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Novltates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novltates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3064, 34 pp., 8 figures, 2 tables June 10, 1993 Philippine Rodents: Chromosomal Characteristics and Their Significance for Phylogenetic Inference Among 13 Species (Rodentia: Muridae: Murinae) ERIC A. RICKART1 AND GUY G. MUSSER2 ABSTRACT Karyotypes are reported for 13 murines be- bers of 50 and 88, respectively, indicating longing to the endemic Philippine genera Apomys, substantial chromosomal variability within that Archboldomys, Batomys, Bullimus, Chrotomys, genus. Archboldomys (2N = 26, FN = 43) has an Phloeomys, and Rhynchomys, and the widespread aberrant sex chromosome system and a karyotype genus Rattus. The karyotype of Phloeomys cum- that is substantially different from other taxa stud- ingi (2N = 44, FN = 66) differs from that of P. ied. The karyotype ofBullimus bagobus (2N = 42, pallidus (2N = 40, FN = 60), and both are chro- FN = ca. 58) is numerically similar to that of the mosomally distinct from other taxa examined. Two native Rattus everetti and the two non-native spe- species of Batomys (2N = 52), Chrotomys gon- cies ofRattus, R. tanezumi and R. exulans. Chro- zalesi (2N = 44), Rhynchomys isarogensis (2N = mosomal data corroborate some phylogenetic re- 44), and Apomys musculus (2N = 42) have FN = lationships inferred from morphology, and support 52-53 and a predominance of telocentric chro- the hypothesis that the Philippine murid fauna is mosomes. Two other species ofApomys have dip- composed of separate clades representing inde- loid numbers of30 and 44, and fundamental num- pendent ancestral invasions of the archipelago. INTRODUCTION The Philippine Islands support a remark- Heaney and Rickart, 1990; Musser and Hea- ably diverse murine rodent fauna, including ney, 1992). This assemblage is also predom- at least 46 indigenous species representing 22 inantly endemic: 43 species (93%) are re- genera (Heaney, 1986; Heaney et al., 1987; stricted to the archipelago, and 36 species Curator of Vertebrates, Utah Museum of Natural History, University of Utah, Salt Lake City, Utah 84112. 2 Archbold Curator, Department of Mammalogy, American Museum of Natural History. Copyright © American Museum of Natural History 1993 ISSN 0003-0082 / Price $5.30 2 AMERICAN MUSEUM NOVITATES NO. 3064 (78%) belong to 16 endemic genera. The high data for the endemic Batomys, Apomys, degree ofendemism suggests a pattern ofvery Chrotomys, Rhynchomys, Archboldomys, and few colonizations, followed by repeated spe- Bullimus are reported for the first time. Data ciation within the archipelago (Heaney and also are reported for endemic species of Rickart, 1990). Our ability to discern more Phloeomys and Rattus, and for two non-na- detailed biogeographic patterns among Phil- tive species of Rattus. We discuss the signif- ippine murids is hampered by incomplete un- icance of these chromosomal traits for phy- derstanding of their phylogenetic relation- logenetic inference within the context of the ships. These can be clarified only through alternative hypotheses of relationships out- acquisition of additional specimens, precise lined by Musser and Heaney (1992). diagnosis of taxa, and analyses of multiple data sets. MATERIALS AND METHODS In a survey of morphological characteris- tics, Musser and Heaney (1992) provided a Animals were collected from wild popu- series of alternative hypotheses of possible lations and killed with sodium pentobarbital phylogenetic relationships among murids na- (Nembutal) or chloroform within 24 hours tive to the Philippines (exclusive of the Pa- of capture. Bone marrow preparations were lawan faunal region). They recognized three obtained following the methods of Patton major divisions: (I) "OLD ENDEMICS" (1967) with the exception that 0.4 percent (species ofPhloeomys, Crateromys, Batomys, potassium chloride was used for the hypo- Carpomys, Apomys, Crunomys, Archboldo- tonic treatment. Cells were processed and mys, Chrotomys, Celaenomys, and Rhyncho- fixed in the field, and standard karyotypes mys), (II) the distinctive monotypic genus prepared in the laboratory from stored cell Anonymomys, and (III) "NEW ENDEM- suspensions following methods of Rickart et ICS" (species of Tryphomys, Abditomys, Bul- al. (1989). limus, Tarsomys, Limnomys, and indigenous Four terms are used to describe the shape species of Rattus). The genera in Division I, ofchromosomes based on the position ofthe each characterized by a combination ofprim- centromere: metacentric (biarmed chromo- itive murine traits and uniquely derived spe- somes with arms ofnearly equal length); sub- cializations, comprise six distinctive clusters metacentric (biarmed chromosomes with a with uncertain phylogenetic relationships; the short arm at least one third the length of the affinities of Anonymomys appear to be with other); subtelocentric (biarmed chromo- members of the Sundaic fauna; and each ge- somes with a short arm less than one third nus in Division III is diagnosed by autapo- the length