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Bonn zoological Bulletin

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Koenig (ZFMK), Ichthyology Section, Adenauerallee 160,

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

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

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K1A0C6, tel. + 1 613-759-1787;

bradley.sinclair@inspection.gc.ca

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

Poland, tel. +48 22-628-7304, fax: +48 22-629-6302;

wieslawb@miiz.waw.pl

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niz-Institut fur 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;

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

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

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National University of Singapore, Department of Biological

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

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

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State Collection of Zoology, Miinchhausenstr. 21, 81247

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meine & Systematische Zoologie, Anklamer Str. 20, 17489

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erich.weber@uni-tuebingen.de

I

, 3 7cZ 70*

fj-tt

FORSCHUNGS

museum

KOENIG

Bonn

zoological

Bulletin

formerly: Bonner zoologische Beitrage

Index 2014

Volume 63

Issue 1-2

KOENIG

Bonn

zoological

Bulletin

Volume 63

Issue 1

2014

■v^L KOENIG

Bonn Volume 63

zoological lss2u0e,4

Bulletin

Editor-in-Chief

Fabian Herder

Editorial Board

Dirk Ahrens

Wolfgang Bohme

Jan Decher

Netta Dorchin

Bernhard A. Huber

Rainer Hutterer

Ximo Mengual

Gustav Peters

Ralph Peters

Bradley Sinclair

Dieter Stuning

Till Topfer

Philipp Wagner

An open access journal of organismal zoology, published by

Zoologisches Forschungsmuseum Alexander Koenig - Leibniz-lnstitut fur Biodiversitat derTiere, Bonn

Contents Volume 63

Grebennikov, Vasily V.: 123-147

Morimotodes, a new genus for two minute wingless litter species from

southwest China and Taiwan with an illustrated overview of Molityna and Plithina genera (Coleoptera: Curculionidae:

Molityni)

Vol. 63 Issue 1 : June 2014

Vol. 63 Issue 2: December 2014

Bonn zoological Bulletin 63 (2): 123-147

December 2014

Morimotodes , a new genus for two minute wingless litter species

from southwest China and Taiwan with an illustrated overview of

Molytina and Plinthina genera (Coleoptera: Curculionidae: Molytini)

Vasily V. Grebennikov

Canadian Food Inspection Agency, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, ON K1A 0Y9, Canada;

E-mail: vasily. grebennikov@inspection.gc. ca

Abstract. The new weevil genus Morimotodes g. n. is described for two new species: M. ismene sp. n. (Yunnan and Sichuan)

and M. polynices sp. n. (Taiwan). Morimotodes ismene sp. n. is designated as the type species of Morimotodes g. n. Species

are small (1. 6-2.1 mm), wingless inhabitants of leaf litter in montane deciduous forests with characteristically globular

body and obliterate elytral striae and dorsal sculpture. Analysis of DN A barcodes and external similarity suggest that Mo-

rimotodes g. n. is a sister to the West Palaearctic genus Leiosoma Stephens, 1829 with 32 similarly small-bodied and lit-

ter-inhabiting species. Dated intraspecific phylogeographic analysis utilizing DNA barcodes of M. ismene sp. n. reveals

seven strongly supported geographical clades, four of which are allopatric and three others are parapatric on Mt. Emei

in Sichuan. Potentially allied genera belonging to two of the three Molytinini subtribes, Molytina and Plinthina, are il-

lustrated and briefly discussed. Type specimens of five exceptionally poorly known genera ( Carbonomassula Heller, 1908,

Clarkantlms Alonso-Zarazaga & Lyal, 1999, Gonotrachelus Champion, 1914, Vouauxia Hustache, 1920, Pterotomus Que-

denfeldt, 1888) are re-examined and the original labels illustrated.

Key words: Yunnan, Sichuan, Taiwan, COl, DNA barcoding, phylogeography

INTRODUCTION

In 2008, a first long series of conspicuous but unknown

beetles was discovered by sifting forest leaf litter on the

eastern slope of the Cang Shan Mountain Range in Yun-

nan (China). The peculiarly small, lustrous and rounded

body in combination with the rounded elytral shoulders

strongly resembled other litter-inhabiting wingless

Palaearctic Molytinae, i.e. Otibazo Morimoto, 1961 or

Leiosoma Stephens (Fig. 7). The subsequent four years of

intensive sampling in the same and nearby forests of

Southwest China revealed additional similarly shaped

specimens. A long series of another seemingly congener-

ic species was detected among unidentified specimens col-

lected two decades earlier in Taiwan by Ales Smetana (Fig.

6). The generic identity of these beetles, however, re-

mained elusive.

Since the original discovery in 2008, these beetles were

repeatedly compared to many others, mainly belonging to

the wingless members of Molytinae. Eventually it became

evident that these finds represented a previously undetect-

ed clade presumably deserving a formal status of a new

genus. This thought intensified the search for its possible

relatives. That pursuit was long inconclusive, since numer-

ous unrelated wingless Molytinae were infamous for their

convergent similarity linked to the loss of hind wings (Lyal

2014). DNA-based search for relatives of the new genus

was similarly inconclusive, since the dataset did not

Received: 07.08.2014

Accepted: 12.11.2014

contain many critical and important taxa. Moreover, due

limitations in funding, the DNA work was done with the

single marker of the DNA barcode (part of cytochrome

oxidase I, Hebert et al. 2003), which had significant lim-

itations in its applicability for detecting relatively old evo-

lutionary events (Grebennikov & Heiss 2014). Eventual-

ly, however, a representative of the genus Leiosoma be-

came available for DNA analysis and consistently clus-

tered with a clade representing the new genus. At that point

the project’s results were judged mature enough to be for-

mally presented, even though some shortcomings linked

to a shortage of the material and data remained (see Dis-

cussion).

Another coincidental event further triggering the com-

pletion of this project was the recent publication of the

long-awaited “Phytophaga” volume of the Handbook of

Zoology series, most specifically the chapter on the wee-

vil subfamily Molytinae (Lyal 2014). This subfamily has

long been and still is a phylogenetic challenge. In the

aforementioned chapter the tribes of Molytinae were sig-

nificantly reorganized, as compared to the earlier standard

works of reference (Alonso-Zarazaga & Lyal 1999;

Bouchard et al. 2011; Alonso-Zarazaga 2013). The most

noticeable novelty, however, was the incorporation of the

entire former “subfamily Cryptorhynchinae” into Molyti-

nae, the latter presently containing 37 tribes plus ca. 28

Corresponding editor: D. Ahrens

124

Vasily V. Grebennikov

incertae sedis genera (Lyal 2014: 530, 531). This decision

is in line with the pivotal work of Kuschel (1987) amal-

gamating a multitude of former “subfamilies” into a broad-

ly defined Molytinae (then excluding “Cryptorhynchi-

nae”). Both steps are seen as a much needed attempt to

make the formal classification more in line with the cur-

rent phylogenetic hypothesis (for example, see the remark-

able mosaic of both former “subfamilies” in McKenna et

al. 2009). Given an acute lack of the phylogenetic frame-

work for the subfamily and for the great majority of its

tribes, numerous further changes in Molytinae classifica-

tions will undoubtedly follow. To move forward, a prac-

tical approach was outlined (Lyal 2014: 549) as to “... pro-

pose initial hypotheses for monophyletic groups within the

Molytinae and then analyze them with both molecular and

morphological characters...” This paper presents an at-

tempt to answer this call for actions on Molytinae sub-

groups and to shed light on a small subsection of this

grand, if indeed monophyletic, radiation.

This work has three main goals. First, to name formal-

ly the newly discovered genus and its two species. Sec-

ond, to utilize available mtDNA sequences to test the

monophyly of the genus and to search for the possible sis-

ter group. Third, to discuss and delimit species boundaries

and to undertake a dated phylogeographical analysis.

Forth, to advance understanding of Molytinae genera cur-

rently attributed to the subtribes Molytina and Plinthina

by providing an annotated list of taxa, illustrating a rep-

resentative species for all but three genera and clarifying

the identity of five obscure genera by re-examination of

the type specimens of their type species. Overall, this pa-

per continues the recently initiated efforts to assess phy-

logeographical patterns detected from the low-dispersing

Hightless weevils in southwestern China and adjacent

lands and to clarify the taxonomy and diversity of wing-

less Old World Molytinae (Grebennikov 2014).

MATERIAL AND METHODS

Museum abbreviations, followed by the name of the cu-

rator:

BMNH Natural History Museum, London, UK

(M. Barclay);

CMN Canadian Museum of Nature, Ottawa, Canada

(R. Anderson, F. Genier);

CNC Canadian National Collection of Insects,

Arachnids and Nematodes, Ottawa,

Canada (P. Bouchard);

IZCAS Institute of Zoology, Chinese Academy of

Science, Beijing, P.R. China (R. Zhang);

MNHN Museum national d’Histoire naturelle, Paris,

France (H. Perrin);

MRAC Musee Royal de l’Afrique Centrale, Tervuren,

Belgium (M. De Meyer);

MTD Senckenberg Naturhistorische Sammlungen,

Dresden, Germany (K.-D. Klass, O. Jager);

NHMB Naturhistorisches Museum, Basel, Switzerland

(E. Sprecher);

NMW Naturhistorisches Museum, Wien,

Austria (H. Schillhammer);

ZIN Zoological Institute, Russian Academy of

Sciences, St. Petersburg,

Russia (B.A. Korotyaev).

This study is based on examination of a total of 97 dry-

mounted adult specimens of the new genus. Concepts of

the Curculionoidea family-group names and assignment

of weevil genera to tribes follow Bouchard et al. (201 1 )

and Alonso-Zarazaga & Lyal (1999), respectively, with

subsequent significant modifications by Lyal (2014). The

chronostratigraphic timing is that of Cohen et al. (2013)

with the Pliocene-Pleistocene boundary set at 2.588 MY.

Nomenclature of male genitalia follows that of Wanat

(2007). The length of the body was measured in dorsal as-

pect from the elytral apex to the front of the pronotum.

Geographical labels of the newly described species are cit-

ed verbatim in quotation marks. The hypothetical most re-

cent common ancestor of a monophyletic group referred

to in the discussion is abbreviated as MRCA. The distri-

bution map was generated using the online SinrpleMap-

pr tool (Shorthouse 2010). Species identification of the

two sequenced specimens of Leiosoma reitteri (Bedel) was

done based on the assumption that this is the only species

of the genus known from Georgia (Alonso-Zarazaga

2013). The type species are cited using their presently

valid names, which are not necesserely the same when the

type species were designated; the latter can be found in

Alonso-Zarazaga & Lyal ( 1999).

DNA dataset. All specimens submitted to DNA barcod-

ing have a unique identifier label with the code CNC-

COLVG0000XXXX; this format is shortened to #XXXX

when specimens are cited in the text. All details regard-

ing sequencing, primers, PCR protocols as well as the list

of specimen information including provenience and

voucher number can be seen by accessing the online

dataset at: dx.doi.org/10.5883/DS-MORIMO90. In a few

cases a specimen failed to amplify DNA from the first at-

tempt and was later resubmitted; in such cases a digit “1”

after a dot follows the original four digit number (like

#0251.1 on Fig. 7). The dataset used for the first DNA

analysis contained 90 sequences and 658 aligned positions.

The ingroup was formed by 46 newly generated se-

quences, each representing a specimen of the continental

new species of the new genus (GenBank accessions:

HM4 1 7727-32, HQ986799, HQ986876-8, HQ987105,

Bonn zoological Bulletin 63 (2): 123-147

©ZFMK

Morimotodes, a new weevil genus from China

125

HQ9871 34-5, HQ987 1 59, HQ987 1 79-8 1 , KJ87 1 620-32,

KJ87 1634-5, KJ87 1637-40, KJ87 1 642-5 1 ). All but one

specimen representing another new species collected in

Taiwan in 1992-1998 were submitted to DNA barcoding

but failed to amplify, likely because of either their age,

collecting methods or storage degraded DNA. No fresh

specimens of the target genus were recovered in Taiwan

during a month-long fieldwork in 2013. The majority of

the ingroup sequences were 658 basepairs (bp) long and

no one shorter than 407 bp. The outgroup consisted of 44

sequences and represented all Molytinae genera with avail-

able DNA barcode data (Fig. 7), while three sequences of

Graptus circassicus Solari, 1945 (Entiminae) were used

to root the topology in the first analysis (see below). All

outgroup sequences were of the full DNA barcode length

(658 bp). Three of the outgroup sequences representing

the genus Plinthus Germar, 1817 (KJ871633, KJ871636,

KJ871641 ) were newly generated. The remaining 41 out-

group sequences were taken from previous works

(HM386450, HM4 17677-9, HQ986888-9, HQ987002-3,

KJ427730-2, KJ427736-9, KJ427744, KJ427746,

KJ427748, KJ445683-4, J445687, KJ445690-1,

KJ445693-4, KJ445698-6, KJ445708-9, KJ445712-3,

KJ445715 in Grebennikov 2014; KJ841729, KJ841731 in

Grebennikov & Pham 2014). Specimen images (when

present), geographical data, primers, original chro-

matograms and other relevant data for all 90 sequenced

specimens can be found online in the publicly accessible

dataset “ Morimotodes ” on the Barcode of Life Database

portal (doi: dx.doi.org/10.5883/DS-MORIMO90).

DNA-based phylogenetic analyses. Two separate tree

search analyses were run. The first analysis aimed at test-

ing monophyly of the new genus represented by the type

species, search for its sister-group and to examine its in-

ternal relationships. The original DNA dataset containing

90 sequences was analysed using the Maximum Likeli-

hood (ML) approach with statistical branch support test-

ed by 1000 replications of bootstrapping in MEGA 5

(Tamura et al. 2013). The search for the optimal substi-

tution model for both analyses was done in MEGA 5 and

resulted in the same model (GTR+G+I). The second tree

search analysis was performed to date the branching

events in the continental species of the new genus and to

re-test the interrelationships hypothesis, as already sug-

gested in the first analysis. The original matrix of 90 se-

quences was reduced to 33 to include only the full-length

DNA barcodes (658 nt) of 3 1 ingroup specimens and two

specimens of the sister group (the genus Leiosoma , as sug-

gested in the first analysis). The second analysis was per-

formed in BEAST vl.8.0 (Drummond et al. 2012) utiliz-

ing the Bayesian inference (BI) approach with no a pri-

ori internal grouping, all default priors and options,

GRT+G+I nucleotide substitution model, strict linear mo-

lecular clock and nucleotide substitution rate of 0.0 1 8 ( Pa-

padopoulou et al. 2010). The “burn in’’ option was imple-

mented eliminating the first 2500 of the 10000 obtained

trees. Two resulting topologies, each representing either

ML or BEAST analysis, were visualized in FigTree vl.4

(Rambaut 2013) and presented by Figs 7 and 8, respec-

tively.

Use of morphological data. A formal morphology-based

phylogentic analysis of Molytinae genera to search for the

sister group of the new genus was beyond the scope of

the present paper. Even restricted to 8 1 genera of the tribe

of Molytini (80 mentioned in Lyal 2014, and the new one),

the task was judged as too daunting. An attempt was made

to illustrate a representative of all genera currently as-

signed to Molytina and Plinthina (13 and 17, respective-

ly; see Lyal 2014: 530, 53 1 ; images of three genera were

not obtainable). Five particularly poorly known genera

were elucidated in more details, with their type specimens

and original labels imaged. The third remaining Molyti-

ni subtribe, Flylobiina, historically considered as a tribe

of its own and incorporating 5 1 winged genera (Lyal 2014)

was not included in this treatment. This decision is part-

ly due to the large size of this tribe. The second reason

for exclusion is that until very recently, Hylobiina were

formally in a separate tribe and, therefore, not specifical-

ly targeted since 2008, when this project was in prepara-

tion.

Morimotodes g. n.

Type-species: Morimotodes ismene sp. n., by original des-

ignation.

Diagnosis. Adult specimens of Morimotodes g. n. can be

immediately recognized from other Curculionoidea known

both from the Eastern Palaearctic and the Oriental Region

by their habitus. They are small (about 2 mm) brown or

reddish beetles with lustrous dorsum, rounded body and

nearly obliterated elytral shoulders and striae. In the area

of their known distribution, namely in Yunnan, Sichuan

and Taiwan, as well as in the adjacent regions, adults of

Morimotodes externally resemble those of Otibazo Mo-

rimoto. The latter, however, has not yet been discovered

in China (although its presence is possible, since the first

continental species has been recently detected in north-

ern Vietnam, see Grebennikov & Pham, 2014). Adults of

Otibazo are immediately distinguishable from nearly all

weevils by their unique fields of short dense pubescence

forming characteristic whitish tufts on flagella, femora and

tibiae (Fig. 7), not found in Morimotodes. Field recogni-

tion of Morimotodes can be facilitated by stressing their

biological preferences. All known specimens have been

found in wet leaf litter of primary deciduous forests in the

altitudinal range of 1947^4072 m in southwestern China

and 1110-2550 m in Taiwan. From the most closely re-

Bonn zoological Bulletin 63 (2): 123-147

©ZFMK

126

Vasily V. Grebennikov

Fig. I. Morimotodes ismene g. & sp. n., holotype, male, #0928 from Gaoligong Mt. Range, Yunnan. A-D: habitus; E-G: aedea-

gus and tegment, dorsal (E), ventral (F), lateral; (G); H: stemites 8 and 9.

lated genus Leiosoma, adults of Morimotodes g. n. can be

easily distinguished by their brown or reddish colour

(Leiosoma adults are black) and, additionally, by a wide

gap in distribution ( Leiosoma is restricted to the West

Palaearctic and, therefore, is widely allopatric with Mo-

rimotodes g. n.).

Description. Small (1.6-2. 1 mm) and uniformly brown

or dark reddish, lustrous, wingless weevils. Dorsal surface

of pronotum and elytra without longitudinal ridges, cari-

nae or irregular sculpture; elytral striae weakly indicated

by individual rows of fine setiferous punctures surround-

ed by clearly restricted zone of darker pigmentation (Figs

1 A-C, 6A-C); setiferous punctures on pronotum shallow-

er, not arranged in longitudinal rows and lacking pigment-

ed surroundings (Figs 1 A-C, 6A-C). Length of dorsal se-

tae variable, from almost indistinguishable under 50x mag-

nification (Figs 6A-C, “polished” specimens) to relative-

ly long and exceeding in length longitudinal distance be-

tween setiferous punctures (Figs 4B, 5B, “hairy” speci-

mens). Integument clean, not covered by incrustation or

wax secretion. Elytra with shoulders entirely levelled, in

dorsal view strongly and evenly rounded, 1 .5-1 .8x as wide

as maximal prothoracic width. Body dorsal outline in lat-

eral view evenly and relatively strongly curved (Fig. 5);

elytra neither flattened dorsally, nor declivate apically.

Head capsule moderately visible from above (Fig. 4). Eyes

laterally, not reduced in size. Antennal attachments in api-

cal third of rostrum; funicle with seven antennomeres;

compact club consisting of three antennomeres. Pronotum

not constricted laterally in anterior quarter; widest at mid-

dle or at base; prosternum without rostral canal or ocular

Bonn zoological Bulletin 63 (2): 123-147

©ZFMK.

Morimotodes, a new weevil genus from China

127

Fig. 2. Morimotodes ismene g. & sp. n., paratype, female, #1020 from Gaoligong Mt. Range, Yunnan. A-D: habitus, E: genital

stemites 8 & 9, ventral view.

lobes; procoxae contiguous. All femora without tooth.

Hind wings absent. Sclerolepidia absent (Fig. 3E). Aedea-

gus weakly sclerotized, dorsally and ventrally membra-

nous and translucent; endophallus without slerotization

(Figs 1 E, F, 6E, F). Female hemistemites 9 each with sep-

arate and clearly distinguishable basal and apical pieces

(Fig. 2E; “coxite” and “stylus”, respectively).

Species composition and distribution. As currently

known, the genus Morimotodes g. n. comprises two

species, each relatively widely distributed in Southwest

China (Yunnan and Sichuan) or in Taiwan, respectively.

Biology. All specimens of Morimotodes were collected by

sifting deciduous leaf litter in montane primary forests

with subsequent specimen extraction using Winkler fun-

nels. These collecting circumstances were further corrob-

orated by the loss of hind wings, which strongly suggest

that Morimotodes species are habitat-dependant low-dis-

persing forest litter inhabitants critically dependant on this

habitat. Immature stages, host plants or any specific bio-

logical preferences remain unknown.

Etymology. Morimotodes , patronymic, derived by adding

the Latin suffix -odes to the family name of Katsura Mo-

rimoto (retired, formerly professor at Kyushu Elniversity,

Fukuoka, Japan), in recognition of his outstanding con-

tribution to the knowledge of weevils from the Asia-Pa-

cific Region. Gender masculine.

Morimotodes ismene sp. n.

Figs 1-5.

Diagnostic description. Holotype, male (Fig. 1). Gen-

Bank accession: HQ987135. Length: 1.85 mm. Dorsal out-

line of pronotum in lateral view evenly rounded; prono-

tum and elytra dorsally with fine punctures and white pi-

losity visible at magnification below 30x; each elytron

with single longitudinal groove adjacent and parallel to

edge. Acdeagus parallel-sided; sternite 9 not subdivided

and not wider than 1.5x width of apodeme; sternal

apodeme 9 with large paddle-shaped extension anterior-

ly. Intraspecific variation. Length: 1.60-2.0 mm. Body

outline in dorsal and lateral views variable (Figs 4, 5),

pronotum in basal half either parallel-sided (Fig. 4D) or

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Fig. 3. Morimotodes ismene g. & sp. n., unsexed, #0427 from Cang Shan Mt. Range, Yunnan. SEM images. A: head, left later-

al; B: head and pronotum, left lateral; C: head, frontal; D: rostrum, frontal; E: meso- and metathorax, left lateral; F: claws, later-

al; G: pronotum and head, dorsal; H: meso- and metathorax, right ventro-lateral; I: elytra, left lateral. Mst: metepistemum, mtv:

metaventrite, col-co3: pro-, meso- and metaxocae, respectively, vtl : ventrite 1 (=abdominal segment 3). Note absence of sclerole-

pidia on metepisternal suture separating metepistemum and metaventrite; the presence of this suture in apterous weevils is unusu-

al.

weakly broadened posterad (Fig. 4E); dorsal outline of ros-

trum-to-head junction even (Fig. 5B), evenly and weak-

ly depressed (Figs 1 A, 5A) or marked more abruptly (Fig.

5D). Dorsal body puncture and pilosity variable, from

mostly faint (Fig. 5D) to markedly developed (Fig. 5B).

Material examined. Holotype male (IZCAS): #0928,

“P.R. CHINA, Yunnan, E slope N Gaoligongshan,

N27°47’22. 1 E098°32’17.7, 24.V.2010, 3027m, sifting20,

V.Grebennikov”. Paratypes (CNC, IZCAS, MTD), 19 in

total; 4 exx, #0114-6, #0503: “P.R. CHINA, Yunnan, E

slope N Gaoligongshan, N27°46.212’ E098°36.221\

12.vi.2009, 2312m, sifting02, V.Grebennikov”; 5 exx,

#0144 and 4 not numbered: “P.R. CHINA, Yunnan, E

slope N Gaoligongshan, N27°46.782’ E098°33.087\

14. vi. 2009, 2956m, sifting04, V.Grebennikov”; 2 exx, not

numbered: “P.R. CHINA, Yunnan, E slope N Gaoligong-

shan, N27°46.665’ E098°33.32 1 ’, 15.vi.2009, 2982m, sift-

ing05, V.Grebennikov”; I ex, not numbered: “P.R. CHI-

NA, Yunnan, E slope N Gaoligongshan, N27°46.635’

E098°33.381’, 16.vi.2009, 2946m, sifting07, V.Greben-

nikov”; 2 exx, #0927 and 1 not numbered: same data as

holotype; 1 ex, #0977: 2010/21 “P.R. CHINA, Yunnan, E

slope N Gaoligongshan, N27°47’22.1 E098°32’17.7,

24.V.2010, 3027m, sifting21, V.Grebennikov”; 4 exx,

#1017-20: “P.R. CHIN A, Yunnan, E slope N Gaoligong-

shan, N27°59’01 .0 E098°32’56.9, 27.V.2010, 3018m, sift-

ing22, V.Grebennikov”.

Additional non-type specimens: Cang Shan Mt. Range

(15 in total, CNC): 14 exx, #0300-1, #0427, #0499-502,

7 not numbered: “P.R. CHINA, Yunnan, Cangshan,

N25°40.381’ E100°08.016\ 30.V.2008, 2600m, sifting,

V.Grebennikov”; 1 ex, #0867: “P.R. CHINA, Yunnan, E

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Morimotodes, a new weevil genus from China

129

Fig. 4. Morimotodes ismene g. & sp. n., unsexed, dorsal view. A: #2409 from Mount Gongga, Sichuan; B: #4483 from Mount

Haba, Yunnan; C: #6185 from Mount Emei, Sichuan; D: #1105 from Mount Emei, Sichuan; E: #0867 from Cang Shan Mt. Range,

Yunnan; F: #0251 from Mount Emei, Sichuan. Scale: 1 mm.

slope Cangshan at Dali, N25°40’15.5 E100°07’45.4,

18.V.2010, 2657m, sifting 1 7, V.Grebennikov”. Mount Ha-

ba (6 in total, CNC): 4 exx, #4481-4: “CHINA, Yunnan,

Haba Shan, N27°21’01“ El 00°05’44”, 21.vi.2012,

4072m, sift26, V. Grebennikov”; 2 exx, #4536-7: “CHI-

NA, Yunnan, Haba Shan, N27°21'01“ E100°05’44”,

28.vi.20 12, 4072m, sift35, V. Grebennikov”. Mount

Gongga (9 in total, CNC): 1 ex, #2409: “P.R. CHINA,

Sichuan, NE slope Gongga Shan, N29°48’I5”

E102°03’44”, 06.vi.201 1, 2765m, siftl 1, V.Grebennikov;

1 ex, #2456: “P.R. CHINA, Sichuan, NE slope Gongga

Shan, N29°50’50” EI02°02’28”, 09.vi.2011, 3170m,

sift 1 4, V.Grebennikov”; 3 exx, #2471-3: “P.R. CHINA,

Sichuan, NE slope Gongga Shan, N29°50’05”

E102°02’53”, 1 1 .vi.201 1, 3019m, siftl 5, V.Grebennikov”;

2 exx, #2559-60: “P.R. CHINA, Sichuan, NE slope Gong-

ga Shan, N29°49’33” E102°03’06”, 21. vi.201 1, 2977m,

sift24, V.Grebennikov”; 3 exx, #2562-3: “P.R. CHINA,

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Vasily V. Grebennikov

Fig. 5. Morimotodes ismene g. & sp. n., unsexed, left lateral view. A: #2409 from Mount Gongga, Sichuan; B: #4483 from Mount

Haba, Yunnan; C: #6185 from Mount Emei, Sichuan; D: #1 105 from Mount Emei, Sichuan; E: #0867 from Cang Shan Mt. Range,

Yunnan; F: #0251 from Mount Emei, Sichuan. Scale: 1 mm.

Sichuan, NE slope Gongga Shan, N29°49’29”

E102°03’24”, 21 .vi.201 1, 2986m, sift25, V.Grebennikov”.

Mount Emei, clade EM-A (10 in total, CNC): 10 exx,

#1216-9: #6184-9: “P.R. CHINA, Sichuan, EmeiShan,

N29°32’37.3 E103°19’57.5, 18.vi.2010, 2440m, sifting37,

V.Grebennikov”; clade EM-B (6 in total, CNC): 4 exx,

#0251-3, #0504: “P.R. CHINA, Sichuan, Emei Shan,

N29°32.932’ EI03°20.466\ 01.vii.2009, 2310m, sift-

ing 1 4, V.Grebennikov”; 1 ex, #0505: “P.R. CHINA,

Sichuan, Emei Shan, N29°32.806’ E103°20.106\

03.vii.2009, 2342m, sifting 1 5, V.Grebennikov”; 1 ex,

#1172 and #4186 (two numbers on the same specimen):

“PR. CHINA, Sichuan, EmeiShan, N29°32’37.3

E103°19’57.5, 18. vi. 2010, 2440m, sifting37, V.Greben-

nikov”; clade EM-C ( 1 in total, CNC): 1 ex, #1105: “P.R.

CHINA, Sichuan, EmeiShan, N29°33’36.3 E103°20’38.0,

15.vi.2010, 1947m, sifting33, V.Grebennikov”.

