Philander canus ( Osgood, 1913 )

Voss, Robert S., Díaz-Nieto, Juan F. & Jansa, Sharon A., 2018, A Revision of Philander (Marsupialia: Didelphidae), Part 1: P. quica, P. canus, and a New Species from Amazonia, American Museum Novitates 2018 (3891), pp. 1-72: 38-46

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Philander canus ( Osgood, 1913 )


Philander canus ( Osgood, 1913)  

Metachirus canus Osgood, 1913: 96   ; type locality Peru, San Martín, Moyobamba (6°03′ S, 76°58′ W; Stephens and Traylor, 1983).

Metachirus opossum crucialis Thomas, 1923: 604   ; type locality Bolivia, Santa Cruz, Santa Cruz de la Sierra (17°48′ S, 63°10′ W; Paynter, 1992).

Philander mondolfii Lew et al., 2006: 229   ; type locality Venezuela, Bolívar, Reserva Forestal de Imataca   GoogleMaps , Unidad V, between Tumeremo and Bochinche (8°00′ N, 61°30′ W).

Philander olrogi   Flores et al., 2008: 17; type locality   Bolivia, Santa Cruz, 7 km N Santa Rosa (17°03′ S, 63°35′ W) GoogleMaps   .

TYPE MATERIAL: The holotype (by original designation, FMNH 19347) consists of the skin and skull of an adult male collected by W.H. Osgood and M.P. Anderson on 4 August 1912. Although the skin is well preserved, the skull is broken and incomplete (the left zygomatic arch, the left squamosal, and the left bulla are all missing).

DISTRIBUTION AND SYMPATRY: Sequenced specimens and other examined material that we refer to Philander canus   have been collected in central and western Brazil, northern Argentina, Paraguay, eastern Bolivia, eastern Peru, northeastern Colombia, and Venezuela ( fig. 9 View FIG ). Although we have not examined specimens from northeastern Peru (Loreto), eastern Ecuador, or southeastern Colombia (Caquetá, Putumayo), future collecting may eventually fill in this geographic hiatus. Philander canus   occurs sympatrically with P. andersoni   in southern Venezuela (at El Platanal in Amazonas state), with P. mcilhennyi   in eastern Peru (e.g., at Balta, in Ucayali department) and western Brazil (at Sobral, in Acre state), and with P. pebas   in eastern Peru (e.g., at Balta, in Ucayali department).

Brazil, Tocantins (50): JF281038 View Materials   Brazil, Tocantins (50): JF281037 View Materials   , Tocantins (50): JF281039 View Materials   , Tocantins (49): JF281036 View Materials   , Pará or Tocantins: JF281034 View Materials   , Pará or Tocantins: JF281040 View Materials   , Pará (38): JF281035 View Materials   , Mato Grosso (29): KT153573 View Materials   , Mato Grosso (29): LPC392 Acre (12)   : MNFS1453 Mato Grosso (28)   : LPC584 Santa Cruz (9): NK11830 Grosso do Sul (27): KT153575 View Materials   , Meta (52): JAWK402 Mato Grosso do Sul (27): KT153576 View Materials   , Trujillo (91): JAWK281 canus   , Chaco (1): JQ778956 View Materials   , Ñeembucú (76): KM188486 View Materials   , Presidente Hayes (78): KM188487 View Materials   Formosa (2): JQ778957 View Materials   Chaco (1): JQ778962 View Materials   Ñeembucú (76): GD066 Grosso do Sul (27): KT153574 View Materials   Alto Paraguay (75): KM188488 View Materials   Formosa (2): JQ778958 View Materials   Grosso do Sul (27): LPC596 8): NK13894: NK13172 Bolívar (90): TK19152 NK13171   : LAR299   : LAR298 Amazonas (19): DQ236271 View Materials   Amazonas (19): DQ236275 View Materials (16)   : JLP15395 View Materials Amazonas (19): DQ236276 View Materials (16)   : U34678 View Materials   , Amazonas (19): DQ236274 View Materials   Amazonas (19): DQ236273 View Materials   Amazonas (19): DQ236272 View Materials   pebas   Amazonas (19): DQ236277 View Materials   Dios (85): NW579 80): TK73935 81): KM 188489 View Materials 80): TK73919   Ecuador, Orellana (57): F40358 View Materials  

