depths of feeding tunnels are 10 to 410 cm; tunnel diame-
ters are 5 to 12 cm; 10 to 200 mounds are formed per indi-
vidual system; and up to 3 tons of soil are displaced (Topa-
chevskii 1969; Nevo 1979; Savic ́ 1982; Heth 1989, 1991).
The superficial foraging burrows are connected to a deeper
tunnel system or an elevated complex breeding mound in-
volving nest, storage, and sanitary chambers (Nevo 1961,
1999; Savic ́ 1973, 1982; fig. 25.3c – e).
Geographical and Ecological DistributionThe Eurasian family Spalacidae lives in East Europe, West
Asia (including Asia Minor), the Near East, and North Af-
rica (fig. 25.1 in Nevo et al. 2001). The distribution of re-
cent and fossil Spalacidae is illustrated in figures 25.1–25.5
in Savic ́ and Nevo (1990), and the ecogeographical distri-
bution of recent Spalacidae is given in table 25.1 in Savic ́
and Nevo (1990). The Spalacidae live in altitude from be-
low sea level to an elevation of 2,600 m in the Balkans
(Savic ́ and Soldatovic ́ 1979) and Asia Minor (Nevo 1961).
Spalacids range in Mediterranean and steppic habitats, in-
habiting grassy and mountainous steppes, semideserts, and
desert steppes. These mole-rats may penetrate open biota in
agricultural plots. However, basically, they are steppic sub-
terranean rodents whose evolutionary history, speciation,
and adaptation are intimately linked to increasingly arid en-
vironments, primarily in the Near East (Nevo 1991, 1999;
Nevo et al. 2001), North Africa (Nevo et al. 1991, 1992,
1994, 1995; Nevo, Simson, Heth, Redi, and Filippucci
1991), and Asia Minor (Kivanc ́ 1988).
In Israel, the S. ehrenbergisuperspecies S. galili, S. go-
lani, S. carmeli,and S. judaei(2n52, 54, 58, and 60, re-
spectively, fig. 25.4) range from mesic to xeric regions, but
they are restricted by the 100 mm isohyet (Nevo 1961) and
rarely colonize true deserts (Nevo 1989; Nevo et al. 1997).
In Asia Minor, Spalaxextend from mesic habitats near the
Aegean Sea (2n38) and the Mediterranean Sea (2n56)
to xeric habitats in central Anatolia (2n60, 62; Nevo
et al. 1994, 1995; Sözen and Kivan, 1998). In Jordan,
Spalaxspeciated into four species, all 2n60, varying in
karyotype morphology, and across mountain ranges from
Gilead through Amon, Moav, and Edom to Ras-e-Naqb,
near Aqaba (Ivanitskaya and Nevo 1998; Nevo et al. 2000).
Spalacids occupy most soil types, including open primary
habitats of scrub in plains, hills, and mountains and in sec-
ondary cultivated fields where they may become serious
pests (Nevo 1961).Phylogeny and SystematicsThe phylogeny and systematics of the family have been
largely intractable. This is true from the familial to the spe-
cific level, and no consensus has been reached (reviewed by
Ognev 1947; Topachevskii 1969; Corbet 1978; Carleton
and Musser 1984; see Savic ́ and Nevo 1990). Molecular
phylogenetics based on single molecules (albumin, transfer-
ring; Sarich 1985), haemoglobin (Kleinschmidt et al. 1985),
crystalline (Hendriks et al. 1987), and ribonuclease (Jekel
et al. 1990) all suggest an early evolutionary divergence294 Chapter Twenty-Five
Figure 25.4 Geographic distribution of the four species belonging to the
Spalax ehrenbergisuperspecies in Israel, separated by narrow hybrid zones. The
abbreviations of the 18 populations and their ecogeographic nature appear in
parentheses: (C Central; M Marginal; NHZ Near Hybrid Zone; SI Semi-
Isolate and I Isolate), are as follows: Spalax galili(2n52) –1Ma’alot (NHZ);
2 Kerem Ben Zimra (C); and 3 Qiryat Shemona (M); Spalax golani(2n54)
- 4 Mt. Hermon (M); 5 Quneitra (C) and 6 El-Al (NHZ) Spalax carmeli
(2n58) – 7 Kabri (NHZ) 8 Zippori, (C); 8a Mt. Carmel, (C); and 9
Afiq, (M; NHZ); and Spalax judaei(2n60)-10 Anza, (NHZ); 11 Jerusalem
(M); 12 Lahav (C); 13 Sede Boqer (I); 14 Wadi Fara (SI); 15 Jiftlik (SI);
16 Dimona, (I); and 17Ramat Hovav (SI). (Compiled from Nevo et al. 1993).