Composition of the Family Didelphidae Gray, 1821 (Didelphoidea

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Zoology NEW SERIES, NO. 86

Composition of the Family Didelphidae Gray, 1821 (Didelphoidea: Marsupialia), with a Review of the Morphology and Behavior of the Included Four-Eyed Pouched Opossums of the Genus Philander Tiedemann, 1808

Philip Hershkovitzf

Curator Emeritus Division of Mammals Department of Zoology Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago. Illinois 60605-2496 USA

Accepted April 4, 1996 BlOlOGY LIBRARY HALL Published May 30, 1997 101 BURRILL

Publication 1485 _ - /» -mfll OCT 161997 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY k © 1997 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents 3. Hindfoot of Chironectes minimus 7 4. Dorsal pigmentation of Philander opossum and P. andersoni 9 Abstract 1 5. Geographic distribution of f/?/7flnrfer .... 10 Introduction 1 6. Chcmdixdi oi Philander opossum 16 Abbreviations 2 7. Caudal segments of the four largest Didelphidae: Taxonomic Status, The American opossums 18 Genera 2 8. Scales and bristles of Philander opos- A Morphological Test 4 sum tail 19 Genus Philander Tiedemann (Four-Eyed 9. Transitional caudal vertebrae of Philan- Pouched Opossums) 8 der opossum 20 Synonymy 8 10. Points of cranial measurements 22 Distribution 8 11. Diagram of didelphid skull 23 Nomenclature 10 12. Left side of Philander opossum skull External Characters and Comparisons 11 and mandible 25 Cranial Characters and Comparisons 15 13. Philander opossum sVnW 26 Dental Characters and Comparisons 17 1 4. Tooth rows of Philander opossum 27 Descriptions of Individual Teeth of Philan- 15. Upper incisors and canine of Philander der 18 opossum, Caluromys philander, Lutreo- Dental Abnormalities 23 lina crassicaudata, Metachirus nudi- Sexual Dimorphism 24 caudatus 28

Karyology 24 16. Sidi^gered Xy m Philander opossum 29 Parasites 25 17. Deciduous premolar 3 and molars 30 Species and Named and Unnamed Sub- 18. Accessory molar cusps 31 species OF Philander opposum 32 19. Conules of m^ of Philander from vari- Philander opossum Linnaeus 33 ous geographic localities 32 Philander opossum opossum Linnaeus 35 20. M^ of Philander opossum 33 Philander opossum quica Temminck 40 21. Lectolypes of Philander opossum 39 Philander opossum frenatus Olfers 51 22. South American collecting locahties 81 Philander opossum melanurus Thomas .... 51 Philander opossum subsp. nov. 1 52 Philander opossum suhsp. nov.? 52 Philander opossum fuscogriseus J. A. Allen 52 List of Tables Philander andersoni Osgood 59 Philander andersoni andersoni Osgood .... 61

Philander andersoni mcilhennyi Gardner 1 . Summarized measurements of Philan- and Patton 63 der opossum and P. andersoni sub- Behavior of the Four-Eyed Pouched species 12 Opossums (Philander opossum) 64 2. Dimensions and proportions of limb Alphabetical List of Collecting Locali- bones 15 ties 80 3. Vertebral bone formulae 19 Gazetteer 85 4. Nipple formulae 21 Acknowledgments 95 5. Symbols for cranial bones and Literature Cited 95 foramina 24 6. Symbols for dental elements 34 7. Conules of m^ 35

8. Karyotypes of largest American mar- List of Illustrations supials 35 9. Measurements of Philander opossum opossum 36

1 . The four genera of largest American 10. Measurements of Philander opossum marsupials quica 42

2. Skulls of the four largest American 1 1 . Measurements of Philander opossum marsupials melanurus 52

lU 1 2. Measurements of Philander opossum 1 6. Measurements of Philander andersoni fuscogriseus 54 mcilhennyi 64 13. Ratios of hairy tail bases to tail 17. Reproductive condition of 22 female lengths 59 Philander opossum from Nicaragua 68 14. Sympatry between Philander species .... 60 18. Body mass and above-ground foraging .. 72 15. MQd&urcxn&nis oi Philander andersoni 19. Duration of residence in study area 78 andersoni 62 Composition of the Family Didelphidae Gray, 1821 (Didelphoidea: Marsupialia), with a Review of the Morphology and Behavior of the Included Four-Eyed Pouched Opossums of the Genus Philander Tiedemann, 1808

Philip Hershkovitzt

Abstract

The generic content of the family Didelphidae as currently conceived consists of Didelphis, Philander, Chironectes, and Lutreolina. It is shown that most defining characters of the Di- delphidae determined from those of the genus Didelphis are highly derived and not shared by Chironectes or Lutreolina. Accordingly, the family content was restricted to Didelphis and the genus Philander, which is smaller and less derived than Didelphis. The genus Didelphis had previously undergone a partial taxonomic revision. A complete revision of the genus Philander now follows. Two species of the genus have been recognized. Comparisons were made with outgroups Chironectes and Lutreolina. Habits of Philander, insofar as known, are described. The more primitive genus Metachirus, which is sometimes regarded as a didelphid, is being treated separately.

Introduction This report is the author's sixth on New World marsupial systematics and behavior. The series The Phi- four-eyed pouched opossums, genus began in 1992 with a taxonomic revision of the lander Tiedemann, 1808, most related to nearly gracile mouse opossum, genus Gracilinanus the of the Lin- larger opossums genus Didelphis (Marmosidae) (Hershkovitz, 1992a). It was fol- naeus, 1 758, are common much of the throughout lowed in the same year by a critical examination neotropics from northern Argentina into Tamau- of the significance of ankle bones as phylogenetic lipas, Mexico. About 20 Linnaean names have indicators (Hershkovitz, 1992b). The description been proposed for four-eyed pouched opossums, of a seemingly abnormal staggered third lower in- but only two species are recognized here. The ear- cisor, which proved to be the hallmark of all di- liest known, P. opossum Linnaeus, 1758, with its delphimorphs since the earliest Cretaceous, if not geographic distribution nearly equal to that of the late Jurassic, was published in 1995. In press is a genus, is the smaller of the two and generally less taxonomic review and life history account of the darkly colored. Philander andersoni Osgood, relict Chilean mouse opossum Dromiciops (Mi- mainly western Amazonian in distribution, con- crobiotheriidae). Also awaiting publication is an- tains two subspecies. One, P. andersoni andersoni other review, that of the brown Osgood, occurs in Peru and in the Territorio Fed- four-eyed pouchless Metachirus. This taxon was eral Amazonas, Venezuela. The wide gap in dis- opossum, genus first for the Philander tribution between the two countries may be for used at as an outgroup As work Metachirus lack of collecting or the result of a climatic event. opus. progressed, however, The blackish P. andersoni mcilhennyi Gardner took on greater importance and is being complet- and Patton is known only from the upper Rio Pu- ed under its own title. nis basin in Amazonian Peru. In preparation is a monograph of living New

RELDIANA: ZOOLOGY, N.S., NO. 86, MAY 30, 1997, PP. 1-103 World marsupials from which the above-men- scansorial, and members of the genus Didelphis. tioned articles have been partially derived and Discovery of the first caenolestid by Tomes in elaborated. 1863 added another dimension to the concept of American marsupials by directing attention to the existence of nonpouched, non-prehensile-tailed and terrestrial American marsupials. Notwith- Abbreviations standing, apart from accommodations made in catalogs for the increasing number of newly dis- The following abbreviations are used for the covered taxa, no significant changes were made institutions where the specimens examined are in the classifications. Recent catalogers (Cabrera, preserved. 1958; Hall & Kelson, 1959; Honecki et al., 1982; Gardner, 1993) the three current orders AMNH - American Museum of Natural His- recognized of American marsupials, each with a single fam- tory, New York its content as in the literature. = ily, found The cat- bm(nh) British Museum (Natural History), with the numbers of their liv- London egories, respective = ing genera and species in parentheses (ex Gardner, FMNH Field Museum of Natural History, 1993), follow: Chicago Mvzuc = Museum of Vertebrate Uni- Zoology, Order Didelphimorphia versity of California, Berkeley 63 = Family Didelphidae (15 genera, species) USNM National Museum of Natural History, Order Paucituberculata Washington, D.C. Caenolestidae - Family (3, 5) USPMZ Universidade de Sao Paulo Museu de Order Microbiotheria Sao Paulo, Brasil Zoologia, Family Microbiotheriidae (1, 1) MPEG = Museu Paraense Emilio Goeldi, Be- lem, Brazil Research by Reig et al. (1977) on chromosomes = UKMNH University of Kansas Museum of of the Didelphidae resulted in significant advanc- Natural History, Lawrence es. Three karyotypes were distinguished among = LSUMZ Louisiana State University Museum living American marsupials. The largest diploid of Zoology, Baton Rouge number of chromosomes was 22, possessed by MNRJ = Museu Nacional, Rio de Janeiro four genera of American marsupials, namely Di- = RMNH Rijksmuseum van Naturalijke Histo- delphis (three species), Chironectes (one), Lutreo- rie, Leiden lina (one), and Philander (two). The other com- - plements were 2n 18 in the short-tailed mouse opossum Monodelphis (three of about 1 5 species) — and In 14 in Caluromys (three species), Calu- Didelphidae: Taxonomic Status, romysiops (one), Marmosa (sensu lata; eight spe- The Genera cies of about 38, since rearranged into four genera), Metachirus (one), and Dromiciops (one). = The family Didelphidae, erected by Gray, 1821, The 2n 14 karyotype is generally regarded as included all then-known American marsupials di- the primitive one. No reclassification of American vided into the genera Didelphis Linnaeus and marsupials was based on their cytogenetic consti- Cheironectes [sic] Illiger. It was not until late in tution. In any case, Kirsch (1977) had already re- the 20th century that critical attention was paid to moved woolly opossums {Caluromys) from the higher systematic categories of living New World Didelphidae and erected the family Caluromyidae marsupials. for them. In 1992, Hershkovitz (1992a) estab- The early arrangement of the taxa persisted, lished the family Marmosidae for most so-called nevertheless, as if all female marsupials were mouse opossums previously known as Marmosa pouched and all individuals prehensile-tailed. and raised Glironia to family rank. Four opossum

Fig. 1. Representatives of the four genera of large American marsupials usually referred to as the family Didel- phidae, subfamily Didelphinae. Here only Didelphis marsupialis and Philander opossum are so classified. (Figures not to scale.)

FIELDIANA: ZOOLOGY Lutreolina crasBicaudata Thomas

HERSHKOVITZ: PHILANDER and soft anatomical does genera were retained in Didelphidae because they logical, characters, place = Metachirus in a sister relationship with Didelphis and are the largest and their karyotype is 2n 22. Philander, in contrast to Marmosa and Monodelphis, These four are Chironectes, genera Didelphis, as do DNA-DNA hybridization data (Kirsch et al., Philander of Linnaeus, and Lutreolina Thomas, 1995; Lapointe and Kirsch, 1995). Nevertheless, of Metachirus either as the sister of 1910 (Figs. 1-3). placements {Di- + with the mouse lin- Biochemical, chromosomal, and anatomical delphis Philander), opossum eage, or outside of both clades are all equally likely characters were considered by Reig et al. (1985) based on an evaluation of these alternative topologies in an to determine the rela- attempt phylogenetic by the log-likelihood test of Kishino and Hasegawa tionships among didelphoid marsupials. The ex- (1989). As a consequence, perhaps Metachirus is best considered as a basal member of the + tinct forms they considered are outside the scope {Didelphis Philander) and {{Marmosa -I- Micoureus) + Mono- of this monograph, as are most of the included delphis) clades. This is the same conclusion reached The of concern here include genera. Didelphidae by Kirsch (1977) and Reig et al. (1987). the three subfamilies Herpetotherinae, Caluromyi- nae, and The latter was further sub- Didelphinae. Returning to Philander, Patton et al. (1995 divided into the tribes Didelphini, Metachirini, [1996]) do not explain why they dropped the prior and Marmosini. Their tree et al., 1985, (Reig Fig. name Philander andersoni Osgood, 1913, and 2, p. 339) shows all American didelphoids and consistently used instead its junior synonym, P. basic Australian forms from Cre- emerging Upper mcilhennyi Gardner and Patton, 1972, in all their taceous Microbiotheriidae, an improbability. It discussions of marsupial relationships. also shows didelphids arising in the Eocene and giving rise to the Marmosini with the primitive = karyotype 2n 14 and pouchless reproductive system in the Oligocene-Miocene. This also A IVIorphological Test seems unlikely. The pouchless and non-prehensile-tailed, terrestrial Metachirini with 2n = 14 The four genera of large American marsupials may have given rise to didelphids rather than the have consistently been regarded as forming a nat- reverse as depicted. ural assemblage within the family Didelphidae. Studies by Kirsch et al. (1993a) of DNA/DNA This can be tested by the 30 morphological char- hybridization resulted in assignment of the five acters outlined below. Philander shares most largest American genera {Didelphis, Philander, characters with Didelphis. Chironectes and Lu- Lutreolina, Chironectes, and Metachirus) to the treolina, however, have diverged widely from the Didelphidae. Morphologically, as shown below, preceding two genera and from a hypothetical Lutreolina and Chironectes share few traits with common ancestor of all four. Two of the few each other and fewer with Didelphis, and the in- = shared characters (large size and In 22) may dicated position of Metachirus is even less tena- have evolved independently; other characters that ble. may appear trivial are diagnostic nevertheless. An Patton et al. (1995 [1996]) regarded the system- appraisal of the evolutionary stage of each char- atic position of Metachirus to be somewhat equiv- acter is shown in parentheses. ocal:

1. Largest of living New World marsupials While the brown four-eyed opossum Metachirus (derived) forms a sister taxon to the {Didelphis + Philander) The four genera including the largest liv- clade in all trees, this relationship is not well sup- ing New World opossums (Fig. 1) range in ported. Bootstrap values for parsimony analyses are size from the only 68 and 54 for DNA and amino acid sequences, smallest. Philander, through respectively, and decay indices indicate that it would Chironectes and Lutreolina, to the largest, take two or three additional to the only steps collapse Didelphis. Size may have evolved indepen- relationship between Metachirus and the other two dently in each genus, and its phylogenetic genera. Similarly, confidence limits for this node are in the context is dubi- only 28% in the DNA neighbor-joining distance tree. significance present = The pouchless, primitively 2n 14 brown four-eyed ous. opossum contrasts markedly with the pouched, de- 2. 2n = 22 = Karyotype (derived) rived 2n 22 Didelphis and Philander pair, although The same karyotype evolved indepen- the three genera apparently do share similarities in dently in unrelated Marmosa canescens bullar structure (Reig et al., 1987). Indeed, the total from a 2n = 14 evidence Wagner tree presented by Reig et al. (1987, karyotype (Engstrom & Fig. 59), based on 45 craniodental, cytological, sero- Gardner, 1988, p. 231). It also occurs in

FIELDIANA: ZOOLOGY Fig. 2. Dorsal and ventral aspects of skulls of four large American marsupials (shown in order of decreasing size from left to right).

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HERSHKOVrrZ: PHILANDER Australian Macropodidae and Potoroidae. 1 9. Spread of zygomatic arches comparatively The phylogenetic significance of the same narrow (plesiomorphic) chromosome number in Chironectes and Excludes Chironectes Lutreolina may be misleading. A different 20. Lacrymal foramina facial (derived) karyotypic pattern in Didelphis virginiana Excludes Lutreolina and Chironectes does not alter the fact that this and species 21. Nasals behind convergent to a single point D. are alike and marsupialis very nearly or with slight spread between points (ple- most related. intimately siomorphic?) 3. Tail as long as or longer than head and Excludes Lutreolina and Chironectes combined body (incipiently derived) 22. Maxillary sheath for insertion of lower ca- Excludes Lutreolina nine shallow (plesiomorphic) 4. Tail fully prehensile (derived) Excludes Lutreolina and Chironectes Excludes Lutreolina and Chironectes 23. Occipital condyles projecting behind ver- 5. Tail terete (plesiomorphic) tical plane of supraorbital bone (plesio- Excludes Lutreolina and Chironectes morphic) 6. Tail base not thickened notably (plesio- Excludes Chironectes morphic) 24. Incisive foramina long, narrow, and ta- Excludes Lutreolina (muscle, not fat) pered (plesiomorphic) 7. Tail thinly pilose, the scales fully exposed Excludes Chironectes dorsally (plesiomorphic) 25. Foramen ovate Excludes Lutreolina magnum (?) Excludes Lutreolina 8. Complete marsupium present in females, 26. Ventral mandibular plane between canine absent in males (derived) and incisors horizontal Excludes Lutreolina and Chironectes (plesiomorphic) Excludes Lutreolina and Chironectes (?) 9. Thumb and first toe fully opposable (de- 27. incisors 2-5 with little or no deflec- rived) Upper Excludes Lutreolina tion inward and upward (plesiomorphic?) Excludes Lutreolina and Chironectes 10. Pedal digits partially or not webbed (incip- 28. P^ as as P' iently derived) nearly long (plesiomorphic?) Excludes Chironectes Excludes Lutreolina 29. Lower incisors not 1 1 . Crown color pattern whitish above eyes, overlapping (staggered not brownish between (derived) ij included) (plesiomorphic) Excludes Chironectes and Lutreolina Excludes Lutreolina and Chironectes

12. Ears large, roundish, leaflike, and when 30. Locomotion terrestrial-scansorial (derived) laid forward extending midway or more to Excludes Lutreolina and Chironectes outer canthus of eye (possibly derived) Excludes Lutreolina and Chironectes Taking into account other characters mentioned reveal Didelphis as a 13. Manual ungues sharp and protruding (ple- beyond, comparisons shag- oversized ''Philander^ The otter-like Chiro- siomorphic) gy, Excludes Chironectes nectes, distinguished mainly by its nonprehensile, naked with both sexes and the ter- 14. Paired posteromedian palatal vacuities tail, pouched, present (plesiomorphic) restrial weasel-like Lutreolina, with its nonpre- Excludes Chironectes hensile, completely hairy tail and nonopposable 15. first have radiated far from roots, Premaxillary symphysis angular (plesio- digit, didelphoid j morphic) Lacking affinities with each other of didelphid Excludes Lutreolina and Chironectes grade, Chironectes and Lutreolina are each treat- 16. Depression at proximal end of temporal ed as a type of a distinct subfamily. The new ar- ridges (?) rangement follows: Excludes Lutreolina and Chironectes 17. Interorbital region not notably constricted Superfamily Didelphoidea (derived?) Family Didelphidae Gray, 1821 | Excludes Lutreolina Subfamily Didelphinae j IS. Supraorbital processes rudimentaiy (incip- Genus Philander Tiedemann, 1 808 I iently derived) Philander opossum Linnaeus, 1758 Excludes Lutreolina and Chironectes Philander andersoni Osgood, 1913

FIELDIANA: ZOOLOGY Cbironectes minimas Zimmerman

Fig. 3. Chironectes minimus Zimmerman. Plantar and dorsal surfaces of fiilly webbed right hindfoot.

Genus Didelphis Linnaeus, 1758 northernmost species, D. virginiana, occurs in Didelphis marsupialis Linnaeus, North and Middle America; the more southern D. 1758 {aurita Wied-Neuwied a syn- marsupialis, in Mexico and Middle and South onym) America; and D. albiventris, in South America. Didelphis albiventris Lund, 1840 The North American representatives of D. virgin- Didelphis virginiana Kerr, 1792 iana and D. marsupialis have been taxonomically Subfamily Chironectinae (new) revised by Gardner (1973).

Genus Chironectes Illiger, 1811 Distinctive characters of the family and sub- Chironectes minimus Zimmer- families are explicit in each of the 30 characters mann, 1780 described above. Detailed accounts of Chironec- Subfamily Lutreolininae (new) tes and Lutreolina await formal taxonomic revi- Genus Lutreolina Thomas, 1910 sion such as given below for Philander. Didelphis iMtreolina crassicaudata is highly derived. Its roots in the past are un- Desmarest, 1804 known. The animal that evolved a prehensile tail, opposable first digit, complete marsupium in Didelphis is the best known and most widely stages, and other derived characters is far re- distributed genus of American marsupials. Its moved in time from Didelphis. It is most unlikely

HERSHKOVITZ: PHILANDER as such in the kovitz, 1981:943—Philander valid generic name; that the genus Didelphis existed - type P. virginianus Tiedemann Didelphis opos- or even Late Tertiary. To refer to a Early Tertiary sum Linnaeus. Gardner, 1981:447—Philander val- as the stock staggered-lower-incisored Didelphis id; Husson opinion rejected. Rodger, 1982:270— from which the Early Paleocene Bolivian micro- testis and excurrent ducts;— paired spermatozoa. Jen- biothere Pucadelphys andinus Marshall and de kins and Knutson, 1983 type specimens in British Museum (Natural Perez-Hernandez, Muizon (1988) could be derived is anachronistic History). 1985:53, 65—generic characters. Perez- Hernandez and morphologically perverted. et al., 1986:14—dental morphology and diet. Gard- revision of the remain- A taxonomic Philander, ner (in Wilson & Reeder, eds.), 1993:22—synony- ing genus of the Didelphinae, follows. Compari- my; species (P. opossum, P. andersoni) distribution. sons are made with Didelphis, Chironectes, and Metachirops Matschie, 1916:262, 267, 268—included Lutreolina. Except for an occasional reference species pallidus Allen, fuscogriseus Allen, grises- Metachirm is dealt with in a here, separate paper cens Allen, melanurus Thomas, opossum Linnaeus, that is in preparation. canus Osgood, andersoni Osgood, quica Tem- minck (designated iy^6), frenata Lichtenstein; taxonomy. Miranda Ribeiro, 1936:340—characters. Krumbiegel, 1941:199—review. Pine, 1973:391— valid generic name for four-eyed pouched opos- Genus Philander Tiedemann sum; Philander regarded a —synonym of Didelphis. (Four-Eyed Pouched Opossums) Holothylax Cabrera, 1919:47 type Didelphis opossum Linnaeus by original designation. Metachirus Burmeister, 1854:135—subgenus of Di- Philander Brisson, 207-214—name from 1762:13, delphys [sic] Linnaeus, part, D. quica Temminck non-Linnaean work and not available. Gilmore, only. Burmeister, 1856:68—part, M. opossum Lin- 1941:316— Matschie antedated; char- Metachirops naeus only. Thomas, 1888:329—part, D. opossum fever acters; distribution; yellow susceptibility. Linnaeus only. Sonntag, 1924:743—comparative 1947:533—names from Brisson not val- Hopwood, tongue anatomy (M. opossum). Hill and Fraser, Philander Gronovius non-Linnaean and not id; 1925:196, PI. 1, Fig. 3 (urogenital organs), PI. 4, available. Figs. 15-17 (vagina)—female urogenital system. Philander 1808:426—included P. T\tdtm2iX\r\, species Boardman, 1952:848—hair tracts (A/, opossum). Tiedemann D. virginianus [= opossum Linnaeus], Lyne, 1959:84—vibrissae (M. opossum). P. murinus Marmosa P. [= murina], brachyurus Metacheirus [sic] Sanderson, 1949:787—misspelling [= Monodelphis brachyura]. Hershkovitz, 1949: of in combination with M. = Metachirus, opossum. 1 1 —type P. virginianus Tiedemann Didelphis opossum Linnaeus; Philander Brisson, 1762, and Philander Gronovius, 1763, not valid. Hildebrand, Type Species—Philander virginianus Tiede- = 1961:239—comparative body proportions. Peron- mann Didelphis opossum Linnaeus. dini and Perondini, 1965:381, Fig. 7 (metaphase chromosomes). Fig. 8 (karyogram)—chromosome Key to the Species (see also Fig. 4) = complement 2n 22. Enders, 1966:195—breeding season; habits. Biggers, 1966:251 —male external Dorsum uniformly dark or grayish, blackish mid- genitalia; regula- spermatozoan types; temperature dorsal stripe absent or poorly defined tion. Enders and Enders, 1969:431, PI. 1 (uterus P. opossum (p. 33) with 5 fetuses). Pis. 2-6 (uterus section)—placenta; Dorsum with well-defined blackish middorsal fetal membranes; fetal size at birth. Hayman and = Martin, 1969:192—chromosomes (In 22); chro- stripe or band P. andersoni (p. 33) mosome phylogeny. Collins, 1973:69—care and maintenance in captivity; reproduction; growth and Distribution—Figure 5. Tropical and subtrop- development; diet; parasites. Tyndale-Biscoe, ical forests including second growth, in South and 1973:25, 230— 38, 40, 61, 65, 189, 192, biology. Middle America, from Misiones and Chaco in Ar- Pine, 1973:391 —not the four-eyed pouched opos- gentina, and in Paraguay east of the Rio Paraguay, sum. Hershkovitz, 1976:295—history, nomencla- Brasil as far as Bahia on ture. McNab, 1978:115—comparative bioenerget- northward into eastern ics (P. opossum). Husson, 1978:27—treated as syn- the southeast, west from the Rio Tocantins-Ara- of Linnaeus of the onym Didelphis by designation guaia into Bolivia, and Peru north into Ecuador, Virginia opossum as lectotype of type species P. Colombia, Venezuela, the Guianan countries, Pan- virginianus Tiedemann. Landsmeer, 1979:337, Costa Figs. 1,4, 11, 12 (cheiridia, extensor assemblies)— ama, Rica, Nicaragua, Honduras, Salvador, digital extensor muscles; claw retraction. Hersh- Guatemala, Belize, and in Mexico, the state of

FiG. 4. Pigmentation of dorsum. A, Philander opossum; B, P. andersoni andersoni; C, P. andersoni mcilhennyi.

FIELDIANA: ZOOLOGY HERSHKOVITZ: PHILANDER

and Cowper (1704). His diagnosis, "Korper roth- Adult wild-caught animals that had been main- lich braun. Ueber jedem Augen ein gelbich weiser tained in cages and fed without stint in the now

flecken. Schwanz so lang als der Leib. 1 Fuss and defunct Rockefeller Laboratories in Rio de Janei- 3 2^11 lang ohne den Schwanz," is of a four-eyed ro became extremely large in body mass and cra- opossum, and Tiedemann's use of Didelphis opos- nial length. Dental size, however, fixed at com- sum Linnaeus in apposition definitely restricts the plete eruption, remains the reliable indicator of name to the pouched four-eyed species. The other "normal" body size. two species are "P. murinus {Did. murina L.)," a Basic Coloration (Fig. 4)—Trunk dark gray- Marmosa, and "P. brachyurus {Did. brachyuros ish or buffy agouti with or without a blackish Penn.)," a Monodelphis. middorsal stripe or band; face dark brown or Philander Tiedemann derives ultimately from blackish with contrasting supraorbital spot, rhi- Seba's (1734) vernacular names, '^Philander, narial tip usually unpigmented; tail dark brown or opossum s. Carigueja/' cited by Linnaeus (1758) with underside slightly paler, terminal half less in his description of D[idelphis]. opossum. Phi- pigmented or unpigmented. lander virginianus, a substitute name for D. opos- The Philander color pattern of a dark, modified sum Linnaeus, is type by subsequent designation agouti coat, dark brown to buffy head, dark brown

(Hershkovitz, 1949, p. 11). tail and ears, could give rise to all colors and color Didelphis philander Linnaeus, 1758, designated combinations present in didelphoids, including Lutreolina, and Chironectes. A dark type by Thomas (1888, p. 336), was not included Didelphis, brown is characteristic of all di- in the original erection of the genus Philander and eye ring nearly delphoids, and supraorbital spots such as those of hence is not valid as type. Philander expand into temporal bands in Chiro- Pine (1973, p. 391) and Husson (1978, p. 27) nectes and the whitish face of many Didelphis. argued against the use of Philander Tiedemann The more or less pigmented ears and tails of di- for the four-eyed pouched opossum and elected delphoids derive from fully pigmented ones. The Metachirops Matschie, a junior synonym, instead. saturate, bleached, or whitish cover hairs evolved The arguments of both authors were discussed from agouti cover hairs. Blackish (melanistic), and Hershkovitz (1976; 1981) and rejected by by grayish, orange, or reddish phases, whitish guard Gardner summarized and laid (1981, p. 44), who hairs and wool hairs, can be derived from the ba- the to rest. dispute sic color pattern of P. opossum. Coloration of body and limbs, tail, and ears of Lutreolina and the brindled body coloration of Chironectes can External Characters and Comparisons also be derived from a primitive Philander model. The saturate, pheomelanic facial pattern of Lu- Body Size and Form—Species of the genus treolina, without markings, however, is rare Philander average smaller than those of Didel- among didelphoids. The nearest common ances- phis, Lutreolina, and Chironectes but larger than tral facial pattern of both Lutreolina and Philan- all other living didelphoids. Average combined der must have been nearly uniformly modified head and body length is about 281 mm (slightly agouti. less in females than males); condylobasal length Color Phases—A "gray" or grayish brown of skull, an indicator of body size, is about 72 phase and a "brown," orange-brown, or orange mm. Trunk is elongate; muzzle long and slender; phase occur in both species of Philander. In the eyes large; ears conspicuous. Philander andersoni gray phase, the subterminal or pheomelanic bands averages slightly larger than P. opossum, and geo- of the cover hairs of dorsum and sides are graphically peripheral races of each species av- bleached to pale buff or nearly colorless. The gen- erage larger than more central ones. eral effect is a grayish brown appearance. In the Measurements—Table 1. brown phase, the subterminal bands are more Growth—Body mass increases after full erup- densely pigmented, giving the animal an ochra- tion of all molars. Largest individuals of a series ceous or orange appearance. Hairs of underparts or subspecies, however, are not necessarily oldest in gray-phase individuals are usually bicolor with and may include individuals classified as subadult the dark eumelanic bases showing through. In on the basis of dental eruption or wear and suture brown-phase individuals, the ventral hairs are closure. Males may not be larger than females at usually or predominantly pheomelanic to roots. birth but evidently outgrow females to maturity. The color term "brown," used by Mustrangi

HERSHKOVITZ: PHILANDER 11 of of Philander and of P. andersoni. Table 1 . Summary of measurements subspecies opossum subspecies Mea- surement means (extremes in parentheses) and number of specimens.

