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Small mammals recovered from owl pellets from Syria
(Mammalia: Chiroptera, Rodentia)
by Adwan Shehab, Ahmad Daoud, Dieter Kock, and Zuhair Amr
Abstract. A total of 23 species of small mammals (5 Chiroptera and 18 Rodentia) were recov-
ered from owl pellets collected from several localities in Syria. The occurrence of Taphozous
nudiventris magnus
on the Syrian Euphrates is part of a northwestern extension of the Iraqi
population, extending into southern Turkey. The present material fills a distribution gap for Ot-
onycteris hemprichii
between the Syrian, Turkish and Iraqi localities. Records for two species of
the genus Microtus, Microtus (Microtus) socialis and the problematic M. (M.) philistinus, are
given. Comparative material was used to confirm the identity of both species. The known range
for M. socialis is thereby extended considerably further eastwards. Similarly, the present records
of Cricetulus migratorius and Mus macedonicus extend the range of both species eastwards. The
record of Eliomys melanurus represents a noteworthy range extension further to the north.
Kurzfassung. Insgesamt 23 Arten von Kleinsäugern (5 Chiroptera, 18 Rodentia) wurden aus Eu-
lengewöllen identifiziert, die an zahlreichen Fundorten in Syrien gesammelt wurden. Das Vor-
kommen von Taphozous nudiventris magnus am syrischen Euphrat ist Teil des Verbreitungsge-
bietes, das sich vom Irak nordwestlich bis in die südliche Türkei erstreckt. Der Fund von Otonyc-
teris hemprichii
fällt in die Lücke zwischen bekannten türkischen, syrischen und irakischen
Nachweisen. Zwei Arten der Gattung Microtus wurden an Hand von Vergleichsmaterial identifi-
ziert: Microtus (Microtus) socialis und die problematische Art M. (M.) philistinus. Das Vorkom-
men von M. socialis erweitert sich damit beträchtlich nach Osten, ebenso auch durch die östlichen
Nachweise für Cricetulus migratorius und Mus macedonicus. Für Eliomys melanurus wird das
Vorkommen in nördlicher Richtung erweitert.
Key words. Middle East, Levant, small mammals, owl pellets, range extension.
In the course of studying the biology and control of social voles of the genus Microtus in Syria, the senior author, AHMAD DAOUD, and ZUHAIR AMR collected owl pellets from sev-eral localities in the country. Previously small mammals identified from Syrian owl pellets have been reported by RZEBIK-KOWALSKA & NADACHOWSKI (1978), KOCK & NADER (1983), NADER & KOCK (1983), AUFFRAY et al. (1990), NADACHOWSKI et al. (1990), KRUPP & SCHNEIDER (1991), KAHILA & HORWITZ (1994), KOCK et al. (1994), EBENAU (1996), KOCK (1998), OBUCH (2001), HUTTERER & KOCK (2002), and SHEHAB et al. (2003). These studies resulted in several additions to the mammal fauna of Syria: Suncus etruscus by RZE-BIK-KOWALSKA & NADACHOWSKI (1978), Crocidura katinka by HUTTERER & KOCK (2002), Rhinolophus mehelyi, Taphozous nudiventris, Myotis capaccinii, and Eptesicus bottae by EBENAU (1996), Musspretoides” [= macedonicus] by AUFFRAY et al. (1990) or Mus cf. abbotti [= macedonicus] by NADACHOWSKI et al. (1990), Microtus irani by KOCK & NADER (1983), Gerbillus henleyi, G. cheesmani, and G. mesopotamiae by KOCK (1998). Rarer spe- Zoology in the Middle East 33, 2004: 27–42. ISSN 0939-7140 Kasparek Verlag, Heidelberg cies were documented by OBUCH (2001: Eliomys melanurus, Dryomys nitedula) and SHEHAB et al. (2003: D. nitedula). Despite these positive results, however, the mammals contained in owl pellets do not ex- ceed a certain size due to the ability of the owls to deal with the prey item. Consequently, of the larger rodents such as Nesokia indica and Tatera indica, mainly subadult individuals are represented in the prey items. Furthermore, diurnal prey species such as Acomys sp. are only rarely caught by the owls. The open habitat preferred for hunting by Tyto alba and Athene noctua also offers few or no chances to prey upon forest species such as Sciurus anomalus and Sylvaemus mystacinus. Finally, both owl species are opportunistic hunters and will prey on those species which are most abundant, e.g. Microtus sp. during a population outbreak or Mus musculus in cultivated areas. The pellets studied for this paper yield additional distribution data for the rather incom- pletely known Syrian small mammals and will help to identify faunal realms and centres of species diversity in Syria. The ranges of mammal species have been detailed and mapped by HARRISON & BATES (1991), so far as they were known to occur in Syria up to 1991. Material and methods
Cave explorers from Germany have investigated the Euphrates valley and have discovered sev-eral places suitable for their research interest. A number of owl pellets were collected and the mammal remains contained therein were deposited in the Senckenberg Museum and identified. Biologists from the latter institution collected owl pellets in the Khabur River region for compari-son with the antique fauna of Tell Sheikh Hammad. Further series of pellets were later collected from various localities (see gazetteer), some of them identical with those sampled by the German cave exploration group. Each pellet was placed in a Petri dish containing water for at least 10 minutes. Pellets were teased apart and their con-tents were carefully isolated into separate vials. For most of the sites, positive identification of the breeding owl(s) was made. Representative materials were compared with reference specimens at the Forschungsinstitut Senckenberg and Museum (Frankfurt a. M., Germany) collected from Syria or the neighbouring areas of Turkey, Iraq, Jordan or Saudi Arabia. The collection of the latter institution houses further unpublished mammal specimens found in owl pellets, and these have been included in the present paper. The initials given with each locality are for the most frequently cited collectors’ names; less frequent ones remain unabbreviated. Unless otherwise specified, all osteological material listed is from owl pellets; if the predator was identified, its name is included as an abbreviation. Abbreviations: A.D. = A. DAOUD; A.S. = A. SHEHAB; A.n. = Athene noctua; btw. = between
(two localities); C.E. = C. EBENAU; D.K. = D. KOCK; lft = left; mand = mandible/-les; maxl =
maxilla/-lae; R.K. = R. KINZELBACH; rt = right; ro = rostrum; T.a. = Tyto alba; SMF = Sencken-
berg Museum, Frankfurt a. M.
