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Bonn zoological Bulletin

EDITOR-IN-CHIEF

Fabian Herder, Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK), Ichthyology Section, Adenauerallee 160,

53113 Bonn, Germany,

tel. +49 228-9122—255, fax: +49 228-9122-212;

f.herder.zfmk@uni-bonn.de

MANAGING EDITOR SUPPLEMENT SERIES

Thomas Wesener, ZFMK, tel. +49 228-9122-425,

fax: +49 228-9122-212; twesener@uni-bonn.de

EDITORIAL BOARD

Dirk Ahrens, Insects: Coleoptera, ZFMK,

tel. +49 228-9122—286, fax: +49 228-9122-332;

d.ahrens.zfmk@uni-bonn.de

Wolfgang Boéhme, Amphibians and Reptiles, ZFMK,

tel. +49 228-9122—250, fax: +49 228-9122-212;

w.boehme.zfmk@uni-bonn.de

Netta Dorchin, Insects: Diptera, Department of Zoology, Tel

Aviv University, Tel Aviv 69978, Israel, tel. +972-3-

6409808, fax: +972-3-6409403; ndorchin@post.tau.ac.il

Renate van den Elzen, Birds, ZFMK,

tel. +49 228-9122-231, fax: +49 2289122212:

r.elzen.zfmk(@uni-bonn.de

Bernhard Huber, Invertebrates except Insects, ZFMK,

tel. +49 228-9122—294, fax: +49 228—-9122-212:;

b.huber.zfmk@uni-bonn.de

Rainer Hutterer, Mammals, ZFMK,

tel. +49 228-9122-261, fax: +49 228-9122-212;

r.hutterer.zfmk@uni-bonn.de

Gustav Peters, Mammals, ZFMK,

tel. +49 228-9122—262, fax: +49 228-9122-212;

g.peters.zfmk@uni-bonn.de

Bradley Sinclair, Canadian National Collection of Insects,

Ottawa Plant Laboratory — Entomology, CFIA, K.W. Neat-

by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada

KIA 0C6, tel. + 1 613-759-1787, fax: + 1 613-759-1927;

bradley.sinclair@inspection.ge.ca

Dieter Stiining, Insects except Coleoptera and Diptera,

ZFMK, tel. +49 228-9122—220, fax: +49 228-9122-212;

d.stuening.zfmk@uni-bonn.de

Philipp Wagner, Villanova University, Department of Biolo-

gy, 800 Lancaster Avenue, Villanova, PA 19085-1699,

USA; philipp.wagner.zfmk@uni-bonn.de

ADVISORY BOARD

Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer-

sitat, Institut fir Evolutionsbiologie & Okologie, 53113

Bonn, Germany, tel. +49 228—73—5130, fax: +49 228-73-

2321; t.bakker@uni-bonn.de

Aaron M. Bauer, Villanova University, Department of Biolo-

gy, 800 Lancaster Avenue, Villanova, PA 19085-1699,

USA, tel. +1 610-519-4857, fax: +1 610-519-7863;

aaron.bauer@villanova.edu

Wieslaw Bogdanowicz, Museum and Institute of Zoology,

Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa,

Poland, tel. +48 22628-7304, fax: +48 22—629-6302;

wieslawb@miiz.waw.pl

Matthias Glaubrecht, Museum fiir Naturkunde Berlin, Leib-

niz-Institut fir Evolutions- und Biodiversitaétsforschung an

der Humboldt-Universitat zu Berlin, Invalidenstrasse 43,

10115 Berlin, Germany, tel. +49 30—2093-8504/ 8400,

fax: +49 030—2093-8565; matthias.glaubrecht@mfn-

berlin.de

Jeremy D. Holloway, The Natural History Museum, Depart-

ment of Entomology, Cromwell Road, London, SW7 5BD,

U.K.; j.holloway@nhm.ac.uk

Tan Heok Hui, Raffles Museum of Biodiversity Research,

National University of Singapore, Department of Biological

Sciences, 6 Science Drive 2, #03-01, Singapore 117546, tel.

+65-6516 1662, heokhui@nus.edu.sg

Boris KrySstufek, Slovenian Museum of Natural History, P.

O. Box 290, Ljubljana, Slovenia; boris.krystufek@zrs.upr.si

Wolfgang Schawaller, Staatliches Museum ftir Naturkunde,

Rosenstein 1, 70191 Stuttgart, Germany,

tel. +49 711-8936—221, fax: +49 711-8936—100;

schawaller.smns@naturkundemuseum-bw.de

Ulrich K. Schliewen, Department of Ichthyology, Bavarian

State Collection of Zoology, Minchhausenstr. 21, 81247

Miinchen, Germany, tel. + 49 89—8107—110;

schliewen@zsm.mwn.de

Michael Schmitt, Ernst-Moritz-Armdt-Universitat, Allge-

meine & Systematische Zoologie, Anklamer Str. 20, 17489

Greifswald, Germany, tel. +49 3834864242, fax: +49

3834-86-4098; michael.schmitt@uni-greifswald.de

W. David Sissom, Dept. of Life, Earth and Environmental

Sciences, W. Texas A. & M. University, WITAMU Box

60808, Canyon, Texas 79016, USA, tel. +1 806-65 1—2578,

fax: +1 806-651-2928; dsissom@mail.wtamu.edu

Miguel Vences, Technical University of Braunschweig, Zoo-

logical Institute, Spielmannstr. 8, 38106 Braunschweig,

Germany, tel. + 49 531-391-3237, fax: + 49 31-391-8198;

m.vences(@tu-bs.de

Erich Weber, Eberhard-Karls-Universitat, Zoologische

Schausammlung, Sigwartstr. 3, 72076 Tubingen,

tel. +49 7071—2972616, fax +49 7071—295170;

erich.weber@uni-tuebingen.de

Editorial

Welcome to the first issue of the Bonn zoological Bulletin

in 2012. The most comprehensive of the papers published

here is a revision of the East Palaearctic beetle genus

Lobrathium, containing descriptions of a total of thirty-

three species new to science, a catalogue of all species of

the Palaearctic, and keys for the Himalaya and China. Re-

visions like this one, written by Volker Assing, set the stage

for further taxonomic work in the respective group of or-

ganisms, and remain a highly valuable reference for

decades. Descriptions of two additional Asian beetle

species, the first record and redescription of a Geometer

Moth from Iran and Turkey, and a morphological study

focusing on the larval urticating apparatus of a Lappet

Moth species add to our knowledge of arthropod diversi-

Other studies published in the present volume deal with

vertebrates, namely fishes and lizards. Two taxonomic pa-

pers give first descriptions of new colourful killifish

species from West Africa, and the study by Wagner and

co-authors provides data on morphology, behaviour and

habitats of a snake-like plated lizard in Zambia. A second

brief herpetological manuscript reports for the first time

the life colouration of a conspicuous, beige to pinkish, bur-

rowing Madagascan skink. Finally, a brief note proposes

in a nomenclatorial act a replacement name for a fish

genus.

SMITHSON, y

AUG 2 1 2012

LIBRARIES

The Bonn zoological Bulletin is an open access jour-

nal, and its contents have so far been published online in

a section of Museum Koenig’s website. These pages have

now been replaced by a new, attractive website available

at: www.zoologicalbulletin.de. | am pleased to announce

that this website will be accessible with publication of this

issue, increasing the visibility of the journal and its con-

tents. The new, comprehensive archive with the complete

set of open access pdfs is currently under development, a

process that will be completed later this year. The “old”

website will of cause be maintained at least until the trans-

fer of BzB contents is completed. I thank Kat Stojanov

and Sebastian Hitillen very much for setting up this new

portal; I am convinced that readers and authors will take

multiple advantage of it.

With best wishes to all readers, authors and colleagues

from the “BzB-team”,

Tibi floc

Fabian Herder (Editor-in-Chief, Fish Curator at ZFMK)

Bonn, July 2012

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Bonn zoological Bulletin 61 (1): 3-12

Aphyosemion pseudoelegans (Cyprinodontiformes: Nothobranchiidae),

a new killifish species from the Cuvette centrale in the Congo Basin

(Democratic Republic of Congo)

Rainer Sonnenberg’’ & Jouke R. Van der Zee’

‘ Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

> Max-Planck-Institut ftir Evolutionsbiologie, Department of Evolutionary Genetics, August-Thienemann-Strasse 2,

D-24306 Plén, Germany. Corresponding author. E-mail: r.sonnenberg.zfmk@uni-bonn.de

> Royal Museum for Central Africa, Vertebrate Section, Ichthyology, B-3080 Tervuren, Belgium.

Abstract. Aphyosemion pseudoelegans, new species, is described, based on collections that have been misidentified for

decades as A. elegans by all authors. The new species superficially resembles A. e/egans, but can be distinguished from

the latter and all other species of the genus by a diagnostic combination of colour pattern characters, most prominently

it has a dark red to black dorsal fin in males, a colour pattern always absent in all known A. e/egans populations, includ-

ing the type specimens, and an asymmetrical colour pattern on caudal fin margins versus symmetrical in A. elegans.

Aphyosemion pseudoelegans, new species, is found in small rivers on the left bank of the middle Congo, where it occurs

sympatric and sometimes even syntopic with A. elegans, A. sp. aff. castaneum, or an undescribed species of Aphyosemion

at Ikela.

Résumé. Aphyosemion pseudoelegans, espéce nouvelle, est décrite, sur base de collections qui furent, depuis des deé-

cades, mal identifiées en tant que A. e/egans par tous les auteurs. La nouvelle espece ressemble superficiellement a A.

elegans, mais peut en étre distinguée, ainsi que de toutes les autres especes du genre, par une combinaison diagnostique

de caracteres du patron de coloration. Principalement, elle présente une nageoire dorsale rouge foncé a noire chez les

males, un patron de coloration toujours absent chez toutes les populations connues de A. e/egans, y inclus les spécimens

types, et un patron de coloration asymétrique des marges de la nageoire caudale alors qu’il est symétrique chez A. ele-

gans. Aphyosemion pseudoelegans, espéce nouvelle, se rencontre dans de petites rivieres de la rive gauche du Congo

Moyen. Elle y est sympatrique, et parfois méme syntopique, de A. elegans, A. sp. aff. castaneum, ou une espéce non-

July 2012

décrite de Aphyosemion a Ikela.

Key words. taxonomy, Aphyosemion elegans, DNA, morphology, distribution.

INTRODUCTION

In some recent publications, the taxonomy of species of

Aphyosemion s.1. (Cyprinodontiformes: Nothobranchiidae)

of the Congo basin was reviewed, leading to a revalida-

tion of a taxon and the description of new species (Huber

1994, 2004, 2005a, b; Huber & Scheel 1981; Van der Zee

& Huber 2006; Van der Zee & Sonnenberg 2010, 2011;

Woeltjes 1984). Here the genus Aphyosemion Myers, 1924

is used in a more restricted sense, as proposed in three re-

cent publications of the authors (Sonnenberg 2007; Van

der Zee & Sonnenberg 2010, 2011). This corresponds with

the subgenus Aphyosemion or the A. elegans species group

of other authors (e.g. Collier 2007; Huber 2007; Murphy

& Collier 1999; Wildekamp 1993) and includes 17 species

currently accepted as valid (Van der Zee & Sonnenberg

2011). However, the species level taxonomy of this genus

is still not settled. In a recent publication, the occurrence

of another undescribed species was indicated, frequently

misidentified as A. elegans (Baensch & Riehl 1985; Hu-

ber 2005 a, b; Ott 2002; Scheel 1968, 1990; Seegers 1997;

Received: 30.01.2012

Accepted: 08.06.2012

Van der Zee & Huber 2006; Wildekamp 1993) from the

central Congo basin, based on diagnostic colour pattern

differences and supported by a preliminary DNA study

(Van der Zee & Sonnenberg 2011).

In 1899, Boulenger described Haplochilus elegans

based on specimens which originated from Bikoro at Lac

Tumba (Democratic Republic of Congo), and Coquil-

hatville, close to the Congo River (currently Mbandaka,

capital of the province Equateur, Democratic Republic of

Congo). Later this species was included in the genus

Aphyosemion, erected by Myers in 1924.

Until recently, it was assumed that A. elegans is vari-

able with respect to colour pattern, including that of the

dorsal fin (Radda & Piirzl 1987; Huber 2004, 2005a, b;

Van der Zee & Sonnenberg 2011; Wildekamp 1993). How-

ever, the known collections currently identified as A. e/-

egans can easily be separated into two groups, distin-

guished by dorsal fin colouration and colour pattern of the

caudal fin. One group has a light dorsal fin with many

Corresponding editor: F. Herder

4 Rainer Sonnenberg & Jouke R. Van der Zee

small red dots and a symmetric colour pattern in the cau-

dal fin, whilst the other has a mainly dark red dorsal fin

(Van der Zee & Sonnenberg 2011) and an asymmetrical

caudal fin pattern.

Within the description of Haplochilus elegans, a draw-

ing of a male syntype was published, that clearly shows

the colour pattern of the median fins. Additionally it was

mentioned in the text, that all median fins are provided

with carmine red dots (Boulenger 1899). A dark dorsal fin

was never mentioned, nor does the drawing of a syntype

show this character. This indicates that the types of A. e/-

egans belong to the group with a narrow red edge and

small red dots on a light background on the dorsal fin.

Aphyosemion elegans is widespread in the central and

northern Congo Basin (Fig. 1). All known populations

from the northern part of the basin have a dorsal fin with

a narrow red edge and red dots on a light background. In

contrast, in the central part of the basin, the Cuvette cen-

trale, several populations show a complete or nearly com-

plete dark red dorsal fin. Both phenotypes occur within

the Cuvette centrale in sympatry, and at Bokuma (Equa-

teur Province) they occur syntopic (Fig. 1; syntopic col-

lection: MRAC 79229-79236 (A. elegans), MRAC 79237

Sees Gerbéreti R Sea

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A. pseudoelegans

A. elegans

Syntopic A. pseudoelegans and A. elegans

Terra typica

<—>100km

Fig. 1.

AMNH, MNHN, MRAC, and ZSM collections.

Bonn zoological Bulletin 61 (1): 3-12

(A. pseudoelegans, new species)). In addition, A. pseudoel-

egans can be found sympatric and syntopic with A. sp. aff.

castaneum in parts of the Cuvette centrale and with an un-

described species of Aphyosemion at Ikela (see Van der

Zee & Sonnenberg 2011; pers. obs. unpub.).

The probably first import of live specimens with dark

red dorsal fins to Europe might have been made by Lam-

bert in the sixties, originating from Boende (Baensch &

Riehl 1985, as A. elegans). Both, Aphyosemion elegans

sensu Boulenger (1899) and specimens with a dark red

dorsal fin were kept and bred by aquarium hobbyists, es-

pecially the recent imports from the Boende area (Ott

2002; Van der Zee & Sonnenberg 2011), so it could be

confirmed that the offspring only shows the parental

colour pattern (pers. obs., H. Ott pers. comm.).

In addition, a recent phylogenetic analysis based on mi-

tochondrial DNA data of 11 species of Aphyosemion in-

dicated that specimens with a dark dorsal fin do not be-

long to A. elegans. Instead, they represent an undescribed

species (Van der Zee & Sonnenberg 2011). In this paper

the species with the dark dorsal fin, called preliminarily

A. sp. Cuvette in Van der Zee & Sonnenberg (2011), is de-

scribed as Aphyosemion pseudoelegans, new species.

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Collection point map of Aphyosemion pseudoelegans and Aphyosemion elegans based on specimens and data from the

©OZFMK

Aphyosemion pseudoelegans from the Congo Basin 5

Table 1. Morphometrics of Aphyosemion pseudoelegans, new species (H = holotype, P = paratypes: 8 males and 3 females). All

measurements in percentages of standard length, standard length in mm. Paratype | MRAC 178017, paratypes 2—11 ZSM 32434.

P19

lal@ Wie PAs PES Pag PSs Iss Pas IG IVS wile

Standard length 33.3 B67 33.3 B20 299 Wi Ads 92 an) ss a) ae

Body depth IO 19. NO les U7 Wis 20s ZO AUIS oe 7 es Ia

Head length De Ne Sie le One) fe lee lee 2S 221 20:6) 22:8 22.8

Eye diameter 7.8 alee flee 7.8 8.0 8.5 8.5 8.3 8.5 Tell 8.8 8.9

Interorbital width OD OG OO Ose OA OKG 24 NOS) 7 10:4 11.4

Pre-dorsal length CORREO SalmOO 4 OSol) mG 2) OS-O) 09:40 05:2) 1694" 165195 69:1 ~~ 70:9

Pre-anal length SAS ae ONS LOR eS son MENTO Sea 728) S34) 3:4 57.8

Pre-ventral length 456 444 44.1 44.7 425 441 422 436 42.2 418 446 48.1

Dorsal fin base 23 W270) NO 7 L220 Oe Ol OD NOLO 9:4 10.4 10.1

Anal fin base Bol B78 Wry AVS AVG! 22g) PA Dsy ADs So I ID)

Caudalipedslenothwae2S a2 On 2 Sale 22-24 al 2228) 23:9" 2106 252 23:33 18:9 22.4

Caudal ped. depth AS en > ee Ol HO mlisHOmm 3-35) 1352" 12257 1259) 13.5

Caudal ped. ratio 189) ES iareled 188 ey ied) 1.8 1.6 1.6 IES. INES) ey

Table 2. Meristics of the types of Aphyosemion pseudoelegans,

new species. Numbers indicate observed values, numbers in

parentheses the frequency of occurrence; data of the holotype

are indicated by an asterisk. Lateral line scale count only for the

nine specimens with no loss of scales.

meristic count number (frequency)

8 (7), 9 (5*)

13 (3), 14 (8*), 15 (1)

8 (4*), 9 (6), 10 (2)

24 (3), 25 (5), 26 (2), 27 (1*), 28 (1)

29 (5), 30 (4*)

Dorsal fin rays

Anal fin rays

D/A

Caudal fin rays

Lateral line scales

MATERIAL AND METHODS

The description of the new species is based on collections

made in the Central Congo Basin, deposited in the Roy-

al Museum for Central Africa (MRAC, Tervuren, Bel-

gium) and the Zoologische Staatssammlung Miinchen

(ZSM, Munich, Germany). Additional non-type and

comparative material is listed in Van der Zee & Sonnen-

berg (2010, 2011) and below.

Counts and measurements follow Amiet (1987) and

were taken with a digital caliper, partly under a dissect-

ing microscope, and rounded to the nearest 0.1 mm. Meas-

urements, including those taken on the head, were present-

ed as percentages of standard length (SL). All visible fin

Bonn zoological Bulletin 61 (1): 3-12

rays were counted, relative position of first dorsal fin ray

to anal fin (D/A) is estimated as in Sonnenberg & Schun-

ke (2010). Nomenclature for neuromast system on head

follows Scheel (1968) and Van den Bergeijk & Alexan-

der (1962), and for supraorbital (or frontal) squamation

Hoedeman (1958). Heads of different species of

Aphyosemion were scanned with a Scanco viva CT40 at

the Max-Planck-Institut ftir Evolutionsbiologie (Plén, Ger-

many). Digital X-ray images were made with a Faxitron

LX-60 Digital Specimen Radiography System at the Zo-

ologisches Forschungsmuseum Alexander Koenig

(ZFMK, Bonn, Germany).

As species concept, we adopted the pragmatic approach

by Moritz et al. (2000) which is similar to the Evolution-

ary or Phylogenetic Species Concept.

RESULTS

Aphyosemion pseudoelegans, new species

(Fig. 2; Table 1—2)

Aphyosemion elegans, non (Boulenger, 1899): Baensch &

Riehl (1985), Huber (2005 a, b), Ott (2002), Scheel (1968,

1990), Seegers (1997), Van der Zee & Huber (2006), Wil-

dekamp (1993).

Aphyosemion sp. Cuvette: Van der Zee & Sonnenberg

(2011).

Aphyosemion sp. aff. elegans: Van der Zee & Sonnenberg

(2011).

©ZFMK

6 Rainer Sonnenberg & Jouke R. Van der Zee

Fig. 2. A) Male of Aphyosemion pseudoelegans, collected at Boende in May 2002, not preserved. Digital copy of a colour slide.

B) Female of Aphyosemion pseudoelegans, collected at Boende in May 2002, not preserved. Photographed by H. Ott three months

after collection.

Holotype. MRAC 178016, male, 33.3 mm SL, Democrat-

ic Republic of Congo, Equateur Province, Tshuapa Basin,

Boende, 0°14’S, 20°50’E, coll. P. Brichard, 1969.

Paratypes. MRAC 178017, male, 36. 7 mm SL, collect-

ed with the holotype; ZSM 40762 (ex ZSM 32434), 7

males, 25.8-33.3 mm SL, 3 females, 23.7—28.7mm SL,

Democratic Republic of Congo, Bandundu Province,

Lokoro Basin, Lui Kotale, 2°45’S, 20°21’E, coll. G.

Hohmann, November 2000.

Diagnosis. (Figs 2-3) Aphyosemion pseudoelegans shows

the diagnostic combination of characters for the genus

Aphyosemion, which are: preopercular neuromast system

with six pores, slender body shape, posterior origin of dor-

sal fin, dorsal fin small with less than ten fin rays, extend-

Bonn zoological Bulletin 61 (1): 3-12

ed fin rays at the edges of the caudal fin in males, and fe-

males with a well visible reticulation due to dark scale bor-

ders.

Males of A. pseudoelegans differ from all other

Aphyosemion species by a diagnostic combination of

colouration characters. Most prominently it differs from

all other species except A. congicum (Ahl, 1924) by the

almost complete dark red to black dorsal fin versus red

dots or stripes on lighter background. It also differs from

all other species except A. e/egans (Fig. 4) by irregular

narrow vertical red bars, based on confluent dots, on flanks

of males.

It differs from A. congicum by having an asymmetrical

colour pattern of the marginal and submarginal bands in

the caudal fin versus colour pattern of dorsal and ventral

marginal and submarginal bands symmetrically.

OZFMK

Aphyosemion pseudoelegans from the Congo Basin a

Fig. 3. Dorsal fin of male specimens preserved in ethanol. In these specimens the red pigmentation is lost, but the difference in

colour pattern between the two species is still visible. A) 4. pseudoelegans, MRAC 80709-80777, Bokuma, with plain greyish to

brownish fin tissue. B) A. elegans, MRAC 93724, Bokuma, fin tissue with light dots which are the remains of the former red pig-

mentation.

Aphyosemion congicum has only a few red dots on the

flanks and an almost dotless anal fin versus many red dots

or bars on flanks, anal fin with many red dots or inter ra-

dial red stripes.

Besides the dark dorsal fin, the body colour pattern re-

sembles that of A. elegans, but A. pseudoelegans has an

asymmetric colour pattern in the caudal fin versus a sym-

metric pattern in A. e/egans. In general, A. pseudoelegans

is more intensely pigmented then A. e/egans, often hav-

ing larger and more red dots on the flanks and a denser

red pigmentation of the head. Aphyosemion elegans al-

ways shows two or more small isolated rounded red dots

underneath the eye, whereas in most A. pseudoelegans

these dots are larger and irregular and often fused, form-

ing a red streak. In very heavily pigmented A. pseudoel-

egans, the red dots in the caudal fin may form vertical

bars; this is never the case in A. e/egans.

Both species may show narrow vertical dark red bars

on the flanks, but in A. e/egans these bars may run all over

the flanks, whereas in A. pseudoelegans these bars in most

cases only cover the ventral part of the flanks, and dor-

sally frequently with nearly regular rows of red dots.

Bonn zoological Bulletin 61 (1): 3-12

Description. See Figures 2—3 for general appearance and

colour pattern, Table 1 for morphometric, and Table 2 for

meristic data of the type series. Aphyosemion pseudoele-

gans 1s a Slender, elongate species. It shows strong sexu-

al dimorphism, males more colourful, unpaired fins larg-

er and with fin extensions. Relatively large species with-

in Aphyosemion (up to 51.4 mm TL), laterally slightly

compressed; dorsal profile slightly convex, greatest body

depth approximately anterior or at pelvic fin insertion.

Ventral profile slightly convex from head to end of anal

fin, straight or slightly concave on caudal peduncle. Cau-

dal peduncle slender, peduncle depth is 1.5—1.9 times in

length.

Snout slightly rounded, mouth directed upwards, low-

er jaw longer than upper jaw. Dentary with outer row of

large and inner irregular rows of smaller unicuspid, curved

teeth; premaxilla with some larger and several smaller uni-

cuspid and curved teeth.

Frontal (after Scheel, 1968) or nasal (after van Bergeijk

& Alexander, 1962) neuromasts in separate grooves, pre-

opercular canal with six pores.

Scales cycloid, entirely scaled except ventral surface of

head; frontal squamation of G-type; scales on mid-longi-

©ZFMK

8 Rainer Sonnenberg & Jouke R. Van der Zee

Fig. 4. A) Male of Aphyosemion elegans, commercial import (2006) from the Tshuapa River near Boende, not preserved. B) Fe-

male Aphyosemion elegans, commercial import (2006) from the Tshuapa River near Boende, not preserved. Photographed by H.

Ott.

tudinal series 28-30, with two scales posterior to hypur-

al plate; seven transversal scales, 12 scales around cau-

dal peduncle.

Small dorsal fin with 8—9 fin rays, first dorsal fin ray

inserts above 8—1 0th anal fin ray; anal fin with 13—15 rays;

posterior dorsal and anal fin rays slightly elongated in

males; caudal fin with 24—28 rays, with extensions on up-

per and lower fin rays.

X-ray images of three males from comparative materi-

al (MRAC 37840—-843) show 29-30 vertebrae and well

separated upper and lower hypural bones.

Live colouration. Males (Fig. 2A). Flanks golden to

brown, with yellow to bluish-green iridescence. Most flank

scales with a red dot. These dots are mainly situated at the

posterior edge of the scales and usually form up to five

Bonn zoological Bulletin 61 (1): 3-12

parallel, interrupted stripes. Many populations have 1n ad-

dition scales that have the red dots situated at the anteri-

or edge of the scale. This causes, in combination with the

adjacent scales above and below with red dots at the pos-

terior edge, vertical narrow red bars. Three red streaks on

opercle in an approximate 45° angle. One or two infra-

buccal bands present on the lower jaw. Red dots under-

neath the eye often united in a red streak. Pectoral fin

translucent yellow to orange with a whitish or light yel-

low edge; red dots present, sometimes concentrated near

the yellow edge forming a red band. Anal fin light blue

or light yellow, provided with many red dots and/or in-

ter-radial red stripes, sometimes with a yellow or light blue

margin. Dorsal fin predominantly dark red to almost black,

with a narrow light band at the base and edged with a

white, yellow, blue or nearly black band, depending on ex-

©OZFMK

Aphyosemion pseudoelegans from the Congo Basin 9

posure to light. Central part of caudal fin with many red

dots and/or inter-radial stripes. Upper margin yellow or

whitish to light blue, sub-marginal band dark red, lower

margin with broad light sub-marginal band and narrow

dark red marginal band. In some populations, including

the types, this red band is interrupted or absent.

Females (Fig. 2B). Flanks grey with strong red reticu-

lation; red streaks on opercle reduced, often only middle

streak clearly visible; infra-buccal band absent. All fins

transparent; edge of anal fin and distal part of ventral fin

light blue or white; small red dots on anal fin, large red

dots and/or short inter-radial stripes in dorsal fin; small

red dots on anterior dorsal edge and at base of caudal fin.

Colour in ethanol. The types from Boende and the com-

parative material from Bokuma show the typical pattern

of preserved Aphyosemion species, in which the red pig-

mentation pattern, after preservation in formalin and trans-

fer into ethanol, leaves corresponding patterns of lighter

areas than the body colouration (Van der Zee & Sonnen-

berg 2010). The dark red band in the dorsal fin becomes

nearly black, grey, or dark brown. After eleven years in

96% ethanol the red colour of the ZSM paratypes is still

present and resembles the live colour pattern except for

the iridescence and the dark red band in the dorsal fin and

upper caudal fin that became nearly black. This is also the

case in one collection originating from Boende and two

of four collections coming from Ikela. Material collected

here in 1958 and 1972 has the red pigmentation perfect-

ly preserved, whereas material collected in 1955 and 1959

shows the typical colouration of preserved Aphyosemion

specimens. Interestingly, the colouration of the formerly

dark red dorsal in A. pseudoelegans leaves, when pre-

served in ethanol, a darker area than the red pigment on

the body; whereas red pigment on the body and dorsal fin

leaves lighter areas in A. elegans (see Fig. 3). Eventual-

ly the red pigment of the dorsal fin differs or the colour

is based on a combination of the more common red pig-

ment as in other species and a darker pigment, which be-

comes nearly black and later greyish in preservation.

Habitat and Distribution. Like all members of the genus

Aphyosemion, A. pseudoelegans inhabits shallow and

small brooks under forest cover. Currently known collec-

tions indicate, that 4. pseudoelegans is restricted to trib-

utaries on the left bank of the middle Congo basin, also

known as the Cuvette or Cuvette centrale. This area is en-

closed by the Congo River in the north, east, and west,

and by the Kasai River in the south of the Democratic Re-

public of Congo (Fig. 1). Aphyosemion pseudoelegans 1s

present in the Tshuapa, Lomani and Lokoro drainages.

Etymology. The name Aphyosemion pseudoelegans

refers to the misidentification as A. elegans, a similar

species, but lacking the typical dark red dorsal fin and the

Bonn zoological Bulletin 61 (1): 3-12

asymmetric caudal fin colour pattern; from the Greek

wevdns , meaning lying or false, the latter is our intend-

ed translation.

DISCUSSION

The study of the killifishes of the Congo Basin by the sen-

ior author indicates, that there are still several undescribed

species in museum collections and the collections made

recently indicate, that further undescribed species are still

to be discovered in this large and mostly not well sam-

pled part of Africa (see also Sonnenberg et al. 2011, Van

der Zee & Sonnenberg 2010, 2011).

We recently made the assumption that A. elegans and

A. pseudoelegans represent different species (Van der Zee

& Sonnenberg 2011). Both were kept and bred by killi-

fish hobbyists for several years and their observations in-

dicate that A. pseudoelegans is not simply a case of poly-

chromy of A. e/egans, which is supported by recently pub-

lished DNA data (Van der Zee & Sonnenberg 2011). In

addition, we did not find any indication of potential hy-

brids in the studied material of both species. The diagno-

sis of A. pseudoelegans from A. elegans is mainly based

on colour pattern differences of unpaired fins. In gener-

al, this genus has a comparatively high variability in

colouration (see also Van der Zee & Sonnenberg 2011),

but beside details, the general pattern of fins, especially

the order of dark and light marginal and submarginal bands

is a comparatively stable character according to our ob-

servations. Aphyosemion elegans and A. pseudoelegans

can easily be distinguished by their different, stable pat-

tern of light and dark areas, and less by their similar, but

not identical, body colour pattern, which in addition can

also be variable between different populations of a species.

For nothobranchiids it is assumed that male colouration

is important for female mate choice within and between

species (Amiet 1987; Brosset & Lachaise 1995; Kullmann

& Klemme 2007) and Van der Zee et al. (2007) presume

that especially the colour pattern of the caudal peduncle

and unpaired fins might have an important role.

With the description of A. musafirii we presented a pre-

liminary mtDNA analysis for some species of

Aphyosemion, including A. pseudoelegans (Van der Zee

& Sonnenberg 2011). We indicated an incongruence of

mtDNA results and phenotype within the samples of the

Cuvette centrale. This is also true for the A. pseudoele-

gans samples, which were labelled within this phyloge-

ny (Van der Zee & Sonnenberg 2011, fig.9) as A. sp. Cu-

vette (sample RS1019) or A. sp. aff. elegans (samples

RS1508 & RS1509). These samples turn out in different

parts of the phylogeny, but are grouped together in a pre-

liminary analysis of a fragment of the nuclear LSU (un-

published data). Additionally, a population originating

from Bandundu belongs to A. pseudoelegans according to

©ZFMK

10 Rainer Sonnenberg & Jouke R. Van der Zee

its phenotype with the diagnostic dark red dorsal fin. This

is well visible on available pictures and specimens deposit-

ed in the MRAC from a collection in 1982. However, the

published mtDNA cytochrome b sequence (Murphy &

Collier 1999, as Aphyosemion cognatum) deviates from

any of our sequences (results not shown).

Due to some incongruencies between mtDNA, nuclear

DNA, and phenotype we hesitate at the moment to dis-

cuss the phylogenetic position of the different species and

the cause of such incongruencies, until further samples and

DNA data are available. A potential cause for incongru-

encies can be mitochondrial introgression, which we dis-

cussed in the recent publication (Van der Zee & Sonnen-

berg 2011) and which is known to occur in the notho-

branchiid genus Chromaphyosemion (Sonnenberg 2007).

But in the case of incongruent DNA and phenotype data,

it has to be considered that also nuclear DNA and mtD-

NA can be affected. An example of homogenization of nu-

clear rRNA genes in F1 hybrids is given by Wang et al.

(2010). This would make it impossible to detect such

crosses, which usually is possible with nuclear sequence

data (see e.g. Sonnenberg et al. 2007). In addition, hybrid

speciation as in some African cichlids (Schliewen & Klee

2004; Schwarzer et al. 2011) might also not be excluded,

especially as the species involved are in contact within the

Cuvette centrale. The available data indicate a complex

pattern within the Aphyosemion of the Congo Basin, and

more data will be necessary to understand the evolution

and phylogeography of these fishes (Van der Zee & Son-

nenberg 2011).

Currently, it appears that A. pseudoelegans is restrict-

ed to the Cuvette central on the left bank of the Congo,

whereas A. elegans is also found further north and on the

right bank of the Oubangui in the Republic of Congo, but

not in the south and southeast of the Cuvette (Fig. 1). Their

distribution areas overlap at least in the region of the low-

er Tshuapa River, were they can occur in sympatry or close

parapatry, e.g. around Boende (Fig. 1). Aphvosemion pseu-

doelegans can also be found in syntopy with other

Aphyosemion, e.g. at Ikela with an undescribed species on-

ly known from preserved specimens in the MRAC, and

near Lui Kotale with A. sp. aff. castaneum (see Van der

Zee & Sonnenberg 2011).

COMPARATIVE AND ADDITIONAL NON-TYPE

MATERIAL

Aphyosemion elegans (Boulenger, 1899): MRAC 990-992, Syn-

types, Democratic Republic of Congo, Equateur Province, Tshua-

pa Basin, Bikoro, coll. P. Delhez, no year, MRAC 993, Syntype,

Democratic Republic of Congo, Equateur Province, Tshuapa

Basin, Coquilhatville (present Mbandaka), coll. P. Delhez, no

year; MRAC 1488, Democratic Republic of Congo, Uélé (not

specified), coll. A. Hutereau, 5.1.1905; MRAC 21628, Demo-

cratic Republic of Congo, Equateur Province, Tshuapa Basin,

Eala, coll. Van Oye, 2.V.1923; MRAC 21636-639, Democratic

Bonn zoological Bulletin 61 (1): 3-12

Republic of Congo, Orientale Province, Mondimbi, coll. J.

Ghesquiére, XI.1927; MRAC 21642—648, Democratic Repub-

lic of Congo, Equateur Province, Tshuapa Basin, Eala, coll. Van

Oye, 2.V.1923; MRAC 21649-653, Democratic Republic of

Congo, Orientale Province, Mondimbi, coll. J. Ghesquiere,

X1.1927; MRAC 21670-682, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Eala, coll. Van Oye,

2.V.1923; MRAC 21683-697, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Eala, coll. Van Oye,

2.V.1923; MRAC 23539-546, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Eala, coll. Van Oye,

2.V.1923; MRAC 44888-894, Democratic Republic of Congo,

Equateur Province, Sources 4 Gemena, coll. P. Henrard, 1935;

MRAC 45973999, Democratic Republic of Congo, Equateur

Province, Tshuapa Basin, Eala, coll. J. Ghesquiere, [V.1936;

MRAC 46000-46049, Democratic Republic of Congo, Equateur

Province, Tshuapa Basin, Eala, coll. J. Ghesquiére, IV.1936;

MRAC 72763, Democratic Republic of Congo, Orientale

Province, Buta, coll. J. Hutsebaut, 1949; MRAC 79229-79236,

Democratic Republic of Congo, Equateur Province, Tshuapa

Basin, Bokuma, coll. P. Lootens, VII.1952; MRAC

80778-80780, Democratic Republic of Congo, Equateur

Province, Tshuapa Basin, Bokuma, coll. P. Lootens, 1952;

MRAC 93724-93773, Democratic Republic of Congo, Equateur

Province, Tshuapa Basin, Bokuma, coll. P. Lootens, HI.1954;

MRAC 96831841, Democratic Republic of Congo, Equateur

Province, Tshuapa Basin, Bonguma 8 km S of Bokuma, coll. P.

Lootens, 1952; MRAC 98696-699, Democratic Republic of

Congo, Equateur Province, Tshuapa Basin, Bonguma, coll. P.

Lootens, 1955; MRAC 73-23-P-11233—302, Democratic Repub-

lic of Congo, Equateur Province, Tshuapa Basin, Boende, coll.

R. Philippe, VIII.1956.

Aphyosemion pseudoelegans: AMNH I-252189, Democratic Re-

public of Congo, Equateur Province, riv. Ta’Simon a Boaugi,

coll. R. Monsembula, 23.V.2010; MRAC 21634-635, Democrat-

ic Republic of Congo, Equateur Province, Flandria, coll. G. Hul-

staert, [V.1928; MRAC 30113, Democratic Republic of Congo,

Equateur Province, riv. Momboyo, Flandria, coll. G. Hulstaert,

15.1V.1930; MRAC 30123, Democratic Republic of Congo,

Equateur Province, riv. Momboyo, Flandria, coll. G. Hulstaert,

15.1V.1930; MRAC 30130-134, Democratic Republic of Con-

go, Equateur Province, riv. Momboyo, Flandria, coll. G. Hul-

staert, 18.X.1931; MRAC 30203-211, Democratic Republic of

Congo, Equateur Province, riv. Momboyo, Flandria, coll. G. Hul-

staert, 16.1V.1931; MRAC 30222259, Democratic Republic of

Congo, Equateur Province, riv. Momboyo, Flandria, coll. G. Hul-

staert, 1931; MRAC 37840-843, Democratic Republic of Con-

go, Equateur Province, riv. Momboyo, Flandria, coll. G. Hul-

staert, 15.VIH.1931; MRAC 47823—830, Democratic Republic

of Congo, Equateur Province, Tshuapa Basin, Flandria, coll. G.

Hulstaert, 1936; MRAC 72531—545, Democratic Republic of

Congo, Equateur Province, riv. Iolo, Flandria, coll. G. Hulstaert,

13.11.1948; MRAC 79237, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Bokuma, coll. P. Lootens,

VII.1952; MRAC 79908-909, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Bokuma, coll. P. Lootens,

10.VIL.1952; MRAC 80709-80777, Democratic Republic of

Congo, Equateur Province, Tshuapa Basin, Bokuma, coll. P.

Lootens, 1952; MRAC 96830, Democratic Republic of Congo,

Equateur Province, Banguma a 8 km Sud de Bokuma, coll. P.

Lootens, 1954; MRAC 96688-693, Democratic Republic of

Congo, Equateur Province, Tshuapa Basin, Bokuma, coll. P.

Lootens, 1.1954; MRAC 98700-701, Democratic Republic of

Congo, Orientale Province, Tshuapa River, Ikela, coll. P. Lootens,

©ZFMK

Aphyosemion pseudoelegans from the Congo Basin 1]

15.1V.1955; MRAC 99000, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Marais de la Kea, Boende,

coll. R. Philippe, IV.1955; MRAC 99753, Democratic Repub-

lic of Congo, Equateur Province, riv. Momboyo, Loselinga, coll.

G. Hulstaert, 5.V.1955; MRAC 99746—752, Democratic Repub-

lic of Congo, Riv. Loe, unknown location, coll. G. Hulstaert,

16.V.1955; MRAC 101891—915, Democratic Republic of Con-

go, Equateur Province, Tshuapa Basin, marches around Boende,

coll. R. Philippe, VII.1955; MRAC 131305-311, Democratic Re-

public of Congo, Orientale Province, Tshuapa River, Losaka

stream, Ikela, coll. H. Matthes, 9.1X.1959; MRAC 175812-842,

Democratic Republic of Congo, Equateur Province, Tshuapa

Basin, Boende, coll. P. Lootens, [.1966; MRAC 178018—023,

Democratic Republic of Congo, Equateur Province, Tshuapa

Basin, Boende, coll. P. Brichard, 1969; MRAC 73-23-P-

1104042, Democratic Republic of Congo, Equateur Province,

Tshuapa Basin, Boende, coll. R. Philippe, 1956; MRAC 73-23-

P-11043-232, Democratic Republic of Congo, Equateur

Province, Tshuapa Basin, Boende, coll. R. Philippe, VI.1956;

MRAC 73-23-P-11303-542, Democratic Republic of Congo,

Equateur Province, Tshuapa Basin, Boende, coll. R. Philippe,

1X.1956; MRAC 75-35-P-37—40, Democratic Republic of Con-

go, Orientale Province, district Ikela, Yetsi, coll. P. Lootens, VI-

11.1972; MRAC 79-09-P-732—740, Democratic Republic of Con-

go, Orientale Province, Ikela territory, Yayama, coll. J. Lambert,

17.V.1958; MRAC 84-25-P-3, Democratic Republic of Congo,

Equateur Province, Boteka, coll. P. Lootens, I.1984; MRAC 85-

47-P-1—4, Democratic Republic of Congo, Bandundu Province,

Kasai drainage, Bandundu, coll. K. Klute, X.1982; MRAC 2010

Congo Expedition, Location 66, stream near Lieki opposite

Yangambi, coll. J. Snoeks & E. Vreven, 27.V.2010.

Acknowledgements. The authors wish to thank the following

persons for their help in realizing this paper: H. Ott

(Monchengladbach, Germany) for providing pictures and spec-

imens of A. pseudoelegans and A. elegans; M. Stiassny for pro-

viding a picture and a drawing of A. pseudoelegans present in

the AMNH (New York, USA) collection, D. Neumann and U.

Schliewen (ZSM, Miinchen, Germany) for loan of specimens un-

der their care and providing data and photos; M. Parrent, J.

Snoeks, and E. Vreven (MRAC, Tervuren, Belgium) for support

and access to the collections under their care, M. Parrent in ad-

dition for the translation of the abstract into French, F. Herder

(ZFMK, Bonn, Germany) for access to the digital X-ray ma-

chine, D. Tautz (MPI, Plén, Germany) for access to the micro

Computer Tomograph, and A.C. Schunke (MPI, Plén, Germany)

for help with earlier versions of the manuscript. In addition, we

are grateful for the critical comments of two anonymous review-

ers.

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' Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

? Max-Planck-Institut ftir Evolutionsbiologie, Department of Evolutionary Genetics, August-Thienemann-Strasse 2,

D-24306 Plon, Germany. Corresponding author. E-mail: r.sonnenberg.zfmk(@uni-bonn.de

> Diederichsstrasse 45, D-42855 Remscheid, Germany.

Abstract. Scriptaphyosemion wieseae, new species, is described from collections in northern Sierra Leone. Results of a

small DNA study indicate, that S. bertholdi from the southeastern part of the country is the closest relative. Both species

can be distinguished by male and female color pattern and male caudal fin shape.

Résumé. Scriptaphyosemion wieseae, une nouvelle espéce, est décrite du Nord de la République de Sierra Leone. Le ré-

sultat d’une petite étude moléculaire ADN montre que son espéce sceur est S. bertholdi, présente au Sud-Est du pays. Les

deux espeéces se distinguent par les couleurs du male et de la femelle et par la forme de la nageoire caudale du male.

July 2012

Key words. Upper Guinean ichthyofaunal province, taxonomy, DNA, phylogeny.

INTRODUCTION

The West African killifish genus Scriptaphyosemion Rad-

da & Puirzl, 1987 is relatively species rich, compared to

the closely related genera Archiaphyvosemion Radda, 1977,

Callopanchax Myers, 1933, and Nimbapanchax Sonnen-

berg & Busch, 2009. At present 18 specific names are

available in Scriptaphyosemion (Eschmeyer 2011; Huber

2007; Lazara 2001), but only 10 (Huber 2007; Wildekamp

& Van der Zee 2003) to 13 (Costa 2008; Lazara 2001)

species are currently accepted as valid, depending on au-

thor and concept (Huber 2007; Eschmeyer 2011; Lazara

2001; Scheel 1990; Wildekamp 1993). Among the West

African killifish genera, species number in Script-

aphyosemion is only exceeded by the Upper Guinean

species of the related genus Epiplatys Gill, 1862 (Costa

2008; Huber 2007; Wildekamp & Van der Zee 2003).

In nothobranchiid killifishes, most species are diagnosed

by male color pattern (Amiet 1987; Sonnenberg 2007),

which is, however, increasingly supplemented by molec-

ular approaches (e.g. Agnése et al. 2009; Legros et al.

2005; Legros & Zentz 2007a, b; Sonnenberg 2007, 2008;

Sonnenberg & Blum 2005; Sonnenberg & Busch 2009,

2010; Sonnenberg et al. 2006, 2011; Van der Zee & Son-

nenberg 2011). In Scriptaphyosemion, female color pat-

tern has also been used as character for diagnosing species

or groups of species (Roloff 1971a).

In most species of Scriptaphyosemion, interspecific vari-

ance in male color pattern is accompanied by intraspecif-

ic variance between and within populations, a phenome-

Received: 06.02.2012

Accepted: 15.06.2012

non which may confuse species identification; however,

some of the most variable species might be composed of

more than one species (see e.g. figures or drawings in

Seegers 1997; Wildekamp 1993). This combination of in-

traspecific variability, inclusion of probably different

species under one taxon, and the lack of a specific diag-

nosis in many species descriptions contributed to some

confusion in Scriptaphyosemion taxonomy (see e.g.

Scheel 1990). More than a decade ago, Murphy et al.

(1999) published the first molecular genetic phylogeny of

Scriptaphyosemion, based on the sequences of the partial

cytochrome b and 12S rRNA genes and including eleven

species. This phylogeny gave a first insight into the rela-

tionships and an additional set of independent characters

for further taxonomic and phylogenetic research.

Species diversity of Scriptaphvosemion is highest in

Sierra Leone, Liberia, and Guinea. However, the distri-

bution area also includes Burkina Faso, Gambia, Guinea-

Bissau, Ivory Coast, Mali, and Senegal (Huber 2007; Pau-

gy et al. 1990; Teugels et al. 1988; Wildekamp 1993;

Wildekamp & Van der Zee 2003). In Sierra Leone, Script-

aphyosemion species are mainly known from the coastal

lowland, the northern and eastern killifish fauna is only

little known (Etzel 1977, 1978a, b, 1984; Huber 2007;

Wildekamp 1993; Wildekamp & Van der Zee 2003).

In 1993, the second author collected together with B.

Wiese nothobranchiids in the northern part of Sierra Leone

(Busch 1995a), including a probably new species near

Corresponding editor: F. Herder

Table 1. List of specimens used for the DNA analyses with locality data and GenBank accession numbers (ND2+tRNAs, mtL-

Rainer Sonnenberg & Eckhard Busch

SU rDNA, nLSU rDNA). * Sequences published in Sonnenberg & Busch (2009).

Collection locality

GenBank acc. no.

Species sample Country

Epiplatys sexfasciatus RS1364 Gabon

Scriptaphyosemion bertholdi RS1402 Sierra Leone

S. brueningi RS1377 Sierra Leone

S. brueningi RS1403 Sierra Leone

S. cauveti RS1680 Guinea

S. chaytori RS1379 Sierra Leone

S. fredrodi RS1398 Sierra Leone

S. geryi RS1381 Sierra Leone

S. guignardi RS1380 Guinea

S. liberiense RS1757 Liberia

S. roloffi RS1376 Sierra Leone

S. schmitti RS1755 Liberia

S. Sp. RS1378 Sierra Leone

S. wieseae RS1382 Sierra Leone

S. wieseae RS1686 Sierra Leone

S. wieseae RS1687 Sierra Leone

G 02/91

SL 93/51

SL 03/04

SL 89, Perie

GM 97/05

SL 93/23 Ngabu

SL 89, Matanga

SL 03/09, Robis 1

GM 97/13

SL 03/10

RL 83/121

SL 03/18

SL 03/16

SL 03/15

SL 03/17

JX124276, FJ872035°, FJ872049°

JX124261, JX124233, JX124247

JX124262, JX124234, JX124248

JX124263, JX124235, JX124249

JX124264, JX124236, JX124250

JX124265, JX124237, JX124251

JX124266, JX124238, JX124252

JX124267, FJ872033°, FJ872047°

JX124268, JX124239, JX124253

JX124269, JX124240, JX124254

JX124270, JX124241, JX124255

JX124271, JX124242, JX124256

JX124272, JX124243, JX124257

JX124273, JX124244, JX124258

JX124274, JX124245, JX124259

JX124275, JX124246, JX124260

Lenghe Curoh. In 2003 they revisited this locality and

found two additional collection localities of this species

(Fig. 1). Preliminary crossing experiments by engaged

aquarists (Vandersmissen 2003) indicated its probable in-

compatibility with S. bertholdi (Roloff, 1965), which was

considered to be the potentially closest relative. Here, we

provide a formal species description and present the phy-

logenetic position of the new species within Script-

aphyosemion, based on mitochondrial and nuclear DNA

sequences.

MATERIAL AND METHODS

The species description is based on collections made in

Sierra Leone by the second author and B. Wiese during

their second visit in this area in November 2003. Speci-

mens were deposited in the Royal Museum for Central

Africa (MRAC, Tervuren, Belgium), the Zoologische

Staatssammlung Miinchen (ZSM, Munich, Germany), and

the Zoologisches Forschungsmuseum Alexander Koenig

(ZFMK, Bonn, Germany).

Bonn zoological Bulletin 61 (1): 13-28

For counts and measurements, see Sonnenberg & Busch

(2009, 2010). All measurements were taken with a digi-

tal caliper, partly under a dissecting microscope, to the

nearest 0.1 mm. Measurements, including those of the

head, are presented as percentages of standard length (SL).

The head region of one specimen was scanned with a

Scanco viva CT40 at the Max-Planck-Institut fiir Evolu-

tionsbiologie in Plon, digital X-ray images were made with

a Faxitron LX-60 Digital Specimen Radiography System

at the ZFMK. Fin rays of unpaired fins were counted on

digital X-ray images; the relative position of the first dor-

sal fin ray to the anal fin (D/A) is estimated following Son-

nenberg & Schunke (2010). Nomenclature of the neuro-

mast system on the head follows Scheel (1968), for supra-

orbital (or frontal) squamation Hoedeman (1958). Male

and female color patterns are compared with that given

in the respective description and on pictures or drawings

published in the literature.

As species concept we employ the pragmatic approach

of Moritz et al. (2000) and recognize species by diagnos-

tic combinations of characters.

©ZFMK

New Scriptaphyosemion from Sierra Leone 1S

Fig. 1. Distribution map of Scriptaphyosemion in Sierra Leone based on data from Huber (2007) and own data of the second au-

thor. White square: S. bertholdi; black triangle: S. brueningi; white circle: S. chaytori; dark grey triangle: S. fredrodi; dark grey

square: S. geryi; light grey circle: S. guignardi; white triangle: S. liberiense; black circle: S. roloffi; black square: S. wieseae. TT

marks the type localities of S. bertholdi and S. wieseae.

A DNA dataset was prepared for 15 Scriptaphyosemion

specimens, including 11 species, and Epiplatys sexfascia-

tus Gill, 1862 as outgroup. We sequenced the complete

mitochondrial ND2 gene and adjacent tRNAs, and frag-

ments of the mitochondrial and nuclear LSU rDNA genes.

Parts of the sequences were published in earlier papers of

the authors; all samples are listed with GenBank acces-

sion numbers and collection locality in Table 1. For de-

tails on the DNA methods, see Sonnenberg & Busch

(2009, 2010) and Sonnenberg et al. (2007).

Abbreviations used in Table 2 and Figures 2—3: BER =

S. bertholdi, BRU = S. brueningi (Roloff, 1971b), CAU

= S. cauveti (Romand & Ozouf, 1995), CHY = S. chay-

tori (Roloff, 1971a), FRE = S. fredrodi (Vandersmissen

et al., 1980), GER = S. geryi (Lambert, 1958), GUI = S.

guignardi (Romand, 1981), LIB = S. liberiense

(Boulenger, 1908), ROL = S. roloffi (Roloff, 1936), SMI

= §. schmitti (Romand, 1979), Ssp =S. sp. SL 03/18, WIE

= S. wieseae, new species, SEX = E. sexfasciatus.

Bonn zoological Bulletin 61 (1): 13-28

Sequences were aligned with ClustalX 1.8 (Thompson

et al. 1997), checked by eye, and merged in a single align-

ment in BioEdit 7.0.5.3 (Hall 1999). As the ND2 se-

quences of three species are not complete, missing sites

were filled with N [S. liberiense (560 bp), S. schmitti (616

bp), and S. cauveti (58 bp)].

The protein coding gene was translated into the corre-

sponding amino acid sequence to check for a functional

coding sequence, and base composition of mitochondri-

al sequences was estimated with MEGA 4.1 beta (Tamu-

ra et al. 2007). Uncorrected p-distances with pairwise ex-

clusion of missing data were calculated in MEGA 4.1 be-

ta (Tamura et al. 2007). Number of constant, variable, and

informative characters were estimated with PAUP 4.0b10

(Swofford 1998).

Data were analyzed using a maximum parsimony (MP)

and a Bayesian approach (BA). Maximum parsimony

analyses were performed with PAUP 4.0b10 (Swofford

1998) with the exclusion of uninformative positions and

©ZFMK

16 Rainer Sonnenberg & Eckhard Busch

BER 1402

99 / 98 / 82 WIE 1382

100 / 100/95 WIE 1687

WIE 1686

92 / 72 /-- CHY 1379

GUI 1380

100 / 100 / 100 Ssp 1378

FRE 1398

100 / 100 / 100 p77 BRU 1377

96/91/53 100 / 100 / 100 BRU 1403

LIB 1757

100 / 85 / 69 ROL 1376

GER 1381

CAU 1680

SM1I1755

SEX 1364

Fig. 2. Results of the maximum parsimony analyses. Shown is one out of three equally short trees from the analysis with the

complete dataset. Values left to nodes are the corresponding bootstrap values of all three analyses (complete dataset / third protein

coding position excluded / alignment columns with missing data excluded). Values for nodes were only given, if at least one analy-

sis has a bootstrap support above 75 %. Tree length = 1262, consistency index (CI) = 0.5166, homoplasy index (HI) = 0.4834, re-

tention index (RI) = 0.5570, rescaled consistency index (RC) = 0.2878.

gaps coded as fifth character, heuristic tree searches were

performed with random addition of sequences for 1,000

replicates. Bootstrap tests were performed with random

addition of sequences for 100 replicates and 10,000 boot-

strap replicates. Three different analyses were made, one

with the complete dataset, one with the exclusion of third

protein coding positions, and a third with exclusion of re-

gions, in which one of the above listed specimens has

missing data. Otherwise, all parameters were identical.

Bayesian analyses were performed with MrBayes 3.1.2

(Huelsenbeck & Ronquist 2001), data were partitioned in

mitochondrial non protein coding regions (tRNA, mtLSU),

protein coding regions (ND2), and the nuclear LSU se-

quences. Number of generation was set to 10,000,000 with

sampling of every 1,000 generations and the results of the

first 10% of the generations were discarded as burn-in time

after checking that the runs had reached the stationary

phase at this point. Four analyses were performed: the two

alignments, one with the complete data set and one with

the exclusion of regions, in which one of the above men-

tioned specimens has missing data, were both performed

with NST=2 and NST=6.

Bonn zoological Bulletin 61 (1): 13-28

All trees were rooted with Epiplatys sexfasciatus as out-

group. Only nodes with bootstrap support values = 75%

or posterior probabilities = 0.95 were considered as sup-

ported by the data. Nodes below these values were con-

sidered as unsupported by the data and not resolved.

RESULTS

DNA Analysis. (Figs 2—3, Table 2). The complete align-

ment of the ND2, tRNA, and both LSU sequences has a

length of 3199 bp including alignment introduced gaps for

a total of 16 specimens (503 bp tRNAs, 1047 bp ND2, 481

bp mtLSU rDNA, 1168 bp nLSU rDNA). Of these, 2278

bp are constant, 456 bp variable but uninformative, and

465 bp parsimony informative. The ND2 sequence trans-

lates into 349 amino acids with no unexpected stop codon,

the base composition of all mitochondrial genes shows the

typical base composition for this organelle genome with

an anti-G bias (Zhang & Hewitt 1996). Average uncorrect-

ed pairwise distance for all sequences is 7.4 %, maximum

observed distance is 16.6 % between outgroup and ingroup

specimens, maximum observed distance between ingroup

specimens is 9.0 %, further details are given in Table 2.

©ZFMK

New Scriptaphyosemion from Sierra Leone

VE MOV O/C SOULE OLY CL LOISEL LO/OEL OOIGEL 99/ECl 9O/EEl LOSI OL/8 Hl “DO/S.El OOS sel SLi9t/ISEl

IS¢T/TIL /EVT/TIL /S HT /E OL /E VT POL PBL /E TL OT /E9T /0'9T /S'ST /9VT/NOT /9HT/DOT (PHT VOL WPCISOT/CHT/I9OT /OPT/E9T /6ET/TOI /PvT/S OT = =—P9ETKAS

T0/r0 10/990 €T1/0E SiG (826 vale Saxe. Cal: / VAS ITIVE GIVE, CKEVIE GOL/ICE O1/E —LOUSC UU Ota

ITT/LO /TIT/LO /06/ES MWOL/8S /BIT/L9 /TIT/S /TOT/T9 /8CL/9L /68/e0S V8IES /EVT/T8 /EIT/79 /61T/S9 /TS/0€ LE9THIM

00/70 E1/0E DGC S/S C™ Calva CHO, SS GiGVee ASIEN Oc/ce B8I/ve 90/97 D101/SaG- Only lel

160/50 /L8/7S /SOT/BS WIT/S9 /CIT/VS /86/68 CTL § /S8/TS V8/IES ISPL/C8 WWIL/C9 (ITO /Sb/LC 989TAHIM

CALNE OG Dive~ SMEG sGaliiCee CMB E GCI. CEN/0ie Oc/OE BI/VE YO/ET 90/ET OT/80

IL8/TS = ISOL/8S /BIL/89 /BOL/TS /S6/8S /TCL/EL /98/eS IS8/ES /EVL/T8 WIL/T9 /BIL/P9 (Lv /8C C8ETAIM

CN Ge HAGIES. (LiQviGy: On/9ie VALS GieiGw. 0:0 /9°0 el/vy 81/cVv 60/TE 60/TE

IQOL/V9 /611/89 /FIT/SS /LOT/99 /TEL/OL /6T/ST /TOT/T9 /OST/98 (VITO /61T/99 /C6/S'S SLetdss

SC/Cb CC/OV Lé/6S 8C/9b LT/IDP Tees, CC/ry VC/TE VC/SE 8C/6V

ITOL/LS /LLIOE /O1T/L9 /SIT/E9 /9OT/09 /601/F9 /66/VS /TIT/6S JOTI /6S /8IT/S9 SSLTTNS

80 /9'P LV Oi ESE LOW Ov. TUE “GrC/8ie. — GiOV/BE GO/BE STL?

