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Bonn Index 2013

Volume 62

zoological sue 12

Bulletin

formerly: Bonner zoologische Beitrage

| Bonn Volume 62

| zoological Sais | | |

aan Editor-in-Chief

\ \ . Fabian Herder

Editorial Board

Dirk Ahrens

= Wolfgang Bohme

Jan Decher

Bonn Volume 62 .

zoological Bera Netta Dorchin

Bulletin

| Renate van den Elzen

Bernhard A. Huber

Rainer Hutterer

Gustav Peters

Bradley Sinclair

Dieter StUning

Till Topfer

Philipp Wagner

An open access journal of organismal zoology, published by

Zoologisches Forschungsmuseum Alexander Koenig — Leibniz-Institut flr Biodiversitat der Tiere, Bonn

Contents Volume 62

Assing, Volker: 1-29

On the Nazeris fauna of China I.

The species of the Qinling Shan, the Daba Shan, and adjacent mountain ranges

(Coleoptera: Staphylinidae: Paederinae) 7 -

Assing, Volker: 30-91

On the Lathrobium fauna of China I.

The fauna of the Qinling Shan, the Daba Shan, and adjacent regions (Coleoptera: Staphylinidae: Paederinae) oe

Assing, Volker: 125-170

On the Nazeris fauna of China Il.

New species and records from Zhejiang, Sichuan, and Yunnan (Coleoptera: Staphylinidae: Paederinae)

Bauer, Aaron M. & Richard Wahlgren: 220-252

On the Linck collection and specimens of snakes figured by Johann Jakob Scheuchzer (1735) —

in the oldest fluid-preserved herpetological collection in the world? ; ee

Jendek, Eduard & Hong-Thai Pham: 111-123

Seven new species of Coomaniella Bourgoin, 1924 (Coleoptera: Buprestidae)

with redefinition of species-groups and remarks on distribution and biology

Kerbis Peterhans, Julian C., Jakob Fahr, Michael H. Huhndorf, Prince Kaleme, 186-202

Andrew J. Plumptre, Ben D. Marks & Robert Kizungu:

Bats (Chiroptera) from the Albertine Rift, eastern Democratic Republic of Congo,

with the description of two new species of the Rhinolophus maclaudi group ; 7

Kerbis Peterhans, Julian C., Michael H. Huhndorf, Andrew J. Plumptre, Rainer Hutterer, 203-219

Prince Kaleme & Benjamin Ndara:

Mammals, other than bats, from the Misotshi-Kabogo highlands (eastern Democratic Republic of Congo),

with the description of two new species (Mammalia: Soricidae) 7 ee

Lehmann, Ingo & Hossein Rajaei Sh.: 100-110

Description of a new genus and three new species of Metarbelidae (Lepidoptera: Cossoidea)

from East and Central Africa, with notes on biography -

Sinclair, Bradley J. & Christel Hoffeins: 92-99

New fossil species of Ragas Walker (Diptera: Empididae) in Baltic amber (Tertiary, Eocene) a

Stiels, Darius & Kathrin Schidelko: 171-176

Records of ‘Indian’ Baya Weaver Ploceus philippinus (Linnaeus, 1766) and

Hooded Wheatear Oenanthe monacha (Temminck, 1825) from Afghanistan (Aves: Passeriformes) 7

Tuyisingize, Deogratias, Julian C. Kerbis Peterhans, Gary N. Bronner & Tara S. Stoinski: 177-185

Small mammal community composition in the Volcanoes National Park, Rwanda

Publication dates:

Vol. 62 Issue 1: May 2013

Vol. 62 Issue 2: December 2013

Bonn zoological Bulletin 62 (2): 125-170

SMITHS ON/aE

December 2013

DEC 02 2014

LIBRARIES

On the Nazeris fauna of China II.

New species and records from Zhejiang, Sichuan, and Yunnan

(Coleoptera: Staphylinidae: Paederinae)

Volker Assing

Gabelsbergerstr. 2, D-30163 Hannover, Germany, E-mail: vassing.hann@t-online.de.

Abstract. Material of the Palaearctic paederine genus Nazeris Fauvel, 1873 from the Chinese provinces Zhejiang, Sichuan,

and Yunnan is revised. Thirty species are described and illustrated, eight of them from Sichuan and the remainder from

Yunnan: Nazeris bilamellatus sp. n. (Sichuan: Emei Shan); NV. foveatus sp. n. (Sichuan: Erlang Shan); N. appendiculatus

sp. n. (Sichuan: Daxue Shan); N. puetzi sp. n. (Sichuan: Erlang Shan); N. cornutus sp. n. (Sichuan: Erlang Shan); N. tri-

furcatus sp. n. (Sichuan: Xiaoxiang Ling); N. wrasei sp. n. (Sichuan: Xiaoxiang Ling); N. schuelkei sp. n. (Sichuan: Laba-

he, Daxiang Ling); NV. wu/iangicus sp. n. (Yunnan: Wuliang Shan and adjacent mountain range); NV. nivimontis sp. n. (Yun-

nan: Xue Shan N Lincang); N. sagittifer sp. n. (Yunnan: Ailao Shan, Wuliang Shan); N. secatus sp. n. (Yunnan: Laobie

Shan); NV. cangicus sp. n. (Yunnan: Diancang Shan); N. acu/eatus sp. n. (Yunnan: Jizu Shan); N. zhemoicus sp. n. (Yun-

nan: Zhemo Shan); N. pungens sp. n. (Yunnan: Xue Shan ENE Lincang); N. /anuginosus sp. n. (Yunnan: Laobie Shan);

N. peniculatus sp. n. (Yunnan: mountain range N Er Hai); N. barbatus sp. n. (Yunnan: Wuliang Shan); N. firmilobatus

sp. n. (Yunnan: Gaoligong Shan); N. spiculatus sp. n. (Yunnan: mountain range E Luxi); N. curvus sp. n. (Yunnan: Gaoligong

Shan); N. infractus sp. n. (Yunnan: Nu Shan); NV. subdentatus sp. n. (Yunnan: Gaoligong Shan); N. meilicus sp. n. (Yun-

nan: Meili Xue Shan); N. vexillatus sp. n. (Yunnan: Gaoligong Shan); N. circumclusus sp. n. (Yunnan: Gaoligong Shan);

N. hastatus sp. n. (Yunnan: Gaoligong Shan); N. bangmaicus sp. n. (Yunnan: Bangma Shan); N. fissuws sp. n. (Yunnan:

Ailao Shan, Wuliang Shan). Records of 17 additional species are reported. Seven of them are represented exclusively by

females and probably undescribed, and the remainder described previously; the latter are from Zhejiang (two species),

Sichuan (four species), and Yunnan (four species). The male sexual characters of some of the previously described species

are illustrated. Species group affiliations are discussed. The distributions of 37 species are mapped. The genus now in-

cludes a total of 171 species and seven subspecies; 94 species have been reported from China (exclusive of Taiwan).

Key words. Taxonomy, Staphylinidae, Paederinae, Nazeris, China, Zhejiang, Sichuan, Yunnan, new species, species groups,

distribution maps, endemism.

INTRODUCTION

According to a recent contribution (Assing 2013a), the

paederine genus Nazeris Fauvel, 1873 has an essentially

Palaearctic distribution and previously comprised a total

of 143 species and seven subspecies. As many as 66

species had been recorded from China (exclusive of Tai-

wan); two of them, however, are synonymous (see below).

All the Nazeris species are flightless and, with one excep-

tion (N. cultellatus Assing, 2013), the species from Chi-

na are locally endemic. In China, Nazeris ranks second

among the paederine genera with respect to the diversity

of micropterous species with restricted distributions, out-

numbered in described species only by Lathrobium

Gravenhorst, 1802, which is currently represented in Chi-

na by 170 described species, most of them locally endem-

ic (Assing 2013b, c, in press; Peng et al. 2013b). The

provinces with the greatest diversity of previously de-

scribed Nazeris species are Zhejiang (13 species), Yun-

nan (11), and Shaanxi (11), followed by Sichuan (9),

Guangxi (6), Anhui (4), Gansu (3), Hubei (3), Chongqing

(3), Jiangxi (2), Fujian (2), Xizang (2), Guizhou (1), and

Received: 21.08.2013

Accepted: 01.10.2013

Henan (1). The sum of species exceeds 64 in this overview

because five species have been reported from more than

one province or from the border region between two or

three provinces.

In all, 15 Nazeris species have been described from Zhe-

jiang province (Assing 2013a; Hu et al. 201 1a; Ito 1996;

Koch 1939). However, according to a revision of type ma-

terial by Hu (e-mail 12.7.2013), two of the names made

available in Hu et al. (201 1a) are junior synonyms, so that

the Nazeris fauna of Zhejiang currently comprises 13

species.

The nine Nazeris species previously known from

Sichuan were recorded from Wolong (one species), the

Gongga Shan (two species), the Emei Shan (two species),

the Micang Shan (one species), the Erlang Shan (one

species), Labahe (one species), and from the Qingcheng

Shan (one species whose description is based on a single

female) (Assing 2013a; Hu et al. 2007; Zheng 1992). Re-

garding the type locality of one of these species, N.

canaliculatus Zheng, the original description (Zheng 1992)

Corresponding editor: D. Ahrens

126 Volker Assing

is contradictory. According to the Chinese description, all

the type specimens are from the Emei Shan, whereas ac-

cording to the English abstract, the holotype is from Wo-

long and the two female paratypes are from the Emei Shan.

Most likely, the type locality given in the abstract is er-

roneous.

The Nazeris fauna of Yunnan previously comprised

eleven species, four from the Gaoligong Shan, two from

the Diancang Shan and the Laohu Shan, two from the Na-

banhe Nature Reserve, and one each from the Yulongxue

Shan, the Jizu Shan, and the Yuan Shan near Kunming (Hu

et al. 2011b; Watanabe & Xiao 1993, 1997, 2000).

Recent studies on the Chinese Lathrobium fauna re-

vealed that the diversity in Yunnan (60 described species)

is far greater than that of any other province and that

Sichuan (39 described species) ranks second (Assing

2013b, c; Peng et al. 2013a); among the remaining

provinces the Lathrobium fauna of Shaanxi (20 described

species) is the most diverse, but still hosts much fewer

species than Yunnan and Sichuan. Since both Lathrobi-

um and Nazeris have a similar distribution in the East

Palaearctic and are often found in the same habitats, it

seemed likely that the diversity trends observed for Lath-

robium should also apply to Nazeris.

The present paper is based primarily on outstanding ma-

terial from the collection of Michael Schiilke (Berlin),

most of which was collected by him and David Wrase

(Berlin) during nine field trips to China in the past two

decades. Additional material came from the collections of

Ales Smetana (Ottawa) and Andreas Piitz (Eisenhiitten-

stadt).

MATERIAL AND METHODS

The morphological studies were conducted using a Ste-

mi SV 11 microscope (Zeiss Germany) and a Jenalab com-

pound microscope (Carl Zeiss Jena). A digital camera

(Nikon Coolpix 995) was used for the photographs. The

maps were created using MapCreator 2.0 (primap) soft-

ware.

Body length was measured from the anterior margin of

the mandibles (in resting position) to the abdominal apex,

the length of the forebody from the anterior margin of the

mandibles to the posterior margin of the elytra, head length

from the anterior margin of the frons to the posterior mar-

gin of the head, elytral length at the suture from the apex

of the scutellum to the posterior margin of the elytra, and

the length of the aedeagus from the apex of the ventral

process to the base of the aedeagal capsule. The “para-

meral” side (1.e., the side where the sperm duct enters) is

referred to as the ventral, the opposite side as the dorsal

aspect.

For a discussion of the terminology of the aedeagal mor-

phology see Assing (2009).

Bonn zoological Bulletin 62 (2): 125—170

COLLECTION MATERIAL DEPOSITORIES

CAS — Chinese Academy of Sciences, Beijing

SMNS _ Staatliches Museum ftir Naturkunde, Stuttgart

(W. Schawaller)

ZFMK_ Zoologisches Forschungsmuseum Alexander

Koenig, Bonn

cAss author’s private collection

cPit private collection Andreas Piitz, Eisenhitittenstadt

cSch private collection Michael Schiilke, Berlin

cSme private collection Ales’ Smetana, Ottawa

RESULTS

Diversity and zoogeography

The revised material is composed of 47 species, ten of

them described previously from Zhejiang (two species),

Sichuan (four species), and Yunnan (four species). The lo-

calities where the material was collected are mapped in

Fig. 1. Thirty species are newly described, eight from

Sichuan and twenty-two from Yunnan. Seven additional,

probably undescribed species remain unnamed since they

were represented exclusively by females. Thus, including

the new species described in this paper, Nazeris compris-

es a total of 171 species and seven subspecies, with 94 of

them recorded from, and confined to, China. In all, 33

species are now known from Yunnan and 16 from Sichuan,

the provinces with the most diverse Nazeris faunas. The

diversity trends previously observed for Lathrobium are

indeed paralleled by Nazeris.

Except for one species (N. ruani), all the species from

Sichuan and Yunnan are locally endemic and known on-

ly from individual or adjacent mountain ranges.

As was observed earlier for the fauna of the Qinling

Shan and the Daba Shan (Assing 2013a), closely related

species, particularly hypothesized adelphotaxa, are gen-

erally at the same time geographically close, suggesting

local speciation events. However, identifying phylogenet-

ic relationships is significantly more difficult in Nazeris

than in Lathrobium, particularly because the external and,

even more importantly, the male sexual characters are less

diverse. Nevertheless, it was possible to assign the ma-

jority of the species to species groups. For details see the

introductions to the regional faunas of Sichuan and Yun-

nan below. Species of one species group generally have

allopatric distributions (exceptions: N. truncatus and N.

bilamellatus of the N. truncatus group in the Emei Shan,

and N. puetzi and N. foveatus of the N. hailuogouensis

group in the Erlang Shan), but often two or three species

of different groups were recorded from the same moun-

tain, not infrequently collected even in the same sample.

OZFMK

On the Nazeris fauna of China II 127

Natural history

As can be inferred from the data specified on the labels

of the material examined and as has been observed already

for the Nazeris species of the Qinling Shan and the Daba

Shan (Assing 201 3a), the Nazeris species in the study re-

gions live in the leaf litter and moss of various forest and

shrub habitats. While the material from Zhejiang was col-

lected at relatively low altitudes (300-1200 m), that from

Sichuan and Yunnan came from significantly higher ele-

vations. In Sichuan, the altitudes range from 1250 to 3250

m, with the vast majority of specimens found between

1600 and 3000 m. Only one species, NV. wrasei, was ob-

served at a lower (1250 m), and one probably undescribed

species at a higher elevation (3250 m). In Yunnan, the al-

titudes range from 1900 to 3150 m, with only one species

(N. barbatus) collected below 2100 m.

Species from Zhejiang

Nazeris rougemonti Ito, 1996

Material examined. China: Zhejiang: 5, 9%, Tianmu

Shan, pass 25 km NNW Linan, 30°26’N, 119°36’E,

620-820 m, stream valley with bamboo and mixed for-

est, litter sifted, 16.V1I.2007, leg. Piitz, Schtilke & Wrase

(cPiit, cSch, cAss); 19, West Tianmu Shan N. R., trail to

peak of immortals, 30°21°N, 119°26’°E, 1100—1200 m, pri-

mary mixed forest, litter and moss sifted, 15.V1.2007, leg.

Piitz (cPiit); 24, Tianmu Shan, 40 km WNW Linan, reser-

voir, 30°21°N, 119°19’E, 300 m, plant refuse and litter sift-

ed, 17.V1.2007, leg. Schtilke (cSch, cAss).

Comment. The original description of N. rouwgemonti is

based on three type specimens from “Mt. Tienmushan”

(Ito 1996). The known distribution is confined to the Tian-

mu Shan.

Nazeris chinensis Koch, 1939

Material examined. China: Zhejiang: 2¢, 19, West

Tianmu Shan N. R., trail to peak of immortals, 30°21°N,

119°26’E, 1100-1200 m, primary mixed forest, litter and

moss sifted, 15.V1.2007, leg. Piitz, Schiilke & Wrase (cPiit,

cSch, cAss).

Comment. Nazeris chinensis has been recorded from the

Tianmu Shan and the Longwang Shan (Hu et al. 201 1a).

Bonn zoological Bulletin 62 (2): 125-170

Species from Sichuan

Species groups. Primarily based on the male sexual char-

acters, the Nazeris fauna of Sichuan is represented by at

least six species groups.

Nazeris dilatatus Assing, 2013 from the Micang Shan

is the sole representative of the N. shaanxiensis group (see

Assing 2013a) in Sichuan.

The evidently widespread N. giganteus group includes

at least two species, NV. giganteus from Yunnan and N. ru-

ani from Sichuan. The species of this group are charac-

terized by conspicuously large body size, a moderately

deep V-shaped posterior excision of the male sternite

VUI, and a rather large aedeagus with a slender and api-

cally acute ventral process (ventral view) and with short,

stout, curved, and apically dilated dorso-lateral apophy-

ses. Based on the small median excision of the posterior

margin of the male sternite VH, a characters otherwise

seen only in N. ruani, the distinctly smaller N. alpinus

from northern Yunnan may belong to this group, too. Un-

like nearly all other Chinese representatives of the genus,

N. ruani 1s not endemic to a single mountain range, but

has a less restricted distribution (Fig. 28).

Nazeris magnus Hu et al., 2007 1s the only species

known from Sichuan that is of similarly large size as the

species of the N. giganteus group, but it has an aedeagus

of different morphology. Closer affiliations to other Naz-

eris species from Sichuan and Yunnan are not evident.

The N. truncatus group comprises six species from

Sichuan, N. truncatus, N. bilamellatus (both from the Emei

Shan), N. bicornis (Gongga Shan), N. cornutus (Erlang

Shan), N. trifurcatus (Xiaoxiang Ling), and N. folicaeus

Zheng, 1992 (Wolong). The species of this group are char-

acterized by an aedeagus with two or three apical appen-

dices of the ventral process of the aedagus, undoubtedly

a synapomorphy. The posterior excision of the male ster-

nite VIII is moderately deep, and the dorso-lateral

apophyses are moderately long (reaching apex of ventral

process at most, not extending beyond it).

The N. hailuogouensis group is represented in Sichuan

by four species, N. hailuogouensis (Gongga Shan), N.

foveatus (Erlang Shan), N. appendiculatus (Daxue Shan),

and N. puetzi (Erlang Shan). The species of this group

share a semi-membranous, short, ventrally sharply edged,

and apically V-shaped (ventral view) ventral process of

the aedeagus (e.g., Figs 5-6, 8—9, 13-14). The dorso-lat-

eral apophyses extend far beyond the apex of the ventral

process, and the posterior excision of the transverse male

sternite VII is rather deep and narrow (e.g., Figs 7, 12).

Nazeris hailuogouensis, N. foveatus, N. appendiculatus,

and N. puetzi are distinguished by clear-cut characters, but

otherwise extremely similar, even in the male sexual char-

acters. This observation suggests that speciation occurred

in evolutionarily more recent times. Similar examples of

closely related taxa have repeatedly been reported from

©ZFMK

128 Volker Assing

Chengd

Bee

Fig. 1.

nan (open circles). The frames illustrate the limits of the distribution maps for the individual species (Figs 28-29, 88-90).

the same region, e.g., for the genus Othius Stephens, 1829

(Staphylinidae: Othiini) (Assing 2003) or for some Lep-

idoptera (Lattin 1967). Nazeris bicornis and N. cornutus

of the N. truncatus group (see above) represent another

example of a pair of closely related Nazeris species oc-

curring in the Gongga Shan and the Erlang Shan (like N.

hailuogouensis and N. foveatus).

The N. schuelkei group includes two species, N.

schuelkei (Labahe, Daxiang Ling) and N. wrasei (Xiaox-

iang Ling). It is distinguished from the other species

groups distributed in Sichuan by a stout and short aedea-

gus with a laterally compressed ventral process and with

short and stout dorso-lateral apophyses (Figs 46-47,

52-53). The posterior excision of the male sternite VIII

is rather broad and moderately deep (Figs 45, 51). Regard-

ing the morphology of the ventral process of the aedea-

gus, the species of the N. schuel/kei group are similar to

the NV. cultellatus group, which is distributed from the Qin-

ling Shan eastwards to Henan and Anhui (see Assing

2013a).

The phylogenetic affiliations of N. gingchengensis

Zheng, 1992 from the Qingcheng Shan (male unknown)

and N. canaliculatus Zheng, 1992 from the Emei Shan

(poorly illustrated) are uncertain.

Bonn zoological Bulletin 62 (2): 125—170

Pooled examined records of Nazeris from the Chinese provinces Zhejiang (triangles), Sichuan (filled circles), and Yun-

Naczeris ruani Hu et al., 2007 (Fig. 28)

Material examined. China: Sichuan: 1, Emei Shan,

29°34°N, 103°21°E, 1800-2400 =m, sifted,

27.VI.—5.VI.2009, leg. Grebennikov (cAss).

Comment. This species was described from Labahe, some

120 km to the northwest of the Emei Shan, suggesting that

it is remarkably widespread (Fig. 28). The male sexual

characters of the above male are identical to those illus-

trated by Hu et al. (2007: figures 15—19).

Naczeris truncatus Zheng, 1992 (Fig. 28)

Material examined. China: Sichuan: 1¢, 49, Emei

Shan, 29°34’N, 103°21’E, 1950 m, sifted, 15.V1.2010, leg.

Grebennikov (CAS, cSme, cAss); 4¢, 82, Emei Shan,

29°34"N, =—:103°21’E, ~=—1800-2400 =m, _ sifted,

27.VI.—5S.VH.2009, leg. Grebennikov (CAS, cSme, cAss).

Comment. The known distribution of NV. truncatus, a

species characterized by an aedeagus of distinctive mor-

phology (see figures 3B—C in Zheng 1992), is confined

to the Emei Shan (Fig. 28).

©ZFMK

On the Nazeris fauna of China II 129

Figs 2-14. Nazeris bilamellatus (2-6), N. hailuogouensis (7-9), and N. foveatus (10-14). 2, 10: forebody; 3, 11: male sternite

VU; 4, 7, 12: male sternite VII; 5-6, 8-9, 13-14: aedeagus in lateral and in ventral view. Scale bars: 2, 10: 1.0 mm; 3—9, 11—14:

0.5 mm.

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

130 Volker Assing

Nazeris bilamellatus sp. n. (Figs 2—6, 28)

Type material. Holotype 3: “P.R. CHINA, Sichuan, Emei

Shan, N29°33.6’, E103°20.6’, 27.vi—5.vii.2009,

1800-2400 m, siftings 11-17, V. Grebennikov / Holoty-

pus & Nazeris bilamellatus sp. n. det. V. Assing 2013”

(CAS). Paratypes: 2%, 62: same data as holotype (CAS,

cSme, cAss).

Etymology. The specific epithet (Latin, adjective: with

two lamellae) alludes to the lamellate pair of apical

processes of the ventral process of the aedeagus.

Description. Body length 5.7—7.3 mm; length of forebody

3.0-3.3 mm. Coloration: body dark—brown to blackish-

brown; legs yellowish; antennae yellowish, with anten-

nomere | usually slightly darker.

Head (Fig. 2) weakly oblong; punctation coarse, dense,

and umbilicate; interstices without microsculpture; eyes

of moderate size and distinctly convex, slightly more than

one third as long as the distance from posterior margin of

eye to posterior constriction of head. Antenna approxi-

mately 1.8 mm long.

Pronotum (Fig. 2) approximately 1.15 times as long as

broad and 0.85—0.90 times as broad as head; punctation

even coarser than that of head, non-umbilicate, and mod-

erately dense; interstices narrower than diameter of punc-

tures, glossy; midline punctate in anterior half, impunc-

tate and somewhat elevated in posterior half; posterior half

with a pair of lateral and somewhat irregularly oblique ob-

long elevations.

Elytra (Fig. 2) approximately 0.55 times as long as

pronotum and slender; humeral angles obsolete; puncta-

tion dense, moderately defined, and coarse; interstices

glossy. Hind wings completely reduced.

Abdomen approximately 1.30—1.35 times as broad as

elytra; punctation dense, defined, and moderately coarse

on tergite III, gradually becoming sparser towards poste-

rior tergites, tergite VII with sparse punctation; interstices

with shallow microsculpture; posterior margin of tergite

VII without palisade fringe; posterior margin of tergite

VIII distinctly convex.

¢: sternite VI-VII unmodified (Fig. 3); sternite VIII

(Fig. 4) with unmodified pubescence, posterior excision

small and V-shaped; aedeagus (Figs 5—6) rather large in

relation to body size, 1.4—1.5 mm long; ventral process

apically with pair of lamellate appendices; dorso-lateral

apophyses curved, apically somewhat dilated and club-

shaped, extending beyond apex of ventral process.

Comparative notes. Based on the morphology of the

aedeagus (ventral process apically with pair of appendices;

dorso-lateral apophyses club-shaped), N. bilamellatus is

Bonn zoological Bulletin 62 (2): 125-170

closely allied to N. truncatus, which too is distributed in

the Emei Shan. It differs from this species by the much

larger and differently shaped aedeagus (N. truncatus:

aedeagus 0.9—1.0 mm long and with apical appendices of

ventral process of completely different shape; apices of

dorso-lateral apophyses far from reaching apex of ventral

process), by slightly larger body size, and by the denser

punctation of the posterior portion of the head and of the

elytra. For illustrations of the aedeagus of N. truncatus see

Zheng (1992).

Distribution and natural history. Nazeris bilamellatus

is presumably endemic to the Emei Shan (Fig. 28), where

the specimens were sifted from leaf litter at an altitude of

1800-2400 m, apparently together with N. truncatus and

N. ruani.

Nazeris hailuogouensis Hu et al., 2007 (Figs 7—9, 28)

Material examined. China: Sichuan: 5<, 39, Gongga

Shan, Hailuogou Glacier Park, Camp 1, 29°36°N,

102°04’E, 2100 m, 27.-31.V.1997, leg. Piitz, Schilke &

Wrase (cPiit, cSch, cAss); 14, 22, Gongga Shan, Hailu-

ogou Glacier Park, Camp 2, 29°35’N, 102°02’E,

2550-2700 m, 30.—31.V.1997, leg. Ptitz & Schiilke (cPit,

cSch); 19, Gongga Shan, Hailuogou Glacier Park, above

Camp 3, 3000 m, 30.V.1997, leg. Putz (cPut).

Comment. The original description of N. hailuogouensis

is based on nine type specimens from Hailuogou in the

Gongga Shan (Hu et al. 2007), where it is probably en-

demic (Fig. 28). The species is characterized particularly

by the shape of the ventral process and the long and slen-

der dorso-lateral apophyses of the aedeagus. The male sex-

ual characters are illustrated in Figs 7-9.

Nazeris foveatus sp. n. (Figs 10-14, 28)

Type material. Holotype 4: “CHINA: W-Sichuan 1999,

Ganzi Tibet. Aut. Pref., Luding Co., W Erlangshan-Pass,

2600 m, 7 km SSE Luding, 29°51N, 102°15E, Laub+ Na-

delstreu, Pilze, 22.VI., leg. M. Schiilke / Holotypus 4 Na-

zeris foveatus sp. n. det. V. Assing 2013” (cAss).

Paratypes: 44, 29: “CHINA: W-Sichuan 1999, Ganzi Ti-

bet. Aut. Pref., Luding Co., W Erlangshan-Pass, 2600 m,

7 km SSE Luding, 29°51N, 102°15E, Laubstreu, Pilze,

29.VI., leg. M. Schiilke” (cSch, cAss); 1d, 19: “CHINA

W-Sichuan (Ganzi Tibet. Aut. Pref., Luding Co.) W Er-

lang Shan Pass, 2600 m, 7 km SSE Luding, 29°S1N,

102°15E, Laubstreu, Pilze, 20.—29.V1.1999, D.W. Wrase”

(cSch, cAss).

©ZFMK

On the Nazeris fauna of China II 131

Etymology. The specific epithet (Latin, adjective: with 1m-

pression) alludes to the distinct impression of the male

sternite VIII, one of the characters distinguishing this

species from its closest relatives.

Description. Species of rather variable size; body length

5.5—7.5 mm; length of forebody 2.9—3.5 mm. Coloration:

body blackish-brown to black; legs yellowish; antennae

yellowish, with antennomere I slightly darker.

Head (Fig. 10) 1.08—1.15 times as long as broad; punc-

tation coarse, dense, and weakly umbilicate; interstices

without microsculpture; eyes small, distinctly less than one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna 1.7—2.0 mm

long.

Pronotum (Fig. 10) 1.20—1.25 times as long as broad and

approximately 0.9 times as broad as head; punctation dis-

tinctly coarser than that of head, non-umbilicate, and very

dense; interstices forming narrow ridges, glossy; posteri-

or half of midline narrowly elevated and impunctate.

Elytra (Fig. 10) approximately 0.6 times as long as

pronotum and slender; humeral angles obsolete; puncta-

tion dense, defined, and approximately as coarse as that

of pronotum; interstices glossy. Hind wings completely re-

duced.

Abdomen 1|.20—1.25 times as broad as elytra; puncta-

tion dense, defined, and rather coarse on tergites II-VI,

only indistinctly sparser and finer on tergite VII; interstices

without microsculpture; posterior margin of tergite VII

without palisade fringe; posterior margin of tergite VIII

distinctly convex.

S: sternite VII with weakly concave posterior margin

(Fig. 11); sternite VIII (Fig. 12) 1.10—1.15 times as broad

as long, with distinct oblong and in the middle weakly

sclerotized defined impression 1n postero-median portion;

posterior excision rather deep and narrow, its depth ap-

proximately 0.3 times the length of sternite; aedeagus

(Figs 13—14) slender, approximately 0.9 mm long (1.2 mm

including dorso-lateral apophyses); ventral process weak-

ly sclerotized, ventrally sharply edged, and with lateral

lamellae forming a “V” in ventral view; basal portion of

ventral process broad, without distinct lateral processes;

dorso-lateral apophyses long and slender, distinctly ex-

tending beyond apex of ventral process.

Comparative notes. As can be inferred from the highly

similar external and male sexual characters, particularly

the similarly derived morphology of the aedeagus, this

species is undoubtedly closely allied to N. hailuogouen-

sis and the two following species. It differs from them par-

ticularly by the shape of the male sternite VII (deeper and

larger postero-median impression; this impression with-

out pubescence and more extensively yellowish in the

middle) and by the morphology of the aedeagus, in par-

Bonn zoological Bulletin 62 (2): 125—170

ticular the shape of the ventral process (apical lamellae

short, ventral portion broad and without lateral process-

es in ventral view). For illustrations of N. hailuogouensis

see Figs 7—9 and figures 10—14 in Hu et al. (2007).

Distribution and natural history. The type locality is sit-

uated in the Erlang Shan, 7 km to the south-southeast of

Luding, Sichuan (Fig. 28). The type specimens were sift-

ed from the litter of broad-leaved and coniferous trees at

an altitude of 2600 m, together with the holotype of N.

cornutus.

Nazeris appendiculatus sp. n. (Figs 15—21, 28)

Type material. Holotype @: “CHINA: W-Sichuan (7),

Daxue Shan, W Kangding, 30.03.13N, 101.57.11E,

2700-2800 m, 24.05.1997, M. Schiilke / Holotypus 2

Nazeris appendiculatus sp. n. det. V. Assing 2013” (cAss).

Paratypes: 1, 59: “CHINA: W-Sichuan (4), Daxue Shan,

2500-2800 m, Bachtal 5 km E Kangding, 30.03.28N,

102.00.15E, 20.05.1997, M. Schiilke” (cSch, cAss); 29:

“CHINA: W-Sichuan (4a), Daxue Shan, Bachtal 5 km E

Kangding, 30.03.28N, 102.00.15E, 2500-2800 m,

23.05.1997, M. Schiilke” (cSch); 14: “CHINA: (W

Sichuan) (4), Daxue Shan, river valley 5 km E Kanding

[sic], 30.03N, 102.00E, 20.&23.05.1997, Wrase” (cAss);

2°:“CHINA: Sichuan, Ganzi pref., Daxue Shan, 102.00E,

30.03N, 5 km E Kangding, river valley, ca. 3000 m,

20.&23.V.1997, leg. A. Piitz” (cPiit); 1¢, 29: “CHINA:

Sichuan, Ganzi pref., Daxue Shan, 101.57N, 30.03E, N

Kangding, 2600-2700 m, 22.&24.V.1997, leg. A. Putz”

(cPut, cAss).

Etymology. The specific epithet (Latin, adjective: with ap-

pendices) alludes to the distinctive baso-lateral projections

(ventral view) of the ventral process of the aedeagus.

Description. Head with punctation not distinctly umbil-

icate. Other external characters (Fig. 15) as in N. hailuo-

gouensis and N. foveatus.

¢: sternite VII (Fig. 16) with posterior margin moder-

ately concave, particularly in the middle; sternite VIII (Fig.

17) approximately 1.05 times as broad as long, without

distinct postero-median impression, posterior excision nar-

rowly V-shaped, its depth approximately 0.23 times the

length of sternite; aedeagus (Figs 18-21) slender, 0.87 mm

long (approximately 1.1 mm including dorso-lateral

apophyses); ventral process weakly sclerotized, ventral-

ly sharply edged, and with lateral lamellae forming a “V”

in ventral view; basal portion of ventral process slender,

with a pair of distinct lateral projections; dorso-lateral

apophyses long and slender, distinctly extending beyond

apex of ventral process.

©ZFMK

132 Volker Assing

Figs 15-27. Nazeris appendiculatus (15-21), and N. puetzi (22-27). 15, 23: forebody; 16, 24: male sternite VII; 17: male stern-

ite VII; 18-21, 25-26: aedeagus in lateral and in ventral view; 22: habitus; 27: apical portion of median lobe of aedeagus in ven-

tral view. Scale bars: 15, 22—23: 1.0 mm; 16-21, 24-26: 0.5 mm; 27: 0.2 mm.

Bonn zoological Bulletin 62 (2): 125—170 ©OZFMK

On the Nazeris fauna of China II 133

Comparative notes. Nazeris appendiculatus undoubted-

ly belongs to the group of species allied to N. hailuogouen-

sis, aS is suggested particularly by the similarly derived

morphology of the aedeagus. It is distinguished from the

two preceding species of this group by the non-umbilicate

punctation of the head, the less deep and relatively broad-

er posterior excision of the male sternite VIII, the absence

of a distinct postero-median impression of the male ster-

nite VIII, and the morphology of the aedeagus (median

lobe of aedeagus relatively longer and of different shape

in lateral view, basally with pair of conspicuous projec-

tions in ventral view).

Distribution and natural history. The species was dis-

covered in three localities in the Daxue Shan in the envi-

rons of Kangding, Sichuan (Fig. 28), at altitudes between

2600 and approximately 3000 m. Additional data are not

available

Nazeris puetzi sp. n. (Figs 22—28, 41)

Type material. Holotype @: “CHINA: W-Sichuan, Ya’an

Prefecture, Tianquan Co., W Erlang Shan Pass / 2900 m,

21.V1.1999, 29.51.13N, 102.17.28E, leg. A. Pitz, sifted

/ Holotypus 3 Nazeris puetzi sp. n. det. V. Assing 2013”

(cAss).

Etymology. The species is dedicated to Anreas Piitz, spe-

cialist of Byrrhidae, who collected the holotype.

Description. Habitus as in Fig. 22. Head with punctation

not distinctly umbilicate (Fig. 23). Other external charac-

ters as in N. hailuogouensis and N. foveatus.

3: sternite VII (Fig. 24) strongly transverse, posterior

margin distinctly concave; sternite VIII (Fig. 41) 1.17

times as broad as long, without distinct postero-median

impression, posterior excision narrowly V-shaped, its

depth approximately 0.3 times the length of sternite;

aedeagus (Figs 25—27) slender, 0.87 mm long (1.08 mm

including dorso-lateral apophyses); ventral process weak-

ly sclerotized, ventrally sharply edged, and with lateral

lamellae forming a “V” in ventral view; basal portion of

ventral process moderately broad, without lateral projec-

tions; dorso-lateral apophyses long and slender, distinct-

ly extending beyond apex of ventral process.

Comparative notes. Nazeris puetzi, too, clearly belongs

to the group of species allied to N. hailuogouensis. It is

distinguished from N. foveatus, another species of this

group probably endemic to the Erlang Shan, by the non-

umbilicate punctation of the head, the distinctly concave

posterior margin of the more transverse male sternite VI,

the shape of the male stenite VII (without distinct pos-

tero-median impression, posterior excision broad), and the

Bonn zoological Bulletin 62 (2): 125-170

morphology of the aedeagus (basal portion of ventral

process more slender in ventral view; apical portion of

ventral process longer; lateral aspect of ventral process of

different shape).

Distribution and natural history. The type locality is sit-

uated in the Erlang Shan in W-Sichuan (Fig. 28). The holo-

type was sifted from leaf litter at an altitude of 2900 m.

Nazeris bicornis Hu et al., 2007 (Fig. 29)

Material examined. China: Sichuan: 1, 2°, Gongga

Shan, Hailuogou Glacier Park, Camp 1, 29°36’N,

102°04’E, 2100 m, 27.-31.V.1997, leg. Schtilke & Wrase

(cSch, cAss); 12, Gongga Shan, Hailuogou Glacier Park,

Camp 2, 29°35’N, 102°02’E, 2550-2700 m,

30.-31.V.1997, leg. Schiilke (cSch).

Comment. Nazeris bicornis is known only from the

Gongga Shan (Fig. 29); its original description is based

on eight type specimens from Hailuogou. Based on the

morphology of the aedeagus (ventral process apically with

a pair of processes), it is closely allied to N. truncatus and

N. bilamellatus.

Nazeris cornutus sp. n. (Figs 29, 30-34)

Type material. Holotype ¢: “CHINA: W-Sichuan 1999,

Ganzi Tibet. Aut. Pref., Luding Co., W Erlangshan-Pass,

2600 m, 7 km SSE Luding, 29°51N, 102°15E, Laubstreu,

Pilze, 29.VI., leg. M. Schiilke / Holotypus ¢ Nazeris cor-

nutus sp. n. det. V. Assing 2013” (cAss).

Etymology. The specific epithet (Latin, adjective: with

horns) alludes to the horn-shaped apices of the ventral

process of the aedeagus and the evident close relationship

with N. bicornis.

Description. Body length 6.0 mm; length of forebody 2.9

mm. Habitus as in Fig. 30. Coloration: body blackish-

brown; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker.

Head (Fig. 31) 1.08 times as long as broad; punctation

coarse, very dense, and umbilicate; interstices without mi-

crosculpture; eyes moderately small, approximately one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna 1.6 mm long.

Pronotum (Fig. 31) 1.2 times as long as broad and 0.9

times as broad as head; punctation somewhat coarser than

that of head, non-umbilicate, and moderately dense; in-

terstices distinctly narrower than diameter of punctures,

glossy; posterior half of midline narrowly elevated and im-

punctate.

OZFMK

134 Volker Assing

(fp

gw Chengdu 4

Fig. 28. Distributions of Nazeris species in Sichuan: N. truncatus (filled star); N. bilamellatus (filled star); N. ruani (filled star:

examined record; open star: type locality); N. hailuogouensis (filled triangles); N. foveatus (filled circle); N. appendiculatus (dia-

monds); N. puetzi (open triangle).

Elytra (Fig. 31) 0.58 times as long as pronotum and slen-

der; humeral angles obsolete; punctation dense, defined,

and approximately as coarse as that of pronotum; inter-

stices glossy. Hind wings completely reduced.

Abdomen 1.2 times as broad as elytra; punctation dense,

defined, and rather coarse on anterior tergites, gradually

becoming sparser and finer towards posterior tergites,

rather sparse and fine on tergite VH; interstices without

microsculpture; posterior margin of tergite VII without

palisade fringe; posterior margin of tergite VIII convex.

@: sternite VII unmodified; sternite VII (Fig. 32) 1.05

times as broad as long, posterior excision V-shaped and

rather small, its depth approximately 0.18 times the length

of sternite; aedeagus (Figs 33-34) slender, 1.05 mm long;

ventral process apically with pair of curved and apically

acute horn-shaped processes in ventral view; dorso-later-

al apophyses long and slender, apically just reaching

apices of ventral process.

Comparative notes. As can be inferred from the similar

external and male sexual characters, particularly the sim-

ilarly derived morphology of the aedeagus, this species is

undoubtedly closely allied to N. bicornis, from which it

differs by the slightly denser punctation of the abdomen,

Bonn zoological Bulletin 62 (2): 125-170

the smaller posterior excision of the male sternite VIII, and

by the distinctly smaller and differently shaped aedeagus

(ventral process basally more slender; apical processes of

ventral process shorter and apically less acute; dorso-lat-

eral apophyses more slender; internal structures of differ-

ent shape). The aedeagus of N. bicornis is 1.2 mm long;

for illustrations of the male sternite VIII and the aedea-

gus of N. bicornis see figures 6—9 in Hu et al. (2007).

Distribution and natural history. The type locality is sit-

uated in the Erlang Shan, 7 km to the south-southeast of

Luding, Sichuan (Fig. 29). The holotype was sifted from

leaf litter at an altitude of 2600 m, together with N. fovea-

tus.

Naczeris trifurcatus sp. n. (Figs 29, 35-40)

Type material. Holotype @: “CHINA: S-Sichuan 1999,

Ya’an Prefecture, Shimian Co., Xiaoxiang Ling, 7 km S

Liziping, 35 km S Shimian, 28°56N, 102°18E, ca. 1600

m, Bambus, Feldrand, 7.VU., leg. M. Schiilke / Holoty-

pus ¢ Nazeris trifurcatus sp. n. det. V. Assing 2013”

(cAss).

©ZFMK

On the Nazeris fauna of China II 135

gw Chengdu

Fig. 29. Distributions of Nazeris species in Sichuan: N. bicornis (triangles); N. cornutus (filled circle); N. infurcatus (star); N.

wrasei (diamond); N. schuelkei (open circles).

Etymology. The specific epithet is an adjective composed

of the Latin prefix tri- (three) and an adjective derived

from the Latin noun furca (fork). It alludes to the trifur-

cate apex of the ventral process of the aedeagus.

Description. Body length 6.0 mm; length of forebody 3.1

mm. Habitus as in Fig. 35. Coloration: body blackish-

brown; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker.

Head (Fig. 36) 1.08 times as long as broad; punctation

coarse, dense, and umbilicate; interstices without mi-

crosculpture; eyes small, less than one third as long as the

distance from posterior margin of eye to posterior constric-

tion of head. Antenna 1.8 mm long.

Pronotum (Fig. 36) 1.20 times as long as broad and 0.93

times as broad as head; punctation even coarser than that

of head, non-umbilicate, and moderately dense; interstices

distinctly narrower than diameter of punctures, glossy;

midline punctate in anterior half, impunctate and some-

what elevated in posterior half; posterior half with a pair

of somewhat irregular glossy oblong elevations.

Elytra (Fig. 36) 0.6 times as long as pronotum and slen-

der; humeral angles obsolete; punctation dense, moderate-

Bonn zoological Bulletin 62 (2): 125-170

ly defined, and nearly as coarse as that of pronotum; in-

terstices glossy. Hind wings completely reduced.

Abdomen 1.25 times as broad as elytra; punctation

dense, defined, and moderately coarse on tergites IH-IV,

sparser on tergite V, gradually becoming sparser towards

posterior tergites, tergite VII with sparse punctation; in-

terstices without microsculpture; posterior margin of ter-

gite VII without palisade fringe; posterior margin of ter-

gite VIII distinctly convex.

&: sternite VI unmodified (Fig. 37); sternite VIII (Fig.

38) weakly oblong with unmodified pubescence, posteri-

or excision V-shaped, 0.17 times as deep as length of ster-

nite; aedeagus (Figs 39-40) slender, 1.1 mm long; ven-

tral process apically trifurcate, with the median process

distinctly shorter than the lateral ones; dorso-lateral

apophyses curved, weakly dilated in apical half, and not

reaching apex of ventral process.

Comparative notes. Nazeris trifurcatus is distinguished

from all its congeners by the male sexual characters, par-

ticularly the trifurcate apex of the ventral process of the

aedeagus. The latter suggests that the species may be

closely related to N. bicornis and allied species.

OZ7FMK

136 Volker Assing

33 34 39 40 41

Figs 30-41. Nazeris cornutus (30-34), N. trifurcatus (35-40), and N. puetzi (41). 30, 35: habitus; 31, 36: forebody; 32, 38, 41:

male sternite VIH; 33-34, 39-40: aedeagus in lateral and in ventral view; 37: male sternite VI. Scale bars: 30-31, 35-36: 1.0 mm;

32-34, 37-41: 0.5 mm.

Distribution and natural history. The type locality is sit- | Nazeris wrasei sp. n. (Figs 29, 42-47)

uated to the south of Shimian in southern Sichuan (Fig.

29). The holotype was collected in a field margin with Type material. Holotype 4: “CHINA S.Sichuan (Ya’an

bamboo at an altitude of approximately 1600 m. Pref., Shimian Co.), Xiaoxiang Ling, side-valley above

Nanya Cun nr. Caluo, 11 km S Shimian, ca. 1250 m,

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

On the Nazeris fauna of China II 137

7.VII.1999, D.W. Wrase / Holotypus ¢ Nazeris wrasei sp.

n. det. V. Assing 2013” (cAss). Paratypes: 19: same da-

ta as holotype (cSch); 29: “CHINA: W-Sichuan, Ya’an

Pref., Shimian Co., Xiaoxiang Ling, side-valley ab. Nanya

Cun nr. Caluo, 11 km S Shimian, 1250 m, 7.VI.1999, leg.

A. Piitz” (cPiit, cAss).

Etymology. The species is dedicated to my friend David

Wrase (Berlin), specialist of Carabidae, who collected two

of the type specimens.

Description. Body length 4.7—5S.2 mm; length of forebody

2.5—2.7 mm. Habitus as in Fig. 42. Coloration: body black-

ish-brown; legs yellowish; antennae yellowish, with an-

tennomere I slightly darker.

Head (Fig. 43) approximately 1.05 times as long as

broad; punctation moderately coarse, very dense, and um-

bilicate; interstices without microsculpture; eyes moder-

ately small, approximately one third as long as the distance

from posterior margin of eye to posterior constriction of

head. Antenna approximately 1.5 mm long.

Pronotum (Fig. 43) approximately 1.15 times as long

as broad and 0.95 times as broad as head; punctation dis-

tinctly coarser than that of head, non-umbilicate, and mod-

erately dense; interstices distinctly narrower than diame-

ter of punctures, glossy; midline punctate in anterior half,

impunctate and narrowly elevated in posterior half.

Elytra (Fig. 43) approximately 0.55 times as long as

pronotum; humeral angles obsolete; punctation dense,

moderately defined, and distinctly less coarse than that of

pronotum; interstices glossy. Hind wings completely re-

duced.

Abdomen 1.20—1.25 times as broad as elytra; puncta-

tion dense, defined, and moderately coarse on anterior ter-

gites; sparser and finer on tergite VII; interstices without

microsculpture; posterior margin of tergite VII without

palisade fringe; posterior margin of tergite VII weakly

convex.

3: sternite VII (Fig. 44) with truncate posterior margin

and with a transverse series of long submarginal setae, oth-

erwise unmodified; sternite VIII (Fig. 45) approximately

as long as broad, posterior excision anteriorly rounded (not

acute) and approximately 0.2 times as deep as length of

sternite; aedeagus (Figs 46-47) stout, 0.85 mm long; ven-

tral process apically concave in ventral view; dorso-lat-

eral apophyses stout and strongly sclerotized, slightly ex-

tending beyond apex of ventral process.

Comparative notes. The morphology of the aedeagus is

somewhat similar to that of species of the NV. cultellatus

group (Assing 201 3a), but the ventral process is less dis-

tinctly compressed laterally and, unlike the species of the

N. cultellatus group, the punctation of the head is umbil-

icate. From the species previously recorded from Sichuan,

N. wrasei is readily distinguished by the stouter aedeagus.

Bonn zoological Bulletin 62 (2): 125-170

Distribution and natural history. The type locality is sit-

uated to the south of Shimian in southern Sichuan (Fig.

29). The type specimens were collected at an altitude of

approximately 1250 m.

Nazeris schuelkei sp. n. (Figs 29, 48-53)

Type material. Holotype @: “CHINA: W-Sichuan 1999,

Ya’an Prefecture, Tianquan Co., Jiajin Shan, Tal oberh.

Labahe N.R. St., 57 km W Ya’an, 30°06N, 102°25E, Streu,

Rinde, Pilze, 1800 m, 12.VII., leg. M. Schiilke / Holoty-

pus ¢ Nazeris schuelkei sp. n. det. V. Assing 2013” (cAss).

Paratypes: 2: same data as holotype (cSch); 14: “CHI-

NA: W-Sichuan 1999, Ya’an Prefecture, Fulin Co., Dax-

iang Ling, Rd. zw. Hanyuanjie u. Siping, 51 km NNE

Shimian, 2300 m, 29°39N, 102°37E, Ufer, Gesiebe,

10.VIL, leg. M. Schiilke” (cAss); 19: “CHINA: W-

Sichuan, Ya’an Pref., Fulin Co., Daxiang Ling, pass N /

Hanyuanjie, 51 km NNE Shimian, smal [sic] stream,

29.39N, 102.37E, 2300 m, 10.VII.1999, leg. A. Piitz”

(cPut).

Etymology. The species is dedicated to my friend and col-

league Michael Schiilke, who collected most of the type

specimens and whose material the present paper 1s large-

ly based on.

Description. Body length 5.0—S.7 mm; length of forebody

2.6—2.9 mm. Habitus as in Fig. 48. Coloration: body dark-

brown to blackish-brown; legs yellowish; antennae yel-

lowish, with antennomere I slightly darker. Abdomen with

microsculpture. Other external characters (Fig. 49) as in

N. wrasei.

&: sternite VII (Fig. 50) with truncate posterior margin

and with a transverse series of moderately long submar-

ginal setae, otherwise unmodified; sternite VIII (Fig. 51)

approximately 1.1 times as broad as long, posterior exci-

sion anteriorly narrowly rounded (not acute) and approx-

imately 0.2 times as deep as length of sternite; aedeagus

(Figs 52-53) stout, 0.85 mm long; ventral process apical-

ly acute in ventral view; dorso-lateral apophyses stout and

strongly sclerotized, subbasally somewhat dilated, distinct-

ly converging in ventral view, and extending nearly to apex

of ventral process.

Comparative notes. The similar external and male sex-

ual characters suggest that NV. schuelkei is closely related

to the geographically close N. wrasei, from which it is dis-

tinguished particularly by the presence of microsculpture

on the abdomen, by the different shape of the male ster-

nite VIII, and by the morphology of the aedeagus (shapes

of ventral process and of dorso-lateral apophyses).

OZFMK

138 Volker Assing

Figs 42-53. Nazeris wrasei (42-47) and N. schuelkei (48-53). 42, 48: habitus; 43, 49: forebody; 44, 50: male sternite VIL; 45,

51: male sternite VIII; 46-47, 52-53: aedeagus in lateral and in ventral view. Scale bars: 42-43, 48-49: 1.0 mm; 44-47, 50-53:

0.5 mm.

Bonn zoological Bulletin 62 (2): 125-170 OZFMK

On the Nazeris fauna of China II 139

Distribution and natural history. The species was dis-

covered in two localities in Ya’an Prefecture, West Sichuan

(Fig. 29). The specimens were sifted from litter, in one lo-

cality near a stream bank, at altitudes of 1800 and 2300

Unnamed and presumably undescribed species

Three probably undescribed species remain unnamed since

they are represented in the examined material exclusive-

ly by females.

Nazeris sp. 1: 19: “CHINA — W-Sichuan, Ya’an Pref.,

Shimian Co., Xiaoxiang Ling, Pass betw. Shimian-Gan-

luo, 27 km SE Shimian, 2450 m, springfed-swamp,

29.02.75N, 102.31.48E, 8.VII.1999, leg. A. Piitz” (cPiit).

This large species is distinguished from the similarly

large N. ruani by more slender body, a more elongate head,

distinctly coarser and non-umbilicate punctation of the

head, and the coarser punctation of the abdomen. The on-

ly other species of similar size known from Sichuan is N.

magnus Hu et al., 2007, which was described from the Er-

lang Shan. Judging from the habitus photo provided by

Hu et al. (2007), N. magnus appears to have a more dense-

ly punctate head and pronotum.

Nazeris sp. 2: 19: “CHINA W.Sichuan (Ganzi Tibet. Aut.

Pref., Yajiang Co.), Shalui Shan, river valley 6 km WSW

Yajiang, 3250 m, 30°0IN, 102°57E (river bank, bank

slope), 4. VII.1999 D.W. Wrase” (cSch).

This moderately large, black species is distinguished

from most geographically close congeners by the non-um-

bilicate punctation of the head. Based on the external char-

acters, it may belong to the N. hailuogouensis group.

