中国杜鹃属映山红亚属植物系统分类研究
摘要
杜鹃属Rhododendron L.全球约1000种,是典型的北温带分布区类型。随着许多新种的发现和先进技术在分类上的应用,这个属的分类问题也日益突出。本文以形态分类为主线,在查阅相关模式标本、非模式标本的基础上,选取在分类上具有重要价值和存在争议的种类,到模式标本产地取样,应用现代分子标记技术、导管形态和叶解剖技术以取得相关的证据,对中国映山红亚属Subgenus Tsutsusi (Sweet) Pojarkova的系统分类进行综合分析,得到如下结论:
1.本文基于ITS序列分析、维管束导管形态分析和叶解剖结构分析等证据,中国映山红亚属有74种2个亚种13个变种和1个变型;其中映山红组有69种
2亚种和12个变种,轮叶组有5种1变种和1个变型。此外,本文提出了一个映山红亚属系统排列。
2.依据ITS分析,映山红亚属应该划分为2个组,即映山红组Sect.Tsutsusi Sweet和轮叶组Sect.Brachycalyx Sweet。
3.根据ITS分析,大武杜鹃Rhododendron tashiroi Maxim.应该放在轮叶组Sect.Brachycalyx Sweet比较合适。
4.ITS分析表示,大字杜鹃Rhododendron schlippenbachii Maxim.没有与映山红亚属聚在同一个分支上,因此它可能有独立的系统地位,需要进一步研究。
5.根据ITS分析、导管形态分析、叶解剖结构分析和表形形态分析,灰齿杜鹃Rhododendron cinereoserratum Tam、戴云山杜鹃Rhododendron daiyunicum Tam不能独立成为一个种,本文研究认为应该把戴云山杜鹃作为满山红的变种处理比较合适,把灰齿杜鹃处理为满山红的异名;保留白花满山红Rhododendron mariesii Hemsl.et Wils.f.albescense B.Y. Ding et G .R.Chen在发表的原文中作满山红的变型;。因此中国映山红亚属轮叶组目前应包括5个种和1个变种、1个变型。
此外,本文研究认为天堂杜鹃Rhododendron tiantangense G .Z .Li是亮毛杜鹃Rhododendron microphyton Franch的新组合;同时把棒柱杜鹃Rhododendron crassimedium Tam处理为背绒杜鹃Rhododendron hypoblematosum Tam的异名;把茶绒杜鹃Rhododendron rufulum Tam处理为龙岩杜鹃Rhododendron florulentum Tam的异名。
6.通过对映山红亚属的区系分析,中国映山红亚属的种数最多,占该亚属种类的69%,是映山亚属的多样化中心,也是映山红组的多样化中心;但日本是轮叶组的多样化中心,因为中国仅有5种1个变种和1个变型,日本达19种。在中国大陆映山红亚属的种数依次是广西>广东>湖南>江西>贵州>福建>四川、云南,其中主要分布在广东、广西、湖南、江西这4省区。映山红亚属的物种多样化中心在广西-广东-湖南南部。
映山红亚属可能由耳叶杜鹃亚组最近的祖先类群演化过来,因此在这个亚属中较原始的类群可能是具刚毛、腺毛的5枚雄蕊类群。这些类群主要分布在广西、广东及湖南南部,因此广西、广东及湖南南部既是映山红亚属的起源中心和物种多样性中心。
7.轮叶组的类群可能直接由映山红组的一些原始类群演化而来,前者在获得的新性状后以较快的速率进化,与映山红组形成姐妹群,说明这两个组有较密切的亲缘关系。
8.映山红亚属在我国广西-广东-湖南南部起源之后,由如下途径向东南、东北部地区散布:起源地→马来西亚诸岛屿→菲律宾;中国大陆→日本;中国大陆→台湾。
9.新的发现
(1)上犹杜鹃新变种Rhododendron seniavinii Maxim. var. Rhododendron shangyounicum R .L. Liu et T. Z.Du,var. nov.
