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A Systematic Review of Sulawesi Bunomys (Muridae, Murinae) with the Description of Two New Species Guy G. Musser

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MUSSER: SULAWESI Scientific Publications of the American Museum of Natural History American Museum Novitates A S YSTEMATIC R EVIEW OF SULAWESI Bulletin of the American Museum of Natural History Anthropological Papers of the American Museum of Natural History BUNOMYS (MURIDAE, MURINAE) WITH THE Publications Committee ESCRIPTION OF WO EW PECIES Robert S. Voss, Chair D T N S Board of Editors Jin Meng, Paleontology Lorenzo Prendini, Invertebrate Zoology BUNOMYS GUY G. MUSSER Robert S. Voss, Vertebrate Zoology Peter M. Whiteley, Anthropology Managing Editor Mary Knight Submission procedures can be found at http://research.amnh.org/scipubs All issues of Novitates and Bulletin are available on the web (http://digitallibrary.amnh. org/dspace). Order printed copies on the web from: http://shop.amnh.org/a701/shop-by-category/books/scientific-publications.html or via standard mail from: American Museum of Natural History—Scientific Publications Central Park West at 79th Street New York, NY 10024 This paper meets the requirements of ANSI/NISO Z39.48-1992 (permanence of paper). AMNH BULLETIN 392 On the cover: Dense, long, and silky-soft fur, brownish- gray upperparts, grayish-white underparts, gray ears, white feet, and a bicolored tail characterize Bunomys penitus, one of eight documented species of Bunomys, all endemic to forested landscapes on the Indonesian island of Sulawesi. Nocturnal and terrestrial, B. penitus lives only in mountain forests where the ambience is cool and wet, the trees and 2014 ground covered with thick moss and epiphytes; its diet in - cludes invertebrates, fungi, and fruit. BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY A SYSTEMATIC REVIEW OF SULAWESI BUNOMYS (MURIDAE, MURINAE) WITH THE DESCRIPTION OF TWO NEW SPECIES GUY G. MUSSER Division of Vertebrate Zoology (Mammalogy) American Museum of Natural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 392, 313 pp., 108 figures, 82 tables Issued December 30, 2014 Copyright E American Museum of Natural History 2014 ISSN 0003-0090 DEDICATION I dedicate this review of Bunomys to the memories of two men of great intelligence and abilities whose sustained sponsorship and unflagging support of my research made possible my survey of endemic murines in the northern part of the west-central region of Sulawesi. Dr. Sampurno Kadarsan, a parasitologist specializing in tick taxonomy, was Director of the Museum Zoologicum Bogoriense during the period I lived in Indonesia. In addition to representing the museum as my sponsor in his country, Dr. Kadarsan (‘‘please call me Sam,’’ he asked) provided access to the Museum’s resources, a place in the Bogor gardens to stay when not in the field, the frequent hospitality of his home, and lively conversations covering a range of topics from science to Indonesian history (see my tribute in the etymology of Kadarsanomys, a Javan endemic murine; Musser, 1981: 32). Dr. P.F. Dirk Van Peenen, a Navy physician and epidemiologist, was in charge of Naval Medical Research Unit No. 2 (NAMRU-2), Jakarta Detachment. We first met at the American Museum of Natural History when Dirk visited to examine Southeast Asian mammal specimens for a handbook on Vietnamese mammals he was preparing. I helped with some of the taxonomic problems, and later identified mammalian hosts collected on epidemiological surveys made in Indonesia by Dr. Van Peenen and his team. Knowing of my desire to begin fieldwork on Sulawesi, Dirk invited me to use the research facilities NAMRU-2 had organized in the village of Tomado, on the shore of Danau Lindu in central Sulawesi, as the starting point for my fieldwork and provided logistical and other kinds of assistance whenever needed, an arrangement heartily condoned by Sam Kadarsan. Through circumstances I could not have anticipated when I joined the Department of Mammalogy at the American Museum of Natural History, a Javanese native and Indonesian scientist united with an American physician and dedicated epidemiologist to facilitate my journey into the tropical rain forests of Sulawesi. 2 CONTENTS Dedication......................................................... 2 Abstract.......................................................... 7 Introduction....................................................... 7 MaterialandMethods............................................... 9 HistoricalSummary................................................. 23 TheGenus........................................................ 26 Bunomys Thomas ................................................. 26 TheSpecies....................................................... 47 The Bunomys chrysocomus Group..................................... 55 GazetteersandSpecimensExamined.................................. 55 Bunomys chrysocomus ............................................ 58 Holotype................................................ 58 TypeLocality............................................. 59 EmendedDiagnosis........................................ 61 Geographic and Elevational Distributions......................... 61 Sympatry with Other Bunomys................................. 69 Description . ............................................. 71 Karyotype............................................... 74 Comparisons ............................................. 74 GeographicVariation....................................... 74 Natural History ........................................... 84 Ectoparasites, Pseudoscorpions, and Endoparasites .................. 97 Synonyms............................................... 99 Subfossils............................................... 103 Bunomys coelestis............................................. 106 Holotype............................................... 107 TypeLocality............................................ 107 EmendedDiagnosis....................................... 107 Geographic and Elevational Distributions........................ 107 Sympatry with Other Bunomys................................ 109 Description . ............................................ 109 Karyotype.............................................. 110 Comparisons ............................................ 110 GeographicVariation...................................... 115 Natural History .......................................... 115 Ectoparasites............................................ 116 Synonyms.............................................. 116 Bunomys prolatus ............................................. 116 Holotype............................................... 116 TypeLocality............................................ 117 EmendedDiagnosis....................................... 117 Geographic and Elevational Distributions........................ 117 Sympatry with Other Bunomys................................ 120 Description . ............................................ 120 Karyotype.............................................. 121 Comparisons ............................................ 121 Natural History .......................................... 123 Ectoparasites............................................ 123 Synonyms.............................................. 124 Bunomys torajae, new species ...................................... 124 Holotype............................................... 124 TypeLocality............................................ 124 3 4 BULLETIN OF THE AMERICAN MUSEUM OF THE NATURAL HISTORY NO. 392 Diagnosis............................................... 124 ReferredSpecimens........................................ 124 Geographic and Elevational Distributions........................ 124 Sympatry with Other Bunomys................................ 125 Etymology.............................................. 125 Description............................................. 125 Karyotype.............................................. 127 Comparisons............................................ 127 Geographic Variation ...................................... 135 NaturalHistory.......................................... 135 Ectoparasites, Pseudoscorpions, and Endoparasites . ................ 136 Synonyms.............................................. 136 Summary of Contrasts among Members of the Bunomys chrysocomus Group. 136 The Bunomys fratrorum group..................................... 139 Gazetteers and Specimens Examined ............................... 139 Bunomys fratrorum............................................ 148 Holotype............................................... 148 TypeLocality............................................ 148 EmendedDiagnosis....................................... 148 Geographic and Elevational Distributions........................ 149 Sympatry with Other Bunomys................................ 149 Description............................................. 149 Karyotype.............................................. 153 Comparisons............................................ 153 Geographic Variation ...................................... 163 NaturalHistory.......................................... 167 Ectoparasites............................................ 167 Synonyms.............................................. 169 Bunomys andrewsi........................................... 169 Holotype............................................... 169 TypeLocality............................................ 169 EmendedDiagnosis....................................... 169 Geographic and Elevational Distributions........................ 171 Sympatry with Other Bunomys...............................
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