Nazeris sp. 3: 19: “CHINA NW Sichuan, 20 km NW

Maowen, 2150 m, Jiuding Shan, coniferous wood,

7-28.V1.2004, leg. R. Fabbri” (cSch).

Species from Yunnan

Species groups. The N. giganteus group is represented 1n

Yunnan by WN. giganteus (Diancang Shan) and possibly the

distinctly smaller NV. a/pinus (Yulongxue Shan). This group

also includes N. ruani from Sichuan, most likely the

adelphotaxon of NV. giganteus. For more details see the sec-

tion on the species groups of Sichuan.

The N. daliensis group comprises at least five species,

N. daliensis (Diancang Shan), N. jizushanensis (Jizu

Shan), N. wuliangicus (Wuliang Shan and adjacent

mountain range), NV. sagittifer (Wuliang Shan, Ailao Shan),

and N. nivimontis (Xue Shan N Lincang). This species

group 1s constituted particularly by the derived morphol-

Bonn zoological Bulletin 62 (2): 125-170

ogy of the aedeagus (ventral process short, weakly scle-

rotized, more or less sharply edged ventrally; dorso-lat-

eral apophyses long and slender, extending far beyond

apex of the ventral process) (e.g., Figs 58-59, 62-63,

65—66). The male posterior incision of the male sternite

VUI is relatively small, 0.25 times as deep as the sternite

at most (e.g., Figs 57, 69, 73, 75), and the male sternite

VI is weakly modified at most (posterior margin truncate

to moderately concave in the middle) (e.g., Figs 68, 72,

74).

The N. vexillatus group includes three species, all of

them distributed in the Gaoligong Shan: N. vexillatus, N.

circumclusus, and N. hastatus. They are characterized by

an aedeagus with a short ventral process and with modi-

fied (dilated or with processes) dorso-lateral apophyses ex-

tending far beyond the apex of the ventral process of the

median lobe (Figs 180-181, 187-188, 193-194). The male

sternite VII is weakly modified (e.g., Figs 185, 191) and

the body colour is brown to dark-brown, but not black.

The N. fissus group is composed of two species, N. fis-

sus (Wuliang Shan, Ailao Shan) and N. caoi Hu et al., 2011

(Nabanhe Nature Reserve). This group is constituted by

a conspicuous synapomorphy, a completely divided (bi-

fid) ventral process of the aedeagus (Figs 205—206).

The male sexual characters of N. secatus (Laobie Shan)

and N. bangmaicus (Bangma Shan) do not suggest clos-

er affiliations to any of the other species known from Yun-

nan.

The remaining species are - partly tentatively - assigned

to the N. cangicus group. They are characterized by an

aedeagus with a more or less slender and apically acute

ventral process (e.g., Figs 96-97, 102-103), usually un-

modified dorso-lateral apophyses not extending distinct-

ly beyond the apex of the ventral process of the median

lobe (exceptions: N. pungens, N. lanuginosus), a more or

less distinctly modified male sternite VII (often with a pos-

tero-median impression, a medially concave posterior mar-

gin, and/or modified pubescence) (e.g., Figs 94, 100), and

a male sternite VIII with a usually deep and narrow pos-

terior excision (often also with an oblong median impres-

sion) (e.g., Figs 95, 101). The colour of the body of ma-

ture specimens is usually blackish. Based on the illustra-

tions provided in their respective original descriptions, WN.

zhangi Watanabe & Xiao, 1993 (Yu’an Shan near Kun-

ming), NV. baihuaensis Watanabe & Xiao, 2000 (Gaoligong

Shan), N. nomurai Watanabe & Xiao, 2000 (Gaoligong

Shan), N. huanxipoensis Watanabe & Xiao, 2000

(Gaoligong Shan), N. ishiianus Watanabe & Xiao, 2000

(Gaoligong Shan), and N. nabanhensis Hu et al., 2011 (Na-

banhe Nature Reserve) may belong to the N. cangicus

group, too. Within this group, two species pairs were iden-

tified. One of them is represented by N. pungens (Xue

Shan N Lincang) and N. /anuginosus (Laobie Shan), both

of which have a male sternite VII with a posterior clus-

ter of long black setae (Figs 113, 119) and an aedeagus

OZFMK

Volker Assing

140

Figs 54-66. Nazeris wuliangicus (54-60), N. daliensis (61-64), and N. jizushanensis (65-66). 54: habitus; 55: forebody; 56: male

sternite VII; 57: male sternite VII; 58-59, 62-63, 65-66: aedeagus in lateral and in ventral view; 60, 64: ventral process of aedea-

gus in ventral view; 61: median dorsal portion of head. Scale bars: 54-55: 1.0 mm; 56-59, 62-63, 65—66: 0.5 mm, 60-61, 64: 0.2

mm.

OZFMK

Bonn zoological Bulletin 62 (2): 125-170

On the Nazeris fauna of China II 141

with the dorso-lateral apophyses extending far beyong the

apex of the ventral process (Figs 115-116, 121-122). The

other species pair comprises N. peniculatus (mountain

range N Er Hai) and N. barbatus (Wuliang Shan), whose

male sternites VI share a unique synapomorphy, a clus-

ter of modified stout black setae (Figs 118, 133).

Nazeris giganteus Watanabe & Xiao, 1997 (Fig. 90)

Material examined. China: Yunnan: 1°, Dali Bai Aut.

Pref., Diancang Shan, 3 km W Dali old Town, “cloud

road”, 25°41’N, 100°07°E, 2750 m, pine forest with

broadleaved undergrowth and bamboo, vinegar trap,

17.-23.VI.2005, leg. Wrase (cAss).

Comment. This large species was described based on

three specimens from “Qinghi Xi, Diancang Shan Mts.”

and one male from “Mt. Laohu Shan, Dali Shi” (Watan-

abe & Xiao 1997). The above specimen is a female, but

no other species of similar size are known from the Di-

ancang Shan. Aside from the conspicuously large body

size, N. giganteus shares similarly derived male charac-

ters with N. ruani from Sichuan (see the section on the

species groups occurring in Sichuan). The above record

is mapped in Fig. 90.

Nazeris alpinus Watanabe & Xiao, 1997 (Fig. 88)

Material examined. China: Yunnan: 2¢, 7%, Lijiang

Naxi Aut. Co., E Yulongxue Shan, 30 km N Lijiang,

27°09°N, 100°15’E, 2800-2900 m, secondary mixed for-

est, 13. VIII.2003, leg. Smetana & Wrase (cSch, cSme,

cAss); 12, 26 km N Lijiang, Ganhaizi pass, 27°07’N,

100°15’E, 3000 m, mixed coniferous forest, litter and

moss under rhododendron sifted, 15.V1.2007, leg. Hajek

& Ruzicka (cAss).

Comment. The original description of N. alpinus is based

on nine type specimens from the Yulongxue Shan

(Watanabe & Xiao 1997), where this species is probably

endemic. The phylogenetic affiliations are unclear. Based

on the short, strongly curved, and strongly sclerotized dor-

so-lateral apophyses and the presence of a minute medi-

an incision of the posterior margin of the male sternite VII

(a character otherwise observed only in N. ruani), N. alpi-

nus is tentatively attributed to the N. giganteus group. The

above records are mapped in Fig. 88.

Bonn zoological Bulletin 62 (2): 125-170

Nazeris daliensis Watanabe & Xiao, 1997 (Figs 61—64,

72-73, 88)

Material examined. China: Yunnan: 4°, Dali Bai Aut.

Pref., Diancang Shan, 5 km SSW Dali, 25°39°N,

100°08’E, 2800 m, 26.VIII.2003, leg. Schilke, Smetana,

Wrase (cSch, cSme, cAss); 14, 29 [all teneral], Diancang

Shan, 3 km W Dali, 25°41’N, 100°07’E, 2600-2650 m,

30.VIII.2003, leg. Smetana (cSme); 1¢, 1% [1 teneral],

Diancang Shan, 3 km W Dali, 25°41°N, 100°07°E, 2750

m, 1.1X.2003, leg. Smetana (cSme, cAss); 1%, Diancang

Shan, 4 km W Dali, 25°41°N, 100°07°E, 2906-3000 m,

21.VIII.2003, leg. Smetana (cSme); 1, Diancang Shan,

3 km W Dali old Town, “cloud road”, 25°41’°N, 100°07’E,

2700 m, 17.VI.2005, leg. Smetana (cAss); 14, 59 [4 ten-

eral], Diancang Shan, 3 km W Dali old Town, “cloud

road”, 25°41°N, 100°07°E, 2650-2750 m, pine forest, pine

litter and moss in ditches sifted, 30. VIII.2003, leg. Schiilke

(cSch); 1 [teneral], same data, but 1.[X.2003, leg. Wrase

(cSch); 12, same data, but 17.VI.2005, leg. Schiilke

(cAss); 12, same data, but 17.-23.VI.2005, leg Wrase

(cSch); 3¢ [all slightly teneral], 1°, Diancang Shan W

Dali, 25°42°N, 100°07°E, 2860 m, pine forest, litter and

moss sifted, 28.V.2007, leg. Piitz & Schiilke (cPtt, cSch,

cAss); 1¢, Diancang Shan, near Dali, 25°40’N, 100°08’E,

2730 m, 11.V.2010, leg. Grebennikov (cSme); 19, Dian-

cang Shan, east slope of Zhonghe Shan, 25°41°N,

100°08’E, 2650 m, mixed forest with pine and rhododen-

dron, litter sifted, 13.VI.2007, leg. Hajek & Ruzicka

(cAss); 1¢, Diancang Shan, E pass 43 km NW Dali,

26°00°N, 100°00°E, 2700 m, secondary pine forest, litter

and moss sifted, 23. VIII.2009, leg. Wrase (cAss); 14, 19,

Diancang Shan above Dali, 2000-2200 m, 4—17.1V.1999,

leg. Schawaller (SMNS, cAss); 2 [1 teneral], same da-

ta, but 2500-2700 m, 8—18.1V.1999 (SMNS, cAss); 19,

same data, but 2700-2900 m, 14.IV.1999 (SMNS).

Comment. The original description of N. daliensis is

based on numerous type specimens from the type locali-

ty (“Mt. Xiaojin Shan, Diancang Shan Mts., Dali Shi’),

two other localities (“Zhonghe Feng”, “Qinghi X17’) in the

Diancang Shan, and from the “Laohu Shan, Dali Shi”.

Watanabe & Xiao (1997) illustrate the slightly different

aedeagi of a male from the type locality and from the Lao-

hu Shan. These illustrations are evidently not quite accu-

rate, particularly regarding the shape of the dorso-lateral

apophyses. Photographs of the head, the aedeagus, and of

the male sternites VII and VII are provided in Figs 61—64,

72—73. The above records are mapped in Fig. 88. Tener-

al specimens were collected in April and May, as well as

in August and September.

©ZFMK

142 Volker Assing

Figs 67-75. Nazeris nivimontis (67-71), N. daliensis (72-73), and N. jizushanensis (74-75). 67: forebody; 68, 72, 74: male ster-

nite VII; 69, 73, 75: male sternite VIII; 70-71: aedeagus in lateral and in ventral view. Scale bars: 67: 1.0 mm; 68-75: 0.5 mm.

Nazeris jizushanensis Watanabe & Xiao, 1997

(Figs 65—66, 74-75, 88)

Material examined. China: Yunnan: 5, 5 [partly te-

neral], Dali Bai Aut. Pref., 37 km NE Dali, Jizu Shan, trail

to cable car, 25°58’N, 100°23’E, 2450 m, mixed forest,

litter an moss sifted, 5.1X.2009, leg. Schtilke & Wrase

(cSch, cAss); 24%, 22, Jizu Shan, along trail to the sum-

mit, 25°58’°N, 100°23’E, 2180-2580 m, mixed forest with

pine, oak, and rhododendron, near — stream,

22.—24.V1.2007, leg. Hajek & Ruziéka (cSch, cAss).

Comment. The original description is based on seven type

specimens from the Jizu Shan (Watanabe & Xiao 1997),

where the above specimens were collected, too (Fig. 88).

The extremely similar external and male sexual charac-

ters suggest that NV. juzushanensis is closely related to, and

possibly the sister species of, N. daliensis, from which it

is best distinguished by the differently shaped dorso-lat-

eral apophyses of the larger aedeagus. The male sexual

characters of N. jizushanensis are illustrated in Figs 65—66,

74-75.

Bonn zoological Bulletin 62 (2): 125-170

7220 74

Nazeris wuliangicus sp. n. (Figs 54—60, 88)

Type material. Holotype @: “CHINA: Yunnan, Dali Bai

Aut. Pref., Wuliang Shan, 9 km SW _ Weishan,

25°10°15.5’’N, 100°14’21.8” E, 2480 m, scrub with (oak,

alder, pine), litter & mushrooms sifted, 14.1X.2009, leg.

M. Schiilke [CH09-51] / Holotypus 4 Nazeris wuliangi-

cus sp. n. det. V. Assing 2013” (cAss). Paratypes: 94, 79

[partly teneral]: same data as holotype (cSch, cAss); 63,

72: “CHINA: Yunnan [CH07-35], Dali Bai Auton. Pref.,

Wuliang Shan, 9 km SW Weishan, 2450-2500 m,

25°10°14"N, 100°14°22”E, oaks and pines, sifted,

13.VI.2007, M. Schiilke” (cSch, cAss); 2, 62: same da-

ta, but leg. A. Piitz (cPiit, cAss); 34, 29: “CHINA (Yun-

nan) Dali Bai Auton. Pref., Wuliang Shan, 9 km SW Wei-

shan, 2450-2500 m, 25°10’714”N, 100°14°22”E, (W.

slope, sec. oak/pine for., pasture, und. stones), 13.V1.2007

D.W. Wrase [35]” (cSch, cAss); 94, 209 [partly tener-

al]: “CHINA: Yunnan, Dali Bai Aut. Pref., mount. range

E Wei-shan, 12 km NE Weishan, 25°17’02-15’N,

100°22’23-30”E, 2630-2660 m, scrub with pines and

bamboo, litter sifted, 15.1X.2009, leg. M. Schiilke [CH09-

©ZFMK

On the Nazeris fauna of China II 143

Figs 76-87. Nazeris sagittifer (76-80) and N. secatus (81-87). 76, 83: male sternite VII; 77, 84: male sternite VIII; 78-79, 85-86:

aedeagus in lateral and in ventral view; 80, 87: ventral process of aedeagus in ventral view; 81: habitus; 82: forebody. Scale bars:

81-82: 1.0 mm; 76-79, 83-86: 0.5 mm; 80, 87: 0.1 mm.

54]” (ZFMK, cSch, cAss); 34 [1 teneral]: “CHINA (Yun-

nan) Dali Bai Aut. Pref., mount. range E Weishan, 12 km

NE Weishan, 2630-2660 m (scrub with pines and bam-

boo, litter sifted) 25°17’02-15”N, 100°22’23-30”E,

15.1X.2009, D.W. Wrase [54A]” (cSch).

Etymology. The specific epithet is an adjective derived

from the mountain range where the type locality is situ-

ated.

Bonn zoological Bulletin 62 (2): 125-170

Description. Body length 5.2—6.3 mm; length of forebody

2.8—3.1 mm. Habitus as in Fig. 54. Coloration: body black-

ish-brown to black, with the elytra occasionally slightly

paler; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker.

Head (Fig. 55) approximately 1.05 times as long as

broad; median dorsal portion indistinctly elevated at most;

punctation moderately coarse, dense, and umbilicate; in-

terstices without microsculpture, forming narrow ridges;

144 Volker Assing

| Kunming

Fig. 88. Distributions of Nazeris species in Yunnan: N. alpinus (filled triangles); N. daliensis (open circles); N. jizushanensis (filled

diamond); N. wuliangicus (filled circles); N. sagittifer (open diamonds); N. nivimontis (star); N. secatus (open triangle).

eyes moderately small, approximately one third as long

as the distance from posterior margin of eye to posterior

constriction of head. Antenna approximately 1.7 mm long.

Pronotum (Fig. 55) approximately 1.2 times as long as

broad and 0.9 times as broad as head; punctation dense

and non-umbilicate, much coarser than that of head; in-

terstices glossy, forming narrow ridges; midline punctate

in anterior half, impunctate and narrowly elevated in pos-

terior half.

Elytra (Fig. 55) approximately 0.55 times as long as

pronotum; humeral angles obsolete; punctation similar to

that of pronotum; interstices glossy. Hind wings complete-

ly reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense, defined, and coarse on tergites II-VI,

much finer and sparser on tergites VII and VII than on

tergite VI; interstices without microsculpture and glossy;

posterior margin of tergite VII without palisade fringe;

posterior margin of tergite VIII strongly convex.

¢: sternite VII (Fig. 56) with posterior margin very

weakly concave in the middle, otherwise unmodified; ster-

nite VIII (Fig. 57) weakly transverse, posterior excision

small and V-shaped; aedeagus (Fig 58-60) approximate-

ly 0.85 mm long; ventral process short, apically acute in

Bonn zoological Bulletin 62 (2): 125-170

ventral view, and laterally somewhat compressed; dorso-

lateral apophyses long and slender, distinctly extending be-

yond apex of ventral process.

Comparative notes. Based on the shapes and chaetotaxy

of the male sternite VII and VII, as well as on the mor-

phology of the aedeagus (short ventral process; long and

slender dorso-lateral apophyses), N. wuliangicus is close-

ly allied to N. daliensis, from which it differs particular-

ly by the more slender dorso-lateral apophyses of the

aedeagus. It is reliably distinguished from the geograph-

ically close N. barbatus, which too was found only in the

Wuliang Shan, only by the completely different male sex-

ual characters. The syntopic N. fissus is smaller and of

more slender habitus, of paler coloration, has a more

coarsely punctate head, a less densely punctate abdomen,

and completely different male sexual characters.

Distribution and natural history. Nazeris wuliangicus

was collected in two localities, one in the Wuliang Shan

and the other in an adjacent mountain range to the east of

Weishan, Yunnan (Fig. 88). The specimens were sifted

from leaf litter in shrub habitats at altitudes of 2450-2660

m. Several specimens collected in September are teneral.

©ZFMK

On the Nazeris fauna of China II 145

Nazeris nivimontis sp. n. (Figs 67—71, 88)

Type material. Holotype @ [teneral]: “CHINA: Yunnan,

Lincang Pref., Xue Shan, 48 km N Lincang, 2070 m,

24°19°03”N, 100°07°13”E, forest remnant, N-slope, lit-

ter & mushrooms sifted, 12.1X.2009, leg. M. Schilke

[CH09-45] / Holotypus ¢ Nazeris nivimontis sp. n. det.

V. Assing 2013” (cAss). Paratype <: [teneral]: same da-

ta as holotype (cSch).

Etymology. The specific epithet is a noun composed of

the Latin noun for snow (nix, nivis) and the genitive of

the Latin noun for mountain (mons). It is derived from the

name of the mountain (Xue Shan = Snow Mountain)

where the species is probably endemic.

Description. Body length 5.6—5.9 mm; length of forebody

3.1-3.3 mm. Head (Fig. 67) 1.00—1.03 times as long as

broad. Pronotum (Fig. 67) 0.85 times as broad as head.

Other external characters as in N. wuliangicus.

3: sternite VII (Fig. 68) with weakly concave posterior

margin, otherwise unmodified; sternite VII (Fig. 69) ap-

proximately 1.1 times as broad as long, posterior excision

rather small and approximately V-shaped, but anteriorly

rounded, not acute, nearly 0.2 times as deep as length of

sternite; aedeagus (Figs 70-71) approximately 0.9 mm

long; ventral process short, apically acute in ventral view;

dorso-lateral apophyses moderately long and slender, dis-

tinctly extending beyond apex of ventral process.

Comparative notes. Based on the male primary and sec-

ondary sexual characters, N. nivimontis is undoubtedly

closely related to N. daliensis and allied species. The shape

of the ventral process of the aedeagus of N. nivimontis

(ventral view) is remarkably similar to that of N. wuliang-

icus, from which N. nivimontis is distinguished by slight-

ly larger body size, a larger head, the different shape of

the posterior excision of the male sternite VII, and by the

relatively shorter dorso-lateral apophyses.

Distribution and natural history. The type locality is sit-

uated in the Xue Shan to the north of Lincang, Yunnan

(Fig. 88). Both type specimens are teneral; they were sift-

ed from forest leaf litter at an altitude of 2070 m.

Nazeris sagittifer sp. n. (Figs 76-80, 88)

Type material. Holotype @: “CHINA: Yunnan, Pu’er

Pref., Ailao Shan, 37 km NW Jingdong, 24°45712”N,

100°41°24.5”E, 2300 m, devastated forest remnant, litter

& dead wood sifted, 13.1X.2009, leg. M. Schiilke [CH09-

48] / Holotypus ¢ Nazeris sagittifer sp. n. det. V. Assing

Bonn zoological Bulletin 62 (2): 125-170

2013” (cAss). Paratypes: 7, 3° [partly teneral]: same da-

ta as holotype (ZFMK, cSch, cAss); 39 [partly teneral]:

“CHINA (Yunnan) Pu’er Pref., Ailao Shan, 37 km NW

Jingdong, 24°45°12”N, 100°41°24.5”E, 2300 m (devas-

tated forest remnant, litter, moss, grass roots sifted)

13.1X.2009 D.W. Wrase [48]” (cSch, cAss); 24: ‘““CHI-

NA: Yunnan, Lincang Pref., Wuliang Shan, old pass road,

W side, 24°42°58.6”N, 100°29°52.0”E, 2200 m, small

creek valley with primary forest remnant, litter sifted,

16.1X.2009, leg. M. Schiilke [CH09-47a]” (cSch, cAss);

1%, 22: same data, but “litter & debris sifted, ...,

12.X.2009 ... [CH09-47]” (cSch, cAss); 19: same data,

as before, but leg D.W. Wrase (cSch); 24: “CHINA: Yun-

nan, Lincang/Dali Pref., Wuliang Shan, old pass road, N

pass, 24°45°16.4”N, 100°29°50.3”E, 2350 m, forest lit-

ter & tea plantation, litter, mushrooms, grass sifted,

16.1X.2009, leg. M. Schiilke [CH09-55]” (cSch, cAss).

Etymology. The specific epithet (Latin, adjective: carry-

ing an arrow) alludes to the shape of the ventral process

of the aedeagus, which in ventral view somewhat resem-

bles an arrowhead.

Description. External characters as in N. wuliangicus.

&: sternite VII (Fig. 76) with posterior margin distinct-

ly concave in the middle, otherwise unmodified; sternite

VII (Fig. 77) approximately as long as broad, posterior

excision narrowly V-shaped, its depth approximately 0.25

times the length of sternite; aedeagus (Figs 78-79) approx-

imately 0.95 mm long; ventral process (Fig. 80) moder-

ately short, apically acute, shaped like an arrowhead in

ventral view, laterally somewhat compressed; dorso-lat-

eral apophyses long and slender, distinctly extending be-

yond apex of ventral process.

Comparative notes. Based on the virtually identical ex-

ternal and the similar male sexual characters, NV. sagittifer

is closely related to the geographically close, apparently

parapatric N. wuliangicus, from which it is reliably dis-

tinguished only by the male sexual characters, especially

the more distinctly concave posterior margin of the male

sternite VII, the deeper posterior excision of the male ster-

nite VIII, and the longer aedeagus with a differently

shaped ventral process and with longer dorso-lateral

apophyses.

Distribution and natural history. Nazeris sagittifer is

known from one locality in the Ailao Shan and two local-

ities in the Wuliang Shan, Yunnan (Fig. 88). The speci-

mens were sifted from forest leaf litter at altitudes of

2200-2350 m, on two occasions together with N. fissus.

Some of the type specimens are teneral.

©ZFMK

146 Volker Assing

Nazeris secatus sp. n. (Figs 81—88)

Type material. Holotype ¢: “CHINA: Yunnan, Lincang

Pref., Laobie Shan, Wei Bo Shan pass, 24°08°167N,

99°42°53”E, 2375 m, creek valley, devastated second. de-

cid. forest, litter & moss sifted, 8.1X.2009, leg. M. Schiilke

[CH09-35] / Holotypus ¢ Nazeris secatus sp. n. det. V.

Assing 2013” (cAss). Paratypes: 34, 49 [3 teneral]: same

data as holotype (cSch, cAss).

Etymology. The specific epithet is the past participle of

the Latin verb secare (to cut off) and refers to the trun-

cate lateral portions of the posterior margin of the male

sternite VIII.

Description. Body length 5.2—5.8 mm; length of forebody

3.0-3.3 mm. Habitus as in Fig. 81. Coloration: body black;

legs yellowish; antennae yellowish, with antennomere I

slightly darker.

Head (Fig. 82) 1.01—1.04 times as long as broad; me-

dian dorsal portion elevated; punctation moderately

coarse, dense, and umbilicate; interstices without mi-

crosculpture; eyes strongly convex and at least one third

as long as the distance from posterior margin of eye to pos-

terior constriction of head, usually somewhat longer. An-

tenna approximately 1.8 mm long.

Pronotum (Fig. 82) approximately 1.15 times as long

as broad and 0.85—0.90 times as broad as head; puncta-

tion dense and non-umbilicate, much coarser than that of

head; interstices glossy, forming narrow ridges; midline

punctate in anterior half, impunctate and narrowly elevat-

ed in posterior half.

Elytra (Fig. 82) approximately 0.55 times as long as

pronotum; humeral angles obsolete; punctation similar to

that of pronotum; interstices glossy. Hind wings complete-

ly reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense, defined, and coarse on tergites HI—VI,

somewhat finer and sparser on tergites VII and VII; in-

terstices without microsculpture and glossy; posterior mar-

gin of tergite VII without palisade fringe; posterior mar-

gin of tergite VIL strongly convex.

3: sternite VII (Fig. 83) with posterior margin weakly

concave in the middle, otherwise not distinctly modified;

sternite VIII (Fig. 84) approximately 1.05 times as broad

as long, lateral portions of posterior margin obliquely trun-

cate, posterior excision moderately deep and U-shaped,

approximately 0.2 times as deep as length of sternite;

aedeagus (Figs 85-86) 0.95—1.0 mm long; ventral process

(Fig. 87) long and apically acute in ventral view, lateral-

ly compressed; dorso-lateral apophyses moderately long

and weakly curved, slightly extending beyond apex of ven-

tral process.

Bonn zoological Bulletin 62 (2): 125-170

Comparative notes. The shapes of the male sternites VII

and VIII are similar to those of the species allied to N.

daliensis, but the ventral process of the aedeagus is much

longer in relation to the dorso-lateral apophyses. Nazeris

secatus is distinguished from the syntopic N. /anuginosus

by slightly smaller body size, the relatively larger and

more convex eyes, and the completely different male sex-

ual characters (shape and chaetotaxy of the male sternite

VII; sternite VIII with less deep posterior excision and

with obliquely truncate lateral portions of the posterior

margin; ventral process of the aedeagus less slender in

ventral view; dorso-lateral apophyses more slender, less

strongly sclerotized, and shorter in relation to ventral

process).

Distribution and natural history. The type locality is sit-

uated in the Laobie Shan (Fig. 88). The partly teneral type

specimens were sifted from litter in a secondary decidu-

ous forest at an altitude of 2375 m, together with NV. /anug-

inOSUS.

Nazeris cangicus sp. n. (Figs 89, 91-97)

Type material. Holotype ¢: “CHINA: N-Yunnan [C03-

19], Dali Bai Nat. Aut. Pref., Diancang Shan, 3 km W Dali

old town, creek valley and pine forest at “Cloud Road”,

right upper chairlift station, 25°41.1°N, 100°06.8’E,

2650-2750 m / [C03-19] litter, pine needles, moss (dry

and wet), mushrooms, 29.VIHI.2003, leg. M. Schilke /

Holotypus ¢ Nazeris cangicus sp. n. det. V. Assing 2013”

(cAss). Paratypes: 1<, 1 2: same data as holotype (cSch);

1¢: “CHINA: N-Yunnan Dali Bai Aut. Pref. Diancang

Shan 3 km W Dali, 25°41.1°N, 100°06.8’E, 2650-2750

m / 29.8.03, A. Smetana [C140]” (cAss); 34: “CHINA:

N-Yunnan [C03-20], Dali Bai Nat. Aut. Pref., Diancang

Shan, 4 km W Dali old town, 25°41.4’°N, 100°06.7’E,

2900-3000 m, E-slope with devastated forest and old pine

forest, mushrooms, 31.VIII.2003, leg. M. Schiilke” (cSch,

cAss); 19 [teneral]: “CHINA (N-Yunnan) Dali Bai Nat.

Aut. Pref., Diancang Shan, 4 km W Dali old town,

2900-3000 m, 25°41.4’N, 100°06.7’E, E-slope, (edge of

road, slope with Salix, bamboo, knotgrass, leaf litter, sift-

ed), 22.VI.2005 D.W. Wrase [15]” (cSch); 23, 19: “CHI-

NA: Yunnan [CH07-03], Dali Bai Auton. Pref., Diancang

Shan W Dali, 25°41°49”N, 100°06°24”E, 2970 m, sift-

ed at rock edges and under shrubs, 28.V.2007, M. Schiilke”

(cSch, cAss); 14, 39: “CHINA: Yunnan [CH07-04], Dali

Bai Auton. Pref., Diancang Shan W Dali, 25°41°47”N,

100°06’732”E, 3016 m, moist escarpment, litter sifted,

28.V.2007, M. Schiilke” (cSch, cAss); 14: “CHINA: Yun-

nan, Dali Bai Aut. Pref., Diacang [sic] Shan, W Dali,

25°41°52”"N, 100°06’28”E, 2960 m, along path, sifted

from litter, moss, flood debris, 6.1X.2009, leg. M.

Schiilke [CH09-31]” (cAss); 1¢, 19: “CHINA (Yunnan)

©ZFMK

On the Nazeris fauna of China II 147

Kunming

Fig. 89. Distributions of Nazeris species in Yunnan: N. cangicus (open circles); N. aculeatus (filled circle); N. zhemoicus (filled

diamond); N. pungens (open diamond); N. /anuginosus (filled triangle); N. peniculatus (filled star); N. barbatus (open star); N.

curvus (open triangles); N. suwbdentatus (open squares).

Dali Bai Aut. Pref., Diancang Shan E pass, 43 km NW

Dali, 2700 m, (secondary pine forest, litter, moss sifted),

25°59’50”N, 100°00°30”E, 23.VIH.2009 D.W. Wrase

[02]” (cSch, cAss); 19: “CHINA: Yunnan, Dali Bai Aut.

Pref., Diancang Shan, E pass, 43 km NW Dali,

25°59’°50”N, 100°00°30”E, 2700 m, secondary pine for-

est, litter, moss and mushrooms sifted, 23. VII.2009, leg.

M. Schiilke [CH09-02]” (ZFMK); 12, 29: “CHINA (Yun-

nan) Dali Bai Aut. Pref., Diancang Shan E pass, 43 km

NW Dali, 3104 m, (oak shrubs, Rhod., bamboo, litter sift-

ed), 25°59’33.5”N,.99°52712.5”"E, 23.VIIL2009 D.W.

Wrase [01A]” (cSch, cAss); 24, 3 [partly slightly ten-

eral]: “CHINA: Yunnan [CH07-08], Dali Bai Auton. Pref.,

Diancang Shan 43 km NW Dali, 3078 m, 25°59°35”N,

99°52’06”E, W pass, Rhodod., oaks, bamboo, sifted,

29.V.2007, M. Schiilke” (cSch, cAss); 34: “P.R. CHINA,

Yunnan, E slope Cangshan at Dali, N25°40°15.5”,

E100°07°45.4”’, 18.v.2010, 2657 m, sifting17, V.Greben-

nikov” (CAS, cAss); 14, 49: “PR. CHINA, Yunnan, E

slope Cangshan at Dali, N25°40713.2”’, E100°07°54.8”,

11.v.2010, 2728 m, sifting05, V.Grebennikov” (CAS,

cSme); 19: “P.R. CHINA, Yunnan, E slope Cangshan at

Dali, N25°407°15.1°, E100°07°39.9”’, 10.v.2010, 2711 m,

sifting04, V.Grebennikov” (CAS); 19: “P.R. CHINA, Yun-

nan, E slope Cangshan at Dah, N25°40°13.2”,

Bonn zoological Bulletin 62 (2): 125—170

E100°07°54.8”, 13.v.2010, 2728 m, sifting08, V.Greben-

nikov” (CAS); 1¢: “P.R. CHINA, Yunnan, E slope Cang-

shan at Dali, N25°40°13.2”, E100°07°54.8”, 9.v.2010,

2728 m, sifting01, V.Grebennikov” (cAss); 19: “CHINA:

Yunnan [CH07-04], Dali Bai Auton. Pref., Diancang Shan

W Dali, 25°41°47”°N, 100°06°32”E, 3016 m, moist es-

carpment, litter sifted, 28.V.2007, leg. A. Piitz” (cPiit); 12,

29: “CHINA: Yunnan, above Dali, 2500-2700 m,

9—18.1V.1999, leg. W. Schawaller” (SMNS, cAss).

Etymology. The specific epithet is an adjective derived

from Cang (= Diancang) Shan, the name of the mountain

range where the species was discovered.

Description. Body length 6.0—7.0 mm; length of forebody

3.1—3.5 mm. Habitus as in Fig. 91. Coloration: body black-

ish; legs yellowish; antennae yellowish, with antennomere

I shghtly darker.

Head (Fig. 92) 1.05—1.10 times as long as broad; me-

dian dorsal portion more or less distinctly elevated, 1.e.,

above the level of the frons; punctation coarse, dense, and

umbilicate (Fig. 93); interstices without microsculpture;

eyes moderately small, approximately one third as long

as the distance from posterior margin of eye to posterior

constriction of head. Antenna 1.7—1.9 mm long.

OZFMK

148 Volker Assing

-Kunmin

= g

Oxe

Fig. 90. Distributions of Nazeris species in Yunnan: N. firmilobatus (open circles); N. spiculatus (open diamond); N. infractus

(filled circle); N. meilicus (filled triangles); N. vexillatus and N. hastatus (open triangles); N. circumclusus (filled diamond); N.

bangmaicus (filled star); N. fissus (open stars); N. giganteus (open square).

Pronotum (Fig. 92) approximately 1.15 times as long

as broad and 0.9 times as broad as head; punctation ap-

proximately as coarse as that of head, non-umbilicate, and

moderately dense; interstices narrower than diameter of

punctures, glossy; midline punctate in anterior half, im-

punctate and narrowly elevated in posterior half.

Elytra (Fig. 92) 0.60—0.65 times as long as pronotum;

humeral angles obsolete; punctation dense, moderately de-

fined, and approximately as coarse as that of pronotum;

interstices glossy. Hind wings completely reduced.

Abdomen approximately 1.25 times as broad as elytra;

punctation dense, defined, and moderately coarse on an-

terior tergites, gradually becoming less dense and finer to-

wards posterior tergites, sparse and fine on tergite VII,

even finer and sparser on tergite VIII; interstices without

microsculpture on tergites IU—VI, with very shallow mi-

croreticulation on tergites VII-VHI; posterior margin of

tergite VI without palisade fringe; posterior margin of ter-

gite VIII weakly convex.

¢: sternite VII (Fig. 94) with posterior margin distinct-

ly concave in the middle; sternite VHI (Fig. 95) approx-

imately as long as broad, posterior excision deep and nar-

row, approximately 0.35 times as deep as length of ster-

Bonn zoological Bulletin 62 (2): 125-170

nite; aedeagus (Figs 96-97) 0.9—-1.0 mm long; ventral

process apically very acute in ventral view, laterally com-

pressed; dorso-lateral apophyses rather short, slightly ex-

tending beyond apex of ventral process.

Comparative notes. Nazeris cangicus is distinguished

from the syntopic N. daliensis, a species of similar size,

by the coarser, deeper, and less dense punctation of the

head, by the more or less distinctly elevated median por-

tion of the head, by the shapes of the male sternites VII

and VIII, as well as by the morphology of the aedeagus,

particularly the much shorter dorso-lateral apophyses. For

illustrations of the head and the male sexual characters of

N. daliensis see Figs 61-64.

Together with several other species recorded from Yun-

nan, N. cangicus forms a group characterized by more or

less blackish body colour, a male sternite VII mostly with

a distinct posterior concavity and/or a postero-median im-

pression, a male sternite VIII often with a median impres-

sion and a rather deep and narrow posterior excision, and

an aedeagus with a slender, laterally more or less com-

pressed, and apically acute ventral process and with short

to moderately long dorso-lateral apophyses.

©ZFMK

On the Nazeris fauna of China II 149

100

Figs 91-104. Nazeris cangicus (91-97) and N. aculeatus (98-104). 91, 98: habitus; 92, 99: forebody; 93: median dorsal portion

of head; 94, 100: male sternite VII; 95, 101: male sternite VII; 96-97, 102-103: aedeagus in lateral and in ventral view; 104: ven-

tral process of aedeagus in ventral view. Scale bars: 91-92, 98-99: 1.0 mm; 94-97, 100-103: 0.5 mm, 93, 104: 0.2 mm.

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

150 Volker Assing

Distribution and natural history. The type specimens

were discovered in several localities in the Diancang Shan

(Fig. 89). They were sifted from litter and moss in vari-

ous kinds of forests and shrub habitats at altitudes of

2650-3100 m, partly together with N. daliensis.

Nazeris aculeatus sp. n. (Figs 89, 98-104)

Type material. Holotype ¢: “CHINA: Yunnan, Dali Bai

Aut. Pref., Jizu Shan, summit plateau, 37 km NE Dali,

25°58°30°N, 100°21°36”E, 3150 m, mixed forest, sifted

from litter, moss & mushrooms, 5.1X.2009, leg. M.

Schiilke [CH09-28] / Holotypus ¢ Nazeris aculeatus sp.

n. det. V. Assing 2013” (cAss). Paratypes: 19: same da-

ta as holotype (cSch); 3%, 1 2: “CHINA (Yunnan) Dali Bai

Aut. Pref., Jizu Shan, summit plateau, 37 km NE Dali

3150 m, (mixed forest, sifted from litter, moss),

25°58°30”N, 100°21°36”E, 5.1X.2009 D.W. Wrase [28]”

(cSch, cAss).

Etymology. The specific epithet (Latin, adjective: with

sting) alludes to the very slender and apically acute ven-

tral process of the aedeagus in ventral view.

Description. Body length 5.8—6.8 mm; length of forebody

3.1—3.4 mm. Habitus as in Fig. 98. Coloration: body black-

ish-brown to blackish; legs yellowish; antennae yellow-

ish, with antennomere | slightly darker.

Head (Fig. 99) 1.05—1.10 times as long as broad; me-

dian dorsal portion weakly elevated; punctation moder-

ately coarse, dense, and umbilicate; interstices without mi-

crosculpture; eyes moderately small, approximately one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna approximately

1.8 mm long.

Pronotum (Fig. 99) approximately 1.15—1.20 times as

long as broad and 0.9 times as broad as head; punctation

approximately as coarse as that of head, non-umbilicate,

and dense; interstices distinctly narrower than diameter of

punctures, glossy; midline punctate in anterior half, im-

punctate and narrowly elevated in posterior half.

Elytra (Fig. 99) approximately 0.6 times as long as

pronotum; humeral angles obsolete; punctation dense,

moderately defined, and approximately as coarse as that

of pronotum; interstices glossy. Hind wings completely re-

duced.

Abdomen 1.2 times as broad as elytra; punctation dense,

defined, and moderately coarse on anterior tergites, grad-

ually becoming less dense and finer towards posterior ter-

gites, sparse and fine on tergite VII, even finer and spars-

er on tergite VIH; interstices without microsculpture on

tergites II-VI, with very shallow microreticulation on ter-

gites VII-—VIII; posterior margin of tergite VII without pal-

isade fringe; posterior margin of tergite VIII convex.

Bonn zoological Bulletin 62 (2): 125-170

&: posterior margin of sternite VII with broadly V-

shaped median excision (Fig. 100); sternite VIII (Fig. 101)

approximately as long as broad, posterior excision deep

and narrow, approximately 0.35 times as deep as length

of sternite; aedeagus (Figs 102—104) approximately 1.0

mm long; ventral process narrow and apically very acute

in ventral view, laterally compressed in dry preparation,

lateral parts somewhat dilated in ventral view (in micro-

scopic preparation); dorso-lateral apophyses of moderate

length, slightly extending beyond apex of ventral process.

Comparative notes. Based on the similar external and

particularly the similar male sexual characters, Nazeris

aculeatus 1s closely related to, and probably the adelpho-

taxon of, N. cangicus, from which it differs by the less

coarse and slightly denser punctation of the forebody, the

shape of the posterior excision of the male sternite VI,

and by the morphology of the aedeagus (ventral face of

the ventral process basally more slender, lateral parts of

ventral process of different shape). It is distinguished from

the sympatric N. jizushanensis by the slightly more coarse-

ly punctate and less matt head, the less densely and less

coarsely punctate pronotum, the less coarsely and less

densely punctate abdomen, the presence of shallow mi-

crosculpture on tergites VII and VIL (absent in N. jizusha-

nensis), the completely different shapes of the male ster-

nites VII and VIII, and by the different morphology of the

aedeagus (shape of venrtral process both in ventral and

in lateral view; dorso-lateral apophyses much shorter in

relation to ventral process).

Distribution and natural history. The type locality is sit-

uated nearly 40 km to the northeast of Dali, on the sum-

mit of the Jizu Shan (Fig. 89) at an altitude of 3150 m.

The specimens were sifted from litter and moss in a mixed

forest.

Nazeris zhemoicus sp. n. (Figs 89, 105—110)

Type material. Holotype ¢ [teneral]: “CHINA: Yunnan,

Dali Bai Aut. Pref., Zhemo Shan, 7 km SW Xiaguan,

25°32-33'N, 100°10-11’E, 2870-2970 m, scrub with bam-

boo, oaks & Rhododendr., litter sifted, 18.1X.2009, leg.

M. Schiilke [CH09-60] / Holotypus 4 Nazeris zhemoicus

sp. n. det. V. Assing 2013” (cAss). Paratypes: 22: same

data as holotype (cSch, cAss).

Etymology. The specific epithet is an adjective derived

from the name of the mountain where the species is prob-

ably endemic.

Description. Body length 5.4—5.7 mm; length of forebody

3.0-3.1 mm. Habitus as in Fig. 105. External characters

(Fig. 106) as in N. cangicus.

©OZFMK

On the Nazeris fauna of China I

107

110 108

109

105

116 117

115

111

113 114 118

Figs 105-118. Nazeris zhemoicus (105-110), N. pungens (AM1-117), and N. peniculatus (118). 105, 111: habitus; 106, 112: fore-

body; 107, 113: male sternite VII; 108, 114: male sternite VIII; 109-110, 115-116: aedeagus in lateral and in ventral view; 117:

ventral process of aedeagus in ventral view; 118: postero-median portion of male sternite VI. Scale bars: 105-106, 111-112: 1.0

©ZFMK

mm; 107-110, 113-116: 0.5 mm, 117-118: 0.2 mm.

Bonn zoological Bulletin 62 (2): 125-170

152 Volker Assing

&: sternite VII (Fig. 107) with small postero-median im-

pression, posterior margin with distinct excision in the

middle; sternite VII (Fig. 108) approximately as long as

broad and with oblong median impression, posterior ex-

cision deep and rather narrow, approximately one third as

deep as length of sternite; aedeagus (Figs 109-110) ap-

proximately 0.9 mm long; ventral process short, lateral-

ly somewhat compressed; dorso-lateral apophyses simple,

distinctly extending beyond apex of ventral process.

Comparative notes. As can be inferred from the highly

similar external and particularly the male sexual charac-

ters (sternite VII with posterior impression and medially

excised posterior margin; sternite VIII with deep and nar-

row posterior excision; morphology of the aedeagus), N.

zhemoicus is closely related to N. cangicus, from which

it differs by the smaller aedeagus with a differently shaped

ventral process and with the dorso-lateral apophyses

longer in relation to the ventral process.

Distribution and natural history. The type locality is sit-

uated in the Zhemo Shan to the southwest of Xiaguan in

Yunnan (Fig. 89). The specimens were sifted from leaf lit-

ter in a shrub habitat with oak, bamboo, and rhododen-

dron at an altitude of 2870-2970 m, together with two fe-

males of a presumably undescribed species. The holotype

is distinctly teneral.

Nazeris pungens sp. n. (Figs 89, 111-117)

Type material. Holotype @: “CHINA: Yunnan, Lincang

Pref., Xue Shan, 11 km ENE Lincang, 2510 m,

23°55’01N, 100°11°17.5”E, second. pine forest with

Rhodod., small cleft with water, litter & mushrooms sift-

ed, 10.1X.2009, leg. M. Schiilke [CH09-39] / Holotypus

3 Nazeris pungens sp. n. det. V. Assing 2013” (cAss).

Paratypes: 214, 17 [partly teneral]: same data as holo-

type (cSch, cAss); 5¢, 49 [partly teneral]: “CHINA (Yun-

nan) Lincang Pref., Xue Shan, 11 km ENE Lincang, 2510

m, 23°55’017N, 100°11°17.5”E (second. pine forest with

Rhodod., small cleft with water, litter sifted) 10.LX.2009

D.W. Wrase [39]” (ZFMK, cSch, cAss).

Etymology. The specific epithet is the present participle

of the Latin verb pungere (to sting) and refers to the sting-

shaped (ventral view) ventral process of the aedeagus.

Description. Body length 5.5—6.5 mm; length of forebody

3.2-3.6 mm. Habitus as in Fig. 111. Coloration: body

black; legs yellowish; antennae yellowish, with anten-

nomere | slightly darker.

Head (Fig. 112) 1.03—1.09 times as long as broad; me-

dian dorsal portion weakly elevated; punctation moder-

ately coarse, dense, and umbilicate; interstices without mi-

Bonn zoological Bulletin 62 (2): 125—170

crosculpture; eyes moderately small, less than one third

as long as the distance from posterior margin of eye to pos-

terior constriction of head. Antenna |.9—2.0 mm long.

Pronotum (Fig. 112) 1.15—1.20 times as long as broad

and approximately 0.9 times as broad as head; punctation

dense and non-umbilicate, distinctly coarser than that of

head; interstices glossy, forming narrow ridges; midline

punctate in anterior half, impunctate and narrowly elevat-

ed in posterior half.

Elytra (Fig. 112) approximately 0.55 times as long as

pronotum; humeral angles obsolete; punctation similar to

that of pronotum; interstices glossy. Hind wings complete-

ly reduced.

Abdomen 1.20—1.25 times as broad as elytra; puncta-

tion dense, defined, and coarse on tergites II-VI, finer and

sparser on tergites VII and VIII; interstices without mi-

crosculpture and glossy; posterior margin of tergite VII

without palisade fringe; posterior margin of tergite VIII

strongly convex.

@: sternite VII (Fig. 113) with small and shallow pos-

tero-median impression, this impression with a cluster of

rather sparse black setae directed postero-mediad, poste-

rior margin weakly concave in the middle; sternite VII

(Fig. 114) approximately 1.1 times as broad as long, pos-

terior excision deep and narrow, approximately 0.35 times

as deep as length of sternite; aedeagus (Figs 115-117) ap-

proximately 1.05 mm long; ventral process long, conspic-

uously narrow and apically very acute in ventral view, lat-

erally compressed; dorso-lateral apophyses long and slen-

der, subapically bent and with lamellate dilatation, distinct-

ly extending beyond apex of ventral process.

Comparative notes. The shape and chaetotaxy of the

male sternite VII and the deep, narrow posterior excision

of the male sternite VIII suggest that NV. pungens belongs

to the group of species allied to N. cangicus. Unlike the

above species of this group, the dorso-lateral apophyses

of the aedeagus are much longer and more slender.

Distribution and natural history. The type locality 1s sit-

uated in the Xue Shan to the east-northeast of Lincang

(Fig. 89). The partly teneral type specimens were sifted

from litter in a secondary pine forest with rhododendron

at an altitude of 2510 m.

Nazeris lanuginosus sp. n. (Figs 89, 118, 119-123)

Type material. Holotype @: “CHINA: Yunnan, Lincang

Pref., Laobie Shan, Wei Bo Shan pass, 24°08°16”N,

99°42°53”E, 2375 m, creek valley, devastated second. de-

cid. forest, litter & moss sifted, 8.1X.2009, leg. M. Schiilke

[CH09-35] / Holotypus ¢ Nazeris lanuginosus sp. n. det.

V. Assing 2013” (cAss). Paratypes: 39 [1 teneral]: same

data as holotype (cSch).

©ZFMK

On the Nazeris fauna of China II 153

121

125

119

127

3 se sr

128 129 123

Figs 119-129. Nazeris lanuginosus (119-123) and N. peniculatus (124-129). 119, 126: male sternite VI; 120, 127: male stern-

ite VHI; 121-122, 128-129: aedeagus in lateral and in ventral view; 123: ventral process of aedeagus in ventral view; 124: habi-

tus; 125: forebody. Scale bars: 124-125: 1.0 mm; 119-122, 126-129: 0.5 mm, 123: 0.2 mm.

Etymology. The specific epithet is an adjective derived

from the Latin noun lanugo (downy beard, downy pubes-

cence) and alludes to the postero-median cluster of setae

on the male sternite VII.

Bonn zoological Bulletin 62 (2): 125-170

Description. Body length 5.8—6.2 mm; length of forebody

3.43.5 mm. Head weakly oblong, 1.01—1.06 times as long

as broad. Other external characters as in NV. pungens.

3: sternite VII (Fig. 119) with postero-median impres-

sion, this impression with a cluster of dense black setae

©OZFMK

154 Volker Assing

directed postero-mediad, posterior margin distinctly con-

cave in the middle; sternite VII (Fig. 120) 1.05 times as

broad as long, posterior excision deep and narrow, 0.34

times as deep as length of sternite; aedeagus (Figs

121-123) 1.05 mm long; ventral process long, narrow and

apically very acute in ventral view, laterally compressed;

dorso-lateral apophyses moderately long, stout, strongly

sclerotized, slightly dilated at basal third, and distinctly

extending beyond apex of ventral process.

Comparative notes. Based on the shapes and chaetotaxy

of the male sternites VII and VHI, as well as on the mor-

phology of the aedeagus, N. /anuginosus is most closely

related to N. pungens, from which it differs by the less ob-

long head, the denser postero-median cluster of black se-

tae and the pronounced posterior concavity of the male

sternite VII, and by the morphology of the aedeagus

(shapes of the ventral process and of the dorso-lateral

apophyses).

Distribution and natural history. The type locality is sit-

uated in the Laobie Shan (Fig. 89). The partly teneral type

specimens were sifted from litter in a secondary decidu-

ous forest at an altitude of 2375 m, together with N. se-

catus.

Nazeris peniculatus sp. n. (Fig. 89, 118, 124-129)

Type material. Holotype @: “CHINA: Yunnan [CH07-

31], Dali Bai Auton. Pref., mtn. range N Er Hai, 42 km

N Dali, 26°04’53”°N, 100°09°39”E, 2500-2550 m, NE

slope with oaks, litter sifted, 12.VI.2007, M. Schiilke /

Holotypus ¢ Nazeris peniculatus sp. n. det. V. Assing

2013” (cAss). Paratypes: 104, 119: same data as holo-

type (ZFMK, cSch, cAss).

Etymology. The specific epithet is an adjective derived

from the Latin noun peniculus (brush) and refers to the

conspicuous cluster of modified setae on the male stern-

ite VIL.

Description. Small species; body length 4.7—5.7 mm;

length of forebody 2.5—2.9 mm. Habitus as in Fig. 124.

Coloration: body black; legs yellowish; antennae yellow-

ish, with antennomere I slightly darker.

Head (Fig. 125) as long as broad or weakly oblong; me-

dian dorsal portion weakly elevated; punctation moder-

ately coarse, dense, and umbilicate; interstices without mi-

crosculpture; eyes moderately small, approximately one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna approximately

1.5 mm long.

Pronotum (Fig. 125) approximately 1.15 times as long

as broad and ().9 times as broad as head; punctures dense

Bonn zoological Bulletin 62 (2): 125—170

and non-umbilicate, of similar diameter as those of head,

but distinctly deeper; interstices glossy, forming narrow

ridges; midline punctate in anterior half, impunctate and

narrowly elevated in posterior half.

Elytra (Fig. 125) approximately 0.6 times as long as

pronotum; humeral angles obsolete; punctation similar to

that of pronotum; interstices glossy. Hind wings complete-

ly reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense, defined, and coarse on tergites II-VI,

much finer and sparser on tergites VII and VII than on

tergite VI; interstices without microsculpture and glossy;

posterior margin of tergite VII without palisade fringe;

posterior margin of tergite VIII strongly convex.

