Taphonomy and abundance of birds from the Lower Eocene Fur Formation of Denmark

The pattern, pace and extent of the evolutionary radiation of modern birds (Neornithes) by the end‐Cretaceous (65 Ma) has long been debated. Well‐dated, taphonomically understood and phylogenetically constrained fossil birds from both sides of the Cretaceous–Paleogene (K–Pg) boundary are required to quantify the shape of this radiation, but have largely been lacking. Here we report on a large collection of fossil birds from the Lower Eocene of Denmark (ca. 54 Ma) that includes three‐dimensionally preserved, articulated specimens from carbonate concretions as well as skeletal imprints and feathers. These birds are from a marine diatomite sequence (the Fur Formation), a low‐energy deep‐water preservational environment unique to the Cretaceous and Paleogene avian fossil record. We present taphonomic and palaeoecological information gleaned from these birds that in combination with phylogenetic data have implications for unravelling avian survivorship across the K–Pg boundary as well as for the pattern of the neornithine evolutionary radiation. Copyright © 2009 John Wiley & Sons, Ltd.     

Paleoenvironmental conditions across the Cretaceous–Paleogene transition at the Apennines sections (Italy): An integrated geochemical and ichnological approach

Two distal Cretaceous–Paleogene (K/Pg) boundary sections in the Central Apennine region (Italy) have been studied: Bottaccione Gorge and Contessa Highway. Geochemical and carbon isotope analyses on the infilling of trace fossils and on the host sedimentary rocks were performed to determine paleoenvironmental conditions during the Cretaceous–Paleogene transition. Major and trace element contents were measured in a 63 cm-thick interval at Bottaccione Gorge (from 22 cm below to 41 cm above the K/Pg boundary) and in a 72 cm-thick interval at Contessa Highway (from 43 cm below to 29 cm above the K/Pg boundary). Even though the K/Pg ejecta layer is now depleted at these sections due to continuous oversampling, the uppermost Maastrichtian and lowermost Danian deposits record the paleoenvironmental conditions prior to and after the K/Pg event. We used redox-sensitive element ratios (V/Al, Cr/Al, Co/Al, Ni/Al Cu/Al, Zn/Al, Mo/Al Pb/Al and U/Mo) and detrital element ratios (K/Al, Rb/Al, Zr/Al and ƩREE/Al) as proxies of certain environmental parameters, used for paleoenvironmental reconstruction. In general, similar values for elemental ratios are registered within Maastrichtian and Danian deposits, which supports similar paleoenvironmental conditions prior to and after the K/Pg event as well as the rapid reestablishment of the pre-impact conditions (i.e., oxygenation, nutrient availability, and/or sedimentary input). An enrichment in certain redox-sensitive elements above the K/Pg at the Bottaccione Gorge section suggests lower oxygenation, as also evidenced by the tracemaker community. Carbon isotope composition data from the infilling material of trace fossils furthermore reveals values similar to those of the host rocks at the corresponding depth, which supports an active infilling by nearly contemporaneous bioturbation during sediment deposition.     

How bioturbation obscured the Cretaceous–Palaeogene boundary record

The Cretaceous–Palaeogene (K/Pg) boundary interval is often penetrated by burrows, which may obscure stratigraphic and micropalaeontological records, leading to misinterpretations of the sequence of events spanning the K/Pg boundary. Here, we assess the role of burrowing organisms in the redistribution of benthic foraminifera across the boundary at Bidart (France), and report a strong relationship between the behaviour represented by pre‐ and post‐K/Pg trace fossils and their benthic foraminiferal content. We further infer a brief interval of eutrophic conditions at the seafloor, as reported from other locations, which disappeared from the lowermost Danian stratigraphic record and is represented only inside post‐K/Pg trace fossils hosted in Cretaceous strata. The combined study of trace fossils and microfossils is a powerful tool in eco‐stratigraphy and event‐stratigraphy, and can yield important insights into the completeness of the K/Pg record, especially at locations such as Bidart where this interval has traditionally been assumed to be complete.     

