phylogenetic relationships and classification of didelphid marsupials ...
phylogenetic relationships and classification of didelphid marsupials ... phylogenetic relationships and classification of didelphid marsupials ...
116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322 seems probable that at least some of the many putative synonyms of Chironectes minimus will be found to represent valid taxa in future analyses of morphological and/or molecular data. Didelphis Linnaeus, 1758 Figure 46 CONTENTS: albiventris Lund, 1840 (including bonariensis Marelli, 1930; dennleri Marelli, 1930; lechei Ihering, 1892; leucotis Wagner, 1847; paraguayensis J.A. Allen, 1902; poecilotis Wagner, 1842; and poecilonota Schinz, 1844); aurita Wied-Neuweid, 1826 (including koseritzi Ihering, 1892; longipilis Miranda-Ribeiro, 1935; and melanoidis Miranda-Ribeiro, 1935); imperfecta Mondolfi and Pérez-Hernández, 1984; marsupialis Linnaeus, 1758 (including battyi Thomas, 1902; cancrivora Gmelin, 1788; caucae J.A. Allen, 1900; colombica J.A. Allen, 1900; etensis J.A. Allen, 1902; insularis J.A. Allen, 1902; karkinophaga Zimmermann, 1780; particeps Goldman, 1917; richmondi J.A. Allen, 1901; and tabascensis J.A. Allen, 1901); pernigra J.A. Allen, 1900 (including andina J.A. Allen, 1902; and meridensis J.A. Allen, 1902); and virginiana Kerr, 1792 (including boreoamericana J.A. Allen, 1902; breviceps Bennett, 1833; californica Bennett, 1833; cozumelae Merriam, 1901; illinensium Link, 1795; pigra Bangs, 1898; pilosissima Link, 1795; pruinosa Wagner, 1843; texensis J.A. Allen, 1901; woapink Barton, 1806; and yucatanensis J.A. Allen, 1901). MORPHOLOGICAL DESCRIPTION: Combined length of adult head and body 310–495 mm; adult weight 600–5100 g. Rhinarium with one ventrolateral groove on each side of median sulcus; dark circumocular mask indistinct in some species (D. aurita, D. marsupialis, D. virginiana) but distinct in others (D. albiventris, D. imperfecta, D. pernigra); pale supraocular spot absent; dark midrostral stripe absent; throat gland absent. Dorsal pelage unpatterned; dorsal underfur white; dorsal guard hairs long, coarse and conspicuous, giving the pelage a distinctively shaggy appearance; ventral fur pale (usually whitish) tipped with black. Manus mesaxonic (dIII . dIV); manual claws usually longer than fleshy apical pads of digits; dermato- glyph-bearing manual plantar pads present; central palmar epithelium smooth or sparsely tuberculate; carpal tubercles absent. Pedal digits unwebbed; dIV slightly longer than other pedal digits in some species (e.g., D. marsupialis) or dII, dIII, and dIV subequal (e.g., in D. albiventris and D. virginiana); plantar surface of heel naked. Pouch well developed, opening anteriorly; mammae normally 3–1–3 5 7 to 6–1–6 5 13 or more 29 ; cloaca present. Tail slightly longer than combined length of head and body, slender and muscular (not incrassate); basal 1/6 to 1/4 densely furred (covered with body pelage); naked caudal integument blackish proximally and abruptly whitish distally; caudal scales in spiral series, each scale usually with three subequal bristlelike hairs emerging from distal margin; ventral caudal surface variably modified for prehension distally (prehensile modifications are more strongly developed in D. auritus and D. marsupialis than in the other species), but dermatoglyph-bearing apical pad consistently present. Premaxillary rostral process absent. Nasals short, not produced anteriorly above I1 (exposing nasal orifice in dorsal view), and conspicuously widened posteriorly near maxillary-frontal suture. Maxillary turbinals elaborately branched. Lacrimal foramina one or two on each side, exposed laterally on orbital margin or on face just anterior to orbit. Postorbital processes bluntly pyramidal, maximally developed in old adults. Left and right frontals co-ossified (midfrontal suture incomplete or absent), but left and right parietals separated by persistent midparietal suture. Parietal and alisphenoid in contact on lateral braincase (no frontalsquamosal contact). Sagittal crest large, well developed, and extending onto frontals. Petrosal not exposed laterally through fenestra in squamosal-parietal suture (fenestra absent). Parietal-mastoid contact absent (interparietal narrowly contacts squamosal). Maxillopalatine and palatine fenestrae present; maxillary fenestrae absent; posterolateral palatal foramina not extending anteriorly lingual to M4 protocones; posterior 29 Occasional records of Didelphis virginiana with up to 17 pouch young (cited by Hamilton, 1958) suggest that teat counts in this species may sometimes exceed 6–1–6 5 13.