(2)在粗柱杜鹃的叶解剖结构中下表皮的上方,发现有一层可能由海绵组织细胞异化为“棉花状”的细胞,这种细胞的名称和功能需要进一步讨论。
(3)在研究中发现,毛果杜鹃、南昆杜鹃、溪畔杜鹃存在“拟穿孔板”(暂时命名),其名称、功能和起源需要进一步研究。
As it is well known , the genus Rhododendron L. is a large one with the number of 1000 species in the world, it is considered as a type of temperate distribution .Since the genus Rhododendron had been established by Linnaeus in 1753,much has been written on the classification of this genus.Because of the collection being in an ever-increasing stream from any place in the world, particularly from China ,more and more new species of Rhododendron have been named ,and at the same time ,problems of systematic classification of Rhododendron are occurring. At the same time,there are same problems occurring in the Subgenus Tsutsusi (Sweet) Pojarkova,for example, some species newly named from China in the recent of 30 years are difficult to be divided ,and whether sect.Brachycalyx Sweet includes Rhododendron tashiroi Maxim. etc. The question now arises: are we to accept the present classification or are we to substitute some other classification in its place? In order to make a clear classification for the subgenus Tsutsusi,a thorough study and a detailed examination of the subgenus Tsutsusi has been made in this paper based on the analysis of ITS sequences ,features of leaf anatomic structure ,features of vessels and characters of morphology.
With modern technique of cassification such as molecular marker and plant anatomy,a study on the systematic classification of subgenus Tsutsus of Rhododend -ron in China is made in this paper, in which 74 species and 2 subspecies and 13 varieties and 1 form belonging to this subgenus have been discussed with the conclusions as follows:
1.Based on various evidences from the analysis of ITS , features of leaf anatomic structure ,features of vessels and characters of morphology in this paper , Subgenus Tsutsusi (Sweet) Pojarkova of Rhododendro has 74species and 2subspecies and 13 varieties and 1 form in China ; Sect.Tsutsusi Sweet of this subgenus has 69species and
2 subspecies and 12 varieties ; Sect.Brachycalyx Sweet of this subgenus has now 5 species and 1 variety and 1 form . In addition , a new arrangement of systemic classification of this subgenus has been offered in this paper.
2.Based on ananlysis of ITS,it is considered to be correct that the Subgenus Tsutsusi (Sweet) Pojarkova of Rhododendron is classified into two sections ,namely section Tsutsusi Sweet and section Brachycalyx Sweet.The Sect.Tsutsusi Sweet and Sect.Brachycalyx Sweet are sister groups belonging to Subgenus Tsutsusi (Sweet) Pojarkova.
3.It is right that Rhododendron tashiroi Maxim. is placed in Sect.Brachycalyx Sweet according to the ananlysis of ITS.
4. Based on ananlysis of ITS ,the systematic position of Rhododendron schlippenbachii Maxim. is in need of further discussion because of its position shown in the phyletic tree in which it has not clustered with Subgenus Tsutsusi .
5.Based on various evidences from the analysis of ITS,features of leaf anatomic structure ,features of vessels and characters of morphology,it is considered to be right that species Rhododendron daiyunicum Tam is treated as a variety of species Rhododendron mariesii Hemsl. et Wils.and and Rhododendron mariesii Hemsl. et Wils. f. albescense B.Y.Ding et G . R.Chen published in 1989 is retained as a form of Rhododendron mariesii Hemsl. et Wils.; Species Rhododendron cinereoserratum is a synonym of Rhododendron mariesii Hemsl. et Wils.; therefore the Sect.Brachycalyx Sweet has now 4 species and 1 variety and 1 form in China .
Species Rhododendron tiantangense G .Z .Li is a new combination of Rhododen -dron microphyton Franch ; Rhododendron crassimedium Tam is treated as a synonym of Rhododendron hypoblematosum Tam and Rhododendron rufulum Tam is a synonym of Rhododendron florulentum Tam
6.Based on the analysis of the flora of subgenus Tsutsusi Sweet in this paper , some important conclusions have been obtained as follows: China is the center of species diversity of subgenus Tsutsusi and of Sect.Tsutsusi ; but Japan is the center of species diversity of section Brachycalyx .
Most of species of subgenus Tsutsusi distribute in China , the number of species of this subgenus is in order as Guangxi> Guangdong>Hunan>Jiangxi>Gui Zhou>Fujian>Sichuan or Yunnan . It is sure that the combination region in Guangxi and Guangdong and Hunan is the center of species diversity of subgenus Tsutsusi in the world .
The subgenus Tsutsusi origins from the combination region in Guangxi and Guangdong and Hunan in China , it evolutes directly from the primitive group of subsect. Auriculata .
7.Based on the analysis of ITS and paleogeography and fig.XIV , Sect. Brachyca -lyx Sweet probably evolutes from the primitive group of Sect.Tsutsus.Sweet , it evolutes quickly with new characters and form a sister group with Sect.Tsutsus.Sweet which show close connection between these two sections .