Results and discussion
A total of 24 species of small mammals representing two orders (Chiroptera and Rodentia) were recovered from owl pellets collected from various localities in Syria. Only two owl species were positively identified, the Barn Owl, Tyto alba, and the Little Owl, Athene noc-tua. Tab. 1. Localities in Syria where owl pellets have been collected, together with the owl species observed. Locality
Owl species
Ebla ruins (Tell Mardikh-Ebla), 26 km SE Idlib Euphrates Valley, 5 km S of, 12 km SE ar’Raqqa Heraqla(btw.) Maharda and Shaizar, 35°57'N, Okersheih, Euphrates Valley, 12 km SE ar’Raqqa Qal`aat al-Hosson (Crak des Chevaliers) 34°47'N, Qal’aat ar’Rahba, S-bank Euphrates 34°58'N, Qal’aat Sukkara, 2 km SW of, Jebel Abdul Aziz Qasret Mohammed Ali, S-bank Euphrates, 7 km S ar’Raqqa 35°52'N, Qater Maghara, near Qasret Moh. Ali Sergiopolis = ar’Rasafeh Shaizar and Maharda (btw.), 25 km NW Hamah Chiroptera: Emballonuridae
Taphozous (Liponycteris) nudiventris magnus Wettstein, 1913
Material. Halabiye, 31.V.1989, colony in crevices of antique city wall, 1 ƃ, 2 Ƃ (3 skulls, 3 in alco-
hol), SMF 74084-6, D.K. – Qater Maghara, 22.II.1993, n2 (T.a.), SMF 80666, C.E. [EBENAU 1994b] –
Qasret Mohammed Ali, 19.III.1996, 2 ad, 4 juv (T.a.), SMF 84474-79, C: EBENAU [EBENAU 1966];
ibid., 28.VI.1998, A.S.
Measurements. HB ƃ 99.3, Ƃ 107-108.3, T ƃ 32.8, Ƃ 31.2-32.3, HF ƃ 19, Ƃ 16.3-18.6, E ƃ 27.2, Ƃ
26.6-28.6, FA ƃ 80.3, Ƃ 77.5-80.2. – GrL skull ƃ 28.2, Ƃ 28.6-29.7, GrL to C1/ ƃ 27.6, Ƃ 28.3-29.4,
CbL ƃ 26.5, Ƃ 26.1-26.2, CbL to 1/ ƃ 25.9, Ƃ 25.5-25.6, Mast ƃ 14.85, Ƃ 15.7-15.83, BB ƃ 12.3, Ƃ
12.2-12.55, Zyg ƃ 16.9, Ƃ 17.63-17.9, C-C1/ ƃ 6.4, Ƃ 6.6-6.8, M-M3/ ƃ 11.7, Ƃ 11.6-11.8, C-M3/ ƃ
alv 11.5, Ƃ 11.1-11.3, dto crown ƃ 12.4, Ƃ 12.0-12.1, IOR ƃ 8.4, ƂƂ 8.6-9.0, POR ƃ 5.1, Ƃ 5.1-5.3,
mand cdl ƃ 22.4, Ƃ 21.9-22.4, mand ang ƃ 22.2, Ƃ 22.6-23.0, C-M/3 ƃ 13.9, Ƃ 13.4-13.6.
Distribution: Three skins of T. nudiventris were collected by the “Expedition to the Euphra-
tes 1850” (DOBSON 1878), but their original data were lost in an accident during that expedi-
tion while on the Euphrates around Dura Europos; the collecting locality could have been
inside presentday Syria. THOMAS (1915) included this material in the description of T. kach-
hensis babylonicus
(a junior synonym of magnus Wettstein 1913) from Mesopotamia on the
Euphrates. EBENAU (1996) reported this species from owl pellets collected on the Euphrates
(see material listed above). The measurements of complete Syrian specimens (see above) prove that they represent the large-sized subspecies T. n. magnus. The occurrence of T. n. magnus on the Syrian Euphrates is part of a northwestern extension of the Iraqi population, which extends into southern Turkey to near Nizp, Gaziantep Prov. (SACHANOWICZ et al. 1999). The Mespotamian population is geographically widely separated from the neighbouring subspecies to the south-west, T. n. nudiventris (Cretzschmar 1830) in Palestine (TRISTRAM 1884, THOMAS 1915, J. AHARONI 1930, HARRISON 1964, ATALLAH 1977, HARRISON & BATES 1991: Lake Kinneret; ALLEN 1915, CLARKE 1977: walls of Jericho; BODENHEIMER 1935: Wadis nr Tiberias; HARRISON 1964: Wadi Ahmud N of Lake Tiberias). Chiroptera: Vespertilionidae
Myotis capaccinii (Bonaparte, 1837)
Material. Qasret Mohammed Ali, 28.VI.1998, n1 (maxl toothrow), A.S.
