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Bonn zoological Bulletin

Editor-in-Chief

Fabian Herder, Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK), Ichthyology Section, Adenauerallee 160,

53113 Bonn, Germany,

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

f-herder.zfmk@uni-bonn.de

Editorial Board

Dirk Ahrens, Insects: Coleoptera, ZFMK,

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

d.ahrens.zfmk@uni-bonn.de

Wolfgang Bohme, 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. +49 228-9122-261, fax: +49 228-9122-212:

r.hutterer.zfmk@uni-bonn.de

Gustav Peters, Mammals, ZFMK,

tel. +49 2289122262, fax: +49 2289122212:

g.peters.zfmk@uni-bonn.de

Bradley Sinclair, Canadian National Collection of Insects,

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

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

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

bradley.sinclair@inspection.gc.ca

Dieter Sttining, Insects except Coleoptera and Diptera,

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

d.stuening.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 22628-7304, fax: +48 22-629-6302;

wieslawb@miuiz.waw.pl

Matthias Glaubrecht, Museum fiir Naturkunde Berlin, Leib-

niz-Institut fiir Evolutions- und Biodiversitatsforschung an

der Humboldt-Universitat zu Berlin, Invalidenstrasse 43,

10115 Berlin, Germany,

tel. +49 30-2093—8504/ 8400, fax: +49 030-2093-8565;

matthias.glaubrecht@mfn-berlin.de

Jurgen Haffer, Tommesweg 60, 45149 Essen, Germany,

tel. +49 201—710426; j.haffer@web.de

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

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

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

Boris Krystufek, Slovenian Museum of Natural History, P.

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

Wolfgang Schawaller, Staatliches Museum ftir Naturkunde,

Rosenstein 1, 70191 Stuttgart, Germany,

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

schawaller.smns@naturkundemuseum-bw.de

Ulrich K. Schliewen, Department of Ichthyology, Bavarian

State Collection of Zoology, Miinchhausenstr. 21, 81247

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

schliewen@zsm.mwn.de

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

meine & Systematische Zoologie, Anklamer Str. 20, 7489

Greifswald, Germany,

tel. +49 3834-86-4242, fax: +49 3834864098;

michael.schmitt@uni-greifswald.de

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

Sciences, W. Texas A. & M. University, 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: + 49 531-391-8198; m.vences@tu-bs.de

Erich Weber, Eberhard-Karls-Universitat, Zoologische

Schausammlung, Sigwartstr. 3, 72076 Tiibingen,

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

erich.weber@uni-tuebingen.de

Editorial

Welcome to the first issue of BzB in 2010. The journal

has been subject to major recent changes, including title

and scope, editor, boards, cover design, and free availabil-

ity of all contributions (open access).

The title “Bonner zoologische Beitrage” changed to “Bonn

zoological Bulletin”. This follows the fact that the jour-

nal’s contributions, like elsewhere in biological sciences,

are nowadays nearly exclusively published in English. The

new title shall also contribute to increasing the journal’s

publicity while keeping the well-known acronym “BzB”.

The journal focuses now explicitly on three fields, sharp-

ening its traditional profile as a museum journal: (1) tax-

onomy, (2) systematics and evolution, and (3) biodiver-

sity and biogeography, all with respect to terrestrial ani-

mals (including fresh or brackish waters).

Publication in BzB continues to be free of charge, includ-

ing colour figures, open access, retention of full rights with

the authors, and production of a widely distributed print

version. Manuscripts will be published online ahead of

print directly after acceptance, allowing immediate pub-

lic access to the latest contents. The BzB webpage (see

interior cover for address) is still under revision, but con-

tains already all published issues back to 1993. Our goal

for the near future is to make the contents of all past vol-

umes available online. In summary, the concept of BzB

will guarantee fast and easy availability of the published

contents.

Some of the editional responsibilities changed include the

editor-in-chief and the boards. Michael (“Theo’’) Schmitt,

responsible for BzB since 2003, left Museum Koenig for

Greifswald University last year. Likewise, the boards have

been subject to some rearrangement. On behalf of the

“BzB-team’’, I thank Theo and the former editorial and ad-

visory board members for their invaluable engagement.

Theo will however stay with BzB as a member of the Ad-

visory Board. A warm “welcome on Board” to those col-

leagues joining or staying with the advisory or editorial

team of Bonn zoological Bulletin, I am looking forwards

to working with you.

JUL 07 2010

LIBRARES

Despite of the changes outlined above, the journal will re-

tain its core strengths as a museum journal. The board

members cover a wide array of expertise in zoology, and

share common interests related to the journal’s focus. [am

convinced that this is a sound basis for the journal’s fu-

ture, building on the history of “Bonner zoologische

Beitrage” since 1950.

Contributions published in the present issue of BzB re-

flect a wide array of approaches and methods applied in

organismal zoology. Besides a checklist and contributions

dealing with morphology, autecology or host-parasite in-

teraction, the contents of the present issue include formal

descriptions of six species new to science, including a

chameleon, a gecko, a killifish (see front cover), two new

species of moths, and a new fossil aquatic dance fly from

Baltic amber. Taxonomic papers like these contribute to

the exploration of our planet’s still only partly known bio-

ta, and are increasingly not in the focus of interest of many

present zoological journals. Publishing sound taxonomic

work remains a core task for museum journals like BzB.

Finaily, some words regarding cover design and text for-

mat. In 2003, the BzB front cover was changed from com-

pletely yellow to the design known until 2009 when the

journal switched to a larger format. At this time, the ad-

ditional cover space was contrasted in light grey. The pres-

ent design by Uwe Vaartjes takes the next step, replacing

the yellow colour completely by grey and neutral ele-

ments, and incorporates Regina Liitzenkirchen’s idea of

including Bonn’s heraldic animal, the “King (Koenig in

German) of Animals”. I am grateful to both for their work.

Some readers and authors will have to get used to the more

simplified literature style, which is a substantial reduction

of the workload in production for both, authors and edi-

tors. I hope that the current revision of BzB in terms of

both, focus and style, will meet the approval of the jour-

nal’s audience.

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

members,

Tibi lect

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

Bonn, April 2010

Bonn zoological Bulletin Volume 57

Issue |

pp. 3-14 Bonn, April 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-53223 Bonn, Germany.

Corresponding author. E-mail: r.sonnenberg.zfmk@uni-bonn.de

2 Max-Planck-Institut fiir Evolutionsbiologie, August-Thienemann-Strasse 2, D-24306 Plon, Germany

3 Diederichsstrasse 45, D-42855 Remscheid, Germany

Abstract. A new species of the annual killifish genus Callopanchax (Nothobranchiidae: Epiplateinae) is described from

coastal Guinea, West Africa. It is distinguished from its congeners by morphological and colour pattern characters in-

cluding up to three irregular and interrupted horizontal stripes on blue flanks in males. A preliminary DNA study indi-

cates that it is closely related with C. occidentalis and C. toddi, which is supported by the following apomorphic char-

acters compared to the more distantly related C. monroviae: a dark red or black postopercular blotch and an origin of the

dorsal fin anterior to or above the first ray of the anal fin. The species first turned up in commercial imports of ornamen-

tal fish from Guinea. The local collectors, the Sidibe family, are acknowledged for sharing their knowledge on this species

and their support to the second author during his collections by naming it Callopanchax sidibei, new species. Currently

it is only known from the type locality and is endemic to Guinea.

Résumé. Une nouvelle espéce annuelle du genre Callopanchax (Nothobranchiidae: Epiplateinae) est décrite des cdtes

maritimes de Guinée, Afrique de l|’Ouest. Elle se distingue de ses congeneres par certains caracteres morphologiques et

un motif coloré comptant jusqu’a trois bandes horizontales irréguliéres et interrompues sur les flancs bleus chez les ma-

les. Une étude ADN préliminaire indique que ce taxon est proche de C. occidentalis et C. toddi. Ceci est également sup-

porté par les caractéres apomorphes suivants — comparés a C. monroviae, de parenté plus éloignée: tache post-opercu-

laire rouge sombre ou noire, et origine de la nageoire dorsale au-dessus ou antérieure au premier rayon de la nageoire

anale. L’espéce a été détectée en premier dans les importations commerciales d’espéces ornementales en provenance de

Guinée. Callopanchax sidibei sp. nov. est dédiée a la famille Sidibe, engagée dans les collectes locales, pour avoir par-

tagé ses connaissances et avoir aidé le second auteur durant sa mission de collecte. Cette espece endémique de Guinée

n’est actuellement connue que de la localité type.

Key words. Cyprinodontiformes, Callopanchax sidibei, new species, taxonomy, endemic, Upper Guinea.

INTRODUCTION

The nothobranchiid genus Callopanchax Myers, 1933

consists of four described species of which three are cur-

rently considered valid (Costa 2008; Romand 1985, 1992;

Seegers 1997; Wildekamp 1996; Wildekamp & Van der

Zee 2003). They are relatively large annual killifishes,

reaching up to 90.0 mm in total length according to

Wildekamp & Van der Zee (2003), living in the coastal

plains from southwestern Guinea and Sierra Leone to

western Liberia. Callopanchax occidentalis (Clausen,

1966) and C. toddi (Clausen, 1966) were described from

Sierra Leone, the latter as a subspecies of C. occidental-

is. Some years later two further species, C. monroviae

(Roloff & Ladiges, 1972) from Liberia, and C. huwaldi

(Berkenkamp & Etzel, 1980) from Sierra Leone, were for-

mally described. The latter species is currently considered

a synonym of C. occidentalis by most authors (Costa

2008; Lazara 2001; Romand 1985; Wildekamp 1996;

Wildekamp et al. 1986).

Callopanchax species are easily separated from all other

species groups within the Epiplateinae by the large num-

ber of anal and dorsal fin rays and an origin of the dorsal

fin anterior or slightly posterior to origin of the anal fin;

in the remaining genera the dorsal fin inserts distinctly

posterior to the anal fin origin. They share, with the re-

maining Epiplateinae, the frontal and preopercular neu-

romast pattern (Clausen 1966, 1967; Grimm 1974; Van der

Zee 2002; Van der Zee & Wildekamp 1994), a red pig-

mentation pattern on the side of the head (Van der Zee

2002), and with Archiaphyosemion guineense (Daget,

Rainer Sonnenberg & Eckhard Busch

Table 1. List of specimens used for DNA sequencing with voucher sample number, collection locality, country, and GenBank ac-

cession numbers. Accession numbers with * are published in Murphy & Collier (1997), with ** in Sonnenberg & Busch (2009).

Aquarium strain means that the origin in nature for this sample is unknown.

Sample no. _ species Collection locality Country GenBank acc. no.

OCC GB Callopanchax occidentalis aquarium strain Sierra Leone U73242*

OCC1530 Callopanchax occidentalis SL 03/11, Malai Sierra Leone GU553007

OCC1531 Callopanchax occidentalis SL 89, Teme Yellah Sierra Leone GU553008

OCC1532 Callopanchax occidentalis SL 03/19, Magbenta Sierra Leone FJ872034**

TOD 1534 Callopanchax toddi GM 97/26, Takhori Guinea GUS553009

TODI535 Callopanchax toddi GM 97/24, Mola Guinea GU553010

cTOD1758 ~~ Callopanchax cf. toddi GM 97/3, Mangata Badjana Guinea GU553011

SID1536 Callopanchax sidibei GM 97/2, Bombokore Guinea GU553012

MON1533 ~~ Callopanchax monroviae L 97, Paynesville Liberia GU553013

GER1381 Scriptaphyosemion geryi SL 03/9, Robis 1 Sierra Leone FJ872033**

GUI1404 Archiaphyosemion guineense SL 93/37, Lenghe Curoh Sierra Leone FJ872031**

50 km

Fig. 1.

Tay,

Map of Callopanchax collection localities in coastal West Africa. The letter T indicates the respective type localities, D

indicates origin of Callopanchax DNA samples used in this study. Black square = C. occidentalis, white square = C. toddi, black

circle = C. cf. toddi, white circle = C. huwaldi, black triangle = C. monroviae, white triangle = C. sidibei.

Bonn zoological Bulletin 57 (1): 3-14

©OZFMK

A new Callopanchax from Guinea 5

Table 2. Uncorrected p-distances for the mitochondrial LSU sequences with pairwise exclusion of missing data among nine sam-

ples of Callopanchax, one of Archiaphyosemion, and one of Scriptaphyosemion. The upper triangle contains the linear geographic

distances between the different collection localities of the sequenced Callopanchax specimens, except for OCC GB, which is ac-

cording to Murphy et al. (1999) an aquarium strain of unknown origin.

sample OCEGBROCES (OKELE OCC TOD TOD cIOD SID MON GER

1530 S31 N37 1534 3S) 1758 1536 1533 1381

OCC 1530 LEV — 204km 40km 196km 64km 106km 95km 370km— —-

O€Een53)1 1S od! = 185km 396km 245km 301 km 289km 175km = —

OC@1532 I) IES 1S — 230km 103km 144km 132km 340km— —

TOD 1534 6.6 5.8 6.8 6.6 — 154km 97km 110km 565km— —

TOD 1535 5) Se 6.2 6.0 ed = S57km 43km 418km —

cTOD 1758 6.8 5.6 6.6 6.4 Del 1S = 14km 472km— —

SID 1536 7.6 6.7 eS Toll HES 6.5 6.2 = 458km —

MON 1533 14.2 W23) 12.8 Salk WS LES IES HSA = =

GER 1381 9.6 Jai) 8.7 9.3) OFS) OF) 10.1 10.6 14.2 —

GUI 1404 IN 10.8 11.3 10.8 10.8 10.8 11.6 12.4 14.8 9.4

1954) and Scriptaphyosemion Radda & Piirzl,1987 a dis-

tinct constriction of the subdistal portion of the posterior

arm of the parasphenoid (Costa 2009). The phylogenetic

placement of Callopanchax within the Epiplateinae and

its sister group relationship with Scriptaphyosemion 1s sup-

ported by molecular data (Murphy & Collier 1997; Mur-

phy et al. 1999; Sonnenberg & Busch 2009).

In 1992 a scientifically unknown Callopanchax species

turned up in a commercial import of ornamental fish from

Guinea to the Netherlands. Its distinctness was first rec-

ognized by the unique colour pattern of this species (Busch

1993; Wildekamp 1996; Wildekamp & Van der Zee 2003)

and was later supported by a study based on molecular da-

ta (Murphy et al. 1999). During several research trips in

Guinea, the second author was able to collect specimens

of this undescribed Callopanchax, which is herein de-

scribed as Callopanchax sidibei, new species.

MATERIAL AND METHODS

The specimens on which the following description is based

were deposited in the ichthyological collections of the

Musée Royal de |’Afrique Centrale (MRAC), Tervuren,

Belgium, and the Zoologisches Forschungsmuseum

Alexander Koenig, Bonn, Germany (ZFMK). Compara-

tive material of the other species is listed below.

Measurements and counts are made as indicated in Son-

nenberg & Busch (2009), transverse scale count was made

between the anterior insertion of the pelvic fin and dor-

sal fin; the very small scales on the belly were not count-

Bonn zoological Bulletin 57 (1): 3-14

ed. Digital X-ray images of the types and some specimens

of the comparative material used for unpaired fin ray and

vertebrate counts were made with a Faxitron LX-60 Dig-

ital Specimen Radiography System at the ZFMK. Caudal

fin ray counts were only made from X-ray images and in-

clude all visible rays.

We prepared a preliminary DNA dataset of a fragment of

the mitochondrial large ribosomal subunit (16S rDNA) for

a small number of specimens, including three of C. occi-

dentalis from Sierra Leone, one sequence of C. occiden-

talis from GenBank (accession number U73242, Murphy

& Collier 1997), two specimens of C. toddi and one of C.

cf. toddi from Guinea, one specimen of each C. monrovi-

ae and C. sidibei, new species, and, as an outgroup, Archi-

aphyosemion guineense and Scriptaphyosemion geryi

(Lambert, 1958). The latter two are the type species of the

respective genera. Specimens are listed with locality in-

formation and GenBank accession number in Table 1,

methods for DNA extraction, amplification and sequenc-

ing of the 16S rDNA fragment are provided in Sonnen-

berg & Busch (2009), where also some of the sequences

were published. Resulting sequences were aligned with

ClustalX 1.8 (Thompson et al. 1997) and checked by eye

in BioEdit 7.0.5.3 (Hall 1999). Sequences were tested for

the anti-G bias characteristic of mitochondrial DNA

(Zhang & Hewitt 1996). Uncorrected p-distances with the

pairwise deletion of missing data were calculated in Mega

4.1 beta 3 (Tamura et al. 2007) and are given in Table 2.

Analyses of sequence data were made with PAUP 4.0b10

(Swofford 1998) by maximum parsimony and with Mr-

Bayes 3.1.2 (Huelsenbeck & Ronquist 2001) by a

©ZFMK

6 Rainer Sonnenberg & Eckhard Busch

Fig. 2.

Bayesian approach. Maximum parsimony analyses were

done with the exclusion of uninformative positions, gaps

were coded as a fifth character state, and heuristic tree

searches were performed with random addition of se-

quences for 1,000 replicates. Bootstrapping was performed

with the random addition of sequences for 100 replicates

and 10,000 bootstrap replicates. Bayesian analyses were

performed with the number of generations set to

10,000,000, with sampling every 1,000 generations, and

the results of the first 1,000,000 generations were discard-

ed as a burn-in time after checking that the runs had

reached the stationary phase at this point of the analysis.

We performed two runs for this dataset, one with two pa-

rameter models (Nst 2) and one with six parameter mod-

els (Nst 6), settings for rate=gamma and ngammacat=4.

We chose to perform these two runs to check if there are

serious differences in the results due to potential under-

or Overparametrization, especially because of the small

dataset with large genetic divergences (Table 2). We con-

sider only nodes with 2 75 % bootstrap support and 2 95

% posterior probabilities as supported by the data.

Additional information for the collection localities in Table

1 can be found in the comparative material list. Collec-

tion points in Figure | are from museum collection data

Bonn zoological Bulletin 57 (1): 3-14

A) Holotype of Callopanchax sidibei, male, ZFMK 41613, 58.5 mm SL. B) Paratype of Callopanchax sidibei, male,

ZFMK 41614, 47.0 mm SL. C) Callopanchax sidibei, male with red dots in the anal and caudal fin, not preserved. D) Paratype of

Callopanchax sidibei, female, ZFMK 41615, 55.4 mm SL.

as accessed via Fishbase (Froese & Pauly 2009), from Hu-

ber (2007), and collection data of the second author. We

adopted, as a species concept, the definition by Moritz et

al. (2000) which ,,...recognize as species geographically

bounded sets of populations that are distinct for morpho-

logical traits or are reproductively isolated from congeners,

with or without corresponding molecular divergence...“.

RESULTS

Callopanchax sidibei, new species

(Fig. 2A—D, Table 3)

Roloffia sp. ,,auinea, CI 92°: Busch (1993), p. 64.

Fundulopanchax species aff. toddi: Wildekamp (1996), pp.

212-213.

Callopanchax sp. Guinea: Wildekamp & Van der Zee

(2003), pp. 381-383.

Callopanchax huwaldi (non Berkenkamp & Etzel, 1980):

Neumann (2005), misidentification in fig. caption p. 48.

Holotype. ZFMK 41613, male, 58.5 mm SL, Guinea,

Guinée maritime, Kindia region, small river and adjacent

pools and ditches in a secondary forest (9°32.71° N,

©ZFMK

A new Callopanchax from Guinea 7

Table 3. Morphometric and meristic data of Callopanchax sidibei. All measurements in percentages of standard length except stan-

dard length in mm. Fin ray counts were made on X-rays and include all visible rays. SD = standard deviation.

Holotype &@ Types @Q Types all Types all Types all Types SD

mean mean mean range

Standard length 58.5 48.4 50.7 49.7 39.0—59.9 6.8

Total length 124.8 124.6 122.7 WBE 117.1-132.5 4.5

Head length 28.4 29.2 DD), 2) Il 27.7-30.5 0.9

Predorsal fin distance S10 // 57.6 60.1 59.0 55.4—63.2 1S,

Prepelvic fin distance 54.1 52.0 Sal 52.4 49.1-56.6 2.0

Preanal fin distance 62.9 62.5 64.2 63.5 60.1—68.9 be

Greatest body depth 24.3 D3 Dre BI} 20.0—25.8 1.6

Depth of caudal peduncle 1333 I WI 12.6 10.7-14.6 1.0

Length of caudal peduncle 23m 21.6 Japa) AN) IE8=2551 lad

Base of dorsal fin 28.3 28.2 Soi 26.8 23.5—-30.7 1:9

Base of anal fin D3) 23rl 18.2 20.3 17.6—24.6 2.6

Eye diameter VP 6.8 6.8 6.8 6.1—7.6 0.4

Interorbital distance IBo7/ 211 Wed IES, 10.3-13.7 0.8

Dorsal fin rays 19 19.8 19.8 19.8 19-21 0.8

Anal fin rays 7) eo 18.1 18.0 17-19 0.6

Dorsal / anal fin position -2 -2.8 -3.2 -3.0 -2— -4 0.7

Caudal fin rays 33 32.0 31.8 Shes) 29-34 72

Total number of vertebrae 33 Saal 32.6 32.8 32-34 0.7

Vertebrae with pleural ribs 15 1522 14.8 15.0 14-16 OW

Vertebrae with haemal spines 18 We 17.8 17.8 WS 0.6

Lateral line scales 3444 Aros) seollasio) SI SollaeSioy Ses V4 riesies) 0.9+0.6

Transverse row of scales WP eZ 10.8 11.0 10-13 0.8

Scales around caudal peduncle 16 NS) 15.8 LY) 15=1'6 0.4

13°14.51° W), close to the small village Bombokoré, about

3 km from the closest larger village Fandi, collection lo-

cality GM 08/2, E. Busch and B. Wiese, 25.10.2008, Fig-

ure 2A.

Paratypes. ZFMK 41614-41615, 1 male, 47.0 mm SL,

1 female, 55.4 mm SL, paratopotypes, collected with holo-

type, Figure 2B & D.

ZFEMK 41616-41628, 5 males, 8 females, 39.0—58.9 mm

SL, paratopotypes, collected with holotype.

MRAC 2010—02—P—1—5, 2 males, 3 females, 46.3—59.9

mm SL, paratopotypes, collected with holotype.

Diagnosis. Callopanchax sidibei (Fig. 2A—D) is, togeth-

er with C. toddi (Fig. 3A—F) and C. occidentalis (Fig. 4A

& B), distinguished from C. monroviae (Fig. SA—C) by

insertion of first dorsal fin ray slightly anterior to first anal

fin ray versus a posterior insertion. The former three

species are also distinguished from C. monroviae by a dark

red to black postopercular blotch above insertion of pec-

toral fin versus absence and a dark or black blotch on the

Bonn zoological Bulletin 57 (1): 3-14

anterior dorsal fin rays instead. The postopercular blotch

is present in females of C. sidibei, C. occidentalis, and C.

toddi but often only faint; the dark blotch on anterior dor-

sal fin in C. monroviae may not be visible in females. Cal-

lopanchax sidibei, C. occidentalis, and C. toddi males have

an anal fin with a white or bluish-white margin and usu-

ally only a small number of red dots or blotches on fin

center; no horizontal red band or only traces of red under

the eye from lower lip to posterior border of opercle;

postopercular red pigmentation on flank formed by up to

three irregular red horizontal bands interrupted into blotch-

es. Males of C. monroviae have a dark or dark-red anal

fin margin, proximal red dots, blotches or broad vertical

streaks, and streaks between fin rays on distal part of anal

fin; a more or less complete red band from snout under

the eye and often extending on side to origin of pelvic fin;

postopercular red pigmentation consists of incomplete

rows of small red dots on anterior side and less and irreg-

ularly distributed red dots or a pattern of incomplete and

irregular vertical stripes on posterior side. Males of C.

sidibei can also be distinguished from C. monroviae by

©ZFMK

Rainer Sonnenberg & Eckhard Busch

Fig. 3. A) Male of Callopanchax cf. toddi, Mangata Badjana, Guinea, collection locality GM 08/3 (= GM 97/3). B) Female of

Callopanchax cf. toddi, Mangata Badjana, Guinea, collection locality GM 08/3 (= GM 97/3). C) Male of Callopanchax toddi, Mo-

la, Guinea, collection locality GM 09/17 (= GM 08/4 & GM 97/24). D) Female of Callopanchax toddi, Mola, Guinea, collection

locality GM 09/17 (= GM 08/4 & GM 97/24). E) Male of Callopanchax toddi, Lamayah, Guinea, collection locality GM 08/11.

F) Male of Callopanchax toddi, Takhori, Guinea, collection locality GM 09/12 (= GM 08/9 & GM 97/26).

body colouration which is mainly blue to blue-green on

sides and dorsally often a golden sheen to the anterior, ver-

sus anteriorly pale blue to blue-green and dorsal and pos-

terior half of body reddish, reddish-brown or orange, or

in the blue form body completely pale to grey-blue (Fig.

S5A-C).

Bonn zoological Bulletin 57 (1): 3-14

Callopanchax sidibei can be distinguished from males of

C. occidentalis and C. toddi by up to three irregular and

interrupted horizontal stripes on flanks, an absence of ver-

tical dark red or reddish-brown stripes or broad blotches

on sides and a mainly blue to blue-green body colour ver-

sus dark brown, brown or red vertical stripes or blotches

©ZFMK

Anew Callopanchax from Guinea 9

Fig. 4.

A) Male of Callopanchax occidentalis, Malai, Sierra Leone, collection locality SL 03/11 (= SL 93/54). B) Female of

Callopanchax occidentalis, Malai, Sierra Leone, collection locality SL 03/11 (= SL 93/54).

Fig. 5. A) Male of Callopanchax monroviae, aquarium strain

from Paynesville, Liberia, collection locality L 97, red form. B)

Male of Callopanchax monroviae, aquarium strain from Pay-

nesville, Liberia, collection locality L 97, blue form. C) Female

of Callopanchax monroviae, aquarium strain from Paynesville,

Liberia, collection locality L 97.

Bonn zoological Bulletin 57 (1): 3-14

and a more reddish brown or copper to golden body colour

with blue only on ventral side. Exceptions are populations

with rare blue males in C. occidentalis and, if valid, C.

huwaldi, which show the same pattern as the ‘typical’ form

but lack orange and yellow pigments, but not the red pig-

mentation and the dark vertical stripes or blotches on pos-

terior side (e.g. Berkenkamp & Etzel 1980; Eberl 1999,

pictures of C. occidentalis from Mabeimah on p. 65;

Roloff 1976).

Large males of C. sidibei show only a slightly rugged anal

and dorsal fin border and rounded or slightly pointed fin

tips versus more pronounced rugged fin borders in most

C. occidentalis and C. toddi, especially the posterior tip

of the anal fin, which can show filamentous extensions.

Females of C. sidibei can be distinguished from C. occi-

dentalis and C. toddi females by mid-laterally blue to blue-

green coloured scales and the absence of the dark brown

vertical stripes or blotches, similar as in males versus blue

to blue-green coloured scales only on anterior side, often

with short vertical dark blotches dorsally on side. They

are distinguished from females of C. monroviae by their

pigmentation pattern of red horizontal stripes with broad

blotches versus more regular horizontal rows of small red

dots and the absence of metallic blue to blue-green

coloured scales on side in C. monroviae.

Description. See Fig. 2A—D for general appearance and

Table 3 for morphometric and meristic data of the types.

Callopanchax sidibei (Fig. 2A—D) shares with its con-

geners the following combination of diagnostic characters

within the Epiplateinae: large and robust species, dorsal

and anal fins with a high number of fin rays, origin of dor-

sal fin close to origin of anal fin. A species with strong

sexual dimorphism, adult males usually larger, with larg-

er fins and more colourful than females. Maximum ob-

©OZFMK

10 Rainer Sonnenberg & Eckhard Busch

served SL in males 59.9 mm, in females 59.0 mm. Body

cylindrical, outline from tip of snout to posterior end of

dorsal and anal fin convex, on caudal peduncle dorsally

and ventrally slightly concave to nearly straight. Greatest

body depth approximately at half distance between ori-

gins of pectoral and pelvic fins.

Snout slightly pointed, mouth directed upwards, lower jaw

longer than upper jaw; many irregularly distributed un1-

cuspid, slightly curved teeth of different size, a small num-

ber of larger ones on the outer row of upper and lower jaw.

Closed frontal neuromast system in one groove, preoper-

cular neuromast system with 5 pores.

Total number of vertebrae 32-34, number of vertebrae

with pleural ribs 14-16, with haemal spines 17—19. The

X-ray images indicate that in about half of the studied

specimens the hypural plates are fused, as no gap or in-

dication of overlap 1s visible.

Fins in males slightly larger than in females, base of dor-

sal mean value 28.2 % of standard length (SL) in males

versus 25.7 % SL, base of anal in males 23.1 % SL ver-

sus 18.2 % SL in females. Dorsal and anal fin rounded or

slightly pointed, no long filamentous extension, fin inser-

tion in both sexes posterior to mid-body. In large adult

males, dorsal and anal fin rays sometimes protrude from

the fin membran, giving the impression of a rugged fin

border. Papillae on anal fin rays in males. First 2—4 dor-

sal fin rays anterior of origin of anal fin. Dorsal fin rays

19-21, anal fin rays 17-19. Caudal fin rounded or slight-

ly truncated, 29-34 fin rays. The single specimen with 29

caudal fin rays has a small gap between the anterior dor-

sally situated rays explaining the comparatively low val-

ue. Pectoral fin origin on ventral half of side, posterior of

opercle, rounded, extending posteriorly, not reaching ori-

gin of pelvic fin. Pelvic fin small and slightly pointed, not

or just reaching anal fin.

Scales cycloid, body and head entirely scaled except

throat, scales dorso-anteriorly larger then ventrally and

posteriorly, frontal squamation of G-type, scales on lat-

eral line 32-34 + 3-5 on caudal fin base, not all scales along

lateral line with small grooves in center. Transverse row

of scales above pelvic fin 10-13, circumpeduncular scale

row 15-16.

Live colouration. Males. (Fig. 2A—C) Dorsal from head

to dorsal fin brown, some scales on back with red dots or

scale borders, forming an irregular reticulated pattern to

the end of the caudal peduncle; flanks blue to blue-green,

dorsolaterally with a metallic copper or golden sheen on

scales, belly blue to bluish-grey.

Bonn zoological Bulletin 57 (1): 3-14

Red pigmentation pattern on head posterior of the eye typ-

ical for Epiplateinae (Van der Zee 2002) with two poste-

rior directed horizontal red stripes, originating from one

point posterior of the eye to posterior border of opercle,

and two more or less complete vertical stripes dorsally.

On flanks three interrupted and irregular red stripes, up-

per and lower stripes only extending up to the posterior

insertion of dorsal fin, only the middle stripe, positioned

slightly above mid-line of body, extends to the end of the

caudal peduncle, in some males the dorsolateral stripe on-

ly consists of a small number of red dots or red scale bor-

ders.

Posterior of the opercle and above insertion of pectoral

fin usually 2—3 scales in two neighbouring rows with

broad black or dark red margin, forming a black or dark

red blotch.

Unpaired fins blue to blue-green, with white or, rarely, yel-

low-white margins and red submarginal stripe. Red pig-

mentation pattern on unpaired fins variable. Dorsal fin

with red dots or short streaks between fin rays, below red

submarginal band a whitish zone. Anal fin without or on-

ly with little red pigmentation beside submarginal red

stripe, in some males a proximal row with a small num-

ber of red dots (Fig. 2C). Caudal fin in some males with

red dots or blotches, lower white margin broader then up-

per and in some specimens slightly yellow-white. In males

with a more rounded caudal fin shape marginal and sub-

marginal stripes can be closed on posterior end of fin, in

males with a more truncate fin shape they end on the pos-

terior fin border (Fig. 2A—C). Pelvic fins bluish, with red

submarginal and white marginal stripe, pectoral hyaline

with a white or blue-white margin.

Females. (Fig. 2D) Body brown to light brown, dorsally

darker then ventrally, sometimes with a golden colour

sheen on anterior side. Scales dorsally and laterally with

dark border, forming a reticulate pattern. Lateral red pig-

mentation pattern similar to males, usually less pro-

nounced and paler. Broad dark brown to brownish-grey

lateral stripe from posterior border of operculum above

origin of pectoral fin to posterior end of caudal peduncle

on upper half. This stripe is not always visible and espe-

cially pronounced in stressed specimens. On upper half

of opercle, posterior of the eye, extending along the dark

stripe on the side irregularly distributed scales with metal-

lic blue or blue-green colour, with the highest density an-

teriorly. Fins except pectoral light brown and slightly hya-

line, unpaired fins with a whitish margin, on dorsal and

caudal with pale red submarginal line. Dorsal and caudal

fin with red dots, on dorsal sometimes restricted to fin

base. Pectoral fin transparent with whitish to light blue

margin.

©ZFMK

Anew Callopanchax from Guinea 1]

Preserved in ethanol. Males. Head dark grey, body dark

brown to grey, some traces of red dots and blotches on

side, belly whitish, beige, or greyish, a ventral dark stripe

posterior to level of pectoral fin insertion to anterior ori-

gin of anal fin. Fins pale grey with a dark submargin and

small whitish margin, except caudal fin with a ventral

broad and dorsal thin whitish margin, posterior border dark

brown to reddish.

Females. Head laterally and dorsally dark grey, chin with

a dark blotch on ventral inner side of lower lip, body dor-

sally darker grey or brown, a dark stripe from posterior

border of opercle to origin of caudal fin, dark stripe dor-

sally with a sharp border, ventrally diffuse; below this

stripe brown, anterior ventral side and belly whitish to light

brown, on side traces of red spots and blotches. Fins light

grey with a dark border, sometimes with a small white

margin, in dorsal and caudal fin traces of red spots, very

rare with proximal single row of red dots in anal.

Etymology. The new species is named after Mr. Samba

Sidibe and his family, who first collected this fish and

made specimens available for the ornamental fish trade.

It is named in acknowledgement of their high interest in

this species, their effort for a sustainable use and protec-

tion of the habitat, and providing important information

for the description.

Distribution and habitat. Currently C. sidibei is only

known from the type locality close to the small village

Bombokoré, despite some effort to find further occur-

rences of this species in the vicinity. Bombokoré is the

spelling in the Sousou language, it might have a different

spelling on maps or gazetteers.

The habitat is a depression with small water bodies, ditch-

es and connected pools in a secondary forest adjacent to

a small creek. The water in the creek is only slowly flow-

ing and stagnant in pools and ditches. During the first col-

lection in 1997, the second author measured an air tem-

perature of 36°C, the water temperature in the creek was

27°C and 29°C in ditches, pH was about 5.2, conductiv-

ity 10uS/cm. Water in the ditches was about 20-30 cm

deep and clear, the bottom consists of of sand and some

stones with a layer of decaying leaves and wood with no

aquatic vegetation. Herbaceous plants were found along

the edges. Callopanchax sidibei and Epiplatys fasciola-

tus (Gunther, 1866) were found in ditches and pools with

standing water. In places with only grass and no leaf cov-

er, only Scriptaphyosemion geryi was recorded.

During the collection in 2008, the depression was flood-

ed. The stagnant or slowly flowing water was shaded by

herbaceous vegetation and young trees on edges and some

aquatic vegetation, especially Nymphea. Air temperature

Bonn zoological Bulletin 57 (1): 3-14

was 31°C, water temperature 27°C, water depth about

60-80 cm maximum. Interestingly the specimens caught

in 2008 were of different size, from juveniles with about

2.5 cm total length to full grown adults, and in good con-

dition, whereas in 1997 all were fully grown and showed

first signs of ageing. During the last visits in 2008 and

2009 only a little human impact on the forest could be de-

tected.

Callopanchax sidibei seems to prefer shaded areas of the

water bodies with slow flowing or standing water. How-

ever, the life cycle of this species depends on the flowing

water of the creek, because there is a seasonal connection

between the creek and the pools and ditches. During the

rain season the creek floods the depression and provides

habitat, fresh water and food sources for the growing ju-

veniles. Later, the inflow is reduced and isolated or par-

tially connected pools and ditches are formed, which dry

out during the dry season. Within these water bodies re-

production takes place and adult fish most likely die dur-

ing the dry season. Callopanchax sidibei 1s, like its con-

geners (e.g. Scheel 1968; Wildekamp 1996), an annual

species with eggs showing a prolonged development of

2—9 month, apparently an adaptation to temporary water

bodies. However, some adults might move during the dry

season into the flowing creek, which is supported by the

observation of the local people.

DNA analyses. The complete alignment of the mitochon-

drial large ribosomal subunit for 11 specimens contains

530 bp including gaps, the sequence lengths without gaps

ranges between 518-526 bp; the GenBank sequence

(U73242) is 46 base pairs shorter, missing positions were

filled with N. Of the 530 bp 133 are variable and 61 were

parsimony informative, calculated without gaps. All se-

quences show an anti-G bias typical for mitochondrial se-

quences (Zhang & Hewitt 1996).

Calculated intraspecific p-distances for the samples of C.

occidentalis as well as C. toddi and C. cf. toddi (Table 2)

are higher than for the corresponding fragment in Nim-

bapanchax (Sonnenberg & Busch 2009), ranging from

1.3—2.7, which is in the range of the interspecific p-dis-

tances within the latter genus. Interspecific p-distances

within Callopanchax range from 5.2—7.6 between C. oc-

cidentalis, C. toddi, and C. sidibei, and up to 14.2 with

the inclusion of C. monroviae. Interestingly the intraspe-

cific distances, even between geographically close collec-

tion localities, are comparatively high (Table 2).

In all resulting phylogenies, C. monroviae splits off first

with respect to the remaining three taxa, which form an

unresolved trichotomy (Fig. 6); this is congruent with the

observed differences in colouration and morphology. As

in Murphy et al. (1999), the distinct position of C. sidibei

©OZFMK

is supported. However, these results should be seen as a

first preliminary insight into the species relationships in

Callopanchax as the dataset is currently too small for de-

veloping well supported, stable phylogenetic hypotheses,

and should be expanded both in number of specimens and

genes in further analyses, especially with regard to the

large genetic distances between the species.

DISCUSSION

The distinctness of Callopanchax sidibei with regard to

its congeneric species was recognized very early after the

first commercial import to the Netherlands (Busch 1993;

Wildekamp 1996), but nothing was known about its ori-

gin, distribution, and habitat. The second author was able

to collect this species in 1997 with the help of the Sidibe

family, who are engaged in the ornamental fish trade and

were probably the first to export this species from Guinea.

Callopanchax sidibei is currently only known from one

locality; other Callopanchax collections in this area con-

tain either C. toddi or C. cf toddi (Fig. 1).

