中国爬行纲动物分类厘定
蔡波, 王跃招*, 陈跃英, 李家堂
中国科学院成都生物研究所, 成都 610041
* 通讯作者 Author for correspondence. E-mail: arcib@cib.ac.cn
摘要

本文对中国爬行纲动物的分类体系和物种进行了系统的评估, 规范了中文学名, 给出了《中国爬行纲校正名录》, 结果表明: 中国现存爬行纲动物3目30科132属462种, 其中鳄形目(Crocodylia)1科1属1种, 龟鳖目(Testudines)6科18属33种, 有鳞目(Squamata)蜥蜴亚目(Lacertilia)10科41属189种, 有鳞目蛇亚目(Serpentes)13科72属239种。与《中国动物志 爬行纲 第一卷(总论、龟鳖目、鳄形目)》、《中国动物志 爬行纲 第二卷(有鳞目: 蜥蜴亚目)》和《中国蛇类》相比, 目和亚目无变化; 科级水平新增5科, 变更2科; 属级水平新增23属, 合并15属, 变更6属; 种级水平新增81种, 变动2种; 未收录同物异名12种、杂交6种、中国无分布7种。形态和分子系统发育研究结果在爬行动物不同分类阶元均有一定差异, 文章对这些争议进行了讨论, 并对名录的选择做了说明。

关键词: 爬行动物; 名录; 中国; 生物多样性
A revised taxonomy for Chinese reptiles
Bo Cai, Yuezhao Wang*, Yueying Chen, Jiatang Li
Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041
Abstract

Based on taxonomic and phylogenetic studies, we presented a comprehensive reassessment of the classification systems, updated corresponding Chinese scientific names and concluded the Checklist of Chinese Reptilia. Our analysis suggested that in China there are a total of 3 orders, 30 families, 132 genera, and 462 reptile species. The order Crocodylia includes one family, one genus, and one species. The order, Testudines includes 6 families, 18 genera, and 33 species. The order Squamata includes the suborder Lacertilia and Serpentes. Lacertilia includes 10 families, 41 genera, and 189 species and Serpentes includes 13 families, 72 genera, and 239 species. Compared to Fauna Sinica (Reptilia 1): General Accounts of Reptilia,Testudoformes and Crocodiliformes, Fauna Sinica (Reptilia 2): Squamata (Lacertilia) and Snakes of China, we added 81 species, revised 2 species and excluded 12 synonymous, 6 hybrid, and 7 undistributed reptile species. At the genus level, there are 23 additional, 6 revised, 15 synonymous genera. At the family level, there are 5 additional and 2 revised families. There is no change at the order or suborder levels. Studies of morphology and molecular phylogeny on taxonomic categories of reptiles reflects several divergences. We discusssed these controversies and explained choices of the Checklist of Chinese Reptilia in this article .

Key words: Reptilia; checklist; China; biodiversity

爬行动物是脊椎动物演化史上首次真正适应陆地环境的动物类群, 中国是少数几个同时跨越古北界与东洋界两大动物区系的国家。第三纪晚期以来发生的地质事件, 使得包括中国在内的区域气候地理环境极其复杂(张荣祖, 2011), 孕育了众多的爬行动物(Huang et al., 2011; 蔡波等, 2012)。

我国爬行动物分类研究起步较晚。《中国爬行动物系统检索》是我国最早的全面而系统整理中国爬行动物的文献(四川省生物研究所两栖爬行动物研究室, 1977), 该书收录我国爬行纲动物316种, 包括鳄目(Crocodiliformes)1科2属2种, 龟鳖目(Testu- doformes) 4科14属24种, 蜥蜴目(Lacertifomes)8科34属117种, 蛇目(Serpentiformes)8科53属173种。其后, 在Herpetology of China(Zhao & Adler, 1993)中, 列举了我国爬行动物388种, 其中鳄目1种、龟鳖目34种、蜥蜴目152种、蛇目201种。而后, 1998-1999年出版的《中国动物志 爬行纲 第一卷(总论、龟鳖目、鳄形目)》(张孟闻等, 1998)、《中国动物志 爬行纲 第二卷(有鳞目: 蜥蜴亚目)》(赵尔宓等, 1998)和《中国动物志 爬行纲 第二卷(有鳞目: 蛇亚目)》(赵尔宓等, 1999)对我国爬行纲动物进行了更为系统的研究和整理, 记录鳄形目(Crocodiliformes)2科3属3种, 龟鳖目6科21属37种, 将蜥蜴目与蛇目分别变更为有鳞目(Squamata)中的蜥蜴亚目(Lacertilia)和蛇亚目(Serpentes), 其中蜥蜴亚目9科39属156种, 蛇亚目8科64属203种。此后, 《中国蛇类》(赵尔宓, 2006)、《中国贸易龟类检索图鉴》(史海涛, 2008, 2011)以及《中国龟鳖分类原色图鉴》(周婷和李丕鹏, 2013)分别对蛇亚目和龟鳖目动物进行了部分整理和总结。

随着分子生物学的发展和进步, 以爬行动物为研究对象的论文和专著日渐增多, 形态学与分子系统学研究的结合也将爬行动物系统发育关系梳理得更为科学合理。国际上爬行纲动物分类体系也发生了很大变化, 并不断出现新分类阶元的报道(Gatesy et al., 2004; Barley et al., 2010; Oaks, 2011; Pyron et al., 2013; Vitt & Caldwell, 2013; Dijk et al., 2014; Wallach et al., 2014)。而我国目前的分类体系与国际公认的分类体系间存在较大差异, 未能及时反映最新的分类成果, 这无疑将对我国的科学研究与学术交流、物种保护及执法等产生不利影响。因此, 我们借国家环境保护部中国脊椎动物红色名录爬行纲研究项目, 邀请国内相关专家对我国的爬行纲分类体系进行了厘定, 并对中文学名做了必要的规范。