Head and Body Tail Hind Foot Ear

Philander opossum opossum Linnaeus

Philander opossum quica Temminck S 6 260 (200-300) 67 292(195-355)63 40 (29-48) 64 34 (22-42) 44 9 9 255 (220-330) 57 279(220-310)55 38 (32-46) 53 33 (22-43) 46 (J

Philander opossum fuscogriseus J. A. Allen 66 281(21 4-308) 73 289 (240-335) 73 45 (37-52) 72 36 (32-40) 57

9 9 276 (237-33 1 ) 36 284 (240-320) 36 42 (37-50) 36 37 (25-40) 28 6S99 279(214-331)109 287(240-335)109 44(37-52) 108 36 (25-40) 85 Philander andersoni andersoni Osgood S6 279(243-380)13 284(255-315)13 42(35-48) 15 37(30-45) 13 9 9 266 (225-300) 14 282 (255-332) 14 41 (35-51) 14 36(33-41) 12 6699 272 (225-380) 27 283 (255-332) 27 41(35-51)29 37 (30-45) 25

Philander andersoni mcilhennyi Gardner and Patton 6 6 299,279 276,313 43, 40 41, 35 9 9 293 (290-298) 5 292 (279-305) 5 41 (36-44)5 37 (35-42) 5 6 6 99 292 (279-299) 7 292 (276-3 1 3) 7 41 (36-44)7 38 (35-42) 7

(1994, p. 252) to describe a pale color phase of P. opossum is common among didelphoids and Marmosops incanus Lund (= M. scapulatus Bur- could give rise to any of the more derived types. meister), may be a tone of pheomelanin, or red- Molt—Changes in pelage with age or season dish. True melanin, or "blackish," is actually cannot be accurately determined on the basis of brown. The female Philander I captured in the material at hand. The few available large series of Parque Nacional do Caparao was reported by Philander opossum, such as the 85 specimens Mustrangi (1994, p. 253), who happened to see (amnh) from Ilha Taiuna, Para, Brasil, collected the prepared specimen while it was en route to between 31 October and 11 November 1931, pro- Chicago for my study. She described it as of "a vide no definite information on seasonal molt and similar, pale coloration" as the Marmosops incan- very little regarding ontogenetic change. Adults us of her report. The pheomelanic or erythristic in both old and new pelage without visible molt color phase and the melanistic phase are rare mu- line may occur at the same time in the same gen- tants among mammals and birds generally (cf. eral area. The fine, thin juvenal pelage is gradu- Hershkovitz, 1968; 1970; 1977; 1992a). ally replaced by coarser, thicker adult pelage. Ju- Pelage—Fur short in P. opossum, longer in P. venal coloration is like that of adults. andersoni, particularly in P. andersoni mcilhennyi, Limb Bones and Locomotion (Table 2)—Us- in which middorsal guard hairs are predominant. ing condylobasal length as an indicator of overall Downy wool hairs are preeminent in the aquatic body size, the cranial length ranges from about 29 Chironectes. Didelphis pelage is shaggy, the long mm in Dromiciops gliroides to about 1 1 2 mm in guard hairs and wool hairs about equally conspic- Didelphis marsupialis, a differential of about 85

uous and almost entirely concealing the cover mm. The hind limb is longer than the forelimb in , hairs. Pelage of LutreoHna is short with underfur, all taxa, and the tibia about as long as, or usually the silky guard hairs weak. A pelage like that of longer than, the femur. In the forelimb, the radius

12 FIELDIANA: ZOOLOGY Table 1. Extended.

Preorbital Postorbital ^ Condylobasal Zygomatic Braincase i Length Breadth Width Width Width

73.0(65.4-82.1)75 38.3 (33.0-43.4) 73 14.1 (11.5-18.9)59 8.8 (7.5-9.7) 74 21.2(18.4-24.0)72 69.7 (64.0-80.8) 39 34.5(31.1-40.7)34 13.1 (11.4-18.4)23 8.7 (8.0-9.7) 34 19.9(18.9-21.4)35 71.9(64.0-82.1) 114 37.0(31.1-43.4) 107 13.8(11.4-18.9)82 8.8(7.5-9.7) 108 20.8(18.4-24.0)107

66.4 (60.0-76.4) 78 35.5 (30.7-43.7) 75 12.6(10.8-15.7)36 8.4(7.4-9.1)77 20.2(18.2-23.1)71 62.8(57.1-72.4)76 31.1 (28.0-38.3)67 11.6(9.7-14.7)42 8.4 (7.5-9.9) 67 19.7(18.0-21.9)67 64.6(57.1-76.4) 154 33.4(28.0-43.7) 142 12.1 (9.7-15.7)78 8.4(7.4-9.9) 144 20.0(18.0-23.0) 138

69.6(64.6-71.9)8 36.6(33.8-38.1)8 —13.2(12.3-13.9)6 9.2 (8.7-9.7) 8 21.5(20.4-23.5)8 64.1, 65.8 33.5, 34.2 , 12.5 9.4, 8.4 20.9, 19.3 68.7(64.1-71.9) 10 36.0(33.5-38.1) 10 13.1(12.3-13.9)7 9.1 (8.4-9.7) 10 21.2(19.3-23.5)10

72.7(61.9-81.5)76 38.4 (32.2-43.0) 73 13.4(10.9-15.7)61 9.0 (8.3-9.5) 73 21.0(19.1-23.5)75 69.7(60.0-80.5)41 36.1 (30.7-43.1)40 12.9(11.5-16.6)31 8.9 (8.0-9.4) 40 20.6(18.7-23.6)40 71.6(60.0-81.5) 117 37.6(30.7-43.1) 113 13.2(10.9-16.6)92 8.7(8.0-9.5) 113 20.9(18.7-23.6) 115

72.3 (66.7-76.5) 17 37.1 (32.3-39.9) 17 13.6(11.8-16.2) 13 9.3(8.0-9.4) 17 21.2(19.6-22.6) 16 68.9(64.6-81.2) 17 34.1 (30.8-40.0) 17 12.6(10.4-14.0) 13 8.8(8.0-10.1) 18 20.6(19.3-23.9) 17 70.6(64.6-81.2)34 35.6 (30.8-40.0) 34 13.1 (10.4-16.2)26 9.1 (8.0-10.1)35 20.9(19.3-23.9)33

74.4,73.1 37.6, 36.5 14.0, 13.5 9.0, 8.4 21.2,20.8 71.8(70.2-73.9)4 35.8 (34.2-36.8) 4 13.7(12.7-14.7)4 9.0 (8.7-9.2) 4 20.7(20.2-21.0)4 72.5 (70.2-74.4) 6 36.2 (34.2-37.6) 6 13.7(12.7-14.7)6 8.9 (8.4-9.2) 6 20.8(20.2-21.2)6

is about as long as the humerus or shorter. In the ronectes is preeminently adapted for aquatic life, semiaquatic Chironectes it averages less. its long legs and palmate hind feet serving for pro- The hind limb is longer than the forelimb in the pulsion through water, the gritty palms and digits Marmosidae, with the greatest differential shown of the front feet for seizing slippery prey and by the Patagonian Lestodelphys halli, apparently climbing slippery rocks and boulders in mountain a leaper or springer. It is closely followed in limb- streams. Limbs of Lutreolina are short and weasel- to-trunk proportions by the terrestrial Metachirus like and apparently better adapted for terrestrial lo- nudicaudatus. No American marsupial is a bur- comotion than for climbing or swimming. rower. Chironectes is the only living American Cheiridia (Fig. 6)—Hands and feet of Philan- marsupial adapted for aquatic life, but all Amer- der are modified for grasping, the pollex and hal- ican marsupials are good swimmers and divers, lux opposable, the latter inunguis; fifth hind toe except possibly the Marmosidae. All are habitual much shorter than fourth, three middle digits sub- tree climbers, except Lutreolina, Metachirus, Chi- equal; toes not webbed. Manual and pedal claws ronectes, Lestodelphys, and Monodelphis, all of of Didelphis stout, hallux inunguis, opposable; which probably can climb but have not been ob- claws sharp, recurved; digits of Lutreolina as in served doing so. Didelphis but claws blunt, little recurved. In Chi- The long hind limbs of Philander are well adapt- ronectes manual digits with short needle-like ed for running and springing. The proportionately claws, hind feet with toes webbed, hallux inun- shorter limbs of Didelphis, especially the anterior, guis (Fig. 3). are better specialized for climbing. Animals of both Tail (Fig. 7)—Tail in Philander and Didelphis genera progress with facility on the ground. Phi- is cylindriform, tapered, prehensile, scaly, practi- lander with moderate to rapid speed and Didelphis cally bare except for densely furred basal 3-9 cm; slowly and with a lumbering gait that does not ex- naked ventrally, the terminal portion plicate be- clude a swift sprint by a frightened animal. Chi- tween vertebral joints to permit clasping, length

HERSHKOVITZ: PHILANDER 13 Table 1. Extended.

Palatal Length

Philander opossum opossum Linnaeus 66 44.0(40.3-49.6)75 38.0(34.9-41.5)75 13.8(12.8-14.9)75 9 9 4L9 (38.4-45.8) 35 36.8 (34.8-39.3) 35 13.7(12.7-14.7)36 SS9 9 43.3(38.4-49.6)110 37.6(34.8-41.5) 110 13.8(12.7-14.9) 111

Philander opossum quica Temminck SS 38.3(34.1-44.8)75 34.4 (32.0-37.7) 78 12.7(11.6-14.4)80 9 9 39.2(35.1-44.6)71 32.9(30.5-38.0)71 12.5(11.2-14.5)78 dd99 38.7(34.1-44.8)146 34.2 (30.5-38.0) 147 12.6(11.2-14.5) 158 Philander opossum melanurus Thomas 6 6 42.0(38.6-43.1)8 36.5 (34.7-37.5) 8 13.6(13.3-14.0)8 9 9 38.8,39.9 34.6, 34.8 13.0, 13.0 6699 41.4(38.6-43.1)10 36.2 (34.6-37.5) 10 13.5(13.0-14.0) 10

Philander opossum fuscogriseus J. A. Allen 66 44.2(37.9-50.2)74 38.1 (34.3-41.8)77 14.2(11.7-15.6)77 9 9 42.5(37.0-48.7)41 36.9(33.9-41.2)42 13.9(12.8-15.3)42 6699 43.6(37.0-50.2)115 37.7(33.9-41.8) 119 14.1 (11.7-15.6) 119 Philander andersoni andersoni Osgood 66 43.4(39.7-48.9)20 38.0 (35.2-43.3) 20 13.9(12.0-16.0)20 9 9 42.0(39.7-49.6)18 36.9(34.2-42.9) 18 13.8(12.6-15.0) 18 6 69 9 42.8(39.7-49.6)38 37.5 (34.2-43.3) 38 13.8(12.0-16.0)38

Philander andersoni mcilhennyi Gardner and Patton 6 6 45.1,44.5 39.8, 38.4 15.3, 14.6 9 9 43.5 (42.0-44.7) 4 38.1 (36.4-39.7)4 14.4(13.7-15.3)5 6 699 44.0(42.0-45.1)6 38.5 (36.4-39.8) 6 14.6(13.7-15.3)7

about equal to that of combined head and body; from the sacrum, and the 5th (or 6th) usually rhomboidal scales arranged spirally; the interscu- sharply differentiated from the succeeding caudal tular bristles usually 5 per scale (Fig. 8). The Di- vertebra. Muscles of the transitional vertebrae delphis tail is more strongly prehensile than that move the tail in particular directions. The action < of Philander, and that of the aquatic Chironectes of a specific muscle may be narrowly restricted, is hardly if at all prehensile, its flattened ventral such as that controlling movements of the tail tip. surface serving as a rudder. The Lutreolina tail is External Ear (Fig. 1)—The auricular lamina the most densely hirsute of the group. Length of Philander is large, rounded, leafy or membra- slightly less than head and body combined and, nous and plicate, entirely blackish or with base of as in Philander, may be nearly the primitive pro- middle portion to total unpigmented; length, when portion for mammals generally. The tails are not laid forward, reaching outer canthus of eye. Ex- fat storing, but that of terrestrial Lutreolina is no- ternal ears of Didelphis and Chironectes are sim- tably thick at base, where the transitional verte- ilar. The Lutreolina auricular lamina is much brae provide insertion for large muscle mass (Fig. smaller, less leafy, hairy, and hardly if at all pli-

1). (More on locomotion in Philander on p. 75.) cate. Auricles of the group could have evolved Transitional Caudal Vertebrae (Table 3)— from a type with lamina little expanded, bare, and Transitional vertebrae of the postcranial skeleton, perhaps slightly motile or plicate. Didelphoid au- usually 4 or 5 in American marsupials, sometimes ricles are comparable to those of many species of 6 depending on interpretation, are present be- shrews, lemurs, and callitrichids (cf. Hershkovitz,) tween the sacrum and typical caudal (coccyxial) 1977, pp. 102-106). vertebrae. Although well differentiated (Fig. 9), Marsupium—A fully formed pouch character- they are usually counted as caudal. The morpho- izes sexually mature females of Philander and Di- logical characters of the transitional vertebrae, delphis and both sexes of the semiaquatic Chiro- i

' however, are well marked, with the first distinct nectes. The pouch opens from the side in the first

14 FIELDIANA: ZOOLOGY "^ •^ Tt On

•- 1 9 3 06X

P « ^^ H 2 « y 0ilfr/¥m'

16 HELDIANA: ZOOLOGY skull length and 60% midline length from tips of ger than that of Philander, sagittal and nuchal nasals to lambdoidal crest; combined tips of pre- crests heavier, postorbital region more elongate, maxillary bones rounded, extending less than 1 braincase more deflected relative to palatal plane, mm beyond base of i". frontal and maxillary sinuses more inflated, muz- Distance from lacrymal foramen to anterior zle relatively shorter, nasals less than 38% greatest edge of glenoid fossa less than zygomatic breadth; skull length and less than 55% midline length supraorbital borders square or rounded, not ridged from nasal tips to lambdoidal crest. or overhanging; postorbital constriction narrower In didelphoids generally, as growth continues than preorbital constriction; well-defined postor- after sexual maturity, crests enlarge and most cra- bital processes short, bluntly pointed; temporal nial proportions shift. ridges uniting behind to form a low sagittal crest connected with a well-developed nuchal or lambdoidal crest; shallow pit or depression at junction Dental Characters and Comparisons of temporal ridges; cranial crests developed early (Table 6, Figs. 13-20) in both sexes.

Palatal vault not markedly arched, its contour The following dental formula for didelphids is nearly flat; palatal extension behind transverse plane the same for all living and known extinct didel- of last molars equal to combined length of last 2 or phoids: 2W molars; paired incisive foramina long, extending 1,2,3,4,5 1 1,2,3 1,2,3,4 behind anterior plane of canines; paired maxillopa- m- (0,2,3,4,5' 1,2,3' 1,2,3,4 latal vacuities long, slitlike and extending fi'om about the level of m' to that of m^; paired postero- The same formula has persisted unchanged palatal vacuities usually present, short, ovate, ex- since no later than Early Cretaceous and likely tending from or behind plane of last molars to raised Early Jurassic. Loss of i,, shown in parentheses, posterior palatal border; posterior choanae narrow, and the staggered or crowded position of i, (Figs. width measured across base of posterior pterygoid 12, 13, 14, 16), evidence of Mesozoic mandibular less than 10% of processes usually condylobasal contraction, are didelphoid hallmarks. The meta- length; tripartite auditory bulla, of tym- composed cone of m'~^ is consistently larger than the para- of bone, from panic wing alisphenoid separated cone. The size relationship between metacone and of bone with tympanic wing petrous ectotympanic paracone in eutherians varies within families and bone between, the suspended components separate sometimes individually. or but not fiised 1992a, touching (Hershkovitz, Fig. According to Luckett (1993), the deciduous or 10, 25; Hershkovitz, in p. press). primary teeth of incisors and canines are nonfunc- Comparisons (Fig. 2)—Skull of Didelphis rel- tional and vestigial. The functional incisors and ca- atively broader, the frontal and sinuses maxillary nines are successional or second generation. The more inflated, crest bladelike. Skull sagittal high, functional premolars are mixed: pi and p2 are the of Chironectes heavier, broader, braincase width unreplaced deciduous teeth; p3, the last tooth to about 35% of condylobasal length (less than 30% erupt, succeeds a functional, fully molarized decid- in Philander); angle between sagittal and frontal uous p3. This tooth is also the first to erupt. Molars planes greater, nasals relatively shorter, less ta- are unreplaced primary teeth (Fig. 14). pered behind, the terminal as in Lu- points spread The sequence of dental eruption revealed by di- treolina; breadth wider, zygomatic greater, palate delphids at hand and by studies of prenatal and the posterior vacuities rectangular, posteromedian early postnatal Didelphis virginiana by McCrady vacuities not usually absent; occipital condyles (1938) and Berkovitz (1978) indicated the follow- projecting behind vertical of plane supraoccipital ing (teeth enclosed by parentheses erupt at about bone, their combined width about half that of pal- the same time): ate between molars; turbinate bones enlarged and i5, il, (c, ml, m4 produced behind to nearly posterior border of pal- dp3, p2, i(2^), pi), m2, m3, p3, atal vacuities. Lutreolina braincase relatively lon- dp3, p2, i(2^), i5, -, (c, pi), ml, m2, m3, p3, m4

Fig. 6. Cheiridia of Philander opossum. A, Right hand, dorsal and plantar aspects; B, right foot, dorsal and plantar aspects.

HERSHKOVITZ: PHILANDER 17 plica

plica ventral Philander opossum

ventral

Dideli^s marsupialis

dorsal

ventral ventral Chironectes minimus

iutreolina crassicaudata

Fig. 7. Basal (dorsal) and terminal (dorsal and ventral) caudal segments of Didelphis, Philander, Chironectes, and Lutreolina. Plicas are cutaneous folds of prehensile tails.

Descriptions of Individual orthodont or slightly prodont, the opposing pair Teeth of Philander convergent for about two-thirds their length, their pointed tips often divergent; unworn i" uniform- Upper Incisors (Fig. 15)—Five in number, first ly premolariform (tridentate), crowns slightly to (i') usually largest, sometimes as small as any of broadly overlapping except in old, worn teeth; i^

i' i^-^; most distinctive of series, subcylindroform, from half to as high as i', slightly less than that

18 FIELDIANA: ZOOLOGY Fig. 8. Section of Philander opossum tail, ventral surface, showing spiral arrangement of scales and interscutular bristles, usually five per scale.

of a small i' i''*; diastema present between and i^; ond (phylogenetic i,) staggered between ij and i4 mesiostyle (a or A) of i" weak, distostyle (b or as in all didelphoids (Figs. 13, 14, 16). E) less defined or obsolete. Remarks—Marsupial dental evolution appears Lower Incisors—Four in number (2-5), the to have been most manifest in the incisor field. first all in phylogenetic lost; uniform structure but The staggered i, marks divergence of the Didel- decreasing in size from numerical first (iz) to last phimorphia from the basic metatherian stock. (is); all incisors markedly recumbent seen from Only microbiotheres, including Pucadelphys an- lingual aspect; crowns usually well separated in dinus Marshall and de Muizon, 1988 (Marshall et adults but sometimes touching in young; each al., 1995), of the Early Paleocene of Bolivia are tooth with labial surface more or less spatulate, the known members of the unstaggered-incisor- crown ovate, root cylindrical, tapered; lingual sur- toothed Microbiotheriomorphia (cf. Hershkovitz, face smooth, the median ridge or torus broadly 1992; 1995). sloping; occlusal groove often present on lingual Upper Canine—Long, slender, smooth, re- surface; terminal styles obsolete; numerical sec- curved with axis directed slightly forward, its

Table 3. Vertebral count of largest American marsupials. Number of transitional vertebrae also included in count of caudal vertebrae as customary. Number of cervical vertebrae is consistently 7, of sacral, 2, in all marsupial taxa examined. Sample number in parentheses.

Taxon growth continuous past sexual maturity; cingulum and accessory conules absent; diastema between canine and last incisor approximately equal to or less than greatest diameter of canine. Lower Canine—Shorter, weaker than upper, /\_™ more or less recurved, its bulk about half that of upper; well differentiated from incisors, about twice their height, without diastema. Upper Premolars—Teeth two-rooted, unicus- pidate with well-developed ledgelike lingual and buccal cingula; first premolar about half or less /sacra] the bulk of second, the diastema between approximately equal to crown length in old adults, about one-half to one-third crown length in young adults; middle premolar larger than last, often ^— 1 equal, infrequently smaller; unworn first premolar with terminal styles usually present and well defined; middle premolar with terminal styles poorly defined or absent except style b or E, the mesio- a or less 1—2 style (parastyle, A) frequently present than the distostyle {b or £); eruption of replace- ment p^ precedes that of m"*, styles poorly differentiated or absent; fmnh 1 14685 with an incipient but well-defined metacone. /— 3 Upper Deciduous Third Premolar (the only upper deciduous tooth) (Fig. 17)—Size about equal to that of m^, general outline and morphol- P: -4 ogy as in m', conules A, B often well developed, conule C larger than B, D greatly reduced or absent, E indicated. Lower Premolars—Teeth two-rooted, unicus- pidate with ledgelike lingual and buccal cingulids ;ni poorly defined, talonid basin simple; lower first premolar similar in size and shape to upper first and likewise separated by diastema from middle premolar, distostylid {b or E) more or less defined, mesiostylid {a or A) obsolete or absent; middle premolar as high or higher than incisors and nearly as high as canine, distostylid present, mesio- caudal stylid minuscule; last premolar like middle but smaller, buccal cingulum usually poorly defined, talonid basin slightly more developed; permanent p, erupts before m4. Lower Deciduous Third Premolar (the only lower deciduous tooth) (Fig. 17)—Like first per-

FiG. 9. Transitional caudal vertebrae 1-5 (untinted) j

of Philander opossum (fmnh 60501) between tinted sa- 1

cral and first typical caudal (coccyxial) vertebrae. Tran- > Philander sitional vertebra 5 differs notably from both the caudaJ opossum j FMNH 60501 behind and transitional 4 ahead. "

20 FIELDIANA: ZOOLOGY Table 4. Nipple formulae of the largest didelphoids. Number of samples more than one shown in parentheses; specific names are those of cited source; formulae from Hershkovitz field notes are of captured animals, the prepared specimens preserved in fmnh or usnm; other preserved specimens in the museum collection are shown as fmnh.

Prime Nipple Formulae [Functional formulae Taxon in brackets] Source

DiDELPHIDAE Fig. 10. Philander opossum, dorsal and ventral aspects, showing points of cranial measurements, a, nasals; b, greatest length of skull; c, least interorbital breadth or postorbital constriction; d, zygomatic breadth; e, postorbital width; f, braincase, greatest width; g, condylobasal length; h, basal length; i, palatal length;], length upper tooth row (i'-m'*); k, length upper molar row (m' *).

soni have not been observed, but geographic dif- the genus, however, reveals that variation from ferentiation does occur. In a comparison of ''Phi- present to absent, or degree of molarization of lander mcilhennyi" from Balta, Peni, with sym- conule C, is about the same for both species (Ta- patric Philander opossum quica, Gardner and ble 7). Patton (1972, p. 3) noted a relatively deep inden- Conule C, like any other derived excrescence tation of the labial margin of m^ present in the of the buccal shelf, is firmly established, cusplike, first species but not in the second. The indenta- and usually subequal to conule B in m' and m^. or is it varies tion, so-called "notch," the angle or valley In m\ however, geographically from a 1 between conule C cone C) and rudiment or to a fair-sized (stylar adjacent anlage cusp nearly ]

conule B (Fig. 19). equal in bulk to conule B (Fig. 19). Its molariza- I In four at hand of Balta Philander tion increases from the most specimens gradually primitive ! andersoni mcilhennyi, conule C of m"* is well de- and geographically central forms of P. andersoni '

veloped or molarized in three females but quite and P. opossum to their most derived or geo^ ;

worn in the single male. In 8 adults of sympatric graphically peripheral representatives. . Balta P. I is opossum quica, find conule C distinctly Comparisons—Dentition of Didelphis essen- 1 less developed in three females and like that of Philander. That of Chironectes rudimentary tially j or virtually absent in 5 males. Examination of all is generally heavier, average transverse width oft available specimens from throughout the range of m^ about 6 nmi (4.5 in Philander), arcade of up- '

22 FIELDIANA: ZOOLOGY Fig. 1 1 . Diagram of didelphid skull, ventral and dorsal aspects, showing topographic features. See Table 5 for explanation of symbols.

per and lower incisors more broadly curved with AMNH 72017, adult 9, Rfo Curaray, Loreto, second lower (i,) less abruptly staggered; unworn Peru incisors of Chironectes triangular in lateral out- Left supernumerary m', slightly smaller line, but shape more nearly equilateral than ob- than normal m', rotated 180° with buc- tuse; upper incisors of Chironectes and Lutreolina cal shelf lingual. notably inflected inward and upward; first pre- Philander opossum fuscogriseus molars less reduced, gap between first and middle AMNH 34373, adult 6, Bagadd, Choc6, Co- premolars narrower in Chironectes, varying from lombia less than half the crown length of anterior tooth Left supernumerary m^, fully erupted, to absent; molar crowns more hypsodont in Chi- smaller than normal m'*, otherwise ronectes and Lutreolina, outline of occlusal sur- slightly similar. face more nearly square or rectangular than tri- Right supernumerary m^ nearly fully angular, the outer posterior angle, particularly of erupted but impacted and rotated about 90° m\ less produced. with buccal side anterior and overlapped by posterior portion of normal m". Lower molars normal in number and form. Dental Abnormalities Philander opossum quica Supernumeraries i^UMZ 12009, adult $, Balta, Ucayali, Peru Philander andersoni andersoni Right supernumerary pm', in diastema be-

HERSHKOVITZ: PHILANDER 23 Table 5. Explanation of cranial symbols 1-23 and talonid presumably a result of crowding by a-z in Figures 1 1 and 1 2 A. developing supernumerary. \ FMNH 114702, adult 6, San Ramon, El Beni, Bones Bolivia 1. Nasal Right supernumerary i,, behind normal i,, 2. Frontal occludes with normal i'; normal oc- 3. Parietal i; cludes with i^ and so on to which bites 4. Supraoccipital ij, 5. Premaxillary into diastema between i' and i,; position 6. Maxillary anteriad of i, more medial and compared 7. Lacrymal to left homologue; jaw, teeth, and occlu- 8. Jugal, zygomatic sion otherwise normal. 9. Squamosal, temporal 10. Sphenoid (includes orbitosphenoid, alisphenoid, FMNH 114685, 9 presphenoid, basisphenoid, pterygoid) Left upper premolar with incipient but 1 1 . Palatine well-defined metacone. 12. Orbitosphenoid 13. Pterygoid process Missing molar 14. Presphenoid Philander opossum opossum 15. Basisphenoid USNM adult Brasil 16. Alisphenoid 393600, 9, Utinga, Para, 17. Alisphenoidal wing of auditory bulla m'' missing; no sign of alveolus but arcade 18. Ectotympanic otherwise normal; m4 normal; teeth mod- 19. Periotic wing of auditory bulla erately worn. 20. Mastoid (temporal) 21. Basioccipital 22. Occipital condyle 23. Exoccipital Sexual Dimorphism Foramina, Fissures, Processes, Fossae, and Crests a. External nares Mature males with four molars fully erupted b. Infraorbital foramen average larger than comparable females in body c. Lacrymal foramina or canals skull and canine size. Postcanine d. Canine fossa mass, length, teeth also but relative to skull e. Posterolateral vacuity or foramen average larger, f. Sphenorbital fissure (concealed above II) length no larger than those of females or even Foramen rotundum g. smaller; the braincase is relatively narrower in h. Foramen ovale males, cranial crests more developed, but the sag- i. Tympanic membrane and auditory meatus (see 18) ittal crest of old males is sometimes equaled by j. Postglenoid foramen k. Stylomastoid foramen that of extremely large or old females. 1. Jugular foramen m. Hypoglossal foramen and or condylar foramen n. Carotid foramen or canal o. Anterior lacerate foramen or petrotympanic fissure Karyology p. Foramen magnum Glenoid fossa q. Descriptions of karyotypes of the present and r. Premaxillary or incisive foramen supposed species of Didelphidae (Table 8) were s. Maxillopalatine or mesolateral vacuity first in 20 All t. Posteromedial or palatine vacuity published nearly papers. publica- u. Ascending postorbital (zygomatic) process tions were critically reviewed, with new data add- V. Ascending postorbital (zygomatic) process ed by Reig et al. (1977). The authors studied and w. Postglenoid process described the chromosomes of 177 species ol X. Temporal ridge American marsupials representing nine gener? y. Sagittal crest z. Lambdoidal crest and 22 species, including those considered in thi^ .

report (Table 8). To avoid repetition and extr* I limital discussion, only Reig et al. (1977) is cite(j

* tween normal first and second premolars, here for documentation. ! fully erupted, normal in appearance except The "standard" karyotype as understood bji rotated about 100° with border et al. is in all treated as di posterior Reig (1977) present { linguad and slightly anteriad; heel with delphids with few minor exceptions. All listed ii as in Table 8 share the number of 22. The fun pronounced distostylid (hypoconulid) diploid | left pm'. "Normal" first premolar without damental number is 20, except in D. virginianal

24 FIELDIANA: ZOOLOG^ Fig. 12. A, Diagram of left side of Philander opossum skull; B, left mandible, buccal side; C, lingual side; D, posterior view. See Table 5 for names of cranial features and below for those of buccal and lingual sides of mandible: a, horizontal ramus; b, ascending ramus and masseteric fossa; c, symphysis; d. angular process; e, condyloid process; f, coronoid process; g, superior notch; h, inferior or lunate notch; i. superior masseteric line; j, inferior masseteric line; k, mylohyoid line (not shown); I, horizontal masseteric line; m, mental foramen; n. mandibular foramen.

where it is 32. Sex chromosomes are acrocentric parasites usually leave a live caged animal shortly in all but are metacentric in the X chromosome after capture and desert the host almost inmiedi- of D. virginiana and L crassicaudata. It appears ately at death. that the most variable complement among the four The list of parasites is not exhaustive. It was genera of Table 8 is D. virginiana. compiled mainly as a source of reference for discussions on marsupial zoogeography. The taxonomic names used are those of the sources cited. Parasites Viruses

Captured marsupials are mostly parasitized by Mucambo (Potkay, 1977; Pard, Brasil; fleas, lice, and staphylinid beetles. These mobile Shope, 1967b (in Potkay, 1977, q.v.))

