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Article(id=1246833313126700032, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1246833307606995898, articleNumber=null, orderNo=null, doi=10.3969/j.issn.0253-4193.2019.08.006, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1554048000000, receivedDateStr=2019-04-01, revisedDate=1560355200000, revisedDateStr=2019-06-13, acceptedDate=null, acceptedDateStr=null, onlineDate=1775197826891, onlineDateStr=2026-04-03, pubDate=1566662400000, pubDateStr=2019-08-25, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1775197826891, onlineIssueDateStr=2026-04-03, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1775197826891, creator=13701087609, updateTime=1775197826891, updator=13701087609, issue=Issue{id=1246833307606995898, tenantId=1146029695717560320, journalId=1149651085930835976, year='2019', volume='41', issue='8', pageStart='1', pageEnd='140', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1775197825576, creator=13701087609, updateTime=1775200503343, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1246844539051332282, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1246833307606995898, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1246844539051332283, tenantId=1146029695717560320, journalId=1149651085930835976, issueId=1246833307606995898, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=63, endPage=71, ext={EN=ArticleExt(id=1246833313520963603, articleId=1246833313126700032, tenantId=1146029695717560320, journalId=1149651085930835976, language=EN, title=Molecular phylogeny of the sesarmid crab based on the partial sequences of mitochondrial COI and 16S rRNA genes from the coast of China, columnId=1243954927383462170, journalTitle=Haiyang Xuebao, columnName=Marine Biology, runingTitle=null, highlight=null, articleAbstract=

The species in the family of Sesarmidae are among the species mostly difficult to be differentiated taxonomically in the Superfamily Grapsoidea because of their morphological similarity. The analysis of partial sequences of mitochondrial COI gene and 16S rRNA gene of 14 species in 8 genera of Sesarmidae for their molecular phylogenetics reveals that the percentages of sequence divergences of 14 species of Sesarmidae are from 5.7% to 14.5% and from 1.5% to 12.1%, both indicating that the species difference are significant. The phylogenetic trees show that the 14 species are separate and valid species and 4 species of Parasesarma and 3 species of Perisesarma are not 2 independent clades but clustered into 1 clade. Chiromantes dehaani and Sesarmops sinensis are clustered into one clade and then clustered with C. haematocheir into another clade, inconsistent with their morphological classification. The complicated molecular phylogenetic relationship among the species of Sesarmidae suggests that the species in the family of Sesarmidae might have polyphyletic origin, and their interspecific relationship or intergenus relationship need to be clarified.

, correspAuthors=Mingliu Yang, authorNote=null, correspAuthorsNote=null, copyrightStatement=Haiyang Xuebao, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Yan Xu, Hongping Pan, Bing Yan, Tingwei Gao, Bin Wu, Mingliu Yang), CN=ArticleExt(id=1246833314250772544, articleId=1246833313126700032, tenantId=1146029695717560320, journalId=1149651085930835976, language=CN, title=基于线粒体COI和16S rRNA基因的中国沿海相手蟹系统发育研究, columnId=1243954927517679901, journalTitle=海洋学报, columnName=海洋生物, runingTitle=null, highlight=null, articleAbstract=

相手蟹科的诸多种类因其形态极其相似成为方蟹总科分类中疑问较多的一个类群。通过对中国沿海相手蟹线粒体COI和16S rRNA基因序列进行分子系统发育分析，结果表明14种相手蟹COI和16S rRNA基因序列之间差异分别为5.7%～14.5%和1.5%～12.1%，均达到了种间差异水平。构建的系统发育树显示，14种相手蟹分别为独立有效物种，但分属于拟相手蟹属和近相手蟹属的4种拟相手蟹和3种近相手蟹，没有分别形成2个独立的支系，而是混合聚成一大支系。而属于螳臂相手蟹属的无齿螳臂相手蟹则首先与属于中相手蟹属的中华中相手蟹聚成一支，再与红螯螳臂相手蟹聚为一大支，表现出与形态分类的不一致。错综复杂的分子系统关系预示着相手蟹类为多系起源，也表明它们之间的种间关系乃至于属间关系尚有诸多问题有待进一步厘定。

, correspAuthors=杨明柳, authorNote=null, correspAuthorsNote=
*杨明柳，助理研究员，主要从事海洋生物研究，E-mail：
, copyrightStatement=版权所有©《海洋学报》编辑部 2023
徐岩，潘红平，阎冰，等. 基于线粒体COI和16S rRNA基因的中国沿海相手蟹系统发育研究[J]. 海洋学报，2019，41(8)：63–71，doi:10.3969/j.issn.0253−4193.2019.08.006Xu Yan，Pan Hongping，Yan Bing, et al. Molecular phylogeny of the sesarmid crab based on the partial sequences of mitochondrial COI and 16S rRNA genes from the coast of China[J]. Haiyang Xuebao，2019, 41(8)：63–71，doi:10.3969/j.issn.0253−4193.2019.08.006
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徐岩（1992—），男，山东省日照市人，主要从事动物资源保护研究。E-mail：

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徐岩（1992—），男，山东省日照市人，主要从事动物资源保护研究。E-mail：

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U. arcuata is used as the outgroup

The value of BI/ML is marked on the corresponding branch. If the support rate is less than 50%, it will not be displayed. "–" means that the branch does not exist

, figureFileSmall=TGEpBOEVvDnFfuZ4UPlMtg==, figureFileBig=yBW6PRNFeHdtgbq/z4F+Jg==, tableContent=null), ArticleFig(id=1254506142332285733, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=CN, label=图1, caption=基于相手蟹及外群弧边招潮COI基因构建的贝叶斯分析方法（BI）和最大似然法（ML）系统树

