中国中生代的鸟类:介绍及综述

最近十来年 ,中国辽宁发现的早白垩世的鸟类化石超过了世界上其它任何一个地区。中国的中生代鸟类化石代表了始祖鸟化石之后鸟类历史上第一次显著的分异。它们不仅包括了带有明显恐龙祖先特征的长尾的鸟类 ,而且还包括了许多进步或特化的种类 ,如早白垩世最大的鸟类 ,最原始的反鸟类 ,以及保存最好的、飞行结构和现生鸟类几乎一样的今鸟类。这些早期鸟类在诸如飞行、大小和食性等所反映的演化、形态和生态学特征等方面出现了重大的分异。具有长尾骨骼的原始基干鸟类热河鸟和驰龙类具有的相似性 ,进一步支持了鸟类起源于恐龙的学说。中国发现的早白垩世的鸟类以及树栖的恐龙化石还为鸟类飞行的树栖起源假说提供了十分重要的证据。“恐龙下树”的假说结合了鸟类起源于恐龙的学说和鸟类飞行的树栖起源学说 ,因此也得到了化石证据的支持。由于多种恐龙带有羽毛 ,因此羽毛不一定代表了恒温。恒温的鸟类可能到了早白垩世的进步鸟类中才开始出现     

中生代反鸟类研究的新进展

反鸟类是繁盛于中生代的鸟类类群,它们在鸟类演化史上第一次成功地实现了全球性辐射,化石发现于除极地以外的各大陆白垩纪地层中,目前全世界被命名的反鸟类1属种超过了50个,其中我国辽西早白垩世"热河生物群"被命名的反鸟类新属种占总数的一半以上。最新的研究表明,在白垩期早期反鸟类的个体大小和相对翼长已有很大的分化,肩带和前肢上与飞行相关的骨骼特征已与现代鸟类基本相似,进步程度甚至高于同时期的今鸟类,表明适应飞行的演化已达到了很高的水平;吻部和牙齿的改变也较同时期的今鸟类多样,说明在食性也有了大的分异。新的发现也进一步减少了反鸟类和今鸟类——这两个中生代进步鸟类姊妹群之间的特征差异。     

热河生物群中两类飞行脊椎动物——翼龙与鸟类的生态竞争

早在1.2亿年前的早白垩世,辽西及其周边地区不但发现了大量的鱼类、两栖类、带羽毛恐龙、哺乳动物,同时也发现了数量惊人的早期鸟类、翼龙化石,以及各种陆生无脊椎动物(包括大量昆虫)和植物(包括被子植物)化石等等。这些化石的发现,揭示了白垩纪地球生命演化历史,同时,也为人们     

翼龙蛋与胚胎化石揭秘翼龙生命史

正翼龙是地球上第一类真正飞行也是唯一绝灭的飞行脊椎动物。由于飞行的需要,翼龙演化出了纤细中空的骨骼,因此翼龙化石十分稀少,而翼龙蛋和胚胎化石更是罕见。经过十余年连续的野外科考工作,中国科学院古脊椎动物与古人类研究所汪筱林等人在新疆哈密戈壁早白垩世地层中发现并抢救性采集了一件超过200枚翼龙蛋、胚胎和骨骼化石三维一体保存的重要标本,     

辽宁早白垩世今鸟类一新属种(Jianchangornis microdonta gen.et sp.nov.)(英文)