HERSHKOVITZ: PHILANDER 25 Fig. 13. Skull of Philander opossum. Upper row, dorsal, ventral, lateral aspects; middle row, anterior, posterior and left lateral — aspects i, pm, (note staggered i, with buttress); bottom row, two posterior portions of basicranium (see Figs. 11, 12 and Table 5 for identification of parts).

Venezuelan equine encephalitis (Almirante, Group B (mosquito borne) Panama; Grayson & Galindo, 1968, in Ilheus (Bahia, Brasil; Laemmert, 1967) Potkay, 1977) Yellow fever (Bahia, Brasil; Laenmiert

Eastern equine encephalitis (Bahia, Brasil; al., 1946) Shope et al., 1966, in Potkay, 1977) I: St. Louis encephalitis (Bahia, Brasil; Shope Group C et al., 1966) Itaqui (Bahia, Brasil; Shope, 1967a)

26 FIELDIANA: ZOOLOGli Fig. 14. Upper and lower jaws with tooth rows of Philander opossum.

Capim Group Sparganum reptans (Imperial Bureau of Ag- Acara (Bahia, Brasil; Belem Virus Labo- ricultural Parasitology, 1933) ratory, 1976c) Protozoa Mosquito-borne group Babesia brasiliensis and Kikuth Turlock (Bahia, Brasil; Shope et al., 1966) Regendanz (Brasil; Regendanz & Kikuth, 1928, p. Phlebotomus-bome 1567) Itaporanga (Bahia, Brasil; Trapp & Shope, Sarcocystis garnhami (British Honduras; 1967) Lainson & Shaw, 1969) cruzi Wood &. Vescicular stomatitis group Trypanosoma (Brasil; Wood, 1941; Costa Rica; Deane, 1961, 1964) New Jersey strain (Panamd; Tesh et al., 1969) Trypanosoma rangeli-like (Pard, Brasil; Deane, 1964) Indiana strain (Panamd; Tesh et al., 1969)

Unknown vectors Fungus Piry (Bahia, Brasil; Belem Virus Labora- Histoplasma capsulatum (Canal Zone, Pana- tory, 1967a) md; Taylor & Shacklette, 1962) Pacui (Bahia, Brasil; Bel^m Virus Labo- ratory, 1967b) Mallophaga (lice)

Oochoristica braziliensis (Colombia; Baer, Gliricola porcelli (S§o Paulo, Brasil; Hop- 1927) kins, 1949) lERSHKOVrrZ: PHILANDER 27 A

Philander opossum Caluromys philander FMNH 16398 FMNH 21725

Philander opossum Lutreolina crassicaudata FMNH 11A698 FMNH 5394A

Philander opossum Metachirus nudicaudatus FMNH 11A711 FMNH 207%

Fig. 15. Upper incisors and canine of Philander opossum compared with outgroups. A, P. opossum (fmnh 16398), i'"' variably convergent proximally, divergent distally; Caluromys philander (fmnh 21725), i'' convergent to tip. B, P. opossum (fmnh 1 14698), premaxillary fossa for lower canine shallow, ventral margin of premaxillary bone nearly horizontal; Lutreolina crassicaudata (fmnh 53944), premaxillary fossa for lower canine deep, ventral margin upturned, premaxillary bone slender, inclined. C, P. opossum (fmnh 1 1471 1), outer margins of nares steep, incisive distostyles obsolete or absent; Metachirus nudicaudatus (fmnh 20796), outer margins of nares gently sloping, nasal tips produced forward, distostyles present, the incisors trident.

Gryopus ovalis (Sao Paulo, Brasil; Hopkins, Amblyomma auricularium Conil (Panama; ! 1949) Fairchild et al., 1966, pp. 191, 209)

j Trimenopon hispidium (Sao Paulo, Brasil; Amblyomma geayi Neumann (Panama; Fair* et child i Hopkins, 1949) al., 1966, pp. 195, 209) , Fairchild et Amblyomma sp. (Panama; al.^ Jones et 1966, pp. 209; Venezuela; al., | Acarina (mites, ticks, chiggers) 1972) Archemyobia pectinata Mendez (Panamd; Androlaelaps fahrenholzi Belese, 1911 (Ven^ Mendez, 1972, p. 615) ezuela; Furman, 1972)

28 FIELDIANA: ZOOLOGY The staggered marsupial incisor

staggered alveolus

I mm buttress PHILANDER OPOSSUM

Fig. 16. Alveolus of and staggered ij, buttress, in Philander opossum (from Hershkovitz, 1982).

Crotiscus disdentatus Boshell and Kerr (Pan- Haemolaelaps glasgowi Ewing (Panam^; Brennan amd; & Yunker, 1966, p. 260) Tipton et al., 1966, p. 34) Brennan Pseudoschoengastia bulbifera (Pan- Nematoda am^; Brennan & Yunker, 1966, p. 260) Trombicula dunni Ewing (Panamd; Brennan Aspidodera sp. (Venezuela; Guerrero, 1985) & Yunker, 1966, p. 260) Capillaria sp. (Venezuela; Guerrero, 1985) Trombicula keenani Brennan and Yunker Travassastrongylus sp. (Venezuela; Guerrero,

(Panamd; Brennan & Yunker, 1966, p. 260) 1985) Eutrombicula alfreddugesi Oudeman, 1910 Moennigia sp. (Venezuela; Guerrero, 1985) (Venezuela; Brennan & Reid, 1974; Pana- Strongvloides sp. (Venezuela; Guerrero,

md; Brennan &. Yunker, 1966, p. 260) 1985) Eutrombicula goeldii Oudeman, 1910 (Ven- Viarinaia barusi Guerrero, 1983 (Venezuela; ezuela; Brennan & Reid, 1974; Panamd; Guerrero, 1983) Viannaia Brennan & Yunker, 1966, p. 260) conspicua (Brasil; Imperial Bureau Eutrombicula tropita Ewing, 1925 (Venezue- of Agricultural Parasitology, 1933, ex Pot- la; Brennan & Reid, 1974) kay, 1977) Euschoengastia nunezi Hoffmann (Panam^; Rhopalia horridus (Angra dos Reis; Imperial Bureau of Brennan &. Yunker, 1966. p. 260) Agricultural Parasitology, 1933, Ixodes lasallei Mendez and Ortiz, 1958 (Ven- ex Potkay, 1977) ezuela; Jones et al., 1972) Viannaia minispicula Guerrero, 1983 (Vene- Ixodes luciae Senevet, 1940 (Venezuela; zuela; Guerrero, 1983) Jones et a!., 1972) Viannaia skrjabini Lent and Freitas, 1937 Ixodes venezuelensis Kohls, 1953 (Venezue- (Venezuela; Guerrero, 1983) la; Jones et al., 1972) Viannaia tenorai Guerrero, 1983 (Venezuela; Ixodes luciae Senevet (Panam^; Fairchild et Guerrero, 1983) Viannaia vianniai al., 1966, p. 209) Travassos, 1914 (Venezue- Tur apicalis Furman and Tipton, 1961 (Ven- la; Guerrero, 1983) ezuela; Furman, 1972) Cestoda Tur uniscutatus Turk (Panama; Tipton et al., Linistowia iheringi (Brasil; 1982, 1966, pp. 41, 42) Beveredge, 107) Omithonyssus wemecki Fonseca (Panam^; p. Yunker & Radovsky, 1966, p. 92) Uniramia (Pterygota)

HERSHKOVITZ: PHILANDER 29 Fig. 17. Occlusal surface of deciduous premolar 3 and molars 1 and 4. A, Left dp"* and m' of Philander opossum (FMNH 55411). B, Right dpj and mi of P. opossum (fmnh 66332). C, Left m^ (composite fmnh 90086, 14015).

Coleoptera Adoratopsylla intermedia copha Jordan (Pan- ama; Tipton & Mendez, 1966, p. 326; SW Amblyopinus henseli (Serra dos 6rgaos, An- Colombia; Mendez, 1977, p. 166) gra dos Reis, Brasil; Tijuca; Seevers, 1955, Adoratopsylla antiquorum cunhai Pinto (Bra- p. 247) sil; Rio de Janeiro, Costa Lima & Hatha-

way, 1946, p. 228) Siphonaptera (Fleas) Ctenocephalides felis Jordan and Rothschild Adoratopsylla intermedia intermedia Wagner, (cosmopolitan; Costa Lima & Hathaway, 1901 (Venezuela; Tipton & Machado, 1946, p. 213) 1972) Xenopsylla cheopsis Rothschild (cosmopoli-

30 FIELDIANA: ZOOLOGY Rhopalias caballeroi Kifune and Uyema (Peni; Tantalean et al., 1992) Rhopalias baculifer Braun (Peni; Tantalean et

al., 1992) Paragonemus amazonicus Miyazaki, Grades, and Uyema (Peni; Tantaledn et al., 1992) Rhopalias coronatus Rudolphi (Peni; Tanta- ex Simpson lean et al., 1992) Amphimeruse ruparupu Kifune and Uyema (Peni; Tantaledn et al., 1992)

Fig. 18. Conules terminal (stylar cusps, ectostyles, Collins (1973, pp. 4, 74) listed the following styles) of m\ 1, buccal view from Bensley (1906, p. 6); genera of parasites of Philander without more in- 2, occlusal view from Simpson (1929, p. 1 19); 3, buccal formation or for affinities. view from Hershkovitz (1977, p. 287); arrow points to regard phylogenetic ectoflexus. See Tables 6-7 for explanation of symbols. The list was culled from the "host catalog index card file of the Parasite Classification Index Cat- alogue of the United States Department of Agri- culture's Parasite Control Center, Beltsville, tan; Costa Lima & Hathaway, 1946, p. 228) Maryland." Polygenis roberti beebei I. Fox (Panamd; Arthropoda Pseudoschongastia, Tipton & M^ndez, 1966, p. 326) flea Polygenis klagesi klagesi Rothschild (Brasil: Adoratopsylla, chigger tick flea Pard, Amazonas, Colombia; Panama; Ven- Amblyomma, Pulix, Ctenocephalides, flea Rhopalias ezuela; Costa Lima & Hathaway, 1946, p. Crotiscus, flea 142) chigger Rhopalopsyllus, Rhopalopsyllus australis tupinus Jordan and Euschoengastia, Schongastia louse Rothschild (Panamd; Tipton & M^ndez, chigger Trimenopon,^ Eutrombicula, chig- Tritopsylla, flea 1966, p. 326) Trombicula, Rhopalopsyllus cacicus saevus Jordan and ger chigger Tur, Acarina Rothschild (Panamd; Tipton & M6ndez, Gigantolaelaps, louse Xenopsylla, flea 1966, p. 326) Gliricola,^ louse Rhopalopsyllus lutzi lutzi Baker (Argentina; louse Acanthocephala Brasil: Minas Gyropus,^ Paraguay; Goias, Gerais; Echinorhynchus Haemolaelaps, mite Costa Lima & Hathaway, 1946, p. 139) Hamanniella Heterothrombidium Neotyplocercus rosenbergi Roth.schild (Ec- Intercutestrix Cestoda uador; Peni; Costa Lima & Hathaway, Ixodes, tick Oorchoristica 1946, p. 229) Leeuwenhoekia Tritopsylla intermedia intermedia Wagner Sparganum Microthrombidium (Brasil; Colombia; Ecuador; Paraguay; Nematoda Central America; Costa Lima & Hathaway, Neothyphloceras, flea Aspidodera 1946. 228) p. Capillaria Omithonyssus, mite Cortiamosoides Pentastoma (see Trematoda Cruzia Porocephalus) Globocephalus Polygenis, flea Duboisiella proloba Baer (Peru; Tantaledn et Gnathostoma Porocephalus, Szi- al., 1992) [Bursotrema tetracotyloides Pentastoma Gongylonemoides dat (Peni; Tantaledn et al., 1992)] Zonorchis allentoshi Foster (Peni; Tantaledn et al., 1992) '"All refer to one captive individual and are certainly Foster Plagiorchis didelphidis (Peni; Tanta- due to contamination," Wemeck, in Hopkins, 1949, lean et al., 1992) p. 439. BIOLOGY LIBRARY HERSHKOVITZ: PHILANDER 101 BURWLL HALL 31 nrT 1^^1997 12010 16393 P. opossum quica P. andersoni andersoni Balta, Peru cf (old) Balta, Peru 9 (adult)

14012 16398 P. opossum quica P. andersoni andersoni Peru 9 9 Balta, (yg) Huanhuachays , Peru (adult)

95313 41446 P. opossum opossum P. andersoni andersoni Makerie, Suriname Montalvo, Ecuador cf (adult)

13799 87123 P. opossum fuscogriseus P. andersoni andersoni Achotal, Mexico 9 (subad) Santa Luisa, Peru

13801 388405 P. opossum fuscogriseus P. andersoni andersoni Achotal, Mexico 9 (subad) Rio Cunucunuma, Venezuela 9 (subad)

Fig. 19. (fnnh) Conules of third upper molar labial shelf of Philander species from various geographic localities. See Table 6 for explanation of symbols.

Macielia Sarcocystis Species and Named and Unnamed Oxysoma Trypanosoma Subspecies of Philander opossum Philostrongylus Trematoda Physaloptera The taxonomic accounts are arranged in the fol- Brachylaemus Skrjabinofilaria lowing order. Maritrema Subulura Opisthorchis 1. Philander opposum Linnaeus Travassostrongylus Philander Linnaeus Trichuris Paragonimus opossum opossum Philander Temminck Viannaia Phaneropsolus opossum quica Plagiorchis Philander opossum frenatus Olfers Protozoa Platynosomum Philander opossum melanurus Thomas Besnoitia Podospathalium Philander opossum subspecies nov. 1 Haemogregaraina Zonorchis Philander opossum subspecies nov. ?

32 FIELDIANA: ZOOLOGY parts of Amazonian Peru, and in central Peni in the departments of Pasco, Junin, and Ayacucho. Philander opossum and P. andersoni are sympatric in the upper Rfo Punis and mid and lower Rio Ucayali basins. Historic Variation—The nearest putative ancestor of P. opossum was perhaps slightly smaller than its living descendants and possibly grayish agouti, likely marked by pale brown superciliary spots. The warm, humid Late Pleistocene climate of the ancestral habitat east of the Andes supported a dense tropical forest that permitted spread of the four-eyed opossum around the northernmost projections of the Andes to the Pa- cific coast and Middle America. With the onset of the cooler, drier climate of Late Pleistocene or Early Recent, shrinkage and fragmentation of the forest resulted in isolated refuges surrounded by superceding or invading savannas (Prance, 1982). The isolated populations of Philander differentiated into the racial forms recognized here. The most widely distributed or largest portion of the erstwhile continuously distributed ancestral population is P. opossum quica. An undescribed population {P. opossum subsp. nov. 1) of northeastern Venezuela (Fig. 5, is said to be very dark. The isolated P. opossum melanurus of coastal Ecuador is also dark. It remains to be seen if the ranges of P. opossum melanurus and P. opossum fuscogriseus meet on the west and between P. opossum quica and the undescribed P. opossum on the northeast. The geographic range of the southeastward- spreading P. andersoni overlaps that of the west- ward-advancing P. opossum quica. Fig. 20. Left third molar and lower third upper right Cranial and Dental Characters—Cranial molar of Philander opossum. See Table 6 for names of and dental characters as for the features. genus, comparisons included.

Coloration (Fig. 4)—Color pattern of head as for the genus; cheeks, chin, throat, neck, chest, Philander opossum fuscogriseus inner sides of limbs ochraceous to J. A. Allen belly, buffy or whitish; brown, scrotum un- Philander andersoni Osgood marsupial region or to Philander andersoni andersoni Osgood pigmented partially completely pigmented; lateral line, if present, usually confined to Philander andersoni mcilhennyi pelvic as a less as a shoul- Gardner and Patton region hip patch, frequently der stripe or patch, color buffy to ochraceous orange, often appearing olivaceous when pale hair ba.ses show crown, midline of and Philander opossum Linnaeus through; nape, dorsum to basal hairy portion of tail brownish; middorsal band, if present, poorly to strongly de- (Synonyms under subspecies heading) fined, individual cover hairs agouti (banded), tip Distribution (Fig. 5)—Distribution as for the brown (eumelanin), buffy (pheomelanin) subter- genus, except replaced by P. andersoni in Ama- minally, followed proximally by a brown band zonian Colombia and Ecuador and contiguous merging into curly and distinctly thinner grayish

HERSHKOVITZ: PHILANDER 33 Table 6. Explanation of symbols used for marsupial dentition (Figs. 1 7-20).

Upper teeth Cones' 1 Eocone (paracone) 2 Protocone 3 4 Metacone 5

Conules and styles-

a {A) Mesiostyle-a (parastyle) b (£) Distostyle-fe (metastyle; hypoconule) j (B) Ectostyle-y it Ectostyle-^ / (Q Ectostyle-/ (mesostyle) m (D) Ectostyle-m

Cristae^

Cristae extend from-to or between cusps; reference cusps in parentheses are not elements of the indicated crista. / Eocrista [l-a-j-k-l-m-b] II Centrocrista [1^] III Epicrista [7-2] IV Postmetacrista [4-b] V Plagiocrista (metaloph) [2-4] VI Protoloph (protocrista; crested portion of cingulum (G) Basins or fossae = Table 7. Molarization of conule C in m'. a barely = rudimentary; b rudimentary conule not always well = - = defined; c low distinct conule; d small cusp; e cusp comparable to conule B in bulk. The tabulation is based on about 95 specimens in the Field Museum, plus about an equal number of borrowed specimens.

Locality Table 9. Measurements of Philander opossum opossum Linnaeus.

Locality Table 9. Extended.

Condylobasal

Taxonomy—Sole basis for the Linnaean Di- delphis opossum is the animals described by the Dutch pharmacist and naturalist collector Albert Seba (1734, in the work entitled Locupletissimi

rerum naturalium thesauri . . . , 1, p. 56, PI. 36, Fig. 1 [male]. Fig. 2 [female]). Upper parts of the body of the male four-eyed opossum shown in the cited figure reproduced here (Fig. 21) are described as dark chestnut, and the tail base is said

to be furred for the length of a human finger, or roughly between 7 and 9 cm. Seba's female. Fig- ure 2 (Fig. 21 here), is fully pouched, but no formal description of the animal appears in the text. Linnaeus (1758, p. 55), in citing Seba, evidently intended that the name D. opossum apply to the female. His diagnosis and description, '' D[idelphis]. cauda semipilosa, superciliorum re- gione pallidore, mammis binis" and ''abdomen circa mammas contrahitur in marsupium; policis postici mutici," leaves no doubt. Color is not mentioned, but the "cauda semipilosa" can be variously translated as hairy for half its length or as partially furred. Whatever the wording, the interpretation must be derived from Seba's wood-

cut figure of the female (Fig. 2 1 ). Variation—The types of P. opossum opossum Linnaeus, as described by Seba (1734, pp. 56, 57) are brown-phase individuals. With respect to other Suriname specimens, a male I collected in La Poule and another in Clevia, Paramaribo, are brown phase and resemble the lectotype preserved Fig. 21. Male and female lectotypes of Didelphis in alcohol in the Leiden Museum. A young female opossum Linnaeus (1758). Upper, "'Philander opossum taken in is One of two sive Carigueja Brasiliensis mas," the male opossum. Lelydorpplan gray phase. Lower, ''Philander americanus, sive Carigueja, cum ca- individuals collected by H. A. Beatty in the Wil- Sacrum ventris intrantibus foemina." and tulus. Figures helmina Mountains is gray phase, the other brown quotations from Seba (1734, p. 57, PI. 36, Fig. 2). phase. Other Suriname skins examined are of juvenals. Gray- and brown-phase individuals and inter- P. andersoni. It may be that during a Pleistocene mediates occur throughout the range of the race. climatic change, the warm, humidity-loving opos- In all, however, underparts are more densely pig- sum disappeared from even more extensive areas mented than in those of neighboring P. opossum where savannas replaced forests. quica. The large series from the Rio Tocantins Characters—General body size larger (Baiao; Ilha do Taiuna) exhibit the full range of throughout than quica or melanurus; molars pro- color variation in P. opossum opossum. portionately smaller; upper parts of trunk grayish Remarks—Under the name Metacherius [sic] to buffy brown, under parts buffy orange to ochra- opossum, Sanderson (1949, p. 787) recorded 10 ceous orange; tail usually parti-colored. males and 1 2 females from Suriname. Of the total, Comparisons—Few individuals of P. opossum 16 are said to have been taken in Paramaribo, and opossum are as small as average-sized quica, and one, a large female, from Zanderij. Other specific the underparts of none of the specimens at hand localities are not mentioned. I found skins of six show the whitish, pale buff or grayish brown un- males and three females in the British Museum, derparts present in many individuals of other all labelled Paramaribo, collected by I. T Sander- races. son. There is also the female from Zanderij (bm Measurements—Tables 1 and 9. 52.122), but it proves to be Metachirus nudicau-

HERSHKOVITZ: PHILANDER 39 datus. Its mammae and a sketch of the mammary (usnm); El Dividiri, 30 km NW Valera, 1 (usnm); formula are by Sanderson (1949, PI. VII, Figs. 3, Motatin, 1 km NNE, 3 (ukmnh); Motata, 5 km 4). The remaining 12 of the 22 collected by San- NNE, 1 (ukmnh); Rio Motatin, 5 (ukmnh). derson are not among the skins in the British Mu- seum. Husson (1978, p. 28) mentions only the Zanderij Metachirus collected by Sanderson but Philander opossum quica Temminck no Philander. — Didelphis quica Temminck, 1824:36. Desmarest, Specimens Examined 370. BRASIL. Amapd: 1827:387—characters. Jentink, 1887:301—BRA- Ferrovia Amapa, km 192, 1 (lsumz); Macapa, Rio SIL: Temminck collection. Jentink, 1888:220— BRASIL; larvata ms. name Amapari, 3 (mzusp); Mazagao, Rio Maraca, 1 part, cotype; Didelphis in synonymy. Hochstetter, 1946:11, PI. 1, Fig. 2 (Mzusp); Serra do Navio, 59 (usnm, 57; mpeg, 2); (brain, sagittal section in skull)—brain, dura mater. Rio 1 Terezinha, Amapari, (lsumz); Amazonas: Avila-Pires and Gouvea, 1977:9—BRASIL: Rio de Auara Igarape, 2 (amnh); Ipixuna, Rio Funis, 1 Janeiro (Monte Serrat; Benfica; Maceiras); local (usnm); Lago do Baptista, 1 (amnh); Lago Sam- name, guaquica. Didelphys quica, Waterhouse, 1841:90—part, BRA- paio, Rosarhino, 2 (amnh); Rio Madeira, 1 SIL. Waterhouse, 1846:480—part, BRASIL. Bur- (amnh); "River 5 (bmnh); Santo Antonio Negro," meister, 1854:136—BRASIL: Rio de Janeiro de 1 Santa Villa Bella Im- Uayara, (amnh); Clara, (Novo Friburgo). Pelzeln, 1883:1 10—BRASIL: peratriz, 3 (amnh); Tefe, 1 (amnh); Pard: Alta- Rio de Janeiro (Sepetiba); Sao Paulo (Ypanema); Mato Grosso mira, 54 km S, 150 km W, 3 (usnm); Altamira, (Mato Grosso). Oudemans, 1892:14, Fig. 25 (Cowper's gland)—male accessory repro- Rio Xingii, 3 (usnm); Ananindeua, 2 (mpeg); Aru- ductive organs. matheua, 1 (mpeg); Baiao, Rio Tocantins, 14 D[idelphys]. quica, Wagner, 1855:225, PI. 18 (fe- 17 (amnh); Belem 2 (amnh, 1; usnm, 1); Cameta, male)—description, figure, and notes from Natterer (mzusp); Canudos, 1 (fmnh); Capim, 6 (amnh, 5; manuscript. Metachirus quica, Burmeister, 1856:70, PI. 7 (male), MPEG, 1); Cuatipuru, Flor do Prado, 2 (mpeg); p. 1 1, Fig. 2 (skull)—part, BRASIL: Rio de Janeiro Fordlandia, 1 (amnh); Gradaus, Rio Fresco, 2 (Novo Friburgo); characters. Hensel, 1872:120— (mzusp); Igarape Tapereba, 4 (mpeg, 2; mzusp, 2); BRASIL: Rio Grande do Sul; characters. Goeldi, Ilha do Taiuna, 85 (amnh); Ipeau-Apez, Belem, 2 1894:460—BRASIL: Rio de Janeiro (Serra dos 6r- J. A. Allen, 1900:195—BRASIL: lo- (usnm); Iriteria, 1 (mpeg); Lazaropolis, 4 (mpeg); gaos). (type cality, "coast region of Brasil, just south of Rio de Macajuba, Rio Tocantins, 3 (amnh); Maraba, Rio Janeiro"). Tocantins, 2 (usnm); Marcos, 1 (mpeg); Mazagao, Metachirus opossum quica, J. A. Allen, 1916c:562— Rio Tocantins, 1 (mpeg); Murutucii, 5 (bm, 2; BRASIL: Mato Grosso (Urucum); Rio de Janeiro FMNH, 3); Peixe-Boi, 1 (mpeg); Rodovia Belem- (Sapitiba [= Sepetiba], restricted type locality). Miller, 1916:589—BRASIL; habits. Carvalho, Brasflia, 3 (mpeg); Santa Maria, Bragan9a, 1 1957:3—BRASIL: Acre (Seringal Oriente, Rio Ju- (usnm); Santarem, 10 (mzusp, 2; usnm, 8); San rua). do 5 Miguel Guama, (mpeg, 2; mzusp, 3); Utinga, [Metachirops] quica, Matschie, 1916:268—BRASIL: 22 (MPEG, 6; usnm, 13; amnh, 1; mzusp, 2); Ro- Rio de Janeiro (type locality, "Sapitibi"). 1927:243—BRASIL: Rio raima: Caracarai, 2 (mzusp). GUYANA. Essequi- Metachirops quica, Pohle, de Janeiro (Barreira; Teresopolis). Schirch, 1932: bo Islands-West Demerara: Buckhall, 2 (bm[nh]); 85—BRASIL: Rio de Janeiro (Teresopolis, 960 m); Upper Kanuku Mts., 5 Takutu-Upper Essequibo: parasite (Amblyopinus). (bm[nh]); Rupununi River, 1 (bm[nh]); Demer- Metachirops opossum quica, Miranda Ribeiro, 1936: do Sul ara-Mahaica: Better Hope, 1 (bm[nh]); George- 340—BRASIL: Rio Grande (Porto Feliz, Rio Rio de Janeiro Santa town, 2 (fmnh); Hyde Park, 4 (bm[nh], 2; fmnh, UrugaO; (Teresopolis); Catarina. Vieira, 1945:421—BRASIL: Mato Gros- 2); Pomeroon-Supenaam: River, 1 Supenaam so (Palmeiras). Vieira, 1949:345—BRASIL: Sao (BM[NH]). GUYANE FRAN^AISE. Guyana: Cay- Paulo; Minas Gerais; Espirito Santo; Mato Grosso; enne, 5 (bm[nh], 1; FMNH, 4). SURINAME. Suri- local names, quica, quaiquica. Ruschi, 1965:2— BRASIL: Santo local cui- name: Paramaribo, Clevia, 2 (fmnh); Lelydorp- Espirito (coast); names, ca, chupata. plan, 1 (fmnh); Paramaribo, 10 (bm[nh], 9; lec- Philander opossum quica, Perondini and Perondini, rmnh); no 3 totype, precise locality, (bm[nh]); 1966:28—BRASIL: Sao Paulo (Cantarera Forestal Nickerie: Kaiserberg Airstrip, 1 (fmnh); Makerie, Preserve); sex chromatin in somatic cells. Carval- Paulo West River, 2 (fmnh); Saramacca: La Poule, 1 ho, 1965:251—BRASIL: Sao (Borageia). [Holothylax] quica, Cabrera, 1919:48—classification. (fmnh); Loksie Hattie, 1 (fmnh). VENEZUELA. Giebel, 1859:227— "der Bolivar: Ciudad 146 km 7 km 2 D[idelphys]. quica [sic], Bolfvar, S, NE, Guica." (usnm); 1 Rfo 10 Maripa, (amnh); Yuruan, Didelphis myosuros Temminck, 1824:38—BRASIL: (amnh); Trujillo: Agua Viva, 18 km N Valera, 1 (type locality); cotypes in the Leiden, Vienna,