8.The routes of emigration of Subgenus Tsutsusi (Sweet) Pojarkova coming from its original region are considered as follows : southern and western part of China→ islands of Malaysia→northern part of Philippine ; southern and western part of China→Japan ; mainland of China→Taiwan.
9.Discovery
(1) A new variety of species : Rhododendron seniavinii Maxim. var. Rhododen -dron shangyounicum R .L. Liu et T. Z.Du,var. nov.
(2) A type of idio-cells which is similar to the“cotton”has been found in mesophyll tissue of Rhododendron crassistylum M.Y.He and it is named as“cotton”cell in this paper. It is necessary to discuss what is the function of this type of cells .
(3) A special type of pits named“perforation-like plates”is found in the vessels of species Rhododendron rivulare Hand. -Mazz. and Rhododendron . naamkwanense Merr.and Rhododendron.seniavinii Maxim.,it is valuable in evolution and in Classification of Rhododendron , its function and origin is in need to further research .
1.本文基于ITS序列分析、维管束导管形态分析和叶解剖结构分析等证据,中国映山红亚属有74种2个亚种13个变种和1个变型;其中映山红组有69种
2亚种和12个变种,轮叶组有5种1变种和1个变型。此外,本文提出了一个映山红亚属系统排列。
2.依据ITS分析,映山红亚属应该划分为2个组,即映山红组Sect.Tsutsusi Sweet和轮叶组Sect.Brachycalyx Sweet。
3.根据ITS分析,大武杜鹃Rhododendron tashiroi Maxim.应该放在轮叶组Sect.Brachycalyx Sweet比较合适。
4.ITS分析表示,大字杜鹃Rhododendron schlippenbachii Maxim.没有与映山红亚属聚在同一个分支上,因此它可能有独立的系统地位,需要进一步研究。
5.根据ITS分析、导管形态分析、叶解剖结构分析和表形形态分析,灰齿杜鹃Rhododendron cinereoserratum Tam、戴云山杜鹃Rhododendron daiyunicum Tam不能独立成为一个种,本文研究认为应该把戴云山杜鹃作为满山红的变种处理比较合适,把灰齿杜鹃处理为满山红的异名;保留白花满山红Rhododendron mariesii Hemsl.et Wils.f.albescense B.Y. Ding et G .R.Chen在发表的原文中作满山红的变型;。因此中国映山红亚属轮叶组目前应包括5个种和1个变种、1个变型。
此外,本文研究认为天堂杜鹃Rhododendron tiantangense G .Z .Li是亮毛杜鹃Rhododendron microphyton Franch的新组合;同时把棒柱杜鹃Rhododendron crassimedium Tam处理为背绒杜鹃Rhododendron hypoblematosum Tam的异名;把茶绒杜鹃Rhododendron rufulum Tam处理为龙岩杜鹃Rhododendron florulentum Tam的异名。
6.通过对映山红亚属的区系分析,中国映山红亚属的种数最多,占该亚属种类的69%,是映山亚属的多样化中心,也是映山红组的多样化中心;但日本是轮叶组的多样化中心,因为中国仅有5种1个变种和1个变型,日本达19种。在中国大陆映山红亚属的种数依次是广西>广东>湖南>江西>贵州>福建>四川、云南,其中主要分布在广东、广西、湖南、江西这4省区。映山红亚属的物种多样化中心在广西-广东-湖南南部。
映山红亚属可能由耳叶杜鹃亚组最近的祖先类群演化过来,因此在这个亚属中较原始的类群可能是具刚毛、腺毛的5枚雄蕊类群。这些类群主要分布在广西、广东及湖南南部,因此广西、广东及湖南南部既是映山红亚属的起源中心和物种多样性中心。
7.轮叶组的类群可能直接由映山红组的一些原始类群演化而来,前者在获得的新性状后以较快的速率进化,与映山红组形成姐妹群,说明这两个组有较密切的亲缘关系。
8.映山红亚属在我国广西-广东-湖南南部起源之后,由如下途径向东南、东北部地区散布:起源地→马来西亚诸岛屿→菲律宾;中国大陆→日本;中国大陆→台湾。
9.新的发现
(1)上犹杜鹃新变种Rhododendron seniavinii Maxim. var. Rhododendron shangyounicum R .L. Liu et T. Z.Du,var. nov.
