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     

鸟类起源与演化百问百答（上）

一、起源篇１ 鸟类起源于恐龙吗 ?目前国际上多数学者已同意鸟类确实起源于一类小型的兽脚类恐龙 ,但仍有一些学者持不同看法。２ “鸟就是恐龙 ,恐龙就是鸟”这句话对吗 ?如果承认鸟类起源于恐龙 ,那么说鸟就是恐龙就相当于说人是哺乳动物 ,从系统关系和分类的角度来讲 ,这句话基本算对 ,但严格来说 ,也不对 ,因为鸟类和恐龙毕竟存在差别正如人和哺乳动物存在很大不同的道理一样 ;但是说恐龙就是鸟则相当于说哺乳动物就是人 ,显然是不对的。３ 鸟类的起源有几大假说 ?历史上关于鸟类的起源 ,主要存在过三种主要的假说 :一是鸟类的恐龙起源…     

被视为恐龙的鸟

尽管于 2 0世纪 2 0年代在亚洲地表层首次发现长有羽毛的恐龙和著名的“龙骨突位点” ,关于鸟类起源的争论仍没有休止。来自化石的证据表明 ,鸟类在进化分支上应归于兽脚类的特殊分支。本文主要阐明完好无损的化石揭示的鸟和非鸟类恐龙的亲密关系和鸟类羽毛及鸟类出现以前的羽毛的起源证据 ,分析体型缩小对飞行进化的重要意义及从新的角度论述鸟类如何飞上了天     

鸟类起源的百问百答(中)

三、鸟类篇３２ 中华龙鸟是鸟吗？中华龙鸟不是鸟 ,而是一类典型的兽脚类恐龙 ,这一点国际上从未有过任何怀疑。中华龙鸟带有的毛状物质不具有鸟类羽毛的基本特征 (如分叉和羽轴 ) ,因此还根本不能称为羽毛。如果因为它具有类似羽毛的特征而称之为鸟 ,这就好比因为鹦鹉会学说话可以被成为人的道理是一样。３３ .尾羽龙 (尾羽鸟 )是鸟吗 ?尾羽龙最初是作为带有真正羽毛的恐龙发表的。对于许多古生物学家来说 ,尾羽龙的主要意义也在于它和原始祖鸟一起第一次证明恐龙具有和鸟类一样的羽毛结构。但是 ,不是所有的学者都同意尾羽龙就是恐龙 ,他…     

我国发现1.2亿年前长着恐龙头骨的鸟类新属种——朱氏克拉通鸷

<正>中生代记录了鸟类如何从恐龙演化出来，并演化出独有的体型特征的过程，这一演化阶段鸟类谱系的多样性主要以反鸟类和今鸟型类构成的鸟胸类为主，而鸟胸类已经演化出大量与现生鸟类相似的形态特征，与最原始的鸟类（始祖鸟）在形态上差异巨大。演化位置介于二者之间的非鸟胸类鸟类（以下简称基干鸟类）则为填补这一鸿沟提供了重要信息，然而，长期以来受限于化石的发现，科学家对基干鸟类早期分异的研究仍十分罕见。     

鸟类羽毛的扫描电镜观察

羽毛是鸟类特有的表皮衍生物,它对保温和完成飞行作用负有特殊功能。以往虽对正羽、绒羽和纤羽做过光镜描述,但因分辨力所限,很难详细描述出在执行功能中,鸟羽结构复杂性的功能意义。为了进一步说明问题,本文采用了扫描电镜观察法,以期为鸟羽结构与功能的相互关系提供一些理论依据。     

鹏鸟(Pengornis)一新材料及其对反鸟特征演化的指示意义（英文）

鹏鸟(Pengornis)是早白垩世已知体型最大的反鸟类,其骨骼兼有反鸟类和今鸟类的特征。报道了辽西九佛堂组新发现的一件鹏鸟的亚成年个体标本,代表了该属鸟类除侯氏鹏鸟(Pengornis houi)正型标本外的已知第二件标本,暂归入鹏鸟未定种(Pengornis sp.)。该标本头骨与头后骨骼近乎完整保存,并附有羽毛印痕,仅缺失部分右前肢和部分左后肢。新标本首次提供了鹏鸟胸骨与基干反鸟类原羽鸟(Protopteryx)及基干今鸟类古喙鸟(Archaeorhynchus)相似的形态特征,肯定了鹏鸟的基干位置,并讨论了其在鸟类胸骨演化中的意义。新标本对前肢和后肢(特别是脚趾)等的许多特征也有补充,表明其应当属于树栖生活的鸟类。     

