国产郁金香属植物分类学问题研究
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摘要
本文通过野外调查及凭证标本的考证，并结合叶表皮形态、花粉、种皮的微形态及比较胚胎学等方面的观察，对国产郁金香属植物分类中存在的问题进行了研究，主要内容如下：
（1）对国产郁金香属植物进行了了修订，确认了国产郁金香属植物共有16种，其中新种1个: Tulipa kuocangshanica D.Y.Tan X.R.Li et D.Y. Hong。
（2）对国产4组16种郁金香植物叶上下表皮的形态学观察，发现叶表皮形态在种间均存在一定程度的差异，但在组间，除了有苞组的4个种与其它种类有明显差异外，其余3组间没有明显的差异，表明郁金香属植物的叶表皮特征在种级水平上具有一定的分类学意义，可以作为种的划分及种间亲缘关系探讨的参考指标之一。
（3）对该属19种植物花粉形态学观察发现，花粉形态和外壁纹饰在组间和种间存在着一定差异。花粉形态有近球形、近舟形和近盾形3种；外壁纹饰可分为五大类：网状纹饰、条纹状纹饰、瘤状纹饰、疣状纹饰和皱波状纹饰，其中网状纹饰细分为2亚类，条纹网状纹饰也可细分为2亚类。花粉形态和外壁纹饰在组间及种间均有一定的分类学意义。
（4）所观察的该属13种植物的种皮表面纹饰可分为网纹、负网纹和皱波状三大类，且各个种之间在种皮微形态上均存在着一定差异，可以作为分种的依据；在组间，除属于有苞组(Sect. Amana) 的T. edulis和其它组的种类有明显差异外，其余3组之间没有明显的界限，说明郁金香属的种皮微形态特征在组的划分上意义不大，但在种的划分上具有一定的分类学意义。
（5）比较胚胎学：国产郁金香属16种植物的胚囊类型共有6种，即 Fritillaria type、Drusa type、T. iliensis type、T. tetraphylla type、Adoxa type、和Eriostemones Type；至少有3种具有两种胚囊类型，如T.iliensis、 T.heteropetala 和T.heterophylla。在所报道的5组共45种郁金香属植物中，发现该属植物的胚囊发育类型具有一定的系统学意义。
In this paper, the taxonomical problems of Tulipa L. from China were studied based on the results of both the field observation and the examination of herbarium specimens,and the analysis of leaf epidermis morphology, pollen morphology, seed coats micromorphology and comparative embryology. The main results are as follows:
(1) Tulipa L. from China is taxonomically revised .There are 16 species distrubuted in China and Tulipa kuocangshanica D.Y.Tan,X.R.Li et D.Y.Hong is described as new.
(2) The morphological characteristics of leaf epidermis of 16 species were studied. The results show that the morphological characteristics of leaf epidermis have not obvious differences among sections except Sect. Amana, but there are some differences between species , and indicate that the morphological characteristics of leaf epidermis have some value for the interspecific classification .
(3)The pollen morphology of 19 species was investigated under SEM . The results show that the morphological characteristics and exine ornamentation are not similar among species and sections. the morphological characteristics are recognized as 3 types: nearly globe,nearly boat shape and nearly shield shape .The exine ornamentation can be divided into 5 types : reticulate, steiate-veticulate, tuberculate,verrucate and rugose-sinuate. The morphological characteristics show some vaviations among sections and species in terms of the shape of pollen and the exine ornamentation.and thus can be used as one of characters in the consideration of the classification of this genus.
(4) The micromorphology of the seed coats were observed in 13 species. The results show that the surface structure of the seeds have obvious differences and can be divided into 3 types including reticulate,areolate and rugose-sinuate. Features of the micromorphology of the seed coats can be used as evidences to classify clearly this genus.
(5) There are 6 types of embryo sac in 16 species from China , such as Eriostemones type, Fritillaria type, Adoxa type, Drusa type ,T. illiensis type and T. tetraphylla type. There are more than one types of embryo sac in 3 species ,sush as T. illiensis,T.heteropetala and T. heterophylla.
According to the analysis of comparative embryology, we think that the types of the embryo sac are of some systematical significances.
