【作者】 韩德民；

【导师】 周开亚；

【作者基本信息】 南京师范大学 ， 动物学， 2003， 博士

【摘要】 壁虎类蜥蜴由睑虎科(Eublepharidae)、鳞脚虎科(Pygopodidae)壁虎科(Gekkonidae)等几个类群组成，全世界现存有116属1110种(2001年统计)。早期对壁虎类系统发生关系的研究主要以形态学性状为依据，同时结合生物学特征及古生物学方面的发现。20世纪中叶以后，在研究中引入了支序分析方法，提出几种系统发生假说，观点虽不尽一致，但一般认为睑虎科最为原始，壁虎科分化较晚。20世纪80年代以来利用染色体尤其蛋白质和核酸等生物大分子证据，对该类动物的系统发生关系作进一步的验证和探讨，取得了不少成绩。本文以核基因C-mos和线粒体基因12S rRNA为分子标记，对壁虎类蜥蜴的分子系统学进行了研究。 1．壁虎类蜥蜴高阶元类群间的系统发生关系 C-mos基因作为分子标记已成功地应用于包括爬行动物在内的某些脊椎动较高阶元类群系统学的研究，我们选择这一基因分析壁虎类蜥蜴高阶元类群间的系统发生关系。从中国、印度、马来西亚、美国、墨西哥、波多黎各、南非、肯尼亚、纳米比亚、莫桑比克、基里巴斯、马达加斯加、澳大利亚、新喀里多尼亚等地共采集样本30个种，分别代表壁虎类蜥蜴已知所有各主要分楚群的30个属。测得该基因约378bp的部分序列，另从GenBank获徘壁虎类11个分类单元该基因片段的序列数据。系统发生分析的结果提示： (1)壁虎类以及睑虎科、壁虎科、鳞脚虎科、鳞脚虎亚科(Pygopodinae)和刺尾虎族(Diplodactylini)(Kluge，1987)各自都是单系的； (2)刺尾虎亚科(Diplodactylinae)和藁趾虎族(Carphodactylini)不是单系—— 的，建议将 KI呛e＊）的鳞脚虎科改订为以下三个科：鳞脚虎科 （Pygopodidae），鼓趾虎科（CarPhodactylidae）和刺尾虎科 （Dipl。dpCtylidae人 （3）g金虎科三个属之间的关系为（oleol7N（Mlo由c trlus十伽7邮ar／d）， 与此前大部分研究结果不同； k）支持壁虎类蝴蝎早期分化与东、西冈瓦纳古陆的分裂相关的假说，现存睑虎 科和东部冈瓦纳分支（刺尾虎科、鳞脚虎科和基趾虎科）的分化时间可能较 壁虎科为早。2．壁虎科晰间的系统发生关系 壁虎科是壁虎类蝴蝎中种类最多、变异最大、系统发生关系最不清楚的一个科。我们测定了壁虎科28个分类单元的核C－。s基因部分序列和线粒体 125 rRNA基因全序列，比对后的序列长依次分别为378hP和 1459hP。将两组数据合并进行系统发生分析，主要结论有： （1）根据 NJ树和 MP树的一致途径，重建的分子系统树将壁虎科样本聚力 6组： l）（肋OptmpUS＋PaCtheCtfiUS）肋OISUffe： 2）NactuS＋LygodactriuS； 3）皿istinus＋Alhacko； 4）肋／IOaC t，dUS（HeNWIOaC tNUS十伽抑）； 5）（（驯ero内ctrius＋Ptenopus）Naru俯sia）（加的有ctrius＋ CalodactriodeS）； 6）HddactriUS＋LopidodactriuS＋GekkO。 以上结果与 Kluge（1987）和 Russell（1972，1976）等基于形态学分析得到的假说有很大的差异。 （2） C－n。os和 12s rRNA合并数据的分析结果表明球趾虎与壁虎科的 Ptenopus和 Narudasia等有较近的亲缘关系．但单用C－。s基因序列分析显示它可能是壁虎 科最原始的一支，该类群的系统发生地位需要增加样本进一步研究。 ③ Tera toscfncus未在自引导支持下与任何分类单元相聚，所以沙虎亚科 （Teratoscincjnae）的系统发生地位在本研究中未得出结论。3．大壁虎的线粒体基因组 采用长片段扩增、克隆测序和PCR引物步行测序等方法，测定了大壁虎（GekkOgecko）线粒体基因组全序列，主要结果如下： ＊）序列全长为16434hP。其基因组成和顺序、基因的方向即编码链的选择、碱 n 壁虎类斯畅的分子系统学研究 基G的低含量和对碱基T的偏好以及GC和 AT偏斜与大部分脊椎动物相同。 GC含量与两栖类和爬行类最为接近，与其他各类脊索动物都有较大程度的差 异。 ② 蛋白编码基因第三编码位置表现为对碱基A的偏好，更接近两栖类以下的较 低等脊椎动物而不是羊膜动物。 （3）蛋白质标准终止密码子（TAA）只有 3个，较大部分脊椎动物为少。 O）从基因组的结构、核着酸组成、编码区和非编码区等的特征来看，壁虎类与 蛇类的差异较大，而与另外两种蛐蝎比较接近。更多还原