(2)在粗柱杜鹃的叶解剖结构中下表皮的上方,发现有一层可能由海绵组织细胞异化为“棉花状”的细胞,这种细胞的名称和功能需要进一步讨论。
(3)在研究中发现,毛果杜鹃、南昆杜鹃、溪畔杜鹃存在“拟穿孔板”(暂时命名),其名称、功能和起源需要进一步研究。
As it is well known , the genus Rhododendron L. is a large one with the number of 1000 species in the world, it is considered as a type of temperate distribution .Since the genus Rhododendron had been established by Linnaeus in 1753,much has been written on the classification of this genus.Because of the collection being in an ever-increasing stream from any place in the world, particularly from China ,more and more new species of Rhododendron have been named ,and at the same time ,problems of systematic classification of Rhododendron are occurring. At the same time,there are same problems occurring in the Subgenus Tsutsusi (Sweet) Pojarkova,for example, some species newly named from China in the recent of 30 years are difficult to be divided ,and whether sect.Brachycalyx Sweet includes Rhododendron tashiroi Maxim. etc. The question now arises: are we to accept the present classification or are we to substitute some other classification in its place? In order to make a clear classification for the subgenus Tsutsusi,a thorough study and a detailed examination of the subgenus Tsutsusi has been made in this paper based on the analysis of ITS sequences ,features of leaf anatomic structure ,features of vessels and characters of morphology.
With modern technique of cassification such as molecular marker and plant anatomy,a study on the systematic classification of subgenus Tsutsus of Rhododend -ron in China is made in this paper, in which 74 species and 2 subspecies and 13 varieties and 1 form belonging to this subgenus have been discussed with the conclusions as follows:
1.Based on various evidences from the analysis of ITS , features of leaf anatomic structure ,features of vessels and characters of morphology in this paper , Subgenus Tsutsusi (Sweet) Pojarkova of Rhododendro has 74species and 2subspecies and 13 varieties and 1 form in China ; Sect.Tsutsusi Sweet of this subgenus has 69species and
2 subspecies and 12 varieties ; Sect.Brachycalyx Sweet of this subgenus has now 5 species and 1 variety and 1 form . In addition , a new arrangement of systemic classification of this subgenus has been offered in this paper.
2.Based on ananlysis of ITS,it is considered to be correct that the Subgenus Tsutsusi (Sweet) Pojarkova of Rhododendron is classified into two sections ,namely section Tsutsusi Sweet and section Brachycalyx Sweet.The Sect.Tsutsusi Sweet and Sect.Brachycalyx Sweet are sister groups belonging to Subgenus Tsutsusi (Sweet) Pojarkova.
3.It is right that Rhododendron tashiroi Maxim. is placed in Sect.Brachycalyx Sweet according to the ananlysis of ITS.
4. Based on ananlysis of ITS ,the systematic position of Rhododendron schlippenbachii Maxim. is in need of further discussion because of its position shown in the phyletic tree in which it has not clustered with Subgenus Tsutsusi .
5.Based on various evidences from the analysis of ITS,features of leaf anatomic structure ,features of vessels and characters of morphology,it is considered to be right that species Rhododendron daiyunicum Tam is treated as a variety of species Rhododendron mariesii Hemsl. et Wils.and and Rhododendron mariesii Hemsl. et Wils. f. albescense B.Y.Ding et G . R.Chen published in 1989 is retained as a form of Rhododendron mariesii Hemsl. et Wils.; Species Rhododendron cinereoserratum is a synonym of Rhododendron mariesii Hemsl. et Wils.; therefore the Sect.Brachycalyx Sweet has now 4 species and 1 variety and 1 form in China .
Species Rhododendron tiantangense G .Z .Li is a new combination of Rhododen -dron microphyton Franch ; Rhododendron crassimedium Tam is treated as a synonym of Rhododendron hypoblematosum Tam and Rhododendron rufulum Tam is a synonym of Rhododendron florulentum Tam
6.Based on the analysis of the flora of subgenus Tsutsusi Sweet in this paper , some important conclusions have been obtained as follows: China is the center of species diversity of subgenus Tsutsusi and of Sect.Tsutsusi ; but Japan is the center of species diversity of section Brachycalyx .
Most of species of subgenus Tsutsusi distribute in China , the number of species of this subgenus is in order as Guangxi> Guangdong>Hunan>Jiangxi>Gui Zhou>Fujian>Sichuan or Yunnan . It is sure that the combination region in Guangxi and Guangdong and Hunan is the center of species diversity of subgenus Tsutsusi in the world .