（1）对国产郁金香属植物进行了了修订，确认了国产郁金香属植物共有16种，其中新种1个: Tulipa kuocangshanica D.Y.Tan X.R.Li et D.Y. Hong。
（2）对国产4组16种郁金香植物叶上下表皮的形态学观察，发现叶表皮形态在种间均存在一定程度的差异，但在组间，除了有苞组的4个种与其它种类有明显差异外，其余3组间没有明显的差异，表明郁金香属植物的叶表皮特征在种级水平上具有一定的分类学意义，可以作为种的划分及种间亲缘关系探讨的参考指标之一。
（3）对该属19种植物花粉形态学观察发现，花粉形态和外壁纹饰在组间和种间存在着一定差异。花粉形态有近球形、近舟形和近盾形3种；外壁纹饰可分为五大类：网状纹饰、条纹状纹饰、瘤状纹饰、疣状纹饰和皱波状纹饰，其中网状纹饰细分为2亚类，条纹网状纹饰也可细分为2亚类。花粉形态和外壁纹饰在组间及种间均有一定的分类学意义。
（4）所观察的该属13种植物的种皮表面纹饰可分为网纹、负网纹和皱波状三大类，且各个种之间在种皮微形态上均存在着一定差异，可以作为分种的依据；在组间，除属于有苞组(Sect. Amana) 的T. edulis和其它组的种类有明显差异外，其余3组之间没有明显的界限，说明郁金香属的种皮微形态特征在组的划分上意义不大，但在种的划分上具有一定的分类学意义。
（5）比较胚胎学：国产郁金香属16种植物的胚囊类型共有6种，即 Fritillaria type、Drusa type、T. iliensis type、T. tetraphylla type、Adoxa type、和Eriostemones Type；至少有3种具有两种胚囊类型，如T.iliensis、 T.heteropetala 和T.heterophylla。在所报道的5组共45种郁金香属植物中，发现该属植物的胚囊发育类型具有一定的系统学意义。
In this paper, the taxonomical problems of Tulipa L. from China were studied based on the results of both the field observation and the examination of herbarium specimens,and the analysis of leaf epidermis morphology, pollen morphology, seed coats micromorphology and comparative embryology. The main results are as follows:
(1) Tulipa L. from China is taxonomically revised .There are 16 species distrubuted in China and Tulipa kuocangshanica D.Y.Tan,X.R.Li et D.Y.Hong is described as new.
(2) The morphological characteristics of leaf epidermis of 16 species were studied. The results show that the morphological characteristics of leaf epidermis have not obvious differences among sections except Sect. Amana, but there are some differences between species , and indicate that the morphological characteristics of leaf epidermis have some value for the interspecific classification .
(3)The pollen morphology of 19 species was investigated under SEM . The results show that the morphological characteristics and exine ornamentation are not similar among species and sections. the morphological characteristics are recognized as 3 types: nearly globe,nearly boat shape and nearly shield shape .The exine ornamentation can be divided into 5 types : reticulate, steiate-veticulate, tuberculate,verrucate and rugose-sinuate. The morphological characteristics show some vaviations among sections and species in terms of the shape of pollen and the exine ornamentation.and thus can be used as one of characters in the consideration of the classification of this genus.
(4) The micromorphology of the seed coats were observed in 13 species. The results show that the surface structure of the seeds have obvious differences and can be divided into 3 types including reticulate,areolate and rugose-sinuate. Features of the micromorphology of the seed coats can be used as evidences to classify clearly this genus.
(5) There are 6 types of embryo sac in 16 species from China , such as Eriostemones type, Fritillaria type, Adoxa type, Drusa type ,T. illiensis type and T. tetraphylla type. There are more than one types of embryo sac in 3 species ,sush as T. illiensis,T.heteropetala and T. heterophylla.
According to the analysis of comparative embryology, we think that the types of the embryo sac are of some systematical significances.