A review of Late Cretaceous fossil birds from Hungary

We review the previously described Late Cretaceous (Santonian) bird remains from the Csehbánya Formation in the Bakony Mountains of Hungary, augmenting initial work by Ősi ( 2008 ), and add a number of newly collected fossils. All together, the eight fossil specimens so far collected from this site are important to our understanding of avian evolution because they document a large range of taxon body sizes from at least one major lineage (Enantiornithes) and come from a critically undersampled time period in the Cretaceous. Globally, very little fossil bird material has been collected from the middle stages of the Late Cretaceous, the Coniacian and Santonian; most known taxa are either Early Cretaceous (ca. 120 Ma) in age or are from the terminal Campanian and Maastrichtian (ca. 70–65 Ma). Indeed, one of the Csehbánya Formation fossil birds is recognized as a new taxon of large enantiornithine, an avisaurid apparently similar in its largely unfused foot morphology to the Argentine Soroavisaurus and to the North American Avisaurus. The Central European records reviewed in this paper highlight the wide distribution of some Late Cretaceous fossil birds, particularly avisaurid enantiornithines, and lead us to a brief discussion of avian biogeography at the end of the Mesozoic. Copyright © 2010 John Wiley & Sons, Ltd.     

Mammalian femora across the Cretaceous–Paleogene boundary in eastern Montana

Our understanding of latest Cretaceous and earliest Paleogene mammalian evolution is based almost entirely on the dental fossil record. Mammalian postcranial fossils are rare and mostly found as isolated elements in latest Cretaceous and earliest Paleogene vertebrate microfossil assemblages of North America. Although placing these fossils in a tooth-based taxonomic framework is difficult, they can provide insight into locomotor diversity and habitat preference to complement diet reconstructions and diversity estimates from dental fossils. Here, we describe 64 femora of mammals recovered from latest Cretaceous (Lancian) and earliest Paleogene (Puercan) localities in eastern Montana. We sorted these based on morphology and size (morphotypes). In some cases, morphotypes were tentatively assigned to dentally based taxa that are known from these strata.Although our resulting femoral dataset is small relative to the study area's dental dataset, we show several key findings. First, there is a greater morphological diversity of multituberculate femora than previously recognized, especially in the latest Cretaceous sample. In contrast, metatherians, which have a high relative abundance in Lancian Hell Creek Formation dental assemblages, are absent from our postcranial samples; eutherian femora are only present in the Puercan assemblages. Second, we record a minor decrease in morphotype richness across the K–Pg boundary that is associated with an increase in mean specimen size, due to the appearance of a few significantly larger-bodied, immigrant taxa. Among the eutherians, there are two specimens of larger-bodied early Puercan archaic ungulates, a very large specimen of a middle/late Puercan taeniodont, pantodont, or triisodontid, as well as a specimen possibly attributed to a “plesiadapiform” archaic primate. Third, preliminary functional morphologic analyses of the more complete specimens suggest that locomotor diversity increased from mainly arboreal or terrestrial/saltatorial multituberculates in the latest Cretaceous to include a fossorial multituberculate and potentially an arboreal eutherian in the early Paleocene. These patterns parallel those previously reported from a dental dataset and indicate that postcranial data are valuable as an independent means to test hypotheses of taxonomic and ecomorphological diversity across the K–Pg boundary.     

An Ichthyornis-like bird from the earliest Late Cretaceous (Cenomanian) of European Russia

Ornithuromorph birds (the clade which includes modern avian radiation) first appeared in the Early Cretaceous in Asia and achieved a great diversity during the latest ages of the Late Cretaceous (Campanian and Maastrichtian). The evolutionary history of orithuromorphs during the first 17 MYAs of the Late Cretaceous (Cenomanian to Santonian ages) remains very poorly known, as the fossil record for this time interval is largely restricted to several isolated finds of the classic avian genus Ichthyornis in North America. Here we describe an isolated distal tibiotarsus of an evolutionary advanced bird, morphologically similar to Ichthyornis, from the middle Cenomanian of Saratov Province, European Russia. This is the first documentation of an Ichthyornis-like bird in the Old World. The find further constitutes only the second pre-Campanian record of the Late Cretaceous Ornithuromorpha in Eurasia, the second record of Cenomanian birds in Russia. This discovery shows that Ichthyornis-like birds enjoyed a wide geographical distribution as early as the beginning of the Late Cretaceous. Given that the earliest and the most primitive ornithuromorph birds are known from Asia, the new find supports a Eurasian origin for Ichthyornithidae.     

