Nannospalax ehrenbergi (Nehring, 1898)

Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Spalacidae, Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona: Lynx Edicions, pp. 108-142 : 141-142

28.

Middle East Blind Mole-rat

Nannospalax ehrenbergi

French: Spalax d'Ehrenberg / German: Ehrenberg-Blindmaus / Spanish: Rata topo ciega de Oriente Medio

Other common names: Ehrenberg’s Mole-rat, Palestine Mole-rat

Taxonomy. Spalax ehrenbergi Nehring, 1898 ,

Jaffa (= Tel Aviv ), Israel.

Included in subgenus Nannospalax . Its distinct phylogenetic position relative to other forms in the genus Nannospalax has long been proposed on the basis of morphological characteristics and has since been strongly supported in molecular analyses. Nannospalax ehrenbergi represents a superspecies composed of complex allopatric, parapatric, and occasionally sympatric forms that are morphologically very similar but clearly represent distinct species. Strongest evidence that multiple good species are present in this complex comes from detailed analysis of N. ehrenbergi in Israel and Palestine by E. Nevo and colleagues in 2001 and in related studies. Four species are clearly present in Israel and Palestine, and these species were each given new names by Nevo and colleagues: golani (diploid number 2n = 54) from the Golan Heights, galili (2n = 52) from north-east of the Sea of Galilee, carmeli (2n = 58) from the Sea of Galilee southward to northern West Bank in the east and farther south along coast to Tel Aviv, and judaei (2n = 60) from Tel Aviv and northern West Bank southward. These four species differ not only in karyotype but also (often subtly) in body and cranial measurements, tooth and bacular morphology, ear ossicle shape, pelage color, habitat specialization, genetics, brain size, basal metabolic rate, urine concentrating ability, water turnover rate, relative kidney size, physiological response to cold, respiratory and circulatory physiology, mortality in captivity, timing of daily activity, exploratory behavior, and even swimming posture. Many of these differences appear to be adaptive. These four species appear to be maintained, even in narrow regions of overlap, by chromosomal incompatibilities, bacular morphology, and various behavioral barriers to reproduction involving olfaction, vocalizations, seismic communication, and differing levels of aggression. These species within the N. ehrenbergi superspecies have emerged as important model systems in evolutionary biology. A series of recent studies even suggests that sympatric speciation has taken place within galili , which suggests that these species warrant further division. Unfortunately, when Nevo and colleagues named golani , galili , carmeli , and judaei , they failed to account for the much older name ehrenbergi , noting only that its type locality, Tel Aviv, is found in the hybrid zone between carmeli and judaei . G. G. Musser and M. D. Carleton in 2005 attempted to resolve the problem of ehrenbergi superspecies taxonomy by adopting the four species of Nevo and colleagues and then applying the name ehrenbergi to populations immediately surrounding the type locality, all populations south of Israel and Palestine, and all populations north of Israel and Palestine. In contrast to their many other taxonomic recommendations, the unorthodox solution of Musser and Carleton in 2005 to the ehrenbergi taxonomy problem has not been widely adopted in subsequent mammal checklists, most of which reluctantly treat N. ehrenbergi as a single species. The name aegyptiacus by A. Nehring in 1898 is available for populations south of Israel and Palestine, and V. A. Topachevskii and others in 1969 recognized the North African forms as a distinct subspecies, aegyptiacus . B. Krystufek and V. Vohralik in 2009 noted that two names are available for populations north of Israel and Palestine, kirgisorum and intermedius , both named by Nehring in 1898. The name kirgisorum has page priority, but acquisition and provenance of type material are disputed. It was originally reported as being from Kazakhstan, well out of the known distribution of Nannospalax , but has subsequently been suggested as being from Syria. Clearly more than just one species exists to the east and north of Israel and Palestine, and future work will yield even more names. Recent molecular work further elucidates relationships among forms within the ehrenbergi superspecies: golani and galili appear to be sister taxa, carmeli and judaei form a clade with aegyptiacus , and Turkish and other northern populations are paraphyletic with respect to the four species from Israel and Palestine. The latter finding supports the hypothesis that N. ehrenbergi , and perhaps the genus Nannospalax , had its origin in Anatolia. Various sources of evidence clearly support existence of multiple species within the ehrenbergi superspecies, but unresolved taxonomic problems mean that it is still prudent to treat it as a single species. Monotypic.