2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 117 Fig. 46. Didelphis marsupialis (based on AMNH 266459, an adult male from Paracou, French Guiana). palate with prominent lateral corners, the choanae abruptly constricted behind. Maxillary and alisphenoid not in contact (separated by palatine) on floor of orbit. Transverse canal foramen present. Alisphenoid tympanic process small and uninflated, usually with posteromedial lamina enclosing extracranial course of mandibular nerve (secondary fora-
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116 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322<br />
seems probable that at least some <strong>of</strong> the<br />
many putative synonyms <strong>of</strong> Chironectes<br />
minimus will be found to represent valid taxa<br />
in future analyses <strong>of</strong> morphological <strong>and</strong>/or<br />
molecular data.<br />
Didelphis Linnaeus, 1758<br />
Figure 46<br />
CONTENTS: albiventris Lund, 1840 (including<br />
bonariensis Marelli, 1930; dennleri Marelli,<br />
1930; lechei Ihering, 1892; leucotis<br />
Wagner, 1847; paraguayensis J.A. Allen,<br />
1902; poecilotis Wagner, 1842; <strong>and</strong> poecilonota<br />
Schinz, 1844); aurita Wied-Neuweid,<br />
1826 (including koseritzi Ihering, 1892; longipilis<br />
Mir<strong>and</strong>a-Ribeiro, 1935; <strong>and</strong> melanoidis<br />
Mir<strong>and</strong>a-Ribeiro, 1935); imperfecta Mondolfi<br />
<strong>and</strong> Pérez-Hernández, 1984; marsupialis<br />
Linnaeus, 1758 (including battyi Thomas,<br />
1902; cancrivora Gmelin, 1788; caucae J.A.<br />
Allen, 1900; colombica J.A. Allen, 1900;<br />
etensis J.A. Allen, 1902; insularis J.A. Allen,<br />
1902; karkinophaga Zimmermann, 1780; particeps<br />
Goldman, 1917; richmondi J.A. Allen,<br />
1901; <strong>and</strong> tabascensis J.A. Allen, 1901);<br />
pernigra J.A. Allen, 1900 (including <strong>and</strong>ina<br />
J.A. Allen, 1902; <strong>and</strong> meridensis J.A. Allen,<br />
1902); <strong>and</strong> virginiana Kerr, 1792 (including<br />
boreoamericana J.A. Allen, 1902; breviceps<br />
Bennett, 1833; californica Bennett, 1833;<br />
cozumelae Merriam, 1901; illinensium Link,<br />
1795; pigra Bangs, 1898; pilosissima Link,<br />
1795; pruinosa Wagner, 1843; texensis J.A.<br />
Allen, 1901; woapink Barton, 1806; <strong>and</strong><br />
yucatanensis J.A. Allen, 1901).<br />
MORPHOLOGICAL DESCRIPTION: Combined<br />
length <strong>of</strong> adult head <strong>and</strong> body 310–495 mm;<br />
adult weight 600–5100 g. Rhinarium with<br />
one ventrolateral groove on each side <strong>of</strong><br />
median sulcus; dark circumocular mask<br />
indistinct in some species (D. aurita, D.<br />
marsupialis, D. virginiana) but distinct in<br />
others (D. albiventris, D. imperfecta, D.<br />
pernigra); pale supraocular spot absent; dark<br />
midrostral stripe absent; throat gl<strong>and</strong> absent.<br />
Dorsal pelage unpatterned; dorsal underfur<br />
white; dorsal guard hairs long, coarse <strong>and</strong><br />
conspicuous, giving the pelage a distinctively<br />
shaggy appearance; ventral fur pale (usually<br />
whitish) tipped with black. Manus mesaxonic<br />
(dIII . dIV); manual claws usually longer<br />
than fleshy apical pads <strong>of</strong> digits; dermato-<br />
glyph-bearing manual plantar pads present;<br />
central palmar epithelium smooth or sparsely<br />
tuberculate; carpal tubercles absent. Pedal<br />
digits unwebbed; dIV slightly longer than<br />
other pedal digits in some species (e.g., D.<br />
marsupialis) or dII, dIII, <strong>and</strong> dIV subequal<br />
(e.g., in D. albiventris <strong>and</strong> D. virginiana);<br />
plantar surface <strong>of</strong> heel naked. Pouch well<br />
developed, opening anteriorly; mammae normally<br />
3–1–3 5 7 to 6–1–6 5 13 or more 29 ;<br />
cloaca present. Tail slightly longer than<br />
combined length <strong>of</strong> head <strong>and</strong> body, slender<br />
<strong>and</strong> muscular (not incrassate); basal 1/6 to<br />
1/4 densely furred (covered with body pelage);<br />
naked caudal integument blackish proximally<br />
<strong>and</strong> abruptly whitish distally; caudal scales in<br />
spiral series, each scale usually with three<br />
subequal bristlelike hairs emerging from<br />
distal margin; ventral caudal surface variably<br />
modified for prehension distally (prehensile<br />
modifications are more strongly developed in<br />
D. auritus <strong>and</strong> D. marsupialis than in the other<br />
species), but dermatoglyph-bearing apical<br />
pad consistently present.<br />
Premaxillary rostral process absent. Nasals<br />
short, not produced anteriorly above I1<br />
(exposing nasal orifice in dorsal view), <strong>and</strong><br />
conspicuously widened posteriorly near maxillary-frontal<br />
suture. Maxillary turbinals<br />
elaborately branched. Lacrimal foramina<br />
one or two on each side, exposed laterally<br />
on orbital margin or on face just anterior to<br />
orbit. Postorbital processes bluntly pyramidal,<br />
maximally developed in old adults. Left<br />
<strong>and</strong> right frontals co-ossified (midfrontal<br />
suture incomplete or absent), but left <strong>and</strong><br />
right parietals separated by persistent midparietal<br />
suture. Parietal <strong>and</strong> alisphenoid in<br />
contact on lateral braincase (no frontalsquamosal<br />
contact). Sagittal crest large, well<br />
developed, <strong>and</strong> extending onto frontals.<br />
Petrosal not exposed laterally through fenestra<br />
in squamosal-parietal suture (fenestra<br />
absent). Parietal-mastoid contact absent (interparietal<br />
narrowly contacts squamosal).<br />
Maxillopalatine <strong>and</strong> palatine fenestrae<br />
present; maxillary fenestrae absent; posterolateral<br />
palatal foramina not extending anteriorly<br />
lingual to M4 protocones; posterior<br />
29 Occasional records <strong>of</strong> Didelphis virginiana with up to 17<br />
pouch young (cited by Hamilton, 1958) suggest that teat counts<br />
in this species may sometimes exceed 6–1–6 5 13.