依据一近完整的相关节的骨骼化石,记述了辽宁建昌早白垩世九佛堂组原始今鸟类一新属种:小齿建昌鸟(Jianchangornis microdonta gen.et sp.nov.)。新鸟个体较大,但从骨化程度分析,正型标本可能属于一亚成年个体。具有一些进步特征,如胸骨及龙骨突加长,乌喙骨具有发育的前乌喙突以及和肩胛骨关联的关节窝,叉骨"U"字型,愈合荐椎包括9-10枚荐椎,尾综骨短小,第二、三掌骨远端愈合,跗跖骨完全愈合等,表明新属无疑属于今鸟类。在以下特征组合上很容易和已知的早白垩世今鸟类化石相区别:齿骨上至少有16枚细小牙齿,从齿骨前端向后沿齿骨大部密集排列;肩胛骨强烈弯曲;第一掌骨粗壮,较其他掌骨宽;第一指长并且远端延伸明显超过第二掌骨;肱骨+尺骨+第二掌骨与股骨+胫跗骨+跗跖骨的长度比例约为1.1。系统发育分析表明新属属于基干的今鸟类。新发现的材料第二、三掌骨远端愈合很好,但近端却未完全愈合,这一特征尚未见于其他已知鸟类,或许表明今鸟类腕掌骨的愈合和现生鸟类的跗跖骨一样是从远端开始的,不同于反鸟类和其他基干鸟类。建昌鸟的下颌还保存了一个前齿骨,这是继早白垩世红山鸟之后的另一例报道,可能进一步表明这一结构在今鸟类中曾普遍出现。新鸟肩带、胸骨和前肢的特征显示了和现代鸟类相近的飞行能力,其后肢、脚趾的比例以及趾爪的形态等显示和燕鸟、义县鸟等相似的地栖特征。保存于标本上的鱼类残骸可能显示了建昌鸟食鱼类的习性。今鸟类新属种的发现进一步表明,早白垩世这一进步鸟类类群的分化已不亚于反鸟类,湖滨环境在今鸟类的早期演化中确实扮演了重要的角色。     

孔子鸟与鸟类的早期演化

大量孔子乌标本的出现,使得我们能够识别这一早期鸟类的一些前所未知的形态特征。依据这些特征,并结合近年来中国发现的其他中生代鸟类资料,我们可以进一步探讨早期鸟类研究中存在的某些问题,诸如鸟类飞行的起源等。孔子鸟头骨的最典型特征是进步的角质像的出现和原始眶后骨的完全保留。孔子鸟的眶后骨和颧骨相连接,这一特征尚未见于其他鸟类。由于始祖鸟也已缺失了眶后骨,因此眶后骨在孔子鸟中的存在,进一步证实了始祖鸟在早期鸟类演化中的旁支地位,同时也表明,鸟类真正的祖先比我们以往所了解的更加原始。孔子鸟保留了三个指爪。第三指爪（中间一个）常较退化,这和飞羽附着第三指有关。发育的趾爪及指爪,显示孔子鸟适应攀援树木的生活。基本愈合的尾椎和尚不完善的飞行器官,不仅支持这一假设,而且可能还表明,孔子鸟尚不能从地面起飞。孔子鸟尚未发育小冀羽,这也更加证实了其飞行的原始性。孔子鸟的某些个体,保存一对长的尾羽,这可能代表雄性的特征。另一些个体的头部还保留装饰性羽毛。数百件个体的集中发现或许还表明,孔子鸟具备了某些现生鸟类集群性的行为方式。个体大小的变化在早期鸟类飞行能力演化中的作用十分显著。早白垩世的反鸟类的成员都明显小于始祖鸟和孔子鸟,加之拥?     

研究揭示利于鸟类取食活动的头骨特征出现于鸟类演化初期

<正>2015年8月22日,在线出版的《系统古生物学》上,中国科学院古脊椎动物与古人类研究所的王敏、胡晗,以及得克萨斯大学的李志恒报道了早白垩世一反鸟类新属种,并讨论了鸟类头骨的早期演化,首次发现有利于鸟类取食活动的头骨特征在鸟类演化初期就已经出现。据王敏介绍,早白垩世的反鸟类标本虽然     

温血动物最直接的证据──皮毛在翼龙中发现

温血动物最直接的证据──皮毛在翼龙中发现吕君昌作为飞行爬行类的翼龙,由于其特殊的构造（前肢特化为翼,第四指为加长的飞行指,骨骼中空,胸骨发育成一个特殊的胸骨突－龙骨,等等这些适用于飞行的特征）和可能与现生的鸟类和蝙蝠类似的生活习性,而鸟类和蝙蝠类均为...     

