Bonn zoological Bulletin (BzB), formerly “Bonner zoologische Beitrage”,

is published by the Zoologisches Forschungsmuseum

Alexander Koenig (ZFMK), Bonn. Two regular issues of BzB are published per year; supplements on focus topics are produced

in irregular succession.

Subscription price is 46 € per volume (year), including supplements and shipping costs. For subscription, back issues and insti-

tutional exchange, please contact the ZFMK library (ZFMK, Bibliothek, Frau Diane Steinebach, Adenauerallee 160, D-53113 Bonn,

Germany, tel. +49 228-9122-216, fax: +49 228-9122-212; d.steinebach.zfmk@uni-bonn.de). The online version of BzB is avail-

able free of charge at the ZFMK homepage: http://www.zfmk.de/web/Forschung/Buecher/Beitraege/index.en.html.

Produced by Eva-Maria Levermann, Kaiserstr. 129, D-53113 Bonn, Germany; emlevermann@netcologne.de.

Printed and bound by DCM, Werner-von-Siemens-Str. 13, D-53340 Meckenheim, Germany.

Bonn zoological Bulletin

Editor-in-Chief

Fabian Herder, Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK), Ichthyology Section, Adenauerallee 160,

53113 Bonn, Germany,

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

f.herder.zfmk@uni-bonn.de

Editorial Board

Dirk Ahrens, Insects: Coleoptera, ZFMK,

tel. +49 2289122286, fax: +49 228-9122-332;

d.ahrens.zfmk(@uni-bonn.de

Wolfgang Boéhme, Amphibians and Reptiles, ZFMK,

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

w.boehme.zfmk@uni-bonn.de

Netta Dorchin, Insects: Diptera, ZFMK,

tel. +49 228-9122—292, fax: +49 228-9122-212

n.dorchin.zfmk@uni-bonn.de

Renate van den Elzen, Birds, ZFMK,

tel. +49 228-9122—231, fax: +49 228-9122-212;

r.elzen.zfmk@uni-bonn.de

Bernhard Huber, Invertebrates except Insects, ZFMK,

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

b.huber.zfmk@uni-bonn.de

Rainer Hutterer, Mammals, ZFMK,

tel. +49 2289122261, fax: +49 2289122212;

r.hutterer.zfmk@uni-bonn.de

Gustav Peters, Mammals, ZFMK,

tel. +49 2289122262, fax: +49 228-9122-212;

g.peters.zfmk@uni-bonn.de

Bradley Sinclair, Canadian National Collection of Insects,

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

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

K1A 0C6, tel. + 1 613-759-1787, fax: + 1 613—759— 1927,

bradley.sinclair@inspection.ge.ca

Dieter Stiining, Insects except Coleoptera and Dive

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

d.stuening.zfmk(@uni-bonn.de a

Philipp Wagner, Villanova University, Department of Biolo-

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

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

Advisory Board

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

sitat, Institut fiir Evolutionsbiologie & Okologie, 53113

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

2321; t.bakker@uni-bonn.de

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

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

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

aaron.bauer@villanova.edu

Wieslaw Bogdanowicz, Museum and Institute of Zoology,

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

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

wieslawb@miiz.waw.pl

Matthias Glaubrecht, Museum ftir Naturkunde Berlin, Leib-

niz-Institut fir Evolutions- und Biodiversitatsforschung an

der Humboldt-Universitat zu Berlin, Invalidenstrasse 43,

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

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

berlin.de

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

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

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

Tan Heok Hui, Raffles Museum of Biodiversity Research,

National University of Singapore, Department of Biological

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

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

Boris Krystufek, Slovenian Museum of Natural History, P.

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

Wolfgang Schawaller, Staatliches Museum fir Naturkunde,

Rosenstein 1, 70191 Stuttgart, Germany,

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

schawaller.smns@naturkundemuseum-bw.de

Ulrich K. Schliewen, Department of Ichthyology, Bavarian

State Collection of Zoology, Minchhausenstr. 21, 81247

Minchen, Germany, tel. + 49 89-8107—110;

schliewen@zsm.mwn.de

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

meine & Systematische Zoologie, Anklamer Str. 20, 17489

Greifswald, Germany, tel. +49 3834—-86-4242, fax: +49

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

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

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

60808, Canyon, Texas 79016, USA, tel. +1 806-651-2578,

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

Miguel Vences, Technical University of Braunschweig, Zoo-

logical Institute, Spielmannstr. 8, 38106 Braunschweig,

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

m.vences(@tu-bs.de

Erich Weber, Eberhard-Karls-Universitat, Zoologische

Schausammlung, Sigwartstr. 3, 72076 Tubingen,

tel. +49 70712972616, fax +49 7071—295170;

erich.weber@uni-tuebingen.de

Editorial

Welcome to the 60" volume of the Bonn zoological Bul-

letin. All contents published since 1950 are now available

online at the ZFMK homepage, providing open access to

numerous contributions to organismal zoology. The jour-

nal has now been added to the Directory of Open Access

Journals, further improving the visibility of papers pub-

lished in the BzB. Besides open access status, publication

remains free of charge for authors, including colour illus-

trations. In sum, the journal is increasingly attractive for

the publication of sound scientific work from its three fo-

cus fields, namely (1) taxonomy, (2) systematics and evo-

lution, and (3) biodiversity and biogeography.

Taken together, 32 scientific papers and 10 catalogues of

type specimens housed at ZFMK were published in more

than 730 pages in the first year after changing the BzB ti-

tle from “Bonner zoologische Beitrage” to “Bonn zoolog-

ical Bulletin”. The papers contain, among other topics, de-

scriptions of more than a dozen species new to science,

ranging from butterflies to fishes, frogs and snakes.

The current issue adds new species records, a comprehen-

sive checklist, descriptions of five new species, and fur-

ther contributions to organismal zoology. A molecular

study suggests that an Amazonian Pecari described a few

years ago likely does not represent a valid species, and a

case study conducted on a small tropical island reports on

the accumulating species richness data in a herpetologi-

cal field survey.

Last but not least, it’s an honour to welcome two new

board members. Philipp Wagner, currently at Villanova

University (USA), is a herpetologist and well known to

BzB readers. He has a special interest in agamid lizards

from Africa, has acted as guest editor previously, includ-

ing last year’s special issue “Herpetologia Koenigiana” in

honour of Wolfgang Bohme, and will now join the Edi-

torial Board. Tan Heok Hui from Raffles Museum of Bio-

diversity Research in Singapore will support the BzB team

as a member of the Advisory Board. He is famous among

ichthyologists, especially for his taxonomic work on Asian

freshwater fishes including sucker loaches, fighting fish,

and the smallest fish species recorded so far, a cyprinid.

His interests include also herpetology and vertebrates as

well as invertebrates occurring in marine, brackish and

freshwater habitats of Asia. A warm “welcome on board”

to both colleagues!

With best wishes to all readers, authors and “BzB-team”

members,

Tibi float

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

Bonn, April 2011

- 5 Ui Hig es iia va cae

ae ah a ae "Giese

hu Dene

ae ba |

yee

Bonn zoological Bulletin Volume 60

Issue | | pp. 3-8 Bonn, May 2011 |

Estimating the herpetofaunal species richness

of Pangkor Island, Peninsular Malaysia

Johan van Rooyen!, Chan Kin Onn3, L. Lee Grismer+ & Norhayati Ahmad>

'Netherlands Centre for Biodiversity Naturalis, section Zoological Museum Amsterdam,

Mauritskade 61, NL-1092 AD Amsterdam, The Netherlands

2Tulpentuin 313, NL-2272 EH, Voorburg, The Netherlands; E-mail: j1.van.rooijen@hetnet.nl

3Institute for Environment and Development (LESTARI), Universiti Kebangsaan Malaysia, 43600 Bangi,

Selangor Darul Ehsan, Malaysia; E-mail: kin_onn@yahoo.com

4Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California,

92515-8247 USA; E-mail: lgrismer@lasierra.edu

5School of Environment and Natural Resource Sciences, Faculty of Science and Technology, Universiti

Kebangsaan Malaysia, 43600 Bangi, Selangor Darul Ehsan, Malaysia; E-mail: yati_68@yahoo.co.uk.

Abstract. Herpetological surveys of Southeast Asian tropical ecosystems rarely, if ever, result in complete inventories.

This is due to the fact that surveying to completion requires huge investments in terms of search effort. As a result, the

presented species lists usually represent subsets of the total herpetofaunal assemblage and consequently do not shed light

on the total species richness of the investigated area. This is regrettable as species richness is an elementary measure of

biodiversity that underlies many ecological models and conservation strategies. By recording not just species but species

per unit of search effort, an extended dataset results which can be used to generate estimates of total species richness. In

this study, the herpetofauna of Pangkor Island, Peninsular Malaysia is used as an example. In 2009 and 2010, the first

herpetological surveys were carried out on this small, 18 km2, island. Those surveys recorded 43 species of reptiles and

13 species of amphibians. In this study, total reptile species richness was estimated by fitting several models to the sam-

ple-based rarefaction curve as well as by application of the nonparametric Chao-I estimator. Of the applied models, the

4-parameter Weibull function was shown to be superior, a finding that is in line with several other studies. Consequent-

ly, the use of this model is recommended. On the basis of the fitted 4-parameter Weibull-function, 69 reptile species are

expected to occur on Pangkor Island. As for amphibians, total species richness was estimated to be 17. As such, a re-

markably extensive herpetofaunal assemblage inhabits this small island.

Key words. Pangkor Island, Malaysia; amphibians, reptiles, species-richness, sample-based rarefaction curve, negative

exponential function, 3-parameter Weibull function, 4-parameter Weibull function, Chapman-Richards model, Chao-I es-

timator

INTRODUCTION

Herpetological surveys of species-rich tropical ecosystems

rarely result in complete inventories (e.g. Lloyd et al.

1968; Murphy et al. 1994; Hofer & Bersier 2001; Van

Rooijen 2009). This is due to the asymptotical nature of

the species accumulation process in combination with lim-

ited survey-investments. As a consequence, the use of es-

timation techniques is unavoidable when the intended ob-

jective is to assess species richness, an elementary meas-

ure of biodiversity that underlies many ecological mod-

els and conservation strategies. One such estimation tech-

nique consists of fitting an appropriate function to the

species accumulation curve. The asymptote of the fitted

functions can then be regarded as an estimate of total

species richness. Species accumulation curves are regu-

larly applied in herpetofaunal surveys but they are most-

ly used to arrive at qualitative judgements about the

exhaustiveness of the survey (e.g. Murphy et al. 1994; Zug

et al. 1998; Ziegler 2002). However, extensive and sophis-

ticated literature exists pertaining to methods and models

used to quantitatively estimate species richness, on the ba-

sis of either rarefied species accumulation curves or abun-

dance patterns (e.g. Colwell & Coddington 1994; Flather

1996; Gotelli & Colwell 2001; Longino et al. 2002; Brose

et al. 2003).

The offshore archipelagos of Peninsular Malaysia have

been the subject of increasing interest in recent years.

Many of these islands have never been surveyed and re-

cent explorations are only beginning to uncover the hid-

den diversity and endemism that they shelter (Chan et al.

2009a; Grismer 2008; Grismer & Norhayati 2008; Gris-

mer & Pan 2008; Grismer et al. 2008, 2009a, b). One such

4 Johan van Rooien et al.

Fig. 1.

al., 2010; D: Dryophiops rubescens (Gray, 1835).

island is Pulau Pangkor, approximately 18 km2 and situ-

ated 3.5 km from the west coast of Peninsular Malaysia.

The island’s interior remains heavily forested and main-

tains a river system, Sungai Pinang, which supplies a sig-

nificant source of permanent fresh water in the form of

multiple streams. Chan et al. (2010) provided the first re-

port on the (non-marine) herpetofauna of this island and

documented 43 reptiles and 13 amphibians. A few illus-

trations are provided in figures | and 2.

The study described in this paper had a dual objective.

First, data collected by Chan et al. (2010) were used to

evaluate the performance of several estimators. Second,

the most appropriate estimator was used to estimate the

total herpetofaunal species richness harboured by Pangkor

Island, West Malaysia.

MATERIAL AND METHODS

The data underlying this study were based on the surveys

carried out by Chan et al. (2010). These were conducted

in the periods March 15—17, May 4-8, June 13 to July 8,

2009, and February 22 to March 8, 2010. During the lat-

ter two periods, only reptiles were surveyed. Marine rep-

Bonn zoological Bulletin 60 (1): 3-8

A: View into the primary forest of Pangkor Island; B: Boiga drapiezii (Bote, 1827); C: Cnemaspis shahruli Grismer et

tiles were ignored altogether. The predominantly applied

survey method corresponds to visual encounter survey

(VES), a simple method which has been shown to be ef-

fective for surveying rainforest herpetofauna (Doan,

2003). VES was carried out both during day and night (e.g.

Coddington et al., 1996; Doan, 2003) in a way similar to

that applied by Minh (2007). Existing trails as well as sev-

eral trails made by the Department of Forestry of Perak

and Peninsular Malaysia were used. These trails traversed

dipterocarp forest, mangrove forest and cultivated areas

and provided ample access to forest streams. The second

collection method consisted of searching for road-kills.

The third entailed turning logs, fallen tree bark and sim-

ilar objects in order to uncover animals hiding underneath.

Sampling effort was expressed in terms of search-days

where one search-day was defined as roughly 4.5 search-

hours. As two of the surveys underlying the data collect-

ed by Chan et al. (2010) focused solely on reptiles, sub-

stantially more information was available regarding rep-

tiles allowing for more thorough analyses. Therefore, this

study focuses predominantly on reptiles, amphibians be-

ing treated separately and in less detail.

Sample-based rarefaction curves (Gotelli & Colwell 2001)

were generated with the program EstimateS (Colwell

©OZFMK

Estimating herpetofaunal species richness

Fig. 2. A: Heosemys spinosa (Gray, 1831); B: Kaloula pulchra Gray, 1831; C: Naja sumatrana (Miller, 1890); D: Tropidolae-

mus wagleri (Boie, 1827).

2005). Four models were fitted to the rarefaction curve.

The first corresponds with a negative exponential model

(Colwell & Coddington 1994; Flather 1996; Van Rooyen

2009). It is based on the assumption that the number of

new species found per search day is proportional to the

number of as yet undiscovered species, in mathematical

terms: dY/dt = c (A—Y) where A is the total number of

species present in the area under investigation, Y is the to-

tal number of species found and c is a constant. This equa-

tion can be represented as a negative exponential function

(e.g. Van Rooyen 2009): Y= A (1—-e~*'). The basic assump-

tion underlying the negative exponential model (hence-

forth NE) may be overly simplistic given that abundance

patterns are usually strongly skewed (e.g. Lloyd et al.

1968; Coddington et al. 1996; Limpert et al. 2001; Longi-

no et al. 2002; Thompson et al. 2003). In order to model

more complex species accumulation processes, the NE can

be refined in various ways by adding one (d) or two pa-

rameters (d and p), resulting in the Chapman-Richards

model (henceforth CR), 3- and 4-parameter Weibull cu-

mulative distribution functions (henceforth 3pW and

4pW):

Y =A (1-e«t)* (CR),

Y=A (1-e()“) pW),

Y= A (1-eep)) (4pW)

Bonn zoological Bulletin 60 (1): 3-8

The four models were fitted to the sample-based rarefac-

tion curve using nonlinear regression analysis (e.g. Noru-

sis and SPSS 1994) with SPSS (release 14 February 1996;

SPSS Inc.).

Extrapolation using different models for the species ac-

cumulation process can provide different asymptotes and

thus predict different values of species richness (e.g. Col-

well & Coddington 1994; Flather 1996). Therefore, care

has to be taken to select the most appropriate model in or-

der to minimize bias. In this study, the appropriateness of

each model was evaluated on the basis of three criteria.

The first criterion was goodness-of-fit. The second crite-

rion entailed the behaviour of the richness-estimate with

increasing cumulative search effort. The final criterion was

the difference with the nonparametric Chao-I estimator

(Chao 1984; Coddington et al. 1996; Hofer & Bersier

2001; Veith et al. 2004). This estimator is an often applied

representative of a class of estimators that uses a differ-

ent approach as they are based on abundance patterns in-

stead of the accumulation curve. The Chao-I estimator is

based on the observed number of rare species,

A = Y + (a?/2b), where A is the total number of species,

Y is the observed number of species, a is the number of

observed species represented by a single specimen and 6

OZFMK

6 Johan van Rooijen et al.

is the number of observed species represented by exact-

ly two specimens.

RESULTS

Figure 3 depicts the deviations of the fitted functions from

the sample-based rarefaction curve (residuals) for reptiles.

As expected, the NE exhibits a very poor goodness-of-fit.

At first, the fitted curve is situated beneath the rarefaction

curve, then above and finally drops beneath the rarefac-

tion curve again. The suboptimal fit of the NE as well as

the pattern of residuals are in agreement with results ob-

tained with regard to avian diversity (Flather 1996) as well

as snake diversity (Van Rooien 2009). Although the RC

and 3pW fit substantially better than the NE, they exhib-

it a similar pattern of residuals. The 4pW function obvi-

ously exhibits a superior, near-perfect, fit.

residual

QRROOS negative exponential

: —— 3-parameter Weibull

-2,5 Miyake - 7 > 4-parameter Weibull

——— Chapman-Richards

search days

Fig. 3. | Deviations of the fitted models from the sample-based

rarefaction curve (residuals).

Figure 4 depicts the sample-based rarefaction curve as well

as the fitted 4pW. The only noticeable deviation of the fit-

ted 4pW from the rarefaction curve lies in the fact that it

passes through (p,0) instead of the origin (0,0). The close

fit of the 4pW is in agreement with results obtained in

studies of avian diversity (Flather 1996), snake diversity

(Van Rooijen 2009) and diversity of small reptiles

(Thompson et al. 2003). The progression of the rarefac-

tion curve clearly indicates that an asymptote has not yet

been reached, thus the surveys have not been exhaustive.

Figure 5 shows how the richness estimates (as opposed

to observed richness or fitted model-values) develop with

increasing search days during the second half of the sur-

vey. The NE is neglected due to its poor fit. Evidently, the

3pW- as well as CR-based estimates are still increasing

at the end of the survey whereas the 4pW-based estimate

Bonn zoological Bulletin 60 (1): 3-8

acne

cumulative number of species

0 1 ZS OO. Fi ACI 9) AOR A 14 201314516 fa Seto m2 mee

cumulative number of search days

Fig. 4. Reptile species sample-based rarefaction curve with

fitted 4-parameter Weibull function.

has reached a more or less stable level. A similar result

(unpublished) was obtained on the basis of a dataset that

underlies a study of Bornean snakes (Van Rooijen 2009):

the 4pW-based estimate reached an approximately stable

plateau rather early with the CR- and 3pW-based estimates

approaching the 4pW-based estimate with increasing

search days.

On the basis of these results, the 4pW-based estimate, 69,

is assumed to be the least biased. Finally, the estimate

based on the nonparametric Chao I estimator, 62, agrees

reasonably well with the 4pW-based estimate. Taking the

average, 65 reptile species are expected to inhabit

Pangkor Island. As for amphibians species richness, the

fitted 4pW and Chao I estimator both resulted in an esti-

mate of 17 species.

2): ete cio cho OO OO OP CLO OOOO OOOO oo aoc o mS

65 |

60 4

@ 504

ze |

B ao

g 35 4

2 304

2 254

20 4

1) 3-parameter Weibull

10 * = * 4-parameter VWeibull

5 | —— Chapman-Richards |

0 oot : ——————

10 1 12 13 14 15 16 7 18 419 20 21

cumulative number of search days

Fig. 5. Richness estimates in relation to cumulative search ef-

fort.

©OZFMK

Estimating herpetofaunal species richness a/

DISCUSSION

Estimation by extrapolation, as opposed to interpolation,

into the unknown entails a high degree of uncertainty.

However, the alternative, surveying until completion, is

simply unfeasible when the survey is aimed at the herpeto-

fauna of a Southeast Asian rainforest ecosystem (Van

Rooijen 2009). As such, there simply is no alternative.

Thus, further refinement of estimation methods is of im-

portance. That said, approximately complete species lists

are obviously still crucial for many zoogeographic stud-

ies. Such species lists can be (and are) compiled by com-

bining results of various surveys. However, this usually

entails the combination of species lists pertaining to dif-

ferent locations. As such, completeness comes at a price

as one has to accept a huge decrease in spatial resolution.

In most zoogeographic studies however, a high resolution

is not essential as species compositions of major land

masses are compared (e.g. In Den Bosch 1985; How and

Kitchener 1997; Inger and Voris 2001). At the spatial scale

which is relevant for conservation though, complete

species lists are a utopia in the majority of cases and es-

timation will therefore gain in importance (Van Rooijen

2009). This study underscores the notion that a combina-

tion of criteria needs to be applied to select the most ap-

propriate estimator. The 4-parameter Weibull function ex-

hibited the highest goodness-of-fit. Moreover, it produced

the most stable estimate near the end of the survey (fig-

ure 5). Finally, it resulted in a richness-estimate that agreed

well with the estimate based on the nonparametric Chao-

I estimator. Even then, the estimate may be downwardly

biased due to suboptimal sampling. First of all, although

searches were carried out during both day and night, no

searches were performed in the early morning (before

dawn). Secondly, sampling of canopy-microhabitat was

obviously unfeasible. Finally, as true species richness

is underestimated by most estimators when sample size

is small (Colwell & Coddington 1994; Canning-Clode et

al. 2008), the estimate of amphibian species richness may

be downwardly biased since it was based on only five

search days. Nevertheless, the amphibian species richness

appears to be rather low which is in agreement with the

impression of herpetologists who carried out the surveys

(Chan et al. 2010).

The fact that the 4-parameter Weibull function exhibits

such a good fit when applied to diverse ecological com-

munities such as small reptiles in desert habitat (Thomp-

son et al. 2003), birds (Flather 1996) and reptiles in rain-

forest habitat (Van Rooijen 2009 and this report) is strik-

ing. Two explanations can be put forward. First, relative

species abundances follow very similar patterns over a

wide range of ecological communities: relatively few

species are abundant whereas most are rare (e.g. Hughes

1986). Thus, the shape of the species accumulation process

Bonn zoological Bulletin 60 (1): 3—8

may also be expected to be rather uniform. Second, the

higher the number of parameters of a model, the better the

fit. Thus, the 4pW may simply be expected to fit better

than similar functions with fewer parameters, irrespective

of the field of application.

65 reptilian species are estimated to inhabit Pangkor Is-

land, 43 of which have been recorded (Chan et al. 2010).

A major part of the as yet unrecorded species will con-

cern snakes as members of this taxon are notoriously hard

to find due to their elusive habits and low densities (e.g.

Lloyd et al. 1968; Inger & Colwell 1977; Hofer & Bersi-

er 2001; Orlov et al. 2003; Van Rooijen 2009). This no-

tion is strengthened by several indirect observations. Lo-

cals provided accounts regarding observations of Boiga

dendrophila (Bote, 1827), Cryptelytrops purpureomacu-

latus (Gray, 1832), Ophiophagus hannah (Cantor, 1836)

and Maticora bivirgata (Boie, 1827). In addition, there has

been a visual record of a Coelognathus radiatus (Boie,

1827) (Schultz, pers. comm.). As amphibian species rich-

ness is estimated to be at least 17, herpetofaunal species

richness is expected to equal at least 82 species, which is

quite impressive given the fact that Pangkor Island encom-

passes merely 18 km?. On the other hand, whether

Pangkor Island harbours a comparatively extensive her-

petofauna is impossible to determine at the moment as

species richness on other Malaysian islands has not yet

been estimated.

Estimating species richness of specific sites such as islands

obviously has added value as species richness is an ele-

mentary criterion a conservationist may use when select-

ing sites and is crucial for many ecological studies. How-

ever, the function fitted to the rarefaction curve not only

provides an estimate of total species richness but also pro-

vides insight in expected return on further investment: how

many previously unrecorded species may be expected to

be found with additional search effort? For instance, on

the basis of the fitted 4-parameter Weibull function, ten

previously unrecorded reptiles are expected to be found

on Pangkor Island when 20 additional search days are in-

vested. Such statistical expectations can be used as input

for a cost-benefit evaluation when choices have to be made

between different sites for the investment of survey-ca-

pacity. Alternatively, one might determine how much

search effort would have to be invested to bring the sur-

vey to some specified level of completeness.

REFERENCES

Brose U, Martinez ND, Williams RD (2003) Estimating species

richness: sensitivity to sampling coverage and insensitivity to

spatial patterns. Ecology 84: 2364-2377

Canning-Clode J, Valdivia N, Molis M, Thomason JC, Wahl M

(2008) Estimation of regional richness in marine benthic com-

©ZFMK

8 Johan van Rooyen et al.

munities: quantifying the error. Limnology and Oceanogra-

phy: Methods 6: 580-590

Chan KO, Grismer LL, Wood PL Jr, Grismer JL, Norhayati A

(2009) Preliminary Checklist of the Herpetofauna of Pulau Be-

sar, Melaka, Malaysia. Tropical Life Sciences Research, 20(1):

81-87

Chan KO, Van Rooyen J, Grismer LL, Daicus B, Akil MAMM,

Jamaludin H, Gregory R, Norhayati A (2010) First report on

the herpetofauna of Pulau Pangkor, Perak, Malaysia. The

Russian Journal of Herpetology 17: 139-146

Chao A (1984) Non-parametric estimation of the number of

classes in a population. Scandinavian Journal of Statistics 11:

265-270

Coddington JA, Young LH, Coyle FA (1996) Estimating spider

species richness in a southern Appalachian cove hardwood for-

est. The Journal of Arachnology 24: 111-128

Colwell RK, Coddington JA (1994) Estimating terrestrial bio-

diversity through extrapolation. Philosophical Transactions of

the Royal Society of London 345: 101-118

Colwell RK (2005) EstimateS: Statistical estimation of species

richness and shared species from samples. Version 7.5. User’s

Guide and application published at: http://purl.oclc.org/esti-

mates

Doan TM (2003) Which methods are most effective for survey-

ing rainforest herpetofauna? Journal of Herpetology 37: 72-81

Flather CH (1996) Fitting species-accumulation functions and

assessing regional land use impacts on avian diversity. Jour-

nal of Biogeography 23: 155-168

Gotelli NJ, Colwell RK (2001) Quantifying biodiversity: pro-

cedures and pitfalls in the measurement and comparison of

species richness. Ecology letters 4: 379-391

Grismer LL (2008) A new species of insular skink (Genus Sphe-

nomorphus Fitzinger 1843) from the Langkawi Archipelago,

Kedah, West Malaysia with the first report of the herpetofau-

na of Pulau Singa Besar and an updated checklist of the her-

petofauna of Pulau Langkawi. Zootaxa 1691: 53-66

Grismer LL, Chan KO, Grismer JL, Wood PL Jr., Daicus B

(2008) Three new species of Cyrtodactylus (Squamata:

Gekkonidae) from Peninsular Malaysia. Zootaxa 1921: 1—23

Grismer LL, Pan KA (2008) Diversity, endemism and conser-

vation of the amphibians and reptiles of Southern peninsular

Malaysia and its offshore islands. Herpetological Review

39(3): 270-281

Grismer LL, Norhayati A (2008) A new insular species of Cyr-

todactylus (Squamata: Gekkonidae) from the Langkawi Ar-

chipelago, Kedah, Peninsular Malaysia. Zootaxa 1924: 53-68

Grismer LL, Wood PL Jr., Grismer JL (2009a) A New Insular

Species of Skink of the Genus Sphenomorphus Strauch 1887

(Squamata: Scincidae) from Pulau Perhentian Besar, Tereng-

ganu, Peninsular Malaysia. Tropical Life Sciences Research

20(1): 51-69

Grismer LL, Norhayati A, Chan KO, Daicus B, Muin, MA,

Wood PL Jr., Grismer JL (2009b) Two new diminutive species

of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from

Peninsular Malaysia. Zootaxa 2019: 40-56

Hofer U, Bersier L-F (2001) Herpetofaunal diversity and abun-

dance in tropical upland forests of Cameroon and Panama.

Biotropica 33: 142-152

Bonn zoological Bulletin 60 (1): 3-8

How RA, Kitchener DJ (1997) Biogeography of Indonesian

snakes. Journal of Biogeography 24: 725-735

Hughes RG (1986) Theories and models of species abundance.

American Naturalist 128: 879-899

In Den Bosch HAJ (1985) Snakes of Sulawesi: checklist, key

and additional biogeographical remarks. Zoologische Verhan-

delingen 217: 1-50

Inger RF, Colwell RK (1977) Organization of contiguous com-

munities of amphibians and reptiles in Thailand. Ecological

Monographs 47: 229-253

Inger RF, Voris HK (2001) The biogeographical relations of the

frogs and snakes of Sundaland. Journal of Biogeography 28:

863-89 1

Limpert E, Stahel WH, Abbt M (2001) Log-normal distributions

across the sciences: keys and clues. Bioscience 51: 341-352

Lloyd M, Inger RF, King FW (1968) On the diversity of reptile

and amphibian species in a Bornean rain forest. The Ameri-

can Naturalist 102: 497-515

Longino JT, Coddington J, Colwell RK (2002) The ant fauna of

a tropical rainforest: estimating species richness three differ-

ent ways. Ecology 83: 689-702

Minh NOM (2007) Estimates of the herpetofaunal diversity in

the Shipstern Nature Reserve, Belize. Biology master degree

thesis, Zoological Institute, University of Neuchatél, Switser-

land

Murphy JC, Voris HK, Karns DL (1994) A field guide to the

snakes of the Danum Valley, a Bornean tropical rainforest

ecosystem. Bulletin of the Chicago Herpetological Society 29:

133-151

Norusis & SPSS Inc. (1993) SPSS Advanced Statistics, SPSS,

Chicago

Orlov NL, Ryabov SA, Nguyen Van Sang, Nguen Quang Truong

(2003) New records and data on the poorly known snakes of

Vietnam. Russian Journal of Herpetology 10: 217—240

Thompson GG, Withers PC, Pianka ER, Thompson SA (2003)

Assessing biodiversity with species accumulation curves; in-

ventories of small reptiles by pit-trapping in Western Australia.

Austral Ecology 28: 361-383

Van Rooijen J (2009) Estimating the snake species richness of

the Santubong Peninsula (Borneo) in two different ways. Con-

tributions to Zoology 78: 141-147

Veith M, Loétters S, Andreone F, Rédel M-O (2004) Measuring

and monitoring Amphibian diversity in tropical forests. II. Es-

timating species richness from standardized transect censing.

Ecotropica 10: 85-99

Ziegler T (2002) Die Amphibien und Reptilien eines Tiefland-

feuchtwald-Schutzgebietes in Vietnam. Minster: Natur und

Tier- Verlag

Zug GR, Win H, Thin T, Min TZ, Lhon WZ, Kyaw K (1998)

Herpetofauna of the Chatthin Wildlife Sanctuary, North-Cen-

tral Myanmar with preliminary observations of their natural

history. Hamadryad 23: 111-120

Received: 21.06.2010

Accepted: 14.03.2011

Corresponding editor: F. Herder

OZFMK

Bonn zoological Bulletin Volume 60 Issue | pp. 9-16 Bonn, May 2011

New records of snakes from Cat Tien National Park,

Dong Nai and Lam Dong provinces, southern Vietnam

Peter Geissler '!, Truong Quang Nguyen !, Nikolay A. Poyarkov 34 & Wolfgang Bohme !

| Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany;

E-mails: pgeissler84@yahoo.de; w.boehme.zfmk(@uni-bonn.de

2 Institute of Ecology and Biological Resources, 18 Hoang Quoc Viet, Hanoi, Vietnam;

E-mail: nqt2@yahoo.com

3 Lomonosov Moscow State University, Biological faculty, Department of Vertebrate Zoology,

Leninskiye Gory, Moscow, GSP-1, 119991, Russia

4 Joint Russian-Vietnamese Tropical Research and Technological Center of the A.N. Severtsov Institute of

Ecology and Evolution, South Branch, 3, Street 3/2, 10 District, Ho Chi Minh City, Vietnam;

E-mail: n.poyarkov@gmail.com.

Abstract. We report ten new records of snakes from Cat Tien National Park, Dong Nai and Lam Dong provinces, south-

ern Vietnam. The specimen of Zyphlops siamensis Gunther, 1864 from Cat Tien represents the second country record in

Vietnam after more than one century. Our new record of Dendrelaphis ngansonensis (Bourret, 1935) leads to a consid-

erable range extension of this species, so far known only from northern and central Indochina. Additional specimens of

Oligodon deuvei David, Vogel & van Rooijen, 2008 are described, along with the first photograph of a living specimen,

showing the coloration including the aposematically red-coloured underside of the tail. An updated checklist of snakes

of the Cat Tien National Park is also provided.

Key words: Squamata: Serpentes: Typhlopidae, Colubridae, Cat Tien National Park.

INTRODUCTION

Among the 545 species of reptiles and amphibians known

from Vietnam, snakes have the highest species diversity

(Nguyen et al. 2009). However, many snake species are

often recorded only by a single or rather few specimens,

and data on their distribution and natural history are still

deficient. Though the Cat Tien National Park is the biggest

and most important National Park for the lowland rain-

forests of southern Vietnam (Polet & Ling 2004), its her-

petofauna is still poorly studied (Nguyen 1988, Le et al.

1998, Nguyen & Ho 2002, Le 2007, Geissler et al. 2009).

The most recent checklist of reptiles and amphibians of

the Cat Tien National Park, which was compiled by

Nguyen & Ho (2002), reported a total of 42 snake species.

We herein provide an updated checklist of snakes from this

national park with ten new records and discussion about

the status of some rare and poorly known species in Viet-

nam.

MATERIAL AND METHODS

Field work was conducted by Peter Geissler in July and

August 2008 and from March until June 2009; by Niko-

lay A. Poyarkov from November until December 2007,

from February until April 2008, from February to April

2009, and in July 2010 in Cat Tien National Park, Dong

Nai Province, southern Vietnam (Fig. 1). Species identi-

fications were also based on the examination of specimens

collected by colleagues from the Appalachian State Uni-

versity (North Carolina, USA). A total of 54 specimens

were examined and they were subsequently deposited in

the collections of the Institute of Ecology and Biological

Resources (IEBR), Hanoi, Vietnam; the Zoologisches

Forschungsmuseum Alexander Koenig (ZFMK), Bonn,

Germany; and the Zoological Museum, Moscow State

University (ZMMU), Moscow, Russia.

The following measurements were taken with a digital

vernier calliper: SVL (snout-vent length); TL (total

10 Peter Geissler et al.

East Sea

(South China Sea)

VI2°€

2D,

len Seat

Paracel Island

x 114€

Elevation (m) |. FI

WF 1200+

Tt Gulf of 4

0: 2 (7S 150

R+—+ +S

Kilometers

Ship teens tail

Spratly,islonds + | *

Thailand 2 Sa hee

ailond f : pe pms

0 150 300 Kilometors

aire

Fig. 1. | Map showing the study site (black diamond) in south-

ern Vietnam: Cat Tien National Park.

length); TaL (tail length). In addition, several scale counts

were taken: VEN (number of ventrals); MD (middorsal

scales); MBS (midbody scale rows, including the ven-

trals); SC (subcaudals); IL (infralabials); SL (supralabi-

als).

TAXONOMIC ACCOUNT

Typhlops siamensis Giinther, 1864

Specimen examined: One adult male (ZFMK 88922),

collected by P. Geissler on the road in a rainy evening on

31 March 2009, at the Headquarters of Cat Tien Nation-

al Park, Dong Nai Province (near 11°25719.3”N,

107°25°42.0”E, 104 m a.s.l.).

Characteristic features: The morphological features of

this specimen fit the descriptions of Giinther (1864) and

Bonn zoological Bulletin 60 (1): 9-16

Wallach (2001): SVL 140.2 mm; TL 4.4 mm; preocular

separated from nasal; inferior nasal suture contacting sec-

ond supralabial; superior nasal suture extending horizon-

tally to rostral, not visible from above; 22 scale rows

around midbody (including ventral scale rows); 308 par-

avertebral scales (306-368 according to Wallach 2001);

dorsum uniformly dark brown, venter cream (see Fig. 2a).

Remarks: Wallach (2001) limited the distribution of this

species to Thailand and Cambodia, although it was record-

ed from Vietnam by Nguyen & Ho (1996), see also

Nguyen et al. (2009). The record is based on a specimen

(ZISP 5426), which was collected by Tramond in 1879 and

subsequently deposited in the collection of the Zoologi-

cal Institute in St. Petersburg (Tirant 1885). Therefore, our

record from Cat Tien National Park represents a rediscov-

ery of this species in Vietnam after 130 years. In main-

land Southeast Asia, both Typhlops diardi Schlegel, 1839

and Zyphlops muelleri Schlegel, 1839 are morphological-

ly similar to Zyphlops siamensis. T: siamensis differs from

T: diardi in having a lower number of scale rows around

midbody (22 vs. 24-28). Bourret (1936) synonymized 7:

siamensis with the subspecies 7. diardi nigroalbus

Dumeril & Bibron, 1844. Typhlops diardi nigroalbus was

recently synonymized with Zyphlops muelleri Schlegel,

1839 by Wallach (2001). 7: siamensis can be distinguished

from 7. muelleri by having a lower count of MBS (22 vs.

24-30) and by the absence of a sharply bounded yellow

ventral surface. However, 7. muelleri was not included in

recent lists of the snake fauna of Vietnam (Ziegler et al.

2007, Nguyen et al. 2009). We thus follow Wallach (2001)

to recognize 7’ siamensis as a distinct species. Future re-

search on 7yphlops from southern Vietnam is required to

show whether 7: muelleri actually occurs in Vietnam as

affirmed by Wallach (2001) or these records are based on

misidentified specimens of the closely related 7: siamen-

sis.

Boiga multomaculata (Boie, 1827)

Specimen examined: One adult male (ZFMK 88923),

collected by K. D. Le in August 2008 in the bamboo for-

est, Cat Loc area, Lam Dong Province (near 11°37’22.5”N

107-17 5SI24Es 3 5emiarsi):

Characteristic features: The morphological features of

this specimen agree with the descriptions of Bourret

(1936), Smith (1943), and Campden-Main (1970): SVL

588 mm; TaL 158 mm; | loreal; | preocular; 2 postocu-

lars; 8 SL; 10 IL; 19 MD; 215 VEN; 93 SC; head with

two dorsal dark brown stripes, from snout to neck. For

colouration in life see Fig. 2b.

©ZFMK

New records of snakes from southern Vietnam 1]

Fig. 2.

Calamaria pavimentata Dumeril & Bibron 1854

Specimen examined: One subadult specimen (ZFMK

88924), collected by K. D. Le in June 2008, in Cat Loc

area, Lam Dong Province (near 11°37’22.5”N

NO fila ie 2 E135 mais):

Characteristic features: The morphological features of

this specimen fit the descriptions of Bourret (1936), Smith

(1943), and Ziegler et al. (2007): SVL 208 mm; TaL 15.9

mm; snout obtuse; no supranasals; no loreals; no tempo-

rals; 13 MD (smooth); 158 VEN; 17 SC; dorsum dark

brown or grey; 6 black dorsolateral stripes; yellow collar,

narrowing dorsally; tail with 2 yellow cross bands. For

colouration in life see Fig. 2 c.

Remarks: This is the southernmost record of this species

from Vietnam (compared with Nguyen et al. 2009).

Bonn zoological Bulletin 60 (1): 9-16

a. Typhlops siamensis (ZFMK 88922); b. Boiga multomaculata (ZFMK 88923); ¢. Calamaria pavimentata (ZFMK 88924);

d. Coelognathus flavolineatus (ZFMK 88898). Photographs: E. Galoyan & P. Geissler.

Coelognathus flavolineatus (Schlegel, 1837)

Specimen examined: One adult female (ZFMK 88898),

collected by P. Geissler in April 2009, on the forest floor,

in a mixed forest near Bau Sau Lake, Cat Tien National

Park, Dong Nai Province (near 11°27°32.9"N

107°20’43.7”E, 167 ma.s.l.).

Characteristic features: The morphological traits of this

specimen agree with the descriptions of Bourret (1936)

and Smith (1943): SVL 1,307 mm; TaL 374 mm; | elon-

gated loreal; 1 preocular; 2 postoculars, 8 SL; 10 IL; 19

MD, keeled; 212 VEN; 96 SC, divided; dorsum brown,

with a orange, black edged vertebral stripe; vertebral stripe

desappearing in the posterior half of body, which is uni-

formly dark brown. For colouration in life see Fig. 2d.

©ZFMK

Fig. 3.

(juvenile, ZFMK 91899); d. Oligodon deuvei(ZMMU NAP-02811); e. Oligodon ocellatus from Cat Tien National Park, Dong Nai

Province (e); f. Xenochrophis flavipunctatuss (ZFMK 88914). Photographs: E. Galoyan, P. Geissler, W. Van Devender.

Dendrelaphis ngansonensis (Bourret, 1935)

Specimens examined: One adult male (ZFMK 88913),

collected by P. Geissler on 22 July 2008, in a secondary

forest near Bau Sau Lake, Cat Tien National Park, Dong

Nai Province (near 11°27°32.9’N, 107°20°43.7”E, 160

m a.s.l.). It was found in the morning, basking on a sun-

ny spot on the lava-rock-covered forest floor.

Bonn zoological Bulletin 60 (1): 9-16

12 Peter Geissler et al.

a. Dendrelaphis ngansonensis (ZFMK 88913); b. Lycodon subcinctus (adult, IEBR A.2010.42); ¢. Lycodon subcinctus

Characteristic features: The characters of the specimen

fit the descriptions given by Bourret (1935, 1936) and

Ziegler & Vogel (1999): SVL 824 mm; TaL 419 mm; 25

dentary teeth; 15 MD; 188 VEN (165-199 in the descrip-

tion of Ziegler & Vogel 1999); 153 SC; vertebral dorsals

enlarged; dorsum bronze-brown; presence of a distinct

black stripe from the posterior margin of the eye to the

neck; scales on dorsilateral neck and anterior part of body

with bluish margin. For colouration in life see Fig. 3a.

©ZFMK

New records of snakes from southern Vietnam 13

Remarks: According to Ziegler & Vogel (1999) and

Nguyen et al. (2009) this species has been known only

from northern and central Vietnam and Quang Nam

Province has been known to be the southernmost locali-

ty of the species’ range. Our record from Cat Tien Nation-

al Park extends the known distribution about 500 km

southwards to the lowland forests of southern Vietnam (see

Fig. 3). The distribution gap seen in the map leads to the

suggestion that D. ngansonensis 1s also present on or along

the Central Highlands as well as the Da Lat Plateau. Like

in other localities in Vietnam (Ziegler & Vogel 1999), Den-

drelaphis pictus (Gmelin, 1789) can be found in the same

habitats with its congener, affirming the species status of

D. ngansonensis, which was originally considered as a

subspecies of D. pictus.

Lycodon subcinctus Boie, 1827

Specimens examined: One adult male IEBR A.2010.42),

collected by W. Van Devender in June 2004, near the head-

quarters of Cat Tien National Park, Dong Nai Province

(near 11°25°19.3”°N, 107°25’42.0”E, 104 m a.s.1.). One

juvenile (ZFMK 91899), collected by W. Van Devender

on 9 June 2004, near Suoi Rang Ranger Station, Cat Tien

National Park, Dong Nai Province. One juvenile (IEBR

A.2010.43), collected by W. Van Devender in 2006 in Cat

Tien National Park, Dong Nai Province.

Characteristic features: The morphological traits of the

specimens (data given in the following order: IEBR

A.2010.42/ZFMK 91899/IEBR A.2010.43) fit the descrip-

tions of Smith (1943); Daltry & Wiister (2002), and

Ziegler et al. (2007): SVL 651 mm / 276 mm / 228 mm;

TaL 159 mm / 65 mm / 55 mm); head with broad blunt

snout; preocular absent; prefrontal and loreal in contact

with the eye; 8 SL, 4 suboculars; 17 MD, smooth;

201/202/201 VEN; precloacal scale divided; 71/72/87 SC,

divided; juveniles black above with white cross bands

(Fig. 3c); adult darker, cross bands absent on posterior part

of body (Fig. 3b).

Oligodon deuvei David, Vogel & van Rooijen, 2008

Specimens examined: One juvenile specimen (IEBR

A.2010.16), collected by local people on 29 May 2004.

An adult male (IEBR A.2010.17), collected by P. Moler

in May 2004. Another adult male (ZFMK 91226), collect-

ed by P. Moler in 2005. These specimens were found near

the Headquarters of Cat Tien National Park, Dong Nai

Province (near 11°27°32.9"N, 107°20’°43.7”E). One

adult male (ZMMU NAP-02811), collected by N. Po-

Bonn zoological Bulletin 60 (1): 9-16

yarkov on 8 November 2007, in Cat Tien Village on the

eastern bank of the Dong Nai River, Dong Nai Province.

Characteristic features: The morphological characters of

four specimens from Cat Tien National Park agree with

the description given by David et al. (2008b): SVL

283-361 mm; TaL 59.0—68.2 mm; 17 MD, 15 MD before

vent; 14 maxillary teeth, the posterior two strongly en-

larged; 8—9 IL; 8 SL; no presubocular; a conspicuous, pale

yellow vertebral stripe, edged with two darker faint par-

avertebral stripes or with lines of dark dots in the paraver-

tebral region; 4-5 markings on dorsal head surface: one

transverse band across the snout, one sagital blotch be-

tween the orbits, two streaks behind the orbit, directed pos-

teriorly downwards, and one broad arrow shaped blotch

on the neck. In one specimen (ZMMU NAP-02811), three

nuchal blotches are fused to one butterfly-shaped mark-

ing, and a number of small irregular dots are present on

the forehead. Measurements and selected scale counts, in

comparison with data of the type specimens provided by

David et al. (2008b), are given in Table 1.

Colouration: In accordance with Deuve’s (1985) unpub-

lished manuscript, our specimens have a grey dorsum. The

yellow vertebral stripe is broad on the neck, narrowing

backwards, and edged by two dark brown paravertebral

stripes. Though not mentioned by Deuve (1985) or David

et al. (2008b) our specimens have a dark grey dorsolater-

al stripe (aligned dark brown dots in preserved specimens).

Large dark blotches are present on upper tail surface in

the holotype but they are lacking in our specimens. Ven-

ter and lower surface of the tail are pinkish red with nu-

merous rectangular and subrectangular blackish-brown

spots. These spots are lacking in the posterior half of the

tail (Fig. 3d).

Remarks: Here we provide the first record for the Cat

Tien National Park and the second collection of the species

after the description of David et al. (2008b). Our data sup-

port the sexual dimorphism observed by David et al.

(2008b). Due to the lower number of subcaudals and the

shorter tail length, we assume that the juvenile specimen

(IEBR A.2010.16) is a female. The specimens in our col-

lection slightly differ from the type series by having a

higher TL/TaL ratio in males and by having eight supral-

abials (this character only occurs in one of 17 specimens

examined by David et al. 2008b). For the first time a

colour photograph of a living specimen is provided show-

ing the coloration of the dorsum as well as the red under-

side of the tail, which is used by the snake for defensive

display (Fig. 3d). Our observations in the field indicate

that this snake species was mostly active during twilight,

but was twice observed actively foraging at day time. O.

deuvei was found along the riverside and neighbouring ru-

ral areas. The stomach of ZMMU NAP-02811 contained

©ZFMK

14 Peter Geissler et al.

Table 1. Measurements and selected scale counts of Oligodon deuvei specimens from Cat Tien National Park in comparison with

the type series (for abbreviations see Material and Methods).

Type specimens JEBR A.2010.17 ZFMK 91226 ZMMU NAP-02811 IEBRA. 2010.16 (juv.)

(David et al. 2008) (3) (3) (3) (3)

TL (mm) up to 333

535 (2) 342 384.2 328.5 108.8

SVL (mm) up to 275(¢)

302(2) 283 316 271 95.1

TaL (mm) up to 58(3)

51(Q) 59 68.2 57.5 BF

TaL/TL (3): 0.158-0.172

(2): 0.132-0.149 0.173 0.178 0.175 0.126

VEN (3): 140-147

(2): 147-155 148 151 144 158

SC (3): 36-73

(Q): 31-38 42 40 40 33

SL 7-8 8/8 8/8 8/8 8/8

IL 8-9 9/9 9/9 9/9 9/9

one juvenile frog (Fejervarya limnocharis) and two

unidentified anuran tadpoles.

Oligodon ocellatus (Morice, 1875)

Specimens examined: Two female specimens (ZFMK

88919-88920), collected by P. Geissler in Cat Tien Na-

tional Park, Dong Nai Province. One female specimen

(IEBR A.2010.54.), collected by W. Van Devender on 28

May 2004, near Bau Sau Lake, Dong Nai Province (near

12773219 CN 109520743. 7 (E67 murassis):

Characteristic features: The morphological traits of the

specimens (data given in the following order: ZFMK

88919 / ZFMK 88920 / IEBR A.2010.54) from Cat Tien

National Park fit the descriptions of David et al. (2008a):

SVL 405 mm / 199 mm / 209 mm; TaL 108 mm / 28.3

mm / 29.7 mm; | presubocular; 8 SL; 9 IL; 19 MD; 173

/ 159/165 VEN; precloacal scale undivided; 59 / 59/41

SC; dorsum light brown, with 11—14 dark brown blotch-

es, edged with black; dark brown cross bands along the

dorsum; head dark brown. For colouration in life see Fig.

Se:

Remarks: The occurrence in the Cat Tien National Park,

Dong Nai Province is the southernmost record of this

species in Vietnam (compare with Nguyen et al. 2009).

Xenochrophis flavipunctatus (Hallowell, 1861)

Specimens examined: One juvenile specimen (IEBR

A.2010.46), collected by W. Van Devender in 2006, in Cat

Tien National Park, Dong Nai Province. One subadult

specimen (ZFMK 88914), collected by P. Geissler on 19

Bonn zoological Bulletin 60 (1): 9-16

May 2009 at the headquarters of the Cat Tien National

Park, Dong Nai Province (near 11°25°19.3”N,

107°25°42.0”E, 104 m a.s.1.).

Characteristic features: The morphological characters of

the specimens (data given in the following order IEBR

A.2010.46./ ZFMK 88914) fit the descriptions of Bour-

ret (1936), Smith (1943), and Ziegler et al. (2007): SVL

206 mm / 244 mm; TaL 76 mm/ 119 mm; | loreal; | pre-

ocular; 3 postoculars; 8 SL; 10 IL; 19 MD, keeled; 136 /

125 VEN; precloacal scale divided; 77 / 85 divided SC;

head and dorsum grey; light band from the eye to the an-

gle of the mouth, edged with black; parietals with a light

blotch, edged with dark brown (in juveniles); lateral sides

of neck yellow; flanks with a series of black blotches. For

colouration in life see Fig. 3f.

DISCUSSION

In their review of the herpetofauna of the Cat Tien Na-

tional Park, Nguyen & Ho (2002) listed 42 species of

snakes, which is almost 21% of the 203 snake species

known from Vietnam (Nguyen et al. 2009, Orlov et al.

2010, Ziegler & Nguyen 2010). These records were based

on their unpublished reports and examination of voucher

specimens. After reviewing the list, we regard one record

as questionable. The record of “Homalopsis fasciatus ”,

which is based on an unpublished report, is obviously in-

correct as this taxon does not exist in current faunal works

in this region (Bourret 1936, Nguyen et al. 2009). Besides

nine new records based on voucher specimens, the pres-

ence of Boiga guangxiensis in the Cat Tien National Park

was proven by a photograph taken by W. Van Devender

(pers. comm.). Nguyen & Ho (2002) reported the occur-

©ZFMK

New records of snakes from southern Vietnam

Table 2. List of snake species recorded from Cat Tien National Park (Dong Nai and Lam Dong provinces).

