FROM FROND TO FAN: ARCHAEOPTERYX AND THE EVOLUTION OF SHORT-TAILED BIRDS

Modern birds have extremely short tail skeletons relative to Archaeopteryx and nonavialian theropod dinosaurs. Long- and short-tailed birds also differ in the conformation of main tail feathers making up the flight surface: frond shaped in Archaeopteryx and fan shaped in extant fliers. Mechanisms of tail fanning were evaluated by electromyography in freely flying pigeons and turkeys and by electrical stimulation of caudal muscles in anesthetized birds. Results from these experiments reveal that the pygostyle, rectrices, rectricial bulbs, and bulbi rectricium musculature form a specialized fanning mechanism. Contrary to previous models, our data support the interpretation that the bulbi rectricium independently controls tail fanning; other muscles are neither capable of nor necessary for significant rectricial abduction. This bulb mechanism permits rapid changes in tail span, thereby allowing the exploitation of a wide range of lift forces. Isolation of the bulbs on the pygostyle effectively decouples tail fanning from fan movement, which is governed by the remaining caudal muscles. The tail of Archaeopteryx, however, differs from this arrangement in several important respects. Archaeopteryx probably had a limited range of lift forces and tight coupling between vertebral and rectricial movement. This would have made the tail of this primitive flier better suited to stabilization than maneuverability. The capacity to significantly alter lift and manipulate the flight surface without distortion may have been two factors favoring tail shortening and pygostyle development during avian evolution.     

Life history of a basal bird: morphometrics of the Early Cretaceous Confuciusornis

Confuciusornis sanctus stands out among the remarkable diversity of Mesozoic birds recently unearthed from China. Not only is this primitive beaked pygostylian (birds with abbreviated caudal vertebrae fused into a pygostyle) much more abundant than other avian taxa of this age but differences in plumage between specimens--some having a pair of long stiff tail feathers--have been interpreted as evidence for the earliest example of sexual dimorphism in birds. We report the results of a multivariate morphometric study involving measurements of more than 100 skeletons of C. sanctus. Our analyses do not show any correlation between size distribution and the presence or absence of blade-like rectrices (tail feathers), thus implying, that if these feathers are sexual characters, they are not correlated with sexual size dimorphism. Our results also provide insights into the taxonomy and life history of confuciusornithids, suggesting that these birds may have retained ancestral dinosaurian growth patterns characterized by a midlife exponential growth stage.     

Coevolution of caudal skeleton and tail feathers in birds

Birds are capable of a wide range of aerial locomotor behaviors in part because of the derived structure and function of the avian tail. The tail apparatus consists of a several mobile (free) caudal vertebrae, a terminal skeletal element (the pygostyle), and an articulated fan of tail feathers that may be spread or folded, as well as muscular and fibroadipose structures that facilitate tail movements. Morphological variation in both the tail fan and the caudal skeleton that supports it are well documented. The structure of the tail feathers and the pygostyle each evolve in response to functional demands of differing locomotor behaviors. Here, I test whether the integument and skeleton coevolve in this important locomotor module. I quantified feather and skeletal morphology in a diverse sample of waterbirds and shorebirds using a combination of linear and geometric morphometrics. Covariation between tail fan shape and skeletal morphology was then tested using phylogenetic comparative methods. Pygostyle shape is found to be a good predictor of tail fan shape (e.g., forked, graduated), supporting the hypothesis that the tail fan and the tail skeleton have coevolved. This statistical relationship is used to reconstruct feather morphology in an exemplar fossil waterbird, Limnofregata azygosternon. Based on pygostyle morphology, this taxon is likely to have exhibited a forked tail fan similar to that of its extant sister clade Fregata, despite differing in inferred ecology and other aspects of skeletal anatomy. These methods may be useful in reconstructing rectricial morphology in other extinct birds and thus assist in characterizing the evolution of flight control surfaces in birds. J. Morphol. 275:1431–1440, 2014. © 2014 Wiley Periodicals, Inc.     

