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phylogenetic relationships and classification of didelphid marsupials ...

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2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 57<br />

Fig. 23. Labial views <strong>of</strong> unworn anterior<br />

m<strong>and</strong>ibular dentitions illustrating taxonomic differences<br />

in lower canine (c1) morphology. A,<br />

Lestodelphys halli (MMNH 17171) with erect,<br />

acutely pointed, <strong>and</strong> simple c1. B, Marmosa<br />

mexicana (AMNH 17136) with semiprocumbent<br />

<strong>and</strong> flat-crowned but simple c1. C, Marmosops<br />

pinheiroi (AMNH 267342) with fully premolariform<br />

c1 (procumbent <strong>and</strong> flat crowned with<br />

posterior accessory cusp).<br />

tendencies are minimally indicated on the<br />

unworn teeth <strong>of</strong> Chacodelphys <strong>and</strong> Metachirus<br />

(which are flattened along their anterior<br />

margin but have no posterior cingulum),<br />

they are increasingly apparent in Cryptonanus,<br />

Gracilinanus, Marmosa (fig. 23B), <strong>and</strong><br />

Thylamys. Premolarization <strong>of</strong> the lower<br />

canine is maximally developed in the smaller<br />

species <strong>of</strong> Marmosops (e.g., M. parvidens <strong>and</strong><br />

M. pinheiroi), where c1 <strong>and</strong> p1–3 form a<br />

series <strong>of</strong> four strikingly similar teeth (fig.<br />

23C). Generally speaking, taxa with welldeveloped<br />

accessory cusps on the upper<br />

canine also have a well-developed posterior<br />

accessory cusp on c1, so variation in these<br />

occluding teeth is at least partially correlated.<br />

Large dasyurids (e.g., Dasyurus) have erect,<br />

unicuspid lower canines, but many small<br />

dasyurids (e.g., Sminthopsis) have procumbent,<br />

premolariform teeth. The lower canines<br />

are small <strong>and</strong> blunt in peramelemorphians. In<br />

Dromiciops, c1 is procumbent <strong>and</strong> more or<br />

less premolariform (Marshall, 1982b: fig. 14).<br />

LOWER PREMOLARS: Didelphid lower premolars<br />

are unremarkable teeth, each having a<br />

single dominant cusp <strong>and</strong> a well-developed<br />

distal cingulum that <strong>of</strong>ten produces a posterior<br />

accessory cusp. Small diastemata are<br />

sometimes present between p1 <strong>and</strong> c1,<br />

between p1 <strong>and</strong> p2, <strong>and</strong> (rarely) between p2<br />

<strong>and</strong> p3. The lower first premolar (p1) is<br />

vestigial <strong>and</strong> occlusally featureless in Caluromys<br />

<strong>and</strong> Caluromysiops, but it is well<br />

developed in all <strong>of</strong> the other genera.<br />

The second lower premolar is distinctly<br />

taller than p3 in most <strong>didelphid</strong>s, but p2 <strong>and</strong><br />

p3 are subequal in Marmosops incanus <strong>and</strong><br />

most examined species <strong>of</strong> Monodelphis <strong>and</strong><br />

Thylamys. In Lestodelphys <strong>and</strong> in certain<br />

other species <strong>of</strong> Monodelphis (e.g., M. emiliae),<br />

however, p3 is distinctly taller than p2.<br />

Most other plesiomorphic <strong>marsupials</strong> have<br />

three lower premolars that are essentially<br />

similar to those <strong>of</strong> <strong>didelphid</strong>s, although some<br />

dasyurids (e.g., Dasyurus) have only two.<br />

Among the taxa we examined, p3 is taller<br />

than p2 in caenolestids, Dromiciops (see<br />

Marshall, 1982b: fig. 14), some dasyurids<br />

(e.g., Murexia), <strong>and</strong> some peramelemorphians<br />

(e.g., Echymipera), but p2 <strong>and</strong> p3 are<br />

subequal in height or p2 is the taller tooth in<br />

other dasyurids (e.g., Sminthopsis) <strong>and</strong> peramelemorphians<br />

(e.g., Macrotis).

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