A molecular phylogeny of scaly tree ferns (Cyatheaceae)

Tree ferns recently were identified as the closest sister group to the hyperdiverse clade of ferns, the polypods. Although most of the 600 species of tree ferns are arborescent, the group encompasses a wide range of morphological variability, from diminutive members to the giant scaly tree ferns, Cyatheaceae. This well-known family comprises most of the tree fern diversity (～500 species) and is widespread in tropical, subtropical, and south temperate regions of the world. Here we investigate the phylogenetic relationships of scaly tree ferns based on DNA sequence data from five plastid regions (rbcL, rbcL-accD IGS, rbcL-atpB IGS, trnG-trnR, and trnL-trnF). A basal dichotomy resolves Sphaeropteris as sister to all other taxa and scale features support these two clades: Sphaeropteris has conform scales, whereas all other taxa have marginate scales. The marginate-scaled clade consists of a basal trichotomy, with the three groups here termed (1) Cyathea (including Cnemidaria, Hymenophyllopsis, Trichipteris), (2) Alsophila sensu stricto, and (3) Gymnosphaera (previously recognized as a section within Alsophila) + A. capensis. Scaly tree ferns display a wide range of indusial structures, and although indusium shape is homoplastic it does contain useful phylogenetic information that supports some of the larger clades recognised.     

中国桫椤科植物叶绿体trnL内含子和trnL-trnF基因间隔区序列的系统发育分析

运用对PCR产物直接测序和克隆后测序的方法测定了蚌壳蕨科1种和桫椤科11种(其中桫椤分别测定19株：小羽桫椤测定2株)植物的叶绿体trnL基因内含子和trnL-trnF基因间隔区序列。12种植物相应的长度介亍l004-l082之间,A T平均含量60．9％,G C平均含量39．1％。计算了不同种间以及种内不同个体间序列的碱基差别(转换值／颠换值)和Kimura遗传距离。序列数据经排列后分别进行最简约法、最大似然法和邻接法分析,结果显示：(1)白桫椤、海南白桫椤和大羽桫椤构成的分支最早和该科内其余植物组成的另一分支分歧,而后者又进一步分为刃个亚分支,分别和桫椤亚属、黑桫椤亚属对应,支持夏群的分类处理：(2)大桫椤～狭羽桫椤～毛轴桫椤～篦齿桫椤、多羽桫椤一白桫椤～海南白桫椤以及小羽桫椤一桫椤各自构成独立、自然的末端分支,再参照分支内植物间的遗传距蔼取值,建议将此3个末端分支依次归并为3种：大桫椤、白桫椤和桫椤;(3)白桫椤属在科内处于基部位置,桫椤属奇桫椤亚属和黑桫椤亚属为衍生分支,赞同Tryon关于桫椤科进化和囊群盖起源的假说。     

SPORE MORPHOLOGY IN THE CYATHEACEAE. I. THE PERINE AND SPORANGIAL CAPACITY: GENERAL CONSIDERATIONS

The literature on cyatheaceous spore morphology relative to the presence of a perine layer is reviewed, and evidence based on a sodium-hydroxide assay is presented indicating that the outer scultpine layer in certain cyatheaceous spores is perine. Perine so defined characterizes Metaxya, paleotropical and certain neotropical species of Sphaeropteris, nearly all species of Alsophila, all species of Nephelea, and certain species of Trichipteris and Cyathea. It is lacking in Lophosoria, many species of Trichipteris and Cyathea, and all species of Cnemidaria. Two major patterns of spore number per sporangium in the family are reported. Lophosoria, Sphaeropteris, Trichipteris, Cyathea, Cnemidaria, and probably Metaxya are characterized by 64-spored sporangia, whereas most species of Alsophila and all species of Nephelea are characterized by 16-spored sporangia. The congruence of this generic distribution of sporangial-capacity types with Tryon's phyletic arrangement of cyatheaceous genera supports the naturalness of his system. The intrasporangial germination of spores retained in dehisced and dispersed sporangia supports the suggestion that decreased spore number per sporangium in Alsophila and Nephelea may relate to the role of the sporangia as dispersal units. The decreased number of spores per sporangium is associated with a trend toward increase in the number of sporangia per sores, with the highest known count approaching 1000 sporangia per sorus. The Alsophila-Nephelea evolutionary line has probably not been ancestral in the phylogeny of the more advanced groups of ferns.     

