寒武纪大爆发与动物树的成型

由“真动物”构成的动物界（不包括海绵）通常被划分为双胚层动物、原口动物和后口动物3个亚界。寒武纪大爆发经历了爆发的前奏—序幕—主幕3个阶段,其中后两个阶段处于早寒武世初期（依次以小壳化石的首次辐射和澄江动物群爆发为代表）,另一个则发生在“寒武前夜”（即前寒武纪末期）,以伊迪卡拉生物群为代表。这次独特的三幕式大爆发分步完成了动物形态演化谱系树（简写为TOA）的成型。已有化石证据显示,动物树的3大主体或3个亚界的起源及其早期辐射分别发生于寒武纪大爆发这3个主要阶段。澄江化石库中发现的早期后口动物亚界涵盖了该亚界中所有6大分支（棘皮类、半索类、头索类、尾索类、脊椎类和绝灭了的古虫类）的原始类群,澄江动物群时代标志着寒武纪大爆发的顶峰,完成了动物树框架的成型,从而宣告了寒武纪大爆发的基本终结。其学术重要性超越了中寒武世的布尔吉斯页岩,因为前者位于动物树形成的“源头”,而后者仅代表此后的“一段流程”。在动物演化历史上4个最具转折意义的重大创新事件（即多细胞动物起源、两侧对称动物起源、后口动物起源和脊椎动物起源）中,第一个发生在伊迪卡拉纪甚至更早的时期,其余3个则可通过梅树村化石群和澄江化石群观察到其完成的主体过程。已灭绝的古虫动物门,是初具鳃裂构造的原始分节动物,作为由原口动物向后口动物过渡的一个珍稀“缺环”,很可能代表着后口动物亚界的一个根。古囊类兼有两侧对称的古虫类和一些原始棘皮动物的镶嵌体征,最可能代表棘皮动物的一个根。在早寒武世动物演化谱系中,后口动物的＂顶端类群＂的代表是被誉为“第一鱼”的昆明鱼目,目前已知包括昆明鱼、海口鱼和钟健鱼,它们具对眼、脊索与串珠状脊椎软骨共存的原始脊椎,?     

澄江化石库中的双胚层动物新知

作为观察寒武纪大爆发主幕奥秘的最佳科学窗口,澄江化石库展示了三胚层动物(即两侧对称动物)首次全面大辐射的历史产物,让学术界终于看到了已知最古老的同时包括原口类和后口类两大支系的动物谱系庞大“树冠”的主体轮廓,从而给许多较高级动物类群的起源和早期演化研究提供了十分重要的信息。然而,该化石宝库中却很少见到作为“树干”的双胚层动物(即辐射动物)的化石记录,这给“真动物”的源头探秘蒙上了阴影。尽管广布于前寒武纪末期全球浅海域的“文德生物群”一直被认为是动物界早期一次“失败的演化试验”的牺牲品,但其中一支成功延续到早寒武世的春光虫(Stromatoveris)具有叠覆状梳齿构造,很可能代表着栉水母类的一类原始祖先。文中描述新的羽毛状动物王氏澄江海笔(Chengjiangopennawangii)不同于春光虫,前者具有彼此分离的羽枝,且羽枝上保存了排列规整的珊瑚个体虫室,因而与现代八射珊瑚中海笔类的形态学特征十分一致;新的谱系分析显示,它应该代表刺胞动物门中的一个原始类群。     

Is Darwin's ‘Abominable Mystery’ still a mystery today?

Darwin described the Cretaceous diversification and subsequent rapid rise of flowering plants (angiosperms) as an “abominable mystery”: how could they have achieved worldwide ecological dominance by early Paleogene times when the oldest angiosperm fossils are only Cretaceous in age? However, recent phylogenetic and palaeobiogeographical analyses have suggested a much earlier, perhaps Triassic origin for the stem angiosperms. We suggest that the fossil record is accurately reflecting the rapid diversification of the crown angiosperms that molecular data suggests occurred in Cretaceous–early Palaeogene times, and which coincides with similar explosive diversifications that occurred in other parts of the terrestrial biota including insects, birds and mammals. Early, stem angiosperms are poorly represented in the fossil record for a combination of taphonomic and ecological reasons. We conclude that the reported evidence shows that the Darwin's “mystery” is in fact no mystery and confirms that it merely requires an appropriate reading of the fossil record.     

