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Bonn

zoological

Bulletin

formerly: Bonner zoologische Beitrage

Volume 64

Issue 1

2015

An open access journal of organismal zoology, published by

Zoologisches Forschungsmuseum Alexander Koenig, Bonn

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Bonn zoological Bulletin

Editor-in-Chief

Fabian Herder, Zoologisches Forschungsmuseum Alexander

Koenig (ZFMK). Ichthyology Section, Adenauerallee 160,

53113 Bonn, Germany, tel. +49 228-9122-255,

fax: +49 228-9122-212; fherder@zfmk.de

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Thomas Wesener, ZFMK, tel. +49 228-9122^25,

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Dirk Ahrens, Insects: Coleoptera, ZFMK,

tel. +49 228-9122-286; d.ahrens@zfmk.de

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tel. +49 228-9122-250; w.boehme@zfmk.de

Jan Decher, Mammals, ZFMK, tel. +49 228-9122-262;

j.decher@zfmk.de

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

Aviv University, Tel Aviv 69978, Israel,

tel. +972-3-6409808; ndorchin@post.tau.ac.il

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Ralph Peters, Insects: Hymenoptera, ZFMK,

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Bradley Sinclair, Canadian National Collection of Insects,

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Greifswald, Gennany, tel. +49 3834-86^242, fax: +49

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erich.weber@uni-tuebingen.de

Bonn zoological Bulletin 64 ( 1 ): 1-15

SEP 1 5 2015

^/SRAR'ES^

July 2015

Wingless Paocryplorrhiniis (Coleoptera: Curculionidae)

rediscovered in Tanzania: synonymy, four new species

and a mtDNA phylogeography

VASILY V. GREBENNIKOV

Cauadian Food Inspection Agency. K.W. Neatby Bldg., 960 Carling Ave., Ottawa. ON KIA 0Y9, Canada.

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

Abstract. The weevil genus Paociyptorrhimis Voss, 1965 is revised. The nominal taxon Anchonidiitm hustachei Hot'f-

mann, 1965 is transfeiTed to Paociyptorrhimis as P. hustachei (Hoffmann, 1965) comb. n. The nominal taxa Anchonid-

ium hustachei Hoffmann, 1 965 and Paociyptorrhimis ohsitus Voss, 1 965 are considered conspecific and the name A. hiis-

tachei Hoffmann, 1965 is a senior subjective synonym of P. ohsitus Voss, 1965 syn. n. The genus, previously known for

a single species from the wet forests on geologically young volcanic mountains in northeastern Tanzania, is reported from

two other forest types: the old crystalline East Arc Mountains and the lowland forest. Four new species, all from Tanza-

nia, are described: P. atropos sp. n. (North and South Pare), P. clotho sp. n. (West Usambara), P. lununigeusis sp. n. (Mt.

Hanang) and P. lachesis sp. n. (Kimboza forest). A single specimen of an unnamed species is reported from East Usam-

bara. The mtDNA barcoding sequences are analysed phylogeographically. Even though the genus was not recovered as

a monophyletic clade in two of the molecular phylogenetic analyses, it is nevertheless hypothesized as being monophylet-

ic based on numerous similarities in adult moiphology, biology and based on the geographical proximity of all records.

The uncertain phylogenetic position of Paociyptorrhimis is brielJy discussed and the genus is transferred from Brachyc-

erinae: Erirhinini to Molytinae iucertae sedis.

Key words. COl, DNA barcoding, Eastem Arc Mountains, East Africa

Introduction

Taxonomic literature contains numerous poorly known

taxa that are mentioned only when described and that are

then seemingly forgotten. A significant, if not the predom-

inant number of genus- and, particularly, species-group

Coleoptera taxa from “exotic” localities (i.e. far from the

historical taxonomic centres of expertise located in Eu-

rope and elsewhere) can be classified as such. Once in-

troduced, the names are repeatedly mentioned in cata-

logues, while their true identity, taxonomic diversity and

distribution, monophyly and phylogenetic position, imma-

ture stages and habitat remain elusive. Such state of af-

fairs might continue indefinitely, unless the taxon sudden-

ly turns out to be of economic significance and gains im-

portance and attention. This was the case with^g/v7i« pla-

nipennis Fainnaire, a jewel beetle from the Asian Pacif-

ic Region. This was a taxonomically neglected and poor-

ly known species until it suddenly spread over most of

eastem North America and became an invasive ash killer

of cosmic proportions (Jendek & Grebennikov 2011).

Such examples fonn the negligible minority within poor-

ly known taxa, while the ovemhelming majority contin-

ues to drag along their ghostly existence decade after

decade. One such example among the weevils is the sub-

ject of the present paper.

The generic name Paociyptorrhimis was introduced a

half century ago by Eduard Voss (1965) for his new

species P. ohsitus from the geologically young volcanic

mountains in northeastern Tanzania. The holotype was

sampled from “humus” (=leaf litter) in the forest on the

eastem slopes of Mount Kilimanjaro at 2700 m, while 12

paratypes came either from the vicinities of Marangu vil-

lage on the south-eastern slope of the same mountain, or

from nearby Mount Mem, Mount Oldeani and the

Ngorongoro highlands (Fig. 1 1). These localities coiTe-

spond closely to the geologically young (<2 Ma) volcanic

highlands of the so-called Ngorongoro-Kilimanjaro Vol-

canic Belt (NKVB, Nonnotte et al. 2008). The type series

was collected by the Belgian entomologists Pieme

Basilewsky and Narcisse Leleup of the Royal Museum for

Central Africa in Tervuren during their epic 1 957 journey

to sample the East African biota. All weevils from their

samples were given to Voss for study, who delivered the

results in two parts published in 1962 and 1965, respec-

tively. The latter contained the description of the new

genus, accompanied by an adult habitus drawing in dor-

sal view and the head and prothorax in lateral view (Voss

1965: Fig. 7). Until present, this was the only time the

genus has been mentioned in original literature (i.e. ex-

cluding catalogs).

Received: 28.07.2014

Accepted: 16.04.2015

Corresponding editor: D. Ahrens

9

Vasily V. Grebennikov

This project was triggered by the necessity to identify

Paoayptoirhinus specimen #3081 included as an out-

group representative in the phylogenetic analysis of new

wingless species of Notaris Germar from southwest Chi-

na (Grebennikov & Kolov, unpublished). Comparison of

the specimen’s moiphology with that of the other seem-

ingly congeneric and newly sampled specimens suggest-

ed existence of other species, aside from the type species.

This was further supplemented by the differences in their

mtDNA barcodes. Another unexpected discovery was that

the type species had already been named and assigned to

a different genus (Hoffmann 1965) about one month be-

fore the genus Paoayptoirhimis was fomially established

by Voss. These discoveries necessitated the completion of

the present paper, which had the following main goals: ( 1 )

to document the course of the work highlighted above; (2)

to revise the genus taxonomically; and (3) to propose a

phylogeographic scenario for the low-dispersing Paoayp-

toirhinus species biologically linked to highly isolated

Afrotropical wet forests of contrasting origin and age.

Material AND Methods

This study is based on examination of 18 adults of

Paoayptorrhiinis sampled from Tanzania and assigned to

six species {P. Inistashei, four fonnally named as new, and

one consciously left unnamed as Paoayptoirhinus sp.

#3081). Sixteen specimens were successfully sequenced

for the Cytochrome Oxidase subunit 1 (5-prime end, i.e.

the Barcoding fragment). Parts of the historical type se-

ries pertaining to two synonymous names of the type

species were examined. Fifteen other specimens represent-

ing various species of mainly Molytinae and Brachyceri-

nae were added to the DNA-based phylogenetic analysis

as an extended outgroup. All 3 1 specimens used for DNA

extraction have at least one unique identifier label pinned

underneath with the code CNCCOLVGOOOOXXXX

(shortened to #XXXX when specimens are cited in the text

below). Some specimens with more than one unique iden-

tifier (for example a specimen with labels #5231, #7270,

#7271) required repeated sequencing attempts. GenBank

accession numbers for 16 Paoctyptorrhymis sequences are

given in the species descriptions; those of the 1 5 outgroup

representatives are: HM4 17724, HQ986868, HQ987003,

HQ987029, KJ445698, K.1445708, K.I672224, KJ672228,

K.I672239, KJ672247, KJ672251, KJ672255, KJ672276,

KJ841714 and KJ841726. A list of all species, specimen

images, georeferences, primers, sequences, original chro-

matograms and other relevant data for all 3 1 sequenced

specimens can be found online in the publicly accessible

dataset Paoayptoirhimis" on the Barcode of Life Data-

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

All methods used in the present work are the same as

those explained in detail for a similarly structured work

on the genus Niphadomimiis (Grebennikov 2015). This in-

cludes specimen collecting and dissection, diagnostic

species descriptions, process of DNA extraction, as well

as the logic, software and implementation of phylogenet-

ic analyses using Maximum Likelihood (ML) and

Bayesian analysis as implemented in MEGA6 (Tamura et

al. 2013) and BEAST v 1.8.0 (Drummond et al. 2012), re-

spectively. The matrix used for both analyses had a length

of 658 base pairs (bp) and contained 31 sequences. The

resulting topologies were visualized in FigTree vl .4 (Ram-

baut 2014). The ML tree was rooted with both represen-

tatives of Anthribidae (Choraginae genus Styphlochora-

gus Frieser and Anthribinae genus Disphaewmi Jordan).

The BEAST analysis was performed without rooting and

without enforcing monophyly of any group. In the absence

of relevant fossil data, time calibration was made by as-

suming strict linear molecular clock and the rate of 0.0 1 8%

of nucleotide substitution per million years (Papadopoulou

et al. 2010). Tracer 1.6 (Rambaut et al., 2014) was used

to graphically determine stationarity and to check conver-

gence of runs.

Labels of holotypes (HT) and paratypes (PT) of the new-

ly described species are cited in quotation marks. Letters

[p] and [h] in square brackets denote printed and hand-

written text on historical labels, respectively, while back

slash \ separates text on individual labels. Weevil taxon-

omy follows Alonso-Zarazaga & Lyal (1999), with sub-

sequent modifications by Oberprieler (2014) and Lyal

(2014). Distribution maps are generated using the on-line

SimpleMappr tool (Shorthouse 2010). The chronostrati-

graphic timing is that of Cohen et al. (2013). Terminolo-

gy for male genitalia follows Wanat (2007). The length

of the body was measured in dorsal aspect from the ely-

tral apex to the anterior edge of the pronotum (standard

length).

The following abbreviations were used to indicate the Mu-

seum depositories (followed by the name of the curator):

BSUM Department of Biological Sciences,

the University of Memphis, Memphis, USA (D.

McKenna);

CNC Canadian National Collection of Insects, Arach-

nids and Nematodes, Ottawa, Canada (P.

Bouchard);

MRAC Musee Royal de FAfrique Centrale, Tervuren,

Belgium (M. De Meyer);

NMW Naturhistorisches Museum, Wien, Austria (H.

Schillhammer).

Bonn zoological Bulletin 64 ( 1 ): 1-15

®ZFMK

Paociyptorrliiniis weevils from Tanzania

3

Taxonomy

Paocryptorrhinus Voss, 1 965

Type species. Paocryptorrhinus obsitus Voss, 1965: 361

fixed by original designation.

Diagnosis. Adult beetles of the genus Paociyptorrliiniis

are rather distinctive in their appearance and can be recog-

nised relatively easily. The presence of postcoxal flanges

on the prostemum (Figs 71, J; Figs 9A-G) forming later-

al walls along the canal for accommodating the rostrum

in repose immediately separates Paociyptorrliiniis from

the majority of weevils not only in East Africa but also

worldwide (except Molytinae: Aedemonini, a few

Sophrorhinini and a few other stray genera). From the lat-

ter, Paociyptorrliiniis can be recognized by having any

two of the following three characters: (A.) eyes much re-

duced in size and seemingly consisting of a single, rela-

tively large ommatidum (Fig. 91) subequal in diameter to

that of any funicular antennomere; (B.) the dorsal outline

of the frons-to-rostrum junction in lateral view not form-

ing an even curve, but marked by a depression (Fig. 3C),

and (C.) elytra flattened, nearly parallel-sided most of their

length (Figs 1 A, C), and not shorter than 1 .55 times their

maximum width. Adults of two other weevil genera ( Oc7<7-

diiis Schoenherr and Isorhainpluis Faust) with a rostral

canal were sifted together with Paociyptorrliiniis. Those,

however, have eyes with several ommatidia and vaulted

elytra which are either rounded (Ocladiiis) or posteriorly

tapered (Isorhamphiis) in dorsal view.

Description. Integument uniformly dark-brown, often

covered by waxy, white-brown coating obscuring colour

and surface texture; standard length 2.31-3.96 mm; body

notably elongate. Head capsule almost completely retract-

ed into prothorax and not visible from above when ros-

trum in repose or directed ventrally. Eyes located latero-

ventrally, small, flat, seemingly consisting of single om-

matidium not exceeding in diameter maximum diameter

of scapus (Fig. 91). Rostral attachment to head capsule

abruptly constricted laterally (in dorsal view) to half ros-

tral width (Fig. 9A) and gently depressed dorsally in lat-

eral view (Fig. 91). Rostrum in lateral view almost straight;

in dorsal view almost parallel-sided and slightly and even-

ly constricted at middle. Antennal attachments in apical

30-40% of rostral length; funicle with seven anten-

nomeres; compact club consisting of three antennomeres.

Pronotum weakly to markedly constricted laterally in an-

terior quarter; widest at middle; prostemum with deep ros-

tral canal delimited laterally by lamellae anterior and pos-

terior to procoxae (Figs 9A-G). Pro- and mesofemora

without ventral tooth; hind femur with or without. Tibiae

mucronate (Fig. 9J). Elytra parallel-sided; elytron with 10

striae each marked by single line of setiferous punctures;

striae 7 and 8 short, not reaching base (Fig. 9J) or apex

of elytra. Hind wings absent. Metaventrite fonning ven-

trally extended posterior wall of rostral canal (Fig. 9H);

metepisternum and metaventrite fused without trace of

metepisternal suture (Figs 7I-J, 9J); sclerolepidia absent

(Fig. 9J). Male tergite 7 (Fig. 21) without deep notch at

posterior margin; stemite 8 (Fig. 2K) entire, not separat-

ed into hemistemites. Aedeagus weakly sclerotized, dor-

sally membranous; endophallus with weak sclerotization

(Figs 2E-G).

Biology and distribution. All known Paociyptorrliiniis

specimens were collected by sifting wet leaf litter on the

floor of wet primary Afrotropical forests. Host plants and

larvae of this genus are unknown, together with all other

more specific biological characteristics of these cryptic

and rarely encountered beetles. The known distribution of

Paociyptorrliiniis is relatively small and limited to 12 lo-

calities in northeastern Tanzania (Fig. 1 1 ) at altitudes be-

tween 288 and 2700 meters.

Key to species of Paocryptorrhinus

1 . Elytral stria 1 or 2 subequal in width to any of striae

4, 5 or 6 (Figs 1 A, C; 6A, C); prosternum with post-

coxal flanges smaller, postcoxal section of rostral

canal half as deep as anterior section (Fig. 9A, B);

pronotum with fine and numerous punctures (Figs

1A,C,2A,C 2

- Elytral stria 1 or 2 markedly narrower than any of stri-

ae 4, 5 or 6 (Fig. 3A); prostemum with postcoxal

flanges greater, post- and antecoxal sections of ros-

tral canal subequal in depth (Fig. 9C-G); pronotum

with large and relatively few punctures (Figs 3A, C,

4A,C, 7A, C) 3

2. Smaller, body length 2.31-3.05 mm (Figs lA-D; 2)

P. hustachei comb. n.

- Greater, body length 3.75-3.92 mm (Fig. 6)

R hanangensis sp. n.

3 Ridges on interstriae 2, 4 and 6 markedly higher and

longer than those on interstriae 1,3,5 (Figs 3, 4A-D);

lateral outline of elytra in dorsal view formed by

raised ridge on interstria 6 (Figs 3 A, 4A) 4

- Ridges on interstriae 1 to 6 subequal in height and

length (Fig. 5); lateral outline of elytra in dorsal view

fonned by raised ridge on interstria 5 (Figs 5A, 7A,

8A) 5

4 Elytral intervals 3 and 5 without raised ridges

P. atropos sp. n. (North Pare)

- Elytral intervals 3 and 5 with raised ridges in poste-

rior half P. atropos sp. n. (South Pare)

Bonn zoological Bulletin 64 (I ): 1-15

®ZFMK

4

Vasily V. Grebennikov

Mt.Meru W-Hang

, Tanganyika

Ig.H. Franz

JLuf^htch^ viA

A, Hoffmann del

1 mm

Fig. 1. Paoayptorrliiinis Intstachei comb, n., A-B: holotype of hustachei, unsexed, with three original labels; C-D: paratype of

ohsitiis, unsexed.

5 Hind femur with large ventral tooth (Fig. 5C)

P. clotho sp. n.

- Hind femur without ventral tooth 6

6 Combined maximal width of elytral striae 4 and 5

subequal to that of striae 1 to 3 (Fig. 8A, C)

Paocryptorrhinus sp. #3081

- Combined maximal width of elytral striae 4 and 5 not

more than 80% of that of striae 1 to 3 (Fig. 7A, C)

P. lachesis sp. n.

Paociyptoirhimis hustachei (Hoffmann, 1965) comb. n.

Figs f, 2, 9A, I, J

Anchouidium hustachei Hoffmann, 1965: 397-398.

Type locality. Tanganyika, Mount Meru [Tanzania].

Type specimens. HT (Figs 1 A-B) stated as male, not dis-

sected (NMW): “Mt. Meru W-Hang Tanganyika Ig. H.

Franz [p] \TYPE [p] [red label] \ Anchonidium hustachei

m. [male sign] [h] A. Hoffmann det [p]”. 2 PT (NMW);

not studied (see Remarks on the status of a specimen la-

belled as PT). Described from three specimens.

Paociyptoirhimis obsitiis Voss, 1965: 361-362, Abb. 7,

syn. n.

Type locality. Mt. Kilimanjaro, Versant Est, 2700 m, for.

mont., humus [Tanzania].

Type specimens. HT (MRAC): not studied. 1 PT

(MRAC) (Figs IC-D): “PARATYPUS [p] [red label] \

etrepage sous Hagenia [p] [blue label] \ l.R.S.A.C. - MUS.

CONGO Kilimanjaro, Marangu 2400 m. 20-11-1956 J. et

N. Eeleup [p] \ R. DET. 7680 [p] A. [h]“; 1 PT (MRAC):

“PARATYPUS [p] [red label] \ foret mont., humus [p]

[blue label] \ COLL. MUS. CONGO Tanganyika Ten: Mt.

Meru, Olkokola, versant N.O. 2600 m. 8-VII-1957 [p] \

R. DET. 7680 [p] A. [h]”; 1 PT (MRAC): “PARATYPUS

[p] [red label] \ etrepage sous foret claire [p] [blue label]

\ COLL. MUS. CONGO Tanganyika Terr. Mt. Meru,

Olkokola, versant N.O. 2500 m. 8-VII-1957 [p] \ R. DET.

7680 [p] A. [h]“; all three paratypes have additional la-

bel: “Mission Zoolog. l.R.S.A.C. en Afrique orientale (P.

Basilewsky et N. Eeleup) [p]”. Described from 13 spec-

imens.

Diagnosis. This species is recognized by the imifonnly

shaped six striae and interstria on the elytral disk in com-

bination with its relatively small body length (not greater

than 3.05 mm).

Description. Holotype, unsexed specimen (Figs lA-B).

Not sequenced for DNA barcode. Length: 2.94 mm.

Smaller prostemal postcoxal flanges fonning rostral canal

twice shallower than that anterior of procoxae; raised ely-

tral interstriae 1 to 6 reaching elytral base, subequal in

height; lateral elytral outline in dorsal view fonned by

ridge on interstria 6; elytral striae 4 and 5 not markedly

wider than others; hind femur without ventral tooth.

Bonn zoological Bulletin 64 ( I ): 1-15

®ZFMK

Paocnptorrilimis weevils from Tanzania

5

Interstria 6

interstria 5

Interstria 6

interstria 5

interstria 6

endophallus

Fig. 2. Paoayptorrhiniis Iwstacliei comb, n., #4893 from Mt. Kilimanjaro, male. A-D: habitus; E-G: aedeagus & legmen, dor-

sal (E), ventral (F) and lateral (G); H: proventriculus; 1: tergite 7; J: tergite 8; K: sternite 8; L: sternite 9.

Intraspecific variation. GenBank accessions: KJ841713,

KJ84 1717, K.I84 1719, KJ84 1 722-KJ84 1 724, KJ84 1 728.

Length: 2.31-3.05 mm.

Additional material examined. 12 exx in total. 5 exx

(NMW): I ex, “Mt. Monduli Tanganyika Ig. H. Franz [p]

\ PARATYPE [p] [rad label] / Anchonidiiim hustachei m.

[male sign] [h] A. Hoffmann del [p]”; 4 exx, “Gebirgs-

wald oc. Marangu Ig. H. Franz [p] \ Kilimanjaro Tanganyi-

ka [p]“. 7 exx (CNC): 1 ex, #4801 “TANZANIA, Mt. Kil-

imanjaro, Marangu route, S3. 1 8424° E37.5 1 344°, 2668 m.

25.xi.20 12, siftOl, V. Grebennikov”; 2 exx, #4844, #4845

“TANZANIA, Mt. Kilimanjaro, Marangu route,

83.21470° E37.5I822°, 2198 m, 28.xi.20 1 2, sift05, V.

Grebennikov”; I ex, #4864, “TANZANIA, Mt. Kiliman-

jaro, Marangu route, 83.20141° E37.51903°, 2370m.

6.xii.2012, sift09, V. Grebennikov”; 2 exx, #4869, #4870

“TANZANIA, Mt. Kilimanjaro, Machame route,

83.14954° E37.24532°, 2124 m, 7.xii.2012, siftll, V.

Grebennikov”; 1 ex, #4893 “TANZANIA, Mt. Kiliman-

jaro, Umbwe route, 83.16197° E37.28464°, 1993 m,

8.xii.2012, sift 13, V. Grebennikov”.

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Fig. 3. Paociyptorrhiniis atropos sp. n., #127 \ from North Pare, female. A-D: habitus.

Distribution. This species is known from five isolated wet

forests on volcanic mountains of the geologically young

NKVB (Fig. 11) in northeastern Tanzania. Elevation:

1993-2700 m.

Remarks. The type specimens of both nominal histori-

cal names pertaining to P. bustachei are similar to each

other (Fig. 1 ), while their type localities are about 60-70

km apart (Fig. 11). Each name was proposed by two au-

thors, respectively, unaware of each other and published

a month apart. These considerations suggest that both

species names are subjective synonymous. Floffmann’s

(1965) paper came out in the issue of the journal published

on May 26, 1965 (endleaf), while the volume containing

Voss (1965) paper was printed in June 1965 (verso of ti-

tle page). Consequently, the name introduced by Hoffman

is a senior subjective synonym of Voss’ name.

Hoffmann ( 1965: 398) stated that he had studied three

bustachei specimens: two males from Mount Meru (one

of them designated by him as the “type” (=holotype, ICZN

Article 73. 1 . 1 , Figs 1 A-B) and a female from Machame,

Mount Kilimanjaro. The series of this species received on

loan from NMW contained six conspecific specimens all

collected by Herbert Franz, two of them with red labels

“TYPE” and “PARATYPE”, respectively. The fonuer

specimen from Mount Meru is considered herein as the

holotype (Figs 1 A-B). The latter is labeled as originating

from Mount Monduli (Fig. 1 1 ), which is a separate, thin-

ly forested and much smaller volcanic cone about 30 km

West of Mount Meru, which itself is situated 60-70 km

West of Mount Kilimanjaro (Fig. 1 1 ). The specimen with

the paratype label does not agree geographically with any

of the two paratypes mentioned in the original description

and, therefore, its paratype status is refuted.

Paocryptorrhinus atropos sp. n.

Figs 3, 4, 9C, D, H

Diagnosis. This species is recognized by the unique

arrangement of elytral interstriae, of which 1 , 3 and 5 are

absent in at least the basal half of the elytra, while inter-

striae 2, 4 and 6 are markedly enlarged along their entire

length.

Description. Holotype, male, (Fig. 5). GenBank acces-

sion: KJ841721. Length: 4.29 mm. Greater prostemal

postcoxal flanges fonning rostral canal subequal in depth

to that anterior of procoxae; elytral interstriae 1 , 3 and 5

without raised ridged entirely (Figs 3A-C, North Pare pop-

ulation) or in anterior half only (Figs 4A-C, South Pare

population); raised ridges on interstriae 2, 4 and 6 marked-

ly enlarged; lateral elytral outline in dorsal view fonned

by ridge on interstria 6; elytral striae 4 and 5 not marked-

ly wider than others; hind femora without ventral tooth.

