Bonn zoological Bulletin 66 (2): 95-106

December 2017

First record of Chrysotoxum baphyrum Walker from West Palaearctic

(Diptera: Syrphidae), with key to species of Chrysotoxum Meigen from Iran

Farzaneh Kazerani', Ali Asghar Taleb? & Ximo Mengual**

' Research Institute of Forests and Rangelands, Agricultural Research Education and Extension Organization (AREEO), Tehran,

Tran

? Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, RO.Box: 14115-336, Tehran, Iran

* Zoologisches Forschungsmuseum Alexander Koenig, Leibniz-Institut fiir Biodiversitat der Tiere, Adenauerallee 160,

D-53113 Bonn, Germany

“Corresponding author. E-mail: x.mengual@ leibniz-zfmk.de

Abstract. A new species of Syrphidae is recorded from Iran, Chrysotoxum baphyrum Walker. This record represents a

new species for the Western Palaearctic Region, as C. baphyrum is predominantly distributed in India and Southeast Asia.

An identification key to the recorded species of Chrysotoxum from Iran is provided, as well as newly obtained DNA se-

quences for C. baphyrum.

Key words. Iran, taxonomy, identification key, new record, DNA barcode.

INTRODUCTION

Flies of the genus Chrysotoxum M eigen, 1803 are excel-

lent wasp mimics, with long antennae (usually as long as

or longer than face), a broad convex abdomen, and a very

distinct black and yellow colour pattern. There are many

observations on habitat and visited flowers by adults

(Speight 2016) and on oviposition (Reemer & Goudsmits

2004), but the precise prey of larvae of Chrysotoxum re-

mains unclear. Some previous authors have reported im-

mature stages of Chrysotoxum near or in ant nests (Dixon

1960; Speight 1976; Rotheray et al. 1996), suggesting that

immatures may live in ant nests and feed on aphids. Oth-

er observations reported a similar behaviour (Inouye 1958;

Luciano et al. 1989), but the truth is that developmental

stages for the majority of species are unknown. Recent-

ly, Patil et al. (2013) reported the species, Chrysotoxum

baphyrum Walker, 1849 feeding on the sugarcane root

aphid from northern Karnataka State, peninsular India.

Chrysotoxum species are so distinct from other genera

that previous workers have given different ranks, i.e. as a

subfamily (Newman 1834; Schiner 1864; Brunetti 1923,

or Hull 1949 among other), as a tribe under Syrphinae

(Vockeroth 1969), or a subtribe due to its ‘aberration’

(Shatalkin 1975). Currently, Chrysotoxum is placed with-

in the tribe Syrphini (Vockeroth 1992; Mengual et al.

2008) without a clear sister group (Rotheray & Gilbert

1999; Mengual 2015; Mengual et al. 2015). Shannon

(1926) divided the genus into two subgenera based on the

ratio of the antennal segments, but subsequent authors

have not followed his subdivision.

Received: 05.02.2017

Accepted: 13.07.2017

In the Palaearctic Region, there are 87 recognized

species of Chrysotoxum at present, from a total of 143 pub-

lished names (Peck 1988; Ghorpadé 2012; NedeljkoviC et

al. 2013, 2015; Thompson 2013; Vujic etal. 2017). There

is currently great need for a revision of the species belong-

ing to this genus. Speight (2016) pointed outa fact already

mentioned by Coe (1953) and Sommaggio (2001), that

“the male genitalia of many European ‘species’ of

Chrysotoxum are equally indistinguishable”. As a conse-

quence, new techniques, molecular data and integrative ap-

proaches are being applied to separate species (M asetti et

al. 2006; NedeljkoviC et al. 2013, 2015) and new species

are described, mostly from China (Huo & Zheng 2004;

Huo etal. 2006; Zhang et al. 2010; Yuan etal. 2011). But

is. still needed» The

Palaearctic species were last revised by Violovitsh (1974),

a sound taxonomic background

but he did not include all the recognized species at the time

and separated females and males in different keys as both

sexes were not known for all species.

The syrphid fauna of Iran has been actively studied dur-

ing the last years (Dousti & Hayat 2006; Gilasian 2007;

Naderloo et al. 2013; Naderloo & Pashaei Rad 2014), and

as aresult some new species (Gilasian & Sorokina 2011;

Gilasian et al. 2015) and multiple new records (Gharali

& Reemer 2008, 2010; Khaganinia 2010, 2011; Bedoreh

& Ansari 2012; Kazerani etal. 2013, 2014a, 2014b; Kha-

ganinia & Hosseini 2013; Vosughian et al. 2013; Kha-

ganinia & Kazerani 2014; Khosravian et al. 2015) have

been reported. In Iran, Dousti & Hayat (2006) reported

12 species of Chrysotoxum in their catalogue, and more

recently, Kazerani et al. (2013) added three more species.

Corresponding editor: B. Sinclair

96 Farzaneh Kazerani et al.

Fig. 1. Map the known geographical distribution of Chrysotoxum baphyrum.

VujiC et al. (2017) described a new species from the Mid-

dle East, Chrysotoxum persicum VujiC, NedeljkoviC &

Hayat, 2017, which is also present in Iran. The present

study reports a new species record of this genus for West

Palaearctic, Chrysotoxum baphyrum. In addition, an

identification key is provided for the recorded Iranian

species of this genus.

MATERIALS & METHODS

Morphological terminology follows Violovitsh (1974) and

Thompson (1999a). In the material examined section, the

use of ellipses follows standard English practice and mere-

ly indicates that the missing information is the same as

that in the preceding record. At the end of each record, be-

tween square brackets ([]) and separated by commas, the

number of specimens and sex, the holding institution, and

the unique identifier or number are given. The abbrevia-

tions used for collections and their equivalents are given

below:

TMUI_ Department of Entomology, Tarbiat M odares

University, Tehran, Iran

ZFMK _ Zoologisches Forschungsmuseum Alexander

Koenig, Bonn, Germany

Photographs were composed using the software Zerene

Stacker® 1.04 (Richland, Washington, USA), based on im-

Bonn zoological Bulletin 66 (2): 95-106

ages of pinned specimens taken with a Canon EOS 7D®

mounted on a P—51 Cam-Lift (Dun Inc., VA, USA) and

with the help of Adobe Lightroom® (version 5.6). Figure

1 was created with the help of SimpleM appr (Shorthouse

2010).

The identification key is based on previous works; i.e.,

Becker (1921), Sack (1932), Violovitsh (1974), Sommag-

gio (2001), van Veen (2006), and Speight (2016), with the

help of the original descriptions and species checklist. The

collected material has been compared with the collections

at TMUI, ZFMK and other major natural history muse-

ums (USNM: National Museum of Natural History, Wash-

ington DC; BMNH: The Natural History Museum, Lon-

don; RMNH: Naturalis Biodiversity Center, Leiden), and

all the recorded species from the literature have been in-

cluded.

RESULTS

During the course of the survey to provide an initial tax-

onomic insight on the genus Chrysotoxum for Iran,

M alaise traps were used during 2010-2012 to collect spec-

imens from different habitats in several Iranian provinces.

Among the available material, five female specimens col-

lected in the southern Hormozgan province were studied

(Fig. 1). They were identified as Chrysotoxum baphyrum

(see Ghorpadé 2012 for the justified emendation of the

original name baphyrus), a widespread species on the In-

©OZFMK

New Chrysotoxum records from Iran 97

Figs 2-7. Chrysotoxum baphyrum Walker, female ZFM K -DIP-00018044. 2 — lateral view. 3 — dorsal view. 4 — frontal view. Chryso-

toxum bicinctum (Linnaeus), female ZFMK-DIP-00018047. 5 — dorsal view. Chrysotoxum parmense Rondani, male ZFM K-DIP-

00017469. 6 — dorsal view. 7 — frontal view. Scale bars = 1 mm.

Bonn zoological Bulletin 66 (2): 95-106 ©ZFMK

98 Farzaneh Kazerani et al.

dian subcontinent, representing the first record of this In-

domalayan

Brunetti (1923), Ghorpadé (1994) and Huo et al. (2007)

were used to key out these specimens, which agreed with

species in the West Palaearctic Region.

the original description and cannot be keyed out using cur-

rent Palaearctic identification keys. The type material of

C. baphyrum was not studied. The material of C. ba-

phyrum was then compared with specimens at the ZFMK

collected in India and Pakistan. Although only five females

were collected, these can be without doubt determined as

C. baphyrum by some morphological characteristics

unique to this taxon, such as basoflagellomere long (longer

than scape and pedicel together), gena yellow, scutum with

a complete lateral yellow vitta from postpronotum to

scutellum, and abdominal pattern reduced, less dark than

other species (see Differential diagnosis below).

Chrysotoxum baphyrum Walker, 1849

Chrysotoxum baphyrus Walker, 1849: 542. Type locality:

North Bengal, between India and Bangladesh.

Figs 2-4

Differential diagnosis. Scape and pedicel light brown to

orange with black hairs dorsally, basoflagellomere dark-

er, black, about 1.5 times as long as scape and pedicel to-

gether (Figs 2,4). Scutum with two dorsomedial, long yel-

lowish pollinose vittae almost reaching the posterior mar-

gin (Fig. 3), with acomplete lateral yellow vitta. Abdomen

mostly orange, dark markings reduced; lateral margin of

abdomen yellow; terga 2—5 each with a pair of medial,

broad, paler maculae, with traces of black fasciate macu-

lae on posterior margin of the pale maculae (Fig. 3).

Material examined. IRAN: Hormozgan province, Ban-

dar Abbas, Zakin, 27°49'37" N 56°18 24” E, 1122 m,

12.11.2012, A. Ameri [1 oO EM UEs..¢ LoavZol2 ty a2

9, TMUI] ..., vi.2011, ... [2 o. ZFMK, ZFMK-DIP-

00018044, ...18045].

Genetics. The GenBank accession numbers for the spec-

imen ZFM K-DIP-00018045 (Lab code ZFM K_D278) are:

28S gene (K Y315697), 18S gene (KY315696), and COI

gene (K Y315698). This is the first time that C. baphyrum

is barcoded, but we hope that more barcodes become avail-

able (especially from near the type locality) in the near fu-

ture and further analyses and comparisons can be done.

Distribution. The type locality of C. baphyrum is N orth

Bengal, a term used for the north-western part of

Bangladesh and northern part of West Bengal in India. The

the Rajshahi

Rangpur Division. This species has been previously

recorded from China (Huo et al. 2007), Pakistan, Sri Lan-

ka, India (Arunachal Pradesh, Chandigarh, Himachal

Bangladesh part denotes Division and

Bonn zoological Bulletin 66 (2): 95-106

Pradesh, Jammu & Kashmir, Karnataka, Maharastra,

Meghalaya, Punjab, Tamil Nadu, Uttarakhand, North Ben-

gal, West Bengal, Uttar Pradesh), Bangladesh, Nepal,

M yanmar, Laos, and Indonesia (Java is mentioned in the

literature but might be present in other regions, although

not recorded eastwards of the Wallace line) (Ghorpadé

1994, 2012, 2014; Huo et al. 2007; Thompson 2013; Mi-

tra et al. 2015).

Key to Chrysotoxum species in Iran

This identification key is based on recorded species from

Iran in the literature plus our new records. The images to

illustrate the key may not be from Iranian specimens.

1 Antennal basoflagellomere long, as long as or longer

than scape and pedicel combined (Figs 2, 4) ...... 13

- Antennal basoflagellomere short, distinctly shorter

than scape and pedicel combined (Figs 5, 7, 9, 11)

2 Median black facial vitta incomplete, not reaching an-

tennal bases; only basal 2/3 of face length occupied

by black vitta, not present on dorsal 1/3 (abdomen as

in Fig. 8)... 00.0.0... 2. Cyrobustum Portschinsky

—- Median black facial vitta complete, reaching or near-

ly reaching-base of antennae (Fre). 7) «20.25 bee leeessee GD

3. Abdomen with distinct yellow fasciate maculae on ter-

ga 2 and 4 only; tergum 3 entirely black or with nar-

row yellow markings (less than half width of those on

tergum 4) (Fig. 5)... ......... C. bicinctum (Linnaeus)

- Abdominal terga 2-5 with yellow, usually interrupt-

ed, fasciate maculae of equal width; tergum 3 with yel-

low markings as wide as those on tergum 4 (Figs 12,

4 Frons yellow (Figs 6,7); scutum shiny black with two

dorsomedial broad pale pollinose vittae that almost

reach scutellum (Fig. 6) ......... C. parmense Rondani

— Frons black (Figs 11,13); scutum shiny black with two

dorsomedial narrow pale pollinose vittae, usually end-

ing at transverse suture or after, but never beyond an-

terior 2/3 of scutum (Figs D224) aos thon cette aeelnes ans

5 Anterior part of frons projecting prominently upwards

(Fig. 9); yellow markings on terga 2-4 very similar,

not arcuate, with anterior margin of fasciate vittae par-

allel to anterior margin of tergum (Fig. 10)

wee. C. coreanum Shiraki

— Frons of usual shape, anterior part of frons not pro-

jecting upwards (Fig. 11); yellow markings on terga

2-4 somehow arcuate (Figs 12, 14) ... 0. ee. 6

6 Lateral margin of abdomen alternately black and yel-

(Pek igg (Sau sacits Be: ANG SH a Ries LD kee Eee as ea. tec eee ae

— Lateral margin of abdomen black, yellow fasciate vit-

fac-noeRieachine im ane ia Ce aes We) es karst. been 7

©OZFMK

New Chrysotoxum records from Iran 99

Hae AG i

hi il NK

Figs 8-13. Chrysotoxum robustum Portschinsky. 8 — abdomen (from Violovitsh 1974). Chrysotoxum coreanum S hiraki. 9 — head,

lateral view (from Violovitsh 1974). 10 — abdomen (from Violovitsh 1974). Chrysotoxum festivum (Linnaeus), female ZFM K-DIP-

00017473. 11 —- lateral view. 12 — dorsal view. Chrysotoxum vernale Loew, female ZFM K-DIP-00017476. 13 — lateral view. Scale

bars = 1 mm.

Bonn zoological Bulletin 66 (2): 95-106 ©ZFMK

100 Farzaneh Kazerani et al.

Figs 14-19. Chrysotoxum octomaculatum Curtis, female CEUA00019589. 14 — dorsolateral view. Chrysotoxum octomaculatum

Curtis, female CEUA00082597. 15 — abdomen, ventral view. Chrysotoxum persicum VujiC, NedeljkoviC & Hayat, paratype female

ZFMK-DIP-00018048. 16 — abdomen, dorsolateral view. 17 — wing and abdomen. 18 — abdomen, ventral view. Chrysotoxum ver-

ralli Collin, female ZFM K-DIP-00017471. 19 — dorsal view. Scale bars = 1 mm.

Bonn zoological Bulletin 66 (2): 95-106 ©ZFMK

New Chrysotoxum records from Iran 101

Figs 20-25. Chrysotoxum elegans Loew, female ZFM K-DIP-00017470. 20 — dorsal view. Chrysotoxum montivagum V iolovitsh,

female ZFMK-DIP-00018049. 21 — abdomen, dorsal view. 22 — wing. Chrysotoxum sp., female from Qazi Chak, ZFMK-DIP-

00018050. 23 — dorsal view. Chrysotoxum cautum (Harris). 24 — male ZFMK-DIP-00017464, lateral view. 25 — female ZFMK-

DIP-00017468, posterior view. Scale bars = 1 mm.

Bonn zoological Bulletin 66 (2): 95-106 ©ZFMK

102 Farzaneh Kazerani et al.

Figs 26-31. Chrysotoxum arcuatum (Linnaeus). 26 — female ZFM K-DIP-00018046, lateral view. 27 — male ZFM K-DIP-00017472,

dorsal view. Chrysotoxum bactrianum V iolovitsh, female. 28 — dorsal view. Chrysotoxum lessonae Giglio-Tos, male ZFM K -DIP-

00017475. 29 — dorsal view. 30 — ventral view. Chrysotoxum intermedium M eigen, male ZFM K-DIP-00017474. 31 — ventral view.

Scale bars = 1 mm.

Bonn zoological Bulletin 66 (2): 95-106 ©ZFMK

New Chrysotoxum records from Iran 103

7 Pro- and mesofemur completely yellow (Fig. 11);

katepisternum usually with dorsal yellow macula (Fig.

{Bile fede die he A ene Oy, Woes ee te C. festivum (Linnaeus)

— At least bases of pro- and mesofemur black; katepis-

ternum usually without dorsal yellow macula; if pres-

ent, size smaller than posterior spiracle (Fig. 13)

wie. C. vernale Loew

8 Terga 3 and 4 with black anterior margin not interrupt-

ed, continuously black. Tergum 5 with anterior black

fascia reaching lateral margin; if not, lateral margin

CEO We CEG yn Pek utc ae gms ce dertated Sak aerherebern sie san

— Terga 3 and/or 4 with black anterior margin interrupt-

ed at lateral margin by yellow vitta; lateral margin part-

ly black (Figs 14, 16); although variable, most speci-

mens exhibit this feature on at least one tergum. Ter-

gum 5 with anterior black fascia also interrupted,

sometimes lateral margin entirely yellow (Fig. 14)

9 Wing with anterior margin yellow, without dark mac-

ula; wing tip hyaline (Fig. 14). Terga 3 and 4 with black

anterior margin broader towards lateral margin because

anterior margin of yellow maculae does not follow ter-

gum anterior margin (Fig. 14); sternum 2 black, ex-

cept posterolateral corners yellow, with two anterior

large yellow maculae (joined medially or not) occu-

pying at least 1/3 of sternum length (Fig. 15). Male

genitalia: surstylus less elongated, about 3 times longer

wee C. octomaculatum C urtis

— Wing with anterior margin yellow, with clear dark mac-

than wide ...... .......

ula between end of Ri and undulation of Ras; wing tip

hyaline (Fig. 17). Terga 3 and 4 with black anterior

margin very narrow, of equal width over much of its

length because anterior margin of yellow macula fol-

lows tergum anterior margin closely (Figs 16, 17); ster-

num 2 black, except posterolateral corners yellow, with

two very small anterior yellow maculae occupying less

than 1/4 of sternum length (Fig. 18). Male genitalia:

surstylus elongate, about 4 times longer than wide

veces. Cy persicum Vujié, Nedeljkovié & Hayat

10 Wing with darkened anterior margin, yellow, without

ca ateia stale

— Wing with darkened anterior margin, yellow to brown-

subapical dark macula (Fig. 19)... 2...

ish, with subapical dark macula (Fig 22)............ 11

11 Scutellum yellow with medial black macula (Figs 21,

22) ce cee eee te ee ee ee Co montivagum V iolovitsh

—- Scutellum black with posterior margin yellow (Fig. 23)

.. a... Chrysotoxum sp. (female from Qazi Chak)

12 Tergum 2 with black anterior margin broader towards

lateral margin because anterior margin of yellow mac-

ulae does not follow anterior margin of tergum (Fig.

POY se oh. uth fh Deed ahi ete, Shah ae CLES ONS Doew

— Tergum 2 with black anterior margin of equal width

over much of length because anterior margin of yel-

low macula follows closely anterior margin of tergum,

separated only at side margin (Fig. 19)

Bonn zoological Bulletin 66 (2): 95-106

Lhe ich hs ie BI a wie iC. verralli C ollin

13 Gena yellow, without black vitta (Figs 2, 4); mesono-

tum with lateral yellow vitta complete, uninterrupted

(Fig. 3)... cc cee cee ee ee cee eee eee» Cy baphyrum W alker

— Gena with black vitta (Figs 24, 29); mesonotum with

lateral yellow vitta interrupted at transverse suture

CRIS BAA) oe en Mean cals ati es 2. i aes news 5 EG

14 Male: genitalia very large, protruding ventrally in lat-

eral view; postabdomen (abdominal segments poste-

rior of sternum 4) greatly enlarged (Fig. 24). Female:

tergum 6 with longitudinal, median membranous slit

(Fig. 25) oc cee cee cee cee cee see vee vee ee Cy cautum (Harris)

—- Male: genitalia smaller, not visible in lateral view;

postabdomen not enlarged (Figs 30, 31). Female: ter-

cum 6 withoutmen-branous st... 6 ares ees phen ILD

15 Abdominal terga with long, erect pile, as long as or

longer than scape; abdomen relatively short and broad

(Figs 26, 27) ... eee ee ee. Cy arcuatum (Linnaeus)

—- Abdominal terga with short, adpressed pile, some ar-

easamray loo kabaresCh ia. 20 late, kan tt andy, logs. yk

16 Scutellum entirely pale yellow (Fig. 28); anterior mar-

gin of wing yellowish-brown with apical dark macu-

La cee ee cee tee ee cee ee ee ee ee Co bactrianum V iolovitsh

—- Scutellum yellow with relatively small translucent,

transparent macula in middle (Fig. 29); anterior mar-

gin of wing yellowish-brown without clear apical dark

meacurla: CRUG 9292): cots ae eeel veuae ce addon eee aetlessireecs eo

17 Sterna: yellow fasciate maculae originated on postero-

lateral corner of sternum 2 do not join medially (Fig.

BO eee Shad i oe A eSSOnde Gisliogl os

— Sterna: yellow fasciate maculae originated on postero-

lateral corner of sternum 2 broadly joint medially

forming fascia (Fig. 31)... ... C. intermedium M eigen

Note: C. intermedium is a species complex (Speight

2016), and the current definition might include more

than one species. Wings of some specimens have a

dark brown apical macula.

DISCUSSION

Although more data is available to establish boundaries

between biogeographic regions, climate change and hu-

man actions alter the ecosystems and their fauna. M ore-

over, researchers use certain geographical or political

boundaries to delimit faunal or biogeographical regions

with considerable controversy (Mengual 2012). Thomp-

son (1999b) illustrates the biotic regional boundaries based

on political subunits, although this was not accurate. One

of the few countries across biogeographical boundaries is

China. Huo et al. (2007) reported several specimens of C.

baphyrum from Cili, Hunan province (Poa 4ge7 ON

111°08/17.6” E). Hunan province is considered Palaearc-

tic by Thompson (1999b), thus our records would not be

the first for the Palaearctic Region.

©OZFMK

104 Farzaneh Kazerani et al.

But other divisions exist, such as Holt et al. (2013),

which includes Hunan province in their Sino-Japanese

zoogeographic realm. On the other side, Hoffman (2001)

mentions a transition zone between Palaearctic and Indo-

malayan Regions, which resembles the Sino-Japanese

realm of Holt et al. (2013), and Hunan province is includ-

ed in this transition zone. The western boundary between

Palaearctic and Indomalayan regions has more agreement,

and somehow divides north-south Pakistan. Based on all

the available divisions, Iran can be considered Palaearc-

tic, making our records the first specimens of C. baphyrum

collected from West Palaearctic. On the other hand, the use

of political or historical names in reference to regions or

realms should not be encouraged, such as Asia (Gilasian

et al. 2015).

The genus Chrysotoxum is distributed mainly in the Ho-

larctic Region, with a few species reported from tropical

areas (Thompson 2013) and probably found in higher al-

titudes (Shannon 1926). Iran is located in the Western

Palearctic Region, with some influences from the Indo-

malayan Region in the south-east, and it has a rich fauna

of insects, acting as a connection between the Palearctic

and Indomalayan faunas. The Hormozgan province,

where C. baphyrum was collected, and the whole south-

ern part of Iran are poorly sampled and understudied. So

far, no long sampling period and large-scale survey has

been done in forests and cultivated lands of Iran. Thus, it

would not be surprising to record more Indomalayan

species in Iran, or even new Species to science due to the

location of Iran, between biodiversity hotspots such as

Caucasus, Anatolian peninsula (as part of the Mediter-

ranean Basin) and the mountains of Central Asia. More

taxonomic studies and field work are needed in this coun-

try, and there is hope to pursue this goal in the next years

as more and more Iranian researchers focus on Syrphidae.

Acknowledgement. We thank the Department of Entomology,

Tarbiat Modares University for providing financial support for

this research. Our cordial thanks are expressed to A. Nadimi and

M. Kheyrandish for collecting some of the studied specimens

in this research.

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Bonn zoological Bulletin 66 (2): 107—133

December 2017

The snakes of Mali

Jean-Francois Trape’" & Youssouph M ané'

' Institut de Recherche pour le Développement (IRD), Laboratoire de Paludologie et de Zoologie Médicale, UMR MIVEGEC,

B.P. 1386, Dakar, Sénégal

“Corresponding author. E-mail: jean-francois.trape @ ird.fr

Abstract. From 2004 to 2010 we sampled snakes in different localities in the Republic of Mali, West Africa. A total of

5,224 snakes belonging to 60 species were collected at 37 localities, including specimens from 18 species not reported

before from this country. Based on a full account of this collection, supplemented with additional museum specimens

and reliable literature reports, we present an annotated checklist of the 65 snake species currently known from M ali. Col-

lecting localities for all specimens are provided and, where necessary, some taxonomical and biogeographical issues are

discussed.

Key words. Reptilia; Ophidia; biogeography; country checklist; venomous snakes; West Africa.

INTRODUCTION

The Republic of M aliis the second largest country of West

Africa with 1,240,000 km’ between latitudes 10°N and

25°N and longitudes 12°W and 4°E. The northern part of

the country is Saharan (Fig. 1), the central part is Sahe-

lian (Fig. 2), and the southern partis Sudanese or Sudano-

Guinean (Figs 3-4). Elevation is low, ranging from 30 m

along the banks of the Senegal River in the westernmost

part of the country, to 1,155 m at the top of the highest

peak inthe HomboriM ounts in central M ali (Fig.5). Even

in the most elevated areas of the country, i.e. the

M andingue and Dogon Plateaux in south-eastern and cen-

tral Mali, and the Adrar des Iforhas in northern M al1, el-

evation rarely exceeds 500 m. The country has a rich hy-

drological system, with the Niger River and its tributar-

ies that form a large inner delta in central Mali before

reaching the Sahara desert then heading southeast to the

Gulf of Guinea, and with four main rivers in the upper

Senegal River basin in the western part of the country (Ar-

naud & Pitte 2010).

Small collections of snakes from M ali, formerly Soudan

francais, have been reported by Pellegrin (1909), Cha-

banaud (1917), Angel (1922, 1933), de Witte (1930), An-

derson (1935), Angel & Lhote (1938), Villiers (1950,

1951,1952, 1953,1954, 1956, 1965), Joger (1981), Schat-

ti (1986), and Bohme et al. (1996). Papenfuss (1969) and

Joger & Lambert (1996) reviewed the literature recording

snake and other reptile species in M ali, reported their own

collections, and provided a country checklist with local-

ity records for each species. Specimens from Mali were

included in works on certain genera or species, in partic-

ular by Roux-Estéve (1974), Hughes (1976, 1977, 1983,

1985), Broadley (1984), W iister & Broadley (2003), Trape

& M ané (2006a), Trape et al. (2006, 2009, 2012), and Cro-

Received: 09.03.2017

Accepted: 14.09.2017

chetetal. (2008). Trape & Mané (2006b) provided square-

degree distribution maps for all species of snakes known

from Mali and neighbouring countries, including a large

part of the material here reported, but precise collecting

localities were not mentioned. Sindaco et al. (2013) pro-

vided of Palaearctic

square-degree distribution maps

species distributed in northern Mali.

MATERIAL AND METHODS

From July 2002 to March 2010, we conducted field work

in several regions of Mali in order to collect snakes. We

deposited cans or buckets half filled with formaldehyde

or ethanol in 32 villages (Fig. 6), which were housed by

the chief of the village. We asked the villagers to deposit

in these containers snakes they killed when they were oc-

casionally encountered in the vicinity of their village. A

modest award (300 CFA, i.e. approximately 0.6 US $) was

given for each preserved specimen. In most parts of Mali,

as in most parts of Africa, all species of snakes are feared

and systematically killed when they are encountered. Thus,

the objective of the award was to acknowledge the effort

of carrying killed snakes from surrounding fields to the

village, but without encouraging snake research and

killing. Visits to the villages were organized in A pril 2003,

January, June and December 2004, February, June and No-

vember 2005, January and May 2006, January, October

and December 2007, January 2008, and March 2010, to

retrieve the specimens. During these trips we also collect-

ed snakes at five additional localities. The 37 collecting

localities (Table 1) were distributed either in the southern

part of the country (7 localities between 11°00’N and

11°59’N), in the

12°00’N and 14°15’N) orin the northern part of the coun-

central part (13 localities between

Corresponding editor: Ph. Wagner

108 Jean-Francois Trape & Youssouph Mané

Fig. lA typical sandy valley and wadi in the mountains of the Adrar des Iforas (19°01’°N,01°50’E).

Fig. 2. View of the Sahelian vegetation in western Mali, near Samé Ouolof study village (14°33’N,11°20’W).

Bonn zoological Bulletin 66 (2): 107-133 ©ZFMK

The snakes of Mali 109

Fig. 3. Sudanese savanna in eastern Mali, west of Niamasso study village (13°02’N, 05°47’W ).

try (17 localities between 14°16’N and 16°45’N), where

average annual rainfall

1,000-1,100 mm, 500-1,000 mm, and 200-500 mm, re-

ranges approximatively from

spectively (M ahé et al. 2012). The most Saharan part of

the country, i.e. north of 17°N, was not surveyed except

during a two-week period in February 2004. Although no

specimen was collected during these two weeks, some in-

formation on records of Cerastes cerastes Linnaeus, 1758

and C, vipera (Linnaeus, 1758) was obtained from locals

(they are indicated as “sight record”).

M ost

specimens were deposited at the Institut de

Recherche pour le Développement (Dakar, Senegal;

acronym: IRD) and several specimens were donated to the

Museum national d’Histoire naturelle (Paris, France;

acronym: MNHN). We also examined selected M alian

specimens from the Institut Fondamental d’A frique N oire

(Dakar, Senegal; acronym: IFAN) and atthe MNHN col-

lections.

Specimens were identified to species according to clas-

sical identification keys for West African snakes (Villiers

& Condamin 2005, Trape & Mané 2006b) and further tax-

onomic analysis (Trape etal. 2009, Trape etal. 2012, Trape

et al., unpublished). For recent changes in snake generic

names, we usually followed those adopted in the reptile

database by Uetz et al. (2016) (http://www.reptile-data-

base.org/).

Bonn zoological Bulletin 66 (2): 107-133

RESULTS

We collected a total of 5,224 specimens belonging to 60

species (Table 1) and examined additional M alian speci-

mens from IFAN (one specimen) and MNHN (13 speci-

mens). Two additional species are known with certainty

from M ali but were not represented among the specimens

that we collected or examined personally. Altogether, the

total number of snake species from Mali is 65. Coordi-

nates of collecting localities are listed in Table 1, and those

obtained from literature data and sight records are in Table

Family Typhlopidae Gray, 1845

Afrotyphlops lineolatus (Jan, 1864)

M aterial: 12 specimens.

Localities: Npiébougou (11), Laminina (1).

Remarks: First record for M ali.

Afrotyphlops punctatus (Leach, 1819)

M aterial: 21 specimens.

Localities: Laminina (1), Mamoroubougou (7), Niamou

(2), Npiébougou (1), Titiéna (4), Zamoko (6).

Literature record: Diafarabé (Villiers 1953).

Remarks: All specimens from Mamoroubougou and one

©ZFMK

110

Table 1. Collecting localities of snakes in M ali (our study).

Jean-Francois Trape & Youssouph M ané

N Locality Latitude Longitude Altitude Ecoregion No.of specimens No.of species

1 A goudoud 15°54’°N 01°13’°W 320 m Sahel 4 2

2 Ballabougou 12°52’N 06°52’°W 338 m Sudanese alg) 7

3 Bamako 12°38’°N 08°00’°W 325 m Sudanese 1 1

4 Bangaya 13°14’N 10°43’°W 135 m Sudanese 118 19

5 Boussouma 15°06’N 02°38’°W 318 m Sahel 19 4

6 Bouyanga 14°30’°N 09°38°>W 278 m Sahel 305 12

i Djinagué T2597 N. 09°52’°W 305 m Sudanese 125 17

8 Donguiba 13°49’N 06°05’W 303 m Sudanese 1 1

9 Doro 16°08’°N 00°50’°W 295 m Sahel 10 2

10 D oussoudiana 11°09°N 07°48’ W 365 m Sudano-Guinean 116 20

iB! Gaoudel 15.95.92 N 04°05°W 270 m Sahel 95 7

12, Gogui (10 km S) 15°35’N 09°20’°W 210 m Sahel 1 1

13 Gouina 14°00’°N 11°06’W 70 m Sudanese 1 1

14 H aoussa-Foulane 15°59’N 00°08’°E 256 m Sahel 7 4

15 Kinani 15°00’°N 03°51°W 268 m Sahel OMT, 2

16 Koundian 13°09°N 10°40’°W 170 m Sudanese 68 18

Lif Koyretao 16°04’°N 03°53’°W 271m Sahel 55 6

18 Laminina PS 127 \N 07°46’°W 361m Sudano-Guinean 160 28

LED) Léré 15°43°N 04°54’°N 268 m Sahel 6 3

20 M amoroubougou 11°14’°N 05°28’°W 386 m Sudano-Guinean 1064 36

21 Niakoni it er Lg 07°48’ W 378 m Sudano-Guinean 14 7

22 Niamasso 12°59’N 05°27°W 281 m Sudanese 21 >

23 Niamou 14°O1’°N 08°02°W 372 m Sudanese 498 23

24 N piébougou P17 2:5:9)" 'N 08°00’°W 367 m Sudano-Guinean 372 29

25 Sadjouroubougou 12:23:55-N 07°44’ W 335 m Sudanese 26 10

26 Samé Ouolof 14°29’°N 11°34’°W 41m Sahel Hil 8

27 Saré-Soma 14°45°N 03°55’°W 271 m Sahel 31 4

28 Sébékourani P2712" 08°42°W 386 m Sudanese 409 28

29 Séoulasso 13°14’°N 04°42’°W 279 m Sudanese 292 16

30 Tacharane 16°09’°N 00°04’°E 257 m Sahel 15 2

3 dl Ténintou 11°20’N 07°44°W 357 m Sudano-Guinean 1 1

3x2 Tinjem ban 16°44’°N 02°50’°W 269 m Sahel 233 4

33 Titiéna PLES Y6ON 06°33’°W 308 m Sudano-Guinean 376 2-4

34 Topokhoné 15°02’N 10°34’W 105 m Sahel 4 3

35 Toum boula 14°20°N 07°47°W 287 m Sahel 170 9

36 Toya 16°39’N 03°03’°W 266 m Sahel 71 3

37 Zamoko 13°09°N 07°57°W 395 m Sudanese 622 25

Total 5,224 60

specimen from Laminina and Titiéna were marbled, the

rest were lineated with three of them blackish (all from

Zamoko).

Family Leptotyphlopidae Stejneger, 1892

Localities: Doussoudiana (1), Mamoroubougou (4).

Remarks: First record for M ali.A picture of the specimen

from Doussoudiana was published in Trape & Mané

(2006b).

Myriopholis algeriensis (Jacquet, 1895)

M aterial: no specimen studied.

Myriopholis adleri (Hahn & Wallach, 1998)

M aterial: | specimen.

Locality: Sébékourani (1).

Remarks: First record for Mali.

Myriopholis albiventer (Hallermann & Rédel, 1995)

M aterial: 5 specimens.

Bonn zoological Bulletin 66 (2): 107-133

Literature records: Bourem (Papenfuss 1969, as Leptoty-

phlops macrorhynchus (Jan, 1860); Hahn & Wallach 1998,

as Leptotyphlops macrorhynchus algeriensis), Tom bouc-

tou (Broadley et al. 2014).

Myriopholis boueti (C habanaud, 1917)

M aterial: 126 specimens.

©OZFMK

The snakes of Mali 111

Table 2. Coordinates of snake records from Mali (literature data and sight records). The asterisk means an approximate.

Locality Latitude Longitude

Adrar des Iforhas 18-20°N* 01-02°E*

A raouane 18°54’N 03°31°W

Bamako 12°39’°N 08°00°W

Bandiagara 14°21°N* 03°37°W *

Bla 12°57°N 05°46’°W

Bougouni 11°25’N 07 32:0-°W:

Bourasso 13°40’°N 04°20°W

Bourem 16°57’°N 00°21’°W

Diafarabé 14°09’°N 05°01°W

Didiéni (toward Kolokani) 13°42’°N 08°02’W

Diré 16°16’N 03°24’°W

Djenné 13°54’N 04°33°W

Dogo 15°10’°N 04°26’°W

Douentza 15°00°N 02°57’°W

Fabiguine 16°44’°N* 03°50°W *

Fatao (9 km N of) 14°24’N 09°29°W

Félou (falls) 14°21’°N 115-201,” We

Gao 16°16’°N 00°03’W

Goundam 1-6'°230-N. 03°40’°W

Gourao 15°19’°N 04°02’W

Kakoulou 14°17’N 11°17’W

K ati 12°43’N 08°04’ W

Katibougou 12°30’N 08°05’°W

Kayo 1.33953." NV: 05°37°W

Ké-Macina 13°58’°N 05°23°W

K idal 18°26’N 01°24’E

K ita 13°03’N 09°29°W

Kokounkourou 12°58’°N 09°37°W

Lobi 16°15’N 00°04’ W *

M adina-Kagoro (10 km W of) 14°20’°N 07°45°W

M opti and vicinity 14°30’N 04°12°W

M ourdiah 14°28’°N 07°28°W

Naréna 12°14’N 08°38°W

Niamiga (W of) 14°30’N 11°12’°W

Nioro du Sahel 15°14’N 09°35°W

Négala (toward K assaro) LQEIS EN. 08°40’°W

San 13°18’°N 04°54°W

Saré M alé 14°05°N 04°26’W

Sévaré (158 km SW of) 13°40’°N 04°20°W

Sikasso (toward Sindi) 11°30’N 05°56’°W

Sormé 14°52’N 04°25’°W

Tabakoro 11°26’N 06°46’°W

Taga Diabozo 14°18’°N 04°57°W

Takabart (SW ) 20°10’N * O01°10°W *

Taoudeni (half way to Araouane) 20°40°N * 03°50°W *

Tessalit (and vicinity) 20°12’°N 01°00°W

Tilem baya 14°09°N 04°59°W

Tim étrine 19°20’°N 00°42°W

Tisserlitine (toward Tim étrine) 20°36’°N 00°11°>W

Toguéré Sanga 14°28°N 03°19°W

Tom bouctou 16°46’N 03°01’°W

Wana Boubou 14°O01°N 04°58°W

Y élimané 15°08’N 10°34’ W

Bonn zoological Bulletin 66 (2): 107-133 ©ZFMK

112 Jean-Francois Trape & Youssouph M ané

Seg ee

"+ _.