引文
1. 陈汉斌,山东植物志（上卷）,青岛出版社, 青岛, 1992,773
2. 陈机等，植物发育解剖学,山东大学出版社, 济南,1996
3. 陈晓亚，胡成华，喻富根，阮嘉蓬，方竹属（竹亚科）叶片表皮微形态特征，植物分类学报，1993，31（3）：227－235
4. 崔乃然,宋宗仁等,新疆草地名录,新疆人民出版社,乌鲁木齐, 1990
5. 额尔特曼著，王伏雄,钱南芬译,花粉形态与植物分类,科学出版社,北京,1962
6. 侯宽昭,中国种子科属词典（修订版）,科学出版社,北京,1998
7. 胡适宜，被子植物胚胎学,高等教育出版社,北京,1982
8. 江苏植物所编,江苏植物志（上）,江苏省人民出版社,江苏, 1977,350
9. 李书心, 辽宁植物志(下册),辽宁科学技术出版社, 辽宁, 1992,698
10. 李正理，植物制片技术（第二版）,科学出版社, 北京, 1987
11. 李锦馨等，郁金香引种试验初报,宁夏农林科技，1991（2）：30－32,
12. 吕海亮，吴世安，杨继，绕广远，叶表皮及种皮特征在黄精族系统中的应用,植物分类学报, 2000,38(1):30-42
13. 梁松筠，百合科（狭义）植物的分布区对中国植物区系的研究的意义,植物分类学报，1995,33 (1) ：27-51
14. 梁凤丽,伊犁郁金香生殖生物学研究,新疆农业大学硕士学位论文,2000
15. 林泉,浙江植物志,浙江人民出版社, 杭州, 1993,434
16. 玛海希瓦里 著，陈机译,被子植物胚胎学引论,科学出版社, 北京, 1966
17. 毛祖美，郁金香属,中国植物志,科学出版社，北京, 1980,14: 86-97
18. 毛祖美，新疆郁金香属植物研究，干旱区研究，1984,1（2）：39－43
19. 毛祖美，郁金香属， 新疆植物志，新疆科技卫生出版社，乌鲁木齐，1996，6: 496-504
20. 宁布,郁金香属,内蒙古植物志(第二版), 内蒙人民出版社, 呼和浩特,1994,5:469
21. 内蒙古植物志编委会(马疏泉主编),内蒙古植物志, 内蒙古人民出版社, 呼和浩特,1985,8：469
22. 裴鉴,江苏南部种子植物手册,北京,科学出版社,1959,146
23. 上海科学技术研究所主编, 上海植物志（上卷）,上海科学技术出版社,上海,1999,656
24. 沈显生，中国郁金香属一新种，云南植物研究，2001，23（1）：39-40
25. 汤彦承等， 中国百合科系统概要及对今后的一些意见，植物研究，1983，3（2）: 56-71
26. 谭敦炎，魏星等， 新疆郁金香属新分类群 ,植物分类学报，2000，38（3）：302-304
27. 谭永平， 新疆郁金香繁育生物学研究，北京师范大学硕士论文,1997
28. 王玉荣,新疆郁金香植物生物学特性及资源评价研究,新疆农业大学硕士论文,2001
29. 王伏雄，钱南芬等,中国植物花粉形态（第二版）,科学出版社,北京,1995
30. 王开发等,孢粉学概论,北京大学出版社,北京,1983
31. 王伟,赵南先,天南星科叶表皮研究,武汉植物学研究,2002,20(5):343-349
32. 熊治廷，洪德元，陈瑞阳，核型不对称的一种定量测定法及其在进化研究中的应用，植物分类学报，1992，30（3）：279－288
33. 新疆八一农学院编，新疆植物检索表（第一册），新疆人民出版社，乌鲁木齐,1982
34. 徐炳声等，染色体研究的进展与植物分类学（上,下），武汉植物学研究，1996，14（2）：177-187
35. 中国科学院新疆生物土壤沙漠研究所编,新疆药用植物志(第二册)，新疆人民出版社，乌鲁木齐，1981,188
36. 中国科学院西北植物研究所编,秦岭植物志（第一卷）种子植物（第一册）,科学出版社,北京,1976,358
37. 中国科学院植物所主编,中国高等植物图鉴,科学出版社,北京,1980,5：445-447
38. 中国科学院兰州沙漠研究所编（主编:刘瑛心),中国沙漠植物志,科学出版社,北京,1985,1：221
39. 郑国锠等编，生物显微技术（第二版），高等教育出版社，北京,1993
40. 张俭 等编著, 郁金香，中国林业出版社，北京,1994
41. 张志耘,卢宝荣,温洁,稻属叶表皮结构及其系统学意义,植物分类学报,1998,36(1):8-18
42. 张树仁,梁松筠,戴伦凯,苔草属复序苔草亚属十四种植物叶片的解剖学研究,植物分类学报,1998,36(4):333-340
43. Baker,J.G. Revision of the genera and species of Tulipeae. J. Linn. Soc. Bot., 1874,14:275-296.
44. Blakey D.H.，Vosa. C.G, Heterochromatin and chromosome variation in Cultivated Species of Tulipa subg.Leiostemones(Liliaceae),Pl.Syst. Evol, 1982,139:163-178
45. Booy G.J. ,Van der Schoot,et al, Heterogeneity of the internal transcribed spacer 1(ITSI)in Tulipa(Liliaceae),Pl. Syst Evol. 2000,225:29-41
46. Boissier E. ,Flora Orientalis, V. Deneve, Basel and Lyon., 1882
47. Botschantzeva Z. P.,Tulips (Taxonomy, morphology, cytology, phytogeography and physiology), translated and edited by Varekamp H. Q. , Netherlands:A. A. Balkema， Rotterdam， 1962
48. BryanD.Ness,SeedMorphology and TaxonomicRelationshipsin Calochortus (Liliaceae),
systematic Botany,1998,14(4):495-505
Chen, S.-C. and Mordik, H. V., Tulipa L. In: Wu Z.-Y. and Raven, P. H. (eds.). Flora
49. of China ，Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis) . USA. ， 2000，24. pp. 123-126
