UNiVLR<- OF AUNOI? MW nitBANA 1BAIQM BIC. .. / 1^^ yy^Ai? ^or ^o.*^^S' '^^sJc^^ )patk ^b\)er em \ v Phil)! PUBLr-.!. r. The FIELDIANA /^ Zoology NEW SERIES, NO. 86 i 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 Published May 30, 1997 101 BURRILL HALL 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 Abstract 1 Introduction 1 Abbreviations 2 Didelphidae: Taxonomic Status, The Genera 2 A Morphological Test 4 Genus Philander Tiedemann (Four-Eyed Pouched Opossums) 8 Synonymy 8 Distribution 8 Nomenclature 10 External Characters and Comparisons 11 Cranial Characters and Comparisons 15 Dental Characters and Comparisons 17 Descriptions of Individual Teeth of Philan- der 18 Dental Abnormalities 23 Sexual Dimorphism 24 Karyology 24 Parasites 25 Species and Named and Unnamed Sub- species OF Philander opposum 32 Philander opossum Linnaeus 33 Philander opossum opossum Linnaeus 35 Philander opossum quica Temminck 40 Philander opossum frenatus Olfers 51 Philander opossum melanurus Thomas .... 51 Philander opossum subsp. nov. 1 52 Philander opossum suhsp. nov.? 52 Philander opossum fuscogriseus J. A. Allen 52 Philander andersoni Osgood 59 Philander andersoni andersoni Osgood .... 61 Philander andersoni mcilhennyi Gardner and Patton 63 Behavior of the Four-Eyed Pouched Opossums (Philander opossum) 64 Alphabetical List of Collecting Locali- ties 80 Gazetteer 85 Acknowledgments 95 Literature Cited 95 List of Illustrations 1 . The four genera of largest American marsupials 2. Skulls of the four largest American marsupials 3. Hindfoot of Chironectes minimus 7 4. Dorsal pigmentation of Philander opossum and P. andersoni 9 5. Geographic distribution of f/?/7flnrfer .... 10 6. Chcmdixdi oi Philander opossum 16 7. Caudal segments of the four largest American opossums 18 8. Scales and bristles of Philander opos- sum tail 19 9. Transitional caudal vertebrae of Philan- der opossum 20 10. Points of cranial measurements 22 11. Diagram of didelphid skull 23 12. Left side of Philander opossum skull and mandible 25 13. Philander opossum sVnW 26 1 4. Tooth rows of Philander opossum 27 15. Upper incisors and canine of Philander opossum, Caluromys philander, Lutreo- lina crassicaudata, Metachirus nudi- caudatus 28 16. Sidi^gered Xy m Philander opossum 29 17. Deciduous premolar 3 and molars 30 18. Accessory molar cusps 31 19. Conules of m^ of Philander from vari- ous geographic localities 32 20. M^ of Philander opossum 33 21. Lectolypes of Philander opossum 39 22. South American collecting locahties 81 List of Tables 1 . Summarized measurements of Philan- der opossum and P. andersoni sub- species 12 2. Dimensions and proportions of limb bones 15 3. Vertebral bone formulae 19 4. Nipple formulae 21 5. Symbols for cranial bones and foramina 24 6. Symbols for dental elements 34 7. Conules of m^ 35 8. Karyotypes of largest American mar- supials 35 9. Measurements of Philander opossum opossum 36 10. Measurements of Philander opossum quica 42 1 1 . Measurements of Philander opossum 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 The four-eyed pouched opossums, genus Phi- lander Tiedemann, 1808, most nearly related to the larger opossums of the genus Didelphis Lin- naeus, 1 758, are common throughout much of the neotropics from northern Argentina into Tamau- lipas, Mexico. About 20 Linnaean names have been proposed for four-eyed pouched opossums, but only two species are recognized here. The ear- liest known, P. opossum Linnaeus, 1758, with its geographic distribution nearly equal to that of the genus, is the smaller of the two and generally less darkly colored. Philander andersoni Osgood, mainly western Amazonian in distribution, con- tains two subspecies. One, P. andersoni andersoni Osgood, occurs in Peru and in the Territorio Fed- eral Amazonas, Venezuela. The wide gap in dis- tribution between the two countries may be for lack of collecting or the result of a climatic event. The blackish P. andersoni mcilhennyi Gardner and Patton is known only from the upper Rio Pu- nis basin in Amazonian Peru. This report is the author's sixth on New World marsupial systematics and behavior. The series began in 1992 with a taxonomic revision of the gracile mouse opossum, genus Gracilinanus (Marmosidae) (Hershkovitz, 1992a). It was fol- lowed in the same year by a critical examination of the significance of ankle bones as phylogenetic indicators (Hershkovitz, 1992b). The description of a seemingly abnormal staggered third lower in- cisor, which proved to be the hallmark of all di- delphimorphs since the earliest Cretaceous, if not late Jurassic, was published in 1995. In press is a taxonomic review and life history account of the relict Chilean mouse opossum Dromiciops (Mi- crobiotheriidae). Also awaiting publication is an- other review, that of the brown four-eyed pouch- less opossum, genus Metachirus. This taxon was used at first as an outgroup for the Philander opus. As work progressed, however, Metachirus took on greater importance and is being complet- ed under its own title. 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- tioned articles have been partially derived and elaborated. Abbreviations The following abbreviations are used for the institutions where the specimens examined are preserved. AMNH - American Museum of Natural His- tory, New York bm(nh) = British Museum (Natural History), London FMNH = Field Museum of Natural History, Chicago Mvzuc = Museum of Vertebrate Zoology, Uni- versity of California, Berkeley USNM = National Museum of Natural History, Washington, D.C. USPMZ - Universidade de Sao Paulo Museu de Zoologia, Sao Paulo, Brasil MPEG = Museu Paraense Emilio Goeldi, Be- lem, Brazil UKMNH = University of Kansas Museum of Natural History, Lawrence LSUMZ = Louisiana State University Museum of Zoology, Baton Rouge MNRJ = Museu Nacional, Rio de Janeiro RMNH = Rijksmuseum van Naturalijke Histo- rie, Leiden Didelphidae: Taxonomic Status, The Genera The family Didelphidae, erected by Gray, 1821, included all then-known American marsupials di- vided into the genera Didelphis Linnaeus and Cheironectes [sic] Illiger. It was not until late in the 20th century that critical attention was paid to higher systematic categories of living New World marsupials. The early arrangement of the taxa persisted, nevertheless, as if all female marsupials were pouched and all individuals prehensile-tailed. scansorial, and members of the genus Didelphis. Discovery of the first caenolestid by Tomes in 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- standing, apart from accommodations made in catalogs for the increasing number of newly dis- covered taxa, no significant changes were made in the classifications. Recent catalogers (Cabrera, 1958; Hall & Kelson, 1959; Honecki et al., 1982; Gardner, 1993) recognized the three current orders of American marsupials, each with a single fam- ily, its content as found in the literature. The cat- egories, with the numbers of their respective liv- ing genera and species in parentheses (ex Gardner, 1993), follow: Order Didelphimorphia Family Didelphidae (15 genera, 63 species) Order Paucituberculata Family Caenolestidae (3, 5) Order Microbiotheria Family Microbiotheriidae (1, 1) Research by Reig et al. (1977) on chromosomes of the Didelphidae resulted in significant advanc- es. Three karyotypes were distinguished among living American marsupials. The largest diploid number of chromosomes was 22, possessed by four genera of American marsupials, namely Di- delphis (three species), Chironectes (one), Lutreo- 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- romysiops (one), Marmosa (sensu lata; eight spe- cies of about 38, since rearranged into four gen- era), Metachirus (one), and Dromiciops (one). The 2n = 14 karyotype is generally regarded as the primitive one. No reclassification of American marsupials was based on their cytogenetic consti- tution. In any case, Kirsch (1977) had already re- moved woolly opossums {Caluromys) from the Didelphidae and erected the family Caluromyidae for them. In 1992, Hershkovitz (1992a) estab- lished the family Marmosidae for most so-called mouse opossums previously known as Marmosa 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 genera were retained in Didelphidae because they are the largest and their karyotype is 2n = 22. These four genera are Didelphis, Chironectes, Philander of Linnaeus, and Lutreolina Thomas, 1910 (Figs. 1-3). Biochemical, chromosomal, and anatomical characters were considered by Reig et al. (1985) in an attempt to determine the phylogenetic rela- tionships among didelphoid marsupials. The ex- tinct forms they considered are outside the scope of this monograph, as are most of the included genera. The Didelphidae of concern here include the three subfamilies Herpetotherinae, Caluromyi- nae, and Didelphinae. The latter was further sub- divided into the tribes Didelphini, Metachirini, and Marmosini. Their tree (Reig et al., 1985, Fig. 2, p. 339) shows all American didelphoids and basic Australian forms emerging from Upper Cre- taceous Microbiotheriidae, an improbability. It 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 seems unlikely. The pouchless and non-prehen- sile-tailed, terrestrial Metachirini with 2n = 14 may have given rise to didelphids rather than the reverse as depicted. Studies by Kirsch et al. (1993a) of DNA/DNA hybridization resulted in assignment of the five largest American genera {Didelphis, Philander, Lutreolina, Chironectes, and Metachirus) to the Didelphidae. Morphologically, as shown below, Lutreolina and Chironectes share few traits with each other and fewer with Didelphis, and the in- dicated position of Metachirus is even less tena- ble. Patton et al. (1995 [1996]) regarded the system- atic position of Metachirus to be somewhat equiv- ocal: While the brown four-eyed opossum Metachirus forms a sister taxon to the {Didelphis + Philander) clade in all trees, this relationship is not well sup- ported. Bootstrap values for parsimony analyses are only 68 and 54 for DNA and amino acid sequences, respectively, and decay indices indicate that it would take only two or three additional steps to collapse the relationship between Metachirus and the other two genera. Similarly, confidence limits for this node are only 28% in the DNA neighbor-joining distance tree. The pouchless, primitively 2n = 14 brown four-eyed opossum contrasts markedly with the pouched, de- rived 2n = 22 Didelphis and Philander pair, although the three genera apparently do share similarities in bullar structure (Reig et al., 1987). Indeed, the total evidence Wagner tree presented by Reig et al. (1987, Fig. 59), based on 45 craniodental, cytological, sero- logical, and soft anatomical characters, does place Metachirus in a sister relationship with Didelphis and Philander, in contrast to Marmosa and Monodelphis, as do DNA-DNA hybridization data (Kirsch et al., 1995; Lapointe and Kirsch, 1995). Nevertheless, placements of Metachirus either as the sister of {Di- delphis + Philander), with the mouse opossum lin- eage, or outside of both clades are all equally likely based on an evaluation of these alternative topologies by the log-likelihood test of Kishino and Hasegawa (1989). As a consequence, perhaps Metachirus is best considered as a basal member of the {Didelphis + Philander) and {{Marmosa -I- Micoureus) + Mono- delphis) clades. This is the same conclusion reached by Kirsch (1977) and Reig et al. (1987). Returning to Philander, Patton et al. (1995 [1996]) do not explain why they dropped the prior name Philander andersoni Osgood, 1913, and consistently used instead its junior synonym, P. mcilhennyi Gardner and Patton, 1972, in all their discussions of marsupial relationships. A IVIorphological Test The four genera of large American marsupials have consistently been regarded as forming a nat- ural assemblage within the family Didelphidae. This can be tested by the 30 morphological char- acters outlined below. Philander shares most characters with Didelphis. Chironectes and Lu- treolina, however, have diverged widely from the preceding two genera and from a hypothetical common ancestor of all four. Two of the few shared characters (large size and In = 22) may have evolved independently; other characters that may appear trivial are diagnostic nevertheless. An appraisal of the evolutionary stage of each char- acter is shown in parentheses. 1. Largest of living New World marsupials (derived) The four genera including the largest liv- ing New World opossums (Fig. 1) range in size from the smallest. Philander, through Chironectes and Lutreolina, to the largest, Didelphis. Size may have evolved indepen- dently in each genus, and its phylogenetic significance in the present context is dubi- ous. 2. Karyotype 2n = 22 (derived) The same karyotype evolved indepen- dently in unrelated Marmosa canescens from a 2n = 14 karyotype (Engstrom & 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). -.Ife HERSHKOVrrZ: PHILANDER Australian Macropodidae and Potoroidae. The phylogenetic significance of the same chromosome number in Chironectes and Lutreolina may be misleading. A different karyotypic pattern in Didelphis virginiana does not alter the fact that this species and D. marsupialis are very nearly alike and most intimately related. 3. Tail as long as or longer than head and body combined (incipiently derived) Excludes Lutreolina 4. Tail fully prehensile (derived) Excludes Lutreolina and Chironectes 5. Tail terete (plesiomorphic) Excludes Lutreolina and Chironectes 6. Tail base not notably thickened (plesio- morphic) Excludes Lutreolina (muscle, not fat) 7. Tail thinly pilose, the scales fully exposed dorsally (plesiomorphic) Excludes Lutreolina 8. Complete marsupium present in females, absent in males (derived) Excludes Lutreolina and Chironectes 9. Thumb and first toe fully opposable (de- rived) Excludes Lutreolina 10. Pedal digits partially or not webbed (incip- iently derived) Excludes Chironectes 1 1 . Crown color pattern whitish above eyes, brownish between (derived) Excludes Chironectes and Lutreolina 12. Ears large, roundish, leaflike, and when laid forward extending midway or more to outer canthus of eye (possibly derived) Excludes Lutreolina and Chironectes 13. Manual ungues sharp and protruding (ple- siomorphic) Excludes Chironectes 14. Paired posteromedian palatal vacuities present (plesiomorphic) Excludes Chironectes 15. Premaxillary symphysis angular (plesio- morphic) Excludes Lutreolina and Chironectes 16. Depression at proximal end of temporal ridges (?) Excludes Lutreolina and Chironectes 17. Interorbital region not notably constricted (derived?) Excludes Lutreolina IS. Supraorbital processes rudimentaiy (incip- iently derived) Excludes Lutreolina and Chironectes 1 9. Spread of zygomatic arches comparatively narrow (plesiomorphic) Excludes Chironectes 20. Lacrymal foramina facial (derived) Excludes Lutreolina and Chironectes 21. Nasals behind convergent to a single point or with slight spread between points (ple- siomorphic?) Excludes Lutreolina and Chironectes 22. Maxillary sheath for insertion of lower ca- nine shallow (plesiomorphic) Excludes Lutreolina and Chironectes 23. Occipital condyles projecting behind ver- tical plane of supraorbital bone (plesio- morphic) Excludes Chironectes 24. Incisive foramina long, narrow, and ta- pered (plesiomorphic) Excludes Chironectes 25. Foramen magnum ovate (?) Excludes Lutreolina 26. Ventral mandibular plane between canine and incisors horizontal (plesiomorphic) Excludes Lutreolina and Chironectes (?) 27. Upper incisors 2-5 with little or no deflec- tion inward and upward (plesiomorphic?) Excludes Lutreolina and Chironectes 28. P^ nearly as long as P' (plesiomorphic?) Excludes Lutreolina 29. Lower incisors not overlapping (staggered ij not included) (plesiomorphic) Excludes Lutreolina and Chironectes 30. Locomotion terrestrial-scansorial (derived) Excludes Lutreolina and Chironectes Taking into account other characters mentioned beyond, comparisons reveal Didelphis as a shag- gy, oversized ''Philander^ The otter-like Chiro- nectes, distinguished mainly by its nonprehensile, naked tail, with both sexes pouched, and the ter- restrial weasel-like Lutreolina, with its nonpre- hensile, completely hairy tail and nonopposable first digit, have radiated far from didelphoid roots, j Lacking affinities with each other of didelphid grade, Chironectes and Lutreolina are each treat- ed as a type of a distinct subfamily. The new ar- rangement follows: Superfamily Didelphoidea Family Didelphidae Gray, 1821 | Subfamily Didelphinae j Genus Philander Tiedemann, 1 808 I Philander opossum Linnaeus, 1758 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 Didelphis marsupialis Linnaeus, 1758 {aurita Wied-Neuwied a syn- onym) Didelphis albiventris Lund, 1840 Didelphis virginiana Kerr, 1792 Subfamily Chironectinae (new) Genus Chironectes Illiger, 1811 Chironectes minimus Zimmer- mann, 1780 Subfamily Lutreolininae (new) Genus Lutreolina Thomas, 1910 iMtreolina crassicaudata Desmarest, 1804 Didelphis is the best known and most widely distributed genus of American marsupials. Its northernmost species, D. virginiana, occurs in North and Middle America; the more southern D. marsupialis, in Mexico and Middle and South America; and D. albiventris, in South America. The North American representatives of D. virgin- iana and D. marsupialis have been taxonomically revised by Gardner (1973). Distinctive characters of the family and sub- families are explicit in each of the 30 characters described above. Detailed accounts of Chironec- tes and Lutreolina await formal taxonomic revi- sion such as given below for Philander. Didelphis is highly derived. Its roots in the past are un- known. The animal that evolved a prehensile tail, opposable first digit, complete marsupium in stages, and other derived characters is far re- moved in time from Didelphis. It is most unlikely HERSHKOVITZ: PHILANDER that the genus Didelphis existed as such in the Early Tertiary or even Late Tertiary. To refer to a staggered-lower-incisored Didelphis as the stock from which the Early Paleocene Bolivian micro- biothere Pucadelphys andinus Marshall and de Muizon (1988) could be derived is anachronistic and morphologically perverted. A taxonomic revision of Philander, the remain- ing genus of the Didelphinae, follows. Compari- sons are made with Didelphis, Chironectes, and Lutreolina. Except for an occasional reference here, Metachirm is dealt with in a separate paper that is in preparation. Genus Philander Tiedemann (Four-Eyed Pouched Opossums) Philander Brisson, 1762:13, 207-214— name from non-Linnaean work and not available. Gilmore, 1941:316 — Metachirops Matschie antedated; char- acters; distribution; yellow fever susceptibility. Hopwood, 1947:533 — names from Brisson not val- id; Philander Gronovius non-Linnaean and not available. Philander T\tdtm2iX\r\, 1808:426 — included species P. virginianus Tiedemann [= D. opossum Linnaeus], P. murinus [= Marmosa murina], P. brachyurus [= Monodelphis brachyura]. Hershkovitz, 1949: 1 1 — type P. virginianus Tiedemann = Didelphis opossum Linnaeus; Philander Brisson, 1762, and Philander Gronovius, 1763, not valid. Hildebrand, 1961:239 — comparative body proportions. Peron- dini and Perondini, 1965:381, Fig. 7 (metaphase chromosomes). Fig. 8 (karyogram) — chromosome complement 2n = 22. Enders, 1966:195 — breeding season; habits. Biggers, 1966:251 — male external genitalia; spermatozoan types; temperature regula- tion. Enders and Enders, 1969:431, PI. 1 (uterus with 5 fetuses). Pis. 2-6 (uterus section) — placenta; fetal membranes; fetal size at birth. Hayman and Martin, 1969:192 — chromosomes (In = 22); chro- mosome phylogeny. Collins, 1973:69 — care and maintenance in captivity; reproduction; growth and development; diet; parasites. Tyndale-Biscoe, 1973:25, 38, 40, 61, 65, 189, 192, 230— biology. Pine, 1973:391 — not the four-eyed pouched opos- sum. Hershkovitz, 1976:295 — history, nomencla- ture. McNab, 1978:115 — comparative bioenerget- ics (P. opossum). Husson, 1978:27 — treated as syn- onym of Didelphis Linnaeus by designation of the Virginia opossum as lectotype of type species P. virginianus Tiedemann. Landsmeer, 1979:337, Figs. 1,4, 11, 12 (cheiridia, extensor assemblies) — digital extensor muscles; claw retraction. Hersh- kovitz, 1981:943 — Philander valid generic name; type P. virginianus Tiedemann - Didelphis opos- sum Linnaeus. Gardner, 1981:447 — Philander val- id; Husson opinion rejected. Rodger, 1982:270 — testis and excurrent ducts; paired spermatozoa. Jen- kins and Knutson, 1983 — type specimens in British Museum (Natural History). Perez-Hernandez, 1985:53, 65 — generic characters. Perez- Hernandez et al., 1986:14 — dental morphology and diet. Gard- ner (in Wilson & Reeder, eds.), 1993:22 — synony- my; species (P. opossum, P. andersoni) distribu- tion. Metachirops Matschie, 1916:262, 267, 268 — included species pallidus Allen, fuscogriseus