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Analyses of qualitative and quantitative variation in bacula and soft parts of the glans penis of 13 species of Ctenomys Blainville, 1926 from Argentina were used to suggest systematic and evolutionary relationships. The 13 species can be divided into spike-bearing {C. australis, C. azarae, C. porteousi, C. rionegrensis, and C. talarum) and spiny bulb-bearing species {C. dorbignyi, C. pearsoni, C. perrensi, C. roigi, Ctenomys sp. from Curuzú Laurel, Ctenomys sp. from M. F. Mantilla, and Ctenomys sp. from San Roque). Ctenomys yolandae is unique because it shows both spikes and spiny bulbs. In addition to spikes and spiny bulbs, some populations of C. pearsoni, C. talarum, and C. yolandae had a new structure, an inner sac of the intromittent sac. The most frequent pattern of occurrence of spikes or spiny bulbs was 1-1 (one at each side of the urethra), an ancestral character state in caviomorphs. Variation in bacular dimensions was limited and differences among species were small, with the exception of C. pearsoni. This species showed a significantly short baculum with a wide base. Sperm and penial morphology suggests that C. talarum is the most plesiomorphic and C. yolandae the most derived species of this group. The pattern of geographic variation among these 13 species rejects the hypothesis of penis morphology contributing to reproductive isolation. Ctenomys pearsoni is the only species with some evidence of reproductive isolation resulting from penis morphology.
Exceptional chromosomal variability makesCtenomys an excellent model for evolutionary cytogenetic analysis. Six species belonging to three evolutionary lineages were studied by means of restriction endonuclease and C-chromosome banding. The resulting banding patterns were used for comparative analysis of heterochromatin distribution on chromosomes. This combined analysis allowed intra- and inter-specific heterochromatin variability to be detected, groups of species belonging to different lineages to be characterized, and phylogenetic relationships hypothesized from other data to be supported. The “ancestral group”,Ctenomys pundti andC. talarum, share three types of heterochromatin, the most abundant of which was also found in C. aff.C. opimus, suggesting that the latter species also belongs to the “ancestral group”. Additionally, within the subspeciesC. t. talarum, putative chromosomal rearrangements distinguishing two of the three chromosomal races were identified. Two species belong to an “eastern lineage”,C. osvaldoreigi andC. rosendopascuali, and share only one type of heterochromatin homogeneously distributed across their karyotypes.C. latro, the only analyzed species from the “chacoan” lineage, showed three types of heterochromatin, one of them being that which characterizes the “eastern lineage”.C. aff.C. opimus, because of its low heterochromatin content, is the most primitive karyotype of the genus yet described. The heterochromatin variability showed by these species, reflecting the evolutionary divergence toward different heterochromatin types, may have diverged since the origin of the genus. Heterochromatin amplification is proposed as a trend withinCtenomys, occurring independently of chromosomal change in diploid numbers.
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