寻找”飞翔”的恐龙

在我们的一般印象中,恐龙是一种身披鳞片、体形巨大的动物,然而事实也许并不如此,至少有一些种类的恐龙体形小巧、动作灵活,而且长有羽毛。那么,这些长羽毛的恐龙与鸟类起源有什么关系？它们是否就是鸟类的祖先？这些问题一直是进化生物学研究人员关注的焦点,也是争议最多的课题。     

寻找”飞翔”的恐龙

在我们的一般印象中,恐龙是一种身披鳞片、体形巨大的动物,然而事实也许并不如此,至少有一些种类的恐龙体形小巧、动作灵活,而且长有羽毛。那么,这些长羽毛的恐龙与鸟类起源有什么关系？它们是否就是鸟类的祖先？这些问题一直是进化生物学研究人员关注的焦点,也是争议最多的课题。     

科学家发现鸟类起源于恐龙的最新证据

鸟类的起源一直是国际古生物学界的热点问题之一,自上个世纪60年代末至70年代初,美国耶鲁大学教授约翰·奥斯罗姆复兴了赫胥黎提出的鸟类兽脚类恐龙起源假说后,鸟类起源的研究取得了巨大的进展,来自世界各地不同地史时期的恐龙和早期鸟类的大量化石证据持续不断地出现,有力地支持着这一假说。1986年美国耶鲁大学教授嘉克斯·高斯特首次用分支系统学的方法系统地分析了鸟类与其他初龙类（一个包括恐龙、翼龙、鳄型动物以及一些绝灭支系的爬行动物类群）的关系,     

Evolution of dinosaur epidermal structures

Spectacularly preserved non-avian dinosaurs with integumentary filaments/feathers have revolutionized dinosaur studies and fostered the suggestion that the dinosaur common ancestor possessed complex integumentary structures homologous to feathers. This hypothesis has major implications for interpreting dinosaur biology, but has not been tested rigorously. Using a comprehensive database of dinosaur skin traces, we apply maximum-likelihood methods to reconstruct the phylogenetic distribution of epidermal structures and interpret their evolutionary history. Most of these analyses find no compelling evidence for the appearance of protofeathers in the dinosaur common ancestor and scales are usually recovered as the plesiomorphic state, but results are sensitive to the outgroup condition in pterosaurs. Rare occurrences of ornithischian filamentous integument might represent independent acquisitions of novel epidermal structures that are not homologous with theropod feathers.     

Development and evolution of the mammalian limb: adaptive diversification of nails, hooves, and claws

SUMMARY Paleontological evidence indicates that the evolutionary diversification of mammals early in the Cenozoic era was characterized by an adaptive radiation of distal limb structures. Likewise, neontological data show that morphological variation in distal limb integumentary appendages (e.g., nails, hooves, and claws) can be observed not only among distantly related mammalian taxa but also among closely related species within the same clade. Comparative analysis of nail, claw, and hoof morphogenesis reveals relatively subtle differences in mesenchymal and epithelial patterning underlying these adult differences in distal limb appendage morphology. Furthermore, studies of regulatory gene expression during vertebrate claw development demonstrate that many of the signaling molecules involved in patterning ectodermal derivatives such as teeth, hair, and feathers are also involved in organizing mammalian distal limb appendages. For example, Bmp4 signaling plays an important role during the recruitment of mesenchymal cells into the condensations forming the terminal phalanges, whereas Msx2 affects the length of nails and claws by suppressing proliferation of germinal epidermal cells. Evolutionary changes in the form of distal integumentary appendages may therefore result from changes in gene expression during formation of mesenchymal condensations ( Bmp4 , posterior Hox genes), induction of the claw fold and germinal matrix ( shh ), and/or proliferation of epidermal cells in the claw matrix ( Msx1 , Msx2 ). The prevalence of convergences and parallelisms in nail and claw structure among mammals underscores the existence of multiple morphogenetic pathways for evolutionary change in distal limb appendages.     

Dinosaur protofeathers: pushing back the origin of feathers into the Middle Triassic?

