硅藻鼠科(哺乳纲,啮齿目)与亚洲中始新世啮齿类的双脊齿(英文)

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Ctenohystrica,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。     

日本西部渐新世-中新世啮齿类及其地质古生物学意义

日本九州渐新－中新世沉积物中新发现的啮齿类,有四种类型：1．河狸科未定属种A;2．晚渐新世Sasebo（佐世保）群的Steneofibersp.;3．河狸科未定属种B;4．早－中中新世Nojima（野岛）群的山东硅藻鼠（Diatomysshantungensis）。河狸科的两个未定属种的标本属于大型河狸,头骨大小接近于巨河狸（Trogontherium）,门齿珐琅质具有明显的纵沟。个体大小和门齿珐琅质特征方面的相似性表明这些标本可能为同一个属。Steneofiber是一个遍布北半球的属,以前还未曾在东亚的晚渐新世地层中发现过。山东硅藻鼠的出现使我们有理由推测,在早中新世晚期,位于东亚边缘的日本和中国东部的哺乳动物同属一个区系。     

始鼠化石在甘肃省党河地区上渐新统的发现

<正>始鼠科(Ｅｏｍｙｉｄａｅ)是一类已绝灭的小型的鼠类,在古近纪和新近纪时广布全北区,以北美(中始新世—晚中新世)和欧洲(早渐新世—晚上新世)发现的化石较丰富。过去在亚洲发现的始鼠化石极少。亚洲的第一个始鼠化石是１９８２年才发现的(郑绍华等,１９８２)。２０世纪９０年代以来,在亚洲陆续发现了一些始鼠化石,表明从中始新世到上新世时在亚洲已有始鼠生存过(Ｗａｎｇ ａｎｄ Ｅｍｒｙ,１９９０;Ｑｉｕ,１９９４,１９９６;Ｔｏｍｉｄａａｎｄ Ｓｅｔｏｇｕｃｈｉ,１９９４;Ｅｍｒｙ ｅｔ     

临夏盆地早中新世上庄组的巨獠犀门齿化石(英文)

记述了在临夏盆地早中新世地层中发现的兰州巨獠犀(Aprotodon lanzhouensis)的下门齿化石,其特点为非常粗壮并强烈弯曲。新材料的发现使巨獠犀在临夏盆地的延续时代跨越渐新世/中新世界线的推测得到完全证实。巨獠犀分布的地质时代和地理范围与巨犀重合,但巨獠犀的化石地点和个体数量都相当稀少。巨獠犀的下颌形态功能特点指示其生活于晚始新世至早中新世中国西北、南亚和中亚干旱环境地带中镶嵌分布的少量近水环境。巨獠犀在中中新世之前彻底绝灭,其原因可能是气候变化的结果,也说明临夏盆地早中新世的环境特征与晚渐新世的疏林系统相似,而不同于中中新世的茂密森林。     

云南禄丰、元谋晚中新世古猿地点始鼠科化石

<正>始鼠科(Eomyidae)是一类绝灭了的啮齿类动物,渐新世和中新世时广布全北区,但亚洲远没有欧洲和北美常见。中国含始鼠类化石地点不多,发现的属只有渐新世的Eomys、Eomyodon和Pseudotheridomys,以及中新世的Keramidomys和Leptodontomys(Zheng and Li,1982;Falhbusch et al．,1983;Wang and Emry,1991;Qiu,1996;Wang,2002)。本文描述的云南始鼠类化石,系1983年在禄丰石灰坝和1999、2000年在元谋雷老两个古猿地点采集到的。材料不多,但至少代表始鼠科的两个新属,为我国惟一采自南方,并与古猿类共生的稀少啮齿类动物。新种的模式产地均为禄丰石灰坝。     