&: sternite VII (Fig. 126) with postero-median impres-

sion, this impression with a conspicuous cluster of distinct-

ly modified short and stout black setae (Fig. 118), poste-

riorly with dense long black setae, posterior margin dis-

tinctly concave in the middle; sternite VIII (Fig. 127)

weakly transverse, posterior excision very deep and nar-

rowly U-shaped, approximately 0.4 times as deep as length

of sternite; aedeagus (Figs 128-129) approximately 0.95

mm long; ventral process long, narrow and apically very

acute in ventral view; dorso-lateral apophyses slender,

somewhat sinuate at basal 2/5, apically obliquely truncate,

and not reaching apex of ventral process.

Comparative notes. Based on the external (punctation of

the forebody) and the male sexual characters (sternite VII

with postero-median impression and with concave poste-

rior margin; sternite VII with deep and narrow posterior

excision; ventral process of aedeagus slender, apically

acute, and extending beyond the apices of the dorso-lat-

eral apophyses), N. peniculatus belongs to the group of

species allied to NV. cangicus. It is distinguished from them

particularly by the deeper posterior excision of the male

sternite VIII, the morphology of the aedeagus, and (ex-

cept for NV. barbatus) by the conspicuous chaetotaxy of the

male sternite VII.

Distribution and natural history. The type locality is sit-

uated in a mountain range to the north of the Er Hai lake

(Fig. 89). The specimens were sifted from oak leaf litter

at an altitude of 2500-2550 m.

Nazeris barbatus sp. n. (Figs 89, 130-136)

Type material. Holotype @: “CHINA: Yunnan, Dali Bai

Aut. Pref., Wuliang Shan, 20 km NW _ Weishan,

25°19’58"N, 100°07°59”E, 1900 m, creek valley, litter

& old flood debris sifted, 17.[X.2009, leg. M. Schiilke

[CH09-58] / Holotypus @ Nazeris barbatus sp. n. det. V.

Assing 2013” (cAss). Paratypes: 14, 12: same data as

holotype (cSch).

©ZFMK

On the Nazeris fauna of China I 155

134

130

133

137 141 142

139 140 132

Figs 130-142. Nazeris barbatus (130-136) and N. firmilobatus (137-142). 130, 137: habitus; 131, 138: forebody; 132, 139: male

sternite VII; 133: postero-median portion of male sternite VII; 134, 140: male sternite VIII; 135-136, 141-142: aedeagus in lat-

eral and in ventral view. Scale bars: 130-131, 137-138: 1.0 mm; 132, 134-136, 139-142: 0.5 mm, 133: 0.2 mm.

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

156 Volker Assing

Etymology. The specific epithet (Latin, adjective: beard-

ed) refers to the conspicuous cluster of modified setae on

the male sternite VII.

Description. Body length 5.2—5.7 mm; length of forebody

2.8—3.1 mm. Habitus as in Fig. 130. Coloration: forebody

dark-brown; abdomen blackish-brown; legs yellowish; an-

tennae yellowish, with antennomere | slightly darker. Oth-

er external characters (Fig. 131) as in N. peniculatus, ex-

cept for the on average shallower punctation of the head.

¢: sternite VII (Fig. 132) with postero-median impres-

sion, this impression with a cluster of distinctly modified

short and stout black setae (Fig. 133), posteriorly with

dense long black setae, posterior margin in the middle with

distinct concavity, middle of this concavity with a trian-

gular projection; sternite VIII (Fig. 134) approximately as

long as broad, posterior excision deep and narrowly U-

shaped, approximately one third as deep as length of ster-

nite; aedeagus (Figs 135-136) approximately 1.05 mm

long; ventral process long, narrow and apically very acute

in ventral view; dorso-lateral apophyses slender, somewhat

dilated in the middle, apically obliquely truncate, and not

reaching apex of ventral process.

Comparative notes. As can be inferred from the similar

external and particularly the similar male sexual charac-

ters, above all the similarly derived chaetotaxy of the male

sternite VII (unique among the Nazeris species known

from Yunnan) and the similarly derived morphology of the

aedeagus (long and slender ventral process), N. barbatus

is most closely related to N. peniculatus, from which it

differs by the median projection in the posterior concav-

ity of the male sternite VII, the less deep posterior exci-

sion of the male sternite VIII, and the longer aedeagus with

differently shaped dorso-lateral apophyses.

Distribution and natural history. The type locality is sit-

uated in the Wuliang Shan, to the northwest of Weishan,

Yunnan (Fig. 89). The specimens were sifted from litter

and old flood debris at an altitude of 1900 m.

Nazeris firmilobatus sp. n. (Figs 90, 137-142)

Type material. Holotype ¢: “P.R. CHINA, Yunnan, E

slope N Gaoligongshan, N27°45°40.8” E098°35’03.2”,

03.v1.2010, 2536 m, sifting 27, V. Grebennikov / Holoty-

pus ¢ Nazeris firmilobatus sp. n. det. V. Assing 2013”

(CAS). Paratypes: 2%, 39: same data as holotype (CAS,

cSme, cAss); 2, 39: “PR. CHINA, Yunnan, E slope N

Gaoligongshan, N27°45°27.1” E098°357°34.5”’,

02.v1.2010, 2600 m, sifting26, V. Grebennikov” (CAS,

cSme, cAss); 24, 19: “P.R. CHINA, Yunnan, E slope N

Gaoligongshan, N27°46.8’ E098°33.1’, 12—15.vi.2009,

2000-3000 m, sifting l—7, V. Grebennikov” (CAS, cSme,

cAss).

Bonn zoological Bulletin 62 (2): 125—170

Etymology. The specific epithet is an adjective composed

of the Latin adjectives firmus (stout, firm) and lobatus

(lobed). It refers to the stout dorso-lateral apophyses of

the aedeagus.

Description. Body length 6.0—7.0 mm; length of forebody

3.3—3.6 mm. Habitus as in Fig. 137. Coloration: body

blackish-brown to blackish; legs yellowish; antennae yel-

lowish, with antennomere | slightly darker.

Head (Fig. 138) approximately 1.05 times as long as

broad; median dorsal portion more or less distinctly ele-

vated; punctation coarse, dense, and umbilicate; interstices

without microsculpture; eyes moderately small, less than

one third as long as the distance from posterior margin of

eye to posterior constriction of head. Antenna approxi-

mately 2.0 mm long.

Pronotum (Fig. 138) approximately 1.2 times as long

as broad and 0.85 times as broad as head; punctation very

dense, coarser than that of head, non-umbilicate; inter-

stices glossy; midline punctate in anterior half, impunc-

tate and narrowly elevated in posterior half.

Elytra (Fig. 138) approximately 0.6 times as long as

pronotum; humeral angles obsolete; punctation dense, de-

fined, and approximately as coarse as that of pronotum;

interstices glossy. Hind wings completely reduced.

Abdomen approximately 1.25 times as broad as elytra;

punctation dense, defined, and coarse on tergites II-VI,

less dense and less coarse on tergite VII, finer and spars-

er on tergite VII; interstices without microsculpture; pos-

terior margin of tergite VII without palisade fringe; pos-

terior margin of tergite VHI convex.

&: sternite VII (Fig. 139) with small postero-median im-

pression, posterior margin distinctly concave in the mid-

dle; sternite VII (Fig. 140) approximately as long as

broad, with narrow, but long and distinct median impres-

sion, posterior excision deep and rather narrow, approx1-

mately one third as deep as length of sternite; aedeagus

(Figs 141-142) approximately 1.1 mm long; ventral

process narrow and apically acute in ventral view, later-

ally compressed; dorso-lateral apophyses stout, somewhat

extending beyond apex of ventral process.

Comparative notes. Nazeris firmilobatus is readily dis-

tinguished from the four previously described species dis-

tributed in the Gaoligong Shan by the distinctly stouter

and nearly straight dorso-lateral apophyses of the aedea-

gus.

Distribution and natural history. The species was found

in three geographically close localities in the northern

Gaoligong Shan (Fig. 90). The specimens with specified

altitudes on the labels were sifted at elevations of approx-

imately 2540 and 2600 m.

©ZFMK

On the Nazeris fauna of China II 157

Nazeris spiculatus sp. n. (Figs 90, 143-149)

Type material. Holotype ¢: “CHINA: Yunnan [CH07-

19], Dehong Dai Aut. Pref., mountain range 31 km E Luxi,

2280 m, 24°29731”N, 98°52’58”E, secnd. pine forest with

old decid. trees, litter sifted, 3.VI.3007, M. Schtilke / Holo-

typus 4 Nazeris spiculatus sp. n. det. V. Assing 2013”

(cAss). Paratypes: 1, 39: same data as holotype (cSch,

cAss); 24, 12: same data, but leg. A. Piitz (cPtit, cAss).

Etymology. The specific epithet is an adjective derived

from the Latin noun spiculum (thorn, spine) and refers to

the shape of the ventral process of the aedeagus in ven-

tral view.

Description. Body length 6.2—7.3 mm; length of forebody

3.4-3.8 mm. Habitus as in Fig. 143. Coloration: body

blackish-brown to blackish, with the elytra occasionally

paler; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker. External characters (Fig. 144) as

in N. firmilobatus, except for the denser punctation of the

abdominal tergite VII.

&: sternite VII (Fig. 145) with small postero-median im-

pression, posterior margin distinctly concave; sternite

VIII (Fig. 146) approximately as long as broad, with dis-

tinct median impression, posterior excision deep and nar-

rowly U-shaped, approximately one third as deep as length

of sternite; aedeagus (Figs 147-149) approximately 1.1

mm long; ventral process narrow and apically acute in

ventral view, laterally strongly compressed; dorso-later-

al apophyses basally strongly dilated and apically slender,

just reaching the apex of the ventral process.

Comparative notes. The similar external and male sex-

ual characters suggest that N. spiculatus is closely allied

to N. firmilobatus, N. curvus, and allied species, from

which it is reliably distinguished only by the shapes of the

male sternites VII and VIII, as well as by the morpholo-

gy of the aedeagus.

Distribution and natural history. The type locality is sit-

uated to the east of Luxi, western Yunnan (Fig. 90). The

specimens were sifted from litter in a secondary pine for-

est with old deciduous trees at an altitude of 2280 m.

Nazerts curvus sp. n. (Figs 89, 150-155)

Type material. Holotype ¢: “CHINA (Yunnan) Baoshan

Pref., Gaoligong Shan nr. Xiaoheishan N.R., 35 km SE

Tengchong, 2110 m, 24°50°16”N, 98°45°43”E (prim. de-

cid. forest, litter, sifted) 30.V.—14.V1.2007 D.W. Wrase

[11] / Holotypus ¢ Nazeris curvus sp. n. det. V. Assing

2013” (cAss). Paratypes: 1: “CHINA: Yunnan [CH07-

13A], Baoshan Pref., Gaoligong Shan, E pass, 36 km SE

Bonn zoological Bulletin 62 (2): 125—170

Tengchong, 2200 m, 24°49°32”N, 98°46°06”E, decid. for-

est, litter, wood, fungi sifted, 4.VI.2007, M. Schiilke”

(cSch); 19: same data, but “[CH07-13] ... 31.V.2007”

(cSch); 19: “CHINA: Yunnan, Baoshan Pref., Gaoligong

Shan, 32 km SE Tengchong, 2150-2250 m, 24°51-53°N,

98°45°E, devast. prim. and second. forest, litter, dead

wood, mushrooms sifted, 26. VIII.2009, leg. M. Schiilke

[CH09-08/09]” (cSch); 19: “CHINA: Yunnan, Baoshan

Pref., Gaoligong Shan, W pass 35 km SE Tengchong, 2100

m, 24°50°18”N, 98°45°43”E, devast. prim. dec. forest, lit-

ter, wood, mushrooms sifted, 28.VIII.2009, leg. M.

Schiilke [CH09-06a]” (cSch); 1¢: “CHINA: Yunnan

[CH07-14A], Baoshan Pref., Gaoligong Shan, 33 km SE

Tengchong, 2100-2200 m, 24°51°22”N, 98°45°36”E, de-

cid. forest, litter, wood, fungi sifted, 4.VI.2007, M.

Schiilke” (cAss); 19: “CHINA: Yunnan [CH07-15],

Baoshan Pref., Gaoligong Shan, 29 km ESE Tengchong,

2350 m, 24°55°37°N, 98°45°09”E, dev. decid forest, lit-

ter, wood, fungi sifted, 1.V1I.2007, leg. A. Piitz” (cPiit).

Etymology. The specific epithet (Latin, adjective: curved)

alludes to the strongly curved dorso-lateral apophyses of

the aedeagus.

Description. Body length 6.0—6.6 mm; length of forebody

3.13.5 mm. Habitus as in Fig. 150. Coloration: body

blackish-brown to black; legs yellowish; antennae yellow-

ish, with antennomere I slightly darker.

Head (Fig. 151) weakly oblong, 1.01—1.03 times as long

as broad; median dorsal portion elevated; punctation mod-

erately coarse and shallow, very dense, and distinctly um-

bilicate; interstices without microsculpture, reduced to

very narrow ridges; eyes rather small, less than one third

as long as the distance from posterior margin of eye to pos-

terior constriction of head. Antenna approximately 2.0 mm

long.

Pronotum (Fig. 151) approximately 1.15—1.20 times as

long as broad and 0.85—0.90 times as broad as head; punc-

tation very dense, coarse, much coarser than that of head,

non-umbilicate; interstices glossy, but very narrow; mid-

line punctate in anterior half, impunctate and narrowly el-

evated in posterior half.

Elytra (Fig. 151) approximately 0.55 times as long as

pronotum; humeral angles nearly obsolete; punctation

dense, defined, and approximately as coarse as that of

pronotum; interstices glossy. Hind wings completely re-

duced.

Abdomen approximately 1.2 times as broad as elytra;

punctation very dense, defined, and coarse on tergites

III-VI, only slightly less dense and less coarse on tergite

VU, finer and sparser on tergite VIII; interstices without

microsculpture; posterior margin of tergite VI without

palisade fringe; posterior margin of tergite VII strongly

convex.

OZFMK

158 Volker Assing

143 444

: 154 155

145 _ 152

Figs 143-155. Nazeris spiculatus (143-149) and N. curvus (150-155). 143, 150: habitus; 144, 151: forebody; 145, 152: male

sternite VII; 146, 153: male sternite VIII; 147-148, 154-155: aedeagus in lateral and in ventral view; 149: ventral process of aedea-

gus in ventral view. Scale bars: 143-144, 150-151: 1.0 mm; 145-148, 152-155: 0.5 mm, 149: 0.1 mm.

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

On the Nazeris fauna of China II 159

&: sternite VII (Fig. 152) with small postero-median im-

pression, posterior margin weakly concave; sternite VII

(Fig. 153) approximately 1.05 times as broad as long and

with oblong median impression, posterior excision rather

deep and moderately narrow, approximately 0.3 times as

deep as length of sternite; aedeagus (Figs 154—155) ap-

proximately 1.05 mm long; ventral process narrow and

apically acute in ventral view, laterally somewhat com-

pressed; dorso-lateral apophyses stout, strongly curved in

lateral view, and slightly extending beyond apex of ven-

tral process.

Comparative notes. The only other species with strong-

ly curved dorso-lateral apophyses in the Gaoligong Shan

is the geographically close N. nomurai Watanabe & X1-

ao, 2000 (type locality: “Lujiangba, Gaoligong Shan Mts.,

Baoshan area’’), from which N. curvus differs by the much

more slender ventral process (ventral view) and by the api-

cally only weakly modified dorso-lateral apophyses. Based

on the illustrations provided by Watanabe & Xiao (2000),

it seems likely that the types of N. nomurai and specimens

from “Dabei” listed as additional material, but not includ-

ed in the type series, are not conspecific.

Distribution and natural history. The known distribu-

tion of N. curvus is confined to several geographically

close localities to the southeast of Tengchong in the

Gaoligong Shan (Fig. 89). The specimens were sifted from

leaf litter in primary and secondary deciduous forests at

altitudes of 2100-2350 m, in one locality together with

N. circumclusus.

Nazeris infractus sp. n. (Figs 90, 156-162)

Type material. Holotype ¢: “CHINA: Yunnan [CH07-

30], Nujiang Lisu Aut. Pref., Nu Shan, 7 km NNW Coa-

jian, 25°43°29”N, 99°07°57”E, 2420 m, second. pine for-

est with shrubs, litter, bark sifted, 11.V1I.2007, M. Schilke

/Holotypus ¢ Nazeris infractus sp. n. det. V. Assing 2013”

(cAss). Paratypes: 29: same data as holotype (cSch); 19:

same data, but leg. Wrase (cSch); 19: same data, but leg.

Piitz (cPiit).

Etymology. The specific epithet is the past participle of

the Latin verb infringere (to bend, to break) and alludes

to the apically abruptly bent dorso-lateral apophyses of the

aedeagus.

Description. Body length 6.0—6.2 mm; length of forebody

3.2-3.5 mm. Habitus as in Fig. 156. Coloration: body

blackish; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker.

Bonn zoological Bulletin 62 (2): 125-170

Head (Fig. 157) as broad as long or weakly oblong; me-

dian dorsal portion elevated; punctation moderately

coarse and shallow, very dense, and distinctly umbilicate;

interstices without microsculpture, reduced to very nar-

row ridges; eyes moderately small, approximately one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna approximately

2.0 mm long.

Pronotum (Fig. 157) approximately |.15—1.20 times as

long as broad and approximately 0.85 times as broad as

head; punctation very dense, much coarser than that of

head, non-umbilicate; interstices glossy, but narrow; mid-

line punctate in anterior half, impunctate and narrowly el-

evated in posterior half.

Elytra (Fig. 157) approximately 0.55 times as long as

pronotum; humeral angles nearly obsolete; punctation

dense, defined, and approximately as coarse as that of

pronotum; interstices glossy. Hind wings completely re-

duced.

Abdomen approximately 1.2 times as broad as elytra;

punctation very dense, defined, and coarse on tergites

III—VI, only slightly less dense and less coarse on tergite

VU, finer and sparser on tergite VIII; interstices without

microsculpture; posterior margin of tergite VI without

palisade fringe; posterior margin of tergite VIII strongly

convex.

3: sternite VII (Fig. 158) with postero-median impres-

sion, posterior margin distinctly concave in the middle;

sternite VIII (Fig. 159) approximately 1.1 times as broad

as long and with oblong median impression, posterior ex-

cision rather deep and narrowly V-shaped, approximate-

ly one third as deep as length of sternite; aedeagus (Figs

160-162) 0.95 mm long; ventral process basally of mod-

erate width and apically acute in ventral view, laterally

somewhat compressed; dorso-lateral apophyses stout, api-

cally abruptly bent, and slightly extending beyond apex

of ventral process.

Comparative notes. In external morphology, this species

is highly similar to N. curvus. It is distinguished from all

its congeners particularly by the conspicuous shape of the

dorso-lateral apophyses of the aedeagus, from N. curvus

additionally by the medially distinctly concave posterior

margin of the male sternite VI, the deeper and different-

ly shaped posterior excision of the male sternite VHI, and

by the much broader ventral process of the smaller aedea-

gus.

Distribution and natural history. The type locality is sit-

uated in the Nu Shan, to the north-northwest of Coajian

(Fig. 90). The specimens were sifted from leaf litter in a

secondary pine forest with shrubs at an altitude of 2420

m. One of the females is teneral.

©ZFMK

160 Volker Assing

159

157 160 161

166

162

167 168

158 165

Figs 156-169. Nazeris infractus (156-162), N. subdentatus (163-168), and N. vexillatus (169). 156, 163: habitus; 157, 164: fore-

body; 158, 165: male sternite VII; 159, 166, 169: male sternite VIII; 160-161, 167-168: aedeagus in lateral and in ventral view;

162: ventral process of aedeagus in ventral view. Scale bars: 156-157, 163-164: 1.0 mm; 158-161, 165-169: 0.5 mm, 162: 0.1

mm.

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

On the Nazeris fauna of China II 161

Nazeris subdentatus sp. n. (Figs 89, 163—168)

Type material. Holotype @: “CHINA: Yunnan, Nujiang

Lisu Pref., Gaoligong Shan, “Cloud pass” 21 km NW

Liuku, 3150 m, 25°58’21”N, 98°41°01”E, shrubs & bam-

boo, litter sifted, 2.1X.2009, leg. M. Schitilke [CH09-22]

/ Holotypus ¢ Nazeris subdentatus sp. n. det. V. Assing

2013” (cAss). Paratypes: 1%: same data as holotype

(cSch); 29: “CHINA: Yunnan [CH07-27], Nujiang Lisu

Aut. Pref., Gaoligong Shan, creek valley 20 km NW

Liuku, 25°58°49”"N, 98°41°48”E, 3000 m, bamboo,

shrubs, litter sifted, 9.VI.2007, M. Schiilke” (cSch,

cAss); 1 [slightly teneral]: “CHINA: Yunnan [CH07-28],

Nujiang Lisu Aut. Pref., Gaoligong Shan, side valley 19

km NW Liuku, 25°59°02”N, 98°42°43”E, 2730 m, dev-

ast. prim. forest, litter sifted, 9.VI.2007, leg. A. Pitz”

(cPit).

Etymology. The specific epithet (Latin, adjective) alludes

to the dentate ventral process of the aedeagus, a charac-

ter distinguishing this species from all its geographically

close congeners.

Description. Body length 6.2—7.5 mm; length of forebody

3.55.7 mm. Habitus as in Fig. 163. Coloration: body

black; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker.

Head (Fig. 164) 1.05—1.10 times as long as broad; me-

dian dorsal portion elevated; punctation coarse, moderate-

ly dense, and non-umbilicate; interstices without mi-

crosculpture, glossy; eyes moderately small, approximate-

ly one third as long as the distance from posterior margin

of eye to posterior constriction of head. Antenna approx-

imately 2.0—2.3 mm long.

Pronotum (Fig. 164) small in relation to head, approx-

imately 1.15 times as long as broad and 0.80—0.85 times

as broad as head; punctation similar to that of head, but

slightly less dense; interstices glossy; midline with very

short elevated impunctate band posteriorly.

Elytra (Fig. 164) approximately 0.6 times as long as

pronotum, of conspicuously trapezoid shape, 1.e., poste-

riorly strongly dilated and with completely obsolete

humeral angles; punctation dense, defined, and somewhat

coarser than that of pronotum; interstices glossy. Hind

wings completely reduced.

Abdomen approximately 1.25 times as broad as elytra,

strongly widened from segment HI to segment VI; punc-

tation dense, defined, and coarse on tergites III—VI, only

slightly less dense and less coarse on tergite VII, finer and

sparser on tergite VIII; interstices without microsculpture;

posterior margin of tergite VII without palisade fringe;

posterior margin of tergite VIII convex.

&: posterior margin of sternite VII with small median

concavity (Fig. 165); sternite VIII (Fig. 166) approximate-

ly 1.1 times as broad as long, posterior excision deep and

Bonn zoological Bulletin 62 (2): 125-170

narrowly V-shaped, approximately one third as deep as

length of sternite; aedeagus (Figs 167—168) approximate-

ly 1.05 mm long; ventral process narrow in basal half,

sharply edged in apical half, and apically very acute in

ventral view, with small subapical tooth in lateral view;

dorso-lateral apophyses simple, rather short, not reaching

apex of ventral process.

Comparative notes. Nazeris subdentatus is distinguished

from all its geographically close congeners by the conspic-

uously trapezoid shape of the elytra, by the shape of the

male sternite VIII, and particularly by the shape of the ven-

tral process of the aedeagus (extending beyond dorso-lat-

eral apophyses; ventral portion sharply edged in apical

half; subapically dentate in lateral view).

Distribution and natural history. The species was found

in two geographically close localities in the northern

Gaoligong Shan, to the northwest of Liuku (Fig. 89). The

specimens were sifted from leaf litter in vegetation com-

posed of shrubs and bamboo and in a degraded primary

forest at altitudes of 2730-3150 m.

Nazeris meilicus sp. n. (Figs 90, 170-176)

Type material. Holotype @: “CHINA: N-Yunnan [C2005-

07A], Diqing Tibet. Aut. Pref., Deqin Co., Meili Xue Shan,

E-side, 12 km SW Deqin, 2890 m, 28°25.30°N,

98°48.47°E / small creek valley, mixed forest with bam-

boo, leaf litter, moss, dead wood, sifted, 13.VI.2005, leg.

M. Schiilke [C2005-07A] / Holotypus @ Nazeris meili-

cus sp. n. det. V. Assing 2013” (cAss). Paratypes: 29:

same data as holotype (cSch, cAss); 1: “CHINA (N-Yun-

nan) Diqing Tibet. Aut. Pref., Deqin Co., Meili Xue Shan,

E-side, 12 km SW Deqin, 2890 m, 28°25.30°N,

98°48.47°E, creek valley (mixed forest, under wood,

stones, in litter/soil) 9.&13.VI.2005, D.W. Wrase [07]”

(cSch); 14, 29: “CHINA: N-Yunnan [C2005-09], Diging

Tibet. Aut. Pref., Deqin Co., Meili Xue Shan, E-side, 14

km W Deqin, 2580 m / 28°27.47°N, 98°46.35°E, creek

valley below glacier, mixed forest, leaf litter, moss, dead

wood, sifted, 11.V1I.2005, leg. M. Schiilke [C2005-09]”

(cSch); 14, 22: “CHINA: N-Yunnan Diqing Tibet. Aut.

Pr. Deqin Co. Meili Xue Shan E-side 12 km SW Degin,

28°25.30°N, 98°48.47°E 2890 m, 13.VI.2005 A. Smeta-

na [C160]” (cSme, cAss); 1: same data, but “9.V1I.2005

... [C156]” (cAss); 12: “CHINA: N-Yunnan Diging Ti-

bet. Aut. Pr. Deqin Co. Meili Xue Shan E-side 14 km W

Degqin, 28°27.47°N, 98°46.35°E 2580 m, 11.VI.2005 A.

Smetana [C158]” (cSme).

Etymology. The specific epithet is an adjective derived

from the name of the mountain where this species was dis-

covered.

©ZFMK

162 Volker Assing

176

170 174 175

182

180 181

179

Figs 170-182. Nazeris meilicus (170-176) and N. vexillatus (177-182). 170, 177: habitus; 171, 178: forebody; 172, 179: male

sternite VII; 173: male sternite VIII; 174-175, 180-181: aedeagus in lateral and in ventral view; 176, 182: ventral process of aedea-

gus in ventral view. Scale bars: 170-171, 177-178: 1.0 mm; 172-175, 179-181: 0.5 mm, 176, 182: 0.1 mm.

Bonn zoological Bulletin 62 (2): 125—170 ©ZFMK

On the Nazeris fauna of China II 163

Description. Body length 5.5—6.7 mm; length of forebody

3.3-3.5 mm. Habitus as in Fig. 170. Coloration: forebody

dark-brown to black; abdomen black; legs yellowish; an-

tennae yellowish, with antennomere | slightly darker.

Head (Fig. 171) 1.05—1.10 times as long as broad; me-

dian dorsal portion weakly elevated; punctation moder-

ately coarse, dense, and umbilicate; interstices without mi-

crosculpture, forming very narrow ridges; eyes relative-

ly weakly convex, weakly projecting from lateral contours

of head, less than one third as long as the distance from

posterior margin of eye to posterior constriction of head.

Antenna approximately 2.0 mm long.

Pronotum (Fig. 171) approximately 1.2 times as long

as broad and 0.85 times as broad as head; punctation very

dense, much coarser than that of head, non-umbilicate; in-

terstices glossy, forming narrow ridges; midline punctate

in anterior half, impunctate and narrowly elevated in pos-

terior half.

Elytra (Fig. 171) approximately 0.6 times as long as

pronotum; humeral angles obsolete; punctation dense, de-

fined, and approximately as coarse as that of pronotum;

interstices glossy. Hind wings completely reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense, defined, and coarse on tergites II-VI,

distinctly finer and sparser on tergites VII and VIII; in-

terstices without microsculpture; posterior margin of ter-

gite VI without palisade fringe; posterior margin of ter-

gite VII convex.

G: sternite VII (Fig. 172) with weakly concave poste-

rior margin, otherwise unmodified; sternite VHI (Fig. 173)

approximately 1.1 times as broad as long, posterior exci-

sion broadly V-shaped, approximately 0.2 times as deep

as length of sternite; aedeagus (Figs 174-176) approxi-

mately 1.0 mm long; ventral process moderately short and

slender, apically acute in ventral view; dorso-lateral

apophyses stout, strongly sclerotized, triangularly dilated

in the middle, curved dorsad in lateral view, and distinct-

ly extending beyond apex of ventral process.

Comparative notes. This species is characterized partic-

ularly by the moderately deep and broadly V-shaped pos-

terior excision of the male sternite VIII and by the dis-

tinctive shape of the dorso-lateral apophyses of the aedea-

gus. The morphology of the aedeagus, in particular the

stout and subapically abruptly bent dorso-lateral apophy-

ses, 1s most similar to that of N. infractus, but the shapes

of the male sternites VII and VIII are rather different.

Distribution and natural history. Nazeris meilicus was

collected in two localities in the Meili Xue Shan in west-

ern Yunnan (Fig. 90). The specimens were sifted from lit-

ter in mixed forests at altitudes of 2580 and 2890 m.

Bonn zoological Bulletin 62 (2): 125-170

Nazeris vexillatus sp. n. (Figs 90, 169, 177-182)

Type material. Holotype @: “CHINA: Yunnan, Lujiang

Lisu Pref., Gaoligong Shan, “Cloud pass”, 21 km NW

Liuku, 25°58’21°N, 98°41’°01”E, 3150 m, shrubs & bam-

boo, litter sifted, 3.1X.2009, leg. M. Schiilke [CH09-22a]

/ Holotypus @ Nazeris vexillatus sp. n. det. V. Assing

2013” (cAss). Paratypes: 4%, 12: same data as holotype

(cSch, cAss); 19: “CHINA (Yunnan) Lujiang Lisu Pref.,

Gaoligong Shan “Cloud pass” 3150 m, 21 km NW Liuku

(shrubs, Vaccinium, bamboo, litter sifted) 25°58’217N,

98°41°01”E, 2.1X.2009 D.W. Wrase [22A]” (cSch); 12:

“CHINA: Yunnan, Lujiang Lisu Pref., Gaoligong Shan,

“Cloud pass”, 21 km NW _ Liuku, 25°58’21”N,

98°41°01”"E, 3150 m, shrubs & bamboo, litter sifted,

2.1X.2009, leg. M. Schiilke [CH09-22]” (cAss); 10, 29:

“CHINA: Yunnan [CH07-28A], Nujiang Lisu Aut. Pref.,

Gaoligong Shan, side valley 19 km NW = Liuku,

25°59’°02”N, 98°42’23”E, 2730 m, devast. prim. for., lit-

ter sifted, 10.VI.2007, M. Schiilke” (cSch, cAss); 12:

“CHINA: Yunnan [CHO07-28], Nujiang Lisu Aut. Pref.,

Gaoligong Shan, side valley 19 km NW _ Liuku,

25°59’°02”N, 98°42’23”E, 2730 m, devast. prim. forest,

litter sifted, 9.VI.2007, M. Schiilke” (cSch); 19: “CHI-

NA: Yunnan [CHO07-27], Nujiang Lisu Aut. Pref.,

Gaoligong Shan, creek valley 20 km NW Liuku,

25°58°49"N, 98°41°48” E, 3000 m, bamboo, shrubs, lit-

ter sifted, 9.VI.2007, M. Schiilke” (cAss); 1¢ [teneral]:

“CHINA (Yunnan) Lujiang Lisu Pref., Gaoligong Shan E

pass 20 km NW Liuku, 3000 m (creek valley with dev-

ast. prim. forest, ferns, litter and moss sift.) 25°58°49”N,

98°41°48”E, 3.1X.2009 D.W. Wrase [25]” (cSch).

Etymology. The specific epithet is an adjective derived

from the Latin noun vexillum (small flag, banneret) and

alludes to the conspicuous lamellate processes of the dor-

so-lateral apophyses of the aedeagus.

Description. Body length 5.2—6.7 mm; length of forebody

2.9-3.4 mm. Habitus as in Fig. 177. Coloration: head and

pronotum reddish-brown to dark-brown; elytra reddish to

dark-reddish; abdomen dark-brown to blackish-brown;

legs yellowish; antennae yellowish, with antennomere I

slightly darker.

Head (Fig. 178) approximately 1.05 times as long as

broad; median dorsal portion more or less distinctly ele-

vated; punctation coarse, dense, and moderately umbili-

cate; interstices without microsculpture; eyes moderate-

ly small, approximately one third as long as the distance

from posterior margin of eye to posterior constriction of

head. Antenna |.7—1.9 mm long.

Pronotum (Fig. 178) approximately 1.15 times as long

as broad and 0.85 times as broad as head; punctation coars-

er and somewhat sparser than that of head, non-umbili-

©OZFMK

164 Volker Assing

195

194

191 ve OM. 492

Figs 183-195. Nazeris circumclusus (183-188) and N. hastatus (189-195). 183, 189: habitus; 184, 190: forebody; 185, 191: male

sternite VII; 186, 192: male sternite VII; 187-188, 193-194: aedeagus in lateral and in ventral view; 195: ventral process of aedea-

gus in ventral view. Scale bars: 183-184, 189-190: 1.0 mm; 185-188, 191-194: 0.5 mm, 195: 0.2 mm.

Bonn zoological Bulletin 62 (2): 125-170 ©ZFMK

On the Nazeris fauna of China I] 165

cate; interstices glossy; midline punctate in anterior half,

impunctate and narrowly elevated in posterior half.

Elytra (Fig. 178) approximately 0.6 times as long as

pronotum; humeral angles obsolete; punctation dense, de-

fined, and approximately as coarse as that of pronotum;

interstices glossy. Hind wings completely reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense, defined, and moderately coarse on ter-

gite III, gradually becoming less dense and finer towards

posterior tergites, moderately sparse and fine on tergite

VII, even finer and sparser on tergite VIII; interstices with-

out microsculpture; posterior margin of tergite VII with-

out palisade fringe; posterior margin of tergite VIII strong-

ly convex.

&: sternite VII (Fig. 179) with truncate posterior mar-

gin; sternite VII (Fig. 169) weakly transverse, posterior

excision moderately deep and rather narrowly V-shaped,

approximately 0.25 times as deep as length of sternite;

aedeagus (Figs 180-182) 0.9-1.0 mm long; ventral

process gradually narrowed apicad, apically acute, and

basally with lateral projections in ventral view, laterally

compressed; dorso-lateral apophyses long, distinctly ex-

tending beyond apex of ventral process, flattened, and

strongly dilated, in the middle with conspicuous lamellate

processes (ventral view).

Comparative notes. This species is characterized by its

brownish coloration and particularly by the highly distinc-

tive morphology of the aedeagus, above all by the con-

spicuous processes of the dorso-lateral apophyses. Togeth-

er with the following two species, N. vexillatus forms a

group characterized by the coloration (body not black), the

coarse punctation of the pronotum and the elytra, a weak-

ly modified male sternite VII (without postero-median im-

pression, posterior margin weakly concave), the shape of

the male sternite VIII (approximately as long as broad;

posterior excision moderately deep), and particularly by

the morphology of the aedeagus (ventral process short and

of triangular shape in ventral view; dorso-lateral apophy-

ses modified: long, distinctly extending beyond the apex

of the ventral process, and dilated in various ways).

Distribution and natural history. Nazeris vexillatus is

currently known only from the region to the northwest of

Liuku in the Gaoligong Shan (Fig. 90). The specimens

were sifted from leaf litter and moss in degraded primary

forests and in shrub habitats with bamboo at altitudes of

2730-3150 m. One male collected in the beginning of Sep-

tember is teneral.

Naczeris circumclusus sp. n. (Figs 90, 183-188)

Type material. Holotype @ [teneral]: “CHINA: Yunnan,

Baoshan Pref., Gaoligong Shan, 32 km SE Tengchong,

Bonn zoological Bulletin 62 (2): 125-170

150-2250 m, 24°51-53’°N, 98°45°E, devast. prim. and

econd. forest, litter, dead wood, mushrooms sifted,

26. VIII.2009, leg. M. Schtilke [CH09-08/09] / Holotypus

& Nazeris circumclusus sp. n. det. V. Assing 2013” (cAss).

Etymology. The specific epithet (Latin, adjective: en-

framed, surrounded) alludes to the ventral process of the

aedeagus, whose ventral portion is enframed by the pos-

terior portion in ventral view.

Description. Body length 5.8 mm; length of forebody 3.3

mm. Habitus as in Fig. 183. Coloration (note that the holo-

type is teneral): body brown, with the apex of the abdomen

paler; legs yellowish; antennae yellowish, with anten-

nomere I slightly darker.

Head (Fig. 184) 1.05 times as long as broad; median

dorsal portion weakly elevated; punctation rather shallow,

dense, and distinctly umbilicate; interstices without mi-

crosculpture; eyes moderately small, approximately one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna 2.0 mm long.

Pronotum (Fig. 184) 1.17 times as long as broad and

0.85 times as broad as head; punctation distinctly coars-

er and somewhat irregularly spaced in postero-lateral por-

tions, non-umbilicate; interstices glossy; midline punctate

in anterior half, impunctate and narrowly elevated in pos-

terior half.

Elytra (Fig. 184) nearly 0.6 times as long as pronotum;

humeral angles obsolete; punctation dense, defined, and

approximately as coarse as that of pronotum; interstices

glossy. Hind wings completely reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense, defined, and moderately coarse on an-

terior tergites, distinctly sparser and finer on tergite VII

than on tergite VI; interstices without microsculpture; pos-

terior margin of tergite VII without palisade fringe; pos-

terior margin of tergite VII produced in the middle, ob-

tusely angled.

*: sternite VII (Fig. 185) with weakly concave poste-

rior margin; sternite VII (Fig. 186) approximately as long

as broad, posterior excision rather deep and narrowly U-

shaped, approximately 0.3 times as deep as length of ster-

nite; aedeagus (Figs 187-188) 0.93 mm long; ventral

process short, apically acute, and ventral portion conspic-

uously enframed by the ovoid posterior portion in ventral

view; dorso-lateral apophyses long, at basal third with di-

latation, and distinctly extending beyond apex of ventral

process.

Comparative notes. Based on the external (coarse punc-

tation, body not black) and the male sexual characters

(weakly modified sternites VII and VHI; sternite VII with-

out median impression; aedeagus with short ventral

process and with long, partly dilated dorso-lateral apophy-

ses), N. circumclusus 1s closely allied to N. vexillatus, from

OZFMK

166 Volker Assing

which it differs by the relatively larger and less convex

head, the punctation of the head (less coarse, denser, and

distinctly umbilicate), the longer and more slender anten-

nae, the U-shaped posterior excision of the male sternite

VIL, and by the morphology of the aedeagus (shapes of

ventral process and of the dorso-lateral apophyses).

Distribution and natural history. The type locality is sit-

uated in the Gaoligong Shan, to the southeast of Teng-

chong (Fig. 90). The teneral holotype was sifted from leaf

litter in a degraded mixed primary and secondary forest

at an altitude of 2150-2250 m, together with N. curvus.

Naczeris hastatus sp. n. (Figs 90, 189-195)

Type material. Holotype @: “CHINA (Yunnan) Nujiang

Lisu Aut. Pref., Gaoligong Shan, side valley 18 km NW

Liuku, 2590 m, 25°58°10”N, 98°42°27°E (devast. prim.

forest, litter sifted) 9-10.VI.2007 D.W. Wrase [29] / Holo-

typus ¢ Nazeris hastatus sp. n. det. V. Assing 2013”

(cAss).

Etymology. The specific epithet (Latin, adjective: armed

with a spear) refers to the spear-shaped dorso-lateral

apophyses of the aedeagus.

Description. Body length 6.2 mm; length of forebody 3.4

mm. Habitus as in Fig. 189. Coloration: body dark-brown,

with the elytra paler brown; legs yellowish; antennae yel-

lowish, with antennomere [| slightly darker.

Head (Fig. 190) 1.05 times as long as broad; median

dorsal portion elevated; punctation moderately coarse,

dense, and weakly umbilicate; interstices without mi-

crosculpture; eyes of moderate size, approximately one

third as long as the distance from posterior margin of eye

to posterior constriction of head. Antenna 2.0 mm long.

Pronotum (Fig. 190) 1.2 times as long.as broad and 0.85

times as broad as head; punctation distinctly coarser and

somewhat irregularly spaced in postero-lateral portions,

non-umbilicate; interstices glossy; midline punctate in an-

terior half, impunctate and narrowly elevated in posteri-

or half.

Elytra (Fig. 190) 0.57 times as long as pronotum;

humeral angles obsolete; punctation dense, defined, and

approximately as coarse as that of pronotum; interstices

glossy. Hind wings completely reduced.

Abdomen 1.25 times as broad as elytra; punctation

dense, coarse on tergites HI—V, somewhat finer on tergite

VI, only slightly finer and sparser on tergite VII than on

tergite VI; interstices without microsculpture; posterior

margin of tergite VII without palisade fringe; posterior

margin of tergite VHI convex.

*: sternite VII (Fig. 191) with weakly concave poste-

rior margin; sternite VII (Fig. 192) approximately as long

Bonn zoological Bulletin 62 (2): 125-170

as broad, posterior excision moderately deep and nearly

U-shaped, 0.24 times as deep as length of sternite; aedea-

gus (Figs 193-195) 1.1 mm long; ventral process short,

of triangular shape, and apically acute; dorso-lateral

apophyses long and spear-shaped, strongly triangularly di-

lated in the middle, and distinctly extending beyond apex

of ventral process.

Comparative notes. As can be inferred from the exter-

nal (coarse punctation, body not black) and the male sex-

ual characters (weakly modified sternites VII and VIII;

sternite VIII without median impression; aedeagus with

short ventral process and with long, partly dilated dorso-

lateral apophyses), N. hastatus is closely related to N. vex-

illatus and N. circumclusus. It is distinguished from these

species by the different punctation of the head (coarser

than in N. circumclusus, less coarse and denser than in N.

vexillatus), by the shape of the posterior excision of the

male sternite VII, and particularly by the morphology of

the aedeagus, above all the conspicuous shape of the dor-

so-lateral apophyses.

Distribution and natural history. The type locality is sit-

uated in the Gaoligong Shan, to the northwest of Liuku

(Fig. 90). The holotype was sifted from leaf litter in a de-

graded primary forest at an altitude of 2590 m.

Nazeris bangmaicus sp. n. (Figs 90, 196—200)

Type material. Holotype ¢ [teneral]: “CHINA: Yunnan,

Lincang Pref., Bangma Shan, 20 km NW Lincang, 2210

m, 23°58’25”’N, 99°54’36”E, water reservoir, devast. for-

est with ferns, litter & ferns sifted, reservoir bank,

9.1X.2009, leg. M. Schiilke [CH09-37] / Holotypus 3

Nazeris bangmaicus sp. n. det. V. Assing 2013” (cAss).

Paratype: 19: same data as holotype (cSch).

Etymology. The specific epithet is an adjective derived

from the name of the mountain where this species was dis-

covered.

Description. Body length 5.6—5.7 mm; length of forebody

3.1—3.3 mm. Habitus as in Fig. 196. Coloration: forebody

blackish-brown; abdomen black; legs yellowish; antennae

yellowish, with antennomere I slightly darker.

Head (Fig. 197) indistinctly transverse, 1.01—1.03

times as broad as long, of subcircular outline in dorsal

view; median dorsal portion very weakly elevated; punc-

tation moderately coarse, dense, and umbilicate; interstices

without microsculpture, forming very narrow ridges; eyes

relatively small, weakly projecting from lateral contours

of head, distinctly less than one third as long as the dis-

tance from posterior margin of eye to posterior constric-

tion of head. Antenna approximately 1.8 mm long.

©ZFMK

On the Nazeris fauna of China II 167

199

205 | 206

198 ~eeeree te 203 | ~ 204

Figs 196-206. Nazeris bangmaicus (196-200) and N. fissus (201-206). 196, 201: habitus; 197, 202: forebody; 198, 203: male

sternite VII; 199-200, 205-206: aedeagus in lateral and in ventral view; 204: male sternite VIII. Scale bars: 196-197, 201-202:

1.0 mm; 198-200, 203-206: 0.5 mm.

Bonn zoological Bulletin 62 (2): 125-170 OZFMK

168 Volker Assing

Pronotum (Fig. 197) approximately 1.15 times as long

as broad and 0.85 times as broad as head; punctation very

dense, much coarser than that of head, non-umbilicate; in-

terstices glossy, forming narrow ridges; midline punctate

in anterior half, impunctate and narrowly elevated in pos-

terior half.

Elytra (Fig. 197) very short, approximately 0.5 times as

long as pronotum; humeral angles obsolete; punctation

dense, defined, and approximately as coarse as that of

pronotum; interstices glossy. Hind wings completely re-

duced.

Abdomen 1.25—1.30 times as broad as elytra; puncta-

tion dense, defined, and coarse on tergites III-VI, some-

what less dense and less coarse on tergite VH, finer and

sparser on tergite VII; interstices without microsculpture;

posterior margin of tergite VII without palisade fringe;

posterior margin of tergite VHI convex.

&: sternite VII (Fig. 198) with small postero-median de-

pression, this depression with denser setae directed diag-

onally postero-mediad, posterior margin weakly concave

in the middle; sternite VIII transverse, posterior excision

moderately deep and V-shaped; aedeagus (Figs 199-200)

probably approximately 1.05 mm long (basal portion of

aedeagus of the teneral holotype somewhat deformed);

ventral process short and broad, with ventral portion of

triangular and dorsal portion of semi-circular outline in

ventral view; dorso-lateral apophyses short, strongly di-

lated in apical three fourths (ventral view), and somewhat

extending beyond apex of ventral process.

Comparative notes. This species 1s characterized partic-

ularly by the conspicuously short elytra and by the dis-

tinctive shapes of the ventral process and of the dorso-lat-

eral apophyses of the aedeagus. Closer affiliations to oth-

er species known from Yunnan are not evident.

Distribution and natural history. The type locality is sit-

uated in the Bangma Shan to the northwest of Lincang,

Yunnan (Fig. 90). The specimens were sifted from leaf lit-

ter in a degraded forest at an altitude of 2210 m. The holo-

type is distinctly teneral.

Naczeris fissus sp. n. (Figs 90, 201-206)

Type material. Holotype ¢@: “CHINA (Yunnan) Pu’er

Pref., Ailao Shan, 37 km NW Jingdong, 24°45°12”N,

100°41°24.5”E, 2300 m (devastated forest remnant, lit-

ter, moss, grass roots sifted), 13.[X.2009 D.W. Wrase [48]

/ Holotypus ¢ Nazeris fissus sp. n. det. V. Assing 2013”

(cAss). Paratypes: 72, 8 [partly teneral]: “CHINA: Yun-

nan, Pu’er Pref., Ailao Shan, 37 km NW _ Jingdong,

24°45°12”N, 100°41°24.5”E, 2300 m, devastated forest

remnant, litter & dead wood sifted, 13.1X.2009, leg. M.

Schiilke [CH09-48]” (ZFMK, cSch, cAss); 14: “CHINA:

Bonn zoological Bulletin 62 (2): 125-170

Yunnan, Lincang/Dali Pref., Wuliang Shan, old pass road,

N pass, 24°45°16.4”N, 100°29°50.3”E, 2350 m, forest lit-

ter & tea plantation, litter, mushrooms, grass sifted,

16.1X.2009, leg. M. Schiilke [CH09-55]” (cAss).

Etymology. The specific epithet (Latin, adjective: split,

divided) alludes to the conspicuously bifid ventral process

of the aedeagus.

Description. Small and slender species; body length

4.5—5.5 mm; length of forebody 2.5—2.8 mm. Habitus as

in Fig. 201. Coloration: forebody dark-reddish to dark-

brown, with the pronotum usually slightly darker; ab-

domen blackish-brown; legs yellowish; antennae yellow-

ish, with antennomere I slightly darker.

Head (Fig. 202) 1.05—1.08 times as long as broad; me-

dian dorsal portion indistinctly elevated; punctation

coarse, dense, and umbilicate; interstices without mi-

crosculpture, forming narrow ridges; eyes moderately

small, distinctly less than one third as long as the distance

from posterior margin of eye to posterior constriction of

head. Antenna approximately 1.5 mm long.

Pronotum (Fig. 202) 1.11—1.15 times as long as broad

and 0.93—1.00 times as broad as head; punctures rather

dense and non-umbilicate, nearly of similar diameter as

those of head, but much deeper; interstices glossy; mid-

line punctate in anterior half, impunctate and narrowly el-

evated in posterior half.

Elytra (Fig. 202) approximately 0.6 times as long as

pronotum; humeral angles obsolete; punctation dense,

somewhat less coarse than that of pronotum; interstices

glossy. Hind wings completely reduced.

Abdomen approximately 1.2 times as broad as elytra;

punctation dense and coarse on tergite III, gradually be-

coming less dense and less coarse towards tergite [V—VI,

fine and sparse on tergites VII and VHI; interstices with-

out microsculpture and glossy; posterior margin of tergite

VII without palisade fringe; posterior margin of tergite

VIII strongly convex.

3: sternite VII (Fig. 203) unmodified, posterior margin

truncate; sternite VIII (Fig. 204) as long as broad, poste-

rior excision small and V-shaped, approximately 0.13

times as deep as the length of sternite VII; aedeagus (Figs

205-206) 0.85—0.87 mm long and of highly distinctive

morphology; ventral process completely divided into two

lamellae; dorso-lateral apophyses almost straight and slen-

der, at basal third with a distinct process directed medi-

ad, not reaching apex of ventral process.

Comparative notes. Nazeris fissus is readily distinguished

from other congeners known from Yunnan by the com-

pletely divided ventral process of the aedeagus, the shape

of the dorso-lateral apophyses, from most species also by

small body size in combination with brownish coloration

of the forebody. Based on the synapomorphically derived

OZFMK

On the Nazeris fauna of China II 169

morphology of the aedeagus (ventral process completely

divided), this species is undoubtedly most closely relat-

ed to N. caoi Hu et al., 2011 from the Nabanhe Nature Re-

serve.

Distribution and natural history. The species was

recorded from two localities, the type locality in the Ailao

Shan and one locality in the Wuliang Shan, Yunnan (Fig.

90). The specimens were sifted from litter in two forest

habitats at altitudes of 2300 and 2350 m, on both occa-

sions together with N. sagittifer. Some of the type speci-

mens are teneral.

Unnamed and presumably undescribed species

Four species were represented in the examined material

only by females:

Nazeris sp. 4: 192: “CHINA: Yunnan, Baoshan Pref.,

Gaoligong Shan, 78 km N Tengchong, 2000 m,

25°44°49”N, 98°33°29”E, cleft with creek and forest rem-

nant, litter & dead wood sifted, 1.1X.2009, leg. M. Schiil-

ke [CH09-21]” (cSch).

Based on the external characters, this species 1s proba-

bly closely related to N. curvus and allied species of the

N. cangicus group.

Nazeris sp. 5: 12: “CHINA: Yunnan prov., 1.3—2.0 km S$

of Haba, 17—20.VI.2007, Haba Xueshan Mts., 2830-3000

m, 27°22.1’N, 100°08.2’E, Hajek & Ruzicka leg.”

(cSch).

This species, too, probably belongs to the NV. cangicus

group.

Nazeris sp. 6: 29: “CHINA: Yunnan, Dali Bai Aut. Pref.,

Zhemo Shan, 7 km SW Xiaguan, 25°32-33’N, 100°10-

11°E, 2870-2970 m, scrub with bamboo, oaks & Rhodo-

dendr., litter sifted, 18.1X.2009, leg. M. Schiilke [CH09-

60]” (cSch).

Based on the external characters, this species is close-

ly related to N. daliensis and allied species. It is readily

distinguished from the syntopic N. zhemoicus by a larg-

er and more robust body, a less oblong head with a prac-

tically completely matt surface and less coarse punctation,

and the denser punctation of the abdomen.

Nazeris sp. 7: 32: “CHINA: Yunnan, Dali Bai Aut. Pref.,

Mao Jiao Shan, E pass, 58 km NE Dali, 25°56’41”N,

100°40°05”E, 2525 m, second. mixed forest, litter, moss

& mushrooms sifted, 4.1X.2009, leg. M. Schiilke [CH09-

26]” (cSch).

This species, too, is presumably closely related to N.

daliensis and allied species, as is suggested by the simi-

lar external characters.

Bonn zoological Bulletin 62 (2): 125-170

Acknowledgements. | am indebted to Michael Schtlke, Ales

Smetana, Andreas Pitz, and Wolfgang Schawaller for the loan

of specimens. In particular, | am grateful to Michael Schiilke for

the generous gift of numerous holotypes, as well as to Andreas

Piitz for his permission to retain the holotype of NV. puetzi. Jia-

Yao Hu (Shanghai) kindly informed me on unpublished syn-

onymies of two species described from Zhejiang, provided a

translation of the Chinese description of NV. canaliculatus, and

helped in clarifying the type locality of this species. The help-

ful comments of the two reviewers are greatly appreciated.

