phylogenetic relationships and classification of didelphid marsupials ...

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100 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322 DISTRIBUTION: Currently known from fewer than a dozen specimens, Hyladelphys has been reported from just nine localities in the rainforested lowlands of eastern Peru, central Amazonian Brazil, southern Guyana, and French Guiana (Astúa, 2007). Extrapolating from these scanty data is obviously problematic, but it would not be surprising to find this elusive taxon anywhere in Amazonia. REMARKS: High levels of molecular divergence between sequenced specimens of Hyladelphys from French Guiana and Peru, together with geographic variation in morphological characters, suggest that additional species remain to be described in this genus (Jansa and Voss, 2005). Subfamily Didelphinae Gray, 1821 CONTENTS: Marmosini, Metachirini, Didelphini, and Thylamyini. DIAGNOSIS: Members of this clade uniquely differ from other didelphids by lacking an anterior cutting edge on P3 (the third upper premolar is double bladed in all basal didelphids). In addition, Didelphinae differs from Glironiinae by having a caudal dorsum that is macroscopically naked, at least near the tip; an unpaired median teat; welldeveloped posterolateral palatal corners; and postorbital processes that (when present) are formed only by the frontals. Didelphinae additionally differs from Caluromyinae by having a shorter fourth manual digit (dIII is subequal to or longer than dIV), an incompletely ossified palate (maxillopalatine and sometimes other fenestrae are invariably present), transverse canal foramina, a wide gap between the alisphenoid tympanic process and the rostral tympanic process of the petrosal, an acute and strongly inflected mandibular angle, an upper canine alveolus that lies within the premaxillary-maxillary suture, a nonvestigial P1, a larger P3 (subequal to or taller than P2), and a distinct ectoflexus on M3. Didelphinae additionally differs from Hyladelphinae by having an unpaired median teat, nonvestigial milk teeth, and a larger P3 (subequal to or taller than P2). REMARKS: The monophyly of Didelphinae is strongly supported by all analyses of BRCA1 sequences (fig. 31), concatenated sequence data from five genes (fig. 33), and combined (nonmolecular + molecular) data when Chacodelphys is excluded (fig. 35). Subfamilial monophyly is also supported by a uniquely shared insertion (not coded for phylogenetic analysis) at the BRCA1 locus (fig. 31). However, only two morphological traits optimize as unambiguous synapomorphies of Didelphinae: an unpaired median teat and a strongly labially inflected centrocrista (subsequently reversed in Didelphini; appendix 5). In addition to genera referred to the tribes named above, the genus {Hyperdidelphys (here unassigned to tribe) clearly belongs to this subfamily based on characters described and illustrated by Goin and Pardiñas (1996) and others subsequently scored for phylogenetic analysis by D.A. Flores and the senior author. Tribe Marmosini Hershkovitz, 1992 CONTENTS: Marmosa, Monodelphis, {Thylatheridium, andTlacuatzin. DIAGNOSIS: Marmosines can be distinguished from other members of the subfamily Didelphinae by lacking a pouch (present in all Didelphini), by lacking an anteromedial bullar process spanning the transverse canal foramen (present in Metachirini and most Thylamyini), by lacking a fenestra in the squamosal-parietal suture (present in most Thylamyini), and by having the supraoccipital form part of the dorsal margin of the foramen magnum (absent in adult Metachirini and most adult Didelphini). REMARKS: The monophyly of Marmosini as construed herein is strongly supported by all analyses of BRCA1 (fig. 31), vWF (fig. 32), concatenated sequence data from five genes (fig. 33), and our combined (nonmolecular + molecular) dataset excluding Chacodelphys (fig. 35). However, no morphological trait optimizes as an unambiguous synapomorphy of this molecularly robust clade. Instead, all of the diagnostic features listed above appear to be plesiomorphic within the subfamily Didelphinae. Although the name Marmosini was used by Reig et al. (1985, 1987) it was not accompanied by an explicit statement of taxonomically differentiating characters. To
