黑龙江省31种蕨类植物的形态解剖学研究
摘要
利用光学显微镜对黑龙江省13科22属31种蕨类植物进行形态解剖学研究。结果如下：
1.木贼科问荆属2亚属植物表皮特征差别明显,问荆亚属植物表皮角质层成瘤状,气孔器不下陷,木贼表皮角质层不成瘤状,气孔器下陷；气孔器均为平列细胞型,由2个保卫细胞及2个副卫细胞构成,副卫细胞位于保卫细胞之上；气孔器上的辐射状纹饰来源于保卫细胞的腹壁(ventral wall),是细胞壁形成的片状凸起；所观察的木贼科5种植物主枝的基本结构一致,内皮层层数在2亚属植物中不稳定,不能将它作为分亚属的证据。
2.26种真蕨类植物叶表皮细胞多为不规则形,上、下表皮细胞垂周壁均呈波浪状,少数种类的上、下表皮细胞呈较规则的近长方形；共有6种气孔器类型,即极细胞型、腋下细胞型、不等细胞型、不规则四细胞型、不规则型和周细胞型,每种植物具1-4种不同类型的气孔器；气孔均为下生型气孔,一般散生,分布较均匀,并且沿着叶的长轴方向分布,只是偶尔在局部地方取向不规则,有时同一叶表皮的不同部分明显分布着不同类型的气孔。
3.蕨类植物鳞片的形状、网眼以及边缘的特征可以作为分类的依据之一
4.叶柄横切面显示,供试真蕨类植物维管束均为周韧型,可分为3种类型：(1)叶柄从基部向上至叶轴,维管束始终为一条,粗壮阴地蕨、桂皮紫萁、银粉背蕨、掌叶铁线蕨、过山蕨和耳羽岩蕨均属于这一类型；(2)叶柄基部具分体中柱2个,呈“八”字形排列,自叶柄基部向上,两个分体中柱于远轴面逐渐靠近,并连合成一个中柱,两束木质部远轴面末端的钩端相连,形成倒“V”字形至“人”字形,蹄盖蕨科供试的所有植物、卵果蕨、荚果蕨、球子蕨和乌苏里瓦韦均属于这一类型,除乌苏里瓦韦以外,其他植物木质部为“海马型”；(3)维管束多条,较规则的分散排列于基本组织之中,蕨、鳞毛蕨科供试的所有植物和有柄石韦属于这一类型。
5.供试真蕨类植物的气孔器均与表皮细胞在同一平面内,保卫细胞2个；多数植物叶肉没有分化为栅栏组织与海绵组织,可以明显的看出上表皮附近的同化组织比靠近下表皮的同化组织数量多,排列紧密；粗壮阴地蕨、桂皮紫萁、银粉背蕨、乌苏里瓦韦与有柄石韦同化组织均分化出栅栏组织与海绵组织。
6.与薄囊蕨纲的蕨类植物相比,木贼科植物、厚囊蕨纲的粗壮阴地蕨和原始薄囊蕨纲的桂皮紫萁的形态解剖学方面均表现出了较为原始的性状。
The epidermal structures, scales and cross section of 31 species from different families in Heilongjiang Province were investigated under light microscopes.The results indicated that:
1.The two subgenus of Equisetum shows apparent differences in the morphology of epidermis and stomatal position. Stomatas are both paracytic type and composed of two guard cells and two subsidiary cells,subsidiary cells located above the guard cells.Radial pattern of the stomatas derived from the apophysis on the ventral wall of the guard cells.The stem structure of five plants are similar,there are metaxylems in the vascular bundle.There are one or two layers of endodermis among the five species.
2.The leaf epidermal cells of 26 species of ferns in Heilongjiang are mostly irregular, the anticlinal walls of the upper and lower epidermal cells are wavy;There are six types of stomatal apparatus:Polocytic type,Axillocytic type,Aisocytic type,Anomotetracytic type,Anomocytic type and Pericytic type,each plant with one to four different types of stomatal apparatus;all the stomatal apparatuses only occur in the lower epidermis and distribution along the long axis of the leaves.
3.The shape, mesh and edge features of the ferns can be used as to classification.