IPTL/6S /8TL/SL /66/6S /VTL/69 /CCL/89 /PST/88 /TCL/69 /TCI1/69 /LIT/99 9LETTON

VI/SS LI/TS OT/BE Sl/pp LI/6ry SO/TT SO/TT VI/9E

I6TL/S9 /BTI/S9 /BIL/9S /BCL/PO /BVI/TL /ev/TC fv /TC WIT/9S = LSLTaYT

GGNAL “WAAC OT/'E IRGiGYA Cully Sav, el/9p LIVE

IZ EL/E8 /6 01/99 /66/T9 /VST/L8 /8TI/PL JOEL /SL /LO/6S § O8ETIND

CT /0'V Ctr ST/RE 61/TV 61T/Cb €T/VE

ISEL/T8 /BTL/LL /SS1/06 /OET/LL MEL /6L /PCL/SL T8eTHao

S/S 8aliOi GiOrivae 60/veE ELE

/66/19 /VST/98 /LIT/99 /TVCL/89 /68/eS s6eTHdt

SiG Sites VANIORE VIE OCISTC

ITVL/98 [TTL /OL /SCL/CL /C8/CS 6LETAVO

OT /8°E CHS Ss MO GIES

IPvVI/T8 /9VIL/T8 /SbI/€8 O89TAWO

00/00 60/L7T

/60/p0 /O1L/T9 ecOrTONa

60 /LC

/CIL/€9 LLeToNa

L89TAIM 989THIM T8ETAIM § SLetdss SSLIINS9LETTON LSLIAIT O8€HINDI8ElNAD 8oEladA OLETAVD O89IAVO cOrInNUa LLEINUA LOFINad ajdures

“VN! SJ Jeajonu oy) / VNC NS [epuoyooyiul oy} / SYNRI+ZON [PEpuoysoyru oy} / oseep dj0[dUIOS oY} IOJ vIEP BUISSIUL JO UOISN[OXO osIMIIed YIM soouRjsIp-d poyoouIOIUy)-*Z AIGUL

©OZFMK

Bonn zoological Bulletin 61 (1): 13-28

Rainer Sonnenberg & Eckhard Busch

A BER 1402

1.0/0) WIE 1382

WIE 1686

1.0/1.0 wre 1687

CHY 1379

GUI 1380

1.0/1.0 p= Ssp 1378

FRE 1398

0.96/0.99 1.0/1.0* Sood

ot — GER 1381

1.0/1.0 = BRU 1377

1.0/1.0 BRU 1403

1.0/1.0 | LIv 91.07

LIB 1757

CAU 1680

SMI 1755

SEX 1364

B BER 1402

1.0/1.0 erences

1.0/1.0] p WIE 1382

0.98/0.98 WIE Fese

GUI 1380

0.95/0.89* EERE IGGO

ROL 1376

0.97/0.99 1.0/1.0 GER 1381

Ssp 1378

1.0/1.0be Fre 1398

1.0/1.0 = BRU 1377

1.0/1.0] L BRU 1403

LIB 1757

CAU 1680

SMI 1755

SEX 1364

0.02

Fig. 3. Results of the Bayesian analyses. Shown is the consensus tree of the analysis with the complete dataset and NST=6 (A)

and the consensus tree of the dataset with excluded alignment columns with missing data and NST=6 (B). Values left to nodes are

the corresponding posterior probability values, except for values marked with * which are on the right side of the corresponding

node (complete dataset, NST=2 / complete dataset, NST=6 in A; alignment columns with missing data excluded, NST=2 / align-

ment columns with missing data excluded, NST=6 in B). Values for nodes were only given, if at least one analysis has a posteri-

or probability above 0.95.

Bonn zoological Bulletin 61 (1): 13-28 ©ZFMK

New Scriptaphyosemion from Sierra Leone 19

All three maximum parsimony analyses result in near-

ly the same tree topology and differ only in branch length

and bootstrap support values; here, only one of three short-

est trees of the complete dataset is presented (Fig. 2). The

Bayesian analyses for the complete and the reduced

datasets show slightly different tree topologies, independ-

ent if the analyses were performed with NST=6 or 2.

Shown are the results for the complete and the reduced

dataset of the NST=6 analyses, on the nodes posterior

probabilities for both analyses for each dataset are present-

ed if the values are equal to or above 0.95 (Fig. 3 A+B).

Scriptaphyosemion schmitti and S. cauveti are support-

ed as basal to all other species of the genus (Figs 2-3).

As S. schmitti was placed in the study of Murphy et al.

(1999) within a clade containing the other Script-

aphyosemion species studied, we checked for possible

causes of this discrepancy. As the sequences studied here

have no overlap with those of Murphy et al. (1999), we

compared a cytochrome b fragment of our sample with

the data deposited in GenBank. However, both samples

differ in only one nucleotide (results not shown), and a

Blast search with our analysed sequences (see Table 1) on-

ly returned the highest similarity with other Script-

aphyosemion sequences. To exclude that the basal posi-

tion of S. schmitti is an artifact due to the missing sequence

data, we also performed the analyses with the reduced

dataset, which did not contain missing data, but gave the

same results (Figs 2-3). However, further samples should

be studied for a confirmation of the basal position, espe-

cially as this node is supported neither by bootstrap nor

posterior probability values.

Our results indicate a closer relationship between S.

liberiense and S. brueningi, S. roloffi and S. geryi, S. fred-

rodi and a probably undescribed species (see under Dis-

cussion), and S. bertholdi and S. wieseae, new species. A

closer relationship between S. chaytori and S. guignardi

occurs in all resulting trees (Figs 2—3), but is well sup-

ported only in the Bayesian analyses of the complete

dataset. Both species were in addition grouped with S.

roloffi and S. geryi in the analyses of the reduced dataset,

but not in the maximum parsimony analyses and the

Bayesian analyses of the complete dataset (Figs 2-3).

All analyses support a monophyletic group formed by

all Scriptaphyosemion except of S. schmitti and S. cau-

veti (Figs 2—3). Another well supported subgroup is

formed by S. bertholdi, S. wieseae, new species, S. chay-

tori, S. guignardi, S. fredrodi, and S. sp. (see under Dis-

cussion); only in the analyses of the reduced dataset S.

roloffi and S. geryi were also included in this group (Fig.

2-3). To this subgroup, S. brueningi and S. liberiense,

probably is the sister group, a hypothesis only significant-

ly supported in the Bayesian analyses (Fig. 3).

The phylogenetic results show that S. bertholdi is the

well supported sister species to S. wieseae, new species,

and indicates no closer relationship to other mentioned

Bonn zoological Bulletin 61 (1): 13-28

species. The genetic distance values (Table 2) show a sim-

ilar divergence as between S. /iberiense and S. bruenin-

gi.

Scriptaphyosemion wieseae, new species

(Fig. 4, Table 3)

Scriptaphyosemion sp. Lenghe Curoh: Busch (1995a),

Vandersmissen (2003)

Holotype. ZFMK 47680, male, 36.1 mm SL, Sierra

Leone, Lenghe Curoh, Sanghi or Sanigi River, field num-

ber SL 03/16 (9°28,04’ N, 11°40,58’ W), E. Busch and B.

Wiese, 18. Nov. 2003.

Paratypes. ZFMK 47681-47692, 8 males, 4 females,

19.5-34.7 mm SL, paratopotypes, collected with the holo-

type.

MRAC B2-06-P-1—2, 1 male, 22.9 mm SL, | female, 21.5

mm SL, paratopotypes, collected with the holotype.

ZFMK 47693-47707, 7 males, 8 females, 18.8—41.3 mm

SL, Sierra Leone, Fadugu, at the bridge over the Mabakah

River, field number SL 03/15 (9°23,52’ N, 11°45,76’ W),

E. Busch and B. Wiese, 18. Nov. 2003.

ZSM 41596 (2), 1 male, 28.4 mm SL, 1 female, 22.3 mm

SL, collected with ZFMK 47693-47707.

Non-type DNA samples. RS1686, | male, DNA vouch-

er, preserved in 96 % ethanol, collected with ZFMK

47693-47707, field number SL 03/15; RS1382, 2 males,

1 female, DNA voucher, preserved in 96 % ethanol, col-

lected with the holotype; RS1687, 1 male, DNA vouch-

er, preserved in 96 % ethanol, Sierra Leone, Kabala, col-

lected in rice swamp, field number SL 03/17 (9°35’ N,

11°33’ W), E. Busch and B. Wiese, 18. Nov. 2003.

Diagnosis. Scriptaphyosemion wieseae is distinguished

from its congeners by a unique combination of male and

female coloration characters. Males have a blue body col-

or, red dots on side forming irregular, interrupted red

stripes, especially on caudal peduncle a more reticulated

pattern, unpaired and pelvic fins centre blue with a nar-

row blue margin and a broad red submargin, on anal and

dorsal fin a small number of basal red blotches, centre of

caudal fin with some red blotches, pectoral transparent to

orange with a blue margin; females with a dark brown

midlateral stripe, interrupted on caudal peduncle, body on

lower half of side and ventrally orange vs. a mainly red

lateral body color in male S. cauveti; head and anterior

body yellowish to brownish, unpaired fin margins white

or yellow, and pectoral fn margin yellow to orange in male

S. schmitti; males with more or less regular oblique red

bars on side, at least on the caudal peduncle, fin margins,

especially of caudal and pectoral, often yellow, and fe-

males with dark brown streaks or blotches, similar to the

OZFMK

20 Rainer Sonnenberg & Eckhard Busch

Fig. 4. A) Male of Scriptaphyosemion wieseae, collection locality SL 03/16, not preserved. B) Female of Scriptaphyosemion wieseae,

collection locality, SL 03/16, not preserved.

red oblique bars of males on side, often with a dark blotch

at the root of the caudal in S. geryi and S. roloffi; males

with broad red blotches, often forming broad irregular ver-

tical stripes, caudal fin margins yellow, often with a small

white submargin followed by a broader red band, females

with irregularly distributed dark brown dots or short

oblique streaks or blotches dorsally on the caudal pedun-

cle, a weak anterior dark lateral band, and a dark blotch

on the posterior dorsal end of the caudal peduncle at the

caudal fin base in S. brueningi; often yellow, sometimes

whitish caudal fin margins in males, females with irreg-

Bonn zoological Bulletin 61 (1): 13-28

ularly distributed dark brown or red dots on side in S.

chaytori; males with dark red blotches on side, caudal fin

margins whitish to light blue or yellow, pectoral fin mar-

gin white, females with dark brown blotches or oblique

short streaks on dorsal half of caudal peduncle and a dark

blotch on posterior end of caudal peduncle at dorsal base

of caudal fin in S. fredrodi; males with light blue to blue-

green body color, many irregularly distributed small red

streaks or dots, submarginal stripes of unpaired fins dark

red to nearly black, margins light blue or whitish, some-

times on dorsal and often on anal absent, females with red

©OZFMK

New Scriptaphyosemion from Sierra Leone 21

Table 3. Morphometrics of Scriptaphvosemion wieseae, new species. Values in the table are based on 15 specimens, for which

all measurements and counts were taken (7 of each male and female paratypes). All measurements in percentages of standard length

(SL) except standard length in mm. TL = total length, HL = length of head, PD = predorsal fin distance, pPED = prepectoral fin

distance, pPD = prepelvic fin distance, pAD = preanal fin distance, DB = greatest body depth, DC = caudal peduncle depth, CL =

length of caudal peduncle, BD = base of dorsal fin, BA = base of anal fin, E = eye diameter, I = interorbital width, SD = standard

deviation.

Paratypes ¢ mean Paratypes 2 mean all Types mean all Types range all Types SD

Holotype

SL 36.0 25.1 24.7 25.6 21.5—36.0 4.0

TL 119.7 124.4 122.5 123.2 119.7-127.0 DED

HL 26.3 26.6 26.9 26.7 25.1-28.9 1.3

PD 66.0 65.1 64.9 65.0 61.2-67.8 2.0

pPED Poll 27.4 27.9 27.6 25.9-29.8 let

pPD 45.3 47.2 46.6 46.8 44.7—-49.4 Ie3)

pAD 57.3 57.8 SHES 5) lad! 54.9-59.8 IES

DB 19.5 19.8 19.3 19.6 18.7-21.7 0.8

DC 12.3 11.9 11.4 Med 10.9-12.3 0.5

CL 3 22.4 DOES) DOs) 21.1-23.6 0.7

BD 17.6 16.3 15.7 16.1 14.4-17.6 1.0

BA 21.1 20.4 19.1 19.8 17.9-21.5 1.0

E 5.4 6.5 6.5 6.4 5.4-7.3 0.5

I 11.2 OS 9.6 Od 8.6—1.2 0.6

or dark brown dots on side, on some specimens nearly lin-

ear arranged, on others more irregular, in S. guignardi;

mostly yellow or whitish caudal fin margins in males, ir-

regular dark blotches on dorsal half of side in females, ac-

cording to the first description (Boulenger 1908), in S.

liberiense.

Scriptaphyosemion wieseae males (Fig. 4A) differ from

its closest relative, S. bertholdi (Fig. 5A), by coarser and

often denser red pigmentation on body and fins and broad-

er dots or short streaks in the caudal fin, caudal fin trun-

cate vs. small dots on body and fins, especially in the cau-

dal fin, caudal rounded in S. bertholdi males. Females of

S. wieseae (Fig. 4B) have a midlateral more or less com-

plete dark stripe, often interrupted posterior to origin of

anal fin on caudal peduncle, ventral half of sides below

midlateral dark stripe often intense orange in wild caught

specimens or orange-brown in aquarium kept specimens

vs. irregularly distributed dark brown dots on scales on

dorsal half of side, pale brownish-orange on anterior side

in females of S. bertholdi (Fig. 5B).

Description. See Figure 4 for overall appearance and

Table 3 for morphometric and meristic values of the type

series. Scriptaphyosemion wieseae shows a strong sexu-

al dimorphism with males more colorful, usually larger,

and with slightly extended fin rays in unpaired fins. Body

cylindrical, elongate, and laterally slightly compressed.

Dorsal profile slightly convex, ventral profile anteriorly

slightly convex, slightly concave to nearly straight on cau-

Bonn zoological Bulletin 61 (1): 13-28

dal peduncle, greatest body depth approximately anteri-

or to origin of pelvic fin.

Snout slightly pointed, mouth directed upwards, lower

jaw longer than upper jaw, posterior end of rictus at the

level of the dorsal third to centre of eye. Teeth curved and

unicuspid, on dentary an outer row of larger teeth, inner

rows more or less regular with smaller teeth. Premaxilla

with an outer row of larger teeth and a small number of

short teeth irregularly on the inner part. Closed frontal neu-

romast system within one groove, preopercular neuromast

system with 5 pores.

Scales cycloid, body and head completely scaled, ex-

cept for ventral surface of head. Frontal squamation of G-

type; scales on lateral line 27-30 + 2-3 scales on caudal

fin base. Transverse rows of scales above pelvic fin 9; cir-

cumpeduncular scale row 12-14.

Tips of dorsal, anal, and, in some males, also caudal fin

slightly pointed, but no filamentous extensions, caudal fin

truncate. Fins in females are generally smaller and more

rounded. Dorsal and anal fin origin posterior to mid-body.

First dorsal fin ray posterior to origin of anal fin, situat-

ed above anal fin ray 4—6. Dorsal fin with 12—13 rays, anal

with 16-17 rays, caudal fin with 25—28 fin rays. Pectoral

fin rounded, fin origin on ventral half of side, just poste-

rior to operculum, directed posteriorly, not reaching pelvic

fin origin. Pelvic fin small, origin ventrally, directed pos-

teriorly, not or just reaching anal fin origin. Total number

of vertebrates 29-30, of which 13—14 have pleural ribs and

15—16 have haemal spines.

OZFMK

Rainer Sonnenberg & Eckhard Busch

Fig. 5.

bertholdi, SL 93/51, Sierra Leone, Victoria, not preserved.

Live coloration. Males. (Fig. 4A) Lateral head blue with

a slightly irregular red pattern typical for the endemic West

African genera Archiaphyosemion, Callopanchax, Nimba-

panchax, and Scriptaphyosemion and the related Epiplatys

(Van der Zee 2002). Lips dark grey, red line around mouth

along upper and lower lip, small dark grey dot at chin, iris

of eye grey, dorso-anterior with blueish or blue-greenish

Bonn zoological Bulletin 61 (1): 13-28

A) Male of Scriptaphyosemion bertholdi, SL 93/51, Sierra Leone, Victoria, not preserved. B) Female of Scriptaphyosemion

hue, a light, reflecting ring around pupil. Dorsal from head

to dorsal fin brownish, scales with some blue in centre and

irregular red margins, forming an interrupted reticulated

pattern. Flanks, unpaired fins and pelvic fins blue, dorsal

and caudal fin centre often with a yellow hue, giving a

more greenish impression of fin color. Red pigmentation

on side in three to four very irregular stripes, consisting

©ZFMK

New Scriptaphyosemion from Sierra Leone 23

of red blotches, especially on caudal peduncle looking

more like an interrupted reticulation. Unpaired and

pelvic fins with a narrow blue margin, sometimes slight-

ly lighter than color of fin centre and body; the blue mar-

gin is followed by a broad red submargin. On base of dor-

sal, anal, pelvic, and centre of caudal fin a small number

of red blotches, pectoral fin transparent with a blue mar-

gin and a, more or less complete, red submargin.

Females. (Fig. 4B) Body brownish, dorsally darker, on

ventral half of side and belly orange. Pigmentation on head

similar to males, but mostly darker brown instead of red.

Midlateral dark brown stripe, interrupted and slightly ir-

regular on caudal peduncle, scales in this dark stripe some-

times with small red scale margins; centre of unpaired and

pelvic fins brown-orange, with a very narrow blueish mar-

gin, and a faint, narrow red submarginal, except on cau-

dal fin. Above submarginal on anal and pelvic fins a small

area with blueish or greenish hue, centre of dorsal fin with

small red dots, basally larger than marginally. Pectoral fin

transparent.

Color in ethanol. Males. Body brownish, darker on back

and lighter on belly, red pigmentation pattern of live spec-

imens 1s now carmine or dark brown, on ventral side of

head a small grey blotch on chin. Fin color light brown,

unpaired fins with a whitish margin and a carmine or dark

brown sub-margin, some carmine or brown dots, pelvic

fins with a dark brown margin.

Females. Similar to males body brown with a darker

dorsal and lighter ventral part, on side the same dark

brown pigmentation as in live specimens, on ventral side

of head a grey blotch as in males. Fins light brown, on

dorsal several small brown dots.

Distribution and habitat. Scriptaphyosemion wieseae is

currently known from three localities in the upland of

northern Sierra Leone between the Sula and Wara Wara

mountains (Fig. 1); all of these are located within the Seli

or Rokel River basin. At the collection localities Fadugu

(SL 03/15) and Lenghe Curoh (SL 03/16), S. wieseae is

present in small rivers, at the former place together with

Epiplatys cf. njalaensis, Archiaphyosemion guineense, un-

determined characids, gobies, and cyprinids, probably

Barbus leonensis, B. stigmatopygus and a small undeter-

mined Barbus specimen; at the latter place it was collect-

ed together with A. guineense. At Kabala (SL 03/17), on-

ly asmall number of mostly juvenile specimens were col-

lected in large rice swamps. The second author visited this

area two times, in 1993 and 2003, and noticed an inten-

sified agricultural land usage.

Collections of the second author during four collection

trips between 1989 and 2003 suggest that there is no over-

lap of the distribution areas of S. bertholdi and S. wieseae

(Fig. 1). Scriptaphyosemion bertholdi inhabits southeast-

ern Sierra Leone and is found in river basins which enter

Bonn zoological Bulletin 61 (1): 13-28

the Atlantic ocean through the southern costal area (Hu-

ber 2007; Wildekamp 1993; Wildekamp & Van der Zee

2003), whereas S. wieseae inhabits the upper tributaries

of the Seli or Rokel River, which enter the Atlantic ocean

more to the north (Fig. 1). The geographically closest

Scriptaphyosemion species with regard to the known col-

lection localities of S. wieseae is S. chaytori (Fig. 1), to

the north in Guinea it is S. guignardi (Busch 1995a;

Wildekamp & Van der Zee 2003; own unpublished col-

lection data)

Etymology. The new species is named after Barbara

Wiese, who accompanied the second author on most of

his research trips in West Africa and was the first who rec-

ognized it as a new Scriptaphyosemion species.

DISCUSSION

Scriptaphyosemion wieseae is distinguished from the

closely related S. bertholdi (Figs 2—3) by several diagnos-

tic characters, and is probably separated by strong genet-

ic incompatibilities (Vandersmissen 2003). As proposed

by Roloff (1971a), female color characters can be useful

for distinguishing between some Scriptaphyosemion

species, here by the diagnostic lateral color pattern differ-

ences between S. wieseae (Fig. 4B) and S. bertholdi (Fig.

5B). However, female pigmentation pattern can be ob-

scured by several factors, as e.g. stress, and vary in inten-

sity (Roloff 1971a; own observations).

The combination of high variability in male coloration

between isolated populations of one species and the of-

ten only small diagnostic color pattern differences between

species can make species identification in Script-

aphyosemion difficult (Roloff 1971a; Scheel 1990).

The recent introduction of molecular methods in taxo-

nomic and phylogenetic studies of nothobranchiid killi-

fishes added an additional and independent set of charac-

ters and helped to resolve complicated cases of species lev-

el taxonomy (e.g. Legros & Zentz 2007b; Legros et al.

2005; Sonnenberg 2007, 2008). This approach can also be

applied to resolve the taxonomy of Scriptaphyosemion by

clarifying the relationships between species and pinpoint-

ing diagnostic characters in terms of morphology and col-

or pattern.

Our present results are largely congruent with those of

Murphy et al. (1999), if only the well supported nodes are

considered. Differences e.g. in the placement of S. rolof-

fiare based on different opinions on the species level tax-

onomy. Incongruences in the placement of some species

and low support of several nodes might be based on in-

sufficient information content in the dataset.

The perceived variability of male coloration in the ab-

sence of other diagnostic characters and different species

concepts led to varied opinions concerning species valid-

OZFMK

24 Rainer Sonnenberg & Eckhard Busch

ity and number between authors (e.g. Costa 2008; Huber

2007; Lazara 2001; Scheel 1968, 1990; Wildekamp 1993).

Even decades after Roloff (1971la) and Scheel’s (1990)

comments on the taxonomic difficulties, the species lev-

el taxonomy of Scriptaphyosemion is still far from being

resolved. Here we provide an overview of species names

in Scriptaphyosemion and their synonyms, with short com-

ments on species whose taxonomy needs a revision. We

mainly follow the species level taxonomy as proposed by

Costa (2008); the species concept employed is that of

Moritz et al. (2000).

The following species, S. bertholdi, S. brueningi, S.

chaytori, S. fredrodi, and S. schmitti have comparatively

smaller distribution areas, and usually show less variabil-

ity in the diagnostic characters than the species listed be-

low. They are, with the exception of S. fredrodi, concor-

dantly considered as valid species by most authors (Cos-

ta 2008; Huber 2007; Lazara 2001; Wildekamp & Van der

Zee 2003). In addition, the studied samples of these

species show considerable genetic distance with regard to

their closest relatives within the molecular phylogenies

(Figs 2-3, Murphy et al. 1999), supporting their status as

separate species.

Scriptaphyosemion geryi is distributed from northern

Sierra Leone to the Senegal (Wildekamp 1993;

Wildekamp & Van der Zee 2003). Within this large area,

differences in coloration pattern between populations are

known (e.g. Cauvet 2008; Etzel & Berkenkamp 1979;

Wildekamp 1993), indicating that probably more than one

species 1s currently included under this name.

Scriptaphyosemion guignardi was the first of three for-

mally described species of a species complex (Romand

1981, 1982; Seegers 1982). According to the DNA study

of Murphy et al. (1999), these three species form a mono-

phyletic group. Costa (2008) listed them as valid species,

but Etzel & Berkenkamp (1989a, b) and Etzel & Vander-

smissen (1984), based on crossing experiments, regard S.

nigrifluvi and S. banforense as junior synonyms of S. guig-

nardi. A short comparison of the published cytochrome b

sequences of Murphy et al. (1999) indicate considerable

genetic divergence between the three studied specimens

(4.9-10.8 %) which is in the range of intraspecific diver-

gence values in killifishes (Murphy et al. 1999; Sonnen-

berg 2007, 2008; Sonnenberg & Blum 2005; Van der Zee

& Sonnenberg 2011; own unpublished data). In addition,

these species share some coloration characters, which in-

dicates their close phylogenetic relationship. Still they are

diagnosable by male coloration characters. Studies em-

ploying crossing experiments (e.g. Etzel & Berkenkamp

1980a, b, c, 1989a, b; Etzel & Vandersmissen 1984; Ro-

mand 1986; Scheel 1968, 1990) show, that between many

populations different levels of genetic incompatibility ex-

ist. However, the conclusion that successful crossings up

to the third generation indicate that they are conspecific

is viewed with strong hesitation. The difficulties of the

Bonn zoological Bulletin 61 (1): 13-28

biospecies concept of Mayr (1963) is discussed in litera-

ture (e.g. Otte & Endler 1989), and recent examples of hy-

brid species and mitochondrial introgressions between

species prove the possibility of fertile hybrids between dis-

tinct species (e.g. Nolte et al. 2005; Schliewen & Klee

2004; Schwarzer et al. 2011; Sonnenberg 2007). At the

moment we follow Costa (2008) in regarding them as three

separate species.

The description of Scriptaphyosemion liberiense 1s

based on preserved females (Boulenger 1908). It has a

large distribution in coastal Liberia (Wildekamp & Van der

Zee 2003). Aphyosemion calabaricus Ahl, 1936, described

on specimens from an ornamental fish import without se-

cure collection locality, and Epiplatys melantereon

Fowler, 1950 were considered as junior synonyms to S.

liberiense (Costa 2008; Eschmeyer 2011; Huber 2007).

However, S. liberiense in its current usage is rather vari-

able in male phenotype between different populations (Et-

zel 1992, 1993; Milkuhn 2007, 2008; Wildekamp 1993).

Roloff (1971b) proposed that S. calabaricum females dif-

fer in coloration pattern from S. liberiense as described

by Boulenger (1908) and might represent a different

species. A careful study of S. /iberiense over its distribu-

tion range will be necessary to search for potentially sep-

arating stable diagnostic characters between groups of

populations and to decide about the status of S$. calabar-

icum. At the moment we tentatively consider this species

as a synonym of S. liberiense.

Scriptaphyosemion roloffi was accidentally described by

Roloff (1936), the intended description by Ahl (1937) fol-

lowed a year later. Both authors gave a detailed color pat-

tern description of the new species, drawings or black and

white pictures, the latter ones from E. Roloff or W. Har-

vey, were published by different authors (e.g. Henzelmann

1937; Mayer 1937; Ott 1997). By later authors (e.g. Et-

zel 1978c; Scheel 1968, 1990; Wildekamp 1993) the iden-

tity of S. roloffi was confused due to the inclusion of dif-

ferent populations, which do not show the diagnostic char-

acters of the specimens described by Roloff (1936) and

Ahl (1937) and probably represent different species. Et-

zel & Berkenkamp (1980a, b, c) presented the results of

crossing experiments between different populations,

which were assumed to be S. roloffi, but most probably

included at least one currently undescribed species, judged

from the published drawings. In 1979, S. etzeli was de-

scribed by Berkenkamp, based on specimens which show

the same coloration characters as given in the description

of S. roloffi (Ahl 1937; Henzelmann 1937; Mayer 1937;

Roloff 1936). The synonymy of S. etze/i with S. roloffi in

the sense of Roloff (1936) and Ahl (1937) was discussed

by Busch (1995b, 1996) and Ott (1997). We are of the

opinion that S. efzeli is a junior synonym of S. roloffi,

based on the diagnostic coloration characters of males and

females, and that S. ro/offi in the current usage

(Berkenkamp 1979; Etzel & Berkenkamp 1980a, b, c; Et-

OZFMK

New Scriptaphyosemion from Sierra Leone

zel 1978a, c; Etzel et al. 2004; Huber 2007; Wildekamp

1993) probably contains at least one undescribed species.

The incongruence of the placement of S. ro/offi between

our study (Figs 2-3) and Murphy et al. (1999) is caused

by different taxonomic concepts. Their S. etze/i is in our

view S. rolofji and the S. roloffi sample in their study most

probably is an undescribed species. The voucher of Script-

aphyosemion sp. in our DNA analyses (Figs 2—3) belongs

to the group of populations currently considered by oth-

er authors as conspecific with S. roloffi. However, it does

not show the diagnostic coloration characters of this

species and belongs to a different genetic lineage, closer

to S. fredrodi, and might probably be an undescribed

species related to that labeled as S. ro/offi in Murphy et

al. (1999). Scriptaphyosemion hastingsi (Wright, 1972),

also was an accidental description, but should be restud-

ied by new collections with regard to the species complex

currently included in S. ro/offi, to which it is currently con-

sidered as a synonym (e.g. Eschmeier 2011; Huber 2007;

Wildekamp 1993). We tentatively consider it as a syn-

onym, but it might turn out to be a valid name for a species

currently included in S. roloffi.

For the two species names, Roloffia caldal Cahalan,

1973 and Aphyosemion muelleri Scheel, 1968, we follow

other authors (Eschmeyer 2011; Huber 2007; Wildekamp

1993), who consider them as invalid or not available, as

they were not mentioned by Costa (2008).

COMPARATIVE MATERIAL

Scriptaphyosemion bertholdi (Roloff, 1965): ZFMK

47708-47714, Sierra Leone, Victoria, collection locality SL

O3 Silk

Scriptaphyosemion brueningi (Roloff, 1971b): ZFMK

47715-47725, Sierra Leone, Perie, collection locality SL 89

Perie; ZFMK 47726-47735, Sierra Leone, Tawai, collection lo-

cality SL 03/2.

Scriptaphyosemion calabaricum (Ahl, 1936): ZMB 21039, Lec-

totype, Nigeria (probably from Liberia); ZMB 31494, Paralec-

totype, collected with the Lectotype.

Scriptaphyosemion cauveti (Romand & Ozouf-Costaz, 1995):

ZFMK 47736-47739, Guinea, Kamara Bounyi, collection local-

ity GM 97/5.

Scriptaphyosemion chaytori (Roloff, 1971la): ZFMK

47740-47755, Sierra Leone, Kamabai, collection locality SL

93/34; ZFMK 47756-47760, Sierra Leone, Nom-Mussiray, col-

lection locality SL 93/38.

Scriptaphyosemion fredrodi (Vandersmissen, Etzel &

Berkenkamp, 1980): ZFMK 47761-47775, Sierra Leone,

Matanga, collection locality SL 89 Matanga.

Scriptaphyosemion geryi (Lambert, 1958): ZFMK 47776-47780,

Sierra Leone, Rotain, collection locality SL 89 Rotain.

Scriptaphyosemion guignardi (Romand,1981): ZFMK

47781-47782, Guinea, Gubi, collection locality GM 97/13.

Scriptaphyosemion liberiense (Boulenger, 1908): BMNH

1903.6.3.17-18, syntypes, Liberia, Monrovia; ZFMK

47783-47785, Liberia, Paynesville, collection locality L97;

ZFMK 12163, Liberia, surrounding of Monrovia, commercial

import; ZFMK 12659-662, Liberia, Totota.

Bonn zoological Bulletin 61 (1): 13-28

Scriptaphyosemion melantereon (Fowler, 1950): ANSP 72125,

Holotype, Liberia, Robertsport; ANSP 72126—27, Paratypes, col-

lected with holotype.

Scriptaphyosemion roloffi (Roloff, 1936): ZMB 21071, Lecto-

type, Sierra Leone, tribuary of Port Loko Creek, ca. 40 km east-

northeast of Freetown, ZMB 31472, Paralectotypes, collected

with Lectotype; ZFMK 47786-47798, Sierra Leone, Malai, col-

lection locality SL 03/11; ZFMK 47799-47813, Sierra Leone,

Lungi Lol, collection locality SL 99/10; ZFMK 12678-80 (as S.

etzeli), Sierra Leone, Port Loko district, Kirma.

Scriptaphyosemion schmitti (Romand, 1979): ZFMK

47814-47815, Liberia, Juarzon, aquarium bred specimens.

Scriptaphyosemion sp. aff. roloffi: ZFMK 47816-47819, Sier-

ra Leone, Devil Hole, collection locality SL 03/18; ZFMK

47820-47825, Sierra Leone, Kundubu, collection locality SL

93/6; ZFMK 12656-58, Sierra Leone, Magbenta; ZFMK 14808-

11, Sierra Leone, 15 km north Bo.

Acknowledgements. EB thanks Mohamed Basiru and family

(Brama Village, Sierra Leone) and the people of Brama Village

for hospitality, organization and help during field work at four

research trips in Sierra Leone, Wilhelm Kugelmann

(Dinkelscherben, Germany) for his support in keeping and breed-

ing many species collected on several journeys, and Barbara

Wiese (Remscheid, Germany) for her support on many research

trips in West Africa since years. RS thanks Peter Bartsch and

Christa Lamour (MEN, Berlin, Germany), and Oliver Crimmen

and James Maclaine (NHM, London, Great Britain) for kind ac-

cess to the collection under their care, Klaus Busse and Fabian

Herder (ZFMK, Bonn, Germany) for access to the collection and

the X-ray device, Kyle Luckenbill, John Lundberg, and Mark

Sabaj Perez (ANSP, Philadelphia, USA) for data, X-rays, and

photos of type specimens under their care, Diethard Tautz (MPI,

Plén, Germany) for support with lab space and access to the mi-

cro Computer Tomograph, Heinz Otto Berkenkamp (Wil-

helmshaven, Germany), Gerhard Schreiber (Liidenscheid, Ger-

many), Thomas Litz (Attenweiler, Germany) for support with

Literature. We thank Giinther Fleck (Saint-Gaudens, France) for

the translation of the Abstract, Anja C. Schunke (MPI, Pl6n, Ger-

many) for comments and corrections on an earlier version of the

manuscript, and Fabian Herder and two anonymous reviewers

for valuable comments.

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Bonn zoological Bulletin 61 (1): 29-30

July 2012

Mercediella nom. nov., a replacement name for

Camposichthys Figueiredo & Silva Santos, 1991

(Pisces: Pycnodontiformes)

Stefan Koerber

Friesenstr. 11, D-45476 Muelheim, Germany; E-mail: pecescriollos@koerber-germany.de; www.pecescriollos.de.

Abstract. Camposichthys has been published three times for genera of bony fishes, generating a triple homonymy which

is described and analyzed in this paper. A replacement name for the youngest homonym is presented.

Camposichthys Travassos, 1946

(Characiformes: Cynodontidae)

Travassos (1946) was the first to errect Camposichthys and

did so on some erroneous assumptions regarding the then

complicated status of Cynodon Spix in Spix & Agassiz,

1829, Raphiodon Agassiz in Spix & Agassiz,1829, and

Rhaphiodonichthys Campos, 1945 (see Kottelat 1988). In

Opinion 2012, the International Commission on Zoolog-

ical Nomenclature (2002) considered Camposichthys

Travassos, 1946, to be a junior objective synonym of Cyn-

odon Spix in Spix & Agassiz, 1829, and placed it in the

‘Official Index of Rejected and Invalid Generic Names in

Zoology’. Despite this nomenclatural act, in accordance

with § 80.7.1 of the current version of the Code (ICZN

1999) this name continues to compete in homonymy.

Camposichthys Whitley, 1953

(Characiformes: Curimatidae)

In 1953, Whitley established Camposichthys as a replace-

ment name for Camposella Fernandez-Yepez, 1948 pre-

occupied by Camposella Cole, 1919 (Insecta: Diptera)

and, when he learned that Camposichthys Whitley, 1953

was also preoccupied, in 1954 he replaced it with a new

name, Stupens.

Camposichthys Figueiredo & Silva Santos, 1991

(Pycnodontiformes: Pycnodontidae)

Camposichthys was used a third time as the name of a new

genus of Pisces by Figueiredo & Silva Santos (1991), for

tCamposichthys riachuelensis, a fossil pycnodontid

species from Brazil described as new in the same paper.

This genus is currently treated as valid (Barbosa et al.

2008; Figueiredo 2009; Poyota-Ariza & Wenz 2002;

Souza Lima et al. 2002) and no junior synonym is avail-

able for this genus that could serve as a replacement name.

Received: 03.01.2012

Accepted: 02.02.2012

To resolve the remaining homonymy between Cam-

posichthys Travassos, 1946, and Camposichthys Figueire-

do & Silva Santos, 1991, Mercediella is here proposed as

a replacement name for the latter junior homonym follow-

ing §§ 53.2, 56.1 and 60.3 of the Code. The only known

member of this genus is the originally designated type

species +Mercediella riachuelensis (Figueiredo & Silva

Santos, 1991).

The new name honours Mercedes Azpelicueta, ichthyol-

ogist and paleoichthyologist from La Plata, Argentina.

Thanks are due to Miguel Alonso-Zarazaga, Madrid, and

Alain Dubois, Paris, for sharing their experience and ex-

pertise regarding the rules of the Code. An earlier version

of this paper benefited from comments of Mark Wilson,

Edmonton, and an anonymous reviewer.

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©ZFMK

Bonn zoological Bulletin 61 (1): 31-34

July 2012

Report on the life colouration of the enigmatic burrowing skink

Voeltzkowia rubrocaudata (Grandidier, 1869)

from southwestern Madagascar

Goncalo M. Rosa’, Paolo Eusebio Bergo , Angelica Crottini- & Franco Andreone

' Durrell Institute of Conservation and Ecology, School of Anthropology and Conservation, University of Kent, Canterbury,

E Kent, CT2 7NR, UK

Centro de Biologia Ambiental, Faculdade de Ciéncias da Universidade de Lisboa, Bloco C2. Campo Grande,

\ P-1749-016 Lisboa, Portugal

* Museo Regionale di Scienze Naturali, Sezione di Zoologia, Via G. Giolitti, 36, I-10123, Torino, Italy

“CIBIO, Centro de Investigacao em Biodiversidade e Recursos Genéticos, Campus Agrario de Vairao, R. Padre Armando Quin-

tas, P-4485-661 Vairao, Portugal

* corresponding author: Gongalo M. Rosa: goncalo.m.rosa@gmail.com

Abstract. Voeltzkowia is a monophyletic genus of burrowing skinks endemic to Madagascar. The fossorial habits of these

species make them hard to see and study, and witness their life history traits. During two herpetological surveys in south-

western Madagascar (in 2009 and 2011) we found several Voeltzkowia rubrocaudata individuals in a corn plantation, a

habitat that differs from the forested habitat reported in the literature. Life colouration for this shy scincid is described

for the first time.

Key words. Southwestern Madagascar, corn plantation, Squamata, Scincidae.

Voeltzkowia Boettger, 1893 is a poorly known mono-

phyletic genus of scincid lizards endemic of Madagascar

(Whiting et al. 2004, Schmitz et al. 2005, Crottini et al.

2009). Species of this genus belong to a phylogenetic clade

that contains species of Amphiglossus sensu strictu, and

the other limb-reduced Pygomeles and Androngo. With a

body elongation and limb reduction, these shy and enig-

matic skinks show burrowing habits (Brygoo 1981, Glaw

& Vences 2007). The genus is currently composed of three

completely limbless species (V. lineata, V. mira and V.

rubrocaudata), and two species with very rudimentary

hindlimbs (V. petiti and V. fierinensis), but a careful genus

revision is currently in act and may increase the number

of species of this enigmatic group (A. Miralles, pers.

comm.). Despite exhaustive field surveys, fossorial

skinks are usually hard to find and some species were not

being observed on the island for more than a century (Bry-

goo 1981, Kohler et al. 2010).

According to the original description, a reddish tail char-

acterizes Voeltzkowia rubrocaudata. This species is

known from many locations in the arid regions of south-

western and western Madagascar (Grandidier 1869,

Glaw & Vences 2007). In this manuscript we report for

the first time on the life colouration of live individuals of

the species Voeltzkowia rubrocaudata, providing also an

update on its known distribution and details on habitat

preferences.

Received: 18.01.2012

Accepted: 05.04.2012

The individuals reported were found during two her-

petological surveys carried out in southwestern Madagas-

car in late 2009 and early 2011. According to available

data summarised by Glaw & Vences (2007), V. rubrocau-

data is known from Ambohimahalevona, Ampoza, Befan-

driana, Beroboka Avaratra, Fierin, Isalo, Toliana and Zom-

bitse Forest (currently integrated in the complex Zombitse-

Vohibasia National Park). Despite recent herpetological

surveys in Zombitse, our team could not confirm its pres-

ence. Additionally, intensive research at Isalo National

Park (e.g., Mercurio et al. 2008) provided similar results

in that area: nevertheless, we observed traces on sand

along dry rivers that probably refer to a burrowing skink

species.

When pictures of other Voeltzkowia species were

shown to local people in Andranomaitso (a small village

on the Route Nationale 7, Commune rurale de Sakaraha),

they informed us that similar animals would be found in

the corn plantation near the village (S22° 53’ 52.2”— E44°

39’ 23.1”, 710 m a.s.1.; fig. 1). An intensive search, per-

formed by digging the earthy dry substrate close to the

stems and roots, and flipping the rocks of the area, led us

to find one individual [MRSN R3726; (corresponding to

the tissue sample ACZC2565 where ACZC refers to field

numbers of A. Crottini); GenBank accession number

JQO005113] on the 11 December 2009 (fig. 2). Such indi-

vidual (probably a male, SVL 52 mm, with 115 ventral

Corresponding editor: P. Wagner

eS)

va P i sf} f

Gongalo M. Rosa et al.

Fig. 1. Habitat of Voeltzkowia rubrocaudata: corn plantation (in foreground) near the village of Andranomaitso, Commune rurale

de Sakaraha.

Table 1. Morphological measurements (all in mm) of specimens

of Voeltzkowia rubrocaudata from southwestern Madagascar.

Used abbreviations are: MRSN (Museo Regionale di Scienze

Naturali, Torino), FAZC (Franco Andreone Zoological Collec-

tion, Field numbers), SVL (snout—vent length), TL (total

lenght), VS (ventral scales), DS (dorsal scales), MBS (midbody

scales).

MRSN FAZC SVL TL VS DS MBS

R3726 14370 52 82.5 WS 113 18

R3738 14776 90 1275 120 119 19

R3742 14777 63.5 100 116 120 19

R3743 14778 = 58 90.5 116 118 19

R3744 14775 78 127 117 119 19

“tip of the tail removed as DNA tissue sample.

scales, 113 dorsals, 18 around midbody) was burrowed

<10 cm deep near a corn root. Nine other individuals were

found during a second visit that took place in the same

area on the 14 February 2011. Of these, four are now host-

ed in the zoological collection of the Museo Regionale di

Scienze Naturali (Turin, Italy) (MRSN_ R3738,

R3742-3744), and five are hosted in the zoological col-

lection of the Parc Botanique et Zoologique de Tzimbaza-

Bonn zoological Bulletin 61 (1): 31-34

za (Antananarivo, Madagascar) (Franco Andreone field

numbers FAZC 14779-14783). Morphological measure-

ments are presented in Table | and overall fit the descrip-

tions provided by Mocquard (1894) and Brygoo (1981).

All these individuals are attributed to V. rubrocaudata

due to the absence of hindlimbs, high number of ventral

scales, and nostril in contact with first suprabial, accord-

ing to the indications by Brygoo (1981). Notwithstanding,

one major diagnostic character, the red tail (from which

the specific epithet) was missing from all the examined

live individuals. Chromatic differences among the indi-

viduals do not seem to be accentuated. The overall live

colouration was light beige with pinkish and yellowish nu-

ances on the back, and dark arrays of small dark points

that form longitudinal stripes on the caudal portion. In par-

ticular, two individuals currently hosted in MRSN, and

featured by a rather small body size (MRSN R3742-3743)

have the dark lines on the tail quite evident and contrast-

ed. The ventral side of the individuals was a slightly

lighter. The five specimens housed in MRSN after about

three years (MRSN R3726) and one year (MRSN R3738,

R3742-3744) of preservation in ethanol (70%) show a

general fade from beige into an almost whitish coloration,

with persistence of the dark point and lines. In such a con-

text it appeared quite unusual the absence of the red

colouration reported by Grandidier (1869) and stressed by

Brygoo (1981).

©ZFMK

Life colouration of Voeltzkowia rubrocaudata 33

fec5 ae

a Se ee oe e

Mee ee om ae

as rs Salt “ < rs

Fig. 2. Voeltzkowia rubrocaudata individual (MRSN R3726) in life from Andranomaitso village, southwestern Madagascar, found

on the 11 December 2009: A. dorso-lateral overview; B. dorsal view; C. head close up; D. individual burrowing in the substrate.

Photos by Gongalo M. Rosa.

Further studies are necessary to understand the popu-

lation variability of the species, and verify the persistence

of the red colouration in the tails of specimens coming

from the type locality (Fierin). Moreover, the habitats and

ecology of this species are still poorly known, but this bur-

rowing skink seems to be tolerant to anthropogenically-

modified areas, benefiting from shaded habitat created by

the platation, as well as soil tillage that makes digging eas-

ier.

Bonn zoological Bulletin 61 (1): 31-34

Acknowledgments. We are grateful to our guide Anicet and all

the kind people from Andranomaitso village. The work was car-

ried out in collaboration with Madagascar National Parks (MNP).

We are grateful to the Malagasy authorities, in particular the Min-

istére de |’ Environnement et des Eaux et Foréts, for research per-

mits. We thank A. Miralles for confirming the species identifi-

cation and for sharing unpublished data. Fieldwork was partial-

ly funded by Mohamed bin Zayed Species Conservation Fund

and EDGE, Zoological Society of London. The work of AC was

supported by a_ postdoctoral grant from FCT

(SFRH/BPD/72908/2010).

©ZFMK

34 Gongalo M. Rosa et al.

REFERENCES

Brygoo ER (1981) Systématique des lézards scincidés de la ré-

gion malgache. VII. Révision des genres Voeltzkowia Boett-

ger, 1893, Grandidierina Mocquard, 1894, et Cryptoscincus

Mocquard, 1894 [sic!]. Bulletin du Muséum National d’ His-

toire Naturelle, Paris 4me Série, 3, Section A 2: 675—688

Crottini A, Dordel J, Kohler J, Glaw F, Schmitz A, Vences M

(2009) A multilocus phylogeny of Malagasy scincid lizards

elucidates the relationships of the fossorial genera Androngo

and Cryptoscincus. Molecular Phylogenetics and Evolution

53: 345-350

Glaw F, Vences M (2007) A field guide to the amphibians and

reptiles of Madagascar. Third edition, Cologne, Vences &

Glaw Verlag, 496 pp

Grandidier MA (1869) Descriptions de quelques animaux nou-

veaux découverts, pendant l’année 1869, sur la cote ouest de

Madagascar. Revue et Magazine de Zoologie (Paris) 2(21):

337-342

Kohler J, Vences M, Erbacher M, Glaw F (2010) Systematics

of limbless scincid lizards from northern Madagascar: mor-

phology, phylogenetic relationships and implications for clas-

sification (Squamata: Scincidae). Organisms, Diversity & Evo-

lution 10: 147-159

Bonn zoological Bulletin 61 (1): 31-34

Mercurio V, Aprea G, Crottini A, Mattioli F, Randrianirina JE,

Razafindrabe TJ, Andreone F (2008) The amphibians of Isa-

lo Massif, southern-central Madagascar: high frog diversity

in an apparently hostile dry habitat. Pp. 143-196 in: Andreone

F (eds.). A Conservation Strategy for the Amphibians of Mada-

gascar. Monografie XLV. Museo Regionale di Scienze Natu-

rali, Torino

Mocquard F (1894) Reptiles nouveaux ou insuffisamment

connus de Madagascar. Compte-Rendu Sommaire des Séances

de la Société Philomathique de Paris 17: 1-8

Schmitz A, Brandley MC, Mausfeld P, Vences M, Glaw F, Nuss-

baum RA, Reeder TW (2005) Opening the black box: phylo-

genetics and morphological evolution of the Malagasy fosso-

rial lizards of the subfamily “Scincinae”. Molecular Phyloge-

netics and Evolution 34: 118-133

Whiting AS, Sites Jr, Bauer AM (2004) Molecular phylogenet-

ics of Malagasy skinks (Squamata: Scincidae). African Jour-

nal of Herpetology 53(2): 135-146

©ZFMK

Bonn zoological Bulletin 61 (1): 35-40

July 2012

New data on the morphology and natural history of

Tetradactylus ellenbergeri (Angel, 1922) (Sauria: Gerrhosauridae) and

Trachylepis ivensii (Bocage, 1879) (Sauria: Scincidae)

in northeastern Zambia

Philipp Wagner” , Dennis Rédder & Thomas M. Wilms

. ‘ Department of Biology, Villanova University, Villanova, Pennsylvania 19085, USA.

_ Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

‘ Zoologischer Garten Frankfurt, Bernhard-Grzimek-Allee 1, D-60316 Frankfurt am Main, Germany

‘corresponding author: philipp.wagner.zfmk@uni-bonn.de.

Abstract. We report new observations regarding the morphology, behaviour and habitat of the two sparsely known lizards

Tetradactylus ellenbergeri (Gerrhosauridae) and Trachylepis ivensii (Scincidae) from northeastern Zambia and review

the available data about the distribution of both species.

Key words. Sauria, Zrachylepis ivensii, Tetradactylus ellenbergeri, habitat, Africa, Ikelenge, Zambia.

In 2008, a field study was conducted in the Ikelenge area

(northern Mwinilunga District) of Zambia to explore the

unique diversity of amphibians and reptiles in this nation-

al biodiversity hotspot. Ikelenge is situated in the extreme

north-western edge of Zambia within a pedicle wedged

between Angola and the Democratic Republic of the Con-

go (DR Congo). Three other studies on the herpetofauna

were conducted in this area in the past. Donald Broadley

(1991) was the first who summarized a checklist of rep-

tiles and amphibians of the northern Mwinilunga district.

He recorded 57 reptiles and 35 amphibian species, includ-

ing Tetradactylus ellenbergeri Angel, 1922 for the first

time in the area. Later, in 1994, Alan Channing, Robert

Drewes and Jens Vindum also did a field survey which

was mainly concentrated on amphibians. The results of

this study were never published but aspects are mentioned

in a book on southern African amphibians (Channing

2001). Also Haagner et al. (2000) collected a series of both

amphibians and reptiles from this area, including some

records of T. ellenbergeri, and the discovery of Trachylepis

ivensii Bocage, 1879 for the first time for Zambia and the

DR Congo.

The herein described field survey was conducted be-

tween July and September 2008 to collect amphibians and

reptiles. In this area the rainy season is nearly over in

March and therefore this study was done, like the Haag-

ner study (W.B. Branch, pers. comm.), in the colder dry

season just after the rainy but before the hot dry season.

During the study time seven specimens of Tetradactylus

ellenbergeri and six specimens of Trachylepis ivensii were

collected (see below). The vouchers are stored and cata-

logued in the herpetological collections of the Zoologi-

Received: 10.10.2011

Accepted: 03.04.2012

sches Forschungsmuseum Alexander Koenig (ZFMK) in

Bonn, Germany and the Muséum d’histoire naturelle

(MHNG), Geneva, Switzerland.

Notes on Tetradactylus ellenbergeri (Angel, 1922)

Tetradactylus ellenbergeri is a snake-like plated lizard

from southern central and eastern Africa and the sole

member of the genus in Zambia. The genus is character-

ized by extremely reduced limbs. In 7: ellenbergeri the

front limbs are lacking and the hind limbs are reduced to

about 2 mm. The tail is long, more than three times longer

than the body. Individuals are slightly bluish above, with

two median vertebral rows of brown scales. The tempo-

ral region is spotted brownish; the underside is pale olive

(see fig. la).

Vouchers. ZFMK 88526-529, ZFMK 89188, ZFMK

89421, ZFMK 92525. Most of the voucher specimens

were found within the Nchila Reserve, Hillwood Farm,

near Ikelenge. One specimen (ZFMK 88529) was found

by local collectors in the surrounding area. Five of the sev-

en preserved specimens are adults and have a snout-vent

length (SVL) between 59.7 and 68.5 mm, with an aver-

age of 64.0 mm (n= 5). The single subadult (ZFMK

88529) measures 41.4 in SVL. The sole juvenile (ZFMK

92525) measures 30.6 mm and has an original tail with a

length (TL) of 93.8 mm. In adults, the TL measured in av-

erage 186.2 mm (114.2—250.0 mm; n=5) but in four spec-

imens the tip is cut or the tail is regenerated. The only

specimen (ZFMK 88528) with an original tail measured

Corresponding editor: F. Herder

Philipp Wagner et al.

Fig. 1. Tetradactylus ellenbergeri. A= Specimen from Nchila Reserve, Hillwood Farm, Ikelenge, Zambia. B= X-ray image of a

pregnant female containing two eggs of 7. ellenbergeri. C= Habitat of 7. ellenbergeri in the Nchila Reserve.

Bonn zoological Bulletin 61 (1): 35—40 OZFMK

New data of Jetradactylus ellenbergeri and Trachylepis ivensii in northeastern Zambia 37

10°

20°

“Tanzania, —

Fig. 2. Distribution of Tetradactylus ellenbergeri and Trachylepis ivensii. T. ellenbergeri. 1= Tanzania: Gendawaki Valley, Udzung-

wa Mountains (Menegon et al. 2006). 2= Tanzania: Tatanda (Spawls et al. 2002). 3= Tanzania: Songea (Spawls et al. 2002). 4=

DR Congo: Kundelungu (Broadley 1971). 5= DR Congo: Kansenia (Broadley 1971). 6= Zambia: Chongwe River (Broadley 1971).

7= Zambia: Lunga Game Reserve (Broadley 1971). 8= Zambia: Ikelenge area. 9= Angola: Lunda (Broadley 1971). 7. ivensii. 10=

DR Congo: Sanolumba Village (Branch & Haagner 1993). 11= Zambia: Ikelenge area, including source of the Zambezi and north

of Ikelenge hospital (Branch & Haagner 1993). 12= Angola: Luena (Branch & Haagner 1993). 13= Angola: Dala, Lunda (Branch

& Haagner 1993). 14= Angola: Alto Cuilo (Branch and Haagner 1993). 15= Angola: Alto Chicapa (Branch & Haagner 1993). 16=

Angola: Luando River (Branch & Haagner 1993). 17= Angola: Curanza River (Branch & Haagner 1993). 18= Angola: Cuando

River (Branch & Haagner 1993). 19= Angola: Huambo (Branch & Haagner 1993).

SVL: 60.5 mm and TL: 231.0 mm. This results in a

TL/SVL ratio of 3.81, and therefore the tail is about four

times longer than the body. Broadley (1991) mentioned a

subadult specimen (National Museums of Zimbabwe,

NMZB 10663) with a SVL of 56 mm and a TL of 200 mm,

a ratio of 3.57.

Distribution. In Tanzania, the species is only known from

three localities: Tatanda and Songea are mentioned by

Spawls et al. (2002), whereas Menegon et al. (2006) found

the species also at Gendawaki Valley in the Udzungwa

Mountains which is the easternmost record. In Zambia it

is known from three localities (Chongwe River, Lunga Ga-

me Reserve, Ikelenge area), and even in DR Congo and

Angola it is only known from few localities (see fig. 2).

Habitat. Spawls et al. (2002) noted that the Tanzanian

specimens live in moist savanna. All specimens from the

Nchila Reserve were found in a similar habitat (fig. 1c),

a swampy grassland, interspersed with grass tussocks. A

typical such habitat is about six metres broad and 300 me-

tres in length, bordering the gallery forest of the Sakeji

Bonn zoological Bulletin 61 (1): 35—40

River. Specimens of 7 e/lenbergeri were captured during

daytime, basking on the top of grass tussocks. If disturbed,

they tried to escape, diving into the water between the tus-

socks. The muddy water is relatively deep, between 10 and

30 cm. Sphagnum moss (Bryophyta, Sphagnaceae) and

sundew plants (Magnoliophyta, Droseraceae) were found

between the tussocks in this habitat. Both plants indicate

a permanent water body. In the more open areas of the wa-

ter body, Sphagnum spec. is dense and overgrows the ex-

panse of water.

Natural history. Spawls et al. (2002) mentioned the species

as diurnal and probably oviparous. Haagner et al. (2000)

recognized ova and eggs in several specimens collected

in April, but one adult female (ZFMK 88526, field no. PW

HF 296) collected much later in the year, on August 18th,

2008 also contained two well developed eggs (see fig. 1b).

If grasped, these lizards readily shed their tails. How-

ever, tail regeneration is rapid, probably because the tail

is essential for locomotion. A captive specimen regener-

ated one centimetre of a freshly lost tail in four weeks.

©ZFMK

38 Philipp Wagner et al.

—_— i ~_

nig eM

WN Ny

FASS Aw

Fig. 3. Trachylepis ivensii. A= Specimen from Nchila Reserve, Hillwood Farm, Ikelenge, Zambia. B= Typical habitat of 7. iven-

sti in the Nchila Reserve. C= Typical habitat of 7 ivensii at the Zambezi north of Ikelenge.

Bonn zoological Bulletin 61 (1): 35-40 ©ZFMK

New data of Tetradactylus ellenbergeri and Trachylepis ivensii in northeastern Zambia 39

Notes on Trachylepis ivensii (Bocage, 1879)

Trachylepis ivensii is a relatively poorly known scincid

lizard from south-western central Africa and has so far on-

ly been recorded from a handful of vouchers worldwide.

Branch & Haagner (1993) reported 21 specimens collect-

ed by them and 17 are preserved at the Port Elizabeth Mu-

seum (PEM). Additionally, two specimens were donated

to the Lambiris private collection and two to the Natural

History Museum of Zimbabwe (NMZB). Nine specimens

are present in the collection of the American Museum of

National History (AMNH) and three at the California

Academy of Science (CAS). Also Boulenger (1887) men-

tioned one specimen present in the collection of the British

Museum (BMNH). Together with the six specimens col-

lected in this study, only 40 vouchers exist in museums

worldwide. The specimens mentioned by Bocage (1879)

including the name bearing type are lost (Branch & Haag-

ner 1993).

T. ivensii 1s a large, elongate species of the genus with

a greatly enlarged tail (see fig. 3a). It is characterized by

having two to three (usually three) ear lobules and 28-32

scales rows around midbody. Body scales are keeled and

heavily keeled over the middle of the back.