Distribution. The species is known from five isolated high

altitude forest regions in Yunnan and Sichuan provinces

(Fig. 9).

Etymology. The species epithet is the Latinized Greek

mythical name of Ismene, a daughter and half-sister of

Oedipus, daughter and granddaughter of Jocasta, and sis-

ter of Polynices; noun in apposition.

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Morimotodes , a new weevil genus from China

131

sternite 9

Fig. 6. Morimotodes polynices g. & sp. n., holotype, male, #5949 from Taiwan. A-D: habitus; E-G: aedeagus and tegment, dor-

sal (E), ventral (F), lateral; (G); H: sternites 8 and 9.

Morimotodes polynices sp. n.

Fig. 4.

Diagnostic description. Holotype, male (Fig. 6). DNAnot

sequenced. Length: 2.00 mm. Dorsal outline of pronotum

in lateral view with hump in basal third; pronotum and ely-

tra dorsally with fine punctures and pilosity not visible at

magnification below 30x; each elytron without longitu-

dinal groove parallel to edge. Aedeagus rounded lateral-

ly, sternite 9 Y-shaped and about 3-5x wider than

apodeme; sternal apodeme 9 without extention anterior-

ly. Intraspecific variation. Length: 1.98-2.12 mm.

Material examined. Holotype male (CMN): #5949 “TAI-

WAN, Kaohsiung Hs. Rd. Abv. Tona For. Sta. Km 16-17,

1700-1 800m 28. V. 1998, A. Smetana (T190)”. Paratypes

(CNC, CMN), 26 in total: 2 exx, #5925-6: “TAIWAN,

Kaohsiung Hsien, Kuanshan trail at Kaunshanchi Riv.

2400 m 24.VI. 1992 A. Smetana (T94)”; 9 exx, #5927-35:

“TAIWAN, Kaohsiung Hsien, Peinantashan trail, 2000m

7. VII. 93 A. Smetana (T144)”; 1 ex, #0587: “TAIWAN,

Kaohsiung Hsien, Kuanshan trail above Kaunshanchi Riv.,

2550m, 22.VII.93 A. Smetana (T160)”; 1 ex, #5936: “TAI-

WAN, Kaohsiung Hsien, Peinantashan trail 2450m,

2. V. 1995 A. Smetana (T170)”; 13 exx, #5940-8, #5950-3:

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Vasily V. Grebennikov

Morimotodes ismene g. et sp. n.

. Mt. Emei, Sichuan, clade EM-B

#4186

#0252.1

#0253.1

#0505

#0251.1

Mt. Gongga, Sichuan

0 94

#2563

#2471

#2559

#2562

#2456

# 2409

#2473

#2472

#2560

. Mt. Haba, Yunnan

0.85

#4483

#4537

#4484

#4482

#4481

#4536

022

0.05 substitutions/site S.--14

0.03

0.01

0,57

0.07

0.24

0.33

0.23

, Mt. Emei, Sichuan, clade EM-A

t Mt. Emei, Sichuan, clade EM-C

#6185

#6186

. #6184

.#6188

■#1216

- #1217

#6189

#1218

#1219

0.37 7

[ Cang Shan Mt. Range, Yunnan

#1105

#0867

#0500

#0502

#0501

#0499

#1017

#1018

#1020

2^3 Gaoligong Shan Mt. Range, Yunnan

0.7

0.62

0.74

#0503

#0115.1

#0114

#0116.1

#0927

#0928 Holotype

#0144

#0977

Leiosoma reitteri

Lobosoma rausense

Niphadomimus spp

0.88

Ectatorhinus adamsi

Typoderus furcatus

Acall/nus tuberculatus

Hylobius sp.

Adexius scrobipermis

Plinlhus spp

Anchonidium unguiculare

Aparopion costatum

Pissodes punctatus

Thalasselephas spp

Euthycus sp.

Otibazo polyphemus

Thromboslernus cuculatus

Fig. 7. Maximum Likelihood inference phylogram positioning monophyletic Morimotodes ismene g. & sp. n. (in black) as a sis-

ter to Leiosoma reitteri among other analysed Molytinae genera (in grey) using the 658 nt of the mtDNA barcoding COl gene

fragment. Digits at intemodes are bootstrap values. The tree is rooted on Graptus circassicus (Entiminae; not shown). Seven ge-

ographically linked lineages of M. ismene g. & sp. n. are marked in red; note that Mount Emei uniquely harbours three unrelated

lineages. Red and italicized numbers are those of imaged specimens (Figs 1, 4, 5) illustrating respective clades. Two pink ovals

with question marks denote two nodes not recovered in the alternative BI analysis (Fig. 8). Inserted image of the forest illustrates

exact spot where the type specimen #0928 was found.

same data as holotype. Additional non-type specimens:

3 exx, (CMN), #5937-9: “TAIWAN, Ilan Hsien Shen-Mi

Lake 24°22’43”N 121°44’12”E, 1110m I0.V.1995 A.

Smetana (T 1 77)”.

Distribution. The species is known from two widely sep-

arated regions in southern and northern Taiwan, respec-

tively (Fig. 9).

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Morimotodes , a new weevil genus from China

133

Pliocene

Pleistocene

Mt. Gongga, Sichuan

Mt. Haba, Yunnan

Mt. Emei, Sichuan, clade EM-A

Cang Shan Mt. Range, Yunnan

Mt, Emei, Sichuan, clade EM-C

Mt. Emei, Sichuan, clade EM-B

Gaoligong Shan Mt. Range,

Yunnan

Fig. 8. Ultrametic tree of Morimotodes ismene g. & sp. n. reconstructed using Baeysian inference method strict linear molecular

clock approach to analyse 658 nt of the mtDNA barcoding COl gene fragment and calibrated at a rate of 0.018

substitutions/site/MY. Numbers at nodes and on the scale below are million years before present. Node bars represent 95% confi-

dence intervals of the age estimate. Two pink ovals with question marks denote two nodes not recovered in the alternative ML

analysis (Fig. 7).

Fig. 9. Known distribution of Morimotodes g. n.

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Vasily V. Grebennikov

Etymology. The species epithet is the Latinized Greek

mythical name of Polynices, the son of Oedipus and Jo-

casta, as well as a grandson of Jocasta; noun in apposi-

tion.

RESULTS

The first tree search resulted in a Maximum Likelihood

tree (Fig. 7) with the highest log likelihood of- 9190.09.

All analyzed specimens of M. ismene g. & sp. n. formed

a weakly supported clade (bootstrap value 25%). The

genus Morimotodes g. n., represented by only its type-

species, formed a moderately supported (52%) clade with

the genus Leiosoma. Forty-six analyzed specimens of M.

ismene sp. n. from five mountain regions formed seven

geographical lineages (one monotypical, six others sup-

ported with 100% bootstrap). Four of these lineages cor-

responded to four mountain regions (Fig. 7), while all 15

analyzed specimens from Mount Emei formed three

groups not most closely related to each other. Temporal

(second) analysis in BEAST (Fig. 8) recovered all Mori-

motodes specimens forming a clade with the same seven

lineages as in the first analysis, although the grouping

among themselves was not identical with the results of the

first analysis. The inferred timing of the splits forming the

seven geographically linked lineages within monophylet-

ic M. ismene g. & sp. n. detected in the second analysis

date between 10.34 MY to 3.46 MY (Fig. 8).

DISCUSSION

Monophyly and sister group of Morimotodes g. n.

As presently defined, the genus Morimotodes g. n. is most

likely monophyletic. This assertion is not tested by the

cladistic methods, and is based entirely on circumstantial

evidence. None of the 29 specimens of M. polynices g. &

sp. n. responded positively to a DNA amplification effort

and, therefore, the species was not represented in the DNA

analyses. Continental populations of Morimotodes g. n.

from Yunnan and Sichuan all assigned to a broadly defined

M. ismene g. & sp. n. were consistently recovered as a

monophyletic clade in the first tree analysis (Fig. 7). The

assumption of both Morimotodes g. n. species forming a

clade is based entirely on their adult external similarities,

further corroborated by the analogous collecting circum-

stances suggesting similar biology. The geographical gap

of about 2000 km separating known distributional areas

of both species is not large enough to shed significant

doubts on their sister group relationships. Such are all

presently available data to justify the implemented taxo-

nomic decisions of grouping both species into a presum-

ably monophyletic new genus.

The most surprising aspect of the first analysis is the rel-

atively weak statistical support for M. ismene g. & sp. n.

(25%, Fig. 7), which can be at least partly attributed to the

known phylogenetic limitations of COl (Maddison

2012), including the saturation effect.

The sister-group relationships between Morimotodes g.

n. and the West Palaearctic Leiosoma is moderately sup-

ported (bootstrap 52% ( Fig. 7). Close affinities of both

genera have been hypothesised ever since the first repre-

sentatives of Morimotodes g. n. came to notice in 2008,

and was grounded entirely on adult morphological simi-

larities (small weevils less than 3 mm in body length with

laterally rounded and not flattened bodies and with shiny

dorsum) and biological preferences (inhabitants of the de-

ciduous leaf litter). Analogous expectations for the simi-

larly shaped genus Otibazo were, however, rejected by the

ML tree, the latter clustered with the genus Euthycus (Fig.

7). At present, it is impossible to suggest phylogenetic

affinities of the clade Morimotodes g. n. + Leiosoma , since

the available data are highly inadequate. Pending further

investigation, the genus Morimotodes g. n. is placed in the

same higher taxonomic group containing its sister group

Molytinae: Molytini: Molytina.

If Morimotodes g. n. is indeed sister to Leiosoma , then

attribution of both new species from China to the former

genus could be a phylogenetically sound taxonomic alter-

native. It was, however, rejected because establishing a

new genus better reflects the significant external dissim-

ilarity between their members and the wide gap in their

known geographical distribution.

Species delimitation of Morimotodes g. n.

Both species of Morimotodes g. n. as presently defined

are much dissimilar to each other. The species from Tai-

wan, M. polynices sp. n., is known from a museum series

of externally remarkably similar specimens, all but three

of which were sampled from a geographically confined

region (Kaohsiung, Fig. 9) not more than 50 km in its

greatest detention and within the limited altitudinal range

( 1 700-2550 m). This evidence suggests that all type spec-

imens most likely share the MRCA. Three additional non-

type specimens from north of Taiwan ( llan. Fig. 9) are ex-

ternally most similar to those from the type locality and,

therefore, are thought to be conspecific, although with

lesser certainty. These two hypotheses could not be test-

ed using DNA data, since the sequences were not obtain-

able. Judging from the morphological and geographical

data, it seems unlikely that the Taiwanese specimens of

Morimotodes g. n. could be grouped within the clade

formed by their continental counterparts. This evidence,

although not tested phylogenetically, strongly suggests

separate evolutionary history for the Taiwanese lineage

worthy, therefore, of establishing a formal species.

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135

Deciding what to all a species for the continental mem-

bers of Morimotodes g. n. is ambiguous. The most inclu-

sive and conservative approach is, therefore, employed.

All specimens of Morimotodes g. n. from continental Chi-

na are presently assigned to the broadly defined M. ismene

sp. n. This uncertainty on species boundaries is not made

any easier by the relatively large amount of available rel-

evant data for the continental species. The implemented

decision to assign all continental Morimotodes g. n. to a

single species is a balanced temporary solution implement-

ed to minimize the conflict among four independent lines

of evidence (in the increasing order of their contribution):

( 1 ) biological characteristics, (2) morphological similar-

ity/dissimilarity (3) geographical distribution, and (4)

mtDNA sequences. The logic of their combined evalua-

tion is described below.

The condition determining the gene flow among Mori-

motodes g. n. populations inhabiting five continental lo-

calities (the Cang Shan and the Gaoligong Mountain

Ranges, as well as Mounts Haba, Gongga and Emei; Fig.

9) can be considered analogous to those acting for terres-

trial organisms confined to oceanic islands. This approx-

imation is commonly used for the low-dispersing and

flightless moisture- and habitat-dependent species found

in relatively wet isolated forests widely separated by seem-

ingly unsuitable lowlands. Such estimate might perhaps

be even an understatement of the dispersal difficulties for

the mountain species, since the salt water barriers are oc-

casionally crossed by seemingly most unseaworthy organ-

isms such as chameleons possibly rafting on dislodged

floating trees, or blind subterranean Arthropods perhaps

surviving the passage in soil attached to their roots (Rax-

worthy et al. 2002 and Peck 1 990, respectively). The very

first task with such allopatric mountainous low-dispers-

ing organisms as Morimotodes g. n. was, therefore, to as-

sess whether the specimens from each separate locality

form distinct evolutionary clusters and whether any of the

localities supports more than one such cluster.

The four aforementioned lines of evidence contributed

conflicting data. Biological data were the least informa-

tive, since all sampled specimens of M. ismene sp. n. were

sifted from seemingly similar forest litter and within com-

parable altitude range. The only exception was shown by

the Mount Haba specimens recovered from the upper for-

est zone at altitude notably higher than known for other

continental populations. Adult external morphology was

the second least informative line of evidence (see below

on the use of genital characters). Although the imaged

specimens representing each of seven clades were some-

what dissimilar to each other (note the hairy specimen

#4484 from Mount Haba, or the large and non-hairy spec-

imen #1105 representing the clade EM-C from Mount

Emei; Figs 4, 5), it was impossible to consistently distin-

guish the others. The most powerful signal came from the

mtDNA sequences, which grouped specimens from four

localities into four strongly supported corresponding

clades (Fig. 7). The exception was Mount Emei, where 14

specimens clustered in two unrelated groups (EM- A and

EM-B on Fig. 7) with an additional third equally unrelat-

ed lineage (EM-C) represented by a single specimen

#1 105. The emerging distributional pattern indicated that

each locality supports at least one lineage of Morimotodes

g. n., while Mount Emei has at least three such lineages.

The main operational problem was that the majority of the

seven detected radiations could neither be uniquely char-

acterized morphologically, nor biologically.

It should be noted that detecting seven evolutionary sep-

arate lineages of continental Morimotodes g. n. inhabit-

ing five discrete mountain regions is not prohibitive in it-

self. There are no reasons to expect that no more than a

single radiation of closely related low-dispersing organ-

isms should exist in any suitable locality. Among all five

continental localities. Mount Emei should be rightfully ex-

pected to be the most diverse in Morimotodes g. n., be-

cause it is overwhelmingly rich in other forms of life and

by that exceeds many other comparable localities. For ex-

ample, its relatively well documented flora condensed on

just 154 km2 and lacking the entire alpine zone, encom-

passes an astonishing 3703 species of higher plants (Wang

et al. 2013). This diversity exceeds Canada (3,156;

Mosquin et al. 1995) and is comparable with that of the

British Isles (3842, Fitter & Peat 2014). Summing up, the

phylogeographic pattern as represented on Fig. 7 did not

pose significant biological inconsistencies. By far the ma-

jor challenge was the necessity to implement the Linnaean

taxonomy and decide on how many nominal species are

there.

At this stage the purely practical limitations of available

material has to be mentioned. Mostly, specimens of Mo-

rimotodes g. n. are relatively rarely encountered. For ex-

ample, the radiation inhabiting the Cang Shan Mountain

Range responded well to the original sifting in 2008, how-

ever, DNA sequencing of the 2008 specimens resulted in

only short fragments of 407 bp (four specimens

#0499-0502). During the subsequent two visits to Cang

Shan in 2010 and 2012, a significant greater sampling ef-

fort resulted in the collection of only a single specimen

(#0867), but provided a full-length DNA barcode. Mount

Emei has also been visited three times with one month of

intense fieldwork in total. Among the 27 sifting samples,

only five contained representatives of Morimotodes g. n.

Four focussed samples were taken in two years within 50

meters from where the unique specimen #1105 (lineage

EM-C, Fig. 7) had been sifted, but 1 failed to discover ad-

ditional specimens. The entire lineage EM-A was recov-

ered from a single sample. Only the lineage EM-B was

detected in three samples, two of them taken in exactly

the same place in different years and the third one some

650 m away. Such sampling results strongly suggest that

specimens of the continental Morimotodes g. n. are most

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Vasily V. Grebennikov

unevenly distributed in time and across seemingly iden-

tical landscape of mountainous forest.

Species delimitation in insects often relies on the shape

of male genitalia. However, due to the scarcity of Mori-

motodes g. n. specimens coupled with possible structur-

al damages during dissection of the minute genitalia, I re-

frained to examine genitalia of all sequenced specimens.

It was impossible to sex specimens externally and, there-

fore, male genitalia dissection had to be attempted with a

randomly chosen specimen, some of them females. Even

more problematic were the difficulties linked to the me-

chanical body properties of these highly compact minute

beetles with globular elytra firmly fused to the hind body

and forming together a most robust, inflexible and, accord-

ingly, highly breakable capsule. After the muscle tissue

was nearly completely dissolved and the remaining inter-

nal structures much loosened, the genitalia were success-

fully dissected for the two holotypes, only. It was, there-

fore, judged inadvisable to perform extensive male gen-

italia dissections.

The practical conclusion reached after considering all

was to define preliminarily a single species for all conti-

nental representatives of Morimotodes g. n., even though

this taxon might be later split into more separate species.

The Haba Shan lineage having distinct external adult mor-

phology further corroborated by the unique biological

preferences of higher altitudes can perhaps merit its own

species even at present. This action, however, will render

the rest of the continental Morimotodes g. n. paraphylet-

ic and, therefore, is not implemented. Considering a like-

lihood of further taxonomic changes in the broadly defined

M. ismene sp. n., the type series is purposely restricted to

the specimens forming a separate lineage uniquely char-

acterised by DNAand geographical data. Specimens from

the Gaoligong Mountain Range forming the extreme west

of the known generic distribution (Fig. 9) were chosen for

this purpose, specifically because they are numerous in the

DNA analysis. Additionally, this lineage is represented by

specimens recovered in at least six separate samples tak-

en from the area measuring 25 km in its greatest dimen-

sion and, therefore, having the least chances of harbour-

ing an undetected sympatric congeneric lineage (i.e., like

Mount Emei with at least three such lineages). Conse-

quently, all specimens of M. ismene sp. n. from localities

other than the Gaoligong Mountain Range are not desig-

nated as paratypes.

Dated phylogeography of Morimotodes ismene g. & sp. n.

Little can be presently said about the temporal aspect of

Morimotodes g. n. evolution. The single and seemingly

most interesting aspect of it, namely the time of the im-

plied separation between continental M. ismene sp. n. and

M. polynices sp. n. from Taiwan cannot be addressed at

present because of the lack of mtDNA data to represent

the latter species. Even though all seven lineages consti-

tuting monophyletic M. ismene sp. n. were consistently re-

covered in the first (ML) and the second (BI) analyses,

their grouping was consistent in only three nodes (Fig. 8):

(a.) the split between lineages inhabiting Mount Haba and

Mount Gongga, (b.) their MCRA separating from the lin-

eage EM-A inhabiting Mount Emei and (c.) the split be-

tween Mount Emei clade EM-B and the lineage from the

Gaoligong Mt. Range. All four consistently recovered evo-

lutionary events (three aforementioned and the origin of

the entire M. ismene sp. n.) date 4.45 MY, 8.08 MY, 3.46

MY and 10.34 MY, respectively. These estimations pro-

vide a very approximate time frame for the origin and di-

versification of the continental Morimotodes g. n., which

is comparable to those of the sympartically distributed

weevil genera Niphadomimus Zherikhin, 1987 (Greben-

nikov 2014) and Notaris Gennar, 1817 (Grebennikov &

Kolov, unpublished) likely linked to the advanced stages

of the uplift of the Tibetan Plateau (Favre et al. 2014).

Illustrated overview of genera of the subtribes Molyti-

na and Plinthina

The catalog of Alonso-Zarazaga & Lyal ( 1 999, 2002) had

all genera currently assigned to Molytina and Plinthina al-

ready listed as such (except for two proposed later) and

all included in the broadly defined Molytini. Since then

the following changes were proposed: ( 1 ) the monotypic

Stenanchonus Voss, formerly of Anchonini, was syn-

onymized with Euthycus Pascoe of Molytina (Meregalli

2003); (2) some species of the genus Minyops Schoenherr

of Plinthina was assigned to the newly erected Paraminy-

ops Osella & Bello (Osella & Bello 2010); (3) the tribe

Sthereini was re-erected to incorporate nine genera, four

of them formerly of Plinthina (Alonso-Zarazaga 2013;

Lyal 2014); (4) the former Molytini subtribe Typoderini

was elevated to tribe level; (5) the former Molytini sub-

tribe Leiosomatina was synonymized with Molytina (Lyal

2014); (6) the former tribe Hylobiini was synonymized

with Molytini by forming its third and the last subtribe

(Lyal 2014); (7) one new genus, Morimotodes g. n., be-

ing described herein. Moreover, the Palaearctic represen-

tatives of both subtribes accounting for about 80% of their

species were recently catalogued (Alonso-Zarazaga 20 1 3).

All 30 genera presently assigned to both subtribes are

listed below and 27 of them are illustrated. This summa-

ry was done with the intention to bring them to the no-

tice of the contemporary scientific community and hope-

fully trigger their reinvestigation in a phylogenetic frame-

work using modem data and analytical methods. Up to

now very little of such work has been done for the entire

subfamily Molytinae, including both Molytina and Plinthi-

na. Most importantly, reciprocal monophyly of both sub-

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Morimotodes, a new weevil genus from China

137

tribes has never been demonstrated and, therefore, their

taxonomic existence as currently defined seems very pre-

carious yet. Moreover, with the exception of Craw’s

(1999) detailed analysis of the New Zealand’s Hadram-

phus & Lyperobius , none of the non-monotypical genera

shown in detail below has ever been tested for its mono-

phyly. Some phylogenetically relevant statements were,

however, made: Meregalli (1985) suggested a sister-group

relation between Minyops (then containing Paraminyops)

and Plinthus. Davidian (2008) proposed that the lineage

presently consisting of Paraminyops and Minyops is most

closely related to a subgroup of the genus Plinthus (ren-

dering the former paraphyletic) and that Hoplopteridius

forms a clade with all of them. Osella & Bello (2010),

when separating their new genus Paraminyops from Miny-

ops, suggested that both genera together with PIo-

plopteridius form a group (“subtribe Minyopina”, a syn-

onym of Plinthina, Bouchard et al. 2011 ). None of these

hypotheses have been analytically tested.

Subtribe Molytina

The tribe includes 13 genera, which are numbered and list-

ed alphabetically below. Nine of them, as noted by Lyal

(2014), are widely scattered from New Zealand and Aus-

tralia, through New Guinea, Borneo, Java, and Indochi-

na, to Sri Lanka, eastern India, China and Japan. Three

other genera ( Anisorhynchus , Leiosoma and Liparus) are

distributed in the Western Palaearctic, with the former al-

so known from Central Asia (Alonso-Zarazaga & Lyal

1999). One genus known only from the holotype (Ptero-

tomus) is restricted to Angola.

Anisorhynchus Sehoenherr, 1842 (Figs 10A, B). This is

a genus of 13 large-bodied species and one non-nomina-

tive subspecies restricted to the West Palaearctic and dis-

tributed mainly around the Mediterranean Sea (Alonso-

Zarazaga 2013).

Apterylobius Marshall, 1942 (Figs 10C, D). Fewer than

a dozen specimens of this genus are known, all exclusive-

ly from Mount Kinabalu in northern Borneo. Since its de-

scription (Marshall 1942), the genus contains two species

and one non-nominative subspecies, and no newly collect-

ed specimens have been since reported.

Carbonomassula Heller, 1908 (Figs 10E, F, 16A). Spec-

imens of this genus, apart from their much greater size,

markedly resemble those of Morimotodes g. n. in having

similarly evenly rounded and dorsally lustrous body of

reddish or brownish color (compare Figs 6A and 1 0E). The

genus contains the single species from Papua New Guinea,

C. glaberrima Heller, 1908. The original description was

based on an unknown number of syntypes and accompa-

nied by a high quality habitus drawing (Heller 1908). The

type locality is “Nova Guinea Britannica, ad flumen

Aroa”. Two syntypes are in MTD, the smaller one best fit-

ting the body length given in the original description is

illustrated herein (Figs 10E, F). Both syntypes have a blue

rectangular label “14523” (Fig. I6A), even though the

original description mentions “14525”. One additional and

seemingly conspecific specimen supplied to BMNH by

M.E. Bacchus in 1965 has the locality label “NEW

GINEA: Morobe Dist., Herzog Mts., Vagau, C.4,000 ft.

4-7.L1965”. No other specimens of this genus have been

reported (Set! iff 2007).

Clarkanthus Alonso-Zarazaga & Lyal, 1999 (Figs 10G,

H, 1 6B). This genus consists of two poorly known species

from Sri Lanka and the adjacent part of India. Its type

species, C. taprobanus (Faust, 1895) was described from

the holotype collected on “Ceylon". The genus has long

been known under a preoccupied generic name Ischno-

pus Faust, year (non Amyot). Second species, C.

madrasensis Pajni & Sood, 1982, was described from a

holotype collected in the South Indian state of Tamil Nadu

just across the Palk Strait from Sri Lanka. Except for the

two type specimens, representatives of the genus have not

been reported in the literature. Two additional historical

specimens are present in MTD: one seemingly conspecif-

ic with the type species and labelled “Ceylon Plason” in

Faust’s handwriting, and another at least congeneric and

with a printed label “India or. Madura”. The locality might

perhaps refer to the Madurai District in Tamil Nadu, which

is the type locality of C. madrasensis. These four speci-

mens are the sole representatives of this obscure genus.

Euthycus Pascoe, 1885 (Figs 11 A, B). This genus em-

braces 12 species distributed at the junction between the

Palaearctic and Oriental Regions and recorded from Japan,

southern China and Taiwan, southwards to Borneo and Ja-

va, and westwards to Assam, Sikkim and Andaman Islands

of India (Meregalli 2003). Little is known of their natu-

ral history, aside from appearing to be associated with mid-

altitude primary mountainous forests. Observation of two

live specimens personally collected by sifting litter in Tai-

wan and represented in the DNA analyses revealed re-

markably strong adhesive power of their tarsi. This sug-

gests that the beetles are climbers rather than true litter

inhabitants (such as Morimotodes g. n.), where they might

be encountered only accidently.

Hadramphus Broun, 1911 (Figs 1 1C, D). This genus in-

cludes four relatively large-bodied species restricted to

New Zealand and some nearby islands and was revised

by Craw (1999, including a summary on the immature

stages and host plants). Together with Lyperobius , this

genus can be considered as a weevd analogous to the moa

or weta (i.e., flightless giant birds or crickets), that evolved

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138

Vasily V. Grebennikov

*

1

Fig. 10. Representatives of the Molytini: Molytina genera. A-B: Anisorhynchus barbatus (Rossi), Italy, Abruzzo, no date, no col-

lector, type species of Anisorhynchus , MTD; C-D: Apterylobius basalis (Heller, 1922), Malaysia, Borneo, no date, no collector,

?syntype, MTD; E-F: Carbonomassula glaberrima Heller, Papua New Guinea, syntype, see text, type species of Carbonomassu-

Icy MTD; G- H: Clarkanthus taprobanus (Faust), Sri Lanka, holotype, see text, type species of Clarkanthus , MTD.

in the unique New Zealand biota mostly deprived of ter- species quickly driven to extinction by invasive humans,

restrial mammals and, therefore, highly vulnerable to in- all but one ( Tymbopiptus valeas Kuschel, 1987) known

vasive predatory species. Unlike the moa with all nine large New Zealand weevils continue to survive, despite

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Morimotodes, a new weevil genus from China

139

Fig. 11. Representatives of the Molytini: Molytina genera. A-B: Euthycus macilentus Pascoe, 1885, Indonesia, Java, Mt. Guntur,

no date. Overbeck, type species of Euthycus, MTD; C-D: Hadramphus tuberculatus (Pascoe), New Zealand, Christchurch, 1910,

no collector, BMNH. image: Vasily Grebennikov, © The Natural History Museum, London; E-F: Leiosoma deflexion (Panzer, 1 795),

Poland, Muszkowice, 6.V.1986, M.Wanat, type species of Leiosoma , CNC; G-H: Lipams glabrirostris (Kiistcr, 1849), Poland, Sudetes,

1 5.vii. 1 975, M.Kupczynski, CNC.

loosing most of their former range (Kuschel & Worthy

1996). Recently an extant population of H. tuberculatus

(Pascoe, 1877), a species thought to be extinct, was dis-

covered (Fountain et al. 2013).