Allen, grises- cens Allen, melanurus Thomas, opossum Linnaeus, canus Osgood, andersoni Osgood, quica Tem- minck (designated iy ^6), frenata Lichtenstein; tax- onomy. Miranda Ribeiro, 1936:340 — characters. Krumbiegel, 1941:199— review. Pine, 1973:391— valid generic name for four-eyed pouched opos- sum; Philander regarded a synonym of Didelphis. Holothylax Cabrera, 1919:47 — type Didelphis opos- sum Linnaeus by original designation. Metachirus Burmeister, 1854:135 — subgenus of Di- delphys [sic] Linnaeus, part, D. quica Temminck only. Burmeister, 1856:68 — part, M. opossum Lin- naeus only. Thomas, 1888:329 — part, D. opossum Linnaeus only. Sonntag, 1924:743 — comparative tongue anatomy (M. opossum). Hill and Fraser, 1925:196, PI. 1, Fig. 3 (urogenital organs), PI. 4, Figs. 15-17 (vagina) — female urogenital system. Boardman, 1952:848 — hair tracts (A/, opossum). Lyne, 1959:84 — vibrissae (M. opossum). Metacheirus [sic] Sanderson, 1949:787 — misspelling of Metachirus, in combination with M. opossum. Type Species — Philander virginianus Tiede- mann = Didelphis opossum Linnaeus. Key to the Species (see also Fig. 4) Dorsum uniformly dark or grayish, blackish mid- dorsal stripe absent or poorly defined P. opossum (p. 33) Dorsum with well-defined blackish middorsal stripe or band P. andersoni (p. 33) Distribution — Figure 5. Tropical and subtrop- ical forests including second growth, in South and Middle America, from Misiones and Chaco in Ar- gentina, and in Paraguay east of the Rio Paraguay, northward into eastern Brasil as far as Bahia on the southeast, west from the Rio Tocantins-Ara- guaia into Bolivia, and Peru north into Ecuador, Colombia, Venezuela, the Guianan countries, Pan- ama, Costa Rica, Nicaragua, Honduras, Salvador, 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 Philander opossum (a) P.O. opossum ® P- o. frenata © p. o. ?uioa @ p. o. fuscogriseus © p. o. raelanurus (J) P. o. subspecies 6 ® P. o. subspecies £ Phliander andersoni (gl P. a. andersoni Q p. a mcilhennyi Overlap between Fig. 5. Distribution of subspecies of Philander opossum (a-g), P. andersoni (h), and P. andersoni mcilhennyi (i) in South America and Middle America (part inset). Tamaulipas on the Atlantic versant, Oaxaca on the Pacific; altitudinal range from sea level to about 2000 m above. Philander is unknown in the Caribbean islands, the Caribbean coast of Venezuela and Colombia, and the inland highlands (tepuis) of Venezuela. The genus has not been recorded from the Sierra Nevada de Santa Marta and the northwestern low- lands in Colombia. Spotty records from the Bras- ilian Amazonas suggest that Philander may be present throughout the basin east of the Rio Negro and east of the Rio Tocantins-Araguaia. Although Philander is generally absent from the drier parts of the cerrado of central Brasil, it does occur in the gallery forests. The habitats are described in greater detail in the species and subspecies ac- counts. Nomenclature Tiedemann (1808, p. 426) included three nom- inal didelphid species in his Philander. The first of these is ''Virginische opossum, P[hilander]. virginianus (Did. opossum L[innaeus])" and is based on bibliographical references to figures of opossums by Buffon (1763) and Schreber (1739- 1810) and anatomical accounts by Tyson (1698) 10 FIELDIANA: ZOOLOGY and Cowper (1704). His diagnosis, "Korper roth- lich braun. Ueber jedem Augen ein gelbich weiser flecken. Schwanz so lang als der Leib. 1 Fuss and 3 2^11 lang ohne den Schwanz," is of a four-eyed opossum, and Tiedemann's use of Didelphis opos- sum Linnaeus in apposition definitely restricts the name to the pouched four-eyed species. The other two species are "P. murinus {Did. murina L.)," a Marmosa, and "P. brachyurus {Did. brachyuros Penn.)," a Monodelphis. Philander Tiedemann derives ultimately from Seba's (1734) vernacular names, '^Philander, opossum s. Carigueja/' cited by Linnaeus (1758) in his description of D[idelphis]. opossum. Phi- lander virginianus, a substitute name for D. opos- sum Linnaeus, is type by subsequent designation (Hershkovitz, 1949, p. 11). Didelphis philander Linnaeus, 1758, designated type by Thomas (1888, p. 336), was not included in the original erection of the genus Philander and hence is not valid as type. Pine (1973, p. 391) and Husson (1978, p. 27) argued against the use of Philander Tiedemann for the four-eyed pouched opossum and elected Metachirops Matschie, a junior synonym, instead. The arguments of both authors were discussed and rejected by Hershkovitz (1976; 1981) and by Gardner (1981, p. 44), who summarized and laid the dispute to rest. External Characters and Comparisons Body Size and Form — Species of the genus Philander average smaller than those of Didel- phis, Lutreolina, and Chironectes but larger than all other living didelphoids. Average combined head and body length is about 281 mm (slightly less in females than males); condylobasal length of skull, an indicator of body size, is about 72 mm. Trunk is elongate; muzzle long and slender; eyes large; ears conspicuous. Philander andersoni averages slightly larger than P. opossum, and geo- graphically peripheral races of each species av- erage larger than more central ones. Measurements — Table 1. Growth — Body mass increases after full erup- tion of all molars. Largest individuals of a series or subspecies, however, are not necessarily oldest and may include individuals classified as subadult on the basis of dental eruption or wear and suture closure. Males may not be larger than females at birth but evidently outgrow females to maturity. Adult wild-caught animals that had been main- tained in cages and fed without stint in the now defunct Rockefeller Laboratories in Rio de Janei- ro became extremely large in body mass and cra- nial length. Dental size, however, fixed at com- plete eruption, remains the reliable indicator of "normal" body size. Basic Coloration (Fig. 4) — Trunk dark gray- ish or buffy agouti with or without a blackish middorsal stripe or band; face dark brown or blackish with contrasting supraorbital spot, rhi- narial tip usually unpigmented; tail dark brown or with underside slightly paler, terminal half less pigmented or unpigmented. The Philander color pattern of a dark, modified agouti coat, dark brown to buffy head, dark brown tail and ears, could give rise to all colors and color combinations present in didelphoids, including Didelphis, Lutreolina, and Chironectes. A dark brown eye ring is characteristic of nearly all di- delphoids, and supraorbital spots such as those of Philander expand into temporal bands in Chiro- nectes and the whitish face of many Didelphis. The more or less pigmented ears and tails of di- delphoids derive from fully pigmented ones. The saturate, bleached, or whitish cover hairs evolved from agouti cover hairs. Blackish (melanistic), grayish, orange, or reddish phases, whitish guard hairs and wool hairs, can be derived from the ba- sic color pattern of P. opossum. Coloration of body and limbs, tail, and ears of Lutreolina and the brindled body coloration of Chironectes can also be derived from a primitive Philander model. The saturate, pheomelanic facial pattern of Lu- treolina, without markings, however, is rare among didelphoids. The nearest common ances- tral facial pattern of both Lutreolina and Philan- der must have been nearly uniformly modified agouti. Color Phases — A "gray" or grayish brown phase and a "brown," orange-brown, or orange phase occur in both species of Philander. In the gray phase, the subterminal or pheomelanic bands of the cover hairs of dorsum and sides are bleached to pale buff or nearly colorless. The gen- eral effect is a grayish brown appearance. In the brown phase, the subterminal bands are more densely pigmented, giving the animal an ochra- ceous or orange appearance. Hairs of underparts in gray-phase individuals are usually bicolor with the dark eumelanic bases showing through. In brown-phase individuals, the ventral hairs are usually or predominantly pheomelanic to roots. The color term "brown," used by Mustrangi HERSHKOVITZ: PHILANDER 11 Table 1 . Summary of measurements of subspecies of Philander opossum and subspecies of P. andersoni. Mea- surement means (extremes in parentheses) and number of specimens. Head and Body Tail Hind Foot Ear 290 (235-343) 55 285(245-350)31 288 (245-350) 86 292(195-355)63 279(220-310)55 291 (195-355) 118 264 (250-276) 7 270, 229 261 (229-276) 9 Philander opossum opossum Linnaeus r) "^^ rj o r-' r-^ 00 00 r- 00 vo vo r- >r) m Tt Tt OV ,6 (55-62) ,63 .9 (47-55) ,4 (52-58) ,1 (56-72) ,44 r~ 00 f<^ w^ ■<* ri m in o Tt «n Tf Tt OV -182) -266) -259) -173) 158.8(134- 134, 145 241 (221- 231 (197- 153.3(141- 180, 205 >n Tt n- ov -62) -85) -90) -62) 9 Tt r~ 00 Tj- to >n ^ r- vo m >n 00 "'^ U-) fS (N vO ■* — d r-' 00 -^ \r) Wi 00 t^ vi in *rt -^ T}- ON \0 00 On vO .rt. 1 111^ t-~ o o 00 r^ in ^ oo r~- >o >o p ^^ >n in p CO Q vo ri — '^ ro ^ m 00 00 nO m m Tf rf On — — ro O •^ f^ fT "^00 .0(44- .43 .5(61- .0 (55- .3(44- .3, 48. vo 00 nd m r~ m Tt r<-) nO vO Tl- Tl- sitional vertebra 5 differs notably from both the caudaJ j 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. Taxon Prime Nipple Formulae [Functional formulae in brackets] Source DiDELPHIDAE Didelphini Philander andersoni [2-1-2] da Silva and Patton. pers. comm. andersoni [2-0-2] da Silva and Patton. pers. comm. andersoni [3-0-3] da Silva and Patton. pers. comm. andersoni 3-1-3 = 7(4) da Silva and Patton. pers. comm. opossum 3-1-3 = 7(4) Thomas, 1888 opossum 7 Osgood. 1921:75 opossum 3-1-3 = 7(4) Hershkovitz. field notes (fmnh) opossum 3-1-3 = 7(4) da Silva and Patton. pers. comm. Didelphis albiventris 5-1-5= 11 (2) Hershkovitz, field notes (usnm) albiventris [4-1-3 = 8] (2) Hershkovitz, field notes (usnm) albiventris 11(2) Carlsson, 1903:490 albiventris 5-1-5= 11 Krieg. 1924:652, Fig. 1 marsupialis 5-1-5= 11 (4) Hershkovitz, field notes (usnm) marsupialis [4-1-4 = 9] (3) Hershkovitz. field notes (usnm) marsupialis [10] O'Connell. 1979:81 virginiana 13 Hartman. 1920:255 virginiana 13 McCrady, 1938:183 Lutreolininae iMtreolina crassicaudata 4-1-4 = 9 Thomas, 1888:335 crassicaudata 5-1-5= 11 Krieg. 1924:652. Fig. c crassicaudata 9 Osgood, 1921:75 crassicaudata [7]-9 Creighton, 1984:24 Chironectinae Chironectes minimus 2-1-2 = 5 Hershkovitz, field notes (usnm) minimus 5 Enders, 1966 minimus [4] 5 Krieg, 1921 minimus 4 to 5 Mondolfi and Medira Padilla, 1958:155 manent molar (m,) except one-fourth to one-third smaller Upper Molars (m' ••) (Figs. 17-20) — Crowns wider than long; second molar (m^) longer and larger than first (m'), slightly shorter to longer than third (m'), fourth molar (m^) about half bulk of third; mesiostyle and distostyle present in m'A but distostyle may be obsolete or absent in m^; trigon cusps of m'-' with metacone largest, para- cone smallest, of m" paracone largest, metacone smallest, often minute; stylocone (conule B) al- ways large and prominent in m'"\ style (conule C) as large or larger than stylocone in m', sub- equal or smaller in m^ subequal or usually small- er and often rudimentary or merely indicated in m\ in m^ buccal cusps small or rudimentary; ac- cessory conules derived from buccal conules B, D, from well defined to mere crenulations, or ab- sent; crest of m' metacrista longest of molar se- ries; paracrista connected with conule B. Lower Molars {n\^_^) — Lingual cingulids and neocingulids absent; second or third molar largest, fourth smallest, largest cusp (paraconid or proto- conid, 1) progressively larger than metaconid from m, to m4; paraconid smaller than or subequal to metaconid, hypoconid larger than entoconid; mesiostylid (a or A) not certainly definable in ev- ery tooth, distostylid (b or E) well defined, its size increasing from first to last tooth, in m4 often as high or higher than paraconid; postentoconid larg- er than distostylid, becoming larger than paracon- id in following teeth. Variation — Consistent dental differences be- tween Philander opossum and Philander ander- HERSHKOVITZ: PHILANDER 21 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- ferentiation does occur. In a comparison of ''Phi- lander mcilhennyi" from Balta, Peni, with sym- patric Philander opossum quica, Gardner and Patton (1972, p. 3) noted a relatively deep inden- tation of the labial margin of m^ present in the first species but not in the second. The indenta- tion, or so-called "notch," is the angle or valley between conule C (stylar cone C) and adjacent conule B (Fig. 19). In four specimens at hand of Balta Philander andersoni mcilhennyi, conule C of m"* is well de- veloped or molarized in three females but quite worn in the single male. In 8 adults of sympatric Balta P. opossum quica, I find conule C distinctly less developed in three females and rudimentary or virtually absent in 5 males. Examination of all available specimens from throughout the range of the genus, however, reveals that variation from present to absent, or degree of molarization of conule C, is about the same for both species (Ta- ble 7). Conule C, like any other derived excrescence of the buccal shelf, is firmly established, cusplike, and usually subequal to conule B in m' and m^. In m\ however, it varies geographically from a 1 rudiment or anlage to a fair-sized cusp nearly ] equal in bulk to conule B (Fig. 19). Its molariza- I tion increases gradually from the most primitive ! and geographically central forms of P. andersoni ' and P. opossum to their most derived or geo^ ; graphically peripheral representatives. . Comparisons — Dentition of Didelphis is essen- 1 tially like that of Philander. That of Chironectes j is generally heavier, average transverse width oft 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 second lower (i,) less abruptly staggered; unworn incisors of Chironectes triangular in lateral out- line, but shape more nearly equilateral than ob- tuse; upper incisors of Chironectes and Lutreolina notably inflected inward and upward; first pre- molars less reduced, gap between first and middle premolars narrower in Chironectes, varying from less than half the crown length of anterior tooth to absent; molar crowns more hypsodont in Chi- ronectes and Lutreolina, outline of occlusal sur- face more nearly square or rectangular than tri- angular, the outer posterior angle, particularly of m\ less produced. Dental Abnormalities Supernumeraries Philander andersoni andersoni AMNH 72017, adult 9, Rfo Curaray, Loreto, Peru Left supernumerary m', slightly smaller than normal m', rotated 180° with buc- cal shelf lingual. Philander opossum fuscogriseus AMNH 34373, adult 6, Bagadd, Choc6, Co- lombia Left supernumerary m^, fully erupted, slightly smaller than normal m'*, otherwise similar. Right supernumerary m^ nearly fully erupted but impacted and rotated about 90° with buccal side anterior and overlapped by posterior portion of normal m". Lower molars normal in number and form. Philander opossum quica i^UMZ 12009, adult $, Balta, Ucayali, Peru Right supernumerary pm', in diastema be- HERSHKOVITZ: PHILANDER 23 Table 5. Explanation of cranial symbols 1-23 and a-z in Figures 1 1 and 1 2 A. Bones 1. Nasal 2. Frontal 3. Parietal 4. Supraoccipital 5. Premaxillary 6. Maxillary 7. Lacrymal 8. Jugal, zygomatic 9. Squamosal, temporal 10. Sphenoid (includes orbitosphenoid, alisphenoid, presphenoid, basisphenoid, pterygoid) 1 1 . Palatine 12. Orbitosphenoid 13. Pterygoid process 14. Presphenoid 15. Basisphenoid 16. Alisphenoid 17. Alisphenoidal wing of auditory bulla 18. Ectotympanic 19. Periotic wing of auditory bulla 20. Mastoid (temporal) 21. Basioccipital 22. Occipital condyle 23. Exoccipital Foramina, Fissures, Processes, Fossae, and Crests a. External nares b. Infraorbital foramen c. Lacrymal foramina or canals d. Canine fossa e. Posterolateral vacuity or foramen f. Sphenorbital fissure (concealed above II) g. Foramen rotundum h. Foramen ovale i. Tympanic membrane and auditory meatus (see 18) j. Postglenoid foramen k. Stylomastoid foramen 1. Jugular foramen m. Hypoglossal foramen and or condylar foramen n. Carotid foramen or canal o. Anterior lacerate foramen or petrotympanic fissure p. Foramen magnum q. Glenoid fossa r. Premaxillary or incisive foramen s. Maxillopalatine or mesolateral vacuity t. Posteromedial or palatine vacuity u. Ascending postorbital (zygomatic) process V. Ascending postorbital (zygomatic) process w. Postglenoid process X. Temporal ridge y. Sagittal crest z. Lambdoidal crest tween normal first and second premolars, fully erupted, normal in appearance except rotated about 100° with posterior border linguad and slightly anteriad; heel with pronounced distostylid (hypoconulid) as in left pm'. "Normal" first premolar without talonid presumably a result of crowding by developing supernumerary. \ FMNH 114702, adult 6, San Ramon, El Beni, Bolivia Right supernumerary i,, behind normal i,, occludes with normal i'; normal i; oc- cludes with i^ and so on to ij, which bites into diastema between i' and i,; position of i, more medial and anteriad compared to left homologue; jaw, teeth, and occlu- sion otherwise normal. FMNH 114685, 9 Left upper premolar with incipient but well-defined metacone. Missing molar Philander opossum opossum USNM 393600, adult 9, Utinga, Para, Brasil m'' missing; no sign of alveolus but arcade otherwise normal; m4 normal; teeth mod- erately worn. Sexual Dimorphism Mature males with four molars fully erupted average larger than comparable females in body mass, skull length, and canine size. Postcanine teeth also average larger, but relative to skull length no larger than those of females or even smaller; the braincase is relatively narrower in males, cranial crests more developed, but the sag- ittal crest of old males is sometimes equaled by that of extremely large or old females. Karyology Descriptions of karyotypes of the present and supposed species of Didelphidae (Table 8) were first published in nearly 20 papers. All publica- tions were critically reviewed, with new data add- ed by Reig et al. (1977). The authors studied and described the chromosomes of 177 species ol American marsupials representing nine gener? 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 here for documentation. * ! The "standard" karyotype as understood bji Reig et al. (1977) is present in all treated as di { delphids with few minor exceptions. All listed ii Table 8 share the diploid number of 22. The fun | 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 in all but are metacentric in the X chromosome of D. virginiana and L crassicaudata. It appears that the most variable complement among the four genera of Table 8 is D. virginiana. Parasites Captured marsupials are mostly parasitized by fleas, lice, and staphylinid beetles. These mobile parasites usually leave a live caged animal shortly after capture and desert the host almost inmiedi- ately at death. The list of parasites is not exhaustive. It was compiled mainly as a source of reference for dis- cussions on marsupial zoogeography. The taxo- nomic names used are those of the sources cited. Viruses Mucambo (Potkay, 1977; Pard, Brasil; 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 aspects and left lateral 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, Panama; Grayson & Galindo, 1968, in Potkay, 1977) Eastern equine encephalitis (Bahia, Brasil; Shope et al., 1966, in Potkay, 1977) St. Louis encephalitis (Bahia, Brasil; Shope et al., 1966) Group B (mosquito borne) Ilheus (Bahia, Brasil; Laemmert, 1967) Yellow fever (Bahia, Brasil; Laenmiert al., 1946) Group C Itaqui (Bahia, Brasil; Shope, 1967a) I: 26 FIELDIANA: ZOOLOGli Fig. 14. Upper and lower jaws with tooth rows of Philander opossum. Capim Group Acara (Bahia, Brasil; Belem Virus Labo- ratory, 1976c) Mosquito-borne group Turlock (Bahia, Brasil; Shope et al., 1966) Phlebotomus-bome Itaporanga (Bahia, Brasil; Trapp & Shope, 1967) Vescicular stomatitis group New Jersey strain (Panamd; Tesh et al., 1969) Indiana strain (Panamd; Tesh et al., 1969) Unknown vectors Piry (Bahia, Brasil; Belem Virus Labora- tory, 1967a) Pacui (Bahia, Brasil; Bel^m Virus Labo- ratory, 1967b) Oochoristica braziliensis (Colombia; Baer, 1927) Sparganum reptans (Imperial Bureau of Ag- ricultural Parasitology, 1933) Protozoa Babesia brasiliensis Regendanz and Kikuth (Brasil; Regendanz & Kikuth, 1928, p. 1567) Sarcocystis garnhami (British Honduras; Lainson & Shaw, 1969) Trypanosoma cruzi (Brasil; Wood &. Wood, 1941; Costa Rica; Deane, 1961, 1964) Trypanosoma rangeli-like (Pard, Brasil; Deane, 1964) Fungus Histoplasma capsulatum (Canal Zone, Pana- md; Taylor & Shacklette, 1962) Mallophaga (lice) Gliricola porcelli (S§o Paulo, Brasil; Hop- kins, 1949) lERSHKOVrrZ: PHILANDER 27 A Philander opossum FMNH 16398 Caluromys philander FMNH 21725 B Philander opossum FMNH 11A698 Lutreolina crassicaudata FMNH 5394A Philander opossum FMNH 11A711 Metachirus nudicaudatus 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. I Gryopus ovalis (Sao Paulo, Brasil; Hopkins, 1949) Trimenopon hispidium (Sao Paulo, Brasil; Hopkins, 1949) Acarina (mites, ticks, chiggers) Archemyobia pectinata Mendez (Panamd; Mendez, 1972, p. 615) Amblyomma auricularium Conil (Panama; ! Fairchild et al., 1966, pp. 191, 209) j Amblyomma geayi Neumann (Panama; Fair* child et al., 1966, pp. 195, 209) , i Amblyomma sp. (Panama; Fairchild et al.