phylogenetic relationships and classification of didelphid marsupials ...

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164 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322 CRANIUM AND MANDIBLE Character 40: Premaxillae not produced anteriorly beyond alveolar margin of I1 (0); or forming a distinct, shelf-like rostral process anterior to I1 (1). Scoring criteria and information about the distribution of these traits among didelphid terminal taxa were provided by Voss and Jansa (2003: character 29). Among our outgroups, only Caenolestes, Dromiciops, andPerameles exhibit state 1. Character 41: Nasals produced anteriorly above or beyond I1, obscuring nasal orifice in dorsal view (0); or nasals posterior to I1, exposing nasal orifice dorsally (1); or nasals posterior to premaxillary facial processes, exposing incisive foramina in dorsal view (2). Scoring criteria and information about the distribution of states 0 and 1 among didelphid terminal taxa were provided by Voss and Jansa (2003: character 32). See text for comments about traits expressed by outgroup taxa. Because state 1 is obviously intermediate to states 0 and 2, this character was treated as ordered (0 « 1 « 2) in all of our analyses. Character 42: Nasals extend posteriorly between lacrimals (0); or truncated anterior to lacrimals (1). Among the taxa examined for this study, nasals that are truncated anterior to the lacrimals (state 1) are peculiar to peramelemorphians (see text). Character 43: Nasals conspicuously wider posteriorly than anteriorly (0); or nasals uniformly narrow, their lateral margins subparallel (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 33). Among our outgroups, only Echymipera and Perameles exhibit state 1. Character 44: Antorbital fenestra between nasal, frontal, and maxillary bones on each side of rostrum absent (0); or present (1). See text for a discussion of this character and its taxonomic distribution. Character 45: Maxilloturbinals large and elaborately branched (0); or small and unbranched (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 31). See text for comments about traits expressed by outgroup taxa. Character 46: Maxillary-jugal suture straight, crescentic, or irregular (0); or suture deeply bifid, dividing jugal into dorsal and ventral processes flanking antorbital fossa (1). See text for a discussion of this character and its taxonomic distribution. Character 47: Distinct frontal process of jugal absent (0); or present (1). See text for a discussion of this character and its taxonomic distribution. Character 48: Lacrimal margin of orbit smoothly rounded (0); or forming a distinct crest (1). See text for a discussion of this character and its taxonomic distribution. Character 49: One lacrimal foramen usually present (0); or two lacrimal foramina usually present (1). See text for a discussion of this character and its taxonomic distribution. Character 50: Foramen rotundum laterally exposed behind sphenorbital fissure (0); or foramen rotundum concealed from lateral view, partly or wholly confluent with sphenorbital fissure in rear of orbit (1). See text for a discussion of this character and its taxonomic distribution. Character 51: Maxillary and alisphenoid separate (0); or in contact on orbital floor (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 43). See text for comments about traits expressed by outgroup taxa. Character 52: Postorbital processes absent or indistinct (0); or present (1). This character was modified from Voss and Jansa’s (2003) character 34 by eliminating our former distinction between ‘‘flattened’’ and ‘‘not flattened’’ postorbital processes, which are not consistently distinguishable among nondidelphid marsupials. See text for comments about traits expressed by outgroup taxa. Character 53: Postorbital processes formed by frontals only (0); or by frontals and parietals (1). Inthe context of this study, state 1 is an autapomorphy of Glironia (see text). Character 54: Left and right frontals separated by ontogenetically persistent median suture (0); or left and right frontals partially or completely co-ossified in adults (1). See text for a discussion of this character and its taxonomic distribution. Character 55: Left and right parietals separated by ontogenetically persistent median suture (0); or left and right parietals partially or completely co-ossified in adults (1). See text for a discussion of this character and its taxonomic distribution. Character 56: Parietal and alisphenoid in contact on lateral aspect of braincase (0); or frontal and squamosal in contact (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 36). See text for comments about traits expressed by outgroup taxa. Character 57: Scars of M. temporalis origin on braincase not joined middorsally to form sagittal crest (0); or sagittal crest small, not extending to frontals (1); or sagittal crest large and extending to frontals (2). Descriptions of character states and an account of their taxonomic distribution among didelphid terminals were provided by Voss and Jansa (2003: character 35), but an intermediate state between absence of a sagittal crest and presence of a large crest extending to the frontals is recognized here. See text for comments about traits expressed by outgroup taxa. Because state 1 is obviously intermediate to states 0 and 2, this character was treated as ordered (0 « 1 « 2) in all of our analyses. Character 58: Fenestra in squamosal-parietal suture absent (0); or present (1). This character is modified from its original formulation (Voss and Jansa, 2003: character 37) by eliminating an intermediate state for polymorphisms, which were recoded as 0/1 for this analysis. See text for comments about traits expressed by outgroup taxa.
