Pliocene – Early Pleistocene Arvicolids of North China: Taxonomy, Phylogeny, and Biochronology （中国北部における鮮新世・前期更新世のハタネズミ類 ̶その分類，系統および生物年代学） Yingqi ZHANG （張 穎奇） Pliocene – Early Pleistocene Arvicolids of North China: Taxonomy, Phylogeny, and Biochronology （中国北部における鮮新世・前期更新世のハタネズミ類 ̶その分類，系統および生物年代学） 理学研究科 生物地球系専攻 平成１９年度 Yingqi ZHANG （張 穎奇） CONTENTS ABSTRACT·················································································································································· 1 I. Introduction ············································································································································ 3 II. Significance of Arvicolid Research ································································································· 5 III. Mammal Chronology in Europe, North America and China ··············································· 6 A. Mammal Neogene zones of Europe (MN) and European Land Mammal Ages (ELMA) ················ 7 B. North American Land Mammal Ages (NALMA) ············································································ 8 C. Chinese Neogene Mammal Ages and Mammal Faunal Units (NMU) ············································· 9 D. What on earth is biochronology? ·································································································· 10 IV. A Review of Arvicolid Biochronology ························································································ 12 V. Introduction of Arvicolid Fossil Localities ················································································· 15 A. Bilike ·············································································································································· 15 B. Gaotege ·········································································································································· 15 C. Renzidong ······································································································································ 17 D. Lingtai ··········································································································································· 18 E. Xiaochangliang ······························································································································ 20 F. Other related localities and species ································································································ 22 VI. Systematic Descriptions ················································································································· 22 A. Descriptive terminology and measuring methods ········································································ 22 B. Calculating method for quantified characters ··············································································· 25 C. Systematic descriptions ················································································································· 25 Order RODENTIA ROCHEBRUNE, 1883 ······························································································ 25 Family CRICETIDAE GRAY, 1821 ····································································································· 25 Subfamily ARVICOLINAE GRAY, 1821 ························································································ 25 Tribe ARVICOLINI KRETZOI, 1954 ···················································································· 27 Genus Mimomys FORSYTH-MAJOR, 1902 ······································································ 27 Mimomys bilikeensis (QIU AND STORCH, 2000) ························································ 28 Mimomys cf. M. bilikeensis (QIU AND STORCH, 2000) ·············································· 32 Mimomys teilhardi LI, 2006 ····················································································· 33 Mimomys cf. M. orientalis YOUNG, 1935 ································································· 35 Mimomys gansunicus ZHENG, 1976 ········································································· 37 Genus Allophaiomys KORMOS, 1932 ············································································· 41 i Allophaiomys deucalion KRETZOI, 1969 ·································································· 42 Omniprocessimys gen. nov. ··························································································· 44 Omniprocessimys parallelus sp. nov. ······································································· 45 Genus Villanyia KRETZOI, 1956 ···················································································· 47 Villanyia sp. nov. ····································································································· 48 Villanyia fanchangensis ZHANG ET AL., in press ······················································ 49 Villanyia cf. V. fanchangensis ZHANG ET AL., in press ·············································· 53 Villanyia sp. 1 ·········································································································· 55 Villanyia sp. 2 ·········································································································· 57 Genus Borsodia JÁNOSSY AND VAN DER MEULEN, 1975 ·················································· 58 Borsodia chinensis (KORMOS, 1934) ········································································· 58 Borsodia sp. ············································································································· 60 Tribe MICROTINI KRETZOI, 1954 ····················································································· 62 Genus Proedromys THOMAS, 1911 ················································································ 62 Proedromys bedfordi THOMAS, 1911 ······································································· 62 Mimomys teilhardi - Villanyia sp. nov. COMPLEX ································································ 64 Mimomys cf. M. orientalis - Villanyia sp. 1 COMPLEX ·························································· 66 Proedromys bedfordi - Allophaiomys deucalion COMPLEX ··················································· 66 Arvicolinae gen. et sp. indet. ···································································································· 67 VII. Phylogenetic Analysis ··················································································································· 70 A. Selected characters and character coding ····················································································· 70 B. One intuitively acceptable phylogenetic hypothesis ······································································ 76 1. Character evolution reconstruction ···················································································· 76 2. Representative arvicolid lineages ························································································· 78 VIII. Pliocene ~ Early Pleistocene Arvicolid Biochronology of North China ······················· 81 A. e age of Renzidong from the “stage of evolution” view of fossil arvicolids ································· 82 B. e age of 93001 section, Lingtai from the “stage of evolution” view of fossil arvicolids ·············· 82 C. Pliocene ~ Early Pleistocene Arvicolid Biochronology of North China ········································· 85 1. Promimomys asiaticus-?>Mimomys bilikeensis->M. teilhardi zone ········································· 85 2. Mimomys orientalis zone ·········································································································· 86 3. Mimomys youhenicus->Omniprocessimys peii zone ································································· 87 4. Mimomys gansunicus-Villanyia fanchangensis-Omniprocessimys parallelus zone ·················· 87 5. Allophaiomys deucalion-Borsodia chinensis zone ···································································· 88 ii Acknowledgements ································································································································· 89 References ·················································································································································· 90 FIGURES [04] Figure 1. Pliocene ~ Early Pleistocene arvicolid fossil localities involved in this study [16] Figure 2. Gaotege section (A: after LI, 2006) and its magnetostratigraphic study result (B: modified from Fig. 7 of XU ET AL., 2007) [19] Figure 3. Lithostratigraphic correlation of three sections in the Leijiahe area, Lingtai, Gansu (after ZHENG AND ZHANG, 2001) [21] Figure 4. Columnar section and paleomagnetic data of the Xiaochangliang site (modified from ZHU ET AL., 2001) [23] Figure 5. Terminology and measuring methods for the occlusal surface of arvicolid molars (occlusal views after CARLS AND RABEDER, 1988) [24] Figure 6. Terminology and measuring methods for sinuous line of arvicolid molars (after CARLS AND RABEDER, 1988) [77] Figure 7. An intuitively acceptable hypothesis about the phylogenetics of the arvicolid taxa studied here [84] Figure 8. Pliocene ~ Early Pleisotcene arvicolid biochronology of North China TABLES [68] Table 1. Measurements of arvicolids from the Pliocene ~ Early Pleistocene of North China [71] Table 2.