1 方法

在《中国动物志 爬行纲 第一卷(总论、龟鳖目、鳄形目)》(张孟闻等, 1998)、《中国动物志 爬行纲 第二卷(有鳞目: 蜥蜴亚目)》(赵尔宓等, 1999)和《中国蛇类》(赵尔宓, 2006)的基础上, 参考了《中国贸易龟类检索图鉴》(史海涛, 2011)、《中国龟鳖分类原色图鉴》(周婷和李丕鹏, 2013)、IUCN中国爬行动物名录(2013)、中国科学院成都生物研究所两栖爬行动物标本馆标本数据库、reptile-database.org和www.herpnet.org网站, 截止到2015年4月30日, 共收集各类有效信息6, 700余条, 形成了中国爬行动物初步编目。

在整理过程中, 我们遵循新种和新记录种须有明确的分类鉴定特征、规范的模式标本描述、明确的模式标本存放地(可公开查询的标本馆、博物馆)、发表的杂志须为公开的学术刊物的原则。厘定分类体系科学性的原则为: (1)符合分类学法规; (2)大部分爬行动物分类专家认同; (3)有网评和会审过程; (4)实际考察验证。

新的分类阶元中物种中文学名的拟定原则为: (1)科学、适用; (2)历史传承; (3)符合拉丁名的原意; (4)约定俗成的习惯; (5)与形态特征吻合。

我们举行了4次网评和2次会审, 反复征求国内27名两栖爬行动物分类专家的意见, 最终确定有效种, 剔除同物异名、杂交种、误定种等无效种, 形成《中国爬行纲校正名录》(以下简称《名录》)。

2 结果

最初, 我们初步归纳、整理截至2012年以来我国爬行纲物种517种。然后, 根据第一轮评审, 确定421种有效种, 18种无效种, 78种未予评估。而后, 第二轮评审, 增加了2013年以来我国发现的新种, 共整理出3目30科131属461种(包括区域灭绝种2种), 形成《中国生物多样性红色名录· 爬行动物红色名录》(以下简称《红色名录》)。而后, 我们在《红色名录》的基础上进行最后订正增删。截至2015年4月30日, 最终确定我国爬行纲共有3目30科132属462种, 其中鳄形目1科1属1种, 龟鳖目6科18属33种, 有鳞目23科113属428种(不包括区域灭绝种, 见附录1)。

2.1 科级分类阶元调整

在《中国动物志 爬行纲 第一卷(总论、龟鳖目、鳄形目)》(张孟闻等, 1998)、《中国动物志 爬行纲 第二卷(有鳞目: 蜥蜴亚目)》(赵尔宓等, 1999)和《中国蛇类》(赵尔宓, 2006)的基础上, 基于已有研究和评估, 对科级分类阶元做了调整(附录2)。

新增5科, 即球趾虎科(Sphaerodactylidae) (Gamble et al., 2008, 2011, 2012; Pyron et al., 2013)、闪皮蛇科(Xenodermatidae)、钝头蛇科(Pareatidae)、水蛇科(Homalopsidae)和鳗形蛇科(Lamprophiidae) (Vidal et al., 2007; Wiens et al., 2008; Zaher et al., 2009; Pyron et al., 2011, 2013; Wallach et al., 2014)。

将1亚科提升为科, 即将鼍亚科(Alligatorinae)提升为鼍科(Alligatoridae)(张孟闻等, 1998; Brochu, 2003; Gatesy et al., 2004; Willis, 2009; Oaks, 2011; Erickson et al., 2012; Jaratlerdsiri et al., 2014)。变更1科, 将水龟亚科(Batagurinae)更改为地龟科(Geoemydidae)(Shaffer et al., 1997; 张孟闻等, 1998; Krenz et al., 2005; Zhang et al., 2008; Barley et al., 2010; Thomson & Shaffer, 2010; Dijk et al., 2014)。

2.2 属、种级分类阶元调整

基于已有研究和评估, 《名录》与《中国动物志 爬行纲 第一卷(总论、龟鳖目、鳄形目)》相比, 龟鳖目合并3属, 新增4种, 未收录同物异名4种、杂交种6种和无分布2种; 鳄形目未收录区域灭绝种2种; 《名录》与《中国动物志 爬行纲 第二卷(有鳞目: 蜥蜴亚目)》相比, 蜥蜴亚目新增6属, 合并3属, 变更4属, 新增40种, 变动1种, 未收录同物异名8种和国内无分布3种; 《名录》与《中国蛇类》相比, 蛇亚目新增17属, 合并9属, 变更2属, 新增37种, 变动1种, 未收录国内无分布2种(附录3)。

2.2.1 龟鳖目

合并了3属。将龟属(Chinemys)与花龟属(Ocadia)并入拟水龟属(Mauremy)(Spinks et al., 2004), 种的中文名不变; 将锯缘摄龟属(Pyxidea)并入闭壳龟属(Cuora)(Stuart & Parham, 2004; Zhang et al., 2008), 锯缘摄龟(Pyxidea mouhotii)中文名相应变更为锯缘闭壳龟(Cuora mouhotii)。

新增或提升了4种。分别是东北鳖(Pelodiscus maackii)(Fritz & Havaš , 2007; Fritz et al., 2010; 周婷和李丕鹏, 2013; Le et al., 2014)、小鳖(P. parviformis)(唐业忠, 1997; Yang P et al., 2011; 周婷和李丕鹏, 2013; Le et al., 2014)、砂鳖(P. axenaria)(周工健等, 1991; 陈合格等, 2006; Fritz et al., 2010; 周婷和李丕鹏, 2013; Le et al., 2014)和欧氏摄龟(Cyclemys oldhamii)(Kou, 1989; Fritz et al., 2008; Stuart & Fritz, 2008; 周婷和李丕鹏, 2013)。

未收录专家会评时被认为同物异名的4种。斑鼋(Pelochelys maculatus)被认为是斯氏鳖(Rafetus swinhoei)的同物异名(Le & Pritchard, 2009; 王剑等, 2010; 史海涛, 2011; 周婷和李丕鹏, 2013); 大头乌龟(Chinemy megalocephala)被认为是乌龟(Mauremy reevesii)的同物异名(Iverson et al., 1989; Barth et al., 2002, 2004; Feldman & Parham, 2004; Shen et al., 2013); 滇南摄龟(Cyclemys tiannanensis)被认为是欧氏摄龟的同物异名(Fritz et al., 2008; Stuart & Fritz, 2008; 周婷和李丕鹏, 2013); 越南三线闭壳龟(Cuora cyclornata)被认为是三线闭壳龟(C. trifasciata)的同物异名(Spinks & Shaffer, 2006), 后被证明为无效种(Spinkset al., 2009)。

