中国鲤形目鱼类的脊椎骨数及其生态适应性

为探讨中国鲤形目鱼类脊椎骨数及其与系统发育和生态习性的相关性,采用X光透视照相法对鲤科(Cyprinidae)、鳅科(Cobitidae)、爬鳅科(Balitoridae)鱼类共157种的脊椎骨数、肋骨数和尾椎数进行了比较分析。结果显示,中国鲤形目鱼类的脊椎骨数30~52枚,均值39.5±4.4;肋骨数8~28对,均值15.3±3.1;尾椎数14~34枚,均值21.1±2.9。依据脊椎骨数、肋骨数、尾椎数对鲤科各亚科进行聚类,显示鲤科12个亚科可归为2大类:Ⅰ类包括雅罗鱼亚科、鲌亚科、鲴亚科、裂腹鱼亚科4亚科;Ⅱ类包括鲢亚科、亚科、亚科、鲃亚科、野鲮亚科、鳅亚科、鲤亚科、亚科8亚科。对鲤科89种所作的单因素方差分析显示,肉食性鱼类的肋骨数与脊椎骨数的比值显著小于植食性鱼类(P0.05),而脊椎骨数、尾椎数则显著多于植食性鱼类(P0.05)。中上层鱼类的脊椎骨数、尾椎数显著多于下层鱼类(P0.05)。极小型鱼类的脊椎骨数、肋骨数以及肋骨数与脊椎骨数之比显著少于或小于大型鱼类(P0.05)。表明鲤科鱼类的脊椎骨数与其生态习性及体型具有明显的相关性。     

用mtDNA 12S rRNA序列变异检验鲤形目鱼类系统发育关系

通过对鲤形目鱼类5个科的代表类群的完整线粒体12S rRNA进行测序和分析,以检验目前的形态学假说。经序列比对后,有1000个位点,其中467个位点在茎区,533个在环区;有395个位点为变化位点,其中267个为系统发育信息位点。采用邻接法和最大简约法进行了系统发育分析,其结果支持鲤科鱼类成为一个单系群,非鲤科的鲤形目鱼类成为另一个单系群的观点,这与Siebert提出的假说相一致。鲤科鱼类包含3个主要的分支,即鱼丹系、鲤系和雅罗鱼系;但在非鲤科鲤形目鱼类中,其关系不能得到很好的解决,其中鳅科是复系,平鳍鳅科、条鳅亚科和花鳅亚科可能有较近的亲缘关系。     

基于细胞色素b基因序列的鲤科鱼类系统发育研究(鱼纲: 鲤形目)

鲤科是鱼类最大的科, 在中国淡水鱼类组成中鲤科鱼类的成分占一半以上. 鲤科鱼类的演化过程代表了东亚淡水鱼类的整体演化过程. 为探讨东亚鲤科鱼类系统发育关系, 共分析了包括18种新测序列在内的54种鲤科鱼类细胞色素b基因的全序列. 分析的物种涵盖了鲤科鱼类的12个亚科并对问题较多的亚科(Danioninae)和雅罗鱼亚科(Leuciscinae)进行了广泛的采样. 系统发育树的建立使用了多种方法, 包括邻接法、最大简约法和最大似然法. 亚口鱼科(Catostomidae)的胭脂鱼(Myxocyprinus asiaticus)被指定为外类群, 3种方法所得结果非常相似, 并支持一些亚科的单系性, 如鲌亚科(Cultrinae)、鯝亚科(Xenocyprinae)、鮈亚科(Gobioninae)(包括鳅鮀亚科Gobiobotinae). 从分子水平上证明亚科不是一个单系类群, 雅罗鱼亚科被分为两个互不相关的类群; 东亚雅罗鱼群与鲌亚科和鯝亚科形成一单系类群; 而欧洲、西伯利亚、北美的雅罗鱼形成另一个单系群. Howes所定义的鲤亚科在NJ树和ML树中得到有力的支持并位于系统树的基部. 高体鳑鲏(Rhodeus sericeus)代表的鳑鲏亚科的系统位置没有得到解决. 显示鲤科鱼类可分为3个大的类群并与其3个进化阶段相对应: 老第三纪类群包含亚科的东南亚种类和Howes的鲤亚科; 北方冷水性类群包含北方雅罗鱼、鮈亚科和鳑鲏; 东亚类群包含鲌亚科、鯝亚科、雅罗鱼东亚种类和亚科东亚种类. 老第三纪类群代表鲤科鱼类的原始阶段, 北方冷水性类群对应于鲤科鱼类对全球变冷的反应, 而东亚类群则显示青藏高原急剧隆升后, 在东亚季风气候条件下鲤科鱼类的大量分化.     