Distribution. NE Libya (N Cyrenaica), N Egypt, Israel, W Jordan, Lebanon, Syria, SE Turkey, and N Iraq.

Descriptive notes. Head-body 130-220 mm, no visible externaltail; weight 73-252 g. Middle East Blind Mole-rats are small; males are slightly larger than females. In Israel and Palestine, individuals in cool mesic environments are larger, but the reverse appears to be true in Turkey. Egyptian individuals are smaller than those found farther north, and Libyan individuals are smaller still. Pelage color varies by soil color; it has slate-gray hairs, with dull buff, brown, or reddish tips. When worn, pelage looks uniform gray. Bristly keel of cream or yellow extends from triangular nose pad halfway to ear openings. Forefeet are brown; hindfeet are covered in gray hairs. Incisors have 2-3 grooves. Upper incisors are yellow to orange; lower incisors are pale yellow. Including cartilaginous base, baculum is 5-7-5 mm long and varies in shape and size by cytotype. There are four nipples: 0 pectoral + 2 pairs of inguinal. More than 30 chromosomal forms are known with diploid numbers of 2n = 52, 54, 56, 58, and 60.

Habitat. Usually coastal Mediterranean and interior steppe habitat but also forest clearings, sandy coastal plains, mountainous areas, and some rocky habitat from sea level to elevations up to 2200 m. Although the Middle East Blind Mole-rat is present where precipitation is very low,it is absent in true desert. It is absent from the Sinai Peninsula and Nile Delta area. North African populations are fragmented and located near the coast.

Food and Feeding. The Middle East Blind Mole-rat feeds primarily on tubers and bulbs of a variety of plants but also eats aboveground green plant parts. These plants are usually accessed from tunnels belowground, but occasionally individuals emerge to clip plants aboveground and carry them to burrows. Seeds and insects also have been recorded. Stored food in caches can be 25 kg.

Breeding. Female Middle East Blind Mole-rats breed once a year, although a second litter may occur in rare instances such as during a long rainy season or in close proximity to irrigation. Males and females can be rather aggressive at first interaction, but this aggression diminishes over time and copulation occurs. Gestation is 28-36 days. Females construct breeding mounds, with grass-lined nests with diameters of ¢.20 cm, located c¢.10 cm belowground. Peak births vary by region: early February at low elevations and early April at high elevations in Lebanon, early February and late March in Israel, and October-November in Egypt. Females give birth to 2-4 young that are 5 g and naked. Young disperse after c.2 months by digging tunnels adjacent to mothers’ burrows. At high densities, young may disperse aboveground. Females probably breed in their second year. Maximum life span in captivity is 20 years.

Activity patterns. Middle East Blind Mole-rats are active year-round and ¢.50% of the day. During the rainy season, they appear to be diurnal and polyphasic. During dry season, they are nocturnal and monophasic. The Middle East Blind Mole-rat is a chiseltooth digger. Although it can dislodge some material with head or forelimbs, almost all digging occurs with its lower incisors. Accumulated dirt is pushed to surface with head or back legs and ejected in mounds. More digging occurs in the rainy season when soil is easier to move. The Middle East Blind Mole-rat lives most ofits life underground, but it emerges and is active aboveground with some frequency in March-May and autumn. Aboveground activities include searching and foraging for food and dispersing.