DESCRIPTION: Dorsal pelage short (usually <14 mm) and uniformly grayish, usually without any trace of darker middorsal pigmentation; fur of crown (between the ears) usually grizzled gray; pale preauricular spot often present; ventral fur continuously self-whitish, -cream, or -buffy, at least along the midline, but sometimes with broad lateral zones of graybased hairs; pinnae pale (unpigmented) basally, but blackish distally; dorsal pelage of hind feet usually pale, but sometimes indistinctly darker over lateral metatarsals (never distinctly marked with black); scaly part of tail usually <½ white distally but seldom <¼ white. Nasal bones short (about 46% of condylobasal length on average), never extending posteriorly to or between postorbital processes. Unworn third upper premolar (P3) apparently always with complete labial cingulum extending along entire base of tooth; crown length of upper molar series 13.0 ± 0.4 mm (sexes combined; observed range 12.2–14.1 mm, N = 99); enameled lingual surfaces of upper molars smooth, not crenulated; pre- and postcingula consistently absent; lower molar postcingulids absent.

PHYLOGEOGRAPHY AND GEOGRAPHIC VARIATION: Our phylogenetic analysis of 32 sequences of Philander canus   reveals no comprehensible phylogeographic structure, with haplotypes from northern populations (in Colombia and Venezuela) mixed in among those from central Brazil, Bolivia, Paraguay, and northern Argentina ( fig. 16 View FIG ). Uncorrected sequence divergence at the cytochrome- b locus among haplotypes that we assign to P. canus   is only about 0.8% despite the very wide geographic dispersion of collecting localities. Although we have not statistically tested for geographic variation in morphology among our samples, this appears to be another phenotypically rather uniform species despite modest sample differences in pelage pigmentation (some populations tending to have self-whitish or -cream underparts, whereas others have self-buffy ventral fur).

COMPARISONS: Morphological comparisons of Philander canus   with P.quica   have already been described (see above) and comparisons with P. pebas   will be described subsequently (see below). It remains to compare this species with members of the P. opossum   complex, which—as defined earlier in this report—includes P. opossum   , P. andersoni   , and P. mcilhennyi   .

Philander canus   is superficially similar to P. opossum   , with which it has long been associated as a subspecies or synonym (e.g., by Cabrera, 1958; Patton and da Silva, 1997, 2008; Gardner, 2005; Chemisquy and Flores, 2012; Hice and Velazco, 2012). Although the geographic ranges of P.canus   and P. opossum   are not currently known to come into contact, it seems plausible that these species are sympatric or parapatric in the Brazilian states of Mato Grosso, Tocantins, and southern Pará, where Cerrado vegetation comes into contact with southeastern Amazonian rainforest. Philander canus   is substantially smaller, on average, than P. opossum   in several same-sex univariate comparisons ( table 11), notably in condylobasal length (CBL), nasal length (NL), least interorbital breadth (LIB), least postorbital breadth (LPB), palatal length (PL), and maxillary toothrow (MTR). Despite some overlap in observed ranges for all dimensions, measured samples of these species have nonoverlapping multivariate distributions ( fig. 17 View FIG ), and general-size-adjusted shape coefficients indicate that nasal morphology accounts for much of the observed divergence ( table 12). Visual comparisons suggest that the posterior portion of the nasals of P. canus   are typically much broader than those of P. opossum   , do not extend as far posteriorly, and lack the deep posterolateral notches that are often present in the latter species ( fig. 18 View FIG ). Philander canus   and P. opossum   both have uniformly grayish dorsal fur, mostly pale hind feet, and self-colored ventral fur, but tail pigmentation might be useful for field identification. Whereas the scaly part of the tail is almost always <½ white in specimens of P.canus   , the scaly part of the tail is typically ≥½ white in specimens of P. opossum   , and this modal difference might be expected to become even more pronounced in sympatry. 9

9 We suspect (although there is no behavioral evidence to support our conjecture) that the black-and-white tail markings of Didelphini   have some social-signaling function that might be coopted for species recognition in sympatry.

