浙江乌岩岭国家级自然保护区叶附生苔类及附主植物多样性

叶附生苔类植物是一类主要生长在热带雨林和亚热带常绿阔叶林维管植物叶片表面的苔类植物, 具有重要的生态价值, 是苔藓植物中最需要关注和保护的一个类群。但由于其个体细小、分类困难等原因, 此前针对我国叶附生苔类植物的采样可能存在低估其多样性的情况。为了解采样方式可能对叶附生苔类植物多样性的影响, 本研究在乌岩岭国家级自然保护区对叶附生苔开展了系统的采样及比较研究。根据叶附生苔附主植物的叶片特性、分布位置等因素, 对保护区内的叶附生苔类植物及附主植物开展沿海拔梯度的系统调查和取样, 分析了叶附生苔类植物和附主植物的多样性及物种组成情况。结果显示, 该保护区共有叶附生苔类植物5科14属49种, 叶附生苔附主植物57科84属119种。与历史数据相比, 本研究区新增叶附生苔2科5属35种, 其中浙江省新记录苔类4种。叶附生苔物种丰富度表现出随海拔升高呈单峰型分布的模式, 在500-900 m海拔段的种数最多。基于叶片数的物种累积曲线及随机抽样分析表明, 该保护区叶附生苔在不同生境及同一附主植物不同叶片间表现出较大的种类组成差异。鉴于本研究采集方法获得的较高叶附生苔物种多样性, 建议在我国叶附生苔多样性其他分布中心开展类似的研究, 以进一步摸清我国叶附生苔类植物物种多样性。     

福建苔类和角苔类植物最新名录与区系分析

福建地处亚热带,其维管植物多样性已经有过广泛的研究,但其苔类和角苔类植物的多样性却鲜为人知。在本研究中,作者基于先前的苔藓植物文献报道和近期的野外工作以及对华东师范大学标本馆部分馆藏标本的研究结果,编写了福建苔类和角苔类植物的最新名录。福建共有苔类植物41科79属351种,角苔类植物1科4属6种。本研究新增福建苔类新记录82种。福建苔类和角苔类植物区系主要由细鳞苔科（94种）,耳叶苔科（32种）,羽苔科（25种）,扁萼苔科（23种）和指叶苔科（21种）等一些热带和亚热带的大科组成。除种数较多的属以外,63个属的种数不足5种,其中包括33个单种属。福建苔类和角苔类植物的区系地理分布主要以东亚、热带亚洲和北温带成分为主。福建特有种仅Solenostoma parviperianthum一种。与台湾相比,尽管两地植物间具有较高的相似性,福建在苔类和角苔类植物的多样性以及特有成分上都还远不及台湾丰富。     

福建苔类和角苔类植物最新名录与区系分析

福建地处亚热带,其维管植物多样性已经有过广泛的研究,但其苔类和角苔类植物的多样性却鲜为人知。在本研究中,作者基于先前的苔藓植物文献报道和近期的野外工作以及对华东师范大学标本馆部分馆藏标本的研究结果,编写了福建苔类和角苔类植物的最新名录。福建共有苔类植物41科79属351种,角苔类植物1科4属6种。本研究新增福建苔类新记录82种。福建苔类和角苔类植物区系主要由细鳞苔科（94种）,耳叶苔科（32种）,羽苔科（25种）,扁萼苔科（23种）和指叶苔科（21种）等一些热带和亚热带的大科组成。除种数较多的属以外,63个属的种数不足5种,其中包括33个单种属。福建苔类和角苔类植物的区系地理分布主要以东亚、热带亚洲和北温带成分为主。福建特有种仅Solenostoma parviperianthum一种。与台湾相比,尽管两地植物间具有较高的相似性,福建在苔类和角苔类植物的多样性以及特有成分上都还远不及台湾丰富。     

秦岭地区苔类植物区系的初步研究

在野外调查和已有资料的基础上,分析了秦岭地区苔类植物区系组成和分布类型.现知秦岭地区苔类、角苔类有35科66属270种.其中指叶苔科、叶苔科、裂叶苔科、合叶苔科、齿萼苔科、羽苔科、光萼苔科、耳叶苔科、细鳞苔科、瘤冠苔科为优势科;指叶苔属、鞭苔属、叶苔属、裂叶苔属、合叶苔属、羽苔属、扁萼苔属、光萼苔属、耳叶苔属、细鳞苔属为优势属.在区系成分上,温带成分最高,东亚、热带亚洲成分次之,中国特有成分也比较高.同时有大量的科中仅一属和属中仅一种的类群存在,说明本地区苔类植物科属结构比较简单,分化程度不高.还分析了秦岭苔类植物区系及我国其他5个地区苔类植物区系之间的关系.     