Belly, throat, underside of tail and limbs are patternless;

limbs are dark brown, with two fine darker lines on the

upper sides of the forelimbs and a white band separating

the upper and lower surfaces of the hindlimbs. Head and

body are covered with a stripe pattern. On the back and

the tail are three (one vertebral, two dorsolateral) pale

brown stripes, arising at the temporal region. A white and

dark framed stripe arises on both sides at the nasal region

and runs beneath the eye and through the ear along the

flanks and fades above the top of the hind limbs. A sec-

ond such stripe arises beneath the ear and runs along the

lateral part of the body, separating the belly from the

flanks, passing the hind limbs and extending on the tail

to the tip. The stripes are lacking on regenerated parts of

the tail. Further characteristics of the species are described

by Branch & Haagner (1993) and can be seen in figure

3a.

Vouchers. ZFMK 88547—551, MHNG 2713.34.

Distribution. This skink is only known from Angola, Zam-

bia and south-western DR Congo. Branch & Haagner

(1993) recorded it for the first time in Zambia, from the

northern Mwinilunga District and the adjacent DR Con-

go, which are the only known localities from these coun-

tries to date. In Zambia the species was only found along

to the Sakeji and Zambezi rivers at Ikelenge area; the DR

Congo record is from Salolumba village, 46 km north east

of Sakeji. In Angola, T. ivensii is known from several

rivers in the centre of the country (see fig. 2).

Bonn zoological Bulletin 61 (1): 35—40

Habitat. This species was always found near water bod-

ies or muddy areas, within open grass- or woodland. Spec-

imens were observed along the Sakeji and Zambezi Riv-

er, only a few metres away from the riverbank (see fig.

3c). If there was gallery forest, individuals were never ob-

served on the riverbank, but along the forest border in

muddy grassland (see fig. 3b). However, the species is not

just known from watercourses. Voucher specimens in the

collection of the California Academy of Sciences (CAS

196639641) came from fishponds on Hillwood Farm,

near Ikelenge. Branch & Haagner (1993) found their

vouchers basking on logs alongside the origin of the Zam-

bezi River and near Ikelenge hospital at the Zambezi

rapids. Additionally, they found individuals along the

Sakeji River at Sakeji School, near Ikelenge.

Natural history. Previous authors recognized the partial-

ly aquatic habits of this species. Laurent (1964) noted that

local people found the lizard in fish traps, whereas Man-

acas (1963) found the lizards in muddy areas alongside

riverbanks. Haagner et al. (2000) gave a detailed descrip-

tion of the behaviour and mentioned specimens basking

on vegetation on the banks of the Sakeji River. If disturbed

the lizards leapt into the water, swam underwater with lat-

eral undulations of body and tail and escaped into sub-

merged vegetation. We also observed this behaviour. One

specimen we tried to catch close to the riverbank jumped

off a small cliff into 1.5 m deep water and escaped. Sur-

prisingly, three individuals were found in holes that pre-

sumably were self-excavated by the skinks; one of these

was dug out by a local man. These burrows were about

20 to 30 cm in length and in a distance from river bank

of about one to two meters. However, despite the fact that

T. ivensii only lives near water sources and also swims,

dives and jumps into the water to escape capture, it is not

an aquatic species, and probably similar to the African se-

mi-aquatic species Varanus niloticus (Varanidae) and the

members of the scincid genus Cophoscincopus from West

Africa (Bohme et al. 2000).

Very recently, Blackburn & Flemming (2012) have

shown that reproducing females ovulate tiny eggs and lat-

er supply the nutrients for development by placental

means. Moreover, the recognized pattern of fetal mem-

brane development in 7: ivensii is unique among verte-

brates and the species represents a new extreme in pla-

cental specializations of reptiles (Blackburn & Flemming

ZO):

Acknowledgements. For assistance during the survey we thank

the Director of Research Victor Siamudala and the Head of Re-

search Wilbroad Chansa, both from the Zambian Wildlife Au-

thority (ZAWA) and Kennedy Chongoo and Humphrey

Simukoko, both University of Zambia (UNZA, School of Vet-

erinary Medicine). The Fisher family, Helen Finnie and Esther

Townsend from Hillwood Farm, Zambia helped a lot in collect-

ing lizards and snakes and especially 7. ivensii.

OZFMK

40 Philipp Wagner et al.

Special thanks to Steven Spawls (Norwich), William R.

Branch (Port Elizabeth) and Andreas Schmitz (Geneva) for their

critical review of an earlier draft of the manuscript.

The study was partly funded by the Deutsche Gesellschaft fiir

Herpetologie & Terrarienkunde (DGHT), the Alexander Koenig

Stiftung (AKS) and the Alexander Koenig Gesellschaft (AKG).

Many thanks for the funding which gave us the possibility to do

this research in Zambia.

REFERENCES

Angel F (1922) Sur un Lézard d’un genre nouveau de la famille

de Gerrhosauridae. Bulletin du Muséum National d’ Histoire

Naturelle (Paris) 28: 150-152

Blackburn DG, Flemming AF (2012) Invasive implantation and

intimate placental associations in a placentotrophic A frican li-

zard, Trachylepis ivensi (Scincidae). Journal of Morphology

273: 137-159

Bocage JVB du (1879) Reptiles et batraciens nouveaux d’ An-

gola. Journal of the Academy of Sciences of Lisbon 7: 97-99

Boulenger GA (1887) Catalogue of the lizards in the British Mu-

seum (Natural History).Vol. HI. Lacertidae. London, 1887:

I-XII + 1-118

Branch WR, Haagner GV (1993) The skink Mabuya ivensii: new

records from Zambia and Zaire, and the status of the sub-

species septemlineata Laurent 1964 and the genus Lubuya

Horton 1972. Amphibia-Reptilia 14: 105-115

Bauer AM, Jackman T (2008) Global diversity of lizards in fresh-

water (Reptilia: Lacertilia). In: Balian EV, Lévéque C, Segers

Bonn zoological Bulletin 61 (1): 35-40

H, Martens K, eds. Freshwater Animal Diversity Assessment.

Springer, Netherlands, pp. 581—586

Bohme W, Schmitz A, Ziegler T (2000) A review of the West

African skink genus Cophoscincopus Mertens (Reptilia: Scin-

cidae: Lygosominae): Resurrection of C. simulans (Vaillant,

1884) and description of a new species. Revue suisse de Zo-

ologie 107: 777-791

Broadley DG (1991) The herpetofauna of northern Mwinilun-

ga District, northwestern Zambia. Arnoldia Zimbabwe 9:

519-538

Channing A (2001) Amphibians of central and southern Africa

Cornell University Press, 470 pp.

Haagner GV, Branch WR, Haagner AJF (2000) Notes on a col-

lection of reptiles from Zambia and adjacent areas of the Dem-

ocratic Republic of the Congo. Annals of the Eastern Cape

Museums |: 1—25

Laurent RF (1964) Reptiles et batraciens de I’ Angola (troisiéme

note). Companhia de Diamantes de Angola (Diamang), Ser-

vicos Culturais, Museu do Dundo (Angola), No. 67, 165 pp,

Managas S (1963) Saurios de Angola. Junta Invest. Ultram. 2

ser. 43: 223-240

Menegon M, Salvidio S, Moyer D (2006) Reptiles and amphib-

ians from a montane grassland: Gendawaki Valley, Udzung-

wa Mountains, Tanzania. African Herp News 40: 8-14

Schmitz A, Brandley MC, Mausfeld P, Vences M, Glaw F, Nuss-

baum RA, Reeder TW (2005) Opening the black box: phylo-

genetics and morphological evolution of the Malagasy fosso-

rial lizards of the subfamily ‘Scincinae’. Molecular Phyloge-

netics and Evolution 34: 118-133

Spawls S, Howell K, Drewes RC, Ashe J (2002) A field guide

to the reptiles of East Africa. Academic Press, 543 pp.

©ZFMK

Bonn zoological Bulletin 61 (1): 41-48

July 2012

New data about some Alticini from Taiwan with descriptions

of two new species (Coleoptera: Chrysomelidae)

Konstantin Nadein'!* & Chi-Feng Lee?

! Department of general and applied entomology, Institute of Zoology, National Academy of Sciences of Ukraine, Bogdana

Khmelnitskogo st. 15, 01601 Kiev, Ukraine; E-mail: luperus@mail.ru

2 Applied Zoology Division, Taiwan Agricultural Research Institute, 189 Chung-Cheng Road, Taichung 41362, Wufeng, Taiwan;

E-mail: cflee@gate.sinica.edu.tw

«corresponding author

Abstract. The distributional and taxonomical data on alticine genera Zaiwanoliprus Komiya, 2006, Pseudoliprus Chuy6

& Kimoto, 1960, Lipromorpha Chujo & Kimoto, 1960, and Psylliodes Latreille, 1829 are presented. Psylliodes palleo-

la Motschulsky, 1866 and Psylliodes angusticollis Baly, 1874 are synonymized with Psylliodes viridana Motschulsky,

1858 based on the examination of the type material. The homonymy of Lipromorpha cyanea L. Medvedev, 2009 from

Laos and Lipromorpha cyanea Chen et Wang, 1980 from Yunnan is revealed. Three species Psy/liodes subrugosa Jaco-

by, 1885, Psylliodes chujoe Madar, 1960, and Psylliodes cantonensis Gruev, 1981 are recorded from Taiwan for the first

time. Two new species are described: Lipromorpha alutacea Nadein, sp. n. and Psylliodes yuae sp. n.

Key words. Alticini, flea beetles, Lipromorpha, Pseudoliprus, Taiwanoliprus, Psylliodes, synonyms, homonymy, new

species.

INTRODUCTION

The alticine genera Zaiwanoliprus Komiya, 2006,

Pseudoliprus Chajo & Kimoto, 1960, Lipromorpha

Chuayj6o & Kimoto, 1960, and Psylliodes Latreille, 1829 are

represented in Taiwan by 14 species (Kimoto & Takiza-

wa 1997; Komiya 2006; Doberl 2010). These are: Tai-

wanoliprus wenroni Komiya, 2006, 7. endonis Komiya,

2006, Pseudoliprus lalashanensis Komiya, 2006, P. kimo-

tonis Komiya, 2006, P. saigusai Kimoto, 1970, Lipromor-

pha montana Choyjo, 1935, L. shirozui Kimoto, 1970, L.

difficilis Chen, 1934, Psylliodes taiwana Takizawa, 1979,

P. taiwanica Chojo, 1935, P. chlorophana Erichson, 1842,

P. brettinghami Baly, 1862, P. angusticollis Baly, 1874,

and P. punctifrons Baly, 1874. The presence of L. difficilis

in the Taiwanese fauna requires confirmation (Medvedev

2009). The range of this species is wider in comparison

with others species of the genus. It is recorded from Viet-

nam, South China, Ryu-Kyu Isl., and Taiwan. In general,

the species of Lipromorpha have tendencies to local en-

demism; mostly this concerns insular species. Except L.

difficilis, no continental species is known from islands of

the Oriental Region. Supposedly, the records of this

species for Japan and Taiwan are based on misidentifica-

tions.

The knowledge on the distribution of the above-men-

tioned species is fragmentary; some of them are known

from the type locality only or from one or two regions of

Taiwan. New distributional data presented here supple-

menting the known records of the species. The data about

probable host plants for some species are also given. Two

new species of the genera Lipromorpha and Psylliodes are

described.

Received: 15.11.2011

Accepted: 02.04.2012

MATERIAL

The material depositories are abbreviated as following:

HNHM Museum of Natural History, Budapest

ZMHB Museum fir Naturkunde, Humboldt Uni., Berlin

BMNH Natural History Museum, London

TARI Taiwan Agricultural Research Insitute, Taichung

ZMUM Zoological Museum, Moscow State University

NC K. Nadein collection, Kiev

RESULTS

Psylliodes brettinghami Baly, 1862

Material examined. Taipei City. Taiwan: Taipei, Yang-

mingshan, 24.11.2008, leg. H. Lee, 2 ex. (TARI); Taiwan:

Taipei, Peihsinchuang, 20.1.2007, leg. S.-F. Yu, | ex.

(TARI). Yilan County. Taiwan: Ilan, Chiuchihtse,

07.XII.2008, leg. M.-H. Tsau, 4 exx. (TARI). Taoyuan

County. Taiwan: Taoyuan, Hsuanyuan, 16.[11.2008, leg.

M.-H. Tsao, | ex. (TARI). Pingtung County. Taiwan:

Pingtung, Tahanshan, 22.11.2007, leg. S.-F. Yu, 1 ex.

(TARI). Taitung County. Taiwan: Taitung, Imalintao,

04.11.2008, M.-H. Tsao, 3 exx. (TARI).

Remarks. Widely distributed in the Oriental region

(Scherer 1969; Déberl 2010); from Taiwan it is recorded

from Taito and Kwarenko (Kimoto & Takizawa 1997).

The material examined was collected on Solanum ameri-

canum Miller and Tubocapsicum anomalum (Franch. &

Sav.) Makino (Solanaceae).

Corresponding editor: D. Ahrens

42 Konstantin Nadein & Chi-Feng Lee

Figs 1-4. 1-3. Lipromorpha alutacea sp. n. 1. male, Hsitou; 2. female, Tengchih; 3. male, Tengchih. 4. Psylliodes palleola Motschul-

sky, lectotype.

Bonn zoological Bulletin 61 (1): 41-48 ©ZFMK

New data about Alticini from Taiwan 43

Psylliodes cantonensis Gruev, 1981

Type material examined. Psylliodes cantonensis,

paratypes: China: Canton W. E. Hoffman, 5 exx. (BMNH).

Additional material examined. Taitung County. Tai-

wan: Taitung, Hsiaolanyu, 25.V1.2009, leg. Y.-T. Chung,

7 exx. (TARI). Tainan County. Taiwan: Tainan, Yuching,

08.1.2009, leg. H.-T. Shih, 1 ex. (TARI).

Remarks. New to Taiwan. This species was described

from China (Guandong Prov.). The material examined dif-

fers from the type specimens by the blue metallic coloura-

tion of dorsum, the completely yellowish-reddish legs in-

cluding posterior femora, and the smaller punctures on the

pronotal disk with flat interstices, whereas the structure

of the aedaegus and hind tibia are identical.

Using the key in Kimoto & Takizawa (1997) these spec-

imens key to Psylliodes balyi Jacoby, 1884. According to

Scherer (1982) P. balyi is a synonym of P. chlorophana

Erichson, 1842, a species which is omitted in the Cata-

logue of Palaearctic Coleoptera by Dober! (2010). As con-

firmed by the study of the type material of the latter tax-

on (Psylliodes chlorophana, holotype and 6 paratypes:

Terra van Diem., Schayer, Nr. 56269. (ZMHB)) the two

species differ in the structure of the aedeagus — evenly

curved at lateral view in P. cantonensis and the apical half

nearly straight and then slightly curved at lateral view in

P. chlorophana. Possibly, at least in some cases this species

is confused with P. cantonensis when reported from Tai-

wan and other regions.

Psylliodes chujoe Madar, 1960

Material examined. Kaohsiung County. Taiwan:

Kaoshiung, Shihshan logging trail, 01—03.X.2008, leg. M.-

H. Tsao, 2 exx. (TARI).

Remarks. New to Taiwan. The species was described

from Japan (Kyushu). The specimens at our disposal dif-

fer somewhat from the type specimen (after original de-

scription) by the denser punctate vertex and pronotum, and

smaller size — 4 mm. In the revision of Japanese Psylliodes

(Takizawa 2005) the key thesis and diagnosis relating to

Psylliodes chujoe differs clearly from the original descrip-

tion in many characters such as much smaller body size,

shape and denser punctation of pronotum. This may sug-

gest that the real P. chujoe remained unknown to the au-

thor. Meanwhile, in the key of Japanese Psylliodes by Ki-

moto & Takizawa (1994) the characters and figures of

aedeagus correspond to the original description.

Bonn zoological Bulletin 61 (1): 41-48

Psylliodes subrugosa Jacoby, 1885

Material examined. Kaohsiung County. Taiwan:

Kaoshiang, Tengchih, 19.X1.2008, leg. C.-T. Yao, | ex.

(TARD); the same locality, 05—13.X1.2008, leg. C.-T. Yao,

1 ex. (TARI); the same locality, 25.X1.—03.X1.2008, leg.

C.-T. Yao, 1 ex. (TARI). Nantou County. Taiwan: Nan-

tou, Tatachia, 17—24.III.2007, leg. C.-S. Tung, | ex.

(TARI).

Remarks. New to Taiwan. The species is distributed in

Japan, Russia (Far East: Sakhalin), and China (Doberl

2010).

Psylliodes viridana Motschulsky, 1858 (Fig. 5)

Psylliodes palleola Motschulsky, 1866, syn. n.

Psylliodes angusticollis Baly, 1874, syn. n.

Type material examined. Psy/liodes viridana: Lectotype,

male: Psylliodes viridana Motch. Ceylon / Lectotypus

Psylliodes viridana Mts. L. Medvedev design. (ZMUM);

Paralectotypes: Psylliodes viridana Motch. Mt. Nuva Ell.

Ceyl. / Paralectotypus Psylliodes viridana Mts. L.

Medvedev design., 5 exx. (ZMUM).

Psylliodes angusticollis: Holotype: Type H.T. / Baly coll.

/ Psylliodes angusticollis Baly Japan (BMNH).

Psylliodes palleola: Lectotype, male: Psylliodes palleola

Motch. Ceylon / Lectotypus Psylliodes pallescens Mts. L.

Medvedev design. (ZMUM)

Additional material examined. Taoyuan County. Tai-

wan: Taoyuan, Lalashan, 22.VII.2008, leg. H.-J. Chen, |

ex. (TARI), the same locality, 02.I'V.2009, leg. C.-F. Lee,

1 ex. (TARI); Taiwan: Taoyuan, Hsuanyuan, 12.V1.2008,

leg. S.-F. Yu, 1 ex. (TART), the same locality, 16-.III.2008,

leg. M.-H.Tsao, | ex. (TARI). Taipei City. Taiwan: Taipei,

Sanchih, 20.11.2008, leg. H.-J. Chen, 1 ex. (TARI); Tai-

wan: Taipei, Shihlim, 13.III.2008, leg. H.-J. Chen, | ex.

(TARI); Taiwan: Taipei, Wulai, 04.111.2008, leg. S.-F. Yu,

1 ex. (TARI). Kaohsiung County. Taiwan: Kaoshiang,

Tengchih, 19.X1.2008, leg. C.-T. Yao, | ex. (TARI), the

same locality, 17.XII.2008, leg. C.-T. Yao, 1 ex. (TART),

the same locality, 05.11.2009, leg. M.-H. Tsau, 1 ex.

(TARI); Taiwan: Kaoshiang, Shihshan, logging trail,

01—03.X.2008, M.-H. Tsao, 17 exx. (TARI), the same lo-

cality, 06.11.2009, leg. S.-F. Yu & M.-H. Tsau, 5 exx.

(TARI), the same locality, 24.11.2009, leg. M.-H. Tsou,

1 ex. (TARI), the same locality, 19—24.X1.2008, leg. C.-

T. Yao, 1 ex. (TARI); Taiwan: Kaoshiang, Chuyunshan

logging trail, 24.111.2009, leg. C.-F. Lee, 6 exx. (TARI).

Taichung District. Taiwan: Taichung, Kukuan,

OZFMK

44 Konstantin Nadein & Chi-Feng Lee

Figs 5—7. 5. Psylliodes viridana Motschulsky: lectotype; 6-7. Psylliodes yuae sp. n.: 6. dorsal view; 7. lateral view.

16.VII.2007, leg. M.-H. Tsau, | ex. (TARI). Hsinchu

County. Taiwan: Hsinchu, Talulintao, 17.11.2008, leg. M.-

H. Tsao, 12 exx. (TARI); Taiwan: Hsinchu, Feifengshan,

O5.111.2009, leg. S.-F. Yu, 6 exx. (TARI); Taiwan:

Hsinchu, Hsinfeng, 31.11.2009, leg. S.-F. Yu, 4 exx.

(TARI); Taiwan: Hsinchu, Wufeng, 17.11.2009, leg. S.-F.

Yu, 1 ex. (TARI); Taiwan: Hsinchu, Paoerhshuiku,

04.XII.2008, leg. S.-F. Yu, 1 ex. (TARI).

Remarks. Comparison of the type material reveals clear-

ly that Psylliodes angusticollis is conspecific with Psyl-

liodes viridana by the characteristic impunctate vertex,

body outline, structure of hind tibia, and aedeagus.

Examination of the lectotype of Psylliodes palleola re-

veals its significant similarity to P. viridana. The only ex-

ception is colouration of the body, which is entirely yel-

low including legs in P. palleola (Fig. 4) in contrast to

greenish dorsum and dark legs in P. viridana (Fig. 5). So

far, no pale species of Psyllodes have been found in the

Oriental Region. The reason of this colouration is very

likely that the type specimen appears to be a teneral adult,

which is not fully coloured. This is additionally confirmed

by the incompletely developed and partly folded right

elytron and the weakly sclerotized abdominal ventrites and

aedeagus. Type material of both species was collected at

the same locality “Nuwara Eliya” in Sri Lanka. The lec-

Bonn zoological Bulletin 61 (1): 41-48

6 7

totype of P. palleola was designated by Medvedev (2006)

and erroneously labeled “Lectotypus Psylliodes pallescens

Mts. L. Medvedev design.”; therefore, a new label with

correct name is added to the specimen.

This species was previously reported for Taiwan from

Kosempo (Kimoto & Takizawa 1997). Distributed also in

S. India, Sri Lanka, Nepal, N. Vietnam, S. China, Korea,

Japan, Russia (Far East: Primorskiy Terr., Sakhalin, South

Kuriles) (Doberl 2010). The material from Taiwan was

collected on Solanum americanum Miller, S. lyratum

Thunberg, S. peikuoensis S. S. Ying (Solanaceae).

Psylliodes yuae sp. n. (Figs 6, 7, 18-21)

Type material. Holotype, male: Taiwan: Taoyuan,

Lalashan, 26.V1.2007, leg. S.-F. Yu (TARD); paratype: the

same label data as holotype, 1 male, 1 female (TARI), 1

male (NC).

Description. Body (Figs 6, 7) elongate-oval, convex; dor-

sum shining black with metallic luster, ventral side black,

antennae dark brown with three basal segments lighter,

yellowish-brownish, tibiae and femora dark brown to near-

ly black, knees and tarsi light brown.

©ZFMK

New data about Alticini from Taiwan 45

Head short, vertex rather convex, medially covered with

sparse punctures, punctation at sides of vertex closely to

eyes denser, surface between punctures smooth; ocular sul-

ci well developed, deep, above eyes very wide and ex-

tremely deep, forming large impression containing large

setiferous pore; frontal calli narrow, separated by frontal

ridge, their surface rough, calli delineated by lines from

vertex, In some cases area containing calli and nearest part

of vertex coarsely puncture-wrinkle sculptured, sometimes

calli indistinct; frontal ridge wide, flattened, nearly

smooth, anterofrontal ridge flattened, weakly convex; eyes

comparatively large, oval, convex; labrum moderately

short and wide.

Pronotum subquadrate, in male |.2—1.3 times as wide

as long, nearly equal in length and width in female, rather

convex, especially in anterior half and with sharply slop-

ing sides; anterior margin straight, posterior margin form-

ing obtuse angle, lateral edges weakly converging anteri-

orly, lateral margins nearly straight, thin and smooth,

pronotum above anterior angles and along lateral margin

distinctly convex, lateral side with clear rounded impres-

sion, anterolateral callosities large with well developed,

large, projecting angle, posterolateral callosities also well

developed, somewhat smaller and angularly projecting;

pronotal surface covered with moderately small and shal-

low punctures, distance between them an average |.5—2

times their diameter, surface between punctures smooth.

Elytra convex, at base together hardly wider than prono-

tal base; 1.5—1.6 times longer than wide, maximal width

before middle; scutellum small, with obtuse apex, impunc-

tate; humeral calli small, not very convex; sutural angle

obtuse, epipleurae smooth; elytra with transversal shallow

and narrow impression at basal margin; punctures in stri-

ae small, hardly larger than pronotal, not deep, distance

between punctures in striae is about diameter of puncture,

distance between striae 2—3 times diameter of puncture,

striae not deep, well developed at whole distance; inter-

stices flat or slightly convex, covered with a regular row

of punctures smaller than strial, interstices between punc-

tures smooth; apical part of second striae (the next after

short scutellar) clearly impressed.

Legs thin and straight, first tarsomere of male protarsus

poorly widened, just somewhat wider than in female; pos-

terior tibia clearly curved (Fig. 18), inner ridge at tarsal

articulation with tooth, first tarsomere of posterior tarsus

short and straight.

Aedeagus (Figs 19, 20) with triangular apex, apical sides

straight, from base to apex nearly parallel-sided, at ven-

tral side with narrow and moderately deep furrow; at lat-

eral view apex sinuate and curved, then straight and weak-

ly gradually widened towards basal opening; tegmen typ-

ically Y-shaped.

Spermatheca (Fig. 22) with short collum, nodulus long

and narrow, duct short and moderately thin, not coiled but

curved at base.

Bonn zoological Bulletin 61 (1): 41-48

Body length: male — 2.5—2.7 mm, female — 3.2; body

width: male — 1.2—1.3 mm, female — 1.6.

Remarks. The new species is similar to P. viridana and

P. cantonensis from China (Guandong). It differs from P.

viridana in colouration, a much more convex vertex with

distinct punctation on its surface, by, a significantly con-

vex pronotum with smooth interstices, much more curved

posterior tibia, and the shape and structure of aedeagus.

The new species differs from P. cantonensis in coloura-

tion, proportions and shape of the body, the more convex

vertex, much sparser punctation of pronotum and elytra,

more curved posterior tibia and also in shape and struc-

ture of aedeagus and spermatheca.

The type material was collected on Lycianthes lysimac-

hioides (Wall.) Bitter (Solanaceae).

Distribution. Taiwan (Taoyuan County).

Etymology. The species is named after Mrs. Su-Fang Yu,

who collected the type specimens.

Taiwanoliprus endonis Komiya, 2006

Material examined. Yilan County. Taiwan: Ilan,

Mingchi, 12.1V.2009, leg. M.-H. Tsou, | male (TARI); the

same locality, 05.1V.2009, 1 male (TARI).

Remarks. Previously known from the type locality in

Taoyuan County. The material was collected on Ampelop-

sis sp. (Vitaceae).

In the original description, differences between 7Zai-

wanoliprus wenroni and of T. endonis as stated by the au-

thor are based mostly on colouration of the body, struc-

ture of elytral apex and aedeagus. Zaiwanoliprus wenroni

is entirely yellowish whereas 7. endonis with dorsum red-

dish brown, underside and apical 2/3 of hind femora black-

ish brown. In the material at our disposal, a specimen iden-

tified as T. endonis possesses colouration of 7. wenroni

but without emarginate apex of elytron and with the shape

of aedeagus as in 7. endonis. The colouration of other

specimen is lighter than in the original description of 7.

endonis. It can be suggested that colouration of the species

of this genus 1s rather variable and could not be a reliable

character for distinguishing of species. The structure of

aedeagi of both species 1s rather similar and normally lies

within the limits of intraspecific variability as often pre-

sented in alticines. Additional study is necessary to either

confirmation for species status or synonymization of T.

wenroni.

©ZFMK

46 Konstantin Nadein & Chi-Feng Lee

15 16 17

18 19 20 21 22

Figs 8-22. 8-17. Lipromorpha alutacea Nadein, sp. n. 8-14. Aedeagus. 8. ventral view, Anmashan; 9. lateral view; 10. ventral

view, Hsitou; 11. lateral view; 12. ventral view, holotype, Tengchih; 13. holotype, dorsal view; 14. lateral view; 15. prothorax, dor-

sal view; 16. head, lateral view; 17. fore tarsus of male. 18-21. Psylliodes yuae sp. n. 18. right posterior tibia, inner side; 19. aedea-

gus, ventral view, holotype; 20. aedeagus, lateral view; 21. spermatheca; 22. Lipromorpha montana Chuj6, aedeagus, ventral view

and lateral view (after Kimoto & Takizawa 1997).

Bonn zoological Bulletin 61 (1): 41-48 ©ZFMK

New data about Alticini from Taiwan 47

Pseudoliprus kimotonis Komiya, 2006

Material examined. Taoyuan County. Taiwan: Taoyuan,

Lalashan, 14.V.2009, leg. C.-F. Lee, 3 exx. (TARI); the

same locality, 02.[V.2009, leg. H.-G. Chen, 2 exx.

(TARI); the same locality, 01.1V.2009, leg. C.-F. Lee, 4

exx. (TARI). Yilan County. Taiwan: Ilan, Mingchi,

05.1V.2009, leg. M.-H. Tsou, 2 exx. (TARI). Taipei Coun-

ty. Taiwan, Taipei Hsien, Fu-Shan LTER site, lake shore,

meadow, swept, 25.11.2003, leg. L. Papp & M. Foldvari,

1 ex. (HNHM). Xitun District. Taiwan, Taichung Hsien,

Anmashan Mts, 24°14’28.9” N 120 °56’45.8” E, 1800 m,

dry creek bed, 5.IV.2003, leg. L. Papp & M. Foldvari, 1

ex. (HNHM).

Remarks. Previously known from Nantou County and

Chiayi (East District) after original description. The ma-

terial was collected on Actinidia callosa Lindl. (Actini-

diaceae).

Pseudoliprus saigusai Kimoto, 1970

Material examined. Nantou County. Taiwan: Nantou,

Tatachia, 20.VII.2009, leg. S.-F. Yu, 7 exx (TARI).

Remarks. Previously known from various localites of

Nantou County (Kimoto & Takizawa 1997). The materi-

al was collected on Polygonum cuspidatum Sieb. & Zucc.

(Polygonaceae).

Lipromorpha shirozui Kimoto, 1970

Material examined. Yilan County. Taiwan: Ilan, Fushan,

Chilwuyuan, 20.11.2009, leg. C.-F. Lee, 4 exx. (TARD;

Taiwan: Ilan, Mingchi, 05.IV.2009, leg. M.-H. Tsou, | ex.

(TARI); the same locality, 16. VIII.2008, leg. M.-H. Tsou,

3 exx. (TARI). Taipei County. Taiwan: Taipei, Yangming-

shan, 07.1X.2007, leg. S.-F. Yu, 1 ex. (TARI); Taiwan,

Taipei county, Pi Hu, 3.1V.2002, at light, Gy. Fabian & O.

Merkl, 1 ex. (HNHM). Kaohsiung County. Taiwan:

Kaoshiang, Tengchih, 05.11.2009, leg. S.-E. Yu, 1 ex.

(TARI).

Remarks. Previously reported from Nantou County (Ki-

moto & Takizawa 1997).

Lipromorpha alutacea Nadein, sp. n. (Figs 1-3, 8-17)

Type material. Holotype, male. Kaohsiung County: Tai-

wan: Kaoshiang, Tengchih, 05.11.2009, leg. S.-F. Yu (TA-

RI); Paratypes: the same locality, 2 exx. (TARI), 3 exx.

(NC); the same locality, 18.11.2007, leg. S.-F. Yu, 4 exx.;

Bonn zoological Bulletin 61 (1): 41-48

the same locality, 26.V.2009, leg. C.-F. Lee, 1 exx. (TA-

RI); Taiwan: Kaoshiang, Shihshan logging trail,

24.111.2009, leg. M.-H. Tsou, 2 exx. (TARI). Nantou

County. Taiwan: Nantou, Hsitou, 06.V.2009, leg. C.-F.

Lee, 1 ex. (TARI). Taichung County. Taiwan, Taichung

Hsien, Anmashan Mts, 24°14’°28.9” N 120 56’ 45.8” E,

1800 m, 1800 m, dry creek bed, 5.1V.2003, leg. L. Papp

& M. Foldvari, | male (HNHM); Taiwan: Taichun, An-

mashan, 22.1X.2007, leg. M.-H. Tsou, | female. (TARI).

Description. Dorsum, antennae, and legs yellowish-light

brown, underside brown except last abdominal segment,

apical two thirds of hind femora blackish brown; varia-

tions of colouration: sides of pronotum under lateral mar-

gin, short stripe on medial part of elytra, short stripe at

lateral side of elytra below humeral calli dark brown to

nearly black (Figs 1-3).

Head (Fig. 16) short, vertex nearly flat, its surface

minutely shagreened, medially impunctate, with a few

punctures bearing long setae at sides; frontal calli small,

triangular, penetrating between antennal sockets, area

above frontal calli triangularly concave and sometimes

with shallow short suture, external angles of frontal calli

sometimes with a shallow and short sulci reaching mar-

gin of eye; frontal part strongly raised, at lateral view ver-

tex and frons forming nearly right angle, frons below an-

tennal sockets triangular, flat, smooth, genae about twice

shorter than longitudinal diameter of eye or shorter;

labrum transverse, short, mandibulae short, their apices

brown to blackish brown; eyes medium-sized, oval,

strongly convex; antennae somewhat shorter than body

length, 6th and 7th segment equal in length, 8th—10th seg-

ment shorter and thicker than previous four segments.

Pronotum (Fig. 15) 1.05—1.13 times longer than wide

in male, and 1.07—1.10 times in female; anterior margin

slightly curved medially, posterior margin weakly round-

ed, pronotal base shorter than anterior edge, sides with

margin at anterior half, basal constriction deep, lateral

sides behind anterolateral callosities more or less deeply

grooved, at lateral view pronotal upperside nearly straight

or feebly convex medially, anterolateral callosities large,

convex, obtuse; pronotal surface densely punctate, dis-

tance between punctures do not exceed diameter of punc-

ture, punctures shallow, comparatively large, interstices

flat, distinctly and strongly shagreened, covered with short,

dense, and decumbent hairs.

Scutellum small, triangular, acute, its surface similar to

that of pronotum. Elytra nearly parallel-sided, barely

widened just behind middle, 1.55—1.68 (1.61) times longer

than width in male, 1.45—1.67 (1.59) times in female;

humeral calli strongly raised, impunctate, hind wings com-

pletely developed; base of elytra strongly convex, then

distinctly concave, apical half convex; apices rounded, su-

tural angle not acute, obtusely-rounded; punctures very

large and deep, arranged in regular striae without tenden-

OZFMK

48 Konstantin Nadein & Chi-Feng Lee

cy to confusion, punctures in stria dense, situated direct-

ly at each other, striae deeply engraved at whole distance,

short scutellar row of punctures usually engraved deeper

than others; interstices strongly costate, costae distinct

from base to apex, short costa between short scutellar row

and next row strongly raised, usually stronger than oth-

ers, surface of costae impunctate, smoothly and minute-

ly shagreened, each costa bears a row of long, semi-erect,

comparatively dense hairs.

Femora distinctly swollen, fore tibiae apically curved;

first tarsomere of male slightly broader than in female (Fig.

17).

Aedeagus (Figs 8—14) short, apical third narrow, weak-

ly narrowing apically, apex rounded, medial third broad,

nearly parallel-sided, basal third with large basal opening;

dorsal opening bears long, thin, and strongly curved fla-

gellum.

Body length: male — 1.9-2.4 mm, female — 2.2-2.5;

width: male — 0.9—1.1 mm, female — 1.0-1.1.

Remarks. Lipromorpha alutacea resembles L. montana

Chtaj6, 1935 known from type locality in Chiayi County

(Alishan), from which it differs in the shape of aedeagus:

in L. montana (Fig. 22) the apical third of aedeagus is dis-

tinctly triangular and acute whereas the apex of L. alutacea

is triangularly-rounded and obtuse or apical third in some

specimens is rounded, not triangular (Figs 8-14); it also

differs in the proportions of pronotum and in colouration,

especially the black legs and basal segments of abdomen

in L. montana, whereas the new species has only apical

two third of hind femora brownish black.

The type material was collected on Polygonum chinen-

se L. (Polygonaceae).

Distribution. Taiwan (Kaohsiung County, Nantou Coun-

ty, Taichung County).

Etymology. The species name refers to characteristic sha-

greened surface of head, pronotum, and elytral interstices.

Taxonomic note. The name Lipromorpha cyanea L.

Medvedev, 2009 (Laos) is a junior primary homonym of

Lipromorpha cyanea Chen et Wang, 1980 (China: Yun-

nan). In the original description the author stated that L.

cyanea Medvedev is the only species with metallic blue

colouration of the body among the continental species of

the genus. That is not correct. According to the original

descriptions the both species are highly similar to each oth-

er; hence, in view of the fact that Laos and Yunnan

Province are bordering it can be suggested that they are

conspecific. We do not to propose a new replacement

name because the genus currently is under taxonomic re-

vision by the first author. The final decision will be based

on the examination of the type material.

Bonn zoological Bulletin 61 (1): 41-48

Acknowledgements. The first author is grateful to O. Merkl

(HNHM), N. Nikitsky (ZMUM), S. Shute (BMNH), J. Frisch

(ZMHB) for the opportunity to study material in their care. We

thank the Taiwan Chrysomelid Research Team for assisting with

field collecting and observations and C. Staines (Smithsonian

Institution, Washington) for the linguistic help.

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©ZFMK

Bonn zoological Bulletin 61 (1): 49-128 July 2012

A revision of East Palaearctic Lobrathium

(Coleoptera: Staphylinidae: Paederinae)

Volker Assing

Gabelsbergerstr. 2, D-30163 Hannover, Germany; E-mail: vassing.hann@t-online.de.

Abstract. East Palaearctic species of Lobrathium Mulsant & Rey, 1878, particularly those of the Himalaya and China,

are revised. Thirty-three species, thirty-one of them in Lobrathium and two of lobrathioid habitus in Lathrobium Graven-

horst, 1802, are described and illustrated: Lobrathium ablectum sp. n (China: Hubei); L. biaculeatum sp. n. (East Nepal);

L. bicarinatum sp. n. (North India, Nepal); L. bicornutum sp. n. (East Nepal); L. bimembre sp. n. (China: Yunnan); L.

bispinosum sp. n. (China: Guizhou); L. cholaicum sp. n. (China: Tibet); L. configens sp. n. (China: Shaanxi, Sichuan,

Hubei, Yunnan); L. cornutum sp. n. (East Nepal); L. daxuense sp. n. (China: Sichuan); L. demptum sp. n. (China: Hubei);

L. discrepans sp. n. (North Vietnam); L. domenoides sp. n. (China: Sichuan); L. duplex sp. n. (China: Sichuan, Yunnan);

L. excisissimum sp. n. (Yunnan); L. feldmanni sp. n. (China: Sichuan); L. flavipenne sp. n. (North India: Himachal Pradesh);

L. integrum sp. n. (East Nepal); L. kleebergi sp. n. (Nepal); L. kosiense sp. n. (East Nepal); L. Jamellatum sp. n. (China:

Sichuan); L. mordens sp. n. (North India); L. radens sp. n. (China: Guizhou); L. retrocarinatum sp. n. (China: Yunnan);

L. schuelkei sp. n. (China: Shaanxi); L. semiflavum sp. n. (Russian Far East); L. sinuatum sp. n. (Central Nepal); L. spathu-

latum sp. n. (China); L. taureum sp. n. (China: Hubei, Beijing); L. tuberosum sp. n. (China: Jiangxi); L. unispinosum sp.

n. (Nepal); Lathrobium lobrathioides sp. n. (China: Sichuan); L. lobrathiforme sp. n. (China: Yunnan). All the previou-

sly described species from the Himalaya and China, except for L. emeiense Zheng, 1988, and some species from other

East Palaearctic regions are redescribed and illustrated. Five new combinations are established: Lobrathium guttula (Fau-

vel, 1895), comb. n. (ex Lathrobium); L. pustulatum (Cameron, 1931), comb. n. (ex Lathrobium); L. kashmiricum (Ca-

meron, 1931), comb. n. (ex Lathrobium); Tetartopeus gracilentus (Kraatz, 1859), comb. n. (ex Lathrobium),; T. wui (Zheng,

2001), comb. n. (ex Lobrathium). Lathrobium sublaeve Motschulsky, 1858 is excluded from both Lathrobium and Lo-

brathium, and treated as Paederini incertae sedis. Lobrathium ochreonotatum (Champion, 1922), previously a synonym

of L. semicaeruleum (Cameron, 1921), is revalidated. Lobratium shibatai varium Ito, 1995 is elevated to species rank.

Eight synonymies are proposed: Lobrathium Mulsant & Rey, 1878 = Ponthrobium Korge, 1971, syn. n.; Lobrathium al-

ticola (Cameron, 1943) = L. pirpanjalense Coiffait, 1982, syn. n.; L. hongkongense (Bernhauer, 1931) = L. sibynium Zheng,

1988, syn. n., = L. ryukyuense Ito, 1996, syn. n.; L. triste (Cameron, 1924) = L. kashmiricum (Cameron, 1931, syn. n.,

= L. afghanicum Coiffait, 1979, syn. n., = L. nouristanicum Corffait, 1979, syn. n.; Tetartopeus wui (Zheng, 2001) = 7.

bimaculatum (Li, Tang & Zhu, 2007), syn. n. Lectotypes are designated for Lathrobium lederi Eppelsheim, 1884, L. ba-

dium Cameron, 1924, L. brunneum Cameron, 1931, L. triste Cameron, 1924, L. semicaeruleum Cameron, 1921, L. ochreo-

notatum Champion, 1922, L. guttula Fauvel, 1895, L. pustulatum Cameron, 1931, L. cylindricolle Cameron, 1924, L.

hongkongense Bernhauer, 1931, and L. gracilentum Kraatz, 1859. Lobrathium, according to currently available evidence

a Holarctic genus, is now represented in the Palaearctic region by 114 species and one subspecies, the vast majority of

which, 87 species and one subspecies, are distributed in the East Palaearctic including Middle Asia, Myanmar, and North

Vietnam. The Himalaya (20 species), China (24 species), Taiwan (20 species), and Japan (18 species and one subspecies)

are the regions with the greatest diversity. Most species, particularly those with reduced wings and those subject to wing

dimorphisms, appear to have remarkably restricted distributions. A catalogue of the Lobrathium species recorded from

the Palaearctic region and keys to the species of the Himalaya and of China are provided. Some species from China and

the Himalaya are subject to pronounced wing dimorphisms. East Palaearctic Lobrathium species are partly found near

lakes or on banks of rivers and streams, and partly in the leaf litter of forest and shrub habitats at altitudes of 450-4400

m. Additional records of several West Palaearctic species are reported.

Key words. Taxonomy, rove beetles, East Palaearctic region, Himalaya, China, new species, new synonymies, new com-

binations, revalidation, lectotype designations, key to species, catalogue, diversity, zoogeography, wing dimorphism.

INTRODUCTION

The lathrobiine genus Lobrathium Mulsant & Rey, 1878, | zoogeographic regions have been attributed to this genus,

which had been treated as a subgenus of Lathrobium _ too, but all those taxa that have been re-examined recent-

Gravenhorst, 1802 until a few decades ago, probably has __ ly, have turned out to belong to other genera (e.g., Ass-

an essentially Holarctic distribution. Species from other ing in press c).

Received: 15.03.2012 Corresponding editor: D. Ahrens

Accepted: 20.04.2012

50 Volker Assing

According to Smetana (2004), Lobrathium is represent-

ed in the Palaeactic region by 62 species and three sub-

species in three subgenera. In the meantime, however, nu-

merous additional species have been described, one sub-

generic and numerous species-group names have been

synonymised, one species was transferred to Tetartopeus

Czwalina, 1888, two to Pseudolathra Casey, 1905, and

one species previously assigned to Lathrobium was moved

to Lobrathium (Assing 2004, 2005a, 2006a, 2007, 2008,

2010, in press a; Bordoni 2009; Ito 2007, 2009a—c;

Shavrin 2008). Moreover, Lobrathium wui Zheng, 2001

from China and L. apogeum (Normand, 1936) from Al-

geria had been omitted in the catalogue.

According to Newton et al. (2001), nearly 70 species

of Lobrathium, two of them adventive, are known from

the Nearctic region. However, this figure includes

Pseudolathra Casey, 1905, which, as most recent authors

agree, represents a distinct genus (Assing in press c). Ac-

cording to a database compiled by Newton (unpubl.), there

are 36 Lobrathium and 33 Pseudolathra species in North

America north of Mexico (Newton, pers. comm.), and on-

ly one of them, Lobrathium multipunctum Gravenhorst,

1802, is adventive.

Previous studies have suggested already that the Lo-

brathium fauna of the East Palaearctic is much more di-

verse than that of the West Palaearctic region. The best-

studied regions are Japan and Taiwan. As many as 20

species and subspecies, most of them with more or less

restricted distributions, have been reported from Japan

alone (Ito 2007, 2009b; Smetana 2004). For the vast ma-

jority of these species, illustrations of the male primary

sexual characters are available in the literature. Accord-

ing to a recent revision, 20 species, 19 of them endemic,

are known from Taiwan (Assing 2010). Remarkably, on-

ly seven species have been recorded from mainland Chi-

na, although this vast, geologically, zoogeographically, and

ecologically diverse country is generally known to host a

remarkably rich fauna. Aside from the recent revision of

the Lobrathium species of Taiwan (Assing 2010) and an

incomplete key to the Lobrathium species of the Himalaya

(Coiffait 1982b), which also includes a species of

Pseudolathra, synoptic taxonomic studies of the East

Palaearctic Lobrathium fauna are absent.

A recent revision of the Himalayan Lathrobium species

(Assing in press b) revealed that the type material of sev-

eral species previously attributed to Lathrobium in fact be-

longed to Lobrathium and other genera of Lathrobiina and

Medonina. Subsequently, in order to investigate the diver-

sity of Lobrathium in the East Palaearctic and to clarify

the taxonomic status of previously unrevised Lobrathium

and Lathrobium species, more type and non-type materi-

al from various public and private collections was exam-

ined. These studies yielded a remarkable number of new

species, new synonymies, new combinations, and other

taxonomic changes. In addition, new West Palaearctic ma-

Bonn zoological Bulletin 61 (1): 49-128

terial examined since the latest instalments (Assing 2007,

2008) to a revision of West Palaearctic Lobrathium is re-

ported.

MATERIAL AND METHODS

The morphological studies were conducted using a Ste-

mi SV 11 microscope (Zeiss Germany) and a Jenalab com-

pound microscope (Carl Zeiss Jena). A digital camera

(Nikon Coolpix 995) was used for the photographs.

Head length was measured from the anterior margin of the

frons to the posterior margin of the head, elytral length at

the suture from the apex of the scutellum to the posteri-

or margin of the elytra, the length of the forebody from

the anterior margin of the mandibles (in resting position)

to the elytral hind margin, and the length of the aedeagus

from the apex of the ventral process to the base of the

aedeagal capsule. The “parameral” side (i.e., the side

where the sperm duct enters) is referred to as the ventral,

the opposite side as the dorsal aspect.

The maps were created using MapCreator 2.0 (primap)

software. The coordinates of some Himalayan localities

were obtained from Ahrens (2004).

COLLECTION MATERIAL DEPOSITORIES

BMNH_ The Natural History Museum, London (R. G. Booth)

FMNH_ Field Museum of Natural History, Chicago (via L. H.

Herman)

MCSNV Museo Civico di Storia Naturale, Verona (L. Latella;

via A. Zanetti)

MHNG Muséum d’Histoire Naturelle, Genéve (G. Cuccodoro)

MMUM The Manchester Museum, The Manchester Universi-

ty (D. Logunov)

MNHNP Muséum National d'Histoire Naturelle, Paris (A.

Taghavian)

NHMB Naturhistorisches Museum Basel (M. Brancucci, E.

Sprecher)

Natural History Museum Denmark/ University of

Copenhagen Zoological Museum (A. Solodovnikov)

NHMW Naturhistorisches Museum Wien (H. Schillhammer)

NHMD

NME _ Naturkundemuseum Erfurt (M. Hartmann, assisted by

W. Apfel)

SDEI —Senckenberg Deutsches Entomologisches Institut,

Miuncheberg (L. Behne)

SF Senckenberg, Frankfurt/M. (A. Hastenpflug-Vesmanis)

SMNS | Staatliches Museum fiir Naturkunde, Stuttgart (W.

Schawaller, K. Wolf-Schwenninger)

SNSD _ Senckenberg Naturhistorische Sammlungen Dresden

(O. Jager)

ZML = Museum of Zoology, Lund University (R. Danielsson)

cAss author’s private collection

cFel private collection Benedikt Feldmann, Munster

©ZFMK

Revision of East Palaearctic Lobrathium

cGia private collection Pier Mauro Giachino, Torino, Italy

cKle private collection Andreas Kleeberg, Berlin, Germany

cSch private collection Michael Schiilke, Berlin, Germany

cSha private collection Alexey Shavrin, Daugavpils, Latvia

cSme private collection Ales Smetana, Ottawa, Canada

cWun private collection Paul Wunderle, Ménchengladbach,

Germany

RESULTS

Systematic position and identification of genus. Lo-

brathium belongs to the paederine subtribe Lathrobiina,

which is represented in the Palaearctic region by twelve

genera: Achenium Leach, 1819, Domene Fauvel, 1873,

Lathrobium Gravenhorst, 1802, Lobrathium Mulsant &

Rey, 1878, Micrillus Raffray, 1873, Platydomene Gan-

glbauer, 1895, Pseudobium Mulsant & Rey, 1878,

Pseudolathra Casey, 1905, Scymbalium Erichson, 1839,

Scymbalopsis Reitter, 1909, Tetartopeus Czwalina, 1888,

and Throbalium Mulsant & Rey, 1878. The phylogenetic

affiliations of the lathrobiine genera have never been com-

prehensively studied, so that the polarity of most distin-

guishing characters is unknown and the identification of

apomorphies difficult. For keys to the lathrobiine genera

represented in the Palaearctic and more detailed diagnoses

see Coiffait (1982c) and Assing (2012).

Lobrathium is distinguished from similar genera such

as Lathrobium, Pseudolathra, Platydomene, Pseudobium,

Pseudolathra, and Tetartopeus by the following charac-

ter combination:

Body size moderate to large, not conspicuously flat-

tened. Head more or less coarsely punctate and with broad

posterior constriction of approximately half the width of

head. Pronotum distinctly oblong, usually 1.15—1.35 times

as long as broad, with relatively coarse punctation, and

without a separate dorsal series of punctures on either side

of the usually distinct, impunctate median band. Elytra

with rather coarse and usually defined, often seriate punc-

tation; above lateral margin (in lateral view) with addition-

al fine line (evidently secondarily reduced in some species

from Taiwan). Male sexual characters: sternite VII often

with pronounced modifications (median impression, pos-

terior excision, presence of strongly modified, short and

stout black setae); sternite VII usually with rather large

posterior excision and mostly with strongly modified,

short and stout black setae; aedeagus without parameres,

without strongly sclerotised spines in internal sac; ventral

process of aedeagus often more or less blade-shaped and

with ventral carinae or tooth-like projections. Female ter-

minalia without conspicuous modifications.

The genera that have been most frequently confused

with Lobrathium are Lathrobium, Tetartopeus,

Pseudolathra, and Pseudobium. In Lathrobium, the punc-

tation of the forebody, particularly that of the elytra, is

Bonn zoological Bulletin 61 (1): 49-128

mostly less dense, finer, and less defined than in Lobrathi-

um, never arranged in series on the elytra, the pronotum

is often less convex in cross-section and less oblong, the

elytra lack the additional epipleural line, the chaetotaxy

of the male sternites VII and VIII is different (often mod-

ified, but not with the clusters of short and stout black se-

tae found in Lobrathium), the aedeagus often has strong-

ly sclerotised sclerites (spines, hooks, etc.) in the internal

sac, and the female terminalia are modified to various de-

grees (see, e.g., Assing in press b). In 7etartopeus, the pos-

terior constriction of the head is more slender (approx1-

mately one-third of head width), the elytra lack the sub-

marginal line, the elytral punctation is finer and not seri-

ate, the pubescence of the male sternite VIII is fine and

dense, the posterior excision of this sternite is very small

and often in more or less asymmetric position, the aedea-

gus often has pronounced spines in the internal sac and a

more or less spine-shaped ventral process, and the female

terminalia are modified to various degrees (see, e.g.,

Assing 2011b). In Pseudolathra, the forebody is much

more sparsely punctate, the pronotum has a distinct series

of punctures on either side of the impunctate median band,

the male sternites VH and VIII lack the short stout setae

typical of Lobrathium, the posterior excision of sternite

VIII is — at least in Palaearctic and Oriental species — al-

most always very deep and narrow, and the aedeagus is

of completely different morphology (see Assing in press

c). Pseudobium species are of more slender habitus, usu-

ally much smaller, have a more oblong pronotum (approx-

imately 1.5 times as long as broad), a more finely and

sparsely punctate forebody, a more oblong and distinctly

subrectangular head with longer postgenae and with the

eyes situated before the middle, a more pronounced seri-

ate arrangement of the elytral punctures, a relatively longer

metatarsomere I (distinctly longer than II), male sternites

VII and VII without strongly modified setae, and an

aedeagus of completely different morphology (for illus-

trations see, e.g., Assing 2006b).

Intrageneric affiliations. Lobrathium is evidently repre-

sented in the Palaearctic region by numerous lineages,

some of them currently including only one species (e.g.,

L. cylindricolle, L. discrepans) and some of them speciose

and widespread. However, since many of the species from

Japan and from the Nearctic region have not been exam-

ined yet, a comprehensive species group concept is nei-

ther attempted nor proposed. On the other hand, the sub-

generic classification currently in use is highly doubtful.

Five species from the Mediterranean and from northeast-

ern Turkey have been attributed to the subgenus Ponthro-

bium Korge, 1971 and the remainder to the nominate sub-

genus, a systematic concept that is highly artificial for sev-

eral reasons. Ponthrobium is most unlikely to form a

monophyletic group, but represented by two lineages, one

©OZFMK

of them including three species (including the type species

of Ponthrobium) from northeastern Turkey and one com-

prising two or three species (affiliations of L. apogeum un-

clear) distributed in North Africa and southern Italy. Even

more importantly, maintaining Ponthrobium as a valid

name would undoubtedly render the nominate subgenus

paraphyletic and completely distort the true intrageneric

phylogenetic affiliations; Ponthrobium is most unlikely to

form the sister group of all other Lobrathium species. Un-

der these circumstances there are two options, either sub-

divide the nominate subgenus into numerous additional

subgenera or synonymise Ponthrobium. The latter option

is deemed to be in better accordance with the stability of

nomenclature, also because the genus has not been fully

revised (see above). Hence the following synonymy re-

sults: Lobrathium Mulsant & Rey, 1878 = Ponthrobium

Korge, 1971, syn. n.

Diversity and zoogeography. After the revision, Lo-

brathium is represented in the Palaearctic region (includ-

ing North Vietnam) by 114 species and one subspecies.

The fauna of the West Palaearctic, exclusive of Middle

Asia, comprises 26 species; the diversity hot spots are lo-

cated in the Balkans, Turkey, and the Caucasus region. The

fauna of the East Palaearctic, including Middle Asia,

Myanmar, and North Vietnam, is much more diverse. At

present, 88 named species and one subspecies are known

from this region, four from Middle Asia (exclusive of

Afghanistan), 20 from the Himalaya (including North

Afghanistan), 24 from mainland China, 20 from Taiwan,

18 species and one subspecies from Japan, and the remain-

der from other regions (Siberia, Russian Far East, Myan-

mar, North Vietnam). The widespread L. hongkongense

is included in the figures for Japan, Taiwan, and mainland

China.

Most of the East Palaearctic Lobrathium species have

more or less restricted distributions. The most widespread

species is L. hongkongense, which has been recorded from

numerous localities in China, from Taiwan, and from the

extreme south of Japan. Interestingly, none of the Hi-

malayan species has been recorded from China, and vice

versa. However, at least one species group containing ex-

clusively micropterous and wing-dimorphic species (L.

kosiense, L. wittmeri, L. cholaicum, L. daxuense, L.

domenoides, L. bimembre, L. lamellatum, L. duplex) is dis-

tributed from Nepal to Sichuan and Yunnan. In the course

of the present revision, none of the species from the main

islands of Japan was recorded also from continental Asia.

The only species that was found to be present in both

Japan and continental Asia is L. hongkongense, whose dis-

tribution in Japan, however, is confined to the extreme

south (Ryukyu Islands). Finally, no Lobrathium species

appears to have a trans-Palaearctic distribution.

Bonn zoological Bulletin 61 (1): 49-128

Volker Assing

The available evidence suggests that the true diversity

of the East Palaearctic Lobrathium fauna, particularly that

of the Himalaya, China, and Taiwan is significantly greater

than currently known. In the course of the present revi-

sion and of the revision of the species of Taiwan (Assing

2010), a considerable number of probably undescribed

species was not described because they were represented

only by females. Moreover, numerous named species have

been recorded only once or very rarely and seem to have

very restricted distributions. Finally, in comparison to the

West Palaearctic, Taiwan, and Japan, the faunas of the Hi-

malaya and mainland China have been poorly studied. The

taxonomic status of one species, L. emeiense, is doubtful,

as its male sexual characters are unknown.

Taxonomic changes. In all, 31 species of Lobrathium and

two species of Lathrobium are described for the first time.

Seven species-group names are syonymised, six of them

in Lobrathium and one in Tetartopeus. One previously

synonymised name, Lobrathium ochreonotatum, 1s reval-

idated. Five new combinations are established: three

names are transferred from Lathrobium to Lobrathium,

and two species, one of them previously in Lobrathium

and one in Lathrobium, are moved to Tetartopeus. One

previous subspecies from Japan, L. varium, is elevated to

species.

Di- and polymorphisms. Some species such as the West

Palaearctic L. multipunctum are subject to remarkable in-

traspecific (polymorphic) variation of coloration, body

size, eye size, elytral length, the length of the hind wings,

and other characters.

Conspicuous dimorphisms were observed in several

species from the East Palaearctic. Lobrathium wittmeri

from Nepal, as well as L. bimembre and L. duplex from

China, are represented by two distinct morphs, a

macropterous morph with long elytra and fully developed

hind wings and a micropterous morph with short elytra,

weakly marked humeral angles, and with reduced hind

wings. The available evidence suggests that these dimor-

phisms are not sex-related. Another example of a wing di-

morphism may be L. domenoides, but the species is cur-

rently represented only by a single micropterous male.

Remarkably, several of the Himalayan species are di-

morphic regarding the presence or absence of elytral spots.

This intraspecific variation is apparently not clinal; on

some occasions both morphs were found together. The

same was not observed for the spotted Lobrathium species

from China, though the size of the elytral spot may be sub-

ject to considerable variation, particularly so in L.

hongkongense.

OZFMK

Revision of East Palaearctic Lobrathium 53

Natural history. East Palaearctic Lobrathium species have

been found in various habitats. Many, usually more wide-

spread species seem to primarily inhabit lakeshores and

river banks, whereas other species, most of them with re-

stricted distributions, were predominantly collected from

the leaf litter of forests.

Unlike the vast majority of East Palaearctic Lathrobi-

um species, which, for instance in the Himalaya, are con-

fined to high altitudes of usually at least 2500 m up to ap-

proximately 5000 m, East Palaearctic Lobrathium species

have been recorded also from lower elevations. In the Hi-

malaya, they were collected at altitudes of 450-3100 m,

most of them between 1000 and 2800 m. In China, the al-

titudes range from 150 to 3350 m; one species from Ti-

bet, L. cholaicum, was even found at 4400 m. In general,

macropterous species inhabiting lakeshores and river

banks are found at lower elevations than micropterous and

wing-dimorphic species living in forest litter. Lobrathium

alticola, L. wittmeri, L. daxuense, L. bimembre, L. lamel-

latum, and L. duplex, for instance, were found at higher

altitudes of 2500-3500 m.

