phylogenetic relationships and classification of didelphid marsupials ...
phylogenetic relationships and classification of didelphid marsupials ... phylogenetic relationships and classification of didelphid marsupials ...
86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322 incomplete taxon sampling (Horovitz, 1999); this notion provides some justification for naming monotypic higher taxa to accommodate future discoveries of new forms. Point (5) acknowledges the necessity for widely accepted rules to govern the use of technical names. Because detailed generic synonymies have recently been published by Gardner (2008), we do not provide them here. However, a chronological list of available names with information about type species and current status (table 15) contains most of the information needed to understand current binomial usage. Under the indented heading ‘‘Contents’’ we list all of the valid lower-level taxa included by each higher-level taxon; for example, the genera referred to a tribe, or the species referred to a genus. In lists of congeneric species, those currently regarded as valid are in boldface, followed by an alphabetic list of available junior synonyms in parentheses; however, names of doubtful application (e.g., brasiliensis Liais, 1872, which might be a junior synonym of more than one currently recognized species of Didelphis; see Cerqueira and Tribe, 2008) are not included. Under ‘‘Diagnosis’’ we list unique combinations of traits that distinguish members of suprageneric taxa from other marsupials, whether or not such traits are unambiguously assignable by parsimony as relevant synapomorphies (listed in appendix 5). Under ‘‘Morphological description’’ (for genera) we list all phenotypic descriptors that apply to included species; traits of particular importance for identification are indicated by italics. Ranges of metrical traits (e.g., 170– 210 mm) are approximations that match or slightly exceed known minimal and maximal adult values that we believe to be reliable based on published and unpublished sources. Except as noted otherwise, all descriptions of cranial traits are based on adult specimens, and all dental descriptions are based on unworn teeth. Under ‘‘Remarks’’ we discuss evidence for monophyly and other issues that relate to taxon recognition and rank. Although ranks are biologically arbitrary, they affect spelling and information retrieval in the prevailing Linnaean system, and they are regulated by widely accepted rules of biological nomenclature. The lack of species-level revisionary studies for most genera is also discussed under this heading. Family Didelphidae Gray, 1821 CONTENTS: Glironiinae, Caluromyinae, Hyladelphinae, and Didelphinae. DIAGNOSIS: Didelphids can be distinguished unambiguously from other marsupials by their soft (nonspinous) body pelage; five subequal claw-bearing manual digits; five separate pedal digits (loosely connected by webbing in Chironectes), of which the hallux (dI) is large and opposable and the second (dII) bears an asymmetrical grooming claw; inguinal cloaca; prehensile tail (secondarily lacking external evidence of prehensility in some taxa); long nasal bones (extending anteriorly beyond the facial processes of the premaxillae and posteriorly between the lacrimals); unfenestrated rostrum; uninflected maxillary-jugal suture; distinct foramen rotundum (not confluent or sharing a common vestibule with the sphenorbital fissure); alisphenoid-parietal contact (except in Metachirus); large interparietal fused to the supraoccipital; complete posterolateral palatal foramina; deeply divided vomer that does not conceal the presphenoid or extend posteriorly into the mesopterygoid fossa; lack of midline contact between the left and right pterygoids; unkeeled basisphenoid; well-developed tympanic process of the alisphenoid; well-developed rostral tympanic process of the petrosal; lack of a squamosal epitympanic sinus; laterally exposed ectotympanic; unspecialized malleus (with long sharply inflected neck, small orbicular apophysis, and well-developed lamina); distinct mastoid and paroccipital processes; unreduced dental complement of 50 teeth; nongliriform incisors; large milk premolars (except in Hyladelphys); tribosphenic upper molars, each with a continuous stylar shelf, uninterrupted centrocrista, and reduced or absent para- and metaconules; and tribosphenic lower molars, each with a welldeveloped anterior cingulid (notched for the hypoconulid of the preceding tooth), notched paracristids, and without any trace of a posterior cingulid.