祁连山地区苔类植物的初步研究

通过对祁连山所采集苔类植物标本的鉴定和统计整理,共报道祁连山苔类植物8科10属12种,均为祁连山地区的首次报道,包括11种甘肃省苔类植物新记录。其中报道的秃瓣裂叶苔（Lophozia obtusa）和卵叶羽苔（Plagiochila ovalifolia）为青海省苔类植物新记录,这也是青海省苔类植物的首次报道。     

角苔类在苔藓植物门中的系统位置

苔藓植物是植物界中由水生向陆生过渡,以孢子繁殖为主要方式的一大类群。全世界约有23000种,遍布于各大洲。关于苔类植物门的分纲,有几种不同的观点,但分歧较主要的集中在角苔类植物的系统位置上。本文综合各学者的意见提出一些肤浅的认识,供大家讨论。通常对苔藓植物门(Bry0phyta)的分纲,植物学界大都沿用了最初的观点,多强调苔藓植物营养体的形态结构,将苔藓植物门分为两纲,即苔纲(Hepaticae)和藓纲(Musci)。这里,藓纲的名称是直接采用了拉丁词中藓类的名称(Muscus,PL musci)。目前藓纲(Bryopsida)的划分界线是由德国植     

祁连山地区苔类植物的初步研究

通过对祁连山所采集苔类植物标本的鉴定和统计整理,共报道祁连山苔类植物8科10属12种,均为祁连山地区的首次报道,包括11种甘肃省苔类植物新记录。其中报道的秃瓣裂叶苔(Lophozia obtusa)和卵叶羽苔(Plagiochila ovalifolia)为青海省苔类植物新记录,这也是青海省苔类植物的首次报道。     

九龙大雾山的叶附生苔类植物

九龙大雾山已知叶附生苔类植物有5科10属12种,其中有5属6种,即双齿护蒴苔、小叶拟大萼苔、东亚指叶苔、四齿异萼苔、双齿异萼苔和台湾角鳞苔,均为我国叶附生苔类的新记录。     

钱江源国家公园叶附生苔类植物的物种多样性

本文报道钱江源国家公园叶附生苔类植物4科9属19种, 较之前记录增加3科8属17种。与钱江源处于同纬度的我国其他地区相比, 钱江源叶附生苔类植物的种数仅次于西藏墨脱, 排名第二。鳞叶疣鳞苔(Cololejeunea longifolia)和尖叶薄鳞苔(Leptolejeunea subacuta)是钱江源国家公园最常见的2种叶附生苔类植物。丰富的叶附生苔类植物表明钱江源国家公园具有较合适的叶附生苔类植物生长环境, 这可能与该地区长期有效的保护密切相关。     

江西齐云山自然保护区苔藓植物研究

通过野外考察和标本鉴定,江西齐云山自然保护区共有苔藓植物52科98属191种(包括2亚种和3变种),其中苔类植物22科27属46种1亚种,藓类植物30科71属140种1亚种3变种.报道江西新记录科3科:裸蒴苔科(Haplomitriaceae)、南溪苔科(Makinoaceae)和带叶苔科(Pallaviciniaceae);江西新记录属5属:裸蒴苔属(Haplomitrium)、南溪苔属(Makinoa)、带叶苔属(Pallavicinia)、黄角苔属(Phaeoceros)和细指苔属(Kurzia),以及24种苔类植物为江西新记录.     

Major transitions in the evolution of early land plants: a bryological perspective