Fig. 4. Sudano-Guinean savanna in the vicinity of Doussoudiana, the southernmost village of the study (11°11’N,07°44’W ).

Fig. 5. The Hombori M ounts, the highest mountains of Mali (15°14’N, 01°48’°W ).

Bonn zoological Bulletin 66 (2): 107-133 ©ZFMK

The snakes of Mali 113

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Fig. 6. Map of Mali with location of villages selected for the study. See Table 1 for locality numbers. Doussoudiana, Laminina

and Niakoni villages, located within a5 km radius, are represented by a single number (n° 10). Colours for vegetation areas: Guinean:

dark green; Sudanese: green; Sahelian: light green; Saharan: yellow for sandy areas, white for stony areas, grey forrocky and moun-

tainous areas.

Localities: Bamako (1), Bouyanga (9), Djinagué (2), Remarks: This species is currently known only from

Koundian (1), Niamou (1), Npiébougou (27), Samé Ouolof Mamoroubougou, the type locality, where it represented

(1), Sébékourani (10), Zamoko (74). 1.5% of the 1,042 fossorial reptiles collected in this lo-

Literature records: Djenné (Chabanaud 1917, type local- cality (Typhlopidae: 7 specimens, Leptotyphlopidae: 731

ity), Bamako (Villiers 1950). specimens, Amphisbaenidae (Cynisca leucura (Dum éril &

Remarks: One specimen from Sébékourani was attributed Bibron, 1839)): 304 specimens).

erroneously to Myriopholis rouxestevde (Trape & Mané,

2004) in the distribution map of Trape & Mané (2006b). Rhinoleptus koniagui (Villiers, 1956)

M aterial: 11 specimens.

Rhinoguinea magna Trape, 2014 Localities: Djinagué (1), Koundian (1), Laminina (3),

M aterial: 16 specimens. Npiébougou (6).

Locality: Mamoroubougou (16). Remarks: First record for Mali. Koundian and Laminina

Bonn zoological Bulletin 66 (2): 107-133 ©Z7FMK

114 Jean-Francois Trape & Youssouph M ané

Fig. 7. The Farako River and its forest gallery near Mamoroubougou, the village where the highest number of snakes and snake

species were collected (11°14’°N, 05°28’°W ).

Fig. 8. The Sudanese savanna between Koundian and Bangaya villages in southwestern Mali (13°10’N, 10°38’°W ).

Bonn zoological Bulletin 66 (2): 107-133 ©ZFMK

The snakes of Mali 115

Fig. 9, The Sudanese savanna near $ébékourani village where 28 snake species were collected (12°02’N, 08°44’W ). The deadly

carpet viper Echis jogeri was abundant.

specimens were reported on a grid map in Trape & Mané

(2006b), where the square degree 1I1°N/5°W was men-

tioned erroneously.

Tricheilostoma bicolor (Jan, 1860)

M aterial: 838 specimens.

Localities: Doussoudiana (4), Laminina (1),

Mamoroubougou (711), Niamou (3), Npiébougou (7),

Sébékourani (9), Titiéna (2), Zamoko (101).

Literature records: Diafarabé (Villiers 1950), Bamako

(Villiers 1950, as Leptotyphlops brevicauda (B ocage,

1887)).

Remarks: The maximum total length was 191 mm (tail

length: 6 mm) inaspecimen from Mamoroubougou (IRD

4289.M), a record for the species. Smallest specimens

from Mamoroubougou were 9-10 cm long.

Family Boidae Gray, 1825

Eryx muelleri (B oulenger, 1892)

M aterial: 174 specimens.

Localities: Bangaya (1), Bouyanga (42), Doro (7),

Gaoudel (7), Haoussa-Foulane (1), Koundian (2), Koyre-

tao (8), Léré (3), Niamasso (1), Niamou (95), Samé Ouolof

(1), Séoulasso (1), Tinjemban (1), Toumboula (4).

Literature records: Bandiagara (Angel 1933), Gao (Angel

& Lhote 1938), Nioro du Sahel (Villiers 1950), Dogo (Vil-

liers 1956), Mourdiah (Joger & Lambert 1996).

Bonn zoological Bulletin 66 (2): 107-133

Family Pythonidae Fitzinger, 1826

Python regius (Shaw, 1802)

M aterial: 55 specimens.

Localities: Ballabougou (1), Djinagué (3), Doussoudiana

(10), Koundian (1), Laminina (7), Niakoni (1), Niamou

(1), Sadjouroubougou (1), Sébékourani (18), Titiéna (8),

Zamoko (4).

Literature record: Southern Mali (Chippaux 1999),

Python sebae (Gmelin, 1788)

M aterial: 23 specimens.

Localities: Bangaya (1), Boussouma (1), Bouyanga (1),

Djinagué (1), Laminina (3), Niamou (5), Npiébougou (1),

Samé Ouolof (1), Sébékourani (7), Titiéna (1), Toumboula

(1).

Literature records: Kati (Angel 1922), Sikasso (Villiers

1965), Bamako (Broadley 1984), Bandiagara (Béhme et

al. 1996).

Family Colubridae Oppel, 1811

Afronatrix anoscopus (Cope, 1861)

M aterial: 17 specimens.

Localities: Mamoroubougou (16), Titiéna (1).

Remarks: First record for Mali. Mamoroubougou record

was reported on a grid map in Trape & Mané (2006b).

©OZFMK

116 Jean-Francois Trape & Youssouph Mané

Fig. 10. Near Bandiagara cliff, an area of transition between the Sudanese savanna and the Sahel (14°02’N,03°46’W ).

Bamanophis dorri (Lataste, 1888)

M aterial: 37 specimens.

Bangaya (24), Koundian (4), Niamou (8),

Toum boula (1).

Literature records: Kati (Angel 1922), Bamako (Villiers

1956, Joger & Lambert 1996), chutes du Félou, between

Localities:

Négala and Kassaro (Joger 1981), Bandiagara (Joger &

Lam bert 1996).

Remarks: Most specimens where included in the descrip-

tion of the genus Bamanophis by Schatti & Trape (2008).

Crotaphopeltis hippocrepis (Reinhardt, 1843)

M aterial: 3 specimens.

Localities: Doussoudiana (1), Mamoroubougou (1),

N piébougou (1).

Remarks: First record for Mali. This species seems rare

in Mali contrary to Burkina Faso and Guinea, two neigh-

bouring countries where C. hippocrepis is both common

and widely distributed (Roman 1974, Trape & Baldé

2014).

Crotaphopeltis hotamboeia (Laurenti, 1768)

M aterial: 225 specimens.

Localities: Ballabougou (1), Bangaya (26), Djinagué (29),

D oussoudiana (12), Laminina (10),Mamoroubougou (25),

Niakoni (2), Niamasso (2), Niamou (8), Npiébougou (11),

Sadjouroubougou (2), Saré-Soma (2), Sébékourani (58),

Bonn zoological Bulletin 66 (2): 107-133

Séoulasso (1), Titiéna (22), Toumboula (5), Zamoko (9).

Literature record: Diafarabé (Villiers 1952).

Dasypeltis confusa Trape & Mané, 2006

M aterial: 3 specimens.

Locality: Diafarabé (1, coll. IFAN),M amoroubougou (2).

Remarks: The Diafarabé specimen was collected in the

early 1950’s by the French ichthyologist J. Dagetand was

first published as Dasypeltis scabra (Linnaeus, 1758) by

Villiers (1953). In fact, its pattern (SL) is typical of D. con-

usd (see Trape & Mané 2006a and Trape etal. 2012), and

there is no doubt that it belongs to this Guinean species

although Diafarabé is located in the Sahel at320 km north

of the nearest known locality (Mamoroubougou). Since

Diafarabé is riverine of the Niger River and only D. sa-

helensis Trape & M ané, 2006 is known in the area, we pre-

sume that this specimen was carried by floods, as it also

probably occurred for the holotype of Echis jogeri C her-

lin, 1990 from Tombouctou.

Dasypeltis gansiTrape & Mané, 2006

M aterial: 3 specimens.

Localities: Ballabougou (2), Séoulasso (1).

Remarks: The other records presented on the grid map in

Trape & M ané (2006b) are referable to D. latericia Trape

& Mané, 2006 (see below).

©OZFMK

The snakes of Mali Rg:

Fig. 11. A view of the Sahelian landscape in the vicinity of Boussouma study village (15°08’N, 02°31’W ). Psammophis aft. sibi-

lans, Naja nigricollis ana Echis leucogaster were the most abundant species, but Python sebae was also collected in a pound.

Dasypeltis latericia Trape & Mané, 2006

M aterial: 25 specimens.

Localities: Bangaya (4), Koundian (1), Niamou (5),

Npiébougou (2), Sébékourani (1), Séoulasso (4), Titiéna

(1), Zamoko (7).

Remarks: Most specimens were included in the descrip-

tion of D. latericia (Trape & Mané 2006a). Dasypeltis la-

tericid was recently given full species rank following amo-

lecular analysis of the genus Dasypeltis (Trape etal. 2012).

Dasypeltis sahelensis Trape & Mané, 2006

M aterial: 23 specimens.

Localities: Bouyanga (5), Gaoudel (5), Koyretao (8),

Séoulasso (2), Toumboula (3).

Remarks: Several specimens were included in the mate-

rial used for the description of this species (Trape & M ané

2006a).All specimens from the type series have an entire

nasal and this character was used in the key of the genus

Dasypeltis (Trape et al. 2012). However, we recently ob-

served a semi-divided nasal in some specimens of D. sa-

helensis from M orocco.

Dispholidus typus (S mith, 1829)

M aterial: 19 specimens.

Localities: Laminina (5),

(2), Titiéna (1),

Doussoudiana (gle) 2

Mamoroubougou (7),

Zamoko (3).

Literature record: Sikasso (Villiers 1965).

N piébougou

Bonn zoological Bulletin 66 (2): 107-133

Grayia smithi (Leach, 1818)

M aterial: 15 specimens.

Locality: Mamoroubougou (15).

Remarks: First record for M ali.

Meizodon coronatus (Schlegel, 1837)

M aterial: 21 specimens.

Localities: Bangaya (1), Bouyanga (1), Mamoroubougou

(1), Sébékourani (12), Titiéna (6).

Literature records: Kati (Angel 1922), Tilembaya (Villiers

1951), Diafarabé (Villiers 1952), Wana Boubou (Villiers

1-9°5°3:).s

Natriciteres olivacea (Peters, 1854)

Material: 1 specimen (coll. MNHN).

Locality: Gao.

Literature record: Gao (Chabanaud 1917).

Philothamnus irregularis (Leach, 1819)

M aterial: 42 specimens.

Localities: Doussoudiana (9), Laminina (15), Niakoni (1),

Npiébougou (5), Sébékourani (8), Titiéna (4).

Literature records: Kati (Angel 1922, as Chlorophis em-

ini (Giinther, 1888)), Kayo (Villiers 1950, as Philotham-

nus nitidus (G inther, 1863)), Diafarabé, Tilem baya (Vil-

liers 1953), Ké-Macina (Villiers 1956), Sikasso (Villiers

1965), Lobi near Gao (Hughes 1983, 1985), Mourdiah

(Joger & Lambert 1996).

©ZFMK

118 Jean-Francois Trape & Youssouph M ané

Fig. 12. Vicinity of Haoussa Foulane village (16°00’N,00°08’W ), an area of Sahelo-Saharan transition where Echis leucogaster,

Eryx muelleri, Psammophis aft. sibilans ana Rhagerhis moilensis were collected.

Philothamnus semivariegatus smithi B ocage, 1882

M aterial: 4 specimens.

Localities: Doussoudiana (2), Laminina (1), Niamou (1).

Literature records: Bandiagara (Angel 1933), Gao (Hugh-

es 1985).

Remarks: Trape & M ané (2006b) attributed West African

populations of P. semivariegatus to a distinct subspecies,

i.e.P semivariegatus ssp., differing from the nominal sub-

species by its dorsal colouration: almost uniformly green

in West Africa, versus green with black crossbars in south-

ern Africa where the type originates. Trape & Baldé (2014)

revalidated Philothamnus semivariegatus smithi B ocage,

1882 for this subspecies. Preliminary molecular studies

(Trape & Mediannikov, unpublished) suggest that full spe-

cific rank may be justified.

Spalerosophis diadema cliffordi (Schlegel, 1837)

M aterial: 8 specimens.

Localities: Gaoudel (2), Koyretao (5), Toya (1).

Literature record: Goundam (Angel 1933).

Telescopus tripolitanus (Werner, 1909)

M aterial: 8 specimens.

Localities: Kinani (3), Toumboula (5),

Literature records: Kidal (Andersson 1935, as Tarbophis

obtusus (Reuss, 1834)), Mourdiah (Joger & Lambert

1996).

Remarks: Several specimens where included in the revi-

sion of this species by Crochet et al. (2008).

Bonn zoological Bulletin 66 (2): 107-133

Telescopus variegatus (Reinhardt, 1843)

M aterial: 8 specimens.

Localities: Koundian (1), Mamoroubougou (2), Zamoko

(5)

Literature record: Kati (Angel 1922).

Fam ily Lam prophiidae Ritzinger, 1843

Amblyodipsas unicolor (Reinhardt, 1843)

M aterial: 5 specimens.

Localities: Mamoroubougou (5).

Remarks: This record, the only one for Mali, appeared on

a grid map without specific documentation in Trape &

M ané (2006b).

Atractaspis aterrimd G iinther, 1863

M aterial: 5 specimens.

Localities: Doussoudiana (2), Laminina @l)s

Mamoroubougou (1), Sadjouroubougou (1).

Literature record: Bougouni (Schatti 1986).

Atractaspis dahomeyensis Barboza du Bocage, 1887

M aterial: 10 specimens.

Localities: Mamoroubougou (7), Npiébougou (2), Titié-

na (1).

Literature record: Tabakoro (Sch§atti 1986).

©OZFMK

The snakes of Mali 119

Atractaspis micropholis G iinther, 1872

M aterial: | specimen.

Locality: Niamou (1).

Literature record: (?) Madina Kagoro (Joger & Lambert

1996, specimen possibly attributable to Atractaspis wat-

SONI B oulenger, 1908).

Atractaspis watsoni B oulenger, 1908

M aterial: 7 specimens.

Localities: Bandiagara (1, coll. MNHN), Diafarabé (1,

coll. MNHN), Douentza (1, coll. MNHN), Samé Oulof

(1), Séoulasso (1), Topokhoné (2).

Literature records: Bandiagara (Angel 1933), Douentza

(Angel & Lhote 1938).

Remarks: This species was resurrected from the synonymy

of Atractaspis microlepidota (G iinther, 1866) by Trape et

al. (2006), who also reviewed previous records of Atrac-

taspis micropholis G iinther, 1872 and provided keys to dis-

tin guish the three species. Atractaspis microlepida may oc-

cur in Mali but this West African species is currently

known only from Senegal, The Gambia and southern M au-

ritania.

Boaedon fuliginosus (B oie, 1827)

M aterial: 92 specimens.

Localities: Bangaya (3), Bouyanga (1), Djinagué (7),

Doussoudiana (4), Kinani (2), Koundian (5), Laminina (8),

Mamoroubougou (16), Niamou (2), Npiébougou (11),

Sébékourani (20), Séoulasso (2), Titiéna (2), Zamoko (9).

Literature record: Diafarabé (Villiers 1951).

Boaedon lineatus Dum éril, Bibron & Duméril, 1854

M aterial: 70 specimens.

Localities: Bangaya (3), Djinagué (8), Doussoudiana (1),

Koundian (4), Laminina (2), Mamoroubougou (12),

N piébougou (5), Sadjouroubougou (1), Sébékourani (15),

Séoulasso (2), Titiéna (11), Zamoko (6).

Literature records: Kati (Angel 1922), Naréna (Schiatti

1986).

Gonionotophis granti (G iinther, 1863)

M aterial: 4 specimens.

Localities: Laminina (1), Mamorougougou (1),

N piébougou (1), Sébékourani (1).

Remarks: First record for M ali.

Lycophidion albomaculatum S§ teindachner, 1870

M aterial: 33 specimens.

Localities: Djinagué (5), Koundian (3), Niamou (3),

Sébékourani (16), Zamoko (6).

Literature records: Y élimané, Bamako (Condamin 1994).

Lycophidion irroratum (Leach, 1819)

M aterial: 5 specimens.

Localities: Mamoroubougou (5).

Bonn zoological Bulletin 66 (2): 107-133

Remarks: First record for M ali.

Lycophidion semicinctum (Dum éril, Bibron & Dum éril,

1854)

M aterial: 24 specimens.

Localities: Laminina (4), M amoroubougou (8),

N piébougou (9), Titiéna (3).

Literature record: Sikasso (Joger & Lambert 1996).

Mehelya crossi (B oulenger, 1895)

M aterial: 33 specimens.

Localities: (1505)

N piébougou (3), Sadjouroubougou (1), Sébékourani (4),

Titiéna (11), Zamoko (2).

Literature record: Sikasso (Villiers 1965).

Laminina (2), Mamoroubougou

Remarks: Four specimens from. three _ localities

(M am oroubougou, Npiébougou and Sébékourani) have 19

ranks of dorsals at midbody instead of 17. A1l other char-

acteristics are typical of Mehelya CYOSS1. We also observed

19 rows of dorsals in a specimen from Kissidougou

(Guinea), which was molecularly similar to a typical M.

CYOSSI (K Tolley and JF Trape, unpublished), suggesting

that Mehelya riggenbachi (Sternfeld, 1910) is a junior syn-

onym of M. crossi as previously mentioned by Broadley

(2007). The generic status of M. crossi is currently under

review (Broadley et al., submitted for publication).

Polemon neuwiedi (Jan, 1858)

M aterial: 3 specimens.

Locality: Mamoroubougou (3).

Remarks: First record for M ali.

Prosymna collaris (Sternfeld, 1908)

M aterial: | specimen.

Locality: Topokhoné (1).

Literature records: Y élimané (Broadley 1980, as Prosym-

na meleagris greigerti M ocquard, 1906; Chirio et al. 2011,

as Prosymna greigerti collaris Sternfeld, 1908), (?) Ba-

mako (Broadley 1980, as Prosymna meleagris greigerti;

Chirio et al. 2011, as Prosymna_ greigerti collaris;

MNHN specimen probably mislabelled).

Remarks: This Sahelian species with a white collar dif-

fers molecularly both from Prosymna greigerti M ocquard,

1906 and from Prosymna meleagris (Reinhardt, 1843), but

was provisionally treated as a subspecies of P. greigerti by

Chirio etal. (2011). We recently collected sympatric spec-

imens of the two taxa in Chad, where no trace of intergra-

dation was observed, and a similar situation was observed

in northern Cameroon. For this reason we prefer to treat

this taxon as separate species as it is easy to distinguish

by its colour pattern.

Prosymna greigerti M ocquard, 1906

M aterial: 48 specimens.

Localities: Bangaya (1), Djinagué (1), Koundian (1),

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120 Jean-Francois Trape & Youssouph M ané

Laminina (6), Cle),

N piébougou (7), Sébékourani (7), Titiéna (6), Toumboula

(1), Zamoko (2).

Mamoroubougou Niamou (1),

Literature records: Soudan francais (Chabanaud 1916, as

Prosymna meleagris), K ati (Angel 1922), San (Angel &

Lhote 1938), Bamako (Broadley 1980).

Remarks: M ost specimens were included in the recent re-

vision of Prosymna greigerti and Prosymna meleagris

(Chirio et al. 2011).

Psammophis elegans (Shaw, 1802)

M aterial: 91 specimens.

Localities: Bangaya (1), Bouyanga (1), Doussoudiana (6),

Kinani (5), Koundian (8), Laminina (8),Mamoroubougou

(7), Npiébougou (15), Sadjouroubougou (4), Samé Ouolof

(2), Saré-Soma (1), Sébékourani (6), Séoulasso (14), Tén-

intou (1), Titiéna (3), Zamoko (9).

Literature records: Kati (Angel 1922, as Psammophis

schokari Forssk al, 1775), Diré, Bandiagara (Angel 1933),

Dogo (Villiers 1953), Kita (Joger 1981), M ourdiah (Joger

& Lambert 1996).

Psammophis lineatus (Dum éril, Bibron & Duméril,

1854)

M aterial: 2 specimens.

Locality: Mamoroubougou (2).

Literature record: Kati (Angel 1922).

Psammophis phillipsi (Hallowell, 1844)

M aterial: | specimen.

Locality: Laminina (1).

Remarks: First record for Mali. This specimen with 17 dor-

sals, 180 ventrals and 102 subcaudals is dorsally uniform

with black spots on the supralabials, four infralabials in

contact with the first pair of mentals, and an entire anal

scale.

Psammophis praeornatus (Schlegel, 1837)

M aterial: 28 specimens.

Localities: Mamoroubougou (2), Niamou (1), Npiébougou

(1), Sadjouroubougou (1),Samé Ouolof (1), Sébékourani

(1), Séoulasso (7), Titiéna (6), Zamoko (8).

Literature records: Toguéré Sanga (Villiers 1956), Sikas-

so (Villiers 1965).

Psammophis schokari Forssk al, 1775

M aterial: 4 specimens.

Localities: Gaoudel (1), Tinjemban (3).

Literature record: Tombouctou (Chabanaud 1917).

Remarks: Loveridge (1940) erroneously attributed Cha-

banaud’s specimen to Psammophis elegans. None of our

specimens has the high ventral count (185 and higher) of

Psammophis aegyptius M arx, 1958.

Bonn zoological Bulletin 66 (2): 107-133

Psammophis aff. sibilans (Linnaeus, 1758)

M aterial: 722 specimens.

Localities: Agoudoud (2), Ballabougou (5), Bangaya (18),

Boussouma (13), Bouyanga (36), Djinagué (27), Dongui-

ba (1), Doussoudiana (13), Gaoudel (31), Haoussa-Foulane

(2), Kinani (17), Koundian (15), Koyretao (2), Laminina

(14), Léré (2), Mamoroubougou (13), Niakoni (3), Nia-

masso (16),Niamou (80), Npiébougou (23),Samé Ouolof

(43), Saré-Soma (27), Sébékourani (57), Séoulasso (82),

Tacharane (4), Tinjem ban (3), Titiéna (46), Toumboula (5),

Toya (66), Zamoko (56).

Literature records: Kati (Angel 1922), Bandiagara (An-

gel 1933, Papenfuss 1969), Diafarabé, Gourao (Villiers

1950), Mopti (Angel & Lhote 1938), Bamako (Villiers

1956), Sikasso (Villiers 1965), Kakoulou (Joger 1981), be-

tween Ségou and Séwaré, 158 km SW Séwaré, between

Séwaré and M opti, Djenné, Bla, Bandiagara (B6hme et

al. 1996), Lac Fabiguine, between Sikasso and Sinndi

(Joger & Lambert 1996).

Remarks: The status of West African specimens of the

Psammophis sibilans complex is currently under review

(Trape etal., in preparation). These specimens from M ali

are characterized by five infralabials in contact with the

first pair of mentals, a divided anal, and a more-or-less

striped dorsal pattern, with at leasta black and white chain

on the scales of the vertebral line.

Rhagerhis moilensis (Reuss, 1834)

M aterial: 8 specimens.

Localities: Gaoudel (5), Gogui (1), Haoussa-Foulane (1),

Koyretao (1).

Literature record: Adrar des Iforhas (Angel & Lhote 1938).

Rhamphiophis oxyrhynchus (R einhardt, 1843)

M aterial: 97 specimens.

Localities: Bangaya (1), Bouyanga (4), Djinagué (1),

D oussoudiana (6), Laminina (3),M amoroubougou (3), Ni-

akoni(2), Niamou (3), Npiébougou (16), Sébékourani (9),

Séoulasso (12), Titiéna (28), Zamoko (9).

Literature records: Kati (Angel 1922), Diafarabé (Villiers

1951), Kita (Joger 1981).

Family Elapidae Boie, 1827

Elapsoidea semiannulata moebiusi (Werner, 1897)

M aterial: 28 specimens.

Localities: Bangaya (1), Djinagué (1), Doussoudiana (2),

Laminina (1), Mamoroubougou (1), Npiébougou (13),

Sébékourani (2), Zamoko (7).

Literature record: southern Mali (Chippaux 1999).

Naja haje (Linnaeus, 1758)

M aterial: 2 specimens (NMZB and ZMUC collections).

Locality: Tombouctou (2).

©OZFMK

The snakes of Mali 121

Literature record: Tombouctou (Hughes 1983).

Remarks: These two museum specimens were examined

by D.G. Broadley as partofa review of the Naja haje com -

plex in West Africa (Trape et al. 2009). They belong to

Naja haje contrary to other reports of this species in M ali,

which are attributable to Naja senegalensis Trape, Chirio

& W ister, 2009 (see Trape et al. 2009 and below).

Naja katiensis Angel, 1922

M aterial: 356 specimens.

Localities: Ballabougou (1), Djinagué (4), Doussoudiana

(8), Laminina (14), Mamoroubougou (53), Npiébougou

(106), Sadjouroubougou (5), Sébékourani (8), Titiéna (69),

Zamoko (88).

Literature records: Kati (Angel 1922, type locality), Kok-

ounkourou, Naréna, Bougouni (Sch§atti 1986).

Naja cf. melanoleuca Hallowell, 1857

M aterial: 12 specimens.

Localities: Bangaya (1), Laminina (1), Mamoroubougou

(3), Npiébougou (4), Sébékourani (1), Titiéna (2).

Literature records: Kati (Angel 1922), Kokounkourou

(Schatti 1986).

Remarks: Our specimens have the banded pattern typical

of the West African savanna form of the Naja melanoleu-

Ca com plex.

Naja nigricollis Reinhardt, 1843

M aterial: 40 specimens.

Localities: Bangaya (7), Boussouma (3), Koundian (6),

Mamoroubougou (2), Niamasso (1), Niamou (4), Sébék-

ourani (6), Séoulasso (2), Toya (4), Zamoko (5).

Literature records: Kati (Angel 1922, Villiers 1951), Sikas-

so (Villiers 1965).

Naja senegalensis Trape, Chirio & W ister, 2009

M aterial: 33 specimens.

Localities: Ballabougou (2), Bangaya (1), Djinagué (2),

Doussoudiana (1), Koundian (1), Laminina (2),

Mamoroubougou (3), Npiébougou (2), Sadjouroubougou

(1), Saré-Soma (1), Sébékourani (2), Titiéna (10), Zamoko

(5).

Literature records: chutes du Félou, M ourdiah, Didieni

(Joger & Lambert 1996, as Naja haje).

Remarks: Our specimens were included in the description

of this species that was previously confounded with N. ha-

Je. Naja senegalensis is currently known from Senegal,

Gambia, Guinea, Guinea Bissau, Ivory Coast, Ghana,

M ali, Burkina Faso, Niger, Benin and Nigeria (Trape et

al. 2009, Trape & Baldé 2014).In M ali, it appears distrib-

uted in the whole country except in Sahelo-Saharan areas

where the occurrence of N. haje in Tom bouctou was con-

firmed (Trape et al. 2009).

Bonn zoological Bulletin 66 (2): 107-133

Family Viperidae Oppel, 1811

Bitis arietans (M errem, 1820)

M aterial: 79 specimens.

Localities: Bangaya (9), Bouyanga (3), Djinagué (2),

Doussoudiana (5), Chutes de Gouina (1), Koundian (2),

Laminina (8),M amoroubougou (5), Niakoni(l1), Niamas-

so (1), Niamou (5), Npiébougou (13), Sébékourani (10),

Titiéna (12), Zamoko (2).

Literature records: Katibougou, Sormé (Villiers 1950), Sé-

varé (Papenfuss 1969), Kokounkourou (Sch§atti 1986), be-

tween Bla and Bandiagara, Bla (Béhme et al. 1996).

Causus maculatus (Hallowell, 1842)

M aterial: 187 specimens.

Localities: Bangaya (11), Djinagué (10), Doussoudiana

(12), Koundian (8), Laminina (11),M amoroubougou (15),

Niamou (1), Npiébougou (7), Sébékourani (21), Titiéna

(36), Zamoko (55).

Literature records: Kati (Angel 1922, Villiers 1951, as

Causus rhombeatus (Lichtenstein, 1823)), Saré Malé (Vil-

liers 1950, as C. rhombeatus), Taga Diabozo (Villiers

1952, as C. rhombeatus), Dogo (Villiers 1953, as C.

rhombeatus), Bamako (Villiers 1956, as C. rhombeatus),

(Villiers 1965, as C. rhombeatus), Naréna,

Bougouni (Schatti 1986), see also map of Hughes (1977).

Sikasso

Cerastes cerastes (Linnaeus, 1758)

M aterial: 2 specimens (coll. MNHN).

Localities: 50 km N of Kidal, rocher d’Eguerer.

Literature and sight records: Timétrine (de Witte 1930),

Tessalit (Joger & Lambert 1996), SW of Adrar Takabart

(sight record).

Cerastes vipera (Linnaeus, 1758)

Material: | specimen (coll. MNHN).

Locality: Between Tisserlitine and Tim étrine.

Literature and sight records: Between Tisserlitine and

Tim étrine (de Witte 1930), vicinity of Araouane, half way

between Araouane and Taoudeni, Tessalit (sight records).

Echis leucogaster R oman, 1972

M aterial: 523 specimens.

Localities: Agoudoud (2), Boussouma (2), Bouyanga

(188), Doro (3), Gaoudel (44), Haoussa-Foulane (3), Ki-

nani (10), Koyretao (31), Léré (1), Niamou (43), Samé

Ouolof (21), Séoulasso (2), Tacharane (11), Tinjemban

(16), Topokhoné (1), Toumboula (145).

Literature records: Bandiagara (Angel 1933,as Echis car-

inatus (Schneider, 1801)), Kidal (Andersson 1935, as

Echis carinatus), A drar des Iforhas (Angel & Lhote 1938,

as Echis carinatus), M opti, M ourdiah (Joger & Lambert

1996), Séoulasso (Pook et al. 2009).

Remarks: Most specimens have a clear venter, but spec-

imens with a spotted venter, at least on each side of the

©ZFMK

122 Jean-Francois Trape & Youssouph Mané

ventrals, were not rare, particularly in western Mali. All

specimens had high scale counts, typical of E. leucogaster

(ventrals: 158-177 in males, 166-189 in females; subcau-

dals 30-40 in males, 27-35 in females).

Echis ocellatus Stem m ler, 1970

Echis jogeri Cherlin, 1990

M aterial: 933 specimens.

Localities: Ballabougou (3), Bangaya (4), Bouyanga (14),

Djinagué (21), Doussoudiana (16), Koundian (4), Lamin-

ina (26), Mamoroubougou (65), Niakoni (4), Niamou

(225), Npiébougou (60), Sadjouroubougou (9), Sébék-

ourani (99), Séoulasso (159), Titiéna (80), Zamoko

(144).

Literature record: Bandiagara (Angel 1933, as Echis car-

inatus), Kati (Villiers 1951, as Echis carinatus, Hughes

1976), Bamako (Villiers 1952, 1956, as Echis carinatus),

Kita (Villiers 1953 as Echis carinatus), between Negala

and Kassaro, W of Kita, W of Niamiga (Joger 1981, as

Echis sp.; Joger & Lambert 1996, as Echis Jogeri),

Tom bouctou (Cherlin 1990, type locality of Echis jogeri),

Niakoni (Pook et al. 2009).

Remark: Cherlin (1990) described Echis Jogeri on the ba-

sis of a clear venter and a much lower ventral count than

Echis ocellatus. The type of E. jogeriis a female with 132

ventrals (123 ventrals were erroneously mentioned by

Cherlin), and such low counts are also observed in Sene-

gal (121-135 in males, 128-143 in females) and north-

western Guinea (127-132 in males, 131-138 in females)

(Trape & Mané 2004, Trape & Baldé 2014). By contrast,

the type of Echis ocellatus from Garango (Burkina Faso)

is a female with 156 ventrals. In this country, ventral

counts of Echis ocellatus range from 134 to 152 in males

and from 140 to 157 in females (Roman 1972, 1976). Pook

et al. (2009) demonstrated that specimens from Senegal

are molecularly divergent from those of other parts of

Africa and hence confirmed that E. jogeri is a valid

species.

In order to separate E. jogeri from E. ocellatus in our

study, we investigated a sample of 274 specimens from the

different localities where we obtained specimens of this

complex. Based on our data from Senegal (all specimens

from this country have low ventral counts and are thus pre-

sumed to belong to E. Jogeri) and Roman’s data from

Burkina Faso (most of them are presumed to belong to E.

ocellatus since there is very little overlap in ventral counts

with Senegalese specimens), we attributed to EF. Jogeri

specimens with 121-133 (males) or 128-139 (females)

ventrals, and to E. ocellatus specimens with 136-152

144-157 M ales

134-135 ventrals and females with 140-143 ventrals were

classified as incertae sedis. Table 3 shows that most spec-

Mali (Bangaya,

Koundian, Sébékourani) can be attributed to E. jogeri, and

(males) or (females) ventrals. w ith

imens from southwestern Djinagué,

most specimens from central-western (Bouyanga) and

south-eastern Mali (Ballabougou, Doussoudiana, Lam in-

Npiébougou, Sad-

jouroubougou, Séoulasso, Titiéna) to E. ocellatus. B oth

ina, Mamoroubougou, Niakoni,

species are present in south-central Mali where the pro-

portion of specimens incertae sedis was much higher than

expected (e.g. Niamou, Zamoko), suggesting that hybrids

may occur and/or that the range of ventral counts for one

or both the two species differs between M aliand its neigh-

bouring countries. The presence of absence of black spots

Table 3. Number of specimens attributable to Echis jogeri ort Echis ocellatus based on the number of ventral scales. E. Jogeri:

121-133 (males) or 128-139 (females); E. ocellatus 136-152 (males) or 144-157 (females); incertae sedis: 134-135 (males) or

140-143 (females). Only specimens with intact body and tail allowing full ventral and subcaudal counts were selected for the study.

Locality (N° specimens) N° studied specimens

N° of E. jogeri

N° of E. ocellatus N° of incertae sedis

Bangaya (4) 3 2 0 1

Djinagué (21) 18 15 0 3

Koundian (4) 3 3 0 0

Sébékourani (99) 60 43 7 10

Ballabougou (3) 1 0 1 0

Bouyanga (14) 6 0 6 0

D oussoudiana (16) 1 0 0 1

Laminina (26) 18 1 7 10

Mamoroubougou (65) 11 0 10 1

Niakoni (4) 1 0 1 0

Niamou (225) 42 16 4 22

N piébougou (60) 1 0 1 0

Sadjouroubougou (9) 1 0 1 0

Séoulasso (159) 20 1 11 8

Titiéna (80) 3 1 2 0

Zamoko (144) 85 20 25 40

Total 274 102 76 96

Bonn zoological Bulletin 66 (2): 107-133

©OZFMK

The snakes of Mali 123

on the venter was not helpful to distinguish the two species

and further investigations to clearly separate E. jogeri from

E. ocellatus in the field are needed.

DISCUSSION

Previous reports on the snake fauna of Mali were based

on small collections, totalizing together abouttwo hundred

specimens. Our collection of snakes from M ali com pris-

es 5,224 specimens belonging to 60 species. With addi-

tional museum material examined and taking into account

reliable literature reports, the snake fauna of Mali com-

prises 65 species, i.e. 18 species more than the previous

checklist published 20 years ago by Joger & Lambert

(1996, 1997).

As expected, maximum diversity was observed in the

of the 11°00’N and

12°00’N (Table 4). In this area of transition between the

southern part country, between

Sudanese and the Guinean savannas 46 species were col-

lected, with amaximum of 36 species atMamoroubougou

(11°14’N), a locality near a small river with a preserved

forest gallery which was extensively investigated both for

fossorial and non-fossorial species (Fig. 7). The most

abundant species between 11°00’N and 12°00’N were

Tricheilostoma bicolor, Naja katiensis, Echis ocellatus,

Psammophis aft. sibilans, Crotaphopeltis hotamboeia and

Causus maculatus, and othercommon species were Rham-

Phiophis oxyrhynchus, Bitis arietans, Boaedon fuliginosus,

Psammophis elegans, Prosymna greigerti, and Boaedon

lineatus. Eight Guinean savanna species — most of them

collected in very low num bers — reach their northern lim -

itin Mali south of 11°30’N, i.e. Psammophis phillipsi, Am-

blyodipsas unicolor, Grayia smithi, Lycophidion irrora-

tum, Polemon neuwiedi, Myriopholis albiventer, Rhinogu-

inea magna, and Afronatrix anoscopus, and four addition-

al Guinean species were not collected north of 12°N,

namely Afrotyphlops lineatus, Atractaspis dahomeyensis,

Crotaphopeltis hippocrepis ana Lycophidion semicinctum.

Between 12°00’N and 14°00’N, an area of typical Su-

danese savanna (Fig. 8), a total of 43 species were collect-

ed, with a maximum of 28 and 25 species, respectively,

in Sébékourani (12°12’N) and Zamoko (13°09’N), two lo-

calities also extensively investigated for fossorial species

(Fig. 9). The most abundant species were Echis ocellatus

(w ith E. jogeri in the western part of the country), Psam-

mophis aft. sibilans, Crotaphopeltis hotamboeia, Causus

maculatus, Tricheilostoma bicolor, Naja katiensis, and

Myriopholis boueti, and othercommon species were Boae-

don fuliginosus, Psammophis elegans, Boaedon lineatus,

Rhamphiophis oxyrhynchus, ana Lycophidion albomacu-

latum (in the western part of the country). Four Sahelian

species were collected south of 14°N, butthe number and

proportion of specimens was low: only five specimens of

Eryx muelleri com pared to 169 specimens north of 14°N,

Bonn zoological Bulletin 66 (2): 107-133

with a southernmost record at Niamasso (12°59’N), and

only two specimens of Echis leucogaster com pared to 521

specimens north of 14°N, with a southernmost record at

Séoulasso (13°14’N). Two other Sahelian species were col-

lected in very low number and proportion south of 14°N

at Séoulasso: Dasypeltis sahelensis (two compared to 21),

and Atractaspis watsoni (one compared to three).

Between 14°N and 15°N, an area of rapid transition with

the Sahel (Fig. 10), a total of 27 species were collected,

but only 17 of these species were collected or are known

north of 14°30’N. Five species represented together 90%

of the snakes collected: Echis leucogaster, Psammophis

aff. sibilans, Echis ocellatus, Echis jogeri, and Eryx muel-

leri. Only two other species reached 1% of the specimens

collected: Crotaphopeltis hotamboeia ana Myriopholis

boueti.

Between 15°N and 16°N, a typical Sahelian area (Fig.