Taxon Nguyen Le et al. Nguyen & Ho This study

(1988) (1998) (2002)

Typhlopidae

Ramphotyphlops braminus (Daudin, 1803) x Xx IEBR A.2010.51—A.2010.52

ZFMK 88925

Typhlops siamensis Gunther, 1864 ZFMK 88922

Cylindrophiidae

Cylindrophis ruffus (Laurenti, 1768) x x —

Pythonidae

Python bivittatus Kuhl, 1820 X x x —

Python reticulatus (Schneider, 1801) xX x XxX —=

Xenopeltidae

Xenopeltis unicolor Reinwardt in Boie, 1827 Xx —

Colubridae

Ahaetulla prasina (Reinhardt, 1827) xX x IEBR A.2010.53

Boiga cyanea (Duméeril, Bibron & Dumeril, 1854) x —

Boiga guangxiensis Wen, 1998

Boiga multomaculata (Boie, 1827) ZEMK 88923

Boiga siamensis Nootpand, 1971 x =

Calamaria pavimentata Dumeéril, ZFMK 88924

Bibron & Dumeril, 1854

Chrysopelea ornata (Shaw, 1802) x Ke ZFMK 88905-88907

Coelognathus flavolineatus (Schlegel, 1837) ZFMK 88898

Coelognathus radiatus (Boie, 1827) x x x —

Dendrelaphis ngansonensis (Bourret, 1935) ZFMK 88913

Dendrelaphis pictus (Gmelin, 1789) x Xx ZFMK 88912

Dryocalamus davisonii (Blanford, 1878) x ZFMK 88929-88930, 91898

IEBR A.2010.46

Gonyosoma oxycephalum (Boie, 1827) x —

Lycodon laoensis Ginther, 1864 x ZFMK 88928

Lycodon subcinctus Boie, 1827 IEBR A.2010.42—A.2010.43

ZFMK 91899

Oligodon cinereus (Giinther, 1864) x ZEMK 88921

Oligodon deuvei David, Vogel & van Rooijen, 2008 IEBR A.2010.16—A.2010.17

ZFMK 91226

ZMMU NAP-02811

Oligodon fasciolatus (Gunther, 1864) xe —

Oligodon ocellatus (Morice, 1875) IEBR A.2010.54

ZFMK 88919-88920

Ptyas korros (Schlegel, 1837) x x x ZEMK 88915

Ptyas mucosa (Linnaeus, 1758) x x x —

Rhabdophis chrysargos (Schlegel, 1837) Xx x TEBR A.2010.45

Rhabdophis subminiatus (Schlegel, 1837) x ZFMK 88908-88909

Sibynophis collaris (Gray, 1853) x —

Xenochrophis flavipunctatus (Hallowell, 1861) IEBR A.2010.46, ZFMK

88914

Homalopsidae

Enhydris bocourti (Jan, 1865) x x IEBR A. 2010.55—A.2010.56

ZFMK 88926

Enhydris enhydris (Schneider, 1799) x x —

Homalopsis buccata (Linnaeus, 1758) xX x ZFMK 88927

Pareatidae

Pareas carinatus (Boie, 1828) x ZFMK 88910-88911

Pareas margaritophorus (Jan, 1866) x TEBR A.2010.44

Lamprophiidae (incertae sedis)

Psammodynastes pulverulentus (Boie, 1827) x ZFMK 88900-88904

Elapidae

Bungarus candidus (Linnaeus, 1758) xe —

Bungarus fasciatus (Schneider, 1801) x xX —_—

Calliophis maculiceps (Gunther, 1858) x IEBR A.2010.48,

ZFMK 91900

Naja kaouthia Lesson, 1831 Ke —

Naja siamensis Laurenti, 1768 x x x —

Ophiophagus hannah (Cantor, 1836) x x x —

Viperidae

Calloselasma rhodostoma (Kuhl, 1824) x —

Cryptelytrops albolabris (Gray, 1842) Xx x Xx IEBR A.2010.49

ZFMK 91897

Cryptelytrops rubeus Malhotra, Thorpe, Mrinalini & Stuart, 2011 Ke IEBR A.2010.50

(listed as C. macrops by Nguyen & Ho 2002) ZFMK 88916-88918

Viridovipera stejnegeri (Schmidt, 1925) Xx —

Bonn zoological Bulletin 60 (1): 9-16

©ZFMK

16 Peter Geissler et al.

rence of Cryptelytrops macrops in the Cat Tien National

Park. However, Malhotra et al. (2011) recently described

the Ruby-eyed Green Pitviper Cryptelytrops rubeus from

southern Vietnam and Cambodia.The distribution of

Cryptelytrops macrops is now restricted to Thailand, Cam-

bodia, and Laos (Malhotra et al. 2011). Based on our spec-

imens with the red eye, we herein confirm the presence

of Cryptelytrops rubeus in the Cat Tien National Park. The

updated checklist of 51 species of snakes of Cat Tien Na-

tional Park is provided in Table 2. For the generic com-

position of snakes we follow Pyron et al. (2010).

Acknowledgements. We cordially thank Tran Van Thanh (Cat

Tien National Park) for his valuable support during the field stud-

ies and for issuing respective permits. We gratefully acknowl-

edge Le Xuan Canh and Ta Huy Thinh (IEBR) for their support

and the loan of specimens. We are grateful to Wayne Van De-

vender and Paul Moler for providing scientific material and pho-

tographs. We thank Gernot Vogel and Patrick David for their tax-

onomic advice. PG thanks Thomas Ziegler, Phung My Trung,

Igor Palko, Anton N. Chekanov, Vladimir A. Zryanin, Alexan-

der Krohn, Le Duc Khanh, Jendrian Riedel, Bernd Geissler, and

Rike Bach for their support during numerous field trips in Cat

Tien. We thank Eleanor Sterling and Kevin Koy for providing

the map and E. Galoyan for providing photographs. We highly

appreciate the comments of Andreas Schmitz and Thomas

Ziegler on an earlier version of our manuscript. The field work

of PG in Vietnam was partly funded by the Alexander Koenig

Gesellschaft and the German Herpetological Society (DGHT).

REFERENCES

Bourret R (1935) Notes herpétologiques sur |’Indochine fran-

caise. XI. Sur quelques serpents recoltés en 1934. Bulletin

général de l’instruction publique. 1934/35 (9. Mai): 289-296

Bourret R (1936) Les serpents de I’Indochine, II, Catalogue sys-

tématique déscriptif. H. Basuyau, Toulouse

Cambden-Main S (1970) A fieldguide to the Snakes of South

Vietnam. Division of Reptiles and Amphibians United States

National Museum Smithsonian Institut, City of Washington

Daltry JC, Wiister W (2002) A new species of Wolf Snake (Ser-

pentes: Colubridae: Zycodon) from the Cardamom Mountains,

Southwestern Cambodia. Herpetologica 58: 498-504

David P, Vogel G, Pauwels OSG (2008a) A new species of the

genus Oligodon Fitzinger, 1826 (Squamata: Colubridae) from

Southern Vietnam and Cambodia. Zootaxa (1939): 19-37

David P, Vogel G, van Rooijen J (2008b) A revision of the

Oligodon taeniatus (Gunther, 1861) group (Squamata: Colu-

bridae), with the description of three new species from the In-

dochinese Region. Zootaxa 1965: 1-49

Deuve J (1970) Serpents du Laos. Mémoires de |’ office de la

recherche scientifique et technique outre-mer, Paris 39: 1-251

Deuve J (1985) Serpents du Laos. Dessins en couleurs et doc-

uments manuscrits. Unpublished manuscript notes, deposit-

ed in the library of the ,, Taxonomie-collection- Reptiles & Am-

phibiens* unit, Muséum National d’Histoire Naturelle, Paris

Geissler P, Nazarov R, Orlov NL, Bohme W, Phung TM, Nguyen

QT, Ziegler T (2009) A new species of the Cyrtodactylus ir-

Bonn zoological Bulletin 60 (1): 9-16

regularis complex (Squamata: Gekkonidae) from southern

Vietnam. Zootaxa 2161: 20-32

Giinther ACLG (1864) The reptiles of British India. Ray Soci-

ety, London

Le XC, Hoang MK, Le DT, Ho TC, Hoang VT, Nguyen VT,

Nguyen TV, Tran VD (1998) Results of zoological survey in

Cat Loc Nature Reserve, Cat Tien District, Lam Dong

Province. Unpublished report of IEBR, Hanoi

Malhotra A, Thorpe RS, Mrinalini, Stuart BL (2011) Two new

species of pitviper of the Genus Cryptelytrops Cope 1860

(Squamata: Viperidae: Crotalinae) from Southeast Asia.

Zootaxa 2757: 1—23

Nguyen QT (1988) Preliminary list of Reptiles and Amphibians

in Nam Cat Tien forest reserve. Garrulax 5: 8-9

Nguyen VS, Ho TC (1996) A checklist of reptiles and amphib-

ians of Vietnam. Science and Technology Publishing House,

Hanoi (in Vietnamese)

Nguyen VS, Ho TC (2002) Species composition of the herpeto-

fauna recorded in the Cat Tien National Park. Journal of Bi-

ology 24: 2-10 (in Vietnamese)

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

nam. Edition Chimaira, Frankfurt am Main

Orlov NL, Nguyen QT, Nguyen TT, Ananjeva NB, Ho TC (2010)

A new species of the genus Calamaria (Squamata: Ophidia:

Colubridae) from Thua Thien- Hue Province, Vietnam. Russ-

ian Journal of Herpetology 17: 236-242

Polet G, Ling S (2004) Protecting mammal diversity: opportu-

nities and contraints for pragmatic conservation management

in Cat Tien National Park, Vietnam. Oryx 38: 1-11

Pyron RA, Burbrink FT, Colli GR, Montes de Oca AN, Vitt LJ,

Kuczynski CA, Wiens JJ (2010) The phylogeny of advanced

snakes (Colubroidea), with discovery of a new subfamily and

comparison of support methods for likelihood trees. Molec-

ular Phylogenetics and Evolution: doi:10.1016/j.ym-

pev.2010.11.006

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

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

tiles and Amphibians, Vol. 3. Serpentes. Taylor and Francis,

London

Tirant G (1885) Notes sur les reptiles et les batraciens de la

Cochinchine et du Cambodge. Imprimérie du Gouvernement,

Saigon

Wallach V (2001) Typhlops roxanae, a new species of Thai blind

snake from the 7: diardii species group, with a synopsis of the

Typhlopidae of Thailand (Serpentes: Scolecophidia). Raffles

Bulletin of Zoology 49: 39-49

Ziegler T, Hendrix R, Vu NT, Vogt M, Forster B, Dang NK

(2007) The diversity of a snake community in a karst forest

ecosystem in the central Truong Son, Vietnam, with an iden-

tification key. Zootaxa 1493: 1-40

Ziegler T, Nguyen QT (2010) New discoveries of amphibians

and reptiles from Vietnam. Bonn zoological Bulletin 57:

137-147

Ziegler T, Vogel G (1999) On the knowledge and specific sta-

tus of Dendrelaphis ngansonensis (Bourret, 1935) (Reptilia:

Serpentes: Colubridae). Russian Journal of Herpetology 6:

199-208

Received: 28.10.2010

Accepted: 09.03.2011

Corresponding editor: F. Herder

©ZFMK

Volume 60

Bonn zoological Bulletin

Issue | pp. 17-24 | Bonn, May 2011

Description of a new species of the genus Dendrelaphis Boulenger, 1890

from Myanmar (Squamata: Serpentes: Colubridae)

Gernot Vogel!*& Johan van Rooijen?

| Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany;

E-mail: Gernot. Vogel@t-online.de

2 Netherlands Centre for Biodiversity Naturalis, section Zoological Museum Amsterdam, Mauritskade 61,

1092 AD Amsterdam, The Netherlands

3 Tulpentuin 313, 2272 EH, Voorburg, The Netherlands; E-mail: j1.van.rooijen@hetnet.nl

«corresponding author.

Abstract. The population systematics of the colubrid snake so far referred to as Dendrelaphis gorei (Wall, 1910) were

investigated by carrying out a multivariate analysis of geographic variation. The results reveal the existence of two dis-

tinct and apparently disjunct phenotypes. These phenotypes are assumed to represent independent evolutionary lineag-

es. The first lineage corresponds with D. gorei which inhabits the Assam Valley and adjoining Himalayan foothills. How-

ever, the name Dendrophis gorei Wall, 1910 is actually a junior synonym of Dendrelaphis biloreatus Wall, 1908. Con-

sequently, the latter name is adopted for this lineage. The second lineage, described in this paper as a new species, in-

habits the South Myanmar lowlands and neighbouring mountain chains. It is distinguished from D. biloreatus on the ba-

sis of its higher ventral count (203—212 versus 190-199), its undivided anal shield (divided in D. biloreatus) and its high-

er average number of anterior temporal shields (usually 2 versus usually 1).

Key words. Dendrelaphis sp. n., Dendrelaphis gorei syn. n., Dendrelaphis biloreatus, Myanmar, analysis of geograph-

ic variation.

INTRODUCTION

The colubrid snakes of the genus Dendrelaphis Boulenger,

1890 are widely distributed, ranging from Pakistan in the

West to the northern and eastern coast of Australia in the

East and South and to southern China in the North (Ziegler

& Vogel 1999). Members of the genus Dendrelaphis are

slender, diurnal species that are predominantly arboreal

and feed mainly on lizards and amphibians.

Boulenger (1894), Wall (1921), Meise & Henning (1932),

Mertens (1934) and Smith (1943) have in turn worked on

the systematics of this genus. Nevertheless, their cumu-

lative effort did not result in an unambiguous and com-

plete taxonomy of this genus, a fact that was underlined

by the recent descriptions and revalidations of several

species (Vogel & Van Rooijen 2007, 2008, 2011; Van

Rooyen & Vogel 2008a, 2008b, 2008c, 2009).

Dendrelaphis gorei (Wall, 1910) has been reported to oc-

cur in Northeast India and Myanmar. It is one of the least

known members of its genus and is scarcely represented

in museum collections. In recent years however, new ma-

terial has become available through the Myanmar Herpeto-

logical Survey, a collaborative effort of the California

Academy of Sciences, Smithsonian Institution and the

Forest Department, Ministry of Forestry, Myanmar (e.g.

Wogan et al. 2008). In this paper, we investigate the pop-

ulation systematics of D. gorei by performing a multivari-

ate analysis of geographic variation based on morpholog-

ical data. Furthermore, the name Dendrelaphis biloreatus

Wall, 1908 is evaluated. Smith (1943) synonymized D.

biloreatus with D. gorei (Wall 1910) as he could not find

any differences between the two species apart from the di-

vided loreal shield in D. biloreatus. However, according

to the rule of priority (Art. 23, ICZN, 1999), the name D.

biloreatus should be adopted for this species.

MATERIALS AND METHODS

Eleven museum specimens were examined for this study.

In addition, data on three additional specimens were tak-

en from Wall (1908, 1910). For each examined specimen,

23 characters including aspects of colour pattern, body

proportions and scalation were recorded (Table 1). Eye-

diameter and distance eye-nostril were measured with a

slide calliper to the nearest 0.1 mm. These measurements

were made on the left and right side and were subsequent-

ly averaged. Snout-vent length was measured to the pos-

18 Gernot Vogel & Johan van Rooijen

terior margin of the anal plate by marking the length on

a piece of string and subsequently measuring the position

of the mark to the nearest 0.5 cm. Tail-length was meas-

ured to the nearest 0.5 cm by straightening the tail against

a ruler. The number of ventrals was counted using DowI-

ing’s method (1951). Subcaudals were counted on one

side, the terminal scute was excluded. The first sublabial

was defined as the scale that starts between the posterior

chin shield and the infralabials and that borders the infral-

abials (see Peters 1964, fig. 7). The last infralabial was

defined as the infralabial still covered completely by the

last supralabial. The posterior most temporal scales were

defined as the scales of which more than half of the area

lies in front of an imaginary line that runs from the apex

of the last supralabial to the posterolateral corner of the

parietal.

Geographic coordinates were taken directly from the field

notes or were obtained by translating locations to coordi-

nates. Relevant variables were included in a Principal

Components Analysis (PCA, e.g. Cramer 2003) in order

to reduce the dimensions of the dataset. The resulting

PCA-scores were then plotted against longitude and lati-

tude in order to visualize the pattern of geographic vari-

ation. Confirmatory geographic analyses of the PCA-score

were carried out by Analysis of Covariance (ANCOVA;

Maxwell & Delaney 1990; Norusis & SPSS 1993) using

longitude and latitude as covariates and phenotype as fac-

tor. Normality of the PC-scores was first tested by means

of Kolmogorov-Smirnovy tests.

All statistical analyses were carried out with the software

SPSS (2006; SPSS for Windows. Release 14.0.2. Chica-

go: SPSS Inc.).

Table 1. List of morphometric, scalation and coloration characters used in this study and their abbreviations.

Abbreviation Character

Morphometrics

EYED Horizontal diameter of the eye

EYEN Distance from centre of the eye to posterior border of the nostril

TAIL Tail-length

WSNOUT Width of the snout measured at the position of the nostrils

LHEAD Head-length measured from the tip of the snout to the rear of the jaw

SVL Snout-vent length

Scalation

VENT Number of ventrals

SUBC Number of subcaudals

DORI Number of dorsal scale rows | head-length behind the head

DOR2 Number of dorsal scale rows at the position of the middle ventral

DOR3 Number of dorsal scale rows | head-length before the tail

SUBL Number of infralabials touched by the first sublabial (L+R)

SLI Number of supralabials (L+R)

SL2 Number of supralabials touching the eyes (L+R)

LOR Number of loreals (L+R)

INFR Number of infralabials (L+R)

ATEMP Number of anterior temporals (L+R)

TEMP Number of temporals (L+R)

POC Number of postoculars (L+R)

VERT Vertebral scales smaller than (0) or larger than (1) scales of the first dorsal row

AN Anal shield divided (0) or undivided (1)

Coloration

TSTRIPE Postocular stripe absent (0), rudimentary (1), present (2)

LSTRIPE Ventrolateral stripe absent (0), present (1)

The collected data were used to carry out an analysis of

geographic variation. The objective of this analysis was

to enable differentiation between clinal variation and phe-

netic discontinuities, the latter being evidence for lineage

separation (e.g. Lenk & Wister 1999; De Queiroz 2007).

Bonn zoological Bulletin 60 (1): 17-24

Material examined. S-037, Guwahati, Assam; BMNH

1940.3.4.24, Assam; BMNH 1940.3.4.25, Samagooting,

Assam; BMNH 1946.1.10.30, Jaipur, Assam (syntype

Dendrophis gorei Wall, 1910); CAS 208429, Myanmar,

Bago Division, Bago Yoma, Sein Yay Camp, 18 51 21.636

©OZFMK

A new Dendrelaphis from Myanmar 19

N, 96 10 21.324 E; CAS 211939, Myanmar, Ayeyarwady

Division, vic Mwe Hauk Village (16 16 39.2 N, 94 45 32.5

E); CAS 222114-222115, Myanmar: Bago Division, Aok

Twin Township, Ka Baung Reserve, Sein Ye Camp, Block

120, 18 51 16.1 N, 96 10 23.6 E; CAS 222339, Myan-

mar: Chin State, Min Dat Township, Min Dat Township,

Nat Ma Taung National Park, 21 21 14.9 N, 93 56 08.3

E; CAS 234880, Myanmar,: Chin State, Mindat Township,

Mindat District, 21 26 43.5 N, 94 00 24.0 E; CAS 244037,

Myanmar, Sagaing Division, Leha and Khante township

boundary, Nana Sa Laing camp, 26 07 23.1 N, 95 32 24.6

Museum abbreviations. BMNH: Natural History Muse-

um, London, Great Britain. CAS: Collection of the Cali-

fornia Academy of Sciences, USA. S: Saibal Senguptas

Collection, Guwahati, Assam

RESULTS

Statistics

Visual inspection of the data suggested a striking differ-

entiation between specimens from Assam (NE India) and

Northwest Myanmar on the one hand and specimens from

the remainder of Myanmar on the other hand. One phe-

notype was characterized by a high number of ventral

counts, an undivided anal shield and usually two anteri-

or temporal shields whereas the other was characterized

by a low number of ventrals, a divided anal shield and usu-

ally one anterior temporal shield. These characters were

included in a PCA. The first component explained 92%

of the total variance. This demonstrates that these char-

acters covary strongly and thus form a suite of characters

that sharply differentiates between the two phenotypes.

The individual scores on this component were plotted

: 1

16 '

0.5

O oC

fi.

0.5

Ae yj 2 2

specie ll | |

es eae

T aay

5 ee 2 ibe eo aes 20

lon; earl 2

Stud % 98 latitude

Fig. 1.

PCA-scores based on a PCA of the characters VENT, AN and ATEMP against longitude and latitude. 1 Phenotype 1, 2

phenotype 2, 2b= type D. biloreatus Wall, 1908; 2a = syntype D. gorei Wall, 1910.

Bonn zoological Bulletin 60 (1): 17-24

©OZFMK

against latitude and longitude to visualize the pattern of

geographic variation. In figure 1, data-points correspon-

ding with the type of Dendrelaphis biloreatus and one of

the two syntypes of Dendrophis gorei are indicated for a

later discussion of the appropriate nomenclature. A phe-

netic discontinuity is in evidence, the difference in PCA-

scores between the phenotypes being highly significant

(ANCOVA, df=1,11; P<0.0001). Subsequent inclusion in

the model of longitude and latitude in the form of covari-

ates demonstrated that geographic coordinates had no

added explanatory power (ANCOVA, df=1,9; P=0.9 and

df=1,9; P=0.4 respectively). Thus, there is no evidence of

clinal variation, the transition from one phenotype to the

other apparently being sudden rather than gradual.

The sudden phenetic transition represents strong evidence

for lineage separation. Thus, the two distinguished phe-

notypes are here interpreted as representing distinct evo-

lutionary sister-lineages. An evaluation of the characters

separating the two phenotypes supports this view. First,

one phenotype exhibits an undivided anal shield where-

as the other has a divided anal shield. An undivided anal

shield is an exceptional character in the genus Den-

drelaphis. The only congeneric species that exhibits this

character (in roughly 40% of the specimens) is Den-

Gernot Vogel & Johan van Rooien

drelaphis proarchos (Wall, 1909) (Vogel & van Rooijen

2011). We examined more than 600 museum specimens

of this genus, representing 24 species and subspecies. On-

ly a single specimen (beside those identified as D. proar-

chos and the material presented in this study) was found

to have an undivided anal shield. Second, the two pheno-

types differ substantially in the number of ventral scales,

the difference being 19 on average (211 in phenotype 1,

192 in phenotype 2). This is a relatively large difference

when compared to established interspecific differences.

For instance, the difference in ventral count between D.

formosus on the one hand and D. kopsteini and D.

cyanochloris on the other hand is 5 and 19 respectively

(Vogel & Van Rooyen 2007). The difference between D.

haasi and D. pictus, D. pictus andamanensis, D. humayu-

ni, D. tristis 1s 3, 20, 10 and 14 respectively (Van Rooi-

jen & Vogel, 2008a). Finally, the difference between D.

chairecacos on the one hand and D. tristis and D. schokari

on the other is 10 and 12 respectively (Van Rooijen & Vo-

gel 2009). The third difference between the two pheno-

types is the number of anterior temporal scales. Pheno-

type | usually has two anterior temporal scales whereas

phenotype 2 has a single anterior temporal in the major-

ity of cases. This character alone already leads to a cor-

rect identification in 85% of the cases.

Table 2. Comparison between the type of Dendrelaphis biloreatus and six specimens of the species so far referred to as Den-

drelaphis gorei.

type Dendrelaphis biloreatus

Dendrelaphis gorei (n=6)

Ventrals

Subcaudals

Relative tail-length

Supralabials

Supralabials touching the eye

Anterior temporals

Total number of temporals

Poctoculars

Loreals

Dorsal rows at midbody

Divided anal shield

Faint ventrolateral line, not bordered by black lines

Postocular stripe that covers the whole

temporal region and extends onto the neck

Total length

vertebrals strongly enlarged

Ground color brown

Maxillary teeth

192 190-199

147 144-152

0.34 0.34—0.35

9 8-9

3 2-3

1 1-2

8 7-12

2 1-2

2 1

1 13

yes yes

70.0 53.5—90.0

yes yes

21 20 (n=1)

Bonn zoological Bulletin 60 (1): 17-24

©ZFMK

A new Dendrelaphis from Myanmar 21

Fig. 2.

Dendrelaphis walli sp. n., holotype (CAS 234880).

Nomenclature

Two available names, Dendrophis gorei Wall, 1910 and

Dendrelaphis biloreatus Wall, 1908 are relevant in the

context of this study. In figure 1, data pertaining to a syn-

type of D. gorei and the type of D. biloreatus are indicat-

ed. It is evident that neither of these types represents phe-

notype 1 (South Myanmar and neighbouring mountain

chains). Thus, we here describe phenotype | as a new

species. Phenotype 2 (Assam and Northwest Myanmar)

is currently known as D. gorei (Wall, 1910), but we here

adopt the older name Dendrelaphis biloreatus Wall, 1908

to represent this lineage. Dendrelaphis biloreatus was de-

scribed by Wall on the basis of a single specimen. Smith

(1943) synonymized D. biloreatus with D. gorei as he

could not find any difference between the two species

apart from the divided loreal shield in D. biloreatus. In

table 2, a more detailed comparison is provided between

Bonn zoological Bulletin 60 (1): 17-24

the morphological characteristics of the type of D. bilore-

atus and six specimens of D. gorei. With the exception of

the loreal shield, the values pertaining to the type of D.

biloreatus all fall within the range of D. gorei. Thus, the

results agree with Smith’s view that the type of D. bilore-

atus represents an exceptional specimen with a split lo-

real shield within D. gorei. In the context of a revision of

the taxonomy of this genus, the authors have come across

several specimens of Dendrelaphis with either a double

or an absent loreal shield. Furthermore D. biloreatus has

a rather long and slender postnasal scale which looks like

second loreal shield at the first glance. So indirectly the

name makes some sense. Consequently, the two names in-

deed refer to the same species. According to the rule of

priority (ICZN, 1999, art. 23.1), the name Dendrelaphis

biloreatus should actually be adopted for this species.

OZFMK

ie)

Gernot Vogel & Johan van Rooyen

Table 3. Morphological and coloration characters of the types of Dendrelaphis walli sp. n.

Collection N° CAS 234880 CAS 208429 CAS 211939 CAS 222114 CAS 222339 CAS 222115

status holotype paratype paratype paratype paratype paratype

Sex f - m m m f

Snout-vent length (cm) 48.5 ~ 49.0 54.5 61.5 595

Tail-length (cm) 22.5 ~ 25.5 29.5 29.0 31.0

Head-length (mm) 13.5 - 15.5 15.5 17.0 17.0

Eye-diameter (mm) 3.1 — 3.5 35) 3.9 3.5

Snout-width 2.8 ~ 3.2 3.4 i) 3.3

Ventrals 213 211 203 210 212 212

Subcaudals 145 - 148 163 147 159

Dorsal formula 13-13-11 ~ 13-13-11 13-13-11 13-13-9 13-13-11

Temporal formula 2222/2122) 9 222/222 222/212 212/112 2212/222 222/212

Supralabials 8/8 8/8 8/8 8/8 8/8 8/8

Supralabials touching the eye 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5 4,5/4,5

Infralabials 10/10 - 10/10 2/9 10/10 11/10

Infralabials touched 6,7/6,7 _ 6,7/6,7 6,7/5,6 6,7/6,7 6,7/6,7

by first sublabial

Loreals 1/1 1/1 1/1 1/1 1/1 1/1

Postoculars 2/2 3/3 2/2 2/2 2/2 2/2

Number of scales bordering 4 = 5 5 6 5

the parietal scales

Vertebrals larger than yes ~ yes yes yes yes

dorsals of the first row

Anal shield entire yes yes yes yes yes yes

Light ventrolateral stripe yes = yes yes yes yes

which is faint and is not

bordered by black lines

Postocular stripe broad, yes — yes yes yes yes

covering the whole temporal

region and extending

onto the neck

Taxonomy

Dendrelaphis walli sp. n. (Fig. 2)

Dendrelaphis gorei (Wall, 1921) (part.)

Holotype. CAS 234880, adult female from Myanmar

(Chin State: Mindat township, Mindat District, 21 26 43.5

N, 94 00 24.0 E, 3582 ft), collected by A.K. Shein and T.

Nyo, 23 Aug 2005.

Paratypes. CAS 208429,

222114222115, CAS 222339.

CAS. 211939, CAS

Bonn zoological Bulletin 60 (1): 17-24

Diagnosis. A species of Dendrelaphis, characterized by

the combination of: 1) 13 dorsal scale rows at midbody;

2) strongly enlarged vertebral scales; 3) 203—213 ventrals;

4) 145-163 subcaudals; 5) 8 supralabials; 6) supralabials

4 and 5 bordering the eye; 7) 2 anterior temporals in the

majority of specimens; 8) a short sublabial that touches 2

infralabials; 9) an undivided anal shield; 10) a black pos-

tocular stripe that covers the majority of the temporal re-

gion and extends onto the neck; 11) black, oblique bars

on the neck region; 12) a pale ventrolateral line, not bor-

dered by black lines.

©ZFMK

A new Dendrelaphis from Myanmar 23

Table 4. Diagnostic differences between Dendrelaphis walli sp. n. and Dendrelaphis biloreatus Wall, 1908.

Dendrelaphis walli n. sp. (n=7)

Dendrelaphis biloreatus (n=7)

Ventrals 211 (203-213)

Anterior temporals

anal shield entire

usually 2 (92% of specimens)

192 (190-199)

usually 1 (86% of specimens)

divided

Description of the holotype. Adult female; body very

slender; snout-vent length 48.5 cm; tail-length 22.5

cm;head distinct from neck; head-length 13.5 mm; snout-

width 2.8 mm; pupil round; eye-diameter 3.1 mm; distance

eye-nostril 3.1 mm; 213 ventrals; 145 subcaudals; dorsal

scales in 13-13-11 rows; 8 supralabials, 4'* and 5‘ bor-

der the eye; 10 infralabials, infralabials 1—5 touch the first

chinshield, infralabials 5 and 6 touch the second chin-

shield; | preocular; 2 postoculars; | loreal; temporal for-

mula 2:2:2:1:2 (L), 2:1:2:2 (R); first sublabial touches in-

fralabials 6 and 7; vertebrals strongly enlarged, with

straight posterior margin, width of the vertebral scale at

the position of the middle ventral scale 2.4 mm; anal en-

tire; parietal scales bordered posteriorly by 4 scales;

ground color brownish; supralabials and throat white;

some black spots on supralabials 2-4, the loreal and the

preocular; a black postocular stripe starts behind the eye,

covers the majority of the temporal region, and extends

onto the neck where it breaks up into narrow, black oblique

bars which fade away further posteriorly; a faint ventro-

lateral line is present, covering the first dorsal row and

lower half of the second dorsal row; the ventrolateral line

is not bordered by black lines; belly whitish.

Bassein RANGOON

». Maui

ee iY THAILAND

| Lf

De a |

Fig. 3. Currently known distribution of D. biloreatus

(white b) and D. walli sp. n. (black W).

Bonn zoological Bulletin 60 (1): 17-24

Variation. Table 3 provides data regarding the types of

D. walli sp. n.

Comparison with congeneric species. Dendrelaphis wal-

li sp. n. differs from all congeners, except Dendrelaphis

biloreatus, Dendrelaphis caudolineatus and Dendrelaphis

caudolineolatus, by its dorsal formula of 13-13-11/9. It dif-

fers from Dendrelaphis caudolineatus in its strongly en-

larged vertebral scales (not enlarged in D. caudolineatus),

its much more slender body (stout in D. caudolineatus),

its undivided anal shield, the number of supralabials (9 vs.

8 in D. walli sp. n.) and in its coloration. It differs from

Dendrelaphis caudolineolatus in the number of ventral

scales (149-175 vs. 203-213 in D. walli sp. n.) and sub-

caudal scales (111-129 vs. 145-163 in D. walli sp. n.) and

its undivided anal shield. Differences between D. walli sp.

n. and D. biloreatus are given in Table 4.

Sexual dimorphism. Females have a wider snout than

males (ANCOVA, df=1,5, P=0.004). In addition, there is

some evidence that females have a larger eye than males

(ANCOVA, df=1,5, P=0.08). No evidence of other sexu-

al dimorphisms was found though this may be due to lack

of statistical power (e.g. Streiner 1990).

Distribution. According to currently known locality

records, D. biloreatus inhabits the Assam Valley and ad-

joining Himalayan foothills whereas D. wailli sp. n. inhab-

its the South Myanmar lowlands and neighbouring

mountain chains. Thus, the two species appear to exhib-

it a disjunct distribution (figure 3). However, this may be

due to collecting gaps. In reality, one or both of these

species may inhabit the intermediate area. On the other

hand, the Myanmar Herpetological Survey did collect in

the central and northern parts of Myanmar, which did not

yield a specimen of either species, with the exception of

one D. biloreatus (CAS 244037) near the border with As-

sam, within the known range of this species. If these

species indeed occur allopatrically, the central dry zone

of Myanmar and adjoining mountain ranges may be the

geographic barrier that separates these species and that

may have enabled the independent evolution of the two.

Etymology. This species is dedicated to Major Frank Wall

(1868-1950), in recognition of his outstanding work on

the genus Dendrelaphis.

©ZFMK

24 Gernot Vogel & Johan van Rooijen

Acknowledgements. We thank J. V. Vindum (CAS, San Fran-

cisco, USA), C. J. McCarthy (BMNH, London, United King-

dom), R. Vonk and H. Praagman (ZMA, Amsterdam, The

Netherlands), Saibal Sengupta and Jayaditya Purkayastha (In-

dian collections) for letting us examine preserved specimens.

REFERENCES

Boulenger GA (1894) Catalogue of the Snakes in the British Mu-

seum (Natural History). Volume II., Containing the Conclu-

sion of the Colubridae Aglyphae. London: Taylor & Francis

Cramer D (2003) Advanced quantitative data analysis. Philadel-

phia: Open University Press

De Queiroz K (2007) Species concepts and species delimitation.

Systematic Biology 56: 879-886

Dowling HG (1951) A proposed standard system of counting

ventrals in snakes. British Journal of Herpetology 1: 97-99

ICZN — International Commission on Zoological Nomenclature

(1999) International Code of Zoological Nomenclature,

Fourth Edition adopted by the International Union of Biolog-

ical Sciences. The International Trust for Zoological Nomen-

clature, London

Lenk P, Witister W (1999) A multivariate approach to the sys-

tematics of Italian Ratsnakes of the Elaphe longissima com-

plex (Reptilia, Colubridae): revalidation of Camerano’s Cal-

lopeltis longissimus var. lineata. Herpetological Journal 9:

153-162

Maxwell SE, Delaney HD (1990) Designing experiments and

analyzing data. California: Wadsworth Inc.

Meise W, Henning W (1932) Die Schlangengattung Dendrophis.

Zoologischer Anzeiger 99: 273-297

Mertens R (1934) Die Schlangengattung Dendrelaphis

Boulenger in systematischer und zoogeographischer

Beziehung. Archiv ftir Naturgeschichte, Berlin (N. F.) 3(2):

187-204

Norusis & SPSS Inc. (1993) SPSS Advanced Statistics, SPSS,

Chicago

Peters JA (1964) Dictionary of herpetology: a brief and mean-

ingful definition of words and terms used in herpetology. New

York: Hafner Publ. Co.

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

ma. Reptilia and Amphibia. Vol. 3 Serpentes. London: Tay-

lor & Francis

Streiner DL (1990) Sample size and power in psychiatric re-

search. Canadian Journal of Psychiatry 35: 616-620

Bonn zoological Bulletin 60 (1): 17-24

Van Rooijen J, Vogel G (2008a) A review of the Dendrelaphis

pictus complex (Serpentes: Colubridae) — I: Description of a

sympatric species. Amphibia-Reptilia 29: 101-115

Van Rooijen J, Vogel G (2008b) A new species of Dendrelaphis

(Serpentes: Colubridae) from Java, Indonesia. The Raffles Bul-

letin of Zoology 56: 189-197

Van Rooijen J, Vogel G (2008c) An investigation into the tax-

onomy of Dendrelaphis tristis (Daudin, 1803): revalidation of

Dipsas schokari (Kuhl, 1820) (Serpentes, Colubridae). Con-

tributions to Zoology 77: 33-43

Van Rooyen J, Vogel G (2009) A multivariate investigation in-

to the population systematics of Dendrelaphis tristis (Daudin,

1803) and Dendrelaphis schokari (Kuhl, 1820): revalidation

of Dendrophis chairecacos Boie, 1827 (Serpentes: Colubri-

dae). The Herpetological Journal 19: 193-200

Vogel G, Van Rooijen J (2007) A new species of Dendrelaphis

(Serpentes: Colubridae) from Southeast Asia. Zootaxa 1394:

25-45

Vogel G, Van Rooijen J (2008) A review of the Dendrelaphis

pictus complex (Serpentes: Colubridae) — 2: the eastern forms.

Herpetozoa 21: 3-29

Vogel G, Van Rooijen J (2011, in press) Contributions to a re-

view of the Dendrelaphis pictus (Gmelin, 1789) complex (Ser-

pentes: Colubridae) — 3. the Indian forms, with the descrip-

tion of a new species from the Western Ghats. Journal of Her-

petology 45

Wall F (1908) Two new snakes from Assam. Journal of the Bom-

bay natural History Society 18: 272-274

Wall F (1910) Notes on snakes collected in Upper Assam. Part

2. Journal of the Bombay Natural History Society 19: 825-845

Wall F (1921) Remarks on the Indian Species of Dendrophis and

Dendrelaphis. Records of the Indian Museum 22: 151-162

Wogan GOU, Vindum JV, Wilkinson JA, Koo MS, Slowinsky

JB, Win H, Thin T, Kyi SW, Oo SL, Lwin KS, Shein AK

(2008) New country records and range extensions for Myan-

mar amphibians and reptiles. Hamadryad 33: 83-96

Ziegler T, Vogel G (1999) On the knowledge and specific sta-

tus of Dendrelaphis ngansonensis (Bourret, 1935) (Reptilia:

Serpentes: Colubridae). Russian Journal of Herpetology 6:

199-208

Received: 20.08.2010

Accepted: 20.02.2011

Corresponding editor: P. Wagner

©ZFMK

Bonn zoological Bulletin

Volume 60 Issue | pp. 25-34 Bonn, May 2011

Studies on African Agama IX.

New insights into Agama finchi Bohme et al., 2005 (Sauria: Agamidae),

with the description of a new subspecies

Philipp Wagner!2*, Wolfram Freund!, David Modry34, Andreas Schmitz5 & Wolfgang Bohme!

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

2 Department of Biology, Villanova University, Villanova, Pennsylvania 19085, USA

3 Dept. of Parasitology, Faculty of Veterinary Medicine, University of Veterinary and Pharmaceutical Sciences

Brno, Palackeho 1-3, 612 42 Brno, Czech Republic

+ Biology Center, Institute of Parasitology, Academy of Sciences of the Czech Republic, BraniSovska 31,

370 05 Ceské Budéjovice, Czech Republic

5 Department of Herpetology and Ichthyology, Muséum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6,

Switzerland

“corresponding author: philipp.wagner.zfmk@uni-bonn.de.

Abstract. We present new information on the distribution and morphology of Agama finchi from eastern Africa. For the

first time, material from three different populations (including the type locality) was available and the question of a pos-

sible subspecies in Uganda was positively answered. Based on the distribution pattern of Agama species groups, a gen-

eral distribution pattern is discussed.

Key words. Reptilia, Sauria, Agamidae, Agama finchi, Agama finchi ssp. n., Africa, Kenya, Uganda, DR Congo, Ethiopia.

INTRODUCTION

Cursorily, the taxonomy and relationships between east-

ern African Agama species were thought to be well known,

but after intensive research on these species, it became ob-

vious that species limits were underestimated and sever-

al taxa were described, revalidated or regarded as syn-

onyms (Bohme et al. 2005; Wagner 2007; Wagner et al.

2008a, 2008b; Wagner 2010). Most Agama species occur-

ring in eastern Africa seem to be part of a monophyletic

group (Wagner et al. unpubl. data). However, the only

member of the Agama agama species group occurring in

eastern Africa is still 4. finchi (Leaché et al. 2009). This

species was described by Bohme et al. (2005) from Mal-

aba in western Kenya close to the Ugandan border. These

authors also mentioned a population from Murchison Falls

in Uganda, but specimens from this population were on-

ly known from photographs. They differed from topotyp-

ical Agama finchi in the colouration of the forelimbs and

until recently, it was not possible to confirm these photo-

graphs with voucher specimens. Additionally, further in-

formation given by Bohme et. al. (2005) show the occur-

rence of A. finchi at the “Yale River in Uganda’, but it was

not possible to trace this locality. Probably the locality

refers to the western Kenyan Yala River, which was in the

‘Uganda’ province of East Africa before 1926 (pers.

comm. Stephen Spawls, 25.1.2011).

A. finchi is a small lizard and is characterized by its bi-

coloured red and black tails, scarlet-red heads and fore-

limbs and the velvet-black bodies of adult males. The

throat colouration is uniform pale reddish, with a pattern

of longitudinal dark reddish lines. The females are differ-

ent in colouration from other Agama species as they pos-

sess a uniform brown body with reddish to yellow dorso-

lateral bands. In both sexes, a white to yellowish supral-

abial line is usually obvious. Nevertheless, differences in

colouration between the specimens from Uganda and topo-

typical A. finchi are obvious, since the former possess

body-like coloured forelimbs. However, without vouch-

er specimens, it was so far not possible to assess the sta-

tus of this population.

Currently, only few Agama species are known from Ugan-

da. Agama lionotus elgonis Lonnberg, 1922 is known to

occur in the Mt. Elgon region, and most probably Agama

finchi, described from an area close to the border to Ugan-

da, should occur there. Spawls et al. (2002) also mentioned

a wide distribution of Agama agama in Uganda, but these

records should be referred to Agama lionotus since A. a.

agama is restricted to Central Africa (Wagner et al. 2009).

26 Philipp Wagner et al.

Agama f. leucerythrolaema ssp. n.

Uganda: Murchison Falls

ZFMK 88809, holotype

Agama f. finchi

Kenya: Malaba

NMK L/2716, topotype

Agama lebretoni

Cameroon: Mamfe

ZFMK 87689, holotype

Agama paragama

Cameroon: Waza

ZFMK 15244

Agama spec.CAR-1

Central African Republic: Koumbala

ZFMK 40252

Agama spec. CHAD

Chad: Lake Chad

MNHN uncatalogued

Agama sp. n.

DR Congo: Nagero

ZFMK 51578

Fig. 1. Throat and left head side of male Agama species.

Bonn zoological Bulletin 60 (1): 25-34 OZFMK

Studies on African Agama IX. New insights into Agama finchi ah

Since the description of A. finchi, specimens from Ugan-

da became available and it was possible to recognize the

species from other countries by images and specimens.

Therefore, the aim of this publication is to summarize and

discuss the distribution of A. finchi and to assess the sta-

tus of the populations possessing body-like coloured fore-

limbs.

MATERIAL AND METHODS

Material and morphological sampling. 21 specimens of

Agama finchi, including type material, were examined.

Specimens from collections of the following institutions

(Institutional abbreviations in parenthesis) were used:

Muséum d’histoire naturelle, Geneve (MHNG); Nation-

al Museums of Kenya (NMK); Zoologisches

Forschungsmuseum Alexander Koenig, Bonn (ZFMK).

Type material of the new species is deposited in the Zo-

ologisches Forschungsmuseum A. Koenig, Bonn.

For each specimen, external characters were recorded:

snout-vent-length (SVL), length of tail (TL), head width

between the anterior margins of the ear openings (HW),

head length from the tip of the snout to the anterior mar-

gin of the ear opening on the left side (HL), head height

at the highest point of the head (HH), number of scales

beneath the 4th toe on the left side (SD), number of scales

around mid-body (MBS), number of scales between gu-

lar- and inguinal fold (V; ventrals), and the number of pre-

cloacal pores (PP). Measurements were taken with a di-

al calliper to the nearest 0.1 mm. Measurements and scale

counts were done according to Grandison (1968) and

Moody & Bohme (1984). Colour patterns in living speci-

mens were described using field notes and life photo-

graphs.

Material examined. Agama f. finchi Kenya. Malaba:

ZFMK 82091-82094, NMK L/2533/3, L/2534/1,3,6. Aga-

ma f. ssp. n. Ethiopia. Gambela: ZFMK 8709-8711,

66271. Uganda. Murchison Falls: ZFMK 88808-814,

88829. Agama lebretoni. Cameroon. Douala, Foyer du

Marin: MHNG 2713.31; Fako (Mt. Cameroon), Limbe

(Victoria): ZFMK 18891—894; Korup, Mundemba: ZFMK

61243; Magba: ZFMK 51686, 54906-907; Makum:

ZMB 55709, 37061; Mamfe: ZFMK 87694—699, 87700;

Mamfe, Mukwecha, Amebisu: ZFMK 87694—-699; Met-

chum, Wum: ZFMK 15194-15200; Mt. Nlonako, Nguen-

gue: ZFMK 69017, Rumpi Hills, Mofako Balue: MHNG

2713.29; Rumpi Hills, Big Massaka: MHNG 2713.30,

2713.32. Equatorial Guinea. Bioko Island, San Carlos:

ZFMK 9353-359. Gabon. Fougamou: ZFMK 73239-245;

Ngouassa: IRSNB 15686—-687. Agama paragama.

Cameroon. Logone et Chari, Waza: ZFMK 15242-256.

Niger. Agadez: ZFMK 36599; Tessaoua: ZFMK

Bonn zoological Bulletin 60 (1): 25-34

33749-750. Agama cf. paragama. Cameroon. Benoue,

Boki: ZFMK 15227-241. Agama cf. sylvanus. Cameroon.

Benoué: ZFMK 33751—754. Agama spec. | CAR-1. Cen-

tral African Republic. Koumbala: ZFMK 40251—260.

Agama spec. | CAR-2. Mélé: ZFMK 33766; Ndeéle:

ZFMK 33755-7654; Sibut: ZFMK 33765. Agama spec.

| Chad Chad. Bol (near Lake Chad): 29171-29221 (will be

inventoried at the collection of the MNHN). Agama sp.

n. Democratic Republic of the Congo. Nagero: ZFMK

51576—-587; Isiro: ZFMK 51588.

RESULTS

Agama finchi leucerythrolaema ssp. n.

Holotype. ZFMK 88809, adult male from Murchison

Falls, Uganda; collected by W. Freund, July 2009.

Paratypes. ZFMK 88810, adult male from Murchison

Falls, Uganda; collected by W. Freund, July 2009.

ZFMK 88811, 88829, adult females from Murchison Falls,

Uganda; collected by W. Freund, July 2009.

ZFMK 88808, 88812—814, juveniles from Murchison

Falls, Uganda; collected by W. Freund, July 2009.

Diagnosis. This is a medium-sized lizard of the genus

Agama (total length of adult males up to 275 mm), which

is characterized by a large gular fold, a reticulated throat

and a bright nuptial coloration of adult males. The throat

colouration (Fig. 1A) is a reticular pattern of red lines,

which thus far is only known from A. paragama (Fig. 1D),

A. sylvanus (no true specimen available), A. /ebretoni (Fig.

1C) and from recently unidentified material from the Cen-

tral African Republic (formerly identified as A. sy/vanus;

Fig. 1E) Lake Chad (Fig. IF) and DR Congo (Fig. 1G).

Females are similar to those of the nominate form.

Differentiated diagnosis. A. finchi leucerythrolaema ssp.

n. not only differs from the nominotypic form by its larg-

er size, but additionally males of the new taxon are dis-

tinct as they possess a large gular fold, a reticulated colour

pattern of the throat (for both see Fig. 1A) and body-like

coloured forelimbs. Differences in pholidosis to the nom-

inate form are only marginal but the new subspecies has

a lower count of body scales (Table 1).

Because of the reticulated throat, the new subspecies is

similar to some other Agama species. From the two Cen-

tral Africa species, A. paragama and A. lebretoni, the new

subspecies differs as follows:

Agama paragama possesses a yellow-whitish to chalk

white head and has a higher number of scale rows around

the midbody (Tab. 1). Additionally, adult males of A.

©OZFMK

ii)

Philipp Wagner et al.

nnes ywiaz —

Fig. 2.

Male holotype (ZFMK 88809) of Agama finchi leucerythrolaema ssp. n. from Murchison Falls, Uganda.

Bonn zoological Bulletin 60 (1): 25-34

©ZFMK

Studies on African Agama IX. New insights into Agama finchi 29

paragama show a black instead of a dark blue tail tip. Sim-

ilar to the herein described new species, Grandison (1968)

described the colouration of the throat of A. paragama as

‘a dark network on a cream ground which takes the form

of isolated, round, cream spots’ (Fig. 1D), which is pres-

ent in both sexes, and therefore similar to A. finchi leucery-

throlaema ssp. n. as the females also have a striated throat

pattern.

A. lebretoni differs from the new subspecies by having a

pale vertebral band and scattered white body scales. Ad-

ditionally, the average snout-vent-length in this species is

larger than in the new subspecies (A. /ebretoni: 117.8 mm;

A. f. leucerythrolaema ssp. n.: 102.4 mm).

The new subspecies differs from the West African, A. sy/-

vanus, in having a higher mid-body scale count. Accord-

ing to MacDonald (1981), A. sylvanus has 59 to 66 scale

rows, whereas A. finchi leucerythrolaema ssp. n. has 71

to 78 rows. The same author described the pattern of the

throat as ‘marked with an irregular pattern of longitudi-

nal whitish lines and (more rarely) small spots’, which is

similar to the herein described subspecies.

A. finchi leucerythrolaema ssp. n. is clearly dissimilar to

the Central African A. a. agama, since the latter possess-

es a uniform to striated red throat, a tri-coloured tail and

a yellow head. However, according to Grandison (1968),

her specimens of A. agama from Nigeria (currently not

traceable if it is A. agama sensu stricto or sensu lato) has

59 to 77 rows, which is similar to A. f- leucerythrolaema

ssp. n. (71 to 78 rows).

A. finchi leucerythrolaema ssp. n. differs from other East

African Agama species as follows:

— from A. lionotus in having a reticulated throat, a tri-

coloured tail and a dark blue body colouration, in hav-

ing lower scale counts around the midbody (A. /. liono-

tus: 67-91 | 75.4; A. 1. elgonis: 79-87 | 81.8; A. finchi

leucerythrolaema ssp. n.: 73.9) and a large gular fold;

— from A. turuensis in having a reticulated throat, a gular

fold and lower count of scale rows around the midbody

(A. turuensis: 71-85 | 77.6; A. finchi leucerythrolaema

ssp. n.: 73.9);

— from A. kaimosae in having a reticulated throat, a gu-

lar fold, a blue body colouration, lower scale counts

around the midbody (A. kaimosae: 79-82 | 80.0; A.

finchi leucerythrolaema ssp. n.: 73.9) and a tricoloured

tail;

— from A. mwanzae in having a reticulated throat, a gular

fold, a blue body colouration, lower scale counts around

the midbody (A. mwanzae: 67-82 | 75.2; A. finchi

leucerythrolaema ssp. n.: 73.9) and a tri-coloured tail;

Bonn zoological Bulletin 60 (1): 25-34

— from A. caudospinosa in having a reticulated throat, a

gular fold, a blue body colouration, lower scale counts

around the midbody (4. caudospinosa: 74-116 | 93.3;

A. finchi leucerythrolaema ssp. n.: 73.9) and a tri-

coloured tail;

— from A. montana, A. mossambica and the much small-

er A. armata in not possessing a heterogeneous body

scalation.

Description of the holotype (ZFMK 88809, Fig. 2)

Habitus stout, snout-vent length 118.2 mm, tail length

150.1 mm, head length 30.3 mm, head width 19.4 mm,

head height 12.8 mm.

Large triangular nasal scale slightly above the canthus ros-

tralis and pierced with the nostril in the posterior part, di-

rected and supplied obliquely upwards. Between the nasal

scales, a single narrow longitudinal smooth scale is visi-

ble, followed by one smooth scale, larger than the other

head scales. Ten supralabial and sublabial scales are on

both sides. Head scales between the eyes are smooth, di-

rected sideward from a midline of two rows of feebly

keeled scale; head scales between posterior end of the eyes

and neck smooth to feebly keeled, directed forwards; head

scales of the temporal region smooth to feebly keeled, not

directed to one side; free anterior margins of head scales

with sensory pits; supraocular scales smooth. Parietal

shield large and more or less pentagonal, pineal organ vis-

ible, pierced more or less in the middle of the shield. Ear

hole large, about the same size as the eye, margin being

composed by spiny scales, surrounded by four tufts of

more or less spiny, mucronate scales; tympanum superfi-

cial. Nuchal crest low, consisting of 13 lanceolate scales.

Gular scales flat, smooth, juxtaposed and becoming small-

er towards the large gular fold. Dorsal body scales strong-

ly keeled, but becoming feebly keeled at the vertebral re-

gion, mucronate, equal in size, in 56 scales from midpoint

of pectoral region to midpoint of the pelvic region. Ven-

tral body scales smooth, slightly imbricate at their poste-

rior margins, in 69 scales from midpoint of pectoral re-

gion to midpoint of pelvic region. There are 72 scales rows

around the midbody. Ten precloacal scales stringed in one

row only. Tail scales strongly keeled and mucronate.

Scales on the upper side of the forelimb strongly keeled,

smooth on the underside, on the upper arm scales twice

as large as the dorsal body scales, becoming smaller to-

wards the underside and the manus. 4" finger longest, dig-

ital length decreasing 3-2-5-1, subdigital lamellae keeled

and mucronate. Scales on the upper side of the hindlimb

strongly keeled, becoming smooth on the underside, on

the upper tights slightly larger in size than the dorsal body

scales, becoming larger towards the lower tights. 4" toe

longest, digital length decreasing 3-2-5-1.

©ZFMK

30 Philipp Wagner et al.

Fig. 3.

Living specimens from the type localities (A) male and (B) female of Agama finchi leucerythrolaema ssp. n. from Murchi-

son Falls, Uganda (photos by David Modry); (C) male and (D) female of Agama finchi finchi from Malaba, Kenya (Photos: Bri-

an Finch).