^ 1966, pp. 209; Venezuela; Jones et al., | 1972) Androlaelaps fahrenholzi Belese, 1911 (Ven^ ezuela; Furman, 1972) 28 FIELDIANA: ZOOLOGY The staggered marsupial incisor staggered alveolus buttress I mm PHILANDER OPOSSUM Fig. 16. Alveolus of staggered ij, and buttress, in Philander opossum (from Hershkovitz, 1982). Crotiscus disdentatus Boshell and Kerr (Pan- amd; Brennan & Yunker, 1966, p. 260) Pseudoschoengastia bulbifera Brennan (Pan- am^; Brennan & Yunker, 1966, p. 260) Trombicula dunni Ewing (Panamd; Brennan & Yunker, 1966, p. 260) Trombicula keenani Brennan and Yunker (Panamd; Brennan & Yunker, 1966, p. 260) Eutrombicula alfreddugesi Oudeman, 1910 (Venezuela; Brennan & Reid, 1974; Pana- md; Brennan &. Yunker, 1966, p. 260) Eutrombicula goeldii Oudeman, 1910 (Ven- ezuela; Brennan & Reid, 1974; Panamd; Brennan & Yunker, 1966, p. 260) Eutrombicula tropita Ewing, 1925 (Venezue- la; Brennan & Reid, 1974) Euschoengastia nunezi Hoffmann (Panam^; Brennan &. Yunker, 1966. p. 260) Ixodes lasallei Mendez and Ortiz, 1958 (Ven- ezuela; Jones et al., 1972) Ixodes luciae Senevet, 1940 (Venezuela; Jones et a!., 1972) Ixodes venezuelensis Kohls, 1953 (Venezue- la; Jones et al., 1972) Ixodes luciae Senevet (Panam^; Fairchild et al., 1966, p. 209) Tur apicalis Furman and Tipton, 1961 (Ven- ezuela; Furman, 1972) Tur uniscutatus Turk (Panama; Tipton et al., 1966, pp. 41, 42) Omithonyssus wemecki Fonseca (Panam^; Yunker & Radovsky, 1966, p. 92) Haemolaelaps glasgowi Ewing (Panam^; Tipton et al., 1966, p. 34) Nematoda Aspidodera sp. (Venezuela; Guerrero, 1985) Capillaria sp. (Venezuela; Guerrero, 1985) Travassastrongylus sp. (Venezuela; Guerrero, 1985) Moennigia sp. (Venezuela; Guerrero, 1985) Strongvloides sp. (Venezuela; Guerrero, 1985) Viarinaia barusi Guerrero, 1983 (Venezuela; Guerrero, 1983) Viannaia conspicua (Brasil; Imperial Bureau of Agricultural Parasitology, 1933, ex Pot- kay, 1977) Rhopalia horridus (Angra dos Reis; Imperial Bureau of Agricultural Parasitology, 1933, ex Potkay, 1977) Viannaia minispicula Guerrero, 1983 (Vene- zuela; Guerrero, 1983) Viannaia skrjabini Lent and Freitas, 1937 (Venezuela; Guerrero, 1983) Viannaia tenorai Guerrero, 1983 (Venezuela; Guerrero, 1983) Viannaia vianniai Travassos, 1914 (Venezue- la; Guerrero, 1983) Cestoda Linistowia iheringi (Brasil; Beveredge, 1982, p. 107) 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 Amblyopinus henseli (Serra dos 6rgaos, An- gra dos Reis, Brasil; Tijuca; Seevers, 1955, p. 247) Siphonaptera (Fleas) Adoratopsylla intermedia intermedia Wagner, 1901 (Venezuela; Tipton & Machado, 1972) Adoratopsylla intermedia copha Jordan (Pan- ama; Tipton & Mendez, 1966, p. 326; SW Colombia; Mendez, 1977, p. 166) Adoratopsylla antiquorum cunhai Pinto (Bra- sil; Rio de Janeiro, Costa Lima & Hatha- way, 1946, p. 228) Ctenocephalides felis Jordan and Rothschild (cosmopolitan; Costa Lima & Hathaway, 1946, p. 213) Xenopsylla cheopsis Rothschild (cosmopoli- 30 FIELDIANA: ZOOLOGY ex Simpson 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- lean et al., 1992) Amphimeruse ruparupu Kifune and Uyema (Peni; Tantaledn et al., 1992) Fig. 18. Conules (stylar cusps, ectostyles, terminal styles) of m\ 1, buccal view from Bensley (1906, p. 6); 2, occlusal view from Simpson (1929, p. 1 19); 3, buccal view from Hershkovitz (1977, p. 287); arrow points to ectoflexus. See Tables 6-7 for explanation of symbols. tan; Costa Lima & Hathaway, 1946, p. 228) Polygenis roberti beebei I. Fox (Panamd; Tipton & M^ndez, 1966, p. 326) Polygenis klagesi klagesi Rothschild (Brasil: Pard, Amazonas, Colombia; Panama; Ven- ezuela; Costa Lima & Hathaway, 1946, p. 142) Rhopalopsyllus australis tupinus Jordan and Rothschild (Panamd; Tipton & M^ndez, 1966, p. 326) Rhopalopsyllus cacicus saevus Jordan and Rothschild (Panamd; Tipton & M6ndez, 1966, p. 326) Rhopalopsyllus lutzi lutzi Baker (Argentina; Paraguay; Brasil: Goias, Minas Gerais; Costa Lima & Hathaway, 1946, p. 139) Neotyplocercus rosenbergi Roth.schild (Ec- uador; Peni; Costa Lima & Hathaway, 1946, p. 229) Tritopsylla intermedia intermedia Wagner (Brasil; Colombia; Ecuador; Paraguay; Central America; Costa Lima & Hathaway, 1946. p. 228) Trematoda Duboisiella proloba Baer (Peru; Tantaledn et al., 1992) [Bursotrema tetracotyloides Szi- dat (Peni; Tantaledn et al., 1992)] Zonorchis allentoshi Foster (Peni; Tantaledn et al., 1992) Plagiorchis didelphidis Foster (Peni; Tanta- lean et al., 1992) Collins (1973, pp. 4, 74) listed the following genera of parasites of Philander without more in- formation or regard for phylogenetic affinities. 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, Maryland." Arthropoda Adoratopsylla, flea Amblyomma, tick Ctenocephalides, flea Crotiscus, chigger Euschoengastia, chigger Eutrombicula, chig- ger Gigantolaelaps, louse Gliricola,^ louse Gyropus,^ louse Haemolaelaps, mite Heterothrombidium Intercutestrix Ixodes, tick Leeuwenhoekia Microthrombidium Neothyphloceras, flea Omithonyssus, mite Pentastoma (see Porocephalus) Polygenis, flea Porocephalus, Pentastoma Pseudoschongastia, chigger Pulix, flea Rhopalias Rhopalopsyllus, flea Schongastia Trimenopon,^ louse Tritopsylla, flea Trombicula, chigger Tur, Acarina Xenopsylla, flea Acanthocephala Echinorhynchus Hamanniella Cestoda Oorchoristica Sparganum Nematoda Aspidodera Capillaria Cortiamosoides Cruzia Globocephalus Gnathostoma Gongylonemoides HERSHKOVITZ: PHILANDER '"All refer to one captive individual and are certainly due to contamination," Wemeck, in Hopkins, 1949, p. 439. BIOLOGY LIBRARY 101 BURWLL HALL 31 nrT 1^^1997 12010 P. opossum quica Balta, Peru cf (old) 16393 P. andersoni andersoni Balta, Peru 9 (adult) 14012 P. opossum quica Balta, Peru 9 (yg) 16398 P. andersoni andersoni Huanhuachays , Peru 9 (adult) 95313 P. opossum opossum Makerie, Suriname 41446 P. andersoni andersoni Montalvo, Ecuador cf (adult) 13799 P. opossum fuscogriseus Achotal, Mexico 9 (subad) 87123 P. andersoni andersoni Santa Luisa, Peru 13801 P. opossum fuscogriseus Achotal, Mexico 9 (subad) 388405 P. andersoni andersoni 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 Oxysoma Philostrongylus Physaloptera Skrjabinofilaria Subulura Travassostrongylus Trichuris Viannaia Protozoa Besnoitia Haemogregaraina Sarcocystis Trypanosoma Trematoda Brachylaemus Maritrema Opisthorchis Paragonimus Phaneropsolus Plagiorchis Platynosomum Podospathalium Zonorchis Species and Named and Unnamed Subspecies of Philander opossum The taxonomic accounts are arranged in the fol- lowing order. 1. Philander Philander Philander Philander Philander Philander Philander opposum opossum opossum opossum opossum opossum opossum Linnaeus opossum Linnaeus quica Temminck frenatus Olfers melanurus Thomas subspecies nov. 1 subspecies nov. ? 32 FIELDIANA: ZOOLOGY Fig. 20. Left upper third molar and right lower third molar of Philander opossum. See Table 6 for names of features. Philander opossum fuscogriseus J. A. Allen Philander andersoni Osgood Philander andersoni andersoni Osgood Philander andersoni mcilhennyi Gardner and Patton Philander opossum Linnaeus (Synonyms under subspecies heading) Distribution (Fig. 5) — Distribution as for the genus, except replaced by P. andersoni in Ama- zonian Colombia and Ecuador and contiguous parts of Amazonian Peru, and in central Peni in the departments of Pasco, Junin, and Ayacucho. Philander opossum and P. andersoni are sympat- ric in the upper Rfo Punis and mid and lower Rio Ucayali basins. Historic Variation — The nearest putative an- cestor 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 sup- ported 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 differenti- ated into the racial forms recognized here. The most widely distributed or largest portion of the erstwhile continuously distributed ancestral pop- ulation is P. opossum quica. An undescribed population {P. opossum subsp. nov. 1) of northeastern Venezuela (Fig. 5, 0 is said to be very dark. The isolated P. opossum me- lanurus 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 unde- scribed P. opossum on the northeast. The geographic range of the southeastward- spreading P. andersoni overlaps that of the west- ward-advancing P. opossum quica. Cranial and Dental Characters — Cranial and dental characters as for the genus, compari- sons included. Coloration (Fig. 4) — Color pattern of head as for the genus; cheeks, chin, throat, neck, chest, belly, inner sides of limbs ochraceous to buffy or whitish; marsupial region brown, scrotum un- pigmented or partially to completely pigmented; lateral line, if present, usually confined to pelvic region as a hip patch, less frequently as a shoul- der stripe or patch, color buffy to ochraceous or- ange, often appearing olivaceous when pale hair ba.ses show through; crown, midline of nape, and dorsum to basal hairy portion of tail brownish; middorsal band, if present, poorly to strongly de- fined, individual cover hairs agouti (banded), tip brown (eumelanin), buffy (pheomelanin) subter- minally, followed proximally by a brown band merging into curly and distinctly thinner grayish HERSHKOVITZ: PHILANDER 33 Table 6. Explanation of symbols used for marsupial dentition (Figs. 1 7-20). Eocone (paracone) Protocone Metacone Upper teeth Cones' 1 2 3 4 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 Ptrf trf taf Pretrigon basin or fossa Trigon basin or fossa Talon basin or fossa Cingula and U cingulids Primary F G H Buccal or external (buccal shelf) Anterolingual or anterior (primary lingual shelf) Posterolingual cingulum Lower teeth Conids' I 2 3 4 5 Eoconid (protoconid) Metaconid Paraconid Hypoconid Entoconid Conulids and stylids- a Mesiostylid (parastylid) Distostylid (hypoconulid) Cristids' Cristids extend from-to or between cuspids; refer- ence cuspids in parentheses are not elements of the indicated cristid. / Eocristid [l-a-j-k-l-m-b] II Centrocristid [7^] 777 Epicristid [7-2] IV Postmetacristid [4-b] V Plagiocristid [2-4] VI Protocristid (crested portion of cingulum B) Basins or fossids ptf "Trigonid" basin or fossid taf Talonid basin or fossid Upper and lower teeth Main enamel folds ex ectoflexus (between 7^) 'Most cones and conids are numbered in the order of their origin and development. -Most conules, conulids, styles, and stylids are listed in the order of their position from buccal to lingual and anterior to posterior. Rare or infrequent elements of the tritubercular trigon are listed opportunistically toward the end. Supernumeraries or gemini of established cusps and cuspids are not identified individually. ""Most cristae (-ids) are numbered in the order of their appearance or development in phylogeny, others are numbered opportunistically; all cristae (-ids) except 7-7 inclusive are modified parts of cingula (ids); homologies of the talonid cristids are not certain in every case. bases; guard hairs uniformly brownish; shoul- ders, sides of body, inner sides of limbs contrast- ingly paler than dorsum, the subterminal bands of cover hairs paler than those of dorsum, the dark guard hairs fewer and shorter; metapodials brown above, digits usually contrastingly paler or unpigmented, plantar surface usually well pig- mented and darker than upper surface; tail naked in appearance except for hairy base, color en- tirely brown or mottled to particolored from tip to as much as distal three-fifths of tail more or i less unpigmented; ears naked and wholly brown ; to nearly entirely unpigmented except for brown- i ish borders. , DiCHROMATiSM — Gray and brown color phases' ! described under the generic heading (p. 11) are { present in P. opossum. In brown phase, the pheo- melanic subterminal band of the cover hairs of ! dorsum and sides is saturate ochraceous or or- ange. The same band in the gray phase is bleached f 34 FIELDIANA: ZOOLOGY 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 a-b b-c c-d d-e Philander andersoni andersoni Venezuela Colombia Ecuador Peru (Andean) Peru (Ucayali) 8 8 4 1 1 2 3 2 2 2 - - - - 6 Philander andersoni mcilhennyi 2 Peni (Ucayali) - - 8 Philander opossum fuscogriseus 2 Mexico Guatemala Honduras Costa Rica Panamd Colombia Ecuador - - 3 1 - - 3 1 2 1 5 1 - 1 Philander opossum melanurus 6 3 1 1 2 Ecuador 2 2- Philander opossum opossum ~ Guianas Brasil (Amazonian) 4 2 1 8 9 3 Philander opossum quica Eastern Brasil Bolivia Peni Venezuela 4 2 1 12 7 3 7 10 2 2 5 8 - to pale buff or nearly white. Old pelage with tips and subterminal bands of hairs eroded, appears brown in either color phase because of greater ex- posure of the dark bases. Hairs of underparts in gray phase are dominantly bicolor with terminal portions pheomelanic, basal portions eumelanic. In brown phase, the hairs are dominantly mono- color pheomelanic. Sexual dichromatism, as seen in P. andersoni, is not evident in P. opossum. Philander opossum opossum Linnaeus Philander, Opassum [sic], sive Carigueja, Bras- iliensis: mas., Seba, 1734:56, PI. 36, Fig. 1 (male [Fig. 21]) — description. Philander, Americanus, sive Carigueja, cum catulus, saccum ventris intrantibus foemina, Seba, 1734:57, PI. 36, Fig. 2 (female with pouch young [Fig. 21]) — description. Carigueja, junior, sive Opassum [sic], cum sacco aperto, Seba, 1734:57, PI. 36, Fig. 3 (young female with open pouch) — de.scription. Sarigue, Buffon [and Daubenton], 1763:279, Pis. 45- 50 (animals, anatomy) — part, anatomical descrip- tion of male (PI. 45) and female (PI. 46) only. Molucca opossum. Shaw, 1800:476, PI. 108. [Didelphis] opossum, Linnaeus, 1758:55. Didelphis opossum, Cuvier, 1798:1 14 — le sarigue. fi. Geoffroy St. Hilaire 1803:141 — characters; habits. Latreille, 1803:304 — sarigue ou opossum. Desma- rest, 1820:256 — characters. Temminck, 1825:41 — characters. Gray, 1843:101. Jentink, 1887:301 — SURINAME; GUYANE FRANgAISE: (.Cayenne). Jentink, 1888:220— SURINAME. J. A. Allen, 1900:195— SURINAME: (type locality). Didelphis oposum [sic], Brongniart, 1792:115 — GUYANE FRANgAISE: {Cayenne). Didelphys opossum, Goldfuss, 1809:215. Waterhouse, 1841:'90— part, GUYANA; SURINAME. Water- house, 1846:485— GUYANA. Thomas, 1888: 329— part, GUYANA: Demerara (Better Hope); SURINAME. Lydekker, 1896:200— part, distribu- tion; characters. Goeldi and Hagmann, 1904:100 — BRASIL: Para; local name, mucurd chichica. Table 8. Karyotypes of largest American marsupials. Taxon In SM A X Y FN N 9,6 Source Philander opossum 22 _ 10 A A 20 7.9 Reig et al. (1977) Philander andersoni 22 - 10 A A 20 4,3 Reiget al. (1977) Didelphis marsupialis 22 - 10 A A 20 19, 11 Reig et al. (1977) Didelphis albiventris 22 - 10 A A 20 3,7 Reiget al. (1977) Didelphis virginiana 22 6 4 M A 32 10,9 Reiget al. (1977) Chironectes minimus 22 - 10 A A 20 -, 1 Reigetal. (1977) Lutreolina crassicaudata 22 - 10 M A 20 1, 2 Reiget al. (1977, p. 199) A = acrocentric; FN = fundamental number of autosomal arms; M = metacentric; SM = submetacentric; ST = subtelocentric; X = $ sex chromosome; Y = cj sex chromosome; In = diploid number of chromosomes; N 9, 6 = number of 9 and 6 samples. HERSHKOVITZ: PHILANDER 35 Table 9. Measurements of Philander opossum opossum Linnaeus. Locality Head and Body Tail Hind Foot Ear Philander opossum opossum Guyana 6 6 All localities - 280,-, 312 -, 298, -, 330 - 42, -, 41 - 38, -,- Guyana 9 9 All localities 254, 287, -, 270 266, 273, -, 264 - — Suriname 6 6 All localities 275 (250-305) 8 271 (254-310)8 41 (40-42) 7 39, 39, 40 Suriname 9 9 Paramaribo 245, 277, 260 262, 264, 269 40, 40,43 32, 36, 30 Guyane Fran^aise 66 Cayenne 272, 257 268, 276 42, 41 37, 34 Guyane Fran9aise 9 Cayenne 292 297 39 41 Brasil 6 6 Serra do Navio 302 (280-320) 4 315(310-320)3 44 (42-45) 4 35 (34-36) 4 Rio Amapari 260, 304 280, 300 42, 50 34, 42 Belem, Utinga 280, 270, 280 280, 290, 285 42, 42, 46 39, 41, 39 Capim 270, 280 290, 320 42, 42 36, 36 Baiao 295, 280 295, 305 41, 42 - Guama 255 280 45 40 Tapareba - - - - Cameta 290 300 35 - Taiuna 290(260-314)24 289 (235-343) 24 45 (41-50) 24 - Santarem Highway 270, 270, 280 300, 285, 310 41, 40,44 30, 30, 32 Altamira - - - - Rosarinho 285 285 35 — Imperatriz 309 321 50 - Baptista - - - - Brasil 9 9 Serra do Navio - _ — — Rio Amapari 270 275 40 35 Murutucu -, 280 -, 258 -, ; ?5 -, 20 Belem, Utinga -, 262, 290 -, 273, 275 -, 41,41 - 38, 40 Mocajuba 260 265 35 - Baiao 268 (252-284) 6 279 (274-290) 5 39.8 (39-40) 5 - Tapareba 273 270 39 38 Cameta 290 350 - - Taiuna 275(261-298) 11 298 (272-334) 1 1 42 (41-43)9 - Santarem Highway 255 290 40 30 D[idelphys]. opossum, Olfers, 1818:204 — part, syn- onymy. Wagner, 1855:226 — listed. Sarigua opossum, Muirhead, 1830(1819):429 — clas- sification. Metachirus opossum, Burmeister, 1856:69, PI. 11, Fig. 1 (skull)— characters. Thomas, 1901b: 153— GUYANA: Rupununi (Rupununi River; Kanuku Mts.). Thomas, 1910:188— GUYANA: Demerara (Supinaam River). Thomas, 191 1:143— SURI- NAME: (type locality); Seba sole reference for opossum Linnaeus. Sonntag, 1924:743, Figs. 39c, 40/ (tongue) — tongue anatomy. Boardman, 1952: 848 — hair tracts. Metachirus opossum opossum, J. A. Allen, 1911: 246— VENEZUELA: Bolivar (Rio Yuruan). Didelphvs (Metachirus) opossum, Menegaux, 1902: 496-er Amazon Basin of Venezuela, probably Colombia, Ecuador to Ucayali Department." Metachirus opossum nigratus Thomas, 1923:603 — PERU: Juni'n (type locality, Utcuyacu, 1600 m; Chanchamayo); type, male, skin and skull, British Museum (Natural History), no. 1900.7.7.62; col- lected 21 April 1900 by P O. Simoms. Thomas, 1928a:250, footnote— PERU: Junin (Inaflez). Gardner and Patton, 1972:5 — "more likely a syn- onym of andersoni" than P. opossum canus, apud Cabrera, 1958. Metachirus opossum Tate, 1939:161— VENEZUELA: Amazonas (Mt. Duida). Philander opossum, Gardner and Patton, part (not Linnaeus), 1972:5— PERU: Ayacucho (San Jos^, Rio Santa Rosa. 10(X) m; Huanhuachayo, 1660 m); comparisons. Reig et al.. 1977:197— PERU: Aya- cucho (San Sos6, Rio Santa Rosa; Huanhuachayo). Philander opossum canus Cabrera (part, not Osgood) 1958:35 — nigratus Thomas only, a synonym. Type — Male, skin and skull. Field Museum of Natural History, no. 19655; collected 11 Septem- ber 1912 by M. P. Anderson. Type Lcx:ality — "Yane Yaku" [= Yanayacu], a chacra, or farm, on the Paranapura River about 8 mi from Yurimaguas. The latter is on the Rio Huallaga at the mouth of the Rio Paranapura (cf. Osgood, 1914, p. 147). Distribution (Fig. 5) — Southeastern Colombia, eastern Ecuador, the Rio Huallaga Valley in the Peruvian departments of Amazonas, Pasco, Junin, and Ayacucho, the Rio Ucayali Valley, Ucayali, and the disjunct population of Territorio Federal Amazonas, Venezuela. Characters — Dark middorsal band well de- fined from sides and continuing over rump to less than basal fifth of tail; hind limbs pale brown, grayish brown or buffy, forelimbs paler with digits usually unpigmented; ventral surface ochra- ceous orange to buffy or grayish buff. Comparisons — Distinguished from Philander andersoni mcilhennyi mainly by paler coloration throughout and narrower, more sharply defined middorsal band; from sympatric representatives of P. opossum by larger size, dark middorsal band and darker coloration throughout. Measurements — Tables 1 and 15. Variation — The Andean andersoni from Huanhuachayo, Ayacucho, Peru, is dark like mcil- hennyi but with dorsal pelage shorter, finer, dorsal band not clearly defined from sides, underparts and limbs paler, tail entirely pigmented except for some mottling on terminal portion. A second An- dean specimen, from San Jos6, Ayacucho, is slightly paler throughout except for a similarly dark brown tail. Its poorly defined dorsal band and nearly uniformly dark tail suggest intergra- dation with P. opossum. Nearest geographic rep- resentatives of P. opossum, however, from Cuzco and Madre de Dios are distinct. The third Andean specimen of andersoni from Chanchamayo, Junin, and another from San Juan, Oxapampa, also lack the well-defined blackish dorsal band of the sub- species. In all other respects, they agree. The con- dylobasal length of the last two, 76.3 and 81.2 mm, respectively, are among the largest of the species. The heavily furred basal portion of the tail is notably extensive in the San Juan individ- HERSHKOVrrZ: PHILANDER 61 Table 15. Measurements of Philander andersoni andersoni. Condylobasal Zygomatic Locality Head and Body Tail Hind Foot Ear Length Breadth Colombia 9 9 La Tagua 270, 274, 264 292, 278, 270 37, 40, 37 34, 36, 39 65.9, 69.1, 64.6 31.6, 34.0, 32.3 Venezuela 66 Capibara -, 270 -, 305 45, 45 42, 45 67.3, 74.5 32.3, 37.6 Tamatama 266, -, - 295, -, - 44, 42, - 43, 42,- -, -, 73.7 36.3, -, 38.2 Acanana _ - - - 71.4 36.4 Mount Duida 243 285 45 - 67.9 36.5 Belen - - - - 75.7 39.9 Venezuela 9 9 Capibara 265 285 41 40 67.3 32.3 Esmeralda 258 267 45 41 67.6 33.7 Tamatama 250 290 36 33 - - Belen 225 300 41 38 67.7 36.5 Merey 250 255 41 - 69.1 35.0 Peru 6 6 Chanchamayo 280 275 47 30 76.3 38.1 San Jose 275 300 48 34 71.3 36.6 Yurimaguas 284 288 40 - 76.3 37.8 Pucallpa 279 313 40 35 73.1 36.5 Santa Luisa 267, 304 265, 282 40, 42 33, 35 -, 73.6 -, 39.3 Rio Curaray - - - - 73.0, 68.9 37.0, - Peru 9 9 Huanhuachayo 270 300 45 35 71.7 36.5 Utcuyacu 265 275 40 35 68.2 32.3 Oxapampa 300 332 51 36 81.2 40.0 Yurimaguas 275 278 35 - 68.6 30.8 Pucallpa 298, 295, 290 279, 280, 290 36, 40, 40 37, 35, 38 -, 70.2, 70.3 -, 35.5, 34.2 Santa Elena 278, 277 270, 263 41, 40 35, 33 68.6, 67.1 33.3, 31.7 Rio Curaray - 68.7, 65.3, 72.3 36.7, 32.9, 35.8 Ecuador 6 6 Pindo Yacu 250, 300 255, 315 40, 40 34, 38 68.3, 76.5 34.0, 39.8 Yana Rumi 278 285 40 32 72.4 38.5 Montalvo 250, 264 280, 262 35, 37 35, 35 74.9, 66.7 37.5, 34.9 Ecuador 9 9 San Jose - - - - 68.5 34.8 ual. A skull from Chanchamayo in the British Museum (Natural History), assigned to nigratus {- andersoni) by Thomas (1923, p. 603), mea- sures 80.7 mm in condylobasal length. Two specimens from Santa Elena, Rio Samiria, between the Ucayali and Huallaga, are interme- diate between typical andersoni, the Pucallpa se- ries of andersoni, and the Balta topotypes of mcil- hennyi. The same departure from heavier pigmen- tation continues westward into the Peruvian An- des (Junfn). In Amazonian Venezuela, however, individuals of P. andersoni andersoni are blackish but with underparts and limbs pale (Capibara, USNM 406960; Belem, USNM 388412). Taxonomic History — The dark Peruvian phi- lander identified by Tschudi (1844, p. 145) as Di- delphys myosuros Temminck may well be the ear- liest mention of the form described by Osgood as P. andersoni. Tschudi (1844, pp. 14, 144, 151) also noted the occurrence in Peni of the then- called D. opossum Linnaeus but mentioned no specimens. In his review of four-eyed pouched opossums, Thomas (1923, p. 603) treated all recognized forms as subspecies of Philander (then Metachirus) epos- < sum. His description of Metachirus opossum nigra- tus (1923, p. 603) conforms to that of a sexually dichromatic male P. andersoni andersoni. Sympatry — H. Hinse, who collected in the Pu- callpa region, took both P. andersoni andersoni 62 FIELDIANA: ZOOLOGY Table 15. Extended. Preorbital Postorbital Braincase Palatal Width Width Width Length i-m'' m'^ 10.4. 