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Turkish Journal of Zoology Turk J Zool (2016) 40: 925-932 http://journals.tubitak.gov.tr/zoology/ © TÜBİTAK Research Article doi:10.3906/zoo-1507-50

Distribution and current status of migratorius (Mammalia: ) in , with comments on its status in the Balkans

Nedko NEDYALKOV* National Museum of Natural History, , Bulgaria

Received: 01.08.2015 Accepted/Published Online: 05.01.2016 Final Version: 06.12.2016

Abstract: During the last decade I collected extensive new information about the distribution of the Grey (Cricetulus migratorius) in Bulgaria, mainly through the study of pellets of birds of prey and owls. New data about the dental morphology and morphometrics, distribution, and current status of the Grey hamster in Bulgaria are presented. The Grey hamster has been found in the diet of the Barn owl (Tyto alba), Little owl (Athene noctua), and (Aquila heliaca). It contributed a negligible part of the studied pellets: only 0.06% of individuals from 27,449 small . The presence of the was confirmed only in south- eastern Bulgaria, while in northern Bulgaria the species went extinct. The species’ distribution in the Balkans is discussed.

Key words: Grey hamster, morphology, birds of prey, owls, distribution

1. Introduction In Bulgaria the Grey hamster was first discovered The Grey hamster (Cricetulus migratorius Pallas, 1773) in 1959 (Peshev et al., 1960). Only single findings were is distributed from the Balkans to China and Mongolia reported afterwards, mainly from south-eastern Bulgaria, (Kryštufek et al., 2008). It can utilise various habitats reported in regional journals (in Bulgarian: Straka, 1962; – steppes, semideserts, and even the surroundings of Markov, 1964; Simeonov, 1964a, 1964b). Here I summarise human settlements – and can go up to 4300 m above sea the distribution and the species’ status in Bulgaria, and level (Gromov and Erbajeva, 1995). This is one of the present new data about the species’ morphology from the three hamster species that occur in Europe (Mitchell- western part of its range. Jones et al., 1999). For all of them significant changes and decreases in their ranges have been observed, mainly due 2. Materials and methods to agroenvironmental practices (e.g., changes in habitat This research is based on collected material from pellets utilisation, use of pesticides) or collection for the pelt and food remains from different birds of prey and owls (Cricetus cricetus) (Nechay, 2000). that I obtained between 2001 and 2014 from various parts During the Pleistocene, the Grey hamster was of Bulgaria. I identified and analysed a total of 27,449 widespread throughout Europe and reached Spain and individual small mammals found at roosts and nests Great Britain [sensu Kowalski (2001); the author joined from eastern and south-eastern Bulgaria of the following under this name besides Cricetulus migratorius a small species: Barn owl (Tyto alba), 20,157 small mammals; fossil form bursae, but see Hír (1993) and Tawny owl (Strix aluco), 472; Little owl (Athene noctua), Cuenca-Bescós (2003) for further comments about the 1861; Long-eared owl (Asio otus), 2709; Long-legged status of these forms]. In the beginning of the Holocene buzzard (Buteo rufinus), 643; and Eastern imperial eagle the species still inhabited Slovenia (Toškan and Kryštufek, (Aquila heliaca), 1607. Some preliminary data about new 2009). In spite of its extensive range, data about its findings of C. migratorius in Bulgaria have already been morphology and variation are scarce (Pradel, 1981; presented (Nedyalkov, 2013). Doğramacı, 1989; Hír, 1993).The cricetids are amongst For morphological comparison I used 6 complete the most poorly studied in the Palaearctic; thus, specimens of the Grey hamster collected from Central studying their morphology and the variation of the species Kazakhstan (46°23′30″N, 74°01′44″E) and 8 mandibles will provide key information about the group’s evolution found in the diet of Barn owl and Long-legged buzzard and its origin. from Central Anatolia, Turkey (37°19′14″N, 33°11′13″E). * Correspondence: [email protected] 925 NEDYALKOV / Turk J Zool