4.The cross sections of the petioles show that the vascular bundle of the ferns investigated are all amphicribral,can be divided into three types:(1)There is only one vascular bundle in the petiole,such as Sceptridium robustum (Rupr.) Lyon,Osmunda cinnamomea L.var.asiatica Fernald,Aleuritopteris argentea (Gmel.) Fee,Adiantum pedatum L.,Camptosorus sibiricus Rupr.and Woodsia polystichoides Eaton;(2)The vascular patterns of the basal part of stipe are two meristeles distributing in the shape of "八",the xylems in the two meristeles are typical hippocampus type and gradually combine into amphicribral vascular bundles upwards, such as all the ferns of Athyriaceae observed,Phegopteris connectilis (Michx.) Watt,Matteuccia struthiopteris (L.) Todaro,Onoclea sensibilis L.var.interrupta Maxim.and Lepisorus ussuriensis (Regel et Maack) Ching;(3)Several vascular patterns regularly arrange in the ground tissue,such as Pteridium aquilinum (L.) Kuhn var.latiusculum (Desv.) Underw.ex Heller,all the ferns of Dryopteridaceae observed and Pyrrosia davidii (Baker) Ching.
5.The stomatas of the ferns observed are in the same plane with the epidermal cells with two guard cells;The mesophyll do not differentiate into palisade and spongy tissue except Sceptridium robustum (Rupr.) Lyon, Osmunda cinnamomea L.var.asiatica Fernald, Aleuritopteris argentea (Gmel.) Fee,Lepisorus ussuriensis (Regel et Maack) Ching and Pyrrosia davidii (Baker) Ching.
6.Compared with the ferns of Leptosporangiopsida,the plants of Equisetaceae,Sceptridium robustum (Rupr.) Lyon and Osmunda cinnamomea L.var.asiatica Fernald are more primitive on the morphological and anatomical characteristics.
1.木贼科问荆属2亚属植物表皮特征差别明显,问荆亚属植物表皮角质层成瘤状,气孔器不下陷,木贼表皮角质层不成瘤状,气孔器下陷；气孔器均为平列细胞型,由2个保卫细胞及2个副卫细胞构成,副卫细胞位于保卫细胞之上；气孔器上的辐射状纹饰来源于保卫细胞的腹壁(ventral wall),是细胞壁形成的片状凸起；所观察的木贼科5种植物主枝的基本结构一致,内皮层层数在2亚属植物中不稳定,不能将它作为分亚属的证据。
2.26种真蕨类植物叶表皮细胞多为不规则形,上、下表皮细胞垂周壁均呈波浪状,少数种类的上、下表皮细胞呈较规则的近长方形；共有6种气孔器类型,即极细胞型、腋下细胞型、不等细胞型、不规则四细胞型、不规则型和周细胞型,每种植物具1-4种不同类型的气孔器；气孔均为下生型气孔,一般散生,分布较均匀,并且沿着叶的长轴方向分布,只是偶尔在局部地方取向不规则,有时同一叶表皮的不同部分明显分布着不同类型的气孔。
3.蕨类植物鳞片的形状、网眼以及边缘的特征可以作为分类的依据之一
4.叶柄横切面显示,供试真蕨类植物维管束均为周韧型,可分为3种类型：(1)叶柄从基部向上至叶轴,维管束始终为一条,粗壮阴地蕨、桂皮紫萁、银粉背蕨、掌叶铁线蕨、过山蕨和耳羽岩蕨均属于这一类型；(2)叶柄基部具分体中柱2个,呈“八”字形排列,自叶柄基部向上,两个分体中柱于远轴面逐渐靠近,并连合成一个中柱,两束木质部远轴面末端的钩端相连,形成倒“V”字形至“人”字形,蹄盖蕨科供试的所有植物、卵果蕨、荚果蕨、球子蕨和乌苏里瓦韦均属于这一类型,除乌苏里瓦韦以外,其他植物木质部为“海马型”；(3)维管束多条,较规则的分散排列于基本组织之中,蕨、鳞毛蕨科供试的所有植物和有柄石韦属于这一类型。
5.供试真蕨类植物的气孔器均与表皮细胞在同一平面内,保卫细胞2个；多数植物叶肉没有分化为栅栏组织与海绵组织,可以明显的看出上表皮附近的同化组织比靠近下表皮的同化组织数量多,排列紧密；粗壮阴地蕨、桂皮紫萁、银粉背蕨、乌苏里瓦韦与有柄石韦同化组织均分化出栅栏组织与海绵组织。
6.与薄囊蕨纲的蕨类植物相比,木贼科植物、厚囊蕨纲的粗壮阴地蕨和原始薄囊蕨纲的桂皮紫萁的形态解剖学方面均表现出了较为原始的性状。
The epidermal structures, scales and cross section of 31 species from different families in Heilongjiang Province were investigated under light microscopes.The results indicated that:
1.The two subgenus of Equisetum shows apparent differences in the morphology of epidermis and stomatal position. Stomatas are both paracytic type and composed of two guard cells and two subsidiary cells,subsidiary cells located above the guard cells.Radial pattern of the stomatas derived from the apophysis on the ventral wall of the guard cells.The stem structure of five plants are similar,there are metaxylems in the vascular bundle.There are one or two layers of endodermis among the five species.
