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phylogenetic relationships and classification of didelphid marsupials ...

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2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 133<br />

<strong>and</strong> unfurred except at base; caudal integument<br />

bicolored (dark above, distinctly paler<br />

below); tail scales difficult to distinguish but<br />

apparently in annular series; caudal hairs all<br />

subequal in length <strong>and</strong> thickness; ventral<br />

caudal surface not modified for prehension.<br />

Premaxillary rostral process absent. Nasals<br />

long, extending anterior to I1 (concealing<br />

nasal orifice in dorsal view), <strong>and</strong> conspicuously<br />

widened posteriorly near premaxillarymaxillary<br />

suture. Maxillary turbinals elaborately<br />

branched. Two lacrimal foramina<br />

present on each side, exposed in lateral view<br />

anterior to orbital margin. Supraorbital<br />

margins smoothly rounded, without beads<br />

or crests; postorbital frontal processes usually<br />

absent (indistinct processes are occasionally<br />

developed in old adult males). Left <strong>and</strong><br />

right frontals <strong>and</strong> parietals separated by<br />

persistent median sutures. Parietal <strong>and</strong> alisphenoid<br />

bones in contact on lateral surface<br />

<strong>of</strong> braincase (no frontal-squamosal contact).<br />

Sagittal crest usually absent but weakly<br />

developed along midparietal suture (not<br />

extending anteriorly to frontals) in some<br />

large specimens. 31 Petrosal exposed through<br />

fenestra in parietal-squamosal suture in some<br />

individuals but not in others. Parietal-mastoid<br />

contact present (interparietal does not<br />

contact squamosal).<br />

Maxillopalatine fenestrae present but<br />

sometimes quite small; palatine fenestrae<br />

variable but usually present (Martin, 2005:<br />

fig. 2); maxillary fenestrae absent; posterolateral<br />

palatal foramina very long, usually<br />

extending anteriorly lingual to M4 protocones;<br />

posterior palatal morphology conforms to<br />

Didelphis morphotype (with strongly produced<br />

posterolateral corners, the choanae<br />

constricted behind). Maxillary <strong>and</strong> alisphenoid<br />

not in contact (separated by palatine) on<br />

floor <strong>of</strong> orbit. Transverse canal foramen<br />

present. Alisphenoid tympanic process<br />

smoothly globular, with well-developed anteromedial<br />

process enclosing extracranial<br />

course <strong>of</strong> m<strong>and</strong>ibular nerve (secondary foramen<br />

ovale present), <strong>and</strong> not contacting<br />

rostral tympanic process <strong>of</strong> petrosal. Anterior<br />

limb <strong>of</strong> ectotympanic directly suspended<br />

from basicranium. Stapes usually columelli-<br />

31 Among the specimens we examined, a small sagittal crest is<br />

best developed in UWZM 22422.<br />

form <strong>and</strong> microperforate or imperforate.<br />

Fenestra cochleae concealed in sinus formed<br />

by rostral <strong>and</strong> caudal tympanic processes <strong>of</strong><br />

petrosal. Paroccipital process small, rounded,<br />

<strong>and</strong> adnate to petrosal. Dorsal margin <strong>of</strong><br />

foramen magnum bordered by supraoccipital<br />

<strong>and</strong> exoccipitals, incisura occipitalis present.<br />

Two mental foramina usually present on<br />

lateral surface <strong>of</strong> each hemim<strong>and</strong>ible (three<br />

foramina are present unilaterally on two<br />

specimens examined); angular process acute<br />

<strong>and</strong> strongly inflected.<br />

Unworn crowns <strong>of</strong> I2–I5 symmetrically<br />

rhomboidal (‘‘premolariform’’), with subequal<br />

anterior <strong>and</strong> posterior cutting edges,<br />

slightly increasing in length (mesiodistal<br />

dimension) from I2 to I5. Upper canine<br />

(C1) alveolus in premaxillary-maxillary suture;<br />

C1 simple, without accessory cusps.<br />

First upper premolar (P1) smaller than<br />

posterior premolars but well formed <strong>and</strong><br />

not vestigial; third upper premolar (P3) taller<br />

than P2; P3 with posterior cutting edge only;<br />

upper milk premolar (dP3) large <strong>and</strong> molariform.<br />

Molars strongly carnassialized (postmetacristae<br />

much longer than postprotocrista);<br />

relative widths M1 , M2 , M3 ,<br />

M4; centrocrista strongly inflected labially on<br />

M1–M3; ect<strong>of</strong>lexus shallow or absent on M1,<br />

consistently present <strong>and</strong> distinct on M2,<br />

consistently deep on M3; anterolabial cingulum<br />

<strong>and</strong> preprotocrista discontinuous (anterior<br />

cingulum incomplete) on M3; postprotocrista<br />

without carnassial notch. Last upper<br />

tooth to erupt is P3.<br />

Lower incisors (i1–i4) with distinct lingual<br />

cusps. Lower canine (c1) erect, acutely<br />

pointed, <strong>and</strong> simple (without a posterior<br />

accessory cusp). Third lower premolar (p3)<br />

taller than p2; lower milk premolar (dp3)<br />

large, but trigonid incomplete (uni- or<br />

bicuspid). Hypoconid lingual to protoconid<br />

(not labially salient) on m3; hypoconulid<br />

twinned with entoconid on m1–m3; entoconid<br />

taller than hypoconulid on m1–m3.<br />

DISTRIBUTION: Most known specimens <strong>of</strong><br />

Lestodelphys have been collected in semidesert<br />

shrubl<strong>and</strong> <strong>and</strong> steppe habitats in Patagonian<br />

Argentina between 41u <strong>and</strong> 47uS latitude,<br />

but there are two outlying records from<br />

the Monte desert (between 32u <strong>and</strong> 38uS) that<br />

may represent a relictual population (Sauthier<br />

et al., 2007); all reported elevations

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