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Fig. 4. Paoayptonhinus atropos sp. n., holotype, #5438 from South Pare, male. A-D: habitus; E-G: aedeagus and tegmen, dor-

sal (E), ventral (F) and lateral (G); H; stemite 9.

Intraspecific variation. GenBank accession: KJ84 1718.

Length: 3.73^.34 mm.

Material examined. HT (Fig. 4) male (CNC): #5488 (=

#5439), “TANZANIA, South Pare Mts., Chome For.,

84.27145° £37.92347°, 2072m, 4.i.2013, sift40,

V.Grebennikov”. PT (CNC, BSUM): 1 ex, #5436

(=#5467), same data as the holotype; 3 exx, #5231 (=

#7270, = #7271 ), #5232 (= #7272, = #7273) and one not

numbered, “TANZANIA, North Pare Mts., Kindoroko

For., 83.74313° £37.65022°, 1987m, 23.xii.2012, sift25,

V.Grebennikov”.

Distribution. This species is known from both South and

North Pares in northeastern Tanzania. £levation:

1987-2072 m.

Etymology. The species epithet is the Latinized Greek

mythical name of Atropos, one of three Morai (= Fates),

who cut the thread of life; noun in apposition.

Remarks. Three attempts to amplify DNA from each of

two North Pare specimens failed (hence three unique iden-

tifiers on each specimen) and, therefore, they are not rep-

resented on the phylogenetic tree (Fig. 10).

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Fig. 5. Paoayptorrhiniis clotlw sp. n., holotype, #5476 from West Usambara, female. A-D: habitus.

Paoayptoirhiniis clotho sp. n.

Figs 5, 9E

Diagnosis. This species is unique by having a markedly

developed ventral tooth on each hind femur (Fig. 5C).

Description. Flolotype, female (Figs 5, 9E). GenBank ac-

cession: KJ841727. Length: 4.13 mm. Greater prosternal

postcoxal flanges forming rostral canal subequal in depth

to that anterior of procoxae; raised elytral interstriae 1 to

6 reaching elytral base, subequal in height; lateral elytral

outline in dorsal view formed by ridge on interstria 5; ely-

tral striae 4 and 5 not markedly wider than others; hind

femur with ventral tooth (Fig. 5C).

Intraspecific variation. GenBank accessions: KJ841712,

KJ84I716. Length: 3.68^.13 mm.

Material examined. HT (Figs 5, 9E) female (CNC):

#5476, “TANZANIA, W. Usambara Mts., Mazumbai For.,

84.81054° E38.49858°, 1618m, ll.i.2013, sift43,

V. Grebennikov”. PT (CNC): 1 ex, #5477, same data as the

holotype; 1 female, #5478, “TANZANIA, W. Usambara

Mts., Mazumbai For., 84.82634° £38.49690°, 1860m,

12.1.2013, sift46, V.Grebennikov”.

Distribution. This species is known only from the

Mazumbai forest in West Usambara, northeastern Tanza-

nia. Elevation: 1618-1680 m.

Etymology. The species epithet is the Latinized Greek

mythical name of Clotho, one of three Moral (= Fates),

who spun the thread of life; noun in apposition.

Paocryptorrhiniis hanansensis sp. n.

Figs 6, 9B

Diagnosis. This species is recognized by the unifomily

shaped six elytral discal striae and interstria, in combina-

tion with its relatively greater body length (not smaller

than 3.75 mm).

Description. Flolotype, female (Figs 6, 9B). GenBank ac-

cession: KJ841725. Length: 3.75 mm. 8maller prosternal

postcoxal flanges forming rostral canal twice shallower

than that anterior of procoxae; raised elytral interstriae 1

to 6 reaching elytral base, subequal in height; lateral ely-

tral outline in dorsal view formed by ridge on interstria

6; elytral striae 4 and 5 not markedly wider than others;

hind femur without ventral tooth.

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Fig. 6. Paoayptorrhinus hamingemis sp. n., holotype, #5086 from Mt. Hanang, female. A-D: habitus.

Intraspecific variation. GenBank accessions: KJ841 720.

Length: 3.75-3.92 mm.

Material examined. HT (Figs 6, 9B) female (CNC):

#5086, “TANZANIA, Mt. Hanang, NE slope, S4.41621°

E35.4028 1 2652m, 1 3.xii.20 1 2, sift 1 5, V.Grebennikov”.

PT(CNC): I female, #5 147 (=#51 48 (“TANZANIA, Mt.

Hanang, NE slope, 84.43077° E35.41593°, 2283m,

15.xii.2012, sift20, V.Grebennikov”.

Distribution. This species is known only from Mount

Hanang, northeastern Tanzania. Elevation: 2283-2652 m.

Etymology. The species epithet is an adjective derived

from the name of its type locality, Mount Hanang in north-

eastern Tanzania.

Paocryptorrhinus lachesis sp. n.

Figs 7, 9F.

Diagnosis. This species is most easily recognized by the

deep and abrupt lateral constriction of the pronotum in the

anterior quarter (Fig. 7A).

Description. Holotype, male (Fig. 7, 9F). GenBank ac-

cession: KJ841715. Length: 3.96 mm. Greater prostemal

postcoxal flanges forming rostral canal subequal in depth

to that anterior of procoxae; raised elytral interstriae 1 to

6 reaching elytral base, subequal in height; lateral elytral

outline in dorsal view fomied by ridge on interstria 5; ely-

tral striae 4 and 5 not markedly wider than others; hind

femur without ventral tooth.

Material examined. HT (Fig. 7) male (CNC): #3603,

“TANZANIA, Kimboza forest, 7°0r20”S 37°48’I3”E,

8. i. 2012, 288m, sift. 24, V.Grebennikov”.

Distribution. This species is known only from the holo-

type found in the Kimboza forest. Elevation: 288 m.

Etymology. The species epithet is the Latinized Greek

mythical name of Lachesis, one of three Morai (=Fates),

who measured the thread of life; noun in apposition.

Remarks. External moiphological peculiarity of the sin-

gle known male specimen (Figs 7, 9F), along with its dis-

tinct DNA sequence (though relatively short and consist-

ing of only 407 nt) and a large geographical gap separat-

ing it from the rest of the genus (Fig. II) triggered its

recognition as a fonnally named species (unlike Paociyp-

torrliiinis sp. #3081; see below).

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Vasily V. Grebennikov

Fig. 7. Paooyptorrhimis lachesis sp. n., holotype, #3603 from Kimboza forest, male. A-D: habitus; E-H: aedeagus and tegmen,

dorsal (E), ventro-lateral (F), lateral (G) and ventral (H); I-J: prostemum, meso- and metaventrum showing rostral canal, ventral

(I) and latero-ventral (J).

interstria 5

1 mm

Interstria 5

E

0 2 mm

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Fig. 8. Paociyptorrhinus sp. #3801, #3801 from East Usambara, male. A-D: habitus; E-G: aedeagus and tegmen, dorsal (E),

ventral (F), lateral (G); H: tergite 8 and sternite 9, ventral.

Paocryptorrhinus sp. #3081

Figs 8, 9G

Diagnosis. This species is unique by having markedly

widened elytral striae 4 and 5 (Fig. 8C).

Description. Male (Fig. 8, 9G). GenBank accession:

KJ672257. Length: 3.10 mm. Greater prostemal postcox-

al flanges fomiing rostral canal subequal in depth to that

anterior of procoxae; raised elytral interstriae 1 to 6 reach-

ing elytral base, subequal in height; lateral elytral outline

in dorsal view formed by ridge on interstria 5; elytral stri-

ae 4 and 5 markedly wider than others; hind femur with-

out ventral tooth.

Material examined. Male (CNC); #308 1 , “TANZANIA,

E Usambara Mts., Amani NR, 5°05'56”S 38°37’18”E,

16.xii.201 1, 970m, sift. 07, V.Grebennikov”.

Distribution. This species is known from a single spec-

imen found in East Usambara, northeastern Tanzania. El-

evation: 970 m.

Remarks, Unlike P. lachesis sp. n. also known from a

male singleton, this potentially new species is conscious-

ly left unnamed, mainly due to uncertainties suggested by

its geographical proximity to the majority of other species

of Paociyptorrhinus.

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Vasily V. Grebennikov

Fig. 9. Paocnptoirliiniis spp., ventral view. (A-G): prothorax in ventral view showing prostemal canal and variously devel-

oped postcoxal flanges forming its lateral walls; (H): meso- and metaventrum with two basal abdominal ventrites in ventral view

showing rostral canal and its posterior vertical wall fomied by metaventmm; (1): head in left lateral view showing small eye; (J):

left elytron and left metathorax showing lack of metepistemal suture and sclerolepidia. (A): P. hustachei comb, n., #4893 from Mt.

Kilimanjaro, male; (B): P. haiiongensis sp. n.. #5086 from Mt. Hanang, female; (C): P. atropos sp. n., #7271 from North Pare, fe-

male; (D, H): F! atropos sp. n., holotype, #5438 from South Pare, male; (E): P. clotho sp. n., #5476 from West Usambara, female;

(F) P. lachesis sp. n., holotype, #3603 from Kimboza Forest, male; (G): Paoayptorrhimis sp. #3081, #3081 from East Usambara,

male; (I, J): P. hustachei, #4801 from Mt. Kilimanjaro, unsexed. Scale: 0.2 mm.

Results of DNA analyses

The model search in MEGA6 detected the generalized

time-reversible model with gamma distributed rate hetero-

geneity and inferred proportion of invariable sites

(GTR+G+I) as the best fit. The best ML tree (not shown

and with Paociyptorrhiniis branching pattern identical to

that in Fig. 10) had the highest log likelihood of

-7223.19. The representatives of Paoayptorrhimis were

not monophyletic being separated in two clades, P. hiis-

tachei comb. n. plus P. hauangensis sp. n. versus all oth-

er species, respectively (as in Fig. 10). The BEAST analy-

sis (Fig. 10) recovered a topology identical to that of the

ML analysis, although the exact placement of both ingroup

clades among the members of the extended outgroup was

different. The time tree hypothesis from BEAST revealed

for the Paoayptorrhinus evolution an interval from 14. 16

MY to present (Fig. 10).

Discussion

While the present report greatly increases the amount of

information available on Paoayptorrhimis weevils, all at-

tempts to inteipret the results are still much hindered by

the acute shortage of data and concomitant phylogenetic

resolution. The results, therefore, should be taken with

caution.

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Paocnptorrhiinis weevils from Tanzania

13

Miocene

Plio- Pleistocene

30 MY

20 MY

10 MY

present

Fig. 10. Relevant part of an ultrametric time tree of Paociyplorrhiinis obtained by using BEAST software calibrated at a rate

0.018 substitutions/site/MY. Note that the genus was not recovered as monophyletic. Values above nodes and on scale below are

million years before present (MY); values below nodes are bootstrap support for identical clades recovered in ML analysis. Node

bars represent 95% confidence interval of the age estimate.

Monophyly of Paociyptorr/iiiws

Even though monophyly of Paooyptorrhinus was reject-

ed in both DNA analyses (Fig. 10), the genus is likely

monophyletic. This assumption is based on three inde-

pendent sources of evidence. The seemingly strongest sup-

port comes from rather unifonn adult moiphological char-

acters: the similar unique arrangement of sclerites form-

ing the rostral canal (Figs 7A-B, 9A-H) potentially rep-

resents a complex synapomorphy. The small adult eyes

formed by a seemingly single and relatively flat omma-

tidium might be another synapomoiphy. The deep retrac-

tion of the head capsule into pronotum (Fig. 1 ), lateral con-

striction of the pronotum in apical fourth, and the pecu-

liar sclerotization of the endophallus (Fig. 2E) might be

other synapomoiphies. Thus, a broadly designed moiphol-

ogy-based phylogenetic analysis, presently outside of the

scope of this study, has the potential of recovering the

genus as a monophyletic clade. Such a hypothesis could

be coherent with two other lines of evidence: (a) the sim-

ilar biological preferences (=inhabitants of wet leaf litter

in primary forests), and (b) the restricted distribution of

the genus, with all species limited to northeastern Tanza-

nia (Fig. 1 1 ).

Phylogenetic position of Pciociyptorrhinus

The sister group of Paoaypton-hiniis and its phylogenet-

ic position among other weevils are presently entirely un-

known. The genus was described as a member of “Notari-

ni” and was retained there ever since (although under a

different tribe name of Brachycerinae: Erirhinini; see

Alonso-Zarazaga & Eyal 1999). This assignment is rather

an historical artefact than an evidence-based hypothesis.

When described by Voss, Paoayptorrhimis was only com-

pared with the genus Pachytychius Jekel, which then was

kept in Brachycerinae: Erirhinini. The latter genus, how-

ever, was transfeiTed from Erirhinini to Curculioninae, and

retained either in Storeini (Alonso-Zarazaga & Eyal 1999)

or incertae sedis (Caldara et al. 2014). Consequently, the

original assignment of Paociyptoirliiniis to Erirhinini via

its similarity with Pacliyp’diius was the only reason why

the former remained in Brachycerinae: Erirhinini (al-

though with some reseiwations; see Obeiprieler 2014).

The cun ent assignment of Paoayptorrhimis to Erirhini-

ni is not corroborated in the present analysis (Fig. 10),

since Paoayptorrhimis did not fonn a clade with Notaiis,

the latter being the type genus of Erirhinini. This is not

surprising, since adults of Paoayptorrhimis neither dis-

play the pedotectal-type of male genitalia characteristic

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Vasily V. Grebennikov

NKVB

P. huslachei

P. atropos sp. n.

Zanzibar

lake Victoria

1 . Mt. Meru “

2. Mt. Kilimanjaro, Marangu B

3. Mt. Kilimanjaro, Machame & Umbwe =

4. Mt, Monduli E

5. Mt. Oldeani -i

6. Ngorongoro 5

7 Mt. Hanang S

Kenya

Tanzania

R hanangensis sp. n.

P. clotho sp. n. I 'Jif

Paocryplorrhinus sp. #3081Jitf

P lachesis sp. n.

Fig. 11. Known distribution of Paociyptorrhinus weevils. NKVB: Ngorongoro-Kilimanjaro Volcanic Belt.

to Notaris and most Brachycerinae, nor do they have a lat-

eral setiferous groove on either side of the rostral apex

characteristic for most of Erirhinini (Obeiprieler 2014,

Figs 3.7.1.2D-G). The presence of prosternal postcoxal

flanges forming lateral walls of the rostral canal seeming-

ly suggests affinities to Molytinae; Aedemonini, a group

of 35 genera from most of the wann and temperate regions

of the World (except Europe, New Zealand and both

Americas; Lyal 2014). Efnlike adults of Aedemonini, those

of Paocryptorrhinus have a rostral canal with a marked-

ly developed transverse posterior metaventral wall (Figs

71, J) and the posterior margin of the male tergite 7 is on-

ly weakly emarginated (Fig. 21). Alternative states of these

two characters were suggested as synapomoiphies for

Aedemonini (Fyal 2014) and, therefore, this tribe cannot

accommodate Paoayptorrhiims. Any attempt to place the

genus in the relatively weakly understood phylogenetic

framework of weevils is at present doomed by the lack of

adequate data and pre-existing hypotheses. With all this

in mind, the most rational temporal action is the herein

proposed transfer of the genus from Brachycerinae:

Erirhininae to Molytinae incertae sedis.

Phylogeography of Paociyptorrhinus

Adults of Paociyptorrhinus are most likely low-dispers-

ing and humidity-dependant, specialized litter inhabitants,

as suggested by sampling circumstances, flightlessness,

and eye reduction. As presently known, all species of

Paociyptorrhinus are strictly allopatric (Fig. 1 1 ) and all

are found within a coherent geographical area in north-

eastern Tanzania (Fig. 1 1 ). These observations appear to

favour the simplified model of vicariant speciation for

low-dispersing habitat-dependent organisms (but see be-

low on P. hustachei comb. n. and P. hanangensis sp. n.).

Remarkably, the genus Paociyptorrhinus has not been de-

tected by us in other seemingly suitable and similarly ex-

tensively sampled parts of the Eastern Arc Mountains in

Tanzania, such as Ngum, Kaguru, Rubeho, Udzungwa and

Uluguru. The lack of Paociyptorrhinus records from

Uluguru is particularly puzzling, since the forest edge is

only 10 km west of the minute (about 3 km square) Kim-

boza forest, the type locality of P. lachesis sp. n.

Among all species, P. hustachei comb. n. has the largest

known distribution on forested slopes of five volcanic

mountains forming a nearly straight latitudinal chain some

220 km long: Mt. Kilimanjaro, Mt. Meru, Mt. Mulundi,

Mt. Oldeani and the adjoining Ngorongoro highlands

(NKVB, Fig. 1 1 ). Only Mt. Kilimanjaro specimens from

two localities (Fig. 1 1 ) were available for DNA analysis

(Fig. 10), while morphological similarity was the main cri-

terion to group all specimens into a single species, includ-

ing those not barcoded. A similar approach was used with

morphologically homogenous P. atropos sp. n. known

from North and South Pare mountains, but only the lat-

ter locality was represented on the DNA-based topology.

The ancient lowland or Eastern Arc Mountains in Tanza-

nia were long believed to harbour numerous ancient mois-

ture-dependent low-dispersal animals (Tolley et al. 2011

and references therein), such as Paociyptorrhinus weevils.

The presence of two species in the geologically young

forests on volcanic cones (Fig. 1 1 ) is, therefore, a remark-

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Paoayptorrhimts weevils from Tanzania

15

able result strongly implying their relatively recent colo-

nization. The same species grouping (i.e. those from old

versus young forests), is upheld with the ML and

Bayesian phylogenetic trees (Fig. 10), and morphologi-

cally (see the first couplet of the key). Only a few other

similarly habitat-dependent and locally abundant radia-

tions such as homed chameleons (Ceccarelli et al. 2014),

brevicipitid frogs (Loader et al. 2014) or bush crickets

(Hemp et al. 2014 and references therein) were studied in

sufficient detail and to evaluate distributional patterns,

which were found to be similar to that reported for

Paoayptorrhimis.

Equally remarkable is the fact that the split between both

species inhabiting young forests on volcanic highlands is

dated at 7.89 MY (Fig. 10). This is about four times old-

er than the origin of these mountains and, subsequently,

predates the origin of the wet forest habitat suitable for

these beetles. The last observation strongly suggests that

(A.) separation between P. hustachei comb. n. and P.

hanangensis sp. n. is not attributable to their currently ob-

served allopatry (Fig. 11) and that (B.) the most recent

common ancestor of each of these two species had means

to colonize both localities across presently unsuitable dry

savannah landscapes. A similar phenomenon was hypoth-

esized for a radiation of flightless Galapaganus Lanteri

endemic to, and markedly older than, the Galapagos Is-

lands (Sequeira et al. 2000).

Acknowledgements. Curators of the collections mentioned

above variously helped in accessing specimens under their care.

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

read an early draft of this work.

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Bonn zoological Bulletin 64 ( 1 ): 16-31

July 2015

Species richness of termites (Blattoidea: Termitoidae)

and ants (Hymenoptera: Formicidae)

along disturbance gradients in semi-arid Burkina Faso (West Africa)

Dorkas Kaiser' *, Tra-Bi Croulaud Sylvain% Kolo Yen’, Souleymane Konate' & Karl Eduard Linsenmair'

'Department ofAjumal Ecology’ and Tropical Biology’, Am Huhland, D-97074 Wiierzhiirg, Germany

‘ Corresponding author: E-mail: dorkas.kaiser@biozentrwn.iini-wuerzburg.de, Tel. +49 1 73-7204729:

E-mail: ke_lins@hiozentriim.iini-wiierzburg.de, Tel. +49 931-3184351

■ Universite Jean Lorougnon Guide de Daloa, BP 150 Daloa, Cote d'Ivoire: E-mail: tcrolaud@yahoo.fr, Tel. +225 05-384115

■'Universite Nangiii Ahrogoua, Research unit on Ecology’ and Biodiversity, Station d'ecologie de Lamto, BP 28 N'Douci,

Cote d’Ivoire: E-mail: koloyeo@yahoo.fr, Tel. +225 05-849545: E-mail: skonate2@yahoo.fr Tel. +225 20-304284

Abstract. From 2004 to 2008, we conducted the first inventory of tennites (Blattoidea: Termitoidae) and ants (Hymenoptera:

Formicidae) in Burkina Faso (West Africa) where these important ecosystem engineers are the only active, quantitative-

ly remarkable soil macrofauna during a long dry season. We combined two standard assessment protocols for tropical

forests and adapted them to semi-arid (agro)ecosystems to representatively characterize termite and ant communities in

two agricultural systems, and assess their response to land-use intensification. In total, 65 ant species and 41 tennite species

were collected along parallel transects replicated during consecutive years. Both taxa were highly sensitive to human im-

pact. Their taxonomic and functional richness strongly decreased with increasing intensification from a near-natural sa-

vannah towards cotton fields with recent pesticide application (North Sudan region), and recovered along a succession

of the traditional rehabilitation technique Zai' from bamen, crusted land towards a long-term fallow with diverse flora and

fauna (sub-Sahel region). Fungus-growers dominated the termite communities in all habitats, with the highest variations

between habitats noted among soil- and grass-feeders.

Key words. Soil fauna, community composition, anthropogenic impact, land-use gradient, traditional soil restoration,

sampling protocol.

INTRODUCTION

Termites (Blattoidea: Termitoidae) and ants {Hy-

menoptera: Formicidae) are of extraordinary ecological

and economic importance throughout the tropics and sub-

tropics, particularly amid arid and infertile environments

(Evans et al. 2011; Lee & Foster 1991 ). In Burkina Faso

(West Africa), termites and ants are the only active, t]uan-

titatively remarkable soil macrofauna during the long diy

season (Rouland et al. 2003; Schuumran 2006). Their rich-

ness and abundance may be severely affected by land-use

practices (for example, see reviews of Davies et al. 2003;

Holldobler & Wilson 1990; Lobry de Bruyn 1993). Agri-

cultural practices are a leading cause of biodiversity loss

(Green et al. 2005; Millennium Ecosystem Assessment

2005) and land degradation (Bai et al. 2008). Particular-

ly existential are the consequences in sub-Saharan Africa

where millions of people depend on agriculture for their

livelihoods and where vast areas are covered with degrad-

ed, baiTen soils (Batjes 2001 ). One approach to develop

more sustainable agriculture is to harness ecosystem serv-

ices of naturally occurring organisms (Evans et al. 2011).

Temrites and ants are major examples for soil-dwelling

Received: 04.08.2014

Accepted: 18.11.2014

‘ecosystem service providers’ (Jouquet et al. 2014) that

influence ecosystem functioning by physically altering

their biotic and abiotic surroundings (Jones et al. 1997).

Both taxa contain species that may become a pest in agri-

cultural systems; however, they are far more often bene-

ficial, especially in di-yland agriculture (Evans et al. 2011).

Through soil turnover (tunnelling and sheeting production)

during their foraging and nesting activities, termites and

ants further the soil’s aeration, enhance absorption and

storage of water in soils, and facilitate carbon fluxes and

storage (for example, Lobry de Bruyn & Conacher 1990;

Sileshi et al. 2010). An impressive example of a tradition-

al soil water conseiwation method in semi-arid West

Africa, making use of the positive effects termites exert

on soil properties, is the Zai' practice (Kabore & Reij 2004;

Reij et al. 2009b).

Despite a widespread recognition of the major impor-

tance of temrites and ants in arid and nutrient-poor envi-

ronments, nothing was known about either their actual di-

versity in Burkina Faso or the effects of the ongoing hu-

man-induced changes on the composition of their com-

munities. Most studies in West Africa concentrated on

forests, the few in savannahs and agroecosystems were

Conesponding editor: R. Peters

Termites and ants in semi-arid Burkina Faso

17

Fig. 1. The geographic location of Burkina Faso in West Africa and of the two study regions - the Yatenga Province in the sub-

Sahel zone ( 1 ), and the Gounna Province in the North Sudan region (2).

conducted in humid regions, such as the Comoe NP and

the Lamto Reserve in the Cote d’Ivoire (Dosso et al. 2013;

Konate & Kampmann 201 1; Kone et al. 2012; TraBi 2013;

Yeo et al. 2011). We, therefore, selected two traditional

agricultural systems characteristic for the region in focus

and assessed the invertebrate’s diversity between 2004 and

2008. We combined two well-established standard proto-

cols for sampling termites and ants in tropical forests, and

adapted them to semi-arid savannahs and agroecosystems.

This newly revised protocol has proven highly efficient

in representatively assessing and monitoring the temiite

and ant fauna.

MATERIAL AND METHODS

Study regions

We selected two traditional agricultural systems charac-

teristic for the two main phytogeographic regions in

Burkina Faso, each representing a disturbance gradient

comprising four distinct habitats, presently differing in

the magnitude of human intervention but formerly hav-

ing the same initial state.

