AN ANNOTATED CHECKLIST OF RECENT OPOSSUMS (MAMMALIA: DIDELPHIDAE) - BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
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A N A NNOTATED C HECKLIST OF R ECENT
O POSSUMS ( M A MMALIA: D IDELPHIDAE)
RO BERT S. VOSS
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORYAN ANNOTATED CHECKLIST OF RECENT
OPOSSUMS (MAMMALIA: DIDELPHIDAE)
ROBERT S. VOSS
Division of Vertebrate Zoology (Mammalogy)
American Museum of Natural History
BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY
Number 455, 74 pp., 4 tables
Issued April 4, 2022
Copyright © American Museum of Natural History 2022 ISSN 0003-0090CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Categories of Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Systematic Accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Subfamily Caluromyinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Genus Caluromys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Subgenus Caluromys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Subgenus Mallodelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Genus Caluromysiops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Subfamily Glironiinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Genus Glironia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Subfamily Hyladelphinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Genus Hyladelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Subfamily Didelphinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Tribe Marmosini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Genus Marmosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Subgenus Eomarmosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Subgenus Exulomarmosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Subgenus Marmosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Subgenus Micoureus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Subgenus Stegomarmosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Genus Monodelphis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Subgenus Microdelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Subgenus Monodelphiops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Subgenus Monodelphis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Subgenus Mygalodelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Subgenus Pyrodelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Genus Tlacuatzin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
Tribe Metachirini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Genus Metachirus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Tribe Didelphini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Genus Chironectes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Genus Didelphis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Genus Lutreolina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Genus Philander . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37
Tribe Thylamyini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40
Genus Chacodelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Genus Cryptonanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Genus Gracilinanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42
Genus Lestodelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Genus Marmosops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Subgenus Marmosops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Subgenus Sciophanes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
2Genus Thylamys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Subgenus Thylamys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Subgenus Xerodelphys . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Appendix 1. On the Phylogenetic Classification of Sparassocynids . . . . . . . . . . . . . . . . . . . . . . . . 69
Appendix 2. Opossum Species Revalidated from Synonymies (1993–Present) . . . . . . . . . . . . . . 71
Appendix 3. Opossum Species Described as New since Gardner (1993) . . . . . . . . . . . . . . . . . . . 73
Appendix 4. Opossum Names Synonymized or Disused since 1993 . . . . . . . . . . . . . . . . . . . . . . . 74
34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
ABSTRACT
Living opossums (Didelphidae) comprise 125 species in 18 genera and 4 subfamilies. This syn-
opsis lists all the didelphid taxa (subfamilies, tribes, genera, subgenera, and species) currently rec-
ognised as valid, summarizes information about typification, synonyms, and geographic distributions,
remarks noteworthy recent changes in usage, and comments on still outstanding problems. A con-
cluding discussion rejects the notion that the almost twofold increase in opossum species from 1993
to the present is “taxonomic inflation” and considers the impact of new kinds of data and new
methods of data analysis on species delimitation.
INTRODUCTION in the recent literature. A concluding discussion
mentions some general trends in recent didel-
The need for an up-to-date taxonomic syn-
phid taxonomic research.
opsis of Recent opossums (Didelphidae) has
long been apparent. The last complete techni-
cal checklist was Gardner’s (2005), although an Categories of Information
important partial synopsis—restricted to the
Several categories of technical information
South American taxa—appeared just a few
provided in this synopsis merit brief explana-
years later (Gardner, 2008). Both were major
tions. Readers unfamiliar with taxonomic termi-
advances over earlier compilations, but these
nology may also wish to consult the helpful
useful publications were immediately followed
glossary section of the International Code of
by a decade-long phase of revisionary research
Zoological Nomenclature (ICZN, 1999).1
that substantially increased the number of spe-
Type species: Type species (of genera or sub-
cies recognized as valid. Subsequent lists that
genera) are nominal taxa, which are sometimes
appeared in volumes written for nonspecialist
junior synonyms of species currently known by
readers (Astúa, 2015; Voss and Jansa, 2021)
other names. In these accounts, type species are
necessarily omitted much relevant nomencla-
identified by their original binomial combina-
tural detail. Although no synopsis can plausi-
tions, with a parenthetical explanation if any is
bly claim to be the last word on the taxonomy
needed. For example, the type species of Mono
of this diverse and geographically widespread
delphis is Didelphis brachyuros Schreber, 1777,
group, most opossum genera have now
which corresponds to the species currently
received at least some critical scrutiny based
known as Monodelphis brevicaudata (Erxleben,
on firsthand examination of type material and
1777). Type species may have been originally
supporting analyses of morphological and
designated as such by the author of the genus-
molecular data. Therefore, a summary of taxo-
group name in question, or they may have been
nomic progress is timely.
designated by the same author or by a different
Here I list all the didelphid subfamilies, tribes,
author at some later date.
