RESUMEN
Species boundaries are difficult to establish in groups with very similar morphology. As an alternative, it has been suggested to integrate multiple sources of data to clarify taxonomic problems in taxa where cryptic speciation processes have been reported. This is the case of the harvest mouse Reithrodontomys mexicanus, which has a problematic taxonomy history as it is considered a complex species. Here, we evaluate the cryptic diversity of R. mexicanus using an integrative taxonomy approach in order to detect candidate lineages at the species level. The molecular analysis used one mitochondrial (cytb) and two nuclear (Fgb-I7 and IRBP) genes. Species hypotheses were suggested based on three molecular delimitation methods (mPTP, bGMYC, and STACEY) and cytb genetic distance values. Skull and environmental space differences between the delimited species were also tested to complement the discrimination of candidate species. Based on the consensus across the delimitation methods and genetic distance values, four species were proposed, which were mostly supported by morphometric and ecological data: R. mexicanus clade I, R. mexicanus clade IIA, R. mexicanus clade IIIA, and R. mexicanus clade IIIB. In addition, the evolutionary relationships between the species that comprise the R. mexicanus group were discussed from a phylogenetic approach. Our findings present important taxonomic implications for Reithrodontomys, as the number of known species for this genus increases. Furthermore, we highlight the importance of the use of multiple sources of data in systematic studies to establish robust delimitations between species considered taxonomically complex.
RESUMEN
The Reithrodontomys tenuirostris species group is considered "the most specialized" within the genus Reithrodontomys from morphological and ecological perspectives. Previous studies based on molecular data recommended changes in the taxonomy of the group. In particular, R. microdon has been the most taxonomically questioned, with the suggestion that it constitutes a complex of cryptic species. We analyzed the phylogenetic relationships of the R. tenuirostris species group using DNA sequences from the mitochondrial Cytochrome b gene and Intron 7 of the nuclear beta fibrinogen gene. In addition, divergence times were estimated, and possible new taxa delimited with three widely used species delimitation methods. Finally, possible connectivity routes based on shared haplotypes were tested among the R. microdon populations. All species were recovered as monophyletic with the exception of R. microdon, whose individuals were grouped into four different haplogroups, one of which included specimens of R. bakeri. Diversification within the R. tenuirostris species group began about 3 Ma, in the Pleistocene. The bGMYC and STACEY delimitation methods were congruent with each other, delimiting at the species-level each haplogroup within R. microdon, while the mPTP suggested a greater number of species. Moreover, none of the haplogroups showed potential connectivity routes between them, evidencing lack of gene flow. Our results suggest the existence of a higher number of species in the R. tenuirostris group, because we show that there are four species within what is currently recognized as R. microdon.
Dentro del género Reithrodontomys, el grupo de especies R. tenuirostris es considerado "el más especializado" morfológica y ecológicamente. Estudios moleculares previos recomendaron cambios en su taxonomía, proponiendo a R. microdon como un complejo de especies crípticas. Se analizaron las relaciones filogenéticas del grupo de especies R. tenuirostris con base en información de un gen mitocondrial, Citocromo b, y uno nuclear, el intrón 7 del beta fibrinógeno. Se estimaron los tiempos de divergencia, y se delimitó a posibles nuevos taxa aplicando tres métodos comúnmente utilizados, y se evaluaron posibles rutas de conectividad con base en los diferentes haplotipos identificados en las poblaciones de R. microdon. Todas las especies del grupo se recuperaron como monofiléticas excepto R. microdon, cuyos individuos formaron cuatro haplogrupos diferentes, uno de los cuales incluyó a especímenes de R. bakeri. Según la datación obtenida, la diversificación del grupo R. tenuirostris comenzó en el Pleistoceno, hace aproximadamente 3 Ma. Los métodos de análisis de delimitación de especies bGMYC y STACEY resultaron congruentes entre sí, logrando delimitar cada haplogrupo dentro de R. microdon a nivel de especie, mientras que con el método de mPTP se delimitó un número mayor de especies. No se identificaron rutas de conectividad entre haplogrupos, lo que resultaría en una ausencia de flujo génico. Se sugiere la existencia de un mayor número de especies en el grupo R. tenuirostris, ya que hay cuatro especies incluidas en lo que actualmente se reconoce como R. microdon.