50. Conover M.,Epidermal patterns of the reticulate-veined Liliiflorae and their Parallel-veined allies,Bot J linn Soc., 1991,107:295-312
51. Davis,G.LSystematic embryology of the angiosperms，1966
52. Gary Filion, W. & David h. Blakey, Differential giemsa staining in plants， VI. Centromeric banding, Can.J.Genet.Cytol， 1979，21: 373-378
53. Gary Filion, W., Differential Giemsa Staining in Plants I， Banding Patterns in Three Cultivars of Tulipa, Chromosoma (Berl.)， 1974 ，49:51-60
54. George W.Johnston, Abnormal Pollen of Tulipa， Phytom orphology，1959，9: 320-325
55. Hall, The genus Tulipa， The Royal Horticultural Society， London, 1940
56. Hoog. M. H.， On the origin of Tulipa-Lilies and other Liliaceae, Royal Horticultural Society， London ，1973， 47-64
57. Kroon,G.H & Jongerius,M.C, Chromosome numbers of Tulipa species and the occurrence of hexaploidy,Euphytica，1986,35:73-76
58. Leningrad:Izd.An.USSR. 1935,32 (4): 289--301
59. Mark W. Woods and Ronald Bamford,Chromosome Morphology and Number in Tulipa， American Journal of Botany，1937，24 (4): 175-184
60. Nieuwhof M., Van Raamsdonk L. W. D. et al.，Figment composition of flowers of Tulipa species as a parameter for biosystematic research， Biochemical systematic and ecology, 1990，18(6): 399－404
61. Guignard L., Lappareil sexuel et le double fecondation dans les Tulipes, Ann.Sci. Nat. ser. Bot., 1900, 11:365-387
62. Hall,A.D., The book of the tulip, London, Polyploid in tulips, Gard. Chron, 1929
63. Hall,A.D.,Polyploid in Tulipa,J.Linn.Soc.Bot. ,1937 ,50:481-489
64. Hall,A.D.,The genus Tulipa,London, 1940, R.H.S.
65. Newton,W. C.F., Chromosome studies in Tulipa and some related genera.,J.Linn. Soc. Bot., 1927,47:339-354
66. Newton,W. C.F. ，C.D.Darlington, Meiosis in a triploid tulip. Nature, 1927,120:13
67. Newton,W. C.F.，C.D.Darlington, Meiosis in polyploid , PartⅠTriploid and pentaploid tulips,PartⅡ,Aneuploid hyacinths, J. Genet.,1929,21(1):1-15
68. Newton,W. C.F.，C.D.Darlington, Meiosis in polyploid tulips, J. Genet,1929,19
69. NairP.K.,M.Sharma,PollenMorphologyofliliaceae,Journ.Palynol.Lucknow ,1965,1:38-6
70. Regel E.A.，Enumeratio specierum hucusgue cognitarum generic Tulipae, 1873,2:437-457
71. Romanov I.D., Novaya forma zarodyshevogo meshka Adoxa Tulipa tetraphylla i T. ostrowskiana (A new Adoxa-type embryo-sac in Tulipa tetraphylla and T. ostrowskiana),C.R.Acad. sci. USSR,1938 ,22(3):113-115
72. Romanov,I.D., Dve novye formy zarodyshevogo meshka v rode Tulipa(Two new types of embryo-sac of the genus Tulipa).,C.R.Acad. sci. USSR, 1939,22(3): 139-141
73. Romanov,I.D. , Evolyutsia zarodyshevogo meshka tsvetkovykh rastenii (The evolution of the embryo-sacin the flowering plants),Aft.diss. Sois. Uch. Step.dokt.boil.nauk.Tashkent, 1944
74. Romanov,I.D.,K.anatomii reliktovykh rastenii, Anatomiya lista Tulipa regelii(A contribution to the anatomy of relict plants, The leaf of Tulipa regelii),Mosk. Obshch. Ispyt. Prir. Otd. Boil, 1952 ,57(5):79-87