genera, subgenera, and species currently regarded
Type material and type locality: Except
as valid. I provide information about typification
as noted, information provided under this
and synonyms for each taxon and remark any
heading is consistent with that provided by the
nomenclatural issues not previously discussed by
original describer, although I have often
Gardner (2005, 2008). For supraspecific taxa, I
updated the spelling of geographic place names
additionally comment on published evidence for
and provided geographic coordinates if none
monophyly. For each species, I summarize infor-
mation about geographic distribution and cite 1 Available online (https://www.iczn.org/the-code/
relevant analyses of genetic and phenotypic data the-code-online/).2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 5 were originally given. Geographic coordinates Abbreviations provided without cited references are usually consistent with gazetteer entries in Gardner The following abbreviations are used for (2008) or with standard references such as the museum collections in which types and other ornithological gazetteers published by the specimens are preserved: Museum of Comparative Zoology (Harvard AMNH, American Museum of Natural History University) or those published by the U.S. gov- (New York, NY) ernment (e.g., by the U.S. Board on Geographic BMNH, Natural History Museum (London, UK) Names and the Defense Mapping Agency). CBF, Colección Boliviana de Fauna (La Paz, However, comments with supporting references Bolivia) are provided if the type material or the type CM, Carnegie Museum of Natural History (Pitts- locality differs from what has previously been burg, PA) reported in the literature. Common misconcep- CTUA, Colección Teriológica, Universidad de tions notwithstanding, the type locality is sim- Antioquia (Medellín, Colombia) ply the place where the name-bearing EBD, Estación Biológica Doñana (Sevilla, Spain) specimen(s)—holotype, lectotype, neotype, or EBRG, Estación Biológica Rancho Grande syntypes—was (or were) collected, so the type (Maracay, Venezuela) locality of a species based on material of ENCB, Escuela Nacional de Ciencias Biológicas, unknown origin is unknown, regardless of any Instituto Politécnico Nacional (Ciudad de “restrictions” subsequently proposed by authors México, Mexico) without definite knowledge of provenance. FMNH, Field Museum (Chicago, IL) Synonyms: Except as noted, only available ICN, Instituto de Ciencias Naturales, Universidad names based on Recent type material are listed Nacional de Colombia (Bogotá, Colombia) under this heading, including both objective syn- INPA, Instituto Nacional de Pesquizas da onyms (based on the same type material as the Amazônia (Manaus, Brazil) currently recognized valid name) and subjective IZH, Institut für Zoologie der Universität Zoolo- synonyms (based on different type material). For gische Sammlungen (Halle, Germany) conciseness, only the epithet (with author and LIVCM-D, World Museum (Liverpool, UK) date) is listed for species-group junior synonyms, LSUMZ, Louisiana State University Museum of which were often published in combination with Natural Science (Baton Rouge, LA) different generic names. In the event that a spe- MACN, Museo Argentino de Ciencias Naturales cies-group junior synonym was originally com- “Bernardino Rivadavia” (Buenos Aires, bined with a generic name that differed in gender Argentina) from the generic name in current use, the gender MBUCV, Museo de Biología, Universidad Cen- of the epithet has been changed to agree with tral de Venezuela (Caracas, Venezuela) current usage. Although I acknowledge the MCZ, Museum of Comparative Zoology, Har- potential usefulness of trinomial nomenclature vard University (Cambridge, MA) in several accounts, I do not formally recognize MHNLS, Museo de Historia Natural La Salle subspecies in this report. (Caracas, Venezuela) Distribution: Information provided under MHNN, Muséum d’Histoire Naturelle de this heading is concise if a published range map Neuchâtel (Neuchâtel, Switzerland) based on accurately identified material can be MN, Museu Nacional (Rio de Janeiro, Brazil) cited, or it may require a lengthy description if MNCN, Museo Nacional de Historia Natural no such map is available. (Madrid, Spain) Remarks: All other relevant information is MNHN, Muséum National d’Histoire Naturelle provided under this heading. (Paris, France)
6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
MPEG, Museu Paraense Emilio Goeldi (Belém, Remarks: See Voss and Jansa (2009) for an
Brazil) emended generic description. Generic mono-
MUSM, Museo de Historia Natural, Universidad phyly is strongly supported by phylogenetic anal-
Nacional Mayor de San Marcos (Lima, Peru) yses of multilocus sequence data (e.g., Voss and
MVZ, Museum of Vertebrate Zoology, Univer- Jansa, 2009; Amador and Giannini, 2016). Two
sity of California (Berkeley, CA) subgenera are currently recognized.
NMW, Naturhistorisches Museum Wien
(Vienna, Austria)
Subgenus Caluromys J.A. Allen, 1900
RMNH, Naturalis Biodiversity Center (Leiden,
the Netherlands) Type species: As for the genus.
ROM, Royal Ontario Museum (Toronto, Synonyms: None.
Canada) Remarks: This subgenus has long been
UFMT, Universidade Federal de Mato Grosso thought to contain only one species (but see
(Cuiabá, Brazil) remarks under Caluromys philander, below).
UFPA, Universidade Federal do Pará (Belém,
Brazil)
Caluromys (Caluromys) philander
UMMZ, University of Michigan Museum of
(Linnaeus, 1758)
Zoology (Ann Arbor, MI)
USNM, National Museum of Natural History Type material and type locality: BMNH
(Washington DC) 67.4.12.414, the holotype by monotypy, consists
ZMB, Museum für Naturkunde der Humboldt- of the fluid-preserved carcass and extracted skull
Universität zu Berlin (Berlin, Germany) of an adult female (Thomas, 1892; Jenkins and
ZMUC, Zoological Museum of the University of Knutson, 1983). The type locality is unknown,
Copenhagen (Copenhagen, Denmark) but it is often assumed to be Surinam (after
Thomas, 1911).
Synonyms: affinis Wagner, 1842; cajopolin
SYSTEMATIC ACCOUNTS Müller, 1776; cayopollin Schreber, 1777; cayopol
Subfamily Caluromyinae Reig et al., 1987 lin Kerr, 1792; dichurus Wagner, 1842; flavescens
Brongniart, 1792; leucurus Thomas, 1904; trini
Type genus: Caluromys J.A. Allen, 1900. tatis Thomas, 1894; venezuelae Thomas, 1903.
Remarks: See Voss and Jansa (2009) for a Distribution: Caluromys philander occurs in
morphological diagnosis. This subfamily includes lowland rainforest and dry forests in north-cen-
only Caluromys and Caluromysiops. An alterna- tral and eastern Venezuela; the Guianas; north-
tive concept of Caluromyinae that once included ern, central, and southeastern Brazil; and eastern
Glironia (e.g., in Gardner, 2008) is not demon- Bolivia (Gardner, 2008: map 2).
strably monophyletic (Jansa and Voss, 2000; Voss Remarks: As currently recognized, Caluro
and Jansa, 2009). mys philander is unrevised and may represent a
species complex, but published results are nei-
Genus Caluromys J.A. Allen, 1900 ther sufficient to distinguish valid taxa among
the synonyms listed above nor among geographic
Type species: Didelphis philander Linnaeus, populations currently lacking available names.
1758, by original designation.2 Although López-Fuster et al. (2008) suggested
Synonyms: None. that populations in Trinidad and northern Ven-
ezuela should be recognized as a distinct species,
2 Voss and Jansa (2009: table 15) incorrectly attributed the this inference was not supported by phylogenetic
type designation to Hershkovitz (1949). analyses of mtDNA sequence data reported by2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 7
Voss et al. (2019). By contrast, the latter authors Colombia and western Ecuador, but it is also
discovered that mtDNA sequences from eastern known from the Caribbean lowlands of north-
Bolivia were highly divergent from sequences western Colombia and from the interAndean
obtained elsewhere in the known range of C. phi valley of the Río Cauca (Gardner, 2008: map 2).3
lander; unfortunately, voucher material from Remarks: Phylogenetic analyses of mtDNA
eastern Bolivia has yet to be examined for pos- sequence data suggest that Caluromys derbianus
sibly diagnostic morphological traits. and C. lanatus are genetically divergent and
reciprocally monophyletic species (Voss et al.,
2019), but most 19th and early 20th century
Subgenus Mallodelphys Thomas, 1920
authors (e.g., Thomas, 1913) regarded these taxa
Type species: Didelphis laniger Desmarest, as conspecfic. Apparently, the first researcher to
1820 (= Caluromys lanatus; see below), by origi- treat them as valid species was Gilmore (in Bug-
nal designation. her et al., 1941), who mentioned ear color and
Synonyms: None. differences in the extent of caudal pelage as dis-
Remarks: The monophyly of this subgenus tinguishing characters. However, because there is
was only weakly supported by the mtDNA said to be some species overlap in caudal pelage
sequence data analyzed by Voss et al. (2019). traits (Gardner, 2008: 5), and because pelage
Two species are currently recognized. markings said to distinguish these species appear
to be similarly unreliable (Voss et al., 2019), only
ear coloration seems to be diagnostically useful
Caluromys (Mallodelphys) derbianus
(the pinnae are unpigmented in derbianus versus
(Waterhouse, 1841)
blackish or purple in lanatus). Although Bucher
Type material and type locality: LIVCM- and Hoffman (1980: 1) claimed that C. derbianus
D 194, the holotype by monotypy, is a female is the “largest species in the genus,” measured
specimen of unstated age, originally mounted for series of C. derbianus and C. lanatus exhibit
exhibition, but subsequently remade as a study broad morphometric overlap (R.S.V., personal
skin and skull (Thomas, 1913; Largen, 1985; obs.). Recent landmark-based multivariate mor-
Fisher, 2002). The type locality is unknown, but phometric analyses of Caluromys have either
it has often been assumed to be somewhere in failed to convincingly distinguish C. derbianus
the Río Cauca watershed of northern Colombia and C. lanatus from one another (López-Fuster
(e.g., by Allen, 1904; Thomas, 1913). et al., 2008) or have simply not addressed the
Synonyms: antioquiae Matschie, 1917; aztecus problem (Fonseca and Astúa, 2015).
