular, with rounded apex.

Elytra. Conjointly about 1.4 times as wide as pronotum, about 3.0 times as long as wide, sides

subparallel; surface densely and finely punctate and without luster.

Legs. Moderately slender; femora mostly straight; tibiae mostly straight, with basal part feebly arcuate;

claws simple.

Aedeagus (Fig. 3A-C). Ventral process expanded apically, apex curved inwards, basal part broad and

extending inwards onto ventral side; dorsal plate of each paramere as long as ventral process, stout

and subtruncated, slightly narrowed apically, apical margin with some hairs on it, concave on inner

margin. Laterophyse very short, with pointed apex, invisible in lateral view. Inner sac swollen apically,

somewhat shorter than tegmen.

Body length 4.00-5.50 mm ; width 1.00-1.50 mm

Female (Fig. 2B)

Similar to male. Body wider than in male. Eyes somewhat smaller than in male, ratio of eye diameter

to interocular space 1:4.9. Antennae distinctly shorter than in male, barely exceeding humeri, ratio of

ante nn omere lengths: 13.5:10:13 :15:15:14:14:14:13 :12:14.

Pronotum. Nearly as wide as head, about 1.10 times as long as wide. Elytra conjointly about 1.5 times

as wide as pronotum, about 2.6 times as long as wide. Claws simple. Abdominal ventrite VII widely

emarginate on each side of terminal margin, forming subtriangular lateral lobes, median lobe moderately

developed and triangularly concave in middle, median lobe indistinct in some specimens (Fig. 6A).

Female genitalia (Fig. 6B). Vagina broad and rounded, abruptly extended apically as a long and thick

duct; diverticulum and spermathecal duct arising from apex of long duct of vagina; diverticulum

moderately thin and spiral; spermathecal duct shorter than diverticulum; spermatheca provided with

thin spiral tube, tube longer than diverticulum; accessory gland thin, shorter than spermatheca.

Body length 5.00-6.00 mm ; width 1.50-1.75 mm

Remarks

Individuals of this species were observed visiting flowers of Lauraceae during the day.

Distribution (Fig. 7)

Taiwan (endemic).

10

HSIAO Y. et al. , Revision of the Lycocerus hanatanii species group

Table 2. Diagnostic characters of Lycocerus pictus (Wittmer, 1983) and L. yitingi Hsiao & Olcushima

sp. nov.

L. pictus

L. yitingi sp. nov.

Coloration

Elytra almost pale yellow,

and smaller black parts on

the vertex and legs

Elytra with large fuscous

area, and larger black parts on

the vertex and legs

Aedeagus

Ventral process thicker, laterophyse

with pointed apex, apical margin

of dorsal plate subtruncated

Ventral process thinner, laterophyse

truncated apically, apical margin

of dorsal plate rounded

Abdominal ventrite

VII of female

Mostly yellowish brown

and median lobe

Mostly fuscous

and median lobe

less protuberant

more protuberant

Female genitalia

Vagina broader and rounded,

spermathecal duct and

accessory gland longer

Vagina stouter,

spermathecal duct and

accessory gland shorter

Lycocerus yitingi Hsiao & Olcushima sp. nov.

um:lsid:zoobank.org:act:B425500E-BCB5-4BD2-A57A-2AB9CA8700E4

Figs 2C-E, 3D-F, 6C-D, 7

Diagnosis

This species resembles L. pictus', the differential diagnosis is given in Table 2.

Etymology

The specific epithet is dedicated to one of its collectors, Mr. Yi-Ting Chung (Pingtung, Taiwan).

Material examined

Holotype

TAIWAN: S, Dahanshan, 1000m, Pingtung, 26 Feb. 2013, Y.-T. Chung leg. (NMNS).

Paratypes

TAIWAN: 3 33, 4 ??, same data as for holotype (2 SS, 2 $$ in NMNS; 1 S, 2 in YHC); 1

5 , Chufengkou, Manchou, Pintung, 15 Jan. 1998, C.-S. Lin leg. (NMNS); 2 SS, 3 $$, Chun Jin

[=Chunrih], 950-1000m, Ping Tung Co., 21 Mar. 2000, C.-L. Li leg. (KURA).

Other material

TAIWAN: 1 c?, 1 $, Chihpen-spa, Peinan Hsiang, Taitung, 5 Apr. 2004, T. Tsuru leg. (KURA); 1 §,

Roshan Waterfall, Fuli, Hualien, 16 Feb. 2014, W.-J. Huang leg. (YHC); 2 $ Majia, Pingtung, 12 Mar.

2014, Y.-T. Chung leg. (YHC).

Description

Coloration

Eyes black. Head dark yellow, with black marking on vertex. Ante nn ae brownish yellow. Pronotum

dark yellow, with black marking in medio-longitudinal part. Elytra fuscous, with wide, pale yellow

longitudinal stripe on each elytron. Legs black, with costae, trochanters and basal parts of femora

dark yellow; tibiae with dark yellow tinge. Prosternum dark yellow, mesoventritres, metaventrites

and abdomen black, with margin of abdominal ventrites pale yellow. Body closely covered with fine

11

European Journal of Taxonomy 170 : 1-33 ( 2016 )

yellowish pubescence; anterior margin of clypeus fringed with pale bristles; antennae, elytra and legs

with some yellowish bristles intermingled with primary pubescence.

Male (Fig. 2C)

Head. Nearly as long as wide; vertex faintly hollowed, depressed along apical margin of clypeus and in

lateral areas before eyes; surface densely and finely punctate and semilustrous; anterior margin of clypeus

Fig. 3. Aedeagus. A-C. Lycoceruspictus (Wittmer, 1983). D-F. L. yitingi Hsiao & Okushima sp. nov.

A, D. Ventral view. B, E. Lateral view. C, F. Dorsal view. Scale bars: 0.5 mm.

12

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

arcuate and faintly indented in middle; eyes not so large, globular and slightly prominent, ratio of eye

diameter to interocular space 1:4.6; terminal labial palpomeres rounded ax-shaped; terminal maxillary

palpomeres rounded ax-shaped; antennae filiform, extending to elytral midlength; antennomere I clavate,

II short and a little expanded apicad, III to XI subcylindrical, all antennomeres without groove, ratio of

ante nn omere lengths: 11:10:12:16:16:16:15:15:14:13:14.

Pronotum. Subquadrate, about 0.90 times as wide as head, about 1.10 times as long as wide; anterior

and posterior margins moderately arcuate; lateral margins feebly sinuate, slightly diverging posteriorly;

anterior angles rounded; posterior angles obtuse; dorsum convex in postero-lateral and hollowed in antero¬

lateral areas; medio-longitudinal groove distinct in posterior half; surface smooth and semilustrous.