Colouration. (in alcohol after three months of preserva-

tion). Head and neck red with a light red vertebral band

extending to the back. Limbs, body and anterior third of

the tail dark blue. Tail tri-coloured, at the base dark blue,

followed by red and dark blue. Belly and underside of the

limbs blue, tail anterior whitish, posterior bluish. Throat

and the large gular fold with a reticulated pattern of red

stripes on a white background.

Colouration in life. Males. Head, neck and parts of the

shoulders red, a broad vertebral red band extends on the

back from head to about half way to hindlimbs. Body,

limbs and anterior third of the tail dark blue. Tail tri-band-

ed: dark blue (extending from the body), followed by red

and dark blue at the tip (Figs 3a, 4a). Belly and underside

of limbs blue; throat with a reticulate pattern of red stripes

on a white background.

Females. Head yellowish brown, with fine yellow stripes

and dots; stripes and dots usually extending to the shoul-

ders. Body and tail light brownish with broad lateral yel-

low bands (Fig. 3b). Underside whitish, with fine dark lon-

gitudinal stripes on the throat.

Bonn zoological Bulletin 60 (1): 25-34

Juveniles. Similar to females but with more distinct yel-

low dots and stripes on the head on the shoulders. Yellow

lateral bands sometimes extending to the back. Underside

whitish, with a fine dark reticulated pattern on the throat.

Variation. Variations in morphology are shown in the ap-

pendix and compared to other agamid species in Table 1.

Colouration of males is not variable and all show the typ-

ical colour pattern of throat and forelimb.

Etymology. The new species is named after its remark-

ably characteristic red and white vermiculated throat as

compared to the nominate form. The name is derived from

the Greek words ‘leukos’ for white, ‘erythros’ for red and

‘laema’ for the throat.

Distribution. Beside the type localities of the two sub-

species, A. finchi was also identified from Gambela in

Ethiopia, represented by material from the collection of

the Zoologisches Forschungsmuseum A. Koenig (ZFMK

8709-8711; ZFMK 66639, 66271—72). In contrast to the

specimens of A. f. finchi from Kenya and DR Congo (see

below and figs 3, 5), the adult males of this population

©ZFMK

Studies on African Agama IX. New insights into Agama finchi 3]

Fig. 4.

(A) Living male and (B) habitat of Agama finchi

leucerythrolaema ssp. n. from Murchison Falls, Uganda (Pho-

tos: Wolfram Freund).

show body-like coloured forelimbs and therefore repre-

sent the new subspecies. The colouration of the throat is

not distinguishable, because of the long preservation time.

However, specimens from Gambela are similar in the av-

erage of their snout-vent-length to topotypical material of

the nominal subspecies (Ethiopia: 86.2 mm | Kenya: 85.7

mm). A. f/ leucerythrolaema ssp. n. is also cryptically men-

tioned in Largen & Spawls (2010). Within the species

chapter on Agama agama (sic., as Agama agama in nei-

ther occurs in Ethiopia nor Eritrea), specimens from Gam-

bela are figured (Largen & Spawls 2010, fig. 147) show-

ing the herein described subspecies. According to Stephen

Spawls (pers. comm. 07.X.2010), the western population

of the former ‘Agama agama’ in Ethiopia and Eritrea be-

longs to A. finchi, whereas the southern ones belong to

Agama lionotus.

Quite recently, the new subspecies was also recorded from

the Lorionotom Range, Ilemi Triangle (approx coordi-

nates: 4°53738.53”N, 35°31°59.44”E) by Miroslav Jirku

(specimens will be inventoried in the collection of the Na-

tional Museums of Kenya, Nairobi), which fills the gap

between the Ethiopian and Ugandan localities.

Habitat. Specimens at Murchison Falls were collected in

open grassland (Fig. 4b), sitting on old houses (Fig. 4a)

Bonn zoological Bulletin 60 (1): 25-34

and on palm trees. The habitat of the Ethiopian popula-

tions is unknown, but is likely to be similar.

Relationships. According to Wagner et al. (unpubl. da-

ta) Agama finchi sensu lato is differentiated in two clades.

Individuals from the type locality of the nominate form

are distinct in colouration of forelimbs, throat, and in body

size to specimens from Uganda and Ethiopia, but there are

few other morphological differences (see Appendix). Ge-

netic analysis indicates (Wagner et al. unpubl. data) that

the new taxon is closely related to A. finchi and both taxa

are members of the Agama agama species group.

Agama f. finchi: new distribution record. Democratic

Republic of the Congo. The nominate species was docu-

mented by photographs from the Ituri forest (Fig. 5) where

it occurs on an inselberg surrounded by rainforest is not

otherwise connected to savannah areas. Ecologically, this

inselberg is an arid area, as rainwater flows off immedi-

ately, and the unshaded dark rocks are heated up by sun-

light during the day. Therefore, fauna and flora (with e.g.

rock hyrax and aloe plants; pers. comm. Reto Kuster) of

this inselberg are very different from those of the surround-

ing rainforest. The status of these forests as a true rain-

forest is supported by the occurrence of Lepidothyris

hinkeli Wagner et al., 2009, which is a character species

of equatorial rainforests (Wagner et al. 2009). As can be

seen from the figures 4 C—D, the specimens are identical

in colouration to the specimens from the type locality of

A. f. finchi in western Kenya. However, the throat coloura-

tion is still unknown. The distribution of both taxa is

shown in figure 6.

DISCUSSION

Preliminary genetic analyses (unpubl. data) separated Aga-

ma finchi sensu lato into two distinct clades: this was sup-

ported by differences in the colouration of the forelimbs

of adult males. However, there are more morphological

differences between the Ugandan and Kenyan populations

than between the Ugandan and Ethiopian. The Ugandan

population is distinct from the Kenyan population because

of the reticulated throat and the much larger size of adult

males. Also, the Ugandan population is larger than the

Ethiopian population. Furthermore, only small series were

collected from all populations, and body-sizes are prob-

ably underestimated. Because the Ethiopian specimens

were preserved a long time ago, the throat colouration is

not identifiable anymore.

Little information is available about the species limits

within the genus Agama. In contrast to genetic distances

in-between other Agama species, the distances within the

A. finchi sensu lato populations are comparatively small.

©ZFMK

32 Philipp Wagner et al.

Fig. 5.

However, identification of the subspecies is possible and

demonstrated by pholidosis, differences in colouration of

adult males, and the colour pattern of the throat. There-

fore, these populations should be recognized as a devel-

oping species and according to article 45.6 of the Inter-

national Code of Zoological Nomenclature (ICZN 1999),

they can be recognized as the taxonomic rank of a sub-

species.

Distribution patterns and relationships are poorly known

in Agama lizards. For decades, the genus was not in the

focus of herpetological interests and most of the taxa were

recognized as subspecies of Agama agama which, because

of this original taxonomic classification, was supposed to

be distributed nearly everywhere in sub-Saharan Africa.

Therefore, in the older literature, A. agama is supposed

to have a distribution range with includes nearly every

Afrotropical country. Additionally, many specimens are

misidentified or simply inaccurately labelled as A. aga-

ma in museum collections. Wagner et al. (2009) described

a neotype and restricted A. agama s. str. to northern

Cameroon. The whole A. agama species complex is now

Bonn zoological Bulletin 60 (1): 25-34

Agama f. finchi occurrence on an inselberg within the Ituri Forest, DR Congo (A) View to the inselberg within the for-

est. (B) Habitat on the inselberg. (C) Living male of 4. f. finchi from the same locality. (D) Living female of 4. f finchi from the

same locality (Photos: Reto Kuster).

distributed along the southern border of the Sahara desert

from western to eastern Africa and along the Atlantic coast

to Namibia. Other than that, former East African sub-

species of A. agama are now recognized as a distinct

species (BoOhme et al. 2005), forming the A. /ionotus

species group, which is distributed from Ethiopia through

Kenya to southern Tanzania. However, the distribution pat-

terns of the 4. agama and A. lionotus species groups gen-

erally follow the theory of African arid corridors and 1s

comparable to distributions of other reptile species

groups, such as Varanus exanthematicus/ albigularis (see

Wagner 2010).

The only representative of the 4. agama species complex

in eastern Africa (including Ethiopia) is A. finchi, as none

of the other A. agama species complex taxa are present

in Ethiopia and Uganda (pers. comm. Steven Spawls). To

date, both subspecies of A. finchi have only been found

west of the eastern branch of the East African Rift, but it

is not possible to recognize this geological barrier as a dis-

tribution limit since this species 1s only known in four lo-

calities.

©ZFMK

Studies on African Agama IX. New insights into Agama finchi 33

Fig. 6. Distribution of Agama finchi. Red dots refer to A. f

finchi, whereas blue dots refer to A. f- leucerythrolaema ssp. n.

(1) Kenya: road to Busia near Malaba (type locality); (2) DR

Congo (photo documention); (3) Uganda: Murchison Falls (type

locality); (4) Ethiopia: Gambela. Other Ethiopian localities ob-

tained from Largen & Spawls 2010; (5) Kenya: Lorionotom

Range, Ilemi Triangle (approx coordinates: 4°53’38.53”N,

35°31°59.44”E).

Distribution was also influenced by the dispersal and re-

traction of the equatorial rain forest in the past. The pop-

ulation in the Ituri Forest is a relict population found on

an inselberg surrounded by rainforest, usually a non-suit-

able habitat for Agama species. The status of the Guineo-

Congolian rainforest is supported by the occurrence of

Lepidothyris hinkeli, which is a character species of this

forest type (Wagner et al. 2009). Therefore, the rainforests

in this area seem to be relatively young as A. finchi was

enclosed on its inselberg during an extension of the rain-

forest. As a result, the species must have been widely dis-

tributed within the area during the time when the rainfor-

est last receded.

Acknowledgements. We are grateful to Reto Kuster for

permission to publish his images of Agama finchi from

DR Congo and his comments on the occurrence of this

species in the DR Congo. Moreover, we are grateful to

Stephen Spawls and Miroslav Jirku for their information

about A. finchi in Ethiopia and Kenya. We thank the two

referees for their comments which have improved the

manuscript, and especially to Shelley Barts-Pankow who

did the English review.

Bonn zoological Bulletin 60 (1): 25-34

REFERENCES

Bohme W, Wagner P, Malonza P, Lotters $, Kohler J (2005) A

new species of the Agama agama group (Squamata: Agami-

dae) from western Kenya, East Africa, with comments on Aga-

ma lionotus Boulenger, 1896. Russian Journal of Herpetology

12: 83-90

Grandison AGC (1968) Nigerian lizards of the genus Agama

(Sauria: Agamidae). Bulletin of the British Museum of Nat-

ural History Zoology 17: 67-90

ICZN (1999) International Code of Zoological Nomenclature.

International Trust for Zoological Nomenclature, Fourth Edi-

tion: 306 pp.

Largen M, Spawls S (2010) The amphibians and reptiles of

Ethiopia and Eritrea. Edition Chimaira, 693 pp.

Macdonald MA (1981) A new species of agamid lizard from

Ghana. Journal of Zoology 193: 191-199

Moody SM, Bohme W (1984) Merkmalsvariationen und taxo-

nomische Stellung von Agama doriae Boulenger, 1885 und

Agama benueensis Monard, 1951 (Reptilia: Agamidae) aus

dem Sudangiirtel Afrikas. Bonn. zool. Beitrage 35: 107-128

Spawls S, Howell K, Drewes RC, Ashe J (2002) Field Guide to

the Reptiles of East Africa: All the Reptiles of Kenya, Tanza-

nia, Uganda, Rwanda and Burundi. Princeton Field Guides:

543 pp.

Wagner P (2007) Studies in African Agama I — On the taxonom-

ic status of Agama lionotus usambarae Barbour & Loveridge,

1928 (Squamata: Agamidae). Herpetozoa 20: 69-73

Wagner P, Burmann A, Bohme W (2008a) Studies on African

Agama IU. Resurrection of Agama agama kaimosae Loveridge,

1935 (Squamata: Agamidae) from synonymy and its elevation

to species rank. Russian Journal of Herpetology 15: 1-7

Wagner P, Krause P, Bohme W (2008b) Studies on African Aga-

ma III. Resurrection of Agama agama turuensis Loveridge,

1932 (Squamata: Agamidae) from synonymy and its elevation

to species rank. Salamandra 44: 35—42

Wagner P, Wilms TM, Bauer A, Bohme W (2009) Studies on

African Agama V. On the origin of Lacerta agama Linnaeus,

1758 (Squamata: Agamidae). Bonner Zoologische Beitrage

56: 215-223

Wagner P (2010) Diversity and distribution of African reptiles.

Unpublished PhD thesis, University of Bonn: 374 pp.

Received: 25.11.2010

Accepted: 25.02.2011

Corresponding editor: F. Herder

OZFMK

Philipp Wagner et al.

“SqUII] UdaMjoq YISUI] =Tq'T ‘seiod [eovopsaid Jo saquiNU =GOd £90} pp YO] Youdg sejjowe] =o], “IOBuLy wh Yo] you

-9q se][oule] =I ‘Apogprur punose sayeog =es ‘el[esiog =YOR ‘eyeNUsA =NAA ‘ysusy peoy =TH Spm peoy =H WYs1oy peoy =H ‘yISugy [ej0} =ToY ‘ySugy prey =Tp ‘yBue]-yuaa-jnous =T AS

qyeulay = | Ep = fii Ol @2 &) —s3 vl TZ ST «OST «LOL LOTT) «6VHEL) «= 9B STP YOINY :epuesq = HZ88 MINAZ = Pwavjo.ysCiaonay fy

ayeuraz = Lp = 6 Mo 7 @ ST €T TW VST $6 BEET VMEPI T06 Sled Younpy :epuesEy 11888 MANAZ Yuanjosyjsduaonay fy

qeu 709 ?@l td 61 SL OL 18 vl OT 98Z OOT OPI PH8LT TI TIT ses YoNpl epues O18 MNAZ MMavjosysdiaonaj fp

qe = 09 I 6 8 “re 09) 08 91 TT 96C OT EI = — P8Il sey yom sepuedssy,) = 60888 MINAZ PManjosyjduaonay f

jaw Ci Gl i Gl Sh to 77 I €% 67 6ST TOT ie =. OSs vjoquiey :eidomg 1L799 MINAZ « PulavjosysAiaonay f -¥

qeur Oh 2ST SsCETSC‘«i‘iSTSC(‘<‘zTLSCSCODC v1 GG Sie UN Ell “PSCC - <Gi6ET — SiS8 vpoquiey ‘vido 60L8 MINAZ = Puavjosysdiaonay fp

gyeuray =F Ob = Ie Oe Wh Wer 82 vl OCe-CCGeetOll “Gil (S90. TRL =7/S8 Bjaquied :eidompg IIL8 MNAZ = Puavjosyjdsaonay fy

geuay = 8 Th = IG “Gl OL =Gi- “08 el LeCaeGiGG- 89 VOL SCG “SLED —iGi88 vpoquiey ‘vido OIL8 MNAZ = Puavjosysdsaonay fp

ja Bide Gi ie Sl GR Ge 4s Gal HiGenGeGe SES “SiG SECYE —x9iVS1, “LL eqeey :eAuay = Z/P€ST/T MINN ryouy f¥

S[EUISNN Sit pao CCOpeSIO NO. iG 198 Gl Ge Sule SAN SIG? “PSG 8169" 29/88 gery :eAuoy = 76078 MINAZ 1youy f-¥

yeu 6h )6=6C7TiCté=‘é‘=CS“OCTY = = 173 a vo SET E91 OO BIO xOEZI 88L eqeieyl :eAuay 16078 MINAZ ryouyf f -¥

geury = LIZ = @ Ol 7 03° G3 eT Ce Gi A9ReM VL €€91 G6 €OI 6S gery eAuay = €/E€ST/T MINN ryouyf f -¥

‘ant 7H = (6 S19 oh Oy 33 ia 17 9El $6 99 FOL «POE 09F gee eAuoyy 1 /P€ST/1 MINN you f -V

ant £81 = 1G Of Of 3) 28 Tl SI TIL €6 €9 OS6 «F9S 98E eqeey :eAuay — €/PEST/T MINN 1youy f¥

‘anf SZ =" 16 Of We 8 083 ST HE -=E9T Sl “819 <9I6ZT. EE 22S que :eAuay = 9/P€ST/T MINN 1youy f¥

‘anf 767 = 1¢-- Br OL = 1 Teal HG CHA Gall 6b 6€8 «£€6€ OP eqeey :eAuay P6078 MINAZ 1youyf of -¥

‘ant 7 = 062 fil. QE ee 1 tl vo Trl Ol 6S E€TIT +€89 Obb eqeeyy :eAuay €6078 MINAZ you fy

X9§ TET dOd 99h) = WS YOM NAA MH/TH HH/TH TH MH HH TOL TL ‘TAS AqeI0T ‘ON uOXxeL

APPENDIX

©OZFMK

Bonn zoological Bulletin 60 (1): 25-34

Bonn zoological Bulletin |

Volume 60 | Issue | pp. 35-61 Bonn, May 2011

The reptiles (Testudines, Squamata, Crocodylia)

of the forested southeast of the Republic of Guinea (Guinée forestiére),

with a country-wide checklist

Wolfgang B6hme!*, Mark-Oliver Rédel?, Christian Brede? & Philipp Wagner!

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

2 Museum fiir Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt

University Berlin, Invalidenstr. 43, D-10115 Berlin, Germany

3 Medizinische Klinik und Poliklinik I, Zentrum fiir Experimentelle Molekulare Medizin, Zinklesweg 10,

D-97078 Wurzburg, Germany

“ Corresponding author: E-mail: w.boehme.zfmk@uni-bonn.de.

Abstract. During several herpetological surveys in the forested southeast of the Republic of Guinea (Guinée Forestiére)

we recorded 64 species of reptiles (two chelonian, 16 lizard and 45 snake species as well as one crocodile species. They

are presented in a commented list with documentation of the respective voucher material and with taxonomic and/or eco-

logical information. Our record of the softshell turtle Zrionyx triunguis is the second one for Guinea and the first for Guinée

Forestiére, the forest-dwelling gecko Cnemaspis occidentalis and the lacertid lizard Holaspis guentheri are likewise the

respective second Guinean record of these species. The Forest Nile Monitor Varanus ornatus is documented for the first

time in Guinea. The ground boa Calabaria reinhardtii is again the second country record but the only voucher specimen

available for study. Moreover, we present a country-wide checklist of the reptiles of the Republic of Guinea combining

literature records with our own material. It documents the occurrence of 128 reptilian species (seven chelonian, 30 lizard,

88 snake and three crocodile species). Remarkable are photographic records of a probably new, undescribed gecko species

of the genus Hemidactylus. The skink Trachylepis keroanensis, formerly a synonym of T. perroteti, proved to be a dis-

tinct species belonging to the 7) buettneri/sudanensis group. Finally, we provide a list of fishes found in Ziama Forest

partly recovered from the stomachs of natricid snakes.

Key words. Reptilia; Republic of Guinea; Ziama Forest; country-wide checklist; list of Ziama Forest fishes.

INTRODUCTION

The Republic of Guinea belongs to the lesser known coun-

tries in West Africa (see Barnett et al. 1994 and references

therein). Despite its variety of landscapes which range

from dry savannas in the northeast to marshy mangrove

areas in the west, and through the remarkable Fouta Djal-

lon highlands to the forested areas in the southeast (Lam-

otte et al. 1962, Porembski et al. 1994, 1995), only few

reptile collections have been made in this country. It is

characteristic of this low level of zoological exploration

that Guinea did not even “exist” in the country list of

Welch’s (1982) “Herpetology of Africa”! The only region

where comparatively much work has been done, is the Mt.

Nimba range in the extreme southeast where Guinea,

Liberia and Ivory Coast meet (in part Villiers 1950, An-

gel et al. 1954 a,b, Lamotte 1983 for a summary, and In-

eich 2003 for a recent summary including also the Ivo-

rian and Liberian parts of this important mountain

range). Whereas the amphibian fauna of Guinea received

considerable attention in recent years (Rédel & Bangoura

2004a; Rodel et al. 2004, 2009, 2010; Hillers et al. 2006,

2008 a, b, c, plus many so far unpublished surveys and

data), the reptiles of this country remained poorly known.

Among the few sources reporting (at least partly) reptil-

ian voucher material from Guinea exceeding the Mt. Nim-

ba range are the papers by Mocquard (1908), Klaptocz

(1913), Chabanaud (1916, 1917, 1918, 1920, 1921), Park-

er (1939), Villiers (1950), Grandison (1956), Condamin

(1959), and Greenbaum & Carr (2005).

In October 1993, the first author (WB) had the opportu-

nity to visit the Republic of Guinea relatively shortly af-

ter major political changes in the country enabling him to

work as the first herpetologist in Guinea again, after the

French workers in the colonial 1950-ies. He was invited

by Dr. Wilfried Bitzler as a so-called short-time expert

to do a survey of amphibians and reptiles within a proj-

ect of PROGERFOR (=Projet de Gestation des Ressources

Forestieres), Conakry, in the two southeastern rain forests

36 Wolfgang Bohme et al.

Forét de Ziama and Forét de Diécké. During this mission,

also the Nimba Mts. were briefly visited. An itinerary can

be found in the project report by Bohme (1994 a). So far,

only few aspects of this mission have been published, viz.

accounts on some remarkable frogs and skinks collected

during the stay (BOhme 1994 b, c), including a skink

which proved to be new to science (B6hme et al. 2000).

Moreover, a study of the speciose snake community found

in Ziama forest was published (B6hme 2000), containing

several new country records. In addition, some more spec-

imens and important voucher photographs of reptiles were

subsequently provided by W. Bitzler and his mammalog-

ical colleague Dr. Henning Vierhaus, Soest. All these ma-

terials are deposited in the Zoologisches Forschungsmu-

seum A. Koenig in Bonn (ZFMK). A few further speci-

mens were traced in the Zoological Museum of the Uni-

versity of Copenhagen (ZMUC) which had been collect-

ed and sent to Denmark in the 1950-ies by the Danish zo-

ologist/entomologist Herold Olsen who was a resident of

N’Zérékoré, SE Guinea (see Bohme 1994 b). He had kept

there, in his “Centre Entomologique” in N’Zérékoré also

some herpetological specimens which were guarded, af-

ter his death, along with his other collections, by two

Guinean women formerly employed by him, still in 1993

(see BOhme 1994 b).

In 2003, the second author (MOR) started visiting

Guinea, and carried out herpetological surveys with a main

focus on amphibians in several southeastern forested are-

as (Forét de Diécké, Mt. Béro, Forét de Déré, Pic de Fon

— Simandou Range, Mt. Nimba: see Rédel et al. 2004,

Rodel & Bangoura 2004a, b, 2006). The third author (CB)

surveyed amphibians on Mt. Nimba (2007), Pic de Fon

and Mt. Teétini (2008) and the Fouta Djallon and Ziama

forest (2010). The amphibian collections have partly been

published (Rédel et al. 2004 and see above), while some

reptile specimens which were also collected (see Rédel

& Bangoura 2004b, 2006) and donated to ZFMK and the

Museum fiir Naturkunde Berlin (ZMB), plus records of

non-collected reptile specimens (photo records) are also

included in the present paper.

Apart from the mostly silvicolous reptile fauna from SE

Guinea, some specimens were found or observed between

Conakry and Macenta by Wilfried Biitzler, Henning Vier-

haus and Wolfgang Bohme. These were Hemidactylus sp.

(nov.?) from Coyah (photographic voucher see Figs 27—

28); Bitis arietans between Kissidougou and Guéckédou:

photographic voucher; Crotaphopeltis hotamboeia

(ZFMK 54897) and Psammophis cf. phillipsi (ZFMK

56137) from Kissidougou; Toxicodryas pulverulentus

from 25 km southeast of Guéckédou (ZFMK 56136).

Museum Koenig had also received a small number of

specimens collected in winter 1996/97 by G. Nikolaus in

Bonn zoological Bulletin 60 (1): 35-61

P SANMICUELLIE

Fig. 1. Map of SE Guinea (Guinée Forestiére) with the for-

est regions surveyed: 1. Ziama Forest, 2. Diécké Forest, 3. Mt.

Nimba, 4. Mt. Béro, 5. Déré Forest, 6. Pic de Fon (map: P. Wag-

ner).

the Haut Niger National Park. These specimens, includ-

ing the first record of the black mamba for Guinea, were

left by us for publication to Eli Greenbaum to complement

his and J.L. Carr’s paper on the herpetofauna of this park

(see Greenbaum & Carr 2005). They are summarized here

Fig. 2.

Village of Sérédou, Ziama Forest, with forested slopes

in background (Photo: W. Béhme).

OZFMK

Amphibians and reptiles from Guinea 37

he Se en SR Ky |

Fig. 3. “Antenna hill” at Sérédou, submontane forest with the

tree fern Cyathea manniana (Photo: W. Bohme).

for completeness’ sake: Agama agama (ZFMK

64473479), Chamaeleo gracilis (ZFMK 64489), Varanus

exanthematicus (ZFMK 64471: head only), V. niloticus

(ZFMK 66470), Crotaphopeltis hotamboeia (ZFMK

64467-468), Grayia smithii (ZFMK 64465—466),

Philothamnus irregularis (ZFMK 64469), Psammophis el-

Fig. 4. Inundated lowland forest at Malweta village, Ziama

Forest (Photo: W. B6hme).

Bonn zoological Bulletin 60 (1): 35—61

egans (ZFMK 64462464), Dendroaspis polylepis

(ZFMK 64459460), Elapsoidea semiannulata (ZFMK

64461), and Bitis arietans (ZFMK 64472).

Some more single specimens from near Kindia had been

donated to ZFMK also by Dr. Guy Kremer, Luxembourg:

Chamaeleo gracilis (ZFMK 87583—584), C. senegalen-

sis (ZFMK 87581—582). Particularly remarkably:

Sébastien Trape, Dakar, kindly donated a specimen of the

rare Agama insularis from the Los islands to ZFMK where

it has been catalogued under ZFMK 88247.

Within forested SE Guinea (Fig. 1), the focus of this pa-

per is on Ziama Forest which turned out to house a par-

ticularly speciose snake community (42 species: see

Bohme 2000), at least in regard of the short periods spend

for collecting. Ziama Forest is a hilly, forested area (ca.

1300 km’, up to ca. 1600 m a.s.1., with about 70.500 ha

dense forest: ATLANTA Consult 1988) ranging from Ma-

centa in the north to N’Zébéla in the south, and encom-

passing the small town Sérédou (Fig. 2) with its so-called

“Mont d’antenne” (on which the local wireless mast is lo-

cated) (Fig. 3) and several villages: Balassou a few km in

the north, Malweta in the south (characterized by swampy

forest: Fig. 4) and Souzunzou in the west of Sérédou. Pri-

mary forest is best preserved on slopes (Fig. 5). Moreover,

Diécké Forest (700 km?) and the western slope of Mt.

Nimba (Fig. 6) were visited during the mission of WB in

Oct. 1993.

Further Guinean sites where surveyed with a main empha-

sis on amphibians. From 27 November to 6 December

2002, in autumn 2004 and in September 2008 we surveyed

the Simandou Mountain Range, which extends for 100 km

from Komodou in the north to Kouankan in the south. The

altitudinal range is about 600 m with the Pic de Fon at the

southern part being the highest peak (1,656 m a.s.1.: Fig.

7). Approximately 25,600 ha of this forest were protec-

Fig. 5. Ziama Forest near Sérédou: reforestation of fire-de-

stroyed foreground with Zerminalia sp., slopes in the background

still with primary forest (Photo: W. Bohme).

©ZFMK

38 Wolfgang Bohme et al.

Fig. 6.

Mt. Nimba, at 1750 m (Photo: W. B6hme).

ted in 1953, but larger parts are currently explorated for

iron ore. The Simandou range is in the transition between

the forest and savanna zones, offering a wide range of dif-

ferent habitat types. Especially the rain and montane

gallery and ravine forest on the western slopes range far

more North than anywhere else in West Africa (for more

details see Rodel & Bangoura 2004a, b and literature cit-

ed therein). Three other forest sites were surveyed in No-

vember/December 2003: The Diécké Classified Forest, sit-

uated about 25 km south of N’Zérékoré, comprises an area

of 59,143 ha, with a mean altitude of 400-500 m a.s.1. The

reserve comprises (almost) primary as well as secondary

and highly degraded rainforest (for more details see Rédel

et al. 2004). Currently it is under mining prospection ac-

tivities. The Mont Béro Classified Forest (26,850 ha) is

situated at the northern limit of the rainforest zone, 56 km

north of N’Zérékoré, 52 km south of Beyla and 40 km

west of Lola. Its highest elevation is 1,210 m a.s.l. The

dominant habitat types are semi-evergreen forest (Fig. 8)

and savanna (Rédel et al. 2004). The Déré Classified For-

est is situated at the eastern base of Monts Nimba and di-

rectly borders Ivory Coast. It comprises lowlands and hill-

sides (highest peak Mont Ticton, 740 m a.s.1.). Original-

ly the vegetation consisted of evergreen rainforest, but

most of the reserve is now ina very degraded state. From

22 April to 12 Mai 2005 we surveyed several sites in the

Préfecture de Boké in north-west Guinea: Sarabaya (Rio

Kapatchez), Kamsar et Boulléré. The survey focused on

the (few) remaining forested sites and humid zones (for

details see Hillers et al. 2006, 2008c). CB surveyed fur-

ther sites in southeastern Guinea incl. Simandou and Mt.

Nimba, the Ziama forest, and several sites in the Fouta

Dyallon.

Fig. 8. Mt. Béro with one of its mountain creeks (Photo: M.-

O. Rodel).

This paper aims to make the faunistic and autecological

data of our material available, and to summarize the cur-

rent state of knowledge of the reptile fauna of the Repub-

lic of Guinea in the form of a checklist.

Fig. 7.

Pic de Fon showing rain forest remnants (Photo: M.-

O. Rédel).

Bonn zoological Bulletin 60 (1): 35-61

Fig. 9.

Rodel).

Kinixys erosa from Diécké Forest (Photo: M.-O.

©ZFMK

Amphibians and reptiles from Guinea 39

COMMENTED SPECIES LIST

Testudines

Testudinidae

Three species of tortoises, all of the genus Kinixys, are

known from Guinea. While K. be/liana is known from the

drier north (Chabanaud 1921, Greenbaum & Carr 2005),

the forest species K. homeana has been recorded by Cha-

banaud (1921) from N’Zébéla just south of Ziama For-

est. The third species is likewise silvicolous:

Kinixys erosa (Schweigger, 1812)

Material examined: voucher photographs only.

Remarks: Several photographs by W. Bitzler, document

the occurrence of this species in Ziama Forest near Séré-

dou where it does not seem to be rare: a hatchling from

December 1991, and two adults in March 1995. MOR

found it in Diécké Forest (Fig. 9). Villiers (1958) gave only

the imprecise information “depuis Sierra Leone jusq’au

Nord de l’ Angola” et a Uganda. Cité aussi de Gambie (?)”.

The latter questionable record is cited as a fact by Wer-

muth & Mertens (1961): “Von Gambia stidwarts bis Bel-

gisch-Congo”. In contrast, Pritchard (1979) listed the

countries with reliable records separately: in West Africa

only Ivory Coast, Liberia and Sierra Leone. Ineich (2003),

however, recorded two specimens from the Guinean part

of Mt. Nimba, thus proving the existence of K. erosa in

this country. Our photographic vouchers provided the sec-

ond Guinean locality and the first site outside of Mt. Nim-

ba. However, recent work revealed this tortoise to be more

broadly distributed in Guinée forestiére, viz. also in Déré

and Diécké Forests as well as on Mt. Béro (Rédel & Ban-

goura 2006).

Fig. 10. Zrionyx triunguis from Ziama Forest (Photo: W. Biit-

zler).

Bonn zoological Bulletin 60 (1): 35-61

Trionychidae

Trionyx triunguis (Forsskal, 1775)

Material examined: voucher photographs only.

Remarks: Again, a photograph by W. Bitzler (Fig. 10)

documents the occurrence of this softshelled turtle in Zia-

ma Forest. In his monograph on this species, Gramentz

(2005) enumerates the distributional records for each

country from which 7? triunguis is known, Guinea not be-

ing among them. However, Guinea Bissau. Sierra Leone

and Liberia are (see also Loveridge & Williams 1957) so

that the presence in Guinea is not unexpected. The first

documented record from Guinea is by Greenbaum & Carr

(2005) who found it in the Pare National Haut Niger, our

record from Ziama representing the second for the coun-

try and the first for Guinée forestiére.

Squamata

Agamidae

There are four nominal species of the genus Agama known

from Guinea: Agama agama (Linnaeus, 1758), A. crista-

ta Mocquard, 1905, A. insularis Chabanaud, 1918, and A.

sankaranica Chabanaud, 1918. While Agama agama is a

species complex common but nonetheless taxonomically

problematic (see below), A. cristata (Pays Sankaran) and

A. insularis (Los Islands off Conakry, Kindia) are geo-

graphically very restricted. Morphologically, A. boulen-

geri from Mali and Mauritania seems to be their sister tax-

on (Wagner et al. 2009). We want to stress here that we

disagree with Barabanov (2008) who proposed a new

name for A. cristata claiming its preoccupation by A.

cristata Merrem, 1820 (= Corythophanes cristatus, Igua-

nia: Corythophanidae) as A. maria. Our argumentation is

found in Wagner & Bohme (2009). A. sankaranika, final-

ly, is known from several localities in Guinea

(Moussaia/Pays Sankaran, Kankan, Kérouané, Beyla, Ma-

centa, and between Macenta and N’Zébela (Chabanaud

1921). The two latter sites are in the forest zone, the last

one even at the southern edge of Ziama Forest which 1s

remarkable for a species considered to be a savanna form

(Grandison 1968, Hoogmoed 1968, Joger 1979, Joger &

Lambert 2002). Grandison (1968) overlooked Cha-

banaud’s (1921) records and cited only the original de-

scription of A. sankaranica. Rodel & Bangoura (2006)

recorded A. sankaranica also from Mt. Beéro.

Agama agama (Linnaeus, 1758) complex

Material examined: ZFMK 56080—088, Sérédou; ZFMK

56127—128, N’Zérékoré, W. Bohme coll. 6—26 Oct., 1993.

©OZFMK

40 Wolfgang Bohme et al.

Remarks: The “margouillat” of the local people is wide-

ly distributed and very common in human habitations (ma-

jor cities as well as small villages), forest edges and clear-

ings which it penetrates from its primary savanna habi-

tats along roads.

A. agama is in urgent need of revision and 1s likely to con-

tain more than one species also in West Africa (for East

Africa members of the species complex see e.g. BOhme

et al. 2005). This view is strongly supported by a female

described by Klaptocz (1913) possessing more than twice

as big scales as compared to other specimens from the

same locality. Similarly big-scaled agamas have been pho-

tographed also by Dr. Guy Kremer (pers. comm.) in the

western parts of Guinea. Attributing available names to

the Guinean population(s) (africana Hallowell, bocourti

Rochebrune, savattieri Rochebrune) seems premature be-

fore completion of a major revision currently carried out

by one of us (PW).

Chamaeleonidae

Chamaeleo gracilis Hallowell, 1844

Material examined: ZFMK 56142, subadult, forest near

Malweta, southern Ziama Forest, W. BOhme coll. 6-26

Oct., 1993.

Remarks: C. gracilis is the only chameleon in West Africa

living also in moist, forested areas where it seems to be

rare. Only four other localities have been reported from

Guinée forestiere, viz. Mt. Nimba (Angel et al. 1954 a),

Mt. Béro (Rédel & Bangoura 2006) and two sites in the

western Guinean region of Boké (Hillers et al. 2006). Al-

so Klaptocz (1913) got only one specimen in the moist

savanna near Dabola (140 km E of Mamou) and called it

rare and even unknown to the locals. In contrast, Dr. Guy

Kremer (pers. comm., see ZFMK vouchers mentioned

above) found it, along with the rarer C. senegalensis, to

be more common near Kindia. A further specimen, also

from a savanna habitat in PN Haut Niger has also been

mentioned above. In the savanna areas of the Simandou

range C. senegalensis was the only chameleon species

recorded so far (R6del & Bangoura 2004b; CB further un-

publ. data).

Gekkonidae

Cnemaspis occidentalis Angel, 1943

Material examined: ZMB 75507, Mt. Nimba, 1274 m

a.s.l., L. Sandberger coll. 5 August 2008.

Bonn zoological Bulletin 60 (1): 35-61

Fig. 11. Hemidactylus fasciatus, juvenile from Mt. Nimba (Pho-

to: C. Brede).

Remarks: Recorded from the Mt. Nimba area already by

Ineich (2003).

Hemidactylus angulatus Hallowell, 1854

Material examined: ZFMK 56368, 56140—141, 62172,

Sérédou; ZFMK 56129-131, N’Zérékoré; ZFMK 60766,

forest near Sérédou, W. Bohme coll. 6—26 Oct. 1993;

ZFMK 82162: Diécké Forest (07E35’46.9”N,

O8ES2’18.8” W), 454 m asl., degraded forest, M.-O. Rédel

& M.A Bangoura coll. 24 Nov. 2003.

Remarks: H. angulatus proved to be common in both for-

est areas visited. It was found on house walls and also

within houses and lived in partial syntopy with H.

mabouia, but seemed to be more numerous than the lat-

ter. The specimens from Diécké Forest were included in

the table by Rédel & Bangoura (2006) as H. brooki of

which H. angulatus figured as a subspecies before. Un-

der the same name the species has been likewise listed

from western Guinea (Hillers et al. 2006).

Hemidactylus fasciatus Gray, 1842

Material examined: ZFMK 82161, Diécké Forest (see

above), 454 ma.s.]., degraded forest, M.-O. Rédel & M.H.

Bangoura coll. 24 Nov. 2003.

Remarks: This record was already mentioned in the table

provided by Rédel & Bangoura (2006). A further speci-

©ZFMK

Amphibians and reptiles from Guinea 4]

men, likewise from Diécké Forest, was found by WB pre-

served in the “Centre Entomologique” of Herold Olsen in

N’Zerékoré. MOR observed the species to be not uncom-

mon within the forests of the Simandou range and CB took

pictures of a juvenile on Mt. Nimba (Fig. 11).

Hemidactylus mabouia (Moreau de Jonnés, 1818)

Material examined: ZFMK 72303, Mt. Nimba, Gbakoro,

N’Zo and Zougueé, 500 m a.s.l., native collector 26—30

Aug. 1999.

Remarks: Not recorded from Mt. Nimba area by Ineich

(2003). As already stated above, H. mabouia lives in syn-

topy with H. angulatus on the same housewalls in

N’Zérékoré, but in lesser numbers. R6del & Bangoura

(2006) found it in Diécké Forest. In Sérédou, where H.

angulatus 1s common in human habitations, H. mabouia

seemed to be completely absent.

Hemidactylus muriceus Peters, 1870

Material examined: ZFMK 58617, Diécké Forest, W.

Bohme coll. 15 Oct. 1993; ZFMK 82171, Mt. Béro

(8E8°20.7”"N, 8E34’23.7W), river and gallery forst,

M.A. Bangoura & M.-O. Rédel coll. 1 Dec. 2003; ZFMK

82163, Diécké Forest (7E35’°46.9”N, 8E52°18.8" W), 454

m asl., degraded forest, M.A. Bangoura & M.-O. Rédel

coll. 21-23 Nov. 2003; ZFMK 82164, Diéckeé Forest,

(7E35’43.6N, 8E51°52.3”W), creek in good forest, M.A.

Bangoura & M.-O. Rédel, 27 Nov. 2003.

Remarks: There has been much confusion as to the iden-

tity of H. muriceus, H. echinus and H. pseudomuriceus;

ZFMK 58617 was the first unambiguous specimen of H.

pseudomuriceus from Guinea (Henle & Bohme 2003), fol-

lowed now by the two additional voucher specimens from

Mt. Béro Classified Forest (Rédel & Bangoura 2006). Fur-

ther specimens of this forest gecko have been observed

in the lowland forests of Mt. Nimba (MOR), and the Pic

de Fon/Simandou range (Rédel & Bangoura 2004b).

Scleroglossa

Lacertidae

Holaspis guentheri Gray, 1863

Material examined: ZFMK 60563, near Sérédou, W. Buit-

zler coll. Jan./Apr. 1995.

Remarks: One further specimen was observed by WB on

a big fallen tree trunk near Malweta but could not be col-

Bonn zoological Bulletin 60 (1): 35-61

lected, despite the unusually low habitat structure of this

otherwise strictly canopy-dwelling lizard. A specimen

from Diécké Forest collected by H. Olsen (ca. 1950) is

kept in ZMUC. For Guinea, this species was before only

recorded from the Mt. Nimba region (Angel et al. 1954;

see also Ineich 2003).

Scincidae

Cophoscincopus durus (Cope, 1862)

Material examined: ZFMK 82165, Diécké Forest, 454 m

asl., 7E35°46.9”N, 8E52’18.8” W, degraded forest, M.A.

Bangoura & M.-O. Rédel coll. 21.—23 Nov. 2003; ZFMK

82166, Diécké Forest, 7E35’43.6’N, 8E51752.3”’, creek

in good forest, M.A. Bangoura & M.-O. Rédel coll. 27

Nov. 2003.

Remarks: For distribution in West Africa and sympatry

with the two congeners listed below see Bohme et al.

(2000: map) and Ineich (2003).

Cophoscincopus greeri Bbhme, Schmitz & Ziegler, 2000

Material examined: ZFMK 57599, male (holotype): Mt.

Nimba, 1800 m asl., W. Bohme coll. Oct. 1993; ZMB

75500-7501, Mt. Nimba, A. Hillers coll. May 2006.

Remarks: The male holotype was earlier tentatively as-

signed to C. simulans by BOhme (1994 c). The C. simu-

lans female figured in the same paper (ZFMK 57843),

however, proved actually to belong to this revalidated

species, so that the photographs showing ZFMK 57599

during alternate copulations with this female document a

voluntary interspecific pairing.

Cophoscincopus simulans (Vaillant, 1884)

Material examined: ZFMK 56148, 57843, females, Mal-

weta south of Sérédou, W. Béhme coll. 2—26 Oct 1993;

ZMK S2G7e) eDiéck ywRorest,.) 7E3SS743°67.N,

8E51°52.3”W, creek in primary forest, M.A. Bangoura &

M.-O. Rédel coll. 27 Nov. 2003; ZFMK 82168—170, Mt.

Bero, 8E8’20.7”N, 8E34’23.7”W, river and gallery for-

est, M.A. Bangoura & M.-O. Rédel coll. Dec. 2003;

ZFMK 82178—180, Déré Forest, 444 m asl., 7E36’°13.2”N,

8E!12°42.3”W, M.A. Bangoura & M.-O. Rédel coll. 17

Dec. 2003; ZMB 75502-77505, Mt. Nimba, A. Hillers

coll. May 2006.

©ZFMK

42 Wolfgang Bohme et al.

Remarks: For the West African distribution and sympa-

try with the two above congeners see BOhme et al. (2000:

map) and Ineich (2003).

Panaspis nimbensis (Angel, 1944)

Material examined: ZFMK 56147, Sérédou, on PROGER-

FOR compound under leaf litter. W. Béhme coll. Oct.

1993.

Remarks: This little skink was described from Nimba Mts.

(Angel 1944, Angel et al. 1954 a), and Ineich (2003) sum-

marized the Guinean voucher material of MNHN which

is entirely from the Mt. Nimba area. After 40 years, our

Ziama specimen is the first from outside of Nimba Mts.

(see BOhme 1994 c). P. nimbensis is otherwise known

from Ivory Coast (Perret 1973).

Trachylepis affinis (Gray, 1838)

Material examined: ZFMK 56143—-145, Sérédou, W.

Bohme coll. 2.—26. Oct 1993; ZFMK 58611—612, Diécké

Forest, W. Bohme coll. 2—26 Oct. 1993.

Remarks: A common species (see also R6del & Bangoura

2006) with a wide variety of habitats occupied, from pri-

mary forest where it lives on the ground as well as on high

tree trunks (see Figs. in BOhme 1994 a, c) to secondary

forests, bushland and human habitations (housewalls). In

primary forests the species is usually restricted to more

open sites such as tree fall gaps (MOR, unpubl. data).

Trachylepis maculilabris (Gray,1845)

Material examined: ZFMK 56146, Sérédou, W. BOhme

coll. 2—26 Oct. 1993.

Remarks: Proved to be much rarer in SE Guinea (in con-

trast to e.g. Cameroon: see Herrmann et al. 2005) than 7.

affinis, and the voucher specimen was the only specimen

seen. In comparison to 7? affinis, this species is more close-

ly connected to forest.

Amphisbaenidae

Cynisca cf. liberiensis (Boulenger, 1878)

Material examined: ZFMK 60564, Ziama Forest near

Sérédou. W. Biitzler coll. Jan./Apr. 1995.

Bonn zoological Bulletin 60 (1): 35-61

Remarks: Found in the stomach of a roadkilled Polemon

acanthias (ZFMK 60567).

The prey as well as its predator have been badly damaged

by the vehicle that killed the snake. The head of the am-

phisbaenian is additonally damaged by beginning diges-

tion within the snake, one mandible being already freely

macerated, without teeth. We assign this specimen with

some reservation to Cynisca liberiensis because this

species is known from two Mt. Nimba localities: Angel

et al. (1954 a) reported it from Pierré-Richaud and from

Kéoulenta as C. /amottei (Angel, 1943), a name which has

been synonymized by Gans (1987) with C. liberiensis and

has been considered to be subspecifically valid by Ineich

(2003). The diagnostic characters given by Gans (1987)

can mostly not be verified any more, however, two char-

acteristics still visible in our specimen (“‘small, relative-

ly slender” and “some specimens showing mottling of the

dorsal segments”’) do at least not argue against our tenta-

tive assignment. Recovery of intact specimens from Zia-

ma Forest has to be awaited for.

Varanidae

Varanus ornatus (Daudin, 1803)

Material examined: ZFMK 56028, Sérédou (Mt. d’An-

tenne), native collector coll. 6-26 Oct. 1993, ZFMK

56029—030 (juvs.), Sérédou (agricultural area), native col-

lectors coll. 18 Oct. 1993.

Remarks: The adult male ZFMK 56028 had been caught

in a poacher’s noose in primary forest on the “Antenna

Hill”, Sérédou. The juveniles were brought by Sérédou lo-

cals to the PROGERFOR compound. Further juveniles

have been observed by WB nearby at Malweta river, and

a juvenile specimen collected in Diécké Forest was also

found in the “H. Olsen Entomological Center” in

N’Zérékoré. All specimens seen showed the dorsal pat-

tern of only five oblique rows of light ocelli, and the

voucher specimens checked have a whitish to flesh-

coloured tongue, both characters being diagnostic for /.

ornatus which meets its parapatric sister species V. niloti-

cus in Guinea (B6hme & Ziegler 1997, see also Green-

baum & Carr 2005). Whereas the latter has already been

termed common by Chabanaud (1921) and is also repre-

sented in our Haut Niger National Park material (see

above, and Greenbaum & Carr 2005), the ornatus speci-

mens are the first representatives of the forest species V.

ornatus documented for the country.

©ZFMK

Amphibians and reptiles from Guinea 43

Fig. 12. Zyphlops liberiensis, Diani River, near Nzérékoré (Pho-

to: W. B6hme).

Typhlopidae

Typhlops liberiensis (Hallowell, 1848)

Material examined: ZFMK 56090, adult female. Sérédou,

W. Bohme coll. 12 Oct. 1993; ZFMK 56135, Diani Riv-

er, W. BOhme coll. 13 Oct. 1993.

Remarks: The taxonomic status of this irregularly speck-

led blind snake (Fig. 12) is not yet stabile. Sometimes it

is also regarded to be a subspecies of 7. punctatus, or even

of T. congestus (see Ineich 2003).

Typhlops punctatus Leach, 1819

Material examined: ZFMK 58620, Sérédou, W. Biitzler

coll. Aug./Nov. 1994.

Remarks: Another specimen of this taxon was seen in the

H. Olsen collection in N’Zérékoré.

Boidae

Calabaria reinhardtii (Schlegel, 1848)

Material examined: ZFMK 56042, north of Sérédou (on

road to Macenta) through primary forest, W. Butzler coll.

POC; 1993.

Remarks: A second individual was seen in the “Centre En-

tomologique” in N’Zérékoré (H. Olsen coll.). According

Bonn zoological Bulletin 60 (1): 35-61

Fig. 13. Python sebae, subadult specimen from Mt. Nimba

(Photo: C. Brede).

to Ineich (2003), a third specimen was collected at Ziéla,

Mt. Nimba; it is not kept in MNHN but “dans les collec-

tions guinéennes du mont Nimba (Lamotte, comm. pers.

09/97)”. Thus, our specimen from Sérédou, though bad-

ly damaged as a roadkill, is the only Guinean specimen

available for study.

Pythonidae

Python sebae (Gmelin, 1789)

Material examined: Photographic voucher.

Remarks: In the Mt. Nimba area, a subadult rock python

could be photographed by CB (Fig. 13).

Lamprophiidae

Bothrophthalmus lineatus Peters, 1863

Material examined: ZFMK 56094, south of Sérédou, Mal-

weta village, W. Bohme coll. 6-26 Oct. 1993; ZFMK

58615, Diécké Forest, H. Vierhaus coll. Apr. 1994; ZFMK

82159, Diécké Forest (7.35 N; 8.51 W), M.A. Bangoura

& M.-O. Rédel coll. 27 Nov. 2003 (Fig. 14).

Remarks: Some authors, including Angel et al. (1954 b)

and Ineich (2003), refer to this snake as B. lineatus lin-

eatus, which implies conspecifity of the central African

©ZFMK

44 Wolfgang Bohme et al.

Fig. 14. Bothrophthalmus lineatus, ZFMK 82159, from Diécké

Forest (Photo: C. Brede).

taxon brunneus. However, as east of the distribution range

of the latter (eastern DRC, Ruanda) again typical /inea-

tus are found, we prefer to regard both distinct forms as

separate species.

ZFMK 56094 contained remains of an unidentified small

rodent.

Lamprophis lineatus (Duméril, Bibron & Duméril, 1854)

Material examined: ZFMK 56125 and 56126, Ziama For-

est south of Sérédou: Malweta village, W. Biitzler coll.

1991/92 and W. Bohme 6-26 Oct. 1993, respectively.

Lamprophis olivaceus (Duméril, 1856)

Material examined: 56121—122, Ziama Forest north of

Sérédou: Balassou village, W. Bohme coll. 6—26 Oct.

1993: 58610, Ziama Forest west of Sérédou: Soundédou

village, W. Bitzler leg. Apr. 1994.

Fig. 15. Lycophidion sp., ZMB 75508 from Mt. Tétini, Déré

Forest (Photo: C. Brede).

Bonn zoological Bulletin 60 (1): 35-61

Lamprophis virgatus (Hallowell, 1854)

Material examined: ZFMK 56123, Ziama Forest south of

Sérédou: Malweta village, W. Bohme coll. 6-26 Oct.

1993; ZFMK 56264, same locality, W. Biitzler coll. Nov.

1993; ZFMK 57809, Sérédou, W. B6hme coll. 6—26 Oct.

1993; ZFMK 60556 and 60770, same locality, W. Biitzler

coll. Jan./Apr. and June/Aug. 1994 respectively.

Remarks: ZFMK 57809 was recovered from the stomach

of a Mehelya guirali (ZFMK 56041).

Lycophidion sp.

Material examined: ZMB 75508, Mont Tétini, N

08°20.348’, W 08°22.741’, gallery forest, small river,

rapids, some swampy areas, C. Brede, M.A. Bangoura, J.

Doumbia coll. 21 Sep. 2008.

On Mt. Tétini we collected a snake (Fig. 15) where pholi-

dosis was closest to L. irroratum. However, we recorded

distinct differences in scalation and colour pattern, com-

pared to Chippaux (2001) or L. irroratum specimens

known to us from other West African localities, e.g. Co-

moé National Park, Ivory Coast (SMNS 8469.1—2, Rédel

et al. 1995, 1999) or Pendjari National Park, Benin (MOR,

unpubl. data). The taxonomic status of this snake needs

further investigation.

Mehelya guirali (Mocquard, 1887)

Material examined: ZFMK 56041, Ziama Forest near

Sérédou, road in primary forest, W. BOhme coll. 6—26 Oct.

1993; ZFMK 58627, Ziama Forest near Sérédou, W. But-

zler coll. Dec. 1994.

Remarks: ZFMK 56041 had a Lamprophis virgatus

(ZFMK 57809) in its stomach.

The above two specimens (ID verified by B. Hughes) were

mentioned by B6hme (2000) as the first country record

of this species. Five more specimens from the Guinean and

Liberian part of Mt. Nimba are kept in MNHN (Ineich

2003).

Mehelya poensis (Smith, 1847)

Material examined: ZFMK 56111, south of Sérédou: Mal-

weta village, W. B6hme coll. 6-26 Oct 1993; ZFMK

OZFMK

Amphibians and reptiles from Guinea 45

Fig. 16. Dasypeltis scabra, ZFMK 75496, from Mt. Nimba

(Photo: M.-O. Rédel).

60569, Ziama Forest near Sérédou, W. Butzler coll.