基于S6K1基因部分序列的鲤科鱼类系统发育关系及内含子1 Indel位点分析

鲤科鱼类在东亚分布广泛且数量丰富, 在物种演化上具有重要的系统发育地位. 本研究基于S6K1基因5′端功能调控序列, 通过PCR扩增、克隆和测序, 共获得30种鱼类S6K1 前端部分DNA序列(外显子1, 外显子2及内含子1), 对序列变异进行分析, 并采用邻接法(NJ)、最大简约法(MP)、最大似然法(ML)和贝叶斯法(Bayesian)重建鲤科鱼类系统发育关系. 亚口鱼科的胭脂鱼(Myxocyprinus asiaticus)作为外类群, 通过4种方法所得系统发育分支图大致相同, 均以较高的节点支持率支持雅罗鱼系和鲃系的划分, 雅罗鱼系包括雅罗鱼亚科东亚类群(East Asian group in Leuciscinae)、鲢亚科(Hypophthalmichthyinae)、鲴亚科(Xenocyprinae)、鲌亚科(Cultrinae)、相似文献   

沙鳅亚科鱼类线粒体DNA控制区结构分析及系统发育关系的研究

对沙鳅亚科鱼类3属14个代表种的线粒体DNA控制区序列的结构进行了分析。通过与鲤形目鱼类的控制区序列进行比较,将沙鳅亚科鱼类的控制区分为终止序列区、中央保守区和保守序列区三个区域。同时识别了沙鳅亚科中一系列保守序列,并给出了它们的一般形式。以胭脂鱼为外类群,对比条鳅亚科、花鳅亚科、以及平鳍鳅科的代表性种类,采用NJ、MP和ML法构建沙鳅亚科的分子系统树。分子系统发育分析表明,沙鳅亚科为一单系,包括3个属:沙鳅属、副沙鳅属和薄鳅属,各属均构成单系。根据分子系统学、形态学的结果及地理分布推断,沙鳅亚科中沙鳅属可能为最为原始的属,副沙鳅属其次,而薄鳅属最特化。       

柴达木盆地渐新世的鲤科鱼类化石

记述了首次发现于柴达木盆地早渐新世晚期至晚渐新世早期(距今27～29 Ma)的鲤科鱼类化石。材料包括咽骨、咽齿、匙骨、腹鳍骨及一些零散的鳍条。咽骨及咽齿化石与原始鲃亚科鱼类及裂腹鱼亚科裂腹鱼属鱼类的相似;腹鳍骨化石与原始鲃亚科鱼类的更相似。鲃亚科鱼类现今分布于北纬35°以南的亚洲、欧洲南部及非洲北部;裂腹鱼属鱼类分布局限于青藏高原东、南、西面的边缘区域,在柴达木盆地没有分布。柴达木盆地水系现生鱼类仅见适于高寒环境的高度特化等级的裂腹鱼亚科鱼类及鳅科高原鳅属鱼类,鱼类组成与早渐新世晚期至晚渐新世早期的不尽相同。     

中国雅罗鱼亚科的系统发育

关于雅罗鱼亚科Leuciscinae的系统发育,只有个别作者对其一、二个属稍有讨论。Howes(1978)将(鱼宗)鱼属Luciobrama等从雅罗鱼亚科分出,建立了赤梢鱼群(aspine group);陈湘粦等(1984)将须(鱼岁)属Tinca从雅罗鱼亚科分出,归入钯系,并建立须(鱼岁)亚科Tincinae,这一观点与Regan(1911)“须(鱼岁)与鲃关系较近”的看法相似。 本文全面考察了分布于中国的雅罗鱼亚科鱼类的骨骼学特征,对本亚科的系统发育、起源等进行了探讨。研究标本名录见附录1。     

大鳞裂峡鲃的年轮特征

大鳞裂峡鲃(Hampala macrolepidota)隶属于鲤形目(Cypriniformes)、鲤科(Cyprinidae)、鲃亚科(Barbinae)、裂峡鲃属(Hampala),在我国属濒危鱼类[1]。国外分布于泰国、印度尼西亚等,国内仅分布于澜沧江下游云南西双版纳自治州勐腊县境内支流[2]。     

浅议《中国动物志：硬骨鱼纲.鲤形目》(中卷)