75. Siro Kitamura ,Gen Murata, Tetsuo Koyama, Coloured illus-trations of Herbaceous Plants Of Japan (Monocotyledoneae),Hoikusha Publishing Co.LTD,Osaka, Japan, 1980,540
76. Stork,A.,Tulipessauvagesetcultivees.,Geneva:ConservatoireetJardin botaniques,Serie documentaire，1984，13
77. Taliev V.I., Protsess vidoobrazovaniya v rode Tulipa sektsii Eriostemones (Species formation in the section Eriostemones of the genus Tulipa), Bull. Appl.Bot.Gen.Pl.-Breed,1930,24(2):57-122
78. Tamura, M. N.. Liliaceae . In : Kubitzki , K. (eds.) The families and genera of vascular plants. Ⅲ. Flowering plants. -Monocotyledons Lilianae. Springer-Verlag, Berlin,Heidelberg ,1998
79. Tan Dunyan,Li Xinrong,Hong Deyuan,A new species of the genus tulipa(Liliaceae)from China(unpublished)
80. Van Creij. M.G.M,etc.,Interspecific Crosses in the Genus Tulipa L.: Identification of Pre-fertilization Barriers, Sex Plant Reprod, 1983,10: 116--123
81. Van Eijk J. P., Van Raamsdonk, L.W.D.et al.,Interspecific crosses between Tulipa gesneriana cultivars and wild Tulipa species, a survey, Sex. Pland Reprod. 1991 4:1-5
82. Van Raamsdonk L. W. D. & T. De Vries, Biosystematic studies in Tulipa sect. Eriostemones (Liliaceae), Plant Systematics and Evolution, 1992 ,179:27-41
83. Van Raamsdonk L. W. D. & T. De Vries, Species relationships and taxonomy in Tulipa subg. Tulipa (Liliaceae), Plant Systematics and Evolution, 1995, 195: 13-44
84. Van Raamsdonk L. W. D.，Flower pigment composion in Tulipa.,Genetic resources and crop evolution 1993,40: 49-54
85. Van Raamsdonk, Joop.Van Eijk .et al.,Crossability analysis in subgenus Tulipa of the Genus Tulipa L. Botanical Journal of the Linnean Society,1995 ,117:147--158
86. Vasilevskaya V.K.,Znachenie,anatomicheskikh priznakov vcheshue lukovits sredne aziats kikh tyul'panovdlya ikh sistematiki (Importance des caracteres anatomigues dans la tunique des bulbes de tulips de l'Asie Centrale pour leur systematique),Trud. Sred. Az. Gos. Un. Ser.8b,Bot, 1936, 33
87. Vvedenskii A.I., Tulipae at Junones novae. Byull. Sred.-aziat, goss. Un. ,1935 ,21: 147-152
88. Vvedenskii A.I., Tulipa L. in:Flora of the USSR, 1935,4:246-280
89. Vvedenskii A.I.,Tulipa binutans Vved. In: Not. Syst. Herb. Inst. Bot. Acad. Sci. Uzbek.,1952,13:33
90. Wafai B.A.& Koul A.K., Impact of Polyploidy on the Gametophytes, Ovules and Seeds in Some Himalayan Tulips, Phytomorpology,1981,34 (4): 64-79
91. Wilhelm B.Epidermai and seed surface characters of plants:systematic applicability and some evolutionary aspects,Nord.J.Bot.,1981,1(3):345-355
92. Woods M.W., The nucleolus in Tulipa. Am. J. Bot., 1937,24:528-536
93. Woods M.,R.Bamford,Chromosome morphology and numbers in Tulipa,Am.J.Bot. 1937 ,24(4):175-184
94. Yoshisuke Satake, Jisaburo Ohwi, Siro Kitamura,Shunji Watari ,Tadao Tominari,WILD FLOOWERS OF JAPAN (Herbaceous Plants), Heibonsha Ltd, Publishers ,Tokyo,1982
95. Zhai S-H et al, The karyotype analysis of Tulipa Spp. In Xinjiang, Proc Sino-Jpn. Symposium Plant Chromosome Research, 1987, 95-100