未收录专家会评时认为杂交的6种。分别是: 地龟科中的费氏花龟(Ocadia philippeni)、广西拟水龟(Mauremys guangxiensis)、艾氏拟水龟(M. iversoni)、拟眼斑水龟(Sacalia pesudocellata)(张孟闻等, 1998)、腊戌拟水龟(Mauremys pritchardi)和缺颌花龟(M. glyphistoma), 被证明分别为黄喉拟水龟(M. mutica)与花龟(M. sinensis)、黄喉拟水龟与艾氏拟水龟、黄喉拟水龟与三线闭壳龟、三线闭壳龟与四眼斑水龟(Sacalia quadriocellata)、乌龟(Mauremys reevesii)与黄喉拟水龟、花龟与安南拟水龟(Mauremys annamensis)产生的杂交种(Parham & Shi, 2001; Parham et al., 2001; Buskirk et al., 2005; Stuart & Parham, 2007; 周婷和李丕鹏, 2013)。

未收录专家会评时认为国内无分布的2种。马来闭壳龟(Cuora amboinensis)也称为安布闭壳龟, Pope于1935年首次提及在中国有分布, 但无详细分布信息, 而1990年后国内陆续出现该种的记载(张孟闻等, 1998), 但都与贸易相关, 且记载地点在边境附近。因此, 本文也认为安布闭壳龟在中国无分布(史海涛, 2011; 周婷和李丕鹏, 2013), 不予收录。汤俊和宗愉于1988年根据市场标本, 首次记录我国分布有齿缘摄龟(Cyclemys dentata); 随后, Kou (1989)发布新种滇南摄龟(C. tiannanensis)(被认为是欧氏摄龟的同物异名), 同时发现有齿缘摄龟, 但未说明标本来自野外采集还是市场购买。此后, 欧氏摄龟在野外被发现尚有分布(史海涛和王剑, 个人通讯), 而齿缘摄龟无可靠野外报道, 在中国分布存疑(史海涛, 2011), 因此, 本《名录》未收录该物种。

此外, 专家会评时还对鳄形目和龟鳖目一些物种的中文名做了调整, 如: Alligator sinensis的中文名由“ 鼍” 变更为“ 扬子鳄” , Rafetus 属中文名规范为“ 斑鳖属” , Rafetus swinhoei中文名规范为“ 斑鳖” , Pelodiscus sinensis的中文名由“ 鳖” 规范为“ 中华鳖” , Chelonia mydas的中文名由“ 海龟” 规范为与体征相符并被广泛认可的“ 绿海龟” , 蠵龟属(Caretta)的物种Caretta caretta的中文名由“ 蠵龟” 规范为更易读写、被广泛认可并与体征相符的“ 红海龟” 。

2.2.2 有鳞目

2.2.2.1 蜥蜴亚目

新增、恢复或提升了6属。将原弯脚虎属(Cyrtopodion)分成隆趾虎属(Altiphylax)、中趾虎属(Mediodactylus)和裸趾虎属(Cyrtodactylus)3个属(时磊, 2010; Bauer et al., 2013), 中文名也随属名作相应改变; 蜥蜴属(Lacerta)中拆分出了胎蜥属(Zootoca), 胎生蜥蜴(Lacerta vivipara)相应改为胎蜥(Zootoca vivipara) (Sindaco & Jeremč enko, 2008; Arribas, 2009; Cornetti et al., 2015); 原树蜥属(Calotes)新拆分出了拟树蜥属(Pseudocalotes) (Hallermann & Bö hme, 2000; Mahony, 2010; Pyron et al., 2013), 中文名随属名作相应改变; 此外还增加了泛蜥属(Ablepharus)(时磊等, 2006)。

合并了3属。蝎虎属(Platyurus)被并入蜥虎属(Hemidactylus)(Carranza & Arnold, 2006; Pyron et al., 2013), 蝎虎(Platyurus platyurus)中文名相应变为蝎尾蜥虎(Hemidactylus platyurus); 地蜥属(Platyplacopus)被并入草蜥属(Takydromus), 中文名也随属名作相应改变(Arnold, 1997; Lin et al., 2002; Pyron et al., 2013); 另外, 石龙子科(Scincidae)中, Pyron等(2013)发现裂睑蜥属(Asymblepharus)与泛蜥属(Ablepharus)互不为单系, 按照国际动物命名法(International Code of Zoological Nomenclature)的优先原则, 本《名录》将裂睑蜥属物种暂时并入泛蜥属(时磊, 个人通讯), 因此阿赖山裂睑蜥更名为阿赖山泛蜥(Ablepharus alaicus)。

变更4属。原Eumeces属被拆分, 分布于东亚(包括中国)与北美的“ Eumeces” 和Neoseps合并为Plestiodon属(Griffith et al., 2000; Schmitz et al., 2004; Brandley et al., 2005, 2011, 2012; Pyron et al., 2013), 该属继续沿用中文属名“ 石龙子属” , 属下种中文名不变; 原Riopa属东南亚的一支全部归入蝘蜓属(Lygosoma)(Geissler et al., 2011, 2012; Pyron et al., 2013), 原中文种名“ 侏蜥” 不变; 原Mabuya属亚洲一支全部归入Eutropis属(Ota & Huang, 2000; Mausfeld et al., 2002; Pyron et al., 2013), 继续沿用中文属名“ 南蜥属” , 属下种中文名不变; 蛇蜥科中, 《中国动物志》中脆蛇属(Ophisaurus)变动为脆蛇蜥属(Dopasia) (Conrad et al., 2011; Nguyen et al., 2011; Pyron et al., 2013), 属下种中文名相应改变。