Kyliparus Moromoto, 1982 (not illustrated). This genus

was proposed to accommodate a single species from

Japan, K. lewisii (Roelofs), previously attributed to Li-

parus. The original description of Kiliparus contains a

habitus dorsal image of a single female studied by Mori-

moto (1982, fig. 46E).

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140

Vasily V. Grebennikov

Leiosoma Stephens, 1829 (Figs 11H, F). All 32 species

and one non-nominative subspecies of this genus are re-

stricted to the West Palaearctic, particularly the northern

Mediterranean, with two species extended eastwards to the

Caucasus (Alonso-Zarazaga 2013). Numerous species de-

scriptions were made for localized allopatric populations

and, therefore, a revision is desirable to assess their sta-

tus as that of biologically valid entities.

Liparus Olivier, 1807 (Figs 1 1 G, FI), the type genus of

the subtribe Molytina, the tribe Molytini and the subfam-

ily Molytinae. The genus includes 1 7 species and two non-

nominative subspecies (Alonso-Zarazaga 2014) of large-

bodied weevils. The genus is most diverse in the West

Palaearctic, while two species of the subgenus Li-

parorhynchus Reitter form the extreme eastward bound-

ary of the generic distribution and are found in Central

Asia (Uzbekistan, Tajikistan, Afghanistan, Iran; Bajtenov

1974; Alonso-Zarazaga 2013).

Lyperobius Pascoe, 1876 (Figs 12A, B). Along with

Hadramphus, Lyperobius is another genus of large-bod-

ied and flightless weevils endemic to New Zealand. The

majority of its 1 6 species have highly restricted distribu-

tions due to human-induced habitat changes, including

predation by introduced rodents (Craw 1999).

Morimotodes g. n. (Figs 1-9). The genus includes two

small-bodied litter species from Southwestern China and

Taiwan, respectively, and is a sister to the West Palaeaerc-

tic Leiosoma. See the taxonomic comments above.

Pinacopus Marshall, 1932 (Figs 12C, D). This genus has

14 named species found in Malaysia (8 spp.), Thailand ( 1

sp. ), Laos (4 spp.) and northern India ( 1 sp.), with at least

some specimens “collected from shrubs, which are more

or less covered with lichens, by beating method” (Koji-

ma 2010).

Pterotomus Quedenfeldt, 1888 (Figs 12E, F, 16C). The

genus includes a single species, P. moebiusi Quedenfeldt,

1888, known from the holotype collected in Angola’s

province Malange and never reported since. The original

description included a high quality habitus drawing (Que-

denfeldt 1888).

Subtribe Plinthina

The subtribe Plinthina consists of 17 genera listed below.

Eleven of them are restricted to the Mediterranean region

and some, like Minyops, are also found in Central Europe,

Ukraine, southern Russia and western Kazakhstan. Two

genera, Kurilio and Metahylobius, with a total of three

species are limited to the eastern Pacific (Ffonshu,

Flokkaido and Kunashir). Four other inadequately known

mono- and oligotypic genera are restricted to the Afrotrop-

ical Region, namely Seychelles ( Gonotrachelus ), East

Africa ( Molytophilus and Plinthinella ) and Madagascar

( Vouauxia).

Adexius Schoenherr, 1834 (Figs 13A, B). This genus con-

sists of three West Palaearctic species, two of them poor-

ly known and narrowly distributed. The third one, A. scro-

bipennis Gyllenhal, 1834 is found in forest leaf litter in

most of Central and Southern Europe between Belgium

and Ukraine (Alonso-Zarazaga 2013).

Alloplinthus Solari, 1941 (Figs 13C, D). This genus con-

sists of three species (originally described as Plinthus)

from Tenerife and Gran Canaria (Alonso-Zarazaga 2013).

Caulomorphus Faust, 1886 (Figs 13E, F). The genus con-

tains 1 1 eyeless species from Turkey, Greece, Iran and

Azerbaijan (Alonso-Zarazaga 2013). Osclla (1970) revised

the genus taxonomically and later described three more

species.

Freyianus Zumpt, 1932 (Figs 13G, H). This is a

monobasic genus for F liparoides Zumpt, 1932. It was de-

scribed based on 10 syntypes (one of them adequately il-

lustrated by a habitus photo) collected by “Kulzer” in “Ak-

Schehir” (Zumpt 1932) and thought to be in Turkey (Alon-

so-Zarazaga & Lyal 1999). Since its description, this genus

has not been reported in the original literature and its iden-

tity is inadequately known. The newly imaged specimen

(Figs 13G, H), even if labelled as a “holotype”, is in fact

a syntype and, together with two other syntypes, is stored

in NHMB.

Gonotrachelus Champion, 1914 (Figs 131, J, 16D). The

genus includes a single species, G. quinquecarinatus

Champion, 1914 described from two syntypes found on

the Seychelles (Long Island, near Mahe). An additional

specimen of this species was reported from Madagascar

(Hustache 1925). The genus is characterized by the funi-

cle consisting of fine antennomeres (Champion 1914, figs

11, lib) and was originally compared with Stypldoderes

Wollaston (Typoderini). The type series (presently in

BMNFI) was taken from seashores and was thought to be

introduced via driftwood (Champion 1914).

Hoplopteridius K. Daniel, 1908 (Figs 14A, B). This genus

consists of five species and three non-nominative sub-

species distributed in the northern and eastern Mediter-

ranean (Alonso-Zarazaga 2013).

Iberoplinthus Meregalli, 1986 (Figs 14C, D). This recent-

ly erected genus (Meregalli 1986) consists of six species

(Alonso-Zarazaga 2013) all found in northern Spain, while

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Morimotodes, a new weevil genus from China

141

Fig. 12. Representatives of the Molytini: Molytina genera. A-B: Lyperobius huttoni Pascoe, 1876, New Zealand, Humboldt Mts.,

30.xii. 1 944, C.E. Clarke, type species of Lyperobius , BMNH, image: Vasily Grebennikov, © The Natural History Museum, Lon-

don; C-D: Pinacopus dentirostris Marshal, 1932, Malaysia, Perak, no date, no collector, MTD; E-F: Pterotomus moebiusi Que-

denfeldt, Angola, holotype, see text, type species of Pterotomus , MNHN.

the type species, I. imbricatus (Dufur, 1851), and is re-

ported also from Andorra and southern France.

Kurilio Zherikhin & Egorov, 1991 (Figs 14E, F). Its on-

ly species, K. monachus Zherikhin & Egorov, 1 99 1 , is re-

stricted to Hokkaido and Kunashir Islands (Zherikhin &

Egorov 1990).

Metahylobius Nakane, 1964 (Figs 14G, H). This genus

consists of two species, both restricted to the alpine zone

of Honshu: M.jonensis Nakane, 1964 and M. rubiginosus

Morimoto, 1982. The adults of the former species “were

captured by turning stones on the ground” (Morimoto

1982).

Minyops Schoenherr, 1823 (Figs 141, J). This genus con-

sists of 29 species and two non-nominative subspecies dis-

tributed through most of southern Europe and also Turkey

(Alonso-Zarazaga 2013). The genus was recently taxo-

nomically revised with 22 of its 3 1 species-group taxa de-

scribed as new and eight others re-validated from syn-

onymy (Osella & Bello 2010).

Mitoplinthus Reitter, 1897 (Figs 15 A, B). This genus con-

sists of two species and one non-nominative subspecies

of somewhat cylindrical medium-sized weevils distributed

in southern parts of Europe between Spain and Ukraine.

Meregalli (1986) provided the most recent treatment of

the genus.

Molytophilus Hartmann, 1904 (Figs 15C, D). The genus

consists of three species, the type species Molytophilus

carinatus Hartmann, 1904, described from Tanzania, and

two more from Ethiopia: M. affinis Hustache, 1936, and

M. puncticollis Hustache, 1936, whose affinities with the

genus have not yet been demonstrated.

Bonn zoological Bulletin 63 (2): 123-147

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142

Vasily V. Grebennikov

Fig. 13. Representatives of the Molytini: Plinthinae genera. A-B: Adexius scrobipennis Gyllenhal, Poland, Holubla ad Przemysl,

4.vi,1997, M.Wanat, type species of Adexius, CNC; C-D: Alloplinthus musicus (Wollaston, 1860), Spain, Teneriffe, 1 7.V.1924, E.Ap-

penhagen, type species of Alloplinthus, BMF1N, image: Vasily Grebennikov, ©The Natural History Museum, London; E-F: Caulo-

morphus lederi (Chevrolat, 1880), Georgia, “Meskisch. Gb.”, no date, H.Leder, ?syntype, type species of Caulomorphus , MTD;

G-H: Freyianus liparoides Zumpt, Turkey, “Ak-Schehir”, 22.V.1926, Kulzer, syntype, type species of Freyianus, NHMB; 1 -J: Gono-

trachelus quinquecarinatus Champion, Seychelles, see text, syntype, type species of Gonotrachelus, BMNH, image: Vasily Greben-

nikov, © The Natural History Museum, London.

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Morimotodes, a new weevil genus from China

143

Fig. 14. Representatives of the Molytini: Plinthinae genera. A-B: Hoplopteridins lutosus (Frivaldszky, 1835) Italy, Rom, no date,

no collector, type species of Hoplopteridius , MTD; C-D: Iberoplinthus curvirostris Meregalli, 1986 Spain, Asturias, 1 4.vii. 1 986,

M.Meregalli, paratype, BMNH, image: Vasily Grebennikov, © The Natural History Museum, London; E-F: Kurilio monachus

Zherikhin & Egorov, Russia, Kunashir Isl., 1 6.ix. 1 975, no collector, type species of Kurilio , CNC; G-H: Metahylobius jonensis

Nakane, Japan, Honshu Isl., 1 .vii. 1 986, S.J. Martin, type species of Metahylobius , BMNH, image: Vasily Grebennikov, © The Nat-

ural History Museum, London; 1 J: Minyops carinatus (Linnaeus, 1767), Austria, Wien, no date, no collector, type species of Miny-

ops , MTD.

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144

Vasily V. Grebennikov

Fig. 15. Representatives of the Molytini: Plinthinae genera. A-B: Mitoplinthus caliginosus (Fabricius, 1775), Germany, Amstadt,

16.x. 1949, W.Liebmann, MTD; C-D: Molytophilus carinatus Hartmann, Tanzania, Usambara, no date, no collector, syntype, type

species of Molytophilus, MTD; E-F: Neoplinthus tigratus porculus (Fabricius, 1801 ), Austria, Graz, no date, no collector, MTD;

G-H: Plinthus confiusus Meregalli, 1985, Russia, Caucasus, 21. vi. 1974, J.Vorisek, paratype, CNC; I-J: Vouauxia onopatroidea Hus-

tache, Madagascar, see text, syntype, type species of Vouauxia , MNITN.

Bonn zoological Bulletin 63 (2): 123-147

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Morimotodes, a new weevil genus from China

145

Q’L.H'.'tyw.lA,

j>. Vivjk*

Stoatl. Museum fur

Tierkunde. Dresden

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TijJurV

(JoH, J. Fausts

Ankauf 1900-

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Trust Expedition.

1913.— 170.

Tr.Linn.S.L 1914.

Goifotrachelus !

D^CQniuituSj Cli.

n

frialat,

MUSEUM PARIS

MADAGASCAR

Collection Uon Fairmaire

1906

COTYPE

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Col. A. HUSTACHE

Fig. 16. Type specimen labels. A: Carbonomassida glaberrima Heller, ST; B: Clarkanthus taprobanus (Faust), holotype; C: Ptero-

tomus moebiusi Quedenfeldt, holotype; D; Gonotrachelus quinquecarinatus Champion, syntype, image: Vasily Grebennikov,

© The Natural History Museum, London; E: Vouauxia onopatroidea Hustache, syntype. Labels within each species are of propor-

tional size; habitus images are not to scale.

Neoplinthus Bedel, 1884 (Figs I5E, F). This genus in-

cludes five species and two non-nominative subspecies of

elongated and parallel-sided weevils distributed in Cen-

tral, Eastern and Southern Europe; all of them recorded

from Italy (Alonso-Zarazaga 2013).

Paraminyops Osella & Bello, 2010 (not illustrated). This

genus was recently separated from Minyops by Osella &

Bello (2010) to accommodate nine species and two non-

nominative subspecies from northern and eastern Mediter-

ranean.

Plinthinella Hofmann, 1965 (not illustrated). This genus

includes its sole member, the type species by original des-

ignation P. tenella Hoffmann, 1965. This is a small bee-

tle of some 2. 0-2. 2 mm in body length and funicle with

seven antennomeres described from the holotype and 12

paratypes collected in 1962 by Herbert Frantz. The type

locality in “Tanganyika: Kilimandjaro” (= Mt. Kiliman-

jaro in Tanzania) and the altitude is unknown. When de-

scribed, this taxon was compared with Matxaldiella

glacialis (Hustache, 1929) (cited as “ Mimus glacialis ”)

from Mount Kenya in Kenya (depicted in Hustache 1929:

fig. 15). Since its description, Plinthinella tenella has not

been reported in the original literature and its identity is

inadequately known. The curatorial search to locate the

type series in NMW failed, while my extensive recent

sampling in the type locality did not rediscover the species.

Both the generic and species names should, therefore, be

considered as nomina dubia.

Bonn zoological Bulletin 63 (2): 123-147

Plinthus Gerinar, 1817 (Figs 15G, H). This genus in-

cludes 68 species and 17 non-nominative subspecies of

medium- to large-bodied weevils widely distributed in the

West Palaearctic. The majority of species are found in

mountains between the Pyrenees and Alborz in northern

Iran (Alonso-Zarazaga 2013). The Caucasus appears to

have disproportionally large number of species normally

restricted to alpine mountaintops. This is one of a few gen-

era receiving significant attention in recent years, includ-

ing detailed (although mainly unpublished) studies on in-

ternal classification, structure of male genitalia, species

distribution and biology (Davidian 2008).

Vouauxia Hustache, 1920 (Figs 151, J, I6E). The genus

includes a single species, O. onopatroidea Hustache, 1920,

described from Madagascar from unknown number of

syntypes. The original brief description (Hustache 1920)

was followed by a more detailed one (Hustache 1925),

both relating this taxon to Gonotrachelus Champion. At

least three syntype are present in MNHN. No other spec-

imens of this genus have been reported in the literature.

CONCLUDING REMARKS

This review revealed, how little is actually known about

wingless Molytinae genera, especially those, which are

presently assigned to the subtribes Molytina and Plinthi-

na. The discovery of a peculiar new genus from China

strongly suggests that similar findings still remain to be

made, especially using specialized collecting techniques

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146

Vasily V. Grebennikov

and addressing specific habitats harbouring cryptic and

small-sized organisms. Equally remarkable is the acute

scarcity of data on the ma jority of other genera herein dis-

cussed. In many cases species and even genera were men-

tioned only once when formally described. Many mono-

and oligobasic genera such as Apteiylobius, Carbonomas-

sula, Clarkanthus, Kyliparus, Pterotomus , Freyianus ,

Gonotrachehis , Molytophilus , Plinthinella and Vouauxia

are particularly poorly known and at least some of them

appear only distantly related to the type genera of their

respective subtribes. A detailed morphological and, par-

ticularly, DNA analysis of Molytinae is indeed long over-

due. Much remains to be done with wingless Molytinae

weevils and hopefully this paper helps to reinforce this

seemingly self-evident conclusion.

Acknowledgement. Ales Smetana collected all specimens of

Morimotodes polynices g. & sp. n. in Taiwan. Curators of the

collections mentioned above variously helped in accessing spec-

imens under their care. Genrikh Ed. Davidian (St. Petersburg,

Russia) identified the sequenced specimens of Plinthus. Marek

Wanat (Wroclaw, Poland) sent imaged specimens of Adexius, Li-

parus and Leiosoma. Eva Sprecher contacted on my behalf Is-

abelle Ziircher-Pfander (both Basel, Switzerland), who counted

syntypes and took images of Freyianus liparoides (Figs 1 3G, H).

Thierry Deuve (Paris, France) sent a reprint of Hustache (1920).

Bradley J. Sinclair and Eduard Jendek (both Ottawa, Canada)

read an early draft of this work.

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Bonn zoological Bulletin 63 (2): 148-156

December 20 1 4

New records of amphibians and reptiles from southern Vietnam

Truong Quang Nguyen12, Trung My Phung3, Nicole Schneider4, Andreas Botov4, Dao Thi Anh Tran5,6

& Thomas Ziegler4*

' Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet, Hanoi,

Vietnam; E-mail: nqt2@yahoo.com

' Department of Terrestrial Ecology, Zoological Institute, University of Cologne, Ziilpicher Strasse 47b,

D-50674 Cologne, Germany

3 Dong Khoi 9 A, Tam Hiep, Bien Hoa, Dong Nai Province, Vietnam; E-mail: pmytnmg@yahoo.com

4 AG Zoologischer Garten Koln, Riehler Strasse 173, D-50735 Cologne, Germany

3 Vietnam National University, Ho Chi Minh City, University of Science, Faculty of Biology, 227 Nguyen Van Cu, District 5,

HCM city, Vietnam ; E-mail: ttadao.hcmuns@gmail.com

6 Zoologisches Forschungsmuseum Alexander Koenig, Adenaueralle 160, D-53113 Bonn, Germany

' corresponding author: E-mail: ziegler@koelnerzoo.de

Abstract. We report six new records of poorly known species of amphibians and reptiles on the basis of a new herpeto-

logical collection from southern Vietnam: Ophryophryne gerti and Raorchestes gryllus from Khanh Hoa Province, Acan-

thosaura coronata from Ba Ria - Vung Tau Province, Gekko grossmanni from Ninh Thuan Province, as well as Cylin-

drophis ruffus and Oligodon cinereus pallidocinctus from Binh Phuoc Province.

Keywords. Ophryophiyne gerti , Raorchestes giyllus, Acanthosaura coronata, Gekko grossmanni, Cylindrophis ruffus,

Oligodon cinereus pallidocinctus, new records, taxonomy.

INTRODUCTION

Geographic distribution records are crucial for assessing

the conservation status and are also evidence to elucidate

species’ evolution and biology. Based on newly collect-

ed specimens of amphibians and reptiles from southern

Vietnam, we herein report six new records of poorly

known species, comprising two anuran species from

Khanh Hoa Province, one agamid species from Ba Ria -

Vung Tau, one geckonid species from Ninh Thuan

Province as well as one cylindrophiid and one colubrid

species from Binh Phuoc Province.

MATERIAL AND METHODS

Field work was conducted by Trung My Phung in Hon Ba

Nature Reserve (Khanh Hoa Province) in July 2010, on

Nui Dinh Mountain (Ba Ria -Vung Tau Province) in No-

vember 2008, in Ca Na forest (Ninh Thuan Province) in

March 2010 and by Trung My Phung and Thang Huu

Khuong in Bu Gia Map National Park in September and

October 201 1 (Fig. 1). After taking photographs, speci-

mens were anaesthetized, fixed in 80-85% ethanol and

subsequently stored in 70% ethanol. Specimens were de-

posited in the collections of the Institute of Ecology and

Biological Resources (1EBR) and Vietnam National Mu-

seum of Nature (VNMN), Hanoi, Vietnam and the Zool-

ogisches Forschungsmuseum Alexander Koenig (ZFMK),

Bonn, Germany.

Received: 07.02.2013

Accepted: 25.1 1.2014

Measurements of specimens were taken with a digital

caliper to the nearest 0. 1 mm. Abbreviation are as follows:

SVL (snout-vent length): from tip of snout to anterior mar-

gin of cloacal; TaL (Tail length): from posterior margin

ofcloacal to tip of tail; TL (total length): SVL + TaL. Ter-

minology of morphological characters followed Nguyen

et al. (2012) for amphibians (except for forearm length,

FAL: from axilla to elbow and hand length, HAL: from

base of outer palmar tubercle to tip of finger III), Phung

& Ziegler (2011) for lizards, and David et al. (2012) for

snakes.

RESULTS AND DISCUSSION

Taxonomic accounts

Megophryidae

Ophryophryne gerti Ollier, 2003

Gerti’s Mountain Toad / Coc nui got (Fig. 2)

Specimens examined: VNMN 983 and ZFMK 94220,

males, collected by T. M. Phung on 4 July 2010 from Hon

Ba Nature Reserve (ca. I2°10’N, 109°02’E), Dien Khanh

District, Khanh Hoa Province.

Corresponding editor: T. Ziegler

New herpetofaunal records from Vietnam

149

Fig. 1. Map of sampling sites in southern Vietnam: 1 ) Khanh

Hoa Province, 2) Ninh Thuan Province, 3) Binh Phuoc

Province, 4) Ba Ria - Vung Tau Province.

Morphological characters: Size small, SVL 28.2-29.7

mm (n = 2); head wider than long (HW 8. 5-8. 9 mm, HL

8. 1-8.7 mm); snout truncate, protruding, shorter than hor-

izontal diameter of eye (SL 2. 6-2. 7 mm, ED 3. 2-3. 7 mm);

canthus rostralis rounded, loreal region convex; interor-

bital distance narrower than upper eyelid and internarial

distance (IOD 4. 0-4. 3 mm, UEW 2. 6-2. 7 mm, IND

2. 5-2. 7 mm); nostrils closer to the eye than to the tip of

snout (EN 1.1-1. 4 mm, SN 1.2-1. 5 mm); pupil rounded;

tympanum oval, greater than half eye diameter and tym-

panum-eye distance (TD 1 .8-2.3 mm, TYE 1 .5-1.9 mm);

supratympanic fold distinct; vomerine teeth absent;

tongue rounded posteriorly.

Forelimbs: FAL 6. 5-7.4 mm, HAL 7. 5-7. 6 mm; fingers

free of webbing, relative length of fingers: I<II=IV<III;

tips of fingers rounded, without discs; dermal fringe along

outer finger absent; subarticular tubercles absent; palmar

tubercles indistinct. Hindlimbs: femur longer than tibia and

foot length (FML 15.3-18.7 mm, TBL 14.9-15.2 mm,

FTL 12.7-13.3 mm); toes long and thin, relative length

of toes: I<II<V<III<IV; tips of toes rounded; webbing

basal; dermal fringe along outer toe absent; subarticular

tubercles absent; inner metatarsal tubercle small (1MT

1.4- 1.5 mm), outer metatarsal tubercle absent, tarsal fold

absent.

Dorsal skin of head, body and limbs granular; dorsum

with several skin ridges, forming a X-shape in the mid-

dle and two other ridges running from behind tympanum

to posterior part of back; lateral sides of head and flank

granular, upper part of flank with some glandular warts;

orbital horn pointed; dorsolateral fold absent; throat, chest,

belly and ventral surface of limbs smooth; pectoral and

femoral glands present, small.

Coloration in life: Dorsal surface of head and body dark

brown; a dark triangular pattern between eyes present;

lower part of flank with some dark brown spots; lateral

side of head dark brown; dorsal surface of limbs with dark

bars; throat, chest, belly and ventral surface of limbs dark

brown with some black spots.

Remarks: Moqrhological characters of our specimens

agreed well with the description of Ohler (2003) but they

are a little smaller than the type series (SVL 28.2-29.7 mm

vs. 32.0-45.8 mm).

Distribution: In Vietnam, this species is known from

Thua Thien - Hue Province southward to Lam Dong

Province. Elsewhere, this species has been reported from

Laos (Nguyen et al. 2009).

Rhacophoridae

Raorchestes gryllus (Smith, 1924)

Langbian Bubble-nest Frog / Nhai cay Langbian (Fig. 3)

Specimens examined: VNMN 985 (adult female), VN-

MN 986 and ZFMK 94221-94222 (adult males), collect-

ed by T. M. Phung on 4 July 2010 from Hon Ba Nature

Reserve (ca. 12°10’N, 109°02’E), Dien Khanh District,

Khanh Hoa Province.

Morphological characters: Size small, SVL 21.9-24.2

mm (n = 4), head as long as wide (HL 9. 2-9. 8 mm, HW

9.2-9. 8 mm); snout pointed in dorsal view, a bit longer

than horizontal diameter of eye (SL 3.6-4. 1 mm, ED

3. 1-3.5 mm); canthus rostralis distinct; interorbital dis-

tance wider than internarial distance and upper eyelid

(IOD 24-2.7 mm, IND 1 .7-2.0 mm, UEW 2.3-2.4mm);

nostril lateral, rounded, nearly midway between tip of

snout and eye (SN 1.6-1. 8 mm); vomerine teeth absent;

tympanum rounded, smaller than half of eye diameter but

Bonn zoological Bulletin 63 (2): 148-156

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150

Truong Quang Nguyen et al.

Fig. 2. Gerti’s Frog Ophyryophryne gerti (a) and its habitat (b). Photos: Trung M. Phung.

Bonn zoological Bulletin 63 (2): 148-156

©ZFMK

New herpetofaunal records from Vietnam

151

Fig. 3. Langbian Bubble-nest Frog Raorchestes giyllus (a) and its habitat (b). Photos: Trung M. Phung.

greater than tympanum-eye distance (TD 1.2-1 .3 mm,

TYE 0.4-0. 6 mm); supratympanic fold distinct; tongue

notched posteriorly.

Forelimbs: FAL 4. 0-4.2 mm, HAL 6. 6-7. 7 mm; finger

webbing basal, relative length of fingers: I<II<1V<I1I; tips

of fingers enlarged into round discs, with circummargin-

al grooves; subarticular tubercles rounded and prominent;

series of tubercles present along outer side of forearm and

finger IV; nuptial pad present in males.

Hindlimbs: femur as long as tibia but longer than foot

length (FML 11.3-12.4 mm, TBL 11.2-12.3 mm, FTL

9.5 1 0.3 mm); relative length of toes: I<II<III<V<IV; tips

Bonn zoological Bulletin 63 (2): 148-156

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152

Truong Quang Nguyen et al.

of toes with enlarged discs; subarticular tubercles promi-

nent; inner and outer metatarsal tubercles present; web-

bing formula: I2-2II 1 y3-2 l/2III 1 %-2I V2- 1 V; tubercles

present along outer margins of tibia, foot and toe V; tibio-

tarsal articulation reaching to eye.

Dorsal skin covered by tiny tubercles; throat finely gran-

ular; belly and ventral surface of thigh coarsely granular,

small tubercle present on heel.

Coloration in life varied among individuals: Back-

ground of dorsal surface of head, body and limbs of males

light brown with reddish brown or bright green marbling;

a bright green triangle patch, covering anterior part of up-

per eyelid and snout, sometimes the patch combined with

the posterior bright green marbling; dorsum with two dark

brown stripes, forming a “)(” shape, from rear of eye to

groin, distinct or indistinct; dorsal surface of limbs with

dark brown transverse bars; throat, chest and belly white.

Female uniform green above and whitish beneath.

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New herpetofaunal records from Vietnam

153

Remarks: Our specimens differ from previous descrip-

tions (Smith 1924, Orlov et al. 2012) by having dermal

fringe along outside of tarsus less developed (versus well

developed in description of Orlov et al. 2012). Iris light

golden with brown or reddish mottles posteriorly, pupil

horizontal oval and black (not mentioned by Smith 1924

and Orlov et al. 2012). Specimens of R. gryllus from

Khanh Hoa Province are morphologically similar to Kurix-

alus viridescens , a newly described species from south-

ern Vietnam by Nguyen et al. (2014). However, R. gryl-

lus can be distinguished from the latter by the following

characters: the presence of a brown )(-shaped marking on

dorsum and upper part of flank (versus uniform solid green

dorsum in K. viridescens ), outer metatarsal tubercle pres-

ent (versus absent in Kurixalus viridescens ); toe webbing

formula I2-2II 1 'A-VAl II 1 %-2IV2- 1 V (versus I2-23/4II 1 Vi-

23/4IlI 1 1/2-3IV2,/2-l3/4V in K. viridescens ), and the iris light

golden with brown or reddish mottles posteriorly (versus

golden with some black reticulations in K. viridescens ).

Distribution: In Vietnam, R. gryllus was orginally discov-

ered from Langbian Plateau, Lam Dong Province by Smith

( 1924). This species was also reported from northern Viet-

nam: Lao Cai, Ha Giang, Cao Bang, Tuyen Quang, Lang

Son, Vinh Phuc, Thai Nguyen and Thanh Hoa provinces

as well as from central provinces: Dak Lak, Gia Lai, Kon

Turn (Nguyen et al, 2009). This is the first record of R.

gryllus from Khanh Hoa Province. Elsewhere, it has been

known from Champasak Province of Laos (Frost 2014,

and Orlov et al. 2012).

Againidae

Acanthosaura coronata Gunther, 1861

Coronated Tree Lizard / O ro vanh (Fig. 4)

Specimen examined: One adult female, collected by T.