REFERENCES

Assing V (2003) A revision of Othius Stephens. [X. New taxa,

new synonyms, additional records, and a checklist of species

(Coleoptera, Staphylinidae, Staphylininae, Othiini), pp.

727-752. In: Cuccodoro, G., Leschen, R.A.B. (eds), System-

atics of Coleoptera: Papers celebrating the retirement of Ivan

Lobl. Memoirs on Entomology International Vol. 17, Associ-

ated Publishers, Gainesville, Florida

Assing V (2009) A revision of the Western Palaearctic species

of Nazeris Fauvel, 1873 (Coleoptera: Staphylinidae: Paederi-

nae). Deutsche Entomologische Zeitschrift 56 (1): 109-131

Assing V (2013a) On the Nazeris fauna of China I. The species

of the Qinling Shan, the Daba Shan, and adjacent mountain

ranges (Coleoptera: Staphylinidae: Paederinae). Bonn Zoolog-

ical Bulletin 62 (1): 1-29

ASSING V (2013b) On the Lathrobium fauna of China V. New

species and additional records from Yunnan (Coleoptera:

Staphylinidae: Paederinae). Contributions to Entomology,

Beitrage zur Entomologie 63 (1) (2013): 53-128

Assing V (2013c) Six new species and additional records of Lath-

robium from the Palaearctic region (Coleoptera: Staphylinidae:

Paederinae). Linzer Biologische Beitrage 45 (1): 247-266

Assing V (in press) New species and records of Lathrobium from

China and Nepal (Coleoptera: Staphylinidae: Paederinae). Lin-

zer biologische Beitrage 45 (2) (2013)

Hu J-Y, Li L-Z, Zhao M-J (2007) Four new species of the genus

Nazeris from Sichuan, China (Coleoptera, Staphylinidae). The

Japanese Journal of Systematic Entomology 13 (2): 349-357

Hu J-Y, Li L-Z, Zhao M-J (2011a) Twelve new species of the

genus Nazeris Fauvel from Zhejiang Province, China

(Coleoptera, Staphylinidae, Paederinae). Zootaxa 2797: 1-20

Hu J-Y, Li L-Z, Zhao M-J (2011b) Notes on the Nazeris fauna

of Yunnan Province, China (Coleoptera, Staphylinidae, Paed-

erinae). ZooKeys 84: 13-21

Ito T (1996) A new species of the genus Nazeris from China

(Coleoptera, Staphylinidae). Entomological Review of Japan

51 (1): 63-65

Koch C (1939) Uber neue und wenig bekannte palaarktische Pa-

ederinae (Col. Staph.). Entomologische Blatter 35: 156-172

Lattin G de (1967) Grundriss der Zoogeographie. Gustav Fischer

Verlag, Stuttgart: 602 pp.

Peng Z, Li L-Z, Zhao M-J (2013a) Eight new apterous Lathro-

bium species (Coleoptera, Staphylinidae) from Sichuan,

Southwest China. ZooKeys 303: 1-21

Peng Z, Li L-Z, Zhao M-J (2013b) Two new species and addi-

tional records of Lathrobium Gravenhorst (Coleoptera:

Staphylinidae: Paederinae) from Guangxi, South China.

Zootaxa 3694 (3): 213-220

Watanabe Y, Xiao N N (1993) A new species of the genus

Nazeris (Coleoptera, Staphylinidae) from Yunnan Province,

Southwest China. Elytra 21 (1) 129-133

©ZFMK

170 Volker Assing

Watanabe Y, Xiao N N (1997) Four new Nazeris (Coleoptera, § Zheng F-K (1992) Four new species of the genus Nazeris Fau-

Staphylinidae) from Yunnan Province, Southwest China. vel from China (Coleoptera: Staphylinidae, Paederinae)

Edaphologia 58: 1-12 [English translation of Chinese title]. Acta Entomologica Si-

Watanabe Y, Xiao N N (2000) Four new species of the genus nica 35: 87-91

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Mountains in Yunnan, Southwest China. Elytra 28 (2):

311-321

Bonn zoological Bulletin 62 (2): 125—170 ©ZFMK

Bonn zoological Bulletin 62 (2): 171-176

December 2013

Records of ‘Indian’ Baya Weaver

Ploceus philippinus philippinus (Linnaeus, 1766) and

Hooded Wheatear Oenanthe monacha (Temminck, 1825) from Afghanistan

(Aves: Passeriformes)

Darius Stiels'’, Kathrin Schidelko'

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

° author for correspondence, E-mail: d.stiels@zfmk.de.

Abstract. Here we present the first record of Baya Weaver Ploceus philippinus for the country of Afghanistan. A spec-

imen was collected on May 29, 1967 in the vicinity of Bari Kowt (Gs 454) in the province of Kunar (S544). In addi-

tion, a Hooded Wheatear Oenanthe monacha was collected on January 17, 1968 in the vicinity of Farah (4)!e) in south-

western Afghanistan. Hitherto the status of the species in Afghanistan is unclear. We describe both specimens and short-

ly discuss the records.

Keywords. Ploceidae, Muscicapidae, range extension

INTRODUCTION

Afghanistan is situated in a zoogeographic transition zone

where Palearctic faunal elements meet those typical for

the Oriental region. For a landlocked country, Afghanistan

is extraordinary species-rich with about 490 bird species

recorded so far (Habibi 2007; according to Lepage 2012:

491 species following taxonomy of IOC World Bird

Names 2012). After decades of political instability,

which hindered ornithological research, the birds of

Afghanistan have received more attention only very re-

cently. In particular the seminal work by Rasmussen & An-

derton (2012) gives a comprehensive overview of the avi-

fauna of the country. Otherwise, mainly single observa-

tions were published in the last years (e.g. Balmer & Mur-

doch 2010a, b, Harrison & Grieve 2012a, b, Kowatsch &

Probst 2006, Mostafawi & Ostrowski 2010, Ostrowski et

al. 2008a, b). However, in north-eastern Afghanistan a

likely breeding population of one of the least known bird

species on the globe, the Large-billed Reed-Warbler Acro-

cephalus orinus, was discovered only recently (Svensson

2008, Timmins et al. 2009, 2010), and breeding was con-

firmed right across the border in Tajikistan (Ayé et al.

2010).

The Zoological Research Museum Alexander Koenig

(ZFMK) harbours a significant collection of birds from

Afghanistan. Most of the specimens were collected dur-

ing zoological and botanical expeditions as well as long-

time stays from the mid-1960s until the beginning of the

1970s. The main collectors were G. and J. Niethammer,

A. and H. Brade, C. M. Naumann as well as E. J. Kull-

mann. In addition, at least parts of the collection of the

Received: 21.12.2012

Accepted: 26.09.2013

former Zoological Museum of Kabul are now held at the

ZFMK. Some important results of the ornithological re-

search at that time were contemporarily published (e.g.

Niethammer 1967, Niethammer & Niethammer 1967,

Niethammer 1973). However, the collection has never

been completely inventoried, and this was only carried out

within the last months. Herein, we present the first results

of this task, namely the discovery of two species new for

Afghanistan or at least with hitherto unproven status for

the country.

RESULTS

Specimen of Ploceus philippinus philippinus

The specimen (ZFMK 2012.937) was collected on May

29, 1967 in east Afghanistan in the province Kunar (S504)

in the vicinity of Bari Kowt (Gos 494) (coordinates:

35.2992° N, 71.5397° E) within two kilometres from the

Pakistan-Afghanistan border (original label: E-

Afghanistan, Prov. Kunar-ha, vic. Barikot, 1300 m, Zo-

ologisch. Museum Kabul/Afghanistan Nr. 942). There is

no collector given, but according to the handwriting the

bird was labelled by C. M. Naumann. However, (re)la-

belling could have happened belatedly as the species name

was added subsequently. Thus, the collector remains un-

known. The bird is a male of unknown age. It is perhaps

not in complete breeding plumage, as the dark area of the

head and throat 1s light brownish and yellow feather mar-

Corresponding editor: T. Tépfer

172 Darius Stiels & Kathrin Schidelko

ue sTuRysyy / [14 «ps180[00Z

@h& IN

wrasnh

yd 311920 fel

is 26 ; ZVvo C IN - uu0g ‘Stusoy -y wunasnyy

496VY “SOS

WOOSYV PLY vy “IN @

s yyd saa;

sae

(b) view of the specimen of Baya Weaver Ploceus philippinus (ZFMK 2012.937) collected near

Bari Kowt, Afghanistan, on May 29, 1967.

Bonn zoological Bulletin 62 (2): 171-176 ©ZFMK

173

Records of Baya Weaver and Hooded Wheatear

2 =

f Oenanthe

‘ 6 (law

W- AF gnats?

i Y ‘ tava

monacha o

Oenanthe

(pe Afghanistan ) Prow Farah =

~ Jt e- Tavel,

en of Hooded Wheatear Oenanthe monacha (ZFMK 2012.942) collected in

Fig. 2. Ventral (a) and dorsal (b) view of the specim

the vicinity of Farah, Afghanistan, on January 17, 1968.

©ZFMK

Bonn zoological Bulletin 62 (2): 171-176

174 Darius Stiels & Kathrin Schidelko

Table 1. Measurements of the specimen of Ploceus philippi-

nus philippinus. Methodology follows Eck et al. (2011).

Table 2. Measurements of the specimen of Oenanthe monacha.

Methodology follows Eck et al. (2011).

Body part

- Size [mm] Size [mm]

Wing (flattened) vy Wing (flattened) 109

Tail 49 Tail 75

Bill length (tip to distal nostril) 1 Be Bill length (tip to distal nostril) 119

Bill height (proximal nostril) 9.5

gins on the back are not outstanding prominent (Fig. 1).

The head cap is yellow, partially with some fine darker

shaft streaks. Mantle feathers are centrally brown with ob-

vious yellow margins. Tail and wings are dark brown with

lighter margins. On the tertials, margins of the outer vanes

are buffy to rusty while they are yellow-olive on the sec-

ondaries forming an unobtrusive wing panel. The throat

is light brown, the breast yellow and the belly 1s light yel-

lowish to whitish while the flanks are rather buffy. In ad-

dition, tarsi and toes are horn-coloured. The bill of the

specimen is largely black with small lighter areas at the

basis of the upper and lower mandible. Note that some

colour fading cannot be excluded for darker feathers and

bare parts. Nevertheless, identification as Ploceus philip-

pinus 1s straightforward given the limited number of pos-

sible confounding species in southern Asia. In contrast to

P. benghalensis and P. manyar, the yellow breast is strik-

ing, and this also allows identification as belonging to the

nominate subspecies. The occurrence of the rather simi-

lar subspecies P. p. travancoreensis would be highly un-

likely, as this taxon is limited to south-western India. The

three outer primaries on both wings are heavily abraded

and faded while the inner primaries are fresh. Neverthe-

less, measurements are within the known range of the

species (Table 1, see Rasmussen & Anderton 2012). For

unknown reasons, wing measurement given on the orig-

inal label exceeds our measurement by 9 mm.

Specimen of Oenanthe monacha

The specimen (ZFMK 2012.942) was collected on Janu-

ary 17, 1968 in south-western Afghanistan in the vicini-

ty of the city of Farah (coordinates: 32.3744° N, 62.1164°

E) in the province of the same name (original label: SW-

Afghanistan, Prov. Farah, vic. Farah, Zoologisch. Muse-

um Kabul/Afghanistan Nr. 1075). The specimen was al-

so once held in the collection of the former museum of

Kabul. As for the weaver, no collector is given on the la-

bel. However, specimens collected between Delaram and

Farah on January 16, 1968 and on January 22, 1968 be-

Bonn zoological Bulletin 62 (2): 171-176

Bill height (proximal nostril) 4.5

tween Djuwein and Farah were collected by Heinrich

Klockenhoff. Thus, he is almost certainly the collector of

the Oenanthe monacha specimen as well as of four addi-

tional specimens from the same day and location (species:

Lanius excubitor pallidirostris, Cettia cetti, Phoenicurus

erythronotus, Phoenicurus ochruros).

Species identification is easy. The pure size (Table 2)

excludes the smaller Oenanthe species, and the bird is

readily identifiable as a male. In contrast to Oenanthe al-

bonigra, the whitish head cap is striking (Fig. 2). Mantle

and wings are black, upper tail coverts, rump and lower

back are white. The black of chin and throat extends to

the upper breast. The belly and under tail coverts are white

with a buffy tinge. Also note that the specimen does not

show a complete terminal dark tail bar as would be the

case in Oenanthe picata capistrata. Whitish tips to breast

and throat feathers as well as on the mantle and on the

remiges indicate a relatively fresh plumage. As there is no

obvious moult limit in the greater coverts detectable, it is

most likely an adult (van Duivendijk 2010). Bare parts are

black. Oenanthe monacha is monotypic (Dickinson

2003).

DISCUSSION

We presented two ornithological records for Afghanistan.

Concerning the specimen of Ploceus philippinus, one can

. only speculate if the incongruence on the label is due to

transcription errors or even some kind of mislabelling, as

field labels do not exist. Unfortunately, there is no hint for

distinct collection activities at the finding locality, since

no other specimen from the same location or from the

same time can be found within our collection. Thus, the

collection history of the specimen cannot be retraced. Due

to its bright plumage and its ability to perform tricks, P/o-

ceus philippinus is frequently caught and sold as cage bird

in Pakistan (Roberts 1992). Furthermore, the bird market

of Kabul still reflects the Afghan tradition of keeping pet

birds, as captured wild birds from throughout Afghanistan

as well as from Pakistan and India are sold here (Ostrows-

©OZFMK

Records of Baya Weaver and Hooded Wheatear 175

ki 2007). A record of Ploceus philippinus on the bird mar-

ket could not be provided, though (Ostrowski 2007). The

abraded primaries could also be a hint to a captive origin

although, alternatively, they might have been heavily worn

out naturally. Therefore, it is impossible to completely rule

out the possibility that the record represents an escaped

bird. On the other hand, date and location of the present-

ed record make perfectly sense. Ploceus philippinus is the

most widespread species of Ploceidae in southern Asia.

It inhabits a wide range of habitats including grasslands

and cultivated areas mostly close to water, although it is

less bound to swamps than other weavers in the region

(Craig 2010). In Pakistan, the species can be “locally abun-

dant” in the Indus basin (Grimmett et al. 2008). Bari Kowt

is also located within the Indus basin, as the town is sit-

uated at the shore of the Kuna River, a tributary of the

Kabul River, which eventually feeds the Indus. Collect-

ed at an altitude of 1300 m a.s.l., the bird occurred near

the upper limit of its altitudinal distribution which reach-

es 1400 ma.s.l. at the Himalayan foothills (Craig 2010).

According to the map in Grimmet et al. (2008, p. 224),

the next known natural occurrence in Pakistan might be

about 100 km away. Thus, habitat as well as the location

of the discovery fit well in the general ecology and the

biogeographical pattern of the distribution of the species.

Ploceus philippinus is generally assumed to be sedentary

(Rasmussen & Anderton 2012), but at least in Pakistan it

widely disperses in non-breeding seasons (Grimmet et al.

2008). We do not know whether the record presented here

only indicates a vagrant or whether the specimen even rep-

resents a breeding population. At least the collection date

fits perfectly within the regional breeding season of the

species (Roberts 1992). We recommend adding Ploceus

philippinus at least tentatively to the Afghan avifauna. To

our knowledge there is no other record of the species from

the country. It is not mentioned by Paludan (1959) or Ayé

et al. (2012), and the map in Rasmussen & Anderton

(2012) does not indicate any occurrences.

Oenanthe monacha is patchily distributed from Egypt

throughout the Middle East to southern Pakistan (Collar

2010). In most places, it seems to be only a sparse breed-

ing resident in remote habitats like desert ravines and

wadis up to 1300 maz.s.|. although it might also occur in

the vicinity of buildings in the desert (Collar 2010, Porter

& Aspinall 2010). Habitats are often too barren and arid

for other Oenanthe species (Collar 2010). In addition, the

species 1s described as shy and unobtrusive (Svensson

2009). Nearest depicted occurrence is eastern Iran where

the Lut desert (Dasht-e-Lut) reaches close to the Afghan

border (see map in Porter & Aspinall 2010, p. 314). How-

ever, there is at least one unproven sight report from Pul-

1 Chakri which even suggests breeding (Kullberg 2002)

and this observation was cited by Habibi (2007). Never-

theless, neither Paludan (1959) nor Ayé et al. (2012) men-

tion Oenanthe monacha, and the map in Rasmussen & An-

Bonn zoological Bulletin 62 (2): 171-176

derton (2012) does not denote any findings from

Afghanistan. Thus, the status of the species in the coun-

try was hitherto categorised as unconfirmed (UNEP 2008).

In conclusion, our findings add to the still fragmentary

knowledge of the Afghan avifauna. The presented records

of a Palaearctic and an Oriental faunal element further

highlight the transitional character of the fauna of

Afghanistan.

Acknowledgements. Renate van den Elzen kindly identified the

handwriting on the label as belonging to C. M. Naumann and

gave valuable comments to an earlier draft of the text. We thank

Sonke Twietmeyer who gave technical support. The manuscript

benefited from constructive improvement suggestions by Till

Topfer and one anonymous reviewer.

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Bonn zoological Bulletin 62 (2): 177-185

December 2013

Small mammal community composition

in the Volcanoes National Park, Rwanda

Deogratias Tuyisingize'*, Julian C. Kerbis Peterhans’, Gary N. Bronner’ & Tara S. Stoinski'

' The Dian Fossey Gorilla Fund International / Karisoke Research Center; BP 105, Musanze, Rwanda;

E-mail: deotuyisingize@yahoo.com.

? Roosevelt University, College of Professional Studies, 430 S Michigan Ave., Chicago, IL 60605 & Science and Education,

Field Museum of Natural History, 1400 S Lake Shore Dr., Chicago, IL 60605, U.S.A.

‘University of Cape Town, Rondebosch 7701, Cape Town, South Africa.

Abstract. Terrestrial small mammal community composition was examined in the Volcanoes National Park (VNP), Rwan-

da with respect to variation in habitat type and altitude. Trapping resulted in the capture of 220 individuals including

eight species of rodents, three species of shrews and one mongoose. Of the species captured, Praomys degraaffi 1s vul-

nerable and Sy/visorex vulcanorum is near threatened (IUCN 2012). Six species (Hylomyscus vulcanorum, Mus bufo,

Praomys degraaffi, Sylvisorex vulcanorum, Lophuromys woosnami and Tachyoryctes ruandae) are endemic to the Alber-

tine Rift. Species richness and diversity varied significantly among the different habitat types as they increased with el-

evation up to the middle altitudes (2860—3255m) and then declined with increasing elevation. Altitude accounted for

20 % of the variation in species diversity. Species variation in habitats and altitude was related to environmental factors.

Endemic species were found mainly in low and middle attitude habitats; thus these habitat types are important for con-

servation of small mammals at Volcanoes National Park.

Key words. Rodentia, Soricidae, Volcanoes National Park, endemism, diversity, elevational gradient, community struc-

ture.

INTRODUCTION

Baseline data are essential for biodiversity monitoring, es-

pecially during this era of anthropogenic and climatic

change. However, the lack of accurate data on the status

of free-ranging wildlife populations may limit the effica-

cy of monitoring programs. The African montane primary

forests of the Albertine Rift are home to many endemic

mammal species (Hutterer et al. 1987; Kerbis Peterhans

et al. 1998; Kaleme et al. 2007). The Volcanoes Nation-

al Park, gazetted as Parc National des Volcans in 1925

(hereafter VNP) is an afromontane forest rich in mammal

species and endemism and is continuous with adjacent vol-

canoes in the Democratic Republic of Congo and Ugan-

da. Many small mammals exhibit narrow habitat prefer-

ences and the distribution and abundance of small mam-

mals may be indicators of ecosystem health (Golley et al.

1975). Habitats with low vegetation diversity house rel-

atively few species (Kaleme et al. 2007), and altitudinal

variation also influences small mammal diversity (Kerbis

Peterhans et al. 1998).

The VNP is thought to support more than 25 small mam-

mal species (Gyldenstolpe 1928; Hutterer et al. 1987).

However, few studies on the ecology and community char-

acteristics of small mammals (rodents and shrews) have

been done in the VNP region (Gyldenstolpe 1928; Hut-

terer et al. 1987). While a considerable amount of work

Received: 26.10.2012

Accepted: 14.12.2012

has been carried out on the vegetation, birds and large

mammals of VNP, including the critically endangered Go-

rilla beringei beringei Matschie, 1914 (Plumptre 1991;

Fischer & Hinkel 1992; Robbins et al. 2001; Owiunji et

al. 2005), small mammals remain poorly documented. The

only scientific publication on small mammals in the VNP

was written over 80 years ago (Gyldenstolpe 1928) when

20 species were noted (data included in Table 1), but de-

tailed small mammal diversity and distribution patterns

were not determined. Since then, there has only been one

unpublished study by Kajonjoli in 1993, rendered incom-

plete by 1990’s Rwandan civil war and the unpublished

report (data included in Table 1) on the Ugandan slope of

Mgahinga Gorilla National Park (MGNP) by Kerbis Pe-

terhans & Austin (1996).

Information on small mammal distribution and abun-

dance across diverse habitats and along an elevational gra-

dient in VNP will be useful to supplement additional sci-

entific data for understanding structure and variation of

small mammal communities. In this paper, we document

small mammal community characteristics at VNP based

on our recent collecting program, and specifically (1) com-

pare small mammal abundance, diversity and distribution

in eight habitats (Bamboo, Hagenia Woodland, Brush

Ridge, Mixed Forest, Herbaceous, Sub Alpine, Alpine

Corresponding editor: R. Hutterer

178 Deogratias Tuyisingize et al.

| Virunga National Park |

—

\ AG

Democratic Republic of Congo

\ Ugand i

i Mgahinga National Park |

. AAs

5y0m_

& = Trapping sites

Rwanda ---—- International border

v Yj; Alpine

Pure and Mixed Bamboo

==-=:1 Brush Ridge

Kees] Hagenia Forest

ee | Herbaceous

f ee vA Volcanoes National Park |

\ “| Meadow

“su-] Mimulopsis

Kilometers

Fig. 1.

zones, and Swamp); (2) assess levels of diversity in rela-

tion to altitude, environmental variables and habitat char-

acteristics. (3) Finally, we provide data from two previ-

ous studies that contribute to assessing changes in Virun-

ga small mammal diversity and abundance over the past.

MATERIAL AND METHODS

The study area

The Volcanoes National Park, Rwanda (between

1°21°—1°35’S, 29°22’—29°44’E) is a mountainous region,

ranging 1n altitude from 2300 m to 4500 m (Weber 1987).

It is located on the eastern edge of the Albertine Rift,

which forms part of the watershed between the Nile and

Congo River systems (Weber 1987). It lies in north-west-

ern Rwanda and borders its sister parks, the Virunga Na-

tional Park (ViNP) in the Democratic Republic of the Con-

go (DRC) and Mgahinga National Park (MGNP) in Ugan-

da. The Volcanoes National Park (VNP), ViNP and MGNP

Bonn zoological Bulletin 62 (2): 177-185

ESE] Mixed Forest

Sub-Alpine

A map of the Virunga massif illustrating the major vegetation zones and the locations of the trapping sites in the VNP.

(Fig. 1) form the Virunga volcanoes area (Virunga mas-

sif). This study was carried out at three sites in the Vol-

canoes National Park, Rwanda (Mts. Visoke or Bisoke,

Sabyinyo or Sabinio and Gahinga) from October 2 to No-

vember 8, 2009. The selection of study sites was based

on altitude and the major vegetation zones of VNP (Fig.

1). The major vegetation zones surveyed include: (1)

Mixed Forest, (2) Bamboo, (3) Hagenia Woodland, (4)

Herbaceous, (5) Brush Ridge, (6) Sub Alpine zone, and

(7) Alpine (Plumptre 1991); swamps were categorized as

an 8th zone.

For Table 1, we add data from two historical collections.

The first reflects the efforts of the Swedish Zoological Ex-

pedition to the ‘Birunga Volcanoes’ from February to

March, 1921 (Gyldenstolpe 1928). The second collection

includes the species records from a more recent report

(Kerbis Peterhans & Austin 1996) that gathered baseline

data on terrestrial small mammal communities surround-

ing Kabiranyuma Swamp on the saddle between Muhavu-

ra and Mgahinga (2980 m) as well as land reclaimed for

the park (2810 m) from evicted settlers, just on the Ugan-

OZFMK

Small mammals of the Volcanoes NP, Rwanda 179

Table 1. Small mammal species recorded in the Virunga massif then and now. Area codes: Burunga (B), Chahafi (Ch), Karisim-

bi (Ka), Kibati (Ki), Lulenga (L), Lake Mutanda (LM), Mgahinga (Mg), Muhavura (Mu), Mikeno (Mi), Ninagongo (Ni), Ngoma

(Ng), Sabinio(Sa), Tamohanga (Ta), Tsitsilonga (Ts), Visoke (Vi). References (Ref.): (1) Gyldenstolpe (1928); (2) Kerbis Peter-

hans & Austin (1996); (3) this study.

Species Elevations Mtn (area) Ref.

Crocidura niobe Thomas, 1906 2980 Mg-Mu 2

Crocidura olivieri kivu Osgood, 1910 up to 3300 B,Ki, Lu,Mi,Ta ]

2810 Mg 2

up to 3700 Vi 3

Crocidura tarella Dollman, 1915 2900 LM 1

Vi 3

Paracrocidura maxima Heim de Balsac, 1959 2810 Mg 2

Suncus megalura (Jentink, 1888) 2810, 2980 Mg, Mg-Mu 2

Sylvisorex lunaris ruandae Loénnberg & Gyldenstolpe, 1925 2600 Sa 1

2810, 2980 Mg, Mg-Mu 2

Sylvisorex vulcanorum Hutterer & Verheyen, 1985 2810, 2980 Mg, Mg-Mu 2

2980 Vi 3

Myosorex babaulti Heim de Balsac & Lamotte, 1956 2980 Mg-Mu 2

Heterohyrax helgei Lonnberg & Gyldenstolpe, 1925 3500-4000 Mi, Sa 1

Hystrix stegmanni Muller, 1910 Mu,Sa l

Aethosciurus ruwenzorii vulcanius Thomas, 1909 Mi- Ni 1

Funisciurus carruthersi birungensis Gyldenstolpe, 1927 Mi,Ki |

Tamsicus vulcanorum vulcanorum Thomas, 1918 Mi 1

Graphiurus murinus vulcanicus Lénnberg & Gyldenstolpe 1925 3900 Ki |

2810 Mg 2

2500-3000 Sa, Vi 3

Delanymys brooksi Hayman, 1962 2980 Meg-Mu 2

Dendromus c.f. insignis kivu Thomas, 1916 2980 Mu ]

Meg-Mu 2

Lophuromys aquilus laticeps Thomas & Wroughton, 1907 up to 4000 B,Ka, Ki,LC l

LM, L,Mi,

2810, 2980 Mu,Ng, Sa,Mg, 2

2400-3700 Mg-Mu 3

Vi, Sa

Lophuromys woosnami Thomas, 1906 2810, 2980 L, Mi, Ki, Mu, I

2400-3600 Sa,Mg, Mg-Mu 2

Vi, Sa 3

Arvicanthis abyssinicus rubescens Wroughton, 1909 LC |

Dasymys incomtus medius Thomas, 1906 up to 2600 B, L, Sa,Ta |

Dasymys c.f. rwandae 2810, 2980 Mg, Mg-Mu 2

Grammomys c.f. dolichurus 2810 Mg 2

Hylomyscus aeta weileri Lonnberg & Gyldenstolpe, 1925 2400 Mi l

Hylomyscus vulcanorum Lonnberg & Gyldenstolpe, 1925 3700-3800 Ka, Mi l

2810, 2980 Mg, Mg-Mu 2

2400-3400 Vi,Sa 3

Lemniscomys striatus cf. massaicus (Pagenstecher, 1885) L 1

Mus bufo bufo Thomas, 1906 2700 B, Sa l

2810 Mg 2

2540, 2850 Sa, Vi 3

Mus gratus gratus Thomas & Wroughton, 1910 Ni l

Mus triton birungensis Lonnberg & Gyldenstolpe, 1925 3400 Mi l

2980 Meg-Mu 2

Oenomys hypoxanthus (Pucheran, 1855) |

2810, 2980 Mg, Mg-Mu 2

2820 Vi 3

Praomys degraaffi Van der Straeten & Kerbis Peterhans, 1999 2810 Mg 2

3220 Vi 3

Praomys jacksoni montis Thomas & Wroughton, 1910 Sa,Ki,LM I

Thamnomys kempi Dollman, 1911 3900 Ki l

2810, 2980 Mg, Mg-Mu 2

Otomys denti kempi Dollman, 1915 2800-3400 B,Mu,Sa,Ki l

2810, 2980 Mg, Mg-Mu 2

Otomys tropicalis vulcanicus Lénnberg & Gyldenstolpe, 1925 2980 Sa 1

Mg-Mu 2

Tachyoryctes ruandae Lonnberg & Gyldenstolpe, 1925 L, LC,Mu l

2810 Mg 2

2700 Vi 3

Bonn zoological Bulletin 62 (2): 177-185

OZFMK

180 Deogratias Tuyisingize et al.

da side of the Virunga Volcanos (Mgahinga Gorilla Na-

tional Park). Authors of taxon names of small mammals

are included in Table | or may be found in Wilson & Reed-

er (2005).

Field methods

Both live and snap traps were used. The live traps were

large Sherman folding traps (3”x3”x10"), and small, non-

folding perforated traps (2”x 2 2”5 x 6- 2”). Both Live

and Sherman traps were laid out in lines, 10 m apart (Tews

et al. 2004), with five lines located randomly in each habi-

tat at least 100 m away from human paths (Sutherland

2008). Snap traps were placed at the same points as live

traps. Each trap was set for three nights. All traps were

baited with crushed ground-nuts, meat and fish, bananas,

and pieces of potatoes. They were checked daily between

0700h and 1000h and again between 1400h and 1600h.

A locally made trap was used to collect mole rats (Tachy-

oryctes ruandae) in private cultivated plots adjacent to the

park.

At each station, trapped animals were collected,

processed and/or released (Nicolas & Colyn 2006). Live

animals were placed in a zip-lock plastic bag and

weighed using either 100 g or 500 g Pesola spring bal-

ances. A field identification was provided, sex determined

and various measurements taken (head-body length, tail

length, ear length, and hind foot length) using a ruler grad-

ed in mm. Each newly captured animal to be released was

marked by fur-clipping to ensure it could be recognized

if subsequently recaptured.

Within each of the habitat types, the major plant species

were recorded from the grids as follows: herbaceous zones

used 2m’ plots; shrub areas used 5m’ plots; areas domi-

nated by trees used 10m’ plots. Plant species were iden-

tified from herbarium material at Karisoke Research Cen-

ter (KRC) and from the Flora of Rwanda (Troupin

1977—1988).Temperature and wind speed were collected

daily between 9:00—9:30 am, allowing comparison be-

tween different sites.

Data analysis

To standardize data for all habitats sampled so that species

richness and diversity could be compared, trap success for

each trapline (the number of animals caught per 100 trap

nights) was calculated. The number of species trapped (per

100 trap nights) was also calculated for each site using the

expression: [Species richness/Trap effort] x 100. The

Shannon-Wiener (H) index of diversity was calculated for

Bonn zoological Bulletin 62 (2): 177-185

all sites sampled and evenness values were derived from

it.

H= SJ-eritinri

with H = the Shannon diversity index; P; = fraction of the

entire population made up of species I; S = numbers of

species encountered; and )) = sum from species | to

species S. The Shannon index increases with the number

of species in a community, and in theory, the Shannon in-

dex cannot exceed 5.0 (Krebs 1989). A high value of Shan-

non-Wiener index may indicate a large number for the

species with similar abundances; a low value indicates

lower species richness or domination by a few species.

Evenness values (E)[E = H / In (S)] indicate how num-

bers of individuals are distributed among species in a com-

munity. When the evenness is high, the mammal fauna is

more diverse and the species are equally abundant (Magur-

ran 2005).

We tested non-parametric estimators available in Esti-

mateS software (available at http://viceroy.eeb.uconn.edu

/estimates/EstimateS) to estimate species richness as an

alternative to the observed number of species in model

Chaol. The classic richness estimators Chaol was com-

puted along with log-linear 95 % confidence intervals

(Chao 2005, Colwell & Coddington 1994).

In order to estimate the number of additional species

necessary to reach an asymptotic plateau, species accu-

mulation rates were examined and were compared among

each habitat type (Colwell et al. 2004). The calculation of

species accumulation rates and the estimation of asymp-

totes/plateaux (species richness) were derived from the

distribution of all individuals caught across habitats;

species richness was estimated as a function of number

of samples. If the curves became flatter with increasing

effort (Chao | confidence bounds met or connected), it be-

comes less likely to detect new species in the additional

samples.

Similarities of the small mammal communities in the

different habitats were assessed using Bray-Curtis simi-

larity indices and linked Cluster Analysis. Tukey’s Hon-

estly-Significant-Difference (Tukey HSD) tests were

used for post-hoc comparisons when significant among-

groups differences were indicated. To investigate the re-

lationship between small mammal species and environ-

mental factors, a Canonical Correspondence Analysis

(CCA) was used. This is an indirect gradient analysis tech-

nique used to study the distribution of species along eas-

ily measured, recognizable environmental variables

(Gauch & Whittaker 1972). The mole rat Tachyoryctes

ruandae was found in cultivated fields outside the park,

and it was not included in statistical analyses.

OZFMK

Small mammals of the Volcanoes NP, Rwanda 181

Table 2. Total numbers of individuals per species captured in the eight habitat types at VNP.

Species /Habitat/Altitude Mixed Swamp Bamboo Hagenia Herbaceous Brush Sub Alpine Alpine Total

(m) Forest Woodland Ridge

2380-2580 2540 2540-2660 2740-3020 2900-3183 3220-3400 3420-3600 3640-3710

Crocidura tarella l I

Crocidura olivieri 3 l 2 1 2 9

Galerella sanguinea | |

Graphiurus murinus 2 l 3 6

Hylomyscus vulcanorum 1 I 2 3: 7 2 3 19

Lophuromys aquilus 8 15 4 8 24 15 15 10 99

Lophuromys woosnami 10 7 1] 1] 13 21 >) 78

Mus bufo l ] 2

Oenomys hypoxanthus l l

Praomys degraaffi | l

Sylvisorex vulcanorum l ]

Tachyoryctes ruandae (2)

TOTALS 24 24 18 29 47 41 23 12 218

Trap nights 300 300 300 300 300 300 300 300 2400

Density (Trap success %) 8.0 8.0 6.0 9.7 15.7 13.7 Tel 4.0 9.1

# species 5 4 4 8 5 6 3 2 12

RESULTS

Species richness

Small mammal trapping resulted in the capture of 220 in-

dividuals (208 rodents, 11 shrews, and one mongoose).

Ninety-nine individuals were female while 121 were male.

A total of eight rodent species, three shrews and one mon-

goose species were recorded. All rodents caught belonged

to the subfamilies Murinae, Deomyinae, Rhizomyinae and

Graphiurinae (Wilson & Reeder 2005). The most common

species were Lophuromys aquilus (n=99) and Lophuromys

woosnami (n=78), combining for 80.54 % of the individ-

uals captured. These species were encountered in all habi-

tats; all other species were rare (<19.46 % of individuals

captured in all habitats).

Small mammal species diversity was highest in Hage-

nia Woodland (8), intermediate in Brush Ridge (6), Herba-

ceous and Mixed Forest (5), Swamp and Bamboo (4) and

lowest in Sub Alpine (3) and Alpine (2). Rodent captures

were always more numerous than shrew captures (Table

2).

For Sub Alpine and Alpine zone Chaol species richness

estimator upper bounds reached horizontal plateaux

(asymptotes), and its upper bound reached the lower bound

for the bamboo and mixed forest (Fig. 2), so sampling ef-

fort can be considered adequate to reflect actual species

richness. In other habitats, plateaux were not attained. Fur-

thermore, species accumulation rates for habitat types and

altitudinal range showed that the most species rich habi-

tats in VNP were mid elevation habitats: Hagenia Wood-

land followed by Brush Ridge and Herbaceous.

Bonn zoological Bulletin 62 (2): 177-185

Species diversity between habitats

A Bray-Curtis dendrogram based on all individuals cap-

tured showed that two species captured in Alpine repre-

sented distinct communities than those from other habi-

tats. The Alpine (highest elevation at over 3600 m) and

Bamboo habitats (lowest elevations of 2540—2660 m) had

the least similar mammal communities (26.66 %) and

species richness was low in both zones.

The effective number of species (Shannon diversity) was

highest (H>1.0) in Hagenia Woodlands and Brush Ridge,

intermediate (0.8>H<0.6) in Herbaceous, Mixed Forest,

Bamboo and Swamp and lowest in the Sub Alpine (H=0.2)

and Alpine habitats (H=0.1). These differences in diver-

sity indices were significant (Kruskal-Wallis test:

Hz 4y9=20.49; p=0.005). The variation in community

species (Evenness) also differed significantly among habi-

tats types (Kruskal-Wallis test: H7_49=20.49; p=0.005). The

variation in community species were generally low (<0.5),

indicating that communities in all habitats were dominat-

ed by two species (Lophuromys aquilus and Lophuromys

woosnami). Species diversity was negatively correlated

with altitude. Altitude accounted for 20 % of the varia-

tion in species diversity with increasing altitude (p=0.004).

Environmental correlates of community structures

The CCA-biplot (the first two axes) for environmental

variables, habitat and small mammal abundances ex-

©ZFMK

182 Deogratias Tuyisingize et al.

Mixed forest Swamp

5 10 15 20 5 10 i 20

Bamboo Hagenia

ae Ts an ae ee ee et ee ee Se

5 10 15 0 5 10 15 20 25 30

Herbaceous

Species richness

10 12 14

0) 10 20 -30 40

Subalpine

5 10 15 20

Number of individuals sampled

Fig. 2. Accumulation rates of observed and estimated (Chao1) richness. The black dots are observed number of species, the round

circle represented Chaol estimate, while dotted line is 95% confidence interval.

Bonn zoological Bulletin 62 (2): 177-185 OZFMK

Small mammals of the Volcanoes NP, Rwanda 183

plained 19.3+39.4 (equals to 58.7 %) of variance in the

data set (Fig. 3). The environmental variables that affect-

ed separation of sites and species along axis | most strong-

ly were temperature, altitude, canopy cover, herbaceous

cover and wind speed.

Lophuromys aquilus, Lophuromys woosnami and Hy-

lomyscus vulcanorum plotted near the intersection of the

axes, suggesting that these species are not strongly influ-

enced by any of the six environmental variables and that

they are habitat generalists. Furthermore, Crocidura

olivieri kivu also seemed to be a generalist, and occurred

in five habitat types. The percentage of canopy cover has

a greater influence on the distribution of Graphiurus mur-

inus vulcanicus, a primarily arboreal species.

DISCUSSION

Our study found few small mammal species at VNP com-

pared to previous studies. The contiguous forest at

Mgahinga National Park has 16 known rodent species and

seven shrew species (Kerbis Peterhans & Austin 1996),

eight of which were recorded during this study: Hy-

lomyscus vulcanorum; Lophuromys aquilus; Lophuromys

woosnami; Oenomys hypoxanthus; Sylvisorex vulcano-

rum, Crocidura olivieri; Graphiurus murinus and Mus bu-

fo. Differences in size of the protected areas, trapping

techniques, time of year, elevation, and habitats sampled

may account for differences in species diversity with this

adjacent site. The differences between sizes of sampled

areas and trapping methods might have influenced differ-

ences. For instance squirrels can be obtained using shot-

guns. Shrews are more easily captured with pitfalls.

Gyldenstolpe (1928) covered the whole Volcanoes Nation-

al Park, while this study has focussed on Mt Bisoke and

the foot of Mt Sabyinyo.

Our study confirms a pattern of variation in species rich-

ness of small mammal communities along an altitudinal

gradient. Habitat heterogeneity and altitudinal variations

are a major factor affecting small mammal diversity (Is-

abirye-Basuta & Kasenene 1987; Stanley & Hutterer

2007). Mid elevation peaks in species diversity are com-

mon in both rodents and shrews whereas increasing ele-

vation leads to fewer species. Lophuromys aquilus and Lo-

phuromys wosnami are adaptable to the VNP habitats be-

cause they require moist scrub, forest, grassy, and tropi-

cal moist montane areas (Kingdon 1984). The elevation-

al diversity might be related to climatic impacted factors

such as rainfall, temperature, productivity, competition, re-

source abundance, habitat complexity, or habitat diversi-

ty (Lomolino 2001).

Species accumulation curves show insufficient sampling

effort in six of eight studied habitats. Further, extreme

weather during our sampling periods may have reduced

trapping success. Some species previously collected in

VNP were not captured during this study including Cro-

cidura niobe, Suncus megalura, Sylvisorex lunaris ruan-

dae, Paracrocidura maxima, Myosorex babaulti and

eleven mice/rat species including Delanymys brooksi,

Dendromus insignis kivu, Dasymys incomptus, Dasymys

Hagenia

Ocoh Asvir

yen

td eee >

Crot MysbCanop jee

Axis 1(39.4%)

Prad Gals Brush Ridge

Lob 1.8+-

— 0.9--

= Altitude Tegpaceous ~«

o Bit ee ~ LU os

—

x 23 “15

Mixed forest

6.9

13--

Wind speed » 1.8}

Fig. 3.

CCA-biplot showing the influence of various environmental variables on small mammal distributions during this study

period. Gram: Graphiurus murinus; Crot; Crocidura tarella; Croc: Crocidura olivieri; Hylv: Hylomyscus vulcanorum; Lapq: Lo-

phuromys aquilus; Lopw: Lophuromys woosnami; Musb: Mus bufo; Oenh: Oenomys hypoxanthus; Prad: Praomys degraaffi; Sylv:

Sylvisorex vulcanorum; Gals: Galerella sanguinea.

Bonn zoological Bulletin 62 (2): 177-185

©ZFMK

184 Deogratias Tuyisingize et al.

c.f. rwandae, Grammomys c.f. dolichurus, Hylomyscus ae-

ta, Thamnomys kempi, Otomys denti kempi, Otomys trop-

icalis vulcanus, Mus triton, Mus gratus (Gyldenstolpe

1928; Hutterer et al. 1987; Kerbis Peterhans & Austin

1996). More trapping over different seasons and with the

added use of pitfall traps in more habitats are needed to

adequately document small mammal communities at VNP.

The study area contains relatively few endemic species

(six) compared with other Albertine Rift sites. This com-

pares with twenty-one endemic species in the Ruwenzori

Mountains of Uganda (Kerbis Peterhans et al. 1998), and

18 endemic species in Kahuzi-Biega forest (Kaleme et al.

2007). Five of the six endemic species and the one threat-

ened species were found at low and middle altitudes at

VNP.

Species from VNP have a high probability of survival

owing to international and governmental protection efforts

directed at the mountain gorilla (Gorilla beringei

beringei), an “umbrella species’ from a conservation per-

spective. Since gorillas and endemic rodents and shrews

depend on the same habitats, the conservation of the first

acts to ensure the conservation of the second. In order to

maximize the maintenance of biodiversity conservation in

general and small mammals in particular, conservation ef-

forts need to target both low elevation and mid elevation

zones at VNP.

Our results have implications for how small mammal

communities in VNP may alter with climate change. With

increasing temperature, formerly low-elevation small

mammal species may expand their ranges upwards while

those of high-elevation species may contract, leading to

changed community composition at mid- and high eleva-

tions (Moritz et al. 2008). More long-term monitoring 1s

desirable to incorporate these factors, and to examine in

detail how to mitigate these risks.

Our study indicates that the small mammal species have

changed much since the pioneer study of Gyldenstolpe

(1928). This may be a signal of concern for the conser-

vation of this forest, and stresses the importance of reg-

ular small mammal surveys with close attention to asso-

ciated habitats.

The problem of spatial scales is also important; 40 trap

lines located at three study areas may not have been suf-

ficient in a protected area of 150 km’ and 40 km of width.

Long-term studies are needed to better understand the ef-

fects of climatic events relative to those of local differ-

ences related to habitat structure. The findings of species

accumulation rates and comparatively low richness dur-

ing this study indicate that the small mammal species list

for VNP is still incomplete and further inventory work is

needed. We recommend that future, longer-term studies

are carried out to confirm our species list, and to exam-

ine demographic processes and the impact of climatic

changes on gorilla habitat.

Bonn zoological Bulletin 62 (2): 177-185

Acknowledgements. Funding for this research was provided by

the Tropical Biology Association and the Dian Fossey Gorilla

Fund International. Permission to conduct this research was giv-

en by the Rwanda Development Board. Many thanks to Dr. Rosie

Trevelyan for her efficient help in reading and commenting ear-

lier draft.

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OZFMK

Bonn zoological Bulletin 62 (2): 186-202

Bats (Chiroptera) from the Albertine Rift,

eastern Democratic Republic of Congo,

with the description of two new species of the Rhinolophus maclaudi group

Julian C. Kerbis Peterhans'’, Jakob Fahr’, Michael H. Huhndorf’, Prince Kaleme**, Andrew J. Plumptre’,

Ben D. Marks’ & Robert Kizungu’

' College of Professional Studies, Roosevelt University, 430 S. Michigan Ave., Chicago, IL, 60605, USA

° Science and Education, Field Museum of Natural History, Chicago, IL 60605-2496, USA, E-mail: jkerbis@fieldmuseum.org

* Division of Evolutionary Biology, Zoological Institute, TU Braunschweig, Mendelssohnstr. 4,

D-38106 Braunschweig, Germany

* Department of Botany & Zoology, Stellenbosch University, Private Bag XI, Matieland, South Africa

> Centre de Recherche des Sciences Naturelles, Lwiro, Democratic Republic of Congo

° Wildlife Conservation Society, Plot 802 Kiwafu Rd, Kansanga, PO Box 7487, Kampala, Uganda

Abstract. Horseshoe bats of the Rhinolophus maclaudi species group were recently revised by Fahr et al. (2002). Known

members of the group are located in the mountainous region of West Africa and the Albertine Rift, east of the Congo

River basin with a major gap (4300 km) between the two recognized sub-groups. Here we describe two additional species

within this species group from the Albertine Rift center of endemism in the eastern Democratic Republic of Congo. One

derives from the Misotschi-Kabogo highlands, a heretofore poorly documented region half-way down the western shore

of Lake Tanganyika. Additional bat records from this locality are also documented. The second new taxon was collect-

ed in Kahuzi-Biega National Park, a World Heritage Site adjacent to the shore of Lake Kivu.

Key words. Rhinolophidae, endemism, new species, Misotschi-Kabogo highlands, Kahuzi-Biega National Park, Alber-

tine Rift, Democratic Republic of Congo.

Résumé. Les espéces de chauve-souris appartenant au groupe de Rhinolophus maclaudi étaient recemment révises par

Fahr et al. (2002). Les membres connus du groupe sont localisés du cété opposée au basin du Congo avec un vide de

4300 km entre les deux sous-groupes. Nous décrivons deux espéces appartenant au méme groupe en provenance de I’est

de la République Démocratique du Congo dans le Rift Albertin. Lune dérive des monts Misotschi-Kabogo, un site trés

mal connu pour sa biodiversité, situé a mi-chemin dans le versant ouest du lac Tanganyika. Les autres chauve-souris de

la méme région sont aussi documentées. L’autre espéce était collectée au Parc National de Kahuzi-Biega, un site du pa-

trimoine mondial de 1 UNESCO bien étudié, adjacent dans le versant ouest du lac Kivu.

Mots clés. Rhinolophidae, endémisme, espéces nouvelles, Misotschi-Kabogo, Kahuzi-Biega NP, Rift Albertine, Répu-

December 2013

blique Démocratique du Congo.

INTRODUCTION

The Albertine Rift montane zone of Central Africa, extend-

ing from the Blue Mountains in the north (ca. 2°N,

30°30’E) to the southern end of Lake Tanganyika (ca. 9°S,

30°30°E), is recognized as one of Africa’s biological

hotspots due to high levels of species richness and en-

demism (Kiiper et al. 2004; Orme et al. 2005; Plumptre

et al. 2007). Although numerous surveys have been con-

ducted and published on the gorilla-inhabited Virunga-

Bwindi heartland, many isolated areas, especially Con-

golese landscapes peripheral to this heartland, have nev-

er been properly surveyed. Here we describe two new

species of the Rhinolophus maclaudi group of horseshoe

bats. The group is readily diagnosed by their large ears

and by the absence of a connecting process to the sella

(Fahr et al. 2002). Specimens of a new species were first

Received: 31.07.2012

Accepted: 21.10.2013

collected in hills overlooking Lake Tanganyika in the Mis-

otschi-Kabogo highlands (5°-6°S). Subsequently, an in-

dependent team collected a second new species of this

group in Kahuzi-Biega National Park, a World Heritage

Site known for its eastern lowland gorilla population (Go-

rilla beringei graueri Matschie). These two new species

bring the total number of species described within the Rhi-

nolophus maclaudi group to six, four of them largely con-

fined to montane habitats within the Albertine Rift.

MATERIALS AND METHODS

Bats, including three individuals representing a new

species, were collected between 14 and 17 February 2007,

Corresponding editor: R. Hutterer

Bats from the Albertine Rift 187

Fig. 1. Above — View of Misotschi-Kabogo foothills from Lake Tanganyika. Note mature forest next to degraded patch near vil-

lage. Below — Mosaic of forest and grassland as seen from summit of Misotschi-Kabogo escarpment (Photographs A. Plumptre).

Bonn zoological Bulletin 62 (2): 186-202 SZ7FMK

188 Julian C. Kerbis Peterhans et al.

during the course of a preliminary small mammal survey

in the Misotschi-Kabogo highlands overlooking the west-

ern shore of Lake Tanganyika, eastern Democratic Repub-

lic of Congo. A fourth individual was subsequently cap-

tured nearby, on the Kilicha River on 17 Feb 2007. All

bat specimens were collected with mist nets, courtesy of

the accompanying ornithologists. With a single exception,

the new material discussed herein was collected during the

course of the Misotschi-Kabogo expedition. The excep-

tion was a single horseshoe bat, also representing a new

species, netted a few months later on the slopes of Mt

Kahuzi during a survey of terrestrial small mammals and

birds of Kahuzi-Biega National Park.

Data collected for all specimens included the macro-

habitat, sex, and reproductive condition. The following

standard external measurements were taken in the field:

total length (Total), tail length (Tail), hind foot length (HF),

ear length (Ear), forearm length (FA), and body mass

(Mass). Head and body length (H&B) was determined by

subtracting Tail from Total. All metrics are given in Ta-

bles 1 and 2. Voucher specimens were prepared as study

skins and skeletons or were preserved in 10% formalin.

For the latter group, skulls were later extracted at the Field

Museum of Natural History (FMNH) in Chicago and

transferred to 70% ethanol. The preservation of the spec-

imen is recorded as ssk (skin, skull and skeleton), alc (al-

coholic carcass), asr (alcoholic carcass with skull re-

moved), or sko (skull and skeleton only).

External metrics of the bat specimens, excepting Rhi-

nolophus, were taken by JCKP, using Mitutoyo CD-

6°CSX calipers to the nearest .01 mm for craniodental

variables and to the nearest 0.1 mm for external variables.

JF took the measurements of the Rhinolophus as well as

comparative museum specimens with Mahr 16 ES digi-

tal calipers and the same precisions. Definition of meas-

urements (Table | & 2) is given in Fahr et al. (2002), with

the following additional variables: HS-W: width of

horseshoe; Trag: length of tragus; CrnC,,,: length of skull

from posterior-most point to front of canine alveolus; Cb-

sC,,,: length of skull from posterior-most point of

condyles to front of canine alveolus; C-C,,,: width across

alveoli of upper canines; M’-M°,,,: width across alveoli

of posterior upper molars; C-M’,,,: length of upper (max-

illary) toothrow from front of canine to back of posterior

molar at alveoli; M'Br: breadth of first upper molar; C-

PM®: length from front of upper canine to back of poste-

rior premolar; PM’-M': length from front of posterior up-

per premolar to back of posterior molar; ManA: length of

mandible from tip of angular process to anterior-most

point, excluding incisors; C-M/;,,,: length of lower

toothrow from front of canine to back of posterior molar

at alveoli. Measurements are given in millimeters, body

mass in grams.

Tissue samples were stored in the field in a saturated

NaCl / EDTA buffer. Upon returning from fieldwork, tis-

Bonn zoological Bulletin 62 (2): 186-202

sues were stored in an ultracold freezer at -70°C. In the

field, animals were handled in accordance with American

Society of Mammalogists guidelines (Sikes et al. 2011).