Background Molecular phylogeny has resolved the liverworts as the earliest-divergent clade of land plants and mosses as the sister group to hornworts plus tracheophytes, with alternative topologies resolving the hornworts as sister to mosses plus tracheophytes less well supported. The tracheophytes plus fossil plants putatively lacking lignified vascular tissue form the polysporangiophyte clade. Scope This paper reviews phylogenetic, developmental, anatomical, genetic and paleontological data with the aim of reconstructing the succession of events that shaped major land plant lineages. Conclusions Fundamental land plant characters primarily evolved in the bryophyte grade, and hence the key to a better understanding of the early evolution of land plants is in bryophytes. The last common ancestor of land plants was probably a leafless axial gametophyte bearing simple unisporangiate sporophytes. Water-conducting tissue, if present, was restricted to the gametophyte and presumably consisted of perforate cells similar to those in the early-divergent bryophytes Haplomitrium and Takakia. Stomata were a sporophyte innovation with the possible ancestral functions of producing a transpiration-driven flow of water and solutes from the parental gametophyte and facilitating spore separation before release. Stomata in mosses, hornworts and polysporangiophytes are viewed as homologous, and hence these three lineages are collectively referred to as the 'stomatophytes'. An indeterminate sporophyte body (the sporophyte shoot) developing from an apical meristem was the key innovation in polysporangiophytes. Poikilohydry is the ancestral condition in land plants; homoiohydry evolved in the sporophyte of polysporangiophytes. Fungal symbiotic associations ancestral to modern arbuscular mycorrhizas evolved in the gametophytic generation before the separation of major present-living lineages. Hydroids are imperforate water-conducting cells specific to advanced mosses. Xylem vascular cells in polysporangiophytes arose either from perforate cells or de novo. Food-conducting cells were a very early innovation in land plant evolution. The inferences presented here await testing by molecular genetics.     

Mosses share mitochondrial group II introns with flowering plants, not with liverworts

Extant bryophytes are regarded as the closest living relatives of the first land plants, but relationships among the bryophyte classes (mosses, liverworts and hornworts) and between them and other embryophytes have remained unclear. We have recently found that plant mitochondrial genes with positionally stable introns are well suited for addressing questions of plant phylogeny at a deep level. To explore further data sets we have chosen to investigate the mitochondrial genes nad4 and nad7, which are particularly rich in intron sequences. Surprisingly, we find that in these genes mosses share three group II introns with flowering plants, but none with the liverwort Marchantia polymorpha or other liverworts investigated here. In mitochondria of Marchantia, nad7 is a pseudogene containing stop codons, but nad7 appears as a functional mitochondrial gene in mosses, including the isolated genus Takakia. We observe the necessity for strikingly frequent C-to-U RNA editing to reconstitute conserved codons in Takakia when compared to other mosses. The findings underline the great evolutionary distances among the bryophytes as the presumptive oldest division of land plants. A scenario involving differential intron gains from fungal sources in what are perhaps the two earliest diverging land plant lineages, liverworts and other embryophytes, is discussed. With their positionally stable introns, nad4 and nad7 represent novel marker genes that may permit a detailed phylogenetic resolution of early clades of land plants.     

TRANSITION TO A LAND FLORA: PHYLOGENETIC RELATIONSHIPS OF THE GREEN ALGAE AND BRYOPHYTES

Abstract— Separate cladistic analyses of the green algae, liverworts, and hornworts are presented. Classificatory and evolutionary implications of these analyses, in addition to our previously published cladistic analyses of mosses and the embryophytes as a whole, are discussed. The embryophytes are monophyletic, and are part of a larger monophyletic group that includes some of the green algae (the "charophytes"). Important evolutionary transformations in the early phylogeny of the land plants include: (1) retention of the zygote on the haploid plant (gametophyte), with the sporophyte generation arising de novo by delaying meiosis, (2) independent elaboration of an elongate sporophyte in some liverworts, some hornworts, and in the moss-tracheophyte clade, (3) independent origin of radial (axial) symmetry in the gametophyte in some liverworts and in the moss-tracheophyte clade, (4) independent origin of leaves on the gametophyte in some liverworts and in mosses, and (5) the unique development of a branching sporophyte with multiple sporangia in the tracheophytes.     

Distribution of cell-wall xylans in bryophytes and tracheophytes: new insights into basal interrelationships of land plants

Xylans are known to be major cellulose-linking polysaccharides in secondary cell walls in higher plants. We used two monoclonal antibodies (LM10 and LM11) for a comparative immunocytochemical analysis of tissue and cell distribution of xylans in a number of taxa representative of all major tracheophyte and bryophyte lineages. The results show that xylans containing the epitopes recognized by LM10 and LM11 are ubiquitous components of secondary cell walls in vascular and mechanical tissues in all present-living tracheophytes. In contrast, among the three bryophyte lineages, LM11 binding was detected in specific cell-wall layers in pseudoelaters and spores in the sporophyte of hornworts, while no binding was observed with either antibody in the gametophyte or sporophyte of liverworts and mosses. The ubiquitous occurrence of xylans containing LM10 and LM11 epitopes in tracheophytes suggests that the appearance of these polysaccharides has been a pivotal event for the evolution of highly efficient vascular and mechanical tissues. LM11 binding in the sporophyte of hornworts, indicating the presence of relatively highly substituted xylans (possibly arabinoxylans), separates these from the other bryophytes and is consistent with recent molecular data indicating a sister relationship of the hornworts with tracheophytes.     