辽西早白垩世今鸟类一基干有喙新属种(英文)

根据发现于辽宁西部建昌早白垩世九佛堂组一件保存较为完好的标本,描述了原始今鸟类一新属种,李氏叉尾鸟(Schizooura lii gen.et sp.nov.)。系统发育分析表明,它较建昌鸟和古喙鸟进步,但又较热河生物群中其他已知的今鸟类原始。这一新种叉骨呈Ⅴ型,且具有一短的叉骨突,这是这一特征首次在早白垩世今鸟中发现。其胸骨加长,后缘无窗孔,且无很深的凹口。肱骨三角肌脊膨大,近乎达到肱骨长的一半。新鸟的头骨保存较好,显示其具喙的特征,前颌骨直接与额骨相关节。值得关注的是,这件标本首次展示了在中生代鸟类中,除扇形尾羽之外的第二种尾羽类型。新标本具叉形尾羽,中间分隔明显。对现生鸟类的研究表明,叉尾型的空气动力学效率比扇尾型要低,但在性选择方面具有优势。这一新的发现说明,在热河生物群生活的森林环境中,今鸟类的基干种类在尾羽性状上或许已经存在着不同的选择策略。     

LOCOMOTOR MODULES AND THE EVOLUTION OF AVIAN FLIGHT

The evolution of avian flight can be interpreted by analyzing the sequence of modifications of the primitive tetrapod locomotor system through time. Herein, we introduce the term “locomotor module” to identify anatomical subregions of the musculoskeletal system that are highly integrated and act as functional units during locomotion. The first tetrapods, which employed lateral undulations of the entire body and appendages, had one large locomotor module. Basal dinosaurs and theropods were bipedal and possessed a smaller locomotor module consisting of the hind limb and tail. Bird flight evolved as the superimposition of a second (aerial) locomotor capability onto the ancestral (terrestrial) theropod body plan. Although the origin of the wing module was the primary innovation, alterations in the terrestrial system were also significant. We propose that the primitive theropod locomotor module was functionally and anatomically subdivided into separate pelvic and caudal locomotor modules. This decoupling freed the tail to attain a new and intimate affiliation with the forelimb during flight, a configuration unique to birds. Thus, the evolution of flight can be viewed as the origin and novel association of locomotor modules. Differential elaboration of these modules in various lineages has produced the diverse locomotor abilities of modern birds.     

四川自贡中侏罗世峨眉龙—新种

记述了产自四川自贡大山铺中侏罗世峨眉龙属一新种——焦氏峨眉龙(Omeisaurus jiaoisp.nov.)。新种为大型蜥脚类恐龙,以如下特征区别于其他蜥脚类:前、中部背椎为典型的后凹型,后部背椎为双平型,背椎神经棘呈高大棒状,不分叉;尾椎为弱双凹型,第一尾椎不具扇形尾肋;第一脉弧短小且与第一尾椎关联;锁骨长大;肱骨和股骨细长而圆实;肱骨与股骨长度之比为0.83,尺骨与肱骨长度之比为0.72,胫骨与股骨长度之比为0.63。     

四川盆地一新蛇颈龙

本文记述了蛇颈龙一新属种——杨氏壁山上龙(Bishanopliosaurus youngi gen.et sp.nov.)。它的形态特征与欧洲里阿斯[Lias]期的 Rhomaleosaurus 的相似。因此可以推测产壁山上龙的地层——自流井组的东岳庙段的时代为早侏罗世。     

蜥脚类骨质尾锤之发现

自贡大山铺恐龙化石坑中出土的李氏蜀龙(Shunosaurus lii)和天府峨眉龙(Omeisaurustianfuensis)均发现由尾端脊椎愈合膨大而成之纺锤形骨质尾锤。这一特征在蜥脚类中尚属首次记述。骨质尾锤是一种适应陆地生活之特殊构造。它作为一种防御武器,主要起防卫身体的作用。它的存在说明R.T.Bakker提出的蜥脚类是一类营陆地生活的恐龙的观点是正确的。     