The subgenus Tsutsusi origins from the combination region in Guangxi and Guangdong and Hunan in China , it evolutes directly from the primitive group of subsect. Auriculata .
7.Based on the analysis of ITS and paleogeography and fig.XIV , Sect. Brachyca -lyx Sweet probably evolutes from the primitive group of Sect.Tsutsus.Sweet , it evolutes quickly with new characters and form a sister group with Sect.Tsutsus.Sweet which show close connection between these two sections .
8.The routes of emigration of Subgenus Tsutsusi (Sweet) Pojarkova coming from its original region are considered as follows : southern and western part of China→ islands of Malaysia→northern part of Philippine ; southern and western part of China→Japan ; mainland of China→Taiwan.
9.Discovery
(1) A new variety of species : Rhododendron seniavinii Maxim. var. Rhododen -dron shangyounicum R .L. Liu et T. Z.Du,var. nov.
(2) A type of idio-cells which is similar to the“cotton”has been found in mesophyll tissue of Rhododendron crassistylum M.Y.He and it is named as“cotton”cell in this paper. It is necessary to discuss what is the function of this type of cells .
(3) A special type of pits named“perforation-like plates”is found in the vessels of species Rhododendron rivulare Hand. -Mazz. and Rhododendron . naamkwanense Merr.and Rhododendron.seniavinii Maxim.,it is valuable in evolution and in Classification of Rhododendron , its function and origin is in need to further research .
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李光照,1995.广西杜鹃花属的修订极其地理分布 .广西植物,15(3):193-208.
李光照,2001.广西杜鹃花属新分类群 .广西植物,23(3):287-290.
李正理译,1973. 植物解剖学 . 北京:科学出版社,97—113 .
刘德生,1986.世界自然地理 (2 ed).北京:高等教育出版社,5-37.
路安民主编,1999.种子植物科属地理 .北京:科学出版社,299-319.
毛子军、杨永富,等,2000.东北杜鹃花属植物花粉形态研究 .植物研究,20(1):58-62.
毛加宁, 2000. 杜鹃属 4 种植物花粉形态特点研究 .西南农业大学学报,22(6):525-529).
闵天禄,方瑞征,1990.杜鹃属的系统发育与进化 .云南植物研究,12(4):356-365.
闵天禄,方瑞征,1979.杜鹃属的地理分布及其起源问题的探讨 .云南植物研究,12(4):356-365.
M.温坎坡,郑卓,1990. 被子植物的花粉形态系列与系统发育 . 植物分类学报,28(2):103-111.
熊子仙,杜青,王启德,2000.国产杜鹃花叶解剖与分类 .广西植物,20(4):335-338.
谭沛祥 , 1983. 华南杜鹃志 . 广州:广东科学技术出版社,2~31.
谭沛祥,1982. 华南杜鹃花研究 .植物研究,2(4): 77~78.
谭沛祥,1984.华南杜鹃花志 . 广州:广东科技出版社,1-82.
唐亚,谢嘉穗,高辉,2005. 锦葵目砚木属和柄翅果属的木材解剖学研究及其系统意义. 云南植物研究,27(3)235-246.
田欣,李德珠,2002. DNA 序列在植物进化研究中的应用 .云南植物研究,24:170-184.
王为义,1982.青海 9 种杜鹃的比较研究和分类位置 . 植物分类学报,20(1):49-57.
王伏雄译,1979.植物解剖学 ( K.伊稍 1965). 北京:科学出版社,109-201.
王辉宽,1999. 湘西杜鹃花植物根中甾醇类化合物的研究 .吉首大学学报,4-20.
张长芹,1993 .杜鹃属的子叶形态分类 .广西植物,13(1):16-22.
张昀,1998.生物进化 .北京:北京大学出版社,1-226.
张宏达主编,2004.种子植物系统学 . 北京:科学出版社,42-162.
郑万钧主编,1997 . 中国树木志 .北京:林业出版社.
中国科学院中国自然地理编委会,1986.中国自然地理(古地理).北京:科学出版社,1-21.
中国科学院大陆漂移翻译组, 1975. 大陆漂移 .(J.T 威儿逊著). 北京:科学出版社,42-162.
中国科学院植物所,1986.植物解剖实验技术 . 北京:科学出版社,1-32.
中国科学院植物研究所,1976. 中国高等植物图鉴 (第 3 册). 北京:科学出版社,1-56.