Interestingly in wild males two different caudal fin shapes

exist: a more elongate and truncate form (Fig. 2A) and a

shorter and rounded one (Fig. 2C), of which only the lat-

ter turns up in aquarium bred specimens. In general the

red pigmentation pattern on the body and fins is quite vari-

able (Fig. 2A—C).

As for C. toddi, currently no other colour morphs are

known from C. sidibei, unlike the orange (or red) and blue

phenotypes of C. occidentalis and C. monroviae (Fig.

5A-C, see also figures e.g. in Eberl 1999; Seegers 1997;

Van den Nieuwenhuizen 1974; Wildekamp 1996). These

colour morphs are usually sympatric with the more com-

mon form which is the blue morph in C. monroviae and

the red in C. occidentalis (Berkenkamp & Etzel 2003;

Wildekamp 1996; Wildekamp & Van der Zee 2003 ). One

species, C. huwaldi, which 1s considered as junior syn-

onym to C. occidentalis, differs from the latter species in

external appearance only by the absence of yellow and or-

ange pigments (see colour figure in Roloff 1976, p.523).

There is still some discussion about the validity of the

species; protein electrophoretic data and cross-breeding

experiments by Berkenkamp & Etzel (2003) and Etzel &

Berkenkamp (1981) show differences between the two

species, but protein electrophoretic data by Romand

(1985) do not differ. DNA data by Murphy et al. (1999)

place C. hwwaldi close to C. occidentalis, but the data set

was to limited for further conclusions.

Up to now, several other collections from an area close

to the type locality of C. huwaldi were made e.g. by the

second author, which contained small numbers of blue

Bonn zoological Bulletin 57 (1): 3-14

12 Rainer Sonnenberg & Eckhard Busch

males between red ones. Roloff (1974) reported a sym-

patric occurrence of C. occidentalis and C. toddi, collect-

ed by Chaytor close to Njala, which is also within this area,

and is most probably an occurrence of sympatric red and

blue C. occidentalis males and not a sympatry of the re-

ported species.

The second author was able to breed the red and the blue

phenotype from Largo (SL 89) and Mabeimah (SL 93/46)

several times in the aquarium and it seems that the blue

phenotype is recessive with regard to the red, which is sup-

ported by the observation that the blue phenotype is usu-

ally rare in nature (pers. observation E. Busch). The fe-

males of both forms show no differences.

In case of C. sidibei it can be excluded that it is just a

colour form of C. toddi or C. occidentalis. Beside differ-

ences in body colouration the red pigmentation pattern al-

so differs from all populations of C. toddi and C. occiden-

talis, whereas in the cases of C. monroviae and C. occi-

dentalis, including C. huwaldi, only the yellow and orange

pigments are lost but the red pigmentation pattern is sta-

ble. This is also supported by DNA data, in which C.

sidibei is placed as separate lineage related to C. occiden-

talis and C. toddi (Fig. 6 and Murphy et al. 1999).

The recent collections of the second author in Guinea pro-

vide additional information about the distribution of Cal-

lopanchax, and extends the areal of the genus further

northwards along the coast (Fig. 1). It should be noted that

several of the populations collected so far show combi-

nations of characters, which do not agree with the current

diagnoses of C. toddi and C. occidentalis (Clausen 1966;

Wildekamp & Van der Zee 2003). Further studies will

show whether these species are more variable than pre-

viously known, or if the divergent populations represent

distinct units which might be accepted as separate

species. These fishes are currently listed here as C. cf. tod-

di (see Fig. 1 and as example Fig. 3A & B).

COMPARATIVE MATERIAL

Callopanchax huwaldi: ZFMK 12663-12667, Sierra

Leone, 20 km von Moyamba, Ngabu (8°10’ N, 12°28’ W).

Callopanchax monroviae: ZFMK 41629-41630, Liberia,

Paynesville, L 97 (6°17? N, 10°43’ W); ZFMK

41631-41634, aquarium bred, strain from Liberia, Pay-

nesville, L 97 (6°17’ N, 10°43’ W); RS1533, DNA sam-

ple, aquarium bred, strain from Liberia, Paynesville, L 97

(6°17’ N, 10°43’ W).

Callopanchax occidentalis: ZFMK 41635-41643, first

generation after collection, Sierra Leone, SL 89 Largo

©ZFMK

A new Callopanchax from Guinea 13

TODI534

TODI535

eTODI758

OCC1532

OCC1530

SID1536

MONI533

GERI381

GUI1404

—

Fig. 6.

B 99 cTOD1758

TOD1I534

TODIS535

OCC1530

OCC1532

98/96 | 100

MONI533

GERI381

GUI1404

—_—i

0.02

Results of the maximum parsimony (A) and Bayesian (B) analyses of the mitochondrial large ribosomal subunit (16S)

fragment. Numbers on nodes are bootstrap values in A and posterior probabilities in B. In the latter, the first values are from the

analysis with Nst=2, the second values from the analysis with Nst=6, if both have the same value, only one value is given. Only

for nodes with 2 75 % bootstrap support and = 95 % posterior probabilities values are shown.

(8°16’ N, 12°09’ W); ZFMK 41644-41647, Sierra Leone,

SL 89 Magbenta (8°24 N, 12°56’ W); ZFMK

41648-41654, aquarium bred strain from Sierra Leone, SL

89 Ngabu (8°15’ N, 12°22’ W); ZFMK 41655-41657,

Sierra Leone, SL 89 Romeni (8°53’ N, 12°43’ W); ZFMK

41658-41671, Sierra Leone, SL 89 Teme Yellah (7°41 N,

11°25’ W); RS1531, DNA sample, Sierra Leone, SL 89

Teme Yellah (7°41’ N, 11°25’ W); ZFMK 41672-41674,

Sierra Leone, SL 89 Waanje River (7°50’ N, 11°22’ W);

ZFMK 41675, Sierra Leone, Ma Barie, SL 93/39 (8°25’

N, 11°45’ W); ZFMK 41676-41681, Sierra Leone, Mog-

bomoh Village, Saspoor Water, SL 99/12 (8°20 N, 12°49’

W); ZFMK 41682, Sierra Leone, Brama School, SL 03/7

(8°22’ N, 12°58’ W); ZFMK 41683-41688, Sierra Leone,

Malai, SL 03/11 (8°45’ N, 12°56’ W); ZFMK

41689-41697, Sierra Leone, Rowala, SL 03/12 (8°47’ N,

11°54’ W); ZFMK 41698-41704, Sierra Leone, Magben-

ta, SL 03/19 (8°24’ N, 12°56’ W).

Callopanchax toddi: ZFMK 41705-41715, Guinea, Mo-

la, GM 08/4 (9°20’ N, 12°55’ W); ZFMK 41716-41723,

Guinea, Takhori, GM 08/9 (10°09’ N, 14°01’ W); ZFMK

41724-41728, Guinea, Katamara, GM 08/10 (10°11? N,

14°03’ W); ZFMK 41729-41735, Guinea, Lamayah, GM

08/11 (10°09’ N, 13°44’ W); ZFMK 41736-41739, Sier-

ra Leone, Sentai, SL 93/15 (9°04’ N, 12°53’ W).

Callopanchax cf. toddi: ZFMK 41740-41744, Guinea,

Gbeteyah Moribayah, GM 08/1 (9°36’ N, 13°20’ W);

ZFMK 41745-41751, Guinea, Mangata Badjana, GM

08/3 (9°38’ N, 13°19’ W); ZFMK 41752-41755, Guinea,

Waralneen GV 08/8 (0237. IN, 13.197 W)3 ZEMIK

41756-41757, Guinea, Kabak, GM 08/12 (9°21’ N, 13°23’

W); ZFMK 41758-41763, Guinea, Kabak, GM 08/13

(ONAN G13 237 W)):

Bonn zoological Bulletin 57 (1): 3-14

Acknowledgements. The authors thank Kouroma Christine Sag-

no, Direction National, Ministere du Development, and Bakary

Keita, Ingenieur des Eaux et Forets, Chef de la Division Faune,

Conakry, Guinea, for providing research permissions and sup-

port, Samba, Ousmane and Seybou Sidibe, Conakry, Guinea, for

help with organisation of the research trip and support in the

field; Barbara Wiese, Wilhelm Kugelmann for support in the

field, Fabian Herder (ZFMK, Bonn) for loan of specimens and

access to the digital X-ray machine; Diethard Tautz for support

and laboratory facilities, Floyd A. Reed and Anja C. Schunke

(all MPI, Plén) and two anonymous reviewers for valuable com-

ments and suggestions on earlier versions of the manuscript, and

Gunther Fleck for the French translation of the abstract.

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©ZFMK

Bonn zoological Bulletin Volume 57 pp. 15-17

| Bonn, April 2010

Gekko ulikovskii Darevsky & Orlov, 1994:

a junior synonym of Gekko badenii Szczerbak & Nekrasova, 1994

Quang Truong Nguyen!3, Andreas Schmitz? & Wolfgang B6hme?3

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

E-mail: nqt2@yahoo.com

2 Muséum d’histoire naturelle, C.P. 6434, CH-1211 Geneva 6, Switzerland;

E-mail: andreas.schmitz@ville-ge.ch

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

E-mail: w.boehme.zfmk@uni-bonn.de

In Vietnam, a total of seven species of Gekko are current-

ly recognized: G. badenii, G. chinensis, G. gecko, G.

grossmanni, G. palmatus, G. scientiadventura, and G.

ulikovskii (Rosler et al. 2004; Nguyen et al. 2009). Both

Gekko badenii and G. ulikovskii were simultaneously de-

scribed in 1994 based on the type series collected from

Ba Den Mountain in Tay Ninh Province and “Gilai-Con

Tum Province” (now Kon Tum Province) near the border

with Laos and Cambodia, respectively (Darevsky & Orlov

1994; Szezerbak & Nekrasova 1994). The diagnostic char-

acters of G. badenii and G. ulikovskii obtained from the

original descriptions are presented in Table 1.

In the key to the species of Gekko from Vietnam, Résler

et al. (2004) distinguished these two species by the num-

ber of interorbital scales (30—37 in badenii versus 40-46

in ulikovskii). Based on the original descriptions, G.

badenii further differs from G. ulikovskii by the follow-

ing characters: smaller SVL (51.5—76.5 mm in G. badenii

vs. 87-108 mm in G. uwlikovskii), smaller ratio of

SVL/TaL (0.82 in G. badenii vs. 0.94—-1.24 in G.

ulikovskii), and larger number of precloacal pores (14-18

in G. badenii vs. 10-15 in G. ulikovskii). During recent

field work in southern Vietnam, a series of G. badenii was

collected in October 2004 by Nguyen Huu Dat from the

type locality on Ba Den Mountain, Tay Ninh Province, and

was subsequently deposited in the Institute of Ecology and

Biological Resources, Hanoi, Vietnam (IEBR 1555,

1562-1564, 1566). Our own examination of this new col-

lection compared with two paratypes of G. ulikovskii de-

posited in the collection of the Zoologisches

Forschungsmuseum Alexander Koenig, Bonn, Germany

(ZFMK 55110-55111) revealed that there are no distinct

differences in morphology between these two species. All

Fig. 1. Gekko badenii (IEBR 1562) from Ba Den Moutain, Tay Ninh Province, Vietnam (alive animal)

16 Quang Truong Nguyen et al.

Fig. 2. Gekko badenii (IEBR 1562) from Tay Ninh Province

(a) and Gekko ulikovskii (Paratype: ZFMK 55110) from Kon

Tum Province (b) (preserved specimens).

diagnostic characters obtained from the original descrip-

tions and examined specimens resemble each other or

rather overlap: SVL (51.5—-106.0 in G. badenii and

87.0-108.0 in G. ulikovskii), ratio of SVL/TaL (0.81—1.28

in G. badenii and 0.94—1.24 in G. ulikovskii), interorbital

scales (30-39 in G. badenii and 35—46 in G. ulikovskii),

precloacal pores (12-18 in G. badenii and 10-15 in G.

ulikovskii), dorsum dark grey with 6—8 light bands in both

species, for further comparison see Table 1. It is noted that

our scalation counts for two paratypes of G. ulikovskii

(ZFMK 55110-55111) are a bit different from the origi-

nal description: interorbitals (number of scales between

middle orbitals, not including granular scales on upper

eyelids) 35-38 versus 40-41 in the original description;

scales around midbody (number of scales around body at

the position between axilla and groin): 114-118 versus

131—136 in the original description. Moreover, the char-

acter “scales around midbody” was erroneously noted as

“dorsals around midbody—DoM” for 11 paratypes [Table

1, p. 74] by Darevsky & Orlov (1994). Although these au-

thors did not mention their method to count the number

of dorsals around midbody, we supposed that this is the

number of scales between lateral folds at midbody. Our

examination of ZFMK 55110-55111 showed that the

Bonn zoological Bulletin 57 (1): 15-17

scales between lateral folds at midbody only range from

84-91 (131-136 scales in the original description).

Therefore, this morphological character should be correct-

ed as “scales around midbody” as it was mentioned for

the holotype (132 scales around midbody).

In terms of priority, the formal description of Gekko

badenii was published in Vestnik Zoologii on 15 April

1994, whereas the description of G. ulikovskii was pub-

lished in the journal Salamandra on 15 May 1994. There-

fore, based on the evidence mentioned above and accord-

ing to the International Commission on Zoological

Nomenclature (ICZN, 1999: Articles 23.1 and 23.3) Gekko

ulikovskii Darevsky & Orlov, 1994 must be regarded as

a junior synonym of Gekko badenii Szczerbak & Nekraso-

va, 1994.

Acknowledgements. NQT acknowledges Nguyen Huu Dat (Ho

Chi Minh City) and Nguyen Van Duc (IEBR) for providing spec-

imens from Tay Ninh Province. We thank Ho Thu Cuc (IEBR)

for translation of the article in Russian. We are grateful to L. Lee

Grismer (La Sierra University) and Thomas Ziegler (Cologne

Zoo) for their comments on the manuscript and providing the

map. This research was partially funded by the German Acad-

emic Exchange Service (DAAD).

Vietnam

Gulf of

Tonkin

Cambodia

Gulfof °

Thailand

Fig. 3. Map showing the type localities of Gekko badenii (Tay

Ninh Province) and Gekko ulikovskii (Kon Tum Province) in

Vietnam.

©OZFMK

Gekko ulikovskii Darevsky & Orlov, 1994: a junior synonym of Gekko badenii Szczerbak & Nekrasova, 1994 ia

Table 1. Comparison of morphological characters of G. badenii Szczerbak & Nekrasova and G. ulikovskii Darevsky & Orlov ba-

ce ee,

Original description

Specimens examined

G. badenii G. ulikovskii G. badenii G. ulikovskii

n=3 n= tt M=S n=2

Sex (M = male, F = female) (2 M, | F) (10 M, 1 F) (3 M, 2 F, (2 M, paratypes)

additional material)

SVL 51.5-76.5 87-108 87-106 102.1-102.7

(mean + SD) (62.3 + 0.7) (101.1 + 5.7) (93.3 + 7.6) (102.4 + 0.4)

Tail length (TaL) 72.0-93.5 80-112 82.8-107.8 95.8-99.6

(mean + SD) (82.7 + 1.07) (94.1411.1,n=9) (96.9412.3,n=4) (97.7+42.7)

SVL/TaL 0.82 0.94-1.24 0.81—1.28 1.03—1.06

Snout length > orbit diameter = yes yes yes

Nostril in contact with rostral, yes yes yes yes

supranasal, two postnasals,

and first supralabial

Suture on upper part of rostral present present present present

Supralabials 12-15 11-14 12-14 11-14

Infralabials 13-14 9-13 11-12 12

Internasals 1-3 0-1 1 0-1

Interobitals 30-37 40-46 36-39 35-38

Gular scales 4-8 - 6-12 7-12

Number of gular scales in - 5 4-5 4-6

contact with infralabials

Scales around midbody ~ 123-136 114-119 114-118

Number of dorsal tubercles 12 = 8-13 11-12

Ventral scales between axilla and groin — 51 52-56 52-54

Ventral scales between latero-ventral folds 29-35 — 31-34 32-34

Subdigital lamellae under fourth toe 18-20 15-20 16-19 18

Precloacal pores 14-18 10-15 12-14 12

Enlarged subcaudals - 56-83 60-68 59-63

Ground color of dorsum brownish grey dark grey dark grey dark grey

(in preservative)

Light dorsal bands 4-8 8 6-8 7-8

REFERENCES

Darevsky IS, Orlov NL (1994) Eine bemerkenswerte neue, gross-

wichsige Art der Gattung Gekko: Gekko ulikovskii sp. nov.

aus Zentralvietnam. Salamandra 30: 71-75

International Comission on Zoological Nomenclature (ICZN)

(1999) International Code of Zoological Nomenclature.

Fourth Edition, Online version

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

nam. Edition Chimaira, Frankfurt am Main

R6sler H, Ziegler T, Vu NT, Herrmann H-W, Bohme, W (2004)

A new lizard of the genus Gekko Laurenti, 1768 (Squamata:

Sauria: Gekkonidae) from the Phong Nha-Ke Bang National

Bonn zoological Bulletin 57 (1): 15—17

Park, Quang Binh Province, Vietnam. Bonner zoologische

Beitrage 53: 135-148

Szezerbak NN, Nekrasova OD (1994) A contribution to the

knowledge of Gekko lizards of Southern Vietnam with descrip-

tion of a new species (Reptilia, Gekkonidae). Vestnik Zoologii

1: 48-52 [in Russian]

Received: 23.04.2009

Accepted: 30.04.2009

Corresponding editor: R. van den Elzen

©OZFMK

18 Book Review

Finfstiick HJ, Ebert A, Weiss I (2010). Taschenlexikon der

Vogel Deutschlands. 684 pp., Quelle & Meyer Verlag

GmbH & Co., Wiebelsheim, ISBN 978-3-494-01471-5;

24,95 €.

For many birdwatchers buying a field guide is the first step

into field ornithology. After identifying their first birds,

prospective birdwatchers might wish to learn more about

the species they just recognized. Reaching this goal is the

aim of the work at hand which tries to inform in a com-

pact format suitable for carrying in a rucksack about all

bird species occurring in Germany. This lexicon 1s basi-

cally a summary of the ,,kompendium der Végel Mitteleu-

ropas™. It contains short chapters on taxonomy, origin of

German and scientific names, occurrence, status, habitat,

and breeding and feeding biology of all species found in

Germany. These are complemented by short descriptions

of appearance, voice, body masses and peculiarities. Each

species is presented with a distribution map and photo-

graphs of almost always-good quality, completed by dia-

grams on phenology and moult. Even species that are

rarely photographed in field guides of birds from Germany

like Black Lark, Siberian Thrush or Northern Parula are

depicted in this book. Generally, the proximity to the

“Kompendium” provides reliability and timeliness of in-

formation. Nevertheless the lexicon is not error-free: Right

in the beginning the explanatory distribution map is in-

correct, as the light blue areas of migration periods and

point occurrences are missing. The distribution area of the

Green Willow Warbler has been displaced to Fennoscan-

dia and Alaska, and superspecies are not always handled

with care, e.g. for the Eastern black-eared Wheatear, Pied

and Cyprus Pied Wheatear are missing as further mem-

bers of the superspecies.

Furthermore, many topics are left out that can be found

yet in the “Kompendium”’, for example information on

population changes, behaviour, moult and conservation

measures. A general preface and introductions to the fam-

ilies are nonexistent. Species are presented in alphabeti-

cal order of the German names, so that possibly longer

lasting thumbing through the book could be necessary if

for example looking up different species of a superspecies

or genus, particularly since there is no register available.

Searching for scientific names is hence impossible. Read-

ers should be familiar with current taxonomic findings and

changes of names, as species like Cory’s Shearwater, Yel-

low Wagtail and Black-eared Wheatear are provided with

Bonn zoological Bulletin 57 (1): 18

Fiinfstiick/Ebert/Weifs 4

Taschenlexikon der Vogel

Deutschlands

Ein kompellier -

Begleiter durch die '

heimische Vogelwelt

a -

ee.

{i}

“

RS eres ace

QUELLE & MEYER |

Qusieiiea eS

new German names (“Sepiasturmtaucher”, ‘“Wiesen-

schafstelze”’, “Mauren- and Balkansteinschmatzer’’).

On the whole the lexicon is well suited for an entry into

bird watching and can be recommended to everyone who

is looking for a supplement to mere field guides. Wish-

ing to seriously deal with field ornithology, though, re-

quires purchasing additional books, such as the more cost-

ly but also more detailed three volumes of “Kompendi-

um der Vogel Mitteleuropas”’.

Kathrin Schidelko

Zoologisches Forschungsmuseum Alexander Koenig,

Bonn

©ZFMK

Bonn zoological Bulletin Volume 57 Issue | pp. 19-29

Bonn, April 2010

A new chameleon of the Trioceros bitaeniatus complex

from Mt. Hanang, Tanzania, East Africa

(Squamata: Chamaeleonidae)

Patrick Krause! & Wolfgang Bohme!

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

E-mails: paddy.krause@web.de; w.boehme.zfmk(@uni-bonn.de

Abstract. A new species of Zrioceros from the montane forests of Mount Hanang in Northwest Tanzania is described.

The only known population of this species is found on Mt. Hanang in Tanzania around 2800 m a.s.l. This chameleon

bears close morphological resemblance to Trioceros sternfeldi (Rand, 1963) from northern Tanzania. In order to infer

phylogenetic relationships within the group, morphological and genetic data (mitochondrial 16S and 12S rRNA genes)

were analysed. The new species could be compared with specimens of different populations of Trioceros sternfeldi (Mts.

Meru, Kilimanjaro and Ngorongoro) and several other representatives of the family (J. hoehnelii, T. ellioti, T: rudis rud-

is, T. bitaeniatus). Results of maximum parsimony, maximum likelihood and Bayesian analyses consistently placed Tri-

oceros n. sp. from Mt. Hanang as a single, independent terminal clade with high genetic divergences towards all other

included members of the group.

Key words. Squamata, Chamaeleonidae, Trioceros, Africa, molecular phylogeny, biogeography

INTRODUCTION

Mt. Hanang represents one of the numerous ancient and

isolated volcanic mountains which characterize the land-

scape of eastern Africa. In northern Tanzania Mts. Kili-

manjaro, Meru, and the Ngorongoro Crater are three of

the better known extinct volcanoes which are frequently

visited by tourists and lie within some of Tanzania’s most

famous national parks. Several biologists identified these

forested East African mountains as centers of endemism

for chameleons of the “rudis group”, a subgroup of the

Trioceros bitaeniatus species complex (Rand 1963; Pil-

ley 2000; NeCas et al. 2003). The Trioceros bitaeniatus

complex comprises various taxa of small, hornless, ovo-

viviparous East African chameleons from the highland re-

gions of eastern Africa, (Uganda, Kenya, Tanzania, south-

ern Ethiopia and Sudan). All members of the group were

placed into the separate subgenus Trioceros by Klaver &

Boéhme (1986) which was recently elevated to full gener-

ic rank by Tilbury & Tolley (2009). In contrast to other

species of the bitaeniatus group, chameleons of the rud-

is subgroup are rather stocky animals with short, broad

heads and heterogeneous scalation. They belong to a se-

ries isolated populations restricted to mountain forests

ranging from the Ruwenzori in the north-west, south to

the mountains east of Lake Malawi with outliers on Mt.

Kilimanjaro, Mt. Meru and Ngorongoro in northern Tan-

zania. The Tanzanian forms of this subgroup were initial-

ly assigned to a rudissubspecies (Chamaeleo rudis stern-

feldi) by Rand (1963), but this was long believed to be of

rather specific than subspecific rank by several authors

(e.g. NeCas et al. 2003). The independent taxonomic sta-

tus of this form was subsequently demonstrated on the

ground of morphological (Klaver & Bohme 1997; Krause

2006) and genetic data (Krause 2006).

The speciation of the bitaeniatus group as a whole seems

closely related to the formation of the East African rift val-

leys when volcanism resulted in a separation of the an-

cestral populations. This can therefore be regarded as a

major factor that gave rise to present day species (Pilley

2000; Wasser & Lovett 1993).

Three specimens of a curious chameleon from the Trio-

ceros bitaeniatus- species complex were collected on Mt.

Hanang at 2800 m altitude in March 2002, morphologi-

cally similar to 7! sternfeldi (Rand, 1963) but with a very

distinct colouration. This form is known only from the Mt.

Hanang Catchment Forest Reserve (4° 25’ to 4° 35’ S and

35° 20’ to 35° 25’ E), situated about 4 km from the dis-

trict capital Katesh and 70 km from the town Babati in

the administrative region of Arusha in northern Tanzania

(Figure 1).

20 Patrick Krause & Wolfgang BOhme

Fig. 1.

Map of Tanzania with distribution of the discussed

species. 1=Mt. Hanang: 7. hanangensis n. sp.; 2=Ngorongoro:

T. sternfeldi; 3=Mt. Meru: T. sternfeldi; 4= Mt. Kilimanjaro: 7.

sternfeldi; 5=Usambara Mts.: 7. sternfeldi.

MATERIALS AND METHODS

All specimens were photographed in situ to record their

natural coloration and habitus. Eversion of hemipenes in

freshly dead specimens was done in the by injecting 70

% Alcohol into the organs (Glaw et al. 1999). The collect-

ed voucher specimens were deposited in the Zoologisches

Forschungsmuseum Alexander Koenig in Bonn (ZFMK).

Additional material was examined from the Zoologische

Staatssammlung Miinchen (ZSM).

In order to test for morphological variation of the

paratypes, chameleon body measurements were obtained

with a digital caliper and evaluated with morphometric

analyses methods (Krause 2006). Statistical analysis was

performed using the GLM Multivariate procedure as im-

plemented in SPSS 12 which provides analysis of vari-

ance for multiple dependent variables by one or more fac-

tor variables or covariates. In the present study the mor-

phometric measurements (Krause 2006) excluding body

length were chosen as dependent variables and the mor-

phometric variable “snout-vent-length” was included as

covariate to exclude the effect of body size on the other

measured parameters. After an overall f-test has shown

significance, post hoc tests (Scheffé) were applied to eval-

uate differences among the specific means. Probability val-

ues less than 0,05 were assumed to infer biological sig-

nificance.

Bonn zoological Bulletin 57 (1): 19-29

Molecular data could be obtained from tissue samples

(muscle or liver, fresh or preserved in 98% ethanol) of the

following taxa: Trioceros n. sp., (Mt. Hanang, Tanzania),

Trioceros rudis (Uganda), Trioceros sternfeldi (Mt. Kili-

manjaro, Mt. Meru, Tanzania). In order to analyse the phy-

logenetic relationships within the bitaeniatus group the

following taxa were also included in the analysis: 7. elli-

oti, T. hoehnelii, T. bitaeniatus, T: narraoica, T. werneri,

T. jacksonii, T. deremensis and T: johnstoni. Kinyongia

uthmoelleri, Chamaeleo dilepis. Uromastyx acanthinura

and Leiolepis belliana were used as outgroups. All spec-

imens included in this study are listed in Tab. 1 and se-

quences were submitted to GenBank (accession numbers

AY927239-AY 927276). DNA was extracted using the

Chelex method (Walsh et al. 1991). In this study a 5%

Chelex solution (2,5 mg Chelating Resin [Iminodiacetic

Acid; Sigma Aldrich] in 50ml ddH 0) was used (Schmitz

A 2003). Two mitochondrial genes which encode for the

16S und 12S subunits of the ribosomes (16S and 12S

rRNA genes) where chosen for molecular analysis.

Primers 16SA (light chain 5’- CGC CTG TTT ATC AAA

AAC AT -3’) and 16SB (heavy chain 5’- CCG GTC TGA

ACT CAG ATC ACG T) of Palumbi et al. 1991 were used

to amplify a section of the 16S rRNA gene. The primers

12SA-L (light chain; 5’ - AAA CTG GGA TTA GAT ACC

CCA CTA T - 3’) and 12SB-H (heavy chain; 5’ - GAG

GGT GAC GGG CGG TGT GT - 3’) of Kocher et al.

(1989) were used to amplify a section of the 12S rRNA

gene. PCR cycling procedure for both gene-fragments was

as follows: Initial denaturation step: 90s at 94°C, 33 cy-

cles: denaturation: 45s at 94°C, primer anealing for 45s

at 55°C, extension for 90s at 72°C.

Purification of the PCR products was conducted follow-

ing standard methods using QIAquick PCR Purification

Kit (Qiagen), sequencing reactions of the obtained ampli-

fied DNA including cycle sequencing (standard proce-

dures) were conducted by MWG-Biotech and Max

Planck Institut fiir Zichtungsforschung (MPI) in Cologne

using ABI Prism DNA automatic sequencer.

Sequences were aligned using the computer program

ClustalX (default parameters, Thompson et a. 1997). The

alignment was subsequently adjusted manually using the

computer program BioEdit (Hall 1999).

33 sequences (combined 16S and 12S rRNA genes) could

be obtained, each comprising 861 bp, referring to the

aligned sequences including gaps (16S: 479 bp; 12S: 382

bp). The alignment gaps were treated as missing charac-

ter. The entire 16S+12S alignment was included in the

analyses. As with two specimens (ZFMK 63223, 70752)

only 12S sequences could be obtained, we conducted a

separate analysis using the 12S dataset only.

©ZFMK

A new chameleon of the Zrioceros bitaeniatus complex from Mt. Hanang, Tanzania 21

Fig. 2. Sulcal (left) and asulcal (right) view of the hemipenis

of Trioceros hanangensis n. sp. from Mt. Hanang, Tanzania.

Data were analysed using maximum likelihood and par-

simony optimality criteria in PAUP 4.0b10 (Swofford

2002), further we used Bayesian inference as implement-

ed in MrBayes 3.1.1 (Ronquist & Huelsenbeck 2003). For

maximum likelihood, a heuristic search was conducted

with 10 replicates of random stepwise addition and tree

bisection-reconnection (TBR) branch swapping. Relative

branch support in phylogenetic analysis was evaluated

with 100 bootstrap pseudoreplicates. Parsimony analyses

used heuristic searches with 100 random additions of taxa

and tree bisection-recombination branch swapping. When

more than a single most parsimonious tree was found, a

strict consensus tree was generated. Confidence in the

nodes was assessed by 2000 bootstrap pseudoreplicates

(Felsenstein 1985) with random addition of taxa. Only

bootstrap supports of 70 % and higher were considered

as reliable, as such values were found to indicate 95 %

probability of correct topology (Hillis & Bull 1993). For

Bayesian analyses a GTR+I+G model was chosen accord-

ing to the optimal model and parameter-values based on

the AIC criteria determined in MrModeltest 2.2 (Nylan-

der 2004). Two MCMC analyses for 10° generations each

were conducted. The topologies of the sampled trees were

used to generate a strict consensus tree. Only clades with

a posterior probability of 95% or greater where consid-

ered to be significantly supported.

RESULTS

Trioceros hanangensis n. sp.

Holotype. ZFMK 82369, adult male with everted hemipe-

nis. Locality: Tanzania: Mt. Hanang Forest Reserve near

the village Jorodom (4° 25’ to 4° 35’ S and 35° 20’ to 35°

25’ E) at 2800 m, collected by P. Krause, March 2002.

Bonn zoological Bulletin 57 (1): 19-29

Paratypes. ZFMK 82368, subadult male, same data as the

holotype; ZSM 111/1981/1- ZSM 111/1981/20 collected

by L. Uthmoeller (1937) from the same locality as the

holotype.

Diagnosis. A medium sized, stocky chameleon resembling

a typical member of the Trioceros bitaeniatus complex

(Rand 1963). The maximum total length is 138 mm; the

tail comprises an average of 45 %, males with longer tails.

Adult males and females have a similar snout-vent-length

(SVL) the females being slightly larger. Body scalation

is heterogeneous, with strongly enlarged denticulate scales

in two distinct rows on the sides of the body. The gular

crest is weak to medium, consisting of scales 1.5 times

the length of their maximum width. A ventral crest is clear-

ly visible in bigger specimens to almost indistinct in small-

er ones. No rostral appendix is visible like in other mem-

bers of the bitaeniatus complex (Trioceros narraioca

Ne€as, Modry & Slapeta, 2003; Trioceros hoehnelii Stein-

dachner, 1891).

This chameleon bears a close morphological resemblance

to Trioceros sternfeldi from northern Tanzania (Mts. Kil-

imanjaro, Meru and Ngorongoro Crater) but shows a dis-

tinct colour pattern, which differs remarkably from 7.

sternfeldi (Sternfeld 1912a, Figures 3, 5, 6). Further, it pos-

sesses scattered melanophores on the sulcus of the

hemipenis which are unique as compared to 7. sternfeldi.

Despite a close genetic relationship with T. e//ioti, the new

form can clearly be distinguished by absence of the dis-

tinct longitudinal groves on the gular pouch, a more het-

erogeneous scalation and a stouter body as compared with

T. ellioti, which is rather slim (Gtinther 1895; Spawls et

al. 2002).

The only known population of this species is found on Mt.

Hanang above 2800 m in montane bushland.

Description of holotype. Head short and broad, distance

from tip of snout to end of parietal crest less than twice

greatest width between the temporal ridges. Parietal crest

not swollen posteriorly, being almost straight in profile.

No rostral projection at the point of fusion of canthi ros-

trales. Occipital flaps absent. Ventral crest moderately de-

veloped, composed of cone-shaped, subequal scales,

longest on throat, about 3/4 of eye diameter. Throat ho-

mogenously covered with small scales, no indication of

longitudinal grooves. Head length (from tip of snout to

posterior margin of casque) 21 mm. Height of casque

(from angle of mouth to top of casque) 16 mm, slightly

less than length of mandible (20 mm). Caudal side of

casque homogenously covered with fine granular scales.

Parietal crest consists of 15 enlarged elongate scales. Two

other diagonal (paraparietal) crests visible on each side of

©OZFMK

22 Patrick Krause & Wolfgang Bohme

Fig. 3. Holotype of Trioceros hanangensis n. sp. (male, ZFMK 82369) from Mt. Hanang, in life.

head, consisting of 4 enlarged tubercular scales. These

originate in the immediate vicinity of the parietal crest and

run rostrolaterally towards the orbit. Lateral crests com-

posed of 4 enlarged, prominent pointed scales, extending

to the supraorbital crest (11 pointed scales) and to canthi

rostrales (5 scales).

Body rather stout. Dorsal crest well developed, composed

of scales in groups of three anteriorly and four posterior-

ly. Increasing posteriorly in size within the group, giving

the back a serrate appearance. Dorsal crest extends to the

tail and becomes indistinct within its terminal third. The

Fig. 4.

Bonn zoological Bulletin 57 (1): 19-29

Trioceros hanangensis n. sp. (subadult male, ZFMK

82368) from Mt. Hanang.

first scales of the series of three just behind the head are

longest, reaching 1.7—2 mm length. Ventral crest moder-

ate to weak, best developed on the throat; posteriorly

scales are shorter but still visible up to posterior belly.

Scalation heterogeneous. Sides of body, tail and upper sur-

faces of limbs covered with irregularly sized, convex,

small scales interspersed with larger convex scales. Lar-

gest of these form a narrow row running from neck just

behind temporal crests to tail. A second row of slightly

smaller scales runs from shoulder to hind leg. This row

is much less distinct and more irregular. Vent, inner side

Trioceros sternfeldi (male) Ngorongoro Crater.

©ZFMK

A new chameleon of the 7rioceros bitaeniatus complex from Mt. Hanang, Tanzania

Fig. 6.

Trioceros sternfeldi (male) Mt. Kilimanjaro.

of legs, soles and underside of the tail almost homogene-

ously scaled. No tarsal spurs.

Hemipenis. Smooth margined calyces cover the asulcal

side of the hemipenis, extending as far as the sulcal lips.

Apical tip characterised by two pairs of finely serrate ro-

tulae of roughly semicircular shape, positioned towards

the sulcal side and extending around lateral side of

hemipenis. Scattered melanophores could be observed in

the sulcus (Figure 2).

Colouration and pattern. Vivid green with lateral yel-

low bar following the row of enlarged scales running from

neck to the base of tail. Head light blue except gular re-

gion. Eyeballs bright blue to turquoise.

Variation of the paratypes. Analysis of morphometric

variation between male and female specimens of Trioceros

n. sp. from Mt. Hanang resulted in males having signifi-

cantly broader heads than females ( x = 13,09 mm vs. x

= 12,59 mm; p = 0,030) and a tendency to increased ru-

gosity in adult males. There is no apparent statistical dif-

ference in snout-vent-length (SVL) comparing both sex-

Fig. 7. Mt. Hanang from Katesh, SSW side of the mountain.

Bonn zoological Bulletin 57 (1): 19-29

es. (males: SVL = 72,30 mm, N = 9 / females: SVL =

75,52 mm, N = 14). No significant differences in relative

tail length (TL) between sexes could be observed (males:

TL = 66,22 mm / females: TL = 62,71 mm).

Distribution. Only known from Mt. Hanang Catchment

Forest Reserve (4° 25’ to 4° 35’ S and 35° 20’ to 35° 25’

E) situated about 4 km from the district capital Katesh and

70 km from the town Babati in the administrative region

of Arusha. The new species is named after the type-local-

ity Mount Hanang.

Ecological remarks and habitat preference.: 7) hanan-

gensis n. sp. was found in upland moorland at 2800 m on

the moister north-eastern side of Mt. Hanang. Despite a

prevaling semi-arid climate with 750-1500 mm annual

rainfall, a pronounced mist effect on the eastern and north-

eastern slopes is leading to a maximum of 2000 mm pre-

cipitation per year at the higher regions of the mountain.

The sampled specimens where observed during daytime

in bushy highland vegetation (with Hypericum sp., Myr-

sine africana, Protea sp.) near the source of the river Him-

it, roosting on twigs at a height ranging from 100—150 cm

from the ground.

Phylogenetic affinities. In order to infer the phylogenet-

ic relationships between the different members of the

bitaeniatus group on the basis of molecular data maximum

parsimony (MP), maximum likelihood (ML) and Bayesian

analyses (PP) were conducted. Maximum likelihood

analysis was conducted with a reduced dataset, due to

computational constraints. Sequences obtained from the

combined 16S+12S rRNA mitochondrial genes were

analysed together with gene-fragments of Trioceros

sternfeldi, T. rudis, T. ellioti, T: hoehnelii, T: bitaeniatus,

T. narraoica, T. werneri, T. jacksonii, T: deremensis and

T: johnstoni. The two included specimens of 7? hanangen-

sis. sp. were found in a well supported clade in the MP

and Bayesian trees (MP: 99; PP: 1.00), separating them

Fig. 8. Type locality of Trioceros at 2800 m altitude.