Disturbance gradient in the Yatenga Province (sub-Sahel

zone). As the first land-use intensification gradient, we se-

lected a temporal cross-section of the traditional restora-

tion system Zai in Ouahigouya (Yatenga Province) in the

sub-Sahel region (altitude 336 m) (Fig. 1, Table 1 ). The

climate is characterized by a rainy season of four to five

months and a long dry season with high temperatures (UN-

ESCO 1977). Between 1998 and 2008, the mean annual

rainfall was about 660 mm, the mean monthly tempera-

ture varied between 19.1°C and 39.1°C.

In the Zai practice, the farmers dig pits (microwater-

sheds) in parallel rows; shortly before the first rains, they

put crop seeds and air-dried compost into the pits. Each

farmer adapts the pits to his farm conditions (pit number,

dimensions, quantity of compost) (Hien & Ouedraogo

2001). Some use Zai to intensify cereal production, oth-

ers for reforestation, or both (Reij et al. 2009a). The or-

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Dorkas Kaiser et al.

Table 1. Abbreviation, characteristics and history of study sites chosen in Burkina Faso (for each gradient, soiled according to

increasing human impact).

First disturbance gradient: four age-stages of the Za'i system in the sub-Sahel region

ZF30 Old Za'i forest (13°32’25”N; 2°22’57”W). In 2009: 30-year Za'i restoration. First cultivated for four years, then 26 years

fallow; diverse woody and herbaceous vegetation; mostly thin but closed canopy; topsoil sandy loam.

ZF20 Young Za'i forest ( 13°32’34”N; 2°22’55”W). In 2009: 20-year Zai restoration. First cultivated for four years, then 16 years

fallow; diverse woody and herbaceous vegetation but still degraded areas; mostly open canopy; topsoil loam.

ZMil Millet fields ( 13°32’29”N; 2°23’07”W). In 2009: both fields 11-year Zai cultivation; millet and green beans. At the time

of sampling, the fields were in the fifth, seventh and eighth year; topsoil loam.

ZDeg Degraded, barren area ( 13°32’36”N; 2°22’59”W). Initial stage of all Zai stages; two plots of ferruginous, infertile Lithosol;

barren, crusted and impemieable surface; no vegetation; topsoil sandy loam.

Second disturbance gradient: four land-use types in the North Sudan region

FRes Pa/na resen’e ( 1 1°38’26”N; 0°3 1 ’45 ’E). Initial stage of all North Sudanese habitats; two plots of protected arboreous and

shnibby savannah; open canopy; dense grass sods; managed by annual fire; topsoil sandy loam.

FPas Pasture area (11°38’40”N; 0°30’03’E). In 2009: exclusively used for grazing cattle since 20 years; most areas with iso-

lated trees and bushes; some barren spots; topsoil sandy loam.

FFal Short-term fallow { 1 1°38' 17”N; 0°30’09’E) from 2003-2006. In 2009: 19 years. At the time of sampling: second and fourth

year fallow. Before fallowing, crop rotation practised every third year, including cotton for four years. All trees had been

felled; dense grass-cover; topsoil clayey loam.

FCot Cotton fields ( 1 1°38’ 15”N; 0°30'27'E). Two fields, in 2009 both 13 years. At the time of sampling: second year cotton cul-

tivation; crop rotation practiced every third year including four years cotton with extensive use of different pesticides (Fan-

ga 500EC, Caiman 500EC, Calthio E, Capt 88EC, Attakan C344SE, Rocky C386EC, Sherphos 230EC) and 200 kg com-

mercial fertilizer per hectare; topsoil loam.

Soil texture was analysed in 3-7 composite samples of the habitat’s topsoil (0-10 cm).

ganic matter attracts temiites which construct galleries in

the crusted soil to commute between the food source and

the nest, thereby increasing the soils’ aeration, water in-

filtration and water holding capacity (Roose et al. 1993).

We selected four age-stages (for the characteristics, see

Table 1). An old forest of about 30 years of age (ZF30,

Fig. 2A) and a younger one about 20 years old (ZF20, Fig.

2B) represented the reforestation (fallow) phase. Two mil-

let fields (ZMil, Fig. 2C) were selected for the cropping

period, and two barren, cmsted areas (ZDeg, Fig. 2D) for

the initial stage. Each site lay within a radius of 300^00

m of the other and comprised an area of about 1 hectare.

Disturbance gradient in the Gourma Province (North Su-

dan region). The second disturbance gradient was chosen

52 km south of Fada N’Gounna (Gounna Province) in the

North Sudan region (altitude 253 m) (Fig. 1, Table 1 ). The

rainy season lasts from around mid-April till October

(Some et al. 2006). Between 1997 and 2008, the mean an-

nual rainfall in Fada N’Gounria was about 875 mm, the

mean monthly temperature ranged between 19.6°C and

37.1°C. The natural vegetation is of the savannah type

(UNESCO 1977).

Two plots in the Pama reserve (FRes, Fig. 2E) with an

area of 1 km^ each represented the characteristic near-nat-

ural savannah habitat for the region (AThiombiano, Oua-

gadougou, pers. comm. 2004) and the initial habitat stage

of this disturbance gradient. A pasture (FPas, Fig. 2F), a

short-tenn fallow (FFal, Fig. 2G) and two cotton fields

(FCot, Fig. 2H) were chosen in a mral area separated from

the park border by a street. Each plot had a size of about

1 hectare.

The characteristics of each habitat are summarized in

Table 1.

The baseline sampling protocol

Our rapid assessment protocol (RAP) (Fig. 3) was based

on two widely accepted standard protocols - the protocol

of Agosti & Alonso (2000) for ants in the leaf litter and

the transect protocol of Jones & Eggleton (2000) for ter-

mites in tropical forests. The protocols were combined and

adjusted to account for the conditions prevailing in our se-

mi-arid agroecosystems; for example, only negligible

amounts of litter covered the ground and most sites had

an area of 1 hectare only.

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Termites and ants in semi-arid Burkina Faso

19

Fig. 2. Study sites in the A-D) sub-Sahel zone, and E-H) North Sudan region. A: old Zai forest (ZF30; one plot); B: young Zai

forest (ZF20; one plot); C: millet field (ZMil, two plots); D: degraded, barren area (ZDeg; two plots); E: Pama reserve (FRes; two

plots); F: pasture area (FPas; one plot); G: short-term fallow (FFal; one plot); H: cotton fields (FCot; two plots).

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Dorkas Kaiser et al.

Fig. 3. Relative position of sampling sites according to the combined and standardized rapid assessment protocol (RAP), fol-

lowed to assess tennite and ant communities.

For termites, the RAP included 10 contiguous 5 x 2 m

sections resulting in a transect length of 50 m. The first

transect in each habitat had a length of 100 m as in Jones

& Eggleton (2000), but many of those run in 1 hectare

plots had to be divided into two parallel parts. In each sec-

tion, tennites were first searched in as many different mi-

crohabitats as possible (such as accumulations of litter, in-

side and underneath dead logs and twigs, epigeal mounds

and mnways on vegetation). Then, eight samples of sur-

face soil (‘soil scrapes’), each 12 x 12 cm and 15 cm deep,

were taken at random locations and searched in situ. The

sampling time per section was 1 man-hour; stopping rules

applied if no microhabitats could be found, resulting in a

shorter sampling time.

Ants were collected on parallel transects in a distance

of 10 m on both sides of the tennite transect at eight sam-

pling sites (four on each side every 12 m). Within an area

of 1 m- the vegetation was at first searched for ants and

thereupon totally removed, and all ants walking on the

ground were collected for a duration of five minutes

(called ‘Winkler-site’). It substituted the mini-Winkler ex-

tractor of Agosti & Alonso (2000), which was applied dur-

ing the first year but failed to capture most of the ants due

to the limited amounts of litter in our study sites. In a

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Temnites and ants in semi-arid Burkina Faso

21

1 m distance, pitfall traps with a diameter of 1 0 cm were

buried in the soil and tilled with 45% ethanol; they were

run for 48 hours. Following the newly revised RAP, ants

were additionally collected when encountered in the sec-

tions of the belt-transects, i.e. in microhabitats and soil

scrapes. For a detailed description and illustration of the

working steps, please refer to Kaiser et al. (2011) and

Kaiser (2014), the relative position of sampling sites ac-

cording to the combined assessment protocol is illustrat-

ed in Fig. 3.

Species identification. Specimen of both taxa were

stored in vials with 90% ethanol; they were identified to

the level of species or, whenever this was impossible, to

numbered moiphospecies - ants by using the taxonomic

works of, for example, Bolton (1994, 1995) and Bolton

et al. (2007); termites following Bouillon & Mathot

( 1965), Emerson (1928), Sands ( 1998), and Webb ( 1961 );

and both with reference to collections from West Africa

held in the research station in Lamto NP (University Nan-

gui Abrogoua, Cote d’Ivoire). Own reference collections

were assembled for both taxa and will be stored in the

Natunnuseum Senckenberg (Frankfurt am Main, Ger-

many). However, presently we still work with the collec-

tion in the research unit on ecology and biodiversity,

UREB (Prof Souleymane Konate), University Nangui Ab-

rogoua (Abidjan, Cote d’Ivoire). According to their diet

and mandible morphology (Deligne 1966) and following

Grasse (1986), termite species were assigned to one of

these four functional groups: soil-feeders, wood-feeders,

grass-feeders and fungus-growers.

All data presented here originate from three to four tran-

sects conducted in each land-use type in the rainy seasons

between 2004 and 2008. Four replicate transects were run

in the Zai stages (sub-Sahel zone) and the Pama reserve,

three in the other habitats of the North Sudan region.

Statistical analyses

Species accumulation and rarefaction cun’es

Incidence-based species accumulation curves were used

to graphically compare the species richness of the habi-

tats belonging to the same disturbance gradient. When the

sampling effort differed between the habitats to be com-

pared (for example, in the North Sudan region), the rar-

efaction method was used to calculate the number of

species expected in a subsample randomly selected from

multiple samples (Chiarucci et al. 2008; Koellner et al.

2004). Smoothed curves were produced by randomizing

the data 500 times. The slopes of the cuiwes illustrate the

rate at which new species were added with an increasing

sampling effort. The increasing sampling effort was equiv-

alent with an increasing number of a) transect sections for

termites and b) sampling units, each comprising one tran-

sect section, one pitfall trap and one Winkler quadrate in

the case of ants. The software Estimates Version 7.5 1 for

Windows (Colwell 2005) was used to calculate the species

accumulation and rarefaction curves.

Comparisons among study sites

The software SPSS 15.0 (SPSS Inc. 2006) was used for

comparisons between the different habitats concerning

their species composition and the number of species ob-

served (or expected at the common sampling effort). Fried-

man’s ANOVA was used to compare land-use types be-

longing to the same gradient. The Wilcoxon sign-rank test

was used for pairwise comparisons between related land-

use types. The Bonfen'oni correction was applied after-

wards to adjust the level of significance.

Disturbance index

To analyse the coiTelation between species richness and

habitat disturbance, we combined the data of both regions

and assigned each habitat type to one of the four distur-

bance classes (ordinal scale), with a disturbance index of

1 used for the near-natural habitats (old Zai Forest and Pa-

ma reserve), 2 for pasture land and fallows (young ZaT For-

est, short-term fallow), 3 for agricultural plots (millet

fields, cotton fields) and 4 for fully degraded, barren land.

A linear regression was then performed in SPSS 15.0

(SPSS Inc. 2006) with the number of a) ant species and

b) termite species collected per site and plot as the depend-

ent variable, and the disturbance index and study region

as independent ones. The study region was defined as a

nominal variable with 1 for the sub-Sahel zone and 2 for

the North Sudan zone.

RESULTS

Combining the sites of both regions, 65 ant species (25

genera and nine subfamilies) and 41 termite species ( 13

genera and four subfamilies) were collected between 2004

and 2008.

Species checklists

Ant communities

A checklist of all ant species present in the different land-

use types is given in Table 2. The lowest species, genus,

and subfamily numbers were found in highly-intensified

areas, the degraded land in the sub-Sahel zone and the cot-

ton fields in the North Sudan region. The ant fauna in

Burkina Faso was dominated by the Mymiicinae, which

comprised 54% of the total number of species collected

along both disturbance gradients, followed by the Formic-

inae (22%) and the Ponerinae (9%).

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Dorkas Kaiser et al.

Table 2. Ant (morpho)species present (X) in the Zai stages in Ouahigouya (sub-Sahel zone) and in the habitats near Fada N’-

Gounna (North Sudan region).

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Tennites and ants in semi-arid Burkina Faso

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Land-use types: old Zai forest (ZF30), young ZaT forest (ZF20), millet fields (ZMil), degraded area (ZDeg); near-natural savannah (FRes), pasture

area (FPas), short-term fallow (FFal), cotton fields (FCot).

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Dorkas Kaiser et al.

Table 3. Termite (moipho)species present (X) in the habitats in Ouahigouya (sub-Sahel zone) and near Fada N’Gourma (North

Sudan region). A two-digit shortcut before the genus name indicates the functional group to which it belongs to: fungus-growers

(fg), grass-feeders (gf), soil-feeders (sO, wood-feeders (wf), unknown (uk).

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Termites and ants in semi-arid Burkina Faso

25

Land-use types: old Zai forest (ZF30). young Zai forest (ZF20), millet fields (ZMil), degraded area (ZDeg); near-natural savannah (FRes), pasture

area (FPas), short-term fallow (FFal), cotton fields (FCot).

Together, the four Za'i stages hosted 41 ant species out of

16 genera and six subfamilies (Table 2). The number of

species decreased with increasing human impact (decreas-

ing habitat restoration) from 35 species ( 14 genera) in the

old Zai' forest to 15 (nine genera) in the degraded area.

Ant assemblages in the North Sudanese sites were rich-

er in species, genera, and subfamilies: 53 ant species out

of 23 genera and nine subfamilies were found in total

(Table 2). The number of species decreased with increas-

ing land-use intensification from 37 species (20 genera)

in the near-natural savannah in the Pama reseiwe to 1 9 ant

species (nine genera) in the cotton fields with recent pes-

ticide application.

Termite communities

The checklist of all termite species present under the dif-

ferent disturbance regimes is given in Table 3. Lowest

numbers of species, genera, subfamilies, and functional

groups were found in areas heavily impacted by humans.

Fungus-growers dominated the tennite fauna in all study

sites. The highest variations between the habitats occuiTed

in soil- and grass-feeding termites.

The four Zai stages together hosted 33 tennite species

out of 1 1 genera and five subfamilies (Table 3). The num-

ber of termite species strongly decreased with increasing

human impact from 25 species (13 genera) in the old ZaV

forest to only one fungus-growing Oclontotennes species

in the most degraded stage. Except for the degraded site,

all four functional groups were represented in the ZaY

stages.

Together, the four habitats in the North Sudan region

hosted 31 termite species out of 13 genera and five sub-

families (Table 3). The decreasing species richness was

obvious along the disturbance gradient: the number of ter-

mite species was highest in the reserve with restricted hu-

man impact (24 species, 1 1 genera) and drastically de-

creased with increasing human pressure to five species

(three genera) in the cotton fields. Only the reserve and

the short-term fallow hosted soil-feeding temiites.

Comparisons among study sites

Incidence-based, randomized rarefaction or accumulation

cuiwes were used to graphically compare the discoveiy rate

and the species richness of related habitats (Figs 4-5). Er-

ror bars correspond to one standard deviation of the mean

after randomization. The colours in Figs 4-5 are identi-

cal to those used for the disturbance classes in Figs 6-7.

Ant communities

For ants collected between 2004 and 2008, rarefaction

curves are presented in Fig. 4 for the habitats located a)

in the sub-Sahel zone, and b) in the North Sudan region.

For each region, we statistically compared the species rich-

ness at the level of the land-use type with the lowest sam-

ple number (degraded area, cotton fields). The number of

ant species predicted at the common sampling effort (39

sampling units, in Figs 4a, b indicated by the dotted line)

differed significantly between the sub-Sahelian (Friedman,

N = 39, X" (3) = 1 17, P < 0.001 ) and the North Sudanese

habitats (Friedman, N = 39, x’(3) = 117, P < 0.001). A

Wilcoxon sign-rank test (with BonfeiToni con'ection) re-

vealed statistical differences also between the 20-year-old

forest and the millet field (Wilcoxon, Z = -5.44, P< 0.001,

R = 0.62). In the sub-Sahel zone (Fig. 4a), the descend-

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Dorkas Kaiser et al.

Ant communities in the sub-Sahel zone

ID

<D

Z5

E

D

O

0

JD

E

D

C

(/)

0

’o

0

Q.

(/)

— old forest

— young forest

— millet field

degraded land

Ant communities in the North-Sudan zone

■D

0

:d

E

Z3

o

i_

0

JD

E

13

C

CO

0

o

0

Q.

CO

— Pama reserve

- -pasture

— fallow

— cotton field

(b)

0

10 20 30 40 50

sampling units

Fig. 4. Incidence-based rarefaction curves (including standard deviations) predicting the accumulation rate of ant species with

increasing sampling effort in the habitats selected along the disturbance gradient in the a) sub-Sahel zone, and b) North Sudan re-

gion. Curves are predicted from richness values assessed along RAP transects; the rarefied number of 39 sampling units (the dot-

ted line) was used to compare the related habitats. In the sub-Sahel zone, the expected richness and discovery rate of ant species

decreased with decreasing restoration stage, and from the short-tenn fallow towards the cotton field in the North Sudan region.

However, both the richness and the accumulation rate were almost identical in the fallow and the reserve (indicated by the over-

lapping error bars).

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Temiites and ants in semi-arid Burkina Faso

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Termite communities in the sub-Sahel zone

— old forest

young forest

— millet field

degraded land

(a)

0

10 20 30 40 50

sampling units

Termite communities in the North-Sudan zone

— Pama reserve

pasture

fallow

— cotton field

(b)

0

10 20 30 40 50

sampling units

Fig. 5. Incidence-based, randomized accumulation (a) and rarefaction curves (b) indicating the accumulation rate of termite species

with increasing sampling effort in the habitats selected along the disturbance gradient in the a) sub-Sahel zone, and b) North Su-

dan region. In the sub-Sahel zone, the richness and discovery rate of termite species decreased with decreasing restoration stage.

In the North Sudan region, the richness predicted at the common sampling effort of 39 sampling units (the dotted line) decreased

from the Pama reseiwe towards the cotton field, although the differences between the pasture and the fallow are not significant (in-

dicated by the overlapping error bars).

Bonn zoological Bulletin 64 ( 1 ): 16-3 1

®ZFMK

28

Dorkas Kaiser et al.

1

2 3 4

disturbance index

0 near-natural

fallow & pasture

0 agricultural fields

degraded land

''' Fit line for total

(/)

■q

0

Q.

C/)

0

E

0

o

0

E

c

25

20

15

10

5

0

disturbance index

0 near-natural

fallow & pasture

0 agricultural fields

^ degraded land

' ' ' Fit line for total

disturbance index

Fig. 6. CoiTelation between habitat disturbance and the number of a) ant species and b) tennite species collected per site and plot

in the sub-Sahel (circles) and the North Sudan region (rhombus). There was a significant negative relationship between the distur-

bance of a habitat and the number of ant species (P < 0.05, F2.9 = 1.22, R’ = 0,71 ) and tennite species (P < 0.001, F2,9 = 37.26,

R- = 0.89).

ing order of all land-uses with regard to the expected num-

ber of ant species is old forest > young forest > millet

fields > degraded soil, and short-term fallow > Pama re-

serve > pasture > cotton field in the North Sudan region

(Fig. 4b). However, ant species richness did not differ sig-

nificantly between the fallow and the reserve (Wilcoxon,

P > 0.025 after BonfeiToni coiTection) - both habitats al-

so exhibited almost identical rates at which new species

were added with the increasing sampling effort.

Termite communities

Quite similar to the trend observed for ants, the number

of tennite species observed in all transect sections differed

significantly between the Zai stages in the sub-Sahel zone

(Friedman, N = 50, f (3) = 141 .94, P < 0.001 ) (Fig. 5a)

and between the four North Sudanese habitats (Friedman,

N = 39, X‘ (3) = 106.68, P < 0.001 ) (Fig. 5b). In the sub-

Sahel zone, the descending order of the habitats with re-

Bonn zoological Bulletin 64 ( 1 ): 16-31

®ZFMK

Teirnites and ants in semi-arid Burkina Faso

29

gard to the number of termite species observed was old

forest > young forest > millet field > barren land. At a low

sampling effort, however, the discovery rate of termites

in the millet field was higher than in the young forest (in-

dicated by the crossing curves at sampling unit 9). In the

North Sudan region, the number of species predicted at

the common sampling effort decreased in the order: re-

serve > pasture > fallow > cotton field although the dif-

ferences between the pasture and the fallow were not sig-

nificant (Wilcoxon sign-rank test, P > 0.025 after Bonfer-

roni correction). The rate at which termite species were

added at a lower sampling effort was higher in the short-

terni fallow than in the pasture (please note the crossing

curves at sampling unit 27).

All rarefaction and accumulation curves roughly ap-

proach an asymptote. Figs 4-5, therefore, indicate that we

representatively characterised the temiite and ant fauna,

also proving that the spectrum of methods we used was

appropriate.

Effect of land-use intensity on species richness

To analyse the correlation between species richness and

habitat disturbance, each habitat type was assigned to one

of four disturbance classes with 1 used for the near-nat-

ural habitats and 4 the degraded, barren land (please re-

fer to the method-section).

Fig. 6 shows the number of ant (Fig. 6a) and termite

(Fig. 6b) species collected in Burkina Faso as a function

of the habitat disturbance. Each point comprises the num-

ber of species collected per study plot; the study region

is indicated by the shape of the point with a rhombus used

for the North-Sudanese sites and circles for the sub-Sa-

helian ones.

A linear regression analysis was perfonned. The scat-

terplots (Figs 6a, b) suggest a strong negative coiTelation

between species richness and habitat disturbance: as the

land-use intensity increases, the species richness of ter-

mites (P < 0.001 , F2,9 = 37.26) and ants (P = 0.05, F2,9

= 1 1 .22) decreases.

For the ant species, richness R- is 0.71, i.e. only 29%

of the variance in species richness could not be explained

with this model. When the ants’ species richness in a habi-

tat was predicted, land-use intensity was found to be a sig-

nificant predictor (P = -0.86, P < 0.05), whereas the study

region was not (P = -0.05, P > 0.05). When the termites’

species richness in a habitat was predicted, both land-use

intensity (P = -1.03, P < 0.001) and study region

(P = -0.39, P < 0.05) were found to be significant predic-

tors. The overall model fit for temiites is = 0.89, i.e.

only 1 1 % of the variance could not be explained with the

model.

DISCUSSION

Combining both regions, 65 ant species (25 genera and

nine subfamilies) and 42 termite species (13 genera and

five subfamilies) were collected according to the combined

RAP (Tables 2 and 3). Depending on the study site, be-

tween 40 and 50 transect sections (Figs 4-5) were sam-

pled between 2004 and 2008. All accumulation and rare-

faction curves in Figs 4-5 roughly approach an asymp-

tote, therefore, indicating that we representatively char-

acterized the termite and ant fauna. To our knowledge,

these findings represent the first records for temiites and

ants in Burkina Faso.

Species richness along disturbance gradients. Termites

and ants were highly sensitive to land-use intensification

- extensively managed or degraded habitats hosted the

lowest numbers of species, genera, subfamilies, and func-

tional groups (Tables 2 and 3, Fig. 6). Similar findings

were also reported from other regions: termite communi-

ties were consistently found to be less diverse in disturbed

habitats (for example, see Davies et al. 1999; Donovan et

al. 2007; Gathome-Hardy et al. 2006; Vaessen et al. 201 1 ),

and crops and plantations hosted less species than near-

by natural vegetation (see Black & Okwakol 1 997 and ref-

erences therein). Although direct comparisons are ham-

pered due to differences in soil type, rainfall or the inten-

sify of agricultural activity, a review of 10 studies revealed

that ant species richness was nearly always lower in farm-

land environments than in naturally vegetated areas (Lo-

bry de Bruyn 1999). Similarly in Cote d’Ivoire, the num-

ber of ant species decreased from relatively well conserved

to the most extensively managed land-use types (Yeo et

al. 2011).

Therefore, and since increasing land-use intensification

progressively simplified the habitats’ physical stmcture in

our study sites (Table I , Figs 2A-H), we did expect that

the species richness of both taxa would decrease with in-

creasing human impact. Very astonishing, however, was

the extent to which their richness decreased in heavily im-

pacted habitats. The termite fauna was especially impov-

erished. Unsustainable land-use and the reduction of veg-

etative cover affect temiites and ants, for instance, by re-

sulting in a loss of available feeding and nesting sites or

by exposing them to greater physiological stress in the pe-

dosphere (for example, see Agosti et al. 2000; Jones et al.