Scutellum triangular with rounded apex.

Elytra. Conjointly about 1.5 times as wide as pronotum, about 3.1 times as long as wide, sides

subparallel; surface covered with minute grains and without luster.

Legs. Moderately slender; femora mostly straight; tibiae mostly straight, with basal part feebly arcuate;

claws simple.

Aedeagus (Fig. 3D-F). Ventral process expanded apically, apex curved inwards, basal part broad and

extending inwards onto ventral side; dorsal plate of each paramere as long as ventral process, stout,

apical margin rounded, with some hairs on it, concave on inner margin. Laterophyse very short and

truncated apically, invisible in lateral view. Inner sac swollen, lengthened behind, and somewhat shorter

than tegmen.

Body length 4.95-6.70 mm (holotype: 5.00 mm ); width 1.00-1.70 mm (1.00 mm).

Female (Fig. 2D)

Similar to male. Body wider than in male. Eyes somewhat smaller than in male, ratio of eye diameter

to interocular space 1:5.1. Antennae distinctly shorter than in male, barely exceeding humeri, ratio of

ante nn omere lengths: 11:10:11:13:13:13:12:12:11:10:12.

Pronotum. Nearly as wide as head, slightly longer than width. Elytra conjointly about 1.5 times as wide

as pronotum, about 2.9 times as long as wide. Claws simple. Abdominal ventrite VII widely emarginate

on each side of terminal margin, forming subtriangular lateral lobes, median lobe moderately developed

and triangularly concave in middle (Fig. 6C).

Female genitalia (Fig. 6D). Vagina stout and abruptly extended apically as a long and thick duct;

diverticulum and spermathecal duct arising from apex of long duct of vagina; diverticulum moderately

thin and spiral; spermathecal duct shorter than diverticulum; spermatheca provided with a thin spiral

tube, tube longer than diverticulum; accessory gland thin, shorter than spermatheca.

Body length 6.00-7.50 mm ; width 1.30-1.80 mm

Remarks

Several examined specimens listed as other material were different from the type by having the head,

antennae, thorax, basal half of the femora and tibiae orange yellow; the black markings on the vertex,

pronotum and legs smaller or indistinct; elytra mostly dark yellow, with suture and lateral sides narrowly

black (Fig. 2E), but display no distinct differences compared to the type specimens on the structure of

the aedeagus. Because insufficient materials were available and distinct diagnostic characters were not

discovered, we herein prefer to regard these specimens as representing a color variation of L. yitingi

sp. nov. rather than giving it a new taxonomic status. Nevertheless, we choose not to place these

13

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Fig. 4. Aedeagus. A-B. Lycocerus araticollis (Fairmaire, 1897). C-D. L. griseopubens (Pic, 1928).

E-G. L. nigripennis (Pic, 1938). H-J. L. aurantiacus Hsiao & Okushima sp. nov. A. Ventrolateral view.

B. Dorsolateral view. E, H. Ventral view. D, F, I. Lateral view. C, G, J. Dorsal view. Scale bars: 0.5 mm.

14

HSIAO Y. et al.. Revision of the Lycocerus hanatanii species group

specimens among the paratypes because they will possibly be confirmed as a new taxon in the future

after more material is examined.

Distribution (Fig. 7)

Taiwan (endemic).

Lycocerus araticollis (Fairmaire, 1897)

Figs 2F-G, 4A-B

Telephorus araticollis Fairmaire, 1897: 216.

Cantharis araticollis - Delkeskamp 1977: 62. — Kazantsev & Brancucci 2007: 239.

Lycocerus araticollis - Okushima 2007b: 257, fig. 1.

Diagnosis

This species resembles the nearly black forms of L. nigripennis and L. evangelium Hsiao & Okushima

sp. nov.; the differential diagnosis is given in Table 3.

Material examined

Lectotype

CHINA: $ (MNHN): [h] Kiangsi [=Tiangxi] / Gonon / D. David // [p] TYPE // [p] LECTOTYPUS // [h]

Telephorus / araticollis / Fairm / Kiangsi [=Jiangxi] // [h] REM / 95 / 11.6 // [h] Athemellus / araticollis

/ (Fairm.) / [p] det. W. Wittmer.

Other material

CHINA: 1 S, Taichow, 19 Apr. 1935, collector unknown (NHMB).

Note

The locality of the additional material possibly referred to Taizhou, but there are two localities in China

named Taizhou in Zhejiang and Jiangsu, respectively.

Description

See Okushima (2007b).

Supplementary description

Aedeagus (Fig. 4A-B). Ventral process expanded and rounded apically, apex curved inwards, basal part

broad and extending inwards onto ventral side; dorsal plate of each paramere somewhat longer than

ventral process, protuberant, apical margin truncated, with some hairs on it. Laterophyse very short and

slender, with pointed apex, barely visible in lateral view.

Remarks

This species should be attributed to L. hanatanii species group because its morphological characteristics

match the definition of this species group, in agreement with the cladogram (Fig. 9), although the female

genitalia have not been examined.

Distribution

China (Jiangxi Province).

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European Journal of Taxonomy 170 : 1-33 ( 2016 )

Lycocerus griseopubens (Pic, 1928)

Figs 2H-I, 4C-D

Malthinus atripennis var. griseopubens Pic, 1928: 55.

Malthinus nigripennis var. griseopubens - Delkeskamp 1977: 354.

Lycocerus atripennis - Kazantsev 2007: 48. — Kazantsev & Brancucci 2007: 249.

Lycocerus griseopubens - Okushima 2007b: 259, figs 2, 6.

Diagnosis

This species resembles the yellowish brown pronotum forms of L. nigripennis and L. aurantiacus Hsiao

& Okushima sp. nov.; the differential diagnosis is given in Table 3.

Material examined

Holotype

CHINA: $ (MNHN): [p] Futschau [=Fuzhou], China / Cos. G. Siemssen lg / vend. 13. VI 1903. // [h]

var. griseopubens / Pic // [p] HOTOTYPUS // [h] Athemellus / araticollis / griseopubens / (Pic) / [p] det.

W. Wittmer.

Other material

CHINA: 1 S, 1 ?, Fiuchou [=Fuzhou], Fujian, date and collector unknown (NHMB).

Description

See Okushima (2007b).

Supplementary description

Male (Fig. 21)

Similar to female (= holotype). Body narrower than in female. Eyes somewhat larger than in female,

ratio of eye diameter to interocular space 1:4.5. Ante nn omeres IV to VII longer than in female, VIII to

XIunavailable, ratio of ante nn omere lengths: 15:10:12:15:15:15:12:-:-:-:-.