differences (Shape) for multivariate analyses of Philander canus   versus P. opossum   a

Philander canus   is much smaller, on average, than either P. andersoni   or P. mcilhennyi   , from which it also differs in nasal shape as described and illustrated above (all members of the P. opossum   complex have long, narrow nasals that are often laterally notched and often extend posteriorly to or between the postorbital processes). Large generalized distances (appendix 4) suggest that multivariate ordinations of P. canus   with either P. andersoni   or P. mcilhennyi   would show nonoverlapping distributions, but we have not performed those analyses because these species are easy to tell apart by other characters. The dorsal pelage pigmentation of P. canus   (uniformly gray; fig. 10 View FIG ) is quite unlike that of P. andersoni   (with a distinctly blackish middorsal stripe) and P. mcilhennyi   (some specimens of which are completely blackish). The self-whitish, -cream, or -buffy underparts of P. canus   likewise contrast with the mostly gray-based ventral pelage of P. andersoni   and the almostblackish ventral fur of P. mcilhennyi   ( fig. 11 View FIG ). Whereas the hind feet of P.canus   are covered dorsally with pale fur, the hind feet of P. andersoni   and P. mcilhennyi   are either completely blackish or have black metatarsals and abruptly whitish digits. Lastly, the scaly part of the tail is almost always <½ white in P.canus   but apparently always ≥ ½ white in P. andersoni   and P. mcilhennyi   .

REMARKS: As understood herein, Philander canus   includes the nominal taxa crucialis, mondolfii   , and olrogi   . The latter are represented in our molecular analyses by: (1) a CYTB sequence we obtained from a Bolivian specimen (AMNH 260034) that was collected near the type locality of crucialis and that resembles the holotype of crucialis in qualitative and morphometric traits; (2) two CYTB sequences, one from a Colombian specimen ( KU 123943 View Materials ) and another from a Venezuelan specimen ( KU 120245 View Materials ) that were part of Lew et al.’s (2006) original material of mondolfii   ; and (3) CYTB sequences that we obtained from two specimens (AMNH 261271, 261272) that were part of Flores et al.’s (2008) original material of olrogi   . All these specimens conform to our morphological diagnosis of P.canus   , so the conclusion that the nominal taxa in question are conspecific seems straightforward, but brief comments on each synonym are appropriate.



Thomas (1923) described crucialis on the basis of a single specimen, which he compared only with azaricus (= P.quica   ; see above). Later, with more Bolivian material at hand for comparison with topotypical specimens of Osgood’s species, he ( Thomas, 1928) judged crucialis and canus   to be indistinguishable. We agree.

Lew et al. (2006) described mondolfii   based on several dozen specimens from Colombia and Venezuela that the authors compared carefully with other species of Philander   known to occur in or near those countries, including P. andersoni   , P.deltae   , P. “ fuscogriseus ” (= P.melanurus   ), and P. opossum   , but they did not compare mondolfii   with P.canus   . Although we have not examined the holotype or paratypes of mondolfii   —all currently inaccessible in Venezuelan museums—we did examine 16 specimens that were part of Lew et al.’s (2006) original material. These specimens (AMNH 16951, 30709, 30711–30714, 133119, 133120, 136163, 136167–136169, 139221; KU 120233 View Materials , 120245, 123943) so closely resemble the type of P.canus   and other referred material from eastern Peru that we could not find any phenotypic basis for retaining mondolfii   even as a subspecies.

Flores et al. (2008) described olrogi   on the basis of seven specimens from Bolivia and Peru. Although we have not seen the holotype, we examined the skull of a paratype (AMNH 246441) as well as several other specimens that were part of Flores et al.’s (2008) original material (AMNH 261269–261272). Despite careful study, we confess ourselves unable to consistently distinguish these specimens from material that the authors referred to P. opossum canus   . Although they reported a principal-components analysis that was said to support the recognition of olrogi   as a distinct taxon, canus   and olrogi   have overlapping distributions in their illustrated results ( Flores et al., 2008: fig. 5 View FIG ). In the absence of compelling evidence for the phenotypic distinctness of these genetically indistinguishable nominal taxa, we interpret the allegedly diagnostic traits of olrogi   to be aspects of intraspecific morphological variation within P. canus   .

