燕尾藓属（Bryhnia）和异叶藓属（Kindbergia）基于分子和形态证据的分类学修订
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摘要
青藓科(Brachytheciaceae)是侧蒴藓类中种类较为丰富、分类学问题较多的类群之一。在青藓科概念趋于稳定后的二十一世纪,青藓科的主要问题则集中在科内形态相近的属间及属内种间关系。本研究选择青藓科中分类学问题较多的燕尾藓属(Bryhnia)和异叶藓属(Kindbergid),采用经典分类学和分子系统学相结合的方法,对两属植物进行了分类学修订。
经典分类学研究主要基于1000余份燕尾藓属和异叶藓属的标本,其中包括模式标本67份,隶属于39种(包括种下单位)。分子系统学研究选择以燕尾藓属和异叶藓属为核心的青藓科3亚科的14属、34种、98份样本作为内群,其中燕尾藓属7种、29份样本,异叶藓属4种、40份样本；蔓藓科(Meteoriaceae)的鞭枝悬藓(Barbella flagellifera)和东亚蔓藓(Meteorium atrovariegatum)为外群；选用1个核基因ITS和3个叶绿体基因rpl16, trnG, trnL-F,运用最大简约法(MP)和贝叶斯分析(BI)两种方法进行了分子系统学分析。主要研究结果如下：
1.燕尾藓属是一个多系类群。
(1)综合经典分类学和分子系统学研究结果,将燕尾藓属处理为青藓属的异名；将燕尾藓(Bryhnia novae-angliae)、短尖燕尾藓(B. hultenii)、 B. scabrida、毛尖燕尾藓(B. trichomitria)移入了青藓属；短枝燕尾藓(B. brachycladula)移入鼠尾藓属,并处理为羽枝鼠尾藓(Myuroclada longiramea)的异名；而Bryhnia graminicolor和密枝燕尾藓(B. serricuspis)分别被移入2个新建属,即小燕尾藓属(Bryhniastrum)和拟异叶藓属(Pseudokindbergia)中。
(2)基于形态学特征,将Bryhnia pflanzii移入青藓属中,B. angustifolia和B. tenerrima移入小燕尾藓属中,多枝燕尾藓(B. polyclada)和长肋燕尾藓(B.longinervis)移入柳叶藓科。
2.异叶藓属植物的分类学修订
(1)将异叶藓属植物的形态特征重新界定为：假鳞毛卵形至三角形,具短或长的毛尖；茎叶、枝叶异形;中肋近及顶：孢蒴下垂至水平；蒴盖具长喙,喙长与孢蒴近等长。
(2)异叶藓属中地位得到确认的种有5种,分别为：异叶藓(Kindbergia praelonga)、树状异叶藓(K. arbuscula)、俄勒冈异叶藓(K. oregana)、硬枝异叶藓(K. stokesii)和糙叶异叶藓(K. squarrifolia)。
(3)印度和不丹的Kindbergia dumosa及中国分布的“树状异叶藓”、“异叶藓”(包括密枝燕尾藓)的主要形态特征及遗传信息均不同于青藓科内现有任一类群,故为其建立一新属,即拟异叶藓属。
3.分类学结论
(1)新建小燕尾藓属(Bryhniastrum M. Li&Y.F., Wang)和拟异叶藓属(Pseudokindbergia M. Li, Y.F., Wang, B.C., Tan&Ignatov)两属。
(2)提出7个新组合、8个新异名和1个新地位：Brachythecium Schimp., Bryol. Eur.6:5.1853.=Bryhnia Kaurin, Bot. Not.1892(2):[60-]61.1892. syn. nov. Brachythecium hultenii (E.B.Bartram) M.Li&Y.F.Wang, comb. nov.=Brachythecium hultenii var. cymbifolia Nog., J. Hattori Bot. Lab.,8:19.1952. syn. nov. Brachythecium pflanzii (Broth.) M. Li&Y.F. Wang, comb. nov. Brachythecium trichomitria (Dix.&Ther.) M. Li&Y.F. Wang, comb. nov. Bryhniastrum graminicolor M. Li&Y.F. Wang, comb. nov.=Bryhnia angustifolia Warnst, Hedwigia57:115.50.1915. syn. nov. Bryhniastrum tenerrima (Broth.&Yasuda) M. Li&Y.F. Wang, comb. nov. Myuroclada longiramea (Miill.Hal.) M.Li, Y.F.Wang, M.S.Ignatov&Huttunen, comb. nov.=Bryhnia brachycladula Cardot, Bull. Soc. Bot., Geneve, Ser.2,4:379.1912. syn. nov. Pseudokindbergia dumosa (Mitt.) M. Li,Y.F. Wang, B.C. Tan&Ignatov, comb. nov. Brachythecium novae-angliae (Sull.&Lesq.) A. Jaeger=Bryhnia tokubuchii (Broth.) Paris, index Bryol.(ed.2)1:77.1904. syn. nov.Cratoneuron filicinum (Hedw.) Spruce=Bryhnia polyclada Ther., Repert. Spec. Nov. Regni Veg.31:26.1932. syn. nov.=Bryhnia longinervis Ther., Repert. Spec. Nov. Regni Veg.31:27.1932. syn. nov. Kindbergia stokesii (Turn.) M. Li&Y.F. Wang, stat. nov.=Kindbergia oedogonium (Mull. Hal.) Ochyra, Beih. Nova Hedwigia88:56.1987. syn. nov.
