Chinese Bulletin of Botany ›› 2020, Vol. 55 ›› Issue (2): 163-176.doi: 10.11983/CBB19118

• EXPERIMENTAL COMMUNICATIONS • Previous Articles     Next Articles

Morphology Characters and Evolution of Ferns Scale Ι: Pteridaceae

Gu Yufeng1,2,3,Jin Dongmei1,Liu Baodong2,Dai Xiling3,Yan Yuehong1,*()   

  1. 1 Shanghai Chenshan Plant Science Research Center, Shanghai Chenshan Botanical Garden, Chinese Academy of Sciences, Shanghai 201602, China
    2 Key Laboratory of Plant Biology, College of Heilongjiang Province, Harbin Normal University, Harbin 150025, China
    3 College of Life Sciences, Shanghai Normal University, Shanghai 200234, China
  • Received:2019-07-01 Accepted:2019-10-24 Online:2020-02-12 Published:2020-03-01
  • Contact: Yan Yuehong


Scales are epidermal appendages on the rhizomes and leaves of many ferns. Features of scales play an important role in classification of ferns. The phylogenetic position and delimitation of Pteridaceae were treated differently by different authors. Here we collected scales of 76 fern species in Pteridaceae, and observed them under a dissecting microscope. By comparing the morphological characters, we found that scales are different among genus and subfamilies. We reconstructed a phylogenetic tree with the plastid rbcL sequence of the species in this study downloaded from GenBank database and reconstruced the ancestral state for two selected characters (margin of the scale and mesh type). The results suggested that homogeneous scale and entire margin were plesiomorphic characters, while non-entire margin and transparent mesh were evolved late in evolutionary process. We also speculated that the formation of transparent or non-transparent mesh may relate to the light intensity in the habitats.

Key words: morphology and anatomy, classification, scales, phylogeny, Pteridaceae


The habitats of Pteridaceae (A) Coniogramme japonica; (B) Ceratopteris pteridoides; (C) Pteris amoena; (D) P. vittata; (E) Aleuritopteris argentea; (F) Pellaea nitidula; (G) Haplopteris flexuosa; (H) Adiantum capillus-veneris. Bars=20 cm"

Figure 2

Scale morphology of Cryptogrammoideae (A) Coniogramme caudiformis; (B) C. robusta; (C) C. emeiensis; (D) C. japonica; (E) C. jinggangshanensis; (F) C. intermedia; (G) C. wilsonii; (H) C. procera; (I) Cryptogramma brunoniana. Bars=0.5 mm"

Figure 3

Scale morphology of Ceratopteridoideae (A) Cystopteris fragilis; (B) Ceratopteris thalictroides. Bars=0.5 mm"

Figure 4

Scale morphology of Pteridoideae (A) Pteris vittata; (B) Pityrogramme calomelanos; (C) Anogramma microphylla; (D) Taenitis blechnoides; (E) Onychium siliculosum; (F) O. japonicum var. lucidum; (G) O. japonicum; (H) P. wallichiana; (I) P. cretica var. nervosa; (J) P. deltodon; (K) P. henryi; (L) P. gallinopes; (M) P. dactylina; (N) P. dispar; (O) P. amoena; (P) P. decrescens; (Q) P. cadieri; (R) P. insignis; (S) P. excelsa; (T) P. aspericaulis var. tricolor; (U) P. setuloso-costulata; (V) P. aspericaulis; (W) P. plumbea; (X) P. stenophylla. Bars=0.5 mm"

Figure 5

Scale morphology of Cheilanthoideae (A) Paragymnopteris marantae; (B) Pa. bipinnata; (C) Pa. vestita; (D) Pa. bipinnata var. auriculata; (E) Doryopteris concolor; (F) Hemionitis arifolia; (G) Notholaena hirsuta; (H) N. chinensis; (I) Cheilosoria tenuifolia; (J) Ch. chusana; (K) Ch. patula; (L) Aleuritopteris argentea; (M) Al. chrysophylla; (N) Al. pseudofarinosa; (O) Al. subvillosa var. tibetica; (P) Pellaea nitidula; (Q) P. mairei. Bars=0.5 mm"