Pronotum. About 0.95 times as wide as head, nearly as long as wide. Elytra conjointly about 1.6 times

as wide as pronotum, about 2.8 times as long as wide. Legs moderately slender; femora mostly straight;

tibiae mostly straight, with basal part feebly arcuate; claws simple.

Aedeagus (Fig. 4C-D). Ventral process moderately expanded and rounded apically, apex curved

inwards, basal part broad and extending inwards onto ventral side; dorsal plate of each paramere as long

as ventral process, broad, apical margin rounded, with some hairs on it. Laterophyse very short, invisible

in lateral view. Inner sac swollen, shorter than tegmen.

Remarks

This species should be attributed to the L. hanatanii species group as its morphological characteristics

match the definition of this species group, in agreement with the cladogram (Fig. 9); however, the female

genitalia remain unexamined.

Distribution

China (Fujian Province).

16

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

Fig. 5. Aedeagus. A-C. Lycocerus evangelium Hsiao & Okushima sp. nov.

Okushima sp. nov. A, D. Ventral view. B, E. Lateral view. C, F. Dorsal view.

D-F. L. kintaroi Hsiao &

Scale bars: 0.5 mm.

17

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Table 3. Diagnostic characters ofL. nigripennis (Pic, 1938) and similar species.

L. araticollis

L. griseopubens

L. nigripennis

L. aurantiacus

sp. nov.

L. evangelium

sp. nov.

Antennomeres

in males

Shorter

More elongated

Shorter

Coloration of

pronotum

Black

Orange, with a

black marking in

middle

Black or orange

to yellowish

brown, with a

black marking

in middle

Orange, with a

black marking in

middle

Black

Surface of

elytra

Lusterless

Semilustrous

Moderately

lustrous

Convexity on

the pronotum

Less angular

Angular

Pubescence on

the elytra

Slightly sparser

Slightly denser

Ventral process

of aedeagus

Moderate and

expanded

apically, the

apex curved

inwards

Moderate and

expanded

apically, the

apex curved

inwards

Moderate and

expanded

apically, the

apex curved

inwards

Thick and

strongly

expanded

apically, the apex

slightly curved

inwards

Moderate and

expanded

apically, the

apex curved

inwards

Dorsal plate of

aedeagus

Protuberant,

somewhat

longer than

ventral

process, the

apical margin

truncated

Broad, as long

as ventral

process, the

apical margin

rounded

Protuberant

and narrowed

apically,

somewhat

longer than

ventral process,

the apical

margin rounded

Protuberant

and narrowed

apically, slightly

longer than

ventral process,

the apical margin

rounded

Broad, as long

as ventral

process, the

apical margin

rounded, and

concave on

inner margin

Lycocerus nigripennis (Pic, 1938)

Figs 2J-M, 4E-G, 6E-F, 8

Malthinus atripennis Pic, 1922: 30.

Malthinus nigripennis Pic, 1938: 121 [new name for Malthinus atripennis Pic, 1922],

Podabrinus taihokuensis Wittmer, 1954: 276.

Athemellus taihokuensis -Wittmer 1972: 126, fig. 1.

Athemellus atripennis -Wittmer 1983: 168.

Lycocerus atripennis - Kazantsev & Brancucci 2007: 249.

Lycocerus nigripennis - Okushima 2007b: 260; 2008: 282, fig. 25. — Sato et al. 2015: 51.

Diagnosis

The nearly black form of this species resembles L. araticollis and L. evangelium Hsiao & Okushima sp.

nov., and the yellowish brown pronotum form resembles L. griseopubens and L. aurantiacus Hsiao &

Okushima sp. nov.; the differential diagnosis is given in Table 3.

18

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

Fig. 6. A-B. Lycocerus pictus (Wittmer, 1983). C-D. L. yitingi Hsiao & Okushima sp. nov.

E-F. L. nigripennis (Pic, 1938). G-H. L. aurantiacus Hsiao & Okushima sp. nov. I-J. L. evangelium

Hsiao & Okushima sp. nov. A, C, E, G, I. Abdominal ventrite VII of female. B, D, F, H, J. Female

genitalia, lateral view. Scale bars: A, C, E, G, I = 1.0 mm; B, D, F, H, J = 0.5 mm.

19

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Material examined

Syntype of Malthinus atripennis Pic, 1922

TAIWAN: 5 (SDEI): [p] Hokuto / [h] III / [p] Formosa / H. Sauter, 1912 // [p] Syntypus // [h] Malthinus

/ atripennis / Pic // [h] Athemellus / atripennis / (Pic) / [p] det. W. Wittmer // [p] DEI Coleoptera / #

200077. [Note: This female syntype of M. atripennis is possibly the female holotype which Wittmer

(1983) referred to.]

Holotype of Podabrinus taihokuensis Wittmer, 1954

TAIWAN: S (CAS): [p] Taihoku / Formosa / [h] III-27-32 // [p] L. Gressitt / Collection // [p] L. Gressitt

Collector // [h] Elolototypns IS [h] Podahrinvis / taihokuensis / Wittmer / [p] det. W. Wittmer IS [p]

California Academy of Sciences / Type No. [h] 7787.

Paratype of Podabrinus taihokuensis Wittmer, 1954

TAIWAN: $ (NHMB): [h] A. / taihokuensis / (Wittmer) / [p] det. W. Wittmer // [h] P. / taihokuensis /

Wittmer / [p] det. W. Wittmer // [h] Athemellus / atripennis / (Pic) / [p] det. W. Wittmer // [p] Paratypus

// [p] Taihoku / Formosa / [h] III-27-32 // [h] 37 // [p] F. Gressitt / Collection // [p] Naturhistorisches /

Museum Basel / Coll. W. Wittmer // [p] CANTHARIDAE / CANTH00003105.

Other material

TAIWAN: 2 c?(?, Junhua, Taoyuan, 11 Mar. 1972, B.-S. Chang leg. (NMNS); 1 Bajie [=Baigi],

Taoyuan, 11 Mar. 1972, B.-S. Chang leg. (NMNS); 1 $, Hsileng, Taoyuan Hsien, 2 Apr. 1982,

N. Ohbayashi leg. (EUMJ); 2 $$, Fushan spa, Nantou Hsien, 2 May 1982, N. Ohbayashi leg. (KURA);

1 5, Fulcukoo, Toen-ken [= Taoyuan], 28 Jan. 1983, K. Baba leg. (EUMJ); 1 S, Tam Shui, N Taiwan,

20 Feb. 1987, K. Baba leg. (EUMJ); 1 Sun Moon Fake, 1000 m, M Taiwan, 14 Mar. 1989, K. Baba

leg. (EUMJ); 2 $$, 1 $, Mu-Tsar, Taipei, 9 Feb. 1991, C.-F. Fi leg. (KURA); 1 $, Baling, Taoyuan

Hsien, 5-8 Apr. 1991, T. Kishimoto leg. (KURA); 1 <$,2 55> Nanshanchi, Nantou Hsien, 10 Apr.