Jan./Apr. 1995.

Remarks: ZFMK 56111 (ID verified by B. Hughes) con-

tained unidentifiable remains of a snake.

Mehelya stenophthalmus (Mocquard, 1887)

Material examined: ZFMK 56112, south of Sérédou: Mal-

weta village, W. Bohme coll. 6—26 Oct. 1993.

Colubridae

Crotaphopeltis hotamboeia (Laurenti, 1768)

Material examined: ZFMK 54896, Sérédou, H. Vierhaus

coll. 4 Aug. 1992; ZFMK 56116—120, Ziama Forest south

of Sérédou: Malweta village, W. Bohme coll. 6—26 Oct.

1993; ZFMK 60568, Ziama Forest near Sérédou, W. Biit-

zler coll. Jan./Apr. 1995; ZFMK 62173-175, Sérédou, W.

Butzler coll. Nov./Dec. 1995.

Remarks: With ten specimens from Ziama Forest the sec-

ond most common snake of our survey. Three specimens

had food items in their intestinal tracts: ZFMK 54896 anu-

ran and insect remains, ZFMK 56116 one Amietophrynus

maculatus, and ZFMK 60568 unidentifiable remains of a

mouse.

Bonn zoological Bulletin 60 (1): 35—61

Dasypeltis fasciata Smith, 1849

Material examined: ZFMK 56048, Ziama Forest south of

Sérédou: Malweta village, W. Bohme coll. 2—26 Oct.

1993.

Remarks: First reported for Guinea by BOhme (2000). In-

eich (2003) recorded three MNHN specimens from the Mt.

Nimba area. One was catalogued in 1943 and reidentified

by C. Gans, the two others in 1962. The identification of

our specimen, the only Guinean one known from outside

of Mt. Nimba, was verified by B. Hughes. Recently, Trape

& Mané (2006 a) reported and figured a specimen from

Ziama Forest.

Dasypeltis scabra (Linnaeus, 1758)

Material examined: ZMB 75496, Mt. Nimba, Grand

Rocher, 1600 m a.s.1., M.-O. Rédel & L. Sandberger coll.

1 Jul. 2007.

Remarks: Reported from Mt. Nimba by Ineich (2003). The

colour pattern of our voucher specimen (Fig. 16) is very

similar to the specimen figured by Trape & Mané (2006

a, Fig. 5c) as D. cf. scabra.

Dipsadoboa brevirostris (Sternfeld, 1908)

Material examined: ZFMK 56096, Ziama Forest south of

Sérédou: Malweta village, W. Béhme coll. 6—26 Oct.

1993; ZFMK 82173, Mont Béro, 8E08’23.7 N, 8E34’23.7

W), M.A. Bangoura & M.-O. Rodel coll. 30 Nov. 2003.

Remarks: ZFMK 56096 is a gravid female with three eggs

inside. The specimens fit the description of Leptodira

guineensis Chabanaud, 1920, described from the neigh-

bouring Diécké Forest. This taxon was synonymized with

brevirostris Sternfeld by Rasmussen (1989, see also Ras-

mussen 1994). Not listed for Mt. Nimba (Angel et al. 1954

b, Ineich (2003).

Dipsadoboa unicolor Giinther, 1858

Material examined: ZFMK 56095 & 56267, Ziama For-

est south of Sérédou: Malweta village, W. Bohme coll.

6-26 Oct. 1993; ZFMK 56267, same locality, W. Biitzler

coll. Nov. 1993; ZFMK 82160, Diécké Forest (7.35 N;

8.51 W), M.A. Bangoura & M.-O. Rédel coll. 27 Nov.

2003, photographic voucher from Mt. Béro (Fig. 17).

©ZFMK

46 Wolfgang Bohme et al.

Fig. 17. Dipsadoboa unicolor from Mt. Béro (Photo: M.-O.

Rédel).

Remarks: ZFMK 56095 and 56267, two males, prove their

identity as true D. unicolor by having 70 and 75 undivid-

ed subcaudals respectively (Rasmussen 1993, 1994). They

are strikingly differently coloured, ZFMK 56095 being

dark bluish-green, and ZFMK 56267 light green (mean-

while light bluish in preservative).

Hapsidophrys lineata Fischer, 1856

Material examined: ZFMK 56089, between Macenta and

Sérédou, W. Biitzler coll. 25 Oct. 1993.

Remarks: The adult, roadkilled specimen is headless.

Hapsidophrys smaragdina (Schlegel, 1837)

Material exmined: ZFMK 56043—046, 56149, Ziama For-

est south of Sérédou: Malweta village, W. Bohme coll.

6-26 Oct 1993; ZFMK 60769, Sérédou, W. Biitzler coll.

June/Aug. 1995.

Remarks: Collected by R. Pujol at Sérédou before (Con-

damin 1959), but — remarkably — not listed for the Mt.

Nimba area (Ineich 2003).

Meizodon coronatus (Schlegel, 1837)

Material examined: ZFMK 56133, N’Zérékore, H. Olsen

coll. ca. 1950.

Remarks: This specimen from N’Zérékoré was donated

to ZFMK by the women responsible for the Olsen collec-

tion in 1993. For the taxonomic and nomenclatural con-

Bonn zoological Bulletin 60 (1): 35-61

fusion between this and the following species see Roux-

Esteve (1969) and below.

Meizodon regularis Fischer, 1856

Material examined: ZFMK 56113—-115, 56261, south of

Sérédou: Malweta village, ZFMK 56260, Sérédou, W.

Bohme coll. 6—26 Oct. 1993.

Remarks: Roux-Estéve (1969) revised the West African

portion of the genus and reached the conclusion that .

coronatus and M. regularis are two distinct species with

broadly overlappping ranges, a view corroborated by

Schatti (1985). She listed also several Guinean localities

for this species she had found 1n various collections in the

course of this work. These localities are spread all over

Guinea. Bohme (2000) and Ineich (2003), not citing Roux-

Estéve (1969) referred to Angel et al. (1954 b) who had

listed all his specimens still under MM. coronatus. There-

fore, Bohme (2000) erroneously believed his specimens

to be new for Guinea, while Ineich (2003) could recheck

the MNHN material and was able to quote M. coronatus

sensu Angel (1954 b) as a partial chresonym of M. regu-

laris.

Philothamnus carinatus (Andersson, 1901)

Material examined: ZFMK 56265, Ziama Forest south of

Sérédou: Malweta village, W. Bohme coll. 6—26 Oct.

1993; ZFMK 57230, Ziama Forest: Gboda village, W. Biit-

zler coll. Feb. 1994.

Remarks: identification verified by B. Hughes, July 1994.

Philothamnus heterodermus (Hallowell, 1857)

Material examined: ZFMK 56138—139, Ziama Forest

south of Sérédou: Malweta village, W. Bohme coll. 8-26

Oct. 1993; ZFMK 58623, Sérédou, W. Biitzler coll.

Aug./Nov. 1994; ZFMK 82172, Mt. Béro (8.08 N; 8.34

W), M.A. Bangoura & M.-O. Rédel coll. 30 Nov. 2003.

Philothamnus irregularis (Leach, 1819)

Material examined: ZFMK 82177, Mt. Nimba area: Mt.

To (7.39 N; 8.29 W), M.A. Bangoura & M.-O. Rédel coll.

21 Nov. 2003.

©ZFMK

Amphibians and reptiles from Guinea 47

Thelotornis kirtlandii (Hallowell, 1844)

Material examined: ZFMK 60767, Ziama Forest near

Sérédou, W. Butzler coll. June/Aug. 1995.

Remarks: Recorded by Villiers (1950, 1954) and Angel

et al. (1954 b) from Mt. Nimba (Ineich 2003). ZFMK

60767 seems to be the first Guinean specimen found out-

side the Mt. Nimba range and has been mentioned and fig-

ured already by Riquier & B6hme (1996). The species was

also found by Rédel & Bangoura (2006) in Diéckeé For-

est.

Thrasops occidentalis Parker, 1940

Material examined: ZFMK 56033, Sérédou, road in pri-

mary forest, W. BOhme coll. 6—26 Oct. 1993.

Remarks: This specimen had a largely digested and thus

unidentifiable anuran in its stomach. Moreover, it was

mentioned as a first country record by Bohme (2000) who

overlooked, however, the record from Dalaba by Villiers

(1950). From 1986-1990 the species was also found in the

Mt. Nimba region of Guinea and Liberia (Ineich 2003).

Toxicodryas blandingii (Hallowell, 1844)

Material examined: ZFMK 54889, Sérédou (in house), H.

Vierhaus coll. 4 Aug. 1992; ZFMK 5603 1—032, Sérédou,

agricultural area, W. Bohme coll. 6—26 Oct. 1993; ZFMK

56040, Sérédou, road in primary forest, W. BoOhme coll.

6-26 Oct. 1993; 56277-278, Ziama Forest south of Séré-

dou: Malweta village, W. Biitzler coll. Nov. 1993.

Remarks: ZFMK 56032 had two weaverbirds in its stom-

ach. The species is very common at and around the for-

mer Nimba research station (1000 m asl.).

Toxicodryas pulverulentus (Fischer, 1856)

Material examined: ZFMK 56039, Sérédou, W. Bohme

coll. 6—26 Oct. 1993; ZFMK 58625—626, same locality,

W. Butzler coll. Aug./Nov. 1994; 60768 and 62178, same

locality, W. Butzler coll. June/Aug. 1995 and Aug./Nov.

1995 respectively.

Remarks: Two specimens had food items in their stom-

ach: ZFMK 56039 several mice, and ZFMK 58265 a

lizard (Agama agama complex).

Bonn zoological Bulletin 60 (1): 35-61

Natricidae

Afronatrix anoscopus (Cope, 1861)

Material examined: ZFMK 56049-079, Ziama Forest

south of Sérédou, Malweta village, W. B6hme coll. 6—26

Oct. 1993; ZFMK 56256, Sérédou (Mt. d’antenne), ZFMK

56268—276 Malweta village, W. Biitzler coll. Nov. 1993;

ZFMK 58609, Ziama Forest west of Sérédou: Soundédou,

W. Bitzler coll. Apr. 1994; ZFMK 82158: Diécké Forest

(7.31 N; 8.50 W), M.A. Bangoura & M.-O. Rédel coll.

27 Nov. 2003.

Remarks: A. anoscopus is one of the most common snakes

in SE Guinea. We recorded them in small streams of al-

most all forests, i.e. in the Pic de Fon area, Diécké, and

on Mt. Nimba. The most common colour morph is uni-

form brown (Fig. 18) to brilliant reddish, fewer individ-

uals have yellowish brown bodies with black occular spots

(Fig. 19). This species proved to be eudominant in the

snake community of Ziama Forest: 42 specimens were col-

lected, followed by the two next-common species (Cro-

taphopeltis hotamboeia and Natriciteres variegata) with

just ten and nine individuals respectively (see BOhme

2000). Because of this sample size of the Ziama popula-

tion, some morphological and nutritional data gathered

from this series seem to be useful:

— Size: snout-vent length 14.6.—45.5 cm, mean 35.54; tail

length 4.6—-15.9 cm, mean 11.6 cm. Largest specimen is

a female (ZFMK 56074): 60.4 cm.

— Scalation: Dorsals around midbody 23 in all but two

(ZFMK 56063, 56079, which both have 25) specimens;

ventrals 139-151; subcaudals 64—82; anal divided in 36

specimens, entire in 4. Lateral keeling of subcaudals dis-

tinct in 36 specimens, indistinct in 4, and lacking in 2;

supralabials 9 on both sides of head in 33 specimens, 8—9

(left/right side of head) in 2, and 9-10 in 3.

— Colour pattern: 32 specimens have a marked pattern of

black spots arranged in longitudinal rows on a lighter

ground colouration; in two specimens, the spots are indis-

tinct, and in eight instances, they are lacking, the respec-

tive individuals being unicoloured.

— Taxonomy: Angel (1932) synonymized Helicops gen-

drii Boulenger, 1910: type locality “Labé, French Guinea”

with Tropidonotus anoscopus Cope, 1861 (type locality

Liberia, but Villiers (1950) used the name Natrix a. gen-

drii (Boulenger, 1910 for the Guinean populations to dis-

tinguish them from the nominotypic N. a. anoscopus

(Cope, 1861: type locality Liberia). The few morphome-

tric data published so far from West Africa show that the

variability of this widely distributed snake only within

©ZFMK

48 Wolfgang Bohme et al.

Fig. 18. Afronatrix anoscopus, unicoloured specimen from Mt.

Nimba (Photo: C. Brede).

Guinea exceeds that of our series in most of the data tak-

en. A modern revision including also the central African

populations seems highly desirable.

— Nutrition: Of the 42 specimens from Ziama Forest, on-

ly three individuals had full stomachs (with fishes), viz.

ZFMK 56066 (Aplocheilichthys schioetzi), 56076 (Krib-

ia cf. nana), and 56275 (Aplocheilichthys schioetzi and

unidentifiable frog remains).

Grayia smithii (Leach, 1818)

Material examined: ZFMK 56036, adult female, south of

Sérédou: Malweta village, W. Bohme coll. 6-26 Oct.

1993; photographic voucher: a big, light-coloured road-

killed specimen from the road between Sérédou and

Zoboroma, W. Biutzler phot. March 1995.

Remarks: This specimen had several fishes (Tilapia

(Coptodon) sp.n.?) in its stomach. The bad shape of the

fishes from the stomach of this snake does not allow a pre-

cise identification, but it seems likely that they belong to

an undescribed species (J. Freyhof, pers. comm.).

Another big female of this species, likewise from Malwe-

ta village, was left for the PROGERFOR teaching collec-

tion at Sérédou.

Bonn zoological Bulletin 60 (1): 35-61

Fig. 19. Afronatrix anoscopus, patterned specimen from Pic de

Fon (Photo: C. Brede).

Natriciteres variegata (Peters, 1861)

Material examined: ZFMK 56097—099, Ziama Forest

south of Sérédou: Malweta village, W. Bohme coll. 6—26

Oct. 1993; ZFMK 56257—258, Sérédou, W. Biitzler coll.

Nov. 1993; ZFMK 56262—263, Malweta village, W. Biit-

zler coll. Nov. 1993; ZFMK 58622 and 60579, Sérédou,

W. Bitzler coll. Aug./Nov. 1994 and Jan./Apr. 1995 re-

spectively.

Remarks: With nine specimens the third most common

snake species in WB’s Ziama Forest survey.

Psammophiidae

Psammophis phillipsi (Hallowell, 1844)

Material examined: ZFMK 56047, Ziama Forest south of

Sérédou: Malweta village, W. Bohme coll. 6—26 Oct.

1993; ZFMK 60792, Sérédou, W. Butzler coll. June/Aug.

1995.

Remarks: Both specimens have the ventrals shaded with

grey, thus representing the true, forest-dwelling P. philipp-

si. Moreover, the smaller specimen is dorsally unicoloured

which is also characteristic. P. cf. phillipsi, often referred

to as P. sibilans in West Africa, has a striped juvenile dress

which fades in adults of this savanicolous species

(Bohme et al. 1996).

©ZFMK

Amphibians and reptiles from Guinea 49

Atractaspididae

Aparallactus lineatus (Peters, 1870)

Material examined: ZFMK 56100, Ziama Forest south of

Sérédou: Malweta village, W. Bohme coll. 6—26 Oct.

1993.

Remarks: The validity of this species and its specific dis-

tinctness from A. niger was proven by Wallach (1994),

lastly based on the sympatric occurence of both species

“from the same rainforest locality” near Mt. Nimba. Here,

again both species are recorded from the same forest lo-

cality (Malweta village, see below).

Aparallactus modestus (Gunther, 1859)

Material excamined: ZFMK 56105, Ziama Forest south

of Sérédou: Malweta village, W. Bohme coll. 6—26 Oct.

1993; ZFMK 82157, Diécké Forest (N 7.35; W 8.12),

M.A. Bangoura & M.-O. Rédel coll. 21/23 Nov. 2003

(Fig. 20).

Remarks: At Malweta village, next to A. lineatus and A.

niger, the sympatric occurrence of the third congener is

proven.

Fig. 20. Aparallactus modestus, ZFMK 82157, from Diécké

Forest (Photo: M.-O. Rédel).

Aparallactus niger Boulenger, 1897

Material examined: ZFMK 56101—102, Ziamam Forest

south of Sérédou: Malweta village, W. Bohme coll. 6—26

Oct 1993; ZFMK 56266, same locality, W. Bitzler coll.

Bonn zoological Bulletin 60 (1): 35-61

Nov. 1993; ZFMK 60565, Sérédou, W. Biitzler coll.

Jan./Apr. 1995.

Remarks: See above under A. /ineatus.

Atractaspis irregularis (Reinhardt, 1843)

Material examined: ZFMK 56106—108, Ziama Forest

south of Sérédou: Malweta village, W. Bohme coll. 6—26

Oct. 1993; ZFMK 58624, Sérédou, W. Biitzler coll.

Aug./Nov. 1994.

Remarks: Ineich (2003) listed several specimens from the

IFAN and MNHN collections from localities in the Mt.

Nimba range, but did not include A. aterrima in his pa-

per, although Rasmussen (2005) found several vouchers

of this last named species in both collections.

Polemon acanthias (Reinhardt, 1860)

Material examined: ZFMK 56103-—14, Ziama Forest south

of Sérédou, W. Béhme coll. 6-26 Oct. 1993; ZFMK

56259, Sérédou (“Mt. d’Antenne’”’), W. Bitzler coll. Nov.

1993; ZFMK 60567, Sérédou, W. Bitzler coll. Jan./Apr.

1995; ZFMK 62176, same locality, W. Butzler coll.

Nov./Dec. 1995.

Remarks: The road-killed specimen ZFMK 60567 con-

tained a Cynisca cf. liberiensis (see above).

Elapidae

Dendroaspis viridis (Hallowell, 1844)

Material examined: ZFMK 54890-891, Sérédou, H. Vier-

haus coll. 4 Aug. 1992; ZFMK 56034—035, Sérédou, W.

Bohme coll. 2—26 Oct. 1993.

Remarks: D. viridis proved to be quite common in Séré-

dou and was regularly observed in the compound of

PROGERFOR.

Naja melanoleuca Hallowell, 1857)

Material examined: ZFMK 56091—092, juvs., Ziama For-

est south of Sérédou: Malweta village; ZFMK 62177, Zia-

ma Forest near Sérédou, W. Bitzler coll. Nov./Dec. 1995.

©ZFMK

50 Wolfgang Bohme et al.

Remarks: Recorded by Rédel & Bangoura (2006) also

from Diécké Forest.

Naja nigricollis Reinhardt, 1843

Material examined: ZFMK 56037: Sérédou. W. Biitzler

coll. 1991/92.

Remarks: Rédel & Bangoura (2006) listed this species al-

so from Mt. Béro.

Pseudohaje nigra Gunther, 1858

Material examined: ZFMK 56134, N’Zérékoré, ex coll.

H. Olsen, received Oct. 1993.

Remarks: Ineich (2003), based on David & Ineich

(1999), claimed that the presence of this species in Guinea

was still uncertain although it was likely to occur, due to

a MNHN specimen from the Liberian part of Mt. Nimba

(Ineich 2003). However, both Roman (1976: “On posséde

en collection un exemplaire de N’Zérékoré, Guinée”’) and

Bohme (2000: based on ZFMK 56134) had reported P. ni-

gra already from Guinea.

Viperidae

Atheris chlorechis (Pel, 1851)

Material examined: ZFMK 56109-110, Ziama Forest

south of Sérédou: Malweta village, W. Bohme coll. 2—26

Oct. 1993; ZFMK 82174, Mt. Béro, M.A. Bangoura & M.-

O. Rédel coll. 5 Dec. 2003.

Remarks: Known in Guinea from the Mt. Nimba area (Vil-

liers 1950, Angel et al. 1954 b, Ineich 2003), but also al-

ready reported from Sérédou (Condamin 1959). A Mt.

Nimba specimen 1s depicted in Fig. 21.

Bitis arietans Merrem, 1820

Material examined: Photographic voucher by MOR from

Pic de Fon, 1600 m.

Remarks: Apart from the altitude, this photographic record

(Fig. 22) is remarkable because it shows a dorsal pattern

where the light chevron marks typical for this species van-

ish already after the first half of the body, passing grad-

Bonn zoological Bulletin 60 (1): 35-61

Fig. 21. Atheris chlorechis from Mt. Nimba (Photo: C. Brede).

ually into subquadrangular light spots. The same pattern

type is seen in the photographic voucher from Kissidougou

mentioned above and seems to be commoner in West

Africa than a chevron-mark pattern along the entire body

as it is typical for eastern and southern African popula-

tions.

Fig. 22. Bitis arietans, specimen from Pic de Fon, 1600 m, note

the restriction of chevron pattern to the anterior half of body

(Photo: M.-O. Rédel).

Bitis nasicornis (Shaw, 1802)

Material examined: ZFMK 56038, Sérédou, W. Biitzler

coll. 1991/92. A further specimen was photographed in

Ziama by CB in 2010.

Bitis rhinoceros (Schlegel, 1855)

Material examined: ZFMK 56126, between Sérédou and

Macenta, W. BOhme & W. Biitzler coll. 23. Oct. 1993.

OZFMK

Amphibians and reptiles from Guinea 5]

Remarks: Only the skin with head and tail inside of this Crocodylia

big male (total length 105 cm) could be preserved in al- Crocodylidae

cohol. A photographic record was made in the Fouta Djal-

lon by CB (Labé/Daralabe/Forét Kokoulo), an extreme — Represented in West Africa by three species, for which the

western locality for this species. slender-snouted species Mecistops cataphractus was re-

ported to occur in the Baffing (= Bafing) river, upper Sene-

gal river) by Klaptocz (1913). The Western Nile Croco-

dile (Crocodylus suchus) (see Schmitz et al. 2003) is sa-

Causus maculatus (Hallowell, 1842) vanicolous and was said to be common in the drier parts

of the country.

Material examined: ZFMK 56093, Ziama Forest south of

Sérédou, W. Bohme coll. 6—26 Oct. 1993; ZFMK 60771

and 62179-—180, Sérédou, W. Buitzler coll. June/Aug. 1995

and Nov./Dec. 1995 respectively. Osteolaemus tetraspis (Cope, 1861)

Remarks: Recorded by Rédel & Bangoura (2006) also. Material examined: voucher photograph.

from Déré Forest (Fig. 23).

Remarks: The existence of the dwarf crocodile in Ziama

Forest 1s proven by a photograph by W. Biitzler (Fig. 24)

taken near Sérédou in summer 1993. A particular site

where the species was observed near Malweta is shown

in Fig. 25. According to Villiers (1958), this species is

Fig. 23. Causus maculatus from Déré Forest (Photo: M.-O.

Rédel).

-_h\. es ee ue AN “a

Fig. 24. Osteolaemus tetraspis killed by local hunter near Mal- __ Fig. 25. Habitat near Malweta village where O. tetraspis was

weta village, Ziama Forest (Photo: W. Butzler). observed (Photo: W. Bohme).

Bonn zoological Bulletin 60 (1): 35-61 ©ZFMK

52 Wolfgang Bohme et al.

characteristic for the “foréts guinéennes” — Rédel & Ban-

goura (2006) recorded it from Diécké Forest — but also

for gallery forests in the savanna, e.g. near Kérouané in

Upper Guinea. Waitkuwait (1986, 1988, 1989) mentioned

for Guinea the rivers Kourai, Sankarani, Boa, Bafing and

Mafou of which only Boa and Bafing belong to the hy-

drographic system of the SE Guinean forest area.

Very recently, Eaton et al. (2009) challenged the conspeci-

fity of West African dwarf-snouted crocodiles with typi-

cal O. tetraspis from the Ogooué basin (Gabon).

Acknowledgements. First of all, WB wants to express his grat-

itude to Dr. Wilfried Biitzler, G6ttingen, in the early nineties head

of PROGERFOR, Conakry, for having invited him as a so-called

short term expert to Guinée forestiére to do an inventory of the

herpetological fauna of the classified forests Ziama and Diéckeé.

Dr. Biitzler provided indispensible hospitality and logistic help,

but took also part in the acquisistion of valuable voucher mate-

rial, even continuing these contribuons in the subsequent years.

Dr. Henning Vierhaus, Soest, having been a short term expert

for small mammals for PROGERFOR, also provided advice and

voucher material. The necessary permits for exporting the col-

lected specimens were kindly provided by M. Sagnah Saténin,

Dép. des Eaux, Forets et Chasses, Conakry. Some additional

Guinean specimens were donated to ZFMK by Dr. Guy Kremer,

Luxembourg. Taxonomic advice was provided by Dr. Barry

Hughes, London, by the late Dr. Jens B. Rasmussen, Copenhagen

(both for snakes), and by Dr. Andreas Schmitz, Geneve (partic-

ularly for scincid lizards). Mrs. Birgit Klasen, Bonn, helped with

recovering and analyzing the stomach contents of the snakes, and

Dr. Jorg Freyhof, Berlin, kindly identified the fishes found in

Afronatrix and Grayia. Last but not least, Ursula Bott (ZFMK

Bonn) provided again her technical skills with the preparation

of the manuscript. MOR and CB are in particular grateful to M.

Mohamad Alhassane Bangoura, M. Joseph Doumbia, Mrs. Lau-

ra Sandberger, Dr. Annika Hillers and M. Kaman Camara for

their great help during field work and/or contributing records.

They likewise thank very much Conservation International,

Washington D.C., Rio Tinto, bhp Billiton and the Société des

Mines de Fer de Guinée and the Declining Amphibian Popula-

tions Task Force for financing (amphibian) surveys in various

Guinean regions. Guinée Ecologie facilitated much some of our

field trips. The ‘Ministere de l’ Agriculture et des Eaux et Foréts’

and the ‘Direction Nationale pour la Protection de la Nature’,

Republic of Guinea, kindly provided collecting and export per-

mits. Last not least we thank Dr. Andreas Schmitz, Genéve, for

his valuable comments on the manuscript.

REFERENCES

Angel F (1932) Les serpents de Il’ Afrique occidentale frangaise.

Bulletin du Comité d’ Etudes historiques et scientifiques de

l Afrique Occidentale Frangaise 15: 613-858

Angel F (1944) Un lézard nouveau du Mont Nimba (Haute Gui-

née francaise) appartenant au genre Lygosoma. Bulletin du

Muséum national d’ Histoire naturelle (2éme sér.) 16: 293-294

Angel F, Guibé J, Lamotte M (1954 a) La Réserve Naturelle In-

tégral du Mont Nimba. Fasc. I (Extrait): Lézards. Mémoires

de I’Institut Fran¢ais d’ Afrique Noire, Dakar, 40: 371-379

Bonn zoological Bulletin 60 (1): 35-61

Angel F, Guibé J, Roy R (1954 b) La Réserve Naturelle Inté-

grale du Mont Nimba. Fasc. I (Extrait): Serpents. Mémoires

de I’Institut Frangais d’Afrique Noire, Dakar 40: 381-402

ATLANTA Consult (1988) Inventaire Forestier de la Guinée

Forestiere. Rapport Technique de Synthése partielle de 1 Mas-

sif de Ziama. Unpublished Report, Hamburg, 92 pp. + 5 an-

nexes

Barabanovy A (2008) A case of homonymy in the genus Agama.

Russian Journal of Herpetology 15: 206

Barnett A, Prangley M, Hayman PV, Diawara D, Koman J (1994)

A preliminary survey of Kounounkan Forest, Guinea, West

Africa. Oryx, London 28: 269-275

Bohme W (1994 a) Rapport sur la mission d’appui herpétolo-

gique (Expertise a court temps) dans les Foréts de Ziama et

de Diécké, Guinée Forestiere, Octobre 1993. Unpublished Re-

port, Conakry (PROGERFOR), 50 pp.

Bohme W (1994 b) Frésche und Skinke aus dem Regenwald-

gebiet Stidost-Guineas, Westafrika. I. Einleitung; Pipidae, Ar-

throleptidae, Bufonidae. Herpetofauna, Weinstadt 16: 11-19

Bohme W (1994 c) Frésche und Skinke aus dem Regenwaldge-

biet Stidost-Guineas, Westafrika. II. Ranidae, Hyperoliidae,

Scincidae; faunistisch-6kologische Bewertung. Herpetofauna,

Weinstadt 16: 6-16

Bohme W (2000) Diversity of a snake community in a Guinean

rainforest (Reptilia, Serpentes. In: Rheinwald G (Ed.), Isolat-

ed vertebrate communities in the tropics: Proceedings of the

4th International Symposium in Bonn, 400 pp. Bonner zool-

ogische Monographien 46: 69-78

Bohme W, Meinig H, Rédel M-O (1996) New records of am-

phibians and reptiles from Burkina Faso and Mali. British Her-

petological Society Bulletin 56: 7-26

Bohme W, Schmitz A, Ziegler T (2000) A review of the West

African skink genus Cophoscincopus Mertens (Reptilia: Scin-

cidae: Lygosominae): resurrection of C. simulans (Vaillant,

1884) and description of a new species. Revue Suisse de Zo-

ologie, Geneve 197: 777-791

Bohme W, Wagner P, Malonza P, Létters S, Kohler J (2005) A

new species of the Agama agama group (Squamata: Agami-

dae) from western Kenya, East Africa, with comments on Aga-

ma lionotus Boulenger, 1896. Russian Journal of Herpetology

12: 143-150

Bohme W, Ziegler T (1997) A taxonomic review of the Varanus

(Polydaedalus) niloticus (Linnaeus, 1766) species complex.

Herpetological Journal, London 7: 155-162

Boulenger GA (1910) Descriptions of four new African snakes

in the British Museum. Annals and Magazines of Natural His-

tory, London, 5 (8th ser.) 512—513

Branch WR, R6del M-O (2003) Herpetological survey of the

Haute Dodo and Cavally forests, western Ivory Coast, Part II:

Trapping results and reptiles. Salamandra 39: 21—38

Brandstatter F (1996) Die Sandrennattern. Magdeburg (West-

arp Wissenschaften) & Heidelberg (Springer) 142 pp.

Chabanaud P (1916) Enumeration des ophidiens non encore

étudiés de |’ Afrique Occidentale, appartenants aux collections

du Muséum, avec la description des espéces et des variations

nouvelles. Bulletin du Muséum d’ Histoire naturelle, Paris 22:

362-382

Chabanaud P (1917) Enumeration des reptiles non encore étudiés

de |’Afrique Occidentale, appartenants aux collections du

Muséum, avec la description des espéces nouvelles. Bulletin

du Muséum d’ Histoire naturelle, Paris 23: 83-105

Chabanaud F (1918) Etude complémentaire de deux Agama de

l’Afrique Occidentale et description de quatre espéces nou-

velles de reptiles de la méme région. Bulletin du Muséum d’

Histoire naturelle, Paris 24: 104-112

©ZFMK

Amphibians and reptiles from Guinea 53

Chabanaud P (1920) Contribution a étude de la faune herpé-

tologique. Note préliminaire sur les résultats d’une mission

scientifique en Guinée francaise (1919-1920). Bulletin du

Comité d’ Etudes historiques et scientifiques de I’ Afrique Oc-

cidentale Francaise 1920: 489-497

Chabanaud P (1921) Contribution a l’étude de la faune herpe-

tologique de Afrique Occidentale, 2iéme note. Bulletin du

Comité d’ Etudes historiques et scientifiques de I’ Afrique Oc-

cidentale Francaise 1921: 445-472

Chippaux J-P (2001) Les serpents d'Afrique occidentale et cen-

trale. Lyons, IRD Editions 292 pp.

Condamin M. (1959) Serpents récoltés a Sérédou (Guinée) par

R. Pujol. Bulletin de l'Institut Francais d’ Afrique Noire,

Dakar, (ser. A) 21: 1351-1366

David O, Ineich I (1999) Les serpents venimeux du monde: sys-

tématique et répartition. Dumerilia, Paris 3: 3-499

Eaton MJ, Martin A, Thorbjarnarson J, Amato G (2009) Species-

level diversification of African dwarf crocodiles (genus Os-

teolaemus): A geographic and phylogenetic perspective. Mo-

lecular Phylogenetics and Evolution, Amsterdam 50: 496-506

Gans C (1987) Studies on amphisbaenians (Reptilia). 7. The

small round-headed species (Cynisca) from western Africa.

American Museum Novitates, New York 2896: 1-84

Golay P, Smith HM, Broadley DG, Dixon JR, McCarthy C, Rage

J-C, Schatti B, Toriba M (1993) Endoglyphs and other major

venomous snakes of the world. A checklist. Geneva (Azemiops

S.A.) xv+478 pp.

Gramentz D (2005) Die Nilweichschildkrote Trionyx triunguis.

Frankfurt/Main (Ed. Chimaira) 166 pp.

Grandison AGC (1956) On a collection of lizards from West

Africa. Bulletin de I’Institut Francais d’ Afrique Noire, Dakar

(sér. A) 28: 224-244

Greenbaum E, Carr JL (2005) The herpetofauna of Upper Niger

National Parc, Guinea, West Africa. Scientific Papers of the

Natural History Museum of the University of Kansas 37: 1—21

Henle K, B6hme W (2003) A new species of Hemidactylus

(Squamata: Gekkonidae) from West Africa, and comments on

species hitherto confused with H. muriceus. African Journal

of Herpetology 52: 23-38

Herrmann H-W, Bo6hme W, Euskirchen O, Herrmann PA,

Schmitz A (2005) African biodiversity hotspots: the reptiles

of Mt. Nlonako, Cameroon. Revue Suisse de Zoologie,

Geneve 112: 1045-1069

Hillers A, Bangoura MA, Loua NS, R6édel M-O (2006) Inven-

taire rapide des amphibians et des reptiles dans la région de

Boke dans le nord-ouest de la Guinée. — pp. 59-64; Rapid sur-

vey of amphibians and reptiles in the Boké region, northwest-

ern Guinea. — pp. 131-136, appendices/annexes 5—7: pp.

178-181, In: Wright HE, McCullough J, Diallo, MS (eds.):

Un inventaire biologique rapide de la préfecture de Boké dans

le nord-ouest de la Guinée/A rapid biological assessment of

Boke Préfecture, northwestern Guinea. RAP Bulletin of Bio-

logical Assessment 41/Bulletin RAP d’Evaluation Rapide 41,

Conservation International, Washington D.C. 190 pp.

Hillers A, Loua N-S, Rédel M-O (2008a) Assessment of the dis-

tribution and conservation status of the viviparous toad Nim-

baphrynoides occidentalis on Monts Nimba, Guinea. Endan-

gered Species Research 5: 13-19

Hillers A, Loua N-S, R6del M-O (2008b) A preliminary assess-

ment of the amphibians of the Fouta Djallon, Guinea, West

Africa. Salamandra 44: 113-122

Hillers A, Zimkus B, R6del M-O (2008c) A new Phrynobatra-

chus (Amphibia: Anura: Phrynobatrachidae) from north-west-

erm Guinea, West Africa. Zootaxa 1815: 43-50

Bonn zoological Bulletin 60 (1): 35-61

Ineich I (2003) Contributions a la connaisance de la biodiver-

sité des régions afro-montagnardes: les Reptiles du Mont

Nimba. In: Lamotte M, Roy R (Eds.) Le peuplement animal

du Mont Nimba (Guinée, Cote d’Ivoire, Liberia). Mémoires

du Muséum National d’ Histoire naturelle, Paris 190: 597-637

Joger U (1979) Zur Okologie und Verbreitung wenig bekannter

Agamen Westafrikas (Reptilia: Sauria: Agamidae). Salaman-

dra, Frankfurt am Main 15: 31—52

Joger U (1980) Eine neue Art der Gattung 7arentola (Reptilia:

Sauria: Gekkonidae) aus Westafrika. Amphibia-Reptilia,

Wiesbaden |: 137—147

Joger U, Lambert MK (2002) Inventory of amphibians and rep-

tiles in SE Senegal, including the Niokolo-Koba National Park,

with observations on factors influencing diversity. Tropical

Zoology, Firenze 15: 165-185

Klaptocz A (1913) Reptilien, Amphibien und Fische aus Fran-

zosisch-Guinea. Zoologische Jahrbiicher ftir Systematik, Je-

na 34: 279-290

Lamotte M (1983) The undermining of Mount Nimba. Ambio

12: 144-179

Lamotte M, Aguesse P, Roy R (1962) Données quantitatives sur

une biocénose ouest-africaine: la prairie montagnarde du Nim-

ba (Guinée). Terre et Vie, Paris 1962: 351-370

Mocquard F (1902) Sur des reptiles et batraciens de Il’ Afrique

Orientale anglaise, du Gabon et de la Guinée francaise. Bul-

letin du Muséum National d’Histoire Naturelle, Paris 8:

404-417

Mocquard F (1905) Description de deux espéces de reptiles. Bul-

letin du Muséum National d’Histoire Naturelle, Paris 11:

288-290

Mocquard F (1908) Description de quelques reptiles et d’un ba-

tracien nouveaux de la collection du Muséum. Bulletin du

Muséum National d’Histoire Naturelle, Paris 14: 259-262

Parker HW (1939) Résultats scientifiques des croisiéres du

Navire-Ecole Belge “Mercator”. V. Reptilia et Amphibia. Mé-

moires du Musée Royal d’Histoire Naturelle de Belgique,

Bruxelles (2) 15: 85—90

Perret JL (1973) Contribution a l’étude des “Panaspis” (Rep-

tilia, Scincidae) d’ Afrique occidentale avec la description de

deux espéces nouvelles. Revue Suisse de Zoologie, Genéve

80: 595-630

Porembski S, Barthlott W, Dérrstock S, Biedinger N (1994) Veg-

etation of rock outcrops in Guinea: granite inselbergs, sand-

stone table mountains and ferricretes — remarks on species

numbers and endemism. Flora, Jena 189: 315-329

Porembski S, Biedinger N, Dorrstock S (1995) Zur Vegetation

Guineas unter besonderer Beriicksichtigung der Inselbergflo-

ra. Natur und Museum, Frankfurt am Main 125: 143-154

Pritchard R (1979) Encyclopedia of turtles. Hongkong (T.F.H.

Publ. Inc. Ltd.) 895 pp.

Rasmussen JB (1989) A taxonomic review of the Dipsadoboa

duchesnii complex. Bonner zoologische Beitrage 40: 249-264

Rasmussen JB (1993) A taxonomic review of the Dipsadoboa

unicolor complex, including a phylogenetic analysis of the

genus (Serpentes, Dipsadidae, Boiginae). Steenstrupia, Koben-

havn 19: 129-196

Rasmussen JB (1994) Studier over afrikanske, furetandede

traesnoges (slaegten Dipsadoboa’s), taxonomi, fylogeni og

zoogeografi (Serpentes, Dipsadidae, Boiginae). Kobenhavn

(Universitetets Zoologiske Museum)16 pp.

Rasmussen JB (2005) A review of the slender stiletto snake,

Atractaspis aterrima Ginther, 1863 (Serpentes Atractaspidi-

dae). Tropical Zoology, Firenze 18: 137-148

Riquier M, Bohme W (1996) Bemerkungen zu Verbreitung und

geographischer Variation sowie zu Freileben und Haltung der

©ZFMK

54 Wolfgang Bohme et al.

Lianennatter, Thelotornis kirtlandii (Hallowell, 1844). Her-

petofauna, Weinstadt 18: 27-34

Rédel M-O, Bangoura MA (2004a) A conservation assessment

of amphibians in the Forét Classée du Pic de Fon, Simandou

Range, southeastern Republic of Guinea, with the description

of anew Amnirana species (Amphibia Anura Ranidae). Trop-

ical Zoology, Firenze 17: 201—232

Rédel M-O, Bangoura MA (2004b) Rapid survey of amphibians

and reptiles in the Forét Classée du Pic de Fon, Guinea. pp.

51-60; Evaluation rapide des amphibiens et des reptiles dans

la Forét Classée du Pic de Fon, Guinée. — pp. 151-164, ap-

pendices/annexes 3—5: pp. 232—237, In: McCullough J. (Ed.):

A rapid biological assessment of the Forét Classée du Pic de

Fon, Simandou Range, South-eastern Republic of Guinea/Une

évaluation biologique rapide de la Forét Classée du Pi de Fon,

chaine du Simandou, Guinée. RAP Bulletin of Biological As-

sessment 35/Bulletin RAP d’ Evaluation Rapide 35, Conser-

vation International, Washington D.C. 248 pp.

Rédel M-O, Bangoura MA (2006) Un inventaire herpétologique

de trois foréts classées dans le sud-est de la République de

Guinée. — pp. 53-58; A herpetological survey of Déré, Diécké

and Mt. Béro classified forests, southeastern Guinea. — pp.

153-158, appendices/annexes 5—7: pp. 229-235, In: Wright

HE, McCullough J, Alonso LE, Diallo MS (Eds.): Une éval-

uation biologique rapide de trois forét classées du sud-est de

la Guinée/A rapid biological assessment of three classified

forests in southeastern Guinea. RAP Bulletin of Biological As-

sessment 40/Bulletin RAP d’Evaluation Rapide 40, Conser-

vation International, Washington D.C. 249 pp.

Rodel M-O, Bangoura MA, Bohme W (2004) The amphibians

of the south-eastern Republic of Guinea (Amphibia:

Gymnophiona, Anura). Herpetozoa, Wien 17: 99-118

Rodel M-O, Grabow K, Béckheler C, Mahsberg D (1995): Die

Schlangen des Comoé-Nationalparks, Elfenbeinktiste (Repti-

lia: Squamata: Serpentes). Stuttgarter Beitrage zur Naturkun-

de, Serie A 528: 1-18

Rodel M-O, Kouadio K, Mahsberg D (1999) Die Schlangenfau-

na des Comoé-Nationalparks, Elfenbeinktiste: Erganzungen

und Ausblick. Salamandra, Rheinbach 35: 165—180

Rédel M-O, Mahsberg D (2000) Vorlaufige Liste der Schlangen

des Tai-Nationalparks/Elfenbeinktiste und angrenzender Ge-

biete. Salamandra, Rheinbach 36: 25-38

Rodel M-O, Sandberger L, Doumbia J, Hillers A (2009) Reval-

idation of Phrynobatrachus maculiventris Guibé & Lamotte,

1958 and description of its aposematic coloured tadpole.

African Journal of Herpetology 58: 15—27

Rédel M-O, Sandberger L, Penner J, Mané Y, Hillers A (2010)

The taxonomic status of Hyperolius spatzi Ahl, 1931 and Hy-

perolius nitidulus Peters, 1875 (Amphibia: Anura: Hyperoli-

idae). Bonn zoological Bulletin 57: 177-188

Roman B (1976) Serpents mortels de l’Ouest A fricain. Les Jour-

nées scientifiques du Centre national de coordination des étu-

des et recherches sur la nutrition et l’alimentation, Paris 6:

1-56

Roux-Estéve R (1969) Etude comparée de Meizodon coronatus

(Schlegel) et de Meizodon regularis Fischer (Colubridés — Ser-

pentes). Bulletin du Muséum National d’ Histoire Naturelle,

Paris 2eme sér., 41: 395-409

Roux-Esteve R (1974) Révision systématique des Typhlopidae

d’ Afrique (Reptilia, Serpentes). Mémoires du Muséum Natio-

nal d’ Histoire Naturelle, Paris (Nouvelle Série) (A) 87: 1-313

Schatti B (1985) Systematics of East African species of

Meizodon Fischer, 1856 (Reptilia Serpentes Colubridae). Mon-

itore Zoologico Italiano, Firenze, n.s., Suppl. 20: 149-175

Bonn zoological Bulletin 60 (1): 35—61

Schatti B, Trape J-F (2008) Bamanophis, a new genus for the

West African colubrid Periops dorri Lataste, 1888 (Reptilia:

Squamata: Colubridae). Revue Suisse de Zoologie, Genéve

115: 595-615

Schmitz A, Mausfeld P, Hekkala E, Shine T, Nickel H, Amato

G, Béhme W (2003) Molecular evidence for species-level di-

vergence in African Nile crocodiles Crocodylus niloticus (Lau-

renti, 1786). Comptes Rendus Palevol, Paris 2: 703-712

Spawls S, Branch WR (1995) The dangerous snakes of Africa:

Natural history, species directory, venom and snakebite. Sani-

bel/Florida (Curtis Books) 192 pp

Stoll F (2008) Taxonomie afrikanischer Skinke: Morphologie

und Variation von Trachylepis affinis und Trachylepis per-

rotetii (Reptilia; Squamata; Scincidae). MSc Thesis, TU Darm-

stadt 128 pp.

Trape J-F, Baldé C (2006) Geographic distribution. Haemorrhois

dorri (Boie’s racer). Herpetological Review 37: 362

Trape J-F, Mané Y (2006 a) Le genre Dasypeltis Wagler (Ser-

pentes: Colubridae) en Afrique de |’Ouest: description de trois

espéces et d’une sous-espece nouvelles. Bulletin de la Société

Herpétologique Frangaise, Paris 119: 27-56

Trape J-F, Mané Y (2006 b) Guide des serpents d’Afrique occi-

dentale. Savane et désert. Paris (IRD éd.) 226 pp

Ullenbruch K, Grell O, BOhme W (2010) Reptiles from south-

ern Benin, West Africa, with the description of a new Hemi-

dactylus (Gekkonidae), and a country-wide checklist. Bonn

zoological Bulletin 57: 31-54

Villiers A (1950) La collection de serpents de I’ Institut Frangais

d’ Afrique Noire, Dakar (Institut Francais d’ Afrique Noire) 155

pp.

Villiers A (1954) La collection de serpents de I’ Institut Frangais

d’Afrique Noire, Dakar (Acquisitions 1953). Bulletin de I’In-

stitut Francais d’Afrique Noire, Dakar (A) 16: 1234-1247

Villiers A (1958) Tortues et crocodiles d’ Afrique Noire francaise.

Initiations africaines, Dakar (IFAN) 15: 1-354

Wagner P, Bohme W (2009) On the nomenclatural status of Aga-

ma cristata Mocquard, 1905; areply to A. Barabanov (2008).

Russian Journal of Herpetology 16: 161-162

Wagner P, Ineich I, Leaché A, Wilms T, Trape S, BGhme W,

Schmitz A (2009) Studies on African Agama VI. Taxonomic

status of the West African Agama (Sauria: Agamidae) with

prominent tail crests: Agama boulengeri Lataste, 1886, Aga-

ma insularis Chabanaud and Agama cristata Mocquard, 1905.

Bonner zoologische Beitrage 56: 239-254

Waitkuwait WE (1986) Contribution a 1’ étude des crocodiles de

l’Afrique de l’ouest. Nature et Faune, Abidjan 1

Waitkuweit WE (1988) Untersuchungen zur Erhaltung und Be-

wirtschaftung von Krokodilen in der Republik Cote d’ Ivoire

(Westafrika). PhD thesis, Universitat Heidelberg 279 pp.

Waitkuwait WE (1989) Present knowledge on the West African

slender-snouted crocodile, Crocodylus cataphractus Cuvier,

1824 and the West African dwarf crocodile, Osteolaemus

tetraspis Cope, 1861. Pp. 260-275 in: Crocodiles, their ecol-

ogy, management and conservation. Gland (IUCN).

Wallach V (1994) Aparallactus lineatus (Peters) and Aparallac-

tus niger Boulenger: two valid species from West Africa. Jour-

nal of Herpetology 28: 95—99

Welch KG (1982) Herpetology of Africa: A checklist and bib-

liography of the orders Amphisbaenia, Sauria and Serpentes.

Malabar/Florida (Krieger Publishing Company) 293 pp.

Wermuth H, Mertens R (1961) Liste der Schildkroten, Kroko-

dile und Briickenechsen (reprinted 1995, with an addendum

by FJ Obst). Jena (Gustav Fischer) 506 pp.

Witte GF, Laurent R (1947) Révision d’un groupe de Colubri-

dae africains. Genres Calamelaps, Miodon, Aparallactus et

©ZFMK

Amphibians and reptiles from Guinea 55

formes affines. Mémoires du Musee Royal d’ Histoire naturelle

de Belgique (2) 29: 1-134

Appendix I

Checklist of reptiles known from the Republic of Guinea

The following checklist does not aim to revise all following

genus and species names. Some are adapted to modern use, but

original use of the respective author’s name is added. Only in

case of small spelling differences e.g. Chamaeleon vs.

Chamaeleo, Dendraspis vs. Dendroaspis, smythii vs. smithii etc.

the original spelling has not been added. Quite a number of

species identifications from the literature had to be taken bona

fide, because a reinvestigation of old vouchers which might be

necessary in view of more recent revisions is beyond the scope

of this list.

Only references giving locality data were included but not those

just enumerating only the country of occureence, except in such

cases where no other and more detailed mentioning of the Re-

public of Guinea is made (see e.g. Welch 1982, Golay et al. 1993,

Spawls & Branch 1995, David & Ineich 1999). A special prob-

lem is that the old reference “Guinea” or Guinée” can also re-

fer to Ghana (David & Ineich 1999).

The families below are arranged systematically, with alphabet-

ical order of genera and species within families.

Pelomedusidae

— Pelomedusa subrufa — Greenbaum & Carr (2005): PN Haut

Niger (Ist country record); Fouta Djallon, Pita/Hore Binti, N

10°51°21.3”, W 12°32°40.2”, photo records, C. Brede.

— Pelusios castaneus — Ineich (2003): Mt. Nimba; Greenbaum

& Carr (2005): Kouroussa; PN Haut Niger; Hillers et al. (2006):

Kolaboui.

— Pelusios sp. — Chabanaud (1921): Kérouaneé (= P. castaneus

22).

Testudinidae

— Kinixys belliana — Greenbaum & Carr (2005): PN Haut Niger.

— Kinixys erosa — Ineich (2003): Mt. Nimba; Rédel &Bangoura

(2006): Diécké Forest; this paper: Sérédou, Macenta.

— Kinixys homeana — Chabanaud (1921): N’Zébéla (as Cinixys

homeana).

Trionychidae

— Trionyx triunguis — Greenbaum & Carr (2005): PN Haut Niger

(1st country record); this paper: Ziama Forest.

Agamidae

— Agama agama complex — Klaptocz (1913): Conakry, Dubre-

ka, Mamou, Dabola, Tinkisso Betaja (as A. colonorum); Cha-

banaud (1921): “Trés commun partout” (as A. colonorum); An-

gel et al. (1954 a): several localities in Mt. Nimba range; Gran-

dison (1956): Mt. Nimba (see also Ineich 2003); Greenbaum &

Carr (2005): PN Haut Niger; Rodel & Bangoura (2004b): Pic

Bonn zoological Bulletin 60 (1): 35—61

de Fon/Simandou range; Rédel & Bangoura (2006): Deré an

Diécké Forests, Mt. Béro; Hillers et al. (2006): Kolaboui, San-

garédi; this paper.

— Agama cristata — Mocquard (1905): “Pays Sankaran” (on

MNHN label: “Bomanesco, Sankaran); Chabanaud (1921):

Sankaran. — Note: We regard the name 4. maria nomen nov.

Barabanov (2008) as invalid. Reasons are given by Wagner &

Bohme (2009). We also disagree with Barabanov’s (2009) view

to locate “Pays Sankaran” in Mali rather than in Guinea.

— Agama insularis — Chabanaud (1918) Los Islands off

Conakry; Parker (1939): Los Islands; Wagner et al. (2009):

Kinkon water falls; Kindia (Pastoria station).

— Agama sankaranica — Chabanaud (1921): “Pays Sankaran”

(see above under A. cristata), N’Zébéla; Rodel & Bangoura

(2006): Mt. Beéro.

Chamaeleonidae

— Chamaeleo gracilis — Klaptocz (1913): Dabola; Chabanaud

(1921): Kérouané, Beyla; Ineich (2003): Mt. Nimba; Greenbaum

& Carr (2005): PN Haut Niger; Rodel & Bangoura (2006): Mt.

Béro; Hillers et al. (2006): Rio Kapatchet (Kamsar), Sangaré-

di; this paper: Kindia; Ziama Forest.

— Chamaeleo senegalensis — Chabanaud (1921): Beyla; Gran-

dison (1956): Koundara; Rédel & Bangoura (2004b): Pic de

Fon/Simandou range; this paper: Kindia; Fouta Djallon,

Labé/Saala, N 11°17713.9", W 12°30°13.5”, photo record, C.

Brede; Pic de Fon/Simandou range, M.A. Bangoura, C. Brede,

M.-O. Rédel.

Eublepharidae

— Hemitheconyx caudicinctus — Mocquard (1908): Kouroussa (as

Psilodactylus caudicinctus); Grandison (1956): Kankan; Green-

baum & Carr (2005): PN Haut Niger.

Phyllodactylidae

— Tarentola parvicarinata — Joger (1980): “nérdliches Guinea”;

this paper: Dabola/ Kouffo, Fouta Djallon, photo record C.

Brede. — Note: The gecko on Fig. 26 was captured but not col-

lected, but its habitus and (phalangeal) foot structure strongly

suggest its allocation to Tarentola parvicarinata. This photo-

graphic voucher 1s the second country record of the genus and

the first with precise locality data. Remarkably, it was found in

a gallery forest habitat where it lived on stones offering it good

camouflage.