40 FIELDIANA: ZOOLOGY Frankfort, and Prince Maximilian zu Wied-Neu- Rio Grande do Sul; local names, guaquica, guaqui. wied Museums. Hershkovitz, 1959:343—in syn- Ihering, 1894:10—BRASIL: Sao Paulo. Thomas, onymy of Philander opossum frenata [sic] Olfers; 1902a:64—BRASIL: Parana (Ro^a Nova, Serra do original description based on four-eyed pouched Mar). J. A. Allen, 1 91 6b:201—COLOMBIA: Meta opossum. (Villavicencio). Bertoni, 1923:51—PARAGUAY: Metachirus nudicaudatus mvosurus [sic], Miranda Ri- Central (Bahia de Asunci6n); Guaranf name, guaki. beiro, 1936:345—BRASIL: Bahia; characters; Por- M[etachirus]. opossum, Miranda Ribeiro, 1905:189—

tuguese translation of original description; ". . . as BRASIL: Rio de Janeiro (Monte Serrat. Itatiaya).

femeas tern sacco . . .". — um completo — Metachirops opossum, Bertoni. 1939:6 PARA- Didelphis lanata Jentink, 1888:220 Natterer ms. GUAY. Cre.spo. 1950:6. Fig. 1 (animal)—ARGEN- name on label of a syntype of quica Temminck. TINA: Misiones (Rio Urugua-i. 39 km from Puerto Metachirus canus Osgood. 1913:96—PERU: San Bemberg); habits; locomotion; local name guaitica. Martin (Moyobamba, type locality); holotype, Philander opossum, Davis. 1945a: 122—BRASIL: Rio male, skin and skull. Field Museum of Natural His- de Janeiro (Fazenda Boa Fe. Teres6polis). Davis, tory, no. 19347; collected 4 August 1912 by W. H. 1947:1—BRASIL: Rio de Janeiro (Fazenda Boa Osgood and M. P. Anderson. Osgood. 1914:148— Fe, Teres6polis); life history. Moojen and Avila- PERU: San Martin (Moyobamba). Sanborn, 1947: Pires, 1966:397—BRASIL: cerrado formation (ab- 215—type history. sent; Didelphis and Monodelphis only). Gardner [Metachirops] canus, Matschie, 1916:268—classifi- and Patton. 1972:5—PERU: Ucayali (Balta, Yari- cation. nacocha); comparisons with mcilhennxi. Reig et al.. [Holothylax grisescens] canus, Cabrera, 1919:47— 1977:197. 212. PI. 1 (karyotype)—part, PERU: classification. Ucavali (Yarinacocha; Balta). Mello and Moojen, Metachirus opossum canus, Thomas. 1927a: 372— 1979:289—BRASIL: Distrito Federal (Brasilia PERU: San Martin (Rioja; Moyobamba; Yurac gallery forest; Fazenda Agua Limpa. gallery; Yacu). Thomas, I927b:606—PERU: Hudnuco Guara, gallery); Mato Grosso (Baliza, gallery; Bar- (Tingo Maria, 2000 ft). Thomas, 1928a:264— ra do Gargas gallery; Pocon^ gallery; Salobra gal- PERU: Ucayali (Cumeria, 1000 ft; Chicosa, 1000 lery); Golds (Aragar^as gallery; Formosa gallery). ft; San Jer6nimo, 500 ft). Correa Gomes, 1984:369—BRASIL: Mato Grosso M[etachirus]. o[possum]. canus, Thomas, 1928b: (Salobra); helminth infection. Da Fonseca and Red- 294—crucialis Thomas a synonym. ford. 1984:517—BRASIL: Distrito Federal Philander opossum canus, Sanborn. 1949:277— (Parque Nacional, Brasilia; humid gallery forest PERU: Vcayali (Yarinacocha). Sanborn. 1951:2— and valley side wet campo; cerrado [uncommon]). PERU: Cuzco (Hacienda Cadena). Cabrera, 1958: Mares et al., 1989: 19—BRASIL: Mato Grosso (Po- 35—crucialis Thomas a synonym. con6). Metachirus opossum azaricus Thomas, 1923:604— Philander opossum opossum, da Fonseca and Kierulff PARAGUAY: Paraguari (Sapucay, type locality): (not Linnaeus), 1989: 118, 143—BRASIL: Minas holotype. female, skin and skull. British Museum Gerais (Fazenda Esmeralda; Fazenda Montes Clar- (Natural History), no. 1903.2.3.36; collected 8 Au- is); reproduction; habits. gust 1892 by W. Foster. Didelphis nudicaudatus, Desmarest (part not £. Geof- Met[achirops]. opossum azaricus, Krumbiegel. 1941: froy), 1827:390—mxosuros Temminck a synonym. 203. 206—PARAGUAY: (Lapango); BRASIL: Rio Waterhouse, 1841:94, PI. 2 (animal). Jentink, 1888: Grande do Sul (Passo Fundo); BOLIVIA: Chiqui- 220—part, syntypes of myosuros Temminck and tos (San Ram6n); crucialis Thomas a synonym. opossum of Temminck collection. Thomas, 1888: Philander opossum azaricus, Cabrera. 1958:34—-clas- 332—part, myosuros in synonymy. sification. Kantis, 1963:54—ARGENTINA: Chaco Philander opossum frenata [sic], Hershkovitz (part (Rio de Oro. mouth); female with three young. not Olfers), 1959:342—Didelphis quica Temminck Crespo. 1974:3—ARGENTINA: Chaco (Rio de in synonymy. Oro. mouth); Misiones (Fracrdn, San Pedro). Metachirops opossum quichua [sic], Krumbiegel. Metachirus opossum crucialis Thomas. 1923:604— 1941:200. 201. 206—name a reiterated lapsus for BOLIVIA: Santa Cruz (type locality. Santa Cruz quica Temminck; BRASIL: Sao Paulo (Juquia, de la Sierra); holotype, female, skin (originally Serra do Mar). mounted) and skull, British Museum (Natural His- tory), no. 1847.11.22.15; collected by Thomas Types—An number of some Bridges. unspecified skins, Didelphxs opossum, Waterhouse, 1841:90—part, mounted, skulls and skeletons of both sexes in the BRASIL. Waterhouse, 1 846:485—part, BRASIL. Vienna, Leiden, Paris, and Prince Maximilian zu Thomas, 1888:329—ARGENTINA: Chaco; BRA- Wied-Neuwied museums; female syntype in the SIL: Rio Grande do Sul (Taquara); BOLIVIA: Vienna Natural History Museum, collected 3 Santa Cruz (Santa Cruz de la Sierra). Winge, 1893: March 1818 Johann lec- 7. 38. PI. 1. Fig. 7 (ear, foot). PI. 3. Fig. 1 (skull). by Natterer, designated PI. 7 4, Fig. (humerus)—BRASIL: Minas Gerais totype by Hershkovitz (1959, p. 337). Recent and com- (Lagoa Santa, fossil); characters; Type Locality— "Bresil," restricted to "Sap- parisons. itiba" (= Sepetiba), Rio de Janeiro (Rio de Ja- D[idelphvs]. opossum, Tschudi. 1844:14, 144, 151 — PERU. neiro, 22°58'S, 43°42'W), by J. A. Allen (1900,

Metachirus opossum, Ihering, 1892:99—BRASIL: p. 195; 1916c, p. 562).

HERSHKOVITZ: PHILANDER 41 Table 10. Measurements of Philander opossum quica. Means (extremes in parentheses) and number of specimens.

Locality Table 10. Extended.

Condylobasal Table 10. Continued.

Locality Table 10. Extended (continued).

Condylobasal

other hand, the nearly equal body and tail pro- Museum, is unquestionably that of the pouchless portions of the same individuals and unpigmented species. Evidently females were not seen, and Wa- tail base indicate the pouched species. The de- terhouse mistakenly included Didelphis nudicau- scriptions of external characters lack precision, data along with D. philander (= Caluromys phi- but some details of the original text apply equally lander) in his Didelphis "Section 1. Opossums in well to both pouched and nonpouched species but which the pouch is well developed." in no case to the latter alone. In his account of the mammals of Rio de Ja- nature of the Despite the apparent composite neiro and Minas Gerais, Burmeister (1854, p. 135) description, Temminck (1824, p. 39) denied that accepted Wagner's (1843) use of the emended his myosure might be identical with D. nudicau- form of the name Didelphis myosurus Temminck data E. Geoffroy, because, he notes, "// est dit and treatment of D. nudicaudata as a synonym. la de cet animal est toute dans le texte que queue Burmeister saw no specimens of myosuros in nue, et que la femelle n'a point de poche.'" Tem- southeastern Brasil, and he drew on Wied-Neu- minck drew attention, nevertheless, (1824, p. 40) wied (1826) for a description of the species. to a skin of a female in poorly preserved young Shortly thereafter, Burmeister (1856, p. 68, PI. 10) the Paris museum labeled D. nudicaudata he that, described a male and female labeled ''Didelphis in with D. said, agreed every respect myosuros. myosurus Temminck" in the Berlin Museum. His that this individual He questioned, however, may characterization of the two animals, believed to have served E. for the Geoffroy original descrip- have originated in Para, is somewhat equivocal tion of D. nudicaudata. authors Notwithstanding, but fits that of Didelphis nudicaudata E. Geoffroy, with Wied-Neuwied (1826) and Des- beginning cited as a synonym. The adult male illustrated in marest concluded that Temminck (1827, p. 390) color is identifiable as Metachirus nudicaudatus. did indeed his D. myosuros with D. nu- identify Thomas (1888) cited "Didelphis myosurus, dicaudata. The mistaken identity gave rise to con- Temm." and its usage by other authors, including fusion in later years. Wied-Neuwied (1826), Wagner (1843), and Bur- Wied-Neuwied (1826, p. 400, PI. 2, Fig. 5 meister (1854; 1856), in the synonymy of "Di- [skull]) described one of two male four-eyed delphys nudicaudata E. Geoffroy." opossums captured in "Comechatiba" (= Com- Jentink (1888, pp. 220, 221) listed the follow- oxatiba or Cumuraxatiba, 17°06' S, 39°ir W), ing skins identified as Didelphis nudicaudata E. between the Rios Prado and Corumbao, Bahia, Geoffroy in his catalog of mammals of the Mu- Brasil, and referred them to D. myosuros Tem- seum d'Histoire Naturelle des Pays-Bas, where minck. The finely detailed characterizations and many of Temminck 's specimens are deposited: the figured skull of the animal are unquestionably those of Metachirus nudicaudatus . Wied-Neu- a. Individu adulte monte, un des types du Didel- wied, however, could not be certain of the pro- phis myosuros Temminck. Bresil. the 1 priety of name myosuros because he ( 826, p. b. Male adulte monte, un des types du Didelphis 405) "had not seen a female which, according to myosuros Temminck. Bresil, 1822. Temminck, has a pouch" (my translation of the c. Male adulte monte. Bresil. original German). d. Male semi-adulte monte. Bresil. Du Cabinet de In his to Schreber's supplement Sdugethiere, M. Temminck. Didelphis opossum. Wagner (1843, p. 43) included Didelphis myosu- It is that most, if of the above ros Temminck, emended to ''Didelphis myosu- unlikely any, male and unsexed of D. actu- rus,'' in his fully pouched opossum group and "types" myosuros served as bases for the of short-haired subgroup (Wagner, 1843, p. 37— ally original description the The females of the indi- ''mastotheca ventrali ampla''; p. 42— ''vellere species. "plusieurs vidus des deux sexes'' mentioned Temminck brevC). His description of D. myosurus, however, by Individual listed is based on the Berlin Museum's type specimen (1825, p. 40) are missing. a, by subse- of Didelphis frenata Lichtenstein (Olfers): Didel- Jentink as a "type," may be a specimen phis nudicaudata E. Geoffroy was listed as a syn- quently labeled "myosuros'' by Temminck. Spec- onym of Didelphis myosurus without explanation. imen b, dated 1822, is certainly one of those col- in do Sai Waterhouse (1846, p. 482) treated Didelphis lected by Natterer the same year Registo myosuros Temminck, 1824, as a junior synonym (April) or Ypanema (January, June) and sent to of Didelphis nudicaudata E. Geoffroy, 1803. His the Vienna Museum (Pelzeln, 1883, p. 111). description, based on a specimen in the British These Natterer specimens represent the opossum

48 FIELDIANA: ZOOLOGY sometimes incorrectly known as Af. nudicaudatus Variation—Material from Rio de Janeiro in- personatus Miranda Ribeiro. cludes individuals of P. opossum quica with sat- Misapplication of the name Didelphis myosuros urate coats like those of P. opossum opossum Temminck for pouchless four-eyed opossums was from Suriname and others with the typical gray pointed out in 1959 (Hershkovitz, 1959. p. 343), coat. A large series (27) from An^polis, Goias, and treatment of the name as a junior synonym of appears to be intermediate between P. opossum Philander opossum frenata [sic] Olfers was sug- opossum and the gray, eastern Brasilian P. opos- gested. On the other hand, my designation of the sum quica. Seven adults in old pelage, taken Oc- Comoxatiba opossum recorded by Wied-Neuwied tober through February, appear brown because of as lectotype of myosuros Temminck is invalid. exposure of the basal portion of the dorsal hairs. The specimen, a male Metachirus nudicaudatus, Eight adults in fresh winter pelage (April-August) was not mentioned by Temminck in his descrip- appear grayish. tion of myosuros and likely was not seen by him In the more southern states of eastern Brasil, at any time. and in Paraguay, Bolivia, and Peru, the animals It is now proposed to designate as lectotype of are predominantly grayish or grayish brown dor- myosuros one of the pouched females used by sally, the underparts pale. Temminck in the original description of the taxon. Dichromatism in nine specimens of quica from In the absence of evidence to the contrary, the Balta, Ucayali, Peru, parallels that of P. andersoni type locality, said to be Brasil, is here restricted mcilhennyi from the same locality. The color dif- to southeastern Brasil. Thus, myosuros {- myo- ference, however, is not sexual. In six specimens a of P. are surus) Temminck becomes synonym opos- (two 9 $ , four S 6), underparts pheomelanic, sum quica Temminck. the hairs buffy or orange to the roots, as in female Metachirus canus Osgood (1913), from Moy- P. andersoni. In three specimens (9, 6 6), un- obamba, Peni, based on a single specimen, was derparts are dominantly eumelanic or grayish, the distinguished from western Colombian grisescens hairs pheomelanic terminally, eumelanic basally, {= P. opossum melanurus) by its generally paler and showing through at the surface, as in male P. coloration and parti-colored tail. Comparisons andersoni. The tail of the single female of the were not made with P. opossum quica, from eumelanic series is entirely pigmented. The ter- which it is indistinguishable, presumably for lack minal whitish portion in all other members of the of material at hand from Bolivia, Paraguay, and Balta series varies from about one-third to one- southeastern Brasil. eighth of total length without significant differ- The Paraguayan Philander opossum azaricus ence otherwise between those of the two color Thomas (1923, pp. 604) was characterized as a phases. Dorsum and sides are likewise nearly the "uniform grey subspecies without blackened same in both, but crown and muzzle are more crown." Thomas' (1923, p. 604) description of deeply and extensively brown in the eumelanic or Philander opossum crucialis from Santa Cruz de gray-phase group. la Sierra, Bolivia, is virtually a paraphrase of that Convergent dichromatism between the Balta P. of azaricus. The type, a "remade skin," "some- opossum and P. andersoni throws more doubt on what faded," was collected about a half-century the validity of the concept of "character displace- earlier by Thomas Bridges. Later, with "good se- ment" in mammals. ries" of the Peruvian canus and Bolivian crucialis An extremely large male and a female from before him, Thomas (1928b, p. 294) opined that Yarinacocha, Rio Ucayali, Peru, agree with P. an- the "two forms should not be considered as dis- dersoni mcilhennyi with respect to body size and tinct from each other" Krumbiegel (1941, p. 204) dark underparts and limbs but lack the diagnostic regarded azaricus and crucialis indistinguishable. dark dorsal band. They are readily distinguished Except for the color phases, seasonal differ- by dorsal coloration and pelage pattern from the ences in pelage, and certain exceptionally large intermediate series of quica taken at nearby Pu- individuals, I find a remarkably high degree of callpa as well as by absence of sexual dichroma- uniformity in the morphology of the common Phi- tism. They also differ from Pucallpa P. opossum lander opossums of eastern Brasil, northern Ar- quica by larger size and darker color throughout gentina, Paraguay, lowland Bolivia, ea.stem Peru, but agree in dental and cranial characters. The and Ecuador Material from Colombia and west- pigmentation of underparts and limbs is interpret- ern Venezuela is similar, but the disjunct distri- ed as the dark-phase coloration of occasional oc- bution suggests convergence. currence in P. opossum quica. The extremely

HERSHKOVITZ: PHILANDER 49 large size of each of the Yarinacocha samples may mor6, 12°26'S, 3 (amnh; fmnh, 2); San Ignacio de be attributed to extreme old age and possibly un- Moxos, 1 (usnm); San Joaquin, 13 (fmnh); San usually favorable living conditions comparable to Pablo, 1 (fmnh); San Ramon, 1 (fmnh); Santa those of the captive quica recorded from eastern Rosa, 1 (fmnh); Santo Dios, 1 (fmnh); Vaca Diez,

Brasil (see above, p. II). Riberalto, 1 (usnm); Cochabamba: Rio Ichilo, 52 The opossums from the Rio Orinoco basin in km S, mouth, Rio Chapare, 1 (amnh); La Paz: Colombia and Venezuela are practically indistin- Chulumani, 1 (bm[nh]); Santa Cruz: Ascension de guishable from Peruvian P. opossum quica or Guarayos, 2 (fmnh); Buena Vista, 12 (amnh; from the Field Museum material from Bolivia and bm[nh], 8; fmnh, 3); El Palmar, 2 (usnm); Ha- Brasil. They may represent a case of parallelism macas, 2 (amnh); Ibanez, Rio Piray, 2 (usnm); Rio or may be a disjunct remnant of an erstwhile Chapare, 2 km S, mouth, 1 (amnh); Santa Cruz, widely distributed subspecies P. opossum quica 3 (bm[nh], holotype of crucialis Thomas; usnm, which may not have differentiated much if any- 2); Santa Rosita, 4 (usnm); Tocomechi, Warnes, 1 thing from the hypothetical ancestral form of the (usnm); Warnes, 3 (usnm). BRASIL. Acre: Ser- species. ingal Oriente, 4 (mpeg, 3; mzusp); Espirito Santo: While this monograph was in preparation, Pat- Cachoeira Bonita, Parque Nacional do Caparao, 1 ton et al. (1995 [1996]) had the opportunity to (mnr); Campinha, 4 (mnrj); Engenheiro Reeve, 3 examine tissues for molecular studies of one spec- (bm[nh]); Pedra Roxa, Parque Nacional do Ca- imen of P. opossum frenatus from "Brasil; Rio parao, 1 (mnrj); Santa Teresa, 7 (mnrj, 5; mzusp, de Janeiro; Naje, Carafao, Est. Teresopolis." Fif- 2); Sierra, 3 (mnrj); Vale Verde, Parque Nacional teen specimens from Teresopolis in the bm[nh], do Caparao, 8 (mnrj); Vila Velha, Morro de An- MNRJ, and USNM I examined were identified on the goles, 3 (mnrj); Vitoria, 2 (mnrj); Golds: Ana- basis of skins and skulls as P. opossum quica. polis, 27 (amnh); Aragar9as, 2 (mpeg); Trinidade, Present knowledge of P. opossum frenatus does 1 (mpeg); Mato Grosso: Caceres, 1 (usnm); Monte not permit its synonymy with any other known Alegre, 5 (mzusp); Santa Teresa, 1 (usnm); Uruc- four-eyed opossum. um, 5 (amnh); Minas Gerais: Alem Paraiba, 4 According to Patton et al. (1995 [1996], pp. 21, (mnrj); Benfica, Serra de Itatiaia, 1 (amnh); Boa 22), P. opossum frenatus "is quite divergent from Esperanga, Serra de Caparao, 3 (amnh); Concei- P. mcilhennyi and P. o. canus {— P. o. quica] (two 9ao de Mato Dentro, Boca de Ulaba, 1 (mnrj); taxa that are sympatric over much of western Fazenda Cardosa, Serra de Caparao, 2 (amnh); Amazonia in Brasil and Peru, with an average di- Fazenda de Floresta, Rio Matipo, 2 (mzusp); Fa- vergence of 16.45%). Consequently, /rena/M5 and zenda S. Francisco de Caparao, 1 (bm[nh]); Juiz {opossum canus + mcilhennyi) form a basal tri- de Fora, 2 (mnrj); Quartel de Sacramento, 1 chotomy with the three species of Didelphis in all (mzusp); "S. Francisco de C," 1 (bm[nh]); Pa- analyses." rand: Rio Paracai, 1 (bm[nh]); Roga Nova, Serra There appears to be a contradiction between the do Mar, 3 (bm[nh]); Rio Grande do Sul: no pre- 15 Teresopolis specimens I identified as P. opos- cise locality, 1 (bm[nh]); Rio de Janeiro: Barro sum quica on geographic and morphological Branco, 2 (mnrj); Itatiaia, Maceira, 2 (mnrj); Ita- grounds and the Teresopolis tissue Patton finds tiaia, Parque Nacional, 1 (mnrj); Mangaratiba, 5 unusually different from the others and the 426 (mnrj); Pedra Branca, Parati, 4 (mnrj); Rio de Ja- specimens I examined over the range of P. opos- neiro, 2 (amnh); Rodico, Serra do Mar, 1 sum quica (Fig. 5). Perhaps the Patton et al. speci- (bm[nh]); Sao Joao Marcos, 2 (mnrj); Teresopolis, men is not quica, but it has not been shown to be 15 (bm[nh]; mnrj, 13; usnm); Rondonla: Porto frenatus. Velho, 1 (usnm); Santa Catarlna: Hansa, 4

Specimens Examined—415. ARGENTINA. (bm[nh]); Joinville, 1 (fmnh); Sao Paulo: Alto da Chaco: Rio Chico, 1 (bm[nh]). BOLIVIA. El Serra, 1 (mzusp); Borageia, 26 (mzusp, 25; usnm, Beni: Arruda, 1 (fmnh); Barranquita, 1 (fmnh); 1); Butanta, Serra da Cantareira, 3 (mzusp); Casa Camiaco, 1 (amnh); Camino Vilches, 1 (usnm); Grande, 3 (mzusp); Cobia, 3 (mzusp); Costao dos

Centinela, 1 (fmnh); El Carmen, 1 (fmnh); Exal- Engenhos, 1 (mzusp); Iporanga, 5 (mnrj); Lageado i tacion, 2 (amnh; fmnh); Guayaramarin, 4 (amnh); de Iporanga, 1 (mzusp); Lageado do Aranhadava, Itonama, 1 (usnm); Magdalena, 5 (fmnh); Mer- 1 (mzusp); Monte Alegre, Amparo, 3 (mzusp); Pi- cedes, 1 (amnh); Palacios, Rio Mamore, 4 km SE, 2 (mzusp); Sao Sebastiao, 4 (bm[nh]); Serra quete, , 1 (amnh); Puerto Caballo, 2 (amnh); Puerto Siles, de Itatiaia, 1 (mzusp); Vila Oliveira, Mogi das 2 (amnh); Rio Ibare, mouth, 4 (amnh); Rio Ma- Cruzes, 1 (mzusp). PERU. Cuzco: Hacienda Cad-

50 FIELDIANA: ZOOLOGY ena, 2 (fmnh); Quincemil, 3 (fmnh); Hudnuco: ''D[idelphys]. Opossum L[innaeus] Tingo Maria, 2 (bm[nh]); Ucayali: Apayacu, 1 "Sarigoy, Sarigue War. (amnh); Balta, Rio Curanja, 9 (lsumz); Chicosa, ''Carigueia Marc. 222. Piso 323 (excl. ic.) 5 (bm[nh]); Cumaria, 1 (bm[nh]); Iquitos, 1 "D. frenata und supercilia 111. gehoren (amnh); Itaya, Iquitos, 1 (amnh); Lagarto Cocha, als varietaten hieher. Das Mannchen 2 (amnh); Pucallpa, 3 (usnm); Rio Urubamba, hat Gelbe Flecken, das Weibchen mouth, 5 (amnh); San Jer6nimo, 1 (bm[nh]); Santa grossere Weisse iiber den Augen." Rosa, 1 (amnh); Sarayacu, 8 (amnh); Yarinacocha, Remarks—A pouched four-eyed opossum per- 4 (fmnh; lsumz, 3); Madre de Dios: Boca Colo- haps representative of P. opossum quica, may rado, 1 (fmnh); Rio Tambo, 1 (usnm); San Martin: well have ranged into the erstwhile coastal forests Moyobamba, 3 (bm[nh], 2; fmnh); Rioja, 1 of southern Bahia and possibly farther north. (BM[NH]); Yurac Yacu, 10 (bm[nh]). PARAGUAY. Pending examination of the type or specimens Sapucay, 21 (bm[nh], 8, including holotype of from Bahia, if any, the name frenatus is retained. azaricus; usnm, 13); Tacuati, 1 (usnm). The specimens identified as P. opossum frenatus by Patton et al. (1995 [1996], pp. 7, 21, 22) are from Teres6polis in the state of Rio de Janeiro Philander opossum frenatus Olfers and well within the range of P. opossum quica. It 1815 (1804-181 1): 107— D[idelphys\. frenata Illiger. has not been shown that the Teres6polis specimen nomen nudum. is not P. opossum quica nor that it is representa- D[idelphys]. frenata Olfers. 1818:204. tive of P. [Metachirops] frenata, Matschie, 1916:268—BRA- opossum frenatus. SIL: Bahia (type locality). Met[achirops]. opossum frenatus, Krumbiegel, 1941: 206 omitted from taxonomic (in text)—explicitly Philander opossum melanurus Thomas review. Thomas, 1888:329— EC- Philander opossum frenata [sic], Hershkovitz, 1959: [l]Didelphis opossum, part, — UADOR: Guayas (Guayaquil). 338, 342 taxonomy; part, synonyms (superciliar- — is Olfers, mvosuros Temminck). Metachirus opossum melanurus Thomas, 1899:285 1913:36—ECUADOR: D[ideiphys]. superciliaris Illiger, 1815(1804-1811): type description. Lonnberg, 107—nomen nudum. Olfers, 1818:204—no locali- Pichincha (Gualea). melanurus, Matschie, 1916:268—clas- ty- [Metachirops] sification. D[idelphis]. mvosurus [sic], Wagner, 1843, footnote melanurus, Cabrera, 1919:48— 25, p. 44—BRASIL: Bahia, type of ''frenata Lich- [Holothylax opossum] tenstein" from Bahia; collected by Kaehne. classification. melanurus, 1921: Metachirus quica, Burmeister (part, not Temminck), Metachirops opossum Lonnberg, ECUADOR: Pichincha 1856:70. PI. 8 (female)—BRASIL: Bahia; type of 68— (Gualea). Philander melanurus, Cabrera, 1958:35— Didelphys frenata Illiger [Olfers] in Berlin Muse- opossum not melantho = in um, figured. part, ( fuscogriseus) synonymy. Metachirus nudicaudatus nudicaudatus, Cabrera P[hilander]. o[possum]. melanurus. Baker. 1974: 135—ECUADOR: Esmeratdas (part, not t. Geoffroy. 1803), 1958:48— "/renam (Quinind^). Lichtenstein (Wagner)" in synonymy. Type—Old male, skin and skull, British Mu- Type—Adult female, skin mounted, Berlin Mu- seum (Natural History), no. 1897.11.7.61; col- seum: collected before 1815 by Herr Kaehne. lected 1 1 April 1897 by W. F H. Rosenberg. Type Locality—Originally given as Brasil; ac- Type Locality—Paramba, Rio Mira, Imbabu- cording to Wagner (1843, footnote 25, p. 44) the ra, western Ecuador; altitude, 1 100 m. specimen labeled D. frenata in the Berlin Mu.se- Distribution (Fig. 5)—The humid forests of um was collected by Herr Kaehne in Bahia, pre- northwestern Ecuador in the provinces of Esmer- sumably in the present Sao Salvador, which may aldas, eastern Manabi, and bordering parts of be taken as the restricted type locality (cf. Mat- western Imbabura and Pichincha. Nothing is schie, 1916, p. 268). known of the opossums in the Colombian depart- Distribution (Fig. 5)—Known only from the ments of Narifio and Cauca, where fuscogriseus ascribed type locality. and melanurus may intergrade or prove to be

Characters—Olfers' (1818, p. 204) descrip- sympatric. tion oi frenata (and superciliaris) reproduced be- Characters—Dorsum dominantly brownish low is little more than the indication required to agouti, underparts buffy to deep orange often with validate the specimen label names first published dark ba.sal portions of hair showing through; tail as nomina nuda by Illiger. brown but extreme tip sometimes whitish.