M. Phung on 1 November 2011 from Nui Dinh Mountain

(ca. 10°30’N, 107°07’E), Tan Thanh District, Ba Ria -

Vung Tau Province. This specimen was released after ex-

amining of morphogical characters.

Morphological characters: SVL 90 mm, TaF 125.9 mm

(n=l); forepart of head deeply concave; scales on upper

head inhomogeneous, obtusely keeled, much smaller in

size in posterior part; canthus rostralis and supraciliary

edge distinct; a long spine behind the eye present; anoth-

er spine on the back of the head present, mid-way between

tympanum and nuchal crest; tympanum one-third to one-

half the diameter of orbit; 10-13 supralabials.

Dorsal scales small, keeled, intermixed with much larg-

er, strongly keeled scale; scales on upper rows larger in

size, pointing upwards, lateral scales pointing backwards

and upwards, sometimes downwards; ventrals as large as

(he largest dorsal scales, strongly keeled; gular sac absent;

gular scales strongly keeled, smaller than the ventrals; a

strong oblique fold present in front of shoulder; nuchal

crest with triangular spines; dorsal crest continuous with

the nuchal crest, reduced as a prominent ridge, composed

of broad scales; hind limb extending to the tip of the snout.

Tail compressed, subtriangular at base, covered with

keeled scales above, strongly keeled and elongated scales

below.

Coloration in life: Dorsal surface of head and body dark

grey; dorsum and tail base with four light bands, edged

by black posteriorly; head light green or yellow; a dark

cross-bar present between eyes; some dark lines radiat-

ing from eyes; belly cream (identification after Smith

1935).

Distribution: In Vietnam, this species has been known

from Dak Fak, Lam Dong, and Dong Nai provinces. Else-

where, it was reported from Cambodia (Nguyen et al.

2009).

Gekkonidae

Gekko grossmanni Gunther, 1994

Grossman’s gecko / Tac ke g-ro-s-man (Fig. 5)

Specimens examined: VNMN 960 (male), VNMN 959

and ZFMK 94125-94126 (females), collected by T. M.

Phung on 27 March 2010 from Ca Na forest (ca. 11°19’N,

108°49’E), Ninh Phuoc District, Ninh Thuan Province.

Morphological characters: SVL 71-86 mm, TaL

84.8-101.6 nun (n=4); head longer than wide (HL 20-23.9

mm, HW 13.1-17.2 mm), axillary-groin distance

29.2-34.4 mm.

Rostral wider than high, rarely with X- or Y-shaped su-

ture, touching nostril; supranasals 2 (rarely 3); internasals

0-1; interorbital scales between anterior corners of eyes

40^15; supralabials 11-12, infralabials 9-10; dorsum cov-

ered by small, unequal granular scales interspersed, flat

or slightly keeled tubercles; midbody scales rows

1 18-122; tubercles surrounded by 9 scales, in 11-13 rows

at midbody; tubercles absent on upper surface of limbs and

tail; lateroventral fold distinct; ventral scale rows between

lateral folds 29-3 1 ; ventral scales along underside of body

from mental to the cloacal slit 156-162; subcaudals 86-91,

enlarged; precloacal pores 1 3 (in male), femoral pores ab-

sent; finger and toe webbing basal; subdigital lamellae

10-14 under the first finger, 16-18 under the fourth fin-

ger, 11-15 under the first toe, 17-18 under the fourth toe.

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Truong Quang Nguyen et al.

Coloration in life: Dorsal surface of head, body and tail

grey intermixed with yellow dots and dark brown mar-

bling; dorsum with several rows of 7-8 light grey spots:

one along vertebral and two others on each side; light

bands present on upper surface of tail (identification af-

ter Gunther 1994, in comparison with Ngo & Gamble

2011).

Distribution: This species has been known only from the

type locality in Khanh Hoa Province, Vietnam (Nguyen

et al. 2009).

Cylindrophiidae

Cylindrophis ruffus (Laurenti, 1768)

Red-tailed Pipe Snake / Ran trim (Fig. 6a)

One specimen was photographed by T. H. Khuong in Oc-

tober 201 1 in Bu Gia Map National Park (ca. 12°10’N,

1 07° 1 2’E), Binh Phuoc Province.

Identification is based on the characteristic body shape

and colour pattern (after Smith 1943): head depressed,

snout round; tail very short. Coloration in life: dorsal head

Bonn zoological Bulletin 63 (2): 148-156

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New herpetofaunal records from Vietnam

155

Fig. 6. Red-tailed Pipe Snake Cylindrophis ruffus (a) and Guenther’s Kukri Snake Oligodon cinereus pallidocinctus (b) and its

habitat (c). Photos: Thang H. Khuong.

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156

Truong Quang Nguyen et al.

and body blackish brown, iridescent, with narrow light

cross-bars, extending to the middle of the back; an orange

ring present near the tip of tail; venter black with white

bars; ventral surface of tail with orange bars.

Distribution: In Vietnam, this species is known from Vinh

Phuc Province in the North southwards to Ca Mau

Province. Elsewhere, it has been recorded from China,

Myanmar, Thailand, Laos, Cambodia, Malaysia, Singa-

pore, and Indonesia (Nguyen et al. 2009).

Colubridae

Oligodon cinereus pallidocinctus (Bourret, 1934)

Guenther’s Kukri Snake / Ran khiem xam (Fig. 6b,c)

Specimens examined: IEBR A. 2012. 8, adult female, col-

lected by T. M. Phung and T. FI. Khuong on 12 Septem-

ber 201 1 from Bu Gia Map National Park (ca. 12°10’N,

107°12’E), Binh Phuoc Province.

Morphological characters: SVL 540 mm, TaL: 72 mm;

maxillary teeth 12/12; loreal 1/1; supralabials 8/8, fourth

and fifth entering orbit; infralabials 8/8; preocular single;

postoculars 2; temporals 1+2; dorsal scale rows 17-17-15,

smooth; cloacal entire; ventral scales 183, subcaudals 34,

divided.

Coloration in life: Dorsal head with a grey chevron; dor-

sal surface of body reddish brown with 26 grey, black-

edged bands on body, 4 bands on tail; ventral surface

cream with dark spots (identification after Bourret 1936

and Smith 1943).

Remarks: The female specimen differs from the descrip-

tion of Bourret ( 1936) in having more ventrals ( 183 ver-

sus 1 76), fewer subcaudals (34 vs. 40). Smith ( 1 943) men-

tioned the number of light bands of this subspecies to be

varying from 27-34 on body and 3-4 on tail. Because the

intraspecific taxonomy of Oligodon cinereus remains un-

resolved (David et al. 2011), we provisionally considered

pallidocinctus as a subspecies of this species.

Distribution: In Vietnam, this subspecies is known from

Thua Thien - Hue and Ba Ria - Vung Tau provinces, and

Ho Chi Minh City (“Form IV’’ in Smith 1943). Elsewhere,

Oligodon cinereus has a wide range from India and Chi-

na throughout the Indochina region southward to Thailand

(Nguyen et al. 2009).

Acknowledgements. T.M. Phung is grateful to T. L. T. Nguyen,

H.T. Vu, M. X. Pham, T. M. H. Vu, and T.X. Bui (Ho Chi Minh

City) for assistance in the field. We thank T. H. Khuong (Bu Gia

Map National Park, Binh Phuoc) for providing photos of snakes,

E. Sterling (New York) and K. Koy (Berkeley) for providing the

map. This study was partially supported by the Project TN3/T07

of the National Program Tay Nguyen III. Research of D. T. A.

Tran in Gennany is funded by the Ministry of Education and

Training of Vietnam (MOET, Project 322) and German Academ-

ic Exchange Service (DAAD). Research of T. Q. Nguyen in Ger-

many was funded by the Alexander von Humboldt

Stiftung/Foundation (VIE 114344).

REFERENCES

Bourret R (1936) Les seipents de l’lndochine, II, Catalogue sys-

tematique descriptif. H. Basuyau, Toulouse

David P, Das I, Vogel, G. (201 1) On some taxonomic and nomen-

clatural problems in Indian species of the genus Oligodon

Fitzinger, 1 826 (Squamata: Colubridae). Zootaxa 2799: 1-14

David P, Nguyen QT, Nguyen TT, Jiang K, Chen TB, Teynie A,

Ziegler T (2012) A new species of the genus Oligodon

Fitzinger, 1826 (Squamata: Colubridae) from northern Viet-

nam, southern China and central Laos. Zootaxa 3498: 45-62

Frost DR (2014) Amphibian Species of the World: an Online

Reference. Version 6.0 (23 September 2014). Electronic Data-

base accessible at http://research.amnh.org/herpetology/am-

phibia/index.html. American Museum of Natural History, New

York, USA

Gunther R ( 1994) Eine neue Art der Gattung Gekko (Reptilia,

Squamata, Gekkonidae) aus dem Siiden Vietnams. Zoologi-

scher Anzeiger, Jena 233: 57-67

Ngo TV, Gamble T (2011) Gekko canaensis sp. nov. (Squama-

ta: Gekkonidae), a new gecko from Southern Vietnam.

Zootaxa 2890: 53-64

Nguyen QT, Le DM, Pham TC, Nguyen TT, Bonkowski M,

Ziegler T(2012) A new species of Gracixalus (Amphibia, Anu-

ra, Rhacophoridae) from northern Vietnam. Organisms Diver-

sity & Evolution. Doi 10. 1007/sl3 127-012-01 16-0

Nguyen VS, Ho TC, Nguyen QT (2009) Herpetofauna of Viet-

nam. Edition Chimaira, Frankfurt am Main

Older A (2003) Revision of the genus Opluyopluyne Boulenger,

1903 (Megophryidae) with description of two new species.

Alytes 21: 23-44

Orlov NL, Poyarkov NA, Vassilieva AB, Ananjeva NB, Nguyen

TT, Sang NN, Geisslcr P (2012) Taxonomic notes on rha-

cophorid frogs (Rhacophorinae: Rhacophoridae: Anura) of

southern part of Annamite mountains (Truong Son, Vietnam),

with description of three new species. Russian Journal of Her-

petology, 19 (1): 23-64

Phung MT, Ziegler T (2011 ) Another new Gekko species (Squa-

mata: Gekkonidae) from southern Vietnam. Zootaxa 3129:

51-61

Smith MA (1924) New tree-frogs from Indochina and Malay

peninsula. Proceedings of the Zoological Society of London

94(1): 225-234

Smith MA (1935) The fauna of British India, including Ceylon

and Burma. Reptilia and Amphibia. Vol. 2-Sauria. Taylor and

Francis, London

Smith MA (1943) The fauna of British India, Ceylon and Bur-

ma, including the whole of the Indo-Chinese subregion. Vol.

3-Serpentes. Taylor and Francis, London

Bonn zoological Bulletin 63 (2): 148-156

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Bonn zoological Bulletin 63 (2): 157-172

December 20 1 4

Towards an improved knowledge of Sericini of the Tibetan highland:

new species and records (Coleoptera: Scarabaeidae)

Wan-Gang Liu12, Ming Bai', Xing-Ke Yang' & Dirk Ahrens'

1 Key Laboratoiy of Zoological Svstematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Box 92, No. 1,

Beichen West Road, Chaoyang District, Beijing, 100101, PR. China

2 Graduate School, Chinese Academy of Sciences, Yuquan Road, Shijingshan, Beijing, 100039, P.R. China

1 Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, D-531I3 Bonn, Germany; E-mail: d.ahrens@zfmk.de

Abstract. Nine new species of Sericini are described from Tibetan highlands: Calloserica zhangmuensis Liu & Ahrens

sp. n., Gynaecoserica hani Liu & Ahrens sp. n., G. vigongensis Liu & Ahrens sp. n., Lasioserica beibengana Liu & Ahrens

sp. n., Nepaloserica nielamuensis Liu & Ahrens sp. n., Neoserica (s.l.) chayuensis Liu & Ahrens sp. n., N. (s.l.) xizan-

gensis Liu & Ahrens sp. n., Serica (s.l.) fashengi Liu & Ahrens sp. n., S. (s.l .) yadongensis Liu & Ahrens sp. n. Records

are given for further 25 species, most of them being for the first time recorded from Tibet. Distribution maps of these

new species and records are shown along illustrations of habitus and male copulatory organ of the newly described taxa.

Key words. Beetles, chafers, China, new species.

INTRODUCTION

The taxonomy of the fauna of sericine chafer beetles

(Sericini) of the Himalayas and Tibetan highland was re-

vised and reviewed in detail by Ahrens (2004). Subse-

quently, new data and taxa were added (Ahrens 2005a-e,

2006c, Ahrens & Fabrizi 2009a,b, 201 1, Ahrens & Pa-

cholatko 2005) and a number of phylogenetic studies ex-

plored the diversification and biogeographic patterns of

the Himalayan Sericini fauna (Ahrens 2005d, 2006a, b,d-f,

2007a, b). But we have to consider the eastern regions of

the Himalaya and the adjacent mountain regions of north-

ern Myanmar and southwest China including the Tibetan

highland still to be relatively unexplored. This fact ham-

pers more rigorous hypotheses on the diversification of

the Himalayan fauna.

In this paper, we survey the material hold in Chinese in-

stitutional collections resulting in nine new species and

records for further 25 species, most of them being the first

time recorded from Tibet.

MATERIAL AND METHODS

The terminology and methods used for measurements,

specimen dissection and genital preparation follow

Ahrens (2004). Data from specimens examined is cited in

the text with original label contents given in quotation

marks, multiple labels are separated by a “/”. Descriptions

are based on holotype specimen if not otherwise stated.

Received: 24.03.2014

Accepted: 14.10.2014

The variation of paratypes is given separately. Male gen-

italia were glued to a small pointed card attached to the

specimen. Descriptions and illustrations of new taxa are

based on the holotype specimen, while the variation of oth-

er specimens is given separately under variation. All de-

scriptions and measurements were made under an Olym-

pus SZX 12 microscope, and all genital and habitus illus-

trations were made with a digital camera ( AxioCam HRc)

attached to a stereo microscope (Zeiss Stereo Discovery

V20) and Axio Version 4.8 software. The distribution map

was generated using Q-GIS 2.0. 1 and Adobe Photoshop

CS4.

Type specimens and other examined material are deposit-

ed in the following institutions:

CAU Department of Entomology,

China Agricultural University, Beijing (China);

HBUM Museum of Hebei University, Baoding

(Hebei Prov., China);

IZAS Institute of Zoology, Chinese Academy of

Sciences, Beijing (China);

NKU Nankai University, (Tianjin Prov., China);

NWAFU Northwest A & F University, Yangling

(Shaanxi Prov., China);

ZFMK Zoologisches Forschungsmuseum A. Koenig,

Bonn (Germany).

Corresponding editor: D. Ahrens

158

Wan-Gang Liu et al.

Fig. 1. A D: Calloserica zhangmuensis Liu & Ahrens sp. n. (holotype), E-H: Gynaecoserica hani Liu & Ahrens sp. n. (holo-

type), I-K: G. yigongensis Liu & Ahrens sp. n. (holotype). A, E, I: Aedeagus, left side lateral view; C, G, K: Aedeagus, light side

lateral view; B, F, J: Parameres, dorsal view; D, H, K: Habitus (not to scale). Scale: 0.5 mm.

Bonn zoological Bulletin 63 (2): 157-172

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Towards an improved knowledge of Sericini of the Tibetan highland: new species and records

159

Calloserica zhangmuensis Liu & Ahrens sp. n.

Type material examined. Holotype: S [China] “Zhang-

mu, Xizang, 28.VI.1975, 2200m, leg. Wang Ziqing”

(IZAS). Paratypes: 1 S [China] “Zhangmu, Xizang,

24.VI.1975, 2200m, leg. Wang Ziqing” (ZFMK), 1 $

[China] “Zhangmu, Xizang, 5. VII. 1 975, 2400m, leg.

Huang Fusheng” (IZAS). 1 $ [China] “Zhangmu, Xizang,

28. VI. 1975, 2200m, leg. Wang Ziqing” (IZAS).

Description. Length: 8.0 mm, length of elytra: 6.2 mm,

width: 4.0 mm. Body oblong and dull, dorsal surface, ven-

tral surface and legs reddish brown, elytral intervals, head

and pronotum darker; dorsal surface with moderately long

and very dense setae being directed anteriorly, with a few

longer erect setae on head and pronotum, even intervals

with large spots composed of minute white, scale-like se-

tae.

Labroclypeus short, trapezoidal, widest at base, lateral

margins convex and moderately convergent from base to

broadly rounded anterior angles, anterior margin strongly

concavely sinuate, labrum strongly produced medially, lat-

eral border and ocular canthus producing a distinct obtuse

angle, all margins moderately reflexed; surface flat and

shiny, finely and densely punctate, with several long erect

setae in larger punctures; frontoclypeal suture moderate-

ly impressed and curved, smooth area anterior to eye 1 .5

times as wide as long; ocular canthus long and narrow,

finely and densely punctate, glabrous, terminal seta lack-

ing. Frons moderately densely and finely punctate, punc-

tures shortly and erectly setose, with a few long erect se-

tae setae beside eyes. Eyes small, ratio of diameter/ inte-

rocular width: 0.61. Antenna missing in holotype. Men-

tion elevated and flattened anteriorly.

Pronotum widest at base, lateral margins evenly convex

and convergent anteriorly, anterior angles moderately

rounded, not produced and nearly obsolete, anterior mar-

gin straight, without marginal line, basal margin without

marginal line; surface with very dense and fine punctures

each bearing a yellow fine and erect seta, partly with

longer erect setae, midline not impressed, with a weak

transverse impression behind middle; anterior border fine-

ly setose, setae of lateral margins lacking; hypomeron car-

inate, basal margin of hypomeron distinctly produced ven-

trally. Scutellum subtriangular, apex moderately rounded,

with fine and dense punctures and fine setae, impunctate

on basal midline.

Elytra oblong, widest at middle, striae moderately im-

pressed, with fine and dense punctures; intervals flat, with

evenly distributed, fine and dense punctures, finely and

erectly setose, with numerous longer setae on each inter-

val, even intervals with large spots composed of minute

white, scale-like setae; epipleural edge moderately strong,

ending at strongly rounded external apical angle of ely-

tra, epipleura densely setose, apical border chitinous, with-

out short microtrichomes.

Ventral surface with large and dense punctures, includ-

ing metacoxa with fine adpressed setae. Abdominal ster-

nites finely and densely punctate and finely setose, each

sternite with an indistinct transverse row of coarser punc-

tures each bearing a short, robust seta. Penultimate

abdominal sternite fiat, without transverse elevation

posteriorly. Mesosternum between mesocoxae as wide as

mesofemur, with numerous strong setae on an indistinct

semicircular carina. Ratio of length of metepistemum/

metacoxa: 1/ 1.38. Pygidium moderately convex, with fine

and dense punctures each bearing a fine short seta, with

a few longer setae on apical half, midline narrowly

smooth, apical margin slightly reflexed.

Legs slender and moderately shiny, femora finely and

densely punctate, densely setose. Tarsomeres glabrous and

impunctate dorsally, with sparse, short setae ventrally. Pos-

terior legs missing in holotype. Protibia moderately long,

bidentate, protarsal claws symmetrical, basal tooth of in-

ner claw sharply truncate at apex.

Aedeagus: Fig. 1A-C.

Diagnosis. The new species resembles in genital morphol-

ogy mostly C. poggii Ahrens, 1 995. It differs by the much

shorter basomedian process of parameres, the smaller left

paramere (in lateral view) as well as the apparently short-

er phallobasis.

Variation. Length: 7.4-8. 9 mm, length of elytra: 6. 2-7. 4

mm, width: 3. 6-4. 8 mm. Anterior edge of metafemur

acute, lacking an adjacent serrated line, posterior margin

weakly convex, with numerous strong setae medially,

weakly widened ventrally in apical half but not serrate,

senate dorsally, with short setae. Metatibia moderately

long, widest at apex, ratio width/ length: 1/4.2, dorsal mar-

gin sharply carinate, with two groups of spines, basal

group shortly before middle, apical one at three quarters

of metatibial length, with a few single spines basally; lat-

eral face longitudinally convex, with a few sparse and

coarse punctures, ventral edge finely serrate, with three

robust equidistant setae; medial face impunctate and

smooth, apex concavely truncate interiorly near tarsal ar-

ticulation. Metatarsomeres smooth dorsally, with a strong-

ly serrated ridge ventrally and a lateral longitudinal cari-

na, first metatarsomere as long as following two tar-

someres combined and little less than twice as long as dor-

sal tibial spur. Female: antennal club with three anten-

nomeres, as long as remaining antennomeres combined.

Etymology. The new species is named after the type lo-

cality, Zhangmu.

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Wan-Gang Liu et al.

Gynaecoserica hani Liu & Ahrens sp. n.

Type material examined. Holotype: S [China] “Beibeng,

Motuo, Xizang, 17.V. 1983, 850m, leg. Han Yinheng”

(IZAS). Paratype: 1 $ [China] “Beibeng, Motuo, Xizang,

17.V.1983, 850m, leg. Han Yinheng” (IZAS).

Description. Length: 5.2 mm, length of elytra: 3.9 mm,

width: 3. 1 mm. Body oblong, shiny, including antenna yel-

lowish brown, pronotum, frons and metasternum darker,

dorsal surface except a few setae on head glabrous.

Labroclypeus narrow, widest at base, lateral margins

strongly convex and convergent from base to moderate-

ly rounded anterior angles, lateral margin and ocular can-

thus producing a distinct obtuse angle, anterior margin

weakly sinuate medially, margins weakly reflexed; surface

weakly convex medially and moderately shiny, coarsely

and densely punctate, with several long erect setae behind

anterior margin; frontoclypeal suture moderately im-

pressed and curved, smooth area anterior to eye 1 .5 times

as wide as long; ocular canthus short and moderately nar-

row, impunctate, with a short terminal seta. Frons with

moderately dense punctures, glabrous, only beside each

eye with a single long seta. Eyes small, ratio of diameter/

interocular width: 0.6. Antenna with ten antennomeres,

club with four antennomeres, as long as remaining anten-

nomeres combined. Mentum elevated and flattened ante-

riorly.

Pronotum widest at base, lateral margins nearly straight

and evenly and moderately convergent anteriorly, in an-

terior third slightly convex and more strongly convergent,

anterior angles distinctly produced and sharp, posterior an-

gles blunt, anterior margin convexly produced medially,

with a fine marginal line, basal margin without marginal

line; surface with dense and coarse punctures, on sides

with minute setae, midline not impressed; anterior and lat-

eral borders sparsely setose; hypomeron carinate, basal

margin of hypomeron not produced ventrally. Scutellum

subtriangular, apex moderately rounded, with fine, dense

punctures and minute setae, smooth medially.

Elytra oblong, widest at middle, striae moderately im-

pressed, with fine and dense punctures; intervals weakly

convex, with fine and dense punctures concentrated along

striae; epipleural edge moderately strong, ending at blunt-

ly rounded external apical angle of elytra, epipleura sparse-

ly setose, apical border chitinous, without short microtri-

chomes.

Ventral surface partly dull laterally, with large and dense

punctures, sparsely setose, metacoxa only laterally with

a few fine adpressed setae. Abdominal sternites finely and

densely punctate, nearly glabrous, each stemite with a dis-

tinct transverse row of coarse punctures each bearing a

short robust seta. Mesostemum between mesocoxae as

wide as mesofemur. Ratio of length of metepisternum/

metacoxa: 1/ 1.59. Pygidium shiny, beside apical margin

dull, surface moderately convex, with fine, dense punc-

tures and fine, moderately long setae beside apical mar-

gin, without smooth midline.

Legs moderately long; femora shiny, with two longitu-

dinal rows of setae, finely and sparsely punctate; anteri-

or edge of metafemur acute and without a submarginal ser-

rate line, posterior margin weakly convex, ventral part

smooth and glabrous, only weakly widened in apical half

and not serrate, dorsal margin finely serrate and with fine

short setae. Metatibia slender and moderately long, uni-

formly widened towards apex, ratio width/ length 1/ 2.94,

dorsal margin sharply carinate, longitudinally convex on-

ly basally, with two groups of spines, basal group at one

third, apical group at two thirds of metatibial length, with

a few single, fine spines basally; external face longitudi-

nally convex, with sparse and fine punctures, glabrous;

ventral margin sharp and finely serrate, with three strong

equidistant spines; internal face finely and sparsely punc-

tate, smooth, apex sharply and deeply truncate interiorly

near tarsal articulation. Tarsomeres dorsally glabrous and

impunctate, ventrally with sparse, short setae; metatar-

sonreres ventrally with a strongly serrate ridge, first

metatarsomere little shorter than following two tarsomeres

combined and one third of its length longer than dorsal

tibial spur. Protibia moderately long, bidentate, protarsal

claws symmetrical.

Aedeagus: Fig. 1E-G.

Diagnosis. The new species differs from all other Hi-

malayan Gynaecoserica Brenske, 1896 species by the

shiny body surface and the short antennal club. In exter-

nal morphology it is quite similar to G. compacta Ahrens

& Fabrizi, 2009 differing, however, in the wide lateral

process of phallobasis (lateral view) and the shorter para-

meres.

Variation. Female: Length: 5.1 mm, length of elytra: 3.8

mm, width: 3.1 mm. Antennal club composed of three an-

tennomeres, shorter than remaining antennomeres com-

bined. Eyes as large as in male.

Etymology. The new species is named after its collector,

Han Yinheng.

Gynaecoserica yigongensis Liu & Ahrens sp. n.

Type material examined. Holotype S [China] “Yigong,

Xizang, 1 1. VI. 1978, leg. Li Fasheng” (CAU). Paratype:

I $ [China] “Yigong, Xizang, 11. VI. 1978, leg. Li

Fasheng” (CAU).

Description. Length: 6.2 mm, length of elytra: 5 mm,

width: 3.7 mm. Body oblong and shiny, surface dark red-

dish brown, frons, abdomen and some spots on elytra dark

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Towards an improved knowledge of Sericini of the Tibetan highland: new species and records

161

brown, legs and antenna yellowish brown, dorsal surface

nearly glabrous.

Labroclypeus subrectangular, widest at base, lateral mar-

gins in basal half subparallel, anterior angles broadly

rounded, lateral border and ocular canthus producing a dis-

tinct obtuse angle, anterior margin weakly sinuate medi-

ally, margins weakly reflexed, surface weakly convex me-

dially and moderately shiny, very coarsely and densely

punctate, with a few long erect setae behind anterior mar-

gin; frontoclypeal suture moderately impressed and

curved, smooth area anterior to eye narrow, 1 .25 times as

wide as long; ocular canthus short and slender, glabrous,

without terminal seta. Frons shiny, with coarse and

irregularly dense punctures. Eyes small, ratio of diame-

ter/ interocular width: 0.54. Antenna with ten anten-

nomeres, club in male with four antennomeres, as long as

remaining antennomeres combined, first joint of club

slightly shorter than club. Mentum elevated and flattened

anteriorly.

Pronotum widest at base, lateral margins nearly straight,

in basal half subparallel, evenly moderately convergent an-

teriorly, in anterior third slightly convex and more

strongly convergent, anterior angles distinctly produced

and sharp, posterior angles blunt, anterior margin convex-

ly produced medially, with a fine marginal line, basal mar-

gin without marginal line; surface with dense and mod-

erately coarse punctures, glabrous, midline not im-

pressed; anterior and lateral borders sparsely setose; hy-

pomeron carinate, basal margin of hypomeron not pro-

duced ventrally. Scutellum subtriangular, apex moderately

rounded, with fine, dense punctures and minute setae,

widely smooth medially.

Elytra oblong, widest at middle, striae moderately im-

pressed, with fine and dense punctures; intervals weakly

convex, with fine and dense punctures concentrated along

striae, some punctures produce transverse wrinkles extend-

ing over more than one interval, impuctate portions slight-

ly darker; epipleural edge moderately strong, ending at

strongly rounded external apical angle of elytra, epipleu-

ra densely setose, apical border chitinous, without short

microtrichomes.

Ventral surface dull, with large and dense punctures,

nearly glabrous except a few erect setae on metastemal

disc, metacoxa laterally with a few short adpressed setae.

Abdominal stemites finely and densely punctate, with fine,

short setae, each stemite with an indistinct transverse row

of coarse punctures each bearing a short robust seta. Ul-

timate and penultimate abdominal stemites with a shal-

low, longitudinal median excavation. Mesostemum be-

tween mesocoxae as wide as mesofemur. Ratio of length

of metepistemum/ metacoxa; 1/ 1.26. Pygidium weakly

convex and entirely shiny, with fine and dense punctures,

without smooth midline, with a few longer setae along api-

cal margin.