新增、恢复或提升了40种。分别是: 大敦弯脚虎(Cyrtopodion dadunensis)(Shi & Zhao, 2011)、瓦氏裸趾虎(Cyrtodactylus wayakonei)(Nguyen et al., 2010; Yuan & Rao, 2011)、赵氏裸趾虎(C. zhaoermii)(Shi & Zhao, 2010)、缅北蜥虎(Hemidactylus aquilonius)(McMahan & Zug, 2007; Bauer et al., 2010)、锯尾蜥虎(H. garnotii)(蒋爱伍等, 2006; 杨大同和饶定齐, 2008)、半叶趾虎(Hemiphyllodactylus typus) (吕光洋等, 1999; 向高世等, 2009)、梅氏壁虎(Gekko melli)(Ota et al., 1995; Rö sler et al., 2011; Yang et al., 2012)、海南壁虎(G. similignum)(Ota et al., 1995; Rö sler et al., 2011)、文县壁虎(G. wenxianensis)(Zhou & Wang, 2008; Nguyen et al., 2013)、鹰氏壁虎(G. adleri)(Nguyen et al., 2013)、吐鲁番沙虎(Teratoscincus roborowskii)(Macey et al., 1997, 1999; Rö sler, 2000)、越南石龙子(Plestiodon tamdaoensis)(Hikida et al., 2001)、北部湾蜓蜥(Sphenomorphus tonkinensis)(Wang et al., 2013b)、沙地泛蜥(Ablepharus deserti)(时磊等, 2006)、喀什麻蜥(Eremias buechneri)(Szczerbak & Golubev, 2003; Sindaco & Jeremč enko, 2008)、方额麻蜥(E. quadrifrons) (Zhao & Adler, 1993; Sindaco & Jeremč enko, 2008; Guo XG et al., 2011)、汶川攀蜥(Japalura zhaoermii) (高正发和侯勉, 2002; Manthey et al., 2012)、巴塘攀蜥(J. batangensis) (李操等, 2001; 巫嘉伟等, 2005; Manthey et al., 2012)、宜兰攀蜥(J. luei) (Ota et al., 1998; 吕光洋等, 1999; 向高世等, 2009)、绿背攀蜥(J. sagittifera)(Kunte & Manthey, 2009)、雾林攀蜥(J. tricarinata) (Macey et al., 2000; Kunte & Manthey, 2009)、短尾攀蜥(J. brevicauda) (Manthey et al., 2012)、玉龙攀蜥(J. yulongensis)(Manthey et al., 2012)、贵德沙蜥(Phrynocephalus putjatai) (王跃招等, 2002; Ananjeva et al., 2011; Jin & Brown, 2013)、贵南沙蜥(P. guinanensis)(Ji et al., 2009)。原Goniurosaurus lichtenfelderi被拆分为多个物种, 我国新增或提升了海南睑虎(Goniurosaurus hainanensis)(Blair et al., 2009; 张洁等, 2014)、凭祥睑虎(G. luii)(Grismer et al., 1999; 张洁等, 2014)、霸王岭睑虎(G. bawanglingensis)(Grismer et al., 2002; 张洁等, 2014)、英德睑虎(G. yingdeensis)(Wang et al., 2010; 张洁等, 2014)和荔波睑虎(G. liboensis)(Wang et al., 2013a; 张洁等, 2014), 以及未列入《红色名录》而增补进《名录》的越南睑虎(G. araneus) (Chen TB et al., 2014)和蛰龙睑虎(G. zhelongi) (Wang YY et al., 2014; 张洁等, 2014), 因此, 我们采纳张洁等(2014)的意见, 将G. lichtenfelderi原中文名由“ 睑虎” 规范为“ 里氏睑虎” 。密点麻蜥莎车亚种(Eremias multiocellata yarkandensis)、密点麻蜥科克沙尔亚种(E. m. kokshaaliensis)和密点麻蜥天山亚种(E. m. stummeri)被提升为莎车麻蜥(E. yarkandensis)、科克沙尔麻蜥(E. kokshaaliensis)和天山麻蜥(E. stummeri) (Jeremč enko & Panfilov, 1999; Ananjeva et al., 2006; Sindaco & Jeremč enko, 2008), 本名录采用该观点。杨大同和饶定齐(2008)认为孟加拉巨蜥指名亚种(Varanus bengalensis bengalensis)伊江居群应提升为种伊江巨蜥(V. irrawadicus), 专家会评时采纳该意见。乌拉尔沙蜥伊犁亚种(Phrynocephalus guttatus alpherakii)和乌拉尔沙蜥黑腹亚种(P. g. melanurus)提升为伊犁沙蜥(P. alpherakii)(Д у н а е в , 1989; Ananjeva et al., 2011; С о л о в ь е в а et al., 2011; Milto & Barabanov, 2012)和黑腹沙蜥(P. melanurus)(Д у н а е в , 1989; Barabanov & Ananjeva, 2007; Ananjeva et al., 2011; С о л о в ь е в а et al., 2011; Milto & Barabanov, 2012)。大壁虎黑疣亚种(Gekko gekko reevesii)被提升为黑疣大壁虎(G. reevesii) (杜世章等, 2002; 秦新民等, 2007a, b; Rö sler et al., 2011; 邓欢欢等, 2011), 被专家会评时采纳。原地蜥属中, 台湾地蜥(Platyplacopus kuehnei)被拆分出了鹿野草蜥(Takydromus luyeanus)和翠斑草蜥(T. viridipunctatus)(Arnold, 1997; Lin et al., 2002; Lue & Lin, 2008; 向高世等, 2009; Pyron et al., 2013), 而Takydromus kuehnei中文名依照台湾名称由“ 台湾草蜥” 改为“ 古氏草蜥” 。

种的属级分类阶元变动1种。杨大同和饶定齐(2008)认为原西藏树蜥巴坡亚种(Pseudocalotes kingdonwardi bapoensis)应归入攀蜥属(Japalura), 变更为独龙江攀蜥(Japalura bapoensis), 专家会评时采纳该建议。