Variation of the number of proximal caudal vertebrae with tail reduction in Old World monkeys

Tail length in primates can vary greatly between species or even between local conspecific populations, and the tail is markedly reduced in several lineages. In Old World monkeys, tail length is considered as an important feature reflecting their phylogeny and adaptations. The number of caudal vertebrae is one of the important factors which determine tail length, and it is known that this number varies with tail length. Caudal vertebrae can be divided into two types (proximal and distal), and tail mobility and function are considered to be different in these two regions. Though the number of vertebrae in each region is important for understanding tail length evolution in Old World monkeys, there have been few attempts to investigate this matter. This study focused only on the proximal caudal vertebrae, which are more easily preserved than the distal ones, and tested if there is variation in their number with tail length or phylogenic differences. As a result, two important findings were obtained: (1) the variation of the number of proximal caudal vertebrae was different among the phylogenic groups, and (2) especially in Papionini, there was a great variation in the number of proximal caudal vertebrae, and it correlated strongly with relative tail length [RTL = (tail length/head and body length (sitting height)) × 100 %]. I speculate that these variations in the number of proximal caudal vertebrae were possibly caused by a change of the embryonic developmental mechanism of tail morphogenesis, a common mechanism of morphological evolution. To clarify the mechanisms and evolutionary trends of the variation in the proximal caudal vertebrae, not only morphological approaches but also developmental biological approaches will be necessary in the future.     

A New Species of Pengornithidae (Aves: Enantiornithes) from the Lower Cretaceous of China Suggests a Specialized Scansorial Habitat Previously Unknown in Early Birds

We describe a new enantiornithine bird, Parapengornis eurycaudatus gen. et sp. nov. from the Lower Cretaceous Jiufotang Formation of Liaoning, China. Although morphologically similar to previously described pengornithids Pengornis houi, Pengornis IVPP V18632, and Eopengornis martini, morphological differences indicate it represents a new taxon of the Pengornithidae. Based on new information from this specimen we reassign IVPP V18632 to Parapengornis sp. The well preserved pygostyle of the new specimen elucidates the morphology of this element for the clade, which is unique in pengornithids among Mesozoic birds. Similarities with modern scansores such as woodpeckers may indicate a specialized vertical climbing and clinging behavior that has not previously been inferred for early birds. The new specimen preserves a pair of fully pennaceous rachis-dominated feathers like those in the holotype of Eopengornis martini; together with the unique morphology of the pygostyle, this discovery lends evidence to early hypotheses that rachis-dominated feathers may have had a functional significance. This discovery adds to the diversity of ecological niches occupied by enantiornithines and if correct reveals are remarkable amount of locomotive differentiation among Enantiornithes.     

Evolution of Sexual Dimorphism in the Number of Tail Vertebrae in Salamanders: Comparing Multiple Hypotheses

The evolution of sexual dimorphism is an important topic of evolutionary biology, but few studies have investigated the determinants of sexual dimorphism over broad phylogenetic scales. The number of vertebrae is a discrete character influencing multiple traits of individuals, and is particularly suitable to analyze processes determining morphological variation. We evaluated the support of multiple hypotheses concerning evolutionary processes that may cause sexual dimorphism in the number of caudal vertebrae in Urodela (tailed amphibians). We obtained counts of caudal vertebrae from >2,000 individuals representing 27 species of salamanders and newts from Europe and the Near East, and integrated these data with a molecular phylogeny and multiple information on species natural history. Per each species, we estimated sexual dimorphism in caudal vertebrae number. We then used phylogenetic least squares to relate this sexual dimorphism to natural history features (courtship complexity, body size dimorphism, sexual ornamentation, aquatic phenology) representing alternative hypotheses on processes that may explain sexual dimorphism. In 18 % of species, males had significantly more caudal vertebrae than females, while in no species did females have significantly more caudal vertebrae. Dimorphism was highest in species where males have more complex courtship behaviours, while the support of other candidate mechanisms was weak. In many species, males use the tail during courtship displays, and sexual selection probably favours tails with more vertebrae. Dimorphism for the number of tail vertebrae was unrelated to other forms of dimorphism, such as sexual ornamentation or body size differences. Multiple sexually dimorphic features may evolve independently because of the interplay between sexual selection, fecundity and natural selection.     