Legs moderately robust and long; femora shiny, with

two longitudinal rows of setae, finely and sparsely punc-

tate; anterior margin of metafemur sharply carinate, with-

out a submarginal serrate line, posterior margin weakly

convex and glabrous, ventral part smooth and glabrous,

only weakly widened in apical half and not serrate, dor-

sal margin finely serrate and with fine and short setae.

Metatibia slender and moderately long, uniformly widened

toward apex, ratio width/ length: 1/ 3.5, dorsal margin

nearly longitudinally convex, indistinctly carinate only in

apical third, with two groups of spines, basal group at one

third, apical group at two thirds of metatibial length, basal-

ly with a few single, fine spines; external face longitudi-

nally convex, with sparse and fine punctures, glabrous;

ventral margin carinate and senate, with four robust equi-

distant spines; internal face finely and sparsely punctate,

smooth, apex sharply and deeply truncate interiorly near

tarsal articulation. Tarsomeres dorsally glabrous and im-

punctate, ventrally with sparse, short setae; metatar-

someres ventrally with a strongly serrate ridge, first

metatarsomere one third of its length longer than dorsal

tibial spur, subsequent tarsomeres lacking in holotype.

Protibia moderately long, bidentate, protarsal claws sym-

metrical.

Aedeagus: Fig. 1I-K.

Diagnosis. Gynaecoserica yigongensis sp. n. is in exter-

nal shape similar to G. compacta. It differs by the longer

and apically extended lateral apophysis of phallobasis as

well as by the long, fused parameres being bent laterally

(to the right). From G. hani sp. n. it may be distinguished

by the slightly larger size, the shape of parameres as well

as the longer lateral apophysis of phallobasis.

Etymology. The new species is named after the type lo-

cality, Yigong.

Variation. Length: 6. 1-6.2 mm, length of elytra; 4. 6-5.0

mm, width: 3. 6-3. 7 mm. Female: Antennal club composed

of three antennomeres, shorter than remaining anten-

nomeres combined.

Lasioserica beibengana Liu & Ahrens sp. n.

Type material examined. Holotype: $ [China] “Beibeng,

Motuo, Xizang, 20.V.1983, 800m, 850m, 800-900m, leg.

Han Yinheng” (IZAS). Paratypes: 1 S [China] “Beibeng,

Motuo, Xizang, 27.V.1983, 800m, 850m, 800-900m, leg.

Han Yinheng” (IZAS), 1 $ [China] “Beibeng, Motuo,

Xizang, 29. V. 1983, 800m, 850m, 800-900m, leg. Han

Yinheng” (ZFMK).

Description. Length: 5.4 mm, length of elytra: 4.0 mm,

width: 3.5 mm. Body oblong, dorsal surface dark brown,

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Fig. 2. A -D: Lasioserica beibengana Liu & Ahrens sp. n. (holotype), E-H: Nepaloserica nielamuensis Liu & Ahrens sp. n. (holo-

type), I K: Neoserica (s.l.) chayuensis Liu & Ahrens sp. n. (holotype). A, E, I: Aedeagus, left side lateral view; C, G, K: Aedea-

gus, right side lateral view; B, F, J: Parameres, dorsal view; D, H, K: Habitus (not to scale). Scale: 0.5 mm.

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163

antenna and ventral surface brown, dorsal surface dull,

pronotum and head with greenish shine, sparsely setose,

with fine minute and white, scale-like setae on elytra and

pronotum.

Labroclypeus subtrapezoidal, widest at base, lateral mar-

gins in basal half subparallel, anterioly strongly convex

and strongly convergent from middle to broadly rounded

anterior angles, lateral border and ocular canthus produc-

ing a distinct obtuse angle, anterior margin concavely sin-

uate, margins weakly reflexed; surface weakly convex me-

dially and moderately shiny, finely and densely punctate,

with a few long erect setae behind anterior margin; fron-

toclypeal suture weakly impressed and moderately curved,

smooth area anterior to eye 1 .5 times as wide as long; oc-

ular canthus moderately long and narrow, finely and

densely punctate, without terminal seta. Frons dull, fine-

ly and in part densely punctate. Eyes small, ratio of di-

ameter/ interocular width: 0.5. Antenna with ten anten-

nomeres, club in male with four antennomeres, 1 .2 times

as long as remaining antennomeres combined, first joint

of club slightly shorter than club. Mentum elevated and

flattened anteriorly.

Pronotum widest at base, lateral margins subparallel,

nearly straight, distinctly and widely sinuate before pos-

terior angles, moderately curved anteriorly and convergent

to acute, distinctly produced anterior angles, posterior an-

gles nearly right-angled, anterior margin nearly straight,

with a fine marginal line, basal margin without marginal

line; surface with dense and coarse punctures each bear-

ing either an adpressed, minute seta or a moderately long,

white scale, with a transverse impression behind middle;

anterior and lateral borders sparsely setose; hypomeron

carinate, basal margin of hypomeron weakly produced

ventrally. Scutellum subtriangular, apex moderately

rounded, with fine and dense punctures and moderate se-

tae, smooth on basal midline.

Elytra oblong, widest shortly behind middle, striae

moderately impressed, with fine and dense punctures; in-

tervals moderately convex, with fine and dense punctures

concentrated along striae, with sparse, short white scales

or minute setae; epipleural edge moderately strong, end-

ing at strongly rounded external apical angle of elytra, epi-

pleura densely setose, apical border narrowly membrane-

ous, with very short microtrichomes.

Ventral surface dull, with large and dense punctures,

sparsely setose, metacoxa only laterally with a few fine,

adpressed setae. Abdominal sternites finely and densely

punctate and minutely setose, each sternite with a distinct

transverse row of coarse punctures each bearing a short,

robust seta. Penultimate abdominal sternite without tuber-

cles but slightly transversely elevated medially. Mesoster-

num between mesocoxae as wide as mesofemur. Ratio of

length of metepisternum/ metacoxa: 1/ 1.38. Pygidium

moderately convex and dull, with coarse, dense punctures

and fine, white setae, with wide smooth midline.

Legs moderately slender and long; femora dull on ven-

tral face, with two longitudinal rows of setae, finely and

sparsely punctate; anterior edge of metafemur acute, with

an adjacent serrated line, ventrally weakly widened ven-

trally in apical half but not serrate, dorsally serrate. Tar-

someres glabrous, finely punctate dorsally, with sparse,

short setae ventrally. Metatibia including tarsi missing in

holotype. Protibia moderately long, bidentate. Protar-

someres missing in holotype. Female unknown.

Aedeagus: Fig. 2A-C.

Diagnosis. The shape of parameres is similar to those of

L. turaensis Ahrens, 2000. It differs significantly by the

at left side ventrally extended phallobasis, whose interi-

or rim of dorsoapical median sinuation produces a sharply

pointed dorsal process, which is unique trait among all oth-

er so far known Lasioserica Brenske, 1896 species.

Variation. Length: 5. 0-5. 6 mm, length of elytra: 3. 8-3. 9

mm, width: 3. 0-3. 2 mm. Protarsomeres short and robust,

claws slightly asymmetrical, basal tooth of interior pro-

tarsal claw widened and bluntly truncate.

Etymology. The new species is named after the type lo-

cality, Beibeng.

Nepaloserica nielantuensis Liu & Ahrens sp. n.

Type material examined. Flolotype S “Zhangmu, Niela-

mu, Xizang, 23. IV. 1974, 2250m, leg. Zhang Xuezhong”

(IZAS). Paratypes: 3 S “Zhangmu, Nielamu, Xizang,

23. IV. 1974, 2250m, leg. Zhang Xuezhong” (IZAS,

ZFMK), 1 <$ “Zhangmu, Nielamu, Xizang, 10. V. 1966,

2200m, leg. Wang Shuyong” (IZAS).

Description. Length: 9.9 mm, length of elytra: 8.2 mm,

width: 5.2 mm. Body oblong, dorsal and ventral surface

reddish brown and dull, antenna brown, except a few se-

tae on head sparsely setose.

Labroclypeus narrow and nearly square, widest at base,

lateral margins weakly convex and convergent from base

to moderately rounded anterior angles, lateral border and

ocular canthus producing a distinct obtuse angle, anterior

margin moderately sinuate but strongly retlexed; surface

nearly flat and shiny, finely and densely punctate, with a

few long erect setae in coarser punctures anteriorly; fron-

toclypeal suture weakly impressed and moderately curved,

smooth area anterior to eye 1 .5 times as wide as long; oc-

ular canthus long and slender, finely and densely punc-

tate, with one to two terminal setae. Frons dull, finely and

densely punctate, punctures vanish under dull toment, with

a few long setae on posterior frons and beside eyes, oth-

erwise only with minute setae. Eyes moderately large, ra-

tio of diameter/ interocular width: 0.66. Antenna with ten

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164

Wan-Gang Liu et al.

antennomeres, club with seven antennomeres, 1.5 times

as long as remaining antennomeres combined, first joint

of club slightly shorter than club. Mentum elevated and

flattened anteriorly.

Pronotum widest at middle, lateral margins in basal half

subparallel, moderately curved and convergent anterior-

ly, anterior angles acute, distinctly produced, posterior an-

gles blunt, anterior margin weakly produced medially, with

a fine marginal line, basal margin without marginal line;

surface with dense and coarse punctures bearing often a

minute, adpressed seta; anterior and lateral borders

sparsely setose; hypomeron carinate, basal margin of hy-

pomeron not produced. Scutellum subtriangular, apex

moderately rounded, with coarse and dense punctures.

Elytra oblong, widest at posterior third, striae distinct-

ly impressed, with fine and dense punctures; intervals

moderately convex, with fine and dense punctures con-

centrated along striae, glabrous except minute setae and

a few short setae on lateral intervals; epipleural edge mod-

erately strong, ending at strongly rounded external apical

angle of elytra, epipleura densely setose, apical border

membraneous, with a fine rim of short microtrichomes.

Ventral surface with large and dense punctures, sparse-

ly setose, on metastemum with moderately dense and long

setae; metacoxa only laterally with a few fine adpressed

setae. Abdominal stemites finely and densely punctate,

with fine, short setae medially on basal stemites and

minute ones only laterally on other stemites, each stern-

ite with a distinct transverse row of coarse punctures each

bearing a short robust seta, abdominal stemites without

any elevation. Mesosternum between mesocoxae as wide

as mesofemur. Ratio of length of metepistemum/ meta-

coxa: 1/ 1.51. Pygidium moderately evenly convex, with

coarse and dense punctures and a few moderately long se-

tae on apical half; without smooth midline.

Legs slender and long; femora with two longitudinal

rows of setae, finely and sparsely punctate; anterior edge

of metafemur acute with an adjacent serrated line,

ventrally weakly widened ventrally in apical half but not

serrate, dorsal ly serrate. Metatibia moderately wide,

widened at base, ratio width/ length: 1/3.9, dorsal margin

sharply carinate, with two groups of spines, basal group

at one third, apical one at three quarters of metatibial

length, with a few single spines basally; lateral face lon-

gitudinally convex, with very sparse and fine punctures,

along the middle nearly impunctate, ventral margin ser-

rate, with three robust equidistant spines; medial face im-

punctate and smooth, apex interiorly near tarsal articula-

tion shallowly concavely truncate. Tarsomeres glabrous

and finely punctate dorsally, with sparse, short setae ven-

trally, metatarsomeres with a serrated ridge ventrally, and

a sharp carina immediately beside it, first metatarsomere

slightly shorter than following two tarsomeres combined

and slightly longer than the dorsal tibial spur. Protibia

short, bidentate, protarsal claws broken. Female unknown.

Aedeagus: Fig. 2 E-G.

Diagnosis. The new species is in shape of parameres rather

similar to N. procera procera Frey, 1965 and N. procera

rufescens Frey, 1965. From both it differs by the setose

frons, the lack of elevations on penultimate stemites, as

well as the broadly inserted basal lobe of right paramere,

which is over half of its length fused with the paramere.

Etymology. The new species is named after the type lo-

cality, Nielamu.

Variation. Length: 9.3-11.3 mm, length of elytra: 7.4-8. 8

mm, width: 5. 3-5. 9 mm. Protarsal claws symmetrical,

basal tooth of inner claw sharply truncate at apex.

Neoserica (s.l.) chayuensis Liu & Ahrens sp. n.

Type material examined. Holotype <$ [China] “Xi-

achayu, Chayu, Xizang, 12-1 3. VII. 2005, leg. Shi Aimin’’

(HBUM). Paratypes: 1 $ [China] “Xiachayu, Chayu,

Xizang, 12-13.VII.2005, leg. Shi Aimin’’ (HBUM), 2 S

[China] “Xiachayu, Chayu, Xizang, 15. VII. 2005, leg. Shi

Aimin’’ (HBUM, ZFMK).

Diagnosis. Length: 8.9 mm, length of elytra: 6.8 mm,

width: 5.4 mm. Body oval, dorsal and ventral surface dull

and dark brown, legs reddish brown, antenna yellow, dor-

sal surface with yellowish, short and fine, dense setae.

Labroclypeus trapezoidal, widest at base, lateral mar-

gins moderately convex and convergent from base to

broadly rounded anterior angles, anterior margin weakly

sinuate and moderately re flexed, lateral border and ocu-

lar canthus producing a distinct obtuse angle, surface flat

and shiny, at base dull, finely and densely punctate, with

numerous long, erect setae; frontoclypeal suture weakly

incised and moderately curved, smooth area anterior to eye

1 .5 times as wide as long; ocular canthus moderately long

and slender, impunctate, with one or two terminal setae.

Frons finely and densely punctate, completely dull, with

dense and long erect setae. Eyes small, ratio of diameter/

interocular width: 0.56. Antenna with ten antennomeres,

club in male with six antennomeres, as long as remain-

ing antennomeres combined, first joint of club slightly

shorter than club. Mentum elevated and flattened anteri-

orly.

Pronotum widest at base, lateral margins weakly

curved and evenly convergent anteriorly, anterior angles

acute, moderately produced, posterior angles wekaly

rounded, anterior margin moderately convex, with distinct

and fine marginal line, basal margin without marginal line;

surface with dense and coarse punctures each bearing a

yellow, semi-erect, short seta being bent slight posterior-

ly, disc without any impression; anterior and lateral bor-

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165

ders sparsely setose, setae slightly longer than those of the

disc; hypomeron carinate, basal margin of hypomeron not

produced. Scutellum subtriangular, apex moderately

rounded, with fine and dense punctures and fine setae.

Elytra oblong, widest shortly behind middle, striae

weakly impressed, finely and densely punctate; intervals

weakly convex, with evenly distributed, fine and dense

punctures, each bearing a fine, semi-erect, yellow seta, odd

intervals with a few longer, erect setae; epipleural edge

moderately strong, ending at strongly rounded external

apical angle of elytra, epipleura densely setose, apical bor-

der finely membraneous, with a fine rim of very short mi-

crotrichomes.

Ventral surface dull, with large, dense punctures and

dense, sparse setae, metacoxa glabrous, with a few robust

setae laterally only. Abdominal stemites finely and

densely punctate, with fine, dense, short setae, each ster-

nite with a distinct transverse row of coarse punctures each

bearing a short robust seta. Mesostemum between meso-

coxae as wide as mesofemur. Ratio of length of metepis-

temum/ metacoxa: 1/ 1 .67. Pygidium weakly convex, with

coarse, dense punctures and shorter and longer, fine se-

tae, without smooth midline.

Legs moderately slender and long, moderately shiny,

femora with fine and dense punctures and sparsely setose;

anterior edge of metafemur acute, lacking an adjacent ser-

rated line, ventral posterior margin weakly convex,

weakly widened ventrally in apical half but not serrate,

with a few fine setae, dorsal posterior margin serrate, with

a few fine setae. Metatibia long and moderately wide,

widest at apex, ratio width/ length: 1/ 4.2, dorsal margin

sharply carinate, with two groups of spines, basal group

shortly before middle, apical one shortly behind three

quarters of metatibial length, with a few single spines

basally on a short and blunt longitudinal elevation; later-

al face longitudinally convex, with sparse and large, shal-

low punctures; ventral margin senate, with three nearly

equidistant robust setae; medial face impunctate and

smooth, apex shallowly sinuate interiorly near tarsal ar-

ticulation. Tarsomeres glabrous and finely and superficial-

ly punctate dorsally, with sparse, short setae ventrally,

metatarsomeres with a strongly serrated ridge ventrally

and a fine longitudinal carina immediately beside it, first

metatarsomere slightly longer than following two

tarsomeres combined and one third of its length longer

than dorsal tibial spur. Protibia short, bidentate, protarsal

claws symmetrical.

Aedeagus: Fig. 2I-K.

Diagnosis. The new species resembles in external and gen-

ital morphology to Neoserica barberi (Sharp, 1903) from

southern India; it shares the subsymmetrical male geni-

talia, the shape of labroclypeus and legs, as well as the

configuration of antennal club. From all species of the N.

barberi group the new species differs by the dense pilos-

ity of the dorsal body surface, as well as by the basal lobe

present at each paramere.

Variation. Length: 8.9-10.5 mm, length of elytra: 6.8-8

mm, width: 5. 4-6. 6 mm. Female. Eyes smaller, ratio of

diameter/ interocular width: 0.51, antennal club composed

of five antennomeres, first joint of club subequal to one

sixth of club length.

Etymology. The new species is named after the type lo-

cality, Chayu.

Remarks. Head and pronotum of one paratype (“Xi-

achayu, Chayu, Xizang, 1 5. VII. 2005, leg. Shi Aimin’’) are

missing.

Neoserica (s.l.) xizangensis Liu & Ahrens sp. n.

Type material examined. Holotype: S [China] “Lang-

cun, Lang County, Xizang, 8.VI.1986, 3080m, No. 581”

(IZAS). Paratype: 1 $ [China] “Langxian County,

Xizang, 5.VII.I981, leg. Chen Tailu” (ZFMK).

Description. Length: 8.0 mm, length of elytra: 5.8 mm,

width: 4.0 mm. Body oblong, including legs yellowish

brown, head slightly darker, antenna yellow, dorsal sur-

face shiny, nearly glabrous.

Labroclypeus trapezoidal, widest at base, lateral margins

moderately convex and convergent from base to moder-

ately rounded anterior angles, anterior margin weakly

sinuate medially, margins moderately reflexed, lateral mar-

gin and ocular canthus producing a distinct obtuse angle;

surface fiat, coarsely and very densely punctate, distance

between punctures less their diameter, with a few short

setae behind anterior margin; frontoclypeal suture weakly

impressed and moderately curved, smooth area anterior

to eye twice as wide as long; ocular canthus moderately

long, finely and densely punctate, without terminal seta.

Frons densely and coarsely punctate, entirely glabrous.

Eyes moderately large, ratio of diameter/ interocular

width: 0.69. Antenna with ten antennomeres, club in male

with four antennomeres, as long as remaining anten-

nomeres combined, first joint of club slightly shorter than

club. Mentum elevated and flattened anteriorly.

Pronotum widest at middle, lateral margins weakly but

evenly curved and convergent anteriorly and posteriorly,

anterior angles right-angled, weakly produced, posterior

angles blunt, anterior margin convexly produced medial-

ly, with distinct marginal line, basal margin without mar-

ginal line; surface with very dense and coarse punctures,

without impressions; anterior and lateral borders sparse-

ly setose; hypomeron only medially slightly carinate, ca-

rina completely reduced laterally, basal margin of hy-

pomeron not produced. Scutellum subtriangular, apex

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Fig. 3. A-D: Neoserica (s.l.) xizangensis Liu & Ahrens sp. n. (holotype), E- I I: Serica (s.l ,)fashengi Liu & Ahrens sp. n. (holo-

type). I K: S. (s.l.) yadongensis Liu & Ahrens sp. n. (holotype). A, E, I: Aedeagus, left side lateral view; C, G, K: Aedeagus, right

side lateral view; B, F, J: Parameres, dorsal view; D, H, K: Habitus (not to scale). Scale: 0.5 mm.

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167

moderately rounded, with coarse and dense punctures,

glabrous.

Elytra oblong, widest at middle, striae weakly im-

pressed, with fine and dense punctures; intervals weakly

convex, with fine and dense punctures concentrated along

striae, punctures with very minute setae, odd intervals with

a few short yellow setae; epipleural edge robust, ending

at strongly rounded external apical angle of elytra, epipleu-

ra densely setose, apical border chitinous, without short

microtrichomes.

Ventral surface dull or weakly shiny, with large and

dense punctures, sparsely setose except on metasternal

disc. Mesostemum between mesocoxae as wide as

mesofemur, with numerous strong setae on an indistinct

semicircular carina. Ratio of length of metepistemum/

metacoxa: 1/ 1.27. Abdominal stemites finely and dense-

ly punctate, with fine, moderately dense setae on basal

stemites, otherwise sparsely setose, each stemite with an

indistinct transverse row of coarse punctures each bear-

ing a short robust seta. Pygidium weakly convex and

shiny, with fine, moderately dense punctures and fine se-

tae, without smooth midline.

Legs long and slender, shiny, femora finely and sparse-

ly punctate, sparsely setose; anterior edge of metafemur

acute, lacking an adjacent serrated line, posterior ventral

margin weakly convex, weakly widened ventrally in api-

cal half but not senate, posterior dorsal margin smooth,

densely and shortly setose. Metatibia slender and very

long, widest at apex, ratio width/ length: 1/ 6, dorsal mar-

gin longitudinally convex, not carinate, with two groups

of spines, basal group at one quarter, apical one at two

thirds of metatibial length, with a few single spines basal-

ly; lateral face longitudinally convex, with dense and

coarse, longitudinally impressed punctures, glabrous; ven-

tral edge carinate and finely serrate, with four equidistant

robust spines, between each of them with a single short

seta; medial face densely punctate and with long, dense

setae, apex sharply truncate interiorly near tarsal articu-

lation. Tarsomeres glabrous and finely punctate dorsally,

with moderately dense, short setae ventrally; metatar-

someres with a strongly serrated ridge ventrally and a fine

longitudinal carina immediately beside it, first metatar-

somere slightly longer than following tarsomere and as

long as dorsal tibial spur. Protibia long, bidentate, protar-

someres missing in holotype.

Aedeagus: Fig. 3A-C.

Diagnosis. At the moment it is difficult to nominate a clos-

er relative of this new species. The reduced carina of hy-

pomeron is found in Asia only in Tetraserica Ahrens, 2004

and Serica (s. str.) MacLeay, 1819 [from whose N. xizan-

gensis differs in shape of aedeagus], while the shape of

legs resemble somewhat those of Sericania Motschulsky,

1 860, however, parameres are simple and do not have

basal lobes as all other Sericania species.

Etymology. The new species is named after its occurrence

in Xizang.

Variation. Length: 7. 3-8.0 mm, length of elytra: 5. 4-5. 8

mm, width: 3. 5-4.0 mm. Protarsomeres long and slender,

claws symmetrical, basal teeth of protarsal claws sharply

pointed.

Serica (s.l. ) fashengi Liu & Ahrens sp. n.

Type material examined. Holotype: <$ [China] “Yigong,

Xizang, 29. VII. 1978, 2300m, leg. Li Fasheng” (CAU).

Paratypes: 1 [China] “Yigong, Xizang, 25.VII.1976,

2050m, leg. Li Fasheng” (ZFMK).

Description. Length: 9.2 mm, length of elytra: 7.7 mm,

width: 5.6 mm. Body oblong-oval including legs dark

reddish brown, dull, antenna yellowish brown, dorsal sur-

face nearly glabrous, head and pronotum moderately

shiny.

Labroclypeus subtrapezoidal, widest at base, lateral mar-

gins nearly straight and strongly convergent to broadly

rounded anterior angles, anterior margin weakly sinuate,

margins weakly reflexed, lateral border and ocular can-

thus producing a distinct obtuse angle, surface fiat, coarse-

ly and very densely punctate, without setae; frontoclypeal

suture weakly impressed and moderately curved, smooth

area anterior to eyel .5 times as wide as long; ocular can-

thus short and wide, finely and densely punctate, with a

single, long terminal seta. Frons dull, coarsely and very

densely punctate, with a few long setae beside eyes. Eyes

moderately large, ratio of diameter/ interocular width:

0.66. Antenna with nine antennomeres, club with three an-

tennomeres, 5 times as long as remaining antennomeres

combined, club strongly reflexed. Mentum elevated and

Battened anteriorly.

Pronotum widest at base, lateral margins nearly straight

and evenly convergent anteriorly, in anterior third

moderately curved and more strongly convergent, anteri-

or angles acute, distinctly produced, posterior angles

moderately rounded, anterior margin weakly convex me-

dially, with a fine marginal line, basal margin without mar-

ginal line; surface with dense and coarse punctures,

glabrous, without impressions; anterior and lateral borders

sparsely setose; hypomeron carinate, basal margin of hy-

pomeron not produced. Scutellum subtriangular, apex

moderately rounded, with dense and coarse punctures,

glabrous.

Elytra oblong, widest at middle, striae weakly im-

pressed, with fine and dense punctures; intervals weakly

convex, with fine and dense punctures concentrated along

striae, except a few minute setae on lateral intervals

glabrous; epipleural edge strong, ending at strongly

rounded external apical angle of elytra, epipleura dense-

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Wan-Gang Liu et al.

ly setose, apical border membraneous, with short microtri-

chomes.

Ventral surface with large and dense punctures, except

a few erect setae on metasternal disc nearly glabrous;

metacoxa only laterally with a few robust, adpressed se-

tae. Mesosternum between mesocoxae as wide as mesofe-

mur, with numerous strong setae on an indistinct semicir-

cular carina. Ratio of length of metepistemum/ metacoxa:

1/ 1 .35. Abdominal sternites finely and densely punctate,

nearly glabrous, each stemite with a distinct transverse row

of coarse punctures each bearing a short robust seta. Py-

gidium weakly convex, with fine, dense punctures and a

few fine setae along apical margin, without smooth mid-

line.

Legs moderately long; femora with two longitudinal

rows of setae, finely and sparsely punctate; anterior edge

of metafemur acute with an adjacent, but partly interrupt-

ed serrated line, posterior ventral margin nearly straight,

weakly widened in apical half and not serrate, posterior

dorsal margin indistinctly serrate, densely setose. Metat-

ibia moderately wide and long, slightly widened behind

middle, ratio width/ length: 1/ 3.7, dorsal margin weakly

carinate, with two groups of spines, basal group shortly

before middle, apical one at three quarters of metatibial

length, with a few single setae in serrated punctures basal-

ly; lateral face longitudinally convex, with fine and sparse

punctures; ventral margin serrate; medial face impunctate

and smooth, apex sharply truncate interiorly near tarsal ar-

ticulation. Tarsomeres glabrous and impunctate dorsally,

with sparse, short setae ventrally, metatarsomeres with a

strongly serrated ridge ventrally, and longitudinal carina

immediately beside it, first metatarsomere as long as fol-

lowing two tarsomeres combined and one third of its

length longer than tibial spur. Protibia moderately long,

bidentate, external edge smooth, claws symmetrical.

Aedeagus: Fig. 3 E-G.

Diagnosis. The new species belongs to the Serica segre-

gatei Arrow, 1946 group. It differs from S.falcifera Ahrens

& Fabrizi, 2009 by the wider basal lobe of the right para-

mere, the sharply hooked dorsal lobe of left paramere, and

the longer antennal club. It is hard to say with a single

known specimen whether also the number of anten-

nomeres constitute a species-specific diagnostic feature to

distinguish from S. segregata and falcifera (both having

an antenna composed of ten antennomeres instead of nine).

Etymology. The new species is named after the collector

of the species, Li Fasheng.

Variation. Length: 9.2-10 mm, length of elytra: 7. 5-7. 7

mm, width: 5. 3-5. 6 mm.

Serica (s.l.) yadongensis Liu & Ahrens sp. n.

Type material examined. Holotype: S [China] “Yadong,

Xizang, 1. VII. 2004, leg. Ba Yibin, Shi Aimin’’ (HBUM).

Paratype: 1 S [China] “Xiasima, Yadong, Xizang,

14. VI. 1983, leg. Wang Jia, Ci Ren, No. 96-2” (IZAS).

Description. Length: 10 mm, length of elytra: 7.4 mm,

width: 4.6 mm. Body oblong, dark reddish brown, head

dark brown, antenna yellowish brown, dorsal surface dull,

elytra with irregular dark spots; head and pronotum with

long erect setae, elytra with minute to short white ad-

pressed setae.