12.5. 10.6 8.3. 8.8. 8.9 19.9. 21.1. 19.7 40.1. 41.1, 39.7 35.5. 35.4. 34.2 13.6. 13.0. 12.6 12.3, 14.3 8.8. 8.5 19.6, 22.0 41.2. 44.6 36.5. 38.1 13.5. 13.8 12.6. -. 13.7 8.6. 8.4. 9.4 19.6. -. 22.4 42.6. 43.8, 42.6 37.8. 38.2. 38.5 14.0. 14.0. 14.1 12.6 8.2 21.0 42.5 37.0 13.4 - 8.9 20.8 40.8 36.8 13.5 14.4 8.6 22.4 44.3 38.2 13.5 12.3 8.8 19.6 41.2 36.5 13.5 12.3 8.7 20.0 41.2 36.7 12.9 12.8 8.8 — 41.3 36.3 13.6 12.6 8.0 19.9 40.5 36.3 13.2 - 8.S 19.8 41.2 36.2 13.4 9.2 _ 48.9 43.3 16.0 _ 9.2 21.6 43.4 39.1 14.5 13.7 9.3 21.5 45.8 39.3 13.7 13.5 8.4 20.8 44.5 38.4 14.6 -, - -.9.0 -. 22.5 40.9, 44.2 36.0. 38.9 13.2. 14.4 -, - 8.0. - 22.6. - 44.0, 42.4 38.9. 37.2 14.4. 14.0 14.0 9.8 21.0 44.9 38.8 14.3 12.5 9.4 19.6 42.5 38.7 14.6 15.3 10.1 23.9 49.6 42.9 15.0 13.0 8.0 19.8 41.8 36.8 14.0 - 13.7. 12.7 -. 8.8, 8.7 -. 20.2. 20.7 -, 42.0. 43.0 -. 36.4. 37.2 14.2. 14.1. 13.7 13.0. 12.9 9.3. 8.2 20.5. 19.3 42.2. 41.0 37.3. 37.0 14.8. 14.2 - 9.1. 8.5, 8.9 22.3, 20.5. 22.1 41.6. 39.7, 44.6 36.5, 35.2. 36.8 14.0. 13.4. 13.5 11.8. 14.5 8.7. 8.7 21.2. 21.0 41.8, 46.5 35.2, 38.6 12.0. 13.4 13.6 8.6 21.2 43.3 38.0 14.3 13.9. 13.2 8.8. 9.0 19.9. 20.0 45.0, 40.3 37.8. 35.8 13.9. 13.6 - 9.0 21.3 42.0 37.5 14.0 and the smaller P. opossum quica 59 km W of Pucallpa. October 1971 and 1972 and November 1971. but only P. opossum 59 km NE of Pucallpa (September 1972) and 59 km SW of Pucallpa (October 1972). Specimens Examined — 82. ECUADOR. Napo: Rio Napo, 3 (bm[nh]); Rio Yana Rumi, 1 (fmnh); San Jose, below, 2 (amnh); Pastaza: Copataza, 3 (amnh, 2; FMNH); Rfo Pindo Yacu, 2 (fmnh); Mon- talvo, Rio Bobonaza, 2 (fmnh); Sarayacu, 1 (amnh). COLOMBIA. Caquetd: Tres Troncos, Rio Caqueta, 4 (fmnh); Putumayo: 17 km N Puer- to Asis, 1 (ukmnh). VENEZUELA. Territorio Federal Amazonas: Capibara, Casiquiare Canal, 7 (usnm); Belen, Rio Cunucunuma, 1 (usnm); Acan- i HERSHKOVITZ: PHILANDER ana, Ri'o Cunucunum^, 1 (usmn); Tamatama, 9 (usnm); Playa del Ri'o Base, Mt. Duida. 2 (amnh); Merey. Rio Casiquiare. 1 (amnh); San Juan, Rio Manapiare, 2 (usnm); Esmeralda. 2 (usnm). PERU. Amazonas: 5 mi W Huampani, Ri'o Ce- nepa, 1 (mvzuc); La Poza, Rfo Santiago, 1 (Mvzuc); Ayacucho: Huanhuachayo, 1 (lsumz); San Jos^, Rio Santa Rosa, 1 (lsumz); Junin: Chanchamayo, 2 (bm(nh]; fmnh); Inanez, 1 (bm[nh]); Utcuyacu, 1, holotype of nigratus Tho- mas (bm[nh]); Ucayali: Iquitos. 3 (bm[nh], 2; AMNH); Santa Elena, Ri'o Samiria, 2 (fmnh); Santa Luisa, 2 (fmnh); Yana Yacu, Yurimaguas, 3, in- cluding holotype of andersoni Osgood (fmnh); Orosa, 1 (fmnh); Ri'o Curaray, mouth, 7 (amnh); 63 Table 16. Measurements of Philander andersoni mcilhennyi. Locality Head and Body Tail Hind Foot Ear Condylobasal Length Zygomatic Breadth Peru Si Balta 299 276 43 41 74.4 37.6 Peru ?9 Balta 290, 292 304, 305 44,43 42, 35 73.9, 73.+ 36.8, 36.8 Lago Miranes, Rio Napo, 2 (bm[nh]); Pucallpa, Rio Ucayali, 8 (usnm); Pasco: San Juan, Oxapam- pa, 1 (USNM). Philander andersoni mcilhennyi Gardner and Patton Philander mcilhennyi Gardner and Patton, 1972:2, Fig. 1 (furred tail base). Fig. 2 (skull). Reig et al., 1977: 197, 212, PI. 2E (karyotype)— PERU: Ucayali (Balta, Rio Curanja); karyotype {2n = 22). Patton et al., 1995 [1996]:7, 20, 21— BRASIL: Amazonas (Alto Rio Urucu); relationships based on molecular evidence. Philander andersoni Engstrom and Gardner, 1988:231 — P. mcilhenni [sic] a synonym. Emmons and Feer, 1990:18— part, PERU: Ucayali; ERASE.: Acre. Type — Adult female, skin and skull, Louisiana State University, no. 16395 (originally no. 11541); collected 10 April 1971 by Alfred L. Gardner. Type Locality — Balta, Rio Curanja, upper Rio Funis, Ucayali, Peru; altitude about 300 m. Distribution (Fig. 5) — Known only from the banks of the upper Rio Punis, Ucayali, Peru. Em- mons and Feer (1990, p. 18) included Acre, Bra- sil, in the geographic range based on specimen in the U.S. National Museum of Natural History. Characters — Darkest of four-eyed opossums; dorsal band broad and more or less differentiated from sides, its long, coarse dorsal pelage often manelike, extending over basal fourth or fifth of tail; fore- and hind limbs brown except for contrastingly pale or unpigmented digits of forefeet; ventral sur- face of body dominantly dark brown, variously mixed with gray; throat, chest, and belly often marked with large, irregular, whitish, buffy, or ochraceous patches; two-fifths to three-fifths of ter- minal portion of tail sharply contrasted pale brown or whitish. Comparisons — Distinguished from P. ander- soni andersoni by darker coloration throughout, pelage of dorsal band longer, wider, coarser; tail base bushy, 4-6 cm; from sympatric representa- tives of P. opossum by larger size, overall darker coloration, and broad, dark dorsal band. Measurements — Tables 1 and 16. Remarks — Morphologically, P. andersoni mcilhennyi intergrades completely with P. ander- soni andersoni through the Pucallpa, Rio Ucayali (8 specimens), and Santa Elena, Rio Samiria (2 specimens). The two groups are assigned to P. andersoni andersoni. The geographic origin of the tissue studied by Patton et al. (1995 [1996]) and identified as that of P. mcilhennyi lies well within the area of pro- jected sympatry between P. andersoni mcilhennyi and P. opossum quica (map. Fig. 5). Patton et al. did not explain why they prefer the junior syn- onym mcilhennyi as the species name for the dark gray four-eyed opossum, P. andersoni. Hutterer et al. (1995, p. 15) believe that Phi- lander mcilhennyi and P. andersoni are distinct species, but comparisons were made only between P. mcilhennyi and P. opossum, which are indeed distinct. They point to differences in the color pat- tern of the head and the clipped genal vibrissae of P. opossum. Normally, all facial vibrissae in P. opossum are as in P. mcilhennyi {—P. andersoni mcilhennyi). They also claim that andersoni and opossum are "probably separately confined to the western and eastern cordillera [sic] of the Andes." In fact, both subspecies are confined to the eastern side of the Andes in Peru (infra map. Fig. 5). Specimens Examined — 4. PERU. Ucayali: Bal- ta, 4, including holotype of mcilhennyi Gardner and Patton (lsumz). Behavior of the Four-Eyed Pouched Opossums {Philander opossum) • ( The activities and traits are arranged alphabet- ically. 64 FIELDIANA: ZOOLOGY Table 16. Extended. Preorbital Width Postorbital Width Braincase Width Palatal Length i-m-* m'-* 14.0 14.7, 13.9 9.0 9.2. 9.2 21.2 21.0. 21.0 45.1 44.5. 44.7 39.8 39.7. 39.3 15.3 15.3. 14.9 Aggression, reaction, defense Associations Barbara: The pale-phase four-eyed opossum Blood values Fat storage Foraging Grooming Habitat Home range and territoriality Hydrotropism Locomotion Longevity Maternal care Mobility Nest Population density Breeding Cannibalism Diet Display Enemies, predators Eye shine Population dynamics Posture Reproduction Sleep Sociability Spontaneous bleeding Stress Tail transport Taming Thermoregulation Vocalization Weight Aggression, Reaction, Defense Enders (1935, pp. 410, 412; Canal Zone, PANA- MA) "Next to the Brown Opossum [Metachirus nu- dicaudatus], this is the fiercest fighter of the opos- sums studied. Allen's opossum is ready to fight at all times and no amount of handling appears to diminish its resentment." One shipped live in the same box with a Me- tachirus of approximately the same size "had been thoroughly cowed by the more aggressive Metachirus and showed signs of having been bit- ten severely about the head and neck, while the aggressor was scatheless." Miller (in J. A. Allen. 1916c, p. 589; Mato Gros- so, BRASIL) "When trapped it does not feign death but fights viciously, biting at any object within reach and holding on until its strength fails. They do not hesitate to bite iron bars or steel traps even though it may break the teeth." Wilson (1970; Barro Colorado Island, PANAMA) "About 8:07 PM on the evening of 3 February, 1969, 1 observed an aggressive encounter between Didelphis marsupialis and Philander opossum. The incident occurred in the laboratory clearing near the animal house on Barro Colorado Island, Panama Canal Zone. A loud hissing noise attract- ed my attention to a small Didelphis and a Phi- lander that were circling slowly and snapping at each other on the lawn. Neither showed signs of wounds and both moved adeptly. Each attempted to grasp the rostrum of its opponent. Whenever the Didelphis secured this grasp, it quickly threw the Philander to the ground and held it down with both forefeet and hindfeet, at the same time biting it in the head. Twice in 2 minutes, the Philander broke loose and tried to escape but each time was attacked by the Didelphis. Soon the Philander was effectively pinned on its back and by 8:22 PM had died from the repeated bites on the head, or at least no longer responded to the attacker." Hershkovitz (field notes; COLOMBIA) The usual reaction to apparent or presumed ag- gression is a bipodal or tripodal stance, mouth- open hissing, often with a lurch forward of the body as if to attack. Goldman (1920, p. 52; PANAMA) "Unlike Didelphis when taken in steel traps these opossums are always ready to fight savage- ly" Associations Hershkovitz (field notes; COLOMBIA) Philander opossum was captured in the same HERSHKOVITZ: PHILANDER 65 trap lines with species of Metachirus, Chironec- tes, Didelphis, Caluromys, Monodelphis, and mouse opossums (Marmosinae). All inhabit the same forests. Barbara: The Pale-Phase Four-Eyed Opossum Hershkovitz (field notes, October 1992) A pale-phase female was live-trapped 17 Oc- tober 1992 in a wooded area about 1200 m above sea level in the Parque Nacional do Caparao, east- em Minas Gerais, Brasil. Her nearly completely closed pouch sheltered six attached young, pos- sibly no more than 1 week old. She was kept in her cage for observation of behavior and litter de- velopment. Dubbed Barbara, she accepted captive life with equanimity, making no attempt to escape even when the door of her cage was opened for feeding or cleaning. She ate everything served, whether beef, chicken legs, insects, peanut butter, bananas, or mice. Live mice dropped into the cage were instantly attacked, the head and neck seized and crunched in her jaws. The mice were com- pletely devoured with nothing left of flesh, en- trails, skin, or bones. Barbara slept most of the day curled up under the cut grass provided for nesting. Prepared food or raw fruit given during the day was usually con- sumed at night. Water was imbibed at all hours. Although normally nocturnal, should a live insect or mouse be dropped into the cage during the day, Barbara would awake with a start and make an instant kill and a fast meal. She easily devoured an entire mouse 10 to 15% of her own weight and looked for more. When wanting food Barbara would fix her eyes on the nearest person, raise her muzzle, and sniff expectantly. On one occasion Barbara left intact hide and fur of a mouse otherwise completely consumed. Barbara washed her face frequently after and during meals. One or both paws were used. Grooming forequarters, sides, and underparts usu- ally followed in that order without particular at- tention 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. Blood Values Tyndale-Biscoe (1980, p. 718; COLOMBIA) Blood values measured at sea level (Buenaven- tura), 500 m above (Villavicencio), and 1600 m above (Call) revealed correlations with age and altitude. Hemoglobin concentrations and hemato- crit values were lowest at lowest altitudes, and those for pouch young lower than for adults. Breeding Natterer (in Pelzeln, 1883, p. 110 — Sepetiba, Rio de Janeiro, BRASIL) Female captured 8 March 1818 with five naked, blind pouch young. Female captured 29 September 1822 in Ipane- ma, Rio de Janeiro, with four blind pouch young ("D. dichrura"). Miller (in J. A. Allen, 1916c, pp. 563, 589- um, Mato Grosso, BRASIL) -Uruc- "Number of young is usually small, between three and eight." Davis (1947, p. 2 — Teresopolis, Rio de Janeiro, BRASIL) Seven of 32 females captured in Teresopolis had pouch young, average number, 4.5. Two fe- males each had seven young and seven nipples in pouch. Breeding begins in August and continues until February. Fonseca and Kierulff (1989, p. 1 19- est, Minas Gerais, BRASIL) -Atlantic for- Female captured February and another in Au- gust had five pouch young each. Cerqueira et al. (1993 — Rio de Janeiro, Restinga de Barra de Marica, BRASIL) Total $9 cap- Lac- tured tant January 12 7 April 16 0 July 15 7 October 9 6 66 FIELDIANA: ZOOLOGY Tyndale-Biscoe (1980, p. 713 — Buenaventura, Valle, 10 m; near Villavicencio, Meta, 500 m; near Call, Valle, 1000 m, COLOMBIA) Four females captured August and September 1971, one pregnant, three lactating, and each with five pouch young. Charles-Dominique et al. (1981, p. 425- Cayenne, GUYANE FRANgAISE) -He de Litter size, 4.24 (1-7). Litters mature in 70 days. Lactating females three of 16 in June, two of 13 in July, zero of nine in August. Pouch young survival and food availability: first litter following dry season, 97% (n = 36); second litters, 97% (n = 38); third litters, food scarce, 28% {n = 67). Interval between litters, 90 days. Sexual maturity, 150 days. Seventy-nine percent of October-November population (n = 42) bom in preceding season. Ovarian cycle inhibited by lactation but not by gestation. Charles-Dominique (1983, p. FRAN(;AISE) 409, GUYANE Didelphoid breeding is continuous throughout the year, but successful reproduction is dependent on a rich diet for the female and is seasonal. Up to three successive litters may be produced in 1 year. The third litter "generally occurs at the be- ginning of the period of food scarcity and in most cases the females lose weight and young die in the pouch at a more or less early stage. Excep- tionally, another litter can follow and abort, but generally reproduction is interrupted until the be- ginning of good recrudescence of fruiting. Among the Didelphidae, P. opossum behaves more like an r-strategist in favoring rapid population in- crease in an unstable environment." Charles-Dominique (1983, p. FRANgAISE) 409, GUYANE Comparison of breeding conditions between Philander opossum and Caluromys philander of Guyane Fran^aise, two similar-sized species oc- cupying different ecological niches: Philander opossum I. Mainly terrestrial Caluromys philander 1. Almost entirely ar- boreal 2. Omnivorous but largely frugivorous 3. Food availability variable in under- growth where for- aging is done 4. Birth to weaning 68-75 days 5. Size weanling, 50- 75 g 6. Age at sexual ma- turity, 6 months 7. Up to three succes- sive litters in same year (third litter mostly pouch- aborted) 8. Behavior as r-strat- egist' 9. Nesting period, 8- 15 days 10. Litter size not more than 7 2. Mostly frugivo- rous, nectarivorous 3. Food availability less variable in up- per strata where foraging is done 4. Birth to weaning 110-125 days 5. Size weanling, 50- 75 g 6. Age at sexual ma- turity, almost 10 months 7. Two successive lit- ters in same year 8. Behavior more like that of /:-strategist^ 9. Nesting period, 30-45 days 10. Litter size not more than 7 ' r strategist: r-selected species, selection favoring a rapid rate of population increase. Typical of species that colonize short-lived environments or of species that un- dergo large fluctuation in population size. - k strategist: ^-selected species, selection producing superior competitive ability in stable, predictable envi- ronments in which rapid population growth is unimpor- tant as the population is maintained at or near the car- rying capacity of the habitat. Atramentowicz (1986a, p. 125, GUYANE FRAN- gAISE) During a study period from September 1978 through October 1979 in Cayenne, two successive litters were produced between September and June. Some females, however, produced three lit- ters in the same period. The difference is attrib- uted to an abundance of food and a shortened nesting period. In the event of a food shortage, milk production ceases. Death of lactating young starved at the nipple is termed pouch abortion. Average adult weight, 400 g. Atramentowicz (1986a, p. 123, GUYANE FRAN- gAISE) During a 26-month (1978-1982) investigation of didelphoid breeding, "animals were live trapped, marked and then released. Caluromys philander. Philander opossum, and Didelphis 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 Weight (g) Pouch Young Measurements' (mm) 16 February _ None — 2 March - 7 20.02 4 March - 7 nm^ 5 March - 7 nm 8 March - 6 nm 8 March - 6 nm 10 March - 7 24.0- 13 March - 3 43.02 17 March 355.0 5(2cJ,39) 44.8 29 March - 4 young, out of pouch (3d, 1 9) 107.0 31 March 420.2 5(M,4$) 35.2 31 March 402.4 5(3d, 29) 31.0 2 April 546.3 7(5d, 29) 63.8 2 April 674.5 6(4d,29) 65.5 2 April 361.8 6(26,49) 38.5 15 April - 7(4(5, 39) 48.0 15 April 521.3 7(36,49) 44.0 15 April 464.9 7(3c?,49) 47.1 24 April 352.1 6(46, 29) 57.0 2 July 256.3 7(2c?,59) 40.5 6 July 550.0 Lactating, no pouch young - 15 July 590.0 1(56, 19, 1?) 