Specimens were compared with material from the fol-

lowing institutions: American Museum of Natural Histo-

ry, New York (AMNH), Natural History Museum, Lon-

don (BMNH), Institut royal des Sciences naturelles de Bel-

gique, Bruxelles (IRSNB), Los Angeles County Museum

of Natural History (LACM), Muséum d’ Histoire naturelle

Geneve (MHNG), Musée Royal de |’Afrique Centrale,

Tervuren (MRAC), Senckenberg Museum Frankfurt

(SMF), and Zoologisches Museum of the University of

Zurich (ZMUZ). The new collections are deposited at the

Field Museum of Natural History, Chicago. D.C.M. refers

to the field notes of David C. Moyer. Taxonomy follows

Simmons (2005) unless stated otherwise.

STUDY AREAS

The Misotschi-Kabogo highlands of the eastern Democrat-

ic Republic of Congo run approximately 100 km along the

escarpment above the western shore of Lake Tanganyika

at an estimated width of 10-20 km. Topographically these

highlands are separated from the Itombwe massif to the

north by the Kilombwe valley, which is ca. 10 km wide

at an altitude of 800-1100 m. The continuous forest cov-

er of the Misotschi-Kabogo highlands on the one hand and

the Itombwe forest on the other hand is separated by a

stretch roughly 100 km wide. To the south, the Misotschi-

Kabogo highlands are separated from the Marungu high-

lands, which lack noteworthy forest cover, by the Luku-

ga valley. With the exception of birds (Prigogine 1960),

these highlands have been virtually unexplored by biolo-

gists. Prigogine (1955) described a new warbler (current-

ly Apalis porphyrolaema kaboboensis) and collected a

squirrel that was later described as a distinct subspecies

(Protoxerus stangeri kabobo Verheyen, 1960). Prigogine

also collected the type specimen of a distinct subspecies

(Groves, 2005) of Angola colobus monkey, Colobus an-

golensis prigoginei (Verheyen 1959) from ‘Mt Kabobo’.

Plumptre et al. (2008) summarized preliminary results of

a biodiversity survey of the Misotschi-Kabogo highlands

and discussed the conservation significance of this region.

The escarpment is continuously forested from the

lakeshore at 770 m up to the highest elevation at ca. 2700

m (Fig. | above). The main forest block is approximate-

ly 1,000 km? in size. On the western side, the highlands

drop to a rather flat area at an altitude of ca. 1000 m, which

is characterized by a heterogeneous mosaic of savannas

and riverine forests of variable width (Fig. | below).

The camps within the Misotschi-Kabogo forest were ac-

cessed from two different fishing villages along the Lake

Tanganyika shoreline. The camp where all but one of the

bats were collected is located approximately 4 km south-

©OZFMK

Bats from the Albertine Rift 189

west of the village of Talama, north of the town of Kalemie

at 1950 m (4°59’29”’S, 29°04’49”E). After exiting the for-

est and climbing a steep grassy ridge, the forest was again

reached and the camp was set within a horseshoe bend of

the Mukungu River. One specimen of the new Rhinolo-

phus was collected at a separate locality referred to here

as the Kilicha River locality at 1880 m (5°06°19"S,

29°03’56”E). Dominant tree species included Syzygium

guineense (Willd.) DC., Garcinia volkensii Engl., and

Tabernaemontana johnstonii (Stapf) Pichon. The climate

of the two survey sites is characterized by a single wet sea-

son from October until April and a dry season from May

until September, with an annual average precipitation of

1500-1550 mm, mean annual minimum temperature of

11.8°C, and mean annual maximum temperature of 21.5°C

(Prigogine 1960, Hijymans et al. 2005).

The specimen from Kahuzi-Biega National Park was

collected on the western slopes of Mt Kahuzi at an ele-

vation of 2600 m (2°15’°09”’S, 28°40’9”E) in secondary

forest with dominant tree species of Nuxia floribunda

(Benth.), Agauria_ salicifolia (Hook.f. ex Oliv.),

Macaranga kilimandscharica (Pax) and Afrocarpus usam-

barensis (Pilg.). The canopy was semi-closed at a height

of 12 meters. The sub-canopy was semi-closed with dom-

inant species of Chassalia subochreata (De Wild.)

Robyns, Polygala ruwenzoriensis (Chodat), Sericostachys

scandens (Gilg & Loper.), Mimulopsis solmsii Schweinf.,

Mikania chevalieri (C.D. Adams) W.C. Holmes & Mc-

Daniel, Sinarundinaria alpina (K. Schum.) C.S. Chao &

Renvoize and Tricalysia sp. (A.Rich. ex DC). The climate

is characterized by a single wet season from September

until May and a dry season from June until August, with

an annual average precipitation of 1850 mm, mean annu-

al minimum temperature of 10.2°C, and mean annual max-

imum temperature of 18.6°C (Hijmans et al. 2005).

RESULTS

Myonycteris angolensis (Bocage) ssp.

FMNH 195081 (original number MHH 844), subadult fe-

male, collected on 16 February 2007 by B. D. Marks. Skin,

skull and skeleton. Collected 4 km southwest of the vil-

lage of Talama, N of Kalemie: 4°59’29’’S, 29°04’°49”E,

1950 m.

The single specimen slightly exceeds the dimensions of

Myonycteris angolensis ruwenzorii (Eisentraut, 1965) and,

despite being a subadult (basioccipital suture incomplete-

ly fused), falls within the size range of the largest sub-

species, M. a. goliath (Bergmans, 1997). The latter author

listed three females of M. a goliath from eastern Zimbab-

we with FA 82.1, 83.8 and 89.6 (vs. FMNH 195081 with

FA = 84.0) while Monadjem et al. (2010) gave a mean FA

length of 83.9 + 2.1 (range: 81.4-87.4) for six females

Bonn zoological Bulletin 62 (2): 186-202

from northern Mozambique. FA-ranges of 83 females of

M. a. ruwenzorii are shorter with 72.8—83.0 (Bergmans

1997). Crn of FMNH 195081 is 43.85 and agrees with

three females of M. a. goliath (45.8, >42.3, >42.2) while

corresponding measurements of M. a. ruwenzorii range

smaller (39.1—43.2, n=58; Bergmans 1997). In terms of

distribution, the present specimen is located within the

range of M. a. ruwenzorii, stretching from southern Su-

dan to southeastern D.R. Congo, while it is rather distant

from the currently known distribution of M. a. goliath (E

Zimbabwe & Mozambique). These data suggest that more

material is needed to assess the taxonomic relationships

of both taxa and their distributions; the elevation of M. go-

liath to species rank proposed by Cotterill (2001) seems

premature. The transfer of Lissonycteris angolensis to

genus Myonycteris follows Nesi et al. (2013).

Hipposideros cf. ruber (Noack)

FMNH 195085 (original number MHH 842), adult female,

collected on 16 February 2007 by M. H. Huhndorf. Teats

small (nulliparous). Skin, skull and skeleton. Collected 4

km southwest of the village of Talama, N of Kalemie:

4°59°29”S, 29°04°49”E, 1950 m.

The taxonomy of the Hipposideros caffer/ruber group

is in flux due to pronounced cryptic diversity revealed by

molecular genetics (Vallo et al. 2009). The latter study

demonstrated that morphometrics previously used to dis-

tinguish species and subspecies show very limited match

with molecular clades; hence we refrain from assigning

the single specimen to any of the available names within

this species complex. Our tentative identification as H. ru-

ber follows the traditional concept, as the specimen is

clearly larger than H. caffer (Sundevall); however, molec-

ular data would be required for an unambiguous identifi-

cation.

Hypsugo cf. eisentrauti (Hill)

FMNH 195086 (original number MHH 845), adult female,

collected on 17 February 2007 by C. Kahindo. Teats small

(nulliparous). Skin, skull and skeleton. Collected 4 km

southwest of the village of Talama, N of Kalemie:

4°59°29”S, 29°04°49”E, 1950 m.

Our specimen is of large size (FA: 37.8), has a small

upper premolar (PM’') visible in lateral view, a broad and

bicuspid inner upper incisor (I'), a bicuspid outer incisor

(’), and IP’ nearly reaches in height the lateral cusp of I’.

It is very similar in measurements to a specimen (SMF

79444) collected in the Nyungwe National Park, Rwan-

da, at 2500 m, referred to as Hypsugo eisentrauti by Vol-

leth & Heller (1994), Heller et al. (1995), and Volleth et

al. (2001). Although both of these specimens have simi-

lar external measurements compared to type specimens of

Hypsugo eisentrauti from Cameroon, they are distinctly

©ZFMK

190

Julian C. Kerbis Peterhans et al.

Table 1. External and craniodental measurements of Chiroptera other than Rhinolophus.

Myonycteris Hipposideros

Hypsugo aff. eisentrauti

Hypsugo eisentrauti

angolensis cf. ruber

Museum FMNH FMNH FMNH SMF ZFMK ZFMK BMNH BMNH

195081 195085 195086 79444 68.5 68.6 67.2129 # 84.1684 §

Country DR Congo DR Congo DR Congo Rwanda Cameroon

Sex e . fd 3 3 a 2 3

Mass 82 10 9.8 12*(full) 7.0 8.3

Total 136 93 93 84 88

H&B 126 59 58 48 48

Tail 10 34 35 36 40

Ear 25 16 12 12.2 11 13

Tragus 4.0 4.7 5.1 4.6

FA 84.0 52.3 37.8 35.8 33.9 36.3 34.2 35.5

3Meta 57.7 38.7 34.6 34.3 30.9 33.2 31.2

3Phal 41.7 16.4 12.5 13.2 10.7 12:5

3Pha2 51.3 17.2 11.2 10.1 10.8 10.3

3Pha3 5.9 6.2 5.1 5.6

4Meta 54.9 36.3 33.2 33.4 30.1 32.8

4Phal 31.7 11.9 11.1 12.1 9.9 11.0

4Pha2 33.1 9.0 9.6 92 D2. 9.0

S5Meta 54.4 31.8 32.6 33.0 28.9 31.7

5Phal 27.0 14.5 8.0 9.1 7.3 7.8

5Pha2 31.2 10.4 5.5 5.0 5.0 5.0

Tibia 13.8 14.2 12.1 13.4

HF 5.5 17.8 7.9 ‘Tal 6.9 7.0 6.8

jclanen 21.6* Oo" 8.6 7.3 8.1

Cm 43.85 18.63 14.76 14.88 13.68 14.16 14.0 14.0

CrnC 42.72 18.35 14.59

Cbs 41.86 16.42 14.10 14.39 13.16 13.63 13.3 1337

CbsC 40.52 16.03 13.76

Mast 16.47 9.93 8.60 8.45 7.77 7.9 8.1

Zyg 24.32 9.53 10.74 10.65 9.61 [9.54]

BcB 17.04 8.44 7.83 7.50 7.36 7.46 Kee 7.0

BcH 12.07 6.79 5.83 5.70 5.54 5.38

C-C 8.01 4.15 4.95 5.20 4.60 4.23 4.5 5.0

M?-M? 12.31 6.37 6.73 6.75 6.06 6.3% 6.3 6.3

C-M? 16.8 6.34 5.34 5.59 5.22 5.1% 5.0 5:3

PoC 8.77 3.11 4.48 4.10 4.27 4.25 4.2 3.9

ManC 32.7 11.32 10.87 11.10 10.33 10.73 9.9 10.9

C-M, 18.44 6.68 5.72 5.98 5.51 5.67 5.5 5.8

* Hill 1968, De Vree, 1972; § measurements courtesy of Dieter Kock. SMF 79444: Rwanda, near Cyangugu, Nyungwe NP,

2500 m, leg. K.-G. Heller & M. Volleth, F-N° 339, 28 Mar 1990, skin & skull; ZFMK 68.5 (holotype eisentrauti): Cameroon,

Rumpi Hills, Dikume-Balue, camp V, leg. M. Eisentraut, F-N° 498, 18 Feb 1967, skin & skull; ZFMK 68.6 (paratype eisen-

trauti): Cameroon, Mt Kupe, camp II, ca. 1100 m, leg. M. Eisentraut, F-N° 198, 30 Nov 1966, skin & skull; BMNH 67.2129

(paratype eisentrauti): Cameroon, Mt Cameroon, Buea, leg. M. Eisentraut, F-N° 643, 15 Mar 1967, skin & skull; BMNH

84.1684: Cameroon, Mt Cameroon, 750 m; leg. M. O. Fedden & H. L. Macleod, F-N° 61, 2 Jan 1984, alc. & skull.

larger in several craniodental measurements (e.g. Crn, Cbs,

Mast, Zyg, M’-M’; Table 1). Similar to true Hypsugo

eisentrauti, the pelage of our specimen is unicolored dark

brown above and slightly bicolored below. However, our

specimen is much paler ventrally with brown roots tipped

with pale brown. Based on available data, specimens

FMNH 195086 and SMF 79444 appear to represent an un-

described species (see also Van Cakenberghe & Happold

2013).

Bonn zoological Bulletin 62 (2): 186-202

Rhinolophus willardi sp. nov. Kerbis Peterhans & Fahr

Willard’s Horseshoe Bat

Holotype. FMNH 195182 (original number D.C.M.

1680). Adult male, all teeth in wear. Collected on 17 Feb-

ruary 2007 by A. J. Plumptre & E. A. Mulungu. Speci-

men preserved in alcohol with skull removed.

©ZFMK

Bats from the Albertine Rift 19]

Fig. 2.

Left — Type locality of Rhinolophus willardi. Note large amount of epiphytes on trees near Kilicha River. Right — Aeri-

al view of artisanal mining site within forest of Misotschi-Kabogo highlands (Photographs A. Plumptre).

Type locality. Misotschi-Kabogo highlands, north of

Kalemie, Kilicha River, above the western shore of Lake

Tanganyika, South Kivu Province, eastern Democratic Re-

public of Congo, 5°06°19"S, 29°03°56”E, 1880 m. The

holotype was captured in a clearing near a stream in a deep

valley (Fig. 2, left). The forest covering the surrounding

slopes was dense with tall trees (40-50 m) covering the

surrounding slopes and had a fairly open understorey. The

clearing was formed in an area where the stream had de-

posited gravel in the wet season but was at a much low-

er level at the time of capture (dry season).

Paratypes. All from 4 km SW of the village Talama,

‘Camp 2’, 4°59°29”S, 29°04’49”E, 1950 m, and all pre-

served as skin, skull and post-cranial skeletons. FMNH

195082, adult male, original number MHH 837, collect-

ed by B. D. Marks on 14 Feb 2007, with convoluted epi-

didymes and abdominal testes (3x2 mm). FMNH 195083,

adult female, original number MHH 838, collected by B.

D. M. on 14 Feb 2007, with enlarged nipples but nulli-

parous. FMNH 195084, adult female, original number PK

754, collected by B. D. M. on 16 Feb 2007 without em-

bryos.

Etymology. The specific epithet honors Dr. David Willard

(Collection Manager, Division of Birds, FMNH) in recog-

nition of his unparalleled 35+ years of service to the Field

Museum of Natural History. While devoting 80+ hours a

week to the job and generously sharing his time and ex-

pertise as an ornithologist and educator, Dave also brings

an unmatched ethical standard. Although he did not col-

lect these particular bats, Dave has contributed to the doc-

umentation of Chiropteran diversity throughout his career

Bonn zoological Bulletin 62 (2): 186-202

and, as a consequence, has documented the distribution

of more mammal species in the Neotropics, Asia and

Africa than the vast majority of field mammalogists.

Diagnosis. Immediately recognized as a part of the R/i-

nolophus maclaudi species group (Fahr et al. 2002) due

to the large ears and the poorly developed connecting

process behind the sella (Fig. 3). Within this group it is

the smallest in most metrical dimensions, particularly FA,

cranium, and palatal length. Twelve internal ear folds.

Description. Secondary horseshoe leaflet present and

trilobate, connecting process and anterior face of sella clad

with long hairs, horseshoe densely clad with short hairs

(Fig. 4); dorsal pelage somewhat wooly (length of hairs

ca. 11.9 mm), hairs unicolored, smoky-brown; ventral

pelage more grayish with a slight sheen (length of hairs

ca. 9.4 mm); membranes dark smoky-brown; skin of nose-

leaf dark grey. Connecting process of sella shoulder-like

in lateral view where meeting sella from behind (point of

insertion of shoulder to tip of sella: 1.45 mm). Ears rela-

tively short (49-56% of FA-length), twelve internal folds.

Bulla length: 4.1—5.0, bulla width: 2.05—2.06. Skull axis

in lateral view more or less straight (as opposed to R.

maclaudi). Profile of parietal, in lateral view, is variable

(Fahr et al. 2002, Fig. 3), with both concave (as R. ruwen-

zorii) and more straight outlines (as R. hil//i). Position of

anterior margin of rostral swelling, in lateral view, ranges

between the front and the center of upper M'. Saddle be-

tween inflation and frontal pronounced, posterior slope of

inflation both illustrating steep (more similar to R. ruwen-

zorii) and moderately steep profiles (more similar to R.

hilli) in lateral view (ibid). Squamosal root of zygoma at

©ZFMK

192 Julian C. Kerbis Peterhans et al.

Table 2. External and craniodental measurements of Rhinolophus willardi sp. nov., R. kahuzi sp. nov., R. hilli, and R. ruwenzorii.

Rhinolophus willardi sp. nov. R. kahuzi sp. nov. R. hilli R. ruwenzorii *

FMNH FMNH FMNH FMNH FMNH ZMUZ MRAC Meant+SD Range, sample size

195182 195082 195083 195084 219793 126639 82006M1

Sex é ¢ ¢ 8 3 a: 19 83,9 99

Mass 14 15 16 13 16.5 17.6+1.5 16.0-19.5, n=7

Total 87 92 95 81 92.0 93.745.3 83.0-104.0, n=24

H&B 61 66 73 57 62.7 63.3+5.1 52.0-72.0, n=27

Tail 23.4 26.0 26 22 24.1 293 29.5+2.5 25.0-35.0, n=25

Ear 24.2 29 28 29 34.5 28.5 35.6+2.1 32.0-40.0, n=27

HS-W 12.1 10.9* 10.7* = 10.3* 11.7 11.8+0.8 10.3—13.0, n=20

FA 49.7 51.5 50.8 51.4 54.5 54.3 54.2 57.6+1.9 — 55.0-61.7, n=28

3Meta 33.2 35.0 35.3 36.8 38.9 37.1 39.8+1.7 37.4-43.6, n=26

3Phal 16.0 16.6 15.7 16.8 17.1 17.2 18.3+1.2 15.9-21.1, n=26

3Pha2 25.7 28.0 28.4 27.2 29.5 29.3 30.641.4 27.8-32.8, n=26

4Meta 36.7 38.2 37.8 38.8 41.0 40.9 42.642.0 39.2-46.9, n=26

4Phal 10.2 9.8 9.5 10.2 11.3 10.9 11.440.8 — 9.5-12.8, n=26

4Pha2 15.9 16.5 16.3 16.7 18.5 19.] 19.3+0.8 17.7-20.9, n=26

5Meta 36.3 37.9 38.7 39.3 43.3 41.0 43.442.0 39.6-46.8, n=26

5Phal 11.4 12.0 12.9 12.7 11.7 12.7 13.0+0.7 11.3-14.1, n=25

5Pha2 15.2 15.7 15.3 15.2 16.0 18.5 17.4+0.8 15.7-18.9, n=24

Tibia 20.7 20.0 20.7 21.9 23.8 23.5+1.1 21.7-26.0, n=26

HF,, 10.7 11.0 11.6 10.7 11.1 11.1 11.6+0.8 10.0-13.0, n=26

HF. 12.0 12.5 12.5 12.1 12.3 12.2 13.2+0.8 11.7-14.5, n=12

Crn 22.60 24.20 23.89 23.3 25.41+0.73 24.20-26.37, n=9

CrnC 21.74 22.38 22.07 22.25 23.43 23.02 24.48+0.68 23.65-25.55, n=13

CrmC,), 21.45 21.96 21.62 21.88 23.19 22.79 24.62+0.48 24.16-25.19, n=4

Cbs 19.96 21.24 21.22 = 20.7 22.90+0.50 22.00—23.62, n=9

CbsC 19.28 19.72 19.39 = 19.76 20.47 20.19 21.64+0.66 20.70—22.70, n=13

CbsC yi, 18.85 19.36 18.93 19.36 20.30 19.97 21.8840.77 21.10—22.82, n=4

Mast 10.37 10.67 10.64 10.69 11.58 10.90 11.2 11.9740.26 11.20-12.30, n=18

Zyg 10.30 10.35 10.85 10.57 10.61 10.60 10.9 11.0740.23 10.50-11.40, n=18

BcB 9.00 9.32 9.45 9.36 9.97 9.96 10.1 10.06+0.26 9.85—10.81, n=12

BcH 6.87 6.93 7.23 7.14 7.07 6.92 7.28+0.27 6.85-7.67, n=12

C-C 5.58 5.66 5.71 5.65 5.04 5.67 5.6 5.6140.15 5.30-5.90, n=19

C-Criy 5.4] 5.44 5.58 5.55 4.96 5.64 5.58+0.10 5.46—-5.65, n=3

M?-M? 7.72 7.74 8.07 7.84 7.44 7.81 8.0 7.7240.23 7.30-8.20, n=20

M-MP ay 7.44 7.48 7.75 7.63 7.28 7.91 7.4440.18 7.05-7.75, n=13

C-M? 7.73 7.99 7.97 7.99 8.04 8.08 7.9 8.44+0.23 8.15-8.90, n=14

C-MP aly 71 7.66 7.55 7.56 7.88 7.78 8.29+0.28 7.90-8.70, n=6

M'Br 2.16 2.03 2.24 2.17 1.78 2.12 1.98+0.10 1.88—-2.12, n=4

C-PM? 3.21 3.38 3.30 3.41 3.33 3:99 3.67+40.08 3.59-3.74, n=3

PM?-M? 5.95 6.19 6.22 6.13 6.07 6.57 6.46+0.21 6.15-6.80, n=9

InflB 4.16 4.25 4.16 4.30 — 4.55 4.5640.29 4.40-4.99, n=4

RostrBing 6.06 6.34 6.20 6.43 6.88 6.41 6.74+0.11 6.60-6.97, n=14

PalateL 3.04 3.15 312 3.09 3.40 3.58 3.65+0.21 3.25-4.20, n=19

PalateB 3.37 3.43 3.22 3.38 3.20 3.90 3.6740.17 3.53-3.86, n=3

PoC 3.00 3.06 2.97 2.83 3:2] 2.82 2.5 2.8340.21 2.50-3.30, n=19

ManC 14.37 14.53 14.84 14.49 14.78 14.96 15.4 15.8340.45 15.10-16.45, n=13

ManA 14.19 14.52 14.64 14.44 14.35 14.86 15.48+0.43 15.00-15.82, n=3

C-M/3 8.16 8.40 8.59 8.42 8.29 8.50 8.5 8.7940.18 8.45-9.10, n=15

C-M/3,1, 7.93 8.21 8.32 8.22 TID 8.73+0.20 8.54-8.93, n=3

# AMNH 82394 (holotype), BMNH 55.1187, 60.99 — 60.101, FMNH 144309 — 144312, 160357, IRSNB 7047, LACM 51751,

57774, 57776, MHNG 1873.10 — 1873.14, MRAC 85006M447, 85006M448, 35170, 35173, 35206, 35208, 35211, 35217,

35218. For further details, see Appendix 2.

* Horseshoe width measured from dried skin, hence shrunk and originally probably larger.

Bonn zoological Bulletin 62 (2): 186—202 ©ZFMK

Bats from the Albertine Rift 193

Fig. 3.

glenoid in ventral view broad and strut-like for a length

of 1.2 mm before it steps down anteriorly. Canine and sec-

ond upper premolar almost in contact, anterior upper pre-

molar extruded from the tooth row. Mast nearly equal to

Zyg (ratio: 0.97—1.02).

Comparisons. The following features are shared with all

members of the Rhinolophus maclaudi group from the Al-

bertine Rift: R. ruwenzorii (SW Uganda, eastern DR Con-

Bonn zoological Bulletin 62 (2): 186-202

Photographs (A. Plumptre) of the holotype (FMNH 195182) of Rhinolophus willardi sp. nov.

go and NW Rwanda), R. hilli (SW Rwanda), and R. willar-

di sp. nov. (western slopes of Lake Tanganyika) and dis-

tinguish them from the two West African taxa (R. ziama

and R. maclaudi). Secondary leaflets present beneath the

horseshoe. The anterior margin of the horseshoe has a me-

dian emargination half the height of the horseshoe rim. The

highest point of braincase is at the height of the glenoid

process. The chambers of rostral swellings in dorsal view

are subcircular with slight but well-defined postero-me-

©ZFMK

194 Julian C. Kerbis Peterhans et al.

Fig. 4. | Noseleaf of the holotype (FMNH 195182) of Rhinolo-

phus willardi sp. nov. (Photograph J. Weinstein).

dian angulation. The infraorbital bridge is short and stout.

All of the eastern taxa are smaller in size. These shared

similarities suggest that the eastern and western taxa rep-

resent two distinct clades within the R. maclaudi group.

Rhinolophus willardi sp. nov. averages smaller than R.

ruwenzorii in most measurements, with no overlap in Ear,

FA, 3Meta, 4Meta, 4Pha2, 5Meta, Tib, CnC, CbsC, Mast,

BcB, C-M’?, C-PM?, InflB, RostrB;,,, PalatL, PalatB, and

ManC. Its body mass is also lower. However, the anteri-

or upper molar (M') of R. willardi sp. nov. appears broad-

er than in R. ruwenzorii while it averages similar to R.

ruwenzorii in a few craniodental measurements (C-C, M?-

M®, PoC). Compared to R. hilli, the smallest currently rec-

ognized member of the group, R. willardi sp. nov. is small-

er in most measurements, with no overlap in FA, 3Meta,

3Phal, 3Pha2, 4Meta, 4Phal, 4Pha2, 5Meta, 5Pha2, Tib,

CrnC, CbsC, Mast, BcB, InflB, and PalatL. However, it

is similar or slightly larger than R. hilli in some cranio-

dental measurements (Zyg, C-C, M*-M’, PoC, M1Br) with

tooth rows that are proportionately long (C-M,, C-M’) and

a proportionately higher braincase (BcH).

Rhinolophus willardi has twelve internal ear folds com-

pared with 8—9 in R. ruwenzorii / hilli. The shape of sel-

la, in frontal view, is constricted at the base and rounded

terminally. Maxillary bone in dorsal view visible along the

entire length, whereas in R. hilli and R. ruwenzorii it is

concealed by the rostral inflations. Anterior portion of ros-

Bonn zoological Bulletin 62 (2): 186-202

tral sinus in lateral view steeply rising, inflation almost

completely included in rostral sinus (more similar to R.

hilli). The new species differs from R. Ailli with a brain-

case that, in dorsal view, is constricted behind the mas-

toid process. Posterior slope of braincase in lateral view

more horizontal than the sharply dropping slope of R. Ailli.

Relative size of lower molars very large (even slightly

larger than in hi//i). Zygomatic breadth is more or less

equal to the mastoid width as opposed to R. hilli/ruwen-

zorii where the zygomatic breadth is narrower.

Rhinolophus kahuzi sp. nov. Fahr & Kerbis Peterhans

Kahuzi Horseshoe Bat

Holotype. FMNH 219793 (original number JCK 5406).

Subadult male. Collected on 28 July 2007 by R. Kizun-

gu. Specimen preserved in alcohol with skull removed.

Type locality. Western slope of Mt Kahuzi, Kahuzi-Bie-

ga National Park, South Kivu Province, eastern Democrat-

ic Republic of the Congo, 2°15’09”’S, 28°40°09”E, 2600

m. Found within mixed Afrocarpus and Nuxia forest on

the slope of Mt Kahuzi.

Etymology. The specific epithet refers to the type local-

ity, and is used as a noun in apposition.

Diagnosis. Immediately recognized as a part of the Rhi-

nolophus maclaudi species group (Fahr et al. 2002) due

Fig. 5.

Partial noseleaf and sella of the holotype (FMNH

219793) of Rhinolophus kahuzi sp. nov. (Photograph J. Wein-

stein).

©ZFMK

Bats from the Albertine Rift 195

Fig. 6. Skulls in dorsal, ventral, and lateral views. From upper left, clockwise: Rhinolophus ruwenzorii, holotype, AMNH 82394;

R. hilli, holotype, ZMUZ 126639; R. kahuzi sp. nov., holotype, FMNH 219793; R. willardi sp. nov., holotype, FMNH 195182.

Bonn zoological Bulletin 62 (2): 186-202 OZ7FMK

196 Julian C. Kerbis Peterhans et al.

to the large ears and the poorly developed connecting

process behind the sella. Within this group, it is recognized

by its large ears with 11 internal ear folds.

Description. Body — Moderately large rhinolophid (FA:

54.5 mm, Mass: 13 g). Pelage somewhat wooly with uni-

colored hairs, dorsally dark smoky-brown (length of hairs

ca. 11 mm), ventrally barely paler and slightly more gray-

ish (length of hairs ca. 9 mm). Membranes dark smoky-

brown; noseleaf dark gray.

Noseleaf & ears — Horseshoe with deep median emar-

gination, margin slightly “wavy” on the left side, ragged

(coarsely serrated) on the right side (probably injured).

One distinct triangular flap at the posterior point before

the horseshoe meets the noseleaf, with a rim leading from

cup of nostril backwards to base of triangular flap. Tip of

sella broad and rounded with its greatest width at 4.9 mm

and with a mid-length constriction of 3.8 mm (Fig. 5). An-

terior face of sella and horseshoe densely clad with very

short hairs. Weakly developed secondary leaflets with

ragged edge, completely concealed by horseshoe. Lancet

triangular with narrow tip; connecting process low. Ears

relatively very long (63 % of FA-length), conch with 11

internal folds.

Skull & dentition — Upper incisors bilobed. Anterior up-

per premolar (PM') present on both sides and small, on-

ly slightly extruded from tooth row, canine and anterior

upper premolar (PM?’) not in contact; PM? clearly not as

broad as first upper molar (M’'). In lateral view (Fig. 6),

upper edge of zygomatic arch without notch. Anterior

slope of rostral sinus very angular below inflations (cf. fig.

6 in Fahr et al. 2002). Moderately developed sagittal crest

along the anterior half of the braincase. Infraorbital bridge

short and very stout. In dorsal view (Fig. 6), braincase con-

stricted behind mastoid process. Premaxillae moderately

broad; molars completely concealed by rostrum. Lower

incisors trilobed, slightly imbricated. Middle lower pre-

molar (PM,) present on both sides and: small, complete-

ly extruded from tooth row. Mastoid width broader than

zygomatic width (ratio: 0.92), width across canines (C-

C) small, and anterior upper molar (M') narrow.

Comparisons. Due to its small size (FA<55) and the pres-

ence of a secondary leaflet beneath its horseshoe, R. kahuzi

sp. nov. needs comparison only with those species of the

R. maclaudi group inhabiting the Albertine Rift: R. willar-

di sp. noy. (described above from the Misotschi-Kabogo

highlands), R. hilli (from SW Rwanda) and R. ruwenzorii

(SW Uganda, eastern DR Congo and NW Rwanda). Rhi-

nolophus kahuzi sp. nov. is larger than R. willardi sp. nov.

in most external (e.g. Ear, FA, 3Meta, 3Phal, 3Pha2,

4Meta, 4Phal, 4Pha2, 5Meta, 5Pha2, Tib) and several

craniodental measurements (CrnC, CbsC, Mast, BcB, Ros-

trB;,,, PalatL, PoC) while smaller in a few dental meas-

urements (C-C, M?-M?, M'Br).

Bonn zoological Bulletin 62 (2): 186—202

In most dimensions, the new taxon is similar to R. hilli,

another small member of the group (Table 2). Rhinolophus

kahuzi boasts 11 internal ear folds compared with 12 in

R. willardi sp. nov. and 8—9 in R. ruwenzorii / hilli, and

has very large ears (34.5 vs. 24.2—29 in R. willardi sp. nov.,

and 28.5 in R. hilli. Compared to hilli, R. kahuzi sp. nov.

has very similar external measurements, but with much

smaller width across upper canines (C-C) and molars (M’-

M’), shorter PM’-M’, shorter palatal breadth, but broader

rostrum at height of inflations. The shape of the sella, in

frontal view, is constricted at the base and rounded ter-

minally. Maxillary bone, in dorsal view, visible along en-

tire length (as in R. willardi) whereas in R. hilli and R.

ruwenzorii, it is obscured by the rostral inflations. As does

R. willardi, the new species differs from R. hilli with a

braincase that, in dorsal view, is constricted behind the

mastoid process. Posterior slope of braincase in lateral

view drops sharply at 45° degree angle unlike the more

horizontal plane of R. willardi.

Conservation status. R. ruwenzorii — VU B Lab(ii,111,1v,v)

(six out of 15 localities in protected areas), R. hilli - CR

Blab(iii,v)+2ab(iii,v) (both localities in one protected

area), R. kahuzi (single locality in protected area) & R.

willardi (both localities in unprotected areas): DD?

DISCUSSION

The Rhinolophus maclaudi complex now boasts six

species, three having been described in the past decade.

Shared characters unite the western and eastern branches

of the complex into sister clades, but these relationships

have not been confirmed with molecular data. There ap-

pears to be a cline of decreasing size as one moves from

west to east and then, in the east, from north to south.

Our survey essentially presents the first bat records for

the Misotschi-Kabogo highlands. Since these records were

collected by the ornithological team, the documented rich-

ness of four bat species is certainly only a glimpse of the

total bat diversity. In addition to our records, there is a sin-

gle specimen of Eidolon helvum (MRAC 27113, leg. A.

Prigogine, 18 Feb 1957) from Mt Kabobo. Given that

these few records included a new horseshoe bat (Rhinolo-

phus willardi sp. nov.), as well as a potentially undescribed

pipistrelle bat, Hypsugo cf. eisentrauti, additional discov-

eries are highly likely. Furthermore, the elevational gra-

dient with continuous forest cover across the eastern slope

of the Misotschi-Kabogo highlands would provide un-

tapped opportunities to study bat diversity in relation to

altitude in this outstanding biogeographic hotspot (see

Curran et al. 2012).

These discoveries, over the course of only a few nights

in collecting effort, highlight the importance of survey-

ing remote, undocumented African habitats and further in-

©ZFMK

Bats from the Albertine Rift 197

26° 27° 28° 29° 30° 31° 32°

Rhinolophus ruwenzorii Protected Areas

Countries

Submontane forest

oe (900-1500 m)

Rhinolophus hilli

Rhinolophus willardi sp. nov.

Montane forest

(> 1500 m)

Rhinolophus kahuzi sp. nov. Vebee

Fig. 7. Distribution of species of the Rhinolophus maclaudi-group in the Albertine Rift. Type localities indicated with a black

central dot. Extent of montane and submontane forest from GLC 2000 (Mayaux et al. 2004), protected areas (IUCN categories II

& IV, World Heritage Sites) from WDPA (IUCN & UNEP 2010). For data on localities, see Appendix 2.

Bonn zoological Bulletin 62 (2): 186-202 ©OZFMK

198

Julian C. Kerbis Peterhans et al.

Table 3. List of mammal species currently recognized as endemic to the Albertine Rift.

Taxon (number of endemics, 43-49)

Primates (1)

Gorilla beringei

Eulipotyphla (16-19)

Crocidura kivuana

Crocidura lanosa

Crocidura maurisca

Crocidura niobe

Crocidura stenocephala

Crocidura lwiroensis (sp. nov.)

Myosorex bururiensis

Myosorex jejei

Myosorex kabogoensis (sp. nov.)

Mvosorex babaulti

Mvosorex blarina

Myosorex schalleri

Paracrocidura graueri

Paracrocidura maxima

Ruwenzorisorex suncoides

Scutisorex somereni \”

Suncus hututs??

Sylvisorex lunaris

Sylvisorex vulcanorum

Afrosoricida (1)

Micropotamogale ruwenzorii

Chiroptera (4)

Rhinolophus hilli

Rhinolophus kahuzi (sp. nov.)

Rhinolophus ruwenzorii

Rhinolophus willardi (sp. nov.)

Rodentia (21)

Funisciurus carruthersi

Heliosciurus ruwenzorii

Tachyoryctes ruandae

Delanymys brooksi

Dendromus kahuziensis

Lophuromys medicaudatus

Lophuromys rahmi

Lophuromys stanleyi

Lophuromys woosnami

Dasymys montanus

Dasymys rwandae

Grammomys dryas

Hybomys lunaris

Hylomyscus denniae

Hylomyscus vulcanorum

Mus bufo

Praomys degraaffi

Thamnomys kempi

Thamnomys venustus

Otomys denti

Otomys dartmouthi

Carnivora (0-1)

Genetta victoriae'

Artiodactyla (0-2)

Syncerus matthewsi

Cephalophus rubidus

taxonomic status in question

(Author) Reference

(Matschie, 1903) Groves 2005

(Heim de Balsac, 1968) Hutterer 2005

(Thomas, 1904) Hutterer 2005

(Thomas, 1906) Hutterer 2005

(Heim de Balsac, 1979) Hutterer 2005

(Kerbis Peterhans & Hutterer, 2013)

(Kerbis Peterhans et al., 2010)

(Kerbis Peterhans & Hutterer, 2013)

(Heim de Balsac & Lamotte, 1956) Hutterer 2005

(Thomas, 1906) Hutterer 2005

(Heim de Balsac, 1966) Hutterer 2005

(Hutterer, 1986) Hutterer 2005

(Heim de Balsac, 1959) Hutterer 2005

(Osgood, 1936) Hutterer 2005

(Thomas, 1910) Hutterer 2005

(Kerbis Peterhans & Hutterer, 2009)

(Thomas, 1906) Hutterer 2005

(Hutterer & Verheyen 1985) Hutterer 2005

(de Witte & Frechkop, 1955) Bronner & Jenkins 2005

(Aellen, 1973) Fahr et al. 2002

(Fahr & Kerbis Peterhans, this volume)

(Hill, 1942) Fahr et al. 2002

(Kerbis Peterhans & Fahr, this volume)

(Thomas, 1906) Thorington & Hoffman 2005

(Schwann, 1904) Thorington & Hoffman 2005

(Lénnberg & Gyldenstolpe, 1925) Musser & Carleton 2005

(Hayman, 1962) Musser & Carleton 2005

(Dieterlen, 1969) Musser & Carleton 2005

(Dieterlen, 1975) Musser & Carleton 2005

(Verheyen, 1964) Musser & Carleton 2005

(Verheyen et al., 2009)

(Thomas, 1906) Musser & Carleton 2005

(Thomas,1906) Musser & Carleton 2005

(Verheyen et al., 2003) Musser & Carleton, 2005

(Thomas, 1907) Musser & Carleton 2005

(Thomas, 1906) Musser & Carleton 2005

(Lonnberg & Gyldenstolpe, 1925) Musser & Carleton 2005

(Thomas, 1906) Musser & Carleton 2005

(van der Straeten & Kerbis Peterhans, 1999) Musser & Carleton 2005

(Dollman, 1911) Musser & Carleton 2005

(Thomas, 1907) Musser & Carleton 2005

(Thomas, 1906) Musser & Carleton 2005

(Thomas, 1901) Wozencraft 2005

(Lydekker, 1904) Groves & Grubb 2011

(Thomas, 1901) Jansen van Vuuren & Robinson 2001

' species extends westward into Congo Basin but distribution centered on Albertine Rift

’ montane status unclear

Bonn zoological Bulletin 62 (2): 186—202

Bats from the Albertine Rift 199

dicate how much there is yet to learn of African biodiver-

sity. Montane communities contain reservoirs of biodiver-

sity that are inherently isolated. Their exploration must be

placed at the forefront of survey and conservation efforts,

especially in this era of dramatic climatic change. It had

been previously claimed that the Albertine Rift did not

warrant recognition as a biodiversity hotspot due to inad-

equate knowledge (Myers et al. 2000). Continued surveys,

such as the ones reported on here, are rapidly dispelling

this notion. Since 1999 our teams have described eight

small mammal species new to science from the Albertine

Rift region (Van der Straeten & Kerbis Peterhans 1999;

Kerbis Peterhans & Hutterer 2009; Kerbis Peterhans et al.

2010; Kerbis Peterhans et al. 2013; this paper). The four

new species from the Misotschi-Kabogo Highlands and

Kahuzi-Biega National Park described in this volume

bring the total of Albertine Rift montane endemics to a

minimum of 43-49 species, by far the largest endemic

montane fauna in Africa (Table 3, see also Kerbis Peter-

hans et al. 1998; Kaleme et al. 2007; Carleton et al. 2006).

Clearly it is time to prioritize the status of this exception-

al reservoir of biodiversity as it also houses the richest en-

demic montane faunas of birds (Strattersfield et al. 1998:

Voelker et al. 2010), butterflies (Carcasson 1964) and per-

haps reptiles and amphibian (i.e. Greenbaum & Kusam-

ba 2012 for a recent review) on the continent.

All of the newly described forms have been uncovered

through traditional field surveys to remote areas followed

up by morphological diagnoses and comparisons with rep-

resentatives from adjacent montane blocks. Such efforts

are not possible without the use of museum collections and

their associated reference materials. None of these forms

was resurrected from synonymy nor first diagnosed

through molecular techniques. Our methods are available

to all parties, especially to those custodians in tropical re-

gions and not just those with access to modern laborato-

ries and facilities.

We are pleased to recognize interest and support from

non-governmental organizations (NGOs), who have pro-

vided various groups with the resources to begin surveys

of the unexplored pockets within the ecoregion (see Ac-

knowledgements). Discoveries of rare species or species

new to science have the potential to serve as flagship

species for areas that lack formal protection. Finally, the

timely turnover from field survey to academic manuscript

(as exemplified by this journal) further illustrates that the

oft-ignored small mammal community should be funda-

mental in biodiversity surveys.

Acknowledgements. The expedition to the Misotschi-Kabogo

highlands was initiated by the Wildlife Conservation Society

(WCS). Field work was spearheaded by A. J. Plumptre and D.

Kujirakwinja in conjunction with the Research Staff at Lwiro

(CRSN, Democratic Republic of Congo), the Field Museum of

Natural History (Chicago) and the World Wide Fund for Nature

(WWF). Additional funding came from the Daniel K. Thorne

Bonn zoological Bulletin 62 (2): 186-202

Foundation, the John D. and Catherine T. MacArthur Founda-

tion, and the US Fish and Wildlife Service. In the Kabogo high-

lands, we acknowledge J. Badesire for logistical support, E. A.

Mulungu provided critical field support, and G. Lukole and G.

M. Mwankasala provided security and field logistics and local

introductions. D. Pazo (CRSN, Lwiro) provided details on the

botanical communities of Mt. Kahuzi. We thank J. Mwanga

(Head, Biology) and B. Bajope (Director) of the Centre de

Recherche en Sciences Naturelles at Lwiro for continuous as-

sistance in collecting and with export permits. ECHO provided

air transport. We recognize FMNH staff for their assistance: M.

Schulenberg, J. Phelps, and W. T. Stanley. The senior author ac-

knowledges the Barbara Brown Fund and the African Council

of the Field Museum for financial support and an award from

Roosevelt University (Dean’s Office), which enabled him to

work on this publication in Germany with JF. We are indebted

to Casar Claude (ZMUZ) for the loan of the type of R. hilli,

Manuel Ruedi (MHNG) for a short-term visitor grant to JF that

allowed the study of collections under his care, which includes

new material of R. ruwenzorii, Eileen Westwig and Darrin Lunde

(AMNH) for photographs of the type specimen of R. ruwenzorii,

and John Weinstein (FMNH) for skull and noseleaf photographs

of the new species. Finally, the cooperation of the ornithologi-

cal community provides vital continuous support in document-

ing Chiropteran diversity.

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Appendix 1.

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Rhinolophus ruwenzorii J. Eric Hill, 1942

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RG 35170, -173, -174, -206, -208, -211, RG 35216 — 219, 3 22,

4 2, 3 unsexed, leg. R. P. M. J. Celis & M. Lejeune,

28.XI1.1966); Matata Cave (1160 m, Kibali-Ituri) (Hayman et

al. 1966; Fahr et al. 2002; Csorba et al. 2003; IRSNB 7048, 2,

skull only, leg. J. Hiernaux, 10.VHI.1947); Matupi Cave (3500

ft., Mt Hoyo, S of Irumu, Ituri Forest) (Hayman 1960; Fahr et

al. 2002; Csorba et al. 2003; BMNH 60.99 — 101, 3 ¢, 2 skin

& skull, 1 alc., leg. A. E. Wright, 14.&16.1X.1959); Mt Hoyo

(1200 m, Ituri Forest) (Hayman et al. 1966; Fahr et al. 2002;

Csorba et al. 2003; IRSNB 7049, &, skull only, leg. J. Hiernaux,

10. VHI.1947); Mt Hoyo (Komanda) (MHNG 1873.11 — .14, 3

3,1 9, alc., leg. F. Meier, 21.VIII.1976); Pahoni (N’Duye)

(MHNG 1873.10, 9, alc., leg. F. Meier, 6. VIII.1976).

RWANDA: Mutura (2200 m) (Fahr et al. 2002; MRAC 85006

M 447, -448, 2 3, alc. & skull, leg. Baeten & Janssens,

16.XII.1982).

UGANDA: Itama Mine (1615 m, Bwindi-Impenetrable-NP,

Kigezi) (Smith & Hood 1980; Fahr et al. 2002; Csorba et al.

2003: as “Kigezi Highlands”; LACM 51750, 57772 — 777, 3 22,

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4 29, 1 skin & skull, 3 ale. & skull, 3 alc., leg. A. L. Archer,

31.01.1967); Luhizha Mine [Ruhizha wolfram mine] (2286

m, Bwindi-Impenetrable-NP, Kigezi) (Smith & Hood 1980; Fahr

et al. 2002; Csorba et al. 2003: as “Kigezi Highlands”; LACM

51747 — 749, 2 23, 1 9, 3 skin & skull, leg. R. Glen & A.

Williams, 27.11.1967); near Mahoma River (cave, 6700 ft.,

above Ibanda, E-slope Rwenzori Mts.) (Hayman 1957; Fahr et

al. 2002; Csorba et al. 2003: as “Ibanda”; BMNH 55.1187, °,

alc. & skull, leg. G. O. Evans, 22.VIH.1952); right bank of

Mubuku River (6900 ft., above confluence with Mahoma Riv-

er, Rwenzori Mts.) (Kityo & Kerbis 1996: as “Rwenzori Mts.

at 2100 m”; Fahr et al. 2002; Thorn et al. 2009; FMNH 144309

[at Makerere Univ.], ¢, skin, skel. & skull, leg. W. T. Stanley,

25.X1.1990); Nteko Parish (1600 m, edge of Bwindi-Impene-

trable-NP, Bufumbira) (Fahr et al. 2002; Thorn et al. 2009;

FMNH 160357 [exchanged], @, skin, skull & skel., leg. R. M.

Kityo, 19.V.1997); Nyabitaba (2591 m, Mubuku Valley, E-slope

Rwenzori Mts.) (Smith & Hood 1980; Fahr et al. 2002; Csorba

et al. 2003; LACM 51751, @, leg. R. Glen & A. Williams,

5.VI.1967); Nyabitaba Hut (8750 ft., ght bank of Mubuku Riv-

er, below confluence with Bujuku River, Rwenzori Mts.) (Ki-

tyo & Kerbis 1996: as “Rwenzori Mts. at 2700 m”; Fahr et al.

2002; Thorn et al. 2009; FMNH 144310, @, alc., leg. W. T. Stan-

ley, 11.XII.1990; FMNH 144311, -312, 2 42, 1 alc., 1 skin, skull

& skel., leg. J. C. Kerbis, 19&20.1V.1991).

Rhinolophus hilli Aellen, 1973

RWANDA: Uwinka (P.N. de Nyungwe, 2512 m) (Aellen 1973;

Fahr et al. 2002; Csorba et al. 2003: as “Cyangugu”; ZMUZ

126639, holotype, °, alc. & skull, leg. U. Goepel, 25. VHI.1964,

F-N° 481); Ruta Bansugera (P.N. de Nyungwe, 1750 m) (Baeten

et al. 1984; Fahr et al. 2002; MRAC 82006 M 1, 9, leg. F. De

Vree et al. 19.X.1981).

OZFMK

202 Julian C. Kerbis Peterhans et al.

Rhinolophus willardi sp. nov.

D.R. CONGO: Kilicha River (Mt Kabogo, near Kalemie, 1880

m) (FMNH 195182, <4, alc. & skull, leg. A. Plumptre & E. A.

Mulungu, 28.11.2007); 2" camp (Mt Kabogo, 4 km SW Talama,

1950 m) (FMNH 195082 — 084, 1 2, 2 29, skin & skull, leg.

B. D. Marks, 14. & 16.11.2007).

Appendix 2.

Rhinolophus kahuzi sp. nov.

D.R. CONGO: Mt Kahuzi (Kahuzi-Biega-NP, 2560 m) (FMNH

219793, 4, alc. & skull, leg. R. Kizungu, 28. VII.2007, F-N° JCK

5406).

Gazetteer of collecting localities (updated from Fahr et al. 2002)

D.R. CONGO:

Kilicha River 5°067°19"S, 29°03°56”E

2™ camp 4°59°29”S, 29°04’°49”E

Mt Kahuzi 2°12°07°S, 28°40°24”E

Butahu Valley 0°19°35”N, 29°49°20”°E

Butembo O0°09"N, 29°17°E

Kibwe Lya Mikako Cave 0°14°42”S, 29°01V’E

Matata Cave

Matupi Cave

Mt Hoyo

Pahoni [= Paoni]

UGANDA:

Itama Mine

Luhizha Mine

0°58°30°N, 29°40°E

1°16712”N, 29°54°36"E

1°13°N, 29°49°E

1°58.5°N, 28°52.5°E

0°59°S, 29°4VE

1°01°53”S, 29°46°42”E

Mahoma River 0°21°39”N, 30°00°06”E

Mubuku River 0°21°50”N, 29°59°53”E

Nteko Parish 1°02°16"S, 29°36°E

Nyabitaba (Hut) 0°21°26”N, 29°58°317E

RWANDA:

“near Cyangugu”

Mutura 1°37°S, 29°23°E

Ruta Bansugera 2°25°S, 29°10°E

Uwinka 2°29’S, 29°12’E

Bonn zoological Bulletin 62 (2): 186—202

near Kalemie, Misotschi-Kabogo highlands, 1880 m

4 km SW Talama, Misotschi-Kabogo highlands, 1950 m

Kahuzi-Biega National Park, 2560 m

W-slope Rwenzori Mts., 7500 ft. [2286 m]

Kivu Province, 1760 m

Kasuo, Lubero Distr., 1500 m

Kibali-Ituri, 1160 m

Mt Hoyo, S of Irumu, Ituri Forest, 3500 ft. [1066 m]

Ituri Forest, 1200 m

N’Duye [= Nduye], near Biasa River,

NE edge of “Réserve de Faune a Okapis”

Bwindi-Impenetrable-NP, Kigezi Highlands, 1615 m

Bwindi-Impenetrable-NP, Kigezi Highlands, 2286 m

above Ibanda, E-slope Rwenzori Mts., 6700 ft. [2042 m]

above confluence with Mahoma River, E-slope Rwenzori Mts.,

6900 ft. [2103 m]

edge of Bwindi-Impenetrable-NP, Bufumbira, 1600 m

right bank of Mubuku River, below confluence

with Bujuku River, E-slope Rwenzori Mts., 8750 ft. [2667 m]

Parc National de Nyungwe, 2500 m

2200 m

Parc National de Nyungwe, 1750 m

Parc National de Nyungwe,

Préfecture de Cyangugu [= Shangugu], 2512 m

OZFMK

Bonn zoological Bulletin 62 (2): 203-219

December 2013

Mammals, other than bats, from the Misotshi-Kabogo highlands

(eastern Democratic Republic of Congo),

with the description of two new species (Mammalia: Soricidae)

Julian C. Kerbis Peterhans'”, Michael H. Huhndorf’, Andrew J. Plumptre’, Rainer Hutterer’,

Prince Kaleme** & Benjamin Ndara‘

' Evelyn T. Stone College of Professional Studies, Roosevelt University, 430 S Michigan Ave., Chicago, IL, 60605, USA

’ Science & Education, Field Museum of Natural History, Chicago, IL, 60605-2496, USA,

Corresponding author: jkerbis@fieldmuseum.org

’ Albertine Rift Programme, Wildlife Conservation Society, PO Box 7487, Kampala, Uganda

* Zoologisches Forschungsmuseum Alexander Koenig, D-53113 Bonn, Germany

> Department of Botany and Zoology, Stellenbosch University, Matieland, South Africa

° Centre de Recherche en Sciences Naturelles (CRSN), Department of Biology, Lwiro, Democratic Republic of Congo

Abstract. The Albertine Rift has recently been declared by Plumptre et al. (2007a) as one of Africa’s biodiversity hotspots

due to the high number of species and high levels of endemism. Here we report on a recent but brief survey of mammals

from the Misotshi-Kabogo highlands of the eastern Democratic Republic of Congo. This represents the first effort to doc-

ument the mammal community of the region. Remarkably, the collection includes at least three species of mammals new

to science: two species of shrews (Soricidae) described here and one species of horse-shoe bat (Rhinolophidae, Kerbis

Peterhans et al. 2013). A total of five mammal taxa are now known exclusively from this forest. One of the shrews (Mvosorex

n. sp.) which we describe here continues to fill in distributional gaps of this typically montane-restricted genus. Several

of the large mammal taxa provide significant range records, especially as southern terminals for species’ distribution. To-

gether the data recorded here demonstrate the need for conserving this isolated and heretofore unrecognized reservoir of

biodiversity.