南海北部琼东南盆地渐新世-上新世沟鞭藻指示的沉积环境北大核心CSCD

通过对南海北部琼东南盆地渐新世—上新世3个钻井剖面412个沟鞭藻样品的系统分析,发现了较丰富的沟鞭藻化石。根据沟鞭藻化石丰度、分异度变化以及特征性的环境指示种,对琼东南盆地渐新世至上新世的沉积环境进行了详细研究。认为研究区从早渐新世早期海水开始入侵,至早渐新世晚期海侵范围进一步扩大,一直持续到晚渐新世;早中新世沟鞭藻化石数量和种类明显出现低谷,发生了一次较为明显的海退;中中新世至晚中新世,沟鞭藻化石较丰富,丰度和分异度增加,海水明显比早中新世沉积时期加深;早上新世早期,指示海侵的化石属种较丰富,反映了温暖浅海的沉积环境;早上新世晚期,指示一种开阔的海洋环境,水体较深;晚上新世早期,沟鞭藻化石属种面貌反映了热带浅海环境,但水体可能比早上新世晚期沉积时要浅;晚上新世晚期,指示海侵的沟鞭藻化石属种达到最繁盛时期,海侵范围进一步扩大。     

南海北部琼东南盆地渐新世—上新世沟鞭藻指示的沉积环境

通过对南海北部琼东南盆地渐新世—上新世3个钻井剖面412个沟鞭藻样品的系统分析,发现了较丰富的沟鞭藻化石。根据沟鞭藻化石丰度、分异度变化以及特征性的环境指示种,对琼东南盆地渐新世至上新世的沉积环境进行了详细研究。认为研究区从早渐新世早期海水开始入侵,至早渐新世晚期海侵范围进一步扩大,一直持续到晚渐新世;早中新世沟鞭藻化石数量和种类明显出现低谷,发生了一次较为明显的海退;中中新世至晚中新世,沟鞭藻化石较丰富,丰度和分异度增加,海水明显比早中新世沉积时期加深;早上新世早期,指示海侵的化石属种较丰富,反映了温暖浅海的沉积环境;早上新世晚期,指示一种开阔的海洋环境,水体较深;晚上新世早期,沟鞭藻化石属种面貌反映了热带浅海环境,但水体可能比早上新世晚期沉积时要浅;晚上新世晚期,指示海侵的沟鞭藻化石属种达到最繁盛时期,海侵范围进一步扩大。     

伊朗新发现的中新世哺乳动物足迹的亲缘关系及其意义

伊朗晚中新世的几个地点新近发现了哺乳动物足迹化石。足迹化石在伊朗中北部分布在一套被称为上红组的厚层、混杂的泻湖-陆相地层序列的数个层位中;在伊朗北部的里海地区南部,则位于一套陆相沉积序列中。产自上红组的足迹以偶蹄动物类型为主,由于个体很小,可归入几种像羚羊一样大小的种;其他足迹则分别归入小型、中型和大型的鼬科和猫科食肉动物。产自伊朗北部的足迹化石主要为长鼻类,有些可能是犀牛,少数为偶蹄类足迹。根据上红组中发现的猫科动物(剑齿虎)足迹,可认为产足迹的地层年代为晚中新世,而产于伊朗北部古地中海边缘区沉积中的长鼻类足迹显示,地层的最大年龄为中新世最早期。这些足迹化石的发现填补了这一重要地区新近纪哺乳动物化石记录的空白。     

新疆准噶尔盆地北缘第三纪地层古生物研究新进展

近两年对准噶尔盆地北缘第三纪地层进行了进一步调查和研究,在索索泉组和哈拉玛盖组内发现了新的化石层位。对新发现的哺乳动物的研究表明在勒布勒津地区索索泉组顶部的沉积物时代为中中新世,下部的沉积物时代不晚于晚渐新世,中新世与渐新世地层的界线应在索索泉组内寻找。新发现的动物群分子Cricetodonsp．nov．和Tachyoryctoidessp．nov.指示哈拉玛盖动物群的时代应重新考虑,有可能早于丁家二沟动物群。     

内蒙古中部敖尔班地区的岩石及生物地层(英文)