【Abstract】 Gekkotan lizards are composed of Eublepharidae, Pygopodidae and Gekkonidae, and current estimates of gekkotan diversity recognize approximately 1110 species in 116 genera. The morphological characters as well as biological and paleontological findings were employed for analysis of systematic relationship in early times. According to the cladistic analysis, which has been introduced since the middle period of 20 century, Eublepharidae was the most primiteve clade and Gekkonidae was differentiated latest. New evidence of chromosome, protein and nucleic acid has been employed to deduce and validate the interrelationships of the taxa since 1980s. C-mos nuclear and 12S rRNA mitochondrial DNA were employed as genetic markers to deduce (he phylogeny of gekkotan lizards in the present study1. Phylogenetic relationships among the higher taxonomic categories of gekkotan lizardsC-mos gene appears very useful for resolving phylogenetic divergences among higher level vertebrate taxa including reptiles. We selected this gene for the phylogenetic relationship analysis among the higher taxonomic categories of gekkotan lizards. Samples were selected to include representatives of all previously recognized higher order groups of gekkotans and to include wide geographic representation. Samples representing 30 genera were collected from China, India, Malaysia, the United States, Mexico, Puerto Rico, South Africa, Kenya, Namibia, Mozambique, Kiribati, Madagascar, Australia, and New Caledonia. The aligned length of 378 base pair (bp) partial sequences was available for the C-mosDNA. Sequences of an additional 11 gekkotan genera retrieved from GenBank were also used in the analysis.(1) Gekkota as well as Eublepharidae, Gekkonidae, Pygopodidae, Pygopodinae, Diplodactylini (sensu Kluge, 1987) are monophyletic;(2) Diplodactylinae and Carphodactylini are not monophyletic, and we propose to recognize three families within the Pygopodidae sensu Kluge (1987): Pygopodidae, Carphodactylidae and Diplodactylidae.(3) The position ofColeonyx as the sister group of the other genera, Holodactylus and Eublepharis is different from most of previous results;(4) Our results support the hypothesis that the early eladogenesis of the Gekkota was associated with the split of Eastern Gondwanaland from Western Gondwanaland. Divergences within living genera in the Eublepharidae .and the Eastern Gondwana lineages (Diplodactylidae, Pygopodidae and Carphodactylidae) may be older than those in the Gekkonidae.2. Phylogenetic relationships of the gekkonid lizardsThe Gekkonidae was the most heterogeneous and unwieldy of all gekkotan lizards. Samples representing 28 genera were collected , and 378 bp C-mos partial and 1459 bp 12S rRNA complete sequences were obtained. Two data sets were combined in the analysis.(1) Majority-role consensus phylogenetic trees cluster the taxa sampled into 6 clades:1) (Rhoptropus + Pachydaclylu\} Phelsumu;2) Nactus + Lygodactylus; 3 ) Christinus + Afrogecko -,4) Phyllodactylus (Hemiphyllodactylns + Gehyra)\5) ((Sphaerodactylus + Plenopits) Narudasia} (Syrtodactylus + Calodactylodes );6 ) Hemidactylus + Lepidodaclylus + Gekko, and the results have many defferences with those of Kluge (1987) and Russell (1972, 1976) based on morphological characters.(2) The result from combind data sets of C-mos and 12S rRNA gene sequences indicates Sphaerodactylus has a close relationship to Ptenopus and Narudasia, but it is in the basal position of gekkonids based on C-mos data alone. The higher order status of the sphaerodactyls will require more intensive sampling of this group.(3) As Teratoscincus dose not cluster with any other taxa with bootstrap supports inboth data sets, so the phylogenetic status of Teratoscincinae has not been resolved in the present study.3. The mitochondria! genome oftokayAnalyses of complete mitochondria! genomes provide not only information about structural arrangements which may serve as genomic evolutional marker but also sequ更多还原