2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 101 the best of our knowledge, the first familygroup name based on Marmosa is technically available from Hershkovitz (1992b). An alternative name for this clade is Monodelphini, which McKenna and Bell (1997) attributed to Talice et al. (1961). However, Talice et al. did not mention any characters purported to differentiate Monodelphini from other didelphids, so their name is unavailable (ICZN, 1999: Article 13). To the best of our knowledge, the first familygroup name based on Monodelphis is also available from Hershkovitz (1992b). As first revisors in the sense of the Code (ICZN, 1999: Article 24), we select Marmosini Hershkovitz, 1992, to have precedence over Monodelphini Hershkovitz, 1992. Our reluctance to recognize any subtribal distinction between Marmosa and Monodelphis is based on the uncertain position of Tlacuatzin, which is not convincingly resolved despite the large amount of data at hand (.2000 parsimony-informative characters; table 12). In the event that a sister-group relationship between Marmosa and Tlacuatzin (weakly indicated by supermatrix analyses; figs. 33–36) were strongly supported by some future dataset, it would then make sense to recognize one subtribe (e.g., Marmosina) for those taxa and another (Monodelphina) for Monodelphis and {Thylatheridium. Although it seems indisputable that Monodelphis and {Thylatheridium are closely related (Goin and Rey, 1997), we note that this hypothesis has yet to be tested analytically, nor is it certainly known whether or not these taxa are reciprocally monophyletic. 25 Marmosa Gray, 1821 Figure 41 CONTENTS: We recognize two subgenera, Marmosa Gray, 1821, and Micoureus Lesson, 1842 (see Remarks, below). The subgenus Marmosa contains andersoni Pine, 1972; lepida Thomas, 1888 (including 25 In fact, it has been suggested that they are not: ‘‘Considero que Thylatheridium es un género derivado, en el Plioceno inferior, de una de las especies de Monodelphis que, ante posibilidades ecológicas favorables, aceleró su ritmo evolutivo, apartándose rápidamente de sus congéneres de evolución lenta …’’ (Reig, 1958: 90). grandis Tate, 1931); mexicana Merriam, 1897 (including mayensis Osgood, 1913; savannarum Goldman, 1917; and zeledoni Goldman, 1917); murina Linnaeus, 1758 (including bombascarae Anthony, 1922; chloe Thomas, 1907; dorsigera Linnaeus, 1758; duidae Tate, 1931; guianensis Kerr, 1792; klagesi J.A. Allen, 1900; macrotarsus Wagner, 1842; madeirensis Cabrera,. 1913; maranii Thomas, 1924; meridionalis Miranda-Ribeiro, 1936; moreirae Miranda-Ribeiro, 1936; muscula Cabanis, 1848; parata Thomas, 1911; roraimae Tate, 1931; tobagi Thomas, 1911; and waterhousei Tomes, 1860); quichua Thomas, 1899 (including musicola Osgood, 1913); robinsoni Bangs, 1898 (including casta Thomas, 1911; chapmani J.A. Allen, 1900; fulviventer Bangs, 1901; grenadae Thomas, 1911; isthmica Goldman, 1912; luridavolta Goodwin, 1961; mimetra Thomas, 1921; mitis Bangs, 1898; nesaea Thomas, 1911; pallidiventris Osgood, 1912; ruatanica Goldman, 1911; and simonsi Thomas, 1899); rubra Tate, 1931; tyleriana Tate, 1931 (including phelpsi Tate, 1939); and xerophila Handley and Gordon, 1979. The subgenus Micoureus contains alstoni J.A. Allen, 1900 (including nicaraguae Thomas, 1905); constantiae Thomas, 1904 (including budini Thomas, 1920); demerarae Thomas, 1905 (including areniticola Tate, 1931; domina Thomas, 1920; esmeraldae Tate, 1931; limae Thomas, 1920; and meridae Tate, 1931); paraguayana Tate, 1931 (including cinerea Temminck, 1824 [preoccupied]); phaea Thomas, 1899 (including perplexa Anthony, 1922); and regina Thomas, 1898 (including germana Thomas, 1904; mapiriensis Tate, 1931; parda Tate, 1931; rapposa Thomas, 1899; and rutteri Thomas, 1924). MORPHOLOGICAL DESCRIPTION: Combined length of adult head and body ca. 100– 210 mm; adult weight ca. 20–170 g. Rhinarium with two ventrolateral grooves on each side of median sulcus; dark circumocular mask present; pale supraocular spot absent; dark midrostral stripe absent; throat gland absent in some species (e.g., M. murina) but present in adult males of other species (e.g., M. mexicana). Dorsal pelage unpatterned, superficially brownish, reddish, or grayish, but dorsal hair bases always dark gray; dorsal guard hairs short and inconspicuous;
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PHYLOGENETIC RELATIONSHIPS AND CLAS
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ABSTRACT This report summarizes a d
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