The measurements of teeth are made in accordance birds’ diet: 0.06% of the analysed material (27,449 small with the methodology described by Pradel (1981), while mammals). the terminology used follows Hír (1989). The teeth The Grey were identified mainly by their measurements were taken by stereomicroscope (accuracy: cranial and mandibular bones. The most complete remains 0.05 mm), and molar row – coronal length, with а digital were found in the remains of the Barn owl (Figure 1). One caliper (accuracy 0.01 mm). The abbreviations of the of the strongest bones, the mandible, remained entirely characteristics are as follows: M/m – upper and lower complete in the Barn owl pellets, in contrast to those from molar, L – greatest length, W – greatest width, MNI the Little owl and the Eastern imperial eagle, where it is – minimum number of individuals, identified by the fragmented in different degrees: broken proc. coronoideus, maximum count identified fragments from the left and proc. articularis, and proc. angularis (Figure 1). This is due right mandibula, the maxilla, or separate teeth. to differences in the feeding biology and habits of these species: the Barn owl swallows its prey (small mammals) 3. Results whole, while birds of prey (eagles, buzzards, falcons) 3.1. Data from the diet of the avian predators tear their prey to pieces (even small mammals) prior to From the analysed 27,449 individual small mammals from swallowing, as do some small owl species (such as Little pellets and prey remains, a minimum of 17 specimens of owl and Scops owl) for which insects are the main food the Grey hamster were found as part of the diet of three (Marti et al., 2007). avian species: Barn owl (Tyto alba), Little owl (Athene 3.2. Morphological description data of the teeth noctua), and Eastern imperial eagle (Aquila heliaca). It was Data from the dental measurements of the Grey hamster not found in the diets of the other 3 studied species. are presented in Table 2. The Grey hamster was represented in different М1. The anterocone is usually deeply split into two proportions in the diets of the avian predators (Table cusps. The anterior connection between the paracone and 1). Altogether, it constitutes a negligible part from the protocone is present or poorly developed, but sometimes

Table 1. Cricetulus migratorius localities and proportion in the diet of the owls and the Eastern imperial eagle.

Predator Location Collection day C. migratorius (MNI) All small mammals Tyto alba Matochina 20.8.2008 1 50 Tyto alba Levka 03.5.2006 14 532 Athene noctua Raykova mogila 10.8.2012 1 11 Aquila heliaca Sliven 15.3.2008 1 7

Figure 1. Degree of fragmentation of Cricetulus migratorius mandibles found in the diets of different avian predators; from left to right: A. heliaca, A. noctua, and T. alba.

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Table 2. Dental measurement (in mm) of Cricetulus migratorius from Bulgaria.