未收录专家会评时被认为是同物异名的8种。台湾脆蛇(Ophisaurus formosensis)被认为是脆蛇蜥(Dopasia harti)的同物异名(Lin et al., 2003; Pyron et al., 2013); 塔里木岩蜥(Paralaudakia tarimensis)被认为是新疆岩蜥(P. stoliczkana)的同物异名(Peters, 1971; Baig et al., 2012); 红原沙蜥(Phrynocephalus hongyuanensis)被认为是青海沙蜥(P. vlangalii)的同物异名(王跃招等, 2002; Guo & Wang, 2007; Ananjeva et al., 2011; Guo XG et al., 2012); 泽当沙蜥(P. zetangensis)被认为是西藏沙蜥(P. theobaldi)的同物异名(Guo & Wang, 2007; Jin et al., 2008; Ananjeva et al., 2011; Guo XG et al., 2012); 白梢沙蜥(P. koslowi)被认为是叶城沙蜥(P. axillaris)的同物异名(Barabanov & Ananjeva, 2007; Ananjeva et al., 2011); 无斑沙蜥(P. immaculatus)是变色沙蜥(P. versicolor)的同物异名(Barabanov & Ananjeva, 2007; Ananjeva et al., 2011); 草原沙蜥(P. frontalis)是荒漠沙蜥(P. przewalskii)的同物异名(Gozdzik & Fu, 2009; Ananjeva et al., 2011); 白条沙蜥(P. albolineatus)是乌拉尔沙蜥黑腹亚种的同物异名(Д у н а е в , 1989; Barabanov & Ananjeva, 2007; Ananjeva et al., 2011; С о л о в ь е в а et al., 2011; Milto & Barabanov, 2012)。

未收录专家会评时认为国内无分布的3种。翠蜥属(Lamprolepis)的翠蜥(L. smaragdina)在台湾的唯一记录是Stejneger在1910年报道的动物标本商收集的一只幼体(向高世和林松霖, 2001), 该记录点的地理位置和生境与翠蜥分布并不一致(Perry & Buden, 1999; Linkem et al., 2012), 加之1910年以后在台湾没有被再次发现, 该种在台湾的分布是误记(周文豪, 林思民, 陈俊海, 个人通讯), 《台湾两栖爬行动物图鉴》和《台湾两栖爬行类图鉴》也未收录(吕光洋等, 1999; 向高世等, 2009), 因此国内无翠蜥属物种分布。莎车隆趾虎(旧称南疆裸趾虎或莎车裸趾虎Altiphylax yarkandensis)被认为是宽斑隆趾虎(Altiphylax stoliczkai)的同物异名或在中国无分布(时磊, 2010; Bauer et al., 2013), 因此莎车隆趾虎未被收录。球趾虎科的托克逊沙虎(Teratoscincus toksunicus)被认为是争议种(陈彬和王跃招, 2001)或是吐鲁番沙虎的同物异名(Macey et al., 1997), 该种也未被收录。

2.2.2.2 蛇亚目

新增、恢复或提升了17属。原竹叶青蛇属(Trimeresurus)中拆分出了绿蝮属(Viridovipera)、喜山蝮属(Himalayophis)和华蝮属(Sinovipera)3个属(Malhotra & Thorpe, 2004; 郭鹏, 2005; Dawson et al., 2008; Guo et al., 2010; Guo & Wang, 2011; Pyron et al., 2013), 中文名随属名作相应改变。多项研究证明原锦蛇属(Elaphe)不是单系, 据此《名录》新增加了三索蛇属(Coelognathus)(Helfenberger, 2001; Utiger et al., 2005; Burbrink & Lawson, 2007; Pyron et al., 2013)、晨蛇属(Orthriophis)(Utiger et al., 2002, 2005; Burbrink & Lawson, 2007; Pyron et al., 2013)、紫灰蛇属(Oreocryptophis)(Utiger et al., 2005; Pyron et al., 2013)、玉斑蛇属(Euprepiophis)(Utiger et al., 2002, 2005; Burbrink & Lawson, 2007; Pyron et al., 2013; Chen et al., 2014a)和绿蛇属(Rhadinophis) (Burbrink & Lawson, 2007; Pyron et al., 2013), 中文名随属名变动作相应改变。Schulz等(2011)对方花小头蛇(Oligodon bellus)研究后发现该种与小头蛇属不在同一支系, 并以该种为模式物种建立了方花蛇属(Archelaphe), 专家会评时采纳该意见。王小荷(2014)研究表明宁陕小头蛇(Oligodon ningshaanensis)无论是从线粒体基因和核基因所表现的系统发育方面, 还是初步探讨其食性、半阴茎方面都表现出与食螺蛇亚科(Dipsadinae)关系密切, 因此建立以该蛇为模式物种的线形蛇属(Stichophanes) (Wang XH et al., 2014), 该蛇更名为宁陕线形蛇(Stichophanes ningshaanensis), 本名录采纳该建议。原广义游蛇属(Coluber sensu lato)中, 黄脊游蛇(Coluber spinalis)被归入鞭蛇属(Hierophis), 称为黄脊鞭蛇(Hierophis spinalis)(Nagy et al., 2003), 但更多基因位点的研究认为Hierophis并不是单系(Nagy et al., 2004), 之后Kharin (2009)将黄脊游蛇作为模式种建立了东方蛇属(Orientocoluber), 专家会评时采纳该结果。我国原游蛇属中的花脊游蛇(Coluber ravergieri), 多被认为应归入秘纹蛇属(Hemorrhois), 称为秘纹蛇(Hemorrhois ravergieri)(Nagy et al., 2004; Schä etti & Monsch, 2004; Pyron et al., 2013), 名录采纳该项研究, 因此游蛇属物种在我国已无分布。分子系统进化关系显示原腹链蛇属(Amphiesma)物种不形成单系, 该类群至少包含3个种组(Guo et al., 2014), 从而Guo等(2014)恢复了东亚腹链蛇属(Hebius)和喜山腹链蛇属(Herpetoreas), 因此, 本名录采纳该结果, 将原广义腹链蛇属拆分为3属(AmphiesmaHebiusHerpetoreas), 中文名不变。分子系统发育研究显示原水蛇属(Enhydris)不是单系(Kumar et al., 2012; Pyron et al., 2013), 从而Kumar等(2012)在此基础上结合形态学研究以中国水蛇(Enhydris chinensis)为模式建立了沼蛇属(Myrrophis), 之后, Murphy和Voris (2014)分别以铅色水蛇(Enhydris plumbea)和腹斑水蛇(E. bocourti)为模式种建立了铅色蛇属(Hypsiscopus)和腹斑蛇属(Subsessor), 本名录采纳该研究结果, 种的中文名随属名作相应改变。

