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Morphology of rhizome scales and soral paraphyses of Lepisorus. (a-l) Scales. (a) Lepisorus obscure-venulosus. (b) Lepisorus oligolepidus. (c) Lepisorus hachijoensis. (d) Lepisorus bicolor. (e) Lepisorus macrosphaerus. (f) Lepisorus (Belvisia) henryi. (g) Lepisorus scolopendrium. (h) Lepisorus ussuriensis. (i) Lepisorus loriformis. (j) Lepisorus (Platygyria) waltonii. (k) Lepisorus longifolius (Paragramma longifolia). (l) Lepisorus jakonensis. (m-u) Paraphyses. (m) Lepisorus loriformis. (n) Lepisorus pseudonudus. (o) Lepisorus (Belvisia) spicatus. (p) Lepisorus hachijoensis. (q) Lepisorus oligolepidus. (r) Lepisorus macrosphaerus. (s) Lepisorus bicolor. (t) Lepisorus jakonensis. (u) Lepisorus (Platygyria) waltonii.
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The fern genus Lepisorus represents one of the most complicated and controversial lineages in Polypodiaceae, with about 80 species which have been classified into several separate genera, and is notorious for its taxonomic difficulty. Despite progress in recent phylogenetic studies of the family Polypodiaceae involving Lepisorus and its allies, the...
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Context 1
... typical Lepisorus species which are mainly in subtropical or even temperate Asia. The distinct morphology of coenosori restricted to the strongly narrowed tip of the leaves in Belvisia leads to the exclusion of this genus from Lepisorus ( Fig. 1), whereas their similarities, such as rhizome scales, sori and paraphyses have been largely overlooked (Fig. 2). Another example is the appearance of hairy rhizome scales, which is the diagnostic character of Tricholepidium, but is shared also by some members of Neolepisorus and Lepidomicrosorium. Compared with solved lineages, the relationship between Lepisorus and Paragramma is still problematic. Based on its distinct hypoderms, paraphyses and ...
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... in the Supporting Information section at the end of the article. Fig. S1. The inferred ancestral states for characters (a) rhizome diameter, and (b) lumina of rhizome scales, based on the Mk1 model. Black and white in pie charts on each branch indicate the probabilities of ancestral states inferred by the analysis based on the 100 MLBS trees. Fig. S2. The inferred ancestral states for characters (a) shape of rhizome scales and (b) colour states of scales, based on the Mk1 model. Black and white in the pie charts on each branch indicate the probabilities of ancestral states inferred by the analysis based on the 100 MLBS trees. Fig. S3. The inferred ancestral states for characters ...
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Scales are epidermal appendages on the rhizomes and leaves of many ferns. Features of scales play an important role in classification of ferns. The phylogenetic position and delimitation of Pteridaceae were treated differently by different authors. Here we collected scales of 76 fern species in Pteridaceae, and observed them under a dissecting micr...
Citations
... To keep Lepisorus monophyletic, two opposite taxonomic treatments have been proposed (Wei et al., 2021). i.e., broadening the circumscription of Lepisorus including Neolepisorus, Lemmaphyllum, Lepidomicrosorium etc. (Zhao et al., 2020b) vs. establishing a new genus Ellipinema for L. jakonensis . We decided to adopt the former idea, judging that there are many disadvantages to admitting allopolyploids across two genera. ...
... Thus, all the species of Neolepisorus, Lemmaphyllum and Lepidomicrosorium are transferred to Lepisorus. Wei and Zhang (2013) treated Lemmaphyllum microphyllum as a variant of L. carnosum (J.Sm) C.Presl based on morphometric data, and the recent transfer into Lepisorus (Zhao et al., 2020b) is based on this idea. However, we transferred Lemmaphyllum microphyllum into Lepisorus, maintaining it at the rank of species. ...
... In our study, we observed epidermal cells with straight walls in Pyrrosia lanceolata (Wei and Zhao 2019). Combined molecular studies showed that Lepisorus were recovered to be monophyletic when Tricholepidium, Lemmaphyllum, Neolepisorus and some other related genera were included (Zhao et al. 2020). The stomatal type of Lepisorus normalis varies from copolocytic to polocytic as in all other Lepisorus species in our study. ...
... Raised several subfamilies like Gymnogrammitis (Gymnogrammitidaceae), Drynaria (Drynariaceae), and Platycerium (Platyceriaceae) to the family level. Later, phylogenetic studies based on multiple parameters provided better comprehension of the familial and generic circumscription of Polypodiaceae Schuettpelz and Pryer, 2007;Kreier et al. 2008;Testo et al. 2019;Zhao et al. 2020). ...