Thomas, 1913; canus Matschie, 1917; centralis
Hollister, 1914; fervidus Thomas, 1913; guayanus 3 Insofar as I am aware, Caluromys derbianus and C. lana
Thomas, 1899; nauticus Thomas, 1913; pallidus tus are allopatric, but several problematic records of C. derbia
Thomas, 1899; pictus Thomas, 1913; pulcher nus mapped by Fonseca and Astúa (2015: fig. 2) merit
comment because they imply geographic range overlap. One
Matschie, 1917; pyrrhus Thomas, 1901; senex such Colombian record (their locality 69) is based on USNM
Thomas, 1913. specimens from the Río Raposo, which is in the Cauca valley,
Distribution: Caluromys derbianus is a not (as mapped) in the Cordillera Oriental. A second Colom-
bian record (locality 68) is based on FMNH specimens from
transAndean species that occurs in lowland rain- the upper Río Sinú, which drains the western slopes of the
forest, dry forest, and montane (“cloud”) forest Cordillera Occidental, not (as mapped) the northeastern
slopes of the Serranía de San Lucas. A third problematic
from Veracruz (Mexico) throughout most of record (locality 71), mapped in the Cordillera Oriental of
Central America to northwestern South America Ecuador, is based on AMNH 10058, a specimen from Costa
(Bucher and Hoffmann, 1980: fig. 3). In South Rica. A fourth anomalous record, from Tingo Maria in eastern
Peru (locality 75), is based on a specimen (LSUMZ 17681) that
America, the species is known from the Pacific is almost certainly misidentified (C. derbianus is not known
littoral and adjacent Andean foothills of western from Peru; Pacheco et al., 2020).8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
Some authors (e.g., Hall, 1981) have recog- nominotypical form) appear to differ morphologi-
nized valid subspecies of Caluromys derbianus, cally from western Amazonian and Cerrado mate-
but there is a striking lack of mtDNA sequence rial (Fonseca and Astúa, 2015; Voss et al., 2019), the
variation among samples collected in Costa Rica, taxonomic significance of such comparisons is
Panama, and Ecuador (Voss et al., 2019). Broader unclear in the absence of genetic data.
geographic sampling of genetic variation is
needed to determine whether any of the nominal
Genus Caluromysiops Sanborn, 1951
taxa herein treated as synonyms of C. derbianus
represent evolutionarily significant units worthy Type species: Caluromysiops irrupta Sanborn,
of taxonomic recognition. 1951, by original designation.
Synonyms: None.
Remarks: For an emended generic description,
Caluromys (Mallodelphys) lanatus (Olfers, 1818)
see Voss and Jansa (2009), who also discussed the
Type material and type locality: MNCN- status of Caluromysiops as a valid genus (distinct
M2630, the holotype by monotypy, consists of from Caluromys), a once controversial topic. Only
the skin and skull of a juvenile male collected at a single species is currently recognized.
Caazapá (26.15° S, 56.40° W), Caazapá depart-
ment, Paraguay (Voss et al., 2009).
Caluromysiops irrupta Sanborn, 1951
Synonyms: bartletti Matschie, 1917; cahyensis
Matschie, 1917; cicur Bangs, 1898; hemiurus Type material and type locality: FMNH
Miranda-Ribeiro, 1936; jivaro Thomas, 1913; 68336, the holotype by original designation, con-
juninensis Matschie, 1917; lanigera Desmarest, sists of the skin and skull of a juvenile male col-
1820; meridensis Matschie, 1917; modestus lected at Quincemil (13.22° S, 70.70° W; 680 m),
Miranda-Ribeiro, 1936; nattereri Matschie, 1917; Cusco department, Peru.
ochropus Wagner, 1842; ornatus Tschudi, 1845; Synonyms: None.
vitalinus Miranda-Ribeiro, 1936. Distribution: Caluromysiops irrupta is
Distribution: Caluromys lanatus occurs in restricted to lowland Amazonia, where it is
rainforest, dry forest, and premontane forest known from scattered localities in Brazil (Mato
from northern Colombia to eastern Bolivia, east- Grosso, Rondônia), Colombia (Amazonas), and
ern Paraguay, and southeastern Brazil (Fonseca Peru (Loreto, Madre de Dios) (Santori et al.,
and Astúa, 2015: fig. 3). Most records are from 2016: fig. 2).
Amazonia, the Cerrado, and the lower slopes of Remarks: See Voss and Jansa (2009) for illustra-
the tropical Andes, but the species is also known tions, analyses of phylogenetic relationships and
to occur in the subtropical Paraguayan extension qualitative morphological descriptors. The only
of the Atlantic Forest (Owen et al., 2018). published morphometric data for Caluromysiops
Remarks: To date, mtDNA sequence data for irrupta are still those in Izor and Pine (1987).