中国北方新近纪地层中,早中新世的哺乳动物化石地点比较稀少,化石面貌也不十分清楚。在内蒙古中部,尽管通古尔台地上有研究历史近百年的新近纪经典地点,但上世纪90年代发现的嘎顺音阿得格,依然是内蒙古目前惟一产出早中新世动物群的小盆地。同内蒙古诸多其他新近纪哺乳动物群一样,嘎顺音阿得格动物群与相邻动物群彼此缺乏时空上的联系,还不易建立可靠的层序关系。位于苏尼特左旗东南约60 km的敖尔班(曾用名"奥尔班",见Liddicoat et al.,2007),红色地层大面积出露,是2004年发现的一个新的哺乳动物化石地点。敖尔班剖面总厚50余米,大、小哺乳动物化石共生,岩性特征易识别,有利层位对比,是研究生物地层学的理想地点。自2004年以来,我们连续在敖尔班地区采集化石并进行地层工作,建立了一个较完整的哺乳动物序列,时代跨越早中新世晚期到晚中新世晚期。更加难得的是,敖尔班同一剖面上含有4个哺乳动物化石层位,其上下层序关系一目了然,在内蒙古新近纪地层中仅此一例。敖尔班哺乳动物序列的建立,无疑将对整个内蒙古中部新近纪,尤其是早中新世动物群面貌的了解具有促进作用。本文着重对敖尔班岩石地层进行描述,并结合我们4年来对脊椎动物化石的积累与认识,试图进一步完善内蒙古中部地区的生物地层层序,进而建立这一地区的年代地层框架。对化石的详细描述将适时发表。根据岩性及接触关系,敖尔班剖面可分为三大段:敖尔班组、巴伦哈拉根层及必鲁图层。敖尔班组(新建组)由一套红色和绿色的泥岩及粉砂岩组成,发育有成熟的古土壤层,厚约42 m,时代属于早中新世中晚期。敖尔班组的层型剖面建立在敖尔班露头出露最厚的中部,下部未见底,顶部与上覆巴伦哈拉根层呈假整合或不整合接触。该组目前已知分布仅局限于敖尔班露头。敖尔班组可进一步划分出三段:下红泥岩段、中绿泥岩段及上红泥岩段。三段呈连续沉积。敖尔班组与巴伦哈拉根层之间的假整合所代表的沉积间断延续了中中新世的大部分时段。巴伦哈拉根层为一套橘红色砂岩、粉砂岩及底砾岩,时代大致是最晚中中新世至最早晚中新世。不整合于巴伦哈拉根层之上是必鲁图层。两层之间似乎缺失了晚中新世的大部分堆积。必鲁图层的底砾岩是一种切割与充填构造,其中所含钙质结核及相当数量的化石都可能是巴伦哈拉根层原生堆积物再沉积的结果。必鲁图层的分布还需做更多工作,其时代可能是晚中新世晚期。主要依据小哺乳动物的组合,在敖尔班剖面中可建立4个动物群。最早为敖尔班组下红泥岩段产出的下敖尔班动物群。该动物群的特征是,小哺乳动物中在渐新世十分兴旺的一些科,如Ctenodactylidae,Tachyoryctoididae,Aplodontidae和Zapodidae还相当繁荣;中新世出现的属,如Mioechinus,Keramidomys,Heterosminthus和Democricetodon等占动物群总量的半数以上;大哺乳动物中残留有Palaeogale和裂爪兽。比下敖尔班动物群稍晚的是上敖尔班动物群,产出于敖尔班组的上红泥岩段。上敖尔班动物群的特征是,小哺乳动物在渐新世中占统治地位的一些科或完全绝迹,如Ctenodactylidae,或在种类和数量上明显减退,如Aplodontidae和Zapodidae;缺少在下敖尔班动物群中还相当常见的一些古老属,如Amphechinus,Tachyo- ryctoides和Sinolagomys等;出现了下敖尔班动物群中所没有的Megacricetodon,Cricetodon和Alloptox属;大哺乳动物中出现了长鼻类和柄杯鹿(Ligeromeryx/Lagomeryx)。经过相当长的一个沉积间断,敖尔班剖面的上部出现了巴伦哈拉根动物群。该动物群中渐新世常见的小哺乳动物科进一步衰落,同时出现了亚洲古北界晚中新世以后常见的跳鼠科(Dipodidae)和鼢鼠科(Siphneidae),具有明显的中中新世晚期或晚中新世早期生物组合的特点。最后是敖尔班剖面顶部必鲁图层中的必鲁图动物群。必鲁图动物群显然带有中中新世及晚中新世的混合特征,很可能是水流作用再沉积的结果。根据其中最进步分子的成分判断,估计必鲁图动物群的年代是晚中新世的晚期,其特征是,鼠科(Muridae)动物高度分化,而野兔科(Leporidae)尚未出现;含有晚中新世宝格达乌拉动物群或最晚中新世二登图动物群中大量出现的属种,如Lophocricetus grabaui,Paralactaga suni,Dipus fraudator和Hansdebruijnia pusilla等;但二登图动物群中很繁荣的一些属在必鲁图动物群中未被发现或者发现的个体数量很少,如Prospermophilus, Paralophicricetus和Microtodon等。     

