Yi-Shan Chao

Molecular phylogeny and biogeography of the fern genus Pteris (Pteridaceae)

BACKGROUND AND AIMS Pteris (Pteridaceae), comprising over 250 species, had been thought to be a monophyletic genus until the three monotypic genera Neurocallis, Ochropteris and Platyzoma were included. However, the relationships between the type species of the genus Pteris, P. longifolia, and other species are still unknown. Furthermore, several infrageneric morphological classifications have been proposed, but are debated. To date, no worldwide phylogenetic hypothesis has been proposed for the genus, and no comprehensive biogeographical history of Pteris, crucial to understanding its cosmopolitan distribution, has been presented. METHODS A molecular phylogeny of Pteris is presented for 135 species, based on cpDNA rbcL and matK and using maximum parsimony, maximum likelihood and Bayesian inference approaches. The inferred phylogeny was used to assess the biogeographical history of Pteris and to reconstruct the evolution of one ecological and four morphological characters commonly used for infrageneric classifications. KEY RESULTS The monophyly of Pteris remains uncertain, especially regarding the relationship of Pteris with Actiniopteris + Onychium and Platyzoma. Pteris comprises 11 clades supported by combinations of ecological and morphological character states, but none of the characters used in previous classifications were found to be exclusive synapomorphies. The results indicate that Pteris diversified around 47 million years ago, and when species colonized new geographical areas they generated new lineages, which are associated with morphological character transitions. CONCLUSIONS This first phylogeny of Pteris on a global scale and including more than half of the diversity of the genus should contribute to a new, more reliable infrageneric classification of Pteris, based not only on a few morphological characters but also on ecological traits and geographical distribution. The inferred biogeographical history highlights long-distance dispersal as a major process shaping the worldwide distribution of the species. Colonization of different niches was followed by subsequent morphological diversification. Dispersal events followed by allopatric and parapatric speciation contribute to the species diversity of Pteris.

Extreme Multiple Reticulate Origins of the Pteris cadieri Complex (Pteridaceae)

The Pteris cadieri complex displays extensive morphological variation and seems to have originated through hybridization. However, the members of this complex reproduce by apogamy, which usually limits genetic variation. To evaluate the hypotheses of hybrid origins, the pattern of evolution in this species complex is reconstructed. Multiple methodologies were used. Diploids, triploids, and tetraploids were identified by chromosome counts and flow cytometry. Nuclear DNA markers (cytosolic phosphoglucose isomerase gene, PgiC) were used, together with chloroplast DNA markers (atpB-rbcL spacer and rbcL gene) to infer the biparental and maternal lineages of the Pteris cadieri complex. The three cpDNA haplotype groups and five PgiC alleles found in this study indicate that the evolution of the Pteris cadieri complex has been extremely reticulate. Up to 11 taxa belonging to eight morphs were identified. By comparing genetic variation in the Pteris cadieri in two independent areas, Hainan and Taiwan, we inferred that hybridization has occurred independently in different areas. Furthermore, we found evidence for phenological divergence (evergreen and deciduous) within Taiwan. We propose that the Pteris cadieri complex originated from different genetic lineages through multiple hybridizations in different geographical areas, leading to its present morphological diversity.

Phylogeny and character evolution of the fern genus Tectaria (Tectariaceae) in the Old World inferred from chloroplast DNA sequences

In this study we provide a phylogeny for the pantropical fern genus Tectaria, with emphasis on the Old World species, based on sequences of five plastid regions (atpB, ndhF plus ndhF-trnL, rbcL, rps16-matK plus matK, and trnL-F). Maximum parsimony, maximum likelihood, and Bayesian inference are used to analyze 115 individuals, representing ca. 56 species of Tectaria s.l. and 36 species of ten related genera. The results strongly support the monophyly of Tectaria in a broad sense, in which Ctenitopsis, Hemigramma, Heterogonium, Psomiocarpa, Quercifilix, Stenosemia, and Tectaridium should be submerged. Such broadly circumscribed Tectaria is supported by the arising pattern of veinlets and the base chromosome number (x=40). Four primary clades are well resolved within Tectaria, one from the Neotropic (T. trifoliata clade) and three from the Old World (T. subtriphylla clade, Ctenitopsis clade, and T. crenata clade). Tectaria crenata clade is the largest one including six subclades. Of the genera previously recognized as tectarioid ferns, Ctenitis, Lastreopsis, and Pleocnemia, are confirmed to be members in Dryopteridaceae; while Pteridrys and Triplophyllum are supported in Tectariaceae. To infer morphological evolution, 13 commonly used characters are optimized on the resulting phylogenetic trees and in result, are all homoplastic in Tectaria.

