湖南
莽山布角蟾属
28
Boulenophrys
29
物种
分类
讨论
兼记省新
记录
DOI:10.16605/ki.1007-7847.2022.12.0235湖南莽山布角蟾属(Boulenophrys)物种分类讨论兼记省新记录三种收稿日期:2022-12-07;修回日期:2023-03-03;网络首发日期:2023-04-27基金项目:国家林业和草原局野生动植物保护项目(2021-HN001);野生动物调查、监管和行业监管项目(2022072-HN-001)作者简介:尚袁凌博(1998),男,湖南长沙人,硕士研究生;*通信作者:杨道德(1965),男,湖南澧县人,教授,博士生导师,E-mail:。尚袁凌博1,罗堯1,钱天宇1,高志伟1,李永辉2,邓国杏2,杨道德1*(1.中南林业科技大学 野生动植物保护研究所,中国湖南 长沙 410004;2.湖南莽山国家级自然保护区管理局,中国湖南郴州 424200)摘要:准确的物种鉴定和编目是开展生物多样性保护与管理的前提。莽山位于湖南省南部、湘粤交界处,是南岭生物多样性优先保护区域,具有丰富的两栖动物物种多样性。湖南动物志 两栖纲 记载莽山分布有 2 种布角蟾属(Boulenophrys)物种:挂墩角蟾(B.kuatunensis)和短肢角蟾(B.brachykolos),但随着角蟾亚科(Megophryi-nae)系统分类研究的深入,目前这 2 个物种在莽山的分布存在争议。2021 年 56 月、78 月、11 月和 2022 年68 月,笔者团队对湖南莽山国家级自然保护区的布角蟾属物种进行了 4 次实地调查,并采集到 20 号标本。经形态和分子鉴定,这些标本包括南岭角蟾(B.nanlingensis)、雨神角蟾(B.cf.ombrophila)、九连山角蟾(B.jiulia原nensis)、石门台角蟾(B.shimentaina),并未发现挂墩角蟾和短肢角蟾。其中,雨神角蟾、九连山角蟾、石门台角蟾为湖南省新记录种。调查结果显示,莽山地区的挂墩角蟾和短肢角蟾记录需要对馆藏标本进一步检视研究。此外,石门台角蟾和南岭角蟾在新种发表时缺乏雌性形态特征描述,本文首次对这 2 个物种的雌性标本进行了补充描述。关键词:角蟾科;系统发育分析;挂墩角蟾;短肢角蟾;湖南莽山国家级自然保护区中图分类号:Q959.5+3文献标志码:A文章编号:1007-7847(2023)04-0349-12Taxonomic Discussion of Boulenophrys in Mangshan Mountain,Hunan,China and Three New Provincial RecordsSHANG Yuanlingbo1,LUO Yao1,QIAN Tianyu1,GAO Zhiwei1,LI Yonghui2,DENG Guoxing2,YANG Daode1*(1.Institute of Wildlife Conservation,Central South University of Forestry and Technology,Changsha 410004,Hunan,China;2.Administration of Hunan Mangshan National Nature Reserve,Chenzhou 424200,Hunan,China)Abstract:Accurate species identification is a prerequisite for biodiversity conservation and management.Mangshan Mountain,located in the south of Hunan and at the border of Hunan and Guangdong provinces,isa priority area for biodiversity protection in Nanling Mountains,with rich diversity of amphibian species.Ac-cording to the Fauna Hunan:Amphibia,there are two Boulenophrys species distributed in Mangshan:B.kua-tunensis and B.brachykolos,but with the further study of the systematic classification of subfamily Mego-phryinae,the distribution of the two species in Mangshan is currently controversial.During May to June,Julyto August,November 2021 and June to August 2022,four field surveys on the species of Boulenophrys inMangshan National Nature Reserve,Hunan Province were conducted,and 20 specimens were collected.Through morphological and molecular identification,these specimens proved to belong to four species:B.nanlingensis,B.cf.ombrophila,B.jiulianensis,and B.shimentaina,and the latter three species are newrecords in Hunan Province.However,the species B.kuatunensis and B.brachykolos were not found to exist第 27 卷 第 4 期生命科学研究灾燥造援27 晕燥援4圆园23 年 8 月蕴蚤枣藻 杂糟蚤藻灶糟藻 砸藻泽藻葬则糟澡Aug.