Teneral adults of East Palaearctic Lobrathium species

were collected during the period from February through

September, with most observations from March through

July, suggesting that the pre-imaginal development of

some species occurs during the cold season. However, the

data vary between species.

ADDITIONAL RECORDS

PALAEARCTIC REGION

FROM THE WEST

Lobrathium multipunctum (Gravenhorst, 1802)

Lathrobium striatopunctatum Motschulsky 1858: 646, preoccu-

pied.

Lathrobium differens Gemminger & Harold 1868: 610, replace-

ment name.

Material examined. Spain: | ex., Navarra, Sierra de Andia, Pto.

de Lizzarraga, 1050 m, 15.V1.2006, leg. Anichtchenko (cAss);

| ex. [pale-coloured, brachypterous], Soria, Santa Cruz de Yan-

guas, Sierra del Hayedo de Santiago, Ayo. de las Monjas,

42.09°N, 2.50°W, 1450 m, 16. VIII.2008, leg. Andtjar & Arri-

bas (cAss); 1 ex. [pale-coloured, brachypterous], Soria, La Po-

veda de Soria, Pto. de Piqueras, 42°06°N, 2.54°W, 1710 m,

16. VUI.2008, leg. Andujar & Arribas (cAss); | ex. [pale-colou-

red, brachypterous], La Rioja, Villoslada de Cameros, Sierra Ce-

bollera, Lomos de Orios, 42.04°N, 2.68°W, 1400 m,

17. VIIL.2008, leg. Andtyjar & Arribas (cAss); | ex. [teneral], Ma-

drid, Sierra de Guadarrama, Navacerrada, 1800 m, 5.1X.2001,

leg. Anichtchenko (cSha); 1 ex., Castilla-La Mancha, Fuenca-

liente (CR), Sierra Madrona, Rio Cereceda, 1.X.2005, leg. Len-

cina & Andtjar (cAss); 1 ex., Valencia, Alicante, E Planes, Bco.

de la Encantada, 38°48’N, 00°18’ W, 480 m, 3. VIII.2008, leg For-

cke (cAss); 2 exs., Valencia, Alicante, NW El Castell de Gua-

dalest, Beniarda, 38°41’N, 00°13’W, 400 m, river bank,

9. VIII.2008, leg Forcke (cAss); 1 ex., Valencia, Alicante, NE Be-

nissa, 38°45’N, 00°03’W, 400 m, 10.VIH.2008, leg Forcke

(cAss).

Italy: Basilicata: 2 exs., Pignola ris. WWF, Lago Pignola (PZ),

11.VII.1992, leg. Angelini (NHMD, cAss); 2 exs., Policoro

(MT), 3.XI.2000, leg. Angelini (NHMD, cAss). Calabria: 5 exs.,

Belvedere Mar., S.S. 18 at Sangineto stream (CS), 15.1V.1994,

leg. Angelini (NHMD, cAss); 3 exs., Lao river, 4 km from estua-

ry (CS), 17.VI.1994, leg. Angelini (NHMD, cAss); 4 exs., San-

ta Maria Cedro, Abatemarco river (CS), 17.VI.1994, leg. Ange-

lini (NHMD); | ex., Crotone, Esaro river, 26.X.2003, leg. An-

gelini (cAss).

Catalogue of the Lobrathium species of the Palaearctic region

In the references column, only a selection of relatively recent articles providing descriptions, illustrations, significant records, or

containing important nomenclatural changes is listed. The references are abbreviated as follows:

App = Assing (present paper); A04 = Assing (2004); AOS = Assing (2005); A06a = Assing (2006a); AO7 = Assing (2007); A08 =

Assing (2008); A10 = Assing (2010); Alla = Assing (201 1a); Al2 = Assing (2012); ASO2 = Assing & Schiilke (2002); BO9 = Bor-

doni (2009); Ca21 = Cameron (1921); Ca24 = Cameron (1924); Ca31 = Cameron (1931); Ca43 = Cameron (1943); Co67 = Coif-

fait (1967); Co79 = Coiffait (1979); Co82a = Coiffait (1982a); Co82b = Coiffait (1982b); Co82c = Coiffait (1982c); CZ00 = Ci-

ceroni & Zanetti (2000); G93 = Gusarov (1993); G95 = Gusarov (1995); 195 = Ito (1995); 196a = Ito (1996a); 196b = Ito (1996b);

107 = Ito (2007); 109b = Ito (2009b); Kh39 = Koch (1939); Ko71 = Korge (1971); N36 = Normand (1936); S01 = Solodovnikov

(2001); ShO8 = Shavrin (2008); W72 = Watanabe (1972); W98b = Watanabe (1998b); WB73 = Watanabe & Baba (1973); Z88 =

Zheng (1988).

Valid names are sorted alphabetically, synonyms by publication year.

Lobrathium Mulsant & Rey, 1878 (type species Lathrobium multipunctum Gravenhorst, 1802)

= Lathrobiella Casey, 1905 (type species Lathrobium collare Erichson, 1840)

= Lathrotaxis Casey, 1905 (type species Lathrobium longiusculum Gravenhorst, 1802)

= Ponthrobium Korge, 1971 (subgenus; type species Lathrobium heinzi Korge, 1971)

= Allobrathium Coiffait, 1972 (type species Lathrobium lethierryi Reiche, 1872)

= Paralobrathium Bordoni, 1999 (type species Lathrobium apicale Baudi di Selve, 1857)

Bonn zoological Bulletin 61 (1): 49-128

©OZFMK

54 Volker Assing

taxon revised distribution references

ablectum sp. n. China (Hubei) App

alaiense Gusarov, 1995 Kyrgyzstan G95

alticola (Cameron, 1943) Kashmir App, Ca43, Co82a

= pirpanjalense Coiffait, 1982; syn. n.

amamiense Ito, 1996a

anale (Lucas, 1846)

= concinnum (Gougelet & Brisout, 1860)

= canariense (Wollaston, 1865)

= oviceps (Fauvel, 1902)

= lostiae (Dodero, 1916)

= rufiventre Coiffait, 1953

= bellesi Bordoni, 1977

= ullbrichi Coiffait, 1978

= rubriventre Herman, 2003

angelinii Ciceroni & Zanetti, 2000

angulatum Assing, 2005

apicale (Baudi di Selve, 1857)

= cypriacum Jarrige, 1949

apogeum (Normand, 1936)

badium (Cameron, 1924)

bettae (Solodovnikov, 2001)

biaculeatum sp. n.

bicarinatum sp. n.

bicornutum sp. n.

bidigitatum Assing, 2010

bilobatum Assing, 2010

bimembre sp. n.

bipeniculatum Assing, 2010

bisagittatum Assing, 2010

bispinosum sp. n.

brunneum (Cameron, 1931)

bureschi (Scheerpeltz, 1937)

candicum Bordoni, 2009

cholaicum sp. n.

ciliciae Bordon, 1980

coalitum Assing, 2010

configens sp. Nn.

cornutum sp. n.

cornutissimum Assing, 2010

cribricolle (Sharp, 1889)

cylindricolle (Cameron, 1924)

daxuense sp. n.

demptum sp. n.

diecki (Saulcy, 1878)

digitatum Assing, 2010

discrepans sp. n.

domenoides sp. n.

duplehamatum Assing, 2010

duplex sp. n.

emeiense Zheng, 1988

excisissimum sp. 0.

extensum Assing, 2010

farsicum Assing, 2007

feldmanni sp. n.

flavipenne sp. n.

frater (Korge, 1971)

Japan (Amami Islands)

NW-Africa; SW-Europe; Canary Islands

S-Italy

Greece

Cyprus

Algeria

N-India (Uttaranchal, Himachal Pradesh)

W-Caucasus

E-Nepal

N-India (Uttaranchal); Nepal

E-Nepal

Taiwan

China (Yunnan)

Taiwan

China (Guizhou)

N-India: Sikkim

Romania, Bulgaria

Greece (Crete)

China (Tibet)

S-Turkey

Taiwan

China (Shaanxi, Sichuan, Hubei, Yunnan)

E-Nepal

Taiwan

Japan (Honshu)

N-India (Uttaranchal)

China (Sichuan)

China (Hube1)

S-Italy (Sicily)

Taiwan

N-Vietnam

China (Sichuan)

Taiwan

China (Sichuan, Yunnan)

China (Sichuan)

China (Yunnan)

Taiwan

SW-Iran

China (Sichuan)

N-India (Himachal Pradesh)

NE-Turkey

App, [96a

A07, Co82c

A07, CZ00

AOS

A07, ASO2, Co82c

N36

App, Ca24, Ca31

S01

App

Al0

A10

App

Al0

Al10

App

App, Ca31

AS, Co82c

App, B09

App

A04, AS02

Al0

App

Al0

App, [96a

App, Ca24, Ca31

App

Co82c

A10

App

Al0

App

App, Z88

App

A10

A07

App

Co82c, Ko71

Bonn zoological Bulletin 61 (1): 49-128

©ZFMK

Revision of East Palaearctic Lobrathium

taxon revised distribution references

furcillatum Assing, 2010 Taiwan Al0

gladiatum Zheng, 1988 China (Sichuan) App, Z88

guttula (Fauvel, 1895); comb. n. Myanmar App

hebeatum Zheng, 1988 China (Sichuan, Shaanxi, Yunnan) App, Z88

heinzi (Korge, 1971) NE-Turkey A07, Co82c, Ko71

hokkaidense Ito, 1996 Japan (Hokkaido) 196b

hongkongense (Bernhauer, 1931)

= sibynium Zheng, 1988; syn. n.

= ryukyuense Ito, 1996; syn. n.

indubium (Eppelsheim, 1893)

= altaicum (Coiffait, 1967)

integrum sp. N.

isamutanakai Ito, 2009

ishidai Ito, 1996

ishizuchiense Ito, 1996

kirgisicum Assing, 2007

kleebergi sp. n.

kosiense sp. n.

kuanicum Assing, 2010

lamellatum sp. n.

lederi (Eppelsheim, 1884)

lethierryi (Reiche, 1872)

mizunoi Ito, 1996

mordens sp. n.

multipunctum (Gravenhorst, 1802)

= testaceum Paykull, 1789 (nom. obl.)

= lineare (Gravenhorst, 1802)

= punctatostriatum (Stephens, 1833)

= striatopunctatum (Kiesenwetter, 1850)

= striatopunctatum (Motschulsky, 1858)

= pyrenaicum (Fairmaire, 1863)

= differens (Gemminger & Harold, 1868)

= hispanicum (Dodero, 1916)

= gallienii (Fagniez, 1917)

= endogeum Coiffait, 1971

= cassolai Coiffait, 1972

nigripenne Assing, 2010

nipponense Ito, 1995

novum (Bernhauer & Schubert, 1912)

= cognatum (Eppelsheim, 1892), preocc.

nudum (Sharp, 1874)

ochreonotatum (Champion, 1922); revalid.

ohkurai Ito, 1996

okamotoi Ito, 1995

partitum (Sharp, 1874)

pedes Assing, 2010

penicillatum Assing, 2010

pravum Assing & Schiilke, 2002

pustulatum (Cameron, 1931); comb. n.

radens sp. n.

reitteri (Czwalina, 1889)

retrocarinatum sp. Nn.

reuteri Assing, 2008

riozoi Watanabe, 1972

rotundiceps (Koch, 1939)

China; Taiwan; S-Japan (Ryukyu Islands)

East Siberia (Baikal region), Russian Far East

NE-Nepal

Japan (Honshu)

Japan (Shikoku)

Kyrgyzstan

Nepal

E-Nepal

Taiwan

China (SW-Sichuan)

Azerbaijan, Russian South European territory, Iran

Italy (Sicily), Algeria, Tunisia

Japan (Honshu)

N-India (Uttaranchal, Himachal Pradesh)

Europe; NW-Africa; Nearctic region (adventive)

Taiwan

Japan (Kyushu, Honshu)

Tajikistan, Uzbekistan

Japan (Kyushu, Honshu, Ryukyu Islands)

N-India; Nepal

Japan (Honshu)

Japan (Shikoku)

Japan (Kyushu, Honshu)

Taiwan

E-Turkey, Iraq, Iran

N-India (Assam)

China (Guizhou)

W-Caucasus

China (Yunnan)

Traq

Japan (Hokkaido)

China (Zhejiang)

A10, App, 196b, Z88

App, C067, Co82c, Sh08

App

109b

196b

196a

A07

App

Al0

App

App, Alla, S01

Co82c

196b

App

A07, A08, A12, Co82c, CZ00

Al0

195

AS02, G95

App, 196a

App

196b

195

App

Al0

A07, Alla, ASO2

App, Ca31

App

A07, SO1

App

A08

W72

App, K39

Bonn zoological Bulletin 61 (1): 49-128

©OZFMK

56 Volker Assing

taxon revised distribution references

rugipenne (Hochhuth, 1851) SE-Europe; Turkey; Caucasus A07, ASO02, Co82c, G93

= meridionale (Korge, 1971)

= vicinum Coiffait, 1972

= messeniacum Bordoni, 1986

sahlbergi (Fauvel, 1900) Kyrgyzstan, Kazakhstan Co82c, G95

sasajii Ito, 2007 Japan (Honshu) 107

schillhammeri Assing & Schilke, 2002 SE-Turkey AS02

schuelkei sp. n. China (Shaanx1) App

semicaeruleum (Cameron, 1921) N-India; E-Nepal App, Ca21, Ca31

semiflavum sp. n. Russian Far East App

shibatai Ito, 1995 Japan (Honshu) 195

sinuatum sp. n. C-Nepal App

smetanai Assing, 2010 Taiwan Al0

solarii (Koch, 1936) S-Italy Kh36, Co82c

sororium Assing, 2010 Taiwan Al0

spathulatum sp. n. China (Sichuan, Shaanxi, Shanxi, Hubei, Zhejiang) App

spinosum Assing & Schiilke, 2002 Albania AS02

spoliatum Assing, 2010 Taiwan Al0

stimulans Assing, 2010 Taiwan Al10

taiwanense (Watanabe, 1998) Taiwan W98b

taureum sp. N. China (Hubei, Beijing) App

tortile Zheng, 1988 China (Sichuan, Shaanxi, Hube1) App, Z88

trapezuntis (Bordoni, 1973) NE-Turkey B73, Co82c

triste (Cameron, 1924) N-India (Uttaranchal, Kashmir), Afghanistan, Pakistan App, Ca24, Ca31, Co79

= kashmiricum (Cameron, 1931); syn. n.

= afghanicum Coiffait, 1979; syn. n.

= nouristanicum Coiffait, 1979; syn. n.

tuberosum sp. n. China (Jiangxi) App

unispinosum sp. Nn. Nepal App

varium Ito, 1995, stat. n. Japan (Honshu) 195

wittmeri Corffait, 1982 C-Nepal App, Co82b

wunderlei Assing, 2006 S-Turkey A06a, A07

vagmuri Assing, 2007 SE-Turkey A07

yoshidai sadoensis Watanabe & Baba, 1973 Japan (Sado Island) WB73

yoshidai yoshidai Adachi, 1955 Japan (Honshu) WB73

Yugoslavia: 3 exs., Serbia, Suva planina, Zorebica, 1350 m, Lobrathium rugipenne (Hochhuth, 1851)

5.V.2006, leg. Stevanovic (cAss).

Material examined: Greece: 3 exs., Chalkidiki, Sithonia, Sar-

Comment. When Lathrobium differens Gemminger & ti, 10 m, 40°05’N, 23°59’E, [X.2007, leg. Frisch (MNHUB,

Harold, 1868, a replacement name for the preoccupied L. —_cAss); 1 ex., Trikala, Pindos, Hinka, 1500 m, 23.V.2005, leg. An-

striatopunctatum Motschulsky, 1858, was placed in syn- __ gelini (NHMD).

onymy with L. multipunctum by Assing (2008), the

Motschulsky name was erroneously given as L. multistria- | Comment. This species is distributed from the southern

tum Motschulsky, 1858 (unavailable). This lapsus is rec- | Balkans to the Caucasus region.

tified here.

Bonn zoological Bulletin 61 (1): 49-128 ©ZFMK

Revision of East Palaearctic Lobrathium 57

Lobrathium anale (Lucas, 1846)

Material examined. Spain: 2 exs., Pais Vasco, Guipuzcos, Irun,

Plaiaundi, 9.1V.2006, leg. Anichtchenko (cSha); 1 ex., Andalu-

cia, Granada, Rio Darro, barranco de Teatino, 6.1I.2005, leg.

Anichtchenko (cSha); 1 ex., Andalucia, Cordoba, Charca, Ca-

stillo de la Albaida, 15.V.2006, leg. Baena (cAss); 2 exs., An-

dalucia, Cadiz, 25 km NW Tarifa, Tahivilla, 36°11°N, 5°45’ W,

5 m, flooded fallow, under stone, 26.XII.2009, leg. Assing &

Wunderle (cAss, cWun); | ex., Andalucia, Cadiz, 15 km NW

Algeciras, 36°13’N, 5°33’ W, 25 m, loamy pasture, under stone,

27.XI1I.2009, leg. Assing (cAss).

Morocco: 9 exs., N Touya, Ifri cave, 23.1V.1997, leg. Casale

(cGia, cAss).

Comment. Lobrathium anale is widespread in the west-

ern Mediterranean and the Canary Islands.

Lobrathium lederi (Eppelsheim, 1884) (Figs 1-4)

Lathrobium lederi Eppelsheim, 1884: 15 f.

Type material examined. Lectotype @, present designa-

tion: “Casp1.-M.-Gebiet, Rasano. Leder (Reitter) / Lederi

Epp, Verh. naturf. Ver. Briinn, Bd XXII. / Typus / c. Ep-

plsh. Steind. d. / Lectotypus ¢ Lathrobium lederi Eppels-

heim, desig. V. Assing 2011 / Lobrathtum lederi (Ep-

pelsheim), det. V. Assing 2011” (NHMW). Paralectotypes:

1¢: “Caspi.-M.-Gebiet, Liryk. Leder (Reitter) / 1881.1 /

Typus” (NHMW); 1: “4 / Kaukas Leder, Berge von Ta-

lysch. Lirik / Typus / c. Epplsh. Steind. d.” (NHMW); 19:

“Caspi.-M.-Gebiet, Liryk. Leder (Reitter) / Typus / c.

Epplsh. Steind. d.” (NHMW).

Comment. The original description is based on an unspec-

ified number of syntypes from “Gebirge von Talysch bei

Lirik und Rasano” (Eppelsheim 1884). Four syntypes were

located in the Eppelsheim collection at the NHMW. A

male in good condition is designated as the lectotype. The

male primary and secondary sexual characters of the lec-

totype are illustrated in Figs 1-4.

Lobrathium candicum Bordoni, 2009

Type material examined. Holotype < [teneral]: ““Kreta,

Kritsi, 13.5.75 / Holotypus Lobrathium candicum n. sp.,

Bordoni det. 2006 / ZML 2010, 168 / Lobrathium candi-

cum Bordoni, det. V. Assing 2010” (ZML).

Additional material examined. Greece: | ¢ [teneral], W-Crete,

bank of Litheos river near Agii Deka, 200 m, 12.V.2001, leg.

Apfel (cAss).

Comment. In external and the male sexual characters, in-

cluding the internal structures of the aedeagus, this species

is highly similar to L. apicale, from which it is distin-

Bonn zoological Bulletin 61 (1): 49-128

guished only by the slightly different shape of the apex

of the ventral process of the aedeagus. In L. candicum, it

is more strongly bent dorsad and has more pronounced lat-

eral carinae and subapical teeth (best visible in lateral

view).

THE LOBRATHIUM SPECIES OF THE HIMALAYA

AND MYANMAR

Key to species

1. Pronotum conspicuously oblong, approximately 1.45

times as long as broad (Figs 119-120). Elytra with-

out submarginal line (lateral view! ). Forebody reddish.

4: sternite VII and VIII as in Figs 121—122; aedea-

gus 0.8 mm long and with apically bifid ventral

process (Figs 123-124). Northern India: Uttaranchal

(EIS IOG) eS Lake ere 2: cylindricolle (Cameron)

— Pronotum much less oblong, at most approximately

1.3 times as long as broad. Elytra with submarginal

line (lateral view!). Male sexual characters different.

Se i ee ee eer a

2. Large species; length of forebody at least approxi-

mately 5.0 mm. Head and pronotum dark-brown.

Pronotum at least approximately 1.25 times as long

as broad. ¢: ventral process of aedeagus divided in-

LO MEWONLOMEAPLOCESSESPen-cea-eee ee eeee eee cee eee cee eee 3

— Small to moderately large species; length of forebody

2.9-4.9 mm. (One species, L. badium, whose fore-

body may be more than 4.6 mm long, has the head

and pronotum bright reddish and a distinctly less ob-

long pronotum.) @: ventral process sometimes apical-

ly bifid, but not divided into two long processes ...5

3. Largest Himalayan representative of the genus;

length of forebody approximately 7 mm. Macropter-

ous: elytra distinctly longer than pronotum (Fig. 12);

hind wings fully developed. Posterior margin of ter-

gite VII with palisade fringe. @’: sternite VIII as in Fig.

14; aedeagus 2.1 mm long and shaped as in Figs

15—16. Northern India: Sikkim (Fig. 23)

Si ep paearee, NETEID fthe «ogo § lg brunneum (Cameron)

— §Smaller species; length of forebody less than 6 mm.

Micropterous: elytra much shorter, 0.70—0.75 times

as long as pronotum (Fig. 18); hind wings reduced.

Posterior margin of tergite VII without palisade fringe.

G: sexual characters different ....................0e0008 4

4. ©: sternite VII more transverse (Fig. 19); sternite

VII less oblong and with larger median impression

(Fig. 20); aedeagus 2.4 mm long, with longer and

more slender bifid ventral process (Figs 21—22). East-

ern Nepal: Taplejung (Fig. 23) ...bicornutum sp. n.

— 4: sternite VII less transverse (Fig. 24); sternite

VIII more oblong and with smaller median impres-

©OZFMK

Volker Assing

sion (Figs 25—26); aedeagus 1.8 mm long, with short-

er and stouter bifid ventral process (Figs 27-28). East-

ern Nepal: Terhathum (Fig. 23) ....... cornutum sp. n.

On average larger species; length of forebody 4.2-4.9

mm. Head and pronotum bright reddish, distinctly

contrasting with the dark-brown to black elytra.

Pronotum broad, approximately 1.15 times as long as

broad. @: sternites VII and VIII as in Figs 7-8; aedea-

gus approximately 1.3 mm long, with large blade-

shaped ventral process (Figs 9-10). Northern India:

Uttaranchal and Himachal Pradesh (Fig. 23)

PRE AcraN Teo a ane ne oeae uuu sad Meee: badium (Cameron)

On average smaller species; length of forebody

2.9-4.8 mm. Head and pronotum blackish, sometimes

with bluish hue. Pronotum in the vast majority of

species at least 1.20 times as long as broad. Male sex-

wal\characters ditterent otis. asan. meceeee eee ere 6

Posterior margin of tergite VII without palisade fringe.

Elytra short, approximately 0.75 times as long as

pronotum (Figs 97, 103); hind wings reduced. Small

species; length of forebody approximately 3.5 mm at

Posterior margin of tergite VII with palisade fringe.

Elytra longer and hind wings fully developed (except

in micropterous morph of L. wittmeri). Mostly larg-

CPISPECICS (iv Sore a see Serer Ree oes 8

Eyes small, approximately 1/4 the length of postoc-

ular region (Fig. 97). Length of forebody approxi-

mately 2.9 mm. @: sternite VII highly distinctive, with

pronounced median impression posteriorly, this im-

pression without pubescence, except for two isolat-

ed posterior setae, but margined with dense black se-

tae, anteriorly with short process (Fig. 98); sternite

VII as in Fig. 99; aedeagus 0.85 mm long and shaped

as in Figs 100-101. Kashmir (Fig. 66)

Scene HEATH LOSSES ae ete alticola (Cameron)

Eyes larger, more than half the distance from poste-

rior margin of eye to neck in dorsal view (Fig. 103).

Length of forebody approximately 3.5 mm. <: ster-

nite VII without such conspicuous modifications (Fig.

104); sternite VII as in Fig. 105; aedeagus 0.95 mm

long and shaped as in Figs 106-107. Eastern Nepal:

Kosn(Figs39) ie peso eee kosiense sp. n.

Wing-dimorphic: usually micropterous with elytra

0.70—0.75 times as long as pronotum (Fig. 90) and

with reduced hind wings, very rarely macropterous

(Fig. 91). Body small; length of forebody 3.0-3.5 mm.

Body uniformly blackish; forebody without bluish hue

and elytra always without posterior spot. 4: sternites

VII and VIII as in Figs 92-93; aedeagus approximate-

ly 1.0 mm long and shaped as in Figs 94-95. Central

INepalk (Bigs 4ulb)\.< an. Pee ee wittmeri Corffait

Monomorphic, macropterous species; elytra at least

0.9 times as long as pronotum. Body mostly larger.

Forebody often with bluish hue and elytra often with

Bonn zoological Bulletin 61 (1): 49-128

10.

13%

yellowish to reddish spots posteriorly. Male sexual

TELS KARI OMGKACH NG cagnaacacds6d09029600000000000009000000 9

Forebody brown to black, without bluish hue. Prono-

tum often less than 1.25 times as long as broad. Ely-

tra in some species shorter than, or as long as elytra.

Length of forebody sometimes greater than 4.0 mm.

: sternite VII with or without cluster(s) of short stout

MOIS SCtAG «cs crane pieosapent rae Pepe 10

Forebody black, usually with bluish hue (rare excep-

tions without bluish hue may be identified only based

on the male sexual characters). Pronotum at least 1.25

times as long as broad. Elytra longer than pronotum.

Length of forebody 3.0-4.0 mm. @: sternite VII nev-

er with short stout modified setae ..................0 7

Forebody uniformly blackish, without elytral spot,

rarely with indistinctly paler posterior margin ....... 11

Forebody brown to black; elytra with elytral spot pos-

1-3) 0 (0) dh Aen seers aa Sanaa cAGGauEm Sonu gdoso0dc00000 13

Pronotum broad, approximately 1.15 times as long as

broad (Fig. 114). Body on average larger; length of

forebody 4.2-4.8 mm. @: sternite VII with two clus-

ters of short stout modified setae (Fig. 115); sternite

VIII with extensive and deep impression and with pair

of distinct carinae (Fig. 116); aedeagus 1.5—1.6 mm

long and shaped as in Figs 117—118. Northern India:

Uttaranchal; central Nepal (Fig. 89)

tle SAEERL ERR beo eee bicarinatum sp. n.

Pronotum at least approximately 1.20 times as long

as broad. Body on average smaller; length of forebody

3.6-4.3 mm. <: sternite VII without distinctly mod-

ified setae; sternite VIII with less pronounced impres-

sion and without carinae; aedeagus of different shape

Coloration of forebody black. On average smaller

species; length of forebody 3.6-3.9 mm. <: sternite

VII strongly modified, with two long processes pos-

teriorly (Fig. 109); sternite VHI with relatively deep

and broad posterior excision (Fig. 110); aedeagus ap-

proximately 0.95 mm long and shaped as in Figs

106-107. Eastern Nepal: Kosi (Fig. 89)

Per een oe a adG adit co0d biaculeatum sp. n.

Coloration of forebody brown to black. On average

larger species; length of forebody 3.8-4.3 mm. <:

sternite VII without conspicuous modifications; ster-

nite VII with even deeper posterior excision (Figs

32-33); aedeagus approximately 1. 3 mm long and

with apically spear-shaped ventral process (Figs

34-35). Widespread from northern Afghanistan to

northern India (Fig. 23)) ..4..2..2...-08 triste (Cameron)

Head, except for median dorsal portion, with conspic-

uously dense punctation. @: sternite VII with distinct-

ly modified, short and stout black setae (Figs 69, 73,

79); posterior excision of sternite VIII of subtrapezoid

shape, basally often bisinuate (Figs 70, 74, 80); aedea-

gus with blade-shaped ventral process .............. 14

Revision of East Palaearctic Lobrathium 59

— Head with less dense punctation dorsally. 4: sternite

VII without distinctly modified setae; posterior exci-

sion of sternite VIII not of subtrapezoid shape; aedea-

SusonGitterentimorpholosyaerceeecee--cetkeee ee cece 16

14. @: sternite VII as in Fig. 69; sternite VIII more trans-

verse and with convex lateral margins, its posterior

excision not distinctly bisinuate basally (Fig. 70);

aedeagus approximately |.1 mm long and shaped as

in Figs 71—72. Nepal (Fig. 41) .........4leebergi sp. n.

— <4: sternite VII of different shape and chaetotaxy; ster-

nite VIII not distinctly transverse and with concave

lateral margins, its posterior excision distinctly

bisinuate basally (Figs 74, 80); aedeagus of different

GINGTOS cco casonGo ues uaEe Doo sor aan ane denOR Aeon decane 15

15. 4: sternites VII and VIII as in Figs 79-80; aedeagus

as in Figs 81—82. Central Nepal (Fig. 90)

Rae eee eines os Sone neeinaatie eiisicntes #2 sinuatum sp. n.

— 6: sternites VII and VIII as in Figs 73-74; aedeagus

as in Figs 75-76. Myanmar (Fig. 140)

oo nd Dn SOS aN EH AR ECR PR AIE ANE Soe E Eh Me aeame guttula (Fauvel)

16. Elytra with large and clear-cut elytral spot (Fig. 84).

S: sternites VII and VIII as in Figs 85-86; aedeagus

1.5 mm long and with long, slender, and apically very

acute ventral process (Figs 87-88). Northern India:

/NGeeVIN (tis; 2) Cooodevenconden pustulatum (Cameron)

— Elytra with indistinct and ill-defined elytral spot at

posterior margin (Fig. 108). @: sternite VIII as in Fig.

32; aedeagus approximately 1.3 mm long and with

apically spear-shaped ventral process (Figs 34-35).

Widespread from northern Afghanistan to northern In-

GUE), a ntuceemioe bseue SeGSa OORT EO CAREC ete triste (Cameron)

17. Posterior 2/3 of the elytra yellowish, anterior 1/3

blackish (Fig. 62). ¢: sternite VII with bidomed tu-

bercle posteriorly (Fig. 63); sternite VIII with deep

posterior excision (Fig. 67); aedeagus 1.1 mm long

and with apically bifid ventral process (Figs 64-65).

Northern India: Himachal Pradesh (Fig. 41)

PPR Eee ce aicicieceiaie sess saesiiae fLAVIPENNEISP. 11.

— Elytra with much smaller yellowish to reddish spot

posteriorly or uniformly blackish with bluish hue.

Male sexualicharacters different....22...0.5.--------- 18

18. 4: ventral process of aedeagus asymmetric. Speci-

mens with spotted elytra unknown .................. 19

— 4: ventral process of aedeagus symmetric. Elytra with

or without yellowish to reddish spots ............... 20

19. 3: sternite VII as in Fig. 48; sternite VIII with very

small posterior excision (Fig. 49); aedeagus 1.2—1.3

mm long, ventral process stout, on right ventral side

(ventral view) with pronounced tooth-shaped process

(Figs 50-51). Nepal (Fig. 23) .....unispinosum sp. n.

— : sternite VII as in Fig. 54; sternite VIII with pro-

found posterior excision (Fig. 55); aedeagus 1.4 mm

long and with slender ventral process without tooth

(Figs 56-57). Northeastern Nepal: Taplejung

a Conte No a oar aa Saini Sees at ere integrum sp. n.

Bonn zoological Bulletin 61 (1): 49-128

20. ¢: aedeagus 1.2—1.3 mm long, ventral process very

slender, sting-shaped in ventral view (Figs 44—45);

sternite VII as in Fig. 42; sternite VII with broad pos-

terior excision of almost semi-circular shape (Fig. 43).

Northern India: Uttaranchal, Himachal Pradesh (Fig.

ANI): sed duasbooao do Ae te eee TORE oe ert mordens sp. n.

— ©: aedeagus shorter, ventral process shorter and

stouter; sternite VII of different shape .............. 21

21 4: aedeagus approximately 1.0 mm long, ventral

process apically bifid (Figs 39-40); sternite VIII with

deep and U-shaped posterior excision (Fig. 37). Ely-

tra usually of uniformly black coloration, rarely with

posterior spots. Northern India, eastern Nepal (Fig.

AI) eee aero necsas: semicaeruleum (Cameron)

— ©: aedeagus approximately 0.9 mm long, ventral

process apically not bifid (Figs 60-61); sternite VIII

with less deep and broader posterior excision (Fig.

59). Elytra usually with, rarely without posterior spots.

Northern India, Nepal (Fig. 66)

NN Thy ener Sele ve scisiees ochreonotatum (Champion)

Lobrathium badium (Cameron, 1924) (Figs 5—10, 23)

Lathrobium (Lobrathium) badium Cameron, 1924: 193 f.

Type material examined. Lectotype <, present designa-

tion: “R. Song, Dehra Dun. / Dr. Cameron. 2.1V.1922. /

Syntype / M.Cameron. Bequest. B.M. 1955-147. / Syn-

type Lathrobium badium Cam., det. R.G. Booth 2011 /

Lectotype 3 Lathrobium badium Cameron, desig. V. Ass-

ing 2011 / Lobrathium badium (Cameron), det. V. Assing

2011” (BMNH). Paralectotype 4 [aedeagus missing]:

“Dhobi Ghat, Mussoorie. Dr. Cameron. 14.IV.22. / Lath-

robium badium Cam. / Type / M.Cameron. Bequest. B.M.

1955-147.” (BMNH).

Comment. The original description is based on an unspec-

ified number of syntypes, among them at least one male,

from “Mussoorie District; Dhobi Ghat, Keyarkuli, 6000

feet above the sea. Dehra Dun District; Nim Nadi, Song

River” (Cameron 1924). Two male syntypes from Dehra

Dun and Dhobi Ghat were located in the Cameron col-

lection at the BMNH. The specimen from Dehra Dun is

designated as the lectotype; no aedeagus was found in the

male from Dhobi Ghat.

Additional material examined. India: Uttaranchal: 3 exs.,

Garhwal, 16 km from Srinagar, 550 m, 29.X.1979, leg. Lobl

(MHNG, cAss); | ex., Garhwal, 20 km S Chamba, 1150 m,

20.X.1979, leg. Lobl (MHNG); 1 ex., Garhwal, between Tehri

and Srinagar, 900 m, 25.X.1979, leg. Lobl (MHNG); | ex.,

Garhwal, 22 km N Rishikesh, 450 m, 30.X.1979, leg. L6bl (MH-

NG); 2 exs., Kumaon, 10 km from Ramnagar, Garjia, 450 m,

15.X.1979, leg. Lobl (MHNG, cAss); 19, Haldwani district,

Kaldhunga, 26.III.1923, leg. Champion (cAss). Himachal

Pradesh: | 9, Mandi, Dhelu [“Dhelu, Mandi, Punjab”’], ca. 1500

m, leg. Champion (BMNH); 19, Baijnath [32°02’N, 76°38’E],

ca. 1100 m, leg. Champion (BMNH).

©ZFMK

60 Volker Assing

2 sai

Figs 1-10. Lobrathium lederi, lectotype (1-4), and L. badium (5-10; 5-6, 8: paralectotype). 1, 7: male sternite VII; 2, 8: male

sternite VIII; 3, 9: aedeagus in lateral view; 4: apex of ventral process of aedeagus in lateral view; 5: habitus; 6: forebody; 10:

aedeagus in ventral view. Scale bars: 5—6: 1.0 mm; 1—3, 7-10: 0.5 mm; 4: 0.1 mm.

Redescription. Body length 8.2—10.5 mm; length of fore-

body 4.24.9 mm. Habitus as in Fig. 5. Coloration: head

and pronotum bright-reddish to dark-reddish; elytra dark-

brown to blackish, sometimes with reddish posterior mar-

gin; abdomen reddish-brown with paler apex or reddish

with infuscate apex; legs yellowish to reddish; antennae

reddish to reddish-brown.

Bonn zoological Bulletin 61 (1): 49-128

Head (Fig. 6) as wide as long; posterior angles broad-

ly rounded, almost obsolete; punctation of lateral and pos-

terior dorsal portion very dense and moderately coarse, in-

terstices reduced to narrow ridges; median dorsal portion

and middle of frons more or less impunctate; interstices

without microsculpture. Eyes large, half as long as dis-

tance from posterior margin of eye to neck, or slightly larg-

er.

©OZFMK

Revision of East Palaearctic Lobrathium 61

Pronotum (Fig. 6) approximately 1.15 times as long as

broad and 0.9 times as wide as head; punctation similar

to that of head, but less dense.

Elytra approximately as long as, and distinctly wider

than pronotum (Fig. 6); punctation coarse and dense, not

arranged in distinct series; interstices without microsculp-

ture. Hind wings present.

Abdomen narrower than elytra; punctation fine and

dense; posterior margin of tergite VII with palisade fringe.

&: posterior margin of tergite VIII weakly convex; ster-

nite VII strongly transverse and with impunctate median

impression posteriorly, on either side of this impression

with cluster of moderately sparse black setae, posterior

margin broadly and distinctly concave (Fig. 7); sternite

VII approximately as long as broad, impressed along the

middle, this impression with numerous short modified

black setae, posterior excision relatively small, on either

side of this excision with cluster of black setae (Fig. 8);

aedeagus approximately 1.3 mm long, with large blade-

shaped ventral process (Figs 9-10).

Distribution and natural history. The known distribu-

tion of L. badium is confined to several localities in Ut-

taranchal and two localities in Himachal Pradesh, North

India (Fig. 23). The species has been collected at altitudes

of 450 to almost 2000 m.

Lobrathium brunneum (Cameron, 1931) (Figs 11—16,

23)

Lathrobium brunneum Cameron, 1931: 252.

Type material examined. Lectotype <, present designa-

tion: “Sikkim: Gopaldhara, Rungbong Vall. H. Stevens. /

Lathrobium brunneum Cam Type ¢ / Type / Lectotypus

3 Lathrobium brunneum Cameron, desig. V. Assing 2011

/ Lobrathium brunneum (Cameron), det. V. Assing 2011”

(BMNH).

Comment. The original description is based on an unspec-

ified number of syntypes, among them at least one male,

from “Sikkim: Rungbong Valley” (Cameron 1931). A male

syntype was located in the collections of the BMNH; it

is designated as the lectotype.

Redescription. Body length 11 mm; length of forebody

7 mm. Habitus as in Fig. 11. Coloration: head and prono-

tum dark-brown; elytra reddish-brown; abdomen dark-

brown with slightly paler apex; legs brown; antennae dark-

brown.

Head (Fig. 12) distinctly oblong, approximately 1.1

times as long as broad; posterior angles weakly marked,

broadly rounded; punctation dense and and not very

coarse, median dorsal portion impunctate; interstices with-

out microsculpture. Eyes large, approximately half the

Bonn zoological Bulletin 61 (1): 49-128

length of distance between posterior margin of eye to neck

in dorsal view. Antenna long and slender, 4.8 mm long.

Pronotum (Fig. 12) 1.28 times as long as broad and 0.87

times as wide as head, lateral margins straight, weakly

converging posteriorly in dorsal view; punctation similar

to that of head.

Elytra long, 1.12 times as long as pronotum (Fig. 12);

humeral angles marked; punctation coarse and dense, in-

distinctly arranged in series; interstices without mi-

crosculpture. Hind wings fully developed.

Abdomen narrower than elytra; punctation very fine and

dense; interstices with fine and shallow microsculpture;

posterior margin of tergite VII with palisade fringe.

¢: tergite VIII obtusely pointed posteriorly (Fig. 13);

sternite VII with weakly concave posterior margin; ster-

nite VII with shallow median impression posteriorly, this

impression with weakly modified pubescence, posterior

excision moderately deep, relatively narrow, and almost

V-shaped (Fig. 14); aedeagus 2.1 mm long, ventral process

conspicuously bifid, of highly characteristic shape (Figs

15-16).

Comparative notes. Lobrathium brunneum is readily dis-

tinguished from its congeners by its enormous size, its

conspicuously long antennae, and the distinctive male sex-

ual characters.

Distribution. This species is currently known only from

the type locality in Sikkim, northern India (Fig. 23).

Lathrobium bicornutum sp. n. (Figs 17-23)

Type material. Holotype ¢ [teneral; somewhat damaged;

dissected prior to present study]: “Nepal-Expeditionen

Jochen Martens / 356 Taplejung Distr., Omje Kharka NW

Yamputhin, mature mixed broad-leaved forest, 2300-2500

m, 1-6 May88 Martens & Schawaller / Lobrathium n. sp.,

det. 1990 G. de Rougemont / Holotypus ¢ Lobrathium bi-

cornutum sp. n., det. V. Assing 2011” (SMNS). Paratype

©: same data as holotype (cAss).

Etymology. The specific epithet (Latin, adjective: with

two horns) refers to the shape of the ventral process of the

aedeagus.

Description. Body length 9.5—10.3 mm; length of fore-

body 5.2—5.6 mm. Habitus as in Fig. 17. Coloration: head,

pronotum, and abdomen dark-brown to blackish-brown;

elytra dark reddish-brown; legs and antennae reddish.

Head (Fig. 18) approximately 1.1 times as long as broad;

posterior angles weakly marked, broadly rounded; punc-

tation dense and moderately coarse, frons and vertex

sparsely punctate; interstices without microsculpture. Eyes

not projecting from lateral contours of head and small, ap-

©ZFMK

62 Volker Assing

Figs 11-22. Lobrathium brunneum, lectotype (11-16), and L. bicornutum (17-22). 11, 17: habitus; 12, 18: forebody; 13: male ter-

gite VIII; 14, 20: male sternite VII; 15-16, 21-22: aedeagus in lateral and in ventral view; 19: male sternite VII. Scale bars: 11-12,

17-18: 1.0 mm; 13-16, 19-22: 0.5 mm.

proximately 0.20—0.25 times as long as distance between

posterior margin of eye to neck in dorsal view. Antenna

3.1—3.6 mm long.

Pronotum (Fig. 18) 1.25—1.30 times as long as broad and

0.90—0.95 times as wide as head, lateral margins convex

Bonn zoological Bulletin 61 (1): 49-128

in dorsal view; punctation as coarse as that of head, but

distinctly sparser; interstices glossy.

Elytra (Fig. 18) short, 0.70—0.75 times as long as prono-

tum; humeral angles weakly marked; punctation coarse

and dense, but shallow, not arranged in series; interstices

©ZFMK

Revision of East Palaearctic Lobrathium 63

Fig. 23. Distributions of Himalayan Lobrathium species, based on revised records: L. triste (open squares); L. badium (filled tri-

angles); L. unispinosum (filled diamonds); L. cornutum (open diamond); L. integrum (open triangle); L. bicornutum (filled square);

L. brunneum (open circle).

without microsculpture. Hind wings reduced. Protarsi

without sexual dimorphism, moderately dilated in both

Sexes.

Abdomen broader than elytra; punctation very fine and

dense, slightly less dense on posterior than on anterior ter-

gites; interstices with fine and shallow, or almost obso-

lete microsculpture; posterior margin of tergite VII with-

out palisade fringe; posterior margin of tergite VIII con-

vex.

S: sternite VII with shallow median impression poste-

riorly, with sparse unmodified pubescence, and with weak-

ly concave posterior margin (Fig. 19); sternite VII with

narrow median impression posteriorly, this impression

with distinctly modified, short and stout black setae, pos-

terior excision moderately deep, relatively narrow, and al-

Bonn zoological Bulletin 61 (1): 49-128

most V-shaped (Fig. 20); aedeagus 2.4 mm long, ventral

process deeply bifid, of characteristic shape (Figs 21—22).

Comparative notes. Lobrathium bicornutum 1s readily

distinguished from the similarly large L. brunneum by the

much smaller eyes, much shorter elytra, reduced hind

wings, the absence of a palisade fringe at the posterior

margin of the abdominal tergite VIII, and by the male sex-

ual characters.

Distribution and natural history. The type locality is sit-

uated in Taplejung district, eastern Nepal (Fig. 23). The

specimens were collected in a mature mixed deciduous

forest at an altitude of 2300-2500 m. The holotype is ten-

eral.

OZFMK

64 Volker Assing

Figs 24-40. Lobrathium cornutum (24-28), L. triste, lectotype (29-35), and L. semicaeruleum, lectotype (36-40). 24: male ster-

nite VII; 25, 32, 37: male sternite VIII; 26, 33, 38: posterior median portion of male sternite VIII; 27-28, 34-35, 39-40: aedea-

gus in lateral and in ventral view; 29: habitus; 30, 36: forebody; 31: tergite VII. Scale bars: 29-30, 36: 1.0 mm; 24—25, 27-28,

31-32, 34-35, 37, 39-40: 0.5 mm; 26, 33, 38: 0.1 mm.

Bonn zoological Bulletin 61 (1): 49-128 ©ZFMK

Revision of East Palaearctic Lobrathium 65

Lathrobium cornutum sp. n. (Figs 23-28)

Type material. Holotype ¢ [teneral; somewhat damaged;

dissected prior to present study]: “Nepal-Expeditionen

Jochen Martens / 2978 Terhathum Dist., Tinjura Dara,

2450-2850 m, artenreicher Laubmischwald, 17 Sep 83

Martens & Daams |. / L. lasallei Coiff.? det. 198 [sic] G.

de Rougemont / Lobrathium n. sp. det. 1990 G. de Rouge-

mont (also in coll. Rgmt.) / Lobrathium n. sp., det. 1990

G. de Rougemont / Holotypus 4 Lobrathium cornutum

sp. n., det. V. Assing 2011” (SMNS). Paratype @ [tener-

al]: same data as holotype (cAss).

Etymology. The specific epithet (Latin, adj.: with horns)

refers to the shape of the ventral process of the aedeagus.

Comment. Both type specimens have been subject to

post-mortem darkening, evidently as a result of exposure

to a chemical of unknown identity.

Description. Length of forebody 5.0—5.4 mm. External

characters highly similar to those of L. bicornutum.

¢: sternite VII moderately transverse, with shallow me-

dian impression posteriorly, with sparse unmodified pu-

bescence, and with weakly concave posterior margin (Fig.

24); sternite VIII strongly oblong, with small median im-

pression posteriorly, this impression with few distinctly

modified, short and stout black setae, posterior excision

small and almost U-shaped (Figs 25—26); aedeagus 1.8

mm long, ventral process deeply bifid, of characteristic

shape (Figs 27—28).

Comparative notes. Lobrathium cornutum 1s reliably dis-

tinguished from the similar L. bicornutum only by the male

sexual characters: the less transverse sternite VII, the more

oblong sternite VIII with a smaller median impression,

with fewer modified setae, and with a smaller posterior

excision, as well as the slightly shorter aedeagus with a

shorter and stouter bifid ventral process.

Distribution and natural history. The type locality is sit-

uated in Terhathum district, Kosi Zone, eastern Nepal (Fig.

23). The teneral specimens were collected in a mixed de-

ciduous forest at an altitude of 2450-2850 m.

Lobrathium triste (Cameron, 1924) (Figs 23, 29-35, 108)

Lathrobium (Lobrathium) triste Cameron, 1924: 193.

Lathrobium kashmiricum Cameron, 1931: 202 f.; syn. n.

Lobrathium afghanicum Coiffait, 1979: 562; syn. n.

Lobrathium nouristanicum Coiffait, 1979: 562 f.; syn. n.

Type material examined. L. triste: lectotype @, present

designation: “Chakrata Dist. Manjgaon 6500, Dr. Came-

ron. 21.V.22 / Lathrobium triste Cam. / Type / M.Came-

ron. Bequest. B.M. 1955-147. / Lectotypus ¢ Lathrobium

Bonn zoological Bulletin 61 (1): 49-128

triste Cameron, desig. V. Assing 2011 / Lobrathium triste

(Cameron), det. V. Assing 2011” (BMNH).

L. kashmiricum: syntype 2 [somewhat damaged, left

hind leg and apical half of right antenna missing]: “Type

/Ramsu 4500, Chenab Valley, Kashmir. 8.V.28. B.M. Bha-

tia. / Under wet stones / 308 / Lathrobium kashmiricum

Cam. Type / M. Cameron. Bequest B.M. 1955-147. / Lo-

brathium triste (Cameron), det. V. Assing 2012” (BMNH).

L. nouristanicum: holotype 9: “entre Camdech et

Kouchtous, 1400-2000 m, 8.76 / Afghanistan L et M /

Holotype / Lobrathium nouristanicum H. Coiffait det.

1978 / Lobrathium triste (Cameron), det. V. Assing 2012”

(MNHNP).

Comment. The original description of L. triste is based

on an unspecified number of syntypes, among them at least

one male, from “Chakrata District; Manjgaon, Dodora

Khud, Chulli Khud, 6500 to 8000 feet above the sea”

(Cameron 1924). A male syntype from Manjgaon was lo-

cated in the Cameron collection at the BMNH and desig-

nated as the lectotype.

The original description of L. kashmiricum 1s based on

an unspecified number of syntypes from “Kashmir: Ram-

su, Chenab Valley, alt. 4500 feet” (Cameron 1931). The

male sexual characters are not described, suggesting that

male syntypes were not available. An examination of a fe-

male syntype, possibly the only type specimen that exists,

yielded no evidence whatsoever that it should not be con-

specific with L. triste. It represents the colour morph with

indistinct posterior spots on the elytra. Moreover, the type

locality is well within the range of L. triste. Therefore, the

available evidence suggests that the syntype of L. kash-

miricum is conspecific with the lectotype of L. triste; hence

the synonymy proposed above. The syntype of L. kash-

miricum 1s illustrated in Fig. 108.

Lobrathium afghanicum was described from a male syn-

type collected in “Darah 1 Nour” and three male paratypes

from “Gorges du Tanguy Garou, Maipar” (Coiffait 1979).

The type material was not examined, but based on the il-

lustration of the characteristic aedeagus provided in the

original description, L. afghanicum is undoubtedly con-

specific with L. triste; hence the synonymy proposed

above.

According to the original description of L. nouristan-

icum, which is based on a unique female from “Nouris-

tan, entre Camdech et Kouchtous” (Coiffait 1979), this

species is distinguished from L. afghanicum (there is no

mention of L. triste) by smaller size, more slender habi-

tus, relatively smaller eyes, longer elytra without poste-

rior spots and with the punctation arranged in more dis-

tinct series. An examination of the holotype revealed that,

in all these characters, it is well within the range of in-

traspecific variation of L. triste, suggesting that it is con-

specific with this species. Consequently, L. nouristanicum

is placed in synonymy with L. triste.

©ZFMK

66 Volker Assing

Fig. 41. Distributions of Himalayan Lobrathium species, based on revised records: L. mordens (filled circles); L. flavipenne (open

diamond); L. wittmeri (filled diamonds); L. kleebergi (open circles); L. semicaeruleum (open triangles).

Additional material examined. Pakistan: |, Chitral, Garam

Chashma, 1900-2800 m, 1.—2.VII.1982, leg. Erber & Heinz

(cAss); 24.3, Swat Marghuzar, 1300 m, 8.V.1983, leg. Besuchet

& Lébl (MHNG, cAss); 14, 629, Chitral, Lotkoh, 2350 m,

29.V.1983, leg. Besuchet & Lébl (MHNG, cAss); 12, North-

west Frontier Province, Hazara, Lower Kagan Valley, ca. 1100

m, 24.V.1927, leg. Champion (BMNH); 2 exs., Kargah valley

near Gilgit, 26.X.2008, leg. Latella (MCSNV, cAss); 2 exs.,

Ghangche district, Hushey valley, 25.—29.VI.2008, leg. Latella

(MCSNYV).

India: 1° [labelled as “Cotype” of L. triste], Chakrata dis-

trict, Binal Gad, 28.V.1922, leg. Cameron (MHNG).

Redescription. Body length 7.3—8.3 mm; length of fore-

body 3.84.3 mm. Habitus as in Fig. 29. Coloration: body

reddish-brown to blackish, elytra sometimes with ill-de-

limited yellowish to reddish spots posteriorly; legs uni-

formly reddish to dark-brown with even darker femora;

antennae reddish, with antennomere I sometimes infus-

cate.

Head (Figs 30, 108) weakly oblong, approximately 1.05

times as long as broad; posterior angles broadly rounded,

practically obsolete; punctation dense and coarse, medi-

an dorsal portion sparsely punctate; interstices without mi-

crosculpture. Eyes large and bulging, slightly more than

half as long as distance from posterior margin of eye to

neck.

Bonn zoological Bulletin 61 (1): 49-128

Pronotum (Figs 30, 108) approximately 1.25 times as

long as broad and nearly as wide as head; punctation sim-

ilar to that of head, but less dense.

Elytra (Figs 30, 108) 0.90—0.95 times as long as, and

distinctly wider than pronotum; punctation coarse and

dense, arranged in somewhat irregular series; interstices

without microsculpture. Hind wings present.

Abdomen narrower than elytra; punctation moderately

fine and moderately dense (less fine and less dense than

in L. badium); posterior margin of tergite VII with palisade

fringe.

G: posterior margin of tergite VIII weakly convex (Fig.

31); sternite VII transverse and with median impression

posteriorly, posterior margin broadly and distinctly con-

cave; sternite VIII approximately as long as broad, pos-

terior excision very deep and rather broad, anterior to this

excision with small cluster of modified, short black setae

(Figs 32—33); aedeagus approximately 1.3 mm long, with

conspicuously spear-shaped ventral process (Figs 34-35).

Distribution. Lobrathium triste has been recorded from

North India (Uttaranchal), Kashmir (Smetana 2004),

Afghanistan, and Pakistan (Fig. 23).

©OZFMK

Revision of East Palaearctic Lobrathium 67

Lobrathium semicaeruleum (Cameron, 1921) (Figs

36-41)

Lathrobium semicaeruleum Cameron, 1921: 271.

Type material examined. Lectotype @, present designa-

tion: “Gopaldhara, Br. Sikkim. H. Steven. / L. semi-

caeruleum / Type / M.Cameron. Bequest. B.M. 1955—147.

/Lectotypus ¢ Lathrobium semicaeruleum Cameron, de-

sig. V. Assing 2011 / Lobrathium semicaeruleum

(Cameron), det. V. Assing 2011” (BMNH).

Comment. The original description of L. semicaeruleum

is based on an unspecified number of syntypes, among

them at least one male, from “Gopaldhara, Sikkim ..., West

Bhatkot, alt. 4000 feet, and West Almora, Kumaon”

(Cameron 1921). A male syntype from Gopaldhara was

located in the Cameron collection at the BMNH and des-

ignated as the lectotype.

Additional material examined. India: | 9 [labelled as “Cotype”

of L. semicaeruleum; identification doubtful], Dehra Dun, 1922,

leg. Cameron (MHNG).

Nepal: 43.4, 299 [5 exs. with, 1 without elytral spots; 1 ex.

teneral]: Khandbari district, Arun valley at Num main bridge,

1000 m, 21.IV.1984, leg. Smetana & L6bl (cSme, cAss).

Redescription. Body length 6.0—7.2 mm; length of fore-

body 3.4-3.9 mm. Coloration variable: forebody black-

ish with distinct bluish hue; abdomen black; elytra with

or without large reddish spot occupying posterior 1/3—2/5

of elytra; legs blackish-brown to blackish, with slightly

paler tarsi; antennae blackish-brown to black.

Head (Fig. 36) approximately as long as broad; poste-

rior angles broadly rounded, practically obsolete; punc-

tation dense and coarse, median dorsal portion sparsely

punctate; interstices without microsculpture. Eyes very

large and bulging, distinctly more than half as long as dis-

tance from posterior margin of eye to neck.

Pronotum (Fig. 36) approximately 1.25—1.30 times as

long as broad and approximately 0.9 times as wide as

head, lateral margins distinctly convex in dorsal view;

punctation very dense, somewhat coarser than that of head.

Elytra (Fig. 36) conspicuously long, 1.10—1.15 times as

long as, and distinctly wider than pronotum; punctation

coarse and dense, not arranged in series; interstices with-

out microsculpture. Hind wings present.

Abdomen narrower than elytra; punctation fine and

dense; posterior margin of tergite VII with palisade fringe.

S: posterior margin of tergite VIII weakly convex; ster-

nite VI with small median tubercle; sternite VI with pro-

nounced median tubercle, posterior margin weakly con-

cave; sternite VIII weakly transverse, with long and broad

median impression, this impression with numerous mod-

ified, short black setae, posterior excision relatively deep

and U-shaped (Figs 37-38); aedeagus approximately |

mm long, with apically bifid ventral process of character-

Bonn zoological Bulletin 61 (1): 49-128

istic shape (Figs 39-40).

Intraspecific variation. The coloration of the elytra is

variable. A posterior elytral spot may be present or absent.

In one locality in eastern Nepal, both colour morphs were

collected together.

Distribution and natural history. Confirmed records of

this species are known only from the type locality in North

India and one locality in eastern Nepal (Fig. 41). Other

previous literature records (Cameron 1921, 1931), as well

as the examined female-based record from Uttaranchal

must be regarded as doubtful and require confirmation;

they may well refer to L. mordens or other similar species.

The examined material was found at altitudes of 1000 and

approximately 1200 m. One specimen collected in April

is teneral.

Lobrathium mordens sp. n. (Figs 41-45)

Type material. Holotype 4: “W. Almora, Kumaon, In-

dia. H. G. C. / G.C. Champion coll. B.M. 1927-409 /

Holotypus ¢ Lobrathium mordens sp. n., det. V. Assing

2012” (BMNH). Paratypes: 240: “2115 / W. Almora

Divn. Kumaon. U.P., Apr, 1918. HGC” (BMNH, cAss);

1 ex.: same data, but “Mar. 1917” (cAss); 19 [teneral]:

“Ranikhet, Kumaon. H. G. C.” (BMNH); 2 exs.:

“Ranikhet Div., Kumaon, U.P., Feb. ‘20, HGC.” (BMNH);

2 exs.: “Ranikhet, Kumaon, India. H.G.C.” (BMNH); 1

ex.: “West Bhatka [?], Kumaon 4000’, Mar ‘20 HGC”

(BMNH)); 2 exs.: “India, Haldwani Dist., Kumaon, H.G.

Champion” (BMNH); 3 exs.: “Haldwani Div., Kumaon,

India. H.G.C.” (BMNH, cAss); 1 ex.: “2745 / W. Almo-

ra Dn, Almora U.P., Mar. ‘19. HGC” (cAss); 1 ex.: “Dhelu

[= Chauntra], Mandi [Mandi District in Himachal

Pradesh], Punjab. 4500ft. H.G.C.” (cAss).

Etymology. The specific epithet (Latin, present participle

of mordere: to sting) refers to the long and slender ven-

tral process of the aedeagus.

Description. Body length 5.5—7.3 mm; length of fore-

body: 3.0-3.8 mm. Forebody uniformly black with

bluish hue. Other external characters as in L. semi-

caeruleum; reliably distinguished only by the male sexu-

al characters.