Caluromys lanatus are available only from western Despite several reported sightings and one recent
Amazonian and Cerrado samples, which exhibit capture (summarized by Santori et al., 2016), no
little genetic divergence and a striking absence of new morphological specimens or genetic samples
phylogeographic structure (Voss et al., 2019: fig. 7); have been obtained for several decades.
these results clearly support Fonseca and Astúa’s
(2015) suggestion that just one taxon (for which
Subfamily Glironiinae Voss and Jansa, 2009
ochropus is the oldest available name if any trino-
mial classification were warranted) occurs through- Type genus: Glironia Thomas, 1912.
out these regions. Although specimens from eastern Remarks: See Voss and Jansa (2009) for a
Paraguay and southeastern Brazil (representing the morphological diagnosis. Glironia was formerly2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 9
placed in the subfamily Caluromyinae (e.g., by b amplified from the holotypes of aequatorialis
Gardner, 2008), but morphological and genetic and criniger (from north of the Amazon in east-
support for Caluromyinae sensu lato is weak or ern Ecuador and northeastern Peru, respectively)
nonexistent (Jansa and Voss, 2000; Voss and but a much larger distance (ca. 6%) between
Jansa, 2009; Amador and Giannini, 2016). those sequences and one from south of the Ama-
zon in western Brazil. Unfortunately, no sequence
data are currently available from eastern Amazo-
Genus Glironia Thomas, 1912
nia. Measurements from five western Amazonian
Type species: Glironia venusta Thomas, 1912, specimens were tabulated by Voss et al. (2019).
by original designation.
Synonyms: None.
Subfamily Hyladelphinae Voss and Jansa, 2009
Remarks: See Voss and Jansa (2009) for a
detailed morphological description of Glironia, Type genus: Hyladelphys Voss et al., 2001.
which exhibits several morphological traits that Remarks: See Voss and Jansa (2009) for a mor-
are unknown among other opossums. Only a phological diagnosis. This taxon represents a very
single species is currently recognized. long branch that is consistently recovered as the
sister lineage of Didelphinae in phylogenetic analy-
ses of multilocus sequence datasets (e.g., by Voss
Glironia venusta Thomas, 1912
and Jansa, 2009; Amador and Giannini, 2016).
Type material and type locality: BMNH
12.1.15.7, the holotype by original designation,
Genus Hyladelphys Voss et al., 2001
consists of the skin and skull of an adult male
collected at Pozuzo (10.07° S, 75.53° W; 800 m), Type species: Gracilinanus kalinowskii Hersh-
Pasco department, Peru. kovitz, 1992, by original designation.
Synonyms: aequatorialis Anthony, 1926; cri Synonyms: None.
niger Anthony, 1926. Remarks: Emended morphological descrip-
Distribution: Glironia venusta has been col- tions were provided by Jansa and Voss (2005)
lected or observed at widely scattered rainforest and Voss and Jansa (2009). Only one species is
localities throughout much of Amazonia, but it currently recognized.
is also known to occur in tropical dry forest in
eastern Bolivia (Santa Cruz) and southwestern
Hyladelphys kalinowskii (Hershkovitz, 1992)
Brazil (Mato Grosso). Noteworthy recent exten-
sions of the geographic range as mapped by Díaz Type material: FMNH 89991, the holotype
and Willig (2004: fig. 1) include records from by original designation, consists of the skin and
southeastern Colombia (Montenegro and skull of an adult female collected at Hacienda
Restrepo, 2018), eastern Brazil (Ardente et al., Cadena (13.33° S, 70.77° W; 890 m), Cusco
2013), and French Guiana (Sant and Catzeflis, department, Peru.
2018). Formerly thought to be a lowland species, Synonyms: None.
G. venusta is now known to occur at elevations Distribution: As currently understood, Hyl
>1500 m in the Andes (Arguero et al., 2017). adelphys kalinowskii is known from eastern Peru
Remarks: The only molecular data available (Cusco, Junín, Loreto), northern Brazil (Amazo-
to assess the taxonomic status of nominal taxa nas, near Manaus), southern Guyana, and French
currently regarded as synonyms of Glironia Guiana (Gardner 2008: map 18).
venusta were analyzed by Voss et al. (2019), who Remarks: The possibility that multiple cryp-
reported a trivial sequence difference (0.2%, tic taxa might be represented among the material
uncorrected) between fragments of cytochrome currently referred to this species was discussed10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
by Jansa and Voss (2005). Measurement data Genus Marmosa Gray, 1821
from recently collected specimens are in Catze-
Type species: Didelphis murina Linnaeus,
flis (2017) and Voss et al. (2019).
1758, by monotypy.
Synonyms: Asagis Gloger, 1841; Grymaeomys
Subfamily Didelphinae Gray, 1821 Burmeister, 1854; Cuica Liais, 1872; Grayium
Kretzoi and Kretzoi, 2000.
Type genus: Didelphis Linnaeus, 1758.
Remarks: For an emended generic descrip-
Synonyms: Chironectinae Hershkovitz, 1997;
tion see Voss et al. (2014), who additionally rec-
Lutreolininae Hershkovitz, 1997.
ognized the five subgenera listed below. Although
Remarks: See Voss and Jansa (2009: 100) for
Micoureus was formerly regarded as a separate
a morphological diagnosis. Monophyly of the
genus (e.g., by Gardner and Creighton, 2008a),
nominotypical subfamily has been consistently
the monophyly of Marmosa—as recognized
and strongly supported by phylogenetic analyses
herein, including Micoureus as a subgenus—has
of multilocus sequence datasets (e.g., Voss and
been consistently and strongly supported by phy-
Jansa, 2009; Amador and Giannini, 2016).
logenetic analyses of multilocus sequence data
sets (e.g., by Voss and Jansa, 2009; Voss et al.,
Tribe Marmosini Hershkovitz, 1992 2014; Amador and Giannini, 2016).
Type genus: Marmosa Gray, 1821.
Remarks: The monophyly of Marmosini as
Subgenus Eomarmosa Voss et al., 2014
constituted herein—including Marmosa, Monodel
phis, and Tlacuatzin, but not the other genera Type species: Marmosa rubra Tate, 1931, by
referred to “Marmosidae” by Hershkovitz (1992)— original designation.
has been consistently and strongly supported by all Synonyms: None.
relevant phylogenetic analyses of multilocus Remarks: Only a single species is currently
sequence datasets (e.g., Voss and Jansa, 2009; May- recognized.
Collado et al., 2015; Vilela et al., 2015; Amador and
Giannini, 2016). Recently, however, Beck and
Marmosa (Eomarmosa) rubra Tate, 1931
Taglioretti (2020) suggested that Marmosini be
restricted to include just Marmosa and Tlacuatzin Type material and type locality: AMNH
(or that Marmosini be restricted to include just 71973, the holotype by original designation, con-
Marmosa with a new tribe for Tlacuatzin), and that sists of the skin and skull of an adult female col-
Monodelphis be placed in a separate tribe (Mono- lected at the mouth of the Río Curaray (ca. 2.37°
delphini). These proposed changes were prompted S, 74.08° W; ca. 200 m), Loreto, Peru. Tate (1931,
by phylogenetic analyses that recovered two highly 1933) thought that the type was collected in
specialized fossil taxa in a clade with Monodelphis. Ecuador, but “Boca Río Curaray” (as this locality
However, as explained elsewhere (appendix 1), was originally recorded by the collectors) is well
Beck and Taglioretti’s phylogenetic results are suf- within the internationally recognized borders of
ficiently open to question that it seems unnecessary Peru (Wiley, 2010).
to disrupt the current classification. Restricting Synonyms: None.