Variability of tail length in hybrids of the Japanese macaque (Macaca fuscata) and the Taiwanese macaque (Macaca cyclopis)

In primates, tail length is subject to wide variation, and the tail may even be absent. Tail length varies greatly between each species group of the genus Macaca, which is explained by climatic factors and/or phylogeographic history. Here, tail length variability was studied in hybrids of the Japanese (M. fuscata) and Taiwanese (Macaca cyclopis) macaque, with various degrees of hybridization being evaluated through autosomal allele typing. Relative tail length (percent of crown–rump length) correlated well with the number of caudal vertebrae. Length profiles of caudal vertebrae of hybrids and parent species revealed a common pattern: the length of several proximal-most vertebrae do not differ greatly; then from the third or fourth vertebra, the length rapidly increases and peaks at around the fifth to seventh vertebra; then the length plateaus for several vertebrae and finally shows a gentle decrease. As the number of caudal vertebrae and relative tail length increase, peak vertebral length and lengths of proximal vertebrae also increase, except that of the first vertebra, which only shows a slight increase. Peak vertebral length and the number of caudal vertebrae explained 92?% of the variance in the relative tail length of hybrids. Relative tail length correlated considerably well with the degree of hybridization, with no significant deviation from the regression line being observed. Thus, neither significant heterosis nor hybrid depression occurred.     

内蒙古上白垩统二连组一长颈的镰刀龙类(英文)

镰刀龙类 (又称“懒龙”)是一类奇特的植食性兽脚类恐龙 ,化石记录主要局限于亚洲白垩纪地层中。由于镰刀龙类极其特化的形态和化石材料的局限性 ,这类恐龙的系统位置存在较多的争议。最近的发现 (RussellandDong ,1 994;Xuetal.,1 999)表明这类恐龙属于虚骨龙类 ,但其更为具体的系统位置依然存在争议 (Sues ,1 997;MakovickyandSues,1 998;Xuetal.,1 999;Sereno,1 999)。新发现于内蒙古苏尼特左旗赛罕高毕上白垩统二连组的镰刀龙类化石材料代表这类恐龙的一个新属种。杨氏内蒙古龙 (Neimongosaurusyangigen .etsp .nov .)的正型标本为一较为完整的骨架 ,是已知镰刀龙类当中第一件在同一个体中保存了大多数脊椎和几乎所有肢骨的标本。依据以下特征将内蒙古龙归入镰刀龙超科 :U形的下颌联合部、齿骨前端向下弯曲、齿骨前部没有牙齿、牙齿有一个收缩的基部、近圆形的齿根和叶形的齿冠、前部颈椎的神经脊低矮而轴向较长、后部颈椎背视呈X形、肱骨近端角状、肱骨有后转子、肱骨的尺骨髁和挠骨髁位于前部并为一狭窄槽分开、肠骨的耻骨柄细长而坐骨柄短以及跖部短。内蒙古龙的以下特征区别于其他镰刀龙类 :前部尾椎的横突下部有一圆形的窝 ,桡骨二头肌结节非常发育 ,后足趾节近端跟部非常发育 ,胫骨的腓骨嵴长     