Labroclypeus narrowly subtrapezoidal, widest at base, lat-

eral margins moderately convergent from base to moder-

ately rounded anterior angles, anterior margin weakly

sinuate, margins strongly reflexed, lateral border and oc-

ular canthus producing a distinct obtuse angle; surface flat

and moderately shiny, very coarsely and densely punctate,

larger punctures each bearing a long erect setae inter-

spersed with fine, glabrous punctures; frontoclypeal su-

ture moderately impressed and curved, smooth area an-

terior to eye 1.5 times as wide as long; ocular canthus

moderate long and robust, finely and densely punctate,

with a long terminal seta. Frons dull, finely and densely

punctate, with numerous long, erect setae. Eyes

moderately large, ratio of diameter/ interocular width: 0.7.

Antenna with ten antennomeres, club with three anten-

nomeres, 1.5 times as long as remaining antennomeres

combined, club slightly reflexed. Mentum elevated and

flattened anteriorly.

Pronotum widest at base, lateral margins in basal half

subparallel and slightly concavely sinuate, moderately

curved and strongly convergent in anterior third, anterior

angles bluntly rounded, moderately produced, anterior

margin nearly straight, with a fine marginal line, posteri-

or angles moderately round, basal margin without margin-

al line; surface with dense and fine punctures, midline im-

punctate, with several long erect setae bent posteriorly,

otherwise laterally with short, white or minute setae; an-

terior and lateral borders densely setose; hypomeron car-

inate, basal margin of hypomeron not produced. Scutel-

lum subtriangular, apex moderately rounded, with fine and

very dense punctures, glabrous, impunctate on midline.

Elytra oblong, widest at middle, striae moderately im-

pressed, with fine and dense punctures, reddish brown; in-

tervals moderately convex, second interval posteriorly

strongly widened, intervals with irregular, fine and dense

punctures, with fine, minute or short, scale-like, adpressed

setae concentrated along striae, punctation partly interrupt-

ed by large and dark impunctate spots; epipleural edge

moderately strong, ending at strongly rounded external

apical angle of elytra, epipleura densely setose, apical bor-

der membraneous, with a very fine rim of short microtri-

chomes.

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169

Ventral surface with large and dense punctures, dense-

ly setose; metacoxa glabrous, only laterally with a few fine

adpressed setae. Mesostemum between mesocoxae as

wide as mesofemur. Ratio of length of metepistemum/

metacoxa: 1/ 1.5. Abdominal stern ites finely and moder-

ately densely punctate, punctures with minute setae, each

sternite with an indistinct transverse row of coarse punc-

tures each bearing a short, robust seta. Pygidium strong-

ly convex, with coarse, moderately dense punctures and

fine, long setae, midline smooth.

Legs slender and long; femora with two longitudinal

rows of setae, finely and sparsely punctate; anterior edge

of shiny metafemur acute, without an adjacent serrated

line, ventral posterior margin nearly straight, weakly

widened in apical half ventrally, not serrate, dorsal pos-

terior margin serrate and densely setose. Metatibia slen-

der and long, abruptly widened at apex, ratio width/ length:

1/ 4.2, dorsal margin longitudinally convex, with two

groups of spines, basal group at one third, apical one at

three quarters of metatibial length, with a few single se-

tae basally; lateral face longitudinally convex, with sparse

and moderately coarse punctures, some of them longitu-

dinally impressed, glabrous; ventral margin serrate, with

four equidistant spines; medial face impunctate and

smooth, apex sharply truncate interiorly near tarsal artic-

ulation. Tarsomeres glabrous and impunctate dorsally, with

sparse, short setae ventrally, metatarsomeres with a strong-

ly serrated ridge ventrally and a longitudinal carina im-

mediately beside it; first metatarsomere distinctly short-

er than following two tarsomeres combined and twice as

long as dorsal tibial spur. Protibia moderately long, biden-

tate, external edge smooth, claws symmetrical. Female un-

known.

Aedeagus: Fig. 3I-K.

Diagnosis. The new species differs from Serica erectose-

tosa Ahrens, 1999 by having nearly straight parameres

(lateral view), and from all other species of S. erectose-

tosa group by having long erect setae on head and prono-

tum, parameres are more strongly asymmetric as in S. kar-

naliensis Ahrens, 1999.

Etymology. The new species is named after the type lo-

cality, Yadong.

Variation. Length: 9.0-10.0 mm, length of elytra: 7. 0-7.4

mm, width: 4.6-5. 1 mm.

ADDITIONAL MATERIAL

Serica (s. str.) guidoi Ahrens, 1999

Additional material examined. I $ “Zhangmukou,

Nielamu, Xizang, 12.V. 1974, 2250m, leg. Zhang

Xuezhong” (IZAS).

Serica (s. str.) khajiaris Mittal, 1988

Additional material examined. 2 S “Yigong, Bomi,

Xizang, 14,22. VIII. 1983, 2300m, leg. Flan Yinheng”

(IZAS).

Serica (s. str.) kingdom Ahrens, 1999

Additional material examined. 1 $ “Qiujiaba, Wen

County, Gansu, 22. VII. 1999, 2350m, leg. Yao Jian”

(IZAS), 3 $ “Qiujiaba, Wen County, Gansu, 21. VI 1. 1999,

2350m, leg. He Tongli” (IZAS), 1 $ “Hailuo Gou, Moxi,

Luding, Sichuan, 9. VIII. 2004, 3000m, leg. Wan Xia,

Zhang Yong” (IZAS), 1 $ “Wolongguan Gou, Wo-

longzhen, Sichuan, 20. VII. 2004, 2150m, leg. Zhang Yong”

(IZAS), I S “Yaojiaping, Lushui, Yunnan, 2.VI.1981,

2500m, light trap, leg. Wang Shuyong” (IZAS), 1 <$ “San-

shenggou, Wolong, Sichuan, 8. VIII. 1983, 2500m, leg. Niu

Chunlai” (IZAS), 1 S “Chayu, Xizang, 22. VII. 2009, leg.

Ren Guodong etc.” (HBUM), 1 $ “Hailuogou, Moxi,

Luding, Sichuan, 8. VIII. 2004, 3000m, leg. Wan Xia”

(IZAS).

Serica (s. str.) montreuili Ahrens, 2005

Additional material examined. 1 <$ “Maxionggou, Xi-

angcheng, Sichuan, 25.VI.1982, 3800m, leg. Wang Shuy-

ong” (IZAS), 1 $ “Haizigou, Mts. Siguniangshan,

Sichuan, 29.VII.2004, 3400-3860m, leg. Bai Ming”

(IZAS), 1 $ “Shuangqiaogou, Mts. Siguniangshan,

Sichuan, 28. VII. 2004, 3840m, leg. Bai Ming” (IZAS), 1

<$ “Jiangda, Xizang, 28.VII.1976, 3400m, leg. Han Yin-

heng” (IZAS), 1 $ “Haitong, Mangkang, Xizang, 7, 8. VI-

II. 1983, 2950-3050m, leg. Lin Zai” (IZAS), 1 $ “Gelong,

Yushu, Qinghai, 10, 12. VII. 1964, 3700m, leg. Wang

Shuyong” (IZAS), 1 <S “Zhongreniao, Xiangcheng,

Sichuan, 17.VII.1982, 4100m, leg. Zhang Xuezhong”

(IZAS).

Serica (s. str.) nepalensis (Frey, 1969)

Additional material examined. 1 S “Zhangmu, Niela-

mu, 21.IV. 1974, 2250m, leg. Zhang Xuezhong” (IZAS),

1 $ “Jitang Youxi, Chaya, Xizang, 16. VIII. 1976, leg. Han

Yinheng” (IZAS), l <$ “Youyi Bridge, Nielamu, Xizang,

21.IV.1974, 1668m, leg. Zhang Xuezhong” (IZAS), 1 $

“Zhangmu, Nielamu, Xizang, 21.IV.1974, leg. Zhang

Xuezhong” (IZAS), 1 S, 1 $ “Zhangmukouan, Xizang,

28.VI.1975, 2250m, leg. Zhang Xuezhong” (IZAS).

Serica (s. str.) thibetana Brenske, 1897

Additional material examined. I $ “Youyi Brigde,

Nielamu, Xizang, 6. V. 1974, 1680m, light trap, leg.

Zhang Xuezhong” (IZAS), I <$ “Yadong, Xizang, 29. VI-

11.1980, 2700m, light trap, leg. Li Aihua” (IZAS), 1 S “Xi-

asima, Yadong, Xizang, 29. V. 1983, leg. Wang Jia, Ci Ren,

No. 98-3”(IZAS), 1 S “Yadong, Xizang, 29.V.1983,

2700m, light trap, leg. Li Aihua” (IZAS).

Bonn zoological Bulletin 63 (2): 157-172

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170

Wan-Gang Liu et al.

O karnaliensis 90

-(- longefoliata

+ montreuili

□ nepalensis

• pygidialis ssp annapurnae

A thibetana

95 00

♦

35.00

30.00

O archolabrata O pacholatkoi

m beibengana a procera

♦ chayuensis ☆ rutilans

A ganeshi O sladeni

V lenangensis □ xizangensis

★ nielamuensis

30.00

0-1500

1500-2000

2000-3000

® 30004000

1 Wm 4000-9000

Fig. 4. Distribution and new records of: A: Calloserica bertiae, C. capillata , C. zhangmuensis, Gynaecoserica hani, G. lohiten-

si.s, G. yigongensis, Maladera modestula, M. siniaevi, Micmserica pruinosa. Mi. steelei , Pachyserica bituberculata. B: Serica (s.l.)

erectosetosa , S. (s.l .) fashengi, Serica (s.str.) guidoi , S. (s.str.) kingdom, S. (s.str.) yadongensis. C: S. (s.l.) karnaliensis, S. (s.str.)

montreuili, S. (s.str.) nepalensis, S. (s.str.) thibetana, Oxyserica longefoliata, O. pygidialis ssp. annapurnae. I): Lasioserica beiben-

gana, L. pacholatkoi, Neoserica lenangensis, N. sladeni, N. chayuensis, N. rutilans, N. xizangensis , Nepaloserica archolabrata,

Ne. ganeshi, Ne. nielamuensis , Ne. procera.

Bonn zoological Bulletin 63 (2): 157-172

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Towards an improved knowledge of Sericini of the Tibetan highland: new species and records

171

Serica (s. 1.) kamaliensis Ahrens, 1999

Additional material examined. 1 $ “Yadong, Xizang,

1. VII. 2004, leg. Ba Yibin, Shi Aimin’’ (HBUM).

Serica (s. I.) erectosetosa Ahrens, 1999

Additional material examined. I $ “Zhangmu, Niela-

mu, Xizang, 12.V.1974, 2250m, leg. Zhang Xuezhong”

(IZAS).

Pachyserica bituberculata Ahrens, 2006

Additional material examined. \<$ “Motuo, Xizang,

14.VIII.2003, 1100m, leg. Ren Guodong”(HBUM), 1 <$

“Maniweng, Motuo, Xizang, 28.VIII.1974, 930m, leg.

Huang Fusheng” (IZAS).

Calloserica bertiae Ahrens, 2000

Additional material examined. 1 <$ “Xiasima, Yadong,

Xizang, 8. VII. 1 983, leg. Wang Jia, Ci Ren” (IZAS).

Calloserica capillata Ahrens, 2005

Additional material examined. 1 “Zhangmu, Xizang,

28,30.VI.1975, 2400m, leg. Huang Fusheng, Wang

Ziqing” (IZAS), 1 <$ “Xiao Jilong, Jilong, Xizang,

20, 22. VII. 1975, 2800m, leg. Huang Fusheng” (IZAS).

Lasioserica (s. I.) pacholatkoi Ahrens, 2000

Additional material examined. 1 S “Yadong, Xizang,

27. VIII. 2003, 2900m, light trap, leg. Xue Huaijun, Wang

Xinpu” (NKU).

Neoserica (s. 1.) lenangensis Ahrens & Fabrizi, 2009

Additional material examined. 1 <$ “Bomi, Xizang,

3. IX. 1982, 3050m, leg. Han Yinheng” (IZAS).

Neoserica (s. 1.) rutilans Ahrens & Fabrizi, 2009

Additional material examined. 1 S, I $ “Yigong,

Xizang, 1 1,1 5. VI. 1978, 2300m. leg. Li Fasheng” (CAU).

Neoserica (s. I.) sladeni Ahrens, 2004

Additional material examined. 1 $ “Sangjiu, Chayu,

Xizang, 23. VI. 1973, 3000m, leg. Huang Fusheng” (IZAS).

Oxy serica longefoliata (Frey, 1965)

Additional material examined. 1 $ “Zhangmu, Xizang,

5-6.' VII. 2004, leg. Ba Yibin, Shi Aimin’’ (HBUM).

Oxyserica pygidialis ssp. annapurnae (Ahrens, 1995)

Additional material examined. 2 S- 2 $ “Jilong District,

Jilong County, Xizang, 19-20. VII. 1975, 2800m, leg.

Zhang Xuezhong” (IZAS), 1 1 9 “Xiaojilong, Jilong,

Xizang, 20.VII.I975, 2800m, leg. Huang Fusheng”

(IZAS), 1 S “Jilonggongshe, Xizang, 27. VII. 1984, leg. Pu

Qiongqiong” (IZAS).

Gynaecoserica lohitensis Ahrens & Fabrizi, 2009

Additional material examined. 1 S “Chayu, Xizang,

16-17.VII.2005, leg. Shi Aimin’’ (HBUM).

Microserica pruinosa (Hope, 1831)

Additional material examined. 1 <$ “Youyiqiao, Niela-

rnu, Xizang, 6. V. 1974, 1680m, leg. Zhang Xuezhong”

(IZAS).

Microserica steelei Ahrens, 2004

Additional material examined. I S, 1 9 “Shajiao,

Chayu, Xizang, 25.VI.1978, 1700m, leg. Li Fasheng”

(CAU).

Nepaloserica archolabrata Ahrens & Sabatinelli, 1996

Additional material examined. I <$ “Youyi Bridge,

Nielamu, Xizang, 7. V. 1974, 1680m, leg. Zhang

Xuezhong” (IZAS).

Nepaloserica ganeshi Ahrens, 1999

Additional material examined. 2 S “Jilong, Xizang,

1 1 .VI. 1978, 3150m, leg. Institute of Agricultural Sciences”

(IZAS), I S “Jilong Gongshe, Xizang, 16.VI.1984, leg.

Yan Zhaoxing, Pu Qiongqiong” (IZAS), 1 S “Tongmai,

Jilong, Xizang, 25.V1.1984, leg. Yan Zhaoxing” (IZAS).

Nepaloserica procera procera Frey, 1965

Additional material examined. 1 $ “Qu County, Zhang-

mu, Xizang, 6. VII. 1 975, 3400m, leg. Zhang Xuezhong”

(IZAS), 2 S “Quxiang, Zhangmu, Xizang, 6. VII. 1 975,

3300m, 3400m, leg. Huang Fusheng, Zhang Xuezhong”

(IZAS), 1 S “Jilong Gongshe, Jilong, Xizang, 22. V. 1982,

leg. Li Baoming, Wangjia, Daci” (IZAS).

Maladera modestula (Brenske, 1902)

Additional material examined. “Beibeng, Motuo,

Xizang, 21 .V.1983, 700-800m, leg. Han Yinheng” (IZAS),

1 ^ “Beibeng, Motuo, Xizang, 25. V.1983, 850m, leg. Han

Yinheng” (IZAS).

Maladera siniaevi Ahrens, 2004

Additional material examined. 1 ex. “Beibeng, Motuo,

Xizang, 29.V.1983, 800-900m, leg. Han Yinheng” (IZAS).

Acknowledgements. This research was supported by the Na-

tional Basic Research Program of China (973 Program) (No.

201 1CB302I02), the National Natural Science Foundation of

China (Nos. 31010103913, 31172143), the Knowledge Inno-

vation Program of Chinese Academy of Sciences (No.

KSCX3-IOZ-1004), and by a Fellowship (to M.B.) from

Alexander von Humboldt Foundation. Additionally we thank

Prof. Wang Xinli (CAU), Prof. Pang Hong (LSSYU) and

Prof. Ren Guodong (HBUM) for the loan of unidentified

Sericini to Yang Xingke and Liu Wangang.

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172

Wan-Gang Liu et al.

REFERENCES

Ahrens D (2004) Monographs der Sericini des Himalaya

(Coleoptera, Scarabaeidae). Dissertation.de. Verlag im Inter-

net GmbH, Berlin, 534 pp.

Ahrens D (2005a) A taxonomic review on the Serica (s. str.)

MacLeay, 1819 species of Asiatic mainland (Coleoptera,

Scarabaeidae, Sericini). Nova Supplementa Entomologica 18:

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Ahrens D (2005b) A preliminary cladistic analysis of Nip-

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Ahrens D (2005c) Taxonomic revision of the genus Anomalo-

phylla Reitter, 1887 (Coleoptera, Scarabaeidae: Sericini).

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Ahrens D (2005d) The diversification of the endemic Himalayan

monsoon-season beetles genus Calloserica inferred from a

cladistic analysis (Coleoptera: Scarabaeidae: Sericini). Inver-

tebrate Systematics 19: 217-230

Ahrens D (2005e) New material of the genus Lasioserica

Brenske, 1896 with the description of four new species

(Coleoptera: Scarabaeidae: Sericini). Linzer Biologische

Beitrage 37: 771-781

Ahrens D (2006a) Phylogenetic analysis of Anomalophylla Re-

itter, 1887 (Coleoptera, Scarabaeidae: Sericini). Insect System-

atics and Evolution 37: 1-16

Ahrens D (2006b) The phylogeny of the genus Lasioserica in-

ferred from adult morphology - implications on the evolution

of montane fauna of the South Asian orogenic belt

(Coleoptera: Scarabaeidae: Sericini). Journal of Zoological

Systematics and Evolutionary Research 44: 34-53

Ahrens D (2006c) Additional data on taxonomy and distribution

on Sericini of the Himalayas, with description a further new

species of Maladera (Coleoptera, Scarabaeidae). pp. 409-^11 8.

In: Hartmann, M. and Baumbach, H. (eds. ): Biodiversitdt and

Naturausstattung im Himalaya. Verein der Freunde und

Forderer des Naturkundemuseums Erfurt e.V., Erfurt

Ahrens D (2006d) Evolution of Asian ‘lowland’ taxa and Alpine-

Himalayan Tertiary orogenic belt - Insight from a preliminary

cladistic analysis of Maladera (Cycloserica) (Coleoptera:

Scarabaeidae: Sericini). Zoologischer Anzeiger 244: 193-203

Ahrens D (2006e) Revision und phylogenetische Analyse der

Gattung Pachyserica Brenske, 1897 (Coleoptera, Melolonthi-

dae, Sericini). Revue Suisse de Zoologie 1 13: 487-557

Ahrens D (2006f) Cladistic analysis of Maladera ( Omaladera ):

implications on taxonomy, evolution and biogeography of the

Himalayan species (Coleoptera: Scarabaeidae: Sericini). Or-

ganisms, Diversity and Evolution 6: 1-16

Ahrens D (2007a) Cladistic analysis of Sericania (Coleoptera,

Scarabaeidae: Sericini) - implications on the evolution of the

xerophilous fauna of the Himalaya. European Journal of En-

tomology 104: 517-530

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sight from a phylogeny of the scarabaeid subgenus Serica

(Coleoptera, Scarabaeidae). Systematic Entomology 32:

450-476

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der Amiserica insperata (Brenske, 1898) aus dem

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Bonn zoological Bulletin 63 (2): 173-187

December 20 1 4

Neotropical primates from the Cologne Zoo in the collections of the

Zoologisches Forschiingsmuseum Alexander Koenig:

noteworthy specimens, taxonomic notes and general considerations

Gustav Peters Tamja Haus 2,3 & Rainer Hutterer 1

1 Department of Vertebrates, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany;

E-mail: g.peters@zfmk.de

2 Primate Genetics Laboratoiy, 3 Cognitive Ethology’ Laboratoiy, German Primate Center, Leibniz Institute for

Primate Research, Kellnerweg 4,

D-37077 Goettingen, Germany; E-mail: THaus@dpz.eu

Abstract. Cologne Zoo was a major place for the captive care of primates of the family Pitheciidae in the second half

of the last century. Various species of the genera Cacajao , Chiropotes and Pithecia , which until then had a poor survival

record in human care, lived at the Cologne Zoo for years. This offered the unique possibility to gather important infor-

mation on their biology and care in captivity. Moreover several individuals were featured in a diverse array of technical

publications and/or were mentioned in respective texts. However, at that time knowledge on the taxonomy of South Amer-

ican primates was still incomplete. New species and subspecies were named, some of which were kept in the zoo unno-

ticed. After their death Cologne Zoo donated specimens as vouchers to the Zoologisches Forschungsmuseum Alexander

Koenig. We evaluate their species identity in the context of the ongoing debate on the taxonomy of these three genera

and also address the potential importance of zoo specimens for the scientific study of taxonomic and biological ques-

tions. Furthermore, based on our data the status of Chiropotes israelita and C. sagulatus as valid species is questioned.

Key words. Primates, Pitheciidae, Neotropics, zoo biology, museum vouchers, taxonomy, identification.

INTRODUCTION

From the very beginning of the existence of zoological

museums, menageries and zoos have been an important

source of specimens for their collections, especially for

exotic species (Jalm 1994; Landsberg 1994; Oppermann

1994). Compared to specimens collected in their natural

distribution area (with appropriate documentation) those

originating from captivity usually have shortcomings in

respect of their informative value, significance as vouch-

ers, and thus also for many research topics typically car-

ried out in zoological museums. Although here is not the

place to discuss this in detail, specimens in museum col-

lections that come from a zoo can be important in vari-

ous aspects. Here we address this issue exemplarily for a

few Neotropical primate specimens in the mammal col-

lections of the Zoologisches Forschungsmuseum Alexan-

der Koenig, Bonn (ZFMK) that were donated by the

Cologne Zoo after the animals had deceased. The present

report is also a supplement to the catalogue of mammalian

types and other important voucher specimens in the ZFMK

(Hutterer & Peters 2010).

Two recent publications (Pauly 2010; Pagel & SpieB

201 1) animated us to write the present report and to put

forward some general considerations in this context. Pauly

(2010) discussed the species determination of a primate

Received: 20.01.2014

Accepted: 08.09.2014

individual that lived in Cologne Zoo between 1974 and

1 992 in view of a recently published taxonomic reassess-

ment of the black-headed uacari ( Cacajao

melanocephalus ; Boubli et al. 2008). Pagel & SpieB (201 1 )

provided a complete list of all mammal (sub-) species kept

at Cologne Zoo since its opening in 1 860 with contem-

porary and current scientific names. By discussing spe-

cific examples within a wider perspective we hope to in-

crease awareness both in zoological museums and zoos

that the lasting deposition of deceased zoo specimens in

a zoological museum and their subsequent scientific cu-

ration is the most reasonable ‘fate’ for zoologically im-

portant/interesting specimens of such origin. Only in do-

ing so their significance can be saved and their sustained

scientific use warranted. The same issue was addressed

by Gippoliti & Bruner (2007) and Gippoliti & Kitchener

(2007) in particular.

BACKGROUND

Cologne Zoo was a major place for the captive care of pri-

mates of the family Pitheciidae (taxonomy as in Groves

2005) outside their natural range in South America for sev-

Corresponding editor: J. Decher

174

Gustav Peters et al.

eral decades in the second half of the last century (cf. Hick

1965b, 1966, 1968a, b, 1973, 1974; Hershkovitz 1985,

1987a, b). Various species of the genera Cacajao , Chi-

vopotes and Pithecia , for which relatively little informa-

tion was available at that time and which until then had

a poor survival record in human care, lived at the zoo for

years and important information on their biology and care

in captivity was gathered and published (Hick 1965b,

1966, 1968a, b). Hershkovitz (1985: 1 7) named a new sub-

species of bearded saki Chiropotes satanas utahicki after

Ms. Uta Hick (subsequently and correctly, the name was

changed to utahickae , following the International Code of

Zoological Nomenclature; ICZN 1999), the then-curator

of primates at Cologne Zoo. Hershkovitz (1985) did so to

acknowledge her achievements in the captive care of

Pitheciidae and of her major contributions to the knowl-

edge of these primates. Photos of several live individuals

of these three genera from Cologne Zoo were presented

in technical publications and/or they were mentioned in

their text (e.g. Hill 1960; Napier & Napier 1967; Mitter-

meier & Coimbra-Filho 1981; van Roosmalen et al. 1981;

Hershkovitz 1985, 1987 a, b). At that time the taxonomy

of Cacajao, Chiropotes and Pithecia was far from being

settled (Hershkovitz 1985, 1987 a, b). In the context of

the recent 'revival’ of the debate on the taxonomy of Chi-

ropotes (Boubli 2002; Silva-Jr. & Figueiredo 2002; Boli-

viano et al. 2003; Veiga et al. 2008; Ferrari et al. 2013;

Silva-Jr. et al. 2013), Cacajao (Boubli et al. 2008; Ferrari

et al. 2013; Silva-Jr. et al. 2013) and Pithecia (Marsh 2004,

2006, 2014), some specimens of these taxa, which the

ZFMK received from Cologne Zoo, regain significance.

Unfortunately, the ZFMK received only a minor fraction

of the individuals that lived in the Cologne Zoo between

1954 and 2005, and for none of these specimens verifi-

able data as to their geographic origin are available. There-

fore statements with respect to agreement/disagreement

of their characteristics with those reported for specimens

from specific geographic areas are made conditionally. A

revision of the genus Pithecia (Flershkovitz 1987b) in-

cludes a photo of a live individual of this genus from

Cologne Zoo. Although it was not given to the ZFMK af-

ter its death, it will be addressed in the present report as

well to indicate the potential importance of such zoo spec-

imens for the scientific community.

MATERIAL AND METHODS

Our study is based on the voucher specimens of Neotrop-

ical primates deposited in the mammal collections of Zo-

ologisches Forschungsmuseum Alexander Koenig

(ZFMK MAM). Species of Cacajao , Chiropotes and

Pithecia that originated from Cologne Zoo were analysed

in the context of published and unpublished zoo records

and photographs made available by the zoo archivists R.

Becker and W. SpieB. We are aware of the limited signif-

icance of photos in drawing fully substantiated conclusions

as to the ‘actual’ colour in cases where (fur) colour char-

acteristics are diagnostic criteria for the differentiation of

taxa and will address this issue where relevant. The same

applies to the fading and possible change of colours in

long-term museum specimens, especially those on exhib-

it.

Nomenclature

English common names of taxa used in the present pub-

lication are those given in the primate chapter of Mam-

mal Species of the World (Groves 2005). Some names

coined in Ferrari et al. (2013) are in conflict with the tax-

onomic views expressed here. Vernacular names with ad-

jective characterizations of the species' body coat colour

are adopted with reservations because they imply that it

is a diagnostic character. This can be delusive, though, as

will be detailed below.

Molecular methods

DNA extraction, amplification and sequencing

To determine a questionable Chiropotes specimen from

Cologne Zoo (ZFMK_MAM_2008.244), whose species

affiliation could not be clearly determined based on ex-

ternal characters, we extracted DNA from a small piece

of skin in the ancient DNA laboratory at York Universi-

ty following the protocol of Haus et al. (2013). We used

0.09 mg sample and incubated it for about 24h in 1.5 ml

of a Guanidinium thiocyanate (GuSCN) buffer (5M

GuSCN, 25 mM NaCl, 50 mM Tris, 20 mM EDTA, 1%

Tween 20, 1% beta-mercaptoethanol), which was modi-

fied from Rohland et al. (2004). To purify the DNA we

applied a combination of a batch-based silica and a col-

umn-based method according to Rohland & Hofreiter

(2007) and Rohland et al. (2010). Finally we eluted the

DNA in 50 pi TE buffer. To monitor for possible contam-

ination we additionally performed a blank extraction with-

out sample.

To compare our data with already published sequences

of other Chiropotes taxa we amplified and sequenced a

390 bp long fragment of the cytochrome (cyt) b gene. Boli-

viano et al. (2003) used a 372 base pair (bp) long frag-

ment of the cyt b gene to distinguish between different

species of Chiropotes. Accordingly, we amplified the same

cyt b fragment using primers Cytb fwTGATATGAAA-

AACCATCGTTG (modified from Bonvicino et al. 2001)

and Cytb rvACCTATAAATGCTGTAGCTAT in a 20 pi

mix (3 U AmpliTaq Gold 360 (Applied Biosystems, Ger-

many), lx reaction buffer, 2 mM MgC^, 0.25 mM for

each dNTP, 0.75 pM for each primer, and 0. 1 mg/ml BSA)

and following PCR conditions: 94°C for 10 min, followed

by 60 cycles of 94°C for 30 s 62°C for 45 s, 72°C 45 s.