11), 14 species were collected, with two of them (name-

ly Psammophis aft. sibilans ana Echis leucogaster) rep-

resenting together 77% of the snakes collected. Other com-

mon species were Eryx muelleri and Rhagerhis moilen-

SIS. Only four species widely distributed in the southern

part of the country were also collected: Naja nigricollis,

Psammophis elegans, Python sebae, and Boaedon fulig-

inosus.

North of 16°N,a total of 18 species are now known from

Mali but only 8 were collected during our study, all be-

tween 16°N and 17°N, an area of Sahelo-Saharan transi-

tion (Fig. 12). The most abundant species were Psam-

mophis aff. sibilans (43% of specimens collected), Echis

leucogaster (35% ), Eryx muelleri (9% ) and Dasypeltis sa-

helensis (5%). The

Spalerosophis diadema, Naja nigricollis, Psammophis

schokari and Rhagerrhis moilensis. The ten species not

collected by us are either Saharan species ( Cerastes vipera,

Cerastes cerastes, Myriopholis algeriensis), or species

other species collected were

with a large distribution in Africa but apparently rare in

Mali (Naja haje, which seems excluded by Naja sene-

galensis from most parts of Mali, and Natriciteres oli-

vaced, a rare species in West Africa), or species that we

collected south of 16°N but not in northern Mali (Tele-

SCOpUus tripolitanus, a species known from Kidal, Echis

Jogeri, a species with Tombouctou as type locality, and

three species collected in the past along the banks of the

Niger River near Gao or Tombouctou, ice. Philothamnus

irregularis, Philothamnus semivariegatus and Psam-

mophis elegans.

In all areas of the country, both the number and the pro-

portion of highly venomous snakes were very high. The

three Echis species represented together 28% of the snakes

collected (34% when excluding Leptotyphlopidae) and

were the mostcommon snakes in all areas of the country,

with E. leucogaster as dominant species north of 14°N,

E. ocellatus east of 8°W and E. jogeri in the M andingue

Plateau west of 8°W. The high mortality due to snakebite

©ZFMK

124 Jean-Francois Trape & Youssouph M ané

Table 4. Latitudinal distribution of snakes in Mali (our study, 5,224 specimens collected). For the five species not collected dur-

ing our study, studied museum specimens, literature data and sight records are indicated in brackets. Latitude for the northernmost

record in Mali is based on whole data including literature data (circa 5,400 specimens).

Species 11°N 12°N 13°N 14°N 15°N 216°N Total Northernm ost record in Mali

Cerastes vipera 0 0 0 0 0 (4) (4) 20°40°N

Cerastes cerastes 0 0 0 0 0 (5) (5) 20°12°N

Echis leucogaster 0 0 2 397 63 61 523 18°26’N

Telescopus tripolitanus 0 0 0 5 3 0 8 18°26’°N

Rhagerhis moilensis 0 0 0 0 7 1 8 18°00°N

Myriopholis algeriensis 0 0 0 0 0 (2) (2) 16°57°N

Psammophis schokari 0 0 0 0 1 3 4 16°46’°N

Naja haje 0 0 0 0 0 (2) (2) 16°46’°N

Echis ocellatus + E. jogeri 254 132 311 239 0 0 933 16°46°N

Eryx muelleri 0 1 4 142 11 16 174 16°44°N

Psammophis aft. sibilans 112 105 FED 191 67 75 722 16°44°N

Spalerosophis diadema 0 0 0 0 2 6 8 16°39°N

Naja nigricollis 2 7 20 4 3 4 40 16°39°N

Psammophis elegans 40 10 32 4 5 0 91 16°16°N

Philothamnus semivariegatus 3 0 1 0 0 0 4 16°16°N

Natriciteres olivacea 0 0 0 0 0 (1) (1) 16°16°N

Philothamnus irregularis 34 8 0 0 0 0 42 16°15°N

Dasypeltis sahelensis 0 0 2 8 5 8 23 16°04’N

Causus maculatus 81 31 74 1 0 0 187 15°10°N

Prosymna collaris 0 0 0 0 I 0 1 15°08°N

Lycophidion albomaculatum 0 21 9 3 0 0 33 15°08°N

Python sebae 5 8 1 8 1 0 23 15°06°N

Atractaspis watsoni 0 0 1 1 2 0 4 15°02’°N

Boaedon fuliginosus 41 27 19 3 2 0 92 15°O1°N

Bitis arietans 44 13 13 9 0 0 79 14°52°N

Crotaphopeltis hotamboeia 82 92 36 15 0 0 225 14°45°N

Naja senegalensis 18 7 7 1 0 0 33 14°45’N

Myriopholis boueti 26 13 75 11 0 0 126 14°30°N

Rhamphiophis oxyrhynchus 58 10 25, 7 0 0 97 14°30’N

Meizodon coronatus 7 12 2 0 0 0 21 14°30°N

Psammophis praeornatus 9 2 15 2 0 0 28 14°29°N

Bamanophis dorri 0 0 BS 9 0 0 37 14°21°N

Prosymna greigerti 34 8 4 2 0 0 48 14°20°N

Atractaspis micropholis 0 0 0 1 0 0 1 14°20°N

Afrotyphlops punctatus 13 0 6 2 0 0 21 14°09°N

Tricheilostoma bicolor 725 9 101 3 0 0 838 14°09°N

Dasypeltis confusa 2 0 0 0 0 0 2 14°09°N

Dasypeltis latericia 3 1 16 5 0 0 25 14°01°N

Python regius 26 23 5 1 0 0 55 14°01°N

Boaedon lineatus 31 24 15 0 0 0 70 13°14°N

Dasypeltis gansi 0 2 1 0 0 0 3 13°14’°N

Elapsoidea semiannulata 17 3 8 0 0 0 28 13°14°N

Naja cf. melanoleuca 10 1 1 0 0 0 12 13°14°N

Dispholidus typus 16 0 3 0 0 0 19 13°09°N

Mehelya crossi 26 5 2 0 0 0 33 13°09°N

Naja katiensis 250 18 88 0 0 0 356 13°09°N

Rhinoleptus koniagui 1 I 0 0 0 11 13°09°N

Telescopus variegatus 2 0 6 0 0 0 8 13°09°N

Psammophis lineatus 2 0 0 0 0 0 2 12°43°N

Atractaspis aterrima 4 1 0 0 0 0 5 12°35’N

Gonionotophis granti 3 1 0 0 0 0 4 12°12°N

Myriopholis adleri 0 1 0 0 0 0 1 12°12°N

Afrotyphlops lineatus 12 0 0 0 0 0 12 11°59°N

Atractaspis dahomeyensis 10 0 0 0 0 0 10 11°59°N

Crotaphopeltis hippocrepis 3 0 0 0 0 0 3 11°59°N

Lycophidion semicinctum 24 0 0 0 0 0 24 11°59'N

Afronatrix anoscopus 17 0 0 0 0 0 17 11°26°N

Amblyodipsas unicolor 5 0 0 0 0 0 5 11°14°N

Grayia smithi 15 0 0 0 0 0 15 11°14°N

Lycophidion irroratum 5 0 0 0 0 0 5 11°14’N

Polemon neuwiedi 3 0 0 0 0 0 3 11°14°N

Myriopholis albiventer 5 0 0 0 0 0 5 11°14°N

Rhinoguinea magna 16 0 0 0 0 0 16 11°14°N

Psammophis phillipsi 1 0 0 0 0 0 I 11°12°N

Number of specimens 2103 597 1103 1074 173 174 5224

Number of species 46 32 36 27 14 8 (+5) 60 (+5)

Bonn zoological Bulletin 66 (2): 107-133 ©ZFMK

The snakes of Mali 125

in south-eastern Senegal was attributed to Echis jogeri

(Trape etal. 2001) and the other species of the Echis ocel-

latus com plex are also known to be a majorcause of death

in savanna areas of Nigeria (Warrell & Arnett 1976). Sev-

eral other dangerous species were both common and wide-

spread, e.g. Bitis arietans, Naja nigricollis, and Naja sene-

galensis. The Katian spitting cobra Naja Katiensis was

among the most abundant snakes in Mali but the severi-

ty of its bite is poorly documented. Together, the poten-

tially deadly snakes represented 38% of the snakes col-

lected during our study and 47% of snakes when exclud-

ing worm snakes of the family Leptotyphlopidae.

Acknowledgments. We thank G. Diatta for assistance during

field work, G. Chauvancy for assistance during preparation of

the appendix, and the anonym ous reviewers for their useful com -

ments.

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©ZFMK

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APPENDIX 1

List of collected specimens, locality,and IRD collection num -

ber

Afronatrix anoscopus. M am oroubougou: 1991.M, 1993.M,

1995.M, 2006.M, 2007.M, 2009.M, 2016.M, 2038.M, 2049.M,

2056.M,3692.M, 3750.M, 3761.M, 3766.M, 3784.M, 3823.M;

Titiéna: 3689.M.

Afrotyphlops lineolatus. N piébougou: 3992.M, 4152.M,

4153.M,4154.M,4155.M,4156.M, 4157.M, 4164.M, 4166.M,

4168.M,TR.2504; Laminina: 174.M.

Afrotyphlops punctatus. Niamou: 1924.M, 4264.M (ligné);

N piébougou: 4286.M (ligné); Titiéna: 2125.M, 4171.M,

4176.M (ligné) ; Zamoko: 4149.M, 4150.M,4151.M (ligné) ;

Laminina: 754.M (marbré); Mamoroubougou: 4134.M,

4145.M,4146.M,4147.M,4148.M,4158.M,TR.2503 (marbré) ;

Titiéna: 1932.M (marbré); Zamoko: 160.M,1916.M, 1921.M

(noir).

Amblyodipsas unicolor. M am oroubougou: 1947.M, 1979.M,

3557.M,3790.M, 3796.M.

Bonn zoological Bulletin 66 (2): 107-133

scription of a new species from West Africa. Zootaxa 2236:

1-25

Trape S,Mediannikov O,Trape JF (2012) When colour patterns

reflect phylogeography —- New species of Dasypeltis (Ser-

pentes: Colubridae: Boigini) from WestA frica. Com ptes-Ren-

dus Biologies 335: 488-501

Uetz P, Freed P, HoSek J (2016) The Reptile Database (Eds.). On-

line at: http://w ww.reptile-database.org last accessed on Sep-

tem ber 10" 2017

Villiers, A (1950) La collection de serpents de I’I.F.A.N. Insti-

tut Frangais d’Afrique Noire, Catalogues VI, Dakar

Villiers A (1951) La collection de serpents de l’?IFAN (acquisi-

tions 1950). Bulletin de |’ Institut Francais d’A frique Noire 13:

813-836

Villiers A (1952) La collection de serpents de l’?IFAN (acquisi-

tions 1951). Bulletin de |’ Institut Francais d’A frique Noire 14:

881-898

Villiers A (1953) La collection de serpents de l’?IFAN (acquisi-

tions 1952). Bulletin de l'Institut Francais d’A frique Noire 15:

1103-1127

Villiers A (1954) La collection de serpents de l’?IFAN (acquisi-

tions 1953). Bulletin de I’ Institut Francais d’A frique N oire 16

(sér. A): 1234-1247

Villiers A (1956) La collection de serpents de 1’?IFAN (acquisi-

tions 1954-1955). Bulletin de l'Institut Francais d’A frique

Noire 18 (sér. A): 877-883

Villiers A (1965) Serpents récoltés au Mali et en Haute-Volta

par le Dr. Lamontellerie. Bulletin de

d’A frique Noire 27 (sér. A): 1192-1195

Villiers A,Condamin M (2005) Les serpents de l’OuestA fricain.

IFAN, Inititiation et Etudes A fricaines II (4 éme édit.), Dakar

WarrellDA,Arnett C (1976) The importance of bites by the saw-

scaled or carpet-viper (Echis carinatus): epidemiological stud-

ies in Nigeria and a review of the world literature. Acta Trop-

ica 33: 307-341

W itte GF de (1930) Mission Saharienne Augiéras-Draper, 1927-

l’Institut Francais

1928. Reptiles et Batraciens. Bulletin du Muséum national

d’Histoire naturelle, Paris, série 2, 2: 614-618

W iister W, Broadley (2003)A new species of spitting cobra (Na-

ja) from north-eastern Africa (Serpentes: Elapidae). Journal

of Zoology 259: 345-359

Atractaspis aterrima. D oussoudiana: 111.M,TR.0712; Lamin-

ina: TR .0649; Mam oroubougou: 3850.M;Sadjouroubougou:

2521.M.

Atractaspis dahomeyensis. M am oroubougou: 1936.M,1941.M,

1989.M, 2029.M, 2036.M, 3734.M, 3767.M; Npiébougou:

1808.M,4165.M; Titiéna: 3659.M.

Atractaspis micropholis. Niam ou: 1923.M.

Atractaspis watsoni. Samé Ouolof: TR.1684; Séoulasso:

1493.M; Topokhoné: 126.M,128.M.

Bamanophis dorri. Bangaya: 1.M, 9.M, 11.M, 14.M, 15.M,

19.M,23.M,26.M, 28.M, 31.M, 228.M, 231.M, 243.M, 259.M,

264.M, 265.M, 266.M, 277.M, 278.M, 279.M, 292.M, 295.M,

296.M,297.M; Koundian: 69.M, 2364.M, 3012.M, 3018.M; Ni-

amou: 44.M, 2977.M, 2978.M, 2983.M, 2987.M, 2995.M,

2997.M,3000.M; Toumboula: 2799.M.

Bitis arietans. Bangaya: 20.M, 25.M, 219.M, 223.M, 226.M,

236.M, 244.M, 290.M, 298.M; 55.M, 400.M,

403.M; Djinagué: 1160.M, 1205.M; Doussoudiana: 90.M,

125.M,1760.M,1771.M,TR.0917; Gouina (chutes): 5299.M ;

Koundian: 64.M, 2373.M; Laminina: 762.M,788.M, 820.M,

©OZFMK

Bouyanga:

The snakes of Mali 127

TR .0922, TR.1100, TR v2.25, TR.1309, TR.1314;

M am oroubougou: 175.M,2060.M,2064.M,4111.M,4118.M;

Niakoni: TR.0932; Niamasso: 1344.M; Niamou: 420.M,

422.M, 436.M, 2988.M, 2993.M; Npiébougou: 1852.M,

1884.M ,3327.M,3375.M, 3425.M, 3434.M,3449.M, 3454.M,

3462.M, 3477.M, 3490.M, 3498.M, 3999.M, Sébékourani:

944.M, 961.M, 1000.M, 1002.M, 1041.M, 1081.M, 1082.M,

3243.M,3246.M, 3270.M; Titiéna: 2112.M,2151.M,2180.M,

2263.M,3675.M,4015.M,4021.M,4023.M,4024.M,4025.M,

4027.M,4033.M; Zamoko: 662.M,2558.M.

Boaedon fuliginosus. Bangaya: 225.M, 239.M, 272.M;

Bouyanga: 2500.M; Djinagué: 1111.M, 1114.M, 1133.M,

1134.M, 1151.M, 1166.M, 1194.M; Doussoudiana: 94.M,

112.M,TR.1203,TR.1235; Kinani: 1564.M,1584.M; Koundi-

an: 71.M,72.M,2369.M,3025.M,3032.M; Laminina: 757.M,

766.M, 776.M, 790.M, 792.M, 819.M, 2622.M, TR.0647;

M am oroubougou: 1939.M,1980.M,1988.M, 3696.M ,3720.M,

3722.M,3724.M,3745.M,3771.M,3780.M,3781.M,3794.M,

3797.M,3800.M,3819.M,3844.M; Niamou: 484.M, 2923.M;

N piébougou: 1809.M, 1810.M, 1843.M, 1868.M, 1869.M,

1870.M, 3313.M, 3376.M, 3386.M, 3416.M, 3478.M; Sébék-

ourani: 876.M,882.M,899.M,920.M,938.M,975.M,1040.M,

1075.M,1079.M,1086.M, 3161.M,3180.M,3208.M, 3230.M,

3232.M,3236.M,3237.M,3239.M,3266.M,3290.M; Séoulas-

so: 1715.M, 3094.M; Titiéna: 2073.M, 2130.M; Zamoko:

564.M,566.M,570.M,670.M,672.M,679.M,696.M,2602.M,

2658.M.

Boaedon lineatus. Bangaya: 221.M,275.M,291.M; Djinagué:

1119.M,1128.M,1145.M,1161.M,1168.M,1186.M,1201.M,

3293.M; Doussoudiana: 119.M; Koundian: 65.M, 66.M,

3001.M,3021.M;Laminina: 824.M,848.M;Mamoroubougou:

2026.M,2046.M,2058.M,2059.M,3751.M,3774.M,3785.M,

3786.M, 3813.M, 3827.M, 3848.M, 3853.M; Npiébougou:

1779.M,1854.M,1875.M,3357.M,3441.M; Sadjouroubougou:

2512.M; Sébékourani: 872.M, 888.M,907.M,908.M, 909.M,

918.M, 925.M, 927.M, 3158.M, 3166.M, 3201.M, 3212.M,

3245.M,3253.M,3267.M; Séoulasso: 3086.M, 3128.M;; T itié-

na: 191.M, 2104.M, 2134.M, 2156.M, 2164.M, 2178.M,

2199.M,2203.M,2256.M, 3668.M,4032.M; Zamoko: 671.M,

2534.M,2593.M,2606.M,2641.M,2702.M.

Causus maculatus. Bangaya: 17.M, 30.M, 224.M, 257.M,

258.M ,260.M,273.M,282.M,286.M,299.M,301.M, Djinagué:

1118.M,1120.M,1123.M,1124.M,1154.M,1156.M,1158.M,

1180.M, 1188.M, 1192.M, Doussoudiana: 84.M, 1761.M,

1763.M,1765.M,1768.M,1769.M,1773.M,TR.0715,TR.0718,

TR.0851,TR.1233,TR.1304; Koundian: 61.M,67.M,2366.M,

3010.M,3019.M,3034.M ,3035.M,3043.M; Laminina: 761.M,

782.M,796.M, 806.M, 812.M, 813.M, 816.M, 836.M, 851.M,

856.M, 858.M; Mam oroubougou: 2033.M, 2035.M, 2050.M,

2063.M,2066.M,3749.M, 3802.M, 3829.M, 3831.M,4103.M,

4123.M,4127.M,4131.M,4132.M,4140.M; Niam ou: 2986.M;

Npiébougou: 1799.M, 1829.M, 1840.M, 3324.M, 3398.M,

3428.M,3465.M; Sébékourani: 864.M,902.M,913.M,940.M,

959.M, 962.M, 1013.M, 1037.M, 1050.M, 1060.M, 1105.M,

3169.M,3203.M,3204.M,3210.M,3218.M,3224.M,3241.M,

3247.M, 3249.M, 3264.M; Titiéna: 195.M, 2082.M, 2084.M,

2137.M,2153.M,2154.M,2165.M,2171.M,2194.M,2202.M,

2234.M,2241.M,2262.M, 3609.M, 3610.M, 3611.M,3641.M,

3674.M,3684.M,4011.M,4012.M,4013.M,4014.M,4016.M,

4018.M,4020.M,4028.M,4030.M,4031.M,4034.M, 4036.M,

4038.M, 4039.M, 4303.M,4304.M,4305.M; Zamoko: 567.M,

583.M,590.M,593.M, 600.M, 614.M,637.M,641.M, 642.M,

648.M,657.M, 664.M, 669.M, 689.M, 692.M,693.M,702.M,

Bonn zoological Bulletin 66 (2): 107—133

704.M,723.M,730.M,735.M,736.M,737.M,2532.M,2533.M,

2537.M,2538.M,2542.M, 2552.M,2555.M,2557.M, 2559.M,

2562.M,2574.M,2575.M,2576.M,2585.M, 2594.M,2595.M,

2605.M,2608.M,2609.M, 2616.M, 2633.M, 2640.M, 2642.M,

2646.M,2647.M, 2648.M, 2672.M, 2690.M, 2691.M, 2692.M,

2693.M,2694.M.

Crotaphopeltis hippocrepis. Mamoroubougou: TR.2502;

Npiébougou: 1856.M.

Crotaphopeltis hotamboeia. B allabougou: 2525.M ; Bangaya:

12.M, 18.M, 21.M, 227.M, 229.M, 232.M, 233.M, 234.M,

237.M, 240.M, 241.M, 242.M, 251.M, 253.M, 254.M, 255.M,

256.M, 261.M, 263.M, 269.M,271.M, 280.M, 281.M, 284.M,

293.M,300.M ; Djinagué: 1109.M,1112.M,1121.M,1126.M,

1127.M,1131.M,1136.M,1137.M,1140.M,1143.M,1146.M,

1147.M,1148.M,1149.M,1150.M,1153.M,1163.M,1169.M,

1174.M,1176.M,1177.M,1184.M,1185.M,1190.M,1195.M,

1196.M, 1198.M, 1200.M, 1208.M ; Doussoudiana: 79.M,

86.M,97.M,104.M,105.M,123.M,1758.M,TR.0717,TR.0844,

TR.0915, TR.0916, TR.0919, TR.1236 ; Laminina: 738.M,

742.M,744.M,746.M,748.M,778.M,841.M, 843.M,TR.1230,

TR.1274 ; Mamoroubougou: 2014.M, 2044.M, 3691.M,

3717.M,3746.M, 3747.M, 3759.M, 3791.M, 3793.M, 3798.M,

3801.M,3809.M, 3820.M, 3824.M, 3826.M, 3830.M, 3839.M,

3841.M,3842.M,4106.M,4107.M,4130.M,4136.M,4138.M,

4161.M; Niakoni: TR.0933, TR.1296; Niamasso: 1341.M,

1343.M; Niamou: 38.M, 40.M, 461.M, 2979.M, 2980.M,

2994.M, TR.0257, TR.0258; Npiébougou: 1780.M, 1806.M,

1815.M,1817.M,1823.M,1861.M,1862.M,3337.M, 3361.M,

3467.M,3491.M; Sadjouroubougou: 2513.M,2514.M ; Saré-

Soma: 1594.M,1614.M;Sébékourani: 867.M, 868.M, 870.M,

874.M,877.M, 887.M, 889.M, 892.M, 893.M, 894.M, 895.M,

901.M,904.M,905.M,906.M,910.M,914.M,919.M,921.M,

924.M,926.M,932.M,933.M,935.M,942.M,948.M, 950.M,

952.M, 956.M, 965.M, 978.M, 990.M, 1001.M, 1009.M,

1014.M,1027.M,1043.M,1069.M,3155.M,3163.M,3168.M,

3171.M,3175.M,3177.M, 3179.M, 3182.M,3199.M,3202.M,

3217.M,3222.M,3225.M, 3231.M,3235.M,3248.M,3262.M,

3263.M,3274.M,3279.M; Séoulasso: 1388.M; Titiéna: 211.M,

215.M, 2074.M, 2075.M, 2077.M, 2081.M, 2085.M, 2088.M,

2093.M,2099.M,2122.M,2123.M,2124.M,2144.M, 2168.M,

2170.M,2174.M,2218.M,2226.M,2235.M,2242.M,3621.M;

Toumboula: 1272.M, 1281.M, 2794.M, 2796.M, 2797.M;

Zamoko: 595.M, 610.M, 649.M, 2563.M, 2572.M, 2580.M,

2654.M,2664.M,2707.M.

Dasypeltis confusa. M am oroubougou: 3731.M, 4104.M.

Dasypeltis gansi. Ballabougou: 2355.M, 2356.M; Séoulasso:

4141.M.

Dasypeltis latericia. Bangaya: 27.M, 276.M, 285.M, 288.M ;

K oundian: 2371.M;Niamou: 138.M,139.M,2357.M,2358.M,

2359.M;Npiébougou: 4315.M,4316.M; Sébékourani: 153.M;

Séoulasso: 1351.M, 1714.M, 1715.M, 4163.M;

1930.M ; Zamoko: 161.M, 162.M, 163.M, 171.M, 172.M,

2619.M,4063.M.

Titiéna:

Dasypeltis sahelensis. Bouyanga: 130.M, 412.M, 2361.M,

2362.M, 2363.M;: G aoudel: 1535.M, 3561.M, 3567.M, 3573.M,

3583.M; Séoulasso: 1395.M, 1505.M; Toumboula: 1230.M,

1231.M, 2360.M.

Dispholidus typus. D oussoudiana: 98.M; Laminina: 743.M,

763.M,784.M,795.M, TR.1227; Mamoroubougou: 1976.M,

2002.M, 2011.M, 3693.M, 3748.M, 3777.M, 3789.M;

N piébougou: 3346.M,3417.M; Sébékourani: 998.M; Titiéna:

©ZFMK

128

2267.M; Zamoko: 170.M,602.M,2554.M.

Jean-Francois Trape & Youssouph Mané

Echis leucogaster. A goudoud: 1679.M, 1680.M; Boussouma:

1741.M,1742.M; Bouyanga: 48.M,49.M,50.M,51.M

135.M, 136.M, 302.M, 303.M, 304.M, 305.M,

309.M,310.M, 311.M,

319.M,

327.M,

336.M,

350.M,

359.M,

368.M,

383.M,

407.M,

416.M, 2382.M, 2384.M, 2387.M, 2431.M,

,2434.M,2435.M, 2436.M

2442.M,

2449.M,

2456.M,

2463.M,

2470.M,

2477.M,

2484.M,

2491.M,

2498.M,

4083.M,

4089.M,4091.M,

4097.M,4306.M, 4307.M,

4312.M,4313.M,TR.2868

1689.M,1690.M; Gaoudel

1540.M, 1542.

3534.M, 3536.

3549.M, 3554.

3575.M, 3576.

3584.M, 3585.

3591.M, 3592.M, 3593.

2273.M,

54.M,

307.M

315.M

323.M,

331.M,

346.M,

354.M,

364.M,

376.M,

390.M,

414.M,

2433.M

2440.M,

2447.M,

2454.M,

2461.M,

2468.M,

2475.M,

2482.M,

2489.M,

2496.M,

4081.M,

4088.M,

4096.M,

4311.M,

1688.M,

1539.M,

3532.M,

3548.M,

3572.M,

3581.M,

sa-Foulane:

1552.M,1555.M,1559.

, 308.M,

,316.M,

324.M,

332.M,

347.M,

356.M,

365.M,

377.M,

391.M,

2441.M,

2448.M,

2455.M,

2462.M,

2469.M,

2476.M,

2483.M,

2490.M,

2497.M,

4082.M,

317.M

325.M

333.M

348.M

357.M

366.M

380.M

396.M

,318.M,

,326.M,

,334.M,

»,349.M,

1 399 $M,

,367.M,

,381.M,

,399.M,

2443.M,

2450.M,

2457.M,

2464.M,

2471.M,

2478.M,

2485.M,

2492.M,

4076.M,

4084.M,

4092.M,

»2437.M

2444.M

2451.M

2458.M

2465.M

2472.M

2479.M

2486.M

2493.M

4077.M

4085.M

4093.M

4308.M

2276.M ;

312.M,

320.M,

328.

393.03

Stile

361.

369.

384.

408.

313.M

321.M

329.M

338.M

352.M

362.M

371.M

385.M

409.M

»2438.M,

»2445.M,

,2452.M,

»2459.M,

,2466.M,

a0r4a} 35M,

,2480.M,

,2487.M,

»,2494.M,

,4078.M,

,4086.M,

,4094.M,

,4309.M,

K inani:

,533.M,

306.M,

,314.M,

, 322.M,

, 330.M,

,339.M,

s385. 32M s

, 363.M,

,374.M,

,389.M,

,410.M,

2432.M,

2439.M,

2446.M,

2453.M,

2460.M,

2467.M,

2474.M,

2481.M,

2488.M,

2495.M,

4080.M,

4087.M,

4095.M,

4310.M,

,TR.2869,TR.2870; Doro:

>1531.M,1532.M,

M,1546.M,1547.M,3527.M,

M ,3538.M,3541.M,3545.M,

M,3559.M,3560.M,3564.M,

M,3577.M,3578.M,3579.M,

M,3586.M,3587.M,3588.M,

M,3594.M, 3595.M, 3596.M; Haous-

2275.M,

M,1570.M,1579.M,1581.M,1582.M,

1536.M,

3531.M,

3547.M,

3569.M,

3580.M,

3589.M,

1548.M,

1583.M,1585.M; Koyretao: 1617.M,1619.M,2319.M,2320.M,

2322.M,2323.M,2324.M,2326.M, 2327.M, 2328.M,

2330.M,2331.M,2334.M,2337.M,2341.M,2342.M,

3504.M,3505.M,3507.M, 3511.M,3513.M,3514.M,

3518.M, 3519.M, 3521.M, 3526.M, TR.1482; Léré:

2321.M,

2329.M,

3502.M,

3517.M,

2346.M ;

2926.M,

2943.M,

2958.M,

3870.M,

3895.M,

4052.M,

1748.M,

1904.M,

Niam ou:

426.M,

438.M,

450.M,

459.M,

2929.M,2930.M,2931.M,2937.M,2939.M,

2944.M,2947.M,2950.M,2952.M

2960.M,2961.M,2962.M,2963.M

3871.M,3875.M,3878.M, 3884.M

,2955.M,

,2965.M,

,3886.M,

3925.M,3935.M,3943.M,3964.M, 3973.M,

4292.M,

4293

.M,4294.M, 4295.M;

Samé

1749.M,1753.M,1896.M,1900.M,1901.M,

1905.M,1907.M,1908.M,1909.M,1913.M,

490.M,

2941.M,

2957.M,

2973.M,

3893.M,

3985.M,

Ouolof:

1902.M,

1914.M,

1284.M,

1291.M,

1298.M,

1306.M,

1313.M,

2724.M,

2731.M,

2738.M,

2745.M,

2752.M,

2759.M,

2766.M,

1285.M,

1292.M,

1299.M,

1307.M,

1314.M,

2725.M,

1286.M,

1293.M,

1301.M,

1308.M,

2719.M,

2726.M,

2732.M,2733.M,

2739.M,2740.M,

2746.M,2747.M,

2753.M,2754.M,

2760.M,2761.M,

2767.M,2768.M,

2773.M,2774.M,2774.M,

2779.M,2780.M,2781.M,

2786.M,2787.M.

1287.M,

1294.M,

1302.M,

1309.M,

2720.M,

2727.M,

2734.M,

2741.M,

2748.M,

2755.M,

2762.M,

2769.M,

2775.M,

2782.M,

1288.M,

1295.M,

1303.M,

1310.M,

272:1iM-,

27-28 M *,

2735.M,

2742.M,

2749.M,

2756.M,

2763.M,

2770.M,

2776.M,

2783.M,

1289.M,

1296.M,

1304.M,

1311.M,

277. 2322M;

2729.M,

2736.M,

2743.M,

2750.M,

2757.M,

2764.M,

2771.M,

2777.M,

2784.M,

1290.M

1297.M

1305.M

1312.M

2723.M

2730.M

2737.M

2744.M

2751.M

2758.M

2765.M

2772.M

2778.M

2785.M

Echis ocellatus (com plex, including E. jogeri). Ballabougou:

2526.M, 283.M

137.M

2527.M, 2528.M; Bangaya: 32.M, 33.M,

294.M; Bouyanga: 131.M, 132.M, 133.M, 134.M,

335.M, 397.M, 402.M, 413.M, 415.M, 4079.M, 4090.M

TR.2874; 1116.M, 1142.M, 1162.M, 1173.M

1178.M,1183.M,1191.M,1203.M,1204.M,1206.M,1207.M

1210.M,1216.M,1219.M,1221.M,1224.M,1225.M,1226.M

1227.M, 1228.M; Doussoudiana: 73.M, 80.M, 92.M, 95.M

Djinagué:

99.M,106.M,107.M,116.M,121.M,122.M,124.M,TR.0716,

TR.0918,TR.1270,TR.1307; Koundian: 68.M,70.M,2370.M

3042.M; 755.M, 775.M, 786.M, 791.M, 794.M

799.M,800.M, 810.M, 811.M, 834.M, 835.M, 837.M, 840.M

844.M,846.M, 849.M, 850.M, 853.M, 855.M, 857.M, 859.M

2621.M, TR.0926, RAD 2315 TR.1276,

M am oroubougou: 179.M,180.M, 1986.M,1994.M, 2021.M

2022.M,2028.M, 2032.M, 2034.M, 2037.M, 2039.M, 2043.M

2045.M,2047.M,2057.M,3704.M,3732.M,3740.M,3744.M

3776.M,3783.M,3788.M,3799.M, 3806.M,3807.M,3808.M

3816.M,3817.M,3843.M,3847.M, 3856.M,3857.M,3818.M

3836.M,3846.M,4098.M,4099.M,4100.M,4101.M,4102.M

4105.M,4108.M,4109.M,4112.M,4113.M,4114.M,4115.M

4116.M,4117.M,4119.M,4120.M,4121.M,4122.M,4124.M

4125.M,4126.M,4128.M,4129.M,4133.M,4135.M,4137.M

4139.M, 4170.M, 4298.M, TR.2879; Niakoni: TR.0930

TR.0931, TR.1232, TR.1299; Niamou: 35.M, 37.M, 39.M

43.M,45.M,46.M,47.M,57.M,435.M,439.M,441.M,446.M

449.M,451.M, 452.M, 453.M, 456.M,457.M, 458.M, 464.M

465.M,466.M,470.M,474.M,475.M,476.M,477.M, 481.M

482.M,483.M,485.M,486.M,487.M,491.M,493.M,494.M

496.M,498.M,500.M,501.M,503.M,504.M,505.M, 506.M

507.M,508.M,509.M,510.M,517.M,518.M,519.M,520.M

521.M,522.M,523.M,525.M,526.M,527.M,528.M,2867.M

2924.M ,2925.M, 2927. 2928.M,2932.M,2933.M,2934.M

2935.M,2936.M, 2938.M,2940.M ,2942.M,2945.M,2946.M

2948.M,2949.M,2951.M,2953.M,2954.M,2956.M,2959.M

2964.M ,2966.M,2967.M,2968.M,2969.M,2970.M,2971.M

Laminina:

TR.1280;

3061.M,3064.M,3069.M,3071.M,3074.M,3077.M,

Séoulasso: 1399.M, 1416.M; Tacharane: 1889.M,

1891.M,1892.M,1893.M,1894.M,1895.M,2277.M,2278.M

2279.M, 2280.M; 1621.M, 1625.M, 1626.M

1627.M,1628.M,1629.M,1630.M,1631.M,1632.M, 1634.M

1635.M, 1636.M, 1637.M, 2282.M, 2283.M, 2284.M

Topokhoné: 127.M; Toumboula: 1232.M, 1233.M, 1234.M

1236.M,1237.M,1238.M,1239.M,1240.M,1241.M,1242.M

1243.M,1244.M,1245.M,1246.M,1247.M,1248.M,1249.M

1250.M,1251.M,1252.M,1253.M,1254.M,1255.M,1256.M

1258.M,1260.M,1261.M,1262.M,1263.M,1265.M,1266.M

1267.M,1268.M,1269.M,1270.M,1271.M,1273.M,1274.M

1275.M,1276.M,1277.M,1278.M,1279.M,1280.M,1282.M

1890.M

Tinjem ban:

Bonn zoological Bulletin 66 (2): 107-133

3078.M ;

2972.M

3385.M

3455.M

3473.M

3486.M

3860.M

3867.M

3877.M

3887.M

3896.M

3903.M

3910.M

3917.M

2974.M,

3443.M,

3456.M,

3474.M,

3487.M,

3861.M,

3868.M,

3879.M,

3888.M,

3897.M,

3904.M,

3911.M,

3918.M,

339° lhy

3446

3457

3479

3492

3862

3869

3880

3889

3898

3905

3912

3919

SS ££

3356.M,

3447.M,

3458.M,

3480.M,

3493.M,

3863.M,

3872.M,

3881.M,

3890.M,

3899.M,

3906.M,

3913.M,

3920.M,

3362.M,

3448.M,

3459.M,

3482.M,

3496.M,

3864.M,

3873.M,

3882.M,

3891.M,

3900.M,

3907.M,

3914.M,

3921.M,

3370.M,

3450.M,

3466.M,

3484.M,

3858.M,

3865.M,

3874.M,

3883.M,

3892.M,

3901.M,

3908.M,

3915.M,

3922.M,

3380.M

3453.M

3470.M

3485.M

3859.M

3866.M

3876.M

3885.M

3894.M

3902.M

3909.M

3916.M

3923.M

’

©OZFMK

The snakes of Mali

3924.M,3926.M, 3927.M,3928.M,3929.M,3930.M, 3931.M,

3932.M,3933.M,3934.M,3936.M, 3937.M,3938.M,3939.M,

3940.M,3941.M,3942.M,3944.M,3945.M,3946.M,3947.M,

3948.M,3949.M,3950.M,3951.M,3952.M,3953.M,3954.M,

3955.M,3956.M,3957.M,3958.M,3959.M,3960.M,3961.M,

3962.M,3963.M,3965.M,3966.M, 3967.M,3968.M, 3969.M,

3970.M,3971.M,3972.M,3974.M,3975.M,3976.M,3977.M,

3978.M,3979.M,3980.M,3981.M,3982.M,3983.M,3984.M,

3986.M,3987.M, 3988.M,3989.M,3990.M,3991.M,4044.M,

4045.M,4046.M,4047.M,4048.M,4049.M,4050.M,4051.M,

TR.0253,TR.0254,TR.0255,TR.2872,TR.2873; N piébougou:

1789.M,1800.M,1814.M,1816.M,1819.M,1833.M,1834.M,

1842.M,1850.M,1864.M,1873.M,1874.M,1877.M,1883.M,

3373.M,3993.M,3995.M,3997.M,4000.M,4001.M,4006.M,

4007.M,4009.M ,4010.M,4194.M,4195.M,TR.2875,TR.2876,

TR.2877; Sadjouroubougou: 1316.M, 1318.M, 1319.M,

1330.M,2515.M, 2516.M, 2517.M, 2518.M, 2519.M; Sébék-

ourani: 151.M,880.M,898.M,911.M,915.M,916.M,917.M,

923.M,929.M,930.M,934.M,936.M,939.M, 941.M, 943.M,

945.M,946.M,947.M,949.M,951.M,954.M,955.M, 960.M,

963.M,968.M,969.M,970.M,973.M,974.M,976.M,977.M,

979.M,1004.M,1005.M,1008.M,1010.M, 1011.M, 1015.M,

4041.M, 4042.M, 4043.M; Zamoko:

529.M,

598.M,

615.M,

626.M,

651.M,

661.M,

675.M,

685.M,

697.M,

709.M,

720.M,

731.M,

2568.M

2643.M

2696.M

2713.M

4057.M

4065.M

4072.M

568 .M

599 .M

616.M

627.M

652.M

663.M

676.M

686.M

698.M

711.M

721.M

574.M

604.M

617.M

631.M

653.M

665.M

677.M

687.M

699.M

714.M

722.M

585

607.