Parallel evolution of theropod dinosaurs and birds

The hypothesis of the direct origin of birds from theropod dinosaurs has recently become widespread. Direct sisterly relationships between theropods and birds were assumed in the basis of random and formal synapomorphies, such as the number of caudal vertebrae, relative length of the humerus, and flattening of the dorsal margin of the pubis. In essence, this hypothesis is supported by the characters of theropods and birds, such as the presence of feathering, furcula, uncinate processes of ribs, pygostyle, double-condyled dorsal joint of the quadrate, and posteriorly turned pubis, which are recognized as homologies. Until recently, these characters have been regarded as avian apomorphies; however, they are presently known in various coelurosaurian groups. At the same time, they occur in various combinations in the Dromaeosauridae, Troodontidae, Oviraptoridae, Therizinosauridae, and Tyrannosauridae. None of the theropod groups possesses the entire set of these characters. This suggests that theropods and birds acquired them in parallel. Theropod dinosaurs and Sauriurae (Archaeornithes and Enantiornithes) show a number of important system synapomorphies, which indicate that they are closely related. Ornithurine birds lack such synapomorphies; however, their monophyly is supported by a large number of diagnostic characters. The hypothesis of independent origin of Sauriurae and Ornithurae is substantiated; the former are considered to have evolved from theropods in the Jurassic, while the latter deviated from a basal archosauromorph group in the Late Triassic. The hypothesis that birds existed in the Early Mesozoic is supported by the findings of small avian footprints in the Upper Triassic and Lower Jurassic of different continents.     

Ventral and sub-caudal scale counts are associated with macrohabitat use and tail specialization in viperid snakes

Viperids are a species rich clade of snakes that vary greatly in both morphology and ecology. Many species in the family express tail specializations used for defensive warnings, prey lures, and stability during locomotion and striking. To examine the relationships among ecology, behavior, and vertebral number in the family Viperidae, morphological data (maximum total length and the number of pre-cloacal and caudal vertebrae), macrohabitat use, and tail specialization for 157 viperids were gleaned from published sources. A composite tree topology was constructed from multiple published viperid phylogenies for independent contrasts analysis. The number of vertebrae was strongly correlated with the total length of the snake. Results of both non-phylogenetic and phylogenetically corrected analysis showed that macrohabitat use did not strongly influence total snake length. However, the number of vertebrae per unit length did vary among species according to macrohabitat. Specifically, vertebral density increased with increasing arboreality. Overall, viperids showed a positive correlation between the number of caudal and pre-cloacal vertebrae, but separately rattlesnakes had a significant negative correlation. Species with prehensile tails and those that caudal lure had the most caudal vertebrae. The increased caudal segments of prehensile and luring tails likely improve performance when grasping small vegetation for support or imitating invertebrate prey. These results illustrate that vertebral number is a primary characteristic involved in the diversification of viper species and ecology.     

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。     

贵州疣螈骨骼系统的研究

本文运用硬骨-软骨双染色技术对贵州疣螈（Tylototriton kweichowensis）的骨骼系统做了较全面的观察研究，包括头骨、脊柱和前后肢带骨，并对各骨块的形状、位置以及与邻近骨块的关系都作了详细的描述。最后，将贵州疣螈与东方蝾螈、有斑肥螈的骨骼系统进行了比较，为揭示蝾螈科属间演化关系提供一定的依据。     

Vertebral numbers in male and female snakes: the roles of natural,sexual and fecundity selection

The relative numbers of trunk (body) and caudal (tail) vertebrae in snakes might be influenced by at least four processes: (1) natural selection for crawling speed, (2) fecundity selection for larger trunk size in females, (3) sexual selection for longer bodies or tails in males and/or (4) developmental constraints (if an increase in the number of body vertebrae requires a decrease in the number of tail vertebrae, or vice versa). These four hypotheses generate different predictions about the relationship between sex differences in the numbers of body vertebrae vs. tail vertebrae. I collated published data to test these predictions, both with raw data and using phylogenetically independent contrasts. Some snake lineages show a negative correlation between the magnitude of sex disparities in trunk vs. caudal vertebrae whereas other lineages show the reverse pattern, or no correlation. Thus, different selective pressures seem to have been important in different lineages. Vertebral numbers in snakes may offer a useful model system in which to explore the conflicts between natural, fecundity and sexual selection.     