1991, Y. Okushima leg. (KURA); 1 §, Fuhsing, Taoyuan, 30 Mar. 1992,1.-S. Hsu leg. (NMNS); 1 S,

1 5, Wanjung, Hualien, 2 Feb. 1992, C.-Y. Fee leg. (NMNS); 2 $$, 3 $$, Yangmingshan, Taipei,

27 Mar. 1993, T. Nonaka leg. (KURA); 2 $ §, Fushan spa, Nantou Hsien, 24 Apr. 1994, T. Kishimoto

leg. (KURA); 1 §, Chikou, 350m, Taoyuan Hsien, 25 Apr. 1996, F.-J. Wang leg. (KURA); 1

I 5, Fushan, 21 Mar. 1998, M. Sato leg. (KURA); 7 $$, Paling, Taoyuan, 31 Mar. 1998, M. Sato leg.

(KURA); 1 c?, Fushan, Nantou, 2 Apr. 1998, F. Hayashi leg. (KURA); 4 2 $ §, Gauvaw [=Fuofu],

380 m, Taovuan, 25 Apr. 1998, F.-J. Wang leg. (KURA); 2 Chunyang, Jenai, Nantou, 8-10 Mar.

1999, C.-S. Fin leg. (NMNS); 12 6 $$, Hushichun, Yuanshan, Ilan Hsien, 4-16 Mar. 1999,

T. Mizusawa leg. (KURA); 7 1 $, Yuanshan, Ilan Hsien, 23 Mar. 2000, T. Mizusawa leg. (KURA);

II 13 55, Yuanshan, Ilan Hsien, 23-25 Mar. 2000, T. Mizusawa leg. (KURA); 1 S, Foshan,

Sanxia, Taipei, 27 Apr. 2009, S.-H. Chang leg. (YHC); 1 5 » Baishihu, Neihu, Taipei, 14 Feb. 2013,

F. Huang leg. (YHC); 5 2 5 5 > Neikouli, Neihu, Taipei, 23 Feb. 2014, F. Huang leg. (YHC); 3

1 5 , Shuanchiyen, Taipei, 7 Mar. 2014, F. Huang leg. (YHC); 1 5 ? Fuchushan, Taipei, 25 Mar. 2014,

C.-J. Fi leg. (YHC); 1 5 ? Baehengonlu [=Northern Cross Island Highway], 9 Apr. 2014, F. Huang leg.

(YHC); 1 5 , Fushan, Taipei, 24 May 2014, F. Huang leg. (YHC); 1 Fuchushan, Taipei, 27 Feb. 2015,

T. Hsieh leg. (YHC); 1 5 » Huangdidian, Shuding, Taipei, 28 Feb. 2015, F. Huang leg. (YHC); 4

6 5 5> Kantoushan, Tainan, 28 Feb. 2015, W.-C. Fiao leg. (YHC); 3 <$<$, 4 5 5> Xankonchuan, Danshui,

Taipei, Y.-J. Fu leg. (YHC); 1 5 , Shanming, Taoyuan, 14 Mar. 2015, Y. Hsiao leg. (YHC).

Redescription

Coloration

Eyes black. Head black, but brown in frons. Antennae black, with yellow tinge in the first two segments.

Mandibles yellowish brown. Pronotum yellowish brown to orange, with a black marking in medio-

longitudinal part. Elytra black. Fegs black, with coxae more or less yellowish brown. Prosternum

20

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

yellowish brown; mesoventrites, metaventrites and abdomen black. Body closely covered with pale

pubescence; anterior margin of clypeus, antennae, elytra and legs fringed with pale bristles.

Male (Fig. 2K-L)

Head. Nearly as long as wide; vertex faintly hollowed, depressed along apical margin of clypeus and

in lateral areas before eyes; surface densely and bnely punctate and semilustrous; anterior margin of

ln l &tA 120 121 122

Fig. 7. Distribution map of Lycocerus pictus (Wittmer, 1983) and L. yitingi Hsiao & Okushima sp. nov.

from Taiwan.

21

European Journal of Taxonomy 170 : 1-33 ( 2016 )

clypeus arcuate and faintly indented in middle; eyes not so large, globular and slightly prominent, ratio

of eye diameter to interocular space 1:4.0; terminal labial palpomeres rounded ax-shaped; terminal

maxillary palpomeres rounded ax-shaped; antennae filiform, somewhat exceeding elytral midlength;

antennomere I clavate, II short and a little expanded apicad. III to XI subcylindrical, all antennomeres

without groove, ratio of antennomere lengths: 15:10:14.5:17.5 :17.5:17.5:17:16:15.5:15:16.5.

Pronotum. Subquadrate, about 0.90 times as wide as head, slightly longer than width; anterior and

posterior margins moderately arcuate; lateral margins feebly sinuate, slightly diverging posteriorly;

anterior angles rounded; posterior angles obtuse; dorsum convex in postero-lateral and hollowed

in antero-lateral areas; medio-longitudinal groove distinct in posterior half; surface smooth and

semilustrous. Scutellum triangular, with rounded apex.

Elytra. Conjointly about 1.6 times as wide as pronotum, about 2.9 times as long as wide, sides

subparallel; surface densely and finely punctate and semilustrous.

Legs. Moderately slender; femora mostly straight; tibiae mostly straight, with basal part feebly arcuate;

claws simple.

Aedeagus (Fig. 4E-G). Ventral process expanded apically, apex strongly or moderately expanded, apex

curved inwards, basal part broad and extending inwards onto ventral side; dorsal plate of each paramere

somewhat longer than ventral process, protuberant and narrowed apically, apical margin rounded, with

some hairs on it. Laterophyse very short, with rounded apex, barely visible in lateral view. Inner sac

swollen, lengthened behind, and shorter than tegmen.

Body length 4.80-6.70 mm; width 1.15-2.30 mm

Female (Fig. 2J, M)

Similar to male. Body wider than in male. Eyes somewhat smaller than in male, ratio of eye diameter

to interocular space 1:4.8. Antennae distinctly shorter than in male, barely exceeding humeri, ratio of

ante nn omere lengths: 12:10:11.5:12.5:12:13 :12:11.5:11:10:12.

Pronotum. Nearly as wide as head, nearly as long as wide. Elytra conjointly about 1.6 times as wide

as pronotum, about 3.1 times as long as wide. Abdominal ventrite VII widely emarginate on each

side of terminal margin, forming subtriangular lateral lobes, median lobe considerably developed and

triangularly concave in middle (Fig. 6E).