621.

633.

654.

666.

678.

688.

701.

PLS:

724

.M,

SSS ESE EEE 8

.M,

589.M

608.M

622.M

634.M

655.M

667.M

680.M

690.M

703.M

716.M

157.M,

591.M,

611.M

623.M

636.M

656.M

668.M

682

691

706

Galw

732.M, 2535.M, 2539.M, 2547.M

»2569.M,2577.M,2581.M,2596.M

,2645.M,2649.M

,2699.M,2700.M

»2714.M,2715.M

»4058.M,4059.M

,4066.M,4067.M

»4073.M,4074.M,4075.M.

»2657.M,2659.M

,2709.M,2710.M

»4053.M,4054.M

,4060.M,4061.M

,4068.M,4069.M

725.M 727,

158.M

594.M

612.M

624.M

645.M

658.M

673.M

683.M

694.M

707.M

718.M

728.M

2548.M,

,2601.M,

,2665.M,

,2711.M,

,4055.M,

,4062.M,

,4070.M,

, 159.M

129

,597.M

,613.M

, 625.M

, 646.M

, 660.M

, 674.M

, 684.M

, 695.M

, 708.M

, 719.M

, 729.M

2564.M

2625.M

2666.M

2712.M

4056.M

4064.M

4071.M

Elapsoidea semiannulata moebiusi. Bangaya: 270.M ; Dji-

nagué: 1117.M; Doussoudiana: 110.M, TR.0713; Laminina:

740.M; Mamoroubougou:

3772.M;

N piébougou:

1787.M

1017.M,1018.M,1019.M,1020.M,1023.M,1024.M,1025.M,

1031.M,1032.M,1034.M,1035.M,1038.M,1039.M,1042.M,

1046.M,1047.M,1049.M,1052.M,1053.M,1056.M,1057.M,

1058.M,1059.M,1063.M,1065.M,1066.M,1067.M,1068.M,

1070.M,1071.M,1072.M,1073.M,1074.M,1076.M,1077.M,

1078.M,1080.M,1083.M,1084.M,1085.M,1087.M,1088.M,

1089.M,1090.M,1091.M,1092.M,1093.M,1094.M,1095.M,

1096.M,1097.M,1098.M,1099.M,1101.M,1102.M,1107.M,

3189.M,3200.M,3227.M,3260.M,3276.M,TR.2880; Séoulas-

so: 1346.M, 1347.M, 1349.M, 1350.M, 1352.M, 1353.M,

1355.M,1356.M,1357.M,1359.M,1360.M,1361.M,1362.M,

1363.M,1365.M,1366.M,1367.M,1368.M,1369.M,1370.M,

1371.M,1373.M,1374.M,1375.M,1376.M,1377.M,1378.M,

1379.M,1383.M,1384.M,1385.M,1386.M,1387.M,1389.M,

1391.M,1393.M,1396.M,1397.M,1401.M,1402.M,1404.M,

1405.M,1406.M,1407.M,1408.M,1409.M,1410.M,1411.M,

1413.M,1414.M,1415.M,1417.M,1418.M,1419.M,1420.M,

1421.M,1422.M,1423.M,1426.M,1431.M,1432.M,1433.M,

1434.M,1435.M, 1436.M,1437.M,1438.M,1439.M,1440.M,

1441.M,1442.M, 1443.M,1444.M,1447.M,1448.M,1449.M,

1451.M,1452.M, 1453.M,1454.M,1455.M,1456.M,1459.M,

1461.M,1462.M, 1464.M,1467.M,1468.M,1469.M,1470.M,

1471.M,1472.M,1473.M,1474.M,1476.M,1478.M,1479.M,

1480.M,1481.M,1482.M,1483.M,1484.M,1486.M,1487.M,

1488.M,1489.M,1490.M,1491.M,1492.M,1496.M,1498.M,

1500.M,1502.M,1503.M,1504.M,1506.M,1507.M,1508.M,

1509.M,1510.M,1514.M,1517.M,1518.M,1519.M,1520.M,

1521.M,1523.M,1524.M,1525.M,1526.M,1527.M,1528.M,

1529.M,1691.M,1692.M,1693.M,1694.M,1695.M,1696.M,

1697.M,1698.M,1699.M,1700.M,1701.M,1702.M,1703.M,

1704.M,1705.M,1706.M,1707.M,1708.M,1709.M,1710.M,

1711.M,1712.M, 3136.M, 3292.M, TR.1736, TR .2243; Titié-

na: 200.M, 201.M, 204.M, 207.M, 208.M, 210.M, 212.M,

2067.M,2069.M,2083.M,2092.M,2094.M,2100.M,2114.M,

2119.M,2120.M,2135.M,2147.M,2150.M,2157.M,2159.M,

2166.M,2167.M,2173.M,2176.M,2181.M,2187.M,2189.M,

2193.M,2198.M,2201.M,2204.M, 2208.M,2211.M,2215.M,

2217.M,2227.M,2229.M,2236.M,2240.M,2245.M,2246.M,

2250.M,2253.M,2254.M,2257.M,2258.M,2264.M, 2266.M,

2268.M,2269.M,3612.M,3614.M, 3615.M, 3623.M, 3644.M,

3646.M,3649.M, 3650.M, 3657.M, 3663.M, 3666.M, 3669.M,

3670.M,3672.M, 3681.M, 3685.M, 3688.M 3673.M, 3687.M,

4017.M,4019.M,4026.M,4029.M,4035.M,4037.M,4040.M

Bonn zoological Bulletin 66 (2): 107-133

1835.M,1837.M,1858.M,1859.M,1876.M, 3329.M, 3330.M,

3343.M, 3384.M, 3426.M, 3451.M, 4317.M; Sébékourani:

143.M, 145.M; Zamoko: 164.M, 165.M, 166.M, 167.M,

2582.M,2591.M,2629.M.

Eryx muelleri. Bangaya: 2.M; Bouyanga: 340.M, 341.M,

342.M, 343.M,370.M, 378.M, 382.M, 386.M, 387.M, 388.M,

405.M, 406.M, 2402.M, 2403.M, 2404.M, 2405.M, 2406.M,

2407.M,2408.M,2409.M,2410.M, 2411.M,2412.M, 2413.M,

2414.M,2415.M,2416.M,2417.M, 2418.M, 2419.M, 2420.M,

2421.M,2422.M,2423.M,2424.M, 2425.M, 2426.M, 2427.M,

2428.M, 2429.M, 2430.M, 2499.M; Doro: 1681.M, 1682.M,

1683.M,1684.M,1685.M,1686.M,1687.M; Gaoudel: 3529.M,

3540.M, 3555.M, 3566.M, 3570.M, 3599.M, 3601.M; Haous-

sa-Foulane: 2274.M; Koundian: 3006.M, 3014.M; Koyretao:

2332.M,2333.M,2338.M, 2339.M,3506.M, 3516.M, 3520.M,

3525.M ;Léré:2344.M,2347.M,2348.M; Niamasso: 1345.M;

Niamou: 41.M,56.M, 418.M, 423.M, 425.M, 427.M, 428.M,

430.M, 432.M, 433.M, 434.M, 440.M, 443.M, 444.M, 445.M,

447.M,448.M,454.M,455.M, 460.M, 462.M, 467.M,471.M,

472.M,473.M,495.M,502.M,511.M,512.M,524.M,2800.M,

2801.M,2802.M,2803.M, 2804.M, 2805.M, 2806.M, 2807.M,

2808.M,2809.M,2810.M,2811.M,2812.M,2813.M,2814.M,

2815.M,2816.M,2817.M,2818.M,2819.M,2820.M,2821.M,

2822.M,2823.M,2824.M, 2825.M, 2826.M, 2827.M, 2828.M,

2829.M,2830.M,2831.M,2832.M, 2833.M, 2834.M, 2835.M,

2836.M,2837.M,2838.M,2839.M, 2840.M, 2841.M,2842.M,

2843.M,2844.M,2845.M, 2846.M, 2847.M, 2848.M, 2849.M,

2850.M,2851.M,2852.M, 2853.M, 2854.M, 2855.M, 2856.M,

2857.M,2858.M,2859.M, 2860.M, 2861.M, 2862.M, 2863.M,

2976.M;Samé Ouolof: 1899.M; Séoulasso: 1460.M; Tinjem -

ban: 1623.M; Toum boula: 1259.M,1283.M,1300.M,2793.M.

SS EE

Gonionotophis granti. Lam inina: 831.M;Mamoroubougou:

4110.M; Npiébougou: 3469.M; Sébékourani: 998.M.

Graya smithi. M am oroubougou: 1984.M, 1985.M, 1997.M,

2004.M,2019.M, 3694.M, 3698.M, 3699.M, 3713.M, 3725.M,

3730.M, 3733.M, 3738.M, 3758.M, 3765.M.

Lycophidion albomaculatum. Djinagué: 1132.M, 1138.M,

1152.M, 1197.M, 1214.M; Koundian: 3020.M, 3022.M,

3028.M; Niamou: 2982.M, 2998.M, 2999.M; Sébékourani:

©ZFMK

130 Jean-Francois Trape & Youssouph Mané

146.M, 147.M, 148.M,922.M,983.M,985.M,986.M,988.M,

989.M, 996.M, 997.M, 1003.M, 1016.M, 1084.M, 3176.M,

3216.M; Zamoko: 155.M, 632.M, 712.M, 713.M, 2590.M,

2667.M.

Lycophidion irroratum. M am oroubougou: 1946.M, 3701.M,

3728.M,3795.M, 3812.M.

Lycophidion semicinctum. Lam inina: 739.M,745.M,750.M,

2620.M;Mamoroubougou: 1981.M,1983.M,3700.M,3756.M,

3757.M, 3792.M, 3825.M, 3840.M, 4301.M; Npiébougou:

1849.M, 1860.M,1882.M, 3348.M, 3379.M, 3436.M, 3463.M,

3489.M, 4002.M; Titiéna: 2128.M, 3653.M,4175.M.

Mehelya crossi. Lam inina: 760.M,765.M;Mamoroubougou:

177.M,1978.M,1992.M, 2008.M, 2018.M, 2031.M, 2055.M,

3723.M, 3739.M, 3764.M; Npiébougou: 1776 M, 1783.M,

3392.M; Sadjouroubougou: 2520.M; Sébékourani: 142.M,

150.M, 869.M, 3153.M; Titiéna: 193.M, 2105.M, 2106.M,

2121.M,2131.M,2136.M,3613.M, 3622.M, 3634.M, 3642.M,

3647.M; Zamoko: 173.M,619.M.

Meizodon coronatus. Bangaya: 10.M; Bouyanga: 2501.M;

3702.M; Sébékourani: 984.M, 999.M,

1012.M,1036.M,1103.M,1104.M,1106.M, 3156.M, 3221.M,

3226.M, 3228.M, 3278.M; Titiéna: 202.M, 213.M, 2141.M,

2252.M, 3604.M, 4302.M.

M am oroubougou:

Myriopholis adleri. séb ékourani: 4260.M.

Myriopholis albiventer. 5298.M;

Mam oroubougou: 4288.M, 4320.M,TR.3476, TR .3477.

Doussoudiana:

Myriopholis boueti. Bamako: TR.1686; Bouyanga: 3304.M,

3305.M,3306.M,4181.M,4182.M,4183.M,4184.M,4185.M,

4186.M; Djinagué: 3294.M ,3295.M; Koundian: 3296.M ; Ni-

4196.M; Npiébougou: 3495.M, 3996.M, 4003.M,

4004.M,4005.M,4008.M,4291.M,4266.M,4267.M,4268.M,

4269.M,4270.M,4271.M,4272.M,4273.M,4274.M,4275.M,

4276.M,4277.M,4278.M,4279.M,4280.M,4281.M,4282.M,

4283.M, 4284.M, 4285.M; Samé Ouolof: TR.1685; Sébé-

kourani: 144.M,1100.M, 3283.M, 3284.M, 3285.M, 3286.M,

3287.M,4261.M,4262.M,4263.M; Zamoko:1920.M,4197.M,

4198.M,4199.M,4200.M,4201.M,4202.M,4203.M,4204.M,

4205.M,4206.M,4207.M,4208.M,4209.M,4210.M,4211.M,

4212.M,4213.M,4214.M,4215.M,4216.M,4217.M,4218.M,

4219.M,4220.M,4221.M,4222.M,4223.M,4224.M,4225.M,

4226.M,4227.M,4228.M,4229.M,4230.M,4231.M,4232.M,

4233.M, 4234.M, 4235.M, 4236.M, 4237.M, 4238.M,

4239.M .4581.M,4582.M,4583.M, 4584.M, 4585.M, 4586.M,

4587.M,4588.M,4589.M,4590.M,4591.M,4592.M,4593.M,

4594.M,4595.M,4596.M,4597.M,4598.M,4599.M, 4600.M,

4601.M,4602.M,4603.M,4604.M,4605.M,4606.M, 4607.M,

4608.M,4609.M,TR.2520.

am ou:

Naja katiensis. Ballabougou: 1323.M; Djinagué: 1108.M,

1125.M,1157.M, 1187.M; Doussoudiana: 77.M, 81.M, 85.M,

93.M,1764.M,TR.0913,TR.1234,TR.1271; Laminina: 764.M,

770.M, 785.M, 787.M, 797.M, 807.M, 829.M, 830.M, 833.M,

845.M ,847.M,TR.0921,TR.0924,TR.1229;Mamoroubougou:

176.M, 1987.M, 1996.M, 1998.M, 2000.M, 2001.M, 2005.M,

2010.M, 2013.M, 2015.M, 2020.M, 2023.M, 2030.M, 2040.M,

2041.M, 2042.M, 2051.M, 2052.M, 2054.M, 2061.M, 3690.M,

3695.M, 3697.M, 3703.M, 3705.M, 3706.M, 3707.M, 3708.M,

3709.M,3710.M, 3714.M, 3715.M, 3726.M, 3729.M, 3736.M,

3742.M,3743.M, 3752.M, 3755.M, 3760.M, 3763.M, 3768.M,

3769.M,3770.M, 3773.M, 3775.M, 3778.M, 3779.M, 3787.M,

3815.M, 3821.M, 3822.M, 4299.M; Npiébougou: 1778.M,

Bonn zoological Bulletin 66 (2): 107-133

1784.M,1785.M,1794.M,1797.M,1798.M,1801.M,1807.M,

1811.M,1812.M,1813.M,1818.M,1820.M,1821.M,1825.M,

1826.M,1832.M,1844.M,1848.M,1851.M,1855.M,1857.M,

3311.M,3314.M,3315.M,3316.M, 3317.M, 3318.M, 3319.M,

3°32, 05M... -3'3'2)1, Mi... 3'3:22..M 3.3: 25-M", 313.26. M_,/333.2'8:.M °3.3:3,22M. ,

3334.M,3335.M, 3336.M, 3338.M, 3339.M, 3342.M, 3344.M,

3345.M,3347.M,3350.M, 3352.M, 3353.M, 3354.M, 3355.M,

3358.M,3359.M, 3360.M, 3363.M, 3364.M, 3365.M, 3366.M,

3367.M,3371.M,3372.M,3377.M, 3378.M, 3381.M, 3383.M,

3387.M,3389.M,3390.M, 3391.M, 3393.M, 3394.M, 3395.M,

3396.M,3397.M,3399.M,3400.M, 3401.M, 3403.M, 3405.M,

3406.M,3407.M, 3408.M, 3409.M, 3410.M, 3411.M,3412.M,

3413.M,3418.M, 3421.M,3424.M , 3429.M, 3430.M, 3431.M,

3433.M,3438.M, 3439.M,3440.M, 3444.M, 3445.M, 3460.M,

3464.M,3468.M, 3472.M, 3475.M, 3483.M, 3488.M, 3497.M,

3994.M;Sadjouroubougou: 2503.M,2504.M,2505.M,2506.M,

2507.M;Sébékourani: 875.M,964.M ,991.M,3152.M,3167.M,

3172.M, 3206.M, 3258.M; Titiéna: 181.M, 182.M, 183.M,

185.M,186.M,187.M,188.M,189.M,190.M,197.M, 203.M,

205.M, 214.M, 216.M, 2071.M, 2076.M, 2079.M, 2080.M,

2095.M,2110.M,2111.M,2113.M,2139.M,2158.M,2179.M,

2182.M,2183.M,2191.M,2206.M,2212.M,2219.M,2223.M,

2243.M,2248.M,2251.M,3603.M, 3605.M, 3608.M, 3620.M,

3624.M,3625.M, 3626.M, 3627.M, 3628.M, 3629.M, 3630.M,

3631.M,3632.M, 3633.M, 3635.M, 3636.M, 3643.M, 3648.M,

3651.M,3654.M, 3655.M, 3656.M, 3658.M, 3661.M, 3662.M,

3664.M,3667.M, 3676.M, 3677.M, 3678.M, 3679.M, 3680.M,

3682.M, 3686.M; Zamoko: 530.M, 531.M, 532.M, 533.M,

534.M,535.M,536.M,537.M,538.M,539.M,540.M,541.M,

542.M,543.M,544.M,545.M,546.M,547.M,548.M,549.M,

550.M,551.M,552.M,553.M,554.M,555.M,556.M,557.M,

558.M,559.M,563.M,569.M,571.M,573.M,576.M,578.M,

579.M,580.M, 603.M,605.M, 618.M, 630.M, 638.M, 639.M,

640.M, 647.M, 681.M, 705.M, 710.M, 2529.M, 2530.M,

2536.M,2540.M,2550.M,2551.M,2566.M,2586.M,2587.M,

2597.M,2598.M,2600.M, 2603.M, 2607.M, 2615.M,2627.M,

2630.M,2631.M, 2632.M, 2634.M, 2635.M, 2638.M, 2650.M,

2651.M,2655.M,2674.M,2675.M, 2676.M, 2678.M, 2680.M,

2681.M,2682.M, 2683.M, 2684.M, 2685.M, 2686.M, 2687.M,

2703.M,2708.M.

Naja melanoleuca. Bangaya: 245.M; Laminina: 767.M;

Mamoroubougou: 2027.M, 2048.M, 3716.M; Npiébougou:

1782.M,3312.M,3420.M,3476.M; Sébékourani: 995.M; Titié-

na: 2087.M,2101.M

Naja nigricollis. Bangaya: 3.M, 7.M, 29.M, 217.M, 218.M,

235.M, 246.M; 1738.M, 1739.M, 1740.M;

K oundian: 60.M, 62.M, 3026.M, 3029.M, 3031.M, 3037.M;

M am oroubougou: 1990.M,3727.M; Niamasso: 1342.M; Ni-

amou: 421.M,2981.M,2989.M,2990.M; Sébékourani: 897.M,

3173.M, 3185.M, 3191.M, 3192.M, 3198.M,;

1390.M, 3132.M; Toya: 1642.M, 2298.M, 2302.M, 2312.M;

Zamoko: 565.M,584.M,609.M,2541.M,2614.M.

Boussouma:

Séoulasso:

Naja senegalensis. Ballabougou: 2353.M, 2354.M; Bangaya:

238.M; Djinagué: 1179.M, 1181.M; Doussoudiana: 103.M;

K oundian: 2368.M; 805.M, TRV 2-723"

Mamoroubougou: 1977.M, 2003.M, 2017.M; Npiébougou:

1796.M, 3419.M; Sadjouroubougou: 2352.M; Saré-Soma:

1591.M;Sébékourani: 878.M ,957.M; Titiéna: 184.M,2102.M,

2109.M,2118.M,2145.M,2186.M, 3606.M,3617.M, 3618.M,

3683.M; Zamoko: 156.M,581.M, 2349.M, 2350.M, 2351.M.

Lam inina:

Philothamnus irregularis. D oussoudiana: 75.M,78.M, 87.M,

89.M, 91.M, 114.M, 1756.M, 1774.M, TR.1305; Laminina:

©OZFMK

The snakes of Mali 131

759.M,771.M,772.M,774.M,777.M,783.M, 821.M, 822.M,

TR.0925, TR.1275, TR.1277, TR.1279, TR.1308, TR.1310,

TR.1311,TR.1332; Niakoni: TR.0927; N piébougou: 1775.M,

1847.M,3309.M ,3349.M, 3414; Sébékourani: 980.M ,982.M,

993.M, 994.M, 3162.M, 3196.M, 3211.M, 3233.M; Titiéna:

2078.M,2098.M,2224.M, 3607.M.

Philothamnus semivariegatus smithi. D oussoudiana: 1766.M,

1767.M; Laminina: 751.M; Niamou: 1918.M.

Polemon neuwiedi. Mamoroubougou

4162.M.

1935.M, 4159.M,

Prosymna collaris. Top okhoné: 129.M.

Prosymna greigerti. Bangaya: 289.M; N piébougou: 4314.M;

Djinagué: 1209.M; Koundian: 2372.M; Laminina: 741.M,

T47.M, 752.M, 756.M, 823.M, 854.M; Mamoroubougou:

1933.M,1938.M,1982.M,3712.M,3719.M,3737.M,3753.M,

3805.M,3832.M, 3835.M,3851.M,4160.M,4296.M,4297.M,

4300.M; Niamou: 1922.M; Npiébougou: 1822.M, 3323.M,

3435.M, 3461.M, 3471.M, 3481.M; Sébékourani: 149.M,

152.M, 967.M, 981.M, 992.M, 3261.M, 3269.M; Titiéna:

1925.M, 1926.M, 1927.M, 1929.M, 1931.M, 4172.M; Toum-

boula: 1235.M; Zamoko: 154.M,1919.M.

Psammophis elegans. Bangaya: 13.M; Bouyanga: 52.M;

Doussoudiana: 82.M,88.M,96.M,100.M,117.M,118.M; Ki-

nani: 1554.M,1568.M,1572.M,1573.M,1577.M; Koundian:

2367.M,3007.M,3015.M,3017.M,3024.M, 3036.M,3040.M,

3041.M; Laminina: 779.M, 780.M, 793.M, 803.M, 826.M,

TR.0923, TR.1226, TR.1228; Mamoroubougou: 178.M,

2012.M, 2025.M, 3711.M, 3735.M, 3741.M, 3837.M;

Npiébougou: 1786.M, 1788.M, 1791.M, 1803.M, 1805.M,

1828.M,1830.M, 1836.M,1839.M,1841.M,1879.M,1880.M,

3341.M,3423.M,3494.M;Sadjouroubougou: 2508.M,2509.M,

2510.M,2511.M;Samé Ouolof: 3045.M ,3062.M;Saré-Soma:

1608.M; Sébékourani: 883.M, 928.M, 1062.M, 3195.M,

3209.M, 3214.M; Séoulasso: 1398.M, 1427.M, 1428.M,

1429.M,1445.M,1463.M,1466.M, 1477.M, 3083.M, 3097.M,

3100.M,3103.M,3145.M,3149.M; Ténintou: TR .0688; Titié-

na: 2160.M,3638.M,3665.M; Zamoko:577.M,582.M,587.M,

620.M,650.M, 2588.M, 2611.M, 2626.M, 2679.M.

Psammophis lineatus. M am oroubougou: 3754.M, 3811.M.

Psammophis philippsi. Lam inina: 822.M .

Psammophis praeornatus. M am oroubougou: 3838.M ,3845.M ;

Niam ou: 2985.M; Npiébougou: 3374.M; Sadjouroubougou:

1317.M; Samé Ouolof: 1915.M; Sébékourani: 3229.M;

Séoulasso: 1358.M, 1392.M, 1403.M, 1446.M, 1501.M,

1522.M,3096.M; Titiéna: 2096.M,2107.M,2127.M,2200.M,

2209.M,3637.M; Zam oko: 2545.M,2567.M,2570.M, 2636.M,

2656.M,2662.M, 2663.M, 2673.M.

Psammophis schokari. G aoudel: 1545.M; Tinjem ban: 1622.M,

1624.M, 2281.M.

Psammophis att. sibilans. 4 goudoud: 1677.M, 1678.M; Bal-

labougou: 1320.M, 1321.M,1322.M, 2523.M, 2524.M; Ban-

gaya: 4.M, 6.M, 8.M, 16.M, 24.M, 220.M, 222.M, 230.M,

247.M,248.M, 249.M,250.M, 252.M, 262.M, 267.M, 268.M,

274.M,287.M; Boussouma: 1724.M,1725.M,1726.M,1727.M,

1728.M,1729.M,1730.M,1731.M,1732.M,1733.M,1734.M,

1735.M,1736.M; Bouyanga: 344.M, 345.M, 355.M, 360.M,

372.M,373.M,375.M,379.M, 392.M, 393.M, 394.M, 395.M,

398.M, 404.M, 411.M, 2375.M, 2377.M, 2378.M, 2379.M,

2380.M, 2383.M,2386.M, 2389.M, 2390.M, 2391.M,2392.M,

Bonn zoological Bulletin 66 (2): 107-133

2393.M,2394.M ,2395.M ,2396.M, 2397.M, 2398.M,2399.M,

2400.M, 2401.M, TR.2871; Djinagué: 1110.M, 1113.M,

1122.M,1129.M,1130.M,1139.M,1144.M,1155.M,1159.M,

1164.M,1165.M,1167.M,1170.M,1171.M,1172.M,1175.M,

1182.M,1193.M,1199.M,1202.M,1211.M,1212.M,1213.M,

1215.M,1217.M,1222.M,1223.M; Donguiba: 3308.M;Dous-

soudiana: 83.M, 101.M, 108.M, 115.M, 1755.M, 1757.M,

1770.M, TR.0720, TR.1237, TR.1238, TR.1268, TR.1272,

TR.1306; Gaoudel: 1530.M,1533.M,1534.M,1537.M,1543.M,

1556.M,3530.M, 3535.M, 3537.M, 3539.M, 3542.M, 3543.M,

3544.M,3546.M,3550.M,3551.M,3552.M, 3553.M, 3556.M,

3557.M,3562.M,3563.M,3565.M, 3568.M, 3574.M, 3582.M,

3590.M ,3597.M ,3598.M ,3600.M ,3602.M; Haoussa-Foulane:

2270.M,2272.M; Kinani: 1549.M,1550.M,1553.M,1557.M,

1558.M,1560.M,1561.M,1562.M,1563.M,1565.M,1566.M,

1567.M,1569.M,1574.M,1575.M,1576.M,1580.M; Koundi-

an: 63.M,3002.M ,3003.M,3004.M,3005.M, 3008.M,3009.M,

3011.M,3013.M,3016.M, 3023.M, 3030.M, 3033.M, 3038.M,

3039.M; Koyretao: 1618.M, 3499.M; Laminina: 773.M,

781.M, 801.M, 802.M, 818.M, 825.M, 827.M, 832.M, 838.M,

839.M, 852.M, 2623.M, TR.1312, TR.1313; Léré : 2343.M,

2345.M;Mamoroubougou: 1999.M, 2024.M,2053.M,2062.M,

2065.M,3718.M,3803.M,3804.M,3810.M,3814.M,3852.M,

3854.M,3855.M; Niakoni: TR.0886, TR.0926, TR.1298; Nia-

masso: 1324.M, 1325.M, 1326.M, 1327.M, 1328.M, 1329.M,

1331.M,1332.M,1333.M,1334.M,1335.M,1336.M,1337.M,

1338.M,1339.M,1340.M;Niamou: 34.M,42.M,58.M,59.M,

419.M,429.M, 431.M, 463.M, 468.M, 469.M, 478.M,479.M,

480.M,488.M, 489.M,492.M,499.M,513.M,514.M,515.M,

516.M, 2864.M, 2865.M, 2866.M, 2868.M, 2869.M, 2870.M,

2871.M,2872.M,2873.M,2875.M,2876.M,2877.M,2878.M,

2879.M,2880.M,2881.M,2882.M, 2883.M,2884.M,2885.M,

2886.M,2887.M,2888.M,2889.M, 2890.M,2891.M,2893.M,

2894.M,2895.M,2896.M,2897.M,2898.M,2899.M,2900.M,

2901.M,2902.M,2903.M,2904.M,2905.M,2906.M,2907.M,

2908.M,2909.M,2910.M,2911.M,2912.M,2913.M,2914.M,

2915.M,2916.M,2917.M,2918.M,2919.M,2920.M,2921.M,

2922.M, 2975.M, 2992.M, TR.0256; Npiébougou: 1777.M,

1793.M,1802.M,1824.M,1827.M,1845.M,1846.M,1863.M,

1865.M,1866.M,1871.M,1872.M, 3310.M, 3331.M,3333.M,

3340.M,3382.M,3402.M, 3404.M, 3432.M, 3437.M, 3442.M,

3452.M; Samé Ouolof: 1743.M, 1744.M, 1745.M, 1746.M,

1747.M,1750.M,1751.M,1897.M,1898.M,1903.M,1906.M,

1910.M,1911.M,1912.M,3044.M, 3046.M, 3047.M, 3048.M,

3049.M,3050.M, 3051.M,3052.M, 3053.M, 3054.M, 3055.M,

3056.M,3057.M,3058.M, 3059.M, 3060.M, 3063.M, 3065.M,

3066.M,3067.M, 3068.M, 3070.M, 3072.M, 3073.M, 3075.M,

3076.M, 3079.M, 3080.M, 3081.M; Saré-Soma: 1586.M,

1587.M,1588.M,1589.M,1590.M,1592.M,1593.M,1595.M,

1596.M,1597.M,1598.M,1599.M,1600.M,1601.M,1602.M,

1603.M,1604.M,1605.M,1606.M,1607.M,1609.M,1610.M,

1611.M, 1612.M, 1613.M, 1615.M, 1616.M; Sébékourani:

863.M,871.M,881.M,884.M,953.M,966.M,1007.M,1021.M,

1022.M,1026.M,1029.M,1044.M,1051.M,1054.M,1055.M,

1061.M,1064.M,3154.M,3157.M, 3160.M, 3165.M,3170.M,

3174.M,3178.M,3184.M, 3186.M, 3187.M, 3188.M, 3190.M,

3193.M,3194.M,3197.M,3205.M, 3207.M, 3213.M, 3215.M,

3219.M,3220.M,3223.M,3234.M, 3238.M, 3240.M, 3242.M,

3244.M,3250.M,3251.M,3252.M,3254.M, 3255.M,3256.M,

3257.M,3259.M,3268.M, 3271.M,3275.M,3277.M, 3291.M;

Séoulasso: 1348.M, 1354.M, 1364.M, 1381.M, 1382.M,

1394.M,1400.M,1424.M,1425.M,1430.M,1458.M,1475.M,

1485.M,1494.M,1495.M,1497.M,1499.M,1511.M,1512.M,

1513.M,1515.M,1516.M,1716.M,1717.M,1718.M,1719.M,

1720.M,1721.M,1722.M,3082.M, 3084.M, 3085.M, 3087.M,

©ZFMK

132 Jean-Francois Trape & Youssouph M ané

3088.M,3089.M,3090.M, 3091.M, 3092.M, 3093.M, 3095.M,

3098.M,3099.M,3101.M,3104.M,3105.M,3106.M, 3108.M,

3109.M,3110.M,3112.M,3113.M,3114.M,3115.M,3116.M,

3117.M,3118.M,3119.M,3120.M, 3121.M,3122.M,3123.M,

3124.M,3125.M,3126.M,3127.M, 3129.M, 3131.M, 3133.M,

3135.M,3137.M,3138.M,3139.M,3140.M, 3141.M,3142.M,

3143.M,3144.M,3146.M,3147.M,3148.M,3150.M,3151.M;

Tacharane: 1885.M, 1886.M, 1887.M, 1888.M; Tinjem ban:

1620.M, 1633.M, 2285.M; Titiéna: 192.M, 196.M, 199.M,

206.M, 209.M, 2068.M, 2070.M, 2072.M, 2086.M, 2090.M,

2091.M,2097.M,2126.M,2129.M,2140.M,2142.M,2143.M,

2146.M,2148.M,2149.M,2152.M,2161.M,2163.M,2169.M,

2172.M,2177.M,2185.M,2188.M,2190.M,2196.M,2197.M,

2210.M,2213.M,2216.M,2222.M,2228.M,2231.M,2238.M,

2239.M,2244.M,2247.M,2249.M, 2259.M,2260.M,2261.M,

2265.M; Toumboula: 1264.M, 1315.M, 2789.M, 2792.M,

2795.M; Toya: 1638.M, 1639.M, 1640.M, 1641.M, 1643.M,

1644.M,1645.M,1646.M,1647.M,1648.M,1649.M,1650.M,

1651.M,1652.M,1653.M,1654.M,1655.M,1656.M,1657.M,

1658.M,1659.M,1660.M,1661.M,1662.M,1663.M, 1664.M,

1665.M,1666.M,1667.M,1669.M,1670.M,1671.M,1672.M,

1673.M,1674.M,1675.M,1676.M,2286.M,2287.M,2288.M,

2289.M,2290.M,2291.M,2292.M ,2293.M,2294.M,2296.M,

2297.M,2299.M,2300.M, 2301.M, 2303.M, 2304.M,2305.M,

2306.M,2307.M,2308.M, 2309.M, 2310.M, 2311.M,2313.M,

2314.M,2315.M,2316.M,2317.M,2318.M; Zamoko: 560.M,

561.M,562.M,572.M,575.M,592.M,601.M, 606.M, 628.M,

644.M, 659.M, 700.M, 733.M, 734.M, 2543.M, 2544.M,

2546.M,2553.M,2556.M,2560.M,2561.M,2565.M,2571.M,

2573.M,2578.M,2579.M,2583.M,2584.M,2589.M, 2604.M,

2610.M,2612.M,2617.M,2618.M, 2628.M, 2637.M, 2639.M,

2644.M,2652.M,2660.M, 2661.M, 2668.M, 2669.M, 2670.M,

2671.M,2677.M,2688.M, 2689.M, 2695.M, 2697.M, 2698.M,

2704.M,2705.M,2716.M,2717.M, 2718.M.

Python regius. Ballabougou: 2522.M; Djinagué: 1115.M,

1135.M,1218.M; Doussoudiana: 74.M,102.M,113.M,120.M,

1754.M, 1759.M, 1772.M, TR.0903, TR.1269, TR.1300;

K oundian: 2365.M; Laminina: 768.M,798.M,809.M,814.M,

815.M,817.M,TR.1278; Niakoni: TR.1281; Niamou : 442.M;

Sadjouroubougou: 2502.M; Sébékourani: 860.M, 861.M,

862.M, 866.M, 873.M, 879.M, 886.M, 896.M, 900.M,903.M,

912.M, 931.M, 971.M, 1033.M, 1045.M, 3164.M, 3265.M,

3273.M; Titiéna: 2117.M,2132.M,2162.M,2214.M, 2221.M,

3616.M, 3619.M, 3671.M; Zamoko: 586.M, 596.M, 635.M,

2624.M.

Python sebae. Bangaya: 22.M; Boussoum a:1737.M; Bouyan-

ga: 401.M ; Djinagué: 1189.M ; Laminina: 758.M, 769.M,

808.M; Niamou: 417.M, 424.M, 437.M, 2984.M, 2992.M;

N piébougou: 3368.M; Samé Ouolof: 1752.M; Sébékourani:

865.M,885.M,890.M,891.M,937.M,1048.M,3272.M; Titié-

na: 2089.M; Toum boula: 1257.M.

Rhagerhis moilensis. G aoudel: 1538.M, 1541.M, 1544.M,

3533.M, 3558.M; Gogui (10 km SS): 3307.M; Haoussa-

Foulane: 2271.M; Koyretao: 3501.M.

Rhamphiophis oxyrhynchus. Bangaya: 5.M; Bouyanga:

2376.M,2381.M,2385.M, 2388.M; Djinagué: 1220.M; Dous-

soudiana: 76.M,109.M,1762.M,TR.0719,TR.0914,TR.1302;

Laminina: 789.M, 804.M, TR.0648; Mamoroubougou:

3828.M, 3834.M, 3849.M; Niakoni: TR.0928, TR.1297; Ni-

amou:497.M,2991.M,2996.M;Npiébougou: 1776.M,1781.M,

1790.M,1792.M,1795.M,1804.M,1831.M,1838.M,1853.M,

1867.M,1878.M,1881.M,3369.M, 3388.M, 3415.M, 3422.M,

Bonn zoological Bulletin 66 (2): 107-133

3427.M;Sébékourani: 958.M,972.M,987.M,1006.M,1028.M

.M, 3183

1380.M,1412.M,1450.M,1457.M,1465.M,1723.M,3102.M

3130.M, 3134.M; Titiéna:

,2103.M,2108.M,2115.M, 2116.M, 2133.M, 2138.M

,2175.M,2184.M,2192.M, 2195.M, 2205.M, 2207.M

,2225.M,2230.M, 2232.M,2233.M, 2237.M, 2255.M

3639.M,3640.M, 3645.M, 3660.M,4022.M; Zamoko: 588.M

643.M, 726.M, 2531.M, 2549.M, 2592.M, 2599.M, 2701.M

1030.M

3107.M

1928.M

2155.M

2220.M

» 3159.M, 3181

, 3111.M,

2706.M.

.M;

Séoulasso:

194.M,

1372.M

198.M

Rhinoguinea magna. M amoroubougou: 1956.M, 4142.M

TR.2501,

TR .3479,

4143.M

TR .2823,

» 4144.M,

TR.3481, TR.3482.

4318.M,

TR.2824, TR.2825,

4319.M,

TR .3478,

TR.2822,

TR .3480,

Rhinoleptus koniagui. D jinagué: 4187.M; K oundian: 2374.M

Laminina:

749.M,

753.M,

828.M ;

4177.M,4178.M,4179.M,4180.M,4287.M.

N piébougou:

3998.M

Spalerosophis diadema. G aoudel: 3528.M ,3571.M; Koyretao:

2325.M, 2336.M, 3508.M, 3515.M, 3522.M; Toya: 1668.M.

Telescopus tripolitanus. kK inani: 1551.M, 1571.M, 1578.M

Toum boula: 1229.M, 2788.M, 2790.M, 2791.M, 2798.M.