Meas. N Mean Min Max SD LM1 31 1.58 1.4 1.7 0.071 WM1 31 1.07 0.95 1.2 0.055 LM2 29 1.18 1 1.3 0.091 WM2 29 0.96 0.85 1.05 0.051 LM3 27 0.93 0.85 1 0.054 WM3 27 0.90 0.8 1 0.055 LM1-3 27 3.70 3.55 3.91 0.104 Lm1 24 1.53 1.4 1.65 0.072 Wm1 24 1.01 0.9 1.1 0.040 Lm2 25 1.25 1.1 1.35 0.048 Wm2 25 1.04 1 1.1 0.031 Lm3 22 1.07 1 1.2 0.047 Wm3 22 0.98 0.85 1.05 0.057 m1-3 20 3.85 3.45 4.02 0.140 it may be absent between the paracone. Usually there is Stara Planina Mountain (the main mountain ridge that a parastyle (morphotype C in Hír, 1993), but sometimes separates the country into north and south) (Straka, 1962). with preanterocone cingulum (morphotype A), or both of The species’ distribution is presented in two periods. them may be absent (morphotype D). These morphotypes The first one covers the time from the first finding of were observed in the following frequencies: morphotype the species in Bulgaria until 1985, when all information A, 20%; morphotype C, 40%; and morphotype D, 40% about the species’ distribution was summarised in the first (n = 20 teeth). Only an anterostyle was not observed edition of Red Data Book of Bulgaria (Christov, 1985). (morphotype B). The teeth are four-rooted. Between 1964 and 2003 there was no record of the M2. Anterior cingulum is well developed, especially on species in Bulgaria, but this was due to a lack of systematic the labial side. Mesolophe is absent. Four-rooted tooth. research in the south-eastern part of Bulgaria. Data about M3. The protocone is well developed; hypocone the species’ distribution after 2003 come from the study and metacone are strongly reduced. The anterolingual of diets of different owl species (Georgiev, 2004; Milchev, cingulum is less pronounced or may be absent. 2009). m1. The anteroconide is divided by a depression into My research confirmed the presence of the species in a lingual cusp and a labial cusp, which are of similar size. two old locations, around Sliven town (Simeonov, 1964a, Preanteroconide cingulum is absent. 1964b) and Levka village (Peshev et al., 1960); and one m2. All the four primary cusps are well developed. The more recent location, Rajkova mogila village (Georgiev, mesolophid is also absent in this element. 2004). m3. Mesolophid may or may not be present. There are 3.4. Habitat description a few teeth with little worn surface. The Grey hamster inhabits the lower and open part of All lower molars have two roots. south-eastern Bulgaria, where the climatic conditions are 3.3. Distribution in Bulgaria mild, with a Mediterranean influence penetrating through Based on this study and previous published data, the Grey the Maritsa valley (Gruev and Kuzmanov, 1999). hamster is found in 15 locations, mainly in south-eastern The vegetation composition is dominated primarily by Bulgaria (Markov, 1960; Peshev et al., 1960; Straka, 1962; two habitats. The open land is cover by “Semi-natural dry Markov, 1964; Simeonov, 1964a, 1964b; Georgiev, 2004; grasslands and scrubland facies on calcareous substrates Milchev, 2009; Figure 2). Only one location is north of (Festuco-Brometalia)” (HD 92/43: 6210). The xerothermic

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Figure 2. Distribution of Cricetulus migratorius in Bulgaria (1- Straka, 1962; 2- Simeonov, 1964a, 1964b; this study; 3–6- Milchev, 2009; 7–8- Markov, 1964; 9- Markov, 1960; 10- Peshev et al., 1960; 11- Peshev et al., 1960; this study; 12- Georgiev, 2004; this study; 13–14- Georgiev, 2004; 15- this study). meadows and pastures are covered by perennial grasses 4. Discussion such as Chrysopogon gryllus, Bothriochloa ischaemum, 4.1. Morphological data and Festuca valesiaca, with some semishrubs and shrubs, The teeth of Bulgarian Grey hamsters are close or almost mainly Paliurus spina-christi. Some authors determine this identical to these from Greece, Turkey, and Syria in habitat as secondary steppe (Bondev, 1991). The second size and morphology, but hamsters from Romania and type is “Pannonian-Balkanic turkey oak-sessile oak forests” Moldavia are bigger than Bulgarian ones (Table 3). (HD 92/43: 91M0), where there are xerothermic and The earliest records of the species in Bulgaria come mesothermic oak forests with diverse floristic structure, from Early Pleistocene, from Cave 15 (Popov, 1994). In usually dominated by Quercus frainetto (Bondev, 1991). the Holocene and Late Pleistocene this species has been The Grey hamster was found to inhabit areas from 40 found in all fossil assemblages in Bulgaria (Peshev et al., m up to 400 m a.s.l. (median: 200 m a.s.l.) in Bulgaria. 2004). The fossil hamsters were slightly larger than recent Characteristic accompanying small fauna found ones; this phenomenon is peculiar compared to other in the diets of different owl and birds of prey species is Pleistocene mammals (Kurtén, 1968). dominated by species inhabiting open habitats such The earliest records of the species come from Anatolia as the White-toothed shrew (Crocidura sp.), Grey (Turkey): Early Pliocene from İğdeli (Alpaslan et al., 2010) (Microtus arvalis/levis), and Macedonian house and Early Biharian (Early Pleistocene) from Bıçakçı (van (Mus macedonicus). In the diet of the Eastern imperial den Hoek Ostende et al., 2015). The first records from eagle I found some steppe elements such as the European Europe are from the Early Pleistocene from Tourkoubounia souslik (Spermophilus citellus) and the Mole (Spalax 2 (Greece) (van der Meulen and , 2001) and Cave leucodon). 15 (Bulgaria) (Popov, 1994).