Gekkonidae

— Cnemaspis occidentalis — Ineich (2003): Mt. Nimba, Ziéla,

Nzo; this paper.

— Hemidactylus angulatus — Klaptocz (1913: Conakry, Mamou;

Chabanaud (1921): Conakry, Kérouané, Beyla (as H. brookii);

Grandison (1956): Kankan, Mamou (as H. brookii angulatus);

Greenbaum & Carr (2005): PN Haut Niger; Rodel & Bangoura

(2006): Diécké Forest (as H. brookii).

©ZFMK

56 Wolfgang Bohme et al.

Fig. 26. Tarentola parvicarinata from Fouta Djallon Mts., W

Guinea (Photo: C. Brede).

— Hemidactylus fasciatus — R6del & Bangoura (2006): Diécké

Forest; this paper: Mt. Nimba, photo record, C. Brede.

— Hemidactylus mabouia — Rédel & Bangoura (2006): Diécké

Forest); this paper.

— Hemidactylus muriceus — Henle & Béhme (2003): Diéckeé For-

est; Rédel & Bangoura (2004b): Pic de Fon/ Simandou range;

Rédel & Bangoura (2006): Diécké Forest and Mt. Béro; this pa-

per; Mt. Nimba, photo records, C. Brede, M.-O. Rédel.

— Hemidactylus sp. (aff. mabouia): Coyah (this paper). — Note:

Three specimens of this Hemidactylus similar to H. mabouia

were captured but not collected on October 2, 1993 at the very

beginning of WB’s mission. They were active at night on the

stony walls of huts in a small tourist resort in the tidal zone NE

of Conakry. Despite our expectation, they were not found again,

and a later comparison of the photographs revealed that they are

different from H. mabouia, not only by the lack of the fine, wavy

dorsal crosslines (which are lacking also in some other

“mabouia” populations, e.g. in Benin, see Ullenbruch et al. 2010)

but also by the presence of two pairs of two large, darkened sad-

dle-like flecks separated by a light middorsal line (Fig. 27 & 28).

Moreover, the lowermost flank tubercles were more strongly de-

veloped than in H. mabouia (see Figs.). We regard this commen-

sal gecko to represent an undescribed species.

Lacertidae

— Heliobolus nitidus — Klaptocz (1913): Iryan on the Niger (as

Eremias nitida).

— Holaspis guentheri — Angel et al. (1954 a): 3 places in the Mt.

Nimba area; this paper: N’Zérékoré; near Sérédou.

Scincidae

— Cophoscincopus durus — Grandison (1956): Mt. Nimba; Ine-

ich 2003): Mt. Nimba; Rédel & Bangoura (2006): Diécké For-

est; this paper.

— Cophoscincopus greeri — Bbhme (1994 c): Mt. Nimba (as C.

simulans); Bohme et al. 2000: Mt. Nimba; Ineich 2003: Mt. Nim-

ba; this paper.

— Cophoscincopus simulans — Chabanaud (1921): N’Zébéla,

N’Zerékoré (as Lygosoma simulans); Bohme (1994 c, Bohme

Bonn zoological Bulletin 60 (1): 35-61

et al. 2000): Ziama Forest near Sérédou; Ineich 2003): Mt. Nim-

ba; Rodel & Bangoura (2006): Déré and Diécké Forests and Mt.

Béro; this paper.

— Lepidothyris fernandi — Chabanaud (1921): N’Zébéla (as Ly-

gosoma (Riopa) fernandi); Grandison (1956): Mt. Nimba (as Ri-

opa fernandi); Ineich (2003); Mt. Nimba (as Lygosoma fernan-

di).

— Panaspis nimbensis — Angel et al. (1954 a): Mt. Nimba; BGhme

(1994 c): Sérédou: Ineich (2003): Mt. Nimba; this paper.

— Panaspis togoensis — Chabanaud (1917, 1921); Type locality

“Haute Guinée fran¢aise” (as Paralygosoma monneti, syn. fide

J.L. Perret & A. Schmitz, pers. comm.); Greenbaum & Carr

Fig. 27-28. Hemidactylus sp. (n.?) from Coyah, SW Guinea

(Photos: W. B6hme).

©ZFMK

Amphibians and reptiles from Guinea 57

(2005): PN Haut Niger (LZ. “africanum” in Chabanaud (1921:

Kankan, Kérouane) certainly refers to this species).

~ Trachylepis affinis — Klaptocz (1913): Conakry, Mamou, Konk-

oure, Iryan (as Mabuia raddonii); Chabanaud (1921): Kérouané,

Beyla, Macenta, N’Zébéla, N’Zerékore, Diécké, Sanikolé (as M.

raddonii); Grandison (1956): Mt. Nimba (as Mabuya blandingii);

photo record, C. Brede; B6hme (1994 c): Ziama Forest (as M.

affinis); Ineich (2003, as Euprepis affinis): Mt. Nimba; Rédel

& Bangoura (2004b): Pic de Fon/ Simandou range (as Mabuya

affinis); Greenbaum & Carr (2005): PN Haut Niger; Hillers et

al. (2006): Kolaboui, Rio Kapatchet (Kamsar), Sangarédi (as Ma-

buya affinis); this paper.

— Trachylepis albilabris — Angel et al. (1954 a, as Mabuya

Blandingii), Hoogmoed (1974, as M. affinis), Ineich (2003, as

Euprepis albilabris): several localities in the Mt. Nimba area;

Rédel & Bangoura (2006): Déré & Diécké Forests and Mt. Béro

(as Mabuya affinis).

— Trachylepis keroanensis — Chabanaud (1921): Kérouané (as

Mabuia perroteti var. keroanensis ssp. n. — Note: The two syn-

types MNHN 1921.0323—0324 are strikingly different from 7.

perroteti (see Stoll 2008 and Fig. 29) and seem to belong to the

elongate, short-legged 7: buettneri Matschie, 1910/7. sudanen-

sis Schmidt, 1919 complex sensu Hoogmoed 1974, which is un-

der review by WB and coworkers).

— Trachylepis maculilabris — Chabanaud (1921): Kérouané, Ma-

centa, N’Zébéla, N’Zérékoré, Beyla, Diécké Forest, Sanikolé (as

Mabuya maculilabris); Angel et al. (1954 a, as M. Blandingii

and M. polytropis (in part.); Grandison (1956): Mt. Nimba (as

Mabuya m. maculilabris); Bohme (1994 c): Ziama Forest (as M.

maculilabris); Ineich (2003, as Euprepis maculilabris): Mt. Nim-

ba; this paper.

— Trachylepis perrotetii — Klaptocz (1913): Conakry, Mamou (as

Mabuia perrotetii) Chabanaud (1921): Kankan, Beyla (as M. per-

Fig. 29. Syntypes of Zrachylepis keroanensis Chabanaud,

from Kérouane (MNHN 9121.0323—0324), a valid species of the

T. buettneri/sudanensis group (Photo: W. Bohme).

rotetii); Fouta Djallon, Labé/Daralabe mountain, N 11°13736.9”,

W 12°16753.2”, photo record, C. Brede.

— Trachylepis quinquetaeniata — Klaptocz (1913): Tinkisso near

Dabola (as Mabuia quinquetaeniata).

Bonn zoological Bulletin 60 (1): 35-61

Amphisbaenidae

— Cynisca cf. liberiensis — Angel et al. (1954 a): Mt. Nimba; this

paper: Ziama Forest.

Varanidae

— Varanus exanthematicus — Klaptocz (1913): “am Niger”;

Greenbaum & Carr (2005): PN Haut Niger.

— Varanus niloticus — Chabanaud (1921): everywhere common;

Greenbaum & Carr (2005): PN Haut Niger; Hillers et al. (2006):

Kolaboui, Rio Kapatchet (Kamsar), Sangarédi; Fouta Dyallon,

Pita/Hore Binti, N 10°51°04.8”, W 12°31714.1”, photo record

(Fig. 30) C. Brede.

— Varanus ornatus — Chabanaud (1921): the former statement

of this author implies this species also to be present in the for-

est region (as V. niloticus); Ineich (2003): Mt. Nimba; Rédel &

Bangoura (2004b): Pic de Fon/Simandou range; this paper: Séré-

dou, Ziama Forest, Diécké Forest.

Leptotyphlopidae

— Leptotyphlops bicolor — Trape & Mané (2006): on grid map

but without specific locality.

— Rhinoleptus koniagui — Trape & Mané (2006): on grid map

but without specific locality.

Typhlopidae

— Typhlops liberiensis — Mocquard (1902): Kouroussa; Villiers

(1954), Angel et al. (1954 b): Mt. Nimba (as 7! (p). punctatus);

Roux-Estéve (1974): Mt. Nimba (as 7. p. liberiensis); Bohme

(2000): Djani River nr. NZébéla, N’Zérékoré (as 7. p. liberien-

sis); Ineich (2003): Mt. Nimba; this paper.

— Typhlops manni — Guibé (1952), Angel et al. (1954 b), Ineich

(2003): Mt. Nimba. (all three papers refer to the single holotype

of 7. angeli Guibé, 1952 which was synonymyized with 7? man-

ni by Roux-Estéve 1974, see Ineich 2003).

— Typhlops punctatus — Chabanaud (1921): N’Zérékoré; Villiers

(1950): Dalaba; Condamin (1959): Sérédou; Greenbaum & Carr

(2005): PN Haut Niger.

Pythonidae

— Python regius — Trape & Mané (2006): on grid map but with-

out specific locality .

— Python sebae — Chabanaud (1921): Macenta; Villiers (1954),

Angel et al. (1954 b), Ineich (2003: Mt. Nimba; R6édel & Ban-

goura (2004b): Pic de Fon/Simandou range; Greenbaum & Carr

(2005): PN Haut Niger; this paper: Mt. Nimba (Fig. 13).

Boidae

— Calabaria reinhardtii — Bohme (2000): Sérédou (1st country

record); Ineich (2003): Mt. Nimba (Ziéla); this paper.

©ZFMK

58 Wolfgang Bohme et al.

Lamprophiidae

— Bothrophthalmus lineatus — Chabanaud (1921): N’Zérékoré;

Villiers (1954), Angel et al. (1954 b, as B. /. lineatus): Mt. Nim-

ba; Condamin (1959): Sérédou; BOhme (2000): Ziama Forest;

Ineich (2003): Mt. Nimba (as B. /. lineatus); R6del & Bangoura

(2006): Diécké Forest); this paper.

— Chamaelycus fasciatus — Chabanaud (1921): N’Zérékoré (as

Lycophidium fasciatum), Villiers (1954), Angel et al. (1954 b):

Mt. Nimba (as Oophilositum fasciatum); Ineich (2003): Mt. Nim-

ba.

— Gonionotophis granti — Trape & Mané (2006 b): on grid map

but without specific locality.

— Gonionotophis klingi — Villiers (1954), Angel et al. (1954 b),

Ineich (2003): Mt. Nimba

— Hormonotus modestus — Viliers (1954), Angel et al. (1954 b),

Ineich (2003): Mt. Nimba.

—Lamprophis fuliginosus — Klaptocz (1913): Konkouré (as Boo-

don fuliginosus); Villiers (1950), Angel et al. (1954 b): Mt. Nim-

ba (as Boaedon fuliginosus); Bbhme 2000): Ziama Forest; Ine-

ich (2003): Mt. Nimba; this paper.

— Lamprophis lineatus — Villiers (1950): Mamou (as Boaedon

lineatus); Villiers (1954), Angel et al. (1954 b): Mt. Nimba (as

B. lineatus); Bohme (2000): Ziama Forest; Ineich: Mt. Nimba;

this paper.

—Lamprophis olivaceus — Chabanaud (1921): Diécké Forest (as

Boodon olivaceus); Angel et al. (1954 b): Mt. Nimba (as Boae-

don olivaceus); Bohme (2000): Ziama Forest; Ineich (2003): Mt.

Nimba; Hillers et al. (2006): Rio Kapatchez, Kamsar; this pa-

per.

— Lamprophis virgatus — Chabanaud (1921):N’Zébéla,

N’Zérékoré (as Boodon virgatus); Villiers (1950), Angel et al.

(1954 b): Mt. Nimba (as Boaedon virgatus); Condamin (1959):

Sérédou; Bohme (2000): Ziama Forest; Ineich (2003): Mt. Nim-

ba; this paper.

Fig. 30. Varanus niloticus, juvenile specimen from the Fouta

Djallon range, W Guinea (Photo: C. Brede).

Bonn zoological Bulletin 60 (1): 35-61

— Lycophidion albomaculatum — Trape & Mané (2006 a): on grid

map but without specific locality.

—Lycophidion irroratum — Villers (1954), Angel et al. (1954 b):

Mt. Nimba (as L. capense (in part); Trape & Mané (2006): Kin-

dia.

—Lycophidion nigromaculatum — Angel et al. (1954 b): Mt. Nim-

ba (as L. irroratum (in part)); Ineich (2003): Mt. Nimba.

—Lycophidion semicinctum — Villiers (1950), Angel et al. (1954

b): Mt. Nimba; Trape & Mane (2006 b): on grid map but with-

out specific locality.

— Mehelya crossi — Trape & Mané (2006 b): on grid map but

without specific locality.

— Mehelya guirali — Bohme (2000): Ziama Forest; Ineich (2003):

Mt. Nimba; this paper.

— Mehelya poensis — Chabanaud (1921): near N’Zébéla (as Simo-

cephalus poensis); Villiers (1950, 1954), Angel et al. (1954 b):

Mt. Nimba; Bohme (2000): Ziama Forest; Ineich (2003): Mt.

Nimba; this paper.

— Mehelya stenophthalmus — Villiers (1954), Angel et al. (1954

b): Mt. Nimba; BOhme (2000): Ziama Forest; Ineich (2003): Mt.

Nimba; this paper.

Colubridae

— Bamanophis dorri — Trape & Baldé (2006): Kalekouré (as

Haemorrhois dorri); Schatti & Trape (2008): Kalekoure.

— Crotaphopeltis hippocrepis — Trape & Mané (2006 b): Kin-

dia.

— Crotaphopeltis hotamboeia — Mocquard (1902): Kouroussa (als

Leptodira hotamboeia); Chabanaud (1921): Macenta, Diécké

Forest (as L. hotamboeia); Condamin (1959): Sérédou; Bohme

(2000): Ziama Forest; Greenbaum & Carr (2005): PN Haut

Niger; Trape & Mané (2006 b): Labé; this paper.

— Dasypeltis confusa — Greenbaum & Carr (2005): PN Haut

Niger (as D. scabra); Trape & Mané (2006 a): Kilissi,

Kouroumaya, Kalekouré, Foulaya, Friguiagbé, Camarabunyi,

Seffan, Madina, Hamdalaye, Sankalabadou, Dalakan.

— Dasypeltis gansi — Trape & Mané (2006 a, b): on grid map

but without specific locality, despite all (but only non-Guinean)

localities listed in Trape & Mané (2006 a).

— Dasypeltis fasciata — Bohme (2000): Ziama Forest; Ineich

(2003): Mt. Nimba; Trape & Mané (2006 a): this paper.

— Dasypeltis cf. scabra — Mocquard (1902): Kouroussa; Ineich

(2003) and this paper: Mt. Nimba; Trape & Mané (2006 a):

Dalakan.

— Dipsadoboa brevirostris — Chabanaud (1921): N’Zérékoré,

Diécké Forest (as Leptodira guineensis); Rédel & Bangoura

(2006): Mt. Béro; Hillers et al. (2006): Sangarédi; this paper.

— Dipsadoboa sp.? — Mocquard (1902): Kouroussa (as L. pobe-

guini).

— Dipsadoboa underwoodi — Angel et al. (1954 b): Mt. Nimba

(as D. unicolor); Rasmussen (1993), Ineich (2003): Mt. Nimba.

— Dipsadoboa unicolor — Klaptocz (1913): Mamou; Chabanaud

(1921): Macenta, N’Zéréjoré; Villiers (1950): Dalaba; Villiers

©OZFMK

Amphibians and reptiles from Guinea 59

(1954), Angel et al. (1954 b): Mt. Nimba; B6hme (1994 a): Zia-

ma Forest; Ineich (2003): Mt. Nimba; Rédel & Bangoura (2006):

Diécké Forest; this paper.

— Dipsadoboa viridis — Villiers 1954), Angel et al. (1954 b): Mt.

Nimba (as D. e/ongata); Ineich (2003): Mt. Nimba.

— Dipsadoboa weileri — Angel et al. (1954 b): Mt. Nimba (as D.

unicolor (part)); Ineich (2003): Mt. Nimba.

— Dispholidus typus — Chabanaud (1921): N’Zeébéla; Villiers

(1950): Dalaba; Ineich (2003): Mt. Nimba; Trape & Mané (2006

b): Kindia.

— Hapsidophrys smaragdina — Chabanaud (1921): Macenta,

N’Zérékoré (as Gastropyxis smaragdina) Condamin (1959):

Sérédou (as G. smaragdina); Bohme (2000): Ziama Forest (as

G. smaragdina); this paper.

— Hapsidophrys lineata — Chabanaud (1921): N’Zébéla, Diécké;

Angel et al. (1954 b), Bohme (2000): Ziama Forest; Ineich

(2003): Mt. Nimba; this paper.

— Meizodon coronatus — Mocquard (1902): Koroussa (als Coro-

nella coronata); Villiers (1950): Dalaba; Condamin (1959): Séré-

dou; Roux-Estéve (1969): Sambailo, Kouroussa; BOhme (2000):

N’Zérékoré; Greenbaum & Carr (2005): PN Haut Niger.

— Meizodon regularis — Angel et al. (1954 b) Mt. Nimba (as M.

coronatus (? in part); Roux-Estéve (1969): Kérouané, Beyla,

Fouta Djallon, Dalaba, Coyah, Sérédou, Mt. Nimba; Bo6hme

(2000): Ziama Forest; Ineich (2003): Mt. Nimba; this paper.

— Philothamnus carinatus — Villiers (1954), Angel et al. (1954

b): Mt. Nimba (as P. heterodermus carinatus); Bohme (2000):

Ziama Forest; Ineich (2003): Mt. Nimba; this paper.

— Philothamnus heterodermus — Chabanaud (1921): N’Zébéla

(as Chlorophis heterodermus); Villiers (1950): Mt. Nimba (as

C. heterodermus); Villiers (1954), Angel et al. (1954): Mt. Nim-

ba (as P. h. heterodermus); Greenbaum & Carr (2005): PN Haut

Niger; Rédel & Bangoura (2006): Mt. Béro; this paper.

— Philothamnus irregularis — Villiers (1954), Angel et al. (1954

b): Mt. Nimba (as P. i. irregularis); Ineich (2003): Mt. Nimba;

Rodel & Bangoura (2004b): Pic de Fon/Simandou range; Green-

baum & Carr (2005): PN Haut Niger; this paper.

— Philothamnus semivariegatus — Klaptocz (1913): Dubreka;

Condamin (1959): Sérédou.

— Telescopus variegatus — Greenbaum & Carr (2005): PN Haut

Niger.

— Thelotornis kirtlandii — Villiers (1950), Angel et al. (1954 b),

Ineich (2003): Mt. Nimba; Riquier & B6hme (1996): Ziama For-

est; Ineich (2003): Mt. Nimba; Rédel & Bangoura (2006):

Diécké Forest; this paper.

— Thrasops occidentalis — Villiers (1950): Dalaba; Bohme

(2000): Ziama Forest; Ineich (2003): Mt. Nimba; Trape & Mané

(2006): N’Zérékoré; this paper.

— Toxicodryas blandingii — Chabanaud (1921): Macenta,

N’Zébéla, Diécké Forest (as Dipsadomorphus blandingii) Vil-

liers (1954), Angel et al. (1954 b): Mt. Nimba (as Boiga

blandingii; Condamin (1959): Sérédou (as B. blandingii); Bohme

(2000): Ziama Forest; Ineich (2003): Mt. Nimba (as B.

blandingii); Greenbaum & Carr (2005): PN Haut Niger; Trape

& Mané (2006): N’Zérékoré; this paper; Mt. Nimba, this paper.

Bonn zoological Bulletin 60 (1): 35—61

— Toxicodryas pulverulentus — Chabanaud (1921): Macenta (as

Dipsadomorphus pulverulentus); Villiers (1954), Angel et al.

(1954 b), Ineich (2003): Diécké Forest; Mt. Nimba (as Boiga

pulverulenta),; this paper: east of Guéckédou, Ziama Forest.

Natricidae

— Afronatrix anoscopus — Boulenger (1910): Labé (as Helicops

gendrii sp. n.); Klaptocz (1913): Mamou (as Tropidonotus fer-

ox); Chabanaud (1921): Macenta, N’Zébéla, N’Zérékoré (as T.

ferox); Villiers (1950): Dalaba, Mt. Nimba (as Natrix anosco-

pus gendrii); Villiers (1954): Mt. Nimba (as N. anoscopus); Con-

damin (1959): Sérédou (as N. anoscopus); Bbhme (2000): Zia-

ma and Diécké Forests; Ineich (2003: Mt. Nimba); Rédel & Ban-

goura (2004b): Pic de Fon/Simandou range; Rédel & Bangoura

(2006): Diécké Forest; this paper; Pic de Fon/Simandou range

(Rodel & Bangoura 2004b), this paper.

— Amblyodipsas unicolor — Trape & Mané (2006): on grid map

but without specific locality.

— Grayia smithii — Klaptocz (1913): Dabola; Chabanaud

(1921): N’Zébéla. N’Zérékoré; Villiers (1950): Mt. Nimba; Con-

damin (1959): Sérédou; BOhme (2000): Ziama Forest; Ineich

(2003): Mt. Nimba; Greenbaum & Carr (2005): PN Haut Niger;

Trape & Mané (2005): Djani River; this paper.

— Grayia tholloni — Trape & Mané (2006 b): on grid map but

without specific locality.

— Natriciteres variegata —Mocquard (1902): Kouroussa (als Mi-

zodon variegatus); Chabanaud (1921): Macenta, N’Zébéla,

N’Zérékoré (as Tropidonotus variegatus); Villiers (1950, 1954),

Angel et al. (1954 b): Mt. Nimba (as Neusterophis variegatus);

Condamin (1959): Sérédou (as Neusterophis variegatus);

Bohme (2000): Ziama Forest; Ineich (2003): Mt. Nimba; Green-

baum & Carr (2005) PN Haut Niger; Trape & Mané (2006):

N’Zérékoreé.

Psammophiidae

—Psammophis lineatus — Villiers (1950): Dalaba; Trape & Mané

(2006 b): Kindia.

— Psammophis praeornatus — Mocquard (1902): Kouroussa;

Greenbaum & Carr (2005): PN Haut Niger.

— Psammophis elegans — Greenbaum & Carr (2005): PN Haut

Niger.

— Psammophis cf. phillipsi — Mocquard (1902): Kouroussa (as

P. sibilans); Klaptocz (1913): Dubreka (as P. sibilans); Cha-

banaud (1921): Macenta, N’Zébéla, N’Zérékoré, Diécké Forest

(as P. sibilans); Villiers (1954), Angel et al. (1954 b): Mt. Nim-

ba (as P. sibilans phillipsii); Brandstatter (1996), Ineich (2003):

Mt. Nimba; this paper (Kissidougou).

— Psammophis phillipsi — Villiers (1950, 1954), Angel et al. (1954

b): Mt. Nimba (as P. sibilans phillipsii); Condamin (1950): Séré-

dou (as P. sibilans phillipsii); this paper.

— Rhamphiophis oxyrhynchus — Trape & Mané (2006): on grid

map, but without specific locality.

©ZFMK

60 Wolfgang Bohme et al.

Atractaspididae

— Aparallactus lineatus — Villiers (1950, 1954), Angel et al. (1954

b), Wallach (1994): Mt. Nimba; Bohme (2000): Ziama Forest;

Ineich (2003): Mt. Nimba; this paper: Ziama Forest.

— Aparallactus modestus — Condamin (1959): Sérédou; Ineich

(2003): Mt. Nimba; Rédel & Bangoura (2006): Diécké; this pa-

per.

— Aparallactus niger — Chabanaud (1917): Sampouyara (as

Rouleophis chevalieri); Chabanaud (1921): Kérouane, Beyla,

N’Zérékore, Diécké Forest; Villiers (1950): Dalaba; Angel et al.

(1954 b): Mt. Nimba; Condamin (1959): Sérédou; Wallach

(1994): Mt. Nimba; Ineich (2003): Mt. Nimba; this paper.

— Atractaspis irregularis — Condamin (1959): Sérédou; this pa-

per.

— Atractaspis aterrima — Mocquard (1906): Conakry; Chabanaud

(1921): Dixine; Rasmussen (2005): Mt. Nimba; Trape & Mané

(2006 b): on grid map but without specific localities. Not listed

by Ineich (2003).

— Polemon acanthias — Villiers (1954), Angel et al. (1954 b): Mt.

Nimba (as Miodon acanthias); Condamin (1959): Sérédou (as

M. acanthias); Ineich (2003): Mt. Nimba; this paper: Ziama For-

est.

— Polemon bocourti — Angel et al. (1954): Mt. Nimba (but not

mentioned by Ineich (2003).

— Prosymna greigerti — Trape & Mané (2006): on grid map but

without specific locality.

— Prosymna meleagris — Trape & Mané (2006): on grid map but

without specific locality.

Elapidae

— Dendroaspis polylepis — Greenbaum et al. (2003): PN Haut

Niger; Trape & Mané (2006): Kindia.

— Dendroaspis viridis — Klaptocz (1913): Konkouré; Chabanauid

(1921): N’Zébéla, N’Zérékore; Villiers (1950), Angel et al. (1954

b): Mt. Nimba; Condamin (1959): Sérédou; B6hme (2000): Zia-

ma Forest; Ineich (2003): Mt. Nimba; Greenbaum et Carr (2005):

PN Haut Niger; Trape & Mané (2006): Kindia; this paper.

— Elapsoidea semiannulata — Greenbaum & Carr (2005): PN

Haut Niger.

— Elapsoidea trapei — Trape & Mané (2006): on grid map but

without specific locality. — Note: We are not yet fully convinced

of the taxonomic distinctness of this species.

— Naja haje — Trape & Mané (2006): on grid map but without

specific locality.

— Naja katiensis — Trape & Mané (2006): on grid map but with-

out specific locality.

— Naja melanoleuca — Chabanaud (1921): Macenta, N’Zébela,

N’Zérékoré; Villiers (1950), Angel et al. (1954 b): Diécké For-

est, Mt. Nimba; Condamin (1959), Bohme (2000): Sérédou; In-

eich (2003): Diécké Forest, Mt. Nimba; Rédel & Bangoura

2004b): Pic de Fon/Simandou range; Rédel & Bangoura

(2006): Diécké Forest; Trape & Mané (2006): Kindia; this pa-

per.

Bonn zoological Bulletin 60 (1): 35-61

— Naja nigricollis — Mocquard (1902): Kouroussa; Villiers

(1950), Angel et al. (1954 b): Mt. Nimba; Condamin (1959):

Sérédou; B6hme (2000): Ziama Forest; Ineich (2003): Mt. Nim-

ba; Rédel & Bangoura (2004b): Pic de Fon/Simandou range;

Greenbaum & Carr (2005): PN Haut Niger; Rodel & Bangoura

(2006): Mt. Béro; this paper.

— Pseudohaje nigra — Roman (1976): N’Zérékoré; Bohme

(2000): Diécké Forest; this paper.

Viperidae

— Atheris chlorechis — Chabanaud (1921): Macenta; Villiers

(1950), Angel et al. (1954 b): Mt. Nimba; Condamin (1959):

Sérédou; B6hme (2000): Ziama Forest; Ineich (2003): Mt. Nim-

ba; Rédel & Bangoura (2006): Déré Forest; this paper: Mt. Nim-

ba, photo record C. Brede.

— Bitis arietans — Klaptocz (1913): “Kampement am Niger”; R6-

del & Bangoura (2004b): Pic de Fon/Simandou range; Green-

baum & Carr (2005): PN Haut Niger; Hillers et al. (2006): Rio

Kapatchez, Kamsar; this paper: Kissidougou.

— Bitis nasicornis — Chabanaud (1921): Macenta, N’Zébéla,

N’Zérékoreé; Villiers (1954), Angel et al. (1954 b): Mt. Nimba;

Condamin (1959): Sérédou; BOhme (2000): Ziama and Diécké

Forests, this paper; Ineich (2003): Mt. Nimba; this paper.

— Bitis rhinoceros — Chabanaud (1921): Macenta, N’Zérékoré,

Diécké Forest (as B. gabonica); Villiers (1950): Dalaba (as B.

gabonica); Villiers (1954), Angel et al. (1954 b): Mt. Nimba (as

B. gabonica); Condamin (1959): Sérédou (as B. g. rhinoceros);

Greenbaum & Carr (2005): PN Haut Niger (as B. gabonica);

Bohme (2000): between Macenta and Sérédou, Diécké Forest

(as B. gabonica); Ineich (2003): Mt. Nimba; Rédel & Bangoura

(2004b): Pic de Fon/Simandou range (as B. gabonica); R6édel

& Bangoura (2006): Fouta Dyjallon, Labé/Daralabe/Forét Kok-

oulo; this paper.

— Causus lichtensteinii — Condamin (1959): Sérédou; Ineich

(2003): Mt. Nimba.

— Causus maculatus — Mocquard (1902): Kouroussa (as C.

rhombeatus); Chabanaud (1921): Macenta, N’Zébéla; N’Zéréko-

ré, Diécké Forest (as C. rhombeatus); Villiers (1950): Dalaba,

Mamou, Mt. Nimba (as C. rhombeatus); Angel et al. (1954 b):

Mt. Nimba (as C. rhombeatus); Bohme (2000): Ziama and

Diécké Forests; Ineich (2003): Mt. Nimba; Greenbaum & Carr

(2005): PN Haut Niger; Hillers et al. (2006): Kolaboui; Rédel

& Bangoura (2006): Deéré Forest; Trape & Mané (2006):

N’Zérékoré; this paper.

— Echis ocellatus — Trape & Mané (2006): on grid map, but with-

out specific locality.

Crocodylidae

— Crocodylus suchus — Klaptocz (1913): Mamou, Konkouré N

of Dubreka (as C. niloticus) Greenbaum & Carr (2005): PN Haut

Niger (as C. niloticus).

— Mecistops cataphractus — Klaptocz (1913): Baffing (reported

to him as “Krokodil mit spitzer Schnauze”; Chabanaud (1921):

Kérouané (as Crocodylus ctataphractus; Greenbaum & Carr

(2005): PN Haut Niger (as C. cataphractus).

— Osteolaemus tetraspis — Chabanaud (1921): Keérouané,

N’Zébéla; R6del & Bangoura (2004b): Pic de Fon/Simandou

range; Rédel & Bangoura (2006): Diécké Forest; this paper.

©ZFMK

Amphibians and reptiles from Guinea 61

Appendix II

List of the fish species collected in Ziama Forest

Along with the few fish specimens recovered from the stomachs

of the (semi)aquatic snake species Afronatrix anoscopus and

Grayia smithii (see above), some more fishes were collected on

the occasion of the herpetological survey. They have been iden-

tified by Dr. Jorg Freyhof (formerly ZFMK Bonn, now Berlin)

and are deposited in the ZFMK ichthyological collection:

Mormyridae

Petrocephalus tenuicaudatus (Steindachner, 1899)

Citharinidae

Neolebias unifasciatus Steindachner, 1894

Cyprinidae

Barbus cf. traorei Léveque, Tengels & Thys, 1987

Kribia cf. nana (aus Afronatrix anoscopus ZFMK 56076)

Schilbeidae

Schilbe mandibularis (Gunther, 1867)

Amphiliidae

Amphilius rheophilus Daget, 1959

Bonn zoological Bulletin 60 (1): 35-61

Clariidae

Heterobranchus longifilis Valenciennes, 1840

Mochokidae (Synodontidae)

Chiloglanis occidentalis Pellegrin, 1933

Synodontus sp. (cf. comoensis?)

Aplocheilidae

Epiplatys olbrechtsi Poll, 1941

Epiplatys lamottei Daget, 1954

Cyprinodontidae

Aplocheilichthys schioetzi Scheel, 1968 (aus Afronatrix anosco-

pus ZFMK 56066 and 56275)

Aplocheilichthys normani Ahl, 1928

Cichlidae

Tilapia (Coptodon) sp. n. ? (aus Grayia smithiti ZFMK 56036)

Mastacembelidae

Aethiomastacembelus liberiensis (Boulenger, 1898).

Received: 25.01.2011

Accepted: 01.03.2011

Corresponding editor: F. Herder

©ZFMK

Volume 60

Bonn zoological Bulletin |

| Issue | | pp. 63-65 Bonn, May 2011

A new record of the Persian Brook Salamander,

Paradactylodon persicus (Eiselt & Steiner, 1970)

(Amphibia: Caudata: Hynobiidae)

in northern Iran

Faraham Ahmadzadeh!:2*, Fatemeh Khanjani3, Aref Shadkam4 & Wolfgang Bohme?

'Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute,

Shahid Beheshti University, Evin, Tehran, G. C., [ran

*Herpetology Section, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK),

Adenauerallee 160, D-53113 Bonn, Germany

3Department of Habitat and Biodiversity, Faculty of Environment and Energy, Science and Research Branch,

Islamic Azad University, Ponak, Tehran, Iran

4Department of Environment, Rezvan Shahr, Province Gilan, [ran

“Corresponding Email address: Fahmadza@uni-bonn.de.

INTRODUCTION

The Persian Brook Salamander, Paradactylodon persicus

(Eiselt & Steiner, 1970) is an endemic and poorly known

species of northern Iran (Baloutch & Kami 1995; Kami

1999). It was originally described as Batrachuperus per-

sicus by Eiselt & Steiner (1970), but has been transferred

to the genus Paradactylodon based on genetic studies by

Zhang et al. (2006). This species has been reported from

two localities only: Weyser, southeast of Chalus, in

Mazandaran Province (36° 30° 35” N and 51° 26’ 38” E)

and Delmadeh village, southeast of Khalkhal, in Ardabil

Province (37° 22’ 34” N and 48° 47’ 35” E) (Kami 2004;

Ebrahimi et al., 2004) (Fig. 1). Here, we report a new

record of this species in Hyrcanian forest, in Gilan

Province, Iran.

RESULTS

During our field survey in June 2010, a single specimen

was collected in Dasht-e-Daman Yeylagi, in Rezvan Shahr

city of Gilan Province (37° 27’ 53” N and 48° 47° 10” E)

at an elevation of 1622 ma. s. I. (Fig. 1). This specimen

is a subadult (Fig. 2a—b) and it was identified as

Paradactylodon persicus based on external morphologi-

cal characters which mentioned by Baloutch & Kami

(1995). Because this is a threatened species in Iran (Pa-

penfuss et al. 2011; Ahmadzadeh & Kami 2009), it was

released into the natural habitat after morphological ex-

amination. The characteristic features of this specimen are

as the following: total length 23 mm; snout-vent length

90 mm; tail length 120 mm; head large, 20 mm in length;

vomerine teeth in two arch-shaped rows; snout rounded;

fore and hind limbs with four digits; tail flattened later-

ally, with round-tapered end; dorsal head and body, as well

as upper surface of tail brownish with yellow spots and

marblings; belly cream without pattern (Fig. 2a—b).

Paradactylodon persicus inhabits the mountainous

streams and brooks, with cool, fast-flowing water

(Baloutch & Kami 1995; Kami 1999; Ahmadzadeh & Ka-

mi 2009). Like the Delmadeh habitat in Ardabil Province,

the habitat reported here is also located in a non-forestry

area (Ahmadzadeh & Kami 2009). The specimen was

found under a stone close by a stream on the steep slope

of Alborz Mountains, exposed to the Caspian Sea.

REFERENCES

Ahmadzadeh F & Kami HG (2009) Distribution and conserva-

tion status of the Persian Brook Salamander, Batrachuperus

(Paradactylodon) persicus (Amphibia: Caudata: Hynobiidae)

in north-western Iran. Iranian Journal of Animal Biosystem-

atics (IJAB) 5: 9-15

Baloutch M & Kami HG (1995) Amphibians of Iran. Tehran uni-

versity Publication, Tehran (in Persian): 177 pp.

Ebrahimi M, Kami HG, Stéck M (2004) First description of egg

sacs and early larval development in hynobiid salamanders

(Urodela, Hynobiidae, Batrachuperus) from north-eastern

Iran. Asiatic Herpetological Research 10: 168-175

Eiselt J, Steiner HM (1970) Erstfund eins hynobiiden Molches

in Iran. Annalen des Naturhistorischen Museum Wien 74:

77-90

64 Faraham Ahmadzadeh et al.

Caspian Sea

Dasht-e-Daman

New Locality

Kordestan

Semnan

Hamedan

mani’

48° 5

Fig. 1. | Map showing the two former valid localities and new reported locality of Paradactylodon persicus in Iran.

Kami HG (1999) Additional specimens of the Persian Mountain in Golestan Province, Iran. Asiatic Herpetological Resaerch

Salamander, Batrachuperus persicus, from Iran (Amphibia : 10: 182-190

Hynobiidae). Zoology in the Middle East 19: 37-42 Papenfuss T, Anderson S, Kuzmin S, Rastegar-Pouyani N (2010)

Kami HG (2004) The biology of the Persian Mountain Salaman- Batrachuperus persicus. In: IUCN Red List of Threatened

der, Batrachuperus persicus (Amphibia, Caudata, Hynobidae) Species, online at www.iucnredlist.org, accessed on January

20, 2011

Fig. 2a—b. — Paradactylodon persicus from Dasht-e-Daman Yeylagi in Rezvan Shahr city of Gilan Province.

Bonn zoological Bulletin 60 (1): 63-65 ©OZFMK

A new record of the Persian Brook Salamander, Paradactylodon persicus in northern Iran 65

Fig. 3a—b. The newly recorded habitat of Paradactylodon persicus in the Dasht-e-Daman Yeylagi in Rezvan Shahr city of Gi-

lan Province.

Zhang P, Yue-Qin C, Hui Z, Yi-Fei L, Xiu-Ling Wg, Theodore — Received: 12.10.2010

JP, David Wake B,Liang-Hu Q (2006) Phylogeny, evolution, WNecepted: 02.03.2011

and biogeography of Asiatic Salamanders (Hynobiidae). Pro- oa eRe

ceedings of the National Academy of Sciences of the United

States of America 103: 7360-7365

Corresponding editor: F. Herder

Bonn zoological Bulletin 60 (1): 63-65 ©ZFMK

Bonn zoological Bulletin Volume 60 Issue | pp. 67-71 | Bonn, May 2011

Type list of amphibians and reptiles

in the Zoologisches Forschungsmuseum A. Koenig, Bonn:

corrections and additions

Wolfgang Bohme

Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany;

E-mail: w.boehme.zfmk(@uni-bonn.de.

The last issue of this journal contained type catalogues of

the vertebrate collections in the Zoologisches

Forschungsmuseum A. Koenig (ZFMK) in Bonn, among

them that of the amphibians and reptiles. It was already

earlier and still is my plan to publish an extensive, com-

prehensive and critically commented herpetological type

catalogue after my nearly four decades of curatorship at

ZFMK, together with a new, illustrated overview of the

history of the Bonn collection, of all herpetological activ-

ities in Bonn at ZFMK and the time before (B6hme in

prep. 2011). Therefore I hesitated to take also part in the

joint catalogue project of the museum, since I was afraid

of redundancies between both publication projects. I be-

came, however, convinced that herpetology should not be

missing among the catalogues of mammals, birds and fish-

es, and so I decided for a compromise which could be ac-

complished in a relatively short time without delaying the

time schedule of the entire issue too much: A reduced list

giving only

—name, author and year of description;

— catalogue numbers only for name bearers in ZFMK;; for

name bearers in other collections and for typoids only the

acronym of the respective collections;

— country of origin and current name if the original name

had changed since the original description, either by dif-

ferent generic assignments or by synonymizations. The

reader was already referred to the extensive catalogue with

all specimen data including figures to follow in 2011

(Bohme 2010: Bonn zoological Bulletin 59: 79-108).

Unfortunately, the time pressure which arose from the

above-mentioned circumstances led to some errors and

misprints which escaped our attention during proofread-

ing. Such mistakes are particularly annoying when scien-

tific names are concerned because a misspelled name 1s

a different combination of letters and thus a different

(mostly nude) name in the sense of the Code (ICZN 1999),

and the need to correct them is obvious. I prefer to pro-

vide such an errata list myself before it has to or will be

done by collegues, but I will not consider matters which

are dependant from opinions (e.g. assignments to supra-

generic categories etc.). Rather, I shall correct only those

mistakes (misspellings, omissions) which may affect

nomenclature, authorships, dates of publication, or places

of deposition. For the institutional abbreviations the read-

er is referred to the first type list cited above.

In order to make this paper a little bit more than just an

errata sheet for the published type list, I take also the op-

portunity to add several taxa recently published and not

yet contained in the list. These are marked in bold, as are

the corrections (followed by the erroneous names, words

or acronyms in parentheses; otherwise they are additions).

Corrections are listed in the order of taxa as published in

the type list. The newly described taxa are —in bold — in-

cluded in this systematic order. New, further acronyms are

AMS: Australian Museum, Sydney; CRES: Center for

Natural Resources and Environmental Studies, Hanoi (=

VNUH) ; NCSM: North Carolina Museum of Natural Sci-

ences, Raleigh; and SNHM-BS: State Natural History Mu-

seum, Braunschweig.

AMPHIBIA

Pelodryadidae (not Pelodrydaidae)

Hemiphractidae (not Amphignathodontidae)

Gastrotheca piperata Duellman & J. Kohler, 2005

Holotype: ZFMK 66838, paratypes in ZFMK and KU

Origin: Bolivia

68 Wolfgang Bohme

Hylidae

Osteocephalus oophagus Jungfer & Schiesari, 1995

Paratypes in ZFMK, NMW, MPEG, AMNH, MZUSP, IN-

PA and SMNS, holotype in MZUSP

Centrolenidae

Hyalinobatrachium carlesvilai Castroviejo-Fisher, Padi-

al, Chaparro, Aguayo & De la Riva, 2009

Paratypes in ZFMK, CBG, MHNC and MNHN, holotype

in MHNCP (not in MNHN)

Origin: Bolivia

Strabomantidae

Eleutherodactylus olivaceus J. Kohler, Morales, Lotters,

Reichle & Aparicio, 1998

Paratypes in ZFMK, CBF, MUSM and USNM, holotype

in CBF

Origin: Bolivia

Current name: Pristimantis olivaceus

Pristimantis koehleri Padial & De la Riva, 2009

Paratypes in ZFMK, MNCN and NMK, holotype in NKA

(not MNCN)

Origin: Bolivia

Pristimantis reichlei Padial & De la Riva, 2009

Paratypes in ZFMK, CBG, KU, MNCN, NMP6V,

NMW, USNM and NMK, holotype in NKA

Origin: Bolivia

Leptodactylidae

Adenomera griseigularis Henle, 1981

Holotype: ZFMK 31800, no paratypes

Origin: Peru

Current name Leptodactylus griseigularis (not griseono-

tus)

Leptodactylus raniformis Werner, 1899

Holotype: ZFMK 28484 (not 80600), no paratypes

Origin: Colombia

Current name: Leptodactylus fuscus.

Bonn zoological Bulletin 60 (1): 67-71

Ceratophryidae

Telmatobius mayoloi Salas & Sinsch, 1996

Paratypes in ZFMK, URP, MHNSM and KU, holotype in

URP

Origin: Peru (not Bolivia)

Dendrobatidae

Dendrobates typographus Keferstein, 1867

Holotype: ZFMK 28115, no paratypes

Origin: Costa Rica

Current name: Oophaga (not -gus) pumilio

Epipedobates bilinguis Jungfer, 1989

Holotype: ZFMK49073, paratypes in ZFMK

Origin: Ecuador

Current name: Ameerega (not Amereega) bilinguis.

Bufonidae

Atelopus cruciger vogli L. Miiller, 1935

Paratypes in ZFMK, HLMD and ZSM, holotype in ZSM.

Origin: Venezuela

Current name: Afelopus vogli

Atelopus flaviventris Werner, 1899

Holotype: ZFMK 28107, no paratypes

Origin: Colombia

Current name: Afelopus subornatus (not ignescens)

Bufo scorteccii Balletto & Cherchi, 1970

Paratypes in ZFMK and IZUG, holotype in IZUG

Origin: Yemen

Current name: Duttaphrynus scorteccii

Werneria submontana Rédel, Schmitz, Pauwels, & Boh-

me, 2004

Holotype: ZFMK 69999, paratypes in ZFMK

Origin: Cameroon

Microhylidae

Plethodontohyla mihanika (not mikanika) Vences, Rax-

worthy, Nussbaum & Glaw, 2003

Paratypes in ZFMK, ZSM, UMMZ and MNHN, holoty-

pe in UMMZ

Origin: Madagascar

©ZFMK

Type list of the amphibians and reptiles in the ZFMK, Bonn: corrections and additions 69

Hyperoliidae

Heterixalus andrakata (not mandrakata) Glaw & Vences,

1991

Holotype: ZFMK 52557, paratypes in ZFMK and ZSM

Origin: Madagascar

Arthroleptidae

Leptodactylodon wildi Amiet (not Amit) & Dowsett-

Lemaire, 2000

Paratypes ZFMK, MNHN and BMNH, holotype in

MNHN

Origin: Cameroon

Phrynobatrachus nlonakoensis Plath, Herrmann &

Bohme, 2006

Holotype: ZFMK 80970, paratypes in ZFMK

Origin: Cameroon

Current name: Arthroleptis nlonakoensis

Phrynobatrachidae

Phrynobatrachus kakamikro Schick, Zimkus, Channing,

J. Kohler & Lotters, 2010

Paratypes in ZFMK and NMK (not in BMNH and NKM),

holotype in NMK (not NKM)

Origin: W Kenya

Mantellidae

Blommersia galani Vences, J. Kohler, Pabijan & Glaw,

2010

Paratypes: in ZFMK, ZMA and ZSM, holotype in ZSM

Origin: Madagascar

Current name: Boophis occidentalis

Boophis blommersae Glaw & Vences 1994 (not 1995)

Holotype: ZFMK 57398, paratypes in ZFMK and ZSM

Origin: Madagascar

Boophis haingana Glaw, J. Kohler, De la Riva, Vietes &

Vences, 2010

Paratypes in ZFMK, ZSM and UADBA, holotype in ZSM

Origin: Madagascar

Boophis luciae Glaw, J. Kohler, De la Riva, Vieites &

Vences, 2010

Bonn zoological Bulletin 60 (1): 67-71

Paratypes in ZFMK, ZSM and UADBA, holotype in ZSM

Origin: Madagascar

Boophis quasiboehmei Vences, J. Kohler, Crottini &

Glaw, 2010

Paratypes in ZFMK and ZSM, holotype in ZSM

Origin: Madagascar

Boophis tampoka J. Kohler, Glaw & Vences, 2007

Paratypes in ZFMK, HLMD, UADBA and ZSM (not in

FGZC), holotype in ZSM

Origin: Madagascar

Mantella expectata Busse & Bohme, 1992

Holotype: ZFMK, paratypes in ZFMK and ZSM

Origin: Madagascar

Mantidactylus corvus Glaw & Vences, 1994 (not 1992)

Holotype: ZFMK 57430, paratypes in ZFMK and ZSM

Origin: Madagascar

Current name: Gephyromantis (Phylacomantis) corvus

Rhacophoridae

Philautus quyeti Nguyen Q.T., Hendrix, BOhme, Vu &

Ziegler, 2008

Holotype: ZFMK 82999, paratype in VNUH

Origin: Vietnam

Current name: Gracixalus quyeti

Rhacophorus orlovi Ziegler & J.Kohler, 2001

Holotype: ZFMK 71368, paratypes in ZFMK (not ZM-

FK), MTKD, CRES and ZMB

Origin: Vietnam

Rhacophorus vampyrus Rowley, Le, Tran, Stuart &

Hoang, 2010

Paratypes in ZFMK, AMS, UNS, NCSM, holotype in

AMS

Origin: Vietnam

Ranidae

Rana batcanica H. Schneider, Sinsch & Sofianidou, 1993

Holotype: ZFMK 52825, paratypes in ZFMK

Origin: Greece

Current name: Pelophylax kurtmuelleri (not kurtklari)

©ZFMK

70 Wolfgang Bohme

Agamidae

Agama pakistanica Baig, 1989

Paratypes in ZFMK and PMNH, holotype in PMNH

Origin: N Pakistan

Current name: Laudakia pakistanica

Calotes nigriplicatus Hallermann, 2001 (‘2000’)

Holotype: ZFMK 26379, no paratypes

Origin: Moluccas, Indonesia

Pseudocalotes ziegleri Hallermann, Nguyen Q.T., Orlov

& Ananjeva, 2010

Paratypes in ZFMK, IEBR and ZISP, holotype in IEBR

Origin: Vietnam

Chamaeleonidae

Bradypodion tavetanum boehmei Lutzmann & NeCas,

2002

Holotype: ZFMK 63373, paratypes in ZFMK

Origin: Kenya

Current name: Kinyongia boehmei

Chamaeleo marsabitensis Tilbury, 1991

Paratypes in ZFMK and BMNH, holotype in BMNH

Origin: N Kenya

Current name: 7rioceros marsabitensis.

Kinyongia uthmoelleri artytor Lutzmann, Stipala,

Lademann, Krause, Wilms & Schmitz, 2010

Paratypes in ZFMK and MHNG, holotype in MHNG

Origin: Tanzania

Phyllodactylidae

Gecko fascicularis Daudin, 1802

Neotype: ZFMK 35631

Origin: Libya

Current name: Tarentola f. fascicularis (not T. mauritan-

ica fascicularis)

Tarentola fascicularis wolfgangi Joger & Bshaenia,

2010

Paratypes in ZFMK, HLMD and SNHM-BS, holotype

in SNHM-BS

Origin: Tunisia

Bonn zoological Bulletin 60 (1): 67-71

Sphaerodactylidae

Pristurus obsti Résler & Wranik, 1999

Paratypes in ZFMK, MTKD, HLMD and pers. coll. H.R.,

holotype in MTKD

Origin: Socotra Id., Yemen

Gekkonidae

Cyrtodactylus cattienensis Geibler, Nazarov, Orlov, Boh-

me, Phung (not Trung), Nguyen Q.T. & Ziegler, 2009

Paratypes in ZFMK IEBR, VNUH, ZMMU and ZISP, ho-

lotype in IEBR

Origin: Vietnam

Cyrtodactylus phongnhakebangensis Ziegler, Résler,

H.W. Herrmann & Vu (not Thanh), 2003 (“2002”)

Holotype: ZFMK 76193, paratypes in ZFMK, VNUH and

MTKD

Origin: Vietnam

Gekko siamensis Grossmann & Ulber, 1990

Paratypes in ZFMK, MTKD and pers. coll. W.G., holo-

type in MTKD

Origin: Thailand

Gekko scientiadventura Rosler, Ziegler, Vu (not Thanh),

H.W. Herrmann & Bohme (2005 (“2004’’)

Holotype: ZFMK 76198, paratypes in ZFMK and VNUH

Origin: Vietnam

Hemiphyllodactylus titiwangsaensis Zug, 2010

Paratypes in ZFMK and ZRC, holotype in ZRC

Origin: Malaysia

Pachydactylus capensis katanganus De Witte, 1953

Paratypes in ZFMK and IRSNB, holotype in IRSNB

Origin: D.R. Congo

Lacertidae

Lacerta cappadocica muhtari Eiselt, 1979

Paratypes in ZFMK, SZE, NMW, ZSM, ZMH, BMNH,

FMNH and CAS, holotype in NMW

Origin: Turkey

Current name Apathya cappadocica muhtari.

©ZFMK

Type list of the amphibians and reptiles in the ZFMK, Bonn: corrections and additions 71

Scincidae

Ablepharus chernovi eiselti Schmidtler, 1997

Paratypes in ZFMK, ZSM, ZDEU and NMW, holotype

in ZSM

Origin: S Turkey

Varanidae

Varanus rasmusseni Koch, Gaulke & Boéhme, 2010

Paratype in ZFMK, holotype in ZMUC

Origin: Tawi Tawi Id. (not UID.), Sulu Ids., Indonesia

Calamariidae

Calamaria thanhi (not thani) Ziegler & Le, 2005

Holotype: ZFMK 82920, no paratypes

Origin: Vietnam

Bonn zoological Bulletin 60 (1): 67-71

Colubridae

Lycodon ruhstrati abditus Vogel, David, Pauwels, Sumon-

tha, Norval, Hendrix, Vu & Ziegler, 2009

Holotype: ZFMK 86451, paratypes in ZFMK, MNHN and

ZMB

Origin: Vietnam

Viperidae: Crotalinae

Trimeresurus nebularis Vogel, David & Pauwels, 2001

Paratypes in ZFMK, IRSNB, MNHN, ZRC and Coll. G.

Vogel, holotype in USNM

Origin: Thailand

Acknowledgements. Thanks are due to Frank Glaw, André

Koch, Joérn Kohler and Nguyen Quang Truong for bringing most

of the errors to my attention.