Reports of primordial feathers (protofeathers) in dinosaurs have received widespread interest. Recently, it was proposed that a novel protofeather in the theropod dinosaur Beipiaosaurus completes the transitional series in the evolution of the feather and provides the first evidence of filamentous feathers as display in nonavian theropods. A more far-reaching evolutionary ramification is the claim that these structures push the origin of monofilamentous integumentary structures into the Middle Triassic or earlier. I discuss problems with the analyses within the broader context of studies concerning the hypothesis of protofeathers, and show that affinity between the integumentary structures in Beipiaosaurus and feathers is improbable. The scientific methodology is questioned by its failure to make phenomena perceivable by objective means, by questionable rationalizations in critical issues, and by lack of consideration of exceptions to the postulated thesis. The notion that primordial feathers occurred in a clade more inclusive than the Coelurosauria and that it is supported by the presence of integumental structures in Psittacosaurus is analyzed and rejected.     

The evolution of sexually dimorphic tail feathers is not associated with tail skeleton dimorphism

Sexual selection can influence the evolution of sexually dimorphic exaggerated display structures. Herein, we explore whether such costly ornamental integumentary structures evolve independently or if they are correlated with phenotypic change in the associated skeletal system. In birds, elongate tail feathers have frequently evolved in males and are beneficial as intraspecific display structures but impart a locomotor/energetic cost. Using the sexually dimorphic tail feathers of several passeriform species as a model system, we test the hypothesis that taxa with sexually dimorphic tail feathers also exhibit sexual dimorphism in the caudal skeleton that supports the muscles and integument of the tail apparatus. Caudal skeletal morphology is quantified using both geometric morphometrics and linear morphometrics across four sexually dimorphic passeriform species and four closely related monomorphic species. Sexual dimorphism is assessed using permutational MANOVA. Sexual dimorphism in caudal skeletal morphology is found only in those taxa that exhibit active functional differences in tail use between males and females. Thus, dimorphism in tail feather length is not necessarily correlated with the evolution of caudal skeletal dimorphism. Sexual selection is sufficient to generate phenotypic divergence in integumentary display structures between the sexes, but these change are not reflected in the underlying caudal skeleton. This suggests that caudal feathers and bones evolve semi‐independently from one another and evolve at different rates in response to different types of selective pressures.     

On the purported presence of fossilized collagen fibres in an ichthyosaur and a theropod dinosaur

Since the discovery of exceptionally preserved theropod dinosaurs with soft tissues in China in the 1990s, there has been much debate about the nature of filamentous structures observed in some specimens. Sinosauropteryx was the first non‐avian theropod to be described with these structures, and remains one of the most studied examples. Despite a general consensus that the structures represent feathers or feather homologues, a few identify them as degraded collagen fibres derived from the skin. This latter view has been based on observations of low‐quality images of Sinosauropteryx, as well as the suggestion that because superficially similar structures are seen in Jurassic ichthyosaurs they cannot represent feathers. Here, we highlight issues with the evidence put forward in support of this view, showing that integumentary structures have been misinterpreted based on sedimentary features and preparation marks, and that these errors have led to incorrect conclusions being drawn about the existence of collagen in Sinosauropteryx and the ichthyosaur Stenopterygius. We find that there is no evidence to support the idea that the integumentary structures seen in the two taxa are collagen fibres, and confirm that the most parsimonious interpretation of fossilized structures that look like feather homologues in Sinosauropteryx is that they are indeed the remains of feather homologues.     

Origin of archosaurian integumentary appendages: the bristles of the wild turkey beard express feather-type beta keratins

The discovery that structurally unique "filamentous integumentary appendages" are associated with several different non-avian dinosaurs continues to stimulate the development of models to explain the evolutionary origin of feathers. Taking the phylogenetic relationships of the non-avian dinosaurs into consideration, some models propose that the "filamentous integumentary appendages" represent intermediate stages in the sequential evolution of feathers. Here we present observations on a unique integumentary structure, the bristle of the wild turkey beard, and suggest that this non-feather appendage provides another explanation for some of the "filamentous integumentary appendages." Unlike feathers, beard bristles grow continuously from finger-like outgrows of the integument lacking follicles. We find that these beard bristles, which show simple branching, are hollow, distally, and express the feather-type beta keratins. The significance of these observations to explanations for the evolution of archosaurian integumentary appendages is discussed.     

The Control of Color in Birds

SYNOPSIS. The colors of birds result from deposition of pigments—mainlymelanins and carotenoids—in integumentary structures,chiefly the feathers. The plumages of birds indicate their age,sex, and mode of living, and play important roles in camouflage,mating, and establishment of territories. Since feathers aredead structures, change of color of feathers is effected throughdivestment (molt) and replacement. The color and pattern ofa feather are determined by the interplay of genetic and hormonalinfluences prevailing in its base during regeneration. Mostbirds replace their feathers at least once annually. Some wearthe same kind of basic plumage all the time butothers alternatea basic and breeding plumage, either in one (the male) or bothsexes. Still others may have more than two molts, adding supplementalplumage at certain times in the plumage cycle. The varietiesof patterns of molt, the kinds of plumage, and the colors andpatterns of feathers among birds apparently are the result ofseveral kinds of selection pressures working through evolution.     

