Chromosomal polymorphisms in subterranean mole rats: origins and evolutionary significance

Two distinct chromosomal polymorphisms, among others, characterize the short (p) and long (qh) arms of chromosome number 1 of the Spalax ehrenbergi complex in Israel. We have studied the geographic distributions of these two polymorphisms in 60 animals belonging to four chromosomal species (2 n = 52, 54, 58 and 60). These comprised 15 populations: 12 from continuously distributed populations, two from semi-isolates, and one isolate in the northern Negev Desert.
Our results indicate that: (i) the two polymorphisms are widespread not only within populations and species but most strikingly between different bone marrow cells within an individual; this may reflect a diversity generating mechanism; (ii) the two polymorphisms of p and qh, primarily of p, are correlated with climatic factors of water availability and temperature, as well as geographical distances; these correlations are significantly above those expected by chance, (iii) The semi-isolates and the isolate display high levels of polymorphism in both p and qh, but particularly in qh, despite the small size population effects.
We suggest that at least the polymorphism in p is involved in the adaptive radiation of mole rats into diverse climatic regimes. Furthermore, we speculate that the forces generating chromosomal polymorphisms relevant to speciation (i.e. Robertsonian mutations), coupled with the evolutionary forces operating in small peripherally isolated populations, may be appropriate for the origin of new chromosomal species through peripatric speciation.     

Odour-genes eovariance and differential investigation of individual odours in the Mus species complex

Previous research using habituation techniques has demonstrated that greater genetic similarity between two individuals is associated with more similarity in the qualities of their individual odours ('odour-genes eovariance'). We assessed odour similarities across species in two pairs of genetically close species within the Mus species complex (M. musculus and M. domesticus; M. spicilegus and M. macedonicus). Subjects treated odours within each species pair as similar compared with an odour from the other species pair. Subjects also treated odours of M. spicilegus males from the same population as similar compared with the odour of M. spicilegus males from a different population. This confirms odour-genes eovariance across species and within populations and also supports previous findings that odour similarities are more salient than specific odour markers. When adult males were presented with odours of females from two different heterospecific species, subjects spent more time investigating the odour from his own species pair than the other species pair, indicating greater interest in the odour of the closer heterospecific and demonstrating that odour-genes eovariance is reflected in behavioural responses to odours. Implications of odour-genes eovariance as a basis for identifying degrees of genetic relatedness of unfamiliar individuals through similarities in individual odours are discussed.     

Genetic relatedness assessment through individual odour similarities in mice

Previous research has demonstrated that closer genetic relatedness between individuals (from kin to across species) is associated with greater similarity in the qualities of their individual odours ('odour-genes covariance'). This predictable relationship between individual genotypes and individual odours could enable animals to assess their degree of genetic relatedness to other individuals by comparing the degree of similarity between another individual's odour and their own odour. In two-choice tests with odours of unfamiliar mice from different populations and species, subjects from two species of wild mice, Mus spicilegus and M. musculus , that had been raised in mixed litters of both species spent significantly more time investigating the ano-genital odour of the more genetically similar individual. This differential interest was not affected by common rearing with heterospecifics. These responses are consistent with a self-referencing mechanism enabling differential responses across a wide spectrum of genetic relatedness from kin through populations to heterospecifics. These assessments depend on the degree of similarity between the donor's and the subject's odours rather than on differences between them. Such a parsimonious mechanism may provide a basis for differential responses to conspecifics as opposed to heterospecifics that may function as a premating isolating mechanism.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78, 595–603.     

Odour–genes covariance within a natural population of subterranean Spalax galili blind mole rats

Previous research, using habituation techniques with multiple rodent species as subjects, has demonstrated (from kin to across species) the greater perceptual similarity in the qualities of individual odours of more closely genetically related individuals ('odour–genes covariance', abbreviated 'OGC'). This predictable relationship between individual genotypes and individual odours has only been assessed in rodents in which the genetic similarities and dissimilarities are known either through laboratory breeding or by selecting odour donors from different populations or species. To study OGC within a natural population of rodents, we genotyped Spalax galili blind mole rats using ten microsatellite markers, and conducted pairwise genetic distance comparisons to estimate genetic similarity and to calculate the relatedness coefficient between pairs of individuals. We then used habituation-generalization techniques to assess whether perceived odour similarities covaried with the genetic similarities we had identified. Indeed, animals treated odours from genetically closer donors as more similar than odours from less genetically similar donors. The results suggest an accurate and subtle ability to resolve genetically determined odour distinctions among familially unrelated animals within a population. Graded differential responses to conspecific odours based on degrees of similarity between other individuals' odours and one's own have been demonstrated in kinship and species discriminations and preferences. The evidence presented here provides the basis for hypothesizing a similar process that could promote optimal outbreeding and inclusive fitness within populations of conspecifics. The evolutionary implications of these findings are discussed.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 483–490.     

