Systematics and Phylogenetics ACTA ZOOLOGICA BULGARICA Research Article Acta zool. bulg., 66 (4), 2014: 461-468

Morphological Morphometric Characterisation of the Eastern Broad-toothed Field Mouse mystacinus (Rodentia: ) from Zagros Mountains, north-western Iran

Jamshid Darvish1,2, Zeinolabedin Mohammadi1*, Fatemeh Ghorbani1, Ehsan Mostafavi3,4

1Department of Biology, Faculty of Sciences, Ferdowsi University of Mashhad, Mashhad, Iran; E-mail: [email protected] 2Applied Institute, Rodentology Research Department, Ferdowsi University of Mashhad, Mashhad, Iran 3Department of Epidemiology, Pasteur Institute of Iran, Tehran, Iran 4Research Centre for Emerging and Reemerging infectious diseases, Pasteur Institute of Iran, Akanlu, Kabudar Ahang, Hamadan, Iran

Abstract: The eastern broad-toothed field mouse Apodemus mystacinus (Danford and Alston, 1877) has been re- ported from the Levant, some Aegean islands, Anatolia, the southern Caucasus and North Iraq. In this study, 15 specimens of A. mystacinus were recorded from the Zagros Mountains, north-western Iran. Four external features, 13 linear measurements of the skull and 14 dental characters were measured. Dental and cranial characters were evaluated based on the frequencies of 27 character states for 12 morphological characters. The oak forests of the north-western slopes of the Zagros Mountains are known as the eastern- most range boundary of A. mystacinus, and the species has been blocked by the north-south extension of the Zagros Mountains. Based on morphological and morphometric characters, we suggest that the popu- lation of A. mystacinus from the Zagros Mountains possibly belongs to a new subspecies but this issue requires further analyses.

Keywords: Apodemus mystacinus, Zagros Mountains, morphometrics, Iran, subspecies

Introduction The Eastern broad-toothed field mouse Apodemus the subgenus Sylvaemus was not accepted by some mystacinus (Danford and Alston, 1877) was original- authors (Ço l a k et al. 2007). Me z h z h e r i n (1997) ly described as Mus mystacinus from Zebil, Bulgar and Pa v l i n o v et al. (1995) assigned this species to Dagh, southern . This species is distributed the subgenus Karstomys as a member of the genus in the Eastern Mediterranean through Anatolia, Sylvaemus. the Levant and the northern part of Iraq. Its range El l e r m a n , Mo r r i s o n -Sc o t t (1951) revised the also extends westward to some Aegean islands and taxonomic status of the subspecies of A. mystacinus the easternmost boundary of its range is known to and accepted two valid subspecies: A. m. mystacinus be North Iraq. It occurs in the South Caucaususin and A. m. euxinus. Vo h r a l i k et al. (2002) considered (Co r b e t 1978, Mu s s e r , Ca r l e t o n 2005). the Asian populations of A. mystacinus as a homoge- A. mystacinus was accounted from morphological nous species based on colouration and morphometric point of view as the most primitive species of the ge- analyses, with no distinct subspecies. A. m. mystaci- nus Apodemus (El l e r m a n , Mo r r i s o n -Sc o t t 1951). nus and A. m. euxinus were validated by Co l a k et It was also recognised as the ancestral member of al. (2004). The related form occurring in the Balkan the subgenus Sylvaemus in the Western Palaearctic Peninsula A. epimelas Nehring, 1902 has been syn- based on molecular phylogeny (Ch e l o m i n a , At o p k i n onymised with A. mystacinus or recognised as its 2010). However, the inclusion of A. mystacinus in subspecies (El l e r m a n 1941, El l e r m a n , Mo r r i s o n -

*Corresponding author: [email protected]