Cabrera (1958: 35) listed nigratus as a synonym of canus   (which he ranked as a subspecies of P. opossum   ), but the holotype (BMNH and other material that we refer to nigratus 10 are larger animals (LM = 14.7–16.4 mm) with much darker dorsal fur, completely gray-based ventral fur, blackish feet, shorter white tail-tips (less than ¼ of the tail is unpigmented in most specimens), and an incomplete labial cingulum on P3. Although we do not know whether nigratus is a valid species, its phenotype more closely resembles those of species in the P.opossum   complex than that of P.canus   .

Hershkovitz (1997) used the name Philander opossum quica   for many specimens that we refer to P. canus   , including all the material he listed from central Brazil (Goiás, Mato Grosso) and Bolivia; among the material that he listed from eastern Peru are specimens that we refer to both P. canus   and P. pebas   . The “dichromatism” that he ( Hershkovitz, 1997: 49) noted among specimens of “ P. o. quica   ” from Balta (in the Peruvian department of Ucayali) is the result of sympatry rather than polymorphism: of the six specimens in question that we examined, three (LSUMZ 12006, 12008, 12009) are P.canus   and the others (LSUMZ 12007, 12010, 14011) are P. pebas   (see Specimens Examined for both taxa, below; a third congener, P. mcilhennyi   , also occurs at Balta).

The western Amazonian specimens that Patton et al. (2000) referred to Philander opossum canus   include examples of both P.canus   and P. pebas   . Of the five that we were able to examine— the others having been returned to Brazil —four (MVZ 190343–190346) are P. pebas   ; only one (MVZ 190347, from the state of Acre, near the Peruvian border) is actually P.canus   . The central Amazonian specimens that Nunes et al. (2006) identified as P. canus   are also examples of P. pebas   . 11 The only other material of P.canus   that we have seen from the Brazilian Amazon (besides the MVZ specimen from Acre) is a small series collected many years ago along the lower Rio Madeira (in Amazonas state) and a single specimen from the upper Madeira (in Rondônia).

HABITATS: The geographic range of Philander canus   extends over a wide range of biomes or ecoregions (including the Cerrado, Chaco, Pantanal, Llanos, and Amazonia), and it is possible that the species occurs in a corresponding variety of habitats, but definite ecological information associated with collected specimens is hard to find. Other species of Philander   are

10 From the eastern Andean foothills of Junín and Ayacucho departments, Peru: BMNH,,; FMNH 65782; LSUMZ 16398, 16399; MUSM 71   .

11 We are indebted to S.E. Pavan, who kindly examined the MPEG voucher specimens from Nunes et al.’s (2006) study at our request.

known to live in rainforest, so collection records from biomes dominated by savanna vegetation and/or dry forest (e.g., the Cerrado, Chaco, and Llanos) seem anomalous, but the literature on Cerrado mammals provides a few relevant observations.

In the Cerrado landscapes of eastern Bolivia, Philander canus   is apparently restricted to tall evergreen gallery forests and does not seem to occur in the savannas and dry forests that cover much of the landscape ( Emmons et al., 2006). In the Cerrado of central Brazil, P. “ opossum   ” (presumably P.canus   ) is also said to be a gallery-forest species ( Mello and Moojen, 1979; Redford and Fonseca, 1986; Alho, 2005), but in one report of a multiyear trapping study P. “ opossum   ” was said to prefer gallery forest but to occur frequently in other local habitats, including open grassland, shrub savanna, and dry forest ( Alho et al., 1986). Following Pulliam’s (1988) ecological terminology, we conjecture that gallery forests are probably the source habitat for P.canus   in Cerrado landscapes, whereas open formations (including dry forests) are likely to be sink habitats. In the Chaco of northern Argentina, Philander   opossum   ” (presumably P. canus   ) is also said to occur in gallery forests ( Huck et al., 2017).