89.0 27 July 282.7 5 nm Head and body combined. "Probably crown-rump length." nm - no measurement. rous and insectivorous diet. The first species is arboreal, the others mainly terrestrial, but may be seen climbing to the canopy. The percentage of lactating females in the studied population has been compared to food supply in terms of the availability of fruiting food plants." "The collected data show a direct relation be- tween these two factors. Breeding occurs through- out the year but food scarcity may interrupt re- production either with no births or 'pouch-abor- tions' (death of pouch-young) by starvation for lack of milk production." Atramentowicz (1986b, p. 140, GUYANE FRAN- gAISE) Youngest lactating female, age 8 months. Pouch young 4.2 (2-7); 179 litters. Two births between September 1978 and Oc- tober 1979. Atramentowicz (1988, p. 55, GUYANE FRAN- gAISE) "Fruits are essential during lactation, the sugars they contain helping the females to cover their increased energy need during this period. On weaning, the inhibition of the ovarian cycle is re- moved and a new litter may be produced during the next 15 days. In fact, the ovarian cycle is not inhibited during gestation as in placental mam- mals, but by the suckling which begins after birth when the embryos attach themselves to the teats." Phillips and Jones (1969, NICARAGUA) Size, weight, length, and dates of pouch young of 22 litters are shown in Table 17. Dates range from 16 February to 27 July; pouch litter size, 6.05 (3-7) 21; sex ratios in 13 of the litters with 78 young was 41 (53%) 6 6, 37 (47%) $ $; av- erage head and body length per litter increased from 20.0 to 107.0 mm within above time frame. Biggers (1966, p. 264, NICARAGUA) Seventy-three live-trapped males had sperma- tozoa in testis and epididymis throughout the year; testis weights are lower from September to December inclusive than in remainder of year. Pouch young found January to October inclu- sive with peaks in January, May, and August. Mature animals obtained January to November inclusive. No males trapped in December. No dif- 68 FIELDIANA: ZOOLOGY ference in weight of testes of mature animals throughout the year. Discrepancies noted in paragraphs one and two above are confusing. Perhaps the mixture of data for Philander and Didelphis is the cause. Biggers refers readers to the original compilation of data. Enders (1935, p. 41 1, PANAMA) Only nonbreeding females captured January to April; starting July, "females with litters outnum- ber those without; litters of two, four, and five were observed." Enders (1966, p. 198, PANAMA) "Observations on 18 captured Philander indi- cate that their breeding season . . . begins with f onset of the dry season in February." I "The largest litter encountered was five." Fleming (1973, p. 444, Canal Zone, PANAMA) Females were reproductively active during June, July, and September through December in 1966 and during February through April and June in 1967. Females carrying pouch young or lactat- ing in October and November were anoestrus. Two, possibly more, litters are produced per sea- son. "Males of P. opossum are sexually mature at a body length of 230 millimeters or greater . . . adult males probably are sexually active through- out the year." Litter size 4.6 (2-7) 34; sex ratio 22 (45%) 6 6, 27 (55%) $9. Seba (1734, p. 57, SURINAME) Eight pouch young were counted. Seba and Miller (above) are the only authors to record eight pouch young. Maximum number in all other cases is seven, in agreement with number of nipples. Philander, unlike Didelphis and oth- ers, may not produce more young than can be suckled. Cannibalism Charles-Dominique (1983, p. 405; Cayenne, GUYANE FRANgAISE) Many cases of cannibalism have been observed in captivity of weaned young being eaten by other young and the mother. Density Charles-Dominique et al. (1981, p. 383, Cayenne, GUYANE FRANgAISE) 100-200 individuals per km-. Diet Temminck (1827, p. 38) It preys on small birds and insects but also eats fruit. In captivity it feeds on meat. Pelzeln (1883, p. Ill, BRASIL) Insects found in stomach. Goldman (1920, p. 52; PANAMA) "The stomach of one taken at Gatun was well filled with fragments of crabs. Stomach contents of several others at the same locality were filled with fragments of birds alone, or of birds includ- ing their feathers, and crabs intermixed." Fleming (1972, p. 623; Canal Zone, PANAMA) "Stomachs of the four-eyed opossum contained unidentified plant material, the pulp of Corozo oleifera nuts, insect and fresh water shrimp re- mains, murid rodent fur, and the tail of a young spiny rat (Proechimys semispinosus). One individ- ual was seen feeding on live bats caught in a mist net." Schomburgk (1840, p. 344; GUYANA) "They are very destructive to poultry and like- wise to fruit. They are often found on those sa- vannahs where the wild pine (Bromellia sp. ?) flourishes, to the fruit of which they appear to be partial." Enders (1935, p. 410; Canal Zone, PANAMA) They "are accused of being poultry- and bird- killers. ... In captivity they were more carnivo- rous than any of the other opossums observed. They ate meat of all kinds including ant-eater, car- casses of rodents, grasshoppers, and eggs in pref- erence to any kind of fruit, although they did eat banana, papaya, pineapple, and figs. One hungry individual, while fearful of the hand offering a ripe banana, nevertheless sniffed from a safe dis- tance while saliva dripped from its jaws." JHERSHKOVrrZ: PHILANDER 69 Dalquest (1953, p. 19; eastern San Luis Potosi, MEXICO) ''One animal became entangled when it at- tempted to climb a net to obtain an entrapped bat, others were taken in traps baited with decayed meat, and two were shot while they were feeding on small wild figs. Remains of insects were usu- ally present in the stomachs of the specimens taken." Anthony in Goodwin (1934, p. 5; GUATEMALA) "I was told that these little opossums are often killed in the sugar house [trapiche] and that they are quite fond of sweets. ... At Finca Capres, several times I surprised them at night, feeding on the ripe [bananas]. ... At Chipore . . . one was attracted by bait composed chiefly of peanut but- ter and rolled oats." Hall and Dalquest (1963, p. 198; Veracruz, MEX- ICO) "It has been seen feeding on sweet-lemons, jobo plums, and the fruit of the Chico Zapote (Sa- pote achras) source of chewing gum." One opos- sum "was seen at the base of a large, hollow fig tree. The upper part of the hollow in the tree served as a retreat for a colony of the large fruit bat, Artibeus jamaicensis. Bats were bringing small green figs into the hollow and feeding on them. Parts of the fruit, varying in size from al- most whole figs to mere shreds, were dropped by the bats. The four-eyed opossum was feeding on these bits of figs." The opossum "followed our trap lines, eating mice and other small mammals that had been cap- tured. These opossums were easily trapped by us- ing flesh, preferably much decayed, for bait." Dubost (in Charles-Dominique, 1971, p. 197, GUYANE FRANgAISE) Philander eats tree exudate, and like the pri- mate Cebuella (Callitrichidae) or Microcebus (Le- muridae) uses its teeth to reopen healed openings in tree trunks to renew the flow of sap. Charles-Dominique (1983, pp. 397, 398; Cay- enne, GUYANE FRAN^AISE) The diet is opportunistic, consisting of fruits, flowers, nectaries, insects, earthworms, small ver- tebrates, and carrion. Sympatric D. marsupialis subsists on about the same; Marmosa murina pre- fers fruits but takes earthworms and small verte- brates. Marmosa cinerea [= Micoureus demer- arae] and C. philander are vegetarian. "The period of food scarcity determined both by records of fruit and insect abundance and also by body weight changes in the marsupials begins in May, is most critical during June-July-August, and ends in September. Data collected in 1976, and from 1976 to 1982 by the Institut Pasteur de Cayenne (variation in weight and reproductive condition of marsupials) suggest that this period of food scarcity is a regular phenomenon, proba- bly correlated with seasonal variations in rain- fafl." Charles-Dominique et al. (1981, p. 376; Cayenne, GUYANE FRAN^AISE) "Eighty percent of stomach contents consisted of a great variety of prey. During the rainy season, earthworms, some of which are as much as 80 cm long, forced above ground, made up a large part of the diet. "Fallen fruit consumed were mainly Attalea re- gia, Virola melinonii, Virola sebifera, Virola sur- inamensis, Ocotea puberula, Richardella macro- phylla, Ficus spp., Cordia exaltata, Simarouba amara, Protium heptaphyllum. In general the pulp is eaten with only the smallest seeds {Ficus, etc.) swallowed, the larger ones spit out. The ground level flowering of the Balanophoracea from Jan- uary to June is actively sought for the nectar. Los- es weight during dry season (September-Novem- ber in the years 1978, 1979) when fruit is scarce, fatten when fruit is abundant." Tyndale-Biscoe (1980, p. 716; COLOMBIA) "They would eat fruit and vegetables provided as well as meat; chicken bones were held in the hands and the ends chewed." Atramentowicz (1986a, p. 125; Cayenne, GUY- ANE FRANgAISE) Eats ripe pulpy fruit, insects, and small prey such as earthworms, frogs, etc. Atramentowicz (1988, p. 48; Cayenne, GUYANE FRAN^AISE) Stomach contents of Philander opossum consist of 50% fruit with the remainder invertebrates, small vertebrates, and carrion. A preference is shown for ripe fruit fallen to the ground. The fruit consumed represented 44 species in 21 families. 70 FIELDIANA: ZOOLOGY It was characterized by a fleshy pulp with a high percentage of water The color may be bright or dull, size large or small, shape variable. The pulp may be rich in sugar or lipids but poor in nitrogen, which the animal hnds in animal prey. Display Hershkovitz (field notes) The supraorbital spots are conspicuous at night, at least to the sharp-eyed (see "Sleep," p. 77). Conspicuous supraorbital marking displayed by both nocturnal and diurnal animals may serve for recognition between consjjecifics. I find no differ- ence in markings between the sexes, and the signs are usually absent or faint in juvenals. Enemies, Predators Most likely predators are the eyra cat {Her- pailurus jagouaroundi) and smaller spotted cats, large mustelids such as the tayra {Eira barbara) and grison (Galictis vittata), foxes, large owls, and snakes. Husson (1978, p. 26) found remains of a Philander opossum in the stomach of a gar- den tree boa (Corallis enhydhs) in Suriname. Schomburgk (1840, p. 344) reported that "they are sometimes eaten by the Creoles and Indians [of Guyana], but as they have a rank and disa- greeable smell I doubt if they would prove pal- atable to us." Wilson (1970, p. 386) reported an encounter in Panam^ between Didelphis marsupialis and Phi- lander opossum with the former overcoming and devouring skin, flesh, bones, and brain of the lat- ter (see details under "Aggression, Reaction, De- fense"). yellow under the hunting lamps while the eyes of [Metachirus] show red." Hershkovitz (field notes) The eyes of Philander, like those of other di- delphoids and Metachirus, usually glow bright or- ange when reflecting the beam of a battery-pow- ered flashlight. There is individual variation in tone of eye shine depending on the amount of environmental light, intensity of directed light, and angle of reflection. These factors and the un- controlled conditions make it difficult to find sig- nificant differences in color of didelphid eye shine. More important for identification is dis- tance between the eyes and animal gait as seen by the reflected light. Fat Storage Hall and Dalquest (1963, p. 198; Veracruz, MEX- ICO) "In the lowlands these opossums seldom were fat, but in the highlands, at 5000 feet elevation and higher, in winter, they had a deep layer of yellow [!] fat immediately beneath the skin." Foraging (Table 18) Charles-Dominique et al. (1981; Cayenne, GUY- ANE FRANgAISE) Of a total of 294 individuals live-trapped or tracked by radio, 1 85 were located from 0 to 6 m above ground in the following situations: on ground 70%; fallen logs 7%; small tree trunks 2%; branches 6%; lianas or vines 15%. Eye Shine Anthony in Goodwin ( 1 934, p. 5; GUATEMALA) "In the beams of the carbide hunting lamps, their eyes were extremely bright, gleaming white, like electric lamps, easily seen at one hundred yards or more." Enders (1935, p. 412; Canal Zone, PANAMA) The eyes of [Philander] show up as a reddish Charles-Dominique et al. (1981, p. 343; Cayenne, GUYANE FRANgAISE) Nearly always forages on the ground; of 100 individuals live-trapped only three were captured above ground. Charles-Dominique (1983, GUYANE FRANgAISE) p. 397; Cayenne, See Table 1 8 for body mass and above-ground 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 Mamiosa cinerea {- Micoureus demerarae] Caluromys philander Philander opossum Didelphis marsupialis 40 80 300 400 1,000 Low (shrub) High High Low (essentially ground level) Low (may climb to feed on fruit) Grooming Hershkovitz (field notes, January 30, 1941; north- ern COLOMBIA) A captive washed face with hands mouselike; groomed remainder of body with tongue catlike. Charles-Dominique (1983, p. 407; Cayenne, GUYANE FRANgAISE) "We have never observed any allogrooming between [caged] adults, even during mating peri- ods." Habitat Hall and Dalquest (1963, p. 196; Veracruz, MEX- ICO) "... ranges throughout the tropics of Veracruz. At the extreme upper edge of the upper humid division of the Tropical Life-zone, it lives along cold, clear streams at the edge of the oak belt. Lower down but still in the upper humid division, it was found along rivers and streams that flowed through dense jungle, where the tall, broad-leafed trees were thickly hung with orchids, vines, moss- es and bromeliads. The four-eyed opossum was found living in the thickets bordering the broad rivers of the coastal plain, in the arid division of the Tropical Life-zone, and along the marshy shores of rivers and streams of the lower humid division of the Tropical Life-zone, in the southern part of the state. "Most of our specimens were taken on the very shores of rivers or streams . . . however . . . the species is not confined to such habitat . . . seven kilometers west of Potero, workers discovered a family of four young animals in a field of sugar cane, several kilometers from the nearest water at that time of the year. At Jimba, 350 feet elevation, in southern Veracruz, a four-eyed opossum was taken from a tree on a hillside fully three kilo- meters from the nearest water. These records are unusual, however. Against them are nearly 30 re- cords from in and near water." Dalquest (1953, p. 19; eastern San Luis Potosi, MEXICO) "Found only in the tropics of the eastern part of the state . . . they are taken in dense vegetation growing along the shores of streams and rivers at elevations between 400 and 12,000 feet [more likely 1200 feet]. At El Salto the water is cold and flows swiftly between rocky canyon walls; opossums were found in thickets of ferns, vines, and low, woody plants. Near Huichihuayan, where the water is warmer and sluggish, opos- sums were taken in thickets of thorny bamboo." Goodwin (1946, p. 284; COSTA RICA) "Frequenting forested country from sea level up to about 4000 feet, he is most common at low elevations." Davis (1947, p. 2; Teresopolis, BRASIL) "Found more commonly in moist situations al- though individuals may wander through nearly any kind of vegetation." Enders (1935, p. 410; Barro Colorado Island, PANAMA) "They are not common on the Island for while they enter traps without hesitation none were taken; but many were captured at Alhajuela in habitats similar to those on the Island . . . why they are not abundant on the Island is a puzzle." Schomburgk (1848, p. 777; GUYANA) "It lives mainly in the coastal forests bordering the plantation. During the day it sleeps most of the time in its haunt under tree roots or in hollow trees." 72 FIELDIANA: ZOOLOGY Natterer (in Pelzeln, 1883, p. Ill; southeastern BRASIL) ". . . Fashions its nest from leaves on low trees." Miller (in J. A. Allen. 1916c, p. 589; Mato Gros- so, BRASIL) ". . . is found in deep forests. I have never known it to venture near houses for the purpose of robbing hen-roosts and nests like its larger rel- ative [Didelphis marsupialis], although it may do so." Handley (1976, p. 8; VENEZUELA) Lowlands of western and southern Venezuela, 46 specimens: on the ground, 98%; on a log, 2%; near streams and other moist areas, 100%; ever- green forest, 91%; orchards, croplands, yards, open areas, 9%; elevation 24-324 m. Hall and Dalquest (1963, p. 198; Vera Cruz, MEXICO) "... are usually seen or trapped on the ground, but are sometimes seen in trees." They prefer the vicinity of streams and seem to be as much at home in rivers as on their banks. Enders (1935, pp. 410, 411; Barro Colorado Is- land, Canal Zone, PANAMA) None "were taken in the Island but one was observed high up on a limb on Zetek Trail. . . . While a good climber [it] was usually encoun- tered upon the ground, on or under logs. It is probably more terrestrial in habit than any of the 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 10 to 16 times greater in secondary than in pri- mary forest. Husson (1978, p. 26; SURINAME) Some of the.se opossums were found under the floor of a home at Lelydorp. Atramentowicz (1986a, p. 123; Cayenne, GUY- ANE FRANgAISE) Philander opossum is mainly terrestrial "but may be seen climbing to the canopy" in the sec- ond growth forest. Da Fonseca and Kierulff (1989, p. 118; Atlantic forest, Minas Gerais, BRASIL) "Fourteen individuals were caught during the [17-month] course of this study. The occurrence of this large didelphid is apparently tied to the presence of standing or running water. ... As only a few transects occurred close to streams, this may explain the low trapping success for this species." "The species also proved to be primarily ter- restrial with only 17% of captures in arboreal traps, and 7% of individuals climbed trees after being released." Unpublished (VENEZUELA) Capture sites recorded by members of the Smithsonian Venezuelan Project (SVP) on field tags of captured specimens are summarized be- low: A^ Live-trapped in banana 1 grove Snap-trapped on forest floor 2 Live-trapped on ground Snap-trapped in river Snap-trapped in tree Shot in tree in forest 9 1 1 1 Shot in hollow tree near 1 river Shot in tree alongside river 11 Total 27 All Philander andersoni were taken in the Ter- ritorio Federal Amazonas. The P. opossum were captured in the state of Bolivar, where different ecological conditions prevail (Handley, 1976). It seems that P. andersoni is at least as arboreal as it is terrestrial, with perhaps a preference for trees HERSHKOVITZ: PHILANDER 73 overhanging rivers. This type of habitat restricts enemy attacks to the forested side of the river and provides the opossum with the opportunity of es- caping by diving into the water. The musicrat-like dorsal pelage of P. andersoni mcilhennyi and many individuals of P. andersoni andersoni seems to be better adapted than that of P. opossum for aquatic life. Where the two species occur to- gether, as in Balta and along the Rio Ucayali in Peru, P. andersoni may be more arboreal-aquatic than terrestrial, whereas P. opossum may be more terrestrial-aquatic than arboreal. Home Range and Territoriality Miles etal. (1981a, BRASIL) A small, transparent plastic spool, 6 cm long with 2.2 mm internal and 2.4 mm external diam- 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 was attached to the capture trap or nearby vege- tation. As the released animal traveled through the night the thread unwound. The thread tracked dur- ing the day led to the diurnal nest or refuge. Dis- tance traveled was calculated from the weight of the spool before release of the animal and after recapture. Of 21 released animals, 16 were retrieved. The calculated distance for 15 retrievals averaged 438 m (93-1000 m). This compared with a mean of 801 m (51-2450 m) for Didelphis marsupialis. Charles-Dominique (1983, p. 401; Cayenne, GUYANE FRANgAISE) Philander explores about 25,000 m- during the night. "By systematically trapping, marking and releas- ing we have observed regular loss of individuals, compensated by immigrants from other areas (ju- veniles and adults of both sexes). Trapping was con- ducted using Tomahawk and Sherman traps placed on the ground and special 'home-made' traps set at 15 to 20 m in trees; traps were baited with bananas . . . The following data concern new-arrivals [in an over 20-ha study area] from May to October 1979, after 8 months of regular trapping and marking": May, eight immigrants; June, 2; July, 1 ; August, 3; September, 4; October, 3. "Migration seems to be more important in P. opossum than in C. philander — after one year, the entire population of P. opossum had been replaced by new individuals, but only one-third of the orig- inal C. philander population was replaced." Atramentowicz (1986b, p. 145; GUYANE FRAN- gAISE) Didelphoids are solitary, not territorial, and many individuals are sedentary. Duration of resi- dence in study area is compared with Caluromys philander. Philander opossum, and Didelphis marsupialis (Table 19). Charles-Dominique (1983, p. 418) All American opossums are nocturnal and sol- itary except during mating and maternal care of young. Territory is not defended. Home ranges overlap between many individuals, male and fe- male, but with close contact avoided. There are no social bonds between adults apart from short copulatory periods; there is no allogrooming. Hydrotropism (see also Habitat) Hershkovitz (1962, field notes; SURINAME) A female with 3 pouch young taken live 20 January 1962 from a snap trap in Lelydorpplan was caged for observation. The animal never re- covered from the effects of the blow on the head received from the trap. She slept most of the time and when awake walked slowly with poor coor- dination. She permitted herself to be petted and seemed to enjoy it, but the docility probably owed . to her weakened condition. She ate the insects fed to her and seemed to require large amounts of| water. A Marmosa murina held captive at the same time derived all its water from fruit. The Philander escaped twice from her cage. On each occasion she sought out the nearest dark, cool, moist refuge. The first time she curled up in an empty but still-wet canvas bucket in the kitch- en. The second