邹喻苹,葛颂,王晓东. 2001. 系统与进化植物学中的分子标记 . 北京:科学出版社,9-183.
吴德邻主编,1998. 广东植物志 (第一卷). 广州:广东科技出版社,296-327.
杨瑞林、魏学智,等, 2003. 杜鹃花属照山白叶表皮及花粉形态 .西北植物学报,23(5):806-810.
赵士洞译,1984. 国际植物命名法规 .北京:科学出版社,1-62.
Almut Seithe,nee von Hohh, 1980. Rhododendron Hairs and Taxonomy. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 89-116.
Avise,1994.Molecular Markers,Natural History and Evolution .New York: Chapman & Hall.
Bruce L., King, 1980.The Systematic Implications of Flavonoids in Rhododendron Subgenus Pentanthera. Contribution Toward a Classification of Rhododendron . New York Botanical Garden, 163-186.
Chamberlain D. F. & Rae S.J. 1990. A Revison of Rhododendron.Ⅳ Subgenus Tsutsusi. Edingburgh Journal of Botany 47(2):144-146.
David G., Leach,1959.Rhodododendron of the World . New York:Scribnors Press.
David F., Chamberlain D.F,1980. The Taxonomyof the Elepidote Rhododendron Excluding Azalea (Subgenus Hymenanthes). Contribution Toward a Classifica tion of Rhododendron.New York Botanical Garden, 39-62.
Davidian H. H, 1982. The Rhododendron Species . New York: Timber Press.
Escaravage N., Pornon A.,Doche B , Taberlet P. ,1998. Clonal Diversity in a Rhododendron ferrugineum L.(Ericaceae)Population Inferred from AFLP Members.M0lecular Ecology,7:975-982.
Evans D., Kathleen Kane H., Kinghts B.A and Math V.B, 1980.Chemical Taxonomy of Genus Rhododendron. Contribution Toward a Classification of Rhododendron . New York Botanical Garden, 187-196.
Fang Ming-Yuan,fang rui-zhen,He ming-you et al. 2005.Flora of China . Beijing : Science Press,260-455.
Fahn A. 1982.Plant Anatomy.(3ed.),Oxford:Pergamon Press.
Frost F H. 1930. Specialization in secondary xylem of dicotyledons:II Evolution of end wallof vessel segments .Botanical Gazette 90:178-196.
Hayes S.F., Keenan J.& Cowan J.M., 1951. A Survey of Anatomy of the Rhododendron Leaf in the Relation of the Taxonomy of the Genus . Notes RBG Edinb., 21(1):1-34).
Hillis,Moritz, 1996. Molecular Systematics(2ed).Sinauer Associates,Sunderland, Massachusettes. James Cullen,1980. Rhododendron Taxonomy and Nomenclature . Contribution Toward a Classifi-cation of Rhododendron .New York Botanical Garden,27-38.
Johannes Hedegaard, 1980.Morphological Studies in the Genus Rhododendron ,Dealing With Seeds Fruits and Seedings and Their Associated Hairs. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 117-114.
Jeffrey ,Harborne B, 1980.Flavonoid Pigments as Both taxonomic and Phyletic Markers in the Genus Rhododendron. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 145-162.
Judy Young & Lu-Sheng Chong,1980.Rhododendron of China .
Kurashige Y, Mine M , Kobayashi N, 1998. Investigation of Sectional Relationship in the Genus Rhododendron (Ericaceae) Based on matK Sequences. Journal of Japanese Botany. 73:143-154.
Kurashge Y,Etoh J I,Handa T, 2001. Sectional Relationship in the Genus Rhododendron (Ericaceae) : Evidences from matK and trnK Intron Sequences . Plant Systematics and Evolution ,228: 1-14。
Melva, 1980.Cotyledons and Rhodododendron Classification. Contribution Toward a Classification of Rhododendron .New York Botanical Garden,75-88.
Philipson W.R, 1980. Problems in Classification of the Azalea Complex. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 53-62.
Ruth Atkinson , Kwiton Jong, George Argent ,2000. Chromosome number of some tropical Rhododendrons (section Vireya). EDBGJ Bot , 57(1) : 1 —7.
Sleumer H, 1949. Ein System der Gattum Rhododendron L. Bot.Jahrb.74(4):511-553.
Sleumer H, 1980. Past and Present Taxonomic Systems of Rhododendron Based on Macromorpholo -gical Characters. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 19-26.
Stevenson J.B.Ed .1930. The Species of Rhododendron . Londen: Rhododendron Society, 1-120.