461 Darvish J., Z. Mohammadi, F. Ghorbani, E. Mostafavi

Sc o t t 1966, Co r b e t 1978, Ni e t h a mm e r 1978, ring to Co r b e t (1978), Et e m a d (1978),Vo r o n t s o v Mu s s e r , Ca r l e t o n 1993). Currently, A. epimelas is et al. (1992), Co l a k et al. (2004) and Kr y š t u f e k , treated as a distinct species based on morphology Vo h r a l i k (2005). The standard voucher specimens and morphometrics (Me z h z h e r i n 1997) or molecu- (skins, skulls, tissue samples) were deposited in the lar studies (Fi l i p p u cc i et al. 2002, Be l l i n v i a 2004, Zoology Museum of the Ferdowsi University of Mi c h a u x et al. 2005). Mashhad (ZMFUM). The specimens were weighted, Until recently, four species of wood mice of the sexed and scored for four external dimensions; fur- genus Apodemus have been known from Iran, i.e. ther, 13 linear measurements on the skull were taken A. witherbyi (Thomas, 1902), A. uralensis (Pallas, using a vernier calliper accurate to the nearest 0.05 1811), A. hyrcanicus Vorontsov, Boyeskorov and mm (Table 1 and Appendix 1). 14 dental measure- Mezhzherin, 1992 and A. flavicollis (Melchior, 1834) ments were performed with a measuring microscope (Ja v i d k a r et al. 2005, Kr y s t u f e k , Hu t t e r e r 2006, to 0.001 mm (Table 2 and Appendix 1). Nomenclature Da r v i s h et al. 2010, Ka r a m i et al. 2008). In addi- of the dental characters followed Mi s s o n e (1969) tion, A. avicennicus Darvish, Javidkar & Siahsarvie, and Ja v i d k a r et al. (2007). The skulls were observed 2006 was described by Da r v i s h et al. (2006) from under a stereomicroscope (magnification 40x) and Iran. Fi r o u z (2008) mentioned A. mystacinus as a drawings were done using a camera lucida (Fig. 1). probable member of the fauna of Iran and considered its possible presence in the Azarbaijan provinces, Results north-western Iran. By the present study, the first records of A. Diagnostic characters mystacinus from the Zagros Mountains are added to Dorsal fur smoky grey, ventral fur pure white in chest the list of the of Iran. and white with grayish base near thighs. Pectoral spot lacking. Tail distinctly bi-coloured. Demarcation line Material and Methods of body present but not clearly visible from lateral view. This is the largest Apodemus species in Iran, In total, 15 specimens of A. mystacinus and 38 speci- with the longest tail (Mean=114.21±8.67) and head- mens of A. witherbyi were collected from Kordestan body length (Mean=97.00±7.95). Hind foot long, Province, western Iran, in June – July 2013, using mean=24.52±3.22 (longer than that in the sympat- live-traps and snack baits (Fig. 1). The specimens ric species A. witherbyi, mean=21.80±0.83) (Table were determined based on cranial and external 1). Skull large. Zygomatic arches weak. Brain-case morphological and morphometric features, refer- relatively large and rounded. Condylobasal length

Table 1. External and cranial dimensions (in mm) of A. mystacinus and A.witherbyi from north-western Iran

Variables A.mystacinus N=15 A.witherbyi N=38 Mean ± SE Min Max Mean ± SE Min Max HBL 97.00±7.95 90 118 90.40±1.18 77 103 TL 114.21±8.67 105 142 99.56±1.17 78 117 HFL 24.52±3.22 20 33 21.37±0.24 15 24 EL 17.58±1.41 16 20 15.79±0.08 11 26 CBL 26.75±0.29 24.82 29.26 23.31±0.10 21.24 24.72 FL 14.41±0.12 13.42 15.60 12.28±0.05 11.32 12.90 PAL 5.90±0.10 5.02 6.70 4.79±0.05 4.12 5.46 ZYGB 14.55±0.18 13.76 16.16 12.72±0.08 11.40 13.88 RW 4.69±0.050 4.34 5.24 4.43±0.04 3.66 4.84 IOC 4.75±0.050 4.26 5.08 4.33±0.02 4.10 4.66 BCW 13.27±0.13 12.06 14.40 11.70±0.04 11.23 12.32 IBW 12.16±0.11 11.16 12.88 10.98±0.50 10.44 12 RH 6.72±.09 6.32 7.46 5.94±0.04 5.40 6.38 BCBH 10.29±0.08 9.86 10.82 8.93±0.04 8.34 9.46 TBL 6.62±.08 6.10 7.16 5.91±0.05 5.32 6.82 TBW 4.99±0.03 4.68 5.22 4.40±0.02 4.08 4.76 ML 13.67±0.14 12.92 15.02 11.57±0.07 10.34 12.52