二纺蛛总科（蜘蛛目：新蛛下目）蜘蛛化石研究进展

二纺蛛总科包括5个现生科、5个化石科以及一些科级位置未定的化石物种,是蜘蛛目中较为古老的一个类群。目前已知最早的二纺蛛总科化石记录可以追溯到早侏罗世,但分子系统发育研究推测其起源时间为二叠纪。二纺蛛总科蜘蛛在中生代最为繁盛,化石物种相对较多,是蜘蛛化石类群的重要组成部分。而现生二纺蛛总科蜘蛛十分稀少,且主要分布在热带和亚热带地区。近年来的系统发育研究认为二纺蛛总科可能是复杂生殖器类的姐妹群,是蜘蛛演化研究的关键类群。因此,本文对二纺蛛总科分类系统的研究历史进行了回顾,总结了二纺蛛总科化石类群在不同地质历史时期的组成和分布,统计了二纺蛛总科蜘蛛化石的名录。目前,二纺蛛总科蜘蛛化石分类研究基础薄弱,部分化石类群的系统位置存疑,二纺蛛总科内部系统发育关系尚不清晰。在将来的研究中,需要利用多种技术手段获取化石材料的形态学信息,打好分类学基础;结合化石类群和现生类群进行系统发育分析;进而联系古地理、古生态、生物学行为等研究,全方位分析二纺蛛总科的自然演化历史。     

二纺蛛总科（蜘蛛目：新蛛下目）蜘蛛化石研究进展

二纺蛛总科包括5个现生科、5个化石科以及一些科级位置未定的化石物种,是蜘蛛目中较为古老的一个类群。目前已知最早的二纺蛛总科化石记录可以追溯到早侏罗世,但分子系统发育研究推测其起源时间为二叠纪。二纺蛛总科蜘蛛在中生代最为繁盛,化石物种相对较多,是蜘蛛化石类群的重要组成部分。而现生二纺蛛总科蜘蛛十分稀少,且主要分布在热带和亚热带地区。近年来的系统发育研究认为二纺蛛总科可能是复杂生殖器类的姐妹群,是蜘蛛演化研究的关键类群。因此,本文对二纺蛛总科分类系统的研究历史进行了回顾,总结了二纺蛛总科化石类群在不同地质历史时期的组成和分布,统计了二纺蛛总科蜘蛛化石的名录。目前,二纺蛛总科蜘蛛化石分类研究基础薄弱,部分化石类群的系统位置存疑,二纺蛛总科内部系统发育关系尚不清晰。在将来的研究中,需要利用多种技术手段获取化石材料的形态学信息,打好分类学基础;结合化石类群和现生类群进行系统发育分析;进而联系古地理、古生态、生物学行为等研究,全方位分析二纺蛛总科的自然演化历史。     

Antiquity and long-term morphological stasis in a group of rove beetles (Coleoptera: Staphylinidae): Description of the oldest Octavius species from Cretaceous Burmese amber and a review of the “Euaesthetine subgroup” fossil record

The Staphylinine group of rove beetle subfamilies is a significant animal radiation, and one subordinate monophyletic clade – the ‘Euaesthetine subgroup’ – includes around 3000 species in subfamilies Euaesthetinae and Steninae and has a fossil record dating to the Early Cretaceous. Detailed morphological study of a new well-preserved Cretaceous Burmese amber fossil revealed strong evidence consistent with its taxonomic placement in the euaesthetine genus Octavius. We thus describe Octavius electrospinosus sp. nov., the first Cretaceous record of the genus and of the tribe Euaesthetini. Previously, the oldest records of Octavius and Euaesthetini were from the Eocene (Baltic amber) and discovery of O. electrospinosus sp. nov. therefore nearly doubles the minimum lineage age of Octavius, increasing it by 50 million years. We also briefly review the known Euaesthetine subgroup fossil record and tabulate summary data for all previously described fossils. All are placed in extant genera, and have visible diagnostic generic-level characters including some putative synapomorphies as judged by recent phylogenetic work. Including O. electrospinosus sp. nov., there are now four known Cretaceous species, all of which belong to either Octavius, Nordenskioldia, or Stenus. To explain the long-term morphological stasis in this group of rove beetles, we suggest that the continuous presence of mesic habitats may have buffered these lineages from strong selection for morphological change. Considering the fossils along with phylogenetic hypotheses we suggest the Euaesthetine subgroup originated in the Late Jurassic– Early Cretaceous and the Staphylinine group in the Early Jurassic. We emphasize the derived status of Cretaceous fossils in assessing possible divergence times and the significance of the pre-Cretaceous taphonomic bias for restricting more robust estimates. Further detailed morphological study of available fossils in a phylogenetic framework is badly needed to clarify the phylogenetic positions of these taxa.     

