European Journal of Taxonomy 97: 1-29

http://dx.doi.org/10.5852/ejt.2014.97

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

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2014 • Yoshitomi H.

Research article

urn:lsid:zoobank.org:pub:D58FA727-9A10-4023-B178-9E6CCDlEA457

Comparative morphology of the endophallic structures of the genus

Laius (Coleoptera, Melyridae), with the descriptions of three new species

Hiroyuki YOSHITOMI

Ehime University Museum, Bunkyo 3, Matsuyama, Ehime Pref. 790-8577, Japan.

E-mail: [email protected] (corresponding author)

urn:lsid:zoobank.org:author:744FA6FC-2D84-41B7-8A6C-C1611CC68EEl

Abstract. The endophallic structure of the genus Laius is studied and discussed based on the

examination of 19 species from Asia to the Indian Ocean. The structure contains two primary sclerites

(named gonoporal piece and ligula), a secondary sclerite on the basal part of the gonoporal piece (named

additional sclerite) in some species, and a membranous basal area closely covered with many spines

(named spinous area). Five species groups are recognized based on the morphology of the endophallic

sclerites. The sympatric species have different body sizes and quite distinguishable endophallic sclerites

(= different species group), while the allopatric species have overlapping body sizes and similar

endophallic sclerites (= same species group). Three new species are described and six previously known

species are redescribed with endophallic sclerites, and the descriptions of endophallic sclerites of the

remaining ten species are added. The larva of Laius rodriguesensis sp. nov. is also described. The genus

Nossibeus Evers, 1994 is synonymised with Laius Guerin-Meneville, 1830.

Keywords. Melyridae, taxonomy, distribution, endophallus, new species.

Yoshitomi H. 2014. Comparative morphology of the endophallic structures of the genus Laius (Coleoptera,

Melyridae), with the descriptions of three new species. European Journal of Taxonomy 97: 1-29. http://dx.doi.

org/10.5852/eit.2014.97

Introduction

Male genitalia are considered one of the most important diagnostic traits in insect systematics (Simmons

2014), and this also applies to the order Coleoptera (e.g., Sharp & Muir 1912). Some groups in this order

have developed endophallic structures (e.g., sclerites, projections, small denticles), and these characters

provide important information both taxonomically and phylogenetically. The endophallic structures

are mainly observed in relatively large body-sized beetle families (e.g., Carabidae, Cerambycidae,

Chrysomelidae), because it is necessary to evert and inflate the endophallus to observe the structures

(Sasalcawa 2005; Yamasako & Ohbayashi 2011). Therefore, it is difficult to dissect and observe the

endophallic structures in smaller-sized groups.

The melyrid genus Laius Guerin-Meneville, 1830 is represented by about 30 species from the Pacific

Islands, Indian Ocean, Australia, and the eastern coast of Africa (see Appendix); however, numerous

species have been erroneously included in this genus and wifi be transferred to other genera (Plonski

European Journal of Taxonomy 97: 1-29 (2014)

2013; Wittmer 1997). The genus has a well-developed endophallic structure consisting of many small

spines and some large sclerites, and these characters have been observed through the aedeagus (Wittmer

1997; Yoshitomi 2008; also see Fig. 2B). However, the characters could not be well understood or

described.

The main aim of this study is to inspect the validity of endophallic structures for the recognition of

species in this genus. I firstly define the terminology of the endophallic structures of the genus Laius and

secondly describe the endophallic sclerites of 19 species, including the descriptions of three new species

and redescriptions of six poorly known species.

Materials and methods

General observations and dissections were made under a Teica MZ95 stereomicroscope. Microstructures

of dissected parts were studied in pure glycerine under an Olympus BH-2 compound microscope. After

observation, the dissected parts were mounted on the same card with the specimen. Photographs were

taken under a Leica MZ95 using a microscopy camera system (Nikon DS-Fil-L2), and combined with

the automontage software Combine ZM (Alan Hadley, UK). Some structures were observed with an

SEM (Hitachi S-225) after coating with gold.

The endophallus was extracted from the ventral part of the aedeagal apex using a pin and tweezers. If the

endophallus could not be extracted from the aedeagal apex, endophallic sclerites were directly removed

from the endophallus. After observation, the structures were preserved in pure glycerine.

The holotypes and paratypes are preserved in the Ehime University Museum, Matsuyama (EUMJ),

Naturhistorisches Museum Wien (NMW), Natural History Museum Basel (NHMB) and Bishop

Museum, Hawaii (BPBM).

Abbreviations

Morphological abbreviations used for measurements are as follows (also see Fig. 2):

AL =

aedeagal length

EL =

length of elytra in suture

EW =

maximum width of elytra

GL =

length of gonoporal piece

HL =

head length

HW =

head width across eyes

length

LL =

length of ligula

LW =

width of ligula

PL =

length of pronotum in median line

PW =

maximum width of pronotum

TL =

total length (HL+PL+EL)

W =

width.

The average is given in parentheses after the range.

Results

Characteristics and terminology of endophallic structures

In the copulation position, only the endophallus of the male genitalia is engaged with the female structures

(Fig. 1). Endophallic structures (Figs 1-2) consist of a membranous basal area (named spinous area),

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

two primary sclerites (named gonoporal piece and ligula), and a secondary sclerite in some species

(named additional sclerite).

Spinous area (Figs 2A, 12D): membranous basal area, closely covered with many short spines (Fig. 12D),

about 0.5 x length of AF. The differences in this character between species cannot be observed, and the

comparison and description of each species is not given in this study.

Gonoporal piece (Fig. 2C; gp in Figs 3—4): longest sclerite, about 0.6 x length of AF, straight to rather

curved, with pointed or rounded apex, closely covered with minute spines in basal portion of some

species, projecting apically and straight from the apical portion of spinous area.

Figula (Fig. 2D; / in Figs 3-4): straight to C-shaped curved sclerite, with pointed apex, shorter than

gonoporal piece, projecting laterally from the apico-lateral portion of spinous area.

Additional sclerite (as in Figs 3-4): secondary sclerite situated in basal part of gonoporal piece, shorter

than ligula, projecting laterally from the basal portion of gonoporal piece.

Taxonomy

Class Hexapoda Blainville, 1816

Order Coleoptera Finnaeus, 1758

Superfamily Cleroidea Fatreille, 1802

Family Melyridae Feach, 1815

Subfamily Malachiinae Fleming, 1821

Genus Laius Guerin-Meneville, 1830

Laius Guerin-Meneville, 1830: 78 [Type species: Laius cyaneus Guerin-Meneville, 1830].

Megadeuterus Westwood, 1833: 678 [Type species: Paussusflavicornis Fabricius, 1801],

Rotamalachius Kono, 1937 in Kono & Esaki 1937: 121 [Type species: Rotamalachius esakii Kono, 1937].

Nossibeus Evers, 1994: 173 [Type species: Laiuspolitus Fairmaire, 1880]. syn. nov.

Remarks

The genus Nossibeus was erected for Laius politus Fairmaire, 1880 (Evers 1994). After observation of

this species, including its endophallic structure, it was clear that this species cannot be recognised as

another species of the genus Laius. In the present paper, Nossibeus Evers, 1994 is treated as a junior

synonym of Laius Guerin-Meneville, 1830 (syn. nov.).

Laius baliensis sp. nov.

urn:lsid:zoobank.org:act:9D51A886-E14B-4DE6-96C3-410AF3CBC669

Figs 1, 3A, 5A-B, 7A, 8A, 9A, 13A-B

Laius sp. - Yoshitomi 2008: 289.

Diagnosis

Body small, TF = 4.10^1.46 mm in male, 4.35M.75 mm in female; male antennomere III oval, deeply

concave in dorso-mesal portion, F/W = 0.95; aedeagal apex rounded; gonoporal piece straight and

slender; ligula slightly curved; additional sclerite absent.

