亚洲始新世啮齿目一新科 ———争胜鼠科 ( Zelomyidae) 1)
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第 41 卷 第 4 期 古 脊 椎 动 物 学 报 pp. 249~270 2003 年 10 月 VERTEBRATA PALASIATICA figs. 1~9 亚洲始新世啮齿目一新科 ———争胜鼠科 ( Zelomyidae) 1) Dawson M. R. 1 黄学诗2 李传夔2 王伴月2 (1 美国卡内基自然历史博物馆古脊椎动物学部 匹茨堡 15213) (2 中国科学院古脊椎动物与古人类研究所 北京 100044) 摘要 继 1990 年王伴月和李春田记述了发现在吉林桦甸中始新世的争胜鼠 ( Zelomys) 之后 , 近年又在山西垣曲 、河南卢氏 、江苏溧阳等地的中始新世到晚始新世 ( Irdinmanhan2late Shara2 murunian or Ergilian) 的 5 个地点中陆续发现了一些与争胜鼠相近的新材料。经研究后 ,将所 有材料归诸于创建的一新科 :争胜鼠科 (Zelomyidae fam. nov. ) 。它包括了争胜鼠及本文新记 述的安氏鼠 ( Andersomys gen. nov. ) 、耗子 ( Haozi gen. nov. ) 和苏鼠 ( Suomys gen. nov. ) 共 4 属 6 种 。新科的特征是 :始啮型头骨 - 松鼠型下颌 ;门齿釉质层散系 ;颊齿具有发育的次尖和下次 脊 ,后期种类上颊齿的外侧齿尖的唇侧发展成平凹到新月形。新科的系统关系不很清楚 ,尽 管与始鼠科 ( Eomyidae) 有某些相似之处 ,但它更可能是独立发展的一个支系。 争胜鼠属化石分布较广 ,在我国吉林 、江苏和山西的中 、晚始新世地层中都有发现 ,它的 特征是下颊齿 p42m2 宽度逐渐增大 ,p42m3 具下前边尖 ,并与短的下原尖前臂相接。属中除属 型种 ———东方争胜鼠 ( Z. orientalis Wang et Li) 外 ,还建立了一个约翰争胜鼠新种 ( Z. joannes sp. nov. ) , 一约翰争胜鼠相似种 ( Z. cf. Z. joannes) 。同时认为 Z. gracilis Wang et Li ,1990 是 Z. orientalis 的同物异名。约翰种与属型种的区别在于个体稍大和下臼齿上缺少自下外脊伸 出的附脊。安氏鼠属化石发现在山西中始新世晚期及有可能是晚始新世地层中 ,其特征是 P4 和 p4 分别大于 M1 和 m1 ;P4 前尖和后尖及 M1 后尖唇侧平 ;与争胜鼠属不同在于前臼齿和 臼齿的大小比例和上颊齿具平的唇侧尖。安氏鼠包括两种 :老文种 ( A. laoweni sp. nov. ) 和 南堡头种 ( A. austrarx sp. nov. ) 。两种的不同在于后者个体稍大 ,P4 唇侧齿带较发育和 p4 前 缘更显圆滑 。耗子属仅有一种 ,简单耗子 ( H. simplex sp. nov. ) ,与科中其他属的不同在于颊 齿齿尖较呈丘形 ,P4 和 M1 大小接近 ,上颊齿的前附尖和后附尖很发育 ,P4 三角凹宽于跟凹 。 该属化石仅发现在河南中始新世晚期地层中。苏鼠属也只有新月苏鼠 ( S. selenis sp. nov. ) 一 种 ,发现在江苏中始新世晚期到晚始新世地层中。该属的特点是 P4 比其他上颊齿大 ,具强的 前齿带和唇侧齿带 ;P42M2 的前尖和后尖及 M3 的前尖唇侧凹平 ,且向前 、后延长 ,形成 W 形 外壁 ,中附尖明显 ;下颊齿的唇侧尖也呈 W 形结构 ,使牙齿为新月形式样 ,这是与科中其他属 明显不同的特征 。 文中还对含化石地层时代问题做了评述。 关键词 亚洲东部 , 始新世 , 争胜鼠科 中图法分类号 Q915. 873 1) 美国国家科学基金项目(编号 :DBS 9221231、DEB 9310509、SBR 9615557) ,中国科学院知识创新工程项目 (编号 : KKZCX32SW2127) 和国家自然科学基金项目(编号 :49372720 及 49772092) 资助。 收稿日期 :2003 - 04 - 23 052 古 脊 椎 动 物 学 报 41 卷 ZELOMYIDAE, A NEW FAMILY OF RODENTIA ( MAMMALIA) FROM THE EOCENE OF ASIA Mary R Dawson1 HUANG Xue2Shi2 LI Chuan2Kui2 WANG Ban2Yue2 (1 Section of Vertebrate Paleontology , Carnegie Museum of Natural History Pittsburgh , PA 15213 , USA) (2 Institute of Vertebrate Paleontology and Paleoanthropology , Chinese Academy of Sciences Beijing 100044) Abstract Six Eocene localities in eastern China , ranging geographically fromJilin to Jiangsu provinces and in age from Irdinmanhan to late Sharamurunian or Ergilian , have yielded four genera ( Haozi , Suomys , and Andersomys named as new here and Zelomys) assigned to the new rodent family Zelomyidae. The family is characterized by a primitive protrogomorphous2sciurognathous zygomasseteric structure and pauciserial incisor enamel , but the cheek teeth are precocious in development of hypocone and hypolophid. Later members of the family exhibit a development of buccal flattening to selenodonty of the cusps of the cheek teeth. Relationships of the new family are unclear , although some dental similarities suggest affinities with the Eomyidae. On the other hand , differences from eomyids may suggest that the zelomyids are an independently derived Asian Eocene family. Key words Eastern Asia , Eocene , Zelomyidae 1 Introduction The Eocene rodent record in Asia differs distinctly from that of the comparable time in Europe and North America. In Asia