... Therefore, it is evident from the study that leaf micro-morphological details and venation patterns can serve as an additional set of data in line with molecular and morphological characters which could help in decoding the existing problems up to generic and specific levels. The combination of diagnostic morphological characters like rhizome scales, leaf shapes, venation patterns, and features of the paraphyses proved crucial in untangling the clades and sub-clades of genus Lepisorus, Pleopeltis, and the grammatid ferns Otto et al. 2009;Wang et al. 2010;Zhao et al. 2020). CONCLUSION The present investigation reveals that leaf architectural traits and venation patterns especially higher degree veins are useful characters in delineating species. ...
The present study insights into the interrelationships and taxonomic segregation of some Polypodiaceous fern taxa based on leaf architecture and foliar micro-morphology. Twenty-seven fern species were examined and valuable qualitative and quantitative data were obtained to generate UPGMA dendrogram. A dichotomous key differentiating the taxa was prepared. Results reveal that even though species have overlapping characters, certain specific traits prove taxonomically valuable. The results proved that traits like leaf shape, higher order leaf venation, stomatal and epidermal features are indeed important diagnostic characters and hence can be used for the identification of fern species in their immature stage or even in absence of sori. These data sets often combined with other morphological as well molecular data would contribute to fern phylogenetic study particularly of the large and complex family Polypodiaceae.
... Sm.) Ching (Polypodiaceae) occurs throughout Eastern and Southern Asia with range extensions towards the Pacific islands including Hawai'i and towards tropical Afromadagascar (Ching 1933;Zink 1993;Wang et al. 2012). Taking into account the various taxonomic studies and the recent proposal to expand the generic circumscription by including all genera of the tribe Lepisoreae such as Lemmaphyllum C. Presl, Lepidomicrosorium Ching & K.H. Shing, Neocheiropteris Christ, Neolepisorus Ching, Paragramma (Blume) T. Moore, and Tricholepidium Ching (Zhao et al. 2020) Lepisorus can be currently recognized as one of the most species-rich genera among genera in Polypodiaceae, comprising ca. 90 species in 18 sections (PPGI 2016;Fujiwara et al. 2020;Zhao et al. 2020). ...
... Taking into account the various taxonomic studies and the recent proposal to expand the generic circumscription by including all genera of the tribe Lepisoreae such as Lemmaphyllum C. Presl, Lepidomicrosorium Ching & K.H. Shing, Neocheiropteris Christ, Neolepisorus Ching, Paragramma (Blume) T. Moore, and Tricholepidium Ching (Zhao et al. 2020) Lepisorus can be currently recognized as one of the most species-rich genera among genera in Polypodiaceae, comprising ca. 90 species in 18 sections (PPGI 2016;Fujiwara et al. 2020;Zhao et al. 2020). Whereas the core of the genus Lepisorus (Lepisorus s.s.) is easily recognized by its unique suite of morphological characters including creeping rhizomes covered by clathrate scales, simple leaves, and sori covered with scale-like paraphyses (Ching 1933;Qi et al. 2013;Zhao et al. 2020), some controversy still exists concerning the broader circumscription to avoid the need to recognize the genus Ellipinema Li Bing Zhang & Liang Zhang . ...
... 90 species in 18 sections (PPGI 2016;Fujiwara et al. 2020;Zhao et al. 2020). Whereas the core of the genus Lepisorus (Lepisorus s.s.) is easily recognized by its unique suite of morphological characters including creeping rhizomes covered by clathrate scales, simple leaves, and sori covered with scale-like paraphyses (Ching 1933;Qi et al. 2013;Zhao et al. 2020), some controversy still exists concerning the broader circumscription to avoid the need to recognize the genus Ellipinema Li Bing Zhang & Liang Zhang . Despite significant progress (Wang et al. 2010a;Wei et al. 2017;Zhang et al. 2020;Zhao et al. 2020), taxonomic uncertainty is arguably not restricted to the generic classification but affects the estimation of the total species diversity that is expected to be still underestimated due to the difficulty in the taxonomic classification of this genus. ...