Anatomy and ultrastructure of the sporophyte of Takakia ceratophylla (Bryophyta)

In this study, morphogenesis and structure of the sporophyte of Takakia ceratophylla are characterized beginning with the late embryo and culminating in the fully dehisced capsule. Information is presented on the development, ultrastructure, and anatomy of the three organographic regions of the sporophyte, namely capsule, seta, and foot. Diagnostic features that identify Takakia as a moss include the gradual elongation of seta, persistence of an apical calyptra, expansion of the capsule after cessation of seta elongation, existence of a columella, monoplastidic meiosis, spore ultrastructure (including a perine layer deposited late in spore wall (development), and the structure of the foot. Commonalities with the capsule of the Andreaeopsida include sporogenous tissue that overarchs a central columella, absence of stomata, and lack of a peristome and operculum. Peculiarities of the genus are seen in the internal structure of the capsule, the disintegration of the columella with spore maturation, and the dehiscence of the capsule along a single, spiralled, longitudinal suture line. Passive spore dispersal through longitudinal splitting of the capsule occurs in andreaeopsid mosses, liverworts, hornworts, and seedless vascular plants. The precise mechanism of dehiscence along a spiralled suture is unparalleled in extant archegoniates but finds counterparts in ancestral land plants such as the pteridophyte Tortilicaulis.     

Vegetative and reproductive innovations of early land plants: implications for a unified phylogeny

As the oldest extant lineages of land plants, bryophytes provide a living laboratory in which to evaluate morphological adaptations associated with early land existence. In this paper we examine reproductive and structural innovations in the gametophyte and sporophyte generations of hornworts, liverworts, mosses and basal pteridophytes. Reproductive features relating to spermatogenesis and the architecture of motile male gametes are overviewed and evaluated from an evolutionary perspective. Phylogenetic analyses of a data set derived from spermatogenesis and one derived from comprehensive morphogenetic data are compared with a molecular analysis of nuclear and mitochondrial small subunit rDNA sequences. Although relatively small because of a reliance on water for sexual reproduction, gametophytes of bryophytes are the most elaborate of those produced by any land plant. Phenotypic variability in gametophytic habit ranges from leafy to thalloid forms with the greatest diversity exhibited by hepatics. Appendages, including leaves, slime papillae and hairs, predominate in liverworts and mosses, while hornwort gametophytes are strictly thalloid with no organized external structures. Internalization of reproductive and vegetative structures within mucilage-filled spaces is an adaptive strategy exhibited by hornworts. The formative stages of gametangial development are similar in the three bryophyte groups, with the exception that in mosses apical growth is intercalated into early organogenesis, a feature echoed in moss sporophyte ontogeny. A monosporangiate, unbranched sporophyte typifies bryophytes, but developmental and structural innovations suggest the three bryophyte groups diverged prior to elaboration of this generation. Sporophyte morphogenesis in hornworts involves non-synchronized sporogenesis and the continued elongation of the single sporangium, features unique among archegoniates. In hepatics, elongation of the sporophyte seta and archegoniophore is rapid and requires instantaneous wall expandability and hydrostatic support. Unicellular, spiralled elaters and capsule dehiscence through the formation of four regular valves are autapomorphies of liverworts. Sporophytic sophistications in the moss clade include conducting tissue, stomata, an assimilative layer and an elaborate peristome for extended spore dispersal. Characters such as stomata and conducting cells that are shared among sporophvtes of mosses, hornworts and pteridophytes are interpreted as parallelisms and not homologies. Our phylogenetic analysis of three different data sets is the most comprehensive to date and points to a single phylogenetic solution for the evolution of basal embryophytes. Hornworts are supported as the earliest divergent embryophyte clade with a moss/liverwort clade sister to tracheophytes. Among pteridophytes, lycophytes are monophyletic and an assemblage containing ferns, Equisetum and psilophytes is sister to seed plants. Congruence between morphological and molecular hypotheses indicates that these data sets are tracking the same phylogenetic signal and reinforces our phylogenetic conclusions. It appears that total evidence approaches are valuable in resolving ancient radiations such as those characterizing the evolution of early embryophytes. More information on land plant phylogeny can be found at: http: //www.science.siu.edu/ landplants/index.html.     