Information about the habitat distribution of Philander canus   appears to be unavailable from trapping studies in the Pantanal and Llanos, but we suspect that it is largely restricted to gallery forests in those ecoregions as well. Nevertheless, Lew et al.’s (2006) summary of macrohabitats where Philander   mondolfii   ” (= P.canus   ) has been collected in Venezuela (including lowland and submontane rainforest, semideciduous forest, and tree savannas) suggest that it has broad ecological tolerances, at least where other sympatric congeners are not known to occur.

We have not found any published accounts of where specimens that can definitely be identified as Philander canus   have been collected in Amazonia. Although the natural climax vegetation throughout this enormous ecoregion can be broadly characterized as lowland rainforest, local disturbance (e.g., from lateral migration of rivers within their meander belts) and edaphic factors can result in a surprising diversity of natural vegetation types at many Amazonian localities ( Pires and Prance, 1985; Puhakka and Kalliola, 1995), and anthropogenic habitats are also scattered throughout the region. The geographic distribution of Amazonian collection localities for P.canus   provides no habitat clues, because these localities are not clustered around savanna enclaves, human settlements, or other obvious landscape features. The known Amazonian range of P.canus   broadly overlaps those of P. andersoni   , P. mcilhennyi   , and P. pebas   , so it would be reasonable to suppose that competitive interactions with sympatric congeners might restrict the habitat occupancy of this species to some extent, but the information compiled for this report is entirely inadequate even for conjecture.