Remarks. Previous records labelled “Syria” (NEHRING 1886: Vespertilio Capacinii [sic];
JENTINK 1888: Vespertilio dasycneme) may have originated from parts of the Near East
which are currently in Turkey, Lebanon or Palestine. The species was first recorded from
presentday Syria by EBENAU (1996) and WALTER & EBENAU (1997), who observed M. ca-
at Qater Maghara and found remains in “Pigeon well cave” near ar’Raqqa. This
occurrence on the Euphrates falls into the distribution gap between Iraq (SANBORN 1956:
Kish ruins, as M. c. bureschi Heinrich 1936; KHAJURIA 1988: near Haditha, as M. capaceini
[sic!: 398] bureschi; ABUL-HAB & SHIHAB 1989: Baghdad), Lebanon, Jordan, and Israel (see
HARRISON & BATES 1991, as M. c. bureschi). Since SANBORN (1956) first identified Iraqi M.
with the subspecies bureschi, described from Bulgaria, most subsequent records
from the Near East have been assigned to this subspecies. However, TOPAL (1997) states that
Iraqi M. capaccinii has significant differences when compared with typical M. c. bureschi.
The subspecific status of the Near East population thus needs to be studied in detail.
Pipistrellus kuhlii (Kuhl, 1817)
Material. ar’Rasafeh (Sergiopolis), 28.III.1996, n1, SMF 84495, C.E. – Desert 5 km S of Euphrates
Valley, 18.III.1996, n16, SMF 84413-21, C.E. – Qater Maghara, 19.III.1996, n1, SMF 84481, C.E. –
Okersheih, 20.III.1996, n4, SMF 84445-48, C.E. – Dura Europos (as’Salahiya), 17.V.1989, n11 (ro;
T.a.), SMF 80623, D.K. – Qasret Mohammed Ali, 28. VI.1998, A.S.
Remarks. This widely distributed bat has previously been found in Syria in owl pellets, as
reported by NADER & KOCK (1983: Qal’aat ar’Rahba) and by EBENAU (1996: Qater
Eptesicus bottae hingstoni Thomas, 1919
Material. Cliffs at S-side of Euphrates at Qater Maghara Cave, 19.III.1996, n1 (fragmented skull &
mand; T.a.), SMF 84480, C.E.
Measurements. Breadth across upper canines (alveoli) 5.1; breadth across crowns of M3/
7.04; length of upper toothrow C-M3/ (alveoli) 5.66; postorbital constriction 3.85; length of
mand from condyle 11.93; ibid., from processus angularis 12.14; length of lower toothrow
C-M/3 (alveoli) 6.36.
Taxonomy. The few measurements available are of little diagnostic value, as most of the
teeth have been lost. But the size of rostrum, breadth of M1/ and short M3/ agree with exten-
sive comparative material of E. b. hingstoni as listed by NADER & KOCK (1990).
Distribution. The species was first recorded for Syria by EBENAU (1966), who observed E.
bottae at Qater Maghara and found its remains in owl pellets at the same locality in 1993.
With the present material, the range of the subsp. hingstoni is extended from the nearest
known locality at al-Hadithah in the Iraqi Euphrates Valley (KHAJURIA 1988), northwest-
wards along this river, midway towards the nearest known occurrence of E. bottae anato-
Felten, 1971, at Toprakkale near Ceyhan, Turkey (NADER & KOCK 1990).
Otonycteris hemprichii Peters, 1859
Material. Qal'aat ar’Rahba, 17.V.1989, n3 (3 rt mand; T.a.), SMF 74083, D.K.
Measurements. Length of mand from condyle 15.0-15.18; ibid. from processus angularis
15.3-15.75; lower toothrow C-M/3 (alveoli) 8.6-8.75.
Remarks. The shape of the mandibles and position of tooth alveoles, and less significantly,
their size agree closely with O. hemprichii from Cairo (SMF 25268: mand condylar length
16.0; ibid., angular length 15.94) and Osh region, Kyrgyzstan (SMF 77781: mand condylar
length 16.4; ibid., angular length 16.8). This desert bat species has also been reported as the
prey of owls in the central Sahara by HEIM DE BALSAC (1965: Djanet, Tassili des Ajjers,
Distribution. Even before 1754, RUSSELL (1794) knew of two forms of “Vespertilio muri-
” in Aleppo Bazaar, one of which was only rarely observed: “it was white and had ears
longer than those of the former”. This might well have been an early observation of O. hem-
in Syria, as the whitish colouration would exclude a Plecotus sp. It was more than
two centuries before HARRISON (1964) could record a reliably identified specimen from the
Syrian Desert and (1972) from Qaryateine [= Karyatein or Karyatin, of authors]. This latter
locality, but based on other specimens, was doubtfully located in Jordan by QUMSIYEH
(1985). The present material fills a distribution gap between the Syrian, Turkish and Iraqi
localities as mapped by NADER & KOCK (1983); other distribution maps are either incom-
plete (HARRISON & BATES 1991) or have overestimated the range (GHARAIBEH & QUMSIYEH
Rodentia: Dipodidae
Jaculus jaculus (Linnaeus, 1758)
Material. Dura Europos (as’Salahiya), 17.V.1989, n2 (ro; T.a.), SMF 80638-9, D.K. – Qal’aat
ar’Rahba, 14.III.1979, n3 (2 upper & 3 lower tooth rows; T.a.), SMF 60373, R.K.; ibid., 17.V.1989, n1
(skull; T.a.), SMF 80650, D.K. – 5 km S of Euphrates Valley, 18.III.1996, n6 (partial skulls), SMF
84434-39, C.E. – ar’Rasafeh (Sergiopolis), 28.III.1996, n3 (partial skulls), SMF 84501-03, C.E.; ibi-
dem, 28.VI.1998, n3 (2 skulls, 3 mxl., 3lft & 3 rt mand), A.S. – Okersheih, 28. VI.1998, n1 (1 rt mand),
Distribution. This species is apparently less common in Syria than Allactaga euphratica
(see below). THOMAS (1921, 1922) had specimens from Karyatein; J. AHARONI (1930) men-
tions it from the northern Syrian Desert. HARRISON (1972) records it from Dayr az’Zawr,
Palmyra, and from 40 km W of Palmyra. NADACHOWSKI et al. (1990) collected it 100 km E
of Damascus at 30°36’N, 36°44’E.
