Sympatric speciation has been controversial since it was first proposed as a mode of speciation. Subterranean blind mole rats (Spalacidae) are considered to speciate allopatrically or peripatrically. Here, we report a possible incipient sympatric adaptive ecological speciation in Spalax galili (2n = 52). The study microsite (0.04 km 2 ) is sharply subdivided geologically, edaphically, and ecologically into abutting barrier-free ecologies divergent in rock, soil, and vegetation types. The Pleistocene Alma basalt abuts the Cretaceous Senonian Kerem Ben Zimra chalk. Only 28% of 112 plant species were shared between the soils. We examined mitochondrial DNA in the control region and ATP6 in 28 mole rats from basalt and in 14 from chalk habitats. We also sequenced the complete mtDNA (16,423 bp) of four animals, two from each soil type. Remarkably, the frequency of all major haplotype clusters (HC) was highly soil-biased. HCI and HCII are chalk biased. HC-III was abundant in basalt (36%) but absent in chalk; HC-IV was prevalent in basalt (46.5%) but was low (20%) in chalk. Up to 40% of the mtDNA diversity was edaphically dependent, suggesting constrained gene flow. We identified a homologous recombinant mtDNA in the basalt/chalk studied area. Phenotypically significant divergences differentiate the two populations, inhabiting different soils, in adaptive oxygen consumption and in the amount of outsidenest activity. This identification of a possible incipient sympatric adaptive ecological speciation caused by natural selection indirectly refutes the allopatric alternative. Sympatric ecological speciation may be more prevalent in nature because of abundant and sharply abutting divergent ecologies.adaptation | ecological stress | radio-tracking | metabolism | microscale T he origin and nature of species, the mystery of mysteries (1) and "the most important single event in Evolution" (2), have always been problematic in evolutionary studies (2-7). We adhere to the Biological Species Concept (2), recognizing its merits and demerits (2). The recent resurgence in speciation studies highlights many past obscurities (4), including sympatric speciation (7-18). Nevertheless, many basic questions related to adaptation and speciation, including sympatric speciation, still await resolution based primarily on the genomic sequence studies, such as in Drosophila (8), or even in species that presumably originated sympatrically, such as in the fly Rhagolites (9) or cichlid fishes in Africa and Neotropical America (10).The mode of species origin is still a major focus of heated debate. Does speciation occur primarily in allopatry, i.e., requiring complete geographic isolation, or can it occur in parapatry and peripatry, where limited gene exchange operates, or even in sympatry, where free gene exchange occurs, as suggested by Darwin (1)? Darwin envisaged allopatric, parapatric, and sympatric modes of speciation, but neither he nor his followers estimated their proportions in nature, which remain enigmatic and limited (4). Moreover, no spe...