A nearly complete skeleton of an early juvenile diplodocid (Dinosauria: Sauropoda) from the Lower Morrison Formation (Late Jurassic) of north central Wyoming and its implications for early ontogeny and pneumaticity in sauropods

A nearly complete skeleton of a juvenile sauropod from the Lower Morrison Formation (Late Jurassic, Kimmeridgian) of the Howe Ranch in Bighorn County, Wyoming is described. The specimen consists of articulated mid-cervical to mid-caudal vertebrae and most appendicular bones, but cranial and mandibular elements are missing. The shoulder height is approximately 67 cm, and the total body length is estimated to be less than 200 cm. Besides the body size, the following morphological features indicate that this specimen is an early juvenile; (1) unfused centra and neural arches in presacral, sacral and first to ninth caudal vertebrae, (2) unfused coracoid and scapula, (3) open coracoid foramen, and (4) relatively smooth articular surfaces on the limb, wrist, and ankle bones. A large scapula, short neck and tail and elongate forelimb bones relative to overall body size demonstrate relative growth. A thin-section of the mid-shaft of a femur shows a lack of annual growth lines, indicating an early juvenile individual possibly younger than a few years old. Pneumatic structures in the vertebral column of the specimen SMA 0009 show that pneumatisation of the postcranial skeleton had already started in this individual, giving new insights in the early ontogenetic development of vertebral pneumaticity in sauropods.

The specimen exhibits a number of diplodocid features (e.g., very elongate slender scapular blade with a gradually dorsoventrally expanded distal end, a total of nine dorsal vertebrae, presence of the posterior centroparapophyseal lamina in the posterior dorsal vertebrae). Although a few diplodocid taxa, Diplodocus, cf. Apatosaurus, and cf. Barosaurus, are known from several fossil sites near the Howe Ranch, identification of this specimen, even at a generic level, is difficult due to a large degree of ontogenetic variation.     

辽西下白垩统九佛堂组尾羽龙类—新属(英文)

记述了一件采自辽西热河群九佛堂组一新的窃蛋龙类:义县似尾羽龙(Similicaudipteryx yixianensis gen.et sp.nov.),并依据其和尾羽龙属的相似特征,及其匕首状的尾综骨,肠骨、踱骨和脚趾的形态等将其归入窃蛋龙类的尾羽龙科,但其所具有的一些特征也不同于该科已建立的尾羽龙属,如尾综骨的形态和较大的个体等。该化石具有许多典型的窃蛋龙类的特征,如短尾、较短的前肢等,有别于其他窃蛋龙类的特征还包括耻骨和肠骨的长度比为1.46,背椎上发育2个大而深的椎体下突、背椎侧部具孔等。义县似尾羽龙是又一类具有真正尾综骨的恐龙,表明尾综骨这一曾经被认为是鸟类特有的结构可能是在恐龙中独自演化的。它具有一些进步的特征如具尾综骨等,但同时也具有一些原始的特征如5个愈合的荐椎,耻骨联合长,表明在窃蛋龙类中存在着特征的镶嵌进化现象。似尾羽龙与驰龙类、其他窃蛋龙类等恐龙一样,脚趾并没有对握,已有的证据说明完全的对握目前还只是出现于鸟类中。根据其短尾、中部收缩的趾骨和灵巧的身体等都表明它是一类适于快速奔跑的动物。目前为止,已知的尾羽龙类化石均发现于北票四合屯地区的义县组下部尖山沟层(段),距今约125Ma。新标本发现于义县西二虎桥地点,属于九佛堂组,距今约120Ma,这也是迄今为止在九佛堂组发现的惟一一件尾羽龙类化石,对研究早白垩世窃蛋龙类的演化和热河生物群恐龙组合的特征具有重要的意义。