2003).

The enormous loss of taxonomic and functional diver-

sity of the focal arthropods may have disastrous conse-

quences for small-scale subsistence farmers, impeding a

sustainable agricultural production. Hence, especially

promising are the observations concerning a resilient agri-

cultural production and the progressive restoration of an

increasingly diverse flora and fauna (particularly temiites

and ants) on initially baneii crusted soils with the Zai prac-

tice. To our knowledge, no other study has investigated

Bonn zoological Bulletin 64 ( 1 ): 1 6-3 1

©ZFMK

30

Dorkas Kaiser et al.

the recovery potential of termite and ant communities in

fully degraded, cmsted soils. The study of Gathorne-Hardy

et al. (2002) in Southeast Asia revealed that an Imperata

cvlindrica grassland and a cassava field, both hosting a

strongly reduced temiite fauna, had the potential to recov-

er when left fallow near a source area of primary forest

for more than 50 years. Dibog et al. (1999) observed that

tree-based agriculture in the Amazon had the potential to

support the function of soil termite fauna even on land pre-

viously used as pasture.

To conclude, the present study provides a baseline for

future monitoring studies in Burkina Faso and the com-

parison of termite and ant diversity between the different

disturbance regimes. We present a combination of stan-

dard sampling methods (Fig. 3) that are easy to learn and

effective to representatively, but rapidly, assess termites

and ants in various (agro (ecosystems of semi-arid tropi-

cal regions (Figs 4-5). Since crop production will have

to increase strongly to meet the growing food demand

driven by the ongoing human population growth (Godfray

et al. 2010), more techniques are needed to increase soil

fertility and maintain the sustainability of agricultural pro-

duction by harnessing ecosystem services of naturally oc-

curring organisms. Given the crucial role of termites and

ants in ecosystem functioning throughout the tropics and

subtropics, special focus should be put on techniques that

support or fasten the recovery of their communities in

highly degraded lands. More studies are, therefore, need-

ed that investigate the responses of tennite and ant com-

munities to habitat perturbations, the succession of their

communities during habitat rehabilitation, and that clari-

fy how the successions relate to the recovery of biodiver-

sity and ecosystem processes.

Acknowledgements. We are especially grateful to the fanners

Yacouba Sawadogo (sub-Sahel zone) and Ousmane Loualy

(North Sudan region), and their families for allowing access to

their land, and to Lardia, Boukare, Ali and Inoussa for their as-

sistance in the field. This study was conducted within the frame-

work of an interdisciplinary project, ‘BIOTA West Africa’

(FZOlLCOOl 7a), funded by the Gennan Federal Ministry of Ed-

ucation and Research (BMBF).

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Bonn zoological Bulletin 64 ( 1 ): 16-31

®ZFMK

32

Book Review

Bohme, W. (2014): Herpetology in Bonn. Mertensiella 2 1 : 256

pp.

The impressive book reviews the history of herpetological re-

search and herpetological collections in Bonn in unexpected de-

tails. It starts with the old Prussian university, which hosted the

former natural history museum. This chapter also includes bio-

graphic sketches about important researchers, who worked in the

field of herpetology and were based in Bonn, or worked with

the Bonn collection. This includes e.g., well known persons like

Georg August Goldfuss, Franz von Leydig, and Georg .Ian. This

chapter ends with a short summary about contemporary institu-

tions involved in research about amphibians and reptiles, like

the Federal Agency of Nature Conservation which is based in

Bonn.

However, the most important chapters of the book are about

the Zoological Museum Alexander Koenig (ZFMK) and its her-

petological research. At this point I have to say that the book

could be easily an auto-biography, because Wolfgang Bohme is

by all means the person who is responsible for the internation-

al reputation of the ZFMK heipetological collection. However,

it is very enjoyable that the person who has the best knowledge

about this collection wrote about the history and the involved

former researchers and recent students, instead about primarily

himself The part about the ZFMK collection starts with a his-

toric sketch about the foundation of the herpetological depart-

ment and its first curator. Karl F. Buchholz. It is followed by

sketches about the succeeded curators, but most importantly by

the history, including important researchers, of the four main col-

lections which were donated to the ZFMK collection. This in-

cludes specimen donation from the University of Kiel and Hei-

delberg, as well as the Gottingen collection and the Paul Muller

collection. This historic part is followed by a complete type cat-

alogue of the ZFMK heipetological collection, a chapter about

national and international herpetological symposia held at the

ZFMK. and is rounded down by a chapter on herpetological

teaching at the ZFMK, and its “scientific output”.

Although the last type catalogue was published lately ago

(Bohme 2010), the type catalogue herein is more than just an

update. It includes not only also detailed infonnation also about

secondary types, but also to specimens erroneously considered

as types, and lost type specimens, which are mainly from the

Gottingen collection. Moreover, many important comments are

given to problematic specimens. Therefore, this type catalogue

is not only the most cun'ent catalogue, but also the most detailed

one of the ZFMK heipetology collection.

By several points this book is an important contribution to her-

petology, It is the first detailed review of herpetological research

in Bonn, also including other institutions than the ZFMK. It in-

cludes a detailed history about the ZFMK collection and the re-

searchers, as well as students who are involved in the “evolu-

tion” of this collection, with a strong international reputation,

and it includes the most updated type catalogue of primary and

secondary types. All of this is written by the person who has not

only an excellent knowledge about the ZFMK itself, but also the

best overview regarding herpetological research in Bonn. Un-

der the authority of Wolfgang Bohme, the collection has grown

from about 10.000 specimens to one the largest collections in

Gcnnany. Therefore it is hard to find any critical points or mis-

takes. There are some minor details in the type catalogue, e.g.

that it could be mentioned that the recent name of Lacerta stel-

lio is Stellagama stellio. Moreover I am very sure that, even that

the book is published in 2014, the type catalogue was out of date

by the time of its publication, largely due to ongoing research

of its author.

Summarizing I can say that everybody who is interested in

the history of the ZFMK collection, or in the history of German

heipetological research, should have this book. It is full of de-

tails and historic herpetological knowledge, and it serves some

enjoyable early photographs of important recent herpetologists

like e.g., Frank Glaw, Ulrich Joger, Jom Kohler, Andreas

Schmitz, Miguel Vences, Thomas Wilms and Thomas Ziegler.

REFERENCES

Bohme, W. (2010) A list of the herpetological type specimens

in the Zoologisches Forschungsnuiseum Alexander Koenig,

Bonn. Bonn zoological Bulletin 59: 79-108

Philipp Wagner, Amphibians and Reptiles, Naturmuseum, Um-

weltschutz-Informationszentrum Lindenhof, Karolinenreuther

StraBe 58, 95448 Bayreuth, tel. +49 921-75942-0;

E-mail: philipp.wagner.zfmk(aiuni-bonn.de

Bonn zoological Bulletin 64 ( 1 ): 32

©ZFMK

Bonn zoological Bulletin 64 ( 1 ): 33-58

July 2015

Checklist of the Mammals of Iraq (Chordata: Mammalia)

Omar F. Al-Sheikhly', Mukhtar K. Haba\ Filippo Barbanera’, Gabor Csorba^ & David L. Harrison’

^Department of Biology’, University of Baghdad, Iraq: E-mail: alsheikhlyomar@,gmail.com

'Department of Biology’, College of Science for Women. University of Baghdad. Iraq

^Department of Biology, Zoology’ and Anthropology Unit, Via Volta 4 I-561 26 Pisa. University of Pisa, Italy

''Department of Zoology’, Hungarian Natural Histoiy Museum, Hungaiy, Baross ii. 13. Budapest, H-1088, Hungaiy

^Harrison Institute. Bowej-M’ood House, 15, St. Botolph's Road. Sevenoaks, Kent TNI 3 3AQ, UK

Abstract. A checklist of the mammals of Iraq including their current status and geographical distribution is provided.

Data were collected from literature and combined with field observations obtained during 2013-2014. The checklist com-

prises 93 species of wild mammals occun ing in Iraq and belonging to eight orders, 28 families, and 65 genera. The sta-

tus and distribution of three Erinaceomorphs, three Soricomoiphs, 20 Bats, 23 Carnivores, eight Artiodactyls, two Lago-

moqjhs, 28 Rodents, and six Cetaceans are reported. The checklist includes 24 species evaluated by the Intemational

Union for the Conseiwation of Nature and Natural Resources: one was listed as Extinct, one as Critically Endangered,

four as Endangered, eight as Vulnerable, seven as Near Threatened, and three as Data Deficient. One subspecies of car-

nivores (Smooth-coated Otter Liitrogale perspiciUata ma.xwelli) and one rodent species (Long-tailed Nesokia Nesokia

hiinnii) endemic to Mesopotamian marshlands were highlighted as taxa with conservation priority to Iraq. We also fo-

cused on main threats to the most common wild mammals with special attention to impacts of illegal hunting and poach-

ing.

Key words. Iraq, systematic checklist, vertebrates, wild mammals, endemics

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INTRODUCTION

The ten'itoiy of Iraq is comprised between latitudes 29°

to 38° N and longitudes 39° to 49° E (a small area lies

west of 39°) and spans over 437,072 km- (168,754 sq mi).

It borders with Turkey to the north, Syria to the northwest,

Iran to the east, Jordan to the west, Saudi Arabia to the

south and southwest, and Kuwait to the south. Border with

the latter is marked by a narrow section of coastline

stretching for 58 km (36 mi) in the northern Arabian Gulf

The landscape includes high mountains in the north (Kur-

distan), desert, arid lands, and sandy steppes in the west-

ern and south-western plateau (Al-Badiyah), and the

Mesopotamian marshlands in the southern alluvial plain.

This wide range of habitats awards Iraq with a noticeable

biodiversity, the wild mammals being not an exception

(Fig.l).

Received: 15.08.2013

Accepted: 16.02.2015

Since Hatt ( 1 959) little was known about the mammals

of Iraq. Most of the observations available to date for Iraq

were due to Robert T. Hatt, indeed. In 1959, he published

“The Mammals of Iraq”, where he included data collect-

ed on wild and domestic animals during surveys carried

out between October 1 952 and March 1 953 under the aus-

pices of the United States Educational Foundation of Iraq

(“Fulbright Foundation”). Hatt’s report is considered the

first checklist of the mammals of Iraq, as it includes not

only author’s own findings but also both material and re-

ports provided to him by Charles A. Reed and Lee Mer-

riam Talbot. Charles A. Reed (School of Pharmacy, Uni-

versity of Illinois, USA) joined to an expedition to study

the historical fauna of Iraq in 1954-1955. He was support-

ed by a grant of the National Science Foundation (USA)

and worked in collaboration with the Oriental Institute of

the University of Chicago. Lee Meniam Talbot visited Iraq

Conesponding editor: R. Hutterer

34

Omar F. Al-Sheikhly et al.

Fig. 1. Map showing the major landscapes in Iraq.

briefly in 1955 under the auspices of the International

Union for the Conservation of Nature and Natural Re-

sources (lUCN) to get insight into the status of a few

threatened large mammals.

Another relevant contribution to the knowledge of the

teiTestrial mammals of Iraq was provided by David L. Har-

rison (co-author; deceased March 2015), who joined the

British Royal Air Force in 1953 and earned out the ma-

jor part of his national service in Iraq. Harrison reviewed

Halt’s records and made further field observations espe-

cially by surveying the suiTOundings of Ramadi and Hab-

baniya in central Iraq (Anbar Province). His work was

published in tliree volumes entitled “The Mammals of Ara-

bia” (1964; 1968; 1972). Further updates were included

in the second edition of his book that was published in

1991. In his books, Han'ison exhaustively addressed the

entire information available for the mammalian fauna of

Iraq during the period 1953-1991.

In 1969, Nuri Mahdi and P.V. George prepared their

checklist of the vertebrates of Iraq, which included wild

and domesticated mammals. Mahdi & George (1969) list-

ed 88 mammalian species according to the systematic

aiTangement of Ellerman & Morrison-Scott (1951). They

included taxa that had been actually recorded in Iraq and

those that were assumed to occur as well, citing previous

literature records and reporting the catalogue of the spec-

imens included in the collection of the Biological Research

Centre in Baghdad (BRCB). However, Mahdi & George’s

checklist failed to provide detailed infonnation on all the

listed species (e.g., distribution range and/or collecting lo-

calities and dates).

The present checklist takes into account eight orders,

28 families, 65 genera and 93 species of mammals of Iraq

including both their taxonomic status and geographical

distribution (Table 1 ). Nomenclature adopted in this check-

list follows third edition of Mammal Species of the World

(Wilson & Reeder 2005) unless otherwise stated.

EXTINCT SPECIES

The historical record of the mammals of Iraq (Hatt 1959)

included many species that are currently extinct. The

Mesopotamian Lion {Pantliem leo persica) was present

along the Euphrates River in the early XIX"' century. Last

representatives of the species were killed at the time of

the military actions of the British forces (1916-1918). His-

torical occurrence of the Tiger (Panthera tigris) in Iraq is

documented by Kock ( 1 990). The Indian One-homed Rhi-

noceros (Rhinoceros unicornis) is known only by a sin-

gle lower tooth from the Paleolithic site of Barda Balka,

and by later sculptural representations that likely referred

to animals of Indian origin. Accurate depictions of Bison

(Bison bonasus) resembled on cylinder seals and potter-

ies were found in Mesopotamia (4000 to 2000 B.C). The

Siva’s Giraffe (Sivatheriinn giganteiini) inhabited Iraq dur-

ing the Paleolithic and was present as late as 3500 B.C..

It is presumed that the Water Buffalo (Biibalus biibalis)

was also native to Mesopotamia. In fact, the wild type was

frequently portrayed until about 2 1 00 B.C. before it went

extinct. Teeth testifying the presence of the Elephant (Ele-

phas niaxinnis) in Iraq in the late Pleistocene are also

known. Bones and representations of Wild Ox or Aurochs

(Bos prirnigenius) are abundant in archeological sites from

the Palaeolithic to the IX'" century (B.C. ). The Syrian Wild

Ass “Onager” (Eqiius hemionus hemippits) was abundant

in the plains of north-western Iraq, but since the middle

of the XIX"' centui-y its number has dramatically decreased

due to over-hunting. The last herd was reported near Ja-

balSinjarin 1927(Raswan 1935; Hatt 1959). A few bone

fragments, teeth, and some reasonably accurate depictions

of the Red Deer ( Cerviis elaphus maral) testify the early

occurrence of this species in Iraq, although there is lack

of evidence for its more recent presence. Nevertheless, the

species still inhabits Turkey and Iran. A Beaver (Castor

fiber) mandible was found in Palaeolithic deposits of Iraq

as well as near the Khabur River in eastern Syria in 1837;

the latter, however, was not reliably assessed and could

be assigned also to an Otter (Liitra sp.) (Hatt 1959). As

part of this work, we revised the taxonomic identification

of the species that were misidentified or overlooked. Fi-

nally, we included the Arabian Oryx (Oryx leiicoiyx) in

the checklist as it has occurred until recently in the south-

ern desert of Iraq (Harrison 1968; Harrison and Bates

1991; Al-Sheikhly 2012c).

Bonn zoological Bulletin 64 (1): 33-58

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Checklist of the Mammals of Iraq

35

TAXONOMIC REMARKS

Erinaceomorpha

Early records of the European Hedgehog (Eri)iaceus eu-

ropaetis) from Iraq can be referred now to the Southern

White-breasted Hedgehog {E. concolor) (Harrison &

Bates 1991).

Soricomorpha

Nader ( 1969) examined specimens of Bicolored White-

toothed Shrew, which were assigned by Hatt ( 1959) to the

Iranian subspecies {Crocidiira leiicodon persica), and re-

assigned them to C. russula. A more recent investigation

carried out by Hutterer & Hamson (1988) has provision-

ally allocated Hatt’s ( 1959) specimens to the Lesser White-

toothed Shrew (C. suaveoleus).

Chiroptera

Kinnear (1916) reported the presence ofRhiiiopoma ara-

bicum" (= R. arabium) in Iraq, a taxon formerly regard-

ed as subspecies of Lesser Mouse-tailed Bat {R. hard-

wickii) (Ellerman & Morrison-Scott 1951, Simmons

2005). Presently, the name arabicum is applicable to the

Levantine and Arabian populations of/?, cystops (Berda

et al. 2009). Khajuria (1988) described R. hadithaeusis as

a new bat species from the western Iraq; however, this tax-

on is now considered as Junior synonym of R. udcrophyl-

lum (Kock et al. 2001 ). Both Taphozous kachhensis bahy-

loiriciis and Taphozous maginis are presently regarded as

synonym of the Naked-bellied Tomb Bat ( Taphozous midi-

ventris) (HaiTison & Bates 1991, Simmons 2005). As far

as Kinnear ’s (1916) record of the Meditenanean Horse-

shoe Bat (Rhiuolophus eiuyale) is concerned, Cheesman

( 1920) and Hatt ( 1959) found no confirmation for its oc-

cuiTence in Iraq. Niazi ( 1967) reported the occurrence of

R. eiuyale in two localities in Iraq. However, according

to the cranial measurements, Han ison & Bates (1991) sug-

gested that the specimens described by Niazi were in fact

Mehely’s Horseshoe Bats (R. mehelyi). Mahdi & George

( 1969) included R. eiuyale in their list without providing

details (e.g., voucher specimens or sampling locality) on

the collected material.

Rzebik-Kowalska et al. (1978) collected the first

known specimens of Natterer’s Bat (Myotis uattereri) in

Iraq; nevertheless, these individuals were larger than the

locally known forms and might represent a distinct tax-

on (Hanison & Bates 1991 ). Hairison ( 1956a) discovered

Serotine Bat (Eptesicus serotinus tiircoiiiaiiiis) in north-

ern Iraq. However, this record was assigned later by Har-

rison & Bates ( 1 99 1 ) to Anatolian Serotine [E. bottae aiia-

toliciis (= E. aiiatoliciis)] (Juste et al. 2013). Another in-

dividual collected in Baghdad and initially identified as

Northern Bat {E. nilssoiii nilssoiii) by Hatt ( 1 959) was sub-

sequently assigned to E. bottae (Harrison 1972; Hanison

& Bates 1991 ). Hatt ( 1959) also reported the occurrence

of the Hingston’s Serotine Bat (E. sodalis hiiigstoiii),

which was considered later on as synonym of E. bottae

(Harrison & Bates 1991 ). Hanison ( 1964) considered the

Sind Serotine Bat (E. iiasutus) and Wall’s Serotine Bat (E.

walli) as synonymous. Records from Iraq of E. walli and

its synonym, Vespertilio uiatschiei pelliiceiis, are report-

ed as E. Iiasutus (= Rhyiieptesiciis iiasutus) by Hanison

& Bates ( 1991 ). Similarly, previous records of the Cox’s

Pipistrel [Pipistrelliis co.xi) are now referred to Ruppell’s

Pipistrelle {R rueppellii), while the Schreiber’s Bat or

Long-winged Bat (Miiiiopteriis schreibersi pidcher) de-

scribed by Hanison (1956) is considered to be synonym

of M. s. pallidiis (= M. pallidus) by Harrison & Bates

( 1991 ). Finally, Metaxas ( 1891 ) reported the occurrence

of both Particolored ( Vespevtilio iiiiiriiius) and Long-eared

(Plecotus auritiis) Bat in Iraq, but Hatt ( 1959) could not

confirm these records.

Carnivora

According to Lawrence ( 1956) both Iranian {Cants lupus

pallipes) and Arabian (C. /. arabs) Wolf did occur in Iraq,

where a zone of hybridization between the two subspecies

is known in the western part of the counti'y (Harrison &

Bates 1991 ). Very recently, the Arabian Wolf was record-

ed in the extreme western Iraq by Al-Sheikhly (2012b).

Al-Sheikhly (2012) recorded the Syrian Jackal (Cants au-

reus syriaciis) for the first time in western Iraq. Similar-

ly, the Indian Grey Mongoose (Herpestes edwardsi) was

recently obseiwed for the first time in northern Iraq. This

record represents a large extension of the species range

in Arabia (Al-Sheikhly & Mallon 2013).

The Red Fox ( Vulpes viilpes) observed in the deserts of

Iraq and assigned by Hatt ( 1959) to the Indian subspecies

( V. V. piisilliis) is now listed as Arabian Fox ( V.v. arabica)

according to Harrison & Bates ( 1991 ). Captured individ-

uals and pictures taken by means of stealth camera traps

referring to Ruppell’s Sand Fox (Vulpes riieppelli) were

published in the Al-Assad au Natural (newsletter of the

37 L‘ Sustainment Brigade of the Ohio Army National

Guard, Al-Assad base, Anbar Province- N-SB). Accord-

ing to the morphological traits of the animals reported in

Al-Assad au Natural newsletter, it seems much more like-

ly that the reported Sand Foxes were in fact Arabian Red

Foxes (V V. arabica). However, V. riieppelli was report-

ed once at Khan Al-Baghdadi in western Iraq by Al-As-

sad au Natural (2009).

Mahdi & George ( 1969) reported the likely occuiTence

of the Fennec Fox (V zerda) in Iraq. Al-Robaae (1982)

collected an individual near Jabal Sanam (c. 30 km south

west of Basra, extreme southern Iraq). The same author

also reported that an individual was captured alive near

Jabal Hafit (United Arab Emirates) and kept at the Al-Ain

Zoological Garden. Later on, Gasperetti et al. (1985)

proved that the Jabal Hatlt fox was in fact V. riieppellii.

Bonn zoological Bulletin 64 (1 ): 33-58

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36

Omar F. Al-Sheikhly et al.

The Middle East Badger (Meles canescens) was shown

recently to represent a separate species distinguishable

from its closest relative, the European Badger (Meles

meles) by a combination of cranial and dental characters

(Abramov & Puzachenko 2013).

The Smooth-coated Otter [{Liitrogale perspicillata

(the endemic subspecies: L. p. maxwelli)} was believed to

be extinct in Iraq by the 1990s. However, during field sur-

veys earned out in 2007-2012, Al-Sheikhly & Nader

(2013) proved the occun-ence of the species in the south-

ern marshes.

Finally, whereas the endangered Persian Eeopard (Pan-

thera parchis saxicolor) has been recently recorded in

many localities of northern Iraq (Kurdistan), there has been

no recent sighting of the Asiatic Cheetah (Aciuonyx jiiha-

tiis venaticus) since Hatt’s (1959) report; hence, the lat-

ter is considered as virtually extinct in Iraq.

Artiodactyla

Details about the Gazelles complex and their distribution

in Iraq was reported by Al-Sheikhly (2012c). However,

further investigation is required to update the distribution

of both the Arabian Sand Gazelle (Gazella subgiitturosa

marica) and Persian Gazelle (G. s. siihguttiirosa) in Iraq.

Lagomorpha

With reference to the status of the Hare in Iraq, Hatt ( 1 959)

regarded coivwri as subspecies of the European Hare (Le-

pus ewopaeus). However, according to some genetic in-

vestigations Harrison & Bates (1991) considered the Eu-

ropean and Cape Hare as closely related taxa, and regard-

ed cowwri as subspecies of the Cape Hare (Lepus capeii-

sis). Sanborn (1940) tentatively identified hares collect-

ed in Baghdad and Camp Rashid as European Hare (L. eu-

ropaeiis couuori), which is now regarded as synonym of

L. c. counori. Hubbard ( 1955) reported the occuiTence of

Lepus babylomeus in Baghdad and Hi 11a, a taxon now con-

sidered as synonym of L. capensis. The Syrian subspecies

of the Cape Hare (L. c. syriacus) was recently recorded

(April 2012) c. 17 km to the north west of Rutba (Anbar

Province) near the Iraqi-Syrian borders (Al-Sheikhly, in

litt.).

Rodentia

Hatt (1959) included the Persian Squirrel (Sciunis anom-

aliis anomaliis) among the Iraqi rodents with comparative-

ly deep red-coloured tails; nevertheless, these records are

now considered as referable to the Caucasian SquiiTel (5.

a. pallescens) (Harrison 1956a; Harrison & Bates 1991).

Khajuria (1981) assigned two rats from Al-Quma (Bas-

ra Province) to a new taxon under genus Eiythronesokia,

but shown to be a distinctive species of Nesokia. Long-

tailed Nesokia (Nesokia bunnii) by Al-Robaae & Felten

(1990). However, later on Hamson & Bates (1991 ) sug-

gested that these individuals could provisionally be re-

ferred to as Short-tailed Bandicoot Rat (Nesokia indica).

The earlier records of the Wagner’s Gerbil [Gerbillus

(Heudecapleiaxi) dasyurus)] mentioned by Cabrera

(1901), Cheesman (1920), and Hatt (1959) are now con-

sidered referable to Harrison’s Gerbil [G. (H.)

mesopotamiae)] (Hamson & Bates 1991). Similarly,

Hatt’s (1959) records of the Turkish Jird (Meriones black-

leri blackleh) are now assigned to the Tristram’s Jird [M.