鸟类起源于基干的初龙类

中亚三叠纪和中国早白垩世的新的重要化石发现或许可以帮助解决有关鸟类起源的争议。鸟类的飞行可能起源于三叠纪一些小型的、四脚滑翔的初龙类。羽毛的起源最初是为了滑翔而不是保温。“手盗龙类”实际上起源于鸟类 ,并至少发育了初级飞羽 ,具备滑翔的能力     

辽宁早白垩世早期—鸟化石

本文记述了发现于辽宁朝阳地区早白垩世早期一新的鸟类。这是继三塔中国鸟(Sinornis santensis)、燕都华夏鸟(Cathayomis yandica)之后报道的辽宁中生代第三种鸟化石,它的脊柱和腰带与始祖鸟和恐龙相似,但其肋骨又具有现代鸟类的性状,这是早期鸟类化石的又一新材料。     

Mesozoic birds of China-a synoptic review

A synoptic review of the discoveries and studies of Chinese Mesozoic birds is provided in this paper.40Ar/39Ar dating of several bird-bearing deposits in the Jehol Group has established a geochronological framework for the study of the early avian radiation.Chinese Mesozoic birds had lasted for at least 11 Ma during about 131 Ma and 120 Ma (Barremian to Aptian)of the middle and late Early Cretaceous,respectively.In order to further evaluate the change of the avian diversity in the Jehol Biota,six new orders and families are erected based on known genera and species,which brings the total number of orders of Chinese Mesozoic birds to 15 and highlights a remarkable radiation ever since the first appearante of birds in the Late Jurassic.Chinese Early Cretaceous birds had experienced a significant differentiation in morphology,flight,diet and habitat.Further examination of the foot of Jeholornis suggests this bird might not have possessed a fully reversed hallux.However,the attachment of metatarsal Ⅰ to the medial side of metatarsal Ⅱ does not preclude trunk climbing,a pre-adaptation for well developed perching life of early birds.Arboreality had proved to be a key adaptation in the origin and early evolution of bird flight,and the adaptation to lakeshore environment had played an equally important role in the origin of omithurine birds and their near-modern flight skill.Many Chinese Early Cretaceous birds had preserved the direct evidence of their diet,showing that the most primitive birds were probably mainly insectivorous and that specialized herbivorous or carnivorous (e.g.,piscivorous)dietary adaptation had appeared only in later advanced forms.The only known Early Cretaceous bird embryo fossil has shown that precocial birds had occurred prior to altricial birds in avian history,and the size of the embryo and other analysis indicate it probably had a short incubation period.Leg feathers probably have a wide range of distribution in early birds,further suggesting that leg feathers had played a key role in the beginning stage of the flight of birds.Finally,the Early Cretaceous avian radiation can be better understood against the background of their unique ecosystem.The advantage of birds in the competitions with other vertebrate groups such as pterosaurs had probably not only resulted in the rapid differentiation and radiation of birds but also the worldwide spreading of pterosaurs and other vertebrates from East Asia in the Early Cretaceous.     

Mesozoic birds of China—a synoptic review

A synoptic review of the discoveries and studies of Chinese Mesozoic birds is provided in this paper. ⁴⁰Ar/³⁹Ar dating of several bird-bearing deposits in the Jehol Group has established a geochronological framework for the study of the early avian radiation. Chinese Mesozoic birds had lasted for at least 11 Ma during about 131 Ma and 120 Ma (Barremian to Aptian) of the middle and late Early Cretaceous, respectively. In order to further evaluate the change of the avian diversity in the Jehol Biota, six new orders and families are erected based on known genera and species, which brings the total number of orders of Chinese Mesozoic birds to 15 and highlights a remarkable radiation ever since the first appearance of birds in the Late Jurassic. Chinese Early Cretaceous birds had experienced a significant differentiation in morphology, flight, diet and habitat. Further examination of the foot of Jeholornis suggests this bird might not have possessed a fully reversed hallux. However, the attachment of metatarsal I to the medial side of metatarsal II does not preclude trunk climbing, a pre-adaptation for well developed perching life of early birds. Arboreality had proved to be a key adaptation in the origin and early evolution of bird flight, and the adaptation to lakeshore environment had played an equally important role in the origin of ornithurine birds and their near-modern flight skill. Many Chinese Early Cretaceous birds had preserved the direct evidence of their diet, showing that the most primitive birds were probably mainly insectivorous and that specialized herbivorous or carnivorous (e.g., piscivorous) dietary adaptation had appeared only in later advanced forms. The only known Early Cretaceous bird embryo fossil has shown that precocial birds had occurred prior to altricial birds in avian history, and the size of the embryo and other analysis indicate it probably had a short incubation period. Leg feathers probably have a wide range of distribution in early birds, further suggesting that leg feathers had played a key role in the beginning stage of the flight of birds. Finally, the Early Cretaceous avian radiation can be better understood against the background of their unique ecosystem. The advantage of birds in the competitions with other vertebrate groups such as pterosaurs had probably not only resulted in the rapid differentiation and radiation of birds but also the worldwide spreading of pterosaurs and other vertebrates from East Asia in the Early Cretaceous. Selected from Vertebrata PalAsiatica 2006, 44 (1): 74–98     