Movements, Home range and Social organization. Burrows of Middle East Blind Mole-rats are 10-40 m long and 10-40 cm deep in winter and up to 1-5 m deep in summer. They contain a nest, toilets, and food caches. Typical mounds are 15-20 cm high, but breeding mounds can be 40 cm high, 160 cm long, and 135 cm wide. The Middle East Blind Mole-ratis solitary and aggressive toward conspecifics. Fights may lead to death of the loser. Densities in Israel and Palestine are 0-91-1-8 ind/ha. Individuals communicate seismically by head drumming and using frequencies of 50-100 Hz. Sounds are received by neighboring individuals pressing mandibles to tunnel walls. If an individual is removed,its territory is generally occupied by another within hours or days. Predators include the Marbled Polecat (Vormela peregusna), domestic cats, the little owl (Athene noctua), the common barn-owl (7yto alba ), the northern long-eared owl (Asio otus), the Eurasian eagle-owl (Bubo bubo), the Egyptian vulture (Neophron percnopterus), the black kite (Milvus migrans), Bonelli’s eagle (Aquila fasciata), the golden eagle (A. chrysaetos), the long-legged buzzard (Buteo rufinus), the hooded crow (Corvus corone cornix), and the white-breasted kingfisher (Halcyon SMYrnensis).

Status and Conservation. Classified as Data Deficient on The IUCN Red List (as Spalax ehrenbergi ). The Middle East Blind Mole-rat is considered vulnerable in Turkey. It is present in agricultural areas and is considered a pest in some regions. Intensive agriculture is a threat. Multiple individual species within this superspecies probably warrant conservation concern as distinct entities. Some forms are found in protected areas, but it is probable that species exist within this complex that are not protected.

Bibliography. Coskun et al. (2012), Ellerman & Morrison-Scott (1951), Hadid, Németh et al. (2012), Hadid, Tzur et al. (2013), Happold (2013d), Kankilic et al. (2013), Krystufek & Vohralik (2009), KryStufek, lvanitsakaya et al. (2012), Li Kexin et al. (2015), Lovy et al. (2015), Méhely (1909), Mendelssohn & Yom-Tov (1999), Musser & Carleton (2005), Nehring (1898), Németh et al. (2016), Nevo (1991, 2013), Nevo et al. (2001), Ranck (1968), Schlitter, Shenbrot et al. (2008), Shanas et al. (1995), Singaravelan et al. (2013), Topachevskii (1969).

Figures

1. Altai Zokor (Myospalax myospalax), 2. Steppe Zokor (Myospalax aspalax), 3. Armand’s Zokor (Mpyospalax armandii), 4. Manchurian Zokor (Myospalax psilurus), 5. Khingan Zokor (Myospalax epsilanus), 6. Fontanier’s Zokor (FEospalax fontaniern), 7. Gansu Zokor (Fospalax cansus), 8. Qinling Zokor (Fospalax rufescens), 9. Plateau Zokor (FEospalax bailey), 10. Rothschild’s Zokor (Eospalax rothschildi), 11. Smith’s Zokor (Eospalax smithu), 12. Chinese Bamboo Rat (Rhizomys sinensis), 13. Hoary Bamboo Rat (Rhizomys pruinosus), 14. Indomalayan Bamboo Rat (Rhizomys sumatrensis), 15. Lesser Bamboo Rat (Cannomys badius), 16. African Root Rat (Tachyoryctes splendens), 17. Giant Root Rat (Tachyoryctes macrocephalus), 18. Oltenia Blind Mole-rat (Spalax istricus), 19. Mehely’s Blind Mole-rat (Spalax antiquus), 20. Bukovina Blind Mole-rat (Spalax graecus), 21. Podolsk Blind Mole-rat (Spalax zemni), 22. Sandy Blind Mole-rat (Spalax arenarius), 23. Greater Blind Mole-rat (Spalax microphthalmus), 24. Giant Blind Mole-rat (Spalax giganteus), 25. Kazakhstan Blind Mole-rat (Spalax uralensis), 26. Lesser Blind Mole-rat (Nannospalax leucodon), 27. Anatolian Blind Mole-rat (Nannospalax xanthodon), 28. Middle East Blind Mole-rat (Nannospalax ehrenbergi)

Distribution. NE Libya (N Cyrenaica), N Egypt, Israel, W Jordan, Lebanon, Syria, SE Turkey, and N Iraq.