2009 VOSS AND JANSA: DIDELPHID MARSUPIALS 87 Didelphids have often been compared with dasyurids, but the literature contains no adequate statement of the many characters that distinguish these superficially similar yet highly divergent clades. Among the external and craniodental features treated in this report, dasyurids consistently differ from didelphids by their small, nonopposable hallux; lack of a grooming claw on pedal digit II; a nonprehensile tail (provided with a terminal tuft of hairs that is never present in opossums); anteriorly truncated and medially notched nasals; a small interparietal that, when present in juvenile skulls, is suturally distinct from the supraoccipital; incomplete posterolateral palatal foramina; an undivided vomer that extends posteriorly to underlie the presphenoid within the mesopterygoid fossa; well-developed squamosal epitympanic sinuses; four upper and three lower incisors; a distinct posterior lobe on the unworn cutting edge of i3; vestigial milk premolars; and a distinct posterior cingulid on the lower molars. Originally described as a didelphid by Thomas (1894) and long maintained in that family by subsequent authors (e.g., Simpson, 1945), Dromiciops differs from opossums by its basicaudal cloaca; a foramen rotundum recessed in a common vestibule with the sphenorbital fissure; frontal-squamosal contact; a large interparietal that is suturally distinct from the supraoccipital; an undivided vomer that underlies the presphenoid within the mesopterygoid fossa; left and right pterygoid bones that contact one another in the midline; keeled basisphenoid; a concealed ectotympanic; a specialized malleus (with a short uninflected neck, no orbicular apophysis, and no lamina); lack of a paroccipital process; a discontinuous stylar shelf on M1; an unstaggered i2 alveolus; and a vestigial anterior cingulid on the lower molars (of which m1 lacks a hypoconulid notch). Caenolestids, peramelids, and other marsupials are sufficiently distinct from opossums that explicit comparisons would be pointless here. Some stem metatherians (e.g., herpetotheriids) that are strikingly similar to didelphids in most respects (including ear morphology; Gabbert, 1998) differ from didelphids by having a distinct posterior cingulid on the lower molars in addition to the diagnostic endocranial and postcranial features described by Sánchez-Villagra et al. (2007). 23 REMARKS: A wide range of fossil taxa have at one time or another been regarded as didelphids (e.g., by Simpson, 1935, 1945; Clemens, 1979; Marshall, 1981; McKenna and Bell, 1997), but phylogenetic analyses suggest that most of the extinct forms once thought to be closely related to Recent opossums (e.g., {Alphadon, {Andinodelphys, {Glasbius, {Herpetotherium, {Jaskhadelphys, {Pediomys) are stem metatherians and not members of the crown group Marsupialia (Rougier et al., 1998; Wible et al., 2001; Luo et al., 2003; Sánchez-Villagra et al., 2007). Herein we explicitly restrict Didelphidae to living didelphimorphians, their most recent common ancestor, and all of its descendants. Even more restrictive concepts of Didelphidae have been proposed, but none is widely accepted. Hershkovitz (1992b), for example, used Didelphidae to include just the large opossums with 2n 5 22 chromosomes (Chironectes, Didelphis, Lutreolina, Philander), whereas Kirsch and Palma (1995) excluded Glironia, Caluromys, and Caluromysiops from the family. Because Didelphidae in any of these applications (ours, Hershkovitz’s, or Kirsch and Palma’s) is monophyletic, the choice among them must be justified by other criteria. In our view, the name Didelphidae and its colloquial equivalent (‘‘didelphids’’) are so deeply entrenched in the literature as referring to all Recent opossums that more restrictive applications would serve no adequate compensatory purpose. Although didelphid monophyly is impressively supported by nucleotide sequence data 23 As noted above, the posterior cingulid is absent in didelphids, caenolestids, microbiotherians, and peramelids, so it optimizes as an unambiguous marsupial synapomorphy on all metatherian topologies in which these groups are recovered as basal clades (e.g., Meredith et al. 2008). Because most Cretaceous and early Tertiary metatherians are known only from teeth, this trait provides a potentially useful criterion for distinguishing stem taxa from members of the crown clade. For example, several of the Paleocene Brazilian fossils classified as didelphids by Marshall (1987) are described by that author as having posterior cingulids and might plausibly be regarded on that basis as stem metatherians. By contrast, Marsupialia (in the crown-group sense) does not seem to be diagnosable by any upper molar trait.