Female genitalia (Fig. 6F). Vagina rounded, abruptly extended apically as a long and thick duct;

diverticulum and spermathecal duct arising from apex of long duct of vagina; diverticulum moderately

thin and spiral, longer than spermathecal duct; spermatheca provided with thin spiral tube, tube nearly

as long as diverticulum; accessory gland thin, shorter than spermatheca.

Body length 6.10-7.00 mm; width 1.30-1.80 mm

Variation

Body nearly black, including pronotum.

Remarks

This species is widely distributed in low-altitude forest in Taiwan and appears mainly in the spring

according to the label data. It was observed visiting flowers of Machilus thunbergii Siebold & Zucc.

(Lauraceae) during the day. Additionally, during our study of the collections of different institutions and

the survey of the first author’s personal field trip, the nearly black pronotum form was found to be typical

22

HSIAO Y. et al. , Revision of the Lycocerus hanatanii species group

of this species, although all of the type specimens including those of Malthinus atripennis Pic, 1922 and

Podabrimis taihokuensis Wittmer, 1954 represent the yellowish fringed pronotum form.

Distribution (Fig. 8)

Taiwan (endemic).

\ATA 120

At

121

122

25

O L. nigripennis

0 L. evangelium

☆ / aurantiacus

A L. kintaroi

24

23

22

120

75 km

Fig. 8. Distribution map of L. nigripennis (Pic, 1938), L. evangelium Hsiao & Okushima sp. nov.,

L. aurantiacus Hsiao & Okushima sp. nov. and L. kintaroi Hsiao & Okushima sp. nov. from Taiwan.

23

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Lycocerus aurantiacus Hsiao & Okushima sp. nov.

um:lsid:zoobank.org:act: 1CD64F32-4749-48C6-80CF-C8981514E12B

Figs 2N-0, 4H-J, 6G-H, 8

Diagnosis

This species resembles L. griseopubens and the yellowish brown pronotum form of L. nigripennis; the

differential diagnosis is given in Table 3.

Etymology

The specific epithet is derived from the Latin aurantiacus (“orange”), referring to its orange fringed

pronotum.

Type material

Holotype

TAIWAN: Majia Township, 900 m, Pingtung, 12 Mar. 2014, Y.-T. Chung leg. (NMNS).

Paratypes

TAIWAN: 9 9 ? $, same data as for holotype (5 SS, 5 ? ? in NMNS; 2 SS, 2 $ $ in TARI; 2

2 $5 i n YHC); 1 1 §, Tengchih, Taoyuan, Kaohsiung Hsien, 29 Mar.-l Apr. 2001, S. Nagashima

leg. (KURA); 1 §, Baedawushan [=North Dawushan], Pingtung, 2 Apr. 2013, R.-H. Liou leg. (YHC);

2 SS, Tienchih, Kaohsiung, 8 Mar. 2014, W.-C. Liao leg. (1 S in NMNS; 1 S in YHC).

Description

Coloration

Eyes and head black. Antennae black, with yellow tinge at base of first segment. Mandibles yellowish

brown. Pronotum orange, with black marking in medio-longitudinal part. Elytra black. Legs black.

Pro-, meso- and metasterna, as well as abdomen black, with margins of abdominal ventrites yellowish

brown. Body closely covered with pale pubescence; anterior margin of clypeus, antennae, elytra and

legs fringed with pale bristles.

Male (Fig. 2N)

Head. Nearly as long as wide; vertex faintly hollowed, depressed along apical margin of clypeus and in

lateral areas before eyes; surface densely and finely punctate and semilustrous; anterior margin of clypeus

arcuate and faintly indented in middle; eyes not so large, globular and slightly prominent, ratio of eye

diameter to interocular space 1:5.3; terminal labial palpomeres rounded ax-shaped; terminal maxillary

palpomeres rounded ax-shaped; antennae filiform, barely exceeding elytral midlength; antennomere I

clavate, II short and a little expanded apicad. III to XI subcylindrical, all antennomeres without groove,

ratio of ante nn omere lengths: 16:10:18:21:21:20:20:20:19:17:19.

Pronotum. Subquadrate, about 0.90 times as wide as head, slightly longer than width; anterior and

posterior margins moderately arcuate; lateral margins feebly sinuate, slightly diverging posteriorly;

anterior angles rounded; posterior angles obtuse; dorsum convex in postero-lateral and hollowed

in antero-lateral areas; medio-longitudinal groove distinct in posterior half; surface smooth and

semilustrous. Scutellum triangular with rounded apex.

Elytra. Conjointly about 1.5 times as wide as pronotum, about 3.0 times as long as wide, sides

subparallel; surface densely and finely punctate and semilustrous.

Legs. Moderately slender; femora mostly straight; tibiae mostly straight, with basal part feebly arcuate;

claws simple.

24

HSIAO Y. et al., Revision of the Lycocerus hanatanii species group

Aedeagus (Fig. 4H-J). Ventral process thick, strongly expanded and rounded apically, apex moderately

curved inwards, basal part broad and extending inwards onto ventral side; dorsal plate of each paramere

slightly longer than ventral process, protuberant and narrowed apically, apical margin rounded, with

some hairs on it. Laterophyse very short, with pointed apex, invisible in lateral view. Inner sac swollen,

lengthened behind, somewhat shorter than tegmen.

Body length 5.00-6.00 mm (holotype: 5.60 mm ); width 0.80-1.30 mm (1.10 mm )

Female (Fig. 20)

Similar to male. Body wider than in male. Eyes somewhat smaller than in male, ratio of eye diameter

to interocular space 1:5.4. Antennae distinctly shorter than in male, barely exceeding humeri, ratio of

antennomere lengths: 14.5:10:13.5:15:15:14:14:13 :13 :12:15.

Pronotum. Slightly wider than head, slightly shorter than width. Elytra conjointly about 1.4 times

as wide as pronotum, about 2.8 times as long as wide. Claws simple. Abdominal ventrite VII widely

emarginate on each side of terminal margin, forming subtriangular lateral lobes, median lobe moderately

developed and triangularly concave in middle (Fig. 6G).

Female genitalia (Fig. 6H). Vagina stout, abruptly extended apically as long and thick duct; diverticulum

and spermathecal duct arising from apex of long duct of vagina; diverticulum moderately thin and spiral;

spermathecal duct considerably shorter than diverticulum; spermatheca provided with thin spiral tube,

tube shorter than diverticulum; accessory gland thin, shorter than spermatheca.

Body length 5.50-6.50 mm ; width 1.20-1.50 mm

Remarks

This species is distributed in the southern area of Taiwan at about 1000 m altitude according to the label

data. It was observed being attracted by light at night.

Distribution (Fig. 8)

Taiwan (endemic).