2.The leaf epidermal cells of 26 species of ferns in Heilongjiang are mostly irregular, the anticlinal walls of the upper and lower epidermal cells are wavy;There are six types of stomatal apparatus:Polocytic type,Axillocytic type,Aisocytic type,Anomotetracytic type,Anomocytic type and Pericytic type,each plant with one to four different types of stomatal apparatus;all the stomatal apparatuses only occur in the lower epidermis and distribution along the long axis of the leaves.
3.The shape, mesh and edge features of the ferns can be used as to classification.
4.The cross sections of the petioles show that the vascular bundle of the ferns investigated are all amphicribral,can be divided into three types:(1)There is only one vascular bundle in the petiole,such as Sceptridium robustum (Rupr.) Lyon,Osmunda cinnamomea L.var.asiatica Fernald,Aleuritopteris argentea (Gmel.) Fee,Adiantum pedatum L.,Camptosorus sibiricus Rupr.and Woodsia polystichoides Eaton;(2)The vascular patterns of the basal part of stipe are two meristeles distributing in the shape of "八",the xylems in the two meristeles are typical hippocampus type and gradually combine into amphicribral vascular bundles upwards, such as all the ferns of Athyriaceae observed,Phegopteris connectilis (Michx.) Watt,Matteuccia struthiopteris (L.) Todaro,Onoclea sensibilis L.var.interrupta Maxim.and Lepisorus ussuriensis (Regel et Maack) Ching;(3)Several vascular patterns regularly arrange in the ground tissue,such as Pteridium aquilinum (L.) Kuhn var.latiusculum (Desv.) Underw.ex Heller,all the ferns of Dryopteridaceae observed and Pyrrosia davidii (Baker) Ching.
5.The stomatas of the ferns observed are in the same plane with the epidermal cells with two guard cells;The mesophyll do not differentiate into palisade and spongy tissue except Sceptridium robustum (Rupr.) Lyon, Osmunda cinnamomea L.var.asiatica Fernald, Aleuritopteris argentea (Gmel.) Fee,Lepisorus ussuriensis (Regel et Maack) Ching and Pyrrosia davidii (Baker) Ching.
6.Compared with the ferns of Leptosporangiopsida,the plants of Equisetaceae,Sceptridium robustum (Rupr.) Lyon and Osmunda cinnamomea L.var.asiatica Fernald are more primitive on the morphological and anatomical characteristics.