《中国动物志:硬骨鱼纲鲤形目》(中卷),陈宜瑜等编著,科学出版社.1998年出版。本卷包括鲤科的、雅罗鱼、、鲴、鲢、、鳅及8亚科,记述达260种和20个亚种;待下卷鲤、、裂腹鱼及野鲮4亚科出版后将达532种,这比我们的师辈王以康编的《鱼类分类学》,1958、张春霖写的《中国系统鲤类志》,1959种数都多。现把个人一些粗浅的不同看法敬书于后,谨供本卷著者和对鱼类学有兴趣的师友们参考评论和指正。1　鱼类分布方面自然地理分布是鱼类分类学的重要内容之一。写各属种的分布时,如有人为的(如引种移殖的)分布区,常予写明,以免混淆。如日本…     

梵净山及邻近地区鱼类资源的现状

为了了解梵净山及邻近地区鱼类资源的现状及变化趋势,2001–2003年对这一地区乌江水系和沅江水系鱼类进行了系统的调查。本调查设立了19个调查点,共采集鱼类标本3614号,经鉴定为85种(亚种),分属5目14科61属。除胡子鲇(Clariasfuscus)为外来种外,其他均为土著鱼类。海南拟餐(PseudohemiculterHainanensis)、峨眉后平鳅(Metahomalopteraomeiensisomeiensis)和福建纹胸鱼兆(Glyptothoraxfukiensisfukiensis)为本地区土著鱼类新记录种。鱼类组成以鲤形目鲤科为主体,并以适应急流生活的鱼类占多数,小型野杂鱼类所占比例较高。乌江水系和沅江水系鱼类物种组成存在差异,分别属于长江上游鱼类区系成分和长江中下游区系成分。结果还表明这一地区土著鱼类物种数呈下降趋势,其中鲤形目鱼类物种数减少较多。导致该地区土著鱼类物种数下降的原因是酷渔滥捕和水域生境的破坏。     

Microanatomical diversity of amniote ribs: an exploratory quantitative study

Bone microanatomical diversity in extant and extinct tetrapods has been studied extensively, using increasingly sophisticated quantitative methods to assess its ecological, biomechanical and phylogenetic significance. Most studies have been conducted on the appendicular skeleton, and a strong relationship was found between limb bone microanatomy and habitat preferences. Few comparative studies have focused on the microanatomy of the axial skeleton and its ecological signal. In the present study, we propose the first exploratory study of the microanatomical diversity of amniote ribs. Our comparative sample comprises 155 species of extant amniotes and encompasses the taxonomic, ecological, and body size diversity of this group. We standardized our sampling location to the midshaft of mid‐dorsal ribs. Transverse sections were obtained from classical petrographic methods, as well as by X‐ray microtomography. Most of the microanatomical and size characters of the ribs display a phylogenetic signal, which is an expected result and is also observed in amniote limb bones and vertebrae. We found a significant relationship between rib cortical thickness, global compactness, and lifestyle. As for the vertebrae, the development of the spongiosa in the medullary region appears to be strongly correlated with size. Even though an ecological signal was found in the inner structure of the ribs, additional work is needed to document the intra‐individual variability of the rib microanatomy along the rib cage and within a single element.     

Phylogenetic relationships of Cyprinidae (Teleostei: Cypriniformes) inferred from the partial <Emphasis Type="Italic">S6K1</Emphasis> gene sequences and implication of indel sites in intron 1

The family Cyprinidae is widely distributed in East Asia, and has the important phylogenetic significance in the fish evolution. In this study, the 5′ end partial sequences (containing exon 1, exon 2 and indel 1) of S6K1 gene were obtained from 30 representative species in Cyprinidae and outgroup using PCR amplification and sequencing. The phylogenetic relationships of Cyprinidae were reconstructed with neighbor joining (NJ), maximum parsimony (MP), maximum likelihood (ML), and Bayesian methods. Myxocyprinus asiaticus (Catostomidae) was assigned to the outgroup taxon. Similar phylogenetic relationships within the family Cyprinidae were achieved with the four analyses. Leuciscini and Barbini were monophyletic lineages respectively with the high nodal supports. Leuciscini comprises Hypophthalmichthyinae, Xenocyprinae, Cultrinae, Gobioninae, Acheilognathinae and East Asian species of Leuciscinae and Danioninae. Monophyly of East Asian clade was supported with high nodal support. Barbini comprises Schizothoracinae, Barbinae, Cyprininae and Labeoninae. The monophyletic lineage consisting of Danio rerio, D. myersi, and Rasbora trilineata was basal in the tree. In addition, the large fragment indels in intron 1 were analyzed to improve the understanding of Cyprinidae relationships. The results showed that the large fragment indels were correlated with the relations among species. Some conserved regions in intron 1 were thought to be involved in the functional regulation. However, no correlation was found between sequence variations and species characteristic size.     