广西蕨类植物新记录属——白桫椤属

报道了广西蕨类植物一新记录属——白桫椤属。该属植物以茎干直立,乔木状;叶大型,簇生于茎干顶端,叶柄平滑、有疣突或皮刺,基部鳞片的细胞一式;叶背灰白色;叶脉分叉,2～3叉;无囊群盖等为主要特征。目前该属植物在广西首次记录到白桫椤。根据原始文献及广西的标本对该种进行了描述,并提供了墨线图。     

Relations between submarginal and marginal sori in ferns

In contrast to the generally favoured hypothesis that the marginal position of sporangia and sori is the primitive condition, Hagemann (e.g. 1984a) holds the view that the superficial position of reproductive organs is primitive. Marginal reproductive organs correlate with a distinct succession of a vegetative and a reproductive developmental phase in the sporophyll (phase differentiation in leaf development). Since phase differentiation means an increase in complexity, marginal reproductive organs are considered advanced. In order to examine the relations between submarginal and marginal sori in the light of Hagemann's hypothesis, various soral forms of ferns were analysed using the methodological framework of construction morphology sensu Weber (1958). The sorus is regarded as a reproductive region of the leaf surface that comprises not only the sporangia but also the receptacle as an essential component. The growth pattern of the receptacle, the developmental succession of the sporangia, and the formation of the indusia lead to the recognition of several sorus types.In the present paper the basipetal marginal sorus is presented and defined as the combination of (a) a ridge-, cushion-, or rod-shaped receptacle, (b) a basipetal sequence of sporangia initiation, and (c) the presence of a lower and an upper indusium or of a cup-shaped indusium. As the first examples of this type, the sori of selected Hypolepidaceae and Dennstaedtiaceae are treated in detail. Regarding the phylogenetic relationships of the present families, the following conclusions can be drawn: (1) Owing to the meristem fusion of the indusia the sorus ofDennstaedtia possesses a new quality as compared to that of the Hypolepidaceae. Therefore, the Hypolepidaceae and the Dennstaedtiaceae should not be combined in a single family, as suggested by some authors. (2) The marginal position of the sori, and, in addition, the highly complex soral construction of the Dennstaedtiaceae, are not in support of the primitiveness of the two families, as sometimes assumed. (3) A sorus construction as found inDennstaedtia glauca can be gradually derived from a less elaborate one, such as that found, e.g., inPteridium. The continuous increase in the perfection of the soral construction (Pteridium Dennstaedtia) suggests a placement of the Dennstaedtiaceae after the Hypolepidaceae.     

Relations between submarginal and marginal sori in ferns IV. The sori of Osmunda and selected Schizaeaceae and their morphological relations

Developmental types of sori in Osmunda and Schizaeaceae are comparatively studied and discussed in a systematic context. (1) The ``simultaneous marginal sorus' is characterized by a minute knob-like receptacle representing the thickened tip of a short costa, simultaneous initiation of massive sporangia, and absence of indusia. (2) The ``acropetal marginal sorus' is characterized as an accessory outgrowth at the end of a costa or at the margin of a sorophore, comprising a minute or elongated bifacial receptacle, acropetal initiation of massive marginal sporangia, and minute or long-extended or hood-shaped upper indusia. The state of superficial singly-arranged massive sporangia such as in Todea may give rise to the Osmunda type of the simultaneous marginal sorus and to the Schizaeaceae types of the acropetal marginal sorus. Phase differentiation in leaf development combined with complete laminar reduction leads to the Osmunda type, while phase differentiation in leaf lamina development and additional processes may give rise to the Mohria, Anemia, and Lygodium types. These morphological results support recent phylogenetic studies by other authors in disclaiming close relationships between Osmundaceae and Gleicheniaceae, and between Marattiaceae and Ophioglossaceae (``Eusporangiatae'). They also potentially support relationships between Osmunda- ceae and Schizaeaceae or between Ophioglossaceae and Schizaeaceae. The high complexity of the sori in Marattiaceae and Ophioglossaceae is surprising if the two families are viewed as basal groups.     