引文
[1]陆树刚.蕨类植物学[M].北京：高等教育出版社,2007
[2]吴兆洪,秦仁昌.中国蕨类植物科属志[M].北京：科学出版社,1991
[3]吴国芳,冯志坚等.植物学下册[M].北京：高等教育出版社,1992
[4]敖志文,李国范.黑龙江省蕨类植物[M].哈尔滨：东北林业大学出版社,1990
[5]Lana Zoric, Ljiljana Merkulov.Leaf epidermal characteristics of Trifolium L. species from Serbia and Montenegro [J].Flora-Morphology, Distribution, Functional Ecology of Plants,2009,204(3):198-209
[6]M. Namaganda, T. Krekling, K.A. Lye.Leaf anatomical characteristics of Ugandan species of Festuca L. (Poaceae)[J].South African Journal of Botany,2009,75(1):52-59
[7]焦瑜,王士俊.三种国产阴地蕨科植物叶的表皮构造[J].植物学通报,2000,17(4)：352-359
[8]张耀甲,于海峰,卢云霞等.国产水龙骨科植物的气孔器类型及其系统学意义[J].兰州大学学报(自然科学版),1999,35(1)：130-139
[9]张丽兵.中国后生耳蕨组和新生耳蕨组(耳蕨属)的叶表皮解剖研究[J].应用与环境生物学报,1996,2(4)：361-368
[10]王秀华.东北鳞毛蕨科植物叶比较解剖的初步研究[J].植物研究,2001,21(2)：202-207
[11]戴锡玲,王全喜等.凤丫蕨属6种植物叶表皮特征的研究[J].植物研究,2009,29(1)33-38
[12]王玛丽,任毅.蹄盖蕨科植物叶表皮特征的比较形态学研究[J].西北植物学报,1997,17(5)：37-43
[13]徐皓.假冷蕨属的系统学研究[D].西北大学,2006
[14]李艳晖,刘斌等.山西产13种蕨类植物叶表皮特征的观察[J].山西大学学报(自然科学版),2004,27(4)：397-401
[15]焦瑜,陈立群,宋书银.七指蕨科和瓶尔小草科植物的叶表皮特征[J].植物研究,1999,19(2)：132-137
[16]巩乙南,刘星等.瓦韦属10种植物的气孔器类型及其系统学意义[J].北京师范大学学报(自然科学版),2006,42(3)：291-293
[17]王晓楠,石雷等.木贼科气孔器的扫描电镜观察[J].植物研究,2006,26(2)：147-150
[18]邢怡,党安志,刘保东.黑龙江产十种蕨类植物孢子囊和叶表皮的扫描电镜观察[J].植物研究,2004,24(4)：413-418
[19]刘家熙.北京蹄盖蕨属孢子形态的研究[J].贵州师范大学学报(自然科学版),1998,16(1)：23-25
[20]张玉龙.剑蕨属孢子形态的研究.植物学报[J],1963,11(1)：26-37
[21]张玉龙.书带蕨属孢子形态的研究.植物学报[J],1965,13(2)：151-158
[22]中国科学院植物研究所古植物研究室孢粉组.中国蕨类植物孢子形态[M].北京：科学出版社,1976
[23]黄增泉.台湾蕨类植物图志[M].台湾：仁和出版社,1981
[24]Wagner W.H.Jr.Structure of spores in relation to fern phylogeny[J].Ann Missouri Bot Gard,1974,61:332-353
[25]Tryon A.F.,Lugardon B.Spores of the Pteridophyta.Springer-Verlag.New York.1991
[26]Blanca Perez-Garcia, Aniceto Mendoza-Ruiz.Effect of light and temperature on germination of spores of four tropical fern species[J]. Acta Oecologica,2007,32(2):172-179
[27]Josef Psenicka, Jin Bek.Description of synangia and spores of the holotype of the Carboniferous fern Lobatopteris miltoni, with taxonomic comments[J].Review of Palaeobotany and Palynology,2009,155(3-4):133-144
[28]J.P. Ramos Giacosa, M.A. Morbelli, G.E. Giudice.Morphology and ultrastructure of the spores of the Grammitidaceae from Argentina[J].Review of Palaeobotany and Palynology,2007,143(3-4):155-166
[29]G.E. Giudice, M.A. Morbelli.Morphology and ultrastructure of the spores of Dennstaedtiaceae from North West Argentina[J].Review of Palaeobotany and Palynology,2006,141(3-4):245-257