Phylogenetic relationships of Cyprinidae (Teleostei: Cypriniformes) inferred from the partial S6K1 gene sequences and implication of indel sites in intron 1

The family Cyprinidae is widely distributed in East Asia, and has the important phylogenetic signifi- cance in the fish evolution. In this study, the 5′ end partial sequences (containing exon 1, exon 2 and indel 1) of S6K1 gene were obtained from 30 representative species in Cyprinidae and outgroup using PCR amplification and sequencing. The phylogenetic relationships of Cyprinidae were reconstructed with neighbor joining (NJ), maximum parsimony (MP), maximum likelihood (ML), and Bayesian meth- ods. Myxocyprinus asiaticus (Catostomidae) was assigned to the outgroup taxon. Similar phylogenetic relationships within the family Cyprinidae were achieved with the four analyses. Leuciscini and Barbini were monophyletic lineages respectively with the high nodal supports. Leuciscini comprises Hy- pophthalmichthyinae, Xenocyprinae, Cultrinae, Gobioninae, Acheilognathinae and East Asian species of Leuciscinae and Danioninae. Monophyly of East Asian clade was supported with high nodal support. Barbini comprises Schizothoracinae, Barbinae, Cyprininae and Labeoninae. The monophyletic lineage consisting of Danio rerio, D. myersi, and Rasbora trilineata was basal in the tree. In addition, the large fragment indels in intron 1 were analyzed to improve the understanding of Cyprinidae relationships. The results showed that the large fragment indels were correlated with the relations among species. Some conserved regions in intron 1 were thought to be involved in the functional regulation. However, no correlation was found between sequence variations and species characteristic size.     

Evolution of the intermuscular bones in the Cyprinidae (Pisces) from a phylogenetic perspective

Intermuscular bones (IBs) are widely present in morphologically generalized teleost fishes and are commonly found in the Cyprinidae. Intermuscular bones are small, hard spicules of bone that are formed by ossification in the myosepta between neighboring myomeres. Why fish have IBs, and whether there is any evolutionary pattern to their occurrence, has been poorly understood. However, the presence of IBs does substantially affect the meat quality and commercial values of many cyprinid fishes in aquaculture. In this study, we sampled 592 individuals of cyprinid fishes to systematically investigate the evolution of IBs from a phylogenetic point of view. We found that the total number of IBs in the Cyprinidae ranged from 73 to 169, and we clarified that only two categories of IBs (epineural and epipleural) were present in all examined cyprinids. Most of the IBs were distributed in the posterior region of the fish, which might be an optimal target for selecting fewer IB strains in aquaculture. There was a positive correlation between IBs and the number of vertebrae, thus making it possible to predict the approximate number of IBs by counting the number of vertebrae. Although the IBs displayed some correlation with phylogenetic relationships in some lineages and to ecological factors such as diet (especially carnivore), in an overall view the variations of IBs in cyprinids were extremely diverse. The number and patterns of IBs in these fishes may reflect their phylogenetic history, but have been shaped by multiple environment factors. In this study, we also confirmed that X‐ray photography remains an optimal and reliable method for the study of IBs.     

Molecular phylogenetics of the family Cyprinidae (Actinopterygii: Cypriniformes) as evidenced by sequence variation in the first intron of S7 ribosomal protein-coding gene: further evidence from a nuclear gene of the systematic chaos in the family