合并了9属。莽山烙铁头蛇属(Zhaoermia)被并入原矛头蝮属(Protobothrops)(Guo et al., 2007; Guo P et al., 2012), 中文名也随属名作相应改变; 棘眦海蛇属(Acalyptophis)、棘鳞海蛇属(Astrotia)、平颏海蛇属(Lapemis)、长吻海蛇属(Pelamis)、海蝰属(Praescutata)和截吻海蛇属(Kerilia)被并入海蛇属(Hydrophis) (Pyron et al., 2013; Sanders et al., 2013), 中文名不变。链蛇属(Dinodon)被并入白环蛇属(Lycodon) (Guo et al., 2013; Pyron et al., 2013; Siler et al., 2013), 由于眼镜蛇科中Bungarus属的中文名为“ 环蛇属” , 为避免中文名混淆, 会评专家同意将“ 链蛇属” 中文名保留给Lycodon, “ 白环蛇属” 名称废弃, 除Lycodon septentrionalis(根据拉丁文原意, septentrionalis有北方的意思, 因此中文名“ 白链蛇” 废弃) 中文名改为“ 北链蛇” 之外, 其他种中文名随属名作相应改变。乌梢蛇属(Zaocys)被认为应并入鼠蛇属(Ptyas)(David & Das, 2004; Utiger et al., 2005), 黄松(个人通讯)认为乌梢蛇属物种间在形态和生理上存在部分差异, 建议继续保留乌梢蛇属各种的中文名以示区别, 专家会评时采纳该建议。

变更2属。Hedges等(2014)利用分子系统发育关系对盲蛇科研究后, 建立亚洲盲蛇属(Asiatyphlops)和印度盲蛇属(Indotyphlops)。但Pyron和Wallach (2014)认为按照国际动物命名法的优先原则, Asiatyphlops属无效, 因为东南亚盲蛇属(Argyro- phis)占先(Gray, 1845), 因此, 名录采纳Pyron和Wallach(2014)的研究, 将我国盲蛇科原有的钩盲蛇属(Ramphotyphlops)和盲蛇属(Typhlops), 更改为印度盲蛇属和东南亚盲蛇属, 原中文名不变。

新增、恢复或提升了37种。如: 黑顶钝头蛇(Pareas nigriceps)(Guo & Deng, 2009)、大别山原矛头蝮(Protobothrops dabieshanensis)(Huang X et al., 2012; Zhang et al., 2013)、喜山原矛头蝮(P. himalayanus)(Pan et al., 2013)、角原矛头蝮(P. cornutus) (龚世平等, 2010)、茂兰原矛头蝮(P. maolanensis) (Yang et al., 2011a; Liu et al., 2012)、冈氏绿蝮(Viridovipera gumprechti)(David et al., 2002; Malhotra & Thorpe, 2004; 郭鹏, 2005; Dawson et al., 2008; Guo et al., 2010; Guo & Wang, 2011; Pyron et al., 2013)、四川华蝮(Sinovipera sichuanensis)(Guo & Wang, 2011)、庙岛蝮(Gloydius lijianlii)(江帆和赵尔宓, 2009)、坎氏晨蛇(Orthriophis cantoris)(Utiger et al., 2005; 罗伟雄等, 2014)、若尔盖锦蛇(Elaphe zoigeensis)(Huang S et al., 2012)、云南华游蛇(Sinonatrix yunnanensis)(Rao & Yang, 1998; 杨大同和饶定齐, 2008)、福清链蛇(Lycodon futsingensis)(Vogel et al., 2009; Guo et al., 2013)、东川链蛇(L. synaptor)(Vogel & David, 2010)、刘氏链蛇(L. liuchengchaoi)(Zhang et al., 2011)、贡山链蛇(L. gongshan)(Vogel & Luo, 2011)、刘氏后棱蛇(Opisthotropis laui)(Yang et al., 2013)、黄斑后棱蛇(O. maculosa)(Stuart & Chuaynkern, 2007; Yang et al., 2011b)、香港过树蛇(Dendrelaphis hollinrakei)(Lazell, 2002)、黄斑异色蛇(Xenochrophis flavipunctatus)(Vogel & David, 2006; Vidal et al., 2007)、四川温泉蛇(Thermophis zhaoermii)(Guo et al, 2008)、方斑小头蛇(Oligodon nagao)(David et al., 2012)和束纹小头蛇(O. fasciolatus)(张君等, 2011), 以及未列入《红色名录》而增补进《名录》的广东颈槽蛇(Rhabdophis guangdongensis) (Zhu et al., 2014)、察隅腹链蛇(Herpetoreas burbrinki)(Guo et al., 2014)和香格里拉温泉蛇(Thermophis shangrila)(Penget al., 2014)。根据相关研究, 会评专家同意将红沙蟒-东方沙蟒复合体(Eryx miliaris-tataricus complex)在我国重新恢复为红沙蟒(E. miliaris)和东方沙蟒(E. tataricus), 将台湾钝头蛇-中国钝头蛇复合体(Pareas formosensis-chinensis complex)重新恢复为中国钝头蛇(P. chinensis)和台湾钝头蛇(P. formosensis)(Guo YH et al., 2011); 将蚺属(Python)中蚺双带亚种(Python molurus bivittatus)提升为种Python bivittatus(Jacobs et al., 2009), 按传统叫法将原中文名“ 蚺” 重新规定为“ 蟒” ; 圆斑蝰泰国亚种(Daboia russellii siamensis)提升为泰国圆斑蝰(D. siamensis)(Thorpe et al., 2007), 草原蝰东方亚种(Vipera ursinii renardi)提升为东方蝰(V. renardi)(Herrmann et al., 1992; Gvozdik et al., 2012), 山烙铁头蛇台湾亚种(Ovophis monticola makazayazaya)提升为台湾烙铁头蛇(O. makazayazaya) (Malhotra et al., 2011), 将赤链蛇南方亚种(Dinodon rufozonatum meridionale)提升为南方链蛇(Lycodon meridionale)(Orlov & Ryabov, 2004; 罗键等, 2010)。根据相关研究, 会评专家同意恢复高原蝮(Gloydius strauchi)种组中的雪山蝮(G. monticola)、六盘山蝮(G. liupanensis)、秦岭蝮(G. qinlingensis)(Xu et al., 2012)。Orlov等(2013)将白头蝰属(Azemiops)物种拆分为两种, 大致以越南的河内以及中国昆明和攀枝花为界, 以东为新种Azemiops kharini, 该种主要在中国分布, 且该类群在未拆分前就因头部特征被称之为“ 白头蝰” , 因此, 将其中文名冠以“ 白头蝰” ; 以西为Azemiops feae, 中文名则改为“ 黑头蝰” 。将台湾华珊瑚蛇(Sinomicrurus sauteri)拆分出新种羽鸟氏华珊瑚蛇(S. hatori), 原S. sauteri的中文名“ 台湾华珊瑚蛇” 已不合适, 则沿用台湾名称“ 梭德氏华珊瑚蛇” (Ota et al., 1999; Slowinski et al., 2001; 向高世等, 2009)。