[30]J.P. Ramos Giacosa, M.A. Morbelli, G.E. Giudice.Spore morphology and wall ultrastructure of Blechnum L. species from North West Argentina[J].Review of Palaeobotany and Palynology,2009,156(1-2):185-197
[31]刘家熙.北京产粉背蕨属孢子形态的研究[J].植物研究,1998,18(1)：28-30
[32]常缨,王全喜等.中国短肠蕨属孢子形态的研究[J].上海师范大学学报(自然科学版),2001,30(3)：56-62
[33]刘保东,包文美,敖志文.中国卷柏科孢子形态的研究[J].植物研究,1989,9(3)：113-131
[34]张刚明,张宪春.薄鳞蕨属(中国蕨科)植物的孢子形态研究[J].植物分类学报,2002,40(4)：345-350
[35]石雷.中国产瘤蕨属植物的孢子形态研究及其分类学意义[J].植物研究,2002,22(4)：428-432
[36]王玛丽,陆树刚,赵桂仿.中国介蕨属孢子形态的研究[J].植物分类学报,2003,40(5)：414--420
[37]戴锡玲,王全喜.中国蕨类植物孢子形态的研究Ⅷ.岩蕨科[J].西北植物学报,2007,27(11)：2210-2216
[38]戴锡玲,曹建国,王全喜.水蕨孢子壁的形成和发育[J].植物学通报,2008,25(1)：72-79
[39]戴绍军,高晶等.蕨类植物孢子与种子植物花粉萌发的比较[J].植物学通报,2008,25(2)：139-148
[40]王全喜,于晶.扫描电镜下真蕨目孢子表面纹饰的分类[J].云南植物研究,2003,25(3)：313-320
[41]张金谈.从孢粉形态特征试论植物某些类群的分类与系统发育[J].植物分类学报,1979,17(2)：1-7
[42]张金谈,王伏雄.孢粉形态研究进展.植物学通报[J],1988,5(2)：77-81
[43]郭庆梅,高慧,郑吉营等.山东产鳞毛蕨属植物形态解剖学的研究[J].纪念秦仁昌论文集,1999,316-327
[44]郭庆梅,周凤琴,李建秀.山东3种金星蕨科植物的比较解剖[J].山东师范大学学报(自然科学版),2007,22(2)：116-119
[45]郭庆梅,周凤琴,王汝新.山东产木贼科草药形态解剖学的研究[J].中草药.2000,31(9)：700-702
[46]孔稚颖,韩月莲等.山东产卷柏属植物形态解剖学研究[J].纪念秦仁昌论文集,1999,335-343
[47]周凤琴,高翠芳等.山东肿足蕨科形态解剖学及其在分类上的意义[J].纪念秦仁昌论文集,1999,357-369
[48]周凤琴,郭庆梅等.山东产蹄盖蕨科2属植物形态解剖学的研究[J].西北植物学报,2006,26(8)：1569-1574
[49]张宪春,刘全儒,徐静.宽带蕨属(水龙骨科)的系统学研究[J].植物分类学报,2003,41(5)：401-415
[50]郑玲.单叶双盖蕨属(蹄盖蕨科)的分类学研究[D].西北大学,2006
[51]王玛丽.蹄盖蕨科系统分类学研究[D].西北大学,2003
[52]David L. Des Marais, Alan R. Smith.Phylogenetic relationships and evolution of extant Horsetails,Equisetum,based on chloroplast DNA sequence data (rbcL AND trnL-F)[J]. Int. J. Plant Sci.,2003,164(5):737-751
[53]Emerson R. Pansarin, Antonio Salatino, Maria L.F. Salatino.Phylogeny of South American Pogonieae (Orchidaceae, Vanilloideae) based on sequences of nuclear ribosomal (ITS) and chloroplast (psaB, rbcL, rps16, and trnL-F) DNA, with emphasis on Cleistes and discussion of biogeographic implications[J].Organisms Diversity & Evolution,2008,8(3):171-181
[54]Warren D. Hauk, Clifford R. Parks, Mark W. Chase.Phylogenetic studies of Ophioglossaceae:evidence from rbcL and trnL-F plastid DNA sequences and morphology[J].Molecular Phylogenetics and Evolution,2003,28(1):131-151
[55]Chun-Xiang Li, Shu-Gang Lu, Qun Yang.Phylogeny and biogeography of Chinese and Australasian Polystichum ferns as inferred from chloroplast trnL-F and rps4-trnS sequence data[J].Palaeoworld,2007,16(4):294-300
