The lycophyte genus Selaginella alone constitutes the family Selaginellaceae, the largest of the lycophyte families. The genus is estimated to contain 700-800 species distributed on all continents except Antarctica, with highest species diversity in tropical and subtropical regions. The monophyly of Selaginella in this broad sense has rarely been doubted, whereas its intrageneric classification has been notoriously contentious. Previous molecular studies were based on very sparse sampling of Selaginella (up to 62 species) and often used DNA sequence data from one genome. In the present study, DNA sequences of one plastid (rbcL) and one nuclear (ITS) locus from 394 accessions representing approximately 200 species of Selaginella worldwide were used to infer a phylogeny using maximum likelihood, Bayesian inference and maximum parsimony methods. The study identifies strongly supported major clades and well resolves relationships among them. Major results include: (i) six deep-level clades are discovered representing the deep splits of Selaginella; and (ii) 20 major clades representing 20 major evolutionary lineages are identified, which differ from one another in molecular, macro-morphological, ecological and spore features, and/or geographical distribution.
No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing.This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ABSTRACTUsing light microscope (LM) and scanning electron microscope (SEM), the megaspores and microspores of 77 samples representing ca. 70 species of Selaginella from China are observed. Combing previous studies, the spore morphology of nearly all documented Selaginella species from China were reviewed. Based on the morphological characteristics in megaspores and/or microspores, we divided the spores of Chinese species into 15 types and three types are further divided into various subtypes. Each type and subtype are described in detail and a key to the types and subtypes of spores is given. For the first time, the systematic significance of microspores of Selaginella are discussed, and the results indicate that microspores of Selaginella are significant in the systematics of Selaginella. Some important morphological characteristics in spores (e.g., color, micro-sculpture, size, etc.), often been neglected in previous studies, are introduced. Some spore-morphological synapomorphies of the clades and subclades, identified by recent molecular work (Zhou et al. 2015a), are well established. Using the spore morphology, the delimitation of some taxonomically difficult species in Selaginella is assessed.
Ginkgo biloba is grown worldwide as an ornamental plant for its golden leaf color. However, the regulatory mechanism of leaf coloration in G. biloba remains unclear. Here, we compared G. biloba gold-colored mutant leaves and normal green leaves in cytological, physiological and transcriptomic terms. We found that chloroplasts of the mutant were fewer and smaller, and exhibited ruptured thylakoid membranes, indistinct stromal lamellae and irregularly arranged vesicles. Physiological experiments also showed that the mutant had a lower chlorophyll, lower flavonoid and higher carotenoid contents (especially lutein). We further used transcriptomic sequencing to identify 116 differentially expressed genes (DEGs) and 46 transcription factors (TFs) involved in chloroplast development, chlorophyll metabolism, pigment biosynthesis and photosynthesis. Among these, the chlorophyll biosynthesis-related PPO showed down-regulation, while chlorophyll degradation-related NYC/NOL had up-regulated expression in mutant leaves. Z-ISO, ZDS, and LCYE, which are involved in carotenoid biosynthesis were up-regulated. Quantitative real-time PCR (RT-qPCR) further confirmed the altered expression levels of these genes at three stages. The alteration of PPO and NYC/NOL gene expression might affect chlorophyll biosynthesis and promote degradation of chlorophyll b to chlorophyll a, while the up-regulated genes Z-ISO, ZDS and LCYE enhanced carotenoid accumulation. Consequently, changes in the ratio of carotenoids to chlorophylls were the main factors driving the golden leaf coloration in the mutant G. biloba.
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.
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