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Mammalia, Rodentia) and Bilophodonty in Middle Eocene Asian Rodents

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Mammalia, Rodentia) and Bilophodonty in Middle Eocene Asian Rodents 第 卷 第 期 古 脊 椎 动 物 学 报 48 摇 4 pp.328-335 年 月 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 摇 2010 10 VERTEBRATA PALASIATICA 摇 figs.1-3 硅藻鼠科( 哺乳纲,啮齿目) 与亚洲中 始新世啮齿类的双脊齿 1 李传夔2 齐 陶2 M美ar国y卡R内基. 自D然A历W史博S物O馆N古脊摇 椎动物学部 匹摇兹堡 摇 (1 摇 摇 PA 15213) 中国科学院古脊椎动物与古人类研究所 北京 (2 摇 摇 100044) 摘要 亚洲特有的啮齿类硅藻鼠科自渐新世以来分布于东亚和南亚 现生硅藻鼠类的分布只 : 。 限于老挝的喀斯特地区 就目前所知 这些具有豪猪型头骨 松鼠型下颌的啮齿类的颊齿都 。 , - 是不同程度的横向双脊齿 时代最早的硅藻鼠类产于巴基斯坦渐新世地层中 其颊齿的双脊 。 , 齿构造上仍保留齿尖残迹 基本符合双脊齿型牙齿结构 至渐新世末期 硅藻鼠科的牙齿出 , 。 , 现分化 中新世及以后硅藻鼠类的化石记录相对较少 分子生物学证据将硅藻鼠类归入 。 。 这种归属也从始新世梳趾鼠类的臼齿形态上得到一定的支持 除此之外 有关 Ctenohystrica, 。 , 硅藻鼠类的起源问题几乎一无所知 亚洲中始新世的 Hydentomys 臼齿表现出轻微的双脊型 。 , 然而其他方面却与硅藻鼠类不同 另一个具双脊齿的啮齿类 Dolosimus 新属 产于江苏中始 。 ( ) 新世 其具有更为发育的双脊齿 特别是臼齿型下牙 新属的不完整记录及其形态不能解决 , , 。 如下问题 它是否与后来出现的像硅藻鼠类和跃兔类这些具有明显双脊齿型颊齿的啮齿类有 : 亲缘关系 或者只是这种形态发育中没有留下后继者的早期试验品 , 。 关键词 亚洲 始新世 啮齿目 硅藻鼠科 双脊齿型颊齿 : , , , , 中图法分类号 文献标识码 文章编号 :Q915. 873摇 :A摇 :1000-3118(2010)04-0328-08 THE DIATOMYIDAE (MAMMALIA, RODENTIA) AND BILOPHODONTY IN MIDDLE EOCENE ASIAN RODENTS 1 2 2 Vertebrate Paleontology MCarnyegRie. MDusAeuWm SofONNatu摇ralLHI isCtohryuan鄄Kui 摇 QI Tao (1 , 4400 Forbes Ave, Pittsburgh PA 15213 USA 摇 dawsonm@ carnegiemnh. org) Institute of Vertebrate Paleontology and Paleoanthropology Chinese Academy of Sciences (2 , Beijing 100044 China) Abstract 摇 The Asian endemic rodent family Diatomyidae is known to have inhabited eastern and south鄄 ern Asia since the Oligocene. Its current distribution is limited to karstic regions of Laos. So far as known these hystricomorphous鄄sciurognathous rodents have some degree of transverse bilophodonty of the cheek teeth. The earliest recognized diatomyids, which are from the Oligocene of Pakistan, retain some traces of cusps on the cheek teeth, overlying the basically bilophodont tooth structure. By the end of the Oligocene there is some dental diversity within the family. Miocene and later diatomyids are relatively rare in the fossil record. Molecular evidence unites the diatomyids in the Ctenohystrica, an assignment that receives some support from the molar morphology of Eocene ctenodactyloids. OtherHtyhdaennttohmisyscon鄄 nection, little is clear regarding the origin of the diatomyids. The middle Eocene Asian ex鄄 hibits a slight degree of bilophodonty, but is otherwise unlike diatomyids. Another taxon of bilophodont 收稿日期 :2010-02-24 期 Dawson et al.: The Diatomyidae (Mammalia, Rodentia) and bilophodonty 4 摇 329 in middle Eocene Asian rodents Dolosimus rodent, n. gen. from the middle Eocene of Jiangsu Province, has still more precocious de鄄 velopment of bilophodonty, especially in the lower molariform teeth. The incomplete record of this new taxon as well as its morphology cannot answer the question of whether this taxon is allied to such later appearing, strongly bilophodont rodents as diatomyids and pedetids, or is an early experiment of Ktheisyswtriokridnsg morphological development that left no successors. 