[56]王玛丽,陈之端,张宪春等.蹄盖蕨科的系统发育：叶绿体DNA trnL-F区序列证据[J]. 植物分类学报,2003,41(5)：416-426
[57]张秀英,范亚文.黑龙江省蹄盖蕨科三属(蹄盖蕨属、羽节蕨属及冷蕨属)的遗传多样性分析[J].植物研究,2004,25(2)：186-191
[58]范亚文,张秀英,门晓宇.黑龙江省蹄盖蕨科(Athyriaceae)植物的RAPD分析及其系统学意义[J].植物研究,2008,28(2)：211-215
[59]林孝辉,王全喜,包文美,敖志文.东北蕨类植物配子体发育的研究XI：蹄盖蕨科[J].植物研究,1996,15(3)：322-340
[60]刘家熙,李雅轩.木贼科精子器比较研究[J].首都师范大学学报(自然科学版),1999,20(3)：62-65
[61]孟宪利,李勇,刘保东.叉蕨科4属5种植物配子体的发育模式及其系统学意义[J].植物学通报,2008,25(3)：298-306
[62]傅沛云.东北植物检索表第二版[M].北京：科学出版社,1995
[63]刘慎谔.东北草本植物志第一卷[M].北京：科学出版社,1958
[64]Tomasi V.H.,Rovasio R.A.Softening of plant specimenic (Equisetaceae) to improve the preparation of paraffin sections [J]. Biotech Histochem,1997,72(4):209
[65]Dilcher D.L.Approaches to the indentification of angiosperm leaf remains[J].Bot Rev, 1974,40(1):154-157
[66]Frvns-Claessens E, Van Cotterm W R J. A new classification of the ontogenetic types of stomata[J]. Bot Rev,1973,39(1):71-138
[67]Copeland E B. Genera filicum,Waltham,1947:247-257
[68]Hauke R L. The stomatal apparatus of Equisetum[J],Bull. Torrey Bot.Club,1957,84:178-181
[69]Johnson M A.Origin and development of tissues in Equisetum scirpoides[J]. Bot. Gaz.,1933,94:469-494
[70]王中仁等.中国植物志第三卷第二分册[M].北京：科学出版社,1999
[2]吴兆洪,秦仁昌.中国蕨类植物科属志[M].北京：科学出版社,1991
[3]吴国芳,冯志坚等.植物学下册[M].北京：高等教育出版社,1992
[4]敖志文,李国范.黑龙江省蕨类植物[M].哈尔滨：东北林业大学出版社,1990
[5]Lana Zoric, Ljiljana Merkulov.Leaf epidermal characteristics of Trifolium L. species from Serbia and Montenegro [J].Flora-Morphology, Distribution, Functional Ecology of Plants,2009,204(3):198-209
[6]M. Namaganda, T. Krekling, K.A. Lye.Leaf anatomical characteristics of Ugandan species of Festuca L. (Poaceae)[J].South African Journal of Botany,2009,75(1):52-59
[7]焦瑜,王士俊.三种国产阴地蕨科植物叶的表皮构造[J].植物学通报,2000,17(4)：352-359
[8]张耀甲,于海峰,卢云霞等.国产水龙骨科植物的气孔器类型及其系统学意义[J].兰州大学学报(自然科学版),1999,35(1)：130-139
[9]张丽兵.中国后生耳蕨组和新生耳蕨组(耳蕨属)的叶表皮解剖研究[J].应用与环境生物学报,1996,2(4)：361-368
[10]王秀华.东北鳞毛蕨科植物叶比较解剖的初步研究[J].植物研究,2001,21(2)：202-207
[11]戴锡玲,王全喜等.凤丫蕨属6种植物叶表皮特征的研究[J].植物研究,2009,29(1)33-38
[12]王玛丽,任毅.蹄盖蕨科植物叶表皮特征的比较形态学研究[J].西北植物学报,1997,17(5)：37-43
[13]徐皓.假冷蕨属的系统学研究[D].西北大学,2006
[14]李艳晖,刘斌等.山西产13种蕨类植物叶表皮特征的观察[J].山西大学学报(自然科学版),2004,27(4)：397-401
[15]焦瑜,陈立群,宋书银.七指蕨科和瓶尔小草科植物的叶表皮特征[J].植物研究,1999,19(2)：132-137
[16]巩乙南,刘星等.瓦韦属10种植物的气孔器类型及其系统学意义[J].北京师范大学学报(自然科学版),2006,42(3)：291-293
[17]王晓楠,石雷等.木贼科气孔器的扫描电镜观察[J].植物研究,2006,26(2)：147-150
[18]邢怡,党安志,刘保东.黑龙江产十种蕨类植物孢子囊和叶表皮的扫描电镜观察[J].植物研究,2004,24(4)：413-418