time, after I had moved to a frame house in La Poule, she snuggled into a dark, wet corner behind a primitive flush toilet. Fonseca and Kierulff (1989, p. 118; Atlantic for- est, Minas Gerais, BRASIL) "The occurrence is apparently tied to the pres-' ence of standing or running water." Tyndale-Biscoe (1980, p. 716; COLOMBIA) J "All these animals were caught in second 74 FIELDIANA: ZOOLOGY r growth forest close to streams, in several caves less than a meter from water. It was thus interest- ing to observe that a pair in captivity would in- variably defecate into the water container rather than elsewhere in the enclosure." Locomotion Dalquest (1953, p. 19; eastern San Luis Potosf, MEXICO) ^ They "are swift in their actions. For the most 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- ing into the water of the Rio Naranjos and swim- ming away beneath the surface." Crespo (1950, p. 6; Misiones, ARGENTINA) They "are excellent swimmers, water appar- ently being their preferred habitat. I have watched them at night swimming swiftly even upstream against the current in rivers such as the Uruguai." Hall and Dalquest (1963, pp. 197, 198; MEXICO) '"Philander is quick and active. Trapped indi- viduals are able to jump about and twirl in sur- prising fashion. . . ." The animal "is an agile climber and a skillful swimmer." "Along the Rio Atoyac several four-eyed opos- sums were taken in a trap set beneath the water level, at the base of a cut-bank. They could have reached the trap only by swimming." "... A four-eyed opossum was seen just before midnight, running swiftly over the larger, rounded boulders (six to 18 inches in diameter) along the river bank. When frightened [by us] the animal turned [from the river's bank] and made a smooth, clean dive into the swift water, and as it did not reappear, must have swum away underwater." Hershkovitz (field notes) The usual didelphid locomotor pattern on the ground is a measured walking gait, and in climb- ing an alternating movement of the grasping hands and feet with support of the clinging, pre- hensile tail. The principal running gait on or above ground is a trot. I have never seen a didel- phid leap or bound except in rapid escape. In de- scending a tree trunk the animal will proceed head first or simply drop to the ground. Hershkovitz (1941, field notes; northern COLOM- BIA) The prehensile tail was used for swinging the body from limb to limb, the hands for 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 grasp the limb above with its hands. Carrying an opossum by its tail can be hazardous. Marmosids habitually use the tail for climbing. When attacking live prey, the opossum springs for throat or head with lightning speed. Longevity Farris(1950, p. 258) Life span in captivity, 3 years, 6 months. Atramentowicz (1986b, p. 140; GUYANE FRAN- gAISE) Based on tooth wear, longevity of wild Philan- der opossum and Didelphis marsupialis was cal- culated as 2.5 years; of Caluromys philander, as 3.5 years. Captive Caluromys philander (Brunoy, France) lived to 6 years. Maximum life expectan- cy of Didelphis marsupialis was said to be less than 36 months. Maternal Care Seba(1734, p. 57) "The mother does not remove the young from her pouch before they are old enough to enjoy the light of day. When the time is right she retreats to a lookout to assure the safety of her young. She then opens the pouch and permits the offspring to emerge into the sunlight and play with her. At the slightest suspicion of danger she calls her young with a warning cry that sounds tik, tik. tik. The young respond immediately by returning to the pouch whereupon the mother runs to a hiding place." Comment — Seba actually beheld the mother and her brood at one time or another. The fanciful description of their relationship (translation), however, must have been confected long after the supposed facts. Didelphids are nocturnal, but in captivity, es- pecially as household pets, as Seba's opossums HERSHKOVITZ: PHILANDER 75 must have been, they generally adjust to the hours they are fed. Nest Goldman (1920, p. 52; PANAMA) "A nest of one of these opossums was found three feet from the ground on a fallen log. The log lay in the dense thicket of an old clearing and was heavily overhung with vines and bushes. The nest, globular in form and about a foot in diam- eter, was placed in a well hidden spot among the vines. It was made entirely of the banana-like leaves of a native plant rather neatly laid together. The opening at one end faced outward along the log. . . . The nest cavity was clean and about the size of the animal's body." Carriker (in J. A. Allen, 1911, p. 247; Yuruan, VENEZUELA) "The female with a litter of young was taken out of the hollow stump in a grass field in the [forest] clearing." Hall and Dalquest (1963, p. 197; Veracruz, MEX- ICO) "Several four-eyed opossums were taken in a trail leading from a dense thicket ... to a stream ten feet away. ... A hole, about five inches in diameter that led downward beneath the roots of a tree was discovered near the center of the thick- et. This seemed to be the home of at least one of the opossums." Two "were found in a nest that they had con- structed in the palm thatch of the roofs of aban- doned houses. These nests consisted of a handful of dry leaves, pushed in between the layers of palm fronds. From outside, a distinct spherical or oval lump in the thatch marked the site of the nest. Inside the house we could see no trace of the nests." A nest "was found in a cavity in the side of a piece of tree trunk, 15 inches in diameter and three feet long, that was suspended in the air by vines. . . . The nest was of dry leaves, about 1 1 inches deep and seven inches in diameter." One animal "was shot from the large hollow in the side of a giant 'ligaron' [higaron?] tree. This tree was fully 1 2 feet in diameter at waist height, and contained a hollow about 60 feet high and five feet in diameter." Dalquest (1953, p. 19; eastern San Luis Potosi, MEXICO) "A burrow near Huichihuayan was in muddy soil beneath a clump of thorny bamboo, and an- other was beneath a decayed log, nearly concealed by vines and succulent plants. This burrow was so shallow that when the log was rolled aside the entire burrow was exposed. A nest at the end of the branched burrow consisted of a formless mass of dry leaves a foot in diameter. Two feet away there was an opening to the dense vegetation be- side the log. Three feet farther on, the burrow emerged from under the log; and a trough like trail led through the vegetation to the river, twenty feet away." Murie (1935, p. 16; GUATEMALA) "In the edges of the pine ridge, especially near the streams, were numerous well worn trails about five inches wide, which were apparently made by these opossums. Many of the trails led from one ground burrow to another. At the end of one of the burrows about fifteen inches from the en- trance, I found a nest ten inches in diameter, com- posed of dry sedges." Enders (1966, p. 201; PANAMA) After the young are born the female moves about more. Unlike habits during the rest of the year (Enders, 1935, p. 397), they frequent the same nest. Later, when the young are able to de- tach themselves and so leave the mother, young may continue to use the nest. Husson (1978, p. 26; SURINAME) Nest with female and seven young found under a tree. Nothing more is said. Miles et al. (1981a, p. 341; BRASIL) "Nests of the opossum were often found 8-10 m above the forest floor in hollow trees or as open nests in tree forks; some nests were, however, ter- restrial, in cavities besides buttressed tree roots." ! Farris (1950, p. 259; PANAMA) i > "The animals usually were found in their nests i in palm trees, at the branching and in the leaves I about 5 feet off the ground where conditions were j relatively dry." 76 FIELDIANA: ZOOLOGY Population Density Charles-Dominique et al. (1981, p. 383; Cayenne, GUYANE FRANgAISE) Philander opossum was one of the most abun- dant mammals on the island. Its biomass, based on 199 observations within a square kilometer, was calculated as between 100 and 200 individ- uals, with a biomass between 40 and 60 kilograms in the same area. The biomass of each of the five other marsupials of the same area was 25 to 50 kg for D. marsupialis, 30 to 60 kg for C. philan- der, 2 to 8 kg for Marmosa cinerea [Micoureus demerarae], and 0.7 to 3.5 for M. murina. Of the six marsupial species inhabiting the is- land. Philander, Didelphis, Caluromys, and M. murina were most abundant. Marmosa cinerea [ = Micoureus demerarae] were moderately abun- dant; Monodelphis brevicaudata, uncommon. Cerqueira et al. (1993, p. 513; Restinga de Barra de Maricd, Rio de Janeiro, 22°57'S, 42°5rw, an impoverished small animal community, At- lantic forest, BRASIL) Average density 1.91/ha; population main- tained mostly by recruitment. Atramentowicz (1986b, p. 144; GUYANE FRAN- gAISE) Density per square kilometer based on obser- vations from September 1978 to October 1979 was 137 P. opossum, 143 C. philander, and 45 D. marsupialis. Population Dynamics ed for comparison. These sympatric Didelphids are nocturnal, with a basically frugivorous and in- sectivorous diet. The three main species have high densities, high reproductive rates, short life-spans, and rapid population turnover. There are some dif- ferences in these parameters, mainly between the arboreal species, C. philander, and both terrestrial species, D. marsupialis and P. opossum. Although all have few social interactions and show a lack of territoriality, C. philander appear to have a lon- ger life-span, a lower reproductive rate and a more sedentary population. M. cinerea [Micou- reus demerarae] were very rare in this area, whereas M[armosa]. murina show large density variation." The well-known tendency of American marsu- pials to enter baited traps, usually the same ones, time and again ensures a high degree of accuracy in estimating their duration of residence in an area. In general, both sexes of P. and C. philander are highly sedentary. Didelphis marsupialis, on the other hand, tends to wander, particularly the males (Table 19). Posture Enders (1935, p. 412; Canal Zone, PANAMA) Philander "used the tripod [sitting] posture more frequently than Metachirus although the tail of the latter is a very effective appendage." Hershkovitz (1941, field notes; northern COLOM- BIA) Sitting was tripodal, the body supported by hind limbs and tail. Atramentowicz (1986b, GUYANE FRAN^AISE) sleep "Five species of Didelphid marsupials were trapped live, marked, and released in a secondary forest in French Guiana during a 26-mo field study. Data on body weight, body length, dental stage, and reproductive state of females were col- lected. A total of 85 1 individuals, including 372 pouch-young, were monitored in 2273 captures. This abundance of data we made led us to a com- parative analysis of the population dynamics of three species: Didelphis marsupialis. Philander opossum, and Caluromys philander, which rep- resented 94 percent of the whole captures (Table 19). Some data on the two other species, Mar- mosa murina and M[icoureus]. cinerea, were add- Temminck (1827, p. 38) It sleeps during the day rolled up into a ball. Its breathing is like that of a ferret, and it leaves its hiding place only at night. Schomburgk (1840, 348; GUYANA) They sleep during the day under tree roots or in hollow trees and hunt at night. Hershkovitz (1941, field notes; northern COLOM- BIA) The animal sleeps rolled up on its side or ex- HERSHKOVITZ: PHILANDER m 77 ,Vi •s •S &. a ^ \^ «j S •2 ■« ^ ^ :s Q V u C !2 s *t« t« v o 0^ §" •s i s ■« _o 5 :§ n u i^ s O, Q h. ^ ■« s :§ is Ci, 5r 5^ g * I. s "3 U o r~ O w-i o — m (N fN (N rf 0>r) — (N — sd ^^ m — g ^ §3 „• _• rn >. >. 00— 'inooTtr-oovo SL, Tt ^ Tt VO \0 vO "^ sp O; (N >> >> — (N r- CN (T) (N in — •— (N — > ON IT) 8 CJ — o V 2-0 « C8 D 3 •:= -a -P isX) 00 00 Jl C C C c w is s Is •« c v; c/3 s/; •— O (U u (U 0/5 -a i_ u. u. 1> • — CS « CS E 3 3 3 3 •o -a -O g '> '> |> -^ "5 "o "5 =« — £ £ S tended backside up. The closed eyes are not vis- ible, but the whitish supraorbital spots give the animal an appearance of being awake with eyes widely opened. Sociability Hall and Dalquest (1963, p. 179; Veracruz, MEX- ICO) "On a few occasions, two four-eyed opossums were seen as close together as 50 feet, but oth- erwise they were solitary." Charles-Dominique (1983, pp. 405-407; GUY- ANE FRAN^AISE) "Didelphids are always solitary day and night. During development immediately after the young have released the nipples, they remain for a short period in a nest (8 to 15 days for P. opossum and 30 to 45 days for C[aluromys]. philander). During the first days (the first weeks for C. philander), the young maintain a certain cohesion between themselves (they often grasp each other) and ex- plore the surroundings of the nest. This gregarious behavior indicates the persistence of a need for body contact, directed indiscriminately towards the litter mates as well as towards the mother. Sometimes one or several young of the litter cling to the mother's fur during nocturnal activity. . . . Individual recognition may not exist between lac- tating females and young. In the middle of the night, and again at dawn, the mother returns to the nest for 2 to 3 h. This cohesive behavior dis- appears after weaning, mother and young becom- ing indifferrent or aggressive when they meet. "We observed no durable social bonds between adult individuals. When two animals met (for ex- ample, in a fruiting tree), they often threatened each other with an aggressive vocalization and then continued their individual activities." Charles-Dominique (1983, pp. 406, 407; Cay- enne, GUYANE FRAN^AISE) A dominance hierarchy or peck order does not ! exist among didelphoids. "Nineteen adult four- eyed opossums (10 females; 9 males) habitually | fed on a 1 m high platform baited daily with ba- j nanas. Animals were identified by collars (radio- i transmitters and color bands) and only had access I to the platform via a single branch. Observations under weak lighting were made from a distance 78 FIELDIANA: ZOOLOGY of 3 m. Conflicts were observed principally dur- ing the first 2 hr of the night when the four-eyed opossums came to take the first meal (12 evenings of observation). Usually an animal which had not 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- changed (hissing, open mouth, and start of chase). No rank order was observed and an individual (male or female) chased by another (male or fe- male) one night could chase it in turn the follow- ing night depending on the situation (degree of hunger)." Spontaneous Bleeding Hershkovitz (1941, field notes; northern COLOM- BIA) I An adult female P. opossum was clubbed by a woodsman and seized as she jumped from its nest in the felled tree. The animal was brought live to my camp, where it recovered without signs of in- jury. When irritated, however, it bled spontane- ously from the tips of the fingers, the tail, and the nose. I had never witnessed or known of this phe- nomenon before. Stress .Hunsaker and Shupe (1977, p. 302) 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 animal is frightened and can be considered a sub- mission posture." The coiled tail reduces exposure from seizure by a predator. Tail Transport Hunsaker and Shupe (1977, p. 301) "Tail coiling has been observed to facilitate carrying nesting material in Didelphis virginiana, Monodelphis domestica, Caluromys derbianus and Marmosa robinsoni. The fact that more spe- cies have not been observed to do so is probably a function of inadequate observations." Philander does coil its tail in the manner described and may indeed do so for transport of building material. According to Layne (1951, p. 464), use of the tail for transport by Didelphis was first described and illustrated by Pray (1921, p. 109). Layne further elaborated on the subject. Taming Hershkovitz (1941, field notes; northern COLOM- BIA) The opossum captured alive after being struck with a machete by a woodsman was leashed and 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 the mouth open. Within a few days, however, the animal became tame and permitted itself to be handled and stroked and seemed to enjoy climb- ing onto my shoulder and head. Thermoregulation Enders and Davis (1936, p. 165; Canal Zone, PANAMA) Body 35.4, environment 26.1 (afternoon). Miller (in J. A. Allen, 1916c, p. 589; Mato Gros- so, BRASIL) "As a general rule opossums cannot stand great heat and will soon die if left exposed to the direct rays of the tropical sun." Vocalization Charles-Dominique (1983, pp. 407, 411, 412; Cayenne, GUYANE FRAN^AISE) "Two types of calls by adults are the weak sex- ual call (rarely by the male) and an agressive call by both sexes. "The young begin emitting very short high- pitched "clicks" about one month before releas- ing the nipples, especially when the mother licks the young attached to the nipples in her pouch, or when the young are experimentally pulled off the nipples. "These clicks progressively disappear when the young release the nipple and the first 'hiss' — as- sociated with the typical posture (opened mouth) — is emitted very early in conditions of fear or when pulled off the nipple." HERSHKOVITZ: PHILANDER 79 In the case of C. philander the clicking sound was particularly loud, audible at 200 m. The same sound, but muted, was emitted by Marmosa mu- rina, Monodelphis hrevicaudata, and D. marsu- picdis. The clicking sound has also been recorded by Reynolds (1952, p. 235) for D. virginiana as be- ing most pronounced during breeding. Kirsh (1979, p. 392) heard the sound from Caenolestes fuliginosus; Thrasher et al. (1971), from Marmosa robinsoni. Bruce Patterson (personal communi- cation) heard Metachirus nudicaudatus clicking as a signal of aggression. Apparently all New World marsupials are ca- pable of three types of calls: the standard hiss, usually uttered when standing erect or tripodally in an aggressive posture, is usually fear stimulat- ed; a mouselike squeak, which may be a sexual call; and the click, which is emitted under various conditions. Weight Tyndale-Biscoe (1980; P. opossum— COLOM- BIA) Weights of adults (g): ^6 6, 416.9 ± 132.1 (230-675); 49 $, 323.8 ± 65.2 (275-420). Alphabetical List of South American Collecting Localities of Philander with Key Numbers to Distribution Map (Fig. 22) and Gazetteer (p. 85) See Hall and Kelson (1959) for Middle Amer- ican portion of range. Acanana; Territorio Federal Amazonas, Venezuela (107) Agua Limpa; Brasilia, Distrito Federal, Brasil (175c) Agua Viva; Trujillo, Venezuela (98a) Albina; Marowijne, Suriname (128) Alem Parafba; Minas Gerais, Brasil (182) Alguacil; Zulia, Venezuela (97a) Altamira; Para, Brasil (151) Alto Bonito; Antioquia, Colombia (1) Alto da Serra; Sao Paulo, Brasil (202) Ananindeua; Para, Brasil (159) Anapolis; Goias, Brasil (174b) Apayacu; Ucayali, Peru (43) Aragar^as; Goias, Brasil (173) Arruda; El Beni, Bolivia (73) Arumateua; Para, Brasil (157) Ascencion de Guarayos; Santa Cruz, Bolivia (84) Asuncion; Central, Paraguay (91) Auara; Igarape; Amazonas, Brasil (144) Avanavero Falls; Nickerie, Suriname (119) Avanhandava (Lajeado); Sao Paulo, Brasil (195) Ayacucho (Ibanez); Santa Cruz, Bolivia (88) Bacaetava; Sao Paulo, Brasil (199) Bagado; Choco, Colombia (4) Bahia; Bahia, Brasil (172) Baiao; Para, Brasil (156) Balisa; Mato Grosso, Brasil (168d) Balta; Ucayali, Perii (55) Barra do Cartas; Mato Grosso, Brasil (168c) Barranquita; El Beni, Bolivia (73) Barrieira; Rio de Janeiro, Brasil (189) Barro Branco; Rio de Janeiro, Brasil (193) Baptista (Lago); Amazonas, Brasil (141) Belem; Para, Brasil (159) Belem-Brasflia (Rodovia); Para, Brasil (159) Belen; Territorio Federal Amazonas, Venezuela (106) Bellavista; Antioquia, Colombia (2) Bemberg (Puerto); Misiones, Argentina (95) Benfica; Minas Gerais, Brasil (181) 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) Boa Fe (Fazenda); Rio de Janeiro, Brasil (189) Boa Vista (Fazenda); Rio de Janeiro, Brasil (189) Boa Vista; Roraima, Brasil (134) Boca Colorado; Madre de Dios, Peril (65) Bom Jardim; Brasil; not located; many so-named throughout most states Boraceia (Ponta); Sao Paulo, Brasil (203) Boraceia, Rio Tiete; Sao Paulo, Brasil (196) Branco (Rio); Amapa, Brasil (133) Brasilia; Distrito Federal, Brasil (175c) Bucay; Guayas, Ecuador (28) Buck Hall; Essequibo Islands- West Demerara, Guyana (116) Buenaventura; Valle del Cauca, Colombia (10) Buenavista; Santa Cruz, Bolivia (86) Butanta; Sao Paulo, Brasil (200) Caceres; Mato Grosso, Brasil (168a) ! Cachavi (= Cachabi); Esmeraldas, Ecuador (18) Cadena (Hacienda); Cuzco, Peru (63) Caixa D'agua; Espirito Santo, Brasil (186) Call; Valle del Cauca, Colombia (11) Cameta; Para, Brasil (153) 80 FIELDIANA: ZOOLOGY 0 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) Camino Vilches; EI Beni, Bolivia (73) Campinho; Espirito Santo, Brasil (185) Caney; Meta, Colombia (16) Cantareira (Serra da); Sao