the Paleogene endemic ctenodactyloids dominate during the Early Eocene , followed by increasing prominence in the Middle Eocene of cricetids , eomyids , and zapodids (Tong , 1997) ; ischyromyids are present but seem never to have been dominant (Dawson and Wang , 2001) . In Europe , on the other hand , the Early Eocene dominance of ischyromyids is followed in the Middle Eocene by the origin of the presumably autochthonous theridomorphs and gliroids ( Escarguel , 1999) . The North American Paleogene faunas include dominant ischyromyids , but show the origin and increased diversity there by the late Early Eocene of the apparent ischyromyid derivatives , the Sciuravidae ( Korth , 1984) . The latter family is characterized by an early formation of quadrate upper molars , having a strongly developed hypocone , and lophate lower molars with a well developed hypolophid. In several Asian Eocene faunas there are rodents that have been referred definitely or with question to the otherwise North American family Sciuravidae (Li , 1963 ; Dawson , 1964 ; Shevyreva , 1976 ; Hussain et al. , 1978 ; Tong and Wang , 1980 ; Wang and Li , 1990) . Later , some of these have been shown to be ctenodactyloids (Dawson , 1977 ; Wood , 1977 ; Dawson et al. , 1984) . One , the rodent Zelomys (Wang and Li , 1990) , has been referred to the Eomyidae ( Korth , 1994) . Reexamination of Zelomys and several other of these problematic rodents , outside of the ctenodactyloids , has now shown that they represent a discrete rodent lineage , here recognized as the new family Zelomyidae. As currently known , six Asian Eocene localities have produced representatives of the new family , ranging in age from probably Irdinmanhan (middle Middle Eocene) through Ergilian (latest Eocene) . Their currently known geographic distribution encompasses southern Jiangsu , western Henan , southern Shanxi , and central Jilin provinces. Abbreviations and terminology : ALMA , Asian Land Mammal Age. IVPP , Institute of Vertebrate Paleontology and Paleoanthropology , Chinese Academy of Sciences , Beijing. MP , Paleogene Mammalian Reference Level ( European) . 4 期 Dawson M. R. et al. :亚洲始新世啮齿目一新科 ———争胜鼠科 (Zelomyidae) 152 NALMA , North American Land Mammal Age. a2p , anteroposterior length ; trw , transverse width of trigon ; taw , transverse width of talon ; tri , transverse width of trigonid ; tal , transverse width of talonid ; mm , millimeter. For the dental notation we use upper case letters for upper teeth and lower case letters for lower teeth ; e. g. M1 indicates an upper first molar , m1 a lower first molar. Hedi is the corrected Pinyin spelling for the formation in the Yuanqu Basin ( Huang et al. , 2001 ) , which has also been spelled Hoti ( Chow et al. , 1973 , using the Wade2Giles transliteration) and Heti (Li and Ting , 1983) . 