Key words. Soricidae, Mammalia, Democratic Republic of Congo, Misotshi-Kabogo Highlands, Albertine Rift, endemism,

new species.

INTRODUCTION

The Albertine Rift montane zone of central Africa, extend-

ing from the Blue Mountains in the north (ca. 2°N,

30°30’E) to the southern end of Lake Tanganyika (ca. 9°S,

30°30’E) has only recently been declared one of Africa’s

biological hot spots due to high levels of species diversi-

ty and endemism (summarized in Plumptre et al. 2007a).

Although many surveys have been conducted and pub-

lished on the gorilla-inhabited Virunga-Bwindi heartland,

many isolated areas, especially Congolese landscapes pe-

ripheral to this heartland, have never been properly stud-

ied. Such areas include the Blue Mountains to the north

and the Itombwe Massif, Misotshi-Kabogo Highlands and

Marungu Plateau to the south. The Misotshi-Kabogo high-

lands (5°—-6° S) overlook the middle of Lake Tanganyika.

The last survey on vertebrates in the region was conduct-

ed by Prigogine in the 1950’s (Prigogine 1960) who doc-

umented the bird community and described one species

new to science, the Kabobo Apalis (Apalis kaboboensis).

As for mammals, we are only aware of Prigogine’s list

(1960) of large mammals and short notes on a single squir-

rel and single colobus monkey both collected by Prigogine

Received: 02.08.2012

Accepted: 05.08.2013

during his time there and both described as sub-specifi-

cally distinct (Verheyen 1959, 1960). Here we report on

a brief survey in these highlands that provides the first

records of terrestrial small mammals for the region, includ-

ing two species (Soricidae) that are new to science. Al-

though some of this information has been published al-

ready (see Plumptre et al. 2007b), these data are so im-

portant that formal publication is warranted.

MATERIALS AND METHODS

Different means were used to survey the small and large

mammal communities. Small mammals were actively col-

lected from three campsites between 28 January, 2007 and

26 February, 2007, in the Misotshi-Kabogo highlands just

north of the town of Kalemie (once known as Albertville,

05° 56’S, 29° 12’E), eastern Democratic Republic of Con-

go. Shrew sampling was attempted using pitfall bucket

lines. These consisted of a total of fifteen to seventeen 5

liter buckets individually set five meters apart. Plastic

Corresponding editor: J. Decher

204 Julian C. Kerbis Peterhans et al.

sheeting, ca 25 cm high, served as a drift fence over the

midline of the buckets (see Voss & Emmons 1996). A pit-

fall line was set at each of the first two camps (68 buck-

et nights and 75 bucket nights respectively) but without

any success. Fourteen conventional ‘rodent’ trap lines col-

lected all shrews and mice. Camp one had six trap lines

while camps two and three had four each. Trap number

ranged from 35 to 63 traps per line. Trap lines varied in

trap number and duration due to shifting camps and vary-

ing personnel availability. The majority of traps were

placed on the ground. Others were set near streams and

others above the ground on vines, tree limbs or fallen logs.

A mixture of peanut butter and oatmeal was used as bait

but occasionally fish and corn kernels were employed. All

of the trap lines and pitfall lines were set within 500 me-

ters of the respective camp site.

We used a combination of methodologies to determine

which large mammal species currently occur in the Mis-

otshi-Kabogo forest (Plumptre et al. 2007b). We inter-

viewed hunters actively working in the forest today as well

as former military who were active there during times of

civil strife. These people were interviewed about which

large mammal species they had seen in the forest and

where they saw them. We used Kingdon’s (1997) Field

Guide to African Mammals to show local hunters pictures

of the animals and then cross-checked their identifications

by asking them about the behavior of the animal. We al-

so asked people in villages to show us any skins of ani-

mals to cross-check the list obtained by talking to hunters.

One hyrax skin and several carnivore skins were pur-

chased from local people. A brush-tailed porcupine

(Atherurus africanus) was snared by one of the camp staff

and a black-fronted duiker (Cephalophus nigrifrons) was

found in a snare while surveying transects (see below).

We also attempted to collect data on large mammals by

walking along transects in fixed compass directions from

one of eight camp sites (Plumptre et al. 2007b). Where

only gallery forest existed, zigzag transects were covered

to maximize the time spent in this forest type before mov-

ing towards the next patch of forest. The zigzag transects

extended beyond the forest by 500 meters before return-

ing back to the forest. In this way some sampling was

made in the woodlands around the gallery forests but most

of the effort was concentrated in the forests. Densities for

certain species could be calculated for the forest and the

surrounding 500 m of woodland from these data. Once a

patch of forest had been surveyed with the zigzag tran-

sects, a walk was made following a fixed compass direc-

tion to the next patch of forest.

At each camp site a transect or reconnaissance walk was

made each day following a compass direction where pos-

sible but deviating when the terrain became impassable

or if in gallery forest. Much of the Misotshi-Kabogo for-

est consists of very steep hillsides which have been cut

by fast running rivers. Many of the rivers have cliffs along

Bonn zoological Bulletin 62 (2): 203-219

their course making traversing them very difficult. We

therefore used reconnaissance walk methods whenever we

reached a point that was impassable and then selected a

new compass direction. GPS positions were taken for any

sighting of an animal or its signs (nest or dung of elephant

and pigs) and also every 250 m a GPS position was tak-

en with a description of the habitat type. This allowed us

to map where teams had visited and also helped ground

truth the satellite classifications.

Data collected for all specimens included the macro-

habitat, sex, and reproductive condition. On the morning

of capture, the following standard external measurements

were taken: total length (TL), tail vertebrae length (TV),

hind foot length (HF), ear length (E) and mass (WT). Head

and body length (HB) is determined by subtracting TV

from TL; due to damage to specimens and tails by ants,

tables display HB rather than TL. Metrics posted in the

Tables are for adult specimens only. For taxa represented

by multiple individuals, metrics are pooled, then averaged.

CR refers to crown-rump length of embryos. Voucher

specimens were prepared as study skins and skeletons or

were preserved in 10 % formalin. For the latter group,

skulls were later extracted at the Field Museum of Natu-

ral History in Chicago and transferred to 70 % ethanol.

The nature of the specimen preserved (cond) is recorded

as ssk (skin, skull and skeleton), alc (alcoholic carcass),

asr (alcoholic carcass with skull removed), or sko (skull

and skeleton only).

Cranio-dental measurements were taken at the Field

Museum of Natural History using Mitutoyo CD-6”CSX

calipers to the nearest .01 mm. These include the follow-

ing: CI (condylo-incisive length), GB (greatest breadth of

skull), MX (breadth of the maxilla), UTR (length of the

upper tooth row), HCC (height of the cranial capsule,

measured in the sagittal plane), PGL (post-glenoid

breadth), IO (inter-orbital breadth), MD (mandible length

from the tip of the anteromost incisor to rear of the

mandibular condyle), LTR (length of the lower tooth row),

COR (height of the coronoid process of the mandible).

Measurements are given in millimetres, body mass in

grams.

Full technical names (based on the respective chapters

in Wilson & Reeder 2005) for larger mammals are listed

in Table 2; for the small mammal species they are given

in the text.

Tissue samples were stored in the field in a saturated

NaCl / EDTA buffer. Upon returning from the field, tis-

sues were stored in an ultra-cold freezer at -70°C. In the

field, animals were handled in accordance with American

Society of Mammalogists guidelines (Animal Care and

Use Committee 1998).

Specimens discussed herein were collected during the

course of the Misotshi-Kabobo expedition (see Acknowl-

edgements), supplementing the records of Prigogine

(1960). Specimens were compared with material from the

OZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands 205

@ suconso Pg

LENDU PLATEAU

UGANDA (

a5 ¢

o BWAMBA ¢

B kiwave j

RUWENZORI

EQUATOR

= ff

- M) IMPENETRABLE

<I LAKE EOWARD jh conEST : LAKE VICTORIA

oO oa 1. Kasae ~.

KAHUZI-BIEGA NP 5) a Pa

iS) Me ; ~

= MT MUHAVURA Bas

= voLCANoES = { ee

oe 4

LAKE KN i

2 4 Oe

\ wa

« \ RWANDA i

= ou \ 7

fs TOMBWE FOREST ¢

o ‘

eo)

|] MISOTSCHI-KABOGO.

} FORES

i] LUKUGAR

oO ee FORESTED AREA

Uvira |

Mookuingu R Talama

25 bmp SW Tatahbs

MisolschiKebego Fores \

i} J hmiW Magen!

| w

| Kalemes |

Fig. 1. Map of the Albertine Rift with the Misotshi-Kabogo highlands (a), and position of the three collecting localities (b).

following institutions: The Field Museum of Natural His-

tory, Chicago (FMNH); the United States National Mu-

seum, Washington D.C. (USNM); and the Zoologisches

Forschungsmuseum Alexander Koenig, Bonn (ZFMK).

All specimens are deposited at the Field Museum of Nat-

ural History, Chicago, IL. JCKP are the abbreviations for

the senior author. Unless otherwise noted, taxonomy and

distributional notes for all mammals are from the relevant

chapters in Wilson & Reeder (2005) and Kingdon (1997).

STUDY AREA

Prigogine (1960) and Plumptre et al. (2007b) reviewed

some of the biology of the area and provided notes on bio-

geographic boundaries. The Misotshi-Kabogo highlands

are separated from the Itombwe massif to the north by a

50 km wide savannah through which crosses the Kilomb-

we River (Fig. la). This savannah constitutes a barrier for

some bird species (Prigogine 1960), which must be true

for some mammals as well. The absence of forest in the

Kilombwe valley may be caused by the low rainfall and

low humidity due to strong winds that favour herbaceous

vegetation. The Marungu highlands lie to the south and

are separated from the Misotshi-Kabogo highlands by the

Lukuga River. Both rivers formed following the tecton-

ic activity and uplift that created these highlands on the

western shore of Lake Tanganyika. To the west, gallery

forest and savannah continue for hundreds of kilometres

south of the main Congo Basin Forest Block (Plumptre

Bonn zoological Bulletin 62 (2): 203-219

et al. 2007b). The Misotshi-Kabogo highlands run approx-

imately 100 km along the escarpment above Lake Tan-

ganyika at an estimated width of 10-20 km. On the west-

ern slopes, the forest descends the valleys as gallery for-

est; while in the east, it is continuous along Lake Tanganyi-

ka where it reaches 2500-2750 m (Prigogine 1960;

Plumptre et al. 2007b). The forested escarpment persists

uninterrupted to the lakeshore at 770 m. The main forest

block is approximately 1,000 kn in size.

Three camps (Fig. 1b) for small mammals within the

Misotshi-Kabogo forest were accessed from two differ-

ent fishing villages along the Lake Tanganyika shoreline

(Huhndorf & Kaleme 2007). The first camp (29 Jan—7

Feb) was established approximately 3 km west of Miz-

imu fishing village (5°28°45”S, 29°16°22”E) at an eleva-

tion of 1250 meters. Traps were placed in primary forest

along the ridge and peripheral to a creek edge. Trap lines

at this location were then relocated along the forest edge

and secondary growth. This habitat can be described as

transitional between lowland and montane and was close

to the forest edge. The other camps were established fur-

ther north and were accessed from Talama fishing village

near the base of Mt. Misotshi. After climbing a steep

grassy ridge, the forest was reached and camp was set

within a horseshoe bend of the Mukungu River. The sec-

ond camp (4°59°29"S, 29°4°49”"E, 1950 m) (13 Feb

2007-17 Feb 2007) was established approximately 4 km

southwest of Talama. Some traps were again set along the

riverbank but mostly along a dry hill slope. Later, traps

were shifted to higher ground with fewer, but larger trees

©ZFMK

206

Julian C. Kerbis Peterhans et al.

Table 1. Small terrestrial mammals of the Misotshi-Kabobo highlands. Bold = Albertine Rift endemic, SL = southern limit of dis-

tribution.

Camp 1 3 2

Habitat Degraded lowland-montane drier forest wet montane forest

forest transition on eastern escarpment

Elevation 790 m 1250 m 1600 m 1950 m

Species

Crocidura c.f. dolichura ]

Crocidura c.f. fuscomurina I

Crocidura olivieri l 7

Crocidura n. sp. ]

Myosorex n. sp. 1

Graphiurus murinus l

Lophuromys aquilus 3 3 PS)

Hybomys aff. univittatus 3 6

Hylomyscus stella 12 l

Mus minutoides 2

Mus triton l

Pelomys minor l

Praomys degraaffi SL 2

Praomys jacksoni 4 43 15 l

Praomys sp. ]

Rattus rattus it

Total specimens (# species) 11 (5) 72 (9) 23 (5) 11 (5)

Trap effort and success Na 72/783= 23/630= 11/520=

2.1%

and more abundant undergrowth. This habitat was typi-

cal wet montane forest. The third camp (4°59’05"S,

29°5°34”E, 1600 m) was located approximately 2.5 km

southwest of Talama between 20 and 25 February, 2007.

Here, traps were set along a ridge adjacent to a fern prairie,

and others along large rocks and a stream along the slope

of the ridge. The camp 3 habitat can be described as dri-

er forest, located on the eastern edge of the escarpment,

also close to the forest edge. Finally, eleven specimens

were collected in a degraded agricultural area on the out-

skirts of Mizimu at 790 m (5°27°59”S, 29°17°16”E) while

switching between camps (February 6—7, 2007).

RESULTS

Results of the trapping efforts for small mammals at the

four camps are presented in Table 1. Trap success marked-

ly declined with elevation. At 1250 m, trap success was

9.2 % (nine species and 783 trap nights); at 1600 m, 3.7%

(five species and 630 trap nights); at 1950 m, 2.1 % (five

species and 520 trap nights). Results from the large mam-

mal surveys are presented in Table 2, along with histori-

cal data from Prigogine (1960). Metrical data on the

shrews collected, including identification, date, sex, age

Bonn zoological Bulletin 62 (2): 203-219

9.2%

and external measurements are presented in Table 3. and

their cranio-dental measurements are presented in Table

4. For the rodents, external measurements are presented

in Table 5 and select cranio-dental measurements are

found in Table 6. Although each camp was not trapped

with identical intensity, trap success and species diversi-

ty was highest at 1250 m. Small mammal abundance de-

creased with elevation although species diversity (n = 5)

was similar at 1600 m and 1950 m. Of the two species

new to science, one was captured at the lowest camp (Cro-

cidura n. sp.) and one (Myosorex n. sp.) at the highest.

The following details some of the more noteworthy re-

productive, taxonomic, and distributional issues posed by

the specimens as well as formal descriptions of the two

new species of Soricidae.

HyYRACOIDEA

Dendrohyrax c.f. dorsalis. This dorsal pelt, recovered

from a local hunter, cannot be definitively identified to

species as diagnostic elements are missing (e.g. skull, chin

and rostrum portion of the pelt) while others show mixed

affinities. The dorsal white patch is long (90-100 mm) and

creamy white. The midline dorsal hair is black, forms a

©ZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands

207

Table 2. Large mammals of the Misotshi-Kabogo Highlands. PL = Plumptre et al. (2007b); PR = Prigogine (1960); FMNH, cat-

alogue #; SL = demarcates southern limit of species; Albertine Rift endemics in bold; S = sight record; F = feces record; N = nest

record; H = reported by at least 3 hunters.

Species

Dendrohyrax c.f. dorsalis (Fraser, 1855)

Loxodonta africana (Blumenbach, 1797)

Cercopithecus ascanius (Audebert, 1799)

Cercopithecus denti (Thomas, 1907)

Cercopithecus lhoesti (P. Sclater, 1899)

Cercopithecus doggetti (Pocock, 1907)

Lophocebus albigena (Gray, 1850)

Papio anubis (Lesson, 1827)

Colobus angolensis prigoginei (Verheyen, 1960)

Pliocolobus foai oustaleti (Trouessart, 1906)

Pan troglodytes (Blumenbach, 1775)

Manis gigantea (Illiger, 1815)

Manis tricuspis (Rafinesque, 1821)

Leptailurus serval (Schreber, 1776)

Profelis aurata (Temminck, 1857)

Civettictis civetta (Schreber, 1776)

Genetta genetta (Linnaeus, 1758)

Genetta maculata (Gray, 1830)

Nandinia binotata (Gray, 1830)

Galerella sanguinea (Riippell, 1835)

Mungos mungo (Gmelin, 1788)

Crocuta crocuta (Erxleben, 1777)

Canis adustus (Sundevall, 1847)

Mellivora capensis (Schreber, 1776)

Hylochoerus meinertzhageni (Thomas, 1904)

Potamochoerus larvatus (F. Cuvier, 1822)

Oreotragus oreotragus (Zimmermann, 1783)

Syncerus caffer (Sparrman, 1779)

Tragelaphus euryceros (Ogilby, 1837)

Tragelaphus scriptus ornatus (Pocock, 1900)

Cephalophus nigrifrons kivuensis

(Lonnberg, 1919)

Cephalophus weynsi (Thomas, 1901)

Philantomba monticola (Thunberg, 1789)

Funisciurus carruthersi chrysippus

(Thomas 1923)

Funisciurus pyrropus akka (de Winton, 1895)

Heliosciurus rufobrachium ssp. (Waterhouse, 1842)

Heliosciurus ruwenzorii vulcanius

(Thomas, 1909)

Paraxerus boehmi vulcanorum (Thomas, 1918)

Protoxerus stangeri kabobo (Verheyen, 1959)

Atherurus africanus (Gray, 1842)

sort of dorsal crest and is quite long (60-80 mm in front

of the dorsal white patch, 45—65 mm behind it), similar

to Dendrohyrax dorsalis sylvestris skins from Ghana, West

Africa in the collections at the FMNH. If confirmed as D.

dorsalis, this would prove to represent the SE terminus

of the species’ distribution.

Bonn zoological Bulletin 62 (2): 203-219

Common Name

Tree hyrax

African elephant

Red-tailed monkey

Dent’s mona monkey

L’Hoest’s monkey

Blue monkey

Grey-cheeked mangabey

Olive baboon

Angola colobus

Red colobus

Chimpanzee

Giant pangolin

Tree pangolin

Serval

Golden cat

African Civet

Common genet

Rusty-spotted genet

2 spotted palm civet

Slender mongoose

Banded mongoose

Spotted hyena

Side striped jackal

Honey badger

Giant forest hog

Bush pig

Klipspringer

Cape buffalo

Bongo

Bushbuck

Black-fronted duiker

Weyn’s duiker

Blue duiker

Carruther’s mountain squirrel

Fire-footed rope squirrel

Red-legged sun squirrel

Ruwenzori sun squirrel

Boehm’s bush squirrel

Giant forest squirel

Brush tailed porcupine

PROBOSCIDEA

FMNH

195080

195087—

195088

195089

195090

195091

195178

Reference

Skin

PR, H

PL (S)

PR, PL (S)

PL (S)

PR, PL (

PL(N,S

PL (H)

PL(H)

PL (H)

PL(H)

Skin

PL (H)

PR, PL (F)

PR, PL (S, H)

)

PR, PL (H)

PL (H)

PR, PL (S, H)

PL (H)

Skull, skin

Loxodonta africana. Elephants no longer occur in the for-

est but occasional crop raiders are reported. Prigogine

(1960) stated that in the 1950’s elephants of small size

(Loxodonta cyclotis’?) were known from the area.

©OZFMK

208 Julian C. Kerbis Peterhans et al.

PRIMATES

Cercopithecus lVhoesti. This Albertine Rift endemic

recorded here and before by Prigogine (1960) reaches its

southern limit in these highlands.

Cercopithecus doggetti. Current taxonomy recognizes this

taxon as a distinct species, adding it to the expanding list

of Albertine Rift endemics. It also reaches its southern lim-

it here.

Colobus angolensis prigoginei. This taxon, notable by its

all-white tail, is only known from these highlands. It is

endangered as only one sight record of this taxon was

made during the recent survey, and none were seen along

transects.

Pan troglodytes. Surprisingly, chimpanzees were not dis-

cussed or listed by Prigogine (1960). Chimpanzees reach

their southern limit on the Marungu highlands to the south

where Noack (1887) described a population under the

name marungensis. However, our aerial reconnaissance

shows grassy highlands with only traces of gallery forest

remaining there. Details on our chimpanzee surveys and

estimates can be found in Plumptre et al. (2007b). Their

surveys documented nests suggesting a density of 1.22

chimpanzees per km’ in the Misotshi-Kabogo Highlands.

With a forest block of approximately 804 km’, this trans-

lates into an estimated population of 977 + 252. It appears

that local taboos against hunting chimpanzees in the for-

est highlands have been effective in their conservation.

PHOLIDOTA

Manis gigantea. The Misotshi-Kabogo highlands appear

to represent the southern terminus for this forest depend-

ent species.

CARNIVORA

Genetta maculata. Two skins (FMNH 195087—195088),

one of a juvenile. Both skins have tails with black tips and

with dorsal crests starting midway down the torso, as de-

scribed in Kingdon (1977). The spots are not in discrete

linear rows.

ARTIODACTYLA

Hylochoerus meinertzhageni. This record represents the

southern limit for this spectacular suid. As it is in Kenya,

the species was only found at high elevation (2500 m). Pri-

Bonn zoological Bulletin 62 (2): 203-219

gogine (1960) also confirmed this high elevation restric-

tion (2400 m) in the Misotshi-Kabogo highlands.

Tragelaphus eurycerus. As for the previous record, this

also represents the southern limit for a magnificent ani-

mal (Ralls 1978). As for Prigogine (1960), our record is

from high elevation at 2500 m, sharing a distinct biotope

with Hylochoerus meinertzhageni in areas at the periph-

ery of its range (Ralls 1978). Given the shared elevation

range of this population with the montane Kenyan popu-

lation, the possibility of taxonomic similarities between

these populations should be investigated.

SORICOMORPHA

Crocidura c.f. fuscomurina (Heuglin, 1865). A single rep-

resentative (FMNH 195071) was taken in a Museum Spe-

cial trap. As it was prepared as a skeleton, no details of

the pelage coloration or caudal bristles are available. The

identification of the specimen remains tentative. Sex was

not determined. In size, it ranks as one of the smaller Cro-

cidura known for the region (4.8 g, CI=17.3 mm). Brain-

case is small in both height (HCC 4.1 mm) and breadth

(GB 7.6 mm); maxilla is narrow (MX 5.3 mm), dentition

light. First upper unicuspid by far the largest, 2-3 times

larger than second and third, both of which are sub-equal

in size. Cingula of unicuspids well developed; parastyle

reduced and not projecting; upper P4 vacuous posterior-

ly; upper M3 short (reduced). Lower incisor smooth and

without denticulations; lower M3 simple and without

talonid basin.

This specimen approximates Crocidura ansellorum

(Hutterer & Dippenaar, 1987) from Northern Zambia in

many cranio-dental dimensions (CI=17.3 mm vs. C. ansel-

lorum with CI of 16.8-17.7 mm) but its small brain case

(GB=7.6 mm, HBC=4.1 mm) distinguishes it (Crocidu-

ra ansellorum, GB=8-8.4 mm, HBC 4.64.7 mm). All cra-

nial measurements fall within the ranges of members of

the Crocidura fuscomurina group, as reviewed in Hutter-

er (1983) who described this as a savannah species; here

it was captured at the forest edge in a transitional brushy

area near a grassy hillside.

Crocidura c.f. dolichura Peters, 1876 (FMNH

195070).This taxon is recognized by its delicate dentition,

three upper unicuspids with heavy cingula, and a naked

tail that is significantly longer then HB. Originally de-

scribed from Cameroon, its range extends from Nigeria

in the west across the Congo basin into western Uganda,

Burundi and Rwanda in the east (Hutterer 2005; FMNH

specimens). Variation within the group is not trivial and

the taxon, as currently recognized, is in need of revision.

OZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands 209

Adult male, testes 3x2 mm. Teeth minimally worn. Cap-

tured in museum special trap; skull broken at capture. Dor-

sal pelage with silvery grey base (70 % of hair length),

followed by light brown (15 %) and tipped with darker

brown (15 %) yielding an appearance that is dark grey

washed with brown. Belly fur silvery grey throughout. Tail

length ca. 130 % of HB. Lower first incisor with two sub-

tle denticulations. Maxillary unicuspids with heavy cin-

gula. First upper unicuspid the largest, twice as large as

second, third unicuspid the next largest, 50 % larger than

second. These proportions differ from those evident in

Brosset et al.’s (1965) figure of the type specimen of Cro-

cidura dolichura where the second and third unicuspids

are sub-equal in size, and only slightly smaller than the

first. Upper P4 lightly built, vacuous posteriorly. Upper

M3 well developed. Dental characteristics similar to

FMNH 162198 from Gabon, but differing in its longer tail

(132 % of HB compared with 111 %) and more grey-like

dorsal pelage (vs. cinnamon brown).

Crocidura — olivieri | (Lesson, 1827) (FMNH

195072—195078, 195180).These ubiquitous ‘giant shrews’

of forested sub-Saharan Africa were quite common at low-

er elevations of the Misotshi-Kabogo Highlands where

they represented 8/83 (9.6 %) of snap trap captures at

camp | (Table 1). Frequency of capture at Camp | was

7/783 (.009). They were readily caught with convention-

al break-back traps baited with peanut butter and oatmeal.

However, not a single specimen was caught at 1600 m and

1950 m (34 specimens captured during 1150 trap nights).

Five of the 8 specimens collected were adult, four of sev-

en sexable specimens were female, two of three adult fe-

males were pregnant, and male testes size averaged 7x3.5

mm.

Using the sub-species recognized by Hutterer (2005)

and the key of Dollman (1915a), these specimens need

comparison with C. olivieri kivu (Osgood, 1910) described

from Lake Kivu to the north, and C. olivieri zuleika (Doll-

man, 1915b) described from southern Zimbabwe. In his

key, and with access to the relevant type material at the

British Museum (Natural History), Dollman (1915a) dis-

tinguished these taxa based on dorsal pelage, the former

being a ‘dark chocolate brown’ while the latter is ‘a dull

coffee-brown’. Our series are more referable to the ‘dull

coffee’ type but the dorsal pelage has grey bases, tipped

with coffee brown. In any case, we realize that this char-

acter must be locally plastic and cannot have great taxo-

nomic weight. However, in his description, Dollman

(1915b) also distinguishes C. olivieri zuleika from C.

olivieri kivu based on the larger teeth of the former. How-

ever, the UTR of our series (adults only) averages 13.6

mm compared to 14.8 mm for the type of C. 0. zuleika.

Therefore, we are unable to associate this series with a cur-

rently recognized sub-species.

Bonn zoological Bulletin 62 (2): 203-219

Crocidura lwiroensis n. sp. Kerbis Peterhans & Hutter-

et:

(Fig. 2; Tables 3, 4); The Misotshi-Kabogo Crocidura.

Holotype. FMNH 195181, original number MHH 828

(Fig. 2). Adult female with swollen mammae (3+3). Ba-

sioccipital suture closed. Dentition with moderate wear.

Left leg (tibia) broken post-mortem. Prepared as an alco-

holic carcass with skull removed (asr). Captured in a ‘mu-

seum special’ mouse trap with peanut butter and oatmeal

as bait. Collected on 06 February, 2007 by B. Ndara.

Type Locality. Approximately 45 km NE of Kalemie and

3 km W of the village of Mizimu above the western shore

of Lake Tanganyika, Katanga Province, eastern Democrat-

ic Republic of Congo (29°16’22”E 5°28°45”S, 1250 m).

Etymology. The specific epithet honors the biological re-

search team based at Lwiro (Centre National de Recherche

Scientifique), the biological research station 25 km north

of Bukavu, South Kivu, eastern Democratic Republic of

Congo. This group has persisted in their academic 1n-

quiries despite difficult socio-economic conditions and

civil strife over the past 15 years. Their academic accom-

plishments have been inspiring and this represents but one

of the team’s impressive findings.

Diagnosis. Crocidura lwiroensis is distinguished from oth-

er members of the genus both externally and cranio-den-

tally. Externally, it features very small body small size (5.6

g) and a tail that is nearly naked with only 8-10 bristles

on the basal 20 %. Dark grey above, slightly lighter be-

low. Skull very small (CI=18.26 mm). Parastyle of P4

long, heavy and broad, projecting forward, and providing

a medial pocket for the large third unicuspid. Third uni-

cuspid large, 40 % larger than second; 30 % of its length

eclipsed by the upper P4.

Description. External measurements are presented in

Table |. Pelage dark grey in appearance. Dorsal fur short

(2.7 mm) with dark grey roots tipped with drab brown.

Occasional silver ticks to dorsum. Belly fur appearing

slightly more light grey as the tips are only faintly brown.

Six swollen inguinal teats. The tail is naked except for 8-

10 bristle hairs located on the bottom 20 % of the tail. Pes

sparsely haired with dark hairs, foot scales darker later-

ally giving an impression of a darker colour on outside

aspect. Foot 11.5 mm with claw, 10.2 mm without.

Cranio-dental measurements are presented in Table 2.

Maxilla short (UTR 7.9 mm) and broad (6.04 mm). In-

fra-orbital bridge narrow (0.69 mm; figured in Meester

1963). First upper incisor lightly built. Cingulum on up-

per unicuspids very heavy, especially on first and third.

The third upper unicuspid is exceptionally large, 25 %

larger than UP2, and projects posteriorly so that fully 1/3

OZFMK

210 Julian C. Kerbis Peterhans et al.

Fig. 2. Crocidura lwiroensis n. sp., holotype FMNH 195181 in dorsal, ventral, and lateral view. Scale is 5 mm.

Bonn zoological Bulletin 62 (2): 203-219 ©ZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands 211

of its length is overlapping with the medial aspect of the

upper P4. Parastyle of upper P4 prominent. Upper P4 fair-

ly weak and vacuous. Upper M3 modest in dimensions

(0.62 x 1.35 mm). Lower incisor with slight denticulation.

Lower M3 simple and without talonid.

Comparisons. The small size (CI <19 mm) and nearly

naked tail aligns the new species with a small subset of

members of the genus (e.g. C. niobe, C. pitmani) with tail

pilosity at or below 50 %. The new species resembles Cro-

cidura niobe due its small size, nearly naked tail and heavy

unicuspids with prominent cingulum. However, it is sig-

nificantly smaller in all cranio-dental dimensions than C.

niobe (CI 18.2 vs 20.0 mm). All maxillary unicuspids

more rectangular than those of C. niobe which are broad-

er than they are long. Upper P4 more slightly built than

the stocky form seen in C. niobe, especially the talonid.

Anterior palatal foramina positioned across from the an-

terior half of the M1 in the new form compared with their

positioning across the posterior half in C. niobe. Lower

incisor relatively smooth compared with the denticulate

appearance of C. niobe. C kivuana is also much larger

(CI>20 mm) and is void of tail bristles. The new species

is approached by Crocidura pitmani (Barclay, 1932) in

cranio-dental dimensions but C. /wiroensis has a more nar-

row braincase (GB: 3.92 vs. 4.4 mm), and a much larger

third unicuspid that overlaps with the upper P4. C. /wiroe-

nesis has shorter fur (2.7 vs. 4-5 mm) and a shorter and

virtually naked tail with very few bristle hairs (vs. hairs

over the basal 50 % in C. pitmani).

In cranial dimensions, Crocidura hildegardeae ssp. is

comparable. However, the new form has long bristle hairs

on only 20 % of its length, whereas in C. hildegardeae

this ranges from 50-75 %. The new species also differs

from it in its darker and more unicolored pelage above and

below, vs. the brown/grey bicoloured pattern of C. hilde-

gardeae. The skull in the new form is shorter and heav-

ier as reflected in the more rounded braincase (vs. the oval

form found in C. hildegardeae) and broader maxillary re-

gion (compared to the length of the upper tooth row). Up-

per P3 much larger compared to P2 in C. /wiroensis com-

pared to the subequal form in C. hildegardeae. One vari-

ety of C. hildegardeae (C. h. lutreola Heller, 1912) also

has a much larger upper third premolar but does not over-

lap greatly with the upper P4 as the new form. It is fur-

ther distinguished by its more bristled tail, narrower max-

illa and broader infra-orbital bridge. The last two unicus-

pids are larger and more elongate in C. /wiroensis than the

more squared/rounded form seen in C. hildegardeae. The

last upper unicuspid in C. /wiroensis protrudes posterior-

ly and contrasts the squared-off form in C. hildegardeae.

As noted above, the upper P3/P4 complex of C. hwiroen-

sis 1S unique among these small Crocidura ssp.

More distant comparisons include forms from West

Africa, Ethiopia, and northern Kenya. C. crossei from

Bonn zoological Bulletin 62 (2): 203-219

Nigeria has a longer (51 vs. 45 mm) and more bristled tail

(‘evenly scattered throughout except at the extreme tip’;

Thomas 1895). It is more unicolored, being slate grey

above and below (Hutterer & Happold 1983). It also has

a longer skull (19.3 vs. 18.26) and upper tooth row (8.3

vs. 7.91) whilst having a more narrow maxilla (5.6 vs.

6.04). The Ethiopian C. harrenna (Hutterer & Yalden,

1990, Hutterer, pers. obs.) is larger (mass 8.5 vs. 5.6 g)

with higher pilosity of the tail (80 % vs. 20 %), a longer

skull (19 vs. 18.26 mm), and a broader braincase (9.2 vs.

8.42 mm) while having a narrower maxilla (5.6 vs. 6.04

mm) (Hutterer & Yalden 1990). Likewise, C. phaeura Os-

good, 1936, also from Ethiopia, has a darker pelage (black-

ish brown to dark brown) with a longer foot (15 mm).

Cranio-dental measurements are longer than the new

species described here: CI (19.3—21.1 mm) and UTR of

8.8, with a longer upper M3 (0.81—0.88 mm). Finally, C.

macowi, known only from Mt. Nyiro in northern Kenya,

is more dark brown (and with more sharp contrast to the

grey of the belly). C. macowi also has a longer tail (up to

58 mm), longer skull (19.7 mm ), and a longer, yet more

narrow maxilla, and with a broader interorbital region (4.6

mm). The third upper unicuspid and its medial eclipse of

the parastyle of the upper P4 are again distinctive in the

new form.

Ecology and accompanying small mammal communi-

ty. The new species of Crocidura was captured along a

creek in primary forest at 1250 m. The small mammals

with which it was caught include the following: Crocidu-

ra olivieri (n = 7), Crocidura c.f. fuscomurina ssp. (n =

1), Lophuromys aquilus (n= 3), Hvbomys aff. univittatus

(n = 3), Hvlomyscus stella (n= 12), Pelomys minor (n =

1), Praomys jacksoni (n = 43), and Rattus rattus (n= 1).

Myosorex kabogoensis n. sp. Kerbis Peterhans & Hutte-

Ter

(Figs 3, 4; Tables 3, 4); The Misotshi-Kabogo Myosorex.

Holotype. FMNH 195079, original number MHH 840.

Adult male, testes not convoluted, 2 x 1 mm. Basioccip-

ital suture closed. Dentition with very little wear. Prepared

as a skin with skull and skeleton (ssk). Captured in a ‘mu-

seum special’ mouse trap with peanut butter and oatmeal

as bait. Collected on 15 February, 2007 by P. Kaleme.

Type locality. Mt. Misotshi area, 4 km SW of the village

of Talama, above the western shore of Lake Tanganyika,

South Kivu Province, eastern Democratic Republic of

Congo (29°4°49”E 4°59°29"S, 1950 m).

Etymology. Named for the montane forest in which it was

found.

©OZFMK

Julian C. Kerbis Peterhans et al.

Fig. 3. Myosorex kabogoensis n. sp., holotype FMNH 195071 in dorsal, ventral, and lateral view. Scale is 10 mm.

Bonn zoological Bulletin 62 (2): 203-219 OZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands

Fig. 4.

Table 3. External measurements of Misotshi-Kabogo Soricidae.

Myosorex kabogoensis n.sp., holotype FMNH 195071, view of palate and upper dentition. Scale is 5 mm.

FMNH Species Date Cond Sex Age TOT TL HF E WT _ Tail bristles

N=5 + Crocidura var. var. 2m Ad 201 74 20 11 40.5 56 %

olivieri 3f

195071 Crocidura c.f. Feb 4 2007 sko ? Ad 113 49 ll 8 48 na

fuscomurina

195181 Crocidura Feb 6 2007 asr F Ad 110 45 11.5 8 5.6 20 %

lwiroensis

195079 Myosorex Feb 15 2007 © ssk M Ad 117 28 14 5 12 0%

kabogoensis

195070 Crocidura c.f. Feb 15 2007 © ssk M Ad 167 95 15 11 7.8 0%

dolichura

* specimen eaten in trap, re-measured in Chicago

+ mean of the five adult specimens (195072, 195073, 195074, 195076, 195077: 2 m, 3 f)

Diagnosis. Among members of the genus, Myosorex kabo-

goensis is easily diagnosed both externally and cranio-den-

tally. Externally, it has an exceptionally short tail in rela-

tion to head and body length (28/89=31.5 %). The pelage

is very dark slate blue/grey above and below, the individ-

ual hairs unicolored to their base. Braincase broad but

short. The maxilla is particularly broad, both absolutely

Bonn zoological Bulletin 62 (2): 203-219

and in relation to the breadth of the braincase. Upper 4"

unicuspid very small (1/6 the size of the 3rd), lies within

the toothrow and is visible in lateral view. Upper 3" uni-

cuspid, *%4 the size of the 1“. Upper M3 proportionately

large. Lower P2 not visible in lateral view. Mandibular

corpus of moderate depth.

©ZFMK

214 Julian C. Kerbis Peterhans et al.

Table 4. Cranio-dental measurements of Misotshi-Kabogo Soricidae.

FMNH Species Cl GB

MX HCC PGL IO MD LTR COR

N=5 + Crocidura olivieri 29.8 12.1 9.6 13.6 6.7 8.4 53) 19.0 12.3 7.85

195071 Crocidura fuscomurina 17.3 7.6 5.32 7.37 4.1 5.0 3.63 10.22. 6.72 4.09

195181 Crocidura lwiroensis 18.26 8.42 6.04 7.91 4.75 6.09 3.92 11.11 7.31 4.32

195079 Myosorex kabogoensis 21.83 11.54 7.18 9.75 6.04 7.33 4.97 13.9 8.72 5.62

195070 Crocidura c.f. dolichura a 5.53 8.04 o 5.95 4.02 11.17 7.27 4.13

® skull broken

Description. Pelage dark slate grey, hairs unicolored to

their base. Color identical above and below. Tail very dark

above and below; extremely short (28 mm), only 31.5 %

of head and body length. Foreclaws long, longest 3.15 mm

(III), hindclaws shorter, longest 2.4 mm (III). Long dark

hairs (1.4 mm) clothing dorsum of pes and manus (see

Table 1).

Braincase broad but short (as reflected in the antero-lat-

eral length of the hexagon), fitting well within the range

of broad-headed Myosorex (GB/CI=>0.5, Table 2, see Ker-

bis Peterhans et al. (2010). Maxilla broad. Interorbital re-

gion bugling in ventral view. Narrow anterior opening (ca.

30 degrees) to the combined foramen opticum, rotundum

and lacerum anterius (Meester, 1963). The foramen mag-

num narrow and oval-shaped.

Upper unicuspids broad. Upper U4 lies within the tooth

row and is visible in lateral view (Fig. 4). Upper P4 heav-

ily built without expansive posterior concavity. Upper M3

large (long). Lower incisor with two clear denticulations.

Lower unicuspids large. Only one lower P2 present (or at

Table 5. External measurements of adult Rodentia.

least visible in medial view). Medial articular facet of the

mandibular condyle short and straight. Lower M3 long and

narrow.

Comparisons. Cranially, the new Myosorex aligns with

the broad-skulled forms of Myosorex (GB/CI=>0.5: M.

blarina, M. babaulti, M. bururiensis, M. geata, M. ki-

haulei, M. okuensis, M. rumpi, and M. zinki (see Kerbis

Peterhans et al., 2010). Its maxillary and braincase breadth

(GB/CI=.529) and broad inter-orbital region distinguish

it from the more narrow-headed forms (GB/CI<0.5: /.

cafer, M. eisentrauti, M. gnoskei, M. longicaudatus, M. je-

Jei, M. schalleri, M. sclateri, M. tenuis, M. varius).

The West African taxa, M. rumpi and M. okuensis are

both only known from a single plateau or mountain peak

2000 miles to the west. M. rumpi is substantially larger,

Cl>23.0 mm, and with a proportionately narrower max-

illa (max/GB: .587 vs. .622 in M. kabogoensis). M. okuen-

sis possesses a much narrower maxilla in absolute

breadth (6.72 mm vs. 7.18 mm). M. zinki has a much

Species FMNH Sex F, M Age HB TV HF WT

Graphiurus murinus 195179 F ya 95 70 18 19

Lophuromys aquilus means 7F, 2M ad 126 — 21 39

Hybomys c.f. univittatus means 2F, 4M ad 128.5 11] 29.7 54

Hylomyscus stella means 4F, SM ad 130 18 21

Mus minutoides 195116 F ad 71.5 40.5 12 9

Mus minutoides 195117 F yad

Mus triton 195118 F ad 88 55 14 20

Pelomys minor 195187 M ad 124 122 27 5]

Praomys degraaffi 195120 M ad 109 126 23.5 30.5

Praomys degraaffi 195122 F - 101 123 23 26

Praomys jacksoni means 3F yad 147.7 105.3 23.7 36.3

Praomys jacksoni means 7F ad 141 113.7 24 40.5

Praomys jacksoni means 4M yad 134 101 Su) 31

Praomys jacksoni means 6M ad 149 118 50.2

Rattus rattus 195177 M Juv 133 152 69

Atherurus africanus 195178 F ad 475 165 75 na

Bonn zoological Bulletin 62 (2): 203-219 OZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands

Table 6. Select cranio-dental measurements of adult Rodentia.

Species FMNH Sex KM Age

Graphiurus murinus 195179 F

Lophuromys aquilus means 3F

Hybomys c.f. univittatus means 3F, 4M

Hylomyscus stella means 3F, 5M

Mus minutoides 195116 F

Mus minutoides 195117 F

Mus triton 195118 F

Pelomys minor 195187 M

Praomys degraaffi 195120 M ya

Praomys degraaffi 195122 F ya

Praomys jacksoni means 3F ya

Praomys jacksoni means SF ad

Praomys jacksoni means 5M ya

Praomys jacksoni means 15M ad

Rattus rattus 195177 M

Atherurus africanus 195178 F

onger skull and narrower maxillae (22.8 mm and 6.5 mm

vs. 21.83 mm and 7.18 mm) and is only known from Mt.

Kilimanjaro.

Although this species is exceeded in breadth by sever-

al others, including neighboring M. bururiensis, M. bla-

rina and M. babaulti, it has the hexagonal ‘gestalt’ and

metrics of the more broad-skulled group. The braincase

is short and smaller overall than these three. In ventral

view, the interorbital region is convex and bulging com-

pared to the parallel-sided or often concave-sided M.

babaulti, M. blarina and M. bururiensis. Narrow anteri-

or opening (ca. 30 degrees) to the combined foramen op-

ticum, rotundum and lacerum anterius (Meester 1963)

compared with ca. 40-45 degrees in M. babaulti. Only

slightly narrower than that seen in M. bururiensis. The

foramen magnum is more narrow and oval-shaped than

either.

Upper unicuspids less broad than those seen in M. bu-

ruriensis but similar in their proportions. The breadth and

size of the upper unicuspids are comparable to those of

M. babaulti and M. blarina but unlike M. babaulti, the up-

per 4" unicuspid is visible in lateral view. This is because

it lies within the tooth row and is not medially displaced

and to a lesser extent, the parastyle of the following up-

per P4 does not protrude forward. Upper M3 much larg-

er and longer than that of M. blarina.

The mandibular corpus, articular facet of the mandibu-

lar condyle and the lower unicuspids of M. kabogoensis

are much less robust than in M. bururiensis. The lower

unicuspids are comparable in size to M. babaulti. The low-

er M3 is clearly more narrow, especially the talonid. The

medial facet of the mandibular condyle is very short and

Bonn zoological Bulletin 62 (2): 203-219

Cl IO BC UTR

23.6 4.4 13.2 3.1

28.5 6.3 12.8 5.0

295 5-7 14.3 522

23.6 4.6 11.4 3.8

17:5 3.4 8.8 3

17.6 Ce, 8.7 3.2

20.9 3.9 9.9 3.5

29.0 4.6 [3:5 533

2125 4.75 12.6 4.95

25.8 4.9 11.9 4.45

Died 4.8 12.1 4.5

27.9 4.7 12.2 4.8

26.4 4.7 11.7 4.6

29.2 5 12.4 4.8

32.1 4.9 15.2 6.7

86.2 24.8 32.3 17.8

straight as opposed to the longer and more concave con-

dition seen in M. babaulti.

The dark slate-grey of the pelage is unicolored and with-

out the tricolored mottling seen in certain members of the

genus (e.g. M. gnoskei, M. geata (?) and the S African M.

cafer and M. varius). This also contrasts with the bicol-

ored pelage (basal 80 % slate grey, distal 20 % dark

brown) seen in the more northern forms (M. blarina, M.

bururiensis and M. babaulti). Externally, its short tail, in

relation to head and body length is met only by M. bu-

ruriensis (Table 2 in Hutterer et al. 2001; Table 2).

Ecology and the small mammal community. The new

species was captured along a dry hill slope in primary for-

est where a few large trees emerge from dense and shrub-

by undergrowth (1950-2000 m). It had rained the previ-

ous early morning, ca 24 hours prior to capture. Rodents

collected in the same general area include Graphiurus

murinus (n= 1), Hybomys univittatus (n = 6), Praomys

degraaffi (n = 2), and Praomys jacksoni (n = 1).

RODENTIA

Funisciurus carruthersi chrysippus. Our observations

and Prigogine (1960) mark the southernmost occurrence

known for this Albertine Rift endemic.

Heliosciurus ruwenzorii vulcanius. The record from Pri-

gogine (1960) marks the most southerly record known for

this Albertine Rift endemic.

©ZFMK

216

Protoxerus stangeri kabobo. This giant squirrel is wide-

spread in tropical Africa and is an indicator of good for-

est. This subspecies is recognized as distinct and is con-

fined to the Kabogo-Misotshi highlands.

Graphiurus murinus (Desmarest, 1822). The sole spec-

imen (FMNH 195179), a young-adult with first premolar

in early wear, is a nulliparous female and was prepared

as a skin with skull and skeleton. We refer this species to

Graphiurus murinus due to its size (external and cranio-

dental), dark coloration towards the rear of the manus and

pes and the lack of a white tip to the tail.

Lophuromys aquilus True, 1892. In addition to Praomys

jJacksoni, members of this species complex are typically

the most abundant taxon in east African forests. They were

not a dominant component of the lower two camps

(7/94=7.4 %) and were not documented at 1950 m. Nine

of the ten captures were adult. The sex ratio was seven

females and three males. Three of the five females inspect-

ed were pregnant (one each with two, three and four em-

bryos) with CR ranging from 5—13mm. Testes from a sin-

gle male measured 7 x 12 mm.

Hybomys atf. univittatus (Peters, 1876). As is typical for

this genus, several specimens were captured near streams

within forest. Six of the nine captures were adult. The

overall sex ratio was four females and five males. Two of

the four females inspected were pregnant (one with two

embryos, the other with three) with CR ranging from 8—20.

Testes of adult males ranged from 7 x 13 mm to 10 x 16

mm.

These specimens are more comparable to Hybomys uni-

vittatus than to Hybomys lunaris, the only two taxa allud-

ed to in eastern DR Congo. As discussed in the original

description (Thomas 1906) and more recently by Muss-

er & Carleton (2005), H. /unaris 1s lightly built compared

to H. univittatus. Comparisons of true Hybomys univitta-

tus with topotypes of Hybomys lunaris at Field Museum

confirm this. The greatest skull length (33.0—35.1 mm) of

the Kabogo series falls within the range of H. univittatus

(33.5—35.6 mm = | standard deviation). These are signif-

icantly larger than the 1. /unaris series at Field Museum

(31.25—32.48 mm). The dentition of the Kabogo series 1s

intermediate: UTR (5.05—-5.35 mm) and M' breadth

(1.73-1.85 mm) compared with Hybomys lunaris

(4.74—-5.08 mm and 1.6)—1.7 mm) and Hybomys univit-

tatus (5.66-6.1 mm and 1.87—1.99 mm = | standard de-

viation).

Hylomyscus stella Thomas, 1911. This species is typical

in lowland forest of the eastern Congo basin. Ten of the

12 aged specimens were adult. The sex ratio was seven

females and six males. One of the four females inspect-

ed was pregnant with two embryos, with a CR of 3 mm.

Bonn zoological Bulletin 62 (2): 203-219

Julian C. Kerbis Peterhans et al.

Testes of adult males measured 6 x 12 mm. The absence

of members of the Hylomyscus denniae group (Carleton

et al. 2006) in an Albertine Rift montane context is curi-

ous. Evidently, Hylomyscus vulcanorum reaches its cur-

rent southern extent on the Itombwe Plateau, just to the

north (Demos, unpublished data). Since these highlands

extend beyond 2700 m, we cannot explain that members

of the Hylomyscus denniae complex were victims of the

last interglacial, ca. 12,000 years before present.

Mus musculoides Temminck, 1853. Two specimens were

captured in tall grass near the village of Mizumu on the

shore of Lake Tanganyika. One female was not pregnant

but displayed six large teats while the other was pregnant

with two embryos (CR 12x17).

Mus triton Thomas, 1909. One specimen caught in tall

grass near the village of Mizumu on the shore of Lake Tan-

ganyika. The adult female, multiparous with six teats, was

pregnant with three embryos with a CR of 20 mm.

Pelomys minor Cabrera & Ruxton, 1926. A single adult,

scrotal male (FMNH 195187) was taken at the forest edge

along the ridge slope.

Praomys degraaffi Van der Straeten & Kerbis Peterhans,

1999. This is the second published record for this Alber-

tine Rift endemic in DRC. FMNH has many specimens

of this taxon from Mt. Tshiabirimu, Kahuzi-Biega NP, and

the Itombwe Forest. The female, FMNH 195122, displays

the charcoal pelage and 8 teats characteristic of the tax-

on while the male, FMNH 195121 shows the charcoal

pelage and long and broad incisive foramina characteris-

tic of the species. Notably, both are recorded at the high-

est camp (1950m) whereas 62/63 members of Praomys

Jacksoni were found at lower elevations (Table 1). This

pattern of altitudinal displacement was discussed in the

original description of the species (Van der Straeten &

Kerbis Peterhans 1999).

Praomys jacksoni de Winton, 1897. As is typical in much

of forested eastern Africa, this is by far the most abundant

murid (63/104=61 %) from the surveys. It comprised over

50 % of all captures except for the highest camp where

its frequency dropped to 10 % (1/10). It is distinguished

from the previous species by its brown vs. black pelage,

additional accessory plantar tubercles, and shorter, more

narrow incisive foramina. The overall sex ratio for this

species was 26 f to 34 m. Forty one out of 61 specimens

were adult. Juvenile sex distribution was skewed towards

females (13/20). However, this seemingly reversed in

adulthood with adult males composing 27 of 40 individ-

uals. Testes size in adult males averaged 9x15 (n=19).

Three of seven adult females inspected were pregnant with

embryos numbering 2 (CR 20), 3 (CR 20) and 4 (CR 14).

OZFMK

Mammals, other than bats, from the Misotshi-Kabogo highlands 217

There appears to be no evidence of sexual dimorphism

(Table 6).

Rattus rattus Linnaeus, 1758. The recovery of this spec-

imen (FMNH 195177) is surprising as a forest find. An

adult scrotal male with testes 13x7 mm was caught in the

same trap line as Crocidura lwiroensis.

Atherurus africanus. One female (FMNH 195178), uterus

enlarged, lactating from two axial mammae, was snared

by a camp attendant in primary forest.