Lycocerus evangelium Hsiao & Okushima sp. nov.

urn:lsid:zoobank.org:act:4AF4EFCB-BB70-44A3-BDD4-5C00B06747AE

Figs 2P-Q, 5A-C, 6I-J, 8

Diagnosis

This species resembles L. araticollis and the nearly black form of L. nigripennis ; the differential

diagnosis is given in Table 3.

Etymology

The specific epithet is derived from the Latin evangelium (“good news”), referring to fact that its

discovery was good news for the team; the specific name is also in memory of the anime “Neon Genesis

Evangelion”, which is one of the greatest animes in Japanese history and had a strong impact on Japanese

popular culture.

Material examined

Holotype

TAIWAN: Meifeng, Nantou, 26 Apr. 2015, Y. Hsiao leg (NMNS).

25

European Journal of Taxonomy 170: 1-33 (2016)

Paratypes

TAIWAN: 1 3,2 same data as for holotype (NMNS); 10 33 , 2 §§, Tsuifeng, Nantou Hsien,

18 Jun. 1989, M. Sato leg. (2 33 , 1 $ in EUMJ; 8 33 , 1 ? in KURA); 1 3 , 2 Tsuifeng, Nantou

Hsien, 24 Jun. 1989, M. Sato leg. (KURA); 1 <f?, 3 §?, Yuanfeng, Nantou Hsien, 24 Jun. 1989,

M. Owada leg. (KURA); 5 33 , 3 §§, Hohuanshan, Nantou Hsien, 25 Jun. 1989, M. Sato leg. (1 3 ,

1 $ in EUMJ; 4 33 , 2 $ $ in KURA); 1 3 , Piluchi, Nantou Hsien, 26 Jun. 1989, M. Sato leg. (KURA);

3 33 , 1 ?, Anmashan, 2600 m, Taichung Hsien, 29 Jul. 1997, B. Tanaka leg. (KURA); 2 §§, Dalu

Forest Trail, Hsinchu, 21 Jun. 2008, L.-P. Hsu leg. (NMNS); 1 3, 1 $, Sungkang, Natou, 31 Mar. 2012,

Y.-T. Wang leg. (YHC); 1 §, Liyuan, Haiduan, Taitung, 19 Apr. 2014, W.-J. Huang (NMNS); 5 §?,

Jianchin Historical Trail, Ilan, 1900 m, 26 Apr. 2014, Y. Hsiao leg. (3 5? in TARI; 2 $$ in YHC);

2 $ ?, Walami Trail, 23.344679° N, 121.201454° E, 862 m, Huangma, Zhuoxi County, Hualian, 28 Feb.

2015, Y.-M. Weng leg. (YHC); 2 33 , 1 ?, Dabaishan, Ilan, 14 Apr. 2015, L. Huang leg. (TARI); 1 $,

Chiarouhu, Ilan, 25 Apr. 2015, F.-C. Hsu leg. (NMNS).

Themus nobilis nobilis

5 14 15

L. maculicollis

16 28 29

p a • L. purpureus

1 23

9 11 19

Ml

DO— L. suture/lus suture//us

11 17 24

oaa— L. adustico/lis

12 23 26

□oc^ L. aegrotus

1 2 6 17 26 28

^□ooooo- L. watanabei

2 13 19 24 30

^C4DOO— L. michiakii

2 10 11 16 30

L. sichuanus

7 16

11

L. vite/Unus

L. yamatensis

n

L. rhagonychiformis

L. nigripennis

L. aurantiacus sp. n.

L. hanatanii

L. ueharaensis

L. pictus

L. yitingi sp. n.

L. evangelium sp. n.

L kintaroi sp. n.

L. araticollis

L. griseopubens

Outgroup

L. maculicollis Group

L. purpureus Group

L. suturellus Group

L. adusticollis Group

L. aegrotus Group

L. oedemerokles Group

L. michiakii Group

L. hickeri Group

L. vitellinus Group

L. lineatipennis Group

L. hanatanii Group

Fig. 9. Phylogenetic reconstruction of the Lycocerus hanatanii species group. Strict consensus tree with

characters mapped on branches using unambiguous character changes in WinClada. Tree length = 72;

Consistency Index = 54; Retention Index =72.

26

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

Other material

TAIWAN: 1 §, Alishan, Chiayi Hsien, 17 May 1968, Y. Watanabe leg. (KURA); 2 55> Mt. Yushan,

Nantou Hsien, 20 May 1968, Y. Watanabe leg. (KURA); 1 8, 3 $$, Mt. Alishan, 27 Jul. 1970,

M. Chujo leg. (1 8 in NHMB; 3 55 in KURA); 1 8, Mt. Alishan, 28 Jul. 1970, M. Chujo leg.

(NHMB); 9 88, 2 §§, Meifeng, Nantou Hsien, 15 Jun. 1976, T. Matsumoto leg. (KURA); 2 55>

Sunglcang-Meifeng, Nantou Hsien, 2 May 1977, W. Suzuki leg. (KURA); 1 8, 1 ?, Tsuifen, Nantou

Hsien, 4 May 1979, S. Tsuyuki leg. (KURA); 1 8, Nanfanshan, Kaoshun [=Kaohsiung], 28 Apr. 1981,

S. Tsuyuki leg. (KURA); 1 8, Lake Yenyanfu, Ilan Hsien, 29 Apr. 1982, N. Ohbayashi leg. (KURA); 1 §,

Kengting Park, Pingtung Hsien, 4 May 1982, N. Ohbayashi leg. (KURA); 1 $, Shyk Shan, near Liu Kui,

S Taiwan, 12 Jun. 1986, K. Baba leg. (KURA); 2 88, 1 ?, Shyk Shan, near Liu Kui, S Taiwan, 28 Jun.

1986, K. Baba leg. (KURA); 4 $ §, Shi Nan Shan, near Liu Kui, S Taiwan, 23 Jul. 1986, K. Baba leg.

(KURA); 2 88, 1 ?, Thu Yun Shan, near Liu Kui, S Taiwan, 28 Jun. 1986, K. Baba leg. (KURA); 1 8,

2 5 J, Hohuanshan, Nan Tow Hsien, M Taiwan, 6 Aug. 1986, K. Baba leg. (KURA); 1 §, Twolin, Ilan

Hsien, 29 Apr. 1987, N. Ohbayashi leg. (KURA); 2 88, Meifeng, 2150 m, M Taiwan, 15 Mar. 1989,

K. Baba leg. (KURA); 6 88, 2 55, 1 ex, Yuanfeng, Nantou Hsien, 24 Jun. 1989, M. Owada leg.

(KURA); 17 88, 2 369 Cottage, Hsueshantungfeng, Taichung Hsien, 30 Jun. 1989, M. Sato leg.