Figure 6

Scale morphology of Vittarioideae (A) Monogramma paradoxa; (B) Antrophyum callifolium; (C) A. obovatum; (D) Haplopteris anguste-elongata; (E) H. amboinensis; (F) H. flexuosa; (G) H. plurisulcata; (H) H. doniana; (I) H. fudzinoi; (J) H. linearifolia; (K) Adiantum myriosorum; (L) Ad. flabellulatum; (M) Ad. capillus-veneris var. dissectum; (N) Ad. capillus-veneris; (O) Ad. erythrochlamys; (P) Ad. fengianum; (Q) Ad. fimbriatum; (R) Ad. philippense; (S) Ad. caudatum; (T) Ad. malesianum; (U) Ad. meishanianum; (V) Ad. juxtapositum; (W) Ad. gravesii; (X) Ad. mariesii. Bars=0.5 mm"

Figure 7

Phylogenetic tree of Pteridaceae obtained from the maximum likelihood analysis of the plastid rbcL sequence The BI analysis was performed in MrBayes 3.2.7 based on the best model Bayesian information criterion (BIC) implemented in jModelTest. Each chains was run for 1 000 000 generations, and was sampled every 1 000 generations. Numbers on branches indicate support values. A: Cryptogrammoideae; B: Ceratopteridoideae; C: Pteridoideae; D: Cheilanthoideae; E: Vittarioideae"

Figure 8

Ancestral state reconstruction for margin of scales and mesh type characters optimized onto the best tree obtained in the plateau phase of the Bayesian with the maximum likelihood (ML) analysis of plastid gene rbcL sequence Only species of Pteridaceae were used for the phylogeny. Pie charts show the percentage of node absence and the average likelihood received by each state across all the trees possessing that node. (L) Mesh type; (R) Margin of scales. A: Cryptogrammoideae; B: Ceratopteridoideae; C: Pteridoideae; D: Cheilanthoideae; E: Vittarioideae"