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.     

中国中生代鸟类研究概述

对最近轰动世界的我国中生代鸟类化石的系统结果表明,中国是目前中生代早期鸟类化石产地和化石发现最多的地区,从晚侏罗世到早白垩世存在两个亚纲的鸟类分子,晰鸟亚纳（Sauriurae)和今鸟亚纲（Ornithurae),鸟类从爬行动物分化出来不久即分为两大支系向前演化,孔子鸟（Confuciusornis)和辽宁鸟（Liaoningornis)的发现证明,始祖鸟（Archaeopteryx)只是鸟类进化的一个旁支,不是最早的鸟类,孔子鸟代表鸟类向飞行进行的主流,辽宁鸟是目前今鸟亚纲最早的代表,鸟类在进化过程中渐变和突变同时并存,鸟类的历史应大大向前推移。     

A large collection of Presbyornis (Aves,Anseriformes, Presbyornithidae) from the late Paleocene and early Eocene of Mongolia

We describe a large collection of fossil ‘waterfowl’ bones that are referable to the extinct clade Presbyornithidae (Anseriformes). All of these fossils were collected between 1971 and 1994 from Upper Paleocene and Lower Eocene sediments at the Tsagaan Khushuu site in the Gobi Desert of southern Mongolia. The collection includes specimens referred to a new small species within the genus Presbyornis Wetmore, 1926 as well as large numbers of bones that we place in the genus Presbyornis. On this basis of the Tsagaan Khushuu collection we suggest that several species of Presbyornis likely coexisted in this region; indeed, the presence of large numbers of middle‐sized, morphologically consistent but probably ecologically disparate species at the Tsagaan Khushuu site is consistent with the range of variation seen, for example, in taxa of extant dabbling ducks (Anatini). Although the anatomy and phylogenetic position of Presbyornithidae (in particular Presbyornis) are well known, this material from Mongolia further demonstrates the prevalence of these birds in aquatic and semi‐aquatic habitats by the earliest Paleogene. Because presbyornithids are also well documented from the late Cretaceous, their palaeoecology and morphological diversity provides a clue to selective avian survivorship across the Cretaceous‐Paleogene (K‐Pg) boundary. Copyright © 2009 John Wiley & Sons, Ltd.     

A new bird track,Koreanaornis lii ichnosp. nov., from the Lower Cretaceous Hekou Group in the Lanzhou-Minhe Basin,Gansu, Northwest China,and implications for Early Cretaceous avian diversity

There are a growing number of Early Cretaceous avian tracks and trackways from around the world, with Asia (China and Korea) having the largest reported number and diversity of Mesozoic avian traces to date, and these new discoveries are increasing the Early Cretaceous avian ichnodivesrity of Laurasia. Here we report on a new Lower Cretaceous avian track locality in the Guanshan area, Yongjing County, Gansu Province, northwest China, and on a novel ichnospecies of Koreanaornis, Koreanaornis lii ichnosp. nov. Koreananornis lii is distinct from other Koreanaornipodidae in that it possesses a consistently wider digit divarication than previously described tridactyl tracks, and possess a short, small, posteromedially oriented hallux that displays a different orientation than that seen in Koreanaornis hamanensis. The lack of linear and angular data reported for digit I traces of many avian ichnotaxa has the potential to give misleading results in multivariate statistical analyses. Also, the wide divarication of Koreanaornis lii causes the ichnotaxon to not group with other Koreanornipodidae in multivariate analyses, but with Ignotornidae. Despite the results of the analyses, K. lii is morphologically distinct from these ichnotaxa. The results demonstrate that relying solely on multivariate statistical analyses without careful examination of footprint morphology will result in erroneous ichnospecies groupings. While new vertebrate ichnotaxa discoveries from Asia may support the hypotheses of the presence of a unique and endemic Asian vertebrate ichnofauna during the Cretaceous, the recent discovery of skeletal remains interpreted to be of a volant wading bird from the Early Cretaceous, and recent reports of tracks from volant avians, could suggest that flighted avians of the shore- and wading bird ecotypes could have had a Laurasian-wide distribution during the Early Cretaceous. However, strong convergence in foot morphology of shore- and wading birds suggests that avian ichnotaxa found in both present-day Asia and North America may have been made by birds endemic to eastern and western Laurasia during the Early Cretaceous.     