4.将小燕尾藓属植物的主要形态特征界定为：植物体细弱；叶卵状披针形至披针形；叶细胞具明显的前角突。拟异叶藓属植物的主要形态特征界定为：茎叶、枝叶异形；假鳞毛半圆形至三角状半圆形,先端钝或平截；孢蒴倾立至水平；蒴盖圆锥形。
5.确定了假鳞毛形态、孢蒴生长方向和蒴盖形态在青藓科内的属级水平分类中具有重要的分类学价值。
总之,本研究不仅解决了燕尾藓属和异叶藓属目前存在的主要问题,同时也为青藓科内与两属有密切联系的青藓属等类群的相关研究提供了有价值的资料,为存在类似问题的类群提供研究方法和思路上的参考。
Brachytheciaceae is one of the largest and most controversial families of pleurocarpous mosses. At the beginning of the21th century, the backbone phylogeny of the Brachytheciaceae has been set up. While problems with some particular morphologically similar genera and species still remain. In this study, the phylogeny of the two problematic genera in the family Brachytheciaceae, Bryhnia Kaurin and Kindbergia Ochyra, was conducted based on molecular and morphological data.
The morphological investigation was based on more than1000specimens of Bryhnia and Kindbergia, including67type materials belonging to39taxa. With Bryhnia and Kindbergia as the core taxa,98samples belonging to34species and14representative genera of the three subfamilies of the family Brachytheciaceae were used as ingroups for the molecular analyses. Among them,29samples from seven species of Bryhnia and40from four of Kindbergia were included. Barbella flagellifera (Cardot) Nog. and Meteorium atrovariegatum Cardot&Then from Meteoriaceae, the closest relative of Brachytheciaceae, were selected as outgroups. One nuclear and three plastid markers were chosen:nuclear ribosomal internal transcribed spacer region ITS1-5.8S-ITS2(ITS), ribosomal protein L16(rpl16), tRNA (Gly)(UCC)(trnG) and trnL-trnF intergenic spacer (trnL-F). Phylogenetic analyses were performed using Maximum parsimony (MP) and Bayesian inference (BI). The results are as follows:
1. Bryhnia is a polyphyletic group.
(1) Based on molecular and morphological data, Bryhnia Kaurin was here treated as a synonym of Brachythecium Schimp. Bryhnia novae-angliae, B. hultenii, B. scabrida and B. trichomitria were transferred to Brachythecium. Bryhnia brachycladula was transferred to the genus Myuroclada and further treated as a synonym of the new combination, Myuroclada longiramea (Mull. Hal.) M. Li, Y.F. Wang, M.S. Ignatov&S. Huttunen. Bryhnia graminicolor and B. serricuspis were transferred to the new genera, Bryhniastrum and Pseudokindbergia, respectively.