Note. J. jaculus is commonly preyed upon by nocturnal raptors, as reported e.g. from Ara-
bian countries by NADACHOWSKI et al. (1978: Hatra, Iraq), KADHIM (1979: Iraq), CLAYTON
(1991: Kuwait), EVANS & BATES (1993: Harrat al Harrah, N-Saudi Arabia), and KOCK &
NADER (1996: Jubail Wildlife Sanctuary, E-Saudi Arabia).
Allactaga euphratica Thomas, 1881
Material. Tell al Budeiri, 23.V.1989, n1 (rt mand; T.a.), SMF 80563, D.K. – Qal’aat ar’Rahba,
17.V.1989, n1 (ro; T.a.), SMF 80651, D.K. – 5 km S of Euphrates Valley, 18.III.1996, n4 (partial
skulls), SMF 84440-43, C.E. – ar’Rasafeh, 28. VI.1998, n1 (skull), A.S.
Distribution. The above records are all within the known Syrian range of this species (HAR-
RISON & BATES 1991), which is to be complemented by records from ar’Raqqa, Bahrat
[Lake] Hatuniya, and al-Hasaka (BAUER 1988) and at 2 km and at 8 km SE Tell Sheikh
Hamad (KOCK et al. 1994), where A. euphratica was found in owl pellets, as it was previ-
ously in Qal’aat ar’Rahba (KOCK & NADER 1983); the latter locality is confirmed by the
present material.
Rodentia: Gliridae
Eliomys melanurus (Wagner, 1839)
Material. 2 km SW Qal’aat Sukkara, Jebel Abd al-Aziz, 2.X.1988, n1 (rt maxl; A.n.), SMF 80664,
Distribution. This record represents a noteworthy range extension to the north for E. mela-
. RUSSELL (1794, as Mus quercinuus) records it at Aleppo before 1754. Most probably
this was the species found by LORTET (1883, as Myoxus glis) as rather common at Zabadani,
upper Barada Valley. J. AHARONI (1917) collected it at Dschebé [= Jebel al-Geba], N of
Qaryatein; however, the skull of this specimen is a Graphiurus sp. (NADER et al. 1983).
Despite these few records, BODENHEIMER (1935) considered E. melanurus to be common in
the Syrian desert though he only cited Mt. Hermon as a precise locality. Later, KAHMANN
(1981) examined several more recent specimens from Mt. Hermon and from the Golan
Heights (1200-2100m) at Mas’ada Forest (1000m), Birket Bab el-Haoua (900m), Ein Qunya,
and Dan. From Damascus it was reported by GOLEMANSKY & DARWISH (1993), KRYŠTUFEK
& KRAFT (1997) recorded it from Halabiye [SMF 74079, D.K.], and finally OBUCH (2001)
found its remains at Halabiye, Yabrud, at Buransk in T.a. pellets, and at Palmyra in Asio otus
pellets. So far is known at present, E. melanurus ranges north to Harran, 30 km S Urfa in S-
Turkey (MISONNE 1957: sight record) and north-west into Iraq to Hatra [al-Hadr, 35°34’N,
42°42’E, 90 km SW Mossul] (NADACHOWSKI et al. 1978: from owl pellets) and to Mossul
(KAHMANN 1978, 1981). Together with its preferred arid, rock habitat, E. melanurus is
probably more widely distributed in Syria than is documented at present.
Rodentia: Spalacidae
Nannospalax ehrenbergi (Nehring, 1898)
Material. Tell al Budeiri, 23.V.1989, n1 subad (skull & mand; T.a.), SMF 80562, D.K.
Distribution. The mole rat is widely distributed in W- Syria including the Golan Heights.
Records are much scarcer in the interior of the country and are available only from Tall
Abiad and Ain Aarous (MISONNE 1957), and at CeylanpÕnar, Turkey, opposite Ras al-Ain (LEHMANN 1969). Despite its subterranean mode of life, N. ehrenbergi is preyed upon by owls (MISONNE 1957: A.n.), also at Crak des Chevaliers (NADACHOWSKI et al. 1990), and in the Lebanon Mts. (BATE 1945: Asio otus). Rodentia: Cricetidae
Cricetulus migratorius (Pallas, 1773)
Material. Btw. Shaizar and Maharda, 26.III.1980, n2 (ro with mand, 2 maxl with paired mand), SMF
60375, R.K. – ar’Rasafeh (Sergiopolis), 28.III.1996, n7 (T.a.), SMF 84497-500, C.E.
Distribution. Known records were summarised by HARRISON (1972) and are concentrated in
the Aleppo region, at Hama and al-Karyatein. Remains of this species in owl pellets found at
Qal’aat al-Hosson (Crak des Chevaliers) were studied by PRADEL (1981) and NADACHOWSKI
et al. (1990). On Mt. Hermon C. migratorius was found by YOM-TOV (1988) and FILIPPUCCI
et al. (1989). The present record from ar’Rasafeh extends the species range eastwards.
Mesocricetus auratus (Waterhouse, 1839)
Material. Ebla ruins [Tell Mardikh-Ebla], 31.V.1998, 1 juv (ro & 2 mand), SMF 88363, A.D.
Taxonomy. The present remains of a juvenile differ from the closely related M. brandti
(Nehring, 1898) by shorter upper and lower molar rows and agree in this character with adult
M. auratus (SMF 82129-31, Aleppo: tooth cusps worn), as already noticed by B. AHARONI
(1932). The juvenile rostrum has the posterior palatal emargination more U-shaped than V-
shaped, and closer to the last upper molar than illustrated by YIöIT et al. (2000: Fig. 2a), but
it is thought that the appearance changes with growth.