DISCUSSION AND CONCLUSION

These discoveries, over the course of a very short period

in the field (2 % weeks), highlight the importance of sur-

veying undocumented African habitats, particularly iso-

lated montane habitats within the Albertine Rift. The Mis-

otshi-Kabogo highlands represent a particularly important

montane forest block as it lies to the south of the main

Kivu Highlands (Kahuzi-Itombwe) and 1s thought to have

been long-isolated.

The two new species described here and a new horse-

shoe bat (Kerbis Peterhans et al. 2013) bring to five the

number of unique mammal taxa known from these high-

lands. It is probable that further new species will be dis-

covered here given the brevity of the 2007 field survey.

Further, the one unique primate (Co/obus angolensis pri-

goginei) is severely threatened with extinction despite a

healthy forest cover. A total of 11 mammal species reach

their southern range terminus on the Misotshi-Kabogo

Highlands (Tables | and 2). These include such remark-

able members of the megafauna as the giant forest hog

(Hylochoerus meinertzhageni) and the bongo (7ragela-

phus euryceros). The next montane system to the south

in the Albertine Rift chain is the Marungu Highlands. Aer-

ial reconnaissance by our team indicated that there is no

remaining forest block here. We cannot claim that the bar-

ren Marungu Highlands have been deforested by anthro-

pogenic activity or whether it has been in this state

throughout historical times. What is clear is that the Mis-

otshi-Kabogo Highlands are a critical montane system at

the SE corner of the Congo forest block and provides the

southern limit for many endemic Albertine Rift taxa. At

present this montane forest has no formal protection; there

is aneed to establish a protected area here if these recent-

ly discovered new species are to be maintained.

These new discoveries indicate how much there is yet

to learn of African biodiversity, especially within montane

environments. Montane communities contain reservoirs of

biodiversity that are inherently isolated. Their exploration

must be placed at the forefront of survey and conserva-

tion efforts. Only 13 years ago (Myers et al. 2000), it was

claimed that the Albertine Rift did not warrant recogni-

Bonn zoological Bulletin 62 (2): 203-219

tion as an international Hot Spot due to inadequate knowl-

edge. We have started filling in this void by describing 8

small mammal species from the Albertine Rift since 1999

that are new to science (Van der Straeten & Kerbis Peter-

hans 1999; Kerbis Peterhans & Hutterer 2009; Kerbis Pe-

terhans et al. 2010; Kerbis Peterhans et al. 2013, this pa-

per). Continued field surveys and collections, such as the

one reported on here, are crucial to this effort. Interest

from non-government organizations (NGO’s) has provid-

ed various groups with the resources to begin surveys of

the unexplored pockets within the ecoregion. Discover-

ies of rare or species new to science have the potential to

serve as flagship species for areas that lack formal pro-

tection. Finally, the fairly rapid turnover from field sur-

vey to published manuscript further illustrates that the oft-

ignored small mammal community should be fundamen-

tal in biodiversity surveys.

Acknowledgements. The expedition to the Misotshi-Kabogo

Highlands was initiated by the Wildlife Conservation Society

(WCS). Field work was spear-headed by AP in conjunction with

the Research Staff at Lwiro (CRSN, Democratic Republic of

Congo), The Field Museum of Natural History (Chicago) and

the World Wide Fund for Nature (WWF). Additional funding

came from the Daniel K. Thorne Foundation, the John D. and

Catherine T. MacArthur Foundation, the Barbara Brown Fund

(Field Museum), and the US Fish and Wildlife Service. In the

Misotshi-Kabogo Highlands, we acknowledge D. Kujirakwin-

ja and J. Badesire for logistical support. E. A. Mulungu provid-

ed critical field support and G. Lukole and G. M. Mwankasala

provided security and field logistics and local introductions. We

thank J. Mwanga (Head, Biology) and B. Bajope (Director) of

the Centre de Recherche en Sciences Naturelles at Lwiro for con-

tinuous assistance in collecting and with export permits. ECHO

provided air transport. The senior author acknowledges the Bar-

bara Brown Fund and the African Council of the Field Muse-

um for financial support and an award from Roosevelt Univer-

sity (Dean’s Office) enabled JCKP to work on this publication

in Germany with RH. We thank L. Gordon and M. Carleton of

the United States National Museum (USNM) for access and

loans of specimens in their care. We recognize FMNH staff for

their assistance: M. Schulenberg, J Phelps, and WT Stanley. D.

Rohwedder (ZFMK) made the photographs and U. Vaartjes for-

matted the plates. SO. Bober (FMNH) prepared the maps.

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Mammals, other than bats, from the Misotshi-Kabogo highlands 219

Appendix

Specimens examined for comparison

Dendrohyrax dorsalis sylvestris (Temminck, 1853). Gabon (5):

FMNH 54448, 54679-681, 62767.

Crocidura dolichura (Peters, 1876). Gabon (1): FMNH 162198.

Crocidura fuscomurina (Heuglin, 1865). Angola, Humpata,

ZFMK 89.48; Kenya, Nairobi, Embakasi Plains, ZFMK 85.65;

FMNH 17719, 17723; Sudan, FMNH 86028.

Crocidura hildegardeae altae (Heller, 1912). Kenya: Mt Gar-

guez, North Creek, 6000’-6500’, USNM 182426, 182429,

182430.

Crocidura hildegardeae hildegardeae (Thomas, 1904). Demo-

cratic Republic of Congo: Mbayu, ZFMK 68.505, Buhenge,

ZFMK 89-450, Bukarabwa, ZFMK 89-455. Kenya: Fort Hall,

USNM 163948. Wambugu, USNM 163952. Nyeri, USNM

163953-955.

Crocidura hildegardeae lutreola (Heller, 1910). Kenya: Mt Mbo-

lolo, summit, USNM 182456, 182459, 182460.

Crocidura hildegardeae phaios (Setzer, 1956). Sudan: Torit,

2300’, FMNH 66712.

Crocidura hildegardeae procera (Heller, 1912). Kenya: Mt. Gar-

guez, south creek 3600’, USNM 182509, FMNH 47357. Mt.

Lololokwi, summit, 6000’, USNM 182512, 182513, 182517,

182518; FMNH 47358. Mt. Lololokwi, 182522.

Crocidura niobe (Thomas, 1906). Uganda (5): Ruwenzori Moun-

tains, 6200’, FMNH 144181, 144183, 144185, 144190; Mt.

Sabinio, FMNH 26477.

Bonn zoological Bulletin 62 (2): 203-219

Myosorex babaulti (Heim de Balsac and Lamotte, 1956): Bu-

rundi, Kibira NP, FMNH 148937 1M, FMNH 148938 1M,

FMNH 148265 1M, 2100-2350 m; DR CONGO, Kahuzi-Bie-

ga NP, Tshibati, FMNH 189275 1M, ZFMK 68.545, IF, Lwiro

Falls, 2100 m, AMNH 180956—180961, 6M; UGANDA,

Bwindi-Impenetrable NP, 1850 m, FMNH 160175 1M,

Mgahinga Gorilla NP, 2980 m, FMNH 157410 1M.

Myosorex blarina (Thomas, 1906): DR CONGO, Rwenzori

Mountains, FMNH 26285-26287 2M, IF; UGANDA, Rwen-

zori Mountains NP, FMNH 144205—144211, 2M, 4F, 1?,

1900-3980 m.

Myosorex bururiensis (Kerbis Peterhans & Hutterer, 2010): Bu-

rundi, Mumushwizi Valley, Bururi Forest, 1880m, FMNH

155923 F.

Myosorex geata (Allen and Loveridge, 1927): Tanzania, Ulugu-

ru Mountains, Uluguru North Forest Reserve 1345-1535 m,

FMNH 158298-158302, FMNH 158487 3M, 3F, Mbete, TRP

2305.

Myosorex gnoskei (Kerbis Peterhans et al., 2008): FMNH

191568, Malawi, Chilinda Rest Camp, Nyika National Park,

2285m.

Myosorex kihaulei (Stanley & Hutterer, 2000): Tanzania,

Udzungwa Mountains, New Dabaga/Ulangambi Forest Re-

serve, 1816-1940 m, FMNH 169509-169516, 1?, 7F; West

Kilombero Scarp Forest Reserve, 1140 m, FMNH 169501, 1?.

Hybomys lunaris (Thomas, 1906): Uganda, Ruwenzori Moun-

tains, FMNH 144400, 144405, 144408, 144412, 144429,

144430, 144433, 144444, 4M, 4F.

©ZFMK

Bonn zoological Bulletin 62 (2): 220-252

On the Linck collection and specimens of snakes

figured by Johann Jakob Scheuchzer (1735) —

the oldest fluid-preserved herpetological collection in the world?

Aaron M. Bauer' & Richard Wahlgren’

‘Department of Biology, Villanova University, 800 Lancaster Avenue

Villanova, Pennsylvania 19085, U.S.A.

°Prennegatan 23B, SE-223 53 Lund, Sweden.

‘Corresponding author: E-mail: aaron.bauer@villanova.edu, telephone: + 1-610-519-4857

Abstract. One of the great private natural history cabinets of the 18" century was that of the Linck family of Leipzig

pharmacists. Parts of the collection have survived to the present and form the core of the Naturalienkabinett Waldenburg

in Saxony, Germany. The collection was particularly rich in reptiles and was documented by Johann Jakob Scheuchzer

in his Physica Sacra (1731-1735), which figured 67 specimens of snakes and amphisbaenians based on a set of unpu-

blished illustrations, the Jcones Serpentum et Viperarum, prepared under the direction of Johann Heinrich Linck the El-

der. We review the original herpetological content of the Linck collection as documented by Johann Heinrich Linck the

Younger in his Index Musae Linckiani (1783-1787) and provide both a summary of earlier identifications (to 1858) of

the species depicted in the /cones and Physica Sacra and new identifications based on our research. Some of these sna-

kes served as holotypes or syntypes of species described by Linnaeus and Blasius Merrem and, thus, are of taxonomic

significance. As many as I|1 of these illustrated specimens (although none of them types), and an unknown number of

others, are still extant in Waldenburg. At a minimum, these specimens were present in the Linck collection in 1729, but

they may be as much as half a century older, as the reptile collection was already large and well-known by the debut of

the century. Even at the minimum age possible, the surviving Linck snakes figured by Scheuchzer are among the oldest

documented fluid-preserved herpetological specimens in the world.

Key words. Johann Heinrich Linck, Naturalienkabinett Waldenburg, Physica Sacra, Johann Jakob Scheuchzer, Serpen-

December 2013

tes, historic collections.

INTRODUCTION

Natural history cabinets, either self-standing, or as part of

more inclusive curiosity cabinets or Kunstkammer, were

a well-established feature of educated society throughout

much of Europe by the late 16" century and flowered in

the 17" century as the expansion of global trade and trav-

el provided ever more subjects for collection and study

(Seifert 1935; MacGregor 2007). Although many of the

most sumptuous cabinets were owned by royal families

and other aristocrats, there were also a great many schol-

arly and private cabinets, many of which were owned by

physicians and apothecaries for whom objects of natural

history were of professional as well as personal interest.

Perhaps the most famous of the early private natural his-

tory cabinets was that of Ulisse Aldrovandi (1522—1605),

today represented by a variety of dried specimens and an

extensive “paper museum” of contemporary paintings of

natural objects (Alessandrini & Ceregato 2007; Bauer et

al. 2013). Many others survive only in the form of pub-

lished catalogues and iconic images, for example the mu-

seums of Francesco Calzolari (1521-1600; Ceruti &

Chiocco 1622), Ferrante Imperato (1550-1625; Impera-

to 1599), Basilius Besler (1561—1629; Besler 1616, 1622,

Received: 02.05.2013

Accepted: 19.11.2013

1642), and Ole Worm (1588-1654; Worm 1655, see also

Schepelern 1987, 1990). Not until the late 17" century

when William Croone [or Croune] (1633-1684) and

Robert Boyle (1627-1691) began experiments involving

the alcohol preservation of organic specimens (Croune

1662 in Birch 1756; Boyle 1663 in Birch 1756; Boyle

1666) did fluid-preserved specimens become a hallmark

of such collections.

Many of the spirit-preserved collections from the first

third of the 18" century or earlier featured human anatom-

ical preparations. Some still survive, including specimens

from the collections of Bernhard Siegfried Albinus

(1697-1770) in Leiden (Habrich 1994) and Frederik

Ruysch (1638-1731), who sold his famous collection of

anatomical specimens to Peter the Great in 1717

(Luyendijk-Elshout 1994; Driessen-van het Reve 2006).

The latter included many skeletal preparations as well as

dry injected mounts, none of which appear to have sur-

vived, but also fluid preserved material, some of which

is still extant in St. Petersburg (Driessen-van het Reve

2006).

Corresponding editor: Ph. Wagner

The Historic Linck Collection of Snakes 221

Fig. 1.

Stuffed anaconda (Eunectes murinus) from the first collection of Albertus Seba on display in the Zoological Institute of

the Russian Academy of Sciences, St. Petersburg. The head of the specimen was missing and a new “head” fashioned from the

front of the skin. Photo courtesy of Daniel A. Melnikov.

Zoological collections were also popular in natural his-

tory cabinets of this period and these often contained large

numbers of fluid-preserved amphibians and reptiles

Fig. 2.

Suspended stuffed crocodile (Crocodylus niloticus)

mount and (below, in cabinets) spirit-preserved collection of am-

phibians, reptiles and other material in the Franckesche Stiftun-

gen in Halle. The painted cabinet fronts are modern but repro-

duce historical designs. Photo: A. M. Bauer.

Bonn zoological Bulletin 62 (2): 220-252

(George 1987). Mostly obtained from the tropics, these

were generally small, easy to store and display, and attrac-

tive to collectors in temperate Europe because of their ex-

otic appearance as well their real and imagined medical

significance. Perhaps the most well-known spirit-pre-

served herpetological collection of the early 18" century

was that of Albertus Seba (1665—1736) who sold his first

collection to Peter the Great in 1716 (Engel 1937;

Driessen-van het Reve 2006) and then built an even larg-

er second collection. All herpetological components of the

first collection, except a stuffed anaconda (Fig. 1) and per-

haps a few other dry preparations appear to have been lost

(Juriev 1981), but remnants of the second collection are

known to exist in a variety of museums, having been dis-

persed through a series of auctions following Seba’s death

(Boeseman 1970; Juriev 1981; Thireau et al. 1998; Bauer

2002; Daszkiewicz & Bauer 2006; Driessen-van het Reve

2006; Bauer & Gtinther 2013).

Older still is the collection of the Franckesche Stiftun-

gen in Halle, established in 1698 as part of the orphan-

age and educational complex founded by August Hermann

Francke (1663-1727) (Kohler 1799; Sauerlandt 1911;

Storz 1962; Jahn 1994; Miiller-Bahlke & Géltz 1998,

Miiller-Bahlke 2004). Today this collection includes 28

©ZFMK

222 Aaron M. Bauer & Richard Wahlgren

[b]

Johann Heinrich Linck der Jiingere (1734—1807) Ce» ctor ae

ie a Prwe «

(Stich von G. G. Endner nach einem Gemdlde von Knorr) ”

[c] [d]

Plate 1. a. Heinrich Linck (1638-1717), founder of the Linck collection and of the Linck apothecary dynasty in Leipzig. Image

from Seifert (1934) from an engraving by Nicolaus Haublein. b. Johann Heinrich Linck the Elder (1674-1734), son of Heinrich

Linck and accomplished naturalist and pharmacist. Image courtesy of U. Budig, Museum Waldenburg — Naturalienkabinett und

Stadtmuseum. ¢. Johann Heinrich Linck the Younger (1734-1807), grandson of H. Linck and son of J. H. Linck the Elder, who

prepared the Index Musae Linckiani. Image from Seifert (1934) from an engraving by G. G. Endner. d. Fiirst Otto Victor I von

Schénburg-Waldenburg (1785-1859), who purchased the remaining portions of the Linck collection in 1839, establishing it in its

existing building in 1844. Image courtesy of U. Budig, Museum Waldenburg — Naturalienkabinett und Stadtmuseum.

Bonn zoological Bulletin 62 (2): 220-252 OZFMK

The Historic Linck Collection of Snakes 223

Fig. 3. Exterior of the Museum Waldenburg Naturalienkabi-

nett und Heimatmuseum in 2005. Photo: A. M. Bauer.

early 18" century alcohol preparations consisting chiefly

of amphibians and reptiles as well as embryonic materi-

al (Altner 1984; Fig. 2). However, perhaps the most im-

portant intact herpetological collection from the dawn of

fluid preservation is that established by the pharmacist

Heinrich Linck (1638-1717; Plate la) in Leipzig, Ger-

many. Linck was one of many European apothecaries who

assembled such collections during the 17" century (Dilg

1994), but his collection, augmented by his son and grand-

son, is one of few that has survived — in part — the vagaries

of more than 300 years of turbulent European history. To-

day the collection forms part of the Museum Waldenburg

(Fig. 3) in the small Saxon town of Waldenburg, 67 km

south of Leipzig, Germany and is regarded as one of the

very oldest surviving museum collections of this kind

(Mohr 1940; Fleck et al. 1990; Zinke 1999).

No records exist documenting the oldest parts of the col-

lection and thus the unambiguous identification of any por-

tions of the collection dating from Heinrich Linck’s time

is not possible. Nonetheless, the collection includes a num-

ber of distinctive specimens of various kinds that date from

the first decades following Heinrich Linck’s death. For ex-

ample, the famous “chicken man” (Hiihnermensch), a mal-

formed human foetus, was described and figured in 1737

by Gottlieb Friderici and is still extant today (Miller

1999). Other specimens from these early days that are still

present in the collection include a number of distinctive

fossils (ROBler 1999a, 1999b), among them examples of

“lying stones,” crude fossil forgeries described by

Beringer (1726), raw and worked mineral samples (Thal-

heim 1999), a collection of wood samples (Beyrich 1990;

Bonn zoological Bulletin 62 (2): 220-252

Otto & Otto 1999), and the dried echinoderms that were

figured by Heinrich Linck’s son, Johann Heinrich Linck

the Elder, in his treatise De Stellis Marinus (1733a,

1733b). The collection at Waldenburg also includes a num-

ber of Linck birds from the 18" century (Steinheimer

2005).

On September 25—26, 2005 and again on June 22-23,

2009 we visited the Naturalienkabinett Waldenburg in or-

der to examine its herpetological collections and to deter-

mine if any of the specimens therein could be confirmed

as having survived from the early 18" century portion of

the collection. Whether any material dates to the 17" cen-

tury founding collection by Heinrich Linck is unknown,

and probably unknowable, but the accurate illustration of

some of the specimens of the Linck collection before 1730

(Scheuchzer 1735) provides a benchmark for establishing

a minimum age of any illustrated specimens that still sur-

vive. Such specimens could, however, be as much as 60

years older. Thus, it is possible that any such extant spec-

imens might represent the oldest fluid-preserved herpeto-

logical specimens in the world.

BUILDING OF THE LINCK COLLECTION

Heinrich Linck was born in 1638 in Danzig, a center of

natural history collections and later home to Jakob

Theodor Klein and other notable naturalists and collec-

tors (Friedrich 2001; Daszkiewicz & Bauer 2006). He

came to Leipzig in 1669 and first managed and then (1671)

leased the pharmacy “Zum Goldenen Lowen”, becoming

its owner in 1686 (Bormann 1909; Seifert 1934, 1935;

Beyrich 1994; Budig 1999b; Friedrich 2001). The phar-

macy, as the L6wenapotheke, is still in business in its orig-

inal position on Grimmaische Strae (Fig. 4) and in 2009

SSS,

A APOTHEKE 3

Fig. 4.

has functioned as a pharmacy for over 600 years and was first

managed by Heinrich Linck in 1669 under the name “Zum Gold-

enen Lowen.” Photo: A. M. Bauer.

The Lowenapotheke, on Grimmaische Strabe, Leipzig

©ZFMK

224 Aaron M. Bauer & Richard Wahlgren

Fig. 5. — View of the Green Room, housing the bulk of the her-

petological collections, on the first floor of the Museum —

Naturalienkabinett Waldenburg. Photo: A. M. Bauer.

celebrated its 600" anniversary. Linck began collecting

natural history specimens around 1671 or 1672 (Bormann

1909), although the details of the foundation of his natu-

ral history cabinet are largely unknown (Beyrich 1994;

Budig 1999b), and continued to enlarge its holdings un-

til his death in 1717. The collection became a family pas-

sion and the collection was enlarged by Heinrich’s son,

Johann Heinrich Linck the Elder (1674-1734; Plate 1b)

and later by his grandson, Johann Heinrich Linck the

Younger (1734-1807; Plate Ic).

Johann Heinrich the Elder established an extensive trad-

ing network with other collectors and also purchased ma-

terial from all over the world. During his travels across

Europe, Linck visited the important collections of Seba

in Amsterdam, Klein in Danzig, and Sloane in London,

amongst others (Seifert 1934, 1935; Beyrich 1994; Budig

1999b). As well as being a collector, Linck was a scien-

tist himself and published an important work on starfish

and related echinoderms (Linck 1733a, 1733b) and con-

ducted a variety of entomological experiments (Seifert

1934; Beyrich 1994). He was also a member of the Kaiser-

lich Leopoldinisch-Carolinische Akademie der Natur-

forscher, and a corresponding member of the Accademia

delle Scienze dell’ Instituto di Bologna and the Royal So-

ciety of London (Lingke & Lingke 1909; Beyrich 1994;

Budig 2007). After his death the collection languished for

a time until his son came of age and took over both the

pharmacy and the collection in 1757.

Johann Heinrich Linck the Younger subsequently en-

riched the collections by the purchase of the large natu-

ral history collection of Johann Christoph Richter

(1689-1751) in 1784 (Beyrich 1994). Most importantly

he reorganized the collection, largely after Linnaeus’s Sys-

tema Naturae and published an extensive catalogue (Linck

1783-1787) listing the material then present, which in-

cluded approximately 3,400 zoological items (Budig

1999b).

Bonn zoological Bulletin 62 (2): 220-252

The Linck collection was already well known for its size

and diversity by the early 18" century (Kanold in Neick-

elius 1727) and for many years thereafter it was noted as

one of the treasures of Leipzig (Schulz 1784; Leonhardi

1799, Klemm 1838) and was visited by important zoolo-

gists, including Marcus Elieser Bloch (1723-1799), Jo-

han Christian Fabricius (1745-1808), and Lorenz Oken

(1779-1851) (Mohr 1940). Linck was honored by Johann

Gottlob Schneider (1750-1822) who dedicated his Am-

phibiorum Physiologiae, Specimen Alterum (1792, 1797)

“ad virum doctissimum Iohann. Henricum Linck Serenis-

simo Saxoniae Electori a Consiliis Commerciorum Acad-

emiae Caesareae Naturae Curiosorum Socium et cet.” and,

indeed, the last four pages of the work extol Linck’s virtues

and the value of his collection.

Following the death, in 1807, of Johann Heinrich Linck

the Younger, his widow Dorothea Linck leased the phar-

macy, eventually selling it to Karl Heinrich August Rhode

in 1818. Upon her death in 1827, the collection was also

purchased by Rhode. Rhode sold the fabulous library (doc-

umented by Linck 1787 and today represented by at least

some volumes in the Universitatsbibliothek Leipzig) sep-

arately as well as some of the spirit-preserved specimens

— including snakes and amphibians, but subsequently sold

the remaining collection to Fiirst Otto Victor I von Schén-

burg-Waldenburg (1785-1859; Plate 1d) in 1839 for 3,300

Thaler (Seifert 1934; Mohr 1940; Fleck et al. 1990;

Beyrich 1994; Budig 1999b; Zinke 1999).

Otto Victor received the material in Waldenburg the fol-

lowing year (Beyrich 1990) and in 1844 moved it into a

newly built museum (Budig 1999a; Fig. 3), where it is still

housed today. Also in 1840, Otto Victor purchased sever-

al other significant natural history collections: the herbar-

ium, mineralogical and geological collection of Dr. Karl

Ferdinand Reichel (1800—1860), a pharmacist from Ho-

henstein (today Hohenstein-Ernstthal), an entomological

collection from Karl Gerhardt, and an important collec-

tion of birds and other animals from Karl Ferdinand Ober-

lander (1805-1866) of Greiz, a baker and confectioner as

well as an accomplished ornithologist (Heyder 1935; Mohr

1941; Beyrich 1990). The last significant addition to the

collection was derived from the African hunting expedi-

tion of Otto Victor I (1882-1914) in 1908-1910, consist-

ing chiefly of large mammals and birds.

On 26 October 1928 the Fiirstlich Sch6nburg-Walden-

burg’scher Familienverein “SchloB Waldenburg” was

founded and subsequently administered the collection.

During the period from early 1933 to | July 1934 the nat-

ural history collection was closed due to major renova-

tions and reorganization of the collection motivated by

First Giinther von Schénburg-Waldenburg (Seifert 1935;

Fischer 1936). On 25 October 1945 control of the collec-

tion passed to the Landesverwaltung Sachsen and in 1951

responsibility was transferred to the town of Waldenburg,

which continues as the custodian of the collection. Today

©OZFMK

The Historic Linck Collection of Snakes

the Naturalienkabinett is housed in the upper floor of the

museum building (Fig. 5), whereas the Stadtmuseum (for-

merly Heimatmuseum) occupies the ground floor.

Its purchase and move to Waldenburg account for the

survival of the Linck collection in World War II, when oth-

er collections in Leipzig were damaged or destroyed by

allied bombing (3-4 December 1943). However, the amal-

gamation of several collections by First Otto Victor I, as

well as his subsequent additions to the collection through

the early 20" century clouded the identity of the original

Linck collection. Nonetheless, it has been argued that his-

torical museum material can be reasonably identified on

the basis of intrinsic evidence in the absence of preserved

labels or other documentation (Faxon 1915) and we em-

ploy this approach to identify some of the oldest extant

specimens in the Linck collection.

HERPETOLOGICAL SPECIMENS IN THE LINCK

COLLECTION

From at least the time of Johann Heinrich Linck the Eld-

er, herpetology was a focal point in the Linck collection.

By 1727 the collection included 800 jars of spirit preserved

specimens, including many snakes and other herpetolog-

ical specimens (Kanold in Neickelius 1727). By 1783,

when the first and only published catalogue of the collec-

tion appeared, amphibians and reptiles constituted approx-

imately 50% of the spirit preserved collection, or almost

450 specimens. This is larger than the herpetological col-

lection of Marcus Elieser Bloch, assembled in the mid-

to late 18" century, which formed the nucleus of the Zo-

ological Museum of Berlin in 1810 and originally includ-

ed about 380 specimens (Bauer & Giinther 2006).

The catalogue of the Linck collection, the /ndex Musae

Linckiani (Plate 2a) was published in three parts between

1783 and 1787 (incorrectly stated to be different editions

by MacGregor 2007; now available online through Die

Sachsische Landesbibliothek — Staats- und Universitats-

bibliothek Dresden at http://digital.slub-dresden.de/

werkansicht/dlf/9369/1/0/cache.off). The first part (Linck

1783) includes the section “Die Amphibien. Amphibia,”

divided into “Schwimmende Amphibien. Nantes,” includ-

ing lampreys, sharks and rays, sturgeons and a diversity

of teleosts, ““Kriechende Amphibien. Reptiles,” including

most amphibians and non-avian reptiles except snakes, and

“Schleichende Amphibien. Serpentes,” including snakes,

as well as amphisbaenians, caecilians, and some limbless

lizards. The latter two sections occupy pages 61—79 in the

catalogue and are followed by a two-page section

“Einzelne Theile von Amphibien” comprising skins, eggs,

and other parts of “Amphibien” as broadly construed. An

appendix in part | (Anhang; pages 275—277) lists speci-

mens that initially were overlooked or obtained while the

catalogue was in press and includes on page 276 a single

Bonn zoological Bulletin 62 (2): 220-252

additional specimen of a snake (Coluber coerulescens) that

appears to have been missed initially. The third and last

part of the /ndex (Linck 1787) contains a supplement (Ad-

ditamenta; pages 245—260) that includes three snakes on

page 248, one being the same listed in the appendix men-

tioned above.

Latin names follow Linnaeus (1766) in nearly all cas-

es (Table 1), and in a few instances, Linck cited a descrip-

tive name from Seba’s Thesaurus (1734, 1735) or anoth-

er source. German vernacular names are also provided,

along with the country or region of origin of the species

(if known), and occasionally more extensive comments.

Linck collection specimen numbers are also provided as

are plate and figure references for those specimens fig-

ured in published works (Plate 2b; Table 2). The /ndex lists

10 species of chelonians, 13 species of frogs, one croco-

dile, two amphisbaenians, one caecilian, 28 species of

lizards, three species of salamanders and 63 species of

snakes (numerous specimens listed by Linck as varieties

of some of these actually represent separate species). In

addition, the /ndex lists three large snake skins, snake ver-

tebrae, rattlesnake rattles, numerous snake, turtle and croc-

odile eggs, and a “snake stone,” supposedly taken from

the head of a large snake.

It is unknown what cataloguing system, if any, was used

prior to the time of Johann Heinrich Linck the Younger.

His system (Linck 1783), however, included several cat-

egories. Dry preparations were numbered separately from

spirit-preserved ones and a separate numbering applied to

specimens kept in numbered drawers. Thus, the cheloni-

ans included specimens I—8 in drawer 25, as well as dry

preparations 95—100 plus 100A and 100B, and spirit spec-

imens 326 and 327 (Table 1). In total, the spirit specimens

range from 293 to 697, but with the addition of a few in-

tercalated specimens with an alphabetical suffix, the to-

tal number of spirit-preserved herpetological specimens

comes to 411. Dry preparations included eight turtles plus

two large snake skins (all probably displayed on the walls

or hung from the ceiling) and smaller specimens stored

in some of the 184 drawers devoted to zoological speci-

mens (eight chelonians in drawer 25, one frog and seven

lizards in drawer 27, two snakes in drawer 24, a third snake

skin and an unspecified number of rattlesnake rattles in

drawer 127, crocodile and turtle eggs in drawer 17, and

snake vertebrae and “snake stone” in drawer 175). Unfor-

tunately, no trace of these earliest labels remains with the

specimens today, nor were any labels apparently present

at the time of the reorganization of the collection in

1933/34 (Seifert 1935), and no correspondence between

these early numbers and those used subsequently can be

established, except in cases where the extant specimens

are both unique and known to have been part of the col-

lection in the late 18" century.

Another copy of the Linck /ndex in the Universitatsbib-

liothek Leipzig bears additions in the hand of J.H. Linck

©ZFMK

Aaron M. Bauer & Richard Wahlgren

LEI TOE OTTO OOOO ETE,

INDEX j 92 Bierte Cintheifung.

| MYSAEI LINCKIANI, Coluber buccatus, rin. Der Breitbarke, ans Suc

nam, Sch. T. 660. 1. Liv. p. 377-4. f.

oder 4 ; (in Spir. No. 651. 652.)

2 - Coluber berus, rrw. Die curopdifche Matter. Diefe

furzes emati ini Sdlangen haben eit fehe zhhes Kebeas denn wenn man

: fof atifaes Betjeidynif ignen gleid) den Kopf abfchmeidet, Gebslten fie doch

Dev vornehnften Stice i Die Geifeude Berwegung. Sie paaren fic) eimmal des

4 Sabres, und find 5 Monate tridtig. Sie find giftig,

Keanken verordnet, — S By G8. c Lip. p. 377+

Lindifden | | sett etme ate, 6.1.65. 3, Aa

Cin Spir. No, 499.)

Natucalien antmun Coluber angulatus, cin. Die ecigte Matter, aus

| Qin. Sch. T. 630.b. Lio. p. 377. LK 5O-

g ee Cin Spir. No. 502.) X }

gu Lei PsiG. ; Gine aus Cenlon decal. ;

‘ * Cin Spir. No, 503, und 654.)

Coluber apis, rin. Die Otter, Sie ift giftig und fol!

- Bitfe Ut fenn, womit ehemals fo viele Sclangenbefwes

rungen vorgenommen morden find. Die Egyptier aber

offen ibce. Kinder mit diefen Sehlangen fpicten, daber

ift pu i Paaggto daft fie nur gu eenife Zeiten iftig

fenn miiffen. Lio. p.. 378,

(in Spic. No, 690.)

Coluber lebetinus, crx. Die Kupfernatcer, aus dens

ss Sit aiftig. Lin. p. 378.

in Spit. No. 576.)

Coluber melanocephalus, urn, Der SH:

aus Umerifa, Sch. T. 737.2. Lin. p,

[SRA

Erfter Theil. ; f (in Spit. NO?574. $75. $78- 673) SK. OS

‘ Coluber reginac, rin. Dle Konigénatter, aud “obien,

| A i Lin. p, 378. :

} Leivzig ey (in Spit, No. 580. 581-)

y in det Buchhandlung ver Gelesrten, 1783. ye 5 es

fal ~ [b]

Oiler Aye tis Ae race eae g

a arta

ee pies Aiport Me nafaa.

ys ;

ge ot

SL fteiinalrs >»

Sh egeays, LY fan.

» Aichecabiss Zp Jong bon Segre

ba i on Ti |

Seo Ferpecse buen. ewy nas,

Laff ot Wet

= a 2

Ceetted ned

CHE Mes hee

x t belies ta

Augerer fragilis ter

YA, Yo Lond he

G + Plow

a pliers Che, dag, 9 hes

a Lhe toes athe / pod

g oe ba Uasbootfeszers

ot Winban Mew

f EE apgsleuagibe LRG ofp emaps

Shot vol Bitch ae

(He op- pp BUD) daptatn fispan j

[c] [d]

Plate 2. a. Title page of volume I of the /ndex Musae Linckiani (1783). Image courtesy of Universitatsbibliothek Leipzig. b. Typ-

ical snake specimen accounts in the /ndex Musae Linckiani (1783) showing Linck’s associations with names from Linnaeus (1766)

and reference to Scheuchzer (1735) plates. “LK” (Linck Kabinett) numbers corresponding to the illustration numbers in the /cones

Serpentum et Viperarum have been added in the hand of J. H. Linck the Younger. Image courtesy of Universitatsbibliothek Leipzig.

c. Reverse of inserted leaf in the Universitatsbibliothek Leipzig copy of the Index Musae Linckiani (1783), with additions in the

hand of J. H. Linck the younger. Image courtesy of Universitatsbibliothek Leipzig. d. Page of reptile entries in Verzeichnis der im

Fiirstlichen Museum zu Waldenburg befindlichen Gegenstdnde (Archiv Museum Waldenburg, 315), the 1886 hand-written versi-

on of the museum catalogue. Photo: A. M. Bauer.

The Historic Linck Collection of Snakes 227

HMittetlungen

des Furftlid oN ta

Sdhonburg-Waldenburgfhen |

Familienvereins

Sdlof Waldenburg

Fig. 6. | Cover of number 8 (Mohr, 1940) of Mitteilungen des

Fiirstlich Schénburg-Waldenburgschen Familienvereins, pub-

lished by the Familienverein “SchloB Waldenburg.” Courtesy of

Museum fiir Naturkunde, Berlin.

himself (Mohr 1940; Budig 1999b). In the herpetological

portion of the catalogue there are three such annotations.

On page 76, specimen number 684 has been stricken out

under the entry for Coluber haie and, on a leaf inserted

after this page, the same number has been assigned to Co/-

uber Dipsas (specimen 684). On the reverse of the same

leaf Coluber Aspis Aegyptiaca (specimen 818) and Col-

uber Indicus putorius gryseo-luteus (specimen 817) have

both been added (Plate 2c). These changes must have been

made very shortly after the publication of the catalogue,

as the third part of the catalogue (Linck 1787) includes

an Additamenta section in which they are printed along

with reference to the originally omitted specimen of Col-

uber coerulescens. Further, the specimen numbers as-

signed to the two new entries closely follow the highest

number for a spirit preserved specimen in the first part of

the catalogue (815, a monkey mentioned in the list of spec-

imens obtained while the catalogue was in press). This al-

so suggests that specimens added after the publication of

the catalogue were given sequential numbers, regardless

of the taxonomic group to which they belonged.

Following Linck’s catalogues (1783-1787) the next

recorded accounting of the herpetological holdings of the

collection was the Verzeichnis der im Fiirstlichen Muse-

um zu Waldenburg befindlichen Gegenstdnde (Archiv Mu-

seum Waldenburg, 315). This handwritten document, pre-

pared in 1886, lists the spirit preserved amphibians and

Bonn zoological Bulletin 62 (2): 220-252

reptiles on 10 pages. In this list, which uses contempo-

rary nomenclature, no specimen numbers are listed, only

the number of jars (Plate 2d). This list certainly includes

newer material, probably from the Oberlander collection —

also purchased by Otto Victor in 1840, as distinctive

species, such as Phrynosoma orbiculata [sic], not from the

Linck collection, are included. However, a diversity of ma-

terial corresponding to the older collection is also listed.

Three handwritten copies of this document exist at the Mu-

seum, differing only in presentation but not content with

respect to “Amphibien in Spiritus.”

In the 1930s a major reorganization of the Naturalienk-

abinett was undertaken in which an attempt to identify the

Linck specimens present since at least the time of Johann

Heinrich the Younger was made. This took place during

the time of Prince Gunther of Schénburg (1887-1960)

who took an active interest in the museum and its collec-

tions and was active in the Fiirstlich Sch6nburg-Walden-

burg’scher Familienverein “Schloss Waldenburg,” which

published its own Mitteilungen (Fig. 6), with contributions

about the historical collections.

The identification of these early specimens was carried

out in 1933 and 1934 by the mineralogist Alfred Seifert

(1906-1953) and the zoologist Konstantin Leopold W6p-

ke (????-1944; Fig. 7) as part of a major reorganization

a eS eae: £

ees

Fig. 7. Zoologist Konstantin Wopke (left), museum guard Al-

bin Clauder (center), and mineralogist Alfred Seifert (right) in

front of the Museum Waldenburg Naturalienkabinett und

Heimatmuseum in 1933 or 1934. Wopke and Seifert reorgan-

ized the Waldenburg Museum collections and attempted to iden-

tify material dating to the time of Johann Heinrich Linck the

Younger. Image courtesy of U. Budig, Museum Waldenburg —

Naturalienkabinett und Stadtmuseum.

©OZFMK

Fiihver

durd) das Fiirftlid) Schonburgifche

Natuvalienkabinett in Waldenburg/Sa.

und die in ihm enthaltenen Sammiungen

Bearbeitet von

Dr. fonftantin Wopke, Naumburg/Saale

1937

Druch: Buchdruckerei €. Adfner, Waldenburg/Sachfen

Fig. 8. Title page of the guide to the reorganized Waldenburg

Museum prepared by K. Wopke (1937).

of the Waldenburg collections (Budig 1999a), although

subsequent workers have identified other zoological spec-

imens from Linck’s collection that were not recognized

as such during the 1930s (Mohr 1940). Exactly how pre-

sumed original Linck specimens were recognized is un-

known. It is unclear if the jars used at the time of the cat-

aloguing in 1783 were actually numbered, or if the posi-

tions of the jars on shelves were numbered. In either case,

no unambiguous numbering system of specimens seems

to have survived to the 20" century (Seifert 1935), nor does

the earlier catalogue of 1886 suggest that any numbering

system was present at that time (W6pke 1937). Rather, it

is likely that W6pke tried to match specimens to those list-

ed in Linck (1783-1787). Seifert (1934, fig. p. 36; 1935,

fig. 1) figured a selection of spirit-preserved specimens,

chiefly snakes, that were supposedly from the Linck col-

lection. No markings can be seen on the jars (although

some numbering system may have been used on the top

of the bladders capping the jars). Wopke, who had actu-

ally published a single paper in herpetology (Wopke 1930)

based on his Inaugural-Dissertation from the University

of Leipzig, later published a visitor’s guide to the collec-

tions (Wo6pke 1937; Fig. 8) in which he stated that he be-

lieved that a large number of the specimens on display dat-

ed to the time of J.H. Linck the Younger.

Specimens believed to be part of the original (pre-1783)

Linck collection were given red labels in the 1933/34 re-

Bonn zoological Bulletin 62 (2): 220-252

Aaron M. Bauer & Richard Wahlgren

organization (Beyrich 1990; Fleck et al. 1990) and these

remain associated with these specimens today. Material

present at the time of the reorganization is recorded in the

Accessions Katalog der zoologischen Abteilung des

Furstlich-Schonburgischen Museums zu Waldenburg

i./Sa. (Archiv Museum Waldenburg 342; Fig. 9) prepared

by Wopke. In this document, ruled columns record Lfde.

Nr. (consecutive number = a different number assigned to

each species), Bezeichnung (description = species and type

of preparation), Fundort bezw. Vorkommen (locality or,

more commonly, distribution), Zahl (number of speci-

mens), Dat. (date — only for “modern” specimens), Stan-

dort (physical position in the collection), and Bemerkun-

gen (remarks — generally correspondence to the page and

specimen number in the /ndex Musae Linckiani). In gen-

eral, the first three columns are typed and others have en-

tries in ink; additions and corrections to the names and lo-

calities are also entered in ink in Wépke’s hand. The old

collection of amphibians and reptiles occupies seven pages

in the catalogue. In addition, two mounted reptiles are list-

ed from the African trip of Otto Viktor I, both collected

in 1908: “2069, Crocodilus [sic, Crocodylus] niloticus

Laur., Nilkrokodil, Afrika” and “2070, Emys orbicularis

anata Vipsrinus Lair,

re *

“as SUODA, Atrike

Pketlnarcor z

Seneoie EeooNeKOs sn Lave. Bostik

arin ae |

Ee MSO SM NON A

Batman any Arata! Lewes 36 18)

4 Wa eecte Moe tase Deans” COA MCE |

4 ADA nents, ln nag Arsen

Qoyunee D: Lara 1 OE

4 ths Ginette asi, Lew ints

Me btippimnatte nu Mtn” Loa Yl

nO. ;

an”

ie heey aro Kory ate oe

O.melenocephalvs A A Vous ie Geneseo

4 Ale Renata tam Prairies CaS

é a 0 SEE [

he moagonan, Main tne Sean

A law Sphter

fhe apg oe TL]

fe iar as ito

dearer

Fig.9. Page of reptile specimens from Accessions Katalog der

zoologischen Abteilung des Fiirstlich-Schonburgischen Museums

zu Waldenburg i./Sa. (Archiv Museum Waldenburg 342), a cat-

alogue of the collection prepared by K. Wépke during the reor-

ganization of the museum in 1933. Photo: A. M. Bauer.

OZFMK

The Historic Linck Collection of Snakes 229

baumirox?

| ss pies |

Rineremanncs

Fig. 10. View of part of the herpetological collection in the Naturalienkabinett Waldenburg showing jars sealed with glass plates

and covered in picein that were prepared during the 1994 reorganization of the collection. Photo: A. M. Bauer.

L., Sumpfschildkréte, Europa u. Westasien” [this speci-

men is actually the large and highly distinctive softshell

turtle, Zrionyx triunguis, the misidentification suggests that

Wopke’s knowledge of systematic herpetology was lim-

ited]. The number of entries for the old collection totals

98 species and approximately 250 specimens, falling in

the number range 527— 645. The next inventory was con-

ducted in 1994 and was recorded on index cards, one per

specimen, each with data on one side and a black and

white photo on the reverse. The specimens were each giv-

en numbers (applied to their jars or to the specimens them-

selves in the case of dry preparations), where the number

is not shared with either the Index Musae Linckiani or the

1933/34 Accessions Katalog. Each number 1s prefaced by

a Roman numeral “I,” signifying a zoological object, and

followed by the designation “A,,” indicating a herpeto-

logical specimen according to the museum inventory sys-

tem devised by Knorr (1957). These same specimen num-

bers were recorded in the most recent inventory (2001)

and small labels bearing these numbers have been uni-

formly affixed to specimen jars and dry specimens or their

mounts.

Bonn zoological Bulletin 62 (2): 220-252

The existing fluid preserved specimens were last

topped-up with ethanol and their containers sealed in

1993-1994 by the then preparator of the Museum, Olaf

Zinke. At least some of the containers of amphibians and

reptiles are conserved in 80% alcohol (presumably

ethanol) and 5% glycerin, as recommended by Arndt

(1937, 1943). The glass jars are sealed with a glass plate

and covered in picein (Fig. 10). Prior to this the prepara-

tions were in 95% alcohol and sealed with a tin plate and

cow’s bladder (Zinke 1999; see Beyrich 1990, fig. 6; Zinke

1990, fig. p. 90), a common sealing technique of the late

18" century (Bauer & Gunther 2006). This and other 1 7th

and 18" century methods for sealing glass containers for

fluid-preserved specimens were presented by Ruysch

(1710), Turgot & Duhamel du Monceau (1758), Pallas

(1781), Pole (1790) and Osiander (1793) and these and

other methods were later summarized by Naumann (1815),

Thomas (1892), Altner (1984), and Carter & Walker

(1999).

The existing spirit-preserved collections in Waldenburg

comprise 120 fish, 18 amphibians, 95 reptiles and 22

anatomical preparations (Zinke 1999). The fish have been

©ZFMK

230

Fig. 11.

Dry preparation of a loggerhead sea turtle (Caretta

caretta) suspended from the ceiling of the “Linckzimmer,” a

room in the Naturalienkabinett containing specimens believed

to be from the period of the Linck family and arranged in the

fashion of the 18" century. Photo: A. M. Bauer.

discussed earlier (Mohr 1940). Herpetological specimens,

both ethanol preserved and dry preparations, are found

chiefly in “das erste griine Zimmer,” whereas a few jars

occupy a glass-fronted cabinet on the stairwell landing,

and several large sea turtles and some additional spirit pre-

served specimens are in the “Linckzimmer’” in an arrange-

ment that has remained essentially the same since the

1933-34 renovations (Fischer 1936; Fig. 11). Two addi-

tional mounted reptile specimens, a Nile crocodile (Croc-

odylus niloticus) and African softshell turtle (Trionyx tri-

unguis) dating from Otto Victor II’s African expedition are

on display in “das braune Zimmer.”

SCHEUCHZER, SEBA, AND LINCK’S ICONES

From 1714 onward, Johann Heinrich Linck the Elder

maintained a correspondence with the Swiss naturalist and

physician Johann Jakob Scheuchzer (1672-1733) (Beyrich

1994; Budig 1999b). Linck had just purchased the collec-

tion of Christian Sigismund Wolf (1685—1737), which in-

cluded material in turn obtained from Scheuchzer. In the

intervening years, the two exchanged specimens, as well

as letters (Beyrich 1994). Linck’s letters to Scheuchzer are

today preserved as Ms. H 304 (pp. 211-306) in the Zen-

tralbibliothek Zurich. Linck had originally planned to pub-

lish one or more works on zoology, including snakes, with

Albertus Seba, and had even sent copper engravings of

snakes and other specimens to him. However, Seba re-

turned these to Linck, although he had made copies, some

of which served as the basis for illustrations in Seba’s own

Thesaurus (1734-1735). As has often been noted, how-

ever, Linck objected to plate layouts like those of Seba,

which combined a variety of unrelated natural history ob-

Bonn zoological Bulletin 62 (2): 220-252

Aaron M. Bauer & Richard Wahlgren

PHYSICA

SACRA

JOHANNIS JACOBI

SCHEVCHZERI,

Medicine Doctoris,& Math. in Lycco J igurino Prof.

Academie Tmperialis Nature Curiofor. LEOP( ILDINO-

ROLIN c. Reg.

ICONIBVS

ENEIS

illuftrata,

procurante & fumtus fuppeditante

JOHANNE ANDREA PFEFFEL,

Auguttano, SACRA CASAREA MAJESTATIS

Chale

TOMVS IV.

A TAB. DXCIV. ad DCCL.

vapho Aultico.

AVGVYSTA VINDELICORVM & VLAMA,

MDCCXXAY,

Fig. 12. Title page to volume 4 of the Physica Sacra

(Scheuchzer 1735), which includes all 24 of the plates depict-

ing Linck’s snake collection.

jects in order to achieve an artistic effect (Miisch 2000).

Ultimately he turned to Scheuchzer in order to have his

snake images published.

Scheuchzer’s Physica Sacra or “Copper Bible”

(1731-1735; Fig. 12) was illustrated by 760 copper en-

graved plates, the vast majority of which depicted bibli-

cal scenes incorporating aspects of natural history, which

were designed by Johann Melchior Fiili (1677-1736) and

engraved by Johann Andreas Pfeffel (1674-1748) (Fisch-

er 1973; Mtisch 2000). The history of this grand under-

taking and its execution are chronicled by Miisch (2000).

Included among the biblical scenes were a number of

plates illustrating actual natural history specimens. Among

these was the famous Homo diluvii testis (first described

by Scheuchzer in 1726) as well as 24 plates (606,

628-630, 647-648, 652-655, 657, 660-662, 678, 715,

717, 737-739, 746-749) depicting 67 snakes and amphis-

baenians from the Linck collection (Table 2), all in the

fourth and final volume of the work. These plates appear

to have been executed by 1730 (Seifert 1934), establish-

©ZFMK

The Historic Linck Collection of Snakes

ing a minimum age of specimens illustrated in the Phys-

ica Sacra.

The Leipzig University Library today holds a set of col-

ored illustrations of fish and reptiles (Jcones Piscium Mu-

sei Linckiani nach daselbst befindlichen Originalen

gemahit and Icones serpentum et viperarum Musei Linck-

iani nach daselbst befindlichen Originalen gemahit — here-

after simply the /cones), rediscovered in 1998, that are be-

lieved to have been the original drawings upon which the

plates of Scheuchzer were ultimately based (Budig

1999b, Zinke 1999). The snakes are illustrated with one

specimen per sheet, except for plate LX VI, on which two

snakes are figured. The paintings were originally prepared

under the direction of Johann Heinrich Linck the Elder and

were sent to Scheuchzer no later than 1730 (they are men-

tioned in a letter sent by Linck to Scheuchzer on 17 March

1730; Zentralbibliothek Ziirich Ms. H 304). In the text ac-

companying the first of the Linck snake plates (pl. 606)

Scheuchzer (1735) specifically mentions that J.H. Linck

had sent him color illustrations for use in preparing the

plates. Johann Heinrich Linck the Younger subsequently

bound the paintings returned by Scheuchzer and append-

ed his own notes to them (Budig 1999b). Linck’s notes,

in Latin, appear on the pages facing the corresponding

plates. They typically provide the Roman numeral of the

plate, a copy of the text from Linnaeus (1766) for the

species to which Linck believed each snake to belong, and,

at the bottom, Scheuchzer’s (1735) plate and figure des-

ignation, a page reference to Linnaeus (1766), and the col-

lection number of the spirit specimen illustrated, as record-

ed in the Index Musae Linckiani. Presumably, when mul-

tiple specimens were listed these also included specimens

referable to not only to the same species as recognized by

Linck, but also the same variety or form (see Table 1). In

all there are 89 plates numbered I-XC (plate LXXXVI is

apparently lacking, see Table 3). The first 67 of these, with

the exception of number LXVI, are all represented by fig-

ures and accompanying text in Scheuchzer (1735) and of

these only IV and XLVI are not explicitly associated with

particular spirit specimens (although the number of the

specimen corresponding to plate XLVII was handwritten

into Linck’s annotated copy of the /ndex). Plates 68-90

generally lack text and references to Linck collection spec-

imens, although plates LX XII, LXXIX, LXXXIV, and

LXXXV do correspond to particular specimens (Table 3).

The Roman numeral plate numbers in the /cones are

equivalent to “Musei Linckiani” specimen numbers rep-

resented in Arabic numerals on the Physica Sacra plates

for each corresponding species and corroborate that the

Icones illustrations were available to Scheuchzer. These

same numbers were also added to the annotated copy of

the /ndex in in Linck’s hand, prefaced by “LK” for Linck

Kabinett (Plate 2b).

Thirteen pen and wash paintings similar to those in the

Icones, were offered at an auction in London on 13 May

Bonn zoological Bulletin 62 (2): 220—252

23]

2004, but went unsold and were offered again for sale in

2013. These each depict multiple specimens, presumably

clustered for Scheuchzer’s use. Eight of these correspond

in content to the finished plates as published in the Phys-

ica Sacra whereas the remaining five images were never

published and possibly never even engraved. These paint-

ings appear to be annotated in Linck’s hand, suggesting

that they were prepared in Leipzig rather than having been

executed in Ziirich from drawings sent by Linck. The cor-

respondence between these paintings and those in the

Icones is exact and the former were probably traced from

the latter.

TAXONOMIC SIGNIFICANCE OF THE LINCK

COLLECTION

During the 18" century, the Linck collection was visited

by a number of well-known taxonomists, who selected

specimens as type material. For example, among ichthy-

ological specimens, the extant holotype of Lutianus linckii

(or Lutjanus linkii; the name is spelled differently in the

text and on the accompanying plate in the description fide

Eschmeyer 1998), a synonym of Symphodus tinca (Lin-

naeus, 1758), was described and figured by Marcus Ele-

sier Bloch (1790), who visited the collection in 1767

(Mohr 1940; Zinke 1999). Although few herpetological

taxonomists appear to have seen or worked with the Linck

collection, Scheuchzer’s plates in the Physica Sacra made

at least some of the collection well-known to a broad au-

dience. As was common at the time, and well into the 19"

century, these images were sometimes chosen as icono-

types and the specimens upon which the images were

based served as holotypes or syntypes of validly described

taxa. Among the Linck snakes at least seven specimens

illustrated by Scheuchzer (1735) represent type specimens.