(2) Based on morphological data, Bryhnia pflanzii was transferred to the genus Brachythecium, B. angustifolia and B. tenerrima to Bryhniastrum, and B. polyclada and B. longinervis to Amblystegiaceae.
2. A revision of the genus Kindbergia Ochyra
(1) Ovate-apiculate or triangular and piliferous pseudoparaphyllia, dimorphic leaves, nearly percurrent costa, horizontal to subpendent capsule, and long-rostrate operculum were re-delimited as the key morphological characters of Kindbergia.
(2) Five species have been confirmed in Kindbergia. They are K. arbuscula, K. oregana, K. praelonga, K. squarrifolia and K. stokesii.
(3) Kindbergia dumosa distributed in India and Bhutan, and "K. arbuscula","K. praelonga", and B. serricuspis previously reported from China did not fit well in the existing genera molecularly and morphologically. A new monotypic genus, namely Pseudokindbergia, was established for them.
3. Taxonomic conclusions:
(1) Two new genera have been established. They are Bryhniastrum M. Li&Y.F., Wang and Pseudokindbergia M. Li, Y.F. Wang, B.C. Tan&Ignatov, respectively.
(2) Seven new combinations, eight synonyms and one new status are proposed: Brachythecium Schimp., Bryol. Eur.6:5.1853.=Bryhnia Kaurin, Bot.Not.1892(2):[60-]61.1892. syn. nov. Brachythecium hultenii (E.B.Bartram) M.Li&Y.F.Wang, comb. nov.=Brachythecium hultenii var. cymbifolia Nog., J. Hattori Bot. Lab.,8:19.1952. syn. nov. Brachythecium pflanzii (Broth.) M. Li&Y.F. Wang, comb. nov. Brachythecium trichomitria (Dix.&Ther.) M. Li&Y.F. Wang, comb. nov. Bryhniastrum graminicolor M. Li&Y.F. Wang, comb. nov.-Bryhnia angustifolia. Warnst., Hedwigia57:115.50.1915. syn. nov. Bryhniastrum tenerrima (Broth.&Yasuda) M. Li&Y.F. Wang, comb. nov. Myuroclada longiramea (Mull.Hal.) M.Li, Y.F.Wang, M.S.Ignatov&Huttunen, comb. nov.=Bryhnia brachycladula Cardot, Bull. Soc. Bot., Geneve, Ser.2,4:379.1912. syn. nov. Pseudokindbergia dumosa (Mitt.) M. Li,Y.F. Wang, B.C. Tan&Ignatov, comb. nov. Brachythecium novae-angliae (Sull.&Lesq.) A. Jaeger=Bryhnia tokubuchii (Broth.) Paris, index Bryol.(ed.2)1:77.1904. syn. nov. Cratoneuron filicinum (Hedw.) Spruce=Bryhnia polyclada Then, Repert. Spec. Nov. Regni Veg.31:26.1932. syn. nov.=Bryhnia longinervis Then, Repert. Spec. Nov. Regni Veg.31:27.1932. syn. nov. Kindbergia stokesii (Turn.) M. Li&Y.F. Wang, stat. nov.=Kindbergia oedogonium (Mull. Hal.) Ochyra, Beih. Nova Hedwigia88:56.1987. syn. nov.
4. Bryhniastrum was characterized by slender plants, ovate-lanceolate to lanceolate leaves and prorate laminal cells, and Pseudokindbergia by bimorphic leaves, semi-orbicular to orbicular-triangular, obtuse or truncate pseudoparaphyllia, inclined to horizontal capsule and conic-apiculate operculum.