Marmosini as they propose would leave the Distribution: Marmosa rubra is known
robustly supported clade that includes Marmosa, from just a few localities in the Amazonian low-
Tlacuatzin, and Monodelphis without a name, and lands of southeastern Colombia (Putumayo),
their alternative tribal usage lacks any compensa- eastern Ecuador (Napo, Orellana, Pastaza, and
tory advantage for communicating phylogenetic Sucumbios), and eastern Peru (Loreto, and
relationships among Recent taxa. Madre de Dios) (Rossi et al., 2010: fig. 30). Addi-2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 11
tional Peruvian records were summarized by 2010) provided compelling evidence that these
Pacheco et al. (2020). are, in fact, genetically distinct species.
Remarks: See Rossi et al. (2010) for an Although Marmosa regina is a senior synonym
emended morphological description, illustra- of M. isthmica, the former name has long been
tions, measurement data, and comparisons with misapplied to species in the subgenus Micoureus.
congeneric species. To preserve long-standing binomial usage of M.
isthmica, Voss and Giarla (2020a) petitioned the
International Commission on Zoological Nomen-
Subgenus Exulomarmosa Voss et al., 2014
clature to suppress usage of M. regina. A ruling
Type species: Marmosa robinsoni Bangs, from the Commission is pending.
1898, by original designation.
Synonyms: None.
Marmosa (Exulomarmosa) mexicana
Remarks: Subgeneric monophyly has been
Merriam, 1897
consistently and strongly supported by phylo-
genetic analyses of multilocus sequence data Type material and type locality: USNM
sets (e.g., by Voss et al., 2014; Amador and 71526, the holotype by original designation, con-
Giannini, 2016). Six species are currently rec- sists of the skin and skull of an adult male col-
ognized as valid. lected at “Juquila” (= Santa Catarina Juquila:
16.23° N, 97.30° S; 1500 m), Oaxaca state,
Mexico.
Marmosa (Exulomarmosa) isthmica
Synonyms: mayensis Osgood, 1913; ruatanica
Goldman, 1912
Goldman, 1911; savannarum Goldman, 1917.
Type material and type locality: USNM Distribution: As currently recognized (see
170969, the holotype by original designation, Remarks), Marmosa mexicana is known from
consists of the skin and skull of an adult male rainforested and dry-forested localities from
collected on the Río Indio (9.25° N, 79.98° W; at northern Mexico (Tamaulipas) southward and
sea level), a tributary of the lower Río Chagres eastward throughout Central America to eastern
near Gatún, Colón province, Panama. Panama (Rossi et al., 2010: fig. 17).
Synonyms: mimetra Thomas, 1921; regina Remarks: See Rossi et al. (2010) for an
Thomas, 1898 (suppressed; see Remarks). emended description, tabulated measurement
Distribution: Marmosa isthmica is known data, and morphological comparisons with conge-
from humid lowland and lower-montane locali- neric species. As reported by Gutiérrez et al.
ties (usually below 1700 m) in Panama, western (2010), mtDNA sequences from specimens that fit
Colombia (including the interAndean Cauca and the morphological description of Marmosa mexi
Magdalena valleys), and western Ecuador (Rossi cana form two highly divergent haplogroups.
et al., 2010: fig. 22). Some authors (e.g., Ramírez-Pulido et al., 2014)
Remarks: Originally described as a full spe- now recognize M. mayensis as a distinct species—
cies, Marmosa isthmica was treated for many presumably based on Gutiérrez et al.’s sequencing
years as a subspecies or synonym of M. robinsoni results—but diagnostic morphological characters
(after Hershkovitz, 1951). Current usage follows are elusive, and no sequence data are available
Rossi et al. (2010), who demonstrated that M. from holotypes or topotypes to confidently assign
isthmica and M. robinsoni are morphologically names to either mtDNA clade. Marmosa mexi
distinct, known to occur in sympatry, and usu- cana was formerly thought to be conspecific with
ally occupy different habitats (M. isthmica in M. zeledoni (e.g., by Tate, 1933), but evidence that
rainforest, M. robinsoni in dry forest). A subse- these are separate species was summarized by
quent mtDNA sequencing study (Gutiérrez et al., Rossi et al. (2010) and Gutiérrez et al (2010).12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
Marmosa (Exulomarmosa) robinsoni such that specimens cannot be assigned to one
Bangs, 1898 or the other without DNA sequence data.
Type material and type locality: MCZ
Marmosa (Exulomarmosa) simonsi
B7749, the holotype by original designation, con-
Thomas, 1899
sists of the skin and skull of an adult male col-
lected at El Valle de Espírito Santo (10.98° N, Type material and type locality: BMNH
63.87° W; ca. 200 m) on Isla Margarita, Nueva 99.8.1.20, the holotype by original designation,
Esparta state, Venezuela (Rossi et al., 2010). consists of the skin and skull of an adult male
Synonyms: casta Thomas, 1911; chapmani collected at Puná (2.73° S, 79.92° W; near sea
Allen, 1900; fulviventer Bangs, 1901; grenadae level), on Isla Puná, Guayas province, Ecuador.
Thomas, 1911; luridivolta Goodwin, 1961; mitis Synonyms: None.
Bangs, 1898; nesaea Thomas, 1911; pallidiventris Distribution: Marmosa simonsi occurs in
Osgood, 1912. mangroves and dry forests of the Pacific lowlands
Distribution: Marmosa robinsoni occurs and Andean foothills of western Ecuador and
primarily in dry forests, but also occasionally in northwestern Peru (Rossi et al., 2010: fig. 28).
other habitats from western Panama to Colom- Remarks: See Rossi et al. (2010) for an
bia, northern Venezuela, and several adjacent emended description, tabulated measurement
continental-shelf islands (including Isla Mar- data, and morphological comparisons with con-
garita, Trinidad, and Tobago); the species is also generic species. Marmosa simonsi was long con-
known from Grenada, a Caribbean island that sidered a synonym or subspecies of M. robinsoni
is not on the continental shelf (Rossi et al., (e.g., by Herskovitz, 1951; Creighton and Gard-
2010: fig. 25). ner, 2008a), but Rossi et al. (2010) and Gutiérrez
Remarks: See Rossi et al. (2010) for an et al. (2010) showed that these taxa are morpho-
emended description, tabulated measurement logically and genetically distinct.
data, and morphological comparisons with
congeneric species. Several taxa that were for-
Marmosa (Exulomarmosa) xerophila Handley
merly treated as subspecies or synonyms of
and Gordon, 1979
Marmosa robinsoni (e.g., by Hershkovitz, 1951;
Hall, 1981; Creighton and Gardner, 2008a) are Type material and type locality: USNM
now recognized as valid species (M. isthmica, 443819, the holotype by original designation,
M. simonsi) or have been relegated to the syn- consists of the skin and skull of an adult male
onymies of other species. Even in its currently collected at La Isla (ca. 11.63° N, 71.83° S; near
restricted sense (Rossi et al., 2010), however, sea level), Guajira department, Colombia.