High-precision Ar/Ar age for the Jehol Biota

Abundant fossils of the terrestrial Jehol Biota, including plants, insects, dinosaurs, birds, mammals and freshwater invertebrates, were discovered from the Yixian Formation and the overlying Jiufotang Formation in Inner Mongolia, Hebei Province and Liaoning Province, northeastern China. Because of the exceptional preservation of fossils, the Jehol Biota is one of the most important Mesozoic lagerstätten and is referred to as a “Mesozoic Pompeii”. The Jehol Biota has provided a rare opportunity to address questions about the origin of birds, the evolution of feathers and flight, the early diversification of angiosperms and the timing of placental mammal radiation. Six tuff samples and two basalt samples collected from the Tuchengzi, the Yixian and the Jiufotang formations near the classic outcrops in western Liaoning, NE China yielded high-precision ⁴⁰Ar/³⁹Ar ages. We obtain an age of 129.7 ± 0.5 Ma for a basaltic lava from the bottom of the Yixian Formation and an age of 122.1 ± 0.3 Ma for a tuff from the lowermost part of the overlying Jiufotang Formation. Our age results provide an age calibration of the whole Yixian Formation and show that the whole formation was deposited entirely within Early Cretaceous time over an interval of ~ 7 Ma.     

Low ecological disparity in Early Cretaceous birds

Ecological divergence is thought to be coupled with evolutionary radiations, yet the strength of this coupling is unclear. When birds diversified ecologically has received much less attention than their hotly debated crown divergence time. Here, we quantify how accurately skeletal morphology can predict ecology in living and extinct birds, and show that the earliest known assemblage of birds (= pygostylians) from the Jehol Biota (≈ 125 Ma) was substantially impoverished ecologically. The Jehol avifauna has few representatives of highly preservable ecomorphs (e.g. aquatic forms) and a notable lack of ecomorphological overlap with the pterosaur assemblage (e.g. no large or aerially foraging pygostylians). Comparisons of the Jehol functional diversity with modern and subfossil avian assemblages show that taphonomic bias alone cannot explain the ecomorphological impoverishment. However, evolutionary simulations suggest that the constrained ecological diversity of the Early Cretaceous pygostylians is consistent with what is expected from a relatively young radiation. Regardless of the proximate biological explanation, the anomalously low functional diversity of the Jehol birds is evidence both for ecological vacancies in Cretaceous ecosystems, which were subsequently filled by the radiation of crown Aves, and for discordance between taxonomic richness and ecological diversity in the best-known Mesozoic ecosystem.     