HERSHKOVITZ: PHILANDER 51 Table 11. Measurements of Philander opossum melanurus. Means (extremes in parentheses) and number of specimens.

Locality Table 11. Extended.

Condyloba.sal Length Table 12. Measurements of Philander opossum fiiscogriseus. Means (extremes in parentheses) and number of specimens.

Locality Table 12. Extended.

Condylobasal Table 12. Continued.

Locality Head and Body Tail Hind Foot Ear

Guatemala 9 9 Los Amatos Mexico 6 6 Palenque 280 316 Achotal 269(252-292)4 289(258-310)4 44(40-47)4 36 (34-37)4 Mexico 9 9 Tekom Achotal Table 12. Continued.

Locality Table 12. Continued.

Locality muelles, 3 (usnm); Los Santos: Cerro Hoya, 5 nyi than in P. andersoni andersoni, points to (usnm); Panama: Cerro Azul, 21 (usnm); Cerro aquatic habits. The notably shorter, more woolly Jefe, 4 (usnm); La Zumbadora, Gorgas Labora- coat of P. opossum is less adapted for an aquatic tory, 3 (USNM); Veraguas: Isia Cebaco, 14 (usnm); environment. Isla Gobernado, 2 (usnm). COLOMBIA. Antio- The long-haired basal portion of the tail, ac- quia: Alto Bonito, 3 (amnh); Bellavista, 4 km NE, cording to Gardner and Patton (1972, p. 3), av- 1 (fmnh); Caldas: Rio Hondo, Samand, 1 (fmnh); erages 26.6% of tail length in nine specimens of Cauca: Cocal, upper Rio San Juan, 1 (amnh); La P. andersoni mcilhennyi from Balta and averages Boca, Rio Saija, 3 (fmnh); Upper Rio Cauca Val- 16.9% in 10 specimens of P. opossum quica from ley, holotype of grisescens J. A. Allen (amnh); the same locality. Ratios of a sampling from both Choco: Bagad6, 8 (amnh); Condoto, holotype of species with m'' fully erupted, measured in the dry melantho Thomas (bm[nh]); Novita, 1 (amnh); skin, are shown in Table 13. The data reveal that Rio Docampado, 3 (fmnh); Rio Sando, Rio Bau- the thickly furred basal portion is most extensive do, 6 (FMNH); Valle del Cauca: Buenaventura, 1 in the Balta series of P. andersoni mcilhennyi and (FMNH); Rfo Fn'o, 2 (amnh). ECUADOR. El Oro: the Brasilian, French Guianan, and Surinam P. El Chiral, 2 (amnh); Punta Santa Ana, 1 (amnh); opossum opossum and, with few individual ex- Salvias, 1 (amnh); Guayas: Bucay, Rio Chimbo, ceptions, is less in all others. The survey did not 8 (amnh); Pinas, 4 (amnh); Puente de Chimbo, 6 take into account seasonal differences, if any, in (amnh); Rios Chimbo-Coco, 6 (amnh); Ventura, 3 hair growth. Collectors' tail measurements record- (amnh); Loja: CeboUal, 1 (amnh); Manavi: La Pa- ed on field tags are usually reliable, or compara- paya, 1 (amnh); Rio de Oro, 1 (amnh); Rio Pes- ble, but may account for a few disparities in the cado, 2 (AMNH); Lim6n, 1200 ft, 3 (amnh, not lo- ratios. cated). Comparisons—The broad, strongly defined dark middorsal longitudinal band or stripe of long coarse, apparently erectile hairs is the principal external character P. andersoni Philander andersoni Osgood distinguishing from P. opossum. Sympatric and parapatric pop- under (Synonymies subspecies headings) ulations of the latter average slightly smaller in bodily and cranial dimensions (Table 14). Dorsum Distribution (Fig. 5)—Amazonian basin of of P. opossum is nearly uniformly grayish brown Peru, Ecuador, Colombia, Venezuela, and Acre in with never more than a faint indication of dark western Brasil; altitudinal range from near sea dorsal band, which often may be an artifact of level to about 1900 m above. dry, folded-over skin, hind limbs grayish or buffy, Cranial Characters—Cranial characters as not dark brown. for the genus. Sexual Dichromatism—Sexual dichromatism Dental Characters—Dental characters as for is most strongly marked in the Pucallpa, Rfo Uca- the genus. yali, series of andersoni. In females, pheomelanin Coloration—Dorsal surface of trunk with is saturate on underparts and dominant in the broad, blackish (dark brown) longitudinal stripe, banding of the agouti hairs of other bodily parts. or entirely blackish, the dominant guard hairs usu- In males, the agouti pattern persists on underparts, ally coarse, elongate and uniformly colored or with the pheomelanic subterminal bands thinly with a pale, narrow subterminal band; neck, chest, pigmented or pale buff. Irregular pheomelanic and belly often dark brown to blackish with gray patches on belly and chest are paler than in fe- basal portions of hairs showing through; lateral males. Underparts of one of two female andersoni line, if present, restricted to a poorly defined hip only from Santa Elena. Rio Samiria. are complete- patch; forefeet usually paler beneath than above, ly saturate (i.e.. monochromatic) pheomelanic. hind feet darker beneath than above; tail usually Sexual dichromatism in remaining material is parti-colored with proximal half to four-fifths not as clear, but in general pheomelanin saturation brown or blackish, remainder unpigmented, rarely in underparts and sides of body is dominant in entirely dark; ears usually blackish except some- females. In males, the trend is toward pigmentary times with unpigmented band or patch extending dilution on all parts except for pheomelanin sat- from base to mid-center or margin. uration of throat, midline of neck and chest, and Pelage—^The muskrat-like pelage, particularly pubic region. The intensity of sexual dichroma- of the dorsum, more so in P. andersoni mcilhen- tism varies from population to population. Adap-

HERSHKOVrrZ: PHILANDER 59 Table 13. Ratio of hairy basal portion of tail, in dry skin, to total length of tail (collector's measurement) in Philander andersoni and P. opossum. Figures are of individuals from each locality.

Taxon Locality Individual Ratios

P. andersoni mcilhennyi Balta, Peru P. andersoni andersoni

P. opossum opossum

P. opossum quica (selected specimens) 1972:5—PERU: Habits—According to notations on specimen Linnaeus), Ayacucho (San Jos^, Rio Santa Rosa. 10(X) m; Huanhuachayo, 1660 m); labels, individuals of Philander andersoni ander- comparisons. Reig et al.. 1977:197—PERU: Aya- soni had been shot out of trees and on the trapped cucho (San Sos6, Rio Santa Rosa; Huanhuachayo). ground. Philander —opossum canus Cabrera (part, not Osgood) At Tres Troncos, Rio Caquetd, Colombia, two 1958:35 nigratus Thomas only, a synonym. live suckling young, each about 6 cm in combined head and body length, were detached from their Type—Male, skin and skull. Field Museum of trapped, dead mother (fmnh 70985) and left to Natural History, no. 19655; collected 11 Septem- fend for themselves in camp, where I could ob- ber 1912 by M. P. Anderson. serve them. At first, they tottered uncertainly on Type Lcx:ality— "Yane Yaku" [= Yanayacu], weak legs, but quickly adapted to life without a chacra, or farm, on the Paranapura River about mother. They ate ripe bananas, plantains, and pa- 8 mi from Yurimaguas. The latter is on the Rio payas, lapped blood from animals being skinned, Huallaga at the mouth of the Rio Paranapura (cf. and ate some of the flesh. When not otherwise Osgood, 1914, p. 147). engaged the two fought with each other, the victor Distribution (Fig. 5)—Southeastern Colombia, putting the loser to flight. Both animals disap- eastern Ecuador, the Rio Huallaga Valley in the peared after 2 days. Peruvian departments of Amazonas, Pasco, Junin, and Ayacucho, the Rio Ucayali Valley, Ucayali, and the disjunct population of Territorio Federal Philander andersoni andersoni Osgood Amazonas, Venezuela. Characters—Dark middorsal band well de- ['!\D[idelph\s\ mxosuros, Tschudi (not Temminck), 1844:14, 145, 151—PERU; female with well-defined from sides and continuing over rump to less veloped pouch. Tschudi, 1844:250—PERU. than basal fifth of tail; hind limbs pale brown, Metachirus andersoni 1913:95; 1914:149— Osgood, grayish brown or buffy, forelimbs paler with PERU: Ucayali (type locality, "Yane Yacu," Rio digits usually unpigmented; ventral surface ochra- Paranapura, 8 mi from Yurimaguas). Sanborn, ceous to or buff. 1947:215—type history. orange buffy grayish M[etachirops] andersoni, Matschie, 1916:268—clas- Comparisons—Distinguished from Philander sification. andersoni mcilhennyi mainly by paler coloration [Holothylax] andersoni, Cabrera, 1919:47—classifi- throughout and narrower, more sharply defined cation. middorsal band; from of M[etachirus] opossum andersoni, Thomas, 1923: sympatric representatives 603—classification. P. opossum by larger size, dark middorsal band Metachirus opossum andersoni, Thomas, 1928b: and darker coloration throughout. 294—PERU: UcayaU (Iquitos). Measurements—Tables 1 and 15. Philander 1958:34— opossum andersoni, Cabrera, Variation—The Andean andersoni from classification. P^rez-Hemdndez, 1989:373—VEN- is dark like mcil- EZUELA: Territorio Federal Amazonas (Cafio Huanhuachayo, Ayacucho, Peru, Majagua; Cerro Cucurito; San Juan, Rio Manapiari; hennyi but with dorsal pelage shorter, finer, dorsal La Neb- Tamatama; Bel6n; Capibara; Campamento band not clearly defined from sides, underparts lina; Mavaca; Sierra Parima). and limbs paler, tail entirely pigmented except for P[hilander] opossum andersoni, Gardner and Patton, some on terminal A second An- 1972:5—PERU: Ayacucho (Huanhuachayo; San mottling portion. dean from San Jos6, is Jos6); characters; compari.sons. — specimen, Ayacucho, P[hitander] o[possum]. andersoni. Baker, 1974:135 slightly paler throughout except for a similarly ECUADOR: Napo-Pastaza (Santa Cecilia). dark brown tail. Its poorly defined dorsal band Philander andersoni, Emmons and Peer, 1990:18— and nearly uniformly dark tail suggest intergra- part. "upf>er Amazon Basin of Venezuela, probably dation with P. Nearest Colombia, Ecuador to Ucayali Department." opossum. geographic rep- Metachirus opossum nigratus Thomas, 1923:603— resentatives of P. opossum, however, from Cuzco PERU: Juni'n (type locality, Utcuyacu, 1600 m; and Madre de Dios are distinct. The third Andean Chanchamayo); type, male, skin and skull, British specimen of andersoni from Chanchamayo, Junin, Museum (Natural History), no. 1900.7.7.62; col- and another from San Juan, Oxapampa, also lack lected 21 April 1900 by P O. Simoms. Thomas, blackish dorsal band of the sub- 1928a:250, footnote—PERU:— Junin (Inaflez). the well-defined Gardner and Patton, 1972:5 "more likely a syn- species. In all other respects, they agree. The con- of andersoni" than P. canus, onym opossum apud dylobasal length of the last two, 76.3 and 81.2 Cabrera, 1958. mm, respectively, are among the largest of the Metachirus opossum Tate, 1939:161—VENEZUELA: The furred basal of the Amazonas (Mt. Duida). species. heavily portion Philander opossum, Gardner and Patton, part (not tail is notably extensive in the San Juan individ-

HERSHKOVrrZ: PHILANDER 61 Table 15. Measurements of Philander andersoni andersoni. Table 15. Extended.

Preorbital Table 16. Measurements of Philander andersoni mcilhennyi.

Locality Table 16. Extended.

Preorbital Width trap lines with species of Metachirus, Chironec- Blood Values tes, Didelphis, Caluromys, Monodelphis, and mouse opossums (Marmosinae). All inhabit the Tyndale-Biscoe (1980, p. 718; COLOMBIA) same forests. Blood values measured at sea level (Buenaven- tura), 500 m above (Villavicencio), and 1600 m above (Call) revealed correlations with age and Barbara: The Pale-Phase altitude. Hemoglobin concentrations and hemato- Four-Eyed Opossum crit values were lowest at lowest altitudes, and those for pouch young lower than for adults.

Hershkovitz (field notes, October 1992)

A pale-phase female was live-trapped 17 Oc- Breeding tober 1992 in a wooded area about 1200 m above sea level in the Nacional do east- Parque Caparao, Natterer (in Pelzeln, 1883, p. 110—Sepetiba, Rio em Minas Gerais, Brasil. Her nearly completely de Janeiro, BRASIL) closed pouch sheltered six attached young, pos- Female captured 8 March 1818 with five naked, sibly no more than 1 week old. She was kept in blind pouch young. her cage for observation of behavior and litter de- Female captured 29 September 1822 in Ipane- velopment. Dubbed Barbara, she accepted captive ma, Rio de Janeiro, with four blind pouch young life with equanimity, making no attempt to escape ("D. dichrura"). even when the door of her cage was opened for feeding or cleaning. She ate everything served, Miller (in J. A. Allen, 1916c, pp. 563, 589- -Uruc- whether beef, chicken legs, insects, peanut butter, um, Mato Grosso, BRASIL) bananas, or mice. Live mice dropped into the cage were the head and neck seized instantly attacked, "Number of young is usually small, between and crunched in her The mice were com- jaws. three and eight." pletely devoured with nothing left of flesh, en- trails, skin, or bones. Davis (1947, p. 2—Teresopolis, Rio de Janeiro, Barbara slept most of the day curled up under BRASIL) the cut grass provided for nesting. Prepared food Seven of 32 females in or raw fruit given during the day was usually con- captured Teresopolis had number, 4.5. Two fe- sumed at night. Water was imbibed at all hours. pouch young, average males each had seven and seven in Although normally nocturnal, should a live insect young nipples in and continues or mouse be dropped into the cage during the day, pouch. Breeding begins August until Barbara would awake with a start and make an February. instant kill and a fast meal. She easily devoured Fonseca and Kierulff 1 19- -Atlantic for- an entire mouse 10 to 15% of her own weight and (1989, p. looked for more. est, Minas Gerais, BRASIL) When food Barbara would fix her wanting eyes Female captured February and another in Au- on the nearest raise her muzzle, and sniff person, gust had five pouch young each. expectantly. On one occasion Barbara left intact hide and Cerqueira et al. (1993—Rio de Janeiro, Restinga fur of a mouse otherwise consumed. completely de Barra de Marica, BRASIL) Barbara washed her face frequently after and during meals. One or both paws were used. Grooming forequarters, sides, and underparts usually followed in that order without particular attention to the pouch and none to the young. On the 26th of October, head-rump length of the six still-attached pouch young ranged from 1 to 2 cm. Mother and offspring were transported the next day to Rio de Janeiro, where facilities were available for their care. Tyndale-Biscoe (1980, p. 713—Buenaventura, 2. Omnivorous but 2. Mostly frugivo- Valle, 10 m; near Villavicencio, Meta, 500 m; largely frugivorous rous, nectarivorous near Call, Valle, 1000 m, COLOMBIA) 3. Food availability 3. Food availability variable in under- less variable in up- Four females captured August and September growth where for- per strata where 1971, one pregnant, three lactating, and each with aging is done foraging is done five pouch young. 4. Birth to weaning 4. Birth to weaning 68-75 days 110-125 days et al. 425- -He de Charles-Dominique (1981, p. 5. Size 50- 5. Size 50- GUYANE weanling, weanling, Cayenne, FRANgAISE) 75 g 75 g 6. at sexual ma- 6. at sexual ma- Litter size, 4.24 (1-7). Age Age 6 months almost 10 Litters mature in 70 days. turity, turity, months Lactating females three of 16 in June, two of 7. to three succes- 7. Two successive lit- 13 in July, zero of nine in August. Up sive litters in same ters in same year Pouch young survival and food availability: = (third litter first litter following dry season, 97% (n 36); year = second litters, 97% (n 38); third litters, food mostly pouch- aborted) scarce, 28% {n = 67). 8. Behavior as r-strat- 8. Behavior more like Interval between litters, 90 days. that of Sexual maturity, 150 days. egist' /:-strategist^ 9. 8- 9. Nesting period, Seventy-nine percent of October-November Nesting period, = 15 days 30-45 days population (n 42) bom in preceding season. 10. Litter size not 10. Litter size not Ovarian cycle inhibited by lactation but not by more than 7 more than 7 gestation.

' r strategist: r-selected species, selection favoring a Charles-Dominique (1983, p. 409, GUYANE rapid rate of population increase. Typical of species that FRAN(;AISE) colonize short-lived environments or of species that un- dergo large fluctuation in population size. - k ^-selected selection Didelphoid breeding is continuous throughout strategist: species, producing superior competitive ability in stable, predictable envi- the year, but successful reproduction is dependent ronments in which rapid population growth is unimpor- on a rich diet for the female and is seasonal. Up tant as the population is maintained at or near the car- to three successive litters may be produced in 1 rying capacity of the habitat. year. The third litter "generally occurs at the beginning of the period of food scarcity and in most Atramentowicz (1986a, p. 125, GUYANE FRAN- cases the females lose weight and young die in gAISE) the pouch at a more or less early stage. Excep- During a study period from September 1978 tionally, another litter can follow and abort, but through October 1979 in Cayenne, two successive generally reproduction is interrupted until the be- litters were produced between September and ginning of good recrudescence of fruiting. Among June. Some females, however, produced three lit- the Didelphidae, P. opossum behaves more like ters in the same period. The difference is attrib- an r-strategist in favoring rapid population in- uted to an abundance of food and a shortened crease in an unstable environment." nesting period. In the event of a food shortage, milk production ceases. Death of lactating young Charles-Dominique (1983, p. 409, GUYANE starved at the nipple is termed pouch abortion. FRANgAISE) Average adult weight, 400 g. Comparison of breeding conditions between Atramentowicz GUYANE FRAN- Philander opossum and Caluromys philander of (1986a, p. 123, Guyane Fran^aise, two similar-sized species oc- gAISE) different niches: cupying ecological During a 26-month (1978-1982) investigation of didelphoid breeding, "animals were live Philander opossum Caluromys philander trapped, marked and then released. Caluromys I. Mainly terrestrial 1. Almost entirely ar- philander. Philander opossum, and Didelphis boreal marsupialis are nocturnal, with a mixed frugivo-

HERSHKOVrrZ: PHILANDER 67 Table 17. "Reproductive condition of 22 adult females of Philander opossum from Nicaragua" (Phillips & Jones, 1969, p. 345).

Date of Capture ference in weight of testes of mature animals Density throughout the year. Discrepancies noted in paragraphs one and two Charles-Dominique et al. (1981, p. 383, Cayenne, above are confusing. Perhaps the mixture of data GUYANE FRANgAISE) for Philander and Didelphis is the cause. Biggers 100-200 individuals per km-. refers readers to the original compilation of data.

Enders (1935, p. 41 1, PANAMA) Diet Only nonbreeding females captured January to

April; starting July, "females with litters outnum- Temminck (1827, p. 38) ber those without; litters of two, four, and five It on small birds and insects but also eats were observed." preys fruit. In captivity it feeds on meat.

Enders (1966, p. 198, PANAMA) Pelzeln (1883, p. Ill, BRASIL) "Observations on 18 captured Philander indi- Insects found in stomach. cate that their breeding season . . . begins with in f onset of the dry season February." Goldman (1920, p. 52; PANAMA) I "The largest litter encountered was five." "The stomach of one taken at Gatun was well Fleming (1973, p. 444, Canal Zone, PANAMA) filled with fragments of crabs. Stomach contents of several others at the same locality were filled Females were reproductively active during with fragments of birds alone, or of birds includ- June, July, and September through December in ing their feathers, and crabs intermixed." 1966 and during February through April and June in 1967. Females carrying pouch young or lactat- Fleming (1972, p. 623; Canal Zone, PANAMA) ing in October and November were anoestrus. Two, possibly more, litters are produced per sea- "Stomachs of the four-eyed opossum contained son. unidentified plant material, the pulp of Corozo "Males of P. opossum are sexually mature at a oleifera nuts, insect and fresh water shrimp re- body length of 230 millimeters or greater . . . mains, murid rodent fur, and the tail of a young adult males probably are sexually active through- spiny rat (Proechimys semispinosus). One individ- out the year." ual was seen feeding on live bats caught in a mist Litter size 4.6 (2-7) 34; sex ratio 22 (45%) 6 6, net." 27 (55%) $9.

Schomburgk (1840, p. 344; GUYANA) Seba (1734, p. 57, SURINAME) "They are very destructive to poultry and like- Eight pouch young were counted. wise to fruit. They are often found on those sa- Seba and Miller (above) are the only authors to vannahs where the wild pine (Bromellia sp. ?) record eight pouch young. Maximum number in flourishes, to the fruit of which they appear to be all other cases is seven, in agreement with number partial." of nipples. Philander, unlike Didelphis and oth- than can be ers, may not produce more young Enders (1935, p. 410; Canal Zone, PANAMA) suckled. They "are accused of being poultry- and bird- killers. ... In captivity they were more carnivorous than of the other observed. Cannibalism any opossums They ate meat of all kinds including ant-eater, car- casses of rodents, grasshoppers, and eggs in pref- Charles-Dominique (1983, p. 405; Cayenne, erence to any kind of fruit, although they did eat GUYANE FRANgAISE) banana, papaya, pineapple, and figs. One hungry Many cases of cannibalism have been observed individual, while fearful of the hand offering a in captivity of weaned young being eaten by other ripe banana, nevertheless sniffed from a safe dis- young and the mother. tance while saliva dripped from its jaws."

JHERSHKOVrrZ: PHILANDER 69 Dalquest (1953, p. 19; eastern San Luis Potosi, fers fruits but takes earthworms and small verte- MEXICO) brates. Marmosa cinerea [= Micoureus demerarae] and C. philander are vegetarian. ''One animal became entangled when it at- "The period of food scarcity determined both tempted to climb a net to obtain an entrapped bat, by records of fruit and insect abundance and also others were taken in traps baited with decayed by body weight changes in the marsupials begins meat, and two were shot while they were feeding in May, is most critical during June-July-August, on small wild figs. Remains of insects were usu- and ends in September. Data collected in 1976, ally present in the stomachs of the specimens and from 1976 to 1982 by the Institut Pasteur de taken." Cayenne (variation in weight and reproductive condition of marsupials) suggest that this period Anthony in Goodwin (1934, p. 5; GUATEMALA) of food scarcity is a regular phenomenon, proba- correlated with seasonal variations in rain- "I was told that these little opossums are often bly fafl." killed in the sugar house [trapiche] and that they are quite fond of sweets. ... At Finca Capres, et al. several times I surprised them at night, feeding on Charles-Dominique (1981, p. 376; Cayenne, GUYANE the ripe [bananas]. ... At Chipore . . . one was FRAN^AISE) attracted by bait composed chiefly of peanut but- "Eighty percent of stomach contents consisted ter and rolled oats." of a great variety of prey. During the rainy season, earthworms, some of which are as much as 80 cm Hall and (1963, 198; Veracruz, MEX- Dalquest p. long, forced above ground, made up a large part ICO) of the diet. "Fallen fruit consumed were Attalea re- "It has been seen feeding on sweet-lemons, mainly Virola melinonii, Virola Virola sur- jobo plums, and the fruit of the Chico Zapote (Sa- gia, sebifera, inamensis, Ocotea Richardella macro- pote achras) source of chewing gum." One opos- puberula, phylla, Ficus spp., Cordia exaltata, Simarouba sum "was seen at the base of a large, hollow fig amara, Protium heptaphyllum. In the tree. The upper part of the hollow in the tree general pulp is eaten with the smallest seeds {Ficus, etc.) served as a retreat for a colony of the large fruit only swallowed, the ones out. The bat, Artibeus jamaicensis. Bats were bringing larger spit ground level flowering of the Balanophoracea from Jan- small green figs into the hollow and feeding on to June is actively sought for the nectar. Los- them. Parts of the fruit, varying in size from al- uary es weight during dry season (September-Novem- most whole figs to mere shreds, were dropped by ber in the 1978, 1979) when fruit is scarce, the bats. The four-eyed opossum was feeding on years fatten when fruit is abundant." these bits of figs." The opossum "followed our trap lines, eating (1980, 716; COLOMBIA) mice and other small mammals that had been cap- Tyndale-Biscoe p. tured. These were us- opossums easily trapped by "They would eat fruit and vegetables provided ing flesh, preferably much decayed, for bait." as well as meat; chicken bones were held in the hands and the ends chewed." Dubost (in Charles-Dominique, 1971, p. 197, GUYANE FRANgAISE) Atramentowicz (1986a, p. 125; Cayenne, GUY- ANE FRANgAISE) Philander eats tree exudate, and like the pri- mate Cebuella or Microcebus (Callitrichidae) (Le- Eats ripe pulpy fruit, insects, and small prey uses its teeth to muridae) reopen healed openings such as earthworms, frogs, etc. in tree trunks to renew the flow of sap.

Atramentowicz (1988, p. 48; Cayenne, GUYANE Charles-Dominique (1983, pp. 397, 398; Cay- FRAN^AISE) enne, GUYANE FRAN^AISE) Stomach contents of Philander opossum consist The diet is opportunistic, consisting of fruits, of 50% fruit with the remainder invertebrates, flowers, nectaries, insects, earthworms, small ver- small vertebrates, and carrion. A preference is tebrates, and carrion. Sympatric D. marsupialis shown for ripe fruit fallen to the ground. The fruit subsists on about the same; Marmosa murina pre- consumed represented 44 species in 21 families.

70 FIELDIANA: ZOOLOGY It was characterized by a fleshy pulp with a high yellow under the hunting lamps while the eyes of percentage of water The color may be bright or [Metachirus] show red." dull, size large or small, shape variable. The pulp be rich in or but in may sugar lipids poor nitrogen, Hershkovitz (field notes) which the animal hnds in animal prey. The eyes of Philander, like those of other didelphoids and Metachirus, usually glow bright or- Display ange when reflecting the beam of a battery-pow- ered flashlight. There is individual variation in Hershkovitz (field notes) tone of eye shine depending on the amount of environmental light, intensity of directed light, and angle of reflection. These factors and the un- The supraorbital spots are conspicuous at night, controlled conditions make it difficult to find sig- at least to the sharp-eyed (see "Sleep," p. 77). nificant differences in color of didelphid eye Conspicuous supraorbital marking displayed by shine. More important for identification is dis- both nocturnal and diurnal animals may serve for tance between the eyes and animal gait as seen recognition between consjjecifics. I find no differ- by the reflected light. ence in markings between the sexes, and the signs are usually absent or faint in juvenals.

Fat Storage Enemies, Predators

Hall and Dalquest (1963, p. 198; Veracruz, MEX- Most likely predators are the eyra cat {Her- ICO) pailurus jagouaroundi) and smaller spotted cats, "In the lowlands these seldom were large mustelids such as the tayra {Eira barbara) opossums fat, but in the at 5000 feet elevation and grison (Galictis vittata), foxes, large owls, highlands, and higher, in winter, had a layer of and snakes. Husson (1978, p. 26) found remains they deep [!] fat beneath the skin." of a Philander opossum in the stomach of a gar- yellow immediately den tree boa (Corallis enhydhs) in Suriname.

Schomburgk (1840, p. 344) reported that "they are sometimes eaten the Creoles and Indians by Foraging (Table 18) [of Guyana], but as they have a rank and disa- greeable smell I doubt if they would prove pal- Charles-Dominique et al. (1981; Cayenne, GUY- atable to us." ANE FRANgAISE) Wilson (1970, p. 386) reported an encounter in Panam^ between and Phi- Didelphis marsupialis Of a total of 294 individuals live-trapped or lander with the former and opossum overcoming tracked by radio, 1 85 were located from to 6 m skin, flesh, bones, and brain of the lat- devouring above ground in the following situations: on ter (see details under Reaction, De- "Aggression, ground 70%; fallen logs 7%; small tree trunks fense"). 2%; branches 6%; lianas or vines 15%.

Charles-Dominique et al. (1981, p. 343; Cayenne, Shine Eye GUYANE FRANgAISE)

in 1 Anthony Goodwin ( 934, p. 5; GUATEMALA) Nearly always forages on the ground; of 100 individuals live-trapped only three were captured "In the beams of the carbide hunting lamps, above ground. their eyes were extremely bright, gleaming white, like electric lamps, easily seen at one hundred yards or more." Charles-Dominique (1983, p. 397; Cayenne, GUYANE FRANgAISE)

Enders (1935, p. 412; Canal Zone, PANAMA) See Table 1 8 for body mass and above-ground The eyes of [Philander] show up as a reddish foraging by five syntopic species of opossums.

HERSHKOVITZ: PHILANDER 71 Table 18. Body mass and above-ground foraging by five syntopic species of opossums.

Taxon Weight (g) Substrate

Marmosa murina 40 Low (shrub) Mamiosa cinerea {- Micoureus demerarae] 80 High Caluromys philander 300 High Philander opossum 400 Low (essentially ground level) Didelphis marsupialis 1,000 Low (may climb to feed on fruit)

Grooming meters from the nearest water. These records are unusual, however. Against them are nearly 30 re- Hershkovitz (field notes, January 30, 1941; north- cords from in and near water." ern COLOMBIA)

Dalquest (1953, p. 19; eastern San Luis Potosi, A washed face with hands mouselike; captive MEXICO) groomed remainder of body with tongue catlike.