摇 Asia, Eocene, Rodentia, Diatomyidae, bilophodont cheek teeth 1摇 Introduction The family Diatomyidae Mein & Ginsburg, 1997, encompasses Asian rodents that are characterized by their combination of an hystricomorphous / sciurognathous zygomasseteric struc鄄 ture; multiserial incisor enamel; dental formula 1 / 1, 0 / 0, 2 -1 / 1, 3 / 3; transversely bilopho鄄 dont cheek teeth lacking conules; premolars usually having 3 roots, molars 4 roots; relatively low temporomandibular joint; reduced coronoid process; and a relatively unspecialized postcra鄄 nial skeleton. Their fossil record, comprehensively survFeaylelodmbuys Flynn (2006), extends into the Paleogene of the Indian subcontinent, with the genus (FlynnFaeltloaml.,us19F86. ;raMzareivaFu.x ganindsbWureglicomme,F.20q0u3r)ai.shTyhiree Oligocene species (28 or 25 Ma) of , , , and Fa,llohmavues clhaedeakkhtenesthis that are bilophodont but relatively more cuspate than in later diatomyids. Nanda & Sahni, 1998, known by lower teeth and jaws from the late Oligocene of northernFaInlldoimauasnd Thailand, is more hypsodont and more distinctly bilophodont than other species of as well as having an enlarMgeadryhmyupsoconulid forming a third, posterior loph on the lower cheek teeth (Marivaux et al., 2004). Flynn, 2007, known from isolated teeth from the later Oligocene (26 ~ 27 Ma) or early Miocene (23 ~ 24 Ma) of Pakistan (de Bruijn et al., 1981; Flynn 2007), is more strictly bilophodont in its upper and lower molar morphology but has large, cuspate premolars. The later DNieaotogmenyes record of diatoWmyilildmsusstems from southern and eastern Asia and includes the Miocene Lai,on1a9s7te4s and Flynn & Morgan, 2005. The single Recent representative of the family, , appearsDtioatboemryesstricted to karstic areas of Laos (Dawson et al., 2006). The Miocene is currently the best known extinct member of its family, being re鄄 presented by beautifully preserved, though compressed, fossils from the diatomaceous lacustrine shales of the Shanwang locality in Shandong ProDviantcoem,yCs hsihnaan,tulnatgeenesairsly Miocene to early mid鄄 dle Miocene (14 ~ 17 Ma; Qiu et al., 1999). was originally based on two relatively complete skeletons (Li, 1974). The well preserved dentitions of these specimens show that the cheek teeth in each jaw quadrant include one premolar and three molars displa鄄 ying a simple, transversely bilophodont occlusal pattern. Details of cranial and mandibular Daniatomyswere difficult to interpret because of the lateral compression of the fossils. However, was originally thought to have a non鄄enlarged infraorbital foramen (the sciuromorphous condition) and a sciurognathous lower jaw. Its postcranial skeleton lacks any obvious morpho鄄 logical adaptations for either leaping or burrowing. D. shantungensis In June 2005, a new and less compressed specimen of was discovered from the type locality in Shandong Province. The new specimen, IVPP V 12692 ( Fig. 1), complete wiDthiawtohmisyksers and traces of pelage, clarifies aspects of the cranial and mandibular anatomy of that were either missing or obscured by postmortem deformation in Dpiraetvoi鄄 mouyssly described specimens oDfiathtoismsypsecies. V 12L6a9o2nashstoewss that the infraorbital foramen of is very large; hence, resembles in having the hystricomorphous condi鄄 tion. V 12692 also shows that the mandible lacks a coronoid process and has a relatively low condyle; the masseteric fossa extends forward to a level below p4; an