Distribution. Since its first mention in Syria (RUSSELL 1794: Mus cricetus Linn.) and its
description from specimens collected at Aleppo (WATERHOUSE 1839), the range of this ham-
ster appeared to be restricted to the Aleppo region (B. AHARONI 1932: Aleppo; Biliramun,
NW of Aleppo; Azaze, N of Aleppo; NEHRING 1902a, b; OSBORN 1965 [sub M. auritus [sic!]
nr Aleppo], MURPHY 1971, HARRISON 1972, KUMERLOEVE 1975, ATALLAH 1977; SMF
82129-31: Al-Collie Farm, SE of Jerablus). Only BODENHEIMER (1920) mentions the species
as straying S to Beirut. However, this was never confirmed, and very likely originates from
TRISTRAM’S (1884) note of Cricetus nigricans Brandt, 1832 [= M. brandti] being collected at
the foot of Mt. Lebanon and of specimens of M. auratus seen in the Museum at Beirut. Nev-
ertheless, M. brandti is recorded by HAMAR & SCHUTOWA (1966) from ”Syria and Turkey”
based on four specimens examined in the Museum für Naturkunde, Berlin, and that species is
listed by MUSSER & CARLETON (1993) as occurring in Syria (and S to Lebanon and Israel).
However, at our request, M. ADE, formerly curator of the mammal collection in Berlin,
searched unsuccessfully for Syrian specimens. We do not know of any confirmed evidence
(see also HARRISON 1972). – Recently, GATTERMANN et al. (2001) have recorded M. auratus
from Jerablus [SMF 82129-31], and from Albel and Sheikh Riek (Rieh), both 50 km NE of
Aleppo. Specimens from the present locality have been published by SHEHAB et al. (1999)
and the range of the species is now known to extend into southern Turkey (DOöRAMACI et al.
1994: Nizip, Gaziantep province; YIöIT et al. 2000: Kilis).
Although the related M. brandti (Nehring, 1898) was found in owl pellets from Turkey (STEINER & VAUK 1966, SICKENBERG 1971) and MURPHY (1971) mentions owls as predators of M. auratus, this hamster has not previously been found in owl pellets in Syria (SHEHAB et al. 1999). Microtus (Microtus) socialis (Pallas, 1773)
Material. Ebla ruins (Tell Mardikh-Ebla), 31.V.1998, n1 (ro with mand), SMF 88357, A.D. – Athar
Maez, VIII.1998, n1 (ro, 1 lft & 1 rt mand), SMF 88361, A.D. – Shah`Ranar, 27.VI.1998, n2 (1 skull, 2
lft & 2 rt mand; A.n.), SMF 88358, A.D. – btw. al’Hamra and Sahl, Al-Ghab, 26.X.1998, n2 (1 skull, 2
lft & 4 rt mand), SMF 88355-56, A.D. – Qal`aat al-Hosson (Crak des Chevaliers), 27.IV.1998, n2 (2 ro,
1 lft & 2 rt mand; T. a.), SMF 88362, A.D. – Deir Khabeih, III.1998, n2 (2 ro, 2 rt & 2 lft mand), SMF
88359-60, A.D. – Okersheih, 28. VI.1998, n2 (2 rt mand), A.D.
Comparative material. Iran: Kurdistan, 25.III.1962, n1 (skull, skin), SMF 44716, Institute Pasteur
Tehran. – Ushan, Elburs Mts., 30 km NNE Teheran, 19.V.1974, 16.-17.V.1975, 12 ƃ, 9 Ƃ (21 skulls,
21 skins), SMF 47580, H. FELTEN, D. K. & K. WALCH. – Van, NE-Azerbaijan, 23.V.1975, 2 ƃ immat,
1 ƃ ad (3 skulls, 3 skins), SMF 48982-84, D. K. – Gedailu, 12 km S Aslandus, N-Azerbaijan,
24.V.1975, ƃ immat (skull, skin), SMF 48985, D.K.
Taxonomy. The skull characters of M. socialis from Turkey and Syria have been described
by STORCH (1971, 1972), MORLOCK (1978) and KOCK & NADER (1983). This species can be
differentiated from the sympatric M. philistinus (see below) by having the bullae still smaller
than M. guentheri (8.4-8.5 in Syrian material), short upper and lower molar rows (upper
tooth row in Syrian material 4.8-5.8), and the suprameatal triangle shorter (2.3-3.0 in Syrian
Distribution. Older records of voles from Syria identified as “M. socialis” are doubtful.
TRISTRAM (1866, 1884), NEHRING (1902b); all as “Arvicola socialis Desmarest”: desert of
Sahara plains N of Damascus [= Es-Sahara plain]. – BODENHEIMER (1920): Arvicola socialis
var. syriaca and var. cinerescens: deserts south of Damascus. – KOCK & NADER (1983) stud-
ied Syrian Microtus sp. and, while rejecting most records previously identified as “M. so-
”, they identified true M. socialis from owl pellets collected between Shaizar and Ma-
harda, 25 km NW Hamah, and accepted as correctly identified material from Aleppo, re-
ported by ATALLAH (1978). The present record extends the species range surprisingly far
eastwards into the ar’Raqqa area, though apparently confined to the Euphrates valley with its
moister soil and only rarely represented in owl pellets. In western Syria, M. socialis occurs
sympatrically with M. philistinus at Ebla ruins (Tell Mardikh-Ebla), Athar Maez, Qal`aat al-
Hosson, and btw. al’Hamra and Sahl.
Microtus (Microtus) philistinus Thomas, 1917
Material. Ebla ruins (Tell Mardikh-Ebla), 31.V.1998, n1 (skull, 2 mand), SMF 88348, A.D. – Athar
Maez, VIII.1998, n1 (ro, lft mand), SMF 88347, A.D. – Qal`aat al-Hosson (Crak des Chevaliers),
27.IV.1998, n3 (1 ro, 3 lft & 2 rt mand; T.a.), SMF 88349, A.D. – btw. al’Hamra and Sahl, Al-Ghab,
26.X.1998, n5 (1 skull, 3 ro, 3 lft & 1 rt mand), SMF 88350-54, A.D.