Tracing the Origin and Diversification of Dipodoidea (Order: Rodentia): Evidence from Fossil Record and Molecular Phylogeny

Dipodoidea are a diverse rodent group whose earliest known record is from the Middle Eocene. The evolution and diversification of this superfamily have been documented by fossils and comparative morphology, but have not yet been studied from the perspective of molecular phylogeny. This study is the first attempt to reconstruct an extensive phylogeny of Dipodidae and estimate divergence times based on a nuclear gene coding for interphotoreceptor retinoid-binding protein. We found that there is a wide measure of agreement with the fossil record. Each of the three ecological groups of the extant Dipodoidea (sicistines, zapodines, and jerboas) has its distinctive distribution; the distribution patterns have been shaped by the dispersal events. The key events of paleogeographic distribution coincided with major paleoenvironmental events in the Cenozoic. The first important diversification phase occurred during the period from the Oligocene to Early Miocene, when global climate underwent major changes beginning with the Eocene/Oligocene boundary. The second adaptive radiation occurred within jerboas and was associated with the expansion of open habitat starting with the late Middle Miocene. The diversification of jerboas can be correlated with habitat changes in response to global and regional climatic events.     

Scolecophidia (Serpentes) of the Late Oligocene and Early Miocene,North America,and a fossil history overview

An abundant fossil record of the snake clade Scolecophidia exists in Europe; however, the minute snake is noticeably absent in reports about the North American Paleogene and Neogene. Presented here are four localities from Florida, USA, that contain scolecophidian remains older than the Pleistocene: Thomas Farm (late Early Miocene, Hemingfordian Land Mammal Age, LMA), Live Oak (Oligocene-Miocene transition, latest Arikareean LMA), White Springs 3B (late Arikareean LMA), and Brooksville 2 (Late Oligocene, middle Arikareean LMA). These remains extend their known existence by about 26 m.y. and are now the oldest reported scolecophidian remains in North America. Molecular evidence on extant scolecophidians concludes that these tiny snakes have a Gondwanan origin. Interestingly, the oldest record of a scolecophidian is from Europe (Belgium) and dates back to the middle Paleocene (MP 1–5). The earliest African record of the snake clade comes from the Paleocene-Eocene boundary in Morocco. The clade is apparently absent from Europe and Middle East deposits dating from the latest Eocene through to the latest Oligocene (MP 19–30) and to the Early Miocene (MN 4). A portion of this time is known as the booid ‘Dark Period’ which represents an apparent response to global aridization and cooling. Scolecophidians appear to re-emerge into the southern Eurasian record in the Early Miocene (MN 4) and become widely dispersed throughout Europe and Middle East. The fossil record of these minute snakes is largely absent in southern Asia and South America. It is possible that the current lack of a decent fossil scolecophidian record outside of Europe and Middle East is due mainly to a bias in the methodology to recover fossils; wet sieving sediments through < 1.0 mm mesh is needed to recover the minuscule vertebrae.     