(M.) tristrami)) by Harrison & Bates ( 1991 ).

The presence of the Afghan Mole Vole (Ellobius fus-

cocapillus) in Iraq is uncertain as it does not occur today

in the south Levant, but fossils document its fonner pres-

ence during Pleistocene, about 220,000-110,000 and

80,000-60,000 years ago (Tchemov 1992; 1994). Hatt

(1959) suggested that Afghan Mole Vole might still be

present in the mountains of Kurdistan. Some fossils found

in a 25,000 years old deposit (Cave of Hazar Merd) as well

as other remains from an older deposit (Shanidar Cave)

were assigned by Bates ( 1930) to E. fitscocapillus. Bal-

tazard & Seydian (1960) reported the presence of the

species in the locality of Penjwin; however, Petter (1961)

excluded such an occuiTence. The Iraqi records of E. fus-

cocapilliis are now considered referable to the Transcau-

casian Mole Vole [Ellobius (Afgauomys) liitescens],

Hatt’s (1959) records of the Persian Vole (Microtus irani)

regarded as Social Vole [Microtus (Microtus) socialist

(Hamson 1972; Hamson & Bates 1991). Harrison &

Bates ( 1991 ) included giientheri and irani under Social

Vole M. socialis. However, M. irani has been separated

as distinct taxon (Kefelioglu & Krystufek 1999; Golen-

ishchev et al. 2002; Krystufek & Kefelioglu 2002).

Cetacea

Most of Iraq marine mammals’ records were provided by

Al-Robaae ( 1969, 1970, 1974, 1975). However, further in-

vestigation is needed to reliably describe the status of this

order in Iraq.

MATERIAL AND METHODS

Evidence for both occurrence and distribution of the Iraqi

mammals were obtained from field observations carried

out in 2013 and 2014, and combined with the following

literature records (listed according to the year of publish-

ing): Layard ( 1 852); Metaxas (1891); Cabrera & Viaje Del

Sr (1901); Wettstein (1913); Lydekker & Blaine (1914);

Wigram & Wigram (1914); Kinnear (1916); Thomas

(1919a, b); Cheesman (1920); Capper (1921); Pitman

(1922); Cheesman (1923); Cheesman & Hinton (1923);

Musil (1927); Corkill (1929); Bate (1930); Pocock

(1930); Corkill (1930); Pocock (1934); Dollman &

Burlace (1935); Pocock (1936,1938); Heptner (1940);

Bonn zoological Bulletin 64 ( 1 ): 33-58

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Checklist of the Mammals of Iraq

37

Sanborn (1940); Pocock (1941,1946); Ellerman (1947,

1948); Drower (1949); Ellennan & Morrison-Scott

( 1951 ); Felid (1951, 1954); Page (1954); Thesiger ( 1954);

Felid (1955); Hubbard (1955); Weber (1955); Harrison

(1956a, b, c); Sanborn ( 1956); Hayman ( 1957); Maxwell

(1957); Petter (1957); Hoogstraal & Kaiser ( 1958); Reed

(1958); Bodenheimer (1959); Harrison (1959); Hatt

(1959); Maxwell (1960); Baltazard & Seydian ( 1960); Pet-

ter ( 1961 ), Harrison ( 1964); Thesiger ( 1 964); Al-Robaae

( 1 966); Von Lehmann ( 1 966); Groves & Hamson ( 1 967);

Mahdi ( 1967); Al-Robaae ( 1968); Harrison ( 1968); Al-

Robaae (1969); Mahdi & George ( 1969); Nader (1969);

Al-Robaae ( 1970); Nasher ( 1970); Felten ( 1971 ); Nader

(1971); Nadler et al.(1971); Andera (1972); Harrison

( 1972); Al-Robaae (1974, 1975); Kadhim ( 1975); Thalen

(1975); Al-Robaae (1976); Niazi (1976); Al-Robaae

(1977); Jawdat (1977); Kadhim et al. (1977); Bhatnager

& El-Azawi ( 1978); Nadachowski et al. (1978); Rzebik-

Kowalska et al. (1978); Shamsuddin & Mohammad

(1978); Young (1978); Darviche et al. (1979); Kadhim

(1979); Kadhim et al.(l979); Nader (1979); Matskasi

(1980); Jawdat & Mahmoud ( 1981 ); Kadhim ( 1981 ); Kha-

juria ( 1981 ); Al-Robaae ( 1982); Al-Shenawi et al. ( 1 982);

Kadhim & Mustafa (1983); Nader & Kock (1983); Al-

Khalili (1984); Gasperetti et al. (1985); Kadhim & Wahid

(1986); Kadhim et al.(1986); Nader (1968 a; b); Groves

(1988); Hutterer & Harrison (1988); Khajuria (1988);

Kadhim et al. (1989); Kock ( 1990); Al-Robaae & Felten

(1990); Nadachowski et al. (1990); Nader & Kock

( 1990a); Harrison & Bates ( 1991 ); Kadhim ( 1 997, 1998);

Kock et al. (2001 ); McLellan & Garshelis (2006); Haba

(2009); Mallon & Kingswood (2001); Al-Sheikhly

(2012b, c); Co§kuna et al. (2012); Omer et al. (2012); Raza

et al. (2012); Abass (2013); Al-Sheikhly & Nader (2013);

Al-Sheikhly & Mallon (2013); Haba (2013); Lahony et

al. (2013); Mohammad et al. (2013); Raza (2013); Al-

Sheikhly et al. (2014); Al-Sheikhly et al. (2015). We al-

so relied our work on mammal collections resident to: Bi-

ological Research Centre, Baghdad, Iraq (BRCB); Bom-

bay Natural History Society, Mumbai, India (BNHS);

Field Museum of Natural History, Chicago, Illinois, USA

(FMNH); Harrison Institute (formerly Hamson Zoologi-

cal Museum), Sevenoaks, Kent, UK (HZM); Iraqi Natu-

ral History Research Centre and Museum, Baghdad, Iraq

(BRCB); Museum National d’Histoire Naturelle, Paris,

France (PM); collection of the University of Michigan,

Michigan, USA (UMC); Natural History Museum, Lon-

don, UK (BMNH); Hungarian Natural History Museum

- Hungary (HNHM).

Recent records were derived from direct visual obser-

vations of live animals in the wild or in captivity, dead

specimens (hunted or trapped), and investigation of study

skins or mounted specimens, tracks, scats, field signs, calls

and vocalizations. Reports and photographs from mem-

bers of the Iraqi Hunters Association (IHA) as well as in-

terview with local people were also considered. In addi-

tion, Baghdad Zoological Garden (BZG) and local animal

markets were visited. Photo-identification was carried out

by means of an EOS 40D and EOS IDs Mark II Canon

digital SLR camera bodies attached to a Canon EF 400

mm F5.6L USM telephoto lens and Bushnell IR remote

camera traps. The identification of the individuals collect-

ed during this study was based on Hairison & Bates ( 1991 )

and Aulagnier et al. (2008). The orders, families, genera,

species and subspecies were listed in the order they ap-

pear in the 3"“^ edition of Wilson & Reeder’s (2005). Dur-

ring 2013-2014 several recent records (RR) were made

and presented in the text as the following: a single record

made in Anbar Province in 20 1 3 (An- 1 3 ); multiple records

made in Basra Province in 2013 and 2014 (Ba-13; 14) and

so on.

RESULTS

Systematic list of the species

Order Erinaceomorpha

Family Erinaceidae

Southern White-Breasted Hedgehog Erinaceiis concolor

(Martin, 1838)

Subspecies: C. c. couco/or (Martin, 1838).

Distribution: Mountains, forests, and grassy steppes of

northem Iraq. Recorded from: Hais 10 km NW Ser Ama-

dia (Hanison 1956a); Banaman (Hatt 1959).

RR: Bradost Mountain (Er-14).

Long-eared Hedgehog Hemiechimis auritus (Gmelin,

1770)

Subspecies: H. a. calligoni (Satunin, 1901 ).

Distribution: Arid steppes, marshes, and riparians along

the Tigris and Euphrates Rivers. Recorded from: Amara

and Basra (Cheesman 1920); Hit, Al-Falluja, and near

Samarra (Pitman 1922); Baghdad (Sanborn 1940; Hub-

bard 1955; Weber 1955); Mansuriya al Shatt (Hatt 1959;

Nader 1968b; Martin & Nader 1975; Bhatnager & El-Aza-

wi 1978; Shenawi et al. 1982); Chebaeish (Haba 2009);

Jabal Makhool, AFAIam, Hussaiba, and 3 km from Al-

Qa-eem (Al-Sheikhly 2012); Hammar Marsh (Abass

2013).

RR: Jadrriyah and Umm Al-Khanazeer Island, Yarmook,

and Dor’ah (B-13); Hammar Marsh (Ba-14).

Desert Hedgehog Paraechimts aethiopicus (Ehrenberg,

1832)

Subspecies: P. a. litdlowi (Thomas, 1919).

Distribution: Arid deserts and steppes of central and south-

ern Iraq. Recorded from: Qasral Helqum (Hairison 1959),

40 km E Rutba (Nadachowski et al. 1990), Hadithah and

Bonn zoological Bulletin 64 ( 1 ): 33-58

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38

Omar F. Al-Sheikhly et al.

Baghdad (Hatt 1959), Hit (type locality of P. a. htdlowi)

and Hamam Al-Alil (Bhatnager & El-Azawi 1978).

RR: Rutba (An-2013).

Order Soricomorpha

Family Soricidae

Asian House Shrew Sunctis minimis (Linnaeus, 1766)

Subspecies: No named subspecies; Iraq population like-

ly originate from north India.

Distribution: Introduced to extreme southern Iraq. Record-

ed from: Al-Qurna (Cheesman 1920); Basra (Hatt 1959);

Chebaeish (Haba 2009); specimens collected in Fao are

resident to the BMNH collection; Hammar Marsh (Abass

2013).

RR: Al-Qurna (Ba-13); Hammar Marsh (Ba-14).

Etruscan Shrew Siuiciis etriisciis (Savi, 1822)

Subspecies: S. e. etnisciis (Savi, 1822).

Distribution: Bushland, cultivated fields, and marshs of

southem and central Iraq. Recorded from: Basra and Ama-

ra (Cheesman 1920); Al-Falluja (Pitman 1922); Kut al

Imaira (Hatt 1 959); Habbaniya and the eastern bank of the

Euphrates River (Hanison 1959; Harrison & Bates

1991 ); Hammar Marsh (Abass 2013).

Lesser White-toothed Shrew Crocidwa siiaveoleiis (Pal-

las, 1811)

Subspecies: Likely C. 5. portali (Thomas, 1920) sub-

species, with individual variation in the colour of pelage.

Distribution: Mountains and steppes of northern Iraq.

Recorded from: Bebaidi and Sarsank (Hamson 1956a;

Hatt 1959); specimens from Cora and Almawan original-

ly listed by Hatt (1959) as Bicolored White- toothed Shrew

(C leiicodon persica), which was referred later to this

species by Hutterer & Harrison (1988); the vicinity of

Mousel (Andera 1972; Hutterer & Hamson 1988).

RR: Safen Mountain (Er-14).

Order Chiroptera

Family Rliinopomatidae

Lesser Mouse-tailed Bat Rhiiiopomci hardwickH (Gray,

1831)

Subspecies: R. h. arahiiim (Thomas, 1913).

Distribution: Apparently rare, restricted to the caverns in

the arid steppes of western and central Iraq. Recorded

from: 19 km E Hilla by Sanborn (1956); near Haditha

(Hatt 1959; Nader 1971). A colony of 83 individuals were

found roosting in a small cave at Sheikli Hadid sink hole

near Haditha (Al-Sheikhly 2012).

Greater Mouse-tailed Bat Rhiuopoma microplivllitm

(Briinnich, 1782)

Distribution: Uncertain. Recorded by Khajuria (1988)

from Haditha in western Iraq as a distinct species

{Rhiuopoma hadithaensis), but regarded as a junior syn-

onym of the Greater Mouse-tailed Bat (R. microphylhim)

by Kock et al. (2001 ).

Family: Emballonuridae

Naked -rumped Tomb Bat Taphozous (Liponycteris) midi-

veiitris (Cretzschmar, 1830)

Subspecies: T. ii. magiiiis {WettsXein, 1913).

Distribution: Widespread, mainly along the Tigris and Eu-

phrates Rivers. Recorded from: banks of Euphrates Riv-

er (Kinnear 1916); Amara, Shaiba, Ctesiphon, and Basra

(Cheesman 1920); Baghdad and Diltawa (Sanborn 1940);

Ctesiphon arch (Weber 1955); between Habbaniya and Ar

Ramadi, Habbaniya, and at Shaiba (Harrison 1956b);

Aqqar-Quf, Baghdad, Fao, and Najaf (Harrison 1964);

recorded also by Al-Robaae ( 1968), Shamsuddin & Mo-

hammad ( 1978), Nadachowski et al. ( 1978), Al-Shenawi

et al. ( 1982) and Haba (2009).

RR: Large colonies roosting in cracks of old buildings at

Chebaeish (Th-14).

Family: Rhinolophidae (Horseshoe bats)

Greater Horseshoe Bat Rhiiwlophiis fernimeqidmtm

(Schreber, 1774)

Subspecies: R. f. iraiii (Cheesman, 1921 ), with distinctive

pale pelage.

Distribution: Limestone caverns and suitable retreats of

northern and central Iraq. According to Hatt (1959) the

species was reported to occur in Iraq only by Metaxas

(1891) and Kinnear (1916). Recorded from: 130 km W

Ramadi (Niazi 1976); Salahadin and Erbil Liwa (Nader

& Kock 1983); a single specimen from Tang-i-dorg in

Duhok is kept in the collection of the BMNH (Harrison

1964). Al-Robaae ( 1976) has tentatively identified some

specimens from Mousel (Hamson & Bates 1991).

Lesser Horseshoe Bat Rhinolophiis hipposideros (Bech-

stein, 1800)

Subspecies: R. h. midas (Anderson, 1905).

Distribution: Three specimens from Baghdad were report-

ed by Cheesman (1920); this represents the only record

for Iraq (Hatt 1959; Hamson & Bates 1991).

Mediterranean Horseshoe Bat Rhinolophiis einyale (Bla-

sius, 1853)

Subspecies: The Iraqi population can not be assigned to

any named subspecies.

Distribution: Previous data of the species (Niazi 1976) rep-

resents R. mehelyi (see remarks below). The only definite

record is from Bjil village, Akre District in northern Iraq

(Al-Sheikhly et al. 2015).

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Checklist of the Mammals of Iraq

39

Mehely’s Horseshoe Bat Rhiuolophiis mehelyi (Matsch-

ie, 1901)

Subspecies: R. m. mehelyi {Matschie, 1901).

Distribution: Caves of arid areas in central and western

Iraq. Niazi ( 1967) reported the occuiTence of the Mediter-

ranean Horseshoe Bat (R. eiuyale) in two localities; how-

ever, the species was proved to be in fact R. mehelyi (Har-

rison & Bates 1991, Al-Sheikhly et al. 2015). Recorded

from: 43 km SW Ash-Shinaffya and 130 km W Ramadi;

specimens collected in Al-Alfasia Cave (8 km S Haditha)

are resident to the BMNH (Hanison & Bates 1991). Spec-

imens collected in El Hashfe and Hadhita areas of Iraq

(Matskasi 1980) are kept in the collection of HNHM and

those of collected in Bjil village, Akre District in north-

ern Iraq will be deposited in BRCB.

RR: Three individuals were found roosting in a tunnel near

Mawat (Su-14).

Family: Hipposideridae

Geoffrey’s Trident Leaf- nosed Bat Asellia trideus (E. Ge-

offroy, 1813)

Subspecies: A. t. murraiami (Anderson, 1881).

Distribution: Common in deserts and arid steppes and

wadis of central Iraq. Recorded from: Mousel (Wettstein

191 3); Al-Falluja and “Lake” Aqar Quf (Cheesman 1 920);

Baghdad (Sanborn 1940; Hubbard 1955); A I Qa-eem near

the Syrian border (Weber 1955); between Ar Ramadi and

Habbaniya and from Uqhaidhir (Hamson 1 956); Mustan-

siriyah in Baghdad (Hatt 1959); recorded at many local-

ities by Al-Robaae (1966), Shamsuddin & Mohammad

( 1978), and Nadachowski et al. ( 1990); at Sheikh Hadid

sink hole near Haditha by Al-Sheikhly (2012).

Family: Molossidae

European Free-tailed Bat Tadarida teiiiotis (Rafmesque,

1814)

Subspecies: T. t. nieppelli (Temminck, 1826), with pale

grey pelage colour.

Distribution: Uncommon, confined to the rocky teirain and

crevices in western Iraq. Recorded from: Between Ar Ra-

madi and Habbaniya (Hanison 1956b; Hamson & Bates

1991).

Family: Vespertilionidae

Lesser Mouse-eared Myotis Myotis blythii (Tomes, 1 857)

Subspecies: M. h. o/nnr/ (Thomas, 1906).

Distribution: Confined to the caverns of northern moun-

tains. Recorded from: Hazar Mard caves (Bate 1930; Hatt

1959); Diyana and near Baradust (Sanborn 1940; Hani-

son & Bates 1991 ).

Long-fingered Myotis Mvotis capaccinii (Bonaparte,

1837)

Subspecies: M. c. hureschi (Heinrich, 1936).

Distribution: Uncertain; a male individual reported from

the ruined city of Kish: this is the only confimied occur-

rence (Sanborn 1956; Hatt 1959; Harrison 1964; Hani-

son & Bates 1991 ).

Geoffroy’s Myotis Mvotis emarginatiis (E. Geoffroy,

1806)

Subspecies: As no voucher specimens exist, the Iraqi pop-

ulation can not be assigned to any named subspecies.

Distribution: Niazi (1976: 172) reported the occurrence

of this species in the mid-west part of Iraq but no exact

locality or any detailed description was given. Two indi-

viduals were observed and photographed in Bjil village

(Akre District, northern Iraq): this represents the only

record for Iraq (Al-Sheikhly et al. 2015).

Natterer’s Myotis Myotis nattereri (Kuhl, 1817)

Subspecies: Iraqi individuals investigated by Rzebik-

Kowalska et al. ( 1978) were clearly smaller than the ge-

ographically closest and related taxon M. nattereri arax-

eniis (= Myotis schauhi) but differed in certain cranio-den-

tal traits from the other subspecies of M. nattereri. These

individuals could be representative of a distinct taxon.

Distribution: Confined to the rocky caverns of the moun-

tains of northern Iraq. Recorded from Gali Ali Beg, which

is the only confirmed occurrence (Rzebik-Kowalska et al.

1978; Harrison & Bates 1991).

Botta’s Serotine Eptesiciis hottae (Peters, 1869)

Subspecies: E. h. hingstoni (Thomas, 1919), restricted to

central and southern Iraq;

Distribution: Widespread. Recorded from: Basra, Bagh-

dad, Nasiriyah, Karbal’a (Hatt 1959); Habbaniya (Harri-

son 1956b); Ramadi and Sulaymaniyah (Felten 1971 ); 35

km N An-Najaf, Al-Mahmudiya, Baqubah, and Al-Khalis

(Nader & Kock 1990a).

Anatolian Serotine Eptesiciis anatolicus (Felten, 1971 )

The Anatolian Serotine, fonnerly regarded as subspecies

of E. bottae has recently been elevated to the rank of

species on the basis of combined evidences derived from

cranial moiphology, molecular DNA markers, and echolo-

cation characteristics (Juste et al. 2013).

Distribution: Confined to the mountains and foothills of

central and northern Iraq. Recorded from: Ser Amadia and

Kani Mase (Harrison 1956a; Harrison & Bates 1991 ); Al-

Alam (Al-Sheikhly 2012).

Sind Serotine Rhyneptesiciis nasiitiis (Dobson, 1877)

Subspecies: R. n. pelliicens (Thomas, 1906). Rhyneptesi-

ens was re-elevated as a separate genus by Juste et al.

(2013) on craniodental and phylogenetic grounds.

Distribution: Inhabits ruins and old buildings of central

and southern Iraq. Recorded from: Zubeir and Shaiba as

Eptesiciis n. pelliicens (Harrison 1964); Basra and

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Omar F. Al-Sheikhly et al.

Fig. 2. Wolf hunting in Iraq, Photo © Hameed A. Al-Habash/ Iraqi Hunting Association.

Nasiriya (Sanborn 1940); Qalat Salih and Al Miqdadiyah

(Nader 1971; Hamson & Bates 1991).

Kiihl’s Pipistrelle Pipistrellus (Pipistrelliis) kiihlii (Kuhl,

1817)

Subspecies; P. k. ikhwanius (Cheesman & Hinton, 1924),

with pale desert pelage.

Distribution: Abundant and ubiquitous. Recorded from:

Karbala, Baghdad, Mousel, Qalat Sharqat, Mejadin, and

Bahrka (Wettstein 1913); Amara, Baghdad, and Sheik

Saad (Cheesman 1920); Baghdad, Amara marshes. Shaikh

Falih as Saihud’s camp, A 1 Halfayah, Balad, Sinjar, Badra,

Diltawa, An Nasiriya, Karbala, and Rustamiyah Fann near

Baghdad (Sanborn 1940); Baghdad, Al Qa-eem, the

pipeline station T-1 and in Dhibban near Habbaniya (We-

ber 1955); Shaiba, Ramadi marshes, south ofAmiriya, Ra-

madi-Habbaniya, Basra, Al Falluja, and Uqhaidhir (Har-

rison 1956b); Baghdad (Hatt 1959); Specimens from Fao,

Kut as Sayyid, Basra, Khaniqin, Wadia south of Majar al

Kabir of Amara, and from Ain Zalah 45 km S Mousel res-

ident to the collection of the BMNH (Hamsonl964);

recorded also by Shamsuddin & Mohammad ( 1978) and

Al-Shenawi et al. ( 1982).

RR: Ramadi (An- 1 3); Al-Adhamiyah; Jadriyah; Al-

Yannook (B-13); Gamiat Ali and Hammar Marsh (Ba-14).

RiippelFs Pipistrelle Vcmsonia riieppellii (Fischer, 1829)

Subspecies: V. r. coxi (Thomas, 1919), with sandy-buff

dorsal pelage. Vcmsonia was elevated to genus level by

Koubinova et al. (2013) based on genetic data.

Distribution: Uncommon, inhabits settlements and ruins

of central and southern Iraq. Recorded from: Al-Ad-

hamiyah (Nader 1971 ); Basra and at Beit Sheikh Mahom-

mad at the marshes of Chahala in Amara: these specimens

are resident to BMNH collection (Cheesman 1920; Hatt

1959; Hamson 1964); Agergoof (Harrison & Bates 1991 ).

Hemprich’s Desert Bat Otonycteris hemprichii (Peters,

1859)

Subspecies: O. h. petersi (Anderson & de Winton, 1902),

in the eastern Euphrates basin and in central and south-

ern Iraq; O. h. jin (Cheesman & Hinton, 1924), confined

to the desert habitat of both extreme western and south-

western Iraq.

Distribution: Local, confined to arid steppes and rocky

deserts of central and southern Iraq. Recorded from: Fao

(Kinnear 1916) as well as by Ellerman & Morrison-Scott

(1951), who listed the taxon as O. petersi, probably a syn-

onym of hemprichi (Hatt 1959); Chamchamal-Jamio val-

ley in Kurkuk Liwa (Nader & Kock 1983; Hairison &

Bates 1991); Al-Cidr and Himreen foothills (Al-Sheikhly

2012).

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Checklist of the Mammals of Iraq

41

Family: Miniopteridae

Pallid Long-fingered Bat Miniopterus paUidus (Thomas,

1907)

Recorded as Miniopterus schreihersii (Kuhl, 1 8 1 7 ) by Halt

( 1959) and Harrison & Bates ( 1991 ). Mitochondrial and

nuclear DNA studies corroborated by slight variations in

size, wing shape, and echolocation call parameters sug-

gested that M. pallidus should be considered as full species

(Furman et al. 2010a, b).

Distribution: Uncertain, confined to the mountain caves

of the extreme northern Iraq. Reported from: Ser Amadia

(type locality of M. p. piilcher. Harrison 1956), the only

record for Iraq (Harrison 1964; Harrison & Bates 1991 ).

Order Carnivora

Family: Canidae

Golden Jackal Canis aureus (Linnaeus, 1758)

Subspecies: C. a. aureus (Linnaeus, 1758), along the Tigris

and Euphrates Rivers; C. a. syriaciis (Hemprich & Ehren-

berg, 1833), restricted to the desert and arid lands of the

extreme western Iraq, yet probably present also in north-

ern Iraq.