Male–female associations and female olfactory neurogenesis with pair bonding in Mus spicilegus

Mound‐building mice Mus spicilegus exhibit life‐history traits that are unique among the Mus species complex, such as the cooperative mound‐building behaviour that gives the species its common name. In this and other socially coordinated activities, such as those associated with reproduction, these mice should be able to recognize individuals (via discrimination based on kinship, population and species) to mediate their interactions. Our previous studies have provided evidence of population and species recognition in M. spicilegus. The aims of the present study were: (i) to study associations of mice during their reproductive period (in outdoor enclosures), (ii) to investigate whether there is an influence of relatedness of females in these associations, (iii) to determine whether female M. spicilegus are able to make kin vs. non‐kin discriminations, and (iv) to study certain neurobiological correlates of male–female bonds. Stable male–female associations were found in almost all the experimental groups, both those with unrelated and unfamiliar females and those with unfamiliar sisters. Kinship between females did not affect female associations in our enclosure experiment, but in our kin discrimination experiment females did distinguish between unfamiliar sisters and unfamiliar unrelated females. Shared kinship may not encourage cooperative rearing of pups but could enhance cooperation in building mounds where mice over‐winter. Male–female associations could be based on a social bond, as hypothesized from previous laboratory experiments. This was substantiated in this study by increased olfactory bulbar neurogenesis in females that preferred (in two choice tests) their sexual partner 3 weeks after their first mating. Based on these results there is clear evidence to suggest that the mating system of M. spicilegus is social monogamy. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 323–334.     

Karyotype and genetic evolution in speciation of subterranean mole rats of the genus Spalax in Turkey

Karyotype (2fl) and allozymc diversity at 37 gene loci were determined in 69 subterranean mole rats in Turkey belonging to the two superspecies: the ancestor Spalax leucodon ( n = 55; 20 populations) and the descendant S. ehrenbergi ( n = 14; lour populations). We identified remarkable variation of diploid chromosome numbers in the S. leucodon superspecies: 2 n = 38. 40. 50, 54, 60 and 62; and in the S. ehrenbergi superspecies: 2 n = 52, 56 and 58. Genetic diversity indices were low on average in both S. leucodon and S. ehrenbergi superspecies: Allele diversity, A = 1.081 and 1.074; polymorphism, P-5%= 0.077 and 0.068; heterozygosity, H = 0.038 and 0.027; and gene diversity, H, = 0.038 and 0.034, respectively. H ranged from 0 in mesic or semimesic regions to 0.088 in arid Anatolia. We consider the populations with different diploid chromosome numbers, 2 n , as good biological species. Karyotypic diversity may mark extensive ecological speciation. Nei's genetic distances, D (average 0.174, range 0.002 0.422) and ecogeographical criteria suggest that almost each population may represent a different biological species, but critical future testing is necessary to support this claim. Karyotypes and allozymes are nonrandomly distributed across Turkey, displaying remarkable correlations with climatic and biolir factors. Both In and H are significantly correlated with aridity stress (2 n /rainfall, r , =–0.74; P < 0.001), and in our region also with climatic unpredictability. These results support the niche-width genetic variation hypothesis in space and time. Climatic selection in Turkey appears to be a major architect of karyotype and genetic (allozymc) diversity and divergence in mole rat evolution, in both speciation and adaptation.     

Burrow patterns of the mole rat Spalax ehrenbergi in two soil types (terra-rossa and rendzina) in Mount Carmel, Israel

Complete burrow systems of the mole rat Spalax ehrenbergi (Rodentia; Spalacidae), from two sites on Mount Carmel in Israel, with different soil types (terra-rossa and rendzina), were excavated and described here for the first time.
A comparison was made of burrow structure patterns in the two soils and of the sexes, with special attention to the features of the feeding tunnels.
The pattern in the rendzina revealed longer burrows with a longer main tunnel and fewer branches per metre of the main tunnel, while the pattern in the terra-rossa revealed shorter burrows with a shorter main tunnel and a relatively higher number of branches. These differences can primarily be related to the different levels of food availability, which is higher in the terra-rossa. It is suggested that each of the patterns reflects the mole rat's ability to optimize foraging efficiency in accordance with its given food availability.
The average total length of the males' burrows was much greater than those of the females' burrows in the rendzina soil. It appears that food requirements determine different burrow features of the sexes rather than reproduction requirements.
Other tunnel features (e.g. structural complexity, depth and width, segment length and spatial arrangement) and the factors which may affect them, as well as burrow structure of young mole rats and evidence of the underground dispersion of young mole rats, were presented and discussed.
Many similarities were found in a comparison of S. ehrenbergi burrow features with those of other solitary subterranean rodents.