Comparative material. Syria: SMF 64210-11, 15 km SE Aleppo, 3.IV.1961, 1 ƃ 1 Ƃ (2 skins), R.E.
LEWIS (V-6205,V-6207, ex AUB M-314, M-316) (= irani sensu KOCK & NADER 1983). – Orontes
River N of Lake Homs, 1.IV.1979, ƃ (skull, skin), SMF 60411, R.K. (= irani sensu KOCK & NADER
1983). – Palestine: Givat Brenner, nr Tel-Aviv, 27.XI.1977, 1ƃ 2ƂƂ (3 skulls, 3 skins), SMF 54960-
62, L. COHEN (= irani sensu KOCK & NADER 1983). – Ayanot, 31°55’N - 34°46’E, 1.VII.1978, 4ƃƃ
(4 skulls, 4 skins), SMF 55565-68, L. COHEN (= irani sensu KOCK & NADER 1983).
M. guentheri (Danford & Alston, 1880): Turkey: Incekum, Antalya Prov., 24.V.1966, Ƃ (skull, skin), SMF 37488. - Abant, Bolu Prov., 7.-8.IX.1960, 2 ƃ 5 Ƃ (7 skulls, 7 skins), SMF 37474-80, H. KAHMANN. – Ankara, see MORLOK (1978). – Greece: Thivai, Boeotia, 21.II.1961, 1 ƃ 2 Ƃ (3 skulls, 3 skins), SMF 26979-81, I. ONDRIAS. M. irani Thomas, 1921: Turkey: Aliúam, ElazÕ÷ Prov., 20.IX.1971, ƃ (skull, skin), SMF 42347. – Iran: Pahlevi Dej, Gorgan Prov., 13.V.1975, 3 ƃ immat 1Ƃ subad (4 skulls, 4 skins), SMF 48986-89, D.K. - 18 km W Karadj, 35°48’N, 51°00’E, Tehran Prov., 16. & 28.IV.1974, 4 Ƃ (4 skulls, 4 skins), SMF 47576-79, H. FELTEN & K. WALCH. Taxonomy. The differentiation of species of the genus Microtus in the Near East has not yet
been definitively resolved. Karyotypes are not available for most specimens in collections.
Skull morphology therefore has to be used to differentiate the Microtus sp. of the Near East
(see references quoted above for M. socialis). Amongst these, M. philistinus according to its
original description (THOMAS 1917) is characterised by a low rostrum (high in M. guentheri
and its subspecies), and larger bullae (in M. guentheri clearly smaller). The present material
shows these characters and is here considered to represent M. philistinus. It agrees with M.
in having the molar tooth rows similar in length (6.0-6.8 in Syrian M. philistinus), the
rostrum lower than in M. guentheri, the bullae larger than in M. guentheri, but still smaller
(8.8-10.1) than M. irani, and the suprameatal triangle longer (3.1-3.45) than in M. socialis
(see above)
Distribution. B. AHARONI (1932: sub M. philistinus; NEUHÄUSER 1936a, b: sub M. (Sum-
) güntheri shevketi; HARRISON 1972: sub M. socialis guentheri; ATALLAH 1978: sub
M. guentheri) records this vole only from Azaze, N Aleppo, and HARRISON (1972: partim:
pl. 191, no. HARR.2.4644; see KOCK & NADER 1983 sub irani) from 15 km SE Aleppo.
A Turkish population of M. socialis from Nizip, Gaziantep Prov., close to the Syrian bor- der, studied by KEFELIOöLU (1995), was assigned to M. guentheri by ÇOLAK et al. (1997). At Kilis, also adjoining the Syrian border, these authors identified M. irani (reddish-brown colouration). However, M. irani has bullae on average smaller than M. guentheri (ÇOLAK et al. 1997: Tab. 1), but based on the reddish hue of their fur colour and the shape of the skull (ÇOLAK et al. 1997: Fig. 4) these M. irani could possibly belong to M. philistinus. Meriones crassus Sundevall, 1842
Material. Ar`Rasafeh, 28.VI.1998, n1 (ro, lft mand; A.n.), A.S.
Distribution. The range in Syria was detailed by KOCK (1998) and the present material
confirms his findings.
Meriones tristrami Thomas, 1892
Material. Bosr al’Sham, 1997, n13 (11 maxl, 12 lft & 13 rt mand), A.S. – Qal`aat al-Hosson,
1.XII.1997, n4 (3 maxl, 3 lft & 4 rt mand), A.S.; ibidem, 27.IV.1998, n5 (5 maxl, 4 lft & 4 rt mand),
A.S.; ibidem, 25.VI.1998, n5 (1 skull, 4 maxl, 3 lft & 5 rt mand), A.S. – Deir Khabeih, 24.III.1998, n3
(3 maxl, 3 lft & 3 rt mand), A.S. – Qal`aat Sheizar, 31.V.1998, n8 (7 maxl, 8 lft & 7 rt mand), A.S. –
Apamea, 31.V.1998, n3 (3 maxl, 2 lft & 2 rt mand), A.S. – al’Hamra, 31.V.1998, n11 (11 skulls, 5
maxl, 6 lft & 6 rt mand), A.S.; ibidem, 27.VI.1998, n16 (6 skulls, 14 maxl, 14 lft & 16 rt mand), A.S. –
Qal`aat Ja’bar, 28.VI.1998, n3 (3 maxl, 2 lft & 1 rt mand), A.S. – ar’Resafeh, 28.VI.1998, n12 (12
skulls, 26 maxl, 30 ft & 30 rt mand), A.S. – Qasret Mohammed Ali, 28.VI.1998, n13 (13 maxl, 8 lft &
10 rt mand), A.S. – Okersheih, 28.6.1998, n7 (7 maxl, 6 lft & 6 rt mand), A.S. – Heraqla, 29.VI.1998,
n4 (4 maxl, 2 rt mand), A.S.