Linnaeus (1758) based two names, in part, on specimens

from the Linck collection (Bauer 2012). Boa scytale, con-

sidered a synonym of Eunectes murinus (Linnaeus, 1758)

by McDiarmid et al. (1999), is based on indications to a

specimen noted by Gronovius (1756:55) and one illustrat-

ed by Scheuchzer (1735) as figure | on plate 737 of Phys-

ica Sacra (Plate 3a). Linck (1783), however, considered

this illustration to be of Coluber aesculapii — nos. 483-484

in the Linck catalogue, a species now called Scaphiodon-

tophis venustissimus (Wied, 1821). Linnaeus (1766) lat-

er revised and expanded the description of B. scytale in

the 12" edition of Systema Naturae so significantly from

that of 1758 as to be unrecognizable as the same species.

Smith & Gloyd (1963) consequently restricted the appli-

cation of the name Boa scytale to the anaconda, E. mur-

inus, the species clearly intended by Linnaeus (1766) and

consistent with the general use of the name over the pro-

ceeding two centuries.

©ZFMK

232 Aaron M. Bauer & Richard Wahlgren

Plate 3. a. Figure 1, plate 737 from Physica Sacra (Scheuchz-

er 1735). The Linck specimen illustrated is one of two syntypes

of Boa scytale Linnaeus 1758 [= Eunectes murinus (Linnaeus

1758)] but is actually referable to Ervthrolamprus aesculapii.

(see text for explanation). In this and other figures from Physi-

ca Sacra elements other than the specimens of interest have been

digitally removed for clarity. b. Figure 4, plate 747 from Phys-

ica Sacra (Scheuchzer 1735). The Linck specimen illustrated 1s

one of two syntypes of Anguis reticulata Linnaeus 1758 [= Ty-

phlops reticulatus]. ¢. Figure 1, plate 647 from Physica Sacra

(Scheuchzer 1735). The Linck specimen illustrated is the holo-

type of Scytale Scheuchzeri Merrem, 1820 [= Cylindrophis ruf-

fus (Laurenti 1768)].

A second Linck specimen serves as a type of the Lin-

naean name Anguis reticulata = Typhlops reticulatus fide

McDiarmid et al. (1999). Linnaeus (1758) cited both a

Gronovius (1756) specimen, and Scheuchzer’s illustration

(pl. 747, fig. 4). Scheuchzer’s illustration clearly shows

the diagnostic pale snout and pale tail ring of this species

(Plate 3b).

Merrem (1820) also cited a number of Scheuchzer plates

in his original descriptions. Scytale Scheuchzeri Merrem,

1820, a synonym of Cylindrophis ruffus, is based solely

on the snake depicted in Scheuchzer’s plate 647, figure |

(Plate 3c). Elaps corallinus Merrem, 1820 (now Micru-

rus corallinus) was based on Scheuchzer’s plate 648, fig-

ure 2 (Plate 4a) and questionably plate 737, figure 1, al-

though the names Coluber fulvius Linnaeus, C. nigro-ru-

fus La Cépéde, and C. fulvus Latreille were also associ-

Bonn zoological Bulletin 62 (2): 220-252

got

Plate 4. a. Figure 2, plate 648 from Physica Sacra (Scheuchz-

er 1735). The Linck specimen illustrated was one of the sources

for the description of Elaps corallinus Merrem, 1820 [= Micru-

rus corallinus]. b. Figure 2, plate 746 from Physica Sacra

(Scheuchzer 1735). The Linck specimen illustrated depicts the

holotype of Coluber (Natrix) cancellatus Merrem, 1820 [= Li-

ophis reginae (Linnaeus 1758)]. ¢. Figure 1, plate 648 from Phys-

ica Sacra (Scheuchzer 1735). The Linck specimen illustrated is

a syntype of Cophias Jararaca Merrem 1820 [= Bothropoides

jararaca].

ated with the latter specimen by Merrem. According to

Roze (1966) Merrem’s description was also based on spec-

imens collected by Wied and published on by him later

in the same year (Wied 1820). Roze (1966), therefore, des-

ignated AMNH 3911, collected by Wied, as the lectotype.

Scheuchzer’s plate 746, figure 2 (Plate 4b) depicts the

holotype of Coluber (Natrix) cancellatus Merrem, 1820,

a synonym of Liophis reginae. Cophias Jararaca (= Both-

ropoides jararaca) also appears to have been described

by Merrem (1820) based, in part, on Schecuchzer’s plate

648, figure | (Plate 4c). However, McDiarmid et al. (1999)

and other sources credit the name to Wied (1824). Al-

though Coluber ornatus (= Chrysopelea ornata) is cor-

rectly credited to Shaw, 1802, Schlegel (1837) cited

Scheuchzer’s plate 606, figure C as an image of a syntype

of Coluber (Natrix) ornatus Merrem, 1820. Schlegel

(1837) also cited Scheuchzer’s plate 630, figure A as the

image of the type of Coluber (Natrix) Scheuchzeri (= ?

Platyceps rhodorachis fide Wallach 2012), although Mer-

OZFMK

TABB. DCLUI

The Historic Linck Collection of Snakes

DCLIV. DCLV.

Mich. Cap. VII. vert 17.

Pulucrem in morem Serpentis lin-

gent, ut que repunt in terra, &

ex habitaculis fuis contremifcent,

ante DOMINUM DEUM no-

ftrum paucbunt, & te timebunt.

Lingent puluerem ficut ferpens: ve-

jut reptilia terre proturbabuntur

de adibus fuis: DOMINUM

DEUM noftrum defiderabunt,&

timebunt te.

Vid. Gen. Cap. HI. werf. 14.

Eque hance pratermitto occafionem

fiftendi Serpentes nonnullos ex fa

eopuracie Linckiano.

Fig. 1. Serpens Americanus car-

tuleus.

Fig. 2. Serpens Americanus ferrugineo dor-

fo, Mactilis per totam laterum longitudinem

femilunatis nigncantibus; medio albis, venere

albido nigris punttis notato, capite ex ferrugi-

neo & albo vario.

Serpens Americanus ferrugineis vacis ma-

culis nigns in dorio, a latere (quamis ex um-

bra circulis albis femilunatis.

Hxmorrhous, Hamorthois, colore eft arc-

noe Pee corpus v cro notis Digris & albis va-

egata. Actiws, Avic. Arab. TM. p.138.

IV. 6. 3. 36, eee

_ Fig. 3. Serpens Surinamenfis aquaricus ¢x

nee > Seraco & albo varius , maculis pra

Bm in dorlo nigtis, (ed vane fieure & ma-

Bnitudinis. : . ue

Serpens vulgaris aquaticus Suri

: ig namenfis ex

mare & albo varius.

aris vani ferpentis fpecies. Giauhari

A « ari.

pk: + Tab. triad Serpens Americanus

Bniatus , cents latioribus cupreis & an;

bus fordide albis. gute:

tpens Americanus annulis major

ajorib:

Supro & minoribus fordide albo. aie

. Serpens minor Americanus ¥

mal pots = annulis nigris.

» $- Diptas Surinamenhi i

futco ke $Surir Afis Capite ex nigro,

albo vatio, diademate ex punstis albis

anegatus, tac~

conftante ornatus, collo fuperné purpurco - fu-

fco, ad latera albo, femitxniis dehine per to-

tam dof longitudinem alternis latioribus fu-

{co-purpurcis, anguftioribus dilutioris purpu-

rx in maculas candidas tcrminats , qux folie

caudam prctenuem longam nigricantcem de-

corant: venter Aauelcic.

Dipfas Surinamenfis perbellé noratus.

"Abr UrQogcutta uihaotras aagedir age.

Semper fabobfiura nigricat extrema causa.

Nicand. de Dipfade v. 337.

Diptadi funt nigredinis vetigia plurima, &

color pottrema partis veelus caudam ad nigre-

dinem declinat. duicenna.

Serpens albus in capite lincas habens co-

ronax inftar. Damir.

Serpens Americanus longus, cratfus, capi-

te lation’, cauda cenui & longa. Mine. Cont Hl

82.

Serpens Americanus longus, tenuis, ¢api-

te craifo, cauda cenui. Hine. Cent IV. 15

Serpens Siamentis longi us VOFICQ UUs,

Capite plano & viperino , tuico - nuANores

Fine Get. IV. 56.

hig. 6. Tab. DCLV. Serpens Amencanus ex

Ineo & grifeo (vel fulco) in capite & colle ve~

ludi reticulatus, reliquo corpure (xMiats , Lx-

Nils veluu tratts.

Ser-

(ws)

[Der bunte Leguan

Lacerla marmerats UN.

Ainorihea, Eure pa

[d]

Plate 5. a. Figure 8, plate 655 from Physica Sacra (Scheuchzer 1735). The Linck specimen illustrated is one of the syntypes of

Pseudoboa fasciatus Schneider 1801 [= Bungarus fasciatus]. b. Figure 2, plate 657 from Physica Sacra (Scheuchzer 1735). The

Linck specimen depicted is a syntype of Dipsas multomaculata Reinwardt in Boie 1827 [= Boiga multomaculata]. ¢. Bottom por-

tion of p. 1296 from Physica Sacra (Scheuchzer 1735) showing Latin text related to plates DCLIH—DCLY, which illustrate Linck

snakes. d. Specimen of Lacerta marmoratus = Polychrus marmoratus (12003A3) in the Naturalienkabinett Waldenburg on a red-

labeled stand, signifiying its recognition as a Linck specimen by Wopke in the 1933/34 reorganization of the collection.

Bonn zoological Bulletin 62 (2): 220-252 OZFMK

234 Aaron M. Bauer & Richard Wahlgren

rem (1820) himself listed this image in his account of Co/-

uber fulgidus. Coluber (Natrix) Linkii, another Merrem

(1820) name, despite its patronymic epithet, was not as-

sociated with a Scheuchzer plate by its author.

Schneider’s (1801) description of Pseudoboa fasciatus

= Bungarus fasciatus is based in part on material in the

Bloch collection (now part of the Museum ftir Naturkunde,

Berlin) and in part on the Linck specimen illustrated on

plate 655, figure 8 by Scheuchzer (1735; Plate 5a). Final-

ly, Dipsas multomaculata Reinwardt in Boie, 1827 (=

Boiga multomaculata) was described in part on the basis

of the snake figured in Scheuchzer’s plate 657, figure 2

(Plate 5b).

THE IDENTITY OF SNAKES FIGURED BY

SCHEUCHZER

The identity of the snakes illustrated in Linck’s /cones and

Scheuchzer’s Physica Sacra has never been adequately re-

solved. The identifications published by Linck

(1783-1787) can only be considered tentative as they were

Linck’s best guesses based on those species known to Lin-

naeus at the time of the 12" edition of Systema Naturae

(1766), which Linck used as his main source. Linnaeus’s

descriptions were often brief and Linck had no special

training or knowledge of snakes, so the margin for error

was great. Linck may also have been swayed in his iden-

tifications by the presumed geographic origin of the

snakes. However, the stated origins of most specimens that

reached natural history cabinets in the early to mid-18th

century were mostly unreliable, as the material generally

came through a series of intermediaries, often via Dutch

commercial vessels that may have visited any of the main

areas from which reptiles were exported, namely Suri-

name, the Cape, Ceylon, and the East Indies. In short,

Linck’s (1783-1787) identifications in the /ndex cannot

be relied upon.

Information from Scheuchzer’s (1735) text is likewise

of little use in identifying the snakes of the Linck collec-

tion. It is possible that Johann Heinrich Linck the Elder

had sent Scheuchzer some information about the speci-

mens, such as their presumed places of origin, but

Scheuchzer seems to have used a combination of classi-

cal sources as well as contemporary biological literature,

including Hernandez (1651), Marcgrave (1648) and Piso

(1658), Kaempfer (1712), Ray (1693), and Kolb (1719),

amongst others, in order to provide text that was usually

not particularly relevant to the particular snake figured

(Plate Sc).

Thankfully, many of the plates in the /cones and Phys-

ica Sacra are lifelike enough that some hope of identifi-

cation is possible on the basis of the images alone. Indeed,

until the mid-19" century many herpetologists attempted

to place Scheuchzer’s images into the synonymies of the

Bonn zoological Bulletin 62 (2): 220-252

snakes then recognized (Table 2). Unfortunately for them,

and for us, at the time that the plates were executed, the

significance ventral and subcaudal scale counts, as first

advocated by Linnaeus (1745 et seq.) and used by some

other mid-18" century workers (e.g., Gronovius 1756),

was not appreciated and so these useful pieces of infor-

mation are not recorded in the images or in the text of the

Physica Sacra. It was not until the work of Merrem

(1790a, 1790b, 1809, 1810, 1821) that the utility of oth-

er scalation features, such as head scales and the condi-

tion of the anal plate was recognized and regularly fig-

ured accurately (Schmidtler 2006).

Because their scale counts could not be ascertained Lin-

naeus ignored many of the snakes illustrated by Scheuchz-

er, despite their otherwise largely accurate portrayal (Shaw

1802). However, other workers, many of whom had the

benefit of having seen many snakes in other collections,

attempted to place Scheuchzer’s images into context

(Table 2). Gronovius (1756) was the first to consistently

reference Scheuchzer’s (1735) images, but this work was

Ue 2A Rept Be eh AR

Plate 6. a. Match of (left) Psammophylax rhombeatus

(12022A;3) to (center) Scheuchzer (1735) plate 739, figure 7, and

(right) Icones XXVI. Left and center images: A. M. Bauer. Right

image courtesy of Universitatsbibliothek Leipzig. b. Match of

(left) Amphisbaena alba (12046A3) to (top right) Scheuchzer

(1735) plate 652, figure 1, and (bottom right) cones XXXIV.

Left and top right images: A. M. Bauer. Bottom right image cour-

tesy of Universitatsbibliothek Leipzig.

©OZFMK

The Historic Linck Collection of Snakes 235

mat Ge

ange |

Der Klapperer

‘olalu: rt: 5

Gite 0 Graton ter

Plate 7. a. Match of (left) Boa constrictor (12013A3) to (right)

Scheuchzer (1735) plate 746, figure 1. Photos: A. M. Bauer. b.

Match of (left) Crotalus durissus (12085A3) to (center)

Scheuchzer (1735) plate 738, figure 4, and (right) cones XLI-

II. Left and center images: A. M. Bauer. Right image courtesy

of Universitatsbibliothek Leipzig.

itself pre-Linnaean and did not use binominal nomencla-

ture. Selected images of Linck’s snakes were subsequent-

ly cited by Linneaus (1758, 1766), Laurenti (1768),

Gmelin (1789), La Cépéde (1789), Bonnaterre (1790),

Schneider (1801, 1821), Shaw (1802), and Daudin

(1802a, 1802b, 1802c). Of these authors, only Schneider,

who was based in eastern Germany and who had praised

Linck and his collection (1792, 1797) is likely to have ac-

tually seen the specimens or perhaps at least the /cones

(see footnotes Table 2). Merrem (1820) was the first au-

thor to formally place the majority of the Scheuchzer

plates into the synonymy of recognized species. Boie

(1827) in his critique of Merrem (1820) also commented

on the identity of the Linck snakes. The most comprehen-

sive evaluations of their identities, however, were made

by Schlegel (1837) in his Physionomie des Serpens, a

global review of snakes, and by Dumeéril & Bibron (1844)

and Dumeéril et al. (1854a, 1854b) in Erpétologie

Générale, the most detailed herpetological treatise pro-

duced to that time (Table 2). These authors largely agreed

on identifications, although there were some differences

Bonn zoological Bulletin 62 (2): 220-252

of opinion and some images were regarded as being too

imprecisely drawn to allow unambiguous identification.

We used the identifications of previous authors as a

guideline for our own determinations of Scheuchzer’s fig-

ures, but did not restrict our considerations to these taxa.

We also compiled lists of possible subjects based on our

own knowledge of snakes and obtained suggestions from

a panel of colleagues (see Acknowledgements) with broad

familiarity of snake systematics and identification. Ulti-

mately our identifications as given in Table 2 were the re-

sult of comparisons of the plates with photographs and

specimens of all candidate taxa compiled. In some in-

stances no identifrication was possible. In a majority of

cases we could confirm that the identifications of Dumeril

& Bibron (1844) and Duméril et al. (1854a, 1854b) were

correct. As predicted for an early 18" century reptile col-

lection, virtually all of the specimens represent species oc-

curring in parts of the world to which access was con-

trolled by the Dutch, who were the source of most com-

mercially traded exotic natural history objects at the time.

In particular, the greatest part of the collection is repre-

sented by South American species that would likely have

been collected in Suriname. However, South African

(Cape) species, such as Causus rhombeatus, Psammophy-

lax rhombeatus, and Pseudaspis cana are represented, as

Plate 8. a. Match of (left) Boiga dendrophila (12068A3) to (top

center) Scheuchzer plate 662, figure 11, (bottom center) /cones

XLV, (top right) Seba (1735) Plate 21, figure 1, and (bottom

right) illustration in Shaw (1802). Bottom center image courtesy

of Universitatsbibliothek Leipzig. All other images: A. M. Bauer

b. Match of (left) Helicops angulatus (12014A3) to (right) cones

plate LXXXIHII. Left photo: A. M. Bauer. Right image courtesy

of Universitatsbibliothek Leipzig.

Ses Sa

©OZFMK

236 Aaron M. Bauer & Richard Wahlgren

are Sri Lankan (Ceylonese) snakes, including Cylindrophis

maculatus, Amphiesma stolatum, and Chrysopelia orna-

ta, and those from Indonesia (former Dutch East Indies),

e.g., Cylindrophis ruffus, Boiga dendrophila, and Homa-

lopsis buccata.

IDENTIFICATION OF ORIGINAL LINCK SPECI-

MENS

We examined the entire collection of the amphibian and

reptile specimens present in the Waldenburg collection in

2005 and 2009. These are arranged chiefly in two glass-

fronted wooden cabinets and are in numbered aisles 52,

53, 56 and 57. Additional specimens are on display in the

Linck Room and in the stairwell between the two main

floors of the building. Unfortunately, the specimen labels,

including red labels meant to denote Linck specimens

identified in the reorganization of the 1930s, are attached

to small wooden bases on which the jars stand (Plate 5d).

The jars are weakly glued to the bases and in some cas-

es were loose and in many cases these bases were clear-

ly associated with the wrong specimens. There are no tags

or other identifiers associated with the specimens within.

However, each jar bears a small, uniquely numbered la-

bel, added in the 1993/1994 reorganization of the collec-

tion. None of the specimens in the stairwell display case

was associated with a numbered label or a labeled base

nor were several of the dry preparations in the Linck

Room. Photographs were taken for subsequent confirma-

tion of identification and comparison with Scheuchzer’s

(1735) plates but these were of limited use as all fluid-

preserved specimens were sealed with a glass plate and

covered in picein from the 1993—94 inventory and reor-

ganization, requiring that photos were taken through the

glass. Identification methods were the same as for the

Scheuchzer plate images (see above).

We examined the eight specimens that had been previ-

ously identified as deriving from the Linck collection by

Konstantin Wopke during his reorganization in the 1930s

(Table 4). Because only snakes and other reduced limbed

reptiles and amphibians (amphisbaenians and caecilians)

were figured by Scheuchzer, we had no way of confirm-

ing the lizards indicated as being part of the original Linck

collection. However, we compared all the snake and am-

phisbaenian specimens now present in the Waldenburg

collection, including those without red labels, to the pub-

lished plates of Scheuchzer and to the original illustrations

prepared by Linck in the /cones. Linck’s images and the

Scheuchzer plates are consistent with one another with re-

spect to the approximate body positions of the snakes fig-

ured and we believe these to be accurate representations

of the specimens as they appeared to Linck. Thus, we con-

sidered specimens showing the same body positions and

color patterns to be possible matches. We found plausi-

Bonn zoological Bulletin 62 (2): 220-252

ble matches for three of the red-labeled specimens

(Ahaetulla prasina; Liophis lineatus; Psammophylax

rhombeatus, Plate 6a). In addition, seven specimens not

indicated by Wépke’s red labeling were considered match-

es to Scheucher figures (Amphishaena alba, Plate 6b; Boa

constrictor, Plate 7a; Croatalus durissus, Plate 7b; Boiga

dendrophila, Plate 8a; Corallus hortulanus; Thamody-

nastes pallidus; Naja naja) and one additional specimen

corresponds to a plate in the unpublished /cones but was

not subsequently illustrated in Physica Sacra (Helicops

angulatus, Plate 8b). Thus, if our identifications are cor-

rect, 11 of the specimens illustrated in the Jcones and 10

figured by Scheuchzer (1735) appear to be extant (Table

4). It is certainly possible that additional, unfigured spec-

imens of spirit-preserved snakes, as well other reptiles and

amphibians, both dry and in fluid, may also date from this

time, but this cannot be confirmed based on the evidence

available.

One of the surviving specimens, here identified as Boiga

dendrophila (and more specifically as B. d. occidentalis

by Wallach 2012), has long been a source of contention.

Shaw (1802) noted that this same specimen was figured

by Seba (1735, pl. 21, fig. 1) and by Scheuchzer (1735,

pl. 630 [sic]) and reproduced the image himself (Plate 8a).

Merrem (1821) correctly cited the image as plate 662, fig-

ure 11 in Scheuchzer and considered that the similarity of

the figures indicated that one of the authors had plagia-

rized the other. He concluded that it is Seba who had

copied without attribution and indeed accused Seba of ly-

ing in claiming that he had received the snake from “a

friend with good standing in society, who greatly valued

it.” In fact, given that Linck had initially provided Seba

with images of his snakes (see above), no plagiarism of

Scheuchzer is implied, although it may be argued that Se-

ba misrepresented the specimen figured as being part of

his own collection. Johann Heinrich Linck the elder him-

self, felt ill-used by Seba, who kept Linck’s snake illus-

trations for half a year and copied them in association with

a planned co-authored work on snakes, but ultimately de-

cided to publish on his own (Zentralbibliothek Ziirich Ms.

H 304).

THE OLDEST FLUID-PRESERVED HERPETO-

LOGICAL COLLECTIONS

The Linck collection is of both historical and taxonomic

relevance. The majority of the specimens that can be as-

sociated with some assurance to the /ndex of the 1780s,

the /cones watercolors in Leipzig, or the plates of the Phys-

ica Sacra, are important as links to one of the most im-

portant German herpetological collections of the 18" cen-

tury. At least those 11 specimens that can be matched to

the cones and Scheuchzer’s plates must have been in the

Linck collection at least since 1729 (Linck noted in March

OZFMK

The Historic Linck Collection of Snakes 237

1730 that Seba had kept the drawings of snakes he had

sent from Easter to Michaelmas, presumably of the pre-

vious year; Zentralbibliothek Ziirich Ms. H. 304) making

them among the oldest fluid-preserved herpetological

specimens in the world.

Nehemiah Grew (1641—1712) was one of the first to re-

port on anatomical specimens preserved in fluid, among

them some of Boyle’s earliest preparations (Jahn 1994) —

including a snake, in his catalogue of the Royal Society

Museum (Grew 1681). Some of these specimens were still

extant more than a century later (Dobson 1956), but much

of the early spirit-preserved material is known to have de-

graded by the first third of the 19" century (Select Com-

mittee 1835; Whitehead 1969). An illustration, made in

1703, of the collection of Levinus Vencent (MacGregor

2007, fig. 51) clearly shows jars of spirit preserved spec-

imens, amongst them lizards, turtles and frogs. Other large

herpetological collections were assembled by James Petiv-

er (1663-1718) and Theodor Jakob Klein (1685-1759),

as well Sir Hans Sloane (1660-1753), who ultimately in-

corporated Petiver’s collection into his own. However, no

fluid-preserved herpetological specimens are believed to

be extant from any of these collections (although records

of what once existed in Sloane’s collection are held by The

Natural History Museum, London (Clutton-Brock 1994)).

Despite the relative stability of fluid-preserved speci-

mens, most private herpetological collections dating to be-

fore the middle of the 18" century were eventually de-

stroyed or greagtly diminished by fire, jar breakage, al-

cohol evaporation, or simply curatorial neglect or indif-

ference. Royal or national collections fared somewhat bet-

ter, as they were less likely to suffer some of these vicis-

situdes, but even the oldest of these, the Swedish Muse-

um of Natural History (with collections dating from 1739),

is younger than the Linck collection.

It is impossible to determine the actual age of the snakes

figured in the Physica Sacra and today housed in the Nat-

uralienkabinett Waldenburg, as no records exist that doc-

ument any herpetological portion of the Linck collection

prior to 1730, the year in which Scheuchzer’s plates are

believed to have been executed (Seifert 1934). However,

the collection of snakes was large and well-known sever-

al years earlier (Kanold in Neickelius 1727) and it is cer-

tainly conceivable that some specimens could date back

to the period of Heinrich Linck, and perhaps to as early

as the 1680s, by which time spirit-preservation of reptiles

had become common.

A similar lack of temporal precision applies to the few

other collections of comparable age. Thus, specimens in

the collection of the Franckesche Stiftungen in Halle, de-

spite its late 17" century origins, cannot be accurately dat-

ed beyond 1736, when Gotfried August Griindler

(1710-1775) catalogued and reorganized the existing col-

lection there. The Seba collection likewise can only be dat-

ed approximately, with specimens having been obtained

Bonn zoological Bulletin 62 (2): 220-252

between 1716, the year of the sale of Seba’s first collec-

tion (of which no spirit-preserved material is known to sur-

vive), and about 1735, the year in which the second of the

two volumes of Seba’s Thesaurus that contain herpetolog-

ical specimens was published. The surviving material from

all three collections could therefore be approximately the

same age, but the material from the Linck collection can

be explicitly proven to date to at least 1729 (versus 1736

for specimens in Halle and 1734—35 for the now scattered

specimens from Seba’s collection) and has the potential

for the greatest possible age, given that Heinrich Linck be-

gan his collection more than 25 years before the founda-

tion of the Franckesche Stiftungen, when Albertus Seba

was only a child.

Unfortunately, none of the type specimens from the

Linck collection appear to be among the surviving spec-

imens in Waldenburg. Even if all of the putative Linck

specimens identified by W6pke are among the extant spec-

imens and the majority of other herpetoloigical specimens

also date from at least the time of Johann Heinrich Linck

the Younger, it is clear that the vast majority of this once

massive collection has been lost or destroyed over the

years. It is probable that many were either sold by Rhode

prior to the purchase of the rest of the collection by Otto

Victor and have subsequently become lost or destroyed,

or that they deteriorated and were destroyed at some point

prior to the collection reorganization of the 1930s. Indeed,

the value of such historical specimens was generally not

appreciated until late in the 19" century, resulting in ex-

tensive losses in the name of curation (Steinheimer 2005).

There is a remote possibility that additional specimens 1l-

lustrated by Scheuchzer (1735) and therefore dating from

the time of Johann Heinrich Linck the Elder, potentially

even some of the missing types, may yet be identified if

specimens sold by Rohde can be identified in other col-

lections.

Acknowledgements. We are indebted to Frau Ulrike Budig of

the Museum und Naturalienkabinett Waldenburg who provided

us with free access to the collections and archives of the Muse-

um and kindly answered our queries and facilitated our access

to material in the Universitatsbibliothek Leipzig. This research

could not have been completed without her support over a pe-

riod of several years. We thank Susanne Dietel (Universitatsbib-

hothek Leipzig) for permission to reproduce images from the

Icones Serpentum et Viperarum and Rainer Walter (Zentralbib-

hothek Ziirich) for providing copies of Johann Heinrich Linck

the Elder’s correspondence with Jakob Scheuchzer. The diffi-

cult 18" century handwriting in these letters was graciously ren-

dered into modern German by Margaret and Gunnar Thiem and

Pr. Adolf Pentz, through the kind assistance of Prof. Annika

Thiem. Travel to Waldenburg by AMB was funded through the

Gerald M. Lemole, M.D. Endowment Fund. We thank Frank

Tillack, Christoph Kucharzewski, Patrick David, Markus

Monzel, Bo Stille, Didrik Claesson, Gernot Vogel, and Van Wal-

lach for providing their insight into the identification of the

snakes figured by Scheuchzer. Hinrich Kaiser kindly assisted

with the translation of a tricky passage from Merrem.

©ZFMK

238 Aaron M. Bauer & Richard Wahlgren

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the History and Bibliography of Herpetology 4 (2): 5—6

Wallach V (2012) Synonymy and preliminary identifications of

the snake illustrations of Albertus Seba’s ‘Thesaurus’

(1734-1735). Hamadryad 35 (2): 1-190

Whitehead PJP (1969) Zoological specimens from Captain

Cook’s voyages. Journal of the Society for the Bibliography

of Natural History 5: 161-201

Wied [Wied-Neuwied] M [Prince Maximilian] (1820) Ueber die

Cobra Coral oder Cobra Coraés der Brasilianer. Nova Acta

Physico-Medica Academiae Caesareae Leopoldino-Carolinae

Naturae Curiosorum 10 (2): 107-110, pl. iv

Wied [Wied-Neuwied] M [Prince Maximilian](1824) Cophias

jararaca, Lieferung 8, fig. 6, in Abbildungen der

Naturgeschichte Brasiliens. Landes-Industrie-Comptoir,

Weimar, 90 pls., [182] pp

Wopke K (1930) Die Kloake und die Begattungsorgane der

mannlichen Zauneidechse (Lacerta agilis L.). Jenaische

Zeitschrift fiir Naturwissenschaft 65: 275-318, pls iv—vi

Wopke K (1937) Fihrer durch das Firstlich Schénburgische Na-

turalienkabinett in Waldenburg/Sa. und die in ihm enthalte-

nen Sammlungen. Firstlich Sch6nburgische Naturalienkabi-

nett in Waldenburg, Waldenburg, 38 pp

Worm O (1655) Museum Wormianum. Seu Historia Rerum Ra-

riorum, tam Naturalium, quam Artificialium, tam Domestica-

rum, quam Exoticarum, quae Hafniae Danorum in Aedibus

Authoris Servantur. Apud Johnnem Elsevirivm, Lugduni Ba-

tavorum [Leiden], [12], 389, [3] pp, 2 pls.

Zinke O (1999) Die Welt im Weingeist — Spirituspraperate des

17./18. Jahrhunderts. Pp 83—90 in Budig U, Zenker R & Stadt-

lander G (eds.) Heimatmuseum und Naturalienkabinett Wal-

denburg, Sachsische Museen 7. Sachsische Landesstelle fiir

Museumswesen, Chemnitz

OZFMK

The Historic Linck Collection of Snakes 243

APPENDIX

Table 1. Herpetological material in the Linck collection as listed in the /ndex Musae Linckiani (1783-1787). Binominal names

used in the /ndex are from the 12" edition of the Systema Naturae (Linnaeus 1766) and reflect the identifications of Johann Hein-

rich Linck the Younger. Specimens differing from the typical named forms in size, color, or provenance were listed by Linck 1m-

mediately following the species they most closely resembled. /ndex page refers to the volume (I: 1783; III: 1787) and page num-

ber in the /ndex upon which each species or variety account begins. Specimen numbers alone refer to spirit-preserved material,

“aufgeh.” (aufgehangt = hung) or “trocken (= dry) aufgeh.” refers to preparations hanging on a wall or from the ceiling, “Schubl.”

refers to dry preparations in drawers (Schubladen). Linck provided citations to the corresponding page number in Linnaeus (1766),

as well as to corresponding images in Scheuchzer (1735; see Table 2), Seba (I: 1734, II: 1735), and Miller (as Mill. or Millers

Ueb. or Uebers.; III: 1774, Suppl.: 1776) and mentions in Ray (1693) and Laurenti (1768). Information in this table may be cross-

referenced with that in Table 2 by comparing the Specimen numbers columns.

Index name Index Specimen numbers Linnaeus Other Citations

Page 1766

Testudo coriacea 1:6] aufgeh. 98, 25, Schubl. No. 6 350

Testudo imbricata 1:61 aufgeh. 100a 350

Testudo mydas 1:61 trocken aufgeh. 95, 96 350

Desgl. eine von der nemlichen Art, Lol 326, 327,

aber kleiner trocken aufgeh. 97

Testudo orbicularis 1:61] 25 Schubl. No. 5 351

Desgl. nur die Oberschaale 1:61 aufgeh. 100b

Testudo scabra 1:61 25 Schubl. No. 4, aufgeh. 100 351

Testudo scorpioides 1:61 aufgeh. 99 352

Testudo graeca 1:62 25 Schubl. No. 2 352

Testudo carolina 1:62 25 Schubl. No. 8 352

Testudo geometrica 1:62 25 Schubl. No. 1,3 353 Mullers Uebers. T.

Il. T. XU, f. 1

Noch eine Verschiedenheit 1:62 25 Schubl. No. 9 353

von der Geometrica

Testudo pusilla 1:62 25 Schubl. No. 7 353

Rana pipa 1:62 322-325 354 Millers Ueb. T. III.

T. XU, f. 2

Desgleichen ein Exemplar woran man 1:62 321

die aufgesprungenen Eyer recht genau

beobachten kann

Rana musica 1:62 303 354

Rana bufo 1:62 320 354

Rana gibbosa 1:63 310, 311, 319 355 Seb. II, 37-3

Rana bombina 1:63 312 355

Rana ventricosa 1:63 309 355

Ein kleinerer 1:63 318

Rana marina Americana 1:63 316 356

Rana ocellata 1:63 315 356 Seb. I, 76-1

Rana marginata 1:63 297 356

Rana paradoxa 1:63 293, 294, 294A—-C 356 Seb. I, 78-6,16,21

Rana temporaria 1:63 302 357

Rana arborea 1:63 295, 296, 298-301, 305, 357

306; 27 Schubl. No. 5

Dergleichen ein ganz groker weiber 1:63 307

Surinamischer Laubfrosch

Einer dergleichen mit einem Fusse 1:63 304

Rana boans 1:64 308 358

Dergleichen Art aus Surinam 1:64 313

Dergleichen aus Brasilien 1:64 314

Dergleichen aus Carolina 1:64 317 Phil. Trans.

210:215

Draco volans 1:64 439-442, 27 Schubl. No. 2 358 Millers Ueb. T. III.

T. I, f. 4

Desgleichen die Amerikanische 1:64 443-447 358 Millers Ueb. T. II.

fliegende Eidechse T. Ef.5

Bonn zoological Bulletin 62 (2): 220-252 OZFMK

244

Index name

Aaron M. Bauer & Richard Wahlgren

Lacerta crocodilus

Dergl. noch einige junge afrik. Krocodile

Lacerta superciliosa

Lacerta monitor

Lacerta principalis

Lacerta cordylus

Desgl. Aus Amerika

Lacerta mauritanica

Lacerta azurea

Lacerta turcica

Lacerta ameiva

Desgl. eine mit getheilten Schwanze

Desgl. eine Verschiedenheit

Lacerta agilis

Desgl. mit einem langlichen Auswuchse

auf der Oberseite des Schwanzes

Lacerta seps

Lacerta de taletec. min. mas et foem.

Noch eine Amerikanische

Lacerta chamaeleon

Dergl. Ein Exemplar aus Morea

Lacerta gecko

Lacerta stincus

Lacerta orbicularis

Lacerta ypsilon

Lacerta iguana

Lacerta calotes

Lacerta agama

Lacerta marmorata

Lacerta teguixin

Die grokte Teguixin

Lacerta punctata

Lacerta chalcides

Lacerta lemniscata

Lacerta arguta

Noch verschiedene Eidechsen

Lacerta aquatica

Lacerta palustris

Lacerta salamandra

Crotalus miliaris Lin.

Crotalus drvinas Lin.

Crotalus durissus Lin.

Boa hipnale Lin.

Boa constrictor Lin.

Boa hortulana Lin.

Eine mit braunen Bandern

Coluber vipera Lin.

Coluber atropos Lin.

Coluber plicatilis Lin.

Coluber domicella Lin.

Coluber buccatus Lin.

Coluber berus Lin.

Desgl. die surinamische Natter

Coluber angulatus Lin.

Eine aus Ceylon dergl.

Bonn zoological Bulletin 62 (2): 220-252

Index Specimen numbers Linnaeus Other Citations

Page 1766

1:64 328-334, 336, 340 359 Seb. I, 103, 104, 106

1:65 335, 337-339

1:65 72, 374, 449, 450 360 Seb. I, 1-9-24

1:65 343-349 360 Seb. I, 94-1-3

1:65 417 360

1:65 341, 391, 393 361

1:65 392, 27 Schubl. No. 7

1:66 458 361

1:66 342 362 Seb. II, 62-6, 97-4

1:66 456 362

1:66 376-384, 384, 412, 419, 362 Seb. I, 88-2

27 Schubl.

1:66 398

1:66 394, 395, 413 Seb. I, 85-2-4, 86-4

1:66 375, 397, 403-405, 407, 363 Seb. I, 79-5, 4-5

418, 428, 429

1:66 386

1:66 399 363

1:66 409, 410 Seb. I, 79-1

1:67 406, 27 Schubl. Seb. I, 92-5

1:67 433, 435, 436, 438, 438A, 364 Seb. I, 82-1-5, 83-5

27 Schubl. No. 3

1:67 437

1:67 452, 455, 457, 459-461, 365 Seb. I, 108

27 Schubl. No. 6

1:67 400-402 365 Seb. I, 105-3

1:67 424, 451 365 Seb. I, 109-6

1:68 290A

1:68 350, 351, 353-363 366 Seb. I, 96-4,5,

97-3, 98-1

1:68 364-371, 373, 396 367

1:68 448 367 Seb. I, 107-1-3

1:68 385, 387-390, 414-416, 434 368 Seb. II, 76-4

1:68 420, 421, 423, 425-427 368

1:68 352

1:68 430, 432 369

1:69 431 369

1:69 408, 422 369

1:69 411 Miill. Uebers. im

Suppl. Bande p.

188.41.A)

1:69 264, 579, 27 Schubl. No. 1, 4

1:69 453 370 Seb. I, 12-7

1:69 454 370 Seb. I, 89-4,5

1:69 462-468 371 Seb. II, 15-5

1:70 469 372

1:70 563 372 Seb. II 95-3

1:70 553 372 Seb. II 95-2

1:70 671 373 Seb. II 34-2

1:71 669-670 373

I:71 661-662 374

:71 488

1:71 552 375

I:71 540 375

1:71 564 376

1:71 530 376

1:72 651-652 377

1:72 609 377

1:72 499

1:72 502 377

1:72 503, 654

©OZFMK

The Historic Linck Collection of Snakes 245

Index name Index Specimen numbers Linnaeus Other Citations

Page 1766

Coluber aspis Lin. 1:72 690 378

Coluber lebetinus Lin. 1:72 576 378

Coluber melanocephalus Lin. 1:72 574-575, 578, 673 378

Coluber reginae Lin. 1:72 580-581 378

Coluber severus Lin. 1:73 668 379

Coluber aurora Lin. 1:73 565 379

Coluber stolatus Lin. 1:73 679 379

Coluber vittatus Lin. 1:73 676-678 380 Seb. T. 661.8!

Coluber aesculapii Lin. 1:73 483-484 380

Desgl. aus Afrika 1:73 491, 493-494, 587

Desgl. Eine sehr grosse dieser Art 1:73 649

Desgl. noch verschiedene 1:73 486-487, 560

Noch eine Verschiedenheit 1:73 485, 592

Coluber rhombeatus Lin. 1:73 482, 562 380

Coluber cvaneus Lin. 1:73 635-636 380

Coluber natrix Lin. 1:73 473 380 Seb. IH. 35-4

Dergl. eine aus Surinam 1:74 475-481, 542

Dergl. kleinere 1:74 548-549, 554, 556-558, 614, 680

Dergl. vom Vorgeb. der guten Hoffnung 1:74 471-472, 500-501, 504—S08, 375

viel verschiedene Arten 520-529, 531, 537, 539, 545-547,

666, 681-683

Coluber agilis Lin. 1:74 655 381

Coluber lacteus Lin. 1:74 534 38]

Coluber jaculatrix Lin. 1:74 643 381

Coluber pallidus Lin. 1:74 519, 536, 566 381

Coluber lineatus Lin. 1:74 551, 642, 644 382

Coluber naja Lin. 1:74 645 382 Seb. II. 94-1

Dergl. noch verschiedene 1:74 646-648 Seb. I. 44-1

Coluber padera Lin. 1:75 689 382

Coluber canus Lin. 1:75 541, 573 382

Coluber sibilans Lin. 1:75 685 383

Coluber atrox Lin. 1:75 538 383

Coluber nebulatus Lin. 1:75 509-512, 516-517 383

Dergl. kleinere 1:75 513-514

Coluber fuscus Lin. 1:75 664-665 383

Dergleichen von heller Farbe 1:75 474

Dergleichen von brauner Farbe 1:75 470

Coluber saturninus Lin. 1:75 561 384 Seb. I. 1-8

Coluber corallinus Lin. 1:75 24 Schubl. No. 2 384

Coluber exoletus Lin. 1:75 638 385

Coluber lemniscatus Lin. 1:76 495, 585-586 386

Coluber annulatus Lin. 1:76 535, 695 386

Dergl. Eine Varietaet von der Annulata 1:76 686

Coluber aestivus Lin. 1:76 633 387

Coluber ahaetulla Lin. 1:76 618-628, 637 387

Eine dergl. Kleine Art 1:76 629

mit schwarzen Flecken

Coluber petalarius Lin. 1:76 496 387 Seb. H. 16-2

Coluber haie Lin. 1:76 684 387

Coluber dipsas [1:76] [684] °

Coluber filiformis Lin. 1:76 640 388

Coluber pullatus Lin. 1:76 667 388

Coluber hippocrepis Lin. 1:76 543, 567-568 388

Coluber cinereus Lin. 1:76 555 388

Coluber mucosus Lin. 1:77 641 388

Coluber cenchoa Lin. 1:77 656, 663, 688, 691 389

Noch dergl. verschiedene 1:77 515, 653, 657-660

Dergl. eine Varietat lev] 692-694, 696

Coluber mycterizans Lin. eed 631 389

Dergl. Pfrischbliithfarbne, aus Ceylon 1:77 616-617

Dergl. groBe blaue und griine 1:77 634

Dergl. ganz dunkelblaue 1:77 639

246 Aaron M. Bauer & Richard Wahlgren

Index name Index Specimen numbers Linnaeus Other Citations

Page 1766

Eine, welche die schénsten Farben 1:77 630, 632

wie ein Opal spielt, aus Surinam

Coluber coerulescens Lin.? 1:77 675 389

Coluber reticulatns 1:77 533

Coluber, s Vipera maculata 1:77 672 Laurenti ?

Coluber, s. Vipera americana, 1:77 518 Raj [Ray]

variis characteribus orientalibus inscriptus

Noch verschiedene Nattern, 1:78 569

vom Capite bonae spei

Noch verschiedene Arten aus 1:78 492, 532, 544, 550, 570,

verschiedenen Landern 24 Schubl.

Anguis meleagris Lin. 1:78 572 350 Seb. II. 21-4

Anguis maculata Lin. 1:78 599 39]

Anguis reticulata Lin. 1:78 608 39]

Anguis laticauda Lin. 1:78 559 392 Seb. IH. 30-3

Anguis scytale Lin. 1:78 489-490, 582-584 392

Dergleichen etwas kleiner 1:78 497-498, 588-590

Dergleichen noch kleiner 1:78 591, 611-612

Noch eine Verschiedenheit aus Amerika 1:79 593-595

Anguis americ. ex albo sub nigro tessellatus 1:79 600, 650

Anguis eryx Lin. 1:79 610 392

Anguis fragilis Lin. 1:79 577, 687 392

Amphisbaena fuliginosa Lin. 1:79 596-598, 615 392

Dergleichen 1:79 601—606

Amphisbaena alba Lin. 1:79 607 393

Caecilia tentaculata Lin. 1:79 571, 613 393 Seb. II. 25-2

exuvia boae contortricis 1:80 aufgeh. 102

exuvia boae contortricis 1:80 aufgeh. 101 Seb. I. 53-1

exuvia boae contortricis 1:80 127 Schubl.

coronae serpentis 1:80 175 Schubl. No. 6

lapidees serpentini magnetici 1:80 175 Schubl. No. 8

spina dorsalis serpentis 1:80 175 Schubl. No. 4

ova serpentis 1:80 697A Seb. I. 7

ova serpentis 1:80 17 oder 18 Schubl.

ova lacertae crocodilli i:81 17 Schubl. No. 4

ova testudinis 1:81 17 Schubl. No. 5

Coluber coerulescens Lin. 1:276, 674

H1:248

Coluber aspis aegyptiaca O1:248 818 Seb. II.188-1

[sic, prob. 88-1]

Coluber Indicus putorius gryseo-luteus — 1:248 817

'This reference to Seba by Linck is a lapsus or printer’s error. The “661.8” actually corresponds to the plate (661) and figure (8)

in Scheuchzer (1735) and the intended citation would have been “Sch. T. (Tafel = plate) 661.8”.

*In the University of Leipzig copy of the /ndex annotated by J. H. Linck the Younger specimen number 684 is crossed-out in as-

sociation with Coluber haie and a note has been added on the facing blank page indicating that this specimen should be associ-

ated with Coluber dipsas, figure 5 on plate 738 in Scheuchzer (1735).

Bonn zoological Bulletin 62 (2): 220-252 OZFMK

247

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Bonn zoological Bulletin 62 (2)

Aaron M. Bauer & Richard Wahlgren

248

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Bonn zoological Bulletin 62 (2): 220-252

The Historic Linck Collection of Snakes 251

Table 3. Identification of the snakes and lizard illustrated in plates of the /cones that do not correspond to published figures in

Scheuchzer (1735). Correspondences to the Linck specimen numbers, species names, and Seba references as used in the /ndex

(1783) are indicated as are subsequent references to the unpublished /cones plates. Current Identification reflects the authors’ best

interpretation of the identity of Scheuchzer images based on extensive comparisons with specimens and photographs (see text).

Current name in bold is a specimen positively identified as appearing in the corresponding /cones plate.

Icones Linck (1783) Name in Seba Schneider (1801) Daudin (1802) Current Identific.

Plate n° number Linck (1783)

LXVI Elaps Duberria Chrysopelia pelias

LXVIII unknown

LXIX Duberria lutrix ?

LXX Crotalus durissus

LXXI Coelognathus radiatus

LXXIP 645 Coluber naja Lin. Seb. II. 94-1 Naja naja

LXXIIl Naja naja

LXXIII Liophis reginae ?

LXXIV Liophis cobella

LXXV Anguis rufus Cylindrophis ruffus

LXXVI unknown

LXXVII Ahaetulla prasina

LXXVII Eryx rufus unknown

LXXIX 641 Coluber mucosus Lin.

LXXIX Ahaetulla prasina

LXXX Crotalus sp.

LXXxI unknown

LXXXII Oxybelis aeneus

LXXXII Helicops angulatus

LXXXIV 572 Anguis meleagris Lin. Seb. Il. 21-4 Anguis meleagris Acontias meleagris

LXXXV_ 495, 585, 586 Coluber lemniscatus Lin. Micrurus lemniscatus

LXXXVI plate missing’

LXXXVII unknown

LXXXVIII unknowm

LXXXIX typhlonectid caecilian

XC Spilotes pullatus

' Scheuchzer plate 606 incorrectly indicates Mus. Linck. N. 66 (rather than 67) as the source of figure B.

> Mislabeled XLII in Jcones.

*The Roman numerals associated with /cones plates LXXXV and XC are clearly visible, but there are no numbers on the three

intervening plates. However, handwritten pagination is continuous across the associated pages, suggesting that one plate is miss-

ing. We interpret the missing plate as LXXXVI.

Aaron M. Bauer & Richard Wahlgren

Table 4. Specimens of reptiles in the Naturalienkabinett Waldenburg identified by K. Wépke as having been in the Linck collec-

tion at the time of the Index Musae Linckiani (1783-1787) and those identified by the authors as the likely subjects of images in

1730 in the cones Serpentum et Viperarum (1730) and volume IV of Physica Sacra by Scheuchzer (1735). Inventory numbers of

specimens identified by a red shelf label (signifying presumed Linck specimens) indicated in bold. An asterisk (*) indicates an

egregious mismatch of the specimen and its identification on the associated shelf label/stand, implying that they became dissoci-

ated from one another sometime after the reorganization of the collection in 1933-1934.

Shelf label Actual Identification Inventory N° [cones plate Physica Sacra Figure in

plate—figure this paper

Coluber vittatus Psammophylax rhombeatus* 12022A, XXVI 739-7 plate 6a

Coluber domicella Helicops angulatus* 12018A, no match no match

Coluber filaformis Ahaetulla prasina 12017A, XI 655-7

(2 specimens)

Coluber melanocephalus Liophis lineatus* 12016A, LHI 715-2

Coluber angulatus Helicops angulatus 12101A, no match no match

Lacerta marmorata Polychrus marmoratus 12003A, no match no match

Lacerta calotes Calotes calotes 12037A, no match no match

Ameiva surinamensis Thecadactylus rapicauda* 12040A, no match no match

Amphisbaena alba Amphisbaena alba 12046A; XXXIV 652-1 plate 6b

Boa constrictor Boa constrictor 12013A; XLIV 746-1 plate 7a

Crotalus durissus Crotalus durissus 12085A; XLItl 738-4 plate 7b

Coluber pullatus Thamodynastes pallidus* 12072A, VII 678-1

Boa hortulana Corallus hortulanus no number, XVII 661-9

in stairwell

Coluber naja Naja naja no number, XL 749-9

in stairwell

Dipsas dendrophila Boiga dendrophila 12068A;3 XLV 662-11 plate 8a

Coluber pallidus Helicops angulatus* 12014A, LXXXHI - plate 8b

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Freshwater fishes of Western Indonesia and Sulawesi. Periplus

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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, 1758), but abbreviated at subsequent mention, e.g. C. au-

ratus. List names in synonymies as follows: Attelabus asparagi

Scopoli, 1763 (Scopoli 1763: 36, fig. 113.), 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 1s 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

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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-

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is in the responsibility of the editor-in-chief. This decision is main-

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tions for a cover image.

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Manuscripts will be published online at www.zoologicalbulletin.de

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sponding author must clearly state so if preprint online publication

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All published content is freely available without charge to the user

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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:

. Is the paper of interest to the readers of BZB?

. How well does it meet the focus of BZB?

. Is it acceptable for publication, or should it be reconsidered af-

ter revision?

. Please grade originality, quality of data, quality of statistical analy-

ses, and quality of interpretation on a scale from | = low to 5 =

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4. Please grade clarity and quality of text, figures and tables from

1 = low to 5 = high.

. 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?

ISONIAN INSTITUT!

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Contents

Assing, Volker:

On the Nazeris fauna of China ll.

New species and records from Zhejiang, Sichuan, and Yunnan

(Coleoptera: Staphylinidae: Paederinae)

ION LIBRARIES

88 01722 14

Stiels, Darius & Kathrin Schidelko:

Records of ‘Indian’ Baya Weaver Ploceus philippinus (Linnaeus, 1766) and

Hooded Wheatear Oenanthe monacha (Temminck, 1825) from Afghanistan

(Aves: Passeriformes)

Tuyisingize, Deogratias, Julian C. Kerbis Peterhans, Gary N. Bronner & Tara S. Stoinski:

Small mammal community composition in the Volcanoes National Park, Rwanda

Kerbis Peterhans, Julian C., Jakob Fahr, Michael H. Huhndorf, Prince Kaleme, Andrew J. Plumptre,

Ben D. Marks & Robert Kizungu:

Bats (Chiroptera) from the Albertine Rift, eastern Democratic Republic of Congo,

with the description of two new species of the Rhinolophus maclaudi group _

Kerbis Peterhans, Julian C., Michael H. Huhndorf, Andrew J. Plumptre, Rainer Hutterer,

Prince Kaleme & Benjamin Ndara:

Mammals, other than bats, from the Misotshi-Kabogo highlands (eastern Democratic Republic of

Congo), with the description of two new species (Mammalia: Soricidae)

Bauer, Aaron M. & Richard Wahlgren:

On the Linck collection and specimens of snakes figured by Johann Jakob Scheuchzer (1735) —

in the oldest fluid-preserved herpetological collection in the world?

Cover illustration:

Rhinolophus willardi Kerbis Peterhans et al. (this volume, pp. 186-202)

Leibniz-Gemeinschaft

fur Bildung Wissenschaft und Forschung

LS Bundesministerium \4inicterium fur Innovation,

und Forschung des Landes Nordrhein-Westfalen

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