The major initial mechanism of speciation in subterranean blind mole rats, Spalacidae, is chromosomal, primarily through Robertsonian rearrangements. Here we highlight another scenario of chromosomal rearrangement leading to ecological speciation and adaptive radiation apparently initiated by pericentric inversions and genic divergence to different ecologies in mole rats in Jordan. We analysed karyotype, allozyme, size and ecological diversity across the range of mole rats in Jordan from mesic Irbid in the north to xeric Wadi Musa (Petra region) in the south, a transect of 250km. We examined mole rats for chromosome (N= 71), size (N= 76), and allozyme (N=67) diversities, encoded by 32 loci, in 12 populations of the Spalax ehrenbep'superspecies in Jordan. By a combination of chromosome morphology, genetic distance, body size and ecogeography, we identified four new putative biological species. All species (except two animals in Madaba) share 2n=60 but vary in chromosome morphology, caused by pericentric inversions and/or centromeric shifts. The 'north Moav' species is karyotypically polymorphic for 2n (2n=60; including locally also two animals with 2n = 62). The distribution of the four species is associated with ecogeographical different domains and climatic diversity. Genetic diversity indices were low, but like chromosome arms (NFa) were positively correlated with aridity stress. Discriminant analysis correctly classified 91% of the individuals into the four species utilizing combinatonally chromosome, allozyme and size diversities. It is hypothesized that mole rat evolution underground is intimately associated with climatic diversity stress above ground.
Chromosome banding (G-, C-and Ag-NOR) analysis was carried out on 27 specimens of Sphalax ehrenbergi from seven localities and two specimens of S. leucodon from one locality, all from Turkey. No chromosomal variation was detected in S. ehrenbergi populations from Elazig, Siverek, Diyarbakir and Birecik having the same diploid numbers (2n = 52) and morphology of chromosomes (NFa = 72). The karyotypes of mole rats from Tarsus and Gaziantep possessed the identical diploid number (2n = 56) but different numbers of autosomal arms: NFa = 68 in the Tarsus and NFa = 78 in the Gaziantep populations. Chromosomes of S. leucodon from Malatya (2n = 60, NFa = 74) differed distinctly in the C-banding pattern from all S. ehrenbergi cytotypes by the almost entire absence of heterochromatin in acrocentric autosomes and the presence of heterochromatin arms in subtelocentric autosomes. Nucleolar organizing regions were found mainly on three pairs of chromosomes, but some differences in their localization were revealed. Comparison of G-banded chromosomes showed, that most chromosomes have a similar pattern. The types of chromosomal rearrangements were revealed due to the banding methods.
The major initial mechanism of speciation in subterranean blind mole rats, Spalacidae, is chromosomal, primarily through Robertsonian rearrangements. Here we highlight another scenario of chromosomal rearrangement leading to ecological speciation and adaptive radiation apparently initiated by pericentric inversions and genic divergence to different ecologies in mole rats in Jordan. We analysed karyotype, allozyme, size and ecological diversity across the range of mole rats in Jordan from mesic Irbid in the north to xeric Wadi Musa (Petra region) in the south, a transect of 250km. We examined mole rats for chromosome (N= 71), size (N= 76), and allozyme (N=67) diversities, encoded by 32 loci, in 12 populations of the Spalax ehrenbep'superspecies in Jordan. By a combination of chromosome morphology, genetic distance, body size and ecogeography, we identified four new putative biological species. All species (except two animals in Madaba) share 2n=60 but vary in chromosome morphology, caused by pericentric inversions and/or centromeric shifts. The 'north Moav' species is karyotypically polymorphic for 2n (2n=60; including locally also two animals with 2n = 62). The distribution of the four species is associated with ecogeographical different domains and climatic diversity. Genetic diversity indices were low, but like chromosome arms (NFa) were positively correlated with aridity stress. Discriminant analysis correctly classified 91% of the individuals into the four species utilizing combinatonally chromosome, allozyme and size diversities. It is hypothesized that mole rat evolution underground is intimately associated with climatic diversity stress above ground.
Comparative studies among seven populations of 2n = 60 S. leucodon employing classic cytogenetics (G- bands, C-bands, AgNOR-staining), fluorochrome staining, and fluorescence in situ hybridization of telomeric and rDNA probes are reported here for the first time. The studied specimens were assigned to two cytotypes: 2n = 60W and 2n = 60R. The basic karyotype of both cytotypes consisted of eight pairs of subtelocentric and 21 pairs of acrocentric autosomes, subtelocentric X and acrocentric Y chromosomes. Both cytotypes had variable numbers of B-chromosomes (1–3) and variable numbers of autosomal arms (NFa = 74–76) caused by amplification (deletion) of heterochromatin short arms in the second pair. The short arms of subtelocentric chromosomes were comprised of heterochromatin in both cytotypes. Nucleolar organizer regions (NORs) and rDNA clusters were detected at telomeric sites of the short arms in pairs Nos. 3, 5, 6, 9, and 13 in cytotype W, and in the short arms of pair No. 6, 8, 12, 13, and 16 in cytotype R. Different locations of rDNA clusters allowed unambiguous discrimination between two S. leucodon cytotypes possessing the same 2n = 60 and similar NFa (74–76) variability. Our findings suggest a high level of chromosomal divergence, which means that it is possible to consider these cytotypes as a well-differentiated, chromosomal lineage within the leucodon group.
Mole rats (genus Nannospalax) display prolific chromosomal variation with more than 50 distinct cytotypes. These cytotypes are largely indistinguishable morphologically, are mainly allopatric and their taxonomic ranking is contradictory. We established a cytochrome b phylogeny for 15 cytotypes belonging to all three species recognized on morphological grounds (morphospecies): N. leucodon, N. xanthodon and N. ehrenbergi. Phylogenetic reconstructions yielded two highly divergent groups which are in agreement with the current division into two subgenera (Nannospalax and Mesospalax). The former comprised samples from south-eastern Turkey, Israel and Egypt (the morphospecies N. ehrenbergi). Basal dichotomy within Mesospalax remained unresolved and the putative sister position of N. leucodon against the two lineages of N. xanthodon was not supported in our analysis. Net divergences between sister cytotypes were low (< 2.0%) and two N. leucodon cytotypes were not even reciprocally monophyletic. Among the three morphospecies, the genetic diversity was lowest in N. leucodon (2.4% ± 0.3%), highest in N. xanthodon (8.8% ± 0.7%) and intermediate in N. ehrenbergi (5.0% ± 0.5%). Our results show that associations between genetic and chromosomal variation are not widespread and common in mole rats, and therefore refute the generalization of a 'cytotype-equals-species' approach.
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