Polyploidy and Speciation in Pteris (Pteridaceae)

The highest frequency of polyploidy among plants is considered to occur in the Pteridophytes. In this study, we focused on polyploidy displayed by a specific fern taxon, the genus Pteris L. (Pteridaceae), comprising over 250 species. Cytological data from 106 Pteris species were reviewed. The base number of chromosomes in Pteris is 29. Polyploids are frequently found in Pteris, including triploids, tetraploids, pentaploids, hexaploids, and octoploids. In addition, an aneuploid species, P. deltodon Bak., has been recorded. Furthermore, the relationship between polyploidy and reproductive biology is reviewed. Among these 106 Pteris species, 60% exhibit polyploidy: 22% show intraspecific polyploidy and 38% result from polyploid speciation. Apogamous species are common in Pteris. Diploids are the most frequent among Pteris species, and they can be sexual or apogamous. Triploids are apogamous; tetraploids are sexual or apogamous. Most Pteris species have one to two ploidy levels. The diverse ploidy levels suggest that these species have a complex evolutionary history and their taxonomic problems require further clarification.

Microsatellite primers in the native perennial cycad Cycas taitungensis (Cycadaceae)

PREMISE OF THE STUDY Microsatellite primers were developed for the native perennial cycad Cycas taitungensis to evaluate the genetic variation of this endangered insular species. METHODS AND RESULTS Using a magnetic bead enrichment method and EST data, 16 primer sets were developed and identified for the native Taiwan cycad C. taitungensis. The primers amplified dinucleotide, trinucleotide, and complex repeats with 1-9 alleles per locus. Most primers also amplified DNA from C. revoluta and C. debaoensis. CONCLUSIONS These results indicate the utility of primers for future studies of the genetic structure of C. taitungensis. In addition, the primers are useful for further phylogeographic studies between C. taitungensis and C. revoluta, which is a closely related species.

New insights on the phylogeny of Tectaria (Tectariaceae), with special reference to Polydictyum as a distinct lineage

The fern genus Tectaria (Tectariaceae) Cav. is morphologically diverse and difficult in terms of recognizing species and species groups. To infer the systematic positions of some species and identity‐unknown collections with special morphological characters, we undertook phylogenetic analyses based on sequences of five plastid regions (atpB, ndhF + ndhF‐trnL, rbcL, rps16‐matK + matK, and trnL‐F). Three analysis methods (maximum parsimony, maximum likelihood, and Bayesian inference) were used to reconstruct the phylogeny of Tectaria. The most surprising result is that T. menyanthidis (C. Presl) Copel., T. ternata (Baker) Copel., and T. variabilis Tardieu & Ching are revealed to represent a distinct lineage from Tectaria, which should be called Polydictyum C. Presl, and is supported as sister to Pteridrys C. Chr. & Ching. Other accessions of Tectaria are well resolved into four major clades, which is consistent with the results of previous studies. Of the four clades, Clade II (T. subtriphylla (Hook. & Arn.) Copel. group) is unpredictable, with morphologically very diverse species clustered there, and is supposed to be a minor evolutionary line within Tectaria in the Old World. In addition, the position of the climbing genus Arthropteris J. Sm. and the utility of molecular data in recognizing species of Tectaria are briefly discussed. As a conclusion, we formally reinstate the genus Polydictyum by providing diagnostic characters, key to species, nomenclature, and information of detailed distribution and habitat for the currently known three species.