2. 陈机等，植物发育解剖学,山东大学出版社, 济南,1996
3. 陈晓亚，胡成华，喻富根，阮嘉蓬，方竹属（竹亚科）叶片表皮微形态特征，植物分类学报，1993，31（3）：227－235
4. 崔乃然,宋宗仁等,新疆草地名录,新疆人民出版社,乌鲁木齐, 1990
5. 额尔特曼著，王伏雄,钱南芬译,花粉形态与植物分类,科学出版社,北京,1962
6. 侯宽昭,中国种子科属词典（修订版）,科学出版社,北京,1998
7. 胡适宜，被子植物胚胎学,高等教育出版社,北京,1982
8. 江苏植物所编,江苏植物志（上）,江苏省人民出版社,江苏, 1977,350
9. 李书心, 辽宁植物志(下册),辽宁科学技术出版社, 辽宁, 1992,698
10. 李正理，植物制片技术（第二版）,科学出版社, 北京, 1987
11. 李锦馨等，郁金香引种试验初报,宁夏农林科技，1991（2）：30－32,
12. 吕海亮，吴世安，杨继，绕广远，叶表皮及种皮特征在黄精族系统中的应用,植物分类学报, 2000,38(1):30-42
13. 梁松筠，百合科（狭义）植物的分布区对中国植物区系的研究的意义,植物分类学报，1995,33 (1) ：27-51
14. 梁凤丽,伊犁郁金香生殖生物学研究,新疆农业大学硕士学位论文,2000
15. 林泉,浙江植物志,浙江人民出版社, 杭州, 1993,434
16. 玛海希瓦里 著，陈机译,被子植物胚胎学引论,科学出版社, 北京, 1966
17. 毛祖美，郁金香属,中国植物志,科学出版社，北京, 1980,14: 86-97
18. 毛祖美，新疆郁金香属植物研究，干旱区研究，1984,1（2）：39－43
19. 毛祖美，郁金香属， 新疆植物志，新疆科技卫生出版社，乌鲁木齐，1996，6: 496-504
20. 宁布,郁金香属,内蒙古植物志(第二版), 内蒙人民出版社, 呼和浩特,1994,5:469
21. 内蒙古植物志编委会(马疏泉主编),内蒙古植物志, 内蒙古人民出版社, 呼和浩特,1985,8：469
22. 裴鉴,江苏南部种子植物手册,北京,科学出版社,1959,146
23. 上海科学技术研究所主编, 上海植物志（上卷）,上海科学技术出版社,上海,1999,656
24. 沈显生，中国郁金香属一新种，云南植物研究，2001，23（1）：39-40
25. 汤彦承等， 中国百合科系统概要及对今后的一些意见，植物研究，1983，3（2）: 56-71
26. 谭敦炎，魏星等， 新疆郁金香属新分类群 ,植物分类学报，2000，38（3）：302-304
27. 谭永平， 新疆郁金香繁育生物学研究，北京师范大学硕士论文,1997
28. 王玉荣,新疆郁金香植物生物学特性及资源评价研究,新疆农业大学硕士论文,2001
29. 王伏雄，钱南芬等,中国植物花粉形态（第二版）,科学出版社,北京,1995
30. 王开发等,孢粉学概论,北京大学出版社,北京,1983
31. 王伟,赵南先,天南星科叶表皮研究,武汉植物学研究,2002,20(5):343-349
32. 熊治廷，洪德元，陈瑞阳，核型不对称的一种定量测定法及其在进化研究中的应用，植物分类学报，1992，30（3）：279－288
33. 新疆八一农学院编，新疆植物检索表（第一册），新疆人民出版社，乌鲁木齐,1982
34. 徐炳声等，染色体研究的进展与植物分类学（上,下），武汉植物学研究，1996，14（2）：177-187
35. 中国科学院新疆生物土壤沙漠研究所编,新疆药用植物志(第二册)，新疆人民出版社，乌鲁木齐，1981,188
36. 中国科学院西北植物研究所编,秦岭植物志（第一卷）种子植物（第一册）,科学出版社,北京,1976,358
37. 中国科学院植物所主编,中国高等植物图鉴,科学出版社,北京,1980,5：445-447
38. 中国科学院兰州沙漠研究所编（主编:刘瑛心),中国沙漠植物志,科学出版社,北京,1985,1：221
39. 郑国锠等编，生物显微技术（第二版），高等教育出版社，北京,1993
40. 张俭 等编著, 郁金香，中国林业出版社，北京,1994
41. 张志耘,卢宝荣,温洁,稻属叶表皮结构及其系统学意义,植物分类学报,1998,36(1):8-18
42. 张树仁,梁松筠,戴伦凯,苔草属复序苔草亚属十四种植物叶片的解剖学研究,植物分类学报,1998,36(4):333-340
43. Baker,J.G. Revision of the genera and species of Tulipeae. J. Linn. Soc. Bot., 1874,14:275-296.
44. Blakey D.H.，Vosa. C.G, Heterochromatin and chromosome variation in Cultivated Species of Tulipa subg.Leiostemones(Liliaceae),Pl.Syst. Evol, 1982,139:163-178
45. Booy G.J. ,Van der Schoot,et al, Heterogeneity of the internal transcribed spacer 1(ITSI)in Tulipa(Liliaceae),Pl. Syst Evol. 2000,225:29-41
46. Boissier E. ,Flora Orientalis, V. Deneve, Basel and Lyon., 1882
47. Botschantzeva Z. P.,Tulips (Taxonomy, morphology, cytology, phytogeography and physiology), translated and edited by Varekamp H. Q. , Netherlands:A. A. Balkema， Rotterdam， 1962
48. BryanD.Ness,SeedMorphology and TaxonomicRelationshipsin Calochortus (Liliaceae),
systematic Botany,1998,14(4):495-505
Chen, S.-C. and Mordik, H. V., Tulipa L. In: Wu Z.-Y. and Raven, P. H. (eds.). Flora
49. of China ，Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis) . USA. ， 2000，24. pp. 123-126
50. Conover M.,Epidermal patterns of the reticulate-veined Liliiflorae and their Parallel-veined allies,Bot J linn Soc., 1991,107:295-312