Telescopus variegatus. K oundian: 3027.M;M am oroubougou:

3721.M,3833.M; Zamoko:

Zam oko: 168.M

169.M, 629.M, 2613.M, 2653.M

Tricheilostoma bicolor. Doussoudiana: 3280.M, 3281.M,

3282.M, TR.1323; Laminina: 842.M; Mamoroubougou:

1934.M,1943.M,1945.M,1948.M, 1949.M,1950.M,1954.M,

1958.M,1962.M, 1963.M,1966.M, 1968.M,1969.M,1970.M,

1971.M,1972.M,1973.M,1974.M, 1975.M,4169.M, 4289.M,

4610.M,4611.M,4612.M, 4613.M, 4614.M, 4615.M, 4616.M,

4617.M,4618.M, 4619.M, 4620.M, 4621.M, 4622.M, 4623.M,

4624.M,4625.M, 4626.M, 4627.M, 4628.M, 4629.M, 4630.M,

4631.M,4632.M, 4633.M, 4634.M, 4635.M, 4636.M, 4637.M,

4638.M,4639.M, 4640.M, 4641.M, 4642.M, 4643.M, 4644.M,

4645.M,4646.M, 4647.M, 4648.M, 4649.M, 4650.M, 4651.M,

4652.M,4653.M, 4654.M, 4655.M, 4656.M, 4657.M, 4658.M,

4659.M,4660.M, 4661.M, 4662.M, 4663.M, 4664.M, 4665.M,

4666.M,4667.M, 4668.M, 4669.M, 4670.M, 4671.M, 4672.M,

4673.M,4674.M, 4675.M, 4676.M, 4677.M, 4678.M, 4679.M,

4680.M,4681.M, 4682.M, 4683.M, 4684.M, 4685.M, 4686.M,

4687.M,4688.M, 4689.M, 4690.M, 4691.M, 4692.M, 4693.M,

4694.M,4695.M, 4696.M, 4697.M, 4698.M, 4699.M, 4700.M,

4701.M,4702.M,4703.M,4704.M, 4705.M,4706.M,4707.M,

4708.M,4709.M,4710.M,4711.M, 4712.M,4713.M,4714.M,

4715.M,4716.M,4717.M,4718.M, 4719.M, 4720.M,4721.M,

4722.M,4723.M,4724.M,4725.M, 4726.M, 4727.M, 4728.M,

4729.M,4730.M, 4731.M, 4732.M, 4733.M, 4734.M, 4735.M,

4736.M,4737.M, 4738.M, 4739.M, 4740.M,4741.M,4742.M,

4743.M,4744.M,4745.M,4746.M, 4747.M, 4748.M,4749.M,

4750.M,4751.M,4752.M,4753.M, 4754.M, 4755.M,4756.M,

4757.M,4758.M,4759.M,4760.M, 4761.M, 4762.M, 4763.M,

4764.M,4765.M,4766.M,4767.M, 4768.M,4769.M,4770.M,

4771.M,4772.M,4773.M,4774.M, 4775.M,4776.M,4777.M,

4778.M,4779.M, 4780.M,4781.M, 4782.M, 4783.M, 4784.M,

4785.M,4786.M,4787.M,4788.M, 4789.M,4790.M,4791.M,

4792.M,4793.M,4794.M,4795.M, 4796.M,4797.M,4798.M,

4799.M,4800.M, 4801.M, 4802.M, 4803.M, 4804.M, 4805.M,

4806.M,4807.M, 4808.M,4809.M,, 4810.M,4811.M,4812.M,

4813.M,4814.M, 4815.M, 4816.M, 4817.M, 4818.M, 4819.M,

4820.M,4821.M, 4822.M, 4823.M, 4824.M, 4825.M, 4826.M,

4827.M,4828.M, 4829.M, 4830.M, 4831.M, 4832.M, 4833.M,

©ZFMK

4834.M

4841.M

4848.M

4855.M

4862.M

4869.M

4876.M

4883.M

4890.M

4897.M

4904.M

4911.M

4918.M

4925 .M

4932.M

4939.M

4946.M

4953.M

4960.M

4967.M

4974.M

4981.M

4988.M

4995 .M

5002.M

5009.M

5016.M

5023.M

5030.M

5037.M

5044.M

5051.M

5058.M

5065.M

5072.M

5079.M

5086.M

5093.M

5100.M

5107.M

5114.M

5121.M

5128.M

4835.

4842.

4849.

4856.

4863.

4870.

4877.

4884.

4891.

4898.

4905.

4912.

4919.

4926.

4933.

4940.

4947.

4954.

4961.

4968.

4975.

4982.

4989.

4996.

5003.

, 5010

5017.

5024.

3.03.1.

5038.

5045.

5052.

5059.

5066.

S073"

5080.

5087.

5094.

S101.

5108.

al bp)

12:2.

29%,

SESS SESS SEES SSS REESE SSE SESE ERE EEE EE EE ESSERE EEE EEE EEE

4836.

4843.

4850.

4857.

4864.

4871.

4878.

4885.

4892.

4899.

4906.

4913.

4920.

4927.

4934.

4941.

4948.

4955.

4962.

4969.

4976.

4983.

4990.

4997.

5004.

20-11

5018.

5025.

5032.

5.039:

5046.

3;0;5,3-.

5060.

5067.

5074.

5081.

5088.

5095.

5102.

5109.

5116.

34123.

5.03102

SSS ESSE EE KE SEESSESEEKE EEE SEEEEEES

SSS ES EES ERE SEES EES

, 4837.

, 4844.

»4851

» 4858.

» 4865.

» 4872.

» 4879.

» 4886.

» 4893.

» 4900.

» 4907.

4914.

»4921.

»4928.

» 4935.

4942.

4949.

4956.

4963.

» 4970.

> 49° FT...

4984.

»4991.

4998.

» 5005.

; 904 2

wd0e1 97

» 5026.

» 5033.

» 5040.

, 5047.

» 5054.

» 5061

, 5068.

10°73:

» 5082.

» 5089.

, 5096.

>9-4,0;3..

xed: LON

Fae ie ee ae

,5124.

Ripe Ba Soles

S22 Ss iz Beg igide ee) eS BS ee Sees 2S Ss Se Se oe eee ee ee) eee ce se

4838.

4845.

4852.

4859.

48066.

4873.

4880.

4887.

4894.

4901

4908.

4915.

4922.

4929.

4936.

4943.

4950.

4957.

4964.

AOR

4978.

4985.

4992.

4999.

5006.

5013

5020.

5027.

5034.

5041.

5048.

5055.

5062

5069.

5076.

5083.

5090.

5097

5104.

oes ks

5118.

S125:

SSi2s

Seep AR eC het gd gl ll RE ey a i agg AR a a A i ag a GR ew ee

Bonn zoological Bulletin 66 (2): 107-133

4839.M

4846.M

4853.M

4860.M

4867.M

4874.M

4881.M

4888.M

4895.M

4902.M

4909.M

4916.M

4923.M

4930.M

4937.M

4944.M

4951.M

4958.M

4965.M

4972.M

4979.M

4986.M

4993.M

5000.M

5007.M

5014.M

5021.M

5028.M

5035.M

5042.M

5049.M

5056.M

5063.M

5070.M

5077.M

5084.M

5091.M

5098.M

5105.M

5112.M

5119.M

5126.M

5133.M

The snakes of Mali

4840.M

4847.M

4854.M

4861.M

4868.M

4875.M

4882.M

4889.M

4896.M

4903.M

4910.M

4917.M

4924.M

4931.M

4938.M

4945.M

4952.M

4959.M

4966.M

4973.M

4980.M

4987.M

4994.M

5001.M

5008.M

5015.M

5022.M

5029.M

5036.M

5043.M

5050.M

5057.M

5064.M

5071.M

5078.M

5085.M

5092.M

5099.M

5106.M

5113.M

5120.M

5127.M

5134.M

31355

5142.

5149.

5156.

D163

14205,

SET

5184.

S191.

5198.

5205.

322 12k.

5219.

5226.

D233"

5240.

5247.

5254.

5261.

5268.

3; 22732.

5282.

5289.

5296

Niam ou:

SESS EEE EES SEES SESE ESEESE EES

5136.M

5143.M

5150.M

5157.M

5164.M

5171.M

5178.M

5185.M

5192.M

5199.M

5206.M

5213.M

5220.M

5227.M

5234.M

5241.M

5248.M

5255.M

5262.M

5269.M

5276.M

5283.M

5290.M

140.M,

»,5291.M,5292.M

.M,5297.M.,TR.2846,TR.2847; Titiéna: 4173.M,

4265.M;

5137.M

5144.M

5151.M

5158.M

5165.M

5172.M

5179.M

5186.M

5193.M

5200.M

5207.M

5214.M

5221.M

5228.M

5235.M

5242.M

5249.M

5256.M

5263.M

5270.M

5277.M

5284.M

141.M,

,5138.M

»5145.M

»5152.M

»,5159.M

,5166.M

,5173.M

,5180.M

,5187.M

»5194.M

,5201.M

,5208.M

,5215.M

,5222.M

,5229.M

,5236.M

,5243.M

,5250.M

,5257.M

,5264.M

,5271.M

,5278.M

»5285.M

5139.M

5146.M

5153.M

5160.M

5167.M

5174.M

5181.M

5188.M

5195.M

5202.M

5209.M

5216.M

5223.M

5230.M

5237.M

5244.M

5251.M

5258.M

5265.M

5272.M

5279.M

5286.M

5293.M

5140.M

5147.M

5154.M

5161.M

5168.M

5175.M

5182.M

5189.M

5196.M

5203.M

5210.M

5217.M

5224.M

5231.M

5238.M

5245.M

5252.M

5259.M

5266.M

5273.M

5280.M

5287.M

5294.M

N piébougou:

4189.M,4190.M, 4191.M, 4192.M, 4193.M, 4290.M

ourani: 3288.M, 3289.M, 3297.M, 3298.M, 3299.M,

»3302.M,3303.M

3301.

4242.

4249.

4256.

4504.

4511

4518.

4525.

4532.

4539.

4546.

4553.

4560.

4567.

4574.

cue Sashes ee eS Sees

4243.M,4244.M

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4554.M

4561.M

4568.M

4575.M

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4258.M

4506.M

4513.M

4520.M

4527.M

4534.M

4541.M

4548.M

4555.M

4562.M

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;Zamoko: 1917.M,4240.

»4245.M

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4557.M

4564.M

4571.M

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4509.M

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4523.M

4530.M

4537.M

4544.M

4551.M

4558.M

4565.M

4572.M

4579.M

133

5141.M,

5148.M,

5155.M,

5162.M,

5169.M,

5176.M,

5183.M,

5190.M,

5197.M,

5204.M,

5211.M,

5218.M,

5225.M,

5232.M,

5239.M,

5246.M,

5253.M,

5260.M,

5267.M,

5274.M,

5281.M,

5288.M,

5295.M,

4174.M ;

4188.M,

; Sébék-

3300.M,

4241.M,

4248.M,

4255.M,

4503.M,

4510.M,

4517.M,

4524.M,

4531.M,

4538.M,

4545.M,

4552.M,

4559.M,

4566.M,

4573.M,

4580.M.

©ZFMK

Bonn zoological Bulletin 66 (2): 135-137

Additional records of the Arabian Sand Cat

Felis margarita harrisoni (Hemmer, Grubb & Groves, 1976)

(Carnivora: Felidae) in Iraq

Omar F, Al-Sheikhly’* & Mukhtra K. Haba’

' Department of Biology, College of Science, University of Baghdad, Baghdad, Iraq

*Department of Biology, College of Science for Women, University of Baghdad, Baghdad, Iraq

* Corresponding author. E-mail: alsheikhlyomar@gmail.com

Abstract. The Arabian Sand Cat Felis margarita harrisoni 1s a rare nocturnal feline associated with desert and arid steppes.

The species has been recently recorded to Iraq from three specimens by Mohammad et al. (2013). However, its current

status and population trends in Iraq still unknown. In 2014 and 2015, additional records with first photographic evidence

December 2017

were made. These new records confirm the resident occurrence of this species in Iraq.

Key words. Arabian Sand Cat, Carnivora, Felidae, Iraq, wild mammals.

INTRODUCTION

Sand Cat Felis margarita is a cryptic nocturnal feline well

adapted to desert and arid plains. It 1s similar to Wild Cat

Felis silvestris but noticeably smaller and paler. The head

is rather flat with broad and large ears that lack the api-

cal dark tufts in their tips. The vibrissae are white; the

limbs are of medium length and marked with by at least

two pronounced black elbow bars and with broad paws;

the palms and soles are covered with a dense mat of long

wavy hair which conceals the pads. The pelage is soft and

dense with abundant soft woolly underfur, and the col-

oration is strikingly pallid with a pale sandy isabelline tint

in the back. The face is marked with dark brownish stripes

from the anterior canthus of each eye backwards across

the cheeks. The tail is tipped with black and has two or

three subterminal black bars (Harrison & Bates 1991).

Four subspecies have been described: F' m. margarita in

North Africa; m. harrisoni in Arabia; F m. thinobia in

Central Asia; and Fm. scheffeli in Pakistan (Sliwa 2013;

Banfield et al. 2014). The Arabian Sand Cat F' m. harrisoni

is known from many localities across the Middle East, but

its current status and distribution across the Arabian Penin-

sula are not known in detail. Its population size is uncer-

tain; it seems to be declining but at an unknown rate (Mal-

lon & Budd 2011). It was recorded from Oman (type lo-

cality), Yemen, Saudi Arabia, United Arab Emirates, Qatar,

Kuwait, Jordan, Syria, Israel, Sinai, and Iran (Harrison &

Bates 1991; Cunningham 2002; Serra et al. 2007; Strauss

et al. 2007; Sher Shah & Cunningham 2008; Mallon &

Budd 2011; Banfield et al. 2014; Ghadirian et al. 2016).

Received: 15.08.2017

Accepted: 12.10.2017

The status of the Arabian Sand Cat in Iraq is still uncer-

tain. It is a cryptic and rarely seen feline, confined to veg-

etated sandy desert, sand dunes, sand/gravel steppes, and

rocky valleys (wadies) of south and western Iraq (AI-

Sheikhly et al. 2015). Iraq’s first record was based on three

specimens (two males and one female). One of the males

and the female were found alive at a private nursery in

Baghdad on 27 January 2012. They were collected from

a desert area situated in the west of Al-Najaf city in Al-

Najaf province. A third specimen (another male) was a

mounted skin which appeared to be has been trapped from

Al-Jufaira oasis in the desert of Al-Najaf province in No-

vember 2012 (Mohammad et al. 2013).

NEW RECORDS

Three records (four specimens: three adult males and a ju-

venile female) were made (Table 1). On 28 February 2014,

an adult male and juvenile female were presented at the

local animal market in Baghdad. They were collected from

desert area near Al-Shabakah (Shbecha) in Al-Najaf

Province (Fig.1). Another adult male was presented at the

local animal market in Baghdad on 7 March 2014. It was

trapped near Takhadid (c.4 0 km) to the southwest of Ni-

grat Al-Salman in Al-Muthana Province (a note on this

record is found in Al-Sheikhly et al. 2015). On 14 April

2015, an adult male was presented at the local animal mar-

ket in Baghdad. It was trapped in a desert area (c. 20 km)

northwest of Al-Nekheab in Al-Anbar Province (Fig. 2).

Sand Cats are vulnerable to indiscriminate trapping, but

Corresponding editor: R. Hutterer

136 Omar F. Al-Sheikhly & Mukhtra K. Haba

Fig. 1. Juvenile female Arabian Sand Cat Felis margarita har-

risoni trapped near Al-Shabakah (Shbecha) in Al-Najaf Province

and presented at the local animal market in Baghdad.

unlikely to be directly targeted. They are sometimes caught

for the international pet trade which is an ongoing activ-

ity, but the scale is not known (Mallon & Budd 2011). H-

legal trapping with weak implementation of the hunting

laws has been highlighted as a major threat on wildlife in

Iraq (Al-Sheikhly et al. 2015). The newly recorded Sand

cats were accidentally trapped by local truffle collectors

who become active during rainy seasons (January —April)

in southern and western deserts of Iraq. Interviews indi-

cated that Sand cats were coincidently observed by truf-

fle collectors at night and early morning. Sand cats acci-

dentally flashed by the collectors’ spotting torchlight,

chased until they get tired and rest among desert vegeta-

tion, and then are caught by hand. After a few days, Sand

cats are brought to Baghdad by the truffle collectors and

presented in the local animal markets in order to be sold

as pets. The trapped cats were stressed, suffering from

careless handling, and exhausted due to long distance

Fig. 2. Adult male Arabian Sand Cat Felis margarita har-

risoni in captivity trapped Al-Nekheab in Al-Anbar Province.

transportation. The young female seemed to be born in the

wild which suggests that Sand cats are breeding in suit-

able habitats in the southern and western deserts of Iraq

(Fig.1).

The ecological requirements of Sand cats in Iraq and

around the region are still poorly understood. Due to the

rarity of the species, its distribution, status and the impact

of threats are difficult to assess. Apart from new spatial-

ly and temporally localized records in a few range coun-

tries, no easily measurable changes within its distribution

area are apparent (Sliwa 2013). Therefore, additional re-

search needs to be performed to document the present con-

tinuous occurrence and to assess its status.

Acknowledgments. We are grateful the Iraqi Green Climate Or-

ganization (IGCO) for providing information on Sand Cats in

Iraq. We would like to thank Hisham Kher Allah for providing

additional information on Sand cat sightings in Iraq.

Table 1. Records of the Arabian Sand Cat Felis margarita harrisoni in Iraq.

Number Site Coordinates Date of trapping Province Reference

1 Male West of Al-Najaf city - January 2012 Al-Najaf Mohammad et al. 2013

Female West of Al-Najaf city - January 2012 Al-Najaf Mohammad et al. 2013

3 Male Al-Jufaira oasis — November 2012. = Al-Najaf Mohammad et al. 2013

in the desert of Al-Najaf

4 Male (Ad.) near Al-Shabakah (Shbecha) 30°48’N February 2014 Al-Najaf New observation

43°40’°E

5 Female (Juv.)

6 Male (Ad.) — Takhadid (c.40km) 30°11’N March 2014 Al-Muthana New observation

southwest Nigr’at Al-Salman 44°13’E

‘l Male (Ad.) — (c. 20) km northeast 32° 6’N April 2015 Al-Anbar —_— New observation

of Al-Nekheab 42° 2’E

Bonn zoological Bulletin 66 (2): 135-137

©OZFMK

Additional records of the Arabian Sand Cat Felis margarita harrisoni in Iraq 137

REFERENCES

Al-Sheikhly OF Haba MK, Barbanera E, Csorba G, Harrison DL

(2015) Checklist of the Mammals of Iraq (Chordata: Mam-

malia). Bonn Zoological Bulletin 64 (1): 33-58

Banfield LM, Al Qahtani H, Mallon D (2014) Arabian Sand Cat

Felis margarita harrisoni Status Reveiw and Conservation

Strategy. Al Ain Zoo, Abu Dhabi, United Arab Emirates

Bunaian F, Mashaqbeh S, Yousef M, Buduri A, Amr Z (1998)

A new record of the Sand Cat, Felis margarita, from Jordan.

Zoology in the Middle East 16: 5—7

Cunningham P (2002) Status of the Sand Cat, Felis margarita,

in the United Arab Emirates. Zoology in the Middle East 25:

9-14

Ghadirian T, Akbari H, Mohammadreza B, Ghoddousi A, Hami-

di A Kh, Dekhordi ME (2016) Sand cat in Iran: present sta-

tus, distribution and conservation challenges. Cat News Spe-

cial Issue 10 (in press).

Harrison DL, Bates PJJ (1991) The Mammals of Arabia. Har-

rison Zoological Museum. Sevenoaks, Kent, U.K.

Mallon D, Budd K (eds) (2011) Regional Red List Status of Car-

nivores in the Arabian Peninsula. Cambridge, UK and Gland

Bonn zoological Bulletin 66 (2): 135-137

Switzerland: IUCN, and Sharjah, UAE: Environment and Pro-

tected Areas Authority. 49 pp

Mohammad MK, Lahony SR, Al-Rammahi HM (2013) First

record of the Sand cat, Felis margarita Loche, 1858 (Mam-

malia: Carnivora, Felidae), from Iraq. Zoology in the Middle

East 59(4): 358-359

Serra G, Abdallah M S, Al Quaim G (2007) Occurrence of Rup-

pell’s fox Vulpes rueppelli and Sand cat Felis margarita in Syr-

ia. Zoology in the Middle East 42: 99-101

Sher Shah M, Cunningham P (2008) Fences as a threat to Sand

cats Felis margarita Loche, 1958, in Saudi Arabia. Zoology

in the Middle East 42: 99-101

Sliwa A (2013) Felis margarita. In: Kingdon J S & Hoffmann

M (eds), The Mammals of Africa, Academic Press, Amster-

dam, The Netherlands

Sliwa A, Breton G, Chevalier F (2013) Sand cat sightings in the

Moroccan Sahara. Cat News 59: 28-30

Strauss WM, Shobrak M, Sher Shah M (2007) First trapping re-

sults from a new sand cat study in Saudi Arabia. Cat News

47: 20-21

©OZFMK

Bonn zoological Bulletin 66 (2): 139-144

December 2017

Six new records of Afrotropical lizard and snake species

(Reptilia: Squamata)

from the Republic of South Sudan

Klaus Ullenbruch' & Wolfgang Bohme?*

' KindtalstraBe 6b, D-56745 Bell, Germany

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

“Corresponding author. E-mail: w.boehme @ leibniz-zfmk.de

Abstract. we report on reptilian specimens collected in southern Sudan (currently the Republic of South Sudan) in 1978

and stored at the Zoologisches Forschungsmuseum Alexander Koenig, Bonn. Six species (one lizard, Leptosiaphos kili-

mensis, and five snakes, Hapsidophrys lineatus, Thrasops jacksoni, Toxicodryas pulverulenta, Amblyodipsas unicolor,

Atheris squamigera) are documented as new records for the fauna of South Sudan and are discussed in a biogeographi-

cal context.

Key words. N ortheastern Africa, new country records, biogeography.

INTRODUCTION

The Herpetology Section of the Zoologisches

Forschungsmuseum Alexander Koenig (ZFMK) in Bonn

was founded in 1951 (Béhme 2014), but half a century

earlier in several missions between 1897 and 1913, the

founder of the museum, Alexander Koenig, had collect-

ed already amphibians and reptiles from all over the for-

mer Sudan, which is today divided into two countries: the

Islamic Republic of Sudan in the north and the Republic

of South Sudan. Between 1976 and 1984 Gerhard N iko-

laus and Hans Rupp explored the avifauna of Sudan (Su-

dan and South Sudan) during several trips (Nikolaus 1987)

and collected a large number of amphibians and reptiles,

as wellas birds and small mammals, from these countries

that were deposited at the ZFMK. The latest acquisition

of Sudanese (Sudan and South Sudan) amphibians and

reptiles reached the ZFMK in the 1980’s, collected by

Hans-Erkmar Back and by Ulrich Joger. In total, speci-

mens representing more than 150 herpetological species

from all over the former entire Sudan are housed atZFMK.

As a first step we studied the preserved snake and lizard

specimens of these collections and examined 485 lizards

belonging to 49 species and 261 snakes belonging to 61

species. In this first note, we recorded and documented

one lizard and five snake species from the Republic of

South Sudan that were not known from this country be-

fore. Other results of our work on this material will fol-

low in further publications.

All specimens cited in the present work were collected

by Gerhard Nikolaus and Hans Rupp during their avifau-

nal surveys in Sudan in 1978. Furthermore, all collecting

Received: 18.08.2017

Accepted: 17.10.2017

sites of the six new country records (Yei, Katire, Gilo,

Kinyeti and surroundings) are situated in the southern part

of South Sudan, i.e. in the historical Equatoria Region (Fig.

1). Yei is situated in Central Equatoria (now Yei River state)

near the border with Uganda and the Democratic Repub-

lic of the Congo (DRC), on the main road that leads from

the South Sudanese capital Juba to Faradje, in the DRC.

The Imatong Mountains, with their highest peak Mt.

Kinyeti (3180 m a.s.l.) and Katire as the major settlement,

are situated in Eastern Equatoria (now Imatong state), al-

so near the Ugandan border. These parts of South Sudan

are characterized by woodland, forests, and highland

forests (Jackson 1956, Friis & Rasmussen 1981, Nikolaus

1987). The Imatong M ountains, in particular, have an im-

portant biogeographical role as a link between the high-

lands of East Africa and Ethiopia, and also as a shelter of

some of the easternmost examples of the Guineo-Congo-

lian rainforests, such as the Lotti and Talanga forests (Fri-

is & Rasmussen 1981).

LIST OF THE NEW COUNTRY RECORDS OF

SQUAMATA FOR SOUTH SUDAN

Order Squamata

Family Scincidae

Leptosiaphos kilimensis (Stejneger, 1891)

Proceedings of the United States National Museum 14

(802): 405-406. Type locality: Kilimanjaro, Tanzania, East

Africa.

Corresponding editor: X. Mengual

140 Klaus Ullenbruch & Wolfgang Bohme

30°30'E 31°E 31°30'E 32°E 32°30'E 33°E

4°30'N -

4. ess) “~

\ tthe ee -e

i #

év - ’

a? - ;

» ‘

J AL | -

4°N d

3°30'N 4

Fig. 1. Map of the Equatoria Region of South Sudan, showing the collection sites of the new records reported in the present

study. 1. Kajiko North (03°47’N, 30°35’E, 1000 m); 2. Iwatoka/Watoka (03°45’N, 30°38’E, 1000 m); 3. Mt. Korobe (03°58’N,

30°52’°E,1590 m); 4. Talanga (04°01’°N, 32°43’E, 950 m); 5. Katire (04°02’N, 32°47’E, 1000 m); 6. Gilo (04°00’N, 31°51’E, 1900

m); 7. Mt. Kinyeti (03°55’N, 32°55’E, 3180 m).

609

Gals ch dail

Fig. 2. Leptosiaphos kilimensis (ZFM K 26059) from Mt. Korobe, South Sudan (locality no. 3 on Fig. 1).

Bonn zoological Bulletin 66 (2); 139-144 ©ZFMK

Six reptilian species new for South Sudan 141

Fig. 3. Hapsidophrys lineata (ZFM K 2626072) from Kajiko

North, South Sudan (locality no. 1 on Fig. 1).

Voucher specimen. South Sudan: south of Yei, Mt. Ko-

robe, collected by G. Nikolaus, 28 July 1978 (ZFMK

26059; Fig. 2).

Remarks. S paw Is et al. (2002) listed this leaf-litter skink

for Tanzania, Kenya, Uganda and elsewhere south-west to

Angola. Identification followed Perret (1982), Broadley

& Howell (1991) and Spawls et al. (2002). Our specimen

of Leptosiaphos kilimensis shows five toes and a lower

eyelid with a central window in contrast to Leptosiaphos

blochmanni, L. meleagris and L. hackarsi with three or

four digits and the lower eyelid scaly, and L. graueri with

the lower eyelid scaly, too. The frontoparietals are not

fused with the interparietal as it is normally the case in L.

rhomboidalis. Specimen ZFM K 26059 (Fig. 2) shows five

digits on forefoot in contrast to L. aloystisabaudiae w ith

only four digits on forefoot.

Family Colubridae

Hapsidophrys lineata Fischer, 1856

A bhandlungen des N aturwissenschaftlichen Vereins Ham-

burg (3) 4: 79-116. Type locality: Elmina, Ghana.

Bonn zoological Bulletin 66 (2): 139-144

Voucher specimen. South Sudan: south of Yei, Kajiko-

North, collected by G. Nikolaus, 20 July 1978 (ZFMK

26072; Fig. 3).

Remarks. S paws et al. (2002) listed this widely distrib-

uted forest species for Uganda, Kenya, DR Congo, else-

where south-westwards to northern Angola and westwards

to Guinea. Schmidt (1923), Pitman (1974) and Broadley

& Howell (1991) reported this species also for Tanzania.

In Uganda, this species is more widely distributed in the

southwest of the country; otherwise there are only a few

scattered records near Entebbe and in the Mabira and

Budongo forests (Spawls et al. 2002). Our voucher spec-

imen from south of Yei (Fig. 3) represents, therefore, a

considerable range extension to the north and not simply

a border-crossing continuation of the Ugandan distribu-

tion range.

Fig. 4. = Thrasops jacksonii (ZFMK 26008) from between

Katire and Talanga, South Sudan (localities nos. 4 and 5 on Fig.

1).

©OZFMK

142 Klaus Ullenbruch & Wolfgang Bohme

Fig.5. = Toxicodryas pulverulenta (ZFM K 26030) from Talan-

ga Forest, South Sudan (locality no. 4 on Fig. 1).

Thrasops jacksoni G iinther, 1895

Annals and Magazine of Natural History (London) (6) 15:

523-529.

Type locality. K avirondo, Kenya.

Voucher specimens. South Sudan: between Katire and Ta-

langa, collected by H. Rupp, 20-25 July 1978 (ZFMK

26008; Fig. 4); South Sudan: Gilo, collected by G. Niko-

laus, 29 October 1978 (ZFMK 29573).

Remarks. Broadley & Wallach (2002) listed “southern

Sudan” as part of the distribution range of this species and

marked this also in a grid map, however, without provid-

ing locality data or a voucher specimen. Upon request, we

were informed by D.G. Broadley (in litt.) that he had based

this statement on an unpublished report by Friis & Ras-

mussen (1981), who collected one specimen of Thrasops

Jacksoni in the Imatong M ountains, South Sudan, between

29 October and 22 December 1980, also without any fur-

ther data. Their voucher specimen is deposited in the Zo-

ological Museum,

ZMUC R 601199 (the “60” being a code number for col-

ubrid snakes). Our two specimens reported here are there-

University of Copenhagen under

fore, together with the ZMC specimen, the first document-

ed records for the South Sudan. Spawls et al. (2002) list-

ed this species only for Uganda, Rwanda, Tanzania, Kenya,

and the Democratic Republic of Congo. According to these

Bonn zoological Bulletin 66 (2): 139-144

authors, this species also seems to be absent from the

northern half of Uganda, so there is a relatively large gap

between the Ugandan records and our new records from

the Imatong area in South Sudan.

Toxicodryas pulverulenta (Fischer, 1856)

A bhandlungen des N aturwissenschaftlichen Vereins Ham-

burg 3 (4): 79-116.

Type locality. Liberia; restricted or “corrected” to Sao

Tomé by Hughes & Barry (1969).

Voucher specimen. South

(Kinyeti), collected by H. Rupp, July 1978 (ZFMK 26030).

Remarks. Spaw ls et al. (2002) listed this forest and wood-

land species (as Boiga pulverulenta) for Kenya, Uganda

Sudan: Talanga-Forest

and the DR Congo, elsewhere westwards to Guinea and

south-westwards to northern Angola. This confirms data

from Pitman (1974) and Chippaux (2001). The Ugandan

localities listed by Spawls et al. (2002, see also the map

by these authors) leave the northern third of the country

without records. Thus, again in this case, the geographic

distance to the new South Sudanese locality of the spec-

imen ZFM K 26030 (Fig. 5) is remarkable.

Fig. 6. Amblyodipsas unicolor (ZFMK 26088) from Gilo,

South Sudan (locality no. 6 on Fig. 1).

©OZFMK

Six reptilian species new for South Sudan 143

Fig. 7.

toka, South Sudan (locality no. 2 on Fig. 1).

Atheris squamigera (ZFM K 26000) from Iwatoka/Wa-

Family Atractaspididae

Amblyodipsas unicolor (Reinhardt, 1843)

Danske Videnskabernes Selskabs Afhandlinger 10:

233-279.

Type locality. Guinea Coast = Ghana.

Voucher specimen. South Sudan: Imatong M ountains, Gi-

lo, collected by G. Nikolaus, 20 June 1978 (ZFMK

26088).

Remarks. The distribution of this species is centred on

West Africa (Spawls etal. 2002, Trape & Mané 2006) with

a few records in East Africa (Tanzania, Kenya, Uganda)

(Schmidt 1923, Pitman 1974). Its presence in the forest-

ed area of the Imatong Mountains in South Sudan, docu-

mented by ZFMK 26088 (Fig. 6), is therefore a remark-

able The

(Spawls et al. 2002: Kampala, Kinja, Masindi) are not

range extension. three Ugandan localities

close to the Ugandan-South Sudanese border.

Bonn zoological Bulletin 66 (2): 139-144

Family Viperidae

Atheris squamigera (Hallowell, 1854)

Proceedings of the Academy of Natural Sciences Philadel-

phia 3(1): 193-194.

Type locality. “near the river Gabon, Guinea”. Accord-

ing to Loveridge (1957), the type locality is in the ‘French

Congo’,a French colony which at one time comprised the

present-day area of the Republic of the Congo, Gabon, and

the Central African Republic.

Voucher specimen. South Sudan: Iwatoka (= Watoka,

near Yei), collected by H. Rupp, September 1978 (ZFMK

26000).

Remarks. In East A frica, this forest-dwelling species has

been documented for U ganda, Kenya and Tanzania; else-

where westwards to Nigeria, Ghana and Ivory Coast and

south-westwards to northern Angola (Spawls & Branch

1995, David & Ineich 1999, Spawls et al. 2002). The

U gandan localities, mostly isolated from each other, are

listed in Spawls et al. (2002), but none of them is close

to the South Sudanese border. These data again highlight

the importance of our new record (Fig. 7) from near Yei.

DISCUSSION

Scale counts of all specimens mentioned above are with-

in the known range characteristic of each species; hence

they are not discussed here. Similarly, the single specimens

on whom our new country records are based would not

be relevant for any scale count comparisons.

All new records are forest species and belong in all cas-

es to known, widely distributed species. Imatong M oun-

tains also harbour endemics, such as the chameleon J7I-

oceros kinetensis (Schmidt, 1943) (Béhme & Klaver

1980), whose relatives are distributed in East Africa, in-

cluding the Eastern Arc M ountains. Only two species from

our new records occur only in East Africa, viz. Leptosi-

aphos kilimensis and Thrasops jacksoni. The former has

a patchy distribution in Uganda, Kenya and Tanzania

(Spawls et al. 2002), the latter reaches also the eastern

forests of the Congo Basin in DRC but is replaced fur-

ther west by other, closely related species (Fischer &

Hinkel 1992, Spawls et al. 2002). The other four species,

viz. Hapsidophrys lineata, Amblyodipsas unicolor, Toxi-

codryas pulverulenta and Atheris squamigera, have a large

distribution through the Congo Basin westwards to

Cameroon, Ghana or even Guinea. Biogeographically, they

may be regarded as Guineo-Congolian faunal elements and

might indicate that the South Sudanese relic rainforests,

including the Imatong Mountains, can be considered as

northernmost outliers of the Guineo-Congolian realm,

with some biogeographic connection to the Eastern Arc

M ountains. There is also one lizard showing a similar bio-

geographic distribution, i.e.a Guineo-Congolian pattern,

©ZFMK

144 Klaus Ullenbruch & Wolfgang Béhme

viz. the forest-dwelling lacertid Adolfus africanus

(Boulenger, 1906), which has been already recorded from

the Imatong Mountains (Kohler et al. 2003).

These six reptile species newly recorded for the fauna

of South Sudan have in common a wide separation from

the nearest Ugandan records, which are not located just

across the Uganda border but considerably further south.

This underscores the importance of the South Sudanese

relic forest patches, including the Imatong M ountains, as

they have biogeographic relationships with both the East

African Arc and the Guineo-Congolian forests. Certain-

ly, the Equatoria Region of South Sudan is greatly under-

studied in terms of its biodiversity, and the number of

squamates newly recorded in this study points out the po-

tential for another hotspot of herpetological diversity in

this country. We hope that the Republic of South Sudan

will soon return to political conditions that make further

basic research and nature conservation possible.

Acknowledgements. We thank Dr. Heinz Grillitsch, Naturhis-

torisches Museum (NHMW), Vienna, and Dr. Van Wallach, Mu-

seum of Comparative Zoology (MCZ), Harvard University, Cam-

bridge, for their support with some references. Thanks are also

due to Daniel Klingberg Johansson, Zoological Museum, Uni-

versity of Copenhagen (ZM UC) Copenhagen, who gave advice

on the ZMUC voucher specimen of T. jacksonii, and to Morris

Flecks (ZFMK) for preparing the map and photographs. We are

grateful to two reviewers whose comments helped to improve

the text.

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©OZFMK

Bonn zoological Bulletin 66 (2): 145—165

December 2017

Review of the Eumerus barbarus species group (Diptera: Syrphidae)

from the western Mediterranean Basin

Jeroen van Steenis'*, Martin Hauser? & Menno P. van Zuijen’

' Research Associate Naturalis Biodiversity Center Leiden. Hof der Toekomst 48,

3823HX Amersfoort, Netherlands

? Plant Pest Diagnostics Centre California, Department of Food and Agriculture. 3294 Meadowview Road Sacramento CA

95832-1448, USA

> Kolkakkerweg 21-2, 6708 RK Wageningen, Netherlands

Corresponding author. E-mail: j.van.steenis @ xmsnet.nl; jvansteenis 1 @ gmail.com

Abstract. The species of the Eumerus barbarus group from the western parts of the Mediterranean Basin are revised.

Two species new to science are described, i.e. Eumerus gibbosus sp.n. (from Portugal and Spain) and Eumerus schmideg-

geri Sp. n. (from Algeria, Morocco and Tunisia). The other two species included in this group are Eumerus barbarus (Co-

quebert, 1804) and Eumerus sulcitibius Rondani, 1868. A neotype is designated for E. barbarus and we also designated

a lectotype for Eumerus iris Loew, 1848. All species are figured, their synonyms are reviewed and an identification key

is presented. Eumerus truncatus Rondani, 1868 is withdrawn from synonymy with E. barbarus and considered a bona

species; additionally, this species is recorded from Morocco, Portugal, Spain and Tunisia for the first time. A short dis-

cussion on the value of the Mediterranean Basin as biodiversity hotspot for hoverflies is given.

Key words. Eumerus barbarus group, new species, description, Mediterranean Basin, hotspot, threat.

INTRODUCTION

Eumerus Meigen, 1822 is a well-defined genus restrict-

ed to the Old World, except for some Australasian species.

It is one of the largest syrphid genera in the world, with

over 250 described species (Pape & Thompson 2013).

Some species have been introduced in other regions, no-

tably the New World, with bulbs transported by man for

commerce (Gibbson 1917; Sasscer 1919; Smit 1928;

Neboiss 1957; Marinoni & Morales 2007; Speight et al.

2013). The larvae are known to feed in or on decaying

roots, bulbs and fleshy matter of Liliaceae, Amerillaceae,

Orobanchaceae and even in New World cacti, and some

species are regarded to be pests of diverse ornamental

bulbs and ginger (Collin 1920; Smit 1928; Martin 1934;

Sathiamma 1979; Pérez-Banén & Marcos-Garcia 1998;

Ricarte et al. 2008).

The present study deals with the species belonging to

the E. barbarus group as defined by Chroni et al. (2017).

Chroni et al. (2017) used the name E. sulcitibius group

for E. aff. barbarus and E. sulcitibius Rondani, 1868. As

the name E. barbarus (Coquebert, 1804) is senior to E.

sulcitibius, the correct name is E. barabarus species group.