462 Morphological Morphometric Characterisation of the Eastern Broad-toothed Field Mouse Apodemus mystacinus...

Fig. 1. Sampling localities of A. mystacinus from the north western Iran Fig. 1. Sampling localities of A. mystacinus from the north western Iran. Table 2. Dental measurements of A. mystacinus and A.witherbyi from north-western Iran; measurements are in millimetres

Variables A.mystacinus N=15 A.witherbyi N=38 Mean ± SE Min Max Mean ± SE Min Max M1.L 2.28±.04 1.83 2.45 1.85±.01 1.69 1.96 M1.W 1.39±0.02 1.16 1.47 1.16±.01 1.02 1.26 M2.L 1.34±0.01 1.19 1.44 1.13±.01 1.03 1.20 M2.W 1.32±0.02 1.13 1.43 1.11±.01 1.01 1.22 M3.L 1.04±0.02 0.87 1.14 0.84±0.01 0.76 0.98

M3.W 1.07±0.02 0.79 1.21 0.81±.01 0.67 0.93 M.1L 2.14±0.02 1.90 2.22 1.66±0.01 1.17 1.78 M.1W 1.27±0.01 1.12 1.38 1.03±0.01 0.88 1.12 M.2L 1.40±0.01 1.24 1.54 1.14±0.01 1.04 1.25 M.2W 1.28±0.01 1.11 1.36 1.04±0.01 0.90 1.19 M.3L 1.20±0.02 0.98 1.31 0.97±.01 0.88 1.06 M.3W 1.12±0.10 0.84 1.33 0.89±0.04 0.80 1.0 MxTR 4.59±0.05 4.02 4.87 3.78±0.01 3.54 3.95 MnTR 4.73±0.05 4.09 4.93 3.76±.01 3.37 3.94 large, mean=26.75±0.29 mm (compared to that of en specimens (49.98%), U-shaped in three (21.42%) A. witherbyi, mean=23.31±0.10). Mean length of and with dens in four (28.56%) (Table 3). Posterior maxillary tooth row 4.59±0.05 mm (compared to margin of incisive foramina reaching line connect- 3.78±0.01 mm in A. witherbyi). ing anterior parts of upper tooth rows in 64.26%, All specimens of A. mystacinus except one while in 35.7% (n=5) same margin not reaching this have tubercle t12 on second upper molar (Table line (Table 3). 3). Number of cingula on first lower molar ranges Molars relatively broad. Upper molars with from 4 to 5, while on second lower molar 2, 3 and labial antrocone (t3) and lingual antrostyle (t1) to 4 cingula present (Table 3). Fronto-parietal sutures protocone (t5) on upper M1 not connected in 100% wide-V-shaped (49.98%), accolade-shaped (35.7%) of specimens. In lower molars, half of specimens or U-shaped (14.28%). Cusps t1 and t5 in all speci- show 4 and the other half 5 labial cingula on lower mens separated; interpterygoid sinus straight in sev- M.1. Number of labial cingula on lower M.2 is 2 in

463

Darvish J., Z. Mohammadi, F. Ghorbani, E. Mostafavi

Table 3. Variation in size, shape and position of the 12 parameters analysed in 15 specimens of A. mystacinus