5. The shape of pseudoparaphyllia, operculum and capsule orientation are reliable morphological characters to resolve genera level relationships in the family Brachytheciaceae.
In summary, this study has solved most of the taxonomic problems with Bryhnia and Kindbergia. And the results provide valuable information and methods for allied genera in Brachytheciaceae and other taxa with similiar taxonomic problems.
经典分类学研究主要基于1000余份燕尾藓属和异叶藓属的标本,其中包括模式标本67份,隶属于39种(包括种下单位)。分子系统学研究选择以燕尾藓属和异叶藓属为核心的青藓科3亚科的14属、34种、98份样本作为内群,其中燕尾藓属7种、29份样本,异叶藓属4种、40份样本；蔓藓科(Meteoriaceae)的鞭枝悬藓(Barbella flagellifera)和东亚蔓藓(Meteorium atrovariegatum)为外群；选用1个核基因ITS和3个叶绿体基因rpl16, trnG, trnL-F,运用最大简约法(MP)和贝叶斯分析(BI)两种方法进行了分子系统学分析。主要研究结果如下：
1.燕尾藓属是一个多系类群。
(1)综合经典分类学和分子系统学研究结果,将燕尾藓属处理为青藓属的异名；将燕尾藓(Bryhnia novae-angliae)、短尖燕尾藓(B. hultenii)、 B. scabrida、毛尖燕尾藓(B. trichomitria)移入了青藓属；短枝燕尾藓(B. brachycladula)移入鼠尾藓属,并处理为羽枝鼠尾藓(Myuroclada longiramea)的异名；而Bryhnia graminicolor和密枝燕尾藓(B. serricuspis)分别被移入2个新建属,即小燕尾藓属(Bryhniastrum)和拟异叶藓属(Pseudokindbergia)中。
(2)基于形态学特征,将Bryhnia pflanzii移入青藓属中,B. angustifolia和B. tenerrima移入小燕尾藓属中,多枝燕尾藓(B. polyclada)和长肋燕尾藓(B.longinervis)移入柳叶藓科。
2.异叶藓属植物的分类学修订
(1)将异叶藓属植物的形态特征重新界定为：假鳞毛卵形至三角形,具短或长的毛尖；茎叶、枝叶异形;中肋近及顶：孢蒴下垂至水平；蒴盖具长喙,喙长与孢蒴近等长。
(2)异叶藓属中地位得到确认的种有5种,分别为：异叶藓(Kindbergia praelonga)、树状异叶藓(K. arbuscula)、俄勒冈异叶藓(K. oregana)、硬枝异叶藓(K. stokesii)和糙叶异叶藓(K. squarrifolia)。
(3)印度和不丹的Kindbergia dumosa及中国分布的“树状异叶藓”、“异叶藓”(包括密枝燕尾藓)的主要形态特征及遗传信息均不同于青藓科内现有任一类群,故为其建立一新属,即拟异叶藓属。
3.分类学结论
(1)新建小燕尾藓属(Bryhniastrum M. Li&Y.F., Wang)和拟异叶藓属(Pseudokindbergia M. Li, Y.F., Wang, B.C., Tan&Ignatov)两属。
(2)提出7个新组合、8个新异名和1个新地位：Brachythecium Schimp., Bryol. Eur.6:5.1853.=Bryhnia Kaurin, Bot. Not.1892(2):[60-]61.1892. syn. nov. Brachythecium hultenii (E.B.Bartram) M.Li&Y.F.Wang, comb. nov.=Brachythecium hultenii var. cymbifolia Nog., J. Hattori Bot. Lab.,8:19.1952. syn. nov. Brachythecium pflanzii (Broth.) M. Li&Y.F. Wang, comb. nov. Brachythecium trichomitria (Dix.&Ther.) M. Li&Y.F. Wang, comb. nov. Bryhniastrum graminicolor M. Li&Y.F. Wang, comb. nov.=Bryhnia angustifolia Warnst, Hedwigia57:115.50.1915. syn. nov. Bryhniastrum tenerrima (Broth.&Yasuda) M. Li&Y.F. Wang, comb. nov. Myuroclada longiramea (Miill.Hal.) M.Li, Y.F.Wang, M.S.Ignatov&Huttunen, comb. nov.=Bryhnia brachycladula Cardot, Bull. Soc. Bot., Geneve, Ser.2,4:379.1912. syn. nov. Pseudokindbergia dumosa (Mitt.) M. Li,Y.F. Wang, B.C. Tan&Ignatov, comb. nov. Brachythecium novae-angliae (Sull.&Lesq.) A. Jaeger=Bryhnia tokubuchii (Broth.) Paris, index Bryol.(ed.2)1:77.1904. syn. nov.Cratoneuron filicinum (Hedw.) Spruce=Bryhnia polyclada Ther., Repert. Spec. Nov. Regni Veg.31:26.1932. syn. nov.=Bryhnia longinervis Ther., Repert. Spec. Nov. Regni Veg.31:27.1932. syn. nov. Kindbergia stokesii (Turn.) M. Li&Y.F. Wang, stat. nov.=Kindbergia oedogonium (Mull. Hal.) Ochyra, Beih. Nova Hedwigia88:56.1987. syn. nov.