The family Cyprinidae is the largest freshwater fish group in the world, including over 200 genera and 2100 species. The phylogenetic relationships of major clades within this family are simply poorly understood, largely because of the overwhelming diversity of the group; however, several investigators have advanced different hypotheses of relationships that pre- and post-date the use of shared-derived characters as advocated through phylogenetic systematics. As expected, most previous investigations used morphological characters. Recently, mitochondrial DNA (mtDNA) sequences and combined morphological and mtDNA investigations have been used to explore and advance our understanding of species relationships and test monophyletic groupings. Limitations of these studies include limited taxon sampling and a strict reliance upon maternally inherited mtDNA variation. The present study is the first endeavor to recover the phylogenetic relationships of the 12 previously recognized monophyletic subfamilies within the Cyprinidae using newly sequenced nuclear DNA (nDNA) for over 50 species representing members of the different previously hypothesized subfamily and family groupings within the Cyprinidae and from other cypriniform families as outgroup taxa. Hypothesized phylogenetic relationships are constructed using maximum parsimony and Basyesian analyses of 1042 sites, of which 971 sites were variable and 790 were phylogenetically informative. Using other appropriate cypriniform taxa of the families Catostomidae (Myxocyprinus asiaticus), Gyrinocheilidae (Gyrinocheilus aymonieri), and Balitoridae (Nemacheilus sp. and Beaufortia kweichowensis) as outgroups, the Cyprinidae is resolved as a monophyletic group. Within the family the genera Raiamas, Barilius, Danio, and Rasbora, representing many of the tropical cyprinids, represent basal members of the family. All other species can be classified into variably supported and resolved monophyletic lineages, depending upon analysis, that are consistent with or correspond to Barbini and Leuciscini. The Barbini includes taxa traditionally aligned with the subfamily Cyprininae sensu previous morphological revisionary studies by Howes (Barbinae, Labeoninae, Cyprininae and Schizothoracinae). The Leuciscini includes six other subfamilies that are mainly divided into three separate lineages. The relationships among genera and subfamilies are discussed as well as the possible origins of major lineages.     

Axial skeletal and Hox expression domain alterations induced by retinoic acid, valproic acid, and bromoxynil during murine development

Retinoic acid (RA) alters the developmental fate of the axial skeletal anlagen. "Anteriorizations" or "posteriorizations," the assumption of characteristics of embryonic areas normally anterior or posterior to the affected tissues, are correlated with altered embryonal expression domains of Hox genes after in utero RA treatment. These "homeotic" changes have been hypothesized to result from alterations of a "Hox cod" which imparts positional identity in the axial skeleton. To investigate whether such developmental alterations were specific to RA, or were a more general response to xenobiotic exposure, CD-1 pregnant mice were exposed to RA, valproic acid (VA), or bromoxynil (Br) during organogenesis. Additionally, the expression domains of two Hox genes, Hoxa7 and Hoxa10, were examined in gestation day (GD) 12.5 embryos obtained from control, RA, VA, or Br, treated gravid dams exposed on GD 6, 7, or 8. The anterior expression boundary of Hoxa7 is at the level of the C7/T1 vertebrae and that of Hoxa10 is at L6/S1. Compound-induced changes in the incidence of skeletal variants were observed. These included supernumerary cervical ribs (CSNR) lateral to C7, 8 vertebrosternal ribs, supernumerary lumbar ribs (LSNR) lateral to L1, extra presacral vertebrae, and the induction of vertebral and/or rib malformations. RA and VA administration on GD 6 caused posteriorization in the cervico-thoracic region (CSNR) while GD 8 exposure to any of the three compounds resulted in anteriorizations in the thoraco-lumbar area (LSNR and an increase in the number of presacral vertebrae). These effects occurred across regions of the axial skeleton. Analysis of gene expression demonstrated changes in the anterior boundaries of Hoxa7 expression domains in embryos treated on GD 6 and 8 with RA. VA and Br did not induce any statistically significant alterations in Hoxa7 and none of the compounds caused alterations in Hoxa10 expression domains. The studies indicate that RA GD 6 treatment-induced Hoxa7 shifts were rostral (posteriorization) while the RA-induced GD 8 anterior expression boundary shift was caudal (anteriorization), correlating with the axial skeletal changes noted. These data suggest that xenobiotic compounds such as VA and Br may induce similar axial skeletal changes by affecting different components of the developmental processes involved in the patterning of the axial skeleton.     

The axial skeleton of the labyrinthodont Eogyrinus attheyi

An articulated length of vertebral column is used as a basis for the reconstruction of the salient features of the axial skeleton of the embolomerous anthracosaur Eogyrinus attheyi Watson, together with other material, including the holotype, in the Hancock Museum, Newcastle upon Tyne.
The trunk vertebrae are typically emboloinerous, with disoshaped notochordal pleuro-centra, firmly attached by broad facets to this neural arches, and much thinner intercentra. Regional variation is chiefly concerned with the span of the transverse processes, which diminishes posteriorly, and the associated separation of the two heads of each rib. A longitudinal series of trunk ribs, of diminishing length from the mid-trunk backwards, is reconstructed.
Eogyrinus has a normal tetrapod sacrum with one characteristic sacral rib. The first few caudal vertebrae bear ribs of unusual form, (of which four are preserved in sequence in the articulated specimen. The fifth caudal intercentrum bears the first and largest haemal arch and the pleurocentrum of the seventh caudal is distinguished by marked muscle origins presumably for the caudifemoral muscles.
The probability that Eogyrinus, like the few other embolomeres known, had an unusually long vertebral column for a labyrinthodont, is supported by an orthometric comparison using Romer's data on the American form Archeria.     