51. Davis,G.LSystematic embryology of the angiosperms，1966
52. Gary Filion, W. & David h. Blakey, Differential giemsa staining in plants， VI. Centromeric banding, Can.J.Genet.Cytol， 1979，21: 373-378
53. Gary Filion, W., Differential Giemsa Staining in Plants I， Banding Patterns in Three Cultivars of Tulipa, Chromosoma (Berl.)， 1974 ，49:51-60
54. George W.Johnston, Abnormal Pollen of Tulipa， Phytom orphology，1959，9: 320-325
55. Hall, The genus Tulipa， The Royal Horticultural Society， London, 1940
56. Hoog. M. H.， On the origin of Tulipa-Lilies and other Liliaceae, Royal Horticultural Society， London ，1973， 47-64
57. Kroon,G.H & Jongerius,M.C, Chromosome numbers of Tulipa species and the occurrence of hexaploidy,Euphytica，1986,35:73-76
58. Leningrad:Izd.An.USSR. 1935,32 (4): 289--301
59. Mark W. Woods and Ronald Bamford,Chromosome Morphology and Number in Tulipa， American Journal of Botany，1937，24 (4): 175-184
60. Nieuwhof M., Van Raamsdonk L. W. D. et al.，Figment composition of flowers of Tulipa species as a parameter for biosystematic research， Biochemical systematic and ecology, 1990，18(6): 399－404
61. Guignard L., Lappareil sexuel et le double fecondation dans les Tulipes, Ann.Sci. Nat. ser. Bot., 1900, 11:365-387
62. Hall,A.D., The book of the tulip, London, Polyploid in tulips, Gard. Chron, 1929
63. Hall,A.D.,Polyploid in Tulipa,J.Linn.Soc.Bot. ,1937 ,50:481-489
64. Hall,A.D.,The genus Tulipa,London, 1940, R.H.S.
65. Newton,W. C.F., Chromosome studies in Tulipa and some related genera.,J.Linn. Soc. Bot., 1927,47:339-354
66. Newton,W. C.F. ，C.D.Darlington, Meiosis in a triploid tulip. Nature, 1927,120:13
67. Newton,W. C.F.，C.D.Darlington, Meiosis in polyploid , PartⅠTriploid and pentaploid tulips,PartⅡ,Aneuploid hyacinths, J. Genet.,1929,21(1):1-15
68. Newton,W. C.F.，C.D.Darlington, Meiosis in polyploid tulips, J. Genet,1929,19
69. NairP.K.,M.Sharma,PollenMorphologyofliliaceae,Journ.Palynol.Lucknow ,1965,1:38-6
70. Regel E.A.，Enumeratio specierum hucusgue cognitarum generic Tulipae, 1873,2:437-457
71. Romanov I.D., Novaya forma zarodyshevogo meshka Adoxa Tulipa tetraphylla i T. ostrowskiana (A new Adoxa-type embryo-sac in Tulipa tetraphylla and T. ostrowskiana),C.R.Acad. sci. USSR,1938 ,22(3):113-115
72. Romanov,I.D., Dve novye formy zarodyshevogo meshka v rode Tulipa(Two new types of embryo-sac of the genus Tulipa).,C.R.Acad. sci. USSR, 1939,22(3): 139-141
73. Romanov,I.D. , Evolyutsia zarodyshevogo meshka tsvetkovykh rastenii (The evolution of the embryo-sacin the flowering plants),Aft.diss. Sois. Uch. Step.dokt.boil.nauk.Tashkent, 1944
74. Romanov,I.D.,K.anatomii reliktovykh rastenii, Anatomiya lista Tulipa regelii(A contribution to the anatomy of relict plants, The leaf of Tulipa regelii),Mosk. Obshch. Ispyt. Prir. Otd. Boil, 1952 ,57(5):79-87