种的属级分类阶元变动1种。Lei等(2014)认为横纹小头蛇(Oligodon multizonatus)应该归入链蛇属(Lycodon), 改为横纹链蛇(Lycodon multizonatum), 专家会评时采纳该建议。

未收录专家会评时认为国内无分布的2种。张君等(2011)查看中国四线小头蛇(Oligodon quadrilineatus)和管状小头蛇(O. cyclurus)新记录所依据的标本, 并对照国内外该属物种相关描述后, 认为四线小头蛇与管状小头蛇实际应该分别为紫棕小头蛇(Oligodon cinereus)和束纹小头蛇(O. fasciolatus), 从而认为四线小头蛇与管状小头蛇在中国无分布, 专家会评时采纳该结果。

此外, 专家会评时还对蛇亚目一些物种的中文名作了调整。评审专家根据Xenochrophis拉丁名原意以及形态特征, 建议将原中文名“ 渔游蛇属” 改为“ 异色蛇属” , 名录采纳。

本名录未收录区域灭绝种, 如食鱼鳄科(Gavialidae)切喙鳄属(Tomistoma)的马来切喙鳄(Tomistoma schlegelii)和鳄科(Crocodylidae)鳄属(Crocodylus)的湾鳄(Crocodylus porosus) (张孟闻等, 1998)。也未收录外来物种, 如佛罗里达鳖(Apalone ferox)、彩龟红耳亚种(Trachemys scripta elegans)等。

3 讨论

确立爬行纲各物种系统发育关系有利于分类学研究。然而形态与分子系统发育研究结果差异很大, 尤其是科级以上分类阶元。

3.1 目、亚目级分类阶元的争议

近年来分子系统学研究认为爬行纲是并系群(Hedges & Poling, 1999; Lu et al., 2013), 龟鳖目、喙头目(Rhynchocephalia)、有鳞目和鳄形目为单系群, 但各目之下亚目的划分存在争议(Fujita et al., 2004; Townsend et al., 2004; Barley et al., 2010; Oaks, 2011; Pyron et al., 2013; Jaratlerdsiri et al., 2014), 特别是有鳞目亚目的划分存在较大分歧(Townsend et al., 2004; Conrad, 2008; Pyron et al., 2013), 主要体现在分子系统学与形态学研究结果的不一致上。因此本文在目与亚目级别采用经典分类体系(Zug et al., 2001; Vidal & Hedges, 2005; Lillywhite & Wray, 2008; Vitt & Caldwell, 2013; Dijk et al., 2014), 即中国爬行纲包括鳄形目、龟鳖目与有鳞目(蜥蜴亚目和蛇亚目)。

有鳞目方面, 目前形态和分子生物学研究认为应将其分为鬣蜥-硬舌类(Iguania-Scleroglossa)、壁虎类(Gekkota)、石龙子类(Scincomorpha)、蛇蜥类(Anguimorpha)、蚓蜥类(Amphisbaenia)、蜥蜴类(Lacertoidea)、蛇类(Serpentes)和其他总科(Townsend et al., 2004; Lee, 2005; Conrad, 2008; Wiens et al., 2010; Gauthier et al., 2012; Pyron et al., 2013), 并且其分类阶元随着研究方法的不同有亚目、下目(Infraorder)和总科的变动, 但学界目前尚无统一认识。有鳞目科级以上分类阶元之间存在较大争议, 尤其是蜥蜴亚目和蚓蜥亚目(Amphisbaenia) (Townsend et al., 2004; Conrad, 2008; 芮金龙等, 2009; Pyron et al., 2013)。

3.2 科级分类阶元的争议

蜥蜴亚目方面, 越来越多的研究认为传统分类学上的蜥蜴亚目是并系群, 科级分类阶元之间的系统发育关系也存在较大分歧(Townsend et al., 2004; Lee, 2005; Conrad, 2008; Wiens et al., 2010; Gauthier et al., 2012; Pyron et al., 2013)。近年来, 分子系统学研究更多认为双足蜥科(Dibamidae)是现生有鳞目动物中最原始的类群, 其次是广义壁虎科(Gekkonidae sensu lato)和石龙子科(Townsend et al., 2004; Pyron et al., 2013)。目前普遍认为广义壁虎科应包括睑虎科(Eublepharidae)、球趾虎科、壁虎科(Gekkonidae)、叶趾虎科(Phyllodactylidae)、鳞脚虎科(Pygopodidae)、藁趾虎科(Carphodactylidae)和澳虎科(Diplodactylidae) (Gamble et al., 2008, 2011, 2012; Pyron et al., 2013)。

蛇亚目中, 近年来无论是形态学还是分子系统学的研究, 都认为蛇亚目是单系群(Townsend et al., 2004; Conrad, 2008; Pyron et al., 2013)。科级分类阶元方面, 不少研究认为原游蛇科(Colubridae)不是单系, 而应该被拆分为游蛇科(Colubridae)、闪皮蛇科、钝头蛇科、水蛇科和鳗形蛇科等(Vidal et al., 2007; Wiens et al., 2008; Zaher et al., 2009; Pyron et al., 2011, 2013)。