(KURA); 8 88, 9 55» 369 Cottage, Hsueshantungfeng, Taichung Hsien, 2 Jul. 1989, M. Sato leg.

(2 88, 2 55 i n EUMJ; 6 88, 7 55 i n KURA); 1 8, Patungkuan, Hsini, Nantou, 20-22 Jun. 1990,

J.-T. Yang (NMNS); 1 5, 230 Forest trail, Anmashan, Taichung, 3 May 1990, C.-C. Chiang (NMNS);

1 8, Yu-shan West peak, Chiayi, 15 May 1991, C.-C. Chiang (NMNS); 1 5, Mt. Anmashan, Taichung,

2 May 1991, A. Shinohara leg. (KURA); 1 8, Chung-hsueh Shan (Erh-san-ling Lin-tao [=230 Forest

trail]), 2450-2500 m, Tai-an Hsiang, Miao-li Hsien, 24 May 1991, A. Saito leg. (CBM); 1 8, An-ma

Shan, 2160-2300 m, Ho-p’ing, T’aichung Hsien, 25 May 1991, K.-F. Shih leg. (CBM); 2 88, Chia-li

Shan, 2600-2850 m, Tai-an Hsiang, Miao-li Hsien, 27 May 1991, A. Saito leg. (CBM); 1 5, Sungkang,

Nantou Hsien, 1 Jul. 1995, C. Fou leg. (KURA); 2 88, 1 5> Guandoushan, Nantou Hsien, 15 Apr.

1996, C. Fou leg. (KURA); 1 8, Tatachia, Nantou, 5 May 1998, M.-M. Yang & S.-Y Yang (NMNS);

1 8, Tengchih, 1400 m, Taoyuan, Kaohsiung Hsien, 24 Mar. 2002, B. Tanaka leg. (KURA); 11 88,

I 5 5? 369 Cottage, 3100 m, Hoping, Taichung, 7 Jun. 2008, W.-T. Yang leg. (NMNS); 1 8, 1 5> Yakou

Country Inn, 23°16.063' N, 120°58.419' E, 2582 m, Taitung County, 11 Aug. 2008, H. Mendel & MVF

Barclay leg. (BMNH).

Description

Coloration

Body and legs nearly black. Antennae dark brown to black. Mandibles yellowish brown. Body closely

covered with pale pubescence; anterior margin of clypeus, antennae, elytra and legs fringed with pale

bristles.

Male (Fig. 2P)

Head. Nearly as long as wide; vertex faintly hollowed, depressed along apical margin of clypeus and in

lateral areas before eyes; surface densely and finely punctate and semilustrous; anterior margin of clypeus

arcuate and faintly indented in middle; eyes not so large, globular and slightly prominent, ratio of eye

diameter to interocular space 1:5.3; terminal labial palpomeres rounded ax-shaped; terminal maxillary

palpomeres rounded ax-shaped; antennae filiform, extending to elytral midlength; antennomere I clavate,

II short and a little expanded apicad. III to XI subcylindrical, all antennomeres without groove, ratio of

antennomere lengths: 16:10:15.5:20.5:20.5:20.5:20.5:20:20:18.5:23.

Pronotum. Subquadrate, about 0.90 times as wide as head, nearly as long as wide; anterior and posterior

margins moderately arcuate; lateral margins feebly arcuate; anterior angles rounded; posterior angles

obtuse; dorsum convex in postero-lateral and hollowed in antero-lateral areas; medio-longitudinal

groove distinct in posterior half; surface smooth and semilustrous. Scutellum triangular with rounded

apex.

27

European Journal of Taxonomy 170: 1-33 (2016)

Elytra. Conjointly about 1.6 times as wide as pronotum, about 2.6 times as long as wide, sides

subparallel; surface densely and finely punctate and moderately lustrous.

Legs. Moderately slender; femora mostly straight; tibiae mostly straight, with basal part feebly arcuate;

claws simple.

Aedeagus (Fig. 5A-C). Ventral process expanded and rounded apically, apex curved inwards, basal

part broad and extending inwards onto ventral side; dorsal plate of each paramere as long as ventral

process, broad and slightly narrowed apically, apical margin rounded, with some hairs on it, concave on

inner margin. Laterophyse very short, with truncated apex, invisible in lateral view. Inner sac swollen,

lengthened behind, and somewhat shorter than tegmen.

Body length 4.50-5.75 mm (holotype: 5.50 mm ); width 0.80-1.25 mm (1.25).

Female (Fig. 2Q)

Similar to male. Body wider than in male. Eyes smaller than in male, ratio of eye diameter to interocular

space F6.4. Antennae distinctly shorter than in male, barely exceeding humeri, ratio of antennomere

lengths: 13:10:14:16:15:16:14:16:14:13:16.5.

Pronotum. About 0.95 times as wide as head, about 0.95 times as long as wide. Elytra conjointly about

E6 times as wide as pronotum, about 2.6 times as long as wide. Claws simple. Abdominal ventrite VII

widely emarginate on each side of terminal margin, forming subtriangular lateral lobes, median lobe

indistinct in some specimens (Fig. 61).

Female genitalia (Fig. 6J). Vagina stout and rounded, abruptly extended apically as long and thick duct;

diverticulum and spermathecal duct arising from apex of long duct of vagina; diverticulum moderately

thin and spiral, longer than spermathecal duct; spermatheca provided with thin spiral tube, tube slightly

longer than diverticulum; accessory gland thin, shorter than spermatheca.

Body length 4.75-6.50 mm ; width 1.50-2.25 mm

Variation

Lateral sides of pronotum somewhat converging or diverging posteriorly; body lusterless.

Remarks

This species is widely distributed in the Central Mountain Range at mid to high altitude except one

specimen collected from the Renting Park, which is at low altitude. It appears mainly from spring to

summer according to label data.

Distribution (Fig. 8)

Taiwan (endemic).

Lycocerus kintaroi Hsiao & Okushima sp. nov.

um:lsid:zoobank.org:act:FE9AC62A-4B2E-49A0-BD5E-58CD46500D58

Figs 2R, 5D-F, 8

Diagnosis

This species can easily be distinguished from other members of this species group by the black pronotum,

with yellowish fringed margins; black elytra, with very slender yellowish stripe on each humeral side;

aedeagus with dorsal plate broad and subtruncated; apex of laterophyse far from apical margin of dorsal

plate in lateral view.

28

HSIAO Y. et al ., Revision of the Lycocerus hanatanii species group

Etymology

The specific epithet is dedicated to its collector, the late Dr. Kintaro Baba who was one of the great

Japanese entomologists.

Type material

Holotype

TAIWAN: Shi Nan Shan, near Liu Kui, S Taiwan, 5 May 1986, K. Baba leg. (NMNS).