BI/ML的值分别标在相应的分支上，值支持率小于50%的不显示，“–”表示分支不存在

, figureFileSmall=TGEpBOEVvDnFfuZ4UPlMtg==, figureFileBig=yBW6PRNFeHdtgbq/z4F+Jg==, tableContent=null), ArticleFig(id=1254506142483280679, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=EN, label=Fig. 2, caption=Inferred phylogenetic relationships based on nucleotide sequence of mitochondrial 16S rRNA gene of Sesarmidae species using Bayesian (BI) and maximum likelihood method (ML) analyses. U. arcuata is used as the outgroup

The value of BI/ML is marked on the corresponding branch. If the support rate is less than 50%, it will not be displayed. "–" means that the branch does not exist

, figureFileSmall=aMTbVSaF9mdpq/weu6jf3Q==, figureFileBig=PBk4c3NQl+IVY1UT+g94Cg==, tableContent=null), ArticleFig(id=1254506142751716138, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=CN, label=图2, caption=基于相手蟹及外群弧边招潮16S rRNA基因构建的贝叶斯分析方法（BI）和最大似然法（ML）系统树

BI/ML的值分别标在相应的分支上，值支持率小于50%的不显示，“–”表示分支不存在

, figureFileSmall=aMTbVSaF9mdpq/weu6jf3Q==, figureFileBig=PBk4c3NQl+IVY1UT+g94Cg==, tableContent=null), ArticleFig(id=1254506142965625646, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=EN, label=Table 1, caption=

The information of 14 Sesarmidae species and outgroup Uca arcuata

, figureFileSmall=null, figureFileBig=null, tableContent=
代码 种类 采集地点 样本量/只 采样时间
Pa 近亲拟相手蟹 Parasesarma affine 北海高德镇草头村 21°33′38.7″N，109°09′21.8″E 3 2012年7月
Pp 斑点拟相手蟹 Parasesarma pictum 汕头澳头苏埃湾红树林 23°19′29.3″N，116°42′58.3″E 1 2013年6月
Pt 三栉拟相手蟹 Parasesarma tripectinis 雷州企水海角村 20°48′13.2″N，109°44′9.7″E 2 2012年10月
Pu Parasesarma ungulatum 海南三亚青梅港 18°13′27.3″N，109°37′7.0″E 9 2012年10月
Pb 双齿近相手蟹 Perisesarma bidens 北海高德镇草头村 21°33′38.7″N，109°09′21.8″E 3 2012年7月
Pe Perisesarma eumolpe 海南洋浦 19°46′8.8″N，109°15′15.0″E 1 2012年10月
Pm 米埔近相手蟹 Perisesarma maipoensis 海南洋浦 19°46′8.8″N，109°15′15.0″E 3 2012年10月
Ev 泡粒上相手蟹 Episesarma versicolor 海南东寨港 19°58′34.2″N，110°33′34.4″E 3 2012年10月
Nm 小相手蟹 Nanosesarma minutum 汕头澳头苏埃湾红树林 23°19′29.3″N，116°42′58.3″E 2 2013年6月
Ss 中华中相手蟹 Sesarmops sinensis 深圳福田红树林 22°31′13.6″N，114°01′32.9″E 3 2013年6月
Cd 无齿螳臂相手蟹 Chiromantes dehaani 珠海淇澳红树林 22°23′45.4″N，113°38′37.8″E 1 2013年6月
Ch 红螯螳臂相手蟹 Chiromantes haematocheir 山东日照两城东河南村 35°33′34.48″N，119°37′51.0″E 2 2004年6月
Sg 吉氏胀蟹 Sarmatium germaini 海南东寨港 19°58′34.2″N，110°33′34.4″E 6 2012年10月
Cs 中华泥毛蟹 Clistocoeloma sinense 龙海市紫泥镇金定村 24°28′6.4″N，117°54′19.5″E 1 2012年9月
Ua 弧边招潮 Uca arcuata 阳江市阳西县程村镇 21°46′12.4″N，111°44′49.8″E 1 2013年6月
), ArticleFig(id=1254506143313752881, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=CN, label=表1, caption=

测序用14种相手蟹及外群弧边招潮取样信息

, figureFileSmall=null, figureFileBig=null, tableContent=
代码 种类 采集地点 样本量/只 采样时间
Pa 近亲拟相手蟹 Parasesarma affine 北海高德镇草头村 21°33′38.7″N，109°09′21.8″E 3 2012年7月
Pp 斑点拟相手蟹 Parasesarma pictum 汕头澳头苏埃湾红树林 23°19′29.3″N，116°42′58.3″E 1 2013年6月
Pt 三栉拟相手蟹 Parasesarma tripectinis 雷州企水海角村 20°48′13.2″N，109°44′9.7″E 2 2012年10月
Pu Parasesarma ungulatum 海南三亚青梅港 18°13′27.3″N，109°37′7.0″E 9 2012年10月
Pb 双齿近相手蟹 Perisesarma bidens 北海高德镇草头村 21°33′38.7″N，109°09′21.8″E 3 2012年7月
Pe Perisesarma eumolpe 海南洋浦 19°46′8.8″N，109°15′15.0″E 1 2012年10月
Pm 米埔近相手蟹 Perisesarma maipoensis 海南洋浦 19°46′8.8″N，109°15′15.0″E 3 2012年10月
Ev 泡粒上相手蟹 Episesarma versicolor 海南东寨港 19°58′34.2″N，110°33′34.4″E 3 2012年10月
Nm 小相手蟹 Nanosesarma minutum 汕头澳头苏埃湾红树林 23°19′29.3″N，116°42′58.3″E 2 2013年6月
Ss 中华中相手蟹 Sesarmops sinensis 深圳福田红树林 22°31′13.6″N，114°01′32.9″E 3 2013年6月
Cd 无齿螳臂相手蟹 Chiromantes dehaani 珠海淇澳红树林 22°23′45.4″N，113°38′37.8″E 1 2013年6月
Ch 红螯螳臂相手蟹 Chiromantes haematocheir 山东日照两城东河南村 35°33′34.48″N，119°37′51.0″E 2 2004年6月
Sg 吉氏胀蟹 Sarmatium germaini 海南东寨港 19°58′34.2″N，110°33′34.4″E 6 2012年10月
Cs 中华泥毛蟹 Clistocoeloma sinense 龙海市紫泥镇金定村 24°28′6.4″N，117°54′19.5″E 1 2012年9月
Ua 弧边招潮 Uca arcuata 阳江市阳西县程村镇 21°46′12.4″N，111°44′49.8″E 1 2013年6月
), ArticleFig(id=1254506143619937076, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=EN, label=Table 2, caption=