[19]刘家熙.北京蹄盖蕨属孢子形态的研究[J].贵州师范大学学报(自然科学版),1998,16(1)：23-25
[20]张玉龙.剑蕨属孢子形态的研究.植物学报[J],1963,11(1)：26-37
[21]张玉龙.书带蕨属孢子形态的研究.植物学报[J],1965,13(2)：151-158
[22]中国科学院植物研究所古植物研究室孢粉组.中国蕨类植物孢子形态[M].北京：科学出版社,1976
[23]黄增泉.台湾蕨类植物图志[M].台湾：仁和出版社,1981
[24]Wagner W.H.Jr.Structure of spores in relation to fern phylogeny[J].Ann Missouri Bot Gard,1974,61:332-353
[25]Tryon A.F.,Lugardon B.Spores of the Pteridophyta.Springer-Verlag.New York.1991
[26]Blanca Perez-Garcia, Aniceto Mendoza-Ruiz.Effect of light and temperature on germination of spores of four tropical fern species[J]. Acta Oecologica,2007,32(2):172-179
[27]Josef Psenicka, Jin Bek.Description of synangia and spores of the holotype of the Carboniferous fern Lobatopteris miltoni, with taxonomic comments[J].Review of Palaeobotany and Palynology,2009,155(3-4):133-144
[28]J.P. Ramos Giacosa, M.A. Morbelli, G.E. Giudice.Morphology and ultrastructure of the spores of the Grammitidaceae from Argentina[J].Review of Palaeobotany and Palynology,2007,143(3-4):155-166
[29]G.E. Giudice, M.A. Morbelli.Morphology and ultrastructure of the spores of Dennstaedtiaceae from North West Argentina[J].Review of Palaeobotany and Palynology,2006,141(3-4):245-257
[30]J.P. Ramos Giacosa, M.A. Morbelli, G.E. Giudice.Spore morphology and wall ultrastructure of Blechnum L. species from North West Argentina[J].Review of Palaeobotany and Palynology,2009,156(1-2):185-197
[31]刘家熙.北京产粉背蕨属孢子形态的研究[J].植物研究,1998,18(1)：28-30
[32]常缨,王全喜等.中国短肠蕨属孢子形态的研究[J].上海师范大学学报(自然科学版),2001,30(3)：56-62
[33]刘保东,包文美,敖志文.中国卷柏科孢子形态的研究[J].植物研究,1989,9(3)：113-131
[34]张刚明,张宪春.薄鳞蕨属(中国蕨科)植物的孢子形态研究[J].植物分类学报,2002,40(4)：345-350
[35]石雷.中国产瘤蕨属植物的孢子形态研究及其分类学意义[J].植物研究,2002,22(4)：428-432
[36]王玛丽,陆树刚,赵桂仿.中国介蕨属孢子形态的研究[J].植物分类学报,2003,40(5)：414--420
[37]戴锡玲,王全喜.中国蕨类植物孢子形态的研究Ⅷ.岩蕨科[J].西北植物学报,2007,27(11)：2210-2216
[38]戴锡玲,曹建国,王全喜.水蕨孢子壁的形成和发育[J].植物学通报,2008,25(1)：72-79
[39]戴绍军,高晶等.蕨类植物孢子与种子植物花粉萌发的比较[J].植物学通报,2008,25(2)：139-148
[40]王全喜,于晶.扫描电镜下真蕨目孢子表面纹饰的分类[J].云南植物研究,2003,25(3)：313-320
[41]张金谈.从孢粉形态特征试论植物某些类群的分类与系统发育[J].植物分类学报,1979,17(2)：1-7
[42]张金谈,王伏雄.孢粉形态研究进展.植物学通报[J],1988,5(2)：77-81
[43]郭庆梅,高慧,郑吉营等.山东产鳞毛蕨属植物形态解剖学的研究[J].纪念秦仁昌论文集,1999,316-327
[44]郭庆梅,周凤琴,李建秀.山东3种金星蕨科植物的比较解剖[J].山东师范大学学报(自然科学版),2007,22(2)：116-119
[45]郭庆梅,周凤琴,王汝新.山东产木贼科草药形态解剖学的研究[J].中草药.2000,31(9)：700-702
[46]孔稚颖,韩月莲等.山东产卷柏属植物形态解剖学研究[J].纪念秦仁昌论文集,1999,335-343
[47]周凤琴,高翠芳等.山东肿足蕨科形态解剖学及其在分类上的意义[J].纪念秦仁昌论文集,1999,357-369