464 Morphological Morphometric Characterisation of the Eastern Broad-toothed Field Mouse Apodemus mystacinus...

Table 3. Continued

465 Darvish J., Z. Mohammadi, F. Ghorbani, E. Mostafavi

Table 3. Continued

35.7%, 3 in 35.7% and 4 in 28.56% of specimens. The Eastern broad-toothed field mice inhabit Only two specimens have cingulum on third upper in the mountainous area or clay foothills with an al- molar (Table 3). titude of more than 1400 m in north-western Iran. In the sympatric species A. witherbyi, dorsal fur The highland oak forests of the Zagros Mountains brown, with red tint and ventral fur completely pure emerged gradually from 6700 to 5500 years ago white. Pectoral spot present in 100% of specimens. (Va n Ze i s t , Bo t t e m a 1977, Wa s y l i k o w a et al. Demarcation line of body distinct. Fronto-parietal 2008) and provided suitable ecological habitats for suture U-shaped (100%); posterior edge of pala- A. mystacinus in Iran. This species was found in the tine straight; first upper molar clearly stephanodont rocks and bushy foothills of the Zagros Mountains (cusps t1 and t5 connected by a ridge). The first in- where this mouse feeds on seeds and seedlings, nuts cisive foramen passing line connecting anterior part and acorns of the most abundant plant species of of tooth rows in 100% of specimens. Cusp t7 on the these forests, including Quercus infectoria, Quercus first upper M/1 is large. Tubercle t12 on the second brantii, Quercus libani, Prunus orientalis, Prunus upper molar not present. The number of cingula on scoparia, Crataegus azarolus, Pyrus glabra and first lower molar 3, while in the second lower molar Cerasus microcarpa (Ha i d a r i et al. 2013). In addi- is 2 (Table 3). tion, the Zagros forests are the easternmost bound- ary of the distributional range of A. mystacinus and Discussion the species does not penetrate the Central Plateau of Iran. Ve r e s h c h a g i n (1959) described the presence of In spite of some efforts undertaken so far for eluci- A. mystacinus euxinus in the Great and the Lesser dating the biodiversity of Iran (Da r v i s h et al. 2006, Caucasus and western Transcaucasia but it has never Sh a h a b i et al. 2011, Gh o r b a n i et al. 2010, Ka r a m i been reported from the Iranian part of the Caucasus. et al. 2008, Da r v i s h et al. 2012), there are still gaps Mi s s o n e (1959) mentioned that valleys are the only in the studies that address the species diversity in the corridors for the penetration of terrestrial species into country as a whole. The north-west Iran is a part of the Iranian Plateau. In this sense, the Shiller valley in the Irano-Anatolian hotspot, which is one of the most Kordestan, located on the boundary between North priorities for conservation in Asia (Mittermeier et al. Iraq and Iran may serve as such a corridor for east- 2012). The region was described as Armano-Kord area ward expansion of A. mystacinus because the species by Mi s s o n e (1959) and is characterised by high level was not collected in the other parts of the Zagros. of endemicity of its fauna and flora (Re c h i n g e r , La c k Our data confirmed the west-east decrease in the 1991, No r o o z i 2008, Mo h a mm a d i et al. in press). molar size comparing to that in Turkey’s specimens,

466 Morphological Morphometric Characterisation of the Eastern Broad-toothed Field Mouse Apodemus mystacinus... which was previously described by some authors 27.8 mm for A. m. smyrnensis and A. m. rhodius. (Sp i t z e n b e r g e r 1973, Vo h r a l i k et al. 2002). The However, this measurement are greater than those in specimens of A. mystacinus from the west of Iran the Transcaucasian specimens (Mean = 26.5 mm) as also demonstrate lower average values in all meas- reported by Sh i d l o v s k i i (1953). Therefore, the popu- urements comparing to the specimens from Turkey. lation of A. mystacinus from the Zagros Mountains For instance, the average value of the condyloba- might be recognised as a new subspecies after imple- sal length in the Iranian population of A. mystaci- menting some further studies. nus is smaller (Mean = 26.75 mm) in comparison with that from Turkey (Mean = more than 26.9 mm, Funding: We appreciate the financial support of the Center for Kr y š t u f e k , Vo h r a l i k 2005). Co l a k et al. (2004) re- Disease Control and Prevention of the Iranian Ministry of Health ported average values of the condylobasal lengths and Pasteur institute of Iran [Grants no. 582] for the sampling 26.8 mm for A. m. mystacinus and A. m. euxinus, of the study.

Appendix 1. Abbreviations

HBL Head and body length ML Mandible length TL Tail length M1.L First upper molar length HFL Hind foot length M1.W First upper molar width EL Ear length M2.L Second upper molar length CBL Condylobasal length M2.W Second upper molar width FL Facial length M3.L Third upper molar length PAL Palatal length M3.W Third upper molar width ZYGB Zygomatic breadth M.1L First lower molar length RW Rostrum width M.1W First lower molar width IOC Interorbital constriction M.2L Second lower molar length BCW Braincase width M.2W Second lower molar width IBW Interbulla width M.3L Third lower molar length RH Height of rostrum M.3W Third lower molar width BCBH Height of braincase with tympanic bulla MxTR Maxillary tooth row length TBL Tympanic bulla length MnTR Mandibular tooth row length TBW Tympanic bulla width

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