The COI gene and 16S rRNA gene information of 14 Sesarmidae species and outgroup U. arcuata

, figureFileSmall=null, figureFileBig=null, tableContent=
种类 单倍型 （COI/16S rRNA）代码 碱基含量/% （COI/16S rRNA） 序列长度/bp （COI/16S rRNA） 基因库登录号 （COI/16S rRNA）
A T G C A+T
近亲拟相手蟹 P. affine Pa/Pa 29.0/37.1 36.9/39.2 17.1/15.0 17.0/8.7 65.9/76.3 631/507 MK611589/KU052882[5]
斑点拟相手蟹 P. pictum Pp/Pp 29.5/37.2 38.0/38.7 17.4/15.7 15.1/8.4 67.5/75.9 631/511 MK611594/KU052884[5]
三栉拟相手蟹 P. tripectinis Pt1/Pt 30.1/37.3 38.2/38.1 16.3/15.8 15.4/8.8 68.3/75.4 631/520 MK611595/KU052885[5]
Pt2 30.0 38.2 16.5 15.3 68.2 MK611596
P. ungulatum Pu1/Pu 29.2/36.6 38.4/38.9 16.5/15.8 15.9/8.7 67.6/75.5 631/517 MK611597/KU052886[5]
Pu2 29.2 38.5 16.5 15.8 67.7 MK611598
Pu3 29.0 38.4 16.6 16.0 67.4 MK611599
Pu4 29.2 38.2 16.5 16.1 67.4 MK611600
双齿近相手蟹 P. bidens Pb1/Pb 28.8/37.4 38.2/39.0 17.3/15.3 15.7/8.3 67.0/76.4 631/497 MK611590/MK967687
Pb2 28.8 38.4 17.1 15.7 67.2 MK611591
P. eumolpe Pe/Pe 28.5/36.8 39.0/38.4 17.3/16.2 15.2/8.6 67.5/75.2 631/513 MK611592/MK967688
米埔近相手蟹 P. maipoensis Pm/Pm 28.7/37.4 37.9/38.9 16.6/14.6 16.8/9.1 66.6/76.3 631/519 MK611593/MK967689
泡粒上相手蟹 E. versicolor Ev/Ev 28.7/35.2 38.8/38.9 16.6/16.4 15.9/9.5 67.5/74.1 631/512 MK611586/MK967685
小相手蟹 N. minutum Nm1/Nm 30.9/37.9 37.2/39.2 16.2/14.8 15.7/8.1 68.1/77.1 631/494 MK611587/MK967686
Nm2 30.9 37.2 16.2 15.7 68.1 MK611588
中华中相手蟹 S. sinensis Ss1/Ss 31.4/36.6 37.3/38.6 16.2/16.1 15.1/8.7 68.7/75.2 631/508 MK611605/MK967691
Ss2 31.4 37.2 16.2 15.2 68.6 MK611606
无齿螳臂相手蟹 C. dehaani Cd1/Cd 30.3/36.6 37.1/38.9 16.9/16.0 15.7/8.5 67.4/75.5 631/506 MK611581/MK967682
Cd2 30.4 37.1 16.8 15.7 67.5 MK611582
Cd3 30.4 36.8 16.8 16.0 67.2 MK611583
红螯螳臂相手蟹 C. haematocheir Ch/Ch 30.0/37.2 38.2/38.8 16.3/15.5 15.5/8.5 68.2/76.0 631/505 MK611584/MK967683
吉氏胀蟹 S. germaini Sg1/Sg 29.8/35.1 38.8/39.4 15.5/16.7 15.9/8.8 68.6/74.5 631/513 MK611601/MK967690
Sg2 30.0 38.7 15.5 15.8 68.7 MK611602
Sg3 29.5 37.2 16.3 17.0 66.7 MK611603
Sg4 29.5 37.4 16.3 16.8 66.9 MK611604
中华泥毛蟹 C. sinense Cs/Cs 29.3/36.4 37.1/39.0 16.6/15.2 17.0/9.4 66.4/75.4 631/500 MK611585/MK967684
平均 29.7/36.8 37.9/38.9 16.5/15.6 15.9/8.7 67.6/75.7 631/509
弧边招潮 U. arcuata Ua/Ua 29.2/34.7 32.2/37.1 17.1/18.5 21.5/9.7 61.4/71.8 631/493 MK611607/MK967692
), ArticleFig(id=1254506143972258618, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=CN, label=表2, caption=