Etymology

After the type locality, Bali Island, Indonesia.

European Journal of Taxonomy 97 : 1-29 ( 2014 )

Type material

Holotype

S (EUMJ), “Nusa Dua, South Island 8°48’6.81”S 115°14’16.07”E, Bali, INDONESIA 8. VIII. 2010

H. Yoshitomi leg.”.

Paratypes

1 s, 6 $ $ (EUMJ), same data as for the holotype; 6 SS, 10 ? ? (EUMJ, NMW, BPBM, NHMB), “Nusa

Dua, North Island 8°47 , 52.72”S 105°14’9.31”E, Bali, INDONESIA 7-8. VIII. 2010 H. Yoshitomi

ovipositor (F)

1.0 mm

aedeagus (M)

gonoporal piece (M)

0. 5 mm

spinous area (M)

stylus (F)

“W 8

■

7 ,

...■iv

■t

I iguI a (M)

aedeagal apex (M)

Fig. 1. Copulation of Laius baliensis sp. nov. A. Endophallus fully engaged with female structures in

dorsal view. B. Endophallus fully engaged with female structures in dorsal view, close up. Abbreviations:

M = male, F = female.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

leg.”; 3 $ $ (EUMJ), “Nusa Dua, Bali Indonesia 5-1-2000 H. Yoshitomi leg.”; 2 $ ? (EUMJ), ditto but

collector “M. Sato”; 1 $ (EUMJ), ditto but collector “N. Ohbayashi”.

Description

Male

Body oblong, shining, densely covered with short black setae. Coloration of body black with strong bluish

luster on head, pronotum and elytra; labrum, maxillary palpomere III and tarsi brown; antennomeres

I—III and maxillary palpomeres I—II yellowish-orange.

Head slightly narrower than pronotum, densely punctate; vertex flat dorsally; eyes large, strongly

prominent; HL/HW 0.73-0.82 (0.78). Antennae (Fig. 7A) stout; antennomere III (Fig. 8A) oval, deeply

concave in dorso-mesal portion, L/W = 0.95; approximate ratio of each antennomeres (n = 1) as 8.0 :

1.0 : 6.0 : 2.2 : 2.0 : 1.8 : 1.8 : 1.8 : 2.0 : 1.8 : 2.8. Pronotum quadrate, widest near anterior margin,

rounded and obscure in antero- and postero-lateral angles, punctate as in head; PW/PL 1.15-1.44 (1.26).

Scutellum semicircular, covered with very fine punctures. Elytra oblong, subparallel-sided near base

to apical 1/4; EL/EW 1.33-1.71 (1.50); EL/PL 2.50-2.89 (2.65); EW/PW 1.30-1.52 (1.41); TL/EW

2.33-2.93 (2.58). Legs relatively long and slender. Caudal margin of tergite VIII shallowly concave.

Aedeagus (Fig. 9A) about 1.12 mm, round at aedeagal apex. Gonoporal piece (Fig. 3A) straight, rather

wide, 0.61 mm in GL; GL/AL 2.39. Ligula (Fig. 3A) short, curved in basal 1/3; LL 0.23 mm; LW

0.06 mm; LL/LW 3.73; GL/LL 2.68.

Female

Similar to male in general appearance; HL/HW 0.71-0.79 (0.76); PW/PL 1.20-1.36 (1.27); EL/

EW 1.53-1.68 (1.59); EL/PL 2.65-2.93 (2.82); EW/PW 1.31-1.44 (1.39); TL/EW 2.60-2.82 (2.68).

Antennae (Fig. 7A) stout; antennomere III short, L/W = 1.88; approximate ratio of each antennomeres

(n = 1) as 8.8 : 1.0 : 4.2 : 2.4 : 2.2 : 2.2 : 2.0 : 1.8 : 2.0 : 2.0 : 3.0.

Measurements

Male (n = 7): TL 4.10M.46 (4.27) mm; HL 0.80-0.90 (0.85) mm; HW 1.00-1.20 (1.10) mm; PW 1.08-

1.30 (1.18) mm; PL 0.90-1.00 (0.94) mm; EL 2.40-2.60 (2.48) mm; EW 1.40-1.82 (1.67) mm Female

(n = 7): TL 4.35—4.75 (4.61) mm; HL 0.85-0.95 (0.90) mm; HW 1.15-1.25 (1.20) mm; PW 1.20-1.30

(1.24) mm; PL 0.90-1.00 (0.97) mm; EL 2.60-2.80 (2.73) mm; EW 1.60-1.80 (1.72) mm

Distribution

Only known from the type locality, Indonesia (Bali Isl.).

Remarks

This species is similar to Laius flavicornis (Fabricius, 1801) and L. sabangensis Wittmer, 1985 in the

shape of male antennomere III, and differs from them in the shape of endophallic sclerites and smaller

body length.

Biological notes

This species was collected together with Laius pankowi Wittmer, 1999 and L. satoi Yoshitomi, 2008 at

the same locality. Based on the data of Yoshitomi (2008) and the present study, Laius baliensis sp. nov.

was collected in January and August, L. pankowi in January and December, and L. satoi in January,

April, and August.

European Journal of Taxonomy 97 : 1-29 ( 2014 )

Laius andamanensis sp. nov.

urn:lsid:zoobank.org:act:AB272070-CAEA-4964-8CFD-A219E93222Fl

Figs 3B, 5C-D, 7B, 8B, 9B, 12

Diagnosis

Male antennomere III distinctly projecting laterally, shallowly concave in dorso-mesal portion, F/W =

0.70; aedeagal apex concave; gonoporal piece straight and slender; ligula long and slightly curved;

additional sclerite absent.

Etymology

After the type locality, Andaman Islands, India.

Type material

Holotype

S (NHMB), “ANDAMAN Is.: Havelock I. env. of village No. 4, 11°59’N, 92 0 58’E, 22. iv-14. V. 1998,

Karel & Simon Majer leg.”.

Fig. 2. Aedeagus of Laius lutaoensis Yoshitomi & Fee, 2010, showing the terminology and abbreviations

for measurement of endophallic structures. A. Aedeagus with extracted endophallus. B. Aedeagus with

holding endophallus (from Yoshitomi & Fee 2010). C. Gonoporal piece. D. Figula.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Fig. 3. Endophallic sclerites of Laius spp. A. Laius baliensis sp. nov. (paratype). B. L. andamanensis sp. nov.

(paratype). C. L. rodriguesensis sp. nov. (paratype). D. L. maai Wittmer, 1973 (paratype). E. L. marchei

Pic, 1922. F. L. politus Fairmaire, 1880. G. L. sabangensis Wittmer, 1985 (paratype). H. L. sericatus

Champion, 1924.1. L. submar in#brmis Wittmer, 1985 (paratype). J. L. asahinai Nakane, 1955.

European Journal of Taxonomy 97 : 1-29 ( 2014 )

Fig. 4. Endophallic sclerites of Laius spp. A. Laius etsukoae Sato, Yoshitomi & Ohbayashi, 2006

(paratype). B. L. flavicornis (Fabricius, 1801). C. L. keiichii Sato, Yoshitomi & Ohbayashi, 2006

(paratype). D. L. lutaoensis Yoshitomi & Lee, 2010 (paratype). E. L. madli Yoshitomi, 2010 (paratype).

F. L. miyamotoi Nakane, 1955. G. L. satoi Yoshitomi, 2008. H. L. taiwanus Yoshitomi & Lee, 2010

(paratype). I. L. pankowi Wittmer, 1999.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Paratypes

22 10?? (NHMB, EUMJ), same data as for the holotype.

Description

Male

Body oblong, shining, densely covered with short black setae. Coloration of body black with dully bluish

luster on elytra; labrum, maxillary palpomere I, and tarsi brown; antennomeres I—III yellowish-orange.