八臂仙母虫(Eoandromeda octobrachiata)新材料及其意义

在中国贵州江口地区陡山沱组新发现的化石材料表明,伊迪卡拉/震旦纪(Ediacaran/Sinian)的宏体化石——八臂仙母虫Eoandromeda octobrachiata的原始生活状态可能为一个较高的圆锥形生命体,具有沿着侧面锥状螺旋排列的八条旋臂.旋臂表面保存有与栉水母动物的栉板可能同源的横向有机质条带,规则排列...     

Phosphatized Punctatus with mouthparts and its embryo fossils from the Lower Cambrian of Ningqiang, south Shaanxi, China

The appearance of coelenterates marks the real beginning of metazoan evolution. It therefore has a prominent position in the origin and evolutionary history of organisms, and is also a pivotal question of evolutionary biology. Punctatus is an extinct, ancient marine animal from the early stage of the Cambrian explosion, occurring at the lowermost Cambrian of both Kuanchuanpu (Ningqiang, Shaanxi) and Maidiping (Emei, Sichuan) areas. Punctatus has been studied for many years since the discovery of its fragments. Systematic and phylogenetic analysis has long been limited because of the rarity of complete specimens. In order to improve research into Punctatus, more than ten thousand globular fossils were recovered by means of “Chemistry Retting”. On the basis of the study of these globular fossils, a series of Punctatus fossils with cone parts and finely preserved soft-tissue mouthparts and fossilized metazoan embryo were recovered. Through research on characteristics such as shape, modality and structure of these fossils symbiotic with Punctatus emeiensis, the author found many possible embryo fossils including the evidence of gastrula-stage animal fossils. The sequence of fetation might have appeared on the corolliform oral region of Punctatus emeiensis. A study of the soft tissues, functional morphology and the sequence of embryo fossils shows evidence that Punctatus resembles coelenterate polyps in systematic classification. Perhaps it also represents an ancestor of the coelenterate with an original tentacle. A comparison with the real “tentacle animal” found in the Chengjiang Fauna, shows that the original tentacle is very tiny and its function range is limited. This reveals the primitive nature of the animal. Although the original tentacle is so small, it does exist, representing the first big step towards the real flexible tentacle with a strong function from the early evolving tentacle. Translated from Journal of Acta Palaeontologica Sinica, 2006, 45(2): 182–194 [译自: 古生物学报]     

Early Cambrian radiation of brachiopods: A perspective from South China

Brachiopods, a group of benthic suspension-feeding marine invertebrates, made their first appearance in the Lower Cambrian. In the Yangtze Platform (South China), well-exposed Lower Cambrian stratigraphic succession represents shallow to deeper water environments. Strata from eastern Yunnan, southern Shaanxi and the Yangtze gorges areas of western Hubei Province, deposited in muddy-siltstone and carbonate lithofacies, contained an abundant variety of brachiopods, including all the representatives of the subphylum Linguliformea and the calcareous-shelled genera of Kutorgina and Nisusia from the subphylum Rhynchonelliformea. Thus the fossil assemblage bears witness to the first major phase of evolutionary radiation of brachiopods during the ‘Cambrian explosion’ interval of metazoans. Brachiopods from the celebrated Chengjiang fauna have exquisitely preserved soft-tissues, which reveal the body plans and evolutionary acquisition of morphological novelties of the early stocks, and also provide a good opportunity for testing the analogies with the stem groups from the extant representatives. These fossils have corroborated the view that brachiopods developed complex organization of tissues, and achieved considerable evolutionary success already by the onset of ‘Cambrian Explosion’. Thus it is not improbable that a large part of this radiation occurred within, or only just before early Cambrian time. Studies of Chengjiang brachiopods suggest that attachment by a pedicle to the substrate was probably the most common relationship of Cambrian brachiopods with the substrate where they inhabited.     