14种相手蟹和外群弧边招潮COI和16S rRNA基因信息

, figureFileSmall=null, figureFileBig=null, tableContent=
种类 单倍型 （COI/16S rRNA）代码 碱基含量/% （COI/16S rRNA） 序列长度/bp （COI/16S rRNA） 基因库登录号 （COI/16S rRNA）
A T G C A+T
近亲拟相手蟹 P. affine Pa/Pa 29.0/37.1 36.9/39.2 17.1/15.0 17.0/8.7 65.9/76.3 631/507 MK611589/KU052882[5]
斑点拟相手蟹 P. pictum Pp/Pp 29.5/37.2 38.0/38.7 17.4/15.7 15.1/8.4 67.5/75.9 631/511 MK611594/KU052884[5]
三栉拟相手蟹 P. tripectinis Pt1/Pt 30.1/37.3 38.2/38.1 16.3/15.8 15.4/8.8 68.3/75.4 631/520 MK611595/KU052885[5]
Pt2 30.0 38.2 16.5 15.3 68.2 MK611596
P. ungulatum Pu1/Pu 29.2/36.6 38.4/38.9 16.5/15.8 15.9/8.7 67.6/75.5 631/517 MK611597/KU052886[5]
Pu2 29.2 38.5 16.5 15.8 67.7 MK611598
Pu3 29.0 38.4 16.6 16.0 67.4 MK611599
Pu4 29.2 38.2 16.5 16.1 67.4 MK611600
双齿近相手蟹 P. bidens Pb1/Pb 28.8/37.4 38.2/39.0 17.3/15.3 15.7/8.3 67.0/76.4 631/497 MK611590/MK967687
Pb2 28.8 38.4 17.1 15.7 67.2 MK611591
P. eumolpe Pe/Pe 28.5/36.8 39.0/38.4 17.3/16.2 15.2/8.6 67.5/75.2 631/513 MK611592/MK967688
米埔近相手蟹 P. maipoensis Pm/Pm 28.7/37.4 37.9/38.9 16.6/14.6 16.8/9.1 66.6/76.3 631/519 MK611593/MK967689
泡粒上相手蟹 E. versicolor Ev/Ev 28.7/35.2 38.8/38.9 16.6/16.4 15.9/9.5 67.5/74.1 631/512 MK611586/MK967685
小相手蟹 N. minutum Nm1/Nm 30.9/37.9 37.2/39.2 16.2/14.8 15.7/8.1 68.1/77.1 631/494 MK611587/MK967686
Nm2 30.9 37.2 16.2 15.7 68.1 MK611588
中华中相手蟹 S. sinensis Ss1/Ss 31.4/36.6 37.3/38.6 16.2/16.1 15.1/8.7 68.7/75.2 631/508 MK611605/MK967691
Ss2 31.4 37.2 16.2 15.2 68.6 MK611606
无齿螳臂相手蟹 C. dehaani Cd1/Cd 30.3/36.6 37.1/38.9 16.9/16.0 15.7/8.5 67.4/75.5 631/506 MK611581/MK967682
Cd2 30.4 37.1 16.8 15.7 67.5 MK611582
Cd3 30.4 36.8 16.8 16.0 67.2 MK611583
红螯螳臂相手蟹 C. haematocheir Ch/Ch 30.0/37.2 38.2/38.8 16.3/15.5 15.5/8.5 68.2/76.0 631/505 MK611584/MK967683
吉氏胀蟹 S. germaini Sg1/Sg 29.8/35.1 38.8/39.4 15.5/16.7 15.9/8.8 68.6/74.5 631/513 MK611601/MK967690
Sg2 30.0 38.7 15.5 15.8 68.7 MK611602
Sg3 29.5 37.2 16.3 17.0 66.7 MK611603
Sg4 29.5 37.4 16.3 16.8 66.9 MK611604
中华泥毛蟹 C. sinense Cs/Cs 29.3/36.4 37.1/39.0 16.6/15.2 17.0/9.4 66.4/75.4 631/500 MK611585/MK967684
平均 29.7/36.8 37.9/38.9 16.5/15.6 15.9/8.7 67.6/75.7 631/509
弧边招潮 U. arcuata Ua/Ua 29.2/34.7 32.2/37.1 17.1/18.5 21.5/9.7 61.4/71.8 631/493 MK611607/MK967692
), ArticleFig(id=1254506144366523197, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=EN, label=Table 3, caption=

Interspecific and intraspecific genetic distances of COI gene among 14 Sesarmidae species

, figureFileSmall=null, figureFileBig=null, tableContent=
Cd1 Cd2 Cd3 Ch Cs Ev Nm1 Nm2 Pa Pb1 Pb2 Pe Pm Pp Pt1 Pt2 Pu1 Pu2 Pu3 Pu4 Sg1 Sg2 Sg3 Sg4 Ss1 Ss2
Cd1
Cd2 0.002
Cd3 0.005 0.003
Ch 0.081 0.079 0.083
Cs 0.121 0.119 0.123 0.106
Ev 0.113 0.111 0.111 0.115 0.102
Nm1 0.113 0.111 0.115 0.109 0.118 0.137
Nm2 0.115 0.113 0.117 0.111 0.117 0.137 0.003
Pa 0.143 0.141 0.145 0.122 0.134 0.132 0.127 0.127
Pb1 0.124 0.122 0.126 0.122 0.134 0.135 0.123 0.123 0.094
Pb2 0.124 0.122 0.126 0.122 0.132 0.134 0.123 0.123 0.094 0.002
Pe 0.130 0.128 0.132 0.111 0.132 0.134 0.102 0.102 0.104 0.073 0.072
Pm 0.119 0.117 0.117 0.122 0.128 0.107 0.126 0.128 0.114 0.117 0.117 0.097
Pp 0.119 0.117 0.121 0.110 0.123 0.113 0.126 0.125 0.113 0.108 0.108 0.106 0.108
Pt1 0.109 0.107 0.111 0.098 0.117 0.105 0.119 0.117 0.110 0.100 0.100 0.102 0.098 0.057
Pt2 0.107 0.105 0.109 0.100 0.119 0.107 0.117 0.115 0.111 0.102 0.102 0.104 0.100 0.058 0.002
Pu1 0.104 0.102 0.106 0.106 0.122 0.120 0.104 0.104 0.099 0.086 0.086 0.090 0.101 0.100 0.097 0.099
Pu2 0.102 0.100 0.104 0.107 0.120 0.119 0.106 0.106 0.098 0.084 0.084 0.088 0.099 0.099 0.095 0.097 0.002
Pu3 0.100 0.098 0.102 0.106 0.122 0.120 0.106 0.106 0.098 0.082 0.082 0.090 0.101 0.100 0.097 0.099 0.005 0.006
Pu4 0.106 0.104 0.107 0.107 0.124 0.122 0.106 0.106 0.099 0.088 0.088 0.092 0.101 0.102 0.099 0.100 0.002 0.003 0.006
Sg1 0.106 0.104 0.104 0.082 0.113 0.115 0.113 0.113 0.132 0.128 0.128 0.124 0.117 0.104 0.082 0.084 0.107 0.106 0.107 0.109
Sg2 0.108 0.106 0.106 0.084 0.115 0.117 0.115 0.115 0.134 0.130 0.130 0.126 0.119 0.106 0.084 0.086 0.109 0.107 0.109 0.111 0.002
Sg3 0.111 0.110 0.110 0.090 0.121 0.123 0.125 0.125 0.142 0.140 0.139 0.132 0.130 0.110 0.096 0.098 0.109 0.108 0.109 0.111 0.029 0.031
Sg4 0.113 0.111 0.111 0.088 0.123 0.125 0.123 0.123 0.140 0.138 0.137 0.130 0.132 0.111 0.095 0.096 0.111 0.109 0.111 0.113 0.028 0.029 0.002
Ss1 0.065 0.064 0.067 0.079 0.123 0.113 0.102 0.104 0.134 0.120 0.120 0.115 0.123 0.111 0.102 0.104 0.107 0.106 0.104 0.109 0.095 0.097 0.108 0.106
Ss2 0.067 0.065 0.069 0.081 0.121 0.111 0.104 0.105 0.132 0.118 0.118 0.117 0.124 0.113 0.104 0.105 0.106 0.104 0.102 0.107 0.093 0.095 0.106 0.104 0.002
), ArticleFig(id=1254506144668513091, tenantId=1146029695717560320, journalId=1149651085930835976, articleId=1246833313126700032, language=CN, label=表3, caption=