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Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 175

and 72°C for 5 min. Due to the length of our amplified

product (390 bp) and the usually high degradation of DNA

retrieved from museum samples, the possibility that our

sequence represents a putative nuclear insertion of mito-

chondrial DNA (numt) is low. To control for possible con-

tamination with modem DNA, the PCR reaction was con-

ducted with two PCR blanks (HPLC-purified water) in ad-

dition to the extraction blank. We ran the PCR product on

a 1.5% agarose gel, and purified it with the Qiagen Gel

Extraction Kit (Qiagen, Germany) after excision. Subse-

quently, forward and reverse sequences were run on an

ABI 3 1 30 xL sequencer using the BigDye Terminator Cy-

cle Sequencing Kit (Applied Biosystems, Germany) and

respective primers. The Cliiropotes sequence can be down-

loaded from GenBank (GenBank accession number:

KF989492).

Phylogenetic analysis

To examine phylogenetic relationships we used se-

quences of six related taxa as outgroups ( Alouatta belze-

bul AF2895 15, Saimiri sciureus U53582, Callicebus per-

sonatus AF289988, Callicebus donacophilus FJ85423,

Cacajao melanocephalus AY226184, Pithecia irrorata

AY226183) and further included all Cliiropotes cyt b se-

quences available from GenBank: Cliiropotes albinasus :

KC757393; C. utahickae : AY226185, AY226186; C. is-

raelita: AY226187-AY226190; C. chiropotes/sagulatus :

FJ53 1667). We used the same Cliiropotes sequences as in

Bonvicino et al. (2003), except for Cliiropotes albinasus

(Finstermeier et al. 2013) and the cyt b gene of C. chi-

ropotes/ sagulatus from Rio Trombetas east of Rio Bran-

co (Figueiredo-Ready et al. 2013). All sequences were

aligned using the program Mega 5.0 (Tamura et al. 201 1 ).

We also used Mega 5.0 to check for putative numts; we

translated all sequences into amino acid sequences to de-

tect possible unexpected stop codons in our alignment,

which was not the case. Best fit nucleotide substitution

models were estimated with jModeltest (Guindon & Gas-

cuel 2003; Posada 2008) and chosen based on the Akaike

Information Criterion (AIC). Correspondingly, we used

the TrN+G model to reconstruct neighbor-joining (NJ;

Mega 5.0, Tamura et al. 2011), maximum-likelihood (ML;

Garli 2.0, Zwickl 2006; Paup*4bl0, Swofford 1993) and

Bayesian (MrBayes 3.1.2, Huelsenbeck et al. 2001, Ron-

quist & Huelsenbeck 2003 ) phylogenetic trees. For NJ and

ML we used 1 000 and 500 bootstrap replications, respec-

tively. We applied 10 million generations for the Bayesian

approach and sampled parameters every 10,000 genera-

tions; the first 25% of sampled trees and parameters were

discarded as bum in. Phylogenetic trees were finally vi-

sualized and edited in FigTree 1.4.0 (Rambaut 2008).

RESULTS & DISCUSSION

Cacajao Lesson, 1840

After the past argument on whether red and white uacaris

are the same or different species (Napier & Napier 1 967;

Chiarelli 1972) had been provisionally settled (Her-

shkovitz 1972, 1987a), the genus was generally consid-

ered to include two species, the black-headed uacari Ca-

cajao melanocephalus (Humboldt, 1811) (not 1 8 12, as in

Groves 2005), and the bald uacari Cacajao calvus (I. Ge-

offrey, 1847), the latter encompassing the white and the

red form (Barnett & Brandon-Jones 1997; Groves 2001,

2005; Barnett 2005). More recently, based on genetic,

morphological and ecological data Boubli et al. (2008) de-

scribed two new species of black-headed uacari, Cacajao

hosomi Boubli et al., 2008, and Cacajao ayresi Boubli et

al.. 2008. Moreover, they argued that ouakary( Spix, 1823)

presents a junior synonym of melanocephalus according

to the original description of pelage colour of both spec-

imens, the uncertainty of the type locality of Humboldt’s

specimen of melanocephalus and to the now knowledge

of the geographical distribution of different pelage patterns

in this group. This taxonomic arrangement was not fol-

lowed by Ferrari et al. (2010) and Silva-Jr. et al. (2013),

who proposed to recognise three species Cacajao calvus ,

C. melanocephalus and C. ouakary (Ferrari et al. 2010),

and additionally the newly described species Cacajao

ayresi Boubli et al., 2008 (Silva-Jr. et al. 2013). Howev-

er, since no published data exist to support findings of Fer-

rari, Silva-Jr. and colleagues, for now we will retain the

taxonomy of Boubli et al. (2008). Their publication is al-

so relevant for a specimen the ZFMK received from

Cologne Zoo, which we will discuss below in more de-

tail.

Cacajao melanocephalus (Humboldt, 1811) sensu lato

- Black-headed uacari

According to the annual animal registers of Cologne Zoo

(SpieB, pers. comm. 2011; Becker, pers. comm. 2011) two

individuals were received in 1958 and one in 1959. All

three individuals only lived for a short time and no more

specific documentation for these specimens is left. Fur-

ther, no material was transferred to a zoological museum

or another scientific institution after their death. Therefore

it is not possible to retroactively assess their actual species

identity in the light of the recent publication of Boubli et

al. (2008).

Cacajao hosomi Boubli ct al. 2008 - Neblina uacari

Material: ZFMK_MAM_2002.032; adult male, received

by Cologne Zoo 15.111. 1974, f 23. IX. 1992, carcass passed

to ZFMK on 12.VI.I997; skull, postcranial skeleton.

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Gustav Peters et al.

In 1974 the zoo received another individual of black-head-

ed uacari from Venezuela, a male (Windecker 1975),

which lived until 1992, the highest longevity recorded for

a black-headed uacari (Weigl 2005). Published photo-

graphs of the animal, then identified as Cacajao

melanocephalus , show this male at various stages of its

life (Hick 1974; Mittermeier and Coimbra-Filho 1981;

Pauly 20 1 0). In a black-and-white photograph of the adult

individual taken in 1985 by Hershkovitz (1987a: 30, fig.

16) it is identified as Cacajao melanocephalus ouakary

(Spix, 1823). The diagnostic back pelage colouration of

this subspecies given by Hershkovitz (1987a: 25) as ‘Mid-

back pale orange, golden or buffy and contrasted with red-

dish brown or tawny of lower back and thighs’ was not

present in this individual, though (cf. colour photographs

of it mentioned subsequently), i.e. its subspecies identi-

fication is unfounded. Colour photographs taken subse-

quently, observations of the live animal by one of us (G.P.)

and of the carcass revealed no orange tinge on the back

at all. It is possible that this mistake slipped in on the oc-

casion of the later subspecies identification based on the

black-and-white photograph of Hershkovitz (1987a) as the

animal’s back seems brightened because of the lighting

conditions when the photo was taken and/or the exposure

time used.

Only after the publication of Boubli et al. (2008) it be-

came evident that this uacari individual at Cologne Zoo

belonged to the newly described species Cacajao hoso-

mi Boubli et al., 2008 (Pauly 2010). Therefore, the com-

mon name proposed by its authors is used here. Various

colour photographs of the live individual in publications

by Cologne Zoo (zoo guides, title page of the journal

‘Zeitschrift des Kolner Zoo’ 26(4), 1983) and in Pauly

(2010: 186, Abb. 6) support its identification as C. hoso-

mi in showing its diagnostic colouration (Boubli et al.

[2008]: fig. 3: 4, 5, p. 737 [description]): ‘ fur with bright

reddish brown mid to lower back, thighs and tail, in con-

trast to Araca black uacari specimens that have a much

darker general coloration and to C. melanocephalus as re-

defined here, which present a golden brown general ap-

pearance.’

Chiropotes Lesson, 1840

Contrary to an earlier statement by Mittermeier & Coim-

bra-Filho (1981: 74) that ‘[t]he genus Chiropotes . . . does

not present any particular problems’, the species status of

various taxa of Chiropotes satanas (Hoffmannsegg,

1 807) sensu lato has been the subject of recurring discus-

sions (Hershkovitz 1985; Rylands et al. 1995; Rylands et

al. 2000; Groves 2001 , 2005; Silva-Jr. & Figueiredo 2002;

Bonvicino et al. 2003; Figueiredo et al. 2006; Veiga et al.

2008; Rylands & Mittermeier 2009; Ferrari et al. 2013;

Silva-Jr. et al. 2013). As the diagnostic characteristics

(nasal region and upper lip flesh-coloured, with whitish

or yellowish hairs [Hershkovitz 1985]) of Chiropotes al-

binasus (1. Geoffroy and Deville, 1848) are clearly defined

and unambiguously observable, its status as a distinct

species has not been debated (Hill 1960; Chiarelli 1972;

Napier 1976; Hershkovitz 1985; Groves 2001, 2005).

Based on the taxonomic review of Hershkovitz ( 1985), in

which he also described the new subspecies Chiropotes

satanas utahicki , all other taxa of the genus Chiropotes

were likewise classified as subspecies of Chiropotes sa-

tanas by subsequent authors (e.g. Rylands et al. 1995,

2000; Rowe 1996; Groves 2001).

By contrast, Chiropotes a/binasus, C. satanas , C. chi-

ropotes, and C. utahickae were regarded as distinct species

by Bonvicino et al. (2003). Moreover, these authors list-

ed Chiropotes israelita (Spix, 1823), the brown-backed

bearded sakis of northwestern Brazil, north of the Rio Ne-

gro and west of the Rio Branco, as an additional species.

In the primate chapter of ‘Mammal Species of the World’

Groves (2005) adopted this view, a procedure not followed

by Norconk (2007) who like Hershkovitz ( 1985) or Groves

(2001 ) listed only two species in the genus, C. albinasus

and C. satanas. Both last-mentioned authors regarded is-

raelita as a synonym of Chiropotes satanas chiropotes.

More recently, Silva-Jr. et al (2013) and Ferrari et al.

(2013) supported a five species concept as proposed by

Bonvicino et al. (2003), but they considered the taxon west

of the Rio Branco to be Chiropotes chiropotes and that

east of the river to be Chiropotes sagulatus (Traill, 1821).

There are no published data so far that clearly underpin

the taxonomic conclusions of Ferrari et al. (2013) and Sil-

va-Jr. et al. (2013) preventing us from drawing any final

conclusions on the taxonomy of Chiropotes.

Bonvicino et al. (2003) explicated in detail why they

consider the genus Chiropotes to include five distinct

species and why the name israelita ought to be applied

to the Rio Negro population, as also done by Boubli &

de Lima (2009). The remarks of Bonvicino et al. (2003)

in this latter context contain two material misapprehen-

sions, though: 1. There is no substantial evidence that

Spix’s (1823: 1 1-12) original description of Brachyurus

israelita (first in Latin and then in French) refers to two

different specimens as claimed by Bonvicino et al. (2003:

1 32). The specification of the type locality in the Latin text

as ‘Habitat ad Rio-Negro’ (p. 11) and that - as opposed

to this - at the end of the more detailed French text (p.

12) as ‘On trouve ce singe vers le Perou dans les forets

de Japura riviere laterale de Solimoens’ must have been

an error in the case of the latter statement since there is

not a single record known for the occurrence of bearded

sakis south-west of the Rio Negro (Hershkovitz 1985;

Veiga et al. 2008; Ferrari et al. 2013; Silva-Jr. et al. 2013).

Therefore we argue that this information cannot be taken

as evidence of the existence of two different specimens

on which the respective descriptions were based. Indeed,

there is also no clear difference between characters de-

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Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 177

tailed in the Latin text of Spix’s ( 1823) original descrip-

tion of Brachyurus israelita and those described in the sub-

sequent French text as claimed by Bonvicino et al. (2003).

Furthermore, there is no indication for the (past) existence

of a second specimen of bearded saki in the Spix collec-

tion (M. Hienneier, ZSM, pers. comm. 2012). 2. Contrary

to the assumption/statement of Bonvicino et al. (2003:

132) that the holotype specimen of Brachyurus israelita

Spix, 1 823 is lost, it is preserved in the collection of the

Zoologische Staatssammlung Miinchen (ZSM) (Kraft

1983); pertinent information as well as its colour photo

are available on the internet (see http:// www.zsm.mwn.de/

mam/ptypes.htm, and http://www.zsm. mwn.de/mam/i/

Brachyurus israelita 34 D.jpg). Additional colour pho-

tos of the holotype specimen of Brachyurus israelita Spix,

1 823, kindly provided by M. Hienneier (cf. Fig. 1 ), show

the typical coat colouration of Chiropotes satanas chi-

ropotes as given in the key to the geographic forms of Chi-

ropotes ( Hershkovitz 1985: 13): ‘Head, nape, lower aims,

and legs blackish, sharply contrasted with orange of dor-

sum’.

Moreover, the statement by Bonvicino et al. (2003: 131)

that ‘... Hershkovitz [1985] commented that there was no

assigned C. s. chiropotes holotype or type locality. ’ is for-

mulated inappropriately and mistakable. In listing the type

locality of Simia chiropotes as ‘Said to be the upper Rio

Orinoco south of the cataracts, Amazonas, Venezuela’ ac-

cording to Humboldt (1811) and in stating that the type

is not preserved (i.e. was lost during its transport from

South America to Europe), as already reported by Hum-

boldt (1811), Hershkovitz (1985: 16) rendered the issue

correctly. Thus, all available evidence supports the per-

ception that Chiropotes israelita (Spix, 1823) is a junior

synonym of Chiropotes chiropotes (Humboldt, 1811).

Shortly before the study by Bonvicino et al. (2003) was

published Silva-Jr. & Figueiredo (2002) had reintroduced

the name Chiropotes sagulatus (Traill, 1821 ), originally

described from Guiana, as a distinct species. Bonvicino

et al. (2003: 132) thereupon argued ‘Silva-Jr. and

Figueiredo (2002) recently proposed species status for C.

sagulatus ; however, they did not provide sufficient data

to justify that taxonomic arrangement. ’ Therefore Bon-

vicino et al. (2003) did not list sagulatus as a distinct

species of Chiropotes. Husson (1957, 1978), in previous

specific surveys of the mammal fauna of Guiana, had treat-

ed sagulatus as a synonym of Chiropotes satanas chi-

ropotes, like most other authors of later pertinent techni-

cal publications (e.g. Hershkovitz 1985; Groves, 2001,

2005). Bonvicino et al. (2003) rightly pointed out that no

substantial data - in effect no specific data at all - was

presented by Silva-Jr. & Figueiredo (2002) in the pub-

lished abstract of their contribution to the Xth Congres-

so Brasileiro dc Primatologia and that therefore this ab-

stract cannot be the proper basis for the resurrection of

sagulatus as a distinct species. On the other hand, this does

not rule out that the study of Silva-Jr. & Figueiredo (2002)

underlying their congress contribution yielded data sup-

porting such a view. In return, it does not have any rele-

vance in supporting the view of Bonvicino et al. (2003)

that the name israelita ought to be applied to the Chi-

ropotes population west of the Rio Branco and north of

the Rio Negro, and that it represents a distinct species.

Unlike Groves (2005) who adopted the concept of Bon-

vicino et al. (2003) in its entirety, a more recent review

of the diversity of New World primates (Rylands & Mit-

termeier 2009: 40-41 ) pointed out that the cause of such

discrepancies may be ‘ . . . a possible confusion concern-

ing the correct names.’ Indeed, the distribution map pub-

lished by Bonvicino et al. (2003: 125, fig. 1) shows lo-

calities of Chiropotes satanas chiropotes only consider-

ably east of the Rio Branco in Brazil. All localities west

of it and north of the Rio Negro are entered as Chiropotes

sp. The locality of a ‘possibly new taxon of Chiropotes''

reported by Boubli (2002) and previously mentioned by

Rylands et al. (2000) is still further northwest, close to the

southernmost border of Venezuela. All these locality

records are within the distribution range of Chiropotes sa-

tanas chiropotes as given by Hershkovitz ( 1985: 2, Fig.

1 ). In applying the name israelita to their ‘new form of

Chiropotes' the taxonomic views of Bonvicino et al.

(2003) were apparently put forward without observance

of the statement of the type locality (‘On les trouve dans

les vastes deserts de l’Alto-Orinoco, an sud et a Test des

Cataractes.’) in Humboldt’s (1811: 313) original descrip-

tion of Simia chiropotes. It is usually rendered as upper

Rio Orinoco, south of the cataracts, Amazonas State,

Venezuela (Cabrera 1961; Hershkovitz 1985; Groves

2001, 2005). Incidentally, the Chiropotes sp. individual

from the lower Rio Marauia, Amazonas, Brazil, reported

and pictured in a colour photograph by Boubli (2002: fig.

3) clearly shows the typical coat colouration of Chiropotes

satanas chiropotes as listed by Hershkovitz (1985: 13).

The conclusion of Bonvicino et al. (2003: 129) that of the

four Chiropotes taxa they studied the western two differ

in dorsal coat colour (tawny-olive to buffy-brown in the

westernmost versus orange or ochraceus in the eastern tax-

on) obviously contravenes that by Hershkovitz ( 1985: 17):

‘All types of chromatic variation are fully intergrading and

may occur in the same population. Geographic variation

is not evident.’ Of the original descriptions of the rele-

vant Chiropotes taxa named, only the one of israelita (Spix

1823: 11-12) is sufficiently detailed (in its subsequent

French version) to provide pertinent information in this

respect: ‘Les poils du dos sont tres epais, peu longs, d’un

roux jaunatre, ceux des cotes d’un fauve plus fonce’. This

clearly indicates a ‘reddish-yellowish’ hue of the back fur

(as still detectable in the type specimen, see Fig. 1 ). Aside

from the question whether israelita is a synonym of chi-

ropotes there can be no doubt that its type locality places

israelita into the westernmost form of bearded saki. So,

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Gustav Peters et al.

Fig. 1-3. 1. (Left) Back view of the mounted type specimen of Brachyuras israelita Spix, 1823 in the Zoologische Staaatssamm-

lung Miinchen ZSM (Photo: M. Hienneier, ZSM). 2. (Center) Back view of mounted adult male Chiropotes chiropotes specimen

ZFMK_MAM_66.001 (Photo: M. Weigt). 3. (Right) Tanned skin of Chiropotes sp. specimen ZFMK_MAM_2008.244 (Photo: M.

Weigt).

a reddish/orange tinge of the dorsal pelage colour may al-

so be present in this population.

Gregory (201 1 ) who kindly provided colour photos of

living bearded sakis from Suriname opines that these an-

imals have ‘a much brighter orange dorsum’ (visible in the

photos) than the more western populations of bearded

sakis (pers. comm. 2012); she uses the name Chiropotes

sagulatus for them (Gregory 2011; Gregory & Norconk

201 1). Concerning dorsal pelage colour the original de-

scription of Simia sagulatus , the Guianan bearded saki,

states ‘back well clothed with an ochry-coloured fur’

(Traill 1821: 167), and in more detail (p. 168) ‘The colour

is brightest on the shoulders, where it is intermediate be-

tween wood-brown and yellowish-brown; and, on the back

and sides, passes into the latter.’ Traill (1821: 1 69) express-

ly states that these characteristics equally apply to live

more specimens from the same geographical area (Demer-

ara/Demerary in today’s Guayana; originally a Dutch col-

onization area, later occupied by the British) as the holo-

type. Husson ( 1 978: 210) describes the colour of the back

pelage of nine Chiropotes satanas chiropotes specimens

from Suriname as ‘. . . a dull brownish colour varying from

yellowish umber to mummy brown.’ which suggests quite

some variation in this (distinctive?) character even

among individuals from within a restricted portion of the

distribution range of this taxon. Apart from colour fading

likely to have occurred in museum specimens, these dis-

crepancies may be due to differences in the relative cov-

erage of the whole distribution range of Chiropotes sa-

tanas sensu lato surveyed and in over-all sample size and

that for specific populations in these studies. Thereby the

study of Hershkovitz (1985) is based on more specimens

from a wider geographic range than any other published

study so far.

Rylands & Mittermeier (2009: 41 ) stated that ‘Silva-Jr.

and Figueiredo (2002) argue that the form west of the Rio

Branco is correctly C. chiropotes , according to its type lo-

cality, and the form to the east of the Rio Branco, extend-

ing through the states of Para and Amapa, and into the

Guianas (Guyana, Suriname, French Guiana) is C. sagu-

latus (Traill, 1821 ) from Demerara.’. However, they pro-

vided more detailed information on the issue than the pub-

lished abstract by Silva-Jr. & Figueiredo (2002) actually

contains. It may be assumed that it was obtained from

these latter authors personally. The same is true of the

statement in Veiga et al. (2008): ‘... Chiropotes sagula-

tus Traill, 1821, the latter representing the eastern form

of C. chiropotes , which occurs to the east of the rio Bran-

co, in Brazil. Suriname and the Guianas ’. Unfortunately

there are also no relevant data presented in the most re-

cent publications of Silva-Jr. et al. (2013) and Ferrari et

al. (2013) , which might support their taxonomic arrange-

ment. Yet, to the best of our knowledge no ‘proper’ per-

tinent study has been published so far presenting evidence

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Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 179

that C. chiropotes and C. sagalatus represent different

species, or that they possess different distributional ranges.

The name has recently been used, though, for bearded

sakis from Suriname (e.g. Gregory & Norconk 2011) and

the state of Para (north of the Amazon), Brazil (e.g.

Oliveira et al. 2009). The karyotypic differences between

three Chiropotes taxa (not including C. s. satanas and C.

albinasus) reported by Bonvicino et al. (2003), which in

their view strongly indicate their status as distinct

species, may actually support this concept. They found

that the Rio Branco is the parting line of the two karyotyp-

ically different Chiropotes forms north of the Amazon.

There is little data on fur colouration in juvenile indi-

viduals (Hershkovitz 1985); so it is not possible to eval-

uate whether, to which extent and how this characteristic

changes during ontogeny in the respective taxa. Anyhow,

an in-depth comprehensive revision based on as large a

number of specimens as possible from the entire distrib-

utional range, integrating classical morphological and

modem genetic methods, in combination with all other ev-

idence, is still required to clarify the taxonomy of the var-

ious forms of Chiropotes satanas sensu lato.

Chiropotes albinasus (I. Geoffroy & Devi lie, 1848) -

White-nosed saki

Material: Three of five females of Chiropotes albinasus

that lived at Cologne Zoo are in the ZFMK mammal col-

lection. ZFMK MAM 7 1.97b: adult female, f 1971, re-

ceived from Cologne Zoo 1971; skull, flat skin. This is

probably the individual the zoo received as a juvenile in

1968 (Windecker 1969). ZFMKMAM81 .1 815: adult fe-

male, f 17.VI.1981. received from Cologne Zoo VI-

11.1981; skull, postcranial skeleton, flat skin. This is the

individual ‘Bella”, the mother of four of the Chiropotes

hybrids bom at the zoo. This female arrived at the zoo in

1965 (Flick 1965a) and thus attained an age of >16 years.

Most of the photos published of Chiropotes albinasus from

Cologne Zoo (with its hybrid offspring) show this indi-

vidual. ZFMK_MAM 89.480: adult female, f

01. IX. 1988, received from Cologne Zoo IX. 1989; skull,

postcranial skeleton, flat skin. This is the female with the

Zoo house name ‘Bianca’ which arrived there 08.X.1970

(Windecker 1971 ) and is the individual with the maximal

longevity (> 18 years) attained by this species in any zoo

( Weigl 2005). During the time this female lived at the zoo

it gave birth to two offspring. Its estimated age of about

20 years noted by Weigl (2005) would imply that it was

roughly 2 years old when it arrived at the zoo.

The first documented individual of Chiropotes albina-

sus in the care of Cologne Zoo, an adult female, arrived

in 1965 (Hick 1965a; Pagel & SpieB 2011). In 1968 the

zoo received a juvenile female (Windecker 1969, anoth-

er female which arrived in 1966 only survived for a few

months) and in 1970 another two juvenile females

(Windecker 1971). Apart from popular print products of

the Cologne Zoo with photos (black-and-white and colour)

of several of these individuals (e.g. Hick 1968a: 35, 1973;

Windecker 1969; front pages of the journal ‘Freunde des

Kolner Zoo’ 9[3], 1966 and 1 1 [2], 1968), photos of them

also appeared in technical publications (Hick 1968b: Fig.

31; Roosmalen et al. 1981: 420; Hershkovitz 1985: 26).

Two of the white-nosed saki females gave birth success-

fully to hybrid young: a male out of a mating of Chiropotes

albinasus female x Chiropotes chiropotes male in 1968

(Hick 1968a) and five more hybrids (2 males, 3 females)

out of matings of Chiropotes albinasus female x Chi-

ropotes satanas male in 1973 (2), 1975, 1976, and 1979;

in 1978 one female had a miscarriage after such a mat-

ing. Like the first hybrid bom in 1968 (Hick 1968a), some

of the others born later were figured in photos, usually to-

gether with their mother (Kullmann 1976). The first hy-

brid bom in 1968 is mentioned and also figured in tech-

nical publications (Roosmalen et al. 1981: 420; Her-

shkovitz 1985: 26). Hershkovitz’s statement is not correct;

the sire of this hybrid offspring was a Chiropotes chi-

ropotes male (Becker, pers. comm. 2014). Four of the hy-

brid offspring were bom by the female ‘Bella’, two by the

female ‘Bianca’, and the same Chiropotes satanas male

was the sire of 4 of the 5 hybrids bom since 1973. A fe-

male hybrid bom in 1975 lived at Cologne Zoo for more

than 26 years (Weigl 2005). Occasionally one of these ju-

venile hybrids was wrongly labelled as Chiropotes albi-

nasus, resp. no mention was made that the juvenile is a

hybrid, e.g. in the publications of Kullmann (1976) and

Mittermeier & Coimbra-Filho (1981: Fig. 53, p. 86) or in

a figure legend (Zeitschrift des Kolner Zoo 1 7 ( 1 ), p. 1 ).

The same is probably tme for a colour photo of an adult

‘female white-nosed saki (Chiropotes albinasus)' in

Pauly (20 1 0: 1 82, Abb. 1 ). If the date when the photo was

taken (24.V.1990 as stated in the figure legend) is correct,

this individual is a hybrid of Chiropotes albinasus female

x Chiropotes satanas male because according to the zoo

archives the last female white-nosed saki at Cologne Zoo

died in 1988. In 1990 only two female hybrids were liv-

ing at the zoo (SpieB, pers. comm. 2011; Becker, per.

comm. 2012). Unfortunately, none of the altogether six hy-

brids bom at the zoo was given to the ZFMK, another zo-

ological museum or a scientific institute after their death.

Chiropotes chiropotes (Humboldt, 1811) —

Red-backed bearded saki

Material: ZFMK MAM 66.001 ; adult male,

f 04.01.1966, received from Cologne Zoo 1966; skull,

taxidermy mount (Fig. 2).

This individual arrived at the zoo as a juvenile in 1961

and was originally registered as a female which was lat-

er corrected. It is mentioned in Anonymous (1962) and

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Gustav Peters et al.

shown at different ages in two black-and-white photo-

graphs, the earlier taken in May 1962, the later in Sep-

tember 1963 (Anonymous 1963: 87). Comparing both

photographs the pronounced growth of the typical thick

split tuft of hair on the head and that of the eponymous

beard during the time span of about one year becomes ob-

vious. Photos of Chiropotes chiropotes individuals kept

at Cologne Zoo and published from 1964 onwards by zoo

employees (Hick 1965a, 1966) very likely show other in-

dividuals of this species living there, as two more indi-

viduals arrived at the zoo in 1964 and 1965.

Description of the specimen (Fig. 2): As the specimen

was on exhibit in the museum for a few decades, a gen-

eral fading of its pelage colouration is likely to have oc-

curred, especially in respect of possible yellowish/reddish

hues that may have been present in the living animal. Fol-

lowing statements are made under this qualification. There

is a marked contrast between the homogeneous light buffy

ochre of the back and the dark brown to black coloura-

tion of the fur of the head, extremities and the tail. The

colouration of the back extends into the flanks and the

arms and thighs. On the upper arms pelage colour grad-

ually changes into blackish brown of the more distal por-

tion of the anns, on the upper side interspersed with lighter

hairs. This colour change is more abrupt on the proximal

part of the thighs. The backs of hands and feet are cov-

ered by brown hair. The venter is black. The fur colour

on the head (with the beard and the split tuft of hair on

top of it), nape and tail is basically black. The change in

fur colour from the trunk to these body regions is abrupt.