A new species of the species-rich fern genus Lepisorus (Polypodiales, Polypodiaceae) has been found to occur in Shan state, Myanmar. Lepisorus medioximus is described based on morphological characters and phylogenetic evidence. Phylogenetic analyses showed that the specimens of L. medioximus formed a distinct clade nested in the Pseudovittaria clade. The morphological comparison demonstrated that the species is distinct from phylogenetically related species, namely L. elegans , L. contortus , and L. tosaensis , in the morphology of the rhizome scales, size, and shape of the lamina, position of sori, and paraphyses.
... and Zealandia Testo from the paraphyletic genus Microsorum. Link, and Zhao et al. (2020) expanded the definition of the genus Lepisorus (J.Sm.) Ching. ...
... gov/genbank/), covering all genera accepted in PPG I (2016) as far as data availability enabled. To reflect recent progress in our understanding of the natural classification of Polypodiaceae, the treatment of some taxa deviated from PPGI by adapting new concepts (Testo et al. 2019;Zhao et al. 2020;Chen et al. 2021). In the Genbank database, some accessions contained indels, which should not be present in rbcL since it is a protein-coding gene; we removed such low-quality accessions. ...
The genus Thylacopteris is a small, phylogenetically isolated genus belonging to the fern family Polypodiaceae. This study describes a new species, Thylacopteris minuta , based on collections obtained during field surveys of Shan State, Myanmar. This new species is distinct from other species of Thylacopteris in its small size and presence of sclerenchyma strands in the rhizome. This species is also distinct from the only other species of Thylacopteris with molecular data available, T. papillosa , in a plastid rbcL phylogeny of Polypodiaceae. This new discovery of Thylacopteris from Myanmar suggests that this genus is still overlooked in Southeast Asia.
... Its close ally Pleopeltis has certain similarities with genera Microsorum and Phymatodes (PPG, 2016). The generic circumscription of Lepisorus and its allies is a matter of controversy since its recognition (Zhao et al., 2020). In reference to the new classification of Lepisorus by Zhao et al. (2020) are now considered as their synonyms, respectively (Wei & Zhao, 2019). ...
... The generic circumscription of Lepisorus and its allies is a matter of controversy since its recognition (Zhao et al., 2020). In reference to the new classification of Lepisorus by Zhao et al. (2020) are now considered as their synonyms, respectively (Wei & Zhao, 2019). In recent works, SEM and LM observations were useful for evaluation of micro-morphological features in certain species (Adeonipekun et al., 2021;Hameed et al., 2022;Rashid et al., 2018;Usma et al., 2020). ...
The genus Lepisorus is often associated with complex lineages in Polypodiaceae, which leads to difficulty in understanding taxonomic relationships among the species. The taxa is mostly epiphytic with some distinct features like rhizomes covered with clathrate scales, simple fronds, sori in a single row, intermixed with peltate paraphyses. In this study, we precisely focus on the Lepisorus species and seek a broader understanding of the taxonomic relationships that prevail in this genus. The morpho-anatomical traits of rhizomes, stipe, lamina, and sporangia were explored in nine species with the aid of light microscopy and scanning electron microscopy techniques for gathering the qualitative and quantitative data sets. These character traits were resolved into groups based on degree of similarity and principal component analysis to comprehend the covariance among the variables. The data were then employed to prepare an artificial dichotomous key and hierarchical cluster analysis was established that revealed five related clades with eight members whereas one member emerged as outgroup. The results correspond with other recent phylogenetic studies of the concerned genus hence confirming immense reliability and thrust of light microscopy, scanning electron microscopy, and morphology-based studies that are being less utilized in fern taxonomy.
... Lemmaphyllum (C. Presl) C.F. Zhao, R. Wei & X.C. Zhang in the most recently phylogeny study of Lepisorus (Zhao et al., 2020). Species of Lemmaphyllum were formed a monophyletic Lemmaphyllum clade, thus, to avoid the use of unpublished new combinations names of each species and text redundancy, we tend to use Lemmaphyllum in this study. ...
... nud.) (the replacement of Pleopeltidoideae) and Pyrrosioideae, and raised Gymnogrammitis (Gymnogrammitidaceae), Drynaria (Drynariaceae), and Platycerium (Platyceriaceae) to the family level. Recent phylogenetic studies based on multi-locus data not only provided better understanding of the familial and generic circumscription of Polypodiaceae (e.g., Schneider & al., 2004b;Schuettpelz & Pryer, 2007;Kreier & al., 2008a;Otto & al., 2009;Wang & al., 2010a;Sundue & al., 2014;He & al., 2018;Testo & al., 2019;Chen & al., 2020;Zhao & al., 2020a), but also shed new light on the deep phylogenetic relationships within the family that further promoted the comprehensive classification of ferns (e.g., Smith & al., 2006a;Christenhusz & al., 2011;PPG I, 2016). All these studies agreed with a broad definition of Polypodiaceae by lumping several closely related groups into it, including loxogrammoids, grammitids, Gymnogrammitidaceae, Drynariaceae, and Platyceriaceae, the infrafamilial classification, however, was not untangled, especially the problem of polyphyly of Polypodioideae to Grammitidoideae (grammitids), the monophyly of Polypodium s.str. ...