anteroposterior ridge, the linea obliqua, separates dorsal and ventral portions of the masseteric fossa; the angular process is in the same vertical plane as the incisor (the sciurognathous condition); the angular process extends posteriorly as far as the condylar process; the ventral side of the angular process is very 古 脊 椎 动 物 学 报 卷 摇 330 摇 摇 摇 摇 摇 摇 48 Diatomys shantungensis Fig. 1摇 , IVPP V 12692, complete with whiskers and traces of pelage, from the Shanwang locality, Shandong Province, China, late early Miocene to early middle Miocene A. lateral view; B. outline drawing with some parts identified Abbreviations: cl. clavicle; fe. femur; h. humerus; lh. left humerus; lm. left mandible; lv. lumbar verte鄄 brae; ma. manubrium; pe. pelvis; ri. ribs; rm. right mandible; ru. radius + ulna; s. scapula; sc. sternal cartilage; sk. skull; st. sternum. tv. thoracic vertebrae; vi. vibrissa slightly inflected; and the incisor root is short, extendDinigatpoomsytesriorlyLtaooanalesvteesl below m2. Thus, the new specimen demonstrates that the mandibles of and are virtually iden鄄 tical in sharing the following derived characters: absence of a coronoid process, masseteric fossa extending forward to below p4, masseteric fossa subdivided into dorsal and ventral sections, condyle low but higher than tooth row, and shortened incisor. 期 Dawson et al.: The Diatomyidae (Mammalia, Rodentia) and bilophodonty 4 摇 331 in middle Eocene Asian rodents In addition to the fossil vertebrates from the Shanwang locality, fossils of abuDnidaatnotmgyrsasses, dicotyledenous plants, and infrequent palms, suggest that the habitat in which lived had a humid, warm temperate to sub鄄tropical climate, more humid and equable than present (Stromberg et al., 2007). Fallomus There is enougMhardyemntuasl diversity within species of the Oligocene diatomyid and between them and to suggest a considerably earlier origin for the family. So far, there has been little support for details of this origin other than the presence throughout much of Asia of a variety of ctenodactyloids (Fig. 2E, F), an old Asian clade having living representatives only in Africa ( Flynn et al., 1986; Marivaux and Welcomme, 2003; Dawson et al., 2006). Such a relationship is favored by morphological evidence from fossil and living diatomyids as well as molecular evidence (Huchon et al., 2007) that lends support for the Ctenohystrica, a clade encompassing Diatomyidae, Ctenodactylidae, and the Hystricognathi. 2摇 Bilophodonty in Rodentia In spite of their reasonable fossil re鄄 cord and moderately secure affinities based on morphological and molecular evidence, the origin of the Diatomyidae remains enig鄄 Dmiaattiocm. yBsilophodont teeth similar to those of have evolved a number of times within the Rodentia, as emphasized by Li (1974) in his original description of the genus, in which he mentioned this condi鄄 tion among geomyoids and pedetids. While bilophodonty is not restricted to these ro鄄 dents, it is a frequently appearing charac鄄 teristic in them. Perhaps the best docu鄄 mented series of changes among fossil ro鄄 dents from a more cuspate occlusal pattern to a bilophodont one has been documented in the transition withinEtohmeyEs uropRehaondagneo鄄 mysoid ERoimtteynideraiea from
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