圆园23窑 分类与生态学 窑生命科学研究圆园23 年莽山位于湖南省南部,处于湘粤交界处,是南岭生物多样性优先保护区域的一部分,具有丰富的两栖动物物种多样性1。有关莽山地区角蟾类物种的记录,最早可追溯到 1922 年德国动物学家鲁道夫 梅尔(Rudolf Mell)有关中国动物区系的论著 Beitr覿ge zur Fauna Sinica2。该书记录了在湘粤边界山地采集到的 1 号淡肩角蟾(Megophrysboettgeri)的蝌蚪,从分布图可知采集点为今湖南莽山与广东南岭交界的范围。1976 年,四川生物所两栖爬行室发表 湖南省宜章莽山两栖动物调查报告,报道该地区有小角蟾(Megophrys minor)分布3。后来,基于 19591981 年间的调查工作,沈猷慧先生于 1983 年发表了 湖南省两栖动物调查及区系分析4,该文报道在莽山共采集到 30种两栖动物,其中两种小型角蟾被鉴定为挂墩角蟾(Boulenophrys kuatunensis)和小角蟾,同时提及未采集到淡肩角蟾。随着小角蟾复合体分类的研究推进5,沈猷慧先生经重新检视标本后,在 2014年出版的 湖南动物志 两栖纲6中将莽山地区分布的几种小型角蟾重新修订为挂墩角蟾和短肢角蟾(B.brachykolos),并删除了小角蟾的记录。近年来,基于大范围采样的分子研究结果显示,挂墩角蟾和短肢角蟾在其模式产地外并无广泛分布7-8,其模式产地外的记录应考虑重新审视9。挂墩角蟾的模式产地位于福建挂墩,距莽山直线距离超过500 km,高志伟等10认为湖南省挂墩角蟾的分布记录存疑。而短肢角蟾在其模式产地外的记录也陆续被修订,如广西大瑶山的种群最近被描述为新种瑶山角蟾(B.yaoshanensis)11。20212022 年,笔者团队在湖南莽山进行了一系列较为全面的角蟾类物种调查,Qian 等12通过对采集的角蟾蝌蚪进行鉴定发现了 3 种布角蟾属(Boulenophrys)物种,分别为南岭角蟾(B.nanlingensis)、石门台角蟾(B.shimentaina)和雨神角蟾(B.cf.ombrophila)。在本研究中,我们将同蝌蚪采集的成体个体,与后续调查新采集的共 20 号标本进行鉴定。结果显示,除以上 3 种外,莽山还分布有第 4 种布角蟾属物种九连山角蟾(B.jiulianensis),但并未发现挂墩角蟾和短肢角蟾。本文对这 4 种角蟾分别报道,并首次描述南岭角蟾和石门台角蟾的雌性标本特征。1研究地自然概况湖南莽山国家级自然保护区(以下简称“莽山保护区”),位于湖南省郴州市宜章县境内,与广东接壤,地处南岭山脉中段南段,地理坐标为112.72毅 113.00毅 E、24.88毅 25.05毅 N,总 面 积 为198.33 km2。海拔范围 4001 902 m,海拔 100 m以上的山峰有 111 座,1 500 m 以上的山峰有 25座。莽山属亚热带湿润季风气候区,是中国最南端的冬季冰雪地区之一,年均气温 17.2 益,年降水量 1 6002 300 mm,年无霜期 290 d1。森林植被垂直分布明显:海拔 500 m 以下主要为常绿阔叶林;5001 200 m 为常绿落叶和阔叶混交林;1 2001 600 m 为针阔混交林;1 600 m 以上为高山灌丛和高山矮林13。2研究方法2.1标本采集和组织取样2021 年 5 月下旬6 月上旬、7 月中旬8 月中旬、11 月上旬和 2022 年 68 月,采用样线法在莽山保护区对角蟾亚科物种进行了 4 次实地调查,平均每次调查 15 d。每天调查时间为 19:0023:3014。结合角蟾生态习性,在莽山保护区内的猴王寨、枞树坝、相思坑、鬼子寨、天台山、猛坑石等地共布设 15 条样线,样线单侧宽为 2 m,样线长在地势较开阔平坦的区域为 7001 000 m,在地形复杂的区域为 300500 m。调查期间共采集了 20 号布角蟾属标本(采集位置见图 1)。取组织样品(肝脏)并将其保存于95%的乙醇溶液中,用于分子测序;标本经 10%的福尔马林溶液固定后转移到 75%的乙醇溶液中保存,标本存放于中南林业科技大学动物标本室。in Mangshan.Therefore,the existence of the two species in Mangshan should be further confirmed.In addi-tion,the morphological characteristics of female B.shimentaina and B.nanlingensis were described for thefirst time,which were lack in the original new record description.Key words:Megophryidae;phylogenetic analysis;Boulenophrys kuatunensis;Boulenophrys brachykolos;Hu-nan Mangshan National Nature Reserve(Life Science Research,2023,27(4):349-360)350第 4 期尚袁凌博等:湖南莽山布角蟾属(Boulenophrys)物种分类讨论兼记省新记录三种图 1湖南莽山布角蟾属标本采集点位置示意图Fig.1Collection locations of Boulenophrys specimens in Mangshan area of Hunan Province2.2分子实验和系统发育分析使用 TSP201-200 DNA 试剂盒(擎科生物科技股份有限公司)对 14 份肝脏样品分别进行 16SrRNA 和细胞色素氧化酶 1(cytochrome oxidase 1,CO1)基因片段扩增,随后测序。