广东西部云开山自然保护区蕨类植物多样性调查

野外调查和标本鉴定结果表明,云开山自然保护区共有185种蕨类植物,其中10种为广东新分布:中华桫椤(Alsophila costularis)、白桫椤(Sphaeropteris brunoniana)、有刺凤尾蕨(Pteris setulosacostulata)、禾秆亮毛蕨(Acystopteris tenuisecta)、高大短肠蕨(Allantodia muricata)、细裂铁角蕨(Asplenium tenuifolium)、无盖肉刺蕨(Nothoperanema shikokianum)、蓝色鳞毛蕨(Dryopteris polita)、云南舌蕨(Elaphoglossum yunnanense)和黑鳞剑蕨(Loxogramme assimilis)。该地蕨类植物区系主要由亚洲热带亚热带分布和东亚分布成分(各占约37%的种类)构成,以鳞毛蕨科(Dryopteridaceae)、水龙骨科(Polypodiaceae)、金星蕨科(Thelypteridaceae)、蹄盖蕨科(Athyriaceae)和铁角蕨科(Aspleniaceae)的植物最为丰富,包含13种中国特有种但缺乏本地特有成分。垂直分布的统计结果表明,绝大多数植物(143种)生活在海拔900~1300 m的常绿阔叶林下,局限分布于海拔800 m以下的植物有15种,而局限分布于海拔1400~1700 m的植物有23种。该地的珍稀蕨类植物有马尾杉(Phlegmariurus phlegmaria)、华南马尾杉(P. austrosinicu)、薄叶阴地蕨(Sceptridium daucifolium)、小黑桫椤(Gymnosphaera metteniana)、中华桫椤、白桫椤、长柄车前蕨(Antrophyum obovatum)、黑鳞复叶耳蕨(Arachniodes nigrospinosa)、两广鳞毛蕨(Dryopteris liangkwangensis)、云南舌蕨、燕尾蕨(Cheiropleuria bicuspis)、锯蕨(Micropolypodium okuboi)和穴子蕨(Prosaptia urceolaris)。     

A new tree fern stem,Heilongjiangcaulis keshanensis gen. et sp. nov., from the Cretaceous of the Songliao Basin,Northeast China: a representative of early Cyatheaceae

A new genus and species of Cretaceous Cyatheacean tree fern, Heilongjiangcaulis keshanensis gen. et sp. nov., is erected for several permineralized stems collected at the Keshan County in Songliao Basin, Heilongjiang Province, Northeast China. The new taxon is characterized by a dictyostelic, erect stem with dense multicellular scales and surrounded by persistent petiole bases and adventitious roots. The stem contains a central pith lacking medullary bundles, which is surrounded by a dictyostele, and the cortex externally. Each meristele of the dictyostelic ring is enclosed by a sclerenchyma sheath. The pith and cortex are parenchymatous. The proximal petiole bases present a frond trace composed of numerous meristeles, arranged in 1 abaxial and 2 adaxial arcs, with internally projecting bundles on the upper and lateral sides. The feature combination of the new genus is nearly identical to the anatomical structures of modern scaly genera of the Cyatheaceae apart from the absence of medullary bundles. It is interpreted as a primitive representative of early Cyatheaceae, that closely resembles the modern scaly genera, which suggests that in the Cretaceous, the tree ferns in this family were already in possession of most of the anatomical characteristics observed in extant taxa. The fossil records of the stems, petioles, and spores indicate that during the Jurassic and Cretaceous, the Northeastern region of Asia may have been one of the distribution centers of early Cyatheaceae.     