Geochemical and isotopic characterization of trace fossil infillings: New insights on tracemaker activity after the K/Pg impact event

Geochemical and isotopic analyses of the Cretaceous–Paleogene (K/Pg) boundary deposits were conducted at the Caravaca section (External Subbetic, southeast of Spain) in order to evaluate the recovery of the macrobenthic tracemaker community and the bioturbational disturbance. Samples from the infilling material of several lower Danian dark-colored trace fossils (Chondrites, Planolites, Thalassinoides and Zoophycos) located in the uppermost 8-cm of the light upper Maastrichtian strata, as well as samples from the host sedimentary rock of these trace fossils, were analyzed and compared with data from the lower Danian deposits. The values of element ratios indicative of extraterrestrial contamination (Cr/Al, Co/Al and Ni/Al) are higher in the infilling trace fossil material than in the upper Maastrichtian and lower Danian deposits, which suggests a contribution of the ejecta layer. Regarding the isotope composition, the δ¹³C values are lower in the infilling material than in the Maastrichtian host sedimentary rocks surrounding the traces, while the δ¹⁸O are higher in the infilling material. The geochemical and isotopic compositions of the infilling material evidence the unconsolidated character of the sediment, including the red boundary layer. Softground conditions confirm a relatively rapid recovery by the macrobenthic tracemaker community, starting a few millimeters above the K/Pg boundary layer. The mixture of the infilling material of the trace fossils moreover reveals a significant macrobenthic tracemaker activity affecting K–Pg boundary transition sediments that may have significantly altered original signatures.     

Post‐impact event bed (tsunamite) at the Cretaceous–Palaeogene boundary deposited on a distal carbonate platform interior

We show crucial evidence for the Cretaceous–Palaeogene (K–Pg) boundary event recorded within a rare succession deposited in an inner‐platform lagoon on top of a Mesozoic, tropical, intra‐oceanic (western Tethys) Adriatic carbonate platform, which is exposed at Likva cove on the island of Bra? (Croatia). The last terminal Maastrichtian fossils appear within a distinct 10–12 cm thick event bed that is characterised by soft‐sediment bioturbation and rare shocked‐quartz grains, and is interpreted as a distal tsunamite. Directly overlying this is a 2 cm thick reddish‐brown clayey mudstone containing planktonic foraminifera typical of the basal Danian, and with elevated platinum‐group elements in chondritic proportions indicating a clear link to the Chicxulub asteroid impact. These results strongly support the first discovery of a “potential” K–Pg boundary tsunamite on the neighbouring island of Hvar, and these two complementary sections represent probably the most complete record of the event among known distal shallow‐marine successions.     

A New Sapeornithid Bird from China and Its Implication for Early Avian Evolution

<正>Recent discoveries of basal birds have greatly improved our understanding of early bird evolution,yet the evolution of several important features such as cranial kinesis and arboreality remain debated.A new sapeornithid bird,Shenshiornis primita gen.et sp.nov.,based on an articulated skeleton from the Early Cretaceous Jehol Group of western Liaoning,China,sheds new light on these issues.Shenshiornis possesses a skull as primitive as or even more primitive than that of Archaeopteryx and hind limbs modified for an improved arboreal capability.A cladistic analysis shows that:1) presence of a diapsid skull is a plesiomorphy of the Aves and a kinetic skull evolved incrementally later in avian evolution;and 2) cursorial capability significantly weakens at the base of the Pygostylia due to a change in locomotor system.     