其中,16S rRNA基因引物选择 L3975 和 H455115,CO1 基因引物选择DgLCO 和 DgHCO16。PCR 过程同参考文献17,PCR 和测序过程由百迈客生物科技有限公司执行。从 GenBank(https:/www.ncbi.nlm.nih.gov/;2022-08-29)下载除水城角蟾(B.shuichengensis)外所有已描述的 63 种布角蟾属物种的序列(相关信息见表 1),并使用腺角蟾(Xenophrys glandulosa)和莽山角蟾(X.mangshanensis)作为外群11。利用 MEGA 7.0中的 MUSCLE 将所有 16S rRNA 和 CO1 基因分别进行比对18,随后将两个片段串联成一个 1 080 bp的数据集。基于贝叶斯信息准则(Bayesian Infor-mation Criterion,BIC),采用 ModelTest-NG19选择最佳核苷酸替代模型,得到 16S rRNA 和 CO1 的最佳拟合核苷酸替代模型为 GTR+I+G。使用MrBayes 3.2.420构建 BI(Bayesian inference)系统发育树,按如下方法运行:两条链同时运行 1 000万代,每 1 000 代取样一次,最初的 25%代作为老化样本(burn-in)舍弃,系统各分支的置信度通过后验概率(Bayesian posterior probability,BPP)进行检验。最后,使用 MEGA 6.0 中的未修正距离模型(uncorrected p-distance)计算物种间遗传距离21。2.3形态测量使用游标卡尺(精度 0.1 mm)对标本的形态特征进行测量。量度包括:头体长(snout-vent length,SVL)、头长(head length,HL)、头宽(head width,HW)、吻长(snout length,SL)、鼻间距(internasal space,INS)、眼径(diameter of eye,ED)、鼓膜径(diameter of tym-panum TD)、眼间距(interorbital space,IOS)、前臂及手长(length of lower arm and hand,LAHL)、鼓膜与眼间距(tympanum-eye distance,TYE)、跗足长(leng-th of foot and tarsus,LFT)、胫长(tibia length,TL)、足长(foot length,FL)。外形测量和外部形态特征及其缩写依据 中国动物志 两栖纲(中卷)22。3结果3.1分子鉴定结果在重建的布角蟾属系统发育结构(图 2)中,莽山新采集的标本分别与南岭角蟾、雨神角蟾、九连山角蟾和石门台角蟾聚为一支。其中,除南岭角蟾与其模式产地的标本分化较小外,其余均表现出一定程度的分化。这可能与地理距离有关。Qian等12指出,莽山地区标本的采集位点同南岭角蟾和石门台角蟾模式标本采集地的最近距离分别约为 10 km 和 70 km,同雨神角蟾 3 个已知分布位点的距离为 70500 km。此外,本研究新发现的九连山角蟾的模式产地包括江西九连山和广东南昆山,两地均距莽山约 170 km。基于 16S rRNA 的遗传距离计算结果显示,莽山分布的待鉴定谱系 1 与南岭角蟾模式标本的遗传距离为 00.8%,待鉴定谱系 2 与雨神角蟾的遗传距离为 2.0%2.8%,待鉴定谱系 3 与已知分布点九连山角蟾的遗传距离为 0.6%0.8%,待鉴定谱系 4 与石门台角蟾的遗传距离为 2.0%。这些112毅52忆30寓E112毅47忆0寓E112毅58忆0寓E112毅47忆0寓E112毅52忆30寓E112毅58忆0寓E南岭角蟾图例保护站九连山角蟾石门台角蟾雨神角蟾地名湖南莽山国家级自然保护区省界高程低:405 m高:1 887 m351生命科学研究圆园23 年表 1样品信息及其 GenBank 登录号Table 1Samples and GenBank accession numbers used in this studySpecies nameBoulenophrysB.acutaB.angkaB.anlongensisB.baishanzuensisB.baolongensisB.binchuanensisB.binlingensisB.boettgeriB.brachykolosB.caobangensisB.caudoproctaB.cheniB.chishuiensisB.congjiangensisB.daiyunensisB.daojiB.daweimontisB.dongguanensisB.fansipanensisB.fengshunensisB.frigidaB.hoanglienensisB.huangshanensisB.hungtaiB.insularisB.jiangiB.jingdongensisB.jinggangensisB.jiulianensisB.jiulianensisB.jiulianensisB.jiulianensisB.jiulianensisB.jiulianensisB.jiulianensisB.kuatunensisB.leishanensisB.liboensisB.liniB.lishuiensisB.lushuiensisB.minorB.mirabilisB.mufumontanaB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisB.nanlingensisVoucher