... Most earlier studies used previously recognized subfamilies in Polypodiaceae for their discussions, but few of them focused on the natural circumscription of these subfamilial lineages (Smith & al., 2006a;Christenhusz & al., 2011;PPG I, 2016). Although we did not include a comprehensive species sampling, we incorporated representatives from nearly all the major clades identified previously (e.g., Schneider & al., 2004bSchuettpelz & Pryer, 2007;Kreier & al., 2008a;Otto & al., 2009;Wang & al., 2010a,b;Sundue & al., 2014;Almeida & al., 2017;Zhou & al., 2017;He & al., 2018;Chen & al., 2020;Zhao & al., 2020a;Wei & al., 2021a), and fully considered the morphological variation and geographic range of this family. Moreover, by incorporating plastome and nuclear ribosomal genome data, we resolved the phylogenetic placement of Synammia and Adetogramma, monophyly of Polypodium, and the infra-subfamilial relationship within subfamily Polypodioideae. ...
... Although frequent parallel evolution and homoplasy in morphology have been reported in Polypodiaceae (e.g., Hovenkamp, 1996;He & al., 2018), one solution for a stable infrafamilial classification might be identifying diagnostic combinations of characters (Table 3) that are predicted to be of systematic value including stele arrangement in rhizomes and petioles, venation pattern, and spore ornamentation (De la Sota, 1973;Van Uffelen & Hennipman, 1985;Van Uffelen, 1992, 1993, 1997Tejero-Díez & al., 2010;, instead of using a single synapomorphy. For instance, the combination of rhizome scales, leaf shapes, venation patterns and features of paraphyses is effective for distinguishing clades/subclades within Lepisorus, Pleopeltis, and grammitid ferns (Ranker & al., 2004;Otto & al., 2009;Wang & al., 2010a,b;Zhao & al., 2020a), and such an approach is also applicable at higher levels of classification. Therefore, considering affinities in morphology, evolutionary pattern and biogeographic history, and for the sake of a natural classification, we accept the six subfamilies previously recognized in Polypodiaceae (PPG I, 2016) by uniting Synammia with Crypsinoideae, keeping Dictymia in Loxogrammoideae, and maintaining Thylacopteris in Microsoroideae. ...
The polygrammoid fern family Polypodiaceae represents one of the most diversified epiphytic fern groups, with more than 1600 species distributed on all continents except Antarctica, with the highest species diversity in tropical and subtropical regions. Despite progress in recent phylogenetic studies of Polypodiaceae, the infrafamilial classification of this group of ferns is still problematic. Here, we explore the phylogenetic relationship within Polypodiaceae using plastid genome (plastome) and nuclear ribosomal cistron genome data obtained from high‐throughput sequencing. Although genome skimming data strongly support the monophyly of many genera and clades of Polypodiaceae, relationships within some clades and along the backbone of the phylogeny remain incongruent between plastome and nuclear data. The explanation is possibly a factor of complex evolutionary history found in these clades, such as rapid radiation, incomplete lineage sorting, ancient hybridization, and recent introgression. Based on the concatenated dataset, our phylogenetic analyses support nine major clades in Polypodiaceae, which merit the recognition as subfamilies, Crypsinoideae, Grammitidoideae, Loxogrammoideae, Microsoroideae, Platycerioideae, Polypodioideae, Adetogrammoideae, Campyloneuroideae, and Serpocauloideae, while the latter three are separated from Polypodioideae as new subfamilies. All of these infrafamilial divisions, identified with molecular data, are further supported by non‐molecular features including leaf dissection, venation, scales and paraphyses, soral features, and geographical distributions. Systematic and taxonomic discussions on the subfamilial treatment are also provided.
... The Polypodiaceae is one of the largest families of ferns (PPG I 2016) and is renowned for morphological homoplasy (Hovenkamp 1996, Ranker et al. 2004, He et al. 2018, Zhao et al. 2019. This is reflected in frequent changes to the circumscriptions of its constituent genera. ...