©OZFMK

100

1.00,

0.99

1.00

87 al

0.99

0.82

Fig. 9.

Patrick Krause & Wolfgang B6hme

1.00)

67 72

0.86 0.99

1.00

100

1.00

100

1.00.

Kinyongia uthmoelleri Mt. Hanang, Tanzania

Chamaeleo dilepis Mt. Hanang, Tanzania (V)

Trioceros jacksonii Nairobi, Kenya

Trioceros narraoica Mt. Kulal, Kenya

Trioceros kinetensis Sudan

Trioceros ellioti Kakamega Forest, Kenya (I)

Trioceros ellioti Kakamega Forest, Kenya (Il)

Trioceros ellioti Nandi Hills, Kenya

Trioceros ellioti Kenya

- Trioceros ellioti Ruwenzori Mts., Uganda (I)

Trioceros ellioti Ruwenzori Mts., Uganda (Il)

Trioceros hanangensis n. sp. Mt. Hanang, Tanzania (I)

Trioceros rudis Ruhiza, Uganda

Trioceros rudis Bwindi Forest, Uganda

Trioceros sternfeldi Mt. Kilimanjaro, Tanzania

- Trioceros sternfeldi Mt. Meru, Tanzania (l)

Trioceros sternfeldi Mt. Meru, Tanzania (II)

Trioceros hoehnelii Eldama Ravine, Kenya

Trioceros hoehnelii Uganda

Trioceros hoehnelii Mt. Kenya (I)

~ Trioceros hoehnelii Mt. Kenya (Il)

Trioceros bitaeniatus Arusha, Tanzania

Trioceros bitaeniatus Mt. Hanang, Tanzania (I)

Trioceros bitaeniatus Mt. Hanang, Tanzania (Il)

Trioceros bitaeniatus Mt. Hanang, Tanzania (III)

Trioceros bitaeniatus Mt. Hanang, Tanzania (IV)

Trioceros werneri Udzungwa Mts., Tanzania

Trioceros deremensis Usambara Mts., Tanzania

Trioceros johnstoni Bwindi Forest, Uganda

— Leiolepis belliana

Uromastyx aegyptia

- Trioceros hanangensis n.sp. Mt. Hanang, Tanzania (Il)

Strict consensus parsimony tree derived from the combined fragment of the mitochondrial 12S and 16S rRNA genes.

Parsimony bootstrap values are given above the nodes (bold), lower values are posterior probabilities produced by Bayesian infe-

rence. Nodes with less than 50 % bootstrap support are not labelled.

Bonn zoological Bulletin 57 (1): 19-29

©ZFMK

A new chameleon of the Zrioceros bitaeniatus complex from Mt. Hanang, Tanzania 25

100

| 74

Uromastyx aegyptia

Chamaeleo dilepis Mt. Hanang, Tanzania (V)

Trioceros jacksonii Nairobi, Kenya

Trioceros narraoica Mt. Kulal, Kenya

Trioceros bitaeniatus Mt. Hanang, Tanzania (IV)

Trioceros rudis Bwindi Forest, Uganda

Trioceros sternfeldi Mt. Kilimanjaro, Tanzania

Trioceros sternfeldi Mt. Meru, Tanzania (Il)

Trioceros hanangensis n.sp. Mt. Hanang, Tanzania (Il)

Trioceros ellioti Kakamega Forest, Kenya (I)

~ Trioceros ellioti Nandi Hills, Kenya

Trioceros ellioti Ruwenzori Mts., Uganda (Il)

Trioceros hoehnelii Mt. Kenya (Il)

- Trioceros hoehnelii Eldama Ravine, Kenya

Trioceros werneri Udzungwa Mts., Tanzania

Trioceros deremensis Usambara Mts., Tanzania

- Trioceros johnstoni Bwindi Forest, Uganda

Fig. 10. ML tree derived from the combined fragment of the mitochondrial 12S and 16S rRNA genes. Bootstrap values are gi-

ven above the nodes, nodes with less than 50 % bootstrap support are not labelled.

Bonn zoological Bulletin 57 (1): 19-29

©ZFMK

26 Patrick Krause & Wolfgang Bohme

0.56

68 98. ee

92 97

0.95

64 74

0.99

93 94

0.84 |

0.99

59 59

0.69

0.98 1.00

0.97 |

7471

1.00

Uromastyx aegyptia

Chamaeleo dilepis Mt. Hanang, Tanzania (V)

Trioceros jacksonii Nairobi, Kenya

Trioceros narraoica Mt. Kulal, Kenya

Trioceros bitaeniatus Arusha, Tanzania

Trioceros bitaeniatus Maralal, Kenya

Trioceros bitaeniatus Mt. Hanang, Tanzania (I)

Trioceros bitaeniatus Mt. Hanang, Tanzania (Il)

Trioceros bitaeniatus Mt. Hanang, Tanzania (III)

Trioceros bitaeniatus Mt. Hanang, Tanzania (IV)

Trioceros rudis Ruhiza, Uganda

Trioceros hoehnelii Uganda

Trioceros hoehnelii Eldama Ravine, Kenya

Trioceros hoehnelii Mt. Kenya (I)

Trioceros hoehnelii Mt. Kenya (Il)

Trioceros sternfeldi Mt. Kilimanjaro, Tanzania

Trioceros sternfeldi Mt. Meru, Tanzania (I)

Trioceros hanangensis n.sp. Mt. Hanang, Tanzania (i)

Trioceros ellioti Kakamega Forest, Kenya (I)

Trioceros ellioti Nandi Hills, Kenya

Trioceros cf. bitaeniatus Sudan

Trioceros rudis Ruwenzori Mts., Uganda

Fig. 11. Consensus tree derived from Bayesian inference using the mitochondrial 12S gene. Posterior probabilities are given abo-

ve, ML bootstrap values (bold) and MP bootstrap values (inverse) below. Nodes with less than 50 % bootstrap support are not la-

belled.

Bonn zoological Bulletin 57 (1): 19-29

©OZFMK

Table 1. Specimens included in this study.

Species

Kinyongis uthm celier’

Cham seleo dilepis

Trioceres ellioti

Trioc eros ellioti

Troceros ellioti

Trocercs ellioti

Trioceros ellioti

Tricc eros hanangensis 5p. nov

Triocercs hanangensis sp. nov

Trioceros rucis

- Thoceros rudis

Trioc eros rucis

Trice eros kinetensis

Trioceros sternfeldi

Trice eros sternfeldi

Triocercs sternfeldi

Trocercs hoehnelii

Trioceros hoehnelit

Trioceros hoehneli

Trioceros hoehnelit

Trioceros hoehnelii

Trocercs bitaeniatus

Trocercs biaenistus

Trocercs bitsernatus

Troceros bitaematus

Tnocercs bitzeniatus

Trocerce biaeniatus

Trioc eros nairaoics

Trioc eros jacksonii

Triocercs wemeri

Troceros derem ensis

Trioc eros johnstoni

Leiolepic bellians

Uram astyx segyptia

Locality

Nt. Hanang, Tanzania

Mt. Hanang, Tanzania (V)

Kakamegs Forest, Kenya [I)

Kakamega Forest, Kenya (II)

Nandi Hills, Kenya

Kenys

Ruwenzori Mts., Uganda (1)

Ruwenzori Nits., Uganda (II)

It. Hanang, Tanzania (1)

Nit. Hanang, Tanzania (II)

Ruhiza, Uganda

Bwindi Forest, Ugands

Uganda

S$ Sudan

Mt. Kilimanjaro, Tanzania

Nit. Meru, Tanzania (I)

Mit. Meru, Tanzania {II}

Mt. Kenya (1)

Ugands

It. Kenya (Il)

Eldama Ravine, Kenya

Ugsnda

Wit. Hanang, Tanzania (1)

Wt. Hanang, Tanzanis {II}

Mit. Hanang, Tanzania (III)

Wit. Hanang, Tanzania (IV)

Arusha, Tanzania

Maralal, Kenya

Mt. Kulsl, Kenya

Nairobi, Kenya

Udzungwa IMits., Tanzania

East Usambara Mits., Tanzania

Bwindi Forest, Uganda

Egypt

Thailand

Collection number

A new chameleon of the Trioceros bitaeniatus complex from Mt. Hanang, Tanzania

Accession number 165/125

ZFMK 82191 DOQ297231 DQz297245

ZFMK 82381 AY927263, AN 927244

ZFMK 82058 DOQ397278 DQ397258

ZFMK 20835 DO397279 DOQ297259

ZFNIK 82057 DO3S7225 DQ397229

ZFMK 70818 DO397280 DOQz297260

ZFMK 638287 DQ397281 DQ397256

ZFMK 63227 DOQ397282 DQ397257

ZFMK 82368 DQ397283 DOQ397254

ZFNIK 82369 DOQ397284 DQ397255

CAS 201781 DOQ397223 DOQ397223

CAS 201711 DOQ397285 DQ397252

ZFMKE3223 - DOQ397277

ZFMK 29714 DOQ397286 DQ397253

ZFMK 68629 DO397287 DOQ397261

TO68 (prov. ID) DO397288 DOQz97263

ZFMK 82250 DO397289 DO397262

ZFMK 70678 DO297290 DOQ397265

voucher not collected DOQ297291 DOQ297266

ZFMK 66578 DO397292 DQ397264

ZFMK 82088 DOQ397224 DQ397238

voucher net collected DQ397293 DQ397267

ZFIMIK 74949 DOQ3297222 DOQ397237

ZFMK 74952 DOQ397294 DO397265

ZFMK 82365 DOQ397295 DQ397270

ZFMK 82266 DO397296 DO397271

ZFNIK 82251 DO397297 DOs9g7272

ZFMK 70752 - DOQ397276

ZFMK 73958 DOQ397298 DO397268

CAS 199070 DO397226 DOQ397240

ZFMK 70452 DOQ397299 DOQ397274

CAS 168888 DOQ397200 DOQ297273

CAS 201597 DOQ397301 DOL97275

GenBank ABN3 1986 ABN3 1969

GenBank AB03 1994 AB03 1977

from all other included taxa. In the tree resulting from the

maximum likelihood analysis, the single included speci-

men of 7. hanangensis n. sp. clustered together with T.

ellioti (ML: 74). Genetic distances to all other included

taxa of the bitaeniatus group were relatively high (2,2—4

%) and were within the range which could be found be-

tween well defined species. This confirms the independ-

ent specific status of Trioceros hanangensis n. sp.

Phylogenetic inference of the combined 16S+12S rRNA

dataset revealed a sistergroup relationship with 7° e/lioti,

which was weakly to moderately supported with all meth-

ods (MP: 67; PP: 0.86; ML: 74). A genetic distance of

2,05-2,18 % could be observed between Trioceros hanan-

gensis n. sp and the included specimens of 7: ellioti. Us-

ing only the 12S fragment for phylogenetic inference, a

much higher support for this sistergroup relationship could

be observed (MP: 89; PP: 0.99; ML: 81). Nevertheless,

the relationships of 7: hanangensis n. sp. towards the oth-

er taxa of the bitaeniatus group, especially towards T.

sternfeldi could not be finer resolved with the available

limited genetic dataset, resulting in a large polytomy in

all trees.

Bonn zoological Bulletin 57 (1): 19-29

Conservation status. The forested area on Mt. Hanang

is classified as a Catchment Forest Reserve covering ap-

prox. 40 km? which is relatively small compared to oth-

er reserves in Tanzania. The genuine forest of Mt. Hanang

has been drawn back to the very base of the mountain and

is under increasing threat by illegal logging, fires and

drought. Due to the small size of the reserve, even minor

disturbances are likely to affect the habitat of forest de-

pendant species like 7. hanangensis n. sp. Along with the

well-known mountains in northern Tanzania like Meru,

Kilimanjaro, Ngorongoro and the Eastern Arc, Mt.

Hanang is a further example of a remnant mountain for-

est with quite unique climatic and ecological conditions.

For this reason the protection and management of the re-

maining forests on Mt. Hanang should receive urgent pri-

ority.

DISCUSSION

Trioceros hanangensis n. sp. is a member of a species

complex which is closely associated with the forest cov-

ering many of the East African mountains. These

©OZFMK

28 Patrick Krause & Wolfgang Bohme

chameleons represent a subgroup of the bitaeniatus-group

(the “rudis group”’sensu Rand 1963). In contrast to other

species within the bitaeniatus group, populations of the

rudis group are restricted to mountain forests with wide-

ly disjunct distribution, each population bearing specific

morphological peculiarities distinguishing it from the oth-

ers as first observed by Rand (1963).

Like the bitaeniatus group as a whole, the different forms

of the rudis group can most easily be understood as relics

of a period equating with the last pluvial period of the

Pleistocene when the rainfall in East Africa was higher and

the forest was much more extensive than today. Drier pe-

riods following the pluvial maxima resulted in a retreat

of the forests from the lower elevations to their present

positions on the higher mountains. As a consequence the

chameleon populations living in these forests (Wasser &

Lovett 1993) were also fragmented. Different selection

pressures formed the various taxa (7. schubotzi, T. stern-

feldi or T. hanangensis n. sp.) which are today restricted

to relatively small areas or certain types of habitat. In con-

trast to the mountains of the Eastern Arc, most volcanoes

in northern Tanzania are relatively young and result from

volcanic activity persisting from the Oligocene (37 myr

ago) to the Quaternary. As a consequence, the origin of

the different rudis-forms cannot lie very far in the past

which makes it a rewarding subject for the investigation

of beginning speciation processes.

Despite a morphological similarity with Trioceros stern-

feldi (Mts. Meru, Ngorongoro and Kilimanjaro) 7) hanan-

gensis n. sp. can be regarded as valid species, chief indi-

cators being a distinct colour-pattern, presence of scattered

melanophores in the sulcus of the hemipenis (not present

in 7: sternfeldi) and the high degree of genetic differen-

tiation towards 7. sternfeldi and the other members of the

group. Molecular analysis of the combined 16S+12S

dataset shows a genetic differentiation between 2,2-4 %,

which is clearly above the intraspecific level and confirms

the independent specific status of 7) hanangensis n. sp.

A weakly supported sister-group relationship towards T.

ellioti is indicated by all the given trees (MP: 67; PP: 0.86;

ML: 74). In contrast to T e/lioti T7 hanangensis n. sp. does

not possess the distinct longitudinal groves on the gular

pouch as present in 7: e//ioti, it has a more heterogeneous

scalation and a stouter body than 7: ellioti (Giinther 1895;

Spawls et al. 2002). Further, 7) e//ioti has a wide distri-

bution ranging from western Kenya, the northern shore of

Lake Victoria in Uganda, western Uganda, Rwanda and

Burundi and occurs in open bushland and grassland.

Limited by the available genetic dataset, phylogenetic re-

lationships towards the other members of the bitaeniatus

Bonn zoological Bulletin 57 (1): 19-29

group could not be sufficiently resolved . Phylogenetic

affinities of 7) hanangensis n. sp. and T. sternfeldi can on-

ly be assumed by means of morphological data: specimens

of 7. sternfeldi from Mt. Kilimanjaro have dark eyeballs,

in contrast to 7. sternfeldi from Ngorongoro and 7! hanan-

gensis n. sp. which both have light blue eyeballs and are

more alike in general colouration. The parietal knob ob-

served by Rand (1963) associated with specimens from

Mt. Meru and Mt. Kilimanjaro could not be observed in

specimens from Mt. Hanang respectively Ngorongoro.

These observations support the assumption that 7’ hanan-

gensis n. sp. 1s more closely related to 7. sternfeldi from

Ngorongoro than with the sternfeldi-populations from

Meru and Kilimanjaro. The geography of the region would

have allowed geneflow between the populations of

Hanang and Ngorongoro in the past: particularly to the

north-west of Mt. Hanang a continous forest could have

existed during moister periods linking the habitat of 7

hanangensis n. sp. with the Nou- and Marang forests and

further north with Oldeani (3188 m) and Ngorongoro

(3000 m). Today, a dry and hot valley separates the moun-

tain from the rift-wall and forms an impenetrable barrier

for forest dependent species.

A similar observation could be made with Kinyongia uth-

moelleri, an endemic of the highlands of Hanang and

Ngorongoro: the two populations show closest morpho-

logical and genetic relationships (Krause 2006) which as

well supports the above presumed close relationship be-

tween 7. hanangensis n. sp. and T. sternfeldi from

Ngorongoro.

Acknowledgements. The work of the senior author (PK) was

supported by a grant of the German Foreign Exchange Service

(DAAD). Herpetological researches in Tanzania were authorized

by Tanzania Commission for Science and Technology

(COSTECH). Many thanks to Mr. Joas Kahembe, Babati, M.

Nikundiwe and K. Howell (University of Dar es Salaam) who

gave support with a lot of practical advice and assisted obtain-

ing research permits.

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©ZFMK

A new chameleon of the Zrioceros bitaeniatus complex from Mt. Hanang, Tanzania 29

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Received: 04.12.2009

Accepted: 10.02.2010

Corresponding editor: F. Herder

©OZFMK

Bonn zoological Bulletin

Volume 57 Issue | pp. 31-54 Bonn, April 2010

Reptiles from southern Benin, West Africa,

with the description of a new Hemidactylus (Gekkonidae),

and a country-wide checklist

Klaus Ullenbruch!, Olaf Grell? & Wolfgang Bohme!

1Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany;

E-mails: Klaus-Ullenbruch@t-online.de; w.boehme.zfmk(@uni-bonn.de;

2Biologenbtro GGV, Stralsunder Weg 16, D-24161 Altenholz-Stift, Germany; E-mail: olaf.grell@t-online.de

Abstract. We report on a collection of reptiles made in southern Benin, mostly in the vicinity of Lama Forest, a relict

rain forest surrounded by savannah habitats within the the so-called Dahomey gap. 48 species (3 chelonian, 20 saurian

and 25 ophidian species were encountered) are listed with the respective voucher material and commented in respect to

taxonomic or ecological information. Five lizard species (Agama sylvanus, Hemidactylus ansorgei, H. lamaensis n. sp.,

Holaspis guentheri, Varanus ornatus) and two snake species (Natriciteres variegata, Amblyodipsas unicolur) are new

for the fauna of Benin, the second Hemidactylus species being even new to science. Some other species are reported from

Benin for only the second time.

This commented list of species collected from southern Benin is followed by a general country-wide checklist, which is

based on literature data and also some unpublished records from northern Benin, among them the remarkable rediscov-

ery of Agama gracilimembris, 9 decades after its original description from this country.

Key words. Reptilia: Chelonia, Crocodylia, Squamata; Benin, West Africa, new country records; Hemidactylus lamaen-

sis N. sp.; country-wide checklist

INTRODUCTION

Herpetologically, Benin is among the most insufficiently

known countries in Africa. Bordered by Togo in the west,

Burkina Faso and Niger in the north, and Nigeria in the

east, Benin is best-known by its old name Dahomey, be-

cause of the denomination of the huge savannah gap that

separates the Upper Guinean rain forests from those of the

Lower Guinean-Congolian forest block as Dahomey Gap.

Among the first authors who dealt especially with Benin

reptile material within the framwework of West African

herpetology is Chabanaud (1916, 1917). During his cu-

ratorship in the Museum of Natural History in Paris he

published on acquisitions from this former part of French

West Africa by the following collectors: Dr. G. Bouet,

Healt Inspector of French West Africa (“Dahomey”,

1910-1913), Dr. Brot (“Haut-Dahomey”, 1908), Mr. A.

Chevalier (“Dahomey”, 1909-1910), Mr. de Gironcourt

(“Haut-Dahomey”, 1910), Mr. Gruvel (“Dahomey”,

1913), Mr. Primot (“Dahomey”, 1914), and Mr. Waterlot

(“Dahomey”, 1914). The next important step in the explo-

ration of this country was the mission to Togo and

Dahomey carried out by A. Villiers in 1950 (see Villiers

1951, Loveridge 1952) which covered all important

biotopes of these two countries. In 1999, Sinsin &

Bergmans (1999) edited a book on rodents and snakes in

Benin under mostly agricultural aspects. The table on the

West African snake fauna including Benin provided for

this book by Meirte is not considered here because it marks

many species in this table as occurring in Benin without

any reference to the source on which the information is

based; on the other hand, several species are marked as

questionably or not at all occurring in Benin (e.g. Scaphio-

phis albopunctatus, Telescopus variegatus, Toxicodryas

blandingii, Psammophis cf. sibilans, P. phillipsi: Meirte

1999) although they have been documented by museum

voucher specimens in the earlier works cited above (see

also the country checklist at the end of this paper).

Where the savannah reaches the southern coast of West

Africa the Dahomey gap stretches from roughly between

Accra (Ghana) to Lagos (Nigeria), thus making Benin a

seemingly pure savannah country. However, some forest

32 Klaus Ullenbruch et al.

remnants such as Forét de Dan, Forét de Lama and Forét

de Lokoli are still present in the southern part of Benin.

The senior author of this paper (KU) had the opportuni-

ty to work on a herpetological survey of the most impor-

tant of these relict forest remnants, viz. Forét de Lama.

This work took place within the framework of a project

(“Biolama’’) of the University of Basel, Switzerland, guid-

ed by Prof. Dr. Peter Nagel (see Lachat et al. 2006). The

herpetological voucher material of this survey, which 1n-

cluded also specimens obtained from neighbouring for-

est and savannah areas (Ullenbruch 2003) is deposited in

the Zoologisches Forschungsmuseum Alexander Koenig

(ZFMK), Bonn, Germany. The amphibian part of this col-

lection contained some important first country records of

rain forest species (e.g. Ptychadena aequiplicata) and has

already been included in a recent paper by Rédel et al.

(2007). Amphibians collected by O. Grell in the Pendjari

National Park, northern Benin (Grell 2003) have been in-

corporated in the study by Nago et al. (2006). The reptil-

ian records of this last-mentioned survey, mostly docu-

mented by photographs, are included here only in the

country-wide checklist at the end of this paper. The fau-

nistic and ecological information on the reptiles of south-

ern Benin (mission K. Ullenbruch) is presented here in a

systematically arranged, commented species list. This list

further contains some voucher material collected and do-

nated to ZFMK by Dr. Werner Schréder, Gottingen, in

1984.

The commented regional species list which contains also

the description of one new species, 1s supplemented by a

general, country-wide checklist that tries to summarize all

literature data for Benin as a whole.

REPTILES FROM LAMA FOREST, SOUTHERN

BENIN, AND NEIGHBOURING AREAS

Lama Forest or Forét de Lama is situated within a depres-

sion bearing the same name, 80 km north of Benin’s cap-

ital Cotonou (Figure 1). It is bordered in the north by the

plateau of Abomey and in the south by the plateau of Al-

lada. Lama Forest comprises a total area of 16.250 ha of

“forét classée”, 9.750 ha of which belong to the Départe-

ment Atlantique (Sous-préfecture Toffo) and 6.500 ha be-

longing to the Départment Zou (Sous-préfecture Zog-

bodomé) (Emrich et al. 1999). The center of this protect-

ed area — the protectional status exists since 1947 — is

formed by the Noyeau Central (4.777 ha) which is com-

posed of primary forest (1.937 ha), degraded forest (1.388

ha) and of teak plantations and fallow grounds (together

1.452 ha) (Figure 2). The Noyeau Central is surrounded

by ca 9.000 ha of forestry plantations. In earlier times, the

primary forest covered the whole Lama depression which

stretches over the south Benin in a west-east direction. Un-

Bonn zoological Bulletin 57 (1): 31-54

Fig. 1.

lenbruch

View on Forét de Lama, South Benin. Photo: K. Ul-

til 1946, large areas of natural forest were already de-

stroyed, and only 11.000 ha remained. Since the 1960ies

reforestations are taking place which favoured, however,

mainly teak plantations. The natural vegetation has been

protected only from 1988 onwards (Emrich et al. 1999).

Mean annual precipitation is 1100 mm with two peaks (big

and small rainy seasons) in March to June and Septem-

ber to October respectively. The annual mean temperature

lies between 25—29°C. The eastern part of the Lama de-

pression is crossed — in north-south direction — by the

Oueémé River valley. Close to Lama Forest, there are on-

ly few smaller creeks. During the rainy seasons, howev-

er, large parts of the area are changed into overflooded

marshlands. Due to these repeated inondations and stag-

nant watres, there are only six dominant tree species in

Llama Forest, viz. Dialium guineense, Diospyros mespili-

formis, Albizia zygia, Afzelia africana, Khaja senegalen-

sis, and Anogeissus leiocarpus. The Noyeau Central con-

sists mainly of a mixture of relicts of a semideciduous for-

est which floristically approaches the “Forét de Samba”

Fig. 2.

Primary forest of Forét de Lama (Noyeau central) (left)

with bordering teak plantations (right) Photo: K. Ullenbruch

©ZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 33

Fig. 3.

Landscape round the village of Didja, southern

Benin. Photo: K. Ullenbruch

type (Mondjkannagni 1969) and of secondary forest in

various developmental stages. Within this dynamic sys-

tem, the South American fast-growing asteracean neophyte

Chromolaema odarata is commonly found along small

roads and paths and at the forest edges. In smaller plan-

tations, next to the larger ones with teak, trees such as

Gmelina arborea, fraké (Terminalia superba), samba

wood (Triplochiton skleroxylon) and, especially in the Tof-

fo sector, the fire-resistent Cassia siamea are cultivated

(Specht 2002).

The remaining collecting sites in southern Benin were:

— Didja village, situated ca. 25 km NW of Bohicon and

largely surrounded by an agriculturally formed land-

scape. The area north of Didja is characterised by bush

and tree savannas (Figure 3).

— Dan Forest or Forét de Dan is situated ca. 15 km NE of

Didja and is regularly visited by hunters from Didja.

Fig. 4.

Gallery forests at the Zou River, southern Benin. Pho-

to: K. Ullenbruch

Bonn zoological Bulletin 57 (1): 31-54

—Za-Kpota, 18 km NE of Bohicon, is likewise largely sur-

rounded by an agricultural area, replaced in the north-

west by large tree savannas. 7 km N of Za-Kpota, the

Zou River is running, being accompanied on its banks

by a small strip of dense and degraded gallery forest

which belongs to the hunting grounds of the villagers

(Figure 4).

— Lokoli Forest or Forét de Lokoli, a 500 ha large forest

on the Hounto River, ca. 100 km N of Cotonou. Numer-

ous sidearms of the Hounto River run through Lokoli

Forest so that it is penetrable only by boat (Figure 5).

For further details, in the context of an amphibian sur-

vey, see Rédel et al. (2007).

Flooded forest of Forét de Lokoli. Photo K. Ullen-

Fig. 5.

bruch

Finally, some reptiles were collected also in Cotonou and

Bohicon, on house walls and in gardens, and markets were

visited to inspect the reptile species offered. Most of the

reptiles were collected by UK by visual encounter and fo-

cal sampling. Moreover, specimens were brought by the

locals, particularly by two snake hunters from Didja and

Za-Kpota respectively who even received an alcohol-filled

bucket in order to keep interesting snake finds for some

days. The Swiss zoologists of the Biolama project provid-

ed us with few, but very important voucher specimens (e.g.

Hemidactylus sp.n., Holaspis guentheri) taken in trunk

eclectortraps used for the entomological survey.

The survey of the reptile fauna of Lama Forest, South

Benin, and the neighbouring localities mentioned above

revealed the existence of 48 species (2 turtle, 1 tortoise,

20 lizard, and 25 snake species) which are listed and dis-

cussed in the following.

OZFMK

34 Klaus Ullenbruch et al.

ANNOTATED LIST OF SOUTHERN BENINIAN

REPTILES

Testudinidae

Kinixys belliana nogueyi (Lataste, 1886)

Material examined. photographic voucher only (Figure 6).

Remarks. This specimen was brought to KU by a local

hunter. It was taken near Bohicon and was released after

photographic documentation.

Fig. 6.

Kinixys belliana nogueyi from near Bohicon, south-

ern Benin. Photo: K. Ullenbruch

Pelomedusidae

Pelomedusa subrufa (Lacépede, 1788)

Material examined. photographic voucher only (Figure 7).

Remarks. Specimen taken by local hunter near Bohicon,

photographed and afterwards released.

Pelusios castaneus (Schweigger, 1812)

Material examined. photographic voucher only (Figure 8).

Remarks. Specimen taken by local hunter near Bohicon,

photographed and afterwards released.

Agamidae

Agama aff. agama (Linnaeus, 1758)

Material examined. ZFMK 42019-024 (3 males, 3 fe-

males), Godomey near Cotonou, coll. by W. Schréder, Au-

Bonn zoological Bulletin 57 (1): 31-54

Fig. 7. Pelomedusa subrufa from near Bohicon, southern

Benin. Photo: K. Ullenbruch

gust/September 1984; ZFMK 77061-062, Bohicon,

ZFMK 77063-064, Cotonou, all coll. by K. Ullenbruch,

May/June 2002.

Remarks. This common and widespread species was ob-

served in all habitats including human settlements where

vertical structures such as tree trunks and/or house walls

were preferred. It was, however, never seen in the closed

forest.

Fig. 8.

Pelusios castaneus from near Bohicon, southern

Benin. Photo: K. Ullenbruch

©ZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 35

Agama sylvanus Macdonald, 1981

Material examined. ZFMK 77065, Forét de Lama (Noyeau

Central), coll. by K. Ullenbruch, 6 June 2002.

Remarks. The single juvenile specimen was obtained in

an arthropod trap fixed at the trunk of a dead tree (trunk

eclector) in dense primary forest. A second juvenile was

observed at about 5 m height and escaped upwards. Dur-

ing the three months of the survey, no further specimen

of this agama was seen which argues for either a secre-

tive lifestyle or a general rarity, at least in this season of

the year.

The morphological characters of ZFMK 77065 lie with-

in the range of variation of A. sylvanus which was de-

scribed by Macdonald (1981) from Ghana. Joger (1990)

recorded the species also from Cameroon (as A. paraga-

ma sylvanus). The disjunction between these two coun-

tries is partially bridged by our specimen which represents

the first country record for Benin.

Chamaeleonidae

Chamaeleo gracilis Hallowell, 1842

Material examined. ZFMK 77056-057, environs of Did-

ja, coll. by a native hunter, 10 and 30 May 2002 resp.

Remarks. The two males were obtained from a native

hunter on the market of Abomey who certified their ori-

gin as Close to Didja. C. gracilis is widespread in the Su-

dan and Guinea savannah belt south of the Sahara. The

rudimentary occipital flaps distinguish it from the follow-

ing, only recently described species.

Chamaeleo necasi Ullenbruch, Krause & Bohme, 2007

Material examined. ZFMK 77058, male, Za-Kpota,

ZFMK 77059-060, 2 females, Houégbo, all coll. by K. Ul-

lenbruch, May/June 2002.

Remarks. The three specimens form part of the type se-

ries of this species which is otherwise known only from

Togo (Ullenbruch et al. 2007). The Togolese localities

(Kpalimé, Missaboué) as well as the two Benin localities

are situated close to relict forest patches within the arid

Dahomey gap. Za-Kpota is an agriculturally shaped site

not far from the river Zou which has some degraded

gallery forest vegetation, while Houégbo is not far from

Lama Forest.

Bonn zoological Bulletin 57 (1): 31-54

C. necasi belongs to the C. dilepis group and differs from

the south Nigerian populations of the closely related C.

quilensis mainly by hemipenial characters (see Ullenbruch

et al. 2007).

Chamaeleo senegalensis Daudin, 1802

Material examined. ZFMK 42025 (male), Bohicon, coll.

by W. Schréder, August/September 1984; ZFMK 77052-

055, environs of Didja, coll. by a native hunter, 10 May

2002.

Remarks. The four specimens were obtained from a na-

tive hunter on the market of Abomey who certified their

origin as close to Didja. C. senegalensis — a species of the

C. dilepis group without any traces of occipital flaps — was

found on all markets in southern Benin and proved to be

much more common than C. gracilis.

Eublepharidae

Hemitheconyx caudicinctus (Dumeéril, 1851)

Material examined. ZFMK 77066, near Didja, ZFMK

77067, Za-Kpota, both coll. by native collectors, 27 May

and 18 June 2002, respectively.

Remarks. A West African species distributed from Sene-

gal to Cameroon which has its southernmost localities in

Benin. In Benin, it is termed “snake with two heads” by

the locals and thought to be very dangerous.

Gekkonidae

Hemidactylus ansorgii Boulenger, 1901

Material examined. ZFMK 77073-075, Lama Forest

(Noyeau Central), coll. by K. Ullenbruch, 18. (77073) and

28. April 2004.

Remarks. This slender, narrow-headed forest gecko with

a long, cylindrical tail has only rarely been encountered

and is known with certainty only from Liberia, Ivory

Coast, Nigeria, and Cameroun (Henle & Bohme 2003).

Bauer et al. (2006) expected H. ansorgii to occur in Benin,

due to its wide West African distribution but stressed that

there are no confirmed records from this country. Thus,

our specimens represent the first record for Benin (Fig-

ure 9). The geckos were found in the late morning

(11.30h—12.30h) on tree trunks in dense primary forest at

about 1.7 m height.

©OZFMK

36 Klaus Ullenbruch et al.

Fig. 9.

Hemidactylus ansorgei from Lama Forest (Noyeau

central), first country record. Photo: K. Ullenbruch

Hemidactylus angulatus Hallowell, 1852

Material examined. ZFMK 42006-009, Godomey near

Cotonou, coll. by W. Schréder, August/September 1984;

ZFMK 77070, Bohicon, coll. by K. Ullenbruch, 17 April

2002; ZFMK 82813-814, Cotonou, coll. by Thibault La-

chet, 22/23 April 2004.

Remarks. Thys van den Audenaerde (1967) distinguished

between Hallowell’s forest-dwelling species Hemidacty-

lus angulatus and the savannah form H. brookii guineen-

sis Peters, 1868. Since the African taxa of the former con-

cept of an Afro-Indian species H. brookii Gray, 1845 are

not at all close relatives of the true H. brooki from India

(Carranza & Arnold 2006), it is still unclear whether the

name guineensis should be treated as a synonym or a sub-

species of angulatus, or possibly even a species. Accord-

ing to Bauer & Giinther (1991), it is a synonym of angu-

latus (understood by these authors still as a subspecies of

H. brookii).

Examination of the lectotype (Figure 10) of H. guineen-

sis Peters, 1868 [described from “Ada Foah, Guinea” (=

Adafoa, Ghana; not Adafer, Mauritania, as suggested by

Loveridge 1947 and Wermuth 1965!) and compared by Pe-

ters (1868) only with H. verruculatus (= H. turcicus)| re-

vealed that despite its provenance from coastal West

Africa, it resembles the savannah form rather than the for-

est form which would strengthen a partial sympatry with

the forest-dwelling H. angulatus. However, the distin-

guishing key character between both forms, i.e. the num-

ber of granules bordering the nostril above rostral and first

supralabial (Thys van den Audenaerde 1967), has been dis-

proved already by B6hme (1978), and other scalation dif-

ferences also seem to be insufficient for a clear separa-

tion of both forms. Habitus differences could possibly be

found in colour pattern, which often seems to consist of

Bonn zoological Bulletin 57 (1): 31-54

Fig. 10. Hemidactylus angulatus / guineensis complex: lecto-

type of Hemidactylus guineensis Peters, 1868. Photo: P. Wagn-

paired dorsal bars interrupted on the dorsal midline in the

savannah populations, combined with more distinctly

white-coloured single tubercles in the flank region. In the

forest-dwelling populations, it consists of smaller, less

symmetrically arranged darker elements, with less obvi-

ous white flank tubercles in the forest populations.

Steindachner (1870) described H. affinis from Gorée and

Dagana, Senegal, and diagnosed it also only against H.

verruculatus (= H. turcicus). He stressed the similarity of

these two species, particularly concerning the size of their

strongly keeled tubercles. The type series of H. affinis

closely resembles the savannah-dwelling H. guineensis.

In his description of H. stellatus, Boulenger (1885) com-

pared it with H. brookii and H. gleadovii and stressed the

presence of pure-white tubercles intermixed with a ma-

jority of dark brown ones which inspired him to the name

stellatus (= starred). These white tubercles are indeed more

obvious in the savannah-dwelling form (Figure 11) and

Fig. 11. Hemidactylus angulatus / guineensis complex: lecto-

type of Hemidactylus stellatus Boulenger, 1885. Photo: C. Mc-

Carthy

©OZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 37

Fig. 12. Hemidactylus angulatus (s.str.) from Godomey near Cotonou (left), H. mabouia from Cotonou (center) and H. aff. mabouia,

also from Cotonou (right). Photo: P. Wagner

brought also Chabanaud (1917 a) to the conclusion that

there are two different species in this group of geckos, viz.

“H. brookii” (certainly meaning angulatus) and “H. stel-

latus” (= H. guineensis), at Agouagon, Benin, even in

sympatry.

Preliminary genetic support for two species within a H.

angulatus/guineensis complex may be deduced from the

tree in Carranza & Arnold (2006) where a Kenyan Rift

Valley sample is closer to dry savannah samples from

Mauritania and Mali (guineensis) than to a forest sample

from Bioko (angulatus s.str.). Moreover, as already point-

ed out by Thys van den Audenaerde (1967), the forest

species H. angulatus does not occur in Central and East

Africa.

Bonn zoological Bulletin 57 (1): 31-54

Bauer et al. (2006) who provided locality data for many

specimens of their H. angulatus (sensu lato) from all over

Benin, did not distinguish between the two forms thus im-

plying morphological uniformity, except their variable

colour pattern. They stressed that the sympatric H.

mabouia could be easily mixed up with H. angulatus what

we also found to be true (but only for the forest form: an-

gulatus s.str.) particularly when the diagnostic (see the key

in Bauer et al. 2006) characters of tubercles on head and

dorsum are concerned. Here, again the colour pattern of

the dark phases of H. angulatus and H. mabouia may

prove to be more reliable than the scalation (Figure 12).

Concluding it becomes clear that Hemidactylus angula-

tus S.]. 1s in urgent need of a taxonomic revision through-

out its range.

©ZFMK

38 Klaus Ullenbruch et al.

ZFMK 77070 was captured on a house wall on 19.30 h,

i.e. 30 min. after sunset.

Hemidactylus fasciatus Gray, 1842

Material examined. ZFMK 82808, Forét de Lama (Noyeau

Central), coll. by Lamidi Konetche, April 2004.

Remarks. This species was first documented for Benin by

Bauer et al. (2006) who recorded specimens from the

vicinity of Kétou. Our record, the second for Benin, is

from dense primary rain forest.

Hemidactylus mabouia Moreau de Jonnés, 1818

Material examined. ZFMK 77068-09, Cotonou, coll. by

K. Ullenbruch, Datum; 82809-812, Cotonou (Maison des

Stagnaires), coll. by Thibault Lachet, 22/23 April 2004.