numberSYSa002159KIZ040592CIBAL21090531017CIBQY20200719001KIZ019216KIZ019442SYSa005313SYSa004149SYSa002258IBER4385SYSa004281SYSa002142CIBCS20190518019GZNU20200706003SYSa001733SYSa004088KIZ048938SYSa001973AMSR186115SYSa004724AMSR186131VNMN2018.02SYSa002703SYSa007575SYSa002168CIBKKS20180426001SYSa003928SYSa004028CSUFT2207M13SYSa002107SYSa002108SYSa002109SYSa004219SYSa003622SYSa003623SYSa003449SYSa002213GNUG20150813001SYSa002381CIBWYF00169CIBYN201909288SYSa003209SYSa002192SYSa006391CSUFT2107M82CSUFT2107M85CSUFT2107M74SYSa001959SYSa001960SYSa001964SYSa002334SYSa002356SYSa002357SYSa002358LocalityHeishiding Nature Reserve,Guangdong,ChinaKew Mae Pan Nature Trail,Doi Inthanon,Chiang Mai,ThailandAnlong County,Guizhou,ChinaBaishanzu National Park,Qingyuan,Zhejiang,ChinaBaolong Town,Chongqing,ChinaMt.Jizu,Yunnan,ChinaMt.Wawu,Sichuan,ChinaMt.Wuyi,Fujian,ChinaHong Kong,ChinaNguyen Binh,Cao Bang,VietnamZhangjiajie,Hunan,ChinaTaoyuandong Nature Reserve,Hunan,ChinaChishui County,Guizhou,ChinaYueliangshan Nature Reserve,Congjiang,Guizhou,ChinaDaiyun Mountain Nature Reserve,Fujian,ChinaFenghua City,Zhejiang,ChinaMt.Dawei,Yunnan,ChinaMt.Yinping,Guangdong,ChinaSa Pa,Lao Cai,VietnamFengshun County,Guangdong,ChinaKy Quan San Mountain,Bat Xat District,Lao Cai,VietnamSa Pa,Lao Cai,VietnamMt.Huangshan,Anhui,ChinaJiexi County,Guangdong,ChinaNan ao Island,Guangdong,ChinaKuankuoshui Nature Reserve,Guizhou,ChinaMt.Wuliang,Yunnan,ChinaMt.Jinggang,Jiangxi,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Jiulian,Ganzhou City,Jiangxi,ChinaMt.Jiulian,Ganzhou City,Jiangxi,ChinaMt.Jiulian,Ganzhou City,Jiangxi,ChinaMt.Jiulian,Ganzhou City,Jiangxi,ChinaMt.Nankun,Ganzhou City,Jiangxi,ChinaMt.Nankun,Ganzhou City,Jiangxi,ChinaMt.Wuyi,Fujian,ChinaMt.Leigong,Guizhou,ChinaLibo,Guizhou,ChinaTaoyuandong Nature Reserve,Hunan,ChinaLishui,Zhejiang,ChinaLangbazhai,Luzhang,Lushui County,Nujiang Prefecture,Yunnan,ChinaDujiangyan,Sichuan,ChinaHuaping Nature Reserve,Guangxi,ChinaMt.Mufu,Pingjiang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaNanling Nature Reserve,Shaoguan City,Guangdong,ChinaNanling Nature Reserve,Shaoguan City,Guangdong,ChinaNanling Nature Reserve,Shaoguan City,Guangdong,ChinaMt.Qiyun,Chongyi County,Jiangxi,ChinaMt.Qiyun,Chongyi County,Jiangxi,ChinaMt.Qiyun,Chongyi