75. Siro Kitamura ,Gen Murata, Tetsuo Koyama, Coloured illus-trations of Herbaceous Plants Of Japan (Monocotyledoneae),Hoikusha Publishing Co.LTD,Osaka, Japan, 1980,540
76. Stork,A.,Tulipessauvagesetcultivees.,Geneva:ConservatoireetJardin botaniques,Serie documentaire，1984，13
77. Taliev V.I., Protsess vidoobrazovaniya v rode Tulipa sektsii Eriostemones (Species formation in the section Eriostemones of the genus Tulipa), Bull. Appl.Bot.Gen.Pl.-Breed,1930,24(2):57-122
78. Tamura, M. N.. Liliaceae . In : Kubitzki , K. (eds.) The families and genera of vascular plants. Ⅲ. Flowering plants. -Monocotyledons Lilianae. Springer-Verlag, Berlin,Heidelberg ,1998
79. Tan Dunyan,Li Xinrong,Hong Deyuan,A new species of the genus tulipa(Liliaceae)from China(unpublished)
80. Van Creij. M.G.M,etc.,Interspecific Crosses in the Genus Tulipa L.: Identification of Pre-fertilization Barriers, Sex Plant Reprod, 1983,10: 116--123
81. Van Eijk J. P., Van Raamsdonk, L.W.D.et al.,Interspecific crosses between Tulipa gesneriana cultivars and wild Tulipa species, a survey, Sex. Pland Reprod. 1991 4:1-5
82. Van Raamsdonk L. W. D. & T. De Vries, Biosystematic studies in Tulipa sect. Eriostemones (Liliaceae), Plant Systematics and Evolution, 1992 ,179:27-41
83. Van Raamsdonk L. W. D. & T. De Vries, Species relationships and taxonomy in Tulipa subg. Tulipa (Liliaceae), Plant Systematics and Evolution, 1995, 195: 13-44
84. Van Raamsdonk L. W. D.，Flower pigment composion in Tulipa.,Genetic resources and crop evolution 1993,40: 49-54
85. Van Raamsdonk, Joop.Van Eijk .et al.,Crossability analysis in subgenus Tulipa of the Genus Tulipa L. Botanical Journal of the Linnean Society,1995 ,117:147--158
86. Vasilevskaya V.K.,Znachenie,anatomicheskikh priznakov vcheshue lukovits sredne aziats kikh tyul'panovdlya ikh sistematiki (Importance des caracteres anatomigues dans la tunique des bulbes de tulips de l'Asie Centrale pour leur systematique),Trud. Sred. Az. Gos. Un. Ser.8b,Bot, 1936, 33
87. Vvedenskii A.I., Tulipae at Junones novae. Byull. Sred.-aziat, goss. Un. ,1935 ,21: 147-152
88. Vvedenskii A.I., Tulipa L. in:Flora of the USSR, 1935,4:246-280
89. Vvedenskii A.I.,Tulipa binutans Vved. In: Not. Syst. Herb. Inst. Bot. Acad. Sci. Uzbek.,1952,13:33
90. Wafai B.A.& Koul A.K., Impact of Polyploidy on the Gametophytes, Ovules and Seeds in Some Himalayan Tulips, Phytomorpology,1981,34 (4): 64-79
91. Wilhelm B.Epidermai and seed surface characters of plants:systematic applicability and some evolutionary aspects,Nord.J.Bot.,1981,1(3):345-355
92. Woods M.W., The nucleolus in Tulipa. Am. J. Bot., 1937,24:528-536
93. Woods M.,R.Bamford,Chromosome morphology and numbers in Tulipa,Am.J.Bot. 1937 ,24(4):175-184
94. Yoshisuke Satake, Jisaburo Ohwi, Siro Kitamura,Shunji Watari ,Tadao Tominari,WILD FLOOWERS OF JAPAN (Herbaceous Plants), Heibonsha Ltd, Publishers ,Tokyo,1982
95. Zhai S-H et al, The karyotype analysis of Tulipa Spp. In Xinjiang, Proc Sino-Jpn. Symposium Plant Chromosome Research, 1987, 95-100