M. robinsoni is a geographically and ecologi- Synonyms: None.
cally widespread species that includes numer- Distribution: Marmosa xerophila occurs in
ous nominal taxa as subjective synonyms. desert thornscrub along the arid Caribbean coast
Analyses of DNA sequence data (Gutiérrez et of northeastern Colombia and northwestern
al., 2014a) have shown that geographic popu- Venezuela (Rossi et al., 2010: fig. 26). Additional
lations of this species sort out into two strongly specimen records and comments on the ecogeo-
supported phylogroups: an eastern clade for graphic distribution of this species were provided
which the oldest available trinomen would be by Gutiérrez et al. (2014b).
M. robinsoni robinsoni, and a western clade for Remarks: See Rossi et al. (2010) for an
which the oldest trinomen would be M. r. emended morphological description, tabulated
mitis. Unfortunately, these putative subspecies measurement data, and morphological compari-
appear to be phenotypically indistinguishable, sons with congeneric species.2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 13
Marmosa (Exulomarmosa) zeledoni Natterer specimen of this species from the Rio
Goldman, 1911 Madeira. Tate (1933: 101) referred to this speci-
men as “Vienna 195,” and called it a “co-type.” It
Type material and type locality: USNM is now cataloged as NMW B-2610 and consists
12885, the holotype by original designation, con- of the skin and skull of an adult male (F. Zachos,
sists of the skin and skull of an adult male col- in litt., 2 December 2021).
lected at Navarro (9.82° N, 83.87° S; ca. 840 m), Synonyms: madeirensis Cabrera, 1913 (an
Cartago province, Costa Rica. invalid replacement name; see Remarks); musi
Synonyms: None. cola Osgood, 1913; quichua Thomas, 1899.
Distribution: Marmosa zeledoni is known Distribution: As recognized by Rossi (2005;
from widely scattered localities, mostly in pre- see Remarks), Marmosa macrotarsus is an Ama-
montane or montane rainforest (to 2200 m) but zonian species that occurs south of the Amazon
sometimes in very wet lowland forests, from and west of the Tapajós; reported collection
north-central Nicaragua southward through Costa localities are in rainforested lowlands and foot-
Rica and Panama to western Colombia and north- hills (below 1900 m) in Peru, Brazil, and Bolivia
western Ecuador (Rossi et al., 2010: fig. 21). (Rossi, 2005: fig. 62).
Remarks: See Rossi et al. (2010) for an Remarks: Marmosa macrotarsus was ranked
emended description, tabulated measurement as a subspecies of M. murina by Tate (1933) who,
data, and morphological comparisons with con- however, used Cabrera’s (1913) replacement
generic species. Marmosa zeledoni was long name for this taxon, and it was treated as a syn-
regarded as a synonym or subspecies of M. mexi onym of M. murina by Creighton and Gardner
cana (e.g., by Tate, 1933), but these are morpho- (2008a). Current recognition of M. macrotarsus
logically and genetically distinct taxa that are as a valid species follows Rossi (2005), who pro-
known to occur sympatrically at several localities vided a morphological description, tabulated
(Rossi et al., 2010; Gutiérrez et al., 2010). measurement data, and carried out morphomet-
ric comparisons with other congeners. Subse-
Subgenus Marmosa Gray, 1821 quently, Gutiérrez et al. (2010) found that
specimens geographically assignable to M. mac
Type species: As for the genus. rotarsus (sensu Rossi, 2005) are highly divergent
Synonyms: As for the genus. from M. murina in cytochrome b sequence com-
Remarks: Monophyly of the nominotypical parisons (>9%, uncorrected), and phylogenetic
subgenus has been consistently and strongly sup- analyses of multilocus sequence data later
ported by phylogenetic analyses of multilocus showed that these species may not even be sister
sequence datasets (e.g., by Voss et al., 2014; taxa (Voss et al., 2014). Nevertheless, the mor-
Amador and Giannini, 2016). Four species are phological criteria by which specimens of M.
currently recognized as valid. macrotarsus and M. murina can be distinguished
remain to be convincingly documented. Voss et
al. (2019) discussed morphological comparisons
Marmosa (Marmosa) macrotarsus
between M. macrotarsus and M. waterhousei,
(Wagner, 1842)
another taxon formerly ranked as a subspecies of
Type material and type locality: No type M. murina (see below).
was designated in the original description, Cabrera (1913) believed that Didelphys mac
although Wagner (1847) mentioned two speci- rotarsus Wagner, 1842, was preoccupied by D.
mens collected by Johann Natterer on the Rio macrotarsos Schreber, 1778 (a tarsier), and this
Madeira that have subsequently been regarded as opinion was endorsed by Creighton and Gardner
syntypes. Pelzeln (1883), however, listed only one (2008a). However, the one-letter difference in14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
spelling is sufficient to prevent homonymy 2014) suggest that geographic populations cur-
(Rossi, 2005). Therefore, although Cabrera’s rently recognized as Marmosa murina include
replacement name is available in the sense of the four strongly supported phylogroups that
Code (ICZN, 1999: Article 12.2.3), it is an objec- might reasonably be recognized as subspecies:
tive junior synonym of Wagner’s macrotarsus and (1) mainland populations north of the Ama-
therefore invalid. zon, for which the oldest available trinomen
would be M. m. murina; (2) an insular popula-
tion on Tobago, which could be called M. m.
Marmosa (Marmosa) murina (Linnaeus, 1758)
tobagi; (3) populations in southeastern Ama-
Type material and type locality: BMNH zonia (east of the Tapajós and south of the
67.4.12.542, the lectotype (designated by Husson, Amazon), for which M. m. parata would seem
1978), consists of a fluid-preserved female speci- to be the appropriate trinomen; and (4) popu-
men from which the skull has been extracted and lations in the Atlantic Forest of southeastern
lost (Voss et al., 2001). Rossi (2005: 95) thought Brazil, which could be referred to M. m.
that BMNH 67.4.12.541 (a male) was the lecto- moreirae. Of these nominal taxa, however, only
type, citing Thomas (1892) as having so desig- tobagi appears to be morphologically diagnos-
nated that specimen, but Thomas merely able from the others (Rossi, 2005). The logic of
identified two probable syntypes without choos- treating tobagi as a subspecies of M. murina
ing either as the unique name-bearer. Jenkins rather than as a valid species was briefly dis-
and Knutson (1983) also appear to have been cussed by Voss et al. (2014), whose phyloge-
unaware of Husson’s lectotype designation. The netic results implied that this phenotypically
type locality is unknown, but it is often assumed divergent insular form is closely related to
to be Surinam (after Thomas, 1911). adjacent mainland populations.