Head narrower than pronotum, very densely punctate; vertex slightly convex dorsally; eyes relatively

large, prominent; HL/HW 0.76-0.90 (0.81). Ante nn ae (Fig. 7B) long; antennomere III (Fig. 8B)

transversal trapezoidal, distinctly projecting laterally, shallowly concave in dorso-mesal portion, L/W

= 0.70; approximate ratio of each antennomeres (n = 1) as 8.3 : 1.0 : 5.0 : 2.0 : 2.2 : 2.0 : 2.0 : 2.0 :

2.0 : 2.0 : 3.5. Pronotum quadrate, widest near anterior margin, rounded and obscure in antero- and

postero-lateral angles, punctate as in head; PW/PL 1.18-1.33 (1.24). Scutellum semicircular, covered

with fine punctures. Elytra oblong, widest at apical 1/4; sides gradually expanded postero-laterally; EL/

EW 1.36-1.55 (1.45); EL/PL 2.52-2.88 (2.67); EW/PW 1.38-1.55 (1.48); TL/EW 2.30-2.56 (2.44).

Legs relatively long and slender; fore tibia (Fig. 12A) enlarged, deeply excavate. Caudal margin of

tergite VII shallowly concave.

Aedeagus (Fig. 9B) about 1.13 mm, concave at aedeagal apex (Fig. 12C). Gonoporal piece (Fig. 3B)

straight, slender, 0.74 mm in GL; GL/AL 2.39. Ligula (Fig. 3B) relatively long, slightly curved; LL

0.27 mm; LW 0.11 mm; LL/LW 2.58; GL/LL 2.71.

Female

Similar to male in general appearance. HL/HW 0.75-0.80 (0.79); PW/PL 1.18-1.30 (1.23); EL/

EW 1.36-1.48 (1.41); EL/PL 2.44-2.73 (2.60); EW/PW 1.45-1.58 (1.51); TL/EW 2.30-2.43 (2.36).

Antennae (Fig. 7B) long; antennomere III (Fig. 12B) distinctly expanded laterally, shallowly concave in

dorsal portion, L/W = 2.42; approximate ratio of each antennomeres (n = 1) as 8.0 : 1.0 : 4.4 : 2.2 : 2.4 :

2.2 : 2.2 : 2.0 : 2.0 : 2.0 : 3.6

Measurements

Male (n = 20): TL 5.52-6.33 (5.79) mm; HL 1.00-1.15 (1.05) mm; HW 1.22-1.38 (1.30) mm; PW 1.48-

1.75 (1.61) mm; PL 1.20-1.48 (1.29) mm; EL 3.28-3.75 (3.45) mm; EW 2.18-2.70 (2.38) mm. Female

(n = 6): TL 4.43^1.96 (4.67) mm; HL 0.75-0.85 (0.81) mm; HW 1.00-1.08 (1.03) mm; PW 1.20-1.42

(1.32) mm; PL 1.00-1.20 (1.07) mm; EL 2.60-3.00 (2.79) mm; EW 1.85-2.15 (1.98) m m

Distribution

Only known from the type locality, India (Andaman Islands).

Remarks

This is distinct species in having transversal antennomere III of male.

Laius rodriguesensis sp. nov.

urn:lsid:zoobank.org:act:5ACC9D36-0154-4FE3-9D27-D5AAF61A9143

Figs 3C, 5E-F, 7C, 8C, 9C, 10, 11

Diagnosis

Male antennomere III transverse, deeply concave in dorso-mesal portion, L/W= 0.74; aedeagal apex

rounded; gonoporal piece slightly curved; ligula strongly curved; additional sclerite absent; coloration

of female ante nn omeres I—III black or dark brown.

European Journal of Taxonomy 97: 1-29 (2014)

Etymology

After the type locality, Rodrigues Island, Republic of Mauritius.

Type material

Holotype

S (NMW), “RODRIGUES Bucht zw. Trou d’Argent und Anse Bouteille intertidal zone 12.-13.4.11, leg.

M. Madl”.

Fig. 5. Dorsal habitus of Laius spp. of $ (holotypes A, C, E) and $ $ (paratypes B, D, F). A-B. Laius

baliensis sp. nov. C-D. L. andamanensis sp. nov. E-F. L. rodriguesensis sp. nov. Scale bars = E0 mm.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Paratypes

3 $ $ (NMW), same data as for the holotype; 1 5 (NMW), “MAURITIUS Rodrigues, Latanier Riviere

Cascade Victoire 7.5.08. leg. M. Madl”; 2 $ ? (NMW), “MAURITIUS Rodrigues Anse Fumier 2.4.09,

leg. M. Madl”; 4 SS, H ? ? (NMW, EUMJ), ditto but “2.5.08”.

Description

Male

Body oblong, shining, densely covered with short black setae. Coloration of body black with dully bluish

luster on elytra; antennomeres I—III yellowish-orange, but anterior margin of antennomere I infiiscate.

Head narrower than pronotum, very densely punctate; vertex slightly depressed dorsally; eyes relatively

large, prominent; HL/HW 0.72-0.89 (0.78). Antennae (Fig. 7C) long; antennomere III (Fig. 8C)

transverse, deeply concave in dorso-mesal portion, F/W = 0.74; approximate ratio of each antennomeres

(n = 1) as 11.2 : 1.0 : 6.8 : 3.2 : 2.6 : 2.2 : 2.2 : 2.4 : 2.0 : 2.2 : 4.0. Pronotum quadrate, widest

near anterior margin, rounded and obscure in antero- and postero-lateral angles; punctate as in head;

PW/PF 1.33-1.49 (1.39). Scutellum semicircular, finely punctate. Elytra oblong, widest at apical 1/3;

sides weakly expanded postero-laterally; EF/EW 1.31-1.50 (1.43); EF/PF 2.62-2.76 (2.69); EW/PW

1.31-1.42 (1.36); TF/EW 2.18-2.52 (2.42). Fegs relatively long and slender. Caudal margin of tergite

VII shallowly concave. Aedeagus (Fig. 9C) about 1.17 mm, round at aedeagal apex. Gonoporal piece

(Fig. 3C) slightly curved, 0.63 mm in GF; GF/AF 1.81. Figula (Fig. 3C) long, strongly curved in half;

FF 0.30 mm; FW 0.11 mm; FF/FW 2.84; GF/FF2.11.

Female

Similar to male in general appearance; bluish luster on elytra weak; coloration of antennomeres I—III

black or dark brown; HF/HW 0.74-0.95 (0.84); PW/PF 1.32-1.42 (1.37); EF/EW 1.29-1.47 (1.39);

EF/PF 2.47-2.86 (2.68); EW/PW 1.30-1.48 (1.41); TF/EW 2.25-2.47 (2.39). Antennae (Fig. 1C) long;

ante nn omere III short, F/W = 2.16; approximate ratio of each ante nn omeres (n = 1) as 8.6 : 1.0 : 4.4 :

2.0 : 2.0 : 1.8 : 1.8 : 2.0 : 2.0 : 2.0 : 3.8.

Measurements

Male (n = 5): TF 5.50-5.75 (5.64) mm; HE 1.00-1.20 (1.08) mm; HW 1.30-1.45 (1.38) mm; PW 1.60-

1.83 (1.72) mm ; PF 1.20-1.30 (1.24) mm ; EF 3.20-3.40 (3.32) mm ; EW 2.20-2.60 (2.33) mm Female

(n = 10): TF 4.10—4.68 (4.41) mm; HE 0.80-1.00 (0.88) mm; HW 1.00-1.10 (1.05) mm; PW 1.20-1.38

(1.32) mm; PF 0.90-1.00 (0.96) mm; EF 2.35-2.80 (2.57) mm; EW 1.70-2.00 (1.85) mm.

