Phylogeny and evolution of grammitid ferns (Grammitidaceae): a case of rampant morphological homoplasy

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Abstract
We conducted phylogenetic analyses of the fern family Grammitidaceae using sequences from two cpDNA genes and from morphological characters. Data were obtained for 73 species from most recognized genera in the family. The genera Adenophorus, Ceradenia, Calymmodon, Cochlidium, Enterosora, and Melpomene were each strongly supported as being monophyletic. Other recognized genera that were not supported as monophyletic included Ctenopteris, Grammitis, Lellingeria, Micropolypodium, Prosaptia, and Terpsichore. Several previously unrecognized clades were identified, some of which are characterized by distinctive morphological features. Analyses of the distribution of morphological character states on our inferred phylogeny showed extremely high levels of homoplastic evolution for many different characters. Homoplasy for morphological characters was considerably greater than for molecular characters. Many of the characters that exhibited high levels of convergent or parallel evolution across the phylogeny are features that have been commonly used to circumscribe genera in this group (e.g., leaf blade dissection, various rhizome scale characters, and glandular paraphyses). Conversely, some of the characters that exhibited relatively low levels of homoplasy have either not been regarded as having taxonomic value or have been ignored (e.g., root insertion, rhizome scale sheen). Our data support a New World origin of Grammitidaceae, with Old World taxa generally being more evolutionarily derived. Several clades are either primarily Neotropical or primarily Paleotropical but also have a few members distributed in the opposite hemisphere. Thus, we postulate multiple, independent dispersal and colonization events in several lineages.