3: sternite VI unmodified; sternite VII with shallow and

extensive median impression, pubescence not distinctly

modified, posterior margin broadly convex (Fig. 42); ster-

nite VII weakly oblong, in the middle with relatively nar-

row, but deep oblong impression, this impression with nu-

merous modified, short and stout black setae, posterior ex-

cision broadly and rather deeply U-shaped (Fig. 43);

aedeagus |.2—1.3 mm long, with conspicuously long, slen-

©ZFMK

68 Volker Assing

Figs 42-51. Lobrathium mordens (42-45) and L. unispinosum (46-51). 42, 48: male sternite VII; 43, 49: male sternite VII; 44—45,

50-51: aedeagus in lateral and in ventral view; 46: habitus; 47: forebody. Scale bars: 46-47: 1.0 mm; 42-45, 48-51: 0.5 mm.

der, and apically dorso-ventrally dilated ventral process

(Figs 44-45).

Comparative notes. From other externally similar Hi-

malayan representatives with a uniformly blackish fore-

body with bluish hue (L. semicaeruleum, L. unispinosum,

L. integrum), L. mordens is distinguished by the male sec-

ondary sexual characters and particularly by the complete-

ly different shape of the ventral process of the aedeagus.

Distribution and natural history. Lobrathium mordens

is currently known from several localities in Uttaranchal

and Himachal Pradesh in northern India (Fig. 41). Two

specimens were collected at altitudes of approximately

1300-1500 m.

Bonn zoological Bulletin 61 (1): 49-128

Lobrathium unispinosum sp. n. (Figs 23, 46—51)

Type material. Holotype 3: “Nepal, 31.10.92, Pokhara-

See, leg. J. Schmidt / Holotypus <4 Lathrobium

unispinosum sp. n., det. V. Assing 2011” (SDEI).

Paratypes: 14, 59 9: same data as holotype (SDEI, cAss);

12: “Nepal, Annapurna, Pokhara Umg. / 800 m,

26.-28.11.1994, leg. Ahrens” (SDEI); 19 [without bluish

hue]: “Nepal, Annapurna Mts., Landruk-Bhari Kharka,

1650-1900 m, 6.5.1999, leg. C. Kriiger, G. Hirthe” (cAss);

19: “Ost-Nepal, Rolwaling Himal / Tama Koshi Tal, Suri

Dhoban, 1200 m, 14.05.2000, leg. A. Kleeberg” (cKle).

Etymology. The specific epithet (Latin, adjective: with

one tooth) refers to the shape of the ventral process of the

aedeagus.

©ZFMK

Revision of East Palaearctic Lobrathium 69

Description. Body length 6.3—7.3 mm; length of forebody

3.4-3.8 mm. Habitus as in Fig. 46. Coloration: forebody

blackish, usually with distinct bluish hue; abdomen black;

legs blackish-brown to blackish, with slightly paler tarsi;

antennae blackish-brown to black.

In external characters highly similar to L. semi-

caeruleum (Figs 46-47), but reliably distinguished by the

male sexual characters:

&: posterior margin of tergite VIII weakly convex; ster-

nite VI without median tubercle; sternite VII with medi-

an impression posteriorly, but without tubercles, posteri-

or margin broadly concave (Fig. 48); sternite VII approx-

imately as long as broad, with long median impression,

this impression with numerous modified, short and stout

black setae, posterior excision relatively small and con-

cave (Fig. 49); aedeagus 1.2—1.3 mm long, ventral process

asymmetric, on right ventral side (ventral view) with pro-

nounced tooth-shaped process, dorsal plate lamellate (Figs

50-51).

Intraspecific variation. One female from the Annapur-

na lacks the bluish hue and has the forebody black, but

other external characters are identical to those of the ma-

terial from other localities.

Comparative notes. From the externally highly similar

L. semicaeruleum, this species 1s reliably distinguished on-

ly by the modifications of the male sternites VI—-VIII (ster-

nites VI and VII without tubercles, posterior excision of

sternite VIII much smaller and less deep), as well as by

the larger and differently shaped, asymmetric aedeagus.

Distribution and natural history. Lobrathium

unispinosum was collected in central Nepal, one female

also in eastern Nepal (Fig. 23), at altitudes of 800-1900

Lobrathium integrum sp. n. (Figs 23, 52-57)

Type material. Holotype ¢ [slightly teneral]: “Nepal, Ta-

plejung, Mewa Khola 1570 m, 02.VI.2011, leg. S.

Taman, 27°33713N, 87°36’ 10E / Collection Naturkunde-

museum Erfurt / Holotypus ¢ Lathrobium integrum sp.

n., det. V. Assing 2011” (NME). Paratypes: 14, 829

[mostly teneral]: same data as holotype (NME, cAss).

Etymology. The specific epithet (Latin, adjective: immac-

ulate, pure) refers to uniform coloration of the elytra.

Description. Body length 6.9—7.9 mm; length of forebody

3.7-4.0 mm. Habitus as in Fig. 52. Coloration: forebody

blackish, with distinct bluish hue; abdomen black; legs

blackish-brown to blackish, with slightly paler tarsi; an-

tennae blackish-brown to black.

Bonn zoological Bulletin 61 (1): 49-128

In external characters highly similar to L. semi-

caeruleum and L. unispinosum (Figs 52-53), but reliably

distinguished by the male sexual characters:

&: sternite VI unmodified; sternite VII with shallow me-

dian impression and with pair of tubercles posteriorly, pos-

terior margin broadly concave (Fig. 54); sternite VIII

weakly oblong, with narrow median impression in ante-

rior half, this impression with numerous modified, short

and stout black setae, posterior excision conspicuously

deep and narrow (Fig. 55); aedeagus 1.4 mm long, ven-

tral process somewhat asymmetric and of distinctive

shape, dorsal plate lamellate (Figs 56-57).

Comparative notes. From the externally highly similar

L. semicaeruleum and L. unispinosum, this species is read-

ily distinguished by the shapes and chaetotaxy of the male

sternites VII and VIII, as well as by the distinctive shape

of the ventral process of the aedeagus.

Distribution and natural history. Lobrathium integrum

is currently known from only one locality in Taplejung dis-

trict in the very northeast of Nepal (Fig. 23). The speci-

mens, most of which are somewhat teneral, were collect-

ed at an altitude of 1570 m.

Lobrathium ochreonotatum (Champion, 1922), revali-

dated (Figs 58-61, 66)

Lathrobium semicaeruleum var. ochreonotatum Champion,

1922: 34.

Type material examined. Lectotype, present designation:

“R. [River] Sarda Gorge, Kumaon, U. P., Dec. 1918, HGC

/ Lathrobium semicaeruleum Cam. v. ochreonotatum Ch.

/ 2632 / Brit. Mus. 1922-71. / Syntype / Lectotypus 3

Lathrobium ochreonotatum Champion, desig. V. Assing

2011 / Lobrathtum ochreonotatum (Champion), det. V.

Assing 2011” (BMNH). Paralectotype: 19: “Ranikhet,

Kumaon, India, H.G.C. / Lathrobium semicaeruleum Cam.

yv. ochreonotatum Ch. / Brit. Mus. 1922—71. / Syntype”

(BMNH).

Comment. Champion (1922) described L. ochreonatum

as a variety of L. semicaeruleum, stating that it was char-

acterised by “a large ochreous patch at the apex of the ely-

tra”, but “differing in no other way from the type”. The

four type specimens of this variety were apparently col-

lected together with a series of specimens with uniform-

ly coloured elytra. The variety was treated as a synonym

of L. semicaeruleum by Cameron (1931).

Two syntypes, a male and a female, from the Champion

collection at the BMNH were made available for study;

the male is designated as the lectotype. An examination

of these types, as well as of additional specimens previ-

ously identified as L. semicaeruleum with red-spotted ely-

©ZFMK

70 Volker Assing

58 3 63

Figs 52-65. Lobrathium integrum (52-57), L. ochreonotatum (58-61), and L. flavipenne (62-65). 52, 62: habitus; 53: forebody

54, 58, 63: male sternite VII; 55, 59: male sternite VII; 56-57, 60-61, 64-65: aedeagus in lateral and in ventral view. Scale bars:

52-53, 62: 1.0 mm; 54-61, 63-65: 0.5 mm.

Bonn zoological Bulletin 61 (1): 49-128 ©ZFMK

Revision of East Palaearctic Lobrathium 71

Fig. 66. Distributions of Himalayan Lobrathium species, based on revised records: L. alticola (open squares); L. cylindricolle (filled

diamond); L. ochreonotatum (open triangles); L. biaculeatum (filled circles).

tra revealed that they clearly represent a distinct species,

so that the name L. ochreonotatum is revalidated.

Additional material examined. Nepal: | ex., Annapurna range,

Landruk-Bhari Kharka, 1650-1900 m, 6.V.1999, leg. Kriiger &

Hirthe (cAss); 2 exs., Annapurna, Tadapani-Chomrong, Kyum-

nu Khola, 1950 m, 4.V.1999, leg. Kriiger & Hirthe (cAss); 4 exs.,

Annapurna, NE Pokhara Sikles, 2000 m, 15.V.1993, leg. Schmidt

(cKle, cAss); 3 exs., SW-Manaslu, Bhara Pokhari Lekh, below

Taksa, 1500-1700 m, 31.III.1999, leg. Jager (SNSD, cAss); 3

exs., Manaslu, Marsyangdi Khola, E-side, Bhachok Khola, 1000

m, 13.1V.1999, leg. Hirthe (cAss); 3 exs., Manaslu, Marsyang-

di Khola, E-side, above Besi Sahar, 1500 m, 31.11.1999, leg.

Hirthe (cAss); 3 exs., Manaslu, Ngadi Khola, 6 km NE Ngadi,

1350 m, 17.V.2005, leg. Schmidt (NME, cAss); | ex., Mahar-

kali/Darchula, Latinath, 29°44’N, 80°46’E, 1030 m, stream bank,

31.V—1.VI.2005, leg. Weigel (NME); | ex., Dhaulagiri, Baglung

Lekh, above Okhle vill., 2460 m, 13.V.2004, leg. Kleeberg

(cAss); 1 ex., Pokhara, southern shore of Phewa lake, 800—900

m, stream bank, sifted, 8.V.2001, leg. Hirthe (cAss); 1 ex.,

Pokhara lake, 31.X.1992, leg. Schmidt (cAss); 5 exs., Kali-Gan-

daki river, between Tatopani and Beni, 1000 m, 29.X.1992, leg.

Schmidt (SDEI, cAss); 2 exs., W Annapurna, between Lete and

Dana, 1400-2400 m, 25.X.1992, leg. Schmidt (SDEI, cAss); 2

exs. [1 teneral], Annapurna, Sikles range, Garlang, 1350 m,

18.1V.1996, leg. Schmidt (SDED; 1 ex., Annapurna, N Sikles,

below Dhara Kharka, 1750 m, 26.1V.1996, leg. Schmidt (cAss);

1 ex., Annapurna, Marsyangdi Khola, Chamje bis valley,

1400-1700 m, 11.VI.1994, leg. Schmidt (SDEI); 13 [teneral;

without elytral spots], E-Nepal, Ramechap district, Khimti Kho-

Bonn zoological Bulletin 61 (1): 49-128

la near Shivalaya, 1800 m, 5.V.1997, leg. Schawaller (SMNS).

India: Uttaranchal: 7 exs., ca. 30 km N Bageshwar, Khati vil-

lage, 2100-2300 m, 27.—30.VI.2003, leg. Kejval & Tryzna (cSch,

cAss); 14 [slightly teneral], West Almora Division, Kumaon,

VI.1917, leg. Champion (BMNH); | ex., Kumaon, Sarda Gor-

ge, XII.1018, leg. Champion (BMNH); 2 exs., Kumaon, West

Bhatkat, V.1920, leg. Champion (BMNH, cAss); 2 exs., N Ku-

maon, Gori R. Gorge, leg. Champion (BMNH, cAss); 2 exs. [1

slightly teneral], Haldwani Division, Kaldhunga, 26.III.1923, leg.

Champion (BMNH).

Redescription. External characters as in L. semi-

caeruleum. Elytra usually with, rarely without large bright-

reddish spots in the posterior portion of the elytra.

3: sternite VI unmodified; sternite VII with pair of tu-

bercles posteriorly, posterior margin weakly concave (Fig.

58); sternite VIII weakly transverse, with long and broad

median impression, this impression with numerous mod-

ified, short and stout black setae, posterior excision rela-

tively deep and broadly U-shaped (Fig. 59); aedeagus ap-

proximately 0.9 mm long, ventral process of characteris-

tic shape, apically not bifid (Figs 60-61).

Intraspecific variation. In the male from Ramechap dis-

trict, the usually large and distinct elytral spots are miss-

ing; the elytra are uniformly black with a distinct bluish

hue.

OZFMK

72 Volker Assing

Figs 67-76. Lobrathium flavipenne (67), L. kleebergi (68-72), and L. guttula, lectotype (73-76). 67, 70, 74: male sternite VIII;

68: habitus; 69, 73: male sternite VII; 71-72, 75-76: aedeagus in lateral and in ventral view. Scale bars: 68: 1.0 mm; 67, 69-76:

0.5 mm.

Comparative notes. As can be inferred from the highly

similar external morphology and particularly the similar

male primary and secondary sexual characters, L. ochre-

onotatum 1s closely related to, and probably the adelpho-

taxon of L. semicaeruleum, from which it is readily dis-

tinguished by the morphology of the aedeagus. In addi-

tion, it is separated by the slightly different modifications

of the male sternites VI and VIII.

Bonn zoological Bulletin 61 (1): 49-128

Distribution and natural history. Lobrathium ochreono-

tatum is apparently widespread in the Himalaya (North In-

dia, Nepal) (Fig. 66), where it has been found at altitudes

of 800-2460 m, at least on one occasion together with L.

unispinosum. Teneral specimens were collected during the

period from March through June.

OZFMK

Revision of East Palaearctic Lobrathium 73

Lobrathium flavipenne sp. n. (Figs 41, 62—65, 67)

Type material. Holotype ¢: “Parbatti V., Kulu. Punjab.

6—8,000 ft. H. G. C./ G.C. Champion coll. B.M. 1927-409

/ Holotypus 3 Lobrathium flavipenne sp. n., det. V. As-

sing 2012” (BMNH).

Etymology. The specific epithet (Latin, adjective: with

yellow wings) alludes to the distinctive coloration of the

elytra.

Description. Body length 7.4 mm; length of forebody 3.8

mm. Habitus as in Fig. 62. Coloration: head and prono-

tum blackish with distinct bluish hue; elytra yellowish,

with the anterior third infuscate; abdomen blackish; legs

and antennae blackish-brown. Other external characters

similar to those of L. semicaeruleum and L. ochreonota-

tum.

&: sternite VII with shallow and extensive median im-

pression, near anterior margin with transverse, bi-domed

tubercle, posterior margin weakly concave (Fig. 63); ster-

nite VIII weakly oblong, median impression extensive and

with numerous modified, stout and short black setae, pos-

terior excision deep and U-shaped (Fig. 67); aedeagus 1.1

mm long, symmetric, and with apically bifid ventral

process in ventral view (Figs 64-65).

Comparative notes. This species is readily distinguished

from the spotted morphs of L. semicaeruleum and L.

ochreonotatum, as well as from other similar species by

the extensively yellowish coloration of the elytra, as well

as by the male primary and secondary sexual characters.

Distribution and natural history. The type locality is sit-

uated near Kulu in Himachal Pradesh, northern India (Fig.

41), at an altitude of approximately 2000-2600 m.

Lobrathium kleebergi sp. n. (Figs 41, 68—72)

Type material. Holotype 3: “Nepal, nordéstl. Kathman-

du-Tal, 06.05.1993, leg. A. Kleeberg / Holotypus ¢ Lo-

brathium kleebergi sp. n., det. V. Assing 2011” (cAss). Pa-

ratypes: 244: same data as holotype (cKle, cAss); 1d

[slightly teneral]: “Nepal-Himalaya, Annapurna Mts.,

Pokhara-See, Flu8miindung, 850 m, 10.5.1993, leg.

Schmidt” (cKle); 1¢ [slightly teneral]: “Nepal, Tama

Koshi Vall., Suri Doban, 1100—1300 m, 04.06.2000, leg.

J. Schmidt” (cKle); 14: “Nepal Manaslu Mts., Bara

Pokhari Leak [sic] above Bhachok Goan vill., 1600—1800

m, leg. J. Schmidt, 29.IV.2005, 28°14°28N 84°24°32E”

(NME).

Bonn zoological Bulletin 61 (1): 49-128

Etymology. This species is dedicated to Andreas Kleeberg,

who collected most of the type specimens.

Description. Body length 6.6—8.0 mm; length of forebody

4.0-4.6 mm. Habitus as in Fig. 68. Coloration: body black-

ish, elytra with relatively small subcircular yellowish spot,

this spot reaching neither suture, nor lateral margin, nor

posterior margin; legs reddish brown with reddish tarsi,

femora sometimes infuscate; antennae reddish.

Head approximately as long as broad; posterior angles

broadly rounded, weakly marked; punctation coarse and

conspicuously dense, interstices reduced to narrow ridges,

median dorsal portion and frons sparsely punctate or im-

punctate; interstices without microsculpture. Eyes mod-

erately large, approximately half as long as distance from

posterior margin of eye to neck.

Pronotum 1.15—1.20 times as long as broad and

0.90—0.95 times as wide as head, lateral margins almost

straight and weakly converging in dorsal view; punctation

moderately dense, distinctly less dense than that of head.

Elytra of variable length, 0.9—1.1 times as long as, and

somewhat wider than pronotum; punctation coarse and

dense, not arranged in distinct series; interstices without

microsculpture. Hind wings apparently present.

Abdomen narrower than elytra; punctation fine and

dense; posterior margin of tergite VII with palisade fringe.

3: posterior margin of tergite VIII strongly convex; ster-

nite VI unmodified; sternite VII strongly transverse and

with median impression posteriorly, this impression with

pair of clusters of numerous modified, stout and short

black setae, posterior margin broadly and rather deeply

concave (Fig. 69); sternite VIII weakly transverse, with

broad and deep median impression, this impression with

numerous modified, stout and short black setae, posteri-

or excision rather broad and of subtrapezoid shape, in the

middle with short acute process, on either side of excision

with dense blackish pubescence (Fig. 70); aedeagus ap-

proximately 1.1 mm long, with large blade-shaped ven-

tral process (Figs 71—72).

Comparative notes. This species is characterised partic-

ularly by the male sexual characters. It is readily distin-

guished from L. ochreonotatum by external characters

alone: much larger and broader body with a less slender

pronotum, extremely dense punctation of the head, ab-

sence of a bluish hue on the forebody, smaller elytral spot.

Distribution and natural history. Lobrathium kleeber-

gi is currently known only from several localities in Nepal

(Fig. 41). Some of the type specimens were collected at

altitudes between 850 and approximately1700 m. The

specimens from the type locality were floated from the

bank of a stream (Kleeberg, pers. comm.).

©OZFMK

74 Volker Assing

83 84

87 88 86

Figs 77-88. Lobrathium sinuatum (77-82) and L. pustulatum, lectotype (83-88). 77, 83: habitus; 78, 84: forebody; 79, 85: male

sternite VII; 80, 86: male sternite VIII; 81-82, 87-88: aedeagus in lateral and in ventral view. Scale bars: 77-78, 83-84: 1.0 mm;

79-82, 85-88: 0.5 mm.

Lobrathium guttula (Fauvel, 1895), comb. n. (Figs

73-76, 140)

Lathrobium guttula Fauvel, 1895: 234 f.

Type material examined. Lectotype 3, present designa-

tion: “Carin, Asciuii Ghect’ [near Loi-kaw; approx.

19°41°N, 97°1S°E], 1400-1500 m, L. Fea. HI-IV.88. /

Coll. R. I. Sc. N. B. / Ex-Typis / Lectotypus ¢ Lathrobi-

Bonn zoological Bulletin 61 (1): 49-128

um guttula Fauvel, desig. V. Assing 2011 / Lobrathium

guttula (Fauvel), det. V. Assing 2011” (IRSNB). Paralec-

totypes: 19: same data as lectotype; 1 9: “Darjeeling / gut-

tula Fvl. / Coll. R. I. Sc. N. B. / Ex-Typis” (IRSNB).

Comment. The original description is based on an unspec-

ified number of syntypes, among them at least one male,

from “Birmanie, Carina Asciuii Ghécu, 1400-1500 m,

©OZFMK

Revision of East Palaearctic Lobrathium 75

Fig. 89. Distributions of Himalayan Lobrathium species, based on revised records: L. bicarinatum (open square); L. sinuatum (filled

triangle); L. kosiense (open circle); L. pustulatum (open diamond).

I-IV (L. Fea). Sikkim, Darjeeling (Christie)” (Fauvel

1895). Three syntypes, a male and two females, were lo-

cated in the collections of the IRSNB; the slightly tener-

al male is designated as the lectotype. It is currently un-

clear whether or not the female syntype from Darjeeling

is conspecific with the lectotype; the possibility that it

refers to L. kleebergi or to an undescribed species cannot

be ruled out. Although Fauvel (1895) had placed L. gut-

tula near L. multipunctum, it had previously been listed

as Lathrobium even in recent catalogues (Smetana 2004).

The species clearly belongs to Lobrathium.

Redescription. External characters as in L. kleebergi (see

preceding section). Distinguished only by the male sex-

ual characters:

S: sternite VI unmodified; sternite VII strongly trans-

verse and with pronounced median impression, this im-

pression impunctate in the middle and laterally with a pair

of clusters of numerous modified, stout and short black

setae, posterior margin broadly and weakly concave (Fig.

73); sternite VIII as long as broad, with broad and deep

median impression, this impression with numerous mod-

ified, stout and short black setae, posterior excision rather

broad and shallow, in the middle triangularly produced and

with row of five short, black marginal setae on either side

(Fig. 74); aedeagus 1.3 mm long and with blade-shaped

ventral process (Figs 75—76).

Comparative notes. As can be inferred from the similar

external (large head with extremely dense punctation and

large impunctate patches) and from the similar male sex-

ual characters, L. guttula is a close relative of L. kleeber-

Bonn zoological Bulletin 61 (1): 49-128

gi. It is distinguished from the latter by the shape and

chaetotaxy of the male sternites VII and VIII, as well as

by the different shape of the ventral process of the aedea-

gus. For characters separating it from L. sinuatum, its pre-

sumable sister species, see the following section.

Distribution and natural history. Lobrathium guttula

was originally described from Myanmar and Darjeeling

(North India: West Bengal). Cameron subsequently report-

ed it from Siwaliks in Uttaranchal. In the Palaearctic Cat-

alogue (Smetana 2004), it is also listed for Himachal

Pradesh and the Oriental region. In view of the external

similarity of Lobrathium species and the fact that the male

sexual characters of the type material were previously un-

known, all records, except those from the type locality,

which is situated in Kaya State close to the town Loi-kaw

(Fig. 140), should be considered doubtful and require con-

firmation.

Two of the type specimens were collected at an altitude

of 1400-1500 m during spring (March/April). The lecto-

type is slightly teneral.

Lobrathium sinuatum sp. n. (Figs 77—82, 89)

Type material. Holotype @: “Nepal, 31.10.92, Pokhara-

See, leg. J. Schmidt / Holotypus 4 Lobrathium sinuatum

sp. n., det. V. Assing 2011” (SDEI).

Etymology. The specific epithet (Latin, adjective) alludes

to the bisinuate posterior margin of the male sternite

VUL.

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76 Volker Assing

Description. Body length 7.2 mm; length of forebody 4.1

mm. Habitus as in Fig. 77. Coloration: body blackish, ely-

tra with relatively small subcircular yellowish spot, this

spot reaching neither suture nor lateral or posterior mar-

gins; legs blackish with reddish tarsi; antennae dark-

brown.

Head (Fig. 78) 1.05 times as long as broad; lateral mar-

gins behind eyes convexly rounded towards neck, poste-

rior angles obsolete; punctation moderately coarse and

very dense, interstices reduced to narrow ridges, transverse

patch in median dorsal portion and frons impunctate; in-

terstices without microsculpture. Eyes moderately large,

approximately half as long as distance from posterior mar-

gin of eye to neck.

Pronotum (Fig. 78) 1.22 times as long as broad and 0.9

times as wide as head, lateral margins almost straight and

weakly converging in dorsal view; punctation moderate-

ly dense, distinctly less dense and somewhat coarser than

that of head.

Elytra (Fig. 78) nearly 1.1 times as long as, and distinct-

ly wider than pronotum; punctation coarse and dense,

arranged in very irregular series; interstices without mi-

crosculpture. Hind wings apparently present.

Abdomen narrower than elytra; punctation fine and

dense; posterior margin of tergite VII with palisade fringe.

3: posterior margin of tergite VIII weakly convex; ster-

nite VI unmodified; sternite VII moderately transverse and

with median impression posteriorly, this impression de-

limited by approximately 20 modified, stout and short

black setae on either side, posterior margin broadly and

weakly concave (Fig. 79); sternite VHI oblong, with broad

median impression, this impression with numerous mod-

ified, stout and short black setae, posterior excision shal-

low and bisinuate (Fig. 80); aedeagus approximately 1.2

mm long, with large blade-shaped ventral process (Figs

81-82).

Comparative notes. Lobrathium sinuatum 1s charac-

terised particularly by the conspicuous shape and chaeto-

taxy of the male sternite VIII, as well as by the shape and

chaetotaxy of the male sternite VII and the morphology

of the aedeagus. It is additionally separated from the sim-

ilar and geographically close L. kleebergi by the darker

legs and antennae, as well as by the more slender prono-

tum. Based on the similarly derived male sexual charac-

ters (shapes and chaetotaxy of the male sternites VII and

VUI; morphology of the aedeagus), L. sinuatum is the

adelphotaxon of L. guttula, from which it is separated by

the slightly different shape of the ventral process of the

aedeagus (both in lateral and in ventral view), the differ-

ent shape of the dorso-apical portion of the aedeagus in

lateral view, the slightly different chaetotaxy of the less

transverse male sternite VII, as well as by the slightly dif-

ferent shape and chaetotaxy of the male sternite VIII.

Bonn zoological Bulletin 61 (1): 49-128

Distribution and natural history. The type locality is sit-

uated near Pokhara in central Nepal (Fig. 89), where the

holotype was collected together with L. unispinosum.

Lobrathium pustulatum (Cameron, 1931), comb. n.

(Figs 83-89)

Lathrobium pustulatum Cameron, 1931: 202.

Type material examined. Lectotype ¢ [somewhat dam-

aged, four legs missing, antennae broken], present desig-

nation: “Naga Hills, Assam (8) / L. pustulatum / M.

Cameron. Bequest. B.M. 1955-147. / Syntype / Syntype

Lathrobium pustulatum Cameron, 1931, det. R.G. Booth

2011 / Lectotypus 4 Lathrobium pustulatum Cameron, de-

sig. V. Assing 2012 / Lobrathium pustulatum (Cameron),

det. V. Assing 2012” (BMNH). Paralectotype 2: same da-

ta as lectotype (BMNH).

Comment. The original description is based on an unspec-

ified number of syntypes, among them at least one male,

from “Assam: Naga Hills, alt. 9000 feet” collected by “S.

N. Chatterjee” (Cameron 1931). Two syntypes, a male and

a female, were located in the collections of the BMNH.

The somewhat damaged male is designated as the lecto-

type. The species undoubtedly belongs to Lobrathium.

Redescription. Body length 7.0—7.5 mm; length of fore-

body 4.0—4.2 mm. Habitus as in Fig. 83. Coloration: body

blackish, elytra with a relatively large yellowish spot not

reaching posterior and lateral margins; legs yellowish with

the apices of the femora and the tibiae infuscate; anten-

nae brown to dark-brown with darker antennomere I.

Head (Fig. 84) approximately as long as broad, widest

accross eyes, convexly narrowed posteriad behind eyes;

posterior angles broadly rounded, weakly marked; punc-

tation coarse and dense, interstices reduced to narrow

ridges in posterior and lateral dorsal portions, median dor-

sal portion and frons more sparsely punctate; interstices

without microsculpture. Eyes large and strongly convex,

distinctly more than half as long as distance from poste-

rior margin of eye to neck.

Pronotum (Fig. 84) 1.25—1.30 times as long as broad and

0.90-0.95 times as wide as head, lateral margins weakly

convex in dorsal view; punctation moderately dense, dis-

tinctly less dense than that of head; impunctate midline

distinct and of moderate breadth.

Elytra (Fig. 84) long and broad, approximately 1.05

times as long as, and much wider than pronotum; punc-

tation coarse and dense, partly obliquely confluent, and

partly arranged in irregular and distinctly oblique series;

interstices without microsculpture. Hind wings present and

apparently fully developed.

Abdomen narrower than elytra; punctation moderately

fine and dense, shallower on posterior than on anterior ter-

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Revision of East Palaearctic Lobrathium 77

gites; microsculpture shallow on anterior tergites, distinct

on tergite VII; posterior margin of tergite VII with palisade

fringe.

&: posterior margin of tergite VIII convex; sternites V

and VI with shallow impression posteriorly; sternite VII

strongly transverse and with pronounced median impres-

sion posteriorly, this impression without pubescence in the

middle and semitransparent posteriorly, posterior margin

broadly and rather strongly concave (Fig. 85); sternite

VII as long as broad, with deep and relatively narrow me-

dian impression, this impression with numerous modified,

stout and short black setae, posterior excision deep and

narrow (Fig. 86); aedeagus 1.5 mm long and with ventral

process of characteristic shape (Figs 87-88).

©: posterior margin of tergite VIII pointed, almost of

triangular shape.

Comparative notes. Lobrathium pustulatum 1s distin-

guished from the similarly coloured L. kleebergi particu-

larly by the coloration of the legs, the shape of the head,

the larger and more bulging eyes, the partly confluent

punctation of the elytra, the shape and chaetotaxy of the

male sternites VII and VIII, and by the completely differ-

ent morphology of the aedeagus.

Distribution and natural history. Lobrathium pustula-

tum has become known only from the type locality in As-

sam, northeastern India (Fig. 89), where the type speci-

mens were collected at an altitude of about 2750 m.

Lobrathium wittmeri Coiffait, 1982 (Figs 41, 90-95)

Lobrathium wittmeri Coiffait, 1982b: 283.

Type material examined. Holotype @: “Phulchoki 2600

m, 11.—14.6. / Nepal, W. Wittmer, C. Baroni U. 1976 /

Holotype / Lobrathitum wittmeri H. Coiffait 1982 / Lo-

brathium wittmeri Coiffait, det. V. Assing 2011” (NHMB).

Comment. The original description of L. wittmeri is based

on a single male holotype and a female paratype from “Neé-

pal, Phulchoki, 2600 m” (Coiffait 1982b). The holotype

is deposited in the collections of the NHMB.

Additional material examined. Nepal: 13 exs. [micropterous],

Dhaulagiri range, Baglung Lekh, ca. 30 km W Baglung, north-

ern Tara Khola, 2800 m, 20.V.2004, leg. Kleeberg (cKle, cAss);

1 ex. [micropterous], same data, but 2700-2900 m (cKle); 7 exs.

[6 exs. micropterous, 12 macropterous], same data, but upper

Tara Khola, 2600 m, 18.V.2004 (cKle, cAss); 19 [micropterous],

Annapurna, Marsyangdi valley, Bagarchhap, 2100 m, 4.V.2007,

leg. Schmidt (NME); 6 exs. [micropterous], Annapurna, Sikles

range, Nyauli Kharka, S Sikles, 2400 m, 21.—24.IV.1996, leg.

Schmidt (SDEI, cAss); 19 [micropterous], Manaslu, Barapok-

hari Lekh, Lake Barapokhara env., 12 km NE Besisahar villa-

ge, 28°18’N, 84°28’E, 3100 m, 10.1X.2000, leg. Hetzel (cFel);

14, Annapurna Himal, Sikles range, 1400-2100 m, V.1996, leg.

J. Schmidt (NME).

Bonn zoological Bulletin 61 (1): 49-128

Redescription. Body length 6—7 mm; length of forebody

3.0—3.5 mm. Coloration: body blackish; legs dark-brown

to blackish-brown with reddish tarsi; antennae dark-

brown, with antennomere | usually even darker and with

the apical antennomeres usually reddish.

Head (Figs 90-91) approximately as long as broad, oc-

casionally weakly transverse or weakly oblong; posteri-

or angles broadly rounded; punctation dense and moder-

ately coarse, median dorsal portion more sparsely punc-

tate; interstices without microsculpture. Eyes rather large,

approximately half as long as distance from posterior mar-

gin of eye to neck, or slightly larger.

Pronotum (Figs 90-91) approximately 1.25 times as

long as broad and approximately 0.95 as wide as head, lat-

eral margins weakly convex in dorsal view; punctation

similar to that of head, but less dense.

Elytra dimorphic, in micropterous morph short (but see

comment below), 0.70—0.75 times as long and approxi-

mately as wide as pronotum (Fig. 90), or nearly so, in

macropterous morph 1.05 times as long as, and much

broader than pronotum (Fig. 91); humeral angles marked

(macropterous morph) or weakly marked (micropterous

morph); punctation coarse, moderately dense, and not

arranged in series; interstices without microsculpture and

glossy. Hind wings reduced (micropterous morph) or ful-

ly developed (macropterous morph).

Abdomen broader than elytra; punctation fine and

dense; posterior margin of tergite VII with palisade fringe;

tergite VIII without sexual dimorphism, posterior margin

weakly convex in both sexes.

&: sternite VII with median impression posteriorly, on

either side of this impression with sparse, weakly modi-

fied, somewhat darker and longer setae, posterior margin

broadly and distinctly concave (Fig. 92); sternite VIII

transverse, with broad median impression, this impression

with approximately 30-40 modified, stout and short black

setae, posterior excision broad and rather shallow, on ei-

ther side of this excision with tuft of long black setae (Fig.

93); aedeagus approximately | mm long or nearly so; ven-

tral process blade-shaped, slender and apically acute in lat-

eral view; apex of acutely triangular shape in ventral view

(Figs 94-95).

Comparative notes. Lobrathium wittmeri belongs to a

group of species distributed from Nepal to Sichuan and

Yunnan, represented in the Himalaya by L. wittmeri and

L. kosiense, and characterised as follows: body small or

of moderate size; coloration more or less uniformly black-

ish, elytral spots absent; elytra and hind wings reduced or

dimorphic; eyes mostly of reduced size; sternite VII mod-

erately transverse, relatively weakly modified, without dis-

tinctly modified setae; sternite VIII with small posterior

excision, on either side of this excision with tuft of black

setae; aedeagus symmetric and with more or less blade-

shaped ventral process. The Chinese representatives of this

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78 Volker Assing

96 97

100

101 99

Figs 90-101. Lobrathium wittmeri (90-95) and L. alticola (holotype of L. pirpanjalense) (96-101). 90-91, 97: forebody (91:

macropterous female); 92, 98: male sternite VII; 93, 99: male sternite VIII; 94-95, 100-101: aedeagus in lateral and in ventral

view; 96: habitus. Scale bars: 90-91, 96-97: 1.0 mm; 92-95, 98-101: 0.5 mm.

group are L. cholaicum, L. daxuense, L. domenoides, L.

bimembre, L. lamellatum, and L. duplex.

Distribution and natural history. The known distribu-

tion of L. wittmeri is confined to central Nepal (Fig. 41),

where it was found at altitudes of 2100-3100 m. Howev-

er, since the species is wing-dimorphic it may be wide-

spread in the Himalaya. The specimens from Baglung

were scraped from crevices in a vertical loamy wall (Klee-

berg, pers. comm. ).

Bonn zoological Bulletin 61 (1): 49-128

Lobrathium alticola (Cameron, 1921) (Figs 66, 96-101)

Lathrobium (Lobrathium) alticola Cameron, 1943: 36.

Lobrathium pirpanjalensis [sic] Coiffait, 1982a: 87 f.; syn. n.

Type material examined. L. alticola: Holotype 9: “Ka-

shmir, Gulmarg, vi-vii-31, Dr. Cameron / L. alticola Cam.

Type / Type / M.Cameron. Bequest. B.M. 1955-147. / Ho-

lotypus 9 Lathrobium alticola Cameron, rev. V. Assing

2011 / Lobrathium alticola (Cameron), det. V. Assing

2011” (BMNH).

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Revision of East Palaearctic Lobrathium 79

L. pirpanjalense: Holotype @: “India: Kashmir: Tang-

marg, Pir Panjal Gebirge, 2600 m, 21.—25.V.1976,

Martens & Schawaller leg. / Type / Holotypus / Lobrathi-

um pirpanjalensis H. Coiffait 1979 / Senckenberg-Muse-

um Frankfurt/Main / Lobrathium alticola (Cameron), det.

V. Assing 2012” (SF).

Comment. The original description of L. alticola is based

on a unique female holotype from “Kashmir: Gulmarg, al-

titude 8000—10,000 feet” (Cameron 1943).

Lobrathium pirpanjalense was described from a single

male holotype collected in “Inde, Cachemire, Tangmarg,

Pir Panjal Gebirge” (Coiffait 1982a) without any reference

to L. alticola, although the type localities of both names

are separated by a distance of less than 10 km. A compar-

ison with the holotype of L. alticola yielded no differences

whatsoever suggesting that L. pirpanjalense should rep-

resent a distinct species. Hence the synonymy proposed

above.

Redescription. Body length 5.1—5.8 mm; length of fore-

body 2.9 mm. Habitus as in Fig. 96. Coloration: body

blackish; legs, except for the paler tarsi and antennae

brown to dark-brown; maxillary palpi dark-yellowish.

Head (Fig. 97) approximately 1.05 times as long as

broad; posterior angles abruptly rounded, almost marked;

punctation dense and coarse, median dorsal portion some-

what less densely punctate; interstices without mi-

crosculpture. Eyes small, approximately 1/4 the length of

postocular region in dorsal view.

Pronotum (Fig. 97) approximately 1.2 times as long as

broad and 0.92—0.94 times as wide as head, lateral mar-

gins weakly convex in dorsal view; punctation slightly less

coarse and somewhat less dense than that of head.

Elytra short, 0.75 times as long as, and only slightly

wider than pronotum, humeral angles weakly marked (Fig.

97); punctation moderately coarse and dense, not arranged

in series; interstices without microsculpture. Hind wings

reduced.

Abdomen slightly broader than elytra; punctation dis-

tinct, moderately coarse, and moderately dense; posteri-

or margin of tergite VII without palisade fringe.

S: posterior margin of tergite VIII convexly produced

in the middle; sternite VI with shallow median impression

posteriorly; sternite VII strongly transverse, with pro-

nounced median impression posteriorly, this impression

without pubescence, except for two isolated posterior se-

tae, but margined by dense black setae, anteriorly with

short process (Fig. 98); sternite VIII moderately transverse,

with pronounced median impression posteriorly, this im-

pression with moderately modified setae, posterior exci-

sion moderately deep and moderately broad (Fig. 99);

aedeagus 0.85 mm long, with dorsally angled (lateral

view) and apically acute ventral process (Figs 100-101).

Bonn zoological Bulletin 61 (1): 49-128

©: posterior margin of tergite VIII produced, obtusely

angled in the middle.

Distribution and natural history. This species is current-

ly known only from Kashmir (Fig. 66). The specimens

were collected at altitudes of 2600 and approximately

3000 m.

Lobrathium kosiense sp. n. (Figs 89, 102—107)

Type material. Holotype ¢ [slightly teneral]: “E. Nepal:

Kosi, Val. Induwa Kola [sic], 2850 m, 15.IV.84, Lobl —

Smetana / Holotypus 4 Lobrathium kosiense sp. n., det.

V. Assing 2011” (MHNG).

Etymology. The specific epithet (adjective) is derived

from the name of the region where the type locality is sit-

uated.

Description. Body length 7.2 mm; length of forebody 3.5

mm. Habitus as in Fig. 102. Coloration (note that the holo-

type is slightly teneral; the coloration of mature specimens

may be darker): head, pronotum, and abdomen blackish-

brown; elytra dark-brown; legs reddish-brown with paler

tarsi; antennae dark-brown.

Head (Fig. 103) almost as broad as long; lateral mar-

gins behind eyes convexly rounded towards neck in dor-

sal view, posterior angles not marked; punctation coarse

and very dense, interstices reduced to narrow ridges; small

patches on median dorsal surface and on frons with sparse

punctation or impunctate. Eyes large, more than half as

long a distance from posterior margin of eye to neck in

dorsal view. Antenna moderately slender, approximately

2.3 mm long.

Pronotum (Fig. 103) approximately 1.25 times as long

as broad and nearly as wide as head, convexly tapering

posteriad in posterior half; punctation dense and coarse;

interstices without microsculpture and glossy, distinctly

narrower than diameter of punctures.

Elytra (Fig. 103) short, 0.77 times as long as pronotum;

humeral angles weakly marked; punctation very coarse

and dense. Hind wings completely reduced.

Abdomen broader than elytra; punctation dense and not

particularly fine on tergites II-VI, somewhat finer and

less dense on tergite VII; interstices with distinct mi-

crosculpture; posterior margin of tergite VII without pal-

isade fringe.

S: posterior margin of tergite VIII indistinctly angled

in the middle; sternite VII strongly transverse and with rel-

atively shallow median impression of triangular shape pos-

teriorly, this impression with weakly modified black se-

tae (Fig. 104); sternite VIII moderately transverse, with

pronounced median impression posteriorly, this impres-

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80 Volker Assing

110

Figs 102-112. Lobrathium kosiense (102-107), L. triste (syntype of L. kashmiricum) (108), and L. biaculeatum (109-112). 102:

habitus; 103, 108: forebody; 104, 109: male sternite VII; 105, 110: male sternite VIII; 106-107, 111-112: aedeagus in lateral and

in ventral view. Scale bars: 102—103, 108: 1.0 mm; 104-107, 109-112: 0.5 mm.

sion with numerous distinctly modified, short and black

setae, posterior excision relatively small and convex an-

teriorly, on either side of this excision with cluster of dense

black setae (Fig. 105); aedeagus approximately 0.95 mm

long, with relatively simple, blade-shaped ventral process

(Figs 106-107).

Bonn zoological Bulletin 61 (1): 49-128

Comparative notes. Based on the modifications of the

male sternites VII and VIII, as well as on the similar mor-

phology of the aedeagus, L. kosiense appears to be close-

ly related to L. wittmeri, from which it is distinguished par-

ticularly by slightly smaller size, paler coloration of the

legs, distinctly coarser punctation of the head and prono-

OZFMK

Revision of East Palaearctic Lobrathium 81

tum, slightly larger eyes, a male sternite VIII with a nar-

rower posterior excision and more numerous modified se-

tae, and by the smaller aedeagus with a ventral process

of different shape. For additional remarks on species group

affiliations see the comparative notes in the section on L.

wittmeri.

Distribution and natural history. The type locality is sit-

uated in the Induwa Khola, to the east of Num, Kosi, east-

ern Nepal (Fig. 89). The slightly teneral holotype was col-

lected at an altitude of 2850 m.

Lobrathium biaculeatum sp. n. (Figs 66, 109-112)

Type material. Holotype <: “Nepal Khandbari Distr. Pass

NE Mangmaya 2300 m 6.1V.84, Smetana & Lob! / Holo-

typus ¢ Lobrathium biaculeatum sp. n., det. V. Assing

2012” (cAss). Paratypes: 1 9: “Nepal, Khandbari District

/ above Sheduva, 3000 m, 31.II[.—1.IV.1982, A. & Z.

Smetana” (cSme); 1 9: “Nepal, Khandbari District / For.

above Ahale [=Ahali], 2400 m, 25.III.82, A. & Z.

Smetana” (cSme).

Etymology. The specific epithet (Latin, adjective) refers

to the two needle-shaped processes of the male sternite

VII, a unique character separating this species from all its

congeners.

Description. Body length 6.6—8.2 mm; length of forebody

3.6-3.9 mm. Coloration: head, pronotum, and abdomen

blackish; elytra uniformly blackish or with indistinctly

paler posterior margins; legs yellowish, with the profemo-

ra, at least the basal portion of the protibiae, the apices of

the meso- and metafemora, and sometimes the bases of

the meso- and metatibiae infuscate; antennae reddish to

brown.

Head weakly oblong or approximately as broad as long;

posterior angles weakly marked; punctation coarse and

dense, more or less distinctly sparser in median dorsal por-

tion. Eyes large and distinctly convex, at least slightly

more than half as long as distance from posterior margin

of eye to neck in dorsal view. Antenna moderately slen-

der, approximately 1.9—2.0 mm long.

Pronotum 1.20—1.25 times as long as broad and approx-

imately as wide as head, weakly tapering posteriad, lat-

eral margins weakly convex; punctation similar to that of

head.

Elytra moderately short, approximately 0.9 times as long

as pronotum; humeral angles marked; punctation coarse

and arranged in oblique, more or less regular series. Hind

wings present, but possibly of reduced length.

Abdomen slightly broader than elytra; punctation dense

and relatively coarse in anterior half of tergite III, gradu-

ally becoming finer and sparser towards the abdominal

Bonn zoological Bulletin 61 (1): 49-128

apex; microsculpture shallow and transverse, sometimes

practically obsolete on anterior tergites; posterior margin

of tergite VII with palisade fringe; posterior margin of ter-

gite VIII indistinctly angled in the middle.

3: sternite VII highly distinctive, strongly transverse,

posterior margin with broad and deep, semi-circular pos-

terior excision, on either side of this incision with long

process (Fig. 109); sternite VIII with extensive median im-

pression without pubescence and with rather deep poste-

rior excision (Fig. 110); aedeagus 1.55 mm long, ventral

process of highly distinctive shape, somewhat asymmet-

ric in ventral view (Figs 111—112).

©: tergite X without evident modifications.

Comparative notes. Lobrathium biaculaeatum is readi-

ly identified based on the conspicuous morphology of the

male sternite VII and of the aedeagus. It is additionally

distinguished from L. wittmeri by the much coarser punc-

tation of the forebody, the larger and more bulging eyes,

the seriate punctation of the elytra, the paler legs, and the

shape of the elytra (L. wittmeri: shorter and with less pro-

nounced humeral angles in the micropterous morph,

longer in the macropterous morph).

Distribution and natural history. The type specimens

were collected in three localities in Khandbari district,

eastern Nepal (Fig. 66), at altitudes of 2300-3000 m. Since

the two paratypes are females, the possibility that they in

fact refer to different species cannot be ruled out with ab-

solute certainty, but no convincing evidence was found

that they should not be conspecific with the holotype.

Lobrathium bicarinatum sp. n. (Figs 89, 113-118)

Type material. Holotype @: “India: Uttaranchal state, ca.

30 km N of Bageshwar, Khati vill. env., 2100-2300 m,

27.30. V1.2003, Z. Kejval & M. Tryzna/ Holotypus 3 Lo-

brathium bicarinatum sp. n., det. V. Assing 2011” (cAss).

Paratypes: 304, 229: same data as holotype (cSch,

cAss); 19: “Sikles-Mts., upp. Khilang & Chipli, 1.8.95,

23-2500 m / Nepal Annapurna Mts., leg. Fabrizi, Jager,

Schmidt” (SDEI).

Etymology. The specific epithet (Latin, adjective: with

two carinae) refers to the shape of the male sternite VIII.

Description. Body length 7.0—-8.7 mm; length of forebody

4.2-4.8 mm. Habitus as in Fig. 113. Coloration: whole

body blackish, except for the dark-reddish tarsi and the

reddish-yellow posterior margin of tergite VII.

Head (Fig. 114) approximately as long as broad; pos-

terior angles broadly rounded; punctation very dense, with

the interstices reduced to narrow ridges, and moderately

coarse, median dorsal portion and frons sparsely punctate

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82 Volker Assing

Figs 113-124. Lobrathium bicarinatum (113-118) and L. cylindricolle, lectotype (119-124). 113, 119: habitus; 114, 120: forebody;

115, 121: male sternite VII; 116, 122: male sternite VII; 117-118, 123-124: aedeagus in lateral and in ventral view. Scale bars:

113-114, 119-120: 1.0 mm; 115-118, 121-124: 0.5 mm.

or impunctate; interstices without microsculpture. Eyes Pronotum (Fig. 114) approximately 1.15 times as long

moderately large, approximately half as long as distance _ as broad and slightly narrower than head, lateral margins

from posterior margin of eye to neck, or slightly shorter. distinctly converging posteriad in dorsal view; punctation

Antenna 2.5—2.9 mm long. distinctly sparser than that of head.

Bonn zoological Bulletin 61 (1): 49-128 ©ZFMK

Revision of East Palaearctic Lobrathium 83

Elytra (Fig. 114) approximately 0.95 times as long as,

and distinctly broader than pronotum; humeral angles

moderately marked; punctation coarse, dense, and not

arranged in series; interstices without microsculpture and

glossy. Length of hind wings not examined.

Abdomen narrower than elytra; punctation very fine and

moderately dense; posterior margin of tergite VII with pal-

isade fringe; tergite VIII without sexual dimorphism, pos-

terior margin weakly convex, almost truncate, in both sex-

ess

3: sternite VI with shallow median impression poste-

riorly; sternite VII with rather deep median impression

posteriorly, this impression with pair of clusters of mod-

ified, short and stout black setae, posterior margin broad-

ly concave (Fig. 115); sternite VIII weakly transverse, with

extensive and deep median impression, this impression

with numerous modified, short and stout black setae, and

on either side delimited by pronounced carinae, posteri-

or excision of trapezoid shape (Fig. 116); aedeagus 1.5—1.6

mm long, with strongly sclerotized blade-shaped ventral

process (Figs 117-118).

Comparative notes. This species is characterised partic-

ularly by the conspicuous shape and chaetotaxy of the

male sternite VIII, as well as by the morphology of the

aedeagus. It is readily distinguished from some other Hi-

malayan representatives of the genus with a uniformly

blackish body (e.g., L. wittmeri, L. alticola) by much larg-

er size and broader body alone.

Distribution and natural history. The type locality is sit-

uated to the north of Bageshwar in Uttaranchal, North In-

dia (Fig. 89), at an altitude of 2100-2300 m. A female

paratype was collected in the Annapurna range, central

Nepal, at an altitude of 2300-2500 m.

Lobrathium cylindricolle (Cameron, 1924) (Figs 66,

119-124)

Lathrobium (Lobrathium) cylindricolle Cameron, 1924: 194 f.

Type material examined. Lectotype <, present designa-

tion: “Nun Nadi, Dehra Dun. / Dr. Cameron. 15.10.22. /

Lathrobium cylindricolle Cam. / Type H.T. / M.Cameron.

Bequest. B.M. 1955-147. / Lectotypus ¢ Lathrobium cy-

lindricolle Cameron, desig. V. Assing 2011 / Lobrathium

(?) cylindricolle (Cameron), det. V. Assing 2011”

(BMNH).

Comment. The original description of L. cylindricolle is

based on an unspecified number of syntypes, among them

at least one male, from “Dehra Dun District; Nun

Nadi’’(Cameron 1924). A male syntype was located in the

Cameron collection at the BMNH; it is designated as the

lectotype. An examination of this specimen revealed that

Bonn zoological Bulletin 61 (1): 49-128

the generic assignment of L. cylindricolle is somewhat

doubtful. The elytra lack the submarginal carina usually

found in the genus and the pronotum is narrower and more

oblong than is usually the case in the genus. In general,

the facies somewhat resembles that of Pseudobium Mul-

sant & Rey, 1878, but the male primary and secondary sex-

ual characters resemble those of Lobrathium. Until the

generic affiliations of L. cylindricolle are clarified, the

species is tentatively retained in Lobrathium.

Redescription. Body length 6.5 mm; length of forebody

3.7 mm. Habitus as in Fig. 119. Coloration: head and ely-

tra reddish; pronotum dark-reddish; abdomen reddish-

brown, except for the reddish apex (segments VII—X); legs

reddish-yellow; antennae reddish. (Note that, according

to the original description, the pronotum and the abdomen,

except for the red apex, are black.)

Head (Fig. 120) distinctly oblong, 1.13 times as long

as broad; posterior angles moderately marked; punctation

dense and coarse, median dorsal portion more sparsely

punctate; interstices without microsculpture. Eyes large,

slightly more than half the length of postocular region in

dorsal view. Antenna long and slender, 2.5 mm long.

Pronotum (Fig. 120) long and slender, 1.47 times as long

as broad and approximately 0.8 times as wide as head, sub-

parallel, lateral margins straight in dorsal view; puncta-

tion similar to that of head.

Elytra (Fig. 120) long, approximately 1.05 times as long

as, and distinctly broader than pronotum; humeral angles

marked; punctation coarse and dense, not arranged in se-

ries; interstices without microsculpture. Hind wings ful-

ly developed.

Abdomen narrower than elytra; punctation fine and

moderately dense; microsculpture almost obsolete on ter-

gites III-VI and distinct on tergites VII—VIII; posterior

margin of tergite VII with palisade fringe.

¢: sternite VII weakly impressed and with weakly mod-

ified pubescence in the middle, posterior margin broadly

concave (Fig. 121); sternite VII with distinct and broad

median impression, this impression with numerous mod-

ified, short and stout black setae, posterior excision broad-

ly concave and not very deep (Fig. 122); aedeagus 0.8 mm

long, ventral process bifid apically (Figs 123-124).

Distribution. This species is currently known only from

the type locality in Uttaranchal, northern India (Fig. 66).

Lobrathium sp. n. 1

Material examined. 1: “E. Nepal: Kosi, Val. Induwa Kola,

2000 m, 2.1V.84, L6bl — Smetana” (MHNG).

This undescribed species is characterised by large body size

(body length: 10 mm; length of forebody: 4.6 mm), uniformly

blackish body with reddish legs and antennae (except for the in-

fuscate profemora), short elytra (0.7 times as long as pronotum),

and completely reduced hind wings.

OZFMK

84 Volker Assing

Lobrathium sp. n. 2

Material examined. | 9: “377 Taplejung Dist., upper Tamur Val-

ley, below Walungchung Gola, mixed forest, open river bank,

2400-2700 m, 20 May 1988, J. Martens & W. Schawaller leg.”

(SMNS).

This undescribed species is characterised by small body size

(body length: 5.7 mm; length of forebody: 3.0 mm), uniformly

blackish body, short elytra, and reduced hind wings. In general

appearance it is most similar to L. alticola.

Lobrathium sp. n. 3

Material examined. | 9: “Nepal, Khandbari District / “Bakan”

W of Tashigaon, 3200 m, 5.IV.1982, A. & Z. Smetana” (cSme).

This unnamed species 1s characterised by rather large body size

(body length: 9.3 mm; length of forebody: 5.5 mm), uniformly

blackish brown coloration, and a strongly convex female tergite

X with a spine-like posterior extension. As can be inferred from

the morphology of tergite X, this species is not conspecific with

L. biaculeatum.

Lobrathium sp. n. 4

Material examined. | 9: “Nepal Khandbari Distr. Induwa Kho-

la Valley 2000 m, 17.I1V.84 Smetana & L6bl” (cSme).

This species is similar to the preceding species, but slightly

smaller (body length: 8.5 mm; length of forebody: 4.9 mm), and

the female tergite X lacks the posterior spine-like extension. The

legs are completely blackish-brown with pale tarsi.

THE LOBRATHIUM SPECIES OF MAINLAND

CHINA

Key to species

The following key does not account for L. emeiense, whose de-

scription is based on a single female holotype from the Emei

Shan in Sichuan. For a key to the Lobrathium species of Tai-

wan see Assing (2010).

1. Elytra with posterior portion partly or completely yel-

lowish or reddish, mostly with yellowish spots, and

often with bluish to purple hue) ae.snay. tees ce Z

Elytra uniformly dark-brown to blackish ............ 16

2. Elytra with more extensive yellowish coloration pos-

teriorly, at least posterior two fifths completely yel-

lowish'\(Figs 187-188, 194-195) o.. 0. cee dsseenas anaes 3

— Elytra less extensively yellowish or reddish posteri-

orly, usually with more or less defined spots often

leaving the lateral and/or posterior margins blackish

3. : posterior excision of sternite VIII smaller and less

deep (Fig. 197); aedeagus 1.1 mm long, with ventral

process apically spattle-shaped (Figs 198-199).

Shaanxi: Qinling Shan .............0.0.- schuelkei sp. n.

Bonn zoological Bulletin 61 (1): 49-128

G: posterior excision of sternite VIII slightly larger

and somewhat deeper (Fig. 190); aedeagus larger,

1.2-1.3 mm long, ventral process apically with two

long processes (Figs 191—192). Hubei, Beijing (Fig.

193) ccc coe eae eer CCE EEE taureum sp. Nn.

Large species; length of forebody at least 4.9 mm.

Pronotum broad, 1.10—1.20 times as long as broad

(Figs 201, 224). Elytra 0.90—0.95 times as long as

pronotum and without bluish hue. @: sternite VIII

with distinctly modified, short and stout black setae;

sternite VHI with conspicuously deep posterior exci-

sion, its depth at least approximately half the length

of sternite; ventral process of aedeagus ventrally with

Fasp=like:strulctures 22.5 seceeceeee eeeee eee 5

Smaller species; length of forebody 4.6 mm at most.

Pronotum more slender, 1.20—1.35 times as long as

broad, only in one species broader (L. radens). Ely-

tra often longer than pronotum and often with bluish

or purple hue. @: sternite VII in most species with-

out strongly modified setae; sternite VII with less

deep posterior excision, except in L. bispinosum and

Dy, GUD Cr OSU inca esto ook Rk ee 6

Pronotum slightly broader, 1.10—1.15 times as long

as broad (Fig. 224). 4: sternite VII as in Fig. 225; pos-

terior excision of sternite VIII extremely deep, reach-

ing well beyond middle of sternite (Fig. 226); aedea-

gus 1.6 mm long, ventral process with two rasp-like

projections and stoutly blade-shaped (Figs 227-228).

Northern Yunnan: Diancang Shan (Fig. 180)

sitha ds cede a eASltg ia oc ROMER Sh eC eC excisissimum sp. N.

Pronotum slightly less broad, 1.15—1.20 times as long

as broad (Fig. 201). ': sternite VII as in Fig. 202; pos-

terior excision of sternite VIII less deep, approximate-

ly reaching middle of sternite (Fig. 203); aedeagus 1.5

mm long, ventral process with more numerous rasp-

like projections, somewhat more slender and apical-

ly more acute in ventral view (Figs 204—205). East-

ern Guizhou: Leigong Shan (Fig. 153)

1s dle oaipiate Matglajers nate als Mee erers ee CEE eee radens sp. n.

Elytral spots situated in anterior portion of posterior

half of elytra (1.e., at some distance from posterior

margin (Fig. 218). Dorsal surface of head uneven,

with median and lateral impressions. Punctation of

head and pronotum extremely dense (Fig. 218). Rel-

atively large species; length of forebody 4.44.6 mm.

Antennae slender (Fig. 219). 4: sternite VII moder-

ately transverse (Fig. 220); sternite VIII as in Fig. 221;

aedeagus 1.35 mm long, ventral process apically bi-

fid (Figs 222—223). Western Hubei: Daba Shan.

PRESTR AOTC ate ncEa ob ecancanaccoNaenus ablectum sp. n.

Elytral spots situated at or near posterior margin of

elytra. Dorsal surface of head without distinct impres-

sions. Punctation of head and pronotum less dense.