Paulo, Brasil (2(X)) Canudos; Pard, Brasil (149) Caoni (Rio); Pichincha, Ecuador (22) Capibara; Territorio Federal Amazonas, Venezue- la(lll) Capim; Para, Brasil (164) ,Caracarai; Roraima, Brasil (135) Cardosa (Fazenda); Minas Gerais, Brasil (180) Carondelet; Esmeraldas, Ecuador ( 1 8) Casa Grande; Sao Paulo, Brasil (203) Catatumbo (Puerto); Zulia, Venezuela (97b) Cauca (Rfo); Valle del Cauca, Colombia (11) Cayenne; Cayenne, Guyane Fran^aise (129) Cebollal; Loja, Ecuador (30) Cerro Cucurito; Territorio Federal Amazonas, Venezuela (109b) Centinela; El Beni, Bolivia (73) Chaco; Argentina (94) HERSHKOVITZ: PHILANDER Chalimana; Bolivar, Venezuela (104c) Chanchamayo; Junin, Peru (61) Chapare (Rio); Santa Cruz, Bolivia (83) Chicosa; Ucayali, Peru (53) Chimbo (Puente); Guayas, Ecuador (28) Chimbo-Coco (Rios); Guayas, Ecuador (28) Chocolatal, Monte, El Beni, Bolivia (75) Chulumani; La Paz, Bolivia (67) Churuli; Zulia, Venezuela (97a) Clevia; Suriname, Suriname (125) Cocal; Cauca, Colombia (14) Coco (Rfo); Guayas, Ecuador (28) Colonia do Prata; Para, Brasil (163) Colonia Hansa; Santa Catarina, Brasil (209) Colorado (Boca); Madre de Dios, Peru (65) Colorado (Rio); Madre de Dios, Peru (65) Commewijne (River); Commewijne, Suriname (127) Comochatiba (= Comoxatiba) 17°06'S, 39°irw. Maximilian Wied-Neuwied, July 1816. Conceigao do Mato Dentro; Minas Gerais, Brasil (176) Condoto; Choco, Colombia (5) Copataza (Rio); Pastaza, Ecuador (37) Corumba; Mato Grosso do Sul, Brasil (170) Costao dos Engenhos; Sao Paulo, Brasil (201) Cotia; Sao Paulo, Brasil (200) Covaria (Rio); Boyaca, Colombia (15) Cucurito (Cerro); Territorio Federal Amazonas, Venezuela (109b) Culturutuin; Suriname, Suriname (125) Culumani; La Paz, Bolivia (67) Cumaria; Ucayali, Peru (52) Cumeria; Ucayali, Peru (52) Cumuruxotiba (= Comoxatiba = Comochatiba - q.v.) Curaray (Rio); Napo, Ecuador (39) Curaray (Boca); Ucayali, Peru (39) De Oro (Rio); Chaco, Argentina (94) De Oro (Rio); Zulia, Venezuela (97) Dividive; Trujillo, Venezuela (98a) Docampado (Rio); Choco, Colombia (7) Duida (Cerro); Territorio Federal Amazonas, Ven- ezuela (108) El Capricho (Finca); 38 km E Villavicencio, Meta, Colombia (16) El Carmen; El Beni, Bolivia (78) El Chiral; El Oro, Ecuador (29) El Dividive; Trujillo, Venezuela (98a) El Palmar; Bolivar, Venezuela (102a) El Palmar; Santa Cruz, Bolivia (88) Encontrados, Rio de Oro; Zulia, Venezuela (97b) Engenheiro Reeve; Espirito Santo, Brasil (188) Engenheiro Rive (see Engenheiro Reeve) Esmeralda (Fazenda); Minas Gerais, Brasil (178) Esmeralda; Territorio Federal Amazonas, Vene- zuela (109a) Esmeraldas; Esmeraldas, Ecuador (19) Estrada de Santarem-Cuiaba; Para, Brasil (148) Exaltacion; El Beni, Bolivia (74) Fazenda Boa Fe; Rio de Janeiro, Brasil (189) Fazenda Cardosa; Minas Gerais, Brasil (180) Fazenda da Floresta; Minas Gerais, Brasil (178) Flor do Prado; Para, Brasil (161) Floresta (Fazenda); Minas Gerais, Brasil (178) Fordlandia; Para, Brasil (147) Formosa; Goias, Brasil (174a) Fracran; Misiones, Argentina (96) Georgetown; Demerara-Mahaica, Guyana (114) Gradaus; Para, Brasil (152) Gualea; Pichincha, Ecuador (23) Guama; Para, Brasil (165) Guaquitas; Barinas, Venezuela (100b) Guayaquil; Guayas, Ecuador (27) Guayaramarin; El Beni, Bolivia (68) Giiiniquina, Territorio Federal Delta Amacuro, Venezuela (98d) Hacienda Cadena; Cuzco, Peru (63) Hamacas; Santa Cruz, Bolivia (88) Hansa; Santa Catarina, Brasil (209) Hato San Jose; Bolivar, Venezuela (103) Hondo (Rio); Caldas, Colombia (3) Huampami; Amazonas, Perii (56b) Huanhuachayo; Ayacucho, Peru (62) Hyde Park; Demerara-Mahaica, Guyana (115) Ibaiiez; Santa Cruz, Bolivia (88) Ibare (Boca); El Beni, Bolivia (79) Ichilo (Rio); Cochabamba, Bolivia (82) Igarape Agu; Para, Brasil (163) Igarape Tapereba; Para, Brasil (150) Ilha do Taiuna; Para, Brasil (153) Imperatriz; Amazonas, Brasil (140) Inafiez; Pasco, Peru (not located) "High ground between Chanchamayo and up- per Rfo Pachitea" (Hendee in Thomas, 1928a, Annals and Magazine of Natural His- tory, Series 10, 2:250). R. W. Hendee, June 1927, at 5000 ft Ipanema; Sao Paulo, Brasil (199) Ipeau-Apez; Para, Brasil (159) Ipitinga; Para, Brasil (160) Ipixuna; Amazonas, Brasil (138) Iporanga; Sao Paulo, Brasil (197) ' Iquitos; Ucayali, Peru (42) Iriteria; Para, Brasil (165) Itaituba; Para, Brasil (148) j Itatiaia (Parque Nacional de); Rio de Janeiro, Bra- j sil (193) 82 FIELDIANA: ZOOLOGY Itatiaia (Serra de); Sao Paulo, Brasil (204) Itaya; Ucayali, Peru (42) Itonama; El Beni, Bolivia (71) Joinville; Santa Catarina. Brasil (209) Juiz de Fora; Minas Gerais. Brasil (183) Juquia; Sao Paulo, Brasil (201) Kaiserberg Airstrip; Nickerie, Suriname (122) Kanuku Mountains; Upper Takutu-Upper Esse- quibo, Guyana (118) Km 19; Para. Brasil (148) Km 90; Pari Brasil (159) Km 216; Para, Brasil (148) La Blanquita; Apure, Venezuela (101a) La Boca, Rio Saija; Cauca, Colombia ( 1 3) La Lengueta; Barinas, Venezuela (100c) La Neblina; Territorio Federal Amazonas, Vene- zuela (112b) La Papaya; Manabi, Ecuador (not located) T. Mena, May 1942, at 50 m La Ponchera; Tdchira, Venezuela (101b) La Poule; Saramacca, Suriname (124) La Poza; Amazonas, Peru (56a) Lagarto Cocha; Ucayali, Peru (54) Lago do Baptista; Amazonas, Brasil (141) Lago Mirafjes; Ucayali, Peru (40) Lagoa Santa; Minas Gerais, Brasil (177) Lapango; Central, Paraguay (92) Las Bonitas; Barinas, Venezuela (100b) Lazaropolis do Prata; Para, Brasil (not located) Lelydorp; Suriname, Suriname (126) Lelydorpplan; Suriname, Suriname (126) Lim6n; Ecuador (not located) G. H. H. Tate, December 1923, at 1200 ft Loksie Hattie; Brokopondo, Suriname (123) Los Guires, Territorio Federal Delta Amacuro, Venezuela (98c) Macapd; Amapd, Brasil (132) Macieiras; Rio de Janeiro, Brasil (193) Madeira (Rio); Amazonas, Brasil (142) Magdalena; El Beni, Bolivia (76) Majagua (Carlo); Territorio Federal Amazonas, Venezuela (105) Makerie; Nickerie, Suriname (121) Maldonado (Puerto); Madre de Dios, Peru (66) Mamor^ (Rio); El Beni, Bolivia (70) Mamore (Rio); Santa Cruz, Bolivia (83) Mangaratiba; Rio de Janeiro. Brasil (192) Marabd; Pari, Brasil (158) Marco; Para, Brasil (159) Maripa; Bolivar, Venezuela (102b) Matipo (Rio); Minas Gerais, Brasil (178) Mato Grosso; Mato Grosso, Brasil (167) Mavaca; Territorio Federal Amazonas, Venezuela (110c) Mazagao; Amapd, Brasil (133) Mazagao; Para. Brasil (154) Mazan (Rio); Ucayali, Peru (40) Mercedes; Santa Cruz, Bolivia (85) Merey; Territorio Federal Amazonas, Venezuela (112a) M^rida; M^rida, Venezuela (99) Merredor; El Beni. Bolivia (69) Mindo; Pichincha, Ecuador (24) Miranda; Mato Grosso do Sul, Brasil (171a) Miraiies (Lago); Ucayali, Peru (40) Mocajuba; Para, Brasil (155) Montalvo; Pastaza, Ecuador (36) Monte Alegre; Sao Paulo, Brasil (198) Monte Alegre; Mato Grosso, Brasil (169) Monte Chocolatal (see San Ram6n); El Beni, Bo- livia (75) Monte Serrat; Rio de Janeiro, Brasil (193) Montes Claras; Minas Gerais, Brasil (178) Motatdn, 1 km E and 5 km NNE; Trujillo, Ven- ezuela (98a) Motatan, Rio; Trujillo, Venezuela (98a) Moyobamba; San Martin, Peru (57) Murutucu; Pard, Brasil (159) Napo (Puerto); Napo, Ecuador (33) Napo (Rio); Napo, Ecuador (33) Nauta; Ucayali, Perii (45) Navio (Serra do); Amapi, Brasil (131) Nickerie (River); Nickerie, Suriname (120) Novita (Rio), Choc6, Colombia (8) Novo Friburgo; Rio de Janeiro, Brasil (189) Nulita; Apure, Venezuela (101a) 6rgaos (Serra dos); Rio de Janeiro, Brasil (189) Oro (Rio de); Chaco, Argentina (94) Orosa; Ucayali, Peru (44) Ouanary (River); Ouanary, Guyane Fran^aise (130) Oxapampa; Pasco, Peru (60) Pacific Virology Field Camp; Valle del Cauca, Colombia (12) Palacios; El Beni, Bolivia (74) Palmeiras; Mato Grosso, Brasil (169) Papelon (Cerro); Monagas, Venezuela (98b) Para; Pari, Brasil (159) Pari (River); Suriname, Suriname (125) Paracai (Rio); Parana, Brasil (207) Paramaribo; Suriname. Suriname (125) Paramba; Imbabura, Ecuador (21) Parima (Sierra); Territorio Federal Amazonas, Venezuela (110b) Parintins; Amazonas, Brasil (139) Parintins (Serra de); Amazonas, Brasil (139) Parque Nacional de Brasilia; Distrito Federal, Brasil (175b) HERSHKOVITZ: PHILANDER 83 Parque Nacional de Itatiaia; Rio de Janeiro, Brasil (193) Passo Fundo; Rio Grande do Sul, Brasil (210) Pedra Branco; Rio de Janeiro, Brasil (194) Peixe-Boi; Para, Brasil (163) Pescado (Rio); Manabf, Ecuador (25) Pinas; El Oro, Ecuador (29) Pindo Yacu (Rio); Pastaza, Ecuador (38) Piquete; Sao Paulo, Brasil (205) Playa del Rio Base; Territorio Federal Amazonas, Venezuela (108) Pocao, Boa Vista; Roraima, Brasil (134) Pocao, Caracarai; Roraima, Brasil (135) Pocone; Mato Grosso, Brasil (168b) Ponta de Boraceia; Sao Paulo, Brasil (203) Porto Velho; Rondonia, Brasil (166) Pucallpa; Ucayali, Peru (51) Puente de Chimbo; Guayas, Ecuador (28) Puerto Asis; Putumayo, Colombia (17a) Puerto Bemberg; Misiones, Argentina (95) Puerto Caballo; El Beni, Bolivia (74) Puerto Catatumbo; Zulia, Venezuela (97b) Puerto Maldonado; Madre de Dios, Peru (66) Puerto Napo; Napo, Ecuador (33) Puerto Siles; El Beni, Bolivia (72) Punta Santa Ana; Loja, Ecuador (29) Quartel de Sacramento; Minas Gerais, Brasil (178) Quincemil; Cuzco, Peru (64) Quininde (Rio); Esmeraldas, Ecuador (20) Raposo (Rio); Valle del Cauca, Colombia (12) Restrepo; Meta, Colombia (16) Rijweg; Suriname, Suriname (125) Rio de Janeiro; Rio de Janeiro, Brasil (190) Rio de Oro; Chaco, Argentina (94) Rio de Oro; Manabi, Ecuador (26) Rio de Oro; Zulia, Venezuela (97b) Rio Frio; Valle del Cauca, Colombia (9) Rio Hondo; Caldas, Colombia (3) Rioja; San Martin, Peru (58) Ro^a Nova; Parana, Brasil (208) Rodeio; Rio de Janeiro, Brasil (191) Rodovia Belem-Brasflia; Para, Brasil (159) Rosarinho; Amazonas, Brasil (143) Rupununi (River); Rupununi, Guyana (117) Saija (Rio); Cauca, Colombia (13) Salesopolis; Sao Paulo, Brasil (203) Salobra; Mato Grosso do Sul, Brasil (171b) Salvador; Bahia, Brasil (172) Salvias; El Oro, Ecuador (29) San Ignacio de Moxos; El Beni, Bolivia (80) San Javier; Esmeraldas, Ecuador (18) San Jeronimo; Cuzco, Perii (63) San Jeronimo; Ucayali, Peru (49) San Joaquin; ElBeni, Bolivia (73) San Jose; Ayacucho, Peru (62) San Jose; (Hato); Bolivar, Venezuela (103) San Jose, Abajo; Napo, Ecuador (32) San Juan; Territorio Federal Amazonas, Venezue- la (105) San Martin de Turumban; Bolfvar, Venezuela (104a) San Pablo; El Beni, Bolivia (77) San Ramon; El Beni, Bolivia (75) San Ramon; Santa Cruz, Bolivia (89) Sando (Rio); Choco, Colombia (6) Santa Ana, Punta; Loja, Ecuador (29) Santa Cecilia; Napo, Ecuador (31) Santa Clara; Amazonas, Brasil (140) Santa Cruz de la Sierra; Santa Cruz, Bolivia (88) Santa Elena; Ucayali, Peru (46) Santa Luisa; Ucayali, Peru (41) Santa Maria; Para, Brasil (162) Santa Rosa; El Beni, Bolivia (72) Santa Rosa; Ucayali, Peru (54) Santa Rosita; Santa Cruz, Bolivia (87) Santa Teresa; Espirito Santo, Brasil (184) Santa Teresa; Mato Grosso do Sul, Brasil (170) Santo Antonio de Uayara; Amazonas, Brasil (145a) Santo Dios; El Beni, Bolivia (73) Santo Isidoro; Amazonas, Brasil (137) Santarem; Para, Brasil (148) Santarem-Cuiaba (Estrada); Para, Brasil (148) Sao Joao de Petropolis; Espirito Santo, Brasil (184) Sao Joao Marcos; Rio de Janeiro, Brasil (191) Sao Luis de Caceres; Mato Grosso, Brasil (168a) Sao Miguel do Guama; Para, Brasil (165) Sao Paulo; Sao Paulo, Brasil (200) Sao Sebastiao; Sao Paulo, Brasil (206) Sapitiba (see Sepetiba); Rio de Janeiro, Brasil (191) Sapucai; Paraquari, Paraguay (93) Sapucajuba; Para, Brasil (159) Sapucay; Paraguari, Paraguay (93) Sarayacu; Pastaza, Ecuador (35) Sarayacu; Ucayali, Peru (48) Seboyal; Loja, Ecuador (30) Sepetiba; Rio de Janeiro, Brasil (191) Seringal Oriente; Acre, Brasil (146) Serra; Espirito Santo, Brasil (185) Serra de Itatiaia; Sao Paulo, Brasil (204) Serra de Parintins; Amazonas, Brasil (139) Serra do Navio; Amapa, Brasil (131) Serra dos Orgaos; Rio de Janeiro, Brasil (189) Serrat Monte; Rio de Janeiro, Brasil (193) 84 FIELDIANA: ZOOLOGY "S. Francisco de C." Minas Gerais, Brasil (not located) A. Robert, March 1900, at 1580 m. Shahuia; Ucayali, Peni (52) Siles (Puerto); El Beni, Bolivia (72) Sitio Minas Gerais; Minas Gerais, Brasil (not lo- cated) Supenam (River) (see Supinaam River); Demer- ara-Mahaica, Guyana (113) Supinaam (River); Demerara-Mahaica, Guyana (1 13) Tacuati; San Pedro, Paraguay (90) Taiuana (Ilha); Para, Brasil (153) Tamatama; Territorio Federal Amazonas, Vene- zuela (110a) Tapereba (Igarap^); Pard, Brasil (150) Taquara; Rio Grande do Sul, Brasil (211) Tefe; Amazonas, Brasil (137) Teresinha; Amapd, Brasil (131) Teres6polis; Rio de Janeiro, Brasil (189) Terezinha (see Teresinha); Amapa, Brasil (131) Theresopolis (see Teresopolis); Rio de Janeiro, Brasil (189) Ticoporo, Reserva Forestal; Barinas, Venezuela (100a) Tingo Maria; Hu^nuco, Peni (59) Tobesobe, Guayo; Territorio Federal Delta Ama- curo, Venezuela (98c) Tocomechi; Santa Cruz, Bolivia (88) Tres Troncos; Caquetd, Colombia (17b) Trinidade; Goids, Brasil (175a) Turumbdn (= San Martin de Turumban); Bolivar, Venezuela (104a) Uragua-i (Rio); Misiones, Argentina (95) Urubamba (Rio); Ucayali, Peni (54) Urucu, Rio; Amazonas, Brasil (145b) Urucum; Mato Grosso do Sul, Brasil (170) Utcuyacu; Juni'n, Peni (61) Utinga; Pard, Brasil (159) Vaca Diez; El Beni, Bolivia (79) Valao de Sao Louren9o; Espirito Santo, Brasil (187) Ventura; Guayas, Ecuador (28) Vila Oliveira; Sao Paulo, Brasil (202) Vila Velha; Espfrito Santo, Brasil (185) Villa Bella Imperatriz; Amazonas, Brasil (140) Villavicencio; Meta, Colombia (16) Vitoria; Espirito Santo, Brasil (185) Warnes; Santa Cruz, Bolivia (87) Xiriviny (Rio); Amazonas, Brasil (136) Yana Rumi (Rio); Napo, Ecuador (34) Yana Yacu; Ucayali, Peni (47) Yarinacocha; Ucayali, Peni (50) Ypanema (= Ipanema); Sao Paulo, Brasil (199) Yurac Yacu; San Martin, Peni (56c) Yurimaguas; Ucayali, Peni (47) Yurudn (Rio); Bolivar, Venezuela (104b). Gazetteer of South American Collecting Localities of Philander Colombia — Philander opossum fuscogriseus, P. opossum quica, P. andersoni andersoni Antioquia — Philander opossum fuscogriseus 1 . Alto Bonito, 7°0rN, 76°17'W, about 400 m, upper Ri'o Sucio, W slope Cordillera Occidental. L. E. Miller and H. S. Boyle, February 1915. 2. Bellavista, 6°33'N, 75°18'W, upper Rio Porce, Cordillera Centra. R Hershkovitz. February 1950, at 1200 m. Caldas — Philander opossum fuscogriseus 3. Rio Hondo, Samand, 5°42'N, 75°0rw, Cordillera Central. R Hershkovitz, March 1951. Choc6 — Philander opossum fuscogriseus 4. Bagad6, 5°25'N, 76°24'W, Rfo Andagueda. E. L. Kerr, November 1912, at 650 m. 5. Condoto, 5°06'N, 76°37'W, Rio Condoto, tributary of Rio San Juan. H. G. F Spurrell, February 1914, at 92 m. 6. Rio Sand6, Rio Baud6, 5°03'N, 76°57'W. K. von Sneidem, October 1958, at 160 m. 7. Rfo Docampad6, 4°45'N, 77°18'W. K. von Sneidem, September 1958, at 75- 160 m. 8. N6vita, 4°57'N, 76°34'W, Rfo Tamanl L. E. Miller, December 1911, at 120 m. Valle del Cauca — Philander opossum fuscogriseus 9. Rfo Frfo, 4°09'N, 76°15'W, enters Rfo Cauca from east. A. A. Allen and L. E. Miller, November- December 1911, at 100 m. 10. Buenaventura, 3°53'N, 77°04'W. K. von Sneidem, February 1958, at near HERSHKOVITZ: PHILANDER 85 sea level. C. H. Tyndale-Biscoe, September 1971. 10. Buenaventura, 28 km NE. J. A. W. Kirsch, September 1969, at 150 m. 11. Call, 3°27'N, 76°3rw, upper Rio Cauca valley. C. H. Tyndale-Biscoe, September 1971. 1 1. "Rio Cauca" (see Cali). J. H. Batty, June 1898. 12. Raposo (Rio), 3°43'N, 77°08'W, Pacific Virology Field Camp. J. Duran, June 1962, at near sea level. Cauca — Philander opossum fuscogriseus 13. Rio Saija, La Boca (= Mouth), 2°52'N, 77°4rW. K. von Sneidern, June 1958, at near sea level. 14. Cocal, 2°3rN, 77°00'W, upper Rio San Juan. L. E. Miller, July 191 1, at 125 and 187 m. Ecuador — Philander opossum melanurus, P. opossum fuscogriseus, P. andersoni andersoni Esmeraldas — Philander opossum melanurus 18. Cachavi (Cachabi), Rio Cachavi, r03'N, 78°50'W. 18. Carondelet, 1°03'N, 78°50"W. G. Fleming, October 1900. 18. San Javier, 1°04'N, 78°47'W. G. Fleming, August 1900, at 20 m. 19. Esmeraldas, 0°59'N, 79°42'W. W. Richardson, November 1912, at near sea level. 20. Rio Quininde, 0°20'N, 79°28'W. Imbabura — Philander opossum melanurus 21. Paramba, 0°49'N, 78°2rw, 1100 m. W. F H. Rosenberg, April 1897; L. Gomez, June 1941. Boyaca — Philander opossum quica 15. Rio Covaria, 7°03'N, 72°04'W, near mouth, above town of Covaria. K. von Sneidern, March 1959, at 350 m. Meta — Philander opossum quica 16. Caney, Rio Guatiquia (see Restrepo). 16. Restrepo, 4°15'N, 73°33'W. 16. Villavicencio, 4°09'N, 73°37'W, upper Rio Guatiquia. R. Gilmore, February, May, June 1939, at 465 m. C. H. Tyndale-Biscoe, 1971. 16. El Capricho (Finca), 38 km E Villavicencio, 4°09'N, 73°16'W. J. A. W. Kirsch, October 1969. Putumayo — Philander andersoni andersoni 17a. Puerto Asis, 17 km N; 0°3rN, 76°3rw. J. A. W. Kirsch, November 1969, at 330 m. Caqueta — Philander andersoni andersoni 17b. Tres Troncos, 0°08'N, 74°4rw, above La Tagua, Rio Caqueta. P Hershkovitz, January 1952, at 182 m. Pichincha — Philander opossum melanurus 22. Rio Caoni, 0°12'N, 79°23'W. M. Olalla, January 1935. 23. Gualea, 0°08'N, 78°48'W, ca. 1250 m. L. Soderstrom, May 1920. 24. Mindo, below 8°02'S, 78°48'W. L. Soderstrom, December 1924, at 1000 m. Manabi — Philander opossum fuscogriseus 25. Rio Pescado, 1°25'S, 80°15'W. G. H. H. Tate, May 1922, at 500 m. 26. Rio de Oro, 2°10'S, 79°22'W. W. Richardson, January 1917. Guayas — Philander opossum fuscogriseus 27. Guayaquil, 2°10'S, 79°50'W, near sea level. E. Belcher, 1838. 28. Puente de Chimbo, 2°10'S, 79°07'W. G. H. H. Tate, August 1922, at 375 m. 28. Rios Chimbo-Coco, 2°10'S, 79°50'W. G. H. H. Tate, July 1922, at 750 m. 28. Bucay, Rio Chimbo, 2°10'S, 79°06'W. G. H. H. Tate, November 1921, at 312 m. 28. Ventura, Rio Chanchan, 2°17'S, 79°24'W. G. H. H. Tate, April 1922, at 750 m. 86 FIELDIANA: ZOOLOGY El Oro — Philander opossum fuscogriseus 29. El Chiral, 2''39'S, 79°43'W. H. E. Anthony, August 1920, at 167 m. 29. Piiias, 3°42'S, 79°42'W. I G. H. H. Tate, September 1921, at 1 125 m. 29. Salvias, 3°47'S, 79°2rw. H. E. Anthony, August 1920, at 1000 m. Loja — Philander opossum fuscogriseus 29. Santa Ana, Punta (?) 3°50'S, 79°25'W, on road from Zaruma to Loja. H. E. Anthony, December 1920, at 1 14 m. 30. Seboyal (see Cebollal). 30. Cebollal, 4°02'S, 80°02'W. G. H. H. Tate, October 1921, at 968 m. Nsipo^Philander andersoni andersoni 31. Santa Cecilia, 0°03'N, 76°58'W. 32. San Jos^, abajo (= below), 0°3rS, 77°20'W. Olalla Brothers, April 1924. 33. Puerto Naop, Rio Napo, 1°03'S, 77°47'W. 33. Rio Napo (near) (see Puerto Napo). L. Soderstrom, March 1921. 34. Rfo Yana Rumi, 1°38'S, 76°59'W. R. Olalla, October 1934. Pastaza — Philander andersoni andersoni 35. Sarayacu, Rio Bobonaza, 1°44'S, 77°29'W. G. H. H. Tate, March 1924, between 450 and 500 m. 36. Montalvo, Rio Bobonaza, 2°04'S, 76°58'W. R. Olalla, February 1932. 37. Rio Copataza, 2°07'S, 77°27'W. R. Olalla, March 1939; G. H. H. Tate, March 1924. 38. Rio Pindo Yacu, 2°08'S, 76°03'W. R. Olalla, October 1934, at 250 m. Peru — Philander opossum quica, P. andersoni andersoni, P. andersoni mcilhennyi Ucayali — Philander opossum quica (42, 43, 48- 51, 54, 55), P. andersoni andersoni (39-41, 44- 47), P. andersoni mcilhennyi (5 1 , 55). 39. Ri'o Curaray. Boca (= mouth), 2°22'S, 74°05'W. Olalla Brothers, 1925. 40. Lago Miranes, Rio Napo (see Rio Mazdn). 40. Rfo Mazan (mouth, 3°28'S, 73°02'W. L. Soderstrom, September 1930. 41. Santa Luisa, Rio Nanay, 3°35'S, 74°30'W. C. Kalinowski, October 1956. 42. Itaya (see Iquitos). H. Bassler, 1927. 42. Iquitos, Rio Maraiion, 3°46'S, 73°15'W, 106 m. H. Bassler, March 1928; C. Kalinowski, October 1956; R. W. Hendee, January 1928; C. Arevalo (H. Bassler collection), March 1930. 43. Apayacu, 3°19'S, 72°07'W. Olalla Brothers, January 1927. 44. Orosa, 3°26'S, 72°08'W. OUalla Brothers, October 1926. 45. Nauta, 4°30'S, 73°25'W, ca. 130 m. 46. Santa Elena, Rio Samiria, 4°42'S, 74°12'W, ca. 130 m. C. Kalinowski, November 1956. 47. Yurimaguas, Rfo Huallaza, 5°54'S, 76°05'W, 180 m. 47. Yana Yacu, below Yurimaguas, 5°52'S, 76°15"W. M. R Anderson, September 1912, at 180 m. 48. Sarayacu, Rfo Ucayali, 6°44'S, 75°06'W. A. M. Olalla, March, April 1927. 49. San Jer6nimo, 7°45'S, 74°50'W, 300 m. R. W. Hendee, December 1927. 50. Yarinacocha, 8°15'S, 74°43'W, 160 m. C. C. Sanborn, March 1946; A. L. Gardner, August 1968, March 1971. 51. Pucallpa, 59 km W, 59 km NE, 59 km SW (see Pucallpa. Rfo Ucayali). H. Hinse, October, November 1971, October 1972; H. Hinse, 59 km NE, September 1972; 59 km SW, October 1972. 51. Pucallpa, Rfo Ucayali, 8°23'S, 74°32'W, 180 m. 52. Cumeria (= Cumarfa, across from Shahufa, q.v.). 52. Shahufa, Rfo Ucayali opposite Cumarfa, 9°52'S, 74°0rW. R. W. Hendee, July 1927, at 312 m. 53. Chicosa, 10°2rS, 74°00'W, ca. 150 m. R. W. Hendee, September 1927. 54. Lagarto Cocha, Rfo Ucayali, 10°4rs, 73°48'W. Olalla Brothers, January 1928. 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. Olaila Brothers, December 1927. 