[1] 顾钰峰 ( 2015). 蕨类植物鳞片形态的研究. 硕士论文. 上海: 上海师范大学. pp. 7.
[2] 顾钰峰, 韦宏金, 卫然, 戴锡玲, 严岳鸿 ( 2014). 中国双盖蕨属一新记录种——Diplazium×kidoi Sa. Kurata. 植物科学学报 32, 336-339.
[3] 柯勇男 ( 2002). 台湾产广义鳞毛蕨科之毛被物研究. 硕士论文. 高雄: 台湾中山大学. pp. 3.
[4] 刘红梅, 王丽, 张宪春, 曾辉 ( 2008). 石松类和蕨类植物研究进展: 兼论国产类群的科级分类系统. 植物分类学报 46, 808-829.
[5] 秦仁昌 ( 1978). 中国蕨类植物科属的系统排列和历史来源. 植物分类学报 16(3), 1-19.
[6] 邵文, 陆树刚, 商清春 ( 2011). 假瘤蕨属(水龙骨科)植物鳞片特征的分类学意义. 广西植物 31, 14-19.
[7] 石雷, 张宪春 ( 1999). 薄唇蕨属的分类研究. 植物分类学报 37, 145-152.
[8] 石雷, 张宪春 ( 2001). 线蕨种下分类和植物地理的研究. 植物研究 21, 360-364.
[9] 于顺利, 林尤兴 ( 1996). 中国产瓦韦属植物的分类学研究. 植物研究 16, 3-24.
[10] 张宪春 ( 2012). 中国石松类和蕨类植物. 北京: 北京大学出版社. pp. 23.
[11] 张宪春, 卫然, 刘红梅, 何丽娟, 王丽, 张钢民 ( 2013). 中国现代石松类和蕨类的系统发育与分类系统. 植物学报 48, 119-137.
[12] Foster AS, Gifford EM (李正理, 张新英, 李荣敖, 崔克明译)( 1983). 维管植物比较形态学. 北京: 科学出版社. pp. 226-312.
[13] Christenhusz MJM, Chase MW ( 2014). Trends and concepts in fern classification. Ann Bot 113, 571-594.
[14] Christenhusz MJM, Zhang XC, Schneider H ( 2011). A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19, 7-54.
[15] Greenfield SS ( 1938). A comparison of the basal scales and Indusia of Dryopteris Goldiana × marginalis with those of its parent species. Am Fern J 28, 55-62.
[16] Holttum ER ( 1957). The scales of Cyatheaceae (With special reference to the genus Schizocaena J.Sm.). Kew Bull 12, 41-45.
[17] Holttum RE ( 1949). The classification of ferns. Biol Rev 24, 267-296.
[18] Holttum RE ( 1963). Cyatheaceae. Flora malesiana—series 2. Pteridophyta 1, 65-176.
[19] Hoshizaki BJ ( 1970). The rhizome scales of Platycerium. Am Fern J 60, 144-160.
[20] Kuo LY, Li FW, Chiou WL, Wang CN ( 2011). First insights into fern matK phylogeny. Mol Phylogenet Evol 59, 556-566.
[21] Labiak PH ( 2003). A new combination in the fern genus Ceradenia(Grammitidaceae). Kew Bull 58, 991-994.
[22] Lehtonen S ( 2011). Towards resolving the complete fern tree of life. PLoS One 6, e24851.
[23] Maddison DR, Maddison WP ( 2018). Mesquite: a modular system for evolutionary analysis. Version 3.61.http://www. .
[24] Nayar BK ( 1970). A phylogenetic classification of the homosporous ferns. Taxon 19, 229-236.
[25] PPG I ( 2016). A community-derived classification for extant lycophytes and ferns. J Syst Evol 54, 563-603.
[26] Pryer KM, Schuettpelz E, Wolf PG, Schneider H, Smith AR, Cranfill R ( 2004). Phylogeny and evolution of ferns (monilophytes) with a focus on the early leptosporangiate divergences. Am J Bot 91, 1582-1598.
[27] Qi XP, Kuo LY, Guo CC, Li H, Li ZY, Qi J, Wang LB, Hu Y, Xiang JY, Zhang CF, Guo J, Huang CH, Ma H ( 2018). A well-resolved fern nuclear phylogeny reveals the evolution history of numerous transcription factor families. Mol Phylogenet Evol 127, 961-977.
[28] Rai HS, Graham SW ( 2010). Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (monilophytes). Am J Bot 97, 1444-1456.
[29] Schneider H ( 2013). Evolutionary morphology of ferns (monilophytes). In: Ambrose BA, Purugganan M, eds. Annual Plant Reviews, Vol. 45. The Evolution of Plant Form. Hoboken, New Jersey: John Wiley & Sons, Ltd. pp. 115-140.
[30] Schuettpelz E, Schneider H, Huiet L, Windham MD, Pryer KM ( 2007). A molecular phylogeny of the fern family Pteridaceae: assessing overall relationships and the affinities of previously unsampled genera. Mol Phylogenet Evol 44, 1172-1185.
[31] Sermolli REGP ( 1977). Tentamen Pteridophytorum genera in taxonomicum ordinem redigendi. Webbia 31, 313-512.