Distribution: Abundant along the river riparians, cultivat-

ed land, marshes, mountains and arid lands. Canis a. au-

reus recorded from: Babylon (Jebb 1908); Sheik Saad

(Cheesman 1920); Jezira, c. 40 km W Mousel (Boden-

heimer 1959); Habbaniya, Al Falluja and Saman'a (Har-

rison 1959); Basra, Diyala and Hilla (Halt 1959); the

marshes near Majar (Thesiger 1964); Chebaeish (Haba

2009); Hammar Marsh (Abass 2013); Hawraman Moun-

tain (Lahony et al. 2013). Specimens collected in Legait,

Shahraban, Baghdad, Amara, near Kut, Shaiba, Fao and

Saqiawiya are included in the collection of the BMNH

(Hanison & Bates 1991 ). Canis a. syriacus recorded from:

Rutba and Hussaiba (Al-Sheikhly 2012).

RR: Canis a. aureus, abundant in Al-Alam ( Sa- 1 3 ); Anah;

Hit (An- 13); Mashab and Hammar Marsh (Ba-14).

Gray Wolf Canis lupus (Linnaeus, 1758)

Subspecies: C. /. pallipes (Sykes, 1831), confined to the

northern mountains and on eastern bank of the Euphrates

River; C. /. arahs (Pocock, 1934), restricted to the arid

steppes and desert of western and south-western Iraq.

Distribution: Widespread, Canis 1. pallipes recorded from:

Al-Shaiba and At Tanumah (Cheesman 1 920); Ur (Pocock

1935); Rawanduz, Diyala, Sulaymaniyah, and Khaniqin

(Sanborn 1940); Sinjar, Karadagh and umm al-Kahab

Wells (Felid 1954); Erbil Liwa, near Jarmo, east Kurkuk

Liwa, Lake Habbaniya and near Haditha (Hatt 1959); Jazi-

ra, c. 40 km W Mousel (Bodenheimer 1959); near Majar

(Thesiger 1964); Central and Hammar Marsh (Haba

2009); Makhmore, Jabal Himreen, Jabal Makhool, and

Wadi Al-Tharthar (Al-Sheikhly 2012); Hammar Marsh

(Abass 2013). Canis 1. arahs recorded from: Rutba (Al-

Sheikhly 2012b).

RR: Canis 1. pallipes, Jabal Himreen (Di- 13; 14) (Fig. 2).

Red Fox Vulpes vulpes (Linnaeus, 1758)

Subspecies: V. v. arahica (Thomas, 1902), around marsh-

es, deserts, and arid steppes of central and southern Iraq;

Vulpes r: flavescens (Gray, 1843), restricted to the

foothills and mountains of northem Iraq.

Distribution: Widespread. Vulpes i’. arahica. Recorded

from: Qalat Luqait, Shatt al Adhaim, and Basra (Cheesman

1920); Hilla, Baghdad, and Baquba (Hatt 1959);

Saqiawiya, Tikrit, Fort Ukhaidir, Haur al Hasa, Falluja,

and Tar al Hazar north west of Iraq ( Hairison 1968). Spec-

imens collected in Baghdad, Falluja, and Ctesiphon are

included in BMNH collection (Handson & Bates 1991);

Hammar Marsh (Abass 2013); Hawraman Mountain (La-

hony et al. 2013). Vulpes v. flavescens recorded from:

Kurkuk, Rawanduz and Jarmo (Hatt 1959); near Amadia

(Hairison 1959), Himreen foothills (Al-Sheikhly 2012).

RR: Vulpes v. arahica recorded from Jadriyah and-

Yarmook districts (Ba-13); Hila and Dalmaj (Qa-

\M-Vulpes V. flavescens recorded from: Ramadi (An- 13);

Jabal Himreen (Sa-13).

RiippeH’s Fox Vulpes rueppellii (Schinz, 1825)

Subspecies: V. r sahaea (Pocock, 1934).

Distribution: Restricted to the arid steppes and desert of

western and south-western Iraq. Recorded from: Haditha

(Hatt 1959); Khan Al-Baghdadi (N-SB 2008); Massad Al-

Rutba, Hussaiba, Ga’ara, Wadi Horan ( 10 km E Hadithah),

Nikheab, and Khan Al-Baghdadi (Al-Sheikhly 2012).

RR: c. 30 km south of Rahaliya - Karbal’a desert (An-

14) (Fig. 3).

Fig. 3. Adult male RiippePs Fox Vulpes rueppellii, a carni-

vore confined to the desert habitat in western Iraq. Photo © Omar

F. Al-Sheikhly 2014.

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Omar F. Al-Sheikhly et al.

Fennec FoxVulpes zerda (Zimmermann, 1780)

Subspecies: V. z. zerda (Zimmemrann, 1780).

Distribution: Uncertain. Mahdi & George (1969) report-

ed the likely occuiTence of this species in Iraq. Al-Robaae

( 1982) collected an individual near Jabal Sanam SW of

Basra, extreme southern Iraq.

Family: Ursidae

Brown Bear Ursiis arctos (Linnaeus, 1758)

Subspecies: U. a. syriaciis (Hemprich & Ehrenberg, 1828).

While bears in northern Iraq show brown-greyish pelage

those inhabiting the district of Jilu are known for their pale

fur. Nevertheless, individual variation and/or seasonal

change in the pelage colour may occur.

Distribution: Confined to the mountains of extreme north-

ern Iraq. Recorded from: Iraqi-Turkish frontier region

( Wigram & Wigram 1914); Bradost Mountain north of Er-

bil Liwa, east of Sulaymaniyah (Halt 1959); north east of

Kurkuk (McLellan & Garshelis 2006); QaraDaq (Raza

2013).

RR: An adult bear was killed by locals nearby Duhok (Du-

13); a specimen from the Zagros Mountain is included in

the collection of the BRCB; two adult males collected in

Haji Omran are presently kept at BZG.

Family: Mustelidae

Beech Marten Martes foina (Erxleben, 1777)

Subspecies: M. f. syriaca (Nehring, 1902).

Distribution: Confined to the mountains of northern Iraq.

Recorded from: Iraq-Iran border (Cheesman 1920);

Rawanduz (Sanbom 1940); Bradost Mountain (Field

1955); Narwa (Hamson 1956a); Hawraman Mountain

(Lahony et al. 2013).

Marbled Polecat Vonuela peregusna {GiddenstMt, 1770)

Subspecies; V. p. syriaca (Pocock, 1936), with individual

variations in pelage color and pattern.

Distribution: Restricted to the high grounds of Mousel.

Recorded from: c. 64 km S Mousel (Pocock 1936);

Mousel (Halt 1959); 45 kmN Mousel (Allouse 1954; Har-

rison 1968; Hamson & Bates 1991); Hawraman Moun-

tain (Lahony et al. 2013).

Least Weasel Miistela nivalis (Linnaeus, 1766)

Subspecies: Probably M. n. boccamela (Bechstein, 1800).

Distribution: Confined to the mountains of northern Iraq.

Hatt (1959) did not find any definitive occun'ence of the

species, but he received hunters’ reports where the pres-

ence of Weasel-like animals at Erbil Eiwa (Zagros Moun-

tain) was recorded. Confinned record from Shirin Moun-

tain in Barzan area (Raza 2013); Hawraman Mountain

(Lahony et al. 2013).

Middle East Badger canescens (Blanford, 1875)

Distribution: Confined to the mountains of northern Iraq,

with local distribution in central Iraq. Recorded from: Qali

Ali Beg (Sanbom 1940); Baghdad (Hatt 1959); 20 km N

Shaqlawa (Smielowski 1978); Hawraman Mountain (La-

hony et al. 2013).

RR: Barzan Area (Er-14); Chami-reazan area (Su-14).

Honey Badger Mellivora capensis (Schreber, 1776)

Subspecies: M .c. wilsoni (Cheesman, 1920).

Distribution: Widespread in the desert and arid steppes of

central and southern Iraq. Recorded from: Baksai

(Cheesman 1920); An Nuhaidam (Hamison 1959); T1 sta-

tion and Rawa (Hatt 1959); Abu Khasib (Anon 1966);

Hammar Marsh (Abass 2013). Specimens from Fuluja are

included in the collection of the BMNH (Hamison & Bates

1991).

RR: Ga’ara and Massad Al-Rutba (An- 13); Nugratt Al-

Salman (Mu- 14).

European Otter Lutra liitra (Linnaeus, 1758)

Subspecies: L. 1. seistanica (Birula, 1912).

Distribution: Lakes, streams, tributaries, and marshes of

Tigris and Euphrates Rivers. Recorded from: Amara

(Cheesman 1920); Qalat Salih (Sanbom 1940); Baghdad

and Hindiya (Hatt 1959); Central marshes, vicinity of Al-

Maimona, Musharah River, Tanniya, Al-Alam, Samama

Lake, Himreen, Khan Al-Baghdadi, Haditha, Der-

bendikhan, Dukan, Little Zab, Barzan and TaqTaq (Al-

Sheikhly & Nader 2013); recorded also in Mashab in east-

ern Hammar Marsh by Abass (2013). Specimens from

Abusakhair (South of Hor Al-Hammar Marsh) are includ-

ed in the collection of the BMNH (Hamison & Bates

1991). Recent records from Mashab River in eastern Ham-

mar Marsh (Al-Sheikhly et al. 2014).

RR: Mashab and Salal River (Ba-14); Taq Taq (Er-14).

Smooth-coated Otter Lutrogale perspicillata (I. Geoffrey

Saint-Hilaire, 1826)

Subspecies: L. p. maxwelli (Hayman, 1956).

Distribution: Confined to the dense reed beds of the south-

ern marshes; one record from northern Iraq (Kurdistan).

Recorded from: Abusakhair and Azair (Hayman 1956;

Maxwell 1957, 1960); Azair by (Hatt 1959); TaqTaq

(Omer et al. 2012); Fao-Rass Al-Beisha, Abu Al-Khasib

and Umm Al Rassas, Abu Ajaj-Hor Al-Hammar, Umm Al-

Na’aj lake-Hawizeh, and TaqTaq (Al-Sheikhly & Nader

2013; Al-Sheikhly et al. 2014).

Family: Herpestidae

Small Asian Mongoose Herpestes javanicus (E. Geoffroy

Saint-Hilaire, 1818)

Subspecies: H. a. pallipes (Blyth, 1845).

Distribution: Widespread along the Tigris and Euphrates

Rivers. Recorded from: Basra (Cheesman 1920); Amara

(Pitman 1922); Baghdad (Sanbom 1940); Hilla and

Mousel (Hatt 1959); Habbaniya (Hamdson 1959); Zeban,

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Checklist of the Mammals of Iraq

43

c. 32 km from Falluja (Harrison 1968); Chebacish (Haba

2009); many localities Iraq (Al-Sheikhly & Mallon 2013);

Al-Hammar Marsh (Abass 2013).

RR; Recorded from many sites in Al-Hammar Marsh (Ba-

14); Chebaeish (Th-14); Hawizeh (Ma-14).

Indian Grey Mongoose Herpestes edwardsi (E. Geoffroy

Saint-Hilaire, 1818)

Subspecies: H. e. fernigiueits (Blanford, 1874)

Distribution: Local, in the rocky highlands of northern and

central Iraq; however, it may be found in southern Iraq as

well. Recorded from: Qashqulai and near Al-Alam (Al-

Sheikhly & Mallon 2013).

RR: Near TaqTaq (Er-14).

Family: Hyaenidae

Striped Hyena Hyaena hyaena (Linnaeus, 1758)

Subspecies: H. /;. syriaca (Matschie, 1900). Pocock ( 1934)

suggested that H. h. syriaca may be synonym of H. h.

hyaena.

Distribution: Common in the lowlands of central and

southern Iraq; uncommon in the northern mountains.

Recorded from: Ur, Basra and Falluja (Cheesman 1920);

between Amara and Sheikh Saad (Capper 1921 ); Falluja,

Nahrwan, and Ad Dawr (Pitman 1922); Tyb River, Jabal

Hamrin, and Ur (Pocock 1941 ); Bradost Mountain (Field

1955); near Haditha (Lawrence 1956); near Samarra (Har-

rison 1959); Jezira, c. 40 km W Mousel (Bodenheimer

1959); Mansuriya al Shatt (Halt 1959); Erbil Liwa, Cte-

siphon, Chamchamal, Baghdad, Baquba, Habbaniya, Ra-

madi and Sulaymaniya (Hairison 1968); Khan Al-Bagh-

dadi (N-SB 2008; 2009); near Hammar Marsh (Haba

2009); Hammar Marsh (Abass 2013).

RR: Al-Ga’ara, Rutba (An-13).

Family: Felidae

Wild Cat Felis silvestris (Schreber, 1777)

Subspecies: F. s. nesterovi (Birula, 1916), confined to the

forests and wooded valleys of northern Iraq; Felis s. ira-

ki (Cheesman, 1921), restricted to the marshes and arid

steppes of central and southern Iraq. Distribution: Wood-

ed mountains, rooky steppes, and nearby marshes; nev-

ertheless, the species appears to be rare. Felis s. iraki was

recorded from: Sheikh Saad (Cheesman 1920); Baghdad

and Diwaniya (Halt 1959). Felis s. nesterovi was record-

ed from Nachr-Chasasch (Handson 1968; Hanison &

Bates 1991) and Peramagroon (Raza et al. 2012).

RR: Felis s. iraki near Habbaniya (An- 13); Auda Marsh

(Ma-14).

Jungle Cat Felis chans (Schreber, 1777)

Subspecies: F. c. fiirax (de, Winton, 1898), although inter-

mediation with F. c. chans (Giildenstadt, 1776) may oc-

cur.

Distribution: Widespread along the Tigris and Euphrates

Rivers. Recorded from: Amara, Qalat Salih, Madij, and

Shahraban (Cheesman 1920); between Al-Quma and Azir,

Shatt al Adhaim, and Ad Dawr (Pitman 1922); Hilla Desert

(Sanborn 1 940); Abu Ghreb, near Akkarkuf (Bodenheimer

1959); Hila and Dujail (Hatt 1959); Khan Al-Baghdadi (N-

SB 2008; 2009); Hammar Marsh (Abass 2013). Speci-

mens collected in Mousel, Baghdad, Ramadi, and Hab-

baniya are included in the collections of the BMNH and

HZM (Harrison 1968).

RR: Hammar Marsh (Ba-14).

Sand Cat Felis niargarita (Loche, 1858)

Subspecies: F. m.harrisoni (Hemmer, Grubb and Groves,

1976).

Distribution: Confined to the desert and arid steppes of

south western Iraq. Recorded from W Al-Najaf city and

Al-Jufaira oasis in the desert of Al-Najaf Province (Mo-

hammad et al. 2013).

RR: Takhadid (Mu-14) (Fig. 4).

Fig. 4. Adult male Sand Cat Felis margarita trapped in Mu-

thana Province in south-western Iraq. Photo © Omar F. Al-Shei-

khly 2014.

Caracal Caracal caracal (Schreber, 1776)

Subspecies: C. c. sclnnifzi {Matschie, 1912).

Distribution: Confined to the desert and arid steppes of

the extreme western and southern Iraq. Recorded from:

Basra (Metaxas 1891); 45 km from Rutba (Corkill 1930).

Uncertain records from southern and western deserts of

Iraq (Thalen 1975). Reported by locals from Faidhatt Al-

Massad (Al-Sheikhly 2012).

RR: Hunters’ reports from Ain Al-Faras in the Tharthar

Valley (An- 1 3).

Eurasian Lynx Lynx lynx (Linnaeus, 1758)

Subspecies: L. /. dinniki {Satunin, 1915).

Distribution: Restricted to the forested mountains of ex-

treme northern Iraq; it would appear to be rare. Record-

ed from: Zakho and Erbil Liwa (Hatt 1959; Hamson &

Bates 1991); Barzan area (Raza 2013).

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44

Omar F. Al-Sheikhly et al.

Leopard Pantheni parchis (Linnaeus, 1758)

Subspecies: P. p. saxicolor {Pocock, 1927).

Distribution: Rare, restricted to the forested mountains of

extreme northern Iraq. Recorded from: near Rawa (Musil

1927); Aqra and Rowanduz (Pocock 1930); Jabal Bradost

(Field 1955); Harir Dagh (Hatt 1959); Bamur Harrison

(1968); Murdka of Darbandikhan, Mandili, and QaraDag

(Al-Sheikhly 2012b); QaraDag (Raza et al. 2012).

RR: Suran Area (Du-13).

Cheetah Acinonyx Jubatiis (Schreber, 1775)

Subspecies: rt.y. ve)iaticiis {Gr'\tT\{h, 1821).

Distribution: Probably extinct. Recorded from: near Bu-

saiya (W Basra southem desert), K3 station, and HI and

H2 pumping stations in Syrian Desert (Corkill 1929); near

Iraq-Saudi Arabia-Jordan borders (Hatt 1959; Hamson &

Bates 1991 ).

Order Artiodactyla

Family: Bovidae

Wild Goat Capra aegagriis (Erxieben, 1777)

Subspecies: C. a. aegagriis (Erxieben, Mil).

Distribution: Confined to the mountain range of north-

eastern Iraq: the largest population is in the area of Barzan.

Recorded from: Bradost, Barzan near Aqra, and near Za-

witta (Layard 1852); Baradost and Sarsank (Field 1951);

Safin Dagh, Zagarta, and Jabal Sinjar (Hatt 1959); Cham-

chamal Valley (Hoogstraal & Kaiser 1958); near

Shaqlawa, Safin Dagh, Harir Dagh, Erbil Liwa, Nahiya,

near Diyana, Rawandoz, and near Darbandikhan (Harri-

son 1968; Harrison & Bates 1991); Barazan, Qara Dag

Mountain; Peramagroon Mountain (Al-Sheikhly 2012b);

Qara Dag Mountain and Peramagroon Mountain (Raza

2013). Specimens collected in Berat Dag, Mousl Liwa, Ja-

bal Zagarta, Sugrimah Dagh, and Kurkuk Liwa are resi-

dent to the collection of the FMNH (Sanborn 1940). Re-

cent observations with remarkable range extention were

made in Al-Qosh in northwestern Mousel (Ali N. Salman

pers. comm. 2014).

Mouflon Ovis orieiitalis (Gmelin, 1774)

Subspecies: O. o. gnielini (Blyth, 1841).

Distribution: Uncommon, restricted to the highlands of

eastern and north-eastern Iraq. Recorded from: Zagros

Mountain and mountains of Derbendiklian (Harrison 1968;

Nadler et al.l971; Hamson & Bates 1991 ); Jabal Himreen

near Mandli in Diyala (Al-Sheikhly 2012b). Specimens

collected in Kanispika, Baranand Dagh are resident to the

collection of the FMNH (Hatt 1959).

Arabian Oryx Oiyx leiicoiyx (Pallas, 1777)

Subspecies: O. I. leucoiyx (PaWas, Mil).

Distribution: Probably extinct; the last individual was shot

in Maqil in southern Iraq in 1914 and kept in collection

at the BMNH (Hatt 1959). Specimens known from Iraq

yet it is unclear if they have occurred on the eastern side

of the Euphrates River (Harrison 1968; Al-Sheikhly

2012c).

Saudi Gazelle Gazella saiidiva (CaiTuthers & Schwarz,

1935)

Subspecies: this species is distinct from Dorcas Gazelle

(G. dorcas) according to Groves (1988). Status reviewed

by Mallon & Kingswood (2001 ).

Distribution: Uncertain, probably extinct in the wild. There

is only one record from the Ramadi-Rutba road in west-

ern Iraqi desert (Harrison 1968; Harrison & Bates 1991;

Al-Sheikhly 2012c). A single specimen from southem Iraq

was assigned to G. siihgiittiirosa iiiarica (Mallon &

Kingswood 2001 ).

Goitred Gazelle Gazella siibgiittiirosa (Guldenstaedt,

1780)

Subspecies: G. s. inarica (Thomas, 1 897), confined to the

desert and arid steppes in western and southem Iraq; G.

s. siibgiittiirosa (Guldenstaedt, 1780) inhabits steppes of

northern and eastern Iraq.

Distribution: Widspread. Recorded from: Euphrates val-

ley (Lydekker & Blaine 1914); Samarra, Basra, Nasiriyah,

and Amara, where was reported as G. s. inarica

(Cheesman 1920; Mallon & Kingswood, 2001); Sheikh

Saad and NE Baghdad (Dollman & Burlace 1 935); Cham-

chamal (Hoogstraal & Kaiser 1958; Hamison 1968); Faid-

hat Al-Massad, Faidhat Al-Dhaba’a, Al-Dheabeat, Al- Jaz-

era Area near Haditha, Rutba, Al Qa- eem, Jabal Makhool,

Al-Sherqat, Baiji, Jabal Sinjar, Rabe’ea; Maklrmoor, Ja-

bal Himreen, Kafri, Kalar, Jalwla, Mandli, Badra,

Khanaqeen, and Ser Qella at Garmyan area (Al-Sheikhly

2012c). Specimens collected in Erbil and Kurkuk are in-

cluded in the collection of the BMNH (Hatt 1959). Spec-

imens from Basra are resident to the collection of the

BNHS, while those collected in Kurkuk, Baghdad, and Hi-

naidia are included in the collection of the FMNH (Har-

rison 1968).

RR: Small herds reported from Ganniyan area (near

Kalar), assigned to G. s. siibgiittiirosa (Su-14).

Family: Ceiwidae

Fallow Deer Daina daina (Linnaeus, 1758)

Subspecies: D. d. mesopotainica (Brooke, 1875).

Distribution: Rare, a few individuals may still survive in

the mountains of the extreme north eastern Iraq. Record-

ed from: Zakho (Ellemian & Morrison-Scott 1951); Bra-

dost Mountain (Field 1955); between Maidan and Halab-

ja (Harrison 1968; Hamison & Bates 1991).

Bonn zoological Bulletin 64 (1): 33-58

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Checklist of the Mammals of Iraq

45

European Roe Deer Capreoliis capreoliis {Linnaeus, 1758)

Subspecies: C. c. coxi (Clieesman & Hinton, 1923).

Distribution: Confined to the forested nrountains of the ex-

treme northern Iraq. Recorded from: Harir Dagh and Mer-

gasor (Hatt 1959). Subspecies C. c. coxi was described by

Cheesman & Hinton ( 1923) on the basis of individuals ob-

served in Zakho and Harir west of Rawandoz (Harrison

1968).

RR: Small herd reported from Barzan Area (M. Taib, Iraq,

pers. comm. 2014).

Family: Suidae

Wild Boar Sus scrofa (Linnaeus, 1758)

Subspecies: S. s. attiki (Thomas, 1912).

Distribution: Abundant, mainly along the Tigris and Eu-

phrates Rivers and the southem marshes. Recorded from:

Diyala, Kut al Imara, Baghdad, Ad Dawr, and Shatt al Ad-

haim (Pittman 1922); Amara, Khaniqin, and Baradost

(Sanborn 1940); Hawizeh (Drower 1949); Tigris River,

south of Amara (Page 1954); north and south of Hilla and

Mansuriyah al Shatt (Hatt 1959); near Majar, Chahala and

Masharia Rivers, and Auaisij (Thesiger 1 964); Lake Hab-

baniya, Hadithah, Badra, Sulaymaniyah, and Sarsank

(Harrison 1968); Fao and Hammar Marsh (Haba 2009);

east of Hammar Marsh (Abass 2013); Hawraman Moun-

tain (Lahony et al. 2013). Thesiger ( 1954) hunted 488 pigs

in two years around Al-Hammar Marsh. Specimens col-

lected in Baghdad and Baquba are resident to the

BMNH, while those collected in K.haniqin, Wadi Hostoc-

ki, Towaka, Zharir, Baradost, Chahala near Amara, and 30

km N Hilla are resident to the UoM (Hairison & Bates

1991).

RR: Recorded from 33 sites thrghout Iraq during 2013-

2014.

Order Lagoinorpha

Family: Leporidae

Cape Hare Lepiis capensis (Linnaeus, 1758)

Subspecies: L. c. arcihicus (Ehrenberg, 1833).

Distribution: Widespread, confined to the desert and arid

steppes west of the Euphrates River. Ellennan & Morri-

son-Scott (1951) reported the occunence of this hare in

Kuwait. Harrison & Bates ( 1991 ) reported an individual

from Saqiawiya in western Iraq; recorded also from Cen-

tral and Al-Hammar marshes (Haba 2009; Abass 2013);

Hawraman Mountain (Lahony et al. 2013).

RR: Rutba, Ga’ara, and Al-Qa-eem (An- 13); Teeb (My-

13).

European Hare Lepiis (Eulagos) europaeus (Pallas, 1778)

Subspecies:/,, e. connor/ (Robinson, 1918), in the steppe,

grassland, and foothill east of the Euphrates River; L. e.

syriacus (Hemprich & Ehrenberg, 1832), confined to the

arid steppes of the extreme north western Iraq.