Larvae

Coloration of head capsule and urogomphi dark brown; thoracic and abdominal segments and legs

cream. Head capsule (Fig. 10B) suboblong, F/W =1.3; four pairs of stemmata situated near antero¬

lateral corners. Antennae missing. Fabrum (Fig. 11B) with a pair of long lateral and five pairs of short

mesal setae in anterior portion; anterior margin of epipharynx bearing short stout setae. Maxillae

(Fig. 11 A) bearing long setae in antero-lateral portion. Mandibles (Fig. 11C) with two apical teeth; mola

with two short teeth. Fabium (Figs 10C, 11 A) relatively wide, with 2-segmented palpi. Thorax wealdy

sclerotized, widest at posterior part of metathorax; pronotum with two pairs of longitudinal sclerotized

bands in mesal portion, bearing short setae in lateral and mesal portions. Abdomen (Fig. 10A) weakly

sclerotized, subparallel-sided, bearing short setae in lateral and postero-mesal portions. Urogomphi

(Fig. 10D-F) subparallel-sided, closely covered with long setae, simply bifurcate in apical 1/2, upturned

in apical portions, pointed at apices, with some very long hairy setae in apical parts. Fegs relatively long.

Additional material examined

2 Farvae (NMW, in ethanol), “MAURITIUS Rodrigues Anse Fumier 10.05.2008, leg. M. Madl”.

European Journal of Taxonomy 97: 1-29 (2014)

Distribution

Only known from the type locality. Republic of Mauritius (Rodrigues Isl.).

Remarks

This species is closely related to Laius madli Yoshitomi, 2010 known from Mauritius, but differs from the

latter in the following characteristics: 1) ligula strongly curved in middle portion (rather weakly curved

Fig. 6. Dorsal habitus of Laius spp. (left: right: §). A. Laius maai Wittmer, 1973 (paratype).

B. L. marchei Pic, 1922. C. L. politus Fairmaire, 1880. D. L. sabangensis Wittmer, 1985 (paratype).

E. L. sericatus Champion, 1924. F. L. submariniformis Wittmer, 1985 (paratype). Scale bars = 1.0 mm.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

basally in madli)] 2) male antennomere III projecting laterally, transverse, LAV = 0.7 (antennomere III

slightly projecting laterally, L/W = 0.9 in madli).

The larvae of two Japanese species, L. asahinai Nakane, 1955 and L. miyamotoi Nakane, 1955, were

described by Sato (1964) and Asano & Kojima (2009). The larva of L. rodrigesensis sp. nov. is similar to

those of the aforementioned species, and differs from them in that the teeth of the mola are rather short,

and that the head is narrower.

Laius maai Wittmer, 1973

Figs 3D, 6A, 7D, 8D, 9D

Laius maai Wittmer, 1973: 336.

Type material

Paratypes 6 cJcJ, 2 $ ?, (BPBM), “New Guinea: NE Sarmi, W Hollandia”, “20.-23.7.1959 T. C. Maa”.

Redescription

Male

HL/HW 0.82; PW/PL 1.36; EL/EW 1.51; EL/PL 2.95; EW/PW 1.43; TL/EW 2.46. Antennae (Fig. 7D)

long; antennomere III (Fig. 8D) oblong, widened in basal 1/3, L/W = 2.08; approximate ratio of each

antennomeres (n= 1) as 7.4 : 1.0 : 3.6 : 2.1 : 1.9 : 1.9 : 1.7 : 1.6 : 1.7 : 1.9 : 3.1. Caudal margin of tergite

VII concave. Aedeagus (Fig. 9D) about 1.32 mm, concave at aedeagal apex. Gonoporal piece (Fig. 3D)

straight, slender, with long and straight basal projection, 0.77 mm in GL; GL/AL 2.01. Ligula (Fig. 3D)

relatively long, slightly curved; LL 0.29 mm; LW 0.09 mm; LL/LW 3.06; GL/LL 2.65.

Female

HL/HW 0.78-0.81 (0.80); PW/PL 1.31-1.37 (1.33); EL/EW 1.42-1.66 (1.50); EL/PL 2.70-2.96 (2.83);

EW/PW 1.35-1.46 (1.41); TL/EW 2.39-2.67 (2.48). Ante nn ae (Fig. 7D) long; ante nn omere III short,

L/W= 1.88; approximate ratio of each antennomeres (n= 1) as 11.3 : 1.0 : 5.3 : 2.8 : 3.0 : 3.0 : 3.0 : 2.5 :

2.8 : 2.5 : 4.5.

Measurements

Male (n = 1): TL 5.05 mm; HL 0.90 mm ; HW 1.10 mm ; PW 1.43 mm; PL 1.05 mm ; EL 3.10 mm; EW

2.05 mm. Female (n = 4): TL 4.50^1.68 (4.58) mm; HL 0.80-0.85 (0.82) mm; HW 1.00-1.05 (1.03) mm;

PW 1.30-1.33 (1.31) mm; PL 0.95-1.00 (0.98) mm; EL 2.70-2.90 (2.78) mm; EW 1.75-1.90 (1.85) mm

Distribution

New Guinea.

Laius marchei Pic, 1922

Figs 3E, 6B, 7E, 8E, 9E

Laius marchei Vic, 1922: 158.

Rotamalachius esakii Kono, 1937 in Kono & Esaki 1937: 121. Synonymised by Nakane (1955).

Laius marchei - Wittmer 1958: 72 [redescription]. — Sato et al. 2006: 344.

Material examined

56 SS, 14 $$ (EUMJ), “(MARIANA Is.) Teteto Beach Rota I. 22. Jan. 1980 M. Tomokuni”; 1 $

(EUMJ), “Is. Rota mariana VIII. 8. 1940 D. Matsusita”, “ Laius marchei Pic det. W. Wittmer”; 1 S

European Journal of Taxonomy 97: 1-29 (2014)

(EUMJ), “(Rota Tatochio) South Sea Islands of Nippon: 8-VIII. 1940 Coll. Dengo Matsusita”; 2

1 $ (BPBM), “Tumon Bay, Guam 24 June 1969”, “A. A. LaPlante Collection BISHOP Museum Ace.

#1977.84”.

Redescription

Male

HL/HW 0.71-0.93 (0.79); PW/PL 1.21-1.36 (1.28); EL/EW 1.28-1.48 (1.42); EL/PL 2.42-2.86 (2.68);

EW/PW 1.41-1.58 (1.48); TL/EW 2.25-2.44 (2.36). Ante nn ae (Fig. 7E) long; antennomere I long and

curved laterally, with small projection in lateral margin; antennomere III (Fig. 8E) trapezoidal, deeply

concave in dorso-mesal portion, L/W = 0.76; approximate ratio of each antennomeres (n = 1) as 11.8 :

1.0 : 7.0 : 2.6 : 2.0 : 2.2 : 2.2 : 2.4 : 2.4 : 2.4 : 3.8. Caudal margin of tergite VII concave. Aedeagus

Fig. 7. Left antennae of Laius spp. (above: male; below: female). A. Laius baliensis sp. nov. (paratype).

B. L. andamanensis sp. nov. (paratype). C. L. rodriguesensis sp. nov. (paratype). D. L. maai Wittmer,

1973 (paratype). E. L. marchei Pic, 1922. F. L. politus Fairmaire, 1880. G. L. sabangensis Wittmer,

1985 (paratype). H. L. sericatus Champion, 1924. I. L. submariniformis Wittmer, 1985 (paratype).

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

(Fig. 9E) about E28 mm, concave at aedeagal apex. Gonoporal piece (Fig. 3E) straight, slender, with

long and straight basal projection, 0.80 mm in GL; GL/AL 1.91. Ligula (Fig. 3E) long and slender,

strongly curved in apical half; FF 0.33 mm; FW 0.17 mm; FF/FW 2.81; GF/FF 2.44.