Pteris borneensis (Pteridaceae), a New Species from Borneo, with Re-Circumscription of Pteris decrescens and Pteris Parviloba

Abstract A new species was collected during a fern expedition in the Nuluhon Trusmadi Forest Reserve, Sabah, Borneo. Morphologically, it is similar to both Pteris decrescens Christ and P. parviloba Christ but can be distinguished by frond characters. Specifically, the new species differs from the other two species by having basiscopic secondary pinnae (vs. without basiscopic secondary pinnae), petiolulate pinnae (vs. sessile pinnae), without bristles on the stipes (vs. with bristles on the stipes), a crenate margin of the pinna-segment apex (vs. a serrate apex), and entire pseudoindusia (vs. ciliate pseudoindusia). Furthermore, P. decrescens and P. parviloba differ in the number of lateral pinnae (5 or 6 vs. 8–16 pairs) and the length of the basal pinna-segments being shortest or longest of all lateral pinnae. Two plastid genes, rbcL and matK, were used to decipher their phylogenetic relationships. The new species was resolved in different clades from P. decrescens and P. parviloba, providing further support of its uniqueness. The morphology, illustrations, ecology, and distribution of the new species are provided. We also found that the type materials of P. decrescens included two different species, P. decrescens and P. parviloba.A lectotype of P. decrescens was therefore designated. Based on the results of this study, P. decrescens and P. parviloba are re-circumscribed, and the new species is described as P. borneensis sp. nov.

Pteris latipinna sp. nov. (Pteridaceae), a new species segregated from Pteris fauriei

Abstract Pteris fauriei is widely distributed in Eastern Asia and has high morphological variation. Some morphologically similar plants related to this species are difficult to distinguish. We showed that the new Pteris species from Taiwan, previously identified as P. fauriei, can be morphologically distinguished by its wide pinnae, larger terminal pinnae than the lateral pinnae in sterile fronds, and triangular basal segments of the lateral pinnae. It was confirmed that this species is phylogenetically separated from the other East Asian Pteris species, except for a morphologically distinct species P. arisanensis, by means of chloroplast genes, rbcL and matK. The new species is named as Pteris latipinna sp. nov., referring to its wide pinnae. Here, we provide a key to facilitate the identification of the morphologically similar Pteris species in Asia. The morphological descriptions, images, ecology, and distribution are also presented.

New Species of the Fern Genus Lindsaea (Lindsaeaceae) from New Guinea with Notes on the Phylogeny of L. sect. Synaphlebium

To determine the taxonomic identities and the systematic positions of some collections of Lindsaea sect. Synaphlebium (Lindsaeaceae) from Papua New Guinea, we conducted morphological comparisons and phylogenetic analyses on the whole section. A total of 22 morphological characters were selected and coded for each of all known taxa in L. sect. Synaphlebium, and were analyzed using maximum parsimony. The datasets containing either of or combined two plastid DNA sequences (trnL-trnF spacer and trnH-psbA spacer) of 37 taxa were analyzed using maximum parsimony, maximum likelihood, and Bayesian inference. Morphological comparisons revealed two new species which are formally published here as L. subobscura and L. novoguineensis. Lindsaea subobscura is similar to sympatric L. obscura and L. modesta but differs in the obviously reduced upper pinnules and other characters. Lindsaea novoguineensis is most similar to L. pacifica from Melanesia but differs in having rhomboid pinnules with truncate apices and concave soral receptacles. Molecular analyses resolved L. sect. Synaphlebium and allied species into five well-supported clades, namely L. rigida clade, L. obtusa clade, L. pulchella clade, L. multisora clade, and L. cultrata clade. The new species L. novoguineensis is included in L. obtusa clade; L. subobscura is in L. pulchella clade; whereas the majority of L. sect. Synaphlebium is clustered in L. cultrata clade. As the section Synaphlebium sensu Kramer is strongly suggested as polyphyletic, we propose the concept of a monophyletic L. sect. Synaphlebium in a broad sense that comprises five lineages. The morphological circumscription of L. sect. Synaphlebium sensu lato and the divergence in morphology, habit, and distribution between the five lineages are briefly discussed. Further molecular study is needed to test the systematic positions of 16 other species which are supposed to be within L. sect. Synaphlebium sensu lato but have not been included in this and previous molecular analyses.