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86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 322<br />
incomplete taxon sampling (Horovitz, 1999);<br />
this notion provides some justification for<br />
naming monotypic higher taxa to accommodate<br />
future discoveries <strong>of</strong> new forms. Point<br />
(5) acknowledges the necessity for widely<br />
accepted rules to govern the use <strong>of</strong> technical<br />
names.<br />
Because detailed generic synonymies have<br />
recently been published by Gardner (2008),<br />
we do not provide them here. However, a<br />
chronological list <strong>of</strong> available names with<br />
information about type species <strong>and</strong> current<br />
status (table 15) contains most <strong>of</strong> the information<br />
needed to underst<strong>and</strong> current binomial<br />
usage. Under the indented heading<br />
‘‘Contents’’ we list all <strong>of</strong> the valid lower-level<br />
taxa included by each higher-level taxon; for<br />
example, the genera referred to a tribe, or the<br />
species referred to a genus. In lists <strong>of</strong><br />
congeneric species, those currently regarded<br />
as valid are in boldface, followed by an<br />
alphabetic list <strong>of</strong> available junior synonyms<br />
in parentheses; however, names <strong>of</strong> doubtful<br />
application (e.g., brasiliensis Liais, 1872,<br />
which might be a junior synonym <strong>of</strong> more<br />
than one currently recognized species <strong>of</strong><br />
Didelphis; see Cerqueira <strong>and</strong> Tribe, 2008)<br />
are not included.<br />
Under ‘‘Diagnosis’’ we list unique combinations<br />
<strong>of</strong> traits that distinguish members <strong>of</strong><br />
suprageneric taxa from other <strong>marsupials</strong>,<br />
whether or not such traits are unambiguously<br />
assignable by parsimony as relevant synapomorphies<br />
(listed in appendix 5). Under<br />
‘‘Morphological description’’ (for genera)<br />
we list all phenotypic descriptors that apply<br />
to included species; traits <strong>of</strong> particular<br />
importance for identification are indicated<br />
by italics. Ranges <strong>of</strong> metrical traits (e.g., 170–<br />
210 mm) are approximations that match or<br />
slightly exceed known minimal <strong>and</strong> maximal<br />
adult values that we believe to be reliable<br />
based on published <strong>and</strong> unpublished sources.<br />
Except as noted otherwise, all descriptions <strong>of</strong><br />
cranial traits are based on adult specimens,<br />
<strong>and</strong> all dental descriptions are based on<br />
unworn teeth.<br />
Under ‘‘Remarks’’ we discuss evidence for<br />
monophyly <strong>and</strong> other issues that relate to<br />
taxon recognition <strong>and</strong> rank. Although ranks<br />
are biologically arbitrary, they affect spelling<br />
<strong>and</strong> information retrieval in the prevailing<br />
Linnaean system, <strong>and</strong> they are regulated by<br />
widely accepted rules <strong>of</strong> biological nomenclature.<br />
The lack <strong>of</strong> species-level revisionary<br />
studies for most genera is also discussed<br />
under this heading.<br />
Family Didelphidae Gray, 1821<br />
CONTENTS: Glironiinae, Caluromyinae,<br />
Hyladelphinae, <strong>and</strong> Didelphinae.<br />
DIAGNOSIS: Didelphids can be distinguished<br />
unambiguously from other <strong>marsupials</strong><br />
by their s<strong>of</strong>t (nonspinous) body pelage;<br />
five subequal claw-bearing manual digits; five<br />
separate pedal digits (loosely connected by<br />
webbing in Chironectes), <strong>of</strong> which the hallux<br />
(dI) is large <strong>and</strong> opposable <strong>and</strong> the second<br />
(dII) bears an asymmetrical grooming claw;<br />
inguinal cloaca; prehensile tail (secondarily<br />
lacking external evidence <strong>of</strong> prehensility in<br />
some taxa); long nasal bones (extending<br />
anteriorly beyond the facial processes <strong>of</strong> the<br />
premaxillae <strong>and</strong> posteriorly between the<br />
lacrimals); unfenestrated rostrum; uninflected<br />
maxillary-jugal suture; distinct foramen<br />
rotundum (not confluent or sharing a common<br />
vestibule with the sphenorbital fissure);<br />
alisphenoid-parietal contact (except in Metachirus);<br />
large interparietal fused to the<br />
supraoccipital; complete posterolateral palatal<br />
foramina; deeply divided vomer that does<br />
not conceal the presphenoid or extend<br />
posteriorly into the mesopterygoid fossa;<br />
lack <strong>of</strong> midline contact between the left <strong>and</strong><br />
right pterygoids; unkeeled basisphenoid;<br />
well-developed tympanic process <strong>of</strong> the alisphenoid;<br />
well-developed rostral tympanic<br />
process <strong>of</strong> the petrosal; lack <strong>of</strong> a squamosal<br />
epitympanic sinus; laterally exposed ectotympanic;<br />
unspecialized malleus (with long<br />
sharply inflected neck, small orbicular<br />
apophysis, <strong>and</strong> well-developed lamina); distinct<br />
mastoid <strong>and</strong> paroccipital processes;<br />
unreduced dental complement <strong>of</strong> 50 teeth;<br />
nongliriform incisors; large milk premolars<br />
(except in Hyladelphys); tribosphenic upper<br />
molars, each with a continuous stylar shelf,<br />
uninterrupted centrocrista, <strong>and</strong> reduced or<br />
absent para- <strong>and</strong> metaconules; <strong>and</strong> tribosphenic<br />
lower molars, each with a welldeveloped<br />
anterior cingulid (notched for the<br />
hypoconulid <strong>of</strong> the preceding tooth), notched<br />
paracristids, <strong>and</strong> without any trace <strong>of</strong> a<br />
posterior cingulid.