Female

HF/HW 0.72-0.87 (0.80); PW/PF 1.13-1.32 (1.24); EF/EW 1.43-1.55 (1.46); EF/PF 2.64-2.96 (2.82);

EW/PW 1.52-1.67 (1.57); TF/EW 2.34-2.45 (2.38). Antennae (Fig. 7E) long; antennomere III short,

F/W = 2.25; approximate ratio of each antennomeres (n = 1) as 9.4 : 1.0 : 5.0 : 2.4 : 2.8 : 2.4 : 2.4 : 2.4 :

2.4 : 2.4 : 4.0.

Measurements

Male (n = 15): TF 4.90-6.75 (5.36) mm; HE 0.80-1.40 (0.94) mm; HW 1.10-1.50 (1.19) mm; PW 1.35-

1.90 (1.53) mm; PF 1.10-1.50 (1.20) mm; EF 2.90-3.85 (3.21) mm; EW 2.10-3.00 (2.27) mm. Female

(n = 6): TF 5.15-5.73 (5.37) mm; HE 0.85-1.00 (0.92) mm; HW 1.10-1.25 (1.15) mm; PW 1.35-1.50

(1.44) mm; PF 1.10-1.30 (1.17) mm; EF 3.15-3.43 (3.28) mm; EW 2.20-2.40 (2.25) mm

Distribution

Mariana Isis. (Rota, Guam).

Remarks

This is the first record of this species from Guam, Mariana Islands.

Laius politus Fairmaire, 1880

Figs 3F, 6C, 7F, 8F, 9F

Laius politus Fairmaire, 1880: 246.

Nossibeus politus - Evers 1994: 173 [seepage 3].

Material examined

18 SS, 40 (NHMB, EUMJ), “MADAGASCAR Katsepy (Majunga) 24-31. xii. 1977 P. Pacholatko

leg”.

Redescription

Male

HF/HW 0.69-0.94 (0.80); PW/PF 1.33-1.57 (1.43); EF/EW 1.39-1.52 (1.46); EF/PF 2.82-3.40 (3.05);

EW/PW 1.37-1.59 (1.46); TF/EW 2.24-2.51 (2.36). Antennae (Fig. 7F) long; antennomere I stout;

antennomere III (Fig. 8F) elongate, widened in apical portion, F/W = 3.08; approximate ratio of each

antennomeres (n = 1) as 8.4 : 1.0 : 7.4 : 2.2 : 2.0 : 2.0 : 2.2 : 2.0 : 2.2 : 2.0 : 3.8. Caudal margin of tergite

VII shallowly concave. Aedeagus about 1.18 mm, round at aedeagal apex. Gonoporal piece (Fig. 3F)

curved, rather wide, 0.54 mm in GF; GF/AF 2.45. Figula (Fig. 3F) short, evenly curved; FF 0.19 mm;

FW 0.09 mm; FF/FW 2.13; GF/FF 2.88.

Female

HF/HW 0.64-0.90 (0.79); PW/PF 1.38-1.47 (1.42); EF/EW 1.30-1.55 (1.44); EF/PF 2.80-3.10 (2.96);

EW/PW 1.37-1.54 (1.45); TF/EW 2.17-2.50 (2.35). Ante nn ae (Fig. 7F) long; ante nn omere III long and

slender, F/W = 3.1; approximate ratio of each antennomeres (n = 1) as 8.0 : 1.0 : 5.2 : 2.4 : 2.0 : 2.2 :

2.0 : 2.2 : 2.2 : 2.2 : 3.4.

European Journal of Taxonomy 97: 1-29 (2014)

Measurements

Male (n = 17): TL 5.15-5.92 (5.55) mm; HL 0.90-1.13 (1.00) mm; HW 1.18-1.33 (1.25) mm; PW

1.50-1.75 (1.61) mm; PL 1.00-1.20 (1.13) mm; EL 3.10-3.62 (3.42) mm; EW 2.15-2.60 (2.35) mm

Female (n = 10): TL 4.05-5.00 (4.48) mm; HL 0.70-0.90 (0.81) mm; HW 0.93-1.10 (1.02) mm; PW

1.18-1.38 (1.32) mm; PL 0.85-1.00 (0.93) mm; EL 2.50-3.10 (2.75) mm; EW 1.62-2.00 (1.92) mm

Distribution

Madagascar.

Laius sabangensis Wittmer, 1985

Figs 3G, 6D, 7G, 8G, 9G

Laius sabangensis Wittmer, 1985: 386.

Fig. 8. Male antennomere III of Laius spp. A. Laius baliensis sp. nov. (paratype). B. L. andamanensis

sp. nov. (paratype). C. L. rodriguesensis sp. nov. (paratype). D. L. maai Wittmer, 1973 (paratype).

E. L. marchei Pic, 1922. F. L. politus Fairmaire, 1880. G. L. sabangensis Wittmer, 1985 (paratype).

H. L. sericatus Champion, 1924. I. L. submariniformis Wittmer, 1985 (paratype).

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Material examined

Holotype, S (EUMJ), “PHILIPPINES Sabang, N. of Mt. St. Paul, Palawan Julu, 11-13, 1977 M. Sato

leg.”; 2 $ $ paratypes, 9 SS, 6 $ $ (EUMJ), same data as for the holotype.

Redescription

Male

HL/HW 0.84-0.91 (0.88); PW/PL 1.10-1.22 (1.16); EL/EW 1.38-1.50 (1.43); EL/PL 2.53-2.67 (2.59);

EW/PW 1.50-1.62 (1.56); TL/EW 2.37-2.60 (2.48). Antennae (Fig. 7G) stout and short; antennomere

I stout; antennomere III (Fig. 8G) trapezoidal, deeply concave in dorso-mesal portion, L/W = 0.86;

approximate ratio of each antennomeres (n = 1) as 6.6 : 1.0 : 6.0 : 2.4 : 2.0 : 2.2 : 2.0 : 2.2 : 2.0 : 2.2 :

3.6. Caudal margin of tergite VII straight. Aedeagus (Fig. 9G) about 0.94 mm, concave at aedeagal apex.

Gonoporal piece (Fig. 3G) straight, slender, 0.72 mm in GL; GL/AL 1.87. Ligula (Fig. 3G) long and

winding; LL 0.41 mm; LW 0.06 mm; LL/LW 7.30; GL/LL 1.77. Additional sclerite (Fig. 3G) present,

0.10 mm

Fig. 9. Aegeagi of Laius spp. A. Laius baliensis sp. nov. (paratype). B. L. andamanensis sp. nov.

(paratype). C. L. rodriguesensis sp. nov. (paratype). D. L. maai Wittmer, 1973 (paratype). E. L. marchei

Pic, 1922. F. L. politus Fairmaire, 1880. G. L. sabangensis Wittmer, 1985 (paratype). H. L. sericatus

Champion, 1924. I. L. submariniformis Wittmer, 1985 (paratype).

European Journal of Taxonomy 97: 1-29 (2014)

Female

HL/HW 0.79-0.86 (0.81); PW/PL 1.16-1.21 (1.17); EL/EW 1.42-1.47 (1.45); EL/PL 2.61-2.78 (2.70);

EW/PW 1.57-1.62 (1.59); TL/EW 2.36-2.47 (2.42). Ante nn ae (Fig. 7G) short; antennomere III short,

L/W = 2.5; approximate ratio of each antennomeres (n = 1) as 8.0 : E0 : 5.4 : 2.0 : 2.2 : 2.4 : 2.6 : 2.2 :

2.2 : 2.2 : 3.4.

Measurements

Male (n = 4): TL 4.03^1.73 (4.23) mm; HL 0.80-0.98 (0.85) mm; HW 0.90-E08 (0.97) mm; PW 1.02-

E20 (E09) mm; PL 0.88-1.05 (0.94) mm; EL 2.35-2.70 (2.44) mm; EW 1.57-1.90 (1.71) mm. Female

(n = 4): TL 4.18-4.30 (4.23) mm; HL 0.75-0.80 (0.76) mm; HW 0.92-0.95 (0.94) mm; PW 1.05-E15

(E10) mm; PL 0.90-0.95 (0.94) mm; EL 2.48-2.60 (2.53) mm; EW 1.70-1.82 (1.75) mm

Distribution

Philippines (Palawan).