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Table 3. Length and width measurements and the number of molars per specimen of Cricetulus migratorius from some recent and fossil populations. EB – Early Biharian, LV – Late Villanyian.

Lm1 Wm1 Locality Age References Mean Min Max N Mean Min Max

Kazakhstan (Melkosopochnik) Recent This study 1.56 1.5 1.7 12 1.00 0.95 1.05

Ukraine (Nikolaecksa obl.) Recent Topachevsky and Skorik, 1992 1.58 1.5 1.7 22 1.00 0.9 1.1

Moldavia Recent Popov, 2000 1.62 1.5 1.75 108 1.03 0.95 1.2

Romania Recent Hamar, 1962 1.65 1.5 1.7 11 0.99 0.9 1.1

Bulgaria Recent This study 1.53 1.4 1.65 24 1.01 0.9 1.1

Greece Recent Niethammer, 1982 1.58 1.55 1.7 16

Turkey (Central Anatolia) Recent This study 1.53 1.5 1.6 8 1.02 1 1.05

Iran Recent Darvish et al., 2014 1.5 4 0.97

Syria Recent Pradel, 1981 1.54 1.45 1.65 150 0.95 0.85 1.03

Turkey (Toros Mountains) Holocene Hír, 1993 1.52 1.33 1.62 48 0.97 0.88 1.04

Slovenia Holocene Toškan and Kryštufek, 2006 1.81 1.7 1.9 4 1.13 1.1 1.15

Bulgaria () Late Pleistocene Pradel, 1989 1.64 1.55 1.73 20 1.03 0.96 1.11

Bulgaria (Temnata - Prododnata cave) Late Pleistocene Popov, 1994 1.64 1.6 1.75 13 1.06 0.95 1.17

Bulgaria (Temnata cave, Cave 16) Late Pleistocene Popov, 2000 1.67 1.57 1.82 19 1.04 0.9 1.12

Greece (Arnissa) Late Pleistocene Mayhew, 1978 1.64 1.55 1.68 6 1.05 1 1.09

Serbia (Baranica cave) Late Pleistocene Bogićević et al., 2011 1.81 1.7 1.9 7 1.16 1.1 1.2

Bulgaria (Morovitsa cave) Middle Pleistocene Popov, 1989 1.62 1.47 1.71 6 1.03 0.99 1.12

Turkey (Emirkaya 2) Middle Pleistocene Montuire et al., 1994 1.59 1.53 1.68 4 1.00 0.9 1.08

Greece (Tourkoubounia 2) Early Pleistocene (EB) van der Meulen, Doukas, 2001 1.86 1.73 1.98 15 1.14 0.99 1.21

Turkey (Bıçakçı) Early Pleistocene (LV) van den Hoek Ostende et al., 2015 1.52 1 1

Turkey (İğdeli) Early Pliocene Alpaslan et al., 2010 1.71 1.65 1.76 4 1.02 1 1.05

