phylogenetic relationships and classification of didelphid marsupials ...
phylogenetic relationships and classification of didelphid marsupials ... phylogenetic relationships and classification of didelphid marsupials ...
134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322 associated with collected specimens are less than 1000 m above sea level (Birney et al., 1996; Martin, 2003). As noted by Flores et al. (2007), the distribution of Lestodelphys extends further south than that of any other living marsupial. Marmosops Matschie, 1916 Figures 52, 53 CONTENTS: bishopi Pine, 1981; cracens Handley and Gordon, 1979; creightoni Voss et al., 2004; fuscatus Thomas, 1896 (including carri J.A. Allen and Chapman, 1897; and perfuscus Thomas, 1924); handleyi Pine, 1981; impavidus Tschudi, 1845 (including caucae Thomas, 1900; celicae Anthony, 1922; madescens Osgood, 1913; oroensis Anthony, 1922; sobrinus Thomas, 1913; and ucayalensis Tate, 1931); incanus Lund, 1840 (including bahiensis Tate, 1931; and scapulatus Burmeister, 1856); invictus Goldman, 1912; juninensis Tate, 1931; neblina Gardner, 1990; noctivagus Tschudi, 1844 (including albiventris Tate, 1931; collega Thomas, 1920; dorothea Thomas, 1911; keaysi J.A. Allen, 1900; leucastrus Thomas, 1927; lugendus Thomas, 1927; neglectus Osgood, 1915; politus Cabrera, 1913; purui Miller, 1913; stollei Miranda-Ribeiro, 1936; and yungasensis Tate, 1931); ocellatus Tate, 1931; parvidens Tate, 1931; paulensis Tate, 1931; and pinheiroi Pine, 1981 (including woodalli Pine, 1981). MORPHOLOGICAL DESCRIPTION: Combined length of adult head and body ca. 90– 195 mm; adult weight ca. 20–140 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 consistently present in adult males of some species (e.g., M. incanus, M. noctivagus) but apparently absent in others (e.g., M. impavidus, M. parvidens). Dorsal pelage unpatterned, usually some shade of dull grayish brown, but distinctly reddish or grayish in some species; dorsal hair bases always dark gray; dorsal guard hairs short and inconspicuous; ventral fur self-colored (whitish, buffy, or brown) or entirely or partially gray based. Manus mesaxonic (dIII . dIV); manual claws small, shorter than fleshy apical pads of digits; dermatoglyph-bearing manual plan- tar pads present; central palmar epithelium smooth or sparsely tuberculate; lateral carpal tubercles externally conspicuous on wrists of large adult males; medial carpal tubercles absent in both sexes. Pedal digits unwebbed; dIV longer than other pedal digits; plantar surface of heel macroscopically naked (a microscopic pelage of very short hairs, not coarse fur, is usually present). Pouch absent; mammae varying among examined species from 3–1–3 5 7 to 7–1–7 5 15, anteriormost pairs ‘‘pectoral’’ when mammae $11; cloaca present. Tail longer than combined length of head and body, slender and muscular (not incrassate), and macroscopically naked (without a conspicuously furred base); naked caudal integument uniformly dark (usually grayish), bicolored (dark above, pale below), or parti-colored (dark distally, paler distally); caudal scales in spiral series, each scale with three bristlelike hairs emerging from caudal margin; median hair of each caudal-scale triplet usually much thicker and darker than lateral hairs; ventral caudal surface modified for prehension distally (with naked median groove and apical pad bearing dermatoglyphs). Premaxillary rostral process long and well developed in some species (e.g., M. parvidens, M. pinheiroi) but short in others (e.g., M. noctivagus) and apparently absent in some (M. incanus). Nasals long, extending anteriorly beyond I1 (concealing nasal orifice in dorsal view), and conspicuously widened posteriorly near maxillary-frontal suture except in M. incanus (which has narrow, more or less parallel-sided nasals). Maxillary turbinals (viewed through nasal orifice) elaborately branched. Lacrimal foramina, usually two on each side, concealed from lateral view inside orbit (e.g., in M. parvidens) or exposed on orbital margin (e.g., in M. pinheiroi). Interorbital region without abrupt constrictions; supraorbital margins rounded in some species (e.g., M. parvidens), beaded in others (e.g., M. noctivagus); postorbital processes usually absent except in old males of some species (e.g., M. impavidus 32 ). Left and right frontals and parietals separated by persistent median sutures. Parietal and alisphenoid in 32 Postorbital processes in this species are best developed in AMNH 272760.
2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 135 Fig. 52. Marmosops creightoni (based on CBF 6552 and UMMZ 155999, adult males from the valley of the Río Zongo, La Paz, Bolivia).