Remarks. Not found by W. Schréder in 1984 and docu-

mented for the first time in Benin by Bauer et al. (2006).

This species was considered to be rare in Nigeria and be-

lieved to be introduced in port cities by Dunger (1968).

Also, R6dder & B6hme (2008) regarded it as endemic on-

ly in Central and East Africa, while it is an invasive species

—next to South America, the Caribbean region and Flori-

da — also in West Africa.

ZFMK 77068-09 differ from typical H. mabouia by lack-

ing the very characteristic dark pattern of oblique curved

cross stripes (Figure 12). Similar phenotypes have been

found in western Guinea (Bohme et al., unpubl. data). As

no scalation differences could be detected, we regard these

specimens currently as a colour morph of H. mabouia,

pending new material for genetic analyses.

The two specimens were collected by UK at noon time

(12.00h) from house walls during a heavy thunderstorm

rain. The geckos were observed to hunt for insects which

were beaten to (or at least close to) the ground by the rain-

water. The reduced daylight — a consequence of the dense

clouds — might have helped the geckos to become active

and to forage during midday. The masses of insects con-

centrating under these conditions within the lower two me-

ters above the ground were also used and exploited by

Agama aff. agama and Mochlus guineensis.

Hemidactylus lamaensis new species

Holotype. ZFMK 77072, adult female, Forét de Lama

(teak plantation), coll. from a pitfall by Klaus Ullenbruch,

16 April 2002.

Bonn zoological Bulletin 57 (1): 31-54

Diagnosis. A small Hemidactylus with a SVL of about 5.0

cm. Characteristic is the dorsal scalation which consists

of numerous irregular rows of large, convex, keeled tu-

bercles which are nearly juxtaposed on the dorsal hind-

part and on the tail root, separated mostly by only one row

of surrounding, rather coarse granular scales. Also the tem-

ple is covered by large, juxtaposed tubercles with only few

or even without granules between them. Moreover, the

subdigital lamellae of the first digit and toe are very low

in number, the terminal unpaired lamella of digit I (manus)

being the only adhesive element while digit I (toe) has two

scansors: one divided proximal and one undivided termi-

nal one.

Description of the holotype (Figure 13). Habitus stout,

head small with an elongated snout. Head & body length

47.7 mm, length of (regenerated) tail 34 mm. Snout elon-

gate: distance from anterior margin of tympanum to hind-

margin of eye 4 mm, from anterior margin of eye to tip

of snout 5.5 mm; eye diameter 2.5 mm. Rostral broad, with

a deep notch on its anterior midline which continues as a

vertical suture but does not separate the rostral complete-

ly. Nostril bordered by rostral, a Ist labial, three postnasal

scales, and a supranasal which is separated from its fel-

low by one single median scale. Scales on snout small but

polygonal rather than granular, keeled, passing backwards

into a tuberculate scalation of the frontal and parietal re-

gion which becomes intermixed with granules from the

occipital region towards the neck and dorsum. Temple

covered with conical tubercles with only few intermixed

granules between them. Mouthline straight and not uplift-

ed in its hindpart. Supralabials (left/right) 10/10, sublabi-

als (left/right 8/8). Mental triangular, followed by two pairs

of postmentals, the first of them broadly meeting at the

midline behind mental. Gulars small, smooth, passing pos-

teriorly gradually into the semicircle-shaped, subimbricate

and likewise smooth chest and ventral scales. The coni-

cal and weakly keeled dorsal tubercles are arranged in

16-18 irregular longitudinal rows, circumdated by rather

coarse granular scales which separate the tubercles by 1-3

rows, while on sacrum and tail root the tubercles are even

nearly juxtaposed with only few scattered granules be-

tween them. Limbs covered above with coarse granular

scales intermixed with conical, weakly keeled tubercles.

Palmar scales smooth. All fingers and toes with distinct

claws, and with divided adhesive lamellae (scansors) be-

neath, except the terminal one which is simple. Also the

basal subdigital scales are undivided and are added to the

number of subdigital lamellae when they are twice as large

than the palmar scales. The values are: 1-5-6-7-6 (left

manus), |-5-6-7-5 (right manus), 2-6-7-8-7 (left pes), and

2-6-6-7-6 (right pes). The regenerated tail is dorsally cov-

ered with very regular subimbricate smooth scales which

gradually pass into several longitudinal rows of oblique-

ly enlarged, imbricate subcaudal plates.

©ZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 39

Fig. 13. Hemidactylus lamaensis n. sp., holotype from Lama Forest (Noyeau central) in dorsal (a), ventral (b) and lateral (c) view.

Photo: P. Wagner

Bonn zoological Bulletin 57 (1): 31-54 OZFMK

40 Klaus Ullenbruch et al.

Colour in preservative. Upper side rather uniformly light

brownish, with a darker brown head, and one indistinct

darker crossband on the neck and on the shoulder respec-

tively. Underside uniform whitish-yellowish.

Etymology. The species name refers to Lama Forest, so

far the only known locality of the new Hemidactylus. It

is intended to stress the importance of this relict forest

within the Dahomey gap also in conservational aspect as

it may harbour still more unknown rain forest species.

Distribution and natural history. Only known from the

type locality. The single female was taken from a pitfall

on the forest floor in a young teak plantation bordering

the primary forest of the Noyeau Central. This does not

argue against an arboricolous lifestyle, because also tree-

dwellers like H. fasciatus were observed to climb down

from their shelter in big fig trees and to forage on the

ground some hours after sunset (BGhme 1975).

Comparisons. This unique single specimen cannot be as-

signed to any West African species (see Bauer et al. 2006).

Its snout is elongated but not to the extent observed in H.

ansorgei. Its most obvious character is the possession of

densely arranged, large, keeled tubercles which are less

numerous as compared to those of the UH.

angulatus/guineensis complex. On the hindpart of the dor-

sum and the tail root, they are even nearly juxtaposed. Al-

so in contrast to the situation found in the former com-

plex is the dense coverage with tubercular scales of the

temporal region. Moreover, it has the lowest number of

subdigital lamellae of all Beninian and West African Hemi-

dactylus.

Assuming that the original tail in this species has also se-

ries of tubercles (not present in the regenerated tail of the

unique holotype), the key given by Bauer et al. (2006) for

the geckos of Benin can be completed as follows:

13a. Dorsal tubercles prominent, keeled ............... 14

13b. Dorsal tubercles not, or only weakly keeled, sepa-

rated by one another by a distance greater than by tuber-

cle width; dorsum of head atuberculate or with few small

tubercles; color variable, the typical pattern of a series of

dark curved crossbands distinct in the dark phase only

BHO tpn Neath etna wen ce aac re: Hemidactylus mabouia

14a. Dorsal tubercles separated by one another by approx-

imately the width of one tubercle; tubercles usually promi-

nent on dorsum of head; temple granular

By eisai Hemidactylus angulatus/guineensis complex

14b. Dorsal tubercles separated by one another by less than

a tubercle width, on sacrum and tail root even abutting;

temple covered with conical tubercles

a Snaislatee Danis Set arers AGES Hemidactylus lamaensis sp. nov.

Bonn zoological Bulletin 57 (1): 31-54

Scincidae

Panaspis togoensis (Werner, 1902)

Material examined. ZFMK 77079-080, Forét de Lama,

coll. by K. Ullenbruch, 24 and 28 April 2002.

Remarks. Both voucher specimens were collected in a

transition zone of primary forest and a Chromolaema

clearing, after heavy rainfalls around noon time. Consid-

ered to be ombrophilous by Perret (1973), activity after

rains was also observed by Rédel et al. (1997). Accord-

ing to Fuhn (1972) and Schmitz et al. (2005), Lygosoma

(Riopa) dahomeyense Chabanaud 1917 is a synonym of

P. togoensis.

Mochlus guineensis (Peters, 1879)

Material examined. ZFMK 77076, Forét de Lama; ZFMK

77077-078, Cotonou, all coll. by K. Ullenbruch, 16 and

13 April 2002 respectively.

Remarks. The specimen from Lama Forest was taken in

a pitfall trap placed in an old teak plantation close to the

primary forest. The two specimens from Cotonou were

caught in the garden of a house during heavy rainfall when

they were foraging for prey negatively influenced by the

rains (see above, under Hemidactylus mabouia).

Remarkably, the specimen from Lama Forest differs in

some characters from the two Cotonou specimens. It is

more delicately built, having thinner limbs and a longer

and slimmer tail. Moreover, the eyes seem to be smaller,

the overall colouration is distinctly darker, and the sub-

caudal scales are spotted with black, while the Cotonou

specimens have the underside of their tails purely white.

More material is necessary to evaluate these differences

(Figure 14).

Trachylepis affinis (Gray, 1838)

Material examined. ZFMK 42004-005, Godomey near

Cotonou, coll. by W. Schréder, August/September 1984;

ZFMK 77081-090, Forét de Lama, coll. by K. Ullenbruch,

16 April to 1 May 2002.

Remarks. The two specimens from Godomey come from

a farmland/garden environment, the Lama Forest individ-

uals, however, were all collected within dense primary for-

est (Noyeau Central), on clearings grown with Chromo-

laema, and in the neighbouring teak plantations. The

lizards were found either in leaf litter and on the grassy

floor, or on tree trunks. When disturbed they tried almost

©OZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 4]

Fig. 14. Mochlus guineensis, from Cotonou (above) and Lama

Forest (Noyeau central) (below). Photo: P. Wagner

invariably to escape upwards. They were mainly active be-

tween 11.00h and 15.00h, at air temperatures of 31—32°C.

Half of the Lama Forest sample has a distinctly striped

colour pattern with a conspicuous, white sublabial stripe

running along the lower flank towards the tail, while the

other half lacks this stripe.

Trachylepis perroteti (Duméril & Bibron, 1839)

Material examined. ZFMK 41998-999 (2 males),

Godomey near Cotonou, coll. by W. Schréder,

August/Septeber 1984.

Remarks. According to Mausfeld-Lafdhyia et al. (2004)

and Chirio & LeBreton (2007), 7: perroteti consists of at

least two distinct species which require further study (see

also Stoll 2008). But because the true 7. perroteti (s.str.)

is the more southernly distributed from of the two cryp-

tic siblings in Cameroon, the two males from near Coto-

nou are most likely belonging to this latter species.

Whether this is also true for the North Beninian popula-

tions (see the country checklist below) remains to be stud-

ied.

Lacertidae

Holaspis guentheri Gray, 1863

Material examined. ZFMK 77071, Forét de Lama (teak

plantation), coll. by K. Ullenbruch, 26 June 2002.

Remarks. The only voucher specimen was obtained from

an arthropod trap (trunk eclector) fixed on a dead teak tree

(Tectona grandis) within an old teak plantation bordering

the primary forest of Forét de Lama. This little lacertid

Bonn zoological Bulletin 57 (1): 31-54

with the ability of gliding flight has not been recorded so

far from Benin, thus representing the first country record.

Varanidae

Varanus exanthematicus (Bosc, 1792)

Material examined. ZFMK 77004-005, near Didja, coll.

by local hunter, 1 May 2002.

Remarks. Both specimens were obtained from a local

hunter at Abomey who certified their origin as being close

to Didja.

Varanus niloticus (Linnaeus, 1766)

Material examined. ZFMK 42200, Godomey near Coto-

nou, coll. by W. Schréder, August/September 1984; ZFMK

77001, juv., Ouémé River near Togbota, coll. by local

hunter, 15 April 2002.

Remarks. ZFMK 77001 was found under a heap of dry

wood on the bank of a small tributary of the Ouémé Riv-

er, close to the settlement of Togbota.

Varanus ornatus (Daudin, 1803)

Material examined. ZFMK 77002-003, near Didja, coll.

by native hunter, 10 and 30 May 2002 respectively.

Remarks. Both specimens were obtained from local

hunters at Abomey who had collected them in the vicin-

ity of Didja. One of them originated from Lakpo, 8 km

NE of Didja, which is in immediate vicinity of a further

relict forest patch, viz. Foret de Dan. A big, adult speci-

men was seen in hectic flight in Lama Forest and could

not be caught.

On markets, V. ornatus was much less commonly seen

than V. niloticus which reflects also the scarcity of relict

forest patches in the Dahomey gap which serve as refu-

gia for this forest species. For Benin, our specimens rep-

resent the first country record (B6hme & Ziegler 1997).

Pythonidae

Python regius (Shaw, 1802)

Material examined. ZFMK 41993-995, Godomey near

Cotonou, coll. by W. Schréder, August/September 1984;

Photographs from Za-Kpota and Forét de Lama.

©OZFMK

42 Klaus Ullenbruch et al.

Remarks. Of ZFMK 41995, only head and neck are pre-

served. This species was seen in great numbers on all mar-

kets visited. These great numbers certainly pose a serious

conservation problem which could be buffered by farm-

ing royal pythons for the international pet trade. Howev-

er, such farms will be very difficult to be controlled as far

as wild-caught specimens are concerned.

Python sebae (Gmelin, 1788)

Material examined. None.

Remarks. Rock pythons could be seen only twice on mar-

kets by UK during his survey. He was told by the people

that they fear the large snakes and normally kill them

whenever possible. This species is subject to superstitious

beliefs by the local people. For instance, a scientific as-

sistant of the BIOLAMA project even told that rock

pythons are capable to cast out a net out of their mouth

to catch humans and to eat them subsequently.

Colubridae (sensu lato)

Afronatrix anoscopus (Cope, 1861)

Material examined. ZFMK 77006 (juvenile), Forét de

Lokoli, coll. by K. Ullemnbruch, 5 May 2002.

Remarks. The existence of this aquatic species in Benin

was considered as likely by Chippaux (2001). Our spec-

imen (Figure 15) was the first documented record from

Benin until very recently Chirio & Ineich (2009) published

a find from Atakora Province which makes our specimen

the 2nd published record. It is, however, still interesting

to note that both new records of this species are very dis-

tant from each other, ranging from a relict, swampy rain

forest to dry Sudan savannah.

Crotaphopeltis hotamboeia (Laurenti, 1768)

Material examined. ZFMK 42001-002 (juveniles),

Godomey near Cotonou, coll. by W. Schréder,

August/September 1984.

Remarks. None.

Dasypeltis gansi Trape & Mané, 2006

Material examined. ZFMK 77014, Za-Kpota, coll. by na-

tive hunter, may 2002.

Bonn zoological Bulletin 57 (1): 31-54

Fig. 15. Afronatrix anoscopus, juvenile, from Lokoli Forest,

first country record. Photo: K. Ullenbruch

Remarks. Trape & Mané (2006 a) partitioned the West

African egg-eater snakes formerly referred to as D. scabra

into 3 different species, one of them with a likewise new

subspecies: D. confusa, D. gansi (with D. g. gansi and D.

g. latericia), and D. sahelensis. According to the original

description and to the key and colour plates of Trape &

Mané (2006 b) the specimen from Za-Kpota clearly rep-

resents the nominotypic subspecies of D. gansi. It is the

second voucher from Benin, the first (Trape & Mané 2006

a) Benin record being from Lanta (where it is, according

to the same authors, sympatric with D. confusa). The

record of Chabanaud (1916) of “D. scabra palmarum”

refers very likely also to D. g. gansi which is the only of

the newly described taxa which tends to be patternless.

Dispholidus typus (Smith, 1829)

Material examined. ZFMK 77029, between Bohicon and

Didja, coll. by K. Ullenbruch, May 2002.

Remarks. Found dead on the road between the two above-

mentioned places.

Lamprophis fuliginosus (Bote, 1827)

Material examined. ZFMK 77011, Didja; 77015-017, Za-

Kpota; 77041, Forét de Lama; 77051, Forét de Dan, all

coll. by K. Ullenbruch, 14-18 June 2002.

Remarks. ZFMK 77041 was found dead on a path with-

in Lama Forest, just between primary forest and the neigh-

bouring teak plantation. Although still rather fresh, its head

was already badly damaged by ants.

©ZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 43

Lycophidion semicinctum Duméril, Bibron & Duméril,

1854

Material examined. 77007, near Didja, coll. by native

hunter, May 2002; ZFMK 77050, near Bohicon, coll. by

K. Ullenbruch, May 2002.

Remarks. The Bohicon specimen was found dead on a

road.

Meizodon regularis Fischer, 1856

Material examined. ZFMK 42003, Godomey near Coto-

nou, coll. By W. Schréder, August/September 1984.

Remarks. Known from, among else, Ghana (where the

type locality is situated), and Togo west of Benin, as well

as from Cameroon further east (Roux Estéve 1969), this

species was to be expected in Benin but was obviously

only indicated for Benin by Chippaux (2001). ZFMK

42003 represents the second country record and first pub-

lished voucher specimen for Benin.

Natriciteres olivacea (Peters, 1854)

Material examined. ZFMK 77039, Forét de Lama, coll.

by K. Ullenbruch, May 2002.

Remarks. Found on a cloudy day at 11.20h on the forest

floor. This underlines that this species is not confined to

savannah habitats as claimed by Broadley (1983) but that

at least in West Africa also forests are inhabited (see Vil-

liers 1975, Chippaux 2001: distribution map).

Natriciteres variegata (Peters, 1861)

Material examined. ZFMK 77042-046, Foret de Lama,

coll. by K. Ullenbruch, May 2002.

Remarks. All five individuals were found on the forest

floor. The species proved to be rather common but was

never seen outside the forest. Also Dunger (1972a),

Broadley (1983) and Chippaux (2001) regard N. variega-

ta as a forest species. The only earlier record of it was Cha-

banaud’s report from “Dahomey”, thus our small series

represents the first documented locality record for Benin.

Philothamnus irregularis (Leach, 1819)

Material examined. ZFMK 77025-026, Za-Kpota, coll. by

native snake hunter, May 2002; ZFMK 77040, Foret de

Lama: Koto village, coll. K. Ullenbruch, May 2002.

Bonn zoological Bulletin 57 (1): 31-54

Remarks. The last mentioned specimen was killed by a

local farmer in the presence of KU on a sun-exposed

grassy area.

Philothamnus semivariegatus (Smith, 1847)

Material examined. ZFMK 77034-035, Bohicon, coll. by

K. Ullenbruch, May 2002.

Remarks. Both snakes were encountered on the compound

of a mill. One of them was found on the floor of a house

belonging to that compound. Encounters in human habi-

tations and even within buildings were also reported from

Nigeria by Dunger (1973a).

Psammophis elegans (Shaw, 1802)

Material examined. One specimen, uncatalogued, Coto-

nou (stored in the private collection of M. Serge Attignon,

Cotonou), coll. by a S. Attignon, 11 May 2002.

Remarks. The only available specimen of this snake was

found in front of a house in the outskirts of Cotonou.

Psammophis phillipsi (Hallowell, 1844)

Material examined. ZFMK 41996, Godomey near Coto-

nou, coll. by W. Schréder, August/September 1984.

Remarks. The large adult, dark-brown unicoloured female

is allocated to Hallowell’s species because the mid-region

of its ventral plates has a broad median zone shaded with

bluish-grey colour. We think that this is a species of forest-

ed and moist savannahs which is unicoloured above al-

ready as a juvenile has and lacks the typical striped juve-

nile dress of the following form (Bohme et al. 1996).

Psammophis cf. phillipsi (Hallowell, 1844)

Material examined. ZFMK 77024 & 77028, Za-Kpota,

coll. by a native snake hunter, May 2002; ZFMK 77047,

Abomey, coll. by K. Ullenbruch, May 2002.

Remarks. The three specimens belong to the big-growing

form of the Psammophis sibilans species complex which

is dorsally striped as a juvenile and unicoloured above and

below as a large, old adult. It inhabits the dry Sudanian

and Sahelian savannahs and is mostly referred to as P. cf.

phillipsi (e.g. Bbhme 1978, Bohme et al. 1996) or, recent-

ly, again as true P. sibilans (Trape & Mané 2006 b). Al-

©OZFMK

44 Klaus Ullenbruch et al.

Fig. 16. Psammophis sudanensis from Bohicon, southern

Benin. Photo: K. Ullenbruch

so earlier records of P. sibilans from West Africa (e.g. Cha-

banaud 1916) refer to this form which, in our opinion, is

not conspecific with the true P. sibilans from Egypt. Many

not yet fully grown but nonetheless adult (mature) spec-

imens may retain more or less distinct reddish dorsolat-

eral stripes along the body. The belly is light whitish to

yellowish and patternless (no dark hairlines along the ven-

tral plates), but supralabials, sublabials and gulars show

some dark pigmented spots (see BOhme et al. 1996). The

anal shield is entire. Its taxonomic status and nomenclat-

ural status needs still to be assessed.

Psammophis sudanensis Werner, 1919

Material examined. ZFMK 77012, Didja, coll. native

hunter; ZFMK 77018-019; ZFMK 77036-038, Bohicon,

and ZFMK 77047, Abomey, all coll. by K. Ulenbruch,

May 2002.

Remarks. The specimens represent various age classes. All

have distinct hairlines along the outer parts of the ventral

shields, however, these hairlines are brownish in younger

and darker in the older individuals. The zone between

these hairlines is differently coloured than the two outer,

ventrilateral zones, the latter being white while the inner

is yellowish-whitish. This ventral pattern is therefore of

the “subtaeniatus type” and agrees with the type speci-

men of sudanensis Werner, 1919 (NMW 1986 from

Kadugli, Kordofan) which actually was described as P.

subtaeniatus sudanensis (Werner 1919, Bohme 1987). In

all specimens, the anal shield is divided.

ZFMK voucher specimens from e.g. northern Senegal

(Dakar, Fété Olé), Algeria (Tamanrasset), North Cameroon

(Mokolo) (see B6hme 1986) and Chad (ZFMK 74520) are

Bonn zoological Bulletin 57 (1): 31-54

Fig. 17. The same specimen from below. Photo: K. Ullenbruch

very close to typical P. sudanensis but differ in having e1-

ther faint, light brown hairlines (instead of blackish ones)

or no hairlines at all; their undersides are in both cases of

a uniform, bright porcellanic white. They correspond to

the type specimen of P. leucogaster Spawls, 1983

(BMNH 1980: 261, from Wa, northern Ghana, see Spawls

1983), and further sampling must show whether both

forms have zones of intergradation. It seems that if there

is a distinction between typical sudanensis and leucogaster

in West Africa, this is a question of a north-south rather

than a west-east disjunction.

ZFMK 77036-038 were found on the compound of a mill

at Bohicon. One of them was lying in the afternoon on the

terrace of a house, the other two were killed during grass

mowing (Figs 16 and 17).

Rhamphiophis oxyrhynchus (Reinhardt, 1843)

Material examined. ZFMK 41997, Godomey near Coto-

nou, coll. by W. Schréder, August/September 1984; ZFMK

77020-023, Za-Kpota, coll. by a native hunter, May 2002.

Remarks. None.

Scaphiophis albopunctatus Peters, 1870

Material examined. ZFMK 77009-010 (juvenile and

adult), Didja, collected by a local snake hunter, May 2002.

Remarks. The juvenile specimen contained 3 newborn

mice which could not be identified to the familial or gener-

ic level.

©OZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 45

Atractaspididae

Amblvodipsas unicolor (Reinhardt, 1843)

Material examined. ZFMK 77027, Za-Kpota, coll. by na-

tive hunter, May 2002.

Remarks. When fixing this snake, some white nematodes

appeared between the scales and tried to leave the speci-

men. This was not observed in any other snake during

preparation.

Elapidae

Dendroaspis viridis (Hallowell, 1844)

Material examined. ZFMK 77049, Bohicon (market), coll.

by K. Ullenbruch, May 2002.

Remarks. Since this snake was found on a market, no fur-

ther information on its precise provenance is available.

Elapsoidea semiannulata Bocage, 1882

ZFMK 77013, near Didja, coll. by a native snake hunter,

May 2002.

Remarks. Represents the West African subspecies E. s.

moebiusi Werner,1897 (Trape & Mané 2006).

Naja nigricollis Reinhardt, 1843

Material examined. ZFMK 77032, Foret de Lama; ZFMK

77033, Bohicon, both coll. by K. Ullenbruch, May 2002.

Remarks. One of the two juvenile specimens was found

dead on a road at the margin of Lama Forest, the other

was killed during grass mowing in the mill compound of

Bohicon.

Viperidae

Bitis arietans (Merrem, 1820)

Material examined. ZFMK 77048, environs of Abomey,

coll. by native hunter, May 2002.

Remarks. B. arietans is very commonly offered on mar-

kets. Usually, the head is cut off, dried and used for

Voodoo ceremonies while the rest of the animal is used

as food.

Bonn zoological Bulletin 57 (1): 31-54

Causus maculatus (Hallowell, 1842)

Material examined. ZFMK 77030, between Bohicon and

Didja, ZFMK 77031, 9 km east of Abomey, both coll. by

K. Ullenbruch, May 2002.

Remarks. The first specimen of this ubiquitous species was

found dead on the asphalt road between Bohicon and Did-

ja, the second one was killed by locals next to the street

to Abomey in a crop field.

Echis ocellatus Stemmler, 1970

Material examined. ZFMK 77008, near Didja, coll. by lo-

cal snake hunter, May 2002.

Remarks. None.

CONCLUSIONS

Of the 48 species recorded above from several localities

in southern Benin, including the relict rain forests Forét

de Lama and Forét de Lokolhi, only eight can be regard-

ed as true forest species. These are: Agama sylvanus,

Hemidactylus ansorgei, H. fasciatus, H. mabouia, H.

lamaensis sp. n., Holaspis guentheri, Varanus ornatus, and

—as the only snake — Psammophis phillipsi. Two of them,

Hemidactylus fasciatus and H. mabouia, are recorded from

Benin for the second time, H. mabouia, however, being

a recent invasive colonizer. The remaining five lizard

species, all from Lama Forest, are first country records for

Benin, one of them, H. Jamaensis sp. n., being new to sci-

ence.

Three snake species, viz. Afronatrix anoscopus, Amblyo-

dipsas unicolor, and Natriciteres variegata, are found in

both forest and savannah habitats. Of these, Afronatrix

anoscopus 1s recorded here for the second time from

Benin, but for the first time from southern Benin (the very

recent first record being from the Atakora chain in the

northwestern part of the country). Amblyodipsas unicol-

or and Natriciteres variegata are both representing the re-

spective first country records documented by voucher

specimens.

The remaining 40 species are widely distributed in the

West African savannas and have in most cases been

recorded from Benin before. Nonetheless, particularly

some snakes have been listed and/or plotted in maps of

review papers and books by e.g. David & Ineich (1999),

Chippaux (1999, 2001, 2006), or Trape & Mané (2006 b)

without locality and voucher documentation of the respec-

tive country records, so that our specimens are in some

©ZFMK

46 Klaus Ullenbruch et al.

cases nonetheless the first documented records, despite

earlier citations of the respective species from Benin.

In the following country-wide checklist, we included al-

so some new data derived from the mission to Pendjari

National Park by one of authors (OG). Among these, the

by far most remarkable find is the rediscovery of Agama

gracilimembris in Benin, from where it was first described

about ninety years ago.

COUNTRY-WIDE CHECKLIST OF THE REP-

TILES OF BENIN

The following list contains several records of species col-

lected by the Lama Forest survey of the senior author

which have not been documented before from the terri-

tory of Benin. However, as nearly all these species were

already known from the neighbouring countries in the west

(Togo) and in the east (Nigeria), their presence in Benin

is not surprising and was already regarded as probable in

several cases by Villiers (1951), Chippaux (2001) or Bauer

et al. (2006). More general books on West Africa or even

Africa as a whole (e.g. Villiers 1976, Welch 1982, Chip-

paux 1999, 2001, 2006, Trape & Mané 2006 b) often pose

the problem that a species is assumed to occur in a spe-

cific country (e.g. formulations as “from Senegal to Ugan-

da’’) rather than being documented from a specific local-

ity within the respective country; or that it is plotted on a

map without documentation of the origin of the respec-

tive data.

For completeness’ sake, we included in this list some

species recorded by Chabanaud (1916) which are definite-

ly lacking in West Africa and thus are probably due to

misidentifications. They are marked with an asterisk (*).

A re-examination of the respective voucher material as far

as still extant was, however, beyond the scope of this pa-

per. It is also noted here that the most important place of

origin of Chabanaud’s material collected by Dr. Bouet is

constantly spelled Agouagon in his papers, as it is also by

Villiers (1951). However, more recent authors (e.g. Hoog-

moed 1974, Salvador 1982) spell it “Agouagou” under

which name it is also found on modern road maps. Here,

we follow the original spelling.

It was reflecting the still poor state of knowledge of the

reptilian fauna of Benin when Loveridge (1952) stated that

among the turtles and lizards from the A. Villiers mission

(1950) no less than 8 species were new for the country,

among them even Agama agama! More than five decades

later Bauer et al. (2006) recorded — next to more first

records of geckos and a new species of Hemidactylus —

Bonn zoological Bulletin 57 (1): 31-54

also the common H. mabouia as new for the country, and

in this paper, we can list — again next to another new Hemi-

dactylus — further 8 species as new for Benin’s reptile fau-

na. Our reference to Pendjari NP and to Bemberéké are

based on data and photographic documentations collect-

ed by one of us (OG) and of a small donation of snakes

to ZFMK by T. & M. Imthurn (2000). For the latter, the

ZFMK catalogue numbers are given.

Geoemydidae

*Mauremys leprosa — Chabanaud (1917): Porto Novo (as

Clemmys leprosa). Not in Fritz & Havas (2007). Accord-

ing to Fritz (pers. comm.) this record is certainly due to

an error and should be deleted from the country’s species

list.

Testudinidae

Kinixys belliana nogueyi — Loveridge (1951): Bassila; this

paper: Bohicon-Abomey area; Pendjari NP.

Pelomedusidae

Pelomedusa subrufa — Loveridge (1951): Bassila; Zag-

nanado; this paper: Lama Forest surroundings; Pendjari

NP.

Pelusios castaneus — Sinsin et al. (1999): Benin, unspec-

ified (as P. subniger); Branch (2008): on map, but with-

out locality data; this paper: Lama Forest surroundings;

Pendjari NP (ZFMK 83613).

Trionychidae

Cyclanorbis elegans — Gramentz (2008): “Benin”, with-

out specific locality.

Cyclanorbis senegalensis — Gramentz (2008): Pendjari NP

(based on Grell 2003); this paper: Pendjari NP (Figure 18).

Crocodylidae

Crocodylus suchus — This paper: Pendjari NP (Figure 19).

Osteolaemus tetraspis — Sinsin et al. (1999): “Sud-Benin”

including the Departments of Zou, Mono, Ouémé, and At-

lantique.

©ZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 47

Fig. 18. Cyclanorbis senegalensis, juvenile from Pendjari NP,

northern Benin. Photo: O. Grell

Eublepharidae

Hemitheconyx caudicinctus — Chabanaud (1917 a):

“Haute Dahomey”; Agouagon (as Psilodactylus caudicinc-

tus); Bauer et al. (2006): Kétou; Diho (=Idiho); this pa-

per: Didja; Za-Kpota.

Gekkonidae

Hemidactylus ansorgei — This paper: Lama Forest (1st

country record).

Hemidactylus angulatus/guineensis complex — Cha-

banaud (1917 a): Agouagon (as H. brookii and H. stella-

tus); Grandison (1956, in part): “Dahomey”; Loveridge

a ry

Fig. 19. Crocodylus suchus, Pendjari NP. Photo: O. Grell

Bonn zoological Bulletin 57 (1): 31-54

(1947): Ajuda; Godomey; Grand Popo; Porto Novo;

Zomai; Loveridge (1952): Abomey; Bassila; Kous-

soukoingou; Bauer et al. (2006): Attogon; Niaouli;

Abomey; Collines du Dassa-Zoume; Manigri; Pendjari

NP; this paper: Godomey; Bohicon.

Hemidactylus beninensis — Bauer et al. (2006): Collines

du Dassa-Zoume (new species description).

Hemidactylus fasciatus — Bauer et al. (2006): near Kétou;

this paper: Lama Forest (2nd country record).

Hemidactylus mabouia — Bauer et al. (2006): Godomey,

Abomay, Niaouli; this paper: Cotonou (2nd country

record).

Hemidactylus muriceus — Bauer et al. (2006): near Kétou.

Hemidactylus lamaensis — This paper: Lama Forest (new

species description).

Lygodactylus conraui — Bauer et al. (2006): Lokoli Fo-

rest.

Phyllodactylidae

Ptyodactylus ragazzii — Bauer et al. (2006): Chutes de

Koudou, PN du W.

Tarentola ephippiata — Bauer et al. (2006): Pendjari NP.

©ZFMK

48 Klaus Ullenbruch et al.

Fig. 20. Agama gracilimembris, Pendjari NP, first rediscovery

in Benin after its first description from this country nine decades

ago. Photo: O. Grell

Agamidae

Agama agama complex — Loveridge (1952): Bassila; Kou-

tiacou; Zagnanado; this paper: Godomey nr. Cotonou; Co-

tonou; Bohicon; Pendjari NP.

Agama sylvanus — This paper: Lama Forest (1st country

record).

Agama gracilimembris — Chabanaud (1918): “Dahomey”

(original description); this paper: Pendjari NP.

Note: This little agama with unresolved intrageneric re-

lationships was subsequently found in Nigeria and in the

Central African Republik (Grandison 1968, 1969, Joger

1990), the gap between these two records being bridged

by a find in northern Cameroon (Béhme 1975). The pho-

tographic record from Pendjari NP (Figure 20) is the first

find of this rare agama in Benin since its original descrip-

tion in 1918, a rediscovery after 9 decades!

Chamaeleonidae

Chamaeleo africanus — This paper: Atakora chain (1st

country record, based on photograph).

Note. A photograph of a “chameleon” published in a pop-

ular book on Benin by Englebert (1973, cited after Con-

rad 1999) without identifying the species shows clearly a

specimen of C. africanus. Its inclusion by Conrad (1999)

into her unpublished thesis means that it is still the first

published country record for Benin.

Chamaeleo gracilis — This paper: Abomey-Didja; Pend-

jar1 NP.

Bonn zoological Bulletin 57 (1): 31-54

Chamaeleo necasi — Ullenbruch et al. (2007), this paper:

Bohicon-Za-Kpota, Houegbo.

Chamaeleo senegalensis — Loveridge (1952): Zagnanado;

this paper: Bohicon; Abomey-Didja; Pendjari NP.

Scincidae

Chalcides thierryi — Loveridge (1952): Koussokoingou (as

Chalcides ocellatus thierryi).

Mochlus guineensis — Chabanaud (1917): Agouagon; this

paper: Lama Forest; Cotonou.

Panaspis togoensis — Chabanaud (1917): Agouagon (as

Lygosoma (Riopa) dahomeyense, syn. fide Schmitz et al.

2005); Loveridge (1952): Bassila (as Lygosoma (Panaspis)

breviceps togoense) Grandison (1956): “Dahomey” (as P.

breviceps togoense); Fuhn (1972): “Dahomey: Agouagou”

(as Panaspis kitsoni); this paper: Lama Forest; Pendjari

NP.

Trachylepis affinis — Chabanaud (1917): Satadougou (as

Mabuia raddonii); Loveridge (1952): Abomey; Bassila;

Koussokoingou; Tanogou (as Mabuya blandingii); this pa-

per: Godomey nr. Cotonou; Lama Forest; Pendjari NP.

Trachylepis perrotetii— Chabanaud (1917): Agouagon (in

part as Mabuia albilabris, fide Hoogmoed 1974);

Loveridge (1952): Abomey; Bassila; Koussokoingou (as

Mabuya perroteti); this paper: Godomey nr. Cotonou;

Pendjari NP.

Trachylepis maculilabris — Chabanaud (1917): Agouagon;

Loverdige (1952): Bassila.

Trachylepis quinquetaeniata sharica Sternfeld, 1917 —

Chabanaud (1917): Agouagon (in part as Mabuia albi-

labris, fide Hoogmoed 1974); Loveridge (1952): Bassila

(as Mabuya quinquetaeniata sharica); this paper: Pend-

jari NP.

Gerrhosauridae

Gerrhosaurus major zechi — Loveridge (1952): Kous-

sokoingou.

Lacertidae

Acanthodactylus boueti — Chabanaud (1917): Agouagon

(as A. (Latastia) boueti); Loveridge (1952), Salvador

(1982): Bassila.

©OZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 49

Heliobolus nitidus — Loveridge (1952): Bassila (as

Eremias n. nitida); this paper: Pendjari NP.

Holaspis guentheri — This paper: Lama Forest (1st coun-

try record).

Amphisbaenidae

Cynisca leucura — Chabanaud (1917a): Agouagon; “Da-

homey” (as Amphisbaena leucura); Loveridge (1952):

Natitingou.

Varanidae

Varanus exanthematicus — This paper: Abomey-Didja.

Varanus niloticus — Chabanaud (1917a): Agouagon;

“Haut Dahomey”; this paper: Godomey nr. Cotonou;

Ouémé River near Togbota; Pendjari NP (photographic

voucher of an unusually pale olive-coloured juvenile, Fig-

ure 21).

Varanus ornatus — This paper: Abomey-Didja; Lama For-

est (1st country record).

Leptotyphlopidae

Leptotyphlops bicolor — Chabanaud (1916): Porto Novo

(as Glauconia. b. gruveli).

Leptotyphlops brevicauda — Bocage (1887): “Dahomey”

(as Stenostoma brevicauda); Hahn (1980): “Dahomey” (as

Leptotyphlops brevicaudus).

*Leptotyphlops nigricans — Chabanaud (1916): “Da-

homey” (Identification probably erroneus, taxon not in

west Afria).

Typhlopidae

Ramphotyphlops braminus — Trape & Mané (2006 b):

Cotonou.

Typhlops punctatus — Chabanaud (1916): “Dahomey”;

Chabanaud (1917a,b): Ouidah, Agouagon.

Boidae

Gongylophis muelleri — Trape & Mané (2006 b): on grid

map but without specific locality.

Bonn zoological Bulletin 57 (1): 31-54

Fig. 21.

Varanus niloticus, subadult, unusally light-coloured

specimen from Pendjari NP. Photo: O. Grell

Python regius — Villiers (1951): “trés commun”, voucher

specimen from Abomey; this paper: Godomey nr. Coto-

nou; Za-Kpota; Lama Forest.

Python sebae — Villiers (1951): “Trés commun partout”,

specimens from Abomey; this paper: South Benin.

Colubridae

Afronatrix anoscopus — Chirio & Ineich (2009): Niangou

NE of Tanguieta; Atakora Prov.; this paper: Lokoli For-

est (2nd published country record, regarded already as

probably present by Villiers 1951 and Chippaux 2001).