Laius sericatus Champion, 1924

Figs 3H, 6E, 7H, 8H, 9H

Laius sericatus Champion, 1924: 30E

Material examined

2 (NMW), “SEYCHELLES: Mahe Anse aux Coubes 23.03.2011 leg. M. Madl”; 1 S, 1 ? (NMW),

ditto but “21.03.2011”; 3 S3, 2 ?? (1 ? in ethanol; NMW, EUMJ), “SEYCHELLES: Mahe Petite

Police intertidal 28.10.2010, leg. M. Madl”.

Redescription

Male

HL/HW 0.74-0.75 (0.75); PW/PL 1.27-1.31 (E29); EL/EW E40-1.52 (E47); EL/PL 2.90-3.04 (2.95);

EW/PW 1.54-1.58 (1.56); TL/EW 2.27-2.42 (2.37). Antennae (Fig. 7H) long; antennomere I stout;

antennomere III (Fig. 8H) transverse, shallowly and narrowly concave in dorsal portion, L/W = 0.72;

approximate ratio of each antennomere (n = 1) as 6.6 : 1.0 : 4.7 : 1.7 : 1.7 : 1.6 : 1.6 : 1.6 : 1.6 : 1.6 :

2.6. Caudal margin of tergite VII concave. Aedeagus (Fig. 9H) about 1.22 mm, round at aedeagal apex.

Gonoporal piece (Fig. 3H) straight, wide, 0.62 mm in GL; GL/AL 2.11. Ligula (Fig. 3H) short, slightly

curved; LL 0.24 mm; LW 0.09 mm; LL/LW 2.69; GL/LL 2.58.

Female

HL/HW 0.77-0.92 (0.85); PW/PL 1.27-1.29 (1.28); EL/EW 1.41-3.53 (2.47); EL/PL 2.73-2.89 (2.81);

EW/PW 0.61-1.59 (1.10); TL/EW 2.37-5.82 (4.10). Ante nn ae (Fig. 7H) long; antennomere III relatively

long, L/W = 2.73; approximate ratio of each antennomeres (n = 1) as 8.0 : 1.0 : 4.8 : 2.0 : 2.0 : 2.0 : 2.0 :

2.0 : 2.0 : 2.0 : 3.4.

Measurements

Male (n = 3): TL 5.20-5.55 (5.43) mm; HL 0.90-0.95 (0.92) mm; HW 1.20-1.28 (1.23) mm; PW 1.40-

1.55 (1.47) mm; PL 1.10-1.18 (1.14) mm; EL 3.20-3.50 (3.37) mm; EW 2.15-2.45 (2.30) mm. Female

(n = 2): TL 4.62^1.95 (4.79) mm; HL 0.85-0.92 (0.89) mm; HW 1.00-1.10 (1.05) mm; PW 1.23-1.40

(1.32) mm; PL 0.95-1.10 (1.03) mm; EL 2.75-3.00 (2.88) mm; EW 0.85-1.95 (1.40) mm.

Distribution

Seychelles (Mahe, Felicite Isis.).

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Laius submariniformis Wittmer, 1985

Figs 31, 6F, 71, 81, 91

Laius submariniformis Wittmer, 1985: 385.

Material examined

2 88, 6 (EUMJ), “Damai, Sarawak 17. IX. 1990 Ken ltd leg.”; 5 88, 29 ?? (including 10

paratypes; EUMJ), “PHILIPPINES Sabang, N. of Mt. St. Paul, Palawan July, 11-13, 1977 M. Sato leg.”;

5 88, 4 ? ? (BPBM), “PHILIPPINES (S.) Cagayan Sulu I. Feb. 27. 1957”, “Yoshio Kondo collector”.

Fig. 10. Larva of Laius rodriguesensis sp. nov. A. Dorsal habitus. B. Head in dorsal view. C. Head

in ventral view. D. Abdominal segment IX in dorsal view. E. Abdominal segment IX in ventral view.

F. Abdominal segment IX in lateral view.

European Journal of Taxonomy 97: 1-29 (2014)

Redescription

Male

HL/HW 0.75-0.86 (0.80); PW/PL 1.27-1.37 (1.32); EL/EW 1.38-1.50 (1.42); EL/PL 2.64-2.94 (2.79);

EW/PW 1.43-1.56 (1.48); TL/EW 2.25-2.49 (2.37). Ante nn ae (Fig. 71) long; antennomere I long and

slightly curved laterally; antennomere III (Fig. 81) oval, deeply concave in dorso-mesal portion, L/W =

0.88; approximate ratio of each antennomeres (n = 1) as 15.0 : 1.0 : 9.0 : 3.3 : 3.3 : 3.0 : 3.3 : 3.3 : 3.3 :

3.3 : 5.0. Caudal margin of tergite VII deeply concave. Aedeagus (Fig. 91) about 1.37 mm, concave at

aedeagal apex. Gonoporal piece (Fig. 31) straight, rather wide, with very short basal projection, 0.77 mm

in GL; GL/AL 1.51. Ligula (Fig. 31) long, evenly curved; LL 0.37 mm; LW 0.14 mm; LL/LW 2.58; GL/

LL 2.06.

Female

HL/HW 0.67-0.85 (0.76); PW/PL 1.32-1.45 (1.36); EL/EW 1.33-1.52 (1.39); EL/PL 2.64-2.91 (2.75);

EW/PW 1.31-1.59 (1.46); TL/EW 2.15-2.48 (2.31). Ante nn ae (Fig. 71) long; antennomere III short,

L/W = 2.14; approximate ratio of each antennomeres (n = 1) as 8.0 : 1.0 : 4.2 : 2.3 : 2.0 : 2.0 : 2.0 : 1.8 :

1.8 : 1.8 : 3.3.

Measurements

Male (n = 4): TL 4.90-6.33 (5.60) mm; HL 0.88-1.20 (1.02) mm; HW 1.05-1.52 (1.29) mm; PW 1.40-

1.82 (1.60) mm; PL 1.02-1.40 (1.21) mm; EL 3.00-3.78 (3.37) mm; EW 2.00-2.70 (2.37) mm. Female

(n = 5): TL 4.95-5.65 (5.30) mm; HL 0.80-1.10 (0.95) mm; HW 1.20-1.32 (1.24) mm; PW 1.45-1.68

(1.58) mm; PL 1.10-1.25 (1.16) mm; EL 3.05-3.30 (3.19) mm; EW 2.10-2.40 (2.30) m m

Distribution

Philippines (Palawan, Sulu Isis.), Malaysia (Sarawak).

Remarks

This is the first record of this species from Sulu Island and Borneo.

Laius asahinai Nakane, 1955

Figs 3J, 13D

Material examined

JAPAN: 1 $ (EUMJ), “Oshima, Okada, Shinkai, Nodahama, 31. V. 2013, H. Kamezawa leg.”; 1 $

(EUMJ), “[IZ60] Ookubohama, Miyakejima, Izu Isis., N34 07.272 E139 30.930, ca 152 m, 29. VI. 2013,

H. Yoshitomi leg.”; 1 9 $$ (EUMJ), “[IZ58] Sabigahama, Miyake-jima, Izu Isis., N34 04.241 E139

28.837, ca 151 m, 28-30. VI. 2013, H. Yoshitomi leg.”; 1 $ (EUMJ), “Shimo-Koshiki Is. Kagoshima,

Japan 25-26. V. 2013 N. Ohbayashi leg.”; 2 SS, 3 ?? (EUMJ), “Hachijo Is. Mitsune 20. V. 2001 Y.