中国发现世界上最原始的鸟类化石：华美金凤鸟（新属、新种）

始祖鸟的发现被认为是19世纪最重要的科学发现之一,迄今为止共发现了7枚保存骨骼、羽毛印痕的化石标本和1枚单根羽毛化石标本。由于始祖鸟既显示了明显的爬行动物的特征又保存了精美的羽毛,140多年来人们一直将其作为介于恐龙与鸟类之间的“中间环节”(“missing link”),但更倾向认为它是世界上最古老的鸟,或鸟类的始祖。由于始祖鸟化石稀少,加之地理分布十分局限,长期以来国际上一直围绕鸟类起源问题展开了激烈的争论,提出了各种各样的假说。直到20世纪90年代,在中国辽西晚中生代地层中发现了大量长羽毛的恐龙和原始鸟类化石,有力地支持了鸟类起源于小型兽脚类恐龙的学术观点,使赫 140多年前提出的“假说”成为当今国际科学界占主流地位的学说和理论,基本解决了长期困扰国际科学界的鸟类起源问题,遗憾的是,在过去的几年中,中国一直没有发现与德国始祖鸟十分接近或与德国始祖鸟处于同样进化水平的原始鸟类化石,这种缺憾使我们难以对鸟类定义问题进行深入的研究和探讨,2002年,笔者等在中国辽西早白垩世断层中发现了2种初鸟类化石,即中华神州鸟和东方吉祥鸟。通过比较解剖学研究和支序分析,这2种初鸟类要比始祖鸟略微进步一些,在研究鸟类的早期演化方面具有重要意义;但仍然难以对鸟类定义问题作进一步研究。本文首次报道了一件采自中国河北省丰宁县龙凤山桥头组的初鸟类化石——华美金凤鸟（新属、新种）(Jinfengopteryx elegans gen.et sp.nov),与其共生的化石有Lycoptera davidi,Peipiaosteus pani,yanosteus longidorsalis（鱼类）;Jibeinia luanhera,Protopteryx fengningensis(鸟类）;Ephemeropsis trisetalis(昆虫）及少量植物。支序分析表明,华美金凤鸟处于初鸟类谱系树的基部,比始祖鸟略微原始,与后者为姊妹群关系。奔龙等兽脚类恐龙是它们的外类群。金凤鸟、始祖鸟是中华神州鸟等原始鸟类的外类群。这说明华美金凤鸟处在恐龙向鸟类演化过程中一个非常关键的位置,是世界上迄今所发现的最原始的初鸟类。华美金凤鸟的发现在研究鸟类起源、恐龙-鸟类的系统关系等方面具有重要的科学意义。     

辽宁北票四合屯地区早白垩世早期鸟类集群死亡原因探析

通过大规模系统挖掘,在对四合屯7个含鸟层进行澄清的基础上,以四合屯剖面为主,黄半吉沟剖面为辅,分别在含鸟层的底板、含鸟层、含鸟层顶板,不含鸟沉积层及含鸟层底部的火山岩系统取样,进行多项相关元素分析。应用元素地球化学理论,根据其环境地球化学场特征,结合相关地质资料,重点对鸟类集群死亡原因进行了探讨。结果表明包括二氧化硫(SO2)、硫化氢(H2S)、铅(Pb)的卤化物盐(PbBr2、PbBrCl、Pb(OH)Br、(PbO)2PbBr2)及汞(Hg)蒸汽等有毒气体是导致鸟类集群死亡的主要原因;含鸟化石密集的原因是由于火山口在四合屯附近。     

A true carinate bird from lower Cretaceous deposits in Mongolia and other evidence of early Cretaceous birds in Asia

A partial, associated skeleton with feather impressions of a bird from early Cretaceous (Neocomian) deposits in Mongolia constitutes a new family and order (Ambiortidae, Ambiortiformes). The specimen presents a mosaic of archaic and specialised characters within the Class Aves and shows that advanced carinate birds existed some 10 to 12 million years after Archaeopteryx, lending tentative support to the idea that Archaeopteryx may not be representative of the state of avian evolution in the late Jurassic. The new specimen and numerous feather impressions from other early Cretaceous localities in Mongolia and the Soviet Union, indicate that birds were probably common in early Cretaceous biotas.     