Swofford D L. 2001. PAUP: Phylogenetic Analysis Using Parsimony. Version4.0b8.Sunderland: Sinauer
Takasi Yamazaki.1996.A Revision of the genus Rhododendron in Japan,Taiwan,Korea and Sakhalin. Tokyo:Tsumura Laboratory.
Zhang G. H and al., 2001. Laboratory and Field Evaluation of Extracts from Rhododendron molle Flowers as Insect Growth Regulator to Imported Cabbage Worm , J.Appl. Ent., 563-569
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李红芳,任 毅,2005.领春木次生木质部中导管穿孔板的变异 (Eupteleaceae) .植物分类学报,43(1):1-11.
李光照,1995.广西杜鹃花属新分类群和新记录 .广西植物,15(4):293-301.
李光照,1995.广西杜鹃花属的修订极其地理分布 .广西植物,15(3):193-208.
李光照,2001.广西杜鹃花属新分类群 .广西植物,23(3):287-290.
李正理译,1973. 植物解剖学 . 北京:科学出版社,97—113 .
刘德生,1986.世界自然地理 (2 ed).北京:高等教育出版社,5-37.
路安民主编,1999.种子植物科属地理 .北京:科学出版社,299-319.
毛子军、杨永富,等,2000.东北杜鹃花属植物花粉形态研究 .植物研究,20(1):58-62.
毛加宁, 2000. 杜鹃属 4 种植物花粉形态特点研究 .西南农业大学学报,22(6):525-529).
闵天禄,方瑞征,1990.杜鹃属的系统发育与进化 .云南植物研究,12(4):356-365.
闵天禄,方瑞征,1979.杜鹃属的地理分布及其起源问题的探讨 .云南植物研究,12(4):356-365.
M.温坎坡,郑卓,1990. 被子植物的花粉形态系列与系统发育 . 植物分类学报,28(2):103-111.
熊子仙,杜青,王启德,2000.国产杜鹃花叶解剖与分类 .广西植物,20(4):335-338.
谭沛祥 , 1983. 华南杜鹃志 . 广州:广东科学技术出版社,2~31.
谭沛祥,1982. 华南杜鹃花研究 .植物研究,2(4): 77~78.
谭沛祥,1984.华南杜鹃花志 . 广州:广东科技出版社,1-82.
唐亚,谢嘉穗,高辉,2005. 锦葵目砚木属和柄翅果属的木材解剖学研究及其系统意义. 云南植物研究,27(3)235-246.
田欣,李德珠,2002. DNA 序列在植物进化研究中的应用 .云南植物研究,24:170-184.
王为义,1982.青海 9 种杜鹃的比较研究和分类位置 . 植物分类学报,20(1):49-57.
王伏雄译,1979.植物解剖学 ( K.伊稍 1965). 北京:科学出版社,109-201.
王辉宽,1999. 湘西杜鹃花植物根中甾醇类化合物的研究 .吉首大学学报,4-20.
张长芹,1993 .杜鹃属的子叶形态分类 .广西植物,13(1):16-22.
张昀,1998.生物进化 .北京:北京大学出版社,1-226.
张宏达主编,2004.种子植物系统学 . 北京:科学出版社,42-162.
郑万钧主编,1997 . 中国树木志 .北京:林业出版社.
中国科学院中国自然地理编委会,1986.中国自然地理(古地理).北京:科学出版社,1-21.
中国科学院大陆漂移翻译组, 1975. 大陆漂移 .(J.T 威儿逊著). 北京:科学出版社,42-162.
中国科学院植物所,1986.植物解剖实验技术 . 北京:科学出版社,1-32.
中国科学院植物研究所,1976. 中国高等植物图鉴 (第 3 册). 北京:科学出版社,1-56.
邹喻苹,葛颂,王晓东. 2001. 系统与进化植物学中的分子标记 . 北京:科学出版社,9-183.
吴德邻主编,1998. 广东植物志 (第一卷). 广州:广东科技出版社,296-327.
杨瑞林、魏学智,等, 2003. 杜鹃花属照山白叶表皮及花粉形态 .西北植物学报,23(5):806-810.
赵士洞译,1984. 国际植物命名法规 .北京:科学出版社,1-62.
Almut Seithe,nee von Hohh, 1980. Rhododendron Hairs and Taxonomy. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 89-116.
Avise,1994.Molecular Markers,Natural History and Evolution .New York: Chapman & Hall.