Frullania pinnata spec. nov. (Frullaniaceae,Porellales), a new leafy liverwort in mid-Cretaceous Burmese amber from Myanmar

An inclusion in Cretaceous amber from Myanmar is described as a new fossil species in the extant liverwort genus Frullania. The name Frullania pinnata is proposed for the taxon that is characterized by entire underleaves paired with a conspicuous pinnate branching pattern, two distinctive and stable morphological features. Entire underleaves are known in several extant Frullania species but had not previously been documented in Frullania in amber. The combination of morphological characters in this new fossil species is unknown in any crown group lineage of Frullania, and may very well represent a stem lineage element of the genus. This discovery is important because it expands our understanding of the diversity of Frullaniaceae in the Burmese amber forest as well as important ramifications for the phylogenetic reconstruction of extant Frullania lineages.     

澄江动物群的核心价值:动物界成型和人类基础器官诞生

澄江动物群以富含多门类精美软躯体构造化石享誉学界,是寒武纪大爆发主幕的见证者;它亲眼目睹了地球动物树上几乎所有主要门类祖先的首次大聚会。对该动物群30多年的大规模调查研究大体上可以划分为3个阶段。始自1984年的第一个10年(1984—1994年),早期动物树上的基础动物亚界和原口动物亚界的大量门类被发现,然而对后口动物亚界全然无知。第二个10年研究(1995—2005年),基础动物亚界和原口动物亚界中的更多门类被发现;而最重要的进展,是西北大学团队主导的早期后口动物大发现使后口动物亚界的基本框架被构建,由此地球三分动物树首次成型。第三个10年及其以后时期(2005年之后),3个动物亚界中进一步添加了一些新的门类认知;综合20世纪40年代以来对埃迪卡拉生物群、寒武纪第一世和第二世的珍稀动物群研究成果,学界开始思考三分动物树分步成型与多幕式寒武纪大爆发事件的内在关联,提出了三幕式寒武纪大爆发新假说。澄江动物群研究具有多方面的科学价值,包括动物群结构研究,生态环境研究等,然其核心学术价值主要集中在如下两个方面:(1)作为寒武纪大爆发的主幕见证者,澄江动物群首次创建了地球上三分动物树的完整谱系框架。三幕式寒武纪大爆发假说揭示了地球上动物门类由低等到高等、分阶段爆发创造的本质内涵,即历时约0.4亿年的三幕式寒武纪大爆发与动物树阶段性快速成型的内在耦合性。第一幕发生于埃迪卡拉纪晚期,它诞生了众多基础动物亚界门类(包括大量绝灭门类),也可能出现了少量原口动物的先驱分子;第二幕发生于寒武纪的第一世,除了延续基础动物门类的繁盛外,更创生了原口动物亚界里的主要门类(包括蜕皮类和触手担轮类);第三幕发生于寒武纪第二世(以澄江动物群为代表),不仅延续了基础动物和原口动物门类的兴盛,更重要的是诞生了后口动物亚界里的所有门类。至此,整个三分动物树的框架成型,寒武纪大爆发创新事件宣告基本结束。文章还对几个重要动物类群的生物学属性进行了评述。第一鱼昆明鱼目是已知唯一的最古老脊椎动物,而云南虫、海口虫既不是脊椎动物(即高等脊索动物),也不属于低等脊索动物,而是一类特殊的低等后口动物;长江海鞘是可信的始祖尾索动物,它支持了经典的尾索动物起源假说;古虫动物门首创鳃裂构造,对后口动物亚界的起源探索提供了关键信息。(2)澄江动物群具有深刻的人文哲学意义,所发现的古虫动物门创造的“第一鳃裂”,昆明鱼目创造的“第一头脑”“第一脊椎”“第一心脏”,为达尔文在《人类的由来》中关于人类的主要基础器官起源悬案的破解提供了关键证据。澄江动物群承上启下,其各种形态解剖学信息对解读埃迪卡拉生物群以及寒武纪相关化石库中绝大多数生物类型都具有重要的参考价值。     

石珊瑚目的早期系统发育和系统分类研究

石珊瑚目是中新生代的重要造礁生物。直到中三叠世才大量出现多种石珊瑚化石，距二叠纪末生物大灭绝、古生代皱壁珊瑚和床板珊瑚灭亡14 Ma之久。石珊瑚的起源和早期系统发育，成为生物大灭绝及复苏的一个难题。文章目的是综述多学科研究石珊瑚早期系统发育的进展，以及多家系统分类，展望前景。研究方法有骨骼构造与微细构造及软体的对比，时空分布分析，寻找早期珊瑚化石及分子生物学等。研究结果主要有石珊瑚可能起源于只有软体的海葵类，发现早期石珊瑚化石，以及石珊瑚在300 Ma之前已演化为两支，但还有不少空白和争议。结论是为了查明石珊瑚的早期系统演化和分类，需要加强古生物与今生物学者多学科交叉的综合研究，努力寻找中间环节，填补研究空白。     