Received: 02.02.2011

Accepted: 21.02.2011

Corresponding editor: F. Herder

©ZFMK

WD Book Review

Paulsen MJ (2010) The Stag Beetles of Southern South

America (Coleoptera: Lucanidae). Bulletin of the Univer-

sity of Nebraska State Museum Vol. 24: 148pp. 130 illus.

+ 29 maps. Gail Littrell, Publications Secretary, W436 Ne-

braska Hall, University of Nebraska, Lincoln, NE 68588-

0514, U.S.A. E-mail: littrell@unlserve.unl.edu. FAX:

(402) 472-8949.$40.

Matt’s book is a monographic compilation of original and

recent research on southern South American stag beetles

(Coleoptera: Scarabaeoidea: Lucanidae) which includes

their taxonomic treatment, the study of their distribution

but involves also the phylogenetic analysis of a selected

genus, Pycnosiphorus. It was part of a research project that

funded by the National Science Foundation (NSF)-PEET

(Partnerships for Enhancing Expertise in Taxonomy; e.g.

http://www-museum.unl.edu/research/entomology/

PEET2Summary.htm) and which have produced a series

of monographs and other taxonomic works but also attrac-

tive web sites. The endemic stag beetle fauna of the study

area includes 31 species in three subfamilies. The author

provides keys to species and genera of southern South

American Lucanidae and updates information on genera

that have been recently revised. All species are redescribed

and keys are provided in English and Spanish when ap-

propriate. The high quality images of male genitalia and

habitus of species makes the book a MUST for South

American Coleopterists but also aesthetically very nice.

Distribution information presented by detailed maps is

completed also by the knowledge of species phenology.

In the context of the taxonomic treatment he discusses the

systematic position of the monogeneric tribe Streptoceri-

ni Kikuta, 1986 which he placed into synonymy with Lam-

primini.

The monograph concludes with an interesting appendix

which represents a first molecular study of southern South

American Lucanidae, namely on the genus Pycnosipho-

rus, using DNA of the 28s D2 region and ITS2 with a

number of ca. 1500 base pairs. Although the sampling of

the group was complete the author dedicated only little

space for this analysis in regard of methodological details

but also the discussion of the interesting results which

Bonn zoological Bulletin 60 (1): 72

The Stag Beetles of

Southern South America

S/S

Volume 24 - 2

would have been very exciting also for a more generally

interested reader. In the meantime the produced sequence

data were submitted to Genbank and though they can be

extended or used in future molecular work by colleagues

what makes the work are an even more valuable piece in

the mosaic of beetle biodiversity exploration.

Dirk Ahrens

Zoologisches Forschungsmuseum Alexander Koenig,

Bonn

©ZFMK

Bonn zoological Bulletin

Volume 60 [ Issue | | pp. 73-87 Bonn, May 2011

Aphyosemion musafirii (Cyprinodontiformes: Nothobranchiidae),

a new species from the Tshopo Province in the Democratic Republic

of Congo, with some notes on the Aphyosemion of the Congo Basin

Jouke R. Van der Zee! & Rainer Sonnenberg?

! Royal Museum for Central Africa, Zoology Department, Ichthyology, Leuvensesteenweg 13,

B-3080 Tervuren, Belgium. Corresponding author. E-mail: joukevdz@upcmail.nl

2 Max-Planck-Institut fiir Evolutionsbiologie, August-Thienemann-Strasse 2, D-24306 Plén, Germany

(current address)

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

Abstract. Aphyosemion musafirii, new species, is described from specimens collected near Ubundu (Ruiki River, Con-

go Basin, Tshopo Province, Democratic Republic of the Congo). Another population of A. musafirii is known from the

Romee River, 50 km West of Kisangani. The Ruiki and Romée Rivers are small tributaries on the left bank of the Con-

go River. Aphyosemion musafirii can be distinguished from its closest relative 4. castaneum by the male colour pattern.

A preliminary DNA analysis demonstrates that Aphyosemion s.s. consists of two major clades. Aphyosemion musafirii is

in a clade with A. castaneum, A. polli, A. lamberti, A. rectogoense, and A. congicum. The distribution of all species of

Aphyosemion s.s. is discussed.

Resumé. Aphyosemion musafirii, nouvelle espéce, est décrite a partir de spécimens récoltés prés de Ubundu (riviére Rui-

ki, bassin du Congo, Province Tshopo, République Démocratique du Congo). Une autre population de A. musafirii est

connue de la riviére Romée, 50 km a l'Ouest de Kisangani. Les rivieres Ruiki et Romée sont de petits affluents rive gauche

du fleuve Congo. Aphyosemion musafirii peut étre distingué de son plus proche parent A. castaneum par le patron de col-

oration male. Une analyse ADN préliminaire demontre que Aphyosemion s.s. consiste en 2 clades majeurs. Aphyosemion

musafirii est dans un clade avec A. castaneum, A. polli, A. lamberti, A. rectogoense et A. congicum. La distribution de

toutes les espéces de Aphyosemion s.s. est discutée.

Key words. Killifish, eastern Congo basin, Ubundu, systematics, taxonomy, biogeography.

INTRODUCTION

The genus Aphyosemion was erected by Myers in 1924

with the type species A. castaneum, described in the same

publication, from Kisangani (Democratic Republic of

Congo). At present the taxonomy of the genus is still not

settled, here we use Aphyosemion as proposed in two re-

cent publications of the authors (Sonnenberg, 2007; Van

der Zee & Sonnenberg, 2010). This is identical with the

subgenus Aphyosemion of other authors (e.g. Collier,

2007; Huber, 2007; Murphy & Collier, 1999; Wildekamp,

1993) and consists of 16 species currently accepted as

valid, which are, with the exception of two species from

Gabon, endemic to the Congo drainage.

Only three species are currently known to occur in the

eastern part of the Congo Basin: A. christyi (Boulenger,

1915), A. schoutedeni (Boulenger, 1920), and A. casta-

neum Myers, 1924 (Fig. 1). A fourth species, A. margare-

tae Fowler, 1936, is currently considered as a synonym

to A. christyi (Van der Zee & Huber, 2006). The majori-

ty of museum collections of these species originate from

the right bank of the Congo River. Aphyosemion schout-

edeni was assumed to be restricted to the type locality

“Medje at the Naya River”, a tributary to the Aruwimi

Basin, about 300 km northeast of Kisangani. Although the

types are in good condition, all colour has disappeared.

Since nothobranchiid species, at least within species

groups or genera, differ little in morphological characters

(Scheel, 1968, 1990), colour pattern of the male is cru-

cial for species identification. Topotypes collected by Lang

and Chapin in 1910, however, still have their colour pat-

tern preserved (Van der Zee & Huber, 2006) and it is close

to that of A. castaneum with the exception of the anal fin

colour pattern. This colour pattern is also present in sev-

eral Aphyosemion collections in the Royal Museum for

Central Africa (MRAC) (Tervuren, Belgium) originating

from the Aruwimi Basin, east of the Kisangani-Buta road

74 Jouke R. Van der Zee & Rainer Sonnenberg

A. castaneum

A. aff. castaneum

A. christyi

A. schoutedeni

A. plagitaenium

A. aff. polli

A. lefiniense

A. lamberti

0®d O00

Pot | Cala

A. aff. pollisouth @

A. rectogoense

A. decorsei

A. ferranti

A. lujae

A. congicum

A. sp. Cuvette

DNA sample

. schioetzi

. “schioetzi”

. chauchei

@ A.cognatum

Fig. 1.

Map showing the distribution of the genus Aphyosemion. Type localities are indicated by a T in the species symbol. The

type locality for A. congicum is unknown and for A. /efiniense and A. plagitaenium only collections from the type locality or in

proximity are currently known.

and may well represent 4. schoutedeni. Aphyosemion

christyi is restricted to the Epulu and Ituri drainages (about

350 km east of Kisangani) and is the only species that can

be identified by morphological characters (i.e., higher

number of dorsal fin rays than all other species)

(Boulenger, 1915; Van der Zee & Sonnenberg, unpubl. da-

ta).

Aphyosemion castaneum has long been misidentified as

A. christyi in literature and only recently Van der Zee &

Huber (2006) demonstrated that 4. christyi is restricted to

elevations above 500 m over a distance of 180 km north-

east of the type locality, Bafwasende. Aphyosemion cas-

taneum 1s widespread around Kisangani, but additional

records of this species are known from Salonga Park by

Scheel (1990), who didn’t specify the exact location with-

in the park, Lompole, and Yaka in the central Congo Basin

(Huber, 2005b). According to Huber (2005b) it is likely

Bonn zoological Bulletin 60 (1): 73-87

that these populations represent a different, undescribed

species. Consequently, A. castaneum might be restricted

to the right bank of the Congo River (Huber, 2005b), with

the exception of some populations that were found close

to the left bank, and the specimens collected on the left

bank with a wide distribution in the central basin were dis-

tinguished here as A. sp. aff. castaneum (Fig. 1).

From the Romée and Ruiki Rivers on the left bank of the

Congo River in the eastern part of the central Congo

drainage, three collections of Aphyosemion are known

from the MRAC, originally identified as A. christyi. The

colour pattern of the anal and caudal fins in preserved

specimens differs from A. castaneum, which is widespread

around Kisangani. In 2007, A. Van Deun (Leuven, Bel-

gium) collected a species of Aphyosemion with the same

colour pattern as the three previous collections from the

Romée and Ruiki Rivers, at two localities just north of

©ZFMK

Anew Aphyosemion from the eastern Congo 75

Table 1. List of specimens used for the DNA analyses with locality information and GenBank accession numbers. Abbreviations:

DRC = Democratic Republic of Congo; RCA = Republic of Central Africa; US = sample provided by Uli Schliewen, ZSM, Mu-

nich; AS = aquarium bred strain; CI = commercial import; WC = wild caught sample.

Species sample no. Country Collection locality GenBank acc. no.

Aphyosemion castaneum AS RS1408 DRC Kisangani JF307802

A. castaneum AS RS1499 Republic Congo Oyo JF307797

A. castaneum WC RS1790 DRC AVD 3 JF307803

A. cf. chauchei AS RS1527 Republic Congo =Olombo JF307796

A. cf. decorsei AS RS1521 RepublicCongo Lobaye JF307795

A. cognatum WC RS1515 Republic Congo =Mbonza II (US 107) JF307791

A. cognatum AS RS1520 DRC Lake Fwa JF307793

A. cognatum AS RS1529 DRC Kinsuka JF307794

A. congicum AS RS1617 DRC Z 82/17 JF307798

A. elegans WC RS1747 DRC Boende CI 2006 JF307792

A. elegans WC RS1513 DRC Inongo (US 24) JF307790

A. elegans WC RS1514 DRC Inongo (US 66) JF307789

A. lamberti AS RS1256 Gabon BSWG 97/9 JF307781

A. musafirii WC RS1787 DRC AVD 1 JF307804

A. polli AS RS1584 DRC Cl JF307800

A. polli AS RS1479 RCA RCA 91/1, Kapou 1 JF307801

A. rectogoense AS RS1419 Gabon PEG 95/16 JF307799

A. sp. aff. castaneum WC RS1506 DRC Lompolé (US 74) JF307782

A. sp. aff. castaneum WC RS1507 DRC Lompoleé (US 79) JF307783

A. sp. aff. castaneum WC RS1510 DRC Yaka (US 33) JF307786

A. sp. aff. castaneum WC RSI511 DRC Yaka (US 45) JF307787

A. sp. aff. castaneum WC RS1512 DRC Yaka (US 61) JF307788

A. sp. aff. elegans WC RS1508 DRC Lui Kotalé (US 75) JF307784

A. sp. aff. elegans WC RS1509 DRC Lui Kotalé (US 77) JF307785

A. sp. Cuvette AS RS1019 DRC Boende 2002 JF307780

Mesoaphyosemion cameronense RS262 Cameroon CMM 40 AY 748282

Ubundu on the left bank of the Congo River. Based on

live male colour pattern and a preliminary mitochondrial

DNA analysis this species is described here as

Aphyosemion musafirii, new species.

MATERIAL AND METHODS

Morphometric measurements were taken with a digital cal-

liper, partly under a dissecting microscope, and rounded

to the nearest 0.1 mm. Counts and methods follow Ami-

et (1987). Measurements, including subunits of head, are

presented as percentages of standard length (SL). The

number of all visible rays of dorsal, anal, caudal, pelvic,

and pectoral fins were counted, the abbreviation D/A

means the relative position of the first dorsal fin ray with

regard to the opposite anal fin ray. Count of scales on the

Bonn zoological Bulletin 60 (1): 73-87

mid-longitudinal series is the number of scales between

the upper attachment of the opercular membrane and the

caudal fin base. Excluded are the scales posterior to the

hypural junction, which were counted separately. Nomen-

clature for the neuromast system on the head follows

Scheel (1968) and Van Bergeijk & Alexander (1962), and

that for the supraorbital (frontal) squamation follows

Hoedeman (1958).

Total DNA was extracted from fin clips or muscle tissue

from the caudal peduncle of ethanol preserved specimens,

following a modified DNA extraction protocol after Gus-

tinicich et al. (1991). Specimens used for DNA analyses

are listed in Table 1 with GenBank accession numbers. A

fragment of the mitochondrial cytochrome b gene was se-

quenced for 25 specimens of Aphyosemion and Meso-

aphyosemion cameronense (GenBank accession number

OZFMK

Og O¢ ST LNG ce TE Lé O¢ O¢ O¢ 9T ST 8ST 6T 8's £9 OL OL c9 r9 89 £9 cL 6L 8Cl IES ES [@}@)

00 9¢ 6¢ Ib IF € oF ot 6¢ SNE ve Le Se 79 99 6L 6L 89 SL OL EL c8 rs Pel 6TS LDOOD

SS OE If It eis 6€ 6€ Of OE re Le sc P9 99 6L 6L 89 CL OWE SL Git Ps vel O0éS LDOD

ol VT PT | 0T OT OT (Et rl Lil sl c9 89 OL ae We ae SL OL ans) 68 Eel LOSIVHO?

€¢ AG Ll 61 61 ol ol 91 ol OT Lg UL 08 08 tL DHL 08 OL 08 £8 Cel Ics TOA

£0 cl Ll Ll Ll rl cl a} 91 99 OL SL CL OL OL VL GE £8 L8 ak | 608 LATAS

ot Pu PL RA A I DE ER) i IB Ol OS 1A TAS

ie G0 BO. 60 0. TRO. 0 O> TB) D E SS O A Lelia

g SOMMEOOMMCIOMMESION © (Si0) Gin S091 Mie See 2S) MIGe aie SC bil pS Uuly6:-mICGIIMETGISIGVOS

is EW 0 GU ERI A VR, AY a A SS ee NG CHa IIS ISVS

= SOON ASIOENICTO MEL OR Sliven GSu cS. Ee eISL. SE) “ib pS) or cis! me OISISVOs

pa Ct), — SW > fF) © a A) GS ST TK = Ey A IR HS pISta Tas

. COMMEPOM MNOS: <COMe tt fie mO0 AS On ache Sie “0 1G) cic] C1S1ATAS

NS Ok ENC9. 919) OF 9 00k BSOn STU HOW “AG, 98. CSE meLOSISvos

= (6) 18) OE RHE 00) a = MI AA FAO OSI OS

& Ol SE kG RYSEer OIG COs ah 9 5 [la meCeN marc] 6IFIOAU

x RS as GRp (AC ERS GIS 7B SH Ay 9STINVT

3 Pir = OKS SES SO RE GI P8S110d

Ss OSGi S:Se eLiSe ICON NLCLy mG MGIC 6LbI 10d

Ge EC Re Se OE TG) L8LISNW

Ee ee ts iew A UFC 06LISVO

O:e SO Oe etal SOFISVO

69 6L OT 66FISVO

6S 6FI LI9INOD

9TI 6101 Nods

6cSI. OZST «LZSI=CIZST «= GOST «80ST «LEAT. «= TISE. =OLIST «OUST. = FIST «= SIST «= LOST. «(90ST GIF. «(STI SCOPSSISGLFI.«CLSLI.«OGLI. SOFT) 6GFL. SC LIDT. «GIO. = 297

909 909 VHO? 3309 JIS AIAS aId SvoOs Svos_ svos YAS SVOS SVOS OAM WWI 10d 10d SAW SVD SVD SVD NOO nods wo satoads

‘amoang ‘ds ‘p = nods ‘sunsaja ‘ye ‘ds ‘p = qqqs ‘wnaunjspo ‘jye ‘ds “fp = SYOS ‘asuaosojoal ‘py = QTY ‘lod “F = 10d ‘U-ifps

“NW = SQW “Uaqup] “Y = WY] ‘supsaja ‘py = Tq ‘1a81ozap “Jd “P= QA) ‘wina1BUu0d "p = NOD ‘uinjpUusod “Y = NOD ‘layonvyd "Jo "P = VWHO9 ‘uinaupjspo ‘py = SW_ ‘asuauos

Xe -AUDI PW = WY) ‘so[duies YN ou) JO} SuOneIADIgQGY “eIep SUISSIW JO UOISN[OXd asiMAted YIM sadUaNbas q dULO.AY9O}AD [RLIPUOYIOUU dy} 1OJ soouRsIp-d poooLIOsUL) *Z BIqUL

©ZFMK

Bonn zoological Bulletin 60 (1): 73-87

Anew Aphyosemion from the eastern Congo 77

Fig. 2.

Aphyosemion musafirii, male, collected with the

types by A. Van Deun, 22.10. 2007, not preserved. Type local-

ity, 67 km on the road from Kisangani to Ubundu, Democratic

Republic of Congo. Photo: H. Ott.

AY 748282, published in Sonnenberg & Blum [2005]) was

used as outgroup, for lab protocols see Sonnenberg et al.

(2006).

Resulting sequences were aligned with ClustalX 1.8

(Thompson et al., 1997) and checked by eye in BioEdit

7.0.5.3 (Hall, 1999). All sequences were translated into the

corresponding amino acids and tested for the anti-G bias

of mitochondrial sequences (Zhang & Hewitt, 1996) to

confirm for functionality and mitochondrial origin. Un-

corrected p-distances with pairwise exclusion of missing

data were calculated in MEGA 4.1 beta 3 (Tamura et al.,

2007) and are given in Table 2.

Fig. 3.

Aphyosemion musafirii, male, 7 km north of Ubundu,

Democratic Republic of Congo, collected by A. Van Deun,

22.10.2007, not preserved. Photo: H. Ott.

Analyses of sequence data were performed with PAUP

4.0b10 (Swofford, 1998) by maximum parsimony and

with MrBayes 3.1.2 (Huelsenbeck & Ronquist, 2001) by

a Bayesian approach, all analyses with the identical respec-

tive settings given in Sonnenberg & Busch (2010). In ad-

dition, we performed a maximum parsimony analysis with

the same parameters as the previous, but with exclusion

of the third protein coding position. Only nodes, which

are supported with bootstrap values above 75% or poste-

rior probabilities above 95% were considered as support-

ed by the data.

We used the software SplitsTree (Huson & Bryant, 2006)

to calculate a split decomposition network representation

of the dataset.

Bonn zoological Bulletin 60 (1): 73-87

As species concept we adopted the approach by Moritz

et al. (2000), which is similar to the Evolutionary or Phy-

logenetic species concepts (as discussed in Kottelat, 1997).

RESULTS

Aphyosemion musafirii, new species

(Figs 2—6, Tables 3—5)

Holotype. MRAC 2011-007-P-1, male, 36.7 mm SL,

Democratic Republic of Congo, Tshopo Province, 67 km

on the road from Kisangani to Ubundu (1°30’ N, 25°21?

E), 450 m altitude, north-eastern Congo basin, A. Van De-

un, 22 October 2007.

Fig. 4.

Aphyosemion musafirii, male, F1 from specimens col-

lected 7 km north of Ubundu, Democratic Republic of Congo,

showing a different anal fin pattern than most of the wild col-

lected males from that population. Photo: W. Grell.

Paratypes. MRAC 2011-007-P-2-5, 4 females, 30.1—33.5

mm SL, collected with the holotype; MRAC 2011-007-

P-6-9, 4 males, 32.9-34.6 mm SL, collected with the holo-

type.

Additional non-type material. MRAC 90-30-P-1471, la-

belled as A. christyi (Boulenger, 1915), Democratic Re-

public of Congo, Ubundu, L. De Vos, February 1990.

MRAC_ 90-47-P-846-853, labelled as A. christyi

(Boulenger, 1915), Democratic Republic of Congo, Riv.

Romée, km 30 route Kisangani-Opala, L. De Vos & C. Da-

nadu, 07.02.1990.

Fig. 5.

Aphyosemion musafirii, female, 7 km north of Ubun-

du, Democratic Republic of Congo, collected by A. Van Deun,

22.10.2007, not preserved. Photo: H. Ott.

©ZFMK

78 Jouke R. Van der Zee & Rainer Sonnenberg

Fig. 6.

Variability of the anal fin pattern in Aphyosemion

musafirii (all examples from wild caught specimens).

MRAC_ 90-47-P-854-863, labelled as A. christyi

(Boulenger, 1915), Democratic Republic of Congo, Riv.

Romée km 3, rive gauche, route Kisangani-Opala, L. De

Vos & C. Danadu, 10.—17.04.1990.

Diagnosis. Aphyosemion musafirii (Fig. 2—6) is placed

within the genus Aphyosemion by the combination of the

following characters: preopercular neuromast system with

6 pores, slender body, posterior origin of dorsal fin with

less than 10 fin rays, females with a strong reticulation due

to dark scale borders, and the extended edges of the cau-

dal fin in males (Huber, 2005a).

Males of A. musafirii differ from all other representatives

of Aphyosemion in the north-eastern Congo Basin by the

in average broader red margin of the dorsal fin. This red

border is narrow (up to 10%) in all other species in this

area and is often absent at the distal end of the fin.

Bonn zoological Bulletin 60 (1): 73-87

Males of A. musafirii differ from A. castaneum by the ab-

sence of a red band in the approximate centre of the anal

fin, absence of a red infra-buccal band, absence of red

edges around the light ventral zone in the caudal fin, ab-

sence of, or only very narrow, yellow margin of pelvic fins

and a higher average number of red dots on side (A.

musafirii: min. 51, max. 117, average = 79, sd = 23, num-

ber of specimens = 11; 4. castaneum: min: 13, max. 57,

average = 30, sd = 15, number of specimens = 20). These

dots are arranged in more or less regular, interrupted rows

in A. musafirii, whereas these dots are irregularly distrib-

uted in A. castaneum. The higher number of red dots on

the side also distinguishes males of A. congicum (25-40),

A. schoutedeni (17-28) and A. polli (11-49) from A.

musafirii. It can also be distinguished by the more or less

regular interrupted rows of red dots from A. sp. Cuvette,

A. elegans (Boulenger 1899), and A. plagitaenium, which

have either a pattern of vertical streaks or oblique bars on

the side in males. Males of A. musafirii differ from A.

schoutedeni by the same characters as A. castaneum, with

the exception that A. schoutedeni lacks a red band in the

anal fin. Males of A. musafirii have less spots in the cau-

dal fin than A. christyi (35—S5 in A. musafirii; over 70 in

A. christyi). The background colour of the flanks is blue-

green in A. musafirii and purple-blue in A. christyi. In ad-

dition to colouration characters, males of A. musafirii can

be distinguished from A. christyi by the lower number of

dorsal fin rays (7-9 in A. musafirii versus 10-11 in A.

christyi).

Description. See Figures 2—6 for general appearance and

Tables 3—5 for morphometric and meristic data of the type

series. Aphyosemion musafirii shows strong sexual dimor-

phism, males more colourful, unpaired fins larger, dorsal

and anal fins with posterior fin rays extended. A medium

sized, slightly laterally compressed species; dorsal profile

slightly convex, greatest body depth approximately at

pelvic fins. Ventral profile slightly convex from head to

end of anal fin, concave on caudal peduncle. Snout slight-

ly rounded, mouth directed upwards, lower jaw longer

than upper jaw. Dentary bears an outer row of large and

inner irregular rows of smaller unicuspid, curved teeth; the

premaxilla bears some larger and several smaller unicus-

pid and curved teeth.

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

& Alexander, 1962) neuromasts in separate grooves, the

preopercular canal with six pores.

Scales cycloid, entirely scaled except ventral surface of

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

tudinal series 29-30, with 1—2 scales posterior to the hy-

pural plate; 7 transversal scales, 12 scales around the cau-

dal peduncle.

©ZFMK

A new Aphyosemion from the eastern Congo 79

Table 3. Morphometrics of Aphyosemion musafirii, new species, (H = holotype, P = paratypes: 4 males and 4 females ). All mea-

surements in percentages of standard length, standard length in mm.

Pp Q

Ho Pd Pd Pd Pd 2 PQ PQ

Standard length 36.7 34.6 34.2 33.3 32.9 3355 32.6 313 30.1

Body depth 21.0 18.4 19.6 19.5 18.8 18.8 20.8 20.1 21.4

Head length 21.5 20.5 21.6 21.0 20.4 19.1 20.5 19.5 19.0

Eye diameter 6.3 6.3 6.4 6.3 6.1 Sil 6.4 6.4 6.9

Interorbital width 12.0 12*77 11.1 11.4 IES 10.7 11.3 ez 13.0

Pre-dorsal length 63.2 61.0 66.0 66.4 62.1 60.2 66.2 65.8 63.5

Pre-anal length 55.0 54.9 535 54.3 50.8 D922 56.1 54.6 S13

Dorsal fin base 11.2 12 10.2 11.1 11.2 11.0 10.1 10.9 11.0

Anal fin base 19.1 18.8 eS 19.2 18.2 17.3 16.1 173 i2

Caudal peduncle depth 12.5 HES 11.1 Ihveg/ 11.6 10.7 11.7 1S 11.9

Small dorsal fin with 7—9 fin rays, first dorsal fin ray in-

serts above the 7—10th anal fin ray; anal fin with 13-15

rays; posterior dorsal and anal fin rays slightly elongated

in males; caudal fin with 21—24 rays, with extensions on

upper and lower fin rays. Pectoral fin with 12-14, pelvic

fin with 5 rays.

Live colouration Males. (Figs 24 & 6) Flanks greyish

brown with blue-green iridescence. Edges of scales on the

flanks with dark pigmentation resulting in a reticulated

pattern. Dorsally the scales have broader pigmented edges

than ventrally. Flanks with approximately 50 to 120 red

spots. These spots are mainly situated at the anterior edge

of the scales and are mostly irregularly distributed in lines,

usually forming up to five parallel lines. Three red streaks

on opercle in an approximate 45° angle. Infra-buccal band

absent or only present at the sides of the jaw.

Table 4. Meristics of Aphyosemion musafirii, new species.

Numbers indicate observed values; numbers in parenthe-

ses frequency of occurrence; values found for the holo-

type are indicated by an asterisk.

meristic count values (frequency)

9 (3*), 8 (4) 7 (2)

13 (5*), 14 (3), 15(1)

+7 (1*), +8 (1), +9 (3), +10 (4 fem.)

21 (3*), 22 (2), 23 (3), 24 (1)

5 (9*)

12 (2*), 13 (6), 14 (1)

29 (2), 30 (6*), 31 (1)

7 (9*)

circumpenduncul. scales 12 (9*)

dorsal fin rays

anal fin rays

D/A

caudal fin rays

pelvic fin rays

pectoral fin rays

lateral line scales

transversal row scales

Bonn zoological Bulletin 60 (1): 73-87

Pectoral fin yellow, unspotted or just with some tiny spots

in the centre. Pelvic fin yellow with several prominent red

spots. Dorsal and anal fin blue iridescent at the base and

yellow distally, provided with red spots; spots more nu-

merous and larger at the base than distally. Dorsal fin

edged with a broad dark red band (up to 30% of fin length

at mid section). Anal fin edged with a narrow red band.

Spots on anal fin rounded to elongated, sometimes miss-

ing at the distal part of the fin, leaving a broad yellow sub-

distal band. Caudal fin light blue with yellowish distal

edges provided with rounded and/or 35—S5 elongated red

spots. Dorsal and ventral edge of the caudal fin provided

with a broad dark red band.

Females. (Fig. 5) Flanks grey with darker reticulation. Red

streaks on opercle reduced and infra-buccal band absent.

All fins transparent. Edge of anal fin and distal part of ven-

tral fins light blue. Faint spots on anal fin, more promi-

nent spots on base of dorsal and dorsal part of caudal fin.

After one year of preservation in ethanol. Males. Flanks

light brown with transition to light ventral side. Reticu-

lation, buccal band, and red streaks on opercle as in live

specimens. Spots on flanks as in live specimens, but small-

er and more vague. All fins transparent greyish, provid-

ed with spots as in live specimens, but vague and pink.

Females. As in live specimens, but spots on anal fin hard-

ly visible.

Distribution. (Fig. 1) Aphyosemion musafirii is restrict-

ed to brooks in the Ruiki and Romée River systems on

the left bank of the Congo River between Kisangani and

Ubundu, Tshopo Province, Democratic Republic of Con-

go.

©ZFMK

80 Jouke R. Van der Zee & Rainer Sonnenberg

Table 5

Comparison of the morphometric values for Aphyosemion musafirti, new species, and members of the sub-

genus Aphyosemion. Abbreviations used in the table: CAS = A. castaneum, CHR = A. christyi, COG = A. cognatum,

CON = A. congicum, LEF = A. lefiniense, MEL = A. melanopteron, MUS = A. musafirii, POL = A. polli, TEU = A.

teugelsi, E =

dorsal fin distance, pA =

sd = standard deviation.

eye diameter, I = interorbital width, BD = body depth, HL = head length, HW = head width, pD = pre-

preanal fin distance, DB = dorsal fin base, AB = anal fin base, CD = caudal peduncle depth,

species location sex SL 13) I BD HL pD pA DB AB CD

COG LNB) 3 33.5 8.0 11.0 229 18.9 67.1 62.2 IES: 21.9 14.3

CHR HZ 85/14, 29:1 7.9 9:3 18.4 PA) 65.7 59.0 11.6 19.6 11.2

Epulu Q 312 ES 9.3 17.1 22.1 64.5 58.7 M3) 19.9 10.9

POL RCA 91/1, 36:0 7ES 10.6 2285 23.3 68.3 59.4 12 Ped 13.1

Kapou Q DIT 16:9 8.2 23.6 DOr 67.0 61.9 HS 7/ 19.9 12.6

CON type of MEL 4 7.8 OD) 21.6 27.8 67.3 61.5 12.1 20.9 13.4

CAS HZ 85/13 3 20.8 7.7 13.4 20.4 20.2 70.2 53.8 12.0 DD Mila

2 19.4 7.7 11-9 20.8 19.8 68.3 ll 11.5 20.0 W273)

LEF Luna River 3 23 8) 12.1 19.0 24.1 66.6 54.9 7.0 13.0 9.6

2 19.2 8.3 11.4 21.0 23.6 69.9 57.0 8.6 16.2 11.8

mean 7.8 10.6 20.8 DDD 67.7 58.1 11.0 19.6 12.2

sd 0.5 1.4 2.0 2.6 19 3.4 1.6 Dei 1.4

range 6.9-8.7 8.2-13.4 17.1-23.6 18.9-27.8 64.5-70.2 52.1-62.2 8.6-12.1 13.0-22.1 9.6-14.3

MUS

mean 6.3 12.1 MS) 2 30.9 68.3 61.1 14.6 19.0 12.4

sd 0.3 0.5 2.2 1.0 1.6 LES 0.7 1.2 0.2

range 5.4-16.9 11.3-12.8 16.6-21.2 29.3-32.0 66.0-70.5 59.2-62.8 13.8-15.3 17.4-20.7 12.1-12.7

Etymology. Aphyosemion musafirii is named after Dr.

Jean Musafiri (Ubundu, Democratic Republic of Congo),

coordinator for the national tuberculosis and leprosy con-

trol programme in the “Province Orientale Occidentale”,

the huge forest area around Kisangani. The name Musafiri

means “traveller” in Swahili. Indeed, he travels around the

area under very difficult circumstances, covering enor-

mous distances by jeep, motorised canoe or small motor-

cycle. Dr. Musafiri was born in Ubundu and has always

stayed in the province to help his people, in spite of the

very difficult living conditions and the atrocities of the war

in the eastern Democratic Republic of Congo. He made

it possible that the type material of A. musafirii and a new

species of Fenerbahce (Sonnenberg, Woeltjes & Van der

Zee, submitted) could be collected by A. Van Deun (In-

Fig. 7. Aphyosemion castaneum, 11 km west of Kisangani,

Democratic Republic of Congo, not preserved. Photo: H. Ott.

Bonn zoological Bulletin 60 (1): 73-87

Fig. 8.

Aphyosemion castaneum, Wani Rukula, 65 km south-

east of Kisangani, Democratic Republic of Congo. Photo: H. Ott.

stitute of Tropical Medicine, Antwerp) at the occasion of

an external evaluation visit of the tuberculosis/leprosy pro-

gramme.

DNA analyses. The resulting sequence alignment has a

final length of 760 bp, the base composition shows the,

for mitochondrial sequences typical, A/T bias (Zhang &

Hewitt, 1996). In two sequences (RS1747 and RS1521),

up to 13 N were introduced at the start of the alignment

for equal sequence length. We found 192 variable and 130

phylogenetic informative positions. The DNA fragment

translates into 253 amino acids, of which 27 are variable

and 15 phylogenetic informative, and contains no unex-

pected stop codon. Uncorrected pairwise distances be-

©ZFMK

Anew Aphyosemion from the eastern Congo 81

62/-/ 100/99 A. sp. Cuvette 1019

A. congicum 1617

A. castaneum 1499

98 / 64/ 100 / 100 A. castaneum 1408

A. castaneum 1790

A. musafirii 1787

A. polli 1584 CAS group

100 / 99/88/87 A. polli 1479

A. lamberti 1256

88 /-/-/68 A. rectogoense 1419

A. sp. aff. castaneum 1506

A. elegans 1513

A. sp. aff. castaneum 1507

A. elegans 1514

D A. sp. aff. castaneum 1510

80 / 93/87/88

E|!A. sp. aff. castaneum 1512

A. sp. aff. castaneum 1511

B A. elegans 1747 COG group

Cc pA. sp. aff. elegans 1508

A. sp. aff. elegans 1509

A. cf. decorsei 1521

A. cf. chauchei 1527

A}A. cognatum 1520

99 /-/ 100/100

A. cognatum 1529

A. cognatum 1515

M. cameronense 262 A) 100/ 90/ 100/100

B) 92/ 65/100/ 100

C) 100/ --/ 100/100

D) 76/ --/ 100/100

10 E) 96/63 /100/ 100

Fig. 9. Phylogenetic tree by maximum parsimony, tree length = 283, consistency index = 0.5194, homoplasy index = 0.4806,

retention index = 0.7467, and rescaled consistency index = 0.3879. Shown is one of 36 equally short trees, on the left side of nodes

support values are shown for maximum parsimony bootstrap analyses and Bayesian posterior probabilities in the following order:

maximum parsimony / maximum parsimony with exclusion of third position / Bayes with Nst = 2 / Bayes with Nst = 6. Only sup-

port values for nodes are shown, for which at least in one analysis a bootstrap value of 75% or posterior probabilities of 95% were

reached. No value given means that the support value in all analyses 1s below the previous given values or the node is not recov-

ered in the analysis. Some values are given below the tree and are indicated on the respective nodes by a capital letter for better

readability.

Bonn zoological Bulletin 60 (1): 73-87 OZFMK

Jouke R. Van der Zee & Rainer Sonnenberg

A_polli_1473

A_lamberti_1256

A_rectogoense_1419

COG group

A_cognatum_1523

A-cognatum_1520

A_cognatum_1515

A_cf_chauchei_1527

A_cf_decorsei_1521

Jil

A_sp_aff_castaneum_1510 Lh

A_sp_aff_castaneum_1512

A_sp_aff_castaneum_1611

A_elegans_1513

Acelegans_1514

A_sp_saff_castaneum_1506

A_sp_atf_castaneum_1507

A_Aegans_1747

A_sp_aff_elegans_15093

A_sp_aftf_elegans_1508

A_polli_1584

CAS group

A_musatirii_1787

A_castaneum_1730

A_castaneum_1408

A_castaneum_1499

A_congicum_1617

A_sp_Cuvette_1019

outgroup

M_cameronense_262

Fig. 10. SplitsTree network representation of the Aphyosemion dataset. Number of taxa = 26, 760 base pairs, fit = 98.01, uncor-

rected p-distances, neighborNet.

tween sequences are presented in Table 2, the observed

values found between Aphyosemion and Mesoa-

phyosemion as the outgroup range from 12.6—14.9%, the

maximum observed value within Aphyosemion is 9.1%.

The samples of A. cognatum from aquarium strains orig-

inating from Kinsuka and Lake Fwa are, despite nearly

900 km distance, identical, and also with the shorter se-

quence of an 4. cognatum from Lake Fwa published by

Murphy & Collier (1999, GenBank acc. no. AF002324),

therefore it might be possible that the strains were erro-

neously mixed or mislabelled since their introduction in

the killifish hobby. A test with a second specimen from

the Kinsuka strain resulted in the same sequence (data not

shown).

The A. cognatum 1515 sample is only represented by a

female, so the species identification is tentative based on

the knowledge of other Aphyosemion collections around

this area and the resulting sequence is identical with a

shorter sequence of Murphy & Collier (1999, GenBank

acc. no. AF002327), which they have published as A. el-

egans from Naoimda, and slightly different from their se-

Bonn zoological Bulletin 60 (1): 73-87

quence of what they call 4. elegans from Madimba (Gen-

Bank acc. no. AF002328). It is not clear if Naoimda is a

misspelling of Madimba. However, the specimens from

Madimba belong to 4. cognatum (e.g. see Seegers, 1997,

p. 74), the sequences differ in only two bases.

The phylogenetic analyses (Fig. 9) give an only partially

resolved tree of the analysed specimens. In addition, there

are differences in support for nodes; excluding the third

protein coding positions gives no bootstrap support above

50% for several nodes, which were recovered by the oth-

er analyses (Fig. 9). Aphyosemion musafirii seems to be

closer to A. castaneum, however, this node is only sup-

ported by the maximum parsimony analyses. Comparing

the phylogenetic hypotheses summarized in the tree (Fig.

9) with the split-decomposition network representation

(Fig. 10), the network indicates a clear separation of the

analysed specimens into two main groups, which is not

that prominent in the dichotomous phylogenetic hypothe-

ses (Fig. 9). One group, in the following called the A. cas-

taneum group (CAS) is characterised by longer branches

between the species (range 1.1—8.2% p-distance), where-

as the second group, the 4. cognatum group (COG) has

©ZFMK

A new Aphyosemion from the eastern Congo 83

shorter internal branch length (0.0-4.1% p-distance), the

divergence between both groups ranges between 5.8—9.1%

p-distance. The CAS group contains, according to the net-

work (Fig. 10), five species, which were found from the

western to the eastern Congo Basin, and two species from

Gabon. However, this group is not recovered by the phy-

logenetic analyses with any significant support (Fig. 9).

The second group in the network (Fig. 10) is recovered

with high support (Fig. 9) at least by three analyses and

contains samples from the western and central Congo

Basin.

DISCUSSION

Colour pattern

The main criterion for distinguishing species in notho-

branchiids and especially Aphyosemion is the male

colour pattern. All Nothobranchiidae show a polygamous

mating system and a high degree of sexual dimorphism.

Amiet (1987) therefore assumed that male colouration

might be important in mate choice of females in notho-

branchiid fishes. Van der Zee et al. (2007) assumed that

especially the colour pattern of the caudal peduncle and

unpaired fins plays an important role in female mating

preferences. Kullmann & Klemme (2007) were able to

demonstrate for Chromaphyosemion that females prefer

to mate with their own males, for Diapteron this was stud-

ied by Brosset & Lachaise (1995). Thus female mate

choice on divergent male colour characters may enable

speciation by sexual selection. After secondary contact of

previously allopatric populations, species cohesion is then

easily maintained in parapatry or sympatry.

The current study shows that the genus Aphyosemion re-

mains taxonomically problematic, which is in part due to

old type material with no traces of the most important

characters, the male colouration pattern, and descriptions,

which do not give detailed information about live or pre-

served colouration. On the other hand, many species show

a certain degree of variation in colour pattern within and

between populations, which makes identification in some

cases difficult. Differing species identification between au-

thors further complicates killifish literature, as for exam-

ple in the case of the A. elegans samples in the study of

Murphy & Collier (1999) mentioned above. A third sam-

ple listed as A. e/egans in their study, an aquarium strain

from Epoma (Murphy & Collier, 1999) was later described

as A. plagitaenium by Huber (2004).

Phylogeny

Our preliminary molecular phylogeny of this group gives

a very complex pattern. The CAS (A. castaneum) group

Bonn zoological Bulletin 60 (1): 73-87

might be not a monophyletic unit, but consists of sever-

al species with deep divergences, which is reflected by the

large p-distances found in this group (Table 2). The root

of the group might be placed close to this species com-

plex (Fig. 10). The COG (A. cognatum) group consists of

species with lower within-group p-distances (Table 2), ap-

pears to be monophyletic and might contain species of

more recent origin. Both groups cover large ranges in the

Congo and adjacent river basins in Gabon (A. /amberti and

A. rectogoense), have a kind of mixed distribution and oc-

cur In some cases 1n sympatry (e.g. A. elegans and A. sp.

Cuvette) or even syntopically (A. sp. Cuvette and A. sp.

aff. castaneum, A. chauchei and A. elegans; A. ‘“‘schioet-

zi’ and A. polli).

Interestingly some phenotypically different species (A. sp.

aff. castaneum and A. elegans / A. sp. aff. elegans) in the

Cuvette centrale turn out to be closely related and not ge-

netically separated by the mitochondrial DNA data (Fig.

9). Compared with some additional nuclear 28S rDNA

(LSU) sequences (unpublished data), it turns out that the

studied samples can be separated into two groups: one in-

cluding A. castaneum and A. sp. aff. castaneum and on

the other hand the samples of A. elegans / A. sp. aff. ele-

gans. This indicates that most probably mitochondrial in-

trogression between species of the CAS and COG group

has occurred in an area were both groups are at least para-

patric. However, despite the potential of hybridization as

is indicated by the mitochondrial introgression, both

species or species groups live at least in sympatry and have

stable distinguishing characters.

Distribution of the genus Aphyosemion

With the addition of A. musafirii the number of species

in the eastern Congo Basin is increased to four (see In-

troduction). The study of the Aphyosemion specimens in

the MRAC by the first author also added some collections

of other species from this area (Fig. 1), in part due to

misidentified samples. However, the maximum number of

eight species is found in the westernmost part of the Con-

go Basin. An artificial factor, which might explain the con-

centration of species here, is the higher number of sam-

ples from this area. A natural cause of high species diver-

sity might be the proposed existence of a forest refuge dur-

ing dryer periods in the past (Leal, 2004; Maley, 1996;

Plana, 2004). Two of these eight species are widely dis-

tributed in the northern part of the basin: A. castaneum

Myers, 1924 and A. polli Radda & Piirzl, 1987. These two

species were probably transported downstream by the

Congo and Ubangui rivers, and both can also be found at

Pool Malebo. Aphyosemion elegans is here present in the

northern tributaries of the Likouala River, but is wide-

spread in the central Congo Basin (Cuvette centrale) and

north of the middle Congo section up to Buta.

©OZFMK

84 Jouke R. Van der Zee & Rainer Sonnenberg

Three species have a more restricted distribution in the

Central Republic of Congo: A. chauchei Huber & Scheel,

1981 is found in the Likouala and Alima drainages, A.

lefiniense Woeltjes, 1984 is restricted to the southern trib-

utaries of the Lefini River, and A. plagitaenium Huber,

2004 is only known from the type locality at Epoma in

the northern Central Republic of Congo.

Aphyosemion schioetzi Huber & Scheel, 1981 has a dis-

junctive distribution: a rather restricted area in the south

around the type locality and a huge area in the north, sep-

arated by a gap of almost 280 km where only A. /efiniense

is found (Huber & Scheel, 1981). The present authors sug-

gest that the northern populations (labelled here in the fol-

lowing “‘schioetzi”’, Fig. 1) do not belong to A. schioetzi

sensu Strictu, since all representatives are lacking the dark

red edge of the anal fin and seem to be built more slen-

der than A. schioetzi. The northern populations might rep-

resent a new species or be conspecific with A. decorsei

(Pellegrin, 1904). The status of A. decorsei has been sub-

ject to discussion for a long time (Huber, 1994, 2004,

2005a; Scheel, 1968, 1990; Wildekamp, 1993). Poll (1951)

even placed it in Epip/atys and in the description of Hap-

lochilus decorsei Pellegrin assumed it to be close to

Aplocheilichthys spilauchen. Myers (1924) placed it in

Aphyosemion with some hesitation. Scheel (1968, 1990),

Huber (1994, 2004, 2005a), and Wildekamp (1993) all

have seen the types and they confirmed Myers’s statement.

The types from the Central African Republic are in poor

condition with the colour pattern lost. Huber (2004, 2005a)

stated that A. decorsei has only few red spots on the flanks

and might be conspecific with A. po/li, the latter being a

junior synonym. Wildekamp (1993), however, is con-

vinced that A. decorsei has many spots on the flanks, based

on the light spots on the scales of the syntypes. Red pig-

mentation pattern, after preservation in formalin and trans-

fer in ethanol, leaves corresponding patterns of lighter ar-

eas than the body colouration (Van der Zee & Sonnenberg,

2010). Aphyosemion polli has not only few spots on the

flanks, but also has very few spots (or no spots) on the

anal fin, arranged near the base of this fin. In the original

description of A. decorsei Pellegrin writes: “la dorsale,

Vanale et des ventrales avec des petits points carmins plus

ou moins nombreux” (dorsal, anal, and ventral fins with

small more or less numerous carmine spots). The present

authors agree with Wildekamp’s argumentation: 4. decor-

sei 1S a Species with many spots at least on the anal fin.

Aphyosemion cognatum Meinken, 1951 has a huge dis-

tribution area in the south from the right bank of the Con-

go River to Lodja in the upper Lukenie basin. Sympatric

with A. cognatum occur A. ferranti (Boulenger, 1910), A.

congicum (Ahl, 1924), and A. /ujae (Boulenger, 1911).

Bonn zoological Bulletin 60 (1): 73-87

In the large Cuvette centrale only three species occur: a

blue species that resembles A. castaneum (as A. sp. aff.

castaneum in Fig. 1), A. elegans, and another undescribed

species (A. sp. Cuvette) that can occur sympatrically with

A. elegans. This species, with a characteristic dark dorsal

fin, has long been taken for A. e/egans in many publica-

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

Wildekamp, 1993). However, the description and origi-

nal drawing do not mention this dark dorsal fin that is very

prominent even in preserved specimens (Boulenger,

1899). Our DNA analysis shows that A. elegans and A.

sp. Cuvette are not closely related (Fig. 9). Additionally,

the specimens from Lui Kotale (Fig. 1) differ by male

colour pattern and are tentatively named A. sp. aff. e/e-

gans to indicate the differences and their potential status

as separate species.

Historical influcences on current pattern

The distribution pattern within the group is very complex.

Several species are found over large areas (Fig. 1), often

mixed with congeners. This complexity is probably relat-

ed to the history of the Congo Basin.

From our data it cannot be determined when Aphyosemion

reached the Congo Basin. The ancestors of the extant

species must have come from the west where their clos-

est relatives live. With the current data it is not possible

to decide if they were already dispersed around the en-

dorheic lake or if they entered the basin after a river cap-

ture to the west. Both scenarios will result in different dis-

tribution and dispersal patterns.

In later stages forest refuges probably played an impor-

tant role in the establishment of the mosaic distribution

pattern of Aphyosemion during glacial dry periods. Almost

all species of Aphyosemion s.|. are strictly bound to for-

est cover (Brosset, 1982; Kamdem Toham & Teugels,

1997, 1998, 1999). During glacial periods, the majority

of the forest in the Congo Basin was replaced by savan-

nah (see Leal, 2004; Maley, 1996). Only in the western

part of the basin a relatively large refuge was present from

where dispersal might have originated during more humid

periods. Leal (2004) proposed that besides larger well

known refuges also micro refuges must have existed, e.g.

in Gabon. A combination of one or more larger and sev-

eral micro refuges from where repeatedly dispersal could

have occurred might explain the very complex distribu-

tion pattern of Aphyosemion in the Congo Basin.

Prospect

Further DNA studies can contribute to alpha taxonomy and

towards a better understanding of the phylogeny and bio-

©ZFMK

Anew Aphyosemion from the eastern Congo 85

geography of this group and add to our understanding of

its evolution. However, much more samples from the huge

distribution area of Aphyosemion will be needed and the

occurrence of mitochondrial introgression makes it nec-

essary to include several nuclear markers into such a study

to get reliable and well supported results.

COMPARATIVE MATERIAL

Part of the comparative material is listed in Van der Zee

& Sonnenberg (2010); additional material is listed here

and in the Online Appendix:

Aphyosemion sp. Cuvette: MRAC 79-09-P-720-722, la-

belled as 4. christvi (Boulenger, 1915), Democratic Re-

public of Congo, Iteli River, terr. Opala, J. Lambert,

12.05.1958.

Aphyosemion sp.: MRAC 90-30-P-1471, labelled as A.

christyi (Boulenger, 1915), Democratic Republic of Con-

go, Yangambi, J. Lambert, 08.05.1957; MRAC 119855-

856, labelled as A. christyvi (Boulenger, 1915), Democrat-

ic Republic of Congo, Yangambi, J. Lambert, 09.05.1957.

Aphyosemion castaneum Myers, 1924: MRAC 22555-

22561, Democratic Republic of Congo, Stanleyville, Dr.

Richard, 1930; MRAC 89-043-P-547-612, labelled as A.

christyi (Boulenger, 1915), Democratic Republic of Con-

go, Libuku River near Kisangani, L. De Vos & M. Katem-

bo, April 1988.

Aphyosemion schoutedeni (Boulenger, 1920):

MRAC15664-15665, identified by David (1936) as Epi-

platys boulengeri, Democratic Republic of Congo, Med-

je, Lang & Chapin, 1910; MRAC 25529, Democratic Re-

public of Congo, Medje, H. Schouteden, no year.

Acknowledgements. The authors wish to thank the following

persons for their help in realizing this paper: J. Musafiri (Ubun-

du, Democratic Republic of Congo) for enabling the collection

of the new species; A. Van Deun (Leuven, Belgium) for the col-

lection of type material and several additional specimens that

were taken to Belgium alive and donated to H. Ott

(Monchengladbach, Germany), who photographed them; W.

Grell (Neustadt-Erlach, Germany) for the picture of the F1 off-

spring of A. musafirii; W. Eberl (Haubersbronn, Germany) for

providing notes about a collecting trip in the northern Demo-

cratic Republic of Congo by H. Bleher and W. Grell in 1986; E.

Vreven and J. Snoeks (MRAC, Tervuren, Belgium) for their kind

hospitality and support; M. Parrent (MRAC) for his services and

ad hoc help during research at the museum; T. Woeltjes (Nij-

megen, The Netherlands) for access to his private library and

for providing data of hobbyist collection trips; U. Schliewen

(ZSM, Munich, Germany), R. Numrich (Cologne, Germany),

and the members of the Aphyosemion elegans AG of the Ger-

man Killifish Association (DKG), for DNA samples and valu-

Bonn zoological Bulletin 60 (1): 73-87

able information. We also thank two anonymous reviewers, F.

Herder (ZFMK, Bonn, Germany), A. C. Schunke (MPI, Plon,

Germany), and B. Sinclair (Canadian National Collection of In-

sects, Ottawa, Canada) for comments and corrections on previ-

ous versions of the manuscript.

The Online Appendix 1s available at http://www.zfmk.de/

web/Forschung/Buecher/Beitraege/index.en.html

REFERENCES

Ahl E (1924) Uber neue afrikanische Zahnkarpfen der Gattung

Panchax. Zoologischer Anzeiger 60: 303-312

Amiet JL (1987) Le Genre Aphyosemion Myers (Pisces, Tele-

ostei, Cyprinodontiformes). Faune du Cameroun, Volume 2.

Sciences Nat, Compiegne, France

Boulenger GA (1899) Matérieaux pour la fauna du Congo. Pois-

sons nouveaux du Congo. Cinquieme Partie. Cyprins, Silures,

Cyprinodontes, Acanthoptérygiens. Annales du Musee du

Congo (Série Zoologie) Volume | (fasc. 5): 97-128, pls. 40-47

Boulenger GA (1910) Poisson Cyprinodonte nouveau du Bas-

Congo. Monatsberichte der Gesellschaft der Luxemburger Na-

turfreunde (Bulletins Mensuels de Société des Naturalistes

luxembourgeois) 4: 285

Boulenger GA (1911) Liste des poissons réprésentés dans une

nouvelle collection de la riviére Sankuru (Kasai), regue de M.