55. Balta, Rio Curanja, 10°08'S, 71°15'W, 300 m. A. L. Gardner, June 1968, March 1971; R. Thomas, 1971; A. L. Gardner, July, August 1966, February 1977, July 1968; J. R O'Neil, February 1971. 62. San Jose, Rio Santa Rosa, 12°44'S, 73°46'W. A. L. Gardner, April 1971, at 1000 m. 62. Huanhuachayo, 12°44'S, 73°47'W. A. L. Gardner, May 1971, at 1660 m. Cuzco — Philander opossum quica Amazonas — Philander andersoni andersoni 56a. La Poza, Rio Santiago, 4°0rS, 1T41'W. J. L. Patton, August 1979, at 180 m. 56b. Huampami, Rio Cenepa. J. L. Patton, July, August 1977, July 1978. San Martin — Philander opossum quica 56c. Yurac Yacu, 5°52'S, 77°14'W. R. W. Hendee, July 1926, at 780 m. 57. Moyobamba, Rio Mayo, 6°03'S, 76°58'W. W. H. Osgood and M. P. Anderson, August 1912; L. Rutter, February 1924, June 1926, at 845 m. 58. Rioja, 6°10'S, 77°10'W, ca. 800 m. R. W. Hendee, September 1926. 63. Hacienda Cadena, Marcapata, 13°24'S, 70°43'W, 890 m. C. Kalinowski, October 1949, August 1950. 63. SanJeronimo, 13°34'S, 7r54'W. 64. Quincemil, 13°16'S, 70°38'W. C. Kalinowski, June 1953, at 680 m. Madre de Dios — Philander opossum quica 65. Colorado, Rio (mouth) (see Boca Colorado). 65. Boca Colorado, 12°30'S, 70°10'W. C. Kalinowski, October 1954, at 279 m. 66. Puerto Maldonado, Rio Tambo, 12°36'S, 69°irW, 256 m. M. L. Kuns, April 1965. Bolivia — Philander opossum quica La Paz Huanuco — Philander opossum quica 59. Tingo Maria, 9°08'S, 75°57'W. R. W. Hendee, January 1927, at 625 m. Pasco — Philander andersoni andersoni 60. Oxapampa, 10°34'S, 75°24'W. C. R. Perez, August 1964, at 280 m. Junin — Philander andersoni andersoni 61. Chanchamayo, 1 1°03'S, 75°19'W. P O. Simons, August 1900; J. M. Schunke, August 1948; M. Kalinowski before 1916. 61. Utcuyacu, 1 n2'S, 75°28'W. P O. Simons, August 1900, at 1600 m. 67. Culumani (see Chulumani). 67. Chulumani, 16°24'S, 67°3rW, 2000 m. P O. Simons, February 1901, at 2200 m. El Beni 68. Guayaramarin, Ri'o Mamore, 10°5rS, 65°23'W. A. Ximenez, June 1964; K. F Koopman, 1964. 69. Merredor, Rio Mamore, ll°4rs, 65°05'W. D. Edwards, July 1965; A. Ximenez, May 1965. 70. Rio Mamore, 12°26'S, ca. 65°W. f D. E. Afiez, October 1965. « 71. Itonama, 12°28'S, 64°24'W. M. L. Kuns, April 1970. 72. Puerto Siles, Rfo Mamore, 12°49'S, 65°00'W. S. Anderson, October 1965. FIELDIANA: ZOOLOGY 72. Santa Rosa, 13°01'S, 65°1 1'W. M. L. Kuns, June 1964. 73. Barranquita (see San Joaquin). M. L. Kuns, August 1963. 73. Arruda, 4 km NE San Joaquin (q.v.). M. L. Kuns, December 1963. 73. Centinela, 1.5 km E San Joaquin (q.v.). M. L. Kuns, June 1964. 73. Camino Vilches, 1 mi E San Joaquin (q.v.). M. L. Kuns, February 1966. 73. Santo Dios, 2 km NE San Joaquin (q.v.). M. L. Kuns. June 1963. 73. San Joaquin. Ri'o Machupo, 13°04'S, 64°49'W. M. L. Kuns, April, June, July 1963; February, May 1964. 74. Exaltacion, Rio Mamord, 13°16'S, 65°15'W. M. L. Kuns, May 1964; S. Anderson, October 1965, at 8 km N. 74. Palacios, 13°34'S, 65°19'W. D. E. Anez. May 1965. 74. Puerto Caballo, 13°34'S, 65°2rw. S. Anderson, September 1965. 75. San Ramon, Ri'o Machupo, 13°18'S, 64°37'W. M. L. Kuns, January 1964. 75. Monte Chocolatal (see San Ram6n). M. L. Kuns, January 1964. 76. Magdalena, 13°20'S, 64°08'W. M. L. Kuns, July 1963, July 1964. 77. San Pablo, 13°52'S, 65°36'W. M. L. Kuns, September 1963. 78. El Carmen, Rio Blanco, 13°57'S, 63°43'W. M. L. Kuns, June 1964. 79. Rio lhar6 (mouth), 14°37'S, 64°57'W. D. E. Anez, August 1965. K. F. Koopman, August 1965. 79. Vaca Diez, 14°47'S, 64°5rw. M. L. Kuns, May 1969. 80. San Ignacio de M6xos, 14°53'S, 65°36'W. M. L. Kuns, June 1965. 81. Camiaco, 15°24'S, 64°46'W. A. Xim6nez, August 1965. Cochabamba 82. Rfo Ichilo, 16°50'S, 64°45'W. D. E. Aiiez, July 1965. Santa Cruz 83. 2 km S mouth, Ri'o Chapar6, Rio Mamor6, 15°58'S, 64°42'W. D. E. Aiiez, July 1965. 84. Ascencion de Guarayos, 14°57'S, 61°24'W. M. L. Kuns, June 1964. 85. Mercedes, Rio Guapore, 15°36'S, 60°22'W, 6 km S and opposite Buena Hora, Brasil. A. Ximenez, May 1965. 86. Buenavista, 17°27'S, 63°40'W. T. Bridges, probably 1846; F. Steinbach, January 1915; J. Steinbach, June 1921, May 1925, July 1926, June, July 1927, July 1928. 87. Wames, 17°3rs, 63°10'W. M. L. Kuns, August 1965. 87. Santa Rosita, Wames, 17°30'S, 63°10'W. M. L. Kuns, July 1965. 88. Tocomechi, 17°35'S, 62°55'W. M. L. Kuns, August 1965. 88. Hamacas, 17°44'S, 63°1 1'W. O. Silva (Rockefeller Institution), July 1938. 88. El Palmar, 17°48'S, 63°10'W. W. Kerr, August 1966, at 500 m; D. R. Hadden, August 1966. 88. Ayacucho, Ibdfiez, 17°5rS, 63°20'W. J. Riddell, August 1966. 88. 18 km SW Santa Cruz (q.v.). F. Becerra, August 1966. 88. Santa Cruz (see Santa Cruz de la Sierra). 88. Santa Cruz de la Sierra, 17°48'S, 63°10'W. F Becerra, September 1966. 89. San Ram6n, 17°33'S, 61°03'S. Paraguay — Philander opossum quica San Pedro 90. Tacuati, 23°27'S, 56°35'W. C. Wharton, May 1950. Central 91. Asunci6n, 25°16'S, 57°40'W. 92. Lapango, 25°2rS, 57°42'W. H. Krieg, August 1925. Paraguarf 93. Sapucay (= Sapucaf), 25°40'S, 56°55'W. W. Foster, September 1902. Argentina — Philander opossum Chaco 94. No precise locality. C. Friend, before 1880. HERSHKOVrrZ: PHILANDER 89 94. Rio de Oro (mouth), 27°04'S, 58°34'W. I. Apostal, October 1962. Misiones 95. Rio Uruguar-i, 30 km from Puerto Bemberg (Libertad). 26°30'S, 54°W. J. A. Crespo, September, October, November 1949. 96. Fracran, San Pedro, 26°46'S, 54°16'W. J. A. Crespo, February 1952. Merida — Philander opossum subspecies 99. "Merida," 8°36'N, 71°08'W. Barinas — Philander opossum subspecies 100a. Ticoporo, Reserva Forestal, 07°48'N, 69°55'W. 100b. Guaquitas, 07°28'N, 71°39'W. 100b. Las Bonitas, Cano Amaru, 07°23'N, 70°44'W. 100c. La Lengueta, 08°30'N, 70°23'W. Venezuela — Philander opossum subspecies, P. andersoni andersoni Zulia — Philander opossum subspecies 97a. Alguacil, Caja Seca, 09°8'N, 71°04'W. 97a. Churulf, Caja Seca, 09°09'N, 71°04'W. 97b. Puerto Catatumbo, 09°07'N, 72°35'W, 50 m. 97b. Boca del Rio de Oro, 9°06'N, 72°45'W. 97b. Encontrados, 9°03'S, 72°14'W. N. E. Peterson, March 1968. 97b. Encontrados, 60 km WNW, 90°03'N, 72°14'W, 73 m. Trujillo — Philander opossum subspecies 98a. Motatan, 9°24'N, 70°36'W, 1 km E at 330 m, 5 km NNE at 290 m. J. A. W. Kirsch, June 1969. 98a. Motatan (Rio), 9°28'N, 70°34'W at 290 m, 9.8 km NNE at 290 m. J. A. W. Kirsch, June 1969. 98a. Agua Viva, 9°34'N, 70°36'W. N. E. Peterson, September 1965, at 164 m. 98a. El Dividive, 9°29'N, 70°44'W. N. E. Peterson, October 1965, at 90 m. Apure — Philander opossum subspecies 101a. Nulita, 07°19'N, 71°55'W. A. Tuttle, January 1968, at 24 m. 101a. La Blanquita, 7°12'N, 71°45'W. A. Tuttle, January 1968, at 24 m. Tachira — Philander opossum subspecies 101b. La Ponchera, 07°26'N, 71°52'S. Bolfvar — Philander opossum opossum 102a. El Palmar, Rio Grande, 08°0rN, 61°55'W. 102b. Maripa, Rio Caura, 7°26'N, 65°09'W. S. M. Klages, 1901. 103. Hato San Jose, 146 km S, 7 km NE Ciudad Bolivar, 6°44'N, 63°27'W. N. Peterson, D. Peacock, R. Peacock, D. Furman, March 1967, at 302 m. 104a. San Martin de Turumban, Rio Cuyuni, 06°59'N, 6r02'W. 104b. Rio Yuruan, 06°48'N, 61°50'W. M. A. Carriker, Jr., March 1910. 104c. Chalimana (Raudal), Rio Paramichi, Rio Paragua, 04°10'N, 62°59'W. Monagas — Philander [?] opossum subspecies 98b. Papelon (Cerro), 10°0rN, 63°54'W. Delta Amacuro (Territorio Federal) — Philander opossum subspecies 98c. Los Guires, 09°15'N, 61°54'W. 98c. Tobesobe, Guayo, 09°00'N, 61°25'W. 98d. Guiniquina, 90°10'N, 61°03'W, 0 m. Amazonas (Territorio Federal) — Philander andersoni andersoni 105. Majagua (Cafio), Rio Ventuari, 05°20'N, 65°40'W. 105. San Juan, Ri'o Manapiare, 05°19'N, 66°03'W. M. D. Tuttle, E L. Harder, July 1967, at 155 m. 106. Belen, Rio Cunucunuma, 03°43'N, 65°42'W. 90 FIELDIANA: ZOOLOGY r M. D. Tuttle, E L. Harder, January 1967, at 150 m. 107. Acanana, Ri'o Cunucunuma, 03°39'N, 65°66'W. M. D. Tuttle, F. L. Harder, June 1967, at 145 m. 108. Playa del Rio Base, Mt. Duida, 03°25'N. 65°40'W. Olalla Brothers, November 1928. 109a. Esmeralda, 03°1 1'N, 65°33'W. M. D. Tuttle, E L. Harder, March 1967, at 135 m. 109b. Cucurito, Cerro, 03°38'N, 66°25'W. 1 10a. Tamatama, Rio Orinoco, 3°08'N. 65°52'W. M. D. Tuttle. E L. Harder, April, May, June 1967, at 135 m; D. S. Bremington, June 1967, at 135 m. 1 10b. Parima (Sierra), 02°40'N, 64°30'W. 1 10c. Mavaca, 02°3rN, 65°10'W. 111. Capibara, Brazo Casiquiare, 2°34'N, 66°18'W. M. D. Tuttle, E L. Harder, June 1967, at 130 m. 1 12a. Merey, opposite, Brazo Casiquiare, 2°17'N, 67°irw. Olalla Brothers, October 1929. 112b. La Neblina (Campamento), Cerro La Neblina, 00°52'N, 66°14'W. Guyana — Philander opossum opossum Demerara-Mahaica 113. Supinaam River (= Supenam River), 6°58'N, 58°3rw. Crozier. 114. Georgetown, Demerara River, 6°48'N, 58°10'W. J. Rodway. June 1929. 115. Hyde Park, Demerara River, 6"'30'N, 58°16'W. S. B. Warren, September 1906. Essequibo Islands-West Demerara 116. Buck Hall, Essequibo River, 6°56'N, 58°33'W. S. B. Warren, March 1906. Upper Takutu-Upper Essequibo 117. Rupununi River, 4°03'N, 58°34'W. J. J. Quelch, September 1900, at 200 ft. 1 18. Kanuku Mountains, 3°N, 59°45'W. E. V. McConnell, J. J. Quelch, November 1900, at 240 ft. Suriname — Philander opossum opossum Nickerie 119. Avanavero Falls, Kabulebo River, 4°49'N, 57°24'W. 120. Nickerie River, upper, ca. 5°59'N, 56°30'W. 121. Makerie, West River, Wilhelmina Mountains. 3°26'N. 56°45'W. H. A. Beatty. December 1961, January 1962. 122. Kaiserberg Airstrip, 3°10'N, 56°15'W. H. A. Beatty. February 1961, at 275 m. Brokopondo 123. Loksie Hattie, 5°09'N. 55°28'W P. Hershkovitz, December 1961. Saramacca 124. La Poule, 5°47'N, 55°25'W. P. Hershkovitz, January, February 1962. Suriname 125. Paramaribo, 5°50'N, 55°1 1'W. I. T. Sanderson, February 1938. 125. Clevia. Paramaribo (q.v.). P. Hershkovitz, February 1962. 125. "Culturutuin," Agricultural Experimental Station in Paramaribo (q.v.). 125. Para River, ca. 10 km SE Paramaribo (q.v.). 125. Rijweg, ca. 9 km W Paramaribo (q.v.). 126. Lelydorp, 5°42'N, 55°16'W. 126. Lelydorpplan (see Lelydorp). P. Hershkovitz, January, February 1962. Commewijne 127. Commewijne River, 5°54'N, 55°05'W. Marowijne 128. Albina, 5°30'N, 54°03'W. HERSHKOVITZ: PHILANDER 91 Guyane Frangaise — Philander opossum opossum Cayenne 129. Cayenne, 4°56'N, 52°19'W. G. K. Cherrie, B. T. Gault, November 1902; S. Klages, January, February 1917; M. Atramentowicz, September 1978- October 1982; P. Charles-Dominique, 1978- 1982. Ouanary 130. Ouanary River, 4°14'N, 51°39'W. Brasil — Philander opossum opossum, P. opossum frenata, P. opossum quica Amapa — Philander opossum opossum 131. Serra do Navio, 0°59'N, 52°03'W. T. P. Woodall, June 1966; Instituto Evandro Chagas, 1967, 1968, 1969. 131. Teresinha (= Terezinha), Rio Amapari, 0°58'N, 52°02'W. 132. Macapa, Rio Amapari, 0°02'N, 51°03'W. M. Moreira, October, November 1952. 133. Mazagao, Rio Maraca, 0°06'S, 5ri8'W. M. Moreira, December 1958. 133. Rio Branco, tributary Rio Maraca, 0°07'S, 5ri7'W. Roraima — Philander opossum opossum 134. Boa Vista, Rio Branco, 2°49'N, 60°40'W. C. T. Carvalho, M. Sobeiro do Amaral, M. Melo, March 1959. 134. Pocao, Boa Vista (see Boa Vista). 135. Caracarai, Rio Mucajai, r50'N, 61°08'W. C. T. Carvalho, M. Sobeiro do Amaral, M. Melo, March 1959. 135. Pocao, Caracarai (see CaracaraO. Amazonas — Philander opossum opossum 136. Rio Xiriviny, 0°59'S, 61°53'W. K. B. Parker, October 1928, at 80 m. 137. Santo Isidoro, Tefe, 3°27'S, 64°47'W. Olalla Brothers, August 1928. 138. Ipixuna (Lago do), 3°52'S, 63°52'W. 139. Serra de Parintins, 2°35'S, 56°25'W. Olalla Brothers, November 1930. 139. Parintins, 2°36'S, 56°44'W. Olalla Brothers, November 1930. 140. Santa Clara, Villa Bella Imperatriz, 2°50'S, 56°55'W. Olalla Brothers, August 1930. 141. Lago do Baptista, 3°18'S, 58°15'W. A. M. Olalla, June 1936. 142. Rio Madeira, mouth, 3°22'S, 58°45'W. Olalla Brothers, February 1930. 143. Rosarhino, 3°43'S, 59°08'W. Olalla Brothers, June, July 1930. 144. Auara Igarape, 4°22'S, 59°43'W. Olalla Brothers, March 1930. 145a. Santo Antonio de Uayara, Rio Eini, 6°43'S, 69°52'W. Olalla Brothers, April 1930. 145b. Rio Urucu, 4°51'S, 65°16'W. M. N. E da Silva. Acre — Philander opossum quica 146. Seringal Oriente, 8°48'S, 72°46'W. M. Moreira, August 1934; M. Moreira, E Novaes, August 1956. Para — Philander opossum opossum 147. Fordlandia, Rio Tapajoz, 3°40'S, 55°30'W. R. M. Gilmore, February 1938. 148. Km 19, Itaituba-Jacareacanga, 4°17'S, 56°05'W. E Ramos, Instituto Oswaldo Cruz, August 1972. 148. Km 216, Estrada de Santarem-Cuiaba (BR 165) (see Itaituba). Instituto Evandro Chagas, May, June 1973. 149. Canudos, 7°16'S, 58°07'W. E Lima, November 1920. 150. Tapereba, Igarape, Chaves, 0°10'S, 49°55'W. C. Carvalho, June, July 1958. 151. Altamira, Rio Xingu, 3°12'S, 52°12'W. Instituto Oswaldo Cruz, August 1971. 152. Gradaus, 7°43'S, 51°11'W. C. T. Carvalho, N. Hidasi, M. Amaral, June, July, August 1957. 153. Cameta, Rfo Tocantins, 2°15'S, 49°29'W. A. M. Olalla, February 1934, March, April, May, November 1935, November 1936. 92 FIELDIANA: ZOOLOGY 153. Ilha do Taiuna, Rfo Tocantins, 49°29'W, opposite Cameta. A. M. Olalla, October, November 1931. 154. Mazagao, Rio Tocantins. 2°25'S, 49°10'W. F. Lima, November 1912. 155. Mocajuba, Rio Tocantins. 2°35'S. 49°30'W. A. M. Olalla. November 1931. 156. Baiao, Rio Tocantins. 2°41'S. 49°41'W. A. M. Olalla. December 1931. 157. Arumateua. Rfo Tocantins. 3°54'S. 49°41 'W. F. Lima. October 1912. 158. Maraba. Rio Tocantins. 5°20'S, 49°10'W. N. Peterson. September 1975. 159. Para (see Bel^m). 159. Km 90. Para. Rodovia E. Snethlage. April 1909. 159. Belem do Pard (see Belem). 159. Belem. 1°27'S. 48°29'W. M. A. Miles. 1975-1980. 159. Ipeau-Apez. Belem (see Belem). E. Snethlage. 159. Marco, suburb of Belem (q.v.). 159. Murutucu (= Institute Agronomico do Norte). Belem (q.v.). E Luna. August 1922, March 1925. 159. Utinga. suburb of Belem (q.v.). Institute Oswaldo Cruz, June 1963; R. H. Pine. June 1968. 159. Ananinddua. E. E Braganqa. 1°22'S, 48°23'W. E Lima, May 1920. 159. Sapucajuba. suburb of Belem (q.v.). 160. Ipitinga. Rio Acara. 1°57'S. 48°1 1'W. 161. Elor do Prado, near Quatipuru, 0°52'S. 46°59'W. E. Snethlage. October 1916. 162. Santa Maria. Braganqa. 1°03'S. 46°46'W. April 1968. 163. Peixe-Boi. R. R. Bragan<;a. n2'S,47°18'W. 163. Igarape Aqu. 1°32'S. 47°03'W. 163. Colonia do Prata, Igarape Aqu (q.v.). 164. Capim. 1°30'S. 48°20'W. Institute Oswaldo Cruz, 1960. 165. Sao Miguel do Guamd (see Guamd). 165. Guamd. 1°37'S. 47°27'W. Departamento do Zoologia, Sao Paulo. October 1959. 165. Iriteria, Sao Miguel do Guamd (see Guamd). M. Amaral. December 1959, January 1960. Rondonia — Philander opossum quica 166. Porto Velho. 8°46'S. 63°54'W. T Hibbs. April 1965. 2 ° 1 5 ' S . Mato Grosso — Philander opossum quica \bl. Mato Grosso. 15°00'S. 59°57'W. J. Natterer. September-November 1824. 168a. Cdceres. 16°04'S, 57°41'W. M. L. Kuns, September 1965. 168b. PcKone. 16°15'S. 56°37'W. 168c. Ban-a do Gardes. 15°53'S. 52°15'W. 168d. Balisa. 16°15'S. 52°26'W. 169. Monte Alegre. Palmeiras. 16°03'S. 55°30'W. A. M. Olalla. June 1944; A. Aggio, June 1944. Mato Grosso do Sul — Philander opossum quica 170. Commbd. Rio Paraguay, 19°0rS, 57°39'W. M. L. Kuns, September 1965. 170. Urucum, 19°13'S, 57°33'W. L. E. Miller. December 1913. at 125 m. 170. Santa Teresa (see Urucum). M. L. Kuns. September 1965. 171a. Miranda. 20°14'S. 56°22'W. A. M. Olalla. 1957; J. Lima. September 1937. 171b. Salobra. 21°14'S. 57°08'W. L. Travassos, May 1942. Bahia — Philander opossum frenata 172. Bahia (= Salvador). 12°59'S. 38°3rw. Herr Kaehne, before 1815. Golds — Philander opossum quica 173. Aragarqas, 15°55'S, 52°15'W. M. Amaral. May 1958. 174a. Formosa. 15°32'S. 47°20'W. 174b. Anapolis. I6°20'S. 48°58'W. R. M. Gilmore. 1936-1937. 175a. Trinidade. I6°40'S. 49°30'W. S. Hidasi. June 1962. Distrito Federal — Philander opossum quica 175b. Parque Nacional. 15°35'S, 48°54'W. 175c. Brasilia, 15°47'S, 47°55'W. 175c. Agua Limpa (Fazenda). 15°57'S, 47°54'W. HERSHKOVITZ: PHILANDER 93 Minas Gerais — Philander opossum quica 176. Concei^ao do Mato Dentro, 19°0rS, 43°25'W. 177. Lagoa Santa, 19°38'S, 43°53'W. P. W. Lund, between 1833 and 1880. 178. Quartel de Sacramento, 19°44'S, 42°3rw. J. Pinto Fonseca, July 1919. 178. Fazenda de Floresta, Rio Matipo, 19°53'S, 42°33'W. J. Pinto Fonseca, July 1919. 178. Esmeralda (Fazenda), ca. 19°20'S, 42°50'W. 178. Monies Claras, 19°25'S, 42°35'W. 179. Boa Esperan9a, Serra de Caparao, 20°10'S, 41°46'W. E. Kaempfer, August 1929. 180. Fazenda Cardosa, Serra de Caparao, 20°22'S, 41°48'W. E. G. Holt, June 1922. 181. Benfica, Serra de Itatiaia, 21°4rS, 43°26'W. E. H. Holt, June 1922, at 2000 ft. 182. Alem Paraiba, 21°52'S, 42°41'W. 183. Juiz de Fora, 21°45'S, 43°20'W. Servigo do Estudos e Pesquisas sobre a Febra Amarela (SEPSFA). Espirito Santo — Philander opossum quica 184. Santa Teresa, 19°55'S, 40°36'W. A. M. Olalla, October 1942. 184. Sao Joao de Petropolis, 19°49'S, 40°40'W. C. Lako, June 1940. 185. Serra, 20°07'S, 40°18'W. C. Lako, September 1949, at 50 m. 185. Campinho, 20°07'S, 40°17'W. C. Lako, April 1940, at 500 m. 185. Vitoria, 20°19'S, 40°2rw. C. Lako, April 1940. 185. Vila Velha, Morro de Angoles, 20°20'S, 40°17'W. C. Lako, March, April 1940, June 1941. 186. Caixa D'agua, 20°38'S, 40°55'W. C. Lako, June 1940. 187. Valao de Sao Lourengo, Santa Teresa, 2o°37's, 4r4rw. C. Lako, June 1940. 188. Engenheiro Reeve (now Rive), 20°46'S, 41°28'W. A. Robert, March, April 1903. G. R. Hancock, September 1923, at 3000 ft; R. Kellogg, March 1943; G. Perreira, September 1942; D. E. Davis, May, September, October 1943; C. Guinle, Perreira, January 1943; C. Guinle, P. M. Britto, November 1942, March 1943, May 1943; C. Guinle, November 1942; C. Guinle, H. W. Laemmart, April 1943; P M. Britto, January, May 1943. 189. Boa Vista, Fazenda, Teresopolis (see Teresopolis). 189. Boa Fe, Fazenda, Teresopolis, 22°22'S, 41°53'W. D. E. Davis, October, September 1943; R M. Britto, January 1943; G. Perreira, September 1942; C. Guinle, November 1942; C. Guinle, R M. Britto, March, May 1943; C. Guinle, H. W. Laemmart, April 1943. 189. Novo Friburgo, 22°16'S, 42°32'W. Herr Beschke. 189. Barrieira, Serra dos Orgaos, 22°56'S, 42°56'W. Schind in Pohle, 1927. 189. Serra dos 6rgaos, 22°56'S, 42°56'W. 190. Rio de Janeiro, 22°54'S, 43°14'W. R. M. Gilmore, January, February 1938. 191. Rodeio, Serra do Mar, 22°33'S, 43°4rw. G. B. Flowers. 191. Sepetiba, 22°58'S, 43°42'W. J. Natterer, March 1818. 191. Sapitiba (see Sepetiba). 191. Sao Joao Marcos, 22°54'S, 43°58'W. C. Lako, October 1938, January 1939. 192. Mangaratiba, 22°57'S, 44°02'W. C. Lako, July, September 1938, February 1939. 193. Itatiaia Parque Nacional, 22°30'S, 44°34'W. F. Gouvea, July 1957. 193. Monte Serrat, Parque Nacional de Itatiaya. C. Moreira, 1901; F. Gouvea, March 1951, at 800 m, August 1957, at 850 m. 193. Macieiras Itatiaia, 22°30'S, 44°34'W. F. Lima, December 1949; F. Gouvea, December 1949. 193. Barro Branco, 22°23'S, 44°30'W. A. Passarell, April 1941. 194. Pedra Branco, Parati, 23°13'S, 44°43'W. C. Lako, July 1943. Rio de Janeiro — Philander opossum quica 189. Theresopolis (see Teresopolis). 189. Teresopolis, 22°27'S, 42°57'W. Sao Paulo — Philander opossum quica 195. Avanhandava, Lajeado, 20°57'S, 48°46'W. E. Garbe, April 1910. 94 FIELDIANA: ZOOLOGY upper Rio Tiete, 22°10'S, 209. Colonia Hansa (see Hansa). 196. Boraceia, 48°45'W. J. Pinto, May 1958; L. Travassos, March, April 1958; see also Boraceia (203). 197. Iporanga, Lajeado, 24°36'S, 48°34'W. E. Dente, December 1944. 198. Monte Alegre, Amparo, 22°40'S. 46°4rw. J. Lima, June 1944. 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 1962, November 1963. 200. Cantareira, Serra da, 23°25'S, 46°39'W. 2(X). Butanta, Serra da Cantereira (q.v.). J. Navas, April 1910. 201. Juquia, 24°19'S, 47°36'W. 201. Costao dos Engenhos, 24°4rs, 47°25'W. A. M. Olalla, July 1964. 202. Alto da Serra, 23°47'S, 46°19'W. 202. Vila Oliveira, Magi das Cruzes, 23°31'S, 46°irw. J. Lima, September 1943. 203. Salesopolis, Boraceia, 23°32'S, 45°5rw. J. Oliveira, June, July 1961. 203. Casa Grande, 20 km S Salesopolis (q.v.). O. de Sousa Lopez, April 1962, May f 1966, May 1968. 203. Boraceia, Ponta de, 23°48'S, 45°49'W. Near Salesopolis Biological Station; not to be confused with Boraceia (196). 204. Serra de Itatiaia, 22°55'S, 45°28'W. 205. Piquete, 22°37'S, 45°10'W. Zech, January 1897. 206. Sao Sebastiao, 23°48'S, 45°25'W. [ A. Hempel, July, August 1900. Parang — Philander opossum quica 207. Rio Paracai, 23°41'S, 53°57'W. E. Dente, S. Siraglia, January 1954. 208. Ro(;a Nova, Serra do Mar, 25°30'S, 48°50'W. A. Robert, August, September 1901. Santa Catarina — Philander opossum quica 209. Joinville, 26°18'S, 48°50'W. E. Steiger, August 1930. 209. Hansa, 26°27'S, 48°50'W. W. Ehrhardt, August 1928. Rio Grande do Sul — Philander opossum quica 210. Passo Fundo, 28°15'S, 52°20'W. 211. Taquara, 29°3rS, 50°47'W. R. Hensel, before 1867; H. von Ihering, before 1888. 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