This grouping does not reflect a phylogenetic ranking as

the taxonomy and phylogenetics of Eumerus still require

revision (Doczkal 1996; Doczkal & Pape 2009). As the

females of this group are difficult to distinguish from each

other and even from other Eumerus species, the diagno-

Received: 29.07.2017

Accepted: 02.11.2017

sis for each species reviewed here is therefore only based

on males. The characters used in the key to the females

are often subtile and a certain identification is not always

possible.

The aim of the present work is to review the species of

the Eumerus barbarus group from the western parts of the

Mediterranean Basin and to describe two species new to

science, namely Eumerus gibbosus sp. n. (from Portugal

and Spain) and Eumerus schmideggeri sp. n. (from Alge-

ria, Morocco and Tunisia). Moreover, we figure all the

species of the barbarus group, designate a neotype for Eu-

merus barbarus, a lectotype for Eumerus iris Loew, 1848

and provide an identification key. Eumerus truncatus Ron-

dani, 1868 is withdrawn from synonymy with Eumerus

barbarus and considered a bona species.

MATERIAL AND METHODS

The countries considered here as part of the western

Mediterranean Basin are Algeria, France, Italy, Morocco,

Portugal, Spain and Tunisia.

Male genitalia and abdomen were dissected by means

of entomological pins and forceps from dry or softened

specimens. Genitalia and abdomen were boiled in a 10%

KOH solution. Once cleared sufficiently, genitalia and ster-

na IV were placed in glycerine and photographed in a

Nikon SMZ dissecting microscope with a Leica MC 190

Corresponding editor: X. Mengual

146 Jeroen van Steenis et al.

HD mounted camera. The photographs of the male ster-

num IV were touched up in Adobe Photoshop©, and the

photographs of the genitalia were used as a template to

draw them in Adobe IIlustrator©. The photos of the ex-

ternal structures were taken with a Canon EOS 6D mount-

ed on a 2.5 times phototube attached to a Wild M10 stere-

omicroscope. The photos of the habitus and body parts

were processed with Zerene Stacker version 1.04 and fur-

ther edited with GNU Image Manipulation Program

2.8.16. SEM images (Fig. 9) are made with a VEGA 3xm

(Tescan).

Morphological terminology follows Thompson (1999),

and terms related to the male genitalia follow Doczkal

(1996). The measurements were made with the use of a

micrometre. The body is measured from the posterior end

of the antennae to the posterior tip of the abdomen; width

of head is measured at its maximum width in dorsal, re-

spectively frontal view; the width of the face is measured

just below the antennae; width of ocellar triangle is meas-

ured over the posterior ocelli in dorsal view; the length of

the ocellar triangle is measured from the anterior end of

the anterior ocellus to the midline posterior of the poste-

rior ocelli; the length of the frontal triangle is measured

from the anterior corner of the eye contiguity to the pos-

terior corner of the lunule in dorsal view; the length of the

vertical triangle is measured from the posterior corner of

the eye contiguity to the anterior ocellus in dorsal view,

the eye contiguity itself is the length in between the pre-

vious two measurements; the width of the vertex in dor-

sal view is measured posteriorly between the posterior cor-

ner of the eyes (a), over the posterior ocelli (b) and over

the anterior ocellus (c); the length of the basoflagellom-

ere is measured in lateral view from the most anterior part

of the pedicel to the apex of basoflagellomere and the

width is measured at its widest point, perpendicular to the

midline; the length of the metafemur as well as the tar-

someres of the metatarsus is measured in posterior view

from the base to the apex along the midline and the width

is measured at its widest point, perpendicular to the mid-

line.

In the description of type labels, the contents of each

66 99

label is enclosed within double quotation (“”’), italics de-

note handwriting, and the individual lines of data are sep-

arated by a forward slash (/ ).

DEPOSITORIES

AET _ private collection of Andre van Eck, Tilburg,

The Netherlands

CSCA California State Collection of Arthropods,

Department of Food & Agriculture, Sacramento,

California, USA

CEUA Coleccién Entomolégica de la Universidad de

Alicante, Centro Iberoamericano de la

Bonn zoological Bulletin 66 (1): 145-165

Biodiversidad, Alicante, Spain

JSA private collection of Jeroen van Steenis,

Amersfoort, The Netherlands

LACM Los Angeles County Museum, Los Angeles,

California, USA

MNCN Museo Nacional de Ciencias Naturales, Madrid,

Spain

MNHN Laboratoire d’Entomologie, Muséum national

d’ Histoire naturelle, Paris, France

MZUF Museo Zoologico “La Specola”, Firenze, Italy

MZW private collection of Menno van Zuijen,

Wageningen, The Netherlands

NBC_ Natural Biodiversity Centre, Leiden,

The Netherlands

NHM_ The Natural History Museum, London,

United Kingdom

ZMHB Museum fiir Naturkunde, Leibniz-Institut fiir

Evolutions- und Biodiversitétsforschung, Berlin,

Germany

RESULTS

The western Mediterranean species of the Eumerus

barbarus group

The barbarus group contains four Eumerus species in the

western parts of the Mediterranean Basin: E. barbarus, E.

sulcitibius, Eumerus gibbosus sp. n., and Eumerus

schmideggeri sp. n. Until recently, Eumerus truncatus

Rondani, 1868 was regarded as a synonym of E. barbarus.

Grkovi€¢ et al. (2015) treated E. truncatus as bona species

(from several Greek Isles), but without giving any refer-

ence to its recognition or former synonymy. We examined

specimens of this species, including the type material, and

concluded that E. truncatus is not a species of the Eumerus

barbarus group as defined here, which can be seen from

the redescription provided below.

Diagnosis of the Eumerus barbarus species group

As recognised here, the species of the Eumerus barbarus

group share the following characteristics: Face with dense

white pollinosity and white to light-yellow pile (Figs 4A,

4B, 4D-41)), with mouth edge narrowly shiny black. Eyes

white pilose, about 1/4-1/2 times as long as facial pile;

male with rather long eye contiguity, 7-9 ommatidia long

(Figs 3A, 3D, 3F, 3H). Vertex broad, ocellar triangle isosce-

les, located anteriorly on vertex, with widely separated

posterior ocelli; anterior part and two small maculae along

eye margin just posterior of posterior ocelli white polli-

nose (Figs 3A, 3B, 3D-I). Basoflagellomere oval with

rounded apico-dorsal corner (Fig. 6). Thorax with two me-

dial narrow white pollinose vittae on nearly entire length

(Figs 1A, 1B, 1D-—11); on supra-alar region with a row of

©OZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 147

8-12 short black setulae; shiny colour with copper to gold-

en purple sheen. Notopleural suture absent. Abdomen

black with different shiny colours (black, copper to gold)

and with white pollinose lunulate maculae on terga II-IV

and short black and white pile.

Males of the barbarus group can be distinguished with

the following characteristics: Metatrochanter with low tu-

bercle to triangular process ventro-medially (Fig. 7).

Metafemur strongly incrassate (thickened) with long pile

ventrally (Fig. 7), apico-posteriorly with pilose lamina of

different size, with long row of black spinae along pos-

tero-ventral margin or with additionally several larger

spinae ventro-medially (Figs 7, 8). Metatibia ventro-basal-

ly with spinose carina along medial surface and a well-

developed lamina posteriorly (Fig. 9B); in E. gibbosus sp.

nov. carina weakly developed and lamina only present at

extreme apex and weakly developed. Genitalia with elon-

gate surstylus with hook shaped apex (Fig. 11).

Eumerus barbarus (Coquebert, 1804)

Figs 1A, 1B, 2A, 2B, 3A, 3B, 4A, 4B, 6A, 6B, 7A, 8A,

9A, 1OA & LIA

Syrphus barbarus Coquebert, 1804: 117. Type locality:

Barbaria Mus. Dom. Desfontaines [Neotype 0, ZMHB,

here designated].

Eumerus australis Meigen, 1838: 110. Type locality:

Spain, Andalusia [Holotype 9, MNHN].

Eumerus iris Loew, 1848: 118. Type locality: Italy, Sici-

ly, Syracuse [Lectotype 0, ZMHB, here designated].

Eumerus truquii Rondani, 1857: 95. Type locality: Italy,

Piemont [Holotype 2, MZUF].

Diagnosis. Metafemur (Figs 7A, 8A, 9A) strongly incras-

sate, with broad flat non pilose area ventrally with pos-

tero-ventral row of spinae covering nearly the entire length

of the metafemur; metafemur apico-posteriorly with low

lamina covered with dense white pile; metatibia (Figs 7A,

8A) ventro-basally along medial surface with broad cari-

na, posteriorly with very broad lamina; sternum IV (Fig.

10A) squarish, in lateral view with characteristic postero-

medial rounded extensions, in ventral view with laterally

projecting rounded postero-lateral corner and deep and

wide incision medially. Genitalia (Fig. 11A): surstylus

with very narrow medial lobe and apical lobe elongate,

gently curved margins with sharp, weakly bent apex; hy-

pandrium with rather broad apical part and with weakly

rounded and smooth ventral margin.

Redescription. MALE (Figs 1A, 2A). Body length:

6.8-8.7 mm, wing length: 4.7-6.3 mm. Head (Figs 3A,

4A). Face nearly parallel sided. Frons light-grey pollinose,

with scattered light yellow pile. Ratio width of head : width

of face 2.9-3.1 : 1. Eye contiguity relatively short, ratio

Bonn zoological Bulletin 66 (1): 145-165

frons : eye contiguity : vertical triangle is as 1.4-1.7 :

1.2-1.4 : 1. Vertical triangle grey-yellow pollinose, nar-

row grey-yellow pollinose along eye margin and two short

oval maculae posterior of ocelli, pile golden-yellow, on

ocellar triangle ranging from yellow with some black pile

intermixed to entirely black pilose. Occiput predominant-

ly grey pollinose, entirely white pilose. Ocelli isosceles

with broad posterior base, length : width 1.3-1.4 : 1; pos-

terior ocelli relatively far from eye margin, ratio width of

frons : width of ocellar triangle 1.3-1.5 : 1; ratio a:b:c as

1.4-1.6 : 1.2-1.4 : 1; width of head : width of vertex is

as 3.7-4.4 : 1. Antennae black to orange-yellow, scape

black, pedicel from orange to black, basoflagellomere

from entirely orange to yellow with apical half brown to

dark-brown. Scape and pedicel with long yellow to black

setae and light pile. Basoflagellomere (Fig. 6A) short rec-

tangular, ratio length : width as 1.2—1.3 : 1 with anterior

margin skewed ventrally. Arista with broad base, total

length 2-3 times longer than basoflagellomere, basal 1/5

yellow to entirely black.

Thorax. Black bronze shiny, pollinose along lateral mar-

gin and posterior and anterior of notopleura, two medio-

lateral pollinose vittae reaching to posterior 2/3—3/4 of scu-

tum, these pollinose vittae with broad triangular anterior

part. Pile light yellow. Pleurae bronze black, pilose on

proepimeron, postero-dorsal margin of anterior anepister-

num, entire posterior anepisternum, anterior half of

anepimeron and a dorsal and ventral pile patch on katepis-

ternum. Grey pollinose on pilose parts, others bronze black

sub shiny. Metasternum pilose. Scutellum rectangular with

narrow marginal rim, shiny black with bronze sheen, gold-

en yellow pilose. Legs. Black and brown-yellow to orange

coloured. Pro- and mesotarsus orange-yellow, connecting

parts of tarsomeres black; pro- and mesotibia basal 1/2—2/3

orange-yellow and with apical black ring sometimes with

brow-yellow anterior part; pro- and meso femur apical

1/10 brown-yellow. Pile all yellow with black spinae on

ventral side of mesotarsus. Metaleg with coxa black and

long white pile; trochanter (Fig. 7A) brown-yellow to

black, short scattered mixed yellow and black pile, api-

coventrally with short rounded tubercle; femur (Figs 7A,

8A, 9A) black, strongly enlarged, ratio length : width is

2.4-2.7: 1, with broad flat non pilose area ventrally with

antero-ventral row of 5—8 black spinae apically and pos-

tero-ventral row of 14—22 black spinae covering nearly the

entire length of the metafemur; apico-posteriorly with

densely white pilose low lamina; pile light-yellow, near-

ly evenly long on dorsal surface and short on ventral sur-

face, except long on apicoventral part. Metatibia (Figs 7A,

8A) orange-yellow on basal 1/2 and apical 1/10; only

slightly curved, ventro-basally along medial surface with

broad carina, posteriorly with very broad lamina; tarsus

orange to dark-brown. Wing. Entirely microtrichose to

partly bare medially on alula and basally on cell bm. Ab-

domen. Shiny bronze black, white pilose, antero-medial

©ZFMK

148 Jeroen van Steenis et al.

Fig. 1. Adult, dorsal view. A. Eumerus barbarus (Neotype, &). B. Eumerus barbarus (W. Batna, Algeria, 9). C. Eumerus trun-

catus (Holotype, 3). D. Eumerus gibbosus sp. n. (Albacete, Spain, Paratype, 7). E. Eumerus gibbosus sp. n. (Albacete, Spain, Q).

F. Eumerus schmideggeri sp. n. (Gafsa, Tunisia, Paratype, “). G. Eumerus schmideggeri sp. n. (Sidi R’ bat, Morocco, Paratype, @).

H. Eumerus sulcitibius (Albacete, Spain, “). I. Eumerus sulcitibius (Albacete, Spain, 9). Scale = 1 mm.

part of terga II-IV with triangular area black pilose; ter- _— lateral view with characteristic postero-medial rounded ex-

ga I-IV with light-grey pollinose and slightly oblique — tensions, in ventral view with laterally projecting round-

maculae, medially slightly wider; sternum IV squarish,in ed postero-lateral corner and deep and wide incision me-

Bonn zoological Bulletin 66 (1): 145-165 ©ZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 149

dially; sternum VI predominantly light yellow to entirely

black pilose; sternum VII from entirely black to predom-

inantly light-yellow pilose. Genitalia (Fig. 11A). Sursty-

lus with squarish basal part with scattered setae ventro-

medially; medial part narrowly elongate with small patch

of setulae at border with apical part of surstylus; apical

lobe elongate and gently curved with sharp, weakly bent

apex; hypandrium with rather broad apical part and with

weakly rounded and smooth ventral margin; hamus elon-

gate to club-shaped with broad apex.

FEMALE (Figs 1B, 2B). Body length: 7.3-10.1 mm,

wing length: 5.3-7.1 mm. Similar to male except for the

usual sexual dimorphism. Head (Figs 3B, 4B). Ratio width

of head : width of face 2.6-—2.9 : 1. Ocelli isosceles with

broad posterior base, length : width 1.2—1.4: 1; ratio width

width of ocellar triangle 1.8—2.0 : 1; width of

head : width of vertex is as 3.0-3.3 : 1. Basoflagellomere

of frons:

(Fig. 6B) large round, orange to light-brown coloured, ra-

tio length : width as 0.93-1.1 : 1. Legs. Metafemur black,

strongly incrassate, ratio length : width is 2.7-2.9: 1.

Type material examined. Neotype & Syrphus barbarus:

Tunisia: “N-Tunesien 27.6.1994 / Ghar el Mehl Strand /

35 km SO Bizerte / leg. Hauser , Tu-Gha”, “Eumerus bar-

barus | (Coquebert, 1804) d'/ det. M. Hauser 1996”, “Neo-

type J / Syrphus barbarus / Coquebert, 1804 / des J. van

Steenis, 2016” [red label] (ZMHB).

Holotype 9 Eumerus australis: Spain: “—-“ [round sil-

ver label], “Hispania / Eumerus / australis /M.” (MNHN).

Lectotype 0 Eumerus iris: Italy: “Sicilia / 1/5 Zeller”,

“coll / H. Loew”, “Typus”[red label], “Zool. Mus. /

Berlin”, “Lectotype “/ Eumerus iris / Loew, 1848 / de-

sign. J. van Steenis, 2016” [red label], (ZMHB). This spec-

imen is here designated as lectotype to fix and ensure the

universal and consistent interpretation of the name. Para-

lectotypes: 1 ¥, 2 99 Eumerus iris, with same data as the

lectotype (ZMHB). These specimens are here designated

as paralectotypes.

Holotype 2 Eumerus truquii: Italy: “ 141‘, [white oval

label with red text], “Museo La Specola / coll. Rondani /

HOLOTYPUS” [red label], “Eumerus barbarus / det. Vu-

ji€ 2014”, “Holotype Q / Eumerus truquii / Rondani, 1857

/ det J. van Steenis, 2016” [red label] (MZUF).

Additional material examined. Algeria: “Algeria,

4.V.1893, A.E. Eaton”, “Eumerus sp, nr. barbarus N.P. Wy-

att det”, 1 d' (NHM); “Algeria, Constantine, 2.VI.1895,

B.M.1896-137, A.E. Eaton” 1 2 (NHM)”; “Algeria, Hip-

pone, 8.1V.1896”, 1 3 (NHM); “Richter / Algers”, “Eu-

merus ruficornis / det v/d Wulp”, “Eumerus barbarus / det.

v. Helsdingen 1964”, 1 “ (NBC); “20 km N de Maghnia

/ Bab Taza/9 IV 1983 st 14”, Algerie /Tlemcen/R. Leys

& / P. v.d. Hurk”, 1 3 (ISA), 1 J (NBC), 1 2 (MZW);

same labels except “4 km E / st 33 / 24 IV 1983”, 1 9

(JSA); same labels except “Tlemcen / camp. municipal /

Bonn zoological Bulletin 66 (1): 145-165

20 IV 1983 st 4.”, 1 J (MZW); same labels except “Man-

sourah / st 22/14 IV 1983”, 1 J (NBC); “Algerije / W.

Tlemcen /Ain Pezza / 11-4-1981 / leg. R. Hensen”, 1 &

(NBC); “Algerije / W. Batna / Timgad / 17-4-1981 / leg.

R. Hensen”, 1 9 (NBC); “Mascara Algeria / Dr. J. Be-

quart”, “Eumerus / barbarus | Coq. 2”, 1 2° (MNHN);

“Palikao / Mascara Algeria / Dr. J. Bequart”, “Eumerus /

iris / Lw.”, 1 0 (MNHN); France: “Banyuls s Mer / France

/ P.O. / 21-iv-1951 v Doesburg”, 1 J (NBC); same label

except 8-iv, 9-iv and 20-iv, 5 JS, 1 9 (NBC); “France

(Corse) / Nonad / 10-v-1963 / HJP Lambeck”, 3 0’, 1 9

(NBC); “France Corse /J.A.W. Lucas”, “Vemaco /600 m

/ 18-7-1967”, 1 0 (NBC); “France Pyr. Or. / St. Cyprien

/ Plage / 3-VII-1956 / exc Zool Museum”, 1 dS (NBC);

“Sardegna / Teuipio P 24-9-1949 / Castellani leg”, 1 J, 1

9 (NBC); Italy: “Sicily, above Trapani, 16.1V.1965, 200m,

K.M. Guichard, B.M. 1965-273” 1 3 (NHM)”; “Sicily,

Selinunte, 13.1V.1965, S.L., K.M. Guichard, B.M. 1965-

273” 2 SS (NHM)”; “Italy / Sardinia Lode / R. Mannu /

April 1989 / leg M. Hauser:, 1 9 (CSCA); “Ragusa_/ Italia

18-5-1954 / FF. Tippmann”, “Eumerus australis / det v

Doesburg”, 1 3’; same label as previous, except “Eumerus

barbarus / det v Doesburg”, 8 3d, 1 2 (NBC), 1 &

(NHM); “Italia / Sicilie / HJP Lambeck”, “Adreno / Pas-

so Zingelo / 650 m/24-v-1966”,6 03, 1 2 (NBC); “Italia

/ Sicilia”, “Mondello / 10-4-1979 / J.A.W. Lucas”, 1 3

(NBC); same label as previous, except 11-4, 13-4 and 15-

4,8 SS, 2 29 (NBC): “Italia / Sicilia / J.A.W. Lucas”,

“Taormina / 23-4-1976”,3 dd (NBC); “I-Sardinien Tem-

pio 1200m / Mt. Limbara 20.7.96 / leg.: Dr. Ch.L. Neu-

mann”, 1 9 (CSCA); “I-Sardinien / april 1989 / Florinas,

417m /M. Hauser leg”, “Eumerus barbarus Coqu. / det

Claussen 1994”, “Eumerus barbarus / (Coquebert, 1804)

& / det.: M. Hauser 1996” 1 3, (CSCA); Morocco:

“Marokko / 11 km NW Tallouine / 15.III.97 30°34’N

8°00’ W /leg M. Hauser’, 1 dS (JSA); “Marokko Antiat-

las / S Ait-Baha / 12.III.97 30°06’N 9°02’ W / leg M.

Hauser”, 1 3 (CSCA); “Marokko Antatlas Amm- / elen-

tal 10km NE Tafraoute / 14.111.[19]97 29°48 ’N 8°53’°W

/leg M. Hauser”, 2 '% (CSCA); Portugal: “Portugal / 24-

4-1985”, 2 99 (NBC); Spain: “Mallorca I Andraixt,

26.Mar.1977 G.E.Bohart”, 2 0% (LAMC); “Espana / Mal-

lorca / 1-24-vi-1954 / leg Klokke Moll”, “Eumerus aus-

tralis / det v Doesburg”, 1 fe (NBC); “W. Malta / Wardi-

jn Ridge / Rooan Baths / 20-4-1986/ J.A.W. Lucas”, 7 3S,

4 99 (NBC); “Calvia / Caphella / 1-5.V.1979”, “Islas

Baleares / Mallorca / W.H. & A.FE. / Gravestein”, 1 &

(NBC); “Palma”, 1 S| (MNCN); “Alicante /Dusmet”, 1 dS

(MNCN); “Vacia Madrid / Dusmet”, “MNCN / Madrid”,

1 J (MNCN); “Spain, Almeria Province / 800 ft. 3 km W

Benhadux / 11-IV-1999 ME Irwin / 36°55.1°N

02°28.64’ W /hand netted”, 1 0 (CSCA); “El Pardo / VI-

1908 / Arias”, “M.N.C.N. / Madrid”, “Eumerus barbarus

/LlCoguebert,. 1304), / Des. A *Rucarte, 2005-4,

“MNCN_Ent/ 142918”, 1 0 (MNCN); “Espafia, Ciudad

©OZFMK

150 Jeroen van Steenis et al.

Fig. 2. Adult, lateral view. A. Eumerus barbarus (Neotype, &). B. Eumerus barbarus (W. Batna, Algeria, 2). C. Eumerus gibbo-

sus Sp.n. (Albacete, Spain, Paratype, “). D. Eumerus gibbosus sp. n. (Albacete, Spain, 9). E. Eumerus schmideggeri sp. n. (Gafsa,

Tunisia, Paratype, 3). F. Eumerus truncatus (Holotype, &). G. Eumerus sulcitibius (Albacete, Spain, “). H. Eumerus sulcitibius

(Albacete, Spain, Oy. Scale = 1 mm.

Real / P.N. de Cabafieros, maR2 / 1/24-VIII-2004 / leg.: (ESCAY ALS (ISA); “Tunisia S M’Saken/5 km N Sidi

A. Ricarte 5160”, “SYRPHIDAE / Eumerus barbarus / Bou Goubrine / 21.V.1999 ~ 1000 m amsl / 35°36.29’N

(Coquebert, 1804) 9 / Det.: A. Ricarte, 2006 (GBIF)”, 1 10°36.04’E/leg. O. & M. Niehuis”, 2 Jd’, 1 9 (CSCA);

@ (CEUA); same label except “maR1 and 5189”, 1 4 “W Tunesia / 5 km N El Kef / Tal m. Eryngium /

(CEUA); Tunisia: same labels as Neotype, 1 #0, 1 Q 22.VI.1994 leg Schmid-Egger Tu-Kef”, 1 9 (CSCA);

Bonn zoological Bulletin 66 (1): 145-165 ©ZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 151

H eS |

Fig. 3. Head, dorsal view. A. Eumerus barbarus (Neotype, “). B. Eumerus barbarus (W. Batna, Algeria, 2). C. Eumerus trunca-

tus (Holotype, “). D. Eumerus gibbosus sp. n. (Albacete, Spain, Paratype, ). E. Eumerus gibbosus sp. n. (Albacete, Spain, Paratype,

Q). K. Eumerus schmideggeri sp. n. (Gafsa, Tunisia, Paratype, “). G. Eumerus schmideggeri sp. n. (Sidi R’ bat, Morocco, Paratype,

Q). H. Eumerus sulcitibius (Albacete, Spain, &). I. Eumerus sulcitibius (Albacete, Spain, 2). Scale = 1 mm.

“Tunesia / Tabarka / Khatairia 15.V.1993”, 1 ’ (CSCA); country: “Beicos / 1/8”, 1 9 (MNHN); “Sa....0 / Sa..d /

“5 km E of / Tabarka 50 m / 26-IV-1980/ flower pasture”, Krausse”, “Eumerus / barbarus / Coq. 2”, [handwritten,

“Tunesia / E. van Nieukerken, /G. Bryan & / P. Ooster- partly unreadable], 2 99 (MNHN).

broek”, 1 9 (NBC); 3 additional specimens without known

Bonn zoological Bulletin 66 (1): 145-165 ©Z7FMK

52 Jeroen van Steenis et al.

Fig. 4. Head, frontal view. A. Eumerus barbarus (Neotype, m#). B. Eumerus barbarus (W. Batna, Algeria, 9). C. Eumerus trun-

catus (Holotype, 3). D. Eumerus gibbosus sp. n. (Albacete, Spain, Paratype, 7). E. Eumerus gibbosus sp. n. (Albacete, Spain, Q).

F. Eumerus schmideggeri sp. n. (Saida, Algeria, Paratype, 7). G. Eumerus schmideggeri sp. n. (Sidi R’ bat, Morocco, Paratype, Q).

H. Eumerus sulcitibius (Albacete, Spain, “). I. Eumerus sulcitibius (Albacete, Spain, 9). Scale = 1 mm.

Remarks. This is the most widespread species and, in

some places, it seems to be sympatric with the other

species. The type of Syrphus barbarus Coquebert, 1804

was deposited in the Desfontaines collection “Mus. Dom.

Desfontaines” and later placed in the MNHN, but this part

of the MNHN collection is presumed to be lost (Zimsen

1964). In the material studied here several specimens of

Bonn zoological Bulletin 66 (1): 145-165

E. barbarus have been named either E. iris or E. australis.

In order to fix the usage of the name of Eumerus barbarus,

we designated a neotype. The locus typicus is given as

“Barbaria”’, and according to Zimsen (1964) the specimens

collected by Desfontaines originate from Algeria and

Tunisia so we picked a specimen from Tunisia which

agreed with the original description.

©OZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 153

Eumerus gibbosus sp. n.

Figs 1D, 1E, 2C, 2D, 3D, 3E, 4D, 4E, 6D, 6E, 7B, 8B, 10B

& 11B

Diagnosis. Metafemur medio-ventrally with area of

light- to dark-brown pile; apico-posterior lamina of

metafemur large and densely dark brown pilose (Figs 7B,

8B); metatibia ventro-basally along medial surface with

very low carina, posteriorly at extreme base with very low

lamina; sternum IV (Fig. 10B) squarish, with normal pos-

tero-lateral corners and wide oval incision medially. Gen-

italia (Fig. 11B): surstylus with wide triangular medial part

and elongate and tapering apical part with slightly round-

ed and sharply bent apex; hypandrium with broad apical

part and several projections along ventral margin.

Description. MALE (Figs 1D, 2C). Body length: 6.1-7.5

mm, wing length: 4.0-5.2 mm. Head (Figs 3D, 4D). Face

nearly parallel sided. Frons light-grey pollinose, with scat-

tered light yellow pile. Ratio width of head : width of face

3.0-3.1 : 1. Eye contiguity relatively short, ratio frons :

eye contiguity: vertical triangle is as 1.4-1.5 : 1.1-1.2:

1. Vertical triangle grey-yellow pollinose, narrow grey-yel-

low pollinose along eye margin and two short oval mac-

ulae posterior of ocelli, pile golden-yellow, on ocellar tri-

angle ranging from yellow with some black pile intermixed

to entirely black pilose. Occiput predominantly grey polli-

nose, entirely white pilose. Ocelli isosceles with broad pos-

terior base, length : width 1.2—1.4 : 1; posterior ocelli rel-

atively close to eye margin, ratio width of frons : width

of ocellar triangle 1.2-1.3 : 1; ratio a:b:c as 1.3-1.4 :

1.2-1.3 : 1; width of head : width of vertex is as 4.1—4.5

: 1. Antennae black to orange-yellow, scape black, pedi-

cel from orange to black, basoflagellomere from entirely

orange to yellow with apical half brown to dark-brown.

Scape and pedicel with long yellow to black setae and light

pile. Basoflagellomere (Fig. 6D) short rectangular, ratio

length : width as 1.1-1.2 : 1 with anterior margin skewed

ventrally. Arista with broad base, total length 2—3 times

longer than basoflagellomere, basal 1/5 yellow to entire-

ly black. Thorax. Black bronze shiny, pollinose along lat-

eral margin and posterior and anterior of notopleura, two

mediolateral pollinose vittae reaching to posterior 2/3—3/4

of scutum, these pollinose vittae with broad triangular an-

terior part. Pile light yellow. Pleurae bronze black, pilose

on proepimeron, postero-dorsal margin of anterior anepis-

ternum, entire posterior anepisternum, anterior half of

anepimeron and a dorsal and ventral pile patch on katepis-

ternum. Grey pollinose on pilose parts, others bronze black

sub-shiny. Metasternum pilose. Scutellum rectangular with

narrow marginal rim, shiny black with bronze sheen, gold-

en yellow pilose. Legs. Black and brown-yellow to orange

coloured. Pro- and mesotarsus orange-yellow, connecting

parts of tarsomeres black; pro- and mesotibia basal 1/2—2/3

orange-yellow and with apical black ring sometimes with

Bonn zoological Bulletin 66 (1): 145-165

Fig. 5.

to de las Crucetillas. B. Cafiada del Provencio.

Habitat of Eumerus gibbosus sp.n., Albacete. A. Puer-

brown-yellow anterior part; pro- and mesofemur apical

1/10 brown-yellow. Pile all yellow with black setulae on

ventral side of mesotarsus. Metaleg with coxa black and

long white pile; trochanter brown-yellow to black, short

scattered mixed yellow and black pile, apicoventrally with

short rounded tubercle (Fig. 7B); femur (Figs 7B, 8B)

black, slightly enlarged, ratio length : width is 3.0-3.5 :

1, with normally curved ventral surface, with ventral sur-

face long white pilose, medio-ventrally with patch of light-

to dark brown pile, antero-ventrally with a row of 7-11

black spinae apically and postero-ventrally with a row of

9-14 black spinae along apical 1/2, apico-posterior lam-

ina large and densely dark brown pilose; metatibia orange-

yellow on basal 1/2 and apical 1/10; only slightly curved,

apically with antero-medial row of black dense spicules

and ventro-basally along medial surface with very weak

carina, posteriorly at extreme base with very low lamina;

tarsus orange to dark-brown. Wing. Microtrichose, except

bare on very small medial part on alula and extreme base

of cell bm. Abdomen. Shiny bronze black, white pilose,

antero-medial part of terga II-IV with triangular area black

pilose; terga II-IV with light-grey pollinose and slightly

oblique maculae, medially slightly wider; sternum IV (Fig.

10B) squarish, with normal postero-lateral corners and

wide oval incision medially; sternum VI predominantly

light yellow to entirely black pilose; sternum VII from

mixed black and light-yellow to predominantly light-yel-

low pilose. Genitalia (Fig. 11B). Surstylus with large

squarish basal part with very scattered setae ventro-me-

dially; medial part widely triangular with scattered setu-

lae; apical part elongate with slightly rounded and

sharply bent apex. Hypandrium with broad apical part with

two squarish projections along ventral margin and with

dense patch of small setulae ventro-apically; hamus short

and with large round apex.

©ZFMK

154 Jeroen van Steenis et al.

Fig. 6. Antennae, lateral view. A. Eumerus barbarus (Neotype, &). B. Eumerus barbarus (Tlemcen, Algeria, 2). C. Eumerus trun-

catus (Holotype, &). D. Eumerus gibbosus sp. n. (Albacete, Spain, Paratype, m#). E. Eumerus gibbosus sp. n. (Albacete, Spain,

Q). F. Eumerus schmideggeri sp. n. (Gafsa, Tunisia, Paratype, m#). G. Eumerus sulcitibius (Albacete, Spain, 7). H. Eumerus sul-

citibius (Albacete, Spain, 2). Scale = 0.5 mm.

FEMALE (Figs 1E, 2D). Body length: 7.8-8.2 mm,

wing length: 5.8-6.1 mm. Similar to male except for the

usual sexual dimorphism. Head (Figs 3E, 4E). Ratio width

of head : width of face 2.7—2.8 : 1. Ocelli isosceles with

broad posterior base, length : width 1.3—1.5 : 1; ratio width

of frons : width of ocellar triangle 1.5-1.7 : 1; width of

head : width of vertex is as 3.8-—4.0: 1. Basoflagellomere

(Fig. 6E) round to slightly oval, dark orange-brown

coloured with darker antero-dorsal margin, ratio length :

width as 0.97-1.1 : 1. Legs. Metafemur black, slightly in-

crassate, ratio length : width is 2.6—2.9 : 1. Abdomen. Ter-

gum IV without pollinose maculae.

Etymology. The characteristics which separate this

species from closely related species are the two humps on

the ventral margin of the hypandrium; “gibbosus”, the Lat-

in word for humped.

Bonn zoological Bulletin 66 (1): 145-165

Biology. The specimens from Spain were collected in

a low mountainous area with open Pine (Pinus halepen-

sis Miller and PR nigra Arnold) dominated forest with un-

dergrowth of Mediterranean maquis with diverse yellow

and white flowering Apiacaea (Fig. 5).

Type material. HOLOTYPE &: Spain: “Spain Albaceta

/ [leg] J. van Steenis & M.P. van Zuijen”, “Sierra de Al-

caraz, 1400 m [a.s.l.] / Puerto de las Crucetillas / [road]

C 415 km 190, open pine forest / 38°31’N 2°26’ W/on

Thapsia villosa / 21-V1-2003” (NBC). Paratypes: Spain:

with same data as Holotype, 1 #1 2 (CSCA),1 4,1 2

(JSA); with same data except “Sierra de Alcaraz, 1000 m

[a.s.l.] / river valley, meadow marsh / Cafiada del Proven-

cio / 38°31’N 2°20’W / on Thapsia villosa | 22-V1-

2003”, 2 JS (ISA); “Spain (E) Albacete, Riopar / Puerto

©OZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 155

Fig. 7. Right metaleg (without tarsi) of #, anterior view. A. Eumerus barbarus (Sicily, Italy). B. Eumerus gibbosus sp. n. (A1-

bacete, Spain, Paratype). C. Eumerus schmideggeri sp. n. (Saida, Algeria, Paratype). D. Eumerus sulcitibius (Albacete, Spain). Scale

= 1 mm. a = apico-posterior lamina of metafemur, b = posterior lamina of metatibia, c = ventro-medial excavation of metatibia, d

= process of metatrochanter.

de las Crucetillas road / C415 km 190, 1400 m [a.s.1.] /

38°31’ NB 2°26’ WL/ [leg] MP van Zuijen, J van Stee-

nis / 21-6-2003 / dry pine forest, [on] yellow Apiaceae”,

1 J (MZW); “Spain(E) Albacete, Riopar, Cafiada / de

Provencio, 1000 m [a.s.1.] / 38°31 “NB 2°20’ WL / [leg]

MP van Zuijen, J van Steenis / 22-6-2003 / stream, open

forest [on] yellow Apiaceae”, 1 J (ISA), 2 3S (MZW);

Portugal: “PORTUGAL mal.trap / Mertola, west of town

/ dry rainwater brooklet / UTM: 29S 6171-4166 / 21-

23.v.2004 [leg] A v. Eck”, 1 ’ (AET).

Eumerus schmideggeri sp. n.

Figs 1F, 1G, 2E 3F, 3G, 4F, 4G, 6F, 7C, 8C, 10C & 11C

Diagnosis. Pollinosity on ocellar triangle with two large

oval maculae along eye margin (Fig. 3F); metafemur (Figs

7C, 8C) apico-posteriorly with hardly visible and scattered

Bonn zoological Bulletin 66 (1): 145-165

white pilose lamina; metatibia ventro-basally along me-

dial surface with rather broad carina, posteriorly with

rather broad lamina; pollinose maculae on terga very wide

(Figs 1F, G); sternum IV (Fig. 10C) trapezoid, posterior

margin concave with triangular posterolateral corners, nar-

row but deep medial incision. Genitalia (Fig. 11C): sursty-

lus with small squarish medial part and straight apical part

with rounded and slightly bent apex; hypandrium with nar-

row and straight apical part.

Description. MALE (Figs 1F, 2E). Body length: 8.4-9.3

mm, wing length: 5.5-6.3 mm. Head. Face nearly paral-

lel sided. Frons light-grey pollinose, with scattered light

yellow pile. Ratio width of head : width of face 2.8-3.0:

1. Eye contiguity relatively short, ratio frons : eye conti-

guity : vertical triangle is as 1.4-1.6 : 1.2-1.3 : 1. Verti-

cal triangle grey-yellow pollinose, broadly grey-yellow

pollinose along eye margin and two broad squarish mac-

©ZFMK

156 Jeroen van Steenis et al.

Fig. 8. Detail of left metaleg in &, posterior view. A. Eumerus barbarus (Corse, France). B. Eumerus gibbosus sp. n. (Holotype).

C. Eumerus schmideggeri sp. n. (Saida, Algeria, Paratype). D. Eumerus sulcitibius (Albacete, Spain). Scale = 1 mm.

ulae posterior of ocelli, pile golden-yellow, on ocellar tri-

angle entirely black pilose. Occiput predominantly grey

pollinose, entirely white pilose. Ocelli isosceles with broad

posterior base, length : width 1.2—1.3 : 1; posterior ocel-

li relatively far from eye margin, ratio width of frons :

width of ocellar triangle 1.4—1.5 : 1; ratio a:b:c as 1.3-1.4

: 1.2-1.3 : 1; width of head : width of vertex is as 4.3-—4.5

: 1. Antennae black to orange-yellow, scape black, pedi-

cel from orange to black, basoflagellomere from entirely

orange to yellow with apical half brown to dark-brown.

Scape and pedicel with long yellow to black setae and light

pile. Basoflagellomere (Fig. 6F) trapezoid, ratio length :

width as 1.0-1.1 : 1 with anterior margin skewed ventral-

ly. Arista with broad base, total length 2—3 times longer

than basoflagellomere, basal 1/5 yellow to entirely black.