Synonyms: chloe Thomas, 1907; dorsigera
Linnaeus, 17584; duidae Tate, 1931; klagesi J.A.
Marmosa (Marmosa) tyleriana Tate, 1931
Allen, 1900; meridionalis Miranda-Ribeiro, 1936;
moreirae Miranda-Ribeiro, 1936; musculus Caba- Type material and type locality: AMNH
nis, 1848; parata Thomas, 1911; roraimae Tate, 76983, the holotype by original designation, con-
1931; tobagi Thomas, 1911. sists of the skin and skull of an adult female col-
Distribution: As currently understood lected at an expeditionary locality known as
(Voss et al., 2014), Marmosa murina is known Central Camp (ca. 3.38° N, 65.58° W; 1400 m)
from northwestern Venezuela and eastern on the Mt. Duida massif, Amazonas state,
Colombia eastward and southward throughout Venezuela.
the Guianas to Brazil; in Brazil, the species is Synonyms: phelpsi Tate, 1939.
known from Amazonia (east of the Rio Negro Distribution: This species is known from
and the Tapajós), the Cerrado, and the Atlantic just a few localities in the Guiana highlands of
Forest. Marmosa murina is also known from southern Venezuela at elevations from 1300 to
Tobago, but not from Trinidad. Rossi (2005: 2100 m (Creighton and Gardner, 2008a: map 24).
fig. 56) mapped the joint distribution of M. Remarks: Marmosa tyleriana is the sister
murina and M. tobagi, which he regarded as taxon of a clade that contains all the other spe-
distinct species. cies in the subgenus Marmosa (Voss et al.,
Remarks: Analyses of cytochrome b 2014). Rossi (2005) provided a detailed mor-
sequence data (Faria et al., 2013a; Voss et al., phological description and tabulated measure-
ment data of the specimens he examined, but
4 For the priority of murina Linnaeus, 1758, over dorsigera additional measurement data were reported by
Linnaeus, 1758, see Husson (1978: 22). Ochoa (1985).2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 15
Marmosa (Marmosa) waterhousei DNA sequence data (Gutiérrez et al., 2010; Voss
(Tomes, 1860) et al., 2014), morphological distinctions between
some pairs of species remain problematic. Appar-
Type material and type locality: BMNH
ently, only measurement data seem to consis-
7.1.1.215, the lectotype (designated by Thomas,
tently distinguish specimens of M. waterhousei
1921a), consists of a skull, said to be that of an
from specimens of M. macrotarsus (see Voss et
adult female, collected at Gualaquiza (3.40° S,
al., 2019).
78.55° W; 914 m), Morona-Santiago province,
This name was originally spelled waterhousii,
Ecuador. The rest of the specimen, originally
but most subsequent authors have spelled it
preserved in fluid, has been lost.5 Jenkins and
waterhousei. The latter spelling would appear to
Knutson (1983) referred to this specimen as the
have been an incorrect subsequent spelling (in
holotype, but Tomes’ (1860) description was
the special sense of the Code) when it was first
based on an adult and an unspecified number
adopted by Thomas (1888a), but waterhousei is
of young individuals, all of which were, in
the spelling in prevailing usage today and should
effect, syntypes.
be maintained (ICZN, 1999: Article 33.3.1).
Synonyms: bombascarae Anthony, 1922;
maranii Thomas, 1924.
Distribution: As currently recognized, Subgenus Micoureus Lesson, 1842
Marmosa waterhousei occurs in the lowlands
Type species: Didelphis cinerea Temminck,
and adjacent Andean foothills of southeastern
1824 (= Marmosa paraguayana; see below), by
Colombia, eastern Ecuador, northeastern Peru
subsequent designation (Thomas, 1888a)
(north of the Amazon), and a few scattered
Synonyms: None.
localities in the Andes of northern Colombia
Remarks: Micoureus was formerly ranked as
and western Venezuela (Gutiérrez et al., 2011:
a genus (e.g., by Gardner and Creighton, 2008a),
fig. 2). A recently published Peruvian record
but this clade has been consistently recovered
from south of the Amazon (in Junín depart-
nested within Marmosa by phylogenetic analyses
ment; Pacheco et al., 2020) merits phenotypic
of multilocus sequence datasets (Voss and Jansa,
and genetic confirmation.
2009; Voss et al., 2014; Amador and Giannini,
Remarks: Marmosa waterhousei was ranked
2016). Thirteen species are currently recognized,
as a subspecies of M. murina by Tate (1933), and
most of which can be sorted into one or another
it was treated as a synonym of M. murina by
of several groups based on robustly supported
Creighton and Gardner (2008a). Current recog-
phylogenetic relationships (table 1).
nition of M. waterhousei as a valid species fol-
lows Rossi (2005), who provided a morphological
description, tabulated measurement data, and Marmosa (Micoureus) adleri Voss et al., 2021
carried out morphometric comparisons with
Type material and type locality: AMNH
other congeners. Despite compelling support for
272942, the holotype by original designation,
currently recognized species limits in the subge-
consists of the skin and skull of an adult female
nus Marmosa from phylogenetic analyses of
collected 1 km north of the Río Mendoza on
Pipeline Road (9.17° N, 79.75° W; 120 m), Parque
5 Tomes’ (1860) assertion that the fluid-preserved type of
Nacional Soberanía, Colón province, Panama.
Marmosa waterhousei had a pouch is impossible to reconcile
with its skull, which clearly belongs to a species in the pouch-
Synonyms: None.
less nominotypical subgenus of Marmosa. However, the appli- Distribution: Marmosa adleri is currently
cation of this name can only be based on what remains of the known only from Panama, where it has been col-
specimen, and on the type locality (eastern Ecuador), where
only a single species of the nominotypical subgenus is known lected from near the Costa Rican border to the
to occur. Colombian frontier (Voss et al., 2021: fig. 2).16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 455
TABLE 1 Synonyms: None.
Species-group Assignments in the
Distribution: As currently understood (see
Subgenus Micoureus of Marmosa Remarks, below), Marmosa alstoni is only known
from a handful of localities in the central high-
Alstoni Groupa lands of Costa Rica (Voss et al., 2021: fig. 2).