"Found only in the tropics of the eastern part

Charles-Dominique (1983, p. 407; Cayenne, of the state . . . are taken in dense GUYANE they vegetation FRANgAISE) growing along the shores of streams and rivers at elevations between 400 and 12,000 feet [more "We have never observed any allogrooming likely 1200 feet]. At El Salto the water is cold between [caged] adults, even during mating periods." and flows swiftly between rocky canyon walls; opossums were found in thickets of ferns, vines, and low, woody plants. Near Huichihuayan, Habitat where the water is warmer and sluggish, opossums were taken in thickets of thorny bamboo."

Hall and Dalquest (1963, p. 196; Veracruz, MEX- ICO) Goodwin (1946, p. 284; COSTA RICA)

"... ranges throughout the tropics of Veracruz. "Frequenting forested country from sea level At the extreme upper edge of the upper humid up to about 4000 feet, he is most common at low division of the Tropical Life-zone, it lives along elevations." cold, clear streams at the edge of the oak belt. Lower down but still in the upper humid division, Davis (1947, p. 2; Teresopolis, BRASIL) it was found along rivers and streams that flowed "Found more commonly in moist situations al- through dense jungle, where the tall, broad-leafed though individuals may wander through nearly trees were thickly hung with orchids, vines, moss- any kind of vegetation." es and bromeliads. The four-eyed opossum was found living in the thickets bordering the broad Enders (1935, p. 410; Barro Colorado Island, rivers of the coastal plain, in the arid division of the Tropical Life-zone, and along the marshy PANAMA) shores of rivers and streams of the lower humid "They are not common on the Island for while division of the Tropical Life-zone, in the southern they enter traps without hesitation none were of the state. part taken; but many were captured at Alhajuela in "Most of our specimens were taken on the very habitats similar to those on the Island . . . why shores of rivers or streams . . . however . . . the they are not abundant on the Island is a puzzle." species is not confined to such habitat . . . seven kilometers west of Potero, workers discovered a Schomburgk (1848, p. 777; GUYANA) family of four young animals in a field of sugar cane, several kilometers from the nearest water at "It lives mainly in the coastal forests bordering that time of the year. At Jimba, 350 feet elevation, the plantation. During the day it sleeps most of in southern Veracruz, a four-eyed opossum was the time in its haunt under tree roots or in hollow taken from a tree on a hillside fully three kilo- trees."

72 FIELDIANA: ZOOLOGY Natterer (in Pelzeln, 1883, p. Ill; southeastern 10 to 16 times greater in secondary than in pri- BRASIL) mary forest.

". . . Fashions its nest from leaves on low Husson (1978, 26; trees." p. SURINAME) Some of the.se opossums were found under the J. A. Miller (in Allen. 1916c, p. 589; Mato Gros- floor of a home at Lelydorp. so, BRASIL)

Atramentowicz (1986a, p. 123; GUY- ". . . is found in forests. I have never Cayenne, deep ANE known it to venture near houses for the purpose FRANgAISE) of hen-roosts and nests like its rel- robbing larger Philander opossum is mainly terrestrial "but ative it do [Didelphis marsupialis], although may may be seen climbing to the canopy" in the sec- so." ond growth forest.

Handley (1976, p. 8; VENEZUELA) Da Fonseca and Kierulff (1989, p. 118; Atlantic Lowlands of western and southern Venezuela, forest, Minas Gerais, BRASIL)

46 specimens: on the ground, 98%; on a log, 2%; "Fourteen individuals were caught during the near streams and other moist areas, 100%; ever- [17-month] course of this study. The occurrence green forest, 91%; orchards, croplands, yards, of this large didelphid is apparently tied to the open areas, 9%; elevation 24-324 m. presence of standing or running water. ... As only a few transects occurred close to streams, this may Hall and Dalquest (1963, p. 198; Vera Cruz, explain the low trapping success for this species." MEXICO) "The species also proved to be primarily terrestrial with 17% of in arboreal "... are usually seen or trapped on the ground, only captures and 7% of individuals climbed trees after but are sometimes seen in trees." They prefer the traps, released." vicinity of streams and seem to be as much at being home in rivers as on their banks. Unpublished (VENEZUELA) Enders (1935, pp. 410, 411; Barro Colorado Is- Capture sites recorded by members of the land, Canal Zone, PANAMA) Smithsonian Venezuelan Project (SVP) on field None "were taken in the Island but one was tags of captured specimens are summarized below: observed high up on a limb on Zetek Trail. . . .

While a good climber [it] was usually encoun- A^ tered upon the ground, on or under logs. It is in banana probably more terrestrial in habit than any of the Live-trapped other opossums studied excepting possibly Didel- phis. This might be surmised from its build, which is well adapted to terrestrial locomotion."

Davis (1947, p. 2; Teres6polis, BRASIL)

"These opossums seldom climb but are partial to fallen logs and windfalls."

Farris (1950, p. 259; PANAMA)

"About eighteen females were collected in grass fields about 47 miles from Panama City in a location that is low and damp all the year."

Charles-Dominique (1983, p. 419; Cayenne, GUYANE FRANgAISE)

The frequency of encounter with Philander is overhanging rivers. This type of habitat restricts by new individuals, but only one-third of the orig- enemy attacks to the forested side of the river and inal C. philander population was replaced." provides the opossum with the opportunity of es- musicrat-like caping by diving into the water. The Atramentowicz (1986b, p. 145; GUYANE FRAN- dorsal pelage of P. andersoni mcilhennyi and gAISE) many individuals of P. andersoni andersoni Didelphoids are solitary, not territorial, and seems to be better adapted than that of P. opossum many individuals are sedentary. Duration of resi- for aquatic life. Where the two species occur to- dence in study area is compared with Caluromys gether, as in Balta and along the Rio Ucayali in philander. Philander opossum, and Didelphis Peru, P. andersoni may be more arboreal-aquatic marsupialis (Table 19). than terrestrial, whereas P. opossum may be more terrestrial-aquatic than arboreal. Charles-Dominique (1983, p. 418)

All American opossums are nocturnal and sol- and Home Range Territoriality itary except during mating and maternal care of young. Territory is not defended. Home ranges Miles etal. (1981a, BRASIL) overlap between many individuals, male and female, but with close contact avoided. There are A small, transparent plastic spool, 6 cm long no social bonds between adults from short with 2.2 mm internal and 2.4 mm external diam- apart copulatory periods; there is no allogrooming. eters, was wound with double-strand teryline thread. The spool was attached to a captive Phi- lander opossum, and the free end of the thread (see also Habitat) was attached to the capture trap or nearby vege- Hydrotropism tation. As the released animal traveled through the Hershkovitz field night the thread unwound. The thread tracked dur- (1962, notes; SURINAME) the led to the diurnal nest or Dis- ing day refuge. A female with 3 pouch young taken live 20 tance traveled was calculated from the weight of January 1962 from a snap trap in Lelydorpplan the before release of the animal and after spool was caged for observation. The animal never re- recapture. covered from the effects of the blow on the head Of 21 released animals, 16 were retrieved. The received from the trap. She slept most of the time calculated distance for 15 retrievals averaged 438 and when awake walked slowly with poor coor- m (93-1000 m). This compared with a mean of dination. She permitted herself to be petted and 801 for m (51-2450 m) Didelphis marsupialis. to seemed enjoy it, but the docility probably owed . to her weakened condition. She ate the insects fed Charles-Dominique (1983, p. 401; Cayenne, to her and seemed to amounts GUYANE require large of| FRANgAISE) water. A Marmosa murina held captive at the same time derived all its water from fruit. Philander explores about 25,000 m- during the The Philander twice from her On night. escaped cage. each occasion she out the nearest dark, "By systematically trapping, marking and releas- sought moist The first time she curled in ing we have observed regular loss of individuals, cool, refuge. up an but still-wet canvas bucket in the kitch- compensated by immigrants from other areas (ju- empty en. The second after I had moved to a frame veniles and adults of both sexes). Trapping was con- time, house in La Poule, she into a wet ducted using Tomahawk and Sherman traps placed snuggled dark, corner behind a flush toilet. on the ground and special 'home-made' traps set at primitive 15 to 20 m in trees; traps were baited with bananas

. . . Fonseca and Kierulff Atlantic for- The following data concern new-arrivals [in an (1989, p. 118; over 20-ha study area] from May to October 1979, est, Minas Gerais, BRASIL) after 8 months of regular trapping and marking": "The occurrence is apparently tied to the pres-' May, eight immigrants; June, 2; July, 1 ; August, 3; ence of standing or running water." September, 4; October, 3. "Migration seems to be more important in P. Tyndale-Biscoe (1980, p. 716; COLOMBIA) J opossum than in C. philander—after one year, the entire population of P. opossum had been replaced "All these animals were caught in second

74 FIELDIANA: ZOOLOGY r

growth forest close to streams, in several caves Hershkovitz (1941, field notes; northern COLOM- less than a meter from water. It was thus interest- BIA) ing to observe that a pair in captivity would in- The prehensile tail was used for swinging the variably defecate into the water container rather from limb to the hands for than elsewhere in the enclosure." body limb, grasping the next higher support. The animal would also use the tip of its tail for clasping an overhead limb, then climb its tail until it could the Locomotion grasp limb above with its hands. Carrying an opossum by its tail can be hazardous. Marmosids habitually Dalquest (1953, p. 19; eastern San Luis Potosf, use the tail for climbing. MEXICO) When attacking live prey, the opossum springs

^ for throat or head with They "are swift in their actions. For the most lightning speed. part they hunt on the ground, but they climb readi-

ly, often to considerable heights, and are skillful swimmers. Animals twice escaped capture by div- Longevity ing into the water of the Rio Naranjos and swimming away beneath the surface." Farris(1950, p. 258)

Life span in captivity, 3 years, 6 months. Crespo (1950, p. 6; Misiones, ARGENTINA)

They "are excellent swimmers, water appar- Atramentowicz (1986b, p. 140; GUYANE FRAN- ently being their preferred habitat. I have watched gAISE) them at night swimming swiftly even upstream Based on tooth wear, longevity of wild Philan- against the current in rivers such as the Uruguai." der opossum and Didelphis marsupialis was calculated as 2.5 years; of Caluromys philander, as Hall and Dalquest (1963, pp. 197, 198; MEXICO) 3.5 years. Captive Caluromys philander (Brunoy, '"Philander is quick and active. Trapped indi- France) lived to 6 years. Maximum life expectan- viduals are able to jump about and twirl in sur- cy of Didelphis marsupialis was said to be less than 36 months. prising fashion. . . ." The animal "is an agile climber and a skillful swimmer." "Along the Rio Atoyac several four-eyed opossums were taken in a trap set beneath the water Maternal Care level, at the base of a cut-bank. They could have reached the trap only by swimming." Seba(1734, p. 57) "... A four-eyed opossum was seen just before "The mother does not remove the from midnight, running swiftly over the larger, rounded young her before are old to the boulders (six to 18 inches in diameter) along the pouch they enough enjoy of When the time is she retreats river bank. When frightened [by us] the animal light day. right to a lookout to assure the of her She turned [from the river's bank] and made a smooth, safety young. then the and the to clean dive into the swift water, and as it did not opens pouch permits offspring into the and with her. At the reappear, must have swum away underwater." emerge sunlight play slightest suspicion of danger she calls her young with a warning cry that sounds tik, tik. tik. The Hershkovitz (field notes) young respond immediately by returning to the The usual didelphid locomotor pattern on the pouch whereupon the mother runs to a hiding ground is a measured walking gait, and in climb- place." ing an alternating movement of the grasping Comment—Seba actually beheld the mother hands and feet with support of the clinging, pre- and her brood at one time or another. The fanciful hensile tail. The principal running gait on or description of their relationship (translation), above ground is a trot. I have never seen a didel- however, must have been confected long after the phid leap or bound except in rapid escape. In de- supposed facts. scending a tree trunk the animal will proceed head Didelphids are nocturnal, but in captivity, es- first or simply drop to the ground. pecially as household pets, as Seba's opossums

HERSHKOVITZ: PHILANDER 75 must have been, they generally adjust to the hours Dalquest (1953, p. 19; eastern San Luis Potosi, they are fed. MEXICO)

"A burrow near Huichihuayan was in muddy soil beneath a of and an- Nest clump thorny bamboo, other was beneath a decayed log, nearly concealed by vines and succulent plants. This burrow was Goldman (1920, p. 52; PANAMA) so shallow that when the log was rolled aside the "A nest of one of these opossums was found entire burrow was exposed. A nest at the end of three feet from the ground on a fallen log. The the branched burrow consisted of a formless mass log lay in the dense thicket of an old clearing and of dry leaves a foot in diameter. Two feet away was heavily overhung with vines and bushes. The there was an opening to the dense vegetation be- nest, globular in form and about a foot in diam- side the log. Three feet farther on, the burrow eter, was placed in a well hidden spot among the emerged from under the log; and a trough like vines. It was made entirely of the banana-like trail led through the vegetation to the river, twenty leaves of a native plant rather neatly laid together. feet away." The opening at one end faced outward along the

. . . The nest was clean and about the log. cavity Murie (1935, p. 16; GUATEMALA) size of the animal's body." "In the edges of the pine ridge, especially near the streams, were numerous well worn trails about Carriker (in J. A. Allen, 1911, p. 247; Yuruan, VENEZUELA) five inches wide, which were apparently made by these opossums. Many of the trails led from one "The female with a litter of was taken young ground burrow to another. At the end of one of out of the hollow stump in a grass field in the the burrows about fifteen inches from the en- [forest] clearing." trance, I found a nest ten inches in diameter, composed of dry sedges." Hall and Dalquest (1963, p. 197; Veracruz, MEX- ICO) Enders (1966, p. 201; PANAMA) "Several four-eyed opossums were taken in a After the young are born the female moves trail leading from a dense thicket ... to a stream about more. Unlike habits during the rest of the ten feet away. ... A hole, about five inches in year (Enders, 1935, p. 397), they frequent the diameter that led downward beneath the roots of same nest. Later, when the young are able to de- a tree was discovered near the center of the thick- tach themselves and so leave the mother, young et. This seemed to be the home of at least one of may continue to use the nest. the opossums." Two "were found in a nest that they had con- Husson (1978, p. 26; SURINAME) structed in the palm thatch of the roofs of aban- doned houses. These nests consisted of a handful Nest with female and seven young found under of dry leaves, in between the of pushed layers a tree. Nothing more is said. palm fronds. From outside, a distinct spherical or oval lump in the thatch marked the site of the Miles et al. (1981a, p. 341; BRASIL) nest. Inside the house we could see no trace of the nests." "Nests of the opossum were often found 8-10 A nest "was found in in a cavity the side of a m above the forest floor in hollow trees or as open of tree piece trunk, 15 inches in diameter and nests in tree forks; some nests were, however, ter- three feet long, that was in the air in suspended by restrial, cavities besides buttressed tree roots." ! vines. . . . The nest was of dry leaves, about 1 1 inches deep and seven inches in diameter." i > Farris (1950, p. 259; PANAMA) One animal "was shot from the large hollow in the side of a tree. giant 'ligaron' [higaron?] This "The animals usually were found in their nests i tree was 1 2 feet in at fully diameter waist height, in palm trees, at the branching and in the leaves I and contained a hollow about 60 feet high and about 5 feet off the ground where conditions were j five feet in diameter." relatively dry."

76 FIELDIANA: ZOOLOGY Population Density ed for comparison. These sympatric Didelphids are nocturnal, with a basically frugivorous and in- Charles-Dominique et al. (1981, p. 383; Cayenne, sectivorous diet. The three main species have high GUYANE FRANgAISE) densities, high reproductive rates, short life-spans, and rapid population turnover. There are some dif- Philander opossum was one of the most abun- ferences in these parameters, mainly between the dant mammals on the island. Its biomass, based arboreal species, C. philander, and both terrestrial on 199 observations within a square kilometer, species, D. and P. was calculated as between 100 and 200 individ- marsupialis opossum. Although all have few social interactions and show a lack uals, with a biomass between 40 and 60 kilograms of territoriality, C. to have a lon- in the same area. The biomass of each of the five philander appear ger life-span, a lower reproductive rate and a other marsupials of the same area was 25 to 50 more sedentary population. M. cinerea [Micou- kg for D. marsupialis, 30 to 60 kg for C. philan- reus demerarae] were very rare in this area, der, 2 to 8 kg for Marmosa cinerea [Micoureus whereas M[armosa]. murina show demerarae], and 0.7 to 3.5 for M. murina. large density variation." Of the six marsupial species inhabiting the is- The well-known tendency of American marsu- land. Philander, Didelphis, Caluromys, and M. = pials to enter baited traps, usually the same ones, murina were most abundant. Marmosa cinerea [ time and again ensures a high degree of accuracy Micoureus demerarae] were moderately abun- in estimating their duration of residence in an dant; Monodelphis brevicaudata, uncommon. area. In general, both sexes of P. and C. philander are highly sedentary. Didelphis marsupialis, on Cerqueira et al. (1993, p. 513; Restinga de Barra the other hand, tends to wander, particularly the de Maricd, Rio de Janeiro, 22°57'S, 42°5rw, males (Table 19). an impoverished small animal community, At- lantic forest, BRASIL)

Average density 1.91/ha; population main- Posture tained mostly by recruitment. Enders (1935, p. 412; Canal Zone, PANAMA)

Atramentowicz (1986b, p. 144; GUYANE FRAN- Philander "used the tripod [sitting] posture gAISE) more frequently than Metachirus although the tail Density per square kilometer based on obser- of the latter is a very effective appendage." vations from September 1978 to October 1979 was 137 P. opossum, 143 C. philander, and 45 D. Hershkovitz (1941, field notes; northern COLOM- marsupialis. BIA)

Sitting was tripodal, the body supported by hind limbs and tail. Population Dynamics

Atramentowicz GUYANE (1986b, FRAN^AISE) sleep

"Five species of Didelphid marsupials were Temminck (1827, 38) trapped live, marked, and released in a secondary p. forest in French Guiana a 26-mo field during It sleeps during the day rolled up into a ball. Data on dental study. body weight, body length, Its breathing is like that of a ferret, and it leaves and state of females were col- stage, reproductive its hiding place only at night. lected. A total of 85 1 individuals, including 372 pouch-young, were monitored in 2273 captures. Schomburgk (1840, 348; GUYANA) This abundance of data we made led us to a com- They sleep during the day under tree roots or parative analysis of the population dynamics of in hollow trees and hunt at night. three species: Didelphis marsupialis. Philander opossum, and Caluromys philander, which rep- Hershkovitz (1941, field notes; northern COLOM- resented 94 percent of the whole captures (Table BIA) 19). Some data on the two other species, Mar- mosa murina and M[icoureus]. cinerea, were add- The animal sleeps rolled up on its side or ex-

HERSHKOVITZ: PHILANDER 77 m

of 3 m. Conflicts were observed principally dur- According to Layne (1951, p. 464), use of the tail ing the first 2 hr of the night when the four-eyed for transport by Didelphis was first described and to first opossums came take the meal (12 evenings illustrated by Pray (1921, p. 109). Layne further of observation). Usually an animal which had not elaborated on the subject. eaten chased one which was already eating on the platform. I observed direct fighting (bites) on only three occasions, but usually only threats were ex- Taming changed (hissing, open mouth, and start of chase). No rank order was observed and an individual Hershkovitz (1941, field notes; northern COLOM- (male or female) chased by another (male or fe- BIA) male) one night could chase it in turn the follow- The opossum captured alive after being struck ing night depending on the situation (degree of with a machete by a woodsman was leashed and hunger)." brought to me. It objected to being handled, was suspicious of anyone's approach, and reacted like all other didelphoids I've known by hissing with Spontaneous Bleeding the mouth open. Within a few days, however, the animal became tame and permitted itself to be Hershkovitz (1941, field notes; northern COLOM- handled and stroked and seemed to enjoy climb- BIA) ing onto my shoulder and head. adult female P. clubbed a I An opossum was by woodsman and seized as she jumped from its nest in the felled tree. The animal was brought live to Thermoregulation my camp, where it recovered without signs of in- Enders and Davis jury. When irritated, however, it bled spontane- (1936, p. 165; Canal Zone, ously from the tips of the fingers, the tail, and the PANAMA) nose. I had never witnessed or known of this phe- Body 35.4, environment 26.1 (afternoon). nomenon before.

Miller (in J. A. Allen, 1916c, p. 589; Mato Gros- so, BRASIL) Stress "As a general rule opossums cannot stand great heat and will soon die if left to the direct .Hunsaker and Shupe (1977, p. 302) exposed rays of the tropical sun." r "Tight coiling of the tail . . . occurs in Philan- der opossum, Didelphis marsupialis and Caluro- mys as a function of stress. It occurs when an Vocalization animal is frightened and can be considered a sub- mission posture." Charles-Dominique (1983, pp. 407, 411, 412; The coiled tail reduces from seizure exposure Cayenne, GUYANE FRAN^AISE) by a predator. "Two types of calls by adults are the weak sexual call (rarely by the male) and an agressive call by both sexes. Tail Transport "The young begin emitting very short high- pitched "clicks" about one month before releas- Hunsaker and Shupe (1977, p. 301) ing the nipples, especially when the mother licks "Tail coiling has been observed to facilitate the young attached to the nipples in her pouch, or carrying nesting material in Didelphis virginiana, when the young are experimentally pulled off the Monodelphis domestica, Caluromys derbianus nipples. and Marmosa robinsoni. The fact that more spe- "These clicks progressively disappear when the cies have not been observed to do so is probably young release the nipple and the first 'hiss' —as- a function of inadequate observations." Philander sociated with the typical posture (opened does coil its tail in the manner described and may mouth)—is emitted very early in conditions of indeed do so for transport of building material. fear or when pulled off the nipple."

HERSHKOVITZ: PHILANDER 79 In the case of C. philander the clicking sound Apayacu; Ucayali, Peru (43) was particularly loud, audible at 200 m. The same Aragar^as; Goias, Brasil (173) sound, but muted, was emitted by Marmosa mu- Arruda; El Beni, Bolivia (73) rina, Monodelphis hrevicaudata, and D. marsu- Arumateua; Para, Brasil (157) picdis. Ascencion de Guarayos; Santa Cruz, Bolivia (84) The clicking sound has also been recorded by Asuncion; Central, Paraguay (91) Reynolds (1952, p. 235) for D. virginiana as be- Auara; Igarape; Amazonas, Brasil (144) ing most pronounced during breeding. Kirsh Avanavero Falls; Nickerie, Suriname (119)

(1979, p. 392) heard the sound from Caenolestes Avanhandava (Lajeado); Sao Paulo, Brasil (195) fuliginosus; Thrasher et al. (1971), from Marmosa Ayacucho (Ibanez); Santa Cruz, Bolivia (88) robinsoni. Bruce Patterson (personal communi- Bacaetava; Sao Paulo, Brasil (199) cation) heard Metachirus nudicaudatus clicking Bagado; Choco, Colombia (4) as a signal of aggression. Bahia; Bahia, Brasil (172) Apparently all New World marsupials are ca- Baiao; Para, Brasil (156) pable of three types of calls: the standard hiss, Balisa; Mato Grosso, Brasil (168d) usually uttered when standing erect or tripodally Balta; Ucayali, Perii (55) in an aggressive posture, is usually fear stimulat- Barra do Cartas; Mato Grosso, Brasil (168c) ed; a mouselike squeak, which may be a sexual Barranquita; El Beni, Bolivia (73) call; and the click, which is emitted under various Barrieira; Rio de Janeiro, Brasil (189) conditions. Barro Branco; Rio de Janeiro, Brasil (193) Baptista (Lago); Amazonas, Brasil (141) Belem; Para, Brasil (159) Weight Belem-Brasflia (Rodovia); Para, Brasil (159) Belen; Territorio Federal Amazonas, Venezuela Tyndale-Biscoe (1980; P. opossum—COLOM- (106) BIA) Bellavista; Antioquia, Colombia (2) Bemberg (Puerto); Misiones, Argentina (95) of adults ^6 416.9 ± 132.1 Weights (g): 6, Benfica; Minas Gerais, Brasil (181) 49 323.8 ± 65.2 (230-675); $, (275-420). Better Hope; East Demerara, Guyana (not located but likely near Georgetown [114] and perhaps same as Hope [6°42'N, 57°57'W]) Boa Esperanga; Minas Gerais, Brasil (179) Alphabetical List of South American Boa Fe (Fazenda); Rio de Janeiro, Brasil (189) Collecting Localities of Boa Vista (Fazenda); Rio de Janeiro, Brasil (189) Philander with Key Numbers to Boa Vista; Roraima, Brasil (134) Distribution Map (Fig. 22) and Boca Colorado; Madre de Dios, Peril (65) Gazetteer (p. 85) Bom Jardim; Brasil; not located; many so-named throughout most states See Hall and Kelson (1959) for Middle Amer- Boraceia (Ponta); Sao Paulo, Brasil (203) ican portion of range. Boraceia, Rio Tiete; Sao Paulo, Brasil (196) Branco (Rio); Amapa, Brasil (133) Acanana; Territorio Federal Amazonas, Venezuela Brasilia; Distrito Federal, Brasil (175c) (107) Bucay; Guayas, Ecuador (28) Agua Limpa; Brasilia, Distrito Federal, Brasil Buck Hall; Essequibo Islands-West Demerara, (175c) Guyana (116) Agua Viva; Trujillo, Venezuela (98a) Buenaventura; Valle del Cauca, Colombia (10) Albina; Marowijne, Suriname (128) Buenavista; Santa Cruz, Bolivia (86) Alem Parafba; Minas Gerais, Brasil (182) Butanta; Sao Paulo, Brasil (200)

Alguacil; Zulia, Venezuela (97a) Caceres; Mato Grosso, Brasil (168a) ! Altamira; Para, Brasil (151) Cachavi (= Cachabi); Esmeraldas, Ecuador (18) Alto Bonito; Antioquia, Colombia (1) Cadena (Hacienda); Cuzco, Peru (63) Alto da Serra; Sao Paulo, Brasil (202) Caixa D'agua; Espirito Santo, Brasil (186) Ananindeua; Para, Brasil (159) Call; Valle del Cauca, Colombia (11) Anapolis; Goias, Brasil (174b) Cameta; Para, Brasil (153)

80 FIELDIANA: ZOOLOGY PHILANDER

L ocalit y Records

Fig. 22. South American collecting localities of Philander species. See Gazetteer (p. 85) for numbered locality names.