2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 165 Character 59: Interparietal absent (0); or present (1). See text for a discussion of this character and its taxonomic distribution. Character 60: Interparietal and supraoccipital separated by an open suture, at least in juvenile specimens (0); or interparietal-supraoccipital suture fused in juveniles and adults (1). See text for a discussion of this character, which was scored as inapplicable (‘‘-’’) for taxa lacking an interparietal, and as missing (‘‘?’’) for taxa in which it is not known whether an interparietal is present or absent (see above). Character 61: Parietal-mastoid contact present (0); or absent (1). See text for a discussion of this character and its taxonomic distribution. Character 62: ‘‘Accessory’’ fenestrae absent (0); or present (1). These nonvascular openings (first apparently named by Lyne and Mort, 1981: 124) perforate the maxillary palate at the level of P1 and/or P2, occupying a position in between the incisive foramina and the maxillopalatine fenestrae. In the context of this study, state 1 is an autapomorphy of Perameles. Character 63: Maxillopalatine fenestrae absent (0); or present (1). This character was modified from its original formulation (Voss and Jansa, 2003: character 38) by eliminating an intermediate state for polymorphism, which was recoded as 0/1 for this analysis. See text for comments about traits expressed by outgroup taxa. Character 64: Left and right maxillopalatine fenestrae separated by a median septum (0); or confluent, septum absent (1). See text for a discussion of this character and its taxonomic distribution. Character 65: Palatine fenestrae absent (0); or present (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 39). See text for comments about traits expressed by outgroup taxa. Character 66: Maxillary fenestrae absent (0); or present (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 40). See text for comments about traits expressed by outgroup taxa. Character 67: Posterolateral palatal foramina present, with complete bony margins (0); or absent, bony margins incomplete, represented by open notches in posterolateral palate (1). See text for a discussion of this character and its taxonomic distribution. Character 68: Posterolateral palatal foramina posterior to fourth molars (0); or extending lingual to M4 protocones (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 41). See text for comments about traits expressed by outgroup taxa. Although posterolateral palatal foramina are coded as absent in dasyurids, their position behind the M4 protocones in those taxa is indicated by the open notches mentioned in the preceding character description. Character 69: Posterior palate without distinct lateral corners, the choanae unconstricted behind (0); or posterior palate with distinct lateral corners, choanae constricted (1). Descriptions of the states of this character have been slightly reworded from their original phrasing in Voss and Jansa (2003: character 42), where the distribution of these traits was described for didelphid terminals. See text for comments about traits expressed by outgroup taxa. Character 70: Presphenoid exposed in roof of nasopharyngeal orifice above posterior palatal margin (0); or presphenoid concealed by vomer above posterior palatal margin (1). See text for a discussion of this character and its taxonomic distribution. Character 71: Left and right pterygoids separate (0); or in midline contact and forming a sagittal keel (1). See text for a discussion of this character and its taxonomic distribution. Character 72: Transverse canal foramen absent (0); or present (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 44). See text for comments about traits expressed by outgroup taxa. Character 73: Extracranial course of mandibular nerve not enclosed by bone (0); or consistently enclosed by anteromedial strut of alisphenoid bulla (1); or usually enclosed by posteromedial bullar lamina (2). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 45). See text for comments about traits expressed by outgroup taxa. In the absence of any compelling evidence for intermediacy among these alternative conditions, this character was treated as unordered in all of our analyses. Character 74: Alisphenoid tympanic process and rostral tympanic process of petrosal separated by a distinct gap (0); or in contact (1). See text for a discussion of this character and its taxonomic distribution. Character 75: Epitympanic recess small, snugly enclosing the mallear-incudal joint (0); or inflated, arching high above mallear-incudal joint (1). See text for a discussion of this character and its taxonomic distribution. Character 76: Rostral and caudal tympanic processes of petrosal separated by a distinct gap (0); or in contact and separated by a suture (1); or indistinguishably fused (2). States 0 and 1 of this character represent the same pattern of ingroup variation that Voss and Jansa (2003: character 47) described in terms of concealment of the fenestra cochleae. See text for comments about traits expressed by outgroup taxa. Because state 1 is plausibly intermediate to states 0 and 2, this character was treated as ordered (0 « 1 « 2) in all of our analyses. Character 77: Squamosal epitympanic sinus absent (0); or present (1). See text for a discussion of this character and its taxonomic distribution. Character 78: Anterior limb of ectotympanic directly attached to skull (0); or indirectly attached via malleus (1). Scoring criteria and information about the distribution of these traits among didelphid terminals were provided by Voss and Jansa (2003: character 46). See text for comments about traits expressed by outgroup taxa. Character 79: Ectotympanic laterally exposed (0); or concealed from lateral view inside bulla (1). See text
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PHYLOGENETIC RELATIONSHIPS AND CLAS
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CONTENTS Abstract .................
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ABSTRACT This report summarizes a d
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2009 VOSS AND JANSA: DIDELPHID MARS
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