Notes on the divergence time of the fern genus Asplenium from fossil and molecular evidence

The extant fern genus Asplenium Linnaeus, 1753 is widely distributed in tropical-temperate regions and shows a high diversity. The oldest fossils assigned to it comprise fronds and spores preserved in situ from the Lower Cretaceous of northeast Asia. However, molecular dating suggests that Asplenium diverged during the Paleocene (∼57.7 Ma). Here we present some explanations for the disparity in fossil and molecular age estimates, and new insights into the origin and evolution of the genus. We suggest that Cretaceous Asplenium most likely represents a stem-group member, and that lineage extinction during the early evolutionary stages of the genus resulted in missing nodes.     

鄂尔多斯盆地北岔沟砂岩沉积特征及其对天然气成藏的控制作用*

北岔沟砂岩为华北地台二叠系山西组与太原组划分对比的主要标志层,在鄂尔多斯盆地呈大面积连片分布,是上古生界重要的天然气储集层。依据地面露头调查结果与勘探资料,结合前人有关古构造和岩相古地理研究成果,笔者对鄂尔多斯盆地北岔沟砂岩的沉积特征进行了详细分析。结果表明,受益于物源区大量高石英含量的粗碎屑物质在短时间内的集中供应,北岔沟砂岩在地层中位置稳定,以含砾粗粒石英砂岩为主,石英含量高达90%左右,砂体叠置紧密,单砂体之间很少见泥岩隔夹层。这些砂体分别隶属于冲积扇、砾质辫状河和辫状河三角洲沉积体系,在鄂尔多斯盆地北部高、南部次高、中南部低的古地理背景控制下,由盆地南、北边缘向中部依次推进,沉积中心区以北发育4个规模宏大的三角洲沉积主砂带,以南发育6个规模较小的主砂带。受各主砂带砂体规模、矿物组分含量及其成岩作用差异影响,各砂带的储集物性具有较大变化,其中砂体规模最大、石英含量最高、长石等易溶矿物在酸性地层水作用下溶蚀彻底的杭锦旗—榆林—子洲—清涧—永和—吉县主砂带,储集能力最好,最有利于天然气的大规模聚集成藏,是鄂尔多斯盆地北岔沟砂岩天然气进一步勘探开发的有利场所。     

化石及现生真蕨类植物的根系:形态演化及古环境意义*

根是植物的重要营养器官,对维持植物生理活动及生态系统物质循环和能量流动极为重要。真蕨类植物的根系以可预测的方式起源于枝,被称为枝生根,通常也被称为不定根。相比于对种子植物根系的广泛研究,对真蕨类植物根系的认识较为薄弱。通过广泛收集各类文献中有关真蕨类根系的信息,以及观察大量化石、现生植物标本,对若干化石及现生真蕨类植物根系的形态学特征进行了综述。在繁盛于中、晚泥盆世的早期真蕨类植物中,密集的枝生根着生于茎轴、或推测的根状茎、或主干膨大的基部上;根系的分枝罕见,若存在,则仅表现为等二歧式分枝;而在真蕨纲冠群植物中,枝生根密集或稀疏,根系呈单轴式分枝,侧根发育,可达2级甚至更多。真蕨类根系统的形态演化可采用“顶枝学说”进行解释: 真蕨类植物的根系具有一定的演化保守性,均为枝生根;早期真蕨类的原始根系可视为不分枝的或偶见二歧式分枝的顶枝束;经历顶枝束的越顶生长,侧根系统在真蕨纲冠群中演化出现。真蕨类根系的构型、形态、解剖结构、生物量分配等信息可在化石记录中得以揭示,这些信息可为探讨植物根系的演化及研究地史时期的植物—土壤系统提供重要依据。最后,简述了这方面研究的1个案例,即北京西山侏罗纪地层中的真蕨类根系化石及根系—土壤系统。     