14种相手蟹COI基因种间和种内的遗传距离

, figureFileSmall=null, figureFileBig=null, tableContent=
Cd1 Cd2 Cd3 Ch Cs Ev Nm1 Nm2 Pa Pb1 Pb2 Pe Pm Pp Pt1 Pt2 Pu1 Pu2 Pu3 Pu4 Sg1 Sg2 Sg3 Sg4 Ss1 Ss2
Cd1
Cd2 0.002
Cd3 0.005 0.003
Ch 0.081 0.079 0.083
Cs 0.121 0.119 0.123 0.106
Ev 0.113 0.111 0.111 0.115 0.102
Nm1 0.113 0.111 0.115 0.109 0.118 0.137
Nm2 0.115 0.113 0.117 0.111 0.117 0.137 0.003
Pa 0.143 0.141 0.145 0.122 0.134 0.132 0.127 0.127
Pb1 0.124 0.122 0.126 0.122 0.134 0.135 0.123 0.123 0.094
Pb2 0.124 0.122 0.126 0.122 0.132 0.134 0.123 0.123 0.094 0.002
Pe 0.130 0.128 0.132 0.111 0.132 0.134 0.102 0.102 0.104 0.073 0.072
Pm 0.119 0.117 0.117 0.122 0.128 0.107 0.126 0.128 0.114 0.117 0.117 0.097
Pp 0.119 0.117 0.121 0.110 0.123 0.113 0.126 0.125 0.113 0.108 0.108 0.106 0.108
Pt1 0.109 0.107 0.111 0.098 0.117 0.105 0.119 0.117 0.110 0.100 0.100 0.102 0.098 0.057
Pt2 0.107 0.105 0.109 0.100 0.119 0.107 0.117 0.115 0.111 0.102 0.102 0.104 0.100 0.058 0.002
Pu1 0.104 0.102 0.106 0.106 0.122 0.120 0.104 0.104 0.099 0.086 0.086 0.090 0.101 0.100 0.097 0.099
Pu2 0.102 0.100 0.104 0.107 0.120 0.119 0.106 0.106 0.098 0.084 0.084 0.088 0.099 0.099 0.095 0.097 0.002
Pu3 0.100 0.098 0.102 0.106 0.122 0.120 0.106 0.106 0.098 0.082 0.082 0.090 0.101 0.100 0.097 0.099 0.005 0.006
Pu4 0.106 0.104 0.107 0.107 0.124 0.122 0.106 0.106 0.099 0.088 0.088 0.092 0.101 0.102 0.099 0.100 0.002 0.003 0.006
Sg1 0.106 0.104 0.104 0.082 0.113 0.115 0.113 0.113 0.132 0.128 0.128 0.124 0.117 0.104 0.082 0.084 0.107 0.106 0.107 0.109
Sg2 0.108 0.106 0.106 0.084 0.115 0.117 0.115 0.115 0.134 0.130 0.130 0.126 0.119 0.106 0.084 0.086 0.109 0.107 0.109 0.111 0.002
Sg3 0.111 0.110 0.110 0.090 0.121 0.123 0.125 0.125 0.142 0.140 0.139 0.132 0.130 0.110 0.096 0.098 0.109 0.108 0.109 0.111 0.029 0.031
Sg4 0.113 0.111 0.111 0.088 0.123 0.125 0.123 0.123 0.140 0.138 0.137 0.130 0.132 0.111 0.095 0.096 0.111 0.109 0.111 0.113 0.028 0.029 0.002
Ss1 0.065 0.064 0.067 0.079 0.123 0.113 0.102 0.104 0.134 0.120 0.120 0.115 0.123 0.111 0.102 0.104 0.107 0.106 0.104 0.109 0.095 0.097 0.108 0.106
Ss2 0.067 0.065 0.069 0.081 0.121 0.111 0.104 0.105 0.132 0.118 0.118 0.117 0.124 0.113 0.104 0.105 0.106 0.104 0.102 0.107 0.093 0.095 0.106 0.104 0.002
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基于线粒体COI和16S rRNA基因的中国沿海相手蟹系统发育研究
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徐岩 1, 2 , 潘红平 1 , 阎冰 2 , 高霆炜 2 , 吴斌 2 , 杨明柳 2, *
海洋学报 | 海洋生物 2019,41(8): 63-71
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海洋学报 | 海洋生物 2019, 41(8): 63-71
基于线粒体COI和16S rRNA基因的中国沿海相手蟹系统发育研究
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徐岩1, 2 , 潘红平1, 阎冰2, 高霆炜2, 吴斌2, 杨明柳2, *
作者信息
  • 1 广西大学 动物科学技术学院，广西 南宁 530004
  • 2 广西科学院 广西红树林研究中心 广西红树林保护与利用重点实验室，广西 北海 536000