Distribution: Widespread. Recorded from: Hindiya, Kut

al Imara, Shatt al Adhaim, Jalam, the frontier of Arabia,

the plain north of SamaiTa, Al Falluja, Hit, Amara, Al Ku-

mait on the Tigris, Shahraban, and Twin Canals

(Cheesman 1920); along the Tigris north of Ad Dam (Pit-

man 1922); Baghdad and from Camp Rashid on its out-

skirts (Sanborn 1940); Baghdad and Hilla (Hubbard 1955);

Hilla, Mansuriya al Shatt, and Habbaniya (Hatt 1959);

Chebaeish (Haba 2009).

RR: L. e. connori from Hor Dalmaj (Qa-13), Bahr Al-mil-

ih ( Ka- 13);/-. e. syhacns from 1 7 km NW Rutba (An- 1 3 ).

Order Rodentia

Family: Sciuridae

Caucasian Squirrel Sciuriis (Teues) cmomaliis (Gmelin,

1778)

Subspecies: 5. a. pallesceus (Gray 1867), with seasonal

variation in pelage colour.

Distribution: Abundant in the forested mountains of north-

ern Iraq. Recorded from many localities in north eastern

Iraq (Hatt 1959); Penjwin (Petter 1961); Koshek (Von

Lehman 1966); Derbendikhan Dam (Hairison & Bates

1991); Hawraman Mountain (Lahony et al. 2013).

RR: Recoded from 34 sites in northern Iraq (12-14).

Family: Hystricidae

Indian Crested Porcupine Hystrix (Hystrix) iiuiica (Kerr,

1792)

Subspecies: H. i. iiulica (Kerr 1792).

Distribution: Abundant along the Tigris and Euphrates val-

leys, deserts and arid steppes, and mountains of northern

Iraq. Recorded from many localities by Hatt ( 1959), Har-

rison (1959), Smielowski (1978), Nadachowski et al.

(1990) and Kadhim (1997); Peramagroon (Raza et al

2012); Hawraman Mountain (Lahony et al. 2013).

RR: Jabal Makhool, Al-Alam, Al-Cidir, Al-Tharthar Val-

ley (Sa-13); Rutba, Ga’ara, Wadi Horan, Khan Al-Bagh-

dadi, Al-Jezera (An- 13); .labal Himreen and near Hor

Shwija (Di-13), Hor Dalmaj (Qa-13); Central Marshes

(Th-14).

Family: Dipodidae

Euphrates Jerboa AUaclaga (Puralaciaga) eiipliratica

(Thomas, 1881)

Subspecies: /4. e. eitphrafica (Thomas, 1881).

Distribution: Relatively local in central and southern Iraq.

Records were provided by Hanison ( 1959), Hatt ( 1959),

Al-Robaac (1976), Kadhim & Wahid (1986) and Nada-

chowski ct al. (1990).

RR: Western Hammar Marsh (Th-13).

Lesser Egyptian Jerboa Jacnhisjaciiliis (Linnaeus, 1758)

Subspecies: J. j. loftusi ( Blanford, 1 875 ), in the arid steppe

of the eastern Euphrates basin; ,/. /. vocater (Thomas,

Bonn zoological Bulletin 64 (1 ): 33-58

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46

Omar F. Al-Sheikhly et al.

1921), in western and southern desert of Iraqi.

Distribution: Widespread, in the desert and rocky steppes

of central and southern Iraq. Recorded from: Az Zubeir

(Ellenuan & Morrison-Scott 1951 ); W Basra (Hatt 1959);

40 km E Rutba, Elatra (Nadachowski et al. 1990); Elaur

al Hasa, S Habbaniya (Harrison & Bates 1991 ); Al-Cidir

(Al-Sheikhly 2013).

RR: Wadi Al-Tharthar (An- 1 3).

Family: Gliridae

Large-eared Garden Dormouse Eliomys melanunis (Wag-

ner 1840)

Subspecies: E. in. melanunis (Wagner, 1840).

Distribution: Restricted to the rocky steppe of north west-

ern Iraq. Recorded from: Hatra (Nadachowski et al. 1978);

from c. 90 km SW of Mousel (Harrison & Bates 1991).

Forest Dormouse Diyomys nitediila (Pallas, 1779)

Subspecies: Possibly close to the Iranian subspecies D. n.

(Blanford. 1875).

Distribution: Confined to the Oak forests of Kurdistan.

Recorded from: Sulaymaniyah in northern Iraq (Jawdat

1977; Harrison & Bates 1991 ).

Family: Spalacidae

Middle East Blind Mole Rat Nannospalax ehrenbergi

(Nehring, 1898)

Iraqi populations of this variable taxon belong to the same

karyotype (Co§kun et al. 2012). Although N. ehrenbergi

is better regarded as a superspecies comprising several bi-

ological species (Hadid et al. 2012), uncertainty remains

about the taxonomic rank as well as the nomenclature of

the Iraqi karyotype, which is widely distributed also in

southeastern Anatolia.

Distribution: Found in the steppe of northern Iraq. First

description provided by Bate ( 1930), who reported bones

of N. ehrenbergi from the Hazar Mard caves near Sulay-

maniyah. Also recorded by Reed ( 1 958) and Petter (1961)

(Hairison & Bates 1991). Individuals collected in Ser

Amadiya, Tim near Bermaneh (Harrison 1956) and

Sarsank (Hatt 1959) were also assigned to this species. Re-

cently recorded as Spalax leucodon from Hawraman Mts.

(Fahony et al. 2013). The distribution in Iraq is sum-

marised by Co§kun et al. (2012).

Family: Muridae

Eastern Broad-toothed Field Mouse Apodennis mystaci-

niis (Danford & Alston 1877)

Subspecies: A. m. mystaciniis (Danford & Alston, 1877).

Distribution: Confined to the Oak forests and wooded

roclcy cliffs of northern Iraq. Recorded from: Sarsank (Hatt

1959); Qali Ali Beg (Nadachowski et al. 1990; Harrison

& Bates 1991 ).

Yellow-necked Field Mouse Apodennis flavicollis (Mel-

chior, 1834)

Subspecies; A. f. argyropnloi (Heptner, 1948).

Distribution: Confined to the Oak forests of northern Iraq.

Recorded from: Haj Onuan and Sarsank (Hatt 1959); Sala-

hadin and Sulaymaniyah (Al-Shenawi et al. 1982; Harri-

son & Bates 1991 ).

Steppe Field }Aoy\%Q Apodennis witherbyi (Thomas, 1902)

Subspecies: May be the species is represented by the sam-

ple collected in Qazvin (northern Iraq), which separated

electrophoretically from both A. sylvaticiis and A. flavi-

collis. To date, however, the specimens from Qazvin are

considered closer to the latter (Darviche et al. 1979).

Distribution; Confined to the rocky mountain slopes of the

extreme northern Iraq. Harrison (1956) reported this

species from Ser Amadiya as A. sylvaticiis. Hatt (1959)

confirmed its occurrence at the same locality. Also record-

ed by Petter ( 1961 ) (Harrison & Bates 1991 ).

Roof Rat Rattiis rattiis (Finnaeus, 1758)

Subspecies: R. r. rattiis (Finnaeus, 1758).

Distribution: Widespread, an urban pests mainly in settle-

ments along the Tigris and Euphrates Rivers and their trib-

utaries. Recorded from many localities in central and

southern Iraq (Cheesman 1920; Hatt 1959; Nader 1968b;

Al-Shenawi et al. 1982).

RR: Old Rasafa; Yarmook (B-13; 14).

Brown Rat Rattiis norvegiciis (Berkonhout, 1769)

Subspecies: R. n. noixegiciis (Berkonhout, 1769).

Distribution: Widespread, mainly along the Tigris and Eu-

phrates Rivers and their tributaries. Recorded from; Bas-

ra (Cheesman 1920; Hatt 1959); Baghdad (Kadhim et al.

1979; Al-Shenawi et al. 1982; Harrison & Bates 1991);

Hammar Marsh (Abass 2013).

RR: 13 localities in Baghdad (B-13); Mashab River and

Garmatt Ali (Ba-14).

House Mouse Mas (Mas) niiisciiliis (Finnaeus, 1758)

Subspecies: M. m. praetextiis (Brants, 1827) and M. in.

bactrianus (Blyth, 1846) occur in both eastern and west-

ern Iraq (Harrison 1972). However, further investigation

is required.

Distribution: Widespread. Recorded from: Sinn Abtar and

Twin Canals from Kut al Imara, Basra, Shaikh Saad, and

Amara (Cheesman 1920); Habbaniyah, Baghdad, Sarsank,

K-3, Hilla, Haj Omran, the region of Salahuddin, and Jar-

mo (Hatt 1959); Chebaeish and Ashar (Haba 2009); also

recorded by Nader (1968b), Al-Shenawi et al. (1982);

Hammar Marsh (Abass 2013); Hawraman Mountain (Fa-

hony et al. 2013).

RR: Al-Alam (Sa-13), Ramdi (An-13), and Yarmook area

(B-14).

Bonn zoological Bulletin 64 ( 1 ): 33-58

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Checklist of the Mammals of Iraq

47

Short-tailed Nesokia Nesokia indica (Gray, 1830)

Subspecies; TV. i. hoxtoni (Thomas, 1919).

Distribution: Widespread in the riverine plains of both cen-

tral and southern Iraq. Recorded from: Amara, A1 Qur-

na, Aqar Quf, Shaikh Saad, and An Nasiriya (Cheesman

1920); Baghdad (Sanborn 1940); Cheesman’s sites,

Maqil and Hinaidi (Ellerman 1948); Ur (Pocockl934);

Baghdad, Mansuriya al Shatt, Ramadi, and Babylon (Hatt

1959); recorded also by Fetter ( 1961 ), Nader ( 1968b), Al-

Robaae ( 1977), Kadhiin (1981 ) Al-Shcnawi et al. ( 1982);

and Abass (2013).

Long-tailed Nesokia Nesokia bunnii (Khajuria, 1981 )

Originally described under the genus Eiythrouesokia by

Khajuria ( 1981 ), later on it was proved to be a distinctive

species of Nesokia (Al-Robaae & Felten 1990).

Distribution: Current status is unknown. Recorded only

in the marshes at the confluence of the Tigris and Eu-

phrates Rivers in south eastern Iraq (Khajuria 1981; Al-

Robaae & Felten 1990), the species could also possibly

occur in Al-Hawizeh Marsh straddling the Iraq-Iran south

eastern border.

Family: Cricetidae

Gray Dwarf Hamster Cricetulus migratoriiis ( Pallas, 1 773 )

Subspecies: C. /;;. cinerascens (Wagner, 1848).

Distribution; Abundant in cultivated lands, forests, and

steppes mainly of northern Iraq. Individuals were collect-

ed in Ser Amadiya by Harrison ( 1 956). A specimen includ-

ed in the collection of the FMNH was trapped at Jarmo

(Hatt 1959). Also recorded by Fetter ( 1961 ) and Kadhim

(1975) (Han'ison & Bates 1991).

Baluchistan Gerbil Gerbilliis (Heudecapleura) nainis

(Blanford, 1875)

Subspecies; G. ii. nanus (Blanford, 1875), in eastern Iraq;

G. n. arabiinn (Thomas, 1918), confined to the desert of

south western Iraq.

Distribution; Widespread, in deserts and arid steppes of

central and southern Iraq. Recorded from: Ali Al-Gharbi

by Fetter et al. ( 1957); specimens from Basra Liwa are in-

cluded in the collection of the UMC ( Hanison & Bates 1991 ).

Wagner’s Gerbil GerbiUus (Hendecapleiira) dasyiiriis

(Wagner, 1842)

Subspecies: G. d. dasyunts (Wagner, 1842).

Distribution: Widespread in the steppic deserts of central

and western Iraq. Recorded from: c. 1 1 km SE H2 pump-

ing station in western Iraq (Hanison 1956c); Haditha (Hatt

1959; Harisson & Bates 1991 ).

RR: Wadi Al-Tharthar (An- 13).

Mesopotamian Gerbil GerbiUus (Hendecapleiira)

mesopotamiae (Hanison, 1956)

Subspecies: G. ni. niesopotaniiae (Harrison, 1956), with

Bonn zoological Bulletin 64 ( 1 ): 33-58

some individual and local variation.

Distribution: Widespread in arid terrains of central and

southern Iraq. Harrison ( 1956) reported the species in the

suiTOundings of Amiriya. Recorded from: Lake Hab-

baniya, Baghdad, K-3, Basra; Haur Al Hasa, Al-Jadriyah,

Ain Tina, 4 km S Falluja, 10 km W Baghdad, Rashidiya,

4 km S Abu Ghraib, and Um al Khanazeer Island (Hatt

1959); 5 km SE Najaf (Nasher 1970); Ramadi Liwa

(Nadachowski et al. 1990). Specimens from Ali Gharbi

are included in the collection of the PM, while those from

Baghdad and Baquba are included in the collection of the

BRCB (Harrison & Bates 1991 ).

Cheesman’s Gerbil GerbiUus (GerbiUus) cheesniani

(Thomas, 1919)

Subspecies: G. c. cheesniani (Thomas, 1919).

Distribution: Confined to open sandy hummocks with

sparse vegetation of central and southern Iraq. Recorded

from: Along the west bank of the Euphrates River (SW

Al Falluja, near Haur al Hasa, Amiriya: Harrison 1956);

Lake Habbaniya (Hatt 1959; Harrison & Bates 1991).

Indian Gerbil Tatera indica (Hardwicke, 1807)

Subspecies: T. i. taeniura (Wagner, 1843).

Distribution: Widespread. Recorded from: Amara, Sinn

Abtar, Kut al Imara, Baghdad, Aqarkuf, Sheikh Saad,

Shahraban, and Baiji (Cheesman 1920); Kasimain and

Baghdad (Ellerman 1948); near Habbniya (Harrison

1956c); Ali Gharbi (Fetter 1957); Tell Imlihiye (Kock &

Nader 1983); Hatra (Nadachowski et al. 1978; Harrison

& Bates 1991 ). Further records were provided by Kadhim

(1998).

Persian Jird Meriones (Paranieriones) persiciis (Blanford,

1875)

Subspecies; M. p. persiciis (Blanford, 1875).

Distribution: Restricted to the mountains of northern Iraq.

Recorded from: Kani Mase and Ser Amadia (Hanison

1956c); Penjwin and Jabal Batiwa (Fetter et al. 1957);

Salahadin, Shaqlawa, and Jarmo (Hatt 1959).

Tristram’s T\xd Meriones (Pallasioinvs) Iristranii (Thomas,

1892)

Subspecies: M. t. lycaon (Thomas, 1919), with ochraceous

buff dorsal pelage.

Distribution: Confined to the Oak forests of northern Iraq.

Recorded from: Dohuk (Ellerman 1948); Bakrajo in Su-

laymaniyah and Sarsank (Hatt 1959). Fetter et al. ( 1957),

Kock & Nader (1983) and Nadachowski et al. ( 1990) pro-

vided further records. A few specimens are included in the

collection of the HZM (Harrison & Bates 1991 ).

Libyan Jird Meriones (Pallasioinvs) libvciis (Lichtenstein,

1823)

Subspecies: M. 1. syrius (Thomas, 1919), confined to the

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48

Omar F. Al-Sheikhly et al.

desert steppe of western Iraq; M /. eiythwunis (Gray,

1 842), occurs in the eastern region of the Euphrates basin.

Distribution: Widespread. Recorded from: Az Zubair W

Basra (Ellemian 1948); Amiriya and Al Jazira SW of

Samarra (Harrison 1956c); At Tuba, Basra (Hatt 1959).

Also reported by Heptner ( 1 940) and Fetter et al. ( 1 957);

Al-Cidir (Al-Sheikhly 2013).

Sundevall’s Jird Meriones (Pallasiomys) crassiis (Sunde-

vall, 1842)

Subspecies: M. c. crassiis (Sundevall, 1842), confined to

the arid steppes of western Iraq; M. c. charoii (Thomas,

1919) occurs in the eastern region of the Euphrates basin.

Distribution: Widespread. Recorded from: Kasimain and

Az Zubeir (Cheesman 1920); Haur al Hasa and Jazira

(Harrison 1956c); Ali Gharbi and Kurkuk (Fetter et al.

1 957); 1 50 km NW Basra, Haditha, 24 km W Ramadi, 32

km W Baghdad, and Balad (Hatt 1959); Balad (Hanison

1972; Harrison & Bates 1991 ).

Transcaueasian Mole Vole EUohiiis (Afgaiioiiiys) hitesceiis

(Thomas, 1897)

Subspeeies: Fossibly £. /. hitesceiis (Thomas, 1897).

Distribution: Frobably still present on the mountains of

northern Iraq. Fossils were discovered in the caves of Haz-

ar Mard ( Bate 1 930) and Shanidar ( Hatt 1 959). Hatt ( 1 959)

reported that the species might still inhabit the mountains

of northern Iraq. Baltazard & Seydian ( 1960) reported the

occun ence of some speeimens from Fenjwin, whereas Fet-

ter (1961), who examined the same collection, did not

(Harrison & Bates 1991).

Eurasian Water Vole Arvicola ainphibius (Linnaeus,

1758)

Subspeeies: A. a. persicits (de Filippi, 1865).

Distribution: Restricted to the mountains of northeastern

Iraq; reeorded from Fenjwin ( Fetter 1961; Harrison &

Bates 1991).

RR: Chebaeish (Central Marshes, southern Iraq) yet with-

out supporting material (N. Abod. pers. comm. 2014).

Social Vole Microtiis (Microtiis) socialis (Falas, 1773)

Subspecies; Hairison and Bates (1991 ) included giieiitheri

and iraiii under M. socialis. However, M. iraiii has been

separated as a distinct taxon (Kefelioglu & Krystufek

1999; Golenishchev et al. 2002).

Distribution: Confined to the mountains and foothills of

northern Iraq. Recorded from: Ser Amadiya (Harrison

1956a); Sulaymaniyah and Sarsank (Hatt 1959); Fenjwin

(Fetter 1961 ); Hawraman Mountain (Lahony et al. 2013).

A few specimens are included in the collection of both

BMNH and HZM (Hamson & Bates 1991). Ellennan

(1948) reported the occurnce of some specimens from

“Baghdad”, but it is wisely questioned whether they re-

ally came from that locality (Hatt 1959).

Order Cetacea

Family: Delphinidae

Indo-Facific Humpbacked Dolphin Sousa cbineiisis (Os-

beck, 1765)

Distribution: Marine coasts of southern Iraq. The species

(under the name Sotalia leiitigiiiosa) was recorded from

Khor Al Zubeir (Al-Robaae 1970a).

Indo-Facific Bottlenose Dolphin Titrsiops aditncus (Ehren-

berg 1833)

Distribution: Marine coast of southern Iraq. Reported from

Rass Al-Mataf (near Fao) in the northern part of the Ara-

bian Gulf (Al-Robaae 1974).

Finless Foipoise Neopliocaena phocaeiioides (G. Cuvier,

1829)

Distribution: Marine coast of southern Iraq. Recorded

from the Iraqi tciTitorial water c. 37 km S Fao; another

specimen was reported from Fao (Al-Robaae 1975).

Family: Balaenopteridae

Blue Whale Balaenoptera iiuisciihis (Linnaeus 1758)

Distribution: Marine coast of southern Iraq. Uncertain sta-

tus; listed by Mahdi & George ( 1969) but without provid-

ing further details.

Biyde’s Whale Balaenoptera edeni (Anderson 1879)

Distribution: Marine coast of southern Iraq. Recorded

from: Khor Al Zubeir (6.4 km NW of port Umm Qasr)

(Mahdi 1967; Al-Robaae 1969).

Humpback Whale Megaptera iiovaeangliae (Borowski

1781) (Arabian Sea sub-population)

Distribution: Marine coast of southern Iraq. Specimens

collected near Fao and Shatt Al-Arab are resident to the

collection of the BRCB (Hatt 1959; Al-Robaae 1970b).

DISCUSSION

Historical framework

A checklist of the mammals of Iraq was urgently required

in order to support conseiwation plans for the Iraqi fauna.

We reviewed eight major orders including 28 families, 65

genera and 93 species. This manuscript represents the first

systematic checklist of wild mammals prepared for Iraq

since the 1950s, and includes many new species record-

ed since Hatt’s masterpiece. Moreover, Hatt (1959) includ-

ed 12 species that were re-evaluated and eventually as-

signed to other taxa by Harrison & Bates (1991): Bicol-

ored White-toothed Shrew (Crocidiira leiicodoii persica)

assigned to Lesser White-toothed Shrew (C. siiaveoleiis),

the European Hedgehog (Erinaceiis eiiropaeus) assigned

to East European Hedgehog (E. concolor), Babylonian

Bonn zoological Bulletin 64 ( 1 ): 33-58

©ZFMK

Checklist of the Mammals of Iraq

49

Sheath-tailed Bat ( Taphozous kachhensis babylouiciis) as-

signed to Naked-bellied Tomb Bat {T. midiveutris mag-

niis). Northern Bat (Eptesiciis nilssoni nilssoni) assigned

to Botta’s Serotine {E. bottae); V: Hingston’s Serotine

{Eptesiciis sodalis liiiigstoiii) assigned to Botta’s Serotine

(E. bottae), Cox’s Pipistrelle (PipistreUiis coxi) assigned

to Riippell’s Pipistrelle (P. riippellii), the Indian Red Fox

( Vidpes V. pusillits) assigned to the Arabian Fox ( V. v, ara-

bica), the European Hare (Lepiis eiiropaeiis coiinori) as-

signed to Cape Hare (L. capeiisis comiori), the Persian

Squin'el (Sciurits anoiiialiis) assigned to the subspecies

S. a. pallesceiis, Wagner’s Gerbil [Gerbillits {Hende-

capleiira) dasyiiriis] assigned to Han ison’s Gerbil [G. {H. )

luesopotamiae], Turkish Jird (Merioiies blackleri black-

leri) assigned to Tristram’s Jird [M. (M.) tristrami), and

the Persian Vole (Microtiis iraiii) assigned to the Social

Vole (M. sodalis).

Later on, further observations were provided by Harri-

son (1964; 1968; 1972) and Harrison & Bates (1991).

Nevertheless, eight new records obtained in 2012 and 2013

were added to the present checklist: Syrian Jackal {Can-

is aureus syriacus) from Hussaiba, 7 km N W Al-Qa-eem,

western laq (Al-Sheikhly 20 1 2a), Arabian Wolf (Cm; A lu-

pus arabs) from Rutba, western Iraq (Al-Sheikhly 2012b),

Indian Grey Mongoose (Herpestes edwadsi) from Kurdis-

tan, northern Iraq (Al-Sheikhly & Mallon 2013), Least

Weasel (Miistela nivalis) from Kurdistan, northern Iraq

(Raza 2013), Sand Cat (Felis niargarita) from desert of

Al-Najaf Province (Mohammad et al. 2013). Syrian Cape

Hare (Lepiis capeiisis syriacus) from Rutba, western Iraq

(Al-Sheikhly. in litt.), Mediterranean Horseshoe Bat {Rlii-

iiolophus euiyale) from Kurdistan, northern Iraq (Al-

Sheikhly et al. 2015), and Geoffroy’s Myotis (Myotis

emargiiiatiis) from Kurdistan, northern Iraq (Al-Sheikhly

et al. 2015).

Present checklist

Among the 93 species recorded in Iraq, 24 were evaluat-

ed by the lUCN (one species listed as extinct, one species

as Critically Endangered, four species as Endangered,

eight species as Vulnerable, seven species as Near

Threatened, and three species as Data Deficient) (Table

2). In this context, the above-mentioned Smooth-coated

Otter, a flagship species of Iraq is worth of mention. Dur-

ing the 1980s and 1990s this species was guessed to be

extinct due to habitat loss and fragmentation as well as to

over-hunting, but it has been rediscovered very recently

(Omer at al. 2012; Al-Sheikhly & Nader 2013; Al-

Sheikhly et al. 2014).

The strategic geographical position of Iraq provided the

local mammals with a historical natural connection to the

fauna of southern Turkey, western Iran, eastern Syria and

the north eastern part of the Arabian Peninsula. Hence, it

did not come as a surprise that six Arabian species were

found west of the Euphrates River: the Arabian subspecies

of the Lesser Mouse-tailed Bat (Rliiiiopoina hardwickii

arabium), the Arabian Wolf (Cr//rA lupus arabs), the Ara-

bian Fox ( Vidpes viilpes arabica), the Arabian Cape Hare

(Lepiis capeiisis arabicus), the Arabian Sand Gazelle

(Gazella subguttiirosa iiiarica), and the Arabian sub-

species of Baluchistan’s Gerbil (GerbiUiis iiaiiits arabi-

uiii). In addition, the Sand Cat (Felis niargarita liarrisoni),

a species confined to the Arabian Desert, was recently

recorded in Iraq as well. Furthennore, the Syrian Desert,

the monotonic landscape shared between easter n Syria and

wester n Iraq, which extends until the western bank of the

Euphrates River, is inhabited by five species: the Syrian

Jackal (Cam's aureus syriacus), the Syrian Stone Marten

(Martes foiiia syriaca), the Syrian Marbled Polecat

( Vorniela peregiisna syriaca), the Syrian Striped Hyaena

(Hyaena hyaena syriaca) and the Syrian Cape Hare (Le-

pus capeiisis syriacus). Finally, the mammalian fauna of

Iraq is also naturally connected to the zoogeographic realm

of Persia, a teiritoi'y stretching mainly east of the Tigris

River and along the border with Ii'an. Eight species can

be found so far in the forested mountains, rocky valleys,

and grassy steppes of this I'egion: the Iranian Wolf (Can-

is lupus pallipes), the Persian Leopard (Pantliera pardus

saxicolor), the Iranian (Asiatic) Cheetah (Acinonyx Juba-

tus venaticus), the Persian Goitred Gazelle (Gazella

subguttiirosa subguttiirosa), the Wild Goat (Capra aega-

griis), the Persian SquiiTel (Sciurus anoinaliis), the Per-

sian Jird [Merioiies (Paranieriones) persiciis\, and the Pei‘-

sian Water Vole (Anicola ainpliibius persicus).