The evolution of the modern avian digestive system: insights from paravian fossils from the Yanliao and Jehol biotas

The avian digestive system, like other aspects of avian biology, is highly modified relative to other reptiles. Together these modifications have imparted the great success of Neornithes, the most diverse clade of amniotes alive today. It is important to understand when and how aspects of the modern avian digestive system evolved among neornithine ancestors in order to elucidate the evolutionary success of this important clade and to understand the biology of stem birds and their closest dinosaurian relatives: Mesozoic Paraves. Although direct preservation of the soft tissue of the digestive system has not yet been reported, ingested remains and their anatomical location preserved in articulated fossils hint at the structure of the digestive system and its abilities. Almost all data concerning direct evidence of diet in Paraves comes from either the Upper Jurassic Yanliao Biota or the Lower Cretaceous Jehol Biota, both of which are known from deposits in north-eastern China. Here, the sum of the data gleaned from the thousands of exceptionally well-preserved fossils of paravians is interpreted with regards to the structure and evolution of the highly modified avian digestive system and feeding apparatus. This information suggests intrinsic differences between closely related stem lineages implying either strong homoplasy or that diet in each lineage of non-ornithuromorph birds was highly specialized. Regardless, modern digestive capabilities appear to be limited to the Ornithuromorpha, although the complete set of derived feeding related characters is restricted to the Neornithes.     

The quality of the fossil record of Mesozoic birds

The Mesozoic fossil record has proved critical for understanding the early evolution and subsequent radiation of birds. Little is known, however, about its relative completeness: just how 'good' is the fossil record of birds from the Mesozoic? This question has come to prominence recently in the debate over differences in estimated dates of origin of major clades of birds from molecular and palaeontological data. Using a dataset comprising all known fossil taxa, we present analyses that go some way towards answering this question. Whereas avian diversity remains poorly represented in the Mesozoic, many relatively complete bird specimens have been discovered. New taxa have been added to the phylogenetic tree of basal birds, but its overall shape remains constant, suggesting that the broad outlines of early avian evolution are consistently represented: no stage in the Mesozoic is characterized by an overabundance of scrappy fossils compared with more complete specimens. Examples of Neornithes (modern orders) are known from later stages in the Cretaceous, but their fossils are rarer and scrappier than those of basal bird groups, which we suggest is a biological, rather than a geological, signal.     

Pelvis morphology suggests that early Mesozoic birds were too heavy to contact incubate their eggs

Numerous new fossils have driven an interest in reproduction of early birds, but direct evidence remains elusive. No Mesozoic avian eggs can be unambiguously assigned to a species, which hampers our understanding of the evolution of contact incubation, which is a defining feature of extant birds. Compared to living species, eggs of Mesozoic birds are relatively small, but whether the eggs of Mesozoic birds could actually have borne the weight of a breeding adult has not yet been investigated. We estimated maximal egg breadth for a range of Mesozoic avian taxa from the width of the pelvic canal defined by the pubic symphysis. Known elongation ratios of Mesozoic bird eggs allowed us to predict egg mass and hence the load mass an egg could endure before cracking. These values were compared to the predicted body masses of the adult birds based on skeletal remains. Based on 21 fossil species, we show that for nonornithothoracine birds body mass was 187% of the load mass of the eggs. For Enantiornithes, body mass was 127% greater than the egg load mass, but some early Cretaceous ornithuromorphs were 179% heavier than their eggs could support. Our indirect approach provides the best evidence yet that early birds could not have sat on their eggs without running the risk of causing damage. We suggest that contact incubation evolved comparatively late in birds.     

The patterns and modes of the evolution of disparity in Mesozoic birds

The origin of birds from non-avian theropod dinosaurs is one of the greatest transitions in evolution. Shortly after diverging from other theropods in the Late Jurassic, Mesozoic birds diversified into two major clades—the Enantiornithes and Ornithuromorpha—acquiring many features previously considered unique to the crown group along the way. Here, we present a comparative phylogenetic study of the patterns and modes of Mesozoic bird skeletal morphology and limb proportions. Our results show that the major Mesozoic avian groups are distinctive in discrete character space, but constrained in a morphospace defined by limb proportions. The Enantiornithines, despite being the most speciose group of Mesozoic birds, are much less morphologically disparate than their sister clade, the Ornithuromorpha—the clade that gave rise to living birds, showing disparity and diversity were decoupled in avian history. This relatively low disparity suggests that diversification of enantiornithines was characterized in exhausting fine morphologies, whereas ornithuromorphs continuously explored a broader array of morphologies and ecological opportunities. We suggest this clade-specific evolutionary versatility contributed to their sole survival of the end-Cretaceous mass extinction.