Cyathea cranhamii sp. nov. (Cyatheaceae), anatomically preserved tree fern sori from the Lower Cretaceous of Vancouver Island, British Columbia

Permineralized cyatheaceous sori occur among remains of conifers, fungi, and other plants in newly discovered calcareous concretions from Early Cretaceous (Barremian) marine sediments of Vancouver Island, British Columbia, Canada. Sori are superficially attached in two rows to narrow pinnules and display a globose sphaeropteroid indusium. Annulate sporangia with multicellular stalks diverge from a basal, vascularized receptacle. The nearly vertical uniseriate annulus is not interrupted by the stalk. The sporangia bear 64 trilete spores with perispore sculpturing that ranges from irregular granulate/echinate to prominent rodlets. These specimens, described as Cyathea cranhamii sp. nov., are the first anatomically preserved tree fern sori from the fossil record. They represent the most ancient evidence for fertile structures of the Cyatheaceae and demonstrate that essentially modern species of cyatheaceous tree ferns had evolved by the Early Cretaceous.     

桫椤科三种植物配子体发育的研究

用土壤培养桫椤Alsophila spinulosa(Wall.exHook.)R.M.Tryon、中华桫椤A.costularis Baker和白桫椤Sphaeropteris brunoniana(Hook.)R.M.Tryon的孢子,利用光学显微镜对其配子体发育的各个阶段进行了观察,包括孢子形态及其萌发、原丝体发育特点、片状体和生长点的形成及分化、假根特征、精子器和颈卵器的分化及发育,以及原叶体感染真菌后的显微特征。初步讨论了桫椤科不同属(种)间的配子体发育的系统学意义和有性世代的濒危原因。     

中国黑桫椤属植物(桫椤科)一新组合种

根据来自叶绿体trnL内含子和DNAtrnL F间隔区序列等的证据 ,滇南桫椤Al sophilaaustro yunnanensisS .G .Lu将组合到黑桫椤属Gymnosphaera之下更趋合理。因此 ,本文报道中国黑桫椤属植物一新组合种 :滇南黑桫椤Gymnosphaeraaustro yunnanensis (S .G .Lu)S .G .LuetC .X .Li。     

The Cyatheaceae (Polypodiopsida) of Peru

A summary is presented of all Peruvian species of scaly tree ferns (Cyatheaceae). A total of 83 species is native to Peru (Sphaeropteris 1 sp., Alsophila sect. Alsophila 9 spp., Alsophila sect. Gymnosphaera 1 sp., Cyathea 72 spp.), with 13 endemic species in Cyathea and one in Alsophila. The following new species are described and illustrated: Alsophila gastonyi, Cyathea chontilla, C. nephele, C. polliculi, and C. praetermissa. The genera Cnemidaria and Trichipteris are no longer recognized and merged in Cyathea. Necessary combinations are made for Cyathea alatissima, C. nervosa, and C. uleana. Cyathea serpens is presented in detail and its relationship is discussed. An artificial key to the Peruvian species of Cyatheaceae is provided.     

内蒙古乌达地区早二叠世晋囊蕨属一新种

Chansitheca (Sphenopteris) wudaensis sp. nov. is described based on the materials collected from the Lower Permian Shansi Formation of Wuda, Nei Mongol, China. The frond is small and at least two times pinnate. The pinnule is sphenopteroid. The sori are borne on the abaxial surface of the pinnule, arranged in two rows parallel to the midvein, elliptic,and composed of 4-10 round sporangia. The indusium is absent. The annulus is transverse and complete, and consists of about 18 thickened cells. The features of the fertile parts, sori, sporangia and annulus and the absence of indusium indicate that the new species belongs to the Gleicheniaceae.     