    • 徐岩（1992—），男，山东省日照市人，主要从事动物资源保护研究。E-mail：

通讯作者:

*杨明柳，助理研究员，主要从事海洋生物研究，E-mail：
Molecular phylogeny of the sesarmid crab based on the partial sequences of mitochondrial COI and 16S rRNA genes from the coast of China
Yan Xu1, 2 , Hongping Pan1, Bing Yan2, Tingwei Gao2, Bin Wu2, Mingliu Yang2, *
Affiliations
  • 1 College of Animal Science and Technology, Guangxi University, Nanning 530004, China
  • 2 Guangxi Key Lab of Mangrove Conservation and Utilization, Guangxi Mangrove Research Center, Guangxi Academy of Sciences, Beihai 536000, China
出版时间: 2019-08-25 doi: 10.3969/j.issn.0253-4193.2019.08.006
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摘要
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相手蟹科的诸多种类因其形态极其相似成为方蟹总科分类中疑问较多的一个类群。通过对中国沿海相手蟹线粒体COI和16S rRNA基因序列进行分子系统发育分析，结果表明14种相手蟹COI和16S rRNA基因序列之间差异分别为5.7%～14.5%和1.5%～12.1%，均达到了种间差异水平。构建的系统发育树显示，14种相手蟹分别为独立有效物种，但分属于拟相手蟹属和近相手蟹属的4种拟相手蟹和3种近相手蟹，没有分别形成2个独立的支系，而是混合聚成一大支系。而属于螳臂相手蟹属的无齿螳臂相手蟹则首先与属于中相手蟹属的中华中相手蟹聚成一支，再与红螯螳臂相手蟹聚为一大支，表现出与形态分类的不一致。错综复杂的分子系统关系预示着相手蟹类为多系起源，也表明它们之间的种间关系乃至于属间关系尚有诸多问题有待进一步厘定。

相手蟹  /  线粒体DNA  /  COI基因  /  16S rRNA基因  /  系统发育

The species in the family of Sesarmidae are among the species mostly difficult to be differentiated taxonomically in the Superfamily Grapsoidea because of their morphological similarity. The analysis of partial sequences of mitochondrial COI gene and 16S rRNA gene of 14 species in 8 genera of Sesarmidae for their molecular phylogenetics reveals that the percentages of sequence divergences of 14 species of Sesarmidae are from 5.7% to 14.5% and from 1.5% to 12.1%, both indicating that the species difference are significant. The phylogenetic trees show that the 14 species are separate and valid species and 4 species of Parasesarma and 3 species of Perisesarma are not 2 independent clades but clustered into 1 clade. Chiromantes dehaani and Sesarmops sinensis are clustered into one clade and then clustered with C. haematocheir into another clade, inconsistent with their morphological classification. The complicated molecular phylogenetic relationship among the species of Sesarmidae suggests that the species in the family of Sesarmidae might have polyphyletic origin, and their interspecific relationship or intergenus relationship need to be clarified.