Cell and molecular biology of bryophytes: ultimate limits to the resolution of phylogenetic problems

Ultrastructure, biochemistry and 5S rRNA sequences link tracheophytes, bryophytes and charalean green algae, but the precise interrelationships between these groups remain unclear. Further major clarification now awaits primary sequence data. These are also needed to determine directionality in possible evolutionary trends within the bryophytes, but are unlikely to overturn current schemes of classification or phylogeny. Comparative ultrastructural studies of spermatogenesis, sporogenesis, the cytoskeleton and plastids reinforce biochemical and morphogenetic evidence for the wide phyletic discontinuities between mosses, hepatics and hornworts, and also rule out direct lines of descent between them. Direct ancestral lineages from charalean algae to bryophytes and to tracheophytes are also unlikely. EM studies of gametophyte/sporophyte junctions, plus immunological investigations of bryophyte cytoskeletons, are likely to accentuate the differences between mosses, hepatirs and hornworts. Other priorities for systematics include elucidation of oil body ultrastructure, analysis of the changes in nuclear proteins during spermatogenesis and a detailed comparison of bryophyte and charalean plastids. The combined evidence from ultrastrueture, biochemistry, morphology and morphogenesis warrants general acceptance of the polyphyletic origin of the bryophytes. Ultrastructural attributes should be more widely used in bryophyte systematics.     

Phylogeny and diversification of bryophytes

The bryophytes comprise three phyla of embryophytes that are well established to occupy the first nodes among extant lineages in the land-plant tree of life. The three bryophyte groups (hornworts, liverworts, mosses) may not form a monophyletic clade, but they share life history features including dominant free-living gametophytes and matrotrophic monosporangiate sporophytes. Because of their unique vegetative and reproductive innovations and their critical position in embryophyte phylogeny, studies of bryophytes are crucial to understanding the evolution of land plant morphology and genomes. This review focuses on phylogenetic relationships within each of the three divisions of bryophytes and relates morphological diversity to new insights about those relationships. Most previous work has been on the mosses, but progress on understanding the phylogeny of hornworts and liverworts is advancing at a rapid pace. Multilocus multigenome studies have been successful at resolving deep relationships within the mosses and liverworts, whereas single-gene analyses have advanced understanding of hornwort evolution.     

The origin of the sporophyte shoot in land plants: a bryological perspective

Background

Land plants (embryophytes) are monophyletic and encompass four major clades: liverworts, mosses, hornworts and polysporangiophytes. The liverworts are resolved as the earliest divergent lineage and the mosses as sister to a crown clade formed by the hornworts and polysporangiophytes (lycophytes, monilophytes and seed plants). Alternative topologies resolving the hornworts as sister to mosses plus polysporangiophytes are less well supported. Sporophyte development in liverworts depends only on embryonic formative cell divisions. A transient basal meristem contributes part of the sporophyte in mosses. The sporophyte body in hornworts and polysporangiophytes develops predominantly by post-embryonic meristematic activity.

Scope

This paper explores the origin of the sporophyte shoot in terms of changes in embryo organization. Pressure towards amplification of the sporangium-associated photosynthetic apparatus was a major driver of sporophyte evolution. Starting from a putative ancestral condition in which a transient basal meristem produced a sporangium-supporting seta, we postulate that in the hornwort–polysporangiophyte lineage the basal meristem acquired indeterminate meristematic activity and ectopically expressed the sporangium morphogenetic programme. The resulting sporophyte body plan remained substantially unaltered in hornworts, whereas in polysporangiophytes the persistent meristem shifted from a mid-embryo to a superficial position and was converted into an ancestral shoot apical meristem with the evolution of sequential vegetative and reproductive growth.

Conclusions

The sporophyte shoot is interpreted as a sterilized sporangial axis interpolated between the embryo and the fertile sporangium. With reference to the putatively ancestral condition found in mosses, the sporophyte body plans in hornworts and polysporangiophytes are viewed as the product of opposite heterochronic events, i.e. an anticipation and a delay, respectively, in the development of the sporangium. In either case the result was a pedomorphic sporophyte permanently retaining juvenile characters.