title: Leaf evolution in early-diverging ferns: insights from a new fern-like plant from the Late Devonian of China.
abstract: OBJECTIVE
  With the exception of angiosperms, the main euphyllophyte lineages (i.e. ferns sensu lato, progymnosperms and gymnosperms) had evolved laminate leaves by the Late Devonian. The evolution of laminate leaves, however, remains unclear for early-diverging ferns, largely represented by fern-like plants. This study presents a novel fern-like taxon with pinnules, which provides new insights into the early evolution of laminate leaves in early-diverging ferns.
  
  
  METHODS
  Macrofossil specimens were collected from the Upper Devonian (Famennian) Wutong Formation of Anhui and Jiangsu Provinces, South China. A standard degagement technique was employed to uncover compressed plant portions within the rock matrix.
  
  
  RESULTS
  A new fern-like taxon, SHOUGANGIA BELLA GEN ET SP NOV: , is described and represents an early-diverging fern with highly derived features. It has a partially creeping stem with adventitious roots only on one side, upright primary and secondary branches arranged in helices, tertiary branches borne alternately or (sub)oppositely, laminate and usually lobed leaves with divergent veins, and complex fertile organs terminating tertiary branches and possessing multiple divisions and numerous terminal sporangia.
  
  
  CONCLUSIONS
  Shougangia bella provides unequivocal fossil evidence for laminate leaves in early-diverging ferns. It suggests that fern-like plants, along with other euphyllophyte lineages, had independently evolved megaphylls by the Late Devonian, possibly in response to a significant decline in atmospheric CO2 concentration. Among fern-like plants, planate ultimate appendages are homologous with laminate pinnules, and in the evolution of megaphylls, fertile organs tend to become complex.
authors.affiliation: Key Laboratory of Orogenic Belts and Crustal Evolution, Department of Geology, Peking University, Beijing 100871, China, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China and State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China dmwang@pku.edu.cn hhxu@nigpas.ac.cn.
authors.forename: De-Ming
authors.initials: DM
authors.lastname: Wang
authors.affiliation: Key Laboratory of Orogenic Belts and Crustal Evolution, Department of Geology, Peking University, Beijing 100871, China, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China and State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China dmwang@pku.edu.cn hhxu@nigpas.ac.cn.
authors.forename: Hong-He
authors.initials: HH
authors.lastname: Xu
authors.affiliation: Key Laboratory of Orogenic Belts and Crustal Evolution, Department of Geology, Peking University, Beijing 100871, China, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China and State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China.
authors.forename: Jin-Zhuang
authors.initials: JZ
authors.lastname: Xue
authors.affiliation: Key Laboratory of Orogenic Belts and Crustal Evolution, Department of Geology, Peking University, Beijing 100871, China, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China and State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China.
authors.forename: Qi
authors.initials: Q
authors.lastname: Wang
authors.affiliation: Key Laboratory of Orogenic Belts and Crustal Evolution, Department of Geology, Peking University, Beijing 100871, China, State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China and State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China.
authors.forename: Le
authors.initials: L
authors.lastname: Liu
mesh_terms: D005075:Biological Evolution; D002681:China; D029624:Ferns / Q000033:anatomy & histology* / Q000145:classification; D005580:Fossils / Q000033:anatomy & histology*; D010802:Phylogeny; D018515:Plant Leaves / Q000033:anatomy & histology*
keywords: Adventitious root; Late Devonian; Shougangia bella; Wutong Formation; atmospheric CO2; euphyllophytes; fern-like plants; ferns sensu lato; fertile organ; growth habit; leaf evolution; monilophyte; stem