Kawabata”. Others see Yoshitomi & Lee (2010).

Description of endophallic sclerites

Gonoporal piece (Fig. 3J) straight, slender, 0.81 mm in GL; GL/AL 3.96. Ligula (Fig. 3J) short, slightly

curved; LL 0.17 mm; LW 0.04 mm; LL/LW 3.88; GL/LL 4.71.

Distribution

Japan (Honshu, Izu Isis. [Oshima, Miyake-jima, Hachijo-jima], Shikoku, Kyushu, Shimo-koshiki-jima,

Yakushima).

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Fig. 11. Larva of Laius rodriguesensis sp. nov. A. Maxillae and labium in ventral view. B. Labram in

dorsal (left) and ventral (right) views. C. Mandible in dorsal (left) and ventral (right) views.

European Journal of Taxonomy 97: 1-29 (2014)

Remarks

This is the first record of this species from Oshima, Miyake-jima, and Shimo-koshiki-jima. This species

is distributed in the northernmost area in the genus, and widely spread. The local variation of the

endophallic sclerites was not observed in this study, but the specimens in the Izu Isis, have somewhat

darker elytral coloration.

Laius etsukoae Sato, Yoshitomi & Ohbayashi, 2006

Fig. 4A

Material examined

Same as in Sato et al. (2006).

Description of endophallic sclerites

Gonoporal piece (Fig. 4A) straight, slender, 0.59 mm in GL; GL/AL 2.49. Ligula (Fig. 4A) short,

distinctly curved in apical portion; LL 0.24 mm; LW 0.10 mm; LL/LW 2.39; GL/LL 2.49. Additional

sclerite (Fig. 4A) present, 0.05 mm.

Laius flavicornis (Fabricius, 1801)

Fig. 4B

Material examined

Same as in Yoshitomi (2008).

Description of endophallic sclerites

Gonoporal piece (Fig. 4B) straight, slender, with short and curved basal projection, 0.74 mm in GL; GL/

AL 2.14. Ligula (Fig. 4B) short, strongly curved in half; LL 0.26 mm ; LW 0.12 mm ; LL/LW 2.14; GL/

LL 2.85.

Laius keiichii Sato, Yoshitomi & Ohbayashi, 2006

Fig. 4C

Material examined

Same as in Sato et al. (2006).

Description of endophallic sclerites

Gonoporal piece (Fig. 4C) straight, slender, 0.54 mm in GL; GL/AL 1.48. Ligula (Fig. 4C) long and

slender, evenly and strongly curved; LL 0.31 mm; LW 0.16 mm; LL/LW 1.90; GL/LL 1.76. Additional

sclerite (Fig. 4C) present. 0.04 mm

Laius lutaoensis Yoshitomi & Lee, 2010

Figs 2, 4D

Material examined

3 SS, 7 ? ? (EUMJ), “Taitung, TAIWAN Is. Ludao 4-6. IV. 2012 N. OHBAYASHI leg.”. Other material

studied is already mentioned in Yoshitomi & Lee (2010).

Description of endophallic sclerites

Gonoporal piece (Fig. 4D) straight, slender, 0.64 mm in GL; GL/AL 2.09. Ligula (Fig. 4D) short,

strongly curved in half; LL 0.23 mm ; LW 0.12 mm; LL/LW 2.00; GL/LL 2.74.

YOSHITOMIFL, Endophallic structures of Laius (Coleoptera)

Laius madli Yoshitomi, 2010

Fig. 4E

Material examined

Same as in Yoshitomi (2010).

Description of endophallic sclerites

Gonoporal piece (Fig. 4E) curved, rather wide, 0.63 mm in GL; GL/AL 2.10. Ligula (Fig. 4E) relatively

short, curved in basal portion; FF 0.28 mm; FW 0.09 mm; FF/FW 2.94; GF/FF 2.26.

Laius miyamotoi Nakane, 1955

Fig. 4F

Material examined

Same as in Yoshitomi & Fee (2010).

Description of endophallic sclerites

Gonoporal piece (Fig. 4F) straight, slender, 0.72 mm in GF; GF/AF 4.13. Figula (Fig. 4F) short, strongly

curved in half; FF 0.15 mm; FW 0.08 mm ; FF/FW 1.80; GF/FF 4.81.

Fig. 12. Laius andamanensis sp. nov., paratype. A. Inner part of fore leg, showing enlarged femur and

tibia. B. Antennomere III. C. Aedeagal apex. D. Spinous area (SEM photographs).

European Journal of Taxonomy 97: 1-29 (2014)

Laius pankowi Wittmer, 1999

Fig. 41

Material examined

Same as in Yoshitomi (2008).

Description of endophallic sclerites

Gonoporal piece (Fig. 41) curved, wide, 1.04 mm in GL; GL/AL 0.85. Ligula (Fig. 41) long and straight,

with reticulate and rounded apex; LL 0.77 mm; LW 0.09 mm; LL/LW 8.18; GL/LL 1.35.

Laius satoi Yoshitomi, 2008

Figs 4G, 13C

Material examined

2 88, 12 ¥¥ (EUMJ), “NosaDua, North Island 8°47 , 52.72 ,, S 105°14 , 9.31 ,, E, Bali, INDONESIA7-8.

VIII. 2010 H. Yoshitomi leg.”; 1 ? (EUMJ), “Nosa Dua, South Island 8°48’6.81”S 115°14 , 16.07 ,, E,

Bali, INDONESIA 8. VIII. 2010 H. Yoshitomi leg.”. Other material studied is already mentioned in

Yoshitomi (2008).

Fig. 13. Habitat and habits of Laius spp. A. Type locality (Nusa Dua, Bali, Indonesia) of Laius baliensis

sp. nov. and L. satoi Yoshitomi, 2008, three Laius species living sympatrically. B. L. baliensis sp. nov.,

female. C. L. satoi , male. D. L. asahinai Nakane, 1955 (at Miyake-jima). Photographs by HY.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Description of endophallic sclerites

Gonoporal piece (Fig. 4G) straight, slender, 0.94 mm in GL; GL/AL 1.25. Ligula (Fig. 4G) long, slightly

curved apical portion; FF 0.49 mm; FW 0.07 mm; FF/FW 7.33; GF/FF 1.92.

Laius taiwanus Yoshitomi & Fee, 2010

Fig. 4H

Material examined

1 s (EUMJ), “Taitung, TAIWAN Is. Fudao 4-6. IV. 2012 N. OHBAYASHI leg.”. Other material studied

is already mentioned in Yoshitomi & Fee (2010).

Description of endophallic sclerites

Gonoporal piece (Fig. 4H) curved, rather wide, 0.67 mm in GF; GF/AF 1.68. Figula (Fig. 4H) long,

slightly curved; FF 0.38 mm; FW 0.08 mm; FF/FW 4.60; GF/FF 1.74. Additional sclerite (Fig. 4H)

present, 0.09 mm

Remarks

Yoshitomi & Fee (2010) pointed out that this species has local variations between the populations from

mainland Taiwan, Futao and Fanyu Isis, in body size and elytral luster. For the present study, I examined

the endophallic structures of these populations, and it became clear that they certainly belong to the

same species because of the highly similar features of the endophallic structures.

Discussion

The endophallic structure of the genus Laius

The results in this paper show that the endophallic structures of the genus Laius are important

taxonomically. For close and precise examination of the endophallic sclerites, it is necessary to remove

the sclerites from the aedeagus directly or extract them from the ventral part of the aedeagal apex.

Based on the constructions of endophallic sclerites in 19 species, the genus is subdivided into the

following five species groups:

Group 1 ( sericatus , etsukoae, keiichii , taiwanus , sabangensis)'. additional sclerite present.

Group 2 ( maai , marc he i, submar inifor mi s,flavicornis)\ gonoporal piece with basal projection.