Remarks. This widely distributed species is often found in owl pellets (e.g. KOCK & NADER
1983, KOCK et al. 1994, KOCK 1998).
Meriones lybicus Lichtenstein, 1823
Material. ar’Rasafeh, 28.VI.1998, n2 (2 ro; A.n.), A.S.
Remarks. This is the first record of the remains of the Lybian jird from owl pellets in Syria.
Several studies in Jordan have failed to report this species among pellets of the three owls
occurring in Jordan (AMR et al. 1997, AL-MELHIM et al. 1997, RIFAI et al. 1998, 2000).
Gerbillus mesopotamiae Harrison, 1956
Material. Okersheih, 28.VI.1998, n2 (2 ro, 1 lft mand; A.n.), A.S.
Remarks. Recently recorded from several localities in Syria by KOCK (1998).
Tatera indica taeniura (Wagner, 1843)
Material. Qasret Mohammed Ali, 28.VI.1998, n3 (3 maxl, 2 lft & 2 rt mand), A.S. – Okersheih,
28.6.1998, n4 (4 maxl, 4 lft & 3 rt mand), A.S. – Heraqla, 29.VI.1998, n1 (1 maxl), A.S.
Distribution. The range in Syria has been detailed by KOCK (1998), including several re-
cords from owl pellets.
Mus musculus praetextus Brants, 1827
Material. Bosr al’Sham, 1997, 4 maxl, 5 lft, 4 rt, A.S. – Qal`aat al-Hosson, 1.XII.1997, n13 (2 maxl,
13 lft & 11 rt mand), A.S.; ibidem, 27.IV.1998, n5 (5 maxl, 4 lft & 4 rt mand), A.S.; Qal`aat al-Hosson,
25.VI.1998, n19 (2 skulls, 7 maxl, 19 lft & 10 rt mand), A.S. – Deir Khabeih, 24.III.1998, n11 (5 maxl,
6 lft & 11 rt mand), A.S. – Qal`aat Sheizar, 31.V.1998, n8 (7 maxl, 8 lft & 7 rt mand), A.S. – Apamea,
31.V.1998, n34 (29 maxl, 34 lft & 26 rt mand), A.S. – al’Hamra, 31.V.1998, n34 (22 skulls, 34 maxl,
33 lft & 28 rt mand), A.S.; ibidem, 27.VI.1998, n65 (65 skulls, 27 maxl, 37 lft & 36 rt mand), A.S. –
Qal`aat Ja’bar, 28.VI.1998, n1 (1 maxl, 1 lft & 1 rt mand), A.S. – ar’Rasafeh, 28.VI.1998, n31 (27
skulls, 28 maxl, 26 lft & 31 rt mand), A.S. – Qasret Mohammed Ali, 28.VI.1998, n108 (69 skulls, 78
maxl, 103 lft & 108 rt mand), A.S. – Okersheih, 28.VI.1998, n22 (13 skulls, 11 maxl, 22 lft & 22 rt
mand), A.S. – Heraqla, 29.VI.1998, n15 (10 skulls, 6 maxl, 7 lft & 15 rt mand), A.S. – Palmyra, 30.
VI.1998, n4 (1 skull, 1 maxl, 4 lft & 2 rt mand), A.S. – Btw. Shaizar and Maharda, 26.III.1980, n2,
SMF 60388, R.K. – ar’Resafeh (Sergiopolis), 28.III.1996, n9, SMF 84550-54, C.E. – 5 km S of Euph-
rates Valley, 18.III.1996, n6, SMF 84444, C.E. – Euphrates Valley nr Qasret Mohammed Ali,
19.III.1996, n100, SMF 84486-93, C.E. – Okersheih, 20.III.1996, n61, SMF 84457-71, C.E. – Qater
Maghara, 22.II.1993, n76 (16 Skulls & maxl, 31 ro, 9 lft & 8 rt maxl, 51 lft & 60 rt mand; T.a.), SMF
80678, C.E.; ibid., 19.III.1996, n100 (many fragments), SMF 84486-93, C.E. – Halabiye, 31.V.1989,
n47 (5 ro & mand, 11 ro, 7 lft & 5 rt maxl; 39 lft & 42 rt mand; T.a.), SMF 80609-11, D.K. – Qal’aat
ar’Rahba, 14.III.1979, n9 (T.a.), SMF 60378-86, R.K.; ibidem, 17.V.1989, n20 (6 skulls, , 4 lft & 2 rt
maxl, 20 lft & 15 rt mand; T.a.), SMF 80661, D.K. – Dura Europos (as’Salahiya), 17.V.1989, n69
(T.a.), SMF 80640-49, D.K. – Wadi al-Agig, E of Tell Sheikh Hamad, 29.X.1986, n1, SMF 69141, F.
KRUPP & W. SCHNEIDER. – Tell Sheikh Hamad, 28.X.1986, n1, SMF 69131, F. KRUPP & W. SCHNEI-
DER. – 2 km SE Tell Sheikh Hamad, 26.X.1986, n1, SMF 69130, F. KRUPP & W. SCHNEIDER. – Tell al
Budeiri, 23.V.1989, n404 (247 ro, 7 lft & 6 rt maxl; 385 lft & 404 rt mand; T.a.), SMF 80608, D.K.
Remarks: This widespread and very common species is found regularly in owl pellets.
Mus macedonicus Petrov & Ruzic, 1983
Material. Qater Maghara, 22.II.1993, n1 (left ro with malar process broad; T.a.), SMF 80677, C.E. –
Okersheih, 20.III.1996, n1, SMF 84456, C.E.