Hox genes as synchronized temporal regulators: implications for morphological innovation

This special issue on the development and evolution of the amniote integument begins with a discussion of the adaptations to terrestrial conditions, the acquisition of water-impermeability of the reptilian integument, and the initial formation of filamentous integumentary appendages that prepare the way towards avian flight. Recent feather fossils are reviewed, and a definition of feathers is developed. Hierarchical models are proposed for the formation of complex structures, such as feathers. Molecular signals that alter the phenotype of integumentary appendages at different levels of the hierarchy are presented. Tissue interactions and the roles of keratins in evolution are discussed and linked to their bio-mechanical properties. The role of mechanical forces on patterning is explored. Elaborate extant feather variants are introduced. The regeneration/gene mis-expression protocol for the chicken feather is established as a testable model for the study of biological structures. The adaptations of the mammalian distal limb end organs to terrestrial, arboreal and aquatic conditions are discussed. The development and cycling of hair are reviewed from a molecular perspective. These contributions reveal that the structure and function of diverse integumentary appendages are variations that are superimposed on a common theme, and that their formation is modular, hierarchical and cyclical. They further reveal that these mechanisms can be understood at the molecular level, and that an integrative and organismal approach to studying integumentary appendages is called for. We propose that future research should foster interdisciplinary approaches, pursue understanding at the cellular and molecular level, analyze interactions between the environment and genome, and recognize the contributions of variation in morphogenesis and evolution.     

Horse hooves and bird feathers: Two model systems for studying the structure and development of highly adapted integumentary accessory organs--the role of the dermo-epidermal interface for the micro-architecture of complex epidermal structures

Accessory organs of the integument are locally modified parts of the potentially feather-bearing skin in birds (e.g., the rhamphotheca, claws, or scales), and of the potentially hairy skin in mammals (e.g., the rhinarium, nails, claws, or hooves). These special parts of the integument are characterised by a modified structure of their epidermal, dermal and subcutaneous layers. The developmental processes of these various integumentary structures in birds and mammals show both similarities and differences. For example, the development of the specialised epidermal structures of both feathers and the hoof capsule is influenced by the local three-dimensional configuration of the dermis. However, in feathers, in contrast to hooves, the arrangement of the corneous cells is only partially a direct result of the particular arrangement and shape of the dermal surface of the papillary body. Whereas the diameter of the feather papilla, as well as the number, length, and width of dermal ridges on the surface of the feather papilla influence the three-dimensional architecture of the feather rami, there is no apparent direct correlation between the dermo-epidermal interface and the development of the highly ordered architecture of the radii and hamuli in the feather vane. In order to elucidate this morphogenic problem and the problem of locally different processes of keratinisation and cornification, the structure and development of feathers in birds are compared to those of the hoof capsule in horses. The equine hoof is the most complex mammalian integumentary structure, which is determined directly by the dermal surface of the papillary body. Perspectives for further research on the development of modified integumentary structures, such as the role of the dermal microangioarchitecture and the selective adhesion and various differentiation pathways of epidermal cells, are discussed.     

The colour of fossil feathers

Feathers are complex integumentary appendages of birds and some other theropod dinosaurs. They are frequently coloured and function in camouflage and display. Previous investigations have concluded that fossil feathers are preserved as carbonized traces composed of feather-degrading bacteria. Here, an investigation of a colour-banded feather from the Lower Cretaceous Crato Formation of Brazil revealed that the dark bands are preserved as elongate, oblate carbonaceous bodies 1-2mum long, whereas the light bands retain only relief traces on the rock matrix. Energy dispersive X-ray analysis showed that the dark bands preserve a substantial amount of carbon, whereas the light bands show no carbon residue. Comparison of these oblate fossil bodies with the structure of black feathers from a living bird indicates that they are the eumelanin-containing melanosomes. We conclude that most fossil feathers are preserved as melanosomes, and that the distribution of these structures in fossil feathers can preserve the colour pattern in the original feather. The discovery of preserved melanosomes opens up the possibility of interpreting the colour of extinct birds and other dinosaurs.