E. Luja. Monatsberichte der Gesellschaft der Luxemburger

Naturfreunde (Bulletins Mensuels de Société des Naturalis-

tes luxembourgeois) 5: 223-224

Boulenger GA (1915) Catalogue of the freshwater fishes of

Africa in the British Museum (Natural History) Volume 3,

London

Boulenger GA (1920) Poissons recueillis au Congo belge par

Vexpedition du Dr. C. Christy. Annales du Musée du Congo

belge: 1-38, pls 23-25

Brosset A (1982) Le peuplement de cyprinodontes du bassin de

V’Ivindo, Gabon. Terre Vie 36: 233-292

Brosset A, Lachaise D (1995) Evolution as a lottery conflicting

with evolution via sexual selection in African rainforest-

dwelling killifishes (Cyprinodontidae, Rivulinae, Diapteron).

Evolutionary Biology 28: 217-264

Collier GE (2007) The genus Aphyosemion: Taxonomic histo-

ry and molecular phylogeny. Journal of the American Killi-

fish Association 39: 147-168

Fowler HW (1936) Zoological Results of the George Vander-

bilt African Expedition of 1934. Part 3. The Freshwater Fish-

es. Proceedings of the Academy of Natural Science Philadel-

phia 88: 243-335, figs 102—152

Gustinicich S, Manfioletti G, Del Dal G, Schneider C, Caninci

P (1991) A fast method for high quality genomic DNA extrac-

tion from whole human blood. BioTechniques 11: 298-302

Hall TA (1999) BioEdit: a user-friendly biological sequence

alignment editor and analysis program for Windows 95/98/NT.

Nucleic Acids Symposium Series 41: 95—98

Hoedeman J (1958) The frontal scalation pattern in some groups

of toothcarps (Pisces: Cyprinodontiformes). Bulletin of

Aquatic Biology 1: 23-28

Huber J-H (1994) Aphyosemion passaroi, espéce inédite du

Gabon sud-oriental, au patron de coloration unique et descrip-

tion complémentaire de Aphyosemion decorsei (Pellegrin).

Cyprinodontiformes, Aplocheilidae. Revue frangaise d’ Aquar-

iologie 20: 77-80

©ZFMK

86 Jouke R. Van der Zee & Rainer Sonnenberg

Huber J-H (2004) Description of a new Aphyosemion species

from Congo, A. plagitaenium n. sp., exhibiting a probable in-

tra-generic color convergence with oblique bars. Freshwater

and Marine Aquarium 27: 70-74

Huber J-H (2005a) Identifikation einer kleinen Sammlung von

Aphyosemion aus Zaire im Mtinchener Museum (Z.S.M.), mit

weiteren Kommentaren tiber die Validitat der bekannten Kom-

ponenten der e/egans-Superspezies. |. Teil. Deutsche Killi-

fisch Gemeinschaft Journal 37: 8-21

Huber J-H (2005b) Identifikation einer kleinen Sammlung von

Aphyosemion aus Zaire im Miinchener Museum (Z.S.M.), mit

weiteren Kommentaren tiber die Validitat der bekannten Kom-

ponenten der e/egans-Superspezies. 2. Teil. Deutsche Killi-

fisch Gemeinschaft Journal 37: 35—45

Huber J-H (2007) Killi-Data 2007. Updated checklist of taxo-

nomic names, collecting localities and bibliographic references

of oviparous cyprinodont fishes (Atherinomorpha, Pisces). So-

ciéte fran¢aise d’Ichthyologie, Paris (German edition)

Huber J-H, Scheel JJ (1981) Revue systématique de la super-

espece Aphyosemion elegans. Description de A. chauchei n.

sp. et A. schioetzi n. sp. Revue frangaise d’Aquariologie 8:

33-42

Huelsenbeck JP, Ronquist F (2001) MRBAYES: Bayesian in-

ference of phylogeny. Bioinformatics 17: 754-755

Huson DH, Bryant D (2006) Application of Phylogenetic Net-

works in Evolutionary Studies. Molecular Biology and Evo-

lution 23: 254-267

Kamdem Toham A, Teugels GG (1997) Patterns of microhabi-

tat use among fourteen abundant fish species from the lower

Ntem River Basin in Cameroon. Aquatic Living Resources 10:

289-298

Kamdem Toham A, Teugels GG (1998) Diversity patterns of fish

assemblages in the Lower Ntem River Basin (Cameroon), with

notes on potential effects of deforestation. Archiv fiir Hydro-

biologie 141: 421-446

Kamdem Toham A, Teugels GG (1999) First data on an index

of Biotic Integrity (IBI) based on fish assemblages for the as-

sessment of the impact of deforestation in a tropical West-

African river. Hydrobiologia 397: 29-38

Kottelat M (1997) European freshwater fishes. An heuristic

checklist of the freshwater fishes of Europe (exclusive of for-

mer USSR), with an introduction for non-systematists and

comments on nomenclature and conservation. Biologia,

Bratislava, Section Zoology 52 (Supplement 5): 1-271

Kullmann H, Klemme B (2007) Female mating preference for

own males on species and population level in Chroma-

phyosemion killifishes (Cyprinodontiformes, Nothobranchi-

idae). Zoology 110: 377-386

Leal ME (2004) The African rain forest during the Last Glacial

Maximum, an archipelago of forests in a sea of grass. PhD

Thesis Wageningen University, The Netherlands

Maley J (1996) The African rain forest — main characteristics of

changes in vegetation and climate from the Upper Cretaceous

to the Quarternary. Proceedings of the Royal Society of Edin-

burgh 104B: 31-73

Meinken H (1951) Aphyosemion cognatum, eine neue Zahnkarp-

fenart aus dem Kongo Gebiet. Die Aquarien und Terrarien

Zeitschrift (DATZ) 4: 255-257

Moritz C, Patton JL, Schneider CJ, Smith TB (2000) Diversifi-

cation of rainforest faunas: an integrated molecular approach.

Annual Reviews in Ecology and Systematics 31: 533-563

Murphy WJ, Collier GE (1999) Phylogenetic Relationships of

African Killifishes in the Genera Aphyosemion and Fundu-

lopanchax inferred from Mitochondrial DNA Sequences. Mo-

lecular Phylogenetics and Evolution 11: 351-360

Bonn zoological Bulletin 60 (1): 73-87

Myers GS (1924) A new poeciltid fish from the Congo, with re-

marks on funduline genera. American Museum Novitates 116:

1-9

Pellegrin J (1904) Cyprinodontidés nouveaux du Congo et de

?Oubanghi. Bulletin du Muséum National d’Histoire Naturelle

10: 221-223

Plana V (2004) Mechanisms and tempo of evolution in the

African Guineo-Congolian rainforest. Philosophical Transac-

tions of the Royal Society, London, B 359: 1585-1594

Poll M (1951) Notes sur les Cyprinodontidae du Musée du Con-

go belge. Premiere partie: les Rivulini. Revue de Zoologie et

de Botanique Africaines 45: 157-171

Radda AC, Ptirzl E (1987) Colour Atlas of Cyprinodonts of the

Rain Forests of Tropical Africa; Hofmann Verlag, Wien

Scheel JJ (1968) Rivulins of the Old World. TFH Publications,

New Jersey

Scheel JJ (1990) Atlas of the Kullifishes of the Old World. TFH

Publications, New Jersey

Seegers L (1997) Killifishes of the World. Old World Killies I.

Aqualog, A.C.S. GmbH, M6rfelden-Walldorf

Sonnenberg R (2007) Description of three new species of the

genus Chromaphyosemion Radda, 1971 (Cyprinodontif-

ormes: Nothobranchiidae) from the coastal plains of

Cameroon with a preliminary review of the Chroma-

phyosemion splendopleure complex. Zootaxa 1591: 1-38

Sonnenberg R, Blum T (2005) Aphyosemion (Mesoaphyosemion)

etsamense (Cyprinodontiformes: Aplocheiloidei: Notho-

branchiidae), a new species from the Monts de Cristal, North-

western Gabon. Bonner zoologische Beitrage 53: 211—220

Sonnenberg R, Blum T, Misof B (2006) Description of a new

Episemion species (Cyprinodontiformes: Nothobranchiidae)

from northern Gabon and southeastern Equatorial Guinea.

Zootaxa 1361: 1-20

Sonnenberg R, Busch E (2010) Description of Callopanchax

sidibei (Nothobranchiidae: Epiplateinae), a new species of kil-

lifish from southwestern Guinea, West Africa. Bonn zoolog-

ical Bulletin 57: 3-14

Swofford DL (1998) PAUP*.Phylogenetic Analysis Using Par-

simony (*and Other Methods). Version 4. Sinauer Associates,

Sunderland, Massachusetts

Tamura K, Dudley J, Nei M, Kumar S (2007) MEGA4: Molec-

ular Evolutionary Genetics Analysis (MEGA) software ver-

sion 4.0. Molecular Biology and Evolution 24: 1596-1599

Thompson JD, Gibson TJ, Plewniak F, Jeanmougin F, Higgins

DG (1997) ClustalX windows interface: flexible strategies for

multiple sequence alignment aided by quality analysis tools.

Nucleic Acids Research 24: 4876-4882

Van Bergeijk WA, Alexander S (1962) Lateral line canal organs

on the head of Fundulus heteroclitus. Journal of Morpholo-

gy 110: 333-346

Van der Zee JR, Huber J-H (2006) Zur Identitaét von

Aphyosemion christyi (Boulenger, 1915), Aphyosemion

schoutedeni (Boulenger, 1920) und Aphyosemion castaneum

Myers, 1924 (Pisces, Cyprinodontiformes, Aplocheilidae).

Deutsche Killifisch Gemeinschaft Journal 38: 131-142

Van der Zee JR, Sonnenberg R (2010) Aphyosemion teugelsi

(Cyprinodontiformes: Nothobranchiidae), a new species from

a remote locality in the southern Democratic Republic of the

Congo. Zootaxa 2724: 58-68

Van der Zee JR, Woeltjes T, Wildekamp RH (2007) Aplochei-

lidae. Pp. 80-240 in: Stiassny MLJ, Teugels GG, Hopkins CD

(eds.) The Fresh and Brackish Water Fishes of Lower Guinea,

West-Central Africa. Volume II. IRD Editions, Paris

©ZFMK.

A new Aphyosemion from the eastern Congo 87

Wildekamp RH (1993) A World of Killies. Atlas of the Oviparous

Cyprinodontiform Fishes of the World. Volume |. The Amer-

ican Killifish Association, Mishawaka, Indiana

Woeltjes T (1984) Aphyosemion lefiniense spec. nov. (Pisces,

Atheriniformes, Rivulineae) aus dem Lefini Subsystem des

Kongo-Flusses in der Volksrepublik Kongo. Deutsche Killi-

fisch Gemeinschaft Journal 16: 3—14

Bonn zoological Bulletin 60 (1): 73-87

Zhang D-X, Hewitt GM (1996) Nuclear integrations: challenges

for mitochondrial DNA markers. Trends in Ecology and Evo-

lution 11: 247-251

Received: 18.01.2011

Accepted: 04.03.2011

Corresponding editor: F. Herder

©ZFMK

Bonn zoological Bulletin Volume 60

Issue | pp. 89-93 | Bonn, May 2011

Nothobranchius seegersi (Cyprinodontiformes: Nothobranchiidae),

a new annual killifish from the Malagarasi River drainage, Tanzania

Stefano Valdesalici! & Kiril Kardashev?

| Via Ca Bertacchi 5, I-42030 Viano (RE), Italy; E-mail: valdekil@tin.it

? Stramna 4, BG-2600 Dupnitsa, Bulgaria.

Abstract. A new annual killifish species, Nothobranchius seegersi, is described based on specimens collected in ephe-

meral water bodies in the Wulua River drainage system, Malagarasi River basin, central Tanzania. The new species be-

longs to the V. neumanni species group and is distinguished from N. neumanni by different male colouration and longer

predorsal length in males and females.

Key words. Nothobranchius neumanni, new species, taxonomy, seasonal pool, Tanzania.

INTRODUCTION

The killifish genus Nothobranchius occurs in the subtro-

pical and tropical parts of eastern Africa, from Sudan to

South Africa, and from Chad to Zanzibar and Mafia is-

lands in Tanzania. All known species are annual fishes,

living in temporary pools and swamps formed during the

rainy season (Wildekamp 2004). Six Nothobranchius spe-

cies are currently known from central Tanzania: N. neu-

manni from the Great Ruaha, Bubu, Wembere, and Ma-

lagarasi River basins, and Lakes Manyara and Victoria

drainage systems; NV. robustus from the Lake Victoria drai-

nage system; N. taeniopygus from the Wembere, Malaga-

rasi, and Bubu River basins and the Lake Victoria drai-

nage system; and at least three undescribed species: N.

spec. aff. newmanni “Malagarasi Type”, N. spec. aff. neu-

manni “Mbeya Type’, and N. spec. “Lake Victoria” (De

Vos et al. 2001; Seegers 1997; Shidlovsky 2010; Wilde-

kamp 1990, 2004).

Nothobranchius spec. aff. neumanni “Malagarasi Type”

and N. spec. aff. neumanni “Mbeya Type” share with N.

neumanni a unique combination of characters in male co-

lour pattern: the caudal fin is red or partially red, the pec-

toral fins are hyaline, the anal fin is yellow with red stri-

pes, its proximal portion light bluish grey. These synapo-

morphic colour characters are diagnostic for the N. neu-

manni species group.

In May 2008, the second author and Iva Ivanova (Dup-

nitsa, Bulgaria) collected during a Nothobranchius spe-

cies survey specimens of N. spec. aff. neumanni “Mala-

garasi Type” from small pools in the drainage system of

the Wulua River, which flows into the Limba Limba

River. Based on these specimens we herein describe Not-

hobranchius seegersi, new species.

MATERIAL AND METHODS

Measurements and counts were taken as described in

Amiet (1987), Huber (1992), and Valdesalici (2010). Mea-

surements were made with a digital calliper, partly under

a dissecting microscope, and rounded to the nearest 0.1

mm. If not stated otherwise, measurements are presented

as percentages of standard length (SL), except for eye dia-

meter and snout length, which are given as percentage of

head length (HL). Terminology for the cephalic neuromast

series follows Scheel (1968), for the frontal squamation

Hoedeman (1958). Osteological preparations (clearing and

staining, below: C&S) were made according to Taylor and

Van Dyke (1985), but not stained for cartilages. We con-

sider as Nothobranchius neumanni only the populations

from the type locality in the Bubu River drainage (see fig.

A50201-4 in Seegers 1997) and the geographically clo-

se Bahi Swamp area (see figs 5 a—b in Wildekamp 1990),

which were also studied by Wildekamp (1990) in the re-

description of the species. Morphological data from Wil-

dekamp (1990) were used here for comparisons. Numbers

in brackets following counts indicate the frequency of oc-

currence. Type material is deposited in the following in-

stitutions: Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK), Bonn, Germany, Museo Civico di Sto-

ria Naturale “Giacomo Doria” (MSNG), Genova, Italy,

and Royal Museum for Central Africa (MRAC), Tervu-

ren, Belgium.

90 Stefano Valdesalici & Kiril Kardashev

RESULTS

Nothobranchius seegersi, new species

(Figs 1—2, Table 1)

Nothobranchius spec. aff. neumanni: Seegers (1997), p.

75, figs A50205—4

Nothobranchius neumanni: Wildekamp (2004), p. 206, po-

pulation from south-west of Ipole.

Holotype. ZFMK 41848, 1 male, 44.5 mm SL, seasonal

pool on the right side of the main road T8 from Ipole to

Rungwa, near Mabangwe village, close by the bridge over

the Limba Limba River, altitude 1114 m, Malagarasi Ri-

ver drainage, Tanzania, 5°59’1” S, 32°48’5” E, 2 June

2008, Kiril Kardashev and Iva Ivanova.

Fig. 1.

Nothobranchius seegersi, adult male, not preserved,

Tanzania, seasonal pool on right side of the main road T8 from

Ipole to Rungwa, near Mabangwe village. Photo: Iva Ivanova.

Paratypes. ZFMK 41849- 41850, 2 females, 49.5 & 54.7

mm SL, same data as holotype; ZFMK 41851, 1 male,

45.9 mm SL, same data as holotype; MRAC 2010-33-P-

1, | male, 43.8 mm SL, same data as holotype; MSNG

56046, | male, 57.5 mm SL, C&S, same data as holoty-

pe; MRAC 2010-33-P-2-3, 2 males, 42.5 & 44.3 mm SL,

flooded grasslands on the right side of the main road T8

from Ipole to Rungwa, Mkola area, altitude 1107 m, Tan-

zania, 5°54°54” §, 32°45°54” E, Kiril Kardashev and Iva

Ivanova, 2 June 2008; MSNG 56045A-B, 2 males, 36.6

& 38.1 mm SL, large pool on left side of the main road

T8 from Ipole to Rungwa, near Ngoywa village, altitude

1114 m, Tanzania, 5°57’49” S, 32°46’50” E, Kiril Kar-

dashev and Iva Ivanova, 2 June 2008.

Non-type material. Private collection of the first author:

1 male, 62.00 mm SL, C&S, same data as holotype.

Diagnosis. Nothobranchius seegersi males share with the

other members of the V. neumanni species group a com-

bination of colouration characters, which distinguish them

from all other species of the genus: caudal fin red or par-

Bonn zoological Bulletin 60 (1): 89-93

Fig. 2.

Tanzania, seasonal pool on right side of the main road T8 from

Ipole to Rungwa, near Mabangwe village. Photo: Iva Ivanova.

Nothobranchius seegersi, adult female, not preserved,

tially red; pectoral fin hyaline; anal fin yellow with red

stripes, proximal portion light bluish grey. It is distinguis-

hed from N. neumanni by bright light blue scales (vs. pa-

le bluish grey), an irregular pattern formed by red scale

margins on lateral body and head (vs. uniform), and a sin-

gle, blue male colour morph (vs. two colour morphs, blue

and red). Male and female N. seegersi have a relatively

higher predorsal length compared to NV. neumanni (60.3—

68.8 (7) vs. 54.8-61.0 % (13) SL in males, 68.0—-68.4 (2)

vs. 57.1-64.6 % (24) SL in females). In osteological cha-

racters, Nothobranchius seegersi differs from N. neuman-

ni by having a conspicuously longer lateral process of the

post-temporal, and by having short antero-dorsal process

of the urohyal (examined in 2 C&S N. seegersi and 5 N.

neumannt).

Description. See Figs 1—2 for overall appearance and Ta-

ble 1 for morphometric data of the type series. Robust

Nothobranchius with rounded body, maximum length re-

corded in males 62.0 mm SL. Dorsal profile straight to

slightly concave on head, convex from nape to end of dor-

sal fin base. Ventral profile convex, slightly concave on

caudal peduncle posterior to dorsal and anal fin. Snout

slightly pointed, mouth directed upwards, lower jaw lon-

ger than upper, posterior end of rictus at same level as or

slightly above centre of eye. Branchiostegal membrane

projecting posteriorly from opercle. Dorsal and anal fin

located far posterior, tips rounded with short filamentous

rays. Both fins with papillate contact organs on fin rays.

Dorsal fin tip reaching caudal fin. Number of dorsal fin

rays 16-17, anal fin rays 17—18. Pectoral fin approxima-

tely triangular; in some specimens pectoral fins reach pel-

vic fins, and pelvic fins reach the anal fin origin. Caudal

fin rounded.

Scales cycloid, body and head entirely scaled, except for

ventral surface of head. Scales in median lateral series 28—

34 + 3—5 on caudal fin base. Cephalic squamation pattern

variable. Anterior neuromast series of the ‘open’ type.

Central supra-orbital series in shallow grooves, each with

©ZFMK

Nothobranchius seegersi, a new annual killifish from the Malagarasi River drainage, Tanzania 9]

Table 1. Morphometric data of Nothobranchius seegersi. Measurements except of standard length (in mm), eye diameter and head

length (in percent of head length) are percentages of, standard length. Cleared and stained material is not included.

Holotype Males (n = 7) Females (n = 2)

Standard length (mm) 44.5 36.6-45.9 49 .5—54.7

Body depth at pelvic fin 29.2 28.9-33.8 29.2-29.2

Predorsal length 62.2 60.3—68.8 68.0—68.4

Preanal length 58.2 57.4-66.8 69.2-69.6

Prepelvic length 48.0 47.2-56.3 56.6—57.3

Caudal peduncle length 21.5 18.0—23.3 17.1-18.9

Caudal peduncle depth IBZ 13.2-16.2 13.7-13.9

Head length 30.7 30:7-36.0 34.7-35.7

Snout length IWES 17.5—27.2 23.7-24.7

Eye diameter 27.0 22.2-27.0 18.9-22.0

two or three neuromasts. Posterior cephalic neuromast se-

ries curved with three or four neuromasts. One neuromast

on each scale of median longitudinal series. Basihyal bo-

ne sub-triangular. Six branchiostegal rays. Vomerine teeth

present in a large patch. Lateral process of post-temporal

long. Single short antero-dorsal process of urohyal. Num-

ber of vertebrae 31—32. Premaxilla and dentary with ma-

ny irregularly distributed unicuspid, slightly curved teeth

of different size, a small number of larger ones on the ou-

ter row of upper and lower jaw. Females are smaller than

males, maximum observed size 54.7 mm SL. In females,

the dorsal fin is rounded, the anal fin is triangular with

rounded tip. Branchiostegal membrane not projecting from

opercle.

Colour in life. Males. (Fig. 1) Scales on body and head

bright light blue with very thin and variable red margin,

creating an irregular reticulated pattern on body and head.

Branchiostegal membrane light blue with white margin.

Dorsal fin grey greenish to grey bluish with with irregu-

lar rows of dark red spots proximally, which become elon-

gate over fin rays distally. Anal fin bright yellow with light

blue base, separated by a dark red narrow stripe or row

of dark red irregular spots, and black margin. Some spe-

cimens with a median dark red narrow stripe or row of

dark red irregular spots in anal fin. A sub-distal light blue

narrow stripe present in anal fin of some specimens. Pel-

vic fins yellow with red spots. Pectoral fins hyaline with

light blue margin. Caudal fin dark red, with black margin

and light blue submargin. Iris golden, with faint black ver-

tical bar through centre of eye.

Females. (Fig. 2) Scales on body and head pale greyish

blue, with golden to light blue iridescence on scale cen-

tre. Opercular region silvery to golden. Abdomen silvery

to golden. All fins hyaline. Iris golden, with faint black

vertical bar through centre of eye.

Bonn zoological Bulletin 60 (1): 89-93

Colour in ethanol. Males. Scales on body light brown to

whitish, almost all scales on frontal and dorsal area with

distinct dark red margin, ventral scales with irregular dark

red spots. Dorsal fin whitish with dark red to brown spots.

Anal fin pale yellowish with light brown to whitish base,

separated by an irregular stripe formed by dark red to

brown spots, black margin. Some specimens with a me-

dian dark red to brown narrow stripe or row of dark red

irregular spots in anal fin. Pelvic fins yellowish with dark

red proximal spots and distal black spots. Pectoral fins

hyaline. Caudal fin dark red, with black margin and di-

stinct yellowish submargin. Iris bluish.

Females. Body light brown to whitish. Opercular and ven-

tral area yellowish. Unpaired and paired fins whitish. Iris

bluish.

Etymology. The species is named in dedication to its first

collector, the enthusiastic aquarist and ichthyologist Lo-

thar Seegers, Germany.

Distribution and habitat. Nothobranchius seegersi is cur-

rently only known from seasonal pools in the drainage sy-

stem of the Wulua and Mungu Rivers, Malagarasi River

basin, central Tanzania (Fig. 3). The type locality was at

the time of collection a small and very shallow pond, about

3 x 5m wide and 0.25 m deep, without aquatic vegetati-

on, in the open woodland (Fig. 4). The water was brown

and very turbid. No other fish species was found in this

pool.

DISCUSSION

Nothobranchius seegersi was collected for the first time

in 1992 for aquaristic purposes, but apparently no mate-

rial was deposited in public or institutional collections at

©OZFMK

92 Stefano Valdesalici & Kiril Kardashev

Fig. 3.

(black rhomboids, open black rhomboid: type locality), NV. neu-

manni (red rhomboids, open red rhomboid: type locality N. neu-

manni), and N. spec. aff. neumanni “Mbeya Type” (blue rhom-

boids). Map prepared by Béla Nagy.

Geographic distribution of Nothobranchius seegersi

that time. Until the new collections in 2008, only pictu-

res were available (Seegers 1997). Nothobranchius neu-

manni was diagnosed in Wildekamp (1990) by the straight

dorsal profile and marked, angular transition between head

and body along the dorsal profile. The dorsal profile is ho-

wever variable within NV. neumanni, so we did not use this

character for diagnosing the species described here.

Nothobranchius neumanni appears to be restricted to the

Bubu River drainage and the Bahi Swamps. There are in-

dications that the remaining populations so far assigned

to N. neumanni (namely from the areas of the Chipongo-

la, Manyara, Tabora, and Lake Victoria) differ substanti-

ally in terms of colouration, osteology, and genetics from

the Bubu drainage and Bahi Swamp material (prelimina-

ry data from A Dorn & A Cellerino, Jena, pers. comm.),

and might be considered as valid species; this is however

beyond the scope of the present paper. Tanzania is with

21 out of the 44 currently recognized species the hotspot

of Nothobranchius diversity (Wildekamp 2004), and furt-

her species descriptions from this region are to be expec-

ted.

Bonn zoological Bulletin 60 (1): 89-93

COMPARATIVE MATERIAL

Nothobranchius neumanni: MSNG 56051A-—B, 2 males,

52.7 & 57.3 mm SL; Seneki, Tanzania, 5°11’S, 33°17’ E;

MSNG 56052A-B, Sukamahela, Tanzania, 5°8’25” S,

32°46°44” E; 2 males, 37.9 & 47.8 mm SL. MSNG 56053,

1 male, 49,3 mm SL, C&S, Bahi Swamp — Lusilile TZ

2008-19, Tanzania, 5°53’S, 35°12’ E; first authors priva-

te collection: 1 male C&S, 49.7 mm SL, Manyara area,

Tanzania, 3°35’ S, 35°50’ E; 1 male C&S, 44.2 mm SL

Tabora area, Tanzania, 5°1’ S, 32°48’ E; 1 male, 54.1 mm

SL, Magiri, Tanzania, 4°55’ S, 33°1’ E; 2 males C&S, 47.8

& 52.8 mm, Bahi Swamp — Itigi, Tanzania, 05°53’ S,

35°12’ E; | male, 43.9 mm SL, Bahi Swamp-Lusilile, Tan-

zania, 5° 53’ S, 35°12’ E.

Fig. 4.

pool on right side of the main road T8 from Ipole to Rungwa,

near Mabangwe village, Tanzania.

Type locality of Nothobranchius seegersi, seasonal

Acknowledgements. We thank Francesca Fontana for the assi-

stance with the first draft of the manuscript, Aarn Aarn for pro-

viding C&S specimens, Béla Nagy for preparing the distributi-

on map, Iva Ivanova for fish pictures, Ian Sainthouse for cor-

rections of the language, Alexander Dorn and Alessandro Cel-

lerino for providing unpublished genetic data. Thanks to Rainer

Sonnenberg for improving the text and for valuable comments.

We are grateful to Fabian Herder and two anonymous reviewers

for comments, suggestion and critical analysis, and to Fabian

Herder (ZFMK), Giuliano Doria (MSNG), and Miguél Parrent

(MRAC) for access to material under their care.

REFERENCES

Amiet JL (1987) Faune du Cameroun. Vol. 2. Le genre Aphyo-

semion Myers (Pisces: Teleostei: Cyprinodontiformes). Sci-

ences Naturales, Compiegne

De Vos L, Seegers L, Taverne L, Thys Van Der Audenaerde D

(2001) L’ichtyofaune du basin de la Malagarasi (Systeme du

Lac Tanganyka) une synthése de la connaissance actuelle An-

©ZFMK

Nothobranchius seegersi, a new annual killifish from the Malagarasi River drainage, Tanzania 93

nales, Sciences Zoologiques, Musée Royal de I’ Afrique Cen-

trale 285: 117-135

Hoedeman JJ (1958) The frontal scalation pattern in some groups

of tooth carps (Pisces, Cyprinodontiformes). Bulletin of Aqua-

tic Biology 1: 23-28

Huber J-H (1992) Review of Rivulus. Ecobiogeography — Re-

lationships. Cybium, Paris

Scheel JJ (1968) Rivulins of the Old World. Tropical Fish Hob-

byist Publication, Neptune City

Seegers L (1997) Killifishes of the World. Old World Killis IT.

Verlag A.C.S. Morfelden-Walldorf

Shidlovsky K (2010) Collecting Nothobranchius in South-We-

stern Tanzania. Nothobranchius archives 1: 18-27

Taylor WR & Van Dyke GC (1985) Revised procedures for sta1-

ning and clearing small fishes and other vertebrates for bone

and cartilage study. Cybium 9: 107-109

Bonn zoological Bulletin 60 (1): 89-93

Valdesalici S (2010) Nothobranchius boklundi (Cyprinodonti-

formes: Nothobranchiidae): a new annual killifish with two

male colour morphs from the Luangwa River basin, Zambia.

Aqua, International Journal of Ichthyology 16: 51—60

Wildekamp RH (1990) Redescription of two lesser known Not-

hobranchius from Central Tanzania, N. taeniopygus and N.

neumanni (Cyprinodontiformes: Aplocheilidae). Ichthyologi-

cal Exploration of Freshwaters 1: 193-206

Wildekamp RH (2004) A world of killies. Atlas of the oviparous

cyprinodontiform fishes of the world. Vol. 4. The American

Killifish Association, Elyria

Received: 06.12.2010

Accepted: 14.04.2011

Corresponding editor: F. Herder

©ZFMK

caine

a A

be pia

Bonn zoological Bulletin Volume 60 Issue | | pp. 95-101 | Bonn, May 2011

Revisiting the species status of

Pecari maximus van Roosmalen et al., 2007 (Mammalia)

from the Brazilian Amazon

Jaime Gongora!, Cibele Biondo?, Jennifer D. Cooper?, Andrew Taber+, Alexine Keuroghlian>, Mariana

Altrichter®, Fabricia Ferreira do Nascimento’, Amanda Y. Chong!, Cristina Yumi Miyaki’,

Richard Bodmer?, Pedro Mayor!? & Susana Gonzalez!!

'Faculty of Veterinary Science, Gunn Building B19, University of Sydney, NSW 2006, Australia;

E-mail: jaime.gongora@sydney.edu.au

2Departamento de Ecologia, Instituto de Biociéncias, Universidade Estadual Paulista, Avenida 24-A 1515,

Rio Claro, SP, 13506-900, Brazil

3Department of Biological Science, Purdue University, West Lafayette, IN 47907 USA

4Center for International Forestry Research (CIFOR), P.O. Box 0113 BOCBD, Bogor 16000, Indonesia

SWildlife Conservation Society, Brazil, Rua Jardim Botanico 674, 22461-000, Rio de Janeiro, RJ, Brazil

6Environmental Studies Center, University of Redlands, 1200 East Colton Avenue, P.O. Box 3080,

Redlands, California, 92373, USA

7Genetics Division, Instituto Nacional de Cancer, Rua André Cavalcanti 37, 4° andar. Bairro de Fatima,

20231-050, Rio de Janeiro, RJ, Brazil

8Departamento de Genética e Biologia Evolutiva, Instituto de Biociéncias, Universidade de Sao Paulo, Rua do

Matao 277, Sao Paulo, SP 05508090, Brazil

°Wildlife Conservation Society, 2300 Southern Boulevard Bronx, New York 10460

10Pepartament de Sanitat 1 d’ Anatomia Animals, Facultat de Veterinaria, Universitat Autonoma de Barcelona,

Edifici V, E-08193 Bellaterra, Barcelona, Spain

‘Genetica de la Conservacion, Instituto de Investigaciones Biologicas Clemente Estable,

Facultad de Ciencias, Universidad de la Republica Oriental del Uruguay, Av. Italia 3318-11600. Montevideo,

Uruguay

Abstract. Three extant species of peccaries (Tayassuidae) are currently recognized and are distributed in the Americas:

Pecari tajacu, Tayassu pecari and Catagonus wagneri. The recent claim for a new peccary species, Pecari maximus, rais-

es questions about whether there is sufficient molecular or other evidence to support it as a new species. Here, we revis-

it that evidence using 91 published and 50 novel DNA sequences mainly from P. tajacu, as well as comment on the mor-

phological and ecological evidence used to support such a claim. Phylogenetic analyses show that the single specimen

of P. maximus used clustered within the South American lineage of P. tajacu closely related to Brazilian specimens. In

addition, the morphological and ecological data used to claim the species status of P maximus appears to be deficient

and inconclusive, in the light of available literature.

Key words. Collared peccary, control region, mitochondrial DNA, Pecari maximus, Pecari tajacu, Tayassuidae.

INTRODUCTION

Three extant species of peccaries (Tayassuidae) are cur-

rently recognized: Pecari tajacu (Collared peccary),

Tayassu pecari (White-lipped peccary) and Catagonus

wagneri (Chacoan peccary) (Wetzel et al. 1975; Wood-

burne 1968; Wright 1989, 1993). These species have been

identified according to morphological and cytogenetic pa-

rameters and more recently using DNA sequences. Clas-

sification has generally relied on biological concepts of

reproductive and genetic isolation. One confounding fac-

tor is that P. tajacu and T. pecari are known to hybridize

in captivity (Sowls 1997) and in the wild with the latter

being reproductively sterile (Andrea et al. 2001). Cyto-

genetic studies have confirmed the existence of three

species, showing significant chromosomal differences in

number and structure: P tajacu (2n=30), 7) pecari

(2n=26) and C. wagneri (2n=20) (Benirschke & Kumamo-

96 Jaime Gongora et al.

to 1989). Although chromosome painting across species

(Bosma et al. 2004) has provided some chromosomal evo-

lutionary insights into this family, a full understanding of

the species relationships from a cytogenetic perspective

remains hampered by a lack of information on C. wag-

neri. However, phylogenetic analyses of mitochondrial

and nuclear DNA sequences have revealed that T. pecari

and C. wagneri grouped in separate clades from P. tajacu

(Gongora & Moran 2005; Theimer & Keim 1998). These

conclusions obtained from molecular markers contrast

with cladistic analyses of morphological data which pro-

posed two different scenarios of relationships: First, P. ta-

jacu and C. wagneri are more closely related whereas 7:

pecari 1s considered to be a member of a separate clade

(Wright 1989, 1993, 1998); and second, P. tajacu and T.

pecari are more closely related to each other than to the

C. wagneri (Wetzel et al. 1975; Wetzel 1977).

The new species Pecari maximus. The recent claim of a

new peccary species by van Roosmalen et al. (2007) rais-

es questions about whether there is sufficient molecular

and/or other evidence to recognise P. maximus as a new

species. The morphological and genetic evidence used by

van Roosmalen et al. (2007) to support the species status

for P. maximus appears to be controversial. They analysed

a small number of specimens from deceased animals (five

skins and three skulls, two of which were from adults),

and the body weight (40—50 kg) and size of the animals

(total body length 127 cm; ear length 13 cm; shoulder

height 85 cm) were estimated from reports and photo-

graphs by local hunters. Morphometric analyses were al-

so estimated from skin measurements, and they relied on

pelage colour to point the differences with other species.

In this study, the mitochondrial control region and two nu-

clear SINE PRE-1 sequences of a single individual were

used to assess the phylogenetic position of P. maximus

among recognised species of peccaries. In addition, van

Roosmalen et al. (2007) used morphological and behav-

ioural data to support a separate status for those pecca-

ries. These authors argue that in contrast with the other

peccary species, which they suggest roam semi-nomadi-

cally ina highly variable landscape in a noisy herd, P.: max-

imus appears to walk silently through its habitat in small

family groups that contain a single adult pair with or with-

out 1-2 offspring. They also claim that P. maximus per-

forms little or no uprooting and has been seen feeding

mainly on freshly fallen fruits and seeds exposed on the

forest floor, which appears to be result of direct observa-

tions. The claim by van Roosmalen et al. (2007) under-

lines the urgent need to complete an ongoing modern tax-

onomic assessment of the family Tayassuidae. A revision

of the peccaries is imperative to orient conservation man-

agement and planning as well as to increase understand-

ing of the basic biology, ecology and evolution of this

group (Taber 1993).

Bonn zoological Bulletin 60 (1): 95-101

Differentiation within P tajacu. Highly relevant to the

claim by van Roosmalen et al. for a fourth species in the

genus Pecari 1s that previous phylogenetic mitochondri-

al DNA studies have shown genetic variation within Col-

lared peccary to be higher than that observed between C.

wageneri and T. pecari, and was also higher than that ob-

served between recognised species of the family Suidae

(Gongora & Moran 2005; Gongora et al. 2006). Pheno-

typic, morphological, chromosomal and DNA data have

been used to assess the levels of differentiation within P

tajacu. This lineage has a broad distribution in the sub-

tropical and tropical ecosystems of the Americas compris-

ing a wide variety of environments (rainforest, semi-arid

thorn forest, coastal forest, cloud forest, deserts, islands,

rangelands, scrublands, savannas, and freshwater wetland)

between the south-western United States of America and

northern Argentina (Grubb & Groves 1993). Its ancestry

also seems to have an early divergence from the other pec-

cary lineages in the Americas (Gongora et al. 2006). Ear-

ly morphological studies provided the first evidence of cra-

nial and dental variation between P. tajacu from through-

out the Americas (Kiltie 1985; Woodburne 1968), although

specimens were ultimately grouped into a single species.

Variations in size and pelage colour, coupled with distri-

bution data, have been the basis for proposing the exis-

tence of 14 subspecies of P. tajacu (Grubb & Groves 1993;

Hall 1981). However, the inheritance of these traits has

not been tested or substantiated by other methods.

Conventional cytogenetic (Builes et al. 2004; Gongora et

al. 2000; Vasart et al. 1994) and cross-species chromosome

painting (Adega et al. 2006; Bosma et al. 2004) studies

showed variation in the structure of two chromosomes be-

tween P. tajacu specimens from Arizona, Colombia and

Brazil, providing additional evidence for differentiation

within this species. A single captive specimen from

Colombia, heterozygous for a balanced translocation may

represent a hybrid between the two different P. tajacu lin-

eages (Builes et al. 2004). This may contrast with other

cytogenetic studies (Adega et al. 2006) which propose a

‘significant barrier’ for hybridization between major Col-

lared peccary lineages/species. However, germ line cyto-

genetic and/or reproductive studies are required to assess

whether this possible hybrid is fertile before any conclu-

sion can be addressed.

In this paper we revisit the genetic data from P. maximus

(van Roosmalen et al. 2007) using a DNA dataset of the

three peccary species generated by Gongora et al. (2005,

2006) and including 50 new DNA P. tajacu sequences

from United States, Mexico, Colombia and Brazil. We al-

so analyse the morphological and ecological evidence used

to support the species status of P. maximus in the light of

available literature.

©ZFMK

Revisiting the status of Pecari maximus 97

MATERIAL AND METHODS

Sampling and DNA extraction. Blood, tissue and hair

samples from 50 captive and wild P. tajacu from south-

eastern and central-western Brazil (BRAI—BRAS; n=5);

southern and northern Colombia (COL11—COL40; n=30);:

central and southern-central Mexico (MEX41—MEX48;

n=8); and Texas in the United States (USA6—USA 12; n=7)

were used in this study (Appendix I). DNA was extract-

ed using either a standard proteinase K and phenol-chlo-

roform protocol (Sambrook et al. 1989) or QIAamp DNA

mini kits (Qiagen Pty Ltd).

PCR and Sequencing. We used previously published

primers to amplify approximately 1,200 bp of the com-

plete mitochondrial control region (Kim et al. 2002) at the

University of Sydney, Australia. PCR was carried out us-

ing methods outlined in Gongora et al. (2005; 2006) ex-

cept for sequencing of forward and reverse strands which

were directly sequenced rather than clone inserts. Control

region sequences (450 bp) from the Texan specimens

(n=7) were independently generated at Purdue Universi-

ty. Primers, PCR and sequencing conditions were per-

formed as described in Cooper et al. (2010). Although

there is a difference in length between the above subsets

of sequences, the corresponding regions are informative

for assessing the major clades of P. tajacu described in pre-

vious studies (Gongora et al. 2006).

Data analysis. Forward and reverse sequences were over-

lapped in order to obtain a single double-stranded se-

quence for each animal using CodonCode Aligner v3.5.5

(CodonCode Corporation). After excluding flanking

primer regions, the novel sequences (~1,120 bp, GenBank

accession numbers HM102371 through HM102419)

were aligned using the program Muscle (version 3.6;

Edgar, 2004) along with those P tajacu sequences

(AY546539—AY 546569) published by Gongora et al.

(2005; 2006) and the single sequence (DQ009006) from

P. maximus published by van Roosmalen et al. (2007). The

control region from 7. pecari (AY546516—AY 546519) and

C. wagneri (AY 546520, AY546521) were included as an

outgroup. The alignment used for analyses described con-

sisted of 1,140 bp after excluding the tandem repeat mo-

tifs, except for a single motif, due to heteroplasmy accord-

ing to Gongora et al. (2006). Because few gaps were ob-

served they were kept in the alignment.

A separate dataset of concatenated nuclear PRE-1 642 and

PRE-1 27 sequences was generated and aligned as de-

scribed above. It consisted of available sequences from

GenBank (P. maximus, DQ016372 and DQ016371; P. ta-

Jacu, DQ190931—DQ190883, AY569340—AY 569339, and

AY568052—AY568047; 7. pecari, AY546331 and

AY546528; and C. wagneri, AY546530 and AY546527).

Bonn zoological Bulletin 60 (1): 95—101

The best fit DNA substitution model for the control region

and PRE-1 alignments were selected by the program Mod-

elgenerator (version 0.85; Keane et al. 2006) and imple-

mented in Maximum Likelihood (ML) to reconstruct a

phylogenetic tree using the software PhyML (version 3.0;

Guindon & Gascuel 2003). The tree topology space was

searched by using the best of Nearest Neighbour Inter-

change and Subtree Pruning and Regrafting starting from

five random starting trees generated by BioNJ ( Guindon

& Gascuel 2003; Guindon et al. 2010). Branch support

was calculated using the approximate likelihood ratio test

(aLRT) with SH-like interpretation, as it is as conserva-

tive and accurate test compared with bootstrapping but less

computationally intensive (Anisimova & Gascuel 2006;

Guindon et al. 2010).

RESULTS AND DISCUSSION

Clustering of P. maximus within recognised major clades

of P. tajacu. Novel and published Collared peccary mito-

chondrial control region sequences representing 44 hap-

lotypes clustered in two major clades: North/Central

American clade and South American clade (Fig. 1) as de-

scribed by Gongora et al. (2006). The novel P. tajacu se-

quences from Brazil clustered within the South American

clade closely related to two sequences from Argentina,

while those from Texas clustered within the North/Cen-

tral American clade closely related to specimens from Ari-

zona, Mexico and Colombia. Sequences from Colombian

specimens split between the two major clades consistent

with previous studies of being a paraphyletic group

(Gongora et al. 2006). The current results corroborate that

the Gongora et al. (2006) dataset is suitable for assessing

major subcontinental geographical origins of captive and

wild specimens.

Unexpectedly for a sequence from what is considered as

a different species, Pecari maximus clustered within the

major South American clade closely related to central-

western Brazilian and northern Argentinean sequences

supported by high aLRT values (Fig. 1) rather than in a

separate new clade. Consistent with control region analy-

ses, ML tree of PRE-1 sequences shows that those from

P. maximus cluster within the P. tajacu closely related to

sequences from Bolivia, Colombia and the United States

(data not shown). The clustering of Pecari maximus with-

in a previously recognized South American clade of P. ta-

Jacu (Gongora et al. 2006) do not support the new species

status of those specimens studied by van Roosmalen et al.

(2007). Although there is no agreement as to which par-

ticular sequences accurately and universally reflect the

species’ boundaries (Vogler & Monaghan 2007), other mo-

lecular markers, such as cytochrome b or cytochrome c

oxidase / are usually recommended as being more inform-

©OZFMK

COL10

0.92

0.86

Fig. 1.

Jaime Gongora et al.

North/Central America

P. tajacu

South America

Outgrup

Maximum Likelihood tree of the mitochondrial control region sequence representing forty-four haplotypes observed in

eighty P. tajacu specimens, using both C. wagneri (CHP) and 7! pecari (WLP) sequences as outgroups. Brackets indicate the two

major clades found in P. tajacu with Colombian (COL) specimens showing paraphyletic relationships. Pecari maximus clustered

within P. tajacu closely related to those specimens from Brazil (BRA) and Argentina (ARG). Details of samples used in this study

are described in Appendix 1. Numbers close to branches are aLRT values. For clarity, only aLTR values for the main internal groups

higher than 0.80 are shown.

ative to assess genetic species distinction (Blaxter 2004;

Blaxter et al. 2005; Bradley & Baker 2001). Phylogenet-

ic conclusions based on a single specimen are insufficient

and uncertain. Under the current DNA evidence it is like-

ly that single specimens studied by van Roosmalen et al.

(2007) correspond to one of the P. tajacu lineages iden-

tified by Gongora et al. (2006). Species status has come

and gone for some putative taxa within the distantly re-

lated suid family. For instance, the possible extinct S. buc-

Bonn zoological Bulletin 60 (1): 95-101

culentus (Indochinese or Vietnam Warty pig) was initial-

ly considered as a separate species within the genus Sus

(Groves 1997), but after DNA analyses showed that its se-

quences clustered closely with S. scrofa (domestic pig and

wild boar), it has been suggested that it could correspond

to a geographically restricted variant of S. scrofa rather

than a distinct species (Mona et al. 2007; Robins et al.

2006).

©OZFMK

Revisiting the status of Pecari maximus 99

Comments on the morphological, behavioural and eco-

logical evidence of P. maximus. The body weights report-

ed for PR. maximus (van Roosmalen et al. 2007) are with-

in the range reported for P. tajacu, albeit at the large end.

For instance, Sowls (1997) reports exceptional weights of

wild Collared peccaries in Arizona in excess of 42 kg, and

Bodmer (1989) reports weights close to 40 kg in Peru.

Bodmer et al. (pers. obs.) have observed considerable vari-

ation in the skull sizes of both P. tajacu and T. pecari from

the Tahuayo, Yavari and Tamshiyacu regions of north-east-

ern Peru, well within the range reported by van Roosmalen

et al. (2007). These preliminary observations are based on

the collection held at the zoology museum of the Univer-

sidad Nacional de la Amazonia Peruana (UNAP), which

consists of several thousand skulls collected by local sub-

sistence hunters over a 10 year period. In addition, Bod-

mer et al. (per. obs.) have recorded body weights from both

P. tajacu and T. pecari hunted by local people in the

Yavari river valley. Again, there is considerable variation,

and the body weights are well within the range reported

by van Roosmalen (Bodmer pers. obs.). Comparing 7.

pecari with the proposed new species, the former may sim-

ilarly reach head-body lengths over 120 cm and weights

as high as 40 kg in the Brazilian Pantanal (Keuroghlian

et al. 2006) and 50 kg in the northern Amazon (Fragoso

1998). It would be desirable that skull measurements of

the proposed P. maximus be statistically compared with

those from extremely large P. tajacu found elsewhere in

their range. Further, using skin pelage characteristics to

distinguish species can be problematic since bristle colour

may differ substantially within peccary species even in the

same area (Gongora et al. 2006). In Collared peccaries,

their lighter-haired collar pattern may vary from distinct

in some individuals to barely noticeable in others. The

most compelling morphological lines of evidence present-

ed by van Roosmalen et al. (2007) are three photographs

from three individuals of exceptionally long-legged pec-

caries which appear different from peccaries we are fa-

muiliar with, but these are not backed up by measurements.

Ecological and behavioural differences have also been

used to characterize peccary species (Sowls 1997). van

Roosmalen et al. (2007) have attributed a number of dif-

ferentiating traits for their peccary in line with this rea-

soning. However these claims are not based on intensive

field observations, nor do they draw on recent literature

concerning the ecology and behaviour of both P. tajacu

and 7. pecari to inform contrasts. For instance, while they

claim that the new peccary uniquely lives in adult pair

groups, sometimes with young, P. tajacu is also known

to forage alone or in small groups, even while belonging

to herds typically of six to twelve individuals (Keurogh-

lian et al. 2004; Sowls 1997; Taber et al. 1994), although

larger groups have been reported from the Amazon

(Fragoso 1994). Furthermore, the number of individuals

Bonn zoological Bulletin 60 (1): 95-101

seen together may depend on the time of day. Radio

telemetry studies have shown that 7. pecari has distinct

seasonal movements that are related to habitat and fruit

availability within their large home ranges, and P. tajacu

has relatively small stable home ranges (Altrichter et al.

2001; Fragoso 1998; Keuroghlian et al. 2004; Keurogh-

lian & Eaton 2008; Sowls 1997). Neither species can be

described as roaming semi-nomadically as van Roosmalen

et al. (2007) has suggested. Instead they commonly for-

age on freshly fallen fruits (Altrichter et al. 2000, 2001;

Beck 2005; Keuroghlian & Eaton 2008). Therefore, these

are not unique behaviours of P- maximus. Whether pec-

caries need to dig with their noses during foraging depends

on the type of habitat where fruits are available, i.e. buried

within the sediments of a swamp or exposed on the for-

est floor (Keuroghhan & Eaton 2008). With only three

skulls attributed to the new species, sample size is too low

to comment on tooth wear patterns. Also, the lack of scent

from the new peccary skins may be related to the skill of

the hunters (that remove scent glands during butchering;

Keuroghlhian pers. obs.). We note that subjectively there

seems to be a considerable range between species and in-

dividuals with Chacoan peccaries having the least intense

scent which may contribute to them being harder for the

dogs of local hunters to track (Taber et al. 1993). Final-

ly, observations are too limited, and overly dependent on

local hunter hearsay, to draw conclusions that this possi-

ble new species has abandoned social groupings, group

defence and territorial scent marking.

The possible discovery of a new peccary species from the

Amazon basin is very exciting, and plausible, consider-

ing recent discoveries of new mammal species in this re-

gion mainly by van Roosmalen et al. (1998, 2000, 2002,

2003). However, based on the scant information report-

ed by van Roosmalen et al. (2007), and the results of our

own genetic analysis, we conclude that there is not enough

evidence to support the claim of a new species, and that

the specimens studied by van Roosmalen are most likely

P. tajacu. We also consider that further DNA, cytogenet-

ic and morphological research is necessary to better un-

derstand peccary taxonomy. Additional studies using oth-

er genetic molecular markers, cytogenetic, and morpho-

logical analyses incorporating new samples from all across

the Amazon basin, will provide the basic information for

understanding the genetic and morphology variation un-

der the extant recognized species.

Acknowledgements. This study was funded by the University

of Sydney International Program Development Fund to J.

Gongora to support collaboration between the University of Syd-

ney (Australia) and the Universidade de Sao Paulo (Brazil).

Brazilian peccary samples were sourced under permit IBAMA

13601 from Instituto Brasileiro do Meio Ambiente e dos Recur-

sos Naturais Renovaveis and imported into Australia under per-

mits AQIS IP07011173 and CITES, 09BR002827/DF. Other pec-

©ZFMK

100 Jaime Gongora et al.

cary samples were sourced from wild and captive specimens col-

lected for other studies from J. Gongora and J.D. Cooper. We

thank Peter Waser and J. Andrew DeWoody for their support to

J.D. Cooper to access the Collared peccary sequences from

Texas. We also would like to thank all the institutions that col-

laborated in obtaining samples listed in Appendix I.

REFERENCES

Adega F, Chaves R, Kofler A, Krausman PR, Masabanda J,

Wienberg J, Guedes-Pinto H (2006) High-resolution compar-

ative chromosome painting in the Arizona Collared peccary

(Pecari tajacu, Tayassuidae): a comparison with the karyotype

of pig and sheep. Chromosome Research 14: 243-251

Altrichter M, Saenz J, Carrillo E, Fuller T (2000) Dieta estacional

de Tayassu pecari (Artiodactyla: Tayassuidae) en el Parque

Nacional Corcovado, Costa Rica. Revista de Biologia Trop-

ical 48: 689-702

Altrichter M, Carrillo E, Saenz J, Fuller T (2001) White-lipped

peccary (Zayvassu pecari, Artiodactyla: Tayassuidae) diet and

fruit availability in a Costa Rican rain forest. Revista de Bi-

ologia Tropical 49: 1183-1192

Andrea MV, Oliveira C, Rocha GT, Foresti F (2001) Cytogenet-

ical and histological studies in testis of Tayvassu tajacu (Cate-

to), Zavassu pecari (Queixada) and a natural interspecific hy-

brid. Journal of Animal Breeding and Genetics 118: 125-133

Anisimova M, Gascuel O (2006) Approximate likelihood-ratio

test for branches: a fast, accurate, and powerful alternative.

Systematic Biology 55: 539-552

Beck H (2005) Seed predation and dispersal by peccaries

throughout the Neotropics and its consequences: a review and

synthesis. Pp. 77-115. In Seed Fate: Predation, Dispersal and

Seedling Establishment (P-M. Forget, J. E. Lambert, P.E.

Hulme and S. B. Vander Wall, eds.).CABI Publishing,

Wallingfort

Benirschke K, Kumamoto AT (1989) Further studies on the chro-

mosomes of three species of peccary. Pp. 309-316 in Ad-

vances in Neotropical Mammalogy (K.H. Redford and J.F.

