BackgroundThe fern genus Dryopteris (Dryopteridaceae) is among the most common and species rich fern genera in temperate forests in the northern hemisphere containing 225–300 species worldwide. The circumscription of Dryopteris has been controversial and various related genera have, over the time, been included in and excluded from Dryopteris. The infrageneric phylogeny has largely remained unclear, and the placement of the majority of the supraspecific taxa of Dryopteris has never been tested using molecular data.ResultsIn this study, DNA sequences of four plastid loci (rbcL gene, rps4-trnS spacer, trnL intron, trnL-F spacer) were used to reconstruct the phylogeny of Dryopteris. A total of 122 accessions are sampled in our analysis and they represent 100 species of the expanded Dryopteris including Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema. All four subgenera and 19 sections currently recognized in Dryopteris s.s. are included. One species each of Arachniodes, Leptorumohra, and Lithostegia of Dryopteridaceae are used as outgroups. Our study confirms the paraphyly of Dryopteris and provides the first strong molecular evidence on the monophyly of Acrophorus, Diacalpe, Dryopsis, Nothoperanema, and Peranema. However, all these monophyletic groups together with the paraphyletic Acrorumohra are suggested to be merged into Dryopteris based on both molecular and morphological evidence. Our analysis identified 13 well-supported monophyletic groups. Each of the 13 clades is additionally supported by morphological synapomophies and is inferred to represent a major evolutionary lineage in Dryopteris. In contrast, monophyly of the four subgenera and 15 out of 19 sections currently recognized in Dryopteris s.s is not supported by plastid data.ConclusionsThe genera, Acrophorus, Acrorumohra, Diacalpe, Dryopsis, Nothoperanema, and Peranema, should all be merged into Dryopteris. Most species of these genera share a short rhizome and catadromic arrangement of frond segments, unlike the sister genus of Dryopteris s.l., Arachniodes, which has anadromic arrangement of frond segments. The non-monophyly of the 19 out of the 21 supraspecific taxa (sections, subgenera) in Dryopteris strongly suggests that the current taxonomy of this genus is in need of revision. The disagreement between the previous taxonomy and molecular results in Dryopteris may be due partly to interspecific hybridization and polyplodization. More morphological studies and molecular data, especially from the nuclear genome, are needed to thoroughly elucidate the evolutionary history of Dryopteris. The 13 well-supported clades identified based on our data represent 13 major evolutionary lineages in Dryopteris that are also supported by morphological synapomophies.
The Asian keelback snakes (genus Amphiesma) are a widely distributed group of Old World natricines, inhabiting a variety of niches and exhibiting significant morphological variation. Recent molecular phylogenies suggest that this genus is not monophyletic, and that additional cryptic diversity is also likely present. We conducted a phylogenetic analysis of the group based on 3162 bp of one mitochondrial gene (Cyt. b) and three nuclear genes (C-mos, Rag1, NT3), sampling 18 species in addition to those sequenced in previous works. All analyses consistently show that Amphiesma consists of three distinct, monophyletic lineages with strong support. We divide Amphiesma into three genera, Amphiesma, Hebius, and Herpetoreas. The genus Amphiesma is monotypic, Herpetoreas contains three species, and Hebius comprises the remaining 39 species. On the basis of a combination of molecular analyses and external morphological comparisons, we describe a new species in the Herpetoreas group from China as H. burbrinki sp. nov. Several other species are shown to be non-monophyletic or contain significant levels of intraspecific genetic diversity. Another Old World natricine genera, Xenochrophis is also found to be non-monophyletic. Our results indicate that further taxonomic revisions are needed in Natricinae, at multiple levels.