4.将小燕尾藓属植物的主要形态特征界定为：植物体细弱；叶卵状披针形至披针形；叶细胞具明显的前角突。拟异叶藓属植物的主要形态特征界定为：茎叶、枝叶异形；假鳞毛半圆形至三角状半圆形,先端钝或平截；孢蒴倾立至水平；蒴盖圆锥形。
5.确定了假鳞毛形态、孢蒴生长方向和蒴盖形态在青藓科内的属级水平分类中具有重要的分类学价值。
总之,本研究不仅解决了燕尾藓属和异叶藓属目前存在的主要问题,同时也为青藓科内与两属有密切联系的青藓属等类群的相关研究提供了有价值的资料,为存在类似问题的类群提供研究方法和思路上的参考。
Brachytheciaceae is one of the largest and most controversial families of pleurocarpous mosses. At the beginning of the21th century, the backbone phylogeny of the Brachytheciaceae has been set up. While problems with some particular morphologically similar genera and species still remain. In this study, the phylogeny of the two problematic genera in the family Brachytheciaceae, Bryhnia Kaurin and Kindbergia Ochyra, was conducted based on molecular and morphological data.
The morphological investigation was based on more than1000specimens of Bryhnia and Kindbergia, including67type materials belonging to39taxa. With Bryhnia and Kindbergia as the core taxa,98samples belonging to34species and14representative genera of the three subfamilies of the family Brachytheciaceae were used as ingroups for the molecular analyses. Among them,29samples from seven species of Bryhnia and40from four of Kindbergia were included. Barbella flagellifera (Cardot) Nog. and Meteorium atrovariegatum Cardot&Then from Meteoriaceae, the closest relative of Brachytheciaceae, were selected as outgroups. One nuclear and three plastid markers were chosen:nuclear ribosomal internal transcribed spacer region ITS1-5.8S-ITS2(ITS), ribosomal protein L16(rpl16), tRNA (Gly)(UCC)(trnG) and trnL-trnF intergenic spacer (trnL-F). Phylogenetic analyses were performed using Maximum parsimony (MP) and Bayesian inference (BI). The results are as follows:
1. Bryhnia is a polyphyletic group.
(1) Based on molecular and morphological data, Bryhnia Kaurin was here treated as a synonym of Brachythecium Schimp. Bryhnia novae-angliae, B. hultenii, B. scabrida and B. trichomitria were transferred to Brachythecium. Bryhnia brachycladula was transferred to the genus Myuroclada and further treated as a synonym of the new combination, Myuroclada longiramea (Mull. Hal.) M. Li, Y.F. Wang, M.S. Ignatov&S. Huttunen. Bryhnia graminicolor and B. serricuspis were transferred to the new genera, Bryhniastrum and Pseudokindbergia, respectively.