Remarkable phylogenetic resolution of the most complex clade of Cyprinidae (Teleostei: Cypriniformes): A proof of concept of homology assessment and partitioning sequence data integrated with mixed model Bayesian analyses

Despite many efforts to resolve evolutionary relationships among major clades of Cyprinidae, some nodes have been especially problematic and remain unresolved. In this study, we employ four nuclear gene fragments (3.3 kb) to infer interrelationships of the Cyprinidae.A reconstruction of the phylogenetic relationships within the family using maximum parsimony, maximum likelihood, and Bayesian analyses is presented. Among the taxa within the monophyletic Cyprinidae, Rasborinae is the basal-most lineage; Cyprinine is sister to Leuciscine. The monophyly for the subfamilies Gobioninae, Leuciscinae and Acheilognathinae were resolved with high nodal support. Although our results do not completely resolve relationships within Cyprinidae, this study presents novel and significant findings having major implications for a highly diverse and enigmatic clade of East-Asian cyprinids. Within this monophyletic group five closely-related subgroups are identified. Tinca tinca, one of the most phylogenetically enigmatic genera in the family, is strongly supported as having evolutionary affinities with this East-Asian clade; an established yet remarkable association because of the natural variation in phenotypes and generalized ecological niches occupied by these taxa.Our results clearly argue that the choice of partitioning strategies has significant impacts on the phylogenetic reconstructions, especially when multiple genes are being considered. The most highly partitioned model (partitioned by codon positions within genes) extracts the strongest phylogenetic signals and performs better than any other partitioning schemes supported by the strongest 2Δln Bayes factor. Future studies should include higher levels of taxon sampling and partitioned, model-based analyses.     

Characterization of the Skeletal Fusion with Sterility (sks) Mouse Showing Axial Skeleton Abnormalities Caused by Defects of Embryonic Skeletal Development

The development of the axial skeleton is a complex process, consisting of segmentation and differentiation of somites and ossification of the vertebrae. The autosomal recessive skeletal fusion with sterility (sks) mutation of the mouse causes skeletal malformations due to fusion of the vertebrae and ribs, but the underlying defects of vertebral formation during embryonic development have not yet been elucidated. For the present study, we examined the skeletal phenotypes of sks/sks mice during embryonic development and the chromosomal localization of the sks locus. Multiple defects of the axial skeleton, including fusion of vertebrae and fusion and bifurcation of ribs, were observed in adult and neonatal sks/sks mice. In addition, we also found polydactyly and delayed skull ossification in the sks/sks mice. Morphological defects, including disorganized vertebral arches and fusions and bifurcations of the axial skeletal elements, were observed during embryonic development at embryonic day 12.5 (E12.5) and E14.5. However, no morphological abnormality was observed at E11.5, indicating that defects of the axial skeleton are caused by malformation of the cartilaginous vertebra and ribs at an early developmental stage after formation and segmentation of the somites. By linkage analysis, the sks locus was mapped to an 8-Mb region of chromosome 4 between D4Mit331 and D4Mit199. Since no gene has already been identified as a cause of malformation of the vertebra and ribs in this region, the gene responsible for sks is suggested to be a novel gene essential for the cartilaginous vertebra and ribs.     

Differential occupation of axial morphospace

The postcranial system is composed of the axial and appendicular skeletons. The axial skeleton, which consists of serially repeating segments commonly known as vertebrae, protects and provides leverage for movement of the body. Across the vertebral column, much numerical and morphological diversity can be observed, which is associated with axial regionalization. The present article discusses this basic diversity and the early developmental mechanisms that guide vertebral formation and regionalization. An examination of vertebral numbers across the major vertebrate clades finds that actinopterygian and chondrichthyan fishes tend to increase vertebral number in the caudal region whereas Sarcopterygii increase the number of vertebrae in the precaudal region, although exceptions to each trend exist. Given the different regions of axial morphospace that are occupied by these groups, differential developmental processes control the axial patterning of actinopterygian and sarcopterygian species. It is possible that, among a variety of factors, the differential selective regimes for aquatic versus terrestrial locomotion have led to the differential use of axial morphospace in vertebrates.