Grammitidoideae are the largest subfamily in Polypodiaceae and contain about 911 species. Progress has been made in understanding the overall phylogeny and generic boundaries in the light of recent molecular works. However, the majority of species, especially Asian species, and some critical type species of genera remain unsampled. In this study, a dataset of six plastid markers of 1003 (112 new) accessions representing ca. 412 species of Grammitidoideae including the type species of Ctenopterella, Grammitis, Moranopteris, Radiogrammitis, and Themelium, was assembled to infer a phylogeny. Our major results include: (1) the type species of Grammitis is successfully sequenced using a next‐generation sequencing technique and is resolved in Grammitis s.str. as expected; (2) Ctenopterella is found to be polyphyletic and a new clade consisting of C. khaoluangensis is resolved as sister to Tomophyllum; (3) the type species of Ctenopterella is resolved in a clade sister to the C. lasiostipes clade; (4) Oreogrammitis is found to be polyphyletic and three clades outside of the core Oreogrammitis are identified containing O. subevenosa and allies, O. orientalis, and O. beddomeana (+ O. cf. beddomeana); (5) Prosaptia is found to be paraphyletic with P. nutans being sister to a clade containing the rest of Prosaptia and Archigrammitis; (6) the intergeneric and major relationships within the Asia‐Pacific clade are well resolved and strongly supported except for a few branches; (7) extensive cryptic speciation is detected in the Asia‐Pacific clade; and (8) based on the polyphyly of Ctenopterella we describe three new genera, Boonkerdia, Oxygrammitis, and Rouhania, for species formerly in Ctenopterella; because the type species of Grammitis belongs to Grammitis s.str., we describe five new genera, Aenigmatogrammitis, Grammitastrum (stat. nov.), Howeogrammitis, Nanogrammitis, and Thalassogrammitis for species formerly in Grammitis s.l. A key to the 35 Old‐World genera is given, a taxonomic treatment is presented, and the morphology of all new genera is shown with either a color plate and/or a line drawing.
A new fern species, Athyrium aberrans (Athyriaceae), currently only known from southeastern Xizang, China is described. Morphologically, A. aberrans is superficially similar to A. falcatum in the habit and pinna shape, but the new species can easily be distinguished from the latter by having purplish petioles and rachises, a bulbil on the rachis below the lamina apex, and finely toothed pinna margins. A phylogenetic analysis based on five plastid markers demonstrates that A. aberrans is most closely related to but obviously distinct from A. epirachis.
We here describe two new species Arachniodes libingii and A. vietnamensis (Dryopteridaceae) from southern China and central Vietnam, respectively. Our previous phylogenetic analysis found a close relationship between the two new species. Further morphological studies showed that the two species share short creeping rhizomes, blackish brown and lanceolate scales, papery laminae, aristate pinnules, dense scales along fronds, and entire indusia. However, A. libingii has deltoid-oblong laminae and apices abruptly acuminate-caudate, while A. vietnamensis has shortly ovate laminae and apices gradually acuminate.
A new lycophyte species, Selaginella subchaetoloma, is described. This species is relatively abundant in Southwest China. Materials of this species were previously identified as S. chaetoloma, but a detailed morphological study shows that the new species differs by plants with creeping sterile part and ascending fertile part and dorsal leaves with abruptly short-acuminate apex. Molecular phylogenetic results indicate that S. subchaetoloma and S. chaetoloma are not closely related.
Three new species, Polystichum pseudodangii, P. xuansonense, and P. unicum, and a new record, P. omeiense, are described and reported here, respectively. These four species were found in limestone habitat in northern Vietnam, with the first species collected from Phu Tho Province and the rest from Ha Giang Province. Detailed descriptions, images, taxonomic notes, type information, examined specimens, distribution, and habitat are provided for the new species and new record.
Loxogramme yigongensis, so far known only from the type specimens collected in 1965 and treated as a synonym of L. grammitoides, is reinstated. Morphologically, these two species are closely related but L. yigongensis differs in having succulent lamina, becoming rugose when dry, lamina lanceolate with acute apex, widest near middle of lamina, and costa not or slightly raised on lower surface. Molecular analyses based on five plastid loci resolved these two species as sister to each other with distinct divergence. In addition, the ploidy level of the two species are reported for the first time.
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