Fertile organs and in situ spores of a new dipteridaceous fern Hausmannia sinensis from the Jurassic of northern China

As a representative fossil member of the dipteridaceous fern, genus Hausmannia was reported worldwide from the Mesozoic strata; however, little is known about the fertile structures, including sporangia and in situ spores, of this genus. In this study, a new species Hausmannia sinensis was identified from the Middle Jurassic of Nei Mongol (Inner Mongolia), northern China. The specimens are compressions and are well preserved with details of sporangia and in situ spores. The leaf laminae are broadly fan-shaped, with an almost entire margin. Primary and lateral veins dichotomously branch to form square or polygonal meshes. Each ultimate mesh bears one to two circular sori of 0.4 mm in diameter. Sori are exindusiate; each sorus contains three to six round to ovoid sporangia. The annulus is developed and oblique, with stomial region present in proximal position. Spores are trilete, circular to oval in shape. Both proximal and distal surfaces are covered with baculate to subverrucate sculptures. Spores range from 20 to 30 μm in diameter (average 28 μm), and are comparable to the dispersed genera Baculatisporites Thomas and Pflug and Apiculatisporis Potonié and Kremp. Hausmannia sinensis represents the first compression species of genus Hausmannia form Eurasia, which shows the combination of well-preserved sori, sporangia, annuli and in situ spore characters, and is therefore helpful for further understanding the diversity and evolution of the Dipteridaceae fern lineage through time.     

Complete chloroplast genome sequence of a tree fern Alsophila spinulosa : insights into evolutionary changes in fern chloroplast genomes

Background  

Ferns have generally been neglected in studies of chloroplast genomics. Before this study, only one polypod and two basal ferns had their complete chloroplast (cp) genome reported. Tree ferns represent an ancient fern lineage that first occurred in the Late Triassic. In recent phylogenetic analyses, tree ferns were shown to be the sister group of polypods, the most diverse group of living ferns. Availability of cp genome sequence from a tree fern will facilitate interpretation of the evolutionary changes of fern cp genomes. Here we have sequenced the complete cp genome of a scaly tree fern Alsophila spinulosa (Cyatheaceae).     

内蒙古乌达地区早二叠世晋囊蕨属一新种

通过扫描电镜研究,建立了内蒙古中部乌达地区下二叠统山西组晋囊蕨诉一个新种Ｃｈａｎｓｉｔｈｅｃａ ｗｕｄａｅｎｓｉｓ ｓｐ ．ｏｎｖ．。新种的小心片Ｓｐｈｅｎｏｐｔｅｒｉｓ型,囊群椭圆形,由４～１０个孢子囊组成,着生一示羽片背面,位于中脉的两侧。环带横列于孢子囊的上部,完全,单排细胞,细胞数目约为１８个,这些特征表明新种属于里白科。     

SPORE MORPHOLOGY IN THE CYATHEACEAE. II. THE GENERA LOPHOSORIA,METAXYA, SPHAEROPTERIS,ALSOPHILA, AND NEPHELEA

Scanning electron microscopy and transmitted light microscopy are used in a palynological study of Lophosoria, Metaxya, Sphaeropteris, Alsophila, and Nephelea of the tree fern family Cyatheaceae. The monotypic American genera Lophosoria and Metaxya each have a unique spore morphology which reinforces the taxonomic distinctness of these genera as indicated by their other characters. All investigated paleotropical species of Sphaeropteris develop a single type of perine characterized by coarse, pointed projections. In the neotropics, the Sphaeropteris horrida group shares this perine type, whereas all other neotropical Sphaeropteris species appear to have a different kind of perine with fine hair-like processes. The exine in paleotropical Sphaeropteris appears uniformly unsculptured, whereas in the neotropics several exine morphologies are found. In Alsophila all investigated neotropical species and the vast majority of the paleotropical species are characterized by a basically ridged perine morphology and an unsculptured exine. In several paleotropical Alsophila species, however, a perine with hair-like processes similar to those in neotropical Sphaeropteris is found, and the exine in several species is variously pitted. In at least one paleotropical Alsophila species, the porate exine morphology is indistinguishable from that in the neotropical genus Cnemidaria. The spores of the American genus Nephelea are similar to those of the majority of Alsophila species in ridged perine morphology and unsculptured exine. Several new instances of atypical spore numbers per sporangium are reported in Sphaeropteris and Alsophila. These and the palynological data are discussed in a taxonomic framework. The spore morphology in these genera is consistent with Tryon's recent generic revision of the family.     