Group 3 ( baliensis sp. nov., andamanensis sp. nov., rodriguesensis sp. nov. ,politus , sericatus , asahinai ,

lutaoensis, miyamotoi , madli ): ligula short, curved.

Group 4 ( satoi ): ligula long and straight; gonoporal piece long.

Group 5 (pankowi ): ligula long, straight, microcuticulate in apical portion.

Distribution

The species of Laius inhabit rocky seashores (Fig. 13 A), and 1-3 species are distributed in the same

place. Table 1 shows the distribution pattern in five areas (Japan, Taiwan, Palawan Isl., Palau, and Bali

Isl.). Two species in Taiwan (Futao Isl.) and the Philippines (Palawan Isl.), and three in Indonesia (Bali

Isl.) are sympatric, and have different body sizes and endophallic structures (= different species group)

in each area. On the other hand, two species in Japan and Palau are allopatric (or parapatric), their body

sizes overlap and the endophallic structures are also similar to each other (= same species group) in each

area.

All members of Laius have developed hind wings and are capable of flying, and in fact they often fly

in the field. In addition, they are able to stay submerged in seawater for up to 30 hours (Hashimoto

European Journal of Taxonomy 97 : 1-29 ( 2014 )

Table 1 . Distribution of Laius spp. in five areas.

Distribution

Pattern

Species

group

Measurement in male (mm)

Reference

Locality

Japan

Honshu to

Yakushima

allopatric

asahinai

3.95-4.65 (4.27)

1.19

0.81

0.17

Yoshitomi & Lee

2010

Ryukyu Isis.

miyamotoi

3.50M.20 (3.88)

1.17

0.72

0.15

Yoshitomi & Lee

2010

Taiwan

Lutao Isl.

sympatric

tarn anus

3.90-4.88(4.18)

1.03

0.67

0.38

Yoshitomi & Lee

2010

lutaoensis

4.80-5.45 (5.18)

1.31

0.64

0.23

Yoshitomi & Lee

2010

Philippines

Palawan

sympatric

submar iniformis

4.90-6.33 (5.60)

1.37

0.77

0.37

present study

sabangensis

4.03-4.73 (4.23)

0.94

0.72

0.41

present study

Palau

Merir, Sonsorol Isis.

allopatric

etsukoae

3.4M.5

1.00

0.59

0.24

Sato et at. 2006

Peleliu Isl.

keiichii

3.8

1.19

0.54

0.31

Sato et at. 2006

Indonesia

Bali Isl.

sympatric

satoi

5.7

1.53

0.94

0.49

Yoshitomi 2008

pankowi

6.13-6.55 (6.34)

1.58

1.04

0.77

Yoshitomi 2008

baliensis

4.10-4.46(4.27)

1.12

0.61

0.23

present study

1970). Therefore it is supposed that the distribution of each species is widely spread through flight

and tidal currents. However, most of the species show narrow distributions (see Appendix). The genus

Aegialites Mannerheim, 1853 (Coleoptera, Salpingidae, Aegialitinae) also lives in rocky seashores, and

the distribution of each of its species is generally restricted as the ones of the species of the genus Laius

(see Doyen 1976; Zerche 2004). However, Aegialites is flightless with degenerated hind wings, and

its distribution is limited in the Northern Pacific (latitude c. 30-60°N; Northern Japan, Kuril Isis., to

North America). The speciation and distribution pattern of Laius provide interesting zoogeographical

and evolutionary themes.

Acknowledgements

I thank Dr. Manfred Jach (NMW), Dr. Michael Geiser (NHMB), Mr. Ryo Ogawa (Kobe Univ.), Dr.

Junsulce Yamasako and Dr. Nobuo Ohbayashi (both EUMJ), Miss Makoto Asano, Mr. Shepherd Myers

(BPBM), Dr. Nugroho Susetya Putra (Gadjah Mada University), and the late Dr. Masataka Sato for

supplying the materials used in this paper, and Mr. Dennis Murphy (The United Graduate School of

Agricultural Sciences, Ehime University) for his critical reading of the draft. A part of this study is

supported by the KAKENHI (24510333; head investigator: H. Kojima).

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Manuscript received: 22 March 2014

Manuscript accepted: 25 June 2014

Published on: 25 September 2014

Topic editor: Koen Martens

Desk editor: Charlotte Thionois

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, Denmark.

YOSHITOMIH., Endophallic structures of Laius (Coleoptera)

Appendix. Updated list of the true Laius species of the world. * Laius heterocerus Boisduval, 1835

was treated as a junior synonym of this species, and recorded from New Guinea, India, Key Isis.,

Karoline, New Caledonia (Evers 1994). ** This species has also been recorded from Larat, Key Isis.,

Borneo, Lundu, Sarawak, and China (Evers 1994); however, these records were probably based on the

misidentification of another species.

Laius alleni Lea, 1909.

Laius ambonensis Wittmer, 1996.

Laius andamanensis Yoshitomi sp. nov.

Laius asahinai Nalcane, 1955.

Laius baliensis Yoshitomi sp. nov.

Laius cyaneus Guerin-Meneville, 1830*.

Laius cyanoxanthus Bourgeois, 1897.

Laius etsukoae Sato, Yoshitomi & Ohbayashi, 2006

Laiusflavicornis (Fabricius, 1801)**.

Laius impressus Wittmer, 1985.

Laius keiichii Sato, Yoshitomi & Ohbayashi, 2006..

Laius lutaoensis Yoshitomi & Lee, 2010.

Laius miyamotoi Nalcane, 1955.

Laius maai Wittmer, 1973.

Laius madli Yoshitomi, 2010.

Laius marchei Pic, 1922.

Laius miles Bourgeois, 1905.

Laius mindanaonus Wittmer, 1985.

Laius obscurus (Pic, 1908).

Laius pankowi Wittmer, 1999.

Laius politus Fairmaire, 1880.

Laius purpureipennis Lea, 1916.

Laius riedeli Evers, 1994.

Laius rodriguesensis Yoshitomi sp. nov.

Laius rufipes Montrouzier, 1860.

Laius sabangensis Wittmer, 1985.

Laius satoi Yoshitomi, 2008.

Laius sericatus Champion, 1923.

Laius submarinus Champion, 1921.

Laius submariniformis Wittmer, 1985.

Laius taiwanus Yoshitomi & Lee, 2010.

Laius tibialis Gahan, 1900.

Laius velutinus Gerstaecker, 1873 .

Laius wallecei Pic, 1937.

Australia (Queensland) [after Evers 1994]

Indonesia (Ambon Isl.) [after Plonski 2013]

India (Andaman Islands)

Japan (Honshu to Yakushima)

Indonesia (Bali Isl.)

India, New Guinea [after Evers 1994])

New Hebrides (Mallicolo) [after Evers 1994]

Palau (Merir, Sonsorol Isis.)

Indonesia (Java Isl.)

Philippines (Mindanao Isl.) [after Evers 1994]

Palau (Peleliu Isl.)

Taiwan

Japan (Ryukyu Isis.)

New Guinea

Republic of Mauritius (Mauritius Isl.)

Mariana (Rota, Guam Isis.)

Ceylon [after Evers 1994]

Philippines (Mindanao Isl.)

Kenya [after Evers 1994]

Indonesia (Bali Isl.)

Madagascar

Australia [after Evers 1994]

Malaysia (Sarawak)

Republic of Mauritius (Rodrigues Isl.)

New Caledonia [after Evers 1994]

Philippines (Palawan Isl.)

Indonesia (Bali Isl.)

Seychelles (Mahe, Felicite Isis.)

Philippines (Mindanao Isl.), New Guinea

Philippines (Palawan Isl.)

Taiwan (including Lutao Isis.)

Christmas Isl.

Zanzibar Coast [after Champion 1921]

Malaysia (Borneo)