of the other); telocentric (chro- morphic features and a suite of synapomor- mosomes with a very minute second arm or phies shared with Rattus, but otherwise no discernable second arm). Determinations phylogenetic relationships within the divi- ofdiploid number were based on counts from sion are unclear (Musser and Heaney, 1992). a minimum of 10 cells per individual. Fun- In addition to the indigenous fauna, there damental numbers (FN) refer to the total is a separate group of murines that includes number ofmajor chromosome arms (exclud- several species of Rattus (tanezumi minda- ing very minute arms). Specimens examined nensis, rattus, argentiventer, nitidus, norveg- were prepared as skeletons, skins with partial icus, and exulans) as well as one species of skeletons, or preserved in fluid and are de- Mus (musculus castaneus) with wide distri- posited in the National Museum of Natural butions outside the Philippines (Musser and History, Washington, D.C. (USNM), and the Newcomb, 1983). These species have been Philippine National Museum, Manila (PNM). introduced, most likely through human ac- The species we sampled are in the subfam- tivity (Musser, 1977). ily Murinae of the family Muridae, as out- Against this framework of groupings de- lined by Carleton and Musser (1984) and rived from morphology, we present standard Musser and Carleton (1993), and the scien- karyotypes for 13 species of Philippine mu- tific names we use are those employed by rines belonging to 8 genera. Chromosomal Musser and Carleton (1993) and Musser and 1993 RICKART AND MUSSER: PHILIPPINE RODENTS: MURINE KARYOTYPES 3 Heaney (1992). Identifications of the species (09022'N, 123009'E), two males, one female ofApomys from the central Philippines differ (USNM 458751, 458753, and 458754). from those presented in Musser's (1982a) re- Archboldomys luzonensis Musser, 1982: view of the genus, and are based on exami- Luzon Island, Camarines Sur Province, nation of recently acquired specimens. Spec- Mount Isarog, 4 km N, 21.5 km E Naga, imens from Leyte and Biliran are identified 1350-1750 m (13°40'N, 123°22'E), three as Apomys littoralis, whereas those from Ne- males, one female (USNM 573505, 573835, gros probably represent an undescribed spe- 573837, and 573840). cies (Heaney et al., 1989; Musser and Hea- Chrotomys gonzalesi Rickart and Heaney, ney, 1992). 1991: Luzon Island, Camarines Sur Prov- ince, Mount Isarog, 4 km N, 21.5 km E Naga, 1350-1750 m (13°40'N, 123°22'E), two fe- SPECIMENS EXAMINED males (USNM 458955 and 458956). Phloeomys cumingi (Waterhouse, 1839): Rhynchomys isarogensis Musser and Free- Catanduanes Island, Catanduanes Province, man, 1981: Luzon Island, Camarines Sur vicinity of Barrio Summit, 250 m (13°47'N, Province, Mount Isarog, 4 km N, 21.5 km E 124019'E), one male, four females (USNM Naga, 1350-1750 m (13°40'N, 123°22'E), 573154, 573330, and 573332; PNM speci- three males, one female (USNM 573575, mens (EAR field numbers 1662 and 1663); 573901, 573905, and 573910). Luzon Island, Camarines Sur Province, Bullimus bagobus Meams, 1905: Leyte Is- Mount Isarog, 4 km N, 16 km E Naga, 400 land, Leyte Province, Mount Pangasugan, m (13°40'N, 123°19'E), one female (USNM 10.5 km N, 4 km E Baybay, 900 m (10°47'N, 573509). 124°50'E), one male (USNM 458789). Batomys granti Thomas, 1895: Luzon Is- Rattus everetti (Gunther, 1879): Biliran Is- land, Camarines Sur Province, Mount Isarog, land, Leyte Province, 5 km N, 10 km E Na- 4 km N, 22 km E Naga, 1550-1750 m val, 850 m (11036'N, 124°29'E), two males (13040'N, 123022'E), two males, two females (USNM 458800 and 459901); Catanduanes (USNM 458914, 458944, 458946, and Island, Catanduanes Province, 8.5 km W 458949). Gigmoto, 250 m (13047'N, 124°l9'E), two Batomys salomonseni (Sanbom, 1953): males (USNM 573173 and 573343); Leyte Leyte Island, Leyte Province, Mount Pan- Island, Leyte Province, Mount Pangasugan, gasugan, 10.5 km N, 4 km E Baybay, 700- 10.5 km N, 4 km E Baybay, 700-1000 m 1000 m (10°47'N, 124050'E), three males, (10047'N, 124050'E), four males, one female three females (USNM 458783, 458784, (USNM 458816, 458817, and 458820- 459515, 459872, 459873, and 459886). 458822); Luzon Island, Camarines Sur Prov- Apomys littoralis (Sanborn, 1952): Biliran ince, Mount Isarog, 5 km N, 20 km E Naga, Island, Leyte Province, 5 km N, 10 km E 900 m (13040'N, 12321'E), two males Naval, 850 m (1 1036'N, 124029'E), one male (USNM 573856 and 573857). (USNM 459842); Leyte Island, Leyte Prov- Rattus exulans (Peale, 1848): Negros Is- ince, Mount Pangasugan, 10.5 km N, 4 km land, Negros Oriental Province, Silliman E Baybay, 700-1000 m (10047'N, 124050'E), Farm, Dumaguete, 5 m (09018'N, 123018'E), one male, three females (USNM 458762, one female (USNM 459930); Mount Guin- 458764, 459853, and 459861). sayawan, 3 km N, 17 km W Dumaguete City, Apomys musculus Miller, 1911: Luzon Is- 1470 m (09°22'N, 123°09'E), four males, three land, Camarines Sur Province, Mount Isarog, females (USNM 458835, 458837, 458840, 4.5 km N, 20.5 km E Naga, 1125-1350 m 458841, 459931, 459932, and 459936).
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