Bamanophis dorri — Chippaux 1999, 2006; N Benin: un-

specified (as Coluber dorri); Trape & Mané (2007): on

grid map but without specific locality (as Hemorhhois dor-

ri; Schatti & Trape (2008): unspecified.

Crotaphopeltis hippocrepis — Rasmussen (2000): Soubrou-

kou.

Crotaphopeltis hotamboeia — Chabanaud (1916): “Daho-

mey” (as Leptodira hotamboeia); Chabanaud (1917 b):

Agouagon (as Leptodira hotamboeia); this paper: Godo-

mey nr. Cotonou.

Dasypeltis gansi — Chabanaud (1916): “Dahomey” (as D.

scabra palmarum); Trape & Mané (2006 a): Lanta; this

paper: Za-Kpota.

Dispholidus typus — This paper: Bohicon-Didja (1st coun-

try record; regarded as probably present by Villiers 1950).

Grayia smythii — Chabanaud (1916): near Sakété, Porto-

Novo.

©ZFMK

50 Klaus Ullenbruch et al.

Hormonotus modestus — Chabanaud (1916): “Dahomey”;

Chabanaud (1917 b): near Lac Azzi.

Lamprophis fuliginosus — Chabanaud (1916): “Dahomey”

(as Boaodon fuliginosus); Chabanaud (1917b): Agouagon

(as Boodon fuliginosus); Villiers (1951): Koussokoin-

gou/Atakora (as Boaedon fuliginosus); this paper: Dan

Forest; Lama Forest; Didja, Za-Kpota.

Lamprophis lineatus — Chabanaud (1916): “Dahomey” (as

Boaodon linatus); Chabanaud (1917b): Agouagon (as

Boodon lineatus); Villiers (1951): Abomey (as Boaedon

lineatus).

*Lycophidion capense — Chabanaud (1916): “Dahomey”

(listing of capense and capense multimaculatum proba-

bly due to identification error: both taxa not in West

Africa).

Lycophidion semicinctum — Villiers (1951): Abomey; this

paper: Bohicon; Didja.

Mehelya crossi — Chabanaud (1916): “Haut-Dahomey” (as

Simocephalus capensis); Loveridge (1940): Bassila; Vil-

liers (1951): Abomey.

Meizodon coronatus — Trape & Mané (2006 b): on grid

map but without specific locality.

Fig. 22. Psammophis elegans from Pendjari NP, dorsal pattern.

Photo: O. Grell

Bonn zoological Bulletin 57 (1): 31-54

Meizodon regularis — Chippaux (2001): Benin (on map,

without documented locality); this paper: Godomey nr.

Cotonou (2nd country record).

Natriciteres olivacea — Chabanaud (1916): “Dahomey” (as

Tropidonotus olivaceus); this paper: Lama Forest (1st doc-

umented locality record for Benin).

Natriciteres variegata — This paper: Lama Forest (1st

country record).

Philothamnus irregularis — Trape & Mané: on grid map

but without specific locality; this paper: Lama Forest.

Philothamnus semivariegatus — Chabanaud (1916): “Da-

homey”; Chabanaud (1917a): Cotonou, Agouagon; Villiers

(1951): Abomey; this paper: Lama Forest.

Polemon neuwiedii — Chabanaud (1916): “Dahomey” (as

Miodon neuwiedi); Agouagon (as Miodon neuwiedi).

Prosymna meleagris — Chabanaud (1916): “Dahomey”;

Chabanaud (1917a): Agouagon.

Psammophis elegans — Loveridge (1941): Ouidah; this pa-

per: Cotonou; Bembéréké ZFMK 72885); Pendjari NP

(Figure 22).

Psammophis lineatus — Chabanaud (1916): “Dahomey”;

Chabanaud (1917 b): Agouagon.

Psammophis cf. phillipsi— Chabanaud (1916) “Dahomey”

(as P. sibilans); this paper: Abomey; Za-Kpota; Pendjari

NP (Figure 23).

Psammophis phillipsi (s.str.) — Chabanaud (1916): “Da-

homey” (as P. regularis); this paper: Godomey near Coto-

nou.

Psammophis praeornatus — Chabanaud (1916): “Da-

homey” (according to Villiers 1951 not documented).

Psammophis schokari — Chabanaud (1916): “Dahomey”

(as P. shokari: considered to be of doubtful provenance

by this author).

Psammophis sudanensis — This paper: Didja; Za-Kpota,

Abomey, Bohicon (1st country record).

Rhamphiophis oxyrhynchus — Chirio & Ineich (1991):

Benin, unspecified; Trape & Mané (2006): on grid map,

but without specific locality; this paper: Godomey nr.

Cotonou; Za-Kpota.

©OZFMK

Reptiles from southern Benin, West Africa, with the description of a new Hemidactylus 51

Fig. 23. Psammophis cf. phillipsi from Pendjari NP, showing

the characteristic reddish dorsilateral stripes. Photo: O. Grell

Scaphiophis albopunctatus — Bocage (1895, 1896): Aju-

da; Chabanaud (1916): “Dahomey”; Chabanaud (1917a):

Agouagon; Broadley (1994): Parakou, and Segbana; this

paper: Lama Forest (Didja).

Telescopus variegatus — Chabanaud (1916): “Dahomey”

(as Zarbophis variegatus), Chabanaud (1917a): Agouagon

(as Tarbophis variegatus); Villiers (1951): Abomey (as

Tarbophis variegatus).

*T. semiannulatus — Chabanaud (1916): “Dahomey” (as

Tarbophis semiannulatus. Repeated by Villiers (1951), but

probably due to misidentification, as taxon not in present

in West Africa).

Thelotornis kirtlandii — Villiers (1951): Bassila.

Toxicodryas blandingii — Chabanaud (1916): “Dahomey”

(as Dipsadomorphus blandingii); Villiers (1951): Zag-

nanado, Abomey (as Boiga blandingii).

Toxicodryas pulverulentus — Chabanaud (1917b): Porto

Novo (as Dipsadomorphus boueti, syn. fide Villiers 1951).

Atractaspididae

Amblyodipsas unicolor — Considered to be probably in

Benin by Villiers (1950) and Meirte (1999); Chippaux

(2001): On map, but without documented locality; this pa-

per: Za-Kpota (first documented country record).

Aparallactus modestus — Boettger (1898): Grand Popo;

Witte & Laurent (1947): ’Gross Popo”.

Atractaspis aterrima — Villiers (1951): Abomey; Ras-

mussen (2005): “Benin, without locality”.

Bonn zoological Bulletin 57 (1): 31-54

Atractaspis dahomeyensis — Barboza du Bocage (1887):

Zomai; Villiers (1951): Abomey; Trape & Mané (2006b):

Parakou.

Atractaspis watsoni — Trape & Mané (2006 b): on grid

map but without specific locality.

Elapidae

Dendroaspis viridis — Chabanaud (1916): “Dahomey” (as

Dendraspis viridis); this paper: Bohicon.

Elapsoidea semiannulata moebiusi — Chabanaud (1916):

“Dahomey” (as Elapechis guentheri); Chabanaud (1917

b): Agouagon (as Elapechis guentheri); this paper: Lama

(Didja).

Naja melanoleuca — Chabanaud (1916): “Dahomey”; Vil-

liers (1951): Abomey.

Naja nigricollis — Chabanaud (1916): “Dahomey”; Cha-

banaud (1917): Agouagon; Villiers (1951): Abomey; this

paper: Lama Forest; Pendjari NP.

Naja senegalensis — Trape et al. (2009): Niénié.

Viperidae

Bitis arietans — Chabanaud (1916): “Dahomey”; Cha-

banaud (1917 b), Villiers (1951): Koussokoingou/Atako-

ra, Zagnanado (as B. lachesis); this paper: Abomey; Be-

mbéréké (ZFMK 72886).

Bitis nasicornis — Chabanaud (1916): “Dahomey”.

Causus maculatus — Chabanaud (1916): “Dahomey” (as

C. rhombeatus); Chabanaud (1917b): Agouagon (as C.

rhombeatus); Villiers (1951): Koussokoingou/Atakora (as

C. rhombeatus); this paper: Bohicon/Didja, Abomey.

Echis. ocellatus — Chabanaud (1916): “Dahomey” (as E.

carinatus); Chabanaud (1917 b): Agouagon (as E. cari-

natus); Villiers (1951): Koussokoingou/Atakora (as E. car-

inatus); this paper: Didja; Bembéréké (ZFMK 72867,

73091); Pendjari NP.

Acknowledgements. We thank Prof. Dr. Peter Nagel and Dr.

Ralf Peveling, University of Basel, that they provided the op-

portunity to KU to work within the BIOLAMA project. Thanks

are also due to Lamidi Konetche and Thibault Lachet for some

additional gecko specimens. WB thankfully acknowledges Dr.

Werner Schréder, Gottingen, and Mr. Thomas & Mrs. Marianne

Imthurn, Bembéréké/Benin, for some reptile from Godomey and

Bembéreékeé, respectively.

©OZFMK

We further thank PD Dr. Mark-Oliver Rédel and Mr. Frank

Tillack, Museum ftir Naturkunde Berlin, for the loan of relevant

gecko type material, and Dr. Colin McCarthy, Natural History

Museum London, and Dr. Heinz Grillitsch, Naturhistorisches

Museum Vienna, for photos of further important type material

of geckos and a snake. Prof. Dr. Uwe Fritz, Senckenberg

naturhistorische Sammlungen Dresden, kindly commented on a

chelonogical issue while Dr. Andreas Schmitz, Muséum d’His-

toire Naturelle, Geneva, helpfully commented on skink taxon-

omy and on the manuscript as a whole. Mr. Philipp Wagner

(MSc) and Mrs. Ursula Bott (both from ZFMK Bonn) provid-

ed again support with their photographic and technical skills.

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©ZFMK

Bonn zoological Bulletin Volume 57

Issue | pp. 55-64 Bonn, April 2010

Fitness components in the relationship between

Rhopalapion longirostre (Olivier, 1807) (Insecta: Coleoptera: Apionidae) and

Alcea rosea (Linnaeus, 1758) (Malvaceae).

Analysis of infestation balance of a herbivorous weevil and its host plant

Gertha Wilhelm', Hans L. Nemeschkal', John Plant? & Hannes F. Paulus?

' Faculty of Life Sciences, Department of Theoretical Biology, University of Vienna, Althanstr. 14,

A-1090 Vienna, Austria;

E-mails: a9404775@unet.univie.ac.at; Hans.Leo.Nemeschkal@univie.ac.at;

* Faculty of Life Sciences, Department of Evolutionary Biology, University of Vienna,

Althanstr. 14, A-1090 Vienna, Austria; E-mails: john.plant@aon.at; hannes.paulus@univie.ac.at

Abstract. Multivariate statistics (principal components analysis, path analysis) were used to investigate fitness compo-

nents of the interactions between the weevil Rhopalapion longirostre (Olivier, 1807), Apionidae, Coleoptera and its host

plant Alcea rosea (Linnaeus, 1758), Malvaceae. We focused on the activities of the larvae such as the choice of seeds for

consumption, the preparation of seed chambers as a site for pupation, as well as the construction of escape holes through

which the adults later emerge.

The analyses revealed that the optimal conditions for successful development of the weevils depended on the availabil-

ity of seed capsules characterized by a high number of well developed seeds, few undeveloped and few spoiled seeds.

The high number of larvae, pupae and not emerged adults found in the seed capsules corresponds with the successful

emergence of adults. Egg deposition by the females in appropriate flower buds of the host plant, together with larval con-

tribution to overall reproductive success are important fitness components in the life cycle of the weevil. The reproduc-

tive success of the plant is partly guaranteed by a high number of seed chambers, many well developed seeds, many not

infested seeds, and few undeveloped seeds. Furthermore, the extended blooming period of the host plant serves as an es-

cape strategy by developing buds early or late in the season that cannot become entangled in the life cycle of R. lon-

girostre.

Key words. Alcea rosea, Malvaceae, Rhopalapion longirostre, Apionidae; Host plant-parasite interaction, infestation bal-

ance, larval contribution, path analysis, PCA, reproductive success

INTRODUCTION

Life history theory provides a framework for analyzing

the contribution of major features of the life cycle to re-

productive fitness and seeks to place these features in an

evolutionary context (Roff 1992; Stearns 1992; Stearns &

Hoekstra 2000). The life history events of most of the

62,000 described species of weevils or snout beetles (Cur-

culionoidea, Coleoptera) (Alonso-Zarazaga & Lyal 1999),

and their immature stages are incompletely documented

and more or less episodic. As holometabolous insects,

weevils undergo a complete metamorphosis which pro-

duces larval and adult stages each with their own life

habits and physiological and behavioral traits thus enhanc-

ing the possibility for adaptation, specialization and evo-

lution. The phenomenal diversity of weevils was discussed

recently in evolutionary terms by Oberprieler et al. (2007).

Considered as a group, weevils utilize nearly all parts of

host plants from almost all plant taxa (Anderson 1995).

The larvae of most Apionidae, with about 1800 described

species (Kuschel 1995), feed internally in stems, leaves,

buds, galls, fruits or seeds of their specific host plants. The

evolution of larval habits and host plant associations are

taken for analysis of larval role in weevil’s diversification

(Marvaldi et al. 2002).

In this paper we examine several fitness components of

the weevil Rhopalapion longirostre (Olivier, 1807) which

is monophagous on its host plant Alcea rosea (Linnaeus,

1758), with special focus on larval contribution to repro-

ductive success (Reavey & Lawton 1991). Seeds of A.

56 Gertha Wilhelm et al.

rosea, presumably derived from Anatolia (Hammer

1994), were brought to France in the 18 century. The

beauty and popularity of this garden plant, known as the

hollyhock, passe-rose, rose trémiére, or Stockrose, was

lovingly recorded in verse by King Ludwig XVI of France

in the 18‘ century: L- Amour de moi s’y est éclosé / dedans

un petit jardinet, / oi croit la rose et le muguet / et aussi

bien la passe-rose.

Today, the plant occurs in many parts of Europe, North

and South America, mainly due to its cultivation in gar-

dens. The development of the larvae of R. longirostre takes

place in the buds and seeds of A. rosea and the expansion

of the weevil’s distribution range has proceeded in inti-

mate association with that of the host plant (Perrin 1995;

Gonget 1997).

Females of R. longirostre have a body size from 2.4 to 3.4

mm excluding the rostrum (Freude et al. 1983). They em-

ploy their extremely elongated rostrum (average size 2.2

mm, standard deviation +/- 0.2 mm) (Wilhelm 2004), to

bore holes into developing buds of A. rosea. The plant at-

tains a height of 1 to 3 meters and can persist for more

than two years. The buds appear in a long inflorescence

composed of 30 to 50 buds. Every bud is surrounded by

thick inner and outer sepals which have stiff grey-green

hairs (Adler et al. 1994). After, or even during copulation,

with the male sitting on her back, the female bores a chan-

nel into a selected bud using her rostrum. She turns around

and inserts her ovipositor into the bore channel penetrat-

ing the outer and inner sepals of the plant to lay three to

four eggs into it. Female weevils prefer to bore into buds

located in the middle of the inflorescence. These buds will

usually not bloom in the following 2 weeks thus allow-

ing enough time for the development of the larvae. If a

female is among the first to bore a hole into the bud, she

chooses the side of the plant which receives the morning

sunlight. If several females have oviposited in the same

bud, the boring channels are widely spaced from each oth-

er, presumably to avoid competition among the larvae for

food. The first instar larvae are legless, as most of wee-

vils larvae have lost the development of legs (Crowson

1955; Stehr 1991; Marvaldi 1997; Farrell 1998), about 0.4

mm long, and they hatch after 3 to 4 days. If the bore chan-

nel is in the upper part of the bud then the larvae of the

first stage consume the surrounding pollen grains. How-

ever, if the bore channel is located at the bottom side of

the bud, then the larvae feed only on the soft tissue of the

bud. All larvae bore feeding channels toward the direc-

tion of the ovules in the developing seed capsule, using

their strong mandibles to bite and chew. By the time the

bud has blossomed and before it falls off, all larvae must

have migrated below the ovary. Within a few days the up-

per part of the blossom wilts and falls to the ground. When

the seeds of A. rosea are fully developed, each larva tries

Bonn zoological Bulletin 57 (1): 55-64

to enter a seed chamber from the bottom of the seed cap-

sule and consume the entire contents of the seed from

within. Prior to pupation, the last instar bores a hole on

the broadest side of the seed chamber to later escape when

it reaches the adult stage. The escape hole is filled in with

a secretion produced by the larva to protect the pupa and

the developing weevil against outside influences, e.g. wet-

ness and sunlight. Pupation lasts between 2 to 3 weeks de-

pending on weather conditions. The darkening of the adult

weevil’s exoskeleton is visible through the transparent

membrane of the pupa beginning at the apex of the ros-

trum and the elytra. The adult beetle slowly emerges from

the dry and stiff seed chamber over a period of several

hours using its rostrum to get out through the escape hole.

A duration of 8 to 10 weeks is required from egg deposi-

tion to the emergence of adult weevils.

Additional details on the biology of R. /ongirostre are

found in Dieckmann (1977), Behne (1998), Pupier

(1997), Sprick et al. (2002) and Wilhelm (2004).

The interaction between the host plant and the weevil is

reflected in a successful joint expansion in Europe over

the last twenty years. To demonstrate this a null hypoth-

esis was postulated which denied significant interactions

between the life cycles of host plant and weevil. To test

null hypothesis in detail, we posed the following ques-

tions:

(1) What parameters lead to optimal prerequisites for suc-

cessful development of weevils in seed capsules? In par-

ticular, what are the conditions of capsules which contain

larvae, pupae and immature adults? Do these conditions

correspond with successful emergence of adults?

(2) Which counter strategies are taken by the host plant

to survive or to ensure its successful growth and repro-

duction?

(3) How effective is larval contribution to the overall re-

productive success of the weevil?

MATERIALS AND METHODS

To address these questions we examined dry seed capsules

collected from 20 Alcea rosea plants growing on the ter-

race at the University of Vienna, Althanstrasse 14, in the

first week of Nov. 2006. For measurements we examined

only dry capsules (n=205) having a complete set of seed

chambers. Seed capsules, which were incomplete because

some seed chambers had already fallen to the ground, were

not investigated. The plants, 1-20, contained the follow-

ing number of seed capsules: 7, 10, 11, 4, 5, 6, 21, 4, 16,

oh, (UNS 3}, 24 IS, 13), 7, OO, O

©ZFMK

Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 57

infested bud

host plant

seed capsule

(whole)

seed capsule

(schema)

adult weevil

Fig. 1. Brief outline of life history stages of the weevil Rhopalapion longirostre on its host plant, Alcea rosea. Schema showing

various conditions of a seed capsule: seed chambers with 1) fully developed seeds, 2) seeds infested with larvae, 3) pupae and clo-

sed escape holes, 4) not emerged adults and closed escape holes, 5) escape holes, 6) spoiled seeds, 7) undeveloped seeds. Abbre-

viations of infested bud: b bore channel, 0 ovules, p pollen, s sepal; sc seed chamber.

The diameter of each capsule was measured 5 times to

cancel out error. The number of dry seeds per capsule and

the condition of every capsule were documented. The seed

chambers were opened under a dissecting microscope to

determine the inner condition. The infested condition was

indicated by the presence of larvae, pupae or not emerged

adults, or empty seed chambers with open escape holes.

The uninfested condition was indicated by the presence

of seed chambers which are fully developed, undeveloped

or spoiled. In the following, the data were analyzed thor-

oughly by multivariate statistics to estimate the importance

of animal and plant related factors and their interactions

on the reproductive fitness of weevil and plant.

For the following case study nine character variables of

the seed capsules were measured for the statistical analy-

ses.

Diameter of capsules in mm: diam (plant related: PR)

Number of seed chambers per capsule: nrse (PR)

Bonn zoological Bulletin 57 (1): 55-64

Number of open beetle escape holes per capsule: nreh (an-

imal related: AR), i.e. estimate of the number of beetles

which have successfully escaped.

Number of dead larvae per capsule: nrla (AR)

Number of dead pupae per capsule: nrpu (AR)

Number of dead, not emerged adults per capsule: nrne

(AR)

Number of fully developed seeds per capsule: nrds (PR)

Number of undeveloped seeds per capsule: nrus (PR)

Number of spoiled seeds per capsule: nrss (PR)

In preparation of multivariate analyses, the original data

were transformed as follows: The first variate was loga-

rithmically transformed (natural log). In order to fulfill the

©ZFMK

58 Gertha Wilhelm et al.

Table 1. Correlation matrix of transformed variates and significant levels: *** = highest significant P-value < 0.001, ** = highly

significant P-value < 0.01, * = significant P-value < 0.05. Bonferroni corrected. Variates: diam diameter of seed capsule, nrse

number of seed capsules, nreh number of escape holes, nrla number of larvae, nrpu number of pupae, nrne number of not emer-

ged adults, nrds number of fully developed seeds, nrus number of undeveloped seeds, nrss number of spoiled seeds. The signi-

ficance of correlation coefficients was determined by a random permutational test (Nemeschkal 1999).

diam nrsc nreh nrla nrpu nrne nrds nrus nrss

diam 1.0000

nrse 0.075632 1.0000

nreh 0.057043 0.167632* 1.0000

nrla 0.032532 0.293509*** - 0.135968 1.0000

nrpu 0.013449 0.2803 19*** - 0.044457 0.538041 *** 1.0000

nme = - 0.07478 0.129488 0.128997 0.161018 0.212016*** 1.0000

nrds 0.036070 - 0.046134 - 0.247253%** — - 0.304999%** = - 0.31991*** - 2.242189** 1.0000

nrus 0.000937 0.163294 - 0.257581*** — - 0.028526 - 0.088356 - 0.046297 — - 0.306250*** 1.0000

nrss - 0.001917 0.254107*** - 0.26796 1*** 0.096727 0.208869** 0.088184 - 0.402289*** 0.165270* 1.0000

Table 2. Factor matrix, axis | to 7, and significant axes of loadings. *** = highest significant, P-value < 0.001, ** = highly si-

gnificant P-value < 0.01, * = significant P-value < 0.05, ns= no significance P-value > 0.05, eigenvalues, and percentages of the

total variance. The significant levels were determined by a random permutational test (NEMESCHKAL 1999, 10000 iterations).

Potential win for animal and plant. + A, + P; potential loss: - A, - P. Abbreviations of axes: SUPLA for ‘successful plant (seed)

production’; SUAN for ‘successful animal production’; SUMIX for ‘integration of the two trends, successful animal and plant

production’; LIFA for ‘late infestation axis’ and DASA for ‘damaged seed and animal axis’.

Factor axes Fl F2 F3 F4 F5 F6 F7

Abbreviations SUPLA SUAN SUMIX LIFA DASA nn. nn.

var 3 nreh - 0.083229ns_ - 0.72391 6*** 0.568846*** - 0.237805** 0.174577ns_ - 0.024872ns 0.2403 88*««

var 4 nrla 0.680778 - 0.168066« - 0.46383 1** - 0.297076*** -0.174923ns 0.394877*** 0.135561ns

var 5 nrpu 0.726492*** - 0.245330 — - 0.380804 - 0.114225ns 0.074140ns_ - 0.498353*** 0.010510ns

var 6 nrne 0.430808*** — - 0.379749 +«« O.211681** 0.631854 - 0.475213*** 0.012534ns 0.012746ns

var 7 nrds - 0.743359%* — 0.012772ns - 0.532765*** 0.217806** -0.116773ns -0.103023ns 0.302864++«

var 8 nrus 0.217084+« 0.687623 **« 0.397469» - 0.274868** - 0.443069 -0.147826ns 0.168382ns

var9 nrss 0.559095*** 0.45827 1 ««« 0.118490ns 0.384199 0.525592** 0.073805ns 0.184544 +«

Eigenvalues 2.096067 1.439697 1.184405 0.827693 0.778592 0.442973 0.230572

percentage of 29.94 20.57 16.92 11.82 11.12 6.33 3.29

total variance

animal win/loss -A +A +A +A -A

Plant win/loss +P -P +P +P -P

linearity requirement, variates 2 to 9 were substituted by

their square roots. Finally, a product moment correlation

matrix was calculated between all variates over the 205

capsules (Table 1).

The correlation structure of variates 3 to 9 was then ana-

lyzed by a principal components analysis (PCA, for de-

tails see Morrison 2004). An orthogonalization was

reached via PCA by calculating a full analysis and extract-

ing all possible, i.e. 7, axes (Stat. Graphics — Version 7.3

for MS DOS). The eigenvectors were scaled according to

Bonn zoological Bulletin 57 (1): 55—64

eigenvalues leading to a factor matrix (Table 2). The sig-

nificance of loadings on principle axes was determined by

a random permutational test, 10000 random permutations

each, (Nemeschkal 1999). Note that the original variates

are represented on factor axes by their loadings. A load-

ing is the product moment correlation of an original vari-

ate with the factor axis. For details and further statistical

explanation, see Morrison (2004).

Path analysis, a special type of regression analysis, (Stat.

Graphics — Version 7.3 for MS DOS) was applied to rep-

©ZFMK

Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 59

resent the complicated animal-plant relations in diagram-

matic form (Path diagrams, Figs. 2 and 3). In addition, path

analysis has the advantageous option to focus dependen-

cies either on plant or for animal variables (for theoreti-

cal details on path analysis, see Sokal & Rohlf (1995).

For reasons of simplicity, our analyses are mainly based

on procedures of the general linear model. In order to test

whether a loss of information in the data structure has oc-

curred due to data transformations, we calculated, in par-

allel log-linear analyses with the non-transformed data (see

Appendix).

RESULTS

Of the 205 examined seed capsules, each contained an av-

erage of 35 individual seed chambers (ll 34.7758, ul

35.6798; ll: lower limit, ul: upper limit of 95% confidence

of average). Of the 35 seeds chambers, an average of 13

(Il 11.6393, ul 13.4734) were fully developed, inferring

the reproductive success of the host plant, 7 (11 6.0002, ul

7.6396) had open escape holes, inferring the potential re-

productive success of the weevils. The remaining 15 seed

chambers (on average per capsule) were either infested

(documented by dead larvae, pupae or not emerged adults),

undeveloped or spoiled by mold or mites.

The results from the PCA and the path analyses reveal de-

tailed information on the complex system of interactions

between weevil and host plant. For the sake of simplifi-

cation, the diameter and number of seed chambers were

omitted from the PCA and path analyses since the simple

correlation between the diameter (diam) and the remain-

ing variates are not significant (Table 1) and since the

number of seed chambers (nrsc) is a summation of vari-

ates 3 to 9.

A total of 7 axes are significant (Table 2). Factor axes |

to 5 are the most important, accounting for over 90% of

the total variance.

Results summarized for the five most important factor ax-

es:

F1 represents nearly 30% of total variance. Positive and

negative signs of the loadings indicate the reverse propor-

tionality in the variational trends: nrla, nrpu, nrne, nrss (

positive and very highly significant) and nrus (highly sig-

nificant) versus nrds (negative and very highly significant).

F2 represents 21% of total variance. Cumulatively, the ax-

es F1 and F2 explain half of the total variance. The load-

ings are contrasted to each other by their signs (+/-); i.e.

nrus and nrss (very highly significant) and with positive

Bonn zoological Bulletin 57 (1): 55-64

signs, versus nreh, nrpu, nrne (very highly significant) and

nrla (significant), both with negative signs.

F3 explains 17% of the total variance, and by accumula-

tion of axis Fl to F3 more than two thirds of total vari-

ance are explained. The signs of loadings, nreh, nrne and

nrus (very highly significant) contrast with those of nrla

and nrpu (very highly significant).

F4 represents about 12% of total variance. Axes F1 to F4

cumulatively explain more than three quarters of total vari-

ance. The loadings are grouped by positive signs for nrne,

nrss (very highly significant) and nrds (highly significant)

and negative signs for nrla, nrus (very highly significant)

and nreh (highly significant).

F5 explains 11% of total variance. The positive sign of

the loading nrss (very highly significant) contrasts with

those of nrne and nrus (very highly significant).

Factor axes 6 and 7 together provide 9% of total infor-

mation and will not be further considered in this study.

Path Analyses

Two path analyses were conducted to focus on the main

influences affecting the animal’s or plant’s life cycle. Vari-

ables 3 to 9 served as the basis for both path analyses. The

first analysis determines the influence of variables 4 to 9

on variable 3 (nreh: number of escape holes per capsule,

AR = animal related) (Fig. 2). In the second analysis vari-

able 7 (nrds: number of fully developed seeds per capsule,

PR = plant related) the variables 3 to 6, 8 and 9 take in-

fluence on variable 7 (Fig. 3).

Path Analysis I (Fig. 2): Since variable 3, nreh, number

of escape holes per capsule, may be an indicator of fit-

ness for the weevil, it was subjected to a multivariate path

analysis. Simple arrows indicate the directions of influ-

ences of the predictors on criterion (= direct paths), dou-

ble arrows indicate the correlation between predictor vari-

ables. The strength of influence is given by the value of

the path coefficient. The path coefficient is a standardized

multiple regression coefficient (path coefficient = regres-

sion coefficient times standard deviation of predictor di-

vided by the standard deviation of criterion). The path co-

efficient quantifies the amount of change in criterion when

progressing in independent variables. There are 4 direct

paths from the predictors to the criterion. The first direct

path proceeds from var 4, nrla, to the criterion and is neg-

ative and highly significant. It infers that the more dead

larvae per capsule, the fewer escape holes. The second di-

rect path from var 7, nrds, to the criterion is negative and

very highly significant: the more well developed seeds,

the fewer open escape holes. The third direct path proceeds

OZFMK

60 Gertha Wilhelm et al.

ky 0.5380 I

ee OFS 19310

** 0.208869

* 0.165270

*** _0 306250

Variables

nreh number of escape holes

nrla number of larvae

nrpu number of pupae

nrne number of not emerged adults

nrds number of fully developed seeds

nrus number of undeveloped seeds

nrss number of spoiled seeds

Fig. 2. Path diagram | showing 6 predictor variables and a residual effecting one criterion (var 3, nreh, number of escape holes,

an animal related variate),

2= ().414525. The path diagram explains 41.45% of the variance of the criterion. Only significant paths

are shown. The significance of paths was determined by random permutational test; 10000 iterations each; Bonferroni Correction.

Simple arrows indicate the direction of influences of the predictors to the criterion (= direct paths), double arrows indicate the cor-

relation between predictor variables. The strength of influence is given by the value of the path coefficient.

from var 8, nrus, to the criterion and is negative and very

highly significant: the more undeveloped seeds, the few-

er escape holes. The fourth direct path proceeds from var

9, nrss, to the criterion and is negative and very highly sig-

nificant: the more spoiled seeds per capsule, the fewer

open escape holes. In addition, several indirect effects are

apparent. Both direct and indirect paths in the diagram

must be combined to indicate an overall influence on the

criterion. The most important indirect path proceeds from

nrla to the criterion via nrds, and is very highly signifi-

cant: the more larvae, the fewer well developed seeds and

the more escape holes (indirect effect: -0.304999 «

-0.624623 = 0.190509). Direct and indirect paths are mu-

tually counteracting, therefore it is the summation of all

effects that indicates the overall influence on the criteri-

on, which in this case is negative: direct effect: nrla > nreh

-0.277150 plus indirect effect: nrla > nrds > nreh

0.190509 results in an overall effect of -0.086641. A sec-

ond indirect effect proceeds from nrla to the criterion via

Bonn zoological Bulletin 57 (1): 55-64

nrpu, nrne and nrds: the more larvae, the more pupae and

the more not emerged adults means that there are fewer

well developed seeds and more escape holes in the seed

capsule (0.538041 * 0.212016 * -0.242189 * -0.624623

= 0.017257). Another indirect effect between nrpu and

nrds to the criterion is very highly significant: the more

pupae, the fewer well developed seeds and the more es-

cape holes (-0.31931* -0.624623 = 0.19945). The indirect

effect which proceeds from nrds and nrus to the criterion

is very highly significant: the more developed seeds, the

fewer undeveloped seeds and the more escape holes (-

0.30625) * -0.389241 = 0.11921. The indirect effect which

proceeds from nrds and nrss to the criterion is very high-

ly significant: the more developed seeds, the fewer spoiled

seeds and the more escape holes (-0.402289 * -0.421838

= (0.1697). Overall, the indirect effects indicate a decrease

of the negative influence from the direct paths to the cri-

terion.

©ZFMK

Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 61

*** -0.267961

Variables

nreh number of escape holes

nrla number of larvae

nrpu number of pupae

nrne number of not emerged adults

nrds number of fully developed seeds.

nrus number of undeveloped seeds

nrss number of spoiled seeds

Fig. 3. Path diagram 2 showing 6 predictor variables and a residual effecting on criterion (var 7, nrds, plant related variate, num-

ber of fully developed seeds per seed capsule), R? = 0.545329. The path diagram explains 54.53% of the variance of the criterion.

The significance of paths was determined by random permutational test, 10000 iterations each, Bonferroni Correction.

Path Analysis II (Fig. 3): The second path analysis, using

the same variables as in the first analysis, was calculated

to determine the influence of variables 3—6, 8 and 9 on

variable 7, nrds, number of fully developed seeds per cap-

sule, which is the most important plant related factor. Since

variable 7 may be an indicator of fitness for the plant, it

was subjected to a multivariate path analysis.

There are 4 direct paths from the predictors to the crite-

rion. The first direct path proceeds from var 3, nreh, to

the criterion and is negative and very highly significant:

the more escape holes, the less fully developed seeds per

capsule. The second direct path from var 4, nrla, to the

criterion is also negative and very highly significant: the

more larvae, the fewer well developed seeds. The third di-

rect path, from var 8, nrus, to the criterion is negative and

very highly significant: the more undeveloped seeds, the

fewer developed seeds per capsule. The fourth direct path,

from var 9, nrss, to the criterion is also negative and very

highly significant: the more spoiled seeds, the fewer well

developed seeds.

Bonn zoological Bulletin 57 (1): 55—64

There are several indirect effects between the predictors

and the criterion. The indirect effect proceeds from var 3

and var 8 to the criterion and is very highly significant:

the more escape holes, the less undeveloped seeds and the

more fully developed seeds (-0.257581 * -0.387095 =

0.09971). The indirect effect between nreh and the crite-

rion, via nrss, is very highly significant: the more escape

holes, the fewer spoiled seeds and the more fully devel-

oped seeds (-0.267961 * -0.407705 = 0.10925). Both paths

show a decrease of the negative influence of escape holes

on the criterion. Another indirect effect from nrne, via nr-

pu, via nrla to the criterion is very highly significant: the

more not emerged adults, the more pupae and the more

larvae, the fewer well developed seeds will be found

(0.212016 * 0.538041 * -0.256493 = -029259). This indi-

rect effect shows a negative influence on the criterion.

In summary, both the PCA and the path analyses explain

animal - plant interactions; path diagram | focuses on the

animal, while path diagram 2 centers on the plant.

OZFMK

62 Gertha Wilhelm et al.

DISCUSSION

The relationships between insects and plants are of utmost

importance for terrestrial global ecology. Many of these

relationships involve specialized herbivorous insects and

their host plants on which the females oviposit and on

which the larvae feed (Schoonhoven et al. 1998). The

choice of an oviposition site is instrumental for the fitness

of many weevils (Messina 2004). In a case study approach

we investigated fitness components of the interactions be-

tween the weevil, R. longirostre and its host plant, A.

rosea.

The life cycles of the weevil and its host plant must be

considered in connection with each other. With the help

of multivariate statistics it was possible to isolate sever-

al main factors which contribute toward the survival of

the host plant, as well as the successful development of

the weevil. The lengthy inflorescence of A. rosea may con-

tain of more than 30 buds in different development stages

ranging from fully developed buds at the bottom, which

will blossom soon, to immature tiny buds at the top. Host

plant preferences of the female weevils for oviposition

sites are buds located in the middle of the inflorescence.

Exactly these offsetting development stages of the buds

are reflected in the PCA (Table 2). In summary, the PCA

provides an overview of the interaction system between

the plant and the weevil.

Factor axis 1, which is the most important one, is inter-

preted as the axis which reflects successful seed develop-

ment: The variate of final production outcome in the plant,

i.e. nrds, number of developed seeds, is contrasted to the

counteracting variables of numbers of undeveloped seeds,

nrus, and spoiled seeds, nrss, and indicators of weevil de-

velopment, nrla, nrpu and nrne. The axis is to be read as

follows: the more developed seeds, the less dead larvae,

pupae and not emerged adults in the capsules but also the

less undeveloped and spoiled seeds. In contrast to this in-

terpretation, axis | could be read as: the more dead lar-

vae, pupae and not emerged adults inside the capsules and

the more undeveloped and spoiled seeds, the less devel-

oped seeds. Note that the variable of plant final success,

nrds, is exclusively represented on axis | but not the vari-

able of the animal final success, nreh. Consequently, we

assigned to axis | the abbreviation SUPLA for “success-

ful plant” (i.e. seed) production.

The next most important axis, factor axis 2, is to be in-

terpreted as mainly reflecting weevil development and pro-

duction. The variate of final production outcome in the an-

imal, i.e. nreh, number of escape holes, is combined in

variation with the variates of animal development, i.e. nr-

la, nrpu, and nrne. Whereas the first variate, nreh, signi-

fies production success, the remaining variates nrla, nrpu

Bonn zoological Bulletin 57 (1): 55—64

and nrne stand for the unsuccessful weevil development

in the host plant. However, all the animal related variates

are contrasted to plant variates which stand for unsuccess-

ful seed development, i.e. nrus and nrss — undeveloped and

spoiled seeds. The axis loading read as: The less undevel-

oped and spoiled seeds, the more escape holes but also the

more dead larvae, pupae and not emerged adults. Note that

the variate of final production outcome in the plant, nrds,

is not significantly represented on this axis. Consequent-

ly, we designate this axis SUAN, “successful animal” pro-

duction, as a main infestation axis. Highly infested cap-

sules were located in the middle of the plant’s inflores-

cence. Early infestation of seed capsules guarantees high

reproductive success for the weevil as shown by the high

proportion of open escape holes. In cases where a high

number of seeds were infested, the number of undevel-

oped and spoiled seeds was very low.