Sesarmidae  /  mitochondrial DNA  /  COI gene  /  16S rRNA gene  /  phylogeny
徐岩, 潘红平, 阎冰, 高霆炜, 吴斌, 杨明柳. 基于线粒体COI和16S rRNA基因的中国沿海相手蟹系统发育研究. 海洋学报, 2019 , 41 (8) : 63 -71 . DOI: 10.3969/j.issn.0253-4193.2019.08.006
Yan Xu, Hongping Pan, Bing Yan, Tingwei Gao, Bin Wu, Mingliu Yang. Molecular phylogeny of the sesarmid crab based on the partial sequences of mitochondrial COI and 16S rRNA genes from the coast of China[J]. Haiyang Xuebao, 2019 , 41 (8) : 63 -71 . DOI: 10.3969/j.issn.0253-4193.2019.08.006
正文
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1 引言
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相手蟹隶属于方蟹总科（Grapsoidea），相手蟹科（Sesarmidae），种类繁多，多数生活于红树林生态系统中[1-2]。中国记录的相手蟹有37种，隶属于16个属[3]；杨明柳等[4-5]、梁晓莉[6]分别于2014年、2016年、2017年报道了分布于北部湾、海南三亚和长江河口的3种相手蟹：即Perisesarma eumolpe、Parasesarma ungulatum和隐秘螳臂相手蟹（Chiromantes neglectum）。
由于线粒体DNA具有分子量小、结构简单、进化速度快、母性遗传等特点[7]，已成为研究动物群体遗传分化、分类及系统发育的理想分子标记。在线粒体基因组中，线粒体细胞色素氧化酶I亚基（COI）基因属于变异性较大的区域，适合于种间区分和种内的分子系统学研究；16S rRNA基因相对保守，适合于远缘物种研究[8]。已有较多利用COI和16S rRNA基因序列来进行相手蟹等蟹类分子系统学的研究[9-13]。
尽管已有一些针对相手蟹的分类学研究[14-18]，但因早期研究者对一些新种或新记录的描述不够详细，或者因一些模式标本的缺失，致使出现同物异名或异物同名等分类混乱现象，加之对一些种类缺少了解，因此，相手蟹科的分类尚存在较多疑问。
首先对2004年6月至2013年6月采集的中国沿海相手蟹进行形态分类；在此基础上，运用线粒体DNA的COI和16S rRNA基因片段对其进行种间和种内序列分析并构建系统发育树，探讨其属间、种间和种内的分子系统关系，以期利用分子生物学手段，解决相手蟹科蟹类存在的分类疑问，为其分属及种类厘定提供基础资料。
2 材料与方法
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2.1 材料
研究用的14种相手蟹和外群弧边招潮的相关信息见表1，所有样品均浸泡于95%酒精中保存，所有样品均保存在广西红树林保护与利用重点实验室。
2.2 方法
取约20 mg步足肌肉用基因组DNA提取试剂盒（OMEGA公司）进行提取，提取的DNA保存于–20℃。分别以COIL1490：5′-GGTCAACAAATCATAAAGATATTGG-3′，COIH2198：5′-TAAACTTCAGGGTGACCAAAAAATCA-3′和16Sar：5′-CGCCTGTTTAACAAAAACAT-3′，16Sbr：5′-CCGGTCTGAACTCAGATCA CGT-3′为引物扩增COI[19]和16S rRNA[20]基因片段。PCR扩增反应体积为50 µL，其中Taq10 2× MasterMix (北京奥赛博公司)19 µL，模板DNA 2 µL，引物各2 µL，Milli-Q-Water补足至50 µL。
COI基因片段PCR循环参数为：94℃预变性2 min后，94℃变性45 s，47℃退火50 s，72℃延伸1 min，循环35次，然后在72℃延伸7 min；16S rRNA基因片段反应条件为：94℃预变性1.5 min后，94℃变性30 s，49℃退火45 s，72℃延伸1 min，39个循环，最终72℃延伸5 min。产物经1%琼脂糖凝胶电泳，溴化乙锭（EB）染色检测，样品送至广州英潍捷基测序公司进行纯化，并用ABI 3730XL测序仪进行正反链双向测序。
序列由DNAstar软件包的SeqMan、MegAlign和EditSeq程序进行编辑、校对和排序；DNAsp5.0检测多态位点数、简约信息位点数。采用Mega4.0软件计算遗传距离。使用最大似然法和贝叶斯两种分析方法重建系统发育树：利用RAxML v 8.0.0软件对矩阵进行最大似然法分析建树。用100次重复的自展分析来评估树图拓扑结构的可靠性。所有运算都是在CIPRES网站完成[21]。通过MrModeltest筛选出COI和16S rRNA的最佳替换模型分别为GTR+I+G和GTR+I，利用MrBayes v3.0b4软件包[22]进行贝叶斯法分析建树，采用马尔科夫链的蒙特卡洛（Markov Chain Monte Carlo）方法，以随机树为起始树，运行1 000 000，每1 000代取样1次，每次运算开始的250个样本作为老化样本舍弃，最后获得各分支的后验概率。
3 结果
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3.1 基因序列及变异
对14种相手蟹40个样本及外群1个样本共计41个个体的线粒体COI基因片段进行PCR扩增和序列测定，所有序列长度均为631 bp，其中三栉拟相手蟹、P.ungulatum、双齿近相手蟹、小相手蟹、中华中相手蟹、无齿螳臂相手蟹和吉氏胀蟹分别出现了2、4、2、2、2、3和4个单倍型（表2）。
对10种相手蟹25个样本及外群1个样本共计26个个体的线粒体16S rRNA基因片段进行PCR扩增和序列测定，序列长度为494～519 bp，每种相手蟹只有1个单倍型（表2）。
由表2可见，14种相手蟹的A、T、G、C含量只有略微的差异，表现出与其他蟹类COI和16S rRNA基因相同的特点[10, 23-24]，即A+T含量（平均67.6%/75.7%）明显高于G+C含量。本研究14种相手蟹的COI和16S rRNA基因序列比对分别获得631 bp（无插入/缺失位点）和509 bp（含插入/缺失位点）的同源序列，二者共分别检测到196和107个变异位点，其中简约信息位点分别为165个和67个。
3.2 种间和种内遗传差异
利用木村（Kimura）–双参数法计算得到了14种相手蟹种间和种内的遗传差异，COI基因的遗传距离如表3所示，相手蟹种类之间序列差异为5.7%～14.5%，种内之间序列差异为0.2%～3.1%。而相手蟹16S rRNA基因种类之间的序列差异为1.5%～12.1%。
3.3 分子系统发育
以弧边招潮（Ua）为外群，基于14种相手蟹和C. neglectum(代码：Cn；基因库登录号：KX156954)[25]的COI和16S rRNA基因片段序列，构建BI树和ML树，它们的拓扑结构基本一致（图1，图2），14种相手蟹（含C. neglectum）可分为两大分支，其中无齿螳臂相手蟹、C. neglectum、中华中相手蟹和红螯螳臂相手蟹聚为一大支；另一大支中4种拟相手蟹和3种近相手蟹聚成一支，泡粒上相手蟹和中华泥毛蟹聚成一支，吉氏胀蟹单独成支。