Camiaco; El Beni, Bolivia (81) Cardosa (Fazenda); Minas Gerais, Brasil (180)

Camino Vilches; EI Beni, Bolivia (73) Carondelet; Esmeraldas, Ecuador ( 1 8) Campinho; Espirito Santo, Brasil (185) Casa Grande; Sao Paulo, Brasil (203) Caney; Meta, Colombia (16) Catatumbo (Puerto); Zulia, Venezuela (97b) Cantareira (Serra da); Sao Paulo, Brasil (2(X)) Cauca (Rfo); Valle del Cauca, Colombia (11) Canudos; Pard, Brasil (149) Cayenne; Cayenne, Guyane Fran^aise (129) Caoni (Rio); Pichincha, Ecuador (22) Cebollal; Loja, Ecuador (30) Capibara; Territorio Federal Amazonas, Venezue- Cerro Cucurito; Territorio Federal Amazonas, la(lll) Venezuela (109b) Capim; Para, Brasil (164) Centinela; El Beni, Bolivia (73) ,Caracarai; Roraima, Brasil (135) Chaco; Argentina (94)

HERSHKOVITZ: PHILANDER Chalimana; Bolivar, Venezuela (104c) Esmeralda (Fazenda); Minas Gerais, Brasil (178) Chanchamayo; Junin, Peru (61) Esmeralda; Territorio Federal Amazonas, Vene- Chapare (Rio); Santa Cruz, Bolivia (83) zuela (109a) Chicosa; Ucayali, Peru (53) Esmeraldas; Esmeraldas, Ecuador (19) Chimbo (Puente); Guayas, Ecuador (28) Estrada de Santarem-Cuiaba; Para, Brasil (148) Chimbo-Coco (Rios); Guayas, Ecuador (28) Exaltacion; El Beni, Bolivia (74) Chocolatal, Monte, El Beni, Bolivia (75) Fazenda Boa Fe; Rio de Janeiro, Brasil (189) Chulumani; La Paz, Bolivia (67) Fazenda Cardosa; Minas Gerais, Brasil (180) Churuli; Zulia, Venezuela (97a) Fazenda da Floresta; Minas Gerais, Brasil (178) Clevia; Suriname, Suriname (125) Flor do Prado; Para, Brasil (161) Cocal; Cauca, Colombia (14) Floresta (Fazenda); Minas Gerais, Brasil (178) Coco (Rfo); Guayas, Ecuador (28) Fordlandia; Para, Brasil (147) Colonia do Prata; Para, Brasil (163) Formosa; Goias, Brasil (174a) Colonia Hansa; Santa Catarina, Brasil (209) Fracran; Misiones, Argentina (96) Colorado (Boca); Madre de Dios, Peru (65) Georgetown; Demerara-Mahaica, Guyana (114) Colorado (Rio); Madre de Dios, Peru (65) Gradaus; Para, Brasil (152) Commewijne (River); Commewijne, Suriname Gualea; Pichincha, Ecuador (23) (127) Guama; Para, Brasil (165) Comochatiba (= Comoxatiba) 17°06'S, 39°irw. Guaquitas; Barinas, Venezuela (100b) Maximilian Wied-Neuwied, July 1816. Guayaquil; Guayas, Ecuador (27) Conceigao do Mato Dentro; Minas Gerais, Brasil Guayaramarin; El Beni, Bolivia (68) (176) Giiiniquina, Territorio Federal Delta Amacuro, Condoto; Choco, Colombia (5) Venezuela (98d) Copataza (Rio); Pastaza, Ecuador (37) Hacienda Cadena; Cuzco, Peru (63) Corumba; Mato Grosso do Sul, Brasil (170) Hamacas; Santa Cruz, Bolivia (88) Costao dos Engenhos; Sao Paulo, Brasil (201) Hansa; Santa Catarina, Brasil (209) Cotia; Sao Paulo, Brasil (200) Hato San Jose; Bolivar, Venezuela (103) Covaria (Rio); Boyaca, Colombia (15) Hondo (Rio); Caldas, Colombia (3) Cucurito (Cerro); Territorio Federal Amazonas, Huampami; Amazonas, Perii (56b) Venezuela (109b) Huanhuachayo; Ayacucho, Peru (62) Culturutuin; Suriname, Suriname (125) Hyde Park; Demerara-Mahaica, Guyana (115) Culumani; La Paz, Bolivia (67) Ibaiiez; Santa Cruz, Bolivia (88) Cumaria; Ucayali, Peru (52) Ibare (Boca); El Beni, Bolivia (79) Cumeria; Ucayali, Peru (52) Ichilo (Rio); Cochabamba, Bolivia (82) = - Cumuruxotiba (= Comoxatiba Comochatiba Igarape Agu; Para, Brasil (163) q.v.) Igarape Tapereba; Para, Brasil (150) Curaray (Rio); Napo, Ecuador (39) Ilha do Taiuna; Para, Brasil (153) Curaray (Boca); Ucayali, Peru (39) Imperatriz; Amazonas, Brasil (140) De Oro (Rio); Chaco, Argentina (94) Inafiez; Pasco, Peru (not located) De Oro (Rio); Zulia, Venezuela (97) "High ground between Chanchamayo and up- Dividive; Trujillo, Venezuela (98a) per Rfo Pachitea" (Hendee in Thomas, Docampado (Rio); Choco, Colombia (7) 1928a, Annals and Magazine of Natural His- Duida (Cerro); Territorio Federal Amazonas, Ven- tory, Series 10, 2:250). ezuela (108) R. W. Hendee, June 1927, at 5000 ft El Capricho (Finca); 38 km E Villavicencio, Ipanema; Sao Paulo, Brasil (199) Meta, Colombia (16) Ipeau-Apez; Para, Brasil (159) El Carmen; El Beni, Bolivia (78) Ipitinga; Para, Brasil (160) El Chiral; El Oro, Ecuador (29) Ipixuna; Amazonas, Brasil (138)

' El Dividive; Trujillo, Venezuela (98a) Iporanga; Sao Paulo, Brasil (197) El Palmar; Bolivar, Venezuela (102a) Iquitos; Ucayali, Peru (42) El Palmar; Santa Cruz, Bolivia (88) Iriteria; Para, Brasil (165) Rio de Brasil Encontrados, Oro; Zulia, Venezuela (97b) Itaituba; Para, (148) j Reeve; Santo, Brasil Itatiaia Nacional Rio de Janeiro, Bra- Engenheiro Espirito (188) (Parque de); j Engenheiro Rive (see Engenheiro Reeve) sil (193)

82 FIELDIANA: ZOOLOGY Itatiaia (Serra de); Sao Paulo, Brasil (204) Mazagao; Amapd, Brasil (133) Itaya; Ucayali, Peru (42) Mazagao; Para. Brasil (154) Itonama; El Beni, Bolivia (71) Mazan (Rio); Ucayali, Peru (40) Joinville; Santa Catarina. Brasil (209) Mercedes; Santa Cruz, Bolivia (85) Juiz de Fora; Minas Gerais. Brasil (183) Merey; Territorio Federal Amazonas, Venezuela Juquia; Sao Paulo, Brasil (201) (112a) Kaiserberg Airstrip; Nickerie, Suriname (122) M^rida; M^rida, Venezuela (99) Kanuku Mountains; Upper Takutu-Upper Esse- Merredor; El Beni. Bolivia (69) quibo, Guyana (118) Mindo; Pichincha, Ecuador (24) Km 19; Para. Brasil (148) Miranda; Mato Grosso do Sul, Brasil (171a) Km 90; Pari Brasil (159) Miraiies (Lago); Ucayali, Peru (40) Km 216; Para, Brasil (148) Mocajuba; Para, Brasil (155) La Blanquita; Apure, Venezuela (101a) Montalvo; Pastaza, Ecuador (36)

La Boca, Rio Saija; Cauca, Colombia ( 1 3) Monte Alegre; Sao Paulo, Brasil (198) La Lengueta; Barinas, Venezuela (100c) Monte Alegre; Mato Grosso, Brasil (169) La Neblina; Territorio Federal Amazonas, Vene- Monte Chocolatal (see San Ram6n); El Beni, Bo- zuela (112b) livia (75) La Papaya; Manabi, Ecuador (not located) Monte Serrat; Rio de Janeiro, Brasil (193) T. Mena, May 1942, at 50 m Montes Claras; Minas Gerais, Brasil (178) La Ponchera; Tdchira, Venezuela (101b) Motatdn, 1 km E and 5 km NNE; Trujillo, Ven- La Poule; Saramacca, Suriname (124) ezuela (98a)

La Poza; Amazonas, Peru (56a) Motatan, Rio; Trujillo, Venezuela (98a) Lagarto Cocha; Ucayali, Peru (54) Moyobamba; San Martin, Peru (57) Lago do Baptista; Amazonas, Brasil (141) Murutucu; Pard, Brasil (159) Lago Mirafjes; Ucayali, Peru (40) Napo (Puerto); Napo, Ecuador (33) Lagoa Santa; Minas Gerais, Brasil (177) Napo (Rio); Napo, Ecuador (33) Lapango; Central, Paraguay (92) Nauta; Ucayali, Perii (45) Las Bonitas; Barinas, Venezuela (100b) Navio (Serra do); Amapi, Brasil (131) Lazaropolis do Prata; Para, Brasil (not located) Nickerie (River); Nickerie, Suriname (120) Lelydorp; Suriname, Suriname (126) Novita (Rio), Choc6, Colombia (8) Lelydorpplan; Suriname, Suriname (126) Novo Friburgo; Rio de Janeiro, Brasil (189) Lim6n; Ecuador (not located) Nulita; Apure, Venezuela (101a) G. H. H. Tate, December 1923, at 1200 ft 6rgaos (Serra dos); Rio de Janeiro, Brasil (189) Loksie Hattie; Brokopondo, Suriname (123) Oro (Rio de); Chaco, Argentina (94) Los Guires, Territorio Federal Delta Amacuro, Orosa; Ucayali, Peru (44) Venezuela (98c) Ouanary (River); Ouanary, Guyane Fran^aise Macapd; Amapd, Brasil (132) (130) Macieiras; Rio de Janeiro, Brasil (193) Oxapampa; Pasco, Peru (60) Madeira (Rio); Amazonas, Brasil (142) Pacific Virology Field Camp; Valle del Cauca, Magdalena; El Beni, Bolivia (76) Colombia (12) Majagua (Carlo); Territorio Federal Amazonas, Palacios; El Beni, Bolivia (74) Venezuela (105) Palmeiras; Mato Grosso, Brasil (169) Makerie; Nickerie, Suriname (121) Papelon (Cerro); Monagas, Venezuela (98b) Maldonado (Puerto); Madre de Dios, Peru (66) Para; Pari, Brasil (159) Mamor^ (Rio); El Beni, Bolivia (70) Pari (River); Suriname, Suriname (125) Mamore (Rio); Santa Cruz, Bolivia (83) Paracai (Rio); Parana, Brasil (207) Mangaratiba; Rio de Janeiro. Brasil (192) Paramaribo; Suriname. Suriname (125) Marabd; Pari, Brasil (158) Paramba; Imbabura, Ecuador (21) Marco; Para, Brasil (159) Parima (Sierra); Territorio Federal Amazonas, Maripa; Bolivar, Venezuela (102b) Venezuela (110b) Matipo (Rio); Minas Gerais, Brasil (178) Parintins; Amazonas, Brasil (139) Mato Grosso; Mato Grosso, Brasil (167) Parintins (Serra de); Amazonas, Brasil (139) Mavaca; Territorio Federal Amazonas, Venezuela Parque Nacional de Brasilia; Distrito Federal, (110c) Brasil (175b)

HERSHKOVITZ: PHILANDER 83 Parque Nacional de Itatiaia; Rio de Janeiro, Brasil San Joaquin; ElBeni, Bolivia (73) (193) San Jose; Ayacucho, Peru (62) Passo Fundo; Rio Grande do Sul, Brasil (210) San Jose; (Hato); Bolivar, Venezuela (103) Pedra Branco; Rio de Janeiro, Brasil (194) San Jose, Abajo; Napo, Ecuador (32) Peixe-Boi; Para, Brasil (163) San Juan; Territorio Federal Amazonas, Venezue- Pescado (Rio); Manabf, Ecuador (25) la (105) Pinas; El Oro, Ecuador (29) San Martin de Turumban; Bolfvar, Venezuela Pindo Yacu (Rio); Pastaza, Ecuador (38) (104a) Piquete; Sao Paulo, Brasil (205) San Pablo; El Beni, Bolivia (77) Playa del Rio Base; Territorio Federal Amazonas, San Ramon; El Beni, Bolivia (75) Venezuela (108) San Ramon; Santa Cruz, Bolivia (89) Boa Brasil Pocao, Vista; Roraima, (134) Sando (Rio); Choco, Colombia (6) Pocao, Caracarai; Roraima, Brasil (135) Santa Ana, Punta; Loja, Ecuador (29) Pocone; Mato Grosso, Brasil (168b) Santa Cecilia; Napo, Ecuador (31) Ponta de Boraceia; Sao Paulo, Brasil (203) Santa Clara; Amazonas, Brasil (140) Porto Velho; Rondonia, Brasil (166) Santa Cruz de la Sierra; Santa Cruz, Bolivia (88) Ucayali, Peru (51) Pucallpa; Santa Elena; Ucayali, Peru (46) Puente de Chimbo; Guayas, Ecuador (28) Santa Luisa; Ucayali, Peru (41) Puerto Asis; Colombia (17a) Putumayo, Santa Maria; Para, Brasil (162) Puerto Misiones, Argentina (95) Bemberg; Santa Rosa; El Beni, Bolivia (72) Puerto Caballo; El Beni, Bolivia (74) Santa Rosa; Ucayali, Peru (54) Puerto Catatumbo; Zulia, Venezuela (97b) Santa Rosita; Santa Cruz, Bolivia (87) Puerto Maldonado; Madre de Dios, Peru (66) Santa Teresa; Espirito Santo, Brasil (184) Puerto Napo; Napo, Ecuador (33) Santa Teresa; Mato Grosso do Sul, Brasil (170) Puerto Siles; El Beni, Bolivia (72) Santo Antonio de Uayara; Amazonas, Brasil Punta Santa Ana; Loja, Ecuador (29) (145a) Quartel de Sacramento; Minas Gerais, Brasil Santo Dios; El Beni, Bolivia (73) (178) Santo Isidoro; Amazonas, Brasil (137) Quincemil; Cuzco, Peru (64) Santarem; Para, Brasil (148) Quininde (Rio); Esmeraldas, Ecuador (20) Santarem-Cuiaba (Estrada); Para, Brasil (148) Raposo (Rio); Valle del Cauca, Colombia (12) Sao Joao de Petropolis; Espirito Santo, Brasil Restrepo; Meta, Colombia (16) (184) Rijweg; Suriname, Suriname (125) Sao Joao Marcos; Rio de Janeiro, Brasil (191) Rio de Janeiro; Rio de Janeiro, Brasil (190) Sao Luis de Caceres; Mato Grosso, Brasil (168a) Rio de Oro; Chaco, Argentina (94) Sao Miguel do Guama; Para, Brasil (165) Rio de Oro; Manabi, Ecuador (26) Sao Paulo; Sao Paulo, Brasil (200) Rio de Oro; Zulia, Venezuela (97b) Sao Sebastiao; Sao Paulo, Brasil (206) Rio Frio; Valle del Cauca, Colombia (9) (see Rio de Janeiro, Brasil Rio Hondo; Caldas, Colombia (3) Sapitiba Sepetiba); (191) Rioja; San Martin, Peru (58) Ro^a Nova; Parana, Brasil (208) Sapucai; Paraquari, Paraguay (93) Brasil Rodeio; Rio de Janeiro, Brasil (191) Sapucajuba; Para, (159) Rodovia Belem-Brasflia; Para, Brasil (159) Sapucay; Paraguari, Paraguay (93) Ecuador Rosarinho; Amazonas, Brasil (143) Sarayacu; Pastaza, (35) Peru Rupununi (River); Rupununi, Guyana (117) Sarayacu; Ucayali, (48) Ecuador Saija (Rio); Cauca, Colombia (13) Seboyal; Loja, (30) Salesopolis; Sao Paulo, Brasil (203) Sepetiba; Rio de Janeiro, Brasil (191) Salobra; Mato Grosso do Sul, Brasil (171b) Seringal Oriente; Acre, Brasil (146) Salvador; Bahia, Brasil (172) Serra; Espirito Santo, Brasil (185) Salvias; El Oro, Ecuador (29) Serra de Itatiaia; Sao Paulo, Brasil (204) San Ignacio de Moxos; El Beni, Bolivia (80) Serra de Parintins; Amazonas, Brasil (139) San Javier; Esmeraldas, Ecuador (18) Serra do Navio; Amapa, Brasil (131) San Jeronimo; Cuzco, Perii (63) Serra dos Orgaos; Rio de Janeiro, Brasil (189) San Jeronimo; Ucayali, Peru (49) Serrat Monte; Rio de Janeiro, Brasil (193)

84 FIELDIANA: ZOOLOGY "S. Francisco de C." Minas Gerais, Brasil (not Yurudn (Rio); Bolivar, Venezuela (104b). located) A. Robert, March 1900, at 1580 m. Shahuia; Ucayali, Peni (52) Siles (Puerto); El Beni, Bolivia (72) Gazetteer of South American Sitio Minas Gerais; Minas Gerais, Brasil (not lo- Collecting Localities of Philander cated) Supenam (River) (see Supinaam River); Demer- Colombia—Philander opossum fuscogriseus, P. ara-Mahaica, Guyana (113) opossum quica, P. andersoni andersoni Supinaam (River); Demerara-Mahaica, Guyana (1 13) San Tacuati; Pedro, Paraguay (90) Antioquia—Philander opossum fuscogriseus Taiuana (Ilha); Para, Brasil (153) Tamatama; Territorio Federal Amazonas, Vene- 1 . Alto Bonito, 7°0rN, 76°17'W, about 400 m, zuela (110a) upper Ri'o Sucio, W slope Cordillera Tapereba (Igarap^); Pard, Brasil (150) Occidental. Rio Grande do Sul, Brasil (211) Taquara; L. E. Miller and H. S. Boyle, February Tefe; Amazonas, Brasil (137) 1915. Teresinha; Brasil Amapd, (131) 2. Bellavista, 6°33'N, 75°18'W, upper Rio Rio de Brasil Teres6polis; Janeiro, (189) Porce, Cordillera Centra. Terezinha Teresinha); Brasil (see Amapa, (131) R Hershkovitz. February 1950, at 1200 m. Theresopolis (see Teresopolis); Rio de Janeiro, Brasil (189) Ticoporo, Reserva Forestal; Barinas, Venezuela Caldas—Philander opossum fuscogriseus (100a) Tingo Maria; Hu^nuco, Peni (59) 3. Rio Hondo, Samand, 5°42'N, 75°0rw, Tobesobe, Guayo; Territorio Federal Delta Ama- Cordillera Central. curo, Venezuela (98c) R Hershkovitz, March 1951. Tocomechi; Santa Cruz, Bolivia (88) Tres Troncos; Caquetd, Colombia (17b) Trinidade; Goids, Brasil (175a) Choc6—Philander Turumbdn (= San Martin de Turumban); Bolivar, opossum fuscogriseus Venezuela (104a) 4. 5°25'N, 76°24'W, Rfo Uragua-i (Rio); Misiones, Argentina (95) Bagad6, Andagueda. E. L. Kerr, November 1912, at 650 m. Urubamba (Rio); Ucayali, Peni (54) Rio Urucu, Rio; Amazonas, Brasil (145b) 5. Condoto, 5°06'N, 76°37'W, Condoto, of Rio San Juan. Urucum; Mato Grosso do Sul, Brasil (170) tributary H. G. F 1914, at 92 m. Utcuyacu; Juni'n, Peni (61) Spurrell, February 6. Rio Sand6, Rio Baud6, 5°03'N, 76°57'W. Utinga; Pard, Brasil (159) K. von October at 160 m. Vaca Diez; El Beni, Bolivia (79) Sneidem, 1958, 7. Rfo 4°45'N, 77°18'W. Valao de Sao Louren9o; Espirito Santo, Brasil (187) Docampad6, K. von Sneidem, 1958, at 75- Ventura; Guayas, Ecuador (28) September 160 m. Vila Oliveira; Sao Paulo, Brasil (202) 8. N6vita, 4°57'N, 76°34'W, Rfo Tamanl Vila Velha; Espfrito Santo, Brasil (185) L. E. Miller, December 1911, at 120 m. Villa Bella Imperatriz; Amazonas, Brasil (140) Villavicencio; Meta, Colombia (16) Vitoria; Espirito Santo, Brasil (185) Warnes; Santa Cruz, Bolivia (87) Valle del Cauca—Philander opossum fuscogriseus Xiriviny (Rio); Amazonas, Brasil (136) Yana Rumi (Rio); Napo, Ecuador (34) 9. Rfo Frfo, 4°09'N, 76°15'W, enters Rfo Cauca Yana Yacu; Ucayali, Peni (47) from east. Yarinacocha; Ucayali, Peni (50) A. A. Allen and L. E. Miller, November- Ypanema (= Ipanema); Sao Paulo, Brasil (199) December 1911, at 100 m. Yurac Yacu; San Martin, Peni (56c) 10. Buenaventura, 3°53'N, 77°04'W. Yurimaguas; Ucayali, Peni (47) K. von Sneidem, February 1958, at near

HERSHKOVITZ: PHILANDER 85 sea level. Ecuador—Philander opossum melanurus, P. C. H. Tyndale-Biscoe, September 1971. opossum fuscogriseus, P. andersoni andersoni 10. Buenaventura, 28 km NE. J. A. W. Kirsch, September 1969, at 150 m. Esmeraldas—Philander opossum melanurus 11. Call, 3°27'N, 76°3rw, upper Rio Cauca valley. 18. Cachavi (Cachabi), Rio Cachavi, r03'N, C. H. Tyndale-Biscoe, September 1971. 78°50'W.

1 1. "Rio Cauca" (see Cali). 18. Carondelet, 1°03'N, 78°50"W. J. H. Batty, June 1898. G. Fleming, October 1900. 12. Raposo (Rio), 3°43'N, 77°08'W, Pacific 18. San Javier, 1°04'N, 78°47'W. Virology Field Camp. G. Fleming, August 1900, at 20 m. J. Duran, June 1962, at near sea level. 19. Esmeraldas, 0°59'N, 79°42'W. W. Richardson, November 1912, at near sea level. Cauca—Philander opossum fuscogriseus 20. Rio Quininde, 0°20'N, 79°28'W.

13. Rio Saija, La Boca (= Mouth), 2°52'N, 77°4rW. Imbabura—Philander opossum melanurus K. von Sneidern, June 1958, at near sea level. 21. Paramba, 0°49'N, 78°2rw, 1100 m. 14. Cocal, 2°3rN, 77°00'W, upper Rio San W. F H. L. Juan. Rosenberg, April 1897; Gomez, June 1941. L. E. Miller, July 191 1, at 125 and 187 m.

Boyaca—Philander opossum quica Pichincha—Philander opossum melanurus

15. Rio Covaria, 7°03'N, 72°04'W, near mouth, 22. Rio Caoni, 0°12'N, 79°23'W. above town of Covaria. M. Olalla, January 1935. K. von Sneidern, March 1959, at 350 m. 23. Gualea, 0°08'N, 78°48'W, ca. 1250 m. L. Soderstrom, May 1920. 24. Mindo, below 8°02'S, 78°48'W. Meta—Philander opossum quica L. Soderstrom, December 1924, at 1000 m.

16. Caney, Rio Guatiquia (see Restrepo). 16. Restrepo, 4°15'N, 73°33'W. Manabi—Philander 16. Villavicencio, 4°09'N, 73°37'W, upper Rio opossum fuscogriseus Guatiquia. 25. Rio 80°15'W. R. Gilmore, February, May, June 1939, at Pescado, 1°25'S, 465 m. G. H. H. Tate, May 1922, at 500 m. 26. Rio de 79°22'W. C. H. Tyndale-Biscoe, 1971. Oro, 2°10'S, W. 1917. 16. El Capricho (Finca), 38 km E Villavicencio, Richardson, January 4°09'N, 73°16'W. J. A. W. Kirsch, October 1969. Guayas—Philander opossum fuscogriseus

—Philander andersoni andersoni Putumayo 27. Guayaquil, 2°10'S, 79°50'W, near sea level. E. Belcher, 1838. 17a. Puerto Asis, 17 km N; 0°3rN, 76°3rw. 28. Puente de Chimbo, 2°10'S, 79°07'W. J. A. W. Kirsch, November 1969, at 330 m. G. H. H. Tate, August 1922, at 375 m. 28. Rios Chimbo-Coco, 2°10'S, 79°50'W. Caqueta—Philander andersoni andersoni G. H. H. Tate, July 1922, at 750 m. 28. Bucay, Rio Chimbo, 2°10'S, 79°06'W. 17b. Tres Troncos, 0°08'N, 74°4rw, above La G. H. H. Tate, November 1921, at 312 m. Tagua, Rio Caqueta. 28. Ventura, Rio Chanchan, 2°17'S, 79°24'W. P Hershkovitz, January 1952, at 182 m. G. H. H. Tate, April 1922, at 750 m.

86 FIELDIANA: ZOOLOGY El Oro—Philander opossum fuscogriseus 39. Ri'o Curaray. Boca (= mouth), 2°22'S, 74°05'W.

29. El Chiral, 2''39'S, 79°43'W. Olalla Brothers, 1925. H. E. Anthony, August 1920, at 167 m. 40. Lago Miranes, Rio Napo (see Rio Mazdn). 29. Piiias, 3°42'S, 79°42'W. 40. Rfo Mazan (mouth, 3°28'S, 73°02'W.

I G. H. H. Tate, September 1921, at 1 125 m. L. Soderstrom, September 1930. 29. Salvias, 3°47'S, 79°2rw. 41. Santa Luisa, Rio Nanay, 3°35'S, 74°30'W. H. E. Anthony, August 1920, at 1000 m. C. Kalinowski, October 1956. 42. Itaya (see Iquitos). H. Bassler, 1927. 42. Rio Loja—Philander opossum fuscogriseus Iquitos, Maraiion, 3°46'S, 73°15'W, 106 m. H. Bassler, March 1928; C. Kalinowski, 29. Santa Ana, Punta (?) 3°50'S, 79°25'W, on October 1956; R. W. Hendee, 1928; road from Zaruma to Loja. January C. Arevalo (H. Bassler collection), March H. E. Anthony, December 1920, at 1 14 m. 1930. 30. Seboyal (see Cebollal). 43. 72°07'W. 30. Cebollal, 4°02'S, 80°02'W. Apayacu, 3°19'S, Olalla 1927. G. H. H. Tate, October 1921, at 968 m. Brothers, January 44. Orosa, 3°26'S, 72°08'W. OUalla Brothers, October 1926. 45. Nauta, 4°30'S, 73°25'W, ca. 130 m. Nsipo^Philander andersoni andersoni 46. Santa Elena, Rio Samiria, 4°42'S, 74°12'W, ca. 130 m. 31. Santa Cecilia, 0°03'N, 76°58'W. C. Kalinowski, November 1956. 32. San Jos^, abajo (= below), 0°3rS, 77°20'W. 47. Yurimaguas, Rfo Huallaza, 5°54'S, 76°05'W, Olalla Brothers, 1924. April 180 m. 33. Puerto Naop, Rio Napo, 1°03'S, 77°47'W. 47. Yana Yacu, below Yurimaguas, 5°52'S, 33. Rio (near) (see Puerto Napo Napo). 76°15"W. L. Soderstrom, March 1921. M. R Anderson, September 1912, at 180 m. 34. Rfo Yana Rumi, 1°38'S, 76°59'W. 48. Sarayacu, Rfo Ucayali, 6°44'S, 75°06'W. R. Olalla, October 1934. A. M. Olalla, March, April 1927. 49. San Jer6nimo, 7°45'S, 74°50'W, 300 m. R. W. Hendee, December 1927. Pastaza—Philander andersoni andersoni 50. Yarinacocha, 8°15'S, 74°43'W, 160 m. C. C. Sanborn, March 1946; A. L. 35. Rio Bobonaza, 1°44'S, 77°29'W. Sarayacu, Gardner, August 1968, March 1971. G. H. H. Tate, March 1924, between 450 51. Pucallpa, 59 km W, 59 km NE, 59 km SW and 500 m. (see Pucallpa. Rfo Ucayali). 36. Rio 76°58'W. Montalvo, Bobonaza, 2°04'S, H. Hinse, October, November 1971, R. Olalla, February 1932. October 1972; H. Hinse, 59 km NE, 37. Rio 77°27'W. Copataza, 2°07'S, September 1972; 59 km SW, October 1972. R. Olalla, March 1939; G. H. H. Tate, 51. Pucallpa, Rfo Ucayali, 8°23'S, 74°32'W, 180 March 1924. m. 38. Rio Pindo 76°03'W. Yacu, 2°08'S, 52. Cumeria (= Cumarfa, across from Shahufa, R. Olalla, October 1934, at 250 m. q.v.). 52. Shahufa, Rfo Ucayali opposite Cumarfa, 9°52'S, 74°0rW. Peru—Philander P. opossum quica, andersoni R. W. Hendee, July 1927, at 312 m. andersoni, P. andersoni mcilhennyi 53. Chicosa, 10°2rS, 74°00'W, ca. 150 m. R. W. Hendee, September 1927. 54. Lagarto Cocha, Rfo Ucayali, 10°4rs, Ucayali—Philander opossum quica (42, 43, 48- 73°48'W. 51, 54, 55), P. andersoni andersoni (39-41, 44- Olalla Brothers, January 1928.

P. 1 47), andersoni mcilhennyi (5 , 55). 54. Rfo Urubamba, mouth, 10°42'S, 73°42'W.

HERSHKOVrrZ: PHILANDER 87 Olaila Brothers, October, November Ayacucho—Philander andersoni andersoni 1927. 54. Santa Rosa, 10°43'S, 73°53'W. 62. San Jose, Rio Santa Rosa, 12°44'S, 73°46'W. Olaila Brothers, December 1927. A. L. Gardner, April 1971, at 1000 m. 55. Balta, Rio Curanja, 10°08'S, 71°15'W, 300 62. Huanhuachayo, 12°44'S, 73°47'W. m. A. L. Gardner, May 1971, at 1660 m. A. L. Gardner, June 1968, March 1971; R. Thomas, 1971; A. L. Gardner, July, August 1966, February 1977, July 1968; Cuzco—Philander opossum quica J. R O'Neil, February 1971. 63. Hacienda Cadena, Marcapata, 13°24'S, 70°43'W, 890 m. Amazonas—Philander andersoni andersoni C. Kalinowski, October 1949, August 1950. 63. 7r54'W. 56a. La Poza, Rio Santiago, 4°0rS, 1T41'W. SanJeronimo, 13°34'S, 64. 70°38'W. J. L. Patton, August 1979, at 180 m. Quincemil, 13°16'S, C. June at 680 m. 56b. Huampami, Rio Cenepa. Kalinowski, 1953, J. L. Patton, July, August 1977, July 1978.

Madre de Dios—Philander opossum quica San Martin—Philander opossum quica 65. Colorado, Rio (mouth) (see Boca Colorado). 65. Boca Colorado, 12°30'S, 70°10'W. 56c. Yurac Yacu, 5°52'S, 77°14'W. C. Kalinowski, October 1954, at 279 m. R. W. Hendee, July 1926, at 780 m. 66. Puerto Maldonado, Rio Tambo, 12°36'S, 57. Moyobamba, Rio Mayo, 6°03'S, 76°58'W. 69°irW, 256 m. W. H. Osgood and M. P. Anderson, M. L. Kuns, 1965. August 1912; April L. Rutter, February 1924, June 1926, at 845 m. Bolivia—Philander opossum quica 58. Rioja, 6°10'S, 77°10'W, ca. 800 m. R. W. Hendee, September 1926. La Paz

67. Culumani (see Chulumani). Huanuco—Philander opossum quica 67. Chulumani, 16°24'S, 67°3rW, 2000 m. P O. Simons, February 1901, at 2200 m. 59. Tingo Maria, 9°08'S, 75°57'W. R. W. Hendee, January 1927, at 625 m.

El Beni

Pasco—Philander andersoni andersoni 68. Guayaramarin, Ri'o Mamore, 10°5rS, 65°23'W. 60. 75°24'W. Oxapampa, 10°34'S, A. Ximenez, June 1964; K. F Koopman, C. R. Perez, August 1964, at 280 m. 1964. 69. Merredor, Rio Mamore, ll°4rs, 65°05'W. D. Edwards, July 1965; A. Ximenez, May Junin—Philander andersoni andersoni 1965.