Factor axis 3 reflects the mutual interaction of weevils and

plant production. In one group of variates the final pro-

duction outcome of the animal, nreh, and the pre-devel-

opmental stage to that final outcome, i.e. nrne, are com-

bined with the number of undeveloped seeds. In contrast,

in the second group of variates, the final production out-

come of the plant, nrds, is combined with the developmen-

tal variates of the weevil, nrla and nrpu. The axis load-

ings can be read in two different ways, both immediate-

ly indicating a competitive race between animal and plant

to maximize their fitness (as shown for a seed predatory

weevil and its host plant, Tojyu, Sota 2006). On the one

hand, the more escape holes, as well as not emerged adults

and undeveloped seeds in capsules, infer fewer developed

seeds, as well as larvae and pupae. On the other hand, the

more developed seeds, as well as larvae and pupae, infer

fewer escape holes, not emerged adults and undeveloped

seeds. There are two counter acting trends: One, early in-

festation by the weevil which is characterized by a high

number of open escape holes and not emerged adults to-

gether with a high number of undeveloped seeds. Since

larvae only infest well developed seeds, few seeds remain

for plant development. The second trend concerns late in-

festation and incomplete weevil development. It is char-

acterized by a high number of larvae and pupae together

with a high number of developed seeds. The abbreviation

SUMIX (successful animal and plant production) repre-

sents the integration of the two trends.

Factor axis 4 can be interpreted as late infestation. Both

variates, nrds and nreh, are highly significant and contrast

each other. In the one group of variates, nrds is combined

with nrss and nrne. In another group, nreh is combined

with nrla and nrus. The variate loadings are interpreted as

the following: many developed seeds and many spoiled

seeds infer few undeveloped seeds in capsules, while the

presence of fewer escape holes and fewer larvae infer

©ZFMK

Fitness components in the relationship between Rhopalapion longirostre and Alcea rosea 63

fon)

more not emerged adults. A high number of spoiled seeds

indicates that adverse weather conditions, mostly in late

summer, could have influenced negatively both seed and

weevil development, and in particular, the escape of adult

weevils. We abbreviate this axis LIFA, “late infestation

axis”.

Factor axis 5 is an axis representing damaged seeds and

late developmental stages of the weevils. The variate, nrss,

contrasts with nrus and nrne. The loadings are interpret-

ed as follows: The presence of many spoiled seeds infers

fewer undeveloped seeds and fewer not emerged adults,

and vice versa. This axis is termed DASA, “damaged seed

and animal axis”.

The main important axes of the PCA provide a compre-

hensive explanation about the complex system of inter-

action between animal and plant.

The influences and trends of both path diagrams must be

seen in the light of direct paths and indirect effects (see

Figs. 2 and 3). Our null-hypothesis was falsified by the

significant correspondence between the life cycle of host

plant and weevil. In path diagram 1, all direct paths (with-

out exceptions) from the different variates to the criteri-

on have strong negative influences. The sequence of in-

direct effects between the number of larvae > number of

well developed seeds > criterion, as well as the indirect

influence nrla > nrpu > nrne > nrds > criterion, nreh,

are very important. As larval feeding habits are highly con-

servative some of these habits are apparently irreversible,

e.g. feeding on seeds (Marvaldi et al. 2002). Larvae of the

last developmental stage choose well developed seeds to

enter, to consume and to prepare the seed chamber for pu-

pation and future escape. The sequence of indirect effects,

nrds > nrus > criterion, or nrds > nrss > criterion, are

also very important. For the performance of a larva, it is

vital to find a well developed seed instead of an undevel-

oped or spoiled seed.

In path diagram 2, the 4 direct paths (without exceptions)

have also strong negative influences on the criterion nrds.

The indirect effects are advantageous for both, plant and

animal production. The sequence, nrne > nrpu > nrla >

criterion, indicates that the high number of infested seeds

are based on a high number of formerly well developed

seeds. The sequences, nreh > nrus > criterion, nrds, and

nreh > nrss ~ nrds, infer that high numbers of escape

holes and few undeveloped or spoiled seeds increase the

number of well developed seeds. A plant in a healthy con-

dition guarantees the performance of the weevils. The time

of infestation is crucial for the weevils. Our analyses re-

veal that an infestation occurring late in the season in-

volves a high likelihood of poor plant condition, as shown

in the indirect effect nrpu > nrss > nreh.

Bonn zoological Bulletin 57 (1): 55—64

In conclusion, we address the initial questions posed in

the introduction.

1) What are the optimal conditions for the reproduction

of weevils? The most important optimal condition for wee-

vil reproduction is the timely presence of many well de-

veloped seeds.

2) What are the optimal conditions for the successful de-

velopment of plants? The optimal conditions for the suc-

cessful development of plants are few undeveloped and

few spoiled seeds.

3) Is the contribution of larvae to the overall reproductive

success of the weevils important?

Yes, in both path diagrams, the animal related variate, 1.e.

number of larvae, has a highly significant direct path to

the first criterion, number of escape holes, as well as to

the second criterion, well developed seeds. Obviously lar-

vae benefit from capsules with a very high number of well

developed seeds. Since the number of larvae has an ex-

clusively direct impact on both criteria, this influence is

interpreted as a high larval contribution to the reproduc-

tive success of R. longirostre.

Our results strongly support the view of an intricate rela-

tionship between the life cycles of A. rosea and R. lon-

girostre. The plants’ period of bud development lasts from

April to October. The weevils utilize a window within this

range to fulfill their development, beginning in May af-

ter hibernation in the soil and ending in September with

the completion of their life cycle and the emergence of the

next adult generation from the seed chambers. The time

periods before and after weevil activity ensure enough op-

portunity for successful production of seeds.

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Kendall/Hunt, Dubugue, Iowa

Toju H, Sota T (2006) Adaptive divergence of scaling relation-

ships mediates the arms race between a weevil and its host

plant. Biology Letters 2: 539-542

Wilhelm G (2004) Die Lebensgeschichte von Rhopalapien lon-

girostre (Olivier). Diploma dissertation, Department of Evo-

lutionary Biology, University of Vienna, 105 pp

Volume 2.

APPENDIX

Poisson regression analysis

Dependent variate: nreh

Regression coefficients (predictors):

beta- 0 (constant): 4.632211 < 0.0001

beta- | (nrla): -0.170142 0.0002 ***

beta- 2 (nrpu): -0.040228 0.4612 ns.

beta- 3 (nrne): 0.040783 0.5624 ns.

beta- 4 (nrds): -0.405291 < 0.0001 ***

beta- 5 (nrus): -0.267374 < 0.0001 ***

beta- 6 (nrss): -0.258430 < 0.0001 ***

global model, deviance: 530.341739, P-value = < 0.0001, ***

The significance was tested by random permutational tests

(10000 iterations each).

Dependent variate: nrds

Regression coefficients (predictors):

beta- 0 (constant): 3.797212 < 0.0001

beta- 1 (nreh): -0.166088 < 0.0001 ***

beta- 2 (nrla): -0.098854 0.0072 **

beta- 3 (nrpu): -0.061230 0.1904 ns.

beta- 4 (nrne): -0.079595 0.1760 ns.

beta- 5 (nrus): -0.158951 <0.0001 ***

beta- 6 (nrss): -0.170892 <0.0001 ***

global model, deviance: 340.101076, P-value = < 0.0001 ***

The significance was tested by random permutational tests

(10000 iterations each).

Received: 10.10.2008

Accepted: 25.02.2009

Corresponding editor: M. Schmitt

©ZFMK

Bonn zoological Bulletin | Volume 57 | Issue | | pp. 65-73

Bonn, April 2010

Life-history of Gnopharmia kasrunensis Wehrli, 1939

and G. colchidaria Lederer, 1870 (Geometridae, Ennominae)

and their distribution in Iran, with first host-plant records for the genus

Hossein Rajaei Shoorcheh

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

E-mail: rajaei@daad-alumni.de

Abstract. For the first time, observations on the life-history of the little known species Gnopharmia kasrunensis Wehrli,

1939 and G. colchidaria Lederer, 1870 are reported; eggs, larvae, pupae, adults and habitats are illustrated. Suitable dif-

ferential features for a correct identification of both Iranian Gnopharmia species as well as a distribution map are pro-

vided.

Zusammenfassung. Erstmals wird tiber Beobachtungen zur Biologie der wenig bekannten Arten Gnopharmia kasrunen-

sis Wehrli, 1939 und G. colchidaria Lederer, 1870 berichtet. Eier, Larven, Puppen, Imagines und ihre Lebensraéume wer-

den beschrieben. Fiir die korrekte Bestimmung der beiden iranischen Gnopharmia-Arten werden geeignete differential-

diagnostische Merkmale sowie eine Verbreitungskarte vorgelegt.

Key words. Gnopharmia kasrunensis, colchidaria, biology, distribution, Iran

INTRODUCTION

The genus Gnopharmia Staudinger, 1892 belongs to the

tribe Macariini (Ennominae) (Wehrli, 1953: 565; Scoble

& Kriiger 2002) and consists of several species which are

difficult to determine by external means only. Parsons et

al. (1999) list thirteen species in this genus. At present,

Gnopharmia is in the process of taxonomic revision by

the author. By discovery of one new species and five syn-

onymies, the number of valid species in the genus has been

reduced to nine. All species occur in the Middle East and

Asia Minor, their geographic range extending from

Turkey to Pakistan. Five of the species occur in Iran. Our

present knowledge on the biology and distribution of

Gnopharmia species is very poor, partly due to difficul-

ties of their correct identification. In the original descrip-

tions of species in this genus (e.g. Ebert 1965, Erschoff

1874, Lederer 1870, Staudinger 1892, Wehrli 1938, Wehrli

1939, Wehrli 1941, Wiltshire 1967, Wiltshire 1970) no

host plant records or notes about their biology are given.

Prout (1915) listed three Gnopharmia species and six sub-

species, and Wehrli (1953) mentioned seven species (and

11 subspecies). Both described the external features and

Wehrli also characters of their genitalia, but they did not

mention host plants or larval stages. The host-plant index

published by The Natural History Museum, London

(HOSTS, 2007), also does not record any host plant for

species of the genus Gnopharmia.

After a number of unsuccessful breeding experiments with

Prunus spp. and other Rosaceae species by various lepi-

dopterists, the first correct suggestion about the host plant

of Gnopharmia was made by Bernd Miller (Berlin; pers.

comm.), who assumed the wild Almond which occurs

abundant in the habitats where Gnopharmia species oc-

curs (Prunus, subgenus Amygdalus; Rosaceae). The first

breeding of a Gnopharmia was made by Robert Trusch

(Karlsruhe; pers. comm.) in 2008 with material from East-

Iran (Prov. Ostan-e Khorasan, Birjand, Kuh-e Mirza Arab,

2.040 m a.s.1); larvae were fed with leafs of wild Almond

shrubs from the field in Iran, but the experiment had to

be finished because of the lack plant material in Germany;

the species of the reared larvae remained undetermined

During May and July 2009, the author collected females

of G. kasrunensis (Fig. 13) and G. colchidaria (Fig. 14)

at several localities in Iran. The present paper illustrates

the larval stages of both species and gives an overview of

their distribution.

MATERIAL AND METHOD

Females of Gnopharmia species have been collected at six

Localities (Table 1) by using a light trap. For laying eggs,

66 Hossein Rajaei Shoorcheh

they were caged inside small plastic boxes with tissue pa-

per, eventually adding suitable plant material.

In Tole-Heidari valley (province Fars, south of Jahrom,

N 28°28’56” E 053°23’09”, altitude 1112 m a.s.].) May

15, 2009, one female of G. kasrunensis has been collect-

ed (culture A, table 1). This habitat is very dry and there

are scattered bushes of different plant species. Prunus ly-

cioides (Rosaceae; det. H. Akhani and M. Yazbek) as a

possible host plant is common. Three females of G. kas-

runensis were found in Ras-Kuh (province Kerman, road

Baft-Sirjan, 2 km after Baft, N 29°17°27” E 056°35°37”,

altitude 2540 m a.s.l.) May 20 to May 21, 2009 (cultures

B, C and D, table 1). This locality near Ras-Kuh village

has a southeast-facing aspect (Fig. 1). The dominant plant

species in the area is Prunus scoparia (det. H. Akhani and

M. Yazbek) (Fig. 3 and 4). Material of this plant species

has been added to the plastic boxes containing females.

Two females of G. colchidaria have been collected at the

mountains near the village Ab-Asemani, (province Fars,

road from Estahban to Sarvestan, 20 km to Sarvestan, N

29°05’51” E 053°26712”, 1890 m a.s.l., May 22, 2009)

(cultures E and F, table 1). This biotope has a northern ex-

posure. Prunus scoparia was common here too, but there

also were wild Pistacia trees (Anacardiaceae), many bush-

es of Astragalus (Fabaceae) and Echinops (Asteraceae),

and several Umbelliferae spp. (Umbelliferae) (Fig. 2).

One female of G. colchidaria has been collected at Osh-

torankuh (province Lorestan, between Dorud and Gahar

lake, near Cheshmeh-Khorram, N 33°22’41” E 049°11713

females of G. colchidaria were found at Gardaneh-Gar-

rin (province Hamadan, 25 km to Nahavand from

Noorabad, N 34°02’48” E 048°20731”, 2135 ma.s.1., Jun

25, 2009 (cultures H—L table 1) and at Khani-Sefid in Gar-

daneh-Khan near the border of Iran-Iraq (province Ko-

rdestan, 10 km from Saghez to Baneh, N 36°04713” E

045°59’31”, altitude 1976 ma.s.l., June 26, 2009) one fe-

male of G. colchidaria has been collected (culture M, table

1). No eggs or larvae have been observed in nature. Pu-

pation of G. kasrunensis larvae (culture C) took place in

the field (Iran), that of G. colchidaria (culture F) in the

laboratory in Karlsruhe-Germany. In the field, the study

of egg-morphology was done using a hand lens.

For this study the species have been identified by using

male genitalia of freshly emerged adults. Identifying fe-

males by their genitalia 1s still problematic, as differences

between species are minute and not fully understood at

present. In some cases (e.g. G. colchidaria) only a female

emerged and for identification genitalia dissection of si-

multaneously collected males were used. There are clear

characters in the male genitalia, especially in the aedea-

gus, that can be used to separate the species. The aedea-

gus of G. kasrunensis 1s thicker and shorter (length 1.2—1.5

mm) than that of G. colchidaria (length 1.4—1.8 mm) and

bears a group of 2—6 long, rod-like, subapical spines

(length up to 0.3 mm) with rounded tip (Figs 24, 28). In

contrast, the aedeagus of G. colchidaria has a subapical

group of short, more or less cone-shaped spines (max1-

mum 0.15 mm in length), 1—3 of them situated more dis-

tally and 2-3 (rarely up to 5) more basally (Figs 25, 29).

The tips of the ‘octavals’ (a pair of distal processes at ster-

nite 8) are more strongly curved in G. kasrunensis (Figs

22, 26), than in G. colchidaria (Figs 23, 27). The geni-

talia capsules of both species are similar in size and shape,

Table 1. Species, date and number of collected females and their eggs, larvae and pupa, and dates of hatching and pupation, ac-

cording to cultures.

Code of Species Date of Date of N. eggs Date of N. larvae Dateof N. pupa

culture collecting laying egg hatching pupation

A (loc. 1) G. kasrunensis 15.5.09 17.5.09 27 21.-22.5.09 21 — 0

B (loc. 2) G. kasrunensis 20.5.09 23.—24.5.09 37 27.-28.5.09 28 = 0

C (loc. 2) G. kasrunensis 20.5.09 23.—24.5.09 14 27.-28.5.09 12 25.6.09 1

D (loc. 2) G. kasrunensis 21.5.09 24.-25.5.09 28 28.-29.5.09 22 = 0

E (loc. 3) G. colchidaria 22.5.09 28.—29.5.09 24 30.—31.6.09 17 — 0

F (loc. 3) G. colchidaria 22.5.09 28.-29.5.09 33 30.-31.6.09 21 2.7.09 1

G (loc. 4) G. colchidaria 22.6.09 25.6.09 18 29.-30.6.09 16 — 0

H (loc. 5) G. colchidaria 25.6.09 27.-28.09 16 1.-2.7.09 16 _ 0

I (loc. 5) G. colchidaria 25.6.09 27.28.09 45 1.—2.7.09 36 — 0

J (loc. 5) G. colchidaria 25.6.09 27.28.09 42 ].—2.7.09 38 - 0

K (loc. 5) G. colchidaria 25.6.09 27.28.09 U7 1.—2.7.09 14 — 0

L (loc. 5) G. colchidaria 25.6.09 27.28.09 40 1.—2.7.09 36 - 0

M(loc. 6) G. colchidaria 26.6.09 28.-29.6.09 9 2.-3.7.09 7 = 0

Bonn zoological Bulletin 57 (1): 65—73 O©OZFMK

Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 67

Plate 1. Figs 1-2: Biotopes of Gnopharmia species: 1, G. kasrunensis (Ras-Kuh, cultures B, C and D); 2, G. colchidaria (Ab-

Asemani village, cultures E and f); Figs 3-4: host-plant: 3, tree of Prunus scoparia; 4, leaves and fruits of Prunus scoparia; Fig.

5: eggs of G. kasrunensis (culture C); Figs 6-8: larvae of G. kasrunensis: 6, L2; 7, L4; 8, L3; Figs 9-11: larvae of G. colchidaria:

ONES OS WD tle:

Bonn zoological Bulletin 57 (1): 65-73 ©ZFMK

68 Hossein Rajaei Shoorcheh

Plate 2. Figs 12-14: Adults of Gnopharmia: 12, G. colchidaria (female, freshly emerged; culture F); 13, G. kasrunensis (holo-

type, male); 14, G. colchidaria (male) (scale 1 cm); Figs 15-16: Pupa of G. colchidaria: 15, ventral view, 16, lateral view (scale

2mm); Fig. 17: Pupa of G. kasrunensis, ventral view (one day after pupation); Fig. 18: Pupa of G. kasrunensis, posterior part (ma-

le, ventral view; Cr: cremaster); Fig. 19: id., pupa of G. colchidaria, female (scale 1mm); Fig. 20: floricomous setae (distal part)

on female ovipositor (scale on figure); Fig. 21: Setal map of G. colchidaria and G. kasrunensis larvae.

Bonn zoological Bulletin 57 (1): 65—73 OZFMK

Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 69

but the basal process of the sacculus is distinctly longer

in G. colchidaria. In G. kasrunensis both of these projec-

tions seems to be equal in length (Figs 30, 31).

For SEM photography male genitalia have been dehydrat-

ed in 70%—96% and 100% ethanol. For drying with

HMDS (1, 1, 1, 3, 3, 3-Hexamethyldisilazan) the HMDS-

protocol (Oshel 1997) was used. According to this proto-

col the genitalia have been transferred from 100%

ethanol to a 1:1 ethanol-HMDS mixture and then to 100%

HMDS. The specimens were dried by evaporation of

HMDS.

SEM photos were made using a Hitachi S-2460N SEM

at Zoologisches Forschungsmuseum Alexander Koenig.

RESULTS

Rearing report and field observations

The female of culture-A (G. kasrunensis) laid eggs in a

plastic box containing leaves of Prunus lycioides. Most

larvae hatched from these eggs, but died because of lack

of fresh food plants. Leaves of P. lycioides carried with

us had dried out and no P. /ycioides or another possible

host plant was available. Females of cultures B—F (G. kas-

runensis) laid eggs (Prunus scoparia was added to the

containers) just two to three days after collecting and two

days later, L1-larvae started to feed successfully.

Despite attempting to provide relatively stable conditions

to the larvae, many of them died in different stages. Of

79 Ll-larvae of G. kasrunensis in cultures B, C and D,

just 42 survived to L2, and only 18 of them moulted again

(after 7-8 days). L3-larvae were feeding almost continu-

ously, but they were more active in the morning and in

the afternoon. Larvae continued to perish during this stage

as well. 6—8 days later just 7 larvae moulted to L4. They

fed again for 5—6 days, but during this time all but one

died. Fully grown (25.6.2009) the last remaining larva of

this species (culture C) stopped feeding and rested under

pieces of tissue paper in its container, then pupated (see

Fig. 17), without spinning or producing a cocoon.

L1 stage of G. colchidaria took 7—9 days. Many of the

larvae again died later during several instars. Of 38 L1-

larvae (cultures E and F) just 24 moulted to L2. 5 larvae

died in this instar. L3-stage took 7-8 days and of 19 lar-

vae just three reached L4. The feeding period of the last

instar is longer than that of the last species (9 days). Ful-

ly grown, just one larva pupated (Figs 15, 16).

At a temperature of 25—28° C, a male of G. kasrunensis

emerged after 18 days (July 12, 2009; dissected). At a

Bonn zoological Bulletin 57 (1): 65-73

same temperature, one female of G. colchidaria emerged

from pupa after 16 days (July 18, 2009; Fig. 12).

All seven females of the cultures G—M (collected from

Lorestan, Hamadan and Kordestan provinces) laid eggs

(a total of 187 eggs) which were brought to Germany, in

order to find and to test suitable food-plants there. 163 L1-

larvae hatched but none of the plants offered was accept-

ed and all larvae died.

Host-plants

Prunus scoparia has been offered to the L1-larvae of G.

kasrunensis and G. colchidaria (cultures B—F) and was

readily accepted by all of them. This plant may be con-

sidered as the main food resource for both, G. kasrunen-

sis and G. colchidaria, also under natural conditions.

To the L4-larva of G. colchidaria (cultures E and F), feed-

ing on P. scoparia before, I offered two other Prunus

species (Prunus haussknechtii C. K. Schneid. and Prunus

lycioides (det. H. Akhani and M. Yazbek), but no feeding

marks were found in either of them.

To Ll-larvae of G. kasrunensis from Tole-Heidari (cul-

ture A) the common almond (Prunus dulcis (Mill.)

D.A.Webb) and later Prunus haussknechtii C. K. Schneid.

have been offered, but none of them was accepted. Prunus

scoparia was not available at that time.

All larvae of the cultures G—M have been brought to Ger-

many. It took 2 days to find a plant of Prunus dulcis

(Miller) D.A. Webb. During this period I offered Cratae-

gus monogyna Jacq., but it was not accepted by the lar-

vae and more than half of them died. The others died in

the next days because they also did not accept Prunus dul-

cis.

Morphology of preimaginal stages

Ovum

Ovum: Elliptical, surface rough, green in colour (just af-

ter deposition), turning red-ochre after one or two days

(Fig. 5). Most of them were deposited separately in a ver-

tical, upright position, but some were laid in clusters as

shown in Fig. 5.

Under warm conditions, L1-larvae developed very quick-

ly and emerged just after 4—5 days (table 1).

There is no significant difference in the shape or the struc-

ture between the eggs of these two species.

OZFMK

70 Hossein Rajaei Shoorcheh

Plate 3. Figs 22-23: SEM photos of sternite A8, with ‘octavals’: 22, G. kasrunensis; 23, G. colchidaria; Figs 24-25: SEM pho-

tos of apical part of aedeagus: 24, G. kasrunensis; 25, G. colchidaria (scale on figures); Figs 26-27: Sternite A8: 26, G. kasrunen-

sis (holotype, male); 27, G. colchidaria (male); Figs 28-31: male genitalia: 28, G. kasrunensis (aedeagus); 29, G. colchidaria (ae-

deagus); 30, G. colchidaria (genital capsule); 31, G. kasrunensis (genital capsule). Scale 1 mm.

Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 71

Larval stage

G. kasrunensis Wehrli, 1939

L1: 1.42.1 mm (5 days after haching) in length, with two

longitudinal pale-brown bands on green background, start-

ing from the posterior end of the medio-dorsal area and

continuing to the head. Head and end of the body are

brown. A longitudinal brown band, similar to the dorsal

band, runs ventrally from posterior end to the head. The

thoracic legs and abdominal prolegs are dark brown.

L2: (Fig. 6): length 6.2—-7.00 mm (11 days after hatching),

brown medio-dorsal bands change (in compare to L1)

from brown to more green, similar to the ground colour,

but at the end of L2 the rest of brown bands are distinct-

ly visible on two-thirds posterior of body. The brown ven-

tral band vanishes toward head. In this stage, there are two

dorsal processes at the base of the D2 seta dorsally on seg-

ment A2 (see Fig. 21). The tips of these processes are dark

brown or black.

L3 (Fig. 8): 9.6—10.5 mm (20 days after hatching), exhibits

nearly the same colour patterns as found in the fully grown

larvae. The dorsal area is completely green with medio-

dorsal brown bands (from A10 to A6). Ventrally, the green

colour is extended, but on A6—10 there are still soft brown

colour band. During L3, the colour of the dorsal process-

es at the base of the D2 seta (of the A2 segment) changes

from dark brown or black to brown.

L4 (Fig. 7): 19.2—25.8 mm (28 days after hatching), com-

pletely green in colour, with a longitudinal medio-dorsal

brown bands running from the A10 to the A6 segment. The

dorsal processes of the A2 segment are brown, but in two

cases, changed to green.

G. colchidaria Lederer, 1870

L1: 1.2—2.2 mm (7 days after hatching), coloration very

similar to G. kasrunensis, but here longitudinal dorso-me-

dial bands start from the posterior end and continue along

two-thirds of the body dorsally (from A10 to A4—5).

L2: (Fig. 9): 6.5—7.1 mm (12 days after hatching), brown

dorso-medial bands continue to the head, the ventral band

extends slightly to the lateral. The two dorsal processes

of A2 are clearly dark brown to black in colour (like in

G. kasrunensis).

L3: (Fig. 10) 9.7-10.7 mm (20 days after hatching), com-

pletely different in comparison with G. kasrunensis: dor-

sal and ventral longitudinal brown bands extend lateral-

ly and the green colour is reduced to the lateral area. Black

dorsal processes on the A2 segment are clearly visible.

Bonn zoological Bulletin 57 (1): 65—73

L4: (Fig. 11): 19.2-26.1 mm (26 days after hatching), pre-

dominantly brown, with some yellow-green colour later-

ally and dorsally. The A2 dorsal processes are brown, like

the surrounding area.

The position of the primary setae of the L4- larvae of

both species does not exhibit any differences. (see setal

map, Fig. 21). For nomination of seta used McGuffin

(1977) and Stehr (2005).

Pupa

Pupae of both species are obviously very similar in size

and colour. (Figs 15-19). They are slender, about 12.8—13

mm in length, the head section rather acute, almost trian-

gular; the elongated cremaster (length 0.6 mm) bears two

terminal spines of different length (right spine broken in

fig. 19). Until the second day after pupation the colour re-

mains green, and then changes to brown. The pupal exu-

via is mid-brown, too. More material is needed in order

to compare the pupal characters of both species thorough-

ly.

Distribution (Fig. 32). According to our present knowl-

edge, G. kasrunensis and G. colchidaria occur sympatri-

cally in most parts of the Zagros Mountains. G. kasrunen-

sis has been described from Kazeroun (Fars prov., Iran)

and is distributed in the central and eastern Zagros with

some small populations in western Zagros and central Al-

borz. Only two specimens of this species were collected

at Sayhakil in Musandam (Oman). G. colchidaria occurs

in nearly all parts of Zagros, furthermore there are popu-

lations in south-eastern Iran, and in the Alborz (vic.

Tehran, Qazvin, Zanjan, and Gilan). The species has been

described from Helenendorf (East Azerbaijan, Transcau-

casia; present name: Xanlar), and there are additional re-

ports from Armenia (ssp. melanotaenia Wehrli, 1938) and

other regions of S. Caucasus. The map is based on mate-

rial present in the collections studied. Probably these two

species have a wider distribution, but the author so far had

no access to further material proving this.

Bionomic data. According to the analyzed label data, de-

rived from several collections, G. kasrunensis has been

collected between the second week of February and the

second week of September. The flight period of G. colchi-

daria dates from early March to late October. G. kasrunen-

sis flies at altitudes from 200 up to 2200 m, G. colchidaria

from sea level to 2800 m. Both species probably are (at

least) bivoltine in most places, but may be univoltine in

the North of Iran and at higher elevations. Overwintering

stage obviously is the pupa.

©ZFMK

72 Hossein Rajaei Shoorcheh

48° S26

Caspian Sea

56° 60°

36°

32°

Kerman

Ei: A $ ifn

iraz a

A U-~e e@ ie Br :

e ew

28° : e.

Bandar-e Abbas

Persian Gulf

1000-2000 m

> 2000 m

® G.colchidaria

A G.kasrunensis

Fig 32. Distribution map of G. kasrunensis and G. colchidaria in Iran.

DISCUSSION

Despite intensive search on many Prunus shrubs in a num-

ber of Iranian localities for immature stages of

Gnopharmia, no eggs, larvae or pupae were found in the

nature. Of course, it is often difficult to find eggs of moths

in the field, but in case of Gnopharmia these difficulties

may also be related to the existence of a so-called “flori-

comus” on the ovipositor of Gnopharmia females. Numer-

ous specialized (“scoop-shaped”) setae (Fig. 20) on the

papillae anales probably enable the females to scratch par-

ticles from the surface of the bark to cover and camou-

flage the eggs. This behaviour has been observed in the

W. European Ennomine species Theria primaria Haworth,

1809 (Stokoe 1948: 186, pl. 39:1), females of which have

a very similar ovipositor with almost identical, specialized

Bonn zoological Bulletin 57 (1): 65-73

setae. This hypothesis is supported by the fact that the

papillae anales of almost all females studied are spoiled

with dust or bark-particles retained near the setal bases

(Scoble & Kriiger, 2002, D. Stiining, Bonn, pers. comm.,

unpublished).

Prunus scoparia as the most probable natural host plant

of G. kasrunensis and G. colchidaria is well distributed

in many parts of the Zagros Mountains. During the last

years this plant has also been intensively grown by local

people and became more widely distributed, as “bitter al-

monds’ became more frequently used by the pharmaco-

logical industry in Iran. G. kasrunensis and G. colchidaria

fly sympatrically at many localities in Iran (Fig. 32). Lar-

©ZFMK

Life-history of Gnopharmia kasrunensis and G. colchidaria and their distribution in Iran 73

vae of both species are very similar morphologically and

the young larvae (L1, L2) are also without clear differ-

ences in colour and pattern. However, grown larvae (L3,

L4) can be separated.

The reason for the high mortality of the larvae during all

stages is not clear to us. Alteration of climate in the ve-

hicle, low quality of the food plant, obstruction by sun-

light or artificially damp environment (wet paper towels

were provided to keep the food fresh) could have been the

reason for bacterial or viral infection which killed the lar-

vae.

Further studies on the biology of this interesting moth

genus will be carried out during the 2010 collecting sea-

son.

Acknowledgments. I deeply thank Axel Hofmann (Breisach)

and Dr. Jorg-Uwe Meineke (Kippenheim) for their helpful ad-

vice during our common field work, and express my sincere

thanks to Prof. Dr. Hossein Akhani (Tehran) and Mariana Yazbek

(Lebanon) for the determination of host plants. Furthermore, I

do not forget valuable comments by Dr. Robert Trusch and Giin-

ter Ebert (Karlsruhe), Dr. Dieter Stiining (Bonn) and Dr. Axel

Hausmann (Munich). I thank Lily Wescott and Tom Swinehart

(Bonn) for proofreading of the first draft, Stefan Scharf (Karl-

sruhe) for preparing the digital map of Middle-East, Ulla Kreutz

(Meckenheim) for processing of the digital images, Bernd Miiller

(Berlin), Michael Falkenberg (Karlsruhe) and Maryam Bo-

roomandi (Bonn) for their friendly help during different steps

of this project. Special thanks go to Dr. Montazami, Alireza

Naderi, Mohammad-Ebrahim Sehhati and other colleagues of

Iran Department of Environment for their collaboration during

our 2009 excursion and also colleagues of Hayk Mirzayans In-

sect Museum in the Iranian Research Institute of Plant Protec-

tion (HMIM, Tehran) for their friendly collaboration in the proj-

ect Gnopharmia. | am grateful to Hamid Valipoor (Ghoochan)

and Mahdi Nasiri (Dorud) for their aid in the field trip of 2009

in Iran, to Professor Dr. J. W. Wagele (ZFMK, Bonn) for his pro-

motion and DAAD (Deutscher Akademischer Austausch Dienst)

for the financial support of this project.

REFERENCES

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Beitrage zur Naturkunde 142: 1-32, Pl. 14

Erschoff NG (1874) Cheshuekrylyja (Lepidoptera). — In:

Fedchenko AP: Puteshestvie vy Turkestan [Journey to

Turkestan], vol. 2 Zoogeograficheskaja izsledovanija part 5,

p. E-VI, 1-127, Pl. 1-6

HOSTS (a Database of the World’s Lepidopteran Hostplants,

hosted by Natural History Museum, London 2007):

Bonn zoological Bulletin 57 (1): 65—73

http://www.nhm.ac.uk/jdsml/research-curation/research/proj-

ects/hostplants/index.dsml

Lederer J (1870) Contributions a La Faune des Lépidoptéres de

la Transcaucasie. Annales de la Société Entomologique de

Belgique. 13: 39, 48, pl. 2, Fig. 1

McGuffin WC (1977) Guide to the Geometridae of Canada

(Lepidoptera), II subfamily Ennominae 2. Memoirs of the

Entomological Society of Canada. No. 101, Fig. 152

Oshel Ph (1997) HMDS and Specimen Drying for SEM.

Microscopy today 5 (4):16

Parsons MS, Scoble MJ, Honey HR, Pitkin LM, Pitkin RB

(1999) The catalogue. In: Scoble MJ, ed. Geometrid Moths

of the world: a catalogue (Lepidoptera, Geometridae).

Collingwood: CSIRO Publishing. 1—1046

Prout LB (1912-1916) Die spannerartigen Nachtfalter. In: Seitz

A (ed.): Die Gross-Schmetterlinge der Erde 4: i-v, 1-479, pls.

1—25. Stuttgart, Alfred Kernen Verlag

Scoble MJ, Kriiger M (2002) A review of the genera of Macariini

with a revised classification of the tribe (Geometridae:

Ennominae). Zoological Journal of the Linnean Society 134:

257-315

Staudinger O (1892) Neue Arten u. Varietaten von palaarktischen

Geometriden. Deutsche entomologische Zeitschrift Iris.

5:181-188, pl. 2, Fig. 12-13

Stehr FW (2005) Immature Insects. Kendall/Hunt Publishing

Co., U.S. 1-768, p. 290-303

Stokoe GHT (1948) The Caterpillars of British Moths, 2nd se-

ries. London: F. Warne

Wehrli E (1938) Nachtrag zu meiner Arbeit in Nr. 31 d. J. Sei-

te 354 [Neue Gattungen, Untergattungen, Arten und Rassen

von Geometriden (Lep.).]. Entomologische Rundschau 55:

433-434

Wehrli E (1939) Einige neue Arten und Rassen aus dem

sudwestlichen Iran und aus dem Irak, sowie ein neues

Subgenus aus dem letzteren. (Lepid. Geometr.). Mitteilungen

der Miinchner Entomologischen Gesellschaft 29: 70-71

Wehrli in Brandt W (1941) Beitrag zur Lepidopteren-Fauna von

Iran (4). Einige neue Geometriden. Mitteilungen der

Miinchner Entomologischen Gesellschaft 31: 864-886, PI.

28-30

Wehrli E (1939-1954) 6. Subfamilie: Geometrinae. In: Seitz A

(ed.): Die Gross-Schmetterlinge der Erde 4, Die Spanner des

palaearktischen Faunengebietes (Suppl.): 254-722, Pls 19-53.

Stuttgart, Alfred Kernen Verlag

Wiltshire EP (1967) Middle East Lepidoptera XX, A third

contribution to the Fauna of Afghanistan. Beitrage zur

naturkundlichen Forschung in Siidwestdeutschland 26:

137-169

Wiltshire EP (1970) Osterreichische entomologische Expe-

ditionen nach Persien und Afghanistan. Annalen des

Naturhistorischen Museums in Wien 74: 387-398

Received: 15.11.2009

Accepted: 20.02.2010

Corresponding editor: D.Stiining

©ZFMK

Bonn zoological Bulletin | Volume 57 | Issue | | pp. 75-83

Bonn, April 2010

Two new species of the Plutodes costatus-group

from the Philippines and Indonesia

(Lepidoptera, Geometridae, Ennominae)

Dieter Stiining

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

E-mail: d.stuening.zfmk@uni-bonn.de

Abstract. Two new species, Plutodes magdelinae sp. n. from Mindanao Island, the Philippines, and P. thorbeni sp. n.

from Sulawesi, Indonesia, are described. Pattern and coloration clearly indicate that they are members of the P. costa-

tus-group. Besides the well-known, wide-spread and often abundant P. costatus Butler, 1886, other members of this species-

group are P. warreni Prout, 1923, and P. moultoni Prout, 1923. The latter species, described from Peninsular Malaysia,

is redescribed. It is very similar externally to P magdelinae sp. n., but clearly different in the genitalia morphology. Its

known range of distribution is fairly extended, based on new records. P. hermanowskii Stining, 2005 (Luzon, Philip-

pines) is the most recently described species in this group. DNA barcoding of all involved taxa revealed that they are

well differentiated also on the genetic level.

Key words. Plutodes, new species, P. magdelinae sp. n., P. thorbeni sp. n., Philippines, Indonesia, DNA barcoding.

INTRODUCTION

Species of the genus Plutodes Guenée [1858] are charac-

terized by broadly unipectinate antennae in both, males

and females, and their striking wing-pattern of yellow

ground colour with bands and patches in reddish-brown

or grey, the latter often highlighted by margins of irides-

cent scales. Parsons et al. (1999) listed 27 species as a

whole, distributed from the NW Himalayas (Sabathu,

type-locality of PR warreni) and Nepal (Yazaki, 1992,

1993, 1994: eight species) to tropical Australia (P. sig-

nifera Warren, 1896; cf. McQuillan & Edwards, 1996).

Holloway [1994] redescribed the genus and recorded sev-

en species from Borneo. He also provided a definition for

the tribe Plutodini.

Species of the P. costatus-group are distinguished from the

typical species of Plutodes (like the type-species P. cyclar-

ia Guenée and its relatives) externally by larger size and

by the hind wings coloured brown with only some rem-

nants of the yellow ground colour. Also in the forewings

the range of brown colour is enlarged, most definitely so

in P. costatus and P. warreni. P. moultoni has the yellow

areas of the forewings more extended, but has the apical

patch, which is very conspicuous in the two species men-

tioned before, reduced to minor orange-coloured remnants.

Instead, the apical region is marked by a pair of black, out-

wardly pointed spots. PR. hermanowskii is transitional to

the core-group in having oval apical patches to the

forewings, similar to the majority of species related to P

cyclaria.

The two new species described herein clearly belong to

the P. costatus-group, the one from Mindanao being strik-

ingly similar externally to P. moultoni, but clearly differ-

ent in male genitalia. P. thorbeni from Sulawesi is also dis-

tinctive in its habitual features, but closer to P. costatus

externally.

DNA barcodes were obtained at the Canadian Centre for

DNA Barcoding, University of Guelph, Canada, in the

course of a project called ‘All Leps’ (cf. http:// www.lep-

barcoding.org).