广东阳春百涌自然保护区桫椤科植物资源调查

广东阳春百涌自然保护区属云开山脉,区内桫椤科植物资源丰富,但具体种类及其资源量并无准确资料,为此我们开展了相关野外调查和统计。调查表明,保护区内共有4种桫椤科植物,即桫椤Alsophila spinulosa、大叶黑桫椤Gymnosphaera gigantea、小黑桫椤G. metteniana和黑桫椤G. podophylla。依据保护区内桫椤科植物主要沿林道分布的特点,提出了按林道长度估算资源量的方法,据估算,保护区内有桫椤约900株,大叶黑桫椤约1.5万株,小黑桫椤约320株,黑桫椤约4500株。阳春百涌保护区是华南地区大叶黑桫椤的主要保存地。     

基于群落种间关系的桫椤植物保护策略研究进展

桫椤科植物是古老的孑遗植物,现存的桫椤天然群落极为罕见,为了解桫椤(Alsophila spinulosa)及桫椤科(Cyatheaceae)植物在生物群落的中生存与保育状态研究现状,本文通过搜集国内外关于桫椤及桫椤科植物相关文献进行研究。桫椤科植物分布于高温潮湿、光斑直射的生境,常以斑块状分布于常绿阔叶林林缘。现阶段,桫椤科植物保护面临的主要问题是生态旅游、毁林开荒、环境污染、药材的违法采摘和挖掘等,这导致桫椤科植物的生境遭受干扰和破碎,桫椤科植株数量下降。对桫椤科植物的保护方式主要有就地保护、迁地保护,分子生物学技术在桫椤科植物繁育中应用逐渐增加。生物和非生物因素制约着桫椤科植物的分布和多样性,其中,生物因素特别是种间关系对桫椤植物的生长发育和分布有着重要影响,从种间关系的视角出发探讨桫椤科植物的保护或许不失为新时代桫椤科植物保护的有效方式。讨论了桫椤植科物相关的种间关系：桫椤科植物-微生物互作、桫椤科植物-其他植物互作、桫椤科植物-动物互作,探讨了桫椤科植物面临灭绝的因素以及近年来桫椤科植物相关的保护策略及成效,为桫椤科植物的有效利用和保护提供一定理论基础。     

Sporangium Exposure and Spore Release in the Peruvian Maidenhair Fern (Adiantum peruvianum,Pteridaceae)

We investigated the different processes involved in spore liberation in the polypod fern Adiantum peruvianum (Pteridaceae). Sporangia are being produced on the undersides of so-called false indusia, which are situated at the abaxial surface of the pinnule margins, and become exposed by a desiccation-induced movement of these pinnule flaps. The complex folding kinematics and functional morphology of false indusia are being described, and we discuss scenarios of movement initiation and passive hydraulic actuation of these structures. High-speed cinematography allowed for analyses of fast sporangium motion and for tracking ejected spores. Separation and liberation of spores from the sporangia are induced by relaxation of the annulus (the ‘throwing arm’ of the sporangium catapult) and conservation of momentum generated during this process, which leads to sporangium bouncing. The ultra-lightweight spores travel through air with a maximum velocity of ~5 m s^-1, and a launch acceleration of ~6300g is measured. In some cases, the whole sporangium, or parts of it, together with contained spores break away from the false indusium and are shed as a whole. Also, spores can stick together and form spore clumps. Both findings are discussed in the context of wind dispersal.