[48]周凤琴,郭庆梅等.山东产蹄盖蕨科2属植物形态解剖学的研究[J].西北植物学报,2006,26(8)：1569-1574
[49]张宪春,刘全儒,徐静.宽带蕨属(水龙骨科)的系统学研究[J].植物分类学报,2003,41(5)：401-415
[50]郑玲.单叶双盖蕨属(蹄盖蕨科)的分类学研究[D].西北大学,2006
[51]王玛丽.蹄盖蕨科系统分类学研究[D].西北大学,2003
[52]David L. Des Marais, Alan R. Smith.Phylogenetic relationships and evolution of extant Horsetails,Equisetum,based on chloroplast DNA sequence data (rbcL AND trnL-F)[J]. Int. J. Plant Sci.,2003,164(5):737-751
[53]Emerson R. Pansarin, Antonio Salatino, Maria L.F. Salatino.Phylogeny of South American Pogonieae (Orchidaceae, Vanilloideae) based on sequences of nuclear ribosomal (ITS) and chloroplast (psaB, rbcL, rps16, and trnL-F) DNA, with emphasis on Cleistes and discussion of biogeographic implications[J].Organisms Diversity & Evolution,2008,8(3):171-181
[54]Warren D. Hauk, Clifford R. Parks, Mark W. Chase.Phylogenetic studies of Ophioglossaceae:evidence from rbcL and trnL-F plastid DNA sequences and morphology[J].Molecular Phylogenetics and Evolution,2003,28(1):131-151
[55]Chun-Xiang Li, Shu-Gang Lu, Qun Yang.Phylogeny and biogeography of Chinese and Australasian Polystichum ferns as inferred from chloroplast trnL-F and rps4-trnS sequence data[J].Palaeoworld,2007,16(4):294-300
[56]王玛丽,陈之端,张宪春等.蹄盖蕨科的系统发育：叶绿体DNA trnL-F区序列证据[J]. 植物分类学报,2003,41(5)：416-426
[57]张秀英,范亚文.黑龙江省蹄盖蕨科三属(蹄盖蕨属、羽节蕨属及冷蕨属)的遗传多样性分析[J].植物研究,2004,25(2)：186-191
[58]范亚文,张秀英,门晓宇.黑龙江省蹄盖蕨科(Athyriaceae)植物的RAPD分析及其系统学意义[J].植物研究,2008,28(2)：211-215
[59]林孝辉,王全喜,包文美,敖志文.东北蕨类植物配子体发育的研究XI：蹄盖蕨科[J].植物研究,1996,15(3)：322-340
[60]刘家熙,李雅轩.木贼科精子器比较研究[J].首都师范大学学报(自然科学版),1999,20(3)：62-65
[61]孟宪利,李勇,刘保东.叉蕨科4属5种植物配子体的发育模式及其系统学意义[J].植物学通报,2008,25(3)：298-306
[62]傅沛云.东北植物检索表第二版[M].北京：科学出版社,1995
[63]刘慎谔.东北草本植物志第一卷[M].北京：科学出版社,1958
[64]Tomasi V.H.,Rovasio R.A.Softening of plant specimenic (Equisetaceae) to improve the preparation of paraffin sections [J]. Biotech Histochem,1997,72(4):209
[65]Dilcher D.L.Approaches to the indentification of angiosperm leaf remains[J].Bot Rev, 1974,40(1):154-157
[66]Frvns-Claessens E, Van Cotterm W R J. A new classification of the ontogenetic types of stomata[J]. Bot Rev,1973,39(1):71-138
[67]Copeland E B. Genera filicum,Waltham,1947:247-257
[68]Hauke R L. The stomatal apparatus of Equisetum[J],Bull. Torrey Bot.Club,1957,84:178-181
[69]Johnson M A.Origin and development of tissues in Equisetum scirpoides[J]. Bot. Gaz.,1933,94:469-494
[70]王中仁等.中国植物志第三卷第二分册[M].北京：科学出版社,1999