MATERIAL AND METHODS

This study is based mainly on material of the Lepidoptera

collection of the Zoologisches Forschungsmuseum

Alexander Koenig, Bonn, Germany (ZFMK)). In addition,

the collections of the Zoologische Staatssammlung, Mu-

nich (ZSM), including the collection Claude Herbulot, and

two important private collections (coll. M. D. Sommerer,

Munich, and coll. M. Schaarschmidt, Leipzig) were stud-

ied. The relevant material housed at The Natural History

Museum, London (NHM), has been checked by J. D. Hol-

loway, and images of the holotype of P. moultoni Prout

and its labels were provided by the collection manager at

the NHM Lepidoptera department.

76 Dieter Sttining

Figs 1-15. Adults of Plutodes spp. 1-4. Plutodes moultoni Prout, 1: &’, N. Thailand, 2: underside; 3: &@, Sumatra, 4: underside

of 3; S—6. Plutodes magdelinae sp. nov., holotype @, Mindanao; 5: upper side, 6: underside; 7-12. Plutodes thorbeni sp. nov., Su-

lawesi, 7-8: holotype &, 9-12: @ paratypes (8, 10, 12: underside); 13-15. Plutodes costatus Butler; 13: @, Sumatra; 14: 9, Suma-

tra; 15: underside of 9.

Two new species of the Plutodes costatus-group from the Philippines and Indonesia Ta

External features and genitalia of all species were exam-

ined. The illustrations of whole moths (upper side and un-

derside) and moth genitalia were prepared from digital im-

ages, and were subsequently edited in Adobe Photoshop.

The terminology of the genitalia largely follows that of

Klots (1956).

A total of 38 specimens of the ““Plutodes costatus species-

group” has been processed at the Canadian Centre for

DNA Barcoding (CCDB) in order to obtain DNA barcode

sequences (COI, 5’, barcode fragment, 658 bp.), most of

them successfully, though partly 15 years old (for details

of the procedure see Ivanova et al., 2006).

Abbreviations. NHM: The Natural History Museum,

London; ZFMK: Zoologisches Forschungsmuseum

Alexander Koenig, Bonn, Germany; ZSM: Zoologische

Staatssammlung, Munich, Germany.

TAXONOMY

Redescription of Plutodes moultoni Prout (Figs 1-4,

16-18, 19, 20)

Plutodes moultoni Prout, 1923, Annals and Magazine of

Natural History (9) 11: 321.

Type locality. [Peninsular Malaysia] Selangor, Bukit Ku-

tu, 3457 ft.

Holotype o’. NHM, London.

Male (Figs 1-4). Wingspan 31—38mm, length of forewing

17—20mm. Apex rounded, termen evenly curved, tornus

rounded. Fovea lacking. Vein R, of forewing arising from

cell close to the common stem of R,_;, anastomosing with

Sc at varying length. R, branching off from the middle of

the common stem of R;,,4. Forewings yellow, with a grey-

ish-brown basal area and a distal, indistinct, orange patch,

both parts broadly connected. The connecting band is al-

ways present, the orange patch may be more or less dis-

tinctly developed. Distal to and rarely connected with the

orange patch are two blackish-brown, subapical, irregu-

larly triangular spots, their tips pointing towards the ter-

men. Basal area with an ill-defined anterior margin, reach-

ing well into the discal cell, an acute protuberance at the

position of the discal vein and a short concavity prior to

the connection with the apical patch. Posterior half with

indistinct, transverse, greyish striation. Iridescent scales

are found along basal two-thirds of costa, anterior mar-

gin of basal greyish-brown area, around orange patch and

densely covering the triangular spots. Fringe of termen

yellow, that of hind-margin brown.

Bonn zoological Bulletin 57 (1): 75-83

Hind wings with margin evenly rounded, fringe yellow.

A narrow yellow marginal band from apex to tornus,

slightly broader at apical half. Remaining part of hind

wing dark blackish or greyish brown, with extensive trans-

verse striation. Distal margin of dark area lined with iri-

descent scales. Discal spot or streak not visible. In some

specimens a postmedial, double, indistinct zigzag-line

present.

Underside of both wings lighter yellow, forewings with a

transverse, slightly curved, blackish bar from tornus to-

wards middle of costa, but not reaching the latter, less

strongly developed or absent in Malaysian (including the

type) and Sumatran specimens (see Figs 2, 4). A band

proximal to this bar rather greyish-brown than yellow.

Hind wings on underside with a large, blackish, oval patch,

touching or almost reaching the anterior margin near apex.

Basal area of wings light yellow or suffused with darker

scales (the latter character found again in specimens from

Malaysia and Sumatra). Distally a broad, light yellow mar-

ginal band of even width. Fringe yellow.

Head with frons distinctly narrower than eye-diameter,

smoothly scaled, greyish-brown, but with groups of up-

right, narrow scales at dorsal margin. Vertex with larger,

yellow scales and white scales between the antennal bases,

basal segments and dorsal side of lower part of antennae

also white. Palps slightly up-curved, of a light greyish-

brown colour, the third segment small, tapering. Proboscis

well developed. Antennae unipectinate, with longest

branches about 7,5 times as long as the diameter of the

flagellum shaft. Terminal one third of antennae simple.

Patagia and anterior parts of tegulae yellow, posterior part

of tegulae, thorax and abdomen greyish brown. Tibiae of

hindlegs not dilated, thus without hair-pencil. Setal comb

on third abdominal sternite as well as sterno-tympanal

spine lacking. Tympanal organ without lacinia.

Male genitalia (Figs 19, 20). Valves elongate, weakly

sclerotized, apex rounded, lamina internally densely se-

tose. Valve costa, like the whole genitalia capsule, weak-

ly sclerotized, the sacculus being the only strongly scle-

rotized part. It reaches a little more than half the length

of the ventral valve margin, bearing an acute distal tooth-

like process with a second tip at some distance pointing

basally. A strong basal process with a more rounded tip

is directed dorsally. There is also a roundly protruding area

opposite to the sacculus on the external surface of the

valve. Uncus broad, dorso-ventrally flattened, dorsally se-

tose, with a dense group of stiff, narrow, upright scales

near base, slightly curved ventrad, with pointed tip. Socii

present, small, but with long setae. Gnathos with narrow

lateral arms, central part a broad plate at right angles to-

wards the lateral arms, without medial process. Transtil-

©OZFMK

78 Dieter Sttining

ie 7 RE

Figs 16-18. Holotype & of Plutodes moultoni Prout, 1923. 16:

upper side; 17: underside, 18: labels. Images reproduced by kind

permission of the Trustees of the Natural History Museum, Lon-

don.

la a triangular, curved, cap-like plate, its tip pointing dis-

tally, with a pair of short, narrow processes arising from

the proximal corners and directed caudally. Juxta a round

plate, with a v-shaped incision with irregular margins dor-

sally. Saccus narrow, strongly elongate, rounded distally.

The aedeagus straight, club-shaped, with a stout proximal

and a narrow distal half, the vesica with an elongate, se-

mi-cylindrical sclerotization and a small group of cornu-

ti, a long, curved, distal one and 3—4 distinctly smaller

ones, arising from a common base. Bulbus ejaculatorius

moderately large, about half the length of the aedeagus.

Male genitalia of specimens from N Thailand a little small-

er and slightly differing to those from Sumatra in the

length-proportions of the saccular processes (Fig. 20).

Bonn zoological Bulletin 57 (1): 75-83

Female similar to male, wingspan 35—37 mm, length of

forewing 19-21 mm, with forewings a little broader and

termen more strongly curved. Pectinations of antennae

about one fifth shorter than in males and a little narrow-

Material studied. Type material. Holotype, o’, [Peninsu-

lar Malaysia], Bukit Kutu, Selangor, April 1915, 3457’;

coll. NHM (see Figs 16-18). J. D. Holloway (London,

pers. comm.) mentioned two more specimens from

Peninsular Malaysia in coll. NHM, but Prout (1923) ob-

viously only had one specimen at hand when he described

this species (i.e. the holotype).

Further material. 19, [Peninsular] Malaysia, Taiping [~

4°51’N 100°46’E], May [19]79;

40’, N. Sumatra, vic. Tiga-Dolock, 18., 25.v.1972, leg. Ro-

esler & Ktippers; coll. ZFMK; 1c’, N. Sumatra, Dolok Me-

rangir, 21.1x.—5.x.1967, leg. Dr. E. Diehl, ex colli Dr Re

Bender, coll. ZSM. (more specimens from N. Sumatra in

coll. M. D. Sommerer, Munich).

2c", Borneo, Sabah, route Keningau-Kimanis PK 28.5,

1300m, 8., 11.viti.1991, leg. Haxaire. Herbulot collection

in coll. ZSM;

1c’, S. Vietnam, Prov. Quang Tri, Partung, Da Krong,

1.—3.vi.2005; coll. ZFMK;

1o', SE Thailand, Prov. Trat, Kho Chang, 0m, 25.—

30.x1.2004, leg. T. Ihle; coll. ZFMK;

8", N. Thailand, Chiang Mai Prov., Doi Suthep, 1050m,

25.x1.2000, leg. D. Stiining; 4c", same locality, 1100m,

15.-19.1.2008, leg. T. Ihle; 10’, N. Thailand, Chiang Mai

Prov., Doi Angkhang, W Fang, 1600m, 21.v.2004; 19, N.

Thailand, Chiang Mai Prov., Doi Inthanon, 1500m,

28.—30.iv.2006, leg. T. Ihle; 19, N. Thailand, Mae Hong

Son Prov., ca. 30km E Khun Yuam, 950m, 1.v.2006, leg.

T. Ihle; coll. ZFMK;

16 specimens from Doi Suthep, Chiang Mai Prov., are kept

in the collection of Bro. Amnuay Pinratana, St. Gabriel’s

College, Bangkok.

1c, S. China, S. Yunnan Prov., Mangxi Ba Mts., Simao

distr., 18km S Simao city, 1280m, 16.i11—10.1v.2000, ex

coll. Dr. R. Brechlin. Coll. ZFMK.

Distribution and bionomics. Peninsular Malaysia (type-

locality), Borneo, Sumatra; Vietnam, SE & N. Thailand,

S. China (first record for all localities except Peninsular

Malaysia). Specimens have been collected in January,

March-April, May, June, Sept.—October, November; at el-

©ZFMK

Two new species of the P/utodes costatus-group from the Philippines and Indonesia 79

evations between 0 and 1600m. The immature stages are

unknown.

Plutodes magdelinae sp. nov. (Figs 5, 6, 21)

Male (Figs 5—6). Wingspan 37—38mm, length of forewing

21—22mm. Apex rounded, termen evenly curved, tornus

rounded. Fovea lacking. Vein R, of forewing arising from

anterior vein of cell close to the common stem of R_; —

both at rather large distance from upper corner of cell — ,

anastomosing with Sc at varying length. R, branching off

from the middle of the common stem of R3,4. Forewings

yellow, with a greyish-brown basal area and a distal, in-

distinct, orange patch, both parts narrowly connected. The

connecting band is always present, the orange patch may

be more or less distinctly developed. Distal to the orange

patch are two blackish-brown, subapical, elongate trian-

gular spots, their tips pointing towards the termen. Basal

area with an ill-defined anterior margin, reaching just a

little beyond the posterior margin of the discal cell. Pos-

terior half with indistinct, transverse, greyish striation. [ri-

descent scales are not very obvious, weakly covering basal

two-thirds of costa, anterior margin of basal greyish-brown

area, orange patch and the submarginal area near the tri-

angular spots. Fringe of termen yellow, that of hind-mar-

gin brown.

Hind wings with margin evenly rounded, fringe yellow.

A narrow yellow marginal band from apex to tornus, dis-

tinctly broader at apical half. Remaining part of hind wing

dark blackish or greyish brown, with extensive transverse

striation. Distal margin of dark area lined with iridescent

scales. Discal spot visible as a short, curved, white streak.

Postmedial ziczac-line absent.

Underside of both wings (Fig. 6) lighter yellow, forewings

with a broad, posterior, greyish-brown band between hind

margin and posterior vein of cell, but not reaching termen.

Hind wings on underside with a large, blackish-brown,

oval patch, almost reaching the anterior margin near apex.

Basal area of wings strongly suffused with darker scales.

Distally a rather broad, light yellow marginal band of even

width. Fringe yellow.

Head with frons distinctly narrower than eye-diameter,

smoothly scaled, greyish-brown, with thin white lateral

margins, but with groups of upright, narrow scales at dor-

sal margin. Vertex with larger, yellow scales and white

scales between the antennal bases, basal segments and dor-

sal side of lower part of antennae also white. Palps slight-

ly up-curved, basal segment predominantly white, second

and third of a light greyish-brown colour, the third seg-

ment small, tapering, with a white tip. Proboscis well de-

veloped. Antennae unipectinate, with longest branches

Bonn zoological Bulletin 57 (1): 75-83

about 8,5 times as long as the diameter of the flagellum

shaft. Terminal one third of antennae simple. Patagia and

anterior parts of tegulae yellow, posterior part of tegulae,

thorax and abdomen greyish brown. Tibiae of hindlegs not

dilated, thus without hair-pencil. Setal comb on third ab-

dominal sternite as well as sterno-tympanal spine lacking.

Tympanal organ without lacinia.

Male genitalia (Fig. 21). Valves elongate, weakly scle-

rotized, apex rounded, lamina internally densely setose.

Valve costa, like the whole genitalia capsule, weakly scle-

rotized, but sacculus with a prominent and strongly scle-

rotized, plate-like process, inserting in an upright position

close to the ventral margin of the saccular region. It is get-

ting broader distally, with an acute tip, narrower basally,

with a rounded, very short, dorsal bulb. Outer margin of

the plate densely covered with short, appressed setae.

Length a little more than half the length of the ventral

valve margin. Uncus broad, dorso-ventrally flattened, dor-

sally setose, with a dense group of stiff, narrow, upright

scales near base, slightly curved ventrad, with pointed tip.

Soci very small, marked by a few long setae only.

Gnathos with narrow lateral arms, central part a broad

plate at right angles towards the lateral arms, without me-

dial process. Transtilla a triangular, curved, cap-like plate,

its tip pointing distally, with a pair of short, narrow

processes arising from the proximal corners and directed

caudally. Juxta a narrow plate, with a deep, v-shaped in-

cision dorsally. Saccus narrow, strongly elongate, round-

ed distally.

The aedeagus almost straight, club-shaped, with a stout

proximal and a narrow distal half, the vesica with an elon-

gate, semi-cylindrical sclerotization and a small group of

5—6 cornuti, not much differing in size. Bulbus ejacula-

torius moderately large, about half the length of the aedea-

gus.

Diagnosis. P. magdelinae is distinguished from the exter-

nally very similar P. moultoni by slightly larger size, the

wing-shape (forewings more elongate with more strong-

ly oblique termen), a narrower greyish-brown basal area

(anterior margin just reaching the posterior vein of discal

cell, distal margin more strongly concave). The subapi-

cal, black, triangular spots are larger and more distinctly

acute towards the termen. In the hind wings, the yellow

marginal band is much narrower at tornal half. On under-

side, the oblique bar on forewing from tornus to costa is

absent, the basal area of hind wing is much darker.

In the male genitalia, the elongate, simple plate on sac-

culus without a separate basal process is distinctive.

Female unknown.

©OZFMK

80 Dieter Sttining

Figs 19-22. c'-genitalia of Plutodes spp. 19: P. moultoni Prout, Sumatra (vesica partly everted); 20: P. moultoni Prout, N. Thai-

land; 21: P magdelinae sp. nov., paratype, coll. no. LEP 2010/ 469-2; 22: P. thorbeni sp. nov., paratype, coll. no. LEP 2010/ 472-

3. (bottom left: aedeagus, bottom right: enlarged apical part of aedeagus).

Type-material. Holotype, o', “Philippinen, Mindanao

(Prov. Davao del Sur), Mt. Apo, SE-route via Kapatagan,

1570m, 10.-12.July 1996, prim. forest, leg./ ex coll. Dr.

Ronald Brechlin. Coll. ZFMK, coll. no. LEP 2010/468

(Biodat).

Paratypes. 2c’, same data as holotype, coll. nos. LEP

2010/469-1, 469-2; 1c", Philippinen, Mindanao, Mt. Ki-

tanglad, Intavas, 600m, 8° 10’N 124°56’E, 23.24. ii.

2009, leg. J. H. Lourens & K. Knoblich, coll. ZFMK, coll.

no. Lep 2010/470;

Bonn zoological Bulletin 57 (1): 75-83

Io, Philippinen, Mindanao, Mt. Dalongdong, 9-17.

vil.1999. Coll. M. Schaarschmidt, Leipzig. [“Dalong-

dong” correctly is named Dolongdong and 1s a village on

a high plateau at the western slopes of Mt. Kalatungan,

45km SW Talakag, 40km W Maramag, 07°53’N

124°40’E; J.H. Lourens, Luzon, in litt.]

Distribution and bionomics. Probably endemic to Min-

danao Island, the Philippines, only known from Mt. Apo,

Mt. Kitanglad and the Dolongdong high plateau near Mt.

Kalatungan. Specimens have been collected in February,

July and August at 600m and 1570m a.s.l. The immature

stages are unknown.

©OZFMK

Two new species of the Plutodes costatus-group from the Philippines and Indonesia 81

Etymology. The present new species is dedicated to

Magdeline Camille Vlasimsky (formerly Frankfurt a. M.,

Germany, now Dallas/ Texas), in recognition of her and

her parents’ Lezlie and Stan valuable contribution to the

taxonomic research of the author and to conservation of

biodiversity.

Plutodes thorbeni sp. n. (Figs 7-12, 22)

Male (Figs 7-12). Wingspan 34-39mm, length of

forewing 19—23mm. Apex slightly acute, termen evenly

curved, tornus rounded. Fovea lacking. Vein R, of

forewing arising from anterior vein of cell close to the

common stem of R,_;— both at rather large distance from

upper corner of cell- , anastomosing with Sc at varying

length. R, branching off from the middle of the common

stem of R3,,4. Forewings yellow, with a greyish-brown

basal area and a distal, more or less distinct, irregularly

quadrangular, dark brown to orange-brown, rarely light or-

ange patch, both parts broadly or narrowly, sometimes not

connected. Internal surface of the patch with a dark, waved

line. Apical corner of the patch forming an elongate, black

spot, its tip connected with the apex of the forewings. The

spot is most distinctive in specimens with a reduced, or-

ange, apical patch. Basal area with an ill-defined anteri-

or margin, reaching well beyond the posterior margin of

the discal cell, with an acutely triangular protuberance at

the position of the discal vein and a broad concavity pri-

or to the connection with the apical patch. Posterior half

with indistinct, transverse, greyish striation. A margin of

iridescent, leaden-grey scales along the anterior margin

and around the apical patch, including the black apical

streak, but not along distal margin. Fringe of termen yel-

low, that of hind-margin brown.

Hind wings with margin evenly rounded, fringe yellow,

a narrow yellow marginal band from apex to middle of

termen, with rather straight internal border. Larger part of

hind wing dark blackish or greyish brown, with extensive

transverse striation. Distal margin of dark area lined with

iridescent scales. Discal spot weak, visible as a short,

curved, white streak. Indistinct postmedial double waved

or ziczac-line present in most specimens.

Underside of both wings (Figs 8, 10, 12) yellow, fore-

wings with a tiny, black, apical spot and a short, transverse,

almost straight, blackish bar from tornus towards the cen-

tre of the wing. There is a greyish-brown band between

hind margin and posterior vein of cell, more or less agree-

ing with the dark basal area of the upper side and also a

dark patch representing the apical patch of upper side.

Hind wings on underside with a large, blackish-brown,

oval patch, broadly touching the anterior margin near apex.

Basal area of wings yellow, often suffused with darker

Bonn zoological Bulletin 57 (1): 75—83

scales. Distally a narrow, light yellow marginal band from

apex to the middle of termen, basal half represented by

the yellow fringe scales only.

Head with frons distinctly narrower than eye-diameter,

smoothly scaled, greyish-brown, with thin white ventral

margin, but with groups of upright, narrow scales at dor-

sal margin. Vertex with larger, yellow scales and white

scales between the antennal bases, basal segments and dor-

sal side of lower part of antennae also white. Palps slight-

ly up-curved, basal segment predominantly white, second

and third of a light greyish-brown colour, the second edged

white distally, the third segment small, tapering, with a

white tip. Proboscis well developed. Antennae unipecti-

nate, with longest branches about 8,5 times as long as the

diameter of the flagellum shaft. Terminal one third of an-

tennae simple. Patagia and anterior parts of tegulae yel-

low, posterior part of tegulae, thorax and abdomen grey-

ish brown. Tibiae of hindlegs not dilated, thus without

hair-pencil. Setal comb on third abdominal sternite as well

as sterno-tympanal spine lacking. Tympanal organ with-

out lacinia.

Male genitalia (Fig. 22). Valves elongate, weakly scle-

rotized, apex rounded, lamina internally densely setose.

Valve costa, like the whole genitalia capsule, weakly scle-

rotized, but sacculus with a prominent and strongly scle-

rotized, triangular plate with an acute tip distally, insert-

ing in an upright position close to the ventral margin of

the saccular region. Basally, it is getting narrow and

widens again to a rounded, dorsal bulb. Outer margin of

plate and bulb scobinate and slightly setose. Length of sac-

culus and its process a little more than half the length of

the ventral valve margin. Uncus broad, dorsally rounded,

ventrally flattened, dorsally setose, slightly curved ven-

trad, with pointed tip, with a dense group of stiff setae near

base. Socii very small, marked by a few long setae only.

Gnathos with narrow lateral arms, central part a broad

plate at right angles towards the lateral arms, without me-

dial process. Transtilla a triangular, curved, cap-like plate,

its tip pointing distally, with a pair of short, narrow

processes arising from the proximal corners and directed

caudally. Juxta a narrow plate, with a deep, v-shaped 1n-

cision dorsally. Saccus narrow, strongly elongate, round-

ed distally.

The aedeagus club-shaped, with a stout proximal and a

narrow, slightly curved distal half, the vesica with an elon-

gate, semi-cylindrical sclerotization and a small group of

5—6 cornuti, not much differing in size. Bulbus ejacula-

torius moderately large, about half the length of the aedea-

gus.

Diagnosis. P. thorbeni is in no way similar to P. moultoni

or P. magdelinae, but exterrnally quite close to P. costa-

©ZFMK

82 Dieter Stiining

tus, the type of this species-group (Figs 13—15). Both are

of the same size and the arrangement of yellow and brown

pattern elements is very similar. In detail, P thorbeni is

distinguished from P. costatus in the apical forewing patch

being smaller, darker und more strongly irregular. P. costa-

tus has the internal surface of this patch rather grey in-

stead of brown. P. thorbeni has a black, acute apical spot

or streak, best seen in specimens with a reduced apical

patch (Figs 9, 11). It is strongly highlighted by iridescent

scales and absent in P. costatus. There is a curved, black-

ish antemedial line in P. costatus, absent in P. thorbeni.

Fringe of termen of forewings is completely yellow in P.

thorbeni (partly grey in P. costatus) and there is more of

the yellow ground colour distal to the brown pattern ele-

ments. In the hind wing, the tornal half is narrowly yel-

low in P. thorbeni, but brown — including fringe — in P

costatus. The underside is very different in both species,

i.e. the large black patch of the hind wing seen in P. thor-

beni is absent in P. costatus, the surface being light grey

instead in the latter (Fig 15). The underside of P. thorbeni

is much more similar to P. magdelinae and P. moultoni.

Also the male genitalia are strongly different, especially

in the shape of the saccular process, which is also more

similar to P. magdelinae (Fig 21), and cannot be mistak-

en.

Female unknown.

Type material. Holotype, o’, “Indonesia, C. Sulawesi,

Tambusisi mts., 1800m, 1° 45’S 121° 27°E, 11. 1996, na-

tive collector”. Coll. ZFMK, coll. no. LEP 2010/471.

Paratypes. 60’, same data as holotype; 20’, South Sula-

wesi, Quarles Mountains, ca. 30km N Rantepao, Polo

Polo, 2200m, 26.—28.1x.1995, leg. H. Schnitzler; 3c", Cen-

tral Sulawesi, Sempuraga (=Sampuraga), 1500—1700m, vi.

1996, G. Lecourt & J. Martin leg.; 2c", C. Sulawesi, vic.

Mamasa, 2° 57’S 119° 24’E, 17.-18.x.1995, leg. local col-

lector (Gala); 1c, S. Sulawesi, Puncak Palopo,

900—-1300m, x.1997, leg. local collector, ex coll. Dr. R.

Brechlin. Coll. ZFMK, coll. nos LEP 2010/472-476.

2°, S. Sulawesi, Puncak Palopo, 900—1300m, 11.1998, leg.

local collector, coll. M. Schaarschmidt, Leipzig, Germa-

ny.

2c", C. Sulawesi, Mt. Tambusisi, x. 1195, native collec-

tor; 70", same locality, 11. 1996; 2c’, same locality, i1. 1996.

Coll. K. Yazaki, Tokyo.

Further material studied. J. D. Holloway sent images

of three male specimens, collected by him during the ““Pro-

ject Wallace” of the Royal Entomological Society. Col-

Bonn zoological Bulletin 57 (1): 75-83

lecting site: “Sulawesi Utara, Dumoga Bone N.P., Site 15,

1140m, 6.8. vili.1985”. According to his recommenda-

tion, these three undissected males which also show slight

external differences compared to the paratypes mentioned

above and which come from a locality in Northern Sula-

wesi rather far away from the type locality are not recor-

ded as paratypes.

Distribution and bionomics. Probably endemic to Su-

lawesi, but widely distributed on the island. Specimens

have been collected in February, June, September and Oc-

tober at elevations between 900m and 2200m a.s.1. The

immature stages are unknown.

Etymology. The present new species is dedicated to Thor-

ben Briese (Lippstadt, Germany), in recognition of his and

his parents’ Dr. Anette Meyer-Briese and Dr. Harald Briese

valuable contribution to the taxonomic research of the au-

thor and to conservation of biodiversity.

DNA BARCODING RESULTS

Results obtained from the Canadian Centre for DNA Bar-

coding (CCDB) revealed clear specific distances between

all species involved: pairwise distances are between 3.13%

and 7.25% in the P. costatus-group. Of the species treat-

ed in the present paper, P. moultoni and P. thorbeni ex-

hibit the closest relationship (medial pairwise distance

3.54%).

In addition, an unusually deep divergence was found

among the analysed specimens of P. warreni Prout, indi-

cating a cryptic new taxon within material from Laos, Viet-

nam and Thailand. Morphological studies on this group

are carried out at present and will be published in near fu-

ture, together with a more comprehensive account on the

barcoding results of the whole group. Preliminary results

can be viewed under www.boldsystems.org/ Plutodes.

Acknowledgements. Thanks are due to A. Schintlmeister, Dres-

den, for presenting Geometridae material from the Philippines

collected by Karel Cerny, Victor Sinjaev and himself to the Lep-

idoptera & Trichoptera Section of ZFMK. I also thank Jan

Lourens, Lucena City, Luzon, the Philippines, for presenting

valuable specimens of Plutodes, including one paratype of a

species described herein. I am thankful to J. D. Holloway, NHM

London, and M. Sommerer, Munich, for various data and infor-

mation about species of P/utodes in the collections under their

care. The latter also checked, besides his private collection, the

Herbulot collection now housed at the ZSM. K. Yazaki, Tokyo,

and M. Schaarschmidt, Leipzig, provided data and sent images

of specimens in their private collection. I am also very thank-

ful to Mrs. Ulla Kreutz, Meckenheim, for preparing the figures

and plates presented in this article. The careful processing of the

material by the CCDB team (University of Guelph) is grateful-

ly acknowledged as well.

©ZFMK

Two new species of the Plutodes costatus-group from the Philippines and Indonesia 83

REFERENCES

Holloway JD (1993) [1994]. The Moths of Borneo, part 11:

Geometridae, Ennominae. Malayan Nature Journal 47: 1-309,

19 pls

Ivanova NV, de Waard JR, Hebert PDN (2006) An inexpensive,

automation-friendly protocol for recovering high-quality

DNA. Molecular Ecology Notes 6: 998-1002

Klots AB (1956) Lepidoptera. In: Tuxen SL, ed. Taxonomist’s

glossary of genitalia in insects. Copenhagen: Ejnar Munks-

gaard, 97-111

McQuillan PB, Edwards ED (1996) Geometridae. In: Nielsen

ES, Edwards ED, Rangsi TV (eds): Checklist of the Lepi-

doptera of Australia. Monographs on Australian Lepidoptera,

CSIRO Publishing, Collingwood

Parsons M, Scoble MJ, Honey MR, Pitkin LM, Pitkin BR (1999)

In: Scoble MJ (ed.): Geometrid Moths of the World: a Cata-

logue (Lepidoptera, Geometridae). CSIRO Publishing/ Apol-

lo Books, Collingwood/Stenstrup

Bonn zoological Bulletin 57 (1): 75-83

Stiining D (2005) Plutodes hermanowskii sp. n., a new species

of the P. costatus — group from the Philippines. Tinea 21: 9-22

Yazaki K (1992) Geometridae. In: Haruta T (ed.): Moths of

Nepal, part 1. Tinea 13 (Suppl. 2): 5-46

Yazaki K (1993) Geometridae. In: Haruta T (ed.): Moths of

Nepal, part 2. Tinea 13 (Suppl. 3): 103-121

Yazaki K (1994) Geometridae (part). In Haruta T (ed.): Moths

of Nepal, part 3. Tinea 14 (Suppl. 1): 5-40

Yazaki K (2004) Notes on geometrid moths (Lepidoptera, Geo-

metridae) from Nanling Mts, S. China (III). Tinea 18: 117—

126

Received: 10.12.2009

Accepted: 30.01.2010

Corresponding editor: F. Herder

©ZFMK

Bonn zoological Bulletin ] Volume 57 Issue | pp. 85-89 Bonn, April 2010

Proclinopyga ulrichi sp. nov.:

the first fossil aquatic dance fly of the subfamily Clinocerinae

(Diptera: Empididae)

Bradley J. Sinclair

Canadian National Collection of Insects, Ottawa Plant Laboratory — Entomology, CFIA, Ottawa, Canada;

E-mail: bradley.sinclair@inspection.ge.ca

Abstract. Proclinopyga ulrichi sp. nov. is described from Baltic amber, and represents the first definitive clinocerine fos-

sil species. Palaeoparamesia proosti Meunier, 1902, considered a possible species of Clinocerinae, is transferred to the

Trichopezinae.

Key words. Taxonomy, Baltic amber fossil, new species, morphology

INTRODUCTION

The Empidoidea comprise five families and numerous

subfamilies of mostly predaceous flies (Sinclair & Cum-

ming 2006). Some 150 fossil species of Empidoidea (ex-

clusive of Dolichopodidae s.str.) are described (Evenhuis

1994; Solorzano Kraemer et al. 2005). Fossils are de-

scribed from most major empidoid lineages, extending the

origin of these lineages to at least the Cretaceous period

(Grimaldi & Cumming 1999).

The subfamily Clinocerinae consists of primarily aquat-

ic associated taxa, with adults mostly encountered on

emergent rocks in streams and rivers (Sinclair 1995). Two

fossil species have been classified as clinocerines.

Fushunempites furvus Hong, 2002, from Eocene Chinese

amber is possibly a clinocerine on the basis of the long

and narrow cell r,. Unfortunately the photo of the speci-

men (Hong 2002a, b) does not permit a definitive deci-

sion concerning subfamily assignment. Palaeoparamesia

proosti Meunier, 1902 has also been viewed as a possi-

ble clinocerine. A specimen of this Baltic amber species,

identified by Meunier (1908), was examined in this study

and found not to represent a clinocerine (see Discussion).

Consequently, the description herein of a new Baltic am-

ber species of the recent genus, Proclinopyga Melander,

represents the first unequivocal clinocerine fossil. The age

of Baltic amber is assumed to be 40 million years

(Eocene).

MATERIALS AND METHODS

To prepare the pieces for identification, the amber was cut

and polished using an IsoMet® Low Speed Saw (Buehler,

Lake Bluff, Illinois, USA) cutting machine and a

Phoenix® Beta Grinder-Polisher (Buehler) polishing ma-

chine with SiC grinding paper for metallography, grit 800,

1200 and 2500, Microcut® Abrasive Paper (Buehler) plain

backing P 4000, and paperboard. To protect the holotype

of the new species the piece was embedded in synthetic

resin Araldite® 2020 (XW396/XW397) (Huntsman Ad-

vanced Materials, Everberg, Belgium) (M. Solorzano

Kraemer, Bonn, pers. comm. 2008).

Terms used for adult structures primarily follow those of

McAlpine (1981), except for the antenna where terms of

Stuckenberg (1999) were used. Homologies of the male

terminalia follow those of Sinclair & Cumming (2006).

The Baltic amber specimen of Palaeoparamesia examined

in this study was on loan from the Geoscience Centre of

the University of Gottingen (Germany).

Genus Proclinopyga Melander

Proclinopyga Melander, 1928: 220. Type-species, Pro-

clinopyga amplectens Melander, 1928 (original designa-

tion).

86 Bradley J. Sinclair

Diagnosis. Proclinopyga is distinguished from other gen-

era of the Clinocerinae by its protruding face, bare eyes,

labellum with pseudotracheae, broad wings with Sc

evanescent, fifth tarsomere lacking extension, dorsal mar-

gin of the epandrium expanded with setae and female ab-

domen truncate with stout and ridged cercus.

a al

Proclinopyga ulrichi sp. nov. (Figs. 1, 2, 4, 5)

Type Material. Male holotype in Baltic amber, with fol-

lowing label data: “HOLOTYPE/ Proclinopyga/ ulrichi/

Sinclair”; “CNC DIPTERA/ # 12248”: amber is imbed-

ded in resin with following dimensions: 11 x 9 x 6 mm;

Figs 1-3. 1. Proclinopyga ulrichi, habitus. 2. Proclinopyga ulrichi, male terminalia, postero-lateral view of right side. 3. Palaeo-

paramesia proosti, habitus.

Bonn zoological Bulletin 57 (1): 85-89

©ZFMK

Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly of the subfamily Clinocerinae 87

included in amber block is a male chironomid (missing

abdomen). The holotype is housed in the amber collec-

tion of Dr. Hans Ulrich at the Zoologisches

Forschungsmuseum A. Koenig (Bonn, Germany).

Recognition. This species (Fig. 1) is distinguished by the

lengthened, attenuated apical half of the postpedicel, con-

voluted phallus with its hair-pin bend at mid-length and

row of stout posteroventral setae on the male mid-femur.

Description. Male. Eye bare, dichoptic; lower half of face

with slight protuberance in profile. Antenna with scape

slightly longer than pedicel; postpedicel pointed ovate,

longer than pedicel and scape combined, strongly atten-

uated and prolonged in apical half; arista-like stylus 1.5

times longer than postpedicel. Proboscis short; palpus

club-shaped; labellum held in horizontal position, suck-

er-like.

Alternating biserial row of acrostichal setae, ending at

Figs 4-5. 4. Proclinopyga ulrichi, wing. 5. Proclinopyga ulrichi, male terminalia, lateral view of right side. Abbreviations: cl

cere — clasping cercus; epand — epandrium; epand Ib — epandrial lobe; hypd — hypandrium; ph — phallus; sur — surstylus.

Bonn zoological Bulletin 57 (1): 85—89

©OZFMK

88 Bradley J. Sinclair

prescutellar depression; 6—7 dorsocentral bristles, foremost

large and positioned toward postpronotal lobe; 1 post-

pronotal bristle; 1 presutural supra-alar bristle; 2 noto-

pleural bristles; 2 postsutural supra-alar bristles; 1 posta-

lar bristle; 4 scutellar bristles. Laterotergite with short,

black setae in row.

Wing (Fig. 4) (length 3 mm) broad, slightly infuscate;

basal costal bristle long, reaching to near humeral

crossvein; anal lobe well developed; Ry strongly diver-

gent from Rs.

Legs with evenly scattered setae. Mid femur with pos-

teroventral row of long, stout setae, longer than width of

femur. Mid tibia bearing stout, short, erect ventral setae;

one antero- and one posterodorsal setae near base. Hind

tibia with row of erect dorsal and anteroventral setae on

distal two-thirds.

Abdomen with tergite 6 with transverse row of 6 erect dark

setae, length more than half width of abdomen. Termina-

lia (Figs. 2, 5): Hypandrium cone-shaped, as long as epan-

drium is wide. Phallus long, slender and convoluted, with

strong bend at mid-length; apex slightly expanded. Epan-

drium with expanded posterodorsal margin, bearing long

setae and setulae; epandrial lobe continuous with expand-

ed region of epandrium, bearing long setae and attenuat-

ed apically. Surstylus closely associated with epandrial

lobe; shallow notch proximal to slightly expanded and

truncate apex. Clasping cercus digitiform, straight, longer

than surstylus; bearing short setulae.

Female. Unknown.

Geographical Distribution and Phenology. Extant

species of Proclinopyga are confined to the Nearctic, east-

ern Palearctic, and Oriental regions (Sinclair 1995). This

fossil species is the only known representative of this

genus in Europe and the western Palearctic Region. Adults

of Proclinopyga are associated with running water habi-

tats, most commonly collected in streams, creeks, and cas-

cades. The immature stages of this genus are unknown.

Phylogenetic Relationships. Proclinopyga comprises ten

described species and at least this many undescribed

species (Sinclair 1995). The slender, highly convoluted

phallus (Fig. 5) of the fossil species is most similar to ex-

tant species in North America [e.g., P_ flavicoxa Melander,

1928 (see Sinclair 1995, fig. 13)].

Etymology. This species is dedicated to Dr. Hans Ulrich

who has amassed a rich collection of little known amber

species of Empidoidea.

Bonn zoological Bulletin 57 (1): 85-89

DISCUSSION

Proclinopyga ulrichi represents the first definitive

clinocerine fossil taxon. The presence of Clinocerinae in

Baltic amber was first definitively stated by Ulrich (2003,

table, fig. 1) and was based on the specimen described here

(H. Ulrich, Bonn, pers. comm. 2008). As in other subfam-

ilies of the Empidoidea (exclusive of Dolichopodidae

s.str.), Baltic amber fossils often belong to Recent genera

(see discussion by Ulrich 2003). Clinocerines appear to

be underrepresented in amber compared to other groups

of empidoids, likely resulting from their very close asso-

ciation with flowing water habitats. Adults are rarely col-

lected away from emergent rocks in streams and rivers and

they do not form aerial swarms and consequently rarely

would they be associated with tree trunks and other

sources of amber.