Eisenberg, eds.). Sandhill Crane Press, Gainesville

Blaxter ML (2004) The promise of a DNA taxonomy. Philosoph-

ical Transactions of the Royal Society B: Biological Sciences

359: 669-679

Blaxter M, Mann J, Chapman T, Thomas F, Whitton C, Floyd

R, Abebe E (2005) Defining operational taxonomic units us-

ing DNA barcode data. Philosophical Transactions of the Roy-

al Society B: Biological Sciences 360: 1935-1943

Bodmer RE (1989) Frugivory in Amazonian ungulates. Ph.D.

dissertation, Clare Hall, Cambridge University, England

Bosma AA, de Haan NA, Arkesteijn GJ, Yang F, Yerle M, Ziyl-

stra C (2004) Comparative chromosome painting between the

domestic pig (Sus scrofa) and two species of peccary, the Col-

lared peccary (Zayassu tajacu) and the White-lipped peccary

(IZ. pecari): a phylogenetic perspective. Cytogenetic and

Genome Research 105: 115-121

Bradley RD, Baker RJ (2001) A test of the genetic species con-

cept: cytochrome-b sequences and mammals. Journal of Mam-

malogy 82: 960-973

Builes FDP, Loaiza IDD, Lopez JBO (2004) Evidencia de es-

tructuracioOn cromosomica asociada con la distribucion geo-

grafica de pecari de collar (Zayassu tajacu). Proceedings: VI

Congreso Internacional sobre Manejo de Fauna Silvestre en

la Amazonia y Latinoamérica 2004, September 5 — 10, Iqui-

tos, Peri: 152-157

Cooper JD, Vitalis R, Waser PM, Gopurenko D, Hellgren EC,

Gabor TM, DeWoody JA (2010) Quantifying male-biased dis-

Bonn zoological Bulletin 60 (1): 95-101

persal among social groups in the collared peccary (Pecari ta-

jacu) using analyses based on mtDNA variation Heredity 104:

79-87

Edgar, RC (2004) MUSCLE: multiple sequence alignment with

high accuracy and high throughput. Nucleic Acids Res. 32:

1792-1797

Fragoso JMV (1994) Large mammals and the community dy-

namics of an Amazonian rain forest. Ph.D. dissertation. Uni-

versity of Florida, Florida

Fragoso JMV (1998) Home range and movement patterns of

White-lipped peccary (Zayassu pecari) herds in the Northern

Brazilian Amazon. Biotropica 30: 458-469

Gongora J, Bernal JE, Fajardo LC, Moran C, Nicholas F, Mon-

salve H, Gardeazabal J, Orjuela D, Carrillo L, Ramirez M,

Borrero LM, Cristancho L, Cardenas A, Lasso R, Roldan J,

Quimbaya M, Olaya JC (2000) Mayores estudios citogeneti-

cos de peccaries de Collar colombianos. El Astrolabio 2: 6—9

Gongora J, Moran C (2005) Nuclear and mitochondrial evolu-

tionary analyses of Collared, White-lipped, and Chacoan pec-

caries (Tayassuidae). Molecular Phylogenetics and Evolution

34: 181-189

Gongora J, Morales S, Bernal JE, Moran C (2006). Phylogenet-

ic divisions among Collared peccaries (Pecari tajacu) detect-

ed using mitochondrial and nuclear sequences. Molecular Phy-

logenetics and Evolution 41: 1-11

Groves CP, Schaller GB, Amato G, Khounboline K (1997) Re-

discovery of the wild pig Sus bucculentus. Nature 386: 335

Grubb P, Groves CP (1993) The Neotropical Tayassuids Tayas-

su and Catagonus. Pp. 5—7 in Pigs, Peccaries and Hippos Ac-

tion Plan (W. L. R. Oliver, ed.). IUCN, The World Conserva-

tion Union, Gland

Guindon S, Gascuel O (2003) A simple, fast, and accurate al-

gorithm to estimate large phylogenies by maximum likelihood.

Systematic Biology 52: 696-704

Guindon S, Dufayard J, Lefort V, Anisimova M, Hordijk W, Gas-

cuel O (2010) New algorithms and methods to estimate max-

imum-likelihood phylogenies: Assessing the performance of

PhyML 3.0. Systematic Biology 59: 307-321

Hall ER (1981) The Mammals of North America. J. Wiley and

Sons, New York.

Keane TM, Creevey CJ, Pentony MM, Naughton TJ, McIner-

ney JO (2006) Assessment of methods for amino acid matrix

selection and their use on empirical data shows that ad hoc

assumptions for choice of matrix are not justified. BMC Evo-

lutionary Biology 6: 29

Keuroghlian A, Eaton DP (2008) Fruit availability and peccary

frugivory in an isolated Atlantic forest fragment: effects on

peccary ranging behavior and habitat use. Biotropica 40:

62-70

Keuroghlian AD, Eaton P, Freitas TP (2006) Are peccaries sex-

ually dimorphic? Proceedings: VII Congreso Internacional so-

bre Manejo de Fauna Silvestre na Amazonia e America Lati-

na 2006, September 3—7. Universidade Estadual de Santa

Cruz, Ilhéus — Bahia. Brazil

Keuroghlian AD, Eaton P, Longland WS (2004) Area use by

White-lipped and Collared peccaries (Tayassu pecari and

Tayassu tajacu) in a tropical forest fragment. Biological Con-

servation 120: 411-425

Kiltie RA (1985) Craniomandibular differences between rain-

forest and desert Collared peccaries. American Midland Nat-

uralist Journal 113: 384-387

Kim KI, Lee JH, Li K, Zhang Y_P, Lee SS, Gongora J, Moran

C (2002) Phylogenetic relationships of Asian and European

pig breeds determined by mitochondrial DNA D-loop se-

quence polymorphism. Animal Genetics 33: 19-25

OZFMK

Revisiting the status of Pecari maximus 101

Robins JH, Ross HA, Allen MS, Matisoo-Smith E (2006) Sus

bucculentus revisited. Nature 440: E7

Sambrook J, Fritsch EF, Maniatis T (1989) Molecular Cloning:

a Laboratory Manual, 2nd edn. Cold Spring Harbor Labora-

tory Press, New York

Sowls LK (1997) Javelinas and other Peccaries: Their Biology,

Management, and Use, 2nd Edition. Texas A&M University

Press, College Station, Texas

Taber AB, Doncaster CP, Neris NN, Colman FH (1993) Rang-

ing behavior and population-dynamics of the Chacoan pec-

cary, Catagonus wagneri. Journal of Mammalogy 74:

443-454

Taber AB, Doncaster CP, Neris NN, Colman F (1994) Ranging

behaviour and activity of two sympatric peccaries, Catagonus

wagneri and Tayassu tajacu, in the Paraguayan Chaco. Mam-

malia 58: 61-71

Taber AB (1993) The Chacoan peccary (Catagonus wagneri).

Pp. 22-29 in Peccaries, Pigs, and Hippos Action Plan (W. L.

R. Oliver, ed). IUCN, The World Conservation Union, Gland

Theimer TC, Keim P (1998) Phylogenetic relationships of the

peccaries based on cytochrome 6 DNA sequences. Journal of

Mammalogy 79: 566-572

van Roosmalen MGM, Frenz L, Van Hooft P, De Iongh HH,

Leirs H (2007) A new species of living peccary (Mammalia:

Tayassuidae) from the Brazilian Amazon. Bonner zoologische

Beitrage 55: 105—112

van Roosmalen MGM, van Roosmalen T, Mittermeier RA, De

Fonseca GAB (1998) A new and distinctive species of mar-

moset (Callitrichidae, Primates) from the lower Rio Aripuana,

State of Amazonas, central Brazilian Amazonia. Goeldiana Zo-

ologia 22: 1-27

van Roosmalen MGM, van Roosmalen T, Mittermeier RA, Ry-

lands AB (2000) Two new species of marmoset, genus Cal-

lithrix Erxleben, 1777 (Callitrichidae, Primates) from the Tapa-

jos/Madeira interfluvium, south central Amazonia, Brazil.

Neotropical Primates 8: 2-18

van Roosmalen MGM, van Roosmalen T, Mittermeier RA (2002)

A taxonomic review of the titi monkeys, genus Callicebus

Thomas, 1903, with the description of two new species, Cal-

licebus bernhardi and Callicebus stephennashi, from Brazil-

ian Amazonia. Neotropical Primates 10: 1—52

van Roosmalen MGM, van Roosmalen T (2003) The descrip-

tion of anew marmoset genus, Callibella (Callitrichinae, Pri-

mates), including its molecular phylogenetic status. Neotrop-

ical Primates 11: 1-10

Vassart M, Pinton A, Seguela A, Dutertre C (1994) New data

on chromosomes of peccaries. Mammalia 58: 504-507

Vogler AP, Monaghan MT (2007) Recent advances in DNA tax-

onomy. Journal of Zoological Systematics and Evolutionary

Research 45: 1-10

Wetzel RM, Dubos RE, Martin RL, Myers P (1975) Catagonus,

an “extinct” peccary alive in Paraguay. Science 189: 379-381

Bonn zoological Bulletin 60 (1): 95-101

Wetzel RM (1977) The extinction of peccaries and new case of

survival. Annals of the New York Academy of Sciences 288:

538-544

Woodburne MO (1968) The cranial myology and osteology of

Dycotyles tajacu, The Collared peccary, and its bearing on

classification. Memoirs of the Southern California Academy

of Science 7: 1-8

Wright DB (1989) Phylogenetic relationships of Catagonus wag-

neri: sister taxa from the Tertiary of North America. Pp.

281-308 in Advances in Neotropical Mammalogy (J. F. Eisen-

berg and K. H. Redford, eds). Sandhill Crane Press,

Gainesville

Wright DB (1993) Evolution of sexually dimorphic characters

in peccaries (Mammalia, Tayassuidae). Paleobiology 19:

52-70

Wright DB (1998) Tayassuidae. Pp. 389-400 in Evolution of Ter-

tiary Mammals of North America (C. M. Janis, K. M. Scott,

and L. L. Jacobs, eds). Cambridge University Press, Cam-

bridge, United Kingdom

Appendix I.

Origin of Collared peccary samples

Argentina: North, Ciudad de Rio Cuarto ARG1-2; Ola-

varria, ARG3-4; Bolivia: North, La Paz, Mallasa Zoo,

BOL 1-2. Brazil: State of Amazonas, PMAX; South East,

State of Sao Paulo, BRA1—4; Central West, State of Ma-

to Grosso do Sul, BRAS. Colombia: Central, Santa Cruz

Zoo, COL1; Jaime Duque Zoo, COL2; North Central, Ma-

tecana Zoo, COL4, COL20—22; Santafe Zoo, COL5;

North West, Barranquilla Zoo, COL6, COL30—-40; South

Central, La Lagartija Zoo, COL7; La Lagartija Zoo,

COL8; Cali Zoo, COL9; South East, Macagual Farm,

COL10, COL23-29; Central, Santa Cruz Zoo, COL11-19;

Mexico: Central, Guadalajara Zoo, MEX1-—2; North,

Puebla Zoo, MEX3-4, MEX41-42; South Central, Me-

xico DF Zoo, MEX5—8, MEX43-48. USA: Arizona,

USAI1-3; Texas, USA4—-12. Unknown: Adelaide Zoo,

UNK1-2.

Received: 23.10.2010

Accepted: 25.01.2011

Corresponding editor: R. Hutterer

©OZFMK

Bonn zoological Bulletin Volume 60

Issue | pp. 103-107 Bonn, May 2011

Two new species and a new record

of the genus Neolindus Scheerpeltz, 1933

(Coleoptera: Staphylinidae: Paederinae)

Ulrich Irmler

Institute of Ecosystem Research, Dept. Applied Ecology, University, Olshausenstrasse 40, D-24098 Kiel, Ger-

many; E-mail: uirmler@ecology.uni-kiel.de; phone: +49 431 880 4311.

Abstract. The two new species Neolindus verhaaghi from Peru and Neolindus pastazae from Ecuador are described. A

new record from central Amazonia of Neolindus densus Herman, 1991, is presented.

Key words: Paederinae, new species, Ecuador, Peru, new record, central Amazon.

Resumen: Las dos especies nuevas Neolindus verhaaghi de Peru y Neolindus pastazae de Ecuador estan descrito. Uno

lugar nuevo de Amazonica Central de Neolindus densus Herman, 1991, esta presentado.

Palabras clave: Paederinae, especie nueva, Ecuador, Peru, lugar nuevo, Amazonica Central.

INTRODUCTION

The genus Neolindus is restricted to Central and South

America and distributed from Bolivia and southern Brazil

in the south to Costa Rica in the north. Ecuador is the most

species-rich country with 10 species. The first species was

described by Sharp (1876) as Lindus religans Sharp, 1876.

Later, Scheerpeltz (1933) renamed the genus Neolindus,

since the name Lindus was already preoccupied. Irmler

(1981) added five new species from Brazil and Peru, and

Herman (1991) published a review of the genus, added 27

species, and published a cladistic analysis of the genus to-

gether with Cylindroxystus Bierig, 1943. Thus, the genus

contained 33 species in the Neotropical region.

In 2009, I collected a new species of the genus in the Rio

Pastaza basin, Ecuador. Earlier, I had found a further new

species in the collection of my colleague Manfred Ver-

haagh, Natural History Museum in Karlsruhe, Germany.

Inferred from the low number of specimens found, all

species seem to be very rarely collected. Most species have

been described by only one specimen. The two new

species are also known by only one specimen. Overall, 72

specimens have been collected in the Neotropics. There-

fore, a new record of NV. densus Herman, 1991 found near

Manaus in the central Amazon basin is also published,

here.

MATERIAL AND METHODS

The new material is deposited in my private collection

(UIC).

For the photographs, a Makroskop M 420 (Wild, Her-

brugg) was used in combination with a digital camera

(Nikon D100). Head length was measured from anterior

edge of clypeus to posterior edge of head disc, pronotum

along the midline from anterior edge to posterior edge, ely-

tra from anterior edge of shoulders to posterior edge; width

was measured at the widest part of tagmata. In the meas-

urements of total length, the abdominal inter-segmental

space is subtracted.

RESULTS

Neolindus verhaaghi, new species

(Figs 1 A—H)

Holotype. Male; Peru; Huanuco, Panguana, 150 km SW

Pucallpa, tropical rain forest, pitfall trap, 2—16.7.1984, leg.

M. Verhaagh, #pWA65 (UIC).

Diagnosis. The species is attributed to the group of N. agi-

lis Herman, 1991 and N. densus Herman, 1991. The cen-

tral emargination of the sth sternite [Fig. | F] is similar

104 Ulrich Irmler

Fig. 1. Neolindus verhaaghi; antennae (A), aedeagus in lateral (B) and ventral (C) aspect, 7“ and 8¢" visible tergites (D), 6! vis-

ible tergite (E) and sternite (F), dorsal aspect of head and pronotum (G) and elytra (H) showing punctation and surface (scale bar

0.5 mm).

New Neolindus from South America 105

to N. densus Herman, 1991 [Fig. 131 in Herman 1991] and

N. cephalochymus Herman, 1991 [Fig. 139 in Herman

1991]. Antennomere 2 [Fig. | A] is slightly shorter than

3, whereas it is slightly longer in N. densus and N.

cephalochymus. Furthermore, the penultimate anten-

nomeres in N. densus are wider than they are long, but

more or less quadrate in N. verhaaghi. As in N. agilis [Fig.

147 in Herman 1991], N. verhaaghi has no dense patch

of setae on the 8" sternite [Fig. | F]. The apical emargina-

tion on the 8" sternite is deeper in N. verhaaghi than in

N. agilis, and the aedeagus has an apical cavity [Fig. | B,

C] which is absent in all related species.

Description. Length: 4 mm. Colour: red, legs, antennae

and posterior edge of tergites yellow. Head: 0.50 mm long,

0.55 mm wide; eyes as long as temples, temples rounded

in smooth curve without forming angles; extremely fine

micro-punctation and without microsculpture; surface

shiny; one trichobothrium and one setiferous puncture at

front edge of eyes; two setiferous punctures on disc bet-

ween eyes [Fig. | G]; distance between these punctures

slightly wider than between upper edge of eye and adja-

cent puncture; transverse row of punctures at posterior

edge of vertex, and two diagonal rows of punctures be-

tween eyes and posterior edge of vertex. Antennae as long

as head and pronotum; 3" antennomere slightly longer

than 274 antennomere [Fig. | A]; 4** antennomere slight-

ly longer than wide; following antennomeres more or less

quadrate, pubescent and with few apical setae. Pronotum:

0.65 mm long, 0.60 mm wide; with longitudinal row of

11-12 punctures on each side of smooth midline [Fig. 1

G]; several punctures on laterad to paramedial row of

punctures; distance between these punctures at least as

wide as diameter of punctures; surface without mi-

crosculpture, polished and shiny. Elytra: 0.75 mm long,

0.65 mm wide; with sutural row of 11—12 punctures and

three more rows on disc [Fig. | H]; two rows of irregu-

lar punctures laterad to disc. Abdomen densely punctate;

anterior segments distinctly more densely punctate than

posterior segments; 8'" tergite apically rounded [Fig. 1 E];

8th sternite [Fig. 1 F] with triangular emargination, cen-

tral impunctate stripe, and without clusters of setiferous

punctures. Aedeagus with apical cavity and lateral trun-

cate prominences behind cavity [Fig. 1 B, C].

Etymology. The specific name refers to the collector of

the species, Manfred Verhaagh, from the Natural History

Museum in Karlsruhe, Germany.

Neolindus pastazae, new species

(Figs 2 A—I)

Holotype. Male; Ecuador, Tungurahua, 10 km W of

Banos, valley of Rio Pastaza, path near waterfall “Man-

Bonn zoological Bulletin 60 (1): 103-107

to de la Novia” (78°20.16W, 1°24.12S), sifting litter,

28.7.2009, leg. U. Irmler (UIC).

Diagnosis. Neolindus pastazae is certainly closely relat-

ed to N. punctogularis Herman, 1991, as determined by

the transverse cluster of numerous setae at the apical edge

of the gula. It can be easily differentiated from N. punc-

togularis by the darker colour of head and pronotum and

the different punctation of pronotum [Fig. 2 H]. The last

abdominal tergites and sternites [Fig. 2 E, F, G] are very

similar to N. punctogularis Herman, 1991 [Figs 205—207

in Herman 1991]. Besides the development of the trans-

verse row of numerous setae at the anterior edge of the

gula, both species can be differentiated from N. schubar-

ti Irmler, 1981, N. religans Sharp, 1876, and N. bidens

Herman, 1991, since the latter are carinate on 8th stern-

ites or have a ridge at the posterior edge [Figs 211, 220,

223 in Herman 1991].

Description. Length: 10.5 mm. Colour: brown, antennae,

palpae and legs yellow. Head: 0.90 mm long, 1.5 mm

wide; with eyes slightly prominent [Fig. 2 H]; temples 1/4

as long as eyes; temples obtusely angled to posterior edge,

outer part of posterior edge transversely narrowed to neck;

labrum with pair of apically rounded denticles near the

middle, with five setae at anterior edge; disc with fine mi-

cro-punctation; surface polished; two large punctures on

each side of vertex between eyes; distance between these

punctures wider than between each puncture and adjacent

eye; few setiferous supraocular punctures; central punc-

ture with trichobothrium; row of setiferous punctures

along posterior edge; supraocular punctures and posteri-

or row of punctures smaller than two large punctures be-

tween eyes, but much larger than micro-punctures; gula

with transverse row of numerous setae near anterior mar-

gin. Antennae with first antennomere elongate and as long

as antennomeres 2 and 3 combined [Fig. 2 A]; 2™4 anten-

nomere short, only 1/3 as long as 3" antennomere; fol-

lowing antennomeres elongate and decreasing in length;

antennomeres 3 to 11 pubescent. Maxillary palp at 2™4 seg-

ment with several long setae at inner side. Pronotum: 1.35

mm long, 1.65 mm wide; with sides more or less paral-

lel in anterior half [Fig. 2 H]; in posterior half, smoothly

rounded to posterior edge without forming angle; margin

continuing from posterior edge to anterior edge and cov-

ered by anterior angles in dorsal aspect; disc polished and

shiny; irregular row of 11 to 12 punctures on each side of

smooth midline; two transverse rows laterad to parame-

dial rows of punctures and few scattered punctures later-

ad. Elytra: 1.95 mm long, 1.85 mm wide; surface polished

and shiny; with irregular rows of dense punctures [Fig. 2

I]; average distance between punctures distinctly narrow-

er than half diameter of punctures; partly coriaceous. Ab-

domen densely and deeply punctate; anterior tergites more

densely punctate than posterior tergites; 8" tergite trilobed

©OZFMK

106 Ulrich Irmler

ea ‘i 7 AI ry J

Fig. 2. | Neolindus pastazae; antennae (A), aedeagus in lateral (B) and ventral (C) aspect, 5‘ visible sternite in ventral aspect

(D), 78 and 8th visible tergites (E), 6'" visible tergite (F) and sternite (G), dorsal aspect of head and pronotum (H) and elytra (I)

showing punctation and surface (scale bar 0.5 mm).

Bonn zoological Bulletin 60 (1): 103-107 OZFMK

New Neolindus from South America 107

with triangular central prominence and with striate struc-

tures at posterior edge [Fig. 2 F]; 7" sternite with semi-

circular emargination at posterior edge [Fig. 2 D]; 8" ster-

nite with deep central emargination and glabrous central

stripe [Fig. 2 E]. Aedeagus slightly asymmetric, with two

long bifurcate prominences apically and one pair of hooks

in apical half [Fig. 2 B, C]; ventral surface with deep cav-

ity in paramedial position.

Etymology. The specific name derives from the location

in the valley of the Rio Pastaza where it was found.

Neolindus densus Herman, 1991

New record. Brazil, Amazonas, 15 km SW Manaus, on

Ilha de Marchanteria (59°55.21 W, 3°13.59S), inundation

forest in Varzea, tree eclector #50E, 1 male, 1 female,

18.2.1982, leg. J. Adis (UIC).

DISCUSSION

Together with the two new species the total number of Ne-

olindus species increased to 35 species with Ecuador (11)

and Peru (7) as countries with high numbers of species.

Only Brazil, which has a much larger area than these two

countries, has a similarly high number with nine species.

The species have been very rarely collected. As N. den-

sus Herman, 1991 shows, the distribution can neverthe-

less cover a wide area. This species seems to occur along

the Amazon valley from its mouth near Belém to the An-

dean foot hills near Leticia (Columbia). A more detailed

analysis of the ecology is difficult. According to the in-

formation given by Herman (1991) many species have

been found in leaf litter of rain forests. In some labelled

information the habitat was described as “under felled

tree”. The habitat of N. pastazae was also in litter layer

under a felled tree. The collection of N. densus Herman,

1991 by a tree eclector in the central Amazon basin sug-

gests that the tree habitat might be a more important habi-

tat than can be derived from the labelled information.

Thus, soil litter layer might be only an accidental habitat

of the normally inhabited tree trunks, which are rarely in-

vestigated.

Bonn zoological Bulletin 60 (1): 103-107

According to Herman (1991), the Neolindus species can

be separated more or less in three species groups which

are closer related. N. verhaaghi can be attributed to clade

(17) with N. punctventris, N. densus, N. agilis, N.

cephalochymus, N. bullus, N. hamatus, N. procarinatus,

and N. retusus. This group is characterised by the pres-

ence of parallel carinae at the base of sternum VII. The

species of this group are distributed over the whole equa-

torial region, from the lowland rain forest of the Amazon

basin to the western rainforest in Ecuador. Neolindus pas-

tazae seems to be related to clade (24) with N. punctogu-

laris, N, hangarthi, N. schubarti, N. bidens, and N. reli-

gans. In this group, species are characterised by a prono-

tum wider than it is long and tergum VIII being trilobed.

Whereas N. religans and N. schubarti are distributed in

eastern Brazil, N. bidens, H. hanagarthi, and the new N.

pastazae represent a western branch of this group.

Acknowledgements. Dr. Manfred Verhaagh (Natural History

Museum in Karlsruhe, Germany) and Prof. Joachim Adis (for-

merly Max-Planck Institute, Dep. Tropical Ecology, Plén, Ger-

many) gratefully gave me their Neolindus material for the study

and for the deposition in my collection.

REFERENCES

Herman L (1991) Revision of the subtribe Cylindoxystina

(Coleoptera: Staphylinidae: Paederinae). Bulletin of the

American Museum of Natural History 203: 1-83

Irmler U (1981) Neue Arten der neotropischen Gattung Neolin-

dus Scheerpeltz (Coleoptera, Staphylinidae). Studies on

Neotropical Fauna and Environment 16: 209-215

Scheerpeltz O (1933) Coleopterorum Catalogus. Staphylinidae

VI Supplementum I. W. Junk, Berlin, 989-1500

Sharp D (1876) Contributions to an insect fauna of the Amazon

valley. Coleoptera — Staphylindae. Transactions of the Ento-

mological Society of London 1876: 27-424

Received: 03.05.2010

Accepted: 20.10.2010

Corresponding editor: D. Ahrens

©ZFMK

Bonn zoological Bulletin

Volume 60 | Issue | | pp. 109-111 | Bonn, May 2011

A remarkable record of Phaneroptera falcata (Poda, 1761)

(Saltatoria: Phaneropteridae)

from north-eastern Poland

Wolfgang Bohme*, Peter Geissler & Philipp Wagner

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

“ Corresponding author: E-mail: w.boehme.zfmk@uni-bonn.de.

Abstract. A record of Phaneroptera falcata is made from the northeastern Poland, extending the known distribution in

this country for about 300 km to the north and linking it with a newly discovered occurrence in southern Lithuania. The

northern margin of the species’ range in eastern Central Europe and in East Europe is briefly reviewed.

Phaneroptera falcata (Poda, 1761) belongs to those fau-

nal elements which underwent a rapid northward-direct-

ed range extension on the northwestern edge of their dis-

tribution areas, particularly in NW Germany (Northrhine-

Westphalia) and the Benelux countries (Detzel 1998). A

similar trend was observed in Thuringia (Kohler, 2010),

and even the federal states of Lower Saxony and Bran-

denburg have already been reached (Grein 2000, 2007,

Landeck et al. 2005). In the eastern part of its Central Eu-

ropean range, however, P. falcata was believed, despite

some expansional trend in the Czech Republic (Koéarek

et al. 2008), to be restricted to the southern half of Poland,

as indicated by e.g. the sketch maps in Detzel (1998) and

Maas et al. (2002). However, a recent record in southern-

most Lithuania (Lazdijai) close to the Polish northeastern

border (Ivinskis & Rimsaite 2008) suggests a much more

northern distribution also in Poland.

Southern Poland (Galicia) had already been mentioned in

the classic work by Harz (1957) as that part of this coun-

try which is inhabited by P. falcata. Even much earlier,

Zacher (1917) had pointed on a doubtful voucher speci-

men of the Wroclaw (formerly Breslau) Museum from

“Schlesien” (= Silesia), but he claimed that Pylnov (1913)

had already recorded this species more northernly of Wro-

claw, viz. from “Nova Alexandria, Russisch Polen” (=

“Russian Poland”). This place, today Pulawy (51.26N;

21.59E) can be found on older maps as situated between

Radom and Lublin, on the Visla river south of Deblin:

51.34N; 21.50E. Several new Polish references give a

more detailed and also extended picture of the distribu-

tion range of P. falcata in Poland (Bazyluk & Liana 2000,

Koéarek 2000, Orzechowski 2009). The locality data pub-

lished by these authors document this species to be more

widespread in the central part of Poland, the most north-

western records (Lubuskie Province: Orzechowski 2009)

being adjacent to the relatively new records from Bran-

denburg (Landeck et al. 2005). The two northern Polish

regions Pojezierze Pomorskie and Mazurskie which to-

gether roughly comprise the northern third of the coun-

try, were so far lacking Phaneroptera falcata records. The

new and unexpected find of this species in southern

Lithuania (Ivinskis & Rimsaite 2008), however, made the

occurrence of P. falcata likely also in northern Poland .

On August 13, 2010 two of us (WB & PW) passed through

northeastern Poland towards Lithuania. On road no. 16

east of Wigierski National Park, between Serski Las vil-

lage and Sejny, at 53.55N and 23.09E, when searching for

lizards on a spruce forest clearing, we happened to find

an adult female of P. falcata (Fig. 1) which in view of what

is said above seems to be a remarkable record, as it ex-

Fig. 1.

The voucher specimen of Phaneroptera falcata from

E of Wigierski National Park, Northeast Poland. Photograph: P.

Geissler.

110 Wolfgang Bohme et al.

Fig. 2.

Habitat east of Wigierski National Park where our P.

falcata record was found. Photograph: W. Bohme.

tends the known range within Poland for about (appr.) 300

km northeastwards and immediately links it with the first

Lithuanian record from 2008. The locality (Fig. 2) 1s sit-

uated less than 20 km from the Lithuanian border and

closely corresponds to the new and single Lithuanian lo-

cality of this thermophilous species which is situated in

the Lazdiyai district at 54.12N and 23.50E (Ivinskis &

Rimsaite 2008) (Fig. 3). Our specimen is deposited in the

Orthoptera collection of the Zoologisches Forschungsmu-

seum A. Koenig (ZFMK) in Bonn.

According to Zuna-Kratky et al. (2009), adults in Austria

appear about on mid-July but start their main adulthood

season from mid-August. This agrees with our female

voucher specimen, which was also adult, despite its much

more northernly situated locality.

Road no. 16 from Augustow via Serski Las and Sejny runs

in parallel to the main road (no. 8, via Suwalki to Mar-

jampole in Lithuania) and has much less traffic than the

latter. This alone makes it unlikely that the specimen of

P. falcata could have been passively displaced by human

transportation. Rather, this thermophilous species demon-

strated its potential for a natural, northeastward-directed

range extension not only in Germany and the Benelux

countries with a predominantly oceanic climate, but also

in NE Poland, under much more continental climatic con-

ditions and even reached Lithuania. The several individ-

uals registered there underline the existence of a popula-

tion in this area rather than displaced single individuals.

Intensive faunistic search is necessary to assess the dis-

tribution range and its dynamics of Phaneroptera falca-

ta in this region.

The eastward continuation of the northern borderline of

P. falcata through Belarus and the European part of the

Russian Federation also needs more faunistic research ef-

forts. According to the map in Willemse (2007), P. falca-

Bonn zoological Bulletin 60 (1): 109-111

ta is not yet known from Belarus. In Ukraine, the exact

northern borderline has yet to be assessed but runs, accord-

ing to Kotenko (in litt., November 2010), between 52 and

53N. In the European part of the Russian Federation, it is

marked by localities situated approximately on a similar

latitude as the formerly known Polish findings, e.g. Sevsk

(Fig. 3: 12), Kursk (Fig. 3: 13), and Lipetsk (Fig. 3: 14)

in the southern part of the Ryazan area (Bey-Bienko 1954).

A newly collected voucher specimen from the southern

part of the Bryansk area close to the Ukrainian border

(Chukrai village, district of Suzemka, 52.19N; 34.05E,

collected by one of us (PG) in July, 2010 and also deposit-

ed in ZFMK’s Orthoptera collection), roughly fits this dis-

tributional pattern (Fig. 3: 11), which ranges between 51

and 51.40N in the Asiatic part of Russia (Bey-Bienko

1954, 1964). It can be assumed, however, that P. falcata

will extend its northern distribution borderline also in these

parts of its range.

Fig. 3. | Map showing the northern part of the distribution

range of P. falcata in the eastern Central and East European

realm, our new record (2) and the one from Lithuania (1) being

the northernmost sites. 1. Lazdijai District (LIT); 2. Wigierski

National Park (PL); 3. Treuenbrietzen (D); 4. Berkenbriick (D);

5. Lubuski Province (PL); 6. Wroclaw (PL); 7. Nizina San-

domierska (PL); 8. Roztocze National Park; 9. Pulawy (PL); 10.

Polesie National Park (PL); 11. Chukrai (RUS); 12. Sevsk

(RUS); 13. Kursk (RUS); 14. Lipezk (RUS). Map: P. Wagner.

Acknowledgements. The authors are indebted to Prof. Dr. Zbig-

niew Szyndlar and to Dr. Elsbieta Warschowska (both from

Krakow), as well as to Ryszard Orzechowski (Zielona Gora), and

to Tatiana Kotenko (Kiev), for valuable advice and literature ref-

erences. PG thanks Igor Palko (Moscow) for the invitation to

Chukrai village.

OZFMK

A remarkable record of Phaneroptera falcata (Poda, 1761) from north-eastern Poland 111

REFERENCES

Bazyluk, W & Liana A (2000) Prostoskrzydle Orthoptera. Kat-

alogu fauny Polski 58: 1-156

Bey-Bienko GY (1954) Fauna SSR. Pryamokrylye, Moskva,

Leningrad (Izd. Akad,. Nauk SSSR), vol. 2, no. 2:

Kuznechikovye, podsem. Listobye kuznechiki (Phaneropteri-

nae): 394 pp.

Bey-Bienko GY (1964) Opredelityel nasekomykh Evropeskey

chasti SSSR. Tom 1. Moskva, Leningrad (Nauka): 935 pp.

Detzel P (1998) Die Heuschrecken Baden-Wiirttembergs. Stutt-

gart (Ulmer): 580 pp. + 20 pls

Grein G (2000) Zur Verbreitung der Heuschrecken (Saltatoria)

in Niedersachsen und Bremen., Stand 10.4.2000. Informati-

onsdienst Naturschutz Niedersachsen 20: 74-112

Grein G (2007) Zur Ausbreitung von Phaneroptera falcata (Po-

da, 1761) und Conocephalus fuscus (Fabricius, 1793) in Nie-

dersachsen. Articulata 22: 91—98

Harz K (1957) Die Geradfltigler Mitteleuropas. Jena, G. Fischer:

494 pp.

Ivinskis P & Rimsaite J (2008) Phaneroptera falcata (Poda,

1761) (Orthoptera, Phaneropteridae) in Lithuania. Acta Zoo-

logica Lituanica 18: 270-272

Koéarek P (2000) Orthopteroid insects (Orthoptera, Blattaria,

Dermaptera) of the Polesie National Park and its surroundings.

Parki i Rezerwaty Przyrody 19: 89-97

Koéarek P, Holusa J, VIk R, Marhoul P & Zuna-Kratky T (2008)

Recent expansion of the bush-crickets Phaneroptera falcata

and Phaneroptera nana (Orthoptera: Tettigoniidae) in the

Czech Republic. Articulata 23: 67—75

Bonn zoological Bulletin 60 (1): 109-111

Kohler G (2010) Fauna der Heuschrecken (Ensifera et Caelife-

ra) des Freistaates Thiiringen. Naturschutzreport Jena 17: 378

pp.

Maas S, Detzel P & Staudt A (2002) Gefahrdungsanalyse der

Heuschrecken Deutschlands. Verbreitungsatlas, Gefahrdungs-

einstufung und Schutzkonzepte. Bonn (BfN): 401 + xvi pp.

Orzechowski R (2009) Obserwacje wybranych gatunkow pro-

stoskrzydlych (Orthoptera) w poludniowe} czesci woje-

wodztwa Lubuskiego. Przeglad Przyrodnicyyzy 20: 45-50

Pylnov E (1913) Materialy po faunye pryamokrylych (Orthoptera

saltatoria) russkoj Polshi. Russkoye entomologichnoye

obozryenie, St. Peterburg 13: 85—98

Willemse F (2007) Fauna Europaea: Ensifera. In: Heller KG

(ed.): Fauna Europaea, version ileshs

http://www. faunaeur.org.2007

Zacher F (1917) Die Geradfliigler und ihre Verbreitung. Jena,

G. Fischer: 287 pp.

Zuna-Kratky T, Karner-Ranner E, Lederer E, Braun B, Berg, H-

M, Denner M, Bieringer G, Ranner A, Zechner L (2009): Ver-

breitungsatlas der Heuschrecken Ostésterreichs. Wien (Natur-

historisches Museum): 303 pp.

Received: 17.01.2011

Accepted: 23.02.2011

Corresponding editor: F. Herder

©ZFMK

oo et

i Sy

, Bratt i} my _

: 7 TE oC Thi te

ae ’ aa i

' ion ‘ « i

1 ‘Ay i 1

1 Pah F fT

I ;

( ; we

: io ee

Bon zoological Bulletin Volume 60

| Issue | p. 113 Bonn, May 2011

Erratum to:

Sonnenberg & Busch (2010) Description of Callopanchax sidibei

(Nothobranchiidae: Epiplateinae), a new species of killifish

from southwestern Guinea, West Africa

Rainer Sonnenberg !.2 & Eckhard Busch 3

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

Corresponding author; E-mail: r.sonnenberg.zfmk@uni-bonn.de

2 Max-Planck-Institut fiir Evolutionsbiologie, August-Thienemann-Strasse 2, D-24306 Plén, Germany

3 Diederichsstrasse 45, D-42855 Remscheid, Germany

Erroneously the name of the new species was formed as

a noun in genitive case for a single male person (ICZN,

Article 31.1.2), however, in the text it refers to:

Sex. The new species is named after Mr. Samba Sidibe and

his family.... (Sonnenberg & Busch 2010, p. 11, bold in-

troduced here for clarification).

Therefore the species name should be formed in plural

with the ending -orum.

In accordance with the International Code of Zoological

Nomenclature (4th Edition, 1999), Article 32, we propose

as correct species name Callopanchax sidibeorum.

Acknowledgements. We thank Wolfgang Béhme (ZFMK,

Bonn) for alerting us on this mistake in the publication and his

advice for this Erratum.

REFERENCES

International Code of Zoological Nomenclature, 4‘ edition,

1999. International Trust for Zoological Nomenclature, Lon-

don

Sonnenberg R, Busch E (2010) Description of Callopanchax

sidibei (Nothobranchiidae: Epiplateinae), a new species of kil-

lifish from southwestern Guinea, West Africa. Bonn zoolog-

ical Bulletin 57: 3-14

114 Book Review

Brandt T, Jiilch C, Wasmer K, Moning C, Wagner C (2011)

The Top 50 sites for birdwatching in Germany. (Die 50

besten Vogelbeobachtungsplatze in Deutschland). Der

Falke. Aula-Verlag, Wiesbaden. ISBN 978-3-89104-746-

QO: 19:95'€:

This book is published as a special issue of one of the lead-

ing German birdwatcher’s magazines, “Der Falke”, and

large parts of its contents have already been published in

single articles of the journal. It presents the 50 best sites

for birdwatching in Germany. Each single chapter gives

an overview about typical habitats and interesting bird

species and some logistical advices of the where and when

of birdwatching at the spot. Sometimes, interesting addi-

tional information on other cultural or natural study ac-

tivities in the region is given. Boxes show “How to get

there” and give useful local contact addresses. In contrast,

explicitly stated GPS coordinates are often not very help-

ful as they regularly do not show the particular spots from

where to find the birds (e.g. a hide) but the center of the

next village. A register at the end of the volume shows

which rare species occur at each site. A Top 50 list of sites

can always be debated. However, it is a pity that some

species which are much sought after are not covered by

the selected sites (e.g. Rock Bunting) or are not mentioned

at suitable sites (e.g. Cirl Bunting). Nice photos and maps

make it a pleasure to browse in the well illustrated book

and to think about the next short trip. However, if you are

seriously interested in birdwatching in Germany, the three

volume book “Vogel beobachten in Nord-, Siid- und Ost-

deutschland” published by Kosmos and written by mem-

bers of the same author team might be a better and more

comprehensive (but more expensive) choice. Germany still

lacks a concise English “Where to watch birds” guide, and

authors and editors sadly failed to fill this gap. Instead,

recycled material was used to write a nice although not

really essential book for birdwatchers.

Bonn zoological Bulletin 60 (1): 114

Der Falke ur ster

Darius Stiels

Zoologisches Forschungsmuseum Alexander Koenig,

Bonn

©OZFMK

Bonn zoological Bulletin (BzB)

Instructions to authors

Scope

The Bonn zoological Bulletin (BzB), formerly “Bonner zoologi-

sche Beitrage”, is an international, peer-reviewed, open access jour-

nal publishing original research articles, reviews, and scientific notes

dealing with organismal zoology. Focus of the BzB are (1) taxon-

omy, (2) systematics and evolution, and (3) biodiversity and bio-

geography, all with respect to terrestrial animals. Terrestrial animals

as understood here include those inhabiting fresh or brackish wa-

ters. Contributions from related fields like ecology, morphology,

anatomy, physiology or behaviour are welcome when of clear rel-

evance to the focus topics.

Publication in BzB is free of charge, including colour illustrations

or photographs contributing significantly to quality and / or read-

ability of the manuscript. Authors retain full copyright of their pub-

lished papers, may share them with colleagues, and are encouraged

to post the original pdfs on their personal or institutional website

for non-commercial use. All material must be original, unpublished

work and not under consideration for publication elsewhere.

Research articles: Manuscript preparation and submission

Manuscripts should be written in English. For serving readers from

the organism’s country of origin, bilingual abstracts are welcome.

If you plan to submit a manuscript of more than 50 manuscript pages,

please contact the editor-in-chief in advance. Please submit your

manuscript as electronic file (see paragraph on electronic submis-

sion below). Manuscripts should strictly follow the instructions spec-

ified below. Manuscripts not compatible with these guidelines will

not be taken into consideration.

Format Typing should be double-spaced in 12 pt throughout the text,

including tables, figure legends, and reference list. Pages and lines

should be numbered consecutively. Short subheadings may struc-

ture the text into sections. Do not number headings or subheadings.

Structure Research articles should be organized as follows: Title,

Author(s), Address(es) including email address and telephone, Ab-

stract, Key words, Running Title, Introduction, Material and Meth-

ods, Results, Discussion (or combined Results and Discussion), Con-

clusions (optional), Acknowledgements, References, Tables, Figure

Captions, Figures (separately numbered and identified), Appendices

(if necessary). Footnotes should not be used except in Tables.

Title The title should be brief (30 words limit, ideally shorter) and

indicate clearly the field of study and group of animals investigat-

ed. The systematic position of taxa listed in the title must be indi-

cated (e.g. “Squamata: Colubridae’’). A Running Title (maximum 45

characters inclusive of spaces) should describe the paper’s core top-

ic.

Abstract Each manuscript should contain a concise abstract as sin-

gle paragraph (max. 200 words) summarizing the significant find-

ings, followed by not more than six key words.

References In the text, cited sources should be referred to as fol-

lows: Ax (2001), Kim & Lumaret (1989), Holm et al. (1997) — for

three or more authors, or, if cited consecutively: (Ax 2001, 2002;

Holm et al. 1997; Kim & Lumaret 1989) — please follow exactly

the use of commas and semicolons. Do not use commas between

author and year (only exception: taxonomic names, in which a com-

ma is placed between author and year).

The list of references should be arranged alphabetically according

to the surname of the first author; all authors should be included.

Citations of two or more authors should be arranged alphabetical-

ly according to the first- and then the following author’s surname.

When more than one reference is given for a single author or the

same combination of authors, these should be arranged chronolog-

ically. If citing more than one reference by the same author(s) pub-

lished in the same year, use a, b, etc. after the year in both, text and

reference list (e.g. 2006a, b). The names of journals in the

References should be given in full. References “in press” shall on-

ly be cited when they have been accepted for publication. Exam-

ples of reference style are:

Kottelat M, Whitten T, Kartikasari SN, Wirjoatmodjo S (1993)

Freshwater fishes of Western Indonesia and Sulawesi. Periplus

Editions, Hong Kong

Mayr E (2000) The biological species concept. Pp. 17—29 in: Wheel-

er QD & Meier R (eds.) Species Concepts and Phylogenetic The-

ory — A Debate. Columbia University Press, New York

Parenti RP (2008) A phylogenetic analysis and taxonomic revision

of ricefishes, Oryzias and relatives (Beloniformes, Adrianichthyi-

dae). Zoological Journal of the Linnean Society 154: 494-610

Sullivan J (1994) Bufo boreas. In: Fire Effects Information System

(U.S. Department of Agriculture, Forest Service, Rocky Moun-

tain Research Station, Fire Sciences Laboratory). Online at

http://www.fs.fed.us/database/feis/animals/amphibian/bubo/all.ht

ml last accessed on December 28, 2009

Sztencel-Jablonka A, Jones G, Bogdanowicz W (2009) Skull mor-

phology of two cryptic bat species: Pipistrellus pipistrellus and

P. pygmaeus — a 3D geometric morphometrics approach with land-

mark reconstruction. Acta Chiropterologica 11: 113—126

Authors are kindly requested to make sure that sources listed in the

References match those cited in the text. Names of persons who pro-

vided unpublished information should be cited by including the ini-

tial(s) and city as follows: “(JG Frommen, Bonn, pers. comm.

2009)”.

Tables and Figures Tables and Figures should not duplicate infor-

mation provided in the text. Tables should be numbered consecu-

tively in Arabian numbers in order of their mention in the text (Table

1, Tables 1-3; do not abbreviate “Table(s)”). A brief self-explana-

tory title should be typed directly above each table, not on a sepa-

rate page. Do not insert frames, vertical rules, dotted lines or foot-

notes. Figures (maps, graphs, charts, drawings, photographs etc.)

must be numbered consecutively using Arabian numbers in order

of their mention in the text (abbreviate “Fig. 1”, “Figs 1-3” etc. in

the text, but use the full term “Figure(s)” in the captions. If refer-

ring to illustrations in cited sources, use “fig. / figs”. Grouped fig-

ures should be lettered with a lower case block letter in the lower

left corner. Scale should be indicated on figures by a scale bar. A

fully descriptive caption of each figure should contain all relevant

information; captions are to be typed together on a separate page.

Colour figures will be published free of charge if contributing sig-

nificantly to quality and / or readability of the manuscript. The de-

cision to print in colour or black and white any figure submitted in

colour will generally be based on the referee’s and handling editor’s

recommendations, but remains with the editor-in-chief and publish-

er. If appropriate, authors may be asked to group single colour fig-

ures into plates. For review, tables and figures should be enclosed

to text (preferable one electronic file per manuscript), but may al-

ternatively also be submitted as separate files (jpg, pdf, tiff). At this

stage, illustrations of low resolution allowing the referees to follow

the contents are sufficient; high-resolution files (jpg, pdf, tiff of pho-

tographs, maps etc. at > 300 dpi; line artwork etc. at 600 dpi or in

eps format) will be requested after acceptance. Unless otherwise

agreed with the editor, single electronic submissions should not ex-

ceed 6 MB file size.

Appendix Appendices should be numbered consecutively in Roman

numbers, in order of their mention in the text (Appendix I, Appen-

dices I-VI).

Abbreviations Except of very common abbreviations such as mm,

kg etc.; all abbreviations should be explained in the Methods sec-

tion or figure legend if appropriate. Hyphenation or upper case let-

ters for entire words are not permitted. All measurements must be

metric units and given to the same decimal, 1.e., 5.3—6.0 mm (not:

5.3-6 mm).

Instructions to authors

Taxonomy Names of animals and the description of new genera or

species must follow the current version of the International Code

of Zoological Nomenclature (ICZN, available at

http://www.iczn.org/iczn/index.jsp). Type specimens should be de-

posited in recognised institutions; deposition at ZFMK 1s 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 is required.

Review papers

Reviews should be concise, critical and creative, seeking to sum-

marize the state-of-the-art of complex topics, and stimulate debates

and new research initiatives. Review papers should comply with the

instructions for preparation of Research articles, with the exception

that the main body of text may be structured as appropriate.

Electronic submission

Manuscripts should be submitted as text files (*.doc or *.rtf *.pdf

is welcome for the review process, but for production *.doc or *.rtf

will be required) via e-mail (bzb.zfmk@uni-bonn.de) to the editor-

in-chief. Alternatively, send a CD. If neither is possible for any rea-

son, please contact the editor-in-chief. The file name must start with

the submitting author’s last name.

Review process

Each manuscript will be peer-reviewed, generally by two or more

referees. Manuscripts should be submitted to the editor-in-chief, and

will be handled by the editor dealing with the respective taxon or

topic (see list of editors). Authors are encouraged to suggest poten-

tial referees for their manuscripts. Final acceptance for publication

is in the responsibility of the editor-in-chief. This decision is main-

ly based on the referees’ report and the handling editor’s recommen-

dation. Revised manuscripts received more than 6 months after the

reviewers’ comments had been sent will be treated as new submis-

sions.

Ethical and legal aspects

Authors are obliged to state clearly that they followed the legal reg-

ulations and laws for the collection and ethical treatment of animals.

Publication and reprints

Manuscripts will be published online after acceptance as preprint

version, which will be replaced by the final pdf after publication of

the printed issue of BZB. The corresponding author must clearly state

so if preprint online publication of is not desired. In that case, on-

ly title, authors and the information that this manuscript has been

accepted for publication will be made available at that stage. Im-

mediately after publication, the corresponding author will receive

an e-mail containing the final pdf. On request (see order form sub-

mitted with page proofs), 25 reprints are supplied free of charge.

Additional reprints may be purchased on the author’s expense.

Questions we ask our reviewers

Authors preparing manuscripts for BZB should keep in mind the fol-

lowing questions which referees for the journal are asked:

1. Is the paper of interest to the readers of BZB?

2. How well does it meet the focus of BZB?

. Is it acceptable for publication, or should it be reconsidered af-

ter revision?

3. Please grade originality, quality of data, quality of statistical analy-

ses, and quality of interpretation on a scale from | = low to 5 =

high.

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?

. Does the title aptly and correctly describe the paper?

. Does the abstract summarize only the significant findings?

. Is the length of the paper appropriate?

. Are all (colour) figures and tables necessary and appropriate?

Oo on DW

SMITHSONIAN INS

Contents MIMO

Van Rooijen, Johan, Chan Kin Onn, L. Lee Grismer & Norhayati Ahmad:

Estimating the herpetofaunal species richness of Pangkor Island, Peninsular Malaysia

3 9088 01617 336

Geissler, Peter, Truong Quang Nguyen, Nikolay A. Poyarkov & Wolfgang Bohme:

New records of snakes from Cat Tien National Park, Dong Nai and Lam Dong provinces,

southern Vietnam

Vogel, Gernot & Johan van Rooijen:

Description of a new species of the genus Dendrelaphis Boulenger, 1890 from Myanmar

(Squamata: Serpentes: Colubridae)

Wagner, Philipp, Wolfram Freund, David Modry, Andreas Schmitz & Wolfgang Bohme:

Studies on African Agama IX. New insights into Agama finchi Bohme et al., 2005

(Sauria: Agamidae) with the description of a new subspecies

Bohme, Wolfgang, Mark-Oliver Rodel, Christian Brede & Philipp Wagner:

The reptiles (Testudines, Squamata, Crocodylia) of the forested southeast of the Republic Guinea

(Guinée forestiére), wit a country-wide checklist

Ahmadzadeh, Faraham, Fatemeh Khanjani, Aref Shadkam & Wolfgang Bohme:

A new record of the Persian Brook Salamander, Paradactylodon persicus (Eiselt & Steiner, 1970)

(Amphibia: Caudata: Hynobiidae) in northern Iran

Bohme, Wolfgang:

Type list of amphibians and reptiles in the Zoologisches Forschungsmuseum A. Koenig, Bonn:

corrections and additions

Van der Zee, Jouke R. & Rainer Sonnenberg:

Aphyosemion musafirii (Cyprinodontiformes: Nothobranchiidae),

a new species from the Tshopo Province in the Democratic Republic of Congo,

with some notes on the Aphyosemion of the Congo basin

Valdesalici, Stefano & Kiril Kardashev:

Nothobranchius seegersi (Cyprinodontiformes: Nothobranchiidae),

a new annual killifish from the Malagarasi River drainage, Tanzania

Gongora, Jaime, Cibele Biondo, Jennifer D. Cooper, Andrew Taber, Alexine Keuroghlian,

Mariana Altrichter, Fabricia Ferreira do Nascimento, Amanda Y. Chong, Cristina Yumi Miyaki,

Richard Bodmer, Pedro Mayor & Susana Gonzalez:

Revisiting the species status of Pecari maximus van Roosmalen et al., 2007 (Mammalia)

from the Brazilian Amazon

Irmler, Ulrich:

Two new species and a new record of the genus Neolindus Scheerpeltz, 1933

(Coleoptera: Staphylinidae: Paederinae)

Bohme, Wolfgang, Peter Geissler & Philipp Wagner:

A remarkable record of Phaneroptera falcata (Poda, 1761)

(Saltatoria: Phaneropteridae) from north-eastern Poland

Sonnenberg, Rainer & Eckhard Busch:

Erratum to: Sonnenberg & Busch (2010) Description of Callopanchax sidibei

(Nothobranchiidae: Epiplateinae), a new species of killifish from southwestern Guinea, West Africa

Cover illustration:

Naja sumatrana Miller, 1890 from Pangkor Island, Peninsular Malaysia

(this volume, pp. 3-8)

Ministerium fiir Innovation,

Wissenschaft und Forschung

des Landes Nordrhein-Westfalen

P, vay

/)% tf Leibniz

S. Gemeinschaft

TITUTION LIBRAR Ss

103

109

113