SPECIMENS EXAMINED (N = 154): Bolivia — Beni, Arruda ( FMNH 114701 View Materials )   , Camiaco ( AMNH 210402 View Materials )   , Casarabe ( AMNH 261269–261272 View Materials ; MSB 55854 View Materials ), 8 km N Exaltación ( AMNH 210403 View Materials )   , Magdalena ( FMNH 114714 View Materials )   , Mamore River ( AMNH 210409 View Materials )   , 4 km SE Palacios (210410), Puerto Caballo ( AMNH 210411 View Materials )   , Puerto Siles ( AMNH 210413 View Materials , 210414 View Materials )   , Río Tijamuchi ( AMNH 261273 View Materials )   , San Joaquin ( FMNH 114685 View Materials , 114694 View Materials , 114707 View Materials )   ; Pando, Bella Vista ( MSB 57006 View Materials , AMNH 262413 View Materials )   ; Santa Cruz, 7 km E aserradero Moira ( EBD 8736 View Materials )   , 6 km W Asención ( MSB 55855 View Materials )   , Ayacucho ( USNM 390564 View Materials )   , Becerra (390565), 2 km N Chapare River mouth ( AMNH 210416 View Materials )   , 2 km SE Cotoca ( MSB 59887 View Materials )   , Estancia Cachuela Esperanza ( AMNH 260034 View Materials , MSB 55073 View Materials ), Hamecas ( AMNH 135887 View Materials )   , La Laguna ( MSB 55856 View Materials )   , 3 km SE Montero ( AMNH 263964 View Materials , MSB 67025 View Materials ), Palmar ( USNM 390562 View Materials )   , San Miguel Rincón ( AMNH 260037 View Materials , MSB 55074 View Materials , 55075 View Materials ), 10 km N San Ramón ( AMNH 261277 View Materials , 261278 View Materials )   , Santa Cruz de la Sierra ( BMNH [holotype of crucialis]), 15 km S Santa Cruz ( AMNH 263966 View Materials , MSB 58517 View Materials ), 7 km N Santa Rosa ( AMNH 246441 View Materials [paratype of olrogi   ]), near Warnes ( USNM 390005 View Materials , 390009– 390012 View Materials )   . Brazil — Acre, Sobral on Rio Juruá ( MVZ 190347 View Materials )   ; Amazonas, Auara Igarapé on Rio Madeira ( AMNH 91749 View Materials , 91750 View Materials )   , Borba on Rio Madeira ( AMNH 91748 View Materials )   , Lago Sampaio on Rio Madeira ( AMNH 92761 View Materials , 92762 View Materials )   , “Santo Antonio de Uayara” on Rio Madeira (= Santo Antonio de Guajará ; AMNH 92293 View Materials )   ; Goiás, Anápolis ( AMNH 133043 View Materials , 133046 View Materials , 133047 View Materials , 133055 View Materials , 133056 View Materials , 133062 View Materials , 133064 View Materials , 133068–133070 View Materials , 133073–133075 View Materials , 133082 View Materials , 133084–133086 View Materials , 133091–133094 View Materials , 133096–133101 View Materials , 133123 View Materials , 133172 View Materials , 133171 View Materials , 133182 View Materials , 133192 View Materials , 133195 View Materials )   , 24 km SE Formoso ( LACM 10086–10088 View Materials )   ; Mato Grosso, Caceres ( USNM 390014 View Materials )   , Fazenda São Luis ( MVZ 197403 View Materials )   ; Mato Grosso do Sul, Corumba ( USNM 390013 View Materials )   , Passo do Lontra ( MVZ 197402 View Materials )   ; Rondônia, Porto Velho ( USNM 390001 View Materials )   . Colombia — Boyacá, Río Cobaría ( FMNH 92297 View Materials )   ; Meta, Finca El Capricho ( KU 123943 View Materials )   , Restrepo ( AMNH 133119 View Materials )   , Villavicencio ( AMNH 136168 View Materials , 136169 View Materials , 139221 View Materials )   . Paraguay — Alto Paraguay, Estancia Doña Julia ( TTU 79753 View Materials )   ; Central , 17 km E Luque ( MVZ 144304 View Materials )   ; Presidente Hayes, Estancia Loma Porá ( TTU 80404 View Materials )   . Peru — Cusco, Camisea ( MUSM 14150)   , Hacienda Cadena ( FMNH 66412 View Materials , 68332 View Materials )   , Quincemil ( FMNH 75094–75096 View Materials )   ; Huánuco, Moyuna ( MUSM 83)   ; Madre de Dios, “Albergue Lodge Cuzco Amazonico” (= Cusco Amazónico ; MVZ 157613 View Materials , 165927 View Materials )   , Boca Río Colorado ( FMNH 84247 View Materials )   , Lago Sandoval ( MVZ 157614 View Materials )   , mouth of Río La Torre ( LSUMZ 24591 View Materials )   , 6 km W Río Tambopata ( USNM 39002 View Materials )   ; 2.75 km E Shintuya ( FMNH 169815 View Materials )   ; Pasco, San Pablo ( AMNH 230034 View Materials )   , Nevati ( AMNH 230028 View Materials , 230030 View Materials , 230031 View Materials , 254509 View Materials )   ; San Martín, Bellavista ( MUSM 92)   , Moyobamba ( FMNH 19347 View Materials [holotype of canus   ]), Rioja ( MUSM 88)   ; Ucayali, Balta ( LSUMZ 12006 View Materials , 12008 View Materials , 12009 View Materials )   , 59 km SW Pucallpa ( USNM 499001 View Materials , 499002 View Materials )   , Boca Río Urubamba ( AMNH 75906–75908 View Materials )   , Lagarto ( AMNH 76636 View Materials )   , Santa Rosa ( AMNH 75909 View Materials )   . Venezuela — Amazonas, El Platanal ( EBD 8954 View Materials , 8956 View Materials )   ; Apure, 29 km SSW Santo Domingo ( USNM 418545 View Materials , 418546 View Materials )   ; Bolívar, 20 km W La Paragua ( USNM 388403 View Materials )   , Maripa ( AMNH 16951 View Materials )   , Río Yuruán ( AMNH 30709–30714 View Materials )   ; Trujillo, 9.8 km NNE Motatán ( KU 120233 View Materials , 120245, 120246, 120251)   , 19 km W Valera ( USNM 371322 View Materials )   ; Zulia, 60 km WNW Encontrados ( USNM 418548 View Materials )   .


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Philander canus ( Osgood, 1913 )

Voss, Robert S., Díaz-Nieto, Juan F. & Jansa, Sharon A. 2018

Philander olrogi

Flores, D. A. & R. M. Barquez & M. M. Diaz 2008: 17

Philander mondolfii

Lew, D. & R. Perez-Hernandez & J. Ventura 2006: 229

Metachirus opossum crucialis

Thomas, O. 1923: 604

Metachirus canus

Osgood, W. H. 1913: 96