Rossolimo (1979) studied the geographical variation and Long-eared owl from this part of country have been of the Grey hamster based on extensive material (22 checked, but the species was not found, suggesting that it collections and 230 specimens); she found that temperature probably went extinct there. is the main factor that drives its variation. In this species In Dobrogea (where the northern locality is found), there is no clear direction of variation (cline), but rather a there is intensive agriculture with all accompanying “mosaic-cline type of variation” (sensu Rossolimo, 1979). practices (deep ploughing, heavy use of rodenticides, and According to Gromov and Erbajeva (1995), mountainous autumn burning of arable and abandoned fields), which forms are bigger than the forms from lower altitudes. negatively impacts the population of the Grey hamster There is great individual variation in coloration within and the other two hamster species, newtoni the species, even within the same location (e.g., Kabul, and Cricetus cricetus, living in this part of Bulgaria (N. Afghanistan) (Niethammer, 1982). Maybe because of Nedyalkov, unpublished data). In south-eastern Bulgaria the socioeconomic these patterns of variation more than 20 forms were conditions are different; mainly there is less anthropogenic described (Ellerman and Morrison-Scott, 1966; Gromov pressure. This is due to the depopulation of the region, and Erbajeva, 1995). which has been happening since the 1990s, mainly as a 4.2. Distribution of Cricetulus migratorius in Bulgaria, result of demographic tendencies such as migration of with comments on its status in the Balkans people to big cities, aging of population, etc. This region I confirmed the presence of the Grey hamster only in is amongst the least populated in Bulgaria (32–40 people/ south-eastern Bulgaria. In northern Bulgaria there was km2; data from 2013 for the Yambol region, http://www. only one location, which has not been confirmed for more nsi.bg/, National Statistical Institute). Locally, agriculture than 50 years (Straka, 1962). Extensive materials (14,227 is carried out on small and fragmented arable patches, small mammals) from the diets of Barn owl, Little owl, surrounded by abandoned fields.

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There are no new data about the habitat preference of Greece. А revision of the species’ conservation status in the Grey hamster in Bulgaria. Studies that utilised traps the IUCN Red List is needed. revealed that the species is caught predominantly in cereal The localities of my recent findings of the Grey fields, hedges, and orchards (Peshev et al., 1960; Straka, hamster are geographically close to areas in Greece that 1962; Markov, 1964), with an unusual record from a lack records for the species at the moment; especially given forest clearing (Markov, 1960). Multiple authors reported that the species is widespread in the Turkish part of Thrace that the Grey hamster enters houses and grain storage (Kryštufek and Vohralík, 2009), I presume that the lack facilities, including reports from Armenia, Turkey, and of records in this region of Greece is due to insufficient Iran (Sosnikhina, 1950; Kryštufek and Vohralík, 2009 search efforts. and references their). According to Sosnikhina (1950), In conclusion, it can be presumed that, in the western the population of the Grey hamster in natural habitats part of its range (Romania, Bulgaria, and Greece), the in Armenia is very low, in contrast to human settlements species is rare and in low numbers. In terms of further where it was a dominant small mammal. research, additional efforts for determining the species’ In Romania the species occurred in the eastern part on distribution and status in the Balkans are needed. Molecular the border with Moldova (Hamar, 1962); the most recent genetic methods should be used considering the great species distribution was summarised by Murariu (2000). morphological variation of the species, especially in order There are no recent findings on the species (since 1973), to reassess the species’ taxonomical status, which currently but this is likely because of the lack of investigation in the postulates two subspecies, C. m. atticus (for Greece) and C. region (G. Chisamera, personal communication). m. vernula (for Bulgaria) (Niethammer, 1982). In Greece the Grey hamster is reported from the Peloponnese peninsula (Kahmann, 1964; Ondrias, Acknowledgments 1966, Niethammer, 1974). Recently the species has been I thank D Demerdzhiev for providing me with the reported from two locations there (Hymettus and Avlona) materials from the Imperial eagle and help with field (Alivazatos et al., 2005), and from several localities from work, and D Georgiev for the prey items from the Barn Thessaly (Bontzorlos, 2009), all data coming from the diet owl. I am grateful to Y Marinov for data on the vegetation of the Barn owl (Tyto alba). The Grey hamster was present in the region. I thank Hír János, Gabriel Chisamera, and in low numbers in the diet of Barn owl from Greece: 0.56% Ebru Diker for aid with finding published literature. The (from 29,061 prey items) from Thessaly (Bontzorlos, 2009), greatest thanks go to the anonymous reviewers, who 1.1% (from 152 prey items) from Hymettus, and 1.4% provided helpful comments that significantly improved the (from 94 prey items) from Avlona. According to Kryštufek manuscript. Gratitude is also extended to Yurii Kornilev, et al. (2008), the Grey hamster is probably extinct from who significantly improved the language of the paper.

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