DISCOVERY OF APLODONTIDAE (RODENTIA, MAMMALIA) FROM MIDDLE OLIGOCENE OF NEI MONGOL, CHINA

<正> Today only a single species of the aplodontid rodents still survives in the humid region of western North America, but in the Oligocene time the family was widespread over the Northern Hemisphere. Fossils of the Oligocene aplodontids so far have been mainly found in North America and Europe. Their occurrence in Asia is scarce. Only a few examples are known from Kazakhstan, USSR, and Mongolia. No record was known in China until 1977, when some fossil aplodontids were collected from the Middle Oligocene in Nei Mongol.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.     

Systematics and evolutionary significance of the small Abrocomidae from the early Miocene of southern South America

Octodontoidea is the most species-rich clade among hystricomorph rodents, and has a fossil record going back to at least the late Oligocene. Affinities of fossils previous to the late Miocene differentiation of the extant families Abrocomidae, Echimyidae and Octodontidae are controversial, essentially because these fossils may share few apomorphies with modern species. In fact, pre-late Miocene representatives of Abrocomidae had not been recognised until very recently. Here we revise the early Miocene genus Acarechimys, originally assigned to Echimyidae, and alternatively to stem Octodontoidea or to Octodontidae. A systematic and parsimony-based phylogenetic analysis of the species traditionally included in Acarechimys showed that this genus is part of stem Abrocomidae. These results are primarily supported by morphology of the mandible and lower molars. Acarechimys is here restricted to three species, A. minutus, A. pulchellus and Acarechimys pascuali sp. nov., while another species, A. constans, is here transferred to a new abrocomid genus. The remaining species were nested within Octodontidae. According to these results, Abrocomidae might have been as diverse as its sister clade Octodontidae-Echimyidae during the late Oligocene–early Miocene. Extinction of this diversity would have resulted in marked loss of evolutionary history, with extant abrocomids being currently restricted to late-diverged euhypsodont representatives.     

First fossil fruits of Elaeocarpus (Elaeocarpaceae) in East Asia: Implications for phytogeography and paleoecology

The genus Elaeocarpus contains approximately 360 species and occurs in mesic forest communities from India, through to China, Southeast Asia, New Guinea, Australia, and New Caledonia. Elaeocarpus fossils are best known from the Eocene to the Miocene of Australia and the late Pliocene–early Pleistocene of India, but have not been documented from East Asia before. Here we describe six new species of Elaeocarpus, E. nanningensis sp. nov. from the late Oligocene Yongning Formation of the Nanning Basin, E. presikkimensis sp. nov. from the Miocene Erzitang Formation of the Guiping Basin, E. prerugosus sp. nov., E. prelacunosus sp. nov., E. preserratus sp. nov., and E. preprunifolioides sp. nov. from the late Miocene Foluo Formation of the Nankang Basin in Guangxi, South China. This is the first reliable report for the genus occurring in East Asia, and the fossils indicate that Elaeocarpus had colonized this region by the late Oligocene and represented by a morphologically diverse group of species by the late Miocene. This sheds new insights into the timing and migration patterns of the genus in East Asia. Elaeocarpus is typically a rainforest genus occurring in mesic forests. Based on the habitat of their morphologically similar modern relatives we propose that these three sedimentary basins were warm and wet adjacent to mountainous regions with the evergreen or rain forests during the late Oligocene to Miocene.     

A Paleogeographic Overview of Tropical Fossil Sloths: Towards an Understanding of the Origin of Extant Suspensory Sloths?