近代分支系统学的回顾与展望

本文简要介绍分支学近２０年的存在和发展。生物系统学界对求得树系所用方法的讨论由于分支学的出现而变得活跃。广泛的辩论促进了系统学方法的严谨化。分支学已被越来越多的生物系统学家所接受。近年分支学界讨论的内容包括化石在系统学中的作用，简约法则的应用，衡量树系的统计方法，以及如何将分子与形态数据结合等问题。     

寒武纪叶足动物:困惑与思考

叶足动物在显生宙最早期的海洋中曾一度繁盛,但如今茫茫大海中它们已杳无踪影。不过,学术界主流认为它们并没有绝灭——早期叶足动物的一个分支成功演化出原始的节肢动物,成就了当今地球上最庞大的动物家族(六足类、甲壳类、多足类、螯肢类等);叶足动物另一个分支则离开海洋,保持基本躯体构型,变身为陆生叶足动物,然而历经沧桑后,仅余一两脉(有爪动物及缓步动物)残喘幸存至今。这个史诗般的精彩演化故事,深深吸引了众多学者竞先求索。近30年来,以我国云南澄江生物群为代表的一系列寒武纪特异埋藏化石库中发现了大量叶足动物化石。经研究,学术界迄今似已破解了早期叶足动物重重疑团,足以额手称庆。然而治学须严谨,考证不偏狭。作者认为,围绕早期叶足动物的迷雾尚未消散,许多问题还有待解答,部分观点仍需确证。本文对早期叶足类研究的部分主流观点提出一些不同看法,并对尚未破解的疑难问题提出尝试性的见解和猜想。     

Early Cretaceous Aleyrodidae (Hemiptera: Sternorrhyncha) from the Lebanese amber

Four new, monotypic genera of Aleyrodidae from the Lower Cretaceous Lebanese amber are described: Shapashe aithiopa gen. sp. nov.; Milqartis azari gen. sp. nov.; Aretsaya therina gen. sp. nov.; and Yamis libanotos gen. sp. nov. All new taxa represent subfamily Aleurodicinae. The key for identification of genera of Aleurodicinae from the Lebanese amber is provided. The morphological features of newly established taxa are discussed. The importance of the fossils for phylogenetic, evolutionary and ecological studies is presented.     

The emergence of Metazoa in the early history of life

After consideration of currently used time scales and their geochronometric calibration, the views of biologists on evolutionary events in the early history of the Metazoa are discussed, mainly on the basis of two examples from the abundant literature. The phylogenetic views of A.V. Ivanov on the origin and early evolution of Metazoa to the level of sponges, Cnidaria and Platyhelminthes, are outlined. A brief discussion of the subsequent coelomate radiation, based on the work of R.B. Clark follows. It is proposed that these evolutionary events, postulated by biologists, occurred in the time span of about 300 Ma preceding the appearance of large Metazoan body fossils in Ediacarian time, about 600-560 Ma ago. Palaeontological data from this pre-Ediacarian, Late Proterozoic time span are limited to few, mostly small trace fossils. The reasons for this paucity and the consequent need to give new dimensions to trace fossil studies are discussed. It is concluded that there is evidence for a pre-Phanerozoic ‘prehistory’ of the Metazoa. A full explanation of the events and processes leading to their diversification and of its chronologic sequence will only be possible when all interacting systems, including the lithosphere, hydrosphere and atmosphere, relevant to their evolution during the appropriate time interval have been more completely elucidated.     

A new pipid frog from the Upper Cretaceous of Patagonia and early evolution of crown-group Pipidae

Pipid frogs are fully aquatic frogs that today inhabit freshwater environments of southern continents on both sides of the Atlantic Ocean, with a fairly good fossil record dating back to the Cretaceous. Here I report on fossils from the Allen Formation (upper Campanian–lower Maastrichtian), Río Negro Province, Argentina, that are ascribed to a new genus and species of pipid. In order to assess the evolutionary relationships of the new taxon, which is represented by sphenethmoids, otic capsules, ilia, humeri, and vertebrae, cladistic analyses of a data matrix of 165 osteological characters scored for 36 taxa were performed. The results are congruent with previous hypotheses of pipoid interrelationships and consistently place the new taxon as part of the lineage today represented by the African xenopodines. Temporal calibration of the phylogenetic tree based on the fossil record imply that the origin and early diversification of crown-group Pipidae might have occurred during the Early Cretaceous, prior to the final breakup of western Gondwana. This study highlights the importance of including fossils, even fragmentary ones, directly in phylogenetic analyses in order to disentangling how, when, and where pipid frogs diversified.