During the Iraq war in 2003, vandalism against the Bi-

ological Research Centre of Baghdad (BRCB) occuned

and, as a consequence, many valuable specimens of the

Iraqi fauna were lost or damaged. When we examined

mammal specimens kept in the BRCB collection we al-

so noticed that many of them were decomposed due to

long-tei'in storage. Unfortunately, substitute material was

not available especially for Soricomor'pha, Chiroptera and

Rodentia. Nevertheless, we would like herein to focus on

a few valuable mammal specimens collected in Iraq and

described as new taxa. The Haditha Mouse-tailed Bat

(Rliiiiopoina liaditliaensis) was first described by Khajuria

( 1988) based on a few specimens discovered in Haditha

(westerm h'aq) and now resident to the collection of the

BRCB. According to their morphology and, especially, to

the cr'anial features provided by Khajur ia ( 1988), Kock et

al. (2001 ) regarded R. liaditliaensis as junior synonym of

the Greater Mouse-tailed Bat (R. inicropliylliiin). Howev-

er, the status of R. liaditliaensis remains uncertain and fur-

ther investigation is needed. Other valuable specimens

(study skin kept at the BMNH and live cub) belong to the

Iraq Smooth-coated Otter (Lutrogale perspicillata

inaxwelli), collected by Gavin Y. Maxwell (Hayman

1959). The otter skin was found in the village of

Bonn zoological Bulletin 64 ( 1 ): 33-58

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50

Omar F. Al-Sheikhly et al.

Abusakhair (Faraijat tribe, c. 56 km SE of Amara, along

the Tigris River) and included in the collection of the

BMNH. The live cub was probably found in a tumulus is-

land village called Daub, c. 19 km NW of Al-Azair (west

of the Tigris River: Hayman 1956; Maxwell 1957, 1960;

Al-Sheikhly & Nader 2013; Al-Sheikhly et al. 2014). Af-

ter 1980s L. p. maxwelli was believed to be extinct, but

during field surveys cairied out in 2007-2012 the occur-

rence of L. p. maxwelli in southern Iraqi marshes was

proved by Al-Sheikhly & Nader (2013). Additionally, an-

other isolated population of L. perspicillata was found in

Kurdistan (northern Iraq), this finding representing a re-

markable extension of the species range known so far

(Omeret al. 2012; Al-Sheikhly & Nader 2013). The Long-

tailed Nesokia (Nesokia humiii) was described by Kha-

Juria (1981) within the genus Eiythronesokia on the ba-

sis of two individuals collected in Al-Quma (southern Iraq)

and presently included in the collection of the BRCB. This

species can be distinguished from the similar Short-tailed

Nesokia by its large size, cranial features, long tail, red-

ness of the dorsal pelage, and incipient development of

the lower incisor root. Al-Robaae & Felten (1990)

showed that Khajuria’s (1981) N. humiii, indeed, repre-

sents a distinct species. Han ison & Bates (1991) compre-

hensively examined Khajuria’s record and concluded that

it could be a large specimen of Short-tailed Bandicoot Rat

(Nesokia iudica), with an abnormal long tail possibly re-

flecting its ecological adaptation to the aquatic habitat.

However, more samples from the Basra Province are re-

quired to confirm the taxonomic identity of this taxon in

Iraq.

We found that hunting and trapping represent the main

threats to the wild mammals of Iraq. Indian Crested Por-

cupine, Cape Hare, and Jerboas are widely hunted for meat

consumption and fur by local people, especially in cen-

tral and western Iraq. On the contrai'y, such practices are

forbidden in southern Iraq. The spines of the porcupine

are used to decorate houses and cars, the tails of both hares

and jerboas are used to make keychains, while the furiused

to create handmade souvenirs. Goitred Gazelle, Wild Goat,

and Mouflon are heavily hunted to be consumed as food

or trapped to be raised as domesticated pets (Al-Sheikhly

2012b). The skulls of hunted gazelles and goats, especial-

ly those with big horns, are usually mounted and/or hanged

on the front wall of the hunters’ houses to testify the suc-

cess of their hunting trips. Local farmers hunt wild boars

for puipose of eradication, as they considered this species

as a pest. Locals also hunt Fox, Jackal, Wolf, Striped Hye-

na, Honey Badger, Jungle Cat and Leopard according to

traditional practices and/or myths (Hatt 1959; Handson

1972). The Bedouins are known to chase wolves and hye-

nas in the western desert of Iraq in order to collect their

eyes, claws and teeth, which are all used as symbols of

gloiy according to the Arabian tradition. Alternatively,

Bedouins usually trap wolf cubs in order to raise them as

pets. More in general, a conflict between the previous car-

nivore species and shepherds, farmers and hunters is on-

going, the local people being constantly engaged to elim-

inate these species whenever possible. Especially wolves

and leopards are considered dangerous to humans and de-

structive to cattle herds (Fig. 2). Nevertheless, the Euro-

pean Otter and the Smooth-coated Otter are widely hunt-

ed for their fur throughout Iraq. Such a practice led the

populations of both species to the edge of the extinction.

Otters’ fur is used as a waterproof sac, which can be filled

with illegal imports and inserted in the gasoline tank of

smugglers’ vehicles (Al-Sheikhly & Nader 2013; Al-

Sheikhly et al. 2014). Iraqi farmers and hunters regular-

ly collect juvenile Persian SquiiTel and Small Indian Mon-

goose in spring. Squirrels are mainly trapped in northern

Iraq before being canied out to the animal markets of

Baghdad, Mousel and Kurkuk. Many of them usually die

due to serious injuries caused by trapping and/or trans-

portation before they can be sold as tamed pets. Hunters

of central and southern Iraq monitor the lactating female

mongooses and chase them at their dens in order to col-

lect young individuals. These latter are then usually sold

to local farmers. Young mongooses, indeed, are usually

trained to control rodents and insects in farms and settle-

ments.

Unfortunately, illegal trapping and trafficking of au-

tochthonous mammalian species is widely performed in

Iraq. The weakness of the wildlife hunting legislation al-

lows unauthorised local hunters to trap many wild animals

for pui-pose of export to neighbouring countries. For in-

stance, young Sand Gazelle and Wild Goat are trapped

during the breeding season in the western desert and north-

eastern mountains of Iraq, and then shipped to neighbour-

ing countries (Al-Sheikhly 2012c). Youngs of the Brown

Bear are trapped in many localities of northern Iraq ( Kur-

distan) and sold as pets in the animal markets of Bagh-

dad. Nevertheless, the trade of exotic species to Iraq does

frequently occur as well, this representing an additional

threat to the local biota. For instance, the African Lion

(Panthera leo), the North-West African Cheetah (Acinonyx

jubatus hecki), and the Bengal Tiger (Panthera t. tigris)

are regularly imported and sold in the animal markets of

Iraq. The price for each specimen ranges between US$

1 ,700 and US$ 4,000, cubs being the most requested items.

It is clear that the conflict between man and wild species

is leading to a rapid decline of many animal populations

in Iraq. The lack of scientific knowledge and awareness

about the wild fauna among local fanners and hunters, the

ever-increasing hunting pressure, poaching and related il-

legal trade of animals as well as ongoing habitat loss and

fragmentation represent the main threats impacting wild

mammals of Iraq. Despite the huge efforts proposed by

the Iraqi authorities in order to protect the wild mammalian

fauna, further enforcement of present hunting legislation

is essential, as Iraq became a signatoiy countiy to the Con-

Bonn zoological Bulletin 64 ( 1 ): 33-58

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Cliccklist of the Mammals of Iraq

51

vention on International Trade in Endangered Species of

Wild Fauna and Flora (CITES).

Acknowledgments. We are grateful to the National Geograph-

ic Society Conservation Trust (WA, USA) for granting a con-

servation project (#C261-13 to F.B.) on the Smooth-coated Ot-

ter. Fieldwork carried out in the Mesopotamian marshes, indeed,

allowed us to collect some data on wild mammals that were in-

cluded in this checklist. We would like to thank Ali N. Salman

(Iraqi Ministry of Environemnt), Abd Al-Bari Al-Sa’don and

Flameed Al-Habash ( Iraqi Flunters Association ) for providing de-

tails and photographs on wild mammals of Iraq; Najah A. Huss-

ian and Adel F. Abass (University of Basra), Khalid Faiq (Uni-

versity of Kalar, Kurdistan-lraq) and Sarbaz I. Mohamad (Uni-

versiy of Salah Adin, Kurdistan-lraq) for their helpful comments

and assistance during the 2014 fieldwork. We extend our thanks

to the lUCN/SSC Otter Specialist Group (OSG) members:

Nicole Duplaix, Syed A. Hussain, Waseem A. Khan, Arno Gut-

leb, Lesley Wright, Grace Yoxon and Padma de Silva; Paul .1.

Bates and Malcolm Pearch (Han'ison Institute-UK); Tamas Gdr-

tol (Hungarian Natural Histoi^ Museum, Hungary); David Mal-

lon (lUCN/SSC Antelop Specialist Group, UK); lyad Nader

(Saudi Arabia); Masaa M. Al-.lumaily and Abdulkareem Nash-

er (Yemen); Hossein Zohoori (lUCN/SSC Bats Specialists

Group, Islamic Repuplic of Iran). Also, we are grateful to Derek

Gow (Derek Gow consultancy Ltd., UK) for his comments on

the Arvicola amphihiiis identification in southern Iraqi marsh-

es. We are deeply grateful to Rainer Hutterer for his precious

editorial work on a draft version of this checklist.

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Checklist of the Mammals of Iraq

55

GEOGRAPHICAL GAZETTEER

Al- Anbar Province (An)

A1 Faluja (Falluja; Feluja): 33°2 l’N43°46’E; Abu Al Jir:

33°16’N42°53'E;AbuGhreb(Abu Graib): 33°28'N44°17'E; Al-

Dheabeat ( Al-Theabeatt): 32°49’N43°1 1’E; Al-Ga’ra:

33°29’N40°26’E; Al-Jezera (Jezaratt Al-Anbar):

34°11'N42°49’E; AI-Qa-eem: 34°22’N4 1 °06’E; Anah

(Ana):34°28’N41°56’E; Cheleabat: 32°18’N39°47’E; Faidhat Al

Massad: 32°55’N40°13’E; Faidhat Al-Dhaba’a: 32°58’N40°

20'E; HI station: 33°40’N39°57’E; H2 station: 33°16'N40°40'E;

Habbaniya: 33°22’N43°35’E; Hadithah: 34°07'N42°23’E; Haur

al Hasa: 33°12'N43°45'E; Hit: 33°38’N42°49’E; Hussai-

ba:34°23’N40°59’E; K3 station: 32°55’N39°45’E; Lake Hab-

baniya: 33°I7’N43°29'E; Nikheab:32°02’N42°15'E; Qasr al

Helqum: 33°48’N40°35’E; Rahaliyah: 32°46’N43°23'E; Rama-

di (Ar Ramadi): 33° 27’ N 43°I9’ E; Rawa: 34°28’N41°55' E;

Razaza Lake: 32°43’N43°3 1’E; Rutba (ArRutba):

33°03’N40°I8’E; Saqlawiya: 33°24’N 43°41’E; Shamiyah

Desert: 30°20’N46°40’E; Sheikh Hadid: 34°l I ’N42°22’E; Sin

Al Dhibban (Thuban; Zeban): 33°23’N43°36 ‘E: Tlstation:

34°15’N41°20’E; Wadi Arar: 31°27’N4I°38'E; Wadi Horan:

33°54’N41°57’E; Wadi Haqlaniya (Haqian): 34°05'N42°27’E.

Babel (Babylon) Province (Bb)

Al-Mahmodiyah: 32°29N’44°25’E; Hilla: 32°32’N44°13’E;

Hindiya (Sadatt Al-Hindiya): 33°04’N44° 22’E,

Baghdad Province (B)

Agersjoof (AqarQuf; Aqarkuf; Akkarkuf): 33°24’N44°21’E; Al-

Adhamiyah: 33°22’N44°21’E; Al-Kadmiyah: 33°22'N44°20’E;

Al-Rashdiyia (ArRashdiyia): 33°32'N44°2 1'E; Amiriya:

33°22’N44°32’E; Dor’ah (Dora): 33°1 5'N44°23'E; Jadrriyah

and Umm Al-Khanazeer Island: 33°17’N44°23'E; Mnstan-

siriyah: 33°22’N44°24’E; Old Rasafa: 33°20’N44°24’E;

Rashidiya: 33°25'N44°22’E; Rustamiyah: 33°16'N44°32'E;

Yannook: 33°1 7’N44°20’E; Zaafaraniyah: 33°13’N44°33’E.

Basra Province (Ba)

Abu Al-Khasib: 30°26’N48°00’E; Al-Masshab:

30°39’N47°28'E; Al-Qurna (Gurna):3 l°0rN47°27'E; Ashar:

30°30’N47°50'E; At Tanumah: 30°3 l'N47°5 1’E; At Tuba:

30°27’N47°30’E; Basra city: 30°30’N47°48’E; Fao:

29°55'N48°30’E; Jabal Sanam: 30°06'N47°42’E; Khor Al-

Zubeir: 30°02’N47°58’E; Kut as Sayyid: 30°3 1 ’N47°49'E; Ma-

jnoon: 31°12'N47°30’E; Maqil (Ma’qal): 30°33’N47°46’E; Rass

Al-Beisha: 29°55’N48°33’E; Shaiaba (Shaiba; Shuaiba):

30°24’N47°41’E; Shatt Al-Arab: 30°26’N48°09'E; Umm Al

Rassas: 30°26’N48°07’E; Zubeir (Az-Zubeir): 30°24’N47°45’E.

Diyala Province (Di)

Adhaim (Shatt al Adhaim): 34°01 ’N44°19’E; Al Miqdadiyah:

33°58’N44°58’E; Al-Khalis: 33°5 l’N44°32'E; Baladruz:

33°4I’N45°05'E; Baquba (Baqubah): 33°45’N44°40’E; Cte-

siphon: 33°06'N44°36’E; Jalwla: 34°1 7’N45°10’E; Kafri:

34°4rN44°58'E; Khaniqin: 34°20’N45°22’E; Lake Himreen:

34°10’N44°59’E; Madain: 33°05'N44°35’E; Mandli:

33°44’N45°34’E; Mansuriya al Shatt: 33°45’N44°24’E;

Nahrwan (Nahrawan): 33°22’N44°42’E; Shahraban:

33°59’N44°56’E.

Duhok Province (Du)

Banaman: 36°23’N44°08’E; Bebaidi: 37°07’N43°27'; KaniMase

(KaniMasi): 37°13’N43°25’E; Sarsank (Sersang):

37°03’N43°1 5’E; Ser (Ser’) Amadia (Amadiya):

37°08’N43°27’E; Zakho: 37°08’N42°41’E; Zaweta (Zawitta):

36°55’N43°12’E.

Erbil Province (Er)

Bradost Mountain (JabalBradost): 36°46‘N44°21'E; Barzan: 36°

55’N44°03’E; Bola: 36°45’N44°42'E; Diyana:

36°39’N44°32’E; Diyana (Soran): 36°41’N44°35’E; Gali (Qali)

Ali Beg: 36°38’N44°25’E; Hais: 37°13’N43°25’E; Haj Omran:

36°40’N45°05’E; Harir (HarirDagh):36°27’N44°23’E; Jabal

Zhargahta: 35°18’N45°I7’E; Kasimain: 33°22'N44°20’E;

Makhmoor: 35°45’N43°35’E; Mergasor: 36°50’N44° 19’E;

Rawanduz (Rowanduz): 36°38’N44°32E; SafinDaeh (Jabal-

Safin): 36°23’N44° 1 8’E; Shanidr Cave: 36°50’N44°13’E;

Shaqlawa: 36°23’N44°18’E; Shirin Mountain: 36°58’N44°06'E;

Suran (Soran) Area: 36°39’N44°32’E.

KarbaPa Province (Ka)

Uqhaidhir (Ukhaidir): 32°26’N43°36’E; Al -Tar ( Al-Tar caves):

32°28’N32°46’E; EinTamur (Shthatha): 32°33'N32°29'E.

Kurkuk Province (Ku)

Chamchamal-Jarnio: 35°34'N44°55’E; Alton Kobri:

35°44’N44°09'E; TaqTaq: 35°53N44°36’E.

Mayssan Province (Ma)

Abusakhair: 32°32’N47°19’E; Al Kumait: 32°0rN4643’E; Ali

Gharbi: 32°28’N46°42’E; Al-Maimona: 3 1 °4 1 ’N46°58’E; Ama-

ra: 3 1 °50’N47°08’E; Azair: 3 1 °20'N47°26'E; Biet Mohammad:

3I°50’N37°I0’E; Chahala: 31°40’N47°18’E; Daub:

31°22’N47°14’E. Hawizeh: 3 1°39’N47°40’E; Kut al Iniara:

32°30’N45°49’E; Majaral Kabir: 3 1°34'N47°09’E; Musharah

River: 31°50’N47°13’E; Nachr-Chasasch: 30° 50’N47°I0’E;

QalatSalih: 31°32’N47°19'E; Sinn Abtar: 32°30’N45°49’E;

Teeb: 32°03N'47°1 5’E; Umm-Al-Ni’aj: 3 1°37’N47°36’E.

Nenawa (Nineve) Province (Ne)

Ain Tina: 35°35’N42°43’E; Ain Zala (Ain Zalah):

36°44’N42°34’E; Almawan: 36°25’N44°13’E; Aqra (Akre):

36°45’N43°54’E; Aski-kalak: 36°16’N43°38’E; Atrosh:

36°50’N43°20’E; Cora: 36°2 l’N44° 1 0’E; Fishkhaboor:

37°03’N42°22’E; Hamam Al-Alil: 36°10’N43°16’E; Hatra (Al

Hadr): 35°34’N 42°42’E; Jabal Sinjar: 36°23’N41°52’E; Mousel

city: 36°20’N43°06’E; Narwah: 36°42’N44°I0’E.

Muthana Province (IMu)

Busaiyah (Busaiya): 30°5’N46°7’E; Samawa: 31°18’N45°I8’E.

Najaf Province (Na)

Bahr Al-milih (Bahr Al-Najaf): 31°59’N44°15’E; Najaf city:

3I°59’N44°19’E; Seriya: 33°24’N43°4I’E.

Qadissiyab Province (Qa)

Ad Dwaniyah: 3 1 °58’N44°56’E; Al-Hamza: 3 1 °43’N44°58’E;

Al-Shamiyah: 3 1°57’N44°36’E: Ash-Shinaffya:

31°35’N44°39’E; Dalmaj (HorDalmaj): 32°9’N45°3 1’E;

Ghamaz (Gamas): 31°44N’44°37’E.

Salabadin Province (Sa)

Tikrit: 34°35’N43°40’E; Ad Dawr(Daur): 34°29’N43°49’E; Ain

Al-Faras: 34°1 6’N43°1 1 ’E; Al Alam: 34°42’N43°41’E; Al-Jez-

era( Jezeratt Samarra): 35°00’N42°00’E; Baiji (Beaji):

34°56’N43°29’E; Balad: 34°00’N44°09’E; Balad (Beled):

33°58’N44°11’E; Cidir: 34°44’N44°03’E; Dejail:

33°5(VN44°14’E; Dholuiya: 34°05’N44°1 2’E; Duz-khurmato

(Tooz): 34°52’N44°37’E; Jalam (Jalam Albu Ajeel):

34°35’N43°53’E; Jabal Himreen: 34°59’N43°43’E: Makhool

(Jabal Makhool): 35°07’N43°26’E; Qala’t Asharqat:

35°27’N43°13’E; Samara: 34°12’N43°52’E; Sharqat:

35°29’N43°14’E; Tarmiya: 33°40’N44°22’E; Wadi Al-Tharthar:

34°16’N43°27’E.

Bonn zoological Bulletin 64 ( 1 ): 33-58

©ZFMK

56

Omar F. Al-Sheikhly et al.

Sulaymaniyah Province (Su)

Ahmad Awa: 35°17’N46°03’N; Baksai: 32°53’N46°26’E;

Chamchamal: 35°32’N44°50’E; Chami-Rezan:

35°48’N45°01’E; Darbandikhan (Derbendi Khan):

35°18’N45°47’E; Halabja: 35°10’N45°59’E; Hawaraman Area:

35°13’N46°06’E; Hazar Mard: 35°30’N45°20’E; Zagarta (Ja-

belZagarta; Sargat): 35°16’N46°06’E; Kalar: 34°36’N45°19’E;

Kanispika: 35°27’N45°23’E; Koshek: 35°16’N45°47’E; Little

Zab: 35°54'N44°58'E; Maidam: 34°55’N45°37’E; Mavvat:

35°57’N45°24’E; Murdka (Mortka): 35°03’N45°49’E; Parazan:

35°35’N45°49’E; Penjwin: 35°35’N45°57’E; Peramagroon:

35°46’N45°13’E; QaraDaq (Karadagh): 35°17’N45°21’E;

Qashqulai: 35°55’N44°57’E; Sulymaniyah city:

35°33’N45°25’E; Taqia: 35°37’N44°57'E; Zagros Mountains

(Jabal Zagros): 36°00’N45°00'E.

ThiQar Province (Th)

An Nasseriyah: 31°03’N46°16’E; AbuAjaj: 39°49’N46°36’E;

Abu Zirig: 31°06’N46°38’E; Al-Eslah: 31°09’N46°36’E; Al-

Fhood: 30°58’N46°43’E; All Jweaber: 30°54’N46°38’E; Al-

Ta'r:30°56’N56°50’E; Chebaeish: 30°57’N46°59’E; Hor(Hour)

Al Hammar: 30°50’N47°10’E; Kamisiyah: 30°46’N46°29’E;

Kermashiya: 30°47’N46°37’E; Naheat Al Hammar:

30°57’N46°50’E; QannattBani Seed: 30°52’N46°34’E; Suq Al-

Sheiukh: 30°52’N46°29’E; Ur: 30°57’N46°06’E; Zichri (Az

Zichri): 31°03N’47°1 3’E.

Wasit Province (Wa)

Aziziyha: 32°54’N45°05’E; Badra: 33°05’N45°59’E; Kut:

32°29’N45°50’E; Numaniyah: 32°33’N45°25’E; Sewara:

32°54’N44°47’E; Sheik Saad: 32°34’N46°17’E.

Bonn zoological Bulletin 64 (1): 33-58

©ZFMK

Checklist of the Mammals of Iraq

57

58

Omar F. Al-Sheikhly et al.

Table 2. List of mammal species in Iraq evaluated by the lUCN.

Bonn zoological Bulletin 64 (1): 33-58

®ZFMK

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(•

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$

Contents

SMITHSONIAN LIBRARIES

Grebennikov, Vasily V.: 1

Wingless Paocryptorrhinus (Coleoptera: Curcutionidae) rediscovered in Tanzania:

synonymy, four new species and a mtPNA phyloqeoqraphy

Kaiser, Dorkas, Tra-Bi Croulaud Sylvain, Kolo Yeo, Souleymane Konate & Karl Eduard Linsenmair: 16

Species richness of termites (Blattoidea; Termitoidae) and ants (Hymenoptera: Formicidae)

along disturbance gradients in semi-arid Burkina Faso (West Africa)

Al-Sheikhly, Omar F., Mukhtar K. Haba, Filippo Barbanera, Gabor Csorba & David L. Harrison: 33

Checklist of the Mammals of Iraq (Chordata: Mammalia)

Book Review

Bohme, W. (2014) 32

Herpetology in Bonn

(Philipp Wagner)

Cover illustration:

Ruppell's Fox Vulpes rueppellii from Iraq (Photo: Omar F. Al-Sheikhly) (this volume, pp. xxx-xxx).

Bundesministerium

furBildung

und Forschung

Ministerium fiir Innovation,

Wissenschaft und Forschung

des Landes Nordrhein-Westfalen