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134 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322<br />
associated with collected specimens are less<br />
than 1000 m above sea level (Birney et al.,<br />
1996; Martin, 2003). As noted by Flores et al.<br />
(2007), the distribution <strong>of</strong> Lestodelphys extends<br />
further south than that <strong>of</strong> any other<br />
living marsupial.<br />
Marmosops Matschie, 1916<br />
Figures 52, 53<br />
CONTENTS: bishopi Pine, 1981; cracens<br />
H<strong>and</strong>ley <strong>and</strong> Gordon, 1979; creightoni Voss<br />
et al., 2004; fuscatus Thomas, 1896 (including<br />
carri J.A. Allen <strong>and</strong> Chapman, 1897; <strong>and</strong><br />
perfuscus Thomas, 1924); h<strong>and</strong>leyi Pine, 1981;<br />
impavidus Tschudi, 1845 (including caucae<br />
Thomas, 1900; celicae Anthony, 1922; madescens<br />
Osgood, 1913; oroensis Anthony,<br />
1922; sobrinus Thomas, 1913; <strong>and</strong> ucayalensis<br />
Tate, 1931); incanus Lund, 1840 (including<br />
bahiensis Tate, 1931; <strong>and</strong> scapulatus Burmeister,<br />
1856); invictus Goldman, 1912;<br />
juninensis Tate, 1931; neblina Gardner, 1990;<br />
noctivagus Tschudi, 1844 (including albiventris<br />
Tate, 1931; collega Thomas, 1920; dorothea<br />
Thomas, 1911; keaysi J.A. Allen, 1900;<br />
leucastrus Thomas, 1927; lugendus Thomas,<br />
1927; neglectus Osgood, 1915; politus Cabrera,<br />
1913; purui Miller, 1913; stollei Mir<strong>and</strong>a-Ribeiro,<br />
1936; <strong>and</strong> yungasensis Tate,<br />
1931); ocellatus Tate, 1931; parvidens Tate,<br />
1931; paulensis Tate, 1931; <strong>and</strong> pinheiroi Pine,<br />
1981 (including woodalli Pine, 1981).<br />
MORPHOLOGICAL DESCRIPTION: Combined<br />
length <strong>of</strong> adult head <strong>and</strong> body ca. 90–<br />
195 mm; adult weight ca. 20–140 g. Rhinarium<br />
with two ventrolateral grooves on each<br />
side <strong>of</strong> median sulcus; dark circumocular<br />
mask present; pale supraocular spot absent;<br />
dark midrostral stripe absent; throat gl<strong>and</strong><br />
consistently present in adult males <strong>of</strong> some<br />
species (e.g., M. incanus, M. noctivagus) but<br />
apparently absent in others (e.g., M. impavidus,<br />
M. parvidens). Dorsal pelage unpatterned,<br />
usually some shade <strong>of</strong> dull grayish<br />
brown, but distinctly reddish or grayish in<br />
some species; dorsal hair bases always dark<br />
gray; dorsal guard hairs short <strong>and</strong> inconspicuous;<br />
ventral fur self-colored (whitish, buffy,<br />
or brown) or entirely or partially gray based.<br />
Manus mesaxonic (dIII . dIV); manual<br />
claws small, shorter than fleshy apical pads<br />
<strong>of</strong> digits; dermatoglyph-bearing manual plan-<br />
tar pads present; central palmar epithelium<br />
smooth or sparsely tuberculate; lateral carpal<br />
tubercles externally conspicuous on wrists <strong>of</strong><br />
large adult males; medial carpal tubercles<br />
absent in both sexes. Pedal digits unwebbed;<br />
dIV longer than other pedal digits; plantar<br />
surface <strong>of</strong> heel macroscopically naked (a<br />
microscopic pelage <strong>of</strong> very short hairs, not<br />
coarse fur, is usually present). Pouch absent;<br />
mammae varying among examined species<br />
from 3–1–3 5 7 to 7–1–7 5 15, anteriormost<br />
pairs ‘‘pectoral’’ when mammae $11; cloaca<br />
present. Tail longer than combined length <strong>of</strong><br />
head <strong>and</strong> body, slender <strong>and</strong> muscular (not<br />
incrassate), <strong>and</strong> macroscopically naked<br />
(without a conspicuously furred base); naked<br />
caudal integument uniformly dark (usually<br />
grayish), bicolored (dark above, pale below),<br />
or parti-colored (dark distally, paler distally);<br />
caudal scales in spiral series, each scale with<br />
three bristlelike hairs emerging from caudal<br />
margin; median hair <strong>of</strong> each caudal-scale<br />
triplet usually much thicker <strong>and</strong> darker than<br />
lateral hairs; ventral caudal surface modified<br />
for prehension distally (with naked median<br />
groove <strong>and</strong> apical pad bearing dermatoglyphs).<br />
Premaxillary rostral process long <strong>and</strong> well<br />
developed in some species (e.g., M. parvidens,<br />
M. pinheiroi) but short in others (e.g., M.<br />
noctivagus) <strong>and</strong> apparently absent in some<br />
(M. incanus). Nasals long, extending anteriorly<br />
beyond I1 (concealing nasal orifice in<br />
dorsal view), <strong>and</strong> conspicuously widened<br />
posteriorly near maxillary-frontal suture except<br />
in M. incanus (which has narrow, more<br />
or less parallel-sided nasals). Maxillary turbinals<br />
(viewed through nasal orifice) elaborately<br />
branched. Lacrimal foramina, usually<br />
two on each side, concealed from lateral view<br />
inside orbit (e.g., in M. parvidens) or exposed<br />
on orbital margin (e.g., in M. pinheiroi).<br />
Interorbital region without abrupt constrictions;<br />
supraorbital margins rounded in some<br />
species (e.g., M. parvidens), beaded in others<br />
(e.g., M. noctivagus); postorbital processes<br />
usually absent except in old males <strong>of</strong> some<br />
species (e.g., M. impavidus 32 ). Left <strong>and</strong> right<br />
frontals <strong>and</strong> parietals separated by persistent<br />
median sutures. Parietal <strong>and</strong> alisphenoid in<br />
32<br />
Postorbital processes in this species are best developed in<br />
AMNH 272760.