title: Towards a phylogenetic generic classification of Thelypteridaceae: Additional sampling suggests alterations of neotropical taxa and further study of paleotropical genera.
abstract: Thelypteridaceae is one of the largest fern families, having about 950 species and a cosmopolitan distribution but with most species occurring in tropical and subtropical regions. Its generic classification remains controversial, with different authors recognizing from one up to 32 genera. Phylogenetic relationships within the family have not been exhaustively studied, but previous studies have confirmed the monophyly of the lineage. Thus far, sampling has been inadequate for establishing a robust hypothesis of infrafamilial relationships within the family. In order to understand phylogenetic relationships within Thelypteridaceae and thus to improve generic reclassification, we expand the molecular sampling, including new samples of Old World taxa and, especially, many additional neotropical representatives. We also explore the monophyly of exclusively or mostly neotropical genera Amauropelta, Goniopteris, Meniscium, and Steiropteris. Our sampling includes 68 taxa and 134 newly generated sequences from two plastid genomic regions (rps4-trnS and trnL-trnF), plus 73 rps4 and 72 trnL-trnF sequences from GenBank. These data resulted in a concatenated matrix of 1980 molecular characters for 149 taxa. The combined data set was analyzed using maximum parsimony and bayesian inference of phylogeny. Our results are consistent with the general topological structure found in previous studies, including two main lineages within the family: phegopteroid and thelypteroid. The thelypteroid lineage comprises two clades; one of these included the segregates Metathelypteris, Coryphopteris, and Amauropelta (including part of Parathelypteris), whereas the other comprises all segregates of Cyclosorus s.l., such as Goniopteris, Meniscium, and Steiropteris (including Thelypteris polypodioides, previously incertae sedis). The three mainly neotropical segregates were found to be monophyletic but nested in a broadly defined Cyclosorus. The fourth mainly neotropical segregate, Amauropelta, was found to include species considered to be part of Parathelypteris. In Old World thelypteroids, which correspond to nearly half the diversity in the family, an increase in sampling is still needed to resolve relationships and circumscription of genera, particularly in the christelloid clade (i.e., Amphineuron, Chingia, Christella, Pneumatopteris, Pronephrium, and Sphaerostephanos). Based on currently available knowledge, we propose the recognition of 16 genera in the family.
authors.affiliation: Departamento de Botânica, Universidade Federal de Minas Gerais, Av. Antônio Carlos, 6627 - Belo Horizonte, P.O. Box 486, Minas Gerais 30123-970, Brazil. Electronic address: blotiella@gmail.com.
authors.forename: Thaís Elias
authors.initials: TE
authors.lastname: Almeida
authors.affiliation: UMR 7205, CNRS, MNHN, UPMC, EPHE Institut de Systématique, Evolution, Biodiversité, 43 rue de Buffon, 75005 Paris, France; UMR 7207, CNRS, MNHN, UPMC, 'Centre de recherche sur la Paléobiodiversité et les Paléoenvironnements', 43 rue de Buffon, 75005 Paris, France.
authors.forename: Sabine
authors.initials: S
authors.lastname: Hennequin
authors.affiliation: Department of Life Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, England, United Kingdom.
authors.forename: Harald
authors.initials: H
authors.lastname: Schneider
authors.affiliation: The University Herbarium, University of California, Berkeley, 1001 Valley Life Sciences Bldg. #2465, Berkeley, CA 94720-2465, USA.
authors.forename: Alan R
authors.initials: AR
authors.lastname: Smith
authors.affiliation: Departamento de Botânica, Universidade Federal de Minas Gerais, Av. Antônio Carlos, 6627 - Belo Horizonte, P.O. Box 486, Minas Gerais 30123-970, Brazil.
authors.forename: João Aguiar Nogueira
authors.initials: JAN
authors.lastname: Batista
authors.affiliation: Departamento de Botânica, Universidade Federal de Minas Gerais, Av. Antônio Carlos, 6627 - Belo Horizonte, P.O. Box 486, Minas Gerais 30123-970, Brazil.
authors.forename: Aline Joseph
authors.initials: AJ
authors.lastname: Ramalho
authors.affiliation: Departamento de Botânica, Universidade Federal de Minas Gerais, Av. Antônio Carlos, 6627 - Belo Horizonte, P.O. Box 486, Minas Gerais 30123-970, Brazil.
authors.forename: Karina
authors.initials: K
authors.lastname: Proite
authors.affiliation: Departamento de Botânica, Universidade Federal de Minas Gerais, Av. Antônio Carlos, 6627 - Belo Horizonte, P.O. Box 486, Minas Gerais 30123-970, Brazil.
authors.forename: Alexandre
authors.initials: A
authors.lastname: Salino
mesh_terms: D001499:Bayes Theorem; D005075:Biological Evolution; D018742:DNA, Chloroplast / Q000235:genetics; D018744:DNA, Plant / Q000235:genetics*; D029624:Ferns / Q000145:classification* / Q000235:genetics; D010802:Phylogeny; D018087:Plastids / Q000235:genetics; D016415:Sequence Alignment; D017422:Sequence Analysis, DNA
chemical_list: D018742:DNA, Chloroplast; D018744:DNA, Plant
keywords: Ferns; Generic rearrangement; Neotropics; Paleotropics; Phylogeny