[32] Shen H, Jin DM, Shu JP, Zhou XL, Lei M, Wei R, Shang H, Wei HJ, Zhang R, Liu L, Gu YF, Zhang XC, Yan YH ( 2018). Large-scale phylogenomic analysis resolves a backbone phylogeny in ferns. Gigascience 7(2), 1-11.
[33] Smith AR, Pryer KM, Schuettpelz E, Korall P, Schneider H, Wolf PG ( 2006). A classification for extant ferns. Taxon 55, 705-731.
[34] Tryon RM, Tryon AF ( 1982). Ferns and Allied Plants. New York: Springer- Verlag. pp. 166-212.
[35] Tsutsumi C, Kato M ( 2008). Morphology and evolution of epiphytic Davalliaceae scales. Botany 86, 1393-1403.
[36] Tsutsumi C, Zhang XC, Kato M ( 2008). Molecular phylogeny of Davalliaceae and implications for generic classification. Syst Bot 33, 44-48.
[37] Van den Heede CJ, Viane RLL, Chase MW ( 2003). Phylogenetic analysis of Asplenium subgenus Ceterach (Pteridophyta: Aspleniaceae) based on plastid and nuclear ribosomal ITS DNA sequences. Am J Bot 90, 481-495.
[38] Watkins Jr JE, Kawahara AY, Leicht TA, Auld JR, Bicksler AJ, Kaiser K ( 2006). Fern laminar scales protect against photoinhibition from excess light. Am Fern J 96, 83-92.
[39] Wei R, Schneider H, Zhang XC ( 2013). Toward a new circumscription of the twinsorus-fern genus (Diplazium Athyriaceae): a molecular phylogeny with morphological implications and infrageneric taxonomy. Taxon 62, 441-457.
[40] Wolf PG, Robison TA, Johnson MG, Sundue MA, Testo WL, Rothfels CJ ( 2018). Target sequence capture of nuclear-encoded genes for phylogenetic analysis in ferns. Appl Plant Sci 6, e01148.
[41] Zhang GM, Liao WB, Ding MY, Lin YX, Wu ZH, Zhang XC, Dong SY, Prado J, Gilbert MG, Yatskievych G, Ranker TA, Hoope EA, Alverson ER, Metzgar JS, Funston AM, Masuyama S, Kato M ( 2013). Flora of China,Vol. 2-3. Beijing: Science Press; St. Louis: Missouri Botanical Garden Press. pp. 169-256.
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[1] Yan Xiao-hua Cai Zhu-ping. Effects of S-07, PP333 and Triadimefon on Peroxidaseisoentyme of Rice Seedling[J]. Chinese Bulletin of Botany, 1995, 12(专辑3): 109 -112 .
[2] . [J]. Chinese Bulletin of Botany, 1994, 11(专辑): 13 .
[3] Xiaomin Yu;Xingguo Lan;Yuhua Li. The Ub/26S Proteasome Pathway and Self-incompatible Responses in Flowering Plants[J]. Chinese Bulletin of Botany, 2006, 23(2): 197 -206 .
[4] WANG Ling-Li LIU Wen-Zhe. Contents of Camptothecin in Camptotheca acuminata from Different Provenances[J]. Chinese Bulletin of Botany, 2005, 22(05): 584 -589 .
[5] Dai Yun-ling and Xu Chun-hui. Advances in Research on Protein Components of Oxygen-evolving Complex[J]. Chinese Bulletin of Botany, 1992, 9(03): 1 -16 .
[6] . Advances in Research on Photosynthesis of Submerged Macrophytes[J]. Chinese Bulletin of Botany, 2005, 22(增刊): 128 -138 .
[7] Shaobin Zhang;Guoqin Liu. Research Advances in Plant Actin Isoforms[J]. Chinese Bulletin of Botany, 2006, 23(3): 242 -248 .
[9] MA Li-Hui, WU Pu-Te, and WANG You-Ke. Spatial pattern of root systems of dense jujube plantation with jujube age in the semiarid loess hilly region of China[J]. Chin J Plan Ecolo, 2012, 36(4): 292 -301 .
[10] PAN Yu-De, Melillo J. M., Kicklighter D. W., XIAO Xiang-Ming, McGuire A. D.. Modeling Structural and Functional Responses of Terrestria Ecosystems in China to Changes in Climate and Atmospheric CO2[J]. Chin J Plan Ecolo, 2001, 25(2): 175 -189 .