Taxonomy. For the Mediterranean parts of the Near East, AUFFRAY et al. (1990) designated
as Musspretoides” (a name not available according to the International Code of Zoological
Nomenclature) the short-tailed outdoor species of the genus Mus, differing by a zygomatic
index from M. musculus. The malar process at the upper base of the zygomatic plate is wide
in M. macedonicus, but narrower in M. musculus (see also ORSINI et al. 1983, MACHOLAN
Distribution. This species was identified from owl pellets from the Golan Heights by AUF-
FRAY et al. (1990) and from Aleppo by HARRISON & BATES (1991). The present material
extends the range eastwards into steppic regions along the Euphrates.
Rattus rattus (Linnaeus, 1758)
Material. Nahr el-Kabir, 5 km SW Kafariya, road Lattakia - Jisr ech-Choghur, 7.III.1980, 1 juv (rt
mand-fragment), SMF 60376, R.K. – Qal’aat al-Hosson, 25.VI.1998, n1 (skull), A.S. – al’Hamra,
27.VI.1998, n1 (1 skull, 1 maxl, 1 lft & 1 rt mand), A.S. – Shah`Ranar, 27.VI.1998, n1 (1 maxl, 1 lft &
1 rt mand), A.S.
Distribution. Surprisingly few records are available for R. rattus in Syria. RUSSELL (1794,
Mus rattus) lists it from Aleppo; B. AHARONI (1932) records it from Hama and Damascus; it
was found in owl pellets at Crak des Chevaliers (= Qal’aat al-Hosson) (NADACHOWSKI et al.
1990). The species was not found in the Tell Abiad and Ain Aarous regions (MISONNE
Sylvaemus (Sylvaemus) iconicus (Heptner, 1948)
Material. Nahr al-Kabir, 5km SW Kafariya, road Latakia - Jisr ech-Choguhr, 7.III.1980, n3 (3 lft & 3
rt mand), SMF 60377, R.K.
Taxonomy. The species is identified by the skull characters detailed in FILIPPUCCI et al.
(1996) and the specific name used here follows the results of the study by KRYŠTUFEK
(2002), who examined several type specimens relevant to the nomenclatrure of Sylvaemus
species in the Near East. Previous records of this species were either identified with Apode-
mus sylvaticus
(Linnaeus 1758) and its assumed subspecies (tauricus Barrett-Hamilton,
1900, preoccupied by tauricus Pallas, 1811, = iconicus Heptner, 1948 = kilikiae Kretzoi,
1964), or with A. flavicollis (Melchior, 1834) as A. f. argyropuloi Heptner, 1948. Recently
FILIPPUCCI et al. (1989), describing the small-sized S. hermonensis, stated that S. sylvaticus
is absent in Israel and most records (except for Mt. Hermon) refer to S. flavicollis. Without
re-examination of Syrian specimens, the only reliably recorded occurrence is Mt. Hermon
(FILIPPUCCI et al. 1989, FILIPPUCCI 1992) at 1700 to 2200 m. a.s.l. (HAIM et al. 1993).
Nesokia indica myosura (Wagner, 1845)
Material. Dura Europos (as’Salahiya), 17.V.1989, n2 juv (mand; T.a.), SMF 80637, D.K. – Qal’aat
ar’Rahba, 17.V.1989, n2 (1 ro & mand, 2 lft mand), SMF 80659-60, D.K. – Tell al Budeiri, 23.V.1989,
n4 (4 skulls, 3 lft & 3 rt mand; T.a.), SMF 80589-93, D.K. – Qater Maghara, 22.II.1993, n1 (1 lft & 1 rt
mand; T.a.), SMF 80676, C.E. – Halabiye, 31.V.1989, n1 (1 rt maxl, 1 lft mand; T.a.), SMF 80621,
D.K. – Qasret Mohammed Ali, 28.VI.1998, n1 (1 maxl), A.S. – Okersheih, 28.VI.1998, n3 (3 maxl, 2
lft & 1 rt mand), A.S.
Distribution. The few previous records by MISONNE (1957) from Sheikh Hassan, 60 km S
of Tell Abiad, and by KOCK & NADER (1983) from Tell Abu Hurera and at Qal’aat ar’Rahba
(owl pellets), are supplemented by the present material. The Syrian occurrences are confined
to river valleys (Euphrates, Khabur, Balikh). They represent an extension of this Oriental species from the Iraqi populations along the Euphrates. The Syrian population is disjunct from the nearest localities of N. i. bacheri Nehring, 1897, in the Jordan Rift Valley (see HARRISON & BATES 1991). Acknowledgements. Sincere thanks are due to Prof Dr R. KINZELBACH, University of Rostock, Germany, and
Mr. C. EBENAU, Essen, Germany, for depositing reference material in the Senckenberg Museum. Our appre-
ciation is extended to Dr. M. ADE, Museum für Naturkunde of the Humboldt-Universität zu Berlin, for infor-
mation on Syrian hamsters. We also extend our appreciation to Prof. Dr. FAWZI SAMARAH, Damascus Univer-
sity, and the Ministry of Agriculture for their encouragement and support.
This paper is Publication No. 163 of the Zentrale Bibliographie der Ergebnisse des interdisziplinären For-schungsprojekt-Schwerpunktes Tell Sheikh Hamad der Freien Universität Berlin. References
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Authors’ addresses: Dr. Adwan Shehab, General Commission for Scientific Agricultural Research
(GCSAR), Douma, P.O. Box 113, Damascus, Syria. – Prof. Ahmad Daoud, Department of Biology,
Faculty of Sciences, Tishreen University, P.O. Box 4933, Latakia, Syria. – Dr. Dieter Kock, For-
schungsinstitut Senckenberg, Senckenberganlage 25, 60325 Frankfurt a.M., Germany. – Prof. Dr.
Zuhair Amr, Department of Biology, Jordan University of Science and Technology. P.O. Box 3030,
Irbid, Jordan. – Email contact:


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