Thorax. Black bronze shiny, pollinose along lateral mar-

gin and posterior and anterior of notopleura, two medio-

lateral pollinose vittae reaching to posterior 2/3—3/4 of scu-

tum, these pollinose vittae with broad triangular anterior

Bonn zoological Bulletin 66 (1): 145-165

part. Pile light yellow. Pleurae bronze black, pilose on

proepimeron, postero-dorsal margin of anterior anepister-

num, entire posterior anepisternum, anterior half of

anepimeron and a dorsal and ventral pile patch on katepis-

ternum. Grey pollinose on pilose parts, others bronze black

sub shiny. Metasternum pilose. Scutellum rectangular with

narrow marginal rim, shiny black with bronze sheen, gold-

en yellow pilose. Legs. Black and brown-yellow to orange

coloured. Pro- and mesotarsus orange-yellow, connecting

parts of tarsomeres black; pro- and meso tibia basal

1/2—2/3 orange-yellow and with apical black ring, some-

times with brow-yellow anterior part; apical 1/10 of pro-

and meso femur brown-yellow. Pile all yellow with black

setulae on ventral side of mesotarsus. Metaleg with coxa

black and long white pile; trochanter brown-yellow to

black, with short scattered mixed yellow and black pile,

apicoventrally with short rounded tubercle (Fig. 7C); fe-

mur (Figs 7C, 8C) black, slightly enlarged, ratio length :

width is 2.8—3.1 : 1, with normally curved ventral surface,

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Review of the Eumerus barbarus species group from the western Mediterranean Basin 157

Fig. 9. Details of right metaleg of #. A, B SEM pictures; C Stack photo. A. Metafemur ventral view, Eumerus barbarus (Corse,

France). B. Tibia ventral view, Eumerus sulcitibius (Albacete, Spain). C. Femur and tibia, anterior view, Eumerus sulcitibius (Sai-

da, Algeria). A = posterior lamina of metatibia, B = medial carina of metatibia, C = ventro-medial excavation of metatibia.

covered with long white pile, antero-ventrally with a row

of 8-10 black spinae apically and postero-ventrally with

a row of 12-16 black spinae on apical 2/3, apico-posteri-

or lamina large and densely dark brown pilose; metatib-

ia orange-yellow on basal 1/2 and apical 1/10; only slight-

ly curved, apically with antero-medial row of black dense

spicules and posterior ventro-basally along medial surface

with narrow carina; tarsus orange to dark-brown. Wing.

Microtrichose except bare on medial part of alula and

basally on cell bm and br. Abdomen. Shiny bronze black,

white pilose, antero-medial part of terga II-IV with trian-

gular area black to white pilose; terga H-IV with broad

light-grey pollinose and slightly oblique maculae, medi-

Bonn zoological Bulletin 66 (1): 145-165

ally slightly wider; sternum IV (Fig. 10C) trapezoid, pos-

terior margin concave with triangular posterolateral cor-

ners, narrow but deep medial incision; sternum VI pre-

dominantly light yellow pilose; sternum VII predominant-

ly light-yellow pilose. Genitalia (Fig. 11C). Surstylus with

large rectangular basal part with large dorso-medial patch

of setae; medial part squarish with baso-dorsal patch of

setulae; apical part elongate with slightly rounded and

slightly bent apex. Hypandrium with narrow and smooth

apical part; hamus short and with small round apex.

FEMALE (Fig. 1G). Body length: 7.0-9.5 mm, wing

length: 4.8-6.2 mm. Similar to male except for the usual

sexual dimorphism. Head (Figs 3G, 4G). Ratio width of

©OZFMK

158 Jeroen van Steenis et al.

Fig. 10.

Sternum IV J, ventral view. A. Eumerus barbarus (Bizerte, Tunisia). B. Eumerus gibbosus sp. n. (Holotype). C. Eume-

rus schmideggeri sp. n. (Holotype). D. Eumerus sulcitibius (Capri, Italy).

head: width of face 2.7 : 1. Ocelli isosceles with broad pos-

terior base, width : length 1.3 : 1; ratio width of frons :

width of ocellar triangle 1.9 : 1; width of head: width of

vertex is as 3.5: 1. Basoflagellomere oval with ventral part

longer than dorsal, bright orange coloured with anterior

2/3 orange-brown, ratio length : width as 1.1 : 1. Legs.

Metafemur black, slightly incrassate, ratio length : width

is 2.9 : 1. Abdomen. Pollinose maculae oval-shaped, thick,

occupying about 1/3 of length of tergum.

Etymology. Named after Christian Schmid-Egger,

long-time friend, exceptional hymenopterist and avid in-

sect collector, including the Holotype of this species.

Type material. HOLOTYPE &: Morocco: “Morocco,

37 km S Agadir / Sidi R’bat, 30.084 N 9.664 W / 30m

NN leg. Schmid-Egger / 17.06.2014 mal7” (ZMHB).

Paratypes: Morocco: with same labels as holotype, 1 “

(JSA), 2 92 (CSCA); Tunisia: “Tunesien 21.6.2194 /

Bonn zoological Bulletin 66 (1): 145-165

Gafsa, Oasegarten / leg. Hauser, Tu-Gaf”, 1 3’ (CSCA);

Algeria: “Dayet el Kerch / st 5./5 IV 1983”, “Algerie /

Saida/R. Leys & /P. v.d. Hurk”, 1 3’) (NBC); same labels

except “5 km SE de Sfissifa / st. 8./6 IV 1983”,2 3S

(ISA, MZW).

Eumerus sulcitibius Rondani, 1868

Figs 1H, 11, 2G, 2H, 3H, 31, 4H, 41, 6G, 6H, 7D, 8D, 9B,

C,10D & 11D

Eumerus sulcitibius Rondani, 1868: 24. Type locality: Italy,

Parma [Syntype & MZUF, not studied].

Diagnosis. M etatrochanter ventrally with rather long and

triangular process (Fig. 7D); metafemur (Figs 7D, 8D, 9C)

ventro-medially with 2—3 large black spinae, clearly dif-

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Review of the Eumerus barbarus species group from the western Mediterranean Basin 159

Fig, 11.

Genitalia 3, lateral view. A. Eumerus barbarus (Bizerte, Tunisia). B. Eumerus gibbosus sp. n. (Holotype). C. Eumerus

schmideggeri sp. n. (Holotype). D. Eumerus sulcitibius (Capri, Italy). a = apex of surstylus, b = medial part of surstylus, c = basal

part of surstylus, d = hamus, e = ventral surface of hypandrium (here with two “humps’”).

ferentiated from apico-ventral spinae; metafemur with nar-

row, light-brown pilose, apico-posterior lamina; metatib-

ia ventro-medially with narrow but deep sulcus, metatib-

ia (Fig. 9B) ventro-basally along medial surface with

rather broad carina, ending medially in a deep excavation,

posteriorly with broad lamina; sternum IV (Fig. 10D) rec-

Bonn zoological Bulletin 66 (1): 145-165

tangular with large broadly rounded postero-lateral cor-

ner and broad but not deep medial incision. Genitalia (Fig.

11D): surstylus with rectangular medial part; apical part

elongate with slightly curved margins and with sharp, very

weakly bent apex; hypandrium with rather narrow and

slightly wavy apico-ventral margin.

©OZFMK

160 Jeroen van Steenis et al.

Fig. 12... Eumerus truncatus Rondani, &. A. sternum IV, ventral view (Ait-Baha, Morocco). B. Genitalia, lateral view (Ait-Baha,

Morocco). €= teaspoon shaped appendix.

Redescription. MALE (Figs 1H, 2H). Body length:

7.2-8.5 mm, wing length: 5.9-7.0 mm. Head (Figs 3H,

4H). Face slightly triangular widening dorsally. Frons

light-grey pollinose, with scattered light yellow pile. Ra-

tio width of head : width of face 3.1-3.4 : 1; eye contigu-

ity relatively long, ratio frons: eye contiguity : vertical tri-

angle 1.3-1.6: 1.4-1.6: 1. Vertical triangle grey-yellow

pollinose, narrow grey-yellow pollinose along eye margin

and two short oval maculae posterior of ocelli, pile gold-

en-yellow, on ocellar triangle ranging from yellow with

some black pile intermixed to entirely black pilose. Oc-

ciput predominantly grey pollinose, entirely white pilose.

width

1.1-1.3 : 1; posterior ocelli relatively close to eye margin,

Ocelli isosceles with broad base, ratio length :

ratio width of frons : width of ocellar triangle 1.2—1.4 :

1; ratio a:b:c as 1.2—1.4: 1.2—1.3 : 1; width of head: width

of vertex as 4.7-5.2 : 1. Antennae black to basally orange-

yellow, scape and pedicel black, basoflagellomere from

partly orange basally to entirely black. Scape and pedicel

with long yellow to black setae and light pile. Basoflagel-

lomere (Fig. 6G) rectangular, ratio length : width as

1.3-1.4 : 1. Thorax. Black bronze shiny, pollinose along

Bonn zoological Bulletin 66 (1): 145-165

lateral margin and posterior and anterior of notopleura, two

mediolateral pollinose vittae reaching to posterior 2/3—3/4

of scutum, these pollinose vittae with broad triangular an-

terior part. Pile light yellow, posterior intermixed with

longer black hairs. Pleurae bronze black, pilose on

proepimeron, postero-dorsal margin of anterior anepister-

num, entire posterior anepisternum, anterior half of

anepimeron and a dorsal and ventral pile patch on katepis-

ternum. Grey pollinose on pilose parts, others bronze black

sub shiny. Metasternum pilose. Scutellum rectangular with

narrow marginal rim, shiny black with bronze sheen, gold-

en yellow pilose. Legs. Black and brown-yellow to orange

coloured. Pro- and mesotarsus orange-yellow darkened

ventraly, connecting parts of tarsomeres black; pro- and

mesotibia basal 1/2—2/3 orange-yellow and with apical

black ring sometimes with brow-yellow anterior part; pro-

and mesofemur apical 1/10 brown-yellow. Pile all yellow

with black setulae on ventral side of mesotarsus. Metaleg

with coxa black and long white pile; trochanter brown-yel-

low to black, short scattered mixed yellow and black pile,

ventrally with rather long and triangular process (Figs 7D,

9C); metafemur (Figs 7D, 8D, 9C) slightly less incrassate

©OZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 161

compared with E. barbarus, length : width 2.9-3.3 : 1,

medio-ventrally with 2—3 long and strong spinae clearly

differentiated from apico-ventral rows of spinae, apico-

ventrally with long white pile, antero-ventrally with a row

of 8-11 black spinae apically and postero-ventrally with

a row of 8-10 black spinae on apical 1/3, metatibia strong-

ly curved, ventro-basally along medial surface with rather

broad carina, ending medially in a deep excavation, pos-

teriorly with broad lamina. Wing. Microtrichose, partly

bare on cell bm and cup. Abdomen. Shiny bronze black,

white pilose, antero-medial part of terga H-IV with trian-

gular area black pilose; terga I-IV with light-grey polli-

nose and slightly oblique maculae, medially slightly wider;

tergum IV predominantly black pilose, except lateral mar-

gin light yellow pilose; sternum IV (Fig. 10D) rectangu-

lar with large broadly rounded postero-lateral corner and

broad but not deep medial incision; sternum VI predom-

inantly light yellow to entirely black pilose; sternum VII

from entirely white to mixed white and black pilose. Gen-

italia (Fig. 11D). Surstylus with small rectangular basal

part with very scattered setae dorso-medially; medial part

rectangular with a small patch of setulae medially; apical

part elongate with slightly curved margins and with sharp

but weakly bent apex. Hypandrium with rather narrow api-

cal part with slightly wavy ventral margin; hamus long and

with rounded apex.

FEMALE (Figs 11, 21). Body length: 7.5—9.2 mm, wing

length: 5.9-6.8 mm. Similar to male except for the usual

sexual dimorphism. Head (Figs 31, 41). Ratio width of

head : width of face 3.0—3.3 : 1. Ocelli isosceles with broad

1; ratio width of

1; width of head

1. Basoflagellomere (Fig.

posterior base, length : width 1.2-1.4:

frons width of ocellar triangle 1.7-1.9 :

: width of vertex is as 3.8-4.1:

6H) round, black to basally dark-orange coloured, ratio

length : width as 0.95-1.0 : 1. Legs. Metafemur black,

slightly incrassate, ratio length : width is 2.8-3.3 : 1.

Material studied. France: “France (Corse) / Asco (600-

900) / 6-VH-1961 / HJP Lambeck”, 1 % (NBC); Italy:

“Italy / Capri, Mt Solaro /275-589m 25.V1.1992 / leg. P.W.

Lohr, -S.(CSCA)-. 1talia se Sicilia Ac WeyLucas™®,

“Taormina / 23-4-1976”, 4 JS, 1 2 (NBC); Portugal:

“Portugal 260 m / Porto de Mos Alvados / UTM

29S80519-4377 / 12.1V.2007 leg A. van Eck”, 1 ’' (CSCA);

Spain: “Espana / Avila / V.S. v.d. Goot / J.A.W. Lucas”,

“Sierra de Gredos / Navarredonda / de Gredos 1600 m /

9-7-1972”,4 3S, 5 OP (NBC); “Spain, Albaceta /J. van

Steenis & /M.P. van Zuijen”, “Sierra de Alcaraz, 1200 m

/ Puerto de las Crucetillas / C 415 km 186 slope near /

Populus plantation and small / stream 38°21 °N 2°24’ W

/ on Thapsia villosa 21-V1-2003”, 2 JH, 2 QP UISA);

“Spain (E) Albacete, Riopar / Puerto de las Crucetillas

road /C415 km 190, 1400 m [a.s.1.] / 38°31 ’NB 2°26’

WL/ [leg] MP van Zuijen, J van Steenis / 21-6-2003 / dry

pine forest, [on] yellow Apiaceae”, 1 0 (MZW); “Spain,

Bonn zoological Bulletin 66 (1): 145-165

Albaceta / J. van Steenis & /M.P. van Zuijen”, “Sierra de

Alcaraz / Batan del Puerto 1200 m / meadow in open Pine

forest / 38°34’N 2°21’ W / on Thapsia villosa / 21-V1-

2003”,1 3,2 PP (ISA), 1 9 (MZW).

Identification key to the species of the Eumerus bar-

barus group from western Mediterranean Basin

1. Males: eyes holoptic, genital capsule clearly developed

(Figs 1A, 2C, 2G, 3A) .. Aye tt Shela h geet eee

— Females: eyes dichoptic, setiiteaceule absent (Figs 1B,

2B, 3B).. Pere yaNe Ra nace Phe

2, Metatentar metic trae ith a 3 long black

spinae, clearly differentiated from the much shorter api-

co-posterior row of 8-10 spinae (Figs 7D, 8D, 9C), on

apico-posterior corner with densely white pilose, nar-

row lamina (Fig. 8D); metatibia ventro-basally along

medial surface with rather broad carina, ending medi-

ally in a deep excavation and posteriorly with broad lam-

ina (Figs 7D, 8D, 9B, 9C); metatrochanter ventrally with

rather long and triangular process (Figs 7D,

9). cee eee eee. Humerus sulcitibius Rondani, 1868

— Metafemur medio-ventrally without long black spinae,

only with a row of rather short and stout black spinae

(Figs 7A-C, 9A), with differently shaped and pilose api-

co-posterior lamina (Figs 8A—C); metatibia ventro-

basally along medial surface with weak to broad cari-

na, medial carina ending apically in at most a weak ex-

cavation (Figs 7A—C, 8A-—C); metatrochanter ventrally

with low tubercle or with short rounded process (Figs

33 Micra nuun steonly incrassate, mite dds -ventrally with a

broad and flattened scarcely pilose area, and apically

with 5-8 black spinae antero-ventrally and a row of

14-22 black spinae postero-ventrally along nearly the

entire length of the metafemur (Figs 7A, 9A); metafe-

mur at apico-posterior corner with densely white pilose,

narrow lamina (Fig. 8A); metatibia ventro-basally

along medial surface with broad carina, posteriorly with

very wide lamina (Figs 7A, 8A); sternum IV as in Fig.

10A, with two rounded projections postero-medially ..

. Eumerus barbarus (Coquebert, 1804)

_ Metafende slightly to clearly less incrassate, more even-

ly curved and almost entirely pilose, on apical part with

7-11 black spinae antero-ventrally and a row of 9-16

black spinae postero-ventrally along apical 1/2—2/3 (Figs

7B, 7C); metafemur at apico-posterior corner with

densely brown pilose and wide lamina (Fig. 8B) or with

scarcely pilose, hardly visible lamina (Fig. 8C); metat-

ibia ventro-basally along medial surface with very nar-

row to at most rather broad carina, posteriorly with at

most rather wide lamina (Figs 7B, 7C, 8B, 8C); sternum

IV as in Figs 10B-C, postero-medial surface smooth ..

©ZFMK

162 Jeroen van Steenis et al.

4. Metafemur medio-ventrally with an area with brown-

ish pile; metafemur at apico-posterior corner with dense-

ly brown pilose, wide lamina (Fig. 8B); metatibia ven-

tro-basally along medial surface with narrow carina,

posteriorly with very narrow lamina confined to the

most basal part (Figs 7B, 8B); terga with restricted polli-

nosity (Fig. 1D); sternum IV as in Fig. 10B . :

yeah aes eee .. Eumerus eibbOsis sper n.

— Weta eatite inredie ccemeaiie with only light-yellow pi-

losity; metafemur at apico-posterior corner with scarce-

ly pilose, hardly visible lamina (Fig. 8C); metatibia ven-

tro-basally along medial surface with rather broad ca-

rina, posteriorly with rather broad lamina (Figs 7C, 8C);

terga with extensive pollinosity (Fig. 1F); sternum IV

as in Fig. 10C............ Eumerus schmideggeri sp. n.

5. Oblique maculae on terga narrow (Figs 1B, 1E, 11) .. 6

— Oblique maculae on terga broad (Fig. 1G) .

coats eal ERP .. Eumerus schmidepoert sp. n.

6. Aeanoatat aosteis: setnitalty with a row of 4-10 even-

ly spaced spinae (similar to Figs 7A—C); basoflagellom-

ere large, more oval about as wide as long (Figs 6B, 6E)

— Metafemur medio-ventrally with 1—2 spinae, differen-

tiated from the other postero-ventral row of spinae (sim-

ilar to Figs 7D, 8D); basoflagellomere large and round,

almost wider than long (Fig. 6H) . abieee cg

seats . Eumerus sulcinibius Ronauai! 1868

de. REESE slightly incrassate (as in Fig. 3B); basofla-

gellomere large, oval (Fig. 6E); terga III and IV with

pollinose maculae reduced, more or less straight (Fig.

LE) oo. .e cece cee cee ee eee ees. Humerus gibbosus sp. n.

— Metafemur strongly incrassate (as in Fig. 3A); basofla-

gellomere large, rounded (Fig. 6B); pollinose maculae

on tergum III and IV large and strongly oblique (Fig.

1B)............... Eumerus barbarus (Coquebert, 1804)

Excluded species from the E. barbarus group

Eumerus truncatus Rondani, 1868 stat. n.

Figs 1C, 2F, 3C, 4C, 6C, 12A-B

Eumerus truncatus Rondani, 1868: 575. Type locality:

Italy, Sicily, Nebrodes [Holotype &', MZUF].

Diagnosis. Differing from the species of the E. barbarus

group and E. strigatus by the following characters: squar-

ish basoflagellomere (Fig. 6C) (in E. strigatus similar, in

E. barbarus group more oval with rounded apico-dorsal

corner), very slightly pollinose vertex (Figs 1C, 3C, 4C)

(in E. strigatus similar, in E. barbarus group with dense

pollinosity on apical corner and posterolateral from pos-

terior ocelli); ocellar triangle with narrow base (Fig. 3C)

(in E. strigatus slightly wider, in E. barbarus group with

very wide base); mesonotum with hardly visible pollinos-

Bonn zoological Bulletin 66 (1): 145-165

ity, at most very narrow vittae mediolateral reaching to

over 2/3 of scutum (Fig. 1C) (in E. strigatus and E. bar-

barus group with clear and wide medio-lateral pollinose

vittae); metafemur apico-posterior normal, without lam-

ina or pile fringe like in E. strigatus (in E. barbarus group

with pile fringe and sometimes even with carina); water

drop shaped to short straight pollinose maculae on tergum

IV (Fig. 1C) (in E. strigatus and E. sulcitibius with near-

ly straight oblique vittae and in the other three species of

the E. barbarus group with elongate lunulate maculae).

Redescription. MALE (Figs 1C, 2F). Body length:

6.6-6.9 mm, wing length: 4.5-5.1 mm. Head (Figs 3C,

4C). White pilose face and frons; mixed black and light-

brown pile on vertex, with very scattered pollinosity in an-

terior corner; eyes white pilose; ratio width of head : width

of face 3.1-3.3 :

frons: eye contiguity :

1.1-1.2 : 1; ocelli isosceles with narrow posterior base,

length : width 0.84-0.92 :

margin, ratio width of frons : width of ocellar triangle

1.5-1.7 : 1; ratio a:b:c as 1.2-1.3 : 1.2-1.3 : 1; width of

head : width of vertical triangle is as 4.0—-4.6: 1; basofla-

1. Eye contiguity relatively long, ratio

vertical triangle is as 1.1-1.3 :

1; posterior ocelli far from eye

gellomere (Fig. 6C) dark-brown to orange squarish, ratio

length : width as 1.1-1.3 : 1. Thorax. Shiny bronze with

white pile. Legs. Colour predominantly black, only some

tarsomeres and apex and bases of tibiae brown-yellow;

metafemur slightly incrassate with white pile ventrally, the

pile length about 2/3 of the width of femur and with nor-

mal vestiture of short black spinae apically, ratio length :

width is 3.2-3.4 : 1. Abdomen. Abdomen shiny bronze

with pairs of lunulate pollinose maculae on terga II and

III and water drop shaped pollinose maculae on tergum

IV, pile predominantly white; sternum IV (Fig. 12A) with

straight posterior margin, medially with rather deep and

broad notch; sternum VII mixed light-brown to black pi-

lose. Genitalia (Fig. 12B). With characteristic teaspoon

shaped appendix between cerci and surstylus; apical part

of surstylus with large almost squarish base and sudden-

ly narrowed, ventrally sharp tipped apex.

Material studied. Holotype & Eumerus truncatus:

“Maiorni“, “147” [white oval label with red text], “Museo

La Specola / coll. Rondani / HOLOTYPUS” [red label],

“Eumerus truncatus / bona sp./ [det] Vuji¢ 2014”, “Holo-

type 0 / Eumerus truncatus / Rondani, 1869 / det J. van

Steenis, 2016” [red label] (MZLS).

Additional material. Morocco: “Marokko / Antiatlas

/ S. Ait-Baha / 12.11.1997 / 30°00’N 9°02’ W / leg. M.

Hauser”, 3 JS (CSCA); “Marokko /11km NW Taliouine

/ 15.11.1997 / 30°34’N 8°00’ W / leg. M. Hauser”, 1 J

(CSCA); “Morocco 25km NE Tizinit / Massa Riv.

29.890N 9.595W 30m NN / leg. Schmid-Egger /

16.VI.2014 mall5 GS”, 2 0% (CSCA); “Morocco 20km

E Tizinit / Assaka 29.960N 9.530W 190m NN / leg.

©OZFMK

Review of the Eumerus barbarus species group from the western Mediterranean Basin 163

Schmid-Egger 16.V1.2014 mall16”, 1 S| (CSCA); Portu-

gal: “Portugal mal. trap Mertola / west of town / dry rain-

water brooklet/ UTM 29S6171-4166 / 21-23.1.2004 / leg

A.v. Eck”, 1 3 (AET); Spain: “Madrid / Arias Encobet”,

1 J (MNCN); “Belin chon (Cuenca) / 8-VII-1925 / J.M.

Dusmet”, 1 0 (MNCN); “Rio Peyales / Arena de San Pe-

dro / 15.V-4.V1.1976 /550 m Station 7”, “Espafia Avila /

Sierra de Gredos / P. Oosterbroek & / E. Boersma”, 1 &

(JSA); Tunisia: “Tunesia N Kasserine / 24.V.1999 1000

m asl /35°25.14’N 08°44.74°E /leg.O. & M. Niehuis”,

1 & (CSCA); “Tunisia S M’Saken / 5 km N Sidi Bou

Goubrine / 21.V.1999 ~ 1000 m amsl / 35°36.29’N

10°36.04’E/ leg. O. & M. Niehuis”, 1 “7 (CSCA); “Tune-

sia S M’Saken / “Tunesie / J.A.W. Lucas”, “Takrouana /

15-4-1977”, 1 J (NBC).

Remarks. In the synonymy of E. barbarus, Eumerus trun-

catus was mentioned (Pape & Thompson 2013). The lat-

ter is, however, a bona species clearly differentiated by the

characters given in the above description. Grkovi€ et al.

(2015) cited this species from eastern Mediterranean Basin

(Crete, Lesvos and Naxos islands). The species is record-

ed from Morocco, Portugal, Spain and Tunisia for the first

time.

DISCUSSION

The Mediterranean Basin is considered a biodiversity

hotspot and ranked 15" out of the 25 world hotspots (Mé-

dail & Quézel 1999; Myers et al. 2000; Médail & Myers

2004) and, as such, in need for global conservation prior-

ities (Sloan et al. 2014).

The family of Syrphidae has a high species diversity in

the Mediterranean Basin (e.g. Kassebeer 2000, 2001; Vu-

ji€ et al. 1999, 2001; Radenkovi€ et al. 2011; van Steenis

& van Steenis 2014; Pérez-Bafion et al. 2016) but not

many studies have included Syrphidae in analysis of bio-

diversity richness (Petanidou et al. 2013). The genera

Merodon Meigen, 1803 and Eumerus, both dependent on

bulbous plants in the larval and sometimes also adult stage

and with often very specialized life cycle (Ricarte et al.

2008; Rotheray & Gilbert 2011), are key genera for species

diversity in the Mediterranean Basin. The genus Merodon

is well studied with many supposed Mediterranean en-

demics (Hurkmans 1993; Marcos-Garcia et al. 2007; Vu-

ji€ et al. 2007; Radenkovi€ et al. 2011; Popovié et al 2015;

Sasié et al. 2016). The genus Eumerus is less well stud-

ied and, besides the two newly described species in this

paper, several studies indicate that there are many more

new Eumerus species to be expected in the Mediterranean

Basin (Grkovié et al. 2015; Chroni et al. 2017; Grkovié

ef-al.- 2019);

The Mediterranean Basin has many immediate and long-

term threats that put in risk its biodiversity, such as increas-

Bonn zoological Bulletin 66 (1): 145-165

ing tourism and growth of population (Hoekstra et al.

2005; Cuttelod et al. 2008; Riservato et al. 2009). Restric-

tions in water, land and energy resources, and development

of coastal tourism and urbanisation pose risks of further

habitat degradation (Underwood et al. 2009). The predict-

ed climate change will cause warmer summers and less

precipitation during spring, autumn and summer (Lionel-

lo 2012) causing a general drying and possible desertifi-

cation of the land. The warming of the Mediterranean

Basin will be more than the world average making the

global warming an even higher threat to this region. The

decrease in precipitation combined with the increased de-

mands of water for tourism, agriculture and the growing

population lead to an even more rapid loss of natural wa-

ter and drying of natural habitats, with an increasing risk

for forest fires. All these factors influence the extant flo-

ra, with the bulbous plants as most valuable community

of the Mediterranean Basin (Silva et al. 2008; Buerki et

al. 2012; Mendoza-Fernandez et al. 2015), and, conse-

quently, all the species within the specialized genus Eu-

merus are highly threatened too. We believe that the de-

scription of these new species adds to the need for pro-

tective measurements and reappraisal of local refuges (Mé-

dail & Diadema 2009; De la Montafia et al. 2011; Vujic¢

et al. 2016).

Acknowledgments. Christian Schmid-Egger (Berlin, Germany)

is credited for his company during fieldwork and for addition-

al material, and Wouter van Steenis (Breukelen, The Netherlands)

is recognized for his company in the MNHN. Ante Vuji¢ (Novi

Sad, Serbia) is thanked for providing literature. The following

persons are acknowledged for giving the possibility of studying

relevant material: Antonio Ricarte and Cinta Quirce (Alicante,

Spain), Mercedes Paris (Madrid, Spain), Christophe Daugeron

(Paris, France), Luca Bartolozzi (Firenze, Italy), Pasquale Cilib-

erti (Leiden, The Netherlands), Andre van Eck (Tilburg, The

Netherlands) and Joachim Ziegler (Berlin, Germany). Obediah

Sage (Sacramento, USA) is thanked for making the SEM mi-

crographs. The Royal Entomological Society (RES, London) is

acknowledged for the Outreach Fund (ORF) provided for the vis-

it to the MNHN.

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©OZFMK

Bonn zoological Bulletin 66 (2): 167—176

December 2017

George Bass, an early bird collector in the Pacific (1801-1802),

and some notes on early bird collecting on the Pacific Ocean islands

Justin J. F. J. Jansen

c/o Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, Netherlands. E-mail: justin.jansen@ naturalis.nl

Abstract. The arrival of two living Samoan Fruit-doves Ptilinopus fasciatus on 7 June 1803 in the harbour of Le

Havre, France with the return of one of the Baudin expeditionary vessels, the Naturaliste, marked the arrival of

the first live birds from the tropical Pacific in Europe. More specimens from the tropical Pacific followed on 24

March 1804 when the second Baudin vessel, the Géographe, arrived at Lorient, France. The person responsible

for collecting these birds was — most likely —- George Bass (1771-1803), who donated these specimens to the ex-

pedition commanded by Nicolas Baudin at Port Jackson, Australia in November 1802. This paper documents Bass’s

bird collecting activities, his 1801-02 voyage, the role of the Baudin expedition and the Muséum National d’His-

toire naturelle in Paris (e.g., donation, exchanges), and it documents the history of bird collecting in the tropical

Pacific — on locations visited by Bass — pre-1823.

Key words. George Bass; Nicolas Baudin; Pacific; Birds; Australia; Cook Expeditions.

INTRODUCTION

George Bass (1771-1803) is a well-known explorer of

Australia and to a lesser extent of the Pacific (Bowden

1952, Estensen 2005, 2009). Little has been published on

his ornithological activities in Australia (pre-1800), and

in the Pacific and New Zealand (1801-02). His connec-

tion with the expedition commanded by Nicolas Baudin

to Australia (1800-1804) has been published (Bowden

1952: 113, Estensen 2005: 160-161, Starbuck 2009: 105,

Jansen 2014, Jansen & van der Vliet 2015), in particular

the merchandise he sold to the expedition (e.g., Bass sold

pork, salt and other goods to the expedition). In addition,

he donated 160 ethnographic artifacts when the expedi-

tion was moored in Port Jackson in November 1802 (Hamy

1906, Péron 1994: 159-167, Starbuck 2009: 171, 212).

According to Horner (1987: 329), the number of artifacts

was 206, however, this was the total number of artifacts

collected during the entire Baudin expedition (Muséum

d’ Histoire naturelle Lé Havre, France,Ms21001; Starbuck

2009: 171). A part of the whaleboat in which Bass dis-

covered Bass Strait was also given to the expedition (Bow-

den 1952: 68); its current whereabouts, as well as the

ethnographical material he donated, are at present un-

known. Ethnographical material was confiscated by

Napoleon’s wife, the Empress Joséphine de Beauharnais

and incorporated into her private collection at her Malmai-

son estate, and subsequently destroyed in 1814 by invad-

ing troops during the final months of Napoleon’s reign,

while the rest was dispersed and sold in 1829 after

Josephine’s son Eugene de Beauharnais died (cf. Hamy

Received: 26.03.2017

Accepted: 09.11.2017

1906, Horner 1987: 329, Péron 1994: 224, Duyker 2006:

212, 289, Attenbrow 2010: 86, Fornaseiro eft al. 2010:

355). Bass also donated notes to the Baudin expedition on

the vocabulary and grammar of the “savages” at Port Jack-

son (Starbuck 2009: 171), as well as details regarding Bass

Strait. Bass boarded the Géographe at least once for a vis-

it, on 14 November 1802, in the company of Governor

King and Captain Kent (Starbuck 2009: 102).

Until 1823 expeditions in the Tropical Pacific are few

— but included those by Bougainville, Cook, Malaspina,

Bruni d’Entrecasteaux, Vancouver, Krusenstern, Frey-

cinet, Kotzebue (Brosse 1983) — but the first to make no-

table collections after the Cook expeditions was the Co-

quille in 1823 (Holyoak & Thibault 1982). This paper dis-

cusses the birds almost certainty collected by Bass dur-

ing his Pacific travels and documents his early collecting

in Australia.

Birds collected by Bass prior to 1801: We find the first

evidence of Bass collecting natural history specimens in

a letter written to his wife Sarah Bass (1798, State Library

NSW, MLMSS 6544 / 6 letter 2). In this letter, he states

his plans to accompany Mathew Flinders to Tasmania to

collect natural history specimens for Joseph Banks and the

Linnean Society in London (W hittell 1954: 40-46).

In a

(1743-1820) dated 27 May 1799 (State Library NSW, Se-

ries 72.005, CY 3682 11-14), Bass mentions, in describ-

ing his voyage during which he and Mathew Flinders dis-

letter to the famed naturalist Joseph Banks

covered the Bass Strait: ‘In the course of this expedition

I collected at different times some few new subjects, both

Corresponding editor: T. Topfer

168 Justin J. F. J. Jansen

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i Weellington

—»>— voyage to Hawaii Islands

1. Dusky Sound (New Zealand)

12, 13, 14. Huahine, Raiatea, Tahau

—>—_ return voyage 2 & 3. Raivavae, Tubuai (Austral Islands) 15. Marquesas

---»-- possible alternative route 4. Tahiti 16.Tonga

between Samoa and 5.Moorea 17. Tutuila (Samoa)

Cook Islands 6. Kiribati 18. Fiji

7,8,9, 10. Hawaii, Molokai, Kauai, Oahu 19. Vanuatu

Scale Il 000.000

Licrettar Projection

30 et ep a

6 ae

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(Hawaiian Islands)

11. Moorea

20. Cook Islands

Fig. 1. Reconstruction of the journey made by George Bass between 21 November 1801 and 14 November 1802.

animals and plants; but they were from time to time de-

stroyed by rain or by the seas which in bad weather the

boat was constantly moving; so that none remained upon

my arrival at Port Jackson’. With this letter, Bass also sent

to Banks a skull of a Common Wombat Vombatus ursi-

nus (Estensen 2005: 120, Pigott & Jessop 2007: 210), most

likely collected west of Port Jackson, as well as skins of

two birds: a Spotted Quail-thrush Cinclosoma punctatum

dovei (described by Bass as a beautiful “banded runner”

that had been chased and caught by a greyhound), collect-

ed in early January 1799 along the River Derwent in south-

east Tasmania (Bowden 1952: 78, 83), and a Superb Lyre-

bird Menura n. novaehollandiae (described by Bass as a

rare and new pheasant) (Bowden 1952: 83). The last is the

bird mentioned by Collins (1804: 65) and Olsen (2001:

33) as the specimen collected along the Hawkesbury Riv-

er near Port Jackson.

Bonn zoological Bulletin 66 (1): 167-176

Bass regularly went out from Port Jackson in search of

new orrare natural history specimens (Bowden 1952: 33).

William Paterson, Lieutenant-Governor of New South

Wales and a collector of natural history specimens for

Banks, probably encouraged Bass’s collecting activities,

as both men knew each other and notes from Bass’s ob-

servations on birds can be found in Paterson’s diaries (cf.

Bowden 1952: 66, 71-75, 78). Bass met Banks when in

England in 1800, and presented Banks with some addi-

tional information on the items he had sent (it is unknown

if Bass sent Banks more than one shipment of specimens).

In the entry on Bass in the Australian Dictionary of Bi-

ography (http://adb.anu.edu.au/biography/bass-george-

1748), Bowden mentions natural history publications by

Bass, but unfortunately, I have not located them.

While at Port Jackson, Bass was made an associate

member of the Society for Promoting Natural History, in

©OZFMK

George Bass, an early bird collector in the Pacific (1801-1802) 169

the same decade as Paterson (1755-1810) (who subse-

quently became a full member), John William Lewin

(1747-1795) and botanist Robert Brown (1773-1858).

The Society for Promoting Natural History was later

merged with the Linnean Society (Neville 2012: 72).

In addition to the sites mentioned below, Bass could al-

so have collected birds at Norfolk Island during visits in

September—October 1796 and June-July 1798,

Caledonia and Tuvalu in June 1799, the Gilbert Islands

(Abemama, Nonuti, Taputea), Marshall Islands and Mar-

New

iana Islands in July 1799, Saint Helena in early 1800, as

well as some other places visited during his voyages (Bow-

den 1952, Estsensen 2005, 2009).

Pacific Journey: Bass was contracted by Governor Philip

Gidley King (1758-1808) to sail to Tahiti and transport

salted pork back to Port Jackson, as food was scarce in

Port Jackson but pigs were plentiful in Tahiti. From Port

Jackson, Bass sailed into the Pacific on 21 November 1801

with Charles Bishop (1765-1810), and didn’t return un-

til 14 November 1802 (Bowden 1952: 106-113, Bowden

1980: 86-87, Estensen 2005: 149-161). In quotes below

the supposed visits made by Bass (see Fig. 1).

Bass first landed at Dusky Sound, South Island, New

Zealand (6-21 December 1801) (Anon. 1908a,b,c, Hamy

1906: 31-32, Péron 1994: 230-231, Bowden 1980, Rus-

den 2011: 415-416), and then proceeded on to the Aus-

tral Islands (Raivavae, Tubai) (‘December 1801 or/and Jan-

uary 1802’) (Estensen 2005: 150-151), Tahiti (24 Janu-

ary — 6 February 1802, ‘July’-19 August 1802) (Anon.

1967, Anon. 1908b: 226, Hamy 1906: 30, Péron 1994:

229, Bowden 1952, Estensen 2005), Moorea (‘February

1802’) (Estensen 2005: 152), Kiribati (‘February or/and

March 1802’) (Estensen 2005: 154), the Hawalian Islands

(Hawaii, Molokai, Kauai, Oahu) (11 March —- July 1802)

(see below), Moorea (August 1802) (Anon. 1967), French

Polynesia (Huahine, Raiatea, Tahau (August 1802) (Bow-

den 1952: 112)), the Marquesas (island(s) visited un-

1802’) (Estensen

2005: 157), Tonga (island(s) visited unknown, ‘Septem-

ber or October 1802’) (cf. Hamy 1906: 30, Bowden 1980:

86-87, Péron 1994: 229), Samoa (Tutuila) (‘September

or/and October 1802’) (cf. Hamy 1906: 28-32, Maude

1964: 262, Gilson 1970: 67, Pearson 1970: 140, Bowden

1980: 86-87, Linnekin 1991: 16, Péron 1994: 226-231,

Tcherkézoff 2008: 73), (possibly) Fiji (‘October 1802’)

(Bowden 1980: 86-87, Estensen 2005: 159), (possibly)

Vanuatu (‘October 1802’) (cf. Bowden 1980: 86-87) and

Cook Islands (‘October or/and November 1802’) (cf.