70. Rio Mamore, 12°26'S, ca. 65°W. f

61. 1 Chanchamayo, 1°03'S, 75°19'W. D. E. Afiez, October 1965. « P O. Simons, August 1900; J. M. 71. Itonama, 12°28'S, 64°24'W. Schunke, August 1948; M. Kalinowski M. L. Kuns, April 1970. before 1916. 72. Puerto Siles, Rfo Mamore, 12°49'S, 61. Utcuyacu, 1 n2'S, 75°28'W. 65°00'W. P O. Simons, August 1900, at 1600 m. S. Anderson, October 1965.

FIELDIANA: ZOOLOGY 72. Santa Rosa, 13°01'S, 65°1 1'W. 84. Ascencion de Guarayos, 14°57'S, 61°24'W. M. L. Kuns, June 1964. M. L. Kuns, June 1964. 73. Barranquita (see San Joaquin). 85. Mercedes, Rio Guapore, 15°36'S, 60°22'W, M. L. Kuns, August 1963. 6 km S and opposite Buena Hora, Brasil. 73. Arruda, 4 km NE San Joaquin (q.v.). A. Ximenez, May 1965. M. L. Kuns, December 1963. 86. Buenavista, 17°27'S, 63°40'W.

73. Centinela, 1.5 km E San Joaquin (q.v.). T. Bridges, probably 1846; F. Steinbach, M. L. Kuns, June 1964. January 1915; J. Steinbach, June 1921, May 73. Camino Vilches, 1 mi E San Joaquin (q.v.). 1925, July 1926, June, July 1927, July 1928. M. L. Kuns, February 1966. 87. Wames, 17°3rs, 63°10'W. 73. Santo Dios, 2 km NE San Joaquin (q.v.). M. L. Kuns, August 1965. M. L. Kuns. June 1963. 87. Santa Rosita, Wames, 17°30'S, 63°10'W. 73. San Joaquin. Ri'o Machupo, 13°04'S, M. L. Kuns, July 1965. 64°49'W. 88. Tocomechi, 17°35'S, 62°55'W. M. L. Kuns, April, June, July 1963; M. L. Kuns, August 1965. February, May 1964. 88. Hamacas, 17°44'S, 63°1 1'W. 74. Exaltacion, Rio Mamord, 13°16'S, 65°15'W. O. Silva (Rockefeller Institution), July M. L. Kuns, May 1964; S. Anderson, 1938. October 1965, at 8 km N. 88. El Palmar, 17°48'S, 63°10'W. 74. Palacios, 13°34'S, 65°19'W. W. Kerr, August 1966, at 500 m; D. R. D. E. Anez. May 1965. Hadden, August 1966. 74. Puerto Caballo, 13°34'S, 65°2rw. 88. Ayacucho, Ibdfiez, 17°5rS, 63°20'W. S. Anderson, September 1965. J. Riddell, August 1966. 75. San Ramon, Ri'o Machupo, 13°18'S, 88. 18 km SW Santa Cruz (q.v.). 64°37'W. F. Becerra, August 1966. M. L. Kuns, January 1964. 88. Santa Cruz (see Santa Cruz de la Sierra). 75. Monte Chocolatal (see San Ram6n). 88. Santa Cruz de la Sierra, 17°48'S, 63°10'W. M. L. Kuns, January 1964. F Becerra, September 1966. 76. Magdalena, 13°20'S, 64°08'W. 89. San Ram6n, 17°33'S, 61°03'S. M. L. Kuns, July 1963, July 1964. 77. San Pablo, 13°52'S, 65°36'W. —Philander M. L. Kuns, September 1963. Paraguay opossum quica 78. El Carmen, Rio Blanco, 13°57'S, 63°43'W. San Pedro M. L. Kuns, June 1964. 79. Rio lhar6 (mouth), 14°37'S, 64°57'W. 90. Tacuati, 23°27'S, 56°35'W. D. E. Anez, 1965. August C. Wharton, May 1950. K. F. Koopman, August 1965. 79. Vaca Diez, 14°47'S, 64°5rw. Central M. L. Kuns, May 1969. 80. San Ignacio de M6xos, 14°53'S, 65°36'W. 91. Asunci6n, 25°16'S, 57°40'W. M. L. Kuns, June 1965. 92. Lapango, 25°2rS, 57°42'W. 81. Camiaco, 15°24'S, 64°46'W. H. Krieg, August 1925. A. Xim6nez, August 1965. Paraguarf Cochabamba 93. Sapucay (= Sapucaf), 25°40'S, 56°55'W. W. Foster, 1902. 82. Rfo Ichilo, 16°50'S, 64°45'W. September D. E. Aiiez, July 1965.

Argentina—Philander opossum Santa Cruz Chaco 83. 2 km S mouth, Ri'o Chapar6, Rio Mamor6, 15°58'S, 64°42'W. 94. No precise locality. D. E. Aiiez, July 1965. C. Friend, before 1880.

HERSHKOVrrZ: PHILANDER 89 94. Rio de Oro (mouth), 27°04'S, 58°34'W. Merida—Philander opossum subspecies I. Apostal, October 1962. 99. "Merida," 8°36'N, 71°08'W. Misiones Barinas—Philander opossum subspecies 95. Rio Uruguar-i, 30 km from Puerto Bemberg 100a. Reserva (Libertad). 26°30'S, 54°W. Ticoporo, Forestal, 07°48'N, 69°55'W. J. A. Crespo, September, October, November 1949. 100b. Guaquitas, 07°28'N, 71°39'W. 96. Fracran, San Pedro, 26°46'S, 54°16'W. 100b. Las Bonitas, Cano Amaru, 07°23'N, 70°44'W. J. A. Crespo, February 1952. 100c. La Lengueta, 08°30'N, 70°23'W.

Venezuela—Philander opossum subspecies, P. —Philander andersoni andersoni Apure opossum subspecies

101a. Nulita, 07°19'N, 71°55'W. Zulia—Philander opossum subspecies A. Tuttle, January 1968, at 24 m. 101a. La Blanquita, 7°12'N, 71°45'W. 97a. Alguacil, Caja Seca, 09°8'N, 71°04'W. A. Tuttle, January 1968, at 24 m. 97a. Churulf, Caja Seca, 09°09'N, 71°04'W. 97b. Puerto Catatumbo, 09°07'N, 72°35'W, 50 m. Tachira—Philander opossum subspecies 97b. Boca del Rio de Oro, 9°06'N, 72°45'W. 97b. Encontrados, 9°03'S, 72°14'W. 101b. La Ponchera, 07°26'N, 71°52'S. N. E. Peterson, March 1968. 97b. Encontrados, 60 km WNW, 90°03'N, 72°14'W, 73 m. Bolfvar—Philander opossum opossum

102a. El Palmar, Rio Grande, 08°0rN, 61°55'W. Trujillo—Philander opossum subspecies 102b. Maripa, Rio Caura, 7°26'N, 65°09'W. S. M. Klages, 1901. 98a. Motatan, 9°24'N, 70°36'W, 1 km E at 330 103. Hato San Jose, 146 km S, 7 km NE Ciudad m, 5 km NNE at 290 m. Bolivar, 6°44'N, 63°27'W. J. A. W. Kirsch, June 1969. N. Peterson, D. Peacock, R. Peacock, D. 98a. Motatan 9°28'N, 70°34'W at 290 m, (Rio), Furman, March 1967, at 302 m. 9.8 km NNE at 290 m. 104a. San Martin de Turumban, Rio Cuyuni, J. A. W. June 1969. Kirsch, 06°59'N, 6r02'W. 98a. 70°36'W. Agua Viva, 9°34'N, 104b. Rio Yuruan, 06°48'N, 61°50'W. N. E. Peterson, September 1965, at 164 m. M. A. Carriker, Jr., March 1910. 98a. El Dividive, 9°29'N, 70°44'W. 104c. Chalimana (Raudal), Rio Paramichi, Rio N. E. Peterson, October 1965, at 90 m. Paragua, 04°10'N, 62°59'W.

—Philander Monagas [?] opossum subspecies Amazonas (Territorio Federal)—Philander andersoni andersoni 98b. Papelon (Cerro), 10°0rN, 63°54'W. 105. Majagua (Cafio), Rio Ventuari, 05°20'N, Delta Amacuro (Territorio Federal)—Philander 65°40'W. opossum subspecies 105. San Juan, Ri'o Manapiare, 05°19'N, 66°03'W. 98c. Los Guires, 09°15'N, 61°54'W. M. D. Tuttle, E L. Harder, July 1967, at 98c. Tobesobe, Guayo, 09°00'N, 61°25'W. 155 m. 98d. Guiniquina, 90°10'N, 61°03'W, m. 106. Belen, Rio Cunucunuma, 03°43'N, 65°42'W.

90 FIELDIANA: ZOOLOGY r M. D. Tuttle, E L. Harder, January 1967, 1 18. Kanuku Mountains, 3°N, 59°45'W. at 150 m. E. V. McConnell, J. J. Quelch, November 107. Acanana, Ri'o Cunucunuma, 03°39'N, 1900, at 240 ft. 65°66'W.

M. D. Tuttle, F. L. Harder, June 1967, at 145 m. Suriname—Philander opossum opossum 108. Playa del Rio Base, Mt. Duida, 03°25'N. 65°40'W. Nickerie Olalla Brothers, November 1928. 109a. Esmeralda, 03°1 1'N, 65°33'W. 119. Avanavero Falls, Kabulebo River, 4°49'N, M. D. Tuttle, E L. Harder, March 1967, at 57°24'W. 135 m. 120. Nickerie River, upper, ca. 5°59'N, 56°30'W. 109b. Cucurito, Cerro, 03°38'N, 66°25'W. 121. Makerie, West River, Wilhelmina Mountains. 3°26'N. 56°45'W. 1 10a. Tamatama, Rio Orinoco, 3°08'N. 65°52'W. H. A. December 1961, M. D. Tuttle. E L. Harder, April, May, June Beatty. January 1962. 1967, at 135 m; D. S. Bremington, June 1967, at 135 m. 122. Kaiserberg Airstrip, 3°10'N, 56°15'W. H. A. 1961, at 275 m. 1 10b. Parima (Sierra), 02°40'N, 64°30'W. Beatty. February

1 10c. Mavaca, 02°3rN, 65°10'W. 111. Capibara, Brazo Casiquiare, 2°34'N, 66°18'W. Brokopondo M. D. Tuttle, E L. Harder, June 1967, at 123. Loksie 55°28'W 130 m. Hattie, 5°09'N. P. Hershkovitz, December 1961. 1 12a. Merey, opposite, Brazo Casiquiare, 2°17'N, 67°irw. Olalla Brothers, October 1929. Saramacca 112b. La Neblina (Campamento), Cerro La Neblina, 00°52'N, 66°14'W. 124. La Poule, 5°47'N, 55°25'W. P. Hershkovitz, January, February 1962. Guyana—Philander opossum opossum

Demerara-Mahaica Suriname

113. Supinaam River (= Supenam River), 125. Paramaribo, 5°50'N, 55°1 1'W. 6°58'N, 58°3rw. I. T. Sanderson, February 1938.

Crozier. 125. Clevia. Paramaribo (q.v.). 114. Georgetown, Demerara River, 6°48'N, P. Hershkovitz, February 1962. 58°10'W. 125. "Culturutuin," Agricultural Experimental J. Rodway. June 1929. Station in Paramaribo (q.v.).

115. Hyde Park, Demerara River, 6"'30'N, 125. Para River, ca. 10 km SE Paramaribo (q.v.). 58°16'W. 125. Rijweg, ca. 9 km W Paramaribo (q.v.). S. B. Warren, September 1906. 126. Lelydorp, 5°42'N, 55°16'W. 126. Lelydorpplan (see Lelydorp). P. Hershkovitz, January, February 1962. Essequibo Islands-West Demerara

116. Buck Hall, Essequibo River, 6°56'N, 58°33'W. Commewijne S. B. Warren, March 1906. 127. Commewijne River, 5°54'N, 55°05'W.

Upper Takutu-Upper Essequibo Marowijne 117. Rupununi River, 4°03'N, 58°34'W. J. J. Quelch, September 1900, at 200 ft. 128. Albina, 5°30'N, 54°03'W.

HERSHKOVITZ: PHILANDER 91 Guyane Frangaise—Philander opossum opossum 139. Serra de Parintins, 2°35'S, 56°25'W. Olalla Brothers, November 1930. Cayenne 139. Parintins, 2°36'S, 56°44'W. Olalla Brothers, November 1930. 129. Cayenne, 4°56'N, 52°19'W. 140. Santa Clara, Villa Bella Imperatriz, 2°50'S, G. K. Cherrie, B. T. Gault, November 56°55'W. 1902; S. Klages, January, February 1917; Olalla Brothers, August 1930. M. Atramentowicz, September 1978- 141. Lago do Baptista, 3°18'S, 58°15'W. October 1982; P. Charles-Dominique, 1978- A. M. Olalla, June 1936. 1982. 142. Rio Madeira, mouth, 3°22'S, 58°45'W. Olalla Brothers, February 1930. 143. Rosarhino, 3°43'S, 59°08'W. Ouanary Olalla Brothers, June, July 1930. 144. Auara Igarape, 4°22'S, 59°43'W. 130. Ouanary River, 4°14'N, 51°39'W. Olalla Brothers, March 1930. 145a. Santo Antonio de Uayara, Rio Eini, 6°43'S, 69°52'W. Brasil—Philander opossum opossum, P. opossum Olalla Brothers, April 1930. frenata, P. opossum quica 145b. Rio Urucu, 4°51'S, 65°16'W. M. N. E da Silva. Amapa—Philander opossum opossum

131. Serra do Navio, 0°59'N, 52°03'W. Acre—Philander opossum quica T. P. Woodall, June 1966; Instituto Evandro Chagas, 1967, 1968, 1969. 146. Seringal Oriente, 8°48'S, 72°46'W. 131. Teresinha (= Terezinha), Rio Amapari, M. Moreira, August 1934; M. Moreira, E 0°58'N, 52°02'W. Novaes, August 1956. 132. Macapa, Rio Amapari, 0°02'N, 51°03'W. M. Moreira, October, November 1952. 133. Mazagao, Rio Maraca, 0°06'S, 5ri8'W. Para—Philander opossum opossum M. Moreira, December 1958. 133. Rio Branco, tributary Rio Maraca, 0°07'S, 147. Fordlandia, Rio Tapajoz, 3°40'S, 55°30'W. 5ri7'W. R. M. Gilmore, February 1938. 148. Km 19, Itaituba-Jacareacanga, 4°17'S, 56°05'W. Roraima—Philander opossum opossum E Ramos, Instituto Oswaldo Cruz, August 1972. 134. Boa Vista, Rio Branco, 2°49'N, 60°40'W. 148. Km 216, Estrada de Santarem-Cuiaba C. T. Carvalho, M. Sobeiro do Amaral, (BR 165) (see Itaituba). M. Melo, March 1959. Instituto Evandro Chagas, May, June 134. Pocao, Boa Vista (see Boa Vista). 1973. 135. Caracarai, Rio Mucajai, r50'N, 61°08'W. 149. Canudos, 7°16'S, 58°07'W. C. T. Carvalho, M. Sobeiro do Amaral, E Lima, November 1920. M. Melo, March 1959. 150. Tapereba, Igarape, Chaves, 0°10'S, 49°55'W. 135. Pocao, Caracarai (see CaracaraO. C. Carvalho, June, July 1958. 151. Altamira, Rio Xingu, 3°12'S, 52°12'W. Instituto Oswaldo Cruz, August 1971. Amazonas—Philander opossum opossum 152. Gradaus, 7°43'S, 51°11'W. C. T. Carvalho, N. Hidasi, M. Amaral, 136. Rio Xiriviny, 0°59'S, 61°53'W. June, July, August 1957. K. B. Parker, October 1928, at 80 m. 153. Cameta, Rfo Tocantins, 2°15'S, 49°29'W. 137. Santo Isidoro, Tefe, 3°27'S, 64°47'W. A. M. Olalla, February 1934, March, Olalla Brothers, August 1928. April, May, November 1935, November 138. Ipixuna (Lago do), 3°52'S, 63°52'W. 1936.

92 FIELDIANA: ZOOLOGY ° ' 1 — 153. Ilha do Taiuna, Rfo Tocantins, 2 5 S . Mato Grosso Philander opossum quica 49°29'W, opposite Cameta. A. 1931. M. Olalla, October, November \bl. Mato Grosso. 15°00'S. 59°57'W. 154. Rio Tocantins. 2°25'S, 49°10'W. Mazagao, J. Natterer. September-November 1824. F. November 1912. Lima, 168a. Cdceres. 16°04'S, 57°41'W. 155. Rio Tocantins. 2°35'S. 49°30'W. Mocajuba, M. L. Kuns, September 1965. A. M. November 1931. Olalla. 168b. PcKone. 16°15'S. 56°37'W. 156. Baiao, Rio Tocantins. 2°41'S. 49°41'W. 168c. Ban-a do Gardes. 15°53'S. 52°15'W. A. M. Olalla. December 1931. 168d. Balisa. 16°15'S. 52°26'W. 157. Arumateua. Rfo Tocantins. 3°54'S. 49°41 'W. 169. Monte Alegre. Palmeiras. 16°03'S. 55°30'W. F. Lima. October 1912. A. M. Olalla. June 1944; A. Aggio, June 158. Rio 49°10'W. Maraba. Tocantins. 5°20'S, 1944. N. Peterson. September 1975. 159. Para (see Bel^m). 159. Km 90. Para. Rodovia Mato Grosso do Sul—Philander E. Snethlage. April 1909. opossum quica 159. Belem do Pard (see Belem). 159. Belem. 1°27'S. 48°29'W. 170. Commbd. Rio Paraguay, 19°0rS, 57°39'W. M. A. Miles. 1975-1980. M. L. Kuns, September 1965. 159. Ipeau-Apez. Belem (see Belem). 170. Urucum, 19°13'S, 57°33'W. E. Snethlage. L. E. Miller. December 1913. at 125 m. 159. Marco, suburb of Belem (q.v.). 170. Santa Teresa (see Urucum). 159. Murutucu (= Institute Agronomico do M. L. Kuns. September 1965. Norte). Belem (q.v.). 171a. Miranda. 20°14'S. 56°22'W. E Luna. March 1925. August 1922, A. M. Olalla. 1957; J. Lima. September 159. Utinga. suburb of Belem (q.v.). 1937. Institute R. H. Oswaldo Cruz, June 1963; 171b. Salobra. 21°14'S. 57°08'W. June 1968. Pine. L. Travassos, May 1942. 159. Ananinddua. E. E Braganqa. 1°22'S, 48°23'W. E Lima, May 1920. Bahia—Philander 159. Sapucajuba. suburb of Belem (q.v.). opossum frenata 160. Ipitinga. Rio Acara. 1°57'S. 48°1 1'W. 161. Elor do Prado, near Quatipuru, 0°52'S. 172. Bahia (= Salvador). 12°59'S. 38°3rw. 46°59'W. Herr Kaehne, before 1815. E. Snethlage. October 1916. 162. Santa Maria. Braganqa. 1°03'S. 46°46'W. April 1968. Golds—Philander opossum quica 163. Peixe-Boi. R. R. Bragan<;a. n2'S,47°18'W. 163. Igarape Aqu. 1°32'S. 47°03'W. 173. 15°55'S, 52°15'W. 163. Colonia do Prata, Igarape Aqu (q.v.). Aragarqas, 164. Capim. 1°30'S. 48°20'W. M. Amaral. May 1958. Institute Oswaldo Cruz, 1960. 174a. Formosa. 15°32'S. 47°20'W. 165. Sao Miguel do Guamd (see Guamd). 174b. Anapolis. I6°20'S. 48°58'W. 165. Guamd. 1°37'S. 47°27'W. R. M. Gilmore. 1936-1937. Departamento do Zoologia, Sao Paulo. 175a. Trinidade. I6°40'S. 49°30'W. October 1959. S. Hidasi. June 1962. 165. Iriteria, Sao Miguel do Guamd (see Guamd). M. Amaral. December 1959, January 1960. Distrito Federal—Philander opossum quica

Rondonia—Philander opossum quica 175b. Parque Nacional. 15°35'S, 48°54'W. 166. Porto Velho. 8°46'S. 63°54'W. 175c. Brasilia, 15°47'S, 47°55'W. T Hibbs. April 1965. 175c. Agua Limpa (Fazenda). 15°57'S, 47°54'W.

HERSHKOVITZ: PHILANDER 93 Minas Gerais—Philander opossum quica G. R. Hancock, September 1923, at 3000 ft; R. Kellogg, March 1943; G. Perreira, 176. do Mato 19°0rS, Concei^ao Dentro, September 1942; D. E. Davis, May, 43°25'W. September, October 1943; C. Guinle, 177. Santa, 19°38'S, 43°53'W. Lagoa Perreira, January 1943; C. Guinle, P. M. P. W. Lund, between 1833 and 1880. Britto, November 1942, March 1943, May 178. de 42°3rw. Quartel Sacramento, 19°44'S, 1943; C. Guinle, November 1942; C. J. Pinto Fonseca, 1919. July Guinle, H. W. Laemmart, April 1943; P M. 178. Fazenda de Floresta, Rio 19°53'S, Matipo, Britto, January, May 1943. 42°33'W. 189. Boa Vista, Fazenda, Teresopolis (see J. Pinto Fonseca, 1919. July Teresopolis). 178. Esmeralda (Fazenda), ca. 19°20'S, 42°50'W. 189. Boa Fe, Fazenda, Teresopolis, 22°22'S, 178. Monies Claras, 19°25'S, 42°35'W. 41°53'W. 179. Boa Serra de 20°10'S, Esperan9a, Caparao, D. E. Davis, October, September 1943; R 41°46'W. M. Britto, January 1943; G. Perreira, E. 1929. Kaempfer, August September 1942; C. Guinle, November 180. Fazenda Cardosa, Serra de Caparao, 1942; C. Guinle, R M. Britto, March, May 20°22'S, 41°48'W. 1943; C. Guinle, H. W. Laemmart, April E. G. Holt, June 1922. 1943. 181. Benfica, Serra de Itatiaia, 21°4rS, 43°26'W. 189. Novo Friburgo, 22°16'S, 42°32'W. E. H. Holt, June 1922, at 2000 ft. Herr Beschke. 182. Alem Paraiba, 21°52'S, 42°41'W. 189. Barrieira, Serra dos Orgaos, 22°56'S, 183. Juiz de Fora, 21°45'S, 43°20'W. 42°56'W. do Estudos e sobre a Servigo Pesquisas Schind in Pohle, 1927. Febra Amarela (SEPSFA). 189. Serra dos 6rgaos, 22°56'S, 42°56'W. 190. Rio de Janeiro, 22°54'S, 43°14'W. R. M. 1938. Espirito Santo—Philander opossum quica Gilmore, January, February 191. Rodeio, Serra do Mar, 22°33'S, 43°4rw. 184. Santa Teresa, 19°55'S, 40°36'W. G. B. Flowers. A. M. October 1942. Olalla, 191. Sepetiba, 22°58'S, 43°42'W. 184. Sao Joao de Petropolis, 19°49'S, 40°40'W. J. Natterer, March 1818. C. June 1940. Lako, 191. Sapitiba (see Sepetiba). 185. Serra, 20°07'S, 40°18'W. 191. Sao Joao Marcos, 22°54'S, 43°58'W. C. Lako, September 1949, at 50 m. C. Lako, October 1938, January 1939. 185. 40°17'W. Campinho, 20°07'S, 192. Mangaratiba, 22°57'S, 44°02'W. C. at 500 m. Lako, April 1940, C. Lako, July, September 1938, February 185. Vitoria, 20°19'S, 40°2rw. 1939. C. Lako, April 1940. 193. Itatiaia Parque Nacional, 22°30'S, 44°34'W. 185. Vila Velha, Morro de Angoles, 20°20'S, F. Gouvea, July 1957. 40°17'W. 193. Monte Serrat, Parque Nacional de Itatiaya. C. Lako, March, April 1940, June 1941. C. Moreira, 1901; F. Gouvea, March 186. Caixa D'agua, 20°38'S, 40°55'W. 1951, at 800 m, August 1957, at 850 m. C. Lako, June 1940. 193. Macieiras Itatiaia, 22°30'S, 44°34'W. 187. Valao de Sao Lourengo, Santa Teresa, F. Lima, December 1949; F. Gouvea, 2o°37's, 4r4rw. December 1949. C. Lako, June 1940. 193. Barro Branco, 22°23'S, 44°30'W. 188. Reeve Engenheiro (now Rive), 20°46'S, A. Passarell, April 1941. 41°28'W. 194. Pedra Branco, Parati, 23°13'S, 44°43'W. A. 1903. Robert, March, April C. Lako, July 1943.

Rio de Janeiro—Philander opossum quica Sao Paulo—Philander opossum quica

189. Theresopolis (see Teresopolis). 195. Avanhandava, Lajeado, 20°57'S, 48°46'W. 189. Teresopolis, 22°27'S, 42°57'W. E. Garbe, April 1910.

94 FIELDIANA: ZOOLOGY 196. Boraceia, upper Rio Tiete, 22°10'S, 209. Colonia Hansa (see Hansa). 48°45'W. J. Pinto, May 1958; L. Travassos, March, Rio Grande Sul—Philander April 1958; see also Boraceia (203). do opossum quica 197. Iporanga, Lajeado, 24°36'S, 48°34'W. 210. E. Dente, December 1944. Passo Fundo, 28°15'S, 52°20'W. 211. 29°3rS, 50°47'W. 198. Monte Alegre, Amparo, 22°40'S. 46°4rw. Taquara, R. before H. von J. Lima, June 1944. Hensel, 1867; Ihering, = before 1888. 199. Ypanema (Ipanema Bacaetava), 23°26'S, 47°36'W. 200. Sao Paulo, 23°30'S, 46°30'W.

200. Cotia, 12 km W Sao Paulo (q.v.). "I.A.L.," January, July 1961, January Acknowledgments 1962, November 1963. appreciation is for the multitu- 200. Cantareira, Serra da, 23°25'S, 46°39'W. My expressed dinous services performed by research assistants 2(X). Butanta, Serra da Cantereira (q.v.). Eunice Hoshizaki and Barbara Brown, which con- J. Navas, April 1910. tributed substantially to bringing this work to 201. Juquia, 24°19'S, 47°36'W. completion, and to Cameron Pfiffner and Kathleen 201. Costao dos Engenhos, 24°4rs, 47°25'W. Telfer for the illustrations. The unstinted services A. M. Olalla, July 1964. essential to this the Field 202. Alto da Serra, 23°47'S, 46°19'W. production provided by Museum's photography department, headed 202. Vila Oliveira, Magi das Cruzes, 23°31'S, by 46°irw. John Weinstein, and the library staff under Ben- jamin Williams are gratefully acknowledged. J. Lima, September 1943. Thanks are expressed to the authorities of the 203. Salesopolis, Boraceia, 23°32'S, 45°5rw. institutions listed below for permission to study J. Oliveira, June, July 1961. the material in their charge. 203. Casa Grande, 20 km S Salesopolis (q.v.). O. de Sousa Lopez, April 1962, May f 1968. 1966, May Literature Cited 203. Boraceia, Ponta de, 23°48'S, 45°49'W. Alho, C. J. R., L. A. Pereira. and A. C. Paula. 1986. Near Salesopolis Biological Station; not Patterns of habitat utilization by small mammal pop- to be confused with Boraceia (196). ulations in cerrado biome of central Brazil. Mamma- 204. Serra de Itatiaia, 22°55'S, 45°28'W. lia, 5(4): 447-460. 205. 22°37'S, 45°10'W. Piquete, Allen, J. A. 1900. Description of new American mar- Zech, January 1897. supials. Bulletin of the American Museum of Natural 206. Sao Sebastiao, 23°48'S, 45°25'W. History, 13: 191-199.

A. 1900. . 1901a. A preliminary study of the North [ Hempel, July, August American opossums of the genus Didelphis. Bulletin of the American Museum of Natural History, 14: 149-188. Parang—Philander opossum quica . 1901b. Description of two new opossums of the genus Metachirus. Bulletin of the American Mu- 207. Rio Paracai, 23°41'S, 53°57'W. seum of Natural History, 14: 213-218.

. 1911. from Venezuela collected E. Dente, S. Siraglia, January 1954. Mammals by Mr M. A. Carriker. Jr Bulletin of the American Mu- 208. Ro(;a Nova, Serra do Mar, 25°30'S, .seum of Natural 30: 239-273. 48°50'W. HLstory,

. 1912. Mammals from western Colombia. Bul- A. Robert, August, September 1901. letin of the American Mu.seum of Natural History, 31: 71-95.

. 1916a. List of mammals collected for the Santa Catarina—Philander opossum quica American Mu.seum in Ecuador by William B. Rich- ard.son, 1 91 2-1 91 3. Bulletin of the American Museum

of Natural History, 35: 1 13-125. 209. Joinville, 26°18'S, 48°50'W.

. 1916b. List of mammals collected in Colombia E. Steiger, August 1930. by the American Museum of Natural History Expe- 209. Hansa, 26°27'S, 48°50'W. dition I9I0-I9I5. Bulletin of the American Mu.seum W. Ehrhardt, August 1928. of Natural History, 35: 191-238.

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