Taking into account the unsettled taxonomic issues ad-

dressed earlier, the actual species identity of all relevant

specimens that lived at Cologne Zoo cannot be verified

resp. refuted retroactively, even more for those for which

no photos are available and/or no material was given to

the ZFMK or to another scientific institution after the an-

imals’ death. In addition, it is difficult to determine the

species identity of juvenile bearded saki individuals ret-

rospectively on the basis of photographic portraits with

adequate certainty. Therefore we use the scientific names

here under which they are registered in the zoo archives.

According to these (SpieB, pers. comm. 2011; Becker,

pers. comm. 2011) Cologne Zoo received four specimens

of Chiropotes chiropotes between 1961 and 1977, three

males and one individual of unknown sex which died a

few months after its arrival whereas the other individuals

all lived at the zoo for several years. One of these males

(house name ‘Ringo’), which arrived at the zoo as a ju-

venile in 1964, was the father of the first hybrid saki

(mother Chiropotes albinasus ) bom at Cologne Zoo in

1968 (Hick 1968a). This male, given to a private animal

husbandry in 1 982, was figured in Hick ( 1 968a: 38 & 39),

van Roosmalen et al. (1981: 421; fig. 2); photos in Napi-

er and Napier (1967: 121, plate 33) and Hershkovitz

(1985: 14, fig. 8) probably also show it. Unfortunately, no

remains of this individual are conserved in a scientific in-

stitution.

Chiropotes sp. - Bearded saki

Material: ZFMK_MAM_2008.244; adult male, f 1991,

received from Cologne Zoo 5. IX. 1991; skull, postcranial

skeleton, flat skin (Fig. 3). Animal received by Cologne

Zoo as a present from the defunct Dierenpark (zoo) Wasse-

naar, Netherlands, in 1978 and originally listed as Chi-

ropotes satcmas chiropotes in the zoo archives (Kullmann

1979). A colour photo of this individual alive, taken

24. V. 1990 and referred to as Chiropotes chiropotes , was

published by Pauly (20 1 0: 1 83, Abb. 2). However, the dor-

i

0 02

H

SSU53582 Saimiri sdureus

AF289515 Alouatta belzebul

FJ785423 Callicebus donacophilus

■ AF289988 Callicebus personalus

AY226183 Pithecia irrorata

AY226184 Cacajao melanocephalus

Fig. 4. Cytochrome b phylogeny of the genus Chiropotes. A, Posterior probabilities of the Bayesian approach and bootstrap sup-

port values of the maximum-likelihood analysis before and after the slash, respectively. B, Phylogenetic relationships and boot-

strap support values based on the neighbor-joining approach. Sample IDs correspond to GenBank accession numbers.

Bonn zoological Bulletin 63 (2): 173-187

©ZFMK

Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 181

sum of this specimen is light buffy to ochre contrasting

the dark brown to blackish colour of the outer side of the

upper and lower arms, legs, the fur of the head, and the

outer side of the thighs. According to Hershkovitz

( 1985:13) characters of chiropotes and utahickae are de-

scribed as ‘Dorsum dominantly orange or pale brown to

dark brown; outer side of upper arms and proximal part

of outer side of thighs orange to dark brown, not black-

ish’ and as ‘Head, nape, lower arms, and legs blackish,

sharply contrasted with orange of dorsum' in chiropotes

vs. ‘Head, nape, lower arms, and legs pale brown to dark

brown and not sharply contrasted with brown of dorsum’

in utahickae. Based on these external characters and the

fact that no geographical origin is known, the species des-

ignation of the ZFMK specimen remains questionable. The

results of our genetic study of this specimen is shown in

Fig. 4. Except of relationships among Cacajao , Pithecia

and Chiropotes , which cannot be correctly resolved based

on the short cyt b fragment using the ML and Bayesian

approaches, we obtained the same pitheciine phylogeny

as Bonvicino et al. (2003). Chiropotes albinasus forms a

basal taxon to all remaining Chiropotes members and C.

utahickae forms a sister clade to C. israelita. However,

irrespective of the phylogenetic approach and of uncer-

tainties among basal pitheciine relationships, our sample

from the ZFMK clusters together with the C.

chiropotes! sagulatus sample from east of the Rio Branco

causing paraphyletic relationships between C. israelita

(sensu Bonvicino et al. 2003; also Dare et al. 2011) and

C. chiropotes! sagulatus west and east of the Rio Branco,

respectively (Fig. 4).

Chiropotes satanas (Hoffmannsegg, 1807) -

Black bearded saki

Material: ZFMK_MAM_60. 1 08; juvenile male, f

1 1. IV. 1960, received from Cologne Zoo 20.VII.1960; taxi-

dermy mount. The animal died at the zoo shortly after its

arrival. A photo each of a juvenile individual of Chiropotes

satanas from Cologne Zoo published in Hill ( 1960) and

in Anonymous (1963) may be this animal but it may al-

so show one of the other two juveniles of this species

which the zoo received in 1954 resp. 1959, each of each

survived for a short time only.

Description of the specimen: With the exception of the

back which is covered with relatively long chestnut brown

hair, the hair-coat of all other body parts is black.

ZFMK_MAM_81.1819; adult male, | 04.VII.1981, re-

ceived from Cologne Zoo VIII. 1 98 1 ; skull, skeleton, taxi-

dermy mount. When this individual (house name ‘Nicky’)

arrived at the zoo as a young juvenile on 26.VI.1970 it

was originally registered as a female but this was correct-

ed in the archives in 1971. Its black-and-white portrait tak-

en soon after arrival was published (Windecker 1971 : 4).

No other photo published later of a bearded saki from

Cologne Zoo or one kept in the zoo archives can be re-

lated to this individual with certainty; therefore its char-

acters as an adult are not documented. It is the father of

all hybrids of Chiropotes satanas and Chiropotes albina-

sus females bom at Cologne Zoo between 1973 and 1978

(see above).

Description of specimen ZFMK MAM 8I.1819 (Fig.

5): As the specimen was on exhibit in the Museum for

nearly two decades, the same general qualifications as for

the Chiropotes chiropotes specimen ZFMK_MAM_

66.001 apply in respect of an assessment of the coat colour

of this individual. Moreover, it lost hair in the tanning

process, especially on the back. This is likely to result in

a changed perception of the colour of the back coat as

compared to that of a living animal. Generally, there is no

sharp contrast in coat colouration between different parts

of the body. The colour of the head (with beard and split

tuft of hair on top of it), nape, tail, extremities and backs

of hands and feet is black as is that of the venter. On the

back, shoulders, upper arms and thighs a lighter dull brown

shines through the blackish hue, dominating the general

colour impression of the back.

Whereas the last in-depth revision of the genus Chi-

ropotes by Hershkovitz (1985) listed two subspecies of

Chiropotes satanas in addition to the nominate form, the

current notion is that Chiropotes satanas represents a dis-

tinct species with no subspecies (Silva-Jr. & Figueiredo

2002; Bonvicino et al. 2003; Groves 2005; Figueiredo et

al. 2006; Rylands & Mittermeier 2009; Ferrari et al. 2013;

Fig. 5. Back view of mounted adult male Chiropotes satanas

specimen ZFMK MAM 81.1819. The hair on the back is thin-

ner than in the living animal (Photo: M. Weigt).

Bonn zoological Bulletin 63 (2): 173-187

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Gustav Peters et al.

Silva-Jr. et al. 2013). As detailed above, the two remain-

ing former subspecies and an additional one are accord-

ed species rank each as well. We adopt this concept here.

As in the case of Chiropotes chiropotes we have to rely

on the species determinations in the Cologne Zoo

archives for those individuals for which no detailed doc-

umentation and/or collection material is available, appre-

ciating the fact that these may have been wrong, especial-

ly in young juveniles which are difficult to identify as to

species in Chiropotes satanas sensu lato. Between 1954

and 1970 Cologne Zoo received 4 individuals of Chi-

ropotes satanas , all as young juveniles: two males, one

female and one individual the sex of which was not es-

tablished with certainty and which died soon after its ar-

rival. Only the male individual received by the zoo in 1970

lived to adulthood.

Concluding taxonoinical remarks on

Chiropotes specimens in the ZFMk collection

Chiropotes chiropotes ZFMk MAM 66.0111: We con-

sider this species identification to be well-founded. In con-

sideration of the qualifications of such an appraisal the

available evidence still documents an extensive similari-

ty of this ZFMK specimen with the type specimen of

Brachyurus israelita Spix, 1823 and the specimen referred

to as Chiropotes cf. satanas in a colour photograph in Bou-

bli (2002), both of which are synonyms of Chiropotes chi-

ropotes.

Chiropotes sp. ZFMKMAM 2008.244: Our cyt b phy-

logeny shows a close genetic relationship of this speci-

men to an individual from Rio Trombetas east of the Rio

Branco, i.e. within the distribution of C. sagulatus accord-

ing to Ferrari et al. (2013) and Silva-Jr. et al. (2013). How-

ever, their description of sagulatus is ‘Dorsum and upper

limbs ... are orange to reddish-brown. Head, nape, low-

er arms and legs are blackish’ (Ferrari et al. 2013: 481),

which does not fit to the general appearance of the ZFMK

specimen. Despite the small amount of samples analysed,

the unknown origin of the ZFMK specimen and of the low

support values in reconstructed phylogenies, our data al-

low us to draw some preliminary conclusions. Our results

support the existence of different colour variants within

groups of closely related individuals and possibly within

the same geographic region, here within the Chiropotes

form occurring east of Rio Branco. This finding corre-

sponds to the statement by Hershkovitz (1985) that no

clear geographic separation between different pelage vari-

ants exists. Furthermore, our molecular results indicate

that most probably both israelita and sagulatus do not de-

serve species status and may represent synonyms of Chi-

ropotes chiropotes. Therefore, we highly recommend fur-

ther genetic studies on diverse Chiropotes samples from

a variety of geographical areas, including Chiropotes sa-

tanas to clarify the phylogeny of the genus, and most im-

portantly, to allow final taxonomic conclusions.

Chiropotes satanas ZFMk_MAM_60.108: There is very

little reliable information on coat colour of juvenile indi-

viduals of Chiropotes satanas sensu lato. To the best of

our knowledge no such data have been published for ju-

venile Chiropotes satanas sensu stricto; based on the rel-

evant details provided for Chiropotes chiropotes (Her-

shkovitz 1985: 17) it seems possible that this ZFMK in-

dividual actually represents the latter species.

Chiropotes satanas ZFMk_MAM_81.1819: In spite of

the reservations with respect to the assessment of the fur

colouration of the living animal, in light of the condition

of this taxidermy mount we consider this species identi-

fication to be well-founded.

Pithecia Desmarest, 1804

The first revision of the genus Pithecia by Hershkovitz

(1979) recognized four species, a concept also followed

by Buchanan et al. ( 1981 ). The current taxonomy of this

genus as adopted by Groves (2001, 2005), Rylands & Mit-

tenneier (2009) and Ferrari et al. (2013) is based on the

earlier and more detailed revision by Hershkovitz ( 1987b)

in which he recognized five species. However, there is al-

so evidence that a further revision of this genus is required

(Marsh 2004, 2006, pers. comm. 2012). In her final revi-

sion (Marsh 2014) 16 species are recognized.

Pithecia mittermeieri Marsh, 2014 -

Mittermeier’s Tapajos saki

Hershkovitz ( 1987b: 427, Fig. 25) figured an adult male

Pithecia irrorata irrorata (house name Sascha) from

Cologne Zoo. Since its arrival at Cologne Zoo in 1 966 un-

til its death on 20.1.1981 it was registered as Pithecia

monachus in the Zoo archives. Hick ( 1 968b) published its

black-and-white portrait and later on another

photograph/portrait in colour (Hick 1973) under this name.

The best photographic documentations of this individual

referred to as Pithecia monachus are two black-and-white

photographs and a colour photo in Tylinek and Berger

(1984) and colour photos in several official guides of

Cologne Zoo in the 1970es (Fig. 6). According to Weigl

(2005), who listed this animal under the scientific name

Pithecia irrorata irrorata , it reached the highest longevi-

ty known of any individual of this species in human care

(more than 14 years and 7 months). In the recent revision

of the genus this individual is identified as Pithecia mit-

termeieri sp. nov. (Marsh 2014: 5, Table 2). Regrettably

no zoological material of it is preserved in the ZFMK, an-

other zoological museum or a scientific institution.

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Neotropical primates from the Cologne Zoo in the collections of the Zoologisches Forschungsmuseum A. Koenig 183

Fig. 6-7. 6. (Left) Adult male Pithecia mittermeieri Marsh, 2014 in the Zoo Cologne; this animal may be the one shown in fig-

ure 7 (Photo: P. De Prins, reproduced from 'Wegweiser durch den Zoo Koln’, c. 1976). 7. (Right) Adult male specimen of Pithe-

cia aequatorialis (lying individual on the left) (ZFMK_MAM_81.1818) in the Zoo Cologne ( 1973 or later); the individual sitting

on the right may represent Pithecia mittermeieri , possibly the same animal as shown in figure 6 (Photo: G. Peters).

Pithecia aequatorialis Hershkovitz, 1987 -

Equatorial saki

Material: ZFMK_MAM_8 1.1 818; adult male, f 1981 (?),

received from Cologne Zoo VI 11.1981; skull, postcranial

skeleton, flat skin (Fig. 7, the living animal).

This specimen has the diagnostic pelage characteristics of

Pithecia aequatorialis. It was received from Cologne Zoo

as Pithecia monachus and classified as such in the ZFMK

collection then. After the publication of the second revi-

sion of the genus Pithecia by Flershkovitz (1987b) the

species determination of this specimen was reassessed and

subsequently classified as Pithecia aequatorialis. This

identification was confirmed by L. Marsh (pers. comm.

2011) on the basis of several colour photos of the ZFMK

skin. Unfortunately, there is no photograph of the living

animal or written record in the Cologne Zoo archives

which can be unambiguously assigned to this individual

(SpieB & Becker, pers. comm. 2011). However, a photo-

graph by R. Mittermeier (in Marsh 2014: fig. 49) may

show this individual.

Several individuals classified as Pithecia monachus

lived at Cologne Zoo during the second half of the 1970’s.

Apart from the male individual of Pithecia mittermeieri

mentioned above (and then wrongly classified as P.

monachus at the zoo) none of the black-and-white pho-

tographs of ‘ Pithecia monachus' individuals archived at

the zoo shows an animal with the typical pelage charac-

teristics of Pithecia aequatorialis males. In the five year

period from 1977 until 1981 four individuals classified as

P. monachus died at Cologne Zoo, one of which was the

misidentified P. mittermeieri (Becker, pers. comm. 2012).

The only photograph on file of a "Pithecia monachus' in-

dividual at the zoo which died between 1977 and 1981

(other than the P.mittermeieri male mentioned) is one tak-

en in 1969 of a "monachus' that died in 1979. So, there

is definitely no ‘trace’ of the ZFMK Pithecia aequatori-

alis adult male in the Cologne Zoo archives. A colour slide

(Fig. 7) taken by one of us (G.R) shows this individual

alive (or another male of this species if there were more

than one in the zoo at that time) and therefore is addition-

al proof of its species identification (confirmed by L.

Marsh, pers. comm. 2013, based on this slide). The pho-

to can be roughly dated to the time period 1973 to 1977

because it was taken in the Lemur House of Cologne Zoo

which opened in 1973, and on written entries in the ani-

mal records at the zoo.

This species was described as new to science after all

Pithecia individuals (other than P. pithecia ) at Cologne

Zoo had been dead already for several years. In his ear-

lier revision of the genus Hershkovitz (1979: 15, fig. 5)

had figured a study skin of an adult male Pithecia as P.

monachus , only to revise his taxonomic assessment of the

same individual in the later revision and identify it as the

new species Pithecia aequatorialis (Hershkovitz 1987b:

408, fig. 16). Diagnostic characteristics differentiating

males of aequatorialis from male monachus are a ruff of

orange hair and a horseshoe-like area of relatively short

white dense hair around the sparsely haired facial disk in

the former species (L. Marsh, pers. comm. 2011, Marsh

2014).

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184

Gustav Peters et al.

CONCLUDING REMARKS

Our paper is based on only a few Neotropical primate

specimens from Cologne Zoo in the ZFMK mammal col-

lection. Nevertheless, it provides an example of the val-

ue and of the conclusions that can be drawn from collec-

tions obtained from zoos and housed at zoological muse-

ums (or similar scientific institutions). The ZFMK re-

ceived also primate specimens from other zoos, such as

Duisburg and Frankfurt (see Appendix), which may be

equally important. In the first place it shows that even to-

day zoos may have scientifically important specimens in

their care, sometimes unnoticed by the scientific commu-

nity. To avoid such cases as far as possible it is desirable

that zoos and zoological museums intensify their cooper-

ation (cf. Gippoliti & Kitchener 2007). It is important that

zoos are aware of the potential scientific importance of

specimens in their care and try to gather and archive all

pertinent information during the lifetime of such speci-

mens: geographical origin (if known or if it can be traced

back), colour photos at different ages, body mass, meas-

urements, reproductive history, ISIS reports, CITES pa-

pers (if applicable), references of publications dealing with

the specimen (if the specimen is mentioned/figured in a

publication), or other. This is, however, more than usual-

ly recorded. It may be useful to agree on a uniform data-

base, so that information from different zoos can be more

easily combined.

Taking into account relevant veterinary and other legal

regulations the most reasonable procedure would be that

zoos pass such specimens immediately after their demise

to a zoological museum. The condition of the specimen

should thereby allow all standard museum procedures,

e.g., collection of tissue and hair samples, preparation of

skin and skeleton or any other method of conservation. Ap-

propriate storage of such samples should allow subsequent

genetic analyses. Furthermore, copies of all documenta-

tion of that specimen compiled at the zoo during the an-

imal’s lifetime should be provided, including veterinary

treatments. A scientifically important specimen in/from a

zoo can only maintain its ‘voucher status’ and its signif-

icance posthumously as a subject for later studies, if its

remains are housed and curated properly in a zoological

museum or another appropriate scientific institution in the

long run. This ought to be in the genuine interest of the

respective zoo and definitely is so for the scientific com-

munity, especially if specimens have been referred to in

the scientific literature like several Chiropotes individu-

als from Cologne Zoo in the ZFMK mammal collection,

which are listed here. The Pithecia mittermeieri individ-

ual discussed above exemplifies the deplorable ultimate

loss of information and research options if this procedure

is not followed after an animal’s death. On the other hand,

the specimen of Pithecia aequatorialis in the ZFMK col-

lection detailed before illustrates that steady contact and

information exchange between zoos and zoological mu-

seums may have resulted in the timely awareness of the

significance of this individual, especially in the context

of a taxonomical revision of the genus Pithecia.

Zoological museums with their taxonomic expertise

ought to interact with (neighbouring) zoos regularly, in-

cluding the offer to provide advice on matters of taxono-

my if requested. Once a specimen that died in a zoo is giv-

en to a museum, the museum has the responsibility of its

proper long-term documentation and curation, preserving

as much and diverse material of the animal as possible to

enable diverse kinds of (future) studies. This includes the

repeated verification of a specimen’s species identity on

the basis of the most recent pertinent publications to en-

sure its scientific significance. If a specimen is affected

by a taxonomical change, either during its lifetime or

posthumously, this information ought to be circulated

among the institutions involved.

Both zoos and zoological museums can only benefit

from a closer cooperation as a means to increase taxonom-

ic expertise and awareness of taxonomical problems as

well as the proper long-term conservation of scientifical-

ly interesting and important zoological specimens.

Acknowledgements. We are grateful to the many colleagues

who generously helped in gathering all the diverse information

necessary to carry out this study. Our sincere thanks to Wilhelm

SpieB and Ralf Becker (both Cologne Zoo archives) and Uta

Ruempler (nee Hick) (formerly Cologne Zoo) for providing in-

formation on the zoo’s relevant primate individuals and photos,

Andrew Kitchener (National Museums Scotland, Edinburgh) and

Ms. E.M. Lajtos (Library, Naturalis, Leiden) for help with lit-

erature, Laura Marsh (Global Conservation Institute) for expert-

ise in Pithecia identification, Richard Kraft and Michael Hier-

meier (ZSM) for information on and photos of the type speci-

men of Brachyurus israelita. Laura Parker and Wendy Lehkyi

(both Colchester Zoo), Tremaine Gregory (National Zoo, Wash-

ington, D.C.), Marilyn Norconk (Kent State University, Ohio),

and Liza Veiga (deceased) kindly provided photos of living Chi-

ropotes individuals and information. We are indebted to

Melanie Weigt (ZFMK) for photography and to Jorg Adrian

(ZFMK) for technical assistance, to Achim Winkler (Duisburg

Zoo) for information on Pithecia individuals kept at that zoo,

and to Carlos Ruiz Miranda (Universidade Estadual Norte Flu-

minense) for help with contacts in Brazil. We also thank Michael

Hofreiter for giving us the opportunity to conduct the genetic

analysis of the Chiropotes sample in his ancient DNA laborato-

ry at York University. We are also grateful for helpful comments

and suggestions of the two reviewers.

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APPENDIX

List of Pitheciidae in the collections of ZFMK Bonn.

Cacajao calvus. ZFMK MAM 66.612 (Cologne Zoo 1966);

ZFMKM AM_8 1.1816 (Cologne Zoo 1981);

ZFMK MAM 200 1 147 (Cologne Zoo 1993);

ZFMKM AM_20 1 2.427 (Cologne Zoo 2008).

Cacajao hosomi. ZFMK MAM 2002.032

(Venezuela/Cologne Zoo 1992).

Chiropotes albinasus. ZFMK MAM 7 1.097 (Cologne Zoo

1971); ZFMK_MAM_8 1.1815 (Cologne Zoo 1981);

ZFMK MAM 89.480 (Cologne Zoo 1989).

Chiropotes chiropotes. ZFMK MAM 66.001 (Brazil/

Cologne Zoo 1961). ZFMK_MAM_74.155 (Duisburg Zoo

1974).

Chiropotes satanas. ZFMK MAM 60.108 (Cologne Zoo

1960); ZFMK_MAM_ 81.1819 (Cologne Zoo 1981).

Chiropotes sp. ZFMK MAM 2008. 244 (Cologne Zoo).

Pithecia aequatorialis. ZFMK MAM 8 1.1818, male

(Peru/ Cologne Zoo 1981).

Pithecia inusta. ZFMK_MAM_ 87.730, female

(Peru, Panguana 1973; Hutterer et al. 1995).

Pithecia pithecia. ZFMK MAM 71 .097, male (Cologne Zoo

1971); ZFMK_MAM_20 12.009, male

(Frankfurt Zoo 2008).

Pithecia sp. ZFMK MAM 75.108, female

(Duisburg Zoo 1975).

Bonn zoological Bulletin 63 (2): 173-187

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Abbreviations Except of very common abbreviations such as mm,

kg etc.; all abbreviations should be explained in the Methods sec-

tion or figure legend if appropriate. Hyphenation or upper case let-

ters for entire words are not permitted. All measurements must be

metric units and given to the same decimal, i.e., 5. 3-6.0 mm (not:

5.3-6 mm).

9

Instructions to authors

Taxonomy Names of animals and the description of new genera or

species must follow the current version of the International Code

of Zoological Nomenclature (ICZN, available at

http://www.iczn.org/iczn/index.jsp). Type specimens should be de-

posited in recognised institutions; deposition at ZFMK is highly ap-

preciated. Italics are required for species names which are written

in full the first time they appear in the text, e.g. Carassius auratus

(Linnaeus, 1 758), but abbreviated at subsequent mention, e.g. C. au-

ratus. List names in synonymies as follows: Attelabus asparagi

Scopoli, 1763 (Scopoli 1763: 36, fig. 1 13.), and list the source un-

der References. Dichotomous keys are desirable in taxonomic pa-

pers.

Statistics Statistics presented should include the name of the test,

number of observations or degree of freedom, and probability lev-

el (P > 0.05, P < 0.05, P > 0.001, P < 0.001). Values of test statis-

tics are not required.

Scientific Notes

Notes should comply with the instructions given above, but have to

be ( 1) confined to a single point or issue of progress, (2) must be

short (typically no more than 3 printed pages including illustrations

and references), and (3) do not include headings or subheadings. A

brief abstract of no more than three lines is required.

Review papers

Reviews should be concise, critical and creative, seeking to sum-

marize the state-of-the-art of complex topics, and stimulate debates

and new research initiatives. Review papers should comply with the

instructions for preparation of Research articles, with the exception

that the main body of text may be structured as appropriate.

Electronic submission

Manuscripts should be submitted as text files (*.doc or * .rtf; *.pdf

is welcome for the review process, but for production *.doc or *.rtf

will be required) via e-mail (bzb.zfmk@uni-bonn.de) to the editor-

in-chief. Alternatively, send a CD. If neither is possible for any rea-

son, please contact the editor-in-chief. The file name must start with

the submitting author’s last name.

Review process

Each manuscript will be peer-reviewed, generally by two or more

referees. Manuscripts should be submitted to the editor-in-chief, and

will be handled by the editor dealing with the respective taxon or

topic (see list of editors). Authors are encouraged to suggest poten-

tial referees for their manuscripts. Final acceptance for publication

is in the responsibility of the editor-in-chief. This decision is main-

ly based on the referees’ report and the handling editor’s recommen-

dation. Revised manuscripts received more than 6 months after the

reviewers’ comments had been sent will be treated as new submis-

sions. After acceptance, authors are encouraged to submit sugges-

tions for a cover image.

Ethical and legal aspects

Authors are obliged to state clearly that they followed the legal reg-

ulations and laws for the collection and ethical treatment of animals.

Publication and open access policy

Manuscripts will be published online at www.zoologicalbulletin.de

after acceptance as preprint version, which will be replaced by the

final pdf after publication of the printed issue of BzB. The corre-

sponding author must clearly state so if preprint online publication

is not desired. In that case, only title, authors and the infonnation

that this manuscript has been accepted for publication will be made

available at that stage. Immediately after publication, the correspon-

ding author will receive an e-mail containing the final pdf. Reprints

can only be purchased on the author’s cost. Reprint orders must be

submitted to the editor at latest with the approval of the final proofs.

All published content is freely available without charge to the user

or his/her institution. Authors are invited to place the final pdf on

their personal or institutional website.

Questions we ask our reviewers

Authors preparing manuscripts for BZB should keep in mind the fol-

lowing questions which referees for the journal are asked:

1 . Is the paper of interest to the readers of BZB?

2. How well does it meet the focus of BZB?

3. Is it acceptable for publication, or should it be reconsidered af-

ter revision?

3. Please grade originality, quality of data, quality of statistical analy-

ses, and quality of interpretation on a scale from 1 = low to 5 =

high.

4. Please grade clarity and quality of text, figures and tables from

1 = low to 5 = high.

5. Has the welfare of any experimental animals been adequately tak-

en into account?

6. Does the title aptly and correctly describe the paper?

7. Does the abstract summarize only the significant findings?

8. Is the length of the paper appropriate?

9. Are all (colour) figures and tables necessary and appropriate?

Contents

3 9088 01798 0780

Grebennikov, Vasily V.:

Morimotodes, a new genus for two minute wingless litter species from

southwest China and Taiwan with an illustrated overview of Molityna and Plithina genera

(Coleoptera: Curculiomdae: Molityni)

Nguyen, Truong Quang, Trung My Phung, Nicole Schneider, Andreas Botov,

Dao Thi Anh Tran & Thomas Ziegler:

New records of amphibians and reptiles from Southern Vietnam

Liu, Wan-Gang, Ming Bai, Xing-Ke Yang & Dirk Ahrens:

Towards an improved knowledge of Sericim of the Tibetan highland:

new species and records (Coleoptera: Scarabaeidae)

Peters, Gustav, Tanja Haus & Rainer Hutterer:

Neotropical primates from the Cologne Zoo in the collections of the

Zoologisches Forschungsmuseum Alexander Koenig:

noteworthy specimens, taxonomic notes and general considerations

123

148

157

173

Cover illustration:

White-faced saki Pithecia pithecia (Photo: Rolf Schlosser)

(this volume, pp. 173-187).

Bundesministerium

fur Bildung

und Forschung

Ministerium fur Innovation,

Wissenschaft und Forschung

des Landes Nordrhein-Westfalen