Hamy 1906: 31, Péron 1994: 230). During Bass’s stay in

Tahiti, Lewin was also present (Neville 2012: 72-75).

known, ‘August or/and September

Bass’s visit to the Hawaiian Islands is notable. Docu-

mentation of his visit includes several letters (see Bow-

den 1980), including letters from Kealakekua Bay, Hawaii

(George Bass to Sarah Bass, 20 May 1802, ZML MSS

Bonn zoological Bulletin 66 (1): 167-176

6544, ML (Mitchell Library), Sydney) (Bowden 1980:

86-87, Estensen 2009: 154), Oahu (6 letters between Bass

and James Innes) (the letters to Innes, who lived in Port-

sea (Hampshire), England, were sold at Christie’s, Lon-

don on 15 October 2009, then purchased by Hodern

House, Sydney (an antique shop) and subsequently re-sold

to a private collector (M atthew Fishburn in litt. 18 Novem-

ber 2014); one of the letters, dated 23 May 1802, was writ-

ten by Bass at Waikiki Bay, Oahu), Kauai (Bowden 1980:

86-87) and Molokai (George Bass to Elisabeth Bass, 20

May 1802, ZML MSS 6544, ML, Sydney) (Estensen

2009: 151-153). Four ethnographic artifacts donated by

Bass from the “{fles Sandwich” (= Hawaiian Islands) were

listed as no. 74 and nos. 90-92 in the manifest of the

ethnographic artifacts that returned with the Baudin ex-

pedition written by Francois Péron, the expedition’s sur-

viving zoologist (Hamy 1906: 30, Péron 1994: 229-230).

Of special interest is no. 74, which Péron described as a

“Grand piéce d’étouffe de plusieurs aunes de longueuren

trois doubles, dont 1’un noir, l’autre jaune, le 3e jaune strié

de noir” (“Large piece of fabric several yards in length in

three doubles, one black, the other yellow, the third

streaked with black”). The striking colors indicate that this

may have been a feather cloak made from the black and

yellow feathers of the Hawaii Mamo Drepanis pacifica

and Hawaii Oo Moho nobilis.

MATERIAL & METHODS

As part of an ongoing research into the bird collections

made at the Baudin expedition (1800-1804), all known

archival material and specimens brought back are re-

searched (Jansen 2014, 2015, 2016, 2017a). For this, spec-

imens were researched at: Universita degli Studi di Firen-

ze, Italy (C.G.U.); Muséum d’histoire naturelle Blois,

France; Museum-Aquarium de Nancy, France; Muséum

d’ Histoire Naturelle de La Chaux-De-Fonds, Switserland

(MHNC); Muséum d’histoire naturelle de la Ville de

Geneve, Switzerland (MHNG); Musée George Sand et de

la Vallée Noire, La Chatre, France (MLC); Muséum na-

tional d’ Histoire naturelle, Paris (MNHN); Naturalis Bio-

diversity Center, Leiden, Netherlands (N aturalis); N ation-

al Museum of Scotland, Edinburgh, Scotland (NMS);

Naturhistorisches Museum Wien, Austria (NHMW) and

Museo di Zoologia, Universita di Torino, Italy (MZUT).

Additional information was gathered at the MNHN library,

the Archives Nationales, Pierrefitte-sur-Seine, France; and

in Muséum d’histoire naturelle du Havre, France. I fur-

ther examined specimens and documents in the Linnean

Society, London, UK; Natural History Museum, London,

UK; Natural History Museum, Tring, UK; Musée Bouch-

er-de-Perthes, Abbeville, France; Museum fir Naturkunde

Berlin, Berlin, Germany and Naturhistoriska riksmuseet,

Stockholm, Sweden.

©ZFMK

170 Justin J. F. J. Jansen

Fig. 2. MNHN 2005-2563, Samoan Fruit-dove Ptilinopus fasciatus: collected on Samoa, caught in ‘September or October 1802’

and shipped alive on the Naturaliste. It arrived alive in France at7 June 1803, but died soon after. (Photograph by Justin J.F.J. Jansen,

RESULTS

Arrival in Europe ana dispersal of specimens

Donation. W hen supplying the Baudin expedition (that

visited Tenerife, Australia, Timor, Mauritius and South

Africa), Bass donated to the expedition ethnographical

items (see above), and the presence of birds from the same

locations link them. These items were all shipped back lat-

er at the Géographe, but at least two live birds, a pair of

Samoan Fruit-doves Ptilinopus fasciatus (Fig. 2) — most

likely — collected by Bass, arrived with the Naturaliste

(Jangoux etal. 2010). In addition to a variety of other live

birds and mammals, the Naturaliste brought back 599 bird

specimens according to Louis Dufresne, taxidermist at the

Muséum national d’ Histoire naturelle, Paris, France (2 Au-

gust 1803, Archives Nationals AJ/15/590). Possibly be-

cause all the specimens that returned on the Naturaliste

had already been packed (by Péron and Charles-A lexan-

Bonn zoological Bulletin 66 (1): 167-176

dre Lesueur) when Bass arrived back in Port Jackson on-

ly the two Samoan Fruit-doves arrived in France on the

Naturaliste which sailed directly from Sydney to Le Havre

with short stops only at King Island and M auritius. The

G éographe arrived in France in March 1804 with 422 bird

skin specimens and 34 live birds (Dufresne inventory of

28 April 1804,

MNHN).

Collections Mammiferes et Oiseaux

Documentation. Several lists of the ornithological con-

tents from both ships were compiled after the Géographe

specimens arrived at the MNHN on 18 July 1804, join-

ing the material shipped back on the Naturaliste that had

arrived there in July 1803 (Burkhardt 1997: 505, Starbuck

2009: 213). Dufresne compiled three incomplete lists, one

now in the Collections Mammiferes et Oiseaux in the

MNHN (undated), one at the Muséum d’Histoire naturelle,

le Havre (Ms 21036) (dd. 26 June 1804) and one in the

Archives Nationales, Paris (A J/15/592) (dd. 27 June 1804)

©OZFMK

George Bass, an early bird collector in the Pacific (1801-1802) 171

Fig. 3. MNHN 2014-432, Tui Prosthemadera n novaeseelandiae, collected at Dusky Sound, South Island, New Zealand, 6-21

December 1801, arrived at 18 July 1804 in France aboard the Géographe. (Photograph by Phil Koken, February 2015; © Muséum

N ational d’ Histoire naturelle, Paris.)

while a fourth list (undated), also in the Collections Mam-

miferes et Oiseaux MNHN, is probably a transcription by

an unknown author (Jansen 2016). All four lists only iden-

tify the specimens by family and by the number of species

and specimens, but do not identify the individual species

Bonn zoological Bulletin 66 (1): 167-176

within each family listed; moreover, they include only ten

annotations of collecting localities. Even so, we find on

these lists the collecting locality “d’otaiti”, i.e., “from Tahi-

ti”, mentioned twice, first in the entry for “Grive nouveau”

(“new thrush”), possibly the Society Island Flycatcher Po-

©OZFMK

72 Justin J. F. J. Jansen

Fig. 4. MNHN AC. 9818, Polynesian Triller Lalage maculosa ssp, collected in 1801-02 in the Pacific, arrived at 18 July 1804

in France on-board of Géographe. (Photograph by Justin J.F.J. Jansen, February 2015; © Muséum National d’Histoire naturelle,

Paris.)

marea nigra (two specimens listed, both no longer pres-

entin the MNHN), annotated as “tres beau d’otaiti” (“very

beautiful from Tahiti”). Tahiti is mentioned again in the

entry for “Mycteria — Jabiru”, probably referring to the

Black-necked Stork Ephippiorhynchus asiaticus australis

of Australia (one specimen), annotated with “et un bec d’o-

taiti” (“and one beak from Tahiti”), the origin of the lat-

ter species apparently misidentified (list in Collections

Mammiferes et Oiseaux MNHN). Included in the list of

live animals brought back on the WNaturaliste were

“Tourterelles des iles des Navigateurs” (“Turtledoves from

the Navigator Islands”) (cf. Jangoux etal. 2010: 271,281).

Both turtledoves and fruit-doves were referred to as

“tourterelles” at that time (cf. Buffon 1765-1783: plate

142). The only pigeon present to date in MNHN (Fig. 2)

from the ‘iles des Navigateurs’ (= Samoa) is a single

Samoan Fruit-dove, the pedestal underside confirms its

origin (arrived in An XI = via the Naturaliste according

to Louis Dufresne on the pedestal underside). These spec-

imens, of the “Grive nouveau” and the “Tourterelles des

iles des Navigateurs”, almost certainly came from Bass,

as the Baudin expedition did neither visit Tahiti nor Samoa.

However, crew-member Hyacinthe de Bougainville, men-

tioned as origin the Sandwich Islands (de Bougainville,

Archives nationales de France, 155 AP6).

The Birds. Bass most likely collected four specimens of

birds presently in the MNHN; however, as all four lack

their original labels (which applies as well to all specimens

(1000’s) in the MNHN acquired prior to 1850), their ori-

gin remains speculative. The four specimens are: two Tui

Prosthemadera novaeseelandiae (only one, registered as

Bonn zoological Bulletin 66 (1): 167-176

MNHN-ZO-2014-431, is still present (Fig. 3); the other

has not been located by me but it is noted in the acquisi-

tion books and was present in 1856), most likely collect-

ed by Bass Dusky Sound, New

Zealand between 7-21 December 1801 (its origin is doc-

at Facile Harbour,

umented in the pedestal underside that mentions “Nouvelle

Zélande / La Découverte an 12 / Peron et Lesueur / No

80 / 10116 / Prosthemadera novaezeelandiae / Gm);

Samoan Fruit-dove (MNHN-ZO-2005-2563, Fig. 2), col-

lected by Bass in Samoa in ‘September or October 1802’

(this is one of the two captive Samoan Fruit-doves noted

above that arrived on the Naturaliste in 1803; the other is

no longer present in the MNHN) (Jangoux ef al. 2010:

271, 281));

(MNHN-ZO-2016-523, Fig. 4, a skin that was mounted

before) (its new label mentions “Exp Baudin / Ex. Peron

and Polynesian Triller Lalage maculosa

et Lesueur / “Lalage Pacifica”), was collected on an un-

specified Pacific locality. Also, on 27 March 1805 a para-

keet from Tahiti was ‘sold’ to Dufresne, for his private col-

lection (Archives Nationales AJ/15/593).

The notes at the Pigeon and Tui are from Louis

Dufresne’s hand, who mounted birds that arrived in the

museum to be used for the galleries almost directly after

arrival in the museum (Archives Nationales AJ/15/592 &

593, Jansen 2016). This makes these specimens amongst

the best-documented birds from the tropical Pacific col-

lected pre-1823 (Medway 2004). All other Pacific birds

in European collections lack their original label, and on-

ly very few are proven to be collected from the Tropical

Pacific before 1823 (Burton 1968, Medway 2004, Stein-

heimer 2005).

©OZFMK

George Bass, an early bird collector in the Pacific (1801-1802) 173

Exchanges

Only half of the 1.055 specimens (all skins) that arrived

with the Baudin expedition were mounted for exhibition

in the MNHN galleries and about 397 are still present in

European collections (Jansen 2017a). The remainder — es-

pecially those birds that arrived on the Géographe — were

used for exchanges and donations (Jansen 2014, 2016,

Jansen & van der Vliet 2015). It is uncertain how many

bird specimens were in good condition upon the expedi-

tion’s return, considering that the ships were probably in-

fested with swarms of cockroaches and rats (Laurent

1997), or how many deteriorated soon after their arrival

in France, noted is that some had to be thrown away (Louis

Dufresne, 7 May 1811, Laboratory MNHN).

No less than 178 birds were disposed of Empress

Josephine (Geoffroy at 23 September 1804, Archives Na-

tionales AJ/15/593) (93 birds from the Naturaliste and 85

birds from the Géographe, most likely donated in two ship-

ments). Donations were made to schools in Rouen and

M oulins (Jansen 2014, 2016, 2017a) and to Vienna in 1815

as partial repayment or

Napoleonic Wars (Bauer & Wagner 2012; Archives Na-

tionales France, AJ/15/840). Baudin specimens were al-

material looted during the

so probably amongst the 847 birds donated to the Nether-

lands for the same reason (Archives Nationales France,

AJ/15/840) (Jansen 2016: 104).

M ost specimens that arrived on the Géographe were pur-

chased by or donated to the Baudin expedition, as estimat-

ed by 93 specimens from Géographe still at the MNHN;

only 23 birds of which were collected in Australia, a large

portion originated from countries not visited by the Baudin

expedition, for example: Sumatra, Java, Celebes, Mada-

gascar and India. Itis unknown how many birds, from the

total number that arrived on the Géographe, originated

with Bass. That all these precious specimens left the

MNHN was primarily due to the lack of an ornithologist

as curator of the bird collection. At that time a visitor, the

clerk of Bulletin des Lois, Louis Vieillot, described a por-

tion of the birds brought back as new birds (cf. Mearns

& Mearns 1998: 133-134).

The speed with which new specimens were dispersed

is shown by the purchase of 30 skins in 1806 by Leopold

von Fichtel from John Latham; these birds, currently in

Naturhistorisches Museum Wien, Vienna, Austria, were

collected at Dalrymple Bay, Tasmania in April 1802 (Au-

togr. 21/35 L.v. Fichel, Osterreichische Nationalbibliothek,

Vienna, Austria) (Bauernfeind 2004: 557), an area visit-

ed by the Baudin expedition in March-April 1802 (Nat-

uraliste). The area was also visited by Robert Brown on

1-17 January 1804 (cf. Vallance etal. 2001; Jansen 2014),

who collected two bird specimens there (NHMUK Lon-

don, Brown, M.S. Descriptions Animals HMS Investiga-

tor 1801-1805. Aves). From these two specimens for one

its whereabouts remain unknown (one was donated to Lin-

Bonn zoological Bulletin 66 (1): 167-176

nean Society in 1818 (list in Linnean Society (S.P. 156)

and a typed version of the BM donation in NHM, Lon-

don (NHM London, Brown, M.S. Descriptions Animals

HMS Investigator 1801-1805. Aves)). So, part of these

Dalrymple Bay specimens have (if the rumoured date is

correct) — most likely — originated from the Baudin expe-

dition (visited by the Naturaliste in April 1802), the oth-

er from William Paterson who resided here for a few years

between 1804 and 1809.

Other specimens ended up in the private collection of

Coenraad Jacob Temminck, then a private collector, and

later his collection formed the nucleus of the collection

we now know as Naturalis Biodiversity Center, Leiden. In

his collection six pacific specimens arrived between

1803-04 and 1807 and was enlarged to 22 specimens

(Jansen 2017b: 353), and amongst them is a Tonga Fruit-

(RMNH.AVES.213951)

(maybe this is the other ‘Samoan Fruit-dove’ shipped alive

dove Ptilinopus porhyraceus

on the ship Naturaliste). But a fair number ended up in

numerous other collections in Europe (see for more de-

tails Jansen 2017a).

Some specimens from the Baudin expedition may also

have been among specimens from MNHN that entered the

Lever Museum in London (Jansen & van der Vliet 2015).

DISCUSSION

We find various bird specimens from the tropical Pacific

and New Zealand collected prior to 1823 (appendix 1) in

older collections (e.g., Berlin, Cambridge (USA), Edin-

burgh, Gottingen, Hannover, Leiden, LIVCM, MNHN,

Stockholm, NHM, Turin, Vienna) that are still in good con-

dition, probably due to their having been treated with ar-

senic soap either when they were collected or upon their

arrival in Europe (the recipe for arsenic soap was known

to the French and a few others like C.J. Temminck and

Francois Levaillant (cf. Steinheimer 2003, Jansen & Roe

2013, Jansen & van der Vliet 2015)). Few expeditions

and/or collectors are known to have been active in the lo-

cations visited by George Bass (see appendix 1).

In this paper, it has been shown that: 1) George Bass

has been neglected as a natural history collector; 2) There

is evidence that he had direct links with the Baudin expe-

dition, as for ethnographical material (proven) also or-

nithological material could be donated to the expedition

and eventually ended up in the MNHN;; 3) Few, if any, ex-

tant pre-1823 bird skins are unambiguously attributable

to specific collectors, due to inadequate labelling; 4) How-

ever, locality information and known routes of Baudin and

of Bass suggest Bass probably collected some skins that

were taken to France; 5) The fate of Bass’s now poorly-

documented specimens illustrates the problem of deter-

mining the provenance of tropical Pacific specimens that

were circulating in Europe in the early 19th century.

©ZFMK

174 Justin J. F. J. Jansen

I encourage further research to find missing documents

and to locate additional information of tropical Pacific and

Australian specimens collected by George Bass.

Akcnowledgements. | thank Erik Ahlander (Swedish Museum

of Natural History, Stockholm, Sweden), Lynda Brooks (Linnean

Society London), Donna Campbell (State Library of New South

Wales), Alice Cibois, Mariane Estensen, Matthew Fishburn

(Hodern House, Sydney), Paul Martyn Cooper (NHMUK, Lon-

don), Hein van Grouw (NHMUK, Tring), Alison Harding

(NHMUK, Tring), Ulf Johansson (Swedish Museum of Natural

History, Stockholm, Sweden), Adrienne Kaeppler, Murray

Lord, David M oore, Tony Parker (LIVCM), Laurent Raty, Roger

Safford, Jean-Claude Thibault, Benjamin Thomas (Trinity Col-

lege, Melbourne, Australia) and Roland E. van der Vliet, for pro-

viding additional information. Patrick Boussés, Anne Préviato

and Claire Voisin for hosting my visits to MNHN. Andrew Black,

Storrs L. Olson and Bert Theunissen reviewed an earlier man-

uscript and made many useful additions. André van Loon made

the figure in this article. Richard S. Roe found time to discuss

the structure and content of the first editions and to read and com-

ment on these editions. I received support from the SYN-

THESYS Project http://www.synthesys.info/ that is financed by

European Community Research Infrastructure Action under the

FP7 “Capacities” Program.

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176 Justin J. F. J. Jansen

APPENDICES

PREVIOUS COLLECTING IN LOCATIONS VISITED BY BASS

(1766-1823)

To establish the importance of the collecting carried out by Bass,

I analysed ornithological collecting activity up to the early 1800s

in the order of the countries visited by Bass. Up to 1823 expe-

ditions in the Tropical Pacific are few and the first large expe-

dition was the Coquille in 1823 that visited the region (Holyoak

& Thibault 1982).

Australia: See Jansen (2017a).

New Zealand: Cook landed at New Zealand during all three ex-

peditions (cf. Lysaght 1959; Whitehead 1978), and 8 specimens

from New Zealand are known from Banks’s private collection

(Medway 1979). Although few Cook specimens are still extant

(Burton 1968; Medway 2004), specimens known or alleged to

have been collected in New Zealand during the Cook expedi-

tions are (or were) still present in Cambridge (USA) (Paradise

Shelduck Tadorna variegata (1788 collected) not present any-

more, according to http://mcezbase.mcz.harvard.edu/Speci-

menSearch.cfm, accessed 6 June 2015), NHM (three birds), Lei-

den (three birds) and Liverpool Museum (hereafter LIVCM),

(four birds) (cf. Steinheimer 2003). The next expedition to New

Zealand was with the Recherché and Espérance (apparently, no

birds collected in New Zealand, cf. Stresemann 1953b). Captain

A. Malaspina landed in 1793 in New Zealand, but none of his

specimens entered museums and none have survived (Olson

2006). The Coquille visited New Zealand in A pril/M ay 1825 and

approximately 50 birds were collected (MNHN Library, Ms 354).

The Astrolabe took 11 New Zealand birds back to France.

Raivavae: Although Cook sighted this island on his third expe-

dition; he did not land there (Brosse 1983).

Tubai: Cook also sighted this island on the third expedition, but

did not land there either (Brosse 1983).

Kiribati: Cook described this island after a possible visit on 24

December 1777, during the third expedition (Brosse 1983).

Tahiti: Cook visited Tahiti during all three expeditions (Brosse

1983); atleast 15 birds from Tahiti were recorded in Banks’s pri-

vate collection (Medway 1979). Itis unknown if birds are brought

back to Europe/Russia by the frigates Boudeuse and L’etoile (cap-

tained by L.A. de Bougainville, 1766-69, cf. Laissus 1978) and

(captained by O. von Kotzebue,

1823-26), Blossom (captained by F.W. Beechey, 1825-1828, cf.

Beechey 1839, no Pacific specimens are reported in here), Vos-

tok and Mirny (captained by F. Bellinghausen, 1819-1821) and

Senyavin (captained by F.P. Litke, 1826-1829). At least 36 birds

collected in Tahiti during the visit of the Coquille arrived in the

MNHN in 1829.

the ships Predpriyatiye

Moorea: Cook visited Moorea on the third expedition (Brosse

1983).

Hawaiian Islands: Visited by Cook during the third expedition;

at least 53 birds collected in Kauai and Hawaii ended up in

Banks’s private collection (Medway 1979) and were subsequent-

ly dispersed, as were Hawaiian birds from the Cook expedition

deposited in other collections (cf. Steinheimer 2003; Jansen &

Roe 2013). The ship King George (captained by N. Portlock) was

Bonn zoological Bulletin 66 (1): 167-176

mainly at Oahu from 26 May to 13 June 1786, that resulted in

an unknown number of specimens arrived in England (cf. Dixon

1789; Olson & James 1994). From the Uranie voyage, 9+ birds

were collected in August 1819 (Hawaii, Oahu) (archives

MNHN).The ship HMS Blonde (captained by G.A. Byron) vis-

ited various islands in May-July 1825 and collected 21 bird

species (Callcott et al. 1826: 248-252). The ships Nadezhda and

Neva (captained by A.J.R. von Krusenstern) the brig Rurik (cap-

tained by O. von Kotzebue, 1815-1818) expeditions visited the

islands, but apparently, no birds made their way back (maybe

from the later via J.J. Dussemier to the MNHN). Birds may have

been collected when the Blossom visited Oahu in 1826 (Brosse

1983), but no Hawaiian specimens are mentioned in Beechey

(1839).

Huahine: Visited by all three Cook expeditions (Brosse 1983).

Raiatea: Visited by the second and third Cook expeditions (M ed-

way 1979); at least eight birds from these expeditions were in

Banks’s private collection (Medway 1979).

The Marquesas: Visited by the second Cook expedition (Brosse

1983). A lone bird was brought back by Nadezhda and Neva

(Holyoak & Thibault 1982).

Tonga: Visited by the second and third Cook expeditions (Brosse

1983). No less then 40 birds from Tonga are known from Banks’s

private collection (Medway 1979). In 1793, Recherché and Es-

pérance (1791-1794) brought some birds back to France (at least

five birds, Stresemann 1953). Commander Malaspina landed in

Tonga in 1793; some birds were collected, but they were neg-

lected and have vanished (Olson 2006: 43). The Astrolabe ex-

pedition collected 51 birds in Tonga in 1827 (inventory of 520

birds that arrived back from the whole expedition from 25 May

1829, MNHN Ms 2223).

Notable is the description from at least one living Tonga Fruit-

dove by Robert Brown brought back on the HMS Investigator

in 1805 to England, however it “Died from cold on the HMS /n-

vestigator on crossing from New Wales to England” as men-

tioned by Brown (NHM London, Brown, M.S. Descriptions An-

imals HMS Investigator 1801-1805. Aves. No 196). This bird

is not on the list of donations of his 78 specimens in both the

British Museum as to the Linnean Society of London, so it’s

whereabouts are unknown. Its source is unknown, it is not George

Bass, as both men did not meet in Bass’s stint in Sydney after

returning between November 1802 and February 1803 from the

Pacific (Brown returned at 9 June 1803 to Sydney).

Samoa: The islands were visited by the ships Predpriyatiye

(1823-26) and Uranie (1819) but no known birds were brought

back (Brosse 1983).

Fiji: Visited by the second Cook expedition (Brosse 1983). One

bird was collected on the Astrolabe expedition (inventory of 520

birds from 25 May 1829, MNHN Ms 2223).

Vanuatu: Visit at the second Cook expedition, Recherché and Es-

pérance as the Astrolabe (1826-1829) expeditions, butno known

specimens were brought back (cf. Brosse 1983).

Cook Islands: Apparently, a single bird was collected here in

1798 by HMS Discovery (captained by G. Vancouver, cf.

Holyoak & Thibault 1982). HMS Blonde visited Mauke in Au-

gust 1825, and collected were a kingfisher, a pigeon and a star-

ling (Olson 1986).

©OZFMK

Bonn zoological Bulletin 66 (2): 177-181

December 2017

A new species of Meoneura Nitzsch from Japan (Diptera: Carnidae)

Jens-Hermann Stuke'* & Toshihiro Tago’

' Roter Weg 22, D-26789 Leer, Germany

? 29-13-101 Motogou 1-chome kawaguchi Saitama 332-0011 Japan

* Corresponding author. E-mail: jstuke @zfn.uni-bremen.de

Abstract. A new species of the genus Meoneura (Diptera: Carnidae), i.e. Meoneura issunboshii sp.n., is described

from Japan.

Key words. Diptera, Carnidae, Meoneura, sp.n., Japan.

INTRODUCTION

The family Carnidae is a member of the superfamily

Carnoidea (Diptera: Schizophora: Acalyptratae). The

flies are only up to 2 mm long and mainly black. Due to

the similarity to several other, more common acalyptrates,

Carnidae are recognised only by experts and are easily

overlooked in samples. The larvae of Carnidae are

saprophagous and develop in various kinds of dead organ-

ic material as dung, birds’ nests or fungi. To date only five

extant and one fossil genera are recognised. Carnidae com-

prise 116 valid species distributed as follows: 10 A frotrop-

ical species, 26 Nearctic species, 6 Neotropical species,

14 Oriental species, and 77 Palaearctic species (Brake

2011, Stuke unpublished). The highest num ber of species

is known from Europe (52 species; Brake 2011, Stuke un-

published), probably as a result of the long history of

dipterological research, and it is very likely that the lack

of Carnidae described from the Eastern Palearctic Region,

and the low number of species recorded from this region,

is a result of a comparably poor collecting activity.

Only three Carnidae species are reported from Japan so

far. Iwasa et al. (2000) recorded Carnus hemapterus

Nitzsch, 1818 from Fukagawa (Hokkaido), and Iwasa et

al. (2014) found Carnus orientalis Maa, 1968 from Ku-

nigami, (Okinawa). Tago (2011) reported Meoneura tri-

angularis Collin, 1930 from Tokio, representing the first

record of this genus from the region. With 98 valid species,

Meoneura Rondani, 1856 is the genus of Carnidae with

the largest number of species worldwide. Due to an ex-

cellent drawing of the male genitalia provided by Tago

(2011), it became evident that this was a new species and

is herewith described in this study.

Received: 21.10.2017

Accepted: 23.11.2017

MATERIALS AND METHODS

All material reported here was previously published by

Tago (2011), later sent to and examined by JHS. The ab-

domen of three male and two female specimens were dis-

sected, macerated for about three hours in NaOH (4) SO-

lution and stored in a microvial in glycerine together with

the specimen on the insect pin. The morphological termi-

nology follows Cumming & Wood (2009) and Buck &

M arshall (2007). The terms that are used to describe the

postabdomen are additionally illustrated in Figures 1-3.

A separate sclerite, ventrally beneath the protandrium

(syntergosternite 6-8 SEnSU W heeler 2010: 1102, Fig. 5),

is recognised as tergite 7 (Buck & Marshall 2007: 14, Fig.

16). The most anteroventral seta on the face is interpret-

ed as the vibrissa, those above the vibrissa as supravib-

rissal setae, and those posterior to the vibrissal seta on the

gena as genal setae. The labels of the holotype are listed

and numbered in the order found, commencing with the

uppermost. The original content of the label information

is cited between double quotation marks ( “ ) and line

breaks are indicated by a slash mark (/ ).

Digital images were taken using a Zeiss Axioskop 40,

Zeiss A-Plan 10x / 0.25 Ph 1 Varl with a Nikon D7000

mounted ona Zeiss 45 60 05 2.5 x. The pictures were used

as template and the drawings were made with Affinity De-

signer 1.6.0 on Wacom DTZ -1200W.

Corresponding editor: X. Mengual

178 Jens-Hermann Stuke & Toshihiro Tago

DESCRIPTION OF THE NEW SPECIES

Meoneura issunboshii sp. n.

Figs 1-4

Meoneura triangularis Collin, 1930. Tago (2011).

Diagnosis. M ales of Meoneura issunboshii sp. n. are eas-

ily identified by the unique shape of the falcated sursty-

lus with 4-5 dorsal setae (Fig. 2), the characteristic large

lamella (larger than surstylus) covered with long setae

(Figs. 1-2), and the short epandrium (shorter than sursty-

lus, Fig. 1). Females of Meoneura issunboshi sp. n. can-

not be identified at species level with morphological char-

acters.

At first glance the postabdomen of Meoneura issun-

boshii sp. n. resembles several other Meoneura species

with obvious long and setulose structures: Meoneura

amurensis O zerov, 1986 can easily be ruled out because

the large and setulose structure projecting from the epan-

drium is identified as the surstylus by Ozerov (1986), and

with setae of Meoneura

amurensis (see Ozerov 1986: 85) does not occur in Me-

oneura issunboshii sp. n. Meoneura krivosheinae 0 zerov,

1991 and the suspiciously similar Meoneura forcipata

the prominent cercus long

Sabrosky, 1959, also have both obvious large and setose

structures projecting from the epandrium, too. But these

structures are identified as surstyli, they are narrow basal-

ly and have dense setae restricted to their apical half (see

Ozerov 1991: 8, Figs. 4-5, Sabrosky 1959: 21, fig. 8).

There is no additional structure identified in the original

descriptions of both species that resembles the surstylus

of Meoneura issunboshii sp. n. Meoneura stepposa O ze-

rov, 1994 has a shorter, triangular surstylus with only one

seta apically and a longer epandrium (at least twice as long

as surstylus, see Ozerov 1994: 141). Meoneura lamella-

ta Collin, 1930 has a shorter lamella that is only about as

long as surstylus and a larger epandrium (see Collin 1930:

plate III, fig. 7). Meoneura perlamellata Hennig, 1937 has

a large setulose surstylus but no lamella and a longer epan-

drium (see Hennig 1937: 71, fig. 70).

Description. Male Length about 2.4 mm. Wing length =

2.1 mm. Head height = 0.5 mm. Head: black with only

anterior half of frons dark brown. Antenna black. Arista

without pubescence. Eye without ommatrichia. Ratio of

maximum eye length : maximum eye height is 1: 1. Ra-

tio of posteroventral margin of gena closest to eye mar-

gin : maximum eye height is 1 : 2 (= 0.5). Frons micro-

tomentose, frontal triangle shiny. Frontal triangle distinct,

reaching anteriorly less than 1/2 distance from anterior

ocellus to frontal margin. Face slightly microtomentose.

Carina narrow. Postcranium microtomentose. Prementum

about as long as labellum and only slightly wider. Palpus

brown, almost half as long as the haustellum. One pair of

distinct ocellar setae. Supralunular setae convergent. Four

Bonn zoological Bulletin 66 (2): 177-181

fronto-orbital setae, two anterior mesoclinate, two poste-

rior lateroclinate. Two vertical setae. Postocellar setae par-

allel. One strong vibrissal seta; two supravibrissal setae,

the ventral one distinctly smaller. Three strong genal se-

tae, the anterior strongest and upcurved.

Thorax: Scutum completely slightly microtomentose,

and covered with black setulae. Scutellum slightly micro-

tomentose. Pleurae dusted to subshiny. Scutum with on-

ly one long and two inconspicuous smaller dorsocentral

setae (two single setae broken in holotype). Two post-

pronotal setae; one praesutural seta; two notopleural se-

tae; one supraalar seta; two postalar setae; one praescutel-

lar seta; one apical and one lateral scutellar seta. One out-

standing seta and some smaller setulae at posterior mar-

gin of anepisternum. One dorsal seta and one ventral se-

ta on katepisternum. Wing completely covered with mi-

crotrichiae. Costa without obvious setae beyond radial vein

R,. Wing hyaline to light brown, veins light brown to white

yellow. Ry,, almost straight. Knob of halter whitish yel-

low, base of haltere brownish. Legs black to brown. Fore

femur with three strong posteroventral setae. Hind femur

apically with one strong anteroventral seta. Fore coxa with

one outstanding strong seta, middle coxa with two strong

setae, hind coxa with one outstanding seta. Hind metatar-

sus ventrally with scattered yellow golden hairs only. Ra-

tio of length metatarsus 2 : length tibia 2 is 0.7.

Abdomen: Tergites without obvious depressions or se-

tulae tufts. Abdominal pleura with scattered setae on seg-

ments 3-5. Segments 1-5 narrow, ratio of width of ter-

gite 3: length of tergite 3 = 2.7. Tergites 2-5 each with a

small lateral seta at the posterior margin. Tergite 5 with

three longer setae at the posterior margin. Sternite 5 rec-

tangular slightly longer than wide (Fig. 4). Sternite 5 with

two strong laterally curved setae. Midventral tergite 7 dis-

tinctly developed. Protandrium distinct, longer than short

epandrium and about 1/2 of length of tergite 5. Protandri-

um ventrally widely separated. Epandrium with two out-

standing long setae and a few smaller setae (Fig. 1). Ra-

tio of maximum length dorsally in the middle of epandri-

um (including cerci): maximum width of epandrium = 0.2.

Cerci indistinct, slightly projecting, and with a few minute

setulae only. Subepandrial sclerites fused with each oth-

er and fused with the narrow sclerotized hypoproct but the

structures can be distinguished due to a different sclero-

tisation. No setula recognized on subepandrial plate. Hy-

poproct not projecting laterally, with few strong, short se-

tae. No tooth on subepandrial plate. Surstylus as Figs. 1-2:

falcated, dorsobasally slightly concave, with an indistinct-

ly bulging base. Surstylus dorsally with four to five strong

setae, some minute setulae in front of these and one long,

medially directed setulae apically. Lamella large (Figs.

1-2) densely covered with long setae that are dorsally and

apically longer than ventrally. Surstylus and lamella dis-

tinctly fused basally (Figs. 1-2). Postgonite (Fig. 3): dis-

tinctly sclerotized, with a broad base, elongated, pointed,

©OZFMK

A new species of Meoneura Nitzsch from Japan 179

Figs 1-4. Meoneura issunboshii spec. nov. (drawn from holotype). 1. Postabdomen, left, lateral view. 2. Surstylus, left, lateral

view. 3. Postgonite, left, lateral view. 4. Sternites, ventral view: sternite 3 (left), sternite 4 (middle), sternite 5 (right). Abbrevia-

tions: ce - cercus, ep - epandrium, la - lamella, su - surstylus.

180 Jens-Hermann Stuke & Toshihiro Tago

Fig. 5.

apically curved into hook-like prolongation. Distiphallus

(Fig. 5) distinctly longer than epandrium, covered all over

with setulae and apically with a small sclerotized area.

Basiphallus sclerotized, long, narrow, distinctly widened

at base.

Description of female abdomen. Tip of female abdomen

identical to drawing of female abdomen of Meoneura ob-

scurella (Fallén, 1823) as figured by Sabrosky (1987: 911,

fig. 5). Abdominal pleura with dense setae on segments

4-5. Sternite 5 elongated, with two distinct setae on api-

cal half. Sternite 6 elongated, with four setae on apical half.

Sternite 5 almost square. Sternite 5 with one seta. Apical

half of segment 7 ventrally with four setae of which two

are always close together. Segment 8 at apical margin ven-

trally with two pairs of setae. Hypoproct almost square

with one seta. Fused cerci elongate, with several setae.

Bonn zoological Bulletin 66 (2): 177-181

Meoneura issunboshii spec. nov. (paratype, locus typicus, 11.xii.2011), distiphallus, lateral right view.

Etymology. The name issunboshii is derived from the

name of the Inch-High Samurai Issun-b Oshi (—T3HEM)

from a Japanese fairy tale. Issun-bOshi showed that a

minute man can be very successful.

Distribution. To date Meoneura issunboshii sp.n.is on-

ly known from the locus typicus, Mizumoto Park in Tokio,

Katsushika, Japan.

Examined material. Holotype &: (1) “Mizumoto, / Kat-

sushika word, / Tokyo, Japan /10.1.2010/T. Tago leg.”;

(2) “Holotypus / Meoneura issunboshii | spec. nov. S'/ det.

Stuke, 2017”. Male holotype is deposited in the Museum

fiir Naturkunde der Humboldt-Universitaét, Germany,

Berlin (ZMHB). Posterior part of abdomen dissected, mac-

erated and stored in glycerine in a microvial pinned be-

neath the specimen. The remainder of the specimen is

glued on a paper and in an excellent condition.

©ZFMK

A new species of Meoneura Nitzsch from Japan 181

Paratypes: 93'S 1029 with same data as holotype but

different collecting dates stored in the collection of the Os-

aka Museum of Natural History, Japan (OMNH) and in

the private collection of Jens-Hermann Stuke, Germany,

Leer (PJHS): 30S 1Q 10.1.2010 ( PJHS); 200% 299

10.1.2010 (OMNH); 10 299 11.xii.2011 (PJHS); 30S

11.xii.2011 (OMNH); 299 13.i.2013 (PJHS); 19

13.14.2013 (OMNH); ie 8.1.2011 (OMNH); 19 26.11.2012

(OMNH).

Remarks. Meoneura issunboshii sp. n. is found in the

Mizumoto Park at about 5 m above sea level. Adults are

recorded only in the winter months from November to Jan-

uary with temperatures of -3°C to -15°C. This might be

the reason why the species has been overlooked previous-

ly. The flies aggregate on the bark of sun exposed parts

of trees like Zelkova serrata (Thunb.) Makino and

Metasequoia glyptostroboides Hu & W.C. Cheng. There

they sunbathe, walk around, fly a short distance after dis-

turbance, and mate. The forest floor was covered with

woodchips. Although TT looked for, Meoneura issunboshii

sp. n. was not recorded at neighbouring parks or forests

in Tokyo.

Acknowledgements. a first version of the manuscript was kind-

ly checked by Irina Brake (Germany, Burgdorf). Hisashi Oishi

(Japan, Osaka) deposited the paratypes of Meoneura issunboshii

sp.n.in the collection of the Osaka Museum of Natural histo-

ry.

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