Modern sloths are among the more characteristic mammals of South and Central American faunas. Recent discovery in four Paleogene, 22 Neogene, and dozens of Pleistocene fossiliferous localities in the tropics has revealed an unexpected paleobioversity constituted by some 81 fossil sloth species. Probably originating in southern South America near the Eocene/Oligocene transition, sloths were represented in the tropics during the late Oligocene by Pseudoglyptodon, Mylodontidae, and Megalonychidae. The latter occupied the West Indies between at least the late early Miocene and late Pleistocene, and two mylodontid clades, Octodontobradyinae and Urumacotheriinae, were characteristic of Amazonian localities from the Colhuehuapian and the Laventan periods, respectively, until the end of the Miocene. Megatheriinae and Nothrotheriidae appeared during the middle Miocene, colonizing the tropics and then North America, where Mylodontidae and Megalonychidae had already been present since the early late Miocene. Nothrotheriids are more abundant and diversified during the late Miocene in the tropics than in southern South America. Remains closely related to either of the modern sloths are absent from the fossil record, including those in the tropics. The characteristic suspensory posture of Bradypus and Choloepus appeared independently and likely after the Miocene epoch, and thus well after the hypothesized split suggested by molecular studies of the respective clades of these genera. Given their current widespread distribution in and reliance on the tropics, prospecting efforts for the direct fossil kin of suspensory sloths should concentrate on deposits in the Amazonian region, as this area has shown promise in producing fossil sloths.     

A new species of fossil Scutus Montfort, 1810 from New Zealand (Mollusca: Gastropoda: Fissurellidae)

ABSTRACT

A new fossil species of the genus Scutus (Scutus mirus n. sp.) is described from five Late Oligocene to Early Miocene (Waitakian to Altonian; 25.2–15.9?Ma) localities in the South Island, New Zealand. It is one of the oldest fossil species of Scutus known and probably inhabited very shallow, sub-tropical waters surrounding Zealandia during this time. The holotype of Scutus petrafixus Finlay, 1930 is re-examined; it is possibly from All Day Bay, Kakanui (Waitakian 25.2–21.7?Ma). The New Zealand species documented herein significantly expand our understanding of the fossil record of this shallow-marine molluscan lineage, and by proxy, also indicate the presence of very shallow coastal marine environments around the late Oligocene and early Miocene in southern Zealandia.     

The oldest lamprophiid (Serpentes,Caenophidia) fossil from the late Oligocene Rukwa Rift Basin,Tanzania and the origins of African snake diversity

Extant snake faunas have their origins in the mid-Cenozoic, when colubroids replaced booid-grade snakes as the dominant species. The timing of this faunal changeover in North America and Europe based on fossils is thought to have occurred in the early Neogene, after a period of global cooling opened environments and made them suitable for more active predators. However, new fossils from the late Oligocene of Tanzania have revealed an early colubroid-dominated fauna in Africa suggesting a different pattern of faunal turnover there. Additionally, molecular divergence times suggest colubroid diversification began sometime in the Paleogene, although the exact timing and driving forces behind the diversification are not clear. Here we present the first fossil snake referred to the African clade Lamprophiinae, and the oldest fossil known of Lamprophiidae. As such, this specimen provides the only potential fossil calibration point for the African snake radiation represented by Lamprophiidae, and is the oldest snake referred to Elapoidea. A molecular clock analysis using this and other previously reported fossils as calibration points reveals colubroid diversification minimally occurred in the earliest Paleogene, although a Cretaceous origin cannot be excluded. The elapoid and colubrid lineages diverged during the period of global warming near the Paleocene-Eocene boundary, with both clades diversifying beginning in the early Eocene (proximate to the Early Eocene Climate Optimum) and continuing into the cooler Miocene. The majority of subclades diverge well before the appearance of colubroid dominance in the fossil record. These results suggest an earlier diversification of colubroids than generally previously thought, with hypothesized origins of these clades in Asia and Africa where the fossil record is relatively poorly known. Further work in these regions may provide new insights into the timing of, and environmental influences contributing to, the rise of colubroid snakes.