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

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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 E-mail:yhyan@sibs.ac.cn

Abstract:

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

Figure1

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"

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