Palaeoparamesia proosti was described on the basis of a

single male specimen (No. 2118) (Meunier 1902). Meu-

nier (1908) listed two additional specimens of this species

(Nos. 8437, 7786) in his monograph on Baltic amber Em-

pididae. Only specimen No. 8437 from these original se-

ries remains in the Gottingen collection (H. Ulrich, Bonn,

pers. comm. 2001). This male specimen was examined and

found to be in excellent condition (Fig. 3) and conspecif-

ic with P proosti. This examination revealed that

Palaeoparamesia Meunier should be assigned to a group

of several small genera currently found in the western

Nearctic region and assigned to the Trichopezinae; 1.e.,

Boreodromia, Nearctic species of Apalocnemis Philipp

and two winter collected species of a genus with anten-

nae morphologically similar to Gloma Meigen (Sinclair

2008). Palaeoparamesia is characterized by a dichoptic

male head, pair of long frontal bristles (rare in empidoids,

except for Hilarini), postpedicel somewhat similar to Glo-

ma, Trichopezinae-like female terminalia (based on con-

specific specimen in Ulrich collection), hypandrium and

epandrium fused (at least partially), erect male cerci and

ear-like surstyli. On the basis of similarly shaped postpedi-

cel and dichoptic male head, Palaeoparamesia and the

winter species near G/oma are most closely related. These

genera will be more thoroughly examined and described

in a future publication by the author.

Acknowledgements. Many thanks to Monica Solorzano Krae-

mer (University of Bonn) for preparing the amber specimen.

Hans Ulrich (Bonn, Germany) generously permitted the exam-

ination of his amber collection, access to literature and person-

al notes on the Meunier amber collection. Mike Reich (Univer-

sity of Géttingen) kindly arranged the loan of the amber speci-

men, Palaeoparamesia proosti. Henri Goulet (CNC — Canadi-

an National Collection) produced the photographs of the amber

inclusions. Jeff Cumming (CNC), H. Ulrich and three anony-

mous reviewers provided valuable comments on earlier drafts.

©OZFMK

Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly of the subfamily Clinocerinae 89

REFERENCES

Evenhuis NL (1994) Catalogue of the Fossil Flies of the World

(Insecta: Diptera). Backhuys Publishers, Leiden

Grimaldi D, Cumming J (1999) Brachyceran Diptera in Creta-

ceous ambers and Mesozoic diversification of the Eremoneu-

ra. Bulletin of the American Museum of Natural History 239:

1-124

Hong Y-C (2002a) Amber Insects of China. Beijing Science &

Technology Press, Beijing

Hong Y-C (2002b) Atlas of Amber Insects of China. Henan Sci-

entific & Technological Publishing House, Beijin

McAlpine JF (1981) Morphology and terminology — Adults.

[Chapter] 2. Pp 9-63 in: McAlpine JF, Peterson BV, Shewell

GE, Teskey HJ, Vockeroth JR & Wood DM (coords.) Man-

ual of Nearctic Diptera, Volume |. Agriculture Canada Mono-

graph 27

Melander AL (1928) Diptera, Fam. Empididae. Pp. 1-434 in:

Wytsman P (ed.) Genera Insectorum 185 (1927)

Meunier F (1902) Description de quelques dipteres de l’ambre.

Annales de la Société scientifique de Bruxelles 26: 96-104

Meunier F (1908) Monographie des Empidae de l’ambre de la

Baltique et catalogue bibliographique complet sur les dipte-

res fossiles de cette résine. Annales des Sciences Naturelles

Zoologie 7: 81-135, pls HI-XII

Bonn zoological Bulletin 57 (1): 85—89

Sinclair BJ (1995) Generic revision of the Clinocerinae (Empi-

didae), and description and phylogenetic relationships of the

Trichopezinae, new status (Diptera: Empidoidea). The Cana-

dian Entomologist 127: 665—752

Sinclair BJ (2008) Review of three little-known monotypic gen-

era (Diptera: Empidoidea: Brachystomatidae), assigned to Tri-

chopezinae. Zootaxa 1754: 52-62

Sinclair BJ, Cumming JM (2006) The morphology, higher-lev-

el phylogeny and classification of the Empidoidea (Diptera).

Zootaxa 1180: 1-172

Solorzano Kraemer MM, Sinclair BJ, Cumming JM (2005) Five

new species of Tachydromiinae (Diptera: Empididae s.1.) from

New World Tertiary ambers. Zootaxa 1010: 37-52

Stuckenberg BR (1999) Antennal evolution in the Brachycera

(Diptera), with a reassessment of terminology relating to the

flagellum. Studia dipterologica 6: 33-48

Ulrich H (2003) How recent are the Empidoidea of Baltic am-

ber? Studia dipterologica 10: 321-327

Received: 09.01.2009

Accepted: 16.02.2009

Corresponding editor: N. Dorchin

©OZFMK

Bonn zoological Bulletin | Volume 57 Issue | | pp. 91-98

Bonn, April 2010

Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808)

(Coleoptera: Chrysomelidae)

Munna Rani Saikia!; Lakshmi Kanta Hazarika!. 2; Surajit Kalita! & Ramen Lal Baishya!

‘Assam Agricultural University, Department of Entomology, Jorhat-785 013, Assam, India;

2E-mail: Ikhazarika@aau.ac.in

Abstract. Number and types of sensilla on each antennal segment of male and female adult rice hispa Dicladispa armigera

(Olivier) (Coleoptera: Chrysomelidae) were determined based on light and scanning electron microscopic observations.

The males had a significantly greater total number (@ 1828.11) of sensilla than females (@ 1764.43). Five types of sen-

silla, namely, sensilla chaetica, sensilla trichodea I, sensilla trichodea II, sensilla basiconica and pit or coeloconic sensil-

la were distinguished in both sexes. Sensilla trichodea I and II were distributed over the entire length of the antenna,

whereas sensilla chaetica were observed only on the apical five flagellomeres. Methoprene affected antennal morpholo-

gy by producing two-clubbed antenna (additional one at the 34 flagellomere) and alteration in the sensilla of the last fla-

gellomere.

Key words. Rice hispa, sensilla chaetica, sensilla trichodea I, sensilla trichodea II, sensilla basiconica, methoprene.

INTRODUCTION

The rice hispa Dicladispa armigera (Olivier, 1808)

(Coleoptera: Chrysomelidae) occurs in South East Asia

and Africa, and is one of the major pests of rice in many

rice growing states of India (Deka & Hazarika 1996;

Palaszek et al. 2002; Islam et al. 2004; Hazarika et al.

2005). It causes considerable damage to vegetative

stages of rice resulting in yield loss of 28% in India (Nath

& Dutta 1997), between 20-30% in Nepal (Dhaliwal et

al. 1998) and up to 52% in deepwater rice in Bangladesh

(Islam 1989); however, it may be as high as 100% in the

rice transplanted post flood in Assam (Hazarika 2005). In

order to manage this pest, attempts were made to identi-

fy pheromones in this insect, with mixed results (Deka &

Hazarika 1997). Pheromones are not only used for survey

and surveillance of insect pest but also used to manage

them.

During the last three decades, insect communication

through antennal sensilla has received substantial inter-

est (Rao et al. 1990; Kumar et al. 1995; Axtell 1999). Gen-

erally, antennae are covered with huge numbers of sen-

silla, relevant as sensory organs (Chapman 1982). Anten-

nal sensilla are involved in host recognition and mate or

microhabitat choice by pheromone- thermo- and hygrore-

' ception (Hazarika & Bardoloi 1998; Chen et al. 2003;

Ploomi et al. 2003; Marttje et al. 2004). A number of

studies has been conducted on the sensilla of other

coleopteran insects like flea beetles, Phyllotetra crucifer-

ae (Goeze, 1777), Psylliodes punctulata Melsheimer,

1847, P. affinis (Paykull, 1799) and Epitrix cucumeris

(Harris, 1851) (Ritcey & McIver 1990); however, current-

ly there is no information available describing the sensil-

la of the rice hispa. Accordingly, the present study was un-

dertaken in order to determine the number and types of

sensilla on each antennal segment of male and female

adults of D. armigera.

MATERIALS AND METHODS

For determination of number of sensilla, male and female

antennae of field collected adults were fixed separately in

carnoy-lebrun fixative for 30 min and washed for 10 min-

utes in each of the 30%, 50% and 70% alcohol. The an-

tennae were then allowed to remain for twenty minutes

in each of the 90% and absolute alcohol, after which they

were again passed through xylene and cleared in clove oil.

These were then mounted in DPX. Sensilla were observed

under the compound microscope at 100X, 400X and

1000X magnification in oil and were counted on each seg-

ment following the method of Ramaswamy & Gupta

(1981).

92 Munna Rani Saikia et al.

Table 1. Estimated number (Mean+S.E.) of sensilla on each antennal segment of adult male and female D. armigera.

Segment Male Female t-value

Scape 25.25+0.29i 21.23+0.37i 8.95**

Pedicel 28.45+0.32h 24.99+0.28h 8.77**

Flagellum (FI)

1M Il 19.28+0.25k 18.02+0.29) 2.95**

F] 2 22.48+0.30i 21.48+0.30! 2.48

Fl 3 33.66+40.328 26.69+0.308 15.15**

Fl 4 61.67+£0.65f 56.77+0.38 6.82**

Fl 6 265.20+0.414 258.36+0.494 9.7 9**

FL 302.26+0.33°¢ 295.73+0.40¢ 11.99+«

FI8 336.11+0.28> 332.84+0.39> 8.15+*

FI9 490.84+0.534 485.07+0.464 7.A47**

S- Ed) @) 0.05 0.52

C.D. 905 Ost 1.04

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p <

0.05. Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed

by * and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair;

d.f. = 49).

For determination of types of sensilla, the antennae were

fixed for six hours in carnoy-lebrun fixative and mount-

ed in DPX (Schafer & Sanchez 1976). Based on the mor-

phology of the sensilla, sensilla were classified into types.

Sizes of these sensilla were measured with the ocular mi-

crometer and readings were converted into um. We also

bleached the antennae in 2% hydrogen peroxide for 24 hr.

Permeable areas of antennae were examined under bright

field illumination using crystal violet method (Slifer 1960).

One hundred 6—12 h old pupae were treated with 5 ppm

methoprene (Altosid SE, 62.5% RS methoprene, Zoecon,

Palo Alto, CA). Twenty adultoids were randomly select-

ed for this study.

Scanning electron microscopy studies of the antennae of

the adults and adultoids were undertaken by following the

method described in Hazarika & Bardoloi (1998). Anten-

nae were dissected out of the head and cleaned in distilled

water, which were then fixed in buffered glutaraldehyde.

They were dehydrated by passing through a series of ace-

tone starting from 30% to 100%. On drying, antennae were

placed on stubs using double sided, scotch tapes and coat-

ed with gold-palladium in a sputterer (JEOL, JFC 1100,

Japan) for 5—10 min. The specimens were scanned in a

scanning electron microscope (JEOL, 35-CF, Japan) at 15

KV and photographs were taken for each of the specimens.

Bonn zoological Bulletin 57 (1): 91-98

RESULTS

The clubbed antennae of adult specimens consisted of

scape, pedicel and nine sub-segmented flagellum, each

sub-segment is called flagellomere, the first flagellomere

being the longest (0.30+0.02 mm) while eighth is the

shortest (0.15+0.02 mm). A spine is present on the ven-

tral surface of the scape though the rest of the antenna is

free from such spines. Scanning electron microscopic stud-

ies revealed the presence of scales on the scape, pedicel

and first to sixth flagellomere. The numbers of sensilla in

male and female adults of D. armigera are shown in Tab.

1. In male and female antennae, the distal segments are

densely covered with sensilla. There is a distinct differ-

ence in shape of the 9' flagellomere between the male and

female. In each segment as well as sub-segments, the male

adults had a significantly greater number of sensilla than

its female counterpart (Tab. |) except on the fifth flagel-

lomere, where it was reverse.

The mean number of sensilla per unit area is shown in Tab.

2. Both in male and female, the maximum population was

observed on the eighth and ninth flagellomere (7.11+0.03

mm and 8.89+0.03 mm, respectively) and the lowest was

observed on the scape (0.37+0.01 mm? and 0.29+0.01

mm, respectively). Though density of sensilla on each

segment was significantly higher in the male, but on the

fifth to ninth flagellomere, it was reverse (Tab. 2).

©ZFMK

Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808)

Table 2. Mean+S.E. density of sensilla (number/mm2) on each antennal segment of adult male and female D. armigera.

Segment Male Female t-value

Scape 0.37+0.01: 0.29+0.01) 10.28+*

Pedicel 0.60+0.0 1 0.48+0.01! 9.45+**

Flagellum (FI)

FL 0.42+0.011 0.35+0.01) 4.35%

1 0.63+0.025 0.57+0.014 2.80*

13) 0.96+0.012 0.81+0.012 7.77

Fl 4 2.03+0.02f 2.09+0.03f -1.94NS

BLS 4.49+0.02f 5.26+0.02¢ -26.47**

Fl 6 5.98+0.164 6.30+0.044 -1.89NS

ly 6.15+0.02¢ 7.65+0.03° -48.52**

F118 6.95+0.03> 7.75+0.03° -17.18**

FI 9 7.11+0.034 8.89+0.034 -42.61**

S-Ed: @) 0.07 0.03

C.D. 9.95 0.14 0.07

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at P < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different. Means within the rows are followed by

* and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f.

= 49).

Table 3. Size (Mean+S.E., length x width in 1m) of sensilla trichodea I on each antennal segment of adult male and female D.

armigera.

Segment Male Female t-value

Length Width

Scape 41.46+0.032 x 1.86+0.02«4 39.52+0.044 x 1.83+40.02¢4 41.07** -2.27*

Pedicel 39.23+0.08° x 1.80+0.014 38.50+0.07¢ x1.74+0.02¢ 13.70** 2.46%

Flagellum (F1)

ell I 36.50+0.095 x 1.74 +0.02¢ 36.56+0.05f x 1.7440.02¢ -0.53NS 0.01NS

Fl 2 37.18+0.078 x 1.81+0.01¢4 35.21+0.05: x 1.81+0.014 S20 ies 0.42NS

BS 37.46+0.07£ x 1.84+0.02¢4 35.80+0.03" x 1.83+40.024 23.55** OLISNS

Fl 4 37.84+0.12¢ x 1.81+0.01¢¢ 36.37+40.168 x 1.84+0.02¢4 9.53** -1.08NS

FS 37.87+0.18¢ x 1.85+0.01¢ 38.00+0.05¢ x 1.86+0.02>¢ -0.70NS -0.55NS

F16 38.00+0.074 x 1.89+0.01° 38.10+0.044e x 1.90+0.02a 1.91NS 0.31NS

lei 7 38.22+0.064 x 1.90+0.02° 38.22+0.044 x 1.89+40.014 0.01NS 0.41NS

F18 38.51+0.06¢ x 1.92+0.02° 38.52+0.04> x 1.99+0.01a> 26.17NS

0.82NS

F19 38.54+0.07¢ x 1.96+0.014 38.63+0.03> x 1.92+0.014 24.52NS 6.56**

S:Ed. ) 0.13 0.02 0.09 0.02

C.D. 0.05 0.25 0.04 0.18 0.05

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different. Means within the rows are followed by

* and ** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; df.

= 49).

Bonn zoological Bulletin 57 (1): 91-98

©OZFMK

Munna Rani Saikia et al.

Table 4. Estimated number (Mean+S.E.) of sensilla trichodea I on each antennal segment of adult male and female D. armigera.

Segment Male Female t-value

Scape DES EEOIS 1.88+0.42) 1.29NS

Pedicel 5.81+0,.52hi 5.50+0.50h 0.46NS

Flagellum (F1)

FI 1 3.93+0.501 3.45+0.2 14 0.83NS

F2 6.44+0.48h 5.02+0.47h 2.02NS

las} 12.72+0.628 7.6940.588 7.61 **

Fl 4 21.82+0.66f 11.46+0.52£ ISI

FI 5 141.77+0.66¢ 137.22+0.82¢ 3.96**

FI 6 155.43+0.664 176.46+0.844 -16.56**

BL, 181.34+0.79¢ 182.43+0.82¢ -1.13N8

FI8 209.44+41.97> 204.10+0.64> 2.74NS

FL9 235.03+0.694 198.29+0.768 Silo

S.Ed. (4) ih alkg/ 0.89

CAD enos 232 1.76

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different. Means within the rows are followed by

* and ** are significantly different at p < 0.01 and p < 0.05 probability level, respectively (Student t-test; t-values are shown against

each pair; d.f. < 49).

Table 5. Size (Mean4

armigera..

tS.E., length x width in pm) of sensilla trichodea II on each antennal segment of adult male and female D.

Segment Male Female t-value

Length Width

Scape 38.11+40.234 x 1.73+0.01¢4 36.52+0.052 x 1.70+0.03¢ef 6.77** 0.87NS

Pedicel 36.52+0.09> x 1.71+0.014 34.75+0.04> x1.68+0.02¢¢ 20.00+* 1-33NS

Flagellum (FI)

Fl 1 30.03+0.05¢ x 1.69 +0.01¢ 29.12+0.04f x 1.6740.01 11.60+* 1.10NS

Fl 2 31.45+40.014 x 1.73+0.01¢4 31.04+0.064 x 1.73+0.014e 5.93** 0.31NS

FI 3 32.06+0.09¢ x 1.72+0.01¢4 31.73+0.04¢ x 1.74+0.01¢4 2.83+* =1:59NS

Fl4 32.15+40.10¢ x 1.7340.024 30.04+0.04¢ x 1.74+0.02¢4 22.25** -0.46NS

FS 25.05+0.12i x 1.73+40.01¢4 26.51+0.065 x 1.78+0.02ab¢ -8.97** -1.64NS

Fl 6 27.73+0.128 x 1.75+0.01¢ 26.83+0.078 x 1.74+0.02be4 7.44** -0.13NS

Ie 28.24+0.05! x 1.7840.01 25.01+0.04' x 1.78+0.0 1abe 40.29%* -0.43NS

FI8 26.17+0.074 x 1.80+0.015 24.30+0.04) x 1.79+0.01a> 26.57** 1.75NS

FIL9 24.52+0.06) x 1.87+0.014 24.04+0.03* x 1.80+0.014 6.77** 8.11**

SsEds@) 0.15 0.01 0.07 0.02

C.D. 9.05 0.29 0.02 0.13 0.04

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and

*« are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49).

Bonn zoological Bulletin 57 (1): 91-98

©OZFMK

Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) 95

Table 6. Estimated number (Mean+S.E.) of sensilla trichodea II on each antennal segment of adult male and female D. armige-

Segment Male Female t-value

Scape 22.61+0.49! 19.3140.698 4.10**

Pedicel 22.92+0.56f 19.63+0.598 4.27**

Flagellum (Fl)

Jell Il 15.61+0.675 14.76+0.465 0.38NS

BZ 16.01+0.555 16.33+0.53h -0.44NS

Jel 3} 20.10+0.538 20.10+0.628 -0.01NS

Fl 4 39.88+0.61¢ 45.37+0.74! -5.55**

FI 5 63.90+0.614 64.68+0.66° -0).84NS

Fl 6 73.66+0.61¢ 70.81+0.624 -2.52*

ell 7 83.21+0.62> 79.10+0.70¢ 3.28"

F18 83.37+0.84> 90.75+0.82° Els

lal ® [36975200172 156.84+0.732 -19.07+*

S.Ed: (4) 0.89 0.93

C.D. 0.05 1.76 1.84

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and

** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49).

Table 7. Size (Mean+S.E., length x width in um) of sensilla chaetica on each antennal segment of adult male and female D. ar-

migera.

Segment Male Female t-value

Length Width

Scape — — = =

Pedicel — ~ — =

Flagellum (F1)

Fl — _ = =

FL2 — = _ =

Fl 3 = = — -

Fl 4 — - — -

ell) 30.15+0.19¢ x 2.70+0.024 32.24+0.05¢ x 2.74+0.014 -22.17 ** Play

Fl 6 33.41+0.064 x 2.86+0.02° 32.99+0.05¢4 x 2.89+0.05¢ 7.26 ** -0.64NS

F17 34.11+0.06¢ x 2.94+0.02 33.50+0.04¢ x 2.85+0.02° AQ ee 3.83 **

F1 8 37.57+0.06° x 3.00+0.03° 38.40+0.04> x 2.99+0.03> -11.92 + 0.28NS

FI9 3871501062 x3 122-0022 38.70+0.034 x 3.10+0.032 -6.67 ** 0.67NS

S-Ed: @) 0.09 0.03 0.06 0.04

C.D. 9.95 0.18 0.06 0.12 0.08

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and

** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49).

96 Munna Rani Saikia et al.

Table 8. Estimated number (Mean+S.E.) of sensilla chaetica on each antennal segment of adult male and female D. armigera.

Segment Male Female t-value

Scape — - —

Pedicel — _ =

Flagellum (FI)

Fld — — _

Fl 2 = = —

las) = = =

Fl 4 = = ~

ES 14.76+0.604 21.35+0.634 -6.33+*

F1 6 36.58+0.61¢ 10.83+0.62¢ 29.58**

FL 37.84+0.58¢ 32.34+0.61¢ 6.05**

FL8 42.55+0.66> 37.99+0.68> 5.45%»

ial’) 119.01+0.79 130.15+0.708 -12.06**

S.Ed. (+) 0.92 0.91

C.D. 9.05 1.84 1.82

SE = standard error, sample size = 50. Means within columns are separated by Duncan’s Multiple Range Test (DMRT) at p < 0.05.

Means followed by the same letter shown in superscript(s) are not significantly different; means within the rows followed by * and

** are significantly different at p < 0.01 and p < 0.05, respectively (Student t-test; t-values are shown against each pair; d.f. = 49).

Based on morphology, sensilla trichodea (ST) I, ST II, sen-

silla chaetica (SC), sensilla basiconica (SB) I and coelo-

conic sensilla were identified in both sexes of the insect.

The ST I and II were numerous and observed over the en-

tire length of the antennae whereas SC were observed on-

ly on the apical five flagellomeres. In addition, some sen-

silla suspected to be thermoreceptors (Tr) were also ob-

served on scape, pedicel and some flagellomeres.

The ST I pointed distally and curved towards the anten-

nal shaft. It is a slender structure, which tapers gradually

into a very sharp point at the distal end. In males, the

lengths of ST I varied from 36.50 + 0.09 um to 41.46+

0.03um. Likewise, their widths varied from 1.74+ 0.02 um

to 1.96+ 0.01 um. In female, the lengths varied from

35.21+ 0.05 um 39.52+ 0.04 tm and their widths varied

from 1.74+ 0.02 um to 1.99+ 0.01 tm (Tab. 3). The high-

est population of ST I was observed on the ninth flagel-

lomere (235+ 0.03+ 0.69) and the lowest was observed

on the scape (2.51+ 0.37) in males, whereas in females,

the highest population was observed on the eighth flagel-

lomere (204.10+ 0.64), and the lowest was observed on

the scape (1.88 + 0.42) (Tab. 4).

The ST II were also similar to ST I except that they were

unaffected when a solution of crystal violet was applied.

It might be due to non- permeability of the senisilla to

crystal violet. In males, the lengths of ST II varied from

24.52+0.06 to 38.11+0.23 um (Tab. 5). Likewise their

Bonn zoological Bulletin 57 (1): 91-98

widths varied from 1.69+0.01 to 1.87 + 0.01 um. In fe-

males, their lengths varied from 24.04 + 0.03 to 36.52 +

0.05 um and their widths varied from 1.67+0.01 to

1.80+0.01 um.

The highest number of ST II was observed on the ninth

flagellomere (136.75 + 0.77) and lowest on the first fla-

gellomere (15.6+0.67) in males. Likewise, in females al-

so the highest number of ST II was observed on the ninth

flagellomere (156.84 + 0.73) and the lowest was observed

on the first flagellomere (14.76 + 0.46) (Tab. 6).

The SC were pointed distally and projected outward from

a socket at an approximate angle of 50°, thick-walled and

longitudinally grooved. They were restricted to the termi-

nal five flagellomeres and were unaffected when a solu-

tion of crystal violet was applied. In males, their lengths

varied from 30.15+0.09 to 38.17+0.06 um (Tab. 7), where-

as in females they varied from 32.24 +0.05 to 38.70+ 0.03

um. In male and female, the highest number of SC was

observed on the ninth flagellomere and the lowest was ob-

served on the fifth flagellomere (Tab. 8).

A few sensilla basiconica (SB) were present in different

flagellomere of D. armigera. SB are smaller than ST

measuring 5—6 um, wall being porous. Pit or coeloconic

sensilla are also present on the antennae, however, details

of their structure were not studied.

©ZFMK

Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808) 97

After application of growth regulator, methoprene, some

deformities not only on the antennal structure but also on

the sensillar morphology were observed; the 4" flagellom-

ere got deformed. This is very prominent on the tip of the

ninth flagellomere where SC were observed to be disori-

ented (Baishya 1992).

DISCUSSION

Densely covered distal segments of the antennae as ob-

served here are also present in many other insects like

Blatella germanica (Linnaeus, 1767) (Ramaswamy &

Gupta 1981, Wheeler & Gupta 1986), Croesia curvala

(Kearfott, 1907) (Langmaid & Seabrook 1985), Bootet-

tix argentatus Bruner, 1890 (Chapman & Fraser 1989),

Geotrupes auratus Motschulsky, 1858 (Inouchi et al.

1987) and Homoeosoma electellum (Hulst, 1887)

(Faucheux 1995), and four hemipteran species (Usha Rani

& Madhavendra 2005). Palpation conducted with the dis-

tal segments of antenna may provide an explanation for

this pattern.

On the scape, pedicel and three flagellomeres (first to

third), males had significantly greater number of sensilla

per unit area than females. However, from the fourth to

ninth flagellomere, females had a greater number of sen-

silla per unit area than males; a similar pattern was report-

ed by Ritcey & McIver (1990) in case of Psylloides punc-

tulata and P. affinis. A study by Usha Rani & Madhaven-

dra (2005) suggested that sensillae in scape and pedicel

may not be used in sensory perception, which remains

however to be tested critically.

Similar to Dicladispa armigera, terminal segments of an-

tennae covered with ST I and ST II were also reported

in Homoeosoma electellum (Faucheux 1995), Phothori-

maea operculella (Zeller, 1873) (Sharaby et al. 2002), He-

licoverpa armigera (Hiibner, 1808) (Wang et al. 2002).

Bromely et al. (1980) stated that the ST at the antennal

tips of aphids have a contact chemosensory function, and

may be involved in gustation of the plant surface, which

might be the case in the insect studied here as well.

The population of ST I was highest on the ninth flagel-

lomere in males, whereas in the female it was on the eighth

flagellomere. This kind of variation in ST I population may

be associated with specific function performed by the sen-

silla. Similar cases were also recorded in Trichoplusia ni

(Hiibner, 1803) (Mayer et al. 1981), Grapholitha moles-

ta (Busck, 1916) (George & Nagy 1984), Hypera posti-

ca (Gyllenhal, 1813) (Bland 1981). However, in case of

both males and females, the ST II population was high-

est on the ninth flagellomere, which was also reported by

Bland (1981) in Hypera postica, by Kapoor (1985) in

Bonn zoological Bulletin 57 (1): 91-98

Paragnetina media (Walker, 1852) and by Ritcey & Mclv-

er (1990) in flea beetles. SC is a common type usually en-

countered in many insects (Ilango 2000). The presence of

SC on antennae was also reported by Ritcey & Mclver

(1990) in the flea beetles, P. cruciferae, P. punctulata, P.

affinis and E. cucumeris.

As Hazarika & Baishya (1996) showed, application of

methoprene induced morphogenesis in D. armigera; how-

ever, only the fourth and the last flagellomere were affect-

ed. The cause of the selectivity of this effect is unclear up

to now. Hormonal regulation of antennal sensilla is report-

ed for many insects (Wheeler & Gupta, 1986; Yamamo-

to-Kihara et al. 2004). Injection of a neurohormone,

[His’]-corazonin, reduced the number of coeloconic sen-

silla in Locusta migratoria. A similar study on D. armigera

is also required since it could help to clarify the causal net-

work in the development of the antennal sensilla pattern.

Acknowledgements. This work was partly funded by the Na-

tional Agricultural Technology Project, Indian Council of Agri-

cultural Research, New Delhi. We are also grateful to Dr. G. B.

Staal, Zoecon, Palo Alto, CA for providing the methoprene as

a gift for experimental purpose. Scanning electron micrographs

were taken at the Regional Sophisticated Instrumentation Cen-

tre, North Eastern Hill University, Shillong, Meghalaya.

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©OZFMK

98 Munna Rani Saikia et al.

Dhaliwal GS, Arora R, Randhwa NS, Dhawan AK (1998) Eco-

logical Agriculture and sustainable development. In: Proceed-

ings of an International Conference on Ecological Agriculture:

Towards Sustainable Development, Vol 1, Chandigarh, India,

15—17 November, 1997

Faucheux MJ (1995) Sensilla on the antennae, mouthparts, tar-

si and ovipositor of the sunflower moth, Homoeosoma electel-

lum (Hulster) (Lepidoptera: Pyralidae). Annales des Sciences

Naturelles Zoologie et Biologie Animale 11: 121-136

George JA, Nagy BAL (1984) Morphology, distribution and ul-

trastructural differences of sensilla trichodea and basiconica

on the antennae of the oriental fruit moth, Grapholitha mo-

lesta (Busck) (Lepidoptera: Tortricidae). International Jour-

nal of Insect Morphology & Embryology 13: 157-170

Hazarika LK (2005) Rice hispa: How much we really know

about it? In: Ramamurthy VV, Singh VS, Gupta GP, Paul AVN

(eds), Gleanings in Entomology, Division of Entomology, In-

dian Agricultural Research Institute, New Delhi, pp. 1-19

Hazarika LK, Baishya RL (1996) Effect of methoprene on rice

hispa, Dicladispa armigera (Olivier) (Coleoptera: Chrysomel-

idae). Pesticide Resarch Journal 8: 93-95

Hazarika LK, Bardoloi S (1998) Antennal and mouth part sen-

silla of the Muga silkworm, Antheraea assama (Lepidoptera:

Saturnidae). Sericologia 38: 55—63

Hazarika LK, Deka M, Bhuyan M (2005) Oviposition behav-

iour of the rice hispa Dicladispa armigera (Coleoptera:

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hispa. International Rice Research Newsletter 16: 14-15

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25: 163-172

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bution and fine structure of antennal olfactory sensilla in

Japanese dung beetle, Geotrupes auratus Mots. (Coleoptera:

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Scarabaeidae). International Journal of Insect Morphology &

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water rice (DWR). International Rice Research Notes 14: 53

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S, Dutta BC, Bhuyan M (2004) Current status of rice pests

and their management in Assam, India — a discussion with ex-

tension experts. International Rice Research Notes 29: 89-91

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antennal sensilla of the stonefly Paragnetina media (Walker)

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ogy & Embryology 14: 273-280

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three repellents against bed-bugs, Cimex hemipterus Fabr. In-

dian Journal of Medical Research 102: 20-23

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of the antennae of the blueberry leaf tier moth, Croesia cur-

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JA Van Loon, Luise EMVet (2004). Antennal sensilla of two

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cladispa armigera (Oliv.). Journal of the Agricultural Science

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Received: 03.07.2008

Accepted: 08.07.2009

Corresponding editor: M. Schmitt

©ZFMK

Book Review 99

Christoph Moning, Thomas Griesohn-Pfleger & Michael

Horn (2010). Grundkurs Vogelbestimmung: Eine Ein-

fiihrung in die Beobachtung und Bestimmung unserer

heimischen Vo6gel. 422 pp., Quelle & Meyer Verlag GmbH

& Co, Wiebelsheim, ISBN-10: 3494014167, ISBN-13

978-3494014166, 19,95 €.

Facilitating the access to birdwatching by a book is not

particularly new. There are examples from anglophone

countries as well as from Germany (e.g. “Végel beobacht-

en” by E. Bezzel). However, “Grundkurs Vogelbestim-

mung” (Basic course bird identification) is particular up

to date and specific to today’s internet generation. The

book is part of a new series of basic courses on field iden-

tification of different taxonomic groups edited by the same

publishing house.

The book is divided into two major parts. The Chapters

1—12 set the general part; the second offers a surprising-

ly extensive insight into all central European bird fami-

lies. Single chapters are rather short, the text is interrupt-

ed by boxes and figures and invites the reader to skim and

browse.

The general part includes everything from equipment

(binoculars, literature sources, appropriate clothes etc.) to

an introduction to avian behaviour, bioacoustics, and es-

pecially to the Where, How and When of birdwatching.

These chapters are the strengths of the book, and the ex-

perience of the authors who regularly work as tour guides

is quite obvious. They have a good sense of humour when

combining a cold drink and illustrated bird books as nec-

essary equipment to reflect their own observations (fig-

ure on p. 21). However, a Wikipedia definition should nev-

er be cited when explaining complex issues as the hand-

icap principle (p. 76).

The chapter on “Systematics and Taxonomy” is praisewor-

thy. Contributing author Daniel Lingenhohl does not re-

ally give an explanation of these terms, but explains on

five and a half pages — necessarily only rudimentary — “‘bi-

nary nomenclature” or the “biological species concept”.

Some shortcomings can be overlooked here. The author

exemplifies the changing character of this scientific field

by some huge changes in avian higher taxa systematics.

Sadly, his explanation on using italics for scientific names

is not completely correct and it is even more regrettable

that in some parts of the further text not even the correct

parts of his remarks were respected (p. 133 ff. subfami-

lies wrongly italicised, p.137 genus Branta not in italics).

In the conservation chapter, concepts in Germany are com-

pared to those in some neighbouring countries. Many as-

pects the authors rise are certainly true but perhaps it

would have been wiser to generalize a little bit less and

Moning/Griesohn-Pflieger/Horn

—

Grundkurs

Eine Einfiihrung zur

Beobachtung und —

Bestimmung unserer

heimischen Végel

give this issue more space. Feeding birds might be fun but

its value for conservation is at least controversial. The fig-

ure on page 125 is wrongly marked (it is a “Rotkopfwiirg-

er”, “Rotstirnwiirger” is an African species).

The second part is systematically ordered by Central Eu-

ropean bird families although this order does not seem to

be very consistent as non-passerines follow a generally

rather conservative concept while passerines are rather or-

dered by new findings. There are good, compact introduc-

tions for every family and therein focus is on the most

common birds so that beginners are not unnecessarily con-

fused. Short texts and arrows in the pictures complement

information from classical bird guides although some

overlap cannot be denied. Graphics on the topography of

many different birds increase the learning effect; obser-

vation hints enrich the text. However, the authors forgot

the huge population of Melodious warblers in Rhineland-

Palatinate and more important one should never throw

swifts in the air that were found on the ground. The au-

thors may have to take the responsibility for some bro-

ken bones in swifts.

100 Book Review

So, what’s missing? Not much, maybe the unloved glos-

sary but “carpal joint” (Carpalgelenk) and even “taxon”

might not be that easy to understand for the beginner. A

few more internet sources and examples from Austria and

Switzerland would make a broader distribution easier.

There is a growing attention for “citizen science” in or-

ganismic biology. The participation of non-professionals

has a long history in the study of birds — many important

long-term monitoring schemes would be impossible with-

out the help of these so called amateurs. Scientific soci-

eties miss young people and there is hardly a scientist who

Bonn zoological Bulletin 57 (1): 99-100

does not complain about lacking public awareness of his

or her field. Therefore the book is not only recommend-

ed for beginners but also to those who want to know how

to teach their ornithological knowledge and for those Non-

Ornithologists who look for stimulations how to prepare

the popular science aspect of their organisms in an exem-

plary way.

Darius Stiels

Zoologisches Forschungsmuseum Alexander Koenig,

Bonn

©ZFMK

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Kottelat M, Whitten T, Kartikasari SN, Wirjoatmodjo S (1993)

Freshwater fishes of Western Indonesia and Sulawesi. Periplus

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Mayr E (2000) The biological species concept. Pp. 17-29 in: Wheel-

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Parenti RP (2008) A phylogenetic analysis and taxonomic revision

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‘Oo OND

Contents ened det

Sonnenberg, Rainer & Eckhard Busch: 3

Description of Callopanchax sidibei (Nothobranchiidae: Epiplateinae),

a new species of killifish from southwestern Guinea, West Africa

Nguyen, Quang Truong, Andreas Schmitz & Wolfgang Bohme: 15

Gekko ulikovskii Darevsky & Orlov, 1994: a junior synonym of

Gekko badenii Szczerbak & Nekrasova, 1994

Krause, Patrick & Wolfgang Bohme: 19

A new chameleon of the Trioceros bitaeniatus complex

from Mt. Hanang, Tanzania, East Africa (Squamata: Chamaeleonidae)

Ullenbruch, Klaus, Olaf Grell & Wolfgang Bohme: 31

Reptiles from southern Benin, West Africa, with the description of

a new Hemidactylus (Gekkonidae), and a country-wide checklist

Wilhelm, Gertha, Hans L. Nemeschkal, John Plant & Hannes F. Paulus: 55

Fitness components in the relationship between

Rhopalapion longirostre (Olivier, 1807) (Insecta: Coleoptera: Apionidae) and

Alcea rosea (Linnaeus, 1758) (Malvaceae).

Analysis of infestation balance of a herbivorous weevil and its host plant

Shoorcheh, Hossein Rajael: 65

Life-history of Gnopharmia kasrunensis Wehrli, 1939 and

G. colchidaria Lederer, 1870 (Geometridae, Ennominae)

and their distribution in Iran, with first host-plant records for the genus

StUning, Dieter: 75

Two new species of the Plutodes costatus-group

from the Philippines and Indonesia (Lepidoptera, Geometridae, Ennominae)

Sinclair, Bradley J.: 85

Proclinopyga ulrichi sp. nov.: the first fossil aquatic dance fly

of the subfamily Clinocerinae (Diptera: Empididae)

Saikia, Munna Rani, Lakshmi Kanta Hazarika, Surajit Kalita & Ramen Lal Baishya: 91

Antennal sensilla of the rice hispa Dicladispa armigera (Olivier, 1808)

(Coleoptera: Chrysomelidae)

Book Reviews

Funfstlck HJ, Ebert A & Weiss | (2010) 18

Taschenlexikon der Vogel Deutschlands

(Kathrin Schidelko)

Moning C, Griesohn-Pfleger T & Horn M (2010) 99

Grundkurs Vogelbestimmung: Eine EinfUhrung in die Beobachtung und

Bestimmung unserer heimischen Végel

(Darius Stiels)

Cover illustration:

Callopanchax sidibei Sonnenberg & Busch

(this volume, pp. 3-14)

Ministerium fiir Innovation, ray . Fs

Y, Wissenschaft, Forschung Lei bn 1Z

Yi) und Technologie des Landes 5

<8S/_ Nordrhein-Westfalen @ Gemeinschaft