M. adleri Remarks: Marmosa alstoni was once thought
M. alstoni to range from Belize to Colombia (Tate, 1933;
M. nicaraguae Hall, 1981; Gardner and Creighton, 2008a) and
Perplexa Groupb
to include M. nicaraguae as a synonym, but a
recent revision of the Alstoni Group of Marmosa
M. jansae
restricted the application of this name to speci-
M. perplexa
mens collected in the central highlands of Costa
Phaea Groupb Rica (Voss et al., 2021).
M. constantiae
M. demerarae
Marmosa (Micoureus) constantiae Thomas, 1904
M. phaea
Rapposa Groupc Type material and type locality:
M. parda BMNH 3.7.7.157, the holotype by original des-
M. rapposa
ignation, consists of the skin and skull of an
adult male collected at “Chapada” (= Santa
M. rutteri
Ana de Chapada: 15.43° S, 55.75° W; 800 m),
Unaffiliated species
Mato Grosso state, Brazil.
M. germana Synonyms: domina Thomas, 1920; mapirien
M. paraguayana sis Tate, 1931.
a After Voss et al. (2021). Distribution: As currently recognized (see
b New (see text). Remarks), Marmosa constantiae occurs from the
c After Voss et al. (2020).
foothills of the Andes (below about 1100 m) in
eastern Peru and eastern Bolivia eastward across
Remarks: Marmosa adleri is the sister species Amazonia and the Cerrado to central Brazil;
of M. alstoni and a member of the Alstoni Group mtDNA sequencing results (see Remarks) sug-
(Voss et al., 2021). Specimens of M. adleri were gest that the range of this species does not extend
previously misidentified as M. alstoni (e.g., by Pat- north of the Amazon nor east of the Xingu (Silva
ton et al., 2000; Voss et al., 2020) or as M. phaea et al., 2019: fig. 5; Voss et al., 2020: fig. 2).
(e.g., by Handley, 1966; Emmons, 1997), but they Remarks: The name Marmosa constantiae
are unmistakably distinct from both. Illustrations, has long been misapplied to a superficially sim-
a morphological description, measurement data, ilar congener, M. rapposa, that also occurs in
and comparisons with closely related congeners Mato Grosso and eastern Bolivia (see below).
were provided by Voss et al. (2021). Previous reports of M. constantiae from Argen-
tina (Flores et al., 2007) and Paraguay (de la
Sancha et al., 2012; Smith and Owen, 2015)
Marmosa (Micoureus) alstoni (J.A. Allen, 1900)
likewise appear to have been based on speci-
Type material and type locality: AMNH mens of M. rapposa. As recognized by Silva et
11790/16210, the holotype by original designa- al. (2019) and Voss et al. (2020), M. constantiae
tion, consists of the skin and skull of an adult is geographically variable in coloration:
male collected at Tres Ríos (9.90° N, 83.98° W; whereas Cerrado populations have pale dorsal
1219 m), Cartago province, Costa Rica. fur, broadly self-yellow underparts, and parti-2022 VOSS: CHECKLIST OF RECENT OPOSSUMS 17
colored (white-tipped) tails, rainforest popula- of specimens from south of the Amazon has like-
tions are darker dorsally and have mostly wise failed to find compelling evidence that limae
gray-based ventral fur and all-dark tails (Voss is a distinct species.
et al., 2019). The rainforest phenotype of M.
constantiae is difficult to distinguish morpho-
Marmosa (Micoureus) germana Thomas, 1904
logically from M. germana (a distantly related
congener that occurs north of the Amazon; Type material and type locality: BMNH
Voss and Giarla, 2021), so currently recognized 80.5.6.77, the holotype by original designation,
range limits are based, in part, on sequencing consists of the skin and skull of a subadult female
results rather than examined specimens. Phy- collected at Sarayacu (1.73° S, 77.48° W; ca. 700
logenetic analyses of multilocus sequence data m) on the Río Bobonaza, Pastaza province,
recover M. constantiae and M. demerarae as Ecuador.
sister taxa (Voss et al., 2020), and comparisons Synonyms: None.
of sequenced specimens suggest that these taxa Distribution: Marmosa germana has been
are morphologically diagnosable (Silva et al., collected at scattered localities in the Amazonian
2019). lowlands of southeastern Colombia (Caquetá),
eastern Ecuador (Orellana, Pastaza), and north-
eastern Peru (Loreto, north of the Amazon)
Marmosa (Micoureus) demerarae Thomas, 1905
(Voss and Giarla, 2021: fig. 2).
Type material and type locality: BMNH Remarks: This species was long considered a
5.11.1.25, the holotype by original designation, subspecies or synonym of Marmosa regina (e.g.,
consists of the skin and skull of an adult female by Gardner and Creighton, 2008a). Diagnostic
collected at “Comackka” (= Takama: 5.57° N, morphological characters, taxonomic compari-
57.92° W; ca. 100 m), East Demerara-West Coast sons, and phylogenetic relationships were dis-
Berbice, Guyana. cussed by Voss and Giarla (2021). As currently
Synonyms: arenticola Tate, 1931; esmeraldae recognized, this species contains at least two
Tate, 1931; limae Thomas, 1920; meridae Tate, cytochrome b haplogroups that might represent
1931; pfrimeri Miranda-Ribeiro, 1936. distinct taxa (Voss et al., 2020), but too few spec-
Distribution: As currently recognized (see imens are available to assess the constancy of
Remarks), Marmosa demerarae occurs from east- observed phenotypic differences.
ern Venezuela eastward and southward through-
out the Guianas to Brazil. In Brazil, this species
Marmosa (Micoureus) jansae
occurs east of the Rio Negro on the north side of
Voss and Giarla, 2021
the Amazon; it occurs east of the Tapajós along
the south bank of the Amazon, and it occurs as Type material and type locality: ROM
far south as Bahia along the Atlantic coast (Silva 118880, the holotype by original designation,
et al., 2019: fig. 5; Voss et al., 2020: fig. 2). consists of the skin, skull, postcranial skeleton,
Remarks: This concept of Marmosa demer and frozen tissues of an adult male collected 42
arae follows Silva et al. (2019) and includes sev- km south and 1 km east of Pompeya Sur (0.68°
eral cytochrome b haplogroups from south of the S, 76.47° W), Parque Nacional Yasuní, Orellana
Amazon that Voss et al. (2020) recognized as province, Ecuador.
putative species and associated with the name Synonyms: None.
limae. However, the latter authors cautioned that Distribution: Marmosa jansae is currently
their voucher specimens of limae might not be known from the Amazonian lowlands of south-
phenotypically distinguishable from M. demer eastern Colombia (Putumayo), eastern Ecuador
arae, and subsequent examination of large series (Orellana, Pastaza), and northeastern PeruYou can also read