鄂西利川齐岳山高陡背斜带的古应力分析

鄂西利川地区位于湘鄂西构造带与川东构造带的过渡部位,叠加褶皱发育,地处两大构造带分界处的齐岳山高陡背斜带断裂发育。本文以利川地区褶皱和断裂为研究对象,在野外观测和分析的基础上,采用断层滑动数据反演方法,对构造应力场进行了恢复;结合区域构造演化历史,提出该区侏罗纪以来经历了五期构造应力作用,从早到晚分别为:北西-南东向挤压(J3-K1)、近东西向挤压(K1)、近南北向挤压(K1-K2)、北西-南东向引张(K2)和北东-南西向挤压(E3)。该区侏罗纪以来构造变形序列的建立,为深入认识齐岳山高陡背斜带地质灾害形成的地质背景提供了构造地质学证据。     

塔里木盆地麦盖提斜坡皮山北新1井白云质角砾岩地层时代厘定

本文结合岩石薄片、古生物孢粉、同位素、铂族元素、野外露头剖面等资料,对塔里木盆地麦盖提斜坡皮山北新1井钻遇的一套白云质角砾岩地层时代进行了厘定。结果认为：6 884~7 068 m段归属古近系吐依洛克组,化石含量较少;7 068~7 150 m段归属白垩系依格孜牙组,孢粉化石丰富。该井古近纪与白垩纪地层界线深度7 068 m,该位置碳同位素的突变指示海洋环境的剧烈变化,这可以与全球K/E界线处对比。同时,铂族元素在该深度见到明显的异常,其含量明显增大,尤其以Rh、Ru、Ir、Os等最为明显。通过此次厘定,丰富了对麦盖提斜坡晚白垩世—古近纪早期沉积格局的认识。     

大兴安岭─燕山地层分区中、新生代地层

对大兴安岭和燕山北部的中生代地层，进行了系统的总结。侏罗纪早中期为含煤地层，晚期为以兴安岭群为代表的火山堆积；白垩纪早期为火山－沉积含煤地层，晚期为红层。第三纪以二连盆地合大量哺乳动物群的杂色沉积为特征。     

鲁东地区白垩纪地层格架及白垩系与古近系分界问题的探讨

鲁东地区是我国研究晚白垩世和早古近世地层的经典地区,研究历史可追溯到20世纪20年代,积累了大量有关白垩纪和古近纪地层划分与对比的资料。主要介绍在胶州市西郊实施的鲁科一井所揭示的白垩-古近系界线地层的剖面资料。根据岩芯剖面的研究,该区白垩-古近系界线地层自下而上可分为红土崖组、史家屯组和胶州组。对红土崖组的含义重新作了厘定,该组是指介于下伏地层辛格庄组与上覆地层史家屯组之间的1套以产丰富恐龙化石为特征的洪积相、冲积相红色碎屑沉积,客观上包括了原来“金岗口组”的地层。史家屯组是重新恢复使用的一个岩石地层单位,由原先的红土崖组中的“史家屯玄武岩段”提升而来,该组位于王氏群顶部,为一套火山-沉积岩系,厚度可达970余m,与下伏地层红土崖组和上覆地层胶州组均为整合接触。胶州组以1套紫红色粉砂岩为主,夹灰绿色-黄绿色粉砂岩、泥灰岩及少量砂砾岩,与下伏地层王氏群史家屯组为整合接触,但与上覆地层的关系仍不清楚。此外还对鲁东地区晚白垩世地层的划分与对比、恐龙动物群的分布、地质事件、古气候和古环境等进行了探讨,提出白垩-古近纪之交时期,鲁东地区经历了这样一个过程：恐龙繁盛—恐龙消亡—火山喷发—地外撞击（?）—森林大火—白垩纪生物灭绝—古近纪生物群复苏。该过程表明：（1） 将鲁东地区白垩-古近系界线置于胶州组172—148层的这段地层中是合适的;（2） 白垩纪末期生物灭绝与地外撞击事件的关系可能比较密切,而非鸟型恐龙的消亡看来与地外撞击事件的关系不大;（3） 从世界范围来看,非鸟型恐龙均消亡于白垩纪末期撞击事件之前,目前还没有任何证据证明陆地上非鸟型恐龙的消亡与该事件直接相关;（4） 白垩纪晚期曾发生过一系列地质事件,这些事件导致当时的气候、生态、环境等逐渐恶化,生物逐渐失去它们原有的家园,数量和多样性逐步减少,一步一步走上不归之路。非鸟型恐龙也是一样,它们的消亡不是一个事件造成的,而是阶步式的,是多个事件的影响叠加造成的结果。