County,Jiangxi,ChinaMt.Qiyun,Chongyi County,Jiangxi,China16S rRNAMF667869MN508048MT823185MW001150KX811813KX811850MH406892MF667878KJ560403LC483945MH406795KJ560398MN954708MW959773MH406643MH406782KX811870MH406647MH514887MH406848MT364279MH514889MF667883OL635592MF667886MN107744MH406773MH406780OQ517199MK524099MK524100MK524101MH406791MK524102MK524103MF667881MH406673MF285253MF667874KY021418MW001225MF667862MH406669MK524105ON406219ON406220ON406221MK524111MK524112MH406646MH406686MK524113MH406687MH406688CO1MH406099/MT823262MT998291KX812093KX812113MH406354MH406247MH406120/MH406257MH406098MN928959MW959761MH406079MH406242KX812126MH406083MW086548MH406310MW086550MW086551MH406161OL634859MF667923MN107749MH406232MH406239OQ517953MK524130MK524131MK524132MH406253MK524133MK524134MH406206MH406113/MH406135/MW000912MH406194MH406109MK524136/MK524142MK524143MH406082MH406132MK524144MH406133MH406134GenBank accession numberID123456789101112131415161718192021222324252627282930313233343536373839404142434445464748495051525354352第 4 期注:“/”代表无此序列。表 36 相同。Note:“/”represents no such sequence.These are same in the Tables 36.表 1(续)GenBank accession numberID555657585960616263646566676869707172737475767778798081828384858687888990919293949596979899100101102103104Species nameB.nankunensisB.obesaB.cf.ombrophilaB.cf.ombrophilaB.cf.ombrophilaB.cf.ombrophilaB.cf.ombrophilaB.cf.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.ombrophilaB.omeimontisB.palpebralespinosaB.puningensisB.qianbeiensisB.rubrimeraB.sangzhiensisB.sanmingensisB.shimentainaB.shimentainaB.shimentainaB.shimentainaB.shimentainaB.shimentainaB.shimentainaB.shimentainaB.shunhuangensisB.spinataB.tongboensisB.tuberogranulatusB.wugongensisB.wuliangshanensisB.wushanensisB.xiangnanensisB.xianjuensisB.yangmingensisB.yaoshanensisB.yingdeensisB.yunkaiensisXenophrysX.glandulosaX.mangshanensisVoucher numberSYSa004501SYSa002272CSUFT2105M09CSUFT2105M12CSUFT2105M19CSUFT2105M33CSUFT2105M24CSUFT2105M32CIBWY18082308WUYI12015101SYSa004612SYSa004613SYSa004614SYSa004615SYSa004616SYSa001976SYSa001977SYSa001978SYSa001985SYSa004583SYSa005301KIZ011650SYSa005770CIBTZ20190608017AMSR177676SYSa004306SYSa002486CSUFT2105M35CSUFT2107M70CSUFT2107M86CSUFT2107M98SYSa002077SYSa002081SYSa004172SYSa004173HNNU18NS01SYSa002226SYSa002415SYSa004310SYSa004800SYSa003924SYSa003008SYSa002874CIBXJ190505SYSa002877SYSa002189SYSa002100SYSa004637SYSa003758SYSa002177LocalityMt.Nankun,Guangdong,ChinaHeishiding Nature