2 Age and geologic setting All of the localities that have produced members of the new family occur in eastern China (Fig. 1) ; no zelomyids are currently repre2 sented in the relatively well known stratified deposits of Mongolia and Nei Mongol ( Inner Mongolia) on which the Asian Eocene land mammal ages have been based ( Russell and Zhai , 1987) . These rodents have been found in discrete , isolated basins (Lushi , Yuanqu , Huadian) as well as in the Shanghuang fis2 sures , Liyang , and in varying lithologies over a wide geographic area , ranging from the oil shales of central Jilin Province , at about 43° north latitude , to the fissure fillings of southern Jiangsu Province , at about 31°N. These geo2 graphic factors contribute substantially to the problem of correlating both between localities and into the Asian framework. Correlation by biostratigraphy is hampered by several factors , including probable differences in small mam2 mals over this latitudinal span due to paleocli2 matic/ environmental differences and in ecologi2 cal preference between , for example , animals that would be preserved in the oil shales of Jilin , the fresh water marls of Zhaili , and the Fig. 1 Outline map of the eastern part of the People’s karstic fissure fillings of Shanghuang. Republic of China , showing localities in which fossil A recent evaluation of Eocene faunas Zelomyidae have been found from localities in the Yuanqu Basin ( Huang et 1. Huadian Basin , Jilin Province ; 2~4. Yuanqu Basin , al. , 2001) , several of which contain members Shanxi Province ( Huoshipo , Zhaili , Nanbaotou faunas) ; of the new family , provided the following 5. Lushi Basin , Henan Province ( Chugouyu sequence from oldest to youngest : Huoshipo Fm. ) ; 6. Shanghuang fissure fillings , Liyang County , Jiangsu Province (Huang et al. , 1999 ; Zhang et al. , 2001 ; Irdinmanhan ALMA) ; Rencun ( Sharamurunian ALMA; no zelomyids have been recognized from this locality) ; Zhaili ( Naduan ALMA) ; and Nanbaotou ( Ulangochuan ALMA) . While this sequence is almost certainly correct , the correlation to ALMAs outside of the Yuanqu Basin must be viewed with a great deal of caution. All of these ALMAs except Naduan are based on sequences in Nei Mongol. Naduan is an Asian land mammal age that was established (Tong , 1989) on the basis of faunas of mostly large mammals from the Nadu Formation in the Bose Basin of Guangxi Zhuang Autonomous Region in southern China. The Caijiachong fauna of Yunnan Province (Wang , 1985 , 252 古 脊 椎 动 物 学 报 41 卷 1997) , which contains numerous small mammals and has been correlated on a global scale with the North American Chadronian NALMA , was also referred to the Naduan age ( Tong et al. , 1995) . The concept of a Naduan ALMA has been remarkably difficult to establish with precision. To avoid this difficulty , Russell and Zhai (1987) and Emry et al. (1998) have utilized the Ergilian ALMA for post2Sharamurunian , pre2Shandagolian Asian sites. This practice is followed here. The rodents from the Zhaili fauna , which was referred to the Naduan ALMA by Huang et al. (2001) , appear to be older than those of the Caijiachong fauna. The entire Caijiachong fauna may be included in the Ergilian “( older than both the Ardyn Obo local fauna in Asia and MP 21 of Europe”in Wang , 1992 , 1997 ; Emry et al. , 1998) . Russell and Zhai (1987) have already extended the concept of a Sharamurunian ALMA to include the Zhaili fauna , a use that is followed here , with the Zhaili fauna considered to be late Sharamurunian in age. The Nanbaotou local fauna , in the strict sense used here , comes from a limited