The fern genus Microsorum is not monophyletic, with previous phylogenetic analyses finding three lineages to group not with the type species, but to form a grade related to the 13 species of Lecanopteris . These three lineages have recently been recognised as separate genera: Bosmania , Dendroconche , and Zealandia . Here, we explore the morphological characterisation of Lecanopteris and these other three lecanopteroid genera. While the traditional circumscription of Lecanopteris has seemed sacrosanct, its defining morphological character states of rhizome cavities and ant brooding associations occur in other lecanopteroid ferns and elsewhere in the Polypodiaceae . Instead, we suggest that the morphological characterisation of an expanded Lecanopteris including the Dendroconche and Zealandia lineages is just as good, if not better, with the pertinent character states being the absence of sclerenchyma strands in the rhizome and at least some fronds having Nooteboom’s type 5 venation pattern. This wider circumscription is also better able to accommodate phylogenetic uncertainty, and it means that groups of species traditionally placed together in a single genus are not distributed across different genera. General users familiar with the narrower circumscription of Lecanopteris will not be significantly disrupted, because there is little geographic overlap with the lineages added to the genus. Consequently, we make new combinations in Lecanopteris for 11 species and one subspecies.
... The taxonomic decisions that were taken in this study were made once mitochondrial clades were recovered, morphological variation was characterized and analysed, and ecology information was typified. We consider that phylogenetic clades represent independent evolutionary units when, by coalescence processes, they end up depicting concordant clades for mtDNA, morphology and ecology (Kosakyan et al., 2016b;Inoue et al., 2020;Lara et al., 2020;Zhao et al., 2020). ...
Protists are probably the most species-rich eukaryotes, yet their systematics are inaccurate, leading to an underestimation of their actual diversity. Arcellinida (= lobose testate amoebae) are amoebozoans that build a test (a hard shell) whose shape and composition are taxonomically informative. One of the most successful groups is Arcellidae, a family found worldwide in many freshwater and terrestrial environments where they are indicators of environmental quality. However, the systematics of the family is based on works published nearly a century ago. We re-evaluated the systematics based on single-cell barcoding, morphological and ecological data. Overall, test shape appears to be more related to environmental characteristics than to the species’ phylogenetic position. We show several convergences in organisms with similar ecology, some traditionally described species being paraphyletic. Based on conservative traits, we review the synapomorphies of the infraorder Sphaerothecina, compile a list of synonyms and describe a new genus Galeripora, with five new combinations. Seven new species: Arcella guadarramensis sp. nov., Galeripora balari sp. nov., Galeripora bufonipellita sp. nov., Galeripora galeriformis sp. nov., Galeripora naiadis sp. nov., Galeripora sitiens sp. nov. andGaleripora succelli sp. nov. are also described here.
... From the evolutionary emergence of primitive organisms to today's broad variety of organisms, people have been constantly exploring how many species there are on the earth and what kind of evolutionary relationship among species. With the development of open science and technological innovation, methods of species identification range from using morphological characteristics to the integration of various methods (e.g., molecular biology, bioinformatics, bionomics) [1][2][3][4][5], which help us gain a more in-depth understanding of the evolutionary process between organisms and their accurate position in the tree of life. Due to the multi-disciplines combination and the improvement of sharing databases, many misclassifications hidden in the past have been gradually discovered, and their key morphological boundaries have also been rewritten. ...
With the development of open science and technological innovation, using sharing data and molecular biology techniques in the study of taxonomy and systematics have become a crucial component of plants, which undoubtedly helps us discover more hidden outliers or deal with difficult taxa. In this paper, we take Dennstaedtia smithii as an example, based on sharing molecular database, virtual herbarium and plant photo bank, to clarify the outliers that have been hidden in Dennstaedtia and find the key morphological traits with consistent of molecular systematics. In molecular phylogenetic analyses, we used rbcL, rps4, psbA-trnH and trnL-F sequences from 5 new and 49 shared data; the results showed that Dennstaedtia smithii is nested within Microlepia rather than Dennstaedtia. We further studied the morphological characters based on the phylogeny result and found that D. smithii is distinguished from other species of Dennstaedtia by spore ornamentation and the unconnected of grooves between rachis and pinna rachis. According to morphological and molecular phylogenetic studies, our results supported that D. smithii should be a new member of Microlepia and renamed Microlepia smithii (Hook.) Y.H. Yan. Finding hidden outliers can promote the consistency of morphological and molecular phylogenetic results, and making the systematic classification more natural.
