Characterization of the Brazilians species of Myotis Kaup, 1929 (Chiroptera: Vespertilionidae) and test on the phylogeography of Myotis nigricans (Schinz, 1821) and Myotis riparius Handley, 1960 / Caracterização das espécies brasileiras de Myotis Kaup, 1829 (Chiroptera: Vespertilionidae) e ensaio sobre filogeografia de Myotis nigricans (Schinz, 1821) e Myotis riparius Handley, 1960

Caroline Cotrim Aires

Myotis is a highly diverse genus, with 103 species recognized so far, in addition to another 94 subspecies whose taxonomical status has not yet been completely evaluated. Twelve species are known to South America, among which five are considered polytypic, including Myotis nigricans. This taxon in considered a species complex. In order to characterize, both morphologically and molecularly, the taxa of the genus Myotis, I examined 1,018 individuals from national and foreign scientific collections. Twenty-five characters from external and cranial morphology were analyzed and 13 body and 19 cranial measures were checked. In order to investigate inter and intraspecific variations and the existence of sexual dimorphism in each taxon analyzed, I submitted the cranial measures to uni and multivariate statistic tests. To characterize intra and interspecific genetic diversity of the taxa in the genus Myotis, I selected one mitochondrial gene, cytochrome b (402pb), and a nuclear gene, RAG2 (602pb). From these two genes, I obtained 78 sequences: 58 mitochondrial and 20 nuclear. From the data obtained, I recognized as valid eight Brazilian species in the genus Myotis (Myotis albescens, Myotis alter, Myotis levis, Myotis nigricans, Myotis riparius, Myotis ruber, Myotis simus, and Myotis sp.n.). I elevated Myotis osculati, a taxon hitherto considered a subspecies of Myotis nigricans, to specific level and diagnosed Myotis punensis and Myotis chiriquensis as valid species, which were previously considered synonyms of Myotis nigricans nigricans. In this context, I recognize 15 species for the genus Myotis in South America, representing an increase of 25% in the previously known diversity for the group in the continent. I have not found significant sexual dimorphism in the cranial measures for M. albescens, M. levis, M. nigricans, M. riparius and M. ruber. To better delimitate morphometricaly the taxa herein studied, specially M. nigricans from M. riparius, I proposed the use of two indexes: one cranial and one maxillary. Qualitative characters from skull, such as presence or absence of a sagittal crest and the arrangement of upper premolars, associated to those from external morphology, such as the arrangement of the hair along the uropatagium, the shape of external ear, and the angling from the distal portion of the lower maxillae, were fundamental for diagnosing the taxa occurring in Brazil. The phylogenetic relationship among the species of genus Myotis is still imprecise, even after the inclusion of those taxa diagnosed as valid in this thesis. Through Maximum Likelihood (ML), Neighbor-Join (NJ) and Bayesian analysis methods, two clades were defined, one formed by species with absent or shallow sagittal crest (clade A), and another (clade B) with the presence of sagittal crest. I correlate the polytomy found within clade A with a fast diversification of this group in the region, possibly with simultaneous events of speciation. I correlate the inclusion of M. levis in the clade of M. nigricans with an event of mitochondrial introgression, given the nature of this gene and the number of mutations, which are not compatible with the homoplasy for this gene. The estimated divergence time of 1.2 Ma (±0.8), coincides with climatical changes and forest retraction during the Pleistocene, which allowed the hybridization of these two lineages. I ran the Test of Mantel for the populations of M. nigricans and M. riparius from the Atlantic Forest dominium, aiming at verifying the existence of a correlation between genetic distance and geographical distance in the populations, testing the hypothesis of isolation by distance, which was rejected both for M. nigricans (p=0.9449; r=0.23) and for M. riparius (p=0.9997; r=0.60). The populations of Myotis riparius showed a low genetic divergence (0.8 to 1.5%) along their distribution, with the populations from Paraíba, São Paulo, Paraná and Paraguay sharing haplotypes. Regarding Myotis nigricans, I point out that populations from Bahia, São Paulo and Paraná share haplotypes and low divergence (1.9 to 2.4%) between the populations from the western portion of Serra do Mar. Yet, the level of divergence between such populations and those from Serra do Japi and Serra da Mantiqueira is high (5.7 to 8.1%), therefore similar to the interspecific levels for the genus. These data suggest a high gene flow from the North-South direction to eastern Serra do Mar, which may be acting as a barrier to interior populations. The understanding of the processes involved in the diversification of the group will be favored by the inclusion of samples of other populations distributed in the East-West portions of the Atlantic Forest.

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