(2) Based on morphological data, Bryhnia pflanzii was transferred to the genus Brachythecium, B. angustifolia and B. tenerrima to Bryhniastrum, and B. polyclada and B. longinervis to Amblystegiaceae.
2. A revision of the genus Kindbergia Ochyra
(1) Ovate-apiculate or triangular and piliferous pseudoparaphyllia, dimorphic leaves, nearly percurrent costa, horizontal to subpendent capsule, and long-rostrate operculum were re-delimited as the key morphological characters of Kindbergia.
(2) Five species have been confirmed in Kindbergia. They are K. arbuscula, K. oregana, K. praelonga, K. squarrifolia and K. stokesii.
(3) Kindbergia dumosa distributed in India and Bhutan, and "K. arbuscula","K. praelonga", and B. serricuspis previously reported from China did not fit well in the existing genera molecularly and morphologically. A new monotypic genus, namely Pseudokindbergia, was established for them.
3. Taxonomic conclusions:
(1) Two new genera have been established. They are Bryhniastrum M. Li&Y.F., Wang and Pseudokindbergia M. Li, Y.F. Wang, B.C. Tan&Ignatov, respectively.
(2) Seven new combinations, eight synonyms and one new status are proposed: Brachythecium Schimp., Bryol. Eur.6:5.1853.=Bryhnia Kaurin, Bot.Not.1892(2):[60-]61.1892. syn. nov. Brachythecium hultenii (E.B.Bartram) M.Li&Y.F.Wang, comb. nov.=Brachythecium hultenii var. cymbifolia Nog., J. Hattori Bot. Lab.,8:19.1952. syn. nov. Brachythecium pflanzii (Broth.) M. Li&Y.F. Wang, comb. nov. Brachythecium trichomitria (Dix.&Ther.) M. Li&Y.F. Wang, comb. nov. Bryhniastrum graminicolor M. Li&Y.F. Wang, comb. nov.-Bryhnia angustifolia. Warnst., Hedwigia57:115.50.1915. syn. nov. Bryhniastrum tenerrima (Broth.&Yasuda) M. Li&Y.F. Wang, comb. nov. Myuroclada longiramea (Mull.Hal.) M.Li, Y.F.Wang, M.S.Ignatov&Huttunen, comb. nov.=Bryhnia brachycladula Cardot, Bull. Soc. Bot., Geneve, Ser.2,4:379.1912. syn. nov. Pseudokindbergia dumosa (Mitt.) M. Li,Y.F. Wang, B.C. Tan&Ignatov, comb. nov. Brachythecium novae-angliae (Sull.&Lesq.) A. Jaeger=Bryhnia tokubuchii (Broth.) Paris, index Bryol.(ed.2)1:77.1904. syn. nov. Cratoneuron filicinum (Hedw.) Spruce=Bryhnia polyclada Then, Repert. Spec. Nov. Regni Veg.31:26.1932. syn. nov.=Bryhnia longinervis Then, Repert. Spec. Nov. Regni Veg.31:27.1932. syn. nov. Kindbergia stokesii (Turn.) M. Li&Y.F. Wang, stat. nov.=Kindbergia oedogonium (Mull. Hal.) Ochyra, Beih. Nova Hedwigia88:56.1987. syn. nov.
4. Bryhniastrum was characterized by slender plants, ovate-lanceolate to lanceolate leaves and prorate laminal cells, and Pseudokindbergia by bimorphic leaves, semi-orbicular to orbicular-triangular, obtuse or truncate pseudoparaphyllia, inclined to horizontal capsule and conic-apiculate operculum.
5. The shape of pseudoparaphyllia, operculum and capsule orientation are reliable morphological characters to resolve genera level relationships in the family Brachytheciaceae.
In summary, this study has solved most of the taxonomic problems with Bryhnia and Kindbergia. And the results provide valuable information and methods for allied genera in Brachytheciaceae and other taxa with similiar taxonomic problems.
引文
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