Paratype

TAIWAN: 1 same data as for holotype (EUMJ).

Description

Coloration

Eyes and head black. Antennae dark brown to black. Mandibles yellowish brown. Pronotum black,

fringed with yellowish margins. Elytra black, with very slender yellowish stripe on each humeral side.

Legs black. Pro-, meso- and metasterna, as well as abdomen blackish brown, with margins of abdominal

ventrites yellowish brown. Body closely covered with pale pubescence; anterior margin of clypeus,

antennae, elytra and legs fringed with pale bristles.

Male (Fig. 2R)

Head. Nearly as long as wide; vertex faintly hollowed, depressed along apical margin of clypeus and in

lateral areas before eyes; surface densely and finely punctate and lustrous; anterior margin of clypeus

arcuate and faintly indented in middle; eyes not so large, globular and slightly prominent, ratio of eye

diameter to interocular space 1:5.0; terminal labial palpomeres rounded ax-shaped; terminal maxillary

palpomeres rounded ax-shaped; antennae filiform, extending to elytral midlength; antennomere I clavate,

II short and a little expanded apicad, III to XI subcylindrical, all antennomeres without groove, ratio of

ante nn omere lengths: 16:10:13.5:15.5:15.5:15:14.5:15:14:13 :14.5.

Pronotum. Subquadrate, slightly narrower than head, about 0.95 times as long as wide; anterior and

posterior margins moderately arcuate; lateral margins feebly sinuate; anterior angles rounded; posterior

angles obtuse; dorsum convex in postero-lateral and hollowed in antero-lateral areas; medio-longitudinal

groove distinct in posterior half; surface smooth and lustrous. Scutellum triangular, with rounded apex.

Elytra. Conjointly about 1.3 times as wide as pronotum, about 2.5 times as long as wide, sides sub¬

parallel; surface densely and finely punctate and lustrous.

Legs. Moderately slender; femora mostly straight; tibiae mostly straight, with basal part feebly arcuate;

claws simple.

Aedeagus (Fig. 5D-F). Ventral process moderately expanded and rounded apically, apex curved inwards,

basal part broad and extending inwards onto ventral side; dorsal plate of each paramere as long as

ventral process, broad and subtruncated, apical margin with some hairs on it, concave on inner margin.

Laterophyse very short with obtuse apex, invisible in lateral view. Inner sac swollen, lengthened behind,

somewhat shorter than tegmen.

Body length 5.50-6.00 mm (holotype: 5.50 mm ); width 1.50-1.70 mm (1.50 mm).

Female

Unknown.

29

European Journal of Taxonomy 170 : 1-33 ( 2016 )

Remarks

The right antenna of the holotype is missing.

Distribution (Fig. 8)

Taiwan (endemic).

Discussion

Members of the Lycocerus hanatanii species group have a uniform appearance, with only minor, but

diagnostic differences in the structure of genitalia, which implies that these species possibly went through

a relatively young history of speciation, resulting in morphological divergences that are indistinct. Due

to a lack of recognized synapomorphies in the phylogenetic analysis, the relationship between other

species groups and this species group as well as that among the members of L. hanatanii species group

are not well-resolved. A molecular approach could solve the problem of insufficient synapomorphic

characters and give higher resolution to the relationship within this species group and its phylogenetic

position in Lycocerus.

In view of the greater species diversity, it seems that Taiwan is the center of origin of the L. hanatanii

species group. However, in point of the origin of islands, Taiwan is a continental island, which is

generally believed to have a biota related to the mainland of Eurasia (Creer et al. 2001; Jang-Liaw et

Ishigaki

Iriomote

2 spp.

Fig. 10. Range map of the Lycocerus hanatanii species group and the number of species in each

geographic region.

30

HSIAO Y. et al, Revision of the Lycocerus hanatanii species group

al. 2008; Huang & Lin 2011; Tsai et al. 2014). Furthermore, the taxonomy of the cantharids of China

has not been studied completely yet, and it is possible that more members of this species group will be

discovered and the present conjecture will be challenged or invalidated. On the other hand, according

to known distribution information, this species group exhibits a China-Taiwan-Southern Ryukyus

distribution pattern (Fig. 10). The speciation process of this species group, especially insular species

such as L. hanatanii and L. ueharaensis , is possibly associated with the paleogeographic history of the

East China Sea and the islands of eastern Asia during the Pleistocene glaciation, as in the case of the

grass lizards (Lin et al. 2002). Sea level changed during this period and resulted in the appearance of a

larger continental land area to provide animals the opportunity to migrate from continent to islands, then

some populations were isolated when sea levels rose again, and speciation occurred. Further studies of

historical biogeography are needed and will give insight into the evolutionary history of this species

group.

Acknowledgments

We wish to express our hearty thanks to Dr. Darren Pollock (Eastern New Mexico University, USA)

for his critical reading of the original draft of this paper and correcting our English, and to the late

Dr. Masataka Sato (Nagoya, Japan), Dr. Ming-Luen Jeng (NMNS), Dr. Akiko Saito (CBM), Dr. Chi-

Feng Lee (TARI), Dr. Michael Geiser (BMNH), Dr. Hiroyuki Yoshitomi (Ehime University, Japan),

Mr. Hiroyuki Murakami (Ehime University, Japan), Dr. Zhen-Yu Jin (Yangtze University, China),

Mr. Li Huang, Mr. Yi- Min g Weng, Mr. Tien Hsieh, Mr. Ming Liu, Mr. Chi-Jung Li, Mr. Ren-Hou

Liou, Mr. Wen-Chuan Liao, Mr. Yi-Fing Chung, Mr. Wen-Jean Huang, Mr. Yung-Jen Lu, Mr. Chi-Hao

Wu, Mr. Chao-Pu Cheng, Mr. Shih-Hao Chang and Mr. Yu-Tang Wang for their assistance in carrying

out the phylogenetic analysis or in loaning specimens or providing materials. Our thanks also go to

Dr. Norm Pe nn y (CAS), Ms. Isabelle Ztircher-Pfander (NHMB), Dr. Eckhard Groll (SDEI), Dr. Stephan

Blank (SDEI) and the team of the NMNS for providing photographs, as well as to the Ministry of

Science and Technology project of databasing insect type specimens collected from Taiwan that are

housed abroad through grant MOST101-2631-H-178-003. Besides, we are indebted to the editor and

anonymous reviewers for their suggestions in improving the manuscript.

The present study was supported financially by the Ministry of Science and Technology, Republic of

China, grant MOST104-2815-C-002-057-B.

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Manuscript received: 6 September 2015

Manuscript accepted: 3 November 2015

Published on: 5 January 2016

Topic editor: Koen Martens

Desk editor: Danny Eibye-Jacobsen

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

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