3.3 属、种级分类阶元的争议

蜥蜴亚目中, 壁虎科的墨脱弯脚虎(Cyrtopo- dion medogensis)是一个单独的进化支系, 究竟是属于裸趾虎属还是弯脚虎属, 目前尚无定论(时磊, 2010), 因此本文将其暂时归入弯脚虎属(Shi & Zhao, 2011; Bauer et al., 2013)。石龙子科的锡金滑蜥(Scincella sikimmensis)、拉达克滑蜥(S. ladacensis)和喜山滑蜥(S. himalayanus)被认为应归属滑蜥属(王跃招和赵尔宓, 1986; Murthy, 2010), 或裂睑蜥属(Sindaco & Jeremč enko, 2008; Venugopal, 2010), 或喜山蜥属(Himalblepharus)(Jeremč enko, 1987; Sindaco & Jeremč enko, 2008), 但分子系统学研究结果显示锡金滑蜥可与滑蜥属物种聚合在一起(Pyron et al., 2013), 鉴于裂睑蜥属不是单系(Pyron et al., 2013), 因此本文暂将这3个种保留在滑蜥属。鬣蜥科中, 岩蜥属(Lacerta)物种曾被归为岩蜥属、星蜥属(Stellagama)和拟岩蜥属(Paralaudakia)(Baig et al., 2012), 但黄松(个人通讯)认为该文无分子数据, 加之形态上三属区别较弱, 建议维持原岩蜥属不变, 专家会评时采纳该建议; 沙蜥属中, Jin等(2013)认为贵南沙蜥可与贵德沙蜥聚合在一起, 但计翔认为(个人通讯)对贵南沙蜥和贵德沙蜥的抽样和分子标记不同会得出不同的结论, 他们的研究认为贵南沙蜥在形态和空间分布上介于青海沙蜥和贵德沙蜥之间, 贵南沙蜥因适应特殊生境而导致物种分化, 是贵德沙蜥的后裔, 为探讨高原隆起环境变化所致的生态物种分化提供了示例。专家会评时采纳计翔的建议暂时认为贵南沙蜥有效。

蛇亚目的蝰科中, 分子系统学研究显示, 原广义竹叶青蛇属(Trimeresurus sensu lato)中的台湾竹叶青蛇(Trimeresurus gracilis)和冲绳烙铁头蛇(Ovophis okinavensis)是姊妹种, 且与亚洲蝮属(Gloydius)中物种的亲缘关系较近, 与竹叶青蛇属和烙铁头蛇属(Ovophis)的关系较远(Dawson et al., 2008; Guo et al., 2010; Pyron et al., 2013), 但目前尚无形态和其他系统学方面的专门研究, 因此本文暂将其保留在广义竹叶青蛇属中; 另外, Malhotra和Thorpe(2004)认为白唇竹叶青蛇(Trimeresurus albolabris)应当归入腺蝮属(Cryptelytrops), 但David等(2011)认为该属无效, 应该保留在竹叶青属(亚属)(Trimeresurusus)中, 因此名录采纳该建议。游蛇科中, 原锦蛇属的绿锦蛇(Elaphe prasina)和灰腹绿锦蛇(E. frenata)被归入绿蛇属, 称为绿蛇(R. prasinus)和灰腹绿蛇(R. frenatus)(Burbrink & Lawson, 2007), Chen等(2014b)进一步研究认为绿蛇、灰腹绿蛇和尖喙蛇(Rhynchophis boulengeri)应该归入树栖蛇属(Gonyosoma), 但因该文尚未涉及原树栖蛇属所有物种形态学和分子生物学的研究, 因此, 本文暂未采纳该建议; 滇西蛇(Atretium yunnanensis)在一些分子系统学研究中显示聚合在异色蛇属(Xenochrophis)中(Pryon et al., 2013), 但目前该种形态学和系统发育学证据不充分, 因此我们暂采取保守的办法, 保留原分类地位不变(郭鹏, 个人通讯)。

除了分类学方面的问题, 部分物种首次记录源于市场收购或被发现于码头等货运地段等, 而多年后均无再次记录, 尤其是壁虎科和龟鳖目物种。其在国内存在与否, 有待更深一步的调查研究。因此, 我们将这些种类列为存疑种。

在不同生物类群、不同分类学家对种的标准有不同认识。对于一个分类单元的分合与有效性, 往往较难取得一致意见, 本名录通过多次网评和会评, 邀请爬行动物分类专家对中国爬行纲动物的分类体系和物种进行了系统的评估, 形成了较统一的认识, 缩小了我国爬行动物分类体系与国际公认的分类体系间的差异, 获得了目前中国爬行动物多样性最新最全面的数据。同时, 我们建议, 成立由爬行动物分类学专家学者组成的全国的爬行动物物种委员会, 定期评估新种新记录种及新的分类变动等, 规范中文名称, 形成意见统一的爬行动物名录, 以便科学研究与传播和物种保育。

目前, 我国爬行动物生存状况不容乐观, 不少物种处于受威胁状态, 导致个别类群分类研究难以开展, 尤其是龟鳖的分类研究困难重重。我们应该重视广泛收集物种种群变化数据, 定期评估物种受胁状况, 为国家生物多样性保育建言献策, 以改善爬行动物生存状况。

(责任编委: 傅金钟 责任编辑: 闫文杰)

附录1

附录1 中国爬行纲校正名录

Appendix 1 Checklist of Chinese Reptilia

http://www.biodiversity-science.net/fileup/PDF/w2015-037-1.pdf

附录2

附录2 科级分类阶元调整

Appendix 2 Taxonomic adjustment for families

http://www.biodiversity-science.net/fileup/PDF/w2015-037-2.pdf

附录3

附录3 属、种级分类阶元调整

Appendix 3 Taxonomic adjustment for genera and species

http://www.biodiversity-science.net/fileup/PDF/w2015-037-3.pdf

致谢: 在本研究中, 我们咨询了如下专家(按姓氏拼音排序)的意见: 丁利、丁由中、杜卫国、古晓东、顾海军、郭鹏、郭宪光、黄松、黄勇、计翔、蒋志刚、李俊、李丕鹏、林思民、刘惠宁、吕顺清、饶定齐、史海涛、时磊、汪继超、王剑、王小明、王小荷、王英勇、王正寰、肖向红、赵文阁、周文豪等, 在此表示衷心感谢!此外, 我们还要感谢袁思棋、孙晓煜、李大江、刘金龙、原秀云等同学帮助整理爬行动物名录。

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