4 讨论
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4.1 种内分化分析
在14种相手蟹中，三栉拟相手蟹、P. ungulatum、双齿近相手蟹、小相手蟹、中华中相手蟹和无齿螳臂相手蟹种内COI基因单倍型之间序列差异为0.2%～0.6%（表3），这与厚蟹种内0.15%～1.22%的变异度一致[24]；但吉氏胀蟹的4个单倍型分化出来了Sg1/Sg2和Sg3/Sg4两个组群，两个组群内的遗传变异度均为0.2%，但两个组群间的遗传变异度则达到了2.8%～3.1%（表3），表明组群间出现了较大的遗传分化，这可能与滩涂型蟹类扩散能力强有关，但具体原因有待对不同群体进行更深入的研究来揭示。而10种相手蟹的16S rRNA基因片段，尽管有些种类含有多个样本，但也只有1个单倍型；表明该基因较前者相对保守，进化速率相对较慢。
4.2 种间关系分析
本研究的14种相手蟹COI基因序列之间差异为5.7%～14.5%，一般认为COI基因序列的变异速率是16S rRNA基因的2倍以上[9]，若依据16S rRNA基因的1.5%的种间差异标准[26]；这14种相手蟹的序列差异均达到了种间差异，分子数据支持这14种相手蟹分别作为独立物种；同基于16S rRNA针对拟相手蟹的研究结果[5]一样，亦支持P. ungulatum为有效种。相手蟹16S rRNA基因种类之间的序列差异为1.5%～12.1%，也支持其分别为独立物种。
值得注意的是，本研究无齿螳臂相手蟹（C. dehaani）与均产于上海长江口的C. neglectum [25]和无齿螳臂相手蟹(基因库登录号：MH593563)[27]COI/16S rRNA基因的631 bp/506 bp同源序列比对结果显示，二者之间的序列差异分别为0.32%/0.00%、1.42%/0.02%，都属于种内差异范围；即便是均产于上海长江口的无齿螳臂相手蟹[27]与C. neglectum [25] COI和16S rRNA基因的全序列比对结果显示，前者序列均为1 535 bp，序列差异仅为0.65%；而后者除2 bp插入/缺失外的1 338 bp同源序列，序列差异仅为0.30%，也均显示二者属于种内差异范围。系统发育树亦显示无齿螳臂相手蟹（Cd）和C. neglectum（Cn）以很高的置信度首先聚成一支（图1，图2）。
但是，梁晓莉[6]通过对长江口（上海）无齿螳臂相手蟹的形态和COI基因部分序列的研究，认为长江口的无齿螳臂相手蟹为错误鉴定，拟改名为隐秘螳臂相手蟹；需要注意的是，该研究中利用COI基因部分序列进行分子系统分析所用的无齿螳臂相手蟹（基因库登录号：AB457708）标本来自日本[28]，该标本498 bp的COI序列分别与无齿螳臂相手蟹[27]和C. neglectum [25] COI基因的全序列比对结果显示，498 bp的同源序列均只有2 bp（0.40%）的差异，表明三者属于种内差异范围；在梁晓莉[6]基于部分COI基因序列的分子系统研究中，长江口的无齿螳臂相手蟹和日本的无齿螳臂相手蟹之所以序列差异较大，形成两个独立分支，源于使用了非同源序列。
鉴于此，尽管长江口的无齿螳臂相手蟹[6]与日本的无齿螳臂相手蟹[29]在形态上存在一定的差异，但现有的分子数据表明，无论是均产于上海长江口的无齿螳臂相手蟹[27]和C. neglectum [25]，还是本研究来自珠海的无齿螳臂相手蟹和前二者，以及来自日本的无齿螳臂相手蟹[28]和前三者，其遗传差异均属于种内差异范围，揭示C. dehaani和C. neglectum为同一物种，不支持将长江口的无齿螳臂相手蟹改为隐秘螳臂相手蟹[6]。类似结果也存在于其他蟹类如天津厚蟹与侧足厚蟹的相关研究中[24]。
4.3 属间关系分析
本研究的14种相手蟹隶属于现行分类系统相手蟹科中的8个属[1, 3]，系统发育树的拓扑结构显示这14种相手蟹之间的系统关系大致与相手蟹科的现行分类系统相符，但也存在需要进一步阐释的问题（图1，图2）。
首先，4种拟相手蟹（Parasesarma）和3种近相手蟹（Perisesarma）分属于两个属的蟹类，却没有分别形成2个独立的支系，而是混合聚成了1个支系；4种拟相手蟹中，斑点拟相手蟹和三栉拟相手蟹聚成一支且置信度高，表明二者亲缘关系最近（二者COI序列的遗传差异为最小值的5.7%或5.8%），与另外2种拟相手蟹的亲缘关系较远，这与基于16S rRNA的研究结果[5]一致；3种近相手蟹中，双齿近相手蟹与P. eumolpe聚成1支且置信度较高，表明二者亲缘关系较近，但与米埔相手蟹的亲缘关系较远（图1，图2）。此类情况即多种拟相手蟹和近相手蟹紧密的聚类在一起，导致相关种属的重新调整和新属的建立[13]已见于国外的相手蟹类；基于此，中国的拟相手蟹属和近相手蟹属的蟹类是否需要合并为1个复合类群（Parasesarma/Perisesarma），或者2个属所属的种类是否需要重新厘定，或者是否需要建立新属，尚需对该两属更多种类在形态、分子等方面进行更深入全面的研究来确认。
此外，无齿螳臂相手蟹首先与中华中相手蟹聚成一支，表明二者的亲缘关系较近，然后再与红螯螳臂相手蟹聚为一大支（图1，图2）；这与现行分类系统中，中华中相手蟹属于中相手蟹属（Sesarmops），其他两种属于螳臂相手蟹属（Chiromantes）不一致，建议将此两个属合并（Chiromantes/Sesarmops）或者将中华中相手蟹移入螳臂相手蟹属或把无齿螳臂相手蟹移入中相手蟹属，但这有待于对中相手蟹属中其他种类进行更深入的相关研究才能厘定。
总之，14种相手蟹错综复杂的分子系统关系预示着相手蟹类为多系起源，但由于很多种类在形态上有很高的相似度，致使某些相手蟹种类之间的区分难度较大，且国内在其形态、分子方面的相关研究还很少，致使它们的种类、属间关系等存在诸多问题，尚需针对更多种类做更广泛更深入的研究。
基金
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  • 国家重点研发计划科技基础资源调查专项（2017FY100704）；广西自然科学基金项目（2017GXNSFBA198163）。
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2019年第41卷第8期
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doi: 10.3969/j.issn.0253-4193.2019.08.006
  • 接收时间：2019-04-01
  • 首发时间：2026-04-03
  • 出版时间：2019-08-25
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  • 收稿日期：2019-04-01
  • 修回日期：2019-06-13
基金
国家重点研发计划科技基础资源调查专项（2017FY100704）；广西自然科学基金项目（2017GXNSFBA198163）。
作者信息
    1 广西大学 动物科学技术学院，广西 南宁 530004
    2 广西科学院 广西红树林研究中心 广西红树林保护与利用重点实验室，广西 北海 536000

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*杨明柳，助理研究员，主要从事海洋生物研究，E-mail：
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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