Bruce L., King, 1980.The Systematic Implications of Flavonoids in Rhododendron Subgenus Pentanthera. Contribution Toward a Classification of Rhododendron . New York Botanical Garden, 163-186.
Chamberlain D. F. & Rae S.J. 1990. A Revison of Rhododendron.Ⅳ Subgenus Tsutsusi. Edingburgh Journal of Botany 47(2):144-146.
David G., Leach,1959.Rhodododendron of the World . New York:Scribnors Press.
David F., Chamberlain D.F,1980. The Taxonomyof the Elepidote Rhododendron Excluding Azalea (Subgenus Hymenanthes). Contribution Toward a Classifica tion of Rhododendron.New York Botanical Garden, 39-62.
Davidian H. H, 1982. The Rhododendron Species . New York: Timber Press.
Escaravage N., Pornon A.,Doche B , Taberlet P. ,1998. Clonal Diversity in a Rhododendron ferrugineum L.(Ericaceae)Population Inferred from AFLP Members.M0lecular Ecology,7:975-982.
Evans D., Kathleen Kane H., Kinghts B.A and Math V.B, 1980.Chemical Taxonomy of Genus Rhododendron. Contribution Toward a Classification of Rhododendron . New York Botanical Garden, 187-196.
Fang Ming-Yuan,fang rui-zhen,He ming-you et al. 2005.Flora of China . Beijing : Science Press,260-455.
Fahn A. 1982.Plant Anatomy.(3ed.),Oxford:Pergamon Press.
Frost F H. 1930. Specialization in secondary xylem of dicotyledons:II Evolution of end wallof vessel segments .Botanical Gazette 90:178-196.
Hayes S.F., Keenan J.& Cowan J.M., 1951. A Survey of Anatomy of the Rhododendron Leaf in the Relation of the Taxonomy of the Genus . Notes RBG Edinb., 21(1):1-34).
Hillis,Moritz, 1996. Molecular Systematics(2ed).Sinauer Associates,Sunderland, Massachusettes. James Cullen,1980. Rhododendron Taxonomy and Nomenclature . Contribution Toward a Classifi-cation of Rhododendron .New York Botanical Garden,27-38.
Johannes Hedegaard, 1980.Morphological Studies in the Genus Rhododendron ,Dealing With Seeds Fruits and Seedings and Their Associated Hairs. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 117-114.
Jeffrey ,Harborne B, 1980.Flavonoid Pigments as Both taxonomic and Phyletic Markers in the Genus Rhododendron. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 145-162.
Judy Young & Lu-Sheng Chong,1980.Rhododendron of China .
Kurashige Y, Mine M , Kobayashi N, 1998. Investigation of Sectional Relationship in the Genus Rhododendron (Ericaceae) Based on matK Sequences. Journal of Japanese Botany. 73:143-154.
Kurashge Y,Etoh J I,Handa T, 2001. Sectional Relationship in the Genus Rhododendron (Ericaceae) : Evidences from matK and trnK Intron Sequences . Plant Systematics and Evolution ,228: 1-14。
Melva, 1980.Cotyledons and Rhodododendron Classification. Contribution Toward a Classification of Rhododendron .New York Botanical Garden,75-88.
Philipson W.R, 1980. Problems in Classification of the Azalea Complex. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 53-62.
Ruth Atkinson , Kwiton Jong, George Argent ,2000. Chromosome number of some tropical Rhododendrons (section Vireya). EDBGJ Bot , 57(1) : 1 —7.
Sleumer H, 1949. Ein System der Gattum Rhododendron L. Bot.Jahrb.74(4):511-553.
Sleumer H, 1980. Past and Present Taxonomic Systems of Rhododendron Based on Macromorpholo -gical Characters. Contribution Toward a Classification of Rhododendron .New York Botanical Garden, 19-26.
Stevenson J.B.Ed .1930. The Species of Rhododendron . Londen: Rhododendron Society, 1-120.
Swofford D L. 2001. PAUP: Phylogenetic Analysis Using Parsimony. Version4.0b8.Sunderland: Sinauer
Takasi Yamazaki.1996.A Revision of the genus Rhododendron in Japan,Taiwan,Korea and Sakhalin. Tokyo:Tsumura Laboratory.
Zhang G. H and al., 2001. Laboratory and Field Evaluation of Extracts from Rhododendron molle Flowers as Insect Growth Regulator to Imported Cabbage Worm , J.Appl. Ent., 563-569