Smaller species, length of forebody usually 4.0 mm

at most, except for L. spathulatum (3.7-4.5 mm).

©ZFMK

Revision of East Palaearctic Lobrathium 85

Male sexual characters different. A reliable identifi-

cation of the following species is possible only based

onthe malesexualicharacterspnccce-s-scseneee es eeece il

Elytra black, without bluish or purple hue, 0.9—1.0

tinaesyas LOM Syas|PLONOLIMleeeee seein -eeceeeeeees seen 8

Elytra usually with, rarely without bluish or purple

hue, 1.0—1.15 times as long as pronotum ........... 12

@: sternite VIII with small posterior excision in asym-

metric position, posterior margin with tooth-like pro-

jection on either side of the excision (Fig. 136-137);

aedeagus approximately 1.5 mm long, ventral process

very long, slender, and apically asymmetric (Figs

138-139). Widespread: Sichuan, Shaanxi, Hubei

(Bi, AKO) ro aieetas canoadontene Ronn nonRaneteaee tortile Zheng

&: sternite VIII with deep posterior excision in sym-

metric position; aedeagus of different morphology 9

&: aedeagus 0.9-1.0 mm long, ventral process long,

slender, apically acute, and very thin at base (Fig.

152); sternite VHUI with very broad and deep posteri-

or excision, on either side of excision with dense pu-

bescence (Fig. 151); sternite VII as in Fig. 150. Cen-

tral Sichuan: Qingcheng Shan (Fig. 153

sodnoGnoes So ace a aan aeeG Aner ane rten erie ear gladiatum Zheng

Malejsexual'characters!diffierent ©-- 22-4. -2.2---:--2-- 10

. 6: sternite VII anteriorly with tubercle (Figs 182,

184); sternite VIII oblong and with U-shaped poste-

rior excision (Fig. 183); aedeagus 1.2 mm long and

with massive ventral process (Figs 185-186). Jiangxi

RTCA) See ee re oes tuberosum sp. n.

3: sternite VII without tubercle; posterior excision of

sternite VII of different shape; aedeagus longer, at

least approximately 1.4 mm long ..................... 11

. G: sternite VII with shallow posterior excision with

pubescence (Fig. 272); sternite VIII with moderate-

ly deep posterior excision and of characteristic

chaetotaxy (Fig. 273); aedeagus 1.4 mm long and with

ventral process of distinctive shape (Figs 274-275).

Northeastern Hubei (Fig. 180) ........demptum sp. n.

¢: sternite VII with more pronounced posterior im-

pression without pubescence (Fig. 176); posterior ex-

cision of sternite VIII much deeper, broader, and of

subtrapezoid shape (Fig. 177); aedeagus longer, 1.5

mm long, ventral process with two tooth-like projec-

tions ventrally (Figs 178-179). Southeastern Guizhou

CEG, WANE aacics® acocicananase tis dseretess bispinosum sp. n.

. Elytra with weak purple hue; posterior spots relative-

ly small, defined, and of circular shape. Pronotum less

oblong, approximately 1.2 times as long as broad

(Figs 206-207). 2: sternite VII moderately transverse

and with moderately modified setae (Fig. 208); ster-

nite VIII not transverse and with moderately deep pos-

terior excision (Fig. 209); aedeagus 1.0 mm long, ven-

tral process with dorsal carina and apically acute (Figs

210-211). Northern Yunnan (Fig. 153)

SER AA Bre Re a ros AS retrocarinatum sp. n.

Bonn zoological Bulletin 61 (1): 49-128

14.

SY,

16.

hfe

18.

Elytra usually with bluish hue; posterior spots usual-

ly larger and/or of different shape or less defined.

Pronotum more oblong, at least approximately 1.25

times as long as broad. @: sternite VII either with

strongly modified short and stout black setae or with-

out modified setae at all; sternite VII and aedeagus

OMGitiereMtshapememeee erent cance cer eea eee: 13

. G: sternite VII with distinctly modified short and stout

black setae (Fig. 144); sternite VIII with deep U-

shaped posterior excision (Fig. 145); aedeagus ap-

proximately 1.2 mm long, ventral process of distinc-

tive morphology (Figs 146-147). Sichuan, Shaanxi,

Jahbloyen (Oo. 15)S)) os ocoocusesaueddeonse hebeatum Zheng

&: sternite VII without distinctly modified setae; ster-

nite VIII with less deep and differently shaped pos-

terior excision; aedeagus of different morphology 14

G: posterior excision of sternite VII small (Fig. 128);

aedeagus approximately 1.0 mm long and shaped as

in Figs 129-132. Widespread and common species:

China, Taiwan, southern Japan (Fig. 140)

seals ateintiteledis siiticlareciezialslobatdle hongkongense (Bernhauer)

3: posterior excision of sternite VII larger and of

broadly triangular shape; aedeagus of different shape.

Species with more restricted distributions ......... 15

: aedeagus of somewhat variable shape and size,

1.3-1.5 mm long, ventral process very slender, api-

cally acute, and weakly asymmetric (Figs 160-165);

sternites VII and VIII as in Figs 156-159. Widespread

in China: Shaanxi, Sichuan, Hubei, Yunnan

(CRAG alt) ee tceeeen s asee sarin cso cereices configens sp. n.

3: aedeagus longer, 1.6—1.8 mm long, ventral process

distinctly asymmetric and apically distinctly dilated

(ventral view) (Figs 170-173); sternites VII and VI-

II as in Figs 168—169. Widespread in China: Sichuan,

Shaanxi, Shanxi, Hubei, Zhejiang (Fig. 180)

PSB SCE SUBCU OU Romie soca Te mOnc acon aRen: spathulatum sp. n.

Largest representative of the genus in China; length

of forebody approximately 6.0 mm; length of anten-

nae approximately 4.0 mm. Head distinctly oblong,

of ovoid shape, and with conspicuously dense and fine

punctation (Figs 229-230). Zhejiang: Tianmushan

Sic sB Oa ASE OS GEC ae EERE So Oe rotundiceps (Koch)

Distinctly smaller species; length of forebody less

than 5.0 mm; length of antennae 3.3 mm at most.

Head of different shape and with coarser punctation

Posterior margin of abdominal tergite VII without pal-

isade fringe. Eyes small, distinctly less than half the

length of distance from posterior margin of eye to

mecksnedorsalivil warrants emer eee ere 18

Posterior margin of abdominal tergite VII at least with

narrow rudiment of a palisade fringe. Eyes often

Lan O STi see selcueiresn ese wate asi elae aoe aes 19

Slightly smaller species; length of forebody 3.4—3.6

mm. Eyes larger, their length more than one third of

©ZFMK

86 Volker Assing

the distance from posterior margin of eye to neck in

dorsal view (Fig. 232). Punctation of head less dense

and somewhat sparser on vertex (Fig. 232). ¢: ster-

nite VII without modified spines (Fig. 238); sternite

VII as in Fig. 239; aedeagus 0.9 mm long, shaped as

in Figs 240-241. Southeastern Tibet (Fig. 141)

olSojtaitinletarnaten eluate clsisetciag eats See See eee cholaicum sp. n.

— Slightly larger species; length of forebody 3.8 mm.

Eyes smaller, their length little more than one fourth

of the distance from posterior margin of eye to neck

in dorsal view (Fig. 266). Punctation of head (except

frons) conspicuously dense; dorsal surface matt (Fig.

266). S: sternite VII with few modified, short and

stout black setae in median impression (Fig. 267);

sternite VII of different chaetotaxy (Fig. 268); aedea-

gus 1.0 mm long, ventral process slightly asymmet-

ric (ventral view), and more slender both in ventral

and in lateral view (Figs 269-270). Southwestern Si-

Chuanc(hige WSO) seaseeseneeen ener lamellatum sp. n.

19. Larger species; length of forebody 4.1—4.8 mm; length

Of antennae 310 =323emniMl sanceecmaec cece emcee ce nee 20

— Smaller species; length of forebody 3.3-4.1 mm;

length of antennae: 19-22 mann. Saeeneeceices tect es 21

20. Forebody dark-brown. Elytra monomorphic. @: ster-

nites VII and VII as in Figs 214-215; aedeagus 1.8

mm long, ventral process long and slender (Figs

216-217). Northeastern Sichuan (Fig. 193)

— Forebody black. Wing-dimorphic species (Figs

254-255). 3: sternites VII and VIII as in Figs

256-257; aedeagus much smaller, 1.1 mm long, ven-

tral process very slender and apically curved in lat-

eral view (Figs 258—259). Yunnan, Sichuan (Fig. 193)

Io? AOR EEC REE aoc eb aD aad aneunawosatts duplex sp. n.

21. Eyes very small, one third as long as distance from

posterior margin of eyes to neck (Fig. 261). Elytra

short, 0.7 times as long as pronotum (Fig. 261). 3:

sternite VII as in Fig. 262; posterior excision of ster-

nite VIII with semi-transparent margins (Fig. 263);

aedeagus small, 0.75 mm long, ventral process broad

in ventral view (Figs 264-265). Southern Sichuan

(Gea ogsal ies 8 0) Wenetann ner eatin ane dear setter domenoides sp. n.

— Eyes at least slightly larger, at least half as long as dis-

tance from posterior margin of eyes to neck, or near-

ly so. Elytra longer, at least approximately 0.80 times

as long as pronotum. Male sexual characters differ-

ent; posterior excision of sternite VII with fully scle-

FOUSER TMARSIMS cis EAR eae eee lene 22

. Wing-dimorphic species (Figs 243-244). 4: aedeagus

approximately 1.0 mm long, ventral process slender

in ventral view (Figs 247-252); sternites VII and

VII as in Figs 245-246. Yunnan (Fig. 141)

Tea visSuin daaein ose Goa ae eee eRe bimembre sp. n.

— Micropterous species (Fig. 237). 4: ventral process

of aedeagus broader in ventral view (Figs 240-241);

Bonn zoological Bulletin 61 (1): 49-128

sternites VII and VII as in Figs 238—239. Sichuan:

Daxue}Shani(Higa80)ieeeeereseeeereree daxuense sp. n.

Lobrathium hongkongense (Bernhauer, 1931) (Figs

125-132, 140)

Lathrobium (Lobrathium) hongkongense Bernhauer, 1931: 127

Lobrathium sibynium Zheng, 1988: 186 f.; syn. n.

Lobrathium ryukyuense Ito, 1996a: 114 ff.; syn. n.

Type material examined. L. hongkongense: Lectotype 3:

present designation: “7 / Hong Kong. J. J. Walker. /

hongkongensis [sic] Bernh. Cotypus / Chicago NHMus,

M.Bernhauer Collection / Lectotypus 4 Lathrobium hong-

kongense Bernhauer, desig. V. Assing 2012 / Lobrathitum

hongkongense (Bernhauer), det. V. Assing 2012 (FMNH).

Paralectotypes: 1 [dissected prior to present study; some-

what damaged]: “Hong Kong. Walker Coll., 93-58. /

British Museum / hongkongensis Brh. Type / hongkon-

gensis Bernh. Typus / Chicago NHMus, M.Bernhauer Col-

lection” (FMNH); 19: “Hong Kong. Walker Coll.,

93-58. / hongkongensis Bernh. Cotypus / Chicago

NHMus, M.Bernhauer Collection” (FMNH).

L. ryukyuense: Paratypes: 14, 19: “Kametsu Tokuno

Shima Is., 29.II[.1966, T. Ito / Paratype Lobrathium

ryukyuense T. Ito. sp. nov. / Lobrathium hongkongense

(Bernhauer), det. V. Assing 2012” (cAss).

Comment. Lathrobium hongkongense was described from

an unspecified number of syntypes, among them at least

one male, from “Hongkong, gesammelt von J. J. Walker”

(Bernhauer 1931). One of the three syntypes in the Bern-

hauer collection at the FMNH, a male in good condition,

is designated as the lectotype. According to the original

description, additional syntypes, now paralectotypes, are

desposited in the collections of the BMNH.

The original description of L. sibynium is based on nu-

merous specimens from Sichuan (Zheng 1988). Type ma-

terial was not examined, but the descriptive details and the

illustrations provided by Zheng (1988), as well as the fact

that material of this widespread species was seen also from

Sichuan leave little doubt that L. sibynium is conspecific

with L. hongkongense.

Lobrathium ryukyuense was described from various lo-

calities in the Ryuku Islands, southern Japan (Ito 1996a).

An examination of two paratypes made available to me

by Tateo Ito revealed that they are conspecific with the

holotype of L. hongkongense. Hence the synonymy pro-

posed above.

For illustrations of the sexual characters of males from

Taiwan (as L. sibynium) see Assing (2010).

Additional material examined. China: Zhejiang: 17 exs. [part-

ly teneral], Gutianshan National Nature Reserve, 13.—18.III.2008,

leg. ABmann (cFel, cAss, cSch). Jiangsu: 14, 19, Zhenjiang

[“Chinkiang”] (FMNH, cAss). Sichuan: 192, 60 km W Cheng-

©ZFMK

Revision of East Palaearctic Lobrathium 87

Pe 2) Be a) ea) ae

Mathes

125 131 132

135

133 134 138 139

128 137

127

Figs 125-139. Lobrathium hongkongense (125-132; 130-131: lectotype) and L. tortile (133-139). 125, 133: habitus; 126, 134:

forebody; 127, 135: male sternite VII; 128, 136-137: male sternite VIII; 129-132, 138: aedeagus in lateral and in ventral view;

139: apex of ventral process of aedeagus in ventral view. Scale bars: 125-126, 133-134: 1.0 mm; 127-132, 135-138: 0.5 mm;

139: 0.2 mm.

Bonn zoological Bulletin 61 (1): 49-128 ©Z7FMK

88 Volker Assing

Fig. 140. Distributions of Lobrathium guttula (open circle), L. discrepans (filled circle), L. ablectum (open square), and L. hongkon-

gense (triangles), based on revised male-based records (black triangles), revised female-based records (white triangles), and liter-

ature records by Zheng (1988) (grey triangles).

du, 1000 m, V.1997, leg. Reuter (cFel); 19, Daxue Shan, river

valley 5 km E Kangding, 30°03’N, 102°00’E, 2500-2800 m,

20.—23.V.1997, leg. Wrase (cSch). Yunnan: 8 exs., 100 km W

Kunming, Diaolin Nat. Res., 22.V—2.V1.1993, leg. Jendek &

Sausa (NHMW, cAss); 14, Baoshan Pref., Gaoligong Shan, 29

km ESE Tengchong, 24°56’N, 98°45’E, 2350 m, degraded de-

ciduous forest with shrubs, sifted, 1.VI.2007, leg. Wrase

(cAss); 13, 22°, Baoshan Pref., Gaoligong Shan, 33 km SE

Tengchong, 24°51’N, 98°46’E, 2100-2200 m, creek bank in pas-

ture, under gravel and stones, 31.V.2007, leg. Schiilke & Wrase

(cSch, cAss); 14, Nujiang Lisu Aut. Pref., Gongshan Co.,

Gaoligong Shan, tributary of Salween river, 1 km above Gong-

shan, 27°46’N, 98°39’E, 1500 m, old flood debris, 20.V1.2005,

leg. Schiilke (cSch); 23:4, 399 [partly teneral], Nujiang Lisu

Aut. Pref., Gongshan Co., 17 km N Gongshan, small creek val-

ley, 27°55°N, 98°40’E, 1525-1600 m, creek bank, 20.VI.2005,

leg. Schiilke (cAss); 14, Baoshan Pref., mountain range 22 km

S Tengchong, 24°49’N, 98°29’E, 1750 m, loamy banks of fish

ponds, 2.VI.2007, leg. Wrase (cSch); 14', Dali Bai Aut. Pref.,

Diancang Shan, E pass, 44 km NW Dali, 26°02’N, 99°57’E,

2110 m, loamy river bank with gravel and decaying debris,

23.VITI.2009, leg. Wrase (cSch); 12, Yanmen, 13.—23.VI.2005,

leg. Kucera (cSch). Guangxi: 22 exs., 10 km N Liuzhou,

150-200 m, 11.X1.1993, leg. Schillhammer (NHMW, cAss).

Hubei: 16 exs. [partly teneral], Dabie Shan, Wujianshan Forest

Park, 31.1°N, 115.8°E, 17.-18.V1.2003, leg. Turna (NHMW,

cAss). Shaanxi: 19, Lueyang, 23.-26.VI.2004, leg. Kuéera

(cSch); 19, ca. 35 km S Xian, Nan Wutai, 19.[X.1980, leg. Ham-

mond (BMNH).

Bonn zoological Bulletin 61 (1): 49-128

Redescription. Body length 6.3—7.3 mm; length of fore-

body 3.4—-3.9 mm. Habitus as in Fig. 125. Coloration:

head, pronotum, and abdomen black; elytra of very vari-

able coloration, black, mostly with weak bluish hue, pos-

teriorly with pale-reddish to dark-reddish spot of variable

size: small, transverse, weakly delimited, and reaching ne1-

ther posterior margins nor suture in specimens from Tai-

wan, larger, subcircular, well-delimited, often reaching

posterior margin, but not suture and lateral margins in

specimens from mainland China; legs blackish, with red-

dish tibial bases and tarsi; antennae brown with infuscate

antennomere I.

Head (Fig. 126) 1.00—1.10 times as long as broad, usu-

ally widest across eyes, somewhat narrowed behind eyes;

posterior angles broadly rounded; punctation of dorsal sur-

face coarse and moderately dense, in median dorsal por-

tion somewhat sparser (but not impunctate); interstices

without microsculpture. Eyes large and prominent, of

somewhat variable size, slightly to distinctly more than

half as long as distance from posterior margin of eye to

neck. Antenna |.7—2.0 mm long.

Pronotum (Fig. 126) slender, 1.25—1.30 times as long

as broad and slightly narrower than head, lateral margins

very weakly convex, subparallel or weakly converging

posteriad in dorsal view; punctation similar to that of head.

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Revision of East Palaearctic Lobrathium 89

Fig. 141. Distributions of Lobrathium species in China: L. cholaicum (open circle); L. bimembre (filled diamonds); L. domenoides

(open square); L. bispinosum (open diamonds); L. tuberosum (filled square); L. rotundiceps (filled circle); L. tortile (triangles),

based on revised records (filled triangles) and literature records by Zheng (1988) (open triangles).

Elytra (Fig. 126) long, usually at least slightly longer

than pronotum; humeral angles marked; punctation

coarse, dense, and arranged in somewhat irregular series;

interstices without microsculpture and glossy. Hind wings

fully developed.

Abdomen usually slightly narrower than elytra; punc-

tation fine and dense; interstices with fine microsculpture;

posterior margin of tergite VII with palisade fringe; ter-

gite VIII with sexual dimorphism.

6: tergite VIII broadly convex posteriorly; sternite VII

with moderately deep median impression without pubes-

cence posteriorly, posterior margin broadly concave (Fig.

127); sternite VIII weakly transverse, posteriorly with dis-

tinct median impression, this impression with numerous

modified, short and stout black setae, posterior excision

relatively small, with truncate anterior margin (Fig. 128);

aedeagus approximately 1 mm long, ventral process blade-

shaped, symmetric, bisinuate and dorso-ventrally flattened

in lateral view (Figs 129-132).

®: posterior margin of tergite VIII convexly produced

in the middle.

Intraspecific variation. This widespread species is high-

ly variable, more so than most other East Palaearctic con-

geners, particularly regarding the coloration and length of

the elytra, head shape, and eye size.

Bonn zoological Bulletin 61 (1): 49-128

Comparative notes. Lobrathium hongkongense is distin-

guished from its congeners particularly by the shape and

chaetotaxy of the male sternite VIII, as well as by the mor-

phology of the aedeagus, from some Chinese species al-

so by the shape of the female tergite VIII.

Distribution and natural history. This species was orig-

inally described Hong Kong and subsequently reported

from Sichuan and Taiwan (as L. sibynium) (Assing 2010,

Zheng 1988). The above records from Zhejian, Jiangsu,

Yunnan, Guangxi, and Hubei, as well as the localities of

the types of L. ryukyuense suggest that L. hongkongense

is widespread and not uncommon in the southeast of the

East Palaearctic (Fig. 140). The female-based records from

Shaanxi require confirmation. For additional records from

the Ryukyu Islands (as L. ryukyuense) see Watanabe

(1998a) and Ito (2010). Teneral specimens were found in

March and June.

Lobrathium tortile Zheng, 1988 (Figs 133-140)

Lobrathium tortile Zheng, 1988: 187 ff.

Comment. The original description is based on 57 type

specimens from two localities in Sichuan (Zheng 1988).

The types were not examined, but, based on the illustra-

©OZFMK

90 Volker Assing

142

148 149

152

151

Figs 142-152. Lobrathium hebeatum (142-147) and L. gladiatum (148-152). 142, 148: habitus; 143, 149: forebody; 144, 150:

male sternite VI; 145, 151: male sternite VIII; 146-147, 152: aedeagus in lateral and in ventral view. Scale bars: 142-143, 148-149:

1.0 mm; 144-147, 150-152: 0.5 mm.

tions and descriptive details provided by Zheng (1988),

there is no doubt that the material listed below is conspe-

cific with the holotype.

Material examined. China: Sichuan: 53.4, 39 9, Daxue Shan,

W env. Kangding, 30°03’N, 101°57’E, 2600-2700 m,

22.-24.V.1997, leg. Wrase (cSch, cAss); 63:4, 22 9, same da-

ta, but 2700-2800 m, 24.V.1997, leg. Schiilke (cSch, cAss); 13,

Ya’an Pref., Tianquan Co., Jiajin Shan, 54 km W Ya’an, valley

below Labahe N. R. St., 30°03’N, 102°27’E, 1500 m, river bank,

Bonn zoological Bulletin 61 (1): 49-128

12.VII.1999, leg. Wrase (cSch); 14, Ya’an Pref., Baoxing Co.,

Jiajin Shan, 78 km NNW Ya’an, river valley 3 km S Qiaoqi,

30°40°N, 102°45’E, 1950 m, river bank, 11.VII.1999, leg. Wrase

(cSch); 14, 12, Ganzi Tibet. Aut. Pref., Luding Co., 7 km S Lud-

ing, tributary of Dadu He, 29°53’N, 102°13’E, 1250 m, river

bank, 21.VI.1999, leg. Wrase (cSch); 14, Daxue Shan, Hailu-

ogou Glacier Park, Camp 1, 29°36’N, 102°04’E, 2100 m,

27.-31.V.1997, leg. Schiilke (cAss). Shaanxi: 52.4, 19, Qin-

ling Shan, 115 km WSW Xi’an, river bank above Houzhenzi,

33°50’N, 107°47’E, 1450 m, gravel bank, 15.VI.2001, leg.

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Revision of East Palaearctic Lobrathium 9]

Schiilke & Wrase (cSch, cAss). Hubei: 3¢'\¢ [2 teneral], Daba

Shan, creek valley 8 km NW Muyuping, 31°29’N, 110°22’E,

1540 m, stream bank, 18.VII.2001, leg. Schiilke & Wrase (cSch);

cEnAn

544, 22° [4 teneral], same data, but 1550-1650 m (cSch).

Redescription. Body length 6.5—7.5 mm; length of fore-

body 3.4-3.9 mm. Habitus as in Fig. 133. Coloration:

head, pronotum, and abdomen black; elytra black, with-

out bluish hue, posteriorly with reddish spot of variable

size and coloration, moderately large, bright-reddish, and

distinctly contrasting with the remainder of elytral surface

to small, dark-reddish and weakly delimited; legs dark-

brown to blackish, with reddish tarsi; antennae dark-brown

with infuscate antennomere I.

Head (Fig. 134) approximately as long as broad, usu-

ally slightly narrowed behind eyes; posterior angles broad-

ly rounded; punctation of dorsal surface coarse and mod-

erately dense, in median dorsal portion somewhat spars-

er (but not impunctate); interstices without microsculpture.

Eyes moderately large and prominent, usually slightly

more than half as long as distance from posterior margin

of eye to neck. Antenna 1.7—2.0 mm long.

Pronotum (Fig. 134) approximately 1.25 times as long

as broad and approximately as wide as head, lateral mar-

gins straight in the middle and weakly converging poste-

riad in dorsal view; punctation similar to that of head or

slightly coarser.

Elytra (Fig. 134) moderately long, 0.90—0.95 times as

long as pronotum; humeral angles marked; punctation

coarse, dense, and arranged in somewhat irregular series;

interstices without microsculpture and glossy. Hind wings

fully developed.

Abdomen approximately as wide as elytra; punctation

fine and dense; interstices with fine microsculpture; pos-

terior margin of tergite VII with palisade fringe; tergite

VIII without sexual dimorphism, posterior margin weak-

ly convex to indistinctly angled in the middle.

&: sternites V-VI unmodified; sternite VII with shal-

low median impression with sparse pubescence posteri-

orly, posterior margin broadly concave (Fig. 135); stern-

ite VIII weakly transverse, posteriorly with distinct me-

dian impression, this impression with numerous modified,

short and stout black setae, posterior margin of distinc-

tive shape: distinctly asymmetric and with tooth-like pro-

jection on either side of the small sub-median excision

(Figs 136-137); aedeagus approximately 1.5 mm long,

ventral process conspicuously long, asymmetric, and of

distinctive shape (Figs 138-139).

Comparative notes. This species 1s characterised partic-

ularly by the shape of the male sternite VIII and the con-

spicuous morphology of the ventral process of the aedea-

gus. It is additionally distinguished from the widespread

L. hongkongense by the coloration (absence of a bluish

hue) and length of the elytra (distinctly shorter than prono-

tum).

Bonn zoological Bulletin 61 (1): 49-128

Distribution and natural history. This species is current-

ly known from Sichuan, Shaanxi, and Hubei provinces

(Fig. 140), where it was found on banks of rivers and

streams at altitudes of 1250-2800 m. Several specimens

collected in July are teneral.

Lobrathium hebeatum Zheng, 1988 (Figs 142—147, 153)

Lobrathium hebeatum Zheng, 1988: 189 f.

Comment. The original description is based on 16 type

specimens from four localities in Sichuan (Zheng 1988).

The types were not examined, but, based on the illustra-

tions and descriptive details provided by Zheng (1988),

there is no doubt that the material listed below 1s conspe-

cific with the holotype.

Material examined. China: Sichuan: 5¢\3, 19, Daxue Shan,

river valley 5 km E Kangding, 30°03’N, 102°00’E, 2500-2800

m, 20.-23.V.1997, leg. Wrase (cSch, cAss); 734, 19, Ganzi Ti-

bet. Aut. Pref., Kangding Co., Daxue Shan, stream valley 5 km

E Kangding, 30°03’N, 102°00’E, 2500-2600 m, 24.V1.1999, leg.

Wrase (cSch, cAss); 192, Daxue Shan, 10 km S Kangding,

29°59°N, 101°55’E, 3150 m, river valley, 25.V.1997, leg. Wrase

(cSch). Shaanxi: 14, 1Q [teneral], Qinling Shan, 47 km S Xi’an,

autoroute km 70, mountain W pass, 33°51’N, 108°47’E,

2300-2500 m, 26.—29.VHI.1995, leg. Wrase & Schiilke (cSch).

Yunnan: 1, 19, Dali Bai Nat. Aut. Pref., Diancang Shan, 5

km SSW Dali old town, creek valley above cable car, 25°39’N,

100°08’E, 2800 m, 26.VIII.2003, leg Schiilke (cSch).

Redescription. Body length 6.5—7.5 mm; length of fore-

body 3.54.0 mm. Habitus as in Fig. 142. External mor-

phology as in L. tortile (Figs 142—143), except as follows:

Antenna longer and more slender, 2.0—2.2 mm long.

3: sternites II-VI with long, narrow, and glossy me-

dian keel; sternite VI with large and distinct median im-

pression, this impression with numerous distinctly mod-

ified, short and stout black setae, posterior margin distinct-

ly and broadly concave (Fig. 144); sternite VIII weakly

transverse, with distinct median impression, this impres-

sion with numerous modified, short and stout black setae,

posterior excision conspicuously deep and U-shaped (Fig.

145); aedeagus approximately 1.2 mm long, ventral

process straight, with distinct ventral teeth, and apically

of distinctive shape, particularly in ventral view (Figs

146-147).

Comparative notes. This species is characterised partic-

ularly by the presence of median keels on the male ster-

nites II-VI (unique), the shape and chaetotaxy of the male

sternite VII and VIII, as well as by the distinctive shape

of the ventral process of the aedeagus.

Distribution and natural history. Lobrathium hebeatum

is currently known from Sichuan, Shaanxi, and Yunnan

provinces (Fig. 153), where it was found on banks of rivers

©ZFMK

92 Volker Assing

Fig. 153. Distributions of Lobrathium species in China: L. configens (open diamonds); L. retrocarinatum (filled square); L. glad-

iatum (filled circle); L. radens (open square); L. hebeatum (triangles), based on revised records (filled triangles) and a literature

record by Zheng (1988) (open triangle).

and streams at altitudes of 2500-3150 m, in one locality

together with L. hongkongense. Teneral adults were col-

lected in August.

Lobrathium gladiatum Zheng, 1988 (Figs 148-153)

Lobrathium gladiatum Zheng, 1988: 190.

Comment. The original description is based on a unique

holotype from Qingcheng Shan in Sichuan. This specimen

was not examined, but, based on the illustrations and de-

scriptive details provided by Zheng (1988), there is no

doubt that the material listed below, all of which was col-

lected in the Qingcheng Shan, is conspecific with the holo-

type.

Material examined. China: Sichuan: 34.3, 19 [partly tener-

al], Qingcheng Shan, NW Chengdu, 30°54’N, 103°32’E,

650-700 m, 3.4.VI.1997, leg. Schiilke (cSch); 434, 329,

Wenjiang Distr., Dujiangyan Co., Qingcheng Shan, 56 km NW

Chengdu, 30°54’N, 103°32’E, 975 m, stream bank, 975 m,

18.VI.1999, leg. Schiilke (cSch, cAss).

Description. Body length 5.8—6.8 mm; length of forebody

3.2-3.6 mm. Habitus as in Fig. 148. Coloration: head,

pronotum, and abdomen black; elytra black, without bluish

hue, posteriorly with moderately large reddish-yellow spot

reaching posterior, but not lateral margins; legs reddish-

Bonn zoological Bulletin 61 (1): 49-128

brown with reddish tarsi, usually at least the profemora

infuscate; antennae dark-reddish, antennomere I not infus-

cate.

Head (Fig. 149) weakly oblong, widest across eyes, nar-

rowed behind eyes; posterior angles broadly rounded;

punctation of dorsal surface coarse and moderately

dense, in median dorsal portion somewhat sparser (but not

impunctate); interstices without microsculpture. Eyes large

and prominent, slightly more than half as long as distance

from posterior margin of eye to neck. Antenna 1.9—2.0 mm

long.

Pronotum (Fig. 149) slender, 1.30—1.35 times as long

as broad and slightly narrower than head, lateral margins

indistinctly convex in the middle and weakly converging

posteriad in dorsal view; punctation similar to that of head.

Elytra (Fig. 149) moderately long, approximately as

long as pronotum; humeral angles marked; punctation

coarse, dense, and arranged in somewhat irregular series;

interstices without microsculpture and glossy. Hind wings

fully developed.

Abdomen usually slightly narrower than, more rarely

approximately as wide as elytra; punctation fine and dense;

interstices with fine microsculpture; posterior margin of

tergite VII with palisade fringe; tergite VIII without pro-

nounced sexual dimorphism, posterior margin on average

slightly more strongly convex in female than in male.

©OZFMK

Revision of East Palaearctic Lobrathium 93

<: sternites V and VI with shallow median impressions;

sternite VII with pronounced, deep and extensive medi-

an impression (Fig. 150); sternite VIII with pronounced

impression posteriorly, on either side of impression and

of posterior excision strongly bulging, almost carinate,

posterior excision very deep and broad (Fig. 151); aedea-

gus 0.9—1.0 mm long, ventral process with very thin base

and with pronounced median carina dorsally (Fig. 152).

Comparative notes. This species is characterised partic-

ularly by the shape and chaetotaxy of the male sternite VI-

II and the conspicuous morphology of the ventral process

of the aedeagus. It is additionally distinguished from the

externally similar L. hebeatum and L. tortile by the more

slender head and pronotum, and by the longer elytra.

Distribution and natural history. This species 1s current-

ly known only from the Qingcheng Shan in central

Sichuan province, China (Fig. 153). The recently collect-

ed material was found at an altitude of 650-975 m, the

holotype at 1600 m. Teneral adults were observed in June.

Lobrathium emeiense Zheng, 1988

Lobrathium emeiense Zheng, 1988: 191.

Comment. The original description is based on a unique

female holotype from the Emei Shan, Sichuan. Since a re-

liable identification and interpretation of the vast major-

ity of Lobrathium species is possible only based on the

male sexual characters, the identity of this species must

remain doubtful at least until males from the vicinity of

the type locality become available.

Lobrathium configens sp. n. (Figs 153-165)

Type material. Holotype ¢: “China (S-Shaanxi) Qinling

Shan, river bank above Houzhenzi, 115 km WSW Xi’an,

1450 m, 33°50’N, 107°47°E (coarse gravel bank with

plants and leaves) 4.VII.2001 Wrase [06a] / Holotypus 2

Lobrathium configens sp. n., det. V. Assing 2011”

(NHMW). Paratypes: 24:3, 229 [partly teneral]: “Chi-

na: Sichuan Gongga Shan, Hailuogou, in front of Glacier

1, 2850 m, 29°35N, 102°00E, 7.VII.1998, A. Smetana

[C76] / 1998 China Expedition J. Farkaé, D. Kral, J.

Schneider & A. Smetana” (cSme, cAss); 1¢ [slightly ten-

eral]: “China W.Sichuan (Ganzi Tibet. Aut. Pref., Yajiang

Co.), Shalui Shan, river valley 6 km WSW Yajiang, 3250

m, 30°01N, 100°57E (river bank, bank slope), 4. VII.1999

D.W. Wrase” (cSch); 1¢ [slightly teneral]: “China: S-

Sichuan 1999, Ya’an Prefecture, Shimian Co., Xiaoxiang

Ling, Pass zw. Shimian u. Ganluo, 27 km SE Shimian,

29°03N, 102°31E, 2450 m, Quellsumpf, Bachufer, 8.VII.,

leg. M. Schiilke (cSch); 19: “China S.Sichuan (Ya’an

Bonn zoological Bulletin 61 (1): 49-128

Pref., Shimian Co.) Xiaoxiang Ling, road Shimian-Gan-

luo, 20 km SE Shimian (bank limit), 29°05N, 102°29E,

1850 m, 8.VII.1999 D.W. Wrase” (cSch); 404, 399:

“China: W-Hubei, 2002, Dashenongyjia mts., 2100-2900

m, 10.—14.6., 31.5N, 110.3E, leg. J. Turna” (NHMW,

cAss); 1¢: “China, Yunnan prov., 18.6.—4.7.1993,

Heishui=35 km N Lijiang, 27,13 N, 100,19 E, lgt. S. Bec-

var” (NHMW); 14, 59 [partly teneral]: “China, Yun-

nan prov., 1.-19.7.1992, Heishui, 35 km N Lijiang,

27,13N, 100,19E, lgt. S. Becvar” (NHMW, cAss); 19:

same data, but 27.-28.VI.1992 (NHMW); 2¢'¢ [1 tener-

al]: “China (N-Yunnan), Zhongdian Co., 16 km SSE

Zhongdian, 3100 m, 27°40.7’N, 99°°44.2’E (river bank,

gravel), 17. VIII.2003 Wrase [06]” (cSch); 14, 39 9: “Chi-

na (N-Yunnan) Diqing Tibet. Aut. Pref., Deqin Co., Meili

Xue Shan, E-side, 14 km W Deqin, 28°27.47°N,

98°46.35°E, 2700 m / small creek valley, glacier outlet

(under stones and gravel along glacier stream) 11.V1I.2005

D.W.Wrase [09A]” (cSch, cAss).

Etymology. The specific epithet (Latin, present participle

of configere: to sting) refers to the long and acute ventral

process of the aedeagus.

Description. Body length 6.0—7.2 mm; length of forebody

3.64.0 mm. Habitus as in Fig. 155. Coloration: head,

pronotum, and abdomen black; elytra black with bluish

hue, posterior third with yellowish spot reaching posteri-

or and lateral margins; legs blackish with dark-reddish tar-

si; antennae dark-brown.

Head (Fig. 156) approximately as long as broad, widest

across eyes, somewhat narrowed behind eyes; posterior

angles broadly rounded; punctation of dorsal surface

coarse and moderately dense, in median dorsal portion

somewhat sparser (but not impunctate); interstices with-

out microsculpture. Eyes large and prominent, more than

half as long as distance from posterior margin of eye to

neck. Antenna 2.0—2.2 mm long.

Pronotum (Fig. 156) slender, approximately 1.3 times

as long as broad and 0.90—0.95 times as broad as head,

lateral margins almost straight and weakly converging pos-

teriad in dorsal view; punctation similar to that of head.

Elytra (Fig. 156) long, approximately 1.1 times as long

as, and distinctly broader than pronotum; humeral angles

marked; punctation coarse, dense, and arranged in some-

what irregular series; interstices without microsculpture

and glossy. Hind wings fully developed.

Abdomen distinctly narrower than elytra; punctation

fine and dense; posterior margin of tergite VI with pal-

isade fringe; tergite VII without sexual dimorphism, pos-

terior margin strongly convex to obtusely pointed in both

Sexes.

S: sternite V with small and very shallow median im-

pression posteriorly; sternite VI with shallow median im-

pression not confined to posterior half; sternite VII with

OZFMK

94 Volker Assing

162

163

164

159

Figs 154-165. Lobrathium configens. 154: habitus; 155: forebody; 156: male sternite VII; 157-159: sternite VIII of males from

Shaanxi (157), Hubei (158), and Yunnan (159); 160-165: aedeagus in lateral and in ventral view of males from Shaanxi (160-161),

Hubei (162-163), and Yunnan (164-165). Scale bars: 154-155: 1.0 mm; 156-165: 0.5 mm.

moderately deep median impression posteriorly, this im-

pression without pubescence, posterior margin broadly

concave (Fig. 156); sternite VIII weakly transverse, pos-

teriorly with median impression of triangular shape, this

impression with numerous modified, short and stout black

setae, posterior excision broad and not very deep, on ei-

Bonn zoological Bulletin 61 (1): 49-128

ther side of this impression with dense black setae (Figs

157-159); aedeagus of variable size, 1.3—1.5 mm long,

ventral process of somewhat variable shape, long, slen-

der, dorso-ventrally flattened, apically acute, almost

symmetric to distinctly asymmetric, and dorso-ventrally

flattened.

©OZFMK

Revision of East Palaearctic Lobrathium 95

Intraspecific variation. The ventral process of the

aedeagus of this species is remarkably variable. This par-

ticularly applies to the degree of asymmetry, the degree

to which it is twisted apically, and to the width of the base

(ventral aspect) (Figs 160—165). Also, to a lesser extent,

the chaetotaxy of the male sternite VIII is somewhat vari-

able (Figs 157-159). However, in view of the general char-

acter divergence in Lobrathium species and the presence

of transitional conditions, the observed differences are at-

tributed to intra- rather than interspecific variation.

Comparative notes. In external characters, L. configens

is highly similar to the widespread L. hongkongense, but

distinguished from this species by the male primary and

secondary sexual characters, especially the completely dif-

ferent shape of the ventral process of the aedeagus.

Distribution and natural history. The type specimens

were collected in several localities in Shaanxi, Sichuan,

Hubei, and Yunnan provinces (Fig. 153) at altitudes of

1450-3250 m. At least part of the type specimens was

found on stream banks. The specimens from the Gongga

Shan were found in sandy areas near a river leaving a gla-

cier (Smetana pers. comm.) together with L. daxuense.

Teneral adults were repeatedly observed in July and once

in August. The holotype was collected together with nu-

merous specimens of L. tortile.

Lobrathium spathulatum sp. n. (Figs 166-173, 180)

Type material. Holotype @: “China — Sechuan, Pingwu,

2.6.4.6.2001, lgt. E. Kuéera / Holotypus ¢ Lobrathium

spathulatum sp. n., det. V. Assing 2011” (cAss). Paratypes:

14, 2: “China c. 25.V., Xiexian [=Xiaxian], 111,6’-34,8”

[sic], 20 km NE Yuncheng, leg. J. Halada 1996” (NHMW,

Ass); 1¢ [slightly teneral]: “E-China: Zhejiang, Gutian-

shan National Nature Reserve, 13.—18.III.2008, leg. Th.

Assmann” (cFel); 14: “China-Shaanxi, Liieang,

22.5.-25.5.2000, lgt. E. Kuéera (cFel); 19: “China-

N.Sechuan, Wangcang, 18.—20.5.2002, Igt. E. Kucera”

(cFel); 1¢ [teneral], 19: “China: Sichuan (18),

Qingcheng-Shan, NW Chengdu, 600 m, Flussufer,

30.55N, 103.30E, 4.06.1997, M. Schitilke” (cSch, cAss);

14: “China: W-Hubei (Daba Shan), creek valley 8 km NW

Muyuping, 31°29’N, 110°22’E, 1550-1650 m,

18.VII.2001, leg. M. Schtilke [CO1—16A]” (cSch).

Etymology. The specific epithet (Latin, adjective derived

from the noun spathula: spattle, applicator) alludes to the

shape of the ventral process of the aedeagus.

Description. Body length 6.0—8.0 mm; length of forebody

3.7-4.5 mm. Habitus as in Fig. 166. Coloration: head,

pronotum, and abdomen black; elytra black with bluish

Bonn zoological Bulletin 61 (1): 49-128

hue, posterior third with yellowish spot reaching posteri-

or, but not lateral margins; legs blackish with dark-red-

dish tarsi and reddish-brown tibial bases; antennae dark-

brown with blackish antennnomere I.

Head (Fig. 167) approximately as long as broad and of

variable shape; temples behind eyes subparallel or con-

verging posteriad; posterior angles broadly rounded; punc-

tation of dorsal surface coarse and moderately dense, in

median dorsal portion somewhat sparser (but not impunc-

tate); interstices without microsculpture. Eyes large and

prominent, more than half as long as distance from pos-

Pronotum (Fig. 167) slender, approximately 1.3 times

as long as broad and 0.90—0.95 times as broad as head,

lateral margins almost straight and subparallel in dorsal

view; punctation similar to that of head.

Elytra (Fig. 167) long, approximately 1.1 times as long

as, and distinctly broader than pronotum; humeral angles

marked; punctation coarse, dense, and arranged in some-

what irregular series; interstices without microsculpture

and glossy. Hind wings fully developed.

Abdomen distinctly narrower than elytra; punctation

fine and dense; posterior margin of tergite VII with pal-

isade fringe; tergite VIII without sexual dimorphism, pos-

terior margin strongly convex in both sexes.

4: sternite VI, sometimes also sternites [V—V, with small

median impression posteriorly; sternite VII moderately

transverse, with shallow median impression of triangular

shape posteriorly, this impression without pubescence,

posterior margin broadly and not very deeply concave

(Fig. 168); sternite VIII weakly transverse, with long and

not very extensive median impression, this impression

with numerous modified, stout and short black setae, pos-

terior excision broad and moderately deep, margins of ex-

cision with dense black setae (Fig. 169); aedeagus

1.6—-1.8 mm long, ventral process long, slender, apically

acute, distinctly asymmetric, and somewhat twisted.

Comparative notes. As can be inferred from the similar-

ly derived shape and chaetotaxy of the male sternite VI-

II, from the similar general morphology of the aedeagus,

as well as from the similar external characters, L. spathu-

latum is closely allied to L. configens. It is distinguished

from this species by larger size (no overlap), the smaller

elytral spot (not reaching lateral margins), and by the dis-

tinctly larger aedeagus with a ventral process of somewhat

different shape.

Distribution. Lobrathium spathulatum 1s currently known

from several localities in Sichuan, Shaanxi, Shanxi, Hubei,

and Zhejiang (Fig. 180), suggesting that the species is

widespread at least in China. Two specimens were collect-

ed on a river bank. One teneral male was found in March,

another in early June.

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96 Volker Assing

171 172 173

168 169

Figs 166-179. Lobrathium spathulatum (166-173) and L. bispinosum (174-179). 166, 174: habitus; 167, 175: forebody; 168, 176:

male sternite VII; 169, 177: male sternite VIII; 170-173, 178-179: aedeagus in lateral and in ventral view (170-171: male from

Sichuan; 172-173: male from Shaanxi). Scale bars: 166-167, 174-175: 1.0 mm; 168-173, 176-179: 0.5 mm.

Bonn zoological Bulletin 61 (1): 49-128 ©ZFMK

Revision of East Palaearctic Lobrathium 97

Lobrathium bispinosum sp. n. (Figs 141, 174-179)

Type material. Holotype ¢ [slightly teneral]: “China:

Guizhou, Leishan Co., SE Kaili, 15 km E Leishan,

Leigong Shan, S-slope, 26°22.40’N, 108°08.83’E /

12.6.2001, ca. 1000 m, leg. Schillhammer & Wang

(CWBS 431) / Holotypus ¢ Lobrathium bispinosum sp.

n., det. V. Assing 2011” (NHMW). Paratypes: 39 & [part-

ly slightly teneral]: same data as holotype (NHMW, cAss);

1 ex. [abdomen heavily damaged]: “China: Guizhou, Leis-

han Co., SE Kaili, NE Leishan, Leigong Shan, SE-slope,

26°21.42’N, 108°09.36°E / 21.6.2001, ca. 1200 m, nr. Park

gate, leg. Wang (15)” (NHMW).

Etymology. The specific epithet (Latin, adjective: with

two spines) alludes to the pair of spines on the ventral

process of the aedeagus.

Description. Body length 6.0—6.5 mm; length of forebody

3.5—3.8 mm. Habitus as in Fig. 174. Coloration: body

black, posterior third of elytra with yellowish spot reach-

ing posterior and lateral margins; legs reddish (teneral?)

to blackish with dark-reddish tarsi; antennae reddish-

brown to dark-brown, antennomere | infuscate.

Head (Fig. 175) approximately as long as broad, widest

across eyes, behind eyes convexly tapering towards neck,

posterior angles obsolete; punctation of dorsal surface

coarse and moderately dense, in median dorsal portion

somewhat sparser (but not impunctate); interstices with-

out microsculpture. Eyes large and prominent, slightly

more than half as long as distance from posterior margin

of eye to neck. Antenna 1.9—2.2 mm long.

Pronotum (Fig. 175) slender, approximately 1.35 times

as long as broad and 0.95 times as broad as head, lateral

margins in the middle almost straight and subparallel in

dorsal view; punctation similar to that of head.

Elytra (Fig. 175) moderately long and of somewhat vari-

able length, 0.9—1.0 times as long as, and distinctly broad-

er than pronotum; humeral angles marked; punctation

coarse, dense, and arranged in somewhat irregular series;

interstices without microsculpture and glossy. Hind wings

apparently fully developed.

Abdomen distinctly narrower than elytra; punctation

fine and dense, finer and sparser on posterior than on an-

terior tergites; posterior margin of tergite VII with palisade

fringe; tergite VII without sexual dimorphism, posterior

margin strongly convex to obtusely pointed in both sex-

es.

3: sternites III-VI unmodified; sternite VII strongly

transverse and with distinct median impression of trian-

gular shape and without pubescence, posterior margin dis-

tinctly and broadly concave (Fig. 176); sternite VIII ap-

proximately as broad as long, weakly sclerotised (semi-

transparent) anteriorly, and with pronounced median im-

pression without setae, posterior excision conspicuously

Bonn zoological Bulletin 61 (1): 49-128

deep, broad, and of subrectangular shape, on either side

of this excision and of median impression with dense black

setae (Fig. 177); aedeagus 1.5 mm long, ventral process

symmetric, broad (ventral view), apically rounded (ven-

tral view), and bisinuate (lateral view), in the middle with

pair of pronounced tooth-like processes (Figs 178-179).

Comparative notes. Lobrathium bispinosum is readily

distinguished from all its congeners by the conspicuous

morphology of the male sternite VIII and of the aedea-

gus. From L. hongkongense, L. configens, and L. mordens,

it is additionally separated by the absence of a bluish hue

on the elytra, as well as by the shorter elytra.

Distribution and natural history. The species 1s current-

ly known only from two localities in the Leigong Shan in

the southeast of Guizhou province, southern China (Fig.

141). The specimens were collected at altitudes of 1000

and 1200 m; some of them are teneral.

Lobrathium demptum sp. n. (Figs 180, 271-275)

Type material. Holotype ¢: “China: E - Hubei, Dabie

Shan, Wuyjiashan Forest Park, 31.1 N 115.8 E,

17.-18.6.2003, leg. J. Turna / Holotypus ¢ Lobrathium

demptum sp. n., det. V. Assing 2012” (NHMW). Paratypes:

3, 12: same data as holotype (NHMW, cAss).

Etymology. The specific epithet (past participle of the Lat-

in verb demere: to take away) alludes to the small num-

ber of modified setae on the male sternite VIII.

Description. Body length 6.3—7.0 mm; length of forebody

3.6—3.9 mm. Coloration: body black, without bluish hue,

elytra posteriorly with relatively large yellowish spot

reaching posterior and lateral margins; legs dark-brown

with reddish tarsi; antennae reddish with infuscate anten-

nomere I.

Head (Fig. 271) approximately as long as broad, widest

across eyes, behind eyes convexly tapering towards neck,

posterior angles obsolete; punctation of dorsal surface

coarse and moderately dense, in median dorsal portion

somewhat sparser (but not impunctate); interstices with-

out microsculpture. Eyes large and prominent, slightly

more than half as long as distance from posterior margin

of eye to neck. Antenna 2.0—2.2 mm long.

Pronotum (Fig. 271) moderately slender, approximate-

ly 1.25—1.30 times as long as broad and 0.95—1.00 times

as broad as head, lateral margins weakly convex in the

middle and slightly converging in dorsal view; punctation

similar to that of head.

Elytra (Fig. 271) approximately 0.95 times as long as,

and distinctly broader than pronotum; humeral angles

marked; punctation coarse, dense, and arranged in some-

©ZFMK

98 Volker Assing

Fig. 180. Distributions of Lobrathium species in China: L. excisissimum (filled square); L. lamellatum (open diamond); L. dax-

uense (filled circles); L. spathulatum (open triangles); L. schuelkei (filled diamond); L. demptum (filled triangle).

what oblique series; interstices without microsculpture and

glossy. Hind wings apparently fully developed.

Abdomen distinctly narrower than elytra; punctation

fine and moderately dense; interstices without apprecia-

ble microsculpture and glossy, posterior margin of tergite

VII with palisade fringe; tergite VIII without sexual di-

morphism, posterior margin broadly convex in both sex-

es.

&: sternites II-VI unmodified; sternite VII strongly

transverse and with shallow median impression posteri-

orly, pubescence unmodified, posterior margin distinctly

and broadly concave (Fig. 272); sternite VIII approximate-

ly as broad as long, median impression long and broad,

anteriorly and posteriorly with few modified, short and

stout black setae, posterior excision moderately broad and

not very deep (Fig. 273); aedeagus 1.4 mm long, ventral

process symmetric, large (ventral view) and long, some-

what domed and with short median spine ventrally (Figs

274-275).

Comparative notes. Lobrathium demptum is readily dis-

tinguished from all its congeners particularly by the shape

and chaetotaxy of the male sternites VII and VIII, as well

as by the distinctive morphology of the ventral process of

the aedeagus. From L. hongkongense, L. configens, and

L. mordens, it is additionally separated by the absence of

Bonn zoological Bulletin 61 (1): 49-128

a bluish hue on the elytra, as well as by the shorter ely-

tra.

Distribution and natural history. The type locality is sit-

uated in the northeast of Hubei province, very close to the

border with Anhui province (Fig. 180). The specimens ap-

pear to be slightly teneral. Additional data are not avail-

able.

Lobrathium tuberosum sp. n. (Figs 141, 181-186)

Type material. Holotype ¢@ [teneral]: “China Jiangxi W

Jinggang Shan, Ciping env., 2-14. VI.1994 / Holotypus 3

Lobrathium tuberosum sp. n., det. V. Assing 2011”

(NHMW).

Etymology. The specific epithet (Latin, adjective: with tu-

bercles) alludes to presence of minute tubercles on the

male sternites [V—VII.

Description. Body length 6.0 mm; length of forebody 3.6

mm. In external characters highly similar to L. bispinosum

(Fig. 181), reliably distinguished only by the male sexu-

al characters:

©ZFMK

Revision of East Palaearctic Lobrathium 99

188

187

190

184

189

Figs 181-192. Lobrathium tuberosum (181-186) and L. taureum (187-192). 181, 188: forebody; 182, 189: male sternite VII; 183,

190: male sternite VIII; 184: male sternite VII in lateral view; 185-186, 191-192: aedeagus in lateral and in ventral view; 187:

habitus. Scale bars: 181, 187-188: 1.0 mm; 182-186, 189-192: 0.5 mm.

¢: sternites [V—VI with small sharp median tubercle;

sternite VII moderately transverse and with extensive me-

dian impression, this impression without pubescence in the

middle, anterior to impression with tubercle (Fig. 184),

posterior margin broadly concave (Fig. 182); sternite

VIII oblong, weakly sclerotised (semi-transparent) ante-

riorly, and with pronounced median impression without

Bonn zoological Bulletin 61 (1): 49-128

setae, posterior excision conspicuously deep, broad, and

of subrectangular shape, on either side of this excision and

of median impression with dense black setae (Fig. 183);

aedeagus 1.2 mm long, ventral process symmetric, con-

spicuously massive both in lateral and in ventral view, and

apically rounded (ventral view) (Figs 185-186).

©ZFMK

100 Volker Assing

Fig. 193. Distributions of Lobrathium species in China and the Russian Far East: L. duplex (open circles); L. feldmanni (filled di-

amond); L. taureum (open triangles); L. semiflavum (filled square).

Comparative notes. Lobrathium tuberosum is readily dis-

tinguished from all its congeners by the presence of small

sharp tubercles on the male sternites [V—VII, the conspic-

uous morphology of the male sternite VIII, and by the

characteristic shape of the ventral process of the aedea-

gus. The similarly derived shape and chaetotaxy of the

male sternite VIII, as well as the similar external charac-

ters suggest that L. tuberosum 1s closely allied to L.

bispinosum.

Distribution and natural history. The type locality is sit-

uated in Jiangxi province in southeastern China (Fig. 141).

The holotype is teneral.

Lobrathium taureum sp. n. (Figs 187-193)

Type material. Holotype 4: “China: W-Hubei (Daba

Shan), creek valley 8 km NW Muyuping, 31°29’N,

110°22’E, 1550-1650 m, 18.VII.2001, leg. M. Schiilke

[CO1—16A]/ creek valley, deciduous forest, moss (sifted)

[CO1-16A] / Holotypus 3 Lobrathium taureum sp. n., det.

Bonn zoological Bulletin 61 (1): 49-128

V. Assing 2011” (cAss). Paratypes: 17 exs. [partly slight-

ly teneral]: same data as holotype; 1: “China: W-Hubei

(Daba Shan), creek valley 8 km NW Muyuping, 31°29’°N,

110°22’E, 1540 m, 18.VII.2001, moss sifted, leg. M.

Schiilke [CO01-16]” (cSch); 14: same data, but leg. D.

Wrase (cSch); 604, 19 [1¢ teneral]: China: E-Hubei,

Dabie Shan, Wujiashan Forest Park, 31.1 N 115.8 E,

17.-18.6.2003, leg. J. Turna” (NHMW, cAss); 24°35, 32 2

[previously with type labels “Lobrathium pekingi” by V.

Gusarov attached to pins, but these labels removed; | ten-

eral]: “China: B.M. 1980-491, P.M. Hammond / Peking,

Badaling, 9.1x.80” (BMNH, cAss).

Etymology. The specific epithet (Latin, adjective derived

from taurus: bull) alludes to the conspicuous shape of the

ventral process of the aedeagus.

Description. Body length 6.5—7.8 mm; length of forebody

3.7-4.3 mm. Habitus as in Fig. 187. Coloration: head,

pronotum, and abdomen black; elytra black, usually with

faint bluish hue and with almost all of posterior half com-

pletely (including lateral parts and margins) yellowish;

©ZFMK

Revision of East Palaearctic Lobrathium 101

legs blackish-brown with pale-reddish tarsi; antennae red-

dish to dark-brown, antennomere I infuscate.

Head (Fig. 188) approximately as long as broad, widest

across eyes, behind eyes convexly tapering towards neck,

posterior angles obsolete or nearly so; punctation of dor-

sal surface coarse and rather dense, in median dorsal por-

tion and on frons distinctly sparser; interstices without mi-

crosculpture. Eyes large and prominent, more than half as

long as distance from posterior margin of eye to neck. An-

tenna 2.0—2.3 mm long.

Pronotum (Fig. 188) approximately 1.25 times as long

as broad and approximately as broad as head (or nearly

so), lateral margins in the middle almost straight and sub-

parallel or weakly converging in dorsal view; punctation

similar to that of head, but less dense than in lateral dor-

sal portions of head.

Elytra (Fig. 188) 1.05—1.10 times as long as, and dis-

tinctly broader than pronotum; humeral angles marked;

punctation coarse, dense, and arranged in somewhat irreg-

ular series; interstices without microsculpture and glossy.

Hind wings fully developed.

Abdomen distinctly narrower than elytra; punctation

fine and dense, finer and sparser on posterior than on an-

terior tergites; posterior margin of tergite VII with palisade

fringe; tergite VIII without distinct sexual dimorphism,

posterior margin slightly more convex in female than in

male.

¢: sternites V—VI with shallow median impression pos-

teriorly; sternite VII strongly transverse and with distinct

median impression with pubescence, posterior margin

broadly concave (Fig. 189); sternite VIII transverse, with

pronounced long median impression, this impression with

numerous modified, short and stout black setae, posteri-

or excision deep and almost V-shaped (Fig. 190); aedea-

gus 1.2—1.3 mm long, ventral process symmetric and api-

cally with two long processes (Figs 191-192).

Comparative notes. Lobrathium taureum is characterised

particularly by the coloration pattern of the elytra and by

the distinctive shape of the ventral process of the aedea-

gus. It shares the elytral coloration only with L. schuelkei,

with L. semiflavum (Russian Far East), and with L. par-

titum (Japan).

Distribution and natural history. The species is current-

ly known from the Daba Shan (altitude 1540—1650 m) and

the Dabie Shan in Hubei province, as well as from Bei-

jing province, China (Fig. 193). Teneral specimens were

collected in June, July, and September.

Lobrathium schuelkei sp. n. (Figs 180, 194-199)

Type material. Holotype ¢ [teneral]: “China (S-Shaanxi)

Qinling Shan, river bank above Houzhenzi, 115 km WSW

Bonn zoological Bulletin 61 (1): 49-128

Xian, 1450 m, 33°50’N, 107°47°E (coarse gravel bank

with plants and leaves) 4.VII.2001 Wrase [06a] / Holoty-

pus 4 Lobrathium schuelkei sp. n., det. V. Assing 2011”

(cAss).