Reserve,Guangdong,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Wuyi,Fujian,ChinaMt.Wuyi,Fujian,ChinaMt.Jiulian,Jiangxi,ChinaMt.Jiulian,Jiangxi,ChinaMt.Jiulian,Jiangxi,ChinaMt.Jiulian,Jiangxi,ChinaMt.Jiulian,Jiangxi,ChinaRenhua County,Guangdong,ChinaRenhua County,Guangdong,ChinaRenhua County,Guangdong,ChinaRenhua County,Guangdong,ChinaRenhua County,Guangdong,ChinaMt.Le,Sichuan,ChinaPu Hu Nature Reserve,Thanh Hoa,VietnamPuning City,Guangdong,ChinaHuanglian Nature Reserve,Guizhou,ChinaSa Pa,Lao Cai,VietnamZhangjiajie,Hunan,ChinaMt.Emeifeng,Fujian,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaMt.Mang,Yizhang County,Hunan,ChinaShimentai Nature Reserve,Guangdong,ChinaShimentai Nature Reserve,Guangdong,ChinaShimentai Nature Reserve,Guangdong,ChinaShimentai Nature Reserve,Guangdong,ChinaNanshan Forest Park,Hunan,ChinaMt.Leigong,Guizhou,ChinaMt.Tongbo,Jiangxi,ChinaBadagongshan Nature Reserve,Hunan,ChinaMt.Wugong,Jiangxi,ChinaMt.Wuliang,Yunnan,ChinaMt.Wu,Hubei,ChinaMt.Yangming,Hunan,ChinaXianju County,Zhejiang,ChinaMt.Yangming,Hunan,ChinaDayaoshan Nature Reserve,Guangxi,ChinaShimentai Nature Reserve,Guangdong,ChinaYukaishan Nature Reserve,Guangdong,ChinaMt.Gaoligong,Yunnan,ChinaMt.Sanyue,Guangdong,China16S rRNAMH406822KJ579122MZ570079MZ570080MZ570081MZ570082OL711911ON209286MW001159KX856397MH406836MH406837MH406838MH406839MH406840MH406650MH406651MH406652MH406653MH406834MH406887KX811889OL635585MT651554MF536419MH406797MH406697OL347449OL347446OL347447OL347448MH406655MH406656MH406787MH406788MK836023MH406675MH406690MH406801MH406853MH406771MH406732MH406713MN563753MH406716MH406667MH406658MH406843MH406755MH406666CO1MH406284MH406124OL674543OL674544OL674545OL674546/MT998300/MH406298MH406299MH406300MH406301MH406302MH406086MH406087MH406088MH406089MH406296MH406349KX812138OL634853MT654521MW086542MH406259MH406145OQ517954OQ517955OQ517956OQ517957MH406092MH406093MH406249MH406250MK977594MH406115MH406138MH406263MH406315MH406230MH406184MH406165MN563769MH406168MH406107MH406095MH406305MH406214MH406106待鉴定谱系与已知种间的遗传距离均小于 Fou-quet 等23提出的基于 16S rRNA 基因的两栖动物种间遗传距离界限 3.0%,但待鉴定谱系 2 与雨神角蟾、待鉴定谱系 4 和石门台角蟾的遗传距离大于或等于 Chen 等7建议的角蟾亚科物种间的遗传距离界限 2.0%。基于此,我们将莽山的标本进尚袁凌博等:湖南莽山布角蟾属(Boulenophrys)物种分类讨论兼记省新记录三种353生命科学研究圆园23 年一步与以上物种的原始描述进行了比对,发现其形态特征均符合其分子鉴定结果(表 2),因此暂将莽山地区的雨神角蟾、九连山角蟾、石门台角蟾定为湖南省新记录种,现将标本观察结果描述如下。图 2基于线粒体 16S rRNA 及 CO1 基因联合序列构建的贝叶斯系统发育树节点附近的数字表示贝叶斯后验概率。Fig.2The Bayesian phylogenetic tree derived from DNA sequences of the mito