Original InvestigationSystematics of the subgenus of mouse opossums Marmosa (Micoureus) (Didelphimorphia, Didelphidae) with noteworthy records from Paraguay
Introduction
The subgenus Marmosa (Micoureus) Lesson, 1842 includes six species of long-tailed, black masked mouse-opossums: M. (M.) alstoni (J.A. Allen, 1990), M. (M.) constantiae (Thomas, 1904), M. (M.) demerarae (Thomas, 1905), M. (M.) paraguayana (Tate, 1931), M. (M.) phaea (Thomas, 1899), and M. (M.) regina (Thomas, 1898). Micoureus is widely distributed in forested areas of the Neotropics from northern Argentina to Belize (Gardner and Creighton, 2008, Voss and Jansa, 2009). Despite recent systematic assessments (Voss et al., 2009, Rossi, 2005, Rossi et al., 2010, Gutiérrez et al., 2010), most of the nominal forms of Marmosa (Micoureus) have not been revised since Tate (1933), and some currently accepted synonymies are in need of revision (Voss and Jansa 2009). As a logical consequence, distributional and ecological aspects of these species are poorly known, especially towards the southern part of the range of the genus (e.g., Flores et al. 2000). Similarly, no phylogenetic study has focused on the relationships among the species of Micoureus, hampering an understanding of the historical biogeography of the subgenus.
Two species of Marmosa (Micoureus) have been recorded in Paraguay, M. (M.) constantiae Thomas, 1904 and M. (M.) paraguayana Tate, 1931. The former is only known from Parque Nacional Cerro Corá, in eastern Paraguay (Voss et al. 2009), while the second has several records throughout the eastern forested regions of this country (Brown, 2004, Gardner, 2005, Gardner and Creighton, 2008, Krumbiegel, 1941). Herein, we report new and noteworthy locality records for Marmosa (Micoureus) in Paraguay, including the first records for the western Dry Chaco region. These records are identified to the species level (cf. Tate, 1931, Tate, 1933, Voss and Jansa, 2009, Dias et al., 2010, Gutiérrez et al., 2010) on the basis of morphological and molecular data (including the first sequence of a Paraguayan specimen of M. (M.) paraguayana) and by comparison with the original descriptions and photographs of the holotypes. Paraguayan sequences were analyzed in the context of a phylogenetic analysis that includes sequences of five of the six species currently recognized in the subgenus Micoureus. The analyzed matrix incorporates a total of 70 sequences of Micoureus. This constitute the most taxonomically and geographically dense phylogenetic analysis of Micoureus.
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Material and methods
The Paraguayan records of Micoureus reported here are based on specimens housed in the Museo Nacional de Historia Natural del Paraguay (MNHNP), San Lorenzo, Paraguay; Instituto de Bioecología y Investigación Subtropical (IBIS), Pilar, Ñeembucú, Paraguay; The Field Museum of Natural History (FMNH), Chicago, USA; Museum of Southwestern Biology (MSB), Alburquerque, NM, USA; and Natural Science Research Laboratory (TK) and Texas Tech University, Lubbock, TX, USA. External measurements were taken
Results
Levels of observed intraspecific variation range from 0 to 3.7% (alstoni and demerarae respectively (Table 1). Meanwhile, interspecific comparisons reach large values in all cases (Table 1); values range from 8.3 (constantiae vs. regina) to 13.2% (alstoni vs. constantiae). Maximum parsimony analysis recovered 640 shortest trees (789 steps; consistency index = 0.530; retention index = 0.874). The topology of the strict consensus of these trees is fully congruent with that found in the Bayesian
Discussion
The phylogenetic analysis shows that the two main lineages of Micoureus are present in Paraguay. One of this is represented by M. (M.) paraguayana that inhabits the forested areas of eastern Paraguay. Meanwhile, the other main lineage is represented by M. (M.) constantiae, which is distributed in open areas of both Chaco and the Cerrado in eastern Paraguay. This finding highlights the diversity of Paraguay not just at the species level but also phylogenetically. This is important given that the
Acknowledgements
We thank Sharon Jansa and Jim Patton whom generously provided unpublished data and Ronald Pine, Alfred Gardner, Shayna Harris, and an anonymous reviewer for comments and revisions of this manuscript. The assistance of Robert Baker and Heath Garder (TTU), Bruce Patterson (FMNH), Phil Myers (UMMZ), Joe Cook and Jon Dunnum (MSB) and the staff at the Museo Nacional de Historia Natural del Paraguay is greatly appreciated. We thank Tom Husband and the Conservation Genetics Laboratoy at University of
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