Etymology. The species is dedicated to my friend and col-

league Michael Schtilke, whose most remarkable materi-

al from China, which included also the holotype of this

species, represented a major, invaluable contribution to the

present paper.

Description. Body length 7.8 mm; length of forebody 4.2

mm. In external characters practically identical to L. tau-

reum (Figs 194-195), distinguished only by the male sex-

ual characters:

3: sternites V-VI with shallow median impression pos-

teriorly; sternite VII strongly transverse and with distinct

median impression with pubescence, posterior margin

broadly concave (Fig. 196); sternite VIII weakly trans-

verse, with pronounced long median impression, this im-

pression with numerous modified, short and stout black

setae, posterior excision small and almost V-shaped (Fig.

197); aedeagus 1.1 mm long, ventral process symmetric

and apically truncate in ventral view (Figs 198-199).

Comparative notes. Among Chinese Lobrathium, L.

schuelkei is most similar to L. taureum, from which it is

reliably distinguished only by the less deep posterior ex-

cision of the male sternite VIII and by the completely dif-

ferent morphology of the aedeagus.

Distribution and natural history. The type locality is sit-

uated in the Qinling Shan in the south of Shaanxi province,

China (Fig. 180). The somewhat teneral holotype was col-

lected on a gravel bank at an altitude of 1450 m, togeth-

er with L. hongkongense and L. configens.

Lobrathium radens sp. n. (Figs 153, 200-205)

Type material. Holotype @: “China: Guizhou, Leishan

Co., SE Kaili, NE Leishan, Leigong Shan, E-slope,

1700-1800 m, 14.—24.6.2001 / env. of pass between Leis-

han and Fangxiang (26°22.74’°N 108°12.99’E), leg.

Schillhammer (7) / Holotypus ¢ Lobrathium radens sp.

n., det. V. Assing 2011” (NHMW). Paratypes: 232: sa-

me data as holotype (NHMW, cAss).

Etymology. The specific epithet (Latin, present participle

of radere: to scrape) alludes to the presence of distinct

spines and carinae on the ventral process of the aedeagus.

Description. Large species, body length 8.0—9.2 mm;

length of forebody 4.9-5.2 mm. Habitus as in Fig. 200.

Coloration: body black, elytra with large yellowish spot

OZFMK

102

195 198

200

201

204

Volker Assing

205

Figs 194-205. Lobrathium schuelkei (194-199) and L. radens (200-205). 194, 200: habitus; 195, 201: forebody; 196, 202: male

sternite VII; 197, 203: male sternite VII; 198-199, 204—205: aedeagus in lateral and in ventral view. Scale bars: 194-195, 200-201:

1.0 mm; 196-199, 202—205: 0.5 mm.

posteriorly not reaching lateral and posterior margins; legs

reddish; antennae dark-reddish to reddish-brown.

Head (Fig. 201) weakly transverse, behind eyes strong-

ly convex; posterior angles weakly marked; punctation of

dorsal surface moderately coarse and very dense, with in-

terstices reduced to narrow ridges, small spot in median

dorsal portion and frons impunctate and glossy. Eyes mod-

erately large, approximately half as long as distance from

posterior margin of eye to neck. Antenna 2.83.0 mm long.

Pronotum (Fig. 201) relatively short and broad, approx-

imately 1.15—1.20 times as long as broad and almost as

broad as head, lateral margins almost straight in the mid-

dle and converging posteriad in dorsal view; punctation

sparser and slightly coarser than that of head.

Bonn zoological Bulletin 61 (1): 49-128

Elytra (Fig. 201) approximately 0.95 times as long as,

and distinctly broader than pronotum; humeral angles

marked; punctation coarse, dense, and not arranged in se-

ries; interstices without microsculpture and glossy. Hind

wings fully developed.

Abdomen distinctly narrower than elytra; punctation

fine and dense; posterior margin of tergite VII with pal-

isade fringe; posterior margin of tergite VIII weakly con-

vex.

&: sternites II-VI unmodified; sternite VII with small

and V-shaped median impression with cluster of approx-

imately 10 modified, short and stout black setae on either

side, posterior margin distinctly and broadly concave (Fig.

202); sternite VHI weakly transverse, with pronounced

median impression, this impression with numerous mod-

©ZFMK

Revision of East Palaearctic Lobrathium 103

ified, short and stout black setae, posterior excision deep,

broad, and somewhat U-shaped, on either side of this ex-

cision with dense black setae (Fig. 203); aedeagus approx-

imately 1.5 mm long, ventral process blade-shaped and

apically acute, on ventral surface with two tooth-like

processes and additional tranverse carinae (Figs 204—205).

Comparative notes. Lobrathium radens is characterised

particularly by its large size, broad body, the dense punc-

tation of the head, the short and broad pronotum, and par-

ticularly by the shape and chaetotaxy of the male stern-

ite VII and VIII, as well as by the morphology of the

aedeagus.

Distribution and natural history. The type locality is sit-

uated in the Leigong Shan in eastern Guizhou province,

China (Fig. 153). The type specimens were collected at

an altitude 1700-1800 m.

Lobrathium retrocarinatum sp. n. (Figs 153, 206-211)

Type material. Holotype ¢ [teneral]: “China: Yunnan pro-

vince, Shanzhi env., 22.—24.VI.2007, Jizu Shan Mt.,

2180-2580 m, along the path to the summit, 27°57.7—-8’N,

100°22.1—23.6’E, J. Hajek & J. Ruzicka leg. / sifted de-

tritus and leaves, dense mixed forest (with dominant Pi-

nus, Quercus and Rhododendron) near stream [CH45—47]

/ Holotypus 4 Lobrathium retrocarinatum sp. n., det. V.

Assing 2011” (cAss).

Etymology. The specific epithet (Latin, adjective) is com-

posed of retro (back, backwards) and carinatum (carinate);

it refers to the long dorsal carina of the ventral process of

the aedeagus.

Description. Species of moderate size, body length 7 mm;

length of forebody 3.7 mm. Habitus as in Fig. 206. Col-

oration: whole body blackish, elytra with weak purple hue

and with relatively small, circular yellowish spot poste-

riorly not reaching lateral and posterior margins, nor su-

ture.

Head (Fig. 207) weakly transverse, 1.07 times as wide

as long, of subrectangular shape, widest across eyes; pos-

terior angles moderately marked; punctation of dorsal sur-

face coarse and dense; vertex and frons less densely punc-

tate; interstices without microsculpture. Eyes large and

bulging, approximately 0.8 times as long as distance from

posterior margin of eye to neck in dorsal view. Antenna

2.1 mm long.

Pronotum (Fig. 207) moderately oblong, approximate-

ly 1.2 times as long as broad and about 0.95 times as broad

as head, lateral margins almost straight in the middle and

weakly converging posteriad in dorsal view; punctation

similar to that of head.

Bonn zoological Bulletin 61 (1): 49-128

Elytra (Fig. 207) long, 1.05 times as long as, and dis-

tinctly broader than pronotum; humeral angles marked;

punctation coarse, dense, and not arranged in distinct se-

ries; interstices without microsculpture and glossy. Hind

wings fully developed.

Abdomen slightly narrower than elytra; punctation mod-

erately fine and dense; interstices with distinct transverse

microsculpture; posterior margin of tergite VII with pal-

isade fringe; posterior margin of tergite VII weakly con-

vex.

S: sternites V and VI with shallow median impression

posteriorly; sternite VII strongly transverse and with pro-

nounced, extensive median impression, this impression

with a cluster of distinctly modified, short and stout black

setae on either side of the impunctate middle, posterior

margin broadly and distinctly concave (Fig. 208); stern-

ite VUI approximately as long as broad and with pro-

nounced, deep and oblong median impression, this impres-

sion with numerous distinctly modified, short and stout

black setae, except for the impunctate posterior median

portion, posterior excision relatively deep and almost U-

shaped, on either side of this excision with dense black

setae (Fig. 209); aedeagus 1 mm long, ventral process

basally broad, gradually tapering towards apex, apically

acute, without distinct spines, but with pronounced, long

median carina dorsally (Figs 210-211).

Comparative notes. Lobrathium retrocarinatum is dis-

tinguished from other spotted Lobrathium species partic-

ularly by the pronounced modifications of the male ster-

nites VII and VIII, as well as by the distinctive morphol-

ogy of the ventral process of the aedeagus. From many

Chinese representatives of the genus with spotted elytra,

it is additionally distinguished by the relatively small and

subcircular elytral spots.

Distribution and natural history. The type locality is sit-

uated in the Jizu Shan in northern Yunnan, China (Fig.

153). The latitude indicated on the label of the holotype

is evidently erroneous and should read 25° rather than 27°.

The somewhat teneral type specimen was collected in a

mixed forest near a stream at an altitude of 2180-2580 m.

Lobrathium feldmanni sp. n. (Figs 193, 212-217)

Type material. Holotype ¢: “China-N.Sechuan, Wang-

cang, 18.-20.5.2002, Lgt E Kuéera / Holotypus <4 Lo-

brathium feldmanni sp. n., det. V. Assing 2011” (cAss).

Etymology. The species is dedicated to my friend and col-

league Benedikt Feldmann, to whom I owe the holotype.

©OZFMK

104 Volker Assing

213

212

216

217 215

Figs 206-217. Lobrathium retrocarinatum (206-211) and L. feldmanni (212-217). 206, 212: habitus; 207, 213: forebody; 208,

214: male sternite VH; 209, 215: male sternite VIH; 210-211, 216-217: aedeagus in lateral and in ventral view. Scale bars: 206-207,

212-213: 1.0 mm; 208-211, 214-217: 0.5 mm.

Description. Large species, body length 8.6 mm; length

of forebody 4.8 mm. Habitus as in Fig. 212. Coloration:

body dark-brown; legs brown; antennae dark-brown.

Head (Fig. 213) approximately as long as wide, behind

eyes convexly rounded towards neck; posterior angles not

marked; punctation of dorsal surface relatively fine and

dense; interstices narrow, narrower than diameter of punc-

tures, and glossy, without microsculpture; transverse patch

in median dorsal portion and frons impunctate. Eyes rel-

atively small, approximately one third as long as distance

from posterior margin of eye to neck. Antenna slender, 3.2

mm long.

Bonn zoological Bulletin 61 (1): 49-128

Pronotum (Fig. 213) 1.27 times as long as broad and

0.93 times as broad as head, widest in anterior half, con-

vexly tapering posteriorly; punctation dense, slightly

coarser than that of head.

Elytra (Fig. 213) 0.92 times as long as, and distinctly

broader than pronotum; humeral angles marked; puncta-

tion coarse, dense, and not arranged in series; interstices

without microsculpture and glossy. Hind wings apparent-

ly fully developed.

Abdomen as broad as elytra; punctation fine and dense;

posterior margin of tergite VII with palisade fringe; pos-

terior margin of tergite VIII distinctly convex.

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Revision of East Palaearctic Lobrathium 105

219 222

227

228

220

221

226

Figs 218-228. Lobrathium ablectum (218-223) and L. excisissimum (224-228). 218, 224: forebody; 219: antenna; 220, 225: male

sternite VII; 221, 226: male sternite VIII; 222-223, 227-228: aedeagus in lateral and in ventral view. Scale bars: 218-219, 224:

1.0 mm; 220-223, 225—228: 0.5 mm.

¢: sternite VII with shallow median impression poste-

riorly, without modified setae (Fig. 214); sternite VII

moderately transverse and with distinct median impres-

sion posteriorly, this impression with numerous modified,

short and stout black setae, posterior excision shallow, sub-

rectangular, and with somewhat bisinuate anterior margin

(Fig. 215); aedeagus 1.8 mm long, ventral process long,

slender, and bisinuate in lateral view (Figs 216-217).

Comparative notes. Lobrathium feldmanni is distin-

guished from its congeners particularly by the shape and

Bonn zoological Bulletin 61 (1): 49-128

chaetotaxy of the male sternite VIII, as well as by the mor-

phology of the aedeagus. In addition, it is characterised

by rather large body size, slender antennae, and the uni-

formly dark-brown coloration of the body.

Distribution and natural history. The type locality,

Wangcang Xian, is situated in the northeast of Sichuan

province, China (Fig. 193). Additional data are not avail-

able.

©ZFMK

106 Volker Assing

Lobrathium ablectum sp. n. (Figs 140, 218-223)

Type material. Holotype ¢: “China: W-Hubei (Daba

Shan), creek valley 8 km NW Muyuping, 31°29°N,

110°22’E, 1550-1650 m, 18.VII.2001, leg. M. Schiilke

[CHO1—16A] / creek valley, deciduous forest, moss (sift-

ed) [CH01-16A]/ Holotypus 4 Lobrathium ablectum sp.

n., det. V. Assing 2011” (cAss). Paratypes: 29 2: same da-

ta as holotype (cSch).

Etymology. The specific epithet (Latin, past participle of

the verb ablegere: to send or move away) alludes to the

characteristic position of the elytral spot.

Description. Relatively large species, body length 7.7—9.3

mm; length of forebody 4.44.6 mm. Coloration: whole

body black, except for the dark-brown tarsi and apical an-

tennomeres; elytra with a subcircular reddish spot situat-

ed in anterior portion of posterior half (1.e., at some dis-

tance from posterior margin).

Head (Fig. 218) approximately as wide as long or weak-

ly transverse, widest across eyes; lateral margins behind

eyes convexly rounded towards neck; posterior angles ob-

solete; dorsal surface between eyes uneven, with shallow

median and lateral impressions; punctation of dorsal sur-

face moderately coarse, but extremely dense, interstices

reduced to very narrow ridges; frons and a transverse,

somewhat elevated patch on vertex glossy and with spars-

er macropunctures, remainder of dorsal surface matt; in-

terstices without microsculpture. Eyes rather large and

strongly convex, distinctly more than half as long as dis-

tance from posterior margin of eye to neck. Antenna (Fig.

219) slender, approximately 2.5 mm long, antennomeres

VI-IX at least 1.5 times as long as broad.

Pronotum (Fig. 218) long and slender, 1.3 times as long

as broad and approximately 0.9 times as broad as head;

lateral margins in the middle almost parallel, only indis-

tinctly converging posteriad; punctation coarse and con-

spicuously dense, except for a glossy, impunctate, oblong

and somewhat bulging lateral patch on either side and for

the impunctate glossy midline.

Elytra (Fig. 218) long, approximately 1.1 times as long

as, and distinctly broader than pronotum; humeral angles

marked; punctation very coarse, dense, not arranged in se-

ries, and somewhat rugosely confluent near anterior por-

tion of suture; interstices without microsculpture and

glossy. Hind wings apparently fully developed.

Abdomen narrower than elytra; punctation fine and

dense; interstices at most with traces of microsulpture and

glossy; posterior margin of tergite VII with palisade fringe.

S: posterior margin of tergite VIII truncate; sternites

TV—V indistinctly depressed in the middle; sternite VI with

shallow median impression posteriorly; sternite VI mod-

erately transverse, with median impression without pubes-

Bonn zoological Bulletin 61 (1): 49-128

cence, posterior margin distinctly concave and in the mid-

dle weakly convex (Fig. 220); sternite VII weakly trans-

verse, in the middle with long and pronounced impression,

this impression with a cluster of numerous distinctly mod-

ified, short and stout black setae on either side, posterior

excision moderately deep and almost U-shaped (Fig. 221);

aedeagus 1.35 mm long, ventral process strongly sclero-

tised and apically bifid (Figs 222—223).

®: posterior margin of tergite VIII weakly convex.

Comparative notes. Lobrathium ablectum is readily dis-

tinguished from all its congeners not only by the male sex-

ual characters, but also by several distinctive external char-

acters, particularly the punctation and impressions of the

head, the punctation and impunctate lateral patches of the

pronotum, and the position of the elytral spot.

Distribution and natural history. The type locality is sit-

uated in the Daba Shan in the west of Hubei province, Chi-

na (Fig. 140). The type specimens were sifted from moss

in a deciduous forest, together with L. taureum and L.

spathulatum, at an altitude of 1550-1650 m.

Lobrathium excisissimum sp. n. (Figs 180, 224-228)

Type material. Holotype ¢: “China (N-Yunnan) Dali Bai

Nat. Aut. Pref., Diancang Shan, 3 km W Dali old town,

creek valley at “Cloud Road”, right upper chairlift station,

25°41.1’N, 100°06.8’E, 2750 m (under stones, leaf litter

along creek), 17.VI.2005 D.W. Wrase [11C] / Holotypus

3 Lobrathium excisissimum sp. n., det. V. Assing 2011”

(cAss). Paratype 2: same data as holotype (cSch).

Etymology. The specific epithet (Latin, superlative of ex-

cisus) alludes to the pronounced posterior excision of the

male sternite VII.

Description. Large species, body length 9.0-9.5 mm;

length of forebody 4.9-5.1 mm. Coloration: body black,

elytra with subcircular yellowish spot reaching neither

posterior nor lateral margins; legs blackish-brown with

reddish tarsi; antennae blackish-brown.

Head (Fig. 224) approximately 1.1 times as wide as

long; lateral margins behind eyes convexly rounded to-

wards neck; posterior angles not marked; punctation of

dorsal surface relatively fine and dense; interstices nar-

row, narrower than diameter of punctures, and glossy,

without microsculpture; median dorsal portion and frons

impunctate or sparsely punctate. Eyes moderately large

and weakly convex, slightly more than half as long as dis-

tance from posterior margin of eye to neck in dorsal view.

Antenna moderately slender, approximately 3.2 mm long.

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Revision of East Palaearctic Lobrathium 107

Pronotum (Fig. 224) short and broad, 1.10—1.15 times

as long as broad and approximately 0.95 times as broad

as head, widest anteriorly; punctation similar to that of

head, but somewhat less dense; interstices glossy.

Elytra (Fig. 224) approximately 0.9 times as long as, and

distinctly broader than pronotum; humeral angles marked;

punctation coarse, dense, and not arranged in series; in-

terstices without microsculpture and glossy. Hind wings

apparently fully developed.

Abdomen approximately as broad as elytra; punctation

fine and dense; posterior margin of tergite VU with pal-

isade fringe; posterior margin of tergite VIII distinctly con-

vex.

¢: sternite VI with shallow median impression poste-

riorly; sternite VII with extensive median impression, this

impression with numerous distinctly modified, short and

stout black setae, posterior margin strongly and broadly

convex (Fig. 225); sternite VII moderately transverse,

with distinct median impression and remarkably deep, U-

shaped posterior excision, the margins of this impression

and the median excision with numerous distinctly modi-

fied, short and stout black setae (Fig. 226); aedeagus 1.6

mm long, ventral process straight in lateral view, with two

pairs of pronounced ventral teeth (Figs 227-228).

Comparative notes. Lobrathium excisissimum is readi-

ly distinguished from all its congeners by the combina-

tion of large size, broad body (short and compact prono-

tum), the relatively fine and dense punctation of the head,

the coloration, and particularly by the distinctive male pri-

mary and secondary sexual characters. The similarly de-

rived shapes and chaetotaxy of the male sternite VII and

VIII, as well as the similar general morphology of the

aedeagus suggest that L. excisissimum is closely allied to

L. radens.

Distribution and natural history. The type locality is sit-

uated in the Diancang Shan in northern Yunnan, China

(Fig. 180). The type specimens were collected on a stream

bank at an altitude of 2750 m.

Lobrathium rotundiceps (Koch, 1939) (Figs 141,

229-230)

Lathrobium rotundiceps Koch, 1939: 163 f.

Type material examined. Holotype 9 [dissected prior to

present study]: “Tienmuschan, N.W. [sic] China Rtt. / Type

/ Lathrobium rotundiceps Koch, det. C. Koch / Holoty-

pus 1956, det. Kamp / Holotypus Lathrobium rotundiceps

Koch / Domene rotundiceps (Koch) 9, V.I. Gusarov det.

1993 / Lobrathium rotundiceps Koch, det. V. Assing 2012”

(NHMB).

Bonn zoological Bulletin 61 (1): 49-128

Comment. The original descriptions is based on “ein

Weibchen aus Tienmuschan (Nordwestlichen China)”

(Koch 1939). The specimen is deposited in the collections

of the NHMB.

Redescription. Large species, body length 10.2 mm;

length of forebody 6.0 mm. Habitus as in Fig. 229. Col-

oration: body dark reddish-brown; legs dark-yellowish; an-

tennae brown.

Head (Fig. 230) approximately 1.1 times as long as

wide; lateral margins behind eyes convexly rounded to-

wards neck; posterior angles completely obsolete; punc-

tation of dorsal surface relatively fine and conspicously

dense, not sparser in median dorsal portion; interstices re-

duced to narrow ridges everywhere; dorsal surface matt.

Eyes relatively small and weakly convex, barely one third

as long as distance from posterior margin of eye to neck

in dorsal view. Antenna slender, approximately 4 mm long.

Pronotum (Fig. 230) slender, approximately 1.25 times

as long as broad and approximately 0.95 times as broad

as head, widest anteriorly, weakly tapering posteriad;

punctation much coarser and less dense than that of head;

interstices glossy, on average slightly narrower than di-

ameter of punctures.

Elytra (Fig. 230) 0.92 times as long as, and distinctly

broader than pronotum; humeral angles marked; puncta-

tion somewhat coarser and denser than that of pronotum,

not arranged in distinct series, but partly somewhat

obliquely confluent. Hind wings present and probably ful-

ly developed.

Abdomen approximately as broad as elytra; punctation

fine and dense; posterior margin of tergite VII with pal-

isade fringe; posterior margin of tergite VIII obtusely

pointed in the middle.

S: unknown.

Comparative notes. Lobrathium rotundiceps 1s charac-

terised especially by its large body size, the ovoid shape

of the distinctly oblong head (similar to the condition in

many Domene species), as well as by the relatively fine

and very dense punctation of the head.

Distribution and natural history. The type locality Zhe-

jiang is situated in Zhejiang province, northeastern (not

northwestern, as indicated on the label!) China (Fig. 141).

Additional data and records are unknown.

Lobrathium cholaicum sp. n. (Figs 141, 231—236)

Type material. Holotype ¢: “SE Tibet, “Chola Shan”

pass, road Yanjing — Markam, 50 km S Markam, 4400 m,

29°16’N, 98°38’E, mixed forest, 24.-27.V1.1997, Jaroslav

Turna leg. / Holotypus 4 Lobrathium cholaicum sp. n.,

det. V. Assing 2011” (cAss). Paratypes: 543’, 19: same

data as holotype (cSch, cAss).

OZFMK

108 Volker Assing

237 240 241

~ 233 238

Figs 229-241. Lobrathium rotundiceps, holotype (229-230), L. cholaicum (231-236), and L. daxuense (237-241). 229, 231: habi-

tus; 230, 232, 237: forebody; 233, 238: male sternite VII; 234, 239: male sternite VIII; 235-236, 240-241: aedeagus in lateral and

in ventral view. Scale bars: 229-232, 237: 1.0 mm; 233-236, 238-241: 0.5 mm.

Revision of East Palaearctic Lobrathium 109

Etymology. The specific epithet (adjective) is derived

from the name of the pass where the types were collect-

ed.

Description. Species of moderate size, body length 6—7

mm; length of forebody 3.4—3.6 mm. Habitus as in Fig.

231. Coloration: body blackish-brown, with the elytra usu-

ally slightly paler; legs reddish to reddish-brown; anten-

nae reddish.

Head (Fig. 232) approximately as long as broad or weak-

ly oblong; lateral margins behind eyes convexly rounded

towards neck; posterior angles not marked; punctation of

dorsal surface coarse and moderately dense, somewhat

sparser on vertex and on frons; interstices glossy, with-

out microsculpture. Eyes moderately large and moderate-

ly convex, clearly less than half as long as distance from

posterior margin of eye to neck in dorsal view. Antenna

moderately slender, approximately 2 mm long.

Pronotum (Fig. 232) moderately oblong, 1.20—1.25

times as long as broad and approximately 0.95 times as

broad as head, widest anteriorly and noticeably tapering

posteriad; punctation similar to that of head, or slightly

coarser; interstices glossy.

Elytra (Fig. 232) relatively short, approximately 0.85

times as long as pronotum, distinctly widened posteriad;

humeral angles weakly marked; punctation approximate-

ly as coarse as that of pronotum, but distinctly denser; in-

terstices without microsculpture and glossy. Hind wings

reduced.

Abdomen widest at segments VI/VII, distinctly broad-

er than elytra; punctation not particularly fine and defined,

dense on anterior tergites, somewhat sparser on tergite VI;

interstices with very shallow, almost obsolete microsculp-

ture and glossy; posterior margin of tergite VII without

palisade fringe; posterior margin of tergite VIII truncate

to very weakly convex.

S: sternite VI with shallow median impression poste-

riorly; sternite VII moderately transverse, with relatively

shallow median impression posteriorly, without modified

setae, posterior margin broadly and weakly concave (Fig.

233); sternite VII moderately concave, with distinct me-

dian impression, this impression with relatively sparse,

distinctly modified, short and stout black setae, posterior

excision rather shallow and concave, on either side of this

excision with tuft of long black setae (Fig. 234); aedea-

gus approximately 0.9 mm long, ventral process short, al-

most straight in lateral view, and apically acute in ventral

view (Figs 235-236).

Comparative notes. Lobrathium cholaicum is readily dis-

tinguished from all its congeners by the male sexual char-

acters (shape and chaetotaxy of the male sternite VIII,

morphology of the aedeagus), and from other Chinese Lo-

brathium species additionally by the absence of a palisade

fringe at the posterior margin of the abdominal tergite VII.

Bonn zoological Bulletin 61 (1): 49-128

Together with the five following species, L. cholaicum

belongs to a group of species distributed from Nepal to

western China and including also the Himalayan L.

wittmeri and L. kosiense. For characters constituting this

species group see the comparative notes in the section on

L. wittmeri.

Distribution and natural history. The type locality is sit-

uated in the extreme southeast of Tibet, close to the bor-

der with Sichuan (Fig. 141). The type specimens were col-

lected in a mixed forest at an altitude of 4400 m.

Lobrathium daxuense sp. n. (Figs 180, 237-241)

Type material. Holotype @: “China (W Sichuan) (4) Dax-

ue Shan, river valley 5 km E Kangding [sic], 2500-2800

m, 30.03N, 102.00E, 20./23.V.1997 Wrase / Holotypus 4

Lobrathium daxuense sp. n., det. V. Assing 2011” (cAss).

Paratypes: 23.4, 19: same data as holotype (cSch, cAss);

24: “China, Sichuan, Gongga Shan, aby. Camp 3,

3300-3350 m, 23.VII.1994, A. Smetana [C19] (cSme,

cAss); 14: “China: Sichuan Gongga Shan, Hailuogou, in

front of Glacier 1, 2850 m, 29°35N, 102°00E, 7.VII.1998,

A. Smetana [C76]” (cAss).

Etymology. The specific epithet (adjective) is derived

from the name of the mountain range where the types were

collected.

Description. Habitus as in Fig. 266. Externally highly

similar to L. cholaicum, but distinguished as follows:

Head (Fig. 237) with less coarse and distinctly denser

punctation; vertex at most with very small patch with

Sparse punctation. Eyes slightly larger, but still less than

half as long as distance from posterior margin of eye to

neck in dorsal view. Pronotum (Fig. 237) with less coarse

and somewhat denser punctation. Elytra (Fig. 237) with

slightly less coarse and slightly denser punctation. Ab-

domen with much finer and much denser punctation; pos-

terior margin of tergite VII with narrow rudiment of a pal-

isade fringe.

4: sexual characters similar to those of L. cholaicum;

sternite VII as in Fig. 238; sternite VIII in median impres-

sion with denser setae, posterior excision broader and

deeper (Fig. 239); aedeagus with longer and apically less

acute ventral process in different position (base closer to

ventral opening) (Figs 240-241).

Comparative notes. As can be inferred from the similar

external characters, the similar modifications of the male

sternites VII and VIII, as well as from the similar mor-

phology of the aedeagus, L. daxuense is closely related

to L. cholaicum; for characters distinguishing these two

species see the description above. From other Chinese con-

©ZFMK

110 Volker Assing

242 243

247

249

244

250 251 252

Figs 242-252. Lobrathium bimembre. 242: habitus (holotype); 243-244: forebody of micropterous (243) and of macropterous spec-

imen (244); 245: male sternite VH; 246: male sternite VII; 247-252: aedeagus in lateral and in ventral view of males from Yan-

men (247-248), Gaoligong Shan (249-250), and Diancang Shan (251-252). Scale bars: 242—244: 1.0 mm; 245-252: 0.5 mm.

geners, except L. domenoides, L. daxuense is separated ad-

ditionally by the combination of moderate size and the

short and posteriorly widened elytra. For comments on

species group affiliations see the comparative notes in the

sections on L. wittmeri and L. cholaicum.

Distribution and natural history. The species is current-

ly known from two localities in the Daxue Shan in Sichuan

Bonn zoological Bulletin 61 (1): 49-128

province, China (Fig. 180). The specimens from the type

locality were collected in a river valley at an altitude of

2500-2800 m, those from the Gongga Shan at elevations

of 2850-3350 m. In the Gongga Shan, L. daxuense was

collected at the edge of an original coniferous forest (main-

ly fir) by sifting various leaf litter and in sandy areas near

a river leaving a glacier (Smetana pers. comm.) together

with L. configens, close to where L. duplex was found.

©ZFMK

Revision of East Palaearctic Lobrathium 111

Lobrathium bimembre sp. n. (Figs 141, 242-252)

Type material. Holotype ¢@ [micropterous]: “China

(Yunnan), Dali Bai Nat. Aut. Pref., Diancang Shan, 4 km

W Dali old town, 2900-3000 m, 25°41.4’N, 100°06.7’E,

E slope, former stone pit (in overgrown gravel/soil/plant

roots), 31.VIHI.2003 Wrase [20] / Holotypus ¢ Lobrathi-

um bimembre sp. n., det. V. Assing 2011” (cAss).

Paratypes: 12 [micropterous]: same data as holotype

(cSch); 14, 19 [micropterous]: “China (Yunnan), Dali Bai

Nat. Aut. Pref., Diancang Shan, 4 km W Dali old town,

2900-3000 m / 25°41.4°N, 100°06.7’E, E slope, former

stone pit (in overgrown gravel/soil/plant roots/und.

stones), 18.VI.2005 D.W. Wrase [12]” (cSch); 1 9 [mi-

cropterous]: “China (Yunnan), Dali Bai Auton. Pref., Di-

ancang Shan W Dali, 25°41°33”N, 100°06736”E, 2927

m (former stone pit, in gravel /under stones), 28.V.2007

D.W. Wrase [05]” (cSch); 14, 229 [1¢ macropterous,

22° micropterous]: “China (Yunnan) Nujiang Lisu Aut.

Pref., Gaoligong Shan, pass 21 km NW Liuku, 3150 m,

25°58’22”N, 98°41°00’E (bamboo, shrubs, under stones

along road), 9.VI.2007 D.W. Wrase [26]” (cSch, cAss);

12 [micropterous]: “China (Yunnan) Nujiang Lisu Aut.

Pref., Gaoligong Shan, valley 21 km W Gongshan, 3320

m, 27°47°03”N, 98°27°39”E (moss, alder, bamboo,

Rhodod., sifted), 6. VI.2007 D.W. Wrase [22]” (cSch); 19

[micropterous]: “China (Yunnan), Dali Bai Auton. Pref.,

Diancang Shan W Dali, 25°41°49”N, 100°06’24”’E, 2970

m (under stones), 28.V.2007 D.W. Wrase [03]” (cSch); 12

[micropterous]: “China- Yunnan, 13.6.—23.6.2005, Igt. E.

Kucera” (cSch).

Etymology. The specific epithet (Latin, adjective: two-

shaped) alludes to the conspicuous wing-dimorphism.

Description. Wing-dimorphic species of moderately

large size; body length 6.5—7.5 mm; length of forebody

3.7-4.1 mm (micropterous morph) and 4.4 mm

(macropterous morph), respectively. Habitus of mi-

cropterous specimen as in Fig. 242. Externally highly sim-

ilar to the slightly smaller L. daxuense, but distinguished

as follows:

Head (Figs 243-244) with very dense punctation, also

on vertex, almost completely matt. Antenna 2.3—2.5 mm

long. Pronotum (Figs 243—244) with denser and slightly

coarser punctation. Elytra 0.85 times (micropterous

morph, Fig. 243) or 1.13 times as long as pronotum

(macropterous morph, Fig. 244). Posterior margin of ab-

dominal tergite VII with palisade fringe in both morphs.

&: sternites II-VI unmodified; sternite VII moderate-

ly transverse, with shallow median impression posterior-

ly, this impression with numerous short black setae direct-

ed postero-mediad, posterior margin only indistinctly con-

cave (Fig. 245); sternite VIII moderately tranverse, with

rather long and relatively narrow median impression, this

Bonn zoological Bulletin 61 (1): 49-128

impression with numerous distinctly modified, short and

stout black setae, posterior exision small, on either side

with tuft of black setae (Fig. 246); aedeagus approximate-

ly | mm long, ventral process relatively longer than in L.

daxuense, apically more acute (lateral view), and narrow-

er in ventral view (Figs 247-252).

Comparative notes. Lobrathium bimembre is distin-

guished from all its congeners particularly by the male sex-

ual characters. It is additionally separated from the syn-

topic, externally similar, and similarly wing-dimorphice L.

duplex by slightly smaller size, much denser punctation

of the head, shorter antennae, and by the less distinctly

convex posterior margin of the abdominal tergite VIII. For

comments on species group affiliations see the compara-

tive notes in the sections on L. wittmeri and L. cholaicum.

Distribution and natural history. This species is known

from several localities in the Diancang Shan, the

Gaoligong Shan, and one in the environs of Yanmen, all

of them in Yunnan province (Fig. 141). The specimens

were collected from under stones, from gravel, and by sift-

ing litter and debris at altitudes of 2900-3320 m. Only one

male in a total of eleven specimens is macropterous.

Lobrathium duplex sp. n. (Figs 193, 253-259)

Type material. Holotype ¢ [micropterous]: “China

(Yunnan), Dali Bai Auton. Pref., Diancang Shan W Dali,

25°41’49”N, 100°06’24”E, 2970 m (under stones),

28.V.2007 D.W. Wrase [03] / Holotypus 3 Lobrathium du-

plex sp. n., det. V. Assing 2011” (cAss). Paratypes: 14,

329 [micropterous]: same data as holotype (cSch,

cAss); 14, 39 [micropterous]: “China (Yunnan), Dali

Bai Auton. Pref., Diancang Shan W Dali, 25°41°33”N,

100°06’°36”E, 2927 m (former stone pit, in gravel /under

stones), 28.V.2007 D.W. Wrase [05]” (cSch); 1¢ [mi-

cropterous]: “China: N-Yunnan [C03—19], Dali Bai Nat.

Aut. Pref., Diancang Shan, 3 km W Dali old town, creek

valley and pine forest at “Cloud Road”, right upper chair-

lift station, 25°41.1’N, 100°06.8’E, 2650-2750 m /

[C03—19] litter, pine needles, moss (dry and wet), mush-

rooms, 29. VIII.2003, leg. M. Schiilke” (cSch); 53, 12

[micropterous; 4 exs. teneral]: “China (Yunnan), Dali Bai

Nat. Aut. Pref., Diancang Shan, 4 km W Dali old town,

2900-3000 m, 25°41.4’N, 100°06.7’E, E slope, former

stone pit (in overgrown gravel/soil/plant roots), 31.VI-

11.2003 Wrase [20]” (cSch); 39 9 [micropterous]: “Chi-

na (Yunnan), Dali Bai Nat. Aut. Pref., Diancang Shan, 4

km W Dali old town, 2900-3000 m / 25°41.4°N,

100°06.7’E, E slope, former stone pit (in overgrown grav-

el/soil/plant roots/und. stones), 18.V1I.2005 D.W. Wrase

[12]” (cSch, cAss); 93'S, 1192 [14, 12 macropterous]:

“China (W Sichuan) (4) Daxue Shan, river valley 5 km E

OZFMK

112 Volker Assing

253 255 258 259

263

264 265

257

262

Figs 253-265. Lobrathium duplex (253-259) and L. domenoides (260-265). 253, 260: habitus (253: micropterous morph); 254-255,

261: forebody (254: micropterous morph; 255: macropterous morph); 256, 262: male sternite VII; 257, 263: male sternite VIII;

258-259, 264-265: aedeagus in lateral and in ventral view. Scale bars: 253-255, 260-261: 1.0 mm; 256-259, 262-265: 0.5 mm.

Revision of East Palaearctic Lobrathium 113

Kanding [sic], 2500-2800 m, 30.03N, 102.00E,

20./23.V.1997 Wrase” (cSch, cAss); 1043, 19: “China:

W-Sichuan (4a), Daxue Shan, Bachtal 5 km E Kangding,

30.03.28N, 102.00.15E, 2500-2800 m, 23.V.1997, M.

Schiilke” (cSch, cAss); 244, 39 9 [micropterous]: “Chi-

na W.Sichuan (Ganzi Tibet. Aut. Pref., Kangding Co.)

Daxue Shan, brook valley 5 km E Kangding, 2500-2600

m, 30°03N, 102°00E, 24.VI.1999 D.W. Wrase” (cSch,

cAss); 19 [micropterous]: same data, but leg. M. Schiilke

(cSch); 12: “China, S Sichuan 27. VII.1997, Daliang Shan

mts., road Meigu-Leibo, pass 15 km NE Meigu, 28°25°N,

103°17°E, Jaroslav Turna leg.” (cSch); 399: “China

Sichuan, Gongga Shan, Hailuogou, head of Glacier 1,

2850 m, 9.VII.96, 29°35N, 102°00E, C57 / collected by

A. Smetana, J. Farka¢ and P. Kabatek” (cSme, cAss).

Etymology. The specific epithet (Latin, adjective: two-

fold) refers to the conspicuous wing-dimorphism of this

species.

Description. Relatively large, wing-dimorphic species,

body length 7.3—9.5 mm; length of forebody 4.1—4.8 mm

(micropterous morph) and 5.0 mm (macropterous morph).

Habitus of holotype as in Fig. 253. Coloration: body black-

ish; legs with femora dark-brown to blackish, tibiae red-

dish-brown to blackish-brown, and tarsi reddish to dark-

reddish; antennae usually reddish to reddish-brown with

more or less distinctly infuscate antennomere I.

Head (Figs 254-255) usually 1.05—1.10 times as long

as broad; lateral margins behind eyes smoothly curving

towards neck; posterior angles usually obsolete, rarely

weakly marked; punctation of dorsal surface moderately

coarse and dense, slightly less dense on frons and on ver-

tex; interstices without microsculpture, distinctly narrow-

er than diameter of punctures (except on frons and on

small patch of vertex). Eyes slightly larger and more con-

vex in macropterous than in micropterous morph, approx-

imately half as long as distance from posterior margin of

eye to neck, or slightly longer in dorsal view. Antenna long

and slender, 3.0—3.3 mm long; antennomeres IV—VIII at

least twice as long as broad.

Pronotum (Figs 254-255) strongly oblong, usually

1.30—1.35 times as long as broad and approximately

0.90—0.95 times as broad as head, distinctly tapering pos-

teriad; punctation distinctly coarser and somewhat less

dense than that of head; interstices without microsculp-

ture and glossy.

Elytra dimorphic; micropterous morph (Fig. 254): mod-

erately short and slender, approximately 0.9 times as long

as pronotum, weakly widened posteriad, and weakly con-

vex in cross-section; humeral angles practically obsolete;

macropterous morph (Fig. 255): long and broad, approx-

imately 1.1 times as long as pronotum, with parallel lat-

eral margins, and distinctly convex in cross-section; punc-

tation dense and at least slightly coarser than that of prono-

Bonn zoological Bulletin 61 (1): 49-128

tum, deep and well-defined to shallow and ill-defined; in-

terstices without microsculpture and glossy. Hind wings

reduced (micropterous morph) or fully developed

(macropterous morph).

Abdomen widest at segment VI, distinctly broader than

(micropterous morph) or approximately as broad as ely-

tra (macropterous morph); punctation very fine and dense;

interstices with distinct microsculpture and subdued

shine; posterior margin of tergite VII with palisade fringe

in both morphs; posterior margin of tergite VIII convex

to indistinctly angled in the middle.

S: sternites III-V unmodified; sternite VI with shallow

median impression posteriorly; sternite VII strongly

transverse, in the middle with impression, this impression

impunctate in the middle and on either side of middle with

pubescence diagonally directed postero-mediad, posteri-

or margin broadly concave (Fig. 256); sternite VIII weak-

ly transverse and with median impression posteriorly, this

impression with numerous distinctly modified, short and

stout black setae, posterior excision relatively small, on

either side with dense black setae (Fig. 257); aedeagus ap-

proximately 1.1 mm long, with slender, straight, and api-

cally curved (lateral view) ventral process (Figs 258-259).

Comparative notes. Lobrathium duplex is similar to the

syntopic L. daxuense and L. bimembre, but distinguished

by larger body size, the longer and more slender anten-

nae, the more strongly convex posterior margin of the ab-

dominal tergite VII, and by the male sexual characters,

particularly the completely different shape of the ventral

process of the aedeagus, from L. bimembre additionally

by the less coarse and on average less dense punctation

of the head.

Distribution and natural history. The species is current-

ly known from the Diancang Shan (Yunnan), the Daxue

Shan (western Sichuan), and the Daliang Shan (southern

Sichuan) (Fig. 193). Most of the specimens were collect-

ed from under stones and from gravel in a stream valley

and in a stone pit; one male was sifted from moss and lit-

ter. The altitudes range from 2500 to 3000 m. Teneral

adults were found in August. One male and one female,

both from the same sample, are macropterous, the remain-

ing 58 specimens are micropterous.

Lobrathium domenoides sp. n. (Figs 141, 260-265)

Type material. Holotype ¢ [right middle and hind leg

missing]: “China S.Sichuan (Ya’an Pref., Shimian Co.) Xi-

aoxiang Ling, 7 km S Liziping, 35 km S Shimian, 28°56N,

102°18E, 1600 m (field ridge, bamboo) 7.VII.1999 D.W.

Wrase / Holotypus ¢ Lobrathium domenoides sp. n., det.

V. Assing 2011” (cAss).

©ZFMK

114 Volker Assing

267

266

272

274 275

Figs 266-275. Lobrathium lamellatum (266-270) and L. demptum (271-275). 266, 271: forebody; 267, 272: male sternite VII;

268, 273: male sternite VII; 269-270, 274-275: aedeagus in lateral and in ventral view. Scale bars: 266, 271: 1.0 mm; 267-270,

272-275: 0.5 mm.

Revision of East Palaearctic Lobrathium 115

Etymology. The specific epithet (adjective) alludes the

fact that the facies of this species somewhat resembles that

of species of the genus Domene Fauvel, 1873.

Description. Small species, body length 6 mm; length of

forebody 3.3 mm. Habitus as in Fig. 260. Coloration: body

blackish-brown; legs brown with blackish-brown femo-

ra; antennae reddish-brown with infuscate antennomere

Head (Fig. 261) approximately 1.1 times as long as broad,

of ovoid shape; lateral margins behind eyes evenly curv-

ing towards neck; posterior angles obsolete; punctation of

dorsal surface relatively fine and, except for the sparsely

punctate frons, very dense, not sparser on vertex than in

lateral dorsal portions. Eyes small, approximately one third

as long as distance from posterior margin of eye to neck

in dorsal view. Antenna long and slender, approximately

2.2 mm long; antennomeres VI-IX at least 1.5 times as

long as broad.

Pronotum (Fig. 261) strongly oblong, approximately

1.35 times as long as broad and approximately 0.9 times

as broad as head, distinctly tapering posteriad; punctation

distinctly sparser than that of head; interstices without mi-

crosculpture and glossy.

Elytra (Fig. 261) short and very slender, only 1.08 times

as wide and approximately 0.7 times as long as pronotum,

somewhat widened posteriad; humeral angles practically

obsolete; punctation dense and much coarser than that of

pronotum; interstices without microsculpture and glossy.

Hind wings reduced.

Abdomen widest at segment VI, 1.15 times as broad as

elytra; punctation very fine, barely noticeable, relatively

dense on tergites II-VI, sparser on tergite VII; interstices

with distinct microsculpture and subdued shine; posteri-

or margin of tergite VII with palisade fringe; posterior

margin of tergite VIII weakly convex.

&: sternites II-VI unmodified; sternite VII with shal-

low median impression, posterior margin weakly and

broadly concave (Fig. 262); sternite VIII weakly trans-

verse, median impression shallow and with numerous dis-

tinctly modified, short and stout black setae; posterior ex-

cision relatively broad and not very deep, margins semi-

membranous, on either side of excision with tuft of long

black setae (Fig. 263); aedeagus small, 0.75 mm long, with

blade-shaped ventral process (Figs 264-265).

Comparative notes. Lobrathium domenoides is charac-

terised by its small body size, the habitus (ovoid head,

strongly oblong pronotum, slender and short elytra), and

by the male sexual characters. For comments on species

group affiliations see the comparative notes in the sections

on L. wittmeri and L. cholaicum.

Distribution and natural history. The type locality is sit-

uated near Liziping in southern Sichuan province, China

Bonn zoological Bulletin 61 (1): 49-128

(Fig. 141). The holotype was collected in a field margin

at an altitude of 1600 m. The rather low altitude and the

presence of a distinct palisade fringe at the posterior mar-

gin of the abdominal tergite VII suggest that the species

may be wing-dimorphic and more widespread.

Lobrathium lamellatum sp. n. (Figs 180, 266-270)

Type material. Holotype ¢: “China: Sichuan, 24.7.1995,

28°07’N, 101°0S’E, 30 km NW Muli/Bowa, 3500 m, leg.

J. Turna / Holotypus 4 Lobrathium lamellatum sp. n., det.

V. Assing 2012” (NHMW).

Etymology. The specific epithet (adjective) alludes to the

thin ventral process of the aedeagus (lateral view), one of

the characters distinguishing this species from its closest

relatives.

Description. Species of moderate size, body length 7 mm;

length of forebody 3.8 mm. Coloration: body blackish;

legs dark-brown with reddish tarsi; antennae dark-brown.

Head (Fig. 266) approximately 1.05 times as long as

broad, of ovoid shape; lateral margins behind eyes even-

ly curving towards neck; posterior angles obsolete; punc-

tation of dorsal surface relatively coarse and, except for

the sparsely punctate frons, very dense, not distinctly

sparser on vertex than in lateral dorsal portions; dorsal sur-

face matt. Eyes small, little more than one fourth as long

as distance from posterior margin of eye to neck in dor-

sal view. Antenna moderately slender approximately 2.2

mm long; antennomeres VI-IX less than 1.5 times as long

as broad.

Pronotum (Fig. 266) moderately oblong, approximate-

ly 1.25 times as long as broad and approximately 0.9 times

as broad as head, distinctly tapering posteriad; punctation

distinctly sparser and somewhat coarser than that of head;

interstices without microsculpture and glossy.

Elytra (Fig. 266) short and narrow, only 1.05 times as

wide and approximately 0.7 times as long as pronotum,

slightly widened posteriad; humeral angles practically ob-

solete; punctation similar to that of pronotum, but shal-

lower and less defined; interstices without microsculpture

and glossy. Hind wings reduced.

Abdomen widest at segment VI, about 1.15 times as

broad as elytra; punctation fine, dense on tergites III—-VI,

slightly sparser on tergite VII; interstices with distinct mi-

crosculpture and subdued shine; posterior margin of ter-

gite VII without palisade fringe; posterior margin of ter-

gite VII weakly convex.

&: sternites II-VI unmodified; sternite VII with shal-

low median impression posteriorly, this impression with

about 15 modified, short and stout black setae posterior-

ly, anterior to these setae with a small area without pu-

bescence, posterior margin broadly concave (Fig. 267);

©OZFMK

116 Volker Assing

276 wr 577 282

Figs 276-289. Lobrathium partitum (276-279), L. semiflavum (280-285) and L. cribricolle, lectotype (286-289). 276, 282: male

sternite VII; 277, 283, 287: male sternite VII; 278-279, 284-285, 288-289: aedeagus in lateral and in ventral view; 280: habitus;

281, 286: forebody. Scale bars: 280-281, 286: 1.0 mm; 276-279, 282-285, 287-289: 0.5 mm.

Bonn zoological Bulletin 61 (1): 49-128 OZFMK

Revision of East Palaearctic Lobrathium 117

sternite VIII weakly transverse, median impression oblong

and with numerous distinctly modified, short and stout

black setae; posterior excision small (Fig. 267); aedeagus

1.0 mm long, with slender, slightly asymmetric, and, in

lateral view, very thin ventral process (Figs 269-270).

Comparative notes. Lobrathium lamellatum is charac-

terised by numerous characters, particularly its small eyes,

the extremely dense and relatively coarse punctation of

the head, the short elytra, the absence of a palisade fringe

at the posterior margin of tergite VII, the chaetotaxy of

the male sternite VII, and by the morphology of the aedea-

gus. For comments on species group affiliations see the

comparative notes in the sections on L. wittmeri and L.

cholaicum.

Distribution and natural history. The type locality is sit-

uated in the southwest of Sichuan province, not far from

the border with Yunnan (Fig. 180). Morphological char-

acters (small eyes, reduced wings, absence of a palisade

fringe) and the altitude of the type locality (3500 m) sug-

gest that the species may have a restricted distribution.

Lobrathium sp. n. 1

1Q [micropterous]: “China, S Sichuan 27.VII.1997,

Daliang Shan mts., road Meigu-Leibo, pass 15 km NE

Meigu, 28°25’N, 103°17’E, Jaroslav Turna leg.” (cSch).

This evidently undescribed species is similar to L. dax-

uense, but distinguished by broader habitus, slightly larg-

er size, and a denser and less coarse punctation of the head.

Lobrathium sp. n. 2

19: “China, W-Hubei, 20.6.—12.7.2003, Muyuping S eny.,

pitfall traps, 31.45N, 110.4E, ca. 1300 m, J. Turna”

(NHMW).

This species is distinguished from all described Chinese

representatives of the genus by the following characters

combination: body size moderate, 6.5 mm; length of fore-

body 4.0 mm; punctation of head, except for the sparse-

ly punctate frons, extremely dense, without sparsely im-

punctate patches, and matt; eyes large and protruding, lat-

eral margins of head smoothly curved towards neck be-

hind eyes; pronotum slender; coloration of forebody black-

ish-brown, elytra indistinctly reddish in posterior 2/3.

Bonn zoological Bulletin 61 (1): 49-128

SPECIES FROM OTHER EAST PALAEARCTIC

REGIONS

Lobrathium partitum (Sharp, 1874) (Figs 276-279)

Lathrobium partitum Sharp, 1874: 56 f.

Type material examined. Syntype ¥ [dissected prior to

present study]: “Japan / Lathrobium partitum type D.S. /

Type / Japan. G. Lewis. / Sharp Coll 1905-313. / Lecto-

typus Lathrobium partitum Sharp, V. Gusarov des. 1992

/ Lobrathium partitum (Sharp) 2, V.I. Gusarov det. 1992

/ Syntype Lathrobium partitum Sharp, rev. V. Assing 2012

/ Lobrathium partitum (Sharp), det. V. Assing 2012”

(BMNH).

Comment. The original description is based on an unspec-

ified number of syntypes from “under tidal refuse, Nagasa-

ki harbour” (Sharp 1874). It can be inferred from Sharp’‘s

statement that the species is “very local” that he had on-

ly few syntypes, possibly only one specimen before him.

The single syntype in the Sharp collection at the BMNH,

a female, has a lectotype label attached to it, but the des-

ignation has never been published. In view of the fact that

Lobrathium species are usually reliably identified only

based on the male sexual characters, the above type spec-

imen is not designated as the lectotype.

Additional material examined. Japan: Kyushu: 13, Nagasa-

ki Pref., Kunimi, Uematsu, 3.XI.1982, leg. Imasaka (clto).

Shikoku: 1¢: Ehime, Oku-Dogo, 22.XII.1991, leg. Okamoto

(cAss). Honshu: 1 [teneral], Fukui Pref., Mana-gawa river,

1. VIIIL.1965, leg. Hayashi (cAss); 14 [teneral], Wakayama Pref.,

Mt. Iwawaki, 9.VI.1968, leg. Kimura (cAss); 14, Yokohama

(BMNH).

Redescription. Body length 7.0—8.0 mm; length of fore-

body 3.54.1 mm. External characters as in L. taureum.

Distinguished only by the male sexual characters.

&: sternite V with small median depression posterior-

ly; sternite VI with moderately small and shallow medi-

an impression posteriorly; sternite VII strongly transverse

and with distinct median impression with pubescence, pos-

terior margin broadly concave (Fig. 276); sternite VHI

weakly transverse, with pronounced long median impres-

sion, this impression with numerous modified, short and

stout black setae, posterior excision moderately small and

narrowly V-shaped (Fig. 277); aedeagus approximately

1.3—1.4 mm long, ventral process symmetric and of dis-

tinctive shape, in the middle distinctly dilated in ventral

view (Figs 278-279).

Comparative notes. In external characters, L. partitum

is highly similar to L. taureum, L. schuelkei, and L. semi-

flavum. The similar general morphology of the aedeagus

as well as the similar male secondary sexual characters

suggest that the species is most closely related to L. semi-

©ZFMK

118 Volker Assing

291

300 301 303 = 304

290

293 295

Figs 290-304. Lobrathium cribricolle, lectotype (290), L. nudum (291-294), L. amamiense (295-298), L. indubium, syntype (299),

and L. discrepans (300-304). 290, 295, 302: male sternite VH; 291, 300: habitus; 292, 299, 301: forebody; 293, 296: male stern-

ite VIH; 294, 297-298, 303-304: aedeagus in lateral and in ventral view. Scale bars: 291-292, 299-301: 1.0 mm; 290, 293-398,

302-304: 0.5 mm.

Revision of East Palaearctic Lobrathium 119

flavum, from which it is reliably distinguished only by the

shape of the ventral process of the aedeagus both in lat-

eral and in ventral view (L. semiflavum: not distinctly di-

lated in the middle in ventral view) and the slightly nar-

rower posterior excision of the male sternite VIII.

Distribution. Lobrathium partitum was originally de-

scribed from the environs of Nagasaki, Kyushu, southern

Japan, and subsequently also reported from Honshu

(Kameyama et al. 2006); for additional records from these

islands and from Shikoku see the additional material

above.

Lobrathium semiflavum sp. n. (Figs 193, 280-285)

Type material. Holotype ¢ [teneral]: “Russia or: Pri-

morie, Ussurijsky Res., Komarovo Zapovednoe,

132°20°40”E, 43°38°48”N, 20.—29.VII.1999, leg. J.

Sundukov / Holotypus ¢ Lobrathium semiflavum sp. n.,

det. V. Assing 2011” (cAss). Paratypes: 14, 19 [teneral]:

same data as holotype (cSch).

Etymology. The specific epithet (Latin, adjective) refers

to the fact that nearly all of the posterior half of the ely-

tra is bright yellowish.

Description. Body length 5.8—7.8 mm; length of forebody

3.54.0 mm. Habitus as in Fig. 280. Coloration and oth-

er external characters as in L. partitum, except as follows:

Elytra without bluish hue and on average slightly short-

er and narrower (Fig. 281); appendages paler: legs dark-

yellowish; antennae pale-reddish (but note that all the type

specimens are somewhat teneral).

S: sternite V with very small and shallow impression

posteriorly; sternite VI with moderately small and mod-

erately shallow impression posteriorly; sternite VII

strongly transverse and with distinct median impression

with pubescence, posterior margin broadly concave (Fig.

282); sternite VIII weakly transverse, with pronounced

long median impression, this impression with numerous

modified, short and stout black setae, posterior excision

relatively small and almost V-shaped (Fig. 283); aedea-

gus 1.2—-1.4 mm long, ventral process symmetric and of

distinctive shape (Figs 284-285).

Comparative notes. Based on the similar external and

male sexual characters, L. semiflavum is presumably the

adelphotaxon of L. partitum. For distinguishing charac-

ters see the description above and the comparative notes

in the preceding section.

Distribution and natural history. The type locality is sit-

uated in Primorskiy Kray, Russian Far East (Fig. 193). All

type specimens were collected in July and are teneral.

Bonn zoological Bulletin 61 (1): 49-128

Lobrathium cribricolle (Sharp, 1889) (Figs 286-290)

Lathrobium cribricolle Sharp, 1889: 256.

Type material examined. Holotype 3: ““Lathrobium crib-

ricolle. Type D.S. [written on mounting label] / Japan. G.

Lewis. 1910-320. / Suyama. 20.IV.—22.IV.80. / Type /

aedeagus at rest is turned to 90° in abdomen / Lobrathi-

um cribricolle (Sharp) @, V.I. Gusarov det. 1992 / Holo-

typus ¢ Lathrobium cribricolle Sharp, V. Gusarov des.

[sic] 1992 / Lobrathium cribricolle (Sharp), det. V. Ass-

ing 2012” (BMNH).

Comment. The original description is based on a

“unique” holotype from the “Plain under Fujisan” (Sharp

1889).

Redescription. Body length 6.6 mm; length of forebody

3.5 mm. Coloration: body blackish, elytra with yellowish

spot posteriorly; legs and antennae pale-reddish.

Head (Fig. 286) approximately 1.05 times as long as

broad, of subrectangular shape; lateral margins behind

eyes subparallel in dorsal view; posterior angles moder-

ately marked; punctation conspicuously coarse and dense,

even in median dorsal portion; interstices reduced to nar-

row ridges. Eyes bulging and large, approximately

0.6—0.7 times as long as distance from posterior margin

of eye to neck in dorsal view. Antenna relatively short, ap-

proximately 1.8 mm long.

Pronotum (Fig. 286) approximately 1.25 times as long

as broad and 0.95 times as broad as head; punctation con-

spicuously coarse and dense, even along midline; only in

posterior median portion with small impunctate patch.

Elytra (Fig. 286) 1.07 times as long as pronotum; punc-

tation coarse, arranged in irregular series only laterally.

Hind wings apparently fully developed.

Abdomen subparallel and slightly narrower than elytra;

punctation fine and moderately dense, slightly less dense

on tergites VII and VII]; interstices with distinct transverse

microsculpture; posterior margin of tergite VII with pal-

isade fringe.

3: posterior margin of tergite VIII convex in the mid-

dle; sternite VI with shallow impression posteriorly; ster-

nite VI with shallow median impression posteriorly, this

impression with few (approximately 15—20) modified,

short and stout black setae, posterior margin broadly and

rather weakly concave (Fig. 290); sternite VII weakly

transverse, with oblong median impression, this impres-

sion with relatively sparse modified, short and stout black

setae, posterior excision not very deep and of trapezoid

shape (Fig. 287); aedeagus 1.3 mm long, with long ven-

tral process of highly distinctive shape (Figs 288-289).

Comparative notes. Lobrathium cribricolle is charac-

terised particularly by the coarse and dense punctation of

the head and pronotum, by the shape and chaetotaxy of

©ZFMK

120 Volker Assing

the male sternites VII and VII, as well as by the distinc-

tive shape of the ventral process of the aedeagus.

Distribution. The distribution of L. cribricolle is confined

to Japan (Honshu). For additional records see Ito (1996a).

Lobrathium nudum (Sharp, 1889) (Figs 291—294)

Lathrobium nudum Sharp, 1874: 55 f.

Type material examined. Lectotype ¢ and paralectotype

° [glued on same label]: “Japan / Japan. G. Lewis. / Lath-

robium nudum Type D.S. / Sharp Coll 1905-313. / Type

/ aedeagus at rest is turned to 90° in abdomen / Lobrathi-

um nudum (Sharp) 3, V.I. Gusarov det. 1992 / Lectotype

3 Lathrobium nudum Sharp, V. Gusarov des. 1992 / Para-

lectotype 2 Lathrobium nudum Sharp, V. Gusarov des.

1992 / Lobrathium nudum (Sharp), det. V. Assing 2012”

(BMNH).

Comment. The original description is based on an unspec-

ified number of syntypes (“not very rare”), among them

at least one male, from “Nagasaki” (Sharp 1874). In re-

ferring to the above material as lecto- and paralectotype

and illustrating the aedeagus as that of the lectotype, Ito

(1996a) designated the male as the lectotype. The two type

specimens have lectotype and paralectotype labels by V.

Gusarov attached to them, who never published this des-

ignation.

Redescription. Body length 7.0—.7.5 mm; length of fore-

body 3.4-3.6 mm. Habitus as in Fig. 291. Coloration:

head, pronotum, and abdomen blackish; elytra dark-brown

to blackish-brown, posteriorly with a large yellowish spot

reaching posterior margin; legs reddish, with the profemo-

ra and sometimes also the protibia darker; antennae red-

dish.

Head (Fig. 292) approximately as long as broad, weak-

ly narrowed behind eyes in dorsal view; posterior angles

moderately marked; punctation moderately coarse and

moderately dense, rather sparse in median dorsal portion

and on frons. Eyes moderately convex and large, approx-

imately 0.6—0.7 times as long as the distance from poste-

rior margin of eye to neck in dorsal view. Antenna rela-

tively short, approximately 1.8 mm long.

Pronotum (Fig. 292) 1.25—1.30 times as long as broad

and approximately 0.95 times as broad as head; puncta-

tion similar to that of head; midline impunctate.

Elytra (Fig. 292) approximately as long as pronotum;

punctation coarse, arranged in irregular series. Hind wings

fully developed.

Abdomen subparallel and slightly narrower than elytra;

punctation fine and moderately dense; interstices without

distinct microsculpture, except for shallow traces in pos-

terior portions of tergites VII and VII; posterior margin

Bonn zoological Bulletin 61 (1): 49-128

of tergite VII with palisade fringe; posterior margin of ter-

gite VIII convex in both sexes.

3: sternite VI unmodified; sternite VII anteriorly with

pair of tubercles, in the middle with large impression with-

out pubescence, posterior margin broadly and distinctly

concave; sternite VII approximately as broad as long, in

the middle with oblong impression without modified se-

tae, and with relatively deep and V-shaped posterior ex-

cision (Fig. 293); aedeagus approximately 1.0 mm long

and with ventral process of distinctive shape (Fig. 294).

Comparative notes. Lobrathium nudum is characterised

particularly by the relatively large yellowish spot on the

elytra (reaching posterior margin), by the modifications

of the male sternites VII and VIII, as well as by the shape

of the ventral process of the aedeagus.

Distribution. Lobrathium nudum has been recorded from

several localities in Japan (Kyushu, Honshu); for addition-

al records see Ito (1996a), Watanabe (1998c), and Watan-

abe & Shibata (1972). Watanabe & Onoda (1997) report

the species from Kuroshima Island (Ryukyu Islands). It

has been reported also from the Russian Far East

(Smetana 2004), but this record should be considered

doubtful and requires confirmation. In the course of the

present revision, none of the species from the main islands

of Japan was recorded also from continental Asia. The on-

ly species whose presence in both Japan and continental

Asia is confirmed is L. hongkongense, whose distribution

in Japan is confined to the extreme south (Ryukyu Islands).

Lobrathium amamiense Ito, 1996 (Figs 295-298)

Lobrathium amamiense Ito, 1996: 116 ff.

Type material examined. Paratype ¢ [slightly teneral]:

“Hatsuno, Amami Is., 4.IV.1966, T. Ito / Paratype Lo-

brathium amamiense T. Ito. sp. nov. “ (cAss).

Comment. The original description of L. amamiense is

based a male holotype and eleven paratypes from three

localities in the “Amami-Oshima Is., Kagoshima Pref.”

(Ito 1996). For a detailed description of the external char-

acters see Ito (1996).

Redescription of the male sexual characters. Posterior

margin of tergite VIII obtusely pointed in the middle; ster-

nite VII with shallow median impression and without

modified pubescence, posterior margin broadly concave

(Fig. 295); sternite VIII weakly transverse, in posterior half

with well-defined oblong median impression, this impres-

sion with numerous modified, short and stout black setae,

posterior excision moderately deep and moderately broad

(Fig. 296); aedeagus approximately 1.2 mm long, ventral

process blade-shaped and large, symmetric, subapically

OZFMK

Revision of East Palaearctic Lobrathium 121

with median carina, and basally with few lateral teeth (Figs

297-298).

Comparative notes. Lobrathium amamiense 1s distin-

guished from its congeners particularly by the shape and

chaetotaxy of the male sternite VIII, by the shape of the

ventral process of the aedeagus, and from most other

macropterous species distributed in the East Palaearctic

also by the coloration of the elytra (diffusely dark-reddish

in posterior half, without defined spots).

Distribution and natural history. As far as is currently

known, the distribution of L. amamiense is confined to the

Amami Islands, southern Japan. The examined paratype

is slightly teneral.

Lobrathium varium Ito, 1995, stat. n.

Lobrathium shibatai varium Ito, 1995: 43.

Material examined. Japan: 17 exs. [previously labelled as types

of L. kasuganum Gusarov 1.1., labels removed; partly teneral],

Honshu, Nara, foot of Mt. Kasuga, 20. VIII.1980, leg. Hammond

(BMNH, cAss).

Comment. According to the original descriptions of L.

shibatai shibatai and L. shibatai varium, these taxa are

subspecies distinguished by slight differences in the shape

of the ventral process of the aedeagus, the slightly differ-

ent shapes of the impressions of the male sternites VII and

VIII, and the slightly more slender body of L. varium (Ito

1995). However, zoogeographic evidence (see distribution

map in Ito 1995) renders the hypothesis that both morphs

should represent subspecies highly implausible; both

morphs were described from central Honshu. According

to Ito (e-mail, 3.11.2012), who has seen more material in

the meantime, the distributions of both taxa strongly over-

lap, so that they should be considered distinct species

rather than subspecies.

Lobrathium indubium (Eppelsheim, 1893) (Fig. 299)

Lathrobium indubium Eppelsheim, 1893: 52 f.

Platydomene altaicus [sic] Coiffait, 1967: 355 f.; synonymy by

Shavrin (2008).

Type material examined. Syntypes: 29 9: “Ost-Sibirien.

Quellgebiet des Irkut. Leder 1891. / indubium Epp. / c.

Epplsh. Steind. d. / Typus [one labelled as lectotype, the

”

other as paralectotype by V. Gusarov]” (NHMW).

Comment. The original description, which is contained

in a work on the southwestern Baikal region, is based on

two females without specification of the locality (Ep-

pelsheim 1893). Both syntypes are deposited in the Ep-

Bonn zoological Bulletin 61 (1): 49-128

pelsheim collection at the NHMW. They have (para-)lec-

totype labels by V. Gusarov attached to them, but this des-

ignation was never published.

Redescription. Body length: 6.0—6.8 mm; length of fore-

body 3.4-3.9 mm. Coloration: forebody reddish to red-

dish-brown, with the elytra often somewhat paler; elytra

often with indistinct reddish spots posteriorly and/or with

yellowish posterior margins; abdomen dark-brown; legs

and antennae dark-reddish.

Head (Fig. 299) somewhat flattened, as long as broad

or weakly oblong; punctation moderately coarse and

dense; median dorsal portion with few scattered punctures,

almost impunctate. Eyes weakly convex and small, ap-

proximately 1/3 the length of postocular region in dorsal

view. Antenna 2.0—2.4 mm long.

Pronotum (Fig. 299) weakly convex in cross-section, al-

most flattened, and slender, approximately 1.3 times as

long as broad and 0.9 times as broad as head; punctation

similar to that of head, but slightly less dense; midline with

broad and complete impunctate band.

Elytra (Fig. 299) weakly convex in cross-section, ap-

proximately as long as pronotum, or nearly so; punctation

dense and moderately defined, not arranged in distinct se-

ries.

Abdomen approximately as broad as elytra; posterior

margin of tergite VII with palisade fringe; posterior mar-

gin of tergite VUI broadly and rather weakly convex.

¢: sternite VIII with long median impression; this im-

pression with numbers strongly modified, short and stout

black setae, posterior excision of moderate size; aedeagus

approximately 1.3 mm long, with long, symmetric,

basally dilated (ventral view), and apically acute ventral

process. For illustrations of the aedeagus see Coiffait

(1967, 1982c) (as Platydomene altaicus [sic]).

Comparative notes. Lobrathium indubium somewhat re-

sembles L. multipunctum in size and coloration, but is dis-

tinguished from that species by the flatter body and the

irregular punctation of the elytra.

Distribution. The known distribution of this species in-

cludes the Baikal region (Altai, East Sayan) and the Russ-

ian Far East (Primorskiy Kray) (Assing 2007, Schiilke

1990, Shavrin 2008).

Lobrathium discrepans sp. n. (Figs 140, 300-305)

Type material: Holotype ¢: ““N-Vietnam, 120 km SW

Hanoi, Cuc Phuong National Park, 13.04.2010, leg. A.

Kleeberg / Holotypus ¢ Lobrathium discrepans sp. n., det.

V. Assing 2011” (cAss).

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122 Volker Assing

311 312

307 (308

309

305

Figs 305-314. Lobrathium discrepans (305) and Tetartopeus gracilentus (306-314). 305, 308: male sternite VII; 306: forebody;

307: male tergite VII; 309: posterior portion of female tergite VII; 310: female sternite VIII; 311-313: aedeagus in lateral, ven-

tral, and in dorsal view; 314: internal structures of aedeagus extruded in lateral view. Scale bars: 306: 1.0 mm; 305, 307-313: 0.5

mm; 314: 0.2 mm.

Etymology. The specific epithet (Latin, adjective: deviant)

refers to the fact that this species differs in many ways

from the usual morphology of Lobrathium species.

Description. Large species, body length 10.3 mm; length

of forebody 5.6 mm. Habitus as in Fig. 300. Coloration:

body blackish, except for the reddish posterior margins of

the abdominal segments VII and VIII; forelegs blackish

with reddish tarsi; middle and hind legs with pale-yellow-

ish femora, infuscate tibiae, and reddish tarsi; antenna in

basal half blackish-brown, with antennomere I black and

II dark-reddish; antennomeres in apical half gradually be-

coming reddish.

Bonn zoological Bulletin 61 (1): 49-128

Head (Fig. 301) approximately as long as broad, pos-

terior angles broadly convex; punctation moderately

coarse and extremely dense; interstices reduced to narrow

ridges. Eyes large, approximately 0.7 times as long as the

distance from posterior margin of eye to neck in dorsal

view. Antenna slender, approximately 3.2 mm long.

Pronotum (Fig. 301) short and broad, 1.12 times as long

as broad and approximately as broad as head; punctation

as dense as, and slightly coarser than head; impunctate

midline reduced, narrow rudiment present only in poste-

rior half.

Elytra (Fig. 301) as long as pronotum, with parallel mar-

gins in dorsal view, and marked humeral angles; puncta-

©ZFMK

Revision of East Palaearctic Lobrathium 123

tion very coarse and arranged in irregular series. Hind

wings fully developed. Metatarsomere I little more than

half as long as the elongate metatarsomere II.

Abdomen subparallel and distinctly narrower than ely-

tra; punctation very fine and dense; interstices with dis-

tinct microsculpture; posterior margin of tergite VII with

palisade fringe; posterior margin of tergite VIII obtusely

angled in the middle.

&: sternite VII with shallow median impression poste-

riorly, pubescence unmodified, posterior margin weakly

and broadly concave (Fig. 302); sternite VIII as long as

broad, median impression narrow, rather long, and with

approximately 12 distinctly modified, short and stout black

setae, posterior excision deep and almost V-shaped (Fig.

305); aedeagus 1.8 mm long; ventral process conspicuous-

ly bifid, subapically with pronounced ventral teeth (Figs

303-304).

Comparative notes. Among its congeners, L. discrepans

is characterised not only by the male sexual characters,

but also by its large size, the extremely dense punctation

of the head and the pronotum, the broad and short prono-

tum, the conspicuous coloration of the legs, the shape of

the elytra, and by the morphology of the tarsi.

Distribution and natural history. The type locality is sit-

uated some 120 km to the southwest of Hanoi in North

Vietnam (Fig. 140). According to the collector, the type

specimen was found in dead banana trunks saturated with

water, together with Zachinomorphus fulvipes (Erichson,

1840) and Coproporus sp. (Kleeberg, pers. comm.).

SPECIES BELONGING TO OTHER GENERA

Tetartopeus gracilentus (Kraatz, 1859), comb. n. (Figs

306-314)

Lathrobium gracilentum Kraatz, 1859: 115 f.

Type material examined. Lectotype <, present designa-

tion: “Ceylon / Coll. Kraatz / Syntypus / Coll. DEI Ebers-

walde / Lathrobium gracilentum (Kr.) [sic], V.I. Gusarov

det. 1994 / DEI Mincheberg, Col — 02763 / Lectotypus

& Lathrobium gracilentum Kraatz, desig. V. Assing 2011

/ Tetartopeus gracilentus (Kraatz), det. V. Assing 2011”

(SDEI). Paralectotypes: 14, 32 9: same data as lectotype,

but “Col — 02764”, “Col — 02765”, “Col — 02766”, “Col

— 02767”, respectively (SDEI).

Comment. The original description is based on an unspec-

ified number of syntypes from “Ceylan” (Kraatz 1859).

Five syntypes, two males and three females, are deposit-

ed in the Kraatz collection at the SDEI. All of them had

lectotype or paralectotype labels by V. Gusarov (“‘des.

1993”) attached to them. Since a lectotype designation was

Bonn zoological Bulletin 61 (1): 49-128

never published, these labels were removed and new

(para-)lectotype labels were attached to the specimens. The

lectotype is illustrated in Figs 306 and 314. Figs 307-313

are based on paralectotypes.

The species undoubtedly belongs to the genus Tetar-

topeus Czwalina, 1888 and represents clear evidence that

the range of the genus, which was previously considered

to be Holarctic (Assing 2011b), reaches deeply into the

Oriental region.

Redescription. Body length 5.5—6.0 mm; length of fore-

body 3.1—3.4 mm. Coloration: body blackish-brown to

black, elytra with distinct yellowish spots in postero-lat-

eral angles; legs dark-yellowish to yellowish-brown with

darker profemora and protibiae; antennae brown with red-

dish apical and basal antennomeres.

Head (Fig. 306) approximately 1.15 times as long as

broad, widest across eyes; posterior angles not marked;

punctation fine and very dense, dorsal surface almost matt;

interstices reduced to narrow ridges; median dorsal por-

tion often with small impunctate patch. Eyes large and

bulging, approximately 0.7 times as long as distance from

posterior margin of eye to neck in dorsal view. Antenna

long and slender, approximately 2.4 mm long.

Pronotum (Fig. 306) approximately 1.3 times as long

as broad and as wide as head; punctation coarser and less

dense than that of head; interstices without microsculp-

ture and glossy.

Elytra (Fig. 306) approximately 0.95 times as long as,

and distinctly broader than pronotum, humeral angles

marked; punctation dense and slightly finer than that of

pronotum; interstices without microsculpture and glossy.

Hind wings fully developed.

Abdomen narrower than elytra; punctation very fine and

very dense, dorsal surface practically matt; posterior mar-

gin of tergite VII with palisade fringe.

¢: tergite VIII obtusely angled posteriorly (Fig. 307);

sternite VHI with small posterior excision in somewhat

asymmetric position (Fig. 308); aedeagus approximately

0.9 mm long, shaped as in Figs 311-314.

the middle (Fig. 309); sternite VIII much longer than ter-

gite VIII, its posterior margin convexly produced in the

middle (Fig. 310); tergite X distinctly shorter than tergite

IX in the middle.

Comparative notes. The species is highly similar and ev-

idently closely related to 7. fragilis (Sharp, 1889), T: pal-

lipes (Sharp, 1889), and 7? wui (Zheng, 2001), but distin-

guished by the larger eyes, head shape (posterior angles

practically obsolete), the straight ventral process of the

aedeagus, by the shapes of the internal structures of the

aedeagus, and by the posteriorly acutely produced female

tergite VIII.

OZFMK

124 Volker Assing

315

321

324

Figs 315-328. Lobrathium lobrathioides (315-320) and L. lobrathiforme (321-328). 315, 321: habitus; 316, 322: forebody; 317,

323: male sternite VII; 318, 324: male sternite VIII; 319-320, 325-326: aedeagus in lateral and in ventral view; 327: female ster-

nite VIII; 328: female tergites [Xx—X. Scale bars: 315-316, 321-322: 1.0 mm; 317-320, 323-328: 0.5 mm.

Bonn zoological Bulletin 61 (1): 49-128 OZFMK

Revision of East Palaearctic Lobrathium 125

Distribution. This species has become known only from

Sri Lanka.

Tetartopeus wui (Zheng, 2001), comb. n.

Lobrathium wui Zheng, 2001: 324 f.

Lobrathium bimaculatum Li, Tang & Zhu, 2007: 261 f.; syn. n.

Comment. The original description of Lobrathium wui is

based on a male holotype from “Shanmuping, 780 m,

Tianmushan, Zhejiang” and three paratypes from

“Chanyuansi” (Zheng 2001).

Lobrathium bimaculatum was transferred to Tetartopeus

by Assing (2010). The external and sexual characters were

illustrated and additional records from China, among them

also a male from Tianmushan, and Taiwan were reported

by Assing (2011b).

As can be inferred from the illustration of the aedeagus

of L. wui, the species belongs to 7Zetartopeus and there is

little doubt that it is conspecific with 7) bimaculatus.

Hence the synonymy proposed above.

“Lathrobium” sublaeve Motschulsky, 1858

Lathrobium sublaeve Motschulsky, 1858: 647.

Comment. Based on the morphological details specified

in the original description and on the type locality (“In-

des orientales”), this species is most unlikely to belong to

either Lathrobium or Lobrathium. Motschulsky (1858)

compares L. sublaeve with “Sunius filiformis”, today

Astenus procerus (Gravenhorst, 1802), suggesting that L.

sublaeve is not even a member of Lathrobiina.

Lathrobium lobrathioides sp. n. (Figs 315—320)

Type material. Holotype @: “China: SE Sichuan, Jinfo

Shan, 29°01N, 107°14E, 1800 m, 27.VI.1998, A. Smetana

[C70] / 1998 China Expedition J. Farka¢, D. Kral, J.

Schneider & A. Smetana / Holotypus ¢ Lathrobium lo-

brathioides sp. n., det. V. Assing 2011” (cAss).

Etymology. The specific epithet (Latin, adjective) alludes

to the external resemblance of this species with species

of the genus Lobrathium.

Description. Body length 7.2 mm; length of forebody 4.1

mm. Habitus as in Fig. 315. Coloration: body blackish,

elytra with large reddish spot posteriorly reaching lateral

and posterior margins; legs reddish with darker femora;

antennae reddish, with slightly darker antennomere I.

Head (Fig. 316) transverse, 1.15 times as wide as long,

widest across eyes; posterior angles broadly convex, very

weakly marked; punctation coarse and, except in median

Bonn zoological Bulletin 61 (1): 49-128

dorsal portion and on frons, very dense; interstices with

shallow microsculpture. Eyes large and bulging, somewhat

shorter than distance between posterior margin of eye and

neck, but distinctly more than half as long as this distance

in dorsal view. Antenna not particularly slender, 2.1 mm

long.

Pronotum (Fig. 316) 1.17 times as long as broad and

0.92 times as broad as head; punctation distinctly spars-

er and coarser than that of head; interstices without mi-

crosculpture and very glossy.

Elytra (Fig. 316) 0.93 times as long as, and distinctly

broader than pronotum, lateral margins subparallel in dor-

sal view; humeral angles marked; punctation coarse and

rather dense; interstices without microsculpture and

glossy. Hind wings apparently fully developed.

Abdomen approximately as broad as elytra; punctation

relatively coarse and dense on anterior, finer and sparser

on posterior tergites; interstices with very shallow trans-

verse microsculpture and glossy; posterior margin of ter-

gite VII with palisade fringe; posterior margin of tergite

VIII distinctly convex.

3: sternite VII with small and shallow median impres-

sion without modified pubescence, posterior margin

broadly and shallowly concave (Fig. 317); sternite VIII

weakly transverse, with shallow median impression pos-

teriorly, this impression without modified setae, posteri-

or excision small and not very deep (Fig. 318); aedeagus

1.15 mm long, ventral process simple, dorsal plate lamel-

late and weakly sclerotised (Figs 319-320).

Comparative notes. Lathrobium lobrathioides is distin-

guished from other Lathrobium species particularly by its

conspicuous coloration, the strongly transverse head with

large and bulging eyes, and by the male sexual characters.

It is additionally separated from externally similar Lo-

brathium species by the absence of a submarginal carina

of the elytra.

Distribution and natural history. The type locality is sit-

uated in the Jinfo Shan in the south of Chongqing

province, close to the border with Guizhou. The holotype

was collected in an old deciduous forest at an altitude of

1800 m, by sifting moist to wet leaf litter and humus

(Smetana pers. comm.).

Lathrobium lobrathiforme sp. n. (Figs 321-328)

Type material. Holotype @: “China: Yunnan [CH07—24],

Nujiang Lisu Aut. Pref., Gaoligong Shan, valley 18 km

W Gongshan, 3020 m, 27°47°54”N, 98°30°13”E, mixed

forest, litter, moss, wood sifted, 7.VI.2007, M. Schiilke /

Holotypus ¢ Lathrobium lobrathiforme sp. n., det. V. Ass-

ing 2011” (cAss). Paratypes: 299 [1 without head and

pronotum]: same data as holotype (cSch).

OZFMK

126 Volker Assing

Etymology. The specific epithet (Latin, adjective) alludes

to the external resemblance of this species with species

of the genus Lobrathium.

Description. Body length 7.5—8.0 mm; length of forebody

3.7-3.9 mm. Habitus as in Fig. 321. Coloration: body

blackish, elytra with moderately large, transverse reddish

spot in posterolateral angles reaching lateral and posteri-

or margins, but not suture; legs reddish-yellow to reddish

with slightly darker femora; antennae dark-brown to black-

ish-brown.

Head (Fig. 322) transverse, approximately 1.15 times

as wide as long, widest across eyes; posterior angles

broadly convex, weakly marked; dorsal surface uneven,

with shallow lateral impressions; punctation coarse and,

except on frons, very dense; interstices with shallow m1-

crosculpture. Eyes large and bulging, almost as long as dis-

tance between posterior margin of eye and neck in dorsal

view. Antenna not particularly slender, 1.9—2.2 mm long.

Pronotum (Fig. 322) approximately 1.1 times as long

as broad and about 0.95 times as broad as head; puncta-

tion distinctly sparser and slightly coarser than that of

head; interstices without microsculpture and very glossy.

Elytra (Fig. 322) approximately 0.9 times as long as, and

distinctly broader than pronotum, lateral margins subpar-

allel in dorsal view; humeral angles marked; punctation

coarse and rather dense; interstices without microsculp-

ture and glossy. Hind wings fully developed.

Abdomen approximately as broad as elytra; punctation

relatively coarse and dense on tergites III—VI, finer and

sparser on tergite VII; interstices with very shallow trans-

verse microsculpture and glossy; posterior margin of ter-

gite VII with palisade fringe; posterior margin of tergite

VIII distinctly convex.

&: sternites I[V—VI with shallow median impressions,

with dense asperate punctation and dense black setae; ster-

nite VI with median impression and dense black setae,

posterior margin broadly and shallowly concave (Fig.

323); sternite VIII weakly transverse, with median impres-

sion posteriorly, posterior excision small and not very deep

(Fig. 324); aedeagus 1.5 mm long, ventral process apical-

ly acutely pointed in ventral view, dorsal plate lamellate

and weakly sclerotised (Figs 325-326).

@: sternite VII with broadly convex posterior margin

(Fig. 327); anterior portion of tergite [X divided in the

middle, tergite X approximately twice as long as tergite

IX in the middle (Fig. 328).

Comparative notes. Both in external and sexual charac-

ters, L. lobrathiforme is similar to L. lobrathioides. It is

distinguished from this species by the smaller and more

transverse elytral spots, the coarser punctation of the head,

the much denser punctation of the pronotum, the chaeto-

taxy of the male sternite VII, the deeper posterior exci-

sion of the male sternite VIII, and by the morphology of

Bonn zoological Bulletin 61 (1): 49-128

the aedeagus (much longer and more slender, ventral

process apically acutely pointed in ventral view).

Distribution and natural history. The type locality is sit-

uated in the Gaoligong Shan, Yunnan province, China. The

type specimens were sifted from litter and moss in a mixed

forest at an altitude of 3020 m.

Acknowledgements. I am indebted to the colleagues indicated

in the material section for the loan of material under their care.

Special thanks are due to Benedikt Feldmann (Minster), Andreas

Kleeberg (Berlin), Michael Schiilke (Berlin), and Ale’ Smetana

(Ottawa) for the generous permission to retain several holotypes

and single males needed for future reference purposes. More-

over, I am grateful to Tateo Ito (Kyoto) for the generous gift of

paratypes of L. amamiense and L. ryukyuense, the gift of mate-

rial of L. partitum, and help with the identification of Japanese

localities. Lee H. Herman (New York) cross-checked the names

treated in an early draft against his unpublished catalogue of

Paederinae. Alfred F. Newton (Chicago) cross-checked the cat-

alogue and the abstract of the final draft against his unpublished

database. Harald Schillhammer (NHMW) assisted with the iden-

tification of localities in Myanmar. Benedikt Feldmann proof-

read the manuscript.

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Bonn zoological Bulletin 61 (1): 129-134

July 2012

The urticating apparatus in the larva of the Lappet Moth,

Streblote panda Hubner, 1820

(Lepidoptera: Lasiocampidae)

Michel Joél Faucheux

Laboratoire d’Endocrinologie des Insectes Sociaux, Faculté des Sciences et des Techniques,

2 rue de la Houssiniére, B. P. 92208 F-44322 Nantes Cedex 3, France.

Abstract. A morphological study of the urticating apparatus in the last larval instar of Streblote panda Hubner, 1820 (Lep-

idoptera: Lasiocampidae) was undertaken using a scanning electron microscope. It is composed of two meso- and metatho-

racic crevices, accompanied by urticating hairs, which open when the caterpillar is threatened. The urticating hairs taper

at both ends and are mostly smooth but ringed on their distal part, which also features about 400 pores allowing the ur-

ticating liquid to be released.

Key words. Lasiocampidae, Streblote panda, scanning electron microscopy, urticating hairs.

INTRODUCTION

The Lappet Moth Streblote panda Hubner, 1820 (Lasio-

campidae: Lasiocampinae) is distributed in Spain and

North Africa. In Morocco, it is a common species on the

plains and lower mountains of the Atlantic coast, the re-

gion of Souss (Southwestern Morocco) and the Western

and Central High Atlas (Rungs 1981) where adults are ac-

tive in January, June—July and October. Streblote panda

is an eremic species that prefers littoral, sandy and open

scrub areas (Calvo & Molina 2008). The caterpillars are

highly polyphagous, feeding on the leaves of different

plants from a broad spectrum of plant families such as Fa-

gaceae, Euphorbiaceae, Myrtaceae, Fabaceae, Rosaceae,

Salicaceae, Sapindaceae, Rutaceae and Tamaricaceae

(Freina & Witt 1987). In Western Andalusia, it infests sev-

eral perennial plants of ornamental and economic inter-

est such as the Blueberry (Vaccinium sp., Ericaceae; Cal-

vo & Molina, 2004). It has been considered a local pest

for lime and grapefruit and its larvae may also have an

unwanted effect in nurseries when they feed on young

growing plants (Molina 1998).

The caterpillar of S. panda is known for its urticating

properties. The urticating apparatus has not been studied

in detail so far; Calvo & Molina (2008) simply mention

that urticating retractable organs develop beginning from

the second instar and appear as mere cuticle differentia-

tions in the first instar. In the present study, details of the

morphological structures responsible for the urticating

properties are provided for the first time.

Received: 20.11.2011

Accepted: 10.04.2012

MATERIALS AND METHODS

Larvae of Streblote panda were captured at the Atlantic

coast of Morocco at Essaouira in August 2005 and Oua-

lidia in November 2008 on the foliage of Retama monos-

perma (Linné) Boissieu (Fabaceae). Their larval develop-

ment comprises 5—8 instars (Calvo & Molina 2005). On-

ly the last instar larvae have been collected and studied.

For observation with scanning electron microscopy

(SEM), the mesonotum and metanotum were dissected and

isolated, dehydrated in absolute ethanol, mounted on spec-

imen holders and coated with a thin layer of gold and pal-

ladium in a Jeol JFC-1100 sputter coater. Preparations

were examined in a Jeol JSM-6400 SEM at 7 kV.

RESULTS

The urticating retractable apparatus is composed of two

sets of hairs located on the dorsal surface of the mesotho-

racic and metathoracic segments (Fig. 1). In each segment,

more than a thousand of these hairs are inserted in the in-

ternal walls of an integumentary fold composing a deep,

crescent-shaped crevice. At rest, the latter is retracted and

reveals externally only a transverse line whose edges are

lined with a few hairs. When active, the pocket opens, re-

vealing and spreading all the hairs it contains (Fig. 1). A

single type of urticating hair has been observed. It is spear-

shaped, 0.65—1.10 mm long (Fig. 5), with a slender base

(Fig. 6) and pointed distally (Fig. 7). It is delicately stri-

Corresponding editor: D. Stiining

130 Michel Joél Faucheux

Fig. 1. Larva of Streblote panda, last instar. Left: at rest, right: disturbed, showing the two urticating crevices.

ated longitudinally from the base onwards over 14/15" of

its length (Fig. 7). The distal part (1/15 ) comprises an area

with annular ligaments (Figs. 7, 8), tapering to the point-

ed and perforated tip (Figs. 9, 10). The estimated number

of pores per hair is about 400; their diameter reaches 0.3

um. The hairs are inserted in raised cupolae, from which

they may be detached (Fig. 11).

Different types of integumentary structures are associat-

ed with the urticating apparatus:

— Some thirty white scales, 0.30—0.35 mm long, are lined

up along the posterior edge of each urticating pocket

(Fig. 12), whereas dark scales are spread over the rest

Bonn zoological Bulletin 61 (1): 129-134

of the segment (Figs. 13, 14). The white scales, which

have the form of a strongly elongated tennis racket

(some visible in fig. 3), are covered with longitudinal

ribs made up of overlapping elements resembling roof

tiles (Fig. 15). This structure is analogous to that of

scales found on butterflies’ wings.

Five—six nonporous sensilla chaetica, 0.90—1.15 mm

long, are found on the pair of white D1 verrucae in the

anterior area of the meso- and metathorax (Figs. 16, 17).

Other sensilla chaetica, 0.50 mm long, are found among

the urticating hairs (Fig. 18).

Sensilla filiformia, 1.5—2.0 mm long, are found in lat-

eral tufts on the meso- and metathorax (Fig. 16).

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The urticating apparatus in the larva of the Lappet Moth, Streblote panda Hubner, 1820 131

Figs 2-11. Urticating apparatus of S. panda. 2. Urticating crevice on the mesothorax, showing urticating hairs (arrow). 3. Setal

tubercle (T), urticating hairs (arrow) and scales (S). 4. Group of urticating hairs. 5-11. Urticating hairs. 5. Entire hair (arrow). 6.

Base of hair. 7. Tips of hairs. 8. Tip of hair with ringed and perforated area. 9. Detail of the perforated area. 10. Detail of distal

pores. 11. Cupolae of urticating hairs.

Bonn zoological Bulletin 61 (1): 129-134 OZFMK

132 Michel Joél Faucheux

Figs 12-18. Scales, tubercles, sensilla and urticating hairs of S. panda. 12. Scales (arrow) among urticating hairs. 13. Isolated

scales. 14. Filiform scales. 15. Detail of scales. 16. Dorsal surface of the metathoracic segment showing two setal tubercles (T),

urticating hairs (arrows) and filiform sensilla (F). 17. Setal tubercle with sensilla chaetica (arrows). 18. Sensillum chaeticum (ar-

row) among the urticating hairs.

DISCUSSION

Structure. The large number of urticating hairs is char-

acteristic of urticating apparatuses (Bourgogne 1951). Un-

like ordinary long hairs, the urticating hairs of lepidopter-

an larvae are typically not arranged at random and do not

cover the caterpillars’ bodies in a uniform manner. They

are arranged in very dense groups on well defined surfaces

known as ‘mirrors’ that are integumentary folds (Sellier

et al. 1975) or other specific surface areas. The first lar-

val instars lack urticating hairs. In the 5" larval instar of

the ‘brown-tail moth’ Euproctis chrysorrhoea (Linnaeus)

(Lymantriinae), the “pine procession moth’ Thaumetopoea

pityocampa (Denis & Schiffermiiller), and the ‘oak pro-

cession moth’ Thaumetopoea processionea (Linnaeus)

Bonn zoological Bulletin 61 (1): 129-134

(Thaumetopoeinae), these mirrors are located on abdom-

inal segments I-VII whereas in the 3" instar of the same

species, they appear on different segments (Sellier et al.

1975). This difference in location of defense-related se-

tae among instars is rather unusual, given that in most

groups, as in S. panda, the position is constant heoteal

out the larval development. A fter they appear in the 2”

star (according to Calvo & Molina 2008), they remain on

the same thoracic segments, a situation known from many

other species of Lasiocampidae (e.g., in the genera Den-

drolimus and Gastropacha). Other lasiocampid moths,

such as Macrothylacia rubi, Lasiocampa quercus and oth-

ers, are known to possess urticating hairs on the abdom-

inal segments (Carter & Hargreaves 1988).

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The urticating apparatus in the larva of the Lappet Moth, Streblote panda Hiibner, 1820 133

A single type of urticating hairs is known in S. panda.

This is also the case in most other species of different fam-

ilies (Gilmer 1923, 1925; Faucheux 2007). However, there

are also species that have several different types of hairs,

e.g. three types in Latoia thamia Rungs (Limacodidae)

(Faucheux 2000). Among the different forms of urticat-

ing hairs, the most frequent in lepidopteran caterpillars is

a harpoon-shaped hair with pointed spikes directed to-

wards its distal end and arranged in three or four longi-

tudinal rows, e.g. as in TZ pitvocampa, T. processionea, E.

chrysorrhoea (Sellier et al. 1975), Lithosia quadra Lin-

naeus (Faucheux 2007), and in one type of L. thamia (Fau-

cheux 2000). These hairs, though hollow, are not perfo-

rated at any extremity and therefore are unable to inject

the toxic substance (Sellier et al. 1975). Instead, they must

be broken, for example in the skin, in order to release their

urticating substance (Ducombs et al. 1979).

In another type of hair, the nucleus of the poison-gland

cells is located at the base of the hair and the cells devel-

op inside the hair. During contact, the urticating substance

is released as the acute, sharp end of the hair breaks. This

type has been described in the “urticating thorns” of La-

toia (= Parasa) hilarata Staud. (Mills 1925) and in the

“poison apparatus” of Latoia (= Parasa) latistriga Walk-

er (Gilmer 1925). Gilmer (1923) distinguishes two types

of poisonous apparatuses of which the one he considers

more primitive consists of simple hairs that are shorter and

stiffer than others; each hair is connected through its prox-

imal end to a poison gland and opens in a distal pore (e.g.,

Orgyia leucostigma (J. E. Smith)). The structure of the ur-

ticating hairs of S. panda may therefore be close to this

primitive type, as the irritating substance flows through

the numerous pores located on the distal end of the hair.

Function of the mirrors. The mechanism for opening and

closing the ‘mirrors’ (or crevices) that contain the urticat-

ing hairs was described for 7. pitvocampa by Demolin

(1963). In this species, the implantation zone of the hairs

is delimited by a semi-rigid frame made up of chitinous

rods and small pads forming hinges. This arrangement can

open or close the insertion zones of the hairs that are nor-

mally hidden. When the caterpillars feel threatened, this

mechanism opens the fold and the urticating hairs become

entirely exposed. In S. panda, the opening mechanism has

not been studied in detail, but may function in a similar

way.

Urticating function. The urticating, harpoon-shaped hairs

of the ‘brown-tail moth’, the ‘pine procession moth’, and

the ‘oak procession moth’ can penetrate skin and eyes, and

cause irritating cutaneous eruptions or contact dermatitis

and ocular lesions (Blair 1979, Ducombs et al. 1979).

Their urticating properties last for a very long time, some-

times up to several months after their ejection (Ducombs

et al. 1979). In these species, the combination of a me-

Bonn zoological Bulletin 61 (1): 129-134

chanical element (penetration of the skin and breaking of

the hairs caused by scratching) and a chemical element

(discharge of a toxic substance derived from dermal

glands) account for the pathological symptoms. The ac-

tion of the urticating hairs is ascribable, at least in part,

to the secretion of histamine that takes place on the skin

as a result of the penetration of the poison contained in

the hairs (Leclercq 1977).

In S. panda, urticating hairs are released in large num-

bers when the caterpillar is handled. In contact with the

skin, they stick to it by means of their distal circular lig-

aments, but the urticating effect seems to be low. The nu-

merous pores at the distal ends of the hairs indicate that

a liquid is being released, but direct evidence is still miss-

ing. Previous studies do not mention strong irritations or

even dermatitis caused by the caterpillars of the lappet

moth.

Protective function. The aposematic red and dark brown

of the two urticating crevices strongly contrast with the

overall grey colour of the larva. Unlike other urticating

caterpillars, those of S. panda undoubtedly use their ur-

ticating apparatus mainly as a means of warning against

their predators. To my knowledge, no study has tested the

efficacy of this defence mechanism against natural ene-

mies. In the environment in which I observed the Moroc-

can caterpillars (colonies of Retama sp.), they come in fre-

quent contact with likely predators such as the common

chameleon, Chamaeleo chamaeleon Linnaeus and the

stripeless treefrog, Hyla meridionalis Boettger, which may

hesitate to prey on the caterpillars. Similarly, children and

adults are unwilling to touch a caterpillar that manifests

its anxiety by opening and closing its hair slits.

Acknowledgements. It is a pleasure to acknowledge Nicolas

Stephant and Stéphane Grolleau (Centre of SEM, University of

Nantes, France), and Catherine Cerclé for their technical assis-

tance as well as Vittorio Ballardini for help with the translation.

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sectes. Masson, Paris, New-York

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Molina JM (1998) Lepidopteros asociados al cultivo del aran-

dano en Andalucia Occidental. Boletin de Sanidad Vegetal y

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roc. Inventaire faunistique et observations écologiques, Tome

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cantes: biologie, importance économique et médicale. Bulle-

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France 73: 29-41

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Bonn zoological Bulletin 61 (1): 135-139

July 2012

Scotopteryx kuznetzovi (Wardikian, 1957)

(Lepidoptera, Geometridae, Larentiinae),

a new species for the fauna of Iran and Turkey

Hossein Rajaei Sh.’ & Dieter Stiining

Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

* Corresponding author: rajaei@daad-alumni.de; eagle4261@yahoo.com

Abstract. Scotopteryx kuznetzovi (Wardikian, 1957), a rare and habitually remarkable Larentiine species described from

Armenia, is recorded for the first time from Iran and Turkey. Males from both countries are illustrated, the females be-

ing still unknown. A redescription is given and diagnostic characters are presented. External features and genitalia are

mainly compared with those of S. vicinaria hyrcanaria (Staudinger, 1892), described from northern Iran, and other taxa

of the vicinaria species group which all have genitalia very similar to S. kuznetzovi. The taxonomic position of the lat-

ter is briefly discussed.

Key words. Scotopteryx kuznetzovi, redescription, new records, Iran, Turkey.

INTRODUCTION

Scotopteryx Hiibner, 1825, a Larentiine genus belonging

to the tribe Xanthorhoini Pierce, 1914, contains more than

70 species which are distributed in the Palaearctic region

from Europe to Far East Asia, in South Africa and South

America (Parsons et al., 1999). Neither in the latter pub-

lication nor in the updated list of the Geometridae of the

World (Scoble & Hausmann, 2007) Scotopteryx kuznet-

zovi (Wardikian, 1957) was mentioned. Until now, no fur-

ther records of this species, which was described on a sin-

gle male from Armenia, have been published. Here we

record the data of five additional males found in the col-

lections of the Zoologisches Forschungsmuseum Alexan-

der Koenig, Bonn (ZFMK, collection Peter Kuhna) and

Hayk Mirzayans Insect Museum (HMIM) in the Iranian

Research Institute of Plant Protection, Tehran, recently. In

addition, we provide a redescription of this almost un-

known species and compare it with the probably related

taxa of the S. vicinaria-group.

TAXONOMY

Redescription of Scotopteryx kuznetzovi (Wardikian)

Ortholitha kuznetzovi Wardikian, 1957: 281.

Type Material. Holotype <, (collection of the Zoological

Institute, Academy of Science of Armenia).

Type locality. Yerevan (Armenia).

Scotopteryx kusnetzovi: Wardikian, 1985: 27, 52 (incor-

rect subsequent spelling).

Scotopteryx kuznetsovi: Viidalepp, 1996: 13 (incorrect sub-

sequent spelling).

Received: 12.03.2012

Accepted: 31.05.2012

Material examined. 2 ¢: Basmendj [NW Iran, SE

Tabriz], 15.10. [19]74, [leg.] Damanabi, gen. preps 1063

& 1064/2010 H. Rajaei, Iran; coll. HMIM. 2 2: Ost Tiir-

kei, Van, 2700 m, Guizeldere Pass, 28.9.1981, leg. P. Kuh-

na, gen. prep. 3074, P. Kuhna; | @: Turkey, Prov. Bitlis,

Van GOlii, 19 km E of Ahlat, 1700 m, 42°34’ E, 38°46’

N, 18.10.1993, leg. Gy. Fabian, B. Herezig, Gy Laszlo and

K. Szeoke, coll. ZFMK.

Morphology of adult males (Figs 1, 2). Wingspan 29-30

mm. Antenna bipectinate from base to tip, except 2—3 dis-

tal segments, rami moderately long, black, dorsally un-

scaled, arising ventrally from the proximal end of the fla-

gellum segments, the latter scaled white dorsally. Head,

thorax and abdomen covered with mixed greyish-white

and greyish-brown scales. Frons broad, roundly protrud-

ing, lower part smoothly covered with very small, grey-

ish-brown scales, upper margin a broad band of large,

white scales, vertex with large white and brown scales

mixed. Chaetosemata transversally extended. Palpi short

and narrow, acute at tip, just reaching beyond the

clypeus. Haustellum almost completely reduced. Tibia of

hindlegs not dilated, index of spurs 0-2-4. Forewings elon-

gated, length 16-18 mm, apex and tornus rounded. Ground

colour yellowish-brown, surface shining, basal area and

medial band edged with dark brown internally and white

externally. Basal line moderately (in Turkish specimens)

to strongly (in Armenian and Iranian specimens) indent-

ed. Antemedial line roundly curved in the middle, with two

acute incisions anteriorly and posteriorly, the latter en-

larged and completely dividing the medial band in the

Corresponding editor: F. Herder

136 Hossein Rajaei Sh. & Dieter Stiining

3-a

3-b

Figs 1-3. Adults (males) of Scotopteryx. 1-2. Scotopteryx kuznetzovi: 1. Giizeldere Pass, E. Turkey; 2. Basmendj, Iran. 3. Sco-

topteryx vicinaria hyrcanaria, syntype, “Schakuh” [Semnan Prov., NE Iran]; a: dorsal view, b: ventral view. Scale bar: 1 cm.

holotype and in specimens from Iran. Postmedial line

wavy, roundly curved outwards in the middle, also with

two deep incisions at either side. Submarginal line white,

rather smooth in Turkish specimens, wavy in the holotype

Bonn zoological Bulletin 61 (1): 135-139

and the Iranian material. Apical whitish triangular patch

distinct in the holotype, but weakly defined in the other

specimens. Fringe consisting of a row of shorter and dark-

er basal scales and a row of longer and lighter terminal

©ZFMK

Scotopteryx kuznetzovi (Wardikian, 1957), a new species for the fauna of Iran and Turkey 137

Figs 4-5. Male genitalia. 4. S. kuznetzovi (NW. Iran); 5. S. vicinaria hyrcanaria (Schahkuh); a. genital armature, b. aedeagus.

Abbreviations: unc. Uncus; tr. Transtilla; cos. Costa; v. Valva; an.lob. Anellus lobe. Scale bar: 1 mm.

scales, both with distinctly darker areas near the ends of

veins. Discal dots black, narrow streak-like or separated

into dots in the forewings, absent in the hindwings. The

latter oval, elongated, whitish-grey, with a darker grey me-

Bonn zoological Bulletin 61 (1): 135-139

dial band. Fringe as in forewings. Underside generally

paler, but basal area up to postmedial line darker than the

rest, patterns of upper side only partly visible. Underside

of hindwing loosely scattered with single dark brown

OZFMK

138 Hossein Rajaei Sh. & Dieter Stiining

Cyprus :

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lime

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Fig. 6. Distribution map of Scotopteryx kuznetzovi (Wardikian, 1957).

scales. Abdomen long, narrow, light grey, with white pos-

terior margins of the segments. Sternites elongate, ST7 and

ST8 very small, no coremata developed.

Male genitalia (Fig. 4). Valve broad at base, sacculus scle-

rotized, without terminal processes; costa band-like,

curved, longer than the remaining part of the valva and

distally separated from it, with a rounded tip. Sacculus and

valve lamina setose. Transtilla a curved, continous band.

Uncus with a broad base and a narrow, acute tip, curved

ventrad. Anellus lobes elongate, conical, distally round-

ed and setose. Aedeagus only a little longer than valve,

slightly bent, distal one third covered with the densely

spined part of manica (if not removed), vesica with four

broad-based cornut.

Female unknown.

Diagnosis. S. kuznetzovi is easily distinguished from oth-

er members of the genus Scotopteryx by its peculiar ap-

pearance and by a number of distinct characters (here com-

pared mainly to members of the S. vicinaria-group, char-

acters in round brackets): elongated forewings with glossy

surface, rounded apex and tornus in kuznetzovi (forewings

less elongated, not glossy, apex less rounded, angled or

even slightly falcate; fig. 3); hindwing elongated-oval,

Bonn zoological Bulletin 61 (1): 135-139

much lighter than forewings and — except a medial band

— without pattern (hindwings as dark as forewings and

with much more pattern elements); antemedial and post-

medial lines strongly waved and incurved, with deep in-

cisions, sometimes medial band interrupted near hind mar-

gin (transverse lines smooth or — in S. subvicinaria —

slightly waved, not interrupted; fig. 3); transverse lines

highlighted with black inside and white outside, a white

submarginals line present (transverse lines not highlight-

ed in this way, submarginal line unconspicuous); anten-

nae bipectinate from base to apex, except distal 2-3 seg-

ments (antennae bipectinate and agreeing in all details, but

unpectinated distal portion distinctly longer); frons very

broad, palps short and acute at tip, haustellum almost com-

pletely reduced (frons less broad, palps longer, broader and

with rounded tip, haustellum well developed); abdomen

without coremata, 7 segment with a small sternite (ab-

domen with a pair of coremata arising laterally from 7

segment, tergite and sternite of 7 segment reduced, mem-

branous); aedeagus small, only slightly longer than

valve, with a few small cornuti on vesica (aedeagus very

large, at least two times the length of the valve, cornuti

large and numerous; figs 4, 5); tip of uncus short, anellus

lobes long (tip of uncus long, anellus lobes short, round-

ed; figs 4, 5).

©ZFMK

Scotopteryx kuznetzovi (Wardikian, 1957), a new species for the fauna of Iran and Turkey 139

Bionomy. S. Auznetzovi seems to be an autumn-flying

moth (end of September to end of October), based on the

few specimens collected (altitude 1700-2700 m). Food-

plant and larval stages are unknown.

Distribution. Armenia, NW Iran and E Turkey (fig. 6).

Taxonomic note. Habitually and in a number of morpho-

logical characters S. kuznetzovi is very different to and

seems to be widely separated from other members of the

genus Scotopteryx. Wardikian (1957) compared her new

species with S. bipunctaria (Denis & Schiffermiiller, 1775)

which, however, has strongly different genitalia structures

and belongs to another species-group inside Scotopteryx.

We found that the male genitalia of kuznetzovi agree well

and in detail with those of different taxa of the S. vicinar-

ia — group (Figs 4, 5). The taxonomy of the latter (com-

prising S. vicinaria, S. subvicinaria and their subspecies

brunnescens Prout, hyrcanaria Staudinger, libanaria

Prout and probably S. perplexaria Staudinger) is still not

fully understood, but is being revised in the course of fur-

ther studies on Iranian Larentiinae by the senior author.

Male genitalia capsules in all these taxa are extremely sim-

ilar, specific differences can probably be found only in the

number, size and arrangement of cornuti on the vesica, but

also these characters seem to be variable. Molecular stud-

ies have been started, but the number of barcoded speci-

mens is still too small and the results are not yet satisfac-

tory. Fresh specimens of kuznetzovi are urgently needed

to decide upon its position within the genus.

Scotopteryx kuznecovi Herbulot, 1996, mentioned in

Parsons et al. (1999: 871) in contrast to S$. kuznetzovi

Wardikian, is externally similar to S. chenopodiata Lin-

naeus, 1758 and related taxa. Choi (2002) treated it, as well

as S. golovushkini Kostjuk, 1991 (Transbaical region) as

a junior synonym of S. acutangulata Inoue, 1941, de-

scribed from Korea. Despite an identical origin and pro-

nunciation of Auznetzovi Wardikian and kuznecovi Herbu-

lot, the differences in spelling do not explicitly fall under

Bonn zoological Bulletin 61 (1): 135-139

the provisions of the Code (ICZN, 1999, articles

58.1.—58.15), of names deemed to be identical. So they

are here not treated as homonyms.

Acknowledgements. We thank Dr. Helen Alipanah (Tehran) for

loan of material from HMIM, and Dr. Bernd Miiller (Berlin) for

helping us in many ways. Sincere thanks of the senior author

are also due to Professor Dr. J. W. Wagele (ZFMK, Bonn) for

supporting his Ph.D. project and to the DAAD (Deutscher

Akademischer Austauschdienst) for financial help. This paper

is a part of the Ph.D. thesis of Hossein Rajaei at the University

of Bonn, Germany.

REFERENCES

Choi S-W (2002) Taxonomic Review of the Korean Xanthorhoi-

ni (Geometridae: Lepidoptera). Insecta Koreana 19: 213—231

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Wissenschaften der Armenischen SSR], Yerevan, 136 pp. (in

Russian)

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Bonn zoological Bulletin

EDITOR-IN-CHIEF

Fabian Herder, Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK), Ichthyology Section, Adenauerallee 160,

53113 Bonn, Germany,

tel. +49 228-9122-255, fax: +49 228-9122-212;

f-herder.zfmk(@uni-bonn.de

MANAGING EDITOR SUPPLEMENT SERIES

Thomas Wesener, ZFMK, tel. +49 228-9122-425,

fax: +49 228-9122-212; twesener@uni-bonn.de

EDITORIAL BOARD

Dirk Ahrens, Insects: Coleoptera, ZFMK,

tel. +49 2289122286, fax: +49 228—-9122—332:;

d.ahrens.zfmk@uni-bonn.de

Wolfgang Béhme, Amphibians and Reptiles, ZFMK,

tel. +49 228-9122—250, fax: +49 228-9122-212;

w.boehme.zfmk(@uni-bonn.de

Netta Dorchin, Insects: Diptera, Department of Zoology, Tel

Aviv University, Tel Aviv 69978, Israel, tel. +972-3-

6409808, fax: +972-3-6409403; ndorchin@post.tau.ac.il

Renate van den Elzen, Birds, ZFMK,

tel. +49 228-9122-—231, fax: +49 2289122212;

r.elzen.zfmk@uni-bonn.de

Bernhard Huber, Invertebrates except Insects, ZFMK,

tel. +49 228-9122-294, fax: +49 228—-9122—212;

b.huber.zfmk(@un1-bonn.de

Rainer Hutterer, Mammals, ZFMK,

tel. +49 2289122261, fax: +49 228-9122-212;

r.hutterer.zfmk@uni-bonn.de

Gustav Peters, Mammals, ZFMK,

tel. +49 2289122262, fax: +49 2289122212;

g.peters.zfmk@uni-bonn.de

Bradley Sinclair, Canadian National Collection of Insects,

Ottawa Plant Laboratory — Entomology, CFIA, K.W. Neat-

by Bldg., C.E.F., 960 Carling Ave., Ottawa, ON, Canada

K1A 0C6, tel. + 1 613-759-1787, fax: + 1 613-759-1927;

bradley.sinclair@inspection.ge.ca

Dieter Stiining, Insects except Coleoptera and Diptera,

ZFMK, tel. +49 228-9122-220, fax: +49 228-9122-212;

d.stuening.zfmk(@uni-bonn.de

Philipp Wagner, Villanova University, Department of Biolo-

gy, 800 Lancaster Avenue, Villanova, PA 19085-1699,

USA; philipp.wagner.zfmk@uni-bonn.de

ADVISORY BOARD

Theo C. M. Bakker, Rheinische Friedrich-Wilhelms-Univer-

sitat, Institut fiir Evolutionsbiologie & Okologie, 53113

Bonn, Germany, tel. +49 228—73—5130, fax: +49 228—73—

2321; t.bakker@uni-bonn.de

Aaron M. Bauer, Villanova University, Department of Biolo-

gy, 800 Lancaster Avenue, Villanova, PA 19085-1699,

USA, tel. +1 610-519-4857, fax: +1 610-519-7863;

aaron.bauer@villanova.edu

Wieslaw Bogdanowicz, Museum and Institute of Zoology,

Polish Academy of Sciences, Wilcza 64, 00-679 Warszawa,

Poland, tel. +48 22—628—7304, fax: +48 22-629-6302;

wieslawb@muiz.waw.pl

Matthias Glaubrecht, Museum fiir Naturkunde Berlin, Leib-

niz-Institut fiir Evolutions- und Biodiversitatsforschung an

der Humboldt-Universitat zu Berlin, Invalidenstrasse 43,

10115 Berlin, Germany, tel. +49 30—2093-8504/ 8400,

fax: +49 030-2093-8565; matthias.glaubrecht@mfn-

berlin.de

Jeremy D. Holloway, The Natural History Museum, Depart-

ment of Entomology, Cromwell Road, London, SW7 5BD,

U.K.; j.holloway@nhm.ac.uk

Tan Heok Hui, Raffles Museum of Biodiversity Research,

National University of Singapore, Department of Biological

Sciences, 6 Science Drive 2, #03-01, Singapore 117546, tel.

+65-6516 1662, heokhui@nus.edu.sg

Boris Krystufek, Slovenian Museum of Natural History, P.

O. Box 290, Ljubljana, Slovenia; boris.krystufek@zrs.upr.si

Wolfgang Schawaller, Staatliches Museum fiir Naturkunde,

Rosenstein 1, 70191 Stuttgart, Germany,

tel. +49 711-8936-221, fax: +49 711-8936—100;

schawaller.smns@naturkundemuseum-bw.de

Ulrich K. Schliewen, Department of Ichthyology, Bavarian

State Collection of Zoology, Mtinchhausenstr. 21, 81247

Miinchen, Germany, tel. + 49 89-8107—110;

schliewen@zsm.mwn.de

Michael Schmitt, Ernst-Moritz-Amdt-Universitat, Allge-

meine & Systematische Zoologie, Anklamer Str. 20, 17489

Greifswald, Germany, tel. +49 3834—-86-4242, fax: +49

3834864098; michael.schmitt@uni-greifswald.de

W. David Sissom, Dept. of Life, Earth and Environmental

Sciences, W. Texas A. & M. University, WITAMU Box

60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578,

fax: +1 806-651-2928; dsissom@mail.wtamu.edu

Miguel Vences, Technical University of Braunschweig, Zoo-

logical Institute, Spielmannstr. 8, 38106 Braunschweig,

Germany, tel. + 49 531-391-3237, fax: + 49 31-391-8198;

m.vences@tu-bs.de

Erich Weber, Eberhard-Karls-Universitat, Zoologische

Schausammlung, Sigwartstr. 3, 72076 Tiibingen,

tel. +49 70712972616, fax +49 7071—295170;

erich.weber@uni-tuebingen.de

felted veld wt? D

LIBRARIES

Contents WAMU I

Sonnenberg, Rainer & Jouke R. Van der Zee: 3

Aphyosemion pseudoelegans (Cyprinodontiformes: Nothobranchiidae), a new killifish species

from the Cuvette centrale in the Congo Basin (Democratic Republic of Congo)

Sonnenberg, Rainer & Eckhard Busch: 13

Description of Scriptaphyosemion wieseae (Cyprinodontiformes: Nothobranchiidae),

a new species from northern Sierra Leone

Koerber, Stefan: 29

Mercediella nom. nov., a replacement name for Camposichthys Figueiredo & Silva Santos, 1991

(Pisces: Pycnodontiformes)

Rosa, Goncalo M., Paolo Eusebio Bergo, Angelica Crottini & Franco Andreone: 31

Report of the life colouration of the enigmatic burrowing skink

Voeltzkowia rubrocaudata (Grandidier, 1869) from southwestern Madagascar

Wagner, Philipp, Dennis Rodder & Thomas M. Wilms: 35

New data on the morphology and natural history of Tetradactylus ellenbergeri (Angel, 1922)

(Sauria: Gerrhosauridae) and Trachylepis ivensii (Bocage, 1879) (Sauria: Scincidae)

in northeastern Zambia

Nadein, Konstantin & Chi-Feng Lee: 41

New data about some Alticini from Taiwan with descriptions of two new species

(Coleoptera: Chrysomelidae)

Assing, Volker: 49

A revision of East Palaearctic Lobrathium (Coleoptera: Staphylinidea: Paederinae)

Faucheux, Michel Joél: 129

The urticating apparatus in the larva of the Lappet Moth, Streblote pence Hubner, 1820

(Lepidoptera: Lasiocampidae)

Rajaei Sh., Hossein & Dieter Stlining: 135

Scotopteryx kuznetzovi (Wardikian, 1957) (Lepidoptera: Geometridae: Larentiinae),

a new species for the fauna of Iran and Turkey

Cover illustration:

Bundesministerium ,,. . : F :

7X